Fishes of the western North Atlantic. Editorial board: editor-in-chief John Tee-Van [and others]

Fishes of the Western North Atlantic

MEMOIR

SEARS FOUNDATION FOR MARINE RESEARCH

Number I

Fishes of the
Western North Atlantic

PART ONE

L A N C E LE TS

Henry B. Bigelow, Museum of Comparative Zoology and
Isabel Perez Farfante, Museo ^oej>,
University of Havana

CrCLOSTO MES

Henry B. Bigelow a?id William C. Schroeder
Museum of Comparative Zoology

SHARKS

Henry B. Bigelow and William C. Schroeder

NEW HAVEN ig48
SEARS FOUNDATION FOR MARINE RESEARCH, YALE UNIVERSITY

COPYRIGHT 1948

SEARS FOUNDATION FOR MARINE RESEARCH

BINGHAM OCEANOGRAPHIC LABORATORY

YALE UNIVERSITY

Albert E. Parr, Editor
Yngve H. Olsen, A ssistant Editor

Fishes of the
Western North Atlantic

Editorial Board

Editor-in-Chief JOHN TEE- VAN

New York Zoological Society

CHARLES M. BREDER

American Museum of Natural History

SAMUEL F. HILDEBRAND

U. S. Fish and Wildlife Service

ALBERT E. PARR

American Museum of Natural History

WILLIAM C. SCHROEDER

Museum of Comparative Zoology
NEW HAVEN ig48

SEARS FOUNDATION FOR MARINE RESEARCH, YALE UNIVERSITY

Table of Contents

XI

xiii

Preface

Introduction

Maps xvi

LANCELETS, By Henry B. Bigelow and Isabel Perez Farfante i

Acknowledgments I

General Discussion I

Class Leptocardii I

Order Ampihioxi 3

Family Branchiostomidae y

Genus Branchiostoma 8

Branchiostoma bermudae 1 1

Branchiostoma caribaeum 1 3

Branchiostoma -platae 16

Family Epigonichthyidae 1 8

Genus Asymmetron 18

Asymmetron lucayanum 19

Am.'phioxides Larvae 23

A m.'phioxides felagicus 25

Amphioxides valdiviae 27

CYCLOSTOMES, By Henry B. Bigelow and William C. Schroeder 29

Acknowledgments 29

General Discussion 29

Class Agnatha 30

Subclass Cyclostomata 30

Order Myxinoidea 31

Family Myxinidae 3 1

Genus Myxine 32

Myxine glutinosa 34

Order Petromyzonida 43

Family Petromyzonidae 43

Genus Petromyzon 45

Petromyzon marinus 46

SHARKS, By Henry B. Bigelow and William C, Schroeder 59

Acknowledgments ro

General Discussion go

Class Chondrichthyes g2

Subclass Elasmobranchii 63

Order Selachii 64

vii

viii Table of Contents

Suborder Notidanoidea

77

Family Hexanchidae 78

Genus Hexanchus 78

Hexanchus griseus 80

Genus He-ptranchias 87

H e-ptranchias -perlo 88

Suborder Chlamydoselachoidea 93

Suborder Heterodontoidea 95

Suborder Galeoidea 95

Family Carchariidae 98

Genus Carcharias 98

Carcharias taurus lOO

Family Scapanorhynchidae 109

Family Isuridae 109

Genus Lamna 1 1 1

Lamna nasus 112

Gtmxs, I surus 123

Isurus oxyrinchus 1 24

Genus Carcharodon 133

Carcharodon carcharias 134

Family Cetorhinidae 146

Genus Cetorhinus 146

Cetorhinus maximus 147

Family Alopiidae 1 60

Genus A lopias 161

A lopias superciliosus 163

A lopias vulpinus 167

Family Orectolobidae 178

Genus Ginglymostoma 180

Ginglymostoma cirratum 181

Family Rhincodontidae 187

Genus Rhincodon 188

Rhincodon typus 189

Family Scyliorhinidae 1 9 5

Genus Scyliorhinus 202

Scyliorhinus boa 204

Scyliorhinus retifer 207

Scyliorhinus torrei 211

Genus Galeus 214

Galeus arae 2 1 6

Gtnus Aprisiurus 219

j^ pristurus profundorum 222

Apristurus riveri 21s

Table of Contents

Genus Pseudotriakis

Pseudotriakis microdon

Genus Triakis

Triakis barbouri

Genus Mustelus
Mtistelus cants
Mustelus norrisi

Mustelus fasciatus
Mustelus mento
Mustelus schmitti

Scoliodon terrae-novae
Genus Aprionodon

IX

Family Pseudotriakidae ^li

228

229

Family Triakidae 2^^

235
236
240
244

254
Genus Mustelus, Addendum 256

256
259
261

Family Carcharhinidae 262

Genus Galeocerdo 26 c

Galeocerdo cuvier 266

Genus Paragaleus 27 c

Paragaleus fectoralis 276

Genus Prionace ^gQ

Prionace glauca 282
Genus Scoliodon

292
295
303

320

329

A -prionodon isodon nQt

Genus Negafrion -,Qg

Negaprion brevirostris ojq

Genus Hypoprion » j ,

Hypoprion signatus o j g

Genus Carcharhinus

Carcharhinus acronotus
Carcharhinus jalciformis

Carcharhinus foridanus -> o o

Carcharhinus leucas o -> -^

Carcharhinus limbatus o^g

Carcharhinus longimanus 354

Carcharhinus maculipinnis ogj.

Carcharhinus milberti ogg

Carcharhinus nicaraguensis 073

Carcharhinus obscurus '>g2

Carcharhinus oxyrhynchus 30 j

Carcharhinus porosus 'in a

Carcharhinus remotus aqq

Carcharhinus springeri aqa

X Table of Contents

Family Sphyrnidae 407

Genus Sfhyrna 408

Sfhyrna bigelowi 410

Sfhyrna diflana 415

Sfhyrna tiburo 420

Sfhyrna tudes 428

Sfhyrna zy gaena 436

Suborder Squaloidea 449

Family Squalidae 450

Genus Squalus 452

Squalus acanthias 455

Squalus cubensis 473

Genus Squalus, Addendum 478

Squalus fernandinus 478

Genus CentroscylUum 480

Centroscyllium jabricii 482

Genus Eimofterus 487

Etmofterus hillianus 488

Genus Centroscymnus 493

Centroscymnus coelolefis 494

Family Dalatiidae 499

Genus Dalatias 500

Dalatias licha 502

Genus Isistius 508

Isistius brasiliensis 509

Genus 5omwo5MJ 514

Somniosus microcefhalus 5^6

Family Echinorhinidae 526

Genus Echinorhinus 526

Echinorhinus brucus S'^1

Suborder Pristiophoroidea 532

Suborder Squatinoidea 533

Family Squatinidae 534

Genus Squatina 534

Squatina dumeril 53 ^

Genus Squatina, Addendum 544

Squatina argentina 544

Index of Common Names 547

Index of Scientific Names 552

Preface

r

^ "If ^HE inhabitants of the waters of the earth have fascinated human beings ever
since "God created great whales, and every living creature that moveth." Our
interests have by no means been confined to the aesthetic or the gustatory 5 the
reflections of Isaac Walton are an earnest of the composure and rapport with the universe
that exists when fishes and their surroundings are contemplated; the mental relaxation of
fly fisherman or surf caster needs no defense or explanation; the life of fishes, their migra-
tions, their evolution, and the incredibly diverse facets of their activities, aflFord infinite
opportunities for study by the scientist. In latter years man's curiosity about the inhabitants
of "the water in the seas" has been increased and stimulated by his ever greater penetra-
tion into the deeps. Improved apparatus has enabled him to widen his sphere of effort and
to capture fish for his markets farther from shore and deeper down than heretofore. With
goggles and rubber fins he has pushed beneath the surface for momentary glimpses of
those which live below; with diving helmet and diving suit he has gone deeper and investi-
gated more closely; in the bathysphere he has dangled in the sea half a mile down and
checked on the lives of the strange fishes which make their home in that dark and cold
portion of the world.

Expeditions have gone forth with fishes as their prime consideration, and ichthyol-
ogists have studied what the expeditions brought back. Men and women in numerous
laboratories have worked upon fisheries problems, while countless numbers of fishermen,
professional and amateur, have added their bit to the knowledge of the whys and where-
fores of our fishes. All this has produced an enormous quantity of information and lore
which lies scattered in countless publications. The reason for the present series of volumes
is to correlate the contents of the rich storehouse of knowledge relating to the fishes that
live in the waters of the western North Atlantic.

This volume, the first of a series, describes the lancelets, the hagfishes and the lam-
preys, and those most interesting animals, the sharks. It has been written on the premise
that it should be useful to those in many walks of life — to those casually or vitally inter-
ested in the general phenomena of life in our waters, to the sportsman whose interests are
closely associated with pleasure and relaxation, to the fisherman whose livelihood depends
upon knowledge of where fishes are gathered together, as well as to the amateur ichthyol-
ogist and the professional scientist. Special stress has been given to the relationship of the
fishes to ourselves — in most cases this relationship is to man's advantage, but the present
volume also carries this theme in reverse — some sharks will attack man!

Introduction

HALF a century ago Jordan and Evermann's Fishes of North and Middle Amer-
ica was published, and up to the present time these volumes have continued to
be the only comprehensive descriptive account dealing with western Atlantic
fishes. With the progression of years this work has become less available and more obso-
lete, which is understandable in view of the scientific advances made during the intervening
decades.

Vast numbers of papers, both scientific and popular, have appeared since 1 896-1 900
— the dates of issuance of Jordan and Evermann's work. Numerous new genera and spe-
cies have been described 5 many groups of fishes have been subjected to detailed study and
revision, especially within the last two decades; new viewpoints on classification and
phylogeny have been presented; much additional information has been published on life
histories and habits of many species, and some regional studies of the fish faunas have been
made. However, this new information remains widely distributed in numerous books and
periodicals.

Since our knowledge of the fishes on this side of the Atlantic has reached a point of
relative stability, particularly with regard to purely descriptive accounts, the present time
seems especially suitable for a publication which embraces all of our knowledge of the
fish fauna of this region. To bring together and synthesize this scattered ichthyological
information and to make it available to both the public and to marine biologists is the
primary purpose of this work.

The first volume of Fishes of the Western North Atlantic brings to frui-
tion, at least in part, a plan which was conceived at New Haven some years ago. With
the establishment of the Sears Foundation for Marine Research at Yale University in
1937, funds became available for publication, and a group of interested ichthyologists met
to discuss the preparation of a work such as is here presented. To lay a firm groundwork and
to initiate production, the Editorial Board was formed, the members of which are Charles
M. Breder, Jr., Samuel F. Hildebrand, Albert E. Parr, William C. Schroeder, John Tee-
Van, and, until his death in 1944, the late J. R. Norman of the British Museum (Natural
History). Assisting the Editorial Board is an Advisory Committee: William Beebe (New
York Zoological Society), Rolf L. Bolin (Hopkins Marine Station), William K. Gregory
(American Museum of Natural History), Carl L. Hubbs (Scripps Institution of Oceanog-
raphy), Daniel Merriman (Bingham Oceanographic Laboratory), George S. Myers

xiv Introduction

(Stanford University), John T. Nichols (American Museum of Natural History), Luis
Howell-Rivero (University of Havana) and Leonard P. Schultz (U.S. National Mu-
seum).

The articles in this and subsequent volumes, which will be co-operatively produced
by many ichthyologists, are intended to be critical reviews or revisions of each group rather
than perfunctory compilations or mere reprintings of previously published works. An out-
line of the general classification has been prepared, based on widely accepted schemes of
classification (such as that used at the British Museum). Standards for both the text and
the illustrations have been formulated so as to "achieve a fairly uniform treatment for all
volumes. Under each species will be found both the distinctive characters which set it
apart from its nearest relatives, a detailed description, as well as discussions of its color,
size, general habits, abundance, range, relation to man (that is, its economic importance,
danger to man, sporting qualities, etc.), and its occurrence in the western Atlantic. Since
the publication will be used by lay persons as well as by ichthyologists and marine biologists,
the use of highly technical words and phrases has been avoided as far as possible. Because
of the large number of references which are included in a study of this nature, particularly
in the "Synonyms and References," abbreviations have been used throughout. References
to periodicals are listed and abbreviated in accordance with the standards established in
A World List of Scientific Periodicals, Published in the Years i^oo—igjj (Oxford Uni-
versity Press, Second Edition, 1934), and an approximate consistency has been developed
for books and periodicals not listed in that publication. The final volume will contain a
complete and extended bibliography. Common names which are most generally used have
been included} for future volumes it is possible that the recommendations of the Commit-
tee on Common Names of the American Fisheries Society will be available.

The geographical range of Fishes of the Western North Atlantic em-
braces the western half of the North Atlantic, including the adjoining gulfs and seas,
from Hudson Bay southward to the Amazon River. But this range is not strictly ad-
hered to in all instances; a number of species living close to the outer borders of the
region covered by this publication are included, particularly when their inclusion assists
in a more adequate understanding of the group under consideration. Brackish water species
are included, and naturally those which are cosmopolitan. As far as oceanic forms are con-
cerned, pelagic species are treated in full, while the strictly deep-sea (bathypelagic) fishes
are referred to only in keys and by references to the more recent reports describing these
animals. Two factors dictate this decision: i) The relative paucity and incompleteness of
our knowledge of these animals, and 2) the fact that they rarely, if ever, come within the
provenance of the nonspecialist in fishes, since special vessels and gear are required to eflFect
their capture.

The map which accompanies this first volume is by no means complete. Since it was
prepared before the manuscript was finished, all the localities given in the text could not
be included, particularly in such heavily worked areas as New England. However, it will

Introduction xv

serve to give at least a general idea of locations; in future volumes there will be a closer
relationship between the localities given in the text and those included on the map.

The expense incurred in the preparation of this volume has been extensive, and due
appreciation and thanks are extended to the Sears Foundation for its share in making publi-
cation possible and to the institutions that supported the work of the authors and editors.
Income derived from the sales of the volume will be used for the production of the re-
mainder of the publication.

The Editorial Board would like to express its appreciation and gratitude to Yngve H.
Olsen, Assistant Editor of the Sears Foundation for Marine Research, for his diligent and
able editing of the manuscripts and for the guidance of the publication through the press.

To Henry Sears the members of the Editorial Board owe a personal and collective
debt of gratitude for his understanding and for his unswerving continued support.

JOHN TEE-VAN

New York Zoological Society

North America

XVI

50*

45*

70* 65* 60* 55* 50*

South America

XVll

CHAPTER ONE

Lancelets

BY
HENRY B. BIGELOW and ISABEL PEREZ FARFANTE

ACKNOWLEDGMENTS

JVe are indebted to Thomas Barbour and Leonard P. Schultz for putting the
Lancelet collections of the Museum of Comparative Zoology and of the United
States National Museum at our disposal for study. Also, hearty thanks are
due to Gerardo Canet for preparing all the original drawings included here.

GENERAL DISCUSSION

The Lancelets of the western Atlantic Ocean are included in the present volume for con-
venience, following the precedent established in existing manuals of the fishes of various
parts of the world. Actually they are not fishes at all, although fish-like in appearance, but
belong to a separate subphylum (Cephalochordata) of the Chordata, since they are much
simpler in structure than are any of the true vertebrates of the subphylum Euchordata,
or Vertebrata.

Class LEPTOCARDII

The notochord, extending the entire length of the body and persisting throughout
life, is surrounded by a resistant sheath, this notochord and sheath forming a firm but
flexible supporting structure. But there is neither protective skeleton nor cranium for the
anterior part of the neural tube, no bony structures of any sort, and no jaws. The pharynx
in the adult is surrounded by an atrial chamber, formed by the outgrowth and coalescence
of two ridges (the metapleura) of the body wall; the pharynx opens into the atrium by a
double series of gill slits, the number of which continues to increase throughout life; pos-
teriorly, the atrial cavity opens to the exterior by a small aperture, the atriopore. The dorsal
nerve tube terminates anteriorly some distance behind the anterior end of the notochord;
it is much compressed laterally, and the only suggestion of a brain is that its axial canal
widens anteriorly into a cerebral vesicle. The nerves given oflF by the neural tube (except

2 Memoir Sears Foundation for Marine Research

for the first two) are dorsal and ventral in origin, but the dorsal and ventral roots do not
join, and there are no ganglia on the dorsal roots. The muscular system is segmented, the
successive muscle blocks, or myotomes, being separated one from the next by septa of
connective tissue, or myocomma. The final number of myotomes is established early in
life, but the number is somewhat variable in every species. The gonads are segmented. The
circulatory system is very simple; there is no heart, but the larger blood vessels are peri-
staltically contractile. There is a well developed coelom, or body cavity. The outer surface
of the body is clothed with an epidermis consisting of a single layer of columnar epithelial
cells, without scales or other hard epidermal structures, and without cilia except in the
mouth, pharynx, atrial cavity and intestine. There are no eyes and no limbs. The sexes are
usually separate although similar in external appearance, but hermaphrodites have been
reported on several occasions. Development is described below.

The Lancelets differ from all the higher groups of fish-like animals — cyclostomes,
elasmobranchs, chimaeroids, and bony fishes — in the following important morphological
features.

A. Their epidermis consists of a single layer of cells of ectodermal origin in con-
trast to several layers of cells in all higher groups.

B. They have no hard epidermal or tooth-like structures of any sort.

C. They have no eyes, no external nostrils and no true ears.

D. When adult, the pharyngeal region with the gill clefts is enclosed, on the ventral
side, in a so-called atrial cavity.

E. The gill clefts increase in number throughout life whereas in all the higher
groups their number is fixed.

F. They have no specialized internal respiratory structures, no true brain, no heart,
no trace of a cranium and no hard vertebral structures, cartilaginous or bony.

G. The notochord extends forward beyond the anterior end of the dorsal nerve tube.
H. Their blood is colorless, without red corpuscles.

I. The neural canal, entirely closed dorsally in higher vertebrates, extends through
the dorsal wall of the nerve tube as a longitudinal fissure, reminiscent of the ectodermal
infolding by which the tube is formed.

J. The excretory organs are nephridia-like rather than kidney-like, consisting of
numerous (up to 91) pairs of tubules in the pharyngeal region, each discharging inde-
pendently into the atrial cavity.

K. The gonads are numerous, compared to only a single pair in higher groups, and
segmentally arranged; each discharges its products directly into the atrial cavity, there
being no permanent genital ducts.

L. The lining of the intestine bears cilia.

The relationship that the Lancelets bear to the Cyclostomes and to higher fishes has
been actively discussed, one view being that they represent the specialization of some
primitive prevertebrate stage in evolution, another that they are degenerate descendants
of some early type of vertebrate comparable to the Cyclostomes that have developed pe-

Fishes of the Western North Atlantic 3

culiar adaptations for a very special mode of life. Perhaps the most that can be said at
present is that possibly they may be "fairly close to the primitive types from which the
vertebrates have arisen,'" although their atrial cavity has no parallel among the vertebrate
series.^

Order AMPHIOXI

Description. This order includes all known representatives of the subphylum. They
are slender, fish-like in external appearance, the body tapers at both ends and varies in
length from one to eight cm. at maturity; they inhabit tropical and temperate seas. In the
adult the buccal cavity, which leads into the mouth proper, opens on the ventral surface of
the body a little behind the anterior end. It is bounded laterally by a pair of expanded
muscular membranes, the so-called oral hood, the free edge of which bears 20 to 30
slender oral tentacles or cirri, each supported by a cartilaginous rod arising from a
cartilaginous ring situated immediately behind the margin of the hood. Proximally, the
inner surface of the oral hood bears a series of finger-like projections of ciliated epithelium,
jointly forming the wheel organ, the ciliary action of which drives water inward through
the buccal cavity to the mouth, and so to the pharynx. The mouth, at the bottom of the
buccal cavity, is very small and surrounded by a vertical membrane, the so-called velum,
from which several short velar tentacles project inward into the capacious pharynx. The
linings of the pharynx, and of the vertical gill clefts that pierce its two sides, are clothed
with cilia (those of the former having a complex pattern), the joint action of which is to
drive the water from the mouth, along the pharynx, through the gill clefts and so out
through the atrial cavity and atriopore. The pharynx serves chiefly as a feeding organ, as
described below.

The integument is expanded as a single continuous finfold which extends along the
ventral surface from close behind the atriopore, around the posterior end of the body,
thence forward along the dorsal surface and around the anterior end of the latter, where it
forms a snout or rostrum. The finfold thus surrounds the anterior end of the notochord and
contains a lymph space; in the dorsal fin this is segmentally divided by vertical septa into
a series of compartments known as fin-ray chambers and this is sometimes true of the
ventral fin as well. These chambers are partially subdivided by so-called fin rays, the
lateral and apical surfaces of which are free but the bases of which are connected with
the continuous ridge of connective tissue that is derived from the roof of the neural sheath.
The final number of rays and of ray chambers is established early in life, i.e., at a small size,
but is somewhat variable in all species. Anterior to the ventral fin the ventral surface of the
body also bears a pair of prominent longitudinal ridges called the metapleura. As a result
of their presence, the anterior part of the body is roughly triangular in cross section in
adults, the dorsal fin forming the apex of the triangle, the two metapleura its other two

1. Romer, Man and Vert., 19+1 : 10.

2. The atrium of the Lancelets, while analogous to that of the tunicates, cannot be regarded as homologous with the
latter, for the method of formation is very different.

4 Memoir Sears Foundation for Marine Research

angles, and the space between the latter forming its base, which is also the floor of the atrial
chamber.

There is a rather conspicuous pigment spot at the anterior end of the nerve cord,
which has been called an eye spot or median eye, but which appears not to be a light re-
ceptor. Also, an olfactory function has been ascribed to a small diverticulum from the
cerebral vesicle, but it is doubtful whether this is correct.

Habits. Lancelets spend most of the time buried in the sand, in an oblique position,
with the anterior end alone protruding.' If removed from the sand they swim actively,
bending the body from side to side with a sinuous eel-like motion} it is with this same
motion that they bore into the sand, which they do very rapidly. In most cases they burrow
tail foremost, but they have been seen to do this with the anterior end foremost, in which
case they then assume a U-contour to bring the anterior end out again from the sand. It
seems that adults of the genus Branchiostoma seldom emerge spontaneously from the
sand, or only for very brief periods, except at spawning time, for we find no record of their
capture in tow nets.* But Asymmetron has been so taken (p. 21 ).

It has long been known that they feed on microscopic organisms which they strain out
from the current of water that is drawn in through the mouth and driven by ciliary action
through the gill apertures to the atrium, to be expelled through the atriopore. The buccal
tentacles, folding over one another, prevent larger objects from entering. Particles small
enough to pass through this screen are carried inward to the pharynx, where they become
mixed with mucus and are driven against the gill bars. The cilia on the inner faces of the
latter, beating in a ventro-dorsal direction, then drive the mingled food and mucus to the
dorsal pharyngeal groove, along which it is swept to the oesophagus." Feeding appears to
be a continuous process. No doubt the diet includes whatever kinds of microscopic organ-
isms may be available at any given time and place. The intestines of the European Bran-
chiostoma lanceolatum have been found to contain diatoms chiefly, but also desmids, Fora-
minifera. Infusoria, Radiolaria, Cladocera and the eggs of various small invertebrates,
as well as plant detritus.* Diatoms have also been reported from the intestines of Lance-
lets from Ceylon^ and were again the most abundant item in the diet of young Branchios-
toma belcheri at Amoy, China, although the adults also contained the larvae of tunicates,
echinoderms and crustaceans.* At another time° this same species in the same general local-

3. For an excellent photograph of the European Branchiostoma lanceolatum in this situation, see Hagmeier and Hin-
richs (Senckenbergiana, 13, 1931 : fig. 3b, 4b, facing p. 258).

4. Hensen (Ergebn. Plankton-Exped. Humboldt Stiftung, / A, 1892: 24-25) reported the capture of young Lance-
lets up to several centimeters long in plankton nets. But the fact that none so large were to be found subsequently
in the collections (Goldschmidt, Dtsch. Sud-polar Exped., // Zool. 3, 1909: 235) suggests that the stated size
was an error.

5. Condensed from a detailed account of the feeding mechanism in Branchiostoma lanceolatum, by Orton (J. Mar.
biol. Ass. U.K., to [i], 1913 : 19). For an account of the passage of food material through the gut, see Barrington
(Philos. Trans., [B] 228, 1937: 271).

6. For a list of the food of B. lanceolatum compiled from various sources, see Franz (in Grimpe and Wagler, Tier-
welt N- u. Ostsee, Lief 7, 12b, 1927: 26).

7. Tattersall, in Herdman, Rep. Gov't. Ceylon Pearl Oyster Fish., Gulf of Manaar, pt. i, suppl. 6, 1904: 221.

8. Chin, Philip. J. Sci., 75, 1941 : 393- 9- Reeves, Ginling Coll. Mag. for Jan. 193 1 : 29.

Fishes of the Western North Atlantic 5

ity was found feeding chiefly on bacteria, with a few protozoa also. The intestines of some
of the specimens were filled with sand, showing that the oral tentacles do not always bar
entrance to inedible particles.

Development. The larval development of the Lancelets has been the subject of
several major investigations and has been much discussed in relation to the problem of the
ancestry of the vertebrates. The process in the European Branchiostoma lanceolatum,
which may serve as representative of the group, is briefly as follows.

Spawning takes place at sunset. The eggs are minute (o.i mm. in diameter) and float
freely in the water. Segmentation is not only complete but nearly equal and affords one of
the classic examples of endoderm formation by invagination. About twelve hours after
fertilization the embryo, now oval in shape and clothed externally with cilia, breaks out
from the vitelline membrane and swims near the surface by ciliary action. By_ about the
thirty-sixth hour the yolk is entirely absorbed} the mouth has appeared on the left-hand
side; the first gill opening has been formed in the midline, soon to shift to the right side,
however; and the anus has formed at the hinder end of the body a little to the left of the
midline. During subsequent larval development, which may occupy as much as three
months, the larvae live pelagically some distance below the surface of the sea, hanging
for the most part in a vertical position which is maintained by the action of the long cilia, or
flagellae, one of which is borne by each cell of the ectoderm. The larvae (Fig. i), which

Figure i. Branchiostoma lanceolatum Pallas; larva, with 6 1 myotomes, after Franz, a anus, gi gills, i intestine.
nto mouth, n notochord. nc nerve cord.

have a very characteristic appearance because of the swollen gill region in an otherwise
slender body, gradually assume the characters of the adult without any abrupt metamor-
phosis. The most striking of the external accompanying changes are in the numbers and
locations of the gill openings, and the formation of the atrium, of the atriopore and of the
adult mouth. The latter, at first on the left side and forming a most conspicuous feature of
the larva because of its enormous size, shifts to the midline and decreases in relative size
toward the end of larval life, while the preoral hood then develops above it. Additional
gill openings, up to the number of 14 or more, are formed successively along the mid-
ventral line, corresponding at first in number and location to the myotomes in that part of

6 Memoir Sears Foundation for Marine Research

the body but later losing this relationship. Of these primary gill openings, only the second
to ninth persist, however.

After the formation of the primary series of gill openings the number of segments
increases at the posterior end of the body, the final number being attained early in larval
life. In the meantime the embryonic tail fin, a simple ridge of columnar ectoderm cells,
is replaced by the adult fin; this forms as an ectodermal fold, enclosing serial expansions of
the body cavity, the ray chambers; the fin rays develop as columnar outgrowths of meso-
derm upward into these chambers. A secondary series of gill openings, eight or nine in
number, appear on the right side of the body, dorsal to the primary series; and each mem-
ber of each set, except the first, becomes U-shaped and then entirely subdivided by a dorso-
ventral bar. The primary series of gill openings then shift to the left side of the body, so
that from then on the larva is bilaterally symmetrical so far as the location of its gills is
concerned.

The metapleural ridges first appear in larvae with eight to ten gill openings of the
second series. The atrial cavity results from the union of the median sides of these ridges,
commencing posteriorly and progressing anteriorly. The canal so enclosed expands later-
ally in the pharyngeal region to the dimensions of the atrium of the adult, while it con-
tinues open posteriorly as the atriopore. During the formation of the metapleura the larva
abandons its pelagic habit and comes to lie on one side or the other on the bottom. By the
time the mouth has moved to the median position the oral hood has formed and the gills
have assumed the final symmetrical arrangement. The little Lancelet, now resembling the
adult in general appearance, buries itself in the sand; the only further change is the forma-
tion of pairs of tertiary gill openings, a process that continues throughout the life of the
individual. The curious asymmetry of the larval Lancelet has been much discussed, but in
our opinion none of the explanations which have been offered for it is adequate.

Gonads are formed in the second or third year, and the oldest noted among a large
collection of Branchiostoma belcheri was four years old.^°

Relation to Man. Lancelets are neither large enough or numerous enough to be of
any commercial value anywhere in the western Atlantic, except as subjects for biological
investigation; nor are they ever likely to be. However, near Amoy in southern China there
has long been a fishery for Lancelets. Recently this employed about four hundred men in
two hundred boats who fished with shovel- or scoop-shaped dredges from two to four
hours each day on the ebb tide from August until April. This fishing ground is only about
six miles long and less than one mile wide, but it has been estimated that the annual catch
is in the neighborhood of 35 tons, or more than one billion Lancelets. Some of these are
consumed in the near vicinity, while others are dried and shipped to Java and Singapore."
Lancelets are also used occasionally as food in Naples and Sicily.'^

10. See Wells (Science, N.S. 64, 1926: 188) for Branchiostoma caribaeum from Florida; Chin (Philip. J. Sci., 75,
1941 : 400) for B. belcheri from Amoy, China.

1 1. For more detailed accounts, from which the foregoing is condensed, see Light (Science, N.S. 5^, 1923 : 57) *nd
Chin (Philip. J. Sci., t$, 1941 : 369).

12. Franz, in Grimpe and Wagler, Tierwelt N- u. Ostsee, Lief 8, b, 1927 : 44.

Fis/ics of tJic Western North Atlantic j

Fni/iilit's. The order includes two well defined families, Branchiostomidae and
Epigonichtlnidae, separated as indicated in the following key. A third assemblage of pe-
lagic forms, usually grouped together as the genus Amp/iio.xides, have sometimes been
classed as a third family, Amphioxididae. But their chief distinguishing characters — mouth
on the left side, atrial chamber unclosed and gill slits in an unpaired medio-ventral series —
are those of larval Lancelets in general at an early stage of development (p. 23), and it
now seems established in fact that they are larvae that have continued their pelagic exist-
ence for one reason or another until much larger and much further advanced in develop-
ment than is usually the case, rather than taking to the bottom at a smaller size, as most of
them do.*' In fact, we think it is likely that these Amphioxides larvae ne\'er do descend to
the bottom once they are carried out over deep water, but that they simply continue to exist
for an indefinite period as they are swept along with the currents, finally perishing without
producing offspring. On the other hand, it has been suggested that their existence may
provide a means for the dissemination of the species. Up to the present time, none of them
has been positively connected with any particular parent species.

Key to Families'*

I a. Mouth nearly median, with oral cirri; closed atrial chamber and atriopore; a series of
gill clefts on either side.

2a. Series of gonads developed on each side; both metapleura terminating close be-
hind atriopore. Branchiostomidae, p. 7.
2b. Gonads developed on right side only; the right metapleuron continuous with
ventral fin, the left-hand metapleuron terminating behind atriopore.

Epigonichthyidae, p. 18.

lb. Mouth on left side without oral cirri; no closed atrial chamber; gill clefts in a single

series along ventral side. Amphioxididae, p. 23.'^

Family BRANCHIOSTOMIDAE

Description. Mouth nearly in midline, surrounded by oral cirri; tentacles with lateral
sensory papillae, giving them a toothed appearance; closed atrial chamber; a series of gill
slits on each side; gonad pouches developed on both left and right sides; both metapleura
terminate close behind atriopore, including between their posterior ends the anterior end
of the ventral fin; rostral fin continuous with right side of oral hood, but not with left side;
posteriorly the median fin is expanded both dorsally and ventrally in lancet form as a
distinct caudal fin, with its ventral lobe lying to the right of anus; ventral fin-ray cham-
bers, except for the more anterior and more posterior, each contain a pair of fin rays in most
species, although perhaps only a single fin ray in some;"^ but dorsal fin-ray chambers con-

13. For a recent discussion, see Goldschmidt (Biol. Bull. Wood's Hole, 6^, 1933: 321).

14. Ainpliioxididae included to facilitate identification. 15. See above discussion of these.

16. Willey (Quart. J. micr. Sci., 44, 1891: 270) stated that in his Dolic/ior/iync/ins injicus the ventral fin rays arc
sintj'le; but they appear as paired in his illustration.

8 Memoir Sears Foundation for Marine Research

tain a single series of fin rays only; rostral fin, with anterior part of dorsal fin, lacks fin
rays; dorsal fin-ray chambers much more numerous than myotomes, with four or five
chambers to each myotome; the atrial chamber extends posterior to atriopore as a single
blind sac as far as the anus; olfactory pit present.

Genera. Two genera, Branchiostoma and Dolichorhynchus, are commonly recognized
in the family and are separated as indicated in the following icey. In addition, a new sub-
genus of Branchiostoma has recently been proposed under the name Amfhifleurichthys^''
for a species in which "the form is more elongated and less robust" than in Branchiostoma,
"with the myotomes more acutely tapering at each end of the animal," and in which the
"caudal fin is reduced to a low fold."" But the differences appear to us specific, rather
than generic.

Key to Genera

I a. Rostral process, including anterior end of notochord, extends far beyond preoral
hood. Dolichorhynchus Willty, 1901

Ceylon.

lb. Rostral process, including anterior end of notochord, extends only a short distance
beyond preoral hood. Branchiostoma Costa, 1 834, p. 8.

Genus Branchiostoma Costa, 1834

Branchiostoma Costa, Ceni. Zool., 1 834: 49; type species, B. lubricum Costa. Naples.

Generic Synonyms:

Limax Pallas, Specil. Zool., Fasc. 10, 1774: 19, pi. 1, fig. 11; for L. lanceolatus Pallas, Cornwall; not Limax
Linnaeus, 1758.

