Fishes of the Gulf of Maine

UNITED STATES DEPARTMENT OF THE INTERIOR, Douglas McKay, Secretary J2
FISH AND WILDLIFE SERVICE, John L. Farley, Director

FISHES OF THE GULF OF MAINE

BY HENRY B. BIGELOW AND WILLIAM G. SCHROEDER

First Revision

FISHERY BULLETIN 74

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

VOLUME 53

[Contribution No. S92, Woods Hole Oceanographic Institution}

UNITED STATES GOVERNMENT PRINTING OFFICE • Washington : 1953

NOTICE

Bigelow and Schroeder's "Fishes of the Gulf of Maine" was printed in
1953 and went on sale at the U.S. Government Printing Office on February 12,
1954. This was a revision of an earlier work of the same name by Bigelow and
W. W. Welsh (1925); 3,493 copies of the revision were printed. Of these,
2,000 copies were distributed by the Fish and Wildlife Service, the remainder
by the Government Printing Office. The Service supply was exhausted in Janu-
ary 1961 and that of the Printing Office in March 1961.

A photo-offset reprinting was issued in 1964 jointly by the Woods Hole
Oceanographic Institution and the Museum of Comparative Zoology, Harvard
University, organizations with which the authors have been associated for many
years. Since then, additional reprintings have followed. The contents of the book
are precisely the same as published in 1953 save for the addition of this note
and the deletion of two lines at the foot of the title page saying "For sale by the
Superintendent of Documents, U.S. Government Printing Office, Washington 25,
D.C. Price $4.25 (Buckram)."

This reprint may be obtained from the Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts 02138.

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 Petromyzonidae 9

Hag Myxine glutinosa Linnaeus 10

Sea lamprey Petromyzon 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 taurus Rafinesque 18

Mackerel sharks. Family Isuridae 20

Mackerel shark Lamna nasus (Bonnaterre) 20

Sharp-nosed mackerel shark, Mako I sums oxyrinchus Rafinesque 23

Maneater, White shark Carcharodon carcharias (Linnaeus) 25

Basking sharks. Family Cetorhinidae 28

Basking shark Celorhinus maximus (Gunnerus) 28

Thresher sharks. Family Alopiidae 32

Thresher Alopias vulpinus (Bonnaterre) 32

Cat sharks. Family Scyliorhinidae 34

Chain dogfish Scyliorhinus retifer (Garman) 34

Smooth dogfishes. Family Triakidae 34

Smooth dogfish Mustelus canis (Mitchill) 34

Requiem sharks. Family Carcharhinidae 36

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 milberti (Muller 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 Cenlroscymnus coelolepis Bocage and Brito Capello 52

Gurry sharks. Family Dalatiidae 53

Greenland shark Somniosus microcephalus (Bloch and Schneider) 53

Dalalias 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 Mitchill 67

Smooth- tailed skate Raja senla Garman 70

Thorny 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 Dasyatidae 74

Sting ray Dasyatis centroura (Mitchill) 74

Cow-nosed rays. Family Rhinopteridae 76

Cow-nosed ray Rhinoptera bonasus ( Mitchill) 76

Devil rays. Family Mobulidae 77

Devil ray Manta 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 86

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) 106

Shad Alosa sapidissima (Wilson) 108

Thread herring Opislhonema oglinum (LeSueur) 112

Menhaden Brevoortia tyrannus (Latrobe) 113

Anchovies. Family Engraulidae 118

Anchovy Anchoa mitchilli (Cuvier and Valenciennes) 118

Striped anchovy Anchoa hepsetus (Linnaeus) 119

Salmons. Family Salmonidae H»

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 (Mtiller) 134

Smelt Osmerus mordax (Mitchill) 135

Argentines. Family Argentinidae 139

Argentine Argentina silus Ascanius 139

Luminescent fishes ™*

Lanternfishes. Family Myctophidae 141

Headlight-fish Diaphus effulgens (Goode and Bean) 142

Lanternfish Myctophum affine (Liitken) 143

Pearlsides. Family Maurolicidae 143

Pearlsides Maurolicus pennanti (Walbaum) 144

Viper Fishes. Family Chauliodontidae 145

Viperfish Chauliodus sloani Bloch and Schneider 145

Stomiatids. Families Gonostomidae and Stomiatidae 146

Cyclothone Cyclothone signata Garman 146

Stomias Stomias ferox Rheinhardt 147

Stomioides nicholsi Parr 147

Trigonolampa miriceps Regan and Trewavas 148

Hatchet fishes. Family Sternoptychidae 149

Silver hatchet fish Argyropelecus aculealus Cuvier and Valenciennes 149

Eels. Families Anguillidae, Congridae, Simenchelyidae, Synaphobranchidae,

Nemichthyidae, and Ophichthyidae 150

Eel Anguilla rostrata (LeSueur) 151

American conger Conger oceanica ( Mitchill) 154

CONTENTS

Bony fishes. Class Osteichthyes — Continued Paee

Eels — Continued

Slime eel Simenchelys parasiticus Gill 157

Long-nosed eel Synaphobranchus pinnatus (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

Mummichogs 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

Halfbeaks. 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

Cod Gadus callarias Linnaeus 182

Tomcod Microgadus tomcod (Walbaum) 196

Haddock Melanogrammus aeglefinus (Linnaeus) 199

American pollock Pollachius virens (Linnaeus) 213

White hake Urophycis tenuis (Mitchill) 221

Squirrel hake Urophycis chuss (Walbaum) 223

Spotted hake Urophycis regius (Walbaum) 230

Long-finned hake Urophycis chesteri (Goode and Bean) — 232

Blue hake Antimora rostrata Giinther 233

Hakeling Physiculus fulvus Bean 233

Four-bearded rockling Enchelyopus cimbrius (Linnaeus) — 234

Cusk Brosme brosme (Miiller) 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 Reinhardtius hippoglossoides (Walbaum) 258

American dab Hippoglossoides platessoides (Fabricius) 259

Summer flounder Paralichthys dentatus (Linnaeus) 267

Four-spotted flounder Paralichthys oblongus (Mitchill) 270

Yellow-tail Limanda ferruginea (Storer) 271

Winter flounder Pseudopleuronectes americanus (Walbaum) 276

Smooth flounder Liopsetta putnami (Gill) 283

Witch flounder Glypiocephalus cynoglossus (Linnaeus) 285

Sand flounder Lophopsetta maculala (Mitchill) 290

Gulf Stream flounder Citharichthys arctifrons 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

Sil verside 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 wheatlandi Putnam 310

Four-spined stickleback Apeltes 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 scombrus Linnaeus 317

Chub mackerel Pneumatophorus colias (Gmelin) 333

Striped bonito Euthynnus pelamis (Linnaeus) 335

False albacore Euthynnus alleteratus (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 Ruvetius 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 Centrolophus 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 setapinnis (Mitchill) 378

Lookdown Selene vomer (Linnaeus) 279

Leatherjacket Oligoplites saurus (Bloch and Schneider) 380

Threadfin Alectis crinitus (MitcbilO 381

CONTENTS VII

Bony fishes. Class Osteichthyes — Continued P»w

Bluefishes. Family Pomatomidae 382

Bluefish Pomatomus saltalrix (Linnaeus) 383

Sea basses. Family Serranidae 389

Striped bass Roccus saxatilis (Walbaum) 389

White perch Morone americana (Gmelin) 405

Sea bass Centropristes striatus (Linnaeus) 407

Wreckfish Polyprion americanus (Bloch and Schneider) 409

Catalufas or Big Eyes. Family Priacanthidae 410

Short big-eye Pseudopriacanlhus altus (Gill) 410

Porgies. Family Sparidae 411

Scup Stenotomus 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 Lacfipede 423

Kingfish Menticirrhus saxatilis ( Bloch and Schneider) 423

Black drum Pogonias cromis (Linnaeus) 425

Tilefishes. Family Branchiostegidae 426

Tilefish Lopholalilus chamaeleonticeps Goode and Bean 426

Rockfishes. Family Scorpaenidae 430

Rosefish Sebastes marinus (Linnaeus) 430

Black-bellied rosefish Helicolenus dactyloplerus (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 Arlediellus uncinatus (Reinhardt) 440

Mailed sculpin Triglops ommatislius Gilbert 441

Grubby Myoxocephalus aeneus (Mitchill) 443

Shorthorn sculpin Myoxocephalus scorpius (Linnaeus) 445

Longhorn sculpin Myoxocephalus octodecemspinosus (Mitchill) 449

Staghorn sculpin Gymnocanthus tricuspis (Reinhardt) 452

Arctic sculpin Cottunculus microps Collett 453

Sea raven Hemitriplerus americanus (Gmelin) 454

Alligatorfishes. Family Agonidae 457

Alligatorfish Aspidophoroides monopterygius (Bloch) 457

Lumpfishes. Family Cyclopteridae 459

Lumpfish Cyclopterus lumpus Linnaeus 459

Spiny lumpfish Eumicrotremus spinosus (MtiUer) 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 Prionotus carolinus (Linnaeus) 467

Striped sea robin Prionotus evolans (Linnaeus) 470

Armored sea robin Peristedion minialum Goode 471

Flying gurnards. Family Dactylopteridae 472

Flying gurnard Dactyloplerus 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 Ammodytidae 487

Sand launce Ammodytes americanus DeKay 488

VIII CONTENTS

Bony fishes. Class Osteichthyes— Continued Fag»

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 punctaius (Fabricius) 497

Radiated shanny Ulvaria subbifurcala (Storer) 498

Wrymouths. Family Cryptacanthodidae 500

Wry mouth Cryplacanthod.es maculatus Storer 500

WolfBshes. Family Anarhichadidae 502

WolfBsh Anarhichas lupus Linnaeus 503

Spotted WolfHsh 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 reticulalus Reinhardt 516

Cusk eels. Family Ophidiidae 517

Cusk eel Lepophidium cervinum (Goode and Bean) 517

Toadfishes. Family Batrachoididae 518

Toadfish Opsanus tau (Linnaeus) 518

Triggerfishes. Family Balistidae 520

Triggerfish Batistes carolinensis Gmelin 520

Filefishes. Family Monacanthidae 521

Filefish Monacanthus hispidus (Linnaeus) 522

Filefish Monacanthus ciliatus (Mitchill) 523

Orange filefish Alutera 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 Chilomycterus schoepfii (Walbaum) 527

Ocean Sunfishes or Headfishes. Family Molidae 528

Sunfish Mola mola (Linnaeus) 529

Sharp-tailed sunfish Masturus lanceolatus (Lifinard) 53 1

Anglers. Family Lophiidae 532

American Goosefish Lophius americanus Cuvier and Valenciennes 532

Sargassum fishes. Family Antennariidae 541

Sargassum fish Histrio piclus (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 1913 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 ; ' subsequent parts describing the plankton of
the offshore waters of the Gulf and the physical
characteristics of its waters were published in
1926-27, as Part 2.

The preparation of the section on the fishes was
assigned originally to W. W. Welsh, who had
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.

i 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
Bank, 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
amiss 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, cunners, 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

> Kept. U. S. Comm. Fish., (1904) 1905, pp. 163-188.
i Smithsonian Contribution to Knowledge, vol. 30, 1895.
' Bull. U. S. Fish Comm., Vol. 17, 1898, pp. 85-111.
' Bull. U. S. Bur. Fish., vol. 31, Pt. 2, 1913, pp. 549-794.

FISHES OF THE GULF OF MAINE

Hole fishes; Halket's 8 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," 9 and Ehren-
baum's summary of the many scattered accounts
of the eggs and larvae of northern fishes10 have
been especially 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 Scattergood 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 ot the Dominion of Canada and Newfoundland
1913, 138 pp.

» Proc. Nova Scotia Inst, of Science, vol. 19, Pt. 1, 1935, pp. 17-113.

» The fishes of Great Britain and Ireland, by F. Day, Text vol. 1, CXII+
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.

10 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
Tuning; 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,
skates, rays, and chimaera are 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 Oceanographic 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

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 fiah 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.11 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

i' 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 IS.

2nd Donal
Fin i

Procoudal
Pit

Caudal

tst.Ana.1 fin
Ventral fin

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 through 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 fiat, 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 is
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 11

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

11. 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 to 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 18

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 (soft-rayed 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. 301

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," 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

KeyD

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 sallflsh would also come under this heading should one ever be taken In the Gulf of Maine. The distinctions between It and the spearfishts are
Blven 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 unlet 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 Power) 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 Cusk, 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 in 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.13

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

u See Blgelow and Sihroeder, Fishes Western North Atlantic, Pt. 1, ch. 2,
1948, pp. 35-36, for references.

coast of Newfoundland by Dean (1900) ; M from the
neighborhood of the Faroe Islands by Jensen; 1S
from Norway by Hjort; 16 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% 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
Hemisphere 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. Set., vol. 2, Pt. 2, Art. 2, 1900.

" Vlden. Meddel. Dansk naturhlst. Forenlng, 1900, p. 1.

" 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 though It Is to be eipected 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 c f 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 1913
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 when 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

'*

"^M^**

Figure 3. — Sea lamprey {Petromyzon marinus), about 18 inches long, Merrimac 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
Eivers, 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 only a short distance upstream ; even within
the influence of the tide, although invariably in
fresh water. But they are able to ascend falls,
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 Merrimac
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 lampreys 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 November 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.

» Ooode, Fish. Ind. U. S., Sect. 1, 1884, 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, PetitCodiac, and
Shubenacadie Rivers 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 Massachusetts.

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 tho 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 their 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

si Also reported from "West Africa" by Oflnther, 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 surmouating 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.

aa For an account of the lamprey fishery in New England during the first
half of the 19th century, see Qoode, Fish, and Fishery Ind. U. S., Sect. 1, 1884,
p. 680.

" Bailey, Biol. Survey Merrimack Watershed, New Hampshire Fish and
Game Dept., 1938, p. 158.

a* Information gathered for us by Dr. A. G. Huntsman.

a' 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 tbey 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 ( "he tero cereal"), 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, see Bigelow and Schroeder, Fishes Western
North Atlantic, Ft. 1, ch. 3, 1948, p. 64.

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 (Larrma nasus), and the blue
shark (Prionace glauca), occur with us in numbers
sufficient for one to be fairly sure of seeing them
during a summer's boating off the coast north of
Cape Cod.

With the larger sharks generally so scarce (the
mackerel shark is harmless to anything larger than
the fishes on which it feeds, and the blue
shark is also harmless, although better armed),
the danger of attacks on bathers is negligible in
our Gulf. Indeed, not a single well-authenticated
instance of the sort is on record w for the past
80 years for the coast north of Cape Cod, though
the beaches are crowded every summer with
vacationists. But as long as the white shark or
man-eater (Carcharodon carcharias) does stray
occasionally into the Gulf (p. 26), it is always
remotely possible that we may be horrified some
summer by the news of tragedies such as occurred
on the New Jersey coast in July 1916, when
several persons were killed or injured, presumably
by a small shark of this species that was captured
nearby a few days later,28 and near Mattapoisett,
on Buzzards Bav, Mass., on July 25, 1936, when
a swimmer was fatally injured by a shark, species
not determined.29

17 In 1830 (an event often quoted) one Joseph Blaney, fishing from a small
boat In Massachusetts Bay off Swampscott, Mass., was attacked by some
fish that was seen to overset and sink his boat and, presumably, devoured
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
question.

" Murphy and Nichols (Brooklyn Mus. Quart., vol. 3, 1916, No. 4, pp.
145-160) give a detailed account of this occurrence.

» See Oudger (Amer. Midland Natural., vol. 44, 1050, p. 714) for clinical
details of this case.

All Gulf of Maine sharks give birth to young
that are not only practically adult in structure
but of relatively large size at birth, and there is a
placental connection between mother and embryo
in some, but not in others. Still other sharks lay
eggs; this is true of the chain dogfish {Scyliorhinus
retifer, p. 34), which is common out on the conti-
nental shelf from the offing of Cape Cod, south-
ward, and of its immediate relatives; also of the
heterodontids or Port Jackson sharks which are
not represented in the Atlantic.

There is so little market for sharks in Gulf of
Maine ports (attempts to introduce the dogfish as
a food fish having failed so far) that the amounts
landed in Maine and Massachusetts were only
about 240,000 pounds in 1947, and about 309,500
pounds in 1949; they interest fishermen chiefly as
nuisances because of the damage they do to nets
and other gear, except that mackerel sharks are
marketable.

It is possible to identify all the sharks so far
known from the Gulf (and this includes all that
are likely to occur there except strays) bv the
sizes and relative locations of the fins, and by
such tooth characters as may be seen at a glance
at the open mouth or easily felt with the finger
(after the shark is dead!).

We have attempted in the following descriptions
of the several species to include only such features
as will tell what shark is at hand ; for more minute
particulars we refer the reader to our account of
the 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

cusps 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 gref t 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 to 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 10

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. 56

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FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

21. 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

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 base 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, cunners, 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 M 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

» For an account of the embryos, see Springer, Copeia, 1948, No. 3, pp.
153-156.

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 Schroeder, 1948, p. 112.

Garman, 1911, pi. 6, figs. 4-6 (as Isurus ■punctatus).

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, Nahant, Massachusetts. Upper and lower first to
fifth teeth from center of jaw of a larger specimen from Platts Bank, 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 white 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 Eastport, Maine,31 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, unless
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.

» 8cattergood, Copela, 1949, No. 1, pp. 71-72.
n Hubbs, Copela, No. 173, 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 they
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. — la 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-swirnming 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 population 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. O. Jeflers.

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.36 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.38

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 McOonigle and Smith, Proc. Nova Scotia Inst. Sci., vol.
19, 193S, p. 160.

11 Cod tagging cruises of the U. 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 (llro-
phycis) .

In the Gulf of Maine, females containing em-
bryos have been taken in August (near Monhegan
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 in 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 (cf. 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

•'See Scattergood, Copela, 1949, p. 70, for further details as to landings In
Maine and methods of capture.
■ 3.7 meters as calculated from the size of Its jaws.

24

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figube 6. — Sharp-nosed mackerel shark, or Mako (Isurus oxyrinchus), about 64J^ 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). Recorded 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 in 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

" A Soutb 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 Illus.
News, July 14, 1628, p. 83) to hare been a maneater (Corcharodon).

found when landed to contain a large amount of
its flesh*

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 Gulf oj 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 caught
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 Fairlngton (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

*' Informal ion from Dr. W. J. Mixter.

41 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 tad 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

1 1nformation 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 3

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% feet long; ** 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.

11 For further details, see BIgelow 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.46

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 tbree 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 Provincetown
many 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
Fundy, July 2, 1932. And there are 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 the
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 the boat pieces of its teeth, from
which Dr. Garman was able to identify it.50

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.
* A female 12 feet , 8 Inches long, weighing 1 ,299 pounds, reported by Scatter-
good, Trefetben, and Coffin, Copela, 1961, p. 298.
» Putnam, Bull. Essex 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).

Belation 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 " 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 63
wrote of a case where one (apparently the 13-foot
specimen that he had described earlier as atwoodi)
turned furiously on a boat, but was lanced to death
and brought into Provincetown. And a 15-foot
shark, probably this species to judge from the il-
lustration of it that was published,64 that was
killed off Monomoy Point by two fishermen in
November 1928, 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."
Hence, so long as maneaters wander within

»' Proc. Essei Inst. Salem, vol. 6, 18/4. 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 Wltman and Lee Co.'s Market Letter for Nov. 8, 1928; called
to our attention by Dr. Lewis Radcllfle of the U. S. Bureau of Fisheries.

*» For details of this occurrence, see Piers, Proc. Nova Scoila Inst. Scl., vol.
18, 1934, pp. 196-198.

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

» London Illus. News, July 14, 1928, p. 63; 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), 26J4-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 sbark 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 67 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 are 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 59 concluded
from studies of basking sharks taken near the Isle

>' Bull. Boston Soc. Nat. Hist., No. 24, March 1921, p. 5.

" Described by Hussakof, Copela, No. 21, 1915, pp. 25-27.

« Philos. Trans. Roy. Soc. London, Ser. B., vol. 234, 1950, pp. 217-316.

of Skj'e 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 beeo.
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 then-
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

M For further details as to sizes of basking sharks, see Bigelow and
Schroeder, Fishes, Western North Atlantic, Pt. 1, 1948, pp. 151-152.

" Referred to by McKenzie, F-roc. Nova Scotia Hist. Sci.. vol. 20, 1940, p. 42.

" Shown In the film "Men of Arran."

u See Bigelow and Schroeder, Fishes Western North Atlantic, Pt. 1,
1948, pp. 153, 154, for details as to their centers of population and seeular
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, Philos. Trans. Eoy. Soc. London, Ser. B, No. 612, vol.
234, 1960, pp. 347—366 for detailed account.

" Norman and Fraser. Giant Fishes, Whales and Dolphins, 1937, p. 21.

* 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 6: 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

i 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 Fabricius 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 Provincetown, taken in a
fish trap; measured by J. Henry Blake.

1909. One, about 22 feet, in Provincetown Harbor.
1913. One, about 29 feet, Provincetown.

1925. One, about 29 feet, near Monhegan Island, Maine.
1931. Female, 12H 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 Provincetown 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."

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 Menemsha 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.

M McKenzle, Proc. Nova Scotia Inst. Scl., vol. 20, 1939, p. 14.

" McKenzle, Proc. Nova Scotia Inst. Scl., vol. 20, 1939, p. 14.

71 Personal communication from J. W. Burger.

" 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 given here of 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 day 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 6,580 pounds,
taken off Monterey, Calif., had a liver weighing
1,800 pounds, 60 percent of which was oil.7* But,
sad to say, 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 MacGinitie (Science, N. Ser., vol. 73, 1931, p. 496), 21
basking sharks were landed in Monterey, Calif., between November 22, 1930
and February, 1931.

'< MacGinitie, 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.75

'• Ooode, Fish. Ind. U. S., 1884, Sect. 1, p. 669.

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) 1 758

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 many 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 15% 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^r^9

^

\i

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;78 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
(Pomalomus) ; 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

Tfl Several of that size have been taken in the traps at Woods Hole.

11 There is another group of species of the genus, with very large eyes, that
live at greiter depths; for discussion of these, see Bigelow and Schroeder
(Fish. Western North Atlantic, Pt. 1, 1948, pp. 162, 163).

" 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 (e. g. Boston Harbor and Nahant).

Records for it along the coast of Maine include
the vicinity of Monhegan 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; 79 also in the Basin of Minas on the Nova
Scotian 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 McKenzie, Proc. Nova Scotia Inst. Sci., vol. 20, 1939, p. 14.

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. FAMILY SCYLIORHINIDAE

Distinctive features of these little sharks are
that they have five pairs 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

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 retijer), male, about 17 inches long, New Jersey. After Bigelow and Schroeder.

SMOOTH DOGFISHES.

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,

FAMILY TRIAKIDAE

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 canis), 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
Bigelow 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 bays, 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

*> Eipcriments have shown that It requires only 1 to 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 OP 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 fhis 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 Gulf 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 Fundy,
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 incursions 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

" The senses of this shark have been studied by Parker (Bull., U. S. Bur.
of Fish., vol. 29, 1M1, pp. 43-57), and by Sheldon (Jour. Compar. Neurol,
and Psychol., vol. 19, 1909, No. 3, p. 273).

,! Rept. U. S. Coram. Fish., (1906), 1907, Spec. Pap. 6, pp. 14-16.

" Occas. Pap. Mus. Zoo]., I'niv. Mich., No. 56, 1918, p. 15.

the Atlantic species,85 an anal fin, a caudal peduncle
lacking lateral keels, and sharp, bladelike teeth
with a single cusp. All bear "living" young;

B* 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 there 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 Schroeder, Fishes Western
North Atlantic, Part 1, 1948, pp. 250-251.

w The lemon shark (Negaprion brePtrowrU) 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

(%. ID-

Young tiger sharks are 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 in 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% 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.88
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 Gulf 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

M For further details and references, see Bigelow and Schroeder, Fishes
Western North Atlantic, Pt. 1, 1948, p. 269.

»' Bell and Nichols (Copela, 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. 282.

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
Provlncetown was an error. The original description of the specimen In
question (Atwood, Proc. Boston Soc. Nat. Hist., vol. 10, 1865, p. 81) suggests
that it was a mako (Isurus oxyrinchus) .

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 OP 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 90
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 Y% and 2 feet.81 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 inches (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
with 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 Gulf oj Maine and along Nova
Scotia. — Only one blue shark had been reported
definitely from the Gulf 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 18 were reported to us during the summer of
1935,94 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.

81 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. Sci. 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. R. 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.95

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 tbem 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.98

" By the late Walter H. Rich, who was long associated with the U. S. Bu-
reau of Fisheries.

« Cornish, Contr. Canadian Biol. (1902-1905) 1907, p. 81.

« Rept. Newfoundland Fish. Res. Lab., 1935, p. 79.

88 Robert Goffin 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 bad 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-novas
(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 lerrae-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,89 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. Verrill, Is in the Museum of Com-
parative Zoology.

1-1 See Jordan and Evermarm, Bull. 47, V. 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 east 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 Schreeder, 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 natural 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 it9
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, relatively, than
in the brown shark, and sometimes more definitely
sickle-shaped.

The upper teeth are broadly triangular; nearly
erect toward the center of the mouth but weakly
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 their
tips; the pelvics and anal fins grayish white.

Size. — -The usual length at birth is a little more
than three feet.* 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 obscurus 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 platyo-
don).

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 in 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
pair of gfil 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

General range. — 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 oj 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 39 In. (993 mm.) ; see Bigelow and Schroeder, Fishes Western
North Atlantic, Pt. 1, 1948, p. 387.

» Probably this species and not the brown shark because 11-12 feet long.

I In the first edition of this book, the dusky shark was said to have 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 were
(Bigelow and Schroeder, Fishes Western North Atlantic, Pt. 1, 1948, pp.
292, 368).

' Seven feet 10 Inches 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, 1841, must be replaced by plumbeus proposed by Nardo In 1827
for the brown shark of the Adriatic.

44

FISHKRY 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 Cestracion tiburo).

Description. — The peculiar shovel-shaped head
of this shark is enough to distinguish it readdy
from any other shark known from the Gulf of
Maine, except for the hammerhead, from which it
is readdy 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 tad 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 Brazd 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.9

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
tiburo from China and from the Philippines, but without convincing evidence
as to its identity.

10 By Oarman, Mem. Mus. Comp. Zool., vol. 36, 1913, p. 161. Apparently
the specimen is no longer in existence.

11 Personal communication by R. n. Bodrnan, 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
(cf. 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 Ruppell, 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,

11 The larger hammerheads that are sometimes reported probably are not
this species, but the great hammerhead (Sphyrna 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 Fraser-Brunner (Rec. Austral. Mus., vol. 22, No. 3, 1950, pp. 213-214),
has shown that It cannot be separated from the Indo-Paclflc S. leuinl of
Griffith, 1834, a much older name.

i* Tortonese has recently pointed out (Ann. 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 RQppell, 1835, It being Identical with that Indo-Paclflc
species.

n 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 ; I8 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 in 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.

18 This specimen, obtained many years ago by Louis Agasslz, Is In the
Museum of Comparative Zoology.

'• Seen In the flsh market at Portland, Maine, by the late Walter H. Rich.

» McKenzle, Proc. Nova Scotia Inst. Sci., vol. 20, 1939, p. 13.

" Vladykov, Proc. Nova Scotia Inst. Sci., vol. 19, Pt. 1, 1935. p. 8.

" McKenzle, Prov. Nova Scotia Inst. Sci., vol. 20, 1939, p. 13.

FISHES OF THE GULF OF MAINE

THE SPINY DOGFISHES. FAMILY SQUALIDAE

47

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 he close
along the front margins of the two dorsals, the
first not more than 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

Fiqube 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 3% 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 or 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

23 About 14 miles offshore.

24 On Middle Ground about 25 miles off Cape Ann.

" Templeman, 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" (Trachinus 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 are 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 (Pbilos. Trans. Royal Soc., London, Ser. B, vol. 212, 1923, pp. 8,
27) describes the spines and gives clinical records of the effects of wounds
inflicted 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 10%
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 would be simply to list everv seaside
village 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 (50-65 fathoms in late
January 1950, gave birth to young on the deck of the vessel.

» Mclntire, Rept. 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 Stcenstrup, Pt. 2,
No. 30, 1914, p. 7) lists several definite records of this species at Sukkertoppen
and near nolsteinborg. 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-April), 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 until 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 Province town, 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 in 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)

" 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, Squalm cubensis 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 Bigelow and Schroeder, Fishes of the Western North
Atlantic. Ft. 1, 1948, p. 464.

ouuu
7500

7000

6500

eooo

| 5500
2 5000

g 4500

5 4000

I 3500

£ 3000

Si

f 2500

K

2000

•

1500

■

1000

■

500

.. . I

I

■ I.

ll

1

MAR. 'Will ' MAY

JUNE' JULY

'

AUG.

SEPT. ' OCT. ' NOV. '

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

u 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.36 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.
» Templeman, 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 line 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

87 For further discussion of the damage done by dogfish and of their com-
mercial possibilities, see Ann. Rept., Comm. Fish. Oame Mass. (1905), 1908,
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. U. 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. Fischer.

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% 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 weremadedown 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 Dalatias 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.

38 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 (Goode 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 (Cenlroscymnus coelolepis), female about 42% 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 3% 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.39

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 600 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 Cenlroscymnus coelolepis

Bocage and Capello. These sharks, as they were hauled
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 are 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 "
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

»• A male 228 mm. 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
pelvics, 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 16% feet was
reported from the Grand Banks in 1934; one of

« Arctic Regions, 1820, vol. 2, pi. 15, flgs. 3 and 4.
" Jenkins. Fishes British Isles, 1925, p. 325.

16 feet off Portland, Maine, in 1846; one of about
15 feet off Cape Ana 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 intc 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.46 But it is ex-

" Tales to the effect that it attacks Qreenlanders in their kyaks are appar-
ently mythical, and Doctor Porsild, Director of the biological station at
Disko, said that the Eskimos do not tear 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.46

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 Somniosut Tostratus, on the contrary, bears living
young.

*7 For recent discussion of the species of Somniosus, see Bigelow and
Schroeder. Fishes Western North Atlantic, Pt. 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 ;40 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.

19 This one was landed in Boston, where we saw it.

» When they gather to feed on whale, narwhal, and seal carcasses in their
northern home, they may linger for a long time in the 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, Mlndesk.
Jap. Steenstrup, vol. 2, No. 30, 1914, p. 12); also Clark (Science, N. Ser.,
vol.41, 1915, p. 795).

56

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

i\/\/\/\AA

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% 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) ,63

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. 62,
1939, p. 85.

•■ Q 9 w «

Figure 23. — Spiny shark (Echinorhinus brucus), eastern Atlantic specimen about 3 feet long.

der. Drawing by W. P. C. Tenison.

From Bigelow and Schroe-

FISHES OF THE GULF OF MAINE

57

Color. — Described as dark gray, olive or brown
above, with metallic reflections, and with or
without 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.

Occurrence in the Gulf of Maine. — A single
specimen of this little known shark came ashore
at Provincetown in December 1878. This and
one taken near Buenos Aires more recently M
are the only 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, Ser. 2, vol. 10, 1875, p. 4801.

(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 the pelvic
fins into rodlike semitubular claspers (the copula-
tory organs) distinguishes males and females at a
glance. Some bear "living" young, ready for
independent existence; others lay eggs.

" Berg, Com. Ictiol. Comm. Mas. Nac. Buenos Aires, vol. 1, No. 1, 1898,
p. 10.

KEY TO GULF OF MAINE SKATES AND RAYS

1. The front of the head bears a pair of separate, ear-like fins, extending forward Devil ray, p. 77

The front of head does not bear a pair of separate ear-like fins extending forward 2

2. There is a large triangular caudal fin as well as two well developed dorsal fins on the tail Torpedo, p. 58

There is no distinct caudal Ad; the dorsal fins, if any, are very small 3

3. No long dorsal spine on tail Common skates 4

There is a long saw-edged dorsal spine (or spines) on the tail 11

4. The upper surface of the disc is marked with conspicuous black rosettes Leopard skate, p. 66

The markings on the upper surface of the disc are not in the form of black rosettes 5

5. There are no conspicuous thorns along the mid-dorsal zone of disc between the spiracles and 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 with black dots or dashes 6

6. There are no large thorns on the rear Y*-Yi of tail Smooth-tailed or Prickly skate, p. 70

There are one or more rows of large thorns along the rear part of tail as well as farther forward along it 7

7. There are no large thorns on upper side of disc between the spiracles and the level of axils of pectoral fins.

Barndoor skate, very small specimens, p. 61
The upper side of disc, rearward from spiracles, has more or fewer large thorns 8

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 9

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 ray; Numbfish, Crampfish

Bigelow and Schroeder, 1953, p. 96.

Garman, 1913, pi. 25, fig. 2, as Narcacion nobilianus.

Description. — No one would be apt to mistake a

Figttbe 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 naming 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 14K 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 ;w 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

" Comparison of American specimens with one from the North Sea revealed
no differences.

•' 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,

" Reported by Q. F. O. Hansen, then second mate and later master of the
U. S. Fish Commission schooner Oramput, who doubtless was acquainted
with the torpedo at Woods Hole.

•> The most recent record is of one 58 inches long, trawled on the southwest
part in December 1930.

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
objects. 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

w Under aquarium conditions the incubation period for the little skate
(R. trinacea) was 5 to 6 months (p. 69); and it ranged from 4H to about HH
months for 6 common European skates; see Clarke, Jour. Marine Biol.
Assoc. United Kingdom, vol. 12, No. 4, 1927, p. 687.

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.62

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 valued in northwesternEurope for food
with landings for the years just preceding World
War II, running around 90 to 100 million pounds.

Barn-door skate Raja laevis Mitchill 1817

Bigelow and Schroeder, 1953, p. 217.
Garman, 1913, pi. 22, fig. 2, as R. stabuliforis.

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 1% inches on one
side, and 4% by 1% inches on the other in a speci-

11 Three mile hauls with the trawl sweeping a strip about 35 feet wide.

•' Examples are: 15 miles off Monhegan I., Maine, June 24-25, 1913, total
fish caught, 5,463; skates 170. Twenty miles 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, skates 62.

B Scattergood, Copeia, 1950, p. 169.

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 with
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 5 feet; it has been said to
reach 6 feet though there is no definite record of
one that large. One of 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 fish.
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 26% 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 Bigelow and
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 Guff 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 68° 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, butterfish, 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

63

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 winter. 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 7% 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 bails.

Occurrence in the Gulj 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 ocellata Mitchill 1815

Spotted skate; Winter skate; Eyed skate

Bigelow and Schroeder, 1953, p. 240.

Garman, 1913, p. 339, pi. 29, fig. 2, as Rata 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 each
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 fins are close together; the outer
corners of the pectorals are bluntly angular; the
claspers 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

4.

/.'•';

vtx

■

:&\%

&

l

gs

7

Figure 26. — Big skate (Raja ocellala), 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% feet in length, commoidy 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, cbub mackerel, butterfish, cunners,
sculpins, silver hake, tomcod, and hake.96

u From Vina] 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 36 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 are 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
Bay 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 1% 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 Gulf of St. Lawrence, and to the
southern part of the Newfoundland Banks.

Occurrence in the Gulf of Maine. — This, the
second in size of our skates, occurs commonly all
around the Gulf of Maine from Nova 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
about 18 percent (1,116) of the skates taken in 42
trawl hauls by the Eugene H, late June 1951, fish-
ing from Nantucket 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 New
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 cannot 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

• Copela 1951, No. 2, p. 169.
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, pi. 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
eyes, with 1 to 5 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 pectorals, 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 by 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
space on each side of the snout; it is white below.
It is most readily recognized by its color pattern,
with short dark bars as well as spots, 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 largest on record
was about 37 inches long.

General range.— OS the eastern coast of the
United States from Massachusetts Bay to both
coasts of Florida.

Occurrence in the Gulf of 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

Provincetown. 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 garmani Whitley 1939

FiOSETTED 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 tad 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 tad in young specimens,
increasing to 2 to 6 irregular rows ia 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,

87 Garman's illustration is of an abnormal specimen with three dorsal flna.

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 Qulj of Maine. — Our only
reason for mentioning this species is that one
specimen was trawled by the Albatross HI, May
14, 1950, at 52 fathoms southeast of Nantucket
Lightship Gat. 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; Summer 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 K 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, 17% 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. ocellata, 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 16 to 20 inches long;

the maximum recorded length is 21 inches (53
cm.); they weigh about % to 1 pound at 16 to 17
inches and anywhere from 1% to 2 pounds at 18
inches. Females mature sexually when 12%-17
inches (32-43 cm.) long, males at about 14 to
17K 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 75 fathoms or so. Many even follow
up the shelving bottoms of our beaches until they

88 Information supplied by Dr. Daniel Merrimau, Dr. Y. H. Olsen, the
Misses S. B. Wheatland and L. H. Calhoun, who have made a detailed
study of the littlo skate in southern New England waters.

FISHES OF THE GULF OF MAINE

69

strand. And tho 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 southcentral part of Georges Bank, in late
June 1951. Fifty fathoms (off the Bay of Fundy)
is, in fact, the deepest that positively identified
specimens are known, in the inner parts of our
Gulf; 80 fathoms off southern New England.69

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 tiny
objects as copepods have been found in their
stomachs. Probably crabs loom largest in their
diet, for more than 29 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 (Leptocheirus) 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, cunners, silversides, tomcod, silver hake,
have all been found in their stomachs, and they
bite a baited hook readily, affording amusement
to vacationists.

The spawning habits of the little skate have not
been followed in the Gulf of Maine. Studies, at
the Bingham Oceanographic Laboratory, however,
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

n Seventeen that we saw trawled on the Albatross III, May 1950.
" Trans. Roy. Soc. Canada, Ser. 3, vol. 12, sec., 4, 1918, p. 63.
71 Information from Dr. Daniel Merriman, Dr. Y. H. Olsen, and the
Misses S. B. Wheatland and L. H. 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 6 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 \){ to 1% inches by
about 2% to 2% inches, not 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 at the end that has the longer pair
of horns, is about 3% to 4 inches long at hatching;
its abdomen is still 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 Fundy descend to deeper
water the first winter, and this probably applies
to the Gulf of Maine as a whole.

It appears from information of various sorts
that a little skate 8 inches (20 cm.) long may be 1
to 1% years old; one of 11% to 12 inches (30 cm.)
2 to 3 years; one of 15% to 16 inches (40 cm.) 3
to 4 years; one of 19% 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.73

General range. — Atlantic coast of America;
southern side of the Gulf of St. Lawrence and

,3 This summary of the breeding habits is based on extensive information
supplied by Dr. Daniel Merriman, Dr. Y. H. Olsen, and the Misses S. B.
Wheatland and L. H. Calhound.

'3 Information from Dr. Daniel Merriamn, Dr. Y. H. Olsen, and the
Misses S. B. Wheatland and L. H. Calhoun.

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 of 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 prickles 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 prickles 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 20% inches long, Emerald Bank, Nova Scotia.

From Bigelow and Schroeder. Drawing by E. N. Fischer.

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 River in July and August; probably
they 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 skate76

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 (110-140°), 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

11 When the first edition of this book appeared, it was an open question
whether the thorny skate of American waters (named R. 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.

" Figure 1 of Garman's plate 21 is not of a thorny skate, as it is named to the
accompanying caption, but is of a small specimen of the big skate that wc have
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 radiala), female, about 31% inches long. After Garman.

on the other, and are rough with narrow cross-
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,
east 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

77 Doubtfully reported from Belgium and the Bay of Biscay.

offing of New York; and as a stray to the offing
of Charleston, S. C.78

Occurrence in the Gulf of Maine. — The thorny
skate is not often seen close inshore along our coast,
being restricted in general to moderately deep water
(p. 72). But it is now known to be generally dis-
tributed in the deeper waters of the Gulf. Thus
it 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 Ipswicb 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°10' N., long. 77°25' W., in 74 fathoms, by the Albatross
III is in the Museum of Comparative Zoology.

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 only 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
Gulf 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. Should 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, 19S3.

Sting ray Dasyatis centroura (Mitchill) 181580

Stingaree; Clam cracker

Bigelow and Schroeder, 1953, p. 352.

Garman, 1913, pi. 33, figs. 1, 2, aa Dasybatus marinus.

Description. — The most distinctive features of
this sting ray, among other Gulf of Maine fishes,
are its very long, whiplash-like tail without dorsal
fins, and the long, sawedged spine or spines with
which the upper side of its tail is armed. The disc
is rhomboid, about 1% to 1% 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 tail, 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 tail spine for a distance about as

80 This ray was mentioned as Dasybatus marinus and as D. hasiatus 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 1815, as Raja centroura.

FISHES OF THE GULF OF MAINE

75

Fionas 32. — Sting ray (Dasyatis centroura), male, about 55^ 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 smoo th skinned (except for the tail
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 tail but none on the upper side.

Size. — This appears to be the largest sting ray
of the western North Atlantic. The greatest

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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, though 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 i9
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 loDg tail armed with one or more
poisonous sa wedged 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 quadriloba.

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.

" This estimate is based on our own observations near Woods Hole.

11 The eagle rays, family Myliobatidae, and butterfly rays, family
Oymnurldae, are close allies of the cow-nosed rays; none of them has yet
been encountered in our Gulf.

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
back 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.89

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 of the head of one from Rio de Janeiro.

FISHES OF THE GULF OF MAINE

77

Figure 33. — Cow-nosed ray (Rhinoptera bonasus), about 22 inches wide, Newport, Rhode Island.

Schroeder. Drawing by E. N. Fischer.

From Bigelow and

Occurrence in the Gulf of Maine. — The cow-nosed
ray has even less claim than the sting ray to be
called a Gulf of Maine fish, for while schools of
them appear occasionally near Woods Hole where

145 of them were taken in the fish traps in one
day on one occasion, and while it is recorded from
Nantucket, it has never been seen, actually, east
or north of the elbow of Cape Cod.

DEVIL RAYS. FAMILY MOBULIDAE

The devil rays, like the sting rays (p. 74) and
cow-nosed rays (p. 76) have the pectoral fins
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 parts 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 they 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 gill clefts. Some of them are
the largest of the rays and among the largest of

fishes. Being tropical-subtropical in nature they
have no real place in the fish fauna of our Gulf,
but Manta, the largest of them all, has been known
to reach Georges Bank as a stray from warmer
latitudes.

Devil ray Manta birostris (Donndorff) 1798

Bigelow and Schroeder, 1953, p. 502.

Description.- — The so-called cephahc fins of the
devil ray, pointing forward, give it so distinctive
an appearance that it could not be confused with
any other fish, 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

it. WM

Figube 34. — Devil ray (Mania birostris), juvenile male, 11 feet 5H 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 of 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 192 1.84

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 are
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
only one known from within the geographic limits
of the Gulf, is deepest (one-sixth to one-seventh
as 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 probably 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 Gudger (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 31J4 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 17% 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 the 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 fining 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, 1930.8S 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 160 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
{"heterocercal") 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
tail 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.

« Reported by Firth, Bull. Boston Soc. Nat. Hist., 61, 1931, p. 9. It was 49
inches long and weighed 17H pounds dressed.

FISHES OF THE GULF OF MAINE

81

2.

KEY TO GULF OF MAINE STURGEONS

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.

The successive bucklers in the dorsal row are separated one from the next by spaces up to % 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 back (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 these are hard to handle, lower and
blunter on large. On the average there are 10
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 back
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 fin; 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 fin. And each shield in each of the two
lateral rows is separated from the next shield by
a space up to one-half as long as the shields.

The body is rather slender and rendered more
or less pentagonal in cross 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% feet long, depressed
below the level of the forehead, nearly flat below,

M It still is an open question, that we cannot answer, whether the sea
sturgeon of eastern North America is identical with the European sea stur-
geon, Is a recognizable race of the latter, or is a separate species; if the
last, its scientific name is Acipenser oxyrinchus Mitchill, 1816.

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

Color. — 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 New
England, ripe males are up to about 6 to 7 feet
in length, averaging 65 pounds in weight; the
spawning females (which are larger), up to about
10 feet and to about 250 pounds,88 with a larger
one taken from time to time. And the general
run was about the same in the Kennebec, 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 9 weighing between 350 pounds and 600

«' Vladykov and Beaulieu (Natural. Canad., vol. 73, 1946, pp. 143-204),
give a detailed account of the characters that separate the sea sturgeon from
the lake sturgeon (Acipenter fulvescens Raflnesque, 1817).

» According to Cobb, Rept. U. S. Fish Comm. (1899), 1900, p. 277.

*fr~^B^3^ ^si1-^*!*

Figure 36. — 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.

*• According to measurements and weights of 1,592 sturgeons by Vladykov,
Rapp. Gen. Minlstr. Chasse. PGch., Quebec (1948-1949), 1949, pp. 43-54.

•' Ryder, Bull. U. S. Fish Comm., vol. 8, 1890, p. 266.

« Ryder (Bull. D. 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 young 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)i 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.99

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 gained 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. (6% 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.

■< A series of 1,592 sea sturgeons from the lower St. Lawrence River, studied
by Vladykov (Rapp. Gen. Minstr. Chasse, P8ch. Quebec (1948-1949) 1949,
pp. 53-56) included a good representative of sizes from about 4 inches up to
90 inches.

M Prince reports a 6-rnch sturgeon from Hudson Bay (Rept. Sixty-seventh
Meeting, British Assoc. Adv. Sci., Toronto, 1897, p. 687).

•« Greeley, Suppl. 26 Ann. Rept. Conserv. Dept. New York, 1937, p. 82.

•» Vladykov (Rapp. Gen. Minlstr., Chasse, Pech. Quebec, 1948-1949,
pp. 61-63, 66, table 19).

•» 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 11 years for a 75-inch
sturgeon, and 12 years for two others of 88 and
100 inches are anywhere near the truth.09

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 Gvlf 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 are 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," 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

•• See footnote 98.

1 Two sturgeon 44 and 45M Inches long, netted in the Parker River at
Newbury, Mass. , July 23, 1933, are (or were) In the collection of the Boston
Boclety 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; in 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 M&j 1913; at the mouths of the
Essex and Ipswich Rivers, where jumping stur-
geon have been reported recently in the daily
press ; 8 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 Guff.

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.
» The Boston Herald, June 1950.

< Reports collected by the late Walter H. Rich, U. S. Bureau of Fisheries,
and notices in the dally press.

84

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

the catch was about 250 fish in 1880, yielding
12,500 pounds of meat, and not much less in 1898
(10,875 pounds). But the yearly landings were
only about one fourth as great there (2,777 pounds)
by 1919. And the reported landings of sturgeon
from the entire coastline of Maine (including what
few were brought in from offshore) had fallen to
only 300 pounds in 1940, and 400 pounds in
1947. Reported landings in Massachusetts of
5,300 pounds in 1940 (all by otter trawlers) and
of 6,600 pounds (5,000 pounds by otter trawlers,
from off shore), corresponding to some 50 to 70
fish, if they weighed as little as 100 pounds each,
will further illustrate their present-day scarcity.
We have never heard of a large sturgeon hooked
by an angler in the Gulf of Maine. But we hear
from time to time of a small one caught in this
way, as already remarked (p. 83) .5 And the
skill of a woman angler 9 who foul-hooked a stur-
geon about 6 feet long, and beached it on surf-
casting tackle after a long fight, fishing alone at
Wasque Point, Marthas Vineyard, on July 15,
1950, was widely heralded in the daily press.

Short-nosed sturgeon Acipenser brevirostrum
LeSueur 1818

Little sturgeon

Jordan and Evermann, 1896-1900, p. 106.

Description. — The little sturgeon resembles the
sea sturgeon so closely in general appearance that
we need note only the most obvious differences.
These are that the shields in its dorsal row are
relatively smaller, and that each is separated from
the next by a space up to % as long as the shields
themselves (successive dorsal shields in contact or
overlapping in the sea sturgeon); that the space
between its dorsal row of shields and the upper

• We once saw one small one about 23 Inches (575 mm.) long foul hooked In
the side on" South Beach, New York, December 21, 1923, and heard of a simi-
lar experience by the same angler a year later.

• Mrs. George T. Rice. About 30 others were seen by her at the same time
In a deep slew formed by a new bar.

lateral row on each side is only sparsely set with
fine prickles (closely set with coarse prickles in
the sea sturgeon); and that its viscera are black-
ish (pale in the sea sturgeon) ; also the number of
rays in the anal fin averages smaller in the little
sturgeon (19-22) than in the sea sturgeon (23-30).
The snout, too, is considerably shorter relatively,
as well as broader, than it is in young sea stur-
geons of equal size. And while the snout is about
as long, relatively, in the one species as in the
other whea they are full grown, sea sturgeons are
then so much the larger that there is no danger of
confusing the one kind with the other.

Color. — Described as blackish above, tinged
with olive above the upper lateral line of shields,
marked with alternate black and pale bands; sides,
below the upper lateral row of shields, reddish
mixed with violet; abdomen white.7

Size. — This is a much smaller fish than the sea
sturgeon. Males may mature when only 19-20
inches long and most of them do by the time they
pass 21 inches; most of the females at about 24
inches. The largest so far recorded is one of about
36 inches, in the Museum of Comparative Zoology.
One about 31 inches long weighed 7 pounds 4
ounces.8

Habits. — Nothing is known of the habits of the
little sturgeon except that it spawns in rivers and
that it does so late in April in the lower Hudson.
The fact that fair sized specimens are taken there
in summer and also in winter, suggests that it
may not be as regularly migratory as the sea
sturgeon is.9 But the places of capture of the
Gulf of Maine specimens mentioned below show
that some certainly go out into the open sea and
wander for some distance from their parent stream.

General range. — So far as we know, the only

We have not seen a fresh-caught specimen.

■ For further details as to size, see Ryder, Bull. U. S. Fish Comm., vol.8,
1890, p. 238; and Greeley, Suppl. 26 Ann. Kept. Conserv. Dept. New York,
1937, p. 69, table 11, pp. 82, 90.

• Greeley, Suppl. to 26 Ann. Rept. Conserv. Dept. New York, 1937, p. 90,
makes this suggestion.

Figure 37. — Short-nosed sturgeon (Acipenser brevirostrum), Woods Hole specimen. From Goode. Drawing from a

photograph.

FISHES OF THE GULF OF MAINE

85

locality records definitely belonging to this species,
not to young sea sturgeons, are from Province-
town and Waquoit, Mass.; from the Hudson
River, N. Y. ; from Delaware Bay and River;
and from Charleston, S. C.

Occurrence in the Gulf of Maine. — The only
recent record of the little sturgeon in the Gulf is
of one about 23 inches long, taken at Province-

town about 1 907 and now mounted in the Museum
of Comparative Zoology.10 The Museum of the
Essex Institute, Salem, also has — or had — a
stuffed sturgeon from Rockport, Mass., identified
as this species by Goode and Bean.11 Evidently
the sturgeon is now very scarce in our Gulf and
there is no reason to think that it ever has been
more plentiful there.

The Herring and Tarpon Tribes
FAMILIES CLUPEIDAE, DUSSUMIERIIDAE, AND ELOPIDAE

The true herrings (Clupeidae) are soft-finned
fishes wholly lacking spines, with one short dorsal
fin, deeply forked tails, ventral fins situated on
the abdomen far behind the pectorals, teeth small
or lacking in adults, deep bodies flattened side-
wise, and large scales that slip off at a touch.
They are, perhaps, the most familiar of northern
sea fishes and certainly are the most abundant in
number of individuals. Seven species of herring
occur in the Gulf of Maine — the hickory shad (not
very common), the sea herring, the alewife, the
blueback, and the shad (regular and plentiful),
thread herring (scarce), and the menhaden (irreg-
ular in its occurrence) . The shad, menhaden, sea
herring, and thread herring are easily named ; but
the alewife and the blueback resemble one another

so closely that they are often confused, even by the
fishermen who handle them constantly. The round
herrings (Family Dussumieriidae) differ from the
true herrings chiefly in tbeir rounded bellies and
less deep bodies. The members of the Tarpon
Tribe (Family Elopidae) are very closely allied
to the true herrings (Clupeidae), from which they
differ in having a bony plate on the throat between
the branches of the lower jaw. There are only
about five species, all of them tropical. Two
are known from the Gulf, as strays.

10 This Museum also has another of about 36 Inches from Waquoit, on the
southern shore of Massachusetts.

" Bull. Essex Inst., vol. 11, 1879, p. 27. A sturgeon was reported as brt-
virostris from Boston Harbor many years ago, but there is no way now of
checking the Identification.

KEY TO GULF OF MAINE HERRINGS AND TARPONS

1. Last dorsal fin ray prolonged 2

Last dorsal ray not prolonged 3

2. Dorsal fin originates in advance of the ventrals; scales only moderately large Thread herring, p. 112

Dorsal fin originates behind the ventrals; scales very large Tarpon, p. 87

3. Belly rounded 4

Belly sharp edged 5

4. Scales very small; mouth very large with upper jaw-bone extending considerably beyond the rear edge of the eye;

point of origin of dorsal fin about over that of the ventral fins Ten pouDder; p. 86

Scales large; mouth small, with upper jaw-bone extending rearward only about as far as the front edge of the eye;
point of origin of dorsal fin well in advance of that of the ventral fins Round Herring, p. 87

5. Head (tip of snout to edge of gill cover) very large, occupying about one-third the total length of the body to base

of the central rays of the caudal fin; free edges of scales fluted, not rounded Menhaden, p. 113

Head about one-fourth the total length of the body; free edges of the scales rounded 6

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

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 Elops saurus 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 throat 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.11 Ten
pounders are taken from time to time near Woods
Hole.

'« Jordan and Evermann. Bull. 47, U. 8. Nat. Mus., 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
8chroeder (Copela, 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-like 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
the presence of the bony plate on the throat men-
tioned above (p. 85) in the characterization of
the famdy 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.

Color.— 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.18 Its chief center of abundance is in the
West Indies, about Florida, and in the Gulf of
Mexico.

Occurrence in the Gulf 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.
i* Halkett, Check List, Fishes Canada, Newfoundland, 1913, p. 46.
i' Radcliffe, Copeia, No. 26, 1916, p. 3.

Figure 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.

Color. — 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." 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 the U. S. Bureau of Fish-
eries.

" Reported in the newspapers.

» Reported by Leim, Progr. Rept. 21, Atlantic Biol. Sta. Fish. Res. Bd.
Canada, 1937, p. 5; and by McKenzie, Proe. 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 grayish or shading
into green or blue.

Size. — Herring grow to a length of about 17
inches and to a weight of about IK 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-

sl Rept., Norwegian Fish. Mar. Invest. SkriJter, vol. 9, No. 11, Eept. 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," 26 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.26 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 (Meganyctiphanes 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.

M 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 their
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 light.
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. Kept. 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 growth. — 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 3X 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

W Rept. TJ. S. Comm. Fish. (1896), 1898, p. 402.
» A ge and growth of the herrings In Canadian waters.
1914-16 (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. The
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, Rept. U. S. Comm. Fish. (1896), 1888, p. 412.
» Rept. Biol. Board Canada (1930), 1931, p. 23.

Figure 42. — Eggs of the herring (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 stages.

The sea herring is about 5 to 6 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. (1% 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. This 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 3% 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.36 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 in 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.38 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 between 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 160 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) tor an account of
age determination by analysis of the scales, as applied to the herring.

'* Rapp. and Proc. Verb., Cons. Intornat. Erplor. 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 fRapp. 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 9% inches long; usually not until they
are 10 to 10% 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 Gulf of Maine. — To fist 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 38 may be set at about their lower limit.

The distribution of commercial catches, plotted
by Needier (fig. 44) 39 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 of Fundy in May.

" A reliable index, for the herring is a valuable fish.

94

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

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Figtjre 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
flsh 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,**
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.46 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

44 Coast sectors of comparable length.

« Average catches per haul about 56 fish at 22 to 40 fathoms, and 28 at 41 to
50 fathoms, but only 6 at 51 to 60 fathoms.

« W. F. Clapp found no herring in many cod and haddock stomachs
examined by him on Qeorges 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 winter, 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
there 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 they
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 Qeorges Bank, off Seal Island; off Yarmouth,
Nova Scotia; near Macblas, 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.*8

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

Founds Month Founds

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,

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-
rity. 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 w on the average monthly landings of
large herring for Charlotte County, for the period
1920-1931, to the nearest 1,000 pounds, follows:

Month

January

February. .
March

48 No particular attention is paid to sperling around Massachusetts Bay,
for they are too small to be In demand for bait, and they are not plentiful
enough (or not concentrated enough) to support a sardine fishery there.

Pounds Month Pounds

132,000 July. 1,065,000

164,000 August 4,334,000

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 221 623,550

June - -. -. 0 88,657

July 0 0

August 0 1,000

September.. 0 57,287

October 0 9,526

November 0 176,435

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.81

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

11 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.62 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 winter
pasture for them.

Studies carried out from the Atlantic Biological
Station at St. Andrews, chiefly under Dr. A. G.
Huntsman's M leadership, and by the International
Passamaquoddy Fisheries Commission64 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 differential 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 journeys more intelligently, and
because any directive swimming they may carry

u Huntsman. James Johnstone Memorial Vol., 1934, p. 82.
•» For summary, see Huntsman, James Johnstone Memorial Vol., 1934,
pp. 95-96.
« See Graham, Jour. Biol. Board Canada, vol. 2, 1936, No. 2, pp. 93-140.

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 spawners.
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,65 but not commencing
until early August, and ending by early Oc-
tober in others.66 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 Ipswich67 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.

69 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
them, 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.69

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
1890, 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 in 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, 3 1 ,988, 132 pounds, and 32,844,500 pounds,

» Moore, Rept. U. S. Comm. Fish. (1896), 1898, p. 430.

•° Based on Canadian fishery statistics.

•i Earll (Fisheries and Fish. Ind. U. S., sect. 5, 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 Matlnicus 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 offish; 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.*3
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
souDding 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. Scl. Rept. No. 67,
1949, p. 8, for further details.

Figtjbb 45. — Hickory shad (Pomolobus mediocris) , Chesapeake Bay region specimen.

Todd.

From Goode. Drawing by H. L.

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, cunners,
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.94 But it appears not
to do so in the streams tributary to Chesapeake

M Smith (N. C. Geol. 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 possibility that the
"green" fish found in Chesapeake Bay, leave the
Bay, perhaps to spawn in salt water.66

General range. — Atlantic coast of North America
from the Bay of Fundy to Florida.

Occurrence in the Gulf of 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
[approximate 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 13K 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 Schrocder, Bull., U. S. Bur. Fish., vol. 43, 1928, p. 84.

M The Museum of Comparative Zoology received one from this run from
Dr. J. O. 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.

Todd.

From Goode. Drawing by H. L.

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 Whitbome, In "The True Travels of Oapt. John Smith,"
1S16, vol. 2, p. 260.

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
herriDg, 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;

M The average number of eggs In 644 females taken In the Potomac was
102.800 (Smith, N. C. Qeol. and Econ. 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. Biol. (1902-1905), 1907, p. 96).

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
i)i 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 years
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. 91i

" 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, laun.ce, 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 7i 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 its 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° 68' 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.7'

Occurrence in the Gulj 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 78 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 shall 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

'• 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. 3S) and
by Bailey (Biological Survey Merrimac Watershed, New Hampshire Fish
and Oame Dept., 1938, p. 162).

" Belding (Ropt. 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.7*

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 iD
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

'* 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 the
status of the alewife in Massachusetts streams.

" Stocked with 28.000 adult fish in 1949, and fish ladders under construction.

■' Eept. 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 M 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.86 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, Kept. Biol. Board Canada (1931) 1932, p. 34.

13 A special study of the alewife fishery was made for that year, seo Smith,
Eept. U. S. Comm. of Fish. (1896) 1899, pp. 33^13.

•* The Canadian catches for the year were reported in barrels; the conver-
sion factor used is 200 pounds per barrel.

** Reported catches for 1896 were about 4,234,000 pounds for the St. John
River system; 1,390,612 pounds for the Damariscotta River, 385,804 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 884,255 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.88

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. They
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.

» McKenzie, Kept. Biol. Board Canada (1931) 1932, p. 34.

87 5,051,100 pounds and 4,517,500 pounds, respectively.

M 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) 1921,
p. 140.

Blueback Pomolobus aestivalis (Mitchill) 1815

Glut herring; Summer herring; Blackbelly;

Ktack

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 3% 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 sooty 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.

Figure 47. — Blueback (Pomolobus aestivalis), Chesapeake Bay region specimen. From Goode. Drawing by H. L.

Todd.

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°.w 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 adult. 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 II on March 5, 1931 about
100 miles off 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.

Qeneral 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

H The early development and larval stages of the blueback are described
by Kuntz and Radclifle (Boll. U. S. Bur. Fish., vol. 35, 1918, pp. 87-134).

" In Chesapeake Bay, Hildebrand 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 3^ 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: M it spends most of its life in
salt water but runs up into fresh water to spawn.

Occurrence in the Gulf of 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.83 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 3% to bVt 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.

« Dr. A. H. Leim has sent us four typical bluebacki about 12 lnchss long ,
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]

JordaD 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
all, 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 M shad weigh about % pound at 8 inches;
about % pound at 12 inches; about IK pounds at
15 to 16 inches; about 2% pounds at about 20
inches; and about 4 % pounds at 23 to 24 inches,
though with variations according to their condi-
tion. Adult males weigh from IK to 6 pounds;
females from 3K to 8 pounds. Shad are occasion-
ally reported to 12 pounds, and the older writers
mention shad of 14 pounds, 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 their
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.

Figdee 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 " 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." 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.l 10) .

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 fisb as late 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.*

•» Contrib. Canad. Biol., N. Ser., vol. 1, 1923, p. 310.

" Contrib. Canad. Biol.. N. Ser., vol. 2, 1824, p. 231.

» Leldy (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.

< Bean (Bull. 60, Zool., vol. 9, New York State Mus., 1903, p. 207) com-
mented on this long ago.

• Accounts of the breeding habits of the shad have been given by Ryder,
Kept. U. S. Comm. Fish. (1885) 1887, p. 623; 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% 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 Gulf 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 : 8

• Leim (Contr. Canad. Biol., N. Ser., vol. 2, No. 11, 1924, p. 195) gives a
detailed comparison of shad with alewife larvae.

1 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.

• Frcm Stevenson, Rept. U. S. Comm. Fish. (1898) 1899, p. 262.

Number of shad caught, Number of shad caught.

Year reported, or estimated Year reported, or estimated

1789... 830,000 1888 None

1805 540,000 1889 18

1835 365,000 1890-1892 None

1865 50,000 1893 2,020

1871-1873 (aver- 1894 2,750

age) 1,942 1895- — 94

1880 2,139 1896 7

1885 130

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 Fundy; perhaps the St. Croix; 10
the only Maine rivers that see regular run9 of a
few shad are the Nonesuch and the Sheepscot."

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 13a 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

« 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
1816, according to investigations by H. F. Taylor of the U. 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. O. E. Atkinson, U. S. Fish and Wildlife Service.

» Reported to us by John B. Burns of the Massachusetts Division of
Fisheries and Game.

« 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 GTTLF 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 m other rivers there).

Immature shad, up to 2 to 2 K 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.16

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

11 502 barrels (about 100,400 lb.) were taken In one set of mackerel pounds
at Provincetown in June 1910; tbe 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 15W inches long were
taken in one set of traps at Barnstable, on Cape Cod Bay, October 3, 1950.

>• 135,000 pounds of these large spent flsh 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.

18 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 sei organs.

" About 250,000 pounds were brought in to the local freezers yearly in 1913,
IOH. 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 Fundy 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 near 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, N. 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 the 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, how 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. Hollis 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 shad
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 in racial characters for
recognition when taken at sea, but this is a ques-
tion for the future.2*

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-

51 See Vladykov, Nat. Canad., vol. 77, 1950, p. 121, for a detailed account
of his tagging experiments on St. Lawrence River shad.

*> Vladykov, Copeia, 1936, No. 2, p. 168, reports bet-ween 25 and 30 shad of
4-6 pounds, taken per haul, by otter trawlers in March, 1035, southwest of
Middle Ground, about lat. 44*25' N„ long. 61°05' W.. at about 50 fathoms.

n Two shad were trawled by Albatross ///on the eastern part of Nantucket
Shoals at 68 fathoms, and 46 others at 9 stations distributed thence westward
to the offing to Montauk Point (long. 71°52' W.) at 26-64 fathoms, May 11-18,
1950.

It w Vladykov and Wallace (Trans. Amer. Fish. Soo., vol. 67, 1937-1938, pp.
52-66) believe that Sbubenacadie, Delaware River and Chesapeake Bay
shad differ significantly in average number of vertebrae, of mid ventral
scales and of pectoral fln 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 ; x
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

■' Stevenson, Rept. U. S. Comm. Fish (1898) 1899 pp. 265-269.

18 These catches were reported as "barrels" presumably of 200 pounds each.

" Stevenson, Rept. U. S. Comm. Fish (1898) 1S99, p. 121.

« About 460,000 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 1916-17.

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 }i 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}i 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
Brazd, straying northward to Chesapeake Bay,
and occasionally as far as southern Massachusetts.

Occurrence in the Gvlj 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 1885. 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. 29 Being a tropical fish, it is not apt to
reach the Gulf except as the rarest of strays.

Menhaden Brevoortia tyrannus (Latrobe) 1802
Pogy; Mossbtjnker; 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 MscCoy, Bull. Boston Soo. 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, witb 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. — Adult 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 fe«d 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. pla. 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 which 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

" 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 their 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,
suggest that the menhaden is a summer spawner
there also. We have found no eggs in our tow-
nettings north of Cape Cod (young fry were taken
in abundance in Casco Bay in October 1900), prob-
ably because our work there was carried on during
a series of poor menhaden seasons. From Chesa-
peake Bay 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 pilchardus) , but
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

a By Kuntz and Radcliffe. Bull. U. 8. 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.34

Occurrence in the Gulf 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,
1912, were SH to 4 inches (91-99 mm.) long; others taken in Salt Pond, Fal-
mouth. Mass., on November 24, 1949, were 4H to 6 inches long.

« See Hildebrand (Smithsonian Misc. Coll., vol. 107, 1948. No. 18 for a
revision of the genus Breroortia). One named species, P. brericaudala
Goode 1878, is known only from Noank, Conn.; we doubt its validity.

••According to Huntsman (Contr. Canad. Biol., (1921) 1922, p. 69) 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 1891 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.
They 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, Copela, 1961, pp. 93-94.
• Reported by Scattergood, Trefethen, and Coffin, Copela, 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 M 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, partic,
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. New York State Mas., 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; but
they are easily distinguishable from the herrings
by the fact that their 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 fn 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 (Slolephorus 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

■ 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 3K inches long.

General range.- — Coast of the United States
from Maine to Texas, chiefly west and south of
Cape Cod.

10 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) 1758

Jordan and Evermann (Stolephorus brownii), 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; a 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, 1930.43 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), Somers 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

u Bulletin. U. 8. Bur. of Fish., vol. 33, 1915, p. 13.

tribes (p. 141).4* 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, 193S, p. 3).
« Kendall, Bull. Boston Soc. Nat. Hist., No. 58, 1931.
" Sundry other deep-sea fishes have adipose fins.

120

FISHERY BULLETIN OP THE FISH AND WILDLIFE SERVICE

pearlsides, while the lantern, viper, and lancet
fishes are of different general aspect.

Four salmons 46 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 Salvelinus jontinalis (Mitchill) 1815
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 specimon of one of the whitefishes (probably Coreoonus guadrilatiralis
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. Whitensh 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,49 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 the 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.

Figure 53. — Brook trout (JSalvelinus fontinalis) , about 15% 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 the 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 fishing. These, however, he
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

«I Rept. U. S. Comm. Fish., (1904) 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 (Salvclinus alpinus) in northern
Canadian waters which is very plentiful along the coast of northern Labrador.

210941—6

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 hi 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 Nova Scotia streams
tributary to the Bay of Fundy (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 w 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.

Description. — 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 ventrals 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 are taken about Woods Hole, 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 83 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 thesalmon
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 spawns in fresh water.

The salmon are silvery and very fat when they
enter fresh-water on the spawning journey, but

" Huntsman (Bull. Biol. Board Canada, 21, 1931) has published an exten-
sive study of the life history of the salmon of the Maritime 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 creature
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 fife 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 finger 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, 1938, p. 65-88.

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 Mlramlchi 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 greedily 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 % 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.

i 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 Oulf of Maine fish in
particular, with references. Etchelbaum (Cons. Perm. Internal. 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 they 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 Gulf 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" fly; 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.

» 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 grilse
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 Soc. Nat. Hist., vol. 9, No. 1, 1935, p. 32.

" This is the southern European limit given by Berg (Zoogeographies.
vol. 1, Pt. 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. Sor., 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 Salvelinus
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 M 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 beeu
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, 1943, 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, Kept. 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,8* 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

M A few small "salmon" reported of late In the Merrimac probably were
the landlocked form, running down from tributaries stocked with this flsh.

■* 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.

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

'■ Kendall (Mem. Boston Soc. Nat. Hist., vol. 9, No. 1, 1935. pp. 31-33) lists
a number of such cases.

'» Huntsman, Ann. Rept. Fish. Ees. Board Canada, (1947) 194S, p. 37, and
unpublished notes.

" The average was only 3,000 pounds (perhaps 300 flsh) 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

N 1933, 1935, 1937, 1938, 1939, 1940, 1943, 1944, 1945, and 1946.
" 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 Delaware77 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.7'
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 run 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

i» Smith (Bull. V. 9. Fish. Comm., vol. 14, 1896, p. 99) reports salmon
seined among some mackerel off Delaware in 1893.

» Bull. 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) 1948, 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 85 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.88

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.

M 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-fifth 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
Rivers.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 m 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 Soe. Nat. Hist., vol. 9, No. 1, 1935, pp. 58-60)
for age determinations of Penobscot salmon.
•' Radclifle. Rept. U. S. Comm. Fish. (1921) 1922, p. 146.
* 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

Number
offish

Pound!

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 95 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.
- 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

1880-1880 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,88 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 well 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-
are 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.

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 flaplike 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 salmon 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 third 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 5%
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 salmon 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-

« More detailed accounts of these and successive plantings will be found
in tho annual reports of the U. S. Commissioner of Fisheries for the years
1914 to 1928.

•> Reported catch, Washington County, Maine, 1920, 310 pounds.

« It was forwarded to the Massachusetts Commissioners as reported by
C. E. Orant of Gloucester.

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 on
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 Gulf
of 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. 8. Comm. Fish. (1928), Pt. 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.

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 (Miiller) 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-
fine of the belly in the female. In males, too, the
scales in one of the longitudinal rows immediately
above the lateral fine, 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 7%
inches long.

Habits.3 — 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,

• Interesting accounts of the habits of the capelin and of Its rate of growth
In Newfoundland waters have been given recently by Jeffers (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. V. S. Comm. Fish. (1872-1873) 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. Jeff era 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.8

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 fieh 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.

• Tompleman (Bull. Newfoundland Government Lab., 17, Res., 1948>
figs. 18-20) gives a series of eicellent illustrations of capelin larvae at different
stages of growth.

8 According to Jordan and Evermarm the capelin finds its southern limit
at Cape Cod, but we find no actual records of its occurrence farther south than
Is mentioned.

' Quoted from a 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 do 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 (Mitchill) 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
(Copula, No. 42, 1917, pp. 28-30; and Copela, No. 73, 1919, pp. 70-71) give
details.

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FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figube 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. eptrlanus) smells so strong that It Is not held In
very high esteem as a food-fish.

i» Kendall (Bull. V. 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, also 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 five 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 mnmmichogs
(FundvJus) as bait; while dinners, 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." 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

H This method of feeding seems first to have been doscribed In print by
'•Orif' (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 brackish
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 time 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 Eiver, a stream empty-
ing into Boston Harbor, when the temperature of
the water rises to about 40-420.13 This may take
place as early as the first week in March or as
late as the last, about Massachusetts Bay, 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, which 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.

"Atkins (Fish. Ind. U. 8., sect. 5, vol. 1, 1887, pp. 690-693) gives much
Information on the smelt In Maine.

» Kendall (Bull. U. S. Bur. Fish., vol. 42, 1927, pp. 231-233) summarizes
these observations and elves additional information for streams on tho 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

» Kept. U. S. Fish Comm., 1897, p. 188.

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FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

taken in Weir Kiver, 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 smelt 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.1"

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.

18 Smelt fishing has long been restricted to hook and line along this part
of the coast.

*> Maximum 675,700 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, wa»
2.258,030 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 argentines 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 silus 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 silus Is described, and records along the
American coast are given by Kendall and Crawford (Jour. Washington
Acad. Sol., 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 only 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 argeutine is a larger fish than the
smelt or the capelin, growing to a length of about
18 inches.

Habits. — Nothing is known of the life 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 untd the larva is about
50 mm. long.

General range.— North Atlantic, usually in water
as deep as 80 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 Gulf of Maine. — The argentine
was considered rare in our waters until recent-
ly. Some specimens have been brought in from

» For recent records of argentlnes off Nova Scotia, see McKenzle and
Homans, Proc. Nova Scotia Inst. Scl., vol. 19, No. 3, 1938, p. 277 and
McKenzle, Proc. Nova Scotia Inst. Scl., 20, 1939, p. 16.

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. Firth24 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., 61. 1940, p. 10.

FISHES OF THE GULF OF MAINE

LUMINESCENT FISHES

141

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.

KEY TO GULF OF MAINE LUMINESCENT FISHES

1. Trunk at least Yi 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

2. Mouth does not gape back as far as the eye Pearl sides, p. 144

Mouth gapes back beyond the eye 3

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

4. The rayed dorsal fin is far in advance of the ventrals; the jaws are armed with long and conspicuous

fangs 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

5. Eyes very small; no adipose fin behind the rayed dorsal fin; anal fin reaches nearly to the base of the

caudal Cyclothone, 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 fin and the origin of the tail fin 6

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 Lanternfish, p. 143

7. 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

8. Each side has only about 68 luminescent spots; there 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 aearly 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 tbe 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.26

« Parr (Bull. Bingham Oceanog. Coll., vol. 3, art. 3, 1928), and Tanlng
(Vldensk. Meddel., Dansk Naturhlst. Forenlng, vol. 86, p. 49, 1928) have
recently published critical synopses of the lantemflshes.

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.2*

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.28

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, acid 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 3K 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 (Ooode 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 Aethoprora effulgent.

» 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. opallnum.

»> It Is likely that Myctophum alaciale will be found In the Oulf 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. affine very closely In appearance, and In the general arrange-
ment of the luminous organs, but d InYrs from It In that one of the luminescent
spots above the base of the ventral fin 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 in 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
and dorsal fins; its eye very large; its lower jaw pro-
jecting; its mouth oblique; and both its jaws armed
with minute teeth. The dorsal fin (about 1 1 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, Osburn, and Cole, Bull. U. S. Bur. Fish., vol. 31, Pt. 2, 1913,
p. 743.

" Smltt, Scandinavian Fishes, vol. 1, 1892, p. 933, pi. 44, flg. 3.

" Ooode and Bean (Smithsonian Contrlb. 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 of Maine. — The known
occurrences of the pearlsides in the Gulf have been
few. Storer 36 (1867) 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., 1887, p. 160, as Scopelus humboldtil.

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 Chavliodus 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
these if one looks closely, for the viperfish has an

» Coi (Bull. Nat. Hist. Soc. New Brunswick, 14, 1896, append., p. 65)
reported one found dead there, on the shore,
a 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 tbey project when the
mouth is closed. Furthermore, the snout is so
short that the very wide mouth gapes far back of
the eye. 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 rays) 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.3'

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 Gulj 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 " 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 being 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, Tlefsee-Flsche Wlssensch. Ergeb. Deutschen Tlefsee-Exped.,
(1898-1899) 1906, vol. 16, 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.

« Qoode and Bean (Smithsonian Contrlb. Knowl., 1895, p. 97) Ust these
captures.

Figure 64. — Cyclothone (Cyclothone aignata) . After Brauer.

FISHES OF THE GULF OF MAINE

147

irregular flecks and dots along the back and on
the gill covers.41

General range.- — This is an oceanic fish, very
abundant in temperate latitudes in the Atlantic
where it lives pelage 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 fight organ
below each eye.

The tip of tbe 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

a For detailed accounts and colored Illustrations see Garman (Mem. Mus.
Comp. Zool., vol. 24, 1899, p. 246, pi. J, flg. 3), Brauer (Wlssonsch. Ergeb.
Deutschen Tiefsee-Eiped. (1898-1899), 1908, vol. 15, Pt. 1, p. 77, pi. 6, flg. 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 fight 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,*3 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." 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.**

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 ia
swollen at the tip and has two additional filaments
on ODe 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

« Murray and HJort, Depths of the Ocean, 1912, pp. 603, 611, 629.
» For a list of these stations, see Goode and Bean, Smithsonian Contrlb.
Know!., vol. 30, 1898, p. 107.
<• This specimen Is now In the Museum of Comparative Zoology.

Figure 65. — Stomias (Stomias ferox) , Banquereau Bank. From Goode and Bean. Drawing by H. L. Todd.

148

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

--»<■■>--?"*

Figoee 66. — Stomioides nicholsi, Browns Bank. Drawing by Myvanwy M. Dick.

Stomias. Stomioides resembles Stomias in all other
respects so closely that should a specimen of either
be taken, that is not easily identified, we suggest
forwarding it to the U. S. Fish and Wildlife Service
for naming.

Color. — Black below as well as above, the
luminescent organs showing as whitish dots.

Size. — The only specimen yet seen is about 10%
inches long, from tip of snout to base of tail fin.

Range and occurrence in the Gulf of Maine. — The
only known specimen of this species was taken
from the stomach of a swordfish harpooned from
the schooner Barbara, Capt. C. A. Turner, on the
southeastern edge of Browns Bank," over the
250 fathom line, August 3, 1932. Presumably it
had strayed from the mid-depths offshore.

Trigonolampa miriceps Regan and Trewavas 1930

Regan and Trewavas, Danish Dana expeds. 1920-1922,
No. 6, 1930, p. 55, pi. 1, fig. 1.

Trigonolampa resembles Stomias in general
appearance, in the relative sizes and locations of
the fins, and in having a long fleshy barbel on its
chin. But it not only has a small light organ below
the eye (as in Stomias), but also has a small
luminescent patch close behind it, and likewise a
larger triangular patch extending from close behind
the eye back across the top of the cheek; these are

••This specimen, described by Parr (Copela, 1933, p. 177), Is now In the
Museum of Comparative Zoology.

its most distinctive characters. The one species of
the genus yet known differs further both from
Stomias (p. 147) and from Stomioides (p. 147) in a
considerably deeper body (cf. fig. 67 with figs. 65,
66) ; also in that the tip of its lower jaw does not
enclose the tip of its upper jaw when the mouth is
closed; that the point of origin of its dorsal fin is in
advance of its anal fin by a distance about as great
as the diameter of the eye; and that it has only
about 68 light organs in each of its ventral rows, as
against 85 or 86 in Stomias (p. 147).

Color. — Not known, but probably black or very
dark brown.47

Size.- — The largest specimen yet seen (in the
Museum of Comparative Zoology) is about 9
inches (230 mm.) long to the base of the caudal
fin.

Range and occurrence in the Gulf of Maine. —
Only three specimens have been seen yet. The
first was taken in the eastern Atlantic by the
Danish research vessel Thor in 1906 at a depth of
about 600 fathoms; a second was found by Capt.
John Toothaker in the stomach of a swordfish
harpooned on the southern edge of Georges Bank
in the summer of 1922,48 and a third, now in the
Museum of Comparative Zoology, was recorded
simply as taken on Georges Bank about 1913. It
reaches the slope of our outer Banks only as a
stray from the mid-depths offshore.

" One that we have seen Is brown below as well as above wherever the skin
Is intact, with the light organs showing as darker dots.

" Parr (Copela, 1933, No. 4, p. 178) has given a detailed description of this
specimen, which is new in the Museum of Comparative Zoology.

^■^gf-gl-la;'^' PI''1 SPPgggjg

— — ^m^- ^

^^^w

Figure 67. — Trigonolampa miriceps. After Regan and Trewavas.

FISHES OF THE GULF OF MAINE

149

HATCHET FISHES. FAMILY STERNOPTYCHIDAE

These are deep, thin, flat-sided little fishes, with
various spiny projections, large oblique mouths
with small teeth, large eyes which are directed up-
wards in some of them but sidewise in others, and
ventral fins placed far back. Some of them have
an adipose fin behind the rayed dorsal, but others
do not. All of them are silvery, and all of them
have series of large and conspicuous luminescent
organs on the lower part of the body. They are
to be found in the mid-depths in all oceans, some-
times in great abundance.

Silver hatchetfish Argyropelecus aculeatus
Cuvier and Valenciennes 1849

Jordan and Evermann, 1896-1900, p. 604, as A. olfersi
Cuvier, 1829.

Figure 68. — Silver hatchetfish (Argyropelecus aculeatus).
After Brauer.

Description.- — This little fish is of so bizarre an
appearance that once seen it could hardly be mis-
taken for any other species yet known from our
Gulf, or for any that is likely to stray thither,
unless one of its own tribe.49 Its body is very
thin sidewise, with its forward part a little less
than three-fourths (70 percent) as deep as it is
long from snout to base of tail fin, but with the
ventral contour bending upward abruptly about
midway of its length in characteristic contour, so
that the rear half is much less deep than the for-
ward half. This break in the ventral contour is
marked by two short bony spurs, which are out-

*• The several known species of Argyropelecus resemble one another so
closely that their Identification calls for a specialist. They have been re-
viewed by Schultz, Proc. U. 8. Nat. Mus., vol. 86, 1938, pp. 147-153. The
most detailed description of this particular hatchetfish, with the best Illus-
tration (copied here as fig. 68) Is by Brauer, Wlss. Ergeb. Deutschen Valdicta
Tlefsee-Exped., vol. 18, Pt. 1, 1908, p. 110, fig. 47.

growths from the pubic bones, and there is a short
single spur (outgrowth from the pectoral arch) in
front of them in the mid ventral line.

The eyes are large, so high up that the space
between them on the top of the head is very
narrow, and they are directed more upward than
sidewise. The mouth is noticeably large, with
wide gape, and it is so strongly oblique that the
upper jaw is nearly vertical.50 The tips of the two
jaws are about even one with the other when the
mouth is closed, and both jaws are armed with a
large number of tiny sharp teeth. The dorsal fin
is short, about midway of the fish, and of two
parts, separated by a deep but short notch. The
forward subdivision is in the form of a hard,
triangular plate (apex rearward) supported by 8
or 9 hard spines, the rearmost of which is the
stoutest and longest. The rearward subdivision
is supported by 9 soft rays, that are bifid toward
their tips. The adipose fin is long and low. The
pectorals are as long as about two-fifths the
greatest height of the body. The ventral fins,
each with 6 soft rays, stand close behind the break
in the ventral contour of the body, and they are
connected with the anal fin by a thin transparent
ridge. The anal, commencing about under the
rear end of the base of the soft rayed part of the
dorsal, is notched midway of its length ; its forward
part is supported by 7 rays close together, the rear
part by 5 shorter rays spaced more widely. The
caudal fin is forked. A noticeable feature is that
the ventral edge of the deep forward part of the
body, from the pectoral spur to the pubic spines,
is sharp, with a series of 12 hard, plate-like scales
or scutes, that extend for some distance up the
sides, each slightly overlapping the next rearward,
and the profile is saw-edged between the ventral
and anal fins.

The hatchetfishes are provided with a complex
system of conspicuous light-producing spots. The
species aculeatus has one row of 12 very low down
along each side of the deep forward part of the body ;
also, a second row higher up consisting of 6 in front
of each pectoral fin, 2 along the base of the pectoral,
6 between pectoral and ventral fins, 4 between the
ventral and the anal fins, 6 along the anal, and 4
very small ones between anal and tail fins. There
is also one light-organ a little below and behind

1 Most of the published Illustrations of Argyropelecus fall to show this.

150

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

each eye, 2 on the lower part of the gill cover on
each side and about 5 on the lower jaw on each side.

Color. — Perhaps the most conspicuous feature of
the hatchetfishes, as taken from the water, is that
their entire bodies are glistening silvery. On some
specimens the silver is underlaid with velvet black
over the truDk as a whole; on others the black un-
der pigment is confined to a marginal band, broader
or narrower. The luminescent spots are pale yel-
low or white.

Size. — Maximum length probably not more than
3 inches or so.

Range and occurrence in the Gulf oj Maine. — All
the members of this genus are oceanic, and inhabit
the mid-depths. Localities listed by Schultz n
for this species include the Grand Banks, between
Georges and Browns Banks, and the offing of New
Jersey and Virginia in the western Atlantic; the
Gulf of Mexico; West Indies; off the South African
coast; the Indian Ocean; and the Philippines. Our
only reason for mentioning the hatchetfish is that
one specimen was taken on August 31, 1883, by the
Albatross between Georges and Browns Banks
where the depth was 144 fathoms.62

THE EELS. FAMILIES ANGUILLIDAE, CONGRIDAE, SIMENCHELYIDAE, SYNAPHO-
BRANCHIDAE, NEMICHTHYIDAE, AND OPHICHTHYIDAE

Eels have no ventral fins; either they have no
scales or these are so small as to be hardly visible;
their fins are soft, without spines ; the gill openings
are very small; the vertebrae extend in a straight
fine to the tip of the tail ; and a single fin runs over
the back, around the tail and forward on the belly
with no separation into dorsal, caudal, and ventral
portions. All the species of eels known from the
Gulf of Maine have pectoral fins, but most of the
morays of warmer seas are without pectorals.
There are several other fishes of eel-like form in the
Gulf of Maine, viz., the hag and the lampreys the
rock eel (Pholis); the snake blenny (Lumpenus);
the wrymouth (Cryptacanthodes) ; the eel pout
(Macrozoarces); and the sand eel (Ammodytes).
But the jawless, sucker-like mouth of the first two
separates them, at a glance, from the true eels,
while there either is a well-marked separation be-
tween anal and caudal fins in all the rest; or they
have ventral fins (large or small), or the dorsal
fin is spiny, not soft.

Only five true eels are known from the Gulf of
Maine: the common eel (p. 151), the slime eel (p.

157), the conger (p. 154), the snipe eel (p. 159), and
the snake eel (p. 159), which fall into five different
families according to American usage. A sixth spe-
cies, the long-nosed eel (a deep-water form p. 158)
is to be expected in the deepest parts of the Gulf
though it has not actually been recorded there as
yet. The group likewise includes the morays of
warm seas and sundry deep-sea forms, some of
them exceedingly bizarre in appearance.

Common, conger, slime, and long nosed eels look
much alike in general form, but are separated from
one another by the size of the mouth and by the
relative lengths of the fins. In the snipe eels the
two jaws are prolonged into a very long slender
beak, recalling that of a silver gar, the tail is whip-
like, the neck noticeably slimmer than the head,
and the general form extremely slender, while the
snake eel is very slender with a hard pointed tail.

•i Bull. U. S. Nat. Mus.. vol. 86, 1938, pp. 146-147 and pp. 151-152.

« Albairos! station 2063, lat. 42° 23' N., long. 66° 23' W. This specimen was
recorded by Qoode and Bean (Smithsonian Contrlb. Knowl., vol. 30, 1895, p.
127) as A. olfersll. But Schultz (Proc. U. S. Nat. Mus., vol. 86, 1938, p. 151)
has found, on reexamination, that It Is an acuhaiui.

KEY TO GULF OF MAINE EELS

1. Both jaws are prolonged into a long slender bill Snipe eel, p. 159

The jaws are not bill-like 2

2. The anal fin originates well in front of the point of origin of the dorsal fin Long-nosed eel, p. 158

The anal fin originates well behind the point of origin of the dorsal fin 3

3. The dorsal fin originates far behind the tips of the pectorals Eel, p. 151

The dorsal fin originates close behind the tips of the pectorals 4

4. Mouth very small, its gape not reaching back as far as the eye; body very soft Slime eel, p. 157

Mouth large, gaping back as far as the middle of the eye; body firm 5

5. Mouth gaping back only about as far as the middle or rear edge of eye; body moderately stout; tip of tail soft, rounded

Conger, p. 154
Mouth gaping back considerably beyond eye; body very slender, tip of tail hard and pointed Snake eel, p. 159

FISHES OF THE GULF OF MAINE

151

Eel Anguilla rostrata (LeSueur) 1817

American eel; Silver eel; Fresh-water eel;
Elver (young)

Jordan and Evermann 1896-1900, p. 348, A. ehrysypa
Rafinesque 1817.

Description. — In the common American eel the
dorsal fin originates far behind the pectorals,
this character is enou^j to distinguish it from the
conger, from which it also differs in that the lower
jaw projects beyond the tipper or at least equals
it in length, and its eyes are small and round.
Furthermore, it develops scales as it grows, though
these are so small that they might be overlooked.
The eel, however, has a pointed snout, like the
conger, a large mouth gaping back as far as the
middle of the eye or past it; and its gill slits are
set vertically on the sides of the neck, their upper
corners abreast of the center of the base of the
pectoral fin. It is very closely related to the
European eel (Anguilla vulgaris), but has fewer
vertebrae (average about 107 as compared with
about 114 or 115 in the European species).

Color. — The colors of eels vary widely with the
bottom on which they live. As a rule they are
dark muddy brown or olive-brown above, more
or less tinged with yellow on the sides; the lower

surface paler brown and yellower, with dirty yel-
lowish-white belly. It is common knowledge
that eels are dark if living on dark mud but much
paler on pale sand. And Parker M has found that
they can change from pale to dark in about 1%
hours and from dark to pale in a little more than
3 hours, if moved from a white background to a
black or vice versa, under a strong light.

Size. — Eels are said to grow to 4 feet in length
and to 16K pounds in weight. Full-grown females
average only about 2 to 3% feet, however, and
males are smaller. Any eel more than 18 inches
long would probably be a female, and one more
than 24 inches in length would certainly be one.
The smallest mature males are about 11 to 12
inches long, females about 18 inches.

Habits. — The life history of the eel remained a
mystery until very recently. It has been com-
mon knowledge for centuries that young elvers
run up into fresh water in spring, and adults
journey downstream in autumn. A host of myths
grew up to explain the utter absence of ripe eels
of either sex, either in fresh water or along the
seacoast. But it was only a few years ago that
the breeding places of the European and American
eels were discovered and the history of their larvae

« Jour, of Exper. Zool., vol. 88, 1945, No. 3, pp. 211-234.

Figure 69. — Eel (Anguilla rostrata). A, adult, Connecticut River, Massachusetts; from Goode, drawing by H. L.
Todd; B, "Leptocephalus" stage, 49 mm.; C, "Leptocephalus" stage, 55 mm.; D, "Leptocephalus" stage, 58 mm.;
E, transformation stage, 61 mm. B-E, after Schmidt.

152

FISHERY BULLETIN OF THE FISH AND "WILDLIFE SERVICE

traced, chiefly by the persevering researches of
the Danish scientist, Johannes Schmidt.64 Now
we know that the life history of the eel is just the
antithesis of that of the salmon, shad, and alewife,
for eels breed far out at sea, but make their growth
either in estuarine situations or in fresh water.

The young elvers, averaging from 2 to 3% inches
in length, appear along our shores in spring. As
yet we have few data on the exact date of their
arrival on the Gulf 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 this 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 (PhUos. 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. Qeog. Mag., vol. 24, No. 10,
October 1913, p. 1140).

u 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 6S has proved, by experiments
with European eels marked so as to be recognizable
if recaught, that they can carry 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 by 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 67
concluded from a series of measurements taken at
different seasons in lower Chesapeake Bay that those

* Mem. R. Comlt. Talassogr. Ital., vol. 158, 1929.
»' Bull. U. S. Bur. Fish., vol. 43, 1928, p. 114.

FISHES OF THE GULF OF MAINE

153

2}j 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 once
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."

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 beiag that no spent eels ha ye ever
been seen and that large eels have never been

IS Large eels, on their seaward journey, have occasionally been caught by
otter trawlers in the western part of the British Channel, but we know of no
such occurrence on this side of the Atlantic.

« See Schmidt (Ann. Rep. Smithsonian Inst., (1924) 1925, pp. 279-314) for
a readable account of 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, flgs. 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 offshore. 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 1 1 ; 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,81 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 of 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, 1926, Eitr.
Rapp. et Proc. -Verb Cons. Internet. Eipl. 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.

u Reported by Dr. O. W. Jeffers 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 30,000^40,000 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 M

Sea eel

Jordan and Evermann, Leptocephalus conger (Linnaeus
1758), 189&-1900, p. 354.

•* Information from A. H. Lelm.

•* 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 Anouilla.

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 rather
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 anil
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.69

Habits. — The depth range of the conger is from
close to the coastline (they are 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 now 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, 1898
pp. 1016-1017, 1037.

■ 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 OP 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.

'• For photographs of the leptocephalus stage of the European conger, see
Schmidt, Rapp. et Proc. Verb. Cons. Perm. Intemat. Eiplor. Mer, vol. 5,
No. 4, 1906, pi. 9, flgs. 8, 9; and Meddelelser Komm. Havundersiigelser, Ser.
Flskerl, vol. 3, No. 6, pi. 1, flgs. 1-3.

" Conptes Rendus Acad. Scl. Paris, vol. 103, 1886, 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 mo rays, 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 1950.7* 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." 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.

» Schmidtleln (Mittlell, 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.

» 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' W.

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 type 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 known 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 of 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 (Uptosomus)
by Tanaka in 190S, has been classed more recently by him (Fishes of Japan,
vol. 42, 1928, p. 810, pi. 173, flg, 476) as identical with the Atlantic parasiticu*.

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 " mentions
one 26% inches (675 mm.) long from the Azores.

" Result, des Camp. Sci. Prince de Monaco, Pt. 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 in
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 axe described, with
photographs by Schmidt (Rapp. et Proc. Verb. Cons. Perm. Intemat.
Eiplor. Mer, vol. 6, No. 4, 1906, p. 191, pi. 9, flgs. 4-6; and Meddel. Komm-
Havunderstfgelser, Ser, Flskeri, vol. 3, No. 6, 1909, p. 7).

i» This fact is commented on by Schmidt (Rapp. Cons. Perm. Intemat.
Eiplor. Mer, vol. 5, No. 4, 1906, p. 191). For further details as to its distri-
bution see Koefoed, Rept. Michad Surj North Atlantic Eiped., (1910), vol.
4, Pt. 1, 1927, pp. 11, 14.

Figure 72. — Long-nosed eel (Synaphobranchus pinnatus), La Have Bank. From Goode and Eean. Drawing by

H. L. Todd.

FISHES OF THE GTTLF 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 only 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

*> Many such Instances are listed In the Rept. U. S. Comm. Fish. (1879)
1882, p. 787.
>> Goode and Bean, Smithsonian Contrlb. Knowl., vol. 30, 1895, pp. 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 oj 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

" Goode 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

2^22222222222^2;

>a^""""v»"^vi^w>>Y.^^^^^^^^SSi^^

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 Vaillant83 shows both as about
as high throughout their length as the fish is deep,
Goode and Bean84 picture the dorsal as much
higher 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
nothing 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. Scl. Travailleur et Talisman, 1888, pi. 7, figs. 2 and 2a.
M Smithsonian Contrlb. Knowl., vol. 31, 1895, pi. 46, fig. 170.
'« Tiefsee-Flsche, Wlss. Ergeb. Deutsch. Tiefsee-Exped. (189S-1899), vol
16, Pt. 1, 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-

"Copela ,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 Gulf 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 Vladykov and
McKenzie 89 have reported other captures of lan-
cetfishes from La Have Bank, from southeast of
Emerald Bank and Banquereau. Another speci-
men 5% 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. 66) reports several A. ferox from
the halibut banks off the northwestern coast of British Columbia.
M No definite information is available as to this specimen.
'• Proc. Nova Scotia Inst. Sci., vol. 19, 1935, p. 63.

Figube 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 Funduius 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 Fundvlus being sharp-pointed, whereas
they are wedge-shaped in Cyprinodon and in-
cisorlike. The two local species of Funduius
are separable by their markings, majalis of all
ages being barred or streaked with black while
the adult heteroclitus is not.

Common mummichog Funduius 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 eyes.
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 Funduius 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-

" A detailed account of the sexual differences Is given by Newman (Biol.
Bull., vol. 12, No. 5, 1907, pp. 314-348).

Figure 76. — Common mummichog (Funduius heterocliius), Maryland.

A. H. Baldwin.

From Jordan and Evermann. Drawing by

FISHES OF THE GULF OF MAINE

163

form olive to bottle green, darker above, lighter
below, without definite markings though their
sides often show faint and indefinite crossbars of a
deeper tone of the same hue. Their fins are much
paler than those of males. Very young fry of
both sexes show dark transverse bars on the sides,
but these bars are lost with growth.

Killifishes vary in shade from very pale to dark,
according to the color of their surroundings. And
recent experiments 9l have proved that their
ability to change from light to dark depends on the
sense of sight.

Size. — The maximum length is 5 to 6 inches,
but adult mummichogs are seldom more than 3K
to 4 inches long in the Gulf of Maine. Several
varieties of this species have been described, but
they are so closely allied that it is not necessary to
discuss them here.

Habits. — The home of the mummichog is along
sheltered shores where the tide flows over beds of
eelgrass or salt hay (Spartina), among which shoals
of "mummies" may often be seen moving in with
the flood tide. They abound in the tidal creeks
that cut the salt marshes, on the shores of our
harbors, and in the brackish water at the mouths
of our streams and estuaries, particularly in little
muddy pools, creeks, and ditches. So closely,
indeed, do they hug the shore that a line drawn
100 yards out from land would probably inclose
practically all the mummichogs in the Gulf of
Maine. Where the shore is bold and rocky, as it is
about the Bay of Fundy, the mummichog is
practically restricted to brackish water, and it
often goes up into fresh water. At the other
extreme, it is not likely that they ever descend to
a depth of more than a couple of fathoms in their
journeys in and out of the creeks or along the
shore.

Mummichogs are so resistant to a lack of oxy-
gen, to the presence of carbon dioxide, and to un-
favorable surroundings generally, that they can
survive in very foul water. At ebb tide "mum-
mies" are often trapped in little pools where they
remain until the next tide if the water holds, often
huddled together in swarms. Should the pool go
dry, they work their way for the time being into
the mud, where we have often found them.02 And
it is probable that they can flop overland for a few

yards to some other drain as the striped mummi-
chog does.

They winter in a more or less sluggish state on
the bottoms of the deeper holes or creeks, where
they have been found buried 6 or 8 inches deep in
the mud,93 and there is no evidence that they move
out to sea during the cold season; in short, this is
one of the most stationary of fishes.

Food. — "Mummies" are omnivorous, feeding on
all sorts of edible things, vegetable as well as
animal. They have been found full of diatoms,
eelgrass, and other vegetable matter; foraminif-
era; shrimps and other small Crustacea, also
small mollusks; and they sometimes have small
fish in their stomachs.94

They soon congregate about any dead fish or
other bit of carrion, to prey either upon it or upon
the amphipod scavengers that gather on such
dainties and they eat their own or each other's
eggs at spawning time.

Spawning probably takes place at the same
season in the Gulf of Maine as on the southern
coast of New England; i. e., in June, July, and
early August. As sexual activity approaches, the
males (now brilliantly tinted) court and pursue
the females, rivalry among them being very keen;
those that are the most highly colored or most
excited usually driving off the others. Sometimes
they fight fiercely. They spawn in a few inches of
water, seeking shady spots. At the moment of
spawning the male clasps the female with his anal
and dorsal fins just back of her anal and dorsal,
usually forcing her against some stone or against
the bottom, the bodies of both are bent into an S
and their tails vibrate rapidly while the eggs and
the milt are extruded.95 Occasionally, pairs clasp
and spawn free in the water without coming in
contact with any object, and sometimes a female
is seen to pursue and court a male.

The eggs, which are about 2 mm. in diameter,
colorless or pale yellowish and surrounded by a
firm capsule, sink and become so sticky on contact
with the water that they mass together in clumps,
or stick fast to sand grains or to anything else
they chance to rest upon. Incubation occupies
from 9 to 18 days, the exact duration probably

•' Parker and Lanchner, Amer. Jour, of Physiol., vol. 61, 1922, p. 648.
•• This habit Is described by Mast (Jour, of Animal Behavior, vol. 5, No. 5,
1915, p. 351).

•I Chichester, Amer. Naturalist, vol. 64, 1920, p. 651.

•< Lists of stomach contents are given by Field (Rept. U. S. Comm. Fish.,
H906) 1907, p. 29).

« Newman (Biol. Bull., vol. 12, No. 5, April 1907, p. 315) gives an Interest-
ing account of the courtship and spawning, from which the preceding la
condensed.

164

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

depending on temperature. The larva is about 7 to
7.7 mm. long at hatching, its yolk absorbed al-
ready, and its pectoral and caudal fins fully formed.
By the time the little fish has grown to 11 mm. the
dorsal and anal fin rays are present in full number,
and the first trace of the ventrals is to be seen. At
16 mm. the ventrals are apparent, and fry of 20
mm. resemble their parents.

General range. — Coast of North America, from
the Gulf of St. Lawrence to Texas. Port au Port
Bay, on the west coast of Newfoundland,95 is the
most northerly record that we have found for it.

Occurrence %n the Gulf of Maine. — The ' ' mummie' '
is one of the few fish which can fairly be charac-
terized as "universal" in suitable locations around
the entire coastline of the Gulf. We dare say
that there is not a single bit of salt marsh, muddy
creek, harbor, sheltered shore line, or brackish
estuary, where they are not to be found, from the
elbow of Cape Cod around to Cape Sable.

•• Johansen (Canadian Naturalist, vol. 40, February 1926, p. 34).

Importance. — The mummichog is of some com-
mercial value as bait, but only locally. It is
also a favorite for biological experiment.

Striped mummichog Fundulus majalis (Wal-

baum) 1792

Mummichog; Mummy; Killifish

Jordan and Evermann, 1896-1900, p. 637.

Description. — This fish resembles the common
mummichog closely in general form, in the shape
of its dorsal and aDal fins, in its sexual dimorphism,
and in the development of "contact organs" on the
scales of breeding males. But it is more slender,
its snout more pointed, its body more definitely'
fusiform, tapering toward both head and tail,
and its caudal peduncle not so stout. But the
color pattern is the most striking difference
between the two, both sexes of Fundulus majalis
being definitely barred with black at maturity
as well as when young. In the male the barring
is transverse throughout life, the stripes increasing

Figure 77. — Striped mummichog (Fundulus majalis). Upper, male, Woods Hole; lower, female, Maryland.

Jordan and Evermann.

From

FISHES OF THE GULF OF MAINE

165

in number from 7 to 12 in the young to 14 to 20 in
adult fish. When the females reach a length of
about two inches, however, the original 7 to 12
transverse bars are transformed with growth into
two or three longitudinal stripes on each side, the
upper stripe running uninterrupted from gill
opening to tail, the lower stripes in two segments,
the one from close behind the pectoral to above
the ventral, the other thence backward to close
behind the rear edge of the anal fin. One or two
transverse bars persist however on the caudal
peduncle, even on the oldest females.

Color. — This is a decidedly paler fish than the
other "mummy." Apart from the black bars
the male is dark olive green above with silvery
sides, a greenish-yellow belly, and a black spot
on the rear part of the dorsal fin; his pectorals and
caudal are pale yellowish. The male becomes
more brilliant at breeding time, the back turning
almost black, the lower sides and belly changing to
orange or golden, and the fins to bright yellow.
The female is olive green above and white below,
striped as described above.

Size. — This is a larger fish than the common
mummichog, occasionally growing to a length of
7 inches and often to 6 inches.

Habits. — The striped mummie parallels the
common mummie in being restricted to the
immediate neighborhood of the land, and in its
way of life, except that it keeps more strictly to
salt water, and is found more often along open
beaches. Its most interesting habit is its ability
to flop back into the water if it becomes stranded
with the receding tide, jumping unerringly toward
the water in almost every instance, and progressing
from several inches to several feet at each jump.97
And so noted are they for this peculiar ability
that a special article has been devoted to it.88
Their diet consists of small animals including
mollusks, crustaceans, fish, insects, and insect
larvae. Westward and southward from Cape
Cod they spawn from late spring to late summer.

General range. — Coast of the United States,
from the vicinity of Boston, Mass., to Florida.

Occurrence in the Gulf of Maine. — The striped
mummie is very abundant along the southern
shores of New England, westward from Cape
Cod. But the only published records for it in

the gulf are for the vicinity of Boston and Salem,
many years ago, and we had not seen it north of
Cape Cod before 1937. In that autumn, how-
ever, B. Preston Clark brought in four specimens
that he had taken at Cohasset, on the southern
shore of Massachusetts Bay; it was reported to
us as in numbers there in 1939," and we have
seen small schools of them in recent summers
in the salt marshes at the entrance to Cohasset
Harbor, as well as nearby. If this little fish
actually has extended its regular range north-
ward and if its dispersal-route has been via the
Cape Cod Canal, as has been suggested,1 it is to be
expected anywhere in the marshes around Cape
Cod Bay and along the southern shore of Massa-
chusetts Bay, and we suspect that a resident
population is to be found in the Nauset Marshes
and in Pleasant Bay, on the outer shore of Cape
Cod.

Sheepshead minnow Cyprinodon variegatus
Lacep&de 1803

Jordan and Evermann, 1896-1900, p. 671.

Description. — The sheepshead minnow is so
deep bodied (its body is nearly half as deep as
long, not counting the tail fin) that it is not apt
to be mistaken for either of the mummichogs.
And it is separated further from the mummi-
chogs by its teeth, which are large, wedge-shaped
and with tricuspid cutting edges, instead of small
and pointed. It is a flat-sided little fish, with
high arched back, small flat-topped head, small
terminal mouth hardly gaping back to the forward
edge of the eye, and it has the thick caudal
peduncle characteristic of its family. Its tail is
square (rounded in the mummichogs), and the
fact that almost the whole of its dorsal fin is in
front of the anal instead of over it affords an-
other point of difference. The pectorals are
large, reaching back past the base of the ventrals,
which seem very small by contrast. Both its
body and its head are covered with large rounded
scales, largest on top of the head and on the
cheeks, with one much larger than the others just
above the pectoral fin. Young fish are propor-
tionally more slender than old ones. The dorsal,
ventral, and anal fins are higher in the males of
this species than in the females, much as they

•' Hildebrand and Schroeder, Bull. U. 8. Bur. Fish. vol. 48, Pt. 1 1928
p. 141.
•' Mast, Jour .o (Anlma Behavior, vol. 5, No. 5, 1916, pp. 341-340.

« By John W. Lowes.

i Schroeder, Copela, 1937, No. 4, p. 238.

166

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 78. — Sheepshead minnow (Cyprinodon variegatus), Maryland. From Jordan and Evermann. Drawing by

A. H. Baldwin.

are in the mummichogs. The male is deeper
bodied and the average size is larger than that
of the female.

Color. — Out of breeding season both males and
females are olive above (males rather darker and
greener than females) with pale yellow or yellow-
ish-white belly, dusky dorsal fin, and pale orange
pectoral, ventral, and anal fins. The young of
both sexes are irregularly barred with black
transverse stripes, which persist through life in
the female but become obscured in adult males.
Females, furthermore, have a black spot on the
rear corner of the dorsal fin, which is lacking in
males, while the caudal fin of the male is marked
by two black cross stripes, one at the base and
the other at the margin. In breeding season
the male assumes a brilliant coat, his upper
parts turning to steel blue in front of the dorsal
fin with a greenish luster behind it, while his
belly brightens to a deep salmon, his ventrals
and anal change to dusky margined with orange,
and the front edge of his dorsal turns orange.

Size.— The largest specimens are about 3
inches long.

Habits. — The sheepshead minnow (like the com-
mon mummichog) is confined to the shallow waters
of inlets, harbors, and the heads of bays, and salt
marshes, often in brackish water. Its diet is

partly vegetable, partly animal. It is very pug-
nacious, often killing fishes larger than itself,
making repeated attacks with its sharp teeth and
finally devouring its victim. Its breeding habits
recall those of the mummichog (p. 163), the males
fighting fiercely among themselves and clasping
the females just forward of the tail with dorsal
and anal fin, while the eggs and milt are extruded.
Spawning takes place in shallow water from April
to September, the eggs maturing a few at a time,
so that any given female spawns at intervals
throughout the season. The eggs sink and stick
together in clumps by numerous threads. They
are 1.2 to 1.4 mm. in diameter, with one large oil
globule and many minute ones. Incubation oc-
cupies 5 or 6 days, and even at hatching the larvae
(4 mm. long) show alternate light and dark cross-
bands. At a length of 9 mm. all the fins are
formed, and at 12 mm. the fry show most of the
characters of its parents.2

General range. — Atlantic coast of the United
States, Cape Cod to Mexico, in brackish as well
as in salt water.

Occurrence in the Gulf of Maine. — This fish, like
many others, finds its northern limit at Cape Cod

1 An account of courtship and spawning is given by Newman (Biol. Bull.,
vol. 12, No. 8, 1907, p. 336) and of development by Kuntz (Bull., U. S. Bur.
of Fish., vol. 34, (1914) 1916, p. 409).

FISHES OF THE GULF OF MAINE

167

and would not deserve mention here at all were it
not recorded from the Cape by Storer. West and
south of Cape Cod, however, as at Woods Hole,

it is common enough in suitable situations. We
have seined many of them with Fundvlus at the
head of Buzzards Bay.

THE BILLFISHES OR SILVER GARS. FAMILY BELONIDAE

The most noticeable feature of the billfishes is
that both of their jaws are prolonged to form a long
slim beak well armed with teeth. Their bodies
are very slender, and their anal, dorsal, and ventral
fins set far back. They have no finlets between
the dorsal and anal fins and the caudal, the absence
of these being the readiest field mark to separate
the billfishes from the needlefishes (Scomber esox, p.
170). They are swift-swimming, predaceous fishes,
represented by many species, most of them
American. Only two have ever been recorded in
the Gulf of Maine.3

Key to Gulf of Maine Billfishes

Body as thick as it is deep; dorsal, anal, and caudal fins
only moderately concave Silver Gar, p. 167

Body less than }i as thick as it is deep; dorsal, anal, and
caudal fins deeply concave Garfish, p. 168

Silver gar Tylosurus marinus (Walbaum) 1792 4

Billfish; Salt-water gar; Sea pike; and
various other local names

Jordan and Evermann, 1896-1900, p. 714.

Description. — Its long bill and slender body give
the gar so peculiar an aspect that it is not likely to
be confused with any other Gulf of Maine fish

'The closely allied houndflsh (Tylosurus acus Lacepe.de 1803) has been
taken at Nantucket, but has not been found within the Oulf of Maine.
Since It may appear there as a stray from the south , we may point out that it
Is easily distinguished from the sliver gar by Its de eply forked tall and by the
fact that Its dorsal and anal fins are much longer, the former with 23 rays, the
latter with 21. The following characters In combination will serve to Identify
It among the several tropical gars; mouth nearly closable and upper jaw not
arched; dorsal and anal fins long; beak at least twice as long as rest of head;
greatest depth of body not more than two-thirds as great as length of pec-
toral fin; no lateral stripe.

• Jo rdan, Evermann, and Clark (Kept. U. 8. Comm. Fish., (1928) Pt. 2,
1930, p. 196) place this species In the genus Strongylum Van Hasselt 1824.

other than the half beak (p. 169), the needle fish
(p. 170), or its own close relative Ablennes (p. 168).
And it is easily distinguishable from the first of
these by the fact that both of its jaws are prolonged
instead of only the lower; from the second by lack-
ing detached finlets on its back between its dorsal
and caudal fins. The most conspicuous differ-
ences between the silver gar and Ablennes (p. 168)
is that the body of the former is thicker than it is
deep, and that its fins are only moderately con-
cave, whereas the latter is so strongly flattened
sidewise that it is less than one-half as thick as it
is deep with deeply concave fins.

The head of the adult silver gar occupies nearly
one-third of the total length of the fish; the upper
jaw, from the eye forward, is twice as long as the
rest of the head; both jaws are armed with sharp
teeth; and the eyes are large. The long, slender
body is only about one-twentieth as deep as long,
rounded (not laterally flattened) in cross section,
and thicker than deep. Both the body and the
sides of the head are scaly. The dorsal fin, with
13 to 17 rays, and the anal fin, with 17 to 21 rays,
are alike in outline, the anterior rays of both being
much longer than those toward the rear, and the
rear two-thirds of each can be depressed along the
back and nearly concealed in a groove, while the
forward one-third continues erect. Both fins, too,
are situated far back, with the dorsal arising a
little behind the forward end of the anal.

The ventral fins stand about halfway between
a point below the eye and the base of the caudal.
The margin of the caudal fin is only moderately
concave, this fact being the readiest field mark to
separate this particular gar from the only other
species of its genus (Tylosurus acus) taken yet near
the Gulf of Maine (see footnote, p. 167), for the tail
of the latter is deeply forked. There is a distinct

Figure 79. — Silver gar (Tylosurus marinus).

168

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

longitudinal ridge, or low keel, along either side of
the caudal peduncle.6

Color. — Greenish above, silvery on the sides,
white below; a bluish silvery stripe aloDg each side
becoming broader and less distinct toward the tail;
snout dark green; there is a blackish blotch deeper
than long on the upper part of the cheek. The
fins are without markings for the most part; the
dorsal may be somewhat dusky, and the caudal
bluish at its base.

Size. — The silver gar grows to a length of about
4 feet.

General range. — Maine to Texas ; abundant along
the South Atlantic and Gulf coasts of the United
States, often running up fresh rivers above tide
water.

Occurrence in the Gulf oj Maine. — -The silver gar
is common enough along the southern shores of
New England, e. g., in Rhode Island waters and
at Woods Hole where quite a few are found from
June to October. Like many other southern fishes,
however, it seldom journeys eastward past Cape
Cod, the only definite records of it within the
Gulf of Maine being of several collected by Dr.
William C. Kendall at Monomoy Island, forming
the southern elbow of Cape Cod; at Wolfs Neck,
Freeport, and Casco Bay, Maine, and of one
found by Crane 6 in the stomach of a tuna that
she examined at Portland, Maine, in July 1936.
We have not seen it in the Gulf, nor have we heard
even a rumor of its presence there from fishermen,
good evidence that it is as rare a straggler as the
few records indicate, for large silver gars are not

• There are many other species of Ears In tropical seas, any one of which
might stray northward with the Gulf Stream and so to the Gull of Maine.
The silver gar Is Identifiable among them by the following combination of
characters (no one character alone marks It out among Its relatives): mouth
capable of being nearly closed; caudal peduncle with keels; dorsal and anal
fins short, the former with 13-17 rays, the latter with 17-21 rays; caudal fin
only moderately concave; eyes at least one-third as broad as the head Is long
behind the eyes; body not excessively slender but at least one- fifth to one-sixth
as deep as head (including Jaws) Is long; body not very strongly compressed
sldewlse; Jordan and Evermann (Bull. 47, U. 8. Nat. Mus., Pt. 1, 1896. p.
709) gives a useful key to the species of the family.

• Zoologica, New York Zool. Soc, vol. 21, No. 16, 1936, p. 211.

fish to be overlooked. With so little claim to
mention here, we need merely note that it is
voracious, feeding on all sorts of smaller fishes,
and that it runs inshore, possibly even into river
mouths, to spawn. The eggs, described by Ryder,7
are about 3.6 mm. (one-seventh of an inch) in
diameter, and stick together and to any object
they may touch, by long threads scattered over
their surface.

Garfish Ablennes hiana (Cuvier and Valenciennes)
1846

Jordan and Evermann, 1896-1900, p. 718, Athlennes
hians.'

Description. — -This gar resembles the silver gar
(Tylosurus) closely in general appearance, as well
as in the nature and arrangement of its fins;
especially in the fact that the rear parts of its
dorsal and caudal fins can be depressed and almost
completely concealed in a groove, with the for-
ward parts still remaining erect. But its body is so
strongly flattened sidewise as to be less than one-
half as thick as it is deep, instead of about as
thick as deep, or thicker, as it is in the silver gar.
The dorsal fin, also, arises farther back relative to
the anal fin, than is the case in the silver gar.
The tail fin is broadly forked, the dorsal and anal
fins deeply concave.

Color. — -Back greenish with bluish green reflec-
tions; lower part of sides bright silvery, also
the abdomen; snout greenish; dorsal fin mostly
greenish, but with the rays black-tipped; tail fin
greenish. Some individuals have the sides plain
silvery, but others are marked with dark blotches
or indistinct sooty or blue crossbars.9

Size. — Up to 3 feet long, or more.

' Bull. U. S. Fish Comm., vol. 1, 1882, p. 283.

« The original spelling was Athlennes (Jordan and Fordice, Proa U. S. Nat.
Mus., vol. 9, 1886, p. 342). But Jordan and Evermann (Bull. 47, U. S. Nat.
Mus., Pt. 1, 1896, p. 717, footnote) state that "Ablennes" was intended.

• Smith (Sea Fishes of Southern Africa, 1949, pi. 7, fig. 26) gives a colored
Illustration o lone with blue crossbars.

^.iii num iuw.iiihhhu wi.VMwm.1 1 iwp«

Figure 80. — Garfish (Ablennes hians), specimens from North Truro, Mass., and from Acapulco, Mexico. Drawing by

H. B. Bigelow.

FISHES OF THE GULF OF MAINE

169

General range and occurrence in the Gulf of
Maine. — Widespread in tropical seas; 10 Brazil to
Chesapeake Bay in the western Atlantic, and
northward as a stray to Cape Cod. A specimen

of this tropical fish, about 23% inches (594 mm.)
long to the fork of the tail, was taken in a fish
trap on the shore of Cape Cod Bay at North
Truro, Mass., on August 15, 1949.11

THE HALFBEAKS. FAMILY HEMIRAMPHIDAE

The halfbeaks are close allies of the billfishes
(Belonidae, p. 167), but it is only the lower jaw
that is greatly prolonged while the upper jaw is
short in the only species of present concern. They
are largely herbivorous, feeding mainly on green
algae. There are many species in warm seas,
only one of which is known to reach the Gulf of
Maine.

Halfbeak Hyporhamphus unifasciatus

(Kanzani) 1842

Skipjack

Jordan and Evermann, 1896-1900, p. 721 (Hyporham-
phus robeUi (Cuvier and Valenciennes) 1846.

Description. — The most striking feature of this
halfbeak and one which is enough of itself to mark
it off from every other fish known from the Gulf
of Maine, is the fact that while the lower jaw is
very long, the upper jaw is short.12

This is a slender fish, its body only one-sixth to
one-tenth as deep as long (younger fish are still
more slender), tapering slightly toward head and
tail. Its dorsal (14 to 16 rays) and anal (15 to
17 rays) fins are situated far back and opposite
each other, as in the silver gar, and are about
equal in length and alike in outline. There are
no detached finlets between them and the caudal

io We have seen specimens from Acapulco, west coast of Mexico; Panama;
Mauritius; and Zanzibar.

11 This specimen was presented to the Museum of Comparative Zoology by
John Worthlngton of the Pond Village Cold Storage Co., North Truro,
Mass.

" Should a halfbeak be taken in the Gulf of Maine it would be well to con-
sult Meek and Hildebrand (Field Museum of Natural History, Zool. Series,
vol. 15, Pt. 1, p. 232, 1923) for there are several other species that might reach
us as strays, either via the Gulf Stream route or from offshore. One, indeed
(Euleplorhamphus veloz), has been taken at Nantucket. Its lower jaw is even
longer and more slender than that of the halfbeak. its body is more flattened
sldewlse, and Its pectoral flns are longer.

fin. The ventrals stand about midway between
a point below the eye and the base of the caudal.
The teeth are small and the scales are largest on
the upper surface of the head. The beak is much
shorter in young fish than it is in adults.

Color. — Translucent bottle green above with
silvery tinge, each side with a narrow but well-
defined silvery band running from the pectoral fin
to the caudal fin, the sides darkest above and
paler below this band. The tip of the lower jaw is
crimson in life, with a short filament, and three
narrow dark streaks run along the middle of the
back. The forward parts of the dorsal and anal
fins and the tips of the caudal fins are dusky.
The lining of the belly is black.

Size. — Adults are seldom more than 1 foot long.

General range. — Tropical and subtropical on
both coasts of America and in the Gulf of Mexico ;
abundant off the South Atlantic United States,
not uncommon northward to Cape Cod, and stray-
ing to the coast of Maine.

Occurrence in the Gulf of Maine. — In our cool
boreal waters the halfbeak is only a rare stray
from the south, previously recorded only twice in
the Gulf of Maine, i. e., from Machias and from
Casco Bay, Maine, many years ago in each case.
We can now add one taken in Quincy Bay, Boston
Harbor, July 10, 1951, by Gordon Faust; another
off Revere (also in Boston Harbor) on the 19th of
the month, by John M. Hodson; a third, taken
in a trap at Sandwich, September 24 of the same
year,13 and several dozen taken in a pound net at
Small Point, Maine, July 14-15, reportedjby Leslie
Scattergood.

11 These specimens are in the Harvard Museum of Comparative Zoology.

Figure 81. — Halfbeak (Hyporhamphus unifasciatus), Chesapeake Bay.

H. L. Todd.

From Jordan and Evermann. Drawing by

170

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

THE NEEDLEFISHES. FAMILY SCOMBERESOCIDAE

Both jaws (of the adult) are elongated to form a
slender beak in the needlefishes, as in the billfishes
(p. 167), and the anal, dorsal, and ventral fins are
set far back. But the presence of several finlets
between the dorsal and anal fins and the caudal in
the needlefishes (which the billfishes lack) is a
ready field mark for their identification. Further-
more, their teeth are small and weak, and their
bodies only moderately slender. Four or five
species are known in warm seas, one of which is not
uncommon in the Gulf of Maine.

Needlefish Scomberesox saurus (Walbaum) 1792

Billfish; Skipper; Saury

Jordan and Evermann, 1896-1900, p. 725.

Description. — The needlefish resembles the sil-
ver gars in its slender form and in the fact tbat
both its upper and lower jaws are prolonged, but

greenish; the lower parts are silvery with golden
gloss. Young fry, which live in the surface
waters of the open Atlantic, have dark blue backs
and silvery sides.

Size. — Up to 18 inches long. Those caught
along Cape Cod run a foot and more in length.

Habits. — The skipper is an oceanic fish. So
far as known it always lives close to the surface;
so much so indeed that in English waters, where
it is plentiful in summer, few are caught in nets
set as deep as a fathom or two. Its hordes are
preyed upon by porpoises and by all the larger
predaceous fishes; cod and pollock, for instance,
feed greedily upon them, as do bluefish. When
they strand on the beaches, as often happens, it is
probably while they are fleeing from their enemies.
At sea they attempt to escape by leaping, whole
companies of them breaking the surface together
as has often been described, and as we have seen
them doing in Massachusetts Bay.

Figure 82. — Needlefish (Scomberesox saurus) . Adult, Woods Hole. From Goode. Drawing by H. L. Todd.

it differs from the gars in having a series of five or
six little separate finlets, both on its back in the
space between the dorsal fin and the caudal, and
on its lower side between the anal fin and the
caudal. Its body is about nine times as long
(not counting caudal fin) as deep, flattened side-
wise, tapering toward the head and tail, with
slender caudal peduncle, and all its fins are small.
Its dorsal fin originates slightly behind the origin
of its anal; these two fins are alike in outline and
stand far back. Its ventrals are situated about
midway the length of the body. Its caudal is
deeply forked and symmetrical, much like the
tail of a mackerel. Its trunk is covered with
small scales as is a patch on each gill cover. Its
lower jaw projects a little beyond the upper, and
its teeth are pointed but very small.

Color. — Olive green above with a silver band
on each side at the level of the eye and about as
broad as the latter. There is a dark green spot
above the base of each pectoral; the dorsal fin is

©

Figure 83.-

-Needlefish, young, about 2J4 inches long.
After Murray and Hjort.

It is not likely that they ever spawn in the cool
waters of the Gulf of Maine, for we have never
taken their fry in our tow nets, although they are
among the most numerous of young fish in the
open Atlantic between the latitudes of 11° or 12°
N. and 40° N. Although their eggs are covered
with filaments like those of the silver gars,14 they
are not adhesive as the latter are, but drift near
the surface. The most interesting phase in the
development of the skipper is that its jaws do not
commence to elongate until the fry have grown

» Skipper eggs were so described by Haeckel (Archiv fur Anatomie, Phy-
siologic, and Wissenschaftliche Medecin, 1855, p. 23, pi. 5, fig. 15,) 75 years
ago. They were not seen again until 1910 when similar eggs, 2.2 mm. In
diameter, covered with filaments, were towed in the Atlantic by the Michael
Sart (Murray and Hjort. Depths of the Ocean, 1912, p. 742, fig. 531).

FISHES OF THE GULF OF MAINE

171

to about 1% inches (40 mm.), and that the lower
jaw out-strips the upper at first, so that fry of
4 to 6 inches (100 to 150 mm.) look more like
little halfbeaks ("Hemirampbus" itage) than like
their own parents.

European students tell us that the skipper feeds
on the smaller pelagic Crustacea and probably
also on small fish, for it is sometimes caught on
hook and line. One examined by Linton at
Woods Hole contained chiefly annelid worms,
fragments of fish, copepods and crustacean larvae,
with some vegetable debris.

General range. — Temperate parts of the Atlantic,
Pacific, and Indian Oceans, known in the open
sea as far north as northern Norway off the Euro-
pean coast, and to southern Newfoundland and
southern Nova Scotia 16 off the eastern American
coast.

Occurrence in the Gulf of Maine. — -While the
skipper is a straggler to our Gulf from warmer
waters offshore or farther south, it has been
taken along the northern coasts of New England
more often than have any of its relatives ; specific-
ally along Cape Cod; at Provincetown ; at several
locations in Massachusetts Bay where we have
seen schools of them; at Annisquam a few miles
north of Cape Ann; at Old Orchard (Maine); in
Casco Bay; at Monhegan Island; in the central
part of the Gulf; 16 among the islands at the
northern entrance to the Bay of Fundy; and on
the northern part of Georges Bank, where one was
gaffed from the Albatross //on September 20, 1928.
But we find no record of it along the Nova Scotia
shore of the Gulf of Maine. The inner curve of

Cape Cod from Provincetown to Wellfleet seems
to be a regular center of abundance for it, as
Storer long ago remarked, for schools of billfish
are picked up in the traps along that stretch of
beach almost every year, the catch occasionally
amounting to hundreds of barrels, and hosts of
them have been known to strand there. Its num-
bers fluctuate greatly from year to year, however,
and it often fails to appear.17

They are likely to be taken any time from mid-
June to October or November, the largest catches
usually being made late in summer.18 We have
seen several schools skipping, as is their common
habit, off the Scituate shore on the southern side
of Massachusetts Bay. But skippers are so much
less common farther within Massachusetts Bay
that some fishermen had never heard of them
there. They appear only as strays north of Cape
Ann. But it would not be astonishing if a large
school were to be encountered anywhere within
the Gulf; witness their occasional abundance off
northern Nova Scotia.19 When skippers do in-
vade the waters of our Gulf, they may be expected
in multitudes, for they usually travel in vast
schools. Day,20 for example, mentions the cap-
ture of 100,000 in a single haul in British waters.

Commercial importance. — The skipper is not of
much commercial importance, being too sporadic
in its appearances. However, when large catches
are made on Cape Cod they find a ready sale near
by. If too many are caught for the local trade to
absorb, they are sent to Boston, where they are
sold for bait.

THE FLYING FISHES. FAMILY EXOCOETIDAE

The typical flying fishes have one dorsal fin
and one anal fin, both of them soft rayed, both
of them located far rearward, and with the anal
below the dorsal. Their ventral fins are well
behind their pectorals, their tails are very deeply
forked with the extreme tips rounded, the lower
lobe the longer, and they have small mouths and
large rounded scales. Their most distinctive
feature is that their pectorals are so long and so
stiff that their owners can plane through the air

'» Cornish (Contrib. Canadian Biol., (1902-1905) 1907, p. 83) states that
large schools can often be seen at Canso, Nova Scotia, skipping over the water
as they flee from the pollock.

'• The Museum of Comparative Zoology has a specimen, taken 115 miles
southeast of Portland Lightship.

on them, several feet above the water, which they
do mostly in attempts to escape their enemies,
and as has been described, time and again. Jordan
and Evermann have given a popular account of
this so-called "flight" (really not flight at all,
for the flying fish does not flap its wings) in their

" Blake (American Naturalist, vol. 4, 1870, p. 521) remarked that while
years before he saw thousands stranded at Provincetown not one was seen
in 1870. It failed In 1921, also, and no doubt in many intervening years.

" We are Indebted for information on the local abundance of billfish on
Cape Cod to Capt. L. B. Ooodspeed, a fisherman of long experience and
close observation.

a Cornish (Contrib. Canadian Biol., 1902-1905 (1907), p. 83) states that
large schools can often be seen at Canso skipping over the water as they flee
from the pollock.

» The fishes of Great Britain and Ireland, vol. 2, 1880-1884, p. 152.

172

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Guide to the Study of Fishes, 1905, p. 157. For
a more detailed study, based similarly on first-
hand observations, we refer the reader to Hubbs,
Papers of the Michigan Academy of Sciences,
vol. 17, 1933, pp. 575-611. Voyagers in tropical
seas are perhaps more familiar with flying fishes
than with any other fishes. And they are often
seen in the warm ultramarine-blue waters of the
Gulf Stream abreast of our northeastern coast.
But none of them are to be expected in the boreal
waters of our Gulf except as the rarest of strays.
A flying fish could hardly be mistaken for any-
thing else, except possible for a flying gurnard
(p. 472). But a glance should be enough to tell
which of them one has in hand, for the flying
fishes have stiff, narrow, pointed wings, only on
dorsal fin and a very deeply forked tail, whereas
the so-called wings of the flying gurnard are broad,
rounded, and extremely flexible; they have two
dorsal fins, and a tail fin that is only weakly con-
cave in outline.

Flying Fish Cypselurus heterurus(R&finesq\ie) 1810
Jordan and Evermann, 1896-1900, p. 2836.

Distinctive features of flying fishes of the genus
Cypselurus, among its tribe, are pectoral fins so
long that they overlap the anal fin considerably
when laid back; ventrals standing far rearward
and nearly or quite half as long as the pectorals;
anal fin with its point of origin only a litle forward
of the mid-point of the base of the dorsal fin; and
second ray of the pectoral fins branched. The
species heterurus has a plain gray dorsal fin; it has
no teeth on the palatine bonej in the roof of its
mouth; and the pale edging of the outer margin
of its pectoral fins is extremely narrow. The

number of its scales is distinctive, also, 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 Gvlj of 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 thf 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.

FiotTBE 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

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

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

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 110 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

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

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

8,

10.

11

12.

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

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 Radcliffe.

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 Vinal 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 themselves, 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 1920.24 Doctor Huntsman informs
us that spent fish frequently strand on the beaches
on both sides of the Bay of Fundy in September.
We 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.26
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 1936,2"
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. w 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 this 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 29 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 11th of that

» Nichols and Breder, Zoologies, N. Y. Zool. Soc, vol. 9, 1927, p. 163.
■ For details, see Bigelow and Schroeder, 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.

•' Specimens trawled by Albatroti II.

" AlbatroM 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, Zoologies, 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 which 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 Radcliffe
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.

» These catches were all made either at the surface or in oblique hauls
with open nets.

» Kuntz and Radclifle (Bull. V. 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.36

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 third summer.38 Fish of 11
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

» Qoode, Fish, and Fish. Ind. U. 8., Sect. 1, 1884, p. 242.

*• Fry taken In February of different years by Albatross II have ranged
from lii inches (31 mm.) long to i% inches (120 mm.).

« In April 1930 Albatross II trawled young fry ranging in length from 2
Inches (54 mm.) to 6% inches (163 mm.) long at a number of stations from
the offing of Rhode Island to the offing of Chesapeake Bay, at 14 to 85 fathoms.

» For further details, see Bigelow and Schroeder (Biol. Bull., vol. 76, 1939,
pp. 319-320, flg. 8).

" Information supplied by Wm. A. Ellison, Jr.