Giuterobranchus ? Rasch, Mag. Naturvid., Physiogr. Foren. Christiania, 12 (2) 2, 1836: 325, footnote; evi-
dently Branchiostoma, from the excellent account, but only provisionally identified by that author; west-
ern Norway; not Gasterobranchus Bloch, 1795, which is a synonym for the cyclostome Myxine Linnaeus,
1758.

Amfhioxus Yarrell, Brit. Fish., 2, 1836: 468; type, Limax Icmceolatus Pallas, 1774.

Amfhifleurichthys (subgenus) Whitley, Aust. Zool., 7 (3), 1932: 256; type, A. minucauda Whitley. Queens-
land.

Generic Characters. The rostral process, including the anterior end of the notochord,
projects for only a short distance beyond the preoral hood; the characters are otherwise
those of the family.

Range. European coasts from northern Norway to the Mediterranean, the Black Sea
and tropical West Africa; western Atlantic from Chesapeake Bay'* to the Rio de La Plata
(including Bermuda); Pacific coast of the Americas from Middle California to Chile;
Japan; China; East Indies; Philippines; Queensland; India; Ceylon; Madagascar; East
and South Africa.

17. Whitley, Aust. Zool., 7 (3), 1932: 256.

1 8. Lancelets were said by Garman (in Kingsley, Stand. Nat. Hist., 5, 1885 : 62) to range as far north as New York;
but we find no positive record of any member of the group in the western Atlantic farther north than Chesa-
peake Bay.

Fishes of the Western North Atlantic 9

Species. The characters that have been used chiefly in the classification of the species
of the genus are: (i) number of ventral fin-ray chambers; (2) number of dorsal fin-ray
chambers; (3) height of dorsal fin in relation to height from its crest to the margins of
metapleura; (4) shape of caudal fin; (5) location of anus in lower lobe of caudal fin; (6)
number of preatrial myotomes; (7) total number of myotomes. The five species that have
been described from the western Atlantic {B. caribaeum Sundevall, 1853; ^- bennudae,
B. floridae, B. platae and B. virginae Hubbs, 1922) with the anus near, or posterior to, the
midpoint of the ventral lobe of the caudal fin differ sharply from B. lanceolatiim and
B. ajricae of the eastern Atlantic, in which it is considerably farther anterior to it. Among
this western Atlantic group, B. bermudae and B. platae are set apart by the fact that the
lower lobe of the caudal fin originates considerably anterior to the origin of its upper lobe
(Fig. 2 A, F), whereas in the others the two lobes originate opposite one another. B. ber-
mudae is sharply separated from B. platae by a considerably smaller number of dorsal fin-
ray chambers (200-242 vs. 278-330), and fev/er myotomes (not more than 56 vs. at
least 58). But B. fioridae and B. virginae agree with B. caribaeum in the position of the
anus, while counts of fin-ray chambers and myotomes in the specimens we have studied
(Study Material, p. 13), together with those previously published, fail to show any clear
distinctions among the populations of Virginia, North Carolina, Florida (including the
Tortugas) or Porto Rico (representing the West Indian region). The most that can be
said is that some Florida and West Indian specimens have fewer precaudal fin-ray cham-
bers than have yet been recorded for more northerly localities. But this is not always true,
since the maximum recorded counts are in fact for one specimen from Florida and for one
from North Carolina. Therefore it cannot be invoked as a basis for specific separation.

Key to Species of Branchiostonia

la. Caudal fin hardly higher than dorsal and ventral fins. w/«M^^«iirt Whitley, 1932.

Queensland.
lb. Dorsal or ventral lobe of caudal fin, or both, considerably higher than dorsal and ven-
tral fins.
2a. Caudal fin not clearly marked off from ventral fin. capense Gilchrist, 1 902.

South Africa.

2b. Caudal fin clearly marked off from ventral fin.
3a. Anus about at point of origin of caudal fin.

4a. Distance from anus to tip of caudal fin only Y-> distance from anus to
atriopore. bazarutense Gilchrist, 1923.

East Africa.

4b. Distance from anus to tip of caudal fin about as great as from anus to
atriopore. haeckeli Vvd^nz, i<)iz.

Ceylon.

3b. Anus clearly posterior to origin of caudal fin.

lO Memoir Sears Foundation for Marine Research

5a. Anus far in advance of midpoint of lower lobe of caudal fin.

6a. 77 or more myotomes. elongattnn SundcvM, li ^2.

West coast of South America, Chile
to Galapagos Islands.

6b. Not more than 73 myotomes.

7a. 42 to 44. myotomes anterior to atriopore.

africae liuhbs, 1927.
Tropical West Africa.
7b. Not more than 41 myotomes anterior to atriopore.

8a. 68 to 72 myotomes in all. tattersalU Hubbs, 1922."

Ceylon.

8b. Not more than 66 myotomes.

9a. Not more than 62 myotomes; ventral lobe of cau-
dal fin a little longer than distance from its origin
to atriopore; anus clearly anterior to origin of
dorsal lobe of caudal fin.

lanceolatumV3.\\a.i, 1778.
Northern Norway to Mediter-
ranean and Black Sea.
9b. At least 63 myotomes; ventral lobe of caudal fin
only as long as distance from its origin to the atrio-
pore; anus below origin of dorsal lobe of caudal
fin. belcheriGrzy, 1%^-].

Japan, China, the East
Indies, Philippines, In-
dia, and Ceylon to East
Africa.

5b. Anus near midpoint of lower lobe of caudal fin or posterior to it.

lOa. Origin of lower lobe of caudal fin considerably anterior to origin
of its upper lobe.

1 1 a. Not more than 242 dorsal fin-ray chambers or 56 myo-
tomes, bermudae Hubbs, 1922, p. 11.
lib. At least 278 dorsal fin-ray chambers.

1 2a. Rostrum not marked oif from dorsal fin by a notch;

65-74 myotomes. calif orniense Andrews, 1893.

Monterey, California to Gulf of

California.

12b. Rostrum marked off from dorsal fin by a notch; 59-65

myotomes. flatae Hubbs, 1922, p. 16.

lOb. Origin of lower lobe of caudal fin about opposite origin of its

upper lobe. caribaeum SundtvaW, 1853, p. ^3-^°

19. Including gravelyi Prashad (Rec. Indian Mus., ^6, 1934: 333).

20. Including jloridae Hubbs, 1922, and virginiae Hubbs, 1922.

Fishes of the Western North Atlantic ii

Branchiostoma bermudae Hubbs, 1922
Figure 2 A-D

Study Material. Nineteen specimens, 29 to 49 mm. long, from Bermuda (U. S. Nat.
Mus. and Harv. Mus. Comp. Zool.).

Distinctive Characters. Among Atlantic species, B. bermudae differs noticeably from
B. lanceolatum and from B. ajricae in that its anus is about opposite the midpoint of the
lower lobe of its caudal fin. In this respect it closely resembles B. flatae and B. caribaeum,
but it is separable from both of these by a smaller number of myotomes ( 56 at most) as well
as by generally fewer precaudal fin-ray chambers (9 to 24, usually less than 16). The
average number of dorsal fin-ray chambers also is smaller.

Additional Description. Anterior end of notochord extending forward in rostrum in
a straight line; rostral fin marked off from dorsal fin by a subtriangular notch; origin of
lower lobe of caudal fin anterior to origin of its upper lobe by a distance about ^/-j, as great
as length of lower lobe; dorsal fin % to /^ as high as distance from its base to margin of
metapleura in the midregion of body; anus a little behind the midpoint of lower lobe of
caudal fin; origin of lower caudal lobe about midway between its tip and atriopore; dis-
tance from tip of caudal to anus about 0.4 of distance from anus to atriopore; dorsal fin-ray
chambers 204 to 242, the highest 3 to 4 times as high as long; precaudal fin-ray chambers
9 to 24; 35 or 36 myotomes anterior to atriopore; 12 to 14 between atriopore and anus,
5 to 7 posterior to anus, total number 54 to $6; gonads, 22 to 28 pairs.

Color. Living specimens are semitransparent and iridescent, but they become opaque
after preservation.

Size. Maximum recorded length, 53.5 mm."'

Habits. The Bermuda Lancelets are usually found in one-half to six fathoms of water
on coarse sandy bottom into which they burrow tail first and there remain most of the time
with only the anterior part of the body exposed. If disturbed they swim vigorously for a
short time but soon return to the sand. Observations in aquaria have shown that normally
they are no more active by night than by day. Under experimental conditions they usually
swim with the anterior end foremost. If a stimulus is applied to the anterior end, the
Lancelet may dart backward for a short distance, or it may turn end for end. But this
reversal in direction is of short duration, for it soon turns again and proceeds at only a
slight angle from its original course. In taking to bottom after swimming, Lancelets usu-
ally sink quietly through the water to the sand; when in contact with the latter they may
either lie there, passive for some time; or they burrow at once, usually tail first, or head
first on rare occasions. When buried they usually are tortuous in outline, probably from
being crowded among the grains of sand."

Specimens adapted to the summer temperatures of Bermuda (about 31° C.) dart

2 1. Hubbs, Occ. Pap. Mus. Zool. Univ. Mich., 105, 1922: 10.

22. For detailed accounts of the experiments on which the above is based, see Arey (J. exp. Zool., 29 [i], 1915 : 37)
and Parker (Proc. Ainer. Acad. Arts Sci., ^3 [16], 1908: 413).

R 5 "J — ° " ^
fe S 2 fc< CO V. "u

Fishes of the Western North Atlantic 13

rapidly about for a short time if the temperature be either raised or lowered. If heated to
40° C. or higher they diej if chilled to 10° C. they become inactive and may die, as they
invariably do if kept in a temperature of 4° C. for half an hour. But the thermal reactions
are not known for specimens adapted to the winter temperatures that prevail at Bermuda.

It has been found that B. berniudae tends to swim away from a source of light; also it
is stimulated to activity by the presence of light, «.<?., it is photokinetic," and hence it may
be expected to bore deeper into the sand if strongly illuminated, as by the sun. But it is more
sensitive to mechanical than to photic stimulation, as is the European B. lanceolatum.^*
This is especially true of the preoral tentacles and of the outer fringes of the oral hood,
which close and open with a sudden winking motion if touched. It is through this reac-
tion that the Lancelet rids itself of the debris that may accumulate on its preoral tentacles,
for when these become laden they contract sharply to loosen any waste particles, which
are then swept away by water that is expelled simultaneously from the cavity of the oral
hood.

Presumably it spawns chiefly in late spring, for the peak of the breeding season is
passed before June-July.

Range. Bermuda.

Synonyms and References:

Branckiostoma lubricum Goode, Amer. J. Sci., 14, 1877: 293 (Bermuda); not B. lubricum Costa, 1834.

Amfhioxus (no specific name) Brooks, 3rd Annu. Rep. Johns Hopk. Univ., 1878: 54 (Bermuda).

Branchiosloma caribaeum Bristol and Carpenter, Science, N.S. //, 1900: 170 (Bermuda); \'errlll. Trans.
Conn. Acad. Arts Sci., 11, 1901: 55 (Bermuda); Bean, Field Mus. Publ. Zool., 7 (2), 1906: 29 (Ber-
muda); Kutchin, Proc. Amer. Acad. Arts Sci., ./p (10), 1913: 571 (peripheral nervous system).

Amfhioxus caribaeu! Mark, Science, N.S. 20, 1904: 179 (Bermuda).

Branchloitoma caribhacum Barbour, Bull. Mus. comp. Zool. Harv., 46, 1905: 1 10 in part (specimen from Ber-
muda) ; Parker, Proc. Amer. Acad. Arts Sci., 4^ ( 16), 1908; 413 (sensory reactions, Bermuda) ; Arey, J.
exp. Zool., 2p (i), 1915: 37 (swimming habits, Bermuda).

Branchiosloma carribaeum Mark and Crozier, Anat. Rec, 11 (6), 1917: 520 (photo receptors); Conklin, J.
Morph., 54 (l), 1932: 70 (breeding season at Bermuda); not B. caribaeum Sundevall, 1853.

Branchiosloma bermudae Hubbs, Occ. Pap. Mus. Zool. Univ. Mich., 105, 1922: 9 (descr., discus., Bermuda) ;
Jordan, Evcrmann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 7 (Bermuda); Becbe .ind
Tec-Van, Field Bk. Shore Fish. Bermada, 1933: 2 (descr., ill., Bermuda); Goldschmidt, Biol. Bull.
Wood's Hole, 64 (3), 1933: 321 (Bermuda) ; Pratt, Manual Common Invert. Anim., 1935: 757 (no. of
myotomes and gonads, Bermuda) .

Branchiostoma caribaeum Sundevall, 1853

Figure 2 E

Study Material. Numerous specimens, 12 to 66 mm. long, from Maryland, Chesa-
peake Bay, Virginia, North Carolina, eastern and western Florida, the Tortugas, Florida,
and Vieques Island, Porto Rico.

13. Parker, Proc. Amer. Acad. Arts Sci., 4} (16), 1908: 426.
24. Franz, Wiss. Meeresuntersuch. Helgoland, 15 (14), i924'- 6.

14 Memoir Sears Foundation for Marine Research

Distinctive Characters. B. carihaeum differs from B. lanceolatum and from B. africae
in that its anus is about in the middle of the lower lobe of the caudal fin. It is distinguished
from B. bermudae by the shape of the caudal fin and by the origin of the ventral lobe below
that of the dorsal lobe j by the position of the anus, in advance of the midpoint of the lower
lobe of the caudal finj and by the generally greater number of myotomes and dorsal fin-
ray chambers (at least 230 of the latter). It is separated from B. flatae by the shape of the
caudal fin, as well as by the position of the anus and by its tendency to have fewer myotomes
and dorsal fin-ray chambers.

Additional Description. Anterior end of notochord in rostrum extending forward
in a straight line 5 rostrum marked ofF from dorsal fin by a subtriangular notch; caudal fin
symmetrically lanceolate with narrowly rounded tip, its lower lobe considerably higher
than ventral or dorsal fins, its origin opposite origin of its upper lobe and about midway
between tip of caudal fin and atriopore; distance from tip of caudal to anus about V3 dis-
tance from anus to atriopore; dorsal fin Ys as high as distance from its crest to margins of
metapleura in midregion of body; highest dorsal ray chambers 5 to 8 times as high as
long; dorsal ray chambers 230 to 320; precaudal (ventral) fin-ray chambers 18 to 37;
35 to 38 myotomes anterior to atriopore, 13 to 17 between atriopore and anus, and 6 to 9
posterior to anus, recorded totals, 57 to 64; gonads 22 to 29.

Recorded counts for specimens from different localities.

Dorsal

Precaudal

Locality

fin-ray chambers

fin-ray chambers

Myotomes

Virginia and North Carolina

256-320

33-42

58-64"

Florida, including Tortugas

274-330

18-28

57-61

Porto Rico; Bahamas

227-300

15-37

58-61

Color. Live specimens are flesh-color or semitransparent, with a metallic iridescence;
those kept in alcohol become opaque and whitish.

Size. The greatest length so far recorded is 66 mm. (see Study Material, p. 13).

Developmental Stages. In the Chesapeake region the pelagic larvae, of the sort
usual for the group (p. 5), tend to settle to the bottom by the time they have reached
a length of about 7.5 to 8 mm.^*

Habits. The adults, like those of other species, live buried in coarse or fine sand. In
Florida, and presumably elsewhere also, they are most numerous along the edges of sand
bars just below the low tide mark where their presence is indicated by small holes in the
sand. If the sand is laid bare by a low run of tides it appears that they simply burrow more

25. Andrews (Stud. Biol. Lab. Johns Hopk. Univ., 5, 1893: 241) reports a total of only 48 myotomes for a speci-
men from Jamaica. But this is so much fewer than any other recorded count that we judge it to have been erro-
neous; see also Franz (Jena Z. Naturw., 5*, 1922: 399).

26. See Rice (Amer. Nat., t^, 1880: 17, pi. i, fig. 5) for a good illustration of the late larva.

Fishes of the Western North Atlantic 15

deeply for the time being, instead of moving down the slope." If driven out of their holes,
as when a shovel is thrust into the sand close by, they shoot upward into the water and
swim vigorously for a brief period with either the ventral or dorsal side uppermost, but
always .with the anterior end foremost. However, they soon sink to the bottom again.
"Generally as soon as they touched the sand, they would half-arch their bodies and almost
instantaneously disappear from sight . . . after their disappearance, they very rarely
entirely emerged"'* but continue buried in an oblique position, ventral side uppermost,
either with the opening of the oral hood at the surface of the sand or with the anterior
portion of the body protruding. Aquarium observations suggest that they protrude and
feed chiefly at night. No specific information is available as to the diet of this species (p.
4). B. caribaeum has been recorded from the low tide zone down to a depth as great
as 24 fathoms. In Florida, sexually mature males, and females "heavy with eggs,""' have
been reported in March; they are to be expected perhaps two months or so later in the
Chesapeake Bay region, where pelagic larvae are to be found in July and August. Sexual
maturity is attained in the second or third year.

Range. Atlantic coast of America from Chesapeake Bay to the West Indies. Recorded
localities are: several localities in Chesapeake Bay; North Carolina; many localities in
Florida, both on the west coast north to Pensacola and on the east coast; the Tortugas;
the Snapper Banks; Gulf of Mexico; Bahamas; Porto Rico; Jamaica. It is so common
in Florida that one collector reports taking 5,000 of them.

Synonyms and References:

Branchiostoma caribaeum^" Sundevall, Ofvers. Vet. Akad. Forh., Stockholm, lo, 1853: 12 (in part, specimens
from St. Thomas, West Indies) ; Gill, Rep. U.S. Comm. Fish. (1871-1872), 1873: 814 (listed, C. Hat-
teras to Fla.) ; Yarrow, Proc. Acad. nat. Sci. Philad., 1877: 218 (Bird Shoal, N. Carolina); Jordan and
Gilbert, Bull. U.S. nat. Mus., 16, 1883: 3 (Chesapeake Bay to West Indies); Gunther, Rep. Zool. Coll.
"Alert," Brit. Mus., 1884: 32 (in part, specimens from St. Thomas, West Indies, characterization,
discus.); Garman, in Kingsley, Stand. Nat. Hist., 5, 1885: 64 (Gulf of Mexico); Andrews, Stud.
Biol. Johns Hopk. Univ., 5, 1893: 240 (in part, specimens from Florida, Gulf of Mexico and West
Indies, myotome formula); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (l), 1896: 3 (in part,
but B. flatae also included);'' Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 48
(Snapper Banks, Gulf of Mexico, and Tampa, Florida) ; Evermann and Marsh, Bull. U.S. Fish Comm.,
20 (l), 1902: 59 (in part, Porto Rico, but not the ill., which probably is B. flatae because of shape of
caudal); Tattersall, Trans. Lpool. Biol. Soc, 77, 1903: 271, 280 (comp. with lanceolatum) ;
Lonnberg, Bronn's Klassen., 6, Abt. I, Buch 1, 1904: 339 (descr., distrib.) ; Barbour, Bull. Mus. comp.
Zool. Harv., 46, 1905: 1 10 (in part, but bermuiae and flatae also incl. because of distrib.) ; Fowler, Proc.
Acad. nat. Sci. Philad., ^g, 1908: 461 (Vieques Island, Porto Rico) ; Smith, N. C. Geol. econ. Surv., Fishes,
2, 1907: 27 (N. Carolina, but not the ill., which appears to be of B. flatae because of shape of caudal) ;
Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907: 146, 199 (in part, but B. flatae probably included also) ;
Fowler, Proc. biol. Soc. Wash., jj, 1920: 143, footnote (mentioned); Hubbs, Occ. Pap. Mus. Zool.

27. For an interesting account of their occurrence in Florida, and of methods of collecting them, see Wells (Science,
N.S. 64, 1926: 187).

28. Rice, Amer. Nat., i^, 1880: 8. 29. Wells, Science, N.S. 64, 1926: 188.

30. Sometimes spelled caribbaeum.

31. The illustration of caribaeum by Jordan and Evermann (Bull. U.S. nat. Mus., 47 [4], 1900; pi. i, fig. i)
appears from the shape of the caudal to have been based on a specimen of B. flatae.

1 6 Memoir Sears Foundation for Marine Research

Univ. Mich., 105, 1922: 6 (descr.) ; Ribeiro, Fauna brasil., Peixes, 2 (i), Fasc. I, 1923: 4 (in part,
but B. flatae included because of loc.) ; Meel: and Hildebrand, Field Mus. Publ. Zool., 75 (i), 1923: I
(descr., but B. flaiae included because of loc.) ; Nichols, Ann. N. Y. Acad. Sci., 10 (2), 1929: 180, fig. I
(descr., distrib., Porto Rico) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 7
(West Indies); Pratt, Manual Common Invert. Anim., 1935: 757 (no. of myotomes, West Indies);
Longley and Hildebrand, Pap. Tortugas Lab., No. 34, 1941: I (listed for Tortugas, Florida); Fowler,
Monogr. Acad. nat. Sci. Philad., 7, 1945: 262 (Sanibel, Florida).

Branchiosloma lanceolatum Giinther, Cat. Fish. Brit. Mus., 8, 1870: 513 (in part, specimens from Caribbean
Sea) ; Rep. Zool. Coll. "Alert," Brit. Mus., 18S4: 32 (in part, specimens from N. America) ; Jordan and
Gilbert, Bull. U.S. nat. Mus., 16, 1882: 867 (in part, specimens from east coast of N. America) ; Adams
and Kendall, Bull. U.S. Fish Comm., p, 1891: 292, 293, 298 (SW. Florida); Andrews, Stud. Biol.
Johns Hopk. Univ., 5, 1893: 239 (myotome formula of specimens from Chesapeake B.iy) ; not Limax
lanceo/alusTMas, 1 774.

AmpAioxus cariiaeus Jordan and Gilbert, Proc. U.S. nat. Mus., /, 1879: 388; also, Smithson. misc. Coll., tp,
1880: 388 (Bird Shoal, N. Carolina) ; Kirkaldy, Quart. J. micr. Sci., 37, 1895: 313 (in part, descr., and
distrib., but B. fUtae also included); Lonnberg, Bronn's Klassen: 6, Abt. i, Buch I, 1904: 239 (descr.,
distrib.).

Branchiostoma lubricum Goode and Bean, Proc. U.S. nat. Mus., 2, 1880: 121 (E. Florida, name only).

Amphioxus lanceolatus Rice, Amer. Nat., 14, 1880: i, 73, pi. 34, fig. i, 2 (habits, struct., develop.); not
Limax lanceolatus Pallas, 1 774.

Branchiostoma lanceolata Gill, Proc. U.S. nat. Mus., 5, 1883: 515 (Atlant. coast of U.S.) ; not LiTuax lanceo-
latus Pallas, 1774.

Amfhioxus (no specific name) Wright, Amer. Nat., 24, 1890: 1085 (Port Tampa, Florida) ; Andrews, Circ.
Johns Hopk. Univ., //, 1892: 75 (young stages recorded from Jamaica) ; Wells, Science, N.S. 64, 1926:
187 (ecology, habits, breeding season, age at sexual maturity and coll. methods, Florida).

Branchiostoma caribkatum Tattersall, Trans. Lpool. Biol. Soc, 77, 1903: 241, 280 (cf. with B. lanceolatum
and B. belcheri) .

Branchiostoma ftoridae Hubbs, Occ. Pap. Mus. Zool. Univ. Mich., 105, 1922: 7 (descr., Tampa and other
Florida loc); Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 4 (Florida); Jordan, Evermann and
Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 7 (Florida); Pratt, Manual Common Invert. Anim.,
1935: 757 (no. of myotomes, gonads, Florida).

Branchiostoma virginiae Hubbs, Occ. Pap. Mus. Zool. Univ. Mich., 103, 1922: 8 (descr., Chesapeake Bay);
Hildebrand and Schroeder, Bull. U.S. Bur. Fish., 4s, 1928: 42 (descr., habits, Chesapeake Bay) ; Breder,
Field Bk. Mar. Fish. Atlant. Coast, 1929: 4 (Chesapeake Bay) ; Jordan, Manual ^^ert. Anim. NE. U.S.,
1929: 4 (descr., Chesapeake Bay to N. Carolina) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish.
(1928), 2, 1930: 7 (Chesapeake Bay to Florida) ; Cowles, Bull. U.S. Bur. Fish., 46, 1 931: 367 (Chesa-
peake Bay); Pratt, Manual Common Invert. Anim., 1935: 757 (no. of myotomes, Chesapeake Bay to
Florida).

Branchiostoma flatae Hubbs, 1922

Figure 2 F

Study Material. Thirty-six specimens, 31 to 5 1 mm. long, from the vicinity of Rio de
Janeiro and San Sebastiao I., Brazil, and oflF the mouth of the Rio de La Plata, Argentina
(Lat. 36° 43' S.} Long. 56° 23' W.), in the collection of the United States National
Museum.

Distinctive Characters. B. flatae differs from the two eastern Atlantic species of this
genus {lanceolatum, africae), and from caribaeum as well, in having its anus considerably
posterior to the midpoint of the lower lobe of its caudal finj it differs further from cari-
baeum in that the lower lobe of its caudal fin originates considerably anterior to the origin

Fishes of the Western North Atlantic 17

of the upper lobe. The number of myotomes and dorsal fin-ray chambers is often larger
also in B. flatae, although there is no clear distinction between the two in these respects.
B. flatae differs from B. bermudae (which it resembles in the shape of the caudal fin) in
its more numerous myotomes (at least 59) and dorsal fin-ray chambers (at least 275).

Additional Description. Anterior end of notochord in rostrum extends forward in a
straight linej rostrum marked ofF from dorsal fin by a shallow notch j caudal fin lanceolate
but asymmetrical, the origin of its lower lobe anterior to origin of its upper lobe by a dis-
tance V3 to % as great as length of upper lobe, about midway between its tip and atrioporej
anus considerably posterior to midpoint of lower lobe of caudal; distance from tip of
caudal to anus V3 as long as from anus to atriopore; dorsal fin Y^ to ^,4 as high as dis-
tance from its crest to the margins of the metapleura; highest dorsal fin-ray chambers
3 to 6 times as high as long; dorsal fin-ray chambers 280 to 330; precaudal fin-ray cham-
bers 19 to 33; myotomes 37 to 40 anterior to atriopore, 13 to 17 between atriopore and
anus, and 6 to 9 posterior to the anus, the recorded totals from 58 to 65; gonads 26 to 31.

Color. Presumably as in B. caribaeum (p. 14), but no specific inform.ation is avail-
able.

Size. Recorded specimens have ranged from 28 to ^d mm. in length.

D evelof mental Stages. Presumably as in other members of the genus.

Habits. Nothing is known of the habits of B. flatae to differentiate it from its rela-
tives.

Range. Specimens positively identified as B. flatae are known up to the present time
only from oflF the mouth of the Rio de La Plata and from southern Brazil (San Sebastiao I.,
the vicinity of Rio de Janeiro). But it seems probable that the Lancelets that have been
recorded as B. caribaeum from Santos, from Santa Catharina at the mouth of the Amazon,
from the Rio de La Plata and from Buenos Aires, were B. flatae.

Synonyms and References:

Branchiostoma flatae Hubbs, Occ. Pap. Mus. Zool. Univ. Mich., 102, 1922: 10 (descr., off Rio de La Plata);
Jordan, Evermann and Clark, Rep. U.S. Coram. Fish. (1928), 2, 1930: 7, footnote.

Probable References:

Amfhioxus miiUeri Moreau, Bull. Acad. Roy. Belg., (2) 59, 1 875 : 3 1 2, i pL, 1 2 figs. (micr. anat. of notochord,
Rio de Janeiro, Brazil). ^-

Branchiostoma caribaeum^^ Gunther, Rep. Zool. Coll. "Alert," Brit. Mus., 1884: 32 (in part, specimen from
Botafogo, near Rio de Janeiro, Brazil) ; Jordan and Evermann, Bull. U.S. nnt. Mu?., 4.- (l), 1S96: 4 (in
part, Brazil incl. in range) ; Bull. U.S. nat. Mus., 47 (4), 1900: pi. I, fig. I (probably B. flatae because of
shape of caudal) ; Evermann and Marsh, Bull. U.S. Bur. Fish., 20 (l), 1902: 59 (ill., after Jordan and
Evermann 1900, as above, but account is of B. caribaeum, N. Carolina) ; Barbour, Bull. Mus. comp. Zool.
Harv., 46, 1905: no (in part, because Rio de La Plata incl. in distrib.) ; Ribeiro, Arch. Mus. nac. Rio
de J., 14, 1907: 146, 199; Fauna brazil., Peixes, 2(1), Fasc. I, 1923: 4 (in part, because Brazil incl. in

32. The name miilleri would have priority over flatae if the specimens in question actually were identical with
the latter, as the locality suggests. But Moreau gave no account of their external characters, nor is it likely that
the sections on which his studies of microscopic anatomy were based are still in existence.

33. Sometimes spelled "carribaeum."

1 8 Memoir Sears Foundation for Marine Research

range); Schreiner and Ribeiro, Arch. Mus. nac. Rio de J., 12, 1903: 77 (Santa Catharina, Brazil, prob-
ably B. flatae because of loc.) ; Marelli, Elenc. Sist. Fauna B. Aires (Procord. Vert.) ; Mem. Ministr.
Obras Publ. B. Aires (1922-1923), Fishes, 1924: 543 (no descr., Rio de Janeiro, Rio de La Plata,
Buenos Aires, ident. probably because of loc.) ; Luderwaldt, Rev. Mus. paul., 16, 1929: 40 (San Sebastlao
I., Brazil) ; Sawaya and Carvalho, Bol. biol. Fac. Med. S. Paulo, N.S. 2, 1938: 43 (no. of myotomes, be-
havior in aquarium, Santos, Brazil, at 2; meters).
Amfhioxos (no specific name) Luderwaldt, Rev. Mus. paul., 16, 1929: 11,15 ('" plankton, and from bottom
in shallow water, San Sebastiao L, Brazil).

Family EPIGONICHTHYIDAE

Description. Gonads developed on right side onlyj right-hand metapleuron continu-
ous with preanal fin. Characters otherwise those of the order.^*

Genera. The family includes two well defined genera: in Epgonichthys the caudal
fin does not extend as a long narrow process, and the oral tentacles are united, one to the
next, by a uniformly low intertentacular membrane; in Asymmetron the caudal fin, as well
as the notochord, is much prolonged posterior to the myotomes as a narrow process, and the
intertentacular membrane is much higher ventrally than laterally.

Key to Genera

la. Caudal fin prolonged as a narrow process; the intertentacular membrane much higher
ventrally than laterally (Fig. 3). Asymmetron Andrews, 1893, p. 18."

lb. Caudal fin not prolonged as a narrow process; the intertentacular membrane but little
higher ventrally than laterally. Efigonichthys Peters, 1876.^°

New Zealand; Australia; East Indies;
Ceylon; Maldive and Laccadive Ar-
chipelagos, to East Africa.

Genus yii'yww^/row Andrews, 1893

Asymmetron Xnix^yii, Stud. Biol. Lab. Johns Hopk. Univ., 5, 1893: 237; type species, A. lucayanum Andrews.
Bahamas.'^

Generic synonyms:

Branchiostoma (in part) Willey, Amphioxus and Ancest. Vert., 1894: 41 ; including A. lucayanum Andrews,

1893 ; not Branchiostoma Costa, 1834.
Efigonichthys Fowler, Proc. Acad. nat. Sci. Philad., 59, 1907: 461 ; including A. lucayanum Andrews, 1893;

not Efigonichthys Peters, 1876.

34. Whitley (Aust. Zool., 7, 1932: 257, 260) divides this family into Epigonichthyidae and Asymmetrontidae.

35. Including Notasymmetron Whitley (Aust. Zool., 7, 1932: 260, pi. 13, fig. 6). Whitley mentions, as characters
distinguishing this genus from Asymmetron, only that it is larger, with the origin and termination of the dorsal
fin farther forward in relation to the myotomes.

36. Including Bathyamfhioxus and Merscal-pellus Whitley, 1932. The differences on which Whitley (Aust. Zool., 7,
1932: 257—259) has separated these two new genera from Efigonichthys are so small that we hesitate to judge
their validity, not having seen specimens of them. Paramfhioxus Haeckel, 1893, is clearly a synonym of
Efigonichthys Peters, 1876, in our opinion.

37. The characters of this new genus were given also, but without a generic name, by Andrews (Johns Hopk. Univ.
Circ, 72, 1893 : 104).

Fishes of the Western North Atlantic 19

Generic Characters. Median finf old extending far beyond last myotome as a narrow
urostyloid process, with notochord reaching nearly to its tip; intertentacular membrane
much higher ventrally than laterally; ventral fin-ray chambers lacking in type species, but
perhaps present in others;'' caudal sector of median fin not demarked from more anterior
portions dorsal or ventral ; gonad pouches begin at myotomes 1 3 to 1 5 ; rostrum continuous
ventrally with both right and left sides of oral hood, and these in turn with each meta-
pleuron; atrial chamber extending behind atriopore as a pair of blind sacs; preoral tenta-
cles lack sensory papillae; no olfactory pit.

Species. The type species of the genus is A. lucayanum Andrews, 1893, of the West
Indian region and Bermuda with which A. macricaudatum Parker, 1904, of Florida is
doubtless identical (pp. 19, 22); it is also reported from the Philippines. Our examina-
tion of its type specimens leads to this same conclusion for A . orientate Parker, 1 904, of
the Maldive Islands, Indian Ocean; nor does A. caudatum Willey, 1896, from the
Louisiade Archipelago, southeast of New Guinea, appear to have any better claim to spe-
cific recognition.'"

Asymmetron lucayanum Andrews, 1893
Figure 3 A-E

Study Material. Twelve specimens, from North Bimini I., Bahamas, and from
Vieques I., Porto Rico (U. S. Nat. Mus.). Five specimens (all of them types of A. macri-
caudatum Parker, 1904) from Salt Key, Florida (Harv. Mus. Comp. Zool., No. 26282).
Seven specimens (all of them types of A. orientale Parker, 1904) from the Maldive Is.,
Indian Ocean (Harv. Mus. Comp. Zool., No. 32816).