" Belloc, Notes et Memolres No. 21, Office Scientlflque et Technique des
Pfiches 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 (Merluccius 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 41 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 840, niinimum 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 Evermann (Bull. 47, U. S.
Nat. Mas., Ft. 3, 1898, p. 2530). 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. PGches 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 1 1 fish,
contrasting with an average catch of 232 fish per
haul at 60 fathoms and deeper (25 hauls).46

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 6.

*> 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
161 flsh from deeper than 60 fathoms on the northern part of the bank; 90 fish
per haul from shoaler than 30 fathoms and 286 flsh 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 15-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
Jul}7 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

*&T °°

/ .•

/ :

/ :

/ .•

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-drifting 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 hydrographic 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.
1 Ooode, Fish. Ind. U. 8., Sect. 1, 1884, p. 242.

<• Canad. Fish. Eiped. (1914-1916) 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.4'
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 aDd 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, aod 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 beeD able to learn how early silver hake ap-
pear on the coast of Maine east 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: 51 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 Blgelow and Schroeder, Biol. Bull. vol. 76, 1939,
pp. 308, 319-320, fig. 8.

" 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 Fimdy
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 K70 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: M

1,400
2, 255
1, 700
7, 540

May.. _ 860,000

June 1, 158,000

January..
February.

March

April

July 4,444,000

August 4, 879, 000

September 1,974,000

October.. 2,381,000

November 438, 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

u Pounds of round fish and dressed fish combined.

" Practically no trawling is done m 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 w 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,56 and smaller amounts
in Delaware.

Eastward from our limits we find the silver
hake described as abundant67 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 Euaene H., Capt. Henry Kllmm,
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.

" 1942, 5,343,300 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 3,468,200 pounds In 1942; 5,243,700 pounds In 1945; and 7,498,600
pounds In 1947. Delaware trawlers reported 203,500 pounds for 1947.

»' Vladykov and McKenzle, Proc. Nova Scotia Inst. Scl., vol. 19, 1935, p. 72.

w 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 are meant when "hake" are
mentioned In the early publications of the TJ. S. Fish Commission, such as
Balrd's (Eept. TJ. S. Coram. Fish. (1886) 1889, app. A.) report on the fisheries
of eastern North America.

» 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, 875, 000
7, 943, 000
6,936,000
6, 379, 000
8, 678, 000
15, 420, 000
21,038,000

1938

24,851,000

1939

27, 539, 000

1929

1940 _

39, 990, 000

1930

1942-.

45,900,000

1931

1943

48,460,000

1932 .-

1944...

47, 373, 000

1933 ._

1945

73, 866, 000

1935

1946

48,844,000

1937

1947.

58, 936, 000

Cod Gadus ccdlarias Linnaeus 1758 6I
. r , iwc>r)ii*£t 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
wide, 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. TJ. S. Comm. Fish. [1928], Pt. 2,
1930, p. 210) use the species name morrhua Linnaeus 1758. But the use of
caUarias accords better with modern practice, because It preceded morrhua
on the same page of the Systema Naturae.

Number of flnrays

Dorsal

Anal

First

Second

Third

First

Second

13
15
16

19
21
24

18
19
21

20
22
24

17

18

22

Figttee 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 are
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
size. A huge one of 21 1 % pounds and more than 6
feet long, was caught on a long line off the Massa-

chusetts coast in May 1895; 62 one that weighed
138 pounds dressed (hence must have weighed 180
pounds or more alive) was brought in from Georges
Bank in 1838; and Goode M 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

Pounds

19 to 20-

2W-3

3W- 4

4H- 7

5-7

7-9

7H-10

9 -13

12H-17H

16 -23

18 -22

16 -32

29H-32

31 -51

50

54

20 to 21

3 -3H
4-5)4
6M- 8
7 -8M

23 to 24

23 to 24

25 to 26

25 to 26.

27 to 28

27 to 28

30 to 31

30 to 31..

32 to 33

32 to 33. _

34 to 35

34 to 35

36 to 37

1254-17

36to36H

38 to 39

38 to 39

40 to 41

40 to 42

43 to 45

25VS-29

43 to 44

46

48^ to50H

52

57^

A 99%-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

u Jordan and Evermann, American Food and Game Fishes. 1902, p. 514.

•» Fish Ind. U. 8.. Sec. 1,1884, p. 220.

M Based chiefly on measurements given by Earll (Kept. XJ. 3. 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. 8S), 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.96 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 M 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 (1886) 1887, p. 327), Mcintosh and Masterman
(British Marine food fishes, 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 Mlnlst. Agrlc. 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 then-
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
fileus). 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. 184). They
also pick up flounders, cunners, rock eels (Pholis) ,
blennies, 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 M 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 Merrinmn, Copeia, 1944, p. 198.

•' Olsen and Merrlman, Bull. Bingham Oeeanogr. 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.

n Schmidt (Skrift. Komm. Havunderstfgelser, No. 1, 1904, p. 70) and
Paulsen (Meddelel. Kommls. Havunderstfgelser, Serie Plankton, vol. 1,
No. 8, 1909, p. 39).

« Bateson, Jour. Mar. Biol. Assoc. United Kingdom, N. Ser., vol. 1,
1889-90, p. 241.

186

FISHEKY 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, bike 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

" In European seas young cod often live under the disks of the large red
Jellyfish (Ci/anca), but they have not yet been found In this situation in the
Gulf of Maine.

" See especially HJort, Journal du Consel], Cons. Perm. Interaat. Eiplor.
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 1923-1930,76 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 rogion of Nantucket Shoals, and about 30,000 in
other parts of the Gulf of Maine, Including the offshore Banks, 308 fish
recaptured westward from Marthas Vineyard,. For further details, see
Schroeder (Bull. TJ. 8. Bureau of Fisheries, vol. 48, 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.78

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

'• 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.

it Dannevig, Canadian Fisheries Eiped. (1914-1915)., 1919, p. 44.

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 87. — Egg. After Heincke and Ehrenbaum. Figure 88. — Larva, just hatched, 4 mm. After Masterman.

Figure 89. — Larva, 4.5 mm. After Schmidt.

Figure 90. — Larva, 9 mm. After Schmidt.

Figure 91. — Fry, 20 mm. After Schmidt. Figure 92. — Young, 40 mm. After Schmidt.

Cod (fladus 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.

Bate 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

771 771.

29 to 38....

34 to 49....

35 to 51....

771771.

32.9
40

42.8

May 25

June 6..

June 20

771771.

28 to 68....
71 to 76....
73 to 77....

771771.

64

76.5
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 their 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

» Bumpus, Science N. Ser., vol. 8, 1898, p. 852.

» Smith, Bull. U. S. Fish Comm., vol. 19, 1901, p. 307.

™ Damas (Rapp. et Proces-Verb., Cons. Perm. Intern. Explor. Mer, vol.10,
No. 3, 1909) gives an account of the European investigations 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's81 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

3

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
12

4

15

5 ...

19

6...

21

7

24

8

27

g

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 3M 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. U. 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 wbile
the cod are essentially fish of tbe 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 tbe southwestern
part of tbe 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 8S 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

21,698,594

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 88 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. 8. 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.
M Bull. U. S. Bur. Fish., vol. 43, 1929, pp. 266-290.
" Bull. U. 8. 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 1% 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, locky grounds, or among algae, they have
a fair chance of escaping their various enemies,
but that they 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
line 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
majority 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 Manan
(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.81
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.'0
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. 8.
Bureau of Fisheries vessels Halcyon and Albatrcst 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. 2CM5, charts 1-8) for detailed discussion In relation to spawning and
to racial subdivisions of the local stock.

•' 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. Interaat. 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 shoal er, 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 IK 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.96

Cod spawn in shoaler 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 this, 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."
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 was then

" See HJort (Rapp. Prou.-Verb., Cods. Perm. Internat. Ejplor. Mer.,vol.
20, 1914).

" Smith, Rept. U. S. Comm. Fish. (1901) 1902, p. 208; Schroeder, Bull.
U. 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. H. Thomas, formor superintendent of the Woods
Hole hatchery.

" Information from C. G. Corliss, former superintendent of the Gloucester
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

1912-13
1913-14

Off Bockport (Ipswich
Bay).

Feb. 16 to Apr. 7.
Feb. 1 to Apr. 15.

1914-15

1915-16
1916-17

In Ipswich Bay and off
the New Hampshire
coast.

Feb. 9 to Apr. 13.

Feb. 27 to Apr. 13.
Feb. 25 to Apr. 27.
Feb. 27 to Apr. 30.
Dec. 28 to Apr. 30.
Jan. 15 to Apr. 29.

1917-18

1918-19

.... do

1919-20

do

1920-21

do

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 only 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 M reports the
first ripe female as taken near Cape Ann on
September 2 during the season of 1878-79, while
we have taken cod eggs, far enough advanced in
incubation for positive identification as such, off
Shelburne (Nova Scotia) on September 6; near
Mount Desert on September 15; and off Penob-
scot Bay on October 6 (all in 1915).

•' This (act 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. TJ. 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 Gram-pus
(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
prevail 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°— 13° 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°— 12°
(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. l'Eiplor. Mer, Pub. de Clrconstance, No. 8, 1904).

• Dannevlg, Canadian Fish. Eiped. (1914-15) 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°-37°). 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°-40° F. (1.5-4.4° C.) for spawn-
ing, with the chief production of eggs taking place
at 37°-41°F. (3-5° C).*

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
mille, 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. Eiped. (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
Nantucket 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 Albatrosf 11
towed several hundred little cod (4 to 9>4 mm.) off the tip of Cape Cod near-
by, on May 28, 1927. Tho paucity of our other catches of cod larvae (80 to
90 all told) for other parts 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 1944, about 8,226,000 pounds in 1945, and about
8,174,800 pounds in 1946.

1 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 into 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
directus) 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, cockles (Polynices), or on pieces

» Knight, Contrib. to Canad. Biol. (1906-1910) 1912, pp. 23-32.

" For account of cod fishing 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, paliDg 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 \)i 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 u tomcod are 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

» Nichols and Breder (Zoologies, N. Y. Zool. Soc, vol. 9, 1927, p. 166) state
that tomcod up to 10?i inches long were common throughout that summer in
Sandy 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 Gulf of Maine. — The tomcod is
locally common around the entire coastline of the
Gulf. It is reported at Pubnico and in St. Mary Bay,
for example, on the west coast of Nova Scotia ; at
various localities on both shores of theBay 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 " remarks.

Westward and southward from Cape Cod, the
tomcod is plentiful in suitable situations all along
the coast to New Jersey, where Abbott M 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 highly 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 theNova 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; IB 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. 8. Comm. Fish., (188S) 1887, p. 623, pi. 13, fig. 67) de-
scribes and pictures the newly hatched larva of the tomcod.
" Contrib. Canadian Biol., (1921) 1922, p. 67.

» Geol. New Jersey, 1868, p. 818.

•» Contrib. Canadian Biol., N. Ser., vol. 7, No. 16 (Ser. A, general, No.
13). 1932, p. 7.

'• Most recent year when tomcod were mentioned In the United States
catch statistics for the Oulf of Maine coast.

» 35,000 pounds of tomcod were reported for Dlgby County in 1944, bul
this amount Is so much larger than for preceding years, or for 1948, as to suggesi
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 usually 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 only 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 21 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 M 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 35% inches long, weighing about 16%
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 inches, 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 Intemat. 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

WllSHl

in

/

/

/

/

14

/

/

/ J

f /

4

/

13

./

/

/ A

12

/

/

/

II

/

/

• >

/
/

■S

S

«

7

,'

23 26 27 28 29

LUISTH. INCHES

30 31 32 33

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-

» 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. Internat.
Explor. Mer, vol. 10, 1909) and by Schmidt (ibid.).

1 Thompson, Research Bull. No. 6, Newfoundland Dept. Nat. Res., 1939,
p. 9.

" Goode and Bean (Smithsonian Contrlb. Knowl., vol. 30, 1895, p. 364) 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 five 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

In general, the haddock five 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 of 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 chiefly 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 68° 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 ol 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.

» Sec Homans and Needier (Proc. Nova Scotian Inst. Scl., vol. 21, 1946,
pp. 15-49) (or a study of the haddock.
- Needier, Copela, 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,"38 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. 6, 1930, No. 10
p. 7.
" Rept. tJ. 8. 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

Quit of
Maine

North
Sea

Norway

Length,
Inches

Length,
inches

Length,
inches

2_

12

17.5

19

21

22.5

24

25

10
12
16

17.5
20
22
24.5

10 6

3

13

4

15.5

6

17.5

6

19 6

7

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

3K

4H

7Ji

syt.

Average length, Inches

St. Andrews

ISM

20K

22H

24

25H

26Ji

Nantucket
Shoals

18Ji

•mi

22

23^
25
25 Jf

Browns
Bank

18)J
195i
20H
21%
22Ji

Eastern
Nova
Scotia

19Ji

21

22J4

24

25)i

According to Thompson 3g 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 *° gave about 12K inches as the length
of 2+-year-old haddock and 13% inches for 34-
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^ 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 K- 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 %- 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. 54 [295], fig. 17, p. 65 [206].

" 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. 6, No. 19, 1942, p. U.

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
28K 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 M 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 ** 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.

*> Unpublished manuscript.

« Contr. Canadian Biol. (1914-1915) 1916, p. 39.

» 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.48 And while the
experimental trawling campaigns of the Newfound-
land Fishery Kesearch 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 oj Maine. — Haddock are
very plentiful all around the open Gulf, as well as
on all the offshore banks, especially on Georges
where they greatly outnumber 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; therosefishis 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 than5 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,60 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 (Unders<gelser over den Grjlnlandske 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.

" 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
€0 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.61

In 1945 (most recent year for which detailed
statistics are available 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; 62 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

*' Information contributed by Clyde C. Taylor of the U. S. Fish and
Wildlife Service.

*» 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, 1051.

•» Herrington (Fishery Circular No. 23, TJ. 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.68 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-
orn part

Northern
edge

Central and
southeast*
era part

Southwest-
ern part

1942

19
17
20
31
26
19

39
27
37
24

35

40

36
46
35
24
29
33

6

1943

11

1944

9

1945

22

1946

11

1947

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], flg. 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, flg. 1) chart of the distribution of the haddock catch, 1917-
1925.

•' Rich (Rept. U. S. Fish Comm. (1929) 1930, App. 3, pp. 61-117) 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 Gulf
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

Off Chatham

339,000

' 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 studies M
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 8I 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 April
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.

Breedirg 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 tune
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 eggs83 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 flounder, none being
sufficiently advanced in incubation to show the pigment.

13 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."* But
their 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-fathom contour on the southeastern slope of
Georges Bank at the height of the breeding season
(late March 1931)66 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

1919

Apr. 9 to Apr. 23

Feb. 20 to Apr. 23
Mar. 25 to Apr. 25
Jan. 27 to Apr. 14

32, 380. 000
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. Fish., vol. 49, Bull. 29,

8, p. 16, flg. 7.

« Earll, Rept. U. S. Coram. 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 w 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.68

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 flotation of the
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

« Canadian Fish. Eiped. (1914-15) 1919, p. 21.

" Damas, Rapp. et Proc.-Verb. Cons. Internat. Explor. Mer, vol. 10, 1909;
Schmidt, ibid.

any of the haddock eggs produced in the Gulf fail
to rise from the bottom.69

Populations and migrations within the Gulf of
Maine. — Needler's70 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

89 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. V. S. Bureau of Fisheries, vol. 49, Bull. 29, 1938, pp. 13-15.

*> 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.

T3 One fish that was tagged by us near Mount Desert Island was recaptured
in the Nova Scotian side of the Bay of Fundy off 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, " 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-

'< 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).

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.

Nothing 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.78

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

» 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 1 14 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 Stellwagen 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."

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. U. S. Fish Coram, for 1929, 1930, App. 3) gives information
In this respect.

" As happened In 1928 (Needier, Contrlb. Canadian Biol, and Fish., N.
Ser., vol. 6, No. 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 Lauren tian 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 1920.8* 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."

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
area owing to the prevailing currents."88

" For further discussion, see A. W. H. Needier, Contrlb. Canadian Biol,
and Fish., vol. 6, No. 10, 1930, and A. B. Needier, Bull. 25, Biol. Bd. Canada,
1931.

to Research Bull. No. 6, Newfoundland Dept. Nat. Resources, 1939, p. 7.

•J 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 [436].

" 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 data 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."0 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 survival 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 tbe 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

88 From unpublished data for Georges Bank and the South Channel area
supplied by Howard A. Schuck.

•° For details, see Watford'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.

»' Tians. 9th North American Wildlife Conference, 1944, p. 260.

" See especially Thompson's studies for Iceland (Fisheries Scotland, Sci.
Invest. [1928], No. 6, 1929), and Raitt's for the North Sea (Journal du Conseil,
Cons. Internat. Explor. Mer, vol. 11, No. 2, 1936, p. 211).

M Fish that are iced at sea, not salted.

" Herrington, Fishery Circular No. 23, XJ. 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 ?4-}i 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.

M For tabulations of the total catches of haddock in the western Atlantio
by Canadian and United States vessels, 1880-1927, see Needier, Contrib.
No. 2, North American Council on Fish. Investigations, Ottawa, 1929, 13
pp., also Rept. 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."7
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 Gulf

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
6,200.000
4,900,000

11, 600, 000
6, 600. 000
6,600,000
5, 100. 000
4,700,000
8.200,000
7,200.000
7, 100. 000
6,600,000
6,900.000
4,600.000

5, 200. 000

6, 400, 000
6. 200, 000

194, 400, 000

1934

66, 600, 000

1935

89, 700, 000

1936

97, 200, 000

1937

107, 100, 000

1938

113,900,000

1939.

114, 100, 000

1940

104, 100. 000

1941

128,100,000

1942

112,400.000

1943 ..

97, 100. 000

1944 .

96, 400, 000

1945...

83. 900. 000

1946...

109, 700, 000

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. 5, 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 M 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 (Oadus pollachiui)

214

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Length, in

Weight, in

Length, In

Weight in

inches

pounds

inches

pounds

24H

4-5V*

30

8K-9H

26

4

31

10

27

7H

31«

10

27H

iVi

32

10-12

28H

8

33

12

29

8-9

33

14

29H

8^-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 ps 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 1% 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 (Rept. 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
Meganycliphanes and Thysanoessa) are very abun-
dant all summer, Kendall * 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 dn 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 * 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,

• Smltt, Scandinavian Fishes, vol. 1, 1892, p. 503.
« Eept. U. S. Comm. Fish., (1896) 1898, p. 180.

• Proc. Amer. Acad. Arts, Sci., vol. 66, 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 they 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-water 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. 218").

The pollock is a late autumn and early winter
spawner, 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. V. 8. Comm. Fish., vol. 22, 1904, p. 268.

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
the pollock spawns on a falling temperature, with
most of the eggs produced within a comparatively
narrow range and in water several degrees warmer
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 6X 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% years; about 29

» For resume1 see Damas (Rapp. et Proc. Verb., Consell 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 mm. After Schmidt.

Figure 99. — Egg (European). After Mcintosh.

Figure 102. — Larva (European), 12.5 mm. After Schmidt.

Figure 100. — Larva (European), 5 days old, 4.3 mm.

After Mcintosh. Figure 103. — Fry (European), 23 mm. After Schmidt.

American Pollock (Pollachius virens).

inches at 8% years, and about 30 inches at 9%
years. Fish of 3 feet and upward are therefore of
considerable age. The oldest recorded by 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 first
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
and northeastern Nova Scotia to New Jersey;
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

to Sandwich Bay on the southeastern coast of
Atlantic Labrador;11 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 Oulj of Maine. — In our side of
the Atlantic the pollock has its chief center of
abundance in the Gulf of Maine, where it is caught
in large numbers both on the offshore banks, and
all around the coast line, from Nantucket 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 1946,12 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

1 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.
V. S. Bur. Fish., vol. 43, pt. 1, 1928, p. 156) and from Cape Lookout by Coles
(Copeia, No 151, 1926, p [105]).

» 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.

» 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; :3 Nova Scotian
shore of Bay of Fundy, about 500,000 to 1,000,000
pounds; u 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; IS 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, 1946; Canadian catches 7,017,000 pounds In
1944 and 6,642,000 pounds In 1946, besides an Indeterminate amount landed
along this part of the Shelbume County coast line.

» 1944, 513,000 pounds; 1946, 983,000 pounds.

'• Charlotte and St. Johns Counties, about 2,000,000 pounds in 1944, about
3,507,000 in 1946.

'• 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 ofeconomlc reasons
that the fish 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 during 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 fine fishermen from Jeffreys
Ledge throughout the summer, most of them
caught some distance above bottom.

North of the Isles of Shoals, pollock are 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 for
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 towings. 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

•• 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 O. Q. 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 Bay 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-
uary 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...

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

..do

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.l7toJan. 16

Nov. 21 to Jan. 16

2. 081, 400, 000

1918-19-

1,110,470,000

1919-20

954 800 000

1920-21

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 i9
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.
Whether 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 caught 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 }i 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.2*

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 26

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

31 Two pollock tagged on Nantucked Shoals in June and October were
caught off Block Island in the following May and July respectively.

23 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, Pt. 1, 1925,
p. 406.

11 No statistics are available as to what proportions of the Canadian catch
In the Gulf are made with the different kinds of gear.

" The European hake is a Merlucciut (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, with much larger eyes than the cod but with
smaller chin barbels. Furthermore, they have
only two dorsal fins, the second much longer
than the first, and only one anal fin instead of
the three dorsals and the two anals of the pollock,
cod, and haddock. The ventral fins are long,
narrow, and feelerlike.

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 (apparantly 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 alhed 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,26 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 youDgest 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.

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

17 Zoologica, N. Y. Zool. Soc. 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
fives, 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 Research
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 x 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 found in
hake stomachs at Woods Hole,29 such as alewives,
butterfish, cunners, 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.) in 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 off 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. Bu t 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 shrimps (Meganyc-
tiphanes and Thysanoessa) that are so plentiful
there, while the general character of their diet is
sufficient evidence that they 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 fair to assume they do most of
their foraging between sunset and sunrise.

» Cioode, (Fish. Ind. U. S., Sect. 1, 1884, p. 235); Kendall, (Rept. U. S.
Oomm. Fish., (1896) 1898, p. 180); Linton, (Bull. U. S. Fish Comm., vol. 19,
1901, p. 478); Hansen, (Proo. U. S. Nat. Mus., vol. 48, 1915, p. 94); Breder
(Zoologica, N. Y. Zool. Soc, vol. 2, No. 15, 1922, p. 350); and Vinal Edwards'
notes.

" A large white hake taken at Woods Hole in May 1908 had a fish (tepo-
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
Cole, Bull. U. S. Bur. of Fish., vol. 31, pt. 2, 1913, p. 768).

* General Historie of Virginia, New England, and the Summer Isles, 1616,
ed. of 1819, vol. 2, p. 188.

210941—68 18

Herrick 3I 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 five, suggests that they rec-
ognize shrimps, crabs, aDd 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 unit 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-
ficially at the Gloucester hatchery.

Figure 107. — Squirrel hake (Urophycis 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 4K inches (60-1 10 mm.)

" Fry figured by A. Agasstz (Proc. Amer. Acad. Arts, Sci., N. Ser., vol. 9,
1882, pi. 7, fig. 6; pi. 8, figs. 1-3), as Motella argentala 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 inches 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 Bodie 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, many 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.36

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,39
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.

n 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.

ActuaUy, 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 Gulf 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. This 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

17 The tatter were listed by Welsh as V. regiut, but probably they were
white hake.

* About 13,000 pounds of white hake were reported from 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 Gulf, 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 cai-ry 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 year (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. 1 78) 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* 66' 68*

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Figure 109. — Locality records for squirrel hake eggs (•), and for larvae of rockling (A) in the Gulf of Maine.

230

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

30-inch female, with large roe, trawled off New
Jersey (lat. 39°23' N.) by the Albatross II at 88
fathoms, on February 27, 1929; Welsh saw a male
with milt flowing, near Gloucester on April 22,
1913; we saw spent females, but also a male just
ripening, trawled by the Albatross III on the
southwestern part of Georges Bank on May 15,
1950; and Nichols and Breder41 report a ripe
female with ripe eggs at Woods Hole in July.

The height of the spawning season of the squirrel
hake comes at least as early as June south of Cape
Cod, and through early summer in the Massa-
chusetts Bay region, judging from the size (27 to
70 mm.) of the fry mentioned above as found
in scallop shells in late summer and autumn (p.
224). The extreme limits of the season are not
known. But we have towed eggs of this species as
early as June 10 in Petit Passage, Nova Scotia,
and as late as September 20 in various parts of the
Gulf, while captures of fry of 72 mm. as early as
the last week in July (in Sbelburne Harbor, Nova
Scotia), and others as small as 36 mm. in the west-
ern part of the Gulf as late as November 1 (in
1916), point to a spawning season lasting from late
spring until early autumn.

Abundance.— There is nothing in the statistical
picture to suggest that hake of either species
fluctuate very widely in abundance in our Gulf
from year to year, for the ups and downs in the
amounts caught are not greater than can be
charged to market conditions. Neither has any
attempt been made to estimate the periodic varia-
tions in the relative abundance of different year
classes. Earlier characterizations of the numbers
of the two hakes in our waters have been in relative
terms, ranging from "common" to "in immense
numbers."42 More precise information is that the
Atlantis caught an average of about 32 tenuis per
hour's trawling with an 82-foot trawl at 65 to
125 fathoms (120-228 meters) off Cape Cod, west
of Jeffreys Ledge and off Mount Desert, in
August 1936,43 while the Albatross III trawled
about 245 chuss and about 18 tenuis per hour's
hauling on the southwestern part of Georges Bank
and south of Nantucket Shoals in May 1 950. And
we find record of 145 tenuis taken per hour's
trawling,44 on Sable Island Bank, off outer Nova
Scotia.

« Zoologies, N. Y. Zool. Soc, vol. 9. 1927, p. 172.

« Huntsman, Contrib. Canadian Biol., (1921) 1922, p. 68.

« Bigelow and Schroeder, Biol. Bull., vol. 76, 1939, p. 308.

« Report, Newfoundland Fish. Res. Comm., vol. 1, No. 4, 1932, p. 109.

Importance.— The hakes are soft-meated and
have rather poor keeping qualities, but both the
white and the squirrel hake are readily absorbed
by the fish markets if they are large enough, and
great numbers of small squirrel hake are now used
for mink and poultry feed. A quarter of a cen-
tury ago the yearly catch in the Gulf ran between
20 and 35 million pounds, and it has been much
the same of late years (1941-1946), with yearly
landings by Canadian and United States fishermen
of between 19 and 30 million pounds. In 1946,
which may serve as representative, Canadian
fishermen landed about 2,100,000 pounds in outer
Nova Scotian ports (Cape Sable to Cape North),
about 4,800,000 pounds along the southern shore
of the Gulf of St. Lawrence.

At the present time (as represented by 1946 and
1947) 4 to 5 times as much hake is marketed in
Maine and Massachusetts in the form of fresb and
frozen fillets as is marketed there salted, some
are used for fish cakes, and a very small part 46 as
smoked fillets. Hake sounds (swim bladders),
especially of those tbat are caught off Nova Scotia
in deep water, are also used to make isinglass,48 and
increasing amounts of small squirrel hake brought
in from Nantucket Shoals, are utilized from j^ear to
year in the trash-fish industries.47

Roughly two-thirds of the poundage of hake
that is landed in Maine and Massachusetts are
caught in otter trawls nowadays, roughly one-fifth
in gill nets, and only a little more than one-eighth
on long lines.48

The hakes are such dull and inactive fish that
they are of no special interest to the angler. But
a good many fair-sized ones are caught hand-lining
from party boats, for they bite readily, and small
hake are caught from small boats in harbors and
bays, along the Maine coast especially.

Spotted hake Urophycis r^ffkrs (Walbaum) 1792

Jordan and Evermann, 1896-1900, p. 2553.

Description. — This species is distinguishable
among the hakes of the Gulf of Maine by the fact

« About 39,000 pounds reported for 1916, none for 1947.
** For further information see Tressler, Marine Products of Commerce,
1949, pp. 489-191.

" Landings of red hake at New Bedford from the Nantucket Shoals region,
mostly used in this way, were about 6,600,000 pounds in 1947.
" The amounts recorded for 1947 were: Poundt

Otter trawls -- 10,399,800

Gill nets 3,380,200

Long lines.- - 2,097,200

Hand lines 102,200

FISHES OF THE GULF OF MAINE

231

Figure 110. — Spotted hake (Urophycis regius). From Jordan and Evermann. Drawing by H. L. Todd.

that it has no prolonged rays in its first dorsal fin
(which is hardly higher than the second dorsal, and
has 8 or 9 rays) ; by the smaller number of rays in
its second dorsal fin (46 to 51 as against 54 or more
in the squirrel and white hakes); and by having
only 90 to 95 oblique rows of scales along its sides
from gill opening to caudal fin, instead of about
105 to 110 rows and about 140 rows, respectively,
in the other two species. The anal fin has 43 to
49 rays, somewhat fewer, on the average, than the
squirrel or the white hake (48 to 50) .

Convenient field marks are that the outer half
of its first dorsal fin is black with a whitish margin;
that its pectoral fins reach back as far as the origin
of the anal fin, whereas they fall considerably short
of the latter in both the white and the squirrel
hake ; and that its lateral line is darker brown than
the general body color, instead of paler, and is
interrupted by a series of distinct whitish spots.
Otherwise the spotted hake, like the commoner
hakes, is dull brown, darker above than below,
with dorsal and anal fins of the same color as the
back. Its ventrals are whitish.

Size. — The largest of many measured by Welsh
at Atlantic City, in August 1920, were about 16
inches long, and weighed between 1 and \){ pounds;
the usual length is less than 12 inches, and the
longest, of about 600 taken by the Albatross II at
14 stations between the offing of Delaware Bay and
Cape Hatteras, in late winter and spring (1930 and
1931) was 5% inches (130 mm.).

Habits. — The spotted hake resembles the other
hakes in its habits. It may be more of a fish eater,
for Vinal Edwards noted that the few he examined
at Woods Hole contained alewives, menhaden,
launce, and squid. But it also feeds on the crus-
taceans on which the white and squirrel hakes
subsist, for Hildebrand and Schroeder49 found

mysid-shrimps in most of those examined in Chesa-
peake Bay where small spotted hake are very
common. The capture of spawning fish by the
Albatross, off the coast of the Carolinas in
December in 1919, recorded in Welsh's field notes,
is evidence that it is a winter breeder.

General range. — Coast of the United States,
regularly from southern New England and New
York to Cape Hatteras (including Chesapeake
Bay where it is plentiful), and ranging southward
as far as the offing of northern Florida in deep
water.60

Many were trawled as far northward as the
offing of Delaware Bay by the Albatross II, in 1930
and 1931; it is reported as rather uncommon at
New York;61 it has been taken occasionally at
Woods Hole; it has been known to reach the coast
of Maine as a very rare stray; it was reported more
than a century ago off Halifax, Nova Scotia, by
Richardson ; 62 and a single specimen was reported
as taken, pelagic, near Sable Island (lat. 44°10',
long. 59°45') in August 1931."

Occurrence in the Gulf of Maine.- — The spotted
hake strays past Cape Cod so seldom that speci-
mens taken off Seguin Island many years ago,
and four, trawled on the southwestern part of
Georges Bank, by the Albatross III in May 1950,
are the only definite records of it for the Gulf of
Maine.64 But it may well have been overlooked
among the hosts of young hake of the two common
species (white and squirrel) that are caught in

« Bull. U. S. Bur. Fish., vol. 43, pt. 1, 1928, p. 161.

» The U. S. National Museum has specimens taken off 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.

a Report, Newfoundland Fishery Res. Comm., vol. 1, No. 4, 1932, p. 109.

M This species was also listed from Ipswich Bay, from Casco Bay, and off
ofMonhegen Island in the Grampus 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-finned 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.

Habits.- — 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.66 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.5*

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. 276.
■* For early locality records see Goode 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 110 to 140 fathoms, in 1878; and of a few
others that were trawled on the northern edge of
Georges Bank by the Kingfisher, in 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 IIM 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 Gunther 1878

Jordan and Evermann, 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 in 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 in some but forming an acute angle in
others is distinctive, likewise its vent is situated
much farther back than in 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 description of the
species (Proc. U. 8. Nat. Mus., vol. 1, 1878, p. 256).

« Reported by Blgelow and Schroeder, Bull. U. S. Bur. Fish., vol. 48, 1936,
p. 339.

Size. — The longest yet measured was one of
about 2VA 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. w

Hakeling Physiculus jvlvus Bean 1884

Jordan and Evermann, 1896-1900, p. 2547.

Description. — This fish is hakelike in its general
appearance, also in the general arrangement of its

»' See Qoode and Bean (Smithsonian Contrlb. Knowl., vol. 30, 1895, pp.
374-376) for list of stations.
- For a recent summary, see Schroeder, Copela, 1940, No. 4, pp. 236-237.

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

Figure 113. — Hakeling (Physiculus fulvus), outer edge of Continental Shelf off Nantucket.

Drawing by H. L. Todd.

From Goode and Bean.

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 tbe 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 Qulj 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
(lat. 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, the 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 (Loiclla maxillaria) has been taken off 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

^#S&ss

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

•' 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 fining 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 1 2 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 64
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. 63)
and further Information contributed by him.

« Ooode and Bean, (Smithsonian Contrlb. 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 flat. 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 66°.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 only one post anal band
of black pigment; and by the short stocky body-
form. Young hake are more slender and have
scattered pigment; young cusk 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 their 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 (Contrlb. Canadian Biol., N. 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.

97 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 (Qaidropaarus aTgentatut 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 argen-
lotut, with excellent illustrations, see Jensen, Spolia Zool. Mus. Hauniensis,
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), otherwise
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 this 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.

■ A specimen trawled by the Albatross 12 in 12 fathoms off the mouth of
Chesapeake Bay on February 10, 1930, is the only one recorded in shallow
water so far southward.

» Goode and Bean (Smithsonian Contrib. Knowl., vol. 30, 1895, p. 381)
give a list of these localities. For a recent account of G. 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. Essei 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.

Huntsman 72 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
Dannevig75 (1919) records rockling eggs (prob-
ably this species) as early 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 K to 9 fathoms, and
abundantly at 21 fathoms by the Fish Hawk
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 Oat. 35° 40' N.).77

n Contrib. Canadian Biol. (1921) 1922, p. 69.

>» Contrib. Canadian Biol., Fish., N. Ser., vol. 6, No. 6, 1930, p. 13 [1191-
» It spawns from the end of January until August In the Baltic.
« Canadian Fisheries Erped., (1914-1915) 1919, p. 53, table 1C
« Nichols and Breder, Zoologica, N. Y. Zool. Soc, vol. 9, 1927, p. 172.
" For list of early stations, see Goode and Bean (Smithsonian Contrib.
Knowl., vol. 30, 1895, pp. 384-385). They also report a specimen apparently
of this species from the offing of Cape Fear, N. C Oat. 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
Tsle 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 fou^-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 (Muller) 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
3K 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
\% to 2% 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% pounds; 33% 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
1,228 cod; 15 miles southeast of Monhegan Island,
June 24 and 25, 1913, long lines, 580 cusk to 2,880
hake; Jeffreys 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 {Hippoglossoid.es).

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
Guff. 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. Ind. U. S., Sect. 1, 1884, p. 233.

11 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.

•> The records are July 22, 1912, 1 specimen; July 20, 1916, 4 specimens; and
July 22, 1916, 1 specimen.

1 Meddel. Kommis. for Ilavunderstfgelser, 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 mt ms wu wss

,. , • a J i. 1 J „* «„A «„ V,owl Cashes ---- 225,000 98,000 173,000 612,000 1,023,000

tic, chiefly in moderately deep water and on hard Fippemes __ 19000 69,oo0 35,000 47,000 61i000

bottoms; north on the American coast to the riatts - 7,000 6,000 165,000 84,000 45,000

,T - ji j -o 1 J x j.1. a* „•* „f TJ„11„ Jeffrey Ledge- 301,000 143,000 148,000 122,000 53,000

Newfoundland Banks, and to the btrait ot rielle steiiwagen Bank— . 65,000 63,ooo ss.ooo 259.000 78,000

Isle, south regularly to Cape Cod, rarely to

', _, "L, , , , ' , ,T We are mclmed to believe that the wide dif-

southern New England, and occasionally to New ^^ frQm ^ ^ j&^ ^ ^ catcheg oq fchese

Jersey; northern coasts of the British Isles, Den- gmaU ^^^ reflecfc the number of vessels that

mark (Jutland), northern part of the North Sea, fighed there; ratner 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"Matinicus Sou'-

Occurrence in the Gulf of Maine— The cusk is sou'west." In 1921, for example, 43,545 pounds

distributed very generally in the Gulf in water were reported thence, and considerable numbers

deeper than 10 to 15 fathoms, its presence or are taken in the aggregate, on the patches of hard

. , ,. ,. . , , ., bottom that skirt the coast of Maine, as appears

absence depending on the precise type of bottom. . , , , . ,

_, , . , • 1 • / r.o«\ from the approximate amounts landed m the

Because of its preference in this respect (p. 239), smaUer portg 86 ^ ^ m&T&nt Maine counfcies m

it varies greatly in abundance in different parts of igig and ^ ig45. York) g000 pounds and 2600

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 New Brunswick side, and off Brier Island

plentiful off Cape Ann and on Jeffreys Ledge, the on the Nova Scotian Slde' as Docto,r Huntsman

, . , . , ., , ,. , informs us, though none are reported toward the

latter being one of the most productive cusk , , . , ' 6 0 „ , . , , .,

, „ ,, _.. , , , „ , head of the Bay. Small rocky patches along the

grounds of our Gulf. The rocky slopes of Cashes wegfc Nov& g^.^ shore and off geal Island aJso

Ledge, also have long been famous for cusk. In ^ sQme cugk; &nd they &re takeQ regularly OQ
past years when more fishing was done there (as Grand Manan Bank. 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
Platts 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." the rule that cusk, ^ to ,rock/ ^ou^d are ^at
they are at least tolerably plentiful in the co-called

•* 1935 is the most recent year when landings were reported from these

grounds, separately. M 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 contour 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) M and Canadian
fishermen (1944, 1946) combined, of between
215,000 and 250,000 pounds off western Nova
Scotia; " 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 Re-
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 1936^6, 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.96

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

M 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.

•i Recorded by Comlsh (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.

•> For further details and discussion of the status of the cusk as a Greenland
fish, see Jensen (Spolia Zool., Mus. Haunlensis, Copenhagen, vol. 11, 1948,
p. 175).

« Tracy, 40 Ann. Rept. Commiss. Inland Fish. Rhode Island, 1910, p. 159.

•' Abbott, Geol. New Jersey, 1868, p. 819.

FISHES OF THE GULF OF MAINE

243

fact that the cusk (frequenting rough bottom) is
not a good trawl fish. And 80 to 90 percent as
much cusk (pounds) are caught on long lines as
are caught in otter trawls even today (as illus-
trated by 1947), although the yearly landings of
fish of all kinds in Maine and Massachusetts now

are 70 to 80 times as great by otter trawls as by
long lines.96

A few cusk are caught from party boats by
sportsmen hand lining for ground fish in general,
but most of the cusk live too deep to be of any
particular interest to anglers.

THE GRENADIERS. FAMILY MACROURIDAE

The grenadiers are characterized externally by
having large heads, projecting snouts, and slender
bodies that taper to whiplike tails, with no defi-
nitely demarked caudal fin. They have two
dorsal fins, the first high, the second very low
but occupying the greater part of the back. The
anal fin is nearly as long as the second dorsal, or
longer.

The grenadiers are allied to the cod family, in
classification, by the structure of their skull, but
they differ from the cod tribe in having one stout
spine in the first dorsal fin. They are deep-sea
fishes, living on the bottom, loose in texture and
weak swimmers. Many species are known, but
only three of them have ever been taken within
the confines of the Gulf of Maine.

Besides the species described below, three
others, Coryphaenoides rupestris, C. carapinus and
Nematonurus armatus,97 have been taken on the
continental slope abreast of the Gulf and off
southern New England often enough to show
that they are common there below 350 fathoms.
They are typical inhabitants of the deep-sea floor,
never likely to rise shoal enough to come within
the limits of the Gulf of Maine.98 But fish have
a way of straying, and if any grenadier should be
picked up in the Gulf that proves difficult to
identify, we recommend forwarding it either to
the Laboratory of the Fish and Wildlife Service
at Woods Hole, to the U. S. National Museum,
Washington, D. C., or to the Museum of Compar-
ative Zoology, Cambridge, Mass., to be named.
Parr " has recently published a detailed synopsis

" Otter trawlers landed about 499 million pounds of fish of all sorts In
Maine and Massachusetts in 1947; long liners about 7 million pounds.

" According to Parr (Bull. Bingham Oceanogr. Coll., vol. 10, art. 1, 1946,
p. 54) this is the correct name of the grenadier that was reported by Qoode
and Bean (Smithson. Contrib. Knowl., vol. 30, 1895, p. 407) as Hymenoce-
phalui ooodei OQnther, 1887.

M For descriptions and lists of localities where they wore taken during the
early cruises by vessels of the U. S. Fish Commission, see Goode and Bean
(Smithson. Contrib. Knowl., vol. 30, 1895). In June 1949 we trawled about
200 rupettris on the slope off southern Nova Scotia and off the southeastern
face of Georges Bank, at 290-420 fathoms, from Caryn of the Woods Hole
Oceanographic Institution.

" Bull. Bingham Oceanographic Coll., vol. 10, art. 1, 1948.

of all the species known from the western North
Atlantic and from central American seas.

KEY TO GULF OF MAINE GRENADIERS

1. The dorsal spine is perfectly smooth

Long-nosed grenadier, p. 246

The dorsal spine is serrated, with teeth which can be

felt if not seen 2

2. The vent is considerably in front of the origin of the

anal fin; the skin surrounding the vent is naked and

black; the dorsal fin spine is strongly serrated

Common grenadier, p. 243
The vent is close to the origin of the anal fin; the skin
around the vent is scaly and pale colored; the serra-
tions on dorsal fin spine are so fine that they are

hardly visible, though they can be felt

Rough-headed grenadier, p. 245

Common grenadier Macrourus bairdii
Goode and Bean 1877. '

Rat-tail; Marlin-spike

Jordan and Evermann, 1896-1900, p. 2583.

Description. — This grenadier could hardly be
mistaken for any other fish except for one of its
own tribe, so characteristic is its slender body
(flattened sidewise behind the vent and tapering
to a whiplike tail with no definite caudal fin), in
combination with a pointed snout that overhangs
the mouth; very large eyes; and high first dorsal
fin with one large spine ; but very low second dorsal
fin. And it has a chin barbel like a cod (not shown
in the illustration). As noted above, the second
ray of the first dorsal fin is a true spine, serrated
along its front edge with about 15 sharp and very
noticeable teeth pointing upwards.

The first dorsal fin (2 stiff rays, the first very
short, and 1 1 softer rays) is triangular, about twice
as high as it is long; and it originates over the
pectorals, close behind the gill openings. The
space between the two dorsal fins is about as long

' Parr (Bull., Bingham Oceanogr. Coll., vol. 10, art. 1, 1946, p. 37) places
this grenadier in the genus Nezumia of Jordan, 1904. But it seems wiser to
follow the older and more familiar usage here.

244

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

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 foot 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 fathoms 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
macro urid eggs described by European authors8
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 only
two had been recorded there aside from the East-
port and Lubec specimens mentioned above, the

J Proc. U. S. Nat. Mus., vol. 48, 1915, p. 99.

« Ehrenbaum (Nordisches Plankton, vol. I, 1905-1909) summarizes what
little is known of the 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 160 fathoms, the
other from off Gloucester, both of them taken
many years ago. But they must be rather com-
mon on the muddy bottoms of 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.6

Rough headed grenadier Macrourus berglax
Lacepede 1802 8

Rat-tail; Onion-eye

Jordan and Evermann, 1896-1900, p. 2583.

Description. — This species resembles the com-
mon grenadier (p. 243) so closely 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 6 distinct
ridges on the top of its head; that its head is

1 For a list of captures on the continental slope during the early cruises of
the U. S. Fish Commission vessels, see Goode 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 berglai Lacepede, 1802, Jensen (Spolia
Zool., Mus. Hauniensis, Copenhagen, vol. 9, 1948, p. 178) prefers the species-
name fabricii 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, Coryphaenoides Tu-ptstris Gunnerus, 1765, which is commonly
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 fins 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 farther 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

Figure 120. — Rough-headed grenadier (Macrourus berglax), Banquereau Bank.

H. L. Todd.

From Goode and Bean. Drawing by

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, Spitz-
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 9 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 ot
Comparative Zoology.

We have not heard of one, either from Nova
Scotian waters or from the Eastern Channel since
1903; I0 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 their 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 Carj/n of the Woods Hole Oceanographic Institution.

" 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.

and Bean. Drawing by H. L. Todd.

From Goode

FISHES OF THE GULF OF MAINE

247

Color. — Described as silvery gray.

Size. — About 10 inches long.

General range and occurrence in the Gulf oj
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.

-Opah {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, 1888, reporting that a "Sun Fish," identified

'« Reported by Vladykov, Proc. Nova Scotia Inst. Scl., vol. 19, 1935, p. 6.

H This specimen Is In the Museum of Comparative Zoology.

«• Goode and Bean, Smithsonian Contrlb. 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 at Hyamiis, Mass., on September 17, 1928.18

years ago; l8 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

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 Citharichlhys arctifrons, p. 294

The lateral line is arched over the pectoral fin20 3

3. The two ventrals are not alike, the left (upper) being continuous with the anal fin, the rirrht (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 eye-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 6

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

14 We are Indebted to Norman J. Wllimovsky for showing us a copy of
this letter.

'• Vladykov, Proc. Nova Scotia Inst. Scl., vol. 19, 1935, p. 6.

" Radcllffe, Copela, No. 151, 1926, p. 112.

>* Reported by Robert Goffln of the Bureau of Fisheries station at Woods
Hole, Mass.

" Our Eastern Fl3t Fishes, Canadian Fisherman, vol 6, No. 6, 1918,
pp. 788-790.

■ In all the flounders of Ihls 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 Monolene
lartlicauda.

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 bidden by the skin in old fish. The
(Linnaeus) 1758 two pectoral fins are of different shapes, the one

on the upper (eyed) side of the fish being obliquely
Jordan and Evermann, 1896-1900, P. 2661. pointed while the 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; m front 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 are Paler. and are more or less mottled, while
is arched abreast of the pectoral fin. Furthermore large 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-
flatfishes (only about one-third as broad as it is ally is pure 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 to 79 rays), originates close behind the pectorals, among Gulf 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 been looked on as exaggerations we are
glad to be able to give at least one record of a

» Left-handed haUbut have been caught, occasionally. Gulf of Maine halibut in 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,
1941,24 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 of Maine

Length in

Weight in

Length in

Weight in

inches

pounds

inches

pounds

74

215

63

"120

70

16S

42H

30

61

107

42

33

54 to 66

60^

41K

27V£

40 to 42

29

31

12

36

11 to 12

20

2?4

30

9M

27

6'A

24

VA

—

I This fish 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 their 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, however, 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.*8 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 fish was published In the Boston Globe, June 12t
1917. It was bought by the Shore Fish Co.
» Fish. Ind. TJ. S., Sect. 1, 1884, p. 194.
m Reported in the Boston Herald, March 28, 1941.
>• Meddelel. Kommis. Havundersftgelser, Ser-Flskeri, vol. 5, No. 6, 1917.

» Goode and Collins (Fish. Ind. U. S., Sect. 5, vol. 1, 1887, p. 17) give a
readable account.

" Capt. Baldersbeim (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. U. 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° (8° C.)j 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. Bu t 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. Flskerl vol. T,
No. 7, 1925, pp. 17-18) seems to have been the first to bring this to sclentlflo
attention.

* Baldershelm, Happ. Proc. Verb. Conseil Intemat. Explor. Mer, vol. M,
1929, pp. 25-28.

3' For a further discussion of the range and movements of the halibut in
relation to temperature, with references, see Thompson and VanCleve, Rept.
Intemat. Fish. Comm. No. 9, 1936, pp. 22-38.

"Martin and McCracken, Fish. Res. Board Canada, Progress Rept.,
Atlantic Coast Sta: ion, No. 50, 1950, pp. 3-8.

» Capt. Baldersheim described halibut off west Greenland as sometimes
in schools, preying on launce (Rapp. Proc. Verb. Conseil Intemat. Explor.
Mer., vol. 56, 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
grounds.

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) ,36

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.

» Smltt (Scandinavian Fishes, vol. 1, 1882, 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 "ice bird" (probably a dovekie) as taken from a
halibut caught on Georges Bank; and Scudder (Fish. Ind. TJ. S., Sect. 6,
vol. 1, 1887, p. 119) reports Qndlng the skeleton or a gull in tbe stomach of
another.

'« Goode, Fish. Ind. TJ. S., Sect. 1, 1884, p. 196.

•• Meddelelser, Dansk. Komm. Havunders., Ser. Fisk., vol. 7, No. 7,
1925, p. 18.

' Goode and Collins, Fish. Ind. TJ. 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.3' 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
Daturally 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)/3 On the other
hand, halibut spawn regularly in the aquarium at
Trondhjem, Norway, where the eggs have been
fertilized artificially and hatched successfully.*4
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

Fisheries Res. Board Canada, 1946,

» Clemens and Wilby, Bull. No.
p. 312.

* For a general survey of available information, see Taning, Meddelelser
Komm. Danmarks Flsk. 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 (Eapp. Proe. Verb. Cons. Intern. Esplor.
Mer, vol. 39, p. 96, 1926).

<" Goode (Fish and Fishery Industr. TJ. S., Sec. 1, 1884, pp. 196-197),
mentions reports to this effect.

<» Clemens and Wilby, Bull. 68, Fish. Res. Bd. Canada, 1946, p. 312.

« Taning, Meddel. Komm. Danmarks Fisk. Havunders., Ser. Fiskerl,
vol. 10, No. 4, 1936, p. 8.

" Rollefsen, Kgl. Norske Vldensk. 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,46 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 they do not have any oil globule.*9

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 49 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. 409-412.

*' For description of eggs artificially fertilized in the Trondhjem aquarium,
see Rollefsen, Kgl. Norske Vidensk. Selsk. Forh., vol. 7, No. 7, 1934, p. 20-23;
for descriptions of naturally spawned eggs taken In tow nets in Icelandic
waters, see Tanlng, 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, Rept. No. 9,
Internat. Fisheries Comm., 1936.

" 4 to 4.S mm. in diameter according to Jensen, Kgl. Dansk Vidensk.
Selsk. Skr. Nat., afd. 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-sjlgelser, Ser.
Flskeri, vol. I, No. 3, 1904), to Jespersen (Ibid., vol. S, No. fi, 1917), and to
Tanlng, (Meddel. Komm. Danmarks Fisk. Havunders^gleser, Ser. Flskeri,
vol. 10, No. 4, 1936.)

stronger on the right side than on the left, and the
caudal fin (previously rounded) has become square
tipped.

Figure 124. — Larva, 16.2 mm. (European).
Schmidt.

After

Figure 125. — Larva, 22 mm. (European). After Schmidt.

Figure 126. — Larva, 34 mm. (European). After Schmidt.
Halibut (Hippoglossus hippoglossus).

The younger larvae (up to about 25 mm. in
length) are made recognizable as halibut by their
curiously upturned snout. Older ones that are
large enough to show that they belong to some
right-handed large-mouthed flounder are separable
from the American dab larvae (the only other
common Gulf of Maine flatfish with which they
agree in both these respects) by the outlines of the
head and abdomen.

The early life history of the Pacific halibut has
been worked out especially by Thompson and Van
Cleve, who have given an excellent series of illus-
trations of successive stages from newly hatched
larvae to young fry a little more than 1 inch long.60
How long the young halibut lives adrift at the
mercy of the currents, is not known. But the
young fry, so small (47-64 mm long) that they had
evidently been spawned the preceding spring or

*> Rept. No. 9, International Fisheries Commission, 1936, figs. 38-49.

254

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

summer, have been trawled off Iceland during the
last week of July. And the smallest bottom stages
have so far been taken only in water shoaler than
about 27 fathoms (50 meters), evidence that the
larvae of the Atlantic halibut tend both to rise
toward the surface, and to drift inshore during
their pelagic stage, as is also true of the Pacific
halibut.

Fry of 3% to 5% inches (80-150 mm.) such as
have been taken in considerable numbers in Ice-
landic waters in June and July probably are in
their second year. The average relationship be-
tween age and size is as follows according to
Jespersen: 61

Age

1 year

2 years

3 years

4 years

6 years

Average
length

Inches
3.9
9.1

13

18.5

22.4

Extremes of
length

Inches
3. 1 to 5. 9
7.1 to 12.6
8.3 to 18.9
11. 8 to 24. 4
16. 1 to 23

Age

6 years

7 years

8 years

9 years

10 years...

Average
length

Inches
25.6
27.6
29.1
33.9
37.4

Extremes of
length

Inches
20. 9 to 34. 3

21. 7 to 40. 9
22. 8 to 40. 6

26. 8 to 42. 1
29. 5 to 55. 6

Females averaged somewhat longer and heavier
than males of the same age, and the fact that the
oldest was a fish of 20 years, 68 % inches long, sug-
gests that the immense fish of 400 pounds and
more, and upward of 7 feet long, which are occa-
sionally caught, may be half a century old, always
assuming about the same rate of growth for the
Gulf of Maine halibut as for those that are caught
about Iceland.

According to Thompson 52 Pacific halibut grow
at approximately the same rate for the first few
years, more slowly after about the eighth year,
though with wide differences in the rate of growth
on different banks, probably caused by differences
in the food supply.

It is probable that most of the female halibut
do not mature sexually until they are 9 or 10 years
old, some not until they are several years older
still; males mature when they are somewhat
younger.63

General Range. — Boreal and subarctic Atlantic,
in continental waters.

The most southerly record of a halibut, in the
western side of the Atlantic is of a 6-foot fish that

»' Meddelelser fra Komm. Havunders0selser, Ser. Fiskerl, vol. 5, No. 5,
1917; based on a study of the otoliths of more than 2,000 fish caught around
Iceland.

« Report. Comm. Fish. British Columbia, (1914) 1915, pp. 76-99.

H Females of the Pacific form may mature as young as 8 years, or not until
as old as 16 years, with an average of 12; males considerably younger on the
average.

was picked up in a pound net near Reedville, Va.M
Stragglers have been reported off New Jersey and
New York, and off Block Island. And halibut
are caught in commercial quantities (or once were)
from Nantucket Shoals, inner parts of the Gulf of
Maine, Georges Bank, and the Nova Scotian
Banks northward to the northern part of the Gulf
of St. Lawrence (including the west coast of New-
foundland), the southern part of the Grand Bank,
Flemish Cap and to the outer edge of the con-
tinental shelf off outer Labrador at depths of
70-90 fathoms or more. But stray specimens,
only are reported in along the outer coast of Lab-
rador, i. e., in the icy Labrador current.66 And
while the range of the halibut was said by Goode 69
to extend to Cumberland Gulf, we doubt whether
there are any halibut in the icy waters along
Baffins Land, for the halibut is not known off the
Arctic coasts either of Asia or of America, though
the Greenland side of Davis Strait supports a
regular halibut fishery as far north as Disco Bay.

In the eastern Atlantic, halibut have been re-
ported doubtfully from the Gulf of Cadiz, and
definitely from the Bay of Biscay.57 Small catches
are made regularly as far south as the Irish Sea
and English Channel;68 and they are more numer-
ous around Northern Ireland and Scotland and in
the northern part of the North Sea; in the Faroe-
Shetland Region; around Iceland; along the Nor-
wegian Coast; around Spitzbergen and Bear Is-
land; also in Barent's Sea.

The Pacific halibut, an ally so close that it is
hardly to be distinguishable to the untrained eye,
is one of the most important food fishes of the
northeastern Pacific.

Occurrence in the Gvlj of Maine. — The history of
the halibut in the Gulf of Maine, like that of the
salmon, must be written largely in the past tense,
for their numbers have been sadly depleted there
by over-fishing. In Colonial days the halibut was
a familiar fish and seemingly a very abundant one
on the coast of northern New England, but was
considered hardly fit for food. Wood 59 for in-
stance, writes "the plenty of better fish makes

14 Reported by Walford, Copeia, 1946, No. 2, p. 100.

88 One, about 20>S 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.

88 Fish. Ind. U. S., Sect. 1, 1884, p. 190.

81 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.

■ 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
especiaUy 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.

' Goode 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 Gulf
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

« Ooode and Collins (Fish. Ind. U. S., Sect. 6, vol. 1, 1887, p. 3) have col-
lected data on the Oeorges 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 in 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 their 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, 1951.6*

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. M 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 Manan (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.86 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 County 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 avaUable 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 poimds; 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 Kcudder,
Fish. Ind. U. S., Sect. 5, vol. 1, 1887, pp. 90-119, for historical surveys of the
fresh and salt halibut fisheries from New England ports.

» Rcpt. TJ. S. Comm. Fish. (1929) 1930, pp. 85-86, 90.

« Reported in the Boston Globe, June 25, 1951.
" Reported in Salt Water Sportsman for May 29, 1951.
« 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 Skelburne 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 6-year period 1941-1946, plus what
fish may have been landed in Shelburne, Nova
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 1946.69

In the early days of the fishery, halibut were
common in 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

67 The otter trawlers that carried on investigations for 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 fisheryl

a The Gulf of Maine catch was nearly 3 million pounds in 1919.

•• Catch, Cape Sable to Cape Breton in 1946, 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,
1936, p. 21.

'• Goode and Collins, Fish. tad. U. S., Sec. 5, vol. I, 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
fleet 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 Gidf 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 winter 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

210041—53-

-18

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 71 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 readable 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 n 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.76

" Norman (Ann. Mag. Nat. Ilist., ser. 9, vol. 13, p. 539) reports a single
specimen taken southwest of Iceland.

" See Jensen (Meddel. Dansk. Komm. Havunders«lgelser, Ser. Fiskeri,
vol. 7, No. 7, 1927) for a general account of the Greenland halibut off west
Greenland.

H According to Goode (Fish. Ind. U. S., Sect. 1, 1884, p. 198) long liners have
reported it as "very abundant" in and off Fortune Bay, in 60 to 300 fathoms.

» Vladykov and Tremblay, Natural. Canad., vol. 62 (Ser. 3, vol. 6), 1935
p. 82.

It is described as "not uncommon" off Canso,
Nova Scotia,79 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

« Comish, 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, Ser. 3, vol. 17, 1879, p. 40.

" 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" of Europe.

Figure 128. — Canadian plaice, or Dab (Hippoglossoides plaltessoides) , 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 rounded 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. rays) 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
body; large ones are occasionally, though they are
plain colored as a rule.

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 32K 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, 1% 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 19K 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 (plales-
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 they 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 hi 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. H. 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,86 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.

" Ball. 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 plallessoides) .

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 mm.
in other seas, depending on the breadth of the
peri vitelline 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 89 when they are one year old. Thus it
may be assumed that bottom stages 2% to 3%
inches (69-80 mm.) long that we have trawled
off Cape Cod, on May 1, were about one year old;
others of 3% 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 18 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 8 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-

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
extend 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 (Frost M 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; 89 in the Gulf of St. Lawrence as a
whole, and thence westward and southward to
Cape Cod, from close inshore out to the 100-
fathom 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 15M inches long that was
caught off Montauk Point, N. Y.( in 112 fathoms,
February 6, 1930.90

» Flimtsman, Bull. Biol. Board Canada, No. 1, 1918.
° Bull. Biol. Board Canada, No. 1, 1918, p. 28.

91 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 Oat. 54°3T/ 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.

•• 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 Gulj 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 Mount
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
haid 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

•i For an account of it in west Greenland waters, see Jensen (Meddel.
Dansk Komm. Havundersdgelser, vol. 7, No. 7, 1925, p. 24).

w Huntsman (Bull. 1, Biol. Board Canada, 1918) gives an interestingaccount
of this fish in Canadian waters.

•' Bigelow and Schroeder, 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 very 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 M
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.

« Research Bull. 4, Newfoundland Dept. Nat. Resources, 1939, p. 8.
"Jensen, Meddel. Dansk. Komm. H avundersfjgclser, 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 was
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

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 Fundy, 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.

It 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.

■' It is only during the past few years that the landings of this particular
flatfish have been reported separately, as "dab."

•» Bull. 1, Biol. Boald 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. Tt 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 ; " 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 they 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, flgs. 11,
12, and 13.

' See Frost, Res. Bull. No. 4, Newfoundland Dept. Nat. Resources, 1938,
chart 2, for the regional and seasonal distribution of dab eggs in Newfoundland
and Labrador waters.

» Jensen, Meddel. Dansk. Komm. Havundersdgelser, 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 products. 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 dentatus
(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

» The landings for that year were 60,100 pounds for 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 dentatus), Maryland.

Baldwin.

From^Jordan and Evermann. Drawing by A. H.

268

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

orange, pink, and brown to almost black.4 Its
upper surface is variegated with 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. 5 The largest of winch
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 : 9

Average weight,

Length pounds

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 water
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

» Mast, Bull. U. S. Bur. Fish., vol. 34, 1916, p. 177.

■ Nichols and Breder, Zoologica. N. Y. Zool. Soc, vol. 9, 1927, p. 177.

• From Goode, Fish. Ind., U. S., Sect. 1, 1884, p. 179; Hildebrand and
Schroeder, Bull. V. S. Bur. Fish., vol. 43, Ft. 1, 1928, p. 167; and World
Record Marine Game Fishes, Amer. Mus. Nat. Hist., New York, to 1950.

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,s
fishing in the general offing of Marthas Vineyard,
brought in many fares ranging from a few hundred
pounds 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.

' 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, winter, 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. albiguttus
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, but the pig-
mentation on the rear part of their body is less
dense. At a length of 16 mm. the right eye has
nearly completed its migration, and the outlines
of young fry 26 mm. long approach those of the
adult.11

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 H, Capt. Henry Klimm, in 1951.

10 We dare not draw any conclusions as to spawning season from Hilde-
brand and Cable's table (Bull. U. 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 Hildebrand 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,12 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.,"
where we read of one of 1 1 Y% pounds taken as early
as the last week of May, in 1951, 15 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.18
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 Atwood 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 by Goode 17 in 1884 as "only

'• 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. lethostigmtts), a common Florid-
ian fish.

'• 645 pounds reported from the northwest part of Georges in 1947, 100
pounds from the northeastern edge, and 157 pounds from the central and
southeastern part.

" This opens on the outer coast of Cape Cod.

'• Reported in Salt Water Sportsman, June 1, 1951.

'• There is a record of this, by Kendall, in 1896, and we have known of other
cases, of late years.

'I Fish. 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 rare 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 tbat 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.

u Fish trawled by the Eugene Hot! Marthas Vineyard, Jan. 27 to Feb. 3,
1950, at 47 to 67 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 K 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 Gulj oj Maine.- — The four-
spotted flounder is so plentiful along the conti-
nental shelf as far eastward as the general offing
of Nantucket, and to the neighboring part of
Georges Bank that we counted about 1,800 of
them trawled there, by the Eugene H 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

18 Eggs artificially fertilized by O. E. Sette of the Bureau of Fisheries.

M Lefthandness foreshadowed in the larger ones by the fact that it Is the
right eye that had begun to migrate.

ai The most southerly record for It is for five specimens trawled in the
general offing of Charleston, S. C, Iat. 33°00' N., long. 77°44' W., at 92
fathoms, by Albatross III, January 30, 1950. Another flounder, Ancylopsetta
guadroeellata Olll, similar in appearance, for it Is strikingly marked with four
large spots, Is found along the South Atlantic and Quit 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 jerruginea (Storer) 1839

Rusty flounder

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. Scl., Ser. 3, vol. 17, 1879, p. 40) Goode and Bean
state that this specimen was trawled in Gloucester Harbor; in another paper
(Bull. Essex Inst., vol. 11, 1879, p. 7) tbey credit it to the mouth of Salem
Harbor.

272

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

'rrVSlrv

'J-WVS'

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 15K 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

» One was taken at 50 fathoms by Albatross II, September 5, 1926, on the
northwestern part of Oeorges Bank, and two of about 10 inches at 90-95
fathoms on the northern edge of Oeorges 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 hawl, 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-Boone
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 empt3T 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 winter." 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 him 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 in 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
winter flounders resemble one another closely; in
fact, it is probable that some of the young flat-
fishes pictured by A. Agassiz 25 as winter flounders
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 winter 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% to 4% inches long in 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 5 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 (Jeflers, 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.28 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 Gulf 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.

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 the 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).

" Localities are listed by Ooode and Bean (Smithsonian Contrib. Know!.,
vol. 30, 1895, p. 430).
*• To nearest 10,000 pounds.

» Bull. Essex Inst., vol. 11, 1879, p. 6.

"i 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," 3e 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;37
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.

n About 3VS million pounds were landed in New York in 1947.

u Albatross 11 trawled many yellowtails as far southward as the offing of
Delaware Bay Oat. 38° 32' N., long. 74" 24' W.) in 12 to 28 fathoms during
February, April, and June, of 1929 and 1930.

11 The Newfoundland Fishery Research Commission (Rept., vol. 1, No. 4,
1932, p. 1 10) reports 680 yellowtails taken per 10 hours trawlingon Banquercau.

•• Frost, Research Bull. 14, Newfoundland Dept. Nat. Resources, 1940,
p. IS.

" 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 1915.39 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 1 1 ; 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.

» Dannevig (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-fourtb
times as long to the base of the caudal fin as it is
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, Bingham Oceanogr. Coll., vol. II, Art. 2, 1947, pp.
19, 20) gives a detailed tabulation, and graph of the number of dorsal and
anal flu 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 H. 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., V. 3. Fish Comm., vol. 19, 1901, pp. 305-306.

« Copeia 1952, p. 206.

« Zoologica, New York Zool. Soc, vol. 9, 1927, p. 180.

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
dignabilis, by Kendall.46 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 ).49

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. 18-23) in this connection.

« Bull. U. S. Bur. Fish., vol. 30, 1912, p. 391, pi. 57.

<• Perlmutter has already emphasized this point in his detailed study of
the blackback (Bull. Bingham Oceanogr. Coll., vol. 11, Art. 2, 1947, p. 18).

•' Hildebrand 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. In 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)w 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 population 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 Oceanogr. 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." 6I 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,52 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 air tem-
perature rose to 82°-89°, and the temperature of
the water on the very shallow flats nearly as high,
probably.53 But we 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. Bingham Oceanogr. Col]., vol. 11, 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. 56, 1918, pp. 37-39),
also by Nichols and Breder, Zoologlca, 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,64 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 4K 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
1 1-inch flounder at St. Andrews, New Brunswick.53
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. 60, 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 1952. p. 3.

280

FISHERY BULLETIN OF THE FISH AND 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-

*• 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
swimming 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 mm. 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 mm. 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 mm.,
and the left eye has moved upward by then until
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.

•i 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. 38, 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 (Pseudopleuronectes 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}i 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 aeries 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 \% 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.6*

Welsh also concluded, from measurements
gathered from various sources, that the winter
flounders are 5 to 7K inches in length at 2 years of
age, iy% 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
fish 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.

'i Three larvae taken in the Quit in July 1912, were provisionally identified
by Welsh as this species.

210941—53 19

« Hildebrand and Schroeder, Bull. U. S. Bur. Fish., vol. 43, Pt. 1, 1928,
p. 169.
« 28th Rept., New York Conserv. Dept. 1939, Sup., Pt. I, 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

" Jeflers (Contrlb. Canadian Biol., N. ser., vol. 7, No. 16, ser. 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. Mas., vol. 6, 1883, p. 125.

" At 2 stations, see Kept. Newfoundland Fish. Res. Lab., vol. 2, No. 3,
1938, p. 79.

10 Reported from Fort Chlmo, Labrador by Kendall (Proc. 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.

" Reported from Beaufort, N. C. (by Yarrow, Proc. Acad. Nat. Sci., Phila-
delphia, vol. 29, 1877, p. 205); from the Neuse River, near New Bem, N. C.
(by Smith, North Carolina Oeol. and Econ. Surv., vol. 2, 1907, p. 390); and
from Georgia (by Hlldebrand and Schroeder, Bull. V. 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 23,00f>
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 black back.

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 u 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

» 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 Gulf of Maine coasts of the Cape, and how
much off the south shore of Massachusetts.

" 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 fisted under the
heading "spears" in the Massachusetts landings
by gear for 1945 or for 1946."

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 winter 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

Figure 145. — Smooth flounder (Liopsetta pulnami) , 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 young 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; "
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 macy 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. ADthonys, northern Newfound-
land, In the Museum of Comparative Zoology.

" Kept. 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; M 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, generally.

In Casco Bay and in estuaries of the Bay of
Fundy such as the mouths of the St. Croix 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.

M This specimen, formerly in the Museum of Comparative Zoology, is no
longer to be found.

■i Our experience corroborates this to the extent that we have never seen
it 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 Glyplocephalus 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 yellowtail.
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.

snout 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 shoaler 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 1950. but an average

of only about one fish per haul between 31 fathoms
and 80 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,8*
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

M Ooode and Bean (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. Explor. Mer, 1925, p. 3.

» Melander, Pub. de Circonstance, Cons. Internat. Eiplor. 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 tempera ture 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

i 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 Qulf 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.

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 2 % to 4% inches, and of 3 % 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 1936 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 oj 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.

•' See Frost, Res. Bull. No. 4, Newfoundland Dept. Nat. Resources, 1938,
Chart 6, for Newfoundland localities.

*> Ooode and Bean (Smithsonian Contrib. Know]., 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 Guff 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:9* Browns Bank, 44,000
pounds; off western Nova Scotia, 2,000 pounds;
off eastern Maine, 17,000 pounds; off central

w 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, Glyptocepkatus acadianut.

<* 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.85

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 Gulf 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.96 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
[sland.97 And they seem to be moderately plenti-

" An additional 182,000 pounds were landed in Cape Cod fishing porta,
source not stated.

" About 19,000 pounds in New York in 1947, about 28,000 pounds in New
Jersey.

" Only a few 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 M 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 their 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." 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
Julv 8, 1913
July 9, 1913
July 19, 1916....
July 22, 1912....
July 24, 1912....

Aug. 6, 1913

Aug. 9, 1913

Aug. 14, 1912...

109

19

1

100+

1

2

27

7

1

8 to 23.5.

8.5 to 21.5.

14.

5 to 19.

9.5.

8.5 and 16.6

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.

29.6 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 yellowtail, 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 five 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 white 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 )i 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 Oceanogr. Coll., vol. 11, art. 3, 1947, p. 20.
> Bull. Bingham Oceanogr. Coll., vol. 11, art. 3, 1947, p. 63, flg. 12.

Figure 151. — Sand flounder (Lophopsetta maculata). 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 in 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,6 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.*
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 2K-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. 54-5S.

• 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 yellowtail (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 8K 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 1 to 2 months after spawning
commences there. One that was kept in an
aquarium there by Williams u grew from 10 mm.
to 22 mm. in length in 11 days; and Nichols and
Breder's 12 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% inches by late September, is in line
wi thTracy 's statement 13 that the fry are 2 to 3 inches
long in July in Rhode Island waters, growing to

• Zoologies, New York Zool. Soc., vol. 9, 1927, pp. 181-182.

• Hlldebrand and Schroeder, Bull. U. S. Bur. Fish., vol. 43, Ft. 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 flounder.
For photographs of larvae and small fry, see Moore (Bull. Bingham Oceanogr.
Coll., vol. 11, art. 3, 1947, fig. 3).

" Bull. Mus. Comp. Zool., vol. 40, 1902, p. 3.

u Zoologies, New York Zool. Soc, vol. 9, 1927, pp. 181-182.

>» Kept. 40, Comm. Inland Fish. Rhode Island, 1910, p. 166.

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
4K inches long when they are 2 years old (i. e., at
the beginning of their third summer); about 7%
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 Provincetown, 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, 1947, pp. 47-61.

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.18
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, Contrib. Canadian BioL (1921). No. 2.'l922, p. 70.

part. And it is reported as common at Beaufort,
N. C.ie

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 ; n a few
adults have been taken in Chedabucto Bay,
eastern Nova Scotia; 18 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 x 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.31

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 arctijrons
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 I, No. 4, Newfoundland Fishery Res. Comm., 1932, p. 110.

» Comlsh, Contrib. Canadian Biol., (1902-1905) H07, p. 60.

'• Contrib. Canadian Biol.. (1918-1920) 1921, p. 113.

» Trans. Roy. Soc. Canada, Ser. 3, vol. 12, Sect. 4, 1918, p. B3.

a Rept., Newfoundland Fish. Res. Comm., vol. 2, No. 1, 1933, p. 127.

» See Moore (Bull. Bingham Oceanoirr. 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 Wildlife Service.

FISHES OF THE GULF OF MAINE

295

JP

m

mm

tfwM

WMMMzWh

HpMP

p

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 83 rays) originating over the forward
margin of the eye, the anal (left-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 Ihls genus (C. unicornis Goode 18801 may be expected
on the outer slope of Georges Bank in depths of 100 fathoms and more, since
it has been taken off 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 the 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 Ciiharichlhys of the western Atlantic.

We have towed the pelagic larvae of still a third small deep-water flounder
(Monolenc sessiliiauda Goode 1880) ofl 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 fln 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 itsleft-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
years 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 20046, lat. 40° 18' N., long. 68° 09' W., February 22, 1920.

Hogchoker Achirus fasciatus Lac6pede, 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.

H. L. Todd.

Original drawing by

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 whole 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.

THE JOHN DORIES.

ConcniHra. (Lout)
American John Dory M Zenopsis Mellette, (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

» HUdebrand and Schroeder, Bull. U. S. Bur. Fish., vol. 43, pt. 1, 1928,
p. 176.

« Hildebrand and Schroeder, Bull. D. S. Bur. Fish., vol. 43, Pt. 1, 1928,
p. 177.

n 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.

It reaches a length of 2 to 3 inches at one year of
age, and matures when about A)i inches long.
It feeds chiefly on annelid worms and on small
crustaceans.28 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." 27

FAMILY ZEIDAE

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 rayed ; 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 oceUata), 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 18K inches; M 19 inches, weigh-

a This agrees with (he original account of the species (Storer, Proc. Boston
Soc. Nat. Hist., vol. 6, 1858, p. 380) and with a photograph of one about 3H
Inches long, from Campobello, New Brunswick, sent us by Dr. A. H. Lelm.

» Taken ofl Long Island, N. Y., by the Albatross III, lat. 39°39' N., long.
72°08' W., May 12, 1960.

FISHES OF THE GULF OF MAINE

299

ing 3 pounds;31 and 20 inches, weighing 4%
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% 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.3*

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 separates 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 Gulj oj 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 38 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
66 to 75 fathoms; on Emerald Bank off Halifax,
Nova Scotia, in 70 fathoms; and west of Sable
Island, Nova Scotia, at 62 fathoms.

GRAMMICOLEPID FISHES. FAMILY GRAMMIGOLEPIDAE

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 H, May 18,
1960.

J> Double and sometimes triple In the European 7. faber.

" Trawled by the dragger Eugene H from between 65-68 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

M Reported to us, with a photograph, by Dr. A. H. 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 seen by us or reliably reported.

» Reported by Firth, Copeia, 1931, p. 162.

• For a recent account of this family see Myers, Proc, V. 8. National
Museum, vol. 84, 1937, pp. 145-166.

300

FISHERY BULLETIN OF THE FISH AND "WILDLIFE SERVICE

ffiiS

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
along both its front margin and its rear margin for

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 Chilj 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.

« 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 sidcwise 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}i 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; tt 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 size is pictured by Murray and HJort (Depths of the Ocean,
1912, p. 397, fig. 268).

"Goode and Bean (Smithsonian Contrib. Knowl., vol. 30, 1895, p. 483),
without further details.

" Original of Ooode and Bean's illustration (Smithsonian Contrib. Knowl.,
vol. 31, 1895, pi. 127, fig. 396).

Figure 158. — Snipefish (Macrorhamphosus scolopax), 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.46

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 naiTow 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
5% inches, adults usually running 4 to 4% 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

*' 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 in tergradations 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, IT. S. Comm. Fish (1901)
1902, p. 241). For a recent discussion see Bayllffe (Publ. 90, Chesapeake
Biol. Lab., Maryland Dept. Nat. Res.. 1950, p. 5).

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 49 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 the 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 4? 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 silversides 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.

"Kuntzand Radcltffe (Bull. V. S. Bur. Fish., vol. 35, 1918, p. 127) de-
scribe Its development, and Hildebrand (Bull. U. 8. Bur. Fish., vol. 38[ 1921-
22] 1923 that of the southern race. For a recent account of Its life history see
Bayliffe, 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

«Qoode. Fish. Ind. U. S. Sect. 1, 1881, p. 467.

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 Gulj oj 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

« Contrib. Canadian Biol. (1921) 1922, p. 61.

outer coast of Nova Scotia.61 But Leim 62 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 Giinther).

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,

•' 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 fine; 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 M (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 1939,65 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 sdversides,
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 fining 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. 8, 1908, p. 66, as Menidia
beryllina subspecies area Kendall (Rept. U. S. Comzn. Kish. (1901) 1902, p.
261.)

» This specimen is in the Museum of Comparative Zoology.

been known to stray within the confines of the
Gulf of Maine."

Mullet Mugil cephalus 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 (MuUut auratug) 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
Halifax Harbor, Nova Scotia (by Leim, Proc. Nova Scotian Inst. Sci., vol.
17, No. 4, 1930, p. XLVI), hence it may be expected as a stray In our Qulf,
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 fins 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 commoo 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 gray 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)i 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

" See Jaoot (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

u There is (or was) a specimen so labeled in the collection of the Boston
Society of Natural History.
■ 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.

30S

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

•i MacCoy, Bull. 74, Boston Soc. Nat. Hist. 1935, p. 16.
" Huntsman, Contr. Canadian Biol., (1921) 1922, p. 61.

Figure 167. — Three-spined stickleback (Gasterosteus aculeatus). 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. They 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. In 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 fist 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

M Large numbers are sometimes seined In 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 hatcliing-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

87 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 (Proc.
Amer. Acad. Arts Sci., vol. 17, 1882, p. 288, plate 9), and by Ehrenbaum (Nor-
disches Plankton, vol. 1, 1905-1909, p. 319).

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-
culus 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

u This is the Gasterosteus biaculealus of Mitchill 1815 and Storer 1867;
bispinosus of Walbaum 1792; gladiuncutus of Kendall 1896, but not the Q.
bispinosus of Jordan and Evermann 1896, which is a variety of O. 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 Gvlj oj 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.

■ Proo. U. 8. Nat. Mus., vol. 18, 1896, 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, while 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 the three-

Figube 171. — Four-spined stickleback (Apeltes quadracus), Woods Hole.

A. H. Baldwin.

From Jordan and Evermann. Drawing by

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."

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 74 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.

n Bull. U. S. Fish. Comm., vol. I, 1882, p. 24.

71 The early development is described by Ryder (Bull. U. S. Fish Comm.,
vol. I, 1882, p. 24) and by Kuntz and Radcliffe (Bull. U. S. Bur. Fish., vol.
35, 1918, p. 132).

« Contrib. 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 Storer 75
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 (Kept. IT. S. Comm. Fish. [1928], 1930, Pt. 2,
p. 242) refer it to the genus Sj/rictes Jordan and Evermann, 1927. For a syn-
opsis of the various other species of the genus see Jordan and Evermann,
Bull. 47, D. S. Nat. Mus., Pt. 1, 1896. p. 961.

FISHES OF THE GULF OF MAINE

313

Figure 172. — Pipefish (Syngnathus fuscus) . After Bigelow and Welsh.

ninth of the total length (in the trumpetfish it is
nearly one-third) ; the snout 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 back
to the vent, with the marsupial flaps flesh-colored.
The dorsal and pectoral fins are pale, but the caudal
is brown.76 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— 5S 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. 8. 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 then-
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 pouoh 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 Siphostoma floridae see Gudger (Proc. U. S. Nat. Mus., vol. 29, 1906,
pp. 447-600, pis. 6-11).

10 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
fflr die gesamte wissenschaftliche Anatomie, vol. 24, 1904, pp. 192-199).

•> Tracy, 40th Rept., 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 oj 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),
1896-1900, 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 (S. juscus) 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 Gulj oj Maine. — A single
specimen, 3K inches (89 mm.) long, taken on
Georges Bank (Lat. 42° 09' N.; Long. 66° 41' W.),
September 20, 1927, by the Albratross II is 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.

>' Lcim (Proc. Nova Scotian Inst. Sei., 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 Compara-
live Zoology.

FISHES OF THE GULF OF MAINE

315

THE„SEA HORSES. FAMILY HIPPOCAMPIDAE

'-!!■ credos FcrrV
Sea horse Hippocampus hudsonius DeKay 1842

Jordan and Evermann, 1896-1900, p. 777.

Figure 173. — Sea-horse (Hippocampus hudsonius), 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,86 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 time 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.

»> Gill (Proc. TJ. 8. 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 Comra., 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 Gulj of Maine. — The sea horse
is not common much beyond New York. Only a
few are found 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). La 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 tbat 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 H. bretirostris Storer
(1839); later by Jones, (Proc. Nova Scotian Inst. Scl, vol. 5, Part 1, 1882.
p. 95) as H. antiquorum Leach 1814.

'• Proc. Nova Scotian Inst. Scl., 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 finlets
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 98

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 5

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. S) as Fiitularia aerraia.

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. Essex Inst., vol. XI, 1879, p. 4.

B» This specimen is in the Museum of Comparative Zoology.

« 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 Scombridae rather than Jordan, Evermann
and Clark (Kept. U. S. Comm. Fish. (1928) Pt. 2, 1930) who have distributed
them among four families, Cybiidae, Katsuwonidae, Scombridae, and
Thunnidae.

•3 The long finned Albacore (Thunnus alalunga Bonnaterre, 1788) has been
taken at Woods Hole, also on Banquereau Bank, off eastern Nova Scotia
(Qoode and Bean, Bull. Essex Inst., vol. XI, 1879, p. 15), so is likely to show up
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 9l 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% 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

" This interesting fact seems 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 Oeeanographic Institution.

» Fish. Bull. U. S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, p. 264.

" Bull. TJ. 8. 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 by the luminescence of the tiny organisms
that they disturb in their progress. Sette "
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. 8. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, p. 267.
» For observations on the visual transparency of the water of our Gulf, 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 winter in water as cold as 43°— 15°
(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 in August 1896, were packed with crab
larvae, others were full of Sagittae, others, again,
of Sagittae and amphipods (Euthemisto) , 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. 51, 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.

1 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 Gidf of Maine is to the effect that
this would be exceptional there, if it happens at all.

• Nilsson (Publ. de Circ, Conseil Perm. Internat. Explor. Mer, No. 69.
1914) gives a similar list for Swedish waters

210941— 53 -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 6 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 niters but snatching
gulps from different directions and makiDg 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, W2S, fig. 42 C, D for photographs of the
gill rakers. I Til

• Nature, vol. 108, 1921, pp. 12-13.

1 Sette (Fish. Bull. U. S. Fish and Wildlife Service, vol. 51, Bui'. 49, 1950,
pp. 259, 262) 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 off 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.' 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, had 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. Comm.
Fish. (1898) 1899, p. 5; Bigelow and Welsh, Bull. U. S. Bur. Fish., vol. 40,
Pt. 1, 1925, p. 208.

i" 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 I3 (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 oil 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 after 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 IK to 2%
inches obviously spawned that spring, have been
taken at Woods Hole, both in the first half of
June a and in the last 10 days of July,18 fry of
2% to 5 inches in the first half of August, and fish
of about 6% inches at the end of that month.
Similarly, Captain Atwood found fry of 2 inches
and shorter in July in the Massachusetts Bay

» Jour. Gen. Pliysio]., vol. 16, 1933, pp. 841-857.

'* They shrink somewhat when preserved.

'» See Sette (Fish. Bull. U. S. Fish and Wildlife Service, vol. 60, Bull. 38,
1943, pp. 173-178) for detailed statistical analysis of these.

" See Ehrenbaum (Rapp. et Proces Verb., Conseil Perm. Internet. 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 their 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.
549-568.

" Fry spawned in 1927 averaged about 1331 inches but those spawned in
1928 averaged only about 12H inches in their second November according
to Sette.

324

FISHERY BULLETIN OF THE FISH AND "WILDLIFE SERVICE

summer, although it is long lived. The two sexes
grow about equally fast.

Nilsson's studies 21 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.

» Smith, Report U. S. Comm. Fish (1900) 1901, p. 128.

»« Nllsson, 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.

» Jeffers (Contr. Canad. Biol., N. Set'., vol. 7, No. 16 [ser. A, General No.
13], p. 207) reports that several mackerel were caught in 1929 at Raleigh, on
the Newfoundland coast of the Strait of Belle Isle, where none had been seen
In recent years.

1 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 then 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 Gordon32 has

'» The literature dealing with this subject is very extensive. See especially
Goode, Collins, Earll, and Clark (Rept. U. S. Comm. Fish. [1881] 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. 50, Bull. 49, 1950,
pp. 260-261, table 1) lists several such Instances besides those cited previously
by Blgelow and Welsh (Bull. U. S. Bur. Fish. vol. 40, Pt. 1, 1925, p. 196).

•i Three hundred pounds seems to be the largest winter catch definitely
reported up to 1951.

•i 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, Rept. V. 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°^6° 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
tilefish 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. Intemat. 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.

41 Mackerel do not range that far north.

« Cited from Lacepede, Hist. Nat. Poissons, vol. 3, in Buflon, Hist. Nat-
urelle, 1802, p. 32.

11 Ehrenbaum (Rapp. et Proces Verbaux, Cons. Perm. Internat. Eiplor.
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 46 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 49 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

•' 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 taken 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 oft North Carolina (Copela, No. 92, 1921, pp.
18-19), but Dr. Nichols writes us that these were "just Scombroids and
probably not Scomber scombrua."

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
in 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 Bigelow and Welsh (Bull. U. S. Bur. Fish., vol. 40, Pt. 1, 1925, p.
206), for details.
« Dannevlg, 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.86

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.

•» Subfflquent 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,
Pt. 1, 1925, p. 206) believed.

» Sparks, Contrib. Canadian Biol, and Fish., N. Scr., 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. 61, 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 27 th of July, on which date 28 seiners made
catches ranging from 18 to 250 barrels each. And
in 1892, 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°-^7° 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 Senrioe, vol. 51, Bull. 49, 1950. p.
297, flg. 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 Maine 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 1906.58 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

- Sette 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.

71" 70*

69» ea*

67-

SELOOM V//A

ll*

OCCASIONALLY 1 1

USUALLY ES3

^TJF'^

y.

ALMOST ALWAYS MB .

J<."

-'' .'

44-

MM

**'\

4 4*

*i

4J«

*i

^39&Wra£S^

-

42'

*(

0
**>

41*

40

- ..-'"'•-■<

*

x _ i

1 1

I

40*

71* 70-

69* 66'

67*

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 while 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

■ 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 ha ye 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

" 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 off 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; aud 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 (1837^45), 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 16K 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
of the purse seine. Actually, these changes re-
flect the ups and downs in the numbers of the fish
that are in existence from year to year. Mack-
erel, in short, were extremely plentiful in 1885,
very scarce in 1910, moderately plentiful in 1916
and 1917, 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.65

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 No. 4, 1931.

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 11 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 in 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,

• Sec especially Sette, U. S. Bur. Fish., Fishery Circular No. 4, 1931.
« Qulf of Maine catch, 16,391,095 pounds in 1916; 16,021 ,619 pounds In 1917.

« Oulf of Maine catch, 11,007,676 pounds In 1923; 25,475,876 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 Oazette, vol. 60, No. 5, 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
through 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

$7,000,000 (1.4)
4,280,000 (2.3)
3,840,000 (1.3)
3,160,000 (1.3)

$8,800,000 (1.2)
3,940,000 (1.4)
4,750,000 (1.7)
2,340,000 (1.1)

$8,900,000 (1.3)

Cod

2,780,000 (1.4)

4,200,000 (1 4)

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 line 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 7l

Hardhead; Bullsete

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.

'i 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

Figure 177. — Chub mackerel (Pneumatophorus colias), Provincetown, Mass. From Goode. 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 could catch 3,000 in a day on hook and

71 It Is reported from St. Margaret Bay and Halifax by Vladykov (Proc
Nova Scotlan Inst. Set., vol. 19, 1936, p. 7), and Sehmltt (Monographic de
l'lsle de Antlcostl, 1904, p. 285, Paris) credits It with "apparitions lrregu-
Ueres" at Antlcostl.

» 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 1880 there were 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 74
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. 60, New York State Mus., Zool. 9, 1903, p. 383.
« Field notes 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,77
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), in 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 finlet.

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 (Kafinesque)
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
fine 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 Euthynnus. see Fraser
Brunner, Ann. Mag. Nat. Hist., Ser. 12, vol. 3, 1950, p. 150.

Figtjbe 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% 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 Gulf of 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 11, 1949.82 Like various other
tropical fishes they come our way only as strays
from wanner 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

'i 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

*> No one should take a bonito for a large mackerel, its dorsal fins 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,89 on the American coast and to Scandinavia
on the European coast.

Occurrence in the Qvlj 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 the
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 88 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

•» Nichols 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. Selm).

" Reported by Walter H. Rich.

M 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
Corwln, 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
cheeks 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 finlets 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.81

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 excellent 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, 1960).

•' 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 96 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 babitat 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

1931 1

Sep-

•• Internat. Rev. Gesamten Hydroblol. Hydrogr., vol. 25, Pt. 1-2,
p. 60.

• Caught by Comm. D. W. Hodson at Cape Breton, Nova Scotia,
tember 4, 1950.

« Reported In Salt Water Sportsman, for Oct. 1, 1951.

" Caught near Jordan Ferry, Nova Scotia, 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 Prank Mather of
the Woods Hole Oceanographic Institution and Howard Schuck of the V. S.
Fish and Wildlife Service, who have given us much first-hand information
on the habits of the tuna.

" Heldt, 10 Rapp. Comm. Internat. Eiplor. 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 tuna 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.

' See Farrington (Fishing the Atlantic, 1950 [approximate date], upper
photo facing p. 421), for an excellent photograph of a tuna finning.

' See Farrington (Fishing the Atlantic, 1950 [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

» Internat. Game Fish Assoc. Yearbook, 1945, p. 65.

* The tuna that visit the west coast of Newfoundland find summer temper-
tures as high as 59°-60° along the south coast of Newfoundland, and 550-57<'
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.6 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 Oceanographique de Salambo, 1926), and
Sella (Int. Rev. Hydrobiol., Hydrogr., vol. 24, 1930, p. 446) for 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.

7 Frank Mather of the Woods Hole Oceanographic Institution informs us
that a 600-pound tuna has been taken in a trap 200 miles south of Chincc-
teague, Md., and that small ones are taken off Chincoteague. "Tuna" are
reported from time to time off North Carolina, also. But it is not yet certain
whether these actually are "bluefins."

« 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 vertebrae11 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 Reale 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; Heldt (Bulls. 5 and 18, Station Oceano-
graphique Salambo, 1926 and 1930) for summaries of all previous observations
on the breeding habits and larval stages.

» Monogr. 6, Acad. Nat. Sci. Philadelphia, 1944, pp. 261, 373.

» Memoria No. 166, R. Comitato Thalassograflco 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; u north regularly to the western, southern
and southeast coasts of Newfoundland,15 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 the two sides of the Atlantic, and those of different oceans appear, at
most, to represent races of a single wide-ranging species.

" Vcsey- 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: Pounds

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- Essex 50,300

tember off Massachusetts; July and August off Y' .

Casco Bay; through August and September along Cumberlandl

western Nova Scotia. Sagadahoc J 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- £nox (Penobscot Bay) 0

•", ^ , „, , Hancock . 0

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 ; Shelburne to Cape Sable 0

and the vicinity of Wedgeport, on the west coast In most years 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 Dut few larger, if any, are being made again off

in the local weirs causes comment, for the astound- tne 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 oi 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. 1B 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 Gulf 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 the Boston Herald, July 31, 1950. » Reported by Frank Mather of the Woods Hole Oceanographic Institu-

ir Information supplied by Frank Mather, of the Woods Hole Oceano- tion.

graphic Institution. « Frank Mather, of the Woods Hole Oceanographic 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 16, 1951.

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

610

1929

630

1930

620

1931

565

1932

500

1933

495

1934

530

1935

525

Thirty-two fish caught at the mouth of Casco
Bay in 1950 averaged 468 pounds, the heaviest 643
pounds; M 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

ii 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. Soe., vol. 21, No. 16, 1936, p. 207.
u 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 River, the
mouth of the La Have River, 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

* International Oame Fish Assoc, Yearbook, 1943, p. 65.
» Data from Walter 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, w 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, their 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 Latest catch

1943 July 8 Oct. 6

1944 June 29

1945 June 25 Oct. 9

1946 June 15 Oct. 26

1947 June 21 Oct. 28

1948 June 11 Oct. 28

1949 June 7 Nov. 14

1951 July 7 Oct. 15

1952 June 24 Oct. 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.
« In 1950 seven tuna of 200-300 pounds were caught off Boars Head, Maine,
during the first week of October.
81 Reported In Cape Cod Standard Times, October 11, 1950.
•' Information contributed by the Pond Village Cold Storage Co.
210941— 63 23

months. The number of pounds of tuna (dressed
weight) follows:

Month Largest catch Smallest catch

June 17,520 0

July 148,139 12,255

August 185,305 5,029

September 70,125 «• 2, 365

October 43,603 330

November34 2,197 0

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 1928.36 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

u The year 1944 Is omitted from the calculation for September-November,
because the traps were not fished after September 14th that year.
u The only catch recorded for November was 2,197 pounds in 1949.
*» Sella, Internat. Rev. Oesaraten Hydrobiol., Hydrogr., vol. 25, 1931, p. 62.

346

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

abundance, though fishermen are well aware that
their numbers in any part of the Gulf do vary
widely from year to year. Thus it is on record
that they were scarce in the Massachusetts Bay
region for two or three years prior to 1904, but
were plentiful that summer. Commercial land-
ings suggest that they were scarce again in 1943,
when the landings came to only about 380,000
pounds for Maine and Massachusetts combined.39
But they appear to have been much more
plentiful again off the Maine coast in 1945 (catch
there about 850,000 lb.) ; more plentiful than they
have been since, if the commercial landings are a
reliable index to the ups and downs of the tuna
population, which they may not be. The follow-
ing catch statistics of tuna landings (in pounds)
suggest that the stock built up more slowly, from
the 1943 low in Massachusetts waters, to a peak
in 1948, which was a big year on the Ipswich Bay
grounds (p. 343), as well as in Cape Cod Bay.

Year Maine Massachusetts

1943. 25,000 129,500

1944 463, 500 272, 900

1945 859, 500 356, 400

1946 421, 800 571, 100

1947 186, 600 726, 400

1948 229, 100 1, 627, 000

In 1949, the catch by traps in Cape Cod Bay
alone was 811,160 pounds, suggesting a total of
more than a million pounds from the Gulf of
Maine coast of Massachusetts.

During that banner season 2,164 large tuna
were taken on hand lines where the draggers work,
off Ipswich Bay; while 806 fish (305,300 lb.) were
taken off Wedgeport, Nova Scotia, in 1948, 1,760
fish (449,362 lb.) in 1949.

But they were scarce in 1950, to judge from
reports coming in from all along our coast: a
week's fishing, for instance, by the same number of
rods and at about the same date, yielded only
about half as many on the famous Soldiers Reef
off Wedgeport during the International Match
that summer as it had the year before. Many
fewer were caught by anglers in Cape Cod Bay
in 1950 than in 1949, and the tuna disappeared
from the Ipswich Bay region unusually early that
year, as noted above (p. 344). It is too early
(August 5) to forecast how the 1951 season may
develop.

The largest Gulf of Maine catches of which we
have heard were of 336 fish, weighing about 75,000
pounds taken at one lift of 3 traps set for mackerel
on the Barnstable shore of Cape Cod Bay, Aug. 5,
1948; 37 and of 120,000 pounds of fish ranging
from 25 to 30 pounds seined some 50 miles east of
Cape Cod on September 18, 1951, by the Western
Explorer, chartered by the U. S. Fish and Wildlife
Service for experimental fishing for tuna.

Importance. — Horse mackerel were formerly
regarded as a nuisance on the Atlantic coast, for
bands of them made trouble for fishermen by
following herring or mackerel into the traps and
pounds, to tear their way out again through the
net unless harpooned. Many years ago, when fish
oil was more valuable than now, a few were some-
times harpooned for oil, which was tried out of
the heads and bellies, but there was no sale for
their meat. The tuna, however, has been highly
valued as a food fish for many years, not only in
the Mediterranean, but on the west coast of the
United States. And a local demand has developed
on our coast, supplied chiefly by local fisheries
off Casco Bay, in the Cape Ann-Boone Island
region, and in the Cape Cod Bay region.

With this increasing demand, the reported
landings on the Maine and Massachusetts coasts
have risen from about 94,000 pounds in 1919, to
around 250,000 pounds yearly in the early
1930's, and to about 1 to nearly 2 million pounds
for the years 1945 to 1948, this last representing
around 3,000-6,000 fish, if they averaged 300^00
pounds in weight (see table, p. 346). The average
value to the fisherman in 1946 was about 7-9
cents per pound and all that are caught now sell
readily. The annual catches off the entire coast
of Nova Scotia ranged from 152,000 pounds to
about 1,550,000 pounds during the period 1917 to
1928; from 402,000 to 1,820,000 pounds for the
5 years 1942^6.

The commercial catch off the coasts of Maine is
made mostly by harpoon; that off northern
Massachusetts by hook and line and by harpoon;
that off the Cape Cod Bay region mostly in the
traps.38

» The weights given in the Fisheries statistics are for the dressed fish, and
represent about 80 percent of the live weight.

" A photograph of part of the catch was published in the Boston Herald,
August 6, 1948.

» 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 har-
pooned. About 86 percent of the Massachusetts catch was taken in traps of
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 oft 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 ca valla, is that the
pectoral fins of the Spanish are 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.

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 Unlets. 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 oj 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 Geol.
Econ. Surv., vol. 2, 1907, p. 191) for a specimen observed in a Washington,
D. C, fish market. II the identity was correct and this was not the closely
related king mackerel ScombcTomorus 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, 1895-1900, 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
Indies 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.

Figure 183. — King mackerel (Scomberomorus regalis), Key West, Florida. From Goode. Drawing by H. L. Todd

FISHES OF THE GULF OF MAINE

349

Cavalla Scomberomorus cavalla (Cuvier) 1829 *2

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 wliich 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 king 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 oj 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."

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 Ruvettus pretiosus Cocco 1829

Oilfish; Scourfish; Plaintail
Jordan and Evermann, 1896-1900, 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

o Fowler (Proc. Acad. Nat. Sci. Philadelphia, vol. 56, 1905, p. 766) refers
this species to a now subgenus Sierra.

glance from all Gulf of Maine mackerels by the
facts that it has only 2 dorsal finlets 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.

« 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

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 48 and in
East Indian waters. It is plentiful around Cuba
though not reported from Puerto Rico; 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 fife, dark edged
or speckled along the margin with black; the
tips of the jaws diisky.

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. Know]., vol. 30, 1895, p. 197.

" LaMonte, Marine Life, vol. 1, No. 8, 1960, p. 40.

FISHES OF THE GULF OF MAINE

351

Figure 186. — Cutlassfish (Trichiurus lepturus), 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 49 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, 1908, p. 76) are based on the
Wellfleet specimen. He also credits it to Monhegan I., Maine, quoting Storer
as his authority, but Storerstated in his latest mention of the species that only
two had come to his notice; 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.50

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,82 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.63

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 13%
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. H — 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 tbe temperature is
higher than about 60°. But occasional captures

" In the sailfishes 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, oft* Shel-
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,
argen tines, 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,56 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 attackmg
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 of Fisheries reports the
following genera as taken from swordfish on Georges and Browns Banks;
A'episaurus, ChauJiodus, Chiasmodon, Lampadena, Macrostoma, Myctophumt
Notoscopelus, and Stomiaa.

" Kingsley (Science, N. Ser., vol. 56, 1922, pp. 225-226) reports two freshly
swallowed stomiatids (Echiostomi barbatum) being taken from the stomach of
a swordfish harpooned on the offshore slope of Georges Bank.

» Kept. U. S. Comm. Fish., (1880) 1883, pp. 353-354.

354

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

in "fits of temporary insanity," as Goode *•
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.80 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. U. 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 tho rather extensive literature
dealing with the swordfish.

« Rich (Proc. Portland Soc. Nat. Hist., vol. 4, Pt. 2, 1947, p. 71) so Informs

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 swordfisherman 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

M See Farrington (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 oj 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," 66 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 CaDso 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; 6S 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 70 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. 29S-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.

«'Tho weights given are dressed; live-weights would be about 1M time3
as great.

« Forty pounds reported, but this probably is an error, for it is not likely
that a swordfish that small was taken there.

«» Victoria, Cape Breton and Richmond Counties, Nova Scotia.

™ Internat. Game Fish Assoc. Yearbook, 1943, p. 66.

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, off 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.75

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 fine,
on Browns Bank in 1931; a third of 7% pounds,
taken on a long line by the Dacia on Western Bank,
early in September 1931; and a fourth 28% 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
beard. 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
their 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

'« This Is often spoken of as the "Gulf Stream"; Its more accurate name Is
the "slope water."

" Rich, Proc. Portland Soc. Nat. Hist., vol. 4, Part 2, 194", 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 Wiidllfe Service.
The specimen is In Its collection In Woods Hole.

" Fish. Ind. 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.

w Many have been opened with this point in mind; some by us,

» Townsend, Science, N. Ser., vol. 66, 1922, pp. 18-19.

" Spolla Atlantica, in Kong. Danske Vidensk. Selsk. Skrlft, Ser. 6, 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.85 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 would 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.88 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

83 Landings were only about 545,000 pounds in Massachusetts and 7,000
pounds tn 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.

** The Canadian catch statistics for 1947 have not reached us yet.

'• 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 warmer
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 (Proc. 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.'2

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.

•' Hemmlngway (in Vesey-Fltzgerald and Lamonte, Game Fishes of the
World, 1949, p. 158) reports theso striped marlins weighing up to 1,260 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% 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

M Very young marlins have only one continuous dorsal fin, but this separates
later into two.

« 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 martin (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.86

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." 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,

•• 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 flsb 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 large 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.

' 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 loDg was taken on Georges Bank by
the schooner Ethel Merriarn, 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
(any 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
Straits 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, Tetrapterus
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, Farrington5 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 Vesey-Fitzgerald 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

Figure 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
that are deepest forward. Its anal fin, 26 to 30
rays, 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 of Maine. — A dolphin
about Z}{ 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 butterfisb.es (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 which 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 Gulj 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.

• Brama raii has been taken at Woods Hole and on the Grand Banks.

• Bull. Mus. Comp. Zoo]., vol. 69, 1929, pp. 39-50, 1 pi.

FISHES OF THE GULF 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

=■ Pt>pr,lu<; -triaxati THuj

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 fromthescup (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 l)i 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

liss^^^^'

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.11 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.

" 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 Goode13 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,

" Information furnished by O. E. Sette. The illustrations of larvae 2.1
mm. and 3.4 mm. long credited by Kuntz and Radclille (Bull. U. S. Bur.
Fish., vol. 35, 1918, figs. 63 and 64) to the butterfish and reproduced in the
previous edition of this book (Bigelow and Welsh, Bull. U. S. Bur. Fish. vol.
40, Pt. 1, 1925, fig. 116, c and d) have since been proved 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 (26 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
feU into two groups: one ranging from 4 to 5%
inches (averaging about 4%) and the other from
iy2 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 u 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 IS 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

i< Bull. U. S. Bur. Fish. vol. 43, Pt. 1, 1928, p. 214.

u Hoar (Copeia, 1937, p. 238) records two large ones from Margareo 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 I8 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.
*> Comish, 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 2l
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 26-28 th 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 their 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, S50 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

11 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.21 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 tbat summer
off southern New England and farther north and
east. Butterfish, it is true, have been trawled in
February near tbe 90-fathom line abreast of the
eastern part of Long Island, N. Y. ; a6 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.

53 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 flsh taken by Albatross II, February 27, 1929.

Abundance. — During the period 1928 to 1947 2T
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.

>8 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, M
are the only Gulf of Maine records of which we
know.

>° See Meek and Hildebrand (Field Mus. Nat. Hist., Zool. series, vol. 15,
Ft. 2, p. 411, 1925) for discussion.
»' Reported to us by the late Walter H. Rich of the U. S. Bur. Fish.
*> MaoCoy, Bull. 69, Boston Soc. Nat. Hist., 1933, p. 9.
» Reported to us by Edgar Arnold.

Figure 194. — Harvestfish (Peprilus alepidolus), New York. From Goode. Drawing by H. L. Todd.

FISHES OF THE GTJLF 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 percijormis
(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 % 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 gulfweed, 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.36 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;39 most
plentiful south of Cape Cod. Probably it is

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,39
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, 1947.41

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 )i 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

31 Trans. Connecticut Acad. Arts. Sci., vol. 36, 1945, pp. 842-843.

" Notes by Vinal Edwards.

* According to Vladykov and McKenzie (Proc Nova Scotian Inst. Sci.,
vol. 19, 1935, p. 87) occasional specimons are caught off 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).

« Trans. Connecticut Acad. Arts Sci., vol. 36, 1945, pp. 842-843.

» Smith, Copeia, 1921, No. 91, pp. 9-10.

■ 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.

•i 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.

W*a**to»ii*^^

Figure 196. — Black ruff (Centrolophus niger), Dennis, Mass. From Goode and Bean. Drawing by S. F. Denton.

FISHES OF THE GTJLF 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 ** 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; w a second of 2lK
inches was brought in from the northern edge of
Georges Bank by the trawler Thomas Whalen in
September 1936; M 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, 1951.46

Another about 9 inches long was taken in 188848
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. V. S. Nat. Mus., vol. 21,
In the U. S. National Museum.
" Reported by Bigelow and Schroeder, Copeia, 1937, p. 61

, p. 639 and now

« Received through the kindness of John Worthington of the Pond Village
Cold Storage Co.
" Qoode 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

Kudderfish; 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 only
about half as long (16 or 17 rays), 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

Figure 197. — Pilotfish {Naucrates duclor), 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 Gulj of 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 (lat. 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 Scotlan Inst. Scl., vol. 19, 1935, p. 6), 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) 1815.60

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.

M The interrelationships of the several Seriolas that have been described
from our South Atlantic coast still remain in doubt.

•i We have examined specimens ranging from 3 to 9 inches in length taken
in Cape Cod Bay, at Woods Hole, New Bedford, and other localities.
Storer's illustration, reproduced here, was of a 2-inch fish.

Figtjbe 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 oj 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 b}{ 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

o We have no color notes from life.

» Reported by Leim, Proc. Nova Scotian Inst. Sci., vol. 17, 1930, No. IV,
p. xlvi, as S. dumerili.
M Reported to us by Walter H. Rich.

" Reported by Firth, Bull. 61, Boston Soc. Nat. Hist.; 1931, p. 12.
" 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 6T
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.68 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

" 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.

■ 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
line 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 oj 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 fish, a 2H-inch specimen, caught October 10, 1928, was recorded
by Leim (Proc. Nova Scotian Inst. Sci., vol. 17, No. 4, 1930, p. xlvi).

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) ,
hardtail (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
hardtail 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 either side of its
caudal peduncle ; to its flattened oblong form (body

« The yellow tall (Chloroscombrut chrysurut), another species in this group
straggles northward at times and, sometime, may be taken within the Oulf
of Maine. It may bo d istinguished from the crevalle, hardtail, saurel, and big-
oyed 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 {Caranx kippos), 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; 66 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.

Hardtail Caranx crysos (Mitchill) 1815

Yellow jack; Runner; Yellow mackerel

Jordan and Evermann, 1896-1900, p. 921.

Description. — The hardtail resembles the cre-
valle, saurel, and goggle-eyed scad in the rel-

•* Reported near Halifax, Nova Scotia, by Vladykov (Proc. 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 first 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.

Figure 201. — Hardtail (Caranx crysos). Woods Hole. From Goode. Drawing by H. L. Todd.

FISHES OF THE GULF OF MAINE

377

off Gloucester,68 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."

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.

70 A 5-inch specimen is reported from Port Mouton by Leim (Proc. Nova
Scotian Inst. Sci., vol. 17, 1930, No. 4, p. xlvi), and small ones from Pubnico,
and near Halifax, by Vladykov (Proc. Nova Scotian Inst. Sci., vol. 19, 1935,
p. 4).

"Nichols (Bull. Amer. Mus. Nat. Hist., vol. 42, 1920, p. 479) considers
tbe western Atlantic saurel distinct from the eastern Atlantic saurel and
has proposed tbe name lalhumi for it. But this separation has not been
adopted generally.

Color. — Described as bluish green above, silvery
below, with a black spot on the edge of the gill
cover above its rear angle.

Size. — LeDgth 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) 1793

Goggler; Goggle eye jack

Jordan and Evermann, 1896-1900, p. 911.

Description. — This scad resembles the mackerel
scad (p. 374) in general appearance, but it has
larger eyes aDd lacks the detached finlets behind
the dorsal and anal fins. Its high first dorsal fin
separates it readily from the pilotfish (p. 372),
while the fact that the forward half of its lateral
line is only slightly arched instead of strongly so
obviates any danger of confusing it with the
crevalle (p. 375), the hardtail (p. 376), or the saurel
(p. 377). Its first dorsal fin has 8 spines, its second
dorsal is of 1 spine followed by 23 to 26 soft rays;

" Kendall, Bull. No. 58, Boston Soc. of Nat. Hist., 1931, p. 11.
" We received this specimen from Capt. Benjamin Morrow and It is now
in the Museum of Comparative Zoology.

Figure 202. — Saurel (Trachurus trachurus), Rhode Island. From Goode. Drawing by H. L. Todd.
210941—53 25

378

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 203.— Goggle-eyed scad (Trachurops crumenophthalmus) , Woods Hole. From Goode. Drawing by H. L. Todd.

its anal fin has 1 spine and 20 to 23 rays, and is
preceded by 2 stout detached spines. Its caudal
is forked. Its ventrals originate a little behind the
pectorals which reach nearly or quite as far back
as its vent. Its entire breast is scaly, as are parts
of its head.

Color. — Bluish above, silvery below. The fins,
snout, and tip of the lower jaw have dusky
markings.

Size.— Grows to a length of about 2 feet.

General range. — Cosmopolitan in warm seas,
straying as far northward on our Atlantic Coast as
Cape Breton, Nova Scotia.74

Occurrence in the Gvlf of Maine.- — The only
positive records of this species in our Gulf are of
one 5% inches long taken in a trap at Provincetown
on August 27, 1930 ;75 a second 8 miles off Chatham,
Cape Cod ; 76 and a third from Sandwich, on Cape
Cod Bay, in the summer of 1950. 77 It may be
expected to round Cape Cod from time to time,
for it is taken in summer and fall as far northward
and eastward as Woods Hole. ., ....

Moonfish Vomer setapinnis (Mitchill) 1815

Shiner; Horsefish; Bluntnose; Dollakfish

Jordan and Evermann, 1896-1900, p. 934.

Description. — The very deep, thin, sharp-edged
body of the moonfish (adults are scarcely twice as

" Reported from Canso by Cornish, Contrib. Canadian Bio!. (1903-1905)
1907, p. 85. A. H. Lelm advises us that a specimen 145 mm. long was taken
ofl Centre East Pubnico, Nova Scotia, September 12, 1951.

'• Firth, Bull. 61 Boston Soc. Nat. Hist., 1931, p. 11.

» MacCoy, Bull. 70, Boston Soc. Nat. Hist., 1934, p. 6.

" This specimen was received from Capt. Benjamin Morrow and is now in
the Museum of Comparative Zoology.

long as deep, and young fry even deeper, rela-
tively), tapering to a slender caudal peduncle, and
the concave upper anterior profile of its head, are
enough to separate it at a glance from pilotfish,
scad, crevalle, hardtail, saurel, or goggle eye; its
very low dorsal and anal fins distinguish it from the
lookdown (p. 379), which is of something the same
shape (cf. fig. 204 with fig. 205). Its minute
ventral fins, soft dorsal fin and anal fin which are
nearly even in height from end to end, separate it
from the threadfin (p. 381), and from the Cuban
jack (Hynnis cubensis), now thought to be the
adult of the threadfin (p. 381).

The first dorsal of the adult moonfish is reduced
to 8 very short, inconspicuous, detached spines,
but the first two of these are prolonged and fila-
mentous in young fry. Its second dorsal fin (21
to 23 rays) and its anal fin (17 to 19 rays) are about
equal in length, both of them low and tapering
very slightly from front to rear. In very small
fish the second to fourth rays of the second dorsal
fin are more or less prolonged, and the anal fin is
preceded by 3 or 4 short detached spines which
are not to be seen in the adult. The ventrals are
so small that they are likely to be overlooked
except in young fry, in which the ventral rays are
more or less filamentous as are the dorsal spines.
The pectorals are scythe shaped. The scales along
the lateral line are not large enough to be con-
spicuous, and the teeth are very small. There
are no detached finlets, dorsal or anal.

Color. — Bluish green above, bright silvery on
the sides. The second dorsal fin is plain pale
greyish, sometimes light yellow at its base; the

FISHES OF THE GULF OF MAINE

379

Figure 204. — Moonfish {Vomer setapinnis). After Goode. Original drawing by H. L. Todd.

caudal is greenish yellow; the pectorals light
yellow or dusky greenish.

Size. — Said to reach 1 foot in length, but most
of them are less than 9 inches long.

General range. — Warm seas off the east coast of
America from Uruguay to Cape Cod, straying to
Nova Scotia; common from Chesapeake Bay
southward.

Occurrence in the Gvlj of Maine. — This waif from
warmer waters has been recorded from the South
Channel off Cape Cod (one specimen 1% inches
long) ; off Cape Cod (60 miles south by east from
Highland Light), where one was taken in a mack-
erel seine, August 23, 1929 ;78 from Gloucester
(several specimens) ; from Magnolia, Danvers,
Salem, and South Boston (a specimen 2 inches
long) around Massachusetts Bay; from Saco
Beach (fry of about 1 to 3 inches) ; and from Casco
Bay in Maine. It has even been reported once
or twice as far east as Liverpool and Halifax,
Nova Scotia.79 Thus it appears to reach our
Gulf rather more often than any of its relatives
do; not often enough, however, for most of the
fishermen of whom we have inquired to know it
north of Cape Cod. It appears more often3(if
irregularly) at Woods Hole, where young fish
are sometimes common in August and September.

" Reported by Firth, Bull. 61, Boston Society Natural History, 1931, p. 12.
n Leim (Proc. Nova Scotian Inst. Sci., vol. 17, No. 4, 1950, p. xlvi); Vlady-
kov (Proc. Nova Scotian Inst. Sci., vol. 19, 1935, p. 8).

Lookdown Selene vomer (Linnaeus) 1758

Horsehead; Moonfish

Jordan and Evermann, 1895-1900, p. 935.

Description. — The very high second dorsal
(about 22 rays) and anal fins (about 20 rays) of
the lookdown, and their peculiar falcate outline
with the second ray much the longest and the
next 4 or 5 rays successively shorter make dis-
tinction easy between it and the moonfish. And
its peculiar form is hardly less characteristic, for
it shares with the moonfish a deep, rhomboid, but
very thin flat body (the trunk is only about one
and one-quarter times as long as deep), abruptly
truncate in front, with slightly concave upper
anterior profile, and tapering rearward to a slender
caudal peduncle. The mouth is set so low and
the eye so high that the expression of its face is
very characteristic. When adult the first dorsal
is reduced to 7 or 8 short inconspicuous spines,
only the first 3 of which are connected by a mem-
brane, and the ventrals are very small; but some
of the spines of the first dorsal are very long in
fry up to 4 or 5 inches in length, the ventrals are
much longer than in the adults, and the anal fin
is preceded by two short detached spines that
disappear with growth. The caudal fin is deeply
forked like that of other pompanos, and the
pectorals are sharp pointed and falciform, reaching
back behind the middle of the second dorsal fin.

Color. — Small specimens, and northern strays
usually are small, are silvery above as well as

380

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 205. — Lookdown (Selene vomer). From Goode. Drawing by H. L. Todd.

below, with the ground tint of the back leaden;
the sides are barred with several crossbands,
variously described as dark or golden. But these
bands fade out with growth.

Size. — Reaches a weight of about 2 pounds.

General range. — Warm waters on the east and
west coasts of America, north rarely to Cape Cod,
straying to the Gulf of Maine and to Nova Scotia; m
common from Chesapeake Bay southward.

Occurrence in the O-ulf of Maine. — There were
only three records for the lookdown in our Gulf
up to 1933; two of them for Casco Bay, the third
for Boston Harbor (Dorchester) . But many small
ones were reported from the traps at the mouth of
Casco Bay during that autumn, one from Beverly
on the north shore of Massachusetts Bay, and
one from North Truro on Cape Cod. Evidently
this was an unusual incursion, for no one would be
apt to overlook so bizarre a straggler from the
south.

Leatherjacket

Oligoplites saurus
Schneider) 1801

(Bloch and

10 Jones (Proc. and Trans. Nova Scotian Inst. Sci., vol. 5, App., 1879,
p. 89) and Honeyman (Trans. Nova Scotian Inst. Sci., vol. 6, 1886, p. 323)
report young fry as occasionally found in the shore waters of Nova Scotia,
presumably along the outer coast, for tropical fishes are taken oftener there
than along the Gulf of Maine shore of the Province.

Jordan and Evermann, 1896-1900, p. 898.

Description. — The most interesting character of
the leather jacket, and one which places it at
a glance, is that the rear part of its soft dorsal fin
back from the 7th ray, and also its anal fin back
from the 5th ray, is broken, as it were, into a series
of 12 low nearly separate finlets, the ray in each
of which is subdivided at the tip like the hairs of
a little brush. We need only note further that
its body is about 3K times as long as it is deep,
very strongly flattened sidewise, and thin, being
only about one-third as thick as it is deep; its
upper jaw bone reaches back about as far as the
rear edge of the eye; its snout is moderately
pointed; its caudal peduncle very slender, with a
low, inconspicuous keel on either side. Its first
dorsal fin is reduced to about 5 separate spines,
each with small fin membrane and its second
dorsal has about 20 rays; its soft anal fin, also
of about 20 rays, is preceded by two stout and
conspicuous spines, forming, together, a separate
finlet. Its lateral line is nearly straight, and its

FISHES OF THE GULF OF MAINE

381

Figure 206. — Leatherjacket (Oligoplites saurus), Marthas Vineyard. From Jordan and Evermann. Drawing by

H. L. Todd.

scales are very small, and imbedded in the skin,
which is corrugated with a great number of short,
fine, longitudinal ridges, giving it a leathery
appearance, hence its common name.

Color. — Bluish above, silvery below, with yellow
fins.

Size. — The largest are about 12 inches long.

General range. — Common on both coasts of
tropical America; northward to New York and
southern Massachusetts (Woods Hole), reaching
the southwestern part of the Gulf of Maine as a
stray.

Occurrence in the Gulf of Maine. — The only rec-
ord of this southern fish within the Gulf is of one
taken in a trap off the outer beach at Chatham,
Cape Cod.

Threadfin Alectis crinitus (Mitchill) 1826 8I

Jordan and Evermann, 1896-1900, p. 931, as Alectis
ciliaris (Bloch 1788).

Description. — The combination of a head
strongly convex in dorsal profile, with the fact that
the first few rays of its soft dorsal fin, and of its
anal fin also, are extremely long, and threadlike,
places the threadfin at a glance among the ca-
rangoids of our northeastern coast. On small fish
these threadlike rays are considerably longer than
trunk and tail combined, but they shorten with
age, probably to be entirely lost. The trunk of
the threadfin is nearly as high as it is long (to the
caudal peduncle), the dorsal profile of its head is
strongly convex and it is strongly flattened side-

" We follow Smith (Copeia. 1938, p. 146) in using tbe name crimtus Mitchill
1826, proposed for a specimen taken near Block Island, R. I., rather than
ciliaris Bloch 1788 (type locality East Indies), awaiting final decision as to
the true relationship between the threadfins of different oceans.

wise. It has one dorsal fin of 1 stiff ray and about
19 soft rays, preceded by 6 short, separate, in-
conspicuous spines; the anal has 1 stiff ray and
16 soft rays, and is preceded by 2 spines so short
that they are likely to be overlooked. Its lateral
line is strongly arched over the pectoral; and the
rear part armed with a series of bony platelike
scales; the tail fin is deeply forked and the ventrals
are larger than in most other carangoids. The
pectorals are about as long as the head.

Color. — Upper surface bluish, the sides silvery,
with traces of darker bars and blotches that tend
to disappear with age; the prolonged parts of the
dorsal and anal fins are bluish black; ventral fins
mostly black; the fins otherwise more or less
yellowish.

Size. — Specimens with the long threadlike fin
rays have been reported up to about 7 inches long,
in West Indian and Atlantic United States waters.
But it now seems very probable that these are the
young of the Cuban Jack (Hynnis cubensis Poey
1860). Their transformation consists chiefly in
losing the filamentous fin rays; in a decrease
in the depth of the body relative to its length; in
a very considerable decrease in the relative size of
the ventral fins; and in the assumption of a more
falcate shape by the pectorals.

General range.- — The threadfin (or threadfin
stage of the Cuban Jack), is known on both coasts
of tropical America; it strays northward on the
Atlantic coast to southern Massachusetts, and it
has been reported once from the Gulf of Maine.
The adult Cuban Jack has not been reported north
of southern Florida.82

93 We have seen one taken at Key West, and there is one from the east
coast of Florida in the Museum of Comparative Zoology.

382

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 207.— Threadfin (Aleciis crinitus), from
Florida specimen mounted by Al. Pfleuger and
another small specimen from Newport, Rhode
Island. Drawing by H. B. Bigelow.

Occurrence in the Gulf of Maine. — The only
records of this tropical fish for the Gulf of Maine
are of one about 3}i inches (85 mm.) long taken in
a trap at Sagamore, on the southern shore of Cape

Cod Bay, September 1, 1937, and another taken
in a trap at North Truro, Mass., August 16, 1951.
They may have come through the Cape Cod
Canal as suggested by Smith.83

THE BLUEFISHES. FAMILY POMATOMIDAE

The bluefish (the only member of its family)
resembles the pompano family in the general
structure and arrangement of its fins, there being
two dorsals, the first spiny and the second soft,
with the ventrals well forward under the pectorals.
But it lacks the free spines in front of the anal fin
which are characteristic of most pompanos; its
caudal peduncle is deeper; its tail is less deeply
forked; and its teeth are much larger. It bears
a superficial resemblance to certain of the weak-

fish family (p. 417) in its general body form and in
the arrangement of its fins. But it is readily
separable from any of the latter by the fact that
its anal fin is nearly as long as its soft (second)
dorsal, and from the sea-bass family in that its
first (spiny) dorsal is much lower than the second.
Most American ichthyologists look upon the blue-
fish family as closely allied to the pompanos, but

» Copeia, 1938, p. 146.

FISHES OF THE GULF OF MAINE

383

it should be grouped with the sea-bass tribe ac-
cording to another view because of skeletal
characters.

Blucfish Pomatomus saltatrix (Linnaeus) 1758 8*
Snapper (Young)
Jordan and Evermann, 1896-1900, p. 946.

Description. — According to Jordan and Ever-
mann, and to most of their successors, the bluefish
is separable from its closest allies, the pompanos
(Carangidae), by a tail "not deeply forked" and
by larger scales, statements that may easily be
misleading, for while the bluefish certainly bas a
less deeply forked tail than the pompanos, anyone,
we think, would describe it as deeply forked as
compared with any square-tailed fish. And while
its scales are larger than those of most pompanos
there is not much difference in this respect be-
tween a bluefish and a large crevalle (p. 375).
There is, however, one positive point of difference.
The jaws of the bluefish, upper as well as lower,
are armed all around with a single series of stout,
conical, canine teeth (one-eighth to one- fourth of
an inch long in a fish of about 10 pounds), whereas
the crevalle alone of northern pompanos has
canines, and only two of them. Furthermore, the
caudal peduncle of the bluefish is stouter than that
of any pompano. It is sharply differentiated
from all mackerels by the absence of dorsal or
ventral finlets.

The bluefish is moderately stout bodied (large
ones are about one-fourth as deep as long) ; its
belly is flat-sided but blunt-edged below; its caudal
peduncle moderately stout (slimmer, however,

than in many other fish, e. g., striped bass) ; its
head deep; its nose moderately pointed; and its
mouth large and oblique, with projecting lower
jaw, and with prominent canines. "Snappers,"
as small bluefish are called, are relatively deeper
and more flattened sidewise than larger fish. The
first dorsal fin (7 or 8 stout spines), originating
over the middle of the pectorals, is low, rounded,
depressible in a groove. It is separated by only a
very short interval from the second dorsal, which is
more than twice as long as the first (about 23 to 26
soft rays) and about twice as high, tapering back-
ward with slightly concave margin. The anal fin
(25 to 27 rays) is similar in form to the second
dorsal though with a somewhat less concave outer
margin; it originates somewhat farther back and is
preceded by two very short detached spines that
are often hidden in the skin. The caudal is broad
and forked, moderately or deeply according to the
other fish with which it is compared. The ventrals
and pectorals are both of moderate size. The
body, most of the head, and also the second dorsal
and anal fins are clothed with medium-sized scales.
There are no shields or keeled scales along the
lateral line nor is the caudal peduncle keeled.

Color.- — Sea-green above; silvery below. The
second dorsal, caudal, and pectoral fins are of the
general body tint, the latter with a black blotch at
the base.

Size.- — Maximum length about 3}i feet. The
heaviest American fish of which we find definite
record within recent years was 3 feet 9 inches long,
weighing 27 pounds,86 caught off Nantucket in
1903. One of 20 pounds was taken off Montauk,
N. Y., in August 1951.86 It is said that fish of 30
or even 50 pounds were not unheard of during the

M This fish has been known by various vernacular names along the middle
and southern coasts of the United States. But it is the "bluefish" in the
Gulf of Maine.

«' Smith, Forest and Stream, vol. 61, October 10, 1903, p. 283.
» Reported in Salt Water Sportsman, August 17, 1951.

bM%*&&L

f^4|||$3s*-

Figure 208. — Bluefish {.Pomatomus saltatrix). From Jordan and Evermann. Drawing by H. L. Todd, from a cast.

384

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

last half of the eighteenth oentury, but these huge
fish may not have been weighed. And the general
run of the largest fish that are caught off the
American coast is only 10 to 15 pounds. But they
run larger off the African coast where 20-pounders
are not unusual and where one of 45 pounds
has been reported.87 A 1-pound fish is about
14 inches long; a 2-pounder about 17 inches; a
3-pounder about 20 to 21 inches; a 4-pounder,
about 2 feet; and an 8-pounder about 28 to 29
inches long. Fish weighing from 10 to 12 pounds
are about 30 inches long.88

Habits. — The bluefish is oceanic in nature, found
indifferently inshore, offshore, and in many parts of
the ocean (p. 385). It usually travels in schools,
sometimes including many thousands ; in 1901, for
example, a school 4 or 5 miles long was reported
as seen in Narragansett Bay. And it is perhaps
the most ferocious and bloodthirsty fish in the sea,
leaving in its wake a trail of dead and mangled
mackerel, menhaden, herring, alewives, and other
species on which it preys. Goode 89 wrote long ago,
the bluefish, "not content with what they eat,
which is itself of enormous quantity, rush raven-
ously through the closely crowded schools, cutting
and tearing the living fish as they go, and leaving
in their wake the mangled fragments." It is not
only the schooling fish that fall prey to them, but
scup, squeteague, hake, butterfish, cunners, and
small fish of all kinds, besides squid. Baird
writing in the 1870's, when bluefish were at the
height of their abundance, estimated that they
annually destroyed at least twelve hundred million
millions of fish during the four summer months off
southern New England; and while this calculation
surely was wildly exaggerated it will help give
the reader a graphic realization of the havoc that
they wreak during their periods of plenty. They
are also known to eat various Crustacea and even
marine worms on occasion. And the young
"snappers," 6 to 8 inches long, feed largely on
copepods, or crustacean and on molluskan larvae,
as well as on fish fry smaller than themselves.

Bluefish are creatures of warm water, never
found in any numbers in temperatures lower than
about 58° to 60° (at least in summer); and they
appear along the United States coast as warm-

87 By Lt. Commander Henry Lyman (Bluefishing, 1950, p. 9) who also saw
a 22-pounder weighed off northwest Africa, with still larger ones that were not
weighed.

» Goode, Fish. Ind. U. S., Sect. 1, 1884, p. 442.

88 Fish. Ind. U. 8., Sect. 1, 1884, p. 544.

season migrants only. "Bluefish," writes Ly-
man,90 "appear off the southern coast of Florida in
midwinter," and by "late March anglers take
them off the Florida coast in good quantities."
"Large schools pass the Carolinas during March
and April, appear off Delaware during April, and
are first taken off New Jersey and Long Island,
N. Y., during April and May," by commercial
fishermen working well offshore. The earliest
commercial catches are reported off southern
Massachusetts in late May. But it is not until
about a month later that they work inshore in
numbers.

When they do come inshore, midtitudes of little
ones, known as snappers, run up into harbors and
estuaries all along the coast, from Delaware Bay
to Cape Cod. The larger ones, arriving some-
what later, also often come close enough in to the
beach, west and south of Cape Cod, for many to be
caught by anglers casting in the surf. But it is
only in good years that this last holds true in our
Gulf, even in the southern part.91 When they "first
appear offshore, in any locality, almost always they
will be feeding deep, at or near the bottom. This
means that surface lines and baits are practically
worthless." 92 Later in the season schools are often
seen at the surface, harrying other fish; and if they
are deep, they can often be lured to the surface by
throwing out ground bait.

Except for an occasional belated fish (p. 388) , the
bluefish disappear wholly from the entire coast
northward from the Carolinas by early November.
The winter home of this northern contingent has
long been the subject of speculation. But the
fact that we saw one trawled in 55 fathoms off
Marthas Vineyard in mid-January in 1950 by the
Eugene H, and that several hauls of 175 to 1,400
pounds per trip were brought in from the region of
the Hudson Gorge by otter trawlers early that
same February, makes it probable that most of the
members of the northern contingent merely move
offshore on bottom, to the warm zone along the
outer edge of the continent, to pass the winter
there. It is certain, however, that some migrate
far southward (as has often been suggested for the
stock as a whole) for one that was tagged off New
York in August 1936 was recaptured off Matanzas,

" Blueflshing, 1950, pp. 10, 11.

81 We refer the reader to Lt. Comdr. Lyman's recent book (Bluefishing,
1950, pp. 34-19) for an interesting survey of the more-productive blueflshing
grounds, Gulf of Mexico arid Florida to Cape Cod.

•» Quoted from Lyman, Bluefishing, 1950, p. 11.

FISHES OF THE GULF OF MAINE

385

Cuba, in January 1939. 93 Whether wanderers
such as this ever return to the north is unknown.

A few bluefish are caught in winter on both
coasts of Florida, southward from Cape Canaveral
in the east, from Tampa Bay on the west; and
enough are taken near Key West between Decem-
ber 15 and February 15, to yield commercial
catches of 10,000 to 15,000 pounds in most years.94
Some, also, are caught around Cuba by commercial
fishermen in January and February. But these
Florida fish, presumably the Cuban also, vanish at
the end of the winter, not to reappear until early
the next. What their relationship may be to the
northern stock is not known. There are bluefish,
too, off the northern coast of the Gulf of Mexico
and off Texas, but nothing definite is known about
their seasonal movements.

It is not likely that any interchange ordinarily
takes place between the bluefish populations of the
two sides of the Atlantic.

Females with large ova approaching ripeness
are taken off North Carolina in spring, and off
various parts of the coast farther north in sum-
mer; 95 ripe males have even been taken inside
Chesapeake Bay in June and July,96 from which it
appears that they spawn from late spring through
July and perhaps into August. But bluefish have
never been reported actually spawning, though
watch has been kept for them, which makes it
likely either that they interrupt their inshore visit
to move offshore for the purpose, perhaps sinking
deep, or that most of them have spawned out be-
fore they appear along our northern coasts. In
either case, the regular presence of "snappers" in
numbers inshore, and the occasional captures of
smaller fry in Chesapeake Bay 97 and in the Gulf of
Maine (p. 388) make it likely that the spawning
grounds of our northern bluefish are not far
distant.