Distinctive Characters. The long, narrow caudal process marks this species off at a
glance from all other Atlantic Lancelets, from which it differs further in the still more
important morphological respects stated above (Key, p. 7).

38. Whitley's (Aust. 2ool., 7, 1932: pi. 13, fig. 6) illustration of a specimen identified by him as caudatum Willey,
1896, and on which he based the new genus N otasymmetron, shows ventral fin-ray chambers, although he made
no mention of them in his description.

39. A. caudatum Willey (Quart. J. micr. Sci., 59, 1896: 219, pi. 13, fig. 1-4) supposedly differs from A. lucayanum
in that its rostrum is marked off by definite notches or constrictions both dorsally and ventrally. But Goldschmidt
(Biol. Bull. Wood's Hole, 64, 1933 : 323, fig. la) has recently pictured the rostrum as of this same shape for a
specimen of /I. lucayanum from Bermuda, while we have seen one from the latter locality and another from
Porto Rico with a notch on the dorsal side, although with none on the ventral side. A. orientale Parker (Bull.
Mus. comp. Zool. Harv., 46, 1904: pi. i, fig. 4) was separated from A. lucayanum on the basis of a supposedly
narrower caudal fin. But no sharp line can be drawn in this respect between its type specimens, which we have
examined, and A. lucayanum of Florida and the West Indies (Fig. 3). We may also point out that the tail
region of one specimen, a male, described by Willey (1896) as A. caudatum was what may be termed the
"lucayanum" shape, that of the other, a female, of the "orientale" shape. It is possible, however, that the Aus-
tralian form identified by Whitley (Aust. Zool., 7, 1932: 260, pi. 13, fig. 6) as caudatum, and on which he
founded the genus N otasymmetron, may represent a distinct species, in which case a new specific name would be
needed for it; he has pictured it as having ventral fin-ray chambers, although these are not mentioned in his
description of it. We may further note that ventral fin-ray chambers are also indicated in the illustration of
A. lucayanum from the Maldives, by Forster-Cooper (in Gardiner, Fauna Geol., Maldive Laccadive Archip., /,
1903: pi. 18, fig. i). But no trace of such is to be seen in the Maldive specimens that we have examined; nor are
they indicated in Franz' (Jena Z. Naturw., 5J, 1922: 426, fig. 30) figure of a Philippine specimen.

Fishes of the Western North Atlantic 21

Additional Description. Rostrum, continuous with dorsal fin, varies in shape from
very narrow both above and below the notochord to more rounded in shape, and marked
off by definite notches both dorsally and ventrallyj dorsal fin-ray chambers from 170 to
iSoj preoral tentacles 21 to 29; intertentacular membrane much higher around the ven-
tral side of oral hood than laterally, where the tentacles on either side are interconnected
only near their bases/" Median fin (dorsal and ventral), posterior to atriopore, paddle-
shaped in some specimens (wider ventrally than dorsally), narrowing rather abruptly be-
tween anus and last few myotomes; however, it is narrower in some, with a more gradual
transition to the caudal process, there being a wide range of variation in this respect, even
among specimens of a single lot, as illustrated in Fig. 3, D, E; the distance from the anus
to tip of caudal process nearly twice as great as from last myotome to anus; myotomes 42
to 46 anterior to atriopore, 8 to 9 between atriopore and anus, 1 1 to 14 posterior to anus,
total number 62 to 68 ;" gonads 26 to 29, in a single series on the right-hand side.

Color. This has not been described for living specimens.

Size. Nineteen mm. is the greatest length yet recorded for Atlantic specimens.*" If,
however, the Lancelets recorded as A . lucayanum from the Philippines are actually iden-
tical with the western Atlantic form, then the species grows larger in the Far East waters,
for lengths up to 30 mm. have been reported there.

Developmental Stages. In larvae of 6 mm., with only 22 pairs of gill openings, the
caudal extremity is expanded as a rounded fin; by the time the number of gill openings has
increased to 27 pairs it has become pointed, after which it elongates to the adult form."

Habits. This species, like other Lancelets, lives much of the time buried in the sand.
But apparently it emerges more freely to swim about, for large numbers have been taken
in tow nets at or near the surface in Bahaman waters; they are taken most abundantly dur-
ing the early part of the ebb when the tide has been high about nine o'clock in the evening;
rarely are they taken in the daytime, or late at night. In aquaria they seldom leave the
sand in the daytime. Experiments have shown them to be negatively phototropic. The pos-
terior part of the body has considerable power of regeneration if cut off just posterior to the

+0. In the original account of A. lucayanum, Andrews (Stud. Biol. Lab., Johns Hopk. Univ., Zool., 5, 1893 : pi. 13,
fig. 6) pictures the median ventral tentacle as considerably shorter than those next to it, with the membrane join-
ing it to them lower than that which joins the next three or four tentacles; Kirkaldy (Quart. J. micr. Sci., 37,
1895: 318, pi. 34, fig. 3), on the other hand, describes and pictures it as entirely frce'from the neighboring pair.
Forster-Cooper (in Gardiner, Fauna Geogr., Maldive Laccadive Archip., t, 1903: 348, fig. 76) shows the
membrane as notched where it connects with the ventro-median tentacle. But the membrane is higher there in a
specimen from that same region that we have examined; it is so described and pictured also by Franz (Jena Z.
Naturw., 5,?, 1922: 429, 430, fig. 321) for one from the Philippines. Evidently, then, the difference in this
respect is not geographic. Unfortunately, however, our West Indian series are not in good enough condition to
clarify this point.

41. Parker (Bull. Mus. comp. Zool. Harv., ^6, 1904: 48) reported only four or five between atriopore and anus
for the Florida specimens which he named A. tnacricaudatum. But re-examination of these same specimens yielded
counts of eight to nine.

42. Kirkaldy, Quart. J. micr. Sci., 57, 1895: 319.

43. Larval development is described by Andrews (Stud. Biol. Lab., Johns Hopk. Univ., 5, T893: 219, pi. 13,
fig- 3. 5)-

22 Memoir Sears Foundation for Marine Research

anus, although it is not known how far regeneration of the tail can proceed. The feeding
habits are as described for the group in general (p. 4). The time occupied by the passage
of food pellets through the digestive tract, as indicated by carmine particles, may be much
less than an hour. In Bahaman waters sexually mature specimens have been taken in June
and less often in July.**

Range. Circumtropical, with widely separated centers of distribution, and perhaps
with local races; known in the western Atlantic from Bermuda, the Florida Keys, North
and South Bimini in the Bahamas, Vieques I., Culebra I. and Humacao, Porto Rico, and
off Pernambuco, Brazil. Known also from the Maldive Islands (Indian Ocean), the
Philippines, the Louisiade Archipelago southeast of New Guinea, Zanzibar, and perhaps
from North Australia. Evidently it is abundant locally in the tropical belt of the western
Atlantic in suitable situations, for large numbers have been taken both in the Bahamas and
at Castle Harbor, Bermuda.

Synonyms and References:

1. Atlantic:

Asymmetron lucayanum Andrews, Stud. Biol. Lab., Johns Hopk. Univ., 5, 1893: 213, pi. 13, 14, fig. I-25
(descr., anat., habits, sensory reactions, Bahamas) ; Kirkaldy, Quart. J. micr. Sci., 57, 1895: 31-9, pi. 34,
fig. 3 (descr., Bahamas) ; Jordan and Evermann, Bull. U.S. nat. Mus., 47 (l), 1896: 4 (descr., Bahamas) ;
Evermann and Marsh, Bull. U.S. Fish Comm., 20 (l), 1902: 60 (descr., Porto Rico) ; Tattersall, Trans.
Lpool. Biol. Soc, 77, 1903: 291, 297, 302 (descr., discus., distrib.) ; Parker, Bull. Mus. comp. Zool.
Harv., 46, 1904: 49, pi. I, fig. 2 (myotome formula, number of gonads) ; Mark, Science, N.S. 20, 1904:
179 (Bermuda); Bean, in Shattuck, Bahama Islands, Fish., 1905: 296 (Bahamas); Barbour, Bull. Mus.
comp. Zool. Harv., 46, 1905: 1 10 (Bahamas and Bermuda) ; Lonnberg, Bronn's Klassen, 6, Abt. i, Buch
I, 1905: 244 (descr., Bahamas) ; Bean, Field Mus. Publ. Zool., 7, 1906: 29 (Bermuda) ; Gibson, Trans.
Linn. Soc. Lond., Zool., (2) 75, 1910: 241, 242 (number of myotomes; Bahamas, Amfhioxides felagicut
perhaps the neotenic larva of this species); Hubbs, Occ. Pap. Mus. Zool. Univ. Mich., 105, 1922: 16
(off Brazil) ; Franz, Jena Z. Naturw., 58, 1922: 377, 426, fig. 30-32 (descr., discus., distrib.) ; Breder,
Field Bk. Mar. Fish. Atlant. Coast, 1929: 4 (Bahamas) ; Nichols, Mem. N. Y. Acad. Sci., 10, 1929: 181,
fig. 2 (descr., Bahamas, Porto Rico) ; Goldschmidt, Biol. Bull. Wood's Hole, 64, 1933: 231, fig. I A, B
(relation to Amfhioxides larvae, Bermuda) ; Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda, 1933:
21 (descr., distrib., Bermuda); Goodrich, Quart. J. micr. Sci., 75, 1933: 723 (nephridia; Bermuda);
Pratt, Manual Common Invert. Anim., 1935: 757 (no. of myotomes and gonads) ; Andrews, Bigelow and
Morgan, Sci. Mon., 61 (5), 1945: 341, 343 (habits, ill., Bimini, Bahamas).

Branchiostoma lucayanum Willey, Amphioxus and Ancestr. of Vert., 1894: 41 (Bahamas).

Asymmetron macricaudatum Parker, Bull. Mus. comp. Zool. Harv., 46, 1904: 47, pi. 2, fig. 7 (descr., discus..
Salt Key, Florida) ; Pratt, Manual Common Invert. Anim., 1935: 757 (no. of myotomes and gonads).

Efigonichthys leucayanum Fowler, Proc. Acad. nat. Sci. Philad., sg, 1907: 461 (Bermuda).

2. Indo-Pacific:

Asymmetron caudatum Willey, Quart. J. micr. Sci., 59, 1896: 219, pi. 13, fig. 1-4 (descr., ill. of head and
tails of male and female; size; Louisiade Archipel.) ; Quart. J. micr. Sci., 44, 1901: 271 {caudatum a
subspecies of lucayanum) ; Willey's Zool. Res., 6, 1902: 725 (ill., caudatum a subspecies of lucayanum) ;
Hubbs, Occ. Pap. Mus. Zool. Univ. Mich., 105, 1922: 16 (refs.) ; Lonnberg, Bronn's Klassen, 6, Abt. I,
Buch 1, 1905: 244 (doubts if distinct from lucayanum) .

Asymmetron lucayanum Forster-Cooper, in Gardiner, Fauna Geogr., Maldive Laccadive Archip., /, 1 903:

44. The foregoing account is based on observations by Andrews, 1893.

Fishes of the Western North Atlantic 23

348, pi. 18, fig. I (descr., ill., size, Maldive Is., see footnote 39, p. 19) ; Punnett, in Gardiner, as above,
1903: 362 (number of mj'otomes, sizes) ; Tattersall, in Herdman, Rep. Govt. Ceylon Pearl Oyster Fish.,
Gulf of Manaar, suppl. 6, 1903: 222 (listed for Maldives and Zanzibar); Gibson, Trans. Linn. Soc.
Lond., Zool., (2) 13, igio: 24.1 (Maldives, by ref. to Forster-Cooper, 1903, Amfhioxides felagicus
perhaps its neotenic larva); Raff, Zool. Res. "Endeavour," Austral. Dep. Trade. Customs, I (3), 1912:
305 (listed for Louisiade Archipel., Maldives, Zanzibar, Torres Strait) ; Franz, Jena Z. Naturw., 5.?,
1922: 426, 427, fig. 30 (ill., myotome counts, size, Philippine specimens).

Asymmetron orientale Parker, Bull. Mus. comp. Zool. Harv., 46, 1904: 46, pi. I, fig. 4 (descr., ill., Maldive
Is.) ; Hubbs, Occ. Pap. Mus. Zool. Univ. Mich., 105, 1922: 16 (ref.).

Epgonichtkys caudatus Jordan and Evermann, Bull. U.S. Bur. Fish., 25, 1906: 191 (name only, Louisiade
Archipel.) ; Fowler, Mem. Bishop Mus., 10, 1928: 17 (name only, Louisiade Archipel.).

Probable References:*"

H eterofleuron {Asymmetron) lucayanum Haswcll, Rec. Aust. Mus., 7, 1908: 35 (Murray I., Torres Str.,

specimen subsequently named Notasymmetron by Whitley, 1932).
Notasymmetran caudatum Whitley, Aust. Zool., 7, 1932: 260, pi. 13, fig. 6 (descr., ill., Torres Str. spec);

Fish. Aust., /, 1940: 250, fig. 290 (N. Queensland, Murray I., Torres Str.).

Amfhioxides Larvae

Synon}-ms:

Branchiostoma (in part) Gunther, "Challenger" Rep., Zool., j/ (2), 1889: 43, for B. felagicum Gunther;

not Branchiostoma Costa, I 834.
Am.'phioxides Gill, Amer. Nat., 2g, 1895: 458; type species, Branchiostoma felagicum Gunther, 1889.
"Pelagic larvae," Forster-Cooper, in Gardiner, Fauna Flora, Maldive Laccadive Archip., 1 (4), 1903: 354,

pi. 6, fig. 3-6.
Asymmetron (in part) Pietschmann, in Kukenthal and Krumbach, Handb. Zool., 6 (i). Lief i, 1929: 110,

fig. 107, for BranchiostoTna felagicum. Gunther, 1889.

Grouf Characters. Small Lancelets, living pelagically, in which (as in larval Lance-
lets in general) the mouth is on the left side, without oral tentacles, the metapleural folds
are separate, one from another, so that there is no closed atrial cavity, and in which the gill
clefts are in a single row on the ventral side, but which grow to a greater length (up to 2 1
mm.)*' and develop a greater number of gill clefts than is usual for Lancelet larvae before
metamorphosis and which may show at least the rudiments of gonads.

As pointed out above (p. 7), these Amphioxides are juvenile specimens that re-
tain their larval characteristics not only to a greater size than is characteristic of their
parent species*" but to a more advanced stage in their own development j they are not a
primitive group as was originally supposed.*' While they may develop gonads, as just
stated, there is no evidence that Lancelets ever become mature sexually as Amphioxides.

44a. See footnote 35, p. 18.

45. Forster-Cooper (in Gardiner, Fauna Geogr., Maldive Laccadive Archip., i, 1903: 352, pi. 4) reports as
Branchiostoma felagicum a 21 -mm. specimen from the central Indian Ocean that appears to be an Amfhioxides
because no trace of oral tentacles was to be seen.

46. Larvae of this sort are known, technically, as "neotenic."

47. Their larval nature, first suggested by Goldschmidt (Zool. Anz., 50, 1906: 443) and accepted by Gibson (Trans.
Linn. Soc. Lond., Zool., [2] 13, 1910: 239), was substantiated by Goldschmidt (Dtsch. Sud-polar Exped., //,
Zool. 3, 1909: 237), who discovered Amfhioxides in which the secondary series of gill openings had begun to
form, i.e., which had commenced their metamorphosis.

24 Memoir Sears Foundation for Marine Research

Range. Amphioxides larvae have been reported from localities so generally dis-
tributed and so widely separated*' that they are to be expected anywhere on the high seas,
within the latitudinal belt where Lancelets of the family Epigonichthyidae occur in any
abundance.

Species. One specimen of Branchiostoma lanceolatum has been reported in the
Amfhioxides stage, i.e., it still retained its larval characters at a length of $•$ mm., al-
though this species usually undergoes its metamorphosis at about 4.5 mm." The other
Amphioxides larvae that have been described fall in two categories} the dorsal fin- ray
chambers of the one extend forward well beyond the first myotome, those of the other ter-
minate at the dorsal margin of the first myotome. Among the specimens of the second
group, some agree in number of myotomes with the Branchiostoma pelagicum of Gunther,
1889, and have been identified with the latter for this reason." Other specimens of the
group with a larger number of myotomes (70) for only two recorded specimens have been
described as a distinct species, Amphioxides stenurus Goldschmidt, 1905. But it is doubtful
whether the distinction between it and pelagicus is valid. The other category, with dorsal
fin-ray chambers extending far forward, has been named valdiviae Goldschmidt, 1905.

None of these has been definitely connected with any particular parent species. In the
few specimens in which the rudiments of gonads were to be seen, however, these were in a
single series and on the left-hand side, suggesting an Epigonichthys or an Asymmetron
parentage; i.e., that they belong to the family Epigonichthyidae.

Key to Species of Amphioxides

la. Dorsal fin originates opposite 21st to 25th myotome; ventral fin about opposite 40th
myotome ; dorsal fin-ray chambers do not extend forward beyond dorsal edge of i st
myotome.

2a. Not more than 68 myotomes. pelagicus Gunther, 1889, p. 25.

2b. 70 myotomes. j/(?««r«j Goldschmidt, 1905.

Indian Ocean,
lb. Dorsal fin originates opposite 32nd to 33rd myotome or even farther back; ventral fin
about opposite 43rd myotome; dorsal fin-ray chambers extend forward considerably
beyond dorsal edge of ist myotome. valdiviae Goldschmidt, 1 905, p. 27."

48. Reported from the English Channel, Bermuda, off the Amazon and at several other localities in the equatorial
and south tropical Atlantic; mouth of the Red Sea; widespread in the tropical Indian Ocean; from the vicinity
of the Hawaiian Islands. For a distribution chart, to which might be added a few more recent records, see Gold-
schmidt (Dtsch. Sud-polar Exped., //, Zool. 3, 1909: pi. 11).

49. Goldschmidt, Zool. Anz., 50, 1909: 443.

50. So identified by Goldschmidt (Wiss. Ergebn. 'Valdivia,' 12, 1905: 46). But this identification cannot be re-
garded as final until the type specimen of Gunther's felagicum is re-examined, because his illustration of it
("Challenger" Rep., Zool., j/ [2], 1889: pi. 6, fig. B) does not show the anterior termination of the fin-ray
chambers clearly.

51. Previous accounts (Goldschmidt, Wiss. Ergebn. 'Valdivia,' j2, 1905: 46, pi. i, fig. 3, 4; Gibson, Trans. Linn.
Soc. Lond., Zool., [2] 75, 1910: 217, pi. 15, fig. i) base the distinction between valdiviae and pelagicus chiefly
on the shape of the tail fin, which is supposedly more sharply marked off and blunter at the tip, with the notochord
ending more bluntly in the former than in the latter. But the two supposed species appear to intergrade in this
respect.

Fishes of the Western North Atlantic 25

Amphioxides pelagicus (Giinther), 1889

Study Material. None.

Distinctive Characters. Amphioxides larvae differ from adult Lancelets in that they
have neither atrial cavity nor oral cirri 5 their mouths are on the left-hand side, and their
gill clefts are in a single series. Pelagicus is separable from valdiviae by the facts that its
dorsal fin-ray chambers do not extend forward past the first myotome, and that the dorsal
fin originates about opposite the 2ist to 25th myotome, while in valdiviae it commences
opposite the 32nd to 33rd myotome, or even more posteriorly; pelagicus is separated from
stenurus by fewer myotomes (not more than 68 in pelagicus).

Additional Description.^'^ Caudal fin usually not sharply marked off, although some-
times more definitely so, its tip lancet-shaped, usually pointed; notochord tapering to a
narrowly pointed tip; dorsal finfold originates about opposite the 2ist to 25th myotome;
the ventral farther posterior in some (opposite the 40th myotome), but farther forward in
others; the dorsal fin-ray chambers extend forward only to the posterior edge of the first
myotome, anterior to which they are replaced by an undivided tapering canal; two cham-
bers per myotome anteriorly, increasing to 3 or 4 per myotome posteriorly; gill clefts 16
to 18 in specimens of 5 to 6 mm., with 24-26 reported for Bermuda specimens of 8 to
10 mm. or longer, and up to 30 for the Indian Ocean form; myotomes usually 63 to 64
(50 or 51 preanal and 13 postanal) with totals of 67 also reported from Bermuda, and
62 to 68 from Indian Ocean.

Color. No information available.

Size. Pelagicus has been recorded up to 16 mm. in length from Bermuda; up to lO
mm. from the Indian Ocean."

Parentage. It is probable that the pelagicus of the Atlantic is the neotenic larva of
Asymmetron lucayanum, the pelagicus of the Indian Ocean that of the local representa-
tive of lucayanum.^*

Habits. Nothing positive is known of the habits of this or of any other Amphi-
oxides, except that it is planktonic. In the Indian Ocean Amphioxides of the pelagicus
type have been taken in abundance at or near the surface and similarly at several locali-
ties in the tropical Atlantic. On the other hand, many of the records have been from
nets fished at considerable depths.'*^ In most instances, however, there is no certainty that
the specimens were actually taken at the depth at which the major part of the haul was
made, because the nets also fished while being lowered and hauled up again. Consequently,
the depth of chief abundance is still to be learned. We think it probable that the odd speci-

52. Based on previous descriptions and illustrations.

53. A 2i-nim. specimen from the Indian Ocean, reported and pictured by Forster-Cooper (in Gardiner, Fauna Geogr.,
Maldive Laccadive Archip., /, 1903 : 352, pi. 4) was in such poor condition that its specific identity is doubtful.

54. This identity has been maintained by Gibson (Trans. Linn. Soc. Lond., Zool., [2] /j, 1910: 241). Although
Goldschmidt (Biol. Bull. Wood's Hole, 64, 1933: 324) has questioned it, the number of myotomes that he
records for the Bermudian felagicus (50 to 51 preanal and 13 postanal, as well as a stated total of 67) falls
within the limits reported for lucayanum from the Bahamas (62 to 68).

55. Ostensibly down even to 1,000 fathoms (1,829 meters).

26 Memoir Sears Foundation for Marine Research

mens that have been brought up from as deep as 250 to 500 fathoms in closing nets" were
taken while in the process of sinking into the oceanic abyss, as may be the eventual fate of
all the Amfhioxides that drift out into deep water.

The frequency with which pelagkus has been reported from deep hauls makes it
likely that it can exist for a time in considerably cooler water, although it is primarily
tropical in its thermal relationships. But we have yet to learn how low a temperature may
be fatal to it, and how rapidly.

Nothing is known of its feeding habits, nor of those of any Amphioxides.

Range. Specimens showing the characters of felagicus have been reported from the
vicinity of the Hawaiian Islands, the type locality; from numerous localities distributed
across the tropical belt of the Indian Ocean between latitudes 10° 8' S. and 9° 6' N. ; from
five stations between the St. Helena and Ascension Islands and the African Coast (Lat.
about 14° S. to about 4° N.); from one station off the mouth of the Amazon; and from
the vicinity of Bermuda, whence 87 specimens were recorded from 27 townet hauls ;""
perhaps also from the Bahamas."

Synonyms and References:

Branchiostonui felagicum Gunther, "Challenger" Rep., Zool., 5 (2), 1889: 43, pi. 6, fig. B (descr., ill., N.
Pacific near Honolulu); Kirkaldy, Quart. J. micr. Sci., 57, 1895: 320 (mention); Tattersall, Trans.
Lpool. Biol. Soc, ly, 1903: 296 (distrib.) ; in Herdman, Rep. Govt. Ceylon Pearl Oyster Fish., Gulf of
Manaar, Suppl. 6, 1903: 214, plate not numbered, fig. 16 (descr., Indian Ocean) ; Lonnberg, in Bronn's
Klassen, (5, Abt. i, Buch I, 1904: 245 (ref. to type specimen) ; Franz, Jena Z. Naturw., 5<?, 1922: 433
(refs., discus., incl. valdiviae) ; Pietschmann, in Kiikenthal and Krumbach, Handb. Zool., <5 (i). Lief I,
1929: 109 (discus.).

Amfhioxides felagicus Gill, Amer. Nat., 29, 1895: 458 (name); Tattersall, Trans. Lpool. Biol. Soc., 17,
1903: 275 (diagn.) ; Goldschmidt, Wiss. Ergebn. 'Valdivia,' 12, 1905: 45, pi. I, fig. 3, 4 (descr., ill.,
Indian Ocean and trop. Atlantic); Willey, Quart. J. micr. Sci., ^o, 1906: 581 (ref. to Goldschmidt,
1905); Goldschmidt, Dtsch. Sud-polar Exped. (1901-1903), 11, Zool. j, 1909: 234, pi. 27 (discus.,
trop. Atlant., chart of distrib.) ; Gibson, Trans. Linn. Soc. Lond., Zool., (2) 13, 1910: 217, pi. 15, fig. I
(descr., discus., ill., Indian Ocean) ; Hubbs, Occ. Pap. Mus. Zool. Univ. Mich., 105, 1922: 4 (listed);
Goldschmidt, Biol. Bull. Wood's Hole, 64, 1933: 324 (meas., no. of myotomes, discus., Bermuda).

Asymmetron felagicum Pietschmann, in Kiikenthal and Krumbach, Handb. Zool., 6 (l). Lief I, 1929: 1 10,
fig. 107 (ill.).

Doubtful References:

Branchiostonui -pelagicum Forster-Cooper, in Gardiner, Fauna Geogr., Maldive Laccadive Archip., i (4),

1903: 352 (21 mm., Indian Ocean; ident. doubtful because of poor condition).
Not Branchiostoma felagicum Parker, 1 904, Bull. Mus. comp. Zool. Harv., 46, 1904: 40, pi. I, fig. I (this

was valdiviae in reality; see below, p. 28).

56. Gibson (Trans. Linn. Soc. Lond., Zool., [2] i^, 1910: 214) lists two such instances from the Indian Ocean.

57. For a list of Bermuda records, see Goldschmidt (Biol. Bull. Wood's Hole, d^, 1933: 322).

58. A six-mm. specimen from the Bahamas, pictured by Andrews (Stud. Biol. Lab., Johns Hopk. Univ., 5, 1893:
pi. 13, fig. 5) as Asymmetron lucayanum, is classed by Gibson (Trans. Linn. Soc. Lond., Zool., [2] 13, 1910:
241) as Amphioxides. But Andrews' statement (p. 219) that it had "22 branchial clefts on a side" suggests that it
was a specimen in the process of metamorphosis.

Fishes of the Western North Atlantic 27

Amphioxides valdiviae Goldschmidt, 1905
Figure 3 F

Study Material. One specimen, 9 mm. long, with 33 gill clefts, from the Maldive
Islands."

Distinctive Characters. Amphioxides larvae of the valdiviae type are separated
from those of the pelagicus-stenurus type by the following features: their dorsal fin-ray
chambers extend forward well past the first myotome, and the dorsal fin originates about
opposite the 32nd or 33rd myotome (in pelagicus about opposite the 2ist to 25th myo-
tome). Differences in the shape of the tail that have been given specific weight appear not
to be constant.

Additional Description.^" Caudal sector of fin paddle shaped with blunted tip and
rather definitely marked off from more anterior portion (dorsal and ventral) by a con-
striction, about opposite anus j notochord blunt-tipped posteriorly; dorsal finfold originates
opposite 32nd to 33rd myotome, the ventral finfold about opposite 43rd myotome; dorsal
fin-ray chambers extend forward beyond first myotome; about 5 dorsal fin-ray chambers
per myotome; gill clefts 25 to 35 in specimens of 5.7 to 8 mm., 33 to 35 in those of 8 to 9.25
mm.; myotomes SS to 58 anterior to anus, 11 to 15 posterior to it, with recorded totals
of 67 to 70.

Color. No information available.

Size. The maximum recorded length is 9.25 mm.

Parentage. If Amphioxides of this type are the neotenic larvae of species of Epi-
gonichthys, as seems probable," the parentage of valdiviae of the Atlantic presents an
interesting question, because Epigonichthys is not yet known to occur there.

Habits. Nothing is known of the thermal or bathymetric occurrence of valdiviae to
separate it from pelagicus (p. 25),

Range. Tropical Atlantic and Indian Oceans. While valdiviae has not yet been re-
ported from the western Atlantic, it is to be expected in this section of the tropical belt,
many specimens having been taken at the surface off tropical West Africa (Portuguese
Senegal), some of them showing the beginnings of metamorphosis.'" It has been reported
also off the African Coast, south of Tenerife, and at a number of localities in the tropical
Indian Ocean, including the vicinity of Sumatra, Bay of Bengal, Maldive Islands, near the
Chagos Archipelago, southeast of the Seychelles, and in the vicinity of Farquhar Islands.

59. This is the specimen described and pictured by Parker (Bull. Mus. comp. Zool. Harv., 46, 1904: 40, pi. 1, fig. i,
2). The gonads credited to it in the original account prove actually to have been the gill bars.

60. Based on descriptions by Goldschmidt (Wiss. Ergebn. 'Valdivia,' 12, 1905: 47, pi. i, fig. 1), Gibson (Trans.
Linn. Soc. Lond., Zool., [2] /j, 1910: 217), and on the specimen listed above.

61. Gibson (Trans. Linn. Soc. Lond., Zool., [2] /j, 1910: 241) suggests this parentage for Amfhioxides valdiviae
of the Indian Ocean.

62. Goldschmidt, Dtsch. Sud-polar Exped., 11, Zool. 3, 1909: tab. p. 11.

28 Memoir Sears Foundation for Marine Research

Synonyms and References:

Branchiostoma felagicum Parker, Bull. Mus. comp. Zool. Harv., ^6, 1904: 40, pi. I, fig. I (ill. showing blunt
notochord, rounded tail, and dorsal fin-ray chambers extending well beyond the first myotome; this is
clearly valdiviae; see also Study Material, p. 27, and footnote 59, p. 27); not B. felagicum Gunther,
1889.

Branchiostoma felagicum (in part) Franz, Jena Z. Naturw., 58, 1922: 434 {valdiviae incl. in synonymy).

Amfhioxides valdiviae Goldschmidt, Wiss. Ergebn. 'Valdivia,' 12, 1905: 47, pi. I, fig. I (dcscr., ill., trop.
Atlant. and trop. Indian Oceans); Gibson, Trans. Linn. Soc. Lond., Zool., (2) ij, igio: 217 (descr.,
comp. with -pelagicus) ; Goldschmidt, Dtsch. Sud-polar Exped., 11, Zool. 3, 1909: 234, pi. 27 (specimens
commencing metamorphosis, trop. Atlant., chart of distrib.) ; Hubbs, Occ. Pap. Mus. Zool. Univ. Mich.,
105, 1922: 4 (listed) ; Goldschmidt, Biol. Bull. Wood's Hole, 64, 1933: 321 (ref. to spec, showing be-
ginning of metamorphosis).

CHAPTER TWO

Cyclostomes

BY
HENRY B. BIGELOW and WILLIAM C. SCHROEDER

ACKNOWLEDGMENTS

Many persons have contributed information used in the preparation of this paper; to
all of them we are grateful. Our thanks are due in particular to A. A. Blair, Wilfrid
Templeman, A. G. Huntsman, R. A. McKenzie, Raymond T. Kinney, and Ralph
H. Osborn for assembling information as to the status of the Sea Lamprey in the rivers
of Newfoundland, the Maritime Provinces of Canada and Massachusetts; to John
Tee-Van for photostatic copies of bibliographic citations; to A. S. Romer, who assisted
in the classification; to Ludlow Griscom and James A. Peters for assistance on scientific
nomenclature; to H. W . Fowler and Leonard P. Schultz for assistance in numerous
ways, and finally to the late Thomas Barbour, who placed the collections of the
Museum of Comparative Zoology at our disposal.

GENERAL DISCUSSION

Scope of Study. The following pages give descriptions, life histories, geographic distribu-
tion so far as known, and lists of published citations for the genera and species of marine
Cyclostomes that are known to occur on the western side of the North Atlantic. The char-
acterizations of the orders, families and genera cover the Cyclostomes as a whole, as does
the key to the species of the only genus in question that includes more than a single marine
representative.

Descriptions. These are based on the Study Material listed under each species. The ac-
counts of the habits and geographic distribution are taken partly from the published rec-
ords, partly from data of our Study Material, and partly on information from numerous
correspondents, supplemented by our own observations.

Keys. The keys, from the higher groups down to species, have been prepared solely

29

30 Memoir Sears Foundation for Marine Research

for ease of identifying any Cyclostome that may come to hand; for that reason we have,
selected characters which are most easily used.

References. All citations listed among the references, with the few exceptions noted,
were consulted in the original ; for a list of co-operating libraries, see the general discus-
sion for the section on Sharks.

Sources of Material. The collection of Cydostomes in the Museum of Comparative
Zoology has been the chief basis of our studies, but the collections at the United States
National Museum and the Academy of Natural Sciences at Philadaelphia have also been
drawn upon.

Class AGN AT HA
Subclass CYCLOSTOMATA

Fish or fish-like vertebrates,^ eel-like in form, the skeleton cartilaginous or fibrous,
without bone; no definitely developed jaws or bony teeth; at least the rudiments of a
cranium present in the form of a simple trough below the brain in some, but partially
roofed in others; notochord not constricted at all segmentally; vertebral column repre-
sented by a simple notochordal sheath, without vertebral centra, but with rudimentary
neural arches (not joined above) in some; no shoulder or pelvic girdles, no paired limbs
and no true ribs; 6 to 14 pairs of gill pouches opening either directly into the pharynx
internally or into a separate respiratory tube, which in turn opens into the mouth below
the gullet, and opening to the exterior either separately or by a single aperture on each
side; skin without scales; nostril single, either opening into the mouth or not; intestine
with internal longitudinal ridges, or with a slight spiral fold; ear with either i or 2 semi-
circular canals only; no sympathetic nervous system, or spleen; heart without conus arte-
riosus; no cloaca, the genital apertures being separate from anus. Development oviparous,
with or without a definite larval stage ; the sexes separate or not.

The Cydostomes are generally considered the most primitive of true vertebrates;
structurally they are the simplest. They are easily distinguishable from all the higher
fishes by their peculiar jawless mouths, by the fact that there is only one nostril, and by
the very primitive cranium.^

Key to Orders

I a. Snout with prominent barbels; no separate dorsal fin; eyes not visible externally;

nasal opening at tip of snout; mouth not funnel- or disc-like. Myxinoidea, p. 31.

lb. Snout without prominent barbels; one or more dorsal fins separate from caudal; eyes

in adult well developed, and visible externally; nasal opening on upper side of head;

mouth opens as a funnel or disc. Petromyzonida, p. 43.

1. Opinions differ as to whether the Cydostomes are to be regarded as a class distinct from the true fishes, or as a
subclass of the latter.

2. For detailed accounts of the anatomy of the Cydostomes, see especially Lonnberg, Favaro, Mozejko and Rauther
in Bronn's Klassen, 6, Abt. i, Buch i, 1905-1924: 16-39, pl- ''3~3^> ^1*° Pietschmann, in Kukenthal, Handb.
Zool., 6, 1st half, 1929— 1935: 2-5.

Fishes of the Western North Atlantic 31

Order MYXINOIDEA

Description. Six to 15 pairs of gill pouches, opening internally into the pharynx,
those on each side opening either separately to the exterior or by a single common aper-
ture} 2 pairs of barbels on side of nostril and i or 2 pairs at side of mouth; single con-
tinuous fin running posteriorly around tail and anteriorly on lower surface; fin rays
restricted to tail region; nostril at tip of snout opening into mouth and serving as the
entrance for water in respiration; mouth not funnel-like; tongue evertible, with two rows
of horny, rasp-like teeth; prominent row of segmentally arranged mucous pores along
each side; anus near posterior end; eye, without lens or iris, not visible externally, and
apparently degenerate; cranium a simple, unroofed trough below brain; barbels and
tongue supported by cartilaginous bars; branchial basket reduced to a vestige; ear with
one semicircular canal only; a pancreas-like gland well developed; notochordal sheath
without rudimentary neural arches; intestine with internal longitudinal folds, but without
spiral valve.

Development. According to recent studies (see discussion and footnote 14, p. 35) the
myxinoids, although structurally hermaphroditic, are not functionally so. Development
is direct, without a larval stage."

Habitat. Exclusively marine.

Families. Only one, Myxinidae, is known.

Family MYXINIDAE

Hags

Characters. Those of the order.

Discussion of Genera. The members of the family fall in two sharply alternative
groups, depending on whether the gill pouches of each side open to the exterior by a single
common orifice, or separately. By common consent, members of the first group fall in one
genus, Myxine. But the members of the second group have been divided, depending on
the importance given by diflFerent students of classification to the number of gills and the
grouping of their openings. Since none of the latter group occur in the western North
Atlantic we need only point out that the use of the number of gills for generic separation
does not seem permissible, for species occur with 5, 6 to 7, 8, lO, 1 1 to 12, and 14. But
the difference between the close grouping of the gill openings in Paramyxine, and their
wide spacing in all the others, does seem worth generic recognition, as indicated in the
following key.*

3. Dean's detailed description of the early development of Eftatretus stoutl, in "Festschrift von Kupfer's" (1899:
221—277, pis. 15—26) has formed the basis for subsequent accounts in many textbooks.

4. Holly (in Schultze, Kiikenthal, et al., Tierreich, Lief 59, 1933 : 45) includes the shape of the gill openings as an
additional generic character, but Matsubara (J. Imp. Fish. Inst. Tokyo, 32 [1], 1937: 13) has recently shown
that this varies so widely in Paramyxine as not to be reliable.

32 Memoir Sears Foundation for Marine Research

Key to Genera

I a. Gill pouches on each side connect with exterior by single common aperture.

Myxine Linnaeus, 1758, p. 32.
Atlantic and Pacific Oceans.

lb. All gill pouches on each side open independently to exterior.

2a. Gill openings on each side ( 1 6 in number) are close together.

Paramyxine Dean, 1904.

Japan.

2b. Gill openings on each side ( 5-14 in number) separated by interspaces of consider-
able width. Eftatretus Cloquet, 1819.°

Pacific Ocean.

Genus Myxine Linnaeus, 1758
Hags

Myxine Linnaeus, Syst. Nat., 1758: 650; type species, M. glutinosa Linnaeus. Atlantic Ocean.

Generic Synonyms:

Petromyzon (in part) Walbaum, P. Artedi Genera Pise. Emend. Ichthyol., Pt. 3, 1792: 500, for M. glutinosa;
not Petromyzon Linnaeus, 1758.

GaUrobranchu! Bloch, Naturg. Ausland. Fische, Pt. 9, 1793: 66, pi. 413; type species, G. coecus Bloch.
Denmark, Sweden, Norway and Iceland.

Muraenoblenna Lacepede, Hist. Nat. Poiss., 5, 1803: 652; type species, M. olivacea Lacepede. Straits of Ma-
gellan.

Pholis Oken, Lehrb. Naturg., 5 (2), 1816: 122; alternative name for Myxine.

Gasteobranchus Buckland, Nat. Hist. Brit. Fish., 1 881: 144; evident misspelling for Gastrobranchus.

Generic Characters. Five or 6 gill pouches on each side opening to exterior by a single
aperture on ventral surface, close in front of origin of ventral finf old, the left-hand gill
opening, which receives the oesophago-cutaneous duct, being much the larger; fleshy flap
("rostrum" or "labrum") overhanging nostril anteriorly; nostril close to tip of snout;
snout with 6 barbels, flanking both nostril and mouth ; each side with a series of large
mucous pores, segmentally arranged, extending from a short distance behind the mouth
rearward nearly to the caudal extremity. Characters otherwise those of the family and
order.*

Range. Continental shelves and slopes of the North Atlantic in north temperate and

5. This genus has been called most commonly Bdellostoma Miiller, 1835. It has been argued by Apstein (Sitzber.
Gesellsch. Naturf. Berlin, 1915: 187) and Rauther (in Bronn's Klassen, 6, Abt. i, Buch i, Lief 39, 1924: 685)
that it would be well to accept this as a nomen conservandum. But, awaiting action by the International Commis-
sion on Zoological Nomenclature, it seems to us wiser to use the older name. As Rauther points out, the original
description of Eftatretus was based by Cloquet (Diet. Sci. Nat., 1$, 1819: 135) on a combination of two species —
the Chilean dombei and an unnamed species from the South Seas. But even if Eftatretus were to be abandoned as a
generic name on that account, Bdellostoma is long antedated by Homea Fleming (Phil. Zool., 2, 1822: 374) and
by M'Murtrie's Heftatremus (Anim. Kingd. [after Cuvier], 2, 1831: 298).

6. For an excellent account of the general morphology of Myxine, see Smitt (Hist. Scand. Fish., 2, 1895 : 1 196).

Fishes of the Western North Atlantic 33

subarctic latitudes, including the Mediterranean (Adriatic) in moderate depths; coasts of
southern Argentina, Chile, Japan, and South Africa; Gulf of Panama in deep water (1,335
meters), the latter being the only locality where the genus is known to occur in tropical or
subtropical latitudes.

Species. The representatives of the genus fall in two well defined groups, according
to whether the first three lingual teeth of the anterior row are fused at the base, or only the
first two, which is more usual. One member of the first of these groups, M. circifrons, is
further set apart from all others in the genus by the fact that it has only five pairs of gill
sacs. Unfortunately this feature is not apparent from the exterior, and other characters
that have been used to separate supposed species, such as relative length of head and
number of mucous pores, overlap to such an extent that it is doubtful how many of the
named forms will finally stand. For further discussion, see p. 38.

Key to Species of Myxine

I a. First 3 lingual teeth in anterior series fused together at base.

2a. Lingual teeth j| ; head nearly or quite 33.3 % of total length.

circifrons Ga.Tma.n, 1899.'
Gulf of Panama.

2b. Lingual teeth onlyi^ or fewer; head not more than 29% of total length.

3a. 26 or 27 mucous pores anterior to gill openings, and 12 or 13 posterior to
anus. garmani Jordan and Snyder, 1 90 1 .

Japan.

3b. Only 22 mucous pores anterior to gill openings and 9 posterior to anus.

tridentiger Gzrmzn, 1899.
Straits of Magellan,
lb. Only 1st 2 lingual teeth in anterior row fused together at base.

4a. Lingual teeth only | . faucidens Regan, 1 9 1 3.

Japan.
4b. Lingual teeth ^ or more.

5a. 10- 1 1 teeth in anterior series in adult. a finis Giinther, 1 870.

Straits of Magellan.

5b. Not more than 7-9 teeth in anterior series in adult.

glutinosa Linnaeus, 1758, p. 34.*
Both sides of North Atlantic.

7. This species is set apart from all others of the genus by the fact that it has only five pairs of gill sacs. This, how-
ever, is not apparent externally.

8. Including cafensis Regan, 1913, South Africa, and australis Jenyns, 1842, Chile and southern Argentinaj these
species and glutinosa so overlap one another in the' number of teeth and mucous pores and in the relative length
of head that we have not been able to construct a key by which individual specimens could be identified with cer-
tainty. Neither can the presence of seven pairs of gill pouches in cafensis be regarded as a unique specific charac-
ter, since occasional specimens of glutinosa may have this same number (footnote 11, p. 35). Information on
the number of teeth ( jj ) and gill pouches of cafensis, which was not included in the original description of
the species (Regan, Ann. Mag. nat Hist., [8] n, 1913: 398), has been obtained subsequently (Barnard, Ann.
S. Afr. Mus., 31 [i], 1925: 15).

34 Memoir Sears Foundation for Marine Research

Myxine glutinosa Linnaeus, 1758
Hagfish
Figure 4

Study Material. Forty-seven specimens of various sizes up to 610 mm. in length,
from the Grand Banks and localities on both sides of the Gulf of Maine, north slope of
Georges Bank, outer part of the continental shelf off Nantucket Island and off Cape Look-
out. Also 13 specimens from the eastern Atlantic — Norway, Denmark, Kattegat, the Adri-
atic and Liverpool, England.

Distinctive Characters. The combination of jawless mouth, single nasal aperture,
only a single pair of external gill openings, no operculum or covering fold of skin, worm-
like form and lack of paired fins separate the Hag from all other fish-like vertebrates of
the western North Atlantic.

Description. Trunk cylindrical throughout most of its length, its diameter about V24
to ^25 of its total length, tapering rearward from dorsal origin of finfold to narrowly

Figure 4. A Myxine glutinosa, specimen 380 mm. long, from the Gulf of Maine. B Oral view of anterior
part of head of same. C Lingual teeth of same viewed from above, about 3 x. Z) Egg after being laid, after
Dean, about 2 x.

Fishes of the Western North Atlantic 35

rounded caudal extremity 5 a segmentally arranged row of mucous pores low down on
each side, extending from about Vis the way back from snout to beyond anus; 26 to 33
pores in front of gill openings, 57 to 66 between gill openings and anus in those seen (53
to 70 recorded), and 11 to 13 posterior to anus in 9 specimens examined from Grand
Manan Island, New Brunswick."

Length of head to gill openings about 25 to 29% of total length (3.4 to 4 inches total
length); snout obliquely truncate; fleshy rostrum a little higher than wide and broadly
rounded in well preserved specimens, but sometimes more narrowly pointed, possibly due
to contraction in the preservative ; nostril an open pore on ventral surface near tip of snout;
2 pairs of slender, flexible barbels flanking either side of nostril, with a third pair, about
twice as large, flanking the anterior part of mouth; mouth irregularly stellate when closed,
without definite lip, but with a prominent, conical projection on either side of its margin;^"
gill openings close in front of origin of ventral finfold; usually 6 pairs of gill pouches, not
visible externally, but sometimes 7 pairs.^^

Lingual teeth comb-like, with swollen bases and sharp tips, moderately curved rear-
ward, close together, decreasing in size from front to rear, of a strong orange color; those
of anterior series about twice as large as those of posterior series, and partially overlapping
the latter when tongue is retracted within mouth; 7 to 9 on either side in the anterior series
and 8 to 10 in the posterior series; the first 2 in each series fused together at the base.

Ventral finfold originates about Vs of distance back from snout to caudal extremity,
the dorsal fin about % the distance back and slightly anterior to anus," both fins about V^
to V4 as high as the trunk is deep ; ventral fold unsupported anterior to anus, but posterior
to the latter it has a series of very slender, tapering cartilaginous rods, which extend around
caudal extremity and forward along dorsal finfold (decreasing in length) nearly or quite
to origin of latter.

Color. Grayish or reddish brown above, either plain, variously suffused, or mottled,
with darker or paler gray, brown or bluish ; whitish, or pale gray below. The variations in
color may correspond more or less closely with the local color of the sea bottom.

Size. In American waters, on the coast of Maine, Hags are recorded up to 790 mm.
in length, with one series of adults averaging 620 mm." Apparently this is a greater size
than they reach on the opposite side of the Atlantic, where the maximum recorded length
is only 420 mm. (see discussion, p. 38).

Developmental Stages. The Hag was at first believed to be a functional protandrous
hermaphrodite, its single unpaired sex organ first developing sperm in the posterior por-
tion, then eggs later in the anterior portion.^* However, recent detailed studies of the sex

9. 24-34, 54-64 and 10-14 respectively are recorded for European specimens.

10. These projections have sometimes been interpreted as a fourth pair of barbels.

n. Specimens with seven gill pouches on one side, or on both, are recorded by Cole (Anat. Anz., 2y, 1905: 3»6).

12. Its origin is not clear-cut. The first indication of it is nearly as far anterior to the anus as the latter is distant
from the tip of tail, in both American and Norwegian specimens.

13. Conel, J. Morph., 29, 1917: 77.

14. For a summary of earlier studies, see Smitt (Hist. Scand. Fish., 2, 1895 : 1205) and Conel (Dean Memor. Vol.,
Amer. Mus. nat Hist., Art. 3, 1931 : 70).

36 Metiwir Scars Foundation for Marine Research

organ" appear to show that this is not the case; either the male portion of the common sex
organ matures in each individual, with the female portion remaining rudimentary, or vice
versa. It has long been known that the eggs are few in number (only 19 to 30 having been
counted in any one female) and large (up to 25 mm. in length), the horny shell with a
cluster of anchor-tipped filaments at each end very characteristic in appearance.'^ But it
was not until 1900 that any were found which had been laid naturally." The eggs are
deposited on the bottom, where they stick firmly in clusters to some fixed object'' by means
of their filaments and by threads of slime. The newly hatched Hag has not been seen as
yet, but inasmuch as the smallest described, which is about two and one-half inches long
and probably not long out of the egg, resembled the adult, there is no reason to suppose
that the Hag passes through a larval stage.

Habits. The Hag is found chiefly, if not exclusively, where the bottom is soft mud
or clay; its actions in aquaria'" suggest that it spends most of its time imbedded in the clay
or mud, with only the tip of its snout and the nasal barbels projecting, although it swims
actively by an undulating motion in the horizontal plane when disturbed or when aroused
by food in the vicinity; it is most active in the dark. Its depth range is considerable, extend-
ing commonly from 15 to 20 fathoms down to 250 fathoms or so, and it has been taken
as deep as 524 fathoms.^" The fact that it seldom, if ever, attacks hooked or netted fishes
unless they are close to the bottom suggests that it never rises much above the latter.

In aquaria Hags die soon if the salinity is as low as 2.0 to 2.5 per cent;"' survive for
some weeks but do not feed if it is 2.9 to 3.1 per cent;" feed and thrive if it is as high as
3,2 to 3.4 per cent."' Also, it appears to be rather definitely limited in its dispersal toward
the surface by high temperature, since it is rarely if ever found in water warmer than about
50 to S5 degrees, which in all but the most northerly part of its range would confine it to
depths of 1 5 to 20 fathoms or more, except in the cold season. On the other hand, polar
temperatures are probably a barrier to its northward dispersal (p. 40).

By its preference for soft bottom, comparatively high salinity (p. 37) and low
temperature (see above), the Hag is confined within its area of regular occurrence to the
deeper furrows and troughs on the Nova Scotian slope and in the Gulf of Maine, to the
outer parts of the deeper bays, such as Fundy, Passamaquoddy, Massachusetts and prob-

15. Schreiner (Biol. Zbl., 2^, 1904: 91-104, 121— 159, 162-173); Schreiner and Schreiner (Arch. Biol., 11, 1905 :
183-3 14, 8 pis., 3 15-355, 2 pis.; Arch. ZcUforsch., /, 1908: 152); Conel (J. Morph., 29, 1917: 75-163, 12
pis.; Dean Memor. Vol., .4mer. Mus. nat. Hist., Art. 3, 1931 : 70).

16. For reference to early accounts of eggs, see Smitt (Hist. Scand. Fish., 2, 1895: 1206).

17. Dean, Mem. N. Y. Acad. Sci., 2 (2), 1900: 34, pi. 2.

18. To a Bryozoan in one case; see Jensen (Vidensk. Meddel. dansk. Naturhist. Foren., Copenhagen, 1900: i).

19. For an interesting account of the habits of the Hag in aquaria, see Gustafson (Arkiv. f. Zoologi, Stockholm,
28A [2], 1935).

20. Southeast slope of Georges Bank, Lat. 41° 32' N., Long. 65° 55' W. (Goode and Bean, Smithson. Contr.
Knowl., 30, 1895 : 3; Spec. Bull. U.S. nat. Mus., 1895 ; Mem. Harv. Mus. comp. Zool., 22, 1896).

21. Gustafson (Arkiv. f. Zoologi, Stockholm, 2SA [2], 1935), in western Sweden.

22. This is the usual summer range for surface water in Passamaquoddy Bay, where Hags were kept in captivity
by Coonfield (Trans. Amer. micr. Soc, 59, 1940: 398).

Fishes of the Western North Atlantic 37

ably Penobscot, and offshore on the continental slope to the zone deeper than about 100
fathoms.

The Hag is not a parasite, as has sometimes been suggested, there being no reason
to believe that it ever attacks living, uninjured fish. But it is a scavenger, feeding largely
on dead or disabled fish of any sort, into which it bores by means of its rasp-like tongue.
It is best known for its habit of penetrating the body cavities of hooked or gilled fishes,
eating out first the intestines and then the meat, leaving nothing but a bag of skin and bones,
inside of which the Hag itself is often hauled on board} or it may be captured clinging to
the side of a fish it has just attacked. In Norwegian waters as many as six Hags have been
reported in a single haddock." It is also known to prey on marine polychaete worms, at
least in Norwegian waters, and it has been suggested that these may be its normal diet.^*

Being blind, the Hag evidently finds its food by scent, and so successfully that large
numbers are sometimes taken in pots baited with dead fish or other offal j a local instance
is mentioned below.

The fact that the eggs of the Hag have been found off southern Newfoundland at
the mouth of the Bay of Fundy and on Georges Bank on one side of the Atlantic, and on the
other side, near the Faroes, in Norwegian waters and off Morocco, shows that it spawns
throughout its range; also, it spawns throughout the year, for females nearing ripeness,
and others nearly spent, have been recorded for various months, winter and spring, as well
as summer and autumn j in Norwegian waters eggs have been taken from November to
May. The few eggs so far reported have been from depths of 50 to 150 fathoms, and most
of them have been trawled on mud, clay, or sandy bottoms.""

Numerical A bundance. In American waters the Hag has usually been noted as being
not very common. Actually it occurs in very considerable numbers on suitable mud bottoms
at the appropriate depths, though rarely elsewhere, if at all. Thus, in the spring of 19 13
the Hag was so plentiful on the Boon Island-Isles of Shoals fishing grounds that three
to five per cent of all the haddock that we saw taken in gill nets had been attacked by
them. Similarly, fishermen report that in certain areas of soft bottom in the northern part
of the Gulf of Maine they damage a large proportion of the fish caught on long lines,
unless the latter are tended frequently. The vicinity of Grand Manan Island at the mouth
of the Bay of Fundy, and the trough with mud bottom between Jeffrey's Ledge and the
coastline on the western side of the Gulf of Maine, are centers of abundance with which
local fishermen have long been familiar. And evidently they are plentiful locally on the
upper part of the continental slope off southern New England as well, for we took 1 1 large
ones in an hour or less with one set of the Monaco trap off Nantucket at 260 fathoms on
July 8, 1908. But we question whether they ever occur in American waters in such num-
bers as in the fjords of western Sweden and southern Norway, where catches of 100 are

23. S. Nilsson, Prod. Ichthyol. Skand., 1831: 124.

24. Gustafson, Arkiv. f. Zoologi, Stockholm, 28A (2), 1935.

25. Hjort, Rep. Norweg. Fish. Invest, / (i), 1900: 75.

38 Memoir Sears Foundation for Marine Research

usual in eel pots set overnight on suitable bottom, with 1,400 recorded as captured in one
set of 24 hours.^°

Relation to Other Sfecies. The American form has been considered specifically dis-
tinct from the European by some authors {M. limosa Girard, 1859), but not by others.
However, the American form falls well within the limits of the European M. glutinosa
in numbers of lingual teeth and slime pores. Its rostrum is also of the same obtuse shape
in the better preserved specimens we have examined, although it has been pictured as more
acutely pointed in some.^' Nor has our own comparison of specimens from the two sides
of the Atlantic revealed any significant differences in other respects. While the American
form may grow larger than the European (p. 35), we hesitate to use size as a basis for
specific separation unless accompanied by other differences of a sort that could allow any
given individual to be referred to the one species rather than to the other. M. atlantica
Regan, taken off Nova Scotia, seems also clearly referable to glutinosa.

The relationship of glutinosa of the northern hemsiphere to australis, affinis and
capensis of the southern hemisphere is not so clear, but is a question of interest from the
standpoint of geographical distribution. The only clear-cut difference between capensis on
the one hand and the australis-ajfinis group on the other (the former overlaps the latter in
number of teeth and slime pores) is that capensis is described as having seven gill pouches
while there are only six in australis and affinis. However, we doubt whether or not this
apparent difference is of specific importance, for while in glutinosa the usual number is
six, seven also have been recorded (p. 35).

According to Norman's'* recent comparison of australis with affinis, the number of
teeth is less and the average number of abdominal slime pores is smaller in the former
(8 teeth in first series, 8 or 9 in second; s^ to 64 abdominal pores) than in the latter [ 10 or
II (9 in young) teeth in first series, 9-1 1 in second; 63 to 69 abdominal pores] ; and its
rostrum or labrum is longer and more acutely pointed. But this last character, being some-
what variable in glutinosa, may be equally so in the southern hemisphere forms. How-
ever, although the number of pores overlap in the two species, it appears that individual
specimens can be referred to the one or the other, depending on the number of teeth. The
large number of teeth in its anterior series also appears to mark affinis apart from glutinosa
(7-9), although it overlaps the latter in the number of teeth in the posterior series, and
falls within the range of variation recorded for glutinosa in the number of abdominal
pores; however, australis, by Norman's definition, falls within the limits recorded for
glutinosa, both in numbers of teeth and in numbers of pores. Neither have we been able to
separate individual specimens of the one from those of the other by shape of rostrum. But
since none of the considerable series of australis that we have examined are in good condi-
tion, we hesitate to unite the two species, in view of their widely separated areas of dis-
tribution.

26. Lyngnes, Z. Wiss. Biol., Abt. A, Z. Morph. Okol., /p, 1930: 591.

27. Garman, Mem. Harv. Mus. comp. Zool., 2^, 1899: pi. 68, fig. 7.

28. "Discovery" Rep., 16, 1937: 4i see this publication also for the somewhat confused synonymy of the two.

Fishes of the Western North Atlantic 39

Relation to Man. The Hag, being of no value itself, is only a nuisance to the fisher-
men because of its habit of damaging better fish, and a loathsome one, owing to its ability
to discharge slime from its mucous sacs out of all proportion to its size. One Hag, it is said,
can fill a two gallon bucket, and we think this no exaggeration.^'

In American waters the commercial fishes most often damaged by it are the haddock
and the hakes (Urophycis), these being the species most often fished for with long lines
or with gill nets over the particular type of bottom that the Hag frequents. But it some-
times damages cod also, and European authors describe it as attacking ling (Molva) and
other gadoids, herring, mackerel, sturgeon, and even mackerel sharks (Isurus) under
similar circumstances.

Range. Both sides of the northern North Atlantic. In the eastern North Atlantic it
occurs on the Murman coast and in northern Norway'" southward in abundance to the
northern part of the North Sea, the Kattegat (not known from the Baltic) and the Irish
Seaj less commonly to the English Channel (Cornwall) ; occasionally to Portugal. There
are two records of it off Morocco, one just outside the Straits of Gibralter,*^ the other
just inside in the Mediterranean."' It has been credited to the Adriatic" also, no doubt on
the strength of the fact, reported by Garman,'* that there are three specimens labelled
"Trieste" in the collection of the Museum of Comparative Zoology (see Study Material,
p. 34). But so far as we can learn it is not included otherwise in any of the general surveys
of Mediterranean fishes°° that have appeared. This makes it much more probable that the
specimens in question were mislabelled, and that Myxine is actually not a regular member
of the fauna of the inner parts of the Mediterranean.

On the western side of the Atlantic it occurs at least occasionally as far north as the
northern part of Davis Strait (see p. 40), and southward as far as the latitude of Cape
Fear in North Carolina. It is represented in the corresponding thermal belt in the southern
hemisphere (Chile, southern Argentina, Straits of Magellan, Tierra del Fuego, South
Africa) by a form, or forms, so closely allied that it is doubtful whether any sharp line can
be drawn between them (see discussion, p. 33).

Occurrence in the Western Atlantic. While not known for certain along the west
coast of Greenland," so far as we can learn, the Hag has been taken on one occasion in the

29. Linnaeus (Sys. Nat., 1758: 650), referring to this habit, wrote "aquam in glutem mutat."

30. Apparently it does not occur around Iceland, for it is not included by Saemundsson (Skr. Komm. Havunders.
Kbh., No. 5, 1900) in his survey of Icelandic fishes.

31. Eggs; Koefoed (Rep. Sars N. Atlantic Deep Sea Exped., 2k>ol., 4 [1], 1927: 18).

32. Roule, Result. Camp. sci. Monaco, 52, 1919: 129.

33. Schnakenbeck in Grimpe and Wagler, Tierwelt N- u. Ostsee, Lief 7, Teil izd, 1927: 3; Cons, explor. Mer.,
Faune Ichthyol. N. Atlant., 1931.

34. Mem. Harv. Mus. comp. Zool., 34, 1899: 348.

35. MuUer (Vergl. Anat. Myxinoiden, Pt. i, 1835: 17, footnote) long ago rejected Bloch's (Schr. Ges. Naturf.
Freunde Berlin, 10, 1792: 251) suggestion that Myxine is in the Mediterranean, which was based on Aristotle's
account of the slime-producing habit of his Pholis.

36. It has been credited repeatedly to Greenland on the strength of Fabricius' (Fauna Groenl., 1780: 344) charac-
terization of it as "rari in mari Groenlandico." But we find no other record of it among the many subsequent
lists of fishes of Greenland, east or west, except as noted above.

40 Memoir Sears Foundation for Marine Research

northern part of Davis Strait, just south of the Greenland-Baffin Land Ridge." But there
is no report of it either in the region of Hudson Bay, along the Atlantic coast of Labrador,
or on the east coast of Newfoundland j nor did any of the many cod that we saw caught
by hook and line or nets in the summer of 1900 along the outer Labrador coast show any
evidence of attack by Hags. Apart from the Davis Strait record just mentioned, the most
northerly known stations for it on the American coast are the Grand Banks and the south
coast of Newfoundland, where its eggs have been trawled."' Type of bottom, temperature
and salinity are such that it is also to be expected in the deep trough of the Gulf of St. Law-
rence, though we found no definite record of it there.

To the southward it is generally distributed at appropriate depths wherever the bot-
tom is suitable: over the continental shelf and down the continental slope along Nova
Scotia, throughout the Gulf of Maine, along the seaward slope of Georges Bank, and off
southern New England and New York, where specimens have been taken at many locali-
ties by trawl or otherwise, at depths of 100 to 250 fathoms and deeper. Apparently this
marks the limit of its common occurrence in this direction, however, for the only records
of its occurrence south of the latitude of New York are: one specimen taken off Delaware
Bay in 1 26 fathoms, and one or more in 1 78 fathoms off Cape Fear, North Carolina, many
years ago,'"

Synonyms and References:*"

Myxine glutinosa Linnaeus, Syst. Nat., 1758: 650 (Atlant. Oc, grouped among the worms) ; Muller, O. F.,
Prod. Fauna Danica, 1776: 227 (Denmark) ; Pennant, Brit. ZooL, 4, 1777: 39, pi. 20, fig. 5 (habits, ill.) ;
Fabricius, Fauna Groenl., 1780: 344 ("rari in mari Groenlandico") ; Retzius, Fauna Sueciae, I, 1780:
302 (refs., habits, west, seas); Gmelin, Syst. Nat., i (6), 1790-1791: 3082 (descr., Atlant. Oc.) ; Ret-
zius, Svenska. Vet. Akad. Handl., 11, 1790: I lo, pi. 4 {Myxine and Petromyzo?i considered more worm-
like than fish-like; plate referred to is not in copy seen) ; Abildgaard, Schr. Ges. naturf. Freunde, Berlin,
10, 1792: 193, 244, pi. 4 (descr., ill.; a fish, not a worm) ; Bloch, Schr. Ges. naturf. Freunde, Berlin, 10,
1792: 244 (disc, of earlier accounts; believed same as Pholis of Aristotle, therefore in Mediterranean,
Greece); Fleming, Hist. Brit. Anim., 1828: 164 (descr., habits, England); Cuvier, Regne Anim.,
2nd Ed., 1829: 406 (North Sea); Nilsson, Prod. Ichthyol. Skand., 1832: 123 (habits, type of bottom,
feeding, north. Norway) ; Johnston, London's Mag. Nat. Hist., (5, 1833: 15 (Scotland) ; Jenyns, Manual
Brit. Vert. Anim., /, 1835: 413 (Ireland); Muller, J., Vergl. Anat. Myxinoiden, i, 1835: 3 (history),
15 (class., diag.. North Sea, Norway, Sweden, Greenland), 17 (footnote considers Bloch's reference of it
to Mediterranean on basis of Aristotle incorrect) ; Templeton, Charlesworth's Mag. Nat. Hist., 1, I 837:
413 (Ireland) ; Cuvier, Regne Anim., Poiss., 183 8-1 843: 383, pi. 120, fig. 3 (ill.) ; Kr0yer, Danmarks
Fisk., _5, 1838-1853: 1068 (descr., habits, Denmark); Swainson, Nat. Hist. Fish. Amphib. Rept., 2,
1839: 338; Fries, Ekstrom and Sundevall, Skand. Fisk., 6, 1840: 121, pi. 28 (descr., ill., Scandinavia);
Gray, List Fish. Brit. Mus., Chondropt., i, 1851: 147 (Norw.iy, Gt. Brit.); White, List Spec. Brit.
Mus., Fish., 8, 1851: 145 (north. England, Scotland); Nilsson, Skad. Fauna Fisk., 4, 1855: 750 (not
seen) ; Thompson, M., Nat. Hist. Ireland, 4, 1856: 267 (Ireland, Scotland) ; Thomson, .■\., Art. "Ovum,"
in Todd's Cyclop. Anat. Physiol., 5 (suppl. vol.), 1859: 50, fig. 33, c, d (earliest descr. of egg, ill.);

37. Lat. 66° 37' N., 450 meters, temp. 3.12° C; Jensen, Rapp. Cons, explor. Mer., ^9, 1926: 98.

38. Dean, Mem. N. Y. Acad. Sci., 2 (2), 1900: 34.

39. One specimen from this lot is in the Harv. Mus. Comp. Zool.

40. Myxine, as representative of its order (subclass in some schemes of classification), has been the subject of many
anatomical accounts and discussions, in reports of original observations as well as in general textbooks, etc. This
list is confined to such citations as bear directly on its classification, on its habits or on its distribution.