The eggs have not been identified with certainty.
But the possibility is still open that the buoyant
eggs with segmented yolk and large oil globule
from Newport, R. L, provisionally referred to the

•» Reported by Lyman (Bluefishing, 1950, p. 10.)

•' Schroeder, App. 12, Rept. U. S. Commissioner of Fisheries (1923) 1924,
p. 12.

18 Old statements to this effect are corroborated by Lyman (Bluefishing,
1950. p. 10), who reports females with roe and males with milt off North
Carolina and near Nantucket early in summer.

'• Hildebrand and Schroeder, Bull. U. S. Bur. Fish. vol. 43, Pt. 1, 1928, p.
232.

" Hildebrand and Schroeder (Bull. IT. S. Bur. Fish., vol. 43, Pt. 1. 1928, p.
232) report fry as small as 2% inches in Chesapeake Bay.

210941— '53 26

bluefish by Agassiz and Whitman98 were actually
those of this species. And while the identity of
their "bluefish" larvae has likewise been ques-
tioned, we believe that then identification of the
oldest (9 mm. long 99) was correct, though the
younger ones may have belonged to some one of
the mackerel tribe.

At this stage the second dorsal fin is formed, the
first, however, still represented by the rudiments
of the future spines. The anal fin is visible,
also, and the tail is slightly forked. These larvae,
like those of the mackerel (which they much
resemble), have large blue eyes and large pro-
jecting teeth, but they are as far advanced in
development as mackerel twice as large, and as
ferocious in proportion to their size as the adult
bluefish, devouring all other small animals in the
tank with them.

The bluefish fry of three-fourths of an inch to 3
inches long, which have often been taken along
shore in summer not only south of Cape Cod but
even in the Gulf of Maine in some years (p. 388),
are presumably the product of that season's
spawning. And it seems that they grow to a
length of 4 to 9 inches by autumn, fish of that size
being common in October, while general experience
suggests a length of 8 to 12 inches by the following
spring. Nothing definite is known of the rate of
growth of the older fish,1 except that one that
weighed about 1 pound when it was tagged off the
coast of New York on August 10, 1936, was
reported as weighing about 9 pounds when it was
recaptured off Matanzas, Cuba, two years and five
months later (January 15, 1939), which (if not
exaggerated) 2 points to unexpectedly rapid
growth.

The age at which the bluefish matures sexually
is not known.

General range. — Widely but irregularly distrib-
uted in the warmer seas, its known range includ-
ing the eastern coast of the Americas, northward
regularly to Cape Cod, occasionally to outer
Nova Scotia, south to Brazil and Argentina; 3
Bermuda; 4 eastern Atlantic off northwestern

•» Mem. Mus. Comp. Zool., vol. 14. No. 1, Pt. 1, 1885, p. 13, pi. 4, figs 1-6.

»• Mem. Mus. Comp. Zool., vol. 14, No. 1, Pt. 1, 1885, pi. 5, fig. 15.

1 No growth studies based on the scales or on other exact methods have been
undertaken for the bluefish, to our knowledge.

3 Lyman (Bluefishing, 1950, p. 10), who reports the case, suggests that the
fisherman who re-caught the fish "may have been stretching things a bit."

* Frozen bluefish have recently been imported from northern Argentina.

' See Lyman, Bluefishing, 1950, p. 12, for photo from the Bermuda News
Bureau of a 15-pound bluefish caught at Bermuda, February 1949.

386

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Africa ; also Mediterranean ; both coasts of south-
ern Africa; Madagascar; eastern Indian Ocean and
Malay Peninsula; southern Australia and New-
Zealand.

Occurrence in the Gulf of Maine. — Bluefish
have been taken at one time or another wherever
any information is available as to the local fishes
around the western side of the Gulf. But they
have seldom been seen east of Penobscot Bay
(reported at Mount Desert in 1889); we have
heard of only one taken in the Bay of Fundy, a
fish caught in Minas Basin in July 1951,6 and
we have found no record of bluefish off the Nova
Scotian coast of the open Gulf of Maine. But
one was caught off Halifax in 1925, another more
recently near Liverpool on the outer coast of
Nova Scotia,6 and they were reported "common"
near Port Medway, Nova Scotia, in the summer of
1951.7

In our Gulf, too, they seem to be confined to
the vicinity of the coast (they are unknown in the
central basin or on Georges Bank), the small ones
("snappers") running up into brackish water,
as in the Parker River, but the larger sizes
(3 pounds or more) keeping to the outside waters.

The geographic distribution of the places where
they have been recorded would suggest at first
glance that bluefish are practically universal in the
western side of the Gulf. But this is true only
for brief terms of years and at long intervals, for
while they have been known to swarm there for
several summers in succession, they may then
be so rare over periods of many years that the
capture of a single fish causes remark.

Bluefish must have been common at the time
of the first settlement, at least as far north as
what is now southern Maine, for Josselyn, writing
in 1672, referred to them as better meat than the
salmon.

Bluefish were plentiful off southern New
England and also about Nantucket in colonial
times, but they seem to have disappeared thence
about 1764, not to reappear there until about
1810. From that time on they increased in
abundance west and south of Cape Cod, but none
were reported north of the Cape until 1837. And
since a fish as ubiquitous as the bluefish would

certainly have attracted attention and its presence
found its way into print, had it been abundant
in the Massachusetts Bay region, it is safe to say
that very few, if any, visited the Gulf of Maine
during the late eighteenth century, or the first
quarter of the nineteenth.

According to Storer, the first bluefish seen north
of Cape Cod after their long period of absence,
was one caught on October 25, 1837; Captain
Atwood 8 saw them for the first time at Province-
town in 1838. According again to Storer, bluefish
were taken yearly from the wharves at Boston
after 1844. And by 1850 they were so plentiful
about Cape Ann that fishermen complained
of them as driving away most of the other school-
ing fish, while in 1863, which seems to have marked
the culmination of this flood of bluefish, they were
extremely abundant in the Massachusetts Bay
region and especially at Province town.9 They
remained plentiful in the southern part of the
Gulf of Maine for several summers after 1863,
but by 1872 they were reported as much less so,
and there have not been enough bluefish anywhere
in the Gulf since the late 1870's to menace the
local mackerel fishery.

The yearly catch for the Cape Cod Bay region
had fallen to about 22,000 pounds by 1888 (93
pounds for Essex County), to only about 3,000
pounds for 1889. But some bluefish were seen as
far north and east as Mount Desert in that year,
and evidently more of them rounded the Cape
during the next 9 seasons, for the catches for the
years 1890-1898 were between about 26,000 pounds
and 80,000 pounds for Cape Cod Bay; with a few
hundred pounds for the Massachusetts coast
north of Boston. But this period of moderate
plenty was followed by a period of scarcity 10 so
extreme (detailed statistics are wanting) that no
catch as large as 5,000 pounds was reported again
as made anywhere in our Gulf in any year for
which statistics are available from 1900 u down to
the early 1920's. In 1906, in fact, in 1910 and
again in 1919, only an occasional school can have

' Reported to us by Dr. A. H. Lelm.

« Leim. Prcc. Nova Scotian Inst. Sci., vol. 17, Part IV, 1930, p. nvi.
' Information from Dr. A. H. Leim, from report by L. R. Day, Fisheries
h eseareb Board ot Canada.

8 Proc. Boston Soc. Nat. Hist., vol. 9, 1863, p. 189.

• Balrd (Kept. U. S. Comm. Fish (1871-1872) 1873, p. 237-240), and Ooode,
(Fish. Ind. U. S.; Sect. 1, 1884, p. 435-437) have collected much information
about the early history of the bluefish.

" Reported catches for the Cape Cod Bay region by all methods were
only about 3,600 pounds in 1899 and 7,659 pounds In 1900.

II Statistics of the pound net catches, by towns, were published in tbo
Annual Report of the Commissioners on Fisheries and Game of Massa-
chusetts for the years 1906, 1908, 1909, 1910, 1918, and 1919.

FISHES OF THE GULF OF MAINE

387

rounded Cape Cod,12 while bluefish must have
been practically nonexistent north of the Cape in
1918, for the entire reported catch there was only
34 pounds for that year.

We should also point out (we cannot explain
this) that a larger proportion of the bluefish than
usual that did round Cape Cod seem to have
continued on to the northern shore of Massachu-
setts Bay during this period of general scarcity.
Thus about as many (300 lb.) were reported for
Essex County in 1906 as for the Cape Cod Bay
region; about one-fourth as many in 1908, about
one-half to one- third times as many in 1908,
1909, and 1910.13

Bluefish must have come north in greater
numbers in 1927, for they were reported here and
there from Cape Ann northward during that
summer with small catches in the Casco Bay
region,14 and there seem to have been still more of
them in the Gulf during the next two summers, as
reflected in reported catches of 4,825 pounds for
Essex County, Mass., and 140 pounds for the
Casco Bay region, Maine, in 1928; 7,888 pounds
for Essex County and 495 pounds for Casco Bay,
Maine, in 1929. And so many blues invaded the
southwestern part of the Gulf during the next
three years that about 68,000 pounds were re-
ported for Essex County, and 200 pounds for
Casco Bay in 1930; 60,000 pounds for Essex
County and 500 pounds for Casco Bay in 1931;
and 1,414 pounds for the coast of Maine as a
whole in 1932.16

Eighty pounds of bluefish were taken in a set
of traps at North Truro on Cape Cod Bay on
September 9 in 1936;18 we heard of some large
ones caught in the surf on the outer shore of
Cape Cod that same year; we know of one caught
at Cohasset on the south shore of Massachusetts
Bay in 1937, and enough came north again in
1938 to provide a commercial catch of about
1,800 pounds for Essex County. But this slight
upsurge was followed by 6 years, or more, of
scarcity so extreme that only small catches were
reported from Maine (200 pounds in 1944), or

•> The catches north of Cape Cod ran only between about 300 pounds and
600 pounds for those years.

" Pound net catches of 1,015 pounds reported for Essex County, 4,623
pounds for the Cape Cod Bay region in 1908; 600 pounds and 1,342 pounds,
respectively, in 1909; 182 pounds and 419 pounds, respectively, in 1910.

» The Boston Post for July 24, 1927, reported 65 taken near Bald Head
one day, and 35 the next by Capt. Charles F. Pye.

11 No regional breakdown is available for Massachusetts for that year.

" Information contributed by the Pond Village Cold Storage Co.

from northern Massachusetts (200 pounds for
Essex County in 1945). The year 1946 was per-
haps the low point for our Gulf, when the total
catch including the southern coast of Massachu-
setts, was only about 1,200 pounds.

In 1947, however, when the total reported catch
for Massachusetts was only 2,300 pounds (none
reported for Maine)17, schools of small bluefisb, of
a pound or so, appeared along the inner shores of
Cape Cod, near Wellfleet, in August. They are
said to have been more widespread in Cape Cod
during the two next summers, and in 1950 bluefish
of % pound to IK pounds, with a few up to 4 or 5
pounds, rounded Cape Cod in such numbers that
Cape Cod Bay was described by anglers as
"loaded" with them during that August.

Many catches of 60 to 100 pounds were made
from party boats, both in the Wellfleet side of the
Bay and along the Sandwich shore; a set of 8
traps at North Truro made small catches (10 to
160 pounds) at intervals between July 18 and
October 7 of that year, while other traps around
Cape Cod Bay from Provincetown and Sandwich
made various catches between June and October.
Some also worked north across Massachusetts
Bay; witness captures of about 6 bushels of 1 to
1^-pound bluefish in a trap off Marblehead on the
north shore of Massachusetts Bay, July 21 .18 We
heard of at least one taken at Hampton, N. H.,
also a few at Kennebunkport, Maine, early that
September,19 and, at least, one from the lower
Kennebec River. They continued plentiful also
in Cape Cod Bay until the first week of that Sep-
tember. And while few, if any, were seen there
after the severe northeast gale of mid-September,
some were caught along the outer shore of Cape
Cod as late as the first week of that October, and
perhaps until later still.

All we can say, as to the catch in 1951 up to
this writing (August 15) is that the earliest report
of a bluefish in Cape Cod Bay was for June 19; a
small one was taken at Plum Island, northern
Massachusetts, on July 6; good catches (presum-
ably of small fish) were reported in Cape Cod Bay
by the last week of July (2,545 pounds taken in
one set of traps at North Truro, July 7-28); a
tremendous run of small bluefish were reported

>' This is the most recent year for which catch statistics have been published.
>< Reported in the Boston Herald for July 31, 1950.
i' Reported in the Saltwater Sportsman for September 15, and October 6,
1950.

388

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

near Provincetown during the first part of
August,20 and a few large fish, among great num-
bers of small ones were being taken off the south
shore of Massachusetts, and even in the Cape Cod
Canal ; some also were being caught in the rips and
in the surf at the tip of Monomoy Point.

It remains to be seen whether this increasing run
of bluefish in our Gulf is comparable to that of the
1860's. However this may prove, history will no
doubt repeat itself sooner or later, and these sea
pirates will again invade the Gulf in abundance,
probably for several summers in succession.

The disappearance of the bluefish from the Gulf
of Maine following the run of the 1860's was part
of a general shrinkage in the bluefish population
that visits the coast east of New York, and was
to be expected, for the bluefish that reach our
Gulf are only the northernmost fringe of the north-
ern contingent. The increase in the numbers
caught north of Cape Cod in the period 1928-31
was associated, similarly, with a corresponding
rise in the yearly catches made off southern New
England from about 55,000 pounds in 1928 to
about 650,000-920,000 pounds for 1930-33.

The scarcity of bluefish north of Cape Cod from
the early 1930's down to the early 1940's (in-
terrupted in 1938 as noted) was clearly the result
of the general decrease that took place in the
abundance of bluefish over the northern part of
their range as a whole, reflected in the southern
New England catch which fell from nearly a
million pounds in 1933 21 to an all-time low of
only 12,500 pounds in 1945. And there can be
no doubt that the small bluefish that have re-
appeared in the Cape Cod Bay region and north-
ward in increasing numbers during the past 2 or
3 years have been the overflow so to speak, from
an increasing population to the southward, great
numbers of which (mostly small) were being
caught from New Jersey to Nantucket in 1950,
and are being caught there with some large ones,
at this writing (August 15, 1951).

We ought perhaps to add that it is only in the
northern part of its range that the American
bluefish falls periodically to a very low level; in
1945, for example, when the total catch for New
England was only about 26,000 pounds, 223,000
pounds were taken in Chesapeake Bay and about

2% million pounds along the Atlantic and Gulf
coasts. This, again, was to be expected, for it
is near the boreal boundary of its range that any
warm -water fish is subject to the greatest vicissi-
tudes.

An interesting phase of the fluctuations of the
bluefish is that large numbers of very small ones
have visited the southern coast of New England
even in the poorest summers; some have been
reported within the Gulf of Maine. Thus fry so
small as evidently to have been the product of
that season's hatch were taken in Casco Bay,
Maine, in August 1899; slightly larger ones of
4 to 5 inches were caught off Plymouth in the
summer of 1921, and "snappers" are sometimes
reported at Provincetown, at Plymouth, and in
the Parker River in northern Massachusetts.
Almost all the fish, furthermore, that have been
taken within the Gulf of Maine, and the majority
of the larger catches that have been taken off
southern New England during the past few years
have been fish so small (mostly 1 to 2 pounds)
that it is not likely they had reached sexual
maturity. It is only in good bluefish years that
many of the mature fish (weighing upwards of
4 or 5 pounds) appear that far north. In poor
years large fish are caught in numbers only to the
southward of Long Island, N. Y. Prior to August
15, 1951 a number of fish up to 7 pounds had been
caught in southern New England waters, which
may indicate better things to come.

In the years when bluefish pass Cape Cod in
any numbers they usually appear in Cape Cod
and Massachusetts Bays about the middle of
June,22 sometimes as early as the first of that
month, and they are seen off and on all summer.
Most of them depart late in September, but an
occasional fish lingers into late autumn. Blue-
fish have even been caught about Provincetown
as late as December.

Importance. — The bluefish is an excellent table
fish, but it never has been plentiful enough to
support a fishery of any magnitude in the Gulf of
Maine. Nevertheless, its presence or absence
there may be a matter of direct importance to the
fishing interests, for it may drive away the mack-
erel when it swarms, if not the herring and men-
haden as well. Being a favorite game fish, many

■ Reported in Salt Water Sportsman for August 10.
" 920,9fi5 pounds reported in 1933, to be exact.

» Along southern New England the first blues are expected during the last
half of May (p. 384).

FISHES OF THE GULF OF MAINE

389

anglers troll for them in Cape Cod Bay in seasons
when there are enough of them to be worth fol-
lowing; also many are caught in the surf in good

years by anglers casting from the beach, as far
northward along the coast as the outer shore of
Cape Cod.23

THE SEA BASSES. FAMILY SERRANIDAE

The sea basses are an extremely numerous tribe
of perchlike fishes, with both the spiny portion
and the soft rayed portion of the dorsal fin well
developed, either as separate fins, or at least
divided by a deep notch. The ventral fins are
under the pectorals, technically thoracic, in
position. The anal fin is nearly or quite as long
as the soft part of the dorsal; the caudal peduncle
is deep and the tail is broad. In most of the
species the anal fin is preceded by 3 stout spines ;
the margin of the gill cover bears one or two sharp

conical spines in most, and the maxillary bone is
not sheathed nor hidden by the preorbital bone
when the mouth is closed. Smoother cheeks are
a ready field mark to distinguish any of the sea
basses from the rockfish family (p. 430); the 3
anal fin spines distinguish them from the croaker
family (p. 417) which have 1 or 2 anal spines only;
the spiny gill cover from the porgy family (p. 411);
and the large mouth from the cunner and tautog
tribe (p. 473).

KEY TO GULF OF MAINE SEA BASSES

1 . There is one continuous dorsal fin, its front part spiny, its rear part soft rayed 3

There are two separate dorsal fins, the first spiny, the second soft-rayed 2

2. The two dorsal fins are separated by a distinct inter-space; the sides are distinctly striped Striped bass, p. 389

The two dorsal fins are joined at their bases; the sides are not distinctly striped White perch, p. 405

3. The scales are large; the space between the eyes is naked; no bony ridge on the gill covers Sea bass, p. 407

The scales are small; the space between the eyes is scaly; there is a bony ridge on the gill covers.- Wreckfish, p. 409

Striped bass Roccus saxatilis (Walbaum) 1792 and keelless caudal peduncle, stout body, the

presence of two well-developed dorsal fins (spiny
Striper; Rockfish; Rock; Linesides and soft rayedi and the one about as long as the

Jordan and Evermann, 189&-1900, p. 1132, as Roccus other), its lack of dorsal or anal finlets, and a tail

lineatus (Bloch). only moderately forked, separate it from all the

_ . . __ , . . mackerel tribe, from the bluefish, and from the

Description. — No one character alone character- rpi „ *„ + *u„+ :+„ „„„i «„ i,„„ o ™;~,«,

, ... , , , ,. pompanos. Ihe lact that its anal nn has 6 spmes

lzes the striped bass, but rather the combmation

of fin structure and arrangement with general ,T%Tfte^toLJ^mrf-lM^vTf°^™?^t8TJ!!!!

f ° of bluefish tag methods and localities, also of tho natural history of the

outline and structure of the jaw. Its rather deep bluefish.

HinniffiiRi

'-■&&?£**£ 1-7 ' ' l : I ■ > \ * \ ' ■ 1 * * » * * * f * * \X i * i ■'■ '
■. ;■ [ 1 5 m ' i - f , .

§mmm0

P^UwkU'

Figure 209. — Striped bass {Roccus saxatilis), Chesapeake Bay region. From Goode. Drawing by H. L. Todd.

390

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

and is almost as long as the second dorsal, also
(less obvious) that its maxillary (upper jaw) bones
are not sheathed by the preorbital bone, separate
it from all the weakfish tribe (p. 417). Nor is there
any danger of confusing it with the sea bass,
cunner, tautog, or rosefish, for its two dorsal fins
are entirely separate whereas in all these the
spiny and soft-rayed parts are continuous, as a
single fin. The white perch comes closest to it
in general appearance but the two dorsal fins
of the perch have no free space between them
(p. 405), and its fin spines are stiffer.

The trunk of the striped bass is 3% to 4 times
as long (to base of caudal fin) as it is deep, thick
through, its back hardly arched. It has a moder-
ately stout caudal peduncle, a long head (almost
as long as the fish is deep), two spines on the
margin of each gill cover, an oblique mouth gaping
back to the eye, a moderately pointed nose, and a
projecting lower jaw. Young fish are more
slender than old. The two dorsal fins are of about
equal lengths; the first (9 or 10 stiff spines) tri-
angular in outline, originating over the middle
of the pectorals; the second (12 or 13 soft rays)
regularly graduated in height from front to rear,
and separated from the first by a distinct (though
short) space. The anal (about 11 rays preceded
by 3 spines) is of about the same size and form as
the second dorsal, and originates below the middle
of the latter. The caudal is moderately wide and
only slightly forked. The pectorals and ventrals
are of moderate size, the latter somewhat behind
the former.

Color. — Dark olive green varying to bluish
above, paling on the sides, and silvery on the
belly, sometimes with brassy reflections. The
sides are barred with 7 or 8 narrow, sooty, longi-
tudinal stripes, which follow as many rows of
scales and which may be variously interrupted.
The highest stripe is the most distinct, and all of
them but the lowest are above the level of the
pectoral fins. The dorsal, caudal, and anal fins
are somewhat dusky.

Size. — The bass grows to a great size, the
heaviest of which we have found definite record
being several of about 125 pounds that were taken
at Edenton, N. C, in April 189 1.24 One of 112
pounds, which must have been at least 6 feet long,
was caught at Orleans, Mass., many years ago.

One of 100}£ pounds is said to have been taken in
Casco Bay, Maine,26 and fish of 50 to 60 pounds are
not exceptional. Usually bass, as caught, weigh
from 3 to 35 or 40 pounds; the average weight of
ones recorded in the register of the former Glades
Hotel 26 at Scituate, Mass., during the period 1854
to 1858, was about 27 pounds.

Bass weigh about % pound when 12 to 13 inches
long; about 2% to 3 pounds at 18 to 20 inches;
about 5 pounds at 24 inches; about 10-15 pounds
at 30-32 inches; and about 18-20 pounds at 33-36
inches. Twenty-pound bass average about 36
inches in length; 30 pounders about 43 inches;
40 pounders about 47 to 48 inches.27 On the
Pacific coast 50 pounders run about 50 to 51
inches,28 and the relationship between weight and
length runs about the same for very large fish on
the Atlantic coast. The record fish caught on rod
and reel was one of 73 pounds, taken in Vineyard
Sound in August 1913 by C. B. Church.

Females grow larger than males; probably most
bass of 30 pounds and heavier are females.29
Thus the common use of the term "bulls" for the
very large ones might better be replaced by "cows."

Habits.30 — Stripers are powerful fish; so strong
in fact, that they appear to have no difficulty in
handling themselves in the surf, where one is some-
times seen actually in the translucent crest of a
comber just before the latter breaks. But this is
not a very swift fish as compared with the mack-
erel tribe. Bass often swirl conspicuously at the
surface or splash in pursuit of bait fish. They
sometimes roll as the little northern porpoise or
puffing pig (Phocaena) does. And we have heard
of them finning (i. e., with dorsal and tail fins
showing).31 But we have never seen or heard of
one leaping clear of the water as tuna and bonito
so often do unless hooked in shoal water.

During the first two years they live mostly in
small groups. Later they are likely to congregate
in larger schools; this applies especially to those up

" Smith, North Carolina Geol. and Econ. Surv., vol. 2, 1907, p. 271.

" Atkins, Fish. Ind. U. S., Sect. 5, vol. 1, 1887, p. 694j
•* Kindly lent by John Adams.

27 For a detailed tabulation of the length-weight relationship for bass from
H pound to 47M pounds, see Merriman, Fish. Bull. No. 35, 1941, U. S. Fish
and Wildlife Service, p. 7, vol. 50, 1950, pp. 1-77.

"•* As scaled from a graph given by Scofleld, California Fish and Game, vol.
18, 1932, pp. 168-170, fig. 38.

28 Definite information in this regard is scant.

s° Interesting recent studies of the striped bass are by Pearson (Bull. IT. S.
Bur. Fish., vol. 49, 1938, pp. 825-851) and by Merriman (Fishery Bull. 35,
U. S. Fish and Wildlife Service, 1941, 77 pp.).

» Frank Mather of the Woods Hole Oceanographic Institution reports an
instance of this.

FISHES OF THE GULF OF MAINE

391

to 10 pounds or so, which are often spoken of as
"school fish." The larger ones often school, but
the very largest, of 30 to 40 pounds and upward,
are more often found single or a few together.
They are most likely to be in schools while mi-
grating, but more scattered while feeding in one
general locality.

Small fish (2 and 3 years old) in particular,
tend to school densely; also they travel consider-
able distances without scattering but, as Merriman
emphasizes,32 it is not likely that a given school
holds together for any long period, for fish of
various sizes (i. e., ages) up to the very large ones
often school together, showing that different ages
intermingle more or less. Mixed schools running
from 8 or 10 pounds to 30 or 40 pounds were
reported repeatedly in 1950, for example.

The bass is very voracious, feeding on smaller
fishes of whatever kind may be available, and on a
wide variety of invertebrates. Lists of its stomach
contents for one locality or another include alewife,
anchovy, croakers, channel bass, eels, flounders,
herring, menhaden, mummichogs, mullet, rock
eels (Pholis gunnellus), launce, sculpins, shad,
silver hake, silversides, smelt, tomcod, weakfish,
white perch, lobsters, crabs of various kinds,
shrimps, isopods, gammarid crustaceans, various
worms, squid, soft clams (Myra) and small mussels.
In our Gulf the larger bass prey chiefly on herring,
smelt, sand launce, eels, and silver hake, on squid
(on which they gorge when they have the oppor-
tunity) , on crabs large and small, on lobsters, and
on sea worms (Nereis) ; while small ones are said
to feed to a considerable extent on gammarid
crustaceans and on shrimps.

When bass are gorging on any one particular
prey it is common knowledge among fishermen
that they are likely to ignore food of other sorts
for the time being. It seems also that when prey
is plentiful, bass are likely to gorge, then cease
feeding to digest, then to gorge again; also that
all the members of a given school are likely to do
this in unison, with consequent annoyance to the
angler.

Bass, too, seem on the whole to be more active,
and especially to feed more actively, between
sunset and sunrise than while the sun is high.
In estuarine situations this fits with the habits
of their prey, for it is by night that the sea worms
(Nereis) that are the chief item in their diet there

*> Fish. Bull. 35, U. S. Fish and Wildlife Service, 1841, vol. 60, p. 43.

emerge from their burrows to swim about. And
bass fishing is often much more productive by
night than by day off the open coast also, though
schools of bait fish are seen at all hours (else the
terns would starve), while the time when crabs,
etc., are most likely to be stirred up by the surf,
and are most easily caught around the rocks,
depends on the stage of the tide, not on the hour
of the day. So most fishermen (ourselves in-
cluded) believe that it is inherent in the nature of
the larger sized bass to avoid strong sunlight by
sinking to the bottom. A familiar instance is
the regularity with which they desert the surface
soon after sunrise on bright summer days at
places where large numbers are caught by trolling
during the hour or two after daybreak; the
eastern side of Cape Cod Bay is a local example.

It has been discovered recently that trolling
deep with wire lines is often productive, irrespec-
tive of the time of day, at times and places where
bass "show" only during the early morning hours.
This habit, however, is not so deeply engrained
but that schools of bass often rise to the surface
in pursuit of bait fish at any time of day, or come
within easy casting distance of the beach. We
recall seeing several schools of good-sized fish
(those that we landed ran up to 23 pounds)
suddenly splashing all around our boat about
midday, on one occasion off Wellfleet, in Cape Cod
Bay, though it was only for a few hours after
sunrise that the several boats fishing regularly
there had taken any by top-water trolling for
some time previous.

The best advice we can give the surf-caster, in
this regard, is to go fishing whatever time of the
day he is free to do so.

The striper is so strictly an inshore fish that we
have never heard of large catches being made, or
schools seen, more than 4 or 5 miles from the
nearest point of land,33 though the migrating
schools doubtless pass much farther out in crossing
the mouths of the larger indentations of the coast,
such as Delaware Bay and Long Island Sound.
And a few fish may stray far offshore in winter, for
one about 18 inches long was taken in an otter
trawl about 60 miles south of Marthas Vineyard,
in 70 fathoms of water, in February 1949 (p. 400) .3*

" Henry Lyman informs us that bass are caught in numbeis late in the
autumn in the rips east of Nantucket about 4 miles out, but that verbal
reports of some taken during the summer of 1950 on the offshore part of Georges
Bank were actually based on two weakfish (p. 419).

« Reported to us by Capt. Henry W. Klimm of the dragger Eugene H.

392

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

On the landward side, many bass come within
easy casting range of the shore ; we have had a fair
sized one strike our plug not 4 feet from the rock
from which we were casting on the Cohasset
shore. Many (especially the smaller sizes, but
large ones also) run up into estuaries and into
river mouths. In some rivers, good numbers
(large as well as small) are caught so far upstream
as to make it likely that they remain there the
year round. This is notably the case in the
Alabama River system where (we hear) 250 to 300
bass ranging from 5 to 40 pounds were caught near
Tallasseem some 30 miles above Montgomery,
which is at least 300 miles from salt water, follow-
ing the river.36 They are also known to spawn
some 250 miles up the Sacramento River in Cali-
fornia. It would be interesting to know what
proportion of the bass that spawn at Weldon,
N. C, 100 miles or so up the Roanoke, and that
run 60 to 90 miles up the St. John, in New Bruns-
wick,39 ever see salt water. Bass also run up the
Hudson for about 160 miles to Albany.

The great majority of the total population of
bass frequent the coast line, except at breeding
season. Among these, the smaller sizes, up to 15
pounds or so, are found indifferently within en-
closed bays, in small marsh estuaries, in the
mouths of rivers and off the open coast. But we do
not often hear of fish heavier than 20 to 25
pounds caught in situations of these sorts. And
the great majority of the large bass, of 30 pounds
or more, hold to the open coast, except at spawning
time (p. 394), and perhaps in winter (p. 400). But
this is not an invariable rule ; we are familiar with
one narrow inlet where tides run strong, and where
some lucky angler catches a very large bass now
and then (p. 396).

Bass off the open coast are most likely to be
found along sandy beaches, in shallow bays, along
rocky stretches, over and among submerged or
partially submerged rocks and boulders, and at the
mouths of estuaries, the precise situations that
they occupy being governed by the availability of
food. Off the outer beaches they may be any-
where right to the breakers. When they are
close in they frequent the troughs that are

w Information from Henry Lyman, from an angling correspondent in
Alabama. They have long been known up the Alabama as far as Mont-
gomery (Pearson, Bull. U. S. Bur. Fish., vol. 49, 1938, p. 826).

» According to Adams (Field and Forest Rambles, 1873, Pt. 3, Fishes, pp.
248-249), who has given an interesting and readable account of the bass in the
river.

hollowed out by the surf behind off-lying bars, also
the gullies through which the water rushes in and
out across the bars as the rollers break, for it is in
such situations that bait fish are easiest caught,
and that crabs, worms, and clams are most likely
to be tossed about in the wash of the breakers.
When the tide is high, bass often lie on a bar, or
even in the white water along the beach if there is
a good surf running. When the tide falls they
drop down into the troughs or move farther out,
according to the precise topography. In either
case, every surf fisherman knows that his chances
are much better when the sea is breaking at least
moderately heavy so that he can cast into white
water, than when it is smooth.

They also he under rafts of floating rockweed
at times, probably to prey on the smaU animals
they find among the weeds.

The best spots along rocky shores are in the
surf generally, and in the wash of breaking waves
behind offlying boulders and among them, or
where a tidal current flows most swiftly past some
jutting point. In the mouths of estuaries they
are apt to hold to the side where the current is the
strongest, and in the breakers out along the bar on
that side. In shallow bays, they often pursue small
fry among the submerged sedge grass when the
tide is high, dropping back into the deeper chan-
nels on the ebb. And they frequent mussel beds,
both in enclosed waters and on shoal grounds out-
side, probably because these are likely to harbor
an abundance of sea worms (Nereis).

When bass are schooling outside they are
likely to be moving along the coast in the one
direction or in the other. But they may remain
in the same general locality for weeks, or through
the summer. Thus a body of very large fish, of
25 to 50 pounds, stayed close in to the outer
beach near the tip of Cape Cod, through most of
July of 1951 and into that August, yielding
consistent catches to the more skillful surf-
fishermen.

Bass are active over a temperature range from
perhaps 70° down to about 43°-46° F. Present
indications are that if the temperature falls lower
they either withdraw to somewhat warmer water
if off the outer coast, or lie on the bottom in a
more or less sluggish state if they are in some
estuary. On the other hand it is not likely that
they can long survive temperatures higher than
about 77°-80°, for many were found dead in

FISHES OF THE GULF OF MAINE

393

shallow estuaries in Connecticut and in Massa-
chusetts during the abnormally hot August of
1937.37 They are equally at home in fresh or
slightly brackish water, and in coastal salinities
of 3.1 to 3.3 percent. But their usual wanderings
do not take them out into waters of full oceanic
salinities (3.5 percent or higher).

Migrations. — No phase of the life history of
the bass arouses as much discussion among fisher-
men as their migrations. And the picture still
remains so puzzling that we dare not attempt
anything more than a brief summary of what has
been learned to date.

It seems certain that stripers do not ordinarily
travel far until they are 2 years old. Thus the
young fish from the enormous year classes of 1934
and 1942 — apparently produced in the Chesapeake
Bay-Delaware Bay region chiefly — did not appear
in New England waters until 2 years later. But
the fact that they did appear there and in the
Gulf of Maine in hordes in the summers of 1936
and 1944 shows that a bass is capable of very
extensive journeys, once it has reached its third
year.

It has long been known, too, that the pound
nets on Long Island and along southern New
England ordinarily make large catches only in
the spring (peak in May), and again from early
October into November;38 also that large spring
catches are made progressively later in the season,
proceeding from south to north, the reverse being
true in the autumn. This, of course, suggests
that part at least of the bass population follows
the shore line northward and eastward as far as
southern New England in spring, to return
westward and southward in autumn. And this
is verified for bass 2 and 3 years old by the
returns from tagging experiments by Merriman
at the eastern end of Long Island and in Con-
necticut during the years 1936 to 1938,39 for
recaptures of fish that had been tagged there in
May came mostly from farther east along southern
New England, one from Cape Cod Bay, and
another from Cohasset on the southern shore of

» Merriman, Fishery Bulletin No. 35, TJ. S. Fish and Wildlife Service,
1941, vol. 50, p. 43.

« See Merriman, Fish. Bull. No. 35, U. S. Fish and Wildlife Service, 1941,
vol. 60, pp. 33, 34, fig. 24, for details.

w For details we refer the reader to Merriman's original account (Fish. Bull
No. 35, Fish and Wildlife Service, 1941, vol. 50, pp. 36-42, figs. 26-29; also
pp. 71-73, tables 17-20), which is the most authoritative discussion of the
subject that has appeared yet.

the inner part of Massachusetts Bay. But the
recaptures from fish tagged in summer were
mostly from nearby (evidence of a stationary
population), while those for autumn-tagged fish
were scattered along the coast from the eastern
end of Long Island to Chesapeake Bay, with one
from Croatan Sound, one from Albemarle Sound
(Stumpy Point), and one from Pamlico Sound in
North Carolina.

But the picture is by no means so simple as the
foregoing might suggest. To begin with, no evi-
dence is available as to the movements of large
bass, other than the successive dates when they
appear or disappear off different parts of the
coast.40 And it is no less true of bass than it is of
mackerel (p. 330), that successive appearances and
disappearances from place to place are not con-
clusive evidence of along shore migration. Yet
it is now certain that while some bodies of bass
carry out extensive migrations north and east in
spring, west and south in autumn, other bodies
do not. Thus, as Merriman points out,41 the
bass of the northeastern shore of the Gulf of
Mexico are completely isolated, while those of the
Atlantic coast south of Cape Hatteras form an-
other separate population, few of which (if any)
ever spread farther north. The bass of the Gulf
of St. Lawrence and of the lower St. Lawrence
River appear to be wholly isolated also. And
while some interchange may take place between
the populations found in various bays and rivers
around the outer coast of Nova Scotia, it is
doubtful whether these have any regularly
migratory association, either with the Gulf of
St. Lawrence fish or with those of more southern
waters, except in occasional years (p. 398).

Chesapeake Bay, however, harbors both migra-
tory bass,42 as proved by tagging experiments
(p. 393) and other evidence (p. 393), and nonmigra-
tory as proved by the fact that fish of all sizes are
taken there both in summer and in winter, though
not so many of them as in spring and fall. Sim-
ilarly, some bass winter in northern waters though
most of the fish appear to be migrants there; and
perhaps a considerable percentage do so in the
lower reaches of the Hudson River estuary.

« The few returns so far from bass of 5 pounds and upward that have beea
tagged have been from nearby, and soon after they were released.
" Fish. Bull. No. 35, U. S. Fish and Wildlife Service, 1941, vol. 50, p. 42.
« Using this term to mean extensive seasonal journeys.

394

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Merriman 43 has suggested that these northern
wintering fish may be "of two types — the individ-
uals that form the resident more or less isolated
population" and others "that may have had their
origin farther south but spend an occasional winter
in northern waters." It may prove that a good
proportion of these bass that come from the south
when they are 3-^1 years old may remain in the
north for the rest of their lives. And there is no
way for the fisherman to tell in which of these
categories the bass belong, that he lands. The
reader will find some further discussion of migra-
tions in connection with the status of the bass in
the Gulf of Maine (p. 395). We need only add that
the existence of these nonmigratory populations
and the fact that the Pacific coast bass are simi-
larly stationary, are sufficient proof that seasonal
migration is not an essential incident in the life
of the striper.

Bass spawn either in brackish water at the heads
of estuaries*4 (the Hudson, for example) or in
fresh rivers, never off the open coast in salt water
so far as is known. Those that enter fresh rivers
may deposit their eggs only a short distance above
the head of tide as they do in the Potomac, or
they may run much farther upstream. But we
have yet to learn how large a percentage of the
bass that are known to spawn 100 miles up the
Roanoke, near Weldon, N. C. (a major spawning
ground), or still farther up the Alabama,46 and up
the Sacramento River in California, have come
from salt water (p. 392).

The chief requirement for successful spawning
is (it seems) a current turbulent enough to prevent
the eggs from settling on bottom where they would
be in danger of being silted over and smothered.

The spawning season is from late April to early
May in North Carolina; in May, chiefly, in the
Chesapeake Bay region; perhaps equally early in
the waters of New York.46 Any bass that may
spawn in the rivers of Massachusetts, of Maine,
and of the Bay of Fundy, probably do so in June;
those of the southern shore of the Gulf of St.

« Fish. Bull. No. 35, U. S. Fish and Wildlife Service, 1941, vol. 50, p. 42.

<< See Merriman, Fishery Bulletin No. 35, U. S. Fish and Wildlife Service,
1941. vol. 50, p 17, for precise salinites in which bass in their first summer have
been taken in the Hudson River, and in the Parker River, Massachusetts.
See Tresselt (Bull. Bingham Oceanogr. Coll., vol. 14, art. 1, pp. 98-110, 1952)
for a survey of spawning grounds tributary to Chesapeake Bay.

« Pearson (Bull. U. S. Bur. Fish., vol. 49, 1938, p. 829) records a female with
eggs from the Alabama River near Montgomery.

« Greeley (New York Conserv. Dept., Biol. Surv. Lower Hudson Water-
shed, 1937, p. 100) concludes that the spawning season in the Hudson "in-
cludes May."

Lawrence and of the lower St. Lawrence River in
June and July.

A large female during spawning may be sur-
rounded by many small males, and the latter are
described as fighting fiercely47 with one another.

Females stripped at the Weldon, N. C. Hatchery
yielded from 11,000 to 1,215,000 eggs each, during
the period 1928 to 1938, with one of 4K pounds
yielding 265,000. Thus the oft-quoted estimate
of 10 million fish for a really large one is within
reason.48

The eggs average 1.1-1.35 mm. in diameter when
they are deposited in the water, but the pervitel-
line membrane swells during the first hours after
fertilization to an average diameter of about 3.6
mm. They have a large oil globule and are semi-
buoyant; that is, they sink in quiet water, but are
swept up from the bottom by the slightest dis-
turbance, so that they tend to drift downstream
with the current. Consequently the eggs that are
produced far upstream may not hatch until they
have reached tidewater. The eggs are reported
as hatching in about 70 to 74 hours at a temper-
ature of 58-60°; in about 48 hours at 67°; in about
30 hours at 71-72°.

In Chesapeake Bay, the young fry of the year
are about 1}6 inches (30 mm.) long by June; 1% to
2Yi2 inches (45-53 mm.) long in July; 2 to 2% inches
(50-70 mm.) in August; and 3% to 8% inches by
the following April and May; i. e., at the end of
their first year.49 According to Merrimaa,50 most
of the fry of the year taken in the Hudson River
during their first summer are between about 1%
inches (40 mm.) and about 3K inches (90 mm.)
long; a few seined in the Parker River, Newbury,
Mass., were from about 2% inches (71 mm.) to
about 3% inches (85 mm.) long. And this last is
perhaps representative for whatever bass may now
be produced in Gulf of Maine rivers, for we read
that great numbers of fry of 2 to 3 inches were
taken of old in winter in the rivers of Maine in
bagnets set for smelt and tomcod.61

Two-year-old bass taken in Connecticut aver-
aged 1 1 to 1 lji inches (28 or 29 cm.) long in spring,

« See Smith, North Carolina Oeol. Econ. Survey, vol. 2, 1907, p. 272, for
an eyewitness account by S. G. Worth.

i* Merriman (Fish. Bull. No. 35, TJ. S. Fish and Wildlife Service, 1941,
vol. 50, p. 19) gives an excellent summary of information available as to
spawing, characteristics of the eggs, and period of incubation.

« Hildebrand and Schroeder, Bull. U. S. Bur. Fish., vol. 43, Pt. 1, 1928,
pp. 248-249

» Fish. Bull. No. 35, U. S. Fish and Wildlife Service, 1941, vol. 50, p. 17,
fig. 10.

•i Atkins, Fish. Ind. TJ. S., Sect. 5, vol. 1, 1887, p. 693.

FISHES OF THE GULF OF MAINE

395

about 12 inches (30 cm.) in June, and about 14%
inches (37 cm.) in October; the 3-year-olds about
15% inches (40 cm.) in spring and about 18 inches
(46 cm.) in October, while 4-year-olds increased
in length from about 18% inches (48 cm.) to about
20% inches (53 cm.) between spring and autumn,
on the average.62 And the average rate of growth
was about the same for Hudson River fish ex-
amined by Greeley.63 But the rate at which they
grow is governed largely by the food supply.
Bass in captivity have been known to grow from
6 inches long to 20 inches in 11 months, while
some that were kept in a certain pond in Rhode
Island are described as having gained weight from
1 pound in June to 6 pounds in October.64

The later growth rate has not been traced for
our Atlantic bass. But it is generally believed
that the 3 5-50-pound ers that were caught in
considerable numbers in 1950, and are being taken
in 1951, were members of the very successful year
classes of 1940-1942, which fits well with the
growth rate of bass on the Pacific coast, where the
average age is about 7 years for 20-pound fish,
10-11 years for 30-pounders, about 14 years for
40 pounders, and 17 to 18 years for 50-pounders.66

On the Pacific coast females grow faster than
males after the third year, which is probably true
of the Atlantic bass also.66 This certainly is a
long-lived fish for one kept in the New York
Aquarium lived to be 23 years old.67

Merriman M found that "approximately 25 per-
cent of the female striped bass first spawn just
as they are becoming 4 years old, that about 75
percent are mature as they reach 5 years of age,
and that 95 percent have attained maturity by
the time they are 6 years old," among Connecti-
cut fish. But a large percentage of the males
had matured at 2 years, probably nearly all of
them by the time they were 3 years old. And
it is probable that this applies equally to the
Maine bass. Merriman has also made the inter-
esting discovery that only about one-tenth of the
bass of northern waters are males, but that males

* Merriman, Copeia, 1937, p. 23.

*» New York Conscrv. Dept., Biol. Surv. Lower Hudson Watershed, 1937,
p. 62.

» Bean, Bull. New York State Mus., 60, Zool. 9, 1903, p. 527.

" Scaled from Scofleld's graph (California Fish and Game, vol. 18, 1932,
pp. 168-170, fig. 38).

" See Scofield, Fish Bull. No. 29, Div. of Fish and Oame, California,
1931 for growth of bass in California.

« Bull. New York Zool. Soc., vol. 16, No. 60, November 1913, p. 1049.

18 Fish. Bull. No. 35, U. S. Fish and Wildlife Service, 1941, vol. 50, p. 22.

are nearly as numerous as females, southward
from Delaware Bay.

It has been suggested that the striper may not
be a regularly yearly spawner,69 but no positive
evidence is at hand as to this.

General range. — Atlantic coast of eastern North
America, from the lower St. Lawrence River and
the southern side of the Gulf of St. Lawrence
to northern Florida; also along the northern shore
of the Gulf of St. Lawrence to Alabama and
Louisiana; running up into brackish or fresh
water to breed.60 In the last quarter of the 19th
century it was introduced on the Pacific coast,
where its range extends now from Grays Harbor,
Wash.,61 to Los Angeles County, Calif. It is now
a favorite game fish there, and the yearly commer-
cial catch since World War I ran between 500,000
and about 1,000,000 pounds there, until 1935,
wben commercial fishing for stripers was pro-
hibited by the State of California.

Occurrence in the Gulf of Maine. — The range of
the striper includes the coastline of our Gulf from
Cape Cod to western Nova Scotia. But its dis-
tribution there in detail is determined by its very
evident preference for surf-swept beaches and for
particular stretches of rocky or bouldery shoreline;
also for shallow bays, inlets, and estuaries. The
geographic status of bass in our Gulf also depends
on whether it be a good bass year (or run of years)
or a poor one.

When bass are reasonably plentiful, as they have
been during the past 15 years, and with a good
representation of fish of different ages, the outer
shore of Cape Cod provides the most productive
surf casting, with Monomoy Island, the general
vicinity of Nauset Inlet, and the tip of the Cape
northward from Highland Light perhaps the
warmest stretches, in most years. But the topog-
raphy of a beach may be altered to such an extent
during severe storms that a stretch that is good
bass water one summer may be poor the next.
Nauset beach is an example, for very few bass
have been caught or seen there during the present
summer (1951), though this has been one of the
most productive localities on the Cape during the
past few summers. Considerable numbers, mostly

'• Merriman, Fish. Bull. No. 35, U. S. Fish and Wildlife Service, 1941,
vol. 60, p. 16.

» Pearson (Bull. TJ. S. Bur. Fish., vol. 49, 1938, p. 827, fig. 1) charts its
United States range, but does not include its Canadian range.

" Oerlach (Contrlb. 14, Oregon Fish. Comm., 1950) gives an interesting
account of the life history and distribution of the striper in Oregon waters.

396

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

of the smaller sizes, are caught in Pleasant Bay
too, within Nauset Marsh, and in Town Cove,
Orleans.

Considerable catches are made by boats trolling
outside the surf, also, or by casting in toward the
breakers along the outer Cape Cod shore, when
the weather permits. But the most productive
and reliable trolling grounds are along the eastern
and southern sides of Cape Cod Bay in most
summers, especially off the Eastham shore a few
miles southward from Wellfleet, and off the mouth
of Scorton Creek, Barnstable and the Sandwich
shore.62 The shores of Cape Cod and Cape Cod
Bay have, in fact, been the chief center of abund-
ance for bass within the Gulf from as far back as
the record runs. Few bass are reported along the
rocky stretch from the Cape Cod Canal to the
entrance to Plymouth Harbor, though this would
seem to be very good bass water, and schools
must pass by. But many are caught in Plymouth
Harbor, especially off Eel Creek, also up Duxbury
Bay to the salt marsh creeks that open into
its head.

Surf casters account for some along Duxbury
Beach on the outside, for a few also in the boulder-
strewn area at the western end of Humarock
Beach.63 The North and South Rivers in Marsh-
field yield considerable numbers in good years;
we have seen and taken good fish there. Anglers,
casting from the shore, take a few (never any
great number) on boulder-strewn stretches along
the Scituate shore, while Glades Point was famous
for large bass in earlier periods of abundance
(p. 390), when it was common practice to chum
the fish by throwing out chopped lobsters, a
method never likely to be revived because lobsters
are far too costly nowadays. The Cohasset
shoreline (with which we are familiar) yields a few
yearly (mostly caught between sunset and sun-
rise), occasionally a very large one. In seasons
when there is a good run of the smaller sizes,
considerable numbers are taken at various places
within the limits of Boston Harbor; Hull Gut,
Weir River in Hingham, and Wollaston Beach are
well known localities. And in years when there is
a run of little fish, many of them are caught from

« Many are caught by anglers casting in the Cape Cod Canal, but this is
not properly a part of the Gulf of Maine.

•3 The bouldery area at the eastern end at the North River inlet is now
within the limits of the military reservation; hence the only way to fish it is
from a boat by casting in, toward the rocks.

the docks and from the bridges, to the head of
Boston Harbor.

The north shore of Massachusetts Bay seems
not to be as attractive for bass as its succession
of inlets, beaches, and rocky headlands might
suggest, for catches reported are small and
scattering in most summers. But the beaches and
enclosed waters from a few miles north of Cape
Ann to and including the mouth of the Merrimac
River are productive enough to rank second to the
Cape Cod-Cape Cod Bay region . Bass are taken in
the surf from Ipswich Beach, Cranes Beach, and
along the entire length of Plum Island Beach;
many more are caught by boat fishermen over the
flats within the mouth of the Merrimac, as well as
about the jetties at its entrance. Schools are
often reported in Plum Island Sound. And the
Parker River, emptying into the latter, is not only
well known water for bass, especially small fish,
but it holds some bass over the winter (p. 400), and
it is one of the few streams along the New England
shores of our Gulf where very young bass have
been taken within recent years (p. 398).

Some are caught in Hampton Harbor, N. H.
But the next important bass waters (moving
northward) are the lower reaches of the Piscataqua
River system, marking the boundary between
Maine and New Hampshire; a good number, large
and small, are now caught there yearly. In good
years bass are to be caught in several of the
streams that drain the southern part of the Maine
coast, especially in the York, the Mousam, and in
the Saco which is the most productive. Schools
are sighted and a few are caught along the inter-
vening beaches and some in the shallows of
Biddeford Pool.

Information as to the status of bass for the
coastline and streams of northern and eastern
Maine, past or present, is scant, and we have come
to suspect that bass may never have been as
plentiful there as was supposed. A few are caught
here and there around Casco Bay in good years,
product perhaps of the Kennebec. But the esti-
mated catch in the Kennebec was only about
12,760 pounds as far back as 1880; 64 and there have
been far fewer bass there of late years. Our most
recent information is that schools of large fish
were seen in the lower Kennebec, off Popham and
Reed Beaches in early October 1950 with some

« Atkins, Fish. Ind. U. S., Sect. 5, vol. 1, 1887, p. 675.

FISHES OF THE GULF OF MAINE

397

caught up to 26 pounds, and that a few were being
taken daily, in late June 1951.65 Nearly as many
were taken in the Sheepscot, formerly, as in the
Kennebec; the present condition is not known.68
There were bass in the St. George during the period
1936-1940; doubtless there are some there still,
for we heard of some in the surf near Georgetown,
Maine, in August 1951.

Bass are seen in most years in Bangor Pool at
the head of the estuary of the Penobscot, where
some are caught by anglers casting especially for
them, also by salmon fishermen. And many in
the 2- to 4-pound class were reported and caught
in the Belfast River and in Searsport Harbor
farther down Penobscot Bay in 1938. But there
have not been enough of them there during the
past few years to have caused special comment.
Stripers were seen in the tide rips in the narrows
between Mount Desert Island and the mainland
(near the Hancock-Sullivan Bridge) in August
1951, and others were reported driving squid
ashore near Winter Harbor, Maine, a few miles
farther east. Salmon fishermen sometimes "rise"
bass in the Narraguagus, and Atkins 67 speaks of
"a very few" in the St. Croix, though Huntsman
found no recent record of bass in the Passama-
quoddy region.

There may be an occasional bass in Maine rivers
other than those we have mentioned, but there is
nothing in the past record to suggest that there
ever were many. In 1880, for example, the re-
ported catch was nearly as great for the Kennebec
(about 13,000 pounds) as for all the other rivers
and coast of Maine combined (about 15,000
pounds). And there is no reason to suppose that
the regional contrast has altered subsequently in
this respect.68 In the Bay of Fundy region, bass,
as Huntsman has pointed out,69 are confined to
the large warm estuaries and the neighboring
fresh water; i. e., to those of the St. John, Minas
Basin-Cobequid Bay and Shubenacadie River
systems, and of the Annapolis.

Available information suggests that bass always
were more plentiful in St. John River waters than

anywhere along the eastern part of the coast of
Maine, and that they are still. Bass are occa-
sionally caught in St. John Harbor, mostly between
April and June.70 And while they were reported
as already much less numerous in St. John waters
by 1884 than they had been in earlier times,71
there still are enough of them in the St. John and
its tributaries to have yielded commercial catches
of 12,200 pounds in 1944, and 7,400 pounds in
1946. The most recent news that has reached us
from the St. John is that salmon fishermen saw a
school at the surface and caught some that weighed
3 to 11 pounds in late June or early July of 1951. 72

Bass are well known in the Minas Basin-
Cobequid region. According to local fishermen,73
as many as 80 fish are sometimes taken in weirs
there in a day, most of them in the 8- to 10-pound
category, but with occasional fish reported up to
33 pounds.

The status of the bass is especially interesting
in the Shubenacadie River, for they are not only
caught in fresh water there and in Shubenacadie
Lake where they are known to spawn, but some
large fish remain throughout the year in the lake;
i.e., they behave like a land-locked population.74
A thousand or so, in fact, are caught yearly by
anglers in the Lake and in the Shubenacadie
River; 76 and it is said that fish as large as 50 pounds
have been taken,76 though most of them run small
there.

We are informed 77 that the catch by anglers
was about 620 bass (average about 4% pounds)
in the Bass River, tributary to Cobequid Bay in
1950, and that the catches for 1949 and 1950
combined were about 1,350 fish (average about
2% pounds) in the Gaspereau, tributary to Minas
basin; 4,650 fish (average about 5% pounds) in the
Annapolis River; and about 125 fish (average
6 pounds) in the Bear River, tributary to Digby
basin, in 1950. It is interesting, that these fish
ran so small, for the bass caught in Cape Cod and
northern Massachusetts waters during these same

** Reported in Saltwater Sportsman for October 6, 1950.

M Yearly catch about 1880, some 8,000 pounds in the Sheepscot according
to Atkins, Fish. Ind. U. S., Sect. 5, vol. 1, 18S7, p. 716.

" Atkins (Fish. Ind. U. S., Sect. 6, vol. 1, 1887, p. 700) reports one of 20
pounds, taken in the St. Croix in a weir in 1880.

M What few bass were reported from Maine in 1919 were from the Kennebec
(592 pounds) and from Penobscot waters (57 pounds); bass have not been
Included in the fisheries statistics for Maine for any subsequent years.

« Contrib. Canadian Biol. (1921) 1922, p. 63.

70 Information from Dr. A. H. Leim.

» Ooode, Fish. Ind. U. 8., Sect. 1, 1884, p. 425.

'• Saltwater Sportsman for July 6, 1951.

» According to Moore, Boston Herald, August 28, 1950.

" Information from Dr. A. H. Leim.

'• According to Huntsman, Ann. Rept. Fishery Board Canada, (1949)
1950, p. 41.

'• Vladykov and McKenzie, Proc. Nova Scotia Inst. Sci., vol. 19, 1935,
p. 91.

" Information from MaJ. Howard Scott of the Fishery Division of the
Nova Scotian Department of Trade and Industry, received through Henry
Lyman.

398

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

years included a good number of very large fish
(p. 403).

Anglers have also come to realize recently that
bass are to be caught in various bays and river
mouths along the western shoreline of Nova
Scotia. But no definite information has reached
us as to how plentiful they are there, or how large.

The regional contrasts in the abundance of bass
along different sectors of the coastline of our
Gulf may be illustrated more concretely by the
commercial landings for 1945. 78

Outer Cape Cod and Cape Cod Bay7' —

perhaps about 57,000 lbs.

Cape Cod Canal to New Hampshire line 51, 100 lbs.80

New Hampshire 9,000 lbs.

Maine None reported.

St. John River system, New Brunswick 2,400 lbs.

Minas Basin, Cobequid Bay and Shubena-

cadie River region, Nova Scotia 13,800 lbs.

Annapolis County, Nova Scotia 3,100 lbs.

West coast of Nova Scotia

800 pounds in 1944; none reported in 1946.

A regional contrast of another sort, of interest
to anglers, is that really large bass of (say) 30
pounds and upwards, are far more plentiful along
the Massacbusetts coast (especially in Cape Cod
waters) than they are anywhere farther north and
east in our Gulf.

Localities along the outer coast of Nova Scotia
where we have heard (or read) of stripers are the
head of Mahone Bay; head of Chedabucto Bay;
and Mira Bay and other harbors of Cape Breton.
The numbers caught there are so small that they
are not included in the published statistics of the
commercial catches for the counties in question.
The shoal estuaries, however, of the Richibucto
Bay region and also the estuary of the Miramichi
River (on the southern shore of the Gulf of St.
Lawrence) harbor isolated populations of bass
plentiful enough to have yielded commercial
catches of about 3,800 pounds and 9,000 pounds,
respectively, in 1929, 4,100 and 3,000 pounds in
1931.81 And there is also a population (or popu-
lations) below Quebec in the lower St. Lawrence
River, of bass that winter in that same general
region, as proved by marking experiments recently

" The most recent year for which detailed statistics are readily available-

n Assuming that about $i of the Barnstable County catch of 86,200 pounds
was taken along the outer shore of Cape Cod and in Cape Cod Bay (prob-
ably an underestimate).

u Assuming that about H of the Plymouth County catch of 75,000 pounds
was taken on the Massachusetts Bay side.

" This is the most recent year for which information is available for North-
umberland and Kent Counties.

carried out by Vladykov.82 There are enough of
them, in fact, around Isle d'Orleans for bass fishing
to be a favorite sport there. But the commercial
catches are so small as to suggest that the stock of
bass is not very large.83

It has been known for many years that bass
spawn in the St. John River,84 and it is probable
that they also spawn in the small streams trib-
utary to Minas Basin and Cobequid Bay at the
head of the Bay of Fundy; in Grand Lake at the
head of the Shubenacadie River, and probably in
the Annapolis River. It is generally believed,
also, that some bass spawned of old in all the
larger rivers from the Penobscot westward.
Great numbers, so small (2-3 inches) as evidently
to be fry of the year, were caught, for example,
in winter in the 1880's in the Kennebec, where
ripe fish also have been reported85 from the end
of June into July. But the only Maine or Mas-
sachusetts streams where we find evidence of
spawning bass in recent years are the Mousam,
in Maine, where fishermen have reported taking
females with ripe eggs on several occasions;89
and the Parker, in Massachusetts, where Merri-
man87 took three fry of the year 2% to 3% inches
(7.1-8.5 cm. long) on August 4, 1937. Thus it
seems sufficiently established that a great majority
of the bass that summer in the western side of
our Gulf come from spawning grounds to the west
and south.

Merriman's88 painstaking investigations show
beyond reasonable doubt that most of the little
bass of 2 to 5 pounds that appeared in great num-
bers along southern New England and to the
northward in 1936 (p. 402), following a period
of great scarcity of bass there, had been hatched
two years previously (1934) in the region of
Chesapeake Bay, perhaps some of them in the
Delaware Bay region. Some of the abundant
year classes of 1940 and 1942, which appeared in
our Gulf in 1942, and 1944, also may have come

" Rapp. Gen. Minlstr. Chasse et Pecb., Quebec, Pecheries (1945-1947)
1947, p. 50.

M The reported catch for 1948-1949 was only about 1,800 -1,900 pounds (17
quintals; See Rapp. Gen. Ministr. Chasse et Pech., Quebec, Pecheries
(1948-1949) 1949, p. 94).

•« Adams, Field and Forest Rambles. 1873, Pt. 3, Fishes, p. 248.

" Atkins, Fish. Ind. U. S., vol. 5, Sect. 1, 1887, p. 693.

w Towne, State of Maine Striped Bass Survey, Maine Devel. Comm. and
Dept. Sea and Shore Fisheries, 1941 [appro*, date), p. 14.

" Fishery Bulletin No. 35, U. S. Fish and Wildlife Service, 1941, vol. 50,
p. 17.

" Fish. Bull. No. 35, U. S. Fish and Wildlife Service, 1941, pp. 46-52

FISHES OF THE GULF OF MAINE

399

from equally far away; others perhaps from
intermediate spawning areas.

Since the mature bass that visit the coasts of
Cape Cod and northern Massachusetts in such
plenty in good years almost certainly do not
spawn in any numbers in any of the Gulf of Maine
rivers, we caD only suppose that they repair to
more southerly rivers to spawn, perhaps to the
Hudson, in particular. But many of them reach
northern Massachusetts so early in the season,
and so little information is available as to the
condition of their sexual organs when they arrive,
that we still face something of a mystery, here.

In the salt estuaries and open waters of our
Gulf bass are taken only from late spring, through
the summer, and until late in the autumn. In
years when they are plentiful enough to attract
attention, they are likely to be reported about
equally early in the season all along from Cape
Cod to the Merrimac River. In 1950,89 for
example, bass had been reported from the outer
shore of the Cape (Pleasant Bay and Orleans) by
mid-May, from the North and South Rivers, at
Marshneld on the southern side of Massachusetts
Bay, and from the Merrimac at Amesbury by
mid-May; we heard of one caught in Duxbury
Bay as early as May 1 that same year; and in
normally early years they are generally distributed
along the Massachusetts Coast of the Gulf in
May or by the first days of June. The first bass
were reported in and off Hampton Harbor and in
the Piscataqua River about the beginning of the
second week in June (1950), and in Casco Bay
about the middle of the month.

Bass are said to appear as early as the end of
May in Bangor Pool at the head of the estuary of
the Penobscot in some years.90 In 1950 they were
scattered all along Penobscot Bay before the end
of June. And it is probable that the seasonal
schedule is about the same for the bass at the
head of the Bay of Fundy, but information is
scant.81

Once the bass have appeared, they continue in
evidence until well into the autumn (p. 399).
During this part of the year, the bass of the coasts
of Massachusetts and most of those in Maine are
in salt water and in brackish, except for such as
enter fresh water to spawn (p. 398). But they are

89 This is the only year for which we have detailed information.
•° Weston, Field and Stream, March 1932, p. 69.

« Moore (Boston Herald, Aug. 28, 1950) reports that bass are taken in
traps from July on, in the Cobequid Bay region.

caught all summer in fresh water far above the
head of tide in the Shubenacadie in Nova Scotia
(p. 397), 92 also in the Annapolis, and part of the
stock may have here a similar habit in various of
the rivers of Maine, as in the Kennebec, where
they ran up as far as Waterville until they were
prevented by the construction of the dam at
Augusta.93

In rivers where bass winter, they may, of
course, be taken in any month from late autumn
into the spring (p. 400). As autumn approaches the
bass vanish however from the open coast. What
little information we have suggests that most of
them have disappeared along the outer coasts of
Maine by mid-October or the end of that month
in most years. But they may be in evidence in
Maine rivers until later in the autumn, as they
were of old in the Kennebec, where Atkins94
described them as continuing "feeding in weedy
coves until November"; and in the Mousam River
in southern Maine, where fishing is said to have
been good until November during the period
1938-1940, when our Gulf had a spectacular run
of young fish (p. 402).

Farther southward in our Gulf, they may linger
equally late off the open beaches. In 1949, for
example, a set of traps95 located near Province-
town Harbor in 35-45 feet of water, took 3,705
pounds (the only large catch of the year) on
November 3.

In 1950, a late season, Cape Cod Bay eastward
from the Cape Cod Canal was described to us as
"loaded" with bass until the third week in Octo-
ber, fair numbers were still being caught along the
outer shore of Cape Cod at the end of the month,
schools of small fish were reported on November
9, and half a dozen were landed from the surf on
November 18, and one, on December 3.96 Surf
casting is likely to be much more productive along
the outer Cape Cod beaches during 2 weeks or
even 3 weeks of November than it is in July or
August, especially for the smaller fish, and during
the hours of daylight (p. 391).

And the bass in salt water may be in evidence
until equally late in the season in the Minas-

'2 Huntsman, Ann. Kept. Fisheries Res. Board Canada, (1949) 1950, App.
2, pp. 41-42.

•» Atkins, Fish. Ind. TJ. S., Sect. 5, vol. 1, 1887, p. 693.

>< Fish. Ind. U. S., Sect. 5, vol. 1, 1887, p. 693.

•• Property of the Pond Village Cold Storage Co., of North Truro, to whom
we are indebted for much information.

M Reported by Henry Moore, Boston Herald for Dec. 7, 1950.

400

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Cobequid Bay region, at the head of the Bay of
Fundy for fishermen report taking them there
through October and into November.97

The question where the bass that visit the dif-
ferent parts of the coast of the Gulf of Maine spend
their winters still awaits a comprehensive answer.
It has long been known that the Chesapeake Bay
bass winter in the deeper channels near the head,
of the bay as well as in its estuaries, and in the lower
reaches of the rivers, in a more or less inactive state ;
also those of the New Jersey coast run up into
rivers to remain until the following spring, as
described more than a century ago by Mease.98

Knight99 writes too, that as the weather becomes
colder, the bass of the southern side of the Gulf of
St. Lawrence "penetrate into the bays and arms
of the sea and ascend the rivers at some distance,
where they spend the winter resting on the mud in
a half torpid state." The bass also, in Maine
"pass the winter in quiet bays and coves of fresh
water in the rivers," according to Atkins.1 We see
no reason to doubt that the Bay of Fundy bass,
and also those that still frequent the Maine rivers
from the Penobscot westward, still follow this
habit.

It has been known, also, for many years that
some bass winter in the Parker River, in northern
Massachusetts. In fact, some 8,700 pounds were
taken there during the financial depression of 1930
(p. 402). Local fishermen tell us also that a few
bass winter in the deeper parts of the North and
South Rivers in Marshfield, Mass., on the southern
side of Massachusetts Bay, apparently in salt water.
But these and other small streams do not seem
extensive enough to provide wintering grounds for
all the schools of bass that appear in summer be-
tween southern Maine and Boston Harbor in
reasonably good years. Neither is there anything
in the available record to suggest that the Merri-
mac ever was an important wintering ground.
And it is hardly conceivable that the multitude of
bass that sometimes frequent Cape Cod Bay and
the outer shore of the Cape in good bass years can
winter nearby (unless they do so offshore), there
being no large rivers along this section of the coast,
and no local report of bass in winter in the shallow,
partially enclosed bays there, or in the salt marshes.

" Report by Henry Moore, Boston Herald for Aug. 28, 1950.
•» Tians. Litt. Phil. Soc. New York, vol. 1, 1815, pp. 502-504.
•• The River Fisheries of Nova Scotia, 1867, p. 12.
i Fish. Ind. U. S., Sect. 5, vol. 1, 1887, p. 693.

It was generally believed until recently that the
great majority of bass that frequent the Massa-
chusetts coasts of the Gulf (and the Cape Cod
region in particular), and also those that summer
off southern Massachusetts and around the off-
lying islands, move westward along the shore in
autumn: some to contribute to the bodies of fish
that are known to winter in the rivers of Connecti-
cut and in the lower Hudson, and some to journey
perhaps as far as Chesapeake Bay; i. e., to the
region where many of them are hatched. The
capture, however, in 1949, of an 18-inch bass some
60 miles south of Marthas Vineyard in 70 fathoms
of water in February (p. 391) seems to favor the
view, now gaining favor among observant anglers,
that at least a part of the bass of the Cape Cod
region may only move offshore to winter on bottom
well out on the continental shelf in localities where
the otter trawlers do not ordinarily operate, as has
been found of late to be true of the summer floun-
der (p. 268).

If true, this woidd mean that some of the Chesa-
peake-hatched bass that spread northward to
Massachusetts and Maine when 2 or 3 years old
may never return to their home waters. More
definite information in this regard is to be expected
from tagging experiments now in progress.

Periodic fluctuations in abundance. — Nothing re-
garding bass is of greater interest to commercial
fishermen and to anglers than the great fluctua-
tions in its numbers that have taken place in our
Gulf within historic times.

The bass was a familiar fish when New England
and the Maritime Provinces were first colonized,
all along the coast from Cape Cod to the Bay of
Fundy; plentiful and easy to capture, because of
its large size and its habit of coming into the
mouths of streams and creeks; it was also an im-
portant food supply for the early settlers.

Wood,2 for example, tells us that in what is now
a part of Boston Harbor:

The basse is one of the best fishes in the country, and
though men are soon wearied with other fish, yet are they
never with basse. It is a delicate, fine, fat, fast fish, having
a bone in his head which contains a saucerfull of marrow
sweet and good, pleasant to the pallat and wholesome to
the stomach ... Of these fishes some be three and four
foote long, some bigger, some lesser; at some tides a man
may catch a dozen or twenty of these in three hours . . .
When they use to tide in and out of the rivers and creekes
the English at the top of an high water do crosse the creekea

' New Englands Prospect, 1634, p. 37.

FISHES OF THE GULF OF MAINE

401

with long seanes or basse nets, which stop in the fish : and
the water ebbing from them they are left on the dry ground,
sometimes two or three thousand at a set, which are salted
up against winter, or distributed to such as have present
occasion either to spend them in their homes or use them
for their grounds.

He also describes 3 how "shoales of basse have
driven up shoales of mackerel from one end of the
sandie beach to the other," near Salem, and men-
tions them in the Merrimac. In the earliest rec-
ord the chief centers of abundance for them within
the Gulf were Cape Cod Bay and the shore of Cape
Cod; the neighborhood of Boston Bay and harbor;
the vicinity of the Merrimac River; the Kennebec
River and vicinity, and the larger rivers that drain
into the Bay of Fundy. Inexhaustible, however,
though the supply seemed, a decrease was reported
as early as the last half of the eighteenth century.
At first this was apparent only locally. For
example, fewer were seen in the Piscataqua after
about 1792. And very few were reported there
from about 1880 down to 1936 or 1937, when the
young fish, that were hatched in southern waters
in 1934, appeared in our Gulf in such numbers
(p. 402).

They seem to have continued moderately plenti-
ful in Massachusetts Bay and around Cape Cod
during the first half of the nineteenth century,
when bass were still being netted in abundance
along the beaches between Boston and Cohasset;
300 good-sized fish were taken in one seine haul
at the mouth of Barnstable Harbor in July 1829;
while 700 were taken at Provincetown in a day in
October 1859. Fishing for bass from the rocks
with hook and line was a well recognized sport then
around Massachusetts Bay. But Freeman wrote
as long ago as 1862 (in his History of Cape Cod)
that the bass were much less plentiful in Cape
Cod waters then than they had been of old. And
the catch for Cape Cod Bay and the outer shore
of the Cape, combined, reached 2,000 pounds in
only three of the years of record during the last
quarter of the 19th century and the first quarter
of the 20th.4 Bass, in fact, had so nearly van-
ished soon thereafter from the Massachusetts
coast north of Boston that no commercial catches
were reported there for the period 1876 to 1883,
though an occasional fish may have been caught.

There may have been a few more bass along
the coast of northern Massachusetts during the
next 17 years, for yearly catches ranging from
none up to 441 fish (1892) were reported for Essex
County between 1884 and 1900, while a number
were seined in the Merrimac River in 1897. But
this was again succeeded by a period of scarcity
so extreme that no bass were reported for the
Massachusetts coast north of Boston during the
next 30 years.5 And the capture of a single fish
in the inner part of Massachusetts Bay by any
method was so unusual an event then that one of
44% pounds, caught near Brant Rock on the
southern shore of the Bay, in November 1930,
was given wide publicity in the newspapers.

The bass of the coasts of Maine fared no better.
They had practically gone from the Androscoggin
by 1860; the reported catch for the entire coast-
line of Maine (already only about 27,000 pounds
in 1880) had fallen to about 1,600 pounds in 1902,
4,200 pounds in 1905, to 600 pounds in 1919.
And no commercial catches at all have been re-
ported from Maine in any subsequent year,6 ex-
cept for 537 pounds in 1932.

It seems that the bass population of the St.
John River system shared with those of Massa-
chusetts and of Maine in the general decline in
abundance during the first half of the nineteenth
century, for they were reported as much less
plentiful there by 1873 than they had been in
the early 1800's when they were a familiar sight
rolling and splashing at the surface in June.7
But neither the St. John population, nor the
population at the head of the Bay of Fundy, nor
of the Annapolis River, seem ever to have fallen
to as low an ebb as has happened along Maine
and Massachusetts. In 1919, for example, when
no bass were reported from the Massachusetts
coastline of the Gulf (p. 401) and only 600 pounds
for Maine, 2,700 pounds were reported from the
Nova Scotian coastline of the Bay of Fundy,8 1,600
pounds for the St. John River system.

The year 1921 seems to have marked the "turn-
ing of the tide" for the bass in Cape Cod Bay
waters, for 4,784 pounds were taken that year at

• Wood, New Englands Prospect, 1634, p. 47.

« 1878 — 4,974 pounds; 1897 — 4,820 pounds; 1900—6,450 pounds.

' Statistics have been published for 1903 to 1910, 1919, and 1928-1930.

• Statistics published for 1929-1933, and 1935-1947.

' See Adams (Field and Forest Rambles, 1873, pt. 3, Fishes, p. 248), who
described the Indians of the Melicete Tribe as still spearing good-sized bass
from their canoes in the St. John, in 1873.

1 2,000 pounds from the Cobequid-Shubenacadie region (Hants County),
700 pounds from Annapolis County.

402

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Sandwich, near the Cape Cod Canal, through
which they may have come, while some, that did
not find their way into the official returns were
taken in the inlets along the outer shore of Cape
Cod during that summer or the summer before.
And the commercial catches for Barnstable County
rose to 8,060 pounds for 1928, to 18,665 pounds for
1929, and ran between about 27,000 pounds and
about 34,000 pounds9 for 1930, 1931 and 1932.

Surf fishermen, too, did better along the outer
Cape shore from 1930 through the next couple of
summers than they had for many years; (a 33-
pounder was taken in the surf on Cape Cod, and
one of 4.4% pounds on the south shore of Massa-
chusetts Bay, p. 401). And it appears that the
bass spread northward to the estuaries north of
Cape Ann during these years (unless a small stock
had persisted there through the poor period), for
some were taken in the Parker River in the late
1920's while 8,700 pounds were reported thence
in the winter of 1930, when net-fishing was al-
lowed, temporarily.

This upswing was brief (the reported catches for
the entire coastline of Massachusetts were only
4,500 and 5,100 pounds, respectively, for 1933 and
for 1935). I0 But at least it gave a foretaste of
what was to come, for the waters around Cape Cod
were invaded during the summer of 1936 by count-
less schools of little bass, weighing about 2 to 3
pounds. These (as is now known) had been hatched
in 1934 (i. e., 2 years previous) in the Chesapeake
Bay-Delaware Bay region (p. 393), and it is inter-
esting, not only that they came from so far away,
but that this was the largest year's brood that has
been produced in Chesapeake Bay for as far back
as any record is available.11 Unfortunately, there
is no knowing in what numbers they reached the
outer shore of Cape Cod and Cape Cod Bay in
1936, for no record seems to have been kept of
commercial catches of them there in that year.

But they (chiefly) comprised the catches which
were some 5 to 16 times as great in 1936 as in any
of the 8 previous years I2 along the coast of Rhode
Island. And considerable numbers of them were
reported from as far north as the harbors and
rivers along the southern part of the coast of

Maine, where veiy few bass, large or small, had
been caught for many years previous.

In 1937, having now grown to an average weight
of about 3 to 5 pounds, they not only reappeared
in such numbers that a commercial catch of some-
thing like 80,000 pounds was reported from the
Gulf of Maine coast of Massachusetts,13 but so
many of them spread north past Cape Ann that
the catch from the inner part of Massachusetts
Bay to the New Hampshire line (about 55,000
pounds) was perhaps three times as great as that
for the Cape Cod Bay-outer Cape region (in the
neighborhood of 19,000 pounds). And more of
these little bass were caught by anglers in the river
mouths and estuaries of New Hampshire that sum-
mer, and of Maine as far as the Penobscot region,
than had been the case the year before, but not
enough to figure in the official statistics.

The fish of the 1934 year class averaged around
6 pounds by 1939 (many had reached 7-9 pounds) ;
and the bass seemed so well established all along
from Cape Cod to southern Maine that anglers
had largely forgotten the preceding lean years.
And the growth of the individual fish as they ad-
vanced in age, combined with fresh increments
from the south seem to have more than balanced
the death rate (natural or from fishing) for the
next 5 or 6 years, for the coast of Massachusetts
as a whole.14 And a good part of the fish of the
1934 year class (still swimming in good numbers)
grew meantime to 18 to 25 pounds, to the delight
of the anglers.

Bass fishing improved so much in the Hampton
region also, and in the Piscataqua River system
that about 19,000 pounds were reported for 1943
in the commercial statistics for New Hampshire,
where bass had not been mentioned in the fishery
statistics for many ysars. But it is evident that
depletion in numbers outran renewal along the
coast of Maine during this same period, for there
were many fewer fish there in the season 1940
then there had been in 1939 or 1938, though they
ran larger, averaging about 8-10 pounds according
to local reports.

' To the nearest 1,000 pounds.
» No data for 1934.

11 Tiller, Publ. 85, Chesapeake Biol. Lab., 1950, p. 24.
» For details, see Merrlman, Fish. Bull. No. 36, U. S. Fish and Wildlife
Service, vol. 60, 1941, p. 10, flg. 4; p. 13, fig. 8.

'» Assuming that about two-thirds of the catch of 28,700 pounds for Barn-
stable County came from the outer shore of Cape Cod and from Cape Cod
Bay, probably an under estimate

'< Reported catches for Massachusetts as a whole were about 62,500 pounds
for 1939, about 75,700 pounds for 1940, about 99.500 pounds for 1943. no data
available for 1941 or 1942.

FISHES OF THE GULF OF MAINE

403

Small bass (apparently spawned in southern
waters in 1940, 1941, or 1942 16 and now large
enough to be included in the commercial statis-
tics) again entered the southern part of our Gulf
in 1944 in such numbers that the commercial
catch for Massachusetts as a whole was nearly
twice as great for that year (about 191 ,000 pounds)
as it had been the year before (about 99,500
pounds).16

But it seems that very few of the fish of the
1942 year class, if any, spread northward much
beyond the Merrimac River, for the reported
catch for New Hampshire fell from about 19,000
pounds in 1943 to between 10,000 and 11,000
pounds in 1944, and to about 9,000 pounds in
1945, though the proportion of large fish was
greater, while occasional bass, only, were re-
ported in Maine waters in these years, or in the
later 1940's.

Anglers' reports in general, and our own obser-
vations, are to the effect that few young bass ap-
peared in the Cape Cod Bay-outer Cape region
during the four years 1946-1949, or farther north
along the New England coast. While this pre-
vailing scarcity of baby bass caused widespread
fears that the striper might be facing another
serious decline, anglers welcomed an accompany-
ing increase in the numbers of bass weighing up-
wards of 20 to 25 pounds. Thus, a larger num-
ber of fish heavier than 25 to 35 pounds were
landed along the outer shore of the Cape, in Cape
Cod Bay and in northern Massachusetts waters
in 1950 than for many years. We saw one of 45
pounds that was caught by an angling companion
in the inner part of Massachusetts Bay that
September, and one of 51 K pounds, caught in the
surf, was reported from the outer shore of Cape
Cod that August, while others, doubtless in the
50-pound class, were taken of which we did not
chance to hear. A few very large fish were re-
ported, that summer, in the surf at Old Orchard
Beach, Maine. And hundreds of bass of 25 to 45
pounds, with a few running up to 52 or 53 pounds
(few smaller than 25-30 pounds) were caught in
the surf on the outer beach near the tip of Cape
Cod during the summer of 1951, many up to

" The broods of 1940-1942 were large, in Chesapeake Bay, though not as
large as the brood of 1934 (Tiller, Chesapeake Biol. Lab., Pub. 85, 1950, pp.
13, 24-25).

>■ The minimum legal length for bass (snout to fork of tail) having been
set in Massachusetts at 16 inches (fish 3-4 years old).

30-45 pounds in Cape Cod Bay that July; a few
as large as 20 to 30 pounds in Duxbury Bay; at
least two (to our knowledge) weighing a little
more than 50 pounds in the inner part of Mass-
achusetts Bay; a few in the 30 to 45 pound class
on the northern Massachusetts coast.

It is almost certain that most of these large fish
belong to the abundant year classes that were
spawned in the early 1940's or even previously.
Hopes for the future depend, therefore, on renewed
replenishments of the stock. A year ago (in 1950),
prospects seemed good in this respect, for great
numbers of little bass (many smaller than the
legal length in Massachusetts, 16 inches from snout
to fork of tail) were reported that summer and
autumn from various localities along southern
New England, and northward as far as Plymouth
and Duxbury Bays; from the North and South
Rivers in Marshfield; from Boston Harbor; from
the Parker River (Plum Island Sound region);
from the lower Merrimac River; and from Hamp-
ton Harbor, N. H. Many "school" fish of 2 to 8
pounds seen (and some caught) in the Saco in
July and August of 1950, and a few landed
every day from the York and Mousam Rivers late
that June, point similarly to a fresh influx of bass
to southern Maine waters, either that year or the
year before, for it is not likely that these fish had
been spawned in the streams along that part of
the coast.

And reports that Pleasant Bay, on Cape Cod,
the Massachusetts Bay end of the Cape Cod
Canal, and Duxbury Bay, have all been "loaded"
with small bass at times during the present summer
(1951) and also of many too small to keep, off
Wollaston Beach in Boston Harbor, are promising
at least; so is the fact that a good many fish in the
10 to 15-pound class have been caught at various
places along the coast.

An interesting aspect of the bass situation is that
the young bass that invaded the water of Massa-
chusetts and of Maine in 1936 and 1937 seem not
to have spread to the St. John River system for
while commercial catches ran larger there during
the 1930's than they had during the 1920's, the
increase may not have been greater than can be
accounted for by an increasing demand for bass.
And, in any case, it had taken place by 1932, i. e.,
two years before the fish were spawned that re-

404

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

plenished the Massachusetts stock in 1936 and
1937 (p. 402) ,17

Some of these that reached Massachusetts in
1944 may have spread to Nova Scotia, also, for
the average catch was larger there from 1944 to
1946, than it had been for many years. But the
increase was not great enough to suggest than any
large recruitment had taken place from the south.18

Additional evidence that the bass of the Bay of
Fundy region do not intermingle to any great
extent with those of the western side of the Gulf of
Maine is that the fish ran so small in the Nova
Scotian streams in 1949 and 1950 (p. 397) when
there were so many very large ones in Massachu-
setts waters (p. 403). And the fact that those
caught in those years ran considerably larger in
the Annapolis River than in the Shubenacadie
River (p. 397) emboldens us to suggest that the
populations in the several Nova Scotian streams
are more or less separate; and separate also from
the bass of the St. John.

It would, of course, be of the greatest interest,
to commercial fishermen and to anglers alike, to
know what determines that an abundant brood
of bass, or a poor brood is to be reared in any given
year. All we dare say is that the largest brood
on record (that of 1934, in Chesapeake Bay) was
produced when the parent stock was at a very low
ebb, which may prove a general rule, and that
studies by Merriman19 and by Tiller20 suggest
that very large broods are raised only in years
when the temperature of the water is lower than
normal, both before the spawning season and
after it.

Importance. — Striped bass have not been plenti-
ful enough in the Gulf of Maine at any time during
the past 100 years to support a commercial fishery
of any great magnitude. Even in the good years
1944-1946 the reported value of the commercial
catches for Massachusetts as a whole was less
than $50,000 yearly.21 But this does not take
into account bass used for home consumption, or
those sold in small lots. In the seasons of 1937
and 1938, when the yield of the year class of 1934

" Maximum reported catch for St. John River system for period 1922-1943,
was 21,200 pounds in 1932.

u Average yearly catch, Nova Scotian rivers and coast of the Bay of Fundy
was about 3,355 pounds for 1922-1930; about 12,600 pounds for 1932-1943; about
18,300 pounds for 1944-1946.

'• Fish. Bull. 35, U. S. Fish and Wildlife Service, 1941, vol. 60, p. 14.

» Pub. 85, Chesapeake Biol. Lab., 1950, pp. 18, 28.

!l 1944, $29,173; 1944, $34,643; 1945, $48,748; 1946, $34,643.

was at its peak, about 30-40 percent of the re-
ported commercial catch was made by hook and
line, about 40-45 percent in pound nets and traps.
But with the development of increasingly efficient
methods of trolling with hand lines from small
craft, the hook and line catch increased in impor-
tance to about 63-65 percent for the seasons of
1939, 1940, and 1943,22 and to about 89-91 percent
for 1944, 1945, and 1946, while the pound net
and trap catches decreasing correspondingly.23

The striper is the leading game fish in its periods
of plenty all along our coast, from the outer shore
of Cape Cod to New Hampshire waters. The
number of anglers who cast for them in the surf
along the beaches of Cape Cod and northward
from Cape Ann to the mouth of the Merrimac and
at scattered spots elsewhere certainly is in the
thousands. Many party boats troll daily for bass
in Cape Cod Bay, some also along the Plum Island
shore and at the mouth of the Merrimac, while
many are caught by trolling, by live line fishing,
and even by still fishing in the various inlets.

So far as we know, Shubenacadie River and
Lake, and the Annapolis River are the only waters
on the Canadian shores of our Gulf where the
striper attracts attention as a game fish (p. 397);
anglers visiting the St. John are far more interested
in salmon.

So much has been written about the techniques
of surf casting, trolling, choice of lines, and baits,
that we need not delay; but it is interesting, in
comparison, to read, in Wood's New England's
Prospect, published in 1634 (p. 37), that "the way
to catch them is with hook and line, the fisherman
taking a great cod line to which he fasteneth a peece
of lobster and threwes it into the sea. The fish
biting at it, he pulls her to him and knockes her
on the head with a sticke".

We should point out in conclusion, that the
recreational value of the striper is high ; its money
return to the seaside communities where bass are
plentiful is much greater than the price the fish
bring in the market, if the amounts spent for tackle
and bait, boat hire, lodging, guide service, and the
patronage of service garages, and gasoline filling
stations are taken into account.

» No data are available for 1941 or 1942.

» About 15-35 percent for 1939, 1940, and 1943; about 5-11 percent for 1944,
1945. and 1946.

FISHES OF THE GULF OF MAINE

405

White perch Morone americana

(Gmelin) 1789

Sea perch

Jordan and Evermann, 1896-1900, p. 1134.

Description. — The white perch resembles its
larger relative, the striped bass, in the number,
outline, and arrangement of its fins, and in its
deep caudal peduncle without longitudinal keels.
But it is a deeper bodied fish, (only about 2% to 3
times as long as deep, not counting the caudal
fin) ; and it is more flattened sidewise (p. 390) . The
dorsal profile of its body is more convex than that
of a bass, but its head is rather noticeably concave
and its mouth is smaller. Furthermore, there is
no free space between the two dorsal fins of the
white perch, whereas they are separated by a short
interspace in the striped bass. The perch has
fewer rows of scales between gill cover and base of
tail than the bass (about 48 in the white perch,
60 or more in the striped bass), and its anal spines
are much stouter than those of the bass with the
second and third about equal in length (graduated
in the bass); also it usually has only one spine
(sometimes two) at the margin of the gill cover.
Finally, there is a constant difference in color.

The first dorsal fin (9 spines) of the perch is
rounded in outline with its thud and fourth
spines longest, and although there is no free
space between the two dorsal fins they are entirely
separated by a deep notch. The second dorsal

fin (1 spine and 12 rays) is rhomboid in outline
and so short that it leaves a rather long caudal
peduncle bare. The anal fin (8 to 10 rays pre-
ceded by 3 stout spines) originates under the middle
of the second dorsal and is of the same shape as
the latter. The ventrals originate a little way
behind the pectorals and each ventral is armed
with one stout spine at its forward margin. Both
the pectorals and the ventrals of the perch are
larger, in comparison with the size of the fish,
than those of the striped bass.

Color. — The upper surface is variously olive,
dark grayish green, or dark silvery gray, shading
to paler olive or silvery green on the sides and to
silvery white on the belly, while large fish often
show a bluish luster on the head. The fins often
are more or less dusky. The ventral fins and the
anal fin are sometimes rose-colored at the base.
The sides of young specimens are marked with
pale longitudinal stripes but these usually fade
out with growth.

Size. — White perch are occasionally as much as
15 inches long, 5 inches or more deep, and 2 pounds
or a little more in weight; but the average is 8 to
10 inches long and 1 pound in weight, or less.

Habits. — The white perch is much more closely
restricted in its seaward range than the bass, for
while they are taken in undiluted sea water along
southern New England, and at various other locali-
ties thence westward and southward, they are
much more plentiful in ponds connected with the
sea, in the brackish water of bays behind barrier

Figure 210. — White perch (Morone americana). From Goode. Drawing by H. L. Todd.

406

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

beaches, in estuaries, and in river mouths. White
perch also occur landlocked in fresh-water ponds
in many places.

They are ordinarily found in shallow water,
usually not deeper than perhaps a fathom or two,
but sometimes as deep as 10-21 fathoms in
Chesapeake Bay.24 However, they are not bot-
tom fish (except in winter), but wander from place
to place in small schools. Apart from this, they
are resident throughout the year wherever found.
In winter they congregate in the deeper parts of
the bays and creeks, where they either hibernate,
or at least pass the cold season in a sluggish
condition.

When living in salt or brackish water white
perch feed on small fish fry of all kinds, young
squid, shrimps, crabs, and various other inverte-
brates, as well as on the spawn of other fish, of
which they are very destructive. Swarms of
young perch, for instance, have been seen follow-
ing the alewives around the shores of ponds on
Marthas Vineyard, eating their spawn as it was
deposited. They bite freely on almost any bait,
natural or artificial.

Breeding. — Along southern New England the
white perch spawn in April, May, and June.
Presumably the season commences a few weeks
later around the Gulf of Maine, but definite data
are lacking.26 Those living in salt water run up
into fresh or slightly brackish water to spawn.
The eggs (about 0.73 mm. in diameter, with large
oil globule) sink and stick together in masses, or to
any object on which they chance to rest. Incuba-
tion occupies about 6 days at a temperature of
52°. The newly hatched larvae are about 2.3
mm. long with the vent some distance behind the
yolk sac and with very little pigment. In five or
six days after hatching the head begins to pro-
ject forward, the yolk sac has been partly absorbed
and branched pigment cells have appeared on the
oil globule. The late larval and post larval
stages have not been described.26

General range. — Atlantic coast of North America
from the Gulf of St. Lawrence and Nova Scotia to
South Carolina, breeding in fresh or brackish water

« Hildebrand and Schroeder (Bull. U. S. Bur. Fish., vol. 43, Ft. 1, 1928, p.
245) report ripe specimens as deep as 9H to 21 fathoms In Chesapeake Bay.

»» In Chesapeake Bay they spawn chiefly In April and May, but they are
known to do so exceptionally in December (Hildebrand and Schroeder, Bull.
U. S. Bur. Fish., vol. 43, Ft. 1, 1928, p. 245).

» Ryder (Rept. U. S. Comm. Fish., (1885) 1887, p. 518) describes the early
development.

and permanently landlocked in many fresh ponds
and streams.

Occurrence in the Gulf of Maine. — The white
perch inhabit salt, brackish, and fresh water in-
differently along the shores of southern New Eng-
land. But while this is a familiar fish in many
ponds throughout northern New England, New
Brunswick, and Nova Scotia, they are found
regularly in only a few estuarine situations north
of Cape Cod, and they hardly belong to the fish
fauna of the open Gulf. Thus we have heard only
vaguely of them in Duxbury Bay and in the North
and South Kivers in Marshfield; and we had not
been able to satisfy ourselves of their presence in
the salt creeks about Cohasset, Mass. (localities
apparently suited to it) until the summer of 1950,
when white perch running-up stream to a pond
were reported there.27 Storer long ago described
white perch as brought to Boston market from the
mouths of neighboring rivers and from ponds to
which the sea had access. And white perch run in
salt and brackish reaches of the Parker River in
northern Massachusetts, providing fishing for
many small boat anglers in spring and summer.

Ordinarily white perch are so scarce along the
open coast from Cape Cod northward that they
did not figure in the statistics of the shore fisheries
of any part of Massachusetts Bay from 1907 to
1928.28 And ordinarily they are not common
along the coast of Maine ; none was reported from
the shore fisheries of Maine in 1905 or 1919, and
only 400 pounds in 1902; none at all of late years.
But they appear locally, however, in unusual num-
bers on rare occasions. Thus it is probable that
certain unfamiliar fish taken at Beverly on the
north shore of Massachusetts Bay during the
summer of 1950, and in Casco Bay, were white
perch.29 No less than 1,600 pounds of white perch
were reported for the shore fisheries of the short
coast line of New Hampshire in the year 1912:
Casco Bay saw a run of them in the summer of
1901 when local fishermen, not knowing the fish,
dubbed them "sea bass"; and they have been re-
ported at Eastport, Maine. But apparently they
do not occur around the shores of the Bay of
Fundy, either in salt water or in brackish. And
there is no reason to suppose that white perch were
more regularly plentiful along the coast of the
Gulf of Maine than they are today.

» Reported by Lenore Williams, Salt Water Sportsman for June 30, 1950.
* Only recent years from which detailed information is available.
n Moore, Boston Herald, for August 7, 1950.

FISHES OF THE GULF OF MAINE

407

In more eastern Nova Scotian waters, also,
perch are "less often seen" in salt and brackish
water than in fresh30 and they appear to be re-
stricted, in the eastern side of the Gulf of St.
Lawrence, to the "estuarine transition" from salt
water to fresh.31

Importance.- — The white perch is of considerable
commercial importance wherever it is abundant in
tide waters. The commercial catch in Chesa-
peake Bay, for example, was 1,143,700 pounds for
1946, 1,851,000 pounds for 1947. And several
millions of artificially hatched fry are released
there yearly. It also affords good sport to many
anglers wherever it is plentiful, both in brackish
water or in fresh. But it is not important in the
open Gulf of Maine in either of these respects.

Sea bass Centropristes striatus (Linnaeus) 1758

Black sea bass; Blackfish

Jordan and Evermann, 1896-1900, p. 1199.

Description. — The sea bass is easily distinguished
from the striped bass and from the white perch by
the fact that the spiny and soft-rayed portions
of its dorsal fin are continuous, so that there is
only one long fin instead of two short separate fins.
It agrees with its nearer relative the wreck fish
(p. 409) in this; also with the scup (p. 411), with the
rosefish (p. 430), with the cunner (p. 473), and with

» Vladykov and MacKenzie, Proc. Nova Scotia Inst. Sci., vol. 19, 1935,
p. 91.
>i Huntsman, Trans. Roy. Soe. Canada, Ser. 3, vol. 12, Sect. 4, 1918, p. 63.

the tautog (p. 478). But its general form, rounded
caudal and pectoral fins, and its short but high anal
fin are sufficient to separate it from the scup, its
color prevents confusing it with the rosefish; and
no one should take sea bass for tautog or cunner;
its mouth and its pectoral fins are so much larger,
its caudal of different outline (cf . fig. 211 with figs.
249, 250), and the soft portion of its dorsal as long
as the spiny portion. It differs from the wreck fish
(p. 409), in many respects, especially in its much
larger scales; in the smoothness of its head and gill
covers; and in the shape of its tail fin.

It is moderately stout-bodied, about three times
as long (not counting the caudal fin) as it is deep,
with rather high back but flat-topped head, mod-
erately pointed snout, a large oblique mouth, eye
set high up, and one sharp flat spine near the rear
angle of each gill cover. The spiny (10 spines)
and soft (11 rays) portions of its dorsal fin (which
originates slightly in front of the rear corner of the
gill covers) are separately rounded, the latter higher
than long, with the characteristic outline shown in
the illustration (fig. 211). The caudal fin is
rounded. In large fish one of the upper rays is
much the longest, and though the resulting out-
line is a trivial character and variable from fish to
fish it is an extremely characteristic one that is
shared by no other Gulf of Maine species except
the kingfish (p. 423). The anal fin (3 short sharp
spines followed by 7 soft rays) originates under or
very slightly behind the origin of the soft portion
of the dorsal fin, which it resembles in its rounded

Figubb 211. — Sea bass (Centropristes striatus), Connecticut. From Goode. Drawing by H. L. Todd.

408

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

outline and in being higher than long. Both the
anal fin and the soft part of the dorsal are notice-
ably soft and flexible. The pectorals are so long
' that they reach back almost, if not quite to the
anal, broad and round tipped; a good field mark.
The ventrals, too, are larger than in any other fish
with which the sea bass might be confused, and
they originate in front of the pectorals, whereas
they stand slightly behind the latter in scup, rose-
fish, cunner, and tautog. The scales are rather
large, but the top of the head is naked. Adult
males develop a fatty hump on the back in front
of the dorsal fin.

C0I01 . — Sea bass, like most fish that he on rocky
bottom, vary widely in color, the general ground
tint ranging from smoky gray to dusky brown or
blue black, usually more or less mottled. The
belly is only slightly paler than the sides. On
every sea bass we have seen the bases of the ex-
posed parts of the scales are paler than their mar-
gins, giving the fish the appearance of being barred
with longitudinal series of dots of a lighter tint of
brown than the general hue on dark fish, but pearl
gray on pale ones. The dorsal fin is marked with
several series of whitish spots and bands; the other
fins are mottled with dusky. Young fish 2 or 3
inches long are greenish or brownish with a dark
side stripe passing from eye to caudal fin, and
with dark cross bars on the sides.

Size. — Sea bass grow to a length of 2 feet or more
and a few reach a weight of 7K pounds; but north-
ern specimens are seldom heavier than 5 pounds,
and they average only about 1^ pounds. A fish
a foot long weighs about one pound, one of 18 to
20 inches about 3 pounds.

Habits. — The sea bass contrasts with the striped
bass in being strictly confined to salt water. Its
inshore-offshore range extends from close in to the
coast line, in depths of only a few feet, out about
to the 70-fathom contour line, according to the
season of the year. Off New Jersey, Long Island,
and southern New England they appear inshore
during the first or second week in May, withdraw-
ing again late in October or early in November.

They winter offshore along the 30- to 70-fathom
zone; the depth and the distance offshore being
governed, it seems, by a preference for tempera-
tures higher than about 46°-47°.32

It seems, too, that some of the population that
summers off New Jersey and to the northward
may combine this offshore movement with a
southward migration, for sea bass form a consider-
able part of the catches that are made by the winter
trawl fishery off Virginia and northern North Car-
olina from January to April,33 whereas they have
been taken in small numbers only (though widely
dispersed) off southern New England at that time
of year.34

During the part of the year when the sea bass
are inshore they are most plentiful on hard bottom,
in water less than 20 fathoms or so, often around
submerged wrecks and the pilings of wharves.
They are bottom feeders, subsisting chiefly on
crabs, lobsters, shrimp, and various mollusks.
They also eat small fish (e. g., launce and men-
haden), and squid on occasion. And they take
a hook readily.

The sea bass spawn in May along the North
Carolina coast ; from the middle of May to the end
of June off New Jersey, off Long Island, and off
southern New England. The eggs are buoyant.36
The young fry are easily identifiable as sea bass
by the time they have grown to a length of 2%
inches (60 mm.) or so.

General range. — Atlantic coastal waters of the
United States, from northern Florida to Cape Cod,
occasionally to Maine.

Occurrence in the Gulf of Maine. — The sea bass
enters our Gulf only as a rare stray from the south,
Pemaquid Point and Matinicus Island being its
nothernmost known outposts. It has been taken
in Casco Bay; near Gloucester (where a few have
been caught in the traps) ; off Nahant, Salem, and
Beverly in Massachusetts Bay; at North Truro
and at Monomoy on Cape Cod; and 5 miles east
of Pollock Eip Lightship, where a 5-pound fish
was trawled in 24 fathoms, December 1930. 38
But it has never been found in any numbers north
of the elbow of Cape Cod so far as we can learn.
We have never seen it in the Massachusetts Bay
region, nor are fishermen of whom we have in-
quired familiar with it there. Sea bass, it is true,

» Neville, Fishery Circular No. 18, U. S. Bur. Fish., 1935, p. 3-7.

33 For quantities caught and other details, see Pearson, Investigational
Report No. 10, U. S. Bur. Fish., 1932.

« We counted from 1 to 25 sea bass per haul in 31 out of 45 trawl hauls made
by the dragger Eugene H off Rhode Island and off southern Massachusetts,
in 46 to 67 fathoms. Jan. 27 to Feb. 3, 1950.

>> The early development of the sea bass has been described by Wilson (Bull
U. S. Fish Comm., vol. 9, 1891, p. 209).

- Reported by Firth, Bull. 61, Boston Soc. Nat. Hist., 1931, p. 12.

FISHES OF THE GULF OF MAINE

409

occasionally appear in the returns of the local
pound nets and traps.37

But it is doubtful whether these records can be
accepted, for when the name "sea bass" is used
along the northern New England coast it usually
is either striped bass (p. 389), white perch (p. 405),
tautog (p. 478), or even rosefish (p. 430) that is
meant.38 No sooner do we round Cape Cod to
the west, however, than we find the sea bass one
of the important ground fish.

Judging from its season at Woods Hole, where it
is to be caught from May to October (most abun-
dantly in July, August, and September), sea bass
are most likely to be taken in the Gulf of Maine in
summer, if at all, though there is one record for
December. There is no reason to suppose that
they ever succeed in reproducing themselves in
the Gulf or in establishing a temporary foothold
even if the rare migrants should spawn there.

Importance. — Too scarce to be of any importance
in the Gulf, the sea bass is a very valuable food and
game fish in more southern waters.

" For example, 80 pounds at Provincetown for 1896; 146 pounds at Truro
1898; 101 pounds at the same locality for 1900; with occasional fish at Eastham,
Barnstable, Sagamore, Manomet, and Gloucester.

*8 The 3,000 pounds of "sea bass" reported from Manchester, Mass., in 1911
certainly were not this fish.

Wreck fish Polyprion americanus (Bloch and
Schneider) 1801

Wreck bass

Jordan and Evermann, 1896-1900, p. 1139.

Description. — The combination of a sea-bass-
like body with a very rough head having a promi-
nent ridge and strong spines on each gill cover,
and a bony protuberance over the eye and on the
nape, give the wreck fish an aspect so different
from that of any other Gulf of Maine fish (even
from its close relative the sea bass) that it should
be easily recognized if caught. It is strongly
flattened sidewise, about 2% to 3 times as long as
deep (to origin of tail fin), with large mouth. And
the lower jaw projects considerably beyond the
upper. The scales are rough, much smaller rela-
tively than in the sea bass, and they extend over
the bases of the soft-rayed fins. The first part of
the dorsal fin has 11 strong spines, the second
part, 11 or 12 soft rays, and the spiny part of
the dorsal is continuous with the soft-rayed part.
The anal fin. with 3 spines and 8 or 9 rays, is
similar to the soft-rayed part of the dorsal in out-
line. The caudal fin is gently rounded; the pec-

Figcre 212. — Wreckfish (Polyprion americanus), Grand Bank. From Goode and Bean. Drawing by H. L. Todd.
210941—53 27

410

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

torals (about half as long as the head) stand
almost above the ventrals.

Color.— Grayish or blackish brown, the caudal
fin edged with white. Young fish are mottled
above with gray and cream on head and body.

Size. — Reaches a length of 4% to 5 feet at least,
and a weight of more than 100 pounds.

Habits.- — Small wreck fish are most likely to be
found under floating logs or wreckage, as the com-
mon name implies. When larger, they take to
bottom; this, at least, is the case around Madeira
and in the Mediterranean.

General range. — This is a fish of wide distribu-
tion. In the eastern Atlantic it is known as far
north as Norway, as far south as the Canaries;
also in the Mediterranean. It has been reported

only occasionally in the western Atlantic, but its
latitudinal range there extends from the Grand
Banks of Newfoundland to the La Plata River.
It is also known from the Cape of Good Hope and
Indian Ocean.39

Occurrence in the Gulf of Maine. — The only re-
port that has reached us of a wreck fish in any
part of the Gulf of Maine is of one 24 }{ inches long,
weighing 9 pounds 7 ounces (dressed), taken on
the northern edge of Georges Bank, August 13,
1951, by the trawler Winthrop.*0 Another, 6 in-
ches long, was caught on the surface off No Man's
Land Island, near Martha's Vineyard, August 21,
1925; and two have been brought in from the
Grand Banks, one of them many years ago,41 the
second in 1929.42

THE CATALUFAS OR BIG EYES. FAMILY PRIACANTHIDAE

The big eyes are very closely related to the sea
basses (Serranidae), from which they differ chiefly
by the fact that the entire head, including the
snout and upper jaw, is clothed with rough scales.

Short big-eye Pseudopriacanthus alius
(Gill) 1862

Jordan and Evermann, 1896-1900, p. 1239.

Description. — The most striking characters of
this fish are its very large eyes and its brilliant red
color. Apart from these, it is distinguishable from
the seabass tribe by the fact that its whole head,
as well as its body, is clothed with rough scales
and that the anal fin is longer than the soft-rayed
portion of its dorsal fin. Its sidewise flattened
body, unusually stout dorsal fin spines, very large
ventral fins, and small pectorals, are ready field
marks to separate it from the rosefish, the only
common Gulf of Maine species of similar appear-
ance that rivals it in color. The big-eye is ovate
in outline, very thin through, with rounded dorsal
profile, large head, notably oblique mouth, and
enormous eyes. The spiny (10 spines) and soft
(11 rays) portions of its dorsal fin are continuous,
and extend back from the nape nearly to the base
of the caudal fin. The anal (3 stout spines and
9 or 10 rays) originates under the eighth or ninth
dorsal spine and its soft portion is nearly of the
same form as the soft portion of the dorsal, except
that its outer angle is somewhat more rounded.
The caudal is square-cornered and slightly convex.
The ventrals, which originate a little in front of the

pectorals, are much larger than the latter, round
tipped, and each commences with a stiff spine.

Color. — Bright red in life, below and above;
dorsal fin red, the spinous part edged with yellow,
a few blackish dots on the soft rays; caudal fin
pale, with blackish reticulations; anal red, edged
with black; ventrals red at base, dusky on outer
part; pectorals plain red. The iris is gold.

Size. — The largest specimen on record was 11
inches long.

General range. — Caribbean Sea, West Indies,
and Gulf of Mexico in rather deep water, straying
northward to the Woods Hole region and very
rarely rounding Cape Cod.

Occurrence in the Gulf of Maine. — A big-eye
found alive on Marblehead Beach, September 3,
1859; a second, found at Scituate, Mass., in 1932
or 1933 ;a and a third, about 1% inches (38 mm.)
long, picked up in a tide pool at Cohasset, Mass.,
by F. G. Bemis in September 1937," are the only
definite records for this southern fish within the
Gulf. But since it occasionally appears in some
numbers at Woods Hole in summer, it may round
Cape Cod more often than this paucity of actual
records suggests.

» We have given a more detailed statement elsewhere (Copela, 1930, No. 2,
p. 46).

* This specimen, which we have examined, is in the collection of the U. 8.
Fish and Wildlife Service at Woods Hole.

» Ooode and Bean, Smithsonian Contrib. Enowl., vol. 30, 1895, p. 238.

« Schroeder, Copela, June 1930, p. 48.

« Reported by MacCoy, Bull. 67, Boston Soc. Nat. Hist., 1933, p. 9.

» This specimen, reported by Schroeder (Copeia, 1937, p. 238) is in the
Museum of Comparative Zoology.

FISHES OF THE GULF OF MAINE

411

Figure 213. — Short big-eye (Pseudopriacanthus alius), Key West, Florida.

H. L. Todd.

From Jordan and Evermann. Drawing by

THE PORGIES. FAMILY SPARIDAE

The structure of the fins is essentially the same
in this family as in the sea basses; both spiny and
soft portions of the dorsal are well developed and
the ventrals are situated below the pectorals.
There are important anatomic differences, how-
ever, most obvious of which are that the edge of
the gill cover does not end with a sharp spine in
the porgies but is rounded or at most bluntly
angular; and that the maxillary bone (the bone
forming the margin of the upper jaw) is sheathed
and hidden by the preorbital bone when the
mouth is closed. Long, pointed pectoral fins are
likewise characteristic of the family; the spiny
and soft portions of the dorsal fin are continuous,
and the soft rayed anal fin is about as long as the
soft part of the dorsal.

KEY TO THE GULF OF MAINE PORGIES

1. Outline of caudal fin deeply lunate, with sharp

corners _Scup, p. 411

Outline of caudal fin only slightly concave, with
rounded corners. Sheepshead, p. 416

Scup Stenotomus versicolor (MitchiU) 1815

PORGY

Jordan and Evermann, 1896-1900, p. 1346, as Stenotomus
chrysops (Linnaeus), 1766.

Description. — Although the scup is not marked
by any one outstanding character it is made easily
recognizable by the fact that the spiny portion of
its dorsal fin is considerably longer and higher than
the soft-rayed portion, which, with its deeply
lunate caudal fin, separates it from all other Gulf
of Maine fishes of similarly deep and sidewise-
flattened bodies. The scup is about one-half as
deep as it is long (to the base of the tail fin) and
very thin through, recalling a butterfish (p. 363).
But the dorsal profile of its rather short head is
slightly concave instead of convex, and its scales
rather large, thick and firmly attached; not small,
thin and easily detached as they are in the
butterfish.

The mouth of the scup is small, its eyes are
situated high up on the side of the head, and the
margins of its gill covers are rounded. It has one

412

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 214. — Scup (Stenotomus versicolor). Adult, Woods Hole; from Goode, drawing by H. L. Todd; A, egg; B, larva,
3 days old, 2.8 mm.; C, larva, 10.5 mm.; D, larva, 25 mm. A-D, after Kuntz and Radcliffe.

long dorsal fin originating over the pectorals and
preceded by a forward -pointing spine; the spiny
(12 spines) and soft (12 rays) parts are contin-
uous, forming a single fin. As a whole the dorsal
fin is moderately high, its first spine much shorter
than the others, its rear corner rounded, and
it can be laid back in a groove along the mid
line of the back. The anal (3 spines and 11 or 12
rays) is nearly as long as the soft part of the
dorsal, under which it stands, and is almost even
in height from front to rear, but with the first

spine shorter than the others. The anal fin is de-
pressible in a conspicuous groove, like the dorsal.
The caudal is deeply concave with sharp corners.
The pectorals are very long (reaching to even
with the soft part of the dorsal), sharp pointed,
and with slightly concave lower rear margins.
The ventrals, situated below the pectorals, are of
moderate size.

Color.- — Dull silvery and iridescent, somewhat
darker above than below; the sides and back with
12 to 15 indistinct longitudinal stripes, flecked with

FISHES OF THE GULF OF MAINE

413

light blue and with a light-blue streak following
the base of the dorsal fin. The head is silvery,
marked with irregular dusky blotches; the belly
is white. The dorsal, caudal, and anal fins are
dusky, flecked with blue; the pectoral fins of a
brownish tinge; the ventrals white and bluish, and
very slightly dusky; the iris silvery; the pupil
black.

Size.- — The scup is said to reach a length of 18
inches and a weight of 3 to 4 pounds, but adults
usually run only up to about 12 to 14 inches, and
weigh only 1 to 2 pounds.

Habits. — Scup are inshore from early April at
the mouth of Chesapeake Bay, and from early
May northward to southern Massachusetts. Most
of them withdraw from the coast late in October,
though some few linger through November, and an
occasional fish into December even as far north as
the vicinity of Woods Hole.

It has been known for the past 20 years or more
that many scup winter off Virginia and off north-
ern North Carolina, in depths of 20 to 50 fathoms,
where large commercial catches are made yearly
by otter trawlers from January to April,46 with a
few as deep as 90 fathoms or so. And marking
experiments have proved that some of the scup
that summer along southern Massachusetts mi-
grate southward in autumn as far as to the offings
of Chesapeake Bay and of northern North Carolina
for the winter, at least in some years, and vice
versa.46

Scup have, however, been taken during the
past few winters in depths of 45 to 70 fathoms off
southern New England, in numbers large enough
to show that part of the northern contingent
of the species simply moves offshore in autumn,
to come inshore again in spring.47

'• Reported catches for 1930-1931 (the only winter tor which statistics are
readily available) were 9,684 pounds in December, 495,312 pounds in January,
637,595 pounds in February, 653,276 pounds in March, and 76,322 pounds in
April (Pearson, Investigational Report, No. 10, V. S. Bur. Fish., 1932, p. 14,
table 2). In February 1930 Albatross II trawled three off Chesapeake Bay in
93 fathoms.

« One scup, tagged in summer near Woods Hole, was recaptured in winter
off northern Virginia; two off Chesapeake Bay; and one off northern North
Carolina (Neville, Fishery Circular No. 18, U. S. Bur. Fish., 1935, p. 3, flg. 3).
Three tagged in winter off Virginia were recaptured in summer along New
Jersey.

« We counted from 1 to 40 scup per haul in 17 trawl hauls out of a total
of 44 hauls, on the Eugene H off Rhode Island and southern Massachusetts,
Jan. 27 to Feb. 3, 1950, at depths of 47 to 67 fathoms; a dragger that caught
7 to 30 bushels in 3 hauls nearby at the time reported catches of 2,000 to 5,000
pounds as sometimes made in the vicinity at that same season; and the
Priscilla Vreported taking 445 pounds on Jan. 12, also 1, 230 pounds on Jan.
21, 1950, at 62 to 54 fathoms, some 75 to 82 miles south of No Mans Land off
Marthas Vineyard. The Eugene H fishing near Hudson Gorge in about
62 fathoms, caught 30,000 pounds of scup on a trip April 1-6, 1953.

Differences in the locations where the largest
catches are made in cool winters and in warm
make it likely that a preference for water at least
as warm as about 45° F. is the factor that de-
termines how far seaward the scup move off any
part of the coast in any particular winter.48 And
they are so sensitive to low temperatures that
large numbers have been known to perish (both
large ones and small) in sudden cold spells in
shallow water.

It appears that different bodies of scup move
inshore successively in spring, for in 1950 the
Albatross III took 2,700 scup in 15 hauls at 45
to 55 fathoms, in the Hudson Gorge, on May
11-18, which is one or two weeks after the earliest
scup ordinarily appear inshore near New York.
And the fact that scup are more plentiful in June
and July than in May points in the same direction.

It has been said that the first fish to arrive in
spring are the large adults, with the immature
fish following later. But there is no definite
rule in this regard.

During their summer stay inshore, the scup
tend to hug the coast so closely that a line drawn
5 or 6 miles beyond the outermost headlands
would probably enclose the great majority of the
total population at that time of year.

Scup usually congregate in schools. The young
fry come close in to the land in only a few feet of
water. Large fish, however, are seldom caught
in summer in water shallower than 1 or 2 fathoms
(occasionally at the surface), or deeper than 15 to
20 fathoms. They prefer smooth to rocky bot-
tom, which results in a distribution so local that
one trap at Manchester, on the North Shore of
Massachusetts Bay, took small numbers of scup
in 1885, 1886, and 1887, while another trap close
by did not yield as much as one fish. They are
bottom feeders in the main, seldom rising far
above the ground, the adults preying on crusta-
ceans (particularly on amphipods) as well as
on annelid worms, hydroids, sand-dollars, young
squid, and in fact on whatever invertebrates the
particular bottom over which they five may
afford. They also eat fish fry to some extent,
such free-floating forms as crustacean and mollus-
can larvae, appendicularians, and copepods. The
young feed chiefly on the latter and on other small
Crustacea. Adult scup, like most other fish,
cease feeding during spawning time, for which

« For details, see Neville. Fishery Circular No. 18, U. S. Bur. Fish., 1935.

414

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

reason few are caught then, but they bite very
greedily throughout the rest of the summer on
clams, bits of crab, and sea worms (Nereis),
as do the immature fish throughout their stay.

Along southern New England scup spawn
from May to August, but chiefly in June. Prob-
ably spawning both commences later and continues
later for the few fish that manage to summer in
Massachusetts Bay, and it may ba assumed
that they spawn wherever they summer.

The eggs are buoyant, transparent, spherical,
rather small (about 0.9 mm. in diameter), and
have one oil globule. Incubation occupies only
about 40 hours at 72° (probably two to three
days in the June temperatures of Massachusetts
Bay) and judging from the season of spawning at
Woods Hole, it is not likely that development can
proceed normally in water colder than about
50° F. At hatching the larvae are about 2 mm.
long, the yolk is fully absorbed within 3 days
when the larva is about 2.8 mm. long, and there
is then a characteristic row of black pigment
spots along the ventral margin of the trunk. At
25 mm. the pectorals have assumed their pointed
outline and the caudal fin is slightly forked, but
the ventrals are still so small, and the body so
slender, that the little fish hardly suggest their
parentage until they are somewhat larger.49

In southern New England waters fry of 2 to 3
inches, evidently the product of that season's
spawning, have been taken in abundance as early
as September; they are 2J/2 to Z){ inches long in
October, and they may be as long as 4 inches at
Woods Hole in November. Apparently young
scup grow very little during the winter, for many
of 4 inches are seen in the spring, probably the
crop of the preceding season. According to
Neville's unpublished studies,50 scup average about
4K inches (11 cm.) long at one year of age (from
hatching), about 674 inches (16 cm.) at two years,
about 7% inches (20 cm.) at three years, about 9
inches (23 cm.) at four years, and about 9% inches
(25 cm.) at five years. If this age schedule is
correct, the ages of the large fish of 12 to 15 inches,
weighing 1% to 2% pounds are considerably
greater than the 3 to 5 years that have been
credited to them, following Baird's 81 estimate.

General range. — East coast of the United States,
from North Carolina to Cape Cod; casual in the
Gulf of Maine as far as Eastport, Maine.62

Occurrence in the Gulf of Maine. — Although the
scup is one of the most familiar of shore fishes
right up to the elbow of Cape Cod, with the
southern coast of Massachusetts and its off-lying
islands yielding annual catches of 1 million to 2
million pounds in good years, very few find their
way past Monomoy Point into the colder waters
of the Gulf of Maine.

The first definite mention of scup caught north
of Cape Cod is Storer's statement that one was
taken at Nahant in 1835, and another in 1836,
but that it was never seen there before. Possibly
these and one picked up dead at Cohasset in
1833 a were the survivors of a smack load that
had been liberated in Boston Harbor a year or two
earlier, and a similar plant was made in Plymouth
Bay in 1834 or 1835. There is no reason to sup-
pose that these planted fish established themselves.
But when the practice of setting mackerel nets
outside Provincetown Harbor was first adopted
(about 1842) a few scup were taken in them from
year to year; odd fish were caught in Cape Cod
Bay yearly and between Boston and Cape Ann
during the period 1860 to 1867; and a number were
taken in a weir on MUk Island near Gloucester in
1878. It has been learned since (mainly from the
catches of the pound nets and traps) that there
were a few scup in northern Massachusetts waters
in most years (or terms of years) down to the
first decade or so of the present century, alter-
nating with other years, or terms of years, when
only an occasional fish was taken, or none.

In most of the years for which information is
available, and when there have been any scup
north of Cape Cod, the combined catches of the
various traps have run from less than 100 pounds
to 1 to 2 thousand pounds at most, whether for
Cape Cod Bay or for the northern side of Massa-
chusetts Bay (Essex County).64 But Cape Cod
Bay seems to have seen what might almost be
called peaks of abundance in 1879 (catch, about
7,000 fish); in 1882-1885 (yearly catches 2,372-

« Kuntz and Radcliffe (Bull. U. S. Bur. Fish., vol. 35, 1918, p. 106) describe
the early development of the scup.
» Information from James A. Mason, of the U. S. Fish and Wildlife Service.
•' Rcpt. TJ. S. Comm. Fish. (1871-1872) 1873, p. 228.

•» The southern scup, Stenotomus chnjsops (Linnaeus) 1766, which was first
reported from Charleston, S. C, ranges northward about to Cape Hatteras.

" Goode, Fish. Ind. TJ. S., Sect. 1, 1884, p. 387.

" Statistics of the shore fisheries were published by the State of Massa-
chusetts in the Annual Reports of the Commissioner of Fisheries and Game
for 1879-1911 and 1917-1919; of the Division of Fisheries and Game for 1920
and 1921.

FISHES OF THE GULF OF MAINE

415

5,354 fish); in 1887,65 in 1890 (1,890 fish); and in
1895-1896 (14,362 and 5,083 fish, respectively);
also the northern side of Massachusetts Bay in
1909-1910 (8,417 pounds66 and 4,181 pounds);
both Cape Cod Bay (6,000 pounds) and the north
shore of Massachusetts Bay (3,217 pounds) in
1917.

The cataclysmic shrinkage that took place in the
stock of scup off southern Massachusetts between
1896 (prior to which the annual catch there had
usually run from 1 to 3 million pounds) and 1902,
when it fell to only about one-tenth as much (about
200,000 pounds) appears to have involved the scup
in Cape Cod Bay also, for none at all were reported
there from 1907 through 1911, or in 1918-1920,67
except that there was an unusually large run there
in 1917. But 1908, 1909, and 1919 were good
scup years for the north shore of Massachusetts,68
"good," that is, for those northerly waters, suggest-
ing that when conditions favor, a small indepen-
dent population may be present there. Perhaps
the fact that larger catches than usual are not
always registered in both these regions in the same
year may point in this same direction.

No scup were reported from Essex County for
1919, 1928, or 1930; nor were enough taken in
Cape Cod Bay in those years to cause any local
comment.69 Though the fisheries statistics do not
throw any light on the status of the scup north of
Cape Cod subsequently,60 there cannot have been
many of them in Cape Cod Bay regularly at any
time during the past 15 years or so, for the only
scup that were taken in a set of 8 traps at North
Truro from 1935 down through 1950 were 125
pounds taken on June 28, 1938, evidently one
small school of perhaps 100-125 individuals.61
And 33 barrels (about 4,950 pounds) taken in a

H The reported catch for the town of Barnstable for that year was so large
(69,168) as to suggest that it included scup from the south shore.

u If all these really were scup and not some other fish.

" No information is available for the years 1912-1916.

M Pound net catches for Essex County of 1,203 pounds, 8,417 pounds and
4,181 pounds, respectively.

« Catches reported for these years for Barnstable County include not only
such scup as may have been taken in Capo Cod Bay, but the catches (doubt-
less far larger) for the southern coast, which does not fall within the limits of
the Gulf of Maine.

M We are informed by William Royce of the V. S. Fish and Wildlife Service
that catches since 1931 have been credited to the home ports of the vessels
making them, wholly irrespective of whero the fish were caught or landed.
There is no reason to suppose that any significant part of the landings of scup
reported for Essex County since then (which reached a maximum of 7,945,209
pounds for 1938) actually came from Massachusetts Bay, or from anywhere
In Massachusetts waters, for that matter.

•' Information from the Pond Village Cold Storage Co.

trap at Sandwich on the southern shore of Cape
Cod Bay on Sept. 15 or 16, 1944, after a heavy gale,
were the only scup caught in this set of traps from
1944 to 1950.62 It would be interesting to know
whether they came through the Cape Cod Canal
or around the Cape.

lunu

! .a -.ooo
1,400,000
3,20 0,30 0
1.0 OOP 00

ip oop 0 0

2J> 00,000

1,4 CI 0,0 CO
2/00,0 0 0
ip 0 op 0 0
1,0 0 op 0 0

1.6 00.0 0 0

1,400,000

1.7 00.D 0 0

1,0 oop 00
000,0 0 0

* oop 0 0
400,000
loeooo

1

,

\

/

\

/

\

1

---

V

\

/

\

/

\

/

Figure 215. — Scup (Slenotomus versicolor). Annual catch
of scup (pounds) in pound nets and traps in Massa-
chusetts, from statistics published by the State Com-
missioner of Fisheries and Game.

Thus the presence of considerably greater num-
bers of scup on the southern coast of Massachusetts
since about 1928 than had been there during the
preceding decade 63 seems not to have been re-
flected in Cape Cod Bay except in sporadic cases.
And we have not heard of any caught in the north-
ern side of Massachusetts Bay during the past few
summers.

In any case, Cape Ann is the northern boundary
to the usual range of the scup. In 1896, a year of
plenty not only in Massachusetts Bay but to the
south in general, occasional specimens were taken
daily in Casco Bay in the Small Point traps during
the first half of July, and in July 1951, three were
reported from Small Point, Maine, sporadic visits
such as may be expected of any southern stray.
In 1938 about 100 pounds of scup were landed in
Lincoln County, Maine, probably from nearby,

K Information from Benjamin Morrow, who operates these traps.

« The yearly landings of scup for Massachusetts rose from not more than
103,000 pounds for the decade 1912 to 1921 to about 1,100,000 pounds for the 5
years 194.3-1947. But there is no way of knowing how large a part of the
catches reported during the latter period were actually taken in Massachu-
setts waters and not farther west and south along the coast.

416

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

and they have been reported from Eastport.64
But we suspect that porgies in St. Marys Bay,
Nova Scotia, reported to Knight86 were some
other fish.

Probably such scup as spread north of Cape Cod
in favorable summers withdraw southward again
(if they survive) in autumn to the same offshore
wintering grounds to which the much more numer-
ous scup repair from the southern shores of Massa-
chusetts. The fact that small scup, probably
devoured on their way offshore, have been found
in autumn in cod stomachs on Nantucket Shoals,
where scup certainly are not common in summer,
points in this direction. There is no reason to
believe that any of these fish winter in the deep
basin of the Gulf of Maine.

Importance. — Scup are never plentiful enough
anywhere north of the elbow of Cape Cod to be of
importance, whether commercially or to the angler.
But this is an important food fish to the westward
and southward where it is plentiful. Landings
ran, for example, between about 3,300,000 pounds
and 5,600,000 from the southern coast of New
England and between about 3,300,000 and4,300,000
pounds from New York, for the years 1945-47.
The "porgy", as it is commonly called along that
part of the coast, is also a favorite with anglers, for

•* A specimen taken many years ago recorded by Kendall (Occ. Pap.
Boston Soc. Nat. Hist., vol. 7, No. 8, 1908, p. 103).
- Descript. Cat. Fishes Nova Scotia 1866, p. 13.

it bites greedily and is a good pan fish. Great
numbers of them are caught on hook and line for
home consumption.

Sheepshead Archosargus probatocephalus
baum) 1792

Jordan and Evermann, 1896-1900, p. 1361.

(Wal-

Description. — The sheepshead resembles the
scup so closely in its general organization that the
family relations between the two are obvious. Like
the scup it is deep bodied and much flattened side-
wise, with similar profile. It has one long dorsal
fin, scuplike in outline, the anterior two-thirds of
which is spiny (11 or 12 spines) and the posterior
one-third is soft (11 to 13 rays). Its anal fin (3
spines and 10 or 11 rays) is about as long as the
soft portion of its dorsal, under which it stands,
and both the dorsal fin and the anal can be de-
pressed in a deep groove. The pectorals are long
and pointed; the ventrals are situated a little be-
hind the latter; the scales are large; and the eyes
are located high on the sides of the head; in all of
which the sheepshead agrees with the scup. It is
readily recognized, however, by the fact that its cau-
dal fin is not so deeply emarginate as that of the scup,
and has rounded corners instead of pointed ones,
while its dorsal spines are alternately stout and
slender; its second anal spine is much stouter than

Figure 216. — Sheepshead (Archosargus probatocephalus), North Carolina. From Goode. Drawing by H. L. Todd.

FISHES OF THE GULF OF MAINE

417

that of the scup; the dorsal profile of its head is
steeper; its nose is blunter; and its teeth are much
broader. Furthermore, the body of the sheeps-
head is noticeably thicker, its back is rounded, and
its sides show seven broad, dark brown or black
crossbars on a gray or greenish yellow ground,
instead of being plain colored like the sides of the
scup.

Size. — The sheepshead grows to a length of
about 30 inches and to a weight of 20 pounds.

General range. — Atlantic and Gulf of Mexico
coasts of the United States from Texas to Cape
Cod, and reported in the Bay of Fundy as a stray.

Occurrence in the Gulf of Maine. — The sheeps-
head was abundant as far north as New York for-

merly, and not uncommon about Woods Hole.
It is common still to the southward. But it has
been decidedly rare east of New York for many
years past, although a number, about 6 inches
long, were taken off Onset at the head of Buzzards
Bay, in late August 1951.66

The only record of it north or east of the el-
bow of Cape Cod is Cox's 67 statement that it is
occasional in St. John Harbor, New Brunswick.
But no actual specimens are mentioned, and as it
is not known ever to have strayed to Massachu-
setts Bay (a far more likely goal for any southern
coast fish than the Bay of Fundy is), its claim to
mention here is weak.

THE CROAKERS, DRUMS, AND WEAKFISHES. FAMILY SCIAENIDAE

The croakers have both the spiny portion and
the soft portions of the dorsal fin well developed
(either separate or as one continuous fin), and
their ventrals are what is known as thoracic in
position, i. e., about under the pectorals. They
are readily separable from the sea basses (p. 389,)
the porgies (p. 411), and the cunner tribe (p. 473)
by the fact that their anal fin has only 1 or 2 spines
instead of 3, and is much shorter than the soft por-
tion of the dorsal ; from the rockfishes and sculpins
by their relatively smooth head; and from all the
mackerels and the pompano tribe by their stout
caudal peduncles and rounded or only slightly con-
cave caudal fins. Most of them produce loud
drumming sounds by rapid contractions of certain
abdominal muscles against the gas-filled air blad-
der; hence the common names "croaker" and
"drum." The kingfish (p. 423) is an exception
to this rule.

KEY TO GULF OF MAINE CROAKERS
AND WEAKFISHES

1. There is no barbel on the chin 2

The chin bears one or more barbels 3

2. Body only about one-fourth as deep as it is long (to

base of caudal fin) ; anterior profile of head sloping
only moderately; snout pointed; no dark spot be-
hind upper corner of gill opening... Weakfish, p. 417."8

• Information from Mrs. Harold Hatch.

" Bull. Nat. Hist. Soc. New Brunswick, No. 13, 1895, p. 71.

« Jordan (Stanford Univ. Publ., Univ. Series, Biol. Sci., vol. 3, No. 2,
1923, p. 202) placed the weakfish in his new family Otolithidae, which he
separated from the Sciaenidae as having a different arrangement of vertebrae.
But we think it preferable (following Smith, Sea Fishes Southern Africa,
1949, p. 223) to use Sciaenidae In the older and more inclusive sense, because
the only family character marking Otolithidae off from it Is Internal, hence
requires dissection for its recognition.

210041—53 28

Body at least one-third as deep as it is long to base of
caudal fin; anterior profile of head sloping steeply;
snout blunt; there is a dark spot close behind the

upper corner of the gill opening Spot, p. 423

3. Several barbels on chin; snout ends about even with
front of lower jaw; cheek smooth. Black drum, p. 425

Only one barbel on chin; snout projects considerably
beyond lower jaw; cheek with 2 short, tooth-like
serrations Kingfish, p. 423

Weakfish Cynoscion regalis (Bloch and Schneider)
1801

Squeteague; Sea trout; Gray trout

Jordan and Evermann, 1896-1900, p. 1407.