Fishes of the Western North Atlantic 41

Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861 : 63 (Polar regions to Cape Cod) ; Steenstrup, Overs,
danske Vidensk.-Selsk. Forh. (1863), 1864: 233 (eggs); Gunther, Cat. Fish. Brit. Mus., 8, 1870: 511
(descr., coasts of Europe and N. ."Xmer.) ; Gill, Rep. U.S. Comm. Fish. (1871-72), 1873: 814 (listed
Greenland, Polar regions to Cape Cod, Mass.) ; Putnam, Proc. Boston Soc. nat. Hist., 1873: 135 (dimens.
and no. of teeth of Grand Manan specimens); Collett, Vidensk.-Selsk. Forh. Christiania, 1874; also
Norges Fisk., 1 875 : 220 (habits, distrib., depth, Norway) ; Liitken, Cat. Fish. Greenl., in Manual Instr. for
Arctic Exped. by T. R. Jones, Manual Nat. Hist. Geol. Greenl., 1775: 122 (Greenland, by ref. to Fab-
ricius. Fauna Groenl., 1780); Gervals and Boulart, Poiss., 5, 1877: 258, pi. 100 (descr., ill., England to
Scandinavia); Malm, Gciteborgs och Bohuslans Fauna, 1877: 637 (habits, food, west. Sweden); Goode
and Bean, Bull. Essex Inst. Salem, 11, 1879: 31 (occur., depth, off Massachusetts); Winther, Prod.
Ichthyol. Dan. Mar. in Natur. Tidsskr. Copenhagen, (3) 12, 1879: 62 (Skagerrak, north. Kattegat, not in
Baltic); Day, Fish. Gt. Brit., 2, 1880-1884: 364, pi. 179 (refs., descr., ill., habits, Gt. Brit.); Buck-
land, Nat. Hist. Brit. Fish., 1881: 145 (ill.); Mela, Vert. Fennica, 1882: 372, pi. 10 (not seen) ; Jordan
and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 5 (descr., Europe and America) ; Storm, K. norske Vidensk.-
Selsk. Skr., Trondh., 1883:48 (Trondh. Fjord); Bean, Rep. U.S. Comm. Fish. (1882), 1884: 344 (off
Woods Hole); Goode, Fish. Fish. Industr. U.S., Section 1, 1884: 681, pi. 252 (Atlant. coast, U.S.);
Lilljeborg, Sverig. Norg. Fisk., j, 1884: 730 (descr., anat., habits, refs., distrib., Sweden, Norway);
Kingsley, Stand. Nat. Hist., 5, 1885: 67 (habits, Eastport, Maine and Grand Manan); Mcintosh, 3rd
Annu. Rep. Fish. Bd. Scotland, App. F, 1885: 66, 204 (Scotland, not seen) ; Honeyman, Proc. N.S. Inst.
Sci., <S (1), 1886: 230 (off Nova Scotia) ; Cunningham, Quart. J. micr. Sci., N.S. 27, 1887: 49, pi. 6, 7
(habits in aquarium, breathing, reprod. organs, abund., Scotland) ; Jordan, Rep. U.S. Comm. Fish.
(1885), 1887: 791 (in N. Amer. list); Nansen, Bergens Mus. Aarb. (1887), 7, 1888: 5-34,
pi. I, 2 (sex organs, abund. near Bergen, Norway); Vaillant, Exped. Sci. "Travailleur" et "Talis-
man," Poiss., 1888: 384 (off Portugal, 460 meters); Beard, Rep. Brit. Ass. Adv. Sci. (1892), Edin-
burgh, 1893: 789 (Scotland); Goode and Bean, Smithson. Contr. Knowl., 50, 1895; Spec. Bull.
U.S. nat. Mus., 1895; Mem. Harv. Mus. comp. ZooL, 22, 1896: 3, pi. i, fig. i (descr., ill., loc,
depth. Grand Banks, off south. New England and off N. Carolina) ; Smitt, Hist. Scand. Fish., 2nd
Ed., 2, 1895: 1208, pi. 53, fig. 5 (refs., descr., habits, breeding, distr., Scandinavia); Jordan and
Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 211 (Amer. coast south to Delaware); Bull. U.S.
nat. Mus., 47 (i), 1896: 7 (descr., Amer. coast south to Cape Cod); Smith and Kendall, Rep. U.S.
Comm. Fish. (1896), 1898: 169 (off Delaware, Lat. 39° N., Long. 72° W., 75 fath.) ; de Braganza,
Result. Invest. Sci. "Amelia," 1899: 41 (off Portugal, not seen); Garman, Mem. Harv. Mus. comp.
ZooL, 24, 1899: 342, 348, pi. 68, fig. 5 (compar. with other species, rep. on specimens in Mus. Comp.
Zool. Coll., labelled "Trieste"); Dean, Mem. N. Y. Acad. Sci., 2 (2), 1900: 34, pi. 2 (descr. and ill.
of eggs, taken off south. Newfoundland and Georges Bank, 103 and 150 fath., discus.); Hjort, Rep.
Norweg. Fish. Invest., i (1), 1900: 75 (eggs taken in shrimp trawl, Norway, depth and type of bottom;
also young) ; Jensen, Vidensk. Medd. Dansk Naturhist. Foren., Copenhagen, 1900: I, pi. I (descr., ill. of
eggs, off the Faroes) ; Bridge, Camb. Nat. Hist., 7, 1904: 422 (descr., habits, abund. in North Sea, species
of fish preyed on) ; Werner, Zool. Jb., Syst. Abt. i, 2/, 1904: 266 (Norway) ; Jordan, Guide to Study
Fish., /, 1905: 490 (north. Europe) ; Fowler, Proc. Acad. nat. Sci. Philad., ^g, 1908: 461 (old record off
Delaware and Bar Harbor, Maine) ; Kendall, Occ. Pap. Boston Soc. nat. Hist., <? (7), 1908: I (off Maine
and N. Hampshire); Fowler, Proc. Acad. nat. Sci. Philad., 6^, 191 1: 5 (old record off Delaware);
Seabra, Bull. Soc. portug. Sci. nat., 5, igii: 205 (old record off Portugal); Regan, Ann. Mag. nat.
Hist., (8) It, 1913: 397 (class., descr., Gt. Brit., Norway); Sumner, Osburn and Cole, Bull.
U.S. Bur. Fish., 31 (2), 1913: 734 (off Cape Cod); Conel, J. Morph., 2^, 1917: 78, 12 pis.
(urogenital syst., size up to 790 mm., off coast of Maine) ; Roule, Res. Camp. sci. Monaco, 52, 1919: 129
(off Morocco, just inside Strait of Gibraltar) ; Bigelow and Welsh, Bull. U.S. Bur. Fish., 40 (1), 1925:
16 (descr., ill., habits, occur, in Gulf of Maine); Jensen, Rapp. Cons, explor. Mer, jp, 1926: 98;
Koefoed, Rep. Sars N. Atlantic Deep Sea Exped., Zool., 4 (l), 1927: 18 (eggs trawled, off Morocco,
outside Strait of Gibraltar, 535 meters); Nichols and Breder, Zoologica, 9, 1927: 9 (habits, off Cape
Cod); Palingren, Acta zool., 8, 1927: 135 (exper. with pressure on Myxine in aquarium, Drobak,
Norway); Schnakenbeck, in Grimpe and Wagler, Tierwelt N- u. Ostsee, Lief 7, Pt. 12"*, 1927: 3

42 Memoir Sears Foundation for Marine Research

(general); Rey, Fauna Iberica, Feces, /, 1928: 256 (descr., ill., off Portugal); Jordan, Manual Vert.
Anim. NE. U.S., 1929: 5 (in synopsis, Newfoundland to Cape Cod); Breder, Field Bk. Mar. Fish.
Atlant. Coast, 1929: 5 (descr., habits, Arctic to N. Carolina, and east. Atlant.) ; Lyngnes, Z. Morph.
Okol. Tiere, iq, 1930: 591 (descr., and ill. of eggs, abund., type of bottom, Norway); Conel, Dean
Memor. Vol., Amer. Mus. Nat. Hist., j, 1931: 70 (believed not hermaphroditic, see p. 35);
Schnakenbeck, Cons, explor. Mer. Ichthyol. N. Atlant., 1 931: plate not numbered (descr., ill.,
Greenland and Murman coast to Portugal and Adriatic) ; Rep. Newfoundland Fisher. Res. Comm.,
/ (4), 1932: 107 (Newfoundland, no definite loc.) ; Holly, in Schultze, Kiikenthal, et aL, Tierreich,
Lief 59, 1933: 47 (class., refs., descr.); Bigelow and Schroeder, Canad. Atlant. Fauna, 12'', 1934: 2
(descr., ill., comp. with European forms, depth, type of bottom, Greenland and Grand Banks to N. Caro-
lina); Nobre, Faun. Marinh. Portugal, /, 1935: 497 (loc. off Portugal, depth); Gustafson, Arkiv. f.
Zoologi, Stockholm, 28A (2), 1935: l (habits in aquarium, feeding and food, rel. to salinity and light,
abund., west. Sweden) ; Vladykov and McKenzie, Proc. N. S. Inst. Sci., ig (i), 1935: 44 (Nova Scotia
Banks and Bay of Fundy, depth) ; Bigelow and Schroeder, Bull. U.S. Bur. Fish., 4S, 1936: 321 (size of
eggs and number, Georges Bank) ; Liibbert and Ehrenbaum, Handb. Seefisch. Nordeurop., 2, 1936: 323
(habits, eggs, distrib.) ; Norman, "Discovery" Rep., 16, 1937: 5 (comp. with M. australis and M. affinis) ;
Coonfield, Trans. Amer. mici. Soc, 59, 1940: 398—403 (in aquar., St. Andrews, Passamaquoddy Bay;
skin pigment); Jensen, Vidensk. Med., 105, 1942: 55 (Greenland).

Sleep marken, Gunnerus, Trondh. Gesellsch. Schrift., 2, 1766: 230-236, pi. ,3 (descr., ill., habits; con-
sidered a worm; ref. to name Myxine glutinosa Linnaeus. Title page of copy seen is dated 1765, but date
of Gunnerus' paper is given as 1766 by Dean, 1913, Bibliogr. Fishes).

Petramyzon myxine Walbaum, P. Artedi Genera Pise. Emend. Ichthyol., 3, 1792: 500 (diagn., refs.).

GastTobranchus coecus'"- Bloch, Naturg. Ausland Fische, p, 1793: 67, pi. 413 (descr., ill., habits, Denmark,
Sweden, Norway, and Iceland); Bull. Sci. Soc. philom. Paris, i (4), 1797: 26 (equivalent to Myxine
glutinosa Linnaeus, 1758) ; Lacepede, Hist. Nat. Poiss., 2, 1798: 406 (descr., habits) ; Bloch and Schnei-
der, Syst. Ichthyol., 1801 : 534, pi. 104 (descr., ill.) ; Shaw, Gen. ZooL, 5(2), 1804: 264, pi. 134 (descr.,
ill., habits) ; Turton, Brit. Fauna, 1807: 1 10 (brief descr.) ; Cuvier, Regne Anim., 2, 1817: 406; Strack,
Naturg. in Bildern, Fische, Lief 4, 1819-1826: pi. 33 (descr., ill., Norway, America); Yarrell, Hist.
Brit. Fish., 2, 1836: 462 (habits, descr., Gt. Brit.) ; 2, 1841: 612 (same as foregoing) ; Hamilton, Brit.
Fish., 2, 1843: 424 (brief descr.); Buckland, Nat. Hist. Brit. Fish., 1881: 145 (ill., the descr., p. 144,
is as "Gasteobranchus").

Gastrobranche aveugle, Lacepede, Hist. Nat. Poiss., 4° ed., r, 1798: 525, in Buffon, Hist. Nat. (descr., habits) ;
in Sonnini, Hist. Nat. Poiss., 5, 1802-1803: 145, pi. 17 (descr., ill., refs.).

Glutinous gastrobranchus, Shaw and Nodder, Naturalist Misc., ro, 1798: pi. 362 (descr., ill.).

Glutinous hag, Pennant, Brit. Zool., 5, 181 2: 109 (descr., habits).

Myxirut coeca Oken, Lehrb. Naturg., 5 (2), 1816: 127 (descr.; but loc. "Guinea" is no doubt in error) ; Blain-
ville, in Vieillot, Faune Franc, 2, 1825: 2 (descr., meas., north, seas; pi. la not included in copy seen).

Myxine litnosa Girard, Proc. Acad. nat. Sci. Philad., 1852: 224 (descr., off Grand Manan, abund., type of
bottom, depth); Gill, Rep. U.S. Comm. Fish. (1871-72), 1873: 814 (listed Nova Scotia to Massa-
chusetts); Garman, Mem. Harv. Mus. comp. Zool., 24, 1899: 343, pi. 68, fig. 7 (considered distinct
from glutinosa); Dean, Science, N.S. 77, 1903: 433 {limosa considered distinct from glutinosa on
basis of egg case); Jordan, Guide to Study Fish., i, 1905: 490 (retained as distinct from glutinosa);
Halkett, Check List Fish. Canad., 191 3: 38 (Newfoundland south to Cape Cod) ; Regan, Ann. Mag. nat.
Hist., (8) //, 1913: 398 (class., descr.. Bay of Fundy); Fowler, Proc. Boston Soc. nat. Hist., 55, 1917:
no (Bar Harbor, Maine) ; Huntsman, Contr. Canad. Biol. (1921), 1922: 55 (Bay of Fundy, Passama-
quoddy Bay, 18—60 fath., eggs); Holly, in Schultze, Kiikenthal, et al., Tierreich, Lief 59, 1933: 47
(class., refs., descr.); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 8 (con-
sidered distinct from glutinosa, Newfoundland to Cape Cod).

Borer, Couch, Fish. Brit. Isles, 4, 1867: 408 (descr., habits, distrib. in Europe).

Gasteobranchus coecus Buckland, Hist. Brit. Fish., 1881: 144 (descr., habits).

41. Sometimes spelled caecus.

Fishes of the Western North Atlantic 43

Myxine (no specific name) Cole, Anat. Anz., 2y, 1905: 323 (anat., good descr. of teeth, specimens recorded

with 7 gills on one side, or both).
Myxine atlantica Regan, Ann. Mag. nat. Hist., (8) 11, 191 3: 398 (class., descr., off Nova Scotia) ; Holly, in

Schultze, Kukenthal, et al., Tierreich, Lief 59, 1933: 49 (class., descr.).

Order PETROMYZONIDA

Description. In the adult, seven pairs of gill pouches open separately to the exterior,
but open inwardly into a special respiratory tube which is separate from the pharynx and
which ends blind, posteriorly; however, this respiratory tube connects with the mouth
anteriorly/' At the time of metamorphosis this tube loses its connection with the intestine,
while a new pharynx develops above it to form a forward extension of the intestine which
connects with the mouth. Snout without barbels; dorsal and caudal fins separate, supported
by rays; nostril a blind sac, on dorsal surface of head, not opening into mouth; mouth
opens as a funnel or disc surrounded by a circular lip with numerous horny teeth ; sides
of trunk without prominent rows of mucous pores; ear with two semicircular canals; eye
well developed, with lens and iris in adult, although rudimentary in larva; cranium par-
tially roofed over; notochordal sheath with rudimentary neural arches; a complex cartilag-
inous basket around gill pouches; intestine with slight spiral fold, apparently homologous
with the spiral valve of the Chondrichthyes; pancreas represented by scattered follicles.

Development. Sexes separate; eggs small, numerous; development, with larval
(Ammocoete) stage, different in appearance structurally from adult. In some of the
fresh water species the growth stage that normally occurs between the times of metamor-
phosis and sexual maturity is omitted.*'

Habitat. Fresh water, or entering fresh water to breed if marine.

Families. The single family Petromyzonidae, in which the various Lampreys have
been grouped, has been divided recently into two subfamilies, which, in our opinion, may
well be raised to the rank of families as follows:**
la. Upper margin of central mouth with only one dental plate; margin of oral funnel

with a series of fringed, as well as smooth, papillae. Petromyzonidae, p. 43.

lb. Upper margin of central mouth with two separate dental plates; margin of oral disc

with only smooth papillae or cirri. Mordaciidae.

Australia, Tas-
mania, Chile.

Family PETROMYZONIDAE

Characters. Upper margin of the central mouth with only one dental plate, usually
toothed; margin of oral funnel or disc with a series of fringed lappets, as well as a series
of smooth marginal papillae. Characters otherwise those of the order.

42. This respiratory tube represents the pharynx of the larva, into which the gill sacs then open.
4.3. For discussion, see Hubbs, Pap. Mich. Acad. Sci., ^ ist half, 192+: 587.

44. Based on the definitions by Holly (in Schultze, Kukenthal, et al., Tierreich, Lief 59, 1933: 12) and by Piet-
schniann (in Kukenthal and Krumbach, Handb. Zool., 6 [i]. Lief 5, 1935: 54°).

44 Memoir Sears Foundation for Marine Research

Discussion of Genera. Generic characters among the Petromyzonidae, as here lim-
ited, are afforded by the dentition and by the number of dorsal fins, i.e., whether one or
two. Seven genera are recognized in the most recent general synopsis of the family as
limited above/' Five of these are known in the northern hemisphere, but only two, namely
Petromyzon and Lamfetra, occur in the North Atlantic. Petromyzon inhabits only the
western North Atlantic, while Lampetra, which also is marine and anadromous along the
coasts of Europe and northern Asia, is confined to fresh water in North America.

Key to Genera of the Northern Hemisphere*"
la. Only i dorsal fin. Ichthyomyzon Girard, 1858.

Eastern North America.*'

lb. More than i dorsal fin.

2a. Teeth on oral disc, sometimes called labial teeth, close together, arranged in
curvilinear radiating rows (Fig. 2D).

3a. Supraoral dental plate with 2 large teeth; margin of anterior lingual dental
plate deeply indented in the midline.

Petromyzon Linnaeus, 1758, p. 45.

3b. Supraoral dental plate with only i tooth; margin of anterior lingual dental

plate not deeply indented in the midline. Caspiomyzon Berg, 1906.

Caspian Sea.

2b. Teeth on oral disc loosely spaced, not in radiating rows.

4a. Supraoral dental plate with a strongly developed sharp median tooth, as well
as 2 still larger lateral teeth on each side. Entosfhenus Gill, 1862.

Pacific Coast of North Amer-
ica, from California to
Alaska."' *^

4b. Supraoral dental plate without strong, sharp, median tooth, at most with i
or more low, blunt, median denticles. Latnpetra Oken, 1 816.°°

Both coasts of North Atlantic
and western Pacific."'

45. Holly, in Schultze, Kukenthal, et al., Tierreich, Lief 59, 1933: 12.

46. Somewhat amended from the synopsis by Berg- (Annu. Mus. 200I. Acad. Leningrad, 32 [i], 1931: 87). For
synopses of the family as a whole, see Regan (Ann. Mag. nat. Hist., [8] 7, 191 1 : 193) and Holly (in Schultze,
Kukenthal, et al., Tierreich, Lief 59, 1933; 13).

47. In fresh water exclusively.

48. Some of the species that fall in Lamfetra by this key are placed in Entosfhenus by Creaser and Hubbs (Occ. Pap.
Mus. Zool. Univ. Mich., 120, 1922: 6) ; if accepted, this would expand the range of the genus to northeastern
United States, Mexico and Alaska in fresh water; to Japan and the White Sea.

49. Marine, but entering fresh water to breed.

50. Including Eudontomyzon Regan, 191 1, which was classed as a subgenus of Petromyzon by Creaser and Hubbs
(Occ. Pap. Mus. Zool. Univ. Mich., 120, 1922: 2), as a subgenus of Lamfetra by Berg (Annu. Mus. Zool. Acad.
Leningrad, 32 [i], 1931: 92) and by Holly (in Schultze, Kukenthal, et. al., Tierreich, Lief 59, 1933: 22),
with whom we agree.

51. Europe, northern Asia, Japan, North America, Mexico; some species confined to fresh water; others marine,
but entering fresh water to breed.

Fishes of the Western North Atlantic 45

Genus Petromyzon Linnaeus, 1758
Lampreys

Petromyzon (in part) Linnaeus, Syst. Nat., 1758: 230; type species, P. mariiius Linnaeus. European seas.
Generic Synonyms:

A. Adult.

Bathyviyzon Gill, Proc. U.S. nat. Mus., 6, 1883: 254; type species, B. bairJii Gill. Continental slope off Cape

Cod, Lat. 40° 02' N., Long. 68° 51' W., 547 fathoms.">=
Oceanomyzon Fowler, Proc. Acad. nat. Sci. Philad., 59, 1908: 461 ; type species, O. zvihoni Fowler. Atlantic

Ocean.

B. Larva.

Ammocoetus (in part) Dumeril, Dissert. Poiss., 1812: 16; generic diagnosis, no species mentioned.

Atnmocoetes (in part) Cuvier, Regnc .Anim., 2, 1817: 119; emended spelling ior Ammocoetus Blainville, 1 81 2;
type species, Petromyzon branchiaVts Linnaeus, 1758: 230 (larva of Lam.fetra fiuriatilis Linnaeus, 1758;
however, the larva of Petromyzon marinus is not distinguishable from it).°'

Amm.ocaetes (in part) Beithold, in Latreille's Natur. Famil. Tierreich, 1827: 109; emended spelling for
Ammocoetes CM\-\e.T, 1817, and Ammocoetus BLiinville, 1812.

Ammocites (in part) Beithold, in Latreille's Natur. Famil. Tierreich, 1827: 564 (index); evidently a mis-
spelling.

Generic Characters. Two dorsal fins, the ist separated from 2nd by a definite inter-
space, the 2nd demarked from caudal by a deep notch, but continuous with it basally;
teeth renewed periodically by growth, combined with a periodic sloughing off of the outer-
most horny layer, those on oral disc, also called labial teeth, close together in regular
arrangement, the inner series much the largest, the outer series radiating outward in curved
rows: supraoral dental plate small, with 2 teeth; infraoral dental plate broad, with 7 to 9
conical teeth (see footnote 57, p. 47) ; tongue with 3 denticulated plates, the anterior
deeply indented anteriorly in the midline, its toothed margin biconcave; about 70 myo-
meres between rearmost gill opening and anus.

Larva worm-like in appearance, toothless, the oral disc of adult represented in young
by a broad hood-like upper lip and very short lower lip; complexly branched papillae sur-
rounding mouth and present on midzone of upper lip ; eyes rudimentary and not visible ex-
ternally; fins without rays; dorsals not at all, or only faintly, demarked from each other
or from the caudal; muscular segmentation evident externally; gill sacs opening directly
into the pharynx internally; cartilaginous branchial basket rudimentary; pharynx with a
ventral ciliated pocket, a peripharyngeal ciliated groove anteriorly and a dorsal ciliated
tract ; "* gall bladder and bile duct present.

52. This specimen, 275 mm. long, was made the basis of the new genus Bathymyzon because its supraoral and infra-
oral dental plates lacked distinct tubercles. But our own examination of the type specimen (U.S. nat. Mus.,
No. 3331 1) has shown that it simply represents a case where the tubercles have been worn down prior to their
renewal, for a fresh set of very sharp tubercles is exposed when the outer layer of the suboral plate is lifted
free at one end. This appears to apply equally to the type specimen of Oceanomyzon Fowler, 1908.

53. The parentage of the Amm.ocoete larva of Lamfetra fluviatilis was established by A. Miiller (Arch. Anat.
Physiol, wiss. Med., 1856: 323).

54. For description of these, see Dohrn (Mitt. zool. Sta. Neapel, 6, 1886: 59) and Shipley (Quart. J. micr. Sci., 37,
1887:325).

46 Memoir Sears Foundation for Marine Research

During metamorphosis the eyes become functional, the external segmentation dis-
appears, the dorsal fin becomes subdivided, the oral disc and teeth are formed, the bran-
chial basket and skull complete their development, the ventral pharyngeal ciliated pit or
groove becomes the thyroid gland, the pharynx loses its connection with the alimentary
tract, the latter forming a new union with the mouth, while the gall bladder disappears
and the bile duct is obliterated."

Range. Atlantic coasts of Europe and eastern North America; marine but entering
fresh water to breed; also landlocked in certain lakes in the northeastern United States.

Species. It is now generally agreed that the marine Lampreys of this genus represent
only a single species (P. marinus hmnzeus, 1758), and the landlocked form of P. marinus
(dorsatus, Wilder)" appears to be merely a dwarfed race, without any distinguishing
features other than its smaller size.

Pelromyzon marinus Linnaeus, 1758

Sea Lamprey, Lake Lamprey, Stone Sucker

Figure 5

Study Material. Fifty-one American specimens, up to 710 mm. in length, from East-
port, Maine; Exeter, New Hampshire; various localities in Massachusetts and Massachu-
setts Bay; Havre de Grace, Maryland (Chesapeake Bay) ; and the Potomac R., including
the types of Bathymyzon bairdii Gill, 1 884 (U.S. Nat. Mus., No. 3331 1 ) and Oceanomy-
zon wilsoni Fowler, 1907 (Acad. Nat. Sci. Philad., No. 375). Also five Mediterranean
specimens from Nice, Messina and Trieste.

Distinctive Characters. The eel-like appearance of the Lamprey, combined with its
circular oral disc surrounding the jawless mouth and the large number of external gill
openings, places it at a glance among Atlantic fishes.

Description. General form eel-like, the trunk about as thick as high anteriorly, but
somewhat flattened dorsally, hence ovoid in midsection and strongly compressed toward
tail; immature males with a faintly indicated mid-dorsal ridge from about opposite 6th
or 7th gill opening to ist dorsal fin, this much more prominent in large maturing males,
even while still in salt water; females, at maturity, developing a fin-like crest between
anus and caudal fin.

Head, to last gill opening, a little more thank's of total length; nostril prominent,
surrounded by a circular rim, about opposite anterior margin of eye, its distance back from
tip of snout about % to % of length of head to last gill opening. Eye approximately cir-
cular, its diameter about Vi 6 as great as length of head, its anterior margin a little posterior
to posterior edge of oral disc; gill openings round or somewhat oval, about % as long as

55. For a more extensive account, see Bridge (Camb. Nat. Hist., 7, 1904: 429). It has long been realized that the
small Lamprey, repeatedly reported in American waters by early authors as P. nigricans Lesueur, 1818, is merely
the young of P. marinus.

56. In Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 869.

Fishes of the Western North Atlantic

47

horizontal diameter of eye, about equally spaced, the interspaces about as wide as diameter
of eye, the ist gill opening behind eye by a distance about equal to diameter of eye, each
gill opening successively lower on side of head from front to rear. Oral disc circular in out-
line when attached to a fish or other object, but at other times contracted transversely, leav-
ing only a longitudinal fissure open, its diameter when expanded a little greater than
greatest thickness of trunk, or about Va as long as head, its margin with 2 to 4 rows of close-
set fleshy papillae, the inner rows variously fringed and the outermost row also fringed
around posterior part of disc, but smooth around anterior margin.

Figure 5. A Petromyzon marinus, adult about 450 mm. long, from Merrimack River, N. Hampshire (Harv.
Mus. Comp. Zool., No. 35069). B Posterior portion of another specimen of about the same size to illustrate
the variation in the length of the interspace between the dorsal fins. C Oral disc of another adult specimen
from the Merrimack River (Harv. Mus. Comp. Zool., No. 24975), about natural size. D Central mouth of
same with lingual teeth, about 4 x.

Teeth as described above for genus, those on disc about 1 1 2-1 25 in specimens counted,
in curvilinear pattern as illustrated in Fig. 5 C, and marked off in a pavement-like arrange-
ment by narrow furrows of the fleshy tissue, although actually their imbedded bases are
separated, one from the next, by interspaces of considerable width j teeth varying in sharp-
ness in different specimens according to the amount of wear, in extreme cases the supraoral
and infraoral dental plates being nearly smooth."*'

Origin of ist dorsal fin a little posterior to midlength of trunk, its base about V2
as long as head, its height a little more than Y^ as long as its base, with nearly straight
but sloping margins and broadly rounded apexj interspace between ist and 2nd dorsals
varying from very short to about Va as long as base of ist dorsal; 2nd dorsal about twice
as long as ist dorsal basally, but similar in shape, its height a little more than Vs its base,
and separated from caudal by a definite notch," but continuous with latter at its base;

57. The genera Bathymyzon and Oceanomyzon were based on specimens in this condition. See footnote 52, p. 45.

58. Many of the earlier illustrations fail to show this notch, although others do show it

4-8 Memoir Sears Foundation for Marine Research

caudal brush-shaped, with rounded corners, extending forward on ventral side of trunk for
a distance about as long as base of ist dorsal j no separate anal fin; anus anterior to ventral
origin of caudal by a distance about % as long as base of ist dorsal.

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 lead-colored and
as more or less silvery." But large specimens, approaching maturity, are usually olive-
brown above, or of varying shades of yellow-brown, green, red or blue, mottled with a
darker shade of the ground color, although sometimes nearly black, the dark patches con-
fluent; lower surface whitish, gray, or of a pale shade of the same hue as ground color of
back. During the breeding season, at least in the landlocked form, the colors become still
more brilliant, with the ground tint described as turning bright yellow.

Size. The length, at the time of transformation, ranges from about lOO to 200 mm.
Sexually mature specimens, taken in American rivers, average about 2 to 2 V2 feet in length,
the largest of a considerable series from the Navesink River being 33 inches long, weighing
two pounds, four ounces; the maximum recorded length is about three feet.

Developmental Stages. The eggs are small, spherical. A female has been found to
contain 236,000 ova. Segmentation is total, but slightly unequal. The larvae, which differ
widely from the adult in external appearance and habits, as well as in internal morphol-
ogy, are described above (p. 45).

Habits. Since Lampreys never take the hook and are seldom captured in nets, except
close to the beach in pound nets or in estuarine situations with shad nets, they are not often
seen in the open sea; consequently, little is known of their habits in the sea, except that they
are rapid, vigorous swimmers, progressing by an undulating motion, as does an eel, and
that they are exceedingly aggressive in their attacks on other fishes. Occasionally they are
found attached firmly to driftwood and even to boats.

The fact that Lampreys, when encountered in salt water, are usually close to
the land or even in estuarine situations, suggests that most of them remain in compara-
tively shallow water during their sojourn in the sea. But some stray far offshore and
descend to considerable depths. Odd specimens have been caught on the Grand Banks at
86 fathoms north of Emerald Bank; on the seaward slopes of the Nova Scotian Banks off
Nova Scotia, at 200 to 350 fathoms;"" at 85 and at 100 fathoms on the western side of
the Gulf of Maine; at 247 fathoms off Martha's Vineyard and at 547 fathoms off Nan-
tucket, Massachusetts.

The geographic range of the species, combined with observations on the vertical dis-
tribution of temperature at different seasons, shows that it is tolerant of a wide range of
temperature. It is equally tolerant of salinities ranging from fresh water to that of full
oceanic saltness (3.5 per cent or even more).

The normal food of the Sea Lamprey is the blood of other fishes, which it attacks
by sucking with the oral disc. Usually the Lamprey fastens to the side of its victim, where

59. Those that we have seen have lost all color in ...e preservative.

60. Specimens in the U.S. National Museum.

Fishes of the Western North Atlantic 49

it rasps through skin and scales by means of its horny teeth and then sucks the blood. The
secretion of its buccal glands has been found to have an anticoagulating action, thus help-
ing the flow of blood/' Its prey sucked dry, it attacks another. After metamorphosis, young
ones in aquaria attack any fish that may be available and doubtless older Lampreys do the
same. In salt water they have been found preying in this way on mackerel, shad (Alosa),
cod, haddock, American pollock (Pollachius), salmon, basking sharks, the various anad-
romous herrings, swordfish, hake (Urophycis), sturgeons and eels; as many as three or
four sometimes have been found fast to a single shad. Near river mouths the shad and
herring tribes suffer most from them. Judging from their landlocked relatives and from
the frequency with which they have been found attached to marine fish, they must be
extremely destructive to the latter when they are at all plentiful. So far as we are aware,
nothing but fish blood has been found in the stomachs of Lampreys at sea, except fish eggs,
of which they are said to be full occasionally.*^ But it is probable that they take in a certain
amount of solid flesh also, for muscular tissue, as well as blood, has been found in the
stomachs of fresh water Lampreys of another genus."

Before its metamorphosis, the larval Lamprey in fresh water subsists entirely on such
microscopic organisms as may be suspended in the constant stream of water that is drawn
into the pharynx and discharged through the gill chambers, the oral papillae acting as a
sieve to prevent the entrance of grains of sand, etc. When the sieve formed by these papil-
lae becomes clogged, the gill openings are closed and the water is forced back through it."
How the food particles are separated from the water and carried into the oesophagus is not
definitely known.°°

It has been known from early times that the Sea Lamprey is anadromous."'" However,
it does not enter all the streams within its range indiscriminately, but chooses certain ones
and avoids others. As an illustration, we may cite outer Nova Scotia and the Bay of Fundy,
where Lampreys run in the St. Marys, Sackville, Annapolis, Shubenacadie, Petitcodiac
and St. John Rivers, but not in the Margaree, Moser or Apple Rivers, although these
last are also "salmon" rivers.'" For successful reproduction this selectivity is essential
in order to obtain gravelly bottom in rapid water for spawning beds, as well as muddy or
soft sandy bottom in quiet water for the larvae.

The mature Lampreys enter the rivers of the New England and middle Atlantic

61. Gage and Gage, Science, N.S. 66, 1927: 282. 62. Goode, Fish. Fish. Industr. U.S., Sect, j, 1884: 677.

63. Jordan, Guide to Study Fish., /, 1905: 491.

64. For a detailed account of the observations on the larva of the landlocked race, see Gage (Sci. Mon., N. Y., 28,
1929:401).

65. The food particles have been described as being entangled in strings of mucus and swept back with the latter
to the oesophagus by the ciliated tracts on the pharyngeal walls (Bridge, Camb. Nat. Hist., 7, 1904: 429). But
so far as we can learn this has not actually been observed.

65a. See Fontaine (Bull. Inst. Oceanogr. Monaco, No. 848, 1942: 2) for a recent study of the osmotic pressure of
the body fluids of Petromyzon marinus in relation to sexual maturity and to its migrations from salt water into
fresh.

66. The above statement is based on extensive observations made in connection with salmon investigations by the
Biological Board of Canada, communicated to us by A. G. Huntsman.

50 Memoir Sears Foundation for Marine Research

states as early as the end of March or early April. In the rivers tributary to the Gulf of
Maine the runs are at their maximum peak during May and early June. Few, if any, enter
the rivers after that. In New Jersey and Pennsylvania the peak is from late April through
May. Precise seasonal data are lacking for rivers farther south or farther north.°' In many
small streams, and in larger ones also, if their passage is blocked by dams or falls, they may
spawn only a very short distance upstream, even within the influence of the tide, although
invariably in fresh water. They are able to ascend falls, if not too high and steep, by cling-
ing to the rocks with their oral discs and resting, but they do not leap as salmon do in similar
circumstances. They may run up for long distances in large rivers. Such, for instance, was
formerly their habit in the Merrimack and Hudson River drainage systems, while in the
upper tributaries of the Delaware and Susquehanna systems they are still to be found 200
miles or more from the sea, and 1 50 miles upstream in the Savannah River system.

Since the breeding activities of the Sea Lamprey take place in fresh water, a brief
account will suffice here. As the two sexes ripen they become dissimilar in appearance, the
males developing a strong ridge along the back, the females a fin-like crest between the
anus and the caudal fin (p. 46). Analogy with the landlocked form, and dates actually
recorded, suggest that spawning is commenced when the temperature is about 10° C. and
is completed by the time the water has warmed to about 20 to 2 1 ° C.

Spawning takes place in stretches of the stream where the bottom is stony or pebbly.
Working in pairs, a male and a female, with a second female sometimes assisting, make
depressions two to three feet in diameter and about six inches deep in the bed of the stream
by dragging away the stones by means of their oral discs, leaving the stones in a pile down-
stream. 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 de-
scribed as "nests.""* To quote from Regan:'"

The female now secures herself by means of her sucker to some large stone near the upper
end of the nest, and her mate attaches himself to her in the same way near her head, and
winds himself partly round her; then the two together stir up the sand with vigorous move-
ments whilst the eggs and milt are simultaneously deposited. The eggs are covered with an
adhesive substance, and particles of sand stick to them, so that they sink to the bottom of the
nest. The pair now separate and at once commence removing stones from above the nest and
enlarging the pile at the lower end, the sand thus loosened being carried down and covering
all the eggs. The process is repeated at short intervals until the spawning is completed. . . .

After spawning, it seems that the parents die, for not only have they been found dead

67. The landlocked form commences to "run" when the temperature has warmed to about 7 to 9° C. (Surface, 4th
Rep. For. Comm. N. Y., 1899: 227).

68. Bigelow and Welsh (Bull. U.S. Bur. Fish., 40 [i], 1925 : 20) fell into this same error. For an excellent account
of the nesting and spawning of the Sea Lamprey, see Hussakoff (Amer. Nat., ^6, 1912: 729) ; for the land-
locked form, see Surface (4th Rep. For. Comm. N. Y., 1899; 191), Coventry (Publ. Ont. Fish. Res. Lab., Biol.
Ser. No. 20, 1922) and especially Gage (Sci. Mon., N. Y., 2^, 1929: 401).

6g. Regan (Fresh Water Fish. Brit. Isles, 1911: 6), based on accounts of the American landlocked form.

Fishes of the Western North Atlantic 51

repeatedly along the streams/" but their intestines atrophy, they are attacked by fungus,
and they become so debilitated that recovery seems unlikely. The larval stage is believed
to last from three to five years," during which time the larvae live in burrows or under
stones in the mud of the parent stream. Having reached a length of from four to six inches
they undergo transformation to the adult form, an event occupying about two months
(August to September in New England). They then descend the stream to the sea and are
described as reaching salt water in late autumn or early winter in America. The length
of life in the sea is not known, but large ones, not yet mature, are to be found there the year
round.

Numerical Abundance. It is certain that along the American coast as a whole the Sea
Lamprey is now far less numerous than it was, a decrease probably resulting from the
construction of dams that it cannot pass in many of the streams that it enters to spawn.
This decrease has been most severe in the larger rivers of New England. In the Merrimack
River, for example, several cartloads were caught daily for a considerable period in 1847
after the dam was completed there. But so few, if any, now succeed in passing the dams
at Lawrence and Lowell, Massachusetts, notwithstanding the fact that fishways are now
maintained, that a recent survey yielded no evidence that any now breed in the upper
stretches of the river." Similarly, there is a recorded catch of 3,800 in one night at Hadley
Falls in the Connecticut River in 1840, but by 1866 Lampreys had become nearly extinct
in the Connecticut's upper reaches, although still plentiful in its lower part. However,
Lampreys still continue numerous where suitable spawning areas are accessible to them.
For example, we may quote catches of 18, 15, and 119 specimens at three localities on the
Petitcodiac River system, Nova Scotia, during salmon investigations in May and June,
1942 and 1943}" of over 100 on several occasions recently in the lower Exeter River,
New Hampshire;''* and of 98 specimens collected in Swimming River, tributary to Sandy
Hook Bay, New York." While Lampreys, like other anadromous fishes, may seem plen-
tiful when condensed within the narrow bounds of a river's banks, their numbers as a whole
are in no wise comparable with those of the more common salt water fishes.

Relation to Man. In Europe, during the Middle Ages, Sea Lampreys were consid-
ered a great delicacy, and formerly, when they were more plentiful, large numbers were
taken in the rivers of New England for human food, particularly in the Merrimack and
Connecticut Rivers.'" Many were also sold in fish markets in New Jersey as late as the

70. For example, Parley (Rep. Fish. Bay of Fundy, 1851: 156) saw dead Lampreys for miles along the Nerepis
River, New Brunswick, in August, 1840; and he reports a similar situation in the Miramichi (Cat. Fish. N.
Brunsw. and Nova Scotia, 1852).

71. We have no first-hand information to contribute on this point.

72. Bailey (Biol. Surv. Merrimack Watershed, New Hampshire Fish Game Dep., 1938: 155). For an account of
early attempts to restore the Lamprey and other anadromous fish in the Merrimack, see Marston (Biol. Surv.
Merrimack Watershed, New Hampshire Fish Game Dep., 1938: 193).

73. Personal communication from A. G. Huntsman.

74. Collected by R. Witter for the use of the Biological Lab., Harvard College.

75. Nichols and Breder, Zoologica, 9, 1927: 10.

76. For an account of the Lamprey fishery in New England during the first half of the 19th century, see Goode
(Fish. Fish. Industr. U.S., Sect, i, 1884: 680).

52 Memoir Sears Foundation for Marine Research

middle of the 19th century." But so far as we can learn they were never valued in the
more southern part of their American range. For the past half century the Lamprey fishery
has been hardly more than a memory, even in New England, except in a small way for
local home consumption or to supply the needs of biological laboratories. In salt water
they have never been of any commercial importance j the average fisherman might not see
one in a lifetime, nor is there any sale for the few picked up by chance. The larvae are
taken in considerable numbers for bait, however, in the Susquehanna River, and perhaps
in other streams.

Range. Both sides of North Atlantic; northern Norway; only occasional individ-
uals from Iceland;" the Faroes in the east, and southward to Portugal along the coast of
Europe, including the North Sea and the Baltic inward to the Finnish Gulf, the western
Mediterranean (including Algeria),'" and the Adriatic; also reported for West Africa;'"
southern Greenland, Gulf of St. Lawrence and Newfoundland in the west, south to
Florida; breeding exclusively in fresh water, and landlocked in certain American lakes

(P-54)-

Occurrence in the Western Atlantic. The Sea Lamprey has been listed recently for
Greenland,*' where it seems to have been unknown previously. However, apart from this
the estuary and southern side of the Gulf of St. Lawrence (reported from Trois Pistoles,"
Gaspe Basin, Bay of Chaleur and Prince Edward Island) are its northernmost outposts
along the American coast,*' the local stock evidently maintained by reproduction in the
tributary streams, for Lampreys run up the St. Lawrence for at least 40 to 50 miles above
Quebec City." Adults are taken in large numbers also in the Restigouche" and the Mirami-
chi, both in the salt estuary and upstream in fresh water during May and June.*'

Lampreys have never been reported in the rivers of Newfoundland, although these
are fairly well frequented by anglers and wardens." But one specimen was taken 1V2
miles oflF the Newfoundland coast near St. John (found attached to the bottom of a fishing
boat) in November 1946;" one in the U.S. National Museum is recorded for the Grand
Banks south of Newfoundland; also a swordfish, scarred by a Lamprey, was taken ofF Cape
Breton. Earlier characterization of their presence in numbers along outer Nova Scotia is
in line with their presence in the Sackville and St. Marys Rivers, Musquedoboit, Mersey

77. Abbott, in Cook, Geol. N. J., 1868: 830.

78. For list of Icelandic records up to 1909, see Saemundsson (Skr. Komm. Havunders. Kbh., No. 5, 1909: 127).

79. Seurat and Dieuzeide, Bull. Sta. Aquic. Peche Castiglione, 2, 1931 : 83; Algerian record; not seen.

80. Giinther, Cat. Fish. Brit. Mus., 8, 1870: 502.

81. Nordgaard, K. norske Vidensk.-Selsk. Aarsber., 1924: 65; Jensen, Rapp. Cons, explor. Mer, 59, 1926: loi.

82. Personal communication from J. L. Tremblay, conveyed to us by A. G. Huntsman.

83. Bigelow and Schroeder (Canad. Atlant. Fauna, biol. Bd. Canad., 12'', 1934: 4) based its presence on the north
shore of the Gulf on the capture of a small Lamprey taken in the upper Bersimis River, a northern tributary to
the St. Lawrence estuary (Low, Labrador Peninsula, 1896: 329). However, it was not specifically identified and
may have been an Ic/ithyomyzon.

84. Personal communication from J. L. Tremblay, communicated to us by A. G. Huntsman.

85. Specimen in U.S. National Museum.

86. Personal communication from R. A. McKenzie, of the Biological Board of Canada.

87. Information contributed by A. A. Blair, of the Newfoundland Fishery Research Laboratory.

Fishes of the Western North Atlantic 53

and Medway Rivers, and at the mouths of streams flowing into St. Margaret and Mahone
Bays.'* They have also been taken repeatedly as far offshore as the vicinity of Emerald
Bank, the seaward slope of Banquereau Bank and Sable Island Bank, Lahave Bank,
Browns Bank, in the deep gully between the latter and Georges Bank,'* and on the con-
tinental slope off Nantucket and Martha's Vineyard. Lampreys are to be expected any-
where around the shores of the Bay of Fundy, they being recorded from salt water in the
St. Andrew's region; adults were plentiful in the St. John River and its tributaries, for-
merly, and no doubt still are, for small ones were found in the stomach of a Lota maculosa
in Grand Lake, St. John River system, in the winter of 1926-27. They spawn in the An-
napolis and Petitcodiac River systems, as well as in the Shubenacadie River, where larvae
have recently been reported as abundant.'"

They have been reported as being present at many localities along the northwestern
and western shores of the Gulf of Maine and as breeding not only in the Penobscot, Saco
and Merrimack River systems, but in various smaller streams, including the Exeter River,
where they still occur in large numbers, the Lamprey River, a tributary of Great Bay,
New Hampshire, and the Parker River in northern Massachusetts ;°^ no doubt they occur
in other rivers for which there is no published record. In southern Massachusetts they
still run in some numbers in several of the small streams tributary to Buzzards Bay,°^ and
in the Taunton River system.*' There is one record for Nantucket.

They are taken occasionally in pound nets in the Woods Hole region, in Narragansett
Bay where a few breed in the Taunton River, and in Long Island Sound; they spawn in at
least one of the small Long Island tributary rivers which empty into Long Island Sound."
The Connecticut and Housatonic Rivers were famous in past years for their runs of Lam-
preys, although their passage today is barred by dams. Some still enter the Hudson, and
there are records of their presence in the Raritan drainage system. They are common in the
Navesink and Swimming Rivers tributary to Sandy Hook Bay;"" and within the Bay itself
large and small specimens are taken from time to time in pound nets, or found there at-
tached to fish ; they are also taken in Gravesend Bay at the mouth of New York Harbor.

There are numerous recent records for Lampreys, large and small, all along the coast
of New Jersey, north to south ; also up the Delaware River system to the northern part of
Pennsylvania in the Erie River. Although we find no published record of them for the
coastal sector between the mouths of Delaware and Cljesapeake Bays, Lampreys no doubt
occur in this area, for the Bay is a center of abundance for them, with Lampreys recorded

88. Information gathered for us by A. G. Huntsman and R. A. McKenzie of the Biological Board of Canada.

89. Specimens in the U.S. National Museum. 90. Information gathered for us by A. G. Huntsman.

91. Personal communication from Q. A. Arlin, Coastal Warden.

92. Wareham River, Agawam River, Red Brook; also reported in Cape Cod Canal; personal communication from
H. G. Smith, Coastal Warden.

93. Palmer River, personal communication from E. H. Trask, Coastal Warden.

94. The Nissiquague, Hussakoff (J. Amer. Mus. nat. Hist., /j, 1913:323).

95. See especially the account by Hussakoff (Amer. Nat., ^6, 1911: 72) of the nest-building of the Sea Lamprey
in the Navesink.

54 Memoir Sears Foundation for Marine Research

at many localities down to its mouth at Virginia Beach ; they run up the Patuxent, Potomac
and Susquehanna Rivers, the latter a productive spawning region with larvae reported in
abundance in the flats near its mouth.

The next suitable spawning grounds, southward, are the streams discharging via
Pamlico Sound. Correspondingly, Lampreys, both young and adult, are recorded as taken
in shad nets in Albemarle Sound, while they did run up the Neuse River at least as far as
Raleigh, North Carolina, and probably still do. They have been taken in Winyah Bay,
South Carolina,"' and are reported from the Pee Dee and Savannah River systems. Al-
though unreported from Georgia, an early characterization of Lampreys as not uncommon
in the St. Johns River system of northern Florida" is supported by specimens in the
United States National Museum."* But it is not known from the Gulf of Mexico,"'' nor
from the drainage area of the latter.""

It has long been known that a dwarf, landlocked race of the Sea Lamprey occurs
abundantly in Lake Ontario and the lakes tributary to it in northern New York State,
where it is very destructive to other fishes. Formerly it was barred from the upper Great
Lakes by the falls at Niagara. However, with the construction of the Welland Canal, a
passage was opened for it and by 1921 it had reached Lake Erie, where it was unknown
previously^ by 1936 it was in Lake Michigan^ and its spread to Lake Huron and Lake
Superior is to be expected, if it has not already taken place.^""

Synonyms and American References :^''^

Petromyzon marinus Linnaeus, Syst. Nat., /, 1758: 230 (descr., refs., European seas); Mitchill, Trans. Lit.
phil. Soc. N. Y., /, 1815: 461 (descr., N. York); Williams, Hist. Maine, Fish., i, No. 13, 1832 (not
seen) ; Holmes, 2nd Annu. Rep. Nat. Hist. Geol. Maine, 1862: 33, 63 (listed for Maine) ; Goode and
Bean, Bull. Essex Inst. Salem, 11, 1879: 31 (Salem, Massachusetts, specimen attached to American Pol-
lock; also in Massachusetts rivers) ; Jones, List Fish. Nova Scotia, 1879: 1 1 (not uncommon. Nova Scotia) ;
Goode, Proc. U.S. nat. Mas., 2, 1880: 121 (listed for east. Florida) ; Proc. N. S. Inst. Sci., 5, 1882: 97
(same as Jones, 1879); Bull. U.S. Fish Comm., 2, 1883: 349 (American form ident. with European
Nova Scotia to Cape Hatteras; species of fish attacked) ; Jordan and Gilbert, Bull. U.S. nat. Mus., i6
1883: II (descr., Atlant. coast, Europe, N. Amer.) ; Bean, Rep. U.S. Comm. Fish. (1882), 1884
344 (off Woods Hole); Proc. U.S. nat. Mus., 6, 1884: 637 (mouth of Susquehanna R., spring)
Goode, Fish. Fish. Industr. U.S., 1, 1884: 677, pi. 25, upper fig. (descr., habits, former abund.
commercial utilization); Holder, Marvels of Anim. Life, 1885: 5 (abund. in lower Saco R., Maine)
Jordan and Fordice, Ann. N. Y. Acad. Sci., 5, 1885: 283 (class., synonymy, descr., concludes landlocked
form not separable from marinus) ; Lee, Portland (Maine) Adviser (Mar. 3), and Brunswick (Maine)

96. Specimen in the U.S. National Museum.

97. Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900:48.

98. Identification of Lake George, Florida specimens verified by Leonard P. Schultz.

98a. A specimen is listed as Petromyzon castaneous (Girard) by Goode and Bean (Proc. U.S. nat. Mus., 5, 1883:
240), but this is an Ichthyomyzon and probably was taken in fresh water.

99. According to Creaser and Hubbs (Occ. Pap. Mus. Zool. Univ. Mich., 120, 1922) and Gudger (Copeia, No. 4,
1930: 146), a specimen earlier reported as from Muscatine, Iowa (Mississippi drainage system), probably was
in reality from Lake Cayuga, New York, where P. marinus is landlocked.

100. For the history of this expansion of its range, see Hubbs and Pope (Trans. Amer. Fish. Soc. [1936], <5<5, 1937:
172).

101. The Sea Lamprey is also mentioned in most of the larger works on European and American fishes as well as in
a great number of anatomical and embryological papers, zoological textbooks and natural histories.

Fishes of the Western North Atlantic 55

Telegraph (Mar. 13), 1885: (listed for Maine, not seen) ; Gage and Meek, Proc. Amer. Ass. Adv. Sci.,
J5, 1886: 269 (nesting of landlocked form in Lake Cayuga, New York) ; Honeyman, Proc. N. S. Inst.
Sci., 6, 1886: 230 (Nova Scotia, no specific loc.) j Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 792;
Meek, Ann. N. Y. Acad. Sci., 4, 1889: 299 (Lake Cayuga, New York, size when adult); Cox, Bull,
nat. Hist. Soc. New Brunsw., 1893: 42 (south. New Brunswick, Kennebccasis Bay; species of fish
attacked); Gill, Proc. U.S. nat. Mus., ty, 1894: 107 (larva); Cox, Bull. nat. Hist. Soc. New Brunsw.,
13, 1895: 63 (listed for New Brunswick) ; Jordan and Evermann, Bull. U.S. nat. Mus., 47 (l), 1896:

10 (descr., synonyms, Atlant. coast, south to Chesapeake Bay); Rep. LT.S. Comm. Fish. (1895), 1896:
212 (south to Chesapeake Bay); Evermann and Kendall, Rep. U.S. Comm. Fish. (1894), 1896: 584
(small specimens, Vermont) ; Bean, Bull. Amer. Mus. nat. Hist., 9, 1897; 329 (Gravesend Bay, N. York) ;
Mearns, BuU. Amer. Mus. nat. Hist., 10, 1898: 311 (lower Hudson R., juveniles and adults); Smith,
Bull. U.S. Comm. Fish., ly, 1898: 88 (Buzzards Bay) ; Smith, E., Proc. Linn. Soc. N. Y., No. 9, 1898:

11 (fresh and brackish water near N. York); Smith and Bean, Bull. U.S. Comm. Fish., 18, 1899: 180
(District of Columbia); Surface, 4th Ann. Rep. For. Comm. N. Y., I 899: 1 93 (Hudson and Susque-
hanna Rivers) ; Bean, Bull. N. Y. St. Mus., 60, Zool., 9, 1903: 1 1 (descr., habits, Massachusetts, Connecti-
cut, N. York, Delaware, and Susquehanna River systems); Fowler, Rep. N. J. Mus. (1905), 1906: 48
(descr., N. Jersey loc.) ; Roy, Nat. Canad., 23, 1906: 33 (lower St. Lawrence R.) ; Tracy, 36th Annu.
Rep, R. L Comm. inl. Fish., 1906: 44 (Narragansett Bay, breeds in Taunton R.) ; Fowler, Amer. Nat.,
41, 1907: 5 (Penn. loc, Delaware R. system) ; Smith, Geol. Econ. Surv. N. C, 2, 1907: 28 (Albemarle
Sound and Neuse R. system, N. Carolina); Fowler, Rep. N. J. Mus. (1907), 1908: 48 (Delaware R.
system, attacking shad, larva) ; Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: I (New England
loc); Fowler, Proc. Acad. nat. Sci. Philad., 61, 1909: 407 (small specimens attached to anadromous
herring, N. Jersey); Rep. N. J. Mus. (1908), 1909: 351 (descr., size, Delaware River system, species
of fish attacked) ; Tracy, 40th Rep. R. L Comm. inl. Fish., 1910: 58 (same as Tracy, 1906) ; Regan, Ann.
Mag. nat. Hist., (8) 7, 191 1 : 198 (class., descr., Bathymyzoii Gill, also the landlocked form, classed as
synonyms) ; Fresh Water Fish. Brit. Isles, 1911:4 (nesting habits, based on published descr. of landlocked
forms) ; Cornish, Contr. Canad. Biol. (1906-1910), 1912: 79 (Prince Edward I., on mackerel) ; Fowler,
Proc. Acad. nat. Sci. Philad., 64, 191 2: 42, 51,57 (loc, Maryland, Delaware R. and Virginia) ; Hussa-
koff, Amer. Nat.^ 46, 1912: 729 (abund., nesting habits, size at maturity, in Navesink River, N. York) ;
Murray and Hjort, Depths of Ocean, 1912: 644 (Newfoundland Bank, at surface); Stafford, Contr.
Canad. Biol. (1906-1 9 10), 191 2: 54 (Gaspe Bay, Gulf of St. Lawrence); Halkett, Check List Fish.
Canad., 1913: 38 (Maritime Provinces, Prince Edward I. and Gaspe Bay); Hussakoff, J. Amer.
Mus. nat. Hist., 13, 1913: 323 (nest building, Nissiquague R., Long Island, N. York); Nichols, Abstr.
Proc. Linn. Soc. N. Y., No. 20-23, 1913: 90 (off New York) ; Fowler, Proc. Acad. nat. Sci. Philad., 66,
1914: 347 (Delaware R.) ; Kendall, Proc. Portland Soc. nat. Hist., 5 (i), 1914: 9 (loc, Maine); Mc-
Afee and Weed, Proc. biol. Soc. Wash., 28, 191 5: 9 (Potomac River, attached to shad) ; Fowler, Copeia,
No. 31, 1916: 41 (Sandy Hook Bay, N. York season); Proc. Acad. nat. Sci. Philad., 69, 1917: 122
(loc. for Chesapeake Bay) ; Latham, Copeia, 57, 1918: 56 (Long Island, N. York, season) ; Fowler, Proc.
Acad. nat. Sci. Philad., 7/, 1920: 292 (N. Jersey) ; Proc biol. Soc. Wash., 52, 1919: 52 (Delaware and
Susquehanna R. systems); Latham, Copeia, 71, 1919: 53 (Long Island Sd., color of young, season);
Fowler, Proc. biol. Soc. Wash., 5j, 1920: 143 (N. Jersey loc.) ; Breder and Crawford, Copeia, 103, 1922:
II— 15 (Potomac R.) ; Creaser and Hubbs, Occ. Pap. Mus. Zool. Univ. Mich., 120, 1922: 9 (class.,
descr., synonyms); Huntsman, Contr. Canad. Biol. (1921), 3, 1922: (7) 55 (Eastport, Maine, region
of Passamaquoddy Bay, Kennebacasis Bay, St. John R., larvae in Shubenacadie R.) ; Breder, Copeia, 1 14,
1923: 2 (color of young, Sandy Hook Bay; adults in tributaries; nest building in Navesink R.) ; Hubbs, Pap.
Mich. Acad. Sci., 4, 1924: 590 (size of larvae and after transformation, Virginia Beach); Rauther, in
Bronn's Klassen, 6, Abt. I, Buch I, 1924: 677 (class., size, distrib.) ; Bigelow and Welsh, Bull. U.S. Bur.
Fish., 40 (l), 1925: 18 (descr., habits, Gulf of Maine) ; Breder, Copeia, 138, 1925: 4 (Sandy Hook Bay,
New York) ; Jensen, Rapp. Cons, explor. Mer, 59, 1926: loi (listed for Greenland, name only) ; Nich-
ols and Breder, Zoologica, 9, 1927: 10 (descr., habits, season, size at maturity, Sandy Hook Bay; breeding
in tributaries); Fowler, Fish Culturist, 7 (10), 1928 (descr., habits, N. Jersey, Pennsylvania, Mary-
land, Delaware); Hildebrand and Schroeder, Bull. U.S. Bur. Fish., 43, 1928: 43 (loc. for Chesapeake

56 Memoir Sears Foundation for Marine Research

Bay); Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 6 (general account); Dymond, Hart and
Pritchard, Publ. Ont. Fish. Res. Lab., 27, 1929: 37, in Univ. Toronto Stud. Biol, (abund., size,
Lake Ontario; also in Lake Erie); Fowler, Proc. Acad. nat. Sci. Philad., So, 1929: 607 (N. Jersey);
Hubbs and Brown, Trans. Roy. Canad. Inst., 17, 1929: 17 (recent spread to Lake Erie); Jordan,
Manual Vert. Anim. NE. U.S., 1 929: 6 (general account); Truitt, Bean and Fowler, Md. Conserv.
Dep., Conserv. Bull., j, 1929: 27 (Maryland loc.) ; Gudger, Copeia, 4, 1930: 146 (loc. for N.
and S. Carolina); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 8 (in
check list) ; Osburn, Wickliff and Trautman, Ohio J. Sci., 30, 1 930: 120 (listed for Ohio) ; Berg, Annu.
Mus. zool. Acad. St. Petersb., ^2, 1931: 89 (class., descr., refs., distr., alterations at sexual maturity);
Breder, Copeia, 2, 193 1 : 39 (Sandy Hook Bay); Creaser, Copeia, 3, 1932: 157 (Lake Ontario, Lake
Erie) ; Holly, in Schultze, Kiikenthal, et d., Tierreich, Lief 59, 1933: 13 (class., refs., descr., distrib.) ;
Bigelow and Schroeder, Canad. Atlant. Fauna, 12'*, 1934: 3 (descr., distrib.) ; Vladykov and McKenzie,
Proc. N. S. Inst. Sci., rp (l), 1935: 45 (distrib. Nova Scotia and offshore) ; Bigelow and Schroeder, Bull.
U.S. Bur. Fish., 68, 1936: 321 (ref. to Greenland record); Fowler, Bull. Amer. Mus. nat. Hist., yo
(l), 1936: 21 (descr. Amer. and Italian specimens); Vladykov, Copeia, 3, 1936: 168 (N. of Emerald
Bank, off Nova Scotia) ; Hubbs and Pope, Trans. Amer. Fish. Soc. (1936), 66, 1937: 172 (history of its
recent appearance in Lakes Erie and Michigan) ; Bailey, Biol. Surv. Merrimack Watershed, New Hamp-
shire Fish Game Dept., 1938: 151, 155 (former abund. Merrimack R., recent record for Exeter R.) ;
Breder, Bull. N. Y. zool. Soc, 41, 1938: 28 (New York Harbor and lower Hudson R.).

Sea Lamprey, Pennant, Brit. Zool., j, 1776: 76, pi. 8, no. 27.

A'l.imocoetes branchialu (in part) Cuvier, Regne Anim., 2, 181 7: 119 (descr.); GiU, Proc. U.S. nat. Mus.,
ly, 1894: 108. (This is as applicable to the larvae of P. marinus as it is to Lampetra fluviatilis Linnaeus,
1758, the two being indistinguishable at this stage in development; note, however, that Pelromyzon
branchialis Linnaeus, 1 758, is a synonym of L. fiuviatilis Linnaeus, 1758, after transformation.)

Pelromyzon omericanus Lesueur, Trans. Amer. phil. Soc, N.S. /, 1818: 382 (descr., Amer. rivers); Storer,
Rep. Fish. Rept. Birds Mass., 1839: 195 (descr., nesting, Merrimack R., Boston Harbor) ; DeKay, Zool.
N. Y., 4, 1842: 379, pi. 66, fig. 216 (descr., N. York, Hudson R.) ; Linsley, Amer. J. Sci., 47, 1844: 79
(Housatonic R., Connecticut); Storer, Mem. Amer. Acad. Arts Sci., (2) 2, 1846: 517 (descr., Massa-
chusetts, Connecticut, N. York) ; Thoreau, Concord and Merrimack Rivers, 1849: 35 (former occur, in
Concord R., abund. in Merrimack R.; also in subsequent edits.) ; Gray, List Fish. Brit. Mus., Chondropt.,
I, 1851 : 139 (N. Amer. spec.) ; Perley, Cat. Fish. N. Brunsw. and Nova Scotia, 1852: 226 (New Bruns-
wick, St. John, Miramichi R. systems); Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861: 63 (listed
for N. Amer.) ; Storer, Mem. Amer. Acad. Arts Sci., N.S. 9, 1867: 251, pi. 38, fig. 4; also Fishes Mass.:
275, pi. 38, fig. 4 (descr., habits, Massachusetts Bay, Merrimack R.) ; Abbott, in Cook, Geol. N. J., 1868:
830 (N. Jersey, value); Woods, Amer. Nat., j, 1869: 20 (lower Connecticut R.) ; Baird, Rep. U.S.
Comm. Fish. (1871-1872), 1873:827 (Woods Hole) ; Gill, Rep. U.S. Comm. Fish. (1871-72), 1873:
814 (listed Cape Cod to Cape Hatteras) ; Abbott, Rep. U.S. Comm. Fish. (1875-1876), 1878: 827
(abund. in Delaware R. system) ; Goode, Bull. U.S. Fish Comm., 2, 1883: 349 (considered ident. with
marinus) ; Fish. Fish. Industr. U.S., Sect. 1, 1884: 677 (considered probably ident. with marinus).

Pelromyzon nigricans Lesueur, Trans. Amer. phil. Soc, /, 1818: 385 (descr., color, near Philad.; this is young
marinus) ; Storer, Rep. Fish. Rept. Birds Mass., 1839: 197 (size, species of fish attacked, off Massachu-
setts) ; DeKay, Zool. N. Y., 4, 1842: 381, pi. 79, fig. 247 (account from Storer, 1839) ; Linsley, Amer.
J. Sci., 4-/, 1844: 79 (listed for Connecticut) ; Storer, Mem. Amer. Acad. Arts Sci., N.S. 2, 1846: 577
(descr., Massachusetts, Connecticut); Gray, List Fish. Brit. Mus., Chondropt., /, 1851: 139 (refs.);
Thompson, Hist. Vermont, 2, 1857: 150 (descr., habits, small specimens, Vermont); Gill, Proc. Acad,
nat. Sci. Philad., Addend., 1861: 63 (N. Amer.); Storer, Mem. Amer. Acad. Arts Sci., N.S. 9, 1867:
253, pi. 39, fig. 6; also Fishes Mass., 1867: 277, pi. 39, fig. 6 (descr., size, Massachusetts and Connecticut,
fishes attacked) ; Abbott, in Cook, Geol. N. J., 1868: 830 (N. Jersey, with americantis) ; Amer. Nat., 4,
1870: 719 (N. Jersey, attached to an eel); Provancher, Nat. canad., 8, 1876: 262 (descr., common in
Gulf of St. Lawrence, species of fish attacked) ; Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 11
(descr., Atlant. coast U.S. and Cayuga Lake, N. York).

Fishes of the Western North Atlantic 57

Petromyzon (no specific name) Perley, Rep. Fish. Bay of Fundy, i 851 : 156 (abund., season, St. John R. system;
death after spawning).

Ammocoetes unicolor DeKay, Zool. N. Y., 4, 1842: 383, pi. 79, fig. 250 (good descr. and ill. of larva, Con-
necticut R. at Northampton, Massachusetts).

Petromyzon maculosus Gray, Cat. Fish. Coll. Descr. by L. T. Gronow, in Brit. Mus., 1854: 2 (equivalent to P.
marinus, England).

Lamfetra marinus Malm, Vetensk. Handl. Goteborg, 8, 1863: 87; Goteborgs och Bohuslans Fauna, 1877: 630
(occur, west. Sweden).

Ammocoetes fiuviatilis Jordan, Ann. N. Y. Acad. Sci., /, 1879: 120 (Cayuga Lake, N. York) ; not Petromyzon
fluviatilis Llnmeus, 1758.

Petromyzon marinus dorsatus ]ord3n and Gilbert, Bull. U.S. nat. Mus., j6, 1883: 869 (compared with marinus,
Cayuga Lake, Gage Mss.) ; Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 792 (fresh water, east. U.S.) ;
Rauther, Bronn's Klassen, 6, Abt. i, Buch I, 1924: 678 (landlocked, L. Ontario, and lakes of northern
N. York); Berg, Annu. Mus. zool. Acad. St. Petersb., 52 (l), 1931: 89 (ref. to landlocked form);
Vladykov, Trans. Canad. Inst., 20, 1 93 5: 122 (Gt. Lakes).

Petromyzon {Bathymyzon) bairdii Gill, Proc. U.S. nat. Mus., 6, 1884: 254 (continental slope off Nantucket,
547 fath.) ; Goode and Bean, Smithson. Contr. Knowl., 30, 1895: 4 (by ref. to Gill, 1884).

Bathymyzon bairdii jordin, Rep. U.S. Comm. Fish. (1885), 1887: 792 (by ref. to Gill, 1884); Jordan and
Evermann, Bull. U.S. nat. Mus., 47 (l), 1896: 9 (descr., Gulf Stream).

P^/roTOyzow fo«<ro/or Wright, Prelim. Rep. Fish Fish. Rep. Ont. Game Fish Comm. (1892), 1892:439 (Land-
locked race, Gt. Lakes) ; not Ammocoetes concolor Kirtland, 1840, which is an Ichthyomyzon.

Petromyzon unicolor DeKay, or P. dorsatus Wilder; Gage, Wilder Quar. Century Bk., 1 893: 430 (cf. land-
locked form with marinus; descr. and photo of larvae).

Petromyzon marinus (americanus) McClure, Zool. Anz., 16, 1893: 360 (segmentation of egg, N. Jersey,
specimens, season, temp.).

Petromyzon marinus unicolor Jordan and Evermann, Bull. U.S. nat. Mus., 47 (l), 1 896: 10 (descr., the land-
locked race; lakes of north. N. York) ; Rep. U.S. Comm. Fish. (1895), 1896: 212 (lakes of north, and
cent. N. York); Surface, Bull. U.S. Comm. Fish., ty, 1898: 209, pi. 10, upper fig. (life hist., damage
to fishes. Lake Cayuga, N. York) ; 4th Annu. Rep. For. Comm. N. Y., 1899: 194, 200 (extended account
of life history of landlocked form, damage to fishes, lakes of N. York) ; Halkett, Check List Fish. Canad.,
191 3: 38 (Lake Cayuga, reported Lake Champlain) ; Bensley, Contr. Canad. Biol. (i9ll-l9l4),2, 1915:
10 (abund., Lake Ontario, species of fish attacked); Dymond, Publ. Ont. Fish. Res. Lab. 4, in Univ.
Toronto Stud. Biol., Ser. 20, 1922: 60 (first Lake Erie record); Evermann, Bull. U.S. Bur. Fish., 2/,
1922; 95 (listed for Gt. Lakes) ; Gage and Gage, Science, N. S. 66, 1927: 282 (anticoagulation action of
secretion of buccal glands) ; Gage, Biol. Surv. Oswego R. System, 1928: 163 (not seen). Probably not
Ammocoetes unicolor DeKay, 1842, which appears to be an Ichthyomyzon; see Creaser and Hubbs, Occ.
Pap. Mus. Zool. Univ. Mich., 120, 1922: 9.

Lamprey (no scientific name), Bumpus, Science, N.S. 8, 1 898: 850 (date of breeding in Taunton R.) ; Nichols
and La Monte, Amer. Mus. Novit., 901, 1937: I (a swordfish marked by a Lamprey, Cape Breton, Nova
Scotia) ; Stroud, Bull. Bowdoin Coll. (April 1), 6, 1939: 22 (Kent Island, Bay of Fundy, in stomach of
cod).
Oceanomyzon tvilsoni Fowler, Proc. Acad. nat. Sci. Philad., 5p, 1908: 461 (descr., ill., Atlant. Oc).
Petromyzon marinus var. dorsatus Huntsman, Ottawa Nat., 5/, 1917: 25 (Lake Ontario) ; Coventry, Pub. Ont.
Fish. Res. Lab., p, in Univ. Toronto, Stud, biol., ser. 20, 1922: 133 (spawning, nests, Humber R., tribu-
tary to Lake Ontario).
Lake Lamprey, Gage, Sci. Mon., N.Y., 28, 1929: 401 (habits, larvel devel., photos).