Description. — The relative sizes and shapes of
the fins of the weakfish, and its color, are such
ready field marks that it is one of our most easily
identified fishes. Among Gulf of Maine species
with separate spiny and soft-rayed dorsal fins, it is
distinguishable from the mullet by the considerable
length of its dorsals as well as by many other
characters ; its slightly emarginate tail distinguishes
it from any mackerel or pompano ; this same char-
acter, combined with a short anal fin and a first
dorsal fin higher than the second dorsal gives it
an appearance quite different from a bluefish;
and the fact that its second dorsal is much longer
than the first, and that it has only 2 anal spines
and a slender body obviate all possibility of con-
fusing it with striped bass or white perch. The
shape of its dorsal and caudal fins and of its head,
and the absence of a chin barbel make it dis-
inguishable at a glance from the kingfish (p.

418

FISHERY BULLETIN OP THE FISH AND WILDLIFE SERVICE

Figure 217.— Weakfish (Cynoscion regalis). A, adult; from Goode, drawing by H. L. Todd. B, egg; C, larva, 12.4
mm.; D, fry, 32 mm. B and D, from Welsh and Breder; C, after Tracy.

Color. — Dark olive green above with the back
and sides variously burnished with purple, lav-
ender, green, blue, golden, or coppery, and marked
with a large number of smaU black, dark green,
or bronze spots, vaguely outlined and running
together more or less, especially on the back;
thus forming irregular lines that run downward
and forward. The spots are most numerous above
the lateral line, and there are none on the lower
part of the sides or on the beUy. The lower sur-
face, forward to the tip of the jaw, is white, either
chalky or silvery. The dorsal fins are dusky,
usually more or less tinged with yellow ; the caudal
is olive or dusky with its lower edge yellowish at
the base; the ventrals and the anal are yellow; and
the pectorals are olive on the outer side, but
usually yellow on the inner side.

Size. — It is said that weakfish have been taken
as heavy as 30 pounds, but the largest of which we
can find authentic record in recent years was one
of 17 pounds 8 ounces, taken on the New Jersey
coast, on rod and reel, by A. Weisbecker, Jr.,
September 30, 1944. And a fish heavier than 12

423), the absence of barbels on the chin separates
it from a drum (p. 425) ; it has nothing in common
with such bizarre fishes as the John Dory (p. 297),
triggerfish (p. 520) or any member of the sculpin
tribe.

The weakfish is a slim, shapely fish, about four
times as long as deep (to the base of the caudal
fin), only slightly flattened sidewise, with rather
stout caudal peduncle; a head about one-third
as long as body, moderately pointed snout, and
large mouth. Its upper jaw is armed with two
large canine teeth and its lower jaw projects
beyond the upper. The first dorsal fin (10 spines),
originating a little behind the pectorals, is trian-
gular; the second dorsal (26 to 29 rays), originat-
ing close behind the first, is more than twice as
long as the first and roughly rectangular. The
caudal fin is moderately broad and only slightly
concave in outline. The anal fin (2 very slender
spines and 11 or 12 rays) is less than half as long
as the second dorsal, under the rear part of which
it stands. The ventrals are below the pectorals,
which they resemble in their moderate size and
pointed outline.

FISHES OF THE GULF OF MAINE

419

pounds or longer than 3 feet is a rarity. Off
southern Massachusetts the largest fish run 6 to
10 pounds in weight, while most of the larger ones
taken there weigh from 1 to 6 pounds and are 14
to 26 inches long. An average of 5 pounds has
been reported for Massachusetts Bay, but this is
probably excessive. The average proportion be-
tween length and weight of weakfish is about as
follows:

mgih in inches

Weight in pounds

12 to 14

% to 1

14 to 16

1 to 1%

16 to 18

1% to 1%

18 to 20

1% to 2%

22 to 23%

3K to 4%

25K to 27^

5 to 6

30 to 32

9K to 11

The female members of a school usually run
somewhat larger than the males.

Habits. — Although there are very few weakfish
in the Gulf of Maine today, if any, they were for a
time so plentiful in its southwestern waters (and
may at any time reappear there in abundance) that
their habits deserve more attention than the fish's
present status would call for.

In the southern part of its range, (e. g., along the
Carolinas) this is said to be a resident species.
But it is strictly seasonal to the northward, appear-
ing in spring, spending the summer inshore, and
withdrawing again in autumn. Within the mouth
of Chesapeake Bay the fishing season usually is
from the middle of April (commencing a week or
two later up the bay) to the middle of November,
with good catches occasionally made as late as the
first of December. On the southern New England
coast, as illustrated by Woods Hole, weakfish are
caught from May (some years as early as April,
other years not until June) until the middle of
October. Probably they are not to be expected
north of the elbow of Cape Cod until June (in the
years when they come that far north), nor later
than September or October at latest, for most of
the weakfish disappear from the middle Atlantic
coast before the end of October.

The lower limit to the temperature range pre-
ferred by the weakfish has not been determined.
But it has long been known that they are sensitive
to cold. And a case is on record (November 27,
1903) when many were benumbed by a sudden
chilling of the water, near Beaufort, North Caro-

lina.69 Hence seasonal chilling is doubtless the
event that drives them away from the middle
Atlantic and New England coasts in late autumn.

The capture of weakfish in some numbers be-
tween the offings of Chesapeake Bay and of Cape
Hatteras by otter trawlers during the winter
months, during the past twenty-odd years,70 has
dispelled some of the mystery in which the winter
home of this fish was previously shrouded. The
fact that 5 small ones were picked up in the 50 to
55 fathom zone off Rhode Island by the dragger
Eugene H m mid-January 1950, also 6 more south
of Marthas Vineyard in about 55 fathoms,71 and
another 5 pounder on February 20 72 is evidence
that some of those that summer to the northward
only move offshore to escape falling temperature.
Others may move southward in winter for long
distances, and offshore, as some of the northward-
summering scup seem to do (p. 413).

Weakfish tend to hold close inshore during their
summer stay on the coast; we have never heard of
one on Nantucket Shoals, and only once of
weakfish caught on Georges Bank.73 They are
usually found in shallow waters along open sandy
shores and in the larger bays and estuaries, in-
cluding salt marsh creeks. They even run up
into river mouths, but never into fresh water, so
far as we know.

Weakfish move in schools, often small but
sometimes consisting of many thousands.74 They
have been described repeatedly as swimming near
the surface, this being the general rule near New
York and along the southern New England coast,
where great numbers are caught on hook and line
within a few feet of the top of the water. And
their preference for shallow water is reflected in
the large numbers caught in pound nets along the
middle Atlantic coast. Probably few descend
deeper than 5 to 6 fathoms during the summer, but
the precise level at which they are to be caught at

•' Smith, North Carolina Geol. and Economic Survey, vol. 2, 1907, p. 411.

n See Pearson, Investigational Report No. 10, U. S. Bureau of Fisheries,
1932, p. 14, table 2, for the catches for the winter of 1930-1931, by species and
by months. The Albatross III, also, trawled 83 weakfish in 29 fathoms off
Capo Hatteras, and 1 in 14 fathoms off Charleston, S. C, in late January 1950.

" Reported by Capt. Henry Kllmrn. We saw one of them.

7* We saw this fish.

71 Two nsh were reported by an otter trawler from the offshore part of the
Bank in the summer of 1950.

n A notable and oft-quoted instance was off Rockaway Beach, N. Y.,
July 1881, when a school was sighted so large that three menhaden steamers
seined some 200,000 pounds of weakfish from it, averaging Hi to 3 feet In
length.

420

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

any given locality is governed by their food at the
time. On open coasts they often feed on bottom
right in the surf. They also feed on bottom in
estuarine waters when dieting on bottom-living
animals, but in the upper water layers when
preying on small fish.

Weakfish feed on a wide variety of animals,
including crabs, amphipods, mysid and decapod
shrimps, squid, shelled mollusks, and annelid
worms, but chiefly on smaller fish, such as men-
haden, butterfish, herring, scup, anchovies, silver-
sides, and mummichogs, of which they destroy
vast quantities. The precise diet varies with the
locality (that is, with what is most readily avail-
able), but small menhaden are probably the most
important single item. The adult weakfish usu-
ally depend on fish, though occasionally they have
been found feeding exclusively on crabs and
shrimps. The young depend more on shrimp and
on other small crustaceans than the adults.76
Weakfish bite greedily on various kinds of bait,
especially on shedder crabs, clams, shrimp, and
mummichogs or other small fish. And they are
often caught on artificial lures of one kind or
another.

The females do not make any sounds, but the
males have well-developed croaking muscles in the
walls of the abdomen, with which they make a
drumming noise.

Breeding habits.- — On the middle Atlantic coast
the weakfish spawn from May to October, with
the chief production of eggs between mid-May and
mid-June.76 The eggs have been taken in tow
nets at various localities in temperatures ranging
from 60° to 70°, in salinities of 28.01 to 30.9 per
mille. And it is probable that weakfish spawn
locally around the shores of Cape Cod Bay in
years when the fish are plentiful there, as they do
regularly about Woods Hole, if the summer
temperature of the surface is high enough.
Spawning takes place chiefly in the larger estuaries
or close to their mouths, usually at night. The
eggs are buoyant, spherical, 0.74 to 1.1 mm. in
diameter, usually with one, rarely with as mariy
as four, oil globules that coalesce into one large
one as development progresses. Incubation oc-

'• For diet lists of weakfish of various sizes, see especially Welsh and Breder
(Bull. D. S. Bur Fish. vol. 39, 1924, p. 159); also Peck (Bulletin TJ. 8. Fish
Comm., vol. 15, 1S96, p. 352).

'• The following account of the breeding and development of the weakfish
Is condensed from Welsh and Breder (Bull. U. S. Bur. Fish., vol. 39, 1924, p.
150).

cupies 36 to 40 hours at a temperature of 68° to
70°, and the newly hatched larvae are 1.75 mm.
long.

At 30 mm. the young weakfish have attained
most of the structural characters of the adult.
But they continue much deeper and more flat-
tened sidewise until they are 6 to 8 inches long;
the head and eyes are relatively larger; and their
caudal fin is obtusely pointed with the center
rays much the longest, instead of concave. The
smaller fry (lK to 3 inches) are marked with four
dark, saddle-shaped patches extending downward
on the sides to a little below the lateral line, which
are not lost until a length of about 4% inches is
reached. As the young fish grow, other bands of
pigment are interpolated below the lateral line,
the adult coloration not being fully developed until
they are 7 to 8 inches long.77

Bate of growth. — Weakfish fry grow at so variable
a rate during the first summer that they may be
anywhere between 4 inches and 6 inches long in the
fall, when they are about 6 months old. The
smallest fish seen in spring (no doubt yearlings) are
8 to 10 inches long. Thereafter the rate of annual
growth is slower. But the variation in the length
attained by the fry during their first summer and
autumn, consequent on the protracted spawning
season, combined with the fact that scale studies
of this species have proved puzzling, make it diffi-
cult to group the older age classes by size. As far
as known, a weakfish of 10 to 12 inches is likely to
be about 2 years old; one of 13 inches, about 3
years; 15 inches, about 4 to 5 years; 18 inches, about
5 or 6 years; one of 22 inches about G to 7 years
old; 78 24 inches perhaps 9 years; and 30 inches per-
haps as old as 12 years. Both males and females
usually mature at 2 to 3 years of age, i. e., when
10 to 13 inches long.

General range. — Eastern coast of the United
States from the east coast of Florida to Massachu-
setts Bay, straying northward to the Bay of Fundy,
and perhaps to Nova Scotia.79

" Tracy (Thirty-eighth Ann. Rept. Comm. Inland Fish., Rhode Island,
1908, pp. 85-91), Eigenmann (Bull. U. S. Fish Comm., vol. 21, 1902, p. 45), and
Welsh and Breder (Bull. U. S. Bur. Fish., vol. 39, 1924, p. 154) describe the
older larvae and fry.

" According to studies by Taylor (Bull. U. S. Bur. of Fish., vol. 34, 1916,
p. 318); by Welsh and Breder (Bull. U. S. Bur. of Fish., vol. 39, 1924, p. 158);
and by R. A. Nesbit, formerly TJ. S. Bur. Fish, (unpublished).

™ It is credited indefinitely to "Maine" by Holmes (Fishes of Maine, 1862,
p. 74); Ooode (Fish. Ind. U. S. Sect. 1, 1884, p.362), states that scattering indi-
viduals have been caught as far as the Bay of Fundy; and Halkett (Check
List Fishes Canada, Newfoundland, 1913, p. 87) mentions one as probably
caught off Nova Scotia.

FISHES OF THE GULF OF MAINE

421

Occurrence in the Gulf of Maine. — The center of
abundance for the weakfish is along the coast of
the middle Atlantic States from the Virginia Capes
to New York. It also occurs regularly as far
north and east as Cape Cod. But the stock of
weakfish fluctuates widely on the southern New
England coast, and it is only during periods of
great abundance there that weakfish appear in any
numbers in Cape Cod and Massachusetts Bays,
which may be set as the extreme northern limit
for its appearance except as a stray. In the years
when it has passed Cape Cod in appreciable num-
bers it has always been far more plentiful along
the inner side of the Cape and in Cape Cod Bay
than north of Boston, as appears from the following
statement of catches for 1906, a year of great
abundance.

Cape Cod Bay: Pounds

Provincetown 115, 789

Truro 202, 050

Brewster 137, 659

Sandwich 6, 221

North Shore of Massachusetts Bay:

Nahant so 369

Manchester 410

Only once, however, for a period of about
9 years, have there been many weakfish during
the past century and a half, even in the Cape Cod
Bay region.81 Apparently they were plentiful off
southern New England during the last part of the
eighteenth century, and to judge from fishermen's
reports weakfish were well known in Massachusetts
Bay at that time. But they vanished so com-
pletely sometime prior to 1800 that when a stray
specimen was taken at Provincetown in June 1838,
it was sent to Boston for identification. And this
disappearance evidently involved the whole north-
ern part of the range of the species, for weakfish
vanished similarly from the Nantucket-Marthas
Vineyard region sometime between 1800 and
1837. They had reappeared, however, off south-
ern Massachusetts by 1867; they were abundant
there, once more, by 1870; and one or two were
taken off Truro and Provincetown in 1884.
From then on until 1895, a few were returned
yearly from Truro, Provincetown, Plymouth, and
even from as far north as Gloucester and Man-

chester, the annual catch ranging from an odd
fish only (e. g., 1893 and 1894) to 700 or 800
pounds, at most, for Cape Cod Bay and for the
northern part of Massachusetts Bay, combined.

The catch in the Cape Cod Bay-Massachusetts
Bay region was larger for the next few years
(4,892 pounds in 1896,82 1,006 pounds in 1897,
6,046 pounds in 1898, and 11,572 pounds in 1899),
though with the catches localized chiefly on the
outer side of Cape Cod and in Cape Cod Bay, as
might be expected of a stray from the south. And
they appeared in such numbers in Cape Cod Bay
in 1900 that the catch there jumped to upward
of 108,000 pounds for that year,83 while a few
were taken even as far north as Boston Harbor
and Gloucester.

This marked the commencement of a period
of local abundance, which was entirely unexpected
(for nothing like it had been experienced since
the settlement of the country), and which (with
its equally sudden eclipse) is perhaps the most
interesting event in the history of the local
fisheries. Unfortunately definite statistics of the
catches are not available for the crucial years, but
weakfish were so plentiful in Cape Cod Bay in
1901 as to be a drug on the market; while in 1902
and 1903 the pound nets in Cape Cod Bay were
often filled with schools of large weakfish, averag-
ing about 5 pounds. So plentiful were they,
indeed, during the summer of 1903 that the traps
at North Truro alone reported 280,000 pounds.

This abundance continued through 1904, by
which time it seems to have been accepted as the
normal condition of affairs, and no longer worth
comment. But it seems to have culminated in
that summer or the next, for weakfish were
reported as less plentiful in 1906. Nevertheless,
the Cape Cod Bay traps (excluding Barnstable,
Chatham, Yarmouth, and Dennis) reported almost
half a million pounds of weakfish for that year;
the North Shore of Massachusetts Bay, 20,779
pounds, which probably was not more than half
or two-thirds of the actual total, for the returns
were incomplete. This, however, was the last
big year, for the catch north of the elbow of Cape
Cod was less than one-third as great in 1907 as

10 Twenty thousand pounds were also reported from Gloucester, but we
have reason to believe that the fish were actually caught in Cape Cod Bay;
and traps operated at Rockport and at Newburyport took no weakfish.

«> There are intimations in the writings of the early historians of New
England of similar disappearances and returns of the weakfish (Goode, Fish.
Ind. U. 8., Sect. 1, 1884, p. 363).

" Omitting the towns of Yarmouth, Dennis, Chatham, and Barnstable,
where traps have been operated on the Vineyard Sound shore as well as on
the Gulf of Maine shore line.

« Omitting the towns of Yarmouth, Dennis, Chatham, and Barnstable,
where traps have been operated on the Vineyard Sound as well as on the
Cape Cod Bay side.

422

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

it had been in 1906. And this was the beginning
of the end, for only 8,249 pounds were reported
there in 1908, 569 pounds in 1909, and 907
pounds in 1910.

We do not know of the capture of a single
weakfish that can be credited with certainty to
the outer shore of Cape Cod, to Cape Cod Bay,
or to the northern shore of Massachusetts Bay
from that time on, down to 1921 (most recent
year for which the pound net catches were
published in detail), when 21 pounds were reported
for the town of Barnstable.84 We should em-
phasize too that about the same number of pound
nets and traps have been operated from year to
year throughout this period and at about the
same general localities, so that fluctuations in
the catch did actually reflect similar fluctuations
in the stock of fish.

There is no reason to suppose that weakfish have
ever entered Cape Cod Bay in any numbers since
that time. Only one, indeed (a 5-pounder), was
recorded, from one set of 8 traps at North Truro
during the 16 years 1935-1950 ;86 another set of 2
traps at Barnstable, took only 3 weakfish during
the summer of 1950; 86 and 3 other traps at Sand-
wich, Mass., took 2 weakfish in 1948, 1 in 1949,
and none in 1950.87

We doubt whether any weakfish have reached
the northern side of Massachusetts Bay since 1909,
when 200 pounds were reported from a pound net
at Gloucester. Large landings, it is true, have been
reported as from the northern part of the Massa-
chusetts coast (Essex County) in several recent
years, ranging up to some 3,600,000 pounds in
1945. But there is no reason to suppose that any
of them were caught north of Cape Cod for we are
informed by William Royce of the Fish and Wild-
life Service that all fish taken by vessels sailing out
of Gloucester during these years were credited to
that port, irrespective of where caught or where
landed. The fish may have come from as far
south as the North Carolina winter fishery. And
this applies equally to a few that were credited to

w No catch statistics are available for the years 1912-1916, and there is no
knowing whether any of the weakfish reported for Barnstable Co., in 1919
(962 pounds) came from the northern (i. e.. Cape Cod Bay) shore.

88 Information from the Pond Village Cold Storage Co.

88 Information from John E. Vetorino, who operates these traps. One
hundred twenty-three pounds reported from Barnstable County In 1928, and
101 pounds in 1929, may likely have come from the Vineyard Sound shore, not
from the Cape Cod Bay shore.

17 Information from Benjamin Morrow, who operates these traps.

Maine in 1931 (45 pounds) and in 1932 (318
pounds).

We can offer no explanation for this unexpected
invasion of weakfish north of Cape Cod about the
turn of the present century, or for its equally
sudden eclipse, the opportunity having passed long
since for obtaining any information as to the sizes
and ages of the fish, as to their movements, and as
to the physical state of the water at the time. It
was not a local event, however, but part of a
corresponding fluctuation in the population as a
whole existing east and north of New York. Thus
the catch for the southern coast of New England
was more than eight times as great in 1904 (upward
of 7 million pounds) as it had been in 1889 (about
830,000 pounds), but thereafter declined so
markedly that in 1908 both the commercial fisher-
men and the anglers of Rhode Island and of
southern Massachusetts complained of the scarcity
of weakfish. Less than 400,000 pounds were
taken off southern New England in 1919, and the
weakfish had so nearly vanished from the southern
shores of Massachusetts by 1920 and 1921 that the
reported catches for the pound nets of the State
were only 785 and 691 pounds, respectively, for
those years.88 We should emphasize that the
partial recovery that then took place off the
southern Massachusetts coast, where the average
catch was again nearly a quarter of a million
pounds during the period 1931-1938, did not bring
the weakfish back to Cape Cod Bay.

It has often been suggested that weakfish are
plentiful when bluefish are scarce, and vice versa,
and the argument has been advanced that the
latter not only devour fry of the weakfish but its
food also, and hence not only destroy many but
drive others away. But no convincing evidence
has been brought forward that the fluctuations of
these two species of fish are mutually dependent
in any way.

Importance. — At the present time the weakfish
is of no importance in the Gulf of Maine, whether
commercially or to the angler, though it was a very
valuable addition to the shore fisheries of Cape Cod
Bay during its one brief period of plenty there.
However, it is one of the most important of food
fishes along more southern coasts,89 and a favorite

88 No statistics are available for the years 1922-1929.

89 In 1946, the reported catch of weakfish of this species was about 3,252,000
pounds for southern New England; 11,715,000 pounds for the Middle Atlantic
States: 20,657,000 pounds for the Chesapeake Bay region; and 4,770,000 pounds
for the South Atlantic States.

FISHES OF THE GULF OF MAINE

423

game fish which has been the subject of many
accounts from the angler's standpoint.

Spot Leiostomus xanthurus Lac£pede 1802
Lafayette
Jordan and Evermann, 1896-1900, p. 1458.

Description. — The spot agrees closely with the
weakfish in the arrangement and general shapes
and relative sizes of its fins, and in lacking chin
barbels. But it is a much deeper fish relatively
(body about one-third as high as it is long, meas-
ured to base of tail fin), with blunt snout instead
of pointed; it has no large canine teeth; its tail
fin is more forked; and it is marked on either side
with a conspicuous black spot close behind the
upper corner of each gill opening.

The forward (spiny) subdivision of the dorsal
fin, of 10 spines, is triangular, with rounded apex;
the posterior part, of one short spine and 30-34
soft rays, is about one-half as high vertically as
the spiny part. The caudal fin is moderately
concave. The anal fin of two short spines and
12 or 13 soft rays, has a somewhat concave mar-
gin, and the pectorals are pointed.

Color.— Bluish gray above with golden reflec-
tions, silvery below. Medium-sized fish are
marked on each side with 12-15 oblique yellowish
cross bars °° dipping obliquely forward, but these

» Dusky on preserved specimens.

fade with age. And there is a conspicuous black
spot close behind the upper corner of each gill
opening. The fins are partly yellowish, partly
dusky.

Size. — -The spot grows to a length of about 13
to 14 inches and to a weight of 1 pound 6 ounces.'1
But adults average only about 10 to 10K inches
long, and few weigh more than three-quarters of
a pound.

General range. — Inshore waters from Texas 92
to southern New England, and recorded from
Massachusetts Bay as a stray.

Occurrence in the Gulf of Maine. — The spot is
plentiful in some years as far north as New York,
while young ones are described as common in
autumn about Woods Hole. But its normal
range is bounded so sharply by Cape Cod that it
has been reported only once from the Gulf of
Maine; a single specimen, taken in Massachu-
setts Bay, November 1936.93

Kingfish Menticirrhus samtilis
(Bloch and Schneider) 1801
King Whiting; Minkfish; Whiting
Jordan and Evermann, 1896-1900, p. 1475.

n These were the longest of many measured in Chesapeake Bay by Hilde-
brand and Sehroeder (Bull. U. S. Bureau of Fisheries, vol. 43, Part 1, 1028,
p. 272). The maximum length previously recorded was 1194 inches (Nichols
and Breder, Zoologira, New York Zool. Soc , vol. 9, 1927, p. 95).

" Once reported doubtfully from Martinque.

» Reported by Qoffln, Copeia 1937, No. 4, p. 236.

Figure 218. — Spot (Leiostomus xanthurus), Rhode Island. From Goode. Drawing by H. L. Todd.

424

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 219. — Kingfish (Menticirrhus saxatilis), Pensacola, Florida. From Goode. Drawing by H. L. Todd.

Description. — The kingfish resembles the weak-
fish in the general arrangement and the relative
sizes of its fins, the second dorsal being much
longer than the first or than the anal. But its
first dorsal (10 spines) is relatively much higher
and more pointed than that of the weakfish, with
the third spine not only much prolonged but
filamentous at the tip in the adult (a noticeable
character) ; the rather blunt nose, and snout over-
hanging the mouth, give the kingfish a very char-
acteristic cast of countenance (fig. 219). Its upper
jaw, furthermore, projects beyond the lower,
whereas the reverse is the case in the squeteague.
Its chin bears a barbel, which the weakfish lacks,
its lips are fleshy, and it has no canine teeth.
Its tail, too, is of very characteristic outline, with
the lower half rounded but the upper half concave
suggesting (though not exactly parallelling) the
tail of the sea bass (p. 407). Its body is about as
slender, proportionally, as that of a squeteague,
but the kingfish carries its weight farther forward
(it is deepest below the first dorsal fin), and it
has a weak-tailed appearance remotely suggesting
a hake (p. 222). We need merely note further
that the filamentous spine of the first dorsal is
longer in large fish than in small ones; that the
second dorsal (one stout but short spine followed
by 24 to 27 rays) occupies more than one-third
of the length of the back and tapers slightly
from front to rear; that the anal fin (one long
spine and 8 rays) stands under the middle of the

soft dorsal; and the pectorals are pointed and
relatively much longer than those of the sque-
teague.

The Kingfish and its immediate relatives have
no air bladder, hence makes no sounds, in which
they differ from other members of their family.

Color. — Leaden or dusky gray above, (some-
times so dark as to be almost black) with silvery
and metallic reflections; milky or yellowish-white
below. The sides are cross marked irregularly
with dark bars. These run obliquely forward and
downward behind the spiny dorsal fin, but the
foremost one or two bars run in the opposite direc-
tion, so that they form a V-shaped blotch or two
dark V's below the fin. The pale belly is bounded
above by a dark longitudinal streak on either side.
The fins are dusky or blackish; the first dorsal fin
anal, pectorals, and ventrals are tipped with dirty
white.

Size. — Kingfish grow to a maximum length of
17 inches and a weight of about 3 pounds, but the
general run are from 10 to 14 inches long, weighing
y2 to \)'i pounds.

Habits. — Kingfish, like squeteague, are summer
fish, appearing on the coast in May, to vanish in
October. They are confined to the immediate
vicinity of the coast during their stay, frequenting
inclosed as well as open waters, even entering
river mouths, and they are unknown on the off-
shore banks. They run in schools, keep close to
the ground, prefer hard or sandy bottom, and feed

FISHES OF THE GULF OF MAINE

425

on various shrimps (perhaps their chief diet),
crabs, and other crustaceans, small mollusks,
worms, and on young fish.

Breeding habits. — Kingfish spawn in bays and
sounds from June until August, but it is not likely
that any young that might be hatched in the Gulf
of Maine from eggs laid by the occasional visitors
would survive its low temperature. Welsh and
Breder M describe the spawning and early develop-
ment of this species. Young fry of % to 1-inch
already show most of the structural characters of
the adult, including the scales, and so are readily
recognizable as kingfish though they vary widely
in color, ranging from the pattern of the adult to
almost uniform blackish brown. Welsh and
Breder found from an examination of the scales,
confirmed by a large series of measurements, that
kingfish are 4 to 6 inches long by the first winter,
average about 10 inches the second winter, and
13% the third. Many males ripen when 2 years
old, but few females until 3 years old.

General range. — Atlantic coasts of the United
States from Florida (Pensacola, Key West) north-
ward regularly to Cape Cod ; most numerous from
Chesapeake Bay to New York; known as far north
as Casco Bay, Maine, as a stray.

Occurrence in the Gulf of Maine. — This excellent
food and game fish reaches the Gulf of Maine only
as a stray from the south. So far as we can learn
the only positive records of it within our limits are
as follows, south to north: Monomoy and North
Truro on Cape Cod in 1896 (collected by Dr. W. C.
Kendall); one taken at Provincetown, July 1847,
another there in November of that same year and
many small ones, apparently chilled by the cold,
that appeared in that harbor in 1879; one taken
at the entrance of Boston Harbor in a lobster pot
some time before 1833; one at Lynn in 1840; one
8 inches long off Marblehead on October 15, 1872;
one of 6}i inches at Danvers, October 28, 1874;
others at Nahant (one record),95 and in Casco Bay.

Catch statistics, if taken at face value, would
suggest that kingfish reached the northern shore
of Massachusetts Bay (Essex County) in unprece-
dented numbers during the period 1931 to 1938,96

" Bull. U. S. Bur. Fish., vol. 39, 1924, pp. 191-194.

•' Small amounts of "kingfish" appear In tho pound-net returns published
by the State of Massachusetts at various localities in Massachusetts Bay, but
Sshermen Inform us that these were not the true kingfish but some large
jpecies of the mackerel tribe.

•• For Essex County, Mass., 2,029 pounds reported during 1931; 34,981
pounds for 1933; 5,100-10,600 pounds for 1933, 1935, 1937 and 1938.

but we are informed by William Royce of the
U. S. Fish and Wildlife Service that all the fish
taken by vessels sailing out of Gloucester during
these years were credited to that port, irrespective
of where caught or where they were landed.
There is no reason to suppose that any of these
kingfish or "king whiting" actually came from as
far east or north as Cape Cod, or even from any-
where in southern New England waters for that
matter. And this applies equally to 466 pounds
reported in 1932 from Maine. It is, in short,
an unusual event for a kingfish to round the
elbow of Cape Cod, or for a small school of its
fry, nor have we heard of any taken anywhere in
the Gulf during recent years.

Importance. — The kingfish is not plentiful
enough in the Gulf to interest either commercial
fishermen or anglers. It is one of the better table
fishes, and a favorite with surf anglers along the
coasts of New York, New Jersey, and southward,
as it bites readily and fights well. In the words
of a well-known angler, "no fish that swims the
sea makes a better dish. Certainly no bottom
living fish plays such a game for the angler's real
delight." 97

Black drum 98 Pogonias cromis (Linnaeus) 1766

Jordan and Evermann, 1896-1900, p. 1482.

Description. — A short deep body (less than
three times as long as it is deep to the base of the
caudal fin) with high-arched back but flattish belly
is characteristic of the drum. The profile of the
face is even more diagnostic, for the mouth is
horizontal and set very low, the eye high, and the
chin bears several barbels. The arrangement and
sizes of the fins are essentially the same as in the
weakfish, except that the second (soft-rayed)
dorsal is relatively shorter, and that the anal spine
is much stouter. The jaw teeth are small and

" Bhead, Bait Angling for Common Fishes, 1907, p. 145.

11 The channel bass or red drum Sciaenops ocellatu) (Linnaeus) 1766, a south-
ern sciaenid uncommon east or north of New York, is represented in the
collection of the Boston Society of Natural History by a mounted specimen
labeled "near Portland, Me.," but as this fish was probably purchased
in the market, it is likely that it had been shipped from the south than that
it was actually caught nearby. Should this drum ever be taken in the Oulf
of Maine, its relationship to the weakfish, kingfish, and spot would be ap-
parent from the arrangement of its fins, especially from the shortness of the
anal fin relative to the soft (second) dorsal. But it is easily distinguished
from the weakfish by the fact that its upper Jaw extends beyond the lower
Instead of vice versa; from the spot, by the barbels on its chin; and from the
kingfish by having several of these barbels instead of only one; by the shape
of its tall fin; and by the presence of a conspicuous black blotch (sometimes
as many as 4 or more blotches) on each side at the base of the caudal fin
which affords a ready field mark for its identification.

426

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 220. — Black drum (Pogonias cromis). From Goode. Drawing by H. L. Todd.

pointed, but the throat is armed with large, flat,
pavement-like teeth with which the drum crushes
shellfish for food, a character separating it from
its allies the weakfish and the kingfish. The first
dorsal fin (10 spines) is rounded-triangular; the
second (1 short spine and 20 to 22 rays) oblong;
the caudal is square-tipped with moderately high
peduncle; the anal fin (2 spines, the first very short
and the second long and stout, and 6 or 7 soft rays)
is less than half as long as the soft dorsal; the
pectorals are sharp pointed and relatively longer
than those of the weakfish. The second anal spine
is much stouter in young drums than in old ones.
The eyes of the drum are comparatively small and
its scales are large.

Color. — Silvery with a brassy lustre, turning to
a dark gray after death. Young fish have 4 or 5
broad dark vertical bars that fade out with age.
The fins are blackish. This drum occurs in two
color phases, a grayish and a reddish.

Size. — Drums grow to a huge size. The largest
we find positively recorded (caught in Florida)
weighed 146 pounds; adults, as caught, run from
20 to 40 pounds, with 60 pounds not exceptional.
The rod and reel record is 87 pounds 8 ounces, a
fish 4 feet 4 inches long, caught at Cape Charles,
Va., May 6, 1950, by Mrs. H. A. Bradley, Jr.
A fish 40 inches long weighs about 40 pounds.

General range. — Atlantic and Gulf of Mexico
coasts of America from Argentina to southern New
England; common from New York southward and
abundant from the Carolinas to the Rio Grande; a
stray visitor as far north as Massachusetts Bay.

Occurrence in the Gulf of Maine. — -This southern
fish is decidedly uncommon east of New York;
occasional specimens only have been reported from
Woods Hole; and it is only a stray visitor to our
Gulf, where 2 or 3 specimens have been taken at
Provincetown, and 1 in the Mystic River, which
empties into Boston Harbor.

THE TILEFISHES. FAMILY BRANCHIOSTEGIDAE

The tilefishes are sea-bass-like in appearance,
but with the soft (rear) portion of the dorsal fin
much longer that the spiny forward part, and the
ventral fins are under the pectorals (thoracic).
The only species that occurs off the northeastern
United States is characterized by a large fleshy
flap on the nape, suggesting (though not corre-
sponding to) the adipose fin of salmons and smelts.
But this adipose flap or fin is not shared by its
relatives.

Tilefish Lopholatilus chamaeleonticeps Goode and
Bean 1879

Jordan and Evermann, 1896-1900, p. 2278.

Description. — The presence of a thin, high,
fleshy, finlike flap on the nape of the neck in front
of the dorsal fin, close behind the eyes, suggesting
the adipose fin of the salmon tribe in its appearance
though not in its location, serves to identify the
adult tilefish at a glance among Gulf of Maine

FISHES OF THE GULF OF MAINE

427

Figure 221. — -Tilefish (Lopholatilus chamaeleonticeps) , off Marthas Vineyard. From Goode. Drawing by H. L. Todd

fishes. In grown fish this flap is as high as the
dorsal fin, higher than long, and rounded at the
tip. In small fry it is relatively much lower.
Equally distinctive, if less conspicuous, is a smaller
fleshy flap situated on the side of the lower jaw
close to the angle of the mouth, pointing backward
(to be seen in the illustration, fig. 221). The large
head is strongly convex in dorsal profile but nearly
flat in ventral profile, with the eye high up, the
mouth wide, and both the jaws are armed with
an outer series of large conical teeth and inner rows
of smaller teeth. The trunk (moderately flattened
sidewise) is deepest close behind the head, tapering
thence backward to the sidewise-flattened caudal
peduncle. The spiny and soft portions of the
dorsal fin are continuous, extending back from
above the gill opening almost to the base of the
caudal fin, as is the case in cunner, tautog, and
rosefish. But the anal fin (14 or 15 rays) of the
tilefish is about half as long as the dorsal fin, under
the rear (soft-rayed) part of which it stands,
and like the latter it is of nearly even height
throughout most of its length except that its
forward corner is rounded. The ventral fins are
located below the pectorals, which are set low
down on the sides, and both the pectorals and the
ventrals are pointed. The gill covers, as well as
the trunk, have moderately large scales.

Color. — This is a brilliant fish, bluish or olive
green on the back and on the upper part of the
sides, changing to yellow or rose lower down on
the sides; its belly is of the latter tint with white
midline. The head is tinged reddish on the sides;
pure white below. The back and sides above the
level of the pectorals are thickly dotted with

small irregular yellow spots, which are particu-
larly conspicuous below the adipose dorsal flap.
The dorsal fin is dusky, marked with similar but
larger yellow spots, its soft-rayed portion pale
edged. The adipose flap is greenish yellow; the
anal fin pale pinkish clouded with purple and with
bluish iridescence; and the pectorals are pale
sooty brown, with purplish reflections near their
bases.

Size. — Tilefish have been reported up to 50
pounds in weight, but this is unusual. The largest
fish we have seen (an unripe female) weighed
35% pounds and was about 42 inches (108 cm.)
long. Measurements taken by Bumpus" and
more recently by us on the Grampus show that a
40-inch fish may be expected to weigh about 30
pounds; fish of 33 to 36 inches, 20 to 21 pounds;
and 30- to 32-inch fish, 17 to 18 pounds.

Habits. — This is a bottom fish, and its depth
range off our Atlantic coast is a very narrow one,
none ever being taken shoaler than about 45
fathoms,1 and very few much deeper than 100 fath-
oms. The deepest definite record with which we
are acquainted is 170 fathoms (p. 428), and with
the best fishing at 60-90 fathoms. In the Gulf of
Mexico it has been caught at 90 fathoms. The
thermal range to which the tilefish is exposed,
normally, is very narrow also, for the temperature
of the bottom water along the zone inhabited
regularly by it varies only between about 47° and
about 53°, in most years, summer or winter.
And it appears to be very sensitive to chilling;

<> BulJ. U. S. Fish Comm., vol. 18, 1899. p. 329.

• The shoalest we have known any to be trawled was at 43-17 fathoms, by
Albatross III, 35 miles southwest of Nantucket Lightship in mid-May 1950.

428

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

this having been the probable cause of a mass
destruction of tilefish that took place in 1882 (for
further discussion, see p. 429). It is not known
whether the tilefish is equally sensitive to high
temperatures, in any case it could escape such by
descending to a greater depth.

Food. — A great variety of bottom-dwelling
invertebrates have been taken from tilefish
stomachs. Crabs, of which they are often packed
full, are the most important article of diet. The
list 2 also includes squid, shrimp, shelled mollusks,
annelid worms, sea urchins, sea cucumbers, and
sea anemones. Occasionally they catch other fish;
two spiny dogfish, for instance, were found in one,
and an eel (probably a conger or a slime eel) and
unidentified fish bones in others.3 The presence
of pelagic ampbipods (Euthemisto)* and of salpae
in the stomachs of tilefish caught on long lines
proves that they sometimes feed at higher levels,
but they are never known to rise to the surface
voluntarily, and when they are hauled up they
are often "poke blown." Tilefish take any bait,
perhaps menhaden best, salt herring not so
readily.

Although they are strong active fish, it is
probable that they suffer from the attacks of
sharks, for fish caught on the long lines are often
bitten in two. And we have seen numbers of sharks
7 to 8 feet long (species not determined) following
them up to the surface, while the line was being
hauled.

Ever since the tilefish was discovered it has
been known to spawn in July, and eggs were
running from 10 out of 11 females caught by the
Grampus off New York on August 3, 1916, while
the roe of the eleventh was still unripe. How
early the spawning season may open is still to be
learned, but August probably sees its close, for
the majority of 18 females caught on the 26th of
that month in 1914 were spent, only one or two
still having running eggs. Among the fish that
we have examined, the females have greatly
predominated (only 1 male to 29 females in a
total of 39 individuals).

Ripe eggs taken from a tilefish and preserved
in formalin measured about 1.25 mm. in diameter.6

« Linton, Bull. U. S. Fish Comm., vol. 19, 1901, p. 47; Notes by Vina]
Edwards', and our own observations.

» The menhaden credited to the diet of the tilefish by Sumner, Osbum, and
Cole (Bull. U. S. Bur. Fish., vol. 31, Pt. 2, 1913, p. 767) were merely the
pieces ol bait on which the fish had been caugbt.

• Collins, Rept. U. S. Comm. Fish. (18S2) 1884, p. 244.

» Eigenmann, Bull. U. S. Bur. Fish., vol. 21, 1902, p. 37.

As they had an od globule of 0.2 mm., it is safe
to say that they are buoyant, and tow nettings
yielded eggs, indistinguishable from those stripped
from the tdefish, at the station where we caught
the ripe females just mentioned. But the larval
stages have not been seen. The fact that a few
tdefish of 2}i to 3% inches were taken along the
outer edge of the continental slope in April 1930,
and others of 4 to 4^ inches in July, suggests that
4 to 5 inches is the usual length at one year of age.9
Nothing is known of the subsequent rate of
growth, nor at what age the tdefish matures
sexually.

General range. — Outer part of the continental
shelf and upper part of the continental edge off
Nova Scotia and off the North and Middle Atlantic
United States, from Banquereau Bank to the
offing of Chesapeake Bay, in depths of 45 fathoms
to perhaps 200 fathoms; also reported from
southern Florida in more than 100 fathoms,7
and from the Campeche Bank in the southern side
of the Gulf of Mexico, whence the Museum of
Comparative Zoology has received a specimen
taken in 90 fathoms by the schooner Seminole on
February 1, 1946,8 and where local fishermen
report that they have taken a number.

Occurrence off Nova Scotia and off the North and
Middle United States. — The most easterly and
northerly records for the tilefish are of a small one
caught on Banquereau Bank (lat. 44°26' N.,
long. 57°13' W.) in 170 fathoms, December 15,
1902, from the schooner Monitor out of Gloucester,9
and of another of 4}i pounds that was brought in
to Boston in 1933.10

Its chief center of abundance is between the
offings of Nantucket and of Delaware Bay.
And there is some evidence that it ranges farther
east in warm years than in cold. In 1908, for
example, tdefish were caught off the South Channel
(long, about 69°) in September, whde in 1950
the Albatross III trawled a few at 50-80 fathoms
nearly that far east (at longitudes 69°57' to
69°35' W.) in May, whereas the Grampus caught
none off Martha's Vineyard (long, between 70°
and 71° W.) in the very cold July of 1916, but
made a fair catch off New York.

• For details, see Schroeder, Bull. 58, Boston Soc. Nat. Hist., 1931, p. 7.

' One of 23 pounds, caught off Key West in more than 100 fathoms, is
reported by Al Pfleuger, well-known fish taxidermist of Miami.

■ Taken by the schooner Seminole on February 1, 1946. See Bigelow and
Schroeder, Copela, 1947, pp. 62-63, for details.

• Reported by Evormann, Rept. U. S. Comm. Fish. (1903), 1905, p. 85.
'» Reported to us by J. Webster of the U. S. Bureau of Fisheries.

FISHES OF THE GULF OF MAINE

429

On the other hand, none have been reported
alive off the Atlantic coast below lat. 37°29' N., a
few miles north, that is, of the mouth of Chesa-
peake Bay, which makes it likely that the tilefish
of southern Florida and of the Gulf of Mexico are
isolated populations.

The onshore-offshore range of the tilefish off our
northern coasts, being limited in depth (p. 427),
is confined to a bottom belt only some 15 to 25
miles wide — astonishingly narrow for so large a
fish and one that is so plentiful. And presumably
it is a year-round resident wherever it is found
there, for its presence has been established north-
ward to the offing of southern New England as
early in the season as March, and as late as
January, while there was no general falling off
in the catches in autumn and early winter during
the only year (1917-1918) for which monthly
data are available.

Though the tilefish has been reported only once
well within the limits of our Gulf, its history and
its relationship to hydrographic factors are so
interesting that it deserves more attention than
its status as a Gulf of Maine fish would warrant
otherwise.

It is astonishing that the very existence of so
large a fish so close to our coast should have
remained unsuspected until May 1879, when
Captain Kirby, cod fishing in 150 fathoms of
water south of Nantucket Shoals Lightship,
caught the first specimens. Others were caught
at 87 fathoms nearby by the schooner Clara T.
Friend (Capt. William Dempsey) during the fol-
lowing July. And trips by the United States Fish
Commission during the next two summers proved
that the tilefish were plentiful enough to support
an important new fishery. These early investi-
gations likewise proved that it occupies a very
definite environment, along the upper part of
the continental slope and on the outer edge of the
shelf where a narrow band of the sea floor is
bathed with a belt of warm water (about 47° to
53°), varying by only a couple of degrees in tem-
perature from season to season, and that it never
ventures into the lower temperatures on the
shoaling bottom nearer land, nor downward into
the icy Atlantic abyss. The balance, in fact,
between the physiological nature of the tilefish
and its surroundings is so delicate that catastrophe
overtook it within three years of its discovery.
The first news of this disaster came in March

1882; throughout that month and the next vessel
after vessel reported multitudes of dead tilefish
floating on the surface throughout the entire zone
inhabited by it north of Delaware Bay, and it has
been estimated that at least a billion and a half
dead tilefish were sighted.11

It has generally been believed that this destruc-
tion was caused by a temporary flooding of the
bottom along the warm zone by abnormally cold
water.12 Consonant with this is the fact that other
species of fish suffered too, and dredgings carried
on during the following autumn proved that the
peculiar invertebrate fauna that had been found in
abundance along this warm zone in previous
summers had likewise been exterminated.

The destruction of the tilefish was so nearly
complete that fishing trials carried on off southern
New England by the Fish Commission later in
1882; in 1883; 1884 (when a particularly careful
search was made); 1885; 1886; and 1887 did not
yield a single fish. But the species was not quite
extinct, as the Gram-pus proved by catching 8 of
them off Marthas Vineyard in 1892, and 53 in
1893. Tilefish were next heard of in 1897 when
a fishing schooner caught 30 fish of 6 to 15 pounds,
while long-lining for haddock south of Marthas
Vineyard. And tilefish had become so numerous
again by 1898 that the Grampus caught 363
fish, of K to 29 pounds, on three trips of only 1 to
3 days' duration each.

The length of the period which the fish required
to reestablish itself after the mortality of 1882,
together with the fact that in 1898 the catch
included a considerable number of young fish, is
evidence that the replenishment of the stock was
chiefly the result of local reproduction, though
it may have been been recruited to some extent
by immigration from the southern part of the
range, where destruction may not have been so
complete as it was north of Delaware Bay.

The tilefish was kept in view during the next 17
years by occasional trips to the grounds by the
Bureau's vessels. We caught 19, for example,

u Collins (Rept. U. S. Comm. Fish. (1882], 1884, pp. 237-294A) has described
the event in detail, as have many subsequent authors. An account will also
be found in Economic Circular No. 19 of the U. S. Bureau of Fisheries.

u No temperatures were taken on the tilefish ground at the season when the
mortality occurred; and the bottom water was nearly as warm there by the
end of the following August (48°-49°) as it usually is (about 50°-52°). The
temperatures taken in this region during the early years of the Bureau of
Fisheries are discussed elsewhere (Bigolow, Bull, Mus. Comp. Zool., vol. 69,
1915, pp. 238-241.)

430

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

weighing about 350 pounds, on the Grampus on
August 26, 1914, in a set of one hour off Marthas
Vineyard in 105 fathoms. In 1915, the Bureau
undertook to popularize the tilefish in the market,
believing it numerous enough to support an im-
portant fishery, and knowing it to be an excellent
food fish. It proved so plentiful and so easily
caught on long lines that the first trip stocked
38,383 pounds in 27 days. A.nd the landings for
the first 8 months after the inception of the fishery
aggregated upward of 4,388,500 pounds, with a

grand total of 11,641,500 pounds from July 1, 1916
to July 1, 1917. But for some reason the demand
did not hold up; the catches diminished; and in
1947 (most recent year for which information is
available) only 441,000 pounds were landed.13
The tilefish continues, however, to offer a potential
supply of perhaps two to three million pounds
yearly, of fish that is good boiled or baked, and that
is delicious for chowder. It also makes a good
smoked fish, and its sounds are of value for
isinglass.

THE ROCKFISHES. FAMILY SCORPAENIDAE

The rockfisb.es are perch-like or bass-like in
general appearance. But they are related to the
sculpins (p. 439) and to the sea robins (p. 467) by
having a bony stay (an extension of one of the
suborbital bones) stretching across the cheek,
giving the latter a characteristic bony appearance.
Furthermore their cheeks are spiny, and in most
of the species the top of the head is marked by
ridges that terminate in spines. Both the spiny
portion and the soft portion of the dorsal are
well developed, either as a continuous fin or sub-
divided by a deep notch. The ventral fins are
on the chest ("thoracic"). In most of the rock-
fishes (including the rosefish) the eggs are retained
within the mother until they hatch. There are
many species, the temperate Pacific being espe-
cially rich in them. Only one, however, occurs
regularly in the Gulf of Maine, though the range
of another includes its offshore rim.

Key to Gulf of Maine Rockfishes

1. The lower rays of the pectoral fins, like the upper rays,
are connected nearly to their tips by the fin mem-
brane: There are 14 or 15 dorsal fin spines

Rosefish, p. 430

The lower 7-9 pectoral fin rays are free for the outer

half of their length; there are only 12 dorsal fin

spines Black-bellied rosefish, p. 437

Rosefish Sebastes marinus (Linnaeus) 1758

Ocean perch; Redfish; Red sea perch; Red
bream; Norway haddock

Jordan and Evermann, 1896-1900, p. 1760.

Description. — The rosefish is perchlike in its
general appearance, moderately flattened side-
wise, about one-third as deep as it is long (to
base of tail fin), with a large bony head; and its
trunk tapers back from the shoulders to a moder-

ately slender caudal peduncle. The dorsal pro-
file of the head is concave, the mouth is large, very
oblique, and gapes to below the eyes, the lower
jaw projects beyond the upper, and there is a
bony knob at its tip that fits into a corresponding
notch in the upper jaw. Both of the jaws are
armed with many small teeth. The eyes are very
large and set high. The sides of the head are
armed with spines, the most prominent of which
are two near the rear angle of each gill cover, and
a series of five confluent ones on each cheek.
These, with a ridge behind and above each eye
socket, give the head a bony appearance that is
extremely characteristic.

The gill openings are very wide, with pointed
gill covers. There is one continuous dorsal fin
running from nape of neck to caudal peduncle;
the spiny part (14 or 15 spines) is considerably
longer than the soft part (13 to 15 rays), but the
latter is higher than the former. The precise
outline of the fin is easier illustrated (fig. 222)
than described. The anal fin, consisting of three
graduated spines and 7 or 8 longer rays, is shorter
than the soft portion of the dorsal, under which it
stands. The caudal fin is noticeably small, its
rear edge moderately concave, and with angular
corners. The pectoral fins are very large, and the
smaller ventrals are situated below them. Both
head and body are clad with scales of moderate
size. There are about 60 to 70 oblique rows of
scales from the gill opening to the origin of the
caudal fin, just below the lateral line.

The rosefish agrees with the cunner, tautog, and
sea bass in the union of the spiny and soft portions
of its dorsal into a single long fin, and in its gen-

" 62,700 pounds in Massachusetts ports; 128.400 pounds in Rhode Island
and Connecticut; 186,700 pounds in New York and 53,300 pounds in New
Jersey.

FISHES OF THE GULF OF MAINE

431

Figure 222. — Rosefish (Sebastes marinus), Eastport, Maine. From Goode. Drawing by H. L. Todd.

erally perch-like conformation. But it is separable
from the first two by its much larger mouth, spiny
head, large eyes, more slender caudal peduncle,
and larger pectorals ; and from the sea bass by its
large spiny head, by the shape and small size of
its caudal fin, and by the fact that its anal fin
and the soft portion of its dorsal are relatively
much lower. Its brilliant red color is a sufficient
field mark.

Color. — Orange to flame red, occasionally gray-
ish red or brownish red, with the belly a paler red
that fades to white after death. The black eyes
contrast vividly with the brightly colored body.
Medium sized rosefish usually have a dusky blotch
on each gill cover, and several irregularly broken
dusky patches along the back. These dark mark-
ings are more conspicuous on small fish, and young
fry up to 3-4 inches long are only faintly reddish,
if at all so.

Size. — The rosefish matures sexually when 9 to

10 inches long, males when a little smaller than
females. In the Gulf of Maine they are said to
grow ordinarily to a maximum length of perhaps
2 feet. The largest measured specimen taken
recently in the Gulf of Maine of which we have
heard was 22 inches long, and weighed 5 pounds

11 ounces.14 The largest we have seen measured
18% inches.15 But Goode 16 reported one of about

» A fish landed in Gloucester, reported in Maine Coast Fisherman, Janu-
ary 1951, p. 9.

11 One of 63 specimens trawled by Albatross III on the southeastern slope of
Georges Bank at 175-195 fathoms. May 16, 1950.

» Fish. Ind. U. S. Sect. 1, 1884, p. 261.

24 inches, weighing about 14 pounds brought in
to Gloucester; a 27K-inch specimen has been
reported from the southern edge of the Newfound-
land Bank, near the Whale Deep.17 Another
27-inch fish, said to have weighed 13% pounds was
landed in Gloucester by the dragger Estaela on
February 7, 1951, from somewhere off Newfound-
land. And rosefish grow even larger (maximum
about 31-32 inches) in the other side of the
Atlantic and in Arctic Seas.18

The rosefish run smaller near the coast (usually
8 to 12 inches long) than on the offshore banks.
In European waters, where a similar size-rela-
tionship obtains, the small inshore form represents
a separate species (Sebastes viviparus) for it has
many fewer scales than the larger, offshore form
(S. marinus). But no racial distinctions have
been found between the inshore populations and
those offshore among the American rosefish.

The relationship between length and weight
runs about as follows for Gulf of Maine rosefish:
9% inches, % pound; 12 inches, 1 pound; 15 inches,
2 pounds; 17-18 inches, 2^-3 pounds; 20 inches,
perhaps 4 pounds.

Habits. — The young rosefish drift in the upper
and intermediate water layers (p. 435) until they
are nearly an inch long. Fish upwards of a

i' This specimen, reported by McKenzie. (Proc. Nova Scotian Inst.
Sci., vol. 20, 1940, p. 44) was said to have weighed 7H pounds dressed, ap-
parently an error, unless the fish was very thin.

" According to Saemundsson (Faune Ichthyol., Cons. Intemat. Explor.
Mer. 1932, plate). A length of 100 cm. (about 40 inches) has been stated,
but we are inclined to doubt this.

432

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

couple of inches long tend to hold close enough
to the bottom in our Gulf for great numbers of
them to be caught in otter trawls. But some may
also live pelagic over the deep basins as they are
known to do in the Gulf of St. Lawrence; also,
in the Norwegian Sea, where there is a population
of all sizes living mostly at depths of about 50
to 100 fathoms, over much greater depths.19
When they are on bottom the rosefish are chiefly
on rocky or hard grounds or on mud, seldom on
sand, if ever. Their depth range on the bottom
is from within a few feet of tide line (p. 434) down
to 350 fathoms at least; perhaps to 400 fathoms
(p. 434) with the greater part of the commercial
catch trawled at about 40 to 175 fathoms; and
fry, living pelagic, have been taken as deep as 270
fathoms in north European waters.

Our rosefish inhabit a wide range of temperature.
The maximum may be set at about 48° to 50° F.,
and probably it is the low temperature of parts of
the Bay of Fundy, where the upper 10 fathoms or
so may be as cool as 50°-52° even in midsummer
that allows them to remain in shoal water there
the year round (p. 435). At the other extreme
they winter in Massachusetts Bay and in Passa-
maquoddy Bay in water as cold as 33° to 35°, and
perhaps colder, though they could easily avoid
these low temperatures by a short offshore migra-
tion. In fact, the rosefish has often been described
as an Arctic species. But while this is true to the
extent that its range extends to Arctic Seas, it is
a misnomer if taken to mean that it is character-
istic of Polar temperatures, for the records of its
occurrence, horizontal and bathymetric, prove
that the great majority of them inhabit waters
warmer than 35°-36° over the greater part of their
geographic range.

The distribution of the rosefish 20 in the Gulf of
St. Lawrence is especially instructive in this
respect, for it inhabits the comparatively warm
water (39° to 42° F.) in the bottoms of the deep
channels, and not the icy intermediate layer (about
32°) which, generally speaking, is so nearly an
impassable barrier to its upward migration that
it is seldom if ever taken on the shoal banks.
And its vertical range in relation to temperature
seems to be much the same as this off the south-

west coast of Greenland, where rosefish are taken
chiefly deeper than 90 fathoms, in water of about
37°-39°, not in the icy layer above, and where
numbers of them (says Jensen) sometimes come
to the surface dead in winter, apparently having
succumbed to cold.21 In the Norwegian Sea, how-
ever, rosefish of this species are caught only in the
overlying layer of water of Atlantic influence at
temperatures of 37°-39° or higher, never deeper
in the icy cold Polar water.

Temperatures of 37°-39° are the lowest in which
young rosefish are born in any numbers in our
Gulf; there is no water there colder than this by
the time production is well under way, say late
June or early July. At the opposite extreme,
practically the entire production of rosefish takes
place in water colder than 46°-48°, this being the
maximum to which the water warms at the 20-
fathom level and deeper, except in regions of active
vertical mixing where the temperature may rise a
degree or two higher. In the Gulf of St. Lawrence,
rosefish have been found breeding in 39°-42°.
Cursory examination of station data might suggest
that young are born in colder water on the Grand
Banks as well as along the south and east coasts
of Newfoundland, for they have been taken there
in tow nets at many localities where the tempera-
ture was lower than 32°, either on the bottom or
at some intermediate depth. But it is more likely
that the parent fish, and the young fry also, were
living above this icy layer, not in it; i. e., in water
at least as warm as about 35° (1.5° C), and
warmer than about 36°-37° for the most part.

Thus the range of temperature within which
American rosefish fry are produced in one place or
another is from about 37° to 47° or 48°, which is
about the same as for north European waters.22
In fact it is not likely that rosefish breed success-
fully in temperatures lower than 35° anywhere in
either side of the Atlantic.

The salinity in which rosefish breed in our Gulf
is as definitely limited in one direction as is the
temperature, if not in the other, for its young are
produced for the most part in salinities upward of
32 per mille.

19 For studies of the pelagic occurrence of S. marinus in northeastern
Atlantic waters, see Murray and Hjort (Depths of the Ocean, 1912, pp.
647-648) and especially Taning (Journal du Conseil, Cons. Intemat. Explor.
Mer., Vol. 16, 1949, No. 1).

» Huntsman, Trans. Roy. Soc. Canada Ser. 3, vol. 12, Pt. 4, 1918, p. 63.

" See Jensen (Vid. Meddel. Dansk Naturhist. Foren. Copenhagen, vol. 74,
1922, pp. 89-109, for an interesting study of the occurrence of the rosefish in
Greenland waters.

» See Taning (Journal du Conseil, Cons. Internat. Explor. Mer, vol. 16,
No. 1, 1949) for a recent discussion of the thermal relationships and breeding
range of S. marinus.

FISHES OF THE GULF OF MAINE

433

Food. — The diet of the Gulf of Maine rosefish
includes a great variety of crustaceans, especially
mysid, euphausiid, and decapod shrimps; small
mollusks; and various other invertebrates, and
small fish.23 It bites on almost any bait. In
turn, it is the prey of all the larger predaceous
fish, its fry being devoured in quantity by cod,
by older rosefish, and by halibut.

It has long been known that the eggs of the
rosefish develop and hatch within the oviduct
of the mother, and the number produced by large
females may run as high as 25,000-40,000 yearly.
This is a small brood compared to the numbers
produced by many of the marine egg-laying fishes.
But the protection offered the eggs by being
retained inside the mother's body during incuba-
tion gives the young a greater chance for survival.

The larvae are about 6 mm. long at birth
(fig. 223B), with the yolk mostly absorbed, the
mouth already formed, and the first traces of
the caudal rays already visible. At a length of
12 mm. (fig. 223D) the dorsal and anal finrays have
appeared, the ventrals are visible, and the head
spines are prominent. And though the red color
is not developed until the little fishes are about
to take to bottom, or later, all but the very young-
est larvae are recognizable as rosefish by their
large spiny heads, large eyes, short tapering bodies,
very short digestive tract, and by the presence
of two rows of post anal pigment cells, a dorsal
and a ventral row.

This is a very slow-growing fish. Available
information is to the effect that they average
about 2}'2 inches when 1 year old.24 Studies of
the scales of rosefish of different sizes 25 indicate
that 5-inch fish are likely to be 4 years old;
6-inch fish, 6 years old; 7-inch fish 7 or 8 years
old; 8-inch fish 8 or 9 years old; 9 -inch fish 9 or
10 years old, and that many of the largest fish
of 18 inches and upward may be 20 years old,
or older. Thus the mature fish are 8 to 9 years
old and older, with the greater part of the com-
mercial catch 10 years old and upward. And
about as slow a growth rate has been reported

for the immature rosefish of this same species
of Barents Sea, on the other side of the Atlantic.28

>3 Most of the rosefish that we have seen trawled had voided their stomach
contents before they were brought on board.

* According to the sizes of young rosefish collected by us in the Gulf of
Maine, May to August in various years.

» By Perlmutter and Clark, U. S. Fish and Wildlife Service, Fishery
Bull. No. 45, 1949.

Figure 223. — Rosefish (Sebastes marinus). A, egg from
the oviduct of a gravid female; B, larva, 6 mm.; C.
larva, 9 mm.; D, larva, 12 mm.; E, fry, 20 mm. Speci-
mens from Gulf of Maine. From Bigelow and Welsh|

General range. — Both sides of the North Atlan-
tic; northward to Spitzbergen, Nova Zembla,
Iceland, West Greenland, Davis Strait, south-
eastern Labrador, coasts and Banks of New-
foundland, and Gulf of St. Lawrence; southward
to the offing of southern New England and as
far as the offing of New Jersey in deep water
along the American coast,27 to the northern part

■ By Veschezerov, in Knipovitch, Tolar Sci. Inst. Sea Fisheries and
Oceanogr., No. 8, 1941, pp. 238-270 (Russian).

* Taning (Journal du Conseil Cons, Internat. Explor. Mer., vol. 16, 1949,
p. 86) is of the opinion that the American rosefish does not belong to the same
species as the European S. marinus hence he refers to it as S.fasciatus, Storer,
1854. But our own comparison of good-sized specimens from the two sides
of the Atlantic has failed to show any differences that seem sufficient for
specific separation, whether in number of scales, in the spines on the cheefes,
in the fins, or in bodily proportions. We are much indebted to Dr. C. E.
Lucas for sending us a series of rosefish of various sizes that had been landed
in Aberdeen, Scotland.

434

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

of the North Sea and to the southwestern coast
of Iceland along the European coast.

Occurrence in the Gulf of Maine. — This is one
of the most plentiful of the commercially impor-
tant fishes in all but the shoalest parts of the
Gulf: on the offshore banks, in or over the deep
central basin, and along shore. To list its known
occurrences would be to mention practically every
station where hook-and-line or otter-trawl fishing
is carried on deeper than 20 fathoms. Thus con-
siderable numbers are sometimes taken on lines
or trawls in 20 to 35 fathoms or more in the
Massachusetts Bay region both winter and sum-
mer, especially on or near rocky bottom, while
many are caught on and near Jeffreys Ledge
and at other spots between Cape Ann and Boon
Island.

The fact that the Grampus took rosefish in 6
out of 7 hauls in 25 to 60 fathoms with a trawl
only 8 feet across the mouth, between Cape Ann
and Penobscot Bay in July 1912, and that Atlantis
took 2,469 rosefish in 12 hauls with a 30-foot
shrimp trawl in 66 to 96 fathoms in the mud-
floored trough west of Jeffreys Ledge in August
1936, show how universal they are in the western
coastal belt of the Gulf at appropriate depths;
and the depth is not too great for them anywhere
in the troughs of our gidf. The number of rose-
fish there seemed to be independent of the numbers
of shrimp (Pandalus) on which we may assume
they were feeding, the average catch per haul being
almost the same (216 fish) for the group of
stations where shrimp were scarce as for the group
where they were plentiful.28

Rosefish are also caught in plenty all along the
northern shores of the Gulf in depths of 25
fathoms or more; they are common in the Bay
of Fundy, even in such enclosed waters as Passa-
maquoddy Bay; Huntsman found them in St.
Mary Bay; and large commercial catches are made
off the western coast of Nova Scotia (5,253,962
pounds reported in 1946).

Turning offshore, the vaguely outlined trough
known as "South Channel" that separates the
Cape Cod-Nantucket Shoals area from Georges
Bank is one of the most productive and hardest
fished of the rosefish grounds (19,016,052 pounds
taken there in 1946); rosefish are generally
distributed on and around Georges Bank itself,

» For further details, see Bigelow and Schroeder, BioL Bull. vol. 76, 1939,
P. 314.

except perhaps on its shoalest parts; also on and
around Browns Bank, and they range down
to a depth of at least 260 fathoms on the southern
slope of Georges Bank; 29 very likely down to
300 fathoms.

The relative yearly catches, from different
areas, show that the inner and central parts
of the Gulf in general are considerably more
productive of rosefish than the offshore banks,
for the poundage reported from off western
Nova Scotia, from the Bay of Fundy, from the
coasts of Maine and Massachusetts, and from the
west-central part of the Gulf (including the Cashes
Bank region and part of the deep basin) which is
referred to as the "inshore grounds" in the
statistical reports, were about three times as great
as from the South Channel, Georges Bank, and
Browns Bank combined in 1940; about 3% times
as great in 1946, and the regional contrast has
been of this same order in other recent years of
record. The South Channel in turn, has been
many times as productive as the much more
extensive area of Georges Bank,30 though there
were enough of them on Georges formerly for 22
successive trawl hauls to have yielded 3,887
rosefish there, September 26 to 30, 1913 (more
than one-third as many as haddock) .

During 1913, rosefish made up 1.8 percent of
the total catch of fish of all kinds made by several
trawlers operating on Georges Bank, June to
December, and 5.9 percent in the South Channel.

This regional contrast between Bank and
Channel emphasizes the very interesting fact
that the rosefish of our Gulf, and those of outer
Nova Scotian waters as well, are decidedly
more plentiful in the deeper basins and depressions,
and on soft bottom, than they are on the grounds
that are the chief centers of abundance for cod
and haddock, and for most of the commercially
important flat fishes.

The statistics do not suggest any very great
difference in the abundance of rosefish as between
Georges Bank and the Nantucket Shoals-Nan-

» 63 large ones taken In one trawl haul, latitude 40°29' N., longitude 67°lf/
W., at 175-195 fathoms, by the Albatross III, May 16, 1950.

» The total reported catch for the period 1937-1946 (no report for 1942)
was a little more than 15RH million pounds for the South Channel, contrasted
with a little less than 2 million pounds (1,876,000) for the whole of Georges
Bank. The catches for individual years ranged between about 9H million
and about 30W million pounds for South Channel; between a little less than
29,000 pounds to a little more than 625,000 pounds for Georges Bank. For a
chart showing the geographical limits of the statistical areas to which the
catches are referred, see Fishery Statistics of the U. S., 1943, Stat. Digest,
No. 18, U. S. Fish and Wildlife Service, 1943, p. 95.

FISHES OF THE GULF OF MAINE

435

tucket Lightship fishing grounds, for while the
catch has averaged only about one-third as great
for the latter as for the former, the statistical
area in question is about one-third as extensive.
But the catches of rosefish (1937-1946), made by
United States vessels from southern Nova Scotia
out across Browns Bank, are of the same general
order of magnitude 31 as for the South Channel.
And a catch of 1,400 rosefish in two sets of a line
trawl on Browns Bank, April 4, 1913, will illus-
trate how plentiful they were there, before they
were so hard-fished as they have been of late.

Large catches of rosefish are also made all along
the outer Nova Scotian shelf to the eastward.
There is an abundant population on the New-
foundland Banks still awaiting exploitation; some
7,000,000 pounds were taken in Hermitage Bay,
on the south coast of Newfoundland from 1947 to
1950.32 And fry have been taken along both
coasts of Newfoundland; also northward from
Flemish Cap, "where the Gulf Stream and the
Labrador current struggle for mastery.33 The
most northerly record for the rosefish on the
American coast is from the outer coast of Labrador
(Camp Islands), a few miles north of the Strait of
Belle Isle.34

It has been known for many years that there are
rosefish in the deep waters of the Gulf of St.
Lawrence. But we still await information as to
how plentiful they may be there.

The upper limit to the vertical range of the rose-
fish in different parts of our Gulf is clearly cor-
related with temperature. Thus it is only deeper
than 15 to 20 fathoms that rosefish are found
during the warm half of the year in the south-
western part of the Gulf. But they have been
known to run up into Gloucester Harbor in num-
bers in winter 3S (never in summer). Many have
been taken near the surface in the spring in the
drift-nets near the Isles of Shoals where it is only
near bottom that they are reported in summer.
We have taken them as shoal as 10 fathoms in
summer off Mount Desert Island, Maine, where
the water warms to about 52°-54° at that depth,
and they occur in Pasamaquoddy Bay in water no

deeper than 5 fathoms at that season, according
to Huntsman. Verrill,36 in fact, described them
as round the wharves at Eastport, no doubt in late
summer or early autumn, the season he studied
the fauna there.

Apart from shifts in depth of the sort just men-
tioned, with the seasonal rise and fall of tempera-
ture, there is no evidence that the adidt rosefish
of our Gulf carry out any regular migration. But
the larvae may journey for long distances while
they drift helpless in the upper layers of the water
(p. 436).

In 1930, we saw gravid females during the last
half of April, with young nearly ready for birth,
evidence that some rosefish may be born in the
Gulf of Maine as early as the first part of May.
Females also, with well-developed eggs, and males
with well-developed milt, are taken commonly by
mid-May, both within the Gulf and on Georges
Bank,37 while we have towed a few newborn fish
(7 to 10 mm.) off Boothbay and off Mount Desert
on May 31 and on June 14. But July 8 is the
earliest that we have taken them in any numbers
in our tow nets (57 larvae off Cape Cod on that
date in 1913.)

Evidently the production of young continues
right through July and August, for the Albatross II
trawled many gravid females, 10 to 13% inches
long, in the central basin of the Gulf in July (1931),
one of them containing about 20,000 young, 6-7
mm. long, practically ready for birth, while we
have towed newly born larvae (6.5-7 mm.) in one
part of the Gulf or another on July 24 and 29 and
August 4, 7, 12, 14, 16, 22, and 31, and as small as
10 mm. on September 2.38 But it is not likely
that many young are produced after the first week
in September.

Records for rosefish larvae and fry for late June,
July, and August along the outer Nova Scotian
shelf, and in the Gulf of St. Lawrence, as well as
from May until into September around the Grand
Banks and up the two coasts of Newfoundland,
show that the season of production commences
nearly as early in the season in these more nor-
therly waters as it does in the Gulf of Maine and

ii Yearly catches, from about 9H million to about 27H million pounds.

" Twentieth Rept. Dept. Fish. Canada (1949-50) 1951, p. 36.

» Taning, Journal du Conseil, Oons. Internat. Explor. Mer., vol. 16, 1949,
p. 90.

" See Frost, Newfoundland Dept. Nat. Resources, Res. Bull. 4, 1938, Cb.
7, for locality records of rosefish fry in Newfoundland and Labrador waters.

» Fish. Ind. U. S. Sect. 1, 1884, p. 262. We have not heard of them In
sny numbers In any other harbor south of Cape Elizabeth.

'• American Naturalist, vol. 5, 1871, p. 400.

" In 1950 Albatross III trawled a number of large males with well-developed
milt, and large females with young nearly or quite ready for birth, on the
southern slope of Georges Bank on May 16, at 175-195 fathoms.

n For completp list, with station localities, numbers and sizes of larvae,
and depths of the hauls, see Bigelow, Bull, Mus. Comp. Zool., vol. 58, 1914,
p. 108; vol. 61, 1917, pp. 271-272.

436

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

that it continues equally late. In north European
waters young rosefish are produced from mid-April
through August, according to locality.

Seemingly the rosefish fry are ready to sink to
near the bottom when they are about 25-30 mm.
long, for we have not taken any larger than 27
mm. in our tow nets, while fry of 1% inches and
upwards are plentiful on bottom, both in the Bay
of Fundy and in deep water off southern New
England. And our failure to take any young
rosefish in our tow nets off Massachusetts Bay in
November or anywhere in the Gulf in winter
is evidence that their descent to the bottom takes
place early in their first autumn.

In north European waters such of the young
rosefish as are fated to take to the bottom at all
are described as continuing pelagic in the upper
layers until they are 2-2 K inches (to 60 mm.) long.

Apparently rosefish never produce their young
in less than 20 to 30 fathoms west or south of
Penobscot Bay; and while they may perhaps do
so in shoaler water about Mount Desert, and
further east along the coast of Maine, Huntsman 39
reports that the spawning individuals move out
into deep water. With this qualification, we have
taken pelagic young in our tow nets at so many
localities in the northern part of the Gulf including
Southwest Harbor on Mount Desert Island, and
so generally distributed, as to show that rosefish
produce their young wherever they may chance
to be, and do not gather on special grounds for the
purpose. Rosefish (unlike most of the fishes
producing buoyant eggs) also breed successfully
in the Bay of Fundy, their larvae having been
found both at the mouth of the bay and for some
distance up the center, during the late summer.40

In the inner parts of the Gulf, our largest
catches of its drifting young have all been located
within a few miles, one side or the other, of the
50-fathom contour line. Examples are catches
of several hundred off Cape Elizabeth on July 29,
1912; near Cape Sable on August 11, 1914; near
Cashes Ledge on August 10, 1913, and on Sep-
tember 1, 1915; in the sink off Gloucester on
August 9, 1913; on Platts Bank on August 7, 1912.
And Goode and Bean 41 report the fry as caught
"by the bushel" in the trawl by the Fish Hawk at
55 fathoms, presumably off Cape Cod, that being

>' Contr. Canadian Biol. (1920-1921) 1922, p. 64.
• Huntsman, Contrib. Canadian Biol. (1920-1921), 1922, p. 64.
1 8mithsonian Contrib. Knowl., vol. 30, 1895, pp. 260, 261.

the only Fish Hawk station where the rosefish
is listed by them. These last catches rival the
swarms of young Sebastes that have been en-
countered between Iceland and the Faroes.42

On the other hand, most of our records for their
pelagic young outside the 100-fathom contour line
have been based on occasional specimens only.
We have seldom taken young Sebastes in the
western basin, though we have towed there fre-
quently at all seasons, and never in the deep south-
eastern trough of the Gulf nor in the eastern chan-
nel between Georges Bank and Browns. All this
suggests that the chief production of rosefish within
the Gulf of Maine occurs at about 50 fathoms.

The presence of gravid females and ripe males
on Georges bank (p. 435), together with the abund-
ance of mature fish in the so-called "South Chan-
nel," shows that this general region is an important
center of production. And the rosefish also breeds
considerably farther west than this on the outer
edge of the continental shelf, for young fry and
adult females full of eggs were collected in 100 to
180 fathoms off the southern coast of New England
during the early years of the United States Fish
Commission.

The shelf along outer Nova Scotia (especially
the depressions between the banks), the basin of
the Gulf of St. Lawrence, and the waters around
Newfoundland, must be productive nurseries, also,
to judge from the abundance of young drifting
stages that have been collected there.43

Importance and abundance. — The only measure
of abundance of rosefish in our Gulf available be-
fore 1935 was the number taken in a few experi-
mental trawl hauls, or on long lines (p. 434), for
there was so little demand for them that nearly
all of those caught incidentally were thrown back
by the fishermen. Thus the reported catch for
our Gulf was only 54,095 pounds in 1919, rising
to a yearly average of about 209,000 pounds for
the period 1931-1933. But the rosefish is a good
table fish, excellent for quick freezing and filleting.
The marketing of it as frozen fillets in 1935 so in-
creased the demand that the landings from the
Gulf of Maine, plus fish taken from southern Nova

« Schmidt, Skrifter, Kommiss, Havundersflgelser, No. 1, 1904, p. 9; Taning,
Journal du Conseil, Cons. Intemat. Explor. Mer., vol. 16, 1949, p. 93-04.

« See Dannevig (Canadian Fish. Exped. (1914-1915) 1919, pp. 12-14, figs.
8-10), for records of young rosefish along outer Nova Scotia and in the Gulf
of St. Lawrence; Frost (Newfoundland Dept. Nat. Resources, Res. Bull. 4,
1936, Ch. 7) for Newfoundland; also Reports, Newfoundland Fisheries Re-
search Commission, vol. 1, No. 4, 1932; vol. 2, No. 1, 1933; vol. 2, No. 2, 1934,
for details as to exact localities and dates.

FISHES OF THE GULF OF MAINE

437

Scotia out to Browns Bank rose to 17 million
pounds in that year, to about 55 million pounds
in 1936, about 66-89 million pounds in 1938 and
in 1939, to about 106 million pounds in 1940, and
to about 136 million pounds in 1941. The land-
ings fell to about 100 million pounds in 1943, but
rose again in 1945 to a peak of 151 million pounds.
This corresponds to about an equal number of
individual fish, a number larger than that for any
other fish commercially important in our Gulf,
except the herring.

It is now generally believed that this yearly
drain was greater than a fish requiring 8 or 9 years
to reach marketable size could withstand ; the catch
(Gulf of Maine and southwestern Nova Scotia)
fell by about 30 percent the next year, and to only
about one-fourth as much in 1949 as had been
landed from these areas in 1945. 44 And this would
have been calamitous for the fishery had the fleet
not been able to draw on the rosefish to the east-
ward, along the Nova Scotian shelf, whence some-
thing like 133 million pounds were landed in New
England ports in 1949, or between three and four
times as much as from the Gulf of Maine.

We refer the reader to the table on page 333 for
the monetary value of the catches of rosefish in
recent years, as compared with cod, haddock, and
mackerel.

George F. Kelly, writing in the Maine Coast
Fisherman,45 has recently emphasized the prob-
ability that the Nova Scotian catch may also be

«• Landings of 108 million pounds for 1946; only about 36 million pounds
for 1919.
'• Vol. 5, No. 7, Jan. 1951, p. 9.

expected to decline from its present high level as
soon as the accumulated stock of old fish is reduced
there, as it has been in the Gulf of Maine. The
fishery would then have had to depend on the
annual increment of growth of a stock that has
stabilized at a level considerably below its virgin
state, unless operations had been extended to New-
foundland waters, where the same chain of events
will eventually follow. And we must expect this
increment to be far smaller for the slow-growing
rosefish than it is for faster growing fishes, such as
the cod or the haddock.

Finally, almost the entire commercial catch is
taken in otter trawls; also while the rosefish is of
such great importance to the commercial fisher-
men, it offers nothing to the angler; most of them
live too deep to be within his reach, and any
hooked would come in with very little resistance.

Black-bellied rosefish Helicolenus dactylopterus
(De la Roche) 1809

Red bream; Blue mouth

Jordan and Evermann, 1896-1900, p. 1837 as (H. dacty-
lopterus (De la Roche) and H. maderensis Goode and Bean
1895)."

Description. — This species resembles the com-
mon rosefish closely in its general form and in the
outline and arrangement of its fins. But the lower
7 to 9 rays of its pectoral fins are free from the fin
membrane along the outer half to one-third of their
length, and the upper margin of the pectorals is

« We have examined some of Goode and Bean's specimens and agree with
Holt and Byme (Fisheries, Ireland, Sei. Inv. (1906), v. 1908) that the so-
called H. maderensis is identical with //. dactylopterus.

Figure 224. — Black-bellied rosefish (Helicolenus dactylopterus), off southern New England. Drawing by Louella E. Cable.

438

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

nearly straight, which gives the fins an aspect
noticeably different from that of the rosefish. The
space between the eyes (flat in the rosefish) is
grooved in the black-bellied species ; there are only
12 spines in the spiny portion of its dorsal fin (14 or
15 in the rosefish) and only 5 or 6 soft anal rays
instead of 7 (in addition to 3 stiff spines).

Its caudal fin is relatively larger than that of the
rosefish; its eyes closer together, the distance be-
tween them being less than one-half as great as
the diameter of the eye (about two-thirds to three-
quarters in the rosefish); the maximum depth of
its body is somewhat less than the distance from
tip of upper jaw to upper corner of gill cover; and
its scales are larger relatively.47 A more important
difference anatomically is that the red bream has
only 24 or 25 vertebrae, the rosefish 31.

Color.- — More or less vivid reddish or pale
pinkish, usually with some brown and green along
the back and with irregular cross bands of darker
or brighter scarlet on some specimens; the upper
part of the sides marked with a sparse pattern of
narrow, dusky vemiculations, roughly following
the edges of the scales; and each gill cover gener-
ally has a leaden or dusky patch caused by the
black inner surface shining through the bone. The
lower surface is without dark markings. All the
fins are pinkish, the spiny part of the dorsal
mottled with white, and the soft portion of the
dorsal, the ventrals, and the anal edged with white.
The lining of the belly cavity is black, hence one
of its common names.

Size. — Maximum length about 15 inches.

Habits. — Catch records show that the black-
bellied rosefish sometimes are in the mid-depths,
sometimes on bottom or close to it. Beyond
this nothing is known of their daily life. Neither

is it known definitely whether their eggs are
hatched within the oviducts of the mother, as in
the rosefish (p. 433), or whether they are set free
in the water, like those of most fishes.48

General range. — Known from the eastern slope
of Georges Bank westward and southward to
Florida in depths of 68 to 373 fathoms in the
western Atlantic; from Norway to the Canaries
in the eastern; also in the Mediterranean.

Occurrence in the Gulf oj Maine. — This fish
must be generally distributed over the outer part
of the continental shelf and along the upper part
of the continental slope as far east as the general
offing of Nantucket, for it has been reported from
27 stations between longitude 72° and a few
miles east of longitude 70°,49 including one catch
of more than 100 of them, 4% to 11 inches long,
in one haul, by the Albatross III.10 One about
13 inches long was trawled on the eastern edge of
Georges Bank, at 175 fathoms, October 6, 1929."
Subsequent records that fall within the limits set
here for the Gulf of Maine, are of 24 fish, 4-10
inches long, trawled at 5 stations south of Nan-
tucket, at 68-240 fathoms, by the Albatross III,
May 11-18, 1950; of one brought in by the trawler
Red Jacket from the northern slope of Georges
Bank, from 120 fathoms, in 1949 ;82 and of a catch
of about 300 pounds of them, made in the south-
eastern part of the basin of the Gulf, at 120-140
fathoms, July 24, 1948.63

This last catch is especially interesting, for
it shows that schools of black-bellied rosefish
may occasionally come in via the deep channel
between Georges and Browns Banks. But they
have never been reported in the inner parts of
the Gulf, nor are they to be expected there unless
as strays from offshore.

BOAR FISHES. FAMILY CAPROIDAE

Boar fish Antigonia capros Lowe 1843
Jordan and Evermann, 1896-1900, p. 1665.
Description. — This Boar Fish64 is set apart
from all other Gulf of Maine fishes by the fact
that its very thin body is deeper than it is long
(longer than deep in all other species yet recorded
from our Gulf). It resembles the John Dory

"i About 42-48 oblique rows of scales from upper comer of Rill opening to
base of caudal fin in Helicolenus, 60-70 in Sebastes.

" Ehrenbaum (Nordisches Plankton, Zool., vol. 1, 1905, p. 51) thought it
probable that this is an egg-laying species, and Taning (Journal du Conseil,
Conseil Internat. Explor. de la Mer, vol. 16, 1949, p. 86) so characteristics it.
But its ripe eggs have not been seen, so far as we know.

<• For list of stations, with depths, up to 1895, see Goode and Bean, Smith-

(p. 297) in the general arrangement of its fins,
both the spiny portion of the dorsal and the soft
portion being well developed, with the latter much
the longer of the two, but lower; the soft-rayed

sonian Contrib. Knowl., vol. 30, 1895, pp. 251-252, as Helicolenm maderensis;
Albatross III, also, trawled one or more specimens at nine stations off
southern New England in May 1950.

» Latitude 39°42' N., longitude 71°57' W., 145-210 fathoms. May 12, 1950.

« This specimen, reported by Firth (Bull. 61, Boston Soc. Nat. Hist., 1931,
p. 13) is in the Museum of Comparative Zoology.

•> Specimen in Museum of Comparative Zoology.

» Taken by the schooner Alice M. Doughty, Capt. Manual Silva. Six of
these specimens are in the Museum of Comparative Zoology.

« This is the only member of the family that has been reported from the
western side of the North Atlantic.

FISHES OF THE GULF OF MAINE

439

anal is about as long as the soft dorsal and is
preceded by 3 spines with fin membrane. The
ventrals are placed a little behind the pectorals.
It lacks the bony skin plates and the filamentous
prolongations of the dorsal spines so conspicuous
on the John Dory; and its mouth is very small
(larger in the John Dory).

Color. — Color, in life, pink and pinkish white.

Size. — Maximum reported length about 1 foot.

General range. — Tropical and subtropical Atlan-
tic, mostly offshore.66

Occurrence in the Gulf oj Maine. — We mention
this fish because we have seen 8 specimens 66 and
heard of 6 others 67 that were trawled in 55-80
fathoms, south of Nantucket Lightship in May
1950. Other records of it near the American
coast are one trawled by the Albatross III at 50
fathoms and a second at 22 fathoms off North
Carolina, in January 1950. It has also been
taken near Madeira, off the Barbados, and in
Cuban waters.

u Reports of it from Japan, from the Kai Islands and from the Celebes Sea
(Manado) may have been based on a closely allied fish. For descriptions of
the species of this genus, with references, see Fraser-Brunner (Ann. Mag.
Nat. Hist., Ser. 12, vol. 3, No. 32, 1950, pp. 721-724).

M Three trawled by Albatross III; five by the Eugene H.

" Reported by Capt. Henry Klimm, of the dragger Eugene H.

Figure 225. — Boarfish (Antigonia capros), 105 mm. speci-
men, south of Nantucket Lightship. Drawing by H.
B. Bigelow.

THE SCULPINS AND SEA RAVENS. FAMILIES COTTIDAE AND HEMITRIPTERIDAE

The several members of the sculpin and sea-
raven tribe that are known from the Gulf of
Maine are a homogeneous group, characterized by
large spiny heads; very wide gill openings; very
broad mouths; slender bodies; separate spiny and
soft-rayed dorsal fins (united in some rare species) ;
large fanlike pectorals but small caudals; and by
ventrals that are reduced to three long rays. All
of them, too, have a fashion of spreading the gill
covers and of flattening the head when taken in
the hand. They likewise produce grunting sounds,
and some of them have the power of inflating
themselves with air or water when they are
molested. The only other Gulf of Maine fishes
that resemble them in general form, are the sea
robins (p. 467), the toadfish (p. 518), and the goose-
fish (p. 537). But the entire head of the sea robin
is armed with bony plates, different from the
soft-skinned head of a sculpin; in the toadfish the
soft portion of the dorsal fin is many times as long
as the spiny part (at most twice as long as the

spiny part in a sculpin) ; and not only are the fins
of the goosefish small and weak as compared with
the present family, but its lower jaw projects far
beyond the upper, and its mouth is full of very
large pointed teeth, whereas in the sculpins the
teeth are small and the upper and lower jaws are
of approximately equal length.

The sculpin tribe, as a group, are egg-laying
fishes.68 Among the Arctic members of the
family, including the genera Artediellus, Cottun-
culus, Gymnocanthus, and Icelus, the males have
a long anal papilla, through which the urinary
duct and the sperm ducts both pass. The sup-
position is that this serves as a copulating organ,
fertilization taking place within the female, and
the fertilized eggs being laid soon after.69

M Eggs with embryos far advanced in development have been reported
within the ovaries of female short horn sculpins (Myoxocephalus scorpius)
from Finland (Nordquist, Svensk. Fiskeri Tidskr., year 6, 1899). But it is
well established that this sculpin ordinarily lays eggs, as described below
(p. 447).

* See Jensen and Vols0e (Danske Vidensk. Selskab. Biol. Meddel., vol.
21, No. 6, 1949, p. 18) for a detailed account of the anal papula in Icelus.

440

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

KEY TO GULF OF MAINE SCULPINS AND SEA RAVENS

1. There is only one dorsal fin, the spiny and soft parts being continuous, one with the other Arctic sculpin, p. 453

There are two separate dorsal fins 2

2. The first dorsal fin is deeply notched between the spines; the lower jaw and the top of the head are adorned with fleshy

tags Sea raven, p. 454

The first dorsal is not deeply notched between the spines; there are no fleshy tags about the head 3

3. The long spine on each cheek is branched at the tip Staghorn sculpin, p. 452

The long spine on each cheek is simple, not branched at the tip 4

4. The anal fin is long (25 rays) ; there is a series of bony plates along each side of the body Mailed sculpin, p. 441

The anal fin is short (14 rays or fewer) : there are no bony plates along the sides of the body 5

5. The long spine on the cheek is hooked upward Hook-eared sculpin, p, 440

The long spine on the cheek is straight, not hooked 6

The longest (uppermost) cheek spine is four times as long as the one below it, and reaches back to the margin of the

gill cover; all the head spines are very sharp Longhorn sculpin, p, 449

The uppermost cheek spine is not more than twice as long as the one below it, and does not reach more than about

half way to the margin of the gill cover; the head spines are blunter 7

Total length more than 9 inches Shorthorn sculpin, p. 445

Total length less than 8 inches 8

Anal fin with 13 or 14 rays; the soft skin of each side of the throat is pierced by a minute pore close behind the lower

part of the last gill arch Shorthorn sculpin, young specimens, p. 446

Anal fin with only 10 or 11 rays; sides of throat behind last gill arch have no pore Grubby, p. 443

6.

7.

Hook-eared sculpin Artediellus wncinatus (Kein-
hardt) 1833

Arctic Sculpin

Jordan and Evermann, 1896-1900, p. 1906, as Artediellus
allanlicus Jordan and Evermann.

Description. — The most distinctive feature of
this species among local sculpins is the long
hooklike spine on each cheek, pointing backward
and upward, plainly shown in the illustration
(fig. 226). There is also a short backward-pointing
spine covered by a flap of skin at the upper corner
of each gill cover, two short spines on the top of
the nose between the two pairs of nostrils, and a
pair of blunt knobs above the eyes. Head,

mouth, and tapering body are of the usual sculpin
form. The skin is smooth and naked. The
spiny dorsal fin is short (7 to 9 spines) and
rounded in outline, the soft dorsal fin is about
twice as long (13 rays), and the anal (11-12 rays)
is a little shorter than the soft dorsal, which it
resembles in outline and under which it stands.
Each ventral fin consists of three long rays that
reach back nearly to the vent; the pectorals, wide
at the base and rounded in outline, reach beyond
the beginning of the soft dorsal when they are
laid back, and the caudal fin is narrower than it is
in the commoner Gulf of Maine sculpins. The
jaws and the roof of the mouth are armed with
several series of small bristle-like teeth.

Figure 226. — Hook-eared sculpin (Artediellus uncinalus), Massachusetts Bay.

by H. L. Todd.

From Jordan and Evermann. Drawing

FISHES OF THE GULF OF MAINE

441

Color.- — Preserved specimens are mottled with
dark and pale brown, sometimes with a reddish
tinge, and most of them have a blotch at the
base of the caudal fin. All the fins are grayish or
blackish, with oblique or vertical pale cross bands.

Size. — This is one of the smallest of sculpins,
growing to a length of only about 4 inches.

General range. — This is a cold-water fish known
from Labrador and the west coast of Greenland to
Cape Cod in the western Atlantic; also in the
littoral waters of arctic Europe, of Siberia, and
of Greenland.60

Occurrence in the Gulf of Maine. — This sculpin,
formerly thought to be rare in the Gulf of Maine,
is now known to be generally distributed there in
depths greater than 20 to 30 fathoms. It was
dredged in numbers in the deeper parts of Massa-
chusetts Bay many years ago. And we have
since taken it repeatedly near Mount Desert; off
Cape Elizabeth; in the trough between Jeffreys
Ledge and the coast; around Cashes Ledge; along
the northern slopes of Georges Bank; in the south-
eastern part of the basin of the Gulf; and at the
entrance to the deep gully between Georges and
Browns Banks, in depths ranging from 20 to 150
fathoms. Individual trawl hauls have yielded up
to 6 or 8 specimens, both on hard bottom and
on soft.

To the eastward and northward it has been
taken off Cape Sable; at a number of places off
the outer coast of Nova Scotia; and on the New-
foundland Banks, at depths of 50 to 190 fathoms.61

* After examining specimens from New England waters and comparing
them with published drawings of European fish, we can find no significant
differences between the hook-eared sculpins of the two sides of the Atlantic.

ei For localities of record off outer Nova Scotia and on the Newfoundland
Banks, see Qoode and Bean, Smithson. Contrib. Knowl., vol. 30, 1S95, p.
268; also Reports, Newfoundland Fishery Research Commission, Vol. 1,
No. 4, 1932, p. 108; vol. 2, No. 1, 1933, p. 125.

It is common enough in the Gulf of St. Lawrence
for Huntsman to have classed it as a characteristic
inhabitant of the icy intermediate water layer on
the Banks,62 while Vladykov and Tremblay 63
have reported it from the estuary of the St.
Lawrence River near Trois Pistoles; it has been
reported from Hamilton Inlet on the outer coast
of Labrador; 64 and doubtless it will be found
farther north, when the fish fauna of the outer
Labrador coast has been explored more thoroughly,
for it is known from West Greenland.

Presumably, it is resident in small numbers
wherever found, sculpins not being migratory, but
nothing whatever is known of its way of life.

Mailed sculpin Triglops ommatistius Gilbert
1913

Jordan and Evermann, 1896-1900, p. 1923, Triglops
pingeli (Reinhardt), 1832, in part.

Gilbert, Proceedings, U. S. National Museum, vol. 44,
1913, p. 465.

Description. — The most distinctive feature of
this sculpin, apart from its very long anal fin, is
that it has a row of about 45 broad plate-like scales
along its lateral line on each side, with smaller
spiny scales below the dorsal fins, while the skin of
the sides lower down is gathered in obliquely trans-
verse folds. The body, too, is more tapering than
that of our other sculpins, the caudal peduncle
more slender, and the tail fin is smaller. Further-
more, the head is smaller and smoother than in any
of the sculpins that are common in the Gulf of
Maine, with short spines and many prickles. The
first dorsal fin (10 to 12 spines) originates over the

« Trans. Roy. Soc. Canada, Ser. 3, vol. 12, Sect. 4, 1918, p. 63, as Centrtr
dermkhthys uncinatus.
° Natural. Canad., vol. 62, 1935, p. 79.
t Kendall, Proc. Portland Soc. Nat. Hist., vol. 2, 1909, p. 217.

Figure 227. — Mailed sculpin (Triglops ommatistius), off Chebucto, Nova Scotia. From Jordan and Evermann. Drawing

by H. L. Todd.
210941—53 29

442

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

bases of the pectorals and is higher than the second
dorsal, but only about half as long. The second
dorsal has 20 to 25 rays. The anal fin is similar to
the second dorsal in form and stands below it, but
is a little shorter (20 to 22 rays). The ventral fins
(each with 3 rays as is the rule among sculpins)
reach about as far back as the rear end of the first
dorsal, while the pectorals (17 rays) are of the fan-
like shape usual among sculpins. The males have
a very large and noticeable anal papilla.

Color.- — Olive above; white, yellowish or orange
below. There are four dusky blotches above the
lateral line on each side, one on the caudal pe-
duncle, one passing through the first dorsal fin,
and two passing through the second dorsal fin.
The fins are variously marked with yellowish and
with gray-black. The first dorsal of the male has a
dusky blotch between the first and second spines
and another between the seventh and tenth spines;
the second dorsal is marked with three horizontal
olivaceous bars. Females lack the blotches on the
first dorsal fin; and their second dorsal is marked
with narrow lines of dots.

Size. — This is a small species, probably growing
to about 8 inches, the maximum that is recorded
for its European representative.65 The largest
yet recorded for the Gulf of Maine was 6 inches
long.

Habits. — Little is known of its habits beyond
the bare fact that it is a bottom fish, like other
sculpins. Any that breed in the Gulf of Maine
probably spawn in midsummer, Cox 66 having re-
ported a ripe female at Cape Breton in July. Its
eggs were pinkish, 2 mm. in diameter, with many oil
globules. Presumably the eggs sink like those of
other sculpins. The European mailed sculpin is
known to eat worms and various crustaceans, and
the diet of the American form is the same,
probably.

General range. — Sculpins of this general type are
circumpolar, ranging south to Cape Cod along
the American coast and to the Baltic on the
European side of the Atlantic, in rather deep water.
But they show a tendency to split up into local
races, the constancy of which is yet to be tested by
a study of large series. Newfoundland specimens,
for example, differ so much from typical Triglops
ommatistius in the arrangement and number of

folds of skin along the sides that Gilbert w has
dignified them with a separate name (as the
subspecies terranovae of species ommatistius); and
both the eastern American forms are distinguished
from the east Greenland and European mailed
sculpins by the presence of the eyespot on the first
dorsal fin of the male (which the European form
lacks) and by slightly fewer fin rays. We do not
feel convinced, however, that all these forms,
together with the Bering Sea form (Triglops beanii
Gilbert, 1895), will not finally prove to be local
varieties of a single wide-ranging species.

Occurrence in the Gulf oj Maine. — Judging from
the scarcity of records this cold water fish is
uncommon in the Gulf of Maine. Specimens have
been recorded from the neighborhood of St.
Andrews in the Bay of Fundy, in 15 fathoms
(reported by Huntsman); a few from Massa-
chusetts Bay and from off Race Point, Cape Cod
(now or formerly in the collection of the Boston
Society of Natural History); 11 others now in the
United States National Museum were from
Gloucester, Cape Cod, and Georges Bank; we
have trawled them near Mount Desert; in Massa-
chusetts Bay; off Cape Ann; off Cape Cod; and
around the northern slope of Georges Bank,
in depths of 20 to 140 fathoms in various months
from spring to autumn; and two were trawled
on the southeast slope of Georges68 by the
Albatross III, July 17, 1948, in 45 fathoms. Our
most southerly record for it was about 10 miles
east of Chatham, Mass.

The fact that Gilbert found differences be-
tween the Gulf of Maine and Newfoundland
specimens, with others from Chebucto Head
(Nova Scotia) and from Georges Bank inter-
mediate between them, suggests that the mailed
sculpin is a permanent resident of the inner parts
of the Gulf, rather than that it appears there
only as a wanderer, past Cape Sable, from the
east and north.

Eastward and northward from our Gulf, this
sculpin is described as being rather common to
numerous on the outer Nova Scotian fishing
grounds, and as one of the characteristic mem-
bers of the fish fauna of the icy cold water on the
Banks and in the Gulf of St. Lawrence.69

» CoUett, Norske Nordhaus-Expedition, 1876-78, Zool., Fiske, 1880, p. 38.
« Contrib. Canad. Biol. (1818-1920) 1921, p. 111.

•» Proc. U. S. Nat. Mus., vol. 44, 1913, p. 467.

«' Latitute 40°48' N., longitute 66°31' W. (Arnold, Copeia 1949, p. 299).

•• See Huntsman, Trans. Royal Soe. Canada, Ser. 3, vol. 12, Sect. 4, 1918,
pp. 61-67, for a very interesting account of the fishes that are characteristic
of the different water layers in the G ulf of St. Lawrence.

FISHES OF THE GULF OF MAINE

443

It is also reported off Bonne Bay on the west
coast of Newfoundland, in the Strait of Belle
Isle, and from the south coast of Newfoundland.
It is so widespread on the eastern part of the Grand
Banks that it was taken at 18 stations there on
the cruises of the Newfoundland Fisheries Re-
search Commission. It is also reported off the
east coast of the Avalon Peninsula, and off Sand-
wich Bay on the outer coast of Labrador.

Grubby Myoxocephalus aeneus (Mitchill) 1815 70

Little sculpin

Jordan and Evermann, 1896-1900, p. 1972.

Description. — The most distinctive features of
the grubby, as compared with others of its tribe,
are its short, simple head spines, combined with
small size at maturity. It is of the typical
sculpin form, though proportionately a stouter
fish than either the shorthorned or the longhorned
species, that is, about one-fourth as deep as it is
long with smooth skin but showing the head
ridges and spines typical of its genus. Most
noticeable of these are a ridge with two spines
running along the top of the head over each eye;
a pair of spines between the nostrils; and six
short spines on each side of the face between
snout and gill opening. None of the cheek spines
are long (p. 449). The spiny dorsal fin (9 spines),
originating a little in front of the upper corner of
the gill opening, is shorter (front to rear) than the

" Placed in the genus Aeanthocotlm Qirard, 1849, by Jordan, Evermann,
and Clark, Rept. U. S. Comm. Fish (1928), Pt. 2, 1930, p. 386.

second dorsal of 13 or 14 soft rays; its longest
spines, measured from base to tip, are about the
same length as the longest soft rays of the second
dorsal; and the two fins are so close together
that there is no free space between them. The
anal fin (10 or 11 rays) is a little shorter than
the second dorsal, under which it stands. The
pectorals are of the fanlike outline characteristic
of this family, while each ventral fin consists of
one spine and three rays. There is no slit or
pore behind the last gill arch (there is such a slit
or pore in the shorthorn sculpin, at least in most
specimens, p. 445).

Color. — Grubbies, like other sculpins, vary in
color according to the bottoms on which they lie.
All that we have seen, however (this confirms the
published descriptions), have been fight to dark
gray or greenish-gray above, with darker shadings
or irregular barrings that are most evident on
the sides and on the fins. The sides of the head
are usually mottled fight and dark; the belly is
pale gray or white. According to information
supplied by Dr. A. G. Huntsman, the presence
of an uninterrupted pale band of considerable
length along the lower sides of the caudal peduncle
is a useful field character. But we have seen
some specimens intermediate in this respect be-
tween the extreme condition shown in figure 228
and the variable mottlings and cloudings of the
shorthorn sculpin.

Size.- — -This is the smallest of our common
sculpins, few growing to more than 5 or 6 inches
in length, and perhaps none to more than 8 inches.

Figure 228. — Little sculpin (Myoxocephalus aeneus), Staten Island, New York.

by H. L. Todd.

From Jordan and Evermann. Drawing

444

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Remarks. — The differences between the grubby
and the shorthorn sculpin in number of anal rays,
and in the presence or absence of a pore behind
the last gill arch, seem sharp enough to forbid
the possibility that the former may be a dwarf
race of the latter. Determination, however,
of these characters required such close examina-
tion, and grubbies resemble young shorthorns so
closely in all other respects that it is not easy to tell
the one from the other. We therefore suggest that
any small sculpin that may prove difficult to name
be forwarded for identification either to the labora-
tory of the U. S. Fish and Wildlife Service, Woods
Hole, Mass.; to the Division of Fishes, U. S.
National Museum, Washington, D. C; or to the
Department of Fishes, Museum of Comparative
Zoology, Cambridge, Mass.

Habits. — On the southern shores of New Eng-
land, where the grubby is not only more nearly uni-
versal than it is to the east and north of Cape Cod,
but more plentiful, it is found from tide mark down
to 15 fathoms or so. But we have taken it as deep
as 28 fathoms in the Gulf of Maine, and Cox has re-
ported it in the Gulf of St. Lawrence from the stom-
achs of cod caught in 60 to 70 fathoms. It is found
on all sorts of bottoms, most abundantly among eel
grass (Zostera) during the years when this plant
was more plentiful than at present. And it is
the only sculpin that summers in very shoal water
along southern New England (Woods Hole and
up Narragansett Bay) and near New York
Harbor.

In the Gulf of St. Lawrence (e.g., around theMag-
dalen Islands) and on the Nova Scotian coast in
general, it is found in estuaries, as in the southern
part of its range, as well as outside. But it seems
more restricted to the open coast in the Gulf of
Maine, for Huntsman found it rare as far up Pas-
samaquoddy Bay as St. Andrews, though common
at the mouth, and more plentiful in St. Mary Bay
and in Annapolis Basin than it is in Minas Basin on
the Scotian side of the Bay of Fundy. Neither
have we seen it in salt creeks about Massachusetts
Bay.

The known distribution of the grubby in summer
proves that it is certainly at home in water as warm
as 69° F., and perhaps a degree or two warmer;
these temperatures are several degrees higher than
are preferred by its larger relative, the shorthorn.
On the other hand, it survives temperatures as low
as 32° in winter, if not lower, both in the Gulf of St.

Lawrence, on the Nova Scotian coast, and about
Woods Hole. Its presence in the inner parts of
Narragansett Bay on the one hand and off open
coasts on the other also proves it resistant to a wide
range of salinity, but it never runs up into appreci-
ably brackish water, as far as we can learn.

Probably the grubby breeds throughout its geo-
graphic range, certainly as far north as the south-
ern part of the Gulf of St. Lawrence. And the Bay
of Fundy appears to be the site of successful repro-
duction, for Huntsman found grubbies of all sizes
there. The spawning season lasts all winter off
southern New England and until June in the Gulf
of St. Lawrence, Cox having reported a ripe female
on the 18th of that month, at Amherst Island
(Magdalen group). The eggs, which are described
as of a beautiful green color and 1 mm. in diameter,
sink like those of other sculpins and stick to sea-
weeds or to any other objects they chance to rest
upon. Young sculpins (this species among them)
have been caught in tow nets at Woods Hole from
January to May.

This sculpin is omnivorous like its relatives,
feeding on all sorts of small animals which it finds
on the bottom, such as annelid worms, shrimps,
crabs, copepods, snails, nudibranch mollusks,
ascidians, and on small fish, including alewives,
cunners, eels, mummichogs, launce, silversides,
sticklebacks, and tomcod.71 It also scavenges
any kind of animal refuse.

General range. — North American coastal waters,
from New Jersey to northern Nova Scotia and
to the Guff of St. Lawrence, both in the southern
side, where it is common, and the Strait of
Belle Isle.72

Occurrence in the Gulf of Maine. — It is probable
that this little sculpin is to be found in suitable
localities all around the shores of the Gulf of
Maine, for it is reported as common along both
shores of the Bay of Fundy (including St. Mary
Bay) and at various localities in the Massachu-
setts Bay region, such as Cape Ann, Gloucester,
Salem, Cohasset, and Provincetown, and the

" This list of fish fry eaten is from Vinal Edward's notes at Woods Hole.

T2 Maine has sometimes been given as its northern limit. But Doctor
Huntsman writes us that in 1915 he obtained it in tide pools at Souris, Prince
Edward Island; Needier (Proc. Nova Scotian Inst. Sci., vol. 20, 1940, p. 40)
describes it as the common sculpin in Malpeque Bay, Prince Edward Island;
Cox (Contrib. Canadian Biol. (1918-1920) 1921, p. Ill) characterizes it simi-
larly around the Magdalen Islands; Vladkykov and Tremblay (Nat. Canad.,
vol. 62, 1935, p. 80) report it from the estuary of the St. Lawrence, near Trois
Pistoles; and Jeflers (Contrib. Canadian Biol, and Fish., N. Ser., vol. 7, No.
16, Ser. A; No. 13) (1932, p. 208) found two specimens on the beach at
Raleigh, on the Newfoundland side of the Strait of Belle Isle.

FISHES OF THE GULF OF MAINE

445

Albatross II trawled 9 specimens 43-51 mm. long
in 28 fathoms off the outer coast of Cape Cod
abreast of Chatham, May 1, 1930. But it seems
to be decidedly local in its distribution, for the
only places where it has been definitely reported
along the coast between Cape Ann and the Bay of
Fundy is Casco Bay, nor have we caught it in any
of the harbors of Maine where we have fished. In
any case, it is far outnumbered in the Gulf of
Maine by the two larger sculpins to be mentioned
next.

Importance. — Because it is so small the grubby
is of no commercial value. But wherever it is
common it is something of a nuisance to anglers
fishing for flounders and cunners, for it bites as
greedily at any bait as do its larger relatives, and
it serves as a source of food, no doubt, for more
important fishes.

Shorthorn sculpin Myoxocephalus scorpius
(Linnaeus) 1758 "

Daddy sculpin; Black sculpin; Greenland

SCULPIN

Jordan and Evermann,. 1896-1900, as M. scorpius and
M. groenlandicus, p. 1974.

Description. — The shorthorn sculpin, with its
large flat head, vast mouth, weak tapering body,
bat-like pectorals, and insatiable appetite, typifies

n Placed in the genus Acanthocottus, Girard, 1849, by Jordan, Evermann,
and Clark, Rept. U. S. Comm. Fish. (1928), Pt. 2, 1930, p. 386.

the sculpin race in northern seas. It has a longi-
tudinal ridge with 3 knobs or spines running along
each side of its crown; also about 6 (sometimes
5 or 7) short triangular bluntish spines on each
side of the cheek between snout and gill opening,
the uppermost of these less than twice as long as
the one below it, and reaching not much more
than halfway to the edge of the gill cover. And
there is a short but sharp spine at the upper
corner of each gill cover, pointing rearward and
lying on a flap of skin, besides two thornlike spines
on each shoulder close behind the upper corner of
the gill cover.

There is a pore, or small slit, piercing the soft
skin low down on each side of the throat close
behind the last gill arch, easily seen on large speci-
mens and detectable even on small ones on close
examination.

The very large eyes are at least as wide as the
space between them, set high up on the sides of
the head with the upper edges close to the dorsal
profile, and they are directed a little upward as
well as outward. The two parts of the dorsal fin
are entirely separated by a deep notch, but there
is no gap between them. The forward part has
9 to 11 spines, the rear part about 16 or 17
(sometimes 15) soft rays, the longest of which
are only a very little longer, if any, than the long-
est of the spines, each measured from base to
tip. The anal fin, with 13 to 14 rays, is similar
to the second dorsal in shape, but a little smaller;
it originates about under the fourth or fifth soft

Figure 229. — Shorthorn sculpin (Myoxocephalus scorpius), Eastport, Maine. From Goode. Drawing by H. L. Todd

446

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

dorsal ray. The caudal fin is small, its rear
margin weakly rounded; the fanlike pectorals,
of 17 or 18 rays, reach back about as far as the
vent. On large specimens the dorsal, anal, and
pectoral fins are rather noticeably thick and
fleshy. There are two, series of prickly plate-
like scales along each side of the body, one above
the lateral line, the other below it.

Males and females differ in appearance, the
former being the more slender, with higher fins,
and the more brightly colored. Each of the scales,
too, along the lateral line bears three or more
prickles in males, but only one or two at the most
in females, while some of the latter have no scales.
Furthermore, the inner edges of the rays of the
pectoral and ventral fins are armed with teeth
or prickles on the males but not on the females.

Color. — The basic hue of the upper parts is
usually of some shade of brown, ranging from a
warm reddish tint to almost black, with the top
and sides of the head marked with pale blotches
and the back and sides of the body with broad
dark bars on individuals on which the ground
tint is pale. The lower parts of the sides are
more or less spotted with yellow. The belly is
whitish or yellowish in females, usually reddish
orange with large round white spots in males,
this being a good field mark for distinguishing
the sexes. The dorsal fins are mottled dark and
pale, the second dorsal often marked with 3 or 4
definite crossbars, and the caudal fin with various
dark mottlings. The rays of the pectoral and
anal fins are yellow with 2 or 3 irregular dark
crossbars on many specimens, but they are uni-
formly dark in some. Males are more brightly
colored than females in the breeding season, when
their red and yellow tints become very brilliant,
and when an intensification of the red or coppery
ground color of the belly brings out the white
spots more clearly than at other seasons.

Variability. — European studies have shown this
to be a highly variable species, tending to break
up into local races. Whether this is equally
true of it on our side of the Atlantic is not known.74

T* Most American ichthyologists recognize two subspecies of this fish
the true "shorthorn" (scorpiug) and the "Greenland sculpin" (groenlandiais)
And with the prevailing tendency to call American and European fish by
different names it is as the latter that our local sculpin has usually been
recorded. But the differences between the two (size, relative breadth of
the top of the head, and length of the dorsal fin spines) are so very slight
and all of them have proved so variable, that we follow Huntsman (Contrib.
Canadian Biol. (1921) 1922, p. 64) in uniting the two; the more willingly
since both forms have been found on both sides of the Atlantic.

Size. — This is the largest Gulf of Maine sculpin.
It has been said to grow to a length of about 3
feet, but the average run of the adults taken in
our Gulf is only about 8 to 14 inches, the longest
not more than 2 feet. This species increases in
size from south to north, Greenland fish averaging
much larger than those taken off New England
or off the Maritime Provinces.

Remarks. — Young shorthorns, up to 6 or 7
inches long, resemble the little grubby (p. 443) so
closely that they are likely to be confused with it.
Points of difference are that the shorthorn has at
least 13 or 14 rays in its anal fin, and has a pore
piercing each side of the throat close behind the
last gill arch, the grubby only 10 or 11 anal rays,
and no such pores.

Larger specimens of the shorthorn could hardly
be mistaken for any other Gulf of Maine fish,
unless perhaps for the longhorn sculpin. And
even a cursory look is enough to separate one of
these from the other, the upper cheek spine being
less than twice as long as the one below it, and not
reaching more than halfway to the edge of the gill
cover in the shorthorn, but about four times as
long as the one below it in the longhorn, and
reaching back at least as far as the edge of the gill
cover.

Habits. — Bays and the vicinity of ledges that
rise from comparatively smooth bottom in shoal
water are the chief haunts of the shorthorn
sculpin. And it is found indifferently there, on
mud, sand, or pebbles, on bare bottom or among
weeds. Many are also caught off piers and along
our rocky shores by cunner fishermen. Off our
coasts, the great majority live shoaler than 10
fathoms. And while a day's fishing on any of the
ledges northward and eastward from Cape Cod is
likely to yield an occasional shorthorn among
other fish, few are caught on long lines set deeper
than 15-20 fathoms. The deepest records for it
in American waters of which we know are 50
fathoms near Campobello Island, at the mouth of
the Bay of Fundy (reported by Huntsman) , and 57
fathoms in the northern part of the Gulf of St.
Lawrence, just within the Strait of Belle Isle.76

This is a cold-water fish. Even in summer it is
the most plentiful at localities and at depths where
the temperature is lower than 55°-60° F. In
winter it endures temperatures close to the freez-

'« Rept. Newfoundland Fish. Res. Comra, vol. 1, No. 4, 1932, p. 108, sta. 45.

FISHES OF THE GULF OF MAINE

447

ing point of salt water. We have never heard of
one taken in brackish water, at least on our side of
the Atlantic. It is a sluggish fish, often to be seen
lying motionless, and as a rule, it hugs the bottom
so closely that it is hard to tempt one to rise as
much as a few feet by dangling a bait over it.
Neither does it come to the surface voluntarily,
though the surface may drop to the sculpin on the
ebbing tide.

Sculpins usually swim slowly with undulating
motion, spreading their great pectoral fins like
bat's wings. As a rule, they move only a little
way when disturbed, but on occasion they can
dart ahead with folded "wings."

They are among the most voracious fishes,
feeding chiefly on crustaceans, particularly on
crabs, of which they are often full, on shrimps, sea
urchins, and worms; on the fry of various other
fishes; rarely on shellfish. And they are eager
scavengers of any kind of refuse, congregating
about fish wharves and lobster cars to feast on the
debris. Like all sculpins they bite on any bait,
and so greedily that we have caught one time and
again, thrown it back, and seen it bite again
almost as soon as fresh bait reached bottom.

The shorthorn has been described as hiding in
dark crevices or among weeds by day, to emerge at
night. This, however, has not been our ex-
perience, nor did Gill 7e find it doing so at Grand
Manan.

This fish, like the longhorn sculpin, grunts or
gurgles when drawn out of the water, particularly
when handled, and it is also known to grunt in the
water.

We must turn to European sources for the
breeding habits of this sculpin, little attention
having been paid to this phase of its life by Ameri-
can ichthyologists.77 The spawning season is
from November to February, both about Woods
Hole and in north European waters, with the chief
egg production in December, which no doubt
applies equally to the Gulf of Maine. At this
season the adult sculpins have been described as
gathering in schools on sandy or weedy bottom,
with the females greatly outnumbering the males.

Discussion has centered about the manner in
which the eggs are fertilized, it being generally
agreed that this takes place externally as a rule,

but that they may be fertilized within the body of
the mother in some parts of the Baltic Sea. In
either case, the eggs sink 7S and stick together in
irregular spongy masses through which the water
circulates, and which retain considerable moisture
even if they are left bare by the ebbing tide, as
often happens. These egg masses are deposited
on sandy bottoms, in pools in the rocks, among
seaweeds, or in any crevice or hollow, in a tin can,
for instance, or in an old shoe. Sometimes the
male makes a nest of seaweed and pebbles, while
he has been described as sometimes clasping the
egg mass with his pectoral and ventral fins, and
he has been photographed so employed.79

The eggs are of varying shades of red or yellow,
1.5 to 2 mm. in diameter. Incubation is so slow
(occupying 4 to 12 weeks, according to tempera-
ture) that egg masses with advanced embryos
have often been found as late in the spring as
April or even May. Newly hatched larvae are
about 7 to 8 mm. long. In a month they are 10
mm. long and the yolk sac has been absorbed.
The young larvae soon rise to the surface, where
quantities of them have been taken in tow nets in
British waters in March, April, and May. By
May and June some have grown to a length of
22 to 25 mm. They abandon their drifting life
at about this size, or soon after, for the bottom,
and they may be 38 mm. long by July, showing
all the distinctive characters of the adult.80 This
timetable, compiled from European sources, prob-
ably applies equally to the Gulf of Maine, for
larvae are found as early as February in the Bay
of Fundy and thereafter throughout the spring.81

The subsequent rate of growth is not definitely
known. But it is probable that this sculpin is
2 or 3 inches long by the end of its first summer,
for we have taken a few 2-inch fish in late June on
Nantucket Shoals, and 2- to 3%-inch fish in late
September off Boothbay Harbor, Maine. Most
of them, it seems, do not mature sexually until
they are at least 6 inches long.

General range. — -One or another race of this wide
ranging fish is known from Great Britain north-
ward along the coasts of Europe; in Arctic seas

'• Smithsonian Misc. Coll., vol. 47, 1905, p. 352.

« QUI (Smithsonian Misc. Coll., vol. 47, 1905, p. 352) givos a summary of
Its life history.

78 Buoyant eggs taken in the tow net (Agassiz, Proc. Amer. Acad. Arts
Pci., vol. 17, 1882, pi. 3) belonged to some other fish.

» Ehrenbaum, Wiss. Meeresuntersuchungen, Helgoland, Neue Folge, vol.
6, 1904, pi. 8.

» Mcintosh and Masterman, Life-Histories of British Marina Food-Fishes,
1897, p. 129.

•• Huntsman, Contrib. Canadian Biol. (1921) 1922, p 64.

448

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

generally, including Spitzbergen, Nova Zembla,
north Siberia, West Greenland, and northern
Labrador; and southward along the American
coast to southern New England; to New Jersey
as a stray.

Figure 230. — Egg (European). After Ehrenbaum.

Figure 231. — Larva (European), 8.2 mm.
Ehrenbaum.

After

Figure 232. — Larva (European), 10 mm. After
Ehrenbaum.

Figure 233. — Fry, 18 mm. After Ehrenbaum.
Shorthorn sculpin (Myoxocephalus scorpius)

Occurrence in the Gulf of Maine. — This is one
of the most familiar of our shore fishes, common all
around the entire coast line of the Gulf of Maine;
it is not so abundant as the longhorn sculpin
(p. 449), but we doubt if there is a suitable situa-
tion, Cape Cod to Cape Sable, where some short-
horns are not to be found, except perhaps at the
head of the Bay of Fundy.82 But they do not

u Huntsman found none there.

run far up estuaries as a rule, and never into brack-
ish water.

The shorthorn has not been reported positively
either from Georges Bank or from Browns.
Sculpins of some sort, it is true, are so common on
the former that otter trawlers often catch from
20 to 100 per haul, and equally so on Browns
Bank. But fishermen lump this and the next
species together. Also the fact that the few
positively identified on the banks have all proved
to be longhorns, and the general predilection of the
shorthorn for water shoaler than these offshore
grounds, makes it doubtful whether it is to be
found there in any numbers. Further evidence
in this direction (if indirect) is that most of the
shorthorns that were taken during the experi-
mental trawlings of the Newfoundland Fisheries
Research Commission were from hauls shoaler
than 50 fathoms, only one from as deep as 57
fathoms.

Although it is more strictly confined to com-
paratively shoal water than the longhorn sculpin,
the shorthorns are not often seen close to tide
mark in summer, except in the Bay of Fundy
where the temperature of even the uppermost
few feet does not rise above 52°-54° in most
summers, and where there are shorthorns of all
sizes in very shallow water, in summer as well as
at other times of the year.

On the other hand, the shorthorns living around
the open shores of the Gulf show no tendency to
avoid winter chilling by descending to greater
depths in winter, In the Bay of Fundy, for
example, where it is very common, Huntsman has
characterized it as the only fish that remains near
shore during the coldest part of the year; and it
has been described as more plentiful along the
shores of Massachusetts Bay in winter than in
summer, as it certainly is south of Cape Cod.
Certain shallow bays, it is true, such as Duxbury
Harbor, where broad expanses of flats are exposed
at low tide to heating by the sun in summer and
to the formation of ice in winter, are an exception
to this rule; i. e., the shorthorn sculpins tend to
keep to the deeper channels through the coldest
part of the winter as well as during the heat of
midsummer. But we have found no evidence that
they carry out any seasonal migrations more ex-
tensive than this. They are, indeed, among the
most stationary of Gulf of Maine fishes.

FISHES OF THE GULF OF MAINE

449

In Scandinavian waters this fish is said to vary
widely in abundance from year to year, years of
plenty alternating with longer periods of scarcity,
but this does not seem to be the case to any
noticeable extent in the Gulf of Maine where it is
always common.

To the northward and eastward, the shorthorn
is common all along the outer coast of Nova
Scotia, in 10-30 fathoms, and it has been taken
on Banquereau Bank. It has not been reported
in the Magdalen and Prince Edward Island shal-
lows in the southern side of the Gulf of St. Law-
rence, where summer temperatures are high.
But it is to be expected in that side of the Gulf in
slightly deeper water, for it is known all along
the northern shore of the Gulf, from Anticosti to
the Strait of Belle Isle. We find no record of it on
the Newfoundland Banks, probably because of
the depth of water; neither is any definite informa-
tion available as to its status along the south
coast of Newfoundland. But it is recorded off
the east coast, from the trawlings of the New-
foundland Fisheries Research Commission, and
along the outer coast of Labrador, at Battle
Harbor (just north of the Strait of Belle Isle);83
at Rigolet in Hamilton Inlet;84 in the vicinity of
Nain;85 at Fort Chimo, Ungava Bay, and it is
widespread in the Hudson Bay region.86

83 Specimen in Museum of Comparative Zoology.

- See Kendall (Proc. Portland Soc. Nat. Hist., vol. 2, Pt. 8, No. 13, 1909,
pp. 213, 233) for records from outer Labrador.

» Kendall, Proc. U. S. Nat. Mus., vol. 38, 1910, p. 509.

" See Vladykov (Contrib. Canadian Biol., N. Ser., vol. 8, 1933, p. 30 (No. 2,
p. 18], as groenlandicus) for localities where it has been taken in Hudson Bay,
including James Bay.

Importance. — Although this is an edible fish
and accounted a good one, its appearance and
habits will probably close our markets to it as
long as other fish are plentiful. Nevertheless, it
once was of some commercial importance, being
one of the best baits for lobster pots, for which
purpose great numbers were speared formerly on
the Massachusetts coast in spring, and were caught
along the northern coast of the Gulf on hook and
line. But very little use is made of them nowa-
days, if any.

Longhorn sculpin Myoxocephalus octodecimspi-
nosus (Mitchill),87 1815

Gray sculpin; Hacklehead; Toadfish
Jordan and Evermann, 1896-1900, p. 1976.

Description. — This fish resembles the shorthorn
sculpin so closely that the description may be
confined to the points in which it differs. Chief
of these is the great length of its uppermost cheek
spine, which usually is about four times as long
as the spine just below, and which reaches at
least as far back as the edge of the gill cover.
This serves equally to distinguish the young
longhorn from the grubby, which is short-horned.
All the head spines, too, of the longhorn are so
sharp that one must be cautious in grasping one
of these fish, for it turns its spines rigidly outward
by spreading its gill covers. Furthermore the
long spines of the long horn are naked at the tip.
The number and arrangement of the head spines

17 Placed in the genus Acanthocottus Girard, 1849, by Jordan, Evermann,
and Clark (Kept. U. S. Comm. Fish., (1928), Pt. 2, 1930, p. 386.

Figure 234. — Longhorn sculpin (Myoxocephalus octodecimspinosus), New Jersey.
210941—53 30

From Goode. Drawing by H. L. Todd.

450

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

is the same as in the shorthorn sculpin (p. 445),
hence need not be described, and there are two
thorns on each shoulder, with a larger one close
above the origin of tbe pectoral fin. The first
dorsal fin is higher than the second (in the short-
horn sculpin these two fins are of about equal
heights), of rather different shape from that of
the shorthorn (compare fig. 234 with fig. 229), and
proportionately shorter than in the latter though
with about the same number of spines (8 or 9).

The second dorsal fin and the anal have the
same number of rays (15 or 16 dorsal and about
14 anal) as in the shorthorn; but the anal of the
longhorn originates under the second or third
ray of the second dorsal fin instead of under its
fourth or fifth ray. The pectorals are of the
fanlike form usual among sculpins. The lateral
line of the longhorn sculpin is marked by a series
of smooth cartilaginous plates instead of by
prickly scales as it is in the shorthorn, a difference
obvious to tbe touch; its body is more slender
(about five and one-half times as long as it is
deep) ; and its head is flatter.

Color. — The longhorn, like other sculpins, varies
in color with its surroundings. The ground tint
of the back and sides ranges from dark olive to
pale greenish-yellow, greenish-brown, or pale
mouse color, but is never red or black as the
shorthorn so often is. As a rule it is marked
with four irregular, obscure, dark crossbars, but
these are often broken up into blotches and they
may be indistinct. The coarseness of pattern
often corresponds to that of the bottom, as does
the degree of contrast between pale and dark.
On mud and sand bottom this sculpin is often
nearly plain colored, but when it is lying on
pebbles with white corallines its back is often
nearly white with dark-gray blotches, rendering
it almost invisible. The first dorsal fin is pale
sooty with pale and dark mottlings or spots; the
second dorsal is paler olive with three irregular
oblique dark crossbands; the caudal is pale gray;
and the pectorals yellowish. Both caudal and
pectorals are marked with 4 to 6 rather narrow
but distinct dark crossbands. The anal is pale
yellowish with dark mottlings; and there often is
an obscure yellowish band along the lower part
of the sides, marking the transition from the dark
upper parts to the pure white belly.

Size. — This is a smaller fish than the shorthorn
sculpin. It grows to a maximum length of about

18 inches, but only a few of them are more than
10 to 14 inches long. A 10-inch fish weighs about
% pound, one 12 inches long about 1 pound.

Habits.** — Everyone who has fished along the
shores of our Gulf is more or less f amiliar with this
sculpin, for it is a nuisance to cunner and flounder
fishermen. It often is bothersome to the angler to
unhook when it spreads its needle-sharp spines
and erects its spiny dorsal fin. It grunts when
pulled out of the water and bites on any bait.

No doubt it is as omnivorous as the shorthorn.
Specimens examined by Vinal Edwards at Woods
Hole had fed chiefly on shrimps, crabs, amphipods,
hydroids, annelid worms, mussels and sundry
other mollusks, squids, ascidians, and on a con-
siderable list of fish fry, including alewives, din-
ners, eels, mummichogs, herring, mackerel, men-
haden, puffers, launce, scup, silversides, smelts,
tomcod, silver hake, and small fry of other
sculpins. Rock crabs (Cancer irroratus) and
amphipod crustaceans (Leptocheirus) had been
the dominant food of a large series of shorthorns
in Block Island Sound, examined by Morrow;
nearly all of them had eaten shrimps (Crago) but
in small amount; a few contained small lobsters;
and spider crabs (Libinia) were a regular article
of diet in winter, but not in summer. It is
interesting that these particular shorthorns had
eaten only a few mollusks of any kind.89

The longhorn is as useful a scavenger as the
shorthorn, and equally voracious, gathering about
wharves, sardine factories, and under lobster cars,
always keeping to the bottom. Its depth range
is rather wider than that of the shorthorn. At
the one extreme it is abundant in many shoal
harbors and bays, where it comes up on the flats
at high tide, to leave them at low; and it runs up
into estuaries, salt creeks, and river mouths,
though never into fresh water, so far as we know.
At the other extreme it is caught in considerable
numbers down to 50 fathoms or so, and it has
been reported as deep as 105 fathoms.90

The longhorn evidently is at home in tempera-
tures as high as about 65°-66°, for we have seen
many of them in very shallow water that warm, or

» Morrow (Bull. Bingham Oceanographic Coll., vol. 13, Art. 2, 1951)
has recently published a detailed study of this sculpin, as found off southern
New England.

•' For more extensive diet lists see Morrow, Bull. Bingham Oceanographio
Coll., vol. 13, Art. 2, 1951, pp. 60-61, 88-89.

■• In Trinity Bay, east coast of Newfoundland, Kept. Newfoundland Fish.
Res. Comm., vol. I, No. 4, 1932, p. 108, Sta. 35.

FISHES OF THE GULF OF MAINE

451

even a little warmer, in summer in the southern
side of Massachusetts Bay. But in localities
where the temperature of the upper few feet rises
much higher than this they withdraw to some-
what deeper (i. e., cooler) water for the summer
(p. 452), working inshore again in the autumn.

At the other extreme, it is subjected for the
coldest part of the year to water as cold as 32°-33°,
both in our Gulf, along the Nova Scotian shelf, and
in the Gulf of St. Lawrence, while it has been
reported from water of 31°-32° F. (-0.3° C.) in
the bottom of Trinity Bay, Newfoundland. And
it seems that even exposure to freezing tempera-
ture may not be fatal to it, if not too prolonged,
for we find no evidence that these sculpins are
ever killed by cold when they are overtaken on the
flats in severe freezes, a fate that occasionally
overtakes cunners and tautog (pp. 475, 480). But
the fact that the geographic range of the longhorn
does not reach as far north as that of the short-
horn suggests that it is not so well suited as the
latter is to very low temperatures continuing
throughout the year.

Off the southern New England coast the short-
horn deposits its eggs from late November
through January, and perhaps into February, i. e.,
at the coldest time of the year, with the chief
production in late December and January.01
Presumably the spawning season is the same in
the Gulf of Maine. Apparently one locality
serves as well as another, nor is there any evidence
that any particular depth is sought.

Ripe eggs are about 0.85 mm. in diameter before
being laid, but they swell when they come in
contact with the water; they are described as
varying in color, from coppery green to reddish
brown, orange, or purple. A 12%-inch female,
which we examined, taken near Woods Hole on
November 18, 1951, contained about 8,000 choco-
late brown eggs. They sink and they are so
sticky when first laid that they cling together in
clumps, or to anything that they may touch; and
they continue to adhere during the period of in-
cubation, but the surfaces of the eggs that are
exposed to the water lose their stickiness after
about 24 hours. The egg masses have been found
free on the bottom, in empty clamshells or other
cavities, or among the branches of the finger sponge

11 For the most detailed study yet made of the breeding habits of the short-
horn, see Morrow, Bull. Bingham Oceanographic Coll., vol. 13, art. 2, 1951,
p. 30-36.

(Chalina) like the eggs of the sea raven (p. 456) n
and they are sometimes found thrown up on the
beach.

The young fry have been taken in February and
March off southern New England, in April on the
eastern part of Georges Bank and in the channel
between the latter and Browns Bank. These
young stages M have longer cheek spines than the
corresponding stages of the shorthorn sculpin
(p. 447) they are more slender, and they differ fur-
ther in the outline of the dorsal fin, for in the long-
horns (if our identifications be correct) this is
continuous from end to end, only the largest of
them showing a shallow notch between spiny
and soft portions, whereas in the shorthorn the two
sections are separate from the time the fin first
takes definite form.

Captures of many young fry IK to 2 inches long
in September, and 3 to 3}i inches long in February
suggest that the longhorn is about 2 to 2){ inches
long at one year of age. According to Morrow's
studies (based on the otoliths)94 longhorns off
southern New England average about 6K to 7
inches long at 2 years of age; about 10 inches at
4 years; and 11 to 12 inches at 6 years. They are
mature sexually at 3 years or older.

General range. — Coastal waters of eastern North
America from eastern Newfoundland,95 and the
north shore of the Gulf of St. Lawrence, 96 south
regularly to New Jersey, and reported to the
Atlantic coast of Virginia.97

Occurrence in the Gulf of Maine.— This is our
commonest sculpin, to be caught anywhere and
everywhere along the entire coast line of the Gulf
of Maine. We dare venture that there is not a
bay, harbor, estuary, or a fishing station from Cape
Sable to Cape Cod where it is not to be found.
Not only is it more plentiful in most places than
its short-horned relative, but it occupies a wider
depth zone. It is very abundant in many shoal
harbors where it comes up on the flats; it is caught

" Warfel and Merriman (Copeia, 1944, p. 198) were the first to report this
interesting habit.

" The smallest larva we have seen was 13 mm. long, from Georges Bank.

" Bull. Bingham Oceanographic Coll., vol. 13, art. 2, 1951, p. 47, table 6.

» Trinity Bay, Newfoundland; Kept. Newfoundland Fishery Research
Commission, vol. 1, No. 4, 1932, p. 108, sta. 35.

■• Repeated characterizations of this sculpin as ranging to "Labrador" are
based on Storer's (Boston Jour. Nat. Hist., vol. 6, 1852, p. 250) report of 2
young specimens from some point not specified on the northern shore of the
Gulf of St. Lawrence.

,T Albatroit II trawled them in small numbers at 9 stations scattered along
the midbelt of the shelf, from the offing of southern Massachusetts to the
offing of Delaware Bay, in the months of February, April, July, and Septem-
ber, at depths ranging from 11 to 50 fathoms.

452

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

also in considerable numbers down to 50 fatboms
or so (p. 450). We have trawled it at 27 to 33
fatboms in Massachusetts Bay, and at 50 fathoms
off Cape Elizabeth. But it has not been reported
from the basin at greater depths.

It also occurs plentifully on Georges Bank, and
while the composition of the sculpin population
of Georges Bank is yet to be determined, the facts
that this was the only sculpin (except the sea
raven which it greatly outnumbered) taken there
on otter trawling trips in June 1912, or in Septem-
ber 1929, and that the dragger Eugene H took
1,030 of them in 35 trawl hauls on the southwestern
part of Georges, at 25-35 fathoms (but none in
deeper hauls) in late June 1951, are evidence that
it is the commonest member of its tribe on shoal
parts of the bank. It is fair, also, to assume that
this applies equally to Browns Bank, where fisher-
men report sculpins of one sort or another as
plentiful. It is described, also, as very common
along the Nova Scotian coast and banks eastward
from Cape Sable, in suitable depths, and as widely
but irregularly distributed around the southern
shores and islands of the Gulf of St. Lawrence.08

The longhorn is a year-round resident, in the
sense that its only periodic movements are off
and on shore, and of short extent, combined with
movements to and from particular grounds. Near
New York it is commonest near shore from Sep-
tember to May, and is seen only occasionally in
summer. In Long Island Sound they appear to
carry out east-west journeys about which little
is known; in Block Island Sound (off the mouth
of Long Island Sound) they are plentiful on the
productive fishing grounds from November
throughApril, but mostly withdraw thence during
May (either offshore, or onto more rocky grounds
nearby), not to return in force until the next
October." These shifts do not appear to be
connected with temperature.

All that is known of its movements in the Gulf
of Maine is that in partially enclosed and very
shallow situations where the water on the flats
heats to 68°-70° in the warmest part of the
season, but where ice forms in the winter (Dux-
bury Bay, for example), the shorthorns seek
slightly deeper (i. e., cooler) water for the summer,

■• Co*, Contrlb. Canadian Biol. (1918-1920) 1021, p. Ill; Leim, Proc. Nova
Scotian Inst. Sci., vol. 20, 1940, p. 40.

'• For further details see Morrow. Bull, Bingham Oceanographic Coll.,
vol. 13, art. 2, 1951, p. 51

work up again onto the flats in early autumn;
move deeper again in late autumn; then work
back on the flats again in early spring. They
continue common, however, right up to low tide
line all summer in localities where the surface
does not become so warm in summer or so cold
in winter. This is the general rule northward
and eastward around the coast of Maine, includ-
ing the Passamaquoddy region.

The presence of longhorn sculpins of all sizes,
from very young fry to adult, proves that they
breed all along the coasts of Massachusetts and
of Maine, probably along western Nova Scotia
as well. But it seems to be restricted as a breeder
in the Bay of Fundy to the Scotian side. Thus it
appears that the half-grown and adult fish that
are plentiful along the New Brunswick shore are
migrants, either from the Nova Scotian side
across the bay, or from the open Gulf outside.

Importance.— -The only commercial value this
sculpin has had in our Gulf was as bait for
lobster pots, for which they were speared formerly
in some localities, and caught on hook and line
in others. But very few of them are now used
in this way.

Staghorn sculpin Gymnocanthus tricusjns
(Reinhardt) 1832
Jordan and Evermann, 1896-1900, p. 2008.

Description. — This sculpin is easily distinguish-
able from its more common relatives by the shape
of the uppermost of its three cheek spines, which
is broad, flat, and with three short, sharp branches
at its tip ' instead of cylindrical and single pointed;
also its anal fin (16 to 18 rays) originates well in
front of its second dorsal fin instead of behind
the latter, and its two dorsal fins are separated
by a distinct space instead of being practically
continuous at the bottom of the notch that
separates them. Furthermore, the spines char-
acteristic of the top of the head and shoulders of
our other sculpins are either lacking on the stag-
horn or are very short, and the corners of its gill
covers are rounded instead of sharp. Distinctive
also, if less obvious, is the fact that the top of its
head is more or less prickly or warty.

The 3-rayed ventral fins reach only about to
the vent on young fry of 1% to 2 inches, but they

1 These branches are only faintly indicated at the tips of the spatulate
spines on young fry that we have seen, 1H to 2 inches long.

FISHES OF THE GULF OF MAINE

453

Figure 235. — Staghorn sculpin {Gymnocanlhus tricuspis). After Smitt.

become relatively longer with growth until at ma-
turity they reach considerably beyond the point
of origin of the anal fin, farther in males of breed-
ing age than in females.2 The first dorsal fin has
11 or 12 spines; the second dorsal 15 to 17 soft
rays. The caudal and pectoral fins and the gen-
eral shape of the fish are of the usual sculpin type.

Color. — Described as dark brownish or gray
above, the sides as marked with dark crossbands
or with alternate light and dark greenish spots;
the lower surface as white or yellowish with an
irregular line of demarkation between dark sides
and pale belly. The dorsal and pectoral fins are
pale, the former with 3, and the latter with 4 or 5
irregular dark brown or black crossbands. The
ventral and anal fins are yellow rayed, with mem-
branes of the same color as the belly.

Size. — Up to about 10 inches long.

General range. — Arctic Ocean and North At-
lantic, south to northern Norway on the European
coast; on the American coast it ranges southward
along the outer coast of Labrador 3 to the Gulf
of St. Lawrence, where it is generally distributed
along the north shore * and is characteristic of the
icy water on the banks in the southern side, accord-
ing to Huntsman, and it has been reported as far
as Eastport, Maine, as a stray.

Occurrence in the Gulf of Maine. — The most
southerly record for this Arctic sculpin, and the
only one for the Gulf of Maine, is of a specimen
caught at Eastport, Maine, in 1872, and now in
the United States National Museum. It is only

• Small fry that we have examined from Saeglek Bay on the northeastern
coast of Labrador, and an adult male from the estuary of the St. Lawrence
River, agree with Smitt's (Scandinavian Fishes, vol. 1, 1892, p. ICO as O.
veniralis) account of a specimen from Spitzbergen.

3 Presumably it also occurs all around the coasts of Newfoundland; but
it is not Included among the species listed as taken during the trawling cruises
of the N ewfoundland Fisheries Research Commission.

< Stearns, Proc. U. S. Nat. Mus., vol. 6, 1884, p. 125; Kendall, Proc. Port-
land Soc. Nat. Hist., vol. 2, No. 13, 1909, p. 213-214.

as a very rare stray from colder waters to the
north that it ever reaches our Gulf.

Arctic sculpin Cottunculus microps Collett 1875
Jordan and Evermann, 1896-1900, p. 1992.

Description. — The head spines so characteristic
of most sculpins, are reduced in this species to
bony knobs, of which there are four on the top
of the head and several on its sides. The two
portions of the dorsal fin (spiny and soft) are united
into one continuous fin, a feature that marks it
off from all other local sculpins, while the spiny
part (only 6 to 8 spines) is shorter and lower than
the soft part (13 to 15 rays). But the very large
bony head, wide mouth, slender tapering body,
large fan-shaped pectorals, and the location of the
ventrals below the pectorals, give the fish the typ-
ical sculpin aspect. The anal fin (about 10 rays)
is a little shorter than the soft portion of the dor-
sal fin, and the caudal fin is small and rounded.
The skin is roughened with small warts.

Color. — Described as pale with dusky crossbars,
one on the head, two on the body and fins, and
one at the base of the caudal fin. Scandinavian
specimens have been reported as having still an
other band across the tip of the caudal, and as
with the anal and pectoral fins dark mottled.6

Size. — Up to about 8 inches long.

General range and occurrence in the Gulf of
Maine. — This is an Arctic deep-water species,
known off east Greenland and about Spitzbergen
in the Arctic Ocean, and from both sides of the
northern Atlantic. On the eastern side it has been
reported from northern Iceland, from Norwegian
waters southward to the Channel, and doubtfully
from the Skagerak. Off the American coast it

' Smitt, Scandinavian Fishes, vol. 1, 1892, p. 163.

454

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 236. — Arctic sculpin (Cottunculus microps), continental slope off southern New England.

Drawing by H. L. Todd.

From Goode and Bean.

has been taken at numerous localities on the con-
tinental shelf and slope to abreast of New Jersey
in depths of 122 to 487 fathoms. Only two of the
earlier published records fall within the geographic
limits covered by this report, one in the extreme
southeast corner of the basin of the Gulf (latitude
42°23', longitude 66°23') in 141 fathoms, the other
in the eastern channel between Browns and
Georges Banks (latitude 42°15', longitude 65°48')
in 122 fathoms. But we trawled one about 2
inches long, on the northern slope of Georges
Bank, in 120 fathoms of water, on July 24, 1931,
which (with earlier captures) shows that it is to
be expected anywhere in the deep basin of our
Gulf, at depths greater than 100 fathoms.6
Nothing is known of its habits.

Sea raven Hemitripterus americanus (Gmelin)
1789

Red sculpin; Sea sculpin; Raven; King
o'Norway

Jordan and Evermann, 1896-1900, p. 2023.

Description. — No one would be likely to confuse
a sea raven with any other sculpin, for it is a
"most remarkable looking fish," as Jordan and
Evermann remarked.7 It is stouter bodied than
our other common sculpins, about three and three-
fourths times as long as it is deep (counting caudal
fin), with a very large head. Both the jaws of
its wide mouth are armed with several rows of
sharp teeth that are noticeably longer and stouter

• Goode and Bean, Smithsonian Contrib. Knowl; vol. 30, 1895, p. 270, list
the earlier American records.

' Bull. 47, U. S. Nat. Mus., Pt. 2, 1898, p. 2023.

Figure 237. — Sea raven (Hemitripterus americanus), Halifax, Nova Scotia. From Goode. Drawing by H. L. Todd.

FISHES OF THE GULF OF MAINE

455

than the teeth of either the long-horned sculpin or
of the short-horned sculpin. Its most distinctive
features, however, which identify it at a glance,
are the fleshy tabs, simple and branched, on its
head ; the curiously ragged outline of its first dorsal
fin; and the prickly texture of its skin. There is
a series of 4 to 8 of these tabs along each side of
the lower jaw, three pairs on the top of the snout,
and others, variable in number and size, above
and in front of the eyes and along the upper jaw.
There is also a short but high keel on the top of
the snout with a deep hollow behind it, another
high ridge above each eye, and a lower one below
the eye. These ridges, with about 12 rounded
knobs on the crown and several low bosses, and
2 short spines on each cheek, give the head a
peculiarly bony appearance.

The first two or three spines of the first dorsal
fin are the longest, and the fourth and fifth spines
are shorter than those farther back, giving the
fin an outline quite unlike that of any other
sculpin. And the fin membrane is deeply emargi-
nate between every two spines from the third
spine backward, but expanded at the tip of each
spine as an irregular flap of skin. The margin of
the anal fin is similarly, but less deeply scalloped
between the rays. Furthermore, the first dorsal
fin originates further forward than in any other
Gulf of Maine sculpin, i. e., well in front of the
gill opening, and it is much longer (16 spines) than
the second dorsal (1 spine and 12 rays), whereas
in our other sculpins the second dorsal is longer
than the first dorsal. The pectorals are fanlike,
and the caudal brush-shaped, much as in other
sculpins. The ventral fins are fleshy, each with
1 spine and 3 soft rays, with the first 2 rays so
close that there seems to be only 1 soft ray.

The entire skin of the sea raven is prickly,
belly as well as back, with the prickles largest on
the back and along the lateral line; smallest, but
still obvious to the touch, on the lower parts of
the sides and on the belly. In all other Gulf of
Maine sculpins the belly is smooth.

Color. — The sea raven varies in color from
blood red to reddish purple, chocolate, or to yellow-
ish brown, but it is invariably paler below than
above, and it usually has a yellow belly. Many
are plain colored. For instance, one 18 inches
long, which we caught off Mount Desert recently,
was uniform red chocolate on back and sides.

But others are variously mottled with a paler or
darker cast of the general ground tint, or even
with white. The fins are variously barred with
fight and dark, and the pectorals and anal often
are yellow-rayed.

Size. — One of the largest on record, of 25 inches
and 5 pounds, is mentioned by Storer. But this
specimen seems to have been in poor condition
for we have caught one 22^ inches long that
weighed 7 pounds. Warfel and Merriman 8 men-
tion one about 19)2 inches long that weighed 5.8
pounds inflated with water and 5.3 pounds when
emptied. And many sea ravens are 18 to 20
inches long.

Habits. — The sea raven alone, among Gulf of
Maine sculpins has the power of inflating its belly
with water, like a bladder. If released in this
condition it drifts helplessly, feebly waving its
tail to and fro, and we cannot say whether it can
empty itself again at will like a puffer (which,
however, inflates with either air or with water,
p. 526) or whether it must await the gradual escape
of the water it has swallowed. Another way in
which the raven differs from our other sculpins is
that it can bite sharply, having larger teeth.

The raven is quite as voracious as its relatives;
it takes any bait and is said to eat whatever
invertebrates it finds on the bottom, such as
mollusks (both bivalve and univalve), various
crustaceans, sea urchins, and worms. Sea ravens
also eat fish. Vinal Edwards found herring, launce,
sculpins, tautog, silver hake, and both sculpin and
sea-raven eggs in sea ravens taken at Woods Hole.

Our own experience, confirmed by our various
inquiries, is that ravens are to be caught only on
rocky ground (which is their chief haunt from
Massachusetts Bay northward), pebbles, hard
sand, or clay (which they frequent off Cape Cod
and on the offshore Banks), never on soft sticky
mud. There is no definite upper limit to their
vertical wanderings other than the surface. But
they are seldom caught within the smaller estu-
aries, perhaps never on the tidal flats at any time
of year; at least we have never seen them in such
situations in Massachusetts Bay, though they are
not uncommon there about the off-lying ledges.
The majority of them live deeper than a couple of
fathoms at all times. On the other hand their
usual range extends down only to about 50

• Copeia, 1944, p. 204.

456

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

fathoms, although one has been taken as deep
as 105 fathoms.9

The geographic and vertical ranges of the sea
raven suggest that the upper limit to its preferred
temperature is about 58°-60°. At the other
extreme their wide dispersal over the Magdalen
Shallows in the southern side of the Gulf of St.
Lawrence shows that they can winter in tempera-
ture close to the freezing point of salt water,
unless they descend then into considerably deeper
water, a possible shift in depth on their part of
which there is no direct evidence.

Presumably the sea raven breeds throughout its
geographic range. Off southern New England the
eggs are deposited from early October (earliest
date, October 2) on until late December; probably
in autumn and early winter in the more northern
part of its range as well.10 Warfel and Merriman,
who made a special study of the breeding of the
sea raven, have made the very interesting dis-
covery that it deposits its eggs chiefly at the bases
of the finger-like branches of the finger sponge
(Chalina) ; less often on the smaller sponge Hali-
chondria, where they stick together in clusters and
to the sponge. Since the eggs average only about
242 per cluster (minimum 141, maximum 478,
among many clusters counted), whereas adult
females contain something like 15,000 maturing
eggs on the average, and occasionally as many as
40,000, it appears that a female does not lay all
her eggs at one time, but deposits many clusters
during each spawning season.

The eggs are large, averaging 3.9 to 4 mm. in
diameter, with tough egg membrane, yellow when
first spawned, but soon changing to an amber hue;
so heavy that they sink; and very sticky and re-
sistant to injury.11 Eggs brought in to the labora-
tory by Warfel and Merriman hatched a few at a
time, and some of those of a cluster collected on
January 23 and left thereafter in a bottle fastened
to a buoy (i. e., in the normal winter temperature
of Long Island Sound) did not hatch until
March 12.

• A specimen about 14 inches (360 mm.) long was trawled by the U. S.
Fish and Wildlife vessel Delaware August 24, 1951, in 105 fathoms, latitude
42°06' N., longitude 67°50' W.

i° Warfel and Merriman (Copeia, 1944, p. 202) probably were correct in
their suggestion that a 20-inch female, containing ova of 2 sizes, caught by
us off Boothbay, Maine, in April, was simply one that had failed to spawn
at the proper time during the preceding winter.

» Described by Bean (Bull. 60, New York State Mus., Zool. 9, 1903, p. 647).
The buoyant eggs referred to this species by Agassiz and Whitman (Mem.
Mus. Comp. Zool., vol. 14, No. 1, 1885, p. 10) belonged to some other fish.

The sizes of the few young sea ravens that have
been taken in the Gulf of Maine suggest that
they reach a length of 2 to 4 inches by the middle
of their first summer, when 6 to 8 months old,
and about 6 inches by the following April, at
an age of 1% years. Their subsequent rate of
growth has not been followed.

General range. — Atlantic Coast of North America,
southward to Chesapeake Bay; north to Anticosti
in the northern side of the Gulf of St. Lawrence,
to the Strait of Belle Isle on the Newfoundland
side 12 and to the Grand Banks.13

Occurrence in the Gulf of Maine. — Sea ravens
are to be caught all around the coastal belt of the
Gulf, from a fathom or two down to about 50
fathoms, including the passages among the islands
that fringe the coasts of Maine and of Nova
Scotia, as well as the larger estuaries such as
St. Mary, Passamaquoddy, Machias, Penobscot
(it runs up the latter to the head at Bucksport),
and Casco Bays; also in the deeper harbors, for
example Boston, Salem, Eastport, and St. Johns.
Fishermen also report them on Cashes Ledge,
while the otter trawlers and long liners pick them
up in small numbers over the shoaler parts of
Georges Bank; likewise on Browns. But they are
not known to occur on the soft mud bottoms of the
deep troughs and basin of the Gulf.

Presumably the sea raven breeds in the Gulf
wherever it occurs, young fry having been taken
from the Bay of Fundy to Cape Cod.

Off the southern shores of New England sea
ravens work inshore in autumn and out again
into slightly deeper water in spring, but no season-
al movement of this sort (which means merely
that shoal water is too warm for their comfort in
summer) has been reported for them in the cooler
waters of the Gulf of Maine.

Although they are distributed so generally
in the Gulf, sea ravens are not so numerous as the
shorthorn sculpins and longhorn sculpins; this
is as true in the Bay of Fundy as it is in Mas-
sachusetts Bay, where one expects to catch a few
ravens around any of the fishing ledges, but
where it would be unusual for one person to land
any considerable number in a day. Similarly, the
schedules of the catches made bv certain otter

■» Jeflers (Contrib. Canadian Biol. N. Ser., vol. 7, No. 16, (Ser. A, No.
13,) 1932, p. 208).

" Bean (Proc. U. S. Nat. Mus., vol. 3, 1881, p. 86) lists a specimen from the
Grand Banks.

FISHES OF THE GULF OF MAINE

457

trawlers in 1913 and subsequently, added to our
trawling experiences, show that sea ravens are
much less numerous on Georges Bank than the
longhorn sculpin; it is seldom that as many as a
dozen are taken in a haul of the otter trawl.
Thus the dragger Eugene H took only 46 sea
ravens in 38 hawls on the southwestern part of
Georges, at 26-55 fathoms in late June 1951,
and none in her deeper hauls.

To the eastward and northward sea ravens are
described as common all along outer Nova
Scotia to Canso; they have been reported on
Sable Island Bank and on Banquereau Bank in
depths of about 20-30 fathoms; and they are
taken here and there on the Gulf of St. Lawrence
coast of Cape Breton. They are also reported
from Anticosti and in the Strait of Belle Isle
(footnote 12, p. 456); but there cannot be many

of them in the southern side of the Gulf, for they
are not mentioned in the published lists of Fishes
for Prince Edward Island, or around the Mag-
dalens. And there is only one report of a sea
raven on the Grand Banks (see footnote 13, p. 456) ;
and one for the Atlantic coast of Newfoundland,
i. e., from Trinity Bay.1*

To the westward and southward the sea raven
is reported as rather common as far as New York
and New Jersey; and it has been reported from
Chesapeake Bay. 16

Commercial importance. — Although the sea raven
is said to be a good table fish there is no more
market for it than for other sculpins in New Eng-
land or Canada. But it is generally considered
excellent bait for lobster pots, hence shore fisher-
men are likely to save what ravens they catch for
this purpose.

ALLIGATORFISHES. FAMILY AGONIDAE 18

These curious little fishes are related to the
sculpins anatomically, though their general ap-
pearance gives no hint of the fact. Their most
striking external feature is that the body is armed
with several rows of overlapping plates. The
only Gulf of Maine species somewhat suggests a
pipefish in this and in its slender form, but there
is no danger of confusing one with a pipefish,
for its mouth is of the ordinary form; and it has
ventral fins. Some agonids have a spiny dorsal
fin which others lack, while the ventral fins are
situated far forward (only a little rearward of the
pectorals) in all of them. Twenty-six of the
many species included in the family are known
from the western coast of North America from
Bering Sea to southern California; two are known
from the eastern coast of North America.

One of the eastern American species (Leptagonus
decagonus, Bloch and Schneider 1801), with two
dorsal fins, is Arctic, ranging southward only to
northern Nova Scotia;17 the other, with only one

11 Rept. Newfoundland Fish. Eos. Comra., vol. 1, No. 4, 1932, p. 108.

"Jordan, Evermann and Clark's (Rept. U. S. Comm. Fish. 1928 part 2,
1930, p. 393) reference of it to the Tortugas seems to have been an error.

18 Some recent authors separate this gToup into two families, Agonidae or
Sea Poachers for those with two dorsal fins, Aspidopboroididae or Alligator-
fishes for those with only one dorsal fin. But it seems preferable, for the
purposes of the present volume, to follow the older custom of uniting them
in the one family Agonidae.

17 A young specimen has been reported from Banquereau Bank (Rept.
Newfoundland Fish. Res. Comm., vol. 2, No. 1, 1933, p. 127) as Agonus
decagonus.

dorsal fin {Aspidophoroides monopterygius) is a
regular member of the Gulf of Maine fish fauna
(p. 457).

Alligatorfish Aspidophoroides monopterygius
(Bloch) 1786

Sea poacher

Jordan and Evermann, 1896-1900, p. 2091.

Description. — The readiest field marks for the
identification of this curious little fish (so odd in
appearance that it is not likely to be mistaken for
any other) are that its entire head and body are
clad with bony plates; that it has only one dorsal
fin (the soft rayed); aud that it is very slender
(about 12 to 13 times as long as it is deep, not
counting the caudal fin) , rather broader than deep,
and tapering rearward from the head to a very
slender caudal peduncle. The plates are smooth,
arranged in longitudinal rows. There is a double
row along the back running from close behind the
head to close behind the dorsal fin where it unites
into a single mid-dorsal row (altogether 45 to 50
double and single plates along the back); three
rows on the forward part of each side, but only
two rows along the rear part; and two rows along
the lower surface to just behind the anal fin, where
they unite in one row. Thus the trunk is octagonal
in front of the unpaired fins, but hexagonal behind

458

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

*V .

'v$t$,z ■"*t<^~-^^p

Figure 238. — Alligator fish (Aspidophoroides monopterygius). Top and center, adult, side and dorsal views, Halifax,
Nova Scotia; from Jordan and Evermann, drawing by H. L. Todd. Below, larva, 11 mm. long, Gloucester, Mass.

them. There are likewise two large plates and
several small ones in front of each pectoral fin.
The eyes are very large, with prominent ridges
above them, and there are two sharp recurved
spines on the top of the nose. The mouth is small
with minute teeth. The dorsal and anal fins (each
of five or six rays) are fanshaped, one over the
other, and are situated about midway of the truck.
The caudal fin is small and rounded, the pectorals
are larger than the anal, dorsal, or caudal fins, and
the ventral fins are reduced to one spine and two
rays each.

Color.— The many specimens we have seen have
been dark brown above, lighter brown below, with
two darker cross bands between the pectoral fins
and the dorsal fin; one cross band under the dorsal,
and two or three cross bands between the dorsal
and caudal fins. The dorsal and pectoral fins are
more or less barred; the caudal is dusky.

Size. — Five to seven inches long when full
grown.

Habits. — Nothing whatever is known of the life
of the alligatorfish except that it is a bottom fish
and that it has been repeatedly found in the
stomachs of cod, haddock, and halibut although
it is not "much thicker or softer than an iron
spike." I8 The Grampus and the Albatross II have
trawled it both on pebbly bottom, on sand and
broken shells, and on soft mud. So far as known
adults never stray into water shoaler than 10 to
15 fathoms, and the deepest record for it, with
which we are acquainted, is from 104 fath-

u Goode, Fi»h. Ind. U. 8., Sect. 1, 1884, p. 258.

oms." Its range shows that it is a cold water fish.
Its upper temperature limit is about 50°-52°; its
lower limit close to the freezing point of salt water.
Its breeding habits are unknown. Probably its
eggs sink like those of sculpins.

The presence of its larvae in Passamaquoddy
Bay, off Boothbay, and near Seal Island, Nova
Scotia, from April to June, points to late autumn
and early winter as the spawning season. It does
not take to the bottom until of considerable size,
for we have taken young ones as long as 29 mm. in
our towings.20

General range. — From west Greenland and the
east coast of Labrador southward to Cape Cod, and
to northern New Jersey as a stray.

Occurrence in the Gulf of Maine. — Being of no
interest to fishermen, and liviog too deep to strand
on the beach, this fish is seldom reported. It has
been taken in the Bay of Fundy in 15 to 100
fathoms; in the inner part of Passamaquoddy Bay;
near Eastport; off Mount Desert in 60 fathoms;
off Monhegan; near Portland; in 30 fathoms off
Casco Bay; on Cashes Ledge; in Ipswich Bay; off
Gloucester, Nahant, and Boston in Massachusetts
Bay; off Provincetown; off Chatham; and in the
South Channel to the eastward of Cape Cod.
Evidently it may be expected anywhere in the Gulf
in depths of 10 to 100 fathoms, and perhaps deeper.

Goode and Bean 21 described it as "abundant"
in the deeper parts of Massachusetts Bay, but our

» Southeastern slope of Browns Bank, latitude 42°20' N; longitude 65°08'
W; Goode and Bean Smithsonian, Contrib. Knowl., vol. 30, 1895, p. 284.
« Bigelow, Bull. Mus. Comp. Zool., vol. 61, 1917, p. 272.
s> Bull. Essex Inst., vol. 11, 1879, p. 13.

FISHES OF THE GULF OF MAINE

459

experience on the Grampus suggests "not uncom-
mon" as a better description, for our largest catches
were 8 specimens from 32 fathoms in Ipswich Bay;
6 specimens from 27 fathoms in the inner part of
Massachusetts Bay ; and 14 specimens off Chatham,
in May 1930, from 28 fathoms.

Other recent catches in various parts of the
Gulf have been of 1 or 2 fish each. And the fact
that we found it at only 4 out of our 10 trawling
stations of 1912 (all in the western part of the
Gulf) is in line with Huntsman's statement that
it is found only occasionally in the Bay of Fundy.

It is perhaps more plentiful along the Nova
Scotian shelf eastward and northward from Cape
Sable.22 It is numerous enough in the southern
part of the Gulf of St. Lawrence for Huntsman M

to describe it as "characteristic" of the ice-cold
Banks water there; it has been reported in the
estuary of the St. Lawrence River near Trois
Pistoles; 24 and at several localities along the west
coast of Newfoundland. It is so widespread over
the eastern half of the Grand Banks, along
eastern Newfoundland, and off southeastern Lab-
rador that it is listed at 14 stations there from the
experimental trawlings of the Newfoundland
Fisheries Research Commission,25 and is reported
from Hamlton Inlet.26

The only records of it to the westward of Cape
Cod are of the head of one that was dredged off
Watch Hill, N. J., in 1874; " and of one that was
taken off Sandy Hook, N. J., in 1864 .2S

THE LUMPFISHES. FAMILY CYCLOPTERIDAE

The lumps are characterized among Gulf of
Maine fishes by their short, thick, high-arched
bodies; by having a bony sucking disc on the chest
with the very much reduced ventral fins at its
center; and by the fact that the skin is set with
tubercles.

KEY TO GULF OF MAINE LUMPFISHES

1. The body is roughly triangular in end view; the large

tubercles on each side are in widely separated rows:
the lower end of each gill opening is below the level
of the upper edge of the base of the corresponding
pectoral fin; the head (measured to the upper edge
of the bases of the pectoral fins) occupies only about
one-fourth of the length of the trunk; the first dorsal
fin is entirely concealed within the skin after the fish
is 2-3 inches long Common Lump Fish, p. 459

2. The body is nearly round in end view; the large tubercles

on each side are close set, in irregular pattern; the
lower end of each gill opening is considerably above
the level of the upper edge of the base of the cor-
responding pectoral fin; the head (measured to the
upper edge of the bases of the pectoral fins) occupies
about one-third of the length of the trunk; the first
dorsal fin (though more or less fleshy) continues ex-
posed throughout life Spiny Lumpfish, p. 463

Lumpfish Cyclopterus lumpus Linnaeus 1758

Lump; Lump sucker

Jordan and Evermann, 1896-1900, p. 2096.

Description. — The lumpfish is about twice as
long (counting its caudal fin) as it is deep, with a
short head, and the dorsal profile of its trunk is

much more arched than the ventral. There are 7
longitudinal ridges on its body. One of these
ridges runs along the back as a cartilaginous flap
that incloses the first dorsal fin in adults and that
continues rearward as 2 ridges from the first dorsal
fin to the second dorsal fin. There also are 3
ridges along each side, one of them over the eye,
one close above the level of the pectoral fin, and
one marking the line of transition between side
and belly. Each of these ridges is marked by a
line of large pointed tubercles, and the entire skin
between the ridges is thickly studded with small
knobs. The presence of these ridges makes the
trunk of the lumpfish roughly triangular in end
view, with flat belly (except when swollen by milt
or roe) and sharp back, but the caudal peduncle
is rounded. The profile of the head is characteris-
tic, being concave above, convex below, with
mouth at the tip of the snout. The teeth are
small and the eyes and gill openings of moderate
size.

The first dorsal fin (visible only on very small
specimens) is of 6 to 8 spines. The second dorsal
fin and the anal fin below it are alike in outline,
both of them of 9 to 1 1 rays. The caudal is broad

» Vladykov and MacKenzie (Proc. Nova Scotian Inst. Sci., vol. 19, 1935,
p. 97) describe it as "very common" there.

» Trans. Roy. Soc. Canada Ser. 3, vol. 5, sect. 4, 1918, p. 63.

" Vladykov and Tremblay, Natural. Canad., vol. 62, 1935, p. 80.

*■ Annual Reports, Newfoundland Fisheries Research Commission, vol. 1,
No. 4, 1932, p. 108; vol. 2, No. 1, 1933, p. 126; vol. 2, No. 2, 1934, p. 116.

» Kendall, Proc. Portland Soc. Nat. Hist., vol. 2, No. 13, 1909, p. 214.

« Goode and Bean, Smithsonian Contrib. Knowl., vol. 30, 1895, p. 284.

>• Abbot, Geology of New Jersey, 1868, p. 816'

460

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Sir : ■:,*:.- •.'..•■ >5tt& vV^/fclP^

■^ frtA

i.^r-y 'rS?v

Figure 239. — Lumpfish (Cycloplerus lumpus), Eastport, Maine. From Goode. Drawing by H. L. Todd.

based and square tipped or slightly convex; the
pectorals are large, rounded, and so broad based
that they nearly meet on the throat, and they are
larger on males than on females. The ventral
fins are not visible as such, being altered into 6
pairs of fleshy knobs, surrounded by a roughly
circular flap of skin. The entire disk, so formed,
(sucking disk) is about as wide as the width of the
head, and is situated close behind the throat.

Color. — Descriptions of this fish credit it with
a great variety of tints, which we can corroborate.
On adults the ground tint may be bluish gray,
olive, brownish or yellow green, chocolate or kelp
brown, or slaty blue, the belly usually being of a
paler or more yellowish cast of the same hue, but
sometimes whitish. On some specimens the back
and sides are marked with dark blotches and more
or less dotted with black. Others, however, are
plain colored or nearly so, except that the tubercles
are usually dark tipped. Young lumpfish (and it
is with such that we are most familiar) often
match their surroundings very closely in color,
usually being mottled olive green and ochre yellow
with silvery dots and stripings. Males, when
mature, are more vividly colored than females,
and their bellies turn red (brightest near the
sucking disk) during the breeding season.

Size. — The longest lumpfish so far recorded
from the American coast measured 23 inches, and
weighed 13K pounds; the heaviest weighed 20
pounds but measured only 21^ inches (both from

Orient, N. Y.), and the proportion of weight to
length varies similarly in smaller fish. Few are
longer than 14 to 16 inches, or heavier than 3 to 6
pounds while the largest we ourselves have seen
was about 15 inches long.29 Females average
larger than males. Fulton, for example,30 writes
that 39 females taken in the Bay of Nigg (Scot-
land) averaged about 16 inches and 6 pounds, 30
males only 11 inches and a little less than 2 pounds.
Habits. — Although the lumpfish is ungainly, it
can swim more rapidly for a short distance by
vigorous tail strokes than its shape might suggest,
and its young pelagic fry are very active. The
adult lump is primarily a bottom fish, but is also
made semipelagic by its habit of hiding in floating
masses of rockweed. In European seas it ranges
from tide mark down to 150-200 fathoms, but
we have never heard of one taken in more than
a few fathoms in the Gulf of Maine. Perhaps it
is the nature of the bottom that restricts them
there for the soft sticky mud that floors the deeper
basins, at least in the western side,31 can hardly
be a favorable environment for them. Large
lumpfish are often found hiding among rockweed
or holding fast by the sucker to stones or other

26 Smitt (Scandinavian Fishes, vol. 1, 1892, p. 294) gives 24 inches as'the
maximum for Scandinavian and European waters generally, apparently'not
accepting the enormous size (to 48 inches) credited to it by Mobiusjand
Heincke (Vierter Bericht, Komm. wiss. Untersucb. deutschen Meere, Kiel,
18S3, p. 226).

» Twenty-fourth Ann. Report, Fisheries Board Scotland, (1905) 1906, Ft.
3, p. 171.

« The eastern trough of the Gulf has a harder bottom.

FISHES OF THE GULF OF MAINE

461

objects. About Massachusetts Bay, lobster pots
are favorite resorts for them when set on stony
bottom. For instance, W. F. Clapp tells us that
one pot in every 8 or 10 will yield a lumpfish on
the broken ground off the entrance to Duxbury
Harbor, and they often hold to the lower sides of
lobster cars, probably for shade. Occasionally
one is found clinging to one of the poles of a trap
or wen, though this is a much less common event
in the Gulf of Maine than it is in Scottish waters,
where they are frequently caught in salmon nets
set along shore. Welsh notes one entangled in a
gill net set off Great Boars Head in April 1913;
they have (rarely) been found clinging to floating
logs or inside a floating box or barrel; sometimes
they strand on the beach; and there is at least one
record of a lumpfish clinging to a mackerel.

So far as known the only regular migrations
carried out by the lumpfish are the involuntary
drifts of its young fry at the surface, and a general
movement of the adults into shoal water at spawn-
ing time followed by an offshore movement after-
ward.

The young fry swim at the surface, and we
have taken them so often in our tow nets that
we have learned to expect them wherever there
are floating masses of rockweed (a refuge in which
all but the smallest regularly hide, or to the fronds
of which they cling).

Most of the lumpfish taken in tow nets or
dipped up are less than 2 inches long, but large
adults are sometimes seen at the surface; more
often, perhaps, in the Bay of Fundy than else-
where in the Gulf, their presence at the surface
being determined less by the age of the individual
fish than by the presence or absence of floating
seaweed. However, most of the young lumpfish
have left the surface by winter; indeed very few
have been taken at any depth in the Gulf of Maine
during the cold months,32 but we picked up one
on the surface off Lurcher Shoal on April 12, 1920,
and another off Yarmouth, Nova Scotia, on Jan-
uary 4, 1921.

Food. — We have no first-hand information to
offer as to the diet of the lumpfish. In British
waters this has been found to consist chiefly of
isopods, amphipods, and other small crustaceans,
with various other invertebrates, including worms

and soft-bodied mollusks. And its diet is much
the same in the Gulf of Maine for Cox and Ander-
son 33 report euphausiid shrimps (Meganycti-
phanes), fragments of jellyfish (Aurelia), amphipod
crustaceans (Hyperia), caprellid crustaceans, with
the remains of small fish in the stomachs of lumps
from Passamaquoddy Bay. And large numbers
of young clupeids have occasionally been found
in their stomachs. This, too, is one of the few
fish that regularly feed on ctenophores and on
Medusae. Thus 25 specimens examined at Woods
Hole by Vinal Edwards contained nothing but
ctenophores. But lumps, like most other fishes,
cease feeding during the spawning season. The
lumpfish, in its turn, is said to be a favorite food
of seals. Certainly it is so weak a swimmer that
it would fall an easy prey to them.

In Scottish waters, where many observations
have been made on the life of the lumpfish 34
spawning (and the corresponding inshore migra-
tion) takes place from February until near the
end of May. And the evidence afforded by our
tow nettings suggests an equally protracted
spawning season in the Gulf of Maine, for while
we have taken larvae 27 mm. long as early as
May 10, we have taken newly hatched larvae
(6 to 7 mm.) as late as June 19 in the inner parts
of the Gulf and as late as July 23 on the northeast
part of Georges Bank, with one only 10.5 mm.
long on August 22 off Seguin Island. In the
Gulf of St. Lawrence, however, where vernal
warming is later than in the Gulf of Maine, lump-
fish probably do not commence spawning until
the. middle of April, for Cox and Anderson found
no larvae there until late in June, with their
observations pointing to late May as the height
of the breeding season and to mid-June as about
its termination.35 Presumably its period of incu-
bation is about as long with us as it is in European
waters of like temperature, i. e., 6 weeks to 2
months.

On the other side of the Atlantic the lumpfish
spawns in very shallow water, chiefly close to
low-tide mark. But the fact that the egg masses
(more or less familiar objects on European shores)

« Cox and Anderson (Contrib. Canadian Biol., N. Ser., vol. 1, 1922, p. 5)
state that the Canadian Research steamer Prince has taken only two lump-
fish in the Bay of Fundy in winter, both of them small.

» Contrib. Canadian Biol. N. Ser., vol. 1, 1922, p. 9.

« Mcintosh, 14th Ann. Rept., Fishery Board Scotland, (1895) 1898, Pt. 3,
pp. 173-178, and Fulton, 24th Ann. Rcpt., Fishery Board Scotland, (1905)
1906, Pt. 3, pp. 169-178.

M The lumpfish spawns from late May through June on the coast of Green-
land; in April and May in the Baltic; and early in the spring in Norwegian
waters.

462

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

seem never to have been reported along the coast
of the Gulf of Maine, although the local presence
of larvae is proof that lumpfish breed all around
its coast line, suggests that the eggs are deposited
at least a fathom or two down with us. And our
capture of recently hatched larvae over Georges
Bank is evidence that the latter also serves as a
spawning ground in 15 to 25 fathoms or deeper.
But the lower depth limit to spawning is yet to
be determined.

Large females of 18 inches may produce up to
136,000 eggs which sink and stick together in large
spongy masses through which the water circulates
freely. Off the coasts of northern Europe these
egg masses are often found adhering to rocks or
other objects, or in crannies near low- water mark,
and watch should be kept for them along the
rocky coast line of the Gulf of Maine.

The male lumpfish guards the eggs until they
hatch; his courage and devotion has often been
described.36 Throughout the period of guardian-
ship, which he performs fasting, he fans the egg
mass constantly, keeping it free of silt and bathed
in flowing water, never leaving it except to drive
off some intruder. But his vigil ends as soon as the
eggs are hatched, leaving him thin and exhausted.
The females take no part in guarding the eggs,
but are said to move out into deeper water once
they have finished spawning.

The eggs are 2.2 to 2.6 mm. in diameter; pink
when first laid but soon change to pale green or
yellow, and deepen in tint as development pro-
gresses. The larvae are about 4 to 7.4 mm. long at
hatching, shaped like a tadpole with a large head
and slender tail, swimming actively, and soon able
to cling to any bit of weed. At 34 mm. the tuber-
cles begin to appear, and the fry then show most of
the characters of the adult, except for the large
first dorsal fin and slender form.

Lumpfish larvae and fry of all sizes are to be
taken throughout the summer; the smaller ones
undoubtedly are from that season's hatch, but the
larger ones may be either those hatched earliest
that spring, or late in the preceding summer, for
the varying stage of development reached by
different individuals at various sizes proves that
the rate of growth varies widely. Thus Cox and
Anderson describe one Cape Breton specimen that

was only 33 mm. long in July, but that was so
mature in outline and in its dermal armature that
it must have been at least a year old, whereas they
found that in the Bay of Fundy the fry of the
year grow to 40 or 50 mm. by December with the
yearlings averaging about 58 mm. in July and
August. As they remark, the rate of growth is
apparently about the same in the Bay of Fundy
(probably in the Gulf of Maine as a whole) as it is
in Scottish waters, while Gulf of St. Lawrence
lumpfish correspond to those about Helgoland, in
their slower growth.

Figure 240.— Larva, 4-5 mm. After Garman.

•• Fulton (24th Ann. Report, Fishery Board Scotland, (1905) 1906, Pt. 3, p. 169)
gives an Interesting eyewitness account of spawning and/>f the guardianship
by the male parent over the eggs.

Figure 241. — Fry, 34 mm. After Garman.
Lumpfish (Cyclopterus lumpus)

Presumably all Gulf of Maine lumpfish upward
of 2% inches long are in their second year. Accord-
ing to Cox and Anderson lumpfish 3% to 4% inches
long are in their third year, those of about 10
inches in their fifth year.37 But lumpfish grow
much faster if fed to capacity than on the rations
ordinarily available to them; one kept in the
aquarium at St. Andrews increased in length from
about 3.8 inches to about 12 inches in a little more
than 12 months, which is as much as is to expected
in 2 or 3 years under natural conditions.38 Prob-
ably maturity is attained in the third year.

General range. — Both sides of the North At-
lantic; White Sea, northern Norway and Iceland
to the Bay of Biscay and occasionally to Portugal
in the east (including the Baltic); northward in
the west to the Gulf of St. Lawrence, Newfound-
land, outer coast of Labrador, Hudson Bay,
Davis Strait, and Disko (latitude about 70° N.)

" As estimated from the structure of their vertebrae.

« According to McKenzle (Proc. Nova Scotlan Inst. Sci., vol. 20, 1939, p. 17)
this fish was kept well-fed on chopped clams and herring, but perhaps not to
capacity for its appetite seemed Insatiable.

PISHES OF THE GULP OF MAINE

463

in west Greenland; southward to New Jersey, and
to Chesapeake Bay as a stray.

Occurrence in the Qulj of Maine. — The lumpfish
is common along the outer coast of Nova Scotia
and is to be found all around the shores of the
Gulf of Maine. It has been reported at Yarmouth
and in St. Mary Bay on the Nova Scotian side,
and is abundant in all stages at various localities
in the Bay of Fundy. There are many records for
it along the Maine coast, including Eastport,
Penobscot Bay, vicinity of Boothbay, the offing of
Seguin Island, and Casco Bay; also in Massa-
chusetts waters where it has been reported re-
peatedly, as at Nahant, Swampscott, Cohasset,
Plymouth, Truro, along Cape Cod, and at Mono-
moy while we once picked up a lumpfish in the
deep bowl between Jeffreys Ledge and the coast,
probably as the trawl neared the surface.39 It
even enters river mouths, but it is never found
where the water is appreciably brackish, so far as
we can learn. According to fishermen large lump-
fish are seldom seen on the offshore bank?.

Importance. — The lumpfish is never eaten in the
United States, but it sometimes finds its way to
market as a curiosity. At one time a few were
consumed locally in parts of the British Isles, and
may still be.

Spiny lumpfish Eumicrotremus spinosus
(Miiller) 1777

Jordan and Evermann, 1896-1900, p. 2098.

Description. — The spiny lumpfish is easily dis-
tinguishable from its commoner relative by the
fact that its skin tubercles are not only much
larger relatively, and studded from the base to
the tip with rough prickles, but are irregularly
and closely scattered over the body and head.
Furthermore, its gill openings are much shorter,
while its body is not so high-arched, and is nearly
round in cross section, instead of triangular. Also,
its first dorsal (though fleshy in some of them)
retains its fin-like appearance through life, instead
of becoming entirely concealed by the skin.

Color. — Described as olivaceous to brownish.

General Range.- — Arctic and northern parts of
the Atlantic Ocean, south to the Gulf of Maine
as a stray.

Occurrence in the Gulj of Maine. — Stray speci-
mens of this northern fish have been reported
from Eastport, Maine; from off Cape Ann; and
from Salem, in the north side of Massachusetts
Bay. We have also seen three small specimens,
1-lM inches long that were collected about 15
miles southeast of Cape Ann in 23 and 29 fathoms,
by the U. S. Fish Commission in 1878 (now in the
U. S. National Museum.) 40

» Blgelow and Schroeder, Biol. Bull., vol. 76, 1939, p. 309.

» Myers and Boblke (Stanford Ichth. Bui!., vol. 3, No. 4, 1960, p. 199)
have described a new species of spiny lumpfish (B. terrae-novae) from New-
foundland. But the specimens from off Salem and from off Eastport pic-
tured by Goode and Bean (Smithsonian Contrib. Knowl., vol. 31, 1895,
pi. 70, fig. 250) and by Oarman (Mem. Mus. Comp. Zool., vol. 14, 1892,
p. 71, figs. 1-3) were typical spinosuf, as are the three small ones, from about
15 miles southeast of Cape Ann, that are mentioned above.

Figctbb 242. — Spiny lumpfish {Eumicrotremus spinosus), Massachusetts Bay specimen.

Drawing by H. L. Todd.

From Goode and Bean.

464

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

THE SEA SNAILS. FAMILY LIPARIDAE

The sea snails are tadpole-shaped, soft-bodied
little fishes; and like the lumpfish (p. 459) most of
them have a sucking disk on the chest, supported
by the vestigial rays of the ventral fins.41 But
the skin is smooth, and without tubercles, and
the spiny and soft parts of the dorsal fin are
continuous as a single fin. The more than 115
species that are known are widely distributed in
Arctic, North Temperate, and Antarctic Seas,
and from the intertidal zone down to 2,000 fath-
oms or so.42 The Gulf of Maine harbors two
species. A third {Careproctus ranulus Goode and
Beane, 1879) is known only from the vicinity of
Halifax, Nova Scotia; from Middleground off east-
ern Nova Scotia,43 from the Grand Banks, and off
southeastern Newfoundland; perhaps from the es-
tuary of the St. Lawrence River, also.44

We include it in the following Key, on the chance
that it may be encountered in the deeper parts
of our Gulf, sooner or later.

KEY TO GULF OF MAINE SEA SNAILS

1. The spiny (front) and soft (rear) portions of the dorsal

fin are separated by a notch Sea snail, p. 464

There is no notch between the spiny portion and the
soft portion of the dorsal fin 2

2. The anal fin has only about as many rays (26-29) as

the pectoral fin (28-33); there are two separate
nostrils; the body is opaque, variously striped or

spotted Striped Sea Snail, p. 466

The anal fin has many more rays (at least 48)
than the pectoral (27-28) ; there is only a single
nostril; the body is translucent in life and
colorless Careproctus ranulus is

Sea snail Neoliparis atlanticus Jordan and
Evermann, 1898

Jordan and Evermann, 1896-1900, p. 2107.

Description. — Perhaps the most noticeable char-
acter of this and of the striped sea snail (p. 466)

« Some species of the genera Paratiparis and Amitra have lost the sucking
disk.

« See V. Burke (Bull. 150, U. S. Nat. Mus., 1930) for a study of the family
as a whole, giving descriptions and geographic ranges of all known species.

''• McKenzie and Homans, Proc. Nova Scotian Inst. Sci., vol. 19, 1938,
p. 278.

" We cannot judge whether the Careproctus reported by Vladykov and
Tremblay (Natural. Canad., vol. 62 (ser. 3, vol. 6), 1935, p. 81) from the
estuary of the St. Lawrence River as C. longipinnis was indeed Identical with
the fish from north of the Faroe Islands that was described under that name
by C. V. Burke (Ann. Mag. Nat. Hist. Ser. 8, vol. 9, 1912, p. 510), or whether
it is referable to ranulus: as seems the more likely on geographic grounds.

" For a detailed description, with illustrations, see Goode and Bean,
8mithson. Contrib. Knowl., vol. 30, 1895, p. 275, vol. 31, pi. 70, figs. 251-2518,
251b.

is that it is shaped more like a tadpole than like
the conventional fish, thanks to the cylindrical
fore part of the trunk, together with the broad
rounded snout and fat soft belly, and the abrupt
flattening (sidewise) of the body close behind
the vent. It is also provided with a sucking disk
similar to that of the lumpfish (p. 459). In side
view the body is deepest abreast the pectoral fins
(about four times as long as it is deep, not count-
ing the caudal fin), tapering evenly to a moderate
caudal peduncle. The head is flat above, the
mouth is at the tip of the snout and moderately
wide, and the jaws are armed with many small
teeth arranged m bands.

<5

^m^^m^

Figure 243. — Sea snail (Neoliparis atlanticus), side view
(above) and ventral view (below) of adult male. After
Garman.

The dorsal fin (6 spines and 26 soft rays) origi-
nates close behind the pectorals and runs con-
tinuously to the base of the caudal though it is
separate from the latter. The dorsal spines are
longer in males than in females, and project further
beyond the membrane, giving the fin a fringed
appearance. The anal fin (23 to 27 rays) originates
under or a little behind the origin of the soft
portion of the dorsal fin to which it corresponds in
size and outline. The pectoral fins are not only
very large and fanlike, but their bases run forward
under the throat, where they expand into second-
ary lobes or wings with fringed edges. The ven-
tral fins appear only as a circle of low knobs in the
center of the sucking disk, which is situated on
the throat between the pectoral fins. The skin
is scaleless, and it is smooth except at spawning
time, when the male becomes rough with small
prickles.

FISHES OF THE GULF OF MAINE

465

Color. — Olive to reddish brown with lighter and
darker cloudings and dots, the dorsal and anal
fins often with crossbars. Its tints vary widely,
also its markings and the strength of coloration,
whether pale or dark, as is th^. case with so many
fishes that live on the bottom.

Size. — Maximum length about 5 inches.

Remarks. — This sea snail resembles the follow-
ing species (p. 466) so closely that the two are likely
to be confused. The most evident difference
between them is that the spiny portion of the dor-
sal (6 spines hardly stiffer than the soft rays) is
marked off from the much longer soft part (22 to
27 rays) by a slight notch in the present species,
whereas there is no such separation in the striped
sea snail.

Habits. — Sea snails are inconspicuous little fish,
usually found coiled up (tail to head) under stones,
or attached by the sucker to some kelp stalk or
other seaweed. But sometimes they swim to the
upper water layers: thus the Halcyon specimens,
for instance, mentioned below, were taken in a
tow net at 8 fathoms, where the water was about
22 fathoms deep. And they seem to be indifferent
to depth within moderate limits, for while they
have been dredged as deep as 50 fathoms at vari-
ous localities in our Gulf, they are often found
clinging to lobster pots in the Bay of Fundy, (p.
465) , while they have been taken in only a few feet
of water in Massachusetts Bay. One was even
found in a tide pool near St. Andrews, in Passa-
maquoddy Bay46 so it would not be astonishing to
find sea snails left in rock pools elsewhere, or on
pebbly beaches by the ebbing tide, as often hap-
pens with its European representative. Young
ones have been found living within the shells of
the giant scallop (Pecten magellanicus) , a curious
habit that they share with the striped sea snail
(p. 466) and with the hakes of the genus Urophycis
(p. 224). Little else is known of the fife of this
sea snail, except that it is supposed to work in-
shore in winter to spawn. Presumably it feeds
chiefly on small crustaceans and on small shellfish
as its European relative does.

The spawning of this species has not been ob-
served, but probably it takes place from March
until midsummer in our Gulf, for Huntsman
found larvae in Passamaquoddy Bay as early as

April, while we towed one only 7 mm. long on
German Bank as late as September 2 (1915).

The developmental stages of our sea snail have
not been seen. The eggs of the European sea
snail, which are about 1.1 mm. in diameter, and
pale straw color to light salmon pink, sink and
stick together, also to hydroids, seaweeds, and to
debris of any kind. The little clusters are often
brought up on long fines from 4 to 30 fathoms, but
are sometimes found close below tide mark. There
is no reason to suppose that the males care for the
eggs, and the latter are so hardy that they do not
suffer from exposure to the air for hours. Judging
from the dates when newly hatched larvae have
been seen, incubation of the European species
occupies a month, perhaps longer in the case of
the eggs that are spawned earliest in the season,
at winter temperatures. The larvae are about
3.3 to 4.5 mm. long at hatching, with a small
rose-red yolk sac that contains a large oil globule
and that is inclosed in a net of blood vessels.
Most of the characters of the adult are apparent
at 11 to 12 mm. length, but the pectoral fins are
brilliantly pigmented with yellow and black
throughout the larval stage.47

General range. — Rocky shores along the North
American Coast from northeastern Newfoundland,
the northern part of the Gulf of St. Lawrence and
the Grand Banks to southern New England.48 It
is rare west and south of Cape Cod, but has been
taken at Woods Hole, on the coast of Connecticut,
and off New Jersey.49

Occurrence in the Gulf of Maine. — This sea snail
is generally distributed around the shore line of the
Gulf. Thus the Halcyon trawled it off Yarmouth,
Nova Scotia, in January 1921 ; it is rather common
(according to Huntsman) in the Bay of Fundy and
in Passamaquoddy Bay; and it has been definitely
reported at Grand Manan; Eastport; Seguin
Island; off Portland (where many have been col-
lected); off Cape Elizabeth; at Kittery; and at
various localities about Massachusetts Bay. It

«• McKenzie and Homans, Proc. Nova Scotian Inst. Scl., vol. 19, Pt. 3,

1938, p. 278.

« Mcintosh and Mastermann (Life histories of the British Marine Food-
fishes, 1897 p. 191, pi. 2. fig. 9, 10) and Ehrenbaum (Nordisches Plankton,
vol. 1, 1905-1909, p. 109) give descriptions of the eggs and larvae of the Euro-
pean sea snail N. montagui (as Cydogaster montagui) from which the pre-
ceding is condensed.

«» This fish is so closely allied to the north European sea snail, N. montagui
(from which, however, it is quite distinct) that it passed under that name
prior to 1898.

" A specimen was taken by Albatross II. off Atl antic City, lat. 39°24' N.,
long. 74°05' \V\, in 11 fathoms, in April 1930.

466

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

has been taken on Georges Bank also, and on
Browns, living in scallop shells (p. 465).

Vladykov and McKenzie characterize it as "not
uncommon" all around Nova Scotia;60 it is classed
by Huntsman as characteristic of the icy cold
water on the banks in the southern part of the
Gulf of St. Lawrence; and it has been reported
from the estuary of the St. Lawrence River; from
the northeastern part of the Gulf of St. Lawrence;
off the south coast of Newfoundland; on the Grand
Banks, and as far north on the Atlantic coast as
the entrance to the Strait of Belle Isle.61

Striped sea snail Liparis liparis (Linnaeus) 1766

Sea snail
Jordan and Evermann, 1896-1900, p. 2116.

Figure 244. — Striped sea snail (Liparis liparis), side view
(above) and ventral view (below) of adult. After
Garman.

Description.' — This little fish resembles the sea
snail (p. 464) so closely, especially in its tadpole-like
form, in the presence of a sucking disk on its chest,
in which the rays of the ventrals (reduced to mere
knobs) serve as a central support, and in the
peculiar outline of the pectoral fins with secondary
frilled basal lobes, that it is difficult to distinguish
one from the other. The most obvious difference
between the two species is that there is no indenta-
tion between the spiny and the soft parts of the
dorsal fin in the striped sea snail. Furthermore,
it usually has 33-35 rays in the soft portion of its
dorsal fin and 26-29 rays in its anal fin, as against
a maximum of 32 dorsal rays and of 27 anal rays

» Proc. Nova Scotian Inst. Sci., vol. 19, 1935, p. 99.

" From the trawllngs of the Newfoundland Fisheries Research Commission.

in the other sea snail (p. 404) . And the separation
between the dorsal and anal fins and its caudal fin
is not as definite in the striped sea snail as it is
in the preceding species; in fact it is sometimes
difficult to draw a sharp line between the fins.
A minor character, which gives the head a rather
different aspect, is that the dorsal profile is more
arched in the striped sea snail.

Color. — -Many color varieties of this fish have
been described and named. As a rule the ground
tint is of a shade of olive green, gray, or brown,
variously tinged with reddish, with yellowish, or
with lilac, and but little paler below than above.
Red ones with pale and dark stripes have been
seen among kelp in New England waters. And
they are dark and pale in endless variety in vary-
ing situations, some nearly plain, some definitely
striped with few or many narrow longitudinal
bands, others spotted. In fact no two are alike.
Usually the fins are darkly blotched or barred.

Size. — This fish grows to a length of 10 inches
in Arctic seas but very few of them are more than
5 inches long in temperate latitudes.

Habits. — All that is known of its habits in our
Gulf is that it fives on rocky or stony bottom,
usually among the stalks and roots of kelp to
which it sometimes clings fast, a habit which
European writers describe as common. In British
waters it is often to be found hiding in the tiny
pools of water that are left under pebbles by the
ebbing tide, and probably a search of the beaches
would reveal it in similar situations in the Gulf of
Maine also. Small ones often live inside the shells
of the giant scallop (Pecten magellanicus) , and it is
our impression (though not backed by any definite
evidence) that this is a more usual habit with the
striped sea snail than with the preceding one
(p. 465). At any rate, W. F. Clapp informs us
that it is the rule to find at least one or two striped
sea snails in a bushel or so of sea scallops, and
fishermen have told us that sea snails of one species
or the other (probably of both species) are found
in scallop shells on Georges Bank.

Small crustaceans, chiefly amphipods and
shrimps of various kinds, have been found in the
stomachs of striped sea snails on both sides of the
Atlantic; they also feed on small shellfish, and they
were described by Fabricius 62 as eating small fish
fry and algae.

" Fauna Qroenlandica, 1780, p. 137.

FISHES OF THE GULF OF MAINE

467

This is a winter-spring spawner; females are full
of roe at Woods Hole in December and January,
and the collection of the Museum of Comparative
Zoology contains a female distended with eggs that
was taken on April 1 many years ago. Larvae
only 5.5 mm. long, which we towed near the Isles
of Shoals on July 22 and in Massachusetts Bay on
August 31 in 1912, must have been hatched from
eggs spawned at least as late as May, if not in June.

The eggs,63 about 1.5 mm. (0.06-inch) in diame-
ter, sink and stick together in bunches, to hydroids,
seaweeds, or other objects, like those of Neoliparis
atlanticus, and it seems that incubation is about as
long as it is with the latter, i. e., at least a month.
The larvae are about 5.5 mm. long at hatching
and they live adrift until they are upward of 16
mm. long, when the sucking disk is well developed.

General range. — Arctic and North Temperate
Atlantic; north to the White Sea, Spitzbergen,
Greenland, Davis Strait, and northern Labrador,
and reported from the Kara Sea and from the
Arctic Ocean north of Siberia; south to northern
France and to Delaware Bay and Virginia.64

Occurrence in the Gulf of Maine. — The distribu-
tion of this sea snail parallels that of the preceding
species in our Gulf. Thus it has been dredged not
uncommonly in the Bay of Fundy region in from
5 to 100 fathoms and has been recorded from
Grand Manan: from Eastport, as well as from
other localities on the Maine coast; here and there
about Massachusetts Bay; and on Georges Bank;
also at Woods Hole.

In Nova Scotian waters it has been characterized
variously as "common" 66 and as "uncommon." u

It has been described as "common" in the
southern part of the Gulf of St. Lawrence;67 has
been reported in the estuary of the St. Lawrence
River; at Anticosti; and in the northeastern part
of the Gulf; also off the south coast of Newfound-
land; in Conception Bay; off the eastern end
of the Strait of Belle Isle; off the southeastern
Labrador coast,68 and from Fort Chimo, on Ungava
Bay, in northern Labrador,69 as well as from West
Greenland.

It is of no commercial importance.

THE SEA ROBINS OR GURNARDS AND THE ARMORED SEA ROBINS

FAMILIES TRIGLIDAE AND PERISTEDIIDAE

The sea robins and their European relatives, the
gurnards, suggest sculpins in their broad heads,
slender bodies, large fanlike pectoral fins, in
having two separate dorsal fins (a spiny and a
soft rayed), and in the location of their ventral
fins under the pectorals. But their entire heads
are armored with rough bony and spiny plates.
The Gulf of Maine is the northern limit for the
family on the Atlantic Coast of America.

The armored sea robins are close relatives to
the sea robins but they differ from them in four
very noticeable ways: (1) the entire body is en-
closed in an armor of bony plates, each plate with
a spine; (2) it is only the two lower rays of the
pectorals that form separate feelers; (3) each side
of the front of the skull projects forward as a
long flat process, so that the snout appears to be
double; (4) they have 2 long barbels on the chin.
They live on bottom in fairly deep water, and
they are widespread in tropical to boreal seas.
One species is a member of the Gulf of Maine
fish fauna.

« The following lines are condensed from Ehrenbaum's (Nordlsches Plank-
ton, vol. 1, 1905-1909, p. 112) account ofitseggs and larvae In European waters.

KEY TO GULF OF MAINE SEA ROBINS
AND ARMORED SEA ROBINS

1. Front of snout only slightly concave as seen from

above; no barbels on chin 2

Front of snout so deeply concave that it seems to be
double when seen from above (fig. 247) ; two long
barbels on chin Armored Sea Robin p. 471

2. Pectoral fin with 2 broad dusky blotches; there is no

prominent longitudinal stripe on the side of the

body Common Sea Robin p. 467

Pectoral fin with only 1 broad dusky blotch; there is
a prominent longitudinal dark brown stripe on each
side of the body Striped Sea Robin p. 470

Common sea robin Prionotus carolinus
(Linnaeus) 1771 «°

Sea robin; Robin, Green-Eye
Jordan and Evermann, 1896-1900, p. 2156.

M This sea snail, formerly known only as far south as New York, has been
taken off Delaware Bay by Albatross II, and off Assateague, Virginia, by the
Grampus (Welsh, Copeia, No. 18, 1915, p. 2).

« Jones, Proc. Nova Scotian Inst. Sci., vol. 5, pt. 1, 1882, p. 89.

M Vladykov and McKenrie, Proc. Nova Scotian Inst. Sci., vol. 19, 1935,
p. 99.

» Cox, Contrib. Canadian Biol. (1918-1920) 1921, p. 112.

w From the cruises of the Newfoundland Research Commission.

" Packard Proc. Portland Soc. Nat. Hist., vol. 2, No. 13, 1909, p. 112.

"Jordan, Evermann, and Clark (Rept. U. S. Comm. Fish. (1928) Pt. 2,
1930, p. 407, place this species In the Genus Merultnus which was proposed
by Jordan and Evermann in 1898 as a subgenus.

468

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 245. — Common sea robin (Prionotus carolinus). Adult, New Jersey; from Goode, drawing by H. L. Todd.
A, egg; B, larva, just hatched, 2.8 mm.; C, young, 9 mm. A-C, after Kuntz and Radcliffe.

Description. — The large head, tapering body,
and fanlike pectoral fins of the sea robin somewhat
suggest a sculpin. But the robin is distinguished
from all the sculpins by the incasement of its
entire head in bony plates; by its smaller mouth;
by the flat depressed dorsal profile of its snout; by
its large ventral fins; and by the fact that the
three lower rays of each of its pectoral fins are
separate from the rest of the fin and modified into
three independent feelers with slightly dilated
tips, a very noticeable and distinctive feature.
Furthermore, the front margin of the upper jaw
is concave in outline when viewed from above,
not convex as it is in most other fishes, which
gives the nose of the robin a characteristic aspect.
The head plates are rough and there is one sharp
spine on each cheek at the angle of the gill cover;
two short spines over each eye pointing backward;
a spine on either side of the neck ; and one on each
shoulder above the base of the pectoral fin. The
spiny and soft-rayed portions of the dorsal fin are

separate, but they are so close together at their
bases that they are almost in contact. The
spiny dorsal has 10 spines, is rounded in outline,
and higher than the soft dorsal (13 rays); but the
soft dorsal is considerably longer than the spiny
dorsal.

The caudal fin is of moderate size, its margin
slightly concave. The anal fin (1 spine and 11
rays) is similar in outline to the soft dorsal, under
which it stands. The pectorals (their 3 lower rays
as just noted) are rounded in outline and are so
large that they overlap the anal and the second
(soft) dorsal when they are laid back. The ven-
tral fins (each of 1 stiff spine and 5 rays) stand
close behind the pectorals.

Color. — Usually the body is grayish or reddish
brown above, with about five dark saddlelike
blotches along the back, and is dirty white or
pale yellow below. The dorsal fins are grayish,
marked with pale spots and stripes, with a black
spot between the fourth and fifth spines. The

FISHES OF THE GULF OF MAINE

469

caudal fin is uniform grayish or brownish; the
anal plain brown; the ventrals plain yellow to
brown. The pectorals are yellow or orange,
strikingly marked with two broad dusky bars, one
of them crossing the middle of the fin, and the
other crossing its outer third. The pectoral
filaments are orange.

Size. — The maximum length is 15 to 16 inches,
but few of them grow to be more than a foot long.

Habits. — Sea robins, like the sculpins, tend to
keep to the bottom. But they swim actively,
usually with the pectorals closed against the body.
They are often hooked close to the surface; we
have caught them when trolling for mackerel, and
many anglers have told us of similar experiences.
When on the bottom they often lie with the fanlike
pectorals spread. If disturbed they bury them-
selves in the sand, all but the top of the head and
eyes, and they are said to employ the feeler-like
rays of the pectorals in stirring up the weeds
and sand to rout out the small animals upon
which they feed. They are usually found on
smooth hard grounds, less often on mud or about
rocks.

Along southern New England, where robins
are far more plentiful than they are farther north,
a large part of the population appear inshore in
May or June, to pass the summer there; some
close to tide line, but others remaining in depths of
5 to 30 or 40 fathoms, or deeper. Like many
warm-water fishes, they disappear from the coast
in October, to pass the cold season well out on the
shelf, as recently proved by catches made at 50 to
55 fathoms off southern Massachusetts by the
dragger Eugene H in late January,61 in 1950, also
at 21 to 93 fathoms off North Carolina, in that
same month and the next, by the Albatross III.62

The fact that the Albatross III trawled up to
83 sea robins per haul off New York and off south-
ern New England at 22-61 fathoms as late in the
season as mid-May of that same year suggests
that some of them may remain well offshore until
into the summer, if not all summer.

Notwithstanding this inshore and offshore mi-
gration, some at least of the sea robins experience
a temperature range of nearly 30° F. with the
change of the seasons, for those that come closest
inshore are in water as warm as 68°-70° at the end

of the summer, while some that were trawled
along the 30- to 40-fathom zone in May were in
water as cold as 40.2° to 41.4°.

The sea robin is a voracious fish, feeding indif-
ferently on shrimps, crabs of various kinds,
amphipods (crustaceans are its chief diet), squids,
bivalve mollusks, annelid worms, and on small
fish, such as herring, menhaden, and small winter
flounders. Seaweed has also been found in sea
robin stomachs. They bite greedily on any bait,
and are often taken with a spinner, or other
artificial lure.

About Woods Hole the common sea robin
spawns from June to September with July and
August as the peak of the season.63 But some may
spawn earlier, for we have examined females
taken at 50-55 fathoms off southern New Ens-
land at the end of January with eggs so large as
to suggest that they would be spawned by April
or May. Unlike the sculpin tribe, the robin
produces buoyant eggs, which are 0.94 to 1.15
mm. in diameter, slightly yellowish, with a vari-
able number (10 to 25) of oil globules of various
sizes, usually arranged in a more or less definite
ring. Incubation occupies about 60 hours at a
temperature of 72°, but any eggs that might be
spawned in the cooler water of our Gulf would be
slower in hatching. The newly hatched larvae
are 2.5 to 2.8 mm. long, with two transverse
yellow cross bands, one of these close behind the
pectoral fins, the other midway between vent and
tail. The yolk is absorbed, the mouth formed,
and the yellow markings no longer prominent in
5 days, at a length of 3 to 3.4 mm. The dorsal
and anal fin rays are visible and the lower pectoral
rays have separated from the remainder of the
fin at about 9 mm., and young fish of 25 to 30
mm. are darker, with transverse bands, and
already show most of the distinctive characters
of the adult.

General range. — Coastal waters of eastern North
America from the Bay of Fundy to South Carolina;
chiefly west and south from Cape Cod.

Occurrence in the Gulf of Maine. — Plentiful
though the sea robin is off southern New Eng-
land,64 only a few are taken north of Cape Cod.

«i We saw these catches which ranged from 0 up to 5,000 fish per haul, in
54 trawl hauls.
•i One to one hundred and thirty sea robins per haul.

— Kuntz and Radcliffe (Bull. U. S. Bur. Fish., vol. 35, 1918, p. 105-109)
give an account of its embryology and larval stages, subsequently confirmed
and supplemented by Welsh.

" A catch of 1,000, in a day, in one trap, is recorded for Vineyard Sound,
and of as many as 3,000 to 5,000, per trawl haul, at 50 to 55 fathoms off southern
Massachusetts in winter. See footnote 61, p. 469.

470

FISHERY BULLETIN OP THE FISH AND WILDLIFE SERVICE

Following the coast we find them reported at
Truro, Cape Cod; Prof. A. E. Gross has informed
us that he often saw as many as a dozen sea
robins taken in the trap at the entrance to Barn-
stable Harbor in a single tide in the early summer
of 1920; 96 one now in the Museum of Compara-
tive Zoology was trawled 12 to 15 miles off
Plymouth, at 30 fathoms, on November 20, 1943;
the sea robin has been reported off Lynn and
Salem; and in 1913 Welsh saw several in a trap
at Manchester, on the North Shore of Massa-
chusetts Bay on June 29. North of Cape Ann
it has been taken at Anisquam; at Newburyport
at the mouth of the Merrimac River, whence
one about 1 foot long was brought in to the
Museum of Comparative Zoology on August 14,
1949; also at the mouth of the Saco River. And
Dr. W. C. Kendall saw more than 25 taken
from the traps near Small Point, Casco Bay,
between July 4 and 14 in 1896.

The only records for it for the coast east of
Small Point are, however, for a single specimen
caught at Campobello Island in the mouth of the
Bay of Fundy in August 1911, and another in
August 1949,66 one taken in a weir in Passama-
quoddy Bay at St. Andrews, October 2, 1935,"
and of another taken in the Bay of Fundy, near
Minas Channel, during the late summer or early
autumn of 1951. M

Enough sea robins also range eastward across
the South Channel for trawlers to have picked up
a few (never more than a dozen or two on a trip)

« Briefly mentioned in The Auk, vol. 40, No. I, January 1923, p. 24.

on Georges Bank during the summer of 1913.
But it is probable that the deep channel between
Georges and Browns Banks form its easterly
limit, for sea robins are not known on Browns
Bank or off the west coast of Nova Scotia.

It is not hkely that the sea robin ever suc-
ceeds in reproducing itself in the Gulf, unless in
restricted localities such as Casco Bay, where
summer temperatures may be high enough.
We have never taken its rather characteristic
eggs in our tow nets anywhere in the Gulf, nor
have its young fry ever been reported there.
But when wandering fish do find their way
around Cape Cod from the south, they may re-
main there, wintering offshore in deeper water,
as they do farther south.

Importance. — The sea robin is edible, and its
near relatives, the gurnards, are table fish in
Europe, but it is too scarce in the Gulf of Maine
to be of any potential commercial importance
there. Off southern New England, where it is
abundant, it is a nuisance to anglers, taking bait
planned for other fishes.

Striped sea robin Prionotus evolans (Linnaeus)
1766

Jordan and Evermann, 1896-1900, p. 2167, as Prionotus
strigatus Cuvier and Valenciennes, 1829 6'

" This latter flsh was reported by Scattergood, Trefethen, and Coffin
(Copela, 1951, No. 4, p. 298).

*> McGoniglo and Smith, Proc. Nova Scotian Inst. Scl., vol. 19, 1936, p. 160.

« Reported to us by letter by Dr. Huntsman.

• Oinsberg (Texas Jour. Sci., vol. 2, No. 4, 1950, p. 619, 620- 522) has shown
that the P. strigatus of Cuvier and Valenciennes Is not separable from the
evola nsot Linnaeus.

Figure 246. — Striped sea robin (Prionotus evolans), Woods Hole, Mass. From Goode. Drawing by H. L. Todd.

FISHES OF THE GULF OF MAINE

471

Description. — The striped sea robin resembles
the common sea robin so closely that one might
easily be taken for the other. But its mouth is
wider and gapes back almost opposite the front
of the eye, with the maxillary bone more than
one-third as long as the head. Its head is flatter
(compare fig. 246 with fig. 245) ; its pectoral fins
are longer relatively (reaching back to the ninth
or tenth ray of the soft dorsal fin instead of only
to the fifth or sixth ray); its pectoral feelers are
more slender and tapering; its caudal fin is
square-ended instead of concave in rear outline;
and its reddish or olive-brown sides (the general
ground tint varies) are marked longitudinally
with a dusky or bronze-brown stripe below the
lateral line. The first dorsal fin shows the same
black or dusky blotch betweeen the fourth and
fifth spines, so characteristic of the common
robin. The pectorals are orange to brown with
pale edges, their centers washed with dusky, but
without the definite crossbars characteristic of the
common sea robin. The pectoral filaments are
pale brown or orange, marked with narrow
brown bars. (The common sea robin does not
show these bars.)

Size. — This is a larger fish than the common
sea robin, growing to a maximum length of about
18 inches.

General range.- — Shoal water along the Atlantic
coast of North America from South Carolina to
Cape Cod, reaching the Gulf of Maine as a stray
from the south.

Occurrence in the Gulf of Maine. — This southern
fish rounds Cape Cod so seldom that there are
only 6 definite records for it from the Gulf of
Maine: Monomoy; North Truro; Salem; Glouces-
ter; Monhegan Island, Maine (its most north-
erly outpost) where one was taken in an otter
trawl at 40 fathoms, November 19, 1933; and
the eastern part of Georges Bank, whence one
was brought in to the U. S. Fish Commission some-
time between 1877 and 1880. We have never
seen it north or east of the elbow of Cape Cod.

Armored sea robin Peristedion miniatum Goode
1880

Jordan and Evermann, 189&-1900, p. 2178.

Description. — The armored robin resembles the
sea robins in general body form, and in the ar-

Figure 247. — Armored sea robin {Peristedion miniatum),
side view (above) and top view of head (below). Con-
tinental slope off Martha's Vineyard. From Goode and
Bean. Drawings by H. L. Todd.

472

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

rangement of its fins, including the fact that the
two lower rays of each pectoral form separate
feelers. But its body is entirely clothed with
bony plates of considerable size. There are 4
rows of these on each side, from the vent rearward,
with an equal number of rows of thornlike spines,
the latter close set and directed rearward. The
plates on the abdomen have no spines. Thanks
to this armor, the trunk is very stiff. The lower
jaw bears a number of short fleshy barbels; there
is one long barbel with short side branches at each
corner of the mouth (the sea robins have no bar-
bels) ; and the front of its head is given so peculiar
an appearance by the two projections from the
skull (p. 467) that the armored robin could hardly
be mistaken for any other fish, except for one of
its own tribe.

Color. — Bright crimson, below and above.

Size. — Maximum recorded length between 13
and 14 inches (330-355 mm.).

Habits. — This is a ground fish, recorded from
depths ranging from 50 fathoms down to some-
where between 200 and 235 fathoms. And it

seems to be confined to the zone of warm water
along the outer part of the continental shelf and
upper part of the continental slope for the lowest
temperature in which it has been recorded is be-
tween 44° and 45°. n The stomachs of those we
have opened contained shrimps, stomatopods, and
other small crustaceans.

General range. — Outer part of the continental
shelf and upper part of the continental slope ; from
the southwestern face of Georges Bank to the
offing of Charleston, S. C. (lat. 32°24' N., long.
78°44' W.).71

Occurrence in the Gulf of Maine. — Trawlers tell
us they sometimes take these brilliant crimson
fishes on the southwestern part of Georges Bank.
And they must be rather common outside the 60-
fathom contour, for we saw 89 specimens trawled
there and south of Nantucket at depths of 66 to
more than 185 fathoms, by the Albatross III in May
1950. But it is probable that they are barred
from the more easterly parts of the bank and from
the inner parts of the Gulf of Maine by low
temperature.

THE FLYING GURNARDS. FAMILY DACTYLOPTERIDAE

Flying Gurnard Dactylopterus volitans (Linnaeus)
1758

Jordan and Evermann, 1896-1900, p. 2183, as Cephala-

canthus volitans (Linnaeus)

Description. — The flying gurnard (only known
representative of its family), built, in general, on
the "sea robin" plan, is remarkable for its enor-

mous pectorals. When it is about half grown or
older, these reach nearly to the base of the caudal
fin if laid back. When spread, they have the form
of enormous rounded fanlike wings. Other con-

« We have seen it trawled off southern New England In water as shoal as
50 fathoms, and as cold as 44.4°.

" For list of early localities, see Goode and Bean, Smithsonian Contrib.
Knowl., vol. 30, 1895, p. 471.

Figure 248. — Flying gurnard {Dactylopterus volitans), Key West, Florida.

H. L. Todd.

From Jordan and Evermann. Drawing by

FISHES OF THE GULF OF MAINE

473

spicuous respects in which the flying gurnard dif-
fers noticeably from its relatives, the sea robins,
are that the first five or six rays of each pectoral,
with their interconnecting membrane, form a
separate fin, having no connection with the re-
mainder of the pectoral except at the base; that
the few lowermost rays of each pectoral fin are not
separate, in the form of feelers, but are continuous
with the remainder of the fin; that the first two
spines of the dorsal fin are separate, that the bony
armor covering the front and top of the head
reaches rearward considerably beyond the origin
of the dorsal fin on either side to end in a stout
spine, that each gill cover 72 is extended rearward
as a stout spine about as far as the axil of the
pectoral fin, and that the scales are much larger
and each armed with a short stout spine.

Color. — This is a very brilliant fish, varying
widely in color; most of them are of some shade of
brownish to greenish olive above, with the lower
side paler, but marked irregularly with reddish
salmon or salmon yellow. The winglike pectorals
are variously marked with bright blue streaks near

their bases, with blue spots and bars toward their
tips. The caudal fin usually has about three
brownish-red cross bars.

Size. — To about 12 inches.

General range.- — Tropical to warm temperate
latitudes of both coasts of the Atlantic; south to
Brazil and north rather commonly to North Caro-
lina on the American coast; a few to New York
and the southern coast of Massachusetts in most
years (in autumn 73) ; recorded as a stray from
Massachusetts Bay. A dried and hardened speci-
men that was found on the shore near Country
Harbor, Nova Scotia, in September 1939, by
Stanley McKinley, among the kelp and eel grass
that had been washed ashore duriug the night, was
thought by him (no doubt correctly) to have been
carried north on the deck of some steamer from
the south.74

Occurrence in the Oulj of Maine.- — The only
report of this warm-water fish from north or east
of Cape Cod is of one said to have been taken in
Massachusetts Bay.76

THE CUNNER TRIBE, OR WRASSES. FAMILY LABRIDAE

Members of the cunner family have a single
long dorsal fin, its forward part spiny, its rear
part soft rayed, with no evident demarkation
between the two. The ventral fins are located
under the pectorals, and the caudal peduncle is
very deep. The structure of the dorsal fin is
sufficient of itself to distinguish them from all
Gulf of Maine fishes except the scup, sea bass,
rosefish, tilefish, or certain sculpins. And there
is no danger of confusing a cunner or tautog with
any of these, for their rounded tails and pectorals,
and their general body-forms separate them at a
glance from the thin-bodied, fork-tailed scup; their
small mouths and the relative sizes of their fins
are obvious distinctions between them and the sea
bass tribe; their smooth cheeks and broad caudal
fins separate them from the spiny-headed, narrow-
tailed rosefish or from any sculpin; and they do
not in the least resemble the tilefish with its broad
mouth, adipose "fin" on the nape of its neck, con-
cave tail fin, and pointed pectorals. Both the
roof of the mouth and the floor of the throat
(pharynx) is armed with a patch of conical or

" Actually the preopcrcular bone.
210941—53 31

knoblike teeth in the cunner tribe. It is with
these that they grind the hard-shelled mollusks
and crustaceans on which they feed.

KEY TO GULF OF MAINE CUNNERS

1. Gill covers scaly, snout somewhat pointed, dorsal
profile of head rather flat Cunner, p. 473

Gill covers largely naked, snout blunt, dorsal profile
of head high-arched Tautog, p. 478

Cunner Tautoyolabrus adspersus (Walbaum) 1792

Perch; Sea perch; Blue perch; Bergall; Chogset

Jordan and Evermann, 1896-1900, p. 1577.

Description. — The readiest field marks by which
the cunner may be distinguished from its close
relative, the tautog, are mentioned on page 479.
It is moderately deep in body, moderately flat-
tened sidewise, with a very deep caudal peduncle,

'3 The most recent record from Woods Hole, of which we have heard, Is of
two taken there on November 24, 1948, from the deck of Albatross 111 while
she was moored at the dock (Arnold, Copela, 1949, p. 300).

'< McKenzie, Proc. Nova Scotian Inst. Scl., vol. 20, 1940, p. 44.

" This specimen is now in the Museum of Comparative Zoology, to which
It was transferred from the Boston Society of Natural History. There Is no
clue to its origin, except that It was taken many years ago.

474

FISHERY BULLETIN OF THE FISH AND "WILDLIFE SERVICE

Figure 249. — Cunner (Taulogolabrus adspersus). A, adult, Woods Hole, Mass.; from Goode, drawing by H. L. Todd.
B, egg; C, larva, newly hatched, 2.2 mm.; D, larva, 4.2 mm.; E, young, 8 mm. B-E, after Kuntz and Radcliffe.

flat-topped head (in the tautog the dorsal profile
is high arched), small mouth at the tip of the
snout, rather pointed nose, and protractile pre-
maxillary bones. Its lips, too, are thinner than
those of the tautog. It has several rows of conical
teeth of various sizes in each jaw, the outer ones
very stout. Its body and gill covers are covered
with large scales (in the tautog there is a naked
area in front of each gill opening), and its skin is
so tough that the fish must be skinned before
marketing. Its dorsal fin (about 18 spines and
9 or 10 soft rays) originates over the upper corner
of the gill cover, i. e., a little in front of the pecto-
ral fins, and runs back to the caudal peduncle.
The first 4 or 5 rays of the dorsal fin are graduated,
the others are of about equal lengths. The soft
part is only a little more than one-third as long as

the spiny part, and is rounded in outline. The
rear margin of the caudal fin is slightly convex
with rounded corners. The anal fin (3 stout
spines and about 9 rays) originates under or
behind the middle of the dorsal and corresponds
to the soft part of the latter in outline. The
ventrals stand under or a little behind the pecto-
rals; both the ventrals and the pectorals are of
moderate size, and the pectorals are rounded.

Color. — To describe the color of the cunner is to
list all the colors of the bottoms on which it lives,
for it is one of the most variable of fishes. As a
rule the upper parts range from reddish brown
(darker or paler) of a bluish cast to blue with
brownish tinge, variously mottled with blue,
brown, and reddish. Some fish, however, are
uniform brown, while fish caught over mud bot-

FISHES OF THE GULF OF MAINE

475

torn are often very deep sepia. In some situa-
tions they may be dull olive green mingled with
blue, brown, or rust color. Some cunners are
slaty, but reddish or rust tones are apt to prevail
when they are living among red seaweeds about
rocks. Cunners caught in deep water are often
almost as red as the rosefish; on the other hand
we have seen very pale ones, more or less speckled
all over with blackish dots, over sandy bottom.
The belly is invariably of a bluish cast, more or
less vivid, sometimes whitish, sometimes dusky,
sometimes little paler than the sides. Some cun-
ners have the lips and lining of the mouth bright
yellow. Young fry are more or less dark-barred
and blotched.

Size. — In the Gulf of Maine adult cunners
measure about 6 to 10 inches in length and weigh
less than half a pound, and one a foot long is very
large. But a few are caught up to 15 inches long,
and as heavy as 2% pounds.

Habits. — The cunner is chiefly a coastwise fish.
In our northern waters they are the most plenti-
ful from just below tide mark downward. They
swarm among eel grass (Zostera) and about the
piling of wharves and under floats in harbors.
They also run up into the deeper salt creeks, small
fish farther than larger ones, though we have
never heard of one in water that is appreciably
brackish; and young cunners are often found
among eel grass and in rock pools. Southward,
however, from New York or thereabouts, most of
them keep to water at least 15 to 20 feet deep,
hence somewhat farther out, depending on the
topography of the coast line and of the bottom.

At the other extreme, they are common enough
at 10 to 15 fathoms in the inner parts of Massa-
chusetts Bay, and not rare as deep as 25 to 35
fathoms on the offshore ledges and banks, and
we have taken them as deep as 70 fathoms on
Georges Bank. But the great majority live
within 5 or 6 miles of the shore. And while there
are some on the offshore grounds, such as Stell-
wagen Bank, Jeffreys and Cashes Ledges, and even
on Georges and Browns Banks where the otter
trawls frequently pick up a few, we have never
heard of a large catch of them made far out at
sea, whether along southern New England or to
the northward. Most of the cunners that are
caught the deepest and the farthest offshore are
large ones that have probably strayed thither,
and finding good feeding, have remained.

As far as wc know adult cunners never depart
far from the bottom, or from the rocks about
which they make their homes, nor do they school.
Many, it is true, may live together, but they act
quite independently of one another, simply
congregating because the surroundings are attrac-
tive. Cunners, like other rockfish, spend much
of the time resting quietly or swimming slowly
among the bunches of Irish moss (Chondrus) and
fronds of kelp, or in the open spaces among
the eel grass (Zostera), wherever the latter has
reestabhshed itself, always on the lookout for food.

Cunners are year-round residents, broadly
speaking, wherever they are found. At the most,
they may descend into shghtly deeper water to
pass the coldest months,76 or they may desert the
shoalest parts of certain enclosed bays in midsum-
mer to escape the very high temperatures produced
there as the sun strikes the flats at low tide. They
have been described as hibernating in the mud
during the winter, or at least as lying among eel
grass or rocks in a more or less torpid state. But
we find no positive evidence of this; on the
contrary, practical fishermen, among them Capt.
L. B. Goodspeed, to whom we are indebted for
many notes, inform us that cunners are to be caught
in abundance on precisely the same spots in winter
as in summer. In fact a few are landed in Boston
during the cold months, and the only reason more
are not brought in then is that there is so little
demand for them.

It has long been known that the cunner is
vulnerable to very low temperatures. Hazards
of this sort are more frequent south of Cape Cod,
where the fish are more likely to be caught in very
shoal water in a sudden freeze, than in the Gulf of
Maine, where active mixing by the tide usually
prevents the water from chilling to the danger
point, except at the surface. However, this did
take place in Massachusetts Bay in the winter of
1835, when cunners came ashore in quantities
between Marblehead and Gloucester. And the
failure of the cunners to produce young within
the Bay of Fundy (p. 478) suggest that the lower
thermal limit to their successful reproduction is
about 55°-56°, though the young fry as well as
the adults are at home in temperatures close
to the freezing point of salt water. The upper

'• Ambrose (Proe. and Trans., Nova Scotian Inst. Nat. Sci., vol. 2, No. 2,
1870, p. 93) describes the cunners as moving out of Saint Margaret Bay.
Nova Scotia, in autumn, to return early in May.

476

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

thermal limit, for the well being of the cunner, is
something like 70°-72°, to judge from the distri-
bution of the species.

Cunners are omnivorous. As a rule they find
their livelihood browsing among seaweeds, stones,
or dock piles, biting off barnacles and small blue
mussels, with the fragments of which they are
often packed full. They devour enormous num-
bers of amphipods, shrimps, young lobsters, small
crabs, and other small crustaceans of all kinds;
also univalve mollusks and the smaller bivalves,
hydroids, and annelid worms. They sometimes
eat small sea urchins, bryozoans, and ascidians,
and they occasionally capture small fish such as
silversides, sticklebacks, pipefish, mummichogs,
and the fry of larger species. Finally, eel grass is
often found in cunner stomachs besides the animal
food. Small cunner fry taken at Woods Hole
were found by Dr. Linton to have fed chiefly on
minute Crustacea such as copepods, amphipods,
and isopods.

The cunner is a busjr scavenger in harbors, con-
gregating about any animal refuse, to feed on the
latter as well as on the amphipods and other
crustaceans attracted by the same morsels. They
are also said to eat fish eggs, and no doubt feed to
some extent on herring spawn. Our own belief
is that cunners are always hungry, no matter
what the stage of the tide.

The cunner spawns chiefly from late spring
through early summer. The eggs are buoyant,
transparent, 0.75 to 0.85 mm. in diameter, and
they do not have an oil globule. Incubation
occupies about 40 hours at temperatures of 70° to
72°, but it is probable that about 3 days are
required for hatching in the cooler waters of the
Gulf of Maine (55° to 65°). At hatching the
larvae are about 2 to 2.2 mm. long, and at 15 mm.
the young cunner is of practically adult form.
On newly hatched larvae the pigment cells are
scattered uniformly over head and trunk, but by
the 3-mm. stage they have gathered into a pair of
black spots, dorsal and ventral, about halfway
between the vent and the base of the caudal rays,
which are characteristic of the species. And these
spots persist to about the 10- to 20-mm. stage. By
the time the fry have grown to about 25 mm. they
are as variable in color as their parents (it is on
record that Louis Agassiz had 60 colored sketches
of small cunners 3 to 4 inches long, of different

hues, prepared at Nahant during a single sum-
mer).76

Fry of 1 to 1.2 inches have often been taken in
August, and young fish up to 2 inches long in
September in southern New England waters.
Hence we may assume that Gulf of Maine cunners
(probably hatched somewhat later) may average
about 2 to 2% inches by their first autumn, and
2% to 2% inches by the following June when they
are one year old, which Johansen 77 found true
also of the earliest hatched fry in the southern
side of the Gulf of St. Lawrence. The subsequent
rate of growth has not been studied for the cunners
of our Gulf. But Johansen's 78 age determina-
tions for cunners of the Gulf of St. Lawrence make
it likely that Gulf of Maine cunners 3 to 4 inches
long are 2 years old; those of 4 to 5 inches 2 or 3
years old ; those of 5 to 6 inches 3 years old ; those
of 6 to 7 inches 3 or 4 years old; those of 7 to 8
inches 4 or 5 years old; those of 8 to 9 inches 5 or
6 years old; those of 9 to 10 inches about 6 years
old; and those of 10 to 11 inches 6 or 7 years old.
But the relationship is complicated by the fact
that female cunners run larger than males, so that
males may be a year older than females of the
same size.

Most of the cunners mature in their third
summer (i. e., when 2 full years old) when 2% to
3% inches long.

General range. — Atlantic coast of North Amer-
ica and the offshore banks, from Conception Bay,
east coast of Newfoundland, and the western and
southern parts of the Gulf of St. Lawrence,79
southward in abundance to New Jersey, and
occasionallv as far as the mouth of Chesapeake
Bay.

Occurrence in the Gulf of Maine. — The cunner is
one of our most familiar fish, to be found all
around the shore line of the Gulf. The Massa-
chusetts Bay region is perhaps their chief center of
abundance, and they are so numerous there in

r* The embryology and larval development and fry of the cunner have
been described by Agassiz (Proc. Amer. Acad. Arts, Sei., N. Ser., vol. 9,
1882, p. 290, pis. 13 to 15); Agassiz and Whitman (Mem. Mas. Comp. Zool.,
vol. 14, No. 1, Pt. 1, 1885, p. 18, pis. 7-19, and Mem. Mus. Comp. Zool., vol.
40, No. 9, 1915, pis. 32-39); Kuntz and Radcliffe (Bull. U. S. Bur. Fish., vol.
35, 1918, p. 99, flgs. 18-29); and more recently by Johansen (Contr. Canad.
Biol., N. Ser., vol. 2, No. 17, 1925, pp. 44(M50).

" Contrib. Canadian Biol., N. Ser., vol. 2, No. 17, 192B, p. 451.

'• Johansen (Contrib. Canadian Biol., N. Ser., vol. 2, No. 17, 1925, pp.
451-455) worked out the age-length relationship for a large series of Gulf of
St. L3WTence cunners by a study of their scales and otoliths.

'• See Johansen, Contrib. Canadian Biol., Ser. 2, vol. 2, No. 17, 1925,
pp. 5-6 [427-428]), for the distribution of the cunner in Canadian waters.

FISHES OF THE GULF OF MAINE

477

good years, along the rocky shores and around and
over ledges, that no amount of fishing seems to
have any effect on their numbers. Generally
speaking, they are less numerous east of Casco
Bay, and our experience has been that they are
progressively less and less so eastward along the
shore from Penobscot Bay toward the Bay of
Fundy, but average larger. On the outer coast of
Mount Desert, for example, it is unusual to catch
one in the enclosed harbors (precisely the localities
they frequent farther west and south) , and most of
those caught outside are very large. Thus we
took many of 12 to 13 inches, averaging about \lA
pounds, near Baker's Island, off Northeast Harbor,
in August 1922, and no small ones. But young
fish in plenty, as well as adults, have been reported
from Bluehill Bay, nearby,80 where the water is
warmer in summer.

Cunners are also taken, here and there, along the
coast, eastward to the Grand Manan Channel,
sometimes in numbers as in 1928, when so many
were caught "about the rocks and in the coves to
the south of West Quoddy," that they were
reported in the press.81 But they are so scarce
ordinarily around Grand Manan and within
Passamaquoddy Bay that only half a dozen large
specimens had been taken there from the founding
of the Biological Station at St. Andrews in 1906
down to the early 1920's.82 And while the cunner
is reported from Black River east of St. John,
New Brunswick, it seems to be unknown farther in
along the New Brunswick shore of the Bay of
Fundy or in Chignecto Bay and Minas Basin at
the head. But Annapolis Basin on the Nova
Scotian side of the bay, harbors a few, while cun-
ners of all sizes are so numerous in St. Mary Bay
that this must be an important centre of reproduc-
tion and the source of the few large (i.e., old) ones
that are caught farther up the Bay of Fundy.
And they are reported along the western shore of
Nova Scotia, as at Pubnico for example.

There are large cunners in small numbers on the
offshore fishing grounds in our Gulf also, Stell-
wagen at the mouth of Massachusetts Bay,
Cashes Ledge, and Georges and Browns Banks,
as mentioned above (p. 475) in depths down to 50
fathoms or so. But it is not likely that they ever
descend into the deep basins of the Gulf. Cer-

*> Reported to us by Rear Adm. S. E. Morrison, U. S. Navy.

'■ Boston Transcript for August 29, 1928.

" Johansen, Contrib. Canadian Biol. N. Ser., vol. 2, No. 17, 1925, p. 5 [427].

tainly our experimental trawlings have not yielded
any there, 42 fathoms being the greatest depth at
which we have known of a cunner taken anywhere
in the inner parts of the Gulf.83

Extending our survey farther east and north,
we find cunners reported as numerous all along the
outer coast of Nova Scotia, including the many
bays and inlets, also in the southern side of the
Gulf of St. Lawrence from Cape Breton to the
Gaspe Peninsula, including the shallow bays of
Prince Edward Island and the shores of the
Magdalen Islands, also up the west coast of New-
foundland as far as Bay of Islands. And they are
to be expected at the heads of the bays along the
south coast of Newfoundland for they have been
taken in Conception Bay on the east coast. But
this last is their most northerly known outpost on
the Atlantic coast, and they have never been
reported either from the estuary of the St. Law-
rence or anywhere along the north shore of the
Gulf of St. Lawrence.84

Cunners near Newport, Rhode Island, com-
mence spawning by mid-May and June sees the
chief production of eggs there and near Woods
Hole, where most of the fish are spent after the
first days of July, though eggs have been taken in
abundance there until July 15, a few as late as
August 15. 85 Probably spawning does not com-
mence until June in the colder waters of our
Gulf, but continues there through the later
summer, for our towings have yielded many eggs,
apparently of the cunner, in July and August.
And the chief spawning season is about the same
as this in the southern side of the Gulf of St.
Lawrence, according to Johansen 8S and to Reid.87
Cunner eggs have been taken at our tow net
stations along outer Cape Cod; near Race Point
at the tip of the Cape; in Massachusetts Bay
(where we have often towed them in great numbers
in the tideways between the offlying ledges) ; and
at the mouth of Penobscot Bay, as well as in
sundry harbors. Blue Hill Bay inland from
Mount Desert may be a breeding center, for small
fry are reported there.88 And eggs taken off

81 One was trawled at this depth at the mouth of Massachusetts Bay (lat.
42°28' N., long. 70°13' W.) by the Albatross II, July 28, 1931.

>< See Johansen, Contrib. Canadian Biol., N. Ser. 2, vol. 2, No. 17, 1925, pp.
5-6 [427-428] for an account of the status of the cunner in the Gulf of St.
Lawrence, and around Newfoundland.

« Agassiz and Whitman, Mem. Mus. Comp. Zool., vol. 14, No. 1, 1885,
p. 18, Kuntz and Radcllffe, Bull. U. S. Bur. Fish., vol. 35, 1918, p. 99.

» Contrib. Canadian Biol., N. Ser., vol. 2, No. 17, 1925, p. 17 [439].

» Contrib. Canadian Biol, and Fish., N. Ser., vol. 4, No. 27, 1929.

» ByRearAdmiral S.E.Morrlson.U. S.N.

478

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Libbey Island prove that dinners spawn in
diminishing numbers eastward along the Maine
coast nearly to the mouth of the Bay of Fundy.
It is doubtful, however, whether eggs produced
along the coast east of Mount Desert yield more
than a very small proportion of fry, nor do cunners
breed successfully in the cold water of the Bay of
Fundy, where no small ones are ever seen. How-
ever, the Bay is simply a gap in the breeding
range, for St. Mary Bay is a productive nursery.
Both eggs and larvae were taken at various
lo«alities along the outer coast of Nova Scotia by
the Canadian Fisheries Expedition during the
summer; and the shoal inshore waters in the
southern side of the Gulf of St. LawreDce are a
productive spawning area.89

Larval cunners and small specimens generally,
like their eggs, are so closely confined to the coast
line that it is impossible to represent the localities
where we have taken them on a general chart of
the Gulf; in fact, all our catches of 100 or more
have been made either in harbors or at most not
a couple of miles from land.90 There may be some
successful reproduction on Cashes and Jeffreys
Ledges. But we have found no evidence, whether
of eggs or of young fry, that the few large curmers
that wander offshore to Georges Bank produce
any young there.

Variations in abundance. — No evidence is avail-
able as to how much the cunners may vary in
abundance from year to year, along the coasts of
our Gulf as a whole. But they may do so widely
at a given locality. Thus we found very few of
them in 1950 along the Cohasset shore, on the
southern side of Massachusetts Bay, where they
are plentiful ordinarily. And they were so scarce
there during the summer of 1951, that persons
raking Irish Moss (Chondrus) reported seeing
hardly a cunner around the rocks where many are
to be seen in most summers, and another acquaint-
ance who usually baits a lobster pot or two with
cunners taken in a cunner trap caught only one
occasionally in that way.

Importance. — The cunner was a favorite pan
fish once. During the 1870's the annual catch of
the small boats fishing out of Boston was esti-
mated as not much short of 300,000 pounds, while

» See Jobansan (Contrib. Canadian Biol., New Ser., vol. 2, No. 17, 1925,
p. 18 |440]; also Reid, Contrib. Canadian Biol, and Fish. N. Ser., vol. 4,
No. 27, 1929.

» The precise records have been published elsewhere (Bull. Mus. Comp.
Zool., vol. 58. 1914, p. 108, and vol. 61, 1917, p. 271).

the fact that 104,100 pounds of cunners were
reported for Maine in 1889, 148,300 pounds in
1898, and 281,500 pounds in 1905, shows that the
annual harvest was still considerable to that time.
But the reported catch had fallen to 30,695
pounds for Maine by 1919, and to about 10,000
pounds for the entire coast line of Massachusetts,
south as well as north of Cape Cod. And Maine
reported only 10,000 pounds for 1928 and 1,735
pounds for 1929, while the only cunners reported
for Massachusetts were 30 pounds and 45 pounds
for those 2 years, respectively. From that time
down to 1947, commercial catches of cunners
have been reported for Maine in only 3 years out
of the 14.91

The landings reported for Massachusetts during
this period suggests ups and downs so erratic
and so extreme 92 that we hesitate to place any
dependence upon them further than that landings
ranging from 3,100 pounds to 18,700 pounds
(average 7,450 pounds) for the years 1944-1947
show that a small demand continues for cunners.
And we can witness that sizeable ones are very
good pan fish.

Although not regarded as a game fish, the cunner
affords amusement to thousands of vacationists
near our seaside resorts. And the number caught,
of which no record is kept, is so considerable that
this must be classed as a useful little fish from
the recreational standpoint.

Probably more cunners are caught on bits of
clam than on any other bait. But they will take
snails broken from their shells, bits of crab,
lobster, or pieces of sea worms (Nereis) almost
as freely. And we have even caught a few while
trolling near rocks, for mackerel, with a small
spinner tipped with a bit of white fish skin. The
little ones are a great nuisance, often stealing
the bait as fast as it is offered, and because it is
a small-mouthed fish, very small hooks are best.

Tautog Tautoga onitis (Linnaeus) 1758
Blackfish; White chin
Jordan and Evermann, 1896-1900, p. 1578.

•' One hundred and seventy five pounds for 1933, 200 pounds for 1935, 45,300
pounds for 1938. an amount so large that we question its accuracy, especially
sinco the entire catch was reported as made on "lines, trawl." No catch
statistics are available for 1934, 1936, 1941, or 1942.

« Reported catches for Massachusetts jumped from 45 pounds for 1929 to
349,251 pouhds for 1931, dropped to 0 for 1932. 152 pounds for 1933 and 0 again
for 1935; rose to 27,800 pounds for 1937; were 0 again in 1933; but 53,500 pounds
in 1940.

FISHES OF THE GTTLF OF MAINE

479

Figure 250.— Tautog. Tautoga onitis. A, adult, Woods Hole, Mass.; from Goode, drawing by H. L. Todd. B, egg;
C, larva, one day old, 2.9 mm.; D, larva, 5 mm.; E, young fry, 10 mm. B-E, after Kuntz and Radcliffe.

Description. — The tautog suggests an over-
grown cunner, but it is a heavier, stouter fish
(about three times as long as deep, not counting
the caudal fin) with caudal peduncle so broad and
caudal fin so little wider than the peduncle that
it is hard to hold a heavy one by the tail. The
most obvious differences between the two fish are
that the dorsal profile of the head of the tautog
is high-arched, its nose is very blunt, and its lips
are much thicker, giving it a facial aspect quite
different from that of a cunner. A more precise if
less obvious character is that the cheek region
close in front of the gill opening (scaly in the
cunner) is naked in the tautog and velvety to the
touch. The fins of the tautog practically repro-
duce those of the cunner in relative size and

location. The dorsal fin (16 to 17 spines and 10
soft rays) originates over the upper corner of the
gill openings and runs back the whole length of
the trunk; the anal (3 stout spines and 7 or 8
soft rays) corresponds in outline to the soft portion
of the dorsal, under which it stands. The caudal
fin is slightly rounded at the corners, the pectorals
are large and rounded, and each of the ventrals
has one stout spine. The jaw teeth of the tautog
(in two series) are stout, conical, with the two
or three in the front of each jaw larger than the
others. The tautog has, besides, two groups of
flat, rounded, crushing teeth in the rear of the
mouth, as the cunner has also.

Color. — The tautog is a rather dark fish, gener-
ally mouse color, chocolate gray, deep dusky

480

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

green, or dull blackish, with the sides irregularly
mottled or blotched with darker. These mottlings
are more evident in the young than in adults and
usually they are grouped as three pairs of more or
less continuous bars. Large fish are often almost
plain blackish. The belly is only slightly paler
than the sides, but the chin usually is white on
large ones, a very conspicuous character. Tautogs,
like cunners, vary greatly in color on different
bottoms, and also in their markings.

Size. — Maximum length about 3 feet. The 22 }i
pounder, 36^ inches long, mentioned by Goode93
as caught off New York in 1876 and preserved in
the United States National Museum, still remains
the heaviest fish recorded definitely. Fish of
more than 14 pounds are very rare, with 12-
pounders unusual. Tautog average about 2 to 4
pounds as they come to market.

Habits. — The tautog is even more strictly a
coastwise fish than the cunner. Northward from
Cape Cod it is unusual to catch one more than 3
or 4 miles from the land, or deeper than, say 30-60
feet; we have never heard of one caught on a long
line set for cod or haddock, and they are unknown
on the offshore fishing banks. But they range
farther out and deeper to the southward, being one
of the commoner fishes caught in 10-13 fathoms
on the Cholera Bank, 10-12 sea miles offshore from
Long Island, and on Seventeen Fathom Bank, 8
miles off northern New Jersey. At the other ex-
treme, they follow the flood tide up above low-
water level around ledges, to prey on the abundant
supply of blue mussels along the intertidal zone,
dropping back into deeper water during the ebb.
We have helped to seine many small ones close
along the shore in only a few feet of water, at
Provincetown as well as southward, and it is not
unusual for tautog to run up into brackish water,
but we have never heard of them entering fresh
water.

Their favorite haunts are along steep, rocky
shores; around breakwaters, offlying ledges and
submerged wrecks; around the piers and docks;
over boulder strewn bottoms; and on mussel beds.
In some places, however, good numbers are caught
on smooth bottom, far from any rocks (the eastern
side of Cape Cod Bay is an example, see p. 482).
And young fry, 2 to 4 inches long, are often seined
on sandy beaches.94

When tautog are not feeding they are likely to
gather in some hole or cleft among the rocks, where
they lie inert, on their sides, often several crowded
together, until the rising tide stirs them to activity
again.95 And they are extremely local fish, per-
haps more so than any other Gulf of Maine species
that is interesting either to the angler or to the
commercial fisherman.

While tautog are seldom seen before well into
April in any part of their geographic range, or
after November, they do not carry out any exten-
sive migrations with the seasons. At most, those
that find themselves in shoal water in autumn may
drop off into slightly deeper water, to spend the
cold season lying among eelgrass (Zostera), where
this has reestablished itself; in crevices among
rocks; or (in the case of the young ones) in empty
oyster and clam shells. They move and feed little
then, though they have been caught in lobster pots
there and on hook and line off Rhode Island.96

Tautog, like cunners (p. 475), are sometimes
chilled and killed if they are caught in shoal water
by a sudden cold snap, as happened along Rhode
Island and southern Massachusetts in 1841, 1857,
1875, 1901, and no doubt on many other occasions
that have not found their way into print or into
the records of the Bureau of Fisheries.

Food. — Tautog feed on invertebrates, chiefly on
mollusks (both univalves and bivalves), especially
on mussels which are the chief diet of the tautog
living about ledges, and on barnacles that they
pick off the rocks. Crabs and hermit crabs are
favorite morsels. They also eat sand dollars, scal-
lops, amphipods, shrimps, isopods, and lobsters,
swallowing the smaller ones whole, but cracking
the larger with their crushing teeth (p. 479). A
tautog of about 2 pounds that we once caught off
Cohasset, Mass., had made a meal of gammarid
amphipods (sand fleas) gleaned from among the
rockweed with which the ledge was clothed, though
cunners caught at the same time and place were
full of barnacles. We think it likely that tautog
living in shallow bays (Duxbury, for example) prey

•> American Fishes. 1888, p. 292.

M We have seined tautog fry in such situations in localities as far apart as
Provincetown Harbor; Woods Hole, Cape Poge Bay. Marthas Vineyard,
and Cape Charles Beach, Va. And good numbers of larger tautog have been
reported as caught occasionally in nets in the vicinity of Provincetown; 8,700
pounds for example in 1898, and 5,800 pounds in 1899.

« We have often observed this habit of theirs in the large live tank at the
Woods Hole Oceanographic Institution.

•■ Tautog have been described as burying in the mud, but we cannot vouch
for this. And we put no credence whatever in the old myth that the vent
of the tautog closes over in winter.

FISHES OF THE GULF OF MAINE

481

to a considerable extent on sea worms (Nereis);
certainly they take these freely as bait.

Breeding habits. — About Woods Hole the tautog
spawn chiefly in June, and the season for such of
them as breed north of Cape Cod is probably
early and midsummer. The eggs are buoyant,
without oil globule and resemble those of the
cunner, except that they are a little larger (0.9 to
1 mm. in diameter). At a temperature of 68° to
72° incubation occupies 42 to 45 hours, and prob-
ably 10 to 12 hours longer in the cooler water of
Massachusetts Bay. The larvae 97 are about 2.2
mm. long at hatching. When 4 days old (tem-
perature of 68°-72°) they have grown to 3.3 mm.,
the 3rolk has been absorbed, and the mouth is
fully formed. Larvae of 5 mm. show the first
traces of the caudal fin rays; the dorsal and anal
fins are differentiated at 10 mm. and by the time
the little fish are about 30 mm. long they show
the fins, form, deep caudal peduncle, and blunt
nose of the adult tautog. The larvae and youngest
fry of the tautog and of the cunner resemble each
other closely in general form, but the arrangement
of the pigment offers a ready means of identifica-
tion at all but the very earliest stages, for the
black pigment cells remain more or less uniformly
scattered over the whole trunk in the tautog,
whereas they soon cluster in two definite patches
in the cunner as is described elsewhere (p. 476).

Probably Tracy 98 is correct in assuming that the
young tautogs of 3 to 8 inches, which may be seined
in abundance along the shores of southern New
England in summer, are 1 year old. Nothing
definite is known of the rate of growth of older
tautog, nor at what age they mature. But we
suspect that large ones of 8 pounds and more may
be 8 to 10 years old.

General range. — Atlantic coast of North Amer-
ica from the outer coast of Nova Scotia to South
Carolina, chiefly south of Cape Ann; most abun-
dant between Cape Cod and the Delaware Capes,
and restricted to the immediate vicinity of the
coast.

Occurrence in the Gulf of Maine. — The center of
abundance of the tautog lies to the south of Cape
Cod. Most of the authors, in fact, who have
written of it have accepted Mitchill's " statement

that it was not native north of Cape Cod and was
introduced there shortly prior to 1814, there being
no definite record of them in the Gulf of Maine
prior to that date. But it seems far more likely
that the anonymous writer who stated in the
Gloucester Telegraph of May 5, 1860, that tautog
had been plentiful there many years before, and
had merely reappeared after a period of scarcity,
was correct; also that this reappearance would
have taken place in any event, even if none had
been liberated north of Cape Cod.

Apart from Mitchill's statement that by 1814
the Boston market had a full supply (which may
have come from south and not north of Cape
Cod), the first positive record of any in Massa-
chusetts Bay is of several that were caught along
the Cohasset rocks in 1824, ' which the local fisher-
men said was a species new to them. Tautog,
however, were being caught in numbers in the
inner parts of Massachusetts Bay (e. g., Lynn,
Nahant, Boston Harbor) by 1839; they were more
abundant then around Manomet Headland in
Plymouth; and they already supported a con-
siderable hook-and-line fishery at Wellfleet. A
few years later their presence was established for
the coast of Maine, and in 1851 tautog were re-
ported as common (according to Perley) in St.
John Harbor, New Brunswick, though these Bay
of Fundy fish were introduced (not native). In
1876 the weirs north of Cape Cod took 2,274
pounds of tautog, and in 1879 Goode and Bean
described them as abundant in many localities
about Cape Ann.

At present (or within the last few years, for this
fish fluctuates in abundance from year to year),
the regular range of the tautog includes the whole
coast line from Cape Cod around to Cape Ann, in
suitable localities.

Tautog are less regular northward from Cape
Ann, less abundant, and more local. But there
are some tautog grounds about the Isles of Shoals,
off Cape Porpoise, and about Casco Bay, where
Kendall wrote of them in 1931 as having been
"locally numerous" for some time previous.2 We
have also heard of tautog along the ledges near
Boothbay Harbor and in Penobscot Bay. East
of the latter tautog certainly are not common.

« Kuntz and Radclifle (Bull. U. S. Bur. Fish., vol.35, 1918, p. 92) describe
the eggs, larvae, and fry.
•• 40th Ann. Kept. Inland. Fish. Rhode Island, 1910, p. 137.
» Trans. Lit. Philos. Soc., New York, vol. 1. 1815, p. 400.

210941—53 82

' Goode, Fish. Ind., U. S., Sect. 1, 1884 p. 269.

•According to Kendall (Bull. 58, Boston Soc. Nat. Hist., 1931, p. 10-11)
the green crabs (Carcliinides matnas) found In Casco Bay were not native
there but had been Introduced as tautog bait.

482

FISHERY BULLETIN OF THE FISH AND "WILDLIFE SERVICE

And it is so scarce a fish in the Passamaquoddy
region (it has long since vanished from St. John
Harbor) that three specimens, only, are known to
have been taken there within recent years.3

One has been taken near the head of the Bay of
Fundy on the Nova Scotian side (Scotts Bay,
Kings County) one on the Nova Scotian shore of
the open Gulf of Maine (Cranberry Head, Yar-
mouth County), and one on the outer coast of
Nova Scotia near Halifax (Petpeswick Harbor,
Halifax County), this last being the most northerly
record for the tautog.4

The more productive tautog grounds north of
the elbow of Cape Cod of which we chance to
know are the Cape Cod Bay shore southward from
Wellfleet; the Sandwich-Sagamore shore with the
jetties at the mouth of the Cape Cod Canal; the
bouldery ground around Manomet headland and
nearby; Gurnet Point at Duxbury; the ledges off
Scituate and Cohasset and especially those off
Swampscott; the Nahant, Marblehead, and Mag-
nolia Rocks; and here along the rocky shore from
Gloucester Harbor around Cape Ann. The Cape
Cod Bay grounds are exceptional, for the tautog
caught there are on smooth bottom, not among
ledges which are the usual haunts. We have also
known of good-sized tautog taken inside of Nauset
Inlet (where there are scattered boulders only),
one in a lobster pot during the summer of 1949.
And quite a number, large and small, are caught
within Duxbury Bay, especially around the pilings
of Powder Point Bridge.

Although tautog tend to gather in certain choice
spots, they move around enough so that some idea
of their relative importance along different parts
of the coast line can be determined from the
catches made in pound nets. Thus the average
yield per pound net or trap has run from twice to
20 times as great for Cape Cod Bay as for the
north shore of Massachusetts Bay in reasonably
good years 6 during the periods between 1890 and

• One in Passamaquoddy Bay In 1909 or 1910 (Reported by Huntsman
Contrlb. Canadian Biol. (1920-1921) 1922, p. 64); a second in a tidal tributary
of the St. Croix River in the summer of 1934, and another there in August
1935 (reported by McGonigle and Smith, Proc. Nova Scotian Inst. Sci., vol.
19, 1936, p. 160); all of these were taken in herring weirs.

♦ These Nova Scotian specimens are in the Provincial Museum at Halifax;
see Vladykov and McKenzie (Proc. Nova Scotian Inst. Sci., vol. 19, 1935,
p. 100), and Fowler (Proc. Acad. Nat. Science, Philadelphia, vol. 67, 1916,
p. 517) for further details.

' The reported catches for 1895, 1897-1900, and 1910 were 5 to 43 times as
great for Cape Cod Bay as for Essex County, made in 1.5 to 2.3 times as
many pound nets or sets of pound nets.

1921, when the catches for Massachusetts were
reported by towns, hence can be localized.6

The regional discrepancy has not always been
so wide in seasons when the Cape Cod Bay catch
has been smaller; in 1909, for instance, when the
total catch reported for Cape Cod Bay was only
635 pounds of tautog (with 27 pound nets in
operation) the average catch per pound net or
set of pound nets was nearly as great for the coast
from Boston Harbor to Gloucester (total catch
203 pounds with 12 nets or sets of nets in opera-
tion). But the pound nets take a few tautog in
Cape Cod Bay, even in years when they are so
scarce north of Boston that none at all have been
reported for Essex County, despite the fact that
the bottom seems more suited to tautog there
because rockier. The slightly lower temperature
along the north shore of Massachusetts Bay may
have been the contributing factor.

During the peak period 1895-1899, the chief
center of abundance for tautog for Cape Cod Bay
seems to have been along the Sagamore shore,
where the yearly catch, per pound net, then
averaged about 2% times as great as for the eastern
shore of the Bay,7 Brewster to Provincetown.
And catches of 18,100 pounds of tautog by 2 pound
nets at Sandwich in 1895 and 36,010 pounds of
tautog in 12 nets in Brewster in 1898 suggest
concentrations of tautog quite out of the ordinary.
But the best tautog fishing has been reported from
the Wellfleet region in recent years.

Catch statistics suggest, also, that not much
interchange takes place between the populations
of tautog of the Cape Cod Bay region, and of the
the rocky coasts along the north shore of Massa-
chusetts Bay, for the peaks of abundance (as
judged from the reported landings) have fallen in
different years in these two regions.

April 29 (1949) and May 1 (1950) are the earliest
dates at which we have heard of tautog caught
either in Massachusetts Bay or in Cape Cod Bay
(Duxbury in both instances).8 In 1950, which
appears to have been an "early" season, they were
reported as biting well in Cape Cod Bay by May
25 and at Duxbury by the last days of the month;

• 1890-1900; 1906-1911; 1917-1921, In particular. Data are also available for
earlier years.

' Total catch, Sagamore and Sandwich, 41,053 pounds, with 2 to 5 pound
nets or sets of nets working in the different years; Brewster to Provincetown,
18,549 pounds with 14 to 24 pound nets or sets of pound nets in operation
yearly.

' As reported in Salt Water Sportsman for May 25, 1950.

FISHES OF THE GULF OF MAINE

483

further up the Bay, however, as at Cohasset and
Swampscott, very few are caught before July. In
most years the best catches are made in August,
September, and into October, and we have not
heard of a tautog taken anywhere in the Gulf after
early November at latest.

The tautog that frequent any particular ground
in the Massachusetts Bay-Cape Cod region may
be expected to pass the winter in a more or less
inactive state close at hand, as they do farther
south (p. 480). But we have no first-hand
information in this respect.

Presumably the tautog spawn chiefly in June
in Cape Cod and Massachusetts Bays, as they do
along southern Massachusetts; perhaps into July.9
But we have found no tautog eggs nor larvae in
our towings in the Gulf of Maine, nor have any
tautog less than 2 or 3 inches long been credibly
reported in Cape Cod Bay or to the northward (we
may have missed them as tautog spawn close in
to the coast) . And we have yet to learn whether
the fluctuating stock north of Cape Cod is main-
tained wholly by local reproduction, or is rein-
forced from time to time by immigrants from the
south. It would be especially interesting to know
how many tautogs find their way from Buzzards
Bay to Cape Cod Bay through the Cape Cod
canal.

Fluctuations in abundance. — The pound net
catches of tautog (averaged for 3-year periods) sug-
gest that a moderate and irregular rise in abun-
dance took place in the northern side of Massa-
chusetts Bay from 1890-1892 (yearly average, 140
pounds) to 1899-1901 (yearly average, 1,049
pounds), followed by a corresponding decrease so
extreme that none at all were reported from the
pound nets of Essex County for 1917 to 1919, in
the Massachusetts statistics,10 only 42 pounds for
1920, and none for 1921. The local stock seems
next to have built up again about to its former
level, to continue so during the period 1928-1938. u
Our angler-correspondents report that some tautog
are caught along the Essex County rocks every
slimmer, since then. But the fisheries statistics
have not afforded information as to the tautog
situation there for the past few years.

• In 1950 the "spawning run" was reported as about over in Cape Cod Bay
by the end of the first week in June (Salt Water Sportsman for June 9, 1950).

» One hundred and fifty-eight pounds were reported for that year in the sta-
tistics of the U. S. Bureau of Fisheries.

" Landings. Essex County, 1928-1931, 1933, 1935, 1938, 0-803 pounds, average
about 300 pounds. There is no reason to suppose that the 10,700 pounds re-
ported for 1937 came from the Gulf of Maine. See footnotes p. 415; and p. 422.

In the Cape Cod Bay region (again according to
statistics of the landings) tautog seem to have been
scarce for some years through 1890; then to have
increased in numbers to a rather pronounced peak
of abundance in 1895-1900, when the reported
catch averaged about 13,190 pounds yearly (the
maximum 22,264 pounds in 1895), an increase that
came 5 or 6 years earlier than the upswing recorded
for the north shore of Massachusetts Bay. There
appear to have been fewer in 1899 (6,282 pounds
recorded) ; perhaps not more than half as many in
1906 or in 1907 (3,168 pounds and 2,934 pounds
reported) when the publication of the catches by
towns was resumed, and apparently rather fewer
still during the 4-year period 1908-191 1.12

The Cape Cod Bay population seems to have
been at about this same level in 1917, and tautog
seem to have been more plentiful again in 1918,
when the very large catch of 36,000 pounds was
reported from the pound nets along the shore line
of Brewster. But they fell, then, to so low an ebb
that the reported yearly catches for 1919 and 1920
were only 801 and 877 pounds, respectively, and 44
pounds in 1921. Catch records tell nothing as to
the status of the tautog in Cape Cod Bay since
1921. I3

The disappearance of the eel grass (Zostera)
about 1930-1931, must have altered their local
habitat for the worse. But the stock seems to
have built up again with the reappearance of eel
grass here and there. And tautog have been plen-
tiful enough around Cape Cod Bay during recent
summers for party boats, hand-lining, to have made
good catches there day after day. The traps at
Barnstable continue to take some even though
they are set on sand bottom, with their best catches
in autumn when a single lift of 4 traps sometimes
yields as much as 400 pounds.

According to local report, 1950 was a very good
tautog season in Cape Cod waters. But the com-
mercial fishermen took few or none smaller than
one-half pound that year.14 What this presages
for the future remains to be seen.

Importance. — Tautog are not plentiful enough
anywhere north of the elbow of Cape Cod to be of
any great commerical importance, and never have

" Maximum, about 3,900 pounds in 1910; minimum, 635 pounds in 1909;
yearly average, about 1,400 pounds.

■> There is no way of knowing how great a part of the catches reported in
subsequent years from "Barnstable County" came from the Cape Cod Bay
shore; i. e., from the Gulf of Maine.

» Information supplied by Henry Lyman.

484

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

been, but there is a ready sale for all that are
brought to market, most people thinking this a
very good table fish. And with so few fishes in the
Gulf of Maine that can be classed as "game"
(that is, affording sport on rod and reel), we maj'
well wish the tautog were more plentiful there, for
they put up so strong a resistance that tautog fish-
ing is very good sport indeed.

Along the stretch from Manomet Headland,
Plymouth, to Cape Ann, tautog are caught either
from a boat at anchor over submerged ledges or
bouldery bottom, or by casting with a long rod
from dry ledges or from the rocky coast line. In
either case, the fish are so local and irregular in
distribution (depending on the food supply and
also on the contour of the rocks) and so stationary
that it is worth fishing for them only in certain
spots. Even so, a few feet one way or the other
may mean the difference between success and fail-
ure. In Cape Cod Bay, however, where the tautog
are on smooth bottom, they lie in little openings
among eel grass (whenever there is any), "with just
their snouts sticking out" as an angler friend writes
us,15 "and, by lowering a fiddler or hermit crab in
the clear spot in front of them, they will be caught
in very shallow water."

Fishing the Cohasset rocks, we have found
green crabs (Carcinides) the most attractive bait,
whole if small enough, cut if larger; rock crabs
(Cancer), or hermit crabs second best; large
snails or cockles (Polynices) fairly good; lobster
would perhaps be best of all, were it not so
expensive. Mussels are often successful. And
small whole clams are good, hooked through the
"neck", (actually the siphon) with the shell
cracked so as'to let the juices escape, but they are
next to worthless if shelled because they are
stolen almost at once by the swarms of cunners.
Anglers tell us that the same baits are used along
the north shore of Massachusetts Bay. In Cape
Cod Bay, where tautog are caught on smooth
bottom (p. 480), the baits most used are hermit
crabs and fiddler crabs.16 We once had a good-
sized tautog strike a sea worm (Nereis), behind
our boat, while trolling for striped bass.

When a tautog bites, it passes the bait back to
the pharyngeal teeth, to crush the shell before
swallowing; in doing so he gives several distinctive
jerks or twitches. This is the time to hook him;
many are missed by being struck too soon by
anglers not experienced in the ways of the tautog.

THE REMORAS OR SHARK SUCKERS. FAMILY ECHENEIDAE

The several remoras are easi!y distinguished
from all other fishes by the fact that the spiny part
of the dorsal fin is modified into a flat oval sucking
plate, composed of a double series of cartilaginous
crossplates with serrated free edges, and situated
on the top of the head and neck. All the remoras,
too, are slender of form with the lower jaw
projecting well beyond the upper. Their mouths
are armed with many small pointed teeth; their
soft dorsal and anal fins are about the same in
form and size, the one above the other; and their
pectoral fins are set high up on the sides. The
lower surface of the head is convex, the upper flat
(a very conspicuous feature) with the lower surface
of the body nearly as deeply colored as the upper
so that the back is often mistaken for the belly.
The members of this family all attach themselves

to other fishes, or to sea turtles, by their sucking
disk, usually clinging to the sides of their hosts,
but often within the mouth or gill cavities of the
larger sharks and of the giant rays.17 They are
carried about in this way, and they feed on the
scraps from the meals of their transporters. All
the remoras are tropical; they appear only as
strays in boreal seas, usually fast to sharks or
to swordfish.

We follow Sumner, Osburn, and Cole 18 in
uniting under one species the shark sucker
(naucrates), with more than 21 plates but a
sucking disk less than one-fourth as long as the
body, and the pilot sucker (naucrateoides) , with
only 20 or 21 plates but longer, fishes that are
otherwise indistinguishable one from the other.

11 Quoted from a letter from Henry Lyman.

" Not having fished there for tautog, we welcome this information from
Henry Lyman.

" Oudger (Natural History, vol. 22, No. 3, May- June 1922, pp. 243-249)
gives an interesting account of this habit.
>» Bull. U. S. Bur. Fish. vol. 31, Pt. 2, 1913, p. 766.

FISHES OF THE GULF OF MAINE

485

KEY TO GULF OF MAINE REMORAS

1. Pectoral fins pointed; ventral fins attached to the belly for less than one-third of their length.. Shark sucker, p. 485
Pectoral fins rounded; ventral fins attached to the belly for more than half of their length 2

2. Dorsal fin of 29 rays or more; at most 17 plates in the sucker Swordfish sucker, p. 486

Dorsal fin of only about 23 rays; about 18 plates in the sucker Remora, p. 487

Shark sucker Echeneis naucrates Linnaeus 1758 margins are close below the overlapping edge of

the sucking plate.

Pilot sucker; White-tailed sucker n~i~~ tu i j *• ± ■ i * j i

Color. — ine general ground tint is slaty or dark

brownish gray, with the belly nearly as dark as the
Jordan and Evermann, 1896-1900 p,x 2269-2270 as back Each gide ig marked b ft brQad darker

Echeneis naucrates Linnaeus 1758 and E. naucrateoides, , . . . , , . . ,

Zuiew 1789 brown or sooty stripe with white edges, that runs

from the angle of the jaw to the base of the caudal

Description. — The most distinctive characters fin but is interrupted by the eye and by the pectoral

of the shark sucker are mentioned above. It is a fin. The caudal fin is velvety black with white

very slim fish, 11 or 12 times as long as it is deep, corners, a character noticeable enough to give rise

nearly round in cross section, and tapering to a to a vernacular name. The dorsal and anal fins

very slender caudal peduncle. The sucking plate, are dark slate color or black, more or less margined

reaching from close behind the tip of the snout with white. The pectorals and ventrals are

back over the nape of the neck even with the black, either plain or more or less pale edged,

middle of the pectoral fin, is about as broad as the Size.— Reaches about 38 inches,

head, flat, oval, and with 20 or more very con- Gmeml ^^.-Cosmopolitan in warm seas,

spicuous transverse plates. The soft dorsal fin n(Jrth as & tQ Haljf Noya gcoti „ on ^

(32 to 41 rays) and the anal fin (31 to 38 rays) both Atkntic Coagt of North America

ongmate about the mid length of the body, and , , .

they both extend nearly to the base of the caudal Occurrence in the Gulj of Mame.So far as we

fin. Both of them taper, too, from front to rear, can lea™ no sha rke^fe of this sPecies has befn

but the anal is more concave in form than the reported from the Gulf for many years; m fact the

, i mi j i C ■ r i ii • u only positive records of it there are for one taken

dorsal. The caudal fin is slightly concave in old - , , ,. , . . ™ ^.

c i i . • -. ,i ,i from the bottom of a fishing boat m Boston Bay

fish but m young ones its central rays are the . . «, , , , ,

longest. The ventral fins are pointed like the ^e time prior to 1839; » for a second reported by
pectorals below which they stand, and their inner Wheatland from Salem Harbor (^identified by
rays are attached to the skin of the abdomen for .» Leim Proc. Nova scotian inst. sei., vol. 17, pt. 4, 1930, P. xuv.

Only a short distance. The broad-based pectoral * Dumbed and illustrated by Storer (Fishes of Mass., 1867, p. 210, pi. 32.

fins are set so high up on the sides that their upper » jour. Essex Co.. Nat. Hist, soc., vol. 1, No. 3, 1852, P. 125.

WlMffi

IWMfil-iM/

Figure 251. — Shark sucker (Echeneis naucrates), 11-inch
specimen, Tortugas, Florida. Below, top view of head
of a specimen about 18 inches long from Boca Grande
Pass, Florida. Drawings by H. L. Todd.

486

FISHERY BULLETIN OF THE FISH AND "WILDLIFE SERVICE

Goode and Bean as naucrateoides) ; and for a third
reported by Goode and Bean 22 as taken at the
mouth of the Merrimac Eiver in June 1870. And
Leim 23 reports one from Halifax Harbor, Nova
Scotia, during the fall of 1928. It is only as the
rarest of strays that it ever wanders north of Cape
Cod, clinging to some ship (for such is a common
habit in its tropical home) or to some shark.

Swordfish sucker Remora brachyptera (Lowe)
1839 2i

Jordan and Evermann, 1896-1900, p. 2272

Description. — This is a stouter fish than the
shark sucker (p. 485), being only about seven times
as long as it is deep (counting the caudal fin) and
about as thick through the shoulders as it is deep,
with a thicker caudal peduncle. And although
the sucking plate is as long, relatively, it consists
of only 14 to 17 ridges. Furthermore, the pectoral
fins of the swordfish sucker are relatively shorter
than those of the shark sucker, softer, and rounded
instead of pointed, while the upper margins of
these fins are not so close to the edge of the sucking
plate. The ventral fins, too, are attached to the
skin of the abdomen along their inner margins for
at least one-half their length, as noted above
(p. 485). The long dorsal fin (29 to 32 rays) of the

" Bull. Essex Inst., vol. 11, 1879, p. 20.
a Proc. Nova Scotian Inst. Sci., vol. 17, No. 4, 1930, p, XLvi.
« Jordan, Evermann, and Clark (Rept. U. S. Comm. Fish. (1928), Ft. 2,
1930, p. 449) place this species in the genus Remoropsia Gill, 1864.

swordfish sucker serves to separate it from the
remora (p. 487).