Lamper-eel, Marston and Gordon, Biol. Serv. Merrimack Watershed, New Hampshire Fish Game Dep., 1 93 8:
193, 197 (hist, of attempts to restore Lampreys in upper Merrimack R.).

Probable Synonyms:

Ammocoetes bicolor Lesueur, Trans. Amer. phil. Soc, /, 1818: 386 (young, after transformation, Connecticut
R.) ; DeKay, Zool. N. Y., 4, 1842: 383, pi. 79, fig. 248 (descr., ill. after Lesueur, 1818); Linsley, Amer.

58 Memoir Sears Foundation for Marine Research

J. Sci., 47, 1844: 80 (listed for Connecticut) ; Storer, Mem. Amer. Acad. Arts Sci., N.S. 2, 1846: 519

(brief descr., after Lesueur, 1818); Gray, List Fish. Brit. Mus., Chondropt., i, 1851: 146 (by ref. to

Lesueur, 1818) ; Gill, Rep. U.S. Conim. Fish. (1871-72), 1873:814.
Not Ammocoetei affendix DeKay, Zool. N.Y., 4, 1842: 381, pi. 64, fig. 211 (included by Jordan and Ever-

mann, 1896, as a synonym of Petromyzon marinits, but probably a Lamfetra because two dorsal fins are

pictured as continuous at their bases).
Not Petromyzon lamottcni DeKay, Zool. N. Y., 4, 1842: 382, pi. 79, fig. 249 (included by Jordan and Ever-

mann, l 896, as a doubtful synonym of P. marinus, but probably a Lamfetra because two dorsal fins are

pictured as continuous, at their bases).
Not Petromyzon lamfetra Pallas, Zoogr. Rosso Asiat., 5, 1814: 66 (name and loc. only; included in the

synonymy of P. marinus by Holly, 1933, but probably a combination of a Lamfetra with Cajfiomyzon,

because of localities White Sea and Caspian).

CHAPTER THREE

Sharks

BY
HENRY B. BIGELOW and WILLIAM C. SCHROEDER

ACKNOWLEDGMENTS

In preparing the present paper we have received invaluable assistance and co-operation
from many people, both here and abroad. Numerous correspondents have contributed
information of various sorts, including photographs of freshly caught specimens, and
these are noted under the accounts of the respective species. We are particularly grate-
ful to Luis Howell-Rivero and Stewart Springer for contributing much-needed speci-
mens, together with extensive notes on the occurrence of many species from Cuba and
Florida. We wish also to express our gratitude to the following persons: J. L.
Baughman for an extensive series of specimens from Texas; Maj. C. M. Duke,
U.S. Army, for a specimen of the fresh-water Shark from Lake Nicaragua, and
F. B. Richardson for arranging its transportation; Capt. James Whaley for sending
us a "Mako" taken off Ocean City, Maryland; Richard Foster and John Huntington
for a "Mako" from Cat Cay; T)r. Heloisa Alberto Torres for entrusting to us the
type specimen of Scyliorhinus haeckelii (Ribeiro) for study; Lieut. -Commander
J. W. Lowes, U. S.N. R., for records of his own captures (s/' Carcharodon, together
with color notes, measurements and photographs; President Don Anastasio Somozo
of Nicaragua, Capt. W. B. Brinker and Frank Fisher of the National Geo-
graphic Society for photographs of newly caught specimens of the fresh-water Shark
of Lake Nicaragua; Carlos de la Torre for permitting us to have photographs taken
of the late Filipe Poey's unpublished drawings of Cuban sharks, with copies of Poey s
unpublished notes; A. Fraser Brunner and Lieut. Colonel W. P. C. Tenison for
drawings o/" Pseudotriakis and Echinorhinusyrow specimens in the British Museum,
and Miss Ethel wynn Trewavas who enlisted their kind assistance; W. H. Rich
for records of recent captures of the Greenland Shark by Gulf of Maine fisher-

59

6o Memoir Sears Foundation for Marine Research

men; John Tee- Van, who furnished bibliographic citations; A. S. Romer, who assisted
us in determining classifications of major groups and who contributed summaries of the
various genera which occurred in earlier geologic periods; Ludlow Griscom and J. A.
Peters, who assisted us in the solution of puzzling questions regarding scientific nomen-
clature; H. W. Fowler, J. T. Nichols and L. P. Schultz, for their assistaiice; and
to the late Thomas Barbour, not only for putting the collections of the Museum of
Comparative Zoology so freely at our disposal, but for constant encouragement and
personal help in many ways.

GENERAL DISCUSSION

Scope of Study. The following pages give descriptions, illustrations, life histories and geo-
graphic distribution, as well as lists of published citations, for all species of sharks so far
known on the western side of the North Atlantic. In some genera represented within these
geographic limits, a few additional species occurring in the western South Atlantic but not
reported as yet north of the equator are included as addenda. The pertinent characteriza-
tions of the suborders and families, as well as the keys to major groups and genera, cover
the sharks as a whole and in some cases this applies to the species keys within genera. How-
ever, it seems wiser in other cases to limit them geographically until the relationships of
species from the western North Atlantic to those of adjacent parts of the ocean are clarified.

Descriptions. The descriptions are based on the Study Material listed under each
species, except for Pseudotriakis microdon and Echinorhinus brucus, no specimens of which
were available. The discussions of habits and geographic distribution are based on data of
our Study Material, on information submitted to us through the co-operation of numerous
correspondents, checked in many cases by our own observations, and on previously pub-
lished accounts.

Keys. The keys, whether to higher groups or to genera and species, have been ar-
ranged solely for the purpose of facilitating the identification of any shark. Therefore,
we have selected as alternative characters those that are most easily visible and measurable.
Our personal views on phylogeny are not discussed. Species within a genus are presented
in alphabetical sequence.

References. All citations listed in the references, with the few exceptions noted, were
consulted in the original through the kind co-operation of the following libraries and insti-
tutions: Museum of Comparative Zoology and other departments of Harvard University;
American Academy of Arts and Sciences; Yale University; United States Fish and Wild-
life Service; Library of Congress; American Museum of Natural History; American
Philosophical Society; and Massachusetts Institute of Technology.^

Sources of Material. The well-rounded collection of sharks from many parts of the
world, preserved in the Museum of Comparative Zoology, has been our chief source of

I. For pre-Linnaean names and references, see Garman (Mem. Harv. Mus. comp. Zool., 36, 1913).

Fishes of the Western North Atlantic

6i

reference. The extensive collections of the United States National Museum have also been
made available to us, as well as specimens from the Academy of Natural Sciences at Phila-
delphia, the American Museum of Natural History, the Bingham Oceanographic Collec-
tion at Yale University, the Carnegie Museum at Pittsburgh, the Chicago Natural History
Museum, the California Academy of Sciences, the Museu Nacional in Rio de Janeiro and
the Woods Hole Oceanographic Institution. Other specimens and data are acknowledged
on page 59. We regret that war conditions have prevented us from examining the types
of many species of sharks that are in the British and European museums.*^

Proportional Dimensions and Illustrations. The actual measurements from which
the proportional dimensions of the several species have been calculated were taken on
a horizontal line between perpendiculars at given points; for example, the distance from
tip of snout to origin of first dorsal fin is the line BC in the accompanying illustration (Fig.
6), not AC; the length of snout in front of nostril is line ED, not DF. The illustrations
have been drawn on this basis so that the proportions can be scaled from them directly, if

Lowtr
Caudal
lota

Figure 6. Outlines of a typical shark to illustrate terminology and methods of measurement.

desired. In the shark illustrations, the dermal denticles pictured are from high on the sides
of the trunk, below the first dorsal fin, unless otherwise noted. All the illustrations are
original, except as indicated; the great majority were prepared by the well known zoologi-
cal artist E. N. Fischer. Rhincodon was drawn by Janet Roemhild, Pseudotriakis by A.
Fraser-Brunner and Echinorhinus by Lieut. Colonel W. P. C. Tenison.

la. For a list of type specimens of cyclostomes, elasmobranchs and chimaeroids in the Paris Museum, see Bertin
(Bull. Mus. Hist, nat., Paris, [2] //, 1939: 65-93)-

62 Memoir Sears Foundation for Marine Research

Class CHONDRICHTHYES

Elasmobranchs and Chimaeroids

Characters. Fish-like vertebrates with well developed lower jaws and bony teeth;
2 pairs of appendages supported by pectoral and pelvic girdles; a cartilaginous skeleton
which, while more or less calcified, lacks any true bone; scales essentially tooth-like in
structure, the ectoderm as well as the mesoderm sharing in their formation (placoid
scales); two nostrils, each single, partially subdivided; olfactory sacs blind, not opening
into mouth; posterior end of vertebral column either straight or heterocercal ; sympathetic
nervous system, pancreas, spleen and contractile arterial cone present; two, three or more
series of heart valves; swim bladder absent.

Relation to Other Classes. Chondrichthyes are most obviously separated from the
Cyclostomes by their well developed lower jaws and bony teeth, by their much more
highly developed cranium and visceral skeleton, as well as by the presence of pectoral and
pelvic girdles, paired limbs, spleen and a contractile conus arteriosus with two, three
or more series of heart valves. The lack of true bone in the skeleton, which is most
apparent in the skull and pectoral girdle, separates them from all so-called higher fishes,
including the Lung Fishes (Dipnoi). Other features marking them apart from bony fishes
are: (a) cranium, without sutures consequent on its lack of bone; (b) outer margins of fins
supported by horny rays or filaments as contrasted with bony rays or spines among bony
fishes; (c) first gill pouch with a row of gill filaments, which are lacking among bony fishes,
and gill filaments attached to the interbranchial septa except at the tips (free for a greater
or lesser part of their length in bony fishes) ; (d) no true operculum, but at most a fold
of skin serving the same purpose (in Chimaeroids); (e) nostrils single; (f) teeth simply
imbedded in the gums, not firmly attached to jaws or imbedded in the latter; (g) scales
(placoid or dermal denticles) tooth-like in structure, consisting of a hollow cone of
dentine of dermal origin surrounding a pulp cavity; externally this cone is covered with a
layer of an enamel-like substance (vitro-dentine) or possibly true enamel (among Rays)
formed at least partially by the epidermis; i.e., it is of ectodermal origin,''' whereas in bony
fishes the scales are formed by the dermis alone, i.e., they are bone-like in origin; (h) ferti-
lization is internal, and in all modern representatives is effected by cartilaginous appen-
dages, commonly called claspers, which are developed from the inner margins of the pelvic
fins of the males (among such of the bony fishes as have internal fertilization, the intromit-
tent organs are developed either from the genital papilla or urogenital orifice, or in connec-
tion with the anal fin, or as a special structure situated on the chest, but never from the pelvic
fins). Furthermore, the invariable presence among the Chondrichthyes of the so-called
spiral valve in the posterior portion of the intestine separates them from most bony
fishes," as does the presence of a pair of spiracles in many of them (representing vestigial
gill clefts) which open on the dorsal or dorso-lateral side of the head, frequently with a

lb. See Tomes (Philos. Trans., igo, 1898: 460) for further discussion of this question.
1. In a vestigial form in sturgeons, Amia, lung fishes and some others.

Fishes of the Western North Atlantic 63

number of branchial lamellae that probably aid in the oxygenation of blood to the eyes and

brain.*

Key to Subclasses

I a. 5 to 7 pairs of gills and 5 to 7 pairs of gill clefts, each of the latter opening separately
to exterior} dorsal fin or fins, and spines if present, rigid, not erectile; skin with or
without dermal denticles; teeth numerous; upper jaw or palatoquadrate cartilage not
fused to cranium, although it may be locally attached to it; rostral cartilage fused to
cranium; vertebral centra more or less clearly differentiated, and the notochord more
or less constricted segmentally; at least some of vertebrae of trunk region with articu-
lated transverse ribs; the 2 halves of pelvic girdle fused into a single bar; anus and
urogenital canals discharge into a common cloaca; males withou*. prepelvic or frontal
tenacula. Elasmobranchii; Sharks, Skates, Rays, p. 63.

lb. Only 4 pairs of gills and 4 pairs of gill clefts, with only I opening to the exterior on
each side of head;* dorsal fin and spine erectile; skin in adult naked, without dermal
denticles; teeth represented by 6 pair" of grinding plates; upper jaw or palatoquad-
rate cartilage fused with cranium; rostral cartilages articulated to cranium, not fused;
no vertebral centra, and the notochord not constricted segmentally; ribs lacking; the 2
halves of pelvic girdle separate; no cloaca, the urogenital aperture being distinct from
anus and posterior to it; males with an erectile prepelvic tenaculum, and usually with a
frontal tenaculum on the head also." Holocephali; Chimaeroids.

Subclass ELASMOBRANCHII

Sharks, Skates, Rays

Characters. Five to 7 pairs of gill clefts, all opening separately to exterior and not
covered by an opercular fold of skin; dorsal fin or fins, and fin spines if present, rigid, not
erectile; spiracle present or absent; skin armed with numerous placoid scales or "dermal
denticles"; teeth numerous and in several series; no frontal or prepelvic tenacula in males;
notochord more or less constricted segmentally, persisting only between the vertebrae in
many cases; vertebral centra more or less well developed; at least some of vertebrae of
trunk region with short articulated ribs; upper jaw or palatoquadrate cartilage not fused
to cranium, although it may be firmly attached to the latter by i or 2 articular surfaces of
limited extent; lower jaw, or Meckel's cartilage, articulated to upper jaw, and as a rule
attached also to hyomandibular arch, which thus takes part ia the suspension of jaws;'
rostral cartilages ( 1-3) fused to cranium; the 2 halves of pelvic girdle fused into a single
bar; inner margin of each pelvic fin in males modified to form a copulating organ or
"clasper" grooved for the passage of the sperm and supported by an axial cartilage, the

3. Among the skates, the water that reaches the gills is inhaled through the spiracles, at least for the most part.

+. The true gill clefts open into a common branchial chamber (covered by an opercular fold of skin supported by

cartilaginous rays) which opens to the exterior by a single secondary branchial aperture on each side.
5. The frontal tenaculum is lacking in the genus Harriotta,
I. Among notidanoids this arch is much reduced and has no attachment to the lower jaw.

64 Memoir Sears Foundation for Marine Research

latter a rearward extension of the basal cartilaginous element of the fin. Development
either oviparous/ ovoviviparous,' or viviparous;* embryos with transitory external gills.'
For convenience, the modern representatives of this subclass may be grouped in two
orders; the one to include all living sharks as well as the fossil group (hybodonts) from
which they appear to have descended, the other to include the skates and rays, which have
probably descended from the hybodonts also.

Key to Orders of Modern Elasmobranchs

I a. Gill openings at least partly lateral ; edges of pectoral fins not attached to sides of head
anterior to gill openings; upper margin of orbit free from eyeball (eyelid free).

Selachii; Sharks, p. 64.

lb. Gill openings confined to ventral surface; edges of pectoral fins attached to sides of
head anterior to gill openings; upper margin of orbit not free from eyeball (no free
eyelid). Batoidei; Skates, Rays.*

Order SELACHII
Modern Sharks

Characters. Gill openings at least partly lateral; edges of pectoral fins not attached
to sides of head anterior to gill openings; upper edges of orbits free from eyeballs, as free
eyelids. Other than as indicated above, no sharp lines can be drawn between the sharks on
the one hand and the skates and rays on the other, so far as external characters are con-
cerned; the gap between the prevalent cylindrical body shape of the former and the much
flattened form of the latter is bridged by one group of true sharks (Squatinoidea, p. 533).

Skeletal differences between the two groups are considerable, however, correspond-
ing chiefly to the highly specialized external features of the rays. Thus, to mention only
the most obvious, among sharks the propterygial cartilage of the pectoral bears many
fewer radials than the metapterygial and is smaller than the latter,^ while the reverse is
the case among rays. Among sharks the shoulder girdle is neither directly nor firmly at-
tached to the vertebral column, nor are its elements united above, while in rays it is at-
tached above by a separate scapular element, or elements. The suspension of the jaws
differs also between the two groups; in sharks the ceratohyal cartilage is attached to the
lower end of the hyomandibular as well as more or less intimately to the posterior end of
the lower jaw (Meckel's cartilage), thus assisting to support the latter; in rays (typically,

2. Eggs laid before hatcliing.

3. Eggs hatching and embryos developing within the mother, but without placental attachment.

4. Embryos attached to the uterine wall of the mother by a yolk-sac placenta.

5. For an excellent and comprehensive account of the morphology of the elasmobranchs, see Daniel (Elasmobranch
Fishes, Univ. Calif. Press, 1934).

6. The skates and rays are classed as a suborder only by some authors. But the skeletal differences between them and
the typical sharks discussed above (p. 64) seem to us sufficient to set them apart as a separate order.

1. This applies even to the Squatinoidea (pp. 77, 533).

Fishes of the Western North Atlantic 65

at least) it is connected by a ligament with the hyomandibular only, at the upper end of the
latter, and is entirely separate from the lower jaw, hence it does not take a direct part in the
suspension of the lower jaw. The cranium is also much less intimately connected to the
vertebral column in sharks than in rays. It has been stated that while the upper jaw (palato-
quadrate cartilage) bears a transverse process by which it is attached to the cranium by a
ligament in sharks, this is not true in rays. Actually, however, the two groups intergrade in
this respect, for the ligamentary attachment (but not the transverse process) is present in
some skates at least,^ while the transverse process may be represented by an articular area
only, in some sharks (Heterodontidae), or altered to a rounded knob in others (Isuridae).

Replacement of Teeth. The number of series of teeth that are in actual use at any
given time varies from one to four or even five in different sharks, and in different parts of
the jaw of a given shark. There are also one to several additional reserve series lying in a
reversed position (points up in the upper jaw, points down in the lower) against the inner
surfaces of the gums, new series being developed in a deep dental groove along the inner
margin of the jaw and covered over by a fold of the mucous membrane. As functional
teeth are lost, whether by accident or by orderly migration to the outer anterior edge
of the jaw, those of the next younger series move forward to replace them. This process
of replacing older and smaller teeth by younger and larger ones continues throughout life,
there being as many reserve rows in adult specimens of a given species as in the young. It is
this process that provides for the increase in the size of the teeth, which accompanies the
growth of the shark.

Among the majority of galeoid sharks the loss of older teeth is irregular, in part acci-
dental, the older teeth being lost and replaced by younger ones individually. Thus, in
Carcharias^ two days to one week are required for a directly observable tooth to become
detached} sometimes one may be seen dangling from one of the outer corners of its base;
as a rule, too; the teeth are lost singly and not by entire series simultaneously. Evidently this
last feature applies equally to the various carcharhinids, to the White Shark {Carcharo-
don), and to the Hammerheads, for their jaws commonly show various stages in the
progression of teeth. There is no reason to doubt that the replacement is correspondingly
irregular in those squaloids in which the teeth are slender, raptorial and spaced along the
jaw. But in others of that group, in which they form a continuous cutting edge (Figs. 88 A,
97), the process of replacement involves a revolution of the younger series as a unit
from the reversed to the erect position, otherwise gaps would occur in the series in use,
which is seldom, if ever, the case.'' The teeth of the older series, which are being replaced,
do not loosen and fall out forthwith, but continue for some time attached to the outer side
of the gum to which they have moved, although standing meantime at a somewhat lower

2. This is recorded and well illustrated by Parker (Trans, zool. Soc. Lond., ro, 1879: 223, pi. 41, fig. 4) for Raja
clavata.

3. Breder, Copeia, 1924:42.

4. We have never detected such a gap in numerous specimens of Squalus acanthias except as a result of mutilation by
the hook at time of capture; nor among specimens of Centroscymnus, Dalatias, Isistius or Somniosus that we have
examined.

66

Memoir Sears Foundation for Marine Research

level than the series that has replaced them. Consequently, the jaw, as viewed from the
outside, may show two series (perhaps even three or only one), while the series next
younger than those in actual use (one, or possibly two) may be either oblique or may still
lie in the reversed position (Fig. 7), depending on the momentary stage of replacement.
It is probable, also, that replacement of teeth is similar (i.e., by series rather than singly)
in the Smooth Dogfishes (Mustelus), in which they are arranged in mosaic.'

Figure 7. Semidiagrammatic cross-sections of upper
jaws (left) and lower jaws (right) of two adult speci-
mens of Squalus acanthias illustrating difTerent stages
in the replacement of teeth.

Form, Activity, Size. Most sharks are subcylindrical in form 5 some are as beautifully
streamlined as the larger members of the mackerel tribe, giving rise to the vernacular
name "Mackerel Sharks." On the other hand, a few are very much flattened dorso-ven-
trally and expanded laterally, so that they resemble skates or rays in general appearance.
There is a wide variation in their swimming also. The Mackerel Sharks (Isuridae) are
exceedingly active, swift and powerful, whereas others, such as the Greenland and Portu-
guese Sharks (Somniosus, Centroscymnus) are so sluggish and inert that it is a question of
some interest how they succeed in capturing their prey. Sharks also vary widely in size.

5. Cawston, in a series of recent papers, has maintained that the reserve teeth in sharks come into use only when an
individual tooth, lying in front, happens to be lost through injury (Brit. dent. J., 55, 1938: 321 j S. Afr. J. Sci.i
3S> '938: 321 ; Dent. Rec, 59 [10], 1939: i ; Dent. Rec, 60 [11], 1940: 435; S. Afr. dent. ]., 14 [12], 1940:
312; ridskr. Wetensch. en KunS; 2, 1941 ; S. Afr. dent. J., ly, 1943: 117; S. Afr. dent. J., ly, 1943: 295; Copeia,
1944: 184). However, direct observations on the shedding of teeth in Carcharias (Breder, Copeia, 1942: 42)
have proved that the shedding of the older teeth, and their replacement by younger teeth, is a normal process.
Successive stages in the process, such as those illustrated in Fig. 7 for the Spiny Dogfish {Squalus acanthias), are

Fishes of the Western North Atlantic 67

On the one hand the Whale Shark {Rhincodon) reaches a length of at least 45 and prob-
ably 50 to 60 feet, making it by far the largest of fishes, while certain scyliorhinids (p.
213) and triakids (p. 239) mature at lengths of only 300 to 400 mm. (less than 1V2
feet).

Breeding and Development. Fertilization is internal (p. 62). The males have a
pair of copulatory organs (claspers or myxopterygia) that are developed as appendages
from the inner edges of the pelvic fins, supported by cartilages derived from the basiptery-
gial cartilage of the latter, with a groove along the inner side for the guidance of the sperm.
In copulation they are inserted through the cloaca of the female into her two sexual ori-
fices.* In some species at least, as in the European Scyliorhinus caniculus, only one clasper
is inserted at a time and coitus lasts about twenty minutes. As a rule the eggs are enclosed
in horny cases, at least for a time, but the Greenland Shark (Somniosus) may be an excep-
tion (p. 520).

Development is oviparous in some sharks, ovoviviparous in the majority and truly
viviparous in still others. In the first type the horny egg capsules usually (but not always)
bear long tendrils at the corners at one or both ends, by which they are attached to algae,
etc.} in one group (Heterodontidae) there is also a very prominent spiral flange, giving
the egg a very distinctive appearance. In one species representing this category {Scylio-
rhinus caniculus) the period of incubation is 157-178 days.' Among the ovoviviparous
species the embryos, early liberated from the capsule, develop in the oviduct of the
mother; they are nourished from the original yolk alone which is chiefly in the yolk sac,
or from yolk, as well as from nutritive fluids secreted by filaments which are developed
from the walls of the maternal oviducts; these nutrients are absorbed both by the yolk
sac of the embryo and in many cases by appendages borne by its stalk, the so-called um-
bilical or placental cord. The young are not born until fully formed and after the yolk sac
has been absorbed. In the viviparous species the young lie in special uterine dilations of
the oviducts during development; the yolk sac develops folds and processes that inter-
digitate with corresponding folds of the uterine wall, thus forming the so-called yolk-
sac placenta. The number of young is small, as compared with many bony fishes; the
maximum number so far reported in a gravid female of any ovoviviparous shark of which
we have found record is 82.

Intelligence and Senses. It is recognized by common observation that the intelligence
of sharks is of a very low order, alth'ough we cannot find that any significant tests have
been made of their capacity for learning. Their indifference to injury of any kind is pro-
verbial. In numerous recorded instances a shark, severely mutilated or even disembow-

also instructive, since in this case entire series are involved and not merely individual teeth alone; Spiny Dogfish
are so commonly stocked by biological supply houses that large numbers are easily to be had.

6. The method of copulation with which Aristotle was acquainted and which is now a matter of common knowledge,
was rediscovered by Louis Agassiz (Proc. Boston Soc. nat. Hist., i^, 1871 : 340) ; see also Garman and Putnam
(Proc. Amer. Ass. Adv. Sci., ij [2], 1874: 14) and Garman (Proc. Boston Soc. nat. Hist., ly, 1875: 171, 172).
For an eyewitness account of the act of coition in Scyliorhinus caniculus, see Bolaw (Z. Morph. Okol. Ticre, ^5,
1888: 321) i and Liibbert and Ehrenbaum (Handb. Seefisch. Nordeurop., j, 1936: pi. 20, fig. 249) show an ex-
cellent photograph of a pair so engaged.

7. Observations in the Hamburg Aquarium; see Bolaw (Z. Morph. Okol. Tiere, 35, 1888: 324).

68 Memoir Sears Foundation for Marine Research

elled, has returned to continue feeding on the carcass of a whale, or on offal thrown over-
board, or even to take the hook a second time. However, some of their senses are of a much
higher order than the foregoing might suggest, particularly their sense of smell. It has
been shown by experiment that the Smooth Dogfish {Mustelus) seeks its prey chiefly by
smell (p. 248) and it can be only because of their keen scenting ability that sharks gather
so quickly around a whale that is being cut up, or around a dead horse or other carcass
in some tropical harbor. As evidence of the ability of a large shark to scent a comparatively
small object from a considerable distance, we might mention an occasion in the Gulf
Stream, off Key West, Florida, when we saw a large carcharhinid tracking our bait (a
Spanish mackerel) up-current, its dorsal fin cutting the surface as it tacked back and forth
across the trail, and finally dashing forward on a direct line.

Experiments on the Smooth Dogfish {Mustelus canisY have shown that it has at
least fair vision for objects that are close at hand, and this no doubt applies to sharks gen-
erally. In experiments, however, they seldom responded to any object until the latter was
within one foot of them,' thus bearing out the general concept that sight is of very little
importance in the lives of sharks.

No evidence of any response by sharks to vibrations of high frequency (sound)
has been reported, although it seems well established that their auditory (8th) nerves,
as well as the nerves of the lateral-line system, are sensitive to water vibrations of low
frequency.'"

Luminescence. A few genera are luminescent, as noted below (p. 509), but the great
majority are not.

Food. Sharks are carnivorous without exception. Seaweeds have often been found in
the stomachs of one or another species, but no doubt these were taken with the animals on
which they were preying, and the more voracious kinds are so indiscriminate in their feed-
ing that they often swallow any kind of inedible rubbish." A few that have crushing teeth
(e.g., Mustelus and the heterodonts) feed largely on hard-shelled crustaceans (crabs,
lobsters) or on mollusks; but the majority prey chiefly on fishes smaller than themselves,
on squid and to some extent on pelagic Crustacea. In general the size of the prey is relative
to the size of the shark. However, some of the more fiercely predaceous species regularly
attack other fish, including other sharks nearly as large as themselves, if they are in a
position to do soj sea turtles and seals are a regular item in the diet of some sharks. On the
other hand, the two largest species (Whale Shark and Basking Shark) subsist wholly on
minute planktonic forms, chiefly Crustacea, and on small schooling fishes.

Number of Species. In spite of the antiquity of the group, and in spite of the fact
that they appear to be as numerous and as varied now as at any time in the past, there are
many less species of sharks than of bony fishesj not more than 225 to 250 are now known.

Danger to Man. Dependable information on the danger of sharks to man is frag-
mentary 5 nevertheless, we think it necessary to discuss the subject briefly, since it is of

8. Parker, Bull. U.S. Bur. Fish., 29, 191 1 : 46. 9. Parker, Bull. U.S. Bur. Fish., 33, 1914: 64.

10. Parker, Bull. U.S. Bur. Fish., 24, 1905 : 201. 11. For instances, see p. 69.

Fishes of the Western North Atlantic 69

interest to seamen, to fishermen and to seaside visitors who frequent shark-infested regions.

Most species of sharks are either too small, too sluggish, too weakly armed or nor-
mally live at too great a depth to be of any potential danger. This applies also to some of
the larger and better-armed species which feed on small rather than on large prey. On
the other hand, there are unquestionably a considerable number of species, proverbially
voracious, which are large, active and armed with very effective teeth, and which habit-
ually feed on large prey such as other sharks, large fishes and sea turtles; it is equally
true that many persons in various parts of the world have been attacked by sharks. Notable
among dangerous species are the White Shark {Carcharodon), the Tiger Shark (Galeo-
cerdo), certain members of the genus Carcharhinus, the Lemon Shark (Negaprion brev-
irostris) and the larger Hammerheads. All these bear evil reputations as potential man-
eaters and the charge seems to be sufficiently proved against them in one part of the world
or another (see discussions below under the respective species). Perhaps the Makos {I sums
oxyrinchus and /. glaucus), which feed chiefly on small fish, may deserve a similar repu-
tation, but we do not believe that the Blue Shark {Prionace glaucd) does, unless attracted
by blood to a wounded man in the water; under these conditions any shark more than five
or six feet long would be a menace. Among the foregoing list the White Shark {Carcharo-
don carcharias) is beyond question the most dangerous. Fortunately, however, even the
smaller sizes of this species appear not to be common anywhere, while large adults are very
seldom seen, especially close inshore.

In estimating the risk, even from the more dangerous species, we should keep in
mind that man is not the habitual prey of any shark; hence the scent of man in the water
is not likely to prove especially attractive, since it is presumably by scent chiefly that sharks
discover and track down their food. On the other hand, sharks soon learn to gather where
dead animals or garbage are to be expected, as where refuse from a slaughter house drifts
out to sea. When in a feeding mood, some of the more voracious kinds, especially the
"Tiger," will gulp down wholly indigestible objects, such as boots, old clothes, a sack of
coal, tin cans, etc., as readily as a chunk of salt pork or a dead dog. Nor is there any reason
to suppose that the scent of man is repulsive to any shark.

In view of the foregoing it is not astonishing that many shark fatalities are on record,
well attested by hospital reports or otherwise.^' Shark attacks are much more frequent in
warm waters than in cold, as might be expected. For example, from 1919 to 1933, 37
cases were reported for various parts of Australia on seemingly conclusive evidence, with
many more for earlier years." In fact, the shark menace is so real in New South Wales

II. The many reports of fatalities by sharks which are not so attested may be left out of the account; some have
been based on rumor alone (even the individuals concerned may have been imaginary) ; others, involving the
overturning of small boats, etc., or the disappearance of swimmers without trace, may have resulted from quite
other causes; and in still others, an observed attack may have been by a Barracuda {Sphyraena) and not by a
shark.

13. See Coppleson (Med. J. Aust., April 15, 1933 : 449) for a list of these and other such happenings for Australia,
with references for shark attacks in other parts of the world; see also, Whitley (Fish. Aust., /, 1940: 13, 259)
for further details, discussion and list of attacks in Australian waters, with photographs of wounds suffered
by victims.

JO Memoir Sears Foundation for Marine Research

that patrols are maintained on the more popular bathing beaches, some of which are fur-
ther protected by wire netting} in some parts of Cuba bathing areas are similarly pro-
tected with closely spaced palmetto logs. Attacks have been reliably reported from South
Africa, the Red Sea, India," Ceylon, the East Indies, the Philippines, the Pacific coasts of
Mexico and Panama, the coast of Ecuador, the Gulf of Mexico, the West Indies, the
Guianas, the eastern coast of the United States (see below), tropical West Africa, the
eastern Mediterranean, Port Said, and no doubt from other regions as well. However,
the incidence of attacks is very irregular. Sharks, for example, although plentiful enough
along the beaches of Florida, are so slight a menace that we have positive word of only
one or two attacks in recent years (pp. 368, 408), despite the fact that many thousands of
persons bathe there constantly throughout the year. The most recent instance was of a girl
severely bitten while bathing in the surf, only waist deep, at Mayport, Florida, in late May
or early June 1944. The size of the shark's jaws, as outlined by the wounds, showed that
it was only sV^-^V^ feet long, and other circumstantial evidence pointed to a Carcharhinus
maculfpinnis as responsible." Shark attacks appear to be similarly unusual throughout
the West Indian region in general ; although local inhabitants in Porto Rico and among
the Antilles have informed us that, while they would not hesitate to swim by day even if
sharks were about, it would be hazardous in the extreme to do so at night.

Attacks occur from time to time, hov/ever, even to the northward along the Atlantic
coast, although sharks of the dangerous sorts are progressively less numerous in that direc-
tion. Near Charleston, South Carolina, for example, several well-attested cases have been
reported recently.^' More widely heralded was a series of attacks on six bathers on the
New Jersey coast in July 191 6, probably by a small Carcharodon (see p. 139) that was
caught nearby a few days later. More recent still was an attack in Buzzards Bay, Massa-
chusetts, July 26, 1936, on a bather who was so badly injured that he died shortly after-
ward in the New Bedford Hospital. The shark, about six feet long but not identified as to
species, was driven away by the victim's companions who came to his rescue in a boat. How-
ever, these last two instances are the only ones along our northeastern coast that have come
to our attention in a lifetime experience. Such events are certainly no more common along
the bathing beaches of the northern Mediterranean or of northwestern Europe, for we
have not found a single definite case of recent date recorded in the literature of sharks, in
natural history journals or in the press. It also happens that the few large sharks which
are at all common close along the shore north of Cape Hatteras on the one side of the
Atlantic, or of Portugal on the other," are either wholly innocuous, as is the Basking
Shark, or at least have never been proved guilty of attacks on bathers, whatever may be

i+. The Augustine Friar, Sebastiao Manrique, in 1643, was an eyewitness to attacks by sharks on pilgrims wading
out into the sea at Hugli, in Bengal ; see translation by CoUis, The Land of the Great Image, 1943 : 76.