Color. — Described as light reddish brown above
and as darker below, with paler dorsal and anal
fins. A distinctive feature is that it lacks the side
stripes and white fin edgings so characteristic of
the shark sucker.

Size. — A length of 12 inches is the maximum so
far recorded.

General range. —Warm and warm-temperate seas
generally, probably paralleling that of the sword-
fish.

Occurrence in the Oulj of Maine. — Goode and
Bean's 25 description of this sucker as not infre-
quently accompanying swordfish into Massa-
chusetts Bay probably applies to the whole Gulf
except the Bay of Fundy, for specimens have been
brought in from near Matinicus Rock and near
the Isles of Shoals; fishermen occasionally speak
of seeing "suckers" clinging to the swordfish they
harpoon on the offshore Banks; sometimes several
fastened to a single swordfish. But they also
report far more swordfish lacking these uninvited
guests than carrying them, and this has been the
case with the few fish harpooned by the Grampus
during our cruises in the Gulf. Suckers are de-
scribed by eyewitnesses as usually fast to the
shoulder of the swordfish, nor have we heard of one
actually within the gill cover of the latter, though
very likely they refuge there, for one has been

»■ Bull. Esesex Inst., vol. 11, 1879, p. 21.

Figure 252. — Swordfish sucker (Remora brachyptera),
side view and top of head (below), off Halifax, Nova
Scotia. From Jordan and Evermann. Drawings by H.
L. Todd.

FISHES OF THE GULF OF MAINE

487

taken from the gill cavity of a sunfish {Mola mola) ;
others have been found in the mouths and gill
cavities of large sharks.

Nothing beyond this is known of its way of life.
Presumably it feeds on fragments of the fish killed
by its host, as the shark sucker does whose actions
are better known. Presumably, too, it is an active
a swimmer as are its relatives. Nothing is known
of its breeding habits.

Remora Remora remora (Linnaeus) 1758
Jordan and Evermann, 1896-1900, p. 2271.

Description. — The chief distinctions between the
remora and the swordfish sucker is that it has a
larger number of ridges in its sucking plate on the
average (16 to 20, as against 14 to 17), and that
there are only 22 to 25 rays in its dorsal fin, whereas
the swordfish sucker has 29 to 32. Like the latter,
it is a stouter fish than the shark sucker (p. 485),
and its ventral fins are similarly attached to the
skin of the abdomen along their inner edges.

Color.- — -Uniform brownish, blackish, or sooty,
both above and below.

Size. — Maximum length about 18 inches.

Habits. — Very little is known of the life history
of the remoras. The young fry of this, and of
other species of Remora have been taken in the
open Atlantic, usually in June or July which sug-
gests a sharply limited spawning period. A
remora may join a shark, or other host, when
only about 1% inches (3 to 4 cm.) long.26 But we
have yet to learn how long or how constantly one

may accompany a single shark, or how often it
may transfer from one host to another.

General range. — Tropical seas generally; very
common in the West Indies, occasionally north to
New York and to Woods Hole, and only a stray
north of Cape Cod. It is usually attached to large
sharks or to sea turtles.

Occurrence in the Gulf of Maine. — The only
Gulf of Maine records for the remora, up to 1925,
were of one taken many years ago in Salem Har-
bor, no doubt brought thither clinging to the
bottom of some ship in from a southern voyage,
as Goode and Bean 2? remarked; and of one in
the Museum of Comparative Zoology that was
taken at Provincetown long past. More recent
records are of one found clinging to the bottom
of a lobster trap in Portland Harbor in 1931,
probably brought in by some West Indian schooner,
several of which had recently been in the harbor;28
of a second found sucking to the gills of a blue
shark Prionace glauca caught on the northeast
edge of Georges Bank, August 1 of that same
year; and of a third fastened to a shark of the
genus Carcharhinus that was caught at the surface
over the southeast slope of Georges Bank in
July 1939.29

Records from farther east and north along the
American coast are of one taken from a blue
shark, 10 miles off Cape Sable, June 1, 1933;30
of another (also from a blue shark) west of Sable
Island, September 9, 1934; and of two Uken from
sharks on St. Pierre Bank, south of Newfoundland,
one of them on August 13, 1936,31 the other on
October 7, 1937.32

Figure 253. — Remora (Remora remora). After Day.
THE SAND LAUNCES. FAMILY AMMODYTIDAE

The slender, round-bodied sand launces suggest
small eels in general appearance. Eel-like, too,
they lack ventral fins, and they swim with eel-
it Taning, Nature, vol. 20, 1927, p. 224.
>' Bull. Essex Inst., vol. 11, 1879, p. 21.
« Reported to us by the late Walter H. Rich, of the U. S. Bureau of Fisheries.

like undulations from side to side. But they are
not even remote relatives of the true eels, from

" The last 2 are in the collection of the Museum of Comparative Zoology.
» Vladykov, Proc. Nova Scotian Inst. Sci., vol. 19, 1935, p. 7.
" McKenzie and Homans, Proc. Nova Scotian Inst. Scl., vol. 19. 1938, p. 279
» McKenzie, Proc. Nova Scotian Inst. Sci., vol. 20, 1939, p. 18.

488

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

which they are distinguishable at a glance by their
forked caudal fin, separated by an appreciable
space from both dorsal and anal fins; by their wide
gill openings; and by the presence of a large bony
gill cover, not to mention other anatomic charac-
teristics equally important if less obvious. Only
one species inhabits our Gulf.33 The larger North
European launce (A. lanceolatus) , which grows to
12 inches and has 2 stout teeth on the roof of its
mouth, has no representative in our side of the
Atlantic.

Sand launce Ammodytes americanus De Kay
1842 34

Sand eel; Launce; Lant

Jordan and Evermann, 1896-1900, p. 833.

Description. — The sand eel is a slender fish, its
body about one-tenth as deep as it is long (not
counting caudal fin), with long head and sharply
pointed nose, wide gill openings, and large mouth
with the lower jaw projecting far beyond the
upper. The jaws are toothless, and there are
no teeth on the roof of the mouth. There is one
long low dorsal fin, soft-rayed (59 to 64 rays;
no spines), rising somewhat in front of the tips of
the pectorals and running back along the whole
length of the body nearly to the base of the
caudal fin. The anal (28 to 32 rays), similar in
outline and equally lacking spines, originates
slightly behind the middle of the dorsal and runs
equally far back. The tail is forked. The pointed
pectorals are set very low down on the body,
and there are no ventral fins. The scales are

M Asecond species of launce (the Arctic Ammodytes dubius Reinhardt 1838),
thought to be characterized by having more fin rays (65-67 dorsals, 33-36
anals), has been reported from Boston by Giinther (Catalogue Fishes British
Mus., vol. 4, 1862, p. 387), and from Woods Hole by Smith (Bull. U. S. Fish.
Comm., vol. 17, 1898, p. 95), but it is probable that the specimens in question
were merely large Ammodytes americanus. In fact it is doubtful whether
there is any sound distinction between the A. dubius of Greenland and the
European A. tobianus on the one hand, and the American A. americanus
on the other.

1 Our sand eel is so closely allied to the common European launce (Ammo-
dytes tobianus) that we doubt whether the distinction between the two
(more slender form and longer head of americanus) will stand the test of
time.

small, lying in cross series on the sides of the
body between numerous skin folds that run
obliquely downward and backward, and there
is a low ridge of skin on either side along the
belly.

The readiest field marks for the sand eel among
Gulf of Maine fishes are its slender form and
sharply pointed snout, coupled with long dorsal
fin (separated from the caudal) and the absence of
ventral fins. The only fishes with which one
would be apt to confuse it are young eels, but in
these the dorsal, caudal, and ventral fins are
confluent, not separate, and the tail is rounded,
not forked.

Color. — Authors differ in their accounts of the
colors of the sand eel, probably because its
iridescent luster fades at death and because it
varies in shade on different bottoms. The many
we have handled have been olive, brownish or
bluish green above, with silvery lower sides and
a duller white belly. Some have a longitudinal
stripe of steel-blue iridescence 36 along each side,
but others lack this.

Size. — Mature sand eels run from about 4 inches
to about 6 inches in length as a rule, with a few
as long as 7 inches.36

Habits. — Sand eels are found chiefly along sandy
foreshores, also over the shoaler parts of the off-
shore fishing banks; they are seldom seen off
rocky parts of the coast, or over muddy bottoms
in deep water. They usually congregate in dense
schools, often of thousands of individuals, and
they swim as an eel does, by sidewise undulations
that run along the body from front to rear, which
makes them easy to recognize in the water.

The most interesting habit of the sand eel is
the custom it has of digging itself several (4 to 6)
inches deep in the sand, into which it burrows

" In the European sand launce (Ammodytes tobianus), the sides are de-
scribed as with lines of tiny brown dots and the tip of the snout as black-
ish, especially in the young fish.

« According to Bean (Bull. New York State Mus. 60, Zool. 9, 1903, p. 376)
the majority are from 4% to 7 inches; and the largest we measured was about
7 inches long. Storer (Fishes of Mass., 1867, p. 217) credits them with a
maximum length of 12 inches; seemingly this was an error.

Figure 254. — Sand launce (Ammodytes americanus), Nantucket. From Goode. Drawing by H. L. Todd.

FISHES OF THE GULF OF MAINE

489

with great speed, opening the way with its sharp-
pointed snout. It often does this above low-
water mark at high tide to await the return of the
tide. We have often seen them vanish with
surprising rapidity when alarmed by clam diggers,
and we cannot improve on Goode's 37 account of
seeing "a great section of the beach" in Province-
town harbor become "alive with dancing forms
of dozens of these agile fishes" when he stuck his
clam-hoe into the sand. It has been suggested
that they spend a large part of the time so buried,
and that their sudden appearances and disap-
pearances are to be explained thus, rather than
as evidence of their wanderings or migrations.
It is not known whether they follow this habit
only in shoal water where they come under
direct observation, or whether they also burrow
into deeper bottoms. If the burrowing habit is
for refuge, it is not always successful, for, as
Smitt 38 remarks, porpoises have been seen rooting
them out of the sand.

In Scandinavian waters sand eels feed on all
sorts of small marine animals, but chiefly on small
Crustacea, especially on copepods, and on fish fry,
including their own kind. Worms have also been
found commonly in the stomachs of sand eels,
but it is not likely that they catch these while
burrowing, as some writers have suggested.

The sand eel plays a very important role in the
economy of northern seas as food for larger ani-
mals. Finback whales devour them greedily when
they find them in abundance, as happened in
Cape Cod Bay in June 1880, when launce ap- .
peared in swarms early in the month followed by
finbacks a few days later. Porpoises, too, find
them a staple article of food, and sundry preda-
ceous fish such as cod, haddock, halibut, silver
hake, salmon, mackerel, striped bass, and blue-
fish. When sand eels are fleeing from their pur-
suers especially from the silver hake, which does
not hesitate to follow them up on the sand, they
often strand in such numbers as to cover the flats.

Sand eels' noses are so sharp that when they
are swallowed by cod, and perhaps by other fish,
they sometimes work right through the stomachs
and into the body cavities of their captors, to
become encysted in the body wall.

The spawning of the American sand eel has not
been observed so far as we can learn.39

Ripe specimens of the European species (to-
bianus), both male and female, have been taken
throughout the year, a phenomenon that has
given rise to widely differing views as to its spawn-
ing season. But the chief production of its eggs
takes place in autumn and early winter, at least
in the southern part of the North Sea, as Ehren-
baum 40 demonstrated, both by dredging them in
large numbers, and by the fact that its larvae are
extremely abundant there from January to March,
but have seldom been taken at other seasons.

The occurrence of larvae suggests that the sea-
son is about the same for the American form in
the Gulf of Maine. Thus the Fish Hawk towed
numbers of larval launce (identified by R. A.
Goffin of the Bureau of Fisheries and by Mrs. C.
J. Fish) near Provincetown and in Cape Cod Bay,
during December, January, and February, 1924-
1925, evidence both that this part of the Gulf of
Maine (where adults are abundant) is the site of
considerable reproduction, and that spawning
commences as early as November there. Fry
have been taken in March at Woods Hole while
the Albatross towed a number of larvae of 11
to 17 mm. on the western part of Georges
Bank on February 22, 1920. It seems that the
spawning season is progressively later, however, to
the northward, for we took larvae only a few
days old (7 to 8 mm. long), with the yolk still
showing, off Newburyport, Mass., on March 4 in
1921, while the Canadian Fisheries Expedition of
1915 obtained an abundance of slightly older
stages (7 to 15 mm. long) off the southeast coast
of Nova Scotia in May.

Evidently the sand eel breeds successfully
throughout the more northern part of its range,
for its larvae have been found, widespread, over
the Nova Scotian Banks, in the Gulf of St.
Lawrence northward nearly to the Strait of Belle
Isle, throughout the Grand Bank region, off the
east coast of Newfoundland and off the outer coast

« Fish. Ind. D. S.; Sect. 1, 1884, p. 244.
" Scandinavian Fishes, vol. 2, 1895, p. 579.

'• Hind (Fish. Comm. Halifax, 1877, Pt. 2, p. 7) describes the launce in the
Gulf of St. Lawrence as "depositing their large reddish-colored ova on the
sand between high and low water.*' This account, however, is widely at
variance with the spawning habits of their European representative (Am-
modytes tobianus) and with the seasonal occurrence of their larvae; and was
probably borrowed from the larger European sand eel (Ammodytes lanceo-
latus).

« Wissenschaftliche Meeresuntersuchungen, Helgoland, Neue Folge, vol.
6, 1904, p. 184.

490

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

of Labrador, north to Sandwich Bay.41 How far
south it may do so is not known.

Sand eels were formerly thought to spawn on
sandy beaches above low-water mark while bur-
rowing in the sand, but their eggs have never been
found in such a situation, and Ehrenbaum proved,
by dredging them in large numbers, that those of
the European species (Ammodytes tobianus) are
actually deposited in depths of 10 fathoms or so,
on sandy bottom where they stick fast to the grains
of sand. His experience also suggests that they
resort to very definite grounds for spawning, all of
which probably applies as well to the American
form as it does to the European.

The eggs of the American launce have not been
seen. Those of the European tobianus are oval,
0.72 to 0.97 mm. in greatest diameter, with a yel-
low oil globule of 0.25 to 0.31 mm., and they are
described as of an orange tint. The larvae are
very slender, and about 7 mm. long by the time
the yolk is absorbed. The dorsal and anal fin
rays are visible when the larva is about 18 mm.
long, but it is not until the little fish is upward of
25 mm. long that the tail begins to assume its
forked outline; this is a convenient field mark for
distinguishing between the launce and the herring,
in which the tail is deeply forked from a much
earlier stage. The early larval stages are easily
recognizable by their slender form combined with
the fact that the vent opens at one side, just as
among the cod tribe, not at the margin of the larval
fin fold, so that it apparently ends blind.

Figure 255. — Larva of European A. tobianus 6.6 mm.
After Ehrenbaum and Strodtman.

Figure 256. — Larva of European A. tobianus, 20.5 mm.
After Ehrenbaum and Strodtman.

Sand Launce (Ammodytes).

The older larvae resemble the corresponding
stages of the rock eel (p. 493) in their slim form,
and in the location of the vent slightly behind the

« Dannevig, Canadian Fish. Exped. (1914-1915) 1919, p. 29; Frost, Res.
Bull. 4, Newfoundland Dept. Nat. Res., 1938, Chart 8.

middle of the trunk (it is located farther back in
the similarly slender larvae of the herring tribe),
but may be recognized by the row of black pigment
cells along the dorsal side of the intestine instead
of along the ventral side, and by their pointed
noses.

The rate of growth of our launce has not been
studied. But it is probable that the small ones
of 3 to 4 inches, which are plentiful from July until
September, are yearlings; those of 4 inches and
upward probably are 2 years old, or more.

General range. — Atlantic coast of North America
from Cape Hatteras to the Gulf of St. Lawrence,
northern Newfoundland and northern Labrador,
perhaps to Hudson Bay (p. 491). Its European
relative, A. tobianus, occurs from Greenland, Ice-
land, northern Scandinavia and the White Sea
south to Spain.

Occurrence in the Gulf of Maine. — The sand eel
is very plentiful along the coast from Cape Cod to
Cape Sable wherever there are sandy shores,
but it is seldom seen off the rocky parts of the
coast line. Thus it is rather scarce in the Bay of
Fundy except locally, but is common on the sandy
beaches that break the bold northern shores of the
Gulf here and there. They swarm on the strands
of Cape Cod Bay, a phase of their distribution
associated with their burrowing habit. Here one
may see schools of them throughout the summer
in shoal water close in to tide mark, swimming
with the curious undulating motion so character-
istic of them; and they continue plentiful there in
some years during the winter, when great numbers
are sometimes cast on the beach in stormy weather.
Sand eels are to be taken in shallow water on
sandy flats throughout the year about Woods Hole
also, but they are never so plentiful there in winter
as they are in fall and spring. And since a general
decrease in their numbers close inshore seems to
take place during the cold months in the more
northern part of their range as well, it is probable
that a considerable proportion of the local stock
moves out into deeper water for the winter, to
return in spring, as most of the launce do in north
European seas.

In Scandinavian waters this vernal inshore
movement takes place in May as the coast waters
warm up, and it is probable that the seasonal
schedule is much the same in the Gulf of Maine,
judging from its temperature. The sand eels
may also be expected to leave some of the shallow-

FISHES OF THE GULF OF MAINE

491

est bays of our Gulf in midsummer, when the
water is at its warmest, to work in again in early
autumn, such being their custom near Woods
Hole.

There is no reason to suppose that they inhabit
the central deeps of the Gulf of Maine regularly
unless some of them repair thither in winter,
though it would not be astonishing to find an odd
sand eel in deep water at any time; in fact, we
towed a young one about 1% inches long over the
deep basin southeast of Grand Manan on June
10, 1915. But they must be plentiful on Nan-
tucket Shoals, for they were found in the stomachs
of cod caught there by the Halcyon and by the
Albatross II, from time to time throughout the
springs and summers of 1928-1930. There are
also sand eels over the shallows of Georges and
Browns Banks, whence they have been brought
to the Bureau of Fisheries by fishermen on several
occasions.42

Further northward, fishermen are familiar with
them all along the outer coast of Nova Scotia and
on the Scotian Banks; they are so common near
Canso that a seine, dragged on a sandy beach
there "captured hundreds in a short time";43 they

are reported from Prince Edward Island 44 and
from the Magdalens in abundance,46 here and there
along the north shore of the Gulf; from the Strait
of Belle Isle; also from Sandwich Bay and Sloop
Harbor in southeastern Labrador; and they are
to be expected all along the outer Labrador coast,
for we have seen one taken at Eclipse Harbor,
near Cape Chidley.46 Sand eels have also been
found on the southern side of Hudson Strait, and
in the southern and western parts of Hudson
Bay.47 But these northern specimens may repre-
sent a distinct race for they have more fin rays
(63-67 dorsal, 28-33 anal); Vladykov, in fact,
has classed them as a new subspecies (hudsonicus)
of the Greenland launce (A dubius).

Sand eels are locally plentiful southward as far
as northern New Jersey where we have seen
squirrel hake gorged with them; they are reported
as "common" as far as Cape May at the entrance
to Delaware Bay,48 and have been reported as far
south as Cape Hatteras.49

Importance.- — It is only for bait that sand eels
are of any commercial value in our Gulf, for which
purpose 67,800 pounds were landed from the traps
in Massachusetts in 1919, 20,000 pounds in 1946.

BLENNY-LIKE FISHES. FAMILIES LUMPENIDAE, PHOLIDAE, AND STICHAEIDAE

The blenny tribes are characterized among Gulf
of Maine fishes by the position of their ventral
fins, which are under or in front of the pectorals,
combined with a single dorsal fin that is spiny
throughout its length and extends the whole
length of the trunk, and with a slender form, eel-
like in some of them. The only other Gulf of
Maine fishes that resemble them are the wolffishes
(p. 502) and the wrymouth (p. 500) , but both of these
lack ventral fins, which are present in all our blen-
nies, though they may be very small. Further-
more, the tremendous canine tusks and molar teeth

of the wolffish (p. 503) have no counterpart among
the blennies, and the peculiar face of the wrymouth
is equally distinctive for it (p. 501). The eelpout
(Macro zoarces) also is somewhat blennylike in ap-
pearance, but the greater part of its dorsal fin is
soft rayed, not spiny; and its anal fin is continuous
with its caudal fin.

The blennies are a numerous tribe of small
carnivorous shore fishes, widely distributed both
in northern and in tropical seas. Five species
are known in the Gulf of Maine.

KEY TO GULF OF MAINE BLENNY FISHES

Body very slender, about 18-20 times as long as it is high Snake blenny, p. 494

Body only moderately slender, not more than 8 to 10 times as long as it is high 2

There is a row of conspicuous roundish black or dusky spots along the dorsal fin 3

There is only one large and conspicuous dark spot on the dorsal fin, or none 4

« Kept. TJ. S. Comm. Fish. (1879) 1882, pp. 808, 812, 814, 817.
« Cornish (Contrib. Canadian Biol. [1902-1905] 1907, p. 84.)
« Letm, Proc. Nova Scotian Inst. Sci.; vol. 20, Pt. 2, 1940, p. 39.
« Coi, Contrib. Canadian Biol. (1918-1920) 1921, p. 111.

<• This specimen about 3 inches long, collected by C O. Iselin, now in the
Museum of Comparative Zoology, appears to be a typical americanus, for it
has 30 anal fin rays, and only 69 dorsal rays.

«' Vladykov, Contrib. Canadian Biol., N. Ser., vol. 8. No. 2, 1933, pp. 23-25.

« Fowler, Rept. New Jersey State Mus. (1905) 1906, p. 411.

» Jordan and Evermann, Bull. 47, U. S. Nat. Mus., Pt. 1, 1896, p. 833.

492

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

3. The pectoral fins are about as long as the body is high; the ventral fins are well developed, without noticeable spines,

and about as long as one-half the height of the body Arctic shanny, p. 497

The pectoral fins are only about one-half as long as the body is high; the ventral fins are minute (likely to be over-
looked) Rock eel, p. 492

4. Pectoral fins evenly rounded, their middle rays the longest; dorsal fin marked on its forward part with one large and

conspicuous dark blotch; only 43 or 44 dorsal fin spines Radiated shanny, p. 498

Pectoral fins with the lower rays longer than the upper rays and free at their tips; 58 to 61 dorsal fin
spines Shanny, p. 497

Rock eel Pholis gunnellus (Linnaeus) 1758 distances along the middle of the back and spread-

p ing out onto the dorsal fin, are the most character-

istic feature of the color pattern of this fish. The
Jordan and Evermann, 1896-1900, p. 2419. ground tint of the upper part is yellowish, olive

Description.— Tb* slender flexible trunk (only brown- reddisb> or ught red> matching the seaweed

about one-tenth as deep as it is long and about one- or the bottom> Wlth Pale> ^regularly rounded

half as thick as it is deep) , short head, and rounded cloudings on the sides, and with an oblique streak

nose of this little blenny suggest an eel, but the from the eye to the anSle of the Jaw- The belly

spiny nature of its dorsal fin betrays its true rela- vanes from Pale &*? to yellowish white. The

tionship. The dorsal fin (73 to 86 spines) extends pectoral, caudal, and anal fins are yellowish. We

from the nape of the neck back along the whole have seen a specimen (at Boothbay) that was

length of the trunk to the base of the caudal fin, bnck red above and below> USbt and dark mottled,

from which it is marked off by a shaUow notch flecked also Wlth tmy black dots> and Wltb the

only; and it is of uniform height from end to end. sPots on tbe dorsal fin dark red instead of black-

The anal fin (2 very short spines and 37 to 44 rays) Size.— The maximum length is about 12 inches

originates opposite the mid-length of the dorsal, to but few of tbose found are more tban 6 to 8 ^ches

which it corresponds in height and outline, and ionS-

runs back similarly to meet the caudal fin, from Habits.— Rock eels are often found along low
which it is marked off by a distinct notch, though tide mark, left by the ebb in little pools of water
there is no free space between the two fins. The under stones, or among seaweed, where they
caudal fin is small and rounded. The tiny await the return of the tide. But they are not
ventrals, set near together close in front of or confined to very shoal water as formerly supposed,
under the pectorals, are reduced to one very short for we have also collected them in considerable
spine and one rudimentary ray each. The pec- numbers both within the Gulf and on the offshore
toral fins, smaller than in our other blennies, are banks down to 40 fathoms, and one has been taken
a little longer than one-half the length of the head. as deep as 100 fathoms (p. 494) .
The skin of the trunk is clothed with very small They are rather local throughout their range,
scales, hardly visible, however, through the thick In some places one is to be found under almost
laj^er of slimy mucus with which the rock eel is every stone; on others you may turn rocks in
covered. The head is naked; the mouth is small vain; their presence or absence along any particu-
and oblique; the upper jaw is armed with several lar stretch of shore probably depends on the char-
rows of conical teeth, but the lower jaw has a acter of the bottom immediately outside, for this
single row only. fish prefers pebbly, gravelly, or stony ground, or

Color. — A row of about 10 to 14 round, black- shell beds, and not mud or eelgrass. Neither

centered and pale-edged spots, spaced at equal have we found them about the steep ledges so

^^^^t^->.

Figure 257. — Rock eel (Pholis gunnellus), Gloucester, Mass. From Jordan and Evermann. Drawing by H. L. Todd.

FISHES OF THE GULF OF MAINE

493

numerous along the rockbound coasts in the Gulf
of Maine. In Scandinavian waters according to
Smitt 60 they often take refuge inside large empty
mussel shells. But as he remarks, there is no
ground for the accusation that rock eels enter
live bivalves of any sort to devour them. Whether
they seek such places of concealment in deeper
waters is not known.

When disturbed they squirm like eels. Eel-
like, they swim by sidewise undulations, and they
are so active and so slippery (hence the name
"butterfish") that it calls for quick work to catch
one by hand, even in a very small puddle.

Very little is known of the diet of the rock eel,
except that it is carnivorous and that various
molluscan and crustacean fragments have been
found in their stomachs. Vinal Edwards records
small amphipods, shrimps, and worms in the few
that he examined at Woods Hole, but we have no
first-hand information to offer on this point.
In turn, rock eels have been found in the stomachs
of various larger fishes, especially of cod, in New
England waters.

So far as known the rock eel is resident through-
out the year wherever it is found ; at most it may
move out from the beach into slightly deeper
water in winter to escape chilling.

Breeding habits. — It is necessary to turn to
European sources for information about its breed-
ing habits, for its spawning has not been seen in
American waters. In the eastern Atlantic and
in the North Sea 61 it spawns from between tide
marks down to 12 fathoms or more, from Novem-
ber to February or March. And its spawning
season probably is the same in our Gulf, for eggs
apparently belonging to the rock eel have been
found off Rhode Island late in December.62 A
female from Peconic Bay, N. Y., contained 686
eggs.63 These, by European accounts, are about
2 mm. in diameter, opaque, whitish, but iri-
descent on the surface, with a single globule of
about 0.6 mm., and they are laid in holes or cran-
nies. In British waters the rock eel usually
chooses empty oyster shells, or the holes that are
made in the limestone rocks by the boring bivalve

Pholas for the purpose, but there are no oysters
in the Gulf of Maine, except in Cape Cod Bay,
and the local Pholas is unable to bore into the
hard granite rocks of our coast line, so the rock
eels must seek other nesting sites. Perhaps large
mussel shells may serve them, or any crevice.
The eggs are adhesive, and both the parents have
been observed rolling them, by coiling around
them, into balls or clumps an inch or so across,
in which they stick together.

In European waters incubation occupies from
8 to 10 weeks, during which period the parent fish
of both sexes have been seen lying close beside the
egg clumps. But since Ehrenbaum 64 described
the parent as "very negligent" in the aquarium, it
seems that they merely seek the nesting holes as
convenient shelters, and not that they actually
guard the eggs.

» Scandinavian Fishes, vol. 1, 1892. p. 223.

11 Macintosh and Masterman (Life Histories of British Marine Food-
Fishes, 1897, p. 210) and Ehrenbaum (WissenschaftlicheMeeresuntcrs., Hel-
goland, N. Ser., vol. 6, 1904, p. 160) give accounts of its spawning and of its
larval development.

u Tracy, 40th Rept. Comm. Inland Fish., Rhode Island, 1910, p. 151.

" Nichols and Breder, Zoologica, New York Zool. Soc, vol. 9, 1927, p. 159.

Figure 258. — Larva (European), 18 mm. After Ehren-
baum.

Figure 259. — Larva (European), 20 mm. After Ehren-
baum.

Rock Eel (Pholis gunnellus).

The larvae are much larger at hatching (about
9 mm.) and further advanced in development
than those of most of the fishes that lay buoyant
eggs. Older larvae of the rock eel resemble
corresponding stages of the launce and of the
snake blenny in their extremely slender form.
But they are easily distinguishable from both
these species by the presence of a row of small
black pigment spots below the intestine, instead
of above it, and from the herring (the only other
very slender larvae apt to be met in any numbers
in the Gulf at the same season) by the location of
the vent about midway of the body (fig. 259), and
by the fact that their tails are rounded, not forked.
The 12 black dorsal fin spots so characteristic of
the adult are first noticeable against the trans-
parent trunk in young fry of 25 to 30 mm. The

»• Wissenschaftliche Meeresuntersuchungen, Helgoland, N. Ser. vol. 6,
1904, p. 161

494

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

young fish (previously drifting near the surface)
sink to the bottom when 30-40 mm. long, an
event that takes place in late summer or early
autumn in the Gulf of Maine. Nothing definite is
known of the rate of growth of the rock eel after
its first autumn.

General range. — Shoal waters on both sides of
the North Atlantic; from Hudson Strait to the
offing of Delaware Bay on the American coast 65
and south to France on the European coast;
most numerous north of Cape Cod and north of
the English Channel.

Occurrence in the Gulf of Maine. — This little fish
is to be found all around the shores of the Gulf
from Nova Scotia to Cape Cod. Thus it is
definitely recorded at Yarmouth (Nova Scotia);
at various localities on both sides of the Bay of
Fundy where it occurs nearly everywhere (writes
Dr. Huntsman) ; at half a dozen points along the
Maine coast; on Cashes Ledge; at Portsmouth
(N. H.), where A. H. Clark of the U. S. National
Museum reports it plentiful; at Hampton (re-
ported to us by W. F. Clapp); in Ipswich Bay;
here and there along the northern shore of Mas-
sachusetts Bay; at Cohasset, on the southern
shore, where we have seen them; among the
stones and boulders of the Gurnet, off Plymouth,
where W. F. Clapp informs us that he has found
many while shore-collecting for mollusks; also at
Provincetown. And doubtless it is more uni-
versal than these records indicate, for being of no
consequence to the fisherman or to the angler, it
comes to notice only when scientific collections
are made.

We have taken its pelagic fry (20-39 mm. long)
in our tow nets off Seal Island (Nova Scotia) ; on
German Bank; near Mount Desert Island; off
Matinicus Island; and off Ipswich Bay, in April,
May, June, and August (a total, however, of only
6 stations), while Huntsman states that they are
caught in early summer in the Bay of Fundy,
evidence that it breeds all around the Gulf from
early spring on into the summer.

Recent records show that rock eels also occur in
considerable numbers on Nantucket Shoals, as
well as on Georges and Browns Banks, where we
have trawled a number, down to 40 fathoms, one

» In February 1930 Albatross II trawled 2 rock eels, 80 and 84 mm. long, in
23 fathoms and 38 fathoms, at lat. 3S°32' N., long. 74°24' W., and Iat. 38°18
N., long. 73°51' W.

as deep, even, as 100 fathoms,66 where many of
them have been found in the stomachs of cod
and pollock.67 They are described as common
inshore along outer Nova Scotian waters, eastward
from our limits; a and as "rather common" in the
southern side of the Gulf of St. Lawrence,69 where
Huntsman 60 classes them as characteristic of the
warm inshore waters, but where they are exposed
to temperatures close to the freezing point of salt
water in late winter and in spring.

They are reported from the southwest coast of
Newfoundland; from the estuary of the St. Law-
rence near Trois Pistoles,61 from Anticosti; from
the north shore of the Gulf of St. Lawrence; from
both shores of the Strait of Belle Isle; from the
southern and eastern parts of the Grand Banks ; 6*
from Bay Bulls, Conception Bay, and Trinity Bay
on the east coast of Newfoundland; from Hamilton
Inler on the Atlantic Coast of Labrador, and from
Fort Chimo, Ungava Bay, northern Labrador.

Importance.— This little fish is of no commercial
interest. But it appears to be of some importance
on the offshore banks as food for larger fishes.

Snake blenny Lumpenus lumpretaeformis
(Walbaum) 1792

Serpent blenny

Jordan and Evermann, 1896-1900, p. 2438, as Lumpenus
lampetraeformis.**

Description. — This is a very slender little fish
as its name implies, being only about one-twentieth
as deep as it is long, slightly flattened sidewise,
with head about one-eighth as long as body to
base of caudal fin, large eyes, wide mouth, and
blunt snout. It somewhat suggests a launce
(p. 488) in general form, but is much more slender
and eel-like. Also, its rounded tail (that of the
launce is forked), its large pectoral fins, spiny
dorsal fin (the launce has a soft dorsal only), and

w One was caught at this depth on the northeastern part of Georges Bank
August 5, 1932 (Sehroeder, Bull. 66, Boston Soc. Nat. Hist., 1933, pp. 5-6).

n We once found one alive in the mouth of a pollock caught at 34 fathoms
on Browns Bank.

" Vladykov and McKenzie, Proc. Nova Scotian Inst. Sci., vol. 19, 1935,
p. 103.

» Cox, Contrib. to Canadian Biol. (1918-1920) 1921, p. 112.

» Trans. Koy. Soc. Canada, Ser. 3, vol. 12, Sect. 4, 1918, p. 63.

•' Vladykov and Tremblay, Natural. Canad., vol. 62 (Ser. 3, voL 6), 1935,
p. 81).

M The pelagic larvae.

a The original spelling of this name by Walbaum (P. Artedi Qenera
piscium, Ichth., Pt. 3, 1792, p. 184) was lumpTetae-formis.

FISHES OF THE GULF OF MAINE

495

Figure 260. — Snake blenny (Lumpenus lumpretaeformis). Drawing by Louella E. Cable.

the fact that its lower jaw does not project be-
yond the upper, together with its color, serve to
separate it from the launce at a glance.

The chief anatomic feature (apart from its
slenderness) distinguishing it from the rock eel
(p. 492) is that the ventral fins (each of one short
spine and three longer rays) are well developed
and one-third to one-half as long as the pectorals,
slightly in advance of which they stand. The
pectorals, too, aro much larger than those of the
rock eel, and its dorsal fin, with 68 to 85 spines
and its anal with 49 to 62 rays are fully twice as
high, relative to the depth of the body, while
its anal fin originates farther forward ; the separa-
tion of dorsal and anal fins from the caudal is
more evident; and its eyes are noticeably larger.

The very slender body is the most obvious
difference between this species and its allies the
shanny (p. 497), the Arctic shanny (p. 497), and
the radiated shanny (p. 498), which are rather
robust. The outline of the caudal fin, which is
oval (more pointed in large fish than in small),
with the central rays much longer than the outer
ones, is diagnostic, too.

Color. — Whitish or pale brown on the back and
sides, with darker brown markings. On a 12-
inch fish taken off the coast of Maine the head
was pale brown, the sides of the body blotched
with brown, the dorsal fin marked obliquely with
18 pale bars, the caudal marked transversely with
8 bars, the anal rays pale brown against a color-
less membrane, the ventrals white, and the pec-
torals tinged with brown.

Size. — The largest one so far measured was 19
inches long.64

Remarks. — Vladykov's 66 discovery that New-
foundland specimens have a larger number of
dorsal fin-spines (85) and anal fin rays (62) and
a longer caudal fin (longer than the head) than
others from the St. Lawrence estuary (75-79
dorsal spines, 52-56 anal rays) shows that the
snake blenny tends to break up into local races.
Vladykov has made the St. Lawrence race the
basis of one new subspecies, americanus, and the
Newfoundland race the basis of another, terrae
novae; both of which have more spines and rays
than have been recorded for some eastern At-
lantic specimens. Gulf of Maine specimens, with
77-83 dorsal spines, and 56-59 anal rays, are
intermediate between the Newfoundland and Gulf
of St. Lawrence races in this respect.

Habits. — Little is known of the habits of the
snake blenny on either side of the Atlantic.
Although it is not found along the littoral zone,
it is a fish of comparatively shoal water, never
taken as deep as 100 fathoms (so far as we have
been able to learn) and apparently the most
common from a fathom or so below tide mark
down to 40 or 50 fathoms. And as most of the
specimens that have been caught in Scottish
waters were picked up by the foot rope of the
trawl, Sim's 6S suggestion that it burrows in mud

•* In the Museum of Comparative Zoology, trawled about 17 miles off Cape
Ann, lat. 42°28' N., long. 70°13' W., at 42 fathoms, In July 1931.
« Kept. Newfoundland Fish. Res. Comm., vol. 2, No. 3, 1935, p. 75-78.
» Jour. Llnnaean Soc. London, Zoology, vol. 20, 1890, p. 38.

496

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

or clay bottom probably is correct. The Albatross
II has taken it both on mud bottom and on hard .

Food.- — Amphipods, copepods, and other tiny
Crustacea, with very small starfish, small bivalves,
and holothurians have all been found in snake-
blenny stomachs in British seas. These blennies
are eaten in their turn by large fish, by cod and
halibut, for example, in Massachusetts Bay,67 by
pollock in the Bay of Fundy; and by cod in
Northumberland Strait, Gulf of St. Lawrence, as
Capt. Thor Iversen informed Dr. Hunstman
from his experience during the Canadian Fisheries
Expedition of 1915.

The spawning season has been stated as autumn
or winter in north Scandinavian seas, and it may
commence by late summer there, or by early
autumn, for Sim found its roe well advanced in
development as early in the season as the end of
April. Its drifting larvae have been taken in
tow nets from February to March in the Baltic,
and from March to May in the Gulf of Maine.

The eggs of this species have not been seen, but
they probably sink and stick together like those
of the rock eel. Apparently the larvae are of
considerable size at hatching, for the smallest we
have taken (the smallest on record) were about
11 mm. long, though they still lacked any trace of
the dorsal- and anal-fin rays. Snake blenny
larvae are very slender, resembling the correspond-
ing stages of the rock eel and of the launce in
general appearance, but they are distinguishable
from both of these by the fact that the vent is
situated considerably in front of the midlength of
the trunk. There is no danger of confusing them
with the young of the herring, the only other
very slender pelagic fish larva (besides rock eel
and launce) that is apt to be found in any numbers
in the Gulf of Maine in spring, for the tail of the
herring is forked from a very early stage and its
vent is situated much farther back than that of
the blenny (p. 91). Another distinctive feature
of the snake blenny larvae is the presence of a
large black pigment dot at the base of each
pectoral fin, and of a double row of 6 to 9 black
dots along the dorsal surface of the intestine with
several about the vent, which are very conspicuous
by contrast with the colorless body. Our largest
pelagic larva (41 mm. long) showed most of the
characters of the adult, although it was transparent

and had the arrangement of pigment characteristic
of the earlier larval stages.

General range. — Arctic and northern Atlantic
Ocean; south to Scotland, the Baltic, and the
southern part of the North Sea in the eastern side;
to the offing of southern New England in the
western side.

Occurrence in the Gulf of Maine. — It is probable
that this northern fish occurs in small numbers
around the coastline of the Gulf at some little
depth . Thus Hunstman reports it from St. Mary
Bay, Nova Scotia, in August and September ; from
Passamaquoddy Bay from April to August; and
in the open waters of the Bay of Fundy from
January on. It was recorded off Eastport in
1872; Albatross II trawled one specimen (about
12 inches long) 3 miles south of Great Duck
Island, near Mount Desert, Maine, in 28 to 33
fathoms, April 1927; two others (8 and 8% inches
long) 13 miles east of Boone Island, in 88 fathoms
in August 1928; one off the Isles of Shoals at 72-78
fathoms in August 1926; one at 42 fathoms on the
eastern slope of Stellwagen Bank, about 17 miles
off Cape Ann in July 1931. And Goode and
Bean 6S described it long ago as a common resident
in the deeper parts of Massachusetts Bay.

Our tow nettings, also, of 1920 yielded its
drifting larvae off Seguin Island; near Cape
Elizabeth; over Platts Bank; near the Isles of
Shoals; off Ipswich Bay; off Cape Ann; off Boston
Harbor; and in the southwest basin of the Gulf
off Cape Cod during March, April, and May —
evidence that it breeds successfully throughout
the southern part of its range. While it has not
been reported on Browns or Georges banks, it is
to be expected there.

It has never been reported from the banks along
the outer coast of Nova Scotia, so far as we know.
But it is so slender and active a fish that it can
easily pass through the meshes of any of the nets
that are used in our Gulf by commercial fisher-
men, hence is not likely to be brought in unless
it is sought for especially. And the experimental
trawlings by the Newfoundland Fishery Research
Laboratory did take it at several stations on the
Newfoundland Banks, as well as in the Gulf of
St. Lawrence, and also off the southeastern coast
of Labrador, while it has long been known from

" Ooode and Bean, Bull. Essex Inst., vol. 11, 1879, p. 10.

« Bull. Essex Inst., vol. 11, 1879, p. 10.

FISHES OF THE GULF OF MAINE

497

as far north on the Labrador coast as Ungava
Bay.69

Shanny Leptoclinus maculatus (Fries) 1837

Langbarn

Jordan and Evermann, 1896-1900, p. 2433.

Description. — The shanny resembles the snake
blenny somewhat in general appearance and in
the location and shape of its dorsal and anal fins,
but is not so slender (only 10 to 12 times as long
as it is deep instead of about 20 times). The most
important points of difference (aside from its more
robust form) are that the tail of the shanny is
about straight in outline instead of narrowly oval
or pointed as it is in the snake blenny; that the
lower rays of its pectoral are the longest and are
separate at their tips; and that the shanny has
only 58 to 61 dorsal fin spines, and 35 to 38 anal
fin rays.

Color. — Dirty-yellowish, paler below, the back
and sides marked with indistinct yellowish-brown
blotches of various sizes. The dorsal fin is de-
scribed as barred obliquely with about 10 rows of
brownish dots and the pectorals as cross-barred
with about 5 rows. These fins show no distinct
markings on the several preserved specimens we
have examined; the caudal fin, however, shows one
or two dark crossbars, even after preservation.

Size. — Maximum length about 7 inches.

Habits and food. — In Scandinavian waters the
shanny spends most of the year in deep water,
probably coming up to the shallows to spawn.
In the aquarium it keeps close to the bottom,
with the body extended and the pectoral fins ex-
panded, and apparently supports itself on the free
lower rays of those fins.70 Annelid worms and
pelagic amphipods have been found in shanny
stomachs; this is all that is known of their mode
of life. The shanny is supposed to spawn in

•• Kendall, Proc Portland Soo. Nat. Hist., vol. 2, No. 13, 1909, p. 224.
'• Smitt, Scandinavian Fishes, vol. 1, 1892, p. 230.

winter, but neither its eggs nor its larvae have
ever been seen.

General range. — An Arctic fish, known south to
Norway and Sweden in the eastern side of the
Atlantic, and to Cape Cod in the western side.

Occurrence in the Gulf of Maine. — Definite rec-
ords of this Arctic fish for the Gulf of Maine are
of several specimens that were collected in 40 to
90 fathoms in Massachusetts Bay by the U. S.
Fish Commission in 1887 ;71 one that we took in a
tow net near Boone Island on March 4, 1920;
one from the northeast part of Georges Bank,
August 1926, and four (4 to 4% inches long) that
were trawled off Chatham, Cape Cod, in 28 fath-
oms, May 1, 1930, by the Albatross II. This pau-
city of captures suggests that it enters the Gulf
only as a chance straggler from the north, perhaps
maintaining itself in small numbers in the bottoms
of the deep isolated troughs where the water is
coldest.

The nearest records of it to the eastward and
northward are of fish taken off the Atlantic Coast
of Cape Breton,72 from the estuary of the St.
Lawrence River near Trois Pistoles;73 from St.
Mary's Bay on the south coast of Newfoundland,
from the eastern part of the Grand Banks, and off
the east coast of the Avalon Peninsula, New-
foundland.

Arctic shanny Stichaeus punctatus (Fabricius)
1780

Jordan and Evermann, 1896-1900, p. 2439.

Description. — This shanny suggests the rock eel
in its color pattern. But it is easily distinguished
from it by having well developed ventral fins and
considerably larger pectorals, but fewer dorsal fin

71 Presumably the Gulf of Maine specimens reported by Kendall (Proc.
Portland Soc. Nat. Hist., vol. 3, 1914, p. 62), now in the United States Na-
tional Museum, are this lot.

73 By the Newfoundland Fisheries Research Commission; also it is listed
from Nova Scotia without locality, see Vladykov and McKenzie (Proc.
Nova Scotian Inst. Sci., vol. 19, 1935, p. 104).

'i Vladykov and Tremblay, Natural. Canad. vol. 62 (Ser. 3, vol. 6), 1935,
p. 81.

Figure 261. — Shanny (Leptoclinus maculatus). After Collett.

498

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 262. — Arctic shanny (Slichaeus punclatus), about 7% inches, Halifax, Nova Scotia, redrawn by E. N. Fischer from

U. S. Fish Commission wash drawing.

spines (only 48 or 49). The presence of a series
of large roundish spots on the dorsal fin separates
it at a glance from the radiated shanny, which is
similar to it in form, but which has only a single
large blotch on its dorsal; its spotted dorsal and
its evenly rounded pectorals mark it off from the
shanny; and its much less slender body (only
about one-seventh as deep as it is long) from the
snake blenny.

The single long dorsal fin originates directly
over the edge of the gill cover, and is of uniform
height throughout its length, except that the first
2 or 3 spines, and the last 2 or 3, are shorter than
the others. It extends backward nearly to the
caudal, but the fins are separated by a conspicuous
notch. The anal fin (one short spine, closely
followed by about 35 rays) is about two-thirds as
long as the dorsal fin, is of approximately the
same height as the latter, and bears a similar
relation to the caudal. The caudal fin is gently
rounded in outline. The pectorals are broadly
rounded, a little longer than the depth of the
body; the ventral fins are somewhat less than
half as long as the pectorals. The straight dorsal
profile of the head, from tip of snout to origin of
dorsal fin, is an outstanding feature, and the
lateral line is single (double in the radiated shanny,
p. 499), conspicuous, and ends at about the mid-
length of the body.

Size. — Grows to a length of at least 7 inches.

Color. — The single row of 5 to 9 round black
spots with pale margins on the dorsal fin, and
irregular dark bars on the cheeks and chin are the
most conspicuous markings of this shanny. The
ground color of the body of an Alaskan specimen
is„ described as bright scarlet,74 but is brownish
(after preservation) on a Gulf of Maine specimen,
with darker cloudings extending from close behind
the head to the base of the caudal fin. The lower

» Jordan and Evermann, Bull. 47, U. S. Nat. Mus., Pt. HI, 1898, p. 2439.

surface of the body (except the head) is plain
whitish. The anal fin is dusky, edged with
white; the pectorals and caudal are crossed by
pale bars, and the ventrals are plain yellow.

General range. — Arctic and circumpolar, from
Greenland to northern Siberia and the Arctic
Coast of America; 76 southward to Bristol Bay and
Prince of Wales Island, Alaska, in the Pacific, and
to Mount Desert, Maine on the Atlantic coast of
America.

Occurrence in the Gulf of Maine.— A specimen
4% inches (109 mm.) long, of this Arctic species,
taken one-half mile off Little Duck Island near
Mount Desert, Maine, from the stomach of a cod,
on April 30, 1930, was in such good condition that
it unquestionably had been living in the immediate
vicinity.

The next most southerly records are of two
specimens from Chedabucto Bay, Nova Scotia,
near the Atlantic entrance to the Strait of Canso ; 78
and of one taken off Cheticamp," on the Nova
Scotian shore, within the Gulf of St. Lawrence,
where Huntsman 7S classes it as a characteristic
of the icy-cold water on the banks; and of one
from Battle Harbor, on the outer coast of Lab-
rador, a few miles north of the eastern entrance
to the Strait of Belle Isle.

Radiated shanny Ulvaria subbifurcata
(Storer) 1839

Jordan and Evermann, 1896-1900, p. 2440.

Description.79 — The most conspicuous feature
of this little fish, and one that sets it apart from
all the other blennies of our Gulf, is that the
forward part of its dorsal fin is marked with a

" Reported from Hudson Bay by Vladykov, Contrib. Canadian Biol., N.
Ser., vol. 8, No. 3, 1933, p. 35.
» Cornish, Contrib. Canadian Biol. (1902-1905) 1907, p. 87,
" Cox, Contrib. Canadian Biol. (1918-1920) 1921, p. 112.
" Huntsman, Trans. Roy. Soc. Canada, Ser. 3, vol. 12, Sect. 4, 1918, p. 63.
'• Based on 3 specimens from Grand Man3n, the largest 5?i inches long.

FISHES OF THE GULF OF MAINE

499

Figure 263. — Radiated shanny (Vivaria subbifurcata) . From Jordan and Evermann. Drawing by H. L. Todd.

single large dark blotch. More significant morpho-
logical characters are that it is much stouter
of body (about one-sixth to one-seventh as deep
as it is long) than the snake blenny; that its
pectoral fins and especially its ventrals are con-
siderably larger, relatively, than those of the
rock eel (p. 492), while its ventrals are situated
farther in advance of the pectorals; and that its
gill openings are much wider and extend forward
under the throat (confined to the sides of the
neck in the rock eel). The evenly rounded
outline of its pectorals distinguishes it from the
shanny, in which the lower rays are the longest,
and are separate at their tips. It is a stouter
fish than the Arctic shanny (p. 497), and its lateral
line is double, with an upper branch and a lower,
whereas the lateral line of the Arctic shanny is
single. The outline of the edge of its gill covers,
with the upper corner terminating in a rounded
fleshy flap concealing a sharp angle, is diagnostic
also, for they are rounded in all the other Gulf of
Maine blennies.

The dorsal profile of the head is more convex
than the ventral; the lower branch of the lateral
line runs the whole length of the body, but the
upper branch (the more obvious of the two)
reaches only about as far back as the tips of
the pectoral fins. The dorsal fin, of 43 or 44
spines, is higher, relatively, than that of the rock
eel, and is practically continuous with the caudal
fin. The anal fin, of about 30 rays, is about half
as long as the dorsal, and it is separated from the
caudal by a short but definite interspace, made
obvious by the abrupt rear angle of the anal.
The pectoral fins, evenly rounded in outline,
reach back about as far as the eighth dorsal-fin
spine. The rear margin of the caudal fin is evenly
rounded.

Color. — The most distinctive feature of the color
pattern, one which marks this species among local
blennies, is the presence of a large oval dusky
blotch on the dorsal fin extending from the fifth
or sixth spine to the eighth or tenth spine. The
back and the upper parts of the sides are dull
brown, obscurely barred or blotched alternately
with paler and darker; the sides of the head are
marked with a dark bar running obliquely down-
ward and backward from the eye; and the belly
is pale brownish (described also as yellowish
white) . The caudal fin is crossbarred with 3 or 4
series of dark dots, and the dorsal fin is marked
with many tiny dark dots, besides the blotch
just mentioned.

Size. — The largest one we have seen or read of
is 6% inches long, but the maximum size may well
be larger.

Habits. — Nothing is known of the mode of
life of this shanny except that it is a bottom fish
like other blennies, living among seaweed and
stones from low-tide mark down at least to 30
fathoms, and very likely much deeper. Dr.
Huntsman writes in his notes, "It is found under
stones near low tide mark" with the rock eel but
far less abundantly than the latter and only on
the more exposed shores. Cornish M likewise
describes it as taken under stones on the beach,
as well as in the dredge and trawl in 6 to 30
fathoms at Canso, Nova Scotia.

The eggs have not been seen, but the fact that
we have taken larvae as small as 8 to 11 mm. in
our tow nets in June, July, and October 81 points
to a spawning season lasting from late spring
throughout the summer (if our identification is
correct) .

» Contrib. Canad. Biol. (1902-1905) 1907, p. 87.

" Tln>se are listed in Bull. Mus. of Comp. Zool., vol. 68, No. 2, 1914, p. 109:
and vol. 69, 1917, p. 273.

500

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

General range. — So far this fish is known only
off the boreal coasts of eastern North America,
from eastern Newfoundland, the north shore of
the Gulf of St. Lawrence,82 and from northern
Nova Scotia to Nantucket Shoals and southern
Massachusetts.83

Occurrence in the Gulf of Maine. — This shanny,
first described by Storer from a specimen found at
an unusually low tide among the seaweed at
Nahant on the north shore of Massachusetts Bay
in 1838, was long supposed to be rare. But it has
proved to be common in the Bay of Fundy 84 among
seaweed on rocky shores, a number having been
taken both at Campobello Island and at Grand
Manan, and one in the mouth of the St. Croix
River, while the Museum of Comparative Zoology
has a specimen from Grand Manan, collected by
Verrill many years ago. Huntsman also reports
it from St. Mary Bay on the Nova Scotia shore,

and we have found several in the tide pools at
Nahant, on the northern shore of Massachusetts
Bay. Offshore, we have trawled it near Seguin
Island; in Massachusetts Bay; also on the eastern
part of Georges Bank. And we have found so
many in the stomachs of cod caught on Nantucket
Shoals, on Cashes Ledge, and on other offshore
grounds as to show that it is widespread in the
Gulf on hard bottom, from the shore down to at
least 30 fathoms, while we have had one from as
deep as 45 fathoms. Our tow nets also have taken
its pelagic larvae near Seal Island (Nova Scotia) ;
in the Grand Manan Channel; at the mouth of
Casco Bay; near Cape Porpoise; off the Isles of
Shoals; near Cape Ann ; and in Massachusetts Bay.
The only importance of this little fish in the
natural economy of our Gulf is that many of them
are eaten by cod and by other fishes.

THE WRYMOUTHS. FAMILY CRYPTACANTHODIDAE

The wrymouths are slender, eel-like fishes,
close relatives of the blennies but much larger.
Like the blennies they have a long dorsal fin that
is spiny throughout its length, but the demarcation
between dorsal, caudal, and anal fins is so vague
that they can be described as practically contin-
uous. They have no ventral fins, and their mouth
is so strongly oblique that it is nearly vertical.

" Reparted from Trinity Bay, Newfoundland (Rept. Newfoundland
Fishery Research Comm., vol. 1, No. 4, 1932, p. 109, Sta. 39), and from
Canso, Nova Scotia, by Comish (Contrib. Canadian Biol. (1902-1905) 1907,
p.. 87. And Dr. Vladykov writes us that he has collected a specimen at
Pointe du Maurier on the noi th shore of the Gulf of St. Lawrence.

» Sumner, Osburn, and Cole (Bull. U. S. Bur. Fish., vol. 31, Pt. 2, 1913,
p. 768) record 4 living specimens from Vineyard Sound and others found
in the stomach of a sheldrake (Merganser) that was shot near Robinson
Hole in the Woods Hole region.

M Huntsman, Contrib. Canadian Biol. (1921) 1922. p. 66, and unpublished
notes.

Only three species are known, all North American,
of which the Gulf of Maine harbors one.

Wrymouth Cryptacanthodes maculatus Storer 1839

Congo eel; Bastard cusk; Ghostfish

Jordan and Evermann, 1896-1900, p. 2443.

Description. — The wrymouth is eel-like in form,
about thirteen times as long as it is deep but much
flattened sidewise, and (eel-like) it lacks ventral
fins. And (as in the eel) there is no definite
demarcation between dorsal fin and the caudal fin
or between the anal fin (about 47 to 50 rays) and
the caudal, the one merging into the other to form
a continuous fin around the tail, with no inter-

Figure 264. — Wrymouth (Cryptacanthodes maculatus). From Jordan and Evermann. Drawing by H. L. Todd.

FISHES OF THE GULF OF MAINE

501

spaces. But its dorsal fin (which extends from
close behind the pectorals back to the caudal fin)
is spiny (about 75 to 77 spines) for its entire length
like that of its close relatives the blennies. But
the absence of ventral fins separates it from all of
our local blennies, and its peculiar profile is an
equally useful field mark, the head being flat-
topped, the eyes set high up in very prominent
orbits, and the mouth strongly oblique with so
heavy a lower jaw that it gives the face a bulldog-
like expression when the mouth is closed. The
wide gill openings, running forward under the
throat, and the small size and rounded outline of
the pectorals are distinctive, also, as is the fact that
both the dorsal fin and the anal fin are low (less
than half as high as the body is deep in large
specimens, relatively higher in small), and of
uniform heights throughout most of their lengths,
with the anal about two-thirds as long as the dor-
sal. The caudal fin is oval.

Color. — Described (and the few preserved speci-
mens we have seen correspond with this) as of
varying shades of brown or reddish brown, with
the upper part of the sides marked with two or
three irregular rows of small darker brown spots
that run from head to tail; the top of the head is
thickly speckled; the dorsal and anal fins are
spotted with similar but smaller dots, and the belly
is grayish white. A few spotless specimens have
been seen.

Size. — Maximum length about 3 feet.

Habits. — Very little was known of the habits of
the wrymouth until recently, except that it is a
bottom fish living from the intertidal zone down to
considerable depths (where it is sometimes taken
on line trawls in the Bay of Fundy). But in 1910
and again in 1920 Willey and Huntsman 85 found
full-grown wTymouths living in burrows in the mud
on the flats at the mouth of the Magaguadavic
River, a tributary of Passamaquoddy Bay. These
burrows, to quote from their account, "were found
in very soft mud from the lower part of the Fucus
zone downward; that is, as far up as 4 feet above
low-water mark," and "each system of burrows,
inhabited by only one fish, consisted of branching
tunnels about 5 cm. in diameter and from 3 to 8 cm.
below the surface" of the mud, originating from
a more or less centrally placed mound, where the

main entrance was, with other smaller openings
along the tunnels and at their terminations.

It seems that the burrowing instinct is strong,
for one fish kept in a tank constantly inhabited a
piece of hard rubber tubing. Hence it is probable
that wrymouths in other parts of the Gulf likewise
live in burrows or perhaps under stones. And
they seem as likely to be inshore in shoal water
in winter as in summer, for one was speared in
Marblehead Harbor in December many years
ago.86 Within our Gulf wrymouths have been
found from a little above low water mark, as just
remarked, down to about 100 fathoms; and to
somewhere between 245 and 325 fathoms off New
Jersey (see footnote 89, p. 502).

Huntsman and Willey found "beach fleas" or
"sand-hoppers" (Oammarus), shrimps (Crago) and
fragments of winter flounders in several wry-
mouths which they opened, and the one kept in
captivity ate sand-hoppers, hermit crabs, small
herring, and mollusks such as limpets, peri-
winkles, whelks, clams, and mussels. Apparently
it located food as much by sight as by smell.87

Ripe wry mouths are yet to be seen; but the
presence of the larvae early in spring in Passama-
quoddy Bay, as reported by Huntsman, with the
seasonal occurrence of the fry mentioned below
(p. 502), proves it a winter spawner in the Gulf of
Maine. It may breed later in the Gulf of St.
Lawrence, for Dannevig M records a young wry-
mouth only 38 mm. long that was taken there as
late as June 10. The localities where the young
fish have been taken (see p. 502) suggest that wry-
mouths spawn all around the coast of the Gulf of
Maine and wherever they occur on the offshore
banks.

Neither the eggs nor the early larval stages are
known. But by the time the young have grown
to a length of 21 to 22 mm. they show the long
dorsal and anal fins, and the lack of ventral fins
characteristic of their parents, though they are
much less slender, relatively, their caudal fins are
larger and square instead of rounded and their
mouths are nearly horizontal. The pigmentation
of the fry is likewise extremely characteristic, the
upper sides from the eye back to the caudal fin

« Canad. Field Natural., vol. 35, 1921, p. 4.

" Putnam, Bull. Essex Inst., vol. 6, 1874, pp. 11-13.

17 Willey and Huntsman also give interesting data on its respiration and
on its response to various stimuli.

" Canadian Fisheries Expedition, 1914-1915 (1919), p. 16. He gives an
excellent figure of this specimen on pi. 2, fig. 10.

502

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

being thickly speckled with dark brown dots,
which are sparser on the lower part of the sides.

General Range. — Atlantic Coast of North Amer-
ica, from southeastern Labrador, the coasts and
banks of Newfoundland, and the Gulf of St.
Lawrence to Long Island Sound and to the offing
of central New Jersey.89

Occurrence in the Gulf of Maine. — Published
records locate this fish in the Bay of Fundy; at
Eastport; in Casco Bay; at Portland; in the mouth
of the Piscataqua River; at Gloucester; in Marble-
head Harbor; at Swampscott; Nahant; and Dor-
chester in Boston Harbor; and in the outer waters
of Massachusetts Bay; there are specimens in the
Museum of Comparative Zoology from Trenton,
Maine; from outer Boston Harbor; and from near
Provincetown. Two were taken in the central
basin of the Gulf in July 1931 at a depth of 88-95
fathoms; one was trawled by the Atlantis in the
deep trough west of Jeffreys Ledge at 72-78
fathoms, and another in the southwestern basin
of the Gulf off Cape Cod at about 100 fathoms
(183 meters), in August 1936; the Albatross II
trawled one on the eastern slope of Nantucket
Shoals 90 at 52 fathoms, in May 1950. And one
of the crew of the dragger Eugene H reports the
capture of 4 of them on the northeastern part of
Georges Bank on October 12, 1951. We have
also taken its late larvae and fry in tow nets (11
specimens 18 to 40 mm. long) in Massachusetts

Bay off Boston Harbor; over JefferysBankjin the
trough near the Isles of Shoals; in the western
basin a few miles west of Cashes Ledge; off
Penobscot Bay; near Mount Desert Island; and
in the deep basin off Machias, Maine, in May
1915, and in March and April 1920. These
localities are sufficiently scattered to show that it
is to be found, not only all around the coasts of
the Gidf , but on the offshore grounds as well.

However, it seems to be rare or at least very
local, for we have caught few adults ourselves,
nor have we seen it brought in by fishing boats.
In fact, few of the fishermen of whom we have
inquired have been aware of its existence, a fact
no doubt associated with its burrowing habit.
And it has not been reported as yet from Browns
Bank though it is to be expected there.

Following its range to the eastward and north-
ward, we find it described as "rather common"
all along Nova Scotia (taken at 60 fathoms on
Western Bank off Halifax) ;91 it has been reported
from a number of stations on the eastern half of
the Grand Banks region, from the southern and
southeastern coasts of Newfoundland, and from
the outer Labrador coast some 20 miles north of
the Strait of Belle Isle (see footnote 89, p. 502);
also within the Gulf of St. Lawrence from the
Bay of Chaleur in the southwest,92 and from the
entrance to the Strait of Belle Isle in the north-
east.93

THE WOLFFISHES. FAMILY ANARHICHADIDAE

The wolffishes are closely allied to the blennies,
and like the latter they have a single long spiny
dorsal fin running the whole length of the back
from the nape of the neck. But the presence of
large molar teeth and canine tusks, with their
total lack of ventral fins and the fact that all but
the last 10 or 12 of their dorsal fin spines are soft
and flexible at the tips, instead of stiff justify a
separate family for them. They are much larger
fish than any of our blenny tribe. Two species

M The most northern locality-record which we have found is for its drifting
larvae off the outer coast of Labrador, about 20 miles north of Belle Isle
(Rept. Newfoundland Fish. Res. Lab., vol. 2, No. 3, 1035, p. 70, Sta. 422);
the most southern are for one trawled by the Albatross II off northern New
Jersey, lat. 40°04' N., long. 73°32' W„ August 1036, at 35 fathoms; and of
another dredged by the Atlantis 30 miles farther south Oat. 30°31' N., long.
72°10' W.) between 245 and 325 fathoms, that same year.

•• Lat. 40°05' N., long. 69°22' W.

occur in the Gulf of Maine, one, the wolffish,
commonly; the other, the spotted wolffish, only
as a stray from the north. Another cold water
species, Anarhichas latifrons, has been recorded
repeatedly from Nova Scotian waters, hence it is
to be expected in our Gulf sooner or later though
it has not been reported there definitely. It is
included in the following Key.94

•' Vladykov and McKenzie, Proc. Nova Scotian Inst. Sci., vol. 19, 1935,
p. 104.

" Halkett, Checklist Fishes Canada, 1913, p. 112, "Gaspe Bay."

« Rept. Newfoundland Fish. Res. Lab., vol. 2, No. 3, 1935, p. 79, Sta. 370.

•« We think It likely that a specimen of A. latifrons credited by Bean (Proc.
TJ. S. Nat. Mus., vol. 3, 1881, p. 82) to "east coast of United States" was
actually brought In from Nova Scotian waters rat her than I hat it was caught
anywhere west of Cape Sable. Seo Blgelow and Schroeder (Proc. Boston
Soe. Nat. Hist., vol. 41, No. 2, 1035, p. 15) for further remarks on this species.

FISHES OF THE GULF OF MAINE

503

KEY TO GULF OF MAINE AND NOVA
SCOTIAN WOLFFISHES

1. Back and sides definitely black spotted on a pale

ground; the bands of molar teeth in the roof of the
mouth are all of about equal lengths.

Spotted wolffish, p. 507

Back and sides plain colored or dark barred and

blotched, but not definitely black-spotted; the

central band of molar teeth in the roof of the mouth

is longer than the bands that flank it 2

2. The central band of molar teeth originates at about

the same level as the bands on either side of it, but
it extends considerably farther rearward than they
do; the canine teeth are very large and prominent;

the flesh is firm Wolffish, p. 503

The central band of molar teeth originates consider-
ably in advance of the bands on either side of it,
but it terminates about even with them rearward;
the canine teeth are small, not very prominent; the
flesh is noticeably flabby when fresh-caught.

Arctic wolffish, Anarhichas latifrons,
Steenstrup and Hallgrimsson.

Wolffish Anarhichas lupus Linnaeus 1758
Catfish; Ocean whitefish

Jordan and Evermann, 1896-1900, p. 2446.

Description. — The wolffish suggests a huge
blenny in its general make-up, except that its
dorsal fin spines are flexible at their tips instead of
stiff; that it has no ventral fins; and that its mouth
is armed with a set of teeth more formidable than
those of any other Gulf of Maine fishes, except for
its relative, the spotted wolffish (p. 507), and for
some of the sharks. There is a row of about 6 very
large, stout, conical canine tusks with a cluster of 5
or 6 smaller canines behind them in the upper jaw;
and the roof of the mouth back of the latter is
armed with three series of crushing teeth. The
central series of these consists of a double row of
about 4 pairs of large rounded molars that are

united into a solid plate; each of the outer series
consists of two alternating rows of blunt conical
teeth. In the common wolffish the central series,
which is the longest of the three, originates a very
little in advance of the outer series, and it extends
rearward noticeably farther. The lower jaw
has 4 to 6 large tusks in front, behind which are
two longitudinal diverging rows of rounded molars.
And the throat also is armed with small scattered
teeth. The great projecting tusks, rounded nose,
and small eyes give the wolf a singularly savage
aspect.

The body is deepest close behind the head,
tapering back to a slender caudal peduncle and to
a small weak tail fin. The dorsal fin (69-77
spines) is about half as high as the head is long and
uniform in height from end to end except for its
rounded corners, and it extends from the nape of
the neck to the base of the caudal fin. The anal
fin (42-48 rays) is only about half as high as the
dorsal, and a little more than half as long; its rear
corner is angular. The pectoral fins are large and
rounded, and the caudal fin is slightly convex in
outline.

Color. — Wolffishes are dull-colored, but they
vary widely in tint. The upper parts and the
dorsal fins of those taken off the Massachusetts
coast have been described as purplish brown and
we have seen them of this tint. But fish caught on
Georges Bank are invariably dull olive green,
according to Mr. Clapp, while they are described
as purplish, brownish, or bluish gray, or slate
colored in other seas. No doubt the color of the
wolf, like that of many other ground fish, varies
with that of its surroundings, purplish and brown
tints ruling among red seaweeds and olive gray on
clean bottom. Whatever its tint, its sides are
transversely barred with a variable number
(usually 10 or more) of irregular and broken

Figure 265. — Wolffish (Anarhichas lupus), Georges Bank. From Goode. Drawing by H. L. Todd.

504

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

darker bands or blotches, or scattered spots, that
extend out on the dorsal fin. The throat and the
belly back to the vent are dirty-white tinged with
the general ground tint of the upper parts. Wolf-
fish fade so soon after they are caught that those
seen in the markets usually are much paler than
they were in life.

Size. — A length of 5 feet seems about the maxi-
mum in Gulf of Maine waters; one more than 4
feet long is seldom seen, and the larger fish caught
and brought in run less than 3 feet. European
authors speak of wolffish of 6 feet and even longer,
but they average only about 2 feet in Scandinavian
waters,95 i. e., scarcely so large as those in the Gulf
of Maine. A fish 33 inches long weighs about 10
pounds, one of 37 inches between 16 and 17
pounds. The greatest weight reported for Ameri-
can waters so far is about 40 pounds.96

Habits. — The wolffish is solitary, living one
here and one there, and it is not abundant any-
where, in the sense that this term can be applied
to the cod, to the haddock, to the pollock, or others
of our commercially important fishes. It holds
close to the bottom; and it is always caught on
hard ground, never on mud, a preference illus-
trated by the fact that our experimental trawlings
on the soft bottom of the deep troughs within the
Gulf did not catch one wolffish, though they did
yield a variety of other fishes in plenty.97 It is a
weak swimmer, moving by sinuous side to side
undulations like a blenny or an eel; and probably
it spends most of its life hidden among seaweed or
rocks, or nosing about such surroundings for food.
There is no reason to suppose that it ever attacks
other fish in its normal way of life, but when
hauled out of the water it snaps like a bulldog
and with good aim at anything in its way, the
hands, an oar, or at other fish among which it is
thrown, and it can inflict a serious bite. Goode M
remarks that it has been known to make a furious
attack on persons wading among the rock pools at
Eastport, Maine.

The depth zone occupied by the wolffish at one
time or another extends from a fathom or so below
tide mark down to 85 fathoms at least, and very
likely deeper. It has been reported in tide pools
at Eastport, but we have never heard of it in such

" Smitt, Scandinavian Fishes, vol. 1, 1892, p. 232.
« Ooode, Fish. Ind. U. S. Sect. 1, 1884, p. 249.

•» For list of species taken, see Blgelow and Schroeder, Biol. Bull. vol. 76,
1939, p. 309.
» Fish. Ind. U. S. Sect. 1, 1884, p. 249.

situations or at low-water mark anywhere else
in the Gulf, nor does it run up estuaries, and it is
probable that most of the local stock lives in depths
of 10 to 50 or 60 fathoms.

The wolffish is a cool- and cold-water fish, as
might be assumed from the fact that its regular
geographic range extends hardly farther west than
Cape Cod and Nantucket shoals. Those living in
the coastal belt of our Gulf, at depths of 25 fathoms
or less, regularly experience temperatures as low as
34°-36° (locally even as low as 32°) at the end of
the winter, or at some time during the spring,
according to locality.99 They are in temperatures
equally low or even lower, fractionally, in late
spring and early summer on the fishing grounds
along outer Nova Scotia, while the grounds where
they are caught in the Gulf of St. Lawrence are
flooded every spring and early summer, with water
as cold as 32°, which they can avoid only by de-
scending deeper into the Laurentian Trough, a
movement of which we have no direct evidence.
And they have been caught, widespread, on the
Newfoundland Banks (p. 507) in water as cold as
30°-31°. At the other extreme, the highest tem-
perature in which wolffish occur in any numbers is
about 50°-52°, at the end of summer (again for
those living shoalest) in the coastal belt of our
Gulf, and also on the Nantucket shoals grounds.
They are never known to run up into brackish
water.

The wolffish is resident wherever it is found, to
be caught throughout the year. For example,
about as many are brought in from Georges Bank
in one month as in another, allowance being made
for the difficulties and dangers of winter fishing.
And as it passes through only a brief pelagic stage
when it is young (p. 506), it is a comparatively
stationary fish, with much less interchange from
one locality to another than is the case with cod or
with haddock.

The diet of the wolffish consists wholly of hard-
shelled mollusks, crustaceans, and echinoderms.
So far as we can learn fish have never been found
in the stomach of a wolffish. Mr. Clapp found
that the 50 or 60 fish that he opened on Georges
Bank had all eaten large whelks (Buccinum),
cockles (Polynices, Chrysodomus and Sipho), sea
clams (Mactra), and other shellfish, which it
crushes easily in its viselike molars. Sometimes,

« For further details, see Blgelow, Bull. U. S. Bur. Fish., vol. 40, part 2,
1927, p. 542.

FISHES OF THE GULF OF MAINE

505

however, mollusks even as large as these are
swallowed whole, and we have seen many caught
on hooks baited with clams. In north European
waters wolffish are said to subsist largely on
mussels, and one examined by Vinal Edwards at
Woods Hole was full of these; none, however, were
found in stomachs of the fish caught on Georges
Bank, although mussels are plentiful there, which
points to a definite preference for the other shellfish
just mentioned. The wolffish is also known to feed
on large hermit crabs, on ordinary crabs, and other
crustaceans, on starfish, and on sea urchins, a
quart of the latter having been taken from one
caught at Eastport.1 And Mr. Clapp's observa-
tions that every one he has opened contained food
of some sort is good evidence of its constant search
for anything edible. With such a diet it is not
surprising that wolffish are more often caught on
hand lines baited with cockles or clams than on
long lines, which are usually baited with herring.

Breeding habits.- — The breeding habits of the
wolffish have not been followed on this side of the
Atlantic. In north European waters it spawns
chiefly from November until January,2 and ap-
parently the breeding season is about the same
for it in Nova Scotian waters, and in the Gulf of
Maine, for McKenzie and Homans 3 report a mass
of eggs dragged up on February 19, in 1937,
some of them just hatching, while we have taken
larvae of 20 to 22 mm. (fig. 267), that is, 2 to 3
months old from the time the eggs were deposited,
as early as January 30 in 1913, and as late as
March 4 in 1920.

The eggs, 5.5 to 6 mm. in diameter (among the
largest fish eggs known), yellowish, opaque, and
with an oil globule of 1.75 mm., are laid on the
bottom in shoal water where they stick together
in large loose clumps among weeds and stones.
The fish have been described as making an annual
shoreward journey for spawning purposes, but
there is little evidence of this. The precise dura-
tion of incubation is yet to be learned; probably it
is long, as it is for most of the fishes that lay their
eggs on the bottom.

• Verrlll, Amer. Naturalist, vol. 6, 1871, p. 400.

» It was formerly thought to spawn in spring, but Mcintosh and Prince
(Trans. Roy. Soc. Edinburgh, vol. 35, Pt. 3, No. 19, 1890), to whom we owe
all that Is known of its early larval development, proved it an autumn and
winter spawner both by examination of Its ovaries and by the discovery of
its eggs.

> Proc. Nova Scotian Inst. Scl., vol. 19, 1938, p. 279.
210941—53 88

Figure 266. — Larva (European), just hatched. After
Ehrenbaum.

Figure 267. — Larva, 21.5 mm. Gulf of Maine.
Wolffish (Anarhichas lupus).

The slender transparent larvae of the wolffish
of north Europe are described as about half an
inch (12 mm.) long at hatching, but those that
hatched from a mass of eggs dragged up off Nova
Scotia were 17-18 mm. long.4 They have an
enormous baglike yolk sac enclosed in a net of
highly developed blood vessels (see fig. 266, p.
505) , and thus they suggest salmon or trout larvae
remotely, in appearance. Wolffish hatched by
Mcintosh and Prince in the aquarium at St.
Andrews, Scotland, did not absorb the yolk sac
wholly until about 3^ months old and upward of
20 mm. long, but in natural surroundings larvae
as small as 17 mm. have been found free of yolk,
nor was any trace of it visible in the larvae of 21
mm. and upward which we have towed in the
Gulf of Maine. Larvae of 20 to 22 mm. show the
dorsal and anal fin rays in their final number, but
the large head, enormous eyes, and tiny teeth,
with the fact that there is no definite separation
between the anal and dorsal fins and the caudal,
give the young fishes an aspect very different from
that of their parents until they are 1% to 1% inches
long. In life the wolffish is silvery on the sides at
this stage, but this metallic hue fades after preser-
vation, leaving only the dark brown pigment
granules with which the sides are thickly dotted.
The largest fry we have seen (44 mm. long) are
similarly pigmented but somewhat paler.

• McKenzie and Homans, Proc. Nova Scotian Inst. Scl., vol. 19, 1938,

p. 279.

506

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

When first hatched, the larvae lie on bottom
like young trout or salmon, resting on the yolk; it
is not until the latter is considerably reduced in
size (several weeks after hatching) that they swim
much. And they do no more than dart upward
for a few inches and then settle back again until
a month old. Thus the wolffish spends the early
part of its life close to bottom instead of drifting
at the mercy of tide and current, as all fishes do
that produce buoyant eggs. Some of the older
larvae and the young fry adopt a drifting habit
for a time after the yolk is absorbed (we took some
20 specimens of various lengths from 21 to 44 mm.,
in our tow nets during March and April 1920).
But it seems that they seldom rise to the upper-
most water layers, for only 2 of the 8 hauls that
took them were made at the surface, the others
were at depths of 30 to 60 fathoms. And as fry
no larger than this have been trawled on bottom
in European waters, it seems that some do not
leave the ground. It follows, then that the
wolffish probably is not subject to the long in-
voluntary migrations that are carried out by most
of the members of the cod and flatfish tribes, but
that it passes through its entire larval stage near
where it is hatched, hence localities where the
young are taken are evidence of local spawning.
The brevity of the pelagic stage, if any, also im-
plies that the stock of wolffish in any given
locality depends on local reproduction for its
maintenance.

In Scottish waters wolffish fry have been taken
as long as 5 to 6 inches in July, and up to 7 to 8
inches in August, pointing to a rapid rate of growth
for the first summer. Nothing is known of the
later growth.

General range. — Both sides of the North At-
lantic; north to Davis Strait in American waters;
south regulaiTy to Cape Cod; less often to the
westward along southern New England, and
exceptionally to New Jersey;6 also Greenland;
Iceland; and northern Europe southward to
northern France.

Occurrence in the Gulf of Maine. — Although the
wolffish has been recorded at only a few stations

in the Gulf in scientific literature 6 it is a fairly
common fish, to be caught on suitable bottom
in all parts of the Gulf, though nowhere in any
great numbers. Thus, something like 30,000
pounds were taken off western Nova Scotia yearly
during the period 1944-1946; 7 scattering fish are
caught at the mouth of the Bay of Fundy and
around Grand Manan (stray fish only up the Bay,
if any); on one occasion, in April 1930, we saw
35 of them, 2 to 3 feet long, caught on one set of a
long line in 18 fathoms of water off Mount Desert
Island; and enough are caught on the small
grounds thence eastward to Eastport for 16,000
pounds to have been reported from this statis-
tical area in 1945. The various small coastwise
fishing grounds, westward from Mt. Desert,
yield much greater numbers, as reflected in
reported landings, for 1945, of 253,000 pounds for
central Maine; about 1,400 pounds for Cashes
Ledge and the neighboring patches of hard bot-
tom, where we have caught them, as we have on
Platts Bank also; about 26,000 pounds for western
Maine; and about 118,000 pounds reported as
taken by the vessel fishery off the coast of Massa-
chusetts from the New Hampshire fine to Cape
Cod.

Recent statistical surveys have not been of a
sort to localize the catches more precisely than
this. But the small-boat fishermen, landed more
than 37,000 pounds in Essex County, Mass., in
1905, mostly from Jeffreys Ledge, from Stellwagen
Bank, and from the deeper rocky spots near
Gloucester and Nahant, grounds where fishermen
report them as fairly plentiful. And there is no
reason to suppose that the situation in this respect
has altered subsequently.

Considerable catches are also brought in from
Cape Cod waters, as reflected in landings of
233,000 pounds reported for Barnstable County
in 1945, about 224,000 pounds for Cape Cod
in 1947.8

Wolffish are not taken in commercial quantities
either to the westward of Nantucket Shoals, or

• Abbott (Oeol. New Jersey, 1868, p. 818) characterized It as "not unfre-
quently met with" of! the New Jersey coast, and Schnakenbeck (Faune
Ichth.; Cons. Perm. Intemat. Explor. mer. PI. not numbered, 1933) even
outlined Its range as extending southward to Cape Hatteras. But we have
heard of none caught to the westward and southward of Vineyard Sound at
any time during the past half-century.

• The deep channel between Georges Bank and Browns; off Cape Sable;
in St. Mary Bay; at Grand Manan; at Campobello, at the mouth of the Bay
of Fundy; Eastport; Mussel Ridge Channel; Casco Bay; Ipswich Bay;
Annisquam; off Gloucester; Massachusetts Bay; North Truro; Nantucket
Shoals, and Georges Bank.

7 Canadian and United States catches combined.

' We cannot localize these any more precisely.

FISHES OF THE GULF OF MAINE

507

on the southwestern part of Georges Bank.' But
the so-called "south channel" grounds from
Cape Cod out to the northwestern part of Georges,
with the northern and eastern parts of Georges,
and Browns Bank to the eastward, support so
large a population that these grounds, combined,
yielded about 368,000 pounds in 1945, about
840,000 pounds in 1947.10 And it is much
more likely that the differences from year to
year in the catch n are due to the fact that
wolffish are taken only incidentally, so that the
catch depends on the precise grounds fished,
rather than on any periodic alterations in their
abundance.

In 1946 (most recent year for which we have
seen statistics for the Canadian as well as for the
United States catches) something like 1,571,500
pounds of wolffish were reported as caught within
the limits of our Gulf, or something like 260,000
to 270,000 fish, assuming an average weight of
5 to 7 pounds. But it is anyone's guess what
proportion of the total population this may be.

Wolffish appear to be about as numerous od
the various fishing grounds along outer Nova
Scotia (reported catch for 1949, about 800,000
pounds) as they are on Georges and Browns Banks,
(p. 507). But while they are reported at several
localities in the southern side of the Gulf of St.
Lawrence, 12 also off the west coast of Newfound-
land, and at Anticosti, they are not plentiful
enough anywhere in the Gulf to yield commercial
catches. '* And this applies equally to the New-
foundland Banks, though they have been taken
widespread there; also along the south and south-
east coast of Newfoundland, and as far north as
the offing of Hamilton Inlet on the outer Labrador
coast, during the experimental trawlings carried
out by the Fishery Research Board of New-
foundland.

The fact that we have taken wolffish larvae in
the channel between Browns Bank and Cape
Sable ; near Seal Island (Nova Scotia) ; on German
Bank and off its slope; off Lurcher Shoal; off

• Reported landings were about 6,000 pounds for Nantucket Shoals and
about 13.000 pounds for the southwestern part of Georges Bank for 1945;
about 9,000 pounds and 23,000 pounds respectively for these same grounds
in 1947.

io Weight of dressed fish.

" For earlier examples, see Bigelow and Welsh, Bull. U. S. Bur. Fish., vol.
40, Ft. 1, 1925, p. 373.

u Cheticamp; Prince Edward Island; also Gaspe Bay.

11 They are not mentioned in the catch statistics for the Gulf of St. Lawrence
coast of Nova Scotia, for New Brunswick or for Quebec.

Machias (Maine) ; on Jeffreys Bank (off Penobscot
Bay); and in Massachusetts Bay a few miles off
Gloucester, is evidence that the wolffish breeds in
the Gulf wherever it is to be found, as might,
indeed, be expected. And this applies, equally
(it seems) to the more northern parts of the Ameri-
can range of the wolffish, for its pelagic young
have been reported off northeastern Newfound-
land; in the Strait of Belle Isle; and off Sandwich
Bay on the Atlantic coast of Labrador, by the
Newfoundland Fisheries Research Commission.

Commercial importance.- — The market demand
for wolffish is of comparatively recent growth.
It is an excellent table fish, selling readily as
"ocean catfish" or as "whitefish." In 1947 the
average price to the fisherman was between 4 and
5 cents per pound, and the Gulf of Maine catch
was worth about $70,000 to $80,000.

Spotted wolffish Anarhichas minor Olafsen 1774u
Spotted catfish
Jordan and Evermann, 1896-1900, p. 2446.

Description.- — This species resembles the com-
mon wolffish closely in its general form and in
the arrangement of its fins. The chief difference
is that while the central ("vomerine") band
of teeth on the roof of the mouth is longer than
the band on either side ("palatine") in the
common wolffish, these bands are of about equal
lengths in the spotted wolffish, and its teeth are
described as red in life, not white. Furthermore,
the rear end of its dorsal fin is abruptly indented
close to the base of the tail, with its last 3 to 6
spines much shorter than those further forward,
while the rear end of the dorsal is evenly rounded
in the common wolffish.

Color is, however, the most convenient field
mark for the spotted wolffish, its pale olive or
chocolate 16 upper parts together with its dorsal
and caudal fins, being thickly sprinkled with
blackish brown spots, of different sizes and of
irregular shapes.

Size. — Notwithstanding its Latin name this is
fully as large a fish as the common wolffish, said
to grow to a length of 6 feet. One 37 inches long
weighed 13 pounds, eviscerated.

H Jordan, Evermann and Clark (Rept. U. S. Comm. Fish. (1928), Ft. 2,
1930, p. 472) place this species in the genus Lucichlhys Gil] 1876; but it seems
preferable to follow the older usage here.

i' The general ground tint has been variously described.

508

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 268. — Spotted wolffish (Anarhickas minor), off La Have Bank. From Goode. Drawing by H. L. Todd.

Habits. — Very little is known of its habits. Its
diet is much the same as that of its more common
relative. But it is said to keep to rather deeper
waters, having been caught as deep as 200 to 240
fathoms off Banquereau Bank 16, to 200 fathoms
in the eastern Atlantic. And its geographic range
(see below) shows that it is confined to colder
water than is the common wolffish.

General range.- — Chiefly north of the Arctic
circle; north coast of Russia, White and Barents
Seas, and Iceland, south to middle Norway (vicin-
ity of Bergen) on the European coast ; Greenland ;
and southward occasionally to the Gulf of Maine
on the American coast.

Occurrence in the Gulf of Maine.- — Goode and
Bean's 17 statement that "the Fish Commission
has specimens from off the mouth of Gloucester
Harbor and from Eastport, Maine," long remained
the only notice of this northern fish for the Gulf
of Maine,18 and fishermen of whom we have in-
quired have either never seen it there or they
have failed to distinguish it from the common
wolffish, which is unlikely, so striking is its color
pattern. But the late Walter Rich, of the U. S.
Bureau of Fisheries, obtained a specimen that
had been taken in 35 fathoms off Cape Elizabeth
(now in the collection of the Portland Society of
Natural History); another, weighing 3% pounds

was caught on a long line off Portland lightship
on April 23, 1927. Evidently this wolffish reaches
our Gulf only as an accidental waif from its
Arctic home, one to be watched for but hardly
to be expected.

It appears to occur regularly off outer Nova
Scotia however, though in small numbers. Thus
5 to 10 are usually landed at Boston each year
from Sable Island Bank; we have records of 7
caught there in 1934;19 one was brought in many
years ago from 200 fathoms from the deep gulley
between Sable Island Bank and Banquereau; and
one from 250 fathoms from the northeast slope of
the latter bank.20 Another was taken on Ban-
quereau in 50 fathoms, in 1932 ;21 three were taken
on Western Bank in 1937;22 five more were
definitely reported from Banquereau in that
same year; and McKenzie 23 writes that half a
dozen are brought in from that general region
yearly.

With so many records for Nova Scotian waters
it is astonishing that the spotted wolffish has not
been reported at all in the Gulf of St. Lawrence
so far as we can learn, nor is it mentioned among
the fishes listed in Newfoundland waters or off
outer Labrador from the experimental trawlings
by the Fishery Research Commission of New-
foundland.

THE OCEAN POUTS AND WOLF EELS. FAMILY ZOARCIDAE

The ocean pouts and wolf eels are slender
eel-like fishes with the anal fin continuous with
the caudal. In most of the members of the family
the dorsal fin, also, joins the caudal equally,
making one continuous fin extending around the
tip of the tail, but in the only common Gulf of
Maine species the rear portion of the dorsal is so

>• Bean, Proc. U. S. Nat. Mus., vol. 3, 1881, p. 82.
" Bull. Essex Inst., vol. 11, 1879, p. 11.

» Goode's (Fish. Ind. U. S., Sect. 1, 1884, p. 249) statement that it has
been seen in the Bay of Fundy apparently refers to this Eastport record.

low that there seems to be a bare space between
it and the caudal. All the members of the family,
however, known definitely, either from the Gulf
of Maine, or from the outer coast of Nova Scotia,
are readily separable from the true eels by having
ventral fins (small but unmistakable) situated a

'» Blgelow and Schroeder, Bull. U. S. Bur. Fish., vol. 48, 1936, p. 337.
» Bean, Proc. V. 8. Nat. Mus., vol. 3, 1881, p. 82.
>' Vladykov, Proc. Nova Scotian Inst. Scl., vol. 19, pt. 1, 1935, p. 3.
» McKenzie and Homans, Proc. Nova Scotian Inst. Scl., vol. 19, 1938,
p. 279.
» Proc. Nova Scotian Inst. Scl. vol. 20, pt. 1, 1939, p. 18.

FISHES OF THE GTXLF OF MAINE

509

little in advance of the pectorals. But the green
ocean pout (Gymnelis viridis Goode and Bean) of
arctic seas has no ventrals; it ranges southward as
far as the estuary of the St. Lawrence River,24
and perhaps as far as northern Nova Scotia.25
The closest affinities of the ocean pouts, among
Gulf of Maine fishes are with the blennies (p.
491), the wolffishes (p. 500), and the wrymouths
(p. 502). But they are easily separable from the
blennies and wrymouths by the fact that at least
the major part of the dorsal fin is soft-rayed, not
spiny; and from the wolffish by their more slender
form and smaller teeth.

Only two species are known definitely from our
Gulf, one, the common ocean pout (p. 510) very
plentiful; the other, the wolf eel (p. 515), much
less so. A third, the Arctic Ocean pout (p. 516) has
been reported from shoal water to the west of our
Gulf as well as from the Nova Scotian Banks to
the east, though not from the Gulf itself. A
fourth species {Lycodes esmarkii Collett 1875) has
been credited to the Bay of Fundy.26 But the
specimen in question was trawled by the Albatross
I, on the southern slope of the Grand Banks in 244

fathoms of water.27 Two others that have been re-
ported from the Nova Scotian Banks are also in-
cluded in the following key, as they are likely to
be found in our Gulf sooner or later.

Still another species, Lycodes atlanticus Jensen
1904, has been reported from a number of stations
along the continental slope from the offing of
southern Nova Scotia to the offing of northern
North Carolina, in depths of 543 to 1,423 fath-
oms.28 But being a deep-water form, it is not to
be expected either within the limits of the Gulf of
Maine, or on the Nova Scotian Banks.

The various species of the genus Lycodes
resemble one another so closely that their identi-
fication is very difficult. If one should be taken
in the Gulf that does not agree with either of these
that are described on the following pages, we
suggest that it be sent either to the laboratory of
the U. S. Fish and Wildlife Service, Woods Hole,
Mass.; the Division of Fishes, U. S. National
Museum, Washington, D. C. ; or to the Department
of Fishes, Museum of Comparative Zoology,
Cambridge, Mass., to be named.29

KEY TO GULF OF MAINE AND NOVA SCOTIAN OCEAN POUTS AND WOLF EELS

l.

3.

The dorsal fin seems to be separated from the caudal fin by a considerable gap Ocean pout, p. 510

The dorsal, caudal and anal fins form one continuous fin 2

The trunk is extremely slender, at least 14-16 times as long as it is deep; the dorsal fin originates over the tips of the

pectoral fins 3

The trunk is stouter, less than 12 times as long as it is deep 4

Dorsal fin with only about 92 rays and anal fin with about 88 rays; lower surface of body with only a few scales.

Wolf eel, p. 515

Dorsal fin with about 118 rays and anal fin with about 110 rays; lower surface of body uniformly scaly, like upper

surface Lycenchelys paxillus, Goode and Bean 1879 ^

The lateral line runs along the middle of the sides; the vent is only a little nearer to the snout than to the tip of the

tail; there are no scales on the belly or on the forward part of the back Arctic eelpout, p. 516

The lateral lines (or their lower branch if double) run along the lower part of the sides; the vent is considerably

nearer to the snout than it is to the tip of the tail ; the body is covered with scales 5

» Vladykov and Tremblay, Natural. Canad., vol. 62 (Ser. 3, vol. 6), 1935,
p. 82.

» Goode and Bean (Smithsonian Contrib. Knowl., vol. 30, 1895, p. 313)
credit It to Nova Scotia but give no definite locality.

» By Vladykov and McKenzie, Proc. Nova Scotian Inst. Sci., vol. 19, 1935,
p. 109.

■ Reported by Qoode and Bean (Smithsonian Contrib. Knowl., vol. 30,
1895, p. 305) as from lat. 44°47' N.; long. 66°33' W., Albatross Station 2470; but
the actual position o( this station was lat. 44°47' N.; long. 56°33' W. See
Townsend, Kept. U. S. Comm. Fish., (1900), 1901, p. 399.

" Reported by Qoode and Bean (Smithsonian Contrib. Knowl., vol. 30,
1895, pp. 305-306) as L.frigidus Collett 1875, with list of stations.

» Jensen's monograph of the Lycodids of Northern Europe and of Green-
land (Danish Ingolf Exped., vol. 2, Pt. 4, 1904) Includes descriptions, and
beautiful illustrations of all the species of Lycodes that have been reported
from the Gulf of Maine, or from the Nova Scotian Banks. And Vladykov
and Tremblay (sta. Biol. Saint Laurent. Fauna and Flora Laurent, No. 1,
1936) have given a revision of the genus in the western Atlantic with decrip-
tions and photographs of several new species and subspecies from the Gulf
of St. Lawrence and northward.

* Known from several stations on the continental slope abreast of our Gulf
and off southern New England at depths of 365-904 fathoms; also from the
deep gully between LaHave and Sable Island Banks at 200 fathoms (see
Goode and Bean 1895, p. 311 for list).

510

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

5. Lateral line Bingle, running along lower part of side of body; forward part of dorsal fin marked with one or more sooty

patches Lycodes vahlii Reinhardt 1838 *'

Lateral line double, with the more distinct branch of the two running along the lower edge of the side of the body;
the forward part of the dorsal fin is not marked with dark patches Lycodes esmarkii, Collett 1875 B

Ocean pout Macrozoarces americanus (Bloch
and Schneider) 1801

Eelpout; Congo eel; Mdttonfish

Jordan and Evermann, 1896-1900, p. 2457. Zoarces
anguillaris (Peck) 1804.

Description. — The ocean pout is blenny-like or
eel-like in form, its body about 8 times as long as
it is deep (10 to 11 times in young fish up to about
8 inches long), moderately flattened sidewise,
noticeably sway-bellied, and tapering backward
from abreast of the pectorals, where it is deepest,
to a pointed tail. It is very soft, its scales are
very small, and its skin as slimy as an eel. Its
ventral fins are small like those of the rock eel
(p. 492), and they are situated well forward of the
pectorals.

The most useful field mark for the identification
of the ocean pout among the several eel-like fishes
with which it might be confused are its vertical
fins. Its anal fin is continuous with the caudal,
there being no trace of any notch between the two,
as is the case with the true eels. In reality, this is
also true of the dorsal fin of the eelpout. But
about 16 to 24 of the dorsal rays near the rear end
of the fin are so short as to be hardly visible, so
that there seems to be a considerable free gap be-
tween the dorsal fin and the caudal fin. Further-
more, these short rays are spiny instead of soft as
all the other dorsal rays are. The dorsal fin runs
from the nape back along the whole length of the

»' Originally described from Greenland; reported from Banquereau Bank,
at 130-190 fathoms by Ooode and Bean (Smithsonian Contrlb. Knowl., vol.
30, p. 308, as Lycodes zoarckm. See Vladykov and McKemle, Proc. Nova
8ootian Inst. Sci., vol. 19, pt. 1, 1935, p. 109.

B Spitzbergen and Northern Norway; Grand Banks; LaHave Bank, and
southward along the continental slope in depths of 300-420 fathoms to the
offing of Rhode Island.

trunk, and consists of first about 95-100 soft rays;
next of the short spines, then of about 17 more soft
rays. The anal fin (about 105-124 soft rays)
originates a little in front of the mid length of the
fish. Both the dorsal fin and the anal are of nearly
even height from end to end except as just noted,
but the dorsal is nearly twice as high as the anal.
The pectoral fins are large and rounded like those
of the wolffish. The very small ventrals are on
the throat, in front of the pectorals. The upper
jaw projects a little beyond the lower, the soft,
fleshy upper lip somewhat farther still,33 enclosing
the tip of the lower lip when the mouth is closed.

The mouth is wide, gaping back beyond the
small eyes, and it is set low with thick and fleshy
lips that give the profile a distinctive aspect.
Both jaws are armed with two series of strong,
blunt conical teeth, largest in front, but the mouth
lacks the crushing teeth that are so characteristic
of the wolffish tribe (p. 502). There are 131-144
vertebrae.

Color. — -Although this fish has usually been de-
scribed as reddish brown mottled with olive, or as
salmon colored, most of those we have seen caught
have been of some shade of muddy yellow, paler
or darker; some tinged with brownish, some with
salmon, and some with orange; a few have been
pure olive green. Fishermen usually describe
them as yellow, and this is evidently the prevailing
hue in the offshore parts of the Gulf. Other ocean
pouts we have caught inshore along the coast of
Maine, however, have shown yellow only on the
margins of the fins, particularly the lower edge of
the pectorals, with the general ground tint of sides

u The ocean pout has sometimes been described and pictured as with the
upper lip and jaw projecting far beyond the lower; but this is contrary to our
observations.

ss&k

9&g*3K

. . . .• "o»r^VV.%r.v°0'^.>-.'»:;.%*.v.°a.£*W'V°i.v,'.

..^.a****

Figure 269. — Ocean pout (Macrozoarces americanus), Eastport, Maine, specimen. From Goode. Drawing by H. L.

Todd.

FISHES OF THE GULF OF MAINE

511

and back ranging from pale gray (sometimes with
purplish tinge) to dull brown or to dark dusky
olive; the belly ranging from dirty white or yellow-
ish or pinkish to the same dark shade as the back.
One of a pair that were caught side by side in
Northeast Harbor, Mount Desert, was pale
grayish white below while the other was amethyst
pink on the belly and on the lower side of the head.
Whatever the ground tint, the sides are dotted
with small dark spots clustered in irregular cross-
bars, extending out on the dorsal fin. And there
is a dark brown stripe running from eye to edge of
gill cover.

Young fry, up to 3 or 4 inches long, are check-
ered along the sides and irregularly blotched on
the back with light and dark brown, and they have
a small but prominent black spot on the forward
part of the dorsal fin until about 1 foot long, but
this spot fades out with growth.

Size. — The ocean pout is said to reach a length
of 3% feet and to a weight of 12 pounds; Olsen and
Merriman's largest, among some 2,500 specimens,
was 38^ inches long, weighing 11% pounds, and
we have seen one of about 3 feet among the many
we have handled. But only a few grow longer
than 30 inches, with 16 to 28 inches as a fair
average for the general run of those that are
caught.

The average weights of pouts of different size?
were about as follows for a large number taken in
the southern side of the Gulf of St. Lawrence, in
the Bay of Fundy, in the southwestern part of the
Gulf of Maine, and off southern New England:
16 inches, % pound; 18 inches, 1-1 K pounds; 20
inches, 1%-1% pounds; 22 inches, 2-2 % pounds;
24 inches, 2^-3 pounds; 26 inches, 3%-3% pounds;
28 inches, 4% pounds; 30 inches, h% pounds.34

Remarks. — The ocean pout of North Europe
(Zoarces viviparus Linnaeus 1758), a very close
relative, is distinguishable from the American
eelpout, by having fewer fin rays (about 100
dorsal rays and 6 to 10 spines; 80 to 89 anal
rays), fewer vertebrae (101-126), smaller head
and mouth, and only a single row of teeth in the
front of the jaw, while some specimens have no
interruption between the dorsal fin and the caudal.
Also, the European ocean pout is a smaller fish,

" According to graphs by Clemens and Clemens (Contrlb. Canadian Biol.
(1918-1920) 1921, flg. 5, p. 79), for the Gulf of St. Lawrence and Bay of Fundy;
and by Olsen and Merrlman (Bull. Bingham Ooeanographic Coll., vol. 9,
art. 4, 1946, flg. 4, p. 43) for the southwestern part of the Gulf of Maine and for
southern New England.

and its eggs are retained in the ovaries of the
mother until after they have hatched, hence its
specific name viviparus.

It is well known 36 that the European ocean
pout tends to break up into genetic races that are
partly correlated with environmental conditions.
And recent studies by Olsen and Merriman 39
make it likely that there is a slower growing race
of ocean pout in the Bay of Fundy and perhaps
northward, a larger, faster growing race ranging
from Cape Cod southward, with each of these
including minor subpopulations. This interesting
subject would repay further investigation.

Habits.37 — The ocean pout is a ground fish,
as might be expected from the fact that it has
no swim-bladder, as well as from its food (see
p. 512). And the habits of fish kept in aquaria,
where they are described as remaining coiled up
in the darkest parts suggests that they spend
most of their lives hiding among sea weeds and
stones. They are described as moving slowly
backward and forward by undulations of the
fanlike pectoral fins or of swimming more rapidly
by undulating motions of the rear part of the
trunk and tail, with the pectorals wide spread
and held horizontal, and with the dorsal and anal
fins close to the body.38 They swim actively
when disturbed. And it is almost unbelievable
to what a hopeless tangle of cord, fish, and slime
a few ocean pouts can reduce many fathoms of
long line set for other fish.

The vertical range of the ocean pout in one
place or another extends at least as deep as 105
fathoms.89 At the opposite extreme Clemens
and Clemens 40 report that young ones are some-
times found around rocks and in seaweed along
the shore in the Bay of Fundy during the ebb
tide. They are even known to run into rivers
for some distance, though always holding to the
bottom, i. e., to the undercurrent of water of

*' Especially from investigations by Johannes Schmidt and by J. V. O.
Smith (for list of references, see Olsen and Merriman, Bull. Bingham Ocean-
ographic Collection, vol. 9, art. 4, 1948, p. 182).

» Bulletin, Bingham Oceanographic Coll., vol. 9, art. 4, 1946, p. 116-117.

" Olsen and Merriman (Bull. Bingham Oceanogr. Coll., vol. 9, art. 4,
1946) have recently published a detailed study of the life history of the ocean
pout.

" Wllley and Huntsman (Canadian Field Naturalist, vol. 35, 1921, p. 6),
and Clemens and Clemens (Contrib. Canadian Biol [1918-1920], 1921, p. 71)
give some observations on the actions of ocean pouts kept in the aquarium at
the St. Andrews Laboratory.

" Albatrots III trawled 3 specimens from between 10S and 240 fathoms, on
the southwestern slope of Georges Bank in May 1950.

« Contrlb. Canadian Biol. (1918-1920) 1921, p. 72.

512

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

high salinity that tends to move inward along the
bottom from offshore. But most of them live
between 8-10 fathoms and perhaps 45 fathoms in
the waters with which we are most immediately
concerned. During the years when ocean pouts
were in demand (see below), good catches were
made as shoal as 10 to 12 fathoms in the south-
western part of the Gulf, also off southern New
England.41 And we have seen large numbers
caught from party boats, at 8-17 fathoms along
the coast of New Jersey.

We have taken ocean pouts in the Gulf of
Maine on sandy mud, on sticky sand, on broken
bottom, also on pebbles and gravel. They are
caught in large numbers on smooth hard bottom
and we have seen many more of them taken
from party boats off northern New Jersey on
rocky bottom, along with sea bass (p. 407), tautog,
cod, and other fishes, than were taken on soft
bottom when we were fishing for hake (Urophycis).

There is no evidence that they carry out any
extensive migrations. However, information has
accumulated recently to the effect that the adults
congregate through the summer, autumn, and
early winter on rocky bottoms where the eggs
are deposited and guarded, to disperse again in
midwinter (after the eggs have hatched), over
the smoother grounds in the vicinity where food
is more plentiful.42 And this spawning migration
appears to be complicated by an autumnal shift
offshore to deeper water, with a return movement
in spring, in coastal regions where the bottom
water chills in winter to a temperature too low
for their comfort; in the Bay of Fundy, for example
(p. 514), and perhaps in the Gulf of St. Lawrence.

The ocean pout can be classed as a cool-water
fish, for the great majority of them, in whatever
part of their geographic range, are in temperatures
at least no higher than about 62°, throughout
the year. At the other extreme, they have been
taken in 32° in the Bay of Fundy (p. 514); in
about 31° to 32° in Trinity and Conception Bays,
Newfoundland.43 And eelpouts are exposed to
temperatures as low as this, in spring, in the
Gulf of St. Lawrence, unless they descend deeper
into the Laurentian Channel, which they may do.

Food. — The American ocean pout feeds on a
wide variety of shelled mollusks, univalve and
bivalve, on crustaceans large and small, on echino-
derms and on other invertebrates. The Bay of
Fundy fish opened by Clemens and Clemens had
dieted chiefly on the two common mussels, Mytilus
and Modiolaria, on whelks {Buccinum) , periwinkles
(Littorina), and on scallops (Pecten) as well as
on various other bottom-living mollusks, on sea
urchins, brittle stars, and barnacles. A large
specimen caught in Massachusetts Bay, January
1924, was packed full of brittle stars (ophiurans),
spider crabs, and small sea scallops (Pecten magel-
lanicus) ; a number trawled by the Albatross III at
42 fathoms, near Nantucket Lightship, May 17,
1950, were full of small sea scallops (Pecten magel-
lanicus), as many as 100-200 per fish. Olsen
and Merriman 44 write that sand dollars (Echin-
arachnius) were the chief items in the stomach
contents of some 850 ocean pouts taken in the
southwestern part of our Gulf and off southern
New England, with crabs (Cancer) and isopod
crustaceans (Unicola) as seconds; while some had
eaten bivalve mollusks (Yoldia and Pecten) in
large amounts; also the eggs of the longhorn
sculpin (p. 451), which are often laid among the
branches of the finger sponge (Chalina)..

Ocean pouts bite on fish as greedily as they do
on clams or cockles, and pouts kept in the aquar-
ium at St. Andrews took fish as readily as clams; **
while two of the fish opened by Clemens and
Clemens,49 and also Bay of Fundy fish examined
by Olsen and Merriman,47 contained remains of
fish. But in all probability about the only fish
they get are dead ones that have sunk to the
bottom, or occasionally a small one that a pout
may have the good luck to catch. The European
representative of our ocean pout (Zoarces vivi-
parus) is described 4S as taking in mouthfuls of
weeds for the crustaceans and mollusks living
among these, and as swallowing a considerable
amount of the plant material with its animal prey.
But American ocean pouts appear not to feed in

" Olsen and Merriman, Bull. Bingham Oceanographic Coll., vol. 9, art.
4, 1946, p. 37, 38, fig. 3.

• This shift of grounds has been demonstrated recently by Olsen and
Merriman (Bull. Bingham Oceanog. Coll., vol. 9, art. 4, 1949, p. 40-42).

• Rept. Newfoundland Fish. Res. Comm., vol. 1, No. 4, 1932, p. 109.

« See Olsen and Merriman, (Bull. Bingham Oceanog. Coll., vol. 9, art. 4,
1946, p. 124-129) for a list of stomach contents, with discussion.

*• As reported by WUley and Huntsman, Canadian Field Naturalist, vol.
35, 1921, pp. 6-7.

" Contrib. Canadian Biol. (1918-1920) 1921, p. 80, small fish, including a
smelt.

" Bull. Bingham Oceanogr. Coll., vol. 9, art. 4, 1946, p. 129; probably
herring.

" By Blegvad, Report Danish Biol. Stat. (1916), 1917, p. 42.

FISHES OF THE GULF OF MAINE

513

this way to any great extent, for only traces of
plants have been found in their stomachs.

The eggs of the European ocean pout (Zoarces
viviparus) are fertilized within the mother, and
are retained within her oviducts until after they
hatch. But the American eelpout lays eggs, as
proved by the fact that the young were hatched
from a mass of eggs brought up, with two eelpouts,
in an old rubber boot,*' in Blacks Harbor,
Passamaquoddy Bay. And enough small speci-
mens of 1 H inches and upward have been collected
of late, between New Jersey and Maine, to show
that the eelpout breeds successfully throughout
this part of its range, at least; and probably as far
north as northern Nova Scotia and the southern
side of the Gulf of St. Lawrence.

Various lines of evidence 60 show that spawning
takes place in September and October. And the
fact that fish taken in summer differ widely in the
stage of development of their sexual products
suggested to Clemens and Clemens that they do
not breed every year. But it seems more probable
to us that this is evidence simply of a protracted
breeding period, some individuals ripening early
in autumn, others not until later.

Large females lay more eggs than small, the
numbers of maturing eggs actually counted having
ranged from 1,306 in a fish 21 }£ inches (55 cm.)
long to 4,161 in one of about 34% inches (87.5 cm.).

The eggs are yellow, 6-7 mm. in diameter, and
are laid in masses held together by a gelatinous
substance. The only egg mass so far brought in
was in an old rubber boot, suggesting that they
are normally deposited in crevices in rocks or
among stones, which would explain the apparent
tendency of the mature fish to congregate on
rocky bottom as the spawning season approaches.

The fact that eggs brought up in the trawl in the
Passamaquoddy region, where the spawning is sup-
posed to take place from mid-September through
October, hatched in early January, and that ocean-
pout eggs taken off New York in mid-November 6I
still were some weeks short of hatching, makes it
likely that incubation occupies at least 2% to 3
months. And the actions of a captive female
that lay coiled around its mass of eggs, though

these had not been fertilized, M makes it likely
that the eggs are guarded by one or the other
parent during this period, perhaps by both of them.

" For account, see White, Jour. Fish. Res. Bd. Canada, vol. 4, pt. 5, 1939,
pp. 337-338.

» Discussed in detail by Olsen and Merriman, Bull. Bingham Oceanogr.
Coll., vol. 9, art. 4, 1946, pp. 69-77.

n From the stomach of a cod, see Olsen and Merriman, Bull. Bingham.
Oceanog. Coll., vol. 9, art. 4. 1946, p. 76, flg. 9.
210941— 53 34

Figure 270. — Ocean pout (Macrozoarces americanus) .
Above, larva, 48 mm. Below, young fish, 387 mm.
Drawings by Louella E. Cable.

The larvae are about 30 mm. long at hatching,
i. e., much larger than those of most of our com-
mercially important fishes, and they are so far
advanced already in development that they are
easily identified.63 Being so nearly adult in form,
it is probable that they hold to the bottom from
the time they are hatched ; all catches of immature
fish recorded so far have, indeed, been on the
bottom.

The sizes, of the fry in different months, show
that ocean pouts reach a length of about 3 inches
during their first summer, and that they are about
4 to 5 inches long when 1 year old. According to
studies of otoliths by Olsen and Merriman,64 ocean
pouts in southern New England waters may be
expected to reach 6 inches when between 1 and 2
years old; 12 inches at 3 years; and 24 inches when
between 6 and 7 years, the very large fish of 36
inches and upward being 12 years to 16 years old.
Estimates by the same method by Clemens and
Clemens 6S point, however, to a considerably
slower rate of growth in the colder water of the
Bay of Fundy, where a 12-inch fish is likely to be
nearly 5 years old, a 24-inch fish between 12 and 13
years old; and where the 8 oldest fish examined

u See Olsen and Merriman (Bull. Bingham Oceanog. Coll., vol. 9, art.
4, 1946, p. 76, fig. 8) for a photograph of a female so employed, in the
Sbedd Aquarium, Chicago.

'* For further details, see White, Jour. Res. Board Canada, vol. 4, 1939, p.
338.

« Bull. Bingham Oceanogr. Coll., vol. 9, art. 4, 1946, pp. 85-93.

« Contributions to Canadian Biology, 1918-1920 (1921), p. 74.

514

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

(16-18 years) among 190 specimens, were only
26% to 28K inches long.

In one commercial catch, probably representa-
tive, made off Provincetown and analyzed by
Olsen and Merriman, most of the fish were 4 to 10
years old, with only scattered fish of 1 1 to 16 years.

Off southern New England, according to Olsen
and Merriman, a few females mature sexually
when they are about 18 inches long; about half at
22-23 inches, and all of them by the time they are
24-25 inches long; males mature earlier, most of
them by the time they have reached 15-16 inches;
a few not until larger. The smallest females with
large eggs seen in the Bay of Fundy region by
Clemens and Clemens were 16-18 inches long.

General range. — Coast of North America from
the Strait of Belle Isle, Gulf of St. Lawrence, and
southeastern Newfoundland south to Delaware; M
common from the southern side of the Gulf of St.
Lawrence and northern Nova Scotia to New
Jersey.

Occurrence in the Gulf of Maine. — The ocean
pout, known more familiarly as "conger", or
" congo" eel along the coast of Maine,67 is a familiar
fish in the Gulf in moderate depths of water both
near shore and on the offshore banks; abundant
locally off western Nova Scotia; in the Bay of
Fundy ; M all along the coasts of Maine and Massa-
chusetts; and on Georges Bank where considerable
numbers are taken both by otter trawlers and by
long-line fishermen.*8 Very small ones have been
collected off Chatham, Cape Cod; on Stellwagen
Bank at the mouth of Massachusetts Bay; and
near Mount Desert Island, Maine, by us; also in
the Bay of Fundy and in Passamaquoddy Bay by
Clemens and Clemens,69 evidence that it breeds
successfully all around the Gulf.

There seems to be a wide difference in the depth
zone frequented by the ocean pout in different
parts of the Gulf. In the Bay of Fundy some of
them run up into shoal water in summer and young
ones are to be found under stones and among sea-
weed between tide marks. Similarly, one is
always apt to catch several in a half day's flounder
fishing in 1 to 3 fathoms in Penobscot Bay or in

" There Is a doubtful record for North Carolina (Smith, North Carolina
Geological and Economic Survey, vol. 2, 1907, p. 379) .

" Years ago we heard them called "yowlers" by long-line fishermen, but
we doubt that this name is still used for them anywhere.

•' Clemens and Clemens (Contrlb. Canadian Biol. (1918-1920) 1921, p. 69)
give a general account of the ocean pout in the Bay of Fundy, and list the
localities there whence it has been recorded.

" Contrib. Canadian Biol. (1918-1920), 1921, p. 77.

Northeast Harbor, Maine, as we can bear witness.
And this probably applies to bays and harbors all
along the coast of Maine east of Cape Elizabeth.
But we have never seen one taken in less than 10
fathoms of water in the Massachusetts Bay region,
where most fishermen speak of it as a compara-
tively deep-water fish though it has been recorded
from Gloucester Harbor. And the ocean pouts on
the offshore grounds live mostly deeper than 20 to
30 fathoms. Thus Albatross II trawled a number
in the basin of the Gulf down to 90 fathoms in July
1931, while a large number of them have been
trawled on Georges Bank, at depths of 20-60
fathoms.60 And in May 1950, the Albatross III
trawled 3 at 105 fathoms or deeper 8l on the south-
western slope of Georges Bank.

Ocean pouts also frequent different types of bot-
tom in different localities. In Massachusetts Bay
they are seldom caught on the good fishing grounds
on stony or gravelly bottoms, that is, or about
ledges. But if the long line chances to run off
these, the portion of it that is resting on the softer
floor of the deeper parts of the bay often brings in
eelpouts and nothing else except an odd hake.
They are caught regularly on hard bottom, how-
ever, off Cape Cod and to the westward ; we have
trawled them on rather sticky sand in Ipswich Bay
(22 fathoms) among good catches of hake and
plaice; on broken bottom at the mouth of Casco
Bay; on pebbles and mud in Penobscot Bay; they
are commonly caught on stony ground farther
eastward along the coast of Maine; and Huntsman
describes them as taken on hard bottom in the
Bay of Fundy.

In fact, the only type of bottom where we have
not heard of them in our Gulf is the soft oozy
mud with high organic content that floors certain
of the deeper depressions, such as the trough to
the west of Jeffreys Ledge.62

Information as to the seasonal movements of
ocean pouts in different parts of the Gulf is not only
scanty, but perplexing. In the open Bay of Fundy,
Huntsman describes them as working inshore in
spring but moving out again into deeper water in

M Albatross III. for example, trawled 137 of them along the southern slope of
Georges, at 31-60 fathoms, in May 1960; the dragger Engine H trawled an
average of 8 pouts per haul at 26--15 fathoms, and about 2 per haul at 46-75
fathoms on the south central part of Georges Bant, in late June 1961.

" The depth ranged from 105 fathoms to 240 fathoms along the strip of
bottom on which the trawl was working.

K They were not represented among the considerable list of fishes trawled
In such situations by the Atlantis in August 1936 (Bigelow and Schroeder,
Biol. Bulletin, vol. 76, 1939. p. 309.

FISHES OF THE GULF OF MAINE

515

October or early November, whicb is corroborated
by a report of Clemens and Clemens, that set lines
made good catches in the Passamaquoddy region
from early June through September, but caught
none there from January to May. And their abun-
dance in Penobscot Bay in midsummer suggests
that some of them may perform a similar on and
offshore migration there, too. But this may not
apply to the coast south of Cape Elizabeth. Off
southern New England, where they are plentiful on
the commercial fishing grounds in winter and
spring, only stray ocean pouts are taken there in
summer and autumn. But it seems more likely
that they shift, then, to regions of rockier bottom
nearby, than that they move off-shore.63

It is not yet clear to what extent their move-
ments depend on the local food supply, on seasonal
changes in temperature, or on the habit the mature
fish have of congregating on rocky grounds during
the spawning season, and while guarding the eggs
thereafter (p. 513). One must also bear in mind
that failure to catch them on hook and line may
simply mean that they are not biting at the time,
not necessarily that they have moved away. This
is likely to apply to the adult fish in particular
during their spawning and egg-guarding season.

Importance. — -Although the ocean pout has few
bones and is said to be a sweet-meated fish, there
was no regular market for it prior to the early
1930's; only a few, brought in by small boats, were
sold on the streets of Boston, and nearly all of those
that were caught incidentally by the larger vessels
were thrown overboard. A small demand then de-
veloped for them resulting in landings for Massa-
chusetts ranging between 45,600 pounds and

114,700 pounds yearly, for the period 1935 to
1942, 64 though none for Maine.

A concerted attempt was made in 1943 to
market ocean pout as fillets, partly as a war
measure. This was so successful that 3,943,300
pounds were reported as landed in Massachusetts
ports in that year; 4,449,600 pounds in 1944, most
of them caught from the tip of Cape Cod south-
ward, and nearly all of them marketed through
New York. But this popularity was short lived,
for word soon spread that ocean pout are often
afflicted with a protozoan paraeite. Many ship-
ments were condemned for this cause, and the
demand fell off so rapidly that the landings for
Massachusetts were less than one-fourth as great
in 1945 (1,003,700 pounds) as they had been in
1944 «; fell to 613,300 pounds in 1946; were 167,400
pounds in 1947; and dropped to 6,100 pounds in
1948, the most recent year for which statistics of
the catch are available.

Wolf eel Lycenchelys verrillii (Goode and Bean)

1877.

Jordan and Evermann, 1896-1900, p. 2470.

Description. — This fish is eel-like in form and
resembles the ocean pout in most other respects
as well, except that it is more slender (about 14
to 16 times as long as it is deep), and that there is
no separation, apparent or real, between its dorsal,
caudal, and anal fins, but the three form a single
continuous vertical fin running along the back,
around the tail, and forward on the lower surface
to the vent. The dorsal fin not only originates

•> For discussion, see Olsen and Merriman, Bull. Bingham Oceanogr.
Coll. vol. 9, art. 4, 1946, pp. 40-42.

•* Landings (or 1933 to 1940 were listed as "conger eels"; no data are avail-
able for 1934 or 1936.

M For a detailed history of the event, see Olsen and Merriman, Bull. Bing-
ham Oceanogr. Coll., vol. 9, art. 4, 1946, pp. 9-10.

Figure 271. — Wolf eel (Lycenchelys verrillii), off Chebucto, Nova Scotia. From Jordan and Evermann. Drawing by

H. L. Todd.

516

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

farther back than in the ocean pout (over the tip
of the pectoral instead of in front of the base of
the latter), but all the dorsal rays (about 92) are
soft. Furthermore the anal fin (about 88 rays)
extends relatively farther forward than in the
ocean pout. The pectoral fins are rounded like
those of the ocean pout, but smaller relatively,
the small ventrals are similarly located well for-
ward of the pectorals, and in small specimens the
head resembles that of its relative in profile except
for a somewhat wider mouth. Old males, how-
ever (fig. 271), "are transformed almost beyond
specific recognition by an extraordinary develop-
ment of the entire head in advance of the eyes.
The snout becomes shovel-shaped, its length
equal to two-fifths that of the head, while in the
normal condition it is one-fourth".66

Color. — -The sides are light brown above the
lateral line, white below it, with a series of 8 to 10
irregular dark brown patches which the lateral
line bisects. The belly is blue, its lining jet black.

Size. — Maximum length about 10 inches; usual
length about 4 to 6 inches.

Habits. — This is a bottom fish, living on mud or
sand and confined to considerable depths of water.
Normally, 25 to 30 fathoms is its upper limit, but
the fact that the Grampus specimen mentioned
below was taken in a tow net, though close to
bottom, proves that it sometimes rises from the
ground. To the southward, on the continental
slope, it has been trawled down to 603 fathoms.

Nothing is known of its way of life or of its
breeding habits.

General range.- — So far known only off the coasts
of Nova Scotia and of New England, and south-
ward along the continental slope to the offing of
Beaufort, N. C, in rather deep water.

Occurrence in the Gulf of Maine. — The wolf eel
has been trawled at many localities on the con-
tinental slope at 200 to 600 fathoms from the
offing of Cape Fear, North Carolina (lat. 34°40'
N., long. 75°15' W.) to abreast the western end of

• Goode and Bean, Smithsonian Contrib. Knowl.. vol. 30, 1895, p. 310.

Georges Bank (long. 68°22' W.).w It was formerly
regarded as very rare within the Gulf of Maine,
the only records for it there up to 1925 having
been of a few specimens trawled off the mouth of
Passamaquoddy Bay in 35 to 50 fathoms; of one
4 inches long taken off Monhegan Island by the
Grampus on August 2, 1912, in 60 fathoms; and
of several that were collected by the U. S. Fish
Commission many years ago off Cape Ann in 75
to 110 fathoms, in the Western Basin in 115
fathoms, and off Cape Cod. More recent captures,
however, of a number of wolf eels at about 90
fathoms in the trough west of Jeffreys Ledge by
the Albatross II in November 1927, in August 1928,
and in September 1930 (a total of 61 specimens)
show that they are more plentiful in the deeper
parts of the Gulf than the previous record might
have suggested.

The only definite records for the wolf eel east-
ward from Cape Sable are of 5 specimens taken
by the U. S. Fish Commission steamer Speedwell
3 miles off Cape Negro, Nova Scotia, in 90 fathoms,
in the summer of 1877 68, and of one ■• taken 27
miles off the entrance to Halifax Harbor (Chebucto
Head) in 101 fathoms.70

Arctic eelpout Lycodes reticulatus Bernhardt 1838

Jordan and Evermann, 1896-1900, p. 2465.

Description. — This fish resembles the ocean pout
in its general appearance and in the arrangement
of its fins. The readiest field marks for it are
that the dorsal fin is not interrupted, but is con-
tinuous with the caudal fin, and that the dorsal
originates behind the bases of the pectorals instead

i Goode and Bean (Smithsonian Contrib. Knowl., vol. 30, 1895, p. 310)
give a long list of localities from the early cruises of the U. S. Coast Survey
and of the U. S. Fish Commission. And Beebe (Zoologica, N. Y. Zool. Soc,
vol. 12, 1929, p. 18) has reported a wolf eel from the Hudson Gorge, off New
York, at 69 fathoms.

M Reported by Goode and Bean, American Jour. Science, ser. 3, vol. 14,
1877, p. 473.

•' This is the specimen pictured here in flg. 271.

w Probably these specimens were the basis for Jones' (List Fishes Nova
Scotia, 1879, p. 5; Proc. Nova Scotian Inst. Sci., vol. 5, 1882, p. 91) statement
that the wolf eel occurs on the Nova Scotian fishing banks.

Figure 272. — Arctic eelpout (Lycodes reticulatus). Drawing by Louella E. Cable.

FISHES OF THE GULF OF MAINE

517

of in front of them, while the fanlike pectoral fins
are even larger, relatively, than those of the ocean
pout. Furthermore, its upper jaw projects far
beyond the lower, giving it a distinctive cast of
countenance (compare fig. 272 with fig. 269). The
most obvious difference between this Lycodes and
the wolf eel (p. 515) is that the former is much the
stouter bodied of the two, being only about 8 times
as long as it is deep (the wolf eel is 14-16 times as
long as deep), and that the dorsal fin of Lycodes
reticulatus originates farther forward, i. e., close
behind the bases of the pectorals instead of over
the tips of the latter.

Color. — Described as brownish, with a network
of black lines on the head and with several groups
of such lines or with solid dark bands on the body.
The dorsal fin is dark edged. The young fry are
marked with a series of large dark spots od the back
and extending out on the dorsal fin.71

Size. — Specimens of which the measurements
have been definitely recorded have ranged up to
15 inches (380 mm.) in length.

Remarks. — -This lycodid tends to separate into
local races ; one such from northeast Greenland and
Jan Mayen has, in fact, been dignified with a
separate varietal name; var. macrocephalus by
Jensen,72 because seemingly separable from the
West Greenland form. One subspecies, hacheyi,
too, has been described subsequently from Hudson
Bay by Vladykov ; 73 also a second (lavalei) from

the Gulf of St. Lawrence by Vladykov and
Tremblay 74 but none of these call for considera-
tion here.

Habits. — Little is known of its habits except
that it is a ground fish, usually living in moder-
ately deep water, and that worms, crustaceans, and
small fish have been found in the stomachs of
European specimens. In its turn it falls a prey
to larger fishes, and frequently to Greenland
sharks.75

General range. — Both sides of the Arctic Atlantic ;
reported as far south as Vineyard Sound, southern
Massachusetts.

Occurrence along the Atlantic coast of North
America. — This particular Lycodes has been
reported definitely off southeastern Labrador in
the offing of Sandwich Bay; in the Strait of Belle
Isle; in Conception Bay, Newfoundland; on the
Grand Banks; off Placentia Bay, south coast of
Newfoundland; also on the Newfoundland side of
Cabot Strait ; 76 and on the southwest slope of
Banquereau Bank at 300 fathoms ; 77 while
Vladykov and McKenzie report it from Nova
Scotian waters in general.78

It has not yet been recorded from the Gulf of
Maine. But it is to be expected there sooner or
later, for it has not only been taken to the east and
north of our limits, as just stated, but also in
Vineyard Sound to the westward,79 if the specimen
in question was identified correctly.

THE CUSK EELS. FAMILY OPHIDIIDAE

The members of this family are eel-like in
appearance, but they differ very obviously from
the true eels in having well-developed ventral fins
in the form of loDg forked barbel-like structures,
situated on the throat. Their soft-rayed dorsal
and anal fins are low but continuous around the
tip of the tail; the gill openings are very wide, and
the snout projects beyond the mouth. Many
species are known, most of them from warm seas.

" See Smitt (Scandinavian Fishes, vol. 1, 1892, p. 605) for the coloration of
the genus Lycodes in general; Jensen (Danish Ingolf Exped., vol. 2, pt. 4,
pi. 2, figs. 2, 3, and pi. 8) for beautiful illustrations of this species.

" Danish Ingolf Exped., vol. 2, pt. 4, 1904, p. 66, pi. 8.

'■ Contr. Canad. Biol., N. Ser., vol. 8, No. 2, 1933, p. 25.

" Fauna et Flora St. Laurent., Sta. Biol. St. Laurent., No. 1, 1936, p. 34.

" Smitt, Scandinavian Fishes, vol. 1, 1892, p. 613.

'• For records of it in Labrador and Newfoundland waters, see Goode and
Bean (Smithsonian Contrib. Knowl., vol. 30, 1895, p. 305); also the Annual
Reports of the Newfoundland Fisheries Research Commission, vol. 1, No. 4;
vol. 2, Nos. 1-3, 1932-35.

Cusk eel Lepophidium cervinum (Goode and

Bean) 188580
Jordan and Evermann, 1896-1900, p. 2484.

Description.- — The cusk eel is about 12 to 13
times as long as it is deep ; all its fins are soft and
eel-like; and there is no separation between the
dorsal, caudal, and anal fins, but the three form
one continuous fin running backward along the

77 See Goode and Bean, Smithsonian Contrib. Knowl., vol. 31, 1895, List
of plates and figures, p. 17, figs. 273, 281.

" They do not mention any definite locality records but write of it (Proc.
Nova Scotian Inst. Sci., vol. 19, 1935, p. 109) as usually believed to be the most
common Lycodes there.

" Goode and Bean (Smithsonian Contrib. Knowl., vol. 30, 1895, p. 305) ,
Fish-Hawk Station 681; Sumner, Osbum, and Cole (Bull. U. S. Bur. Fish.,
vol. 31, Pt. 2, 1913, p. 768). Goode and Bean also report it from east of the
Bahamas (Albatross Sta. 2652, lat. 24°13' N., long. 77°13' W., 140 fathoms).
But this is so very far to the south of the normal range of this species that we
suspect the record is an error.

» See Jordan and Evermann (Bull. 47, U. S. Nat. Mus., Pt. 3, 1898, p. 2482)
for the nomenclatural history of the name Lepophidium Gill 1895.

518

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

MPPRHI^^W

^^^%o^afessaa^^^

Figure 273. — Cusk eel (Lepophidium cervinum). After Goode and Bean. Drawing by H. L. Todd.

back, around the tip of the tail, and forward on the
lower surface. But it is separated from all the
true eels, by the presence of ventral fins, situated
on the throat far in front of the pectorals, and
reduced to forked barbel-like structures. The
structure of the ventral fins and the uninterrupted
dorsal fin separate the cusk eel from the ocean
pout, its near relative among local fishes. And
the presence of a short sharp spine on the top
of the snout pointing forward and downward,
which is easily felt if not seen (for it is nearly con-
cealed in the skin), likewise differentiates it from
such other Gulf of Maine species as it resembles in
general appearance. The shape of the snout, too,
is distinctive, as are its rather large scales, for the
other genera of its family have naked heads, and
the scales on their bodies are very small.

Color. — Brownish yellow, darker above than
below, the upper part of each side marked with a
row of 14 to 23 roundish white or pale brown spots.

The dorsal and anal fins have narrow black or
dusky margins. It seems that the adult color pat-
tern develops late, for neither the spots nor the
edgings on the fins are visible in a young specimen
of 2% inches.

Size. — The type specimen was about 10% inches
(262 mm.) long.

General range and occurrence in the Gulf of
Maine.- — This fish has been taken at various local-
ities along the outer part of the continental shelf
from off Florida to eastward of Nantucket, in
depths of 38 to 102 fathoms. It is mentioned here
because one specimen has been taken in 76 fath-
oms off Nantucket Shoals,81 while two others,
about V/i and 8% inches long (newly swallowed)
were found in the stomach of a white hake Uro-
phycis tenuis) that was trawled on the south-
western part of Georges Bank (lat. 40°31' N.,
long. 68°55' W.), at 39 fathoms, by the dragger
Eugene II. on June 27, 1951.

THE TOADFISHES. FAMILY BATRACHOIDIDAE

The toadfishes are somewhat sculpin-like in
appearance, but the resemblance is superficial, for
their ventral fins are situated on the throat well
in front of the pectoral fins ("jugular"), and they
have only three gills and three gill arches. Both
the soft and the spiny portions of the dorsal fin
are well developed as separate fins, the former
much longer than the latter. Most of the species
belong to warm seas, only one reaching the Gulf
of Maine.82

Toadfish Opsanus tau (Linnaeus) 1766

Jordan and Evermann, 1896-1900, p. 2315.

Description. — The toadfish, like the sculpins, has
a large flat head, round nose, tremendous mouth,
tapering body with plump belly, and fanlike
pectoral fins. But it differs from all sculpins,

» Goode and Bean, Smithson. Contrib. Know!., vol. 30, 1895, p. 347.
" See Schulti and Reid (Copeia, 1937, No. 4, p. 211) for a synopsis of the
American-Atlantic species of the genus Optanut.

and indeed from all other spiny-finned fishes of
the Gulf of Maine except the blennies (p. 491) in
the location of its ventral fins, which are under the
throat well in front of the pectorals ("jugular")
instead of below the latter or behind them. And
no one could confuse it with any blenny, for it is
not only a totally different looking fish, but its
dorsal fin is mostly soft rayed while that of the
blennies is spiny throughout. The presence of
fleshy flaps of irregular outline on the tip of the
upper jaw and along the edge of the lower jaw,
on the cheeks, and over each eye, gives its head
a peculiar wary appearance. Distinctive, also, is
the fleshy nature of all its fins and the outline of
the dorsal, the soft part of which (26 to 28 rays)
is five to six times as long as the spiny part (3
spines), from which it is entirely separated by a
deep notch, the two together extending the whole
length of the trunk from the nape of the neck
nearly to the base of the caudal fin. The anal

FISHES OF THE GULF OF MAINE

519

Figure 274. — Toadfish (Opsanus lau), Woods Hole. Drawing by Louella E. Cable.

fin (21 or 22 rays) is somewhat shorter than the
second (soft) dorsal, orginates under about the
eighth ray of the latter, and is similar to it in
outline except that its rays are more or less free
at their outer ends, especially in its forward half.
The caudal fin is rounded; the ventrals covered
by thick fleshy skin, are jagged in outline, with
the first ray stouter than the others. There is a
large open pit of unknown function in the axil of
each pectoral fin.

We need only remark, further, that the skin is
covered with a thick layer of slimy mucus. The
toadfish has no scales; its teeth are large and blunt;
and it has two short spines at the upper angle of
each gill cover, hidden however, in the thick skin.

Color. — The general ground tint ranges from
dark muddy olive green to brown or yellow, darker
on back and sides, paler below, and variously and
irregularly marked with darker bars and marblings,
which may be restricted to head and fins or may
extend over the whole fish, belly as well as back.
The toadfish, like many other bottom fishes,
changes color to match the bottom on which it
lies.

Size. — Exceptionally 1 5 inches long, but few are
longer than 12 inches.

Habits. — The toadfish lives in shoal water, and
it is resident the year around wherever found,
probable becoming torpid in winter in the northern
part of its range. It is commonest on sandy or
muddy bottom, hiding among eelgrass or under
stones where it hollows out dens in which it lies
in wait for prey. It is voracious and omnivorous,
Vinal Edwards's diet list for it at Woods Hole
including sea worms (Nereis), amphipods, shrimps,

crabs, hermit crabs, a variety of mollusks both
univalve and bivalve, ascidians, squid, and fish
fry such as alewives, cunners, mummichogs, men-
haden, puffers, sculpins, scup. silversides, smelt,
and winter flounders. No doubt any small fish is
acceptable.

Toadfish snap viciously when caught, and they
often fight among themselves. Like some sculpins
they grunt, especially at night or if handled. And
despite their clumsy appearance, they can dart
out of their hiding places and back again with
considerable speed. They are very tenacious of
life and can live out of water for an astonishingly
long time.

In the northern part of its range the toadfish
spawns in June and early July. The very large
eggs (about 5 mm. in diameter) are laid in holes
under stones, under large shells, in old tin cans,
among sunken logs, or among eelgrass, where they
adhere in a single layer to whatever serves as a
nest, which the male guards during the 3 weeks or
so that are occupied by incubation. Even after
hatching the tadpole-shaped larvae remain at-
tached to the nest by the yolk sac until the latter
is absorbed at a length of 15 to 16 mm. when they
break free.88

General range. — Shoal water along the east
coast of North America from Cuba to Cape Cod,
straying northward to Maine.

Occurrence in the Gulf of Maine. — The toadfish
is common about Woods Hole and thence west-
ward. But it ventures around Cape Cod so

- Ryder (Bull. U. S. Fish Comm., vol. 6, 1887, p. 8) and Gudger (Bull.
U. 8. Bur. Fish., vol. 28, 1910, pt. 2, pp. 1071-1109, pla. 107-113) describe tho
breeding habits, eggs, and larvae of the toadfish. For further accounts
of its habits see 0111 (Smithsonian Mlscell. Coll., vol. 48, 1907, pp. 388-427).

520

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

seldom that none of the fishermen in Massachu-
setts Bay of whom we have inquired have seen or
heard of it there, nor further north. In fact there
are only two definite records of it in the Gulf of
Maine: "Maine"84 and Cohasset on the south

shore of Massachusetts Bay, where one (now or
formerly in the collection of the Boston Society
of Natural History) was caught by Dr. Owen
Bryant. There is also one doubtful record for
Kittery, Maine.85

THE TRIGGERFISHES. FAMILY BALISTIDAE

The triggerfishes are very divergent from the
ordinary spiny-rayed fishes anatomically, and their
external appearance is so characteristic that they
are not apt to be mistaken, unless for their close
relatives, the filefishes (p. 52 1 ) . Tb eir most interest-
ing external characteristics are that the first spine
of the first dorsal fin is not only very much stouter
than the others but it can be locked erect by the
second dorsal spine, and that the large bony
scales are so close set as to form a hard but flexible
armor. Other distinctive features are mentioned
below in the description of the Gulf of Maine
species. Most of the many species of triggerfishes
are purely tropical; it is only as a stray that any
member of the family enters into the Gulf of
Maine fauna. Most of the tropical species are
more or less poisonous if eaten.

" Storer, Mem. Amer. Acad., N. Ser., vol. 2, 1846, p. 384, gives no definite
locality.

"Holmes (2nd Ann Rept. Nat. Hist. Oeol. Maine, Pt. I, 1862, p. 95),
"noticed It while standing on the bridge which connects the Navy Yard at
Kittery with one of the islands."

Triggerfish Batistes carolinensis Gmelin 1789 8fl
Jordan and Evermann, 1896-1900, p. 1701.

Description. — The readiest field marks for the
triggerfish are its deep, sidewise-flattened body
with slender caudal peduncle; its small terminal
mouth with both dorsal and ventral profiles of the
nose nearly straight; its eye situated so high as to
give its face a very peculiar aspect; its large pro-
jecting incisor teeth; its very short gill openings
wholly above the insertions of the pectorals; the
plate armor of thick scales with which its entire
head and body are clad ; and especially its unusu-
ally stout first dorsal spine.

The spiny dorsal fin is triangular, with three
spines, the first so stout that it is more like a horn,
situated close behind the eyes and with the second
spine acting as a trigger to lock the first spine

"Jordan, Evermann, and Clark (Rept. U. S. Fish Comm. (1928), Pt. 2,
1930, p. 491) prefer the name capriscus Gmelin, and correctly so, from the
strictly nomenclatural standpoint. But carolinensis is preferable both be-
cause it appeared on an earlier page of the same publication, and because the
great majority of references to the species have been under that name.

^

Figure 275. — Triggerfish (Batistes carolinensis), New York. From Goode. Drawing by H. L. Todd.

FISHES OF THE GULF OF MAINE

521

erect, whence the common name of the fish. The
soft dorsal fin (27 rays) is separated from the first
by a considerable interspace, is rhomboid in out-
line with the third or fourth ray longest, and
tapers back to the base of the caudal peduncle.
The anal fin (25 rays) corresponds to the soft
dorsal in outline and in location. The caudal fin
is of moderate size, its rear margin moderately
concave, in a very characteristic curve, with
sharp pointed, somewhat prolonged corners. The
pectorals are short, rounded, and situated below the
gill openings. The ventrals are reduced to one
short, stout, blunt spine, mostly embedded in the
skin and they are connected with the general out-
line of the abdomen by a sort of dewlap.

Color. — The colors of this triggerfish vary
widely. A specimen 2 inches long recently taken
on Georges Bank was yellowish, with many small
blue-violet spots on the sides, dusky-blotched
along the back, and with one broad, irregular
dusky band extending from the base of the dorsal
fin almost to the anal. The caudal fin was pale
yellow. Other examples have been described as
olive gray, marked with violet dots and with dark
crossbars, the fins as variously tinted with yellow,
blue, and olive.

Size. — Said to reach a weight of 4 pounds, but
they average only about 1 pound.

General range. — Both sides of the tropical At-
lantic, also the Mediterranean; straying north to
Ireland on the European coast; to the outer coast
of Nova Scotia, in the vicinity of Cape Canso 87 on
the American side.

Occurrence in the Gulf of Maine. — Only one speci-
men of this tropical fish, taken in the Squam River
at Annisquam, near Gloucester, Mass., many
years ago,88 had been reported from the Gulf of
Maine previous to 1925. But it must drift in over
the offshore rim of the Gulf more often than had
been suspected, for two small fry of 2 to 3 inches
were picked up on the northeast part of Georges
Bank among Gulf weed (Sargassum) from the
Albatross II, in mid-September 1927; a large one
about 15 inches long was gaffed at the surface
from the fishing schooner Huntington Sanford 14
miles southeasterly from Highland Light, Cape
Cod, on July 19, 1929; 89 one, now in the Museum
of Comparative Zoology, was picked up at Ply-
mouth, Mass., on September 5, 1932, by the late
C. L. Hauthaway, a well-known angler and a close
observer.90 One was reported from Casco Bay,
(Small Point), Maine, in 1949, and one near
Boothbay Harbor (Linekin Bay), also in 1949.91

THE FILEFISHES. FAMILY MONACANTHIDAE

The filefishes recall the triggerfishes in their gen-
eral form, being similarly deep and flattened
sidewise, with the same peculiar profiles, small
terminal mouths, projecting incisor teeth, eyes set
high up, very stout dorsal spines, and short gill
openings; also in the fact that the ventral fins are
either lacking altogether or at least are reduced to
a single short blunt movable spine at the end of
the very long pelvic bone, forming a keel-like
continuation of the general ventral profile of the
head and connected with that of the belly by a
dewlap of skin. The filefishes differ from trigger-
fishes in having only one dorsal spine instead of
three, and in the fact that their scales are so minute
that the skin is velvety to the touch although very

tough. Most of the species are tropical or sub-
tropical, and none has any commercial or sporting
value. Adults of the four species known from the
Gulf of Maine are separable as follows:

17 Nova Scotian records are from Halifax and from Queensport near Cap©
Canso (Vladykov, Proc. Nova Scotian Inst. Sci., vol. 19, 1935, p. 9; McKenzie
Proc. Nova Scotian Inst. Sci., vol. 20, 1939, p. 18); also 24 miles southeasterly
from Sable Island, where one was picked up by the schooner Wanderer, July
5, 1931 (Firth, Bull. 61, Boston Soc. Natural History, 1931, p. 13).

88 This specimen is now in the Museum of Comparative Zoology.

81 Reported by Firth, Bull. No. 61, Boston Soc. Nat. Hist., 1931, p. 12.

" Another species of trigger fish (Batistes netuia) is more common than
carolinensis at Woods Hole, and is recorded from Nantucket, but it has not
been taken in the Gulf of Maine as yet. It is separable from carolinensis by
the fact that the forward rays of its soft dorsal fin and the corners of its caudal
fin are elongated and filamentous; also by the presence of two blue bars on
each side of its head.

•' These last two fish were reported by Scattergood, Trefethen, and Coffin ,
Copeia, 1951, No. 4, p. 298).

KEY TO GULF OF MAINE FILEFISHES

There is a prominent external ventral spine; the gill openings are nearly vertical.
There is no external ventral spine; the gill openings are very oblique

2
3

522

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

2.

3.

Dorsal profile of head in front of the eyes is straight, or only very slightly concave; there are no thorns on the sides of

the caudal peduncle Filefish (Monacanthus hispidus), p. 522

Dorsal profile of head in front of the eyes is conspicuously concave; there are about 6 stout thorns pointing forward

on each side of the caudal peduncle Filefish (Monacanthus ciliatus) , p. 523

Dorsal fin with about 34 to 38 soft rays; anal with 36 to 41 rays; also, in small specimens, the caudal fin is more than

one-half as long as the body Orange filefish (Alutera schoepfii), p. 524

Dorsal fin with about 44 to 48 soft rays; anal with 47 to 52 rays; also, in small specimens, the caudal fin is less than

one-half as long as the body Unicornfish (Alutera scripta), p. 525

Filefish Monacanthus hispidus (Linnaeus) 1766

Jordan and Evermann, 1896-1900, p. 1715.

Description.- — In this species the rear edge of the
dorsal spine, which is situated over the rear edge
of the eye, is armed with a double series of barbs,
but the sides of the rounded caudal peduncle do
not bear any spines. The point of origin of the
soft dorsal fin (32 to 34 rays) is behind the middle
of the body, while the first soft dorsal ray often is
much prolonged in adults and with a filamentous
tip (young 1 to 2 inches long lack this filamentous
ray.) Otherwise the fin is rounded in outline,
narrowing from the front to the rear. The anal
fin (31 to 34 rays) stands below the soft dorsal,
and is of the same shape except that none of its
rays are prolonged. The caudal fin is rounded.
The pectorals are short, rounded, and situated
lower than the gill openings, like those of trigger-
fishes.

Color. — Greenish, olive, or brownish. The back
and sides of young fish are mottled with irregular
darker blotches but adults are plain colored. The
dorsal spine and the caudal fin are green. The
soft dorsal fin and the anal fin are pale and
translucent.

Size. — Maximum length about 10 inches.

General range. — This is a tropical species, com-
mon along the south Atlantic Coast of the United
States and among the West Indies; also around
the Canaries and Madeira in the eastern Atlantic,
and represented in East Indian waters by a fish
that does not seem to be distinguishable from it
in any way. In the western side of the Atlantic
it is known as far south as Brazil, has been taken
from time to time as far north as Woods Hole, and
has been recorded from St. Margaret Bay and
from Halifax on the outer coast of Nova Scotia.62

" The only recent Nova Scotian record that has come to our notice is of one
taken in Halifax Harbor in the autumn of 1928 (Leim, Proc. Nova Scotian
Inst. Sci., vol. 17, Pt. 4, 1930, p. 46).

Figure 276. — Filefish (Monacanthus hispidus). From Jordan and Evermann. Drawing by W. S. Haines.

FISHES OF THE GULF OF MAINE

523

Occurrence in the Gulf of Maine. — Odd speci-
mens of this filefish have been recorded from
Hingham, Lynn, Nahant, and Boston Harbor in
Massachusetts Bay; and from Cape Cod; all many
years ago. More recent records of it in the Gulf
are of 181 fry, 1-2 inches long, picked up from the
Albatross II on the northeastern part of Georges
Bank among floating Gulf weed (Sargassum), in
September 1927; a larger one taken off Seguin
Island, Maine, September 12, 1927 ;93 one of 6
inches, at Provincetown, November 6, 1929; one
picked up from the schooner Old Glory among
floating rockweed (Fucus or Ascophyllum) and
Gulf weed (Sargassum), on the western part of
Georges Bank, September 15, 1930; 94 one taken
off Portland Lightship, July 17, 1931; one taken
in a trap at Provincetown, October 6, 1950; and
two, about 6 inches long, taken off Wood End,
Provincetown, in 17 fathoms, by the dragger Alary
Magdalyn (Capt. Charles Santos), on October 30,
1951. It is also likely that a "filefish" taken at
Beverly on the north shore of Massachusetts in
1933 95 was of this species. An occasional filefish

straying from the south is thus to be expected
anywhere on Georges Bank, or in the western side
of the Gulf. But we find no evidence that they
ever enter its eastern side, or that they ever reach
the Bay of Fundy.

Filefish Monacanthus ciliatus (Mitchill) 1818
Jordan and Evermann, 1896-1900, p. 1714."

Description. — This filefish resembles the species
hispidus (p. 522) very closely. But its first dorsal
ray is never prolonged, the ventral dewlap extends
somewhat farther behind the tip of the ventral
spine, and the caudal peduncle in the adult is
armed with 2 or 3 pairs of strong forward-curving
hooks on either side.

Color. — Described as varying from olive gray,
or grass green to yellowish brown, with darker
blotches or crossbands. The dorsal and anal fins
are pinkish and they usually have three dark
spots at the base. The ventral dewlap is edged
with scarlet, and the caudal fin is greenish, mottled
dark and pale.

« Kendall, Bull. 58, Boston Soc. Nat. Hist., 1931, p. 11.
•' Firth, Bull. 61, Boston Soc. Nat. Hist., 1931, p. 13.
» MacCoy, Bull. 67, Boston Soc. Nat. Hist., 1933, p. 9.

"The illustration labeled "ciliatus" by Jordan and Evermann (Bull. 47,
U. S. Nat. Mus., Pt. 4, 1900, pi. 259, fig. 634) is actually of hispidus, as is their
figure 635.

Figure 277. — Filefish (Monacanthus ciliatus), Bahamas specimen, 85 mm. long. Drawing by H. B. Bigelow.

524

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Size.— Four to eight inches long.

General range. — Warmer parts of the Atlantic;
from Brazil to Cape Cod on the American coast.
A straggler has even been reported from Argentia,
on the southern coast of Newfoundland, far to the
north of its normal range.97

Occurrence in the Gulf of Maine.- — The only
recent report of this filefish from within the Gulf
of Maine is of a 7-inch specimen taken in a trap
at Provincetown, November 9, 1929.98 We judge
that earlier reports of it from Massachusetts Bay
referred to in Storer's description and illustra-
tion " were actually based on a specimen of
hispidus.

Orange filefish Alutera schoepfii (Walbaum) 1792 i

Filefish; Tttrbot; Hogfish; Sunfish;
Unicornfish

Jordan and Evermann, 1896-1900, p. 1718.

Description. — This fish resembles its relatives of
the genus Monacanthus in most respects (p. 521),
but while it is equally flattened sidewise, it is
relatively shallower, being not over half as deep as
long. The pelvic bone is as prolonged as it is in
the other filefishes, but it does not project ex-

•' Newfoundland Fish. Res. Comm. Rept., vol. I, No. 1, 1933, p. 126.

» Firth, Bull. Boston Soc. Nat. Hist., No. 61, 1931, p. 13.

" The illustration of his Monacanthu* massadiusettensis shows the profile
typical of hupidui, and neither his description nor his illustration suggests
that there were any thorns on the caudal peduncle such as characterize
ciliatus. (See Storer Mem. Amer. Acad. Arts, Sci., N. Ser., vol. 8, 1863,
p. 425, pi. 34, fig. 4; also Fishes of Massachusetts, 1867, p. 231, pi. 34, flg. 4.)

1 Jordan, Evermann, and Clark (Rept. U. S. Comm. Fish. (1928), Ft. 2
1930, p. 495) place the species in the genus CeratacaTithui Gill 1801.

^¥m^^i^.ff<c

ternally, nor is there a ventral dewlap, which is
the readiest field mark by which to distinguish
Alutera from Monacanthus. The eyes, too, are
set lower down on the sides of the head, and the
gill openings are relatively longer and more
oblique. The dorsal spine is relatively shorter
than in Monacanthus, and the lower jaw projects
considerably beyond the upper. The soft dorsal
fin (34-38 rays) originates behind the middle of
the trunk and is rounded in outline, and the anal
fin (36—41 rays) corresponds to the soft dorsal fin
in size, shape, and position. The short rounded
pectorals are situated opposite the lower half of
the oblique gill openings and the tail fin is relatively
narrower than in the other filefishes or trigger-
fishes; its longest rays are more than ){ as long as
the body in small specimens, but only one-fourth
to one-fifth as long as the body in half-grown fish
and larger.

Color. — Described as varying from uniform
olive gray to rich orange yellow or to milky white
above, mottled with darker hues of the same tints;
bluish white beneath. The caudal fin usually is
yellowish on adults but sometimes it is dusky,
edged with white.

Size. — Maximum length about 2 feet.

General range. — Atlantic and Gulf of Mexico
coasts of the United States; not uncommon in
summer as far north as Cape Cod; reported to
Portland, Maine, and perhaps to Halifax, Nova
Scotia.2

1 A specimen, found in Halifax Harbor, August 25, 1938, appears to have
belonged to this species, but it was not In good enough condition for positive
identification (McKenzie, Proc. Nova Scotian Inst. Sci., vol. 20, 1939, p. 19).

^Mm^,

0$

Fiqtjbe 278. — Orange filefish (Alutera schoepfii), Key West, Florida. From Jordan and Evermann. Drawing by

W. S. Haines.

FISHES OF THE GULF OF MAINE

525

Occurrence in the Gulf of Maine. — Although the
orange filefish has been described as "rather
common" at Woods Hole during the summer,3
only 3 specimens have been reported within the
limits of the Gulf of Maine: 1 from Portland,
Maine, and 2 from Salem, Mass., all of them
many years ago.4 Evidently it reaches the Gulf
of Maine only at long intervals, as a waif from the
south.

Unicornfish Alutera scripta (Gmelin) 1789

Jordan and Evermann, 1896-1900, p. 1719.5

Description. — This species much resembles the
orange filefish from which it differs chiefly in its
somewhat more slender body (2 to 3 times as long
as deep), longer dorsal fin (44 to 49 rays), longer
anal fin (47 to 52 rays), shorter caudal fin (in
small unicornfish the caudal is less than half as
long as the body, while in young orange filefish

it is more than half as long), and in color. The
dorsal spine may be serrated in young fish, but it
is smooth in adults.

Color. — The unicornfish is olive on head and
body with light blue reticulations extending from
the snout to the tail; in grown fish there are
numerous small round black spots on the sides of
the body.

Size. — Reaches a length of 3 feet.

General range. — -Tropical seas; northward to
South Carolina on the Atlantic Coast of America,
and to Georges Bank as a stray.

Occurrence in the Gulf of Maine.- — Two speci-
mens of this fish, 5 inches and 5% inches long,
caught on the western edge of Georges Bank Sept.
15, 1930, by the schooner Old Glory,6 are the only
ones that have been reported from the Gulf of
Maine. A third, 5 inches long, was taken by
Atlantis, south of Sable Island (lat. 40°55' N.,
long. 59°55' W.), August 18, 1941.

Figube 279. — Unicorn fish {Alutera scripta), Georges Bank specimen, 143 mm. long. Drawing by H. B. Bigelow.

THE PUFFERS AND PORCUPINE FISHES
FAMILIES TETRAODONTIDAE AND DIODONTIDAE

The members of these two families are so closely
allied one to the other, not only anatomically but
in general appearance, that they may be described
as a unit. They have only one dorsal fin (the soft-
rayed), the spiny dorsal being obsolete, and they
have no ventral fins. Their gill openings are
reduced to short slits like those of their allies, the

» Sumner, Osburn, and Cole, Bull. TJ. S. Bur. Fish., vol. 31, Pt. 2, 1913
p. 762.

* There Is no way to verify the identifications at this late date.

« Jordan and Evermann's (Bull. 47, U. S. Nat. Mus., Pt. 4, pi. 260, fig. 637)
illustration labeled "Scripta" seems to have been based on a specimen of
tchoepfi, in an intermediate stage of development.

triggerfishes and filefishes (pp. 520 and 521); their
teeth are fused into cutting plates; and they have
no scales. The two families are separable by the
structure of the teeth, as described below in the
accounts of the two species concerned, and by
certain anatomical characters.

All of them are capable of inflating their bellies
to balloonlike proportions with air or with water,
if annoyed, and of deflating at will. And it is a
matter of general interest (though not touching

• Reported by MacCoy, Bull. Boston Soc. Nat. Hist., No. 68, 1931 p. 18.

526

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

the Gulf of Maine directly) that the flesh of some
of the species of puffers, and perhaps of all of the
porcupine fishes, is poisonous.

Both groups are warm-water fishes. One species
of puffer reaches the southwestern part of the
Gulf rather commonly; and one porcupine fish has
been reported there as a stray from the south.

KEY TO GULF OF MAINE PUFFERS AND
PORCUPINE FISHES

1. Skin set with large conical spines Burrfish, p. 527

Skin merely prickly Puffer, p. 526

Puffer Sphaeroides maculatus (Bloch and
Schneider) 1801

Swellfish; Swell toad; Balloonfish;
Bellowsfish; Globefish

Jordan and Evermann, 1896-1900, p. 1733.

Description. — When the puffer is not inflated it
it moderately slender (about three times as long
as deep), about as thick as it is deep, and it tapers
from abreast the gill opening to a rather slender
caudal peduncle in one direction, to a rounded
snout in the other. Its very small mouth is
situated at the tip of the snout as it is in the
triggerfishes and filefishes. It has no true teeth
but the bones of its upper and lower jaws form
cutting edges, each divided in the middle by a
suture, giving the appearance of two large incisors
above and two below. The gill openings are very
small and set oblique, but their obliquity is the
reverse of that of the filefishes (p. 521), i. e., back-
ward and downward. The eyes are set very high
and are horizontally oval in outline. The skin has

no scales, but the sides of both head and body, the
back from snout to dorsal fin, and the belly as far
back as the vent are rough with small, stiff, close-
set prickles; those on the back are bluntish and
nearly vertical while those on the sides and belly
are rather sharp, pointing backward when the fish
is not inflated, but erect when it is.

There is no spiny dorsal fin. The soft dorsal fin
is very short (8 rays), rhomboid in outline, about
twice as high as it is long, and set far back close
to the caudal peduncle. The anal fin (7 rays) is
similar to the dorsal in shape and size, and arises
close behind it. The caudal fin is of moderate
size, weakly rounded, with angular corners. The
pectorals are fan-shaped, and are situated close
behind the gill openings. There are no ventral
fins.

The most interesting morphologic character of
the puffer is its ability to inflate itself with air or
with water if it is handled or at the slightest dis-
turbance of any sort, until the skin of the belly is
stretched tight as a football, and the fish is almost
globular. In this condition, it floats at the surface,
belly up, and apparently helpless. Leave it alone,
however, and it soon deflates, discharging the air
or water suddenly, and shrinks back to its normal
dimensions.

Color. — Dark olive green above, sometimes ashy
or dusky, the sides greenish yellow to orange,
crossbarred with 6 to 8 rather indefinite dark
bands or blotches. The belly is white.

Size. — The puffer is said to grow to a length of
14 inches, but few of them are more than 10 inches
long. Females average larger than males.

Habits. — The puffer is an inshore fish, often
coming in to the tide line. It runs up into slightly

M£M

^

'.'•;,'*''

Figure 280. — Puffer (Sphaeroides maculatus), Connecticut. From Jordan and Evermann. Drawing by W. S. Haines.

FISHES OF THE GULF OF MAINE

527

brackish water in various estuaries, and seldom is
caught more than a few fathoms deep, or more than
a mile or two from land. Throughout the north-
ern part of its range it belongs in the rather
numerous and varied category of "summer"
fishes, taken from April to November in Chesa-
peake Bay, from late May or early June to
October or early November along southern New
England. It is probable that when the puffers
disappear from their usual summer haunts, with
the onset of cold weather, they merely descend
into somewhat deeper water nearby, to spend the
winter on bottom in a more or less quiescent state.

Puffers feed on small crustaceans of all sorts
especially on crabs, shrimp, isopods and amphi-
pods, as well as on small mollusks, worms, barna-
cles, sea urchins, and other invertebrates, which
they find on bottom. Young fry of 7 to 10 mm.,
examined by Dr. Linton at Woods Hole, had eaten
copepods as well as crustacean and molluscan
larvae. And they are only too ready to take
bait, if the hook is small enough. Where they
are plentiful they may be nearly as much of a
nuisance in this way as the cunners.

Puffers spawn in shoal water close to the shore,
from mid-May, in Chesapeake Bay, and from
early June through the summer off southern
Massachusetts. And they are prolific. The ova-
ries of a Chesapeake Bay female, 10% inches long
contained (estimated) about 176,000 ova.7 The
eggs (about 0.9 mm. in diameter, with many
small oil globules) sink and stick fast to each
other or to whatever they chance to touch. In-
cubation occupies 3% to 5 days at a temperature of
about 67°-68° F. (20° C). The larvae are about
2.4 mm. long at hatching, and are brilliantly pig-
mented with red, orange, yellow, and black. In
3 days the mouth functions, and when they are 7
mm. long the young fish show most of the diag-
nostic characters of the adults,8 and can inflate
themselves even more, in fact, until the bulging
skin entirely hides the dorsal and anal fins.

General range. — Atlantic coast of the United
States from Florida to Cape Cod in abundance; to
Casco Bay in small numbers, and perhaps to the
Bay of Fundy as a stray.

Occurrence in the Gulj of Maine. — Anglers find
the puffer only too plentiful along the southern

' Hildebrand and Schroeder, Bull. U. S. Bur. Fish., vol. 43, Pt. 1, 1928,
p. 348.

■ Welsh and Breder (Zoologies, New York Zool. Soc., vol. 2, No. 12, Jan-
uary 1822, N. Y.) describe the early stages in the life history of the puffer.

shores of Massachusetts, but the elbow of Cape
Cod marks the eastern and northern limit to their
presence in any numbers. They have been re-
ported at Monomoy, Truro, and Provincetown.
Cape Cod Bay may perhaps support a small resi-
dent population, for Prof. A. E. Gross informs us
that he has seen as many as four or five taken at
one time in a pound net at Sandy Neck, Barn-
stable, at a tide, in the summer of 1920; besides
others stranded there on the beach.9 And we
have heard of others there recently, or nearby.
Storer described them as common at Nahant, a
few miles northeast of Boston, but this seems to
have been an error, for Wheatland (1852, p. 214)
writing about the same period, not only spoke of
them as seldom seen in Massachusetts Bay, but
considered a single specimen taken in Salem Har-
bor in the summer of 1848 as worthy of a note.
And this remained the only positive record for a
puffer for Essex County until August 24, 1920,
when one was caught at Gloucester.10

The only records of puffers north of Cape Ann
that have come to our notice are of two taken in a
trap in Casco Bay in. 1896, and of one taken near
Long Island, off Portland Harbor, Maine, on July
24, 1933. But there may be a small local popu-
lation in Casco Bay, and in the vicinity of Booth-
bay Harbor, Maine, for L. W. Scattergood lI writes
us that the pound net fishermen have long been
acquainted with them there and that he had
received three specimens recently from Pemaquid
Point where the fishermen report them as com-
monest in June. A skeleton, apparently of a
puffer, has been found on the shore of Minas
Basin, at the head of the Bay of Fundy on the
Nova Scotian side.12

Burrfish Chilomycterus schoepjii (Walbaum) 1792 13

Porcupinefish; Rabbitfish; Oysterfish
Jordan and Evermann, 189&-1900, p. 1748.

Description. — The burrfish resembles the puffer
(p. 526) in the positions of its dorsal and anal fins,
but its skin is armed with short, stout, triangular

• The Auk, vol. 40, 1923, p. 24.

u This specimen, reported by MacCoy (Bull. 67, Boston Soc. Nat. Hist.,
1933, p. 9) is in the Museum of Comparative Zoology.

» Letter dated September 19, 1951, U. S. Fish and Wildlife Service.

■ Reported to us (1951) by Dr. A. H. Lelm of the Fisheries Research
Board of Canada.

u Jordan, Evermann, and Clark place this species in the genus CpcliMhi/s •
Kaup 1855.

528

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 281. — Burrfish (Chilomycterus schoepfii), Connecticut. From Goode. Drawing by H. L. Todd.

spines instead of being merely prickly. These
spines are sparsely scattered all over the trunk,
with about 9 or 10 from nose to tail along any
given line. Furthermore, the burrfish is oval in
outline, not fusiform like the puffer; the openings
of its nostrils are prolonged in a single tubular
tentacle; its bony jaw plates are not divided by a
median suture as they are in the puffer, hence each
jaw apparently is armed with a single very broad
incisor tooth instead of with two; the pectoral
fins are not only much larger than in the puffer
but their upper edge is level with the upper corner
of the gill openings in the burrfish (considerable
below it in the puffers) ; its eye is round, not oval;
and its anal fin is below the dorsal, not behind
the latter. We need only add that the soft dorsal
and anal fins (it has no spiny dorsal) are both
rounded, each has 10 to 12 rays; the caudal fin
is very narrow and round-tipped; the pectorals
are much broader than long, and there are no
ventral fins.

Color. — The ground color varies from green to
olive or brownish above, with .pale belly, usually
tinted with yellow or orange. The back and sides
are irregularly striped with olive brown, dusky, or
black lines, running downward and backward,
roughly parallel one with another. There is a
dark blotch on each side at the base of the dorsal
fin, a smaller one between the latter and the anal
fin, one above the base of each pectoral fin, and a
fourth close behind the latter.

Size. — -Length, to about 10 inches.

General range. — Coast of the United States,
from Florida northward regularly about to New
York, occasionally to Cape Cod, and straying as
far as Massachusetts Bay; most plentiful from the
Carolinas southward.

Occurrence in the Gulf of Maine. — The only
record of this southern fish north of the elbow of
Cape Cod is of one taken in Massachusetts Bay
many years ago l* and another caught at West
Point, Maine, August 5, 1949. 16

THE OCEAN SUNFISHES OR HEADFISHES. FAMILY MOLIDAE

Although the ocean sunfishes are allied anatomi-
cally to the puffers and porcupine fishes, with
which they agree in the very small gill openings
and in the fusion of the teeth into a sort of bony
beak, they bear no resemblance whatever to them
in general appearance, for they appear to consist
of nothing but a "huge head to which the fins are
attached," as Jordan and Evermann I6 aptly
express it. They have no spiny dorsal fin; the
soft dorsal and anal fins are short and very high,
and they have no caudal peduncle. The caudal
fin, so short that it is apparently nothing more than

a flap of skin, extends all around the rear outline
of the trunk. Corresponding to their extraordi-
nary conformation the sunfishes have only 16 or 17
vertebrae.

All known members of the family are oceanic in
nature, and they are widely distributed in warm
seas. One (Mola mola, p. 529) is a rather frequent

" This specimen, reported by Kendall (Occas. Paper, Boston Soc. Nat.
Hist., vol. 7, Pt. 8, 1908, p. 118) Is (or was) in the collection of the Boston
Society of Natural History.

" Taken in a fish trap and reported by Scattergood, Trefethen, and Coffin
(Copela, 1951, No. 4, p. 298).

'• Bull. 47, V. S. Nat. Mus., Pt. 2, 1898, p. 1752.

FISHES OF THE GULF OF MAINE

529

visitor to our Gulf from the south; and a second
(Masturus lanceolatus) has, perhaps, a claim to
mention here, on the strength of one very young
suniish that was taken in Massachusetts Bay
many years ago (p. 532).

KEY TO GULF OF MAINE SUNFISHES

1. There is no evident caudal fin Sunfish, p. 529

2. There is an evident caudal fin, extending horizontally

across the posterior edge of the trunk, with a tri-
angular lobelike extension a little above the midlevel
of the body Sharp-tailed sunfish, p. 531

Sunfish Mola mola (Linnaeus) 1758
Jordan and Evermann, 1896-1900, p. 1753.

Figure 282. — Sunfish {Mola mola). From
Drawing by H. L. Todd.

Goode.

Description. — The oblong body of an adult sun-
fish (adults alone are seen regularly in the Gulf of
Maine) suggests the head and fore trunk of some
enormous fish cut off short, for it is truncate im-
mediately back of the dorsal and anal fins, and has
no caudal peduncle. But it tapers in front of the

fins toward the snout so that the forward half of
the trunk is oval in profile. The fish is less than
twice as long as deep, strongly flattened sidewise
(about one-fourth as thick as deep), with a very
small mouth at the tip of the snout; teeth com-
pletely united in each jaw; a very small eye in line
with the mouth; remarkably short gill openings,
and the nose overhangs the upper jaw as a kind
of rough, mobile wart or pad. The soft dorsal fin
(there is no spiny dorsal) stands over the anal
fin, close behind the midlength of the fish. Both
these fins are very much higher than long, tri-
angular, with sharply rounded tips, and each has
15 to 18 rays, with the seventh ray the longest.

The fins cannot be laid back, as they can in most
bony fishes; and the sunfish propels itself along by
waving them from side to side. The caudal fin
extends around the whole posterior margin of the
body. Confluent with the dorsal and anal fins
in the young and hardly separated from them in
the adult, it is so short and its rays so hidden by
the thick opaque skin that it looks more like a
fold of skin than a typical fin. Its general outline
is rounded, paralleling the rear outline of the body,
but its margin is scalloped, with a rounded bony
prominence or knob in line with each caudal ray
(11-14) and with a notch between every two of
these prominences. We have counted 11 such
notches on a fish 3)i feet long, and have record of
8 on one of about 4 feet.17 The pectoral fins are
small, rounded, each with 12 or 13 rays, and are
situated about halfway up the body close behind
the tiny gill openings. There are no ventral fins.
The skin is unusually thick (about 1% inches thick
in one 47 inches long which we harpooned near La
Have Bank on August 7, 1914), very tough and
elastic in texture; it is crisscrossed with low ridges,
and fins as well as trunk are clothed with small
bony tubercles, giving the appearance of shark
skin.

The sunfish is described as glowing luminescent
at night in the water. We cannot verify this first
hand. But we can bear witness that it grunts or
groans when hauled out of the water; that its skin
is covered with a thick layer of tough slime, and
it is the host of a great variety of parasites, ex-
ternal and internal, with copepods and trematodes
clinging to its skin and infesting its gills, with its
muscles harboring round worms and with various

" Taken near Boothbay, Maine, and reported to us by Dr. Austin F. Rlggs.

530

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

round worms and flat worms inhabiting its intes-
tines.

Color. — Dark gray above, the back with a
brownish cast, the sides paler with silvery re-
flections, the belly dusky to dirty white. Some
descriptions mention a broad blackish bar along
the bases of the dorsal and anal fins, but nothing
of the sort was to be seen in the only example we
have handled fresh from the water.

Size. — The sunfish grows to a great size.
Heilner 18 describes the capture of one 10 feet 11
inches long off Avalon (Calif.), while Jordan and
Evermann record another Californian specimen
8 feet 2 inches long, weighing about 1 ,800 pounds.

One measuring 8 feet in length and 1 1 feet from
tip of dorsal fin to tip of anal fin was exhibited in
London in 1883, 19 and an 8-foot specimen was
taken off Cape Lookout (N. C.) in 1904,20 but
large ones such as this are exceptional, the general
run being from 3 to 5 feet (rarely 6 feet) long, and
from 175 pounds to 500 pounds in weight. A fish
4% feet long is about 31 inches across the body and
6% feet from the tip of the dorsal fin to the tip of
the anal. One, 5 feet 3 inches long, was 4 feet 2
inches wide and 14% inches thick.21 A fish 4 feet
1 inch long, caught off Boston Harbor, August 14,
1922, weighed 516 pounds.22

Habits. — The sunfish is a wanderer of the high
seas, drifting at the mercy of the ocean currents;
those that are seen are at the surface (see following
for an exception); how deep they may descend is
not known.

When these unlucky vagrants are sighted in our
cool northern waters they have usually been
chilled into partial insensibility. They float
awash on the surface, feebly fanning with one or
the other fin, the personification of helplessness.
Usually they pay no attention to the approach of
a boat, but we have seen one come to life with
surprising suddenness and sound swiftly, sculling
with strong fin strokes, just before we came within
harpoon range. When one is struck it struggles
and thrashes vigorously while the tackle is being
slung to hoist it aboard, suggesting that they are
far more active in their native haunts than their

'• Bull. New York Zoo]. Soc, vol. 23, No. 6, November 1920, p. 126.
" Smltt, Scandanavian Fishes, vol. 1, 1892, p. 628.

» Smith, North Carolina Geological and Economic Survey, vol. 2, 1907.
p. 363.
■' As reported in the Boston American for June 24, 1930.
a Reported, with photograph, in the Boston Daily Post for August 14, 1922

feeble movements in fatally cold surroundings
might suggest.

The sunfish lives on an unusual diet, for as a
rule the contents of the stomach consists either of
jellyfish, ctenophores, or salpae, or of a slimy liquid
that probably represents the partially digested
remains of these. This has been true of all the
sunfish brought in to the Bureau of Fisheries at
Woods Hole. But various crustacean, molluscan,
hydroid, and serpent-star remains, even bits of
algae and eelgrass (Zostera), have been found in
sunfish stomachs in European waters, proving
that at times they either feed on the bottom in
shoal water, or among patches of floating weed.
And their jaws certainly seem fit for harder fare
than jellyfish.

The spawning habits are not known, nor have
the eggs been seen; presumably these are buoyant,
with many globules, as are those of the sharp-tailed
sunfish Masturus lanceolatus. The young fry differ
from their parents in being armed with 8 short
stout spines on either side, and with a single
median row of 4 spines along the back and 7 along
the ventral margin of the body.23

General range. — Oceanic and cosmopolitan in
tropical and temperate seas; known northward to
northern Norway on the European side of the
Atlantic, to the Newfoundland banks, the Gulf of
St. Lawrence, and the outer coast of Nova Scotia
on the American side.24

Occurrence in the Gulf of Maine.- — -The sunfish is
only a stray visitor to our Gulf, which it enters
now and then from the warmer and more congenial
waters outside the continental slope. There are
published records of its appearance in St. John
Harbor, New Brunswick, near Birch Harbor; near
Seguin Island; off Small Point; and off Cape Eliza-
beth (Maine), where it has been reported re-
peatedly; off Cape Ann; and from various localities
in Massachusetts Bay. Sunfish have even been

■ For a discussion of the young fry of the ocean sunflshes, with illustrations
and references to earlier accounts, see Schmidt, Meddel. Kommisslon
Havunders((gelser, Ser. Fiskeri, Denmark, vol. 6, 1921, No. 6.

•• Localities where sunfish had been reported in the Oulf of St. Lawrence
up to 1947 include north of Cape Breton; Bathurst, New Brunswick; North-
umberland Straits; the north shore of the Gaspe Peninsula; the south shore
of the Gulf opposite the Saguenay River; vicinity of Trois Pistoles; Anticosti;
and Bay of Islands on the west coast of Newfoundland. See Medcofl and
Schiffman (Acadian Naturalist, vol. 2, No. 7, 1947) for list with details.
Dunbar (Canad. Field Naturalist, vol. 64, No. 3, 1950, p. 124) has recently
reported one, 5 feet long, that was found on the beach at Metis on the southern
shore of the Lower St. Lawrence River. A Gulf of St. Lawrence record that
is especially interesting because so late In the season, is of one about 6 feet
long that stranded late in October 1, 1926, at Curling, Bay of Islands, west
coast of Newfoundland (reported in the Boston Traveler for Nov. 2, 1926).

FISHES OF THE GULF OF MAINE

531

seen in Boston Harbor, and on August 18, 1918,
one 4% feet long was killed in a narrow creek at
Quincy, Mass. The Grampus sighted sunfish near
the Isles of Shoals in 1896 (Dr. Kendall's field
notes), in 1912, and in 1914, as well as one in the
eastern basin of the Gulf in 1912. Seaside dwellers
reported one or two near Cape Porpoise in 1921;
one of 7 feet was caught off Boothbay, Maine, in
August 1927; and one 5 feet 3 inches long in the
northern side of Massachusetts Bay, off Bakers
Island, Beverly, in 1940, an especially interesting
case, for the fish in question was taken on a hand
line in 20 fathoms of water.28 And in 1950 several
blundered into one of the traps at Barnstable on
Cape Cod Bay.

An occasional sunfish is, in short, to be expected
anywhere in the western side of the Gulf and along
the coast of Maine. The only record, however, for
a sunfish in the Bay of Fundy is from near its
mouth at St. John Harbor.29 Nor do we find any
report of them along the Nova Scotian side of the
open Gulf of Maine.

In most summers it is something of an event to
see a sunfish anywhere in the inner part of the
Gulf. During July and August of 1912, for exam-
ple, we sighted only one from the Grampus, none
at all in August 1913, and only one in the Gulf
and another near La Have Bank during the mid
and late summer of 1914. They vary, however,
in numbers from year to year; 1928, for example,
was a year of abundance all along the coast, while
in 1950, a single trap at Barnstable on Cape Cod
Bay took 26 sunfish, an astonishing number.
Report also has it (we cannot verify this first-
hand) that sunfish are more plentiful over and
along the southern edge of Georges Bank than
they are within our Gulf, as indeed might be
expected from their oceanic origin.

In the inner parts of our Gulf sunfish are oftenest
sighted in mid or late summer, or early in autumn.
And one has been reported stranded in Bay of
Islands on the west coast of Newfoundland as late
as the end of October.27 But it is not likely that
any can survive the winter in our Gulf, or anywhere
along the coast to the northward. Neither is there

any reason to suppose that the waifs that visit
our Gulf ever spawn there.28

Sharp-tailed sunfish Masturus lanceolatus
(Lienard) 1840 29

Figure 283. — Sharp-tailed sunfish (Maslurus lanceolatus).
Above, adult, Miami, Florida, after Gudger. Below,
young, 54 mm., Massachusetts Bay, after Putnam.

11 Reported In the Boston American, June 24, 1930.
* Cox, Bull. Nat. Hist. Soc. New Brunswick, No. 13, art. 2, 1896, p. 75.
" Reported in the Boston Traveler, November 2, 1926, from the Associated
Press.

» Sunfish fry, about 2 inches long, taken in Massachusetts Bay many years
ago and reported by Putnam (Proc. Amer. Assoc. Advancement of Science,
19th Meeting (1870), 1871, pp. 265, 256, fig. 3) as this species, actually be-
longed to the closely allied sharptailed sunfish (p. 632), as shown by Schmidt
(Meddel. Kommiss. Havundersjtgelser, Denmark, Ser. Fiskeri, vol. 6, 1921,
Pt. 6, p. 6), and by Gudger (Proc. Zool. Soc. London, 1937, Ser A, p. 382).

» It Is an open question still, whether specimens with longer tail fins and
others with shorter tall fins represent two separate species, or whether the
differences between them are sexual ones. See Fraser-Brunner (Bull.
British Mus. Nat. Hist., vol. 1, No. 6, 1951, p. 105) for a recent discussion of
this subject.

532

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Description.- — The sharp-tailed sunfish differs
from the common sunfish (Mola mola) chiefly in
the fact that the rear margin of its body is edged
by a short but evident caudal fin of 18-20 soft rays,
that extends around from close behind the dorsal
fin to close behind the anal fin, with a triangular
lobelike, blunt-tipped projection a little above the
midlevel of the body. Its scales, too, are much
finer and less evident to the touch than those of
Mola, and its skin is less slimy. It resembles Mola
very closely in all other respects.

Color. — Described as with the whole trunk more
or less silvery, the upper parts of the sides grayish
brown to blackish, the lower parts paler; the sides
either plain or variously marked with ill-defined
dark spots; the dorsal and anal fins as dark slaty,
the caudal fin as sometimes with pale blotches.

Size. — This sunfish appears to grow as large as
the more common Mola, perhaps even larger. In
a Florida specimen, 88 inches long (after being
dried somewhat) the tail fin occupied 21 inches,
the body occupied 67 inches and was 38 inches
high.30 The dimensions of a North Carolina
specimen 73 % inches long were: body 54 % inches
long by 37 inches high and 1 1 inches thick, tail fin
19 inches long, dorsal fin 27 inches high, anal fin 25
inches high.31

Habits. — Nothing is known of its habits to dif-
ferentiate it from its more common relative.

General range. — This sunfish, like Mola, appears
to be cosmopolitan in tropical-warm temperate
latitudes, oceanic in nature but coming close in-
shore on occasion, and even into estuarine situ-
ations. Adults have been reported from Japan,
the Hawaiian Islands, Polynesia, Amboina, and
Mauritius in the Indian and Pacific Oceans; from
the Red Sea; from South Africa (Table Bay);
from Madeira, from near the Azores; near Habana,
Cuba (7 specimens); east coast of Florida (9
specimens), and North Carolina (4 specimens) in
the Atlantic. Young fry have been taken off the
Azores; in the Sargasso Sea; west of the Canaries;
in the Caribbean; and in the Gulf of Mexico.32

Occurrence in the Gulf of Maine. — The only
record for the sharp-tailed sunfish for our Gulf is
for 4 young fry, about 2 inches long, that were
taken many years ago in Massachusetts Bay.
These were originally reported by Putnam 33 as-
the young of the common ocean sunfish, but
Schmidt 34 and Gudger 35 have shown that they
were the sharp-tailed species in reality, because
with projecting caudal fins. The nearest locality
record for an adult of the species (to date) is for
Pamlico Sound, N. C. But it would not be
astonishing if one were to drift farther northward
any summer, as so many stray species do from
the south.

THE ANGLERS. FAMILY LOPHIIDAE

This family is the only familiar Gulf of Maine
representative of the small but anatomically re-
markable tribe of pediculate fishes, in which the
base of the pectoral fin takes the form of an arm
("pseudo-brachium") formed by the elongation of
the carpal bones ("actinosts"), which are so short
in all other bony fishes that they are not noticeable
externally. Coupled with this peculiar structure
of the pectorals, the gill openings are reduced to
small apertures in or near the axils ("armpits")
of these fins. The anglers are characterized among
their immediate relatives by a very large and very
much flattened head; by an enormous mouth; and
by the fact that they have only two bones in each
pectoral "arm." The Gulf of Maine harbors one
species.

American goosefish Lophius americanus Cuvier
and Valenciennes 1837

Monkfish; Angler; Allmouth; Molligut;
Fishing frog

Jordan and Evermann, 1896-1900, Lophius piscatorius
Linnaeus 1766 in part, p. 2713.

Description. — The goosefish is so unlike all
other Gulf of Maine fishes that there is no danger
of mistaking it for any other once it is seen. It
is so much flattened, dorso-ventrally, and so soft
in texture that when one is left stranded on the

■ Hubbs and Giovanneli, Copela, 1931, pp. 135-136.

>' Brimley, Jour. Elisha Mitchell Sci. Soc, vol. 55, 1939, p. 295.

•> See Gudger (Proc. Zool. Soc. London, 1937, Ser. A, p. 353), for list of
locality records up to 1937 with references; Brimley (Jour.. Elisha Mitchell
Sci. Soc, vol. 55, 1939, p. 295) for account of North Carolina specimens.

» Proc. Amer. Assoc. Advancement of Science, 19th meeting, Troy.N. Y.
(1870) 1871, pp. 255-256.

»* Meddel. Komm. Havunders0gelser, Denmark, Ser. Fiskeri, vol. 6, Pt. 6,
1921, p. 6.

»» Proc. Zool. Soc. London, 1937, Ser. A, p. 382.

FISHES OF THE GULF OF MAINE

533

Figure 284. — Goosefish (Lophius americanus), oblique-dorsal view, Gulf of Maine specimen. From Bigelow andJWelsh.

shore it collapses until it is hardly thicker than a
skate. Its head is rounded as seen from above,
about as broad as it is long, and enormous in
comparison with the body, which is so narrow
and tapering, back of the pectoral fins as to give
the fish a tadpole-like appearance. The most
noticeable feature is the enormous mouth, which
is directed upward, with the lower jaw projecting
so far beyond the upper jaw that most of the
lower teeth are freely exposed even when the
mouth is closed.

Both jaws are armed with long, slender, curved
teeth, all alike in form but of various sizes, and
very sharp, and all the teeth point inward toward
the gullet. Some of them may be as much as an
inch long in a large fish. The teeth in the lower
jaw are in 1 to 3 rows, mostly large, while in the
upper jaw the few teeth in the middle (there is a
toothless space in the midline) are largest, with a
single row of smaller ones flanking them. And
there are several rows of thorn-like teeth on the
roof of the mouth. The gill openings are behind
the pectoral fins and they lack the gill covers that
are to be seen in most of the bony fishes. The
eyes are on the top of the head, and are directed
upward.

The pectoral fins are exceedingly distinctive, for
their bases have the form of thick fleshy arms as
already described (p. 532) that bear the fins proper
at their outer edge. The finlike parts are fanlike
when spread, and so thick-skinned that the rays
are hardly visible except in the scalloping of the
margins.

The top of the head bears 3 stiff slender spines
(representing the anterior part of the spiny dorsal

fin) hardly thicker than bristles, the first standing
close behind the tip of the snout, the second a
little in front of the eyes, the third on the nape of
the neck. And while the first and second are
movable from recumbent to erect, the third slopes
backward with its basal half imbedded in the
skin.36 The relative lengths of these spines vary,
but the first two have been about equal in length
on most of the fish we have seen, or the second a
little the longer, with the third much the shortest
of the three. The first spine bears an irregular
leaflike flap of skin at its tip, which plays an
important role in the daily life of the goosefish as
a lure for its prey (p. 536) while the second and
third spines have small triangular membranes at
their bases, and one or both of them may be
fringed with short lobes of skin. Besides these
spines there are two well-developed dorsal fins,
the first (of 3 spines) situated over the pectorals
and the second (10 to 11 rays) on the rear part of
the trunk.

The single anal fin (9 to 10 rays) stands below
the second dorsal fin, and the ventral fins (about
5 thick rays) are situated on the lower surface of
the head, well in front of the pectorals. The
caudal fin is small and broom shaped. The dorsal
fins have thin delicate membranes. But the
caudal, anal, and ventral fins are thick and fleshy,
like the pectorals. The skin is scaleless, very
smooth and slippery to the touch, and there is a
row of fleshy flaps of irregular shape running
around the margin of the head and around the
edge of the lower jaw, besides smaller tags that

«• Sometimes more than one-half is so imbedded.

534

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

fringe the sides of the trunk as far back as the
base of the caudal fin. Furthermore, the upper
side of the head bears several low conical tubercles
which vary in prominence from fish to fish.

Color. — The many goosefish we have seen (and
this corroborates the published accounts) have
been chocolate brown above, variously and finely
mottled with pale and dark. The dorsal fins, the
upper sides of the pectoral fins, and the caudal fin
are of a darker shade of the same color as the back,
except nearly black at the tips, while the whole
lower surface of the fish is white or dirty white.
Sometimes, it is said, the upper side is dotted with
white spots but we have seen none that were
marked in that way. Very small ones are de-
scribed as mottled and speckled with green and
brown. Wilson, who watched many in the
aquarium at Plymouth, England,37 writes that the
European species is able to match both its color
and its color pattern closely to the sand and
gravel on which it lies.

Size. — Adults run from 2 to 4 feet long,38 weigh-
ing up to 50 pounds, and heavier ones have been
reported. One 38 inches long, caught at Woods
Hole on July 25, 1923, weighed 32 pounds alive.

Remarks. — The goosefish of eastern North
America has usually been thought identical with
the widespread eastern Atlantic angler (i. pisca-
torius Linnaeus. But as Taning 39 has pointed
out, the late larval stages of our fish do not
resemble those of L. piseatorius as closely as they
do those of the angler of the Mediterranean and
of neighboring parts of the Atlantic that various
authors regard as a separate species, L. budegassa,
Spinosa 1807.40 This suggests that the goosefish
of the western Atlantic is a distinct species, for
which Berrill 41 has revived the old name L.
americanus Cuvier and Valenciennes 1837.

The adults of the three forms in question cer-
tainly resemble one another so closely that we
have not found any external differences that seem
significant to separate Gulf of Maine fish from
two specimens from northern Europe, and others

" Jour. Marine Biol. Assoc. United Kingdom, vol. 21, Pt. 2, 1937, p. 485.

" Rumor has it that goosefish grow to 6 feet, but we find none definitely
recorded (and have seen none) longer than 4 feet.

» Kept. Danish Oceanogr. Expeds. (1908-1910), No. 7, vol. 2, Biol. A. 10.,
1923, p. 7-16.

« See Regan, Ann. Mag. Nat. Hist. Ser. 7, vol. 11, 1903, p. 283 for descrip-
tions of L. piscatorius, of L. budegassa, and of a new species, vailbmti, described
by bim from the Azores and from the Cape Verde Islands.

" Oontrib. Canadian Biol. N. Ser., vol. 4, No. 12, 1929.

from the Mediterranean (all of about the same
size) with which we have compared them. But
it seems wisest to retain the separate name for
our form until the larval differences can be investi-
gated further (which we are not in a position to
do), and until much larger series of grown fish
have been compared.

Habits. — The depth range inhabited by the
goosefish extends from tide line down to at least
365 fathoms on the continental slope off southern
New England, and very likely deeper still. The
adults appear, for the most part, to hold to the
sea floor, where many are taken by the otter
trawlers. And they are found indifferently on
hard sand, on pebbly bottom, on the gravel, sand,
and broken shells of the good fishing grounds, and
on soft mud, where we have trawled them in the
deep basin of our Gulf.

Specimens of the closely allied European goose-
fish kept in the aquarium at Plymouth, England,
spent most of the time resting quietly.42 When
they swam they did so slowly, and they used their
paired fins for walking on the bottom. Wilson
describes one as digging a small hollow in the
bottom when it settled down, using its pelvic fins
to shovel the sand and pebbles forwards-outwards,
and using its pectorals, almost like webbed hands,
to push the sand away to either side until its back
was almost flush with the surrounding bottom.
But the fact that goosefish have been known to
seize and swallow hooked fish as the latter were
being hauled up, and even to capture sea birds
sitting on the surface, proves that while they
ordinarily snap up their prey from ambush, or by a
sudden short rush, they may make considerable
excursions for a meal on occasion.

The American goosefish is at home through a
very wide range of temperature. They have been
trawled on the Newfoundland banks in water as
cold as 320,43 and it is likely that those living
shoalest in the Gulf of St. Lawrence are exposed to
equally low temperatures, in late winter and in
spring. But we doubt if they can survive much
colder water, for many were seen floating dead in
Narragansett Bay, and on the shore, during the

« Wilson (Jour. Mar. Biol. Assoc. United Kingdom, vol. 21, Pt. 2, 1937,
pp. 486-490) has given a very interesting account of the habits of specimens in
the aquarium at Plymouth, England, where some were kept for as long as
11 months.

« Rept., vol. 2, No. 1, Newfoundland Fish. Res. Coram, 1933, p. 125
sta. 97.

FISHES OF THE GULF OF MAINE

535

winter of 1904-1905, apparently killed by the
unusually severe cold.44

At the other extreme, goosefish picked up by net
fishermen near Cape Lookout, N. C, in shoal
water (p. 540) are exposed to temperatures higher
than 70° for part of the season, perhaps as high as
75°. But reports 4S that the inshore contingent of
the goosefish population of Rhode Island waters
works out (i. e., deeper) in July, to work inshore
again in October suggest that they tend to avoid
extreme summer heat, if they can do so by de-
scending into deeper water.

They are tolerant to a wide range of salinity
also, occurring as they do from estuarine situa-
tions out to the upper part of the continental
slope. But we have never heard of one in brack-
ish water.

The larvae of the goosefish, like those of most
sea fishes, feed on various small pelagic animals
such as copepods, crustacean larvae, and glass
worms (Sagitta); and Sagitta is the chief diet of
young goosefish in the Adriatic during the life of
the latter near the surface, hence may serve this
same purpose in the Gulf of Maine.

The goosefish becomes a fish eater in the main
after it takes to the bottom, and the following
Gulf of Maine species have been recorded from its
stomach: spiny dogfish, skates of various kinds,
eels, launce, herring, alewives, menhaden, smelts,
mackerel, weakfish, dinners, tautog, sea bass,
butterfish, puffers, various sculpins, sea ravens,
sea robins, sea snails, silver hake, tomcod, cod,
haddock, hake, witch flounders, American dab,
yellowtail flounders, winter flounders, and various
other species of flatfish unnamed, as well as its own
kind. The goosefish often captures sea birds, as
one of the vernacular names implies, cormorants,
herring gulls, widgeons, scoters, loons, guillemots,
and razor-billed auks are on its recorded dietary,
while we have found grebes and other diving fowl,
such as scaup ducks and mergansers, in goosefish
in Pamlico Sound, N. C. It is questionable,
however, whether even the largest of them would
be able to master a live goose, as rumor has it, nor
do the local fishermen believe it ever does so in
Pamlico Sound, though the abundance of wild
geese there in winter would afford it every op-

« Reported by Tracy, 36th Eept. Comm. Inland Fish. Rhode Island,
1906, p. 92.
" Tracy, 36 Rept. Comm. Inland Fish., Rhode Island 1906, p. 92.

portunity. Goode,46 however, tells of one which
a fisherman saw struggling with a loon. Even a
sea turtle has been found in one.47

Goosefish are also known to devour invertebrates
such as lobsters, crabs of several species, hermit
crabs, squids, annelid worms, shellfish, starfish,
sand dollars, and even eelgrass. Linton's 48 report
of one that was full of mud containing small shell-
fish, crustaceans, and worms is interesting. In
short, nothing edible that strays within reach
comes amiss to a goosefish. And examinations of
stomachs have shown that the relative importance
of various articles in its diet varies widely on
different grounds, depending on what is available.
Thus Field 49 found skates, flounders, and squid
their chief dependence near Woods Hole. The
32-pounder from there, mentioned above, con-
tained 2 menhaden, 1 spiny dogfish a foot long,
and the vertebral columns of 6 others; while goose-
fish diet largely on hakes in the Bay of Fundy; 60
on haddock, flatfish, and on skates on Georges
Bank.

The goosefish has often been cited for its re-
markable appetite. We read, for instance, of one
that had made a meal of 21 flounders and 1 dog-
fish, all of marketable size; of half a pailful of
cunners, tomcod, and sea bass in another; of 75
herring in a third; and of one that had taken 7
wild ducks at one meal. In fact it is nothing
unusual for one to contain at one time a mass of
food half as heavy as the fish itself. And with its
enormous mouth (one 3% feet long gapes about 9
inches horizontally and 8 inches vertically) it is
able to swallow fish of almost its own size. Ful-
ton, for instance, found a codling 23 inches long
in a British goosefish of only 26 inches, while
Field took a winter flounder almost as big as its
captor from an American specimen. One that we
once gaffed at the surface, on Nantucket Shoals,
contained a haddock 31 inches long, weighing 12
pounds, while Captain Atwood long ago described
seeing one attempting to swallow another as large
as itself. Wilson's observations, however, indi-
cate that they are no more gluttonous than any
other rapacious fish, for those that he watched in
the aquarium usually refused food for 2 or 3 days

« Fish. Ind. U. S., Sect. 1, 1884, p. 174.

« Schroeder, Copeia 1947, p. 201.

« Bull. U. 8. Bur. Fish., vol. 19, 1901, p. 487.

" Rept. U. S. Comm. Fish., (1906) 1907, Doc. No. 622, p. 39.

» Connelly, Bull. 3, Biol. Board Canada, 1920, p. 16.

536

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

after a meal. His observation that they evidently
preferred small fish is in line with their normal
habits, for they feed mostly on small fish, not on
large, and even the largest of them take very small
fry on occasion.

In Scottish waters,51 where the habits of the
local goosefish are better known than in the Gulf
of Maine, their local abundance depends on the
supply of small fish. And despite their poor
ability as swimmers goosefish have been found to
congregate near particular shoals of herring.

Goosefish, like most fish of prey, often swallow
indigestible objects. They have even been
credited (on how good evidence we cannot say)
with pouching lobster-pot buoys. And the story
of one whose mouth made a holding ground for
the anchor of a small boat has been related
repeatedly.

The most interesting habit of the goosefish is
that it actually does use the flap of skin at the
tip of its first dorsal spine as a bait to lure small
fishes within seizing distance, much as Aristotle
described. W. F. Clapp (only observer who has
watched the American goosefish feeding, to our
knowledge) has described them to us, in Duxbury
Harbor as lying motionless among the eelgrass,
with the "bait" at the tip of the first dorsal ray
swaying to and fro over the mouth. When a
tomcod (the only fish he saw them take) chances
to approach, it usually swims close up to the
"bait," but never (in his observation) actually
touches the latter, for the goosefish opens its vast
mouth as soon as the victim comes within a few
inches and closes it again, engulfing its prey
instantaneously.

Further details added by observations on Euro-
pean anglers in aquaria at Port Erin, Isle of Man,
by Chadwick,52 and at Plymouth, England, by
Wilson,53 are that the first dorsal spine, with its
terminal "bait" is held down along the top of the
head, to be raised at the approach of a prospective
victim; that the bait may be jerked to and fro
quite actively in front of its owner's head; that
the victim is usually taken in head first; that a
fish swimming close enough may be snapped up
without the bait being brought into play; and
that some anglers use the bait often, others seldom.

Wilson also made the interesting observation that
touching the "bait" does not cause a reflex snap-
ping of the jaws, showing that the angler feeds by
sight.54

Adult goosefish cannot have many enemies.
But small ones are no doubt picked up by various
predaceous fishes. And Lebour's observation M
that goosefish larvae in aquarium jars at Ply-
mouth, England, were devoured by the larvae of
the spiny lobster (Palinurus), by large copepods,
by ctenophores, and by hydroids when they came
close enough to the walls of the jar to be seized
by the latter, is an interesting illustration of the
hazards that larval fishes meet during their free-
drifting stages.

Goosefish spawn in spring, summer, and early
autumn, according to the latitude, and through a
long season. Eggs and larvae have been taken
near Cape Lookout, N. C, in March and April;5*
in May off Cape Hatteras;57 and as early as May
at Woods Hole. But spawning may not com-
mence until early summer in the Gulf of Maine,
for June 24 (Passamaquoddy Bay M) is the earliest
date when eggs have been seen north of Cape Cod.
September 18 (off Seguin Island, Maine) is the
latest recorded date for American waters.

The floating egg-veils of the European angler
have been reported as early as February 18 in
Scottish waters and as late as July 23, 59 while
Taning M concludes from the sizes of larvae taken
at different dates that March-June is the season
of chief production to the west and southwest of
the British Isles in general. In the Mediterranean
(with higher temperatures), anglers spawn from
December and January on, as shown by the pres-
ence of larvae.61

The locality of spawning has been the subject of
discussion, whether inshore in shoal water, or
offshore in deeper. The egg veils reported from
the Bay of Fundy by Connolly; 62 from Passama-

'■ Fulton (Ann. Report, Fish. Board Scotland, (1902) 1903, Pt. 3, p. 195)
lists the stomach contents of 541 goosefish from various localities off Scotland)
•» Nature, vol. 124, 1929, p. 337.
» Jour. Marine Biol. Assoc. United Kingdom, vol. 21, pt. 2, 1937, p. 479,

M Oudger (Amer. Naturalist, vol. 79, 1945, p. 542), has given an interesting
and readable survey of observations, at various hands, on the use of the bait.

« Jour. Mar. Biol. Assoc. United Kingdom, vol. 13, 1925, p. 728.

« Information supplied by the late S. F. Hildebrand.

>> Taning, Rept. Danish Oceanogr. Expeds., 1908-1910, No. 7, vol. 2 (Biol),
A 10, 1923, p. 2B.

« Connolly, Contrib. Canadian Biol. (1921) 1922, p. 116.

•< See Bowman (Fish. Bd. Scotland, Scl. Invest. (1919, No. 2), 1920, p. 23)
for records for angler eggs up to 1919.

« Danish Oceanogr. Eipeds., 1908-1910, No. 7, vol. 2 (Biol.), A 10, 1923,
p. 23.

•> See Stiasny (Arbeit. Zoo!. Inst. Vienna, vol. 19, 1911, p. 70) for Mediter
ranean records, besides which an egg veil has been reported In January near
Naples by Le Bianco (Mltth. Zool. Stat. Neapel., vol. 19, pt. 4, 1909. p. 725)

« Contrib. Canadian Biol. (1921), No. 7, 1922, p. 116.

FISHES OF THE GULF OF MAINE

537

quoddy Bay by Berrill; 63 and from Frenchman
Bay near Mount Desert by Procter and others,64
were in such early stages of incubation tbat they
must have been spawned close at hand. And this
also applies to some isolated eggs that were col-
lected at about the 20-fathom contour line off
northern North Carolina,66 by the Dana. Neither
is there any reason to suppose that veils farther
advanced in incubation, that have been taken in
the inner parts of the Gulf of Maine (p. 541); at
Woods Hole; and at Newport (p. 537), had come
from any great distance. Furthermore, large
adult fish are present in abundance inshore
throughout the spawning season, which would
hardly be the case if they moved offshore or into
deep water to spawn. On the other hand, veils
that could not have been spawned long before
have also been met with near the 1,000 to 1,100
fathom (2,000-meter) contour line over the conti-
nental slope off North Carolina 66 and at about the
same relative position over the slope south of the
Newfoundland Banks.67

It appears, in short, that the American goosefish
spawns indifferently in shoal water and in deep.
It differs in this respect from its European relative,
which moves offshore and down the slope for the
purpose, to near the 1,000-fathom contour, to
judge from the localities where the newly hatched
larvae have been collected in the eastern North
Atlantic.68

The presence of egg veils off North Carolina;
near Newport 69 and near Woods Hole along south-
ern New England; in the Gulf of Maine (p. 541);
and over the continental slope south of the New-
foundland Bank; with the capture of a very small
(4-inch) specimen on the Grand Bank (p. 540)
shows that the American goosefish breeds through-
out its geographic range.

The eggs are shed in remarkable ribband-like
veils of mucus, each probably the product of a

« Contrib. Canadian Biol. Fish., N. Ser., vol. 4, No. 12, 1929, p. 145.

" Biol. Surv. Mount Desert Region, Pt. 2, Fishes, 1928, p. 3.

"Taning, Rept. Danish Oceanogr. Expeds., 1908-1910, No. 7, vol. 2
(Biol.) A. 10, 1923, p. 25.

« Lat. 36°16' N., long. 74°33' W., see Taning, Danish Oceanogr. Expeds.,
1908-1910, vol. 2 (Biol.), A 10, 1923, p. 25.

■ Murray and Hjort, Depths of the Ocean, 1912, p. 108.

" For further discussion, see Bowman (Fishery Bd. Scotland Sci. Invest.
[1919], No. 2, 1920, p. 21) and Taning (Rept. Danish Oceanogr. Expeds.,
1908-1910, vol. 2 (Biol). No. 7, A 10, 1923).

•> It was at Newport that Agassiz, and Agassiz and Whitman, collected the
veils and the larvae on which they based their accounts.

single ovary, up to 25-36 feet long, and said some-
times to be as much as 2 to 3 feet broad, in which
the eggs are arranged in a single layer, lying one to
three or even four in separate hexagonal compart-
ments, with the oil globule uppermost. In an
egg veil found near St. Andrews, New Brunswick,
between 32 and 36 feet long, about 8 inches wide,
about % inch thick, and about 25 quarts (26K
liters) in volume, about 5 percent of the eggs were
single, about 80 percent were in pairs, and about
5 percent were in threes, per compartment. This
veil was estimated to contain about 1,320,000
eggs,70 and Fulton estimated about the same num-
bers (1,345,848 and 1,317,587) in the ovaries of
two in Scottish waters.71

The veils are light violet gray or purplish brown,
made more or less blackish by the embryonic
pigment of the eggs according to the stage of de-
velopment attained by the latter. And they are
so conspicuous when floating at the surface that
fishermen have long been familiar with them,
though it was not until about 1871 that Alexander
Agassiz demonstrated their true parentage.72 The
eggs occasionally become isolated, perhaps when a
storm shreds the mucous veil to pieces, and they
float like any ordinary buoyant fish eggs when this
happens. We have not actually found them in
this condition in the Gulf of Maine, but Agassiz
and Whitman saw isolated eggs at Newport, and
Taning has reported others from North Carolina
waters.

The eggs themselves, large numbers of which
have now been examined, are 1.61 to 1.84 mm. in
longest diameter in the case of the American
species, as they lie in their mucous compart-
ments.73 The yolk is straw-colored, and they have
either one copper-colored or pinkish oil globule
of 0.4 to 0.56 mm., or several smaller ones. Incu-
bation proceeds normally at temperatures from as
low as 41° to as high as 63°-64°, and probably in
higher temperatures. The larvae, which float
with the yolk uppermost at first, have been re-

'» Berrill, Contrib. Canad. Biol, and Fish., N. Ser., vol. 4, No. 12, 1929, pp.
145, 147.

" Sixteenth Annual Report, Fish. Bd. Scotland (1897) 1898, Pt. Ill, pp.
125-134, pis. 2-3.

" Baird, American Naturalist, vol. 5, 1871, pp. 785-786.

T> The eggs of the European L. piscaiorius are described as larger, averaging
about 2.3 mm.

210941—53 — —85

538

FISHERY BULLETIN OF THE FISH AND "WILDLIFE SERVICE

ported as from about 2.5 mm. to about 4.5 mm.
long when they hatch.7*

The first of the dorsal fin rays (which is to form
the second head spine of the adult) appears within

" Larval goosefish from New England, from the Bay of Fundy, and from
Nova Scotian waters have been described and pictured by Agassiz (Proc.
Amer. Acad. Arts, Scl., N. Ser., vol. 9, 1882, p. 280); by Agassiz and Whitman
(Mem. Mus. Comp. Zoo]., vol. 14, No. 1, Ft. 1, 1885, p. 16, pi. 6, figs. 1-10);
by Connolly (Contrib. Canadian Biol. [1921], No. 7, 1922); by Procter and
others (Biol. Surv. Mt. Desert region, Pt. 2, Fishes, 1928); and by Berrill
(Contrib. Canadian Biol, and Fish., N. Ser., vol. 4, No. 12, 1929, pp. 145-149).
For accounts and illustrations of North European and Mediterranean
Lophlus larvae, see especially Lebour (Jour. Marine Biol. Assoc. United
Kingdom, vol. 13, No. 3, 1925, pp. 721-728) who reared them from the eggs at
Plymouth, England; also Bowman (Fishery Board for Scotland, Scl. Invest.
[19191 1920, No. 11), Stiasny (Arbeit, Zool. Inst. Vienna, vol. 19, 1911, p. 71),
and Taning (Report, Danish Oceanogr. Expeds., 1908-1910, No. 7, vol. 2
(Biol.). A 10, 1923).

^A

"%

4 days or so after hatching, as a lobe at the margin
of the embryonic finfold on the nape of the neck.
The pectorals are formed at about 7 days, when
the larva is 5.5 mm. long, the ventral fins have now
appeared as two long conical processes below and
behind the pectorals (fig. 285D) ; and the pigment
has become congregated in 3 or 4 masses behind
the vent, the last being a very conspicuous feature
that the larvae of the European species L. pisca-
torius do not share. The yolk has been absorbed
at a length of 6-8 mm., a second dorsal ray has
formed behind the first, and the ventral fins have
become 2-rayed. The third and fourth dorsal rays
or filaments appear while there are still only two

Figure 285. — Goosefish (Lophius americanus). A, eggs in veil; B, egg with advanced embryo; C, larva, about 5 days
old; D, larva about 12 days old; E, older larva; F, larva, 30 mm. A, E, and F, after A. Agassiz. B, C, and D,
from New England.

FISHES OF THE GULF OF MAINE

539

ventral rays in the American goosefish (fig. 285E),
but they do not do so in the European species
until the third rays have developed, in the ventral
fins. The Mediterranean larvae so far described
have agreed with the American in this respect,
which makes the situation puzzling.

A fifth dorsal ray next appears behind those
that have developed already, and a sixth in front
of these, all of them being interconnected with
membrane at their bases but free at their tips.
The pectoral fins assume a great breadth and
fanlike outline; the second dorsal, the anal, and
the caudal fins take definite form; the ventral rays
become filamentous at their tips, streaming far
out behind the tail; and a complete row of teeth
appears in the lower jaw, with a few in the upper.
The goosefish pictured at this stage by Agassiz
(fig. 285F) was 30 mm. long, and one much like
it taken off Brazil Rock, described by Connolly,
was 27 mm. long, but the larvae of the Medi-
terranean goosefish attain this stage when they
are only 13 to 18 mm. long, according to Stiasny.

The older post-larval stages of the American
goosefish have not been seen yet. But develop-
ment no doubt follows the same course for them
as it does for the Mediterranean form; i. e., the
foremost dorsal ray becomes bristlelike with the
flap appearing at its tip; the last three of the free
rays on the nape of the neck join together as the
future first dorsal fin; the lappets of skin appear
around the margin of the lower jaw and along the
cheeks; and the head broadens and flattens while
the young fish are still living pelagic, with enor-
mous pectoral fins and with threadlike ventrals
(fig. 286).

Figure 286. — Goosefish (Lophius). Larva, Mediterra-
nean, 50 mm. After Stiasny.

The largest free-swimming Mediterranean larva
seen by Stiasny was 2 inches (50 mm.) long.
Probably the young take to the ground shortly

after this stage, for Bowman describes European
goosefish fry of about 2% inches (65 mm.) that
were trawled on the bottom, off Scotland, as of
adult form, except that their pectorals were pro-
portionately larger. To attain this state entails
growth on the part of the head out of proportion
to the rest of the body; enlargement of the mouth;
shrinkage of all the fins (of the ventrals most of
all) ; alteration of the second and third free dorsal
rays into spines (they are soft previously); and a
general flattening of the whole fish. Young of 3
inches taken at Halifax, one of 4% inches from
Campobello (both pictured by Connolly), and
others as small as 4— i% inches that we have
trawled, were at about this stage in their develop-
ment.

The capture of a 2K-inch specimen in October
(presumably), in Halifax Harbor, and of another
of 3 inches there (date not recorded),75 suggests
that the goosefish may be expected to reach about
that length by the onset of their first winter in
our northern waters. One 4% inches long from
Halifax, studied by Connolly, seemed, from the
thickness of its otoliths, to have been in its second
summer or autumn, i. e., one full year old, which
probably applies to three others of 4-4% inches,
trawled in August, that we have seen. But it is
not clear whether 14 others of 7% inches in May,
and of 6%-9 inches in July, were early hatched
fry in their second season, or late hatched speci-
mens in their third season.76

One of the larger fish studied by Connolly
showed 4 concentric rings in its vertebrae; one
31 inches long seemed to have 9 rings; one of 37
inches seemed to have 10 rings; and one of 40
inches seemed to have 12 rings. But it is not
certain whether these vertebral rings are laid down
regularly, one per year, or not.

Fulton's observations " show that the fry of the
North European angler may be 5-5% inches long
by November off Scotland, where spawning com-
mences in March or earlier; which is as large as
the fry of the American species are in their second
summer in our northern waters, where the first
growing season is at least 3 to 4 months shorter.
Fulton's measurements also point to more rapid
growth by the larger Scottish fish than by the

» Connolly (Contrib. Canadian Biol. (1921), No. 7, 1922, pp. 119-120).
n Equal uncertainty applies to two of 10 inches, one of which was trawled
In February, the other in April.

" 21st Ann. Kept. Fish. Bd. Scotland, Pt. 3 (1902) 1903, pp. 190-194.

540

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

American goosefish in the Bay of Fundy, namely
to 9-16 inches at a year and a half; to 14%-18%
inches when 2% years old; and to about 21 inches
at 3 years of age.

Few goosefish mature on either side of the
Atlantic until they are 30 inches long, or longer.78

General range. — Coast of eastern North America
from the southern and eastern parts of the Grand
Banks of Newfoundland, and the northern side of
the Gulf of St. Lawrence n southward to North
Carolina, in shoal and moderately deep water;
also reported (as L. piscatorius) off the Barbadoes
at 209 fathoms, on the Yucatan Bank, southern
part of the Gulf of Mexico, at 84 fathoms,80 and
off Cape Frio, Brazil, in lat. 22°56' S.,81 if these
southern specimens actually belonged to the
same species.

Occurrence in the Gulj of Maine. — This is a
familiar fish in the Gulf of Maine both along shore
and on the outer fishing banks. It has been
recorded in print from the west coast of Nova
Scotia (St. Mary Bay) and from various localities
in the Bay of Fundy, where, according to Hunts-
man, large ones are frequently taken on long lines,
or found stranded on the beach. It is well known,
if not abundant, all along the coast of Maine, and
we once caught 8 (all large) in Ipswich Bay in one
haul of a beam trawl only 8 feet wide. In Massa-
chusetts Bay goosefish are the most common on
the smooth bottom south of Boston; many enter
Duxbury Bay (p. 546) ; and they are so numerous
in Cape Cod Bay that one can hardly walk the
beach for an afternoon without finding a jawbone
bleaching on the sand, which applies equally all
along the outer shores of Cape Cod. Fishermen
speak of them as common on and about Stellwagen
Bank, also. And we have trawled them in the
deep basin of the Gulf.

Goosefish formed about 1 percent (in numbers
of individuals) of the fishes of all kinds taken by
certain otter trawlers in the South Channel and
on Georges Bank in 1913. And most of the trawl
hauls that we have seen made there subsequently

'« The smallest ripe males of the North European species seen by Fulton
were 26-27 inches long, the smallest ripe females 30 inches.

'< Pennant's (Arch. Zoo!., vol. 1, 1784, p. cxci) report of "the Lophiut pitca-
torius or common angler" in Hudson Bay seems to have been based on a
sculpin (for history of the case, see Connolly, Bull. 3, Biol. Bd. Canada,
1920, p. 7). And we think it likely that this applies also to the "Lophivx
laevigatas" reported by Weiz (Proc. Boston Soc. Nat. Hist., vol. 10, 1866,
p. 269) from Okak, northern Labrador.

M Goode and Bean, Smithsonian Contrib. Knowl., vol. 30, 1895, p. 486.

•' Regan, British Antarctic (Terra Noea) Exped. (1910), Zool., vol. I, No.
1, 1914, p. 23.

in depths of 60 to 100 fathoms have brought in
from 1 to 40 of them.

They do not show any evident preference for
any particular depth zone in the inner parts of the
Gulf between tide mark and 100 fathoms or so,
and the Albatross III found them generally dis-
tributed from 22-30 fathoms down to at least as
deep as 150-160 fathoms both on Georges Bank
and off southern New England to the westward,
in May 1950.82 Our failure to take any in the
bowl between Jeffreys Ledge and the coast sug-
gests that they may avoid the very softest mud
bottoms. And it is likely that a rather definite
concentration of them in depths of 26 to 45 fathoms
on the southwestern part of Georges Bank in June
1951, when the Eugene H caught an average of
about 5 per haul there, but only 1 per haul at
46-65 fathoms, was a matter of the food supply,
not of the depth.

Goosefish are said to be as common on Browns
Bank as they are on Georges, also along the outer
Nova Scotian coast and banks as far as Ban-
quereau, though they may not be as common in-
shore there as they are in the Gulf of Maine.
They must be generally distributed in the southern
side of the Gulf of St. Lawrence also, to judge
from the localities of record there, and they have
been reported from Anse des Dunes and from
near Mingan on the north shore.83 They have
also been trawled at a few localities on the south-
ern and eastern part of the Grand Banks.84 And
a 4-inch specimen was brought back from the
Grand Bank in 1856.86 But this seems to be its
northern limit in our side of the open Atlantic,
for they have not been reported from the east
coast of Newfoundland, or reliably from the outer
coast of Labrador (see footnote 79, p. 540).

Goosefish are common westward and southward
also, as far as North Carolina. We have seen
many stranded in winter a few miles north of Cape
Hatteras, both in Pamlico Sound and on the out-
side beach, and Smith 8f> described it as so plentiful

» Catches of 1 to 34 per haul.

« See Cox (Contrib. Canadian Biol. [1902-1905], 1907, p. 90), Cornish
(Contrib. Canadian Biol. [1906-1910], 1912, p. 81), and Connolly (Bull. 3,
Biol. Bd. Canada, 1920, p. 7) for Nova Scotian and Gulf of St. Lawrence
localities; the Annual Reports of the Newfoundland Fishery Research Com-
mission, vol. 1, No. 4, 1932, p. 110, for additional records for the Nova Scotian
Banks.

•< Rept. Newfoundland Fish. Res. Comm., vol. 1, No. 4, 1932, p. 110,
Sta. 17; vol. 2, No. 1, 1933, p. 127, Sta. 97; vol. 2, No. 2, 1935, p. 116, Sta. 204,
205, 274.

" Goode and Bean, Smithsonian Contrib. Knowl., vol. 30, 1895, p. 486.

« North Carolina Econ. Geo]. Surv., vol. 2, 1907, p. 399.

FISHES OF THE GULF OF MAINE

541

near Cape Lookout that "as many as 20 large speci-
mens are sometimes found in a sink net at one
lift," though it has been seen less often of late
years.87

Egg veils have been reported within the Gulf of
Maine from Campobello Island at the entrance to
the Bay of Fundy; from Passamaquoddy Bay (2
instances) ; 88 in Frenchman Bay, Maine ; 89 about
15 miles off Seguin Island, Maine, September 18,
1925 (with eggs nearly ready to hatch, found by
Capt. Greenleaf of the U. S. Bureau of Fisheries) ;
and at Provincetown, where we found a veil
within a few feet of the shore, on June 26, 1925.
The captures of the pelagic larvae within the Gulf
have been fewer, namely 3 taken near Brazil Rock
off southwestern Nova Scotia, and two very small
ones (5 and 6.5 mm. long) collected by us on the
Grampus in Massachusetts Bay, one on July 12,
1912, the other September 29, 1915.

The question why the egg veils should have
been encountered so seldom in our Gulf when they
are so conspicuous in the water, and why so few
larvae have been taken in our tow nets, when the
parent fish are moderately plentiful and very
generally distributed, is an intriguing one that
we cannot answer from present information.

Importance. — No regular commercial use has
been made of the goosefish in America up to the
present time. But it is an excellent food fish,
white-meated, free of bones, and of pleasant flavor,
as Dr. Connolly assures us from personal expe-
rience. In 1948,90 English and Scottish vessels
landed about 7 million pounds of the European
species, as "monk" which fetched nearly as high
a price as haddock in English markets, though it
brings only about one-half as high a price as
haddock in Scotch ports.91 And some were sold
in retail stores during the last year.

THE SARGASSUM FISHES. FAMILY ANTENNARIIDAE

The sargassum fishes resemble their relatives
the anglers (p. 532) in their peculiar armlike pec-
torals and in their large fleshy ventrals, as well as
in the fact that the forward part of their dorsal
fin is represented by a series of separate spines.
But they are very different from the anglers in
general appearance, for their bodies and heads
are flattened sidewise instead of dorso-ventrally;
their soft second dorsal fin is much longer than
their anal fin; their second and third dorsal spines
are clothed with fleshy skin so thick as to obscure
their true nature; and their mouth is much smaller
than that of the anglers.

These curious little tropical-oceanic fishes live
chiefly among masses of floating seaweed, with
which they sometimes drift far to the north of
their normal homes.

Sargassum fish, Histrio pictus (Cuvier and
Valenciennes) 1837

Mousefish

Jordan and Evermann, 1896-1900, p. 2716, as Ptero-
phryne histrio (Linnaeus) 1758.

Description. — The peculiar armlike pectoral fins;
the rather long fleshy ventral fins situated on the

throat; the soft flabby skin; and a body flattened
sidewise (about 2% times as long as it is deep)
distinguish this fish from any other that is known
from the Gulf of Maine, or that is ever likely to be
found there. The goosefish (p. 532), the only one
of its close relatives which normally inhabits the
Gulf, is of such different appearance with its
flattened body form, and enormous mouth, that
it is not necessary to enumerate the smaller
differences between the two.

The head and body of the sargassum fish appear
as one, for each gill opening has the form of a pore
on the lower margin of the pectoral near its base,
so small that it is likely to be overlooked. There
are three detached dorsal spines. The first, stand-
ing over the front margin of the eye, has the form
of a slender tentacle, its tip bearing a bulbous
swelling, known technically as an "illicium,"
which is fringed at the tip. The second spine
(close behind the first) and the third are much
larger than the first; they are enclosed in skin so
thick that they suggest conical horns in then-
general appearance, and they bear several tags or
streamers of skin. All the other fins are also
fleshy. The second (soft-rayed) dorsal fin is more
than twice as long as the anal fin; and the detached

87 Information supplied by the late S. F. Hildebrand.
» Connolly, Contrib. Canadian Biol. (1921) 1922, No. 7, p. 116; Berrill
Contrib. Canadian Biol. Fish. N. Ser., vol. 4, No. 12, 1929, p. 145.
»» Procter and others, Biol. Surv. Mt. Desert, Ft. 2 Fishes, 1928, p. 3.

" Most recent year for which the International fisheries statistics are
readily available.

•i For catches and values, see Bull. Statist., Cons. Internat. Explor. Mer.,
vol. 33, 1951, pp. 14, 15-18.

542

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Figure 287. — Sargassum fish (Histrio pictus). Drawing by Louella E. Cable.

tips of both the dorsal rays and the anal rays form
short fringes. The margin of the caudal fin is
almost straight. The skin feels smooth to the
touch; actually it is finely studded with minute
granulations, and it bears variously shaped fleshy
tags, as appears in the illustration (fig. 287) .

Color. — Creamy white, the fins as well as the
head and body mottled with pale and dark brown.
The fleshy tags are yellowish.

General range. — Tropical and subtropical, living
at the surface among floating seaweed; sometimes
drifting far northward with the Gulf Stream.

Occurrence in the Gulf of Maine. — A specimen
about 4% inches (12 cm.) long, that was picked up

in a purse seine near the surface over the west
central part of Georges Bank, by the Schooner
Old Glory on September 15, 1930,82 and a second
of 2% inches, taken off the southeast slope of
Georges Bank, by the sword fisherman Leonora C,
on June 15, 1937, are the only records of this fish
in the Gulf of Maine; the most northerly records,
in fact, for it for continental waters in this side
of the Atlantic. But it has been picked up from
time to time near Woods Hole.83 Living, as they
usually do, among floating gulf weed (Sargassum),
it is not astonishing that sargassum fish should
drift in over the offshore banks, occasionally.

THE DEEP SEA ANGLERS. FAMILY CERATIIDAE

The members of this family fall with the
anglers and sargassum fishes in the pediculate
tribe. And the first dorsal spine bears a "bait" at
its tip (known technically as an "illicium") as it
does among the anglers. But the wristlike struc-
ture of the pectoral fins is not obvious in the deep
sea anglers. And the members of this family, as
well as those of several other families closely related

> Reported by Firth, Bull. 61, Boston Soc. Nat. Hist., 1931, p. 14.

to them,94 differ from the anglers and from the
sargassum fishes in lacking ventral fins. Their
bodies are somewhat flattened sidewise (not
dorso-ventrally as in the anglers); their dorsal
and anal fins are very short (3 to 5 rays); and
their central four caudal rays are branched. Their

n For early records of It near Woods Hole, see Sumner, Osburn, and Cole,
Bull. U. S. Bur. Fish., Vol. 31, Part 2, 1913, p. 774.

« For a synopsis of the ceratioid fishes, a numerous race, see Regan and
Trewavas, Rept. 2, Danish Dana Eiped. (1928-1930), 1932, p. 48.

FISHES OF THE GULF OF MAINE

543

mouths are oblique when closed, or even vertical.
Associated with their deep-water habitat their
bodies are noticeably soft and flabby. Their eyes
are very small; some appear to be blind. The
ceratioids, too, are unique among the vertebrates in
the fact that the males of many of them (including
those of the Gulf of Maine species) are dwarfs in
size as compared with the females, and live
parasitic, attached to the females by their heads.
They are oceanic as a group, living in the mid
depths, mostly from about 200 fathoms down to
perhaps 750 fathoms. And they are blackish in
color as are so many other pelagic fishes of that
same depth zone. One species has been taken in
the Gulf of Maine as a stray.

Deep sea angler Ceratias holbolli Kr0yer 1844

Jordan and Evermann, 1896-1900, p. 2729 (young as
Mancalias uranoscopus Murray); p. 2730 (young as
Mancalias shufeldti [Gill]; Barbour and Bigelow, Proc.
New England Zool. Club, vol. 23, 1944, p. 16 (adult, as
Reganichthys giganteus, new genus and species) ; Clarke,
Discovery Rept., vol. 26, 1950, p. 1 (adult).

Description. — This deep sea angler is so bizarre
in its appearance that there is no danger of con-

fusing it with any other Gulf of Maine fish, unless
it were with some other member of its own family.
In the large female, the body is strongly flattened
sidewise; the eyes are very small and set high on
the head; and the mouth is nearly vertical when
it is closed. Perhaps their most striking external
feature is the very long and extremely slender
bristlelike spine or "tentacle," that is borne on
the top of the head. This is jointed about two-
thirds the way out along its length, and it ends
in a fleshly, pear-shaped swelling ("illicium"), the
tip of which is described as pierced by a small
pore.95 The illicium is supposed to be luminous,86
and it bears 2 to 4 short filaments.97 This head-
tentacle corresponds to the whiplike head spine of
the goosefish, but is situated farther back, about
abreast of the eyes. It is interpreted as repre-
senting a vestige of the first dorsal fin. The basal

•' So described by Clarke (Discovery Rept., vol. 26, 1950, p. 9) for an Ant
arctic specimen; tbe pore is not visible on the specimen we have examined.

•■ Dablgren (Science, vol. 68, 1928, p. 65) describes the tip of the illicium of
an unnamed species of Ceratias as with an open gland in which light is pro-
duced by bacteria.

" Four in the Gulf of Maine specimen described by Barbour and Bigelow
(Proc. New England zoo!, club, vol. 33, 1944, p. 9) as Reganichthys giganteus:
two (each bifid) in an Antarctic specimen described by Clarke.

Figure 288. — Deep-sea angler (Ceratias holbolli), adult
female (above) and parasitic male (right) that was
attached to her, off Mount Desert Rock. From Bar-
bour and Bigelow.

544

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

joint of the head tentacle is provided with re-
tractor muscles by which it can be withdrawn
rearward into a tunnel-like sheath along the head
and back, bringing the "bait" close to the mouth.

Bertelsen w has found (from dissecting a West
Greenland specimen) that when this happens, the
rear part of the hard axis of the head tentacle,
which is enclosed in the very elastic skin, emerges
from the back of the fish, about midway between
the caudal fin and the base of the pectoral fins,
so as to form the axis of a slender, tapering "dorsal
tentacle." Thus this extraordinary and unique
structure, which has been the subject of much
discussion, is actually the rear end of the head
tentacle which protrudes when the latter is drawn
rearward.

When the cephalic tentacle is moved forward by
its protractor muscles, its protruding rear end is
withdrawn into the tentacular sheath, either par-
tially, when the so-called dorsal "tentacle" appears
as a short fingerlike process, or wholly, leaving
simply an indentation or pore in the midline of
the back, as it is in the Gulf of Maine specimen
pictured in figure 288.

Close behind the so-called "dorsal tentacle" (or
behind the pore representing the latter) are a pair
of low, fleshy appendages or "caruncles," scarcely
noticeable on large specimens, but more con-
spicuous on small. These have been interpreted
as vestiges of the first dorsal fin, for each of them
encloses a spine that can be felt if not seen. Their
function is not known."

The skin is strewn with small prickles on very
small specimens, but is close-set with low conical,
broad-based thorns on larger fish. The eyes are
minute, seemingly functional on small fish, but
covered over by skin and apparently blind on
large ones. The gill slits are very small, C-shaped;
placed below the pectorals and a little behind
them. The small, slender, sharp-pointed teeth
are directed into the mouth. The dorsal and anal
fins each have 4 rays, thick, fleshy, and tapering,
as the caudal rays are also. The central caudal
rays are forked. The caudal fin has been de-
scribed as occupying as much as two-fifths of the

total length of the fish when it is intact.1 But it
has been much damaged in most of the specimens
that have been seen, and the membranes of all
the fins have been mostly torn away.

Color. — Small ones are jet black, but the dermal
prickles, being colorless, show white against the
black skin on large specimens, giving a granulated
black and white appearance.

Size. — The largest specimen seen so far 2 was
26K inches (68 cm.) long to the base of the tail fin,
and about 47 inches (119 cm.) long, counting the
tail fin.3

The parasitic males are fastened to the ventral
side of the female, by two outgrowths from the
front of the head, that are fused at the tip. They
have no teeth, no tentacle-like spine and no eyes,
and the alimentary canal is vestigial; in fact, about
the only important internal organ is a large testis.
But their fins resemble those of their mates, as do
the gill openings; their skins are prickly; and they
are similarly black. Those that have been seen (1
or 2 per female) have ranged from about 3% inches
(85 mm.) long to about 6 inches (150 mm.) long
(Gulf of Maine specimen).

General range. — Oceanic and apparently cosmo-
politan, for adults have been reported from Green-
land; Iceland (where it has been taken the most
often); off Nova Scotia; Gulf of Maine; near the
Azores; and in the sub-Antarctic. Young speci-
mens apparently referable to this species are re-
ported off southern New England; from the
Caribbean; near the Canaries; north of the Cape
Verde Islands; from the South Atlantic (lat.
52°25' S., long. 9°50' E.); and also widespread in
Indo-Pacific waters.*

Occurrence in the Gulf of Maine. — A female,
about 32 inches long to the base of the caudal fin,
and about 40 inches counting what remained of
the latter (fig. 288), with one male attached, and
showing the scar of attachment of another, was
taken 12 miles south of Mount Desert rock, at 125

11 Vid. Meddel. Dansk Naturh. Foren., vol. 107, 1943, pp. 190-193; see
especially his fig. 4, p. 192.

'•See Regan and Trewavas (Rept. 2, Danish Dana Exped. (1928-1930),
1932, pp. 23-24) for an account of the lateral line papillae in different families
of deep sea anglers.

1 This was the case in the specimen about 47 inches (119 cm.) long described
by Krbyer (Naturbist, Tiddsskr., Ser. 2, vol. 1, 1844, pp. 640-642); also in
one pictured by Goode and Bean (Smithsonian Contrib. Knowl., vol. 31,
1895, pi. 117, fig. 399, after Qaimard).

1 Kroyer's original specimen from Greenland.

• See Clarke {Discovers/ Rept., Vol. 26, 1950, p. 14, table 1), for measurements
of several specimens.

• For complete list of localities for adults and young, see Clarke, Discovert
Rept., vol. 26, 1950, pp. 23, 30.

FISHES OF THE GULF OF MAINE

545

fathoms, in October 1943, by the schooner Dorothy
and Ethel II, Capt. Harold Paulsen.6 A second
female, about 18% inches long to the base of the
caudal fin, and about 24% inches counting what
was left of the caudal fin, trawled on the southeast
part of Georges Bank, between 150 and 200
fathoms, in February 1927, appears to belong to

this same species.6 A third probable Gulf of
Maine record is of a fish, about 3 feet long, and
weighing about 20 pounds, that was taken by the
trawler Ebb, in 140 fathoms, on Georges Bank, in
June 1936. Photographs of it appeared in the
Boston Globe and in the Boston Post for June 29
of that year.

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546

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1872. [Note on Coitus groenlandicus Fabr.] Proc.

Acad. Natural Sciences Phila., 1872, pp.
213-214.

1873. On a new American species of Pleuronectoid

(Glyptocephalus acadianus). Proc. Acad.

Natural Sciences Phila., 1873, pp. 360-362.

1873a. Catalogue of the fishes of the east coast of

North America. Report U. S. Comm.

Fish and Fisheries for 1871-72, pp. 779-

822.

1905. The sculpin and its habits. Smithsonian

Misc. Collections, vol. 47, pp. 348-359, figs.

45-55.

Gill, Theodore — Continued

1905a. The life history of the angler. Smithsonian
Misc. Collections, vol. 47, pp. 500-516,
figs. 94-103, pis. 73-75.
1905b. State ichthyology of Massachusetts. Report
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1911. Notes on the structure and habits of the wolf-
fishes. Proc. U. S. Nat. Museum, vol. 39,
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Girard, C. F.

1850. On the genus Coitus Auct. Proc. Boston Soc.
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Godsil, N. C, and E. K. Holmberg.

1950. A comparison of the bluefin tunas, genus
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1875. Albino fishes. Amer. Naturalist, vol. 9, p.

517.

1879. The occurrence of Hippocampus antiquorum,

or an allied form, on Saint George's Banks.

Proc. U. S. Nat. Museum, vol. 1, pp. 45-46.

1879a. The natural and economical history of the

American menhaden. Report U. S. Comm.

Fish and Fisheries for 1877, pp. 1-529, pis.

1-31.

1882. The carangoid fishes of the United States . . .

pompanos, crevalles, amber-fish, and others.
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1883. Materials for a history of the swordfish.

Report U. S. Comm. Fish and Fisheries for
1880, pp. 289-394, pis. 1-24.

1884. Natural history of useful aquatic animals.

Pt. 3, The food fishes of the U. S. In Fish.
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1885. A brief biography of the halibut. Amer.

Naturalist, vol. 19, pp. 953-969.

1887. The swordfish fishery. In Fisheries and

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1888. American fishes . . . xv, 496 pp., illus.
Goode, George Brown, and Tarleton H. Bean.

1877. Descriptions of two new species of fishes
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recently discovered by the U. S. Fish Com-
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several unusual forms. Amer. Jour. Science
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1879. A list of the fishes of Essex County, including
those of Massachusetts Bay, according to
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vol. 11, pp. 1-38.

1879a. The Craig flounder of Europe, Glyptocephalus
cynoglossus, on the coast of North America.
Proc. U. S. Nat. Museum, vol. 1, pp. 19-23.

1879b. The oceanic bonito on the coast of the United
States. Proc. U. S. Nat. Museum, vol. 1,
pp. 24-26.

550

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Goode, Geohge Brown, ond Tarleton H. Bean — Con.

1879c. Descriptions of two gadoid fishes, Phycis
chesteri and Haloporphyrus viola, from the
deep-sea fauna of the northwestern At-
lantic. Proc. U. S. Nat. Museum, vol. 1,
pp. 256-260.

1879d. The identity of Rhinonemus caudacula
(Storer) Gill with Gadus cimbrius Linn.
Proc. U. S. Nat. Museum, vol. 1, pp. 348-
349.

1879e. Note on Platessa ferruginea, D. H. Storer, and
Plalessa rostrata, H. R. Storer. Proc. U. S.
Nat. Museum, vol. 1, pp. 361-362.

1879f. On the identity of Brosmius americanus, Gill,
with Brosmius brosme (Muller) White.
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363.

1879g. Discoveries of the United States Fish Com-
mission: Notices of fifty species of east-
coast fishes . . . Amer. Jour. Science and
Arts, Third Ser., vol. 17, pp. 39-48.

1883. Reports on the results of dredging under the
supervision of Alexander Agassiz ... by
the U. S. Coast Survey steamer Blake . . .
Report on the fishes. Bull. Museum
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1886. Description of Leptophidium cervinum and

L. marmoratum, new fishes from deep water
off the Atlantic and Gulf coasts. Proc. U. S.
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1895. Oceanic ichthyology . . . Smithsonian Contrib.
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Goode, George Brown, and A. Howard Clark.

1887. The menhaden fishery. In Fisheries and

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Goode, George Brown, and Joseph W. Collins.

1882. The winter haddock fishery of New England.
Bull. U. S. Fish Comm., vol. 1, pp. 226-235.
1887. The fresh-halibut fishery. In Fisheries and
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1887a. The Banks hand-line cod fishery; the Lab-
rador and Gulf of St. Lawrence cod
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vol. 1, pp. 123-198.
1887b. Haddock fishery of New England. In
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1887c. The hake fishery. In Fisheries and Fishery
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243.
1887d. The mackerel fishery of the United States.
In Fisheries and Fishery Industries of
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Goode, George Brown, Joseph W. Collins, R. E. Earll,
and A. Howard Clark.

1884. Materials for a history of the mackerel fishery.
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Graham, Michael.

1936. Investigations of the herring of Passama-
quoddy and adjacent regions. Jour. Biol.
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Grant, Isaac H.

1883. Movements of menhaden — catch of herring.
Bull. U. S. Fish Comm., vol. 3, p. 470.
Gf)NTHER, Albert.

1862. Catalogue of the fishes in the British Museum.
Vol. 4, xxi, 534 pp.
Halkett, Andrew.

1913. Check list of the fishes of the Dominion of
Canada and Newfoundland. 138 pp., 14
pis.
Hall, Ansley.

1898. The herring industry of the Passamaquoddy
region, Maine. Rept. U. S. Comm. Fish and
Fisheries for 1896, pp. 443-487.
Haskell, E. H.

1883. Second annual appearance of young cod
hatched by the United States Fish Commis-
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1879-80. Bull. U. S. Fish Comm., vol. 2,
p. 112.
Herrington, W. C.

1936. Decline in haddock abundance on Georges

Bank and a practical remedy. Fishery

Circular No. 23, U. S. Bur. Fisheries, 22 pp.

1944. Factors controlling population size. Trans.

9th N. American Wildlife Conf., pp. 250-263.

Heldebrand, Samuel F., and William C. Schroeder.

1928. Fishes of Chesapeake Bay. Bull. U. S. Bur.

Fisheries, vol. 43, Part 1, 388 pp.

Holmes, Ezekiel.

1862. Report on the fishes of Maine . . . Second
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Homans, R. E. S., and A. W. H. Needler.

1946. Food of the haddock. Proc. Nova Scotian
Inst. Sci., vol. 21, pts. 1 and 2, pp. 15-49.

Honetman, D.

1886. Nova Scotian ichthyology. Proc. Trans.
Nova Scotian Inst. Science, vol. 6, Part 4,
pp. 328-330.

Hubbs, Carl L.

1923. Notes on a small collection of fishes from Mon-
hegan Island, Maine. Copeia, No. 123, pp.
101-103.

Huntsman, A. G.

1918. Histories of new food fishes. 1. The Cana-
dian plaice. Bull. Biol. Board Canada No.
1, 22 pp.

FISHES OF THE GULF OF MAINE

551

Huntsman, A. G. — Continued

1918a. The effect of the tide on the distribution of
the fishes of the Canadian Atlantic coast
Trans. Royal Soc. Canada, Ser. 3, vol. 12,
Sect. 4, pp. 61-67.
1918b. The growth of the scales in fishes. Trans.
Royal Canadian Institute, vol. 12, pp.
61-101.
1918c. Our eastern flat-fishes. Canadian Fisher-
man, June 1918, vol. 5, No. 6, pp. 788-790.
1919. Growth of the young herring (so-called sar-
dines) of the Bay of Fundy. Canadian
Fisheries Exped. 1914-15, Dept. of Naval
Service [Canada], pp. 165-171.
1922. Is winter mackerel fishery possible? Cana-
dian Fisherman, May, vol. 9, No. 5, pp.
88-89.
1922a. The fishes of the Bay of Fundy. Contrib.
Canadian Biol, for 1921, No. 3, pp. 49-72.
1931. The Maritime salmon of Canada. Bull. Biol.
Board Canada, No. 21, 99 pp.

1933. St. John November salmon, the earliest run

known. Progress Rept., No. 6, Atlantic
Biol. Stations, Biol. Board Canada, pp. 7-10.

1934. Herring and water movements. James John-

stone Memorial Volume, pp. 81-96.

1936. Return of salmon from the sea. Bull. Biol.

Board Canada, No. 51, 20 pp.

1937. "Migration" and "homing" of salmon. Sci-

ence, New Ser., vol. 85, pp. 313-314.

1938. "Spawning urge," "homing instinct", and

"waiting" in salmon return. Nature, vol.
141, pp. 421-422.

1948. Atlantic salmon investigation. Ann. Report

Fish. Research Board Canada for 1947, pp.
37-38.

1949. Atlantic salmon investigations. Ann. Report

Fish. Research Board Canada for 1948, pp.
40-41.

1950. Report for 1949 of fish (salmon) management.

Ann. Report Fish. Research Board Canada

for 1949, App. 2, pp. 41-43.
1952. The production of life in the Bay of Fundy.

Trans. Roy. Soc. Canada, vol. 46, ser. 3,

sect. 5, pp. 15-38.
Huntsman, A. G., and A. W. H. Needler.

1927. Fluctuations in the haddock fishery of the

Bay of Fundy. Contrib. Canadian Biol.

Fish., New Ser., vol. 3, No. 18, pp. 423-436.
Huntsman, A. G., and others.

1939. Migration and conservation of salmon. Pub.

No. 8, Amer. Assoc. Adv. Science, 106 pp.
Jackson, C. T.

1852. [Capture of a basking shark off St. John, New
Brunswick.] Proc. Boston Soc. Natural
Hist., vol. 4, pp. 202-205,
1852a. [Capture of a thresher shark in Boston Har-
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vol. 4, pp. 205-206.

Jackson, J. B. S.

1847. Bluefish at Nahant, Mass. Proc. Boston Soc.
Natural Hist., vol. 2, p. 248.
Jeffebs, A. M.

1932. The Fundy shad industry. Ann. Report Biol.

Board Canada for 1931, p. 26.
1932a. The Fundy cod fishery. Ann. Rept. Biol.

Board Canada for 1931, pp. 26-27.
1932b. The Fundy smelt fishery. Ann. Rept. Biol.
Board Canada for 1931, p. 27.
Johansen, Frits.

1925. Natural history of the cunner (Tautogolabrus
adspersus Walbaum). Contrib. Canadian
Biol., New Ser., vol. 2, No. 17, pp. 423-468.
Jones, J. Matthew.

1879. List of the fishes of Nova Scotia. Proc. Trans.
Nova Scotian Inst. Nat. Sci., vol. 5, Pt. 1,
pp. 87-97.
Jordan, David Starr.

1891. A review of the labroid fishes of America and

Europe. Report U. S. Comm. Fish and
Fisheries, 1887, pp. 599-699, pis. 1-11.
Jordan, David Starr, and Bradley Moore Davis.

1892. A preliminary review of the apodal fishes or

eels inhabiting the waters of America and
Europe. Report U. S. Comm. Fish and
Fisheries, 1888, pp. 581-677, pis. 73-80.

Jordan, David Starr, and Charles L. Edwards.

1887. A review of the American species of Tetrao-
dontidae. Proc. U. S. Nat. Museum, vol.
9, pp. 230-247.

Jordan, David Starr, and Carl H. Eigenmann.

1889. A review of the Sciaenidae of America and

Europe. Report U. S. Comm. Fish and
Fisheries, 1886, pp. 343-451, pis. 1-4.

1890. A review of the genera and species of Serrani-

dae found in the waters of America and
Europe. Bull. U. S. Fish Comm., vol. 8,
pp. 329-441, pis. 60-69.
Jordan, David Starr, and Barton Warren Evermann.
1896. A checklist of the fishes and fishlike verte-
brates of North and Middle America.
Report U. S. Comm. Fish and Fisheries for
1895, pp. 207-584.
1896-1900. The fishes of North and Middle America.
Bull. U. S. Nat. Museum, No. 47,
Parts 1-4, 3313 pp., 392 pis.
Jordan, D. S., B. W. Evermann, and H. W. Clark.

1930. Checklist of the fishes and fishlike vertebrates
of North and Middle America . . . Report
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2, pp. 1-670.
Jordan, David Starr, and Charles H. Gilbert.

1882. Synopsis of the fishes of North America.
Bull. U. S. Nat. Museum, No. 16, lvii,
1018 pp.
1884. A review of the American Caranginae. Proc.
U. S. Nat. Museum, vol. 6, pp. 188-207.

552

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Jordan, David Starb, and David Kop Goss.

1889. A review of the flounders and soles (Pleuro-
nectidae) of America and Europe. Rept.
U. S. Comm., Fish and Fisheries, 1886, pp.
225-342, Pis. 1-9.
Jordan, David Starr, and Elizabeth G. Hughes.

1887. A review of the species of the genus Prionotus.
Proc. U. S. Nat. Museum, vol. 9, pp. 327-
338.
Josseltn, John.

1672. New England rarities discovered in birds,
beasts, fishes, serpents, and plants of that
country. Printed for G. Widdowes at the
Green Dragon in St. Paul's Churchyard,
1672. London. [Fishes, p. 23.] Second
edition, 1675, London. Reprint of first
edition, edited by Edward Tuckerman, in
Trans. Amer. Antiquarian Soc, vol. 4, 1860,
pp. 105-238. Another reprint by Edward
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1674. An account of two voyages to New England.
Printed for Giles Widdowes at the Green
Dragon in St. Paul's Churchyard, 1674,
London. [Fishes, pp. 103-115.] Second
edition, 1675, London. Reprint of second
edition in Massachusetts Historical Soc.
Collections, Ser. 3, vol. 3, 1833, pp. 211-396
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Kendall, William Converse.

1896. Description of a new stickleback, Gasterosleus
gladiunculus, from the coast of Maine.
Proc. U. S. Nat. Museum, vol. 18, pp.
623-624.

1898. Notes on the food of four species of the cod
family. Report U. S. Comm. Fish and
Fisheries, 1896, pp. 177-186.

1902. Notes on the silversides of the genus Menidia
of the east coast of the United States, with
descriptions of two new subspecies. Rept.
U. S. Comm. Fish and Fisheries for 1901,
pp. 241-267.

1908. Fauna of New England. List of the Pisces.
Occasional Papers, Boston Soc. Nat. Hist.,
vol. 7, No. 8, 152 pp.

1910. Effects of the menhaden and mackerel fish-
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Bur. Fisheries, vol. 28, Pt. 1, pp. 279-293.

1912. Notes on a new species of flatfish from off the
coast of New England. Bull. U. S. Bur.
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1914. An annotated catalogue of the fishes of Maine.
Proc. Portland Soc. Nat. Hist., vol. 3, Pt.
1, pp. 1-198.

1917. The capelin (Mallotus villosus), with notes on
its occurrence on the coast of Maine.
Copeia, No. 42, pp. 28-30.

Kendall, William Converse — Continued

1919. Second authentic record of capelin {Mallotus
villosus) on the Maine coast. Copeia, No.
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1927. The smelts. Bull. U. S. Bur. Fisheries, vol.

42, pp. 217-375.
1931. Remarks on additions to the marine fauna of
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1935. The fishes of New England. The salmon
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pis.
Kendall, W. C, and Donald R. Crawford.

1922. Notice of a spiral valve in the teleostean fish
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Kingslet, J. S.

1922. The food habits of swordfish. Science, New
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Kingsley, J. S., and H. W. Conn.

1883. Some observations on the embryology of the
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Kneeland, S., Jr.

1847. Dissection of Scymnus brevipinna (LeSueur).
Boston Jour. Nat. Hist., vol. 5, No. 4, pp.
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Knight, Thomas, F.

1866. Descriptive catalog of the fishes of Nova

Scotia. 55 pp. Halifax.

1867. Shore and deep-sea fisheries of Nova Scotia.

vi, 113 pp. Halifax.
Kuntz, Albert, and Lewis Radcliffe.

1918. Notes on the embryology and larval develop-

ment of twelve teleostean fishes. Bull.
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Laighton, Cedric.

1882. The capture of shad at Isles of Shoals, New
Hampshire. Bull. U. S. Fish Comm., vol.
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Lanman, Charles.

1874. The Salmonidae of eastern Maine, New Bruns-
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1874a. The shad and gaspereau or alewife of New
Brunswick and Nova Scotia. Rept. U. S.
Comm., Fish and Fisheries for 1872-73,
pp. 461-462.
Lea, Einar.

1919. Age and growth of herring in Canadian waters.

Canadian Fisheries Expedition of 1914-
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Lee, L. A.

1880. Occurrence of the web-fingered sea-robin on
the coast of Maine. Amer. Naturalist, vol.
14, p. 896.

FISHES OF THE GULF OF MAINE

553

Lee, L. A. — Continued

1885. The fishes of Casco Bay. Portland Adver-
tiser, March 3, 1885, and Brunswick Tele-
graph, Friday, March 13, 1885.

Leim, A. H.

1924. The life history of the shad (Alosa sapidissima
Wilson) with special reference to the factors
limiting its abundance. Contrib. Canadian
Biol., New Ser., vol. 2, No. 11, pp. 163-284,
45 figs.

1930. Unusual fishes and other forms in Nova

Scotian waters. Proc. Trans. Nova Scotian
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1937. Progress Rept. 21, Atlantic Biol. Stations,
Biol. Board Canada, p. 5.
LeSueur, C. A.

1817. A short description of five (supposed) new

species of the genus Muraena . . . Jour.
Acad. Nat. Sciences, Philadelphia, vol. 1,
Pt. 1, pp. 81-83.
1817a. Descriptions of four new species, and two

varieties, of the genus Hydrargira. Jour.

Acad. Nat. Sciences, Philadelphia, vol. 1,

Pt. 1, pp. 126-134.

1818. Descriptions of several new species of North

American fishes. Jour. Acad. Nat. Sci-
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1821. Observations on several genera and species of
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Lyman, Henry.

1950. Bluefishing. 98 pp. Boston.
Lyman, Theodore.

1859. [Account of the habits of some animals recently
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Proc. Boston Soc. Nat. Hist., vol. 7, pp.
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Lyman, Theodore, and Alf. A. Reed.

1866. [Concerning the obstructions to the passage of
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Senate Massachusetts Document No. 8,
Dec. 1865 (1866), pp. 3-41.
MacCoy, Clinton V.

1929. The mackerel in New England. Bull. No. 53,
Boston Soc. Nat. Hist., pp. 3-7.

1931. Museum Notes, Fishes. Bull. No. 58, Boston

Soc. Nat. Hist., pp. 16-18.
1931a. Museum Notes, Fishes. Bull. No. 61,
Boston Soc. Nat. Hist., p. 21.

1933. Museum Notes, Fishes. Bull. No. 67, Boston

Soc. Nat. Hist., p. 9.
1933a. Museum Notes, Fishes. Bull. No. 69,
Boston Soc. Nat. Hist., pp. 8-9.

1934. Museum Notes, Fishes. Bull. No. 70, Boston

Soc. Nat. Hist., pp. 6-7.

1935. Fishes. Bull. No. 74, Boston Soc. Nat. Hist.,

p. 16.

MacDonald, Marshall.

1884. The shad and the alewives. In Fish. Ind.
U. S., Sec. 1, Text, pp. 579-609.
MacFarland, W. E.

1931. A study of the Bay of Fundy herring. Ann.

Rept. Biol. Board Canada for 1930, pp.

23-24.
McGonigle, R. H., and M. W. Smith.

1936. Three rare fishes from Passamaquoddy Bay.

Proc. Nova Scotian Inst. Sciences, vol. 19,

p. 160.
McIntire, Edwin C.

1906. Numbers of dogfish. Rept. Massachusetts

Comm. Fish. Game for 1905, pp. 108-109.
McKenzie, R. A.

1932. The Fundy haddock fishery. Ann. Rept. Biol.

Board Canada for 1931, p. 34.
1932a. The Fundy alewife fishery. Ann. Rept. Biol.

Board Canada for 1931, p. 34.
1934. Cod movements on the Canadian Atlantic

coast. Contrib. Canadian Biol. Fish., New

Ser., vol. 8, No. 31 (Series A, general, No.

46), pp. 433-459.

1939. Some marine fish and salp records. Proc.

Nova Scotian Inst. Sciences, vol. 20, pp.
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1940. Some marine records from Nova Scotian fish-

ing waters. Proc. Nova Scotian Inst.

Sciences, vol. 20, pp. 42-46.
McKenzie, R. A., and R. E. S. Homans.

1938. Rare and interesting fishes and salps in the

Bay of Fundy and off Nova Scotia. Proc.

Nova Scotian Inst. Sciences, vol. 19, Pt. 3,

pp. 277-281.
Maddocks, Luther.

1878. The menhaden fishery of Maine . . . Rept.

Secretary Assoc. Menhaden Oil and Guano

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Portland.
Marston, Philip M.

1938. Notes on fish and early fishing in the Merrimac

River System. Biol. Surv. Merrimac Water

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Martin, S. J.

1882. First appearance of fish at Gloucester, 1881.

Bull. U. S. Fish Comm., vol. 1, p. 66.

1882a. Notes on New England fisheries. Bull. U. S.
Fish Comm., vol. 1, p. 133.

1882b. Notes on New England food-fishes. Bull.
U. S. Fish Comm., vol. 1, p. 202.

1882c. Pollock-fishing in Boston Bay. Bull. U. S.
Fish Comm., vol. 1, p. 342.

1882d. Notes on the fisheries of Gloucester, Massa-
chusetts. Bull. U. S. Fish Comm., vol. 1,
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1883. Movements and catch of mackerel. Bull.

U. S. Fish Comm., vol. 2, pp. 89-90.

554

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Martin, S. J. — Continued

1883a. Notes on the fisheries of Gloucester, Massa-
chusetts. Bull. U. S. Fish Comm., vol. 2,
pp. 91-93.

1883b. Notes on the fisheries of Gloucester, Massa-
chusetts. Bull. U. S. Fish Comm., vol. 3,
pp. 162-178; 297-300.

1884. Notes on the fisheries of Gloucester, Massa-

chusetts. Bull. U. S. Fish Comm., vol. 4,
pp. 89-96; 249-255; 444-448.

1885. Notes on the fisheries of Gloucester, Massa-

chusetts. Bull. U. S. Fish Comm., vol. 5,
pp. 57-62; 203-208.
Martin, W. R., and F. D. McCracken.

1950. Movements of halibut on the Canadian Atlan-
tic coast. Progress Report Atlantic Coast
Stations, Fish. Res. Board Canada, No. 50,
pp. 3-6.
Massachusetts Commissioners of Inland Fisheries.
1877-86. [Annual statistics of the pound and shore
seine fisheries of Massachusetts.] Ann.
Reports, for 1877 to 1885.
Massachusetts Commissioners on Inland Fisheries
and Game.

1887-1902. [Annual statistics of the pound and shore
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Massachusetts Commissioners on Fisheries and
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1903-20. [Annual statistics of the pound and shore
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1906. Report on the damage done by dogfish to the

fisheries of Massachusetts. Ann. Report
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1907. Dogfish. Ann. Report for 1906, pp. 19-20.
Massachusetts Division of Fisheries and Game.

1921-22. [Annual statistics of the pound and shore
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Mavor, James W.

1918. On the age and growth of the pollock in the
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1921. The utilization of dogfish and selachian fishes
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Merriman, Daniel.

1941. Studies on the striped bass (Roccus saxatilis)
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Mitchill, S. L.

1815. The fishes of New York . . . Trans. Literary
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Moore, Henry Frank.

1898. Observations on the herring and herring

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Moore, J. Percy.

1899. Report on mackerel investigations in 1897.

Report U. S. Comm. Fish and Fisheries for
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Morrow, James E.

1951. The biology of the longhorn sculpin . . .
Bull. Bingham Oceanogr. Collection, vol.
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Morton, Thomas.

1637. New English Canaan; or New Canaan, con-
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Printed by C. Green, 1632, 125 pp. 1st
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Moulton, James M.

1951. The unusual behavior of a sand launce.
Copeia, p. 90.

Needler, Alfreda P.

1931. The haddock. Bull. No. 25, Biol. Board

Canada, 28 pp.
Needler, A. W. H.

1929. Studies on the life history of the haddock

(Melanogrammus aeglefinus). Contrib. Ca-
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pp. 265-285.

1930. The migrations and the interrelationships of

haddock populations in North American

waters. Contrib. Canadian Biol. Fisher.,

New Ser., vol. 6, No. 10, pp. 241-314.
Nichols, J. T., and C. M. Breder, Jr.

1927. The marine fishes of New York and southern

New England. Zoologica, New York Zool.

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Nichols, J. T. and F. E. Firth.

1939. Rare fishes off the Atlantic coast, including a

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Washington, vol. 52, pp. 85-88.
Olsen, Y. H., and Daniel Merriman.

1946. The biology and economic importance of the

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Parr, Albert Eide.

1932. On a deep sea devilfish from New England . . .

Bull. No. 63, Boston Soc. Nat. Hist., pp.
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1933. Two new records of deep-sea fishes from New

England with description of a new genus
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FISHES OF THE GULF OF MAINE

555

Peck, William D.

1804. Description of four remarkable fishes, taken

near the Piscataqua in New Hampshire.

Mem. Amer. Acad. Arts and Sciences, vol.

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Pbrley, M. H.

1850. Statements in relation to the fisheries of the

Bay of Fundy. Proc. Boston Soc. Nat.
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1851. Report upon the fisheries of the Bay of Fundy.

v, 176 pp. [Pp. 118-159 reprinted in 1852
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1852. Descriptive catalogue (in part) of the fishes of

New Brunswick and Nova Scotia. Second
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Perlmutter, Alfred.

1947. The blackback flounder and its fishery in New
England and New York. Bull. Bingham
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Perlmutter, Alfred, and G. M. Clark.

1949. Age and growth of immature rosefish (Sebastes
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Piers, Harry.

1934. Accidental occurrence of the man-eater or
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Prince, Edward E.

1907. The eggs and early life-history of the herring,
gaspereau, shad, and other clupeoids.
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1913. The pearlsides. A luminous fish new to
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1917. On serially striped haddock in New Bruns-
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Prince, E. E., and A. H. MacKay.

1901. The paired fins of the mackerel shark. Con-
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Procter, William, and others.

1928. A contribution to the life history of the angler
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1933. Marine fauna. Biol. Survey Mt. Desert
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Putnam, F. W.

1856. [Fishes of Essex County, Mass.] Proc. Essex

Institute, vol. 1, p. 144.
1856a. [Fishes of Essex County, Mass., continued.]

Proc. Essex Institute, vol. 1, p. 148.
1856b. [Fishes of Essex County, Mass., third com-
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1864. [Note on the cod of Massachusetts waters.]
Proc. Boston Soc. Natural Hist., vol. 9,
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1866. [Occurrence of Leplocephalus gracilis at Na-
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1866a. [Description of Gaslerosleus wheatlandi, from
Nahant, Mass.] Proc. Essex Institute,
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1870. [Addition of two species of fishes to the fauna
of Essex County, Mass.] Bull. Essex
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1870a. [On a species of Hemiramphus from Danvers,
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1870b. On the young of Orthagoriscus mola. Amer.
Naturalist, vol. 4, pp. 629-633. Also in
Proceedings, Amer. Assoc. Advancement of
Science, 19th meeting, 1871, pp. 255-260.

1873. List of fishes sent by the Museum to different

institutions . . . Bull. Museum Compar.
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Manan, p. 10].

1874. [Notes on several rare fishes from Essex

County, Mass.] Bull. Essex Institute, vol.
6, No. 1, pp. 11-13.
1874a. [Tooth of a maneater that attacked a dory
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tute, vol. 6, No. 4, p. 72.
1874b. [Chauliodus sloani on George's Bank.] Bull.

Essex Institute, vol. 6, No. 7, p. 111.
1874c. [Liparis linealus and L. montaguii in Salem
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1874d. Notes on the genus Myxine. Proc. Boston

Soc. Natural Hist., vol. 16, pp. 127-135.
1874e. Notes on Liparis, Cyclopterus and their allies.
Proc. Amer. Assoc. Advancement Science,
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Radcliffe, Lewis.

1916. An extension of the recorded range of three
species of fishes in New England waters.
Copeia, No. 26, pp. 2-3.
1922. Fisheries of the New England States in 1919.
App. 5, Report of the U. S. Commissioner
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1926. "Opah" and "Skilligalee" landed at Boston
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556

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Ranet, Edward C, and others.

1952. The striped bass. Bull. Bingham Oceanogi
Coll., vol. 14, art. 1, 177 pp.

Rathbun, R., and W. Wakeham.

1897. Preservation of the fisheries in the waters con-
tiguous to the United States and Canada.
Document No. 315, U. S. House of Repre-
sentatives, 54th Congress, 2d session, pp.
1-178.

Reid, Margaret E.

1929. The distribution and development of the

cunner (Tautogolabrus adspersus) along the
eastern coast of Canada. Contrib. Cana-
dian Biol. Fish., New Ser., vol. 4, No. 27,
pp. 431-441.
Rich, Walter H.

1930. Fishing grounds of the Gulf of Maine. Report

U. S. Commissioner of Fisheries for 1929,
App. 3, pp. 51-117, 5 maps.
1947. The swordfish and the swordsfishery of New
England. Proc. Portland Soc. Natural
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Scattergood, Leslie W.

1949. The production and the fishing methods of
the Maine herring industry, with notes on
the 1947 season. Spec. Sci. Report No. 67,
U. S. Fish and Wildlife Service, 25 pp., 7
graphs and charts.
1949a. Notes on the Maine shark fishery. Copeia,

pp. 69-71.
1951. The occurrence of egg capsules in the winter
skate (Raja diaphanes) in Maine waters.
Copeia, p. 169.
Scattergood, Leslie W., P. S. Trefethen, and G. W.
Coffin.

1951. Notes on the size of menhaden taken in Maine

during 1949. Copeia, pp. 93-94.
1951a. Notes on Gulf of Maine fishes in 1949.
Copeia, pp. 297-298.
Schroeder, William C.

1930. Migrations and other phases in the life history

of the cod off southern New England. Bull.
U. S. Bur. Fish., vol. 46, pp. 1-136.
1930a. A record of Polyprion americanus (Bloch and
Schneider) from the northwestern Atlantic.
Copeia, pp. 46-48.

1931. Notes on certain fishes collected off the New

England coast from 1924 to 1930. Bull.
Boston Soc. Natural Hist., No. 58, pp. 3-8.

1933. Unique records of the brier skate and rock eel
from New England. Bull. No 66, Boston
Soc. Natural Hist., pp. 5-6.

1937. Records of Pseudopriacanthus altus Gill and
Fundulus majalis (Walbaum) from the
Gulf of Maine. Copeia, p. 238.

1939. The Provincetown "sea serpent." New Eng-
land Naturalist, No. 2, pp. 1-2.

1942 Results of haddock tagging in the Gulf of
Maine from 1923 to 1932. Jour. Marine
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Schroeder, William C. — Continued

1947. Notes on the diet of the goosefish Lophius
americanus. Copeia, p. 201.
Schuck, Howard A.

1947. Haddock (Melanogrammus aeglefinus). Fish-

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Service, 9 pp.

1948. Current haddock situation on Georges Bank.

Commercial Fisheries Review, vol. 10, No.
10, 6 pp.

1949. Relationship of catch to changes in population

size of New England haddock. Biometrics,
Amer. Statistical Assoc, vol. 5, No. 3, pp.
213-231.
1951. Northern record for the little tuna Eulhynnus

alletteratus. Copeia, p. 98.
1951a. New Gulf of Maine record for occurrence of
dolphin Coryphaena hippurus, and data on
small specimens. Copeia, p. 171.
Scudder, Newton P.

1887. The salt halibut fishery ... In Fisheries
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vol. 1, pp. 90-119.
Sella, Massimo.

1931. The tuna (Thunnus thynnus L.) of the western
Atlantic. An appeal to fishermen for the
collection of hooks found in tuna fish.
Internat. Revue gesamt. Hydrobiol.
Hydrog., vol. 25, pp. 46-67.
Sette, O. E.

1931. Outlook for the mackerel fishery in 1931.

Fishery Circular No. 4, U. S. Bur. Fish.,
20 pp.

1932. Outlook for the mackerel fishery in 1932.

Fishery Circular No. 10, U. S. Bur. Fish.,
25 pp.

1933. Outlook for the mackerel fishery in 1933.

Fishery Circular No. 14, U. S. Bur. Fish.,
23 pp.

1934. Outlook for the mackerel fishery, 1934.

Fishery Circular No. 17, U. S. Bur. Fish.,
6 pp.
1943. Biology of the Atlantic mackerel (Scomber
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1950. Biology of the Atlantic mackerel (Scomber

scombrus) of North America. Part 2.
Fishery Bull., U. S. Fish and Wildlife Service,
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Sette, O. E., and A. W. H. Needler.

1934. Statistics of the mackerel fishery off the east
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Sharp, Benjamin.

1901. The food of the cod. Proc. Acad. Natural
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Sharp, Benjamin, and Henry W. Fowler.

1904. The fishes of Nantucket. Proc. Acad. Natural
Sciences Philadelphia, vol. 56, pp. 504-512.

FISHES OF THE GULF OF MAINE

557

Sherwood, George H., and Vinal N. Edwards.

1902. Notes on the migration, spawning, abundance,
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Smith, Everett.

1883. A mammoth cod. Bull. U. S. Fish Comm.,

vol. 3, p. 443.
1889. Results of planting shad in the Kennebec
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16.
Smith, Hugh M.

1893. [Eastward movement of menhaden.] Report
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91, pp. 202-203.

1895. Notes on the capture of Atlantic salmon at sea

and in the coast waters of the eastern
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pp. 95-99.

1896. Notes on an investigation of the menhaden

fishery in 1894 . . . Bull. U. S. Fish
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1898. The fishes found in the vicinity of Woods

Hole. Bull. U. S. Fish Comm., vol. 17, pp.
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1898a. The salmon fishery of Penobscot Bay and
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1899. Notes on the extent and condition of the

alewife fisheries of the United States in 1896.
Rept. U. S. Comm., Fish and Fisheries for
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1902. Notes on the tagging of four thousand adult
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1938. Alectis crinitus, a fish new to Massachusetts
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Smith, Hugh M., and R. A. Goffin.

1937. A fish new to Massachusetts Bay. Copeia,
p. 236.
Smith, Hugh M., and William C. Kendall.

1898. Notes on the extension of the recorded range
of certain fishes of the United States coasts.
Report U. S. Comm. Fish and Fisheries for
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Smith, Jerome V. C.

1833. A catalogue of the marine fishes taken on the
Atlantic coast of Massachusetts. In Hitch-
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1833a. Natural history of the fishes of Massa-
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1835. A catalogue of the marine and fresh-water
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Smith, Captaine John.

1616. The generall historie of Virginia, New England,
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Sparks, M. Irving.

1929. The spawning and development of mackerel
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Canadian Biol. Fish., New Ser., vol. 4,
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Spinney, Benjamin H.

1883. Return to Gloucester of young codfish hatched
by United States Fish Commission in 1879.
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Stevenson, Charles H.

1899. The shad fisheries of the Atlantic coast of the
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Storer, David Humphreys.

1836. An examination of the "Catalogue of the

marine and fresh water fishes of Massachu-
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1837. Description of a new species of the genus

Hydrargyra; with some additions to the
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1839. Remarks on the "Natural History of the

Fishes of Massachusetts, embracing a Prac-
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1839a. A report on the fishes of Massachusetts.
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1839b. Reports on the fishes, reptiles, and birds of
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1840. Supplement to the ichthyological report.

Boston Jour. Natural Hist., vol. 3, pp.
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1841. Additional descriptions of, and observations

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1842. Additional descriptions of, and observations

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1842a. [Note on a torpedo taken on Cape Cod.]

Proc. Boston Soc. Natural Hist., vol. 1,

p. 94.

1843. [Description of a new species of flatfish.]

Proc. Boston Soc. Natural Hist., vol. 1,
pp. 130-131.

558

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Storer, David Humphreys — Continued

1843a. [Notes on Sphyraena borealis and Caranx

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vol. 1, pp. 148-149.
1843b. Notice of the discovery of an electrical fish

on our coast. Amer. Jour. Science and

Arts, vol. 44, p. 213; also Ann. Mag.

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1843c. Description of a new species of the torpedo.

Amer. Jour. Science and Arts, vol. 45, pp.

165-170, pi. 3.

1844. [Notes on flat fishes, and on the sunfish,

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1845. [Occurrence of Prionotus tribulus Mitch., In

Massachusetts Bay.] Proc. Boston Soc.
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1845a. [Remarks on a living torpedo taken at

Provincetown.] Proc. Boston Soc. Natural

Hist., vol. 2, p. 71.
1845b. [Alulera schoepfii, as "Balistes aurantiacus

Mitch." at Salem, Mass., and Portland,

Maine.] Proc. Boston Soc. Natural Hist.,

vol. 2, pp. 71-72.
1845c. [Description of a new species of Leptocepha-

lus.] Proc. Boston Soc. Natural Hist.,

vol. 2, pp. 76-77.
1845d. [Description of a new species of Prionotus

from Massachusetts Bay.] Proc. Boston

Soc. Natural Hist., vol. 2, pp. 77-78.
1845e. [Note on a Trichiurus from Massachusetts

Bay.] Proc. Boston Soc. Natural Hist.,

vol. 2, pp. 85-86.

1846. [Occurrence of Palinurus perciformis in Boston

Harbor.] Proc. Boston Soc. Natural Hist.,
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1846a. A synopsis of the fishes of North America.
Mem. Amer. Acad. Arts Sciences, New
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1847. [Descriptions of new species of Alosa and

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1848. [Description of Molella caudacuta.] Proc.

Boston Soc. Natural Hist., vol. 3, p. 5.
1848a. [On a shark taken at Provincetown.] Proc.

Boston Soc. Natural Hist., vol. 3, pp. 11-12.
1848b. [Notes on the Greenland shark, Somniosus

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Natural Hist., vol. 3, pp. 15-16.
1848c. [Occurrence of the hammerhead shark at

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Proc. Boston Soc. Natural Hist., vol. 3,

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1848d. [Description of Carcharias atwoodi.] Proc.

Boston Soc. Natural Hist., vol. 3, pp. 71-72.

Storer, David Hdmphrets — Continued

1853-1867. A history of the fishes of Massachusetts.
Mem. Amer. Acad. Arts and Sciences,
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pp. 309-372, pis. 24-29. Vol. 8, pp.
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263, pis. 36-39.
1854. [Description of Sebastes fasciatus, from Prov-
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1858. [Description of Zeus ocellatus.] Proc. Boston

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1867. A history of the fishes of Massachusetts. 287
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Storer, Horatio Robinson.

1850. Observations on the fishes of Nova Scotia and
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Sullivan, James.

1795. The history of the District of Maine. 421 pp.,
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1913. A biological survey of the waters of Woods
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Swain, Joseph.

1883. A review of the species of Stolephorus found

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Tarr, R. S.

1884. Return to Gloucester Harbor of the young

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TlBBETTS, N. V.

1887. Scarcity of cod and haddock on the coast of

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Towne, Sumner A.

[1942, approx. date.] State of Maine striped bass

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1901. Statistics of the fisheries of the New England

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1910. Annotated list of fishes known to inhabit the

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FISHES OF THE GULF OF MAINE

559

United States Commission of Fish and Fisheries.

1873. Notices in regard to the abundance of fish on

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1882. List of collections made by the fishing vessels
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1894-1905. [Annual statistics of the vessel fisheries
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1898. A manual of fish culture. Report for 1897,
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1905-30. [Statistics of New England fisheries.]
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Verrill, A. E.

1871. On the food and habits of some of our marine

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1872. Marine fauna of Eastport, Maine. Bull.

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1874. Explorations of Casco Bay by the U. S. Fish

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Vladtkov, V. D.

1935. Some unreported and rare fishes from the
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1950. Movements of Quebec shad (Alosa sapidis-
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turaliste Canadien, vol. 77, pp. 121-134, 1
chart; also as Contrib. No. 30, Dept. Fish-
eries, Province Quebec.
Vladtkov, V. D., and R. A. McKenzie.

1935. The marine fishes of Nova Scotia. Proc. Nova
Scotian Inst. Science, vol. 19, pp. 17-113.
Vladtkov, V. D., and D. N. Wallace.

1938. Remarks on populations of shad (Alosa sapi-
dissima) along the Atlantic coast region.
Trans. Amer. Fish. Soc, vol. 67, pp. 52-66.

Walford, L. A.

1938. Effect of currents on distribution and survival

of the eggs and larvae of the haddock (Mel-

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Bull. U. S. Bur. Fish., vol. 49 (Bull. 29),

pp. 1-73.
Wasson, Samuel.

1878. Survey of Hancock County. Ann. Report

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with the 22nd Ann. Report Maine Board

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Welsh, W. W.

1915. Note on the habits of the young of the squirrel

hake and sea snail. Copeia, No. 18, pp. 2-3.
Welsh, William W., and C. M. Breder, Jr.

1924. Contributions to life histories of Sciaenidae of

the eastern United States coast. Bull. U.

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Wheatland, R. H.

1852. Notice of several fishes of rare occurrence.

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1939. The nesting and embryo of Zoarces anguillaris.

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Wilcox, W. A.

1883. Reappearance of young cod hatched by the
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560

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

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INDEX TO COMMON NAMES

Fag*

albacore 338

false 317, 336, 337, 339

great 338

long-finned 317

alewife 86, 101

alligator fish 8, 457

alligator fishes 457

allmouth 532

amberjack 373

American conger 154

American dab 248,253,259

American eel 151

American goosefish 532

American John Dory 297

American pollock 213

American sole 296

anchovies 118

anchovy 118

striped 118, 119

angel shark 17

angler 532

deep-sea 543

anglers 532, 541, 542

deep-sea 542

Arctic eelpout 509, 516

Arctic sculpin 440, 453

Arcticshanny 492, 495, 497, 499

argentine. 133, 139

argentines 139

armored sea robin 467, 471

armored sea robins 467

Atlantic halibut 249

Atlantic mako 23

Atlantic salmon 121

Atlantic slime eel 157

bjlloonfish 526

barn-door skate 57, 61

barracuda, northern 7, 306

barracudas 306

barrelfish 9, 369

basking shark 28

basking sharks 28

bass, channel 425

sea... 389, 407, 424, 430, 473

striped 389, 405, 409, 417

wreck 409

basses, sea 389, 390, 410, 411, 417

bastard cusk 500

bellowsfish 526

bergall 473

big-eye, short 9, 410

big-eyes 410

big skate 58, 63

billfish 167, 170

billfishes 167

Pag*

blackback 276

black-bellied rosefish 430, 437

blackbelly 106

black dogfish 17, 51

black drum 425

blackfish 407,418

black flounder 276

black hake 221

black pilot 369

black ruff 9, 369, 370

black salmon 121

black sculpin 445

black sea bass 407

blenny, serpent 494

snake 491, 494, 497, 498, 499

blenny-like fishes 491, 502, 509, 518

bloody stickleback 311

blueback 86, 106

blue dog.. 20, 38

bluefin tuna 338

bluefish 7, 383, 417

Boston 213

bluefishes 382

blue hake 173, 233

blue martin 358, 360

blue mouth 437

blue perch 473

blue shark 17, 38

bluntnose 378

boar fish 9,438

boar fishes 438

bone shark 28

bonito 336,337, 339, 347

common 317, 337

oceanic 335

striped 317, 335, 336, 339

bonnethead shark 44

bony fishes 80

Boston bluefish 213

Boston hake 221

bramble shark 17, 56

bramble sharks 56

branch herring 101

bream, red 430

breams, sea 361

brier skate 58, 65

brit 88

broadbill 351

brook trout 120

brown shark 17, 26, 43

bullseye 333

burrfish 526, 527

butterfish 363,411

butterfishes 363

561

562

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Page

Canadian plaice 259

capelin 133, 134, 302

cartilaginous fishes 15

cataluf as 410

catfish 503

spotted 507

cat sharks 34

cavalla 317, 347, 349

chain dogfish 17

channel bass 425

chimaera 7, 79

chimaeras 79

chimaeroids 79

chogset 473

Christmas flounder 283

chub 162

chub mackerel 317, 333

clam cracker 74

coalfish 213

cod 173, 182, 196, 199, 203, 216

green 213

rock 182

torn 173, 183, 196

cod family 173

common bonito 317, 337

grenadier 243

hammerhead 45

mullet 305

mummichog 162

pipefish 312

sea robin 467

skate 63, 67

wolfnsh 507

conger, American 150, 154

conger eel 150, 154

congo eel 510

cornetfish 316

cow hake 221

cow-nosed ray 58, 76

cow-nosed rays 76

crampfish 58

crevalle 372, 375, 376, 377, 383

croakers 417

Cuban jack 372, 378, 381

cunner 390, 407, 408, 427, 430, 473, 479

cunner tribe 417, 473

cusk 173,238

bastard 500

cusk eel 7, 173, 517

cusk eels 517

cutlassfish 6, 350

cutlassfishes 350

cyclostomes 9

cyclothone 141, 146

dab, American 248,253,259

deep-water 274

European 259, 274

long rough 259

sand 290

daddy sculpin 445

Page

deep-sea angler 8, 543

deep-sea anglers 542

deep-water dab 274

devil ray 57, 77

devil rays 77

dog, blue 38

smooth 34

dogfish 47

black 17, 51

chain 17

piked 47

shark 18

smooth 17, 34

spiny 17,47, 185

dogfish shark 18

dogfishes, smooth 34

spiny 47

dollarfish 363, 378

dolphin 8, 360

dolphins 360

drum, black 425

red 425

drums 417, 418

dusky shark 17, 41

eagle rays 76

eel 150, 151,492, 500

American 151

conger 154

congo 510

cusk 7, 173,517

European 151

fresh-water 151

long-nosed 150, 518

rock 492,495,497,499

sand 488

sea 154

silver 151,350

slime 150, 157

snake 150, 159

snipe 6, 150, 159

snub-nosed 157

wolf 509,515,517

eelback 283

eelpout 491,510

Arctic 509, 516

eels 150,487,488,517

cusk 517

moray 150

wolf 508

eel sucker 12

electric ray 58

electric rays 58

elver 151

escolar 7, 349

escolars 349

European dab 259, 274

European eel 151

European goosefish 534, 536

European hake 175, 177

European John Dory 297

FISHES OF THE GULF OF MAINE

563

Page

European ocean pout 513

European pilchard 115

European plaice 259

European pollack 213

European sea sturgeon 81

European silver hake 175, 177

European smelt 136

European turbot 290

eyed skate 63

fall herring 100

false albacore 317, 336, 337, 339

fatback 113

filefish 522, 523

orange -- 522, 524, 525

filefishes 520, 521, 525, 526

fish, alligator 457

balloon 526

barrel 369

bellows 526

bill 167, 170

black 407,478

boar 9,438

burr 526,527

butter 363, 411

cat 503

cornet 316

cutlass 6, 350

dollar 363,378

file 522,523

flying 172

fool 283

frost 196

gar.- 168

ghost 500

globe 526

goat 305

handsaw 161

harvest 363, 368

hatchet 141

headlight 141, 142

hog. 524

horse 378

lancet 161

log 369

lump 459, 464

mink 423

monk 532

moon 8,247,378,379

mouse 540

needle 6, 167, 170

oil 349

oyster 527

pilot 370, 372, 373, 375, 377

porcupine 527

rabbit 527

red.- 430

ribband 350

rock 389

rose 211, 390, 407, 408, 409, 430, 438, 473

rudder 369, 372, 373, 375

fish — continued Page

sail 358

sargassum 6, 541

scabbard 350

scour 349

snipe 301

sun 524,529,532

swell 526

tile 426

toad 439,449,518

trigger 520

trumpet 6, 316

unicorn 522, 524,525

viper 145

weak 417, 424, 426

wolf 502,503

wreck 389,407,409

fishes, alligator 457

bill 167

blenny-like 491

boar 438

bony 80

butter 363

cartilaginous 15

cutlass 350

file 520,521,525,526

flat 6,248

flying 171

grammecolepid 299

hatchet 149

head 528

lancet 160

lantern 141

luminescent 141

lump 459

needle 170

pediculate 532

pipe 312

porcupine 525, 526

rock 417,430

rudder 369

sargassum 541, 542

snipe 301

spear 357

sword 351

tile 426

toad 518

trigger 520,521,522, 525

trumpet 316

viper 145

wolf 491,502,509

fishingfrog 532

flatfish 276

flatfishes 6,248

flounder 267,276

black 276

Christmas 283

four-spotted 248,260,270

Georges Bank 249, 276

Gulf Stream 294

pole 285

564

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

flounder — continued Page

rusty 271

sand 248, 260, 290

smooth 249, 283

smooth-back 283

spotted 290

southern 269

summer 248, 260, 267

winter 249,273,276

witch 249, 285

flounders 248

fluke 267

flyingfish 9, 172

flying fishes 171

flying gurnard 7, 172, 472

flying gurnards 472

f oolfish 283

four-bearded rockling 173, 234

four-spined stickleback 307, 311

four-spotted flounder 248, 260, 270

fox shark 32

fresh-water eel 151

fresh-water herring 101

frostfish 196

garfish 167, 168

gar, salt-water 167

silver 167

gars, silver 167

gaspereau 101

Georges Bank flounder 249, 276

ghostfish 500

globefish 526

glut herring 106

goatfish 305

goggle-eyed scad 372, 375, 376, 377

goosefish, American 6,439,532,541

European 534, 536

grammecolepid 299

grayback 101

grayfish 34,47

gray sculpin 449

gray sole 285

gray trout 417

great albacore 338

green cod 213

green-eye 467

green ocean pout 509

green smelt 302

Greenland halibut 248, 258

Greenland sculpin 446

Greenland shark 18, 53

Greenland turbot 258

grenadier, common 243

long-nosed 243, 246

rough-headed 243, 245

grenadiers 243

grilse 121

ground shark 18

grubby 440, 443, 446

Gulf Stream flounder 294

gunnel 492

Pag#

gurnard, flying 172,472

gurnards 467, 470

gurnards, flying 472

gurry shark 53

gurry sharks 53

gymnuridae 76

hacklehead 449

haddock 173, 183, 185, 199, 221, 289

Jerusalem 247

Norway 430

hagfish 5, 10

hagfishes 9

hairtail 350

hake, black 221

blue. 173,233

Boston 221

cow 221

European 175, 177

long-finned 173, 232

mud. 221

New England 173

red 223

silver 173

spotted 173, 230

squirrel 173,221,223,231

white 173,221, 231

hakeling 173, 234

halfbeak 6, 167, 169

halfbeaks 169

halibut 248

Atlantic 249

Greenland 248, 258

Pacific 253,254

hammerhead, common 16, 45

hammerhead sharks 44

handsawfish 161

hardhead 333

hardtail 372, 375, 376, 377

harvestfish 363, 368

hatchetfish, silver 141, 149

hatchetfishes 149

headfishes 528

headlight fish 141, 142

hedgehog skate 67

herring 88, 114, 493, 496

branch 101

fall 100

fresh-water 101

glut 106

Labrador 88

round 85, 87

sea 85, 88, 101, 108

shad 100

smelt 139

summer 106

thread 85, 112

herring tribe 85

heterodontids 16

hickory shad 85, 100

hogchoker 248, 296

FISHES OF THE GULF OF MAINE

565

Page

hogfish -- 524

hook-eared sculpin 440

horsefish 378

horsehead 379

horse mackerel 337, 338

hound, smooth 34

humpback salmon 120, 131

jack 375

Cuban 372,378,381

jacks 371

Japanese slime eel 157

Jerusalem haddock 247

John Dory, American 7, 297, 299,418, 438

European 297

John Dories 297

Johnson's sea bream 361

jumping mullet 305

kelt. 121

killifish 162, 164

killifishes 162

kingfish 348,417,423,426

king mackerel 317, 347, 348, 349

King-o'-Norway 454

king whiting 423

kyak 101, 106

Labrador herring 88

lafayette 423

lake sturgeon 81

lamper 12

lamprey, sea 5, 12

spotted 12

lampreys 9

lancetfish 5, 161

lancetfishes 160

langbarn 497

lant 488

lanternfish 141, 143

lanternfishea 141

launce 8,488,494

sand 488

launces, sand 487

leather jacket 8, 371, 380

lemon sole 276

leopard shark 66

leopard skate 57

linesides 389

ling 221,223

little sculpin 443

little skate. 58, 67

little sturgeon 84

little tunny 336

logfish 369

long-finned albacore 317

long-finned hake 173, 232

longhorn sculpin 440, 447, 449, 455, 456

long-nosed eel 150, 158

long-nosed grenadier 243, 246

long rough dab 259

lookdown 8, 372, 378, 379

luminescent fishes 141

Page

lump 459

lumpfish 459,464

spiny 459,463

lumpfishes 459

lump sucker 459

mackerel 317, 374

chub 317, 333

horse 337, 338

king 317, 347, 348, 349

Spanish 317, 347, 349

yellow 376

mackerels 317,371, 383, 417

mackerel scad 372, 374, 375, 377

mackerel shark 17, 20

sharp-nosed 23

mackerel sharks 20

mailed sculpin 440, 441

mako, Atlantic 23, 354

Pacific 24

man-eater 17, 25

marlin, blue 358, 360

white... 358, 360

marlins 357

marlin-spike 243

menhaden 85, 113

minkfish. 423

minnow, salt-water 162

sheepshead 165

molligut 532

monkfish 532

moonfish 8, 247, 372, 378, 379

moray s 150

mossbunker 113

mousefish 541

mud hake 221

mullet 7,305,417

common 305

jumping 305

red... 305

striped 305

mullets 305

mummichog, common 162

striped 164

mummichogs 162

mummie 162, 164

muttonfish 510

needlefish 6, 167, 170

needlefishes 170

New England hake 173

Newfoundland turbot 258

New York Plaice 290

nine-spined stickleback 307

northern barracuda 306

Norway haddock 430

numbfish 58

ocean catfish 503

perch 430

ocean pout 509, 510

European 513

green 509

566

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Page

ocean pouts 508

ocean sunfishes 528

ocean whitefish 503

oceanic bonito 335, 337

oilfish 349

onion-eye 245

opah 8, 247

opahs 247

orange filefish 522, 524, 525

oysterfish 527

Pacific halibut 253,254

Pacific mako 24

parr 121

pearlfish 144

pearlsides 141, 144

pediculate fishes 532

pelagic pipefish 312, 341

perch 473

blue 473

ocean 430

sea 405,473

white 389, 390, 405, 409, 417

pike, sea 167

piked dogfish 47

pilchard, European 115

pilotfish 9, 370, 372, 373, 375, 377

pilot sucker 485

pipefish, common 312, 357

pelagic 312, 314

pipefishes 312

plaice, Canadian 259

European 259

New York 290

plaicefish 267

plaintail 349

pogy 113

pole flounder 285

pollack, European 213

pollock 173, 183, 185, 200, 203, 213, 221

American 213

pomfrets 36 1

pompanos 374, 375, 382, 383, 417

porbeagle 20

porcupine fish 527

porcupine fishes 525, 526

porgy 411

porgies 411, 417

Port Jackson sharks 16

Portuguese shark 18, 52

pout, eel 491,510

ocean 509, 510

prickly skate 57, 70

puffer 526, 527

puffers 525, 526

rabbitfish 527

radiated shanny 492, 495, 498

rat-tail 243, 245

raven 454

sea 454

Pag»

ray, cow-nosed 58, 74

devil 57,77

electric 58

sting 58, 74

rays 15, 57

cow-nosed 76

devil 77

eagle 76

electric 58

sting 74

red bream 430

red drum 425

redfish 430

red hake 223

red mullet 305

red sculpin 454

red sea perch 430

remora 485, 487

remoras 484

requiem sharks 34

ribbandfish 350

robin 467

rock 389

rock cod 182

rock eel 7, 492,495, 497, 499

rockfish 389

rockfishes 417, 430

rockling, three-bearded 173, 237

rockling, four-bearded 173, 234

rosefish 211,

393, 407, 408, 409, 427, 430, 438, 473

black-bellied 430, 437

rosetted skate 66

rough scad 377

rough-headed grenadier 243, 245

round herring 85, 87

round robin 374

rudderfish 369,372,373, 375

rudderfishes 369

runner 376

rusty flounder 271

sailfish 6, 352, 358

sailfishes 357

saithe 213

salmon 120, 121, 506

Atlantic 121

black 121

humpback 120, 131

sea 121

silver 120, 121, 133

salmons 119

salt-water gar 167

salt-water minnow 162

salt-water smelt 135

Salter 120

sand bar shark 43

sand dab 290

sand eel 488

sandflounder 248,260,290

sand launce 488

FISHES OF THE GULF OF MAINE

567

Page

sand launces 487

sandshark 17, 18

sand sharks 18

sand smelt 302

sardine 88

sargassum fish 6, 541

sargassum fishes 541, 542

saurel 372, 375, 376, 377

saury . . 170

sawbelly 101

scabbardfish 350

scad, goggle-eyed 372, 375, 376, 377

mackerel 372,374, 375, 377

rough 377

scourfish 349

sculpin, Arctic 440, 453

black 445

daddy 445

gray 449

Greenland 446

hook-eared 440

little 443

longhorn 440, 447, 449, 455, 456

mailed 440, 441

red 454

sea 454

shorthorn 439, 440, 444, 445, 449, 455

staghorn 440, 452

sculpins 417, 418, 439, 472, 518

scup 407,408,411,416,473

sea bass 389, 390, 407, 424, 430, 473

black 407

sea basses 383, 389, 410, 411, 417

sea bream, Johnson's 36 1

sea breams 36 1

sea eel 151

sea herring 85,88, 101, 108

sea-horse 6, 315

sea-horses 314, 315

sealamprey 5, 12

seaperch 405, 473

red 430

sea pike 167

seapoacher 457

seapoachers 457

sea raven 6, 440, 454

sea ravens 439

sea robin 467

armored 467, 471

common 467

striped 467, 470

searobins 439,467,471,473

armored 467

sea salmon 121

sea sculpin 454

sea snail 464, 466

striped 464,466

sea snails 464

sea sturgeon 81

European 81

Pag*

sea trout 120, 417

serpent blenny ._ 494

shad 86, 108

herring 100

hickory 85

shanny 492,495,497,498

Arctic 492,495,497,499

radiated 492,495,498

shark

angel 17

basking 28

blue 17,38

bone 28

bonnethead 44

bramble 17,56

brown 17, 26, 43

dogfish 18

dusky 17, 41

fox 32

Greenland 18, 53

ground 18

gurry.. 53

leopard 37

mackerel 17, 20

Portuguese 18, 52

sand 17, 18

sand bar 43

sharp-nosed 17, 40

sleeper 53

spiny 56

tiger 17,37

whale 29

white 17, 25

shark pilot 372

shark sucker 485, 486, 487

shark suckers 484

sharks

basking 28

bramble 56

cat 34

gurry 53

hammerhead 44

mackerel 20

Port Jackson 16

requiem 36

sand 18

thresher 32

sharp-nosed shark 17, 40

sharp-nosed mackerel shark 23

sharp-tailed sunfish 520, 530, 531

sheepshead 363,411,416

sheepshead minnow 165

shiner 302, 363, 378

short big-eye 410

shorthorn sculpin 439, 440, 444, 445, 449, 455, 456

short-nosed sturgeon 81, 84

shovelhead 16, 44

silver eel 151, 350

silver gar 167

silver gars 167

568

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Page

silver hake 173

European 175, 177

silver-hake family 173

silver hatchetfish 149

silversalmon 120, 121, 133

sil verside 302

waxen 302, 304

silversides 118, 302

skate, barn-door 57, 61

big 58,63

brier 58, 65

common 63, 67

eyed 63

hedgehog 67

leopard 57, 66

little 58, 67

prickly 57, 70

rosetted 66

smooth-tailed 70

spotted 63

starry 72

summer 67

thorny 57, 72

winter 63

skates 15, 57, 60

skilligalee 358

skipjack 337, 363

skipper 170

sleeper shark 53

slimeeel 150, 157

Atlantic 157

Japanese 157

smelt 133, 135,302

European 136

green 302

herring 139

salt-water 135

sand 302

smelts 133

smolt 121

smooth dog 34

smooth dogfish 17, 34

smooth dogfishes 34

smooth flounder 249,283

smooth-back flounder 283

smooth hound 34

smooth-tailed skate 70

snake blenny 491, 494, 497, 498, 499

snakeeel 150, 159

snapper 383

snipeeel 6, 150, 159

snipefish 6, 301

snipefishes 301

snub-nosed eel 157

sole 276

American 296

gray 285

lemon 276

soles 248

southern flounder 269

Page

Spanish mackerel 317, 347, 349

spearfishes 351,352,357

sperling 88,302

spiny dogfish 17, 47, 185

spiny dogfishes 47

spiny lumpfish 459, 463

spiny shark 56

spot 417,423

spotted catfish 507

spotted flounder 290

spotted hake 173, 230

spotted lamprey 12

spotted skate 63

spotted turbot 290

spotted wolffish 502, 503, 507

squeteague 417, 424

squirrel hake 173, 221, 223, 231

staghorn sculpin 440, 452

starry skate 72

stickleback 308

bloody 311

four-spined 307, 311

nine-spined 307

three-spined 307, 308

two-spined 307, 308, 310

sticklebacks 307

stingaree 74

sting ray 58, 74

sting rays 74

whip-tailed 74

stomias 141, 147

stomiatids 146

striped anchovy 119

striped bass 389,405,409,417

striped bonito 317, 335, 336, 339

striped mullet 305

striped mummichog 164

striped sea robin 467, 470

striped sea snail 464, 466

striper 389

sturgeon, lake 81

little 84

sea 81

short-nosed 81, 84

sturgeons 80

sucker, lump 459

pilot 485

shark 485,486,487

swordfish 485, 486

white-tailed 485

suckers, shark 4S4

summer flounder 248,260,267

summer herring 106

summer skate 67

sunfish 524, 529, 532

sharp-tailed 529, 530, 531

sunfishes, ocean 528

swellfish 526

swell toad 526

swi veltail 32

FISHES OF THE GULF OF MAINE

569

Page

swordfish 24, 351, 358

swordfishes 351

swordfish sucker 485, 486

tarpon 85, 87

tarpon tribe 85

tautog 369, 390, 407, 408, 409, 427, 430, 473, 478

ten pounder 85, 86

thornback 308

thornfish 308

thorny skate 57, 70

threadfin 8,381

thread herring 85, 112

three-bearded rockling 173, 237

three-spined stickleback 307, 308

thresher 17,32

thresher sharks 32

tiger shark 17, 37

tilefish 5,426,473

tilefishes 426

toadfish- 6,439,449,518

toadfishes 518

tobacco box 67

tomcod 173, 183, 196

torpedo 57, 58

torpedoes 15, 57, 58

torsk 238

triggerfish 7,418,520

triggerfishes 520, 521, 522, 525

trout 120, 506

brook 120

gray 417

sea 120,417

trumpetfish 6, 316

trumpetfishes 316

tuna 317,337, 338

bluefin 338

tunny 338

little 336

turbot 524

European 290

Greenland 258

turbot — continued P»i

Newfoundland 258

spotted 290

tusk 238

two-spined stickleback 307, 308, 310

unicorn fish 522, 524, 525

viperfish 141, 145

viperfishes 145

waxen silverside 302, 304

weakfish 417,424,426

weakfishes 390, 417

whale shark 29

whip-tailed sting rays 74

whitebait 118, 302

white chin 478

white hake 173,221,231

white marlin 358, 360

white perch 389,390,405,409,417

white shark 17,25

whitefish, ocean 503

whitefishes 120

white-tailed sucker 485

whiting 173,423

king 423

windowpane 290

winter flounder 249, 273, 276

winter skate 63

witch flounder 249,285

wolf eel 509,515,517

wolf eels 508

wolffish 502,503

common 502, 503, 507

spotted 502,503,507

wolffishes 491, 502, 509

wrasses 473

wreck bass 409

wreckfish 389, 407, 409

wrymouth 7,9,491,500

wrymouths 500, 509

yellow jack 376

yellow mackerel 376

yellowtail 249, 271, 375

2109U— 63 57

INDEX TO SCIENTIFIC NAMES

Page

Ablennes 167

Ablennes hians 168

acadianus, Glyptocephalus 288

acanthias, Squalus 47

Acanthocottus 443, 445, 449

Achiridae 248

Achirus fasciatus 296

Acipenser brevirostrum 84

f ul vescens 81

oxyrhynchus. 81

sturio 81

Acipenseridae 80

aculeatus, Argyropelacus 149

Gasterosteus 308

acus, Tylosurus 167

adspersus, Tautogolabrus 473

aeglefinus, Melanogrammus 199

aeneus, Myoxocephalus 443

aestivalis, Pomolobus 106

Aethoprora eff ulgens 143

affine, Myctophum 143

affinis, Hydrolagus 79

Agnatha 9

Agonidae 457

Agonus decagonus. 457

alalunga, Thunnus 317

albida, Makaira 360

albiguttatus, Paralichthys 269

Alectis ciliaris 381

crinitus 381

alepidotus, Peprilus 368

Alepisauridae 160

Alepisaurus ferox 161

alleterat us, Euthynnus 336

Alopias vulpinus 32

Alopiidae 32

Alosa sapidissima 108

alpinus, Salvelinus 121

altus, Pseudopriacanthus 410

Alutera schoepfii... 522, 524

scripta .-- 522, 525

americana, Morone 405

americanus, Ammodytes 488

Hemitripterus 454

Istiophorus 357

Lophius 532, 534

Lumpeuus lumpretaef ormis 495

Macrozoarces 510

Polyprion 409

Pseudopleuronectes 276

Xenolepidichthys 299

Amitra 468

Page

Ammodytes americanus 488

dubius 488,491

dubius hudsonicus 491

lanceolatus 488,489

tobianus 488, 489, 490

Ammody tidae 487

ampla, Makaira 358

amplus, Tetrapterus 358

Anarhichadidae 502

Anarhichas latifrons 502, 503

lupus 503

minor 507

Anchoa hepsetus 119

mitchilli 118

Ancylopsetta quadrocellata 271

Anguilla chrysypa 151

rostrata 151

vul garis 151

anguillaris, Zoarces 510

Anguillidae 150

Antennariidae 54 1

Antigonia capros 438

Antimora rostrata 233

viola 233

antiquorum, Hippocampus 316

Apeltes quadracus 311

Archosargus probatocephalus 416

arctifrons, Citharichthys 248, 294

argentata, Motella 226

argentatus, Gaidropsarus 236

Argentina silus 139

Argentinidae 139

Argyropelecus aculeatus 149

olfersi 149

armatus, Nematonurus 243

Artediellus 439

atlanticus 440

uncinatus 440

Aspidophoroides monopterygius 457

Aspidophoroididae 457

Atherinidae 302

Athlennes hians 168

atlanticus, Artediellus 440

Lycodes 509

Neoliparis 464, 467

Tarpon 87

auratus, Mullus 305

bairdii, Macrourus 243,245,353

Balistes capriscus 520

carolinensis 520

vetula 521

Balistidae 520

barbatum, Echiostoma 353

570

FISHES OP THE GULF OF MAINE

571

Page

BatoideL 57

Batrachoididae 518

beanii, Limanda — 274

Triglops 442

Belonidae 167

berglax, Macrourus 245

beryllina, Menidia 304

biaculeatus, Gasterosteus 310

bilinearis, Merluccius 173

birostris, Manta 77

bispinosus, Gasterosteus 310

bonasus, Rhinoptera 76

borealis, Sphyraena 306

Bothidae 248

brachyptera, Remora 486

Brama raii 362

Bramidae 361

Branchiostegidae 426

brevicaudata, Brevootia 116

brevirostris, Hippocampus 316

Negaprion 36

brevirostrum, Acipenser 84

Brevootia brevicaudata 116

tyrannus 113

brownii, Stolephorus 119

Brosme brosme 238

brosme, Brosme 238

brucus, Echinorhinus 56

budegassa, Lophius 534

callarias, Gadus 182

canis, Mustelus 34

capriscus, Balistes 520

Caproidae 438

capros, Antigonia 438

Carangidae 371

Caranx crysos 376

hippos 375

carapinus, Coryphaenoides 243

carcharias, Carcharodon 25

Carcharias taurus 18

Carchariidae 18

Carcharhinidae 36

Carcharhinus 41

milberti 26, 42, 43

obscurus 41

platyodon 43

plumbeus 43

Carcharodon carcharias 25

Careproctus longipinnis 464

ranulus 464

carminatus, Coelorhynchus 246

carolinensis, Balistes 520

carolinus, Prionotus 467

cataphractus, Gasterosteus 310

cavalla, Scomberomorus 349

Centridermichthys unicinatus 441

Centrolophidae 369

Centrolophus niger 370

Centropristes striatus 407

Centroscyllium fabricii 51

Centroscymnus coelolepis 52

Page

centroura, Dasyatis 74

Raja -.- 74

Cephalacanthus volitans 472

cephalus, Mugil... 305

Ceratacanthus 524

Ceratias holbolli 543

Ceratiidae 542

cervinum, Lepophidium 517

Cestracion tiburo 44

Cetorhinidae 28

Cetorhinus maximus 28

chamaeleonticeps, Lopholatilus 426

Chauliodontidae 141, 145

Chauliodus sloani 145

chesteri, Urophycis 232

Chiasmodon 353

Chilomycterus schoepfii 527

Chimaerae 79

Chimaeridae 79

Chloroscombrus chrysurus. . 375

Chondrichthyes 15

chrysops, Stenotomus 411

chrysurus, Chloroscombrus 375

chrysypa, Anguilla 151

chuss, Urophycis 223

ciliaris, Alectis 381

ciliatus, Monacanthus 522,523,524

cimbrius, Enchelyopus 234

Citharichthys arctifrons 248, 294

unicornis 295

Clupea harengus 88

pallasii 93

pilchard us 115

Clupeidae 85

coelolepis, Centroscymnus 52

Coelorhynchus carminatus 246

colias, Pneumatophorus 333

Scomber 333

Conger conger 156

oceanica 154

conger, Conger 156

Leptocephalus 154

Congridae.. 150

Coregonus quadrilateralis 120

Coryphaena hippurus 360

Cory phaenidae 360

Coryphaenoides carapinus 243

rupestris 243,245

Cottidae 439

Cottunculus 439

microps 453

crinitus, Alectis 381

cromis, Pogonias 425

cruentifer, Omochelys 159

Pisodonophis 159

crumenophthalmus, Trachurops 377

Cryptacanthodes maculatus 500

Cryptacanthodidae 500

crysos, Caranx 376

cubensis, Hynnis 378, 381

572

FISHERY BULLETIN OP THE FISH AND WILDLIFE SERVICE

Fan

cubensis, Sqaulus 49

cuvier, Galeocerdo 37

Cybiidae 317

Cyclogaster montagui 465

Cy clopteridae 459

Cyclopterus lumpus 459

Cyclothone signata 146

cy noglossus, Glyptocephalus 285

Cynoscion regalis 417

Cyprinodon variegatus 165

Cypselurus heterurus 172

Dactylopteridae 472

Dactylopterus volitans 472

dactyploterus, Helicolenus 437

Dalatias licha 18, 55

Dalatiidae 53

Dasyatidae 74

Dasyatis centroura 74

Dasybatus hastatus ._ 74

marinus 74

decagonus, Agonus 457

Leptagonus 457

Decapterus maoarellus 374

punctatus 374

dentatus, Paralichthys 59,267

diaphanes, Raia 63

Diaphus effulgens 142

dignabilis, Pseudopleuronectes 277

Diodontidae 525

diplana, Sphyrna 46

dubius, Ammodytes 488

ductor, Nauerates 372

Dussumieridae 85

Echeneidae 484

Echeneis naucrateoides 485

nauerates 485

Echinorhinidae 56

Echinorhinus brucus 56

Echiostoma barbatum 353

effulgens, Aethoprora 143

Diaphus 142

eglanteria, Raja. 65

Elasmobranchii 15

Elopidae 85

Elops saurus. 86

Enchelyopus cimbrius 234

Engraulidae 118

ensis, Gaidropsarus 237

eperlanus, Osmerus 136, 137

erinacea, Raja 67

esmarkii, Lycodes 509,510

Etmopterus princeps 17, 47

Etrumeus sadina 87

Euleptorhamphus velox 169

Eumicrotremus spinosus 463

terrae novae 463

Euthynnus alleteratus 336

pelamis 335, 337

e volans, Prionotus 470

Fag*

Exocoetidae 171

faber, Zeus 299

fabricii, Centroscyllium 51

fasciatus, Achirus 296

Sebastes 433

ferox, Alepisaurus 161

Stomlas 147

ferruginea, Limanda. . 271

Fistularia serrata.. 317

tabacaria 316

Fistulariidae 316

floridae, Siphostoma. . 314

fontinalis, Salvelinus 120

frigidus, Lycodes. 509

fulvescens, Acipenser 81

fulvus, Physiculus 233

Fundulus 162

heteroclitus 162

majalis 164

fuscum, Siphostoma 312

fuscus, Syngnathus 312

Gadidae 173

Gadus callarias 182

pollachius 213

Gaidropsarus argentatus 236

ensis 237

Galeocerdo cuvier 37

Galeorhinus laevis 34

Galeus glaucus 38

garmani, Raja 66

Gasterosteidae 307

Gasterosteus aculeatus 308

biaculeatus 310

bispinosus 310

cataphractus 310

gladiunculus 310

wheatlandi 310

Gempylidae 349

giganteus, Reganichthys 543,545

glaciale, Myctophum 143

glacialis, Liopsetta 285

gladius, Xiphias 351

gladiunculus, Gasterosteus 310

glauca, Prionace 38

glaucus, Galeus 38

Isurus . 24

glutinosa, Myxine 10

Glyptocephalus acadianus 288

cy noglossus 285

Gonostomidae 141, 146

goodei, Hymenocephalus 243

gorbuscha, Oncorhynchus 131

Grammicolepidae 299

groenlandicus, Myoxocephalus 446

gunnellus, Pholis 391,492

Gymnelis viridis 509

Gymnocanthus 439

tricuspis 452

ventralis 453

Gymnosarda pelamis 335

FISHES OF THE GULF OF MAINE

573

Page

hacheyi, Lycodes reticulatus 517

harcngus, Clupea 88

hastatus, Dasybatus 74

Helicolenus dactylopterus 437

maderensis 437, 438

Hemirhamphidae 169

Hemitripteridae 439

Homitripterus americanus 454

hepsetus, Anchoa 119

heteroclitus, Fundulus 162

heterurus, Cj pselurus 172

hians, Ablennes 168

Athlennes 168

Hippocampidae 315

Hippocampus. 314

antiquorum 316

bre virostris 316

hudsonius 315

Hippoglossidae 248

Hippoglossoides limandoides 260

platessoides 259

hippoglossoides, Reinhardtius 258

Hippoglossus hippoglossus 249

hippoglossus, Hippoglossus 249

hippos, Caranx 375

hippurus, Coryphaena 360

hispidus, Monacanthus 522,523

Histrio pictus 541

histrio, Pterophryne 541

holbolli, Ceratias 543

Holocephali 79

hudsonius, Hippocampus 315

humboldtii, Scopelus 144

Hydrolagus affinis 79

Hymenocephalus goodei 243

Hynnis cubensis 378, 381

Hyporhamphus roberti 169

unif asciatus 169

Icelus 439

Istiophoridae 357

Istiophorus americanus 357

Isuridae 20

Isurus glaucus 24

oxyrinchus 23

punctatus 20

japonicus, Pneumatophorus 234

Katsuwonidae 317

kisutch, Oncorhynchus 133

Labridae 473

laevigatus, Lophius 540

laevis, Galeorhinus 34

Raja 61

Lamna nasus 20

Lampadena 353

lampetraef ormis, Lumpenus 494

Lampridae 247

Lampris luna 247

regius 247

lanceolatus, Ammodytes 488,489

Masturus 529, 530, 531

Page

lathami, Trachurus 377

latifrons, Anarhichas 502,503

Leiostomus xanthurus 423

Lepophidium cervinum 517

Leptagonus decagonus 457

Leptocephalus conger 154

Leptoclinus maculatus 497

leptosomus, Simenchelys 157

lepturus, Trichiurus 350

lethostigmus, Paralichthys 269

lewini, Sphyrna 46

licha, Dalatias 18, 55

Limanda beanii 274

ferruginea 271

limanda 274

limanda, Limanda 274

limandoides, Hippoglossoides 260

lineatus, Roccus 389

Liopsetta glacialis 285

putnami 283

Liparidae 464

Liparis liparis 466

liparis, Liparis 466

longipinnis, Careproctus 464

Lopholatilus chamaeleonticeps 426

Lophiidae 532

Lophius americanus 532, 534

budegassa 534

laevigatus 540

piscatorius 532, 534, 537, 540

vaillanti 534

Lophopsetta maculata 290

Lotella maxillaris 234

Lumpenidae 491

Lumpenus lampetraeformis 494

lumpretaeformis 494

lumpretaeformis americanus 495

terrae-no vae 495

lumpretaeformis, Lumpenus 494

Iumpus, Cyclopterus 459

luna, Lampris 247

lupus, Anarhichas 503

Lycenchelys paxillus 509

verrillii 515

Ly cichthys 507

Lycodes atlanticus 509

esmarkii 509, 510

frigidus 509

reticulatus 516

reticulatus hacheyi 517

reticulatus, var. macrocephalus 517

Lycodes vahlii 510

zoarchus 510

macarellus, Decapterus 374

macrocephalus, var. of Lycodes reticulatus 517

Macrorhamphosidae 301

Macrorhamphosus scolopax 301

Macrostoma 353

Macrouridae 243

574

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Page

Macrourus bairdii 243, 245, 353

berglax 245

Macrozoarces 491

americanus 510

maculat a, Lophopsetta 290

maculatus, Cryptacanthodes 500

Leptoclinus 497

Scomberomorus 347

Sphaeroides 526

maderensis, Helicolenus 437, 438

majalis, Fundulus 164

Makaira albida 360

ampla 358

Mallotus villosus 134

Mancalias shufeldti 543

uranoscopus 543, 545

Manta birostris 77

marinus, Dasybat us 74

Petromyzon 12

Sebastes 430

Tylosurus 167

massachusettensis, Monacanthus 524

Masturus lanceolatus 529, 530, 531

Maurolicidae 141

Maurolicus pennanti 144

maxillaris, Lotella 234

maximus, Cetorhinus 28

mediocris, Pomolobus 100

Melanogrammus aeglefinus 199

Menidia beryllina 304

cerea 305

menidia 302

notata 302

menidia, Menidia 302

Menticirrhus saxatilis 423

Merlucciidae 173

Merluccius 1 73-177

bilinearis 173

merluccius 177

merluccius, Merluccius 177

Merulinus 467

microps, Cottunculus 453

microcephalus, Somniosus 53

Microgadus tomcod 196

milberti, Carcharhinus 26,42,43

minjatum, Peristedion 471

minor, Anarhichas 507

miriceps, Trigonolampa 148

mitchilli, Anchoa 118

Stolephorus 118

Mobulidae 77

mokarran, Sphyrna 46

Mola mola 528, 529, 532

mola, Mola 528, 529, 532

Molidae 528

Monacanthidae 521

Monacanthus ciliatus 522, 523, 524

hispidus 522, 523

massachusettensis 524

Monolene sessilicauda 248, 295

Page

monopterygius, Aspidophoroides 457

montagui, Cyclogaster 465

Neoliparis 465

mordax, Osmerus 135

Morone americana 405

Motella argentata. - 226

Mugil cephalus 305

Mugilidae 305

Mullidae 305

Mullus auratus 305

surmulletus 59

Mustelus canis 34

Myctophidae 141

Myctophum 142, 143

Myctophum affine 143

glaciale 143

opalinum 143

Myliobatidae 76

Myoxocephalus aeneus 443

groenlandicus 446

octodecimspinosus 449

scorpius 439, 445

Myxine glutiuosa 10

Myxinidae 9

Narcacion nobilianus 58

nasus, Lamna 20

Naucrates ductor 372

naucrates, Echeneis 485

naucrateoides, Echeneis 485, 486

Negaprion bre virostris 36

Nematonurus armatus 243

Nemichthyidae 150

Nemichthys scolopaceus 159

Neoliparis atlanticus 464, 467

montagui 465

Nezumia 243

nicholsi, Stomioides 147

niger, Centrolophus 370

nobiliana, Torpedo 58

nobilianus, Narcacion 58

notata, Menidia 302

Notoscopelus 353

oblongus, Paralichthys 270

obscurus, Carcharhinus 41

oceanica, Conger 154

ocellata, Raja 63, 68

Zenopsis 297

ocellatus, Sciaenops 425

octodecimspinosus, Myoxocephalus 449

oglinum, Opisthonema 112

olfersi, Argyropelecus 149

Oligoplites saurus 380

ommatistius, Triglops 441

Omochelys cruentifer 159

Oncorhynchus gorbuscha. 131

kisutch 133

onitis, Tautoga 478

opalinum, Myctophum 143

Ophichthyidae 150

Ophidiidae 517

FISHES OF THE GULF OF MAINE

575

Fan

Opisthonema oglinum 112

Opsanus tau 518

Osmeridae 133

Osmerus eperlanus 136, 137

mordax 135

Osteichthyes - 80

Otolithidae 417

oxyrhynchus, Acipenser 81

oxyrinchus, Isurus 23

Palinurichthys percif ormis 369

pallasii, Clupea 93

Paragaleus pectoralis 41

Paralichthyidae 248

Paralichthys albiguttatus 269

dentatus 59, 267

lethostigmus 269

oblongus -- 270

Paraliparis 464

parasiticus, Simenchelys 157

paru, Peprilus 368

paxillus, Lycenchelys 509

pectoralis, Paragaleus 41

pelagicum, Siphostoma 314

pelagicus, Syngnathus 314

pelamis, Euthynnus 335.337

Gymnosarda 335

pennanti, Maurolicus 144

Peprilus alepidotus 368

paru 368

percif ormis, Palinurichthys 369

Peristediidae 467

Peristedion miniatum 471

Petromyzon marinus 12

Petromyzonidae 9

Pholidae 491

Pholis gunnellus 391, 492

Physiculus fulvus 233

pictus, Histrio 541

pilchardus, Clupea 115

pingeli, Triglops 441

pinnatus, Synaphobranchus 158

piscatorius, Lophius 532, 534, 537, 540

Pisodonophis cruentifer 159

platessa, Pleuronectes 59

platessoides, Hippoglossoides 259

platyodon, Carcharhinus 43

Pleuronectes platessa 59

Pleuronectidae 248

plumbeus, Carcharhinus 43

Pneumatophorus colias 333

japonicus 334

Poeciliidae 162

Pogonias cromis 425

pollachius, Gadus 213

Pollachius virens 213

Polyprion americanus 409

Pomatomidae 382

Pomatomus saltatrix 383

Fag*

Pomolobus aestivalis 106

mediocris 100

pseudoharengus 101

Poronotus triacanthus 363

pretiosus, Ruvettus 349

Priacanthidae 410

princeps, Etmopterus 17,47

Taractes 361

Prionace glauca 38

Prionotus carolinus 467

evolans 470

strigatus 470

probatocephalus, Archosargus 416

pseudoharengus, Pomolobus 101

Pseudopleuronectes americanus 276

dignabilis 277

Pseudopriacanthus altus 410

Pterophryne histrio 541

punctatus, Decapterus 374

Isurus 20

Stichaeus 497

Pungitius pungitius 307

pungitius, Pungitius 307

putnami, Liopsetta 283

quadracus, Apeltes 311

quadrilateralis, Coregonus.. 120

quadriloba, Rhinoptera 76

quadrocellata, Ancylopsetta 271

radiata, Raja 72

Raia diaphanes 63

raii, Brama 362

Raja centroura 74

eglanteria 65

erinacea 67

garmani 66

laevis 61

ocellata 63, 68

radiata 72

scabrata 72

senta 70

stabulif oris _. 61

Rajidae 60

ranulus, Careproctus 464

regalis, Cynoscion 417

Scomberomorus 348

Reganichthys giganteus 543, 545

regius, Lampris 247

Urophycis 228, 330

Reinhardtius hippoglossoides 258

Remora brachyptera 486

remora 487

remora, Remora 487

Remoropsis 486

reticulatus, Lycodes 516

retifer, Scyliorhinus 34

Rhinoptera bonasus 76

quadriloba 76

Rhinopteridae 76

roberti, Hyporhamphus 169

576

FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Page

Roccus lineatus 389

saxatilis 389

rostrata, Anguilla 151

Antimora 233

rostratus, Somniosus 55

rupestris, Coryphaenoides 243,245

Ruvettus pretiosus 349

sadina, Etrumeus 87

salar, Salmo 121

Salmo salar 121

Salmonidae 119

saltatrix, Pomatomus 383

Salvelinus fontinalis 120

alpinus 121

sapidissima, Alosa 108

Sarda sarda 337

sarda, Sarda 337

saurus, Elops 86

Oligoplites 380

Scomberesox 170

saxatilis, Menticirrhus 423

Roccus 389

scabrata, Raja 72

schoepfii, Alutera 522,524

Chilomycterus 527

Sciaenidae 417

Sciaenops ocellatus 425

Scoliodon terrae-novae 40

scolopaceus, Nemichthys 159

scolopax, Macrorhamphosus 301

Scomber colias 333

scombrus 317

Scomberesocidae 170

Scomberesox saurus 170

Scomberomorus cavalla 349

maculatus 347

regalis 348

Scombridae 317

scombrus, Scomber 317

Scopelus humboldtii 144

Scorpaenidae 430

scorpius, Myoxocephalus 439,445

scripta, Alutera 522,525

Scyliorhinidae 34

Scyliorhinus 34, 59

Scyliorhinus retifer 34

Sebastes fasciatus 433

marinus 430

viviparus 431

Selachii 15

Selene vomer 379

senta, Raja 70

Seriola zonata 373

Serranidae 389

serrata, Fistularia 317

sessilicauda, Monolene 248, 295

eetapinnis, Vomer 378

6hufeldti, Mancalias 543

signata Cyclothone 146

Pag«

silus, Argentina 139

Simenchelyidae 150

Simenchelys leptosomus 157

parasiticus 157

Siphostoma floridae 314

fuscum 312

pelagicum 314

sloani, Chauliodus 145

Somniosus microcephalus 53

rostratus 55

Sparidae 411

Sphaeroides maculatus 526

Sphyraena borealis 306

Sphyraenidae 306

Sphyrna diplana 46

lewini 46

mokarran 46

tiburo 44, 45

tudes 46

zygaena 45

Sphyrnidae 44

spinosus, Eumicrotremus 463

Squalidae 47

Squalus acanthias 47

cubensis 49

stabuliforis, Raja 61

Stenotomus chrysops 411

versicolor 411

Sternoptychidae 141, 149

Stichaeidae 491

Stichaeus punctatus 497

Stolephorus brownii 119

mitchilli 118

Stomias 141, 145, 147, 148

ferox 147

Stomiatidae 141, 146

Stomioides 141, 145, 147, 148

nicholsi 147

striatus, Centropristes 407

strigatus, Prionotus 470

Stromateidae 363

Strongylura 167

sturio, Acipenser 81

subbif urcata, Ulvaria 498

surmulletus, Mullus 59

Synaphobranchidae 150

Synaphobranchus pinnatus 158

Syngnathidae 312

Syngnathus fuscus 312, 314

pelagicus 314

Syrictes.. 312

tabacaria, Fistularia 316

Taractes princeps 361

Tarpon atlanticus 87

tau, Opsanus 518

taurus, Carcharias 18

Tautoga onitis 478

Tautogolabrus adspersus 473

tenuis, Urophycis 221

FISHES OF THE GULF OF MAINE

577

Pagt

terrae-novae, Lumpenus lumpretaeformis 495

Scoliodon 40

Triglops ommatistius 442

Tetraodontidae 525

Tetrapterus amplus 358

Thunnidae 317

Thunnus thynnus 338

alalunga 317

thynnus, Thunnus 338

tiburo, Cestracion 44

Sphyrna 44, 45

tobianus, Ammodytes -- 488, 489, 490

tomcod, Microgadus 196

Torpedinidae 58

Torpedo nobiliana 58

Trachurops 377

crumenophthalmus 377

Trachurus lathami 377

trachurus 377

trachurus, Trachurus 377

triacanthus, Poronotus 363

Triakidae 34

Trichiuridae 350

Trichiurus lepturus 350

tricuspis, Gymnocanthus 452

Triglidae - 467

Triglops beanii 442

ommatistius 441

ommatistius terrae-novae 442

pingeli 441

Trigonolampa. 141, 145, 148

miriceps 148

tudes, sphyrna 46

Tylosurus acus 167

marinus 167

Typhloeeratias 545

tyrannus, Brevoortia 113

Ulvaria subbifurcata 498

uncinatus, Artediellus 440

Centridermichthys 441

Pan

unicornis, Citharichthys 295

unifasciatus, Hyporhamphua 169

uranoscopus, Mancalias 543, 545

Urophycis chesteri 232

chuss 223

regius 228, 230

tenuis 221

vahlii, Lycodes 510

vaillanti, Lophius 534

variegatus, Cyprinodon 165

velox, Euleptorhamphus 169

ventralis, Gymnocanthus 453

verillii, Lycenchelys 515

versicolor, Stenotomus 411

vetula, Balistes 521

villosus, Mallotus 134

viola, Antimora 233

virens, Pollachius 213

viridis, Gymnelis 509

viviparus, Sebastes 431

Zoarces 513

volitans, Dactylopterus 472

Cephalacanthus 472

Vomer setapinnis 378

vomer, Selene 379

vulgaris, Anguilla 151

vulpinus, Alopias 32

wheatlandi, Gasterosteus 310

xanthurus, Leiostomus 423

Xenolepidichthys americanus 299

Xiphias gladius 351

Xiphiidae 351

Zeidae - 297

Zenopsis ocellata 297

Zeus faber 299

Zoarces anguillaris 510

viviparus 512, 513

zoarchus, Lycodes 510

Zoarcidae 508

zonata, Seriola 372

zygaena, Sphyrna 45

o

U. S. DEPARTMENT OF COMMERCE

Daniel C. Roper, Secretary
BUREAU OF FISHERIES

Frank T. Bell, Commissioner

SUPPLEMENTAL NOTES ON
FISHES OF THE GULF OF MAINE

By HENRY B. BIGELOW and WILLIAM C. SCHROEDER

From BULLETIN OF THE BUREAU OF FISHERIES
Volume XLVIII

Bulletin No. 20

UNITED STATES

GOVERNMENT PRINTING OFFICE

WASHINGTON : 1936

For sale by the Superintendent of Documents. Washington. D. C.

Price 10 cents

320

BULLETIN OF THE BUREAU OF FISHERIES

Page

Saurel, Trachurus trachurus (Linnaeus) 331

Big-eyed scad, Trachurops crumenoplhalma

(Bloch) 331

Hardtail, Caranx hippos (Linnaeus) 331

Hardtail, Caranx crysos (Mitchill) 331

Lookdown, Selene vomer (Linnaeus) 332

Leather jacket, Oligopliles saurus (Bloch

and Schneider) 332

Bluefish, Pomatomus saltatrix (Linnaeus)- 332
Common dolphin, Coryphaena hippurus

Linnaeus 332

Opah, Lampris regius (Bonnaterre) 332

Johnson's sea bream, Taractes princeps

Johnson 332

Butterfish, Poronotus triacanthus (Peck) _ . 332
Harvestfish, Peprilus alepidotus (Lin-
naeus) 333

Striped bass, Roccus lineatus (Bloch) 333

Sea bass, Cenlropristes striatus (Linnaeus) _ 333

Triggerfish, Batistes carotinensis Gmelin__ 333

Filefish, Monacanthus hispidus (Linnaeus) 334

Filefish, Monacanthus ciliatus (Mitchill). 334

Unicornfish, Aluiera scripta (Osbeck) 334

Puffer, Spheroides maculalus (Bloch and

Schneider) 334

Rosefish, Sebastes marinus (Linnaeus) 334

Black-bellied rosefish, Helicolenus dactyl-

oplerus (De la Roche) 334

Hook-eared sculpin, Artediellus uncinatus

(Reinhardt) 335

Mailed sculpin, Triglops ommalistius Gil-
bert 335

Longhorn sculpin, Myoxocephalus octo-

decimspinosus (Mitchill) 335

Deep-sea sculpin, Cottunculus microps

Collett 335

Sea raven, Hemitripterus americanus

(Gmelin) 335

Sea snail, Neoliparis atlanticus Jordan and

Evermann 336

Striped sea snail, Liparis liparis (Lin-
naeus) 336

Red- winged sea robin, Prionotus strigatus

(Cuvier) 336

Remora, Remora remora (Linnaeus) 336

Rock eel, Photis gunnellus (Linnaeus) 336

Snake blenny, Lumpenus lampetraeformis

(Walbaum) 336

Page

Shanny, Leptoclinus 7>iaculatus (Fries) 337

Arctic shanny, Stichaeus punctalus (Fabri-

cius) 337

Radiated shanny, Ulvaria subbifurcata

(Storer) 337

Wrymouth, Cryplacanlhodes maculatus

Storer 337

Spotted wolfRsh, Anarrhichas minor Olaf-

sen 337

Eelpout, Zoarces anguillaris (Peck) 337

Wolf eel, Lycenchelys verritlii (Goode and

Bean) 338

Silver hake, Merluccius bilinearis (Mitch-
ill) 338

Pollock, Pollachius virens (Linnaeus) 338

Cod, Gadus callarias Linnaeus 33S

Haddock, Melanogrammus aeglefinus (Lin-
naeus) 339

Long-finned hake, Urophycis chesteri

(Goode and Bean) 339

Spotted hake, Urophycis regius (Wal-
baum) 339

Four-bearded rockling, Enchelyopus cim-

brius (Linnaeus) 339

Cusk, Brosme brosme (Miiller) 339

Common grenadier, Macrourus bairdii

Goode and Bean 340

American plaice, Hippoglossoides plales-

soides (Fabricius) 340

Four-spotted flounder, Paralichthys oblon-

gus (Mitchill) 340

Rusty dab, Limanda ferruginea (Storer)-. 340

Winter flounder, Pseudopleuronectes ameri-
canus (Walbaum) 340

Georges Bank flounder, Pseudopleuronectes

dignabilis Kendall 341

Witch flounder, Glyptocephalus cynoglossus

(Linnaeus) 34 1

Gulf Stream flounder, Citharichthys arcti-
frons Goode 341

American goosefish, Lophius americanus

Cuvier and Valenciennes 341

Sargassum fish, Histrio histrio (Linnaeus). 341

Deep-sea angler, Mancalias uranoscopus

(Murray) 342

Bibliography 342

Introduction

Since the publication by the Bureau of Fisheries of "Fishes of the Gulf of Maine"
(Document No. 965, U. S. Bureau of Fisheries, Bigelow and Welsh, 1925) enough
new information of general interest has come to hand regarding abundance, dis-
tribution, migrations, breeding habits, and food habits to warrant the issuance of
a supplement to that publication. Many of these data have been obtained during
the investigations carried on by the Bureau; part have been collected from corre-
spondence, while part have beeD gleaned from published material. Brief notes

FISHES OF THE GULF OF MAINE 321

and records of distribution have been taken from the Bulletin of the Boston Society
of Natural History (see Firth (1931), Kendall (1931), MacCoy (1929, 1931a, 1931b,
1933), Schroeder (1931)); from Reports of the Newfoundland Fishery Research
Commission (1932-1933); and from the Proceedings and Transactions of the Nova
Scotian Institute of Science (Leim 1930). For the distribution of certain New
England sharks in South African waters, not referred to in this paper, the reader
is referred to Barnard (1925). For allowing us the use of unpublished notes we
wish to thank F. E. Firth, Dr. G. W. Jeffers, Dr. A. H. Leim, Walter H. Rich, and
O. E. Sette.

The nomenclature used in this supplement is as in "Fishes of the Gulf of Maine."

Hagfish, Myxine glutinosa Linnaeus

Recent detailed studies of the sex organs make it certain that the hag is not
functionally hermaphroditic as was formerly supposed, but that in each individual
either the male portion of the common sex organ matures, with the female organ
remaining rudimentary, or vice versa (Conel, 1931). The fact that a 60 cm speci-
men from Georges Bank contained 30 eggs, 20-25 mm long, shows that large females
may produce somewhat more and slightly larger eggs than previously recorded.

Sea lamprey, Petromyzon marinus Linnaeus

The known range of the sea lamprey in the western Atlantic has been extended
northward to the west coast of Greenland (Jensen, 1926).

Smooth dogfish, Mustelus mustelus (Linnaeus)

The genus Mustelus is established for this species by an opinion rendered by
the International Commission on Zoological Nomenclature (Smithsonian Institution,
1926, p. 8).

Smooth dogfish are taken so seldom in winter that capture of three by a trawler
off Bodie Island, N. C, in 34-45 fathoms, February 1931, is of interest.

Great blue shark, Prionace glauca (Linnaeus)

The International Commission on Zoological Nomenclature (Smithsonian In-
stitution, 1925, p. 27) has rejected Valmont's name, Galeus; consequently the correct
generic name of the species is Prionace Linnaeus.

The blue shark has recently been recorded from the southwest part of the
Grand Bank (Rept., Nfld. Fish. Res. Lab., 1935, p. 79). Although formerly con-
sidered a stray in the Gulf of Maine, recent observations have shown the blue shark
to be common there in August and September, with occasional records for July.
While most often seen offshore, a number were observed and several caught by
J. W. Lowes during the summer of 1935 in Massachusetts Bay.

Young ones are seldom seen along our shores, but Robert Goffin reports one only
20 inches long from Menemsha Bight, near Woods Hole, Mass., August 31, 1925;
while F. E. Firth records another, 38 inches long, taken 65 miles southeast of Highland
Light, Cape Cod, on October 23, 1930.

Dusky shark, Carcharhinus obscurus (LeSueur)

The capture of an 11-foot fish on the northeast peak of Georges Bank, August
10, 1931, extends the known range to the offshore banks (Firth, 1931, p. 9).

322 BULLETIN OF THE BUREAU OF FISHERIES

Shovel-head shark, Cestracion tiburo (Linnaeus)

One specimen of this southern species was recorded by Garman (1913, p. 160)
from Massachusetts Bay.

This record was omitted from "Fishes of the Gulf of Maine" (Bigelow and
Welsh, 1925).

Hammerhead shark, Cestracion zygaena (Linnaeus)

Captures of a 12-foot fish, in August 1928, by the swordfishing schooner Doris M.
Hawes, between Browns and Georges Banks, and of a small one in Halifax Harbor,
September 1932 (Vladykov, 1935, p. 8), extend the known range to the northward
and eastward.

Thresher, Alopias vulpinus (Bonnaterre)

The International Commission on Zoological Nomenclature rejects Valmont's
name, Vulpecula marina, consequently the next oldest name, Alopias vulpinus, must
be substituted (Smithsonian Institution, 1925, p. 27).

Mackerel shark, Isurus nasus (Bonnaterre)

The fact that Isurus punctatus (Storer) is identical with I. nasus (Bonnaterre)
has been pointed out by Bigelow and Schroeder (1927).

The range of the mackerel shark in the western side of the Atlantic is now known
to extend as far north as the Grand Bank of Newfoundland (Rept., Nfld. Fish. Res.
Lab., 1935, p. 79).

Sharp-nosed mackerel shark, Isurus tigris (Atwood)

The many recent fishery investigations in the Gulf of Maine have indicated that
this species is much less common there than /. nasus, for whereas many of the latter
have been observed and captured since 1923, only one record of the sharp-nosed
mackerel shark has come to our attention within that time, a fish 8% feet long taken
10 miles northeast of Nantucket Lightship, June 22, 1930, by the schooner Linta
(Firth, 1931, p. 8).

White shark, Carcharodon carcharias (Linnaeus)

To the few existing Gulf of Maine records of this ferocious shark are added that
of a 13-foot fish taken off Portland in a gill net during October 1931 (identified by
Dr. W. C. Kendall) ; one (identified from a tooth) which attacked a fishing boat off
Digby Gut, Bay of Fundy, July 2, 1932 ;' a somewhat doubtful record from off
Halifax, June 27, 1930 ;3 and another, 15 feet long, apparently of this species, caught
off Monomoy Point, Cape Cod, in the fall of 1928.

Spiny dogfish, Squalus acanthias Linnaeus

The spiny dogfish is now known along the American coast as far northward as
the Straits of Belle Isle. It has also been taken off the west coast of Greenland at
Sukkertoppen and in the vicinity of Holsteinborg (Jensen, 1914, p. 7).

1 Harry Piers, Proc, Nova Scotian Institute ol Science, vol. XVIII, pt. 3, p. 198, 1934.
• Ibid., p. 196.

FISHES OF THE GULF OF MAINE 323

The winter home of the spiny dogfish off the American coast has long been a
subject of uncertainty, hence it is of interest to record that the schooner Victor found
them plentiful about 90 miles southeast of Ambrose Channel Lightship on the tile-
fish grounds the middle of January 1928, and that the Albatross II trawled many
specimens in February 1931, between Cape Hatteras, N. C, and Cape Henry, Va.,
in 16 to 70 fathoms of water. It appears probable, therefore, that the continental
slope to the southward of New England is the chief wintering ground of this species
on this side of the Atlantic.

Analysis of the sizes and of the stages of development of embryos in females
taken at various dates and localities along the coast, and of recent captures of new
born dogfish, also adds to our knowledge (previously scanty) of the breeding habits.

Up until 1925 we had no record of new born dogfish within the Gulf of Maine.
This, together with the facts that females containing large embryos had been often
taken there in early autumn, that dogfish depart entirely from the gulf over the
winter, and that new born young had been reported off Long Island in summer
suggested that the area of reproduction of this species is confined to waters west
and south from Cape Cod.

This is not the case, however, for during the past few years, when special watch
has been kept for new born dogfish, we have learned of their presence in considerable
numbers on Nantucket Shoals and at various localities in the Gulf of Maine from
June to August. Evidently, then, the gulf, as well as the waters off southern New
England, is an important nursery. The fact that embryos, sometimes with yolk sac
nearly absorbed, have repeatedly been found in females off New York in autumn
and on the wintering grounds off Virginia and North Carolina in January and Feb-
ruary might suggest that the coastal waters of the Middle Atlantic States also so
serve. As no new born "dogs" have yet been reported to the southward of New
York at any season, this question remains open, however.

If it should prove that young are born in the southern wintering as well as in
the northern summering grounds, the sizes of the embryos, at different localities and
dates, would suggest that some are set free as early as January or February; in other
words, that the season extends from midwinter right through the spring and summer.

Greenland shark, Somniosus microcephalus (Bloch and Schneider)

So seldom is the Greenland shark captured in the Gulf of Maine that it is of
interest to record a large one taken off Portland Lightship the summer of 1926, and
four others, 4 to 5 feet long, taken in the offing of Portland from 1927 to 1933. A
large one was also taken somewhere in the gulf and brought into Gloucester in Jan-
uary 1929, and another, about 15 feet long, caught in an otter trawl on Jeffreys
Ledge 27 miles northeast of Thatchers Island, off Cape Ann, February 16, 1931.

KEY TO SKATES AND RAYS

Experience has shown that existing keys are not adequate for the identification
of Gulf of Maine skates and rays. The following revision is therefore offered :

1. No long dorsal spire on the tail 2

Tail with long dorsal spines (sting rays) 11

2. Two small dorsal fins, but no distinct caudal fin (includes all our common skates) 3

There is a large triangular caudal fin as well as the two dorsals Torpedo

3. Ventral surface with minute rounded tubercles Raja granulata, 3

Ventral surface smooth 4

' Although Raja granulata is not known from the Quit of Maine it can be eipected there as it has been recorded from La Have
Bank aDd from the continental edge off Halifax, Nova Scotia, from 200 fathoms.

324 BULLETIN OF THE BUREAU OF FISHERIES

4. No thorns along mid-zone of disc between eyes and ventrals.

Barn-door skate, Raja slabuliforis
With one or more rows of thorns along mid-dorsal zone of disc behind eyes 5

5. Posterior third of tail without any large thorns Smooth skate, Raja senta

Posterior third of tail with one or more rows of large thorns 6

6. Tail with only one row of large thorns Young Raja scabrata

Tail with three or more rows of thorns 7

7. Mid-row of tail thorns very much larger than any other thorns on tail.

Prickly skate, Raja scabrata
Mid-row of tail thorns absent or if present not much larger than other thorns on tail 8

8. Three rows of thorns on tail Brier skate, Raja eglanteria

Four or more rows of thorns on tail 9

9. Length of fish more than 2}i feet Big skate, Raja diaphanes

Length of fish less than 2H feet 10

10. Teeth in 70 to 104 rows in each jaw; usually an eye-spot present on each pectoral.

Big skate, Raja diaphanes
Teeth in 38 to 60 rows in each jaw; eye-spot rarely present Little skate, Raja erinacea

11. No dorsal fins on tail Sting ray, Dasybatus marinus

Tail with a dorsal fin ir front of spine Cow-nosed ray, Rhinoptera quadriloba

With regard to the relative abundance of different species of skates on the off-
shore fishing banks of the Gulf of Maine, it is of interest that on a trip to Georges
Bank (chiefly the northeastern part) in September 1929, aboard the otter trawler
Kingfisher, 37 hauls 3uelded from 0 to 105 skates per haul (total 495), as follows:
Raja senta, 57; R. scabrata, 325; R. stabulijoris, 42; and R. diaphanes, 71.

little skate, Raja erinacea Mitchill

This skate has been desciibed as lacking thorns along the midline; but small
specimens 2>){ to 9K inches long and one half-grown specimen of 1Z% inches, recently
examined by us, have this row well developed.

Big skate, Raja diaphanes Mitchill

Recent investigations have shown that the range of this skate extends north-
ward not only to the Gulf of St. Lawrence, as long known, but to the Grand Banks
as well, and southward to Virginia.

The big skate rarely has a median row of thorns except in the very young, so it
is of interest to record a female 18 inches long taken near Jeffreys Ledge, November
1, 1927, which bears a row of large spines along the midline, from the shoulder girdle
to the origin of dorsal on the tail.

Young specimens of R. diaphanes and R. erinacea, especially the females, are not
easily separated from each other by a casual glance, hence the number of rows of
teeth which they possess has been an important means of identification. Investi-
gators have given various tooth counts ranging from 80 to 110 rows for diaphanes and
around 50 rows for erinacea. Several specimens in the Museum of Comparative
Zoology, from New England waters, show 70 to 100 rows of teeth on the jaw of
diaphanes and from 46 to about 60 rows in erinacea.

Prickly skate, Raja scabrata Garman

At the time of publication of "Fishes of the Gulf of Maine" the northern boundary
of this skate (widespread in the Gulf of St. Lawrence) was unknown in the open
Atlantic. Since then it has been found plentifully on the Grand Banks and reported
from the east and north coasts of Newfoundland.

FISHES OF THE GULF OF MAINE 325

Examination of a large number of prickly skates, ranging in size from young,
recently hatched, to the largest recorded, allows us to add the following to previous
descriptions:

The pavementlike teeth are in 41 or 42 rows in the upper jaw, 40 to 44 rows in
lower jaw (4 specimens) and in the male, at least, there are rather sharp cusps on
those teeth situated toward the angles of the jaw. The number of large curved
thorns along the midline of the tail and body was as follows on 23 specimens: 12 (2),
13 (10), 14 (8), 15 (3), with no correlation between the number of thorns and the
size of the specimen. The brownish back is usually marked with small white spots.
The young are more spotted than the adults and have six or seven dark cross bars
on the upper surface of the tail.

This skate grows to 3 feet in length, or slightly larger; a 32-inch fish is about 23
inches wide. The smallest nearly mature male found was 26 inches long.

Since nothing was known of the breeding habits of the prickly skate, it is worth
recording that a specimen 32 inches long taken on the northern part of Georges
Bank, September 22, 1929, had one egg capsule measuring 3 by 2% inches (exclusive
of tendrils) in each oviduct, and that a male, 35% inches long had nearly ripe milt,
but a number of other large females taken at the same time were barren.

Prickly skates caught on Georges Bank in September 1929, had been feeding on
fish, shrimps, spider crabs, anemones, and worms; this is the first definite information
as to the diet of this skate.

Brier skate, Raja eglanteria Bosc

An unusually large one, 37% inches long, was taken off Woods Hole, Mass., in
August 1932.

Additional to the few Gulf of Maine records of this skate already reported are
two specimens taken on Nantucket Shoals, near Round Shoal Buoy, by the Halcyon,
one in July, the other in September 1924. This is a shoal water species, the deepest
capture made by Albatross II between southern New England and the offing of
Chesapeake Bay being in 38 fathoms.

Smooth skate, Raja senta Garman

The smooth skate, formerly believed rare in the Gulf of Maine, is now known
to be quite generally distributed on our offshore fishing banks, as well as on soft
bottom in the deeper parts of the gulf. We have taken it commonly on Georges
Bank, in South Channel, in the deep water (80-100 fathoms) just off Cashes Ledge,
near Jeffreys Ledge, and off Chatham. The shoalest capture was from 25 fathoms.

The largest specimen obtained was 24 inches long.

Barn-door skate, Raja stabuliforis Garman

Young specimens are seldom reported, hence it is of interest to record one of
7}i inches taken on the western edge of Nantucket Shoals July 14, 1930, in 28 fathoms.
This fish had essentially the same characters as the adult. The range of the barn
door skate is now known to extend northward to the western part of the Grand
Bank of Newfoundland.

Torpedo, Narcacion nobilianus (Bonaparte)

As no torpedoes had been reported to the eastward of Cape Cod since 1896,
the captures of a 52-inch specimen weighing 78 pounds, on the southwest part of

80064—36 2

326 BULLETIN OF THE BUREAU OF FISHERIES

Georges Bank, December 8, 1930, and of another of 39 inches at Provincetown,
July 28, 1931, deserve mention here.

Chimaera, Chimaera affinis Capello

It was formerly believed that the chimaera did not exceed a length of about 3
feet, but a specimen measuring 49 inches in length, VI % pounds in weight dressed,
was taken October 15, 1930, 85 miles south by west of Cape Sable in a depth of
about 400 fathoms (Firth, 1931, p. 9).

Common sturgeon, Acipenser sturio Linnaeus

Although sturgeon have seldom been reported from offshore, the recent captures
of a 268-pound fish in South Channel the end of April 1928, of another of 420
pounds in April 1929, of a 335-pound fish trawled on Browns Bank in April 1936,
and of a 435-pound fish on Georges Bank, latitude 41°00' N., longitude 67°45' W.,
on January 7, 1931, indicate that they are to be occasionally found on our outer
fishing banks.

Short-nosed sturgeon, Acipenser brevirostris LeSueur

A 30-inch specimen, taken at Provincetown about 1907, now in the collection
of the Museum of Comparative Zoology, is the only reliable record for the Gulf of
Maine. This record was omitted from "Fishes of the Gulf of Maine" (Bigelow and
Welsh, 1925).

Eel, Anguilla rostrata (LeSueur)

The known range of the American eel in northern waters has been extended to
the west coast of Greenland (Jensen, 1926, p. 101).

American conger, Conger oceanica (Mitchill)

The American conger, long considered identical with the European, has recently
been shown by Schmidt (1931) to be a distinct species, characterized by having fewer
(140-149) vertebrae than the European (154-163 vertebrae); a relationship paralleling
that between the American and European eels of the genus Anguilla.

The American conger ranges along the continental shelf northward to Cape Cod.
Its southern boundary cannot be stated until the congers of the coasts of North and
South America have been critically compared.

Additional to the few records of larvae already reported from the Gulf of Maine
are those of two specimens (4% inches long) picked up on the beach at Newburyport,
Mass., in November 1929, which were sent to us for identification.

Dr. Johannes Schmidt's discovery 4 of very young larvae in the West Indian
region, but nowhere else, points to this as the chief, if not the only, spawning ground
of the American conger.

Snake eel, Pisodonophis cruentifer Goode and Bean

Goode and Bean's (1896, p. 147) record of this species from Jeffreys Bank — the
only one for the gulf — was omitted from "Fishes of the Gulf of Maine." A number
of specimens have been taken recently between the offings of Nantucket and of Cape
Henry, Va., in depths ranging from 24 to 245 fathoms by the Fish Hawk and the
Albatross II.

1 See Schmidt, 1931, p. 602, for a discussion of this question.

FISHES OF THE GULF OF MAINE 327

Herring, Clupea harengus Linnaeus

The northern limit to the known range of the herring in the Western Atlantic
has been extended to the west coast of Greenland by Jensen (1926, p. 101).

Herring are so seldom taken in any large numbers on the offshore banks that it
is of interest to record a catch of 2,800 pounds in South Channel and 3,000 pounds
on the northern edge of Georges Bank, in October 1931.

Mass destruction of young herring, cast up on the beaches has occurred from
time to time in various harbors in the Gulf of Maine. A recent occurrence of this
sort was reported by Dr. Austin H. Clark, who, in Manchester Harbor on the north
side of Massachusetts Bay, early in August 1925, observed that the mud flats were
white with stranded herring which measured 3 to 5 inches in length. Another such
destruction took place in the same harbor in the summer of 1928.

Alewife, Pomolobus pseudoharengus (Wilson)

So little is known about the habits or migrations of the alewife while at sea
that it is of interest to record the capture by Albatross II of 18 adults, 10 to 11 inches
long, by otter trawl, seventy odd miles off Barnegat, N. J., on March 5, 1931.

Blueback, Pomolobus aestivalis (Mitchill)

The maximum length of this herring is usually given as about 1 foot but we have
seen examples of it ranging up to 15 inches.

The capture of seven adult specimens by Albatross II, on March 5, 1931, about
70 miles off Barnegat, N. J., suggests that, like its relative the sea herring, the
blueback moves out from land, and passes the cold season near the bottom, thus
throwing some light on the probable winter home of the Gulf of Maine stock.

Thread herring, Opisthonema oglinum (LeSueur)

The capture of a single specimen, 7 inches long, off Monomoy Point at the
southern angle of Cape Cod in August 1931, extends the known range of this southern
herring to the Gulf of Maine. Occasionally the thread herring is taken off southern
New England; it was even reported as rather common in Buzzards Bay and Vine-
yard Sound in the summer of 1885. As it is essentially a tropical fish it is not apt,
however, to reach the gulf except as the rarest of strays.

Round herring, Etrumeus sadina (Mitchill)

This herring, recorded by Bigelow and Welsh (1925, p. 91) as Etrumeus teres
DeKay, appears very rarely to stray past Cape Cod. Hence, it is of interest to record
the capture of one specimen in Yarmouth River which empties into Casco Bay, and
one in the bay itself, on September 15, 1924.

Anchovy, Anchoviella mitchilli (Cuvier and Valenciennes)

This species is listed by Bigelow and Welsh (1925, p. 124) as Anchovia mitchilli.
The subgenus Anchoviella Fowler differs from the subgenus Anchovia Jordan and
Evermann chiefly by having mucb fewer gillrakers, the former having about 35 to
50 and the latter 100 or more.

328 BULLETIN OF THE BUREAU OP FISHERIES

Striped anchovy, Anchoviella epsetus (Bonnaterre)

A record from off the Presumpscot River, near Portland, October 8, 1930 (Ken-
dall 1931, p. 11) is the first for the Gulf of Maine. This anchovy is now known
from as far northward as Halifax harbor where a number were seined September
29, 1931 (Vladykov 1935, p. 3).

Argentine, Argentina silus Ascanius

Until recently the argentine was considered rare in our waters, for only odd
examples had been brought in from widely scattered localities. The development of
otter trawling proved that argentines are in reality fairly common around the
edge of Georges Bank and off Cape Cod in deep water. Thus, along the northern
and northwestern slopes of the bank and to the eastward of Cape Cod, in depths of
80 to 100 fathoms, it is not unusual for a haul of the trawl to bring in from one to a
dozen, and as much as 15,000 pounds has been reported by one boat during a week's
fishing (Firth 1931, p. 11). It also occurs in the deep central basin of the gulf, for
tae Albatross II has recently (July 1931) trawled a specimen in 115 fathoms off
Mount Desert Rock.

Pearlsides, Maurolicus pennanti (Walbaum)

Additional Gulf of Maine records of this species include 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, from the stomachs of two pollock, caught in 20 fathoms, 7 miles southeast
of Bakers Island, Mount Desert, Maine, July 24, 1930.

Viperfish, Chauliodus sloanei Bloch and Schneider

A specimen found in the stomach of a swordfish caught in the gully between
Browns and Georges Banks in 1931 is the second to be definitely recorded from
within the Gulf of Maine.

lancetflsh, Alepisaurus ferox Lowe

A record of a 5}£-foot specimen of this rare fish caught alive in the surf on Block
Island, R. I., March 12, 1928, is of especial interest even though outside the limits
of the Gulf of Maine. An excellent photograph, sent in by Mrs. Elizabeth Dickens,
shows the upper lobe of the caudal prolonged as a long filament, which most of the
specimens so far seen have lost. This specimen had been feeding on small dogfish.

Needlefish, Scomberesox saurus (Walbaum)

A specimen gaffed at the surface from the Albatross II on northern Georges
Bank, September 20, 1928, is the only definite offshore record for the Gulf of Maine
although the needlefish has been taken in various localities there alongshore.

Trumpetfish, Fistularia tabacaria Linnaeus

Recent reports of the trumpetfish at Port Mouton, Nova Scotia, and on the
south coast of Newfoundland, show that this tropical species may stray much farther
north than previously supposed.

FISHES OF THE GULF OF MAINE 329

Pelagic pipefish, Syngnathus pelagicus Linnaeus

A single specimen 3% inches long, taken on Georges Bank (lat. 42°09' N., long.
66°41' W.) September 20, 1927, by the Albatross II, is 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 tbat was examined.

Common pipefish, Syngnathus juscus Storer

Pipefish are rarely taken on bottom far from the immediate shore waters, hence
it is of interest to report the capture of four specimens 4% to 6 inches long at a depth
of 19 fathoms 10 miles south of No Mans Land, February 5, 1930.

Northern barracuda, Sphyraena borealis DeKay

A specimen about 2 inches long found alive in the surf at Nauset Beach, Cape
Cod, September 26, 1930, by Dr. Edward P. Kichardson, is the only record thus far
reported for the Gulf of Maine. Young fry, a few inches long, are taken, however,
from time to time in the region of Vineyard Sound and Buzzards Bay on the southern
coast of New England, from July to December.

Mackerel, Scomber scombrus Linnaeus

The body length of the mackerel is erroneously given as about three and one-half
times the depth by Bigelow and Welsh (1925, p. 188); actually, it is four to five and
one-half times the depth.

A small mackerel taken at Cape Lookout, N. C, in February 1925 (Coles, 1926,
p. 105), extends the known range southward beyond Cape Hatteras.

Recent captures of a mackerel weighing 7}i pounds,5 and of another of 7% pounds,
26 inches long, both of which we, ourselves, examined, shows that occasional giants
occur, for the weight seldom exceeds 4 pounds or the length 22 inches.

Tuna, Thunnus thynnus (Linnaeus)

Larger catches of tuna have been made within the Gulf of Maine in recent years
partly, at least, because of an increased market demand for the fish. Thus, compared
with the 69,868 pounds recorded for Massachusetts and Maine in 1919, the catch of
1934 amounted to 356,904 pounds, of which 254,076 pounds came from Cape Cod.
The Nova Scotian shore of the Gulf of Maine yielded about 24,000 pounds in 1924
and 10,000 pounds in 1929. At present the annual catch for the gulf is probably
between 300,000 and 400,000 pounds. Assuming an average weight of 300 pounds
(probably too little, for the average weight of about 90 tuna caught off the coast of
Maine in 1926 was about 540 pounds), this would represent a thousand or more fish.
Off the outer coast of Nova Scotia, where tuna have been taken in larger numbers
than within the Gulf of Maine, the annual catches from 1917 to 1933 have fluctuated
between 152,000 and 1,550,000 pounds.

The heaviest New England fish on record, taken off Rhode Island about 1913,
weighed 1,225 pounds, while four or five fish have been brought into Boston that
weighed approximately 1,200 pounds. Another fish weighing 1,300 pounds was
shipped in 1924 from Nova Scotia to Boston (Sella, 1931, p. 61).

' Atlantic Fisherman, August 1925.

330 BULLETIN OF THE BUREAU OF FISHERIES

Small and moderate sized tuna (below 100 pounds) are comparatively rare in
the Gulf of Maine. However, schools composed of individuals estimated to weigh
not more than 40 to 70 pounds were observed around Boston lightship July 13 and
14, 1935.6 None below 20 pounds has been recorded within the gulf, but off southern
New England, especially near Block Island, small tuna are sometimes caught, there
being an unusual run of them (8 to 12 pounds) in 1928. Thus it is probable either
that the lower temperatures of the Gulf of Maine are a barrier to the smaller-sized
tuna, or that they find less favorable feeding grounds there than do the larger sizes.

Off the New England coast the first schools are sighted late in June or early July
to the southward of Block Island, over depths of about 85 fathoms, and a few days
later they appear inshore. At first the fish are hungry, and there is some reason to
believe that their summer migrations follow their breeding period. An example of
their seasonal abundance in the shore waters of the coast of Maine may be had from
the catches made in the vicinity of Casco Bay in 1926, where about 70 fish were taken
in July, 17 in August, 3 in September, and 1 on October 4.

Common bonito, Sarda sarda (Bloch)

Two fish were reported from the mouth of Kennebec River in July and one in
September 1930, and one from southern Nova Scotia (Vladykov, 1935, p. 7) in the
latter month.

In looking through the records of the catches made by a certain set of pound nets
at Provincetown over a period of 10 years, we find the earliest catch for that locality
was in July (1915), and the latest on October 4 (1919).

Spearfish, marlin, Makaira albida (Poey) 7

No spearfish were reported in the Gulf of Maine from about 1880 until 1925.
Since then, however, seven specimens have been brought in, all in summer, one of
them from off Portland, the others from Georges, Browns, and Sable Island Banks,
the last being the most northerly record for the species in the western North Atlantic.
These specimens ranged from 5 feet to nearly 16 feet in length and from 21 to about
700 pounds in weight.

Additional descriptive data based on two New England specimens examined by
us are as follows: The first dorsal fin of one specimen has 47 stiff rays, the other fish
having 48. This fin is separated from the second dorsal by a space equal to the
length of the latter in the one fish, by a shorter space in the other. The first anal fin
(2 spines and about 12 or 13 rays), situated below the rear part of the first dorsal, is
triangular, its first rays forming a sharp angle.

Swordfish, Xiphias gladius Linnaeus

The largest swordfish definitely recorded from the Gulf of Maine was one, caught
in the summer of 1921 by Capt. Irving King and landed at the Boston Fish Pier,
that weighed 915 pounds dressed — hence, upward of 1,000 pounds alive (Fishing
Gazette, September 1921, p. 13). The specimen was not measured, but the sword
being more than 5 feet, the total length of the fish must have approximated 15 feet.

• Data furnished by J. W. Lowes.

' Recorded by Bigelow and Welsh (1925, p. 227) as Tetraplurus imperalor (Bloch and Schneider).

FISHES OF THE GULF OF MAINE 331

In 1931, another large fish was caught, 644 pounds in weight, dressed, 13 feet in
length, with a sword measuring 3 feet 8 inches.

Young swordfish are so rarely reported off the New England coast that it is of
interest to record the capture of a 2-foot fish, weighing 7% pounds, taken by the
Dacia on a trawl line September 2, 1931, on Georges Bank.

Pilotfish, Naucrates ductor (Linnaeus)

Up to 1925 only three definite records for the Gulf of Maine had come to hand.
Since then we have learned of the capture of six more pilotfish, off Portland, in Pro-
vincetown Harbor, to the southeast of Cape Cod, and on the northern edge of Georges
Bank, during the summer and fall months in the years 1921, 1924, 1929, 1931, and
1933. Vladykov (1935, p. 6) reports two specimens from Sable Island Bank and
one from Sambro, off Nova Scotia, in the period 1932-34.

Rudderfish, Seriola zonata (Mitchill)

The known range of the rudderfish has been extended northward to Halifax,
Nova Scotia (Leim, 1930, p. xlvi, as S. dumerili).

One fish was caught on a smelt hook off a Portland wharf in September 1921; a
5K-inch fish was taken off Boston in September 1929; another, 17K inches long, from
South Channel the same month; and a 6-inch specimen on Nantucket Shoals in
August 1930.

Mackerel scad, Decapterus rnacarellus (Cuvier and Valenciennes)

One specimen, 7 inches long, was taken in a trap at Richmond Island, off Cape
Elizabeth, in September 1931, this being only the second recorded for the Gulf of
Maine.

Saurel, Trachurus trachurus (Linnaeus)

One specimen of this fish, rare to the northward of Woods Hole, Mass., was
taken in Casco Bay on August 12 and another near Castine Bay, Maine, on October
15, 1930 (Kendall, 1931, p. 11).

Big-eyed scad, Trachurops crumenopthalma (Bloch)

Two specimens, recently taken off Cape Cod, one at Provincetown, the other
about 8 miles off the beach at Chatham, are the only positive records of this species
for the Gulf of Maine. As it is caught from time to time, however, in the summer and
fall as far northward as Woods Hole, it may be expected to round the cape occasion-
ally. This scad has been recorded from Canso, Nova Scotia by Cornish (1907,
p. 85).

Hardtail, Caranx hippos (Linnaeus)

A hardtail taken off Provincetown in 1933 is the second reported from the Gulf
of Maine. Several specimens about 2 inches long were taken the summer of 1933 in
Musquodoboit Harbor, Nova Scotia (Vladykov, 1935, p. 4).

Hardtail, Caranx crysos (Mitchill)
One fish was taken off Chatham in 1933.

332 BULLETIN OF THE BUREAU OF FISHERIES

Lookdown, Selene vomer (Linnaeus)

During the autumn of 1933 many small lookdowns were reported from traps at
the mouth of Casco Bay, one also from Beverly Farms, and another from North
Truro, an unusual incursion, for only three specimens had previously been recorded
in the Gulf of Maine. Jones (1882 p. 89) and Honeyman (1886 p. 328) record this
species (young) as occasional in the shore waters of Nova Scotia, presumably along
the east coast.

Leatherjacket, Oligoplites saurus (Bloch and Schneider)

A specimen taken in a trap off the outer beach at Chatham is the only record
for the Gulf of Maine.

Bluefish, Pomatomus saltatriz (Linnaeus)

For many years no bluefish had been reported north of Cape Ann, until 1925,
when one was caught off Halifax, Nova Scotia. This seems to have presaged a tempo-
rary extension of range, for numbers of them visited the inner coasts of the gulf
northward to Casco Bay in the summer of 1927, while in 1930 the bluefish was again
reported at Halifax (two specimens) and at Port Mouton, Nova Scotia (one specimen,
Leim, 1930, p. xlvi).

Common dolphin, Coryphatna hippurus Linnaeus

A dolphin about 3% feet long taken 60 miles south-southwest of Cape Sable, in
the deep gully between Browns and Georges Banks by the trawler Natalie Hammond,
August 15, 1930, is the first Gulf of Maine record. The specimen is now in the
collection of tbe Boston Society of Natural History.

Opah, Lampris regius (Bonnaterre) 8

A specimen about 3 feet long was taken in July 1925, on Western Bank, southwest
of Sable Island, by the schooner Falmouth (Radcliffe, 1926), while another of about
the same size stranded on the beach at Hyannis, Mass., on September 17, 1928.

Johnson's sea bream, Taractes princeps Johnson

A fish taken on Browns Bank, off Cape Sable in January 1928 is the first record
of this species for the western Atlantic. This bream previously was known only
from Madeira, in the eastern Atlantic. For a detailed account and comparison with
allied species see Bigelow and Schroeder (1929).

Butterfish, Poronotus triacanthus (Peck)

Recent records show that the northward range of this species extends to the east
coast of Newfoundland, as well as to Nova Scotia as has long been known.

It now seems well established that the butterfish actually withdraw from the
gulf when they disappear in the autumn, as they do from the immediate shore waters
farther south, and from inland waters such as Chesapeake Bay. Until very recently
the winter home of the butterfish was unknown; but as they are now often taken in
the winter otter trawl fishery recently established off the coast between Chesapeake

8 This species was given as Lampris luna (Gmelin) by Bigelow and Welsh (1925, p. 242)i

FISHES OF THE GULF OF MAINE 333

Bay and Cape Hatteras, it appears that they move out to sea to winter on the outer
part of the continental shelf as do several other common Gulf of Maine fishes.

The illustrations of larvae 2.1 and 3.4 mm long credited by Kuntz and Radcliffe
(1918) to the butterfish and reproduced by Bigelow and Welsh (1925, fig. 116, c and d)
have since been proved to belong to one of the hakes.

Harvestfish, Peprilus alepidotus (Linnaeus)

Five or six specimens were reported caught in floating traps at Richmond Island,
off Cape Elizabeth, Maine, in July 1929, while another was taken at the mouth of
the Damariscotta River, Maine, in August 1933, the most northerly record for
the species.

Striped bass, Roccus lineatus (Bloch)

The striped bass considerably increased in abundance along both shores of Cape
Cod between 1928 and 1932, then decreased again as illustrated by the following
catches reported for Barnstable County, Mass.: 1928, 8,060 pounds; 1929, 18,665
pounds; 1930, 27,385 pounds; 1931, 33,600 pounds; 1932, 30,926 pounds; 1933,4,500
pounds. Anglers as well as commercial fishermen have also caught some numbers
along the Eastham-Chatham Beaches and marshes during the past few j^ears, while
a 44% pound bass was caught near Brant Rock on the southern shore of Massachu-
setts Bay, in November 1930. A small stock seems also to have built up in the brack-
ish tributaries of Plum Island Sound north of Cape Ann, for some were taken in
Parker River by anglers during the few years previous to 1930, while in that year
(when fishing restrictions were relaxed) 8,700 pounds were reported thence, though
smaller numbers since then. But this increase did not extend northward beyond
Massachusetts waters, for the commercial reports from the States of New Hampshire
and Maine did not mention bass at all in 1924, or in 192S-33.9

Striped bass so rarely stray away from the immediate shoreline that it is of
interest to mention the capture of a 6-pound fish in a gill net on Cod Ledge, 3 or 4
miles off Cape Elizabeth, Maine, October 15, 1931.

Sea bass, Centropristes striatus (Linnaeus)

Sea bass are seldom taken within the Gulf of Maine, and even on the southern
New England coast are rarely caught later than early November, hence the reported
capture of a 5-pound fish in December 1930, 5 imles east of Pollock Rip Lightship,
in 24 fathoms, is noteworthy.

Trigger fish, Balistes carolinensis Gmelin

Previous to 1925, only one specimen of the trigger fish had been reported from
the Gulf of Maine. Actually, this species must drift over the offshore rim of the
gulf more often than the paucity of early records would suggest, for a specimen was
recorded from Casco Bay in August 1931 ; another was taken in 1932 near Plymouth;
a third, 15 inches long, was gaffed at the surface, on the southeast part of Georges
Bank, from the fishing vessel Huntington Sanford, in July 1929; and two small fry,
2 to 3 inches in length, were picked up on the northeast part of the bank in mid-
September 1927, by the Albatross II. The fact that these last were taken with gulf
weed (Sargassum) suggests that triggerfisli are most apt to appear on the banks with
the latter.

! No statistics are available for 1925-27.

334 BULLETIN OF THE BUREAU OF FISHERIES

Filefish, Monacanthus hispidus (Linnaeus)

The filefish appears in the inner parts of the gulf only as a stray from warmer
seas, recent records being that of a fish taken off Seguin, September 12, 1929, one off
Portland lightship, July 17, 1931, and a 6-inch fish at Provincetown, November 6,
1929. On the offshore banks, however, it is to be expected more frequently (which
accords with its southern origin) for the Albatross II gathered 181 small fry 1 to 2
inches long, on the northeastern part of Georges Bank among floating gulf weed
(Sargassum) in September 1927; while a larger one was picked up to the southeast of
Cape Cod in that same month of 1930.

Filefish, Monacanthus ciliatus (Mitchill)

A 7-inch fish taken in a Provincetown trap in November 1929 is the second (and
only recent) record of this species within the Gulf of Maine (Firth, 1931, p. 13). A
straggler has been reported, however, from Newfoundland — far to the north of its
previously known range.

Unicornfish, Alutera scripta (Osbeck)

Two specimens of this fish, 5 and 5}i inches long, respectively, caught on the west-
ern edge of Georges Bank, constitute the first Gulf of Maine record (Mac Coy, 1931a,
p. 16).

Puffer, Spheroidts maculatus (Bloch and Schneider)

A specimen taken off Long Island, Portland Harbor, on July 24, 1933, is the
first to be recorded from the northern boundary (Casco Bay) of this species since
1896.

Rosefish, Sebastes marinus (Linnaeus)

It is now known that rosefish may be born in the Gulf of Maine as early as the
end of April, for in 1930 we saw gravid females during the last half of that month.
In July 1931 the Albatross II trawled many gravid females, 10 to 13% inches long,
in the central basin of the gulf; one of these, 13 inches long, contained approximately
20,500 young 6 to 7 mm long, ready to be spawned.

The fact that we obtained many young fish 2}i to 5% inches in length, off the coast
of Maine from April to August, suggests that this is the approximate size attained
during their first year of life.

Kecent catches of 75-625 rosefish per haul in a trawl by the Atlantis in 70-130
fathoms in the western and northeastern parts of the gulf are evidence of the abun-
dance of this species over the soft bottoms of the basins, as well as in other parts
of the gulf. The commercial importance of this species has greatly increased of
late, the reported landings having risen from 1,288,000 pounds in 1934 to 14,100,000
pounds in 1935.

Black-bellied rosefish, Ilelicolenus dactylopterus (De la Roche)

A fish 13 inches long, trawled on the eastern edge of Georges Bank in 150 fathoms,
October 6, 1929 (Firth, 1931, p. 13), is the first record for this species within the Gulf
of Maine. In addition to previous records from outside the gulf, a number of small
fish (1% to 3y2 inches) were trawled off southern New England in 80 to 118 fathoms
during 1930.

FISHES OF THE GULF OF MAINE 335

Hook-eared sculpin, Artediellus uncinatus (Reinhardt) l0

This sculpin is now known to be generally distributed in the Gidf of Maine in
depths greater than 20 to 30 fathoms. Thus, in addition to the Massachusetts Bay
records of many years ago, we have recently taken it repeatedly near Mount Desert,
off Cape Elizabeth, near Jeffrey's Ledge, around Cashes Ledge, along the northern
slopes of Georges Bank, in the southeastern part of the basin of the gulf, and at the
entrance to the deep gully between Georges and Browns Banks, in depths ranging
from 20 to 150 fathoms. Individual hauls have yielded up to six or eight specimens,
both on hard and on soft bottom.

After examining specimens from New England waters and comparing published
drawings of European fish, we can find no major differences between the hook-eared
sculpins of the eastern and western Atlantic.11

Mailed sculpin, Triglops ommatistius Gilbert

This sculpin is not as rare in the Gulf of Maine as was formerly supposed, for
during the past few years we have trawled specimens near Mount Desert, in Mas-
sachusetts Bay, off Cape Ann, off Cape Cod, and around the northern slope of Georges
Bank, in depths of 20 to 140 fathoms, in various months from spring to autumn.
The most southerly locality was about 10 miles east of Chatham.

Longhorn sculpin, Myoxocephalus octodecimspinosus (Mitchill)

Numerous young specimens 1% to 2 inches long taken in September, and 3 to 3%
inches in February, suggest that the longhorn sculpin is about 2 to 3 inches long at 1
year of age, spawning as it does in late fall.

Deep-sea sculpin, Cottunculus microps Collett

A specimen, about 2 inches long, trawled by the Albatross II on the northern
slope of Georges Bank, in a depth of 120 fathoms, on July 24, 1931, is the third
record for the Gulf of Maine proper.

Sea raven, Hemitripterus americanus (Gmelin)

The fact that fish of both sexes with gonads only partially developed have recently
been found on Nantucket Shoals late in June, added to previous captures of ripe
females off southern New England in November and December shows this to be a
late fall and early winter spawner. The sea raven is a prolific fish, for a female 20
inches long that we caught off Boothbay Harbor, Maine, in April 1925, contained
about 10,000 eggs. The fact that these were definitely of two sizes, the smaller
averaging 1 .5 mm in diameter, the larger about 3 mm, raises the interesting question
whether individual sea ravens may spawn more than once during the year.

The sizes of the few young sea ravens that have been taken in the Gulf of Maine
suggest that they reach a length of 2 to 4 inches by the middle of the first summer,
when 6 to 8 months old; and about 6 inches by the following April, at an age of 1%
years.

10 Given as Artediellus atlanlkus Jordan and Evermann by Bigelow and Welsh (1925, p. 314).

" Jordan, Evermann, and Clark (193d, p. 377) in the Check List of Fishes placed Artediellus attavticus Jordan and Evermann
in the synonymy of A. uncinatux Reinhard*t

336 BULLETIN OF THE BUREAU OF FISHERIES

Sea snail, Neoliparis atlanticus Jordan and Evermann

The sea snail, previously unknown offshore, has recently been taken on Georges
and on Browns Banks. Its range has recently been found to extend as far southward
as the offing of Atlantic City, N. J. (Lat. 39°20'N.). Most of the specimens were
found living in scallop shells (Pecten magellanicus) , as is so often the case.

Striped sea snail, Liparis liparis (Linnaeus)

This sea snail was formerly known as far southward as New York but the Alba-
tross II has taken it off Delaware Bay and the Grampus off Assateague, Va. (Welsh,
1915, p. 2).

Red-winged sea robin, Prionotus strigatus (Cuvier)

A specimen was taken off Monhegan, Maine, in 40 fathoms, in an otter trawl
November 19, 1933. This is the most northerly record for this straggler in the
Gulf of Maine.

Remora, Remora remora (Linnaeus)

Recent Gulf of Maine records of this species include one found on the bottom of a
lobster trap in Portland Harbor in 1931, probably brought in by a schooner from the
West Indies ; one found sucking to the gills of a blue shark (Prionace glauca) that was
caught on the northeast edge of Georges Bank, August 1, 1931 ; one in Cape Cod Bay
in September 1934, and one off Provincetown in August 1935, taken by C. W. Lowes
on blue sharks; also two specimens, 6 and 17 inches long, respectively, taken on
August 3, 1932, 220 miles east-southeast of Cape Ann. Previously it had been
recorded only once from the Gulf of Maine.

Rock eel, Pholis gunnellus (Linnaeus)

Recent records show that the rock eel occurs in considerable numbers on the offshore
banks in the Gulf of Maine down to at least 40 fathoms and occasionally even to 100
fathoms (Schroeder, 1933, p. 5) as well as inshore. So many have been found in the
stomachs of cod and pollock caught on Nantucket Shoals, Georges Bank, Browns
Bank, Cashes Ledge, etc., that it must be an important food of these two species.

The range of the rock eel recently has been found to extend soutb to the latitude
of Delaware Bay, where in February 1930 Albatross II trawled two specimens in 23
and 38 fathoms, respectively.

Snake blenny, Lumpenus lampetraeformis (Walbaum)

Recent captures, by Albatross II, of adult snake blennies (one specimen each)
off Mount Desert, off Boone Island, and on Stellwagen Bank, in depths ranging from
28 to 88 fathoms, added to earlier records from Massachusetts Bay and from the Bay
of Fundy region, show that this species is generally distributed over the gulf, as records
of its larvae had suggested. So slender and active is tbis fish that it can easily escape
through the meshes of any of the nets used by commercial fishermen, hence it is seldom
reported.

Color notes taken from a 12-inch specimen are as follows: The body had brown
markings on a whitish ground, the head being pale brown. The dorsal fin was
marked obliquely with 18 pale bars, the caudal transversely with 8. The anal rays
were pale brown against a colorless membrane, the ventrals white, while the pectorals
were tinged with brown.

FISHES OF THE GULP OF MAINE 337

One of 19 inches caught on the eastern slope of Stellwagen Bank in 42 fathoms in
July 1931 is the largest on record.

Shanny, Leptoclinus maculatus (Fries)

One specimen of this stray from the north was trawled on the northeast part of
Georges Bank in August 1926 and four (4 to 4% inches long) were taken off Chatham,
Cape Cod, in 28 fathoms, May 1, 1930, by the Albatross II. This is the most southerly
record for the species.

Arctic shanny, Stichaeus punctatus (Fabricius)

A specimen 4}i inches long of this arctic species, taken one-half mile off Little Duck
Island near Mount Desert, Maine, from the stomach of a cod, on April 30, 1930, is the
first record for the Gulf of Maine; the only record indeed to the southward of New-
foundland. This specimen was in such good condition that it unquestionably had
been living in the immediate vicinity.

Radiated shanny, Ulvaria subbifurcata (Storer)

This shanny was previously known to be rather common in the northeastern part of
the gulf, and enough have now been found in the stomachs of cod caught on Cashes
Ledge, Georges Bank, Nantucket Shoals, and other offshore grounds to show that it is
widespread in other parts of the gulf as well, on hard bottom. The deepest capture
was in 45 fathoms.

Wrymouth, Cryptacanthodes maculatus Storer

Recent captures of two specimens in the central basin of the Gulf of Maine, July
1931, in 88-95 fathoms, of three in August 1936, in 72-100 fathoms, and of another
on the continental slope between 245 and 325 fathoms, shows that this species is not
as closely restricted to the vicinity of the coast as previously supposed and that it
reaches considerably greater depths.

The locality of the capture (taken by Atlantis) last mentioned (lat. 39°31' N;
long. 72°16' W.) also extends the known range somewhat farther south.

Spotted wolffish, Anarrhichas minor Olafsen

This Arctic species is seldom taken within the Gulf of Maine, hence the capture
of a small specimen, weighing 3}i pounds, on a trawl, off Portand Lightship on April
23, 1927, is worthy of mention. On the Scotian banks, however, it is not so uncom-
mon, for we have records of 2, 37 and 54 inches long, respectively, caught on Sable
Island Bank in January 1934 and 5 more in March of that year. Usually about 5 to
10 from this general region are landed each year at the Boston Fish Pier.

Eelpout, Zoarces anguillaris (Peck)

Many small specimens from l.S inches long upward, have recently been collected
along our coast between Maine and New Jersey, including (within the gulf) Mount
Desert, Stellwagen Bank, Georges Bank, and the vicinity of Chatham, suggesting
that the eelpout breeds successfully throughout this range. And as all the young
thus far taken have been caught in depths of 20 to 45 fathoms, probably this is the
usual spawning zone. Although eelpouts have seldom been reported deeper than 50

338 BULLETIN OF THE BUREAU OF FISHERIES

fathoms, Albatross //recently (July 1931) trawled a number in the basin of the gulf
as deep as 90 fathoms.

The sizes, in different months, of the young fry show that eelpouts in the Gulf
of Maine grow to a length of about 2 inches in the first 6 months of their lives, and 3
inches in 9 months, agreeing in this respect with the growth-schedule of Bay of Fundy
eelpouts derived by Clemens and Clemens (1921, p. 74) from the annual rings on the
otoliths. Small specimens 5 to 6% inches long taken from February to May are
probably about \)'2 years old.

Young eelpouts, up to 3 or 4 inches long, are checkered along the sides, and
irregularly blotched on the back with light and dark brown, with a small but promi-
nent black spot, which fades out with growth, on the anterior part of the dorsal fin.

Wolf eel, Lycenchelys verrillii (Goode and Bean)

The recorded range of the wolf eel, previously known only off the coasts of New
England and Nova Scotia, has now been extended southward to the offing of New York
(Beebe, 1929, p. IS).

The wolf eel is more common within the Gulf of Maine, in deep water, than was
formerly supposed, for in the autumn and summer of 1928 and 1930 the Albatross II
trawled 61 specimens, 6 to 6% inches long, in the deep basin to the westward of Jeffreys
Ledge, in about 90 fathoms of water. It was also foimd scattered over the central
basin of the gulf, in July 1931, in 95 to 123 fathoms.

Silver hake, Merluccius bilinearis (Mitchill)

The wintering ground of the Gulf of Maine stock of silver hake has been the sub-
ject of so much speculation that the capture by the Albatross II, of many specimens
between the offings of No Mans Land and off Cape Hatteras in depths ranging from
12 to 146 fathoms, in February 1930 at temperatures of 4.2° to 10.6° C. (39.5° to
51° F.), deserves mention. Such wide ranges of temperature indicate that the silver
hake are well distributed on these offshore grounds during the winter.

Young fish are rarely found close to shore within the gulf. Offshore, however, the
Albatross II and Atlantis have trawled large numbers between 2 and about 8 inches
long in widely scattered localities and in depths ranging from 20 to 115 fathoms.

Measurements of young silver hake,12 recently obtained in the Gulf of Maine
indicate that a length of 6-7 inches is attained at about 1 year of age.

Pollock, Pollachius virens (Linnaeus)

Recent tagging experiments verify the earlier view that the pollock which appear
in the cold months of the year off New York and New Jersey are winter migrants from
the region of Nantucket Shoals. In general the pollock in the Gulf of Maine are not
migratory although occasional fish may make long journeys.

Cod, Gadus callarias Linnaeus

Extensive tagging experiments (Schroeder, 1930) have proved that the appear-
ance of cod in winter southward along the coasts of New York and New Jersey in
commercial quantities represents a regular annual mass migration from Nantucket
Shoals followed by a return migration in spring. But only scattering fish join this

" Several hundred specimens.

FISHES OF THE GULF OF MAINE 339

winter migration from the more northerly and easterly parts of the Gulf of Maine.
It has been known that many of these cod spawn on the southern wintering grounds,
but it was not until the spring of 1930 that large numbers of fry were obtained there.
At that time (April) Albatross II trawled hundreds of fry 1% to 2% inches long on
bottom, the most southerly catch being in latitude 36°21' N.13

Haddock, Melanogrammus aeglefinus (Linnaeus)

The haddock, formerly unknown beyond the Straits of Belle Isle, in the western
Atlantic, has now been reported from West Greenland (Jensen and Hansen, 1930, p.
52). From Icelandic waters comes a record of a giant haddock 44 inches long and
weighing about 37 pounds (Thompson, 1929, p. 29).

long-finned hake, Urophycis chesteri (Goode and Bean)

The capture of several specimens on the northern edge of Georges Bank in
September 1929, in 85 to 100 fathoms, and of many to the westward and in the central
basin of the Gulf of Maine the summer of 1931, in 70 to 140 fathoms, suggests that
this species is more plentiful in the gulf than was previously supposed.

This hake is said to be a summer spawner but very little is known concerning its
rate of growth, hence we report captures of 3 fish 57 to 71 mm on April 26, 1931, and
of 16 fish of 74 to 110 mm taken late in July, suggesting that a length of 4 or 5 inches
is reached at 1 year of age.

Spotted hake, Urophycis regius (Walbaum)

The scarcity of this hake within the Gulf of Maine is emphasized by the fact that
not a single one was captured there in the numerous hauls made recently by the
Albatross II. To the southward, however, many were trawled between Cape Hat-
teras and the offing of Delaware Bay in 5 to 45 fathoms (chiefly in less than 20 fath-
oms) from February to May 1930 and 1931.

Although the spotted hake reaches a length of at least 16 inches, large fish are
relatively rare. The longest of about 600 specimens taken on 14 stations by the
Albatross II was only 130 mm (5% inches). In the largest catch (Apr. 8) the dominant
size was 2 to 2}{ inches.

Four-bearded rockling, Enchelyopus cimbrius (Linnaeus)

The rockling has recently (July 1931) been trawled in the central basin of the
Gulf of Maine where it was expected, but heretofore unrecorded. The fact that one
was taken in latitude 36°56' N., off Cape Charles, Va., on February 10, 1930, in only
12 fathoms, shows that in the most southerly parts of its range, it is not restricted to
deep water, as previously supposed.

Cusk, Brosme brosme (Muller)

A fish 40 inches long and weighing 27 pounds, trawled by Albatross II in the
central part of the Gulf of Maine, in 120 fathoms, is the largest definitely recorded
from the Gulf of Maine.

'« These were taken during the course ol O. E. Sette's mackerel investigations.

340 BULLETIN OF THE BUREAU OF FISHERIES

Common grenadier, Macrourus bairdii Goode and Bean

Recent records show that the grenadier is comparatively common on muddy
bottom in the gulf, at depths greater than about 90 fathoms and that it may occasion-
ally be taken shoaler, for one was reported from the slope of Jeffreys Ledge in about
50 fathoms during March 1934. The capture of a ripe male in late September verifies
the earlier suggestion that the grenadier is an autumn spawner. The largest fish
taken by Albatross //was 16 inches long. This grenadier has now been taken as far
eastward as the Grand Banks of Newfoundland (Nfld. Rpt., 1933 (1934), p. 116).

American plaice, Hippoglossoides platessoides (Fabricius)

Recent trawling by Albatross II and Atlantis proves this species to be generally
distributed even in the deeper parts of the central basin of the gulf, to a depth of at
least 120 fathoms. A specimen 15% inches long caught off Montauk Point, N. Y., in
112 fathoms, February 6, 1930, is the most southerly and westerly record.

As this flounder is a spring spawner it may be assumed that bottom stages 69 to 80
mm long trawled off Cape Cod, May 1, were about 1 year old, and 85 to 118 mm fry
found at several localities in July and August were between l}{ and IK years old,
those of 8-10 inches, 2% to 2% years.

Four-spotted flounder, Paralichthys oblongus (Mitchill)

This flounder, formerly thought rare to the east of Cape Cod, has recently been
found here and there on the southern half of Georges Bank. Previously known only as
far southward as New York, many have been trawled by the Albatross //south to the
Virginia Capes (hat. 36°45' N.).

The fact that captures were made in 23 to 112 fathoms in February (7 stations),
31 to 52 fathoms in March (two stations), 10 to 85 fathoms in April (eight stations),
15 to 35 fathoms in May (four stations), 11 to 47 fathoms in June (five stations), and
41 fathoms in July (one station) indicates that it is present and widely distributed in
this general depth zone the year round.

The capture of ripe specimens as late as mid-July shows that the breeding season
is not limited to spring, as formerly supposed, but extends well into the summer.

Rusty dab, Limanda ferruginea (Storer)

Capture of a specimen, in the offing of Hog Island, Va. (lat. 37° 41' N.) consider-
ably extends the known range to the southward. The captures of young dabs 2 to 4
inches long iD February (17 fish), 2% to 4% inches in April (26 fish), 2% to 5% inches in
May (10 fish), 3 to 5 inches in June (3 fish) and 3 to 6% inches in July (13 fish) yield the
first data as to rate of growth. According to this growth schedule the rusty dab
reaches a length of approximately 5 inches at 1 year of age.

Winter flounder, Pseudopleuronectes americanus (Walbaum)

The recovery, off Chatham and on Nantucket Shoals, of winter flounders
tagged and released at Woods Hole proves that some of them, at least, may wander
for longer distances than previously supposed.

FISHES OP THE GULF OF MAINE 341

Georges Bank flounder, Pseudopleuronectes dignabilis Kendall

Tliis flounder, previously known only from the Georges Bank area, is now re-
ported from the eastern edge of the Scotian banks and the western part of the Grand
Banks of Newfoundland (Nfld. Rept., 1934 (1935), p. 79).

Witch flounder, Glyptocephalus cynoglossus (Linnaeus)

The witch flounder has recently been found to be generally distributed in the
central basin of the Gulf of Maine where the Albatross II and Atlantis trawled it
down to 140 fathoms, in July 1931 and in August 1936, respectively. Goode and
Bean's (1896, p. 433) record of it in latitude 34°39' at a depth of 603 fathoms (omitted
in Bigelow arid Welsh, 1925) shows that it ranges southward to the offing of Cape
Hatteras in deep water. But the most southerly record of it in shoal water is a
specimen taken by Albatross II in 10 fathoms off Virginia (lat. 37°50')-

Many specimens from 3 to 5 inches and from 7 to 8% inches long were taken
from July to September suggesting that the witch reaches a length of about 4 inches
at 1 year and about 8 inches at 2 years of age.

Gulf Stream flounder, Citharichthys arctifrons u Goode

This little flounder was formerly believed to reach a length of only about 4
inches but recently the Albatross II collected many specimens up to 7 inches long.

Recent trawling experience extends knowledge of its distribution by showing
that it may occur as shoal as 12 fathoms, and that it finds its northeastern boundary
off the southeastern slope of Georges Bank and its southwestern boundary off Cape
Hatteras. Usually only a few specimens are taken in any given trawl haul, even
further to the west and south where the species appears to be most common; hence,
a catch of about 100 made by the Albatross II, off Montauk Point, N. Y., in 50
fathoms, is noteworthy.

Apparently, it spawns over a long season, from spring through summer, for we
have found females with well-developed ovaries in February while Goode had ripe
ones in September. Although the Gulf Stream flounder is not large enough and
thus far has been found too scarce to be of commercial value, we can witness that
it is excellent on the table.

American goosefish, Lophius americanus Cuvier and Valenciennes

Recent investigations by Berrill (1929) and by Procter et al. (1928) make it
appear that the American goosefish, given as Lophius piscatorius in "Fishes of the Gulf
of Maine" (Bigelow and Welsh, 1925, p. 524), is specifically distinct from the
European.

Very small goosefish are seldom reported, hence captures of 1 of 10 inches in
February, 1 of 10 inches in April, 2 of 7% and 10 inches, respectively, in May, 3 of
6% to 9 inches in July, and 3 of 4 to 4% inches in August between latitudes 43°21' N.
and 37°36' N. in depths ranging from 35 to 140 fathoms, are of interest.

Sargassum fish, Histrio histrio (Linnaeus)

A single specimen about 4% inches long, picked up in a purse seine near the sur-
face over the west central part of Georges Bank, by the schooner Old Glory on Sep-
tember 15, 1930 (Firth, 1931, p. 14), extends the known range of this fish to the Gulf
of Maine.

'< Parr (1931) has made a revision of the genus Citharichthys of the western Atlantic.

342 BULLETIN OF THE BUREAU OF FISHERIES

Deep-sea angler, Mancalias uranoscopus (Murray)

A 24K-inch specimen of this uncommon fish was trawled on Georges Bank Feb-
ruary 9, 1927, by tbe fishing steamer Ripple; this is the only record of a member of
this family (Ceratiidae) from New England waters (Parr, 1932, p. 12).

BIBLIOGRAPHY

Barnard, K. H. 1925. A monograph of the marine fishes of South Africa. Part I. Ann.,

So. African Museum, vol. XXI, 1925, pp. 1-418. Edinburgh.
Beebe, William. 1929. Deep-sea fish of the Hudson Gorge. Zoologica, vol. XII, no. 1, 1929,

19 pp., 1 fig. New York.
Berrill, N. J. 1929. The validity of Lophius americanus Val. as a species distinct from L.

piscatorius Linn, with notes on the development. Contr., Canad. Biol, and Fish., N. S., vol. IV,

no. 12, 1929, pp. 143-155, 7 figs. Ottawa.
Bigelow, Henry B., and William C. Schroeder. 1927. Notes on northwest Atlantic sharks

and skates. Bull., Museum, Comp. Zool., vol. LXVIII, no. 5, September 1927, pp. 239-251.

Cambridge.
Bigelow, Henry B.,and W. C. Schroeder. 1929. A rare Bramid fish (Taractes princeps John-
son) in the northwestern Atlantic. Bull., Museum, Comp. Zool., vol. LXIX, no. 2, February

1929, pp. 39-50. Cambridge.
Bigelow, Henry B., and William W. Welsh. 1925. Fishes of the Gulf of Maine. Bull., U. S.

Bur. Fish., vol. XL, Part I, 1924 (1925), 567 pp., 278 figs.
Clemens, Wilbert A., and Lucy Smith Clemens. 1921. Contribution to the biology of the

muttonfish, Zoarces anguillaris. Cont., Canad. Biol., 1918-1920 (1921), pp. 69-83, 1 pi.

Ottawa.
Coles, Russel J. 1926. Notes on Cape Lookout (North Carolina) fishes — 1925. Copeia, no.

151, February 1926, pp. 105-106.
Conel, J. LeRoy. 1931. The genital system of the Myxinoidea: A study based on notes and

drawings of these organs in Bdellostoma made by Bashford Dean. The Bashford Dean

Memorial Volume, Archaic Fishes, Article III, Amer. Museum, Nat. Hist., 1931, pp. 64-102,

pi. I-IV. New York.
Cornish, George A. 1907. Notes on the fishes of Canso. Further Contr., Canad. Biol., 1902-05

(1907), pp. 81-90. Ottawa.
Firth, Frank E. 1931. Some marine fishes collected recently in New England waters. Bull.,

Boston Soc. Nat. Hist., no. 61, October 1931, pp. 8-14. Boston.
Fish, Charles J. 1927. Production and distribution of cod eggs in Massachusetts Bay in 1924

and 1925. Bull., U. S. Bur. Fish., vol. XLIII, 1927, Part II, pp. 253-296, 16 figs.
Goode, George Brown, and TarletonH. Bean. 1896. Oceanic ichthyology. Mem., Museum,

Comp. Zool., Harvard College, vol. XXII, 1896, xxxv + 553 pp. Cambridge. Also Smith.

Contr. to Knowl., vol. XXX, 1895 (1896), and Spec. Bull. No. 2, U. S. Nat. Museum, 1895

(1896).
Hildebrand, Samuel F., and William C. Schroeder. 1928. Fishes of Chesapeake Bay. Bull.,

U. S. Bur. Fish., vol. XLIII, Part I, 388 pp., 211 figs.
Honeyman, D. 1886. Nova Scotian ichthyology. Proc. and Trans., Nova Scotian Inst. Nat.

Sci., vol. VI, Part IV, 18S6, pp. 328-330. Halifax.
Jensen, Ad. S. 1914. The selachians of Greenland. Mindeskrift for Jepetus Steenstrup, 1914,

40 pp. Copenhagen.
Jensen, Ad. S. 1926. Investigations of the "Dana" in West Greenland waters. 1925. Rapp.

et Proces-Verb., Cons. Perm. Inter. Explor. Mer, vol. XXXIX, 1926, pp. 85-102. Copen-
hagen.
Jensen, Ad. S. 1928. The fauna of Greenland. In Greenland, published by the Commission

for the direction of the Geological and Geographical Investigations in Greenland, vol. I, 1928,

pp. 319-355. Copenhagen.
Jensen, Ad. S., and Paul M. Hansen. 1930. Unders0gelser over den Gr0nlandske Torsk, 1930,

55 pp. Copenhagen.
Jones, J. Matthew. 1879. List of the fishes of Nova Scotia. Proc. and Trans., Nova Scotian

Inst. Nat. Sci., vol. V. Part I, 1879, pp. 87-97. Halifax.

FISHES OF THE GULF OF MAINE 343

Jordan, David Starr, Barton Warren Evermann, and Howard Walton Clark. 1930.

Check list of the fishes and fishlike vertebrates of North and Middle America north of the

northern boundary of Venezuela and Colombia. Report, U. S. Com. Fish., Part II, 1928

(1930), 670 pp.
Kendall, William C. 1931. Remarks on additions to the marine fauna of the coast of Maine.

Bull., Boston Soc. Nat. Hist., no. 58, Jan. 1931, pp. 9-11. Boston.
Leim, A. H. 1930. Unusual fishes and other forms in Nova Scotian waters. In Proc. and Trans.

Nova Scotian Inst. Nat. Sci., vol. XVII, Part 4, 1930, p. xlvi. Halifax.
MacCot, Clinton V. 1929. The mackerel in New England. Bull., Boston Soc. Nat. Hist., no.

53, Oct. 1929, pp. 3-7. Boston.
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Jan. 1931, pp. 16-18. Boston.
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CHANGES IN NAMES AND CLASSIFICATION FOR BIGELOW AND SCHROEDER
Fishes of the Gulf of Maine, 1953

Petromyzonidae = Petromyzontidae

Carchariidae = Odontaspidae; Carcharias taurus = Odontaspis taurus (Rafinesque)

Isuridae = Lamnidae

Cetorhinidae is frequently considered as a subfamily of Lamnidae

Triakidae is frequently included in Carcharhinidae

Scoliodon terrae-novae =Biizoprionodon terraenovae (Richardson)

Carcharhinus milberti = C. plumbeus (Nardo)

Dalatiidae is now included as a subfamily of Squalidae

Echinorhinidae usually considered as a subfamily of Squalidae

big skate = winter skate

brier skate = clearnose skate

Rhinopteridae = Myliobatidae

Acipenser sturio = Acripenser oxyrhynchus oxyrhynchus Mitchill

Tarpon atlantiaus = Megalops attanticus Valenciennes, family Elopidae

Etrumeus sadina = Etrumeus teres (DeKay), family Clupeidae

Clupea harengus = Clupea harengus harengus, Atlantic herring

Pomolobus mediocris = Alosa mediocrxs (Mitchill) ^

Pomolobus pseudoharengus = Alosa pseudoharengus (Wilson)

Pomolobus aestivalis = Alosa aestivalis (Mitchill)

Anguilla (p. 151) belongs to the Anguillidae; Conger (p. 154) to the Congridae;

Simenchelys (p. 157) to the Simenchelyidae; Synaphobranchus (p. 158) to the

Synaphobranchidae; Omochelys (p. 159) to the Opichthidae; and Nemichthys

(p. 159) to the Nemichthyidae

Conger oceanica = Conger oceanicus (Mitchill)

Omocneuys cruenzifer — Oph.ichzh.us cruentifer (Goode hjiu Bean)

Fundulus and Cyprinodon belong to the Cyprinodontidae

billfishes or silver gars = needlefishes

Tylosurus marinus = Strongylura marina (Walbaum)

needlefishes = sauries

Cypselurus heterurus = C. melanurus (Valenciennes)

Merlucciidae is sometimes included in Gadidae but Antimora and Physiculus

here included in Gadidae belong to the Moridae

Gadus callarias = Gadus morhua (Linnaeus)

Urophycis regius = U . regia (Walbaum)

Orophycis chesteri = Phycis chesteri Goode and Bean

Antimora and Physiculus belong to the Moridae

Macrourus bairdii = Nezumia bairdii (Goode and Bean)

Lampris regius = Lampris guttatus (BrUnnich)

Hippoglossidae is now included in the Pleuronectidae, Paralichthyidae in the

Bothidae, and Achiridae (Trinectes) in the Soleidae. Three genera Paralichthys,

Scophthalmus and Citharichthys , are lefteye flounders, Bothidae; the other

genera are righteye flounders, Pleuronectidae

sand flounder, Lophopsetta maculata = window pane, Scophthalmus aquosus (Mitchill)

Achirus fasciatus = Trinectes maculatus (Bloch and Schneider), Soleidae

Zenopsis ocellata = Z. conchifera (Lowe)

Xenolepidichthys americanus = Daramattus americanus (Nichols and Firth)

Macrorhamphosidae is included in Centriscidae by some authors

Hippocampi dae, seahorses are now included in the Syngnathidae

Hippocampus hudsonius = H. erectus Perry, the lined seahorse

Pneumatophorus colias = Scomber japonicus Houttuyn

striped bonito, Euthynnus pelamis = skipjack, Katsuuonus pelamis (Linnaeus)

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false albacore = little tunny

common bonito = Atlantic bonito

tuna = Atlantic bluefin tuna, Thunnus thynnus thynnus (Linnaeus)

Scomberomorus regalis = cero; Scomberomorus cavalla = king mackerel

Makaira ampla - I4a.ka.ira nigricans Lacepede

Makaira albida = Tetrapturus albidus Poey

Poronotus triacanthus = Peprilus triacanthus (Peck)

Centrolophidae is included in Stromateidae by some authors; Palinurichthys

perciformis = Hyperoglyphe perciformis (Mitchill)

Trachurus trachurus = Trachurus lathami (Nichols)

Trachurops crumenopthalmus = Selar crumenopthalmus (Bloch)

Vomer setapinnis = Selene setapinnis (Mitchill)

Alectis crinitis = A. ciliaris (Bloch)

Morone and Polyprion are included in the Percichthyidae by some authors

Roccus saxatilis = Morone saxatilis (Walbaum)

Centropristes striatus = Centropristis striata (Linnaeus)

Pseudopriaeanthus altus = Pristigenys alta (Gill)

Stenotomus versicolor = Stenotomus chrysops (Linnaeus)

Branchiostegidae placed in Malacanthidae by some authors

430 rosefish = redfish or ocean perch. Sebastes marinus is a complex of three
species: S. marinus, S. mentella Travin, and S. fasciatus Storer. A
second couplet can be added to the key:

2. Anal fin rays mostly 7, occasionally 8; total gill rakers on first arch
usually 36 or more; tip of pectoral fin usually not reaching anus . . .
S. fasciatus

Anal fin rays 8 or more; total gill rakers on first arch usually 35 or

less; tip of pectoral fin usually extends past anus

S. mentella

Hemitripteridae is included in Cottidae

Triglops ommatistius = Triglops murrayi Glinther

Myoxocephalus aeneus = M. aenaeus (Michill)

Myoxocephalus octodecimspinosus = M. octodecemspinosus (Mitchill)

Liparidae are included as a subfamily of the Cyclopteridae

Neoliparis atlanticus = Liparis atlanticus (Jordan and Evermann)

Liparis liparis = L. coheni Able, gulf snailfish

Lumpenidae is included in Stichaeidae. Pholis gunnellus is the only member

of the Pholidae in the key; the other species are Stichaeidae

Leptoclinus maculatus = Lumpenus maculatus (Fries)

Monocanthidae is considered as a subfamily of Balistidae by many authors

Balistes carolinensis = B. capriscus Gmelin

Alutera schoepfi = Aluterus schoepfi (Walbaum)

Alutera scriptus = Aluterus scriptus (Osbeck)

Sphaeroides = Sphoeroides, family Tetraodontidae

Chilomycterus belongs to the Diodontidae

Masturus lanceolatus = Mola lanceolata Lienard

Histrio pictus = H. histrio (Linnaeus)

Prepared by Bruce B. Collette, Research Associate in Ichthyology, MCZ; Marine Science
Institute, Nahant; and National Marine Fisheries Service Systematics Laboratory,
Washington, D. C.

December, 1981

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