I 5. This case was reported to us by Stewart Springer and was mentioned in the local press. The victim was treated
at the dispensary of a Naval Base near by.

16. Burton, Sci. Mon., N. Y. 40, 1935 : 279.

17. Sand Shark (Carc/mrias taurus) ; Basking Shark {Cetorhinus maximus) ; Tope (Galeor/iinus galeus) ; Common
Mackerel Shark or Porbeagle {Lamna riasus) ; Blue Shark {Prionace glauca) ; Brown Shark (Carc/iar/iinus mil-
berti) ; Dusky Shark (^Carcharhinus obscurus) .

Fishes of the Western North Atlantic 71

true of them in warm seas. This applies also to the Greenland Shark (Somniosus) of Arctic
seas, for while it preys habitually on living seals it is so sluggish that both Eskimos and
whale fishermen look upon it with contempt (p. 522).

The general conclusion from the foregoing is that in continental waters in temperate
and boreal latitudes on either side of the North Atlantic the danger to a swimmer of
attack by a shark, although existent, is so exceedingly remote as to be wholly negligible,
unless it be known that a shark of some dangerous kind has been seen in the vicinity re-
cently. We believe this to be equally true of the coastwise waters of the North Pacific
south to southern California on the one side and to northern Japan and northern China
on the other, although our personal information is less extensive there than for the North
Atlantic.

Categorical statement is not so safe for warmer seas, because reported attacks have
been much more frequent there, because large sharks of the potentially dangerous kinds
are far more numerous, and because local conditions differ widely between different re-
gions. For coral-reef areas all our sources of information, including personal experience,
agree that while dangerous sharks may be numerous offshore and along the seaward slopes
of the reefs, they seldom enter the lagoons and are much less likely to enter any smaller
pools among banks and coral heads. Large sharks do not often come into wading depth
along open beaches, especially if the swell is breaking heavily a short distance out, as is so
often the case, unless attracted by slaughterhouse wastes, etc., or by corpses, as in India.
Shoal-water bathing is therefore reasonably safe in such situations, at least in the daytime,
unless as just qualified or unless the local inhabitants advise against it. In deeper harbors,
more open to the sea, it is wiser to err on the side of caution, unless the locality is declared
safe by local report, which is usually reliable.

Under normal circumstances the danger of attack to a bather offshore, even in tropical
seas, also appears very slight, for the chances are much against any dangerous sharks being
close at hand or of their being in a feeding mood if present. But if persons in the water are
bleeding from injuries the danger from shark attack may be imminent and the results may
prove fatal. The more voracious of the larger sharks are excited by blood in the water to
such a degree that they will make ferocious attacks, whether the object be fish, whale or
man, dead or alive. Attempts to drive the attacker away by blows or splashing are likely
to be futile, although success might be achieved if the swimmer were uninjured. Instances
are on record, apparently on good evidence, of crews from capsized boats being attacked
and pulled down in tropical seas 5 the southwestern Pacific is reported to have been the site
of such events during the recent war.

So extensive is the resulting laceration likely to be that bites from any large sharks
are extremely dangerous for they are followed by very rapid bleeding and severe shock,
even if the wounds are not still more directly destructive. Thus, "so far as known, about
one-half of Australia's shark attacks have ended fatally.'"'

18. Whitley, Fish. Aust., /, 1940: 16.

72 Memoir Sears Foundation for Marine Research

To class sharks "harmless" as a group, as some authors have done, is contrary to all
the weight of evidence. On the other hand, the danger of attack to the ordinary bather is
very small indeed, except in such special localities and under such circumstances as those
mentioned.

Commercial Imfortance. Sharks are the objects of minor fisheries in the warmer
parts of the world, largely for their liver oil and for their fins (considered a great delicacy
by certain oriental races), and to a lesser extent for their hides and flesh.

Shark liver oil was formerly valued highly in combination with other fish oils for
tanning, the yield from local fisheries being considerable, notably from the Greenland
Shark. Recently a new demand for the liver oil of some species has developed because of
the high vitamin content. This is notably the case in the northeastern Pacific, the California
catch having risen from about 555,000 pounds in 1936 to about 7,800,000 pounds in 1940,
although it dropped to 2,613,431 pounds in 1944, This increase has resulted from the oil
of one vitamin-rich species, the Soupfin or Oil Shark {Galeorhinus galeus). Interest in the
commercial possibilities of shark oil has given impetus to shark fisheries along the eastern
coast of the United States also, but to date no western Atlantic shark that occurs in large
numbers has been found to equal the California Galeorhinus in showing a consistently high
Vitamin A content (nor do representatives of that same species in the eastern Atlantic),
although individual specimens, such as the larger Hammerheads, may give a high yield.
The following table, condensed from a more detailed one,'^ gives the maximum and mini-
mum potencies in Vitamin A (stated in U. S. P. Units) for the liver oil of several Florida
sharks.

Number of Potency

Sfecies
Carcharodon carcharias
Ginglymostofna cirratum
Galeocerdo cuvier
Carcharhinus leucas
Carcharhinus milberti
Carcharhinus obscurus
Carcharhinus Umbatus
Negafrion brevirostris
Sphyrna diplana 34 137,000 5j400

Sphyrna tudes many 340,000 8,250

19. Springer and French (Industr. Engng. Chem., 56 [19], 1944: 190). See Walford (U.S. Fish Wild Life Serv.,
Fish. Mkt. News, 6 [6], 1944: 4) for a detailed table giving the amounts of Vitamin A, both per gram of oil
and per pound of liver, for several species taken in the Gulf of California. For methods of calculating amounts
of Vitamin A in livers, see Sanford (U.S. Fish Wild Life Serv., Fish. Mkt. News, 7 [i], 1945: 6)and Bolomey
and Tompkins (Fish. Bull., Sacramento, 64, 1946: 73) ; for relationship between liver yield of Vitamin A and
the biology of the Soupfin Shark (Galeorhinus galeus) in California waters, see Ripley and Bolomey (Fish.
Bull., Sacramento, 64, 1946: 39).

')ecimens

Maximum'

Minimum

6

7,350

750

many

6,720

641

many

4,625

1,375

many

20,875

1,812

many

15,500

283

6

58,500

6,500

many

22,250

4,250

many

11,425

3,000

Fishes of the Western North Atlantic 73

From earliest times the fins of certain sharks have been highly prized as food in
China and Japan because of their gelatin content, and often the demand has exceeded the
supply. We regret that statistics are lacking for the total amounts marketed. However,
as long ago as 1850 not less than 40,000 sharks were caught yearly in the Arabian Sea,
chiefly for the export of fins to China.^° Until the recent war, supplies were regularly
drawn from as far afield as California. In fact, the species from which the fins are taken
there {Galeorhinus galeus) has been known locally as Soupfin Shark, although at present
the name Oil Shark is more commonly applied to it.

The kinds of sharks which have firm meat are better food fish than is generally ap-
preciated, and various species are regularly placed for sale in the fish markets of the tem-
perate parts of the world. In Chile, for example, 2.7 million pounds of sharks (about lO
per cent of the total catch of fish in all categories) were landed in 1940 to be consumed
locally.''^ Local consumption may be considerable in northern Europe also where the
Spiny Dogfish {Squalus acanthias, p. 462) is in demand. Along the coasts of the United
States the larger sharks have been increasingly marketable of late years. EflForts have also
been made by the United States Bureau of Fisheries to promote the sale of canned meat
from the Spiny Dogfish (p. 462) as "gray fish," but the project failed when discoloration
and spoilage resulted from the generation of ammonia in the cans due to the high content
of urea in shark flesh.

It has long been known that the hides of many of the larger sharks yield leather com-
paring favorably with cowhide, and minor fisheries for this purpose have been carried on in
various parts of the world. In the western North Atlantic these fisheries have been located
off southern Massachusetts, North Carolina, eastern Florida, Key West, Florida, the
Bahamas, and among the Virgin Islands. Up to the present time, however, the amount of
shark leather marketed has been very small, as compared with leather from domestic ani-
mals. In some cases the fisheries have been short-lived, because of depletion of the local
stock of sharks which are large enough to be serviceable (for local instance, see p. 104).
But in regions where a fishery may be expected to draw its supply of sharks from a wide
area, as on the east coast of Florida with the Gulf Stream near at hand, the prospects of
commercial success appear to depend chiefly on an expansion of the demand for shark
leather.

The dermal denticles of many sharks are so sharp and so close set that the skins make
an effective abrasive, and shark skin, often known as "shagreen," was formerly in wide
use by cabinet makers the world over for polishing wood, but it has been almost entirely
supplanted of late by other recently developed abrasives, except perhaps in remote parts
of the world.

At the present time shark scrap, like other fish scrap, is in demand for feeding poultry
and other livestock, and in sum total considerable amounts are marketed. However, we find

io. See Buist (Proc. zool. Soc. Lond., /*, 1850: 100) for an account of the Karachi Fishery in India.
21. Fiedler, Geog. Rev., 5^, 1944: 104.

74 Memoir Sears Foundation for Marine Research

no statistics as to actual amounts of shark scrap for comparison with scrap from other fishes.
Small amounts of shark refuse also find their way into commercial fertilizers, but here
again definite statistics are lacking. Efforts have even been made in the Maritime Provinces
of Canada (p. 462), as well as in the United States and possibly elsewhere, to develop this
industry. But so far as we know all such attempts have been short-lived, because of irregu-
larity in the supply of sharks.

Sharks are not as highly esteemed for food as are various bony fishes that support the
great fisheries, partly because the available supply is only a fraction as great; hence, the
landings of sharks are correspondingly smaller, especially in northern seas, and they are
correspondingly less in value. Thus, the reported catch of sharks (4,417,700 pounds) was
less than one-half of one per cent of the total catch of all kinds of fish (1,458,687,600
pounds) along the Atlantic and Gulf coasts of the United States in 1942, and about one
per cent (10,171,900 pounds out of a total of 1,346,559,600 pounds of fish of all kinds)
on the Pacific Coast of the United States. In warmer regions the shark catch may rank
relatively higher, the catch of bony fishes being much smaller than it is in the northern seas,
e.g., the Chilean catch mentioned above (p. 73). But previous experience suggests that
fisheries for large sharks, if intensive and on a large scale, are likely to be short-lived, seem-
ingly through exhaustion of the local supply.

Recently commercial shark fishing in the western Atlantic has been carried on most
actively off the southern part of the North Carolina coast (Morehead City), along eastern
and southeastern Florida (Mayport, Salerno, Cortez and Key West) and off the Bahamas.
The yield consists chiefly of Tiger Sharks (Galeocerdo), Sand Sharks (Carcharias), vari-
ous species of Carcharhinus, Nurse Sharks {Ginglymostoma) , Hammerheads, and Lemon
Sharks (Negaprion) ; on the whole, the first two rank foremost in commercial importance,
both in quantity and in value. Anchored gill nets with a stretched mesh of about 20 inches,
and anchored set lines (best of chain) with snoods of wire rope every six to eight feet, both
set at depths of 3 to 20 fathoms, are the types of gear chiefly used. The catches of Green-
land Sharks that are made in the waters off Iceland and Greenland are mostly by long lines,
or by hand lines. Basking Sharks have usually been harpooned because of their large size,
and this applies equally to the Whale Sharks that have been fished from time to time in
the Bay of Bengal and in the waters around India.

Habitat and Range. Sharks are marine for the most part, but a few members of the
genus Carcharhinus run far upstream into brackish or even into fresh water in large rivers
such as the Ganges, the Tigris and the Zambezi. We have received two specimens of Car-
charhinus leucas, a well known west tropical Atlantic species, that had been taken in Lake
Yzabal, Guatemala (p. 341), and one landlocked species is known in Lake Nicaragua
(p. 381). Many are oceanic and roam the high seas,"" while others dwell on the ocean
bottom or close to it. In warm latitudes they are often seen following ships for days

2ia. A shark tagged off Ventura, southern California, was recaptured on the west coast of Vancouver Island, it
having migrated about a thousand miles; see Ripley (Calif. Fish Game, [2] 52, 1946: loi).

Fishes of the Western North Atlantic 75

at a time, feeding on garbage thrown overboard. However, the great majority are con-
fined to comparatively shallow water. While a few, which are mentioned below in the
appropriate connections, find their homes on the continental slopes at depths of some hun-
dreds of fathoms, the greatest depth for which there is definite record of the capture
of a shark of any species is about 1,500 fathoms. Nor is it likely that any shark is a
regular inhabitant of the floor of the oceanic abyss. The group is cosmopolitan, but the
great majority inhabit the tropical-subtropical belt. Characteristically temperate species
are much fewer in numbers, and only one genus (Somniosus) is a regular inhabitant of
truly polar waters.

Classification. The question of how to subdivide the modern sharks so as to illustrate
the supposed phylogenetic relationship of diflFerent groups, which has been argued since
the days of the early comparative anatomists, is one that we pass over briefly.

The paleontologic history of the groups of sharks that still exist throws little light on
the matter. Groups as diverse as the heterodonts, orectolobids, galeoids and squatinoids
were all in existence as far back as the Upper Jurassic, and the hexanchids were present
in the middle Jurassic and the squaloids in the Cretaceous 5 while "by the beginning of the
Tertiary period all of the living families of Elasmobranchs appear to have come into
existence.""

Students of living sharks have agreed generally that the most primitive are those
(Hexanchidae and Chlamydoselachus) in which the vertebrae are calcified but weakly,
if at all, and in which the notochord is but little constricted segmentally. The hexanchids
likewise appear to agree with the Mesozoic genus Hybodus both in these features, and
further, in the mode of suspension of the upper jaws (p. 78 ) . However, if these supposedly
primitive groups were actually derived from the hybodoids, as has been suggested, they
have diverged widely from the ancestral stem by a multiplication of gill arches {Hybodus
had five only), by the loss of the second dorsal fin and of fin spines, and by modification in
their dentition. On the other hand the heterodonts, which resemble the ancient hybodoids
so closely in dentition, in number of gills and in the presence of two dorsal fins and fin
spines that they have often been united with them in a single suborder, differ from the
hybodoids in having the vertebrae well calcified, the notochord strongly constricted seg-
mentally and the upper jaw (palatoquadrate cartilage) attached to the cranium in one re-
gion only, without the postorbital connection which has often been regarded as primitive."

Among the remaining, and far more numerous, living members of the order, much
weight in classification has been given to the degree and arrangement of the internal calci-
fications of the ve'-tebral centra. These centra may consist of only a primary ring surround-
ing the notochord ("cyclospondylic"), or of a primary ring with secondary calcifications as
well, either in concentric rings around the primary one or in bars (simple or branched)

12. Romer, Vert. Paleont., 1933: 54-

23. See De Beer (Devel. Vert. Skull, 1937: 421-4.25) for definitions of the rather complex terminology that has
been employed to define the different methods of attachment of upper jaw to skull.

76 Memoir Sears Foundation for Marine Research

radiating out from the primary ring and which may or may not invade the four primary
uncalcined areas that radiate out to the bases of the neural and haemal spines."

Jordan and Evermann," for example, followed in 1930 by Jordan, Evermann and
Clark," classed all sharks, other than the notidanoids, in two orders, Asterospondyli (cor-
responding to our Galeoidea and Heterodontoidea) and Cyclospondyli (including the
squaloids, pristiophoroids and squatinoids) ." The sharks have also been subdivided accord-
ing to the external or the skeletal structure of the male copulatory organs. However, this
results in grouping the notidanoids with the squaloids, and the squatinoids with the Batoi-
dei in one case,"' or Chlamydoselachus with the Holocephali in another.^'

The majority of modern authors'" have given primary consideration to characters
that are visible externally in both sexes, such as the number of gill openings, the presence
or absence of the anal fin, number of dorsal fins and the dentition.

White" classed the Selachii ("Antacea") as a superorder with four orders — Hexan-
chea, Heterodontea, Squalida and Galea, dividing the Squalida into the suborders Squalida
and Rhinida, the Galea into the suborders Isurida and Carcharinida. Still more recently,
Bertin'" classed the skates and rays with the sharks as four suborders under the order
Euselachii, and distributed among three suborders (Scylliformes, Musteliformes and
Lamniformes) the assemblage of families that are united here as the suborder Galeoidea
(White's order Galea).

In our opinion, however, the characters on which these subdivisions of the galeoid
sharks are based — the presence or absence of a nictitating fold or membrane, the position
of first dorsal relative to pelvics, the details of vertebral calcification and the morphology of
the spiral valve — are of a lower taxonomic grade than are those by which the notidanoids,
heterodontoids, squaloids, pristiophoroids and squatinoids can be defined."'

Apart from the names employed, the subordinal classification used in the present
paper follows that of Rey," which in turn is based in its essentials on Garman's" system,

24. Hasse (Nat. Syst. Elasm. Algem. Theil, 1879) proposed the names "cyclospondylic" for the vertebral type
with primary annular calcification only, "tectospondylic" for that with secondary concentric rings of sec-
ondary calcification, and "asterospondylic" for that with radiating bars in addition to the primary ring. Regan
(Proc. zool. Soc. Lond., 1906: 737), however, has more recently limited "asterospondylic" to the type with four
radial bars only, which do not invade the four primary uncalcified areas, and has expanded "tectospondylic" to
include all types that are not either "asterospondylic" as so limited, or "cyclospondylic," an emendation that has
caused some confusion in nomenclature.

25. Bull. U.S. nat. Mus., 47 (1), 1896. 26. Rep. U.S. Comm. Fish. (1928), 2, 1930.

27. In the interim, Jordan (Class. Fish., Stanford Univ. Publ. Biol., 3 [2], 1923) had recognized five orders of
sharks and employed the term "Tectospondyli" in place of "Cyclospondyli" for the order including squaloids,
plus squatinoids.

28. Huber, Z. Wiss. Zool., 70, 1901 : 671. 29. Leigh-Sharpe, J. Morph., ^2, 1926 : 336.

JO. Notably Regan (Proc. zool. Soc. Lond., 1906: 722), Garman (Mem. Harv. Mus. comp. Zool., 5«, 1913),
Rey (Fauna Iberica Peces, r, 1928), White (Bull. Amer. Mus. nat. Hist., 74 [2], 1937) and Bertin (Bull. Inst.
oceanogr. Monaco, 775, 1939).

31. Bull. Amer. Mus. nat. Hist., 74, 1937: 100, loi. 32. Bull. Inst, oceanogr. Monaco, 775, 1939.

33. For a recent discussion of inter-relationships of modern sharks, with resultant proposals as to classification, see
Bertin (Bull. Inst, oceanogr. Monaco, 775, 1939).

34. Fauna Iberica Peces, /, 1928: 280. 35. Mem. Harv. Mus. comp. Zool., 36, 191 3.

Fishes of the Western North Atlantic 77

except that Chlamydoselachus and the Pristiophoridae are each made a distinct suborder
for the reasons stated below (pp. 94, 532).

Key to the Suborders of Modern Sharks
I a. Anal fin present.

2a. 6 or 7 gill openings.

3a. Margins of ist gill openings not continuous across throat 5 upper and lower
teeth notably unlike toward center of mouth. Notidanoidea, p. 77.

3b. Margins of ist gill openings continuous across throat} upper and lower teeth
similar in center of mouth as well as along its sides.

Chlamydoselachoidea, p. 92.
2b. Only 5 gill openings.

4a. Dorsal fins preceded by stout spines j teeth toward center of mouth in each
jaw markedly different from those toward its corners.

Heterodontoidea,°" p. 94.
4b. Dorsal fins not preceded by spines; teeth toward center of mouth of same
basic type as those toward its corners. Galeoidea, p. 95.

lb. No anal fin.

5a. Snout of only moderate length, without lateral teeth or barbels.

6a. Trunk subcylindrical ; eyes lateral; anterior margins of pectorals not over-
lapping gill openings. Squaloidea, p. 449.
6b. Trunk much flattened dorsoventrally; eyes dorsal; anterior margins of pec-
torals far overlapping gill openings. Squatinoidea, p. 533.
5b. Snout greatly elongate, as a narrow beak, armed on either side with sharp teeth,
and with a long fleshy barbel. Pristiophoroidea," p. 5 3 2 .

Suborder NOTIDANOIDEA

Characters. Anal fin present; only one dorsal fin, without spine; either 6 or 7 gill
openings, all anterior to pectorals; margins of ist gill openings not continuous across
throat; snout not beak-like, without lateral teeth or barbels; upper and lower teeth toward
center of mouth widely dissimilar, but essentially similar to those toward corners. Trunk
subcylindrical (shark-like); eyes lateral; anterior margins of pectorals not expanded for-
ward beyond ist gill opening; nostrils separate from mouth, anterior margins without
barbels; eye without nictitating fold or membrane; spiracles present; segmentation of ver-
tebral column incomplete, but centra more or less differentiated, with axial canal somewhat
contracted in its passage through them; notochord partially constricted segmentally in

35a. For a recent account of this group, see Smith, B. G. (Dean Memor. Vol., Amer. Mus. nat. Hist., 8, 1942).

36. These "Saw Sharks" superficially resemble the true "Sawfishes" (Pristidae) which, however, fall among skates

and rays (Batoidei), they having ventral gill openings as well as the edges of the pectorals attached to the sides

of the head anterior to the gill openings.

78 Memoir Sears Foundation for Marine Research

correspondence, more strongly so posteriorly than anteriorly; vertebral centra either with-
out calcification, or those in the tail region with calcareous lamellae radiating from a cen-
tral ring in some forms; neural spines not attached to dorsals; cranium on each side with a
well developed antorbital process, continuous, however, with the auditory capsule; rostral
cartilage single; upper jaw (palatoquadrate cartilage) attached to cranium at two points
{i.e., to the suborbital region and to a postorbital process), but not to the hyomandibular
arch which is much reduced; propterygial cartilage of pectoral without radials; heart
valves in 4 or 5 rows. Development ovoviviparous.

Families. One modern family known, Hexanchidae.

Family HEXANCHIDAE

Characters. Either 6 or 7 gill openings; margins of all gill openings widely inter-
rupted at throat; eyes without nictitating folds or membranes; spiracles present; upper
teeth sharp, with slender, curved, primary cusps; lower teeth blade-like, quadrate or trian-
gular, their margins with several small cusps; caudal fin with well marked subterminal
notch, its axis raised but little; inner margins of pelvics either separated or briefly united
posterior to cloaca; no precaudal pits; dorsal fin posterior to pelvics; lower jaw with either
large or rudimentary labial furrows ; no upper labial furrows ; clasper of male largely en-
closed by a leaf-like expansion of the pelvic fin, its axial cartilage small, simple distally, and
attached to the basipterygial cartilage of fin by 2 small connecting segments.^ Development
ovoviviparous.

Key to Genera

la. 6 gill openings. //^;c«»c^«j Rafinesque, 1810, p. 78.

lb. 7 gill openings.

2a. Head narrow; snout tapering; horizontal diameter of eye considerably greater

than distance between nostrils. Heptranchias Rafinesque, 18 10, p. 87.

2b. Head broad; snout broadly rounded; horizontal diameter of eye considerably

smaller than distance between nostrils. Notorynchus Ayres, 1885.

Mediterranean, South Africa,
Argentina, California to Oregon,
Japan, China, Australia— New
Zealand, Indian Ocean.

Genus Hexanchus Rafinesque, 1 8 1 o
Six-gilled Sharks, Cow Sharks, Mud Sharks

Hexanchus Rafinesque, Caratt. Gen. Spec. Sicil., 18 10: 14; type species, H. griseus Rafinesque, Sicily, equiva-
lent to Squalus griieus Bonnaterre, 1 788.

Generic Synonyms:

Monofterhinus (in part) Blainville, Bull. Soc. philom. Paris, 1816: 121.

I. For illustrations of the cartilag-es of the clasper, see Huber (Z. Wiss. Zool., 70, 1901 : pi. 27, fig. i) and Daniel
(Elasmobranch Fishes, 1934: 51).

Fishes of the Western North Atlantic 'jc)

Notidanus Cuvier, Regne Anim., 2, 1817: 128; type species, Squalus griseus Bonnaterre, 1788, designated by

Jordan, Genera Fish., I, 1 91 7: 97.
NotiJamus Miinster, Beitr. Petrefak., 5, 1842: 66; evident misspelling for Notidanus Cuvier, 1 81 7.
Hexancus L. Agassiz, Nomencl. Zool. Index, 1846: 181 ; equivalent to Hexanchus Rafinesque, 1810.

Generic Characters. Six gill openings, decreasing in length from front to rear; snout
short, rounded; mouth very large, mostly lateral; lower labial furrow well developed;
upper lip widely expanded posteriorly; spiracles small, situated far behind eye; fins of
moderate size; anterior upper teeth slender, pointed; anterior lower teeth broad, quad-
rate; no median upper tooth; lower median tooth present or absent. Characters otherwise
those of the family and suborder.

Range. Both sides of North Atlantic, Mediterranean, Argentina, southern Indian
Ocean, Island of Reunion, Natal and Agulhas Bank, Japan, west coasts of North and South
America. All known representatives of the genus appear to belong to a single wide-ranging
species.

Fossil Teeth. Middle Jurassic to Pliocene, Europe; Upper Cretaceous, western Asia,
New Zealand, Madagascar; Upper Cretaceous to Oligocene, South America; Eocene,
Africa; Miocene, North America.

Species. The representatives of this genus from diflFerent seas resemble one another
closely. But opinions have differed as to whether H. corinus Jordan and Gilbert' of the
Pacific coast of the United States is separable from the well known Six-gilled Shark
{griseus) of Europe. Supposedly, corinus is set apart from griseus by the fact that its
lower teeth other than the median are finely serrate along their inner edges. Actually,
however, no diflFerence exists in this respect between the populations of the two geo-
graphic regions in question, for the lower teeth of the European griseus were long ago
excellently pictured' and described* as having finely serrate inner edges, although this
fact seems to have been overlooked in some of the more recent accounts." On the other
hand, it is expressly stated in the original account of corinus^ that in a small specimen
from Puget Sound the lower teeth were smooth-edged, as they are also in a 2^ -foot
griseus from the Mediterranean that we have examined;^ and our comparison of the latter
with a four-foot specimen from Puget Sound reveals no significant difference in any other
respect. We therefore follow Regan^ and Fowler" in referring corinus, as well as the
Japanese Hexanchus, to griseus. There is nothing in the descriptions or obviously general-
ized illustration to suggest otherwise for the Chilean edulis.^"

2. Proc. U.S. nat. Mus., 3, 1880: 352. 3. L. Agassiz, Poiss. Foss., 3, 1835; pi. e, fig. 2-4.

4. Miiller and Henle, Plagiost., 1841 : 81.

5. Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 17), for example, desoribes them without qualifications as
smooth-edged, and Rey (Fauna Iberica Peces, /, 1928: fig. 83) so pictures them.

6. Jordan and Gilbert, Proc. U.S. nat. Mus., 5, 1880: 352.

7. Similarly, the lower teeth are smooth-edged in a 429-mm. specimen from Cuba; the first large lower tooth is
serrate but the others smooth in one of about five feet ; all the large lateral lower teeth are more or less serrate
along their inner margins in one of 1 1 feet (see Study Material, p. 80) .

8. Ann. Mag. nat. Hist., (7) 16, 1905: 57. 9. Bull. U.S. nat. Mus., 100 (/j), 1941: 57-
10. Perez-Canto, Estud. Escual. Chile, 1886; 8; Philippi, An. Univ. Chile, 71, 1887: 554, pi. 6, fig. i.

8o Memoir Sears Foundation for Marine Research

Hexanchus griseus (Bonnaterre), 1780

Six-gilled Shark, Cow Shark, Grey Shark, Mud Shark

Figures 8, 9

Study Material. Female, 830 mm., from Nice, France (Harv. Mus. Comp. Zool.,
No. 946); male, 429 mm., from off Havana, Cuba, apparently newborn, without trace
of umbilical scar (Harv. Mus. Comp. Zool., No. 35630) ; male, 4 feet, Puget Sound (U.S.
Nat. Mus., No. 104474); male, 1,167 mm-j from off Pacific Beach, California (Harv.
Mus. Comp. Zool., No. 36474) ; also jaws of an i i-foot and of a 5-foot specimen (Harv.
Mus. Comp. Zool., No. 36217, 36216) and of one of 10 feet 2 inches from N. Carolina
(U.S. Nat. Mus., No. 37790).

Distinctive Characters. The presence of six gill openings, combined with the facts
that the lower ends of the members of each pair are widely separated one from the other
in the region of the throat, and that the upper teeth are strikingly unlike the lowers,
separates this species from all other sharks of the North Atlantic.

Description. Proportional dimensions in per cent of total length. Male, 429 mm.,
from Cuba (Harv. Mus. Comp. Zool., No. 35630). Female, 830 mm., from Nice, France
(Harv. Mus. Comp. Zool., No. 946).

Trunk at origin of -pectoral: breadth 10.7, 9.8 ; height 8.2, 7.8.

Snout length in front of: outer nostrils 1.6, 1.7; mouth 7.0, 5.0.

Eye: horizontal diameter: 4.7, 3.0.

Mouth: breadth 9.8, lO.O; height 6.3, 6.1.

Nostrils: distance between inner ends 4.4, 4.6.

Gill opening lengths: ist 6.S, 7-5; 2nd 5.4, 6.Ss 3rd 5.1, 5.8; 4th 4.7, 5.2; 5th

3.4,4.3; 6th 3.5, 4.1.

First dorsal fin: vertical height 3.7, 4.8 ; length of base 5.8, 6.6.

Anal fin: vertical height 1.9, 3.5; length of base 4.4, 6.4.

Caudal fin: upper margin 35.2, 31.3; lower anterior margin 10.2, 8.3.

Pectoral fin: outer margin 13.O, 13.2; inner margin 5.2, 6.0; distal margin lO.O,

10.7.

Distance from snout to: ist dorsal 46.7, 54.1 ; upper caudal 64.8, 68.7; pectorals

20.5, 21.3; pelvics 38.5, 44.6; anal 50.7, 57.2.

Interspace between: ist dorsal and caudal 10.9, 9.0; anal and caudal 6.$, 5-2.

Distance from origin to origin of: pectorals and pelvics 20.0, 25.4; pelvics and anal

12.3,14.4.

Trunk moderately stout anteriorly, its depth opposite origin of pectorals Vr to Vs as
great as its length to origin of caudal fin, tapering rearward and strongly compressed later-
ally posterior to pelvics. Caudal peduncle without precaudal pits. Lateral line clearly
visible as a pale streak from opposite last gill opening rearward out onto caudal and dip-
ping downward abruptly on anterior sector of caudal. Denticles on sides of trunk (Fig.

Fishes of the Western North Atlantic

8i

8 E) usually tridentate (occasionally with one of the lateral points lacking), with a
prominent axial crest and two (occasionally only one) lower lateral crests, loosely to mod-
erately-closely spaced and overlapping but little j those along rear half of upper margin
of caudal fin much larger than on trunk, smooth, ovoid in outline, forming a visible ridge.
Head flattened above. Snout broadly rounded and short, its length in front of mouth

Figure 8. A-E Hexanchus griseus, female, 830 mm. long, from Nice, France (Harv. Mus. Comp. Zool.,
No. 946), with dermal denticles, about 18 x. f Left-hand upper and lower teeth of an 1 1 -foot Cuban speci-
men, about Y2 natural size.

Figure 9. A Hexanchus griseus, new-born female, 429 mm. long, from Havana, Cuba (Harv. Mus. Comp.
Zool., No. 35630). B First lower tooth of the Il-foot Cuban specimen illustrated in Fig. 8f, about 1.5 x.

82 Memoir Sears Foundation for Marine Research

only about Y^ as great as breadth of mouth. Eye oval and noticeably large, its horizontal
diameter about Yj as great as length of head in a 2^/'o-foot specimen, but perhaps relatively
smaller in adults." Spiracle very small, at level of upper margin of eye, about opposite
corner of mouth. Gill openings notably long, extending from high on sides far onto ven-
tral surface of throat, and noticeably oblique, the ist (longest) about 2V2 times as long as
horizontal diameter of eye, successively shorter, rearward, the 6th only a little more than
Y2 as long as the istj the inner margins of 2nd and 3rd gill arches with 2, and 4th to 6th
with 3 to 5, fleshy tubercles, suggesting rudimentary rakers. Nostril much nearer to tip
of snout than to mouth, small, strongly oblique, its anterior margin expanded as a sub-
triangular lobe with blunt tip. Mouth notably large, about % as high as broad, crescentic
and inferior anteriorly, but extending along sides of head for most of its length, the gape
reaching rearward about % of distance to origin of pectoral. Well developed labial furrow
at corner of mouth on lower jaw, visible only when mouth is partly open; none on upper
jaw. Upper lip enclosing posterior part of lower jaw as a free fold extending rearward
past corner of mouth for a distance about equal to horizontal diameter of eye.

Teeth — — ; noticeably diflFerent in the 2 laws; ist 2 to

i2toi6 — I (or 0) • — i2toi6 ' •'

4 uppers simple with slender median cusp curved outward, the ist noticeably smaller, the
2nd slightly smaller and with narrower bases than 3rd, the subsequent teeth to the lOth
or nth with i, 2 or 3 short basal cusps on outer side (number increasing toward corner of
mouth and with growth), the outermost 7 or 8 teeth rounded, with only very small cusp
or none, and much lower than the others. Lower jaw usually with i symmetrical median
tooth, having i median cusp and i, 2 or 3 lower cusps on each side," the next 6 (occasion-
ally 5) teeth trapezoid, about twice as broad as high, with 7 to 8 pointed cusps in small
specimens, increasing in number to 8 to 10 in large, the innermost cusp the longest, the
others progressively shorter, the inner margins smooth in newborn specimens, but finely
serrate in large, with intermediate sizes showing intermediate states,'^ the 7th lower tooth
(6th in specimens which have only 5 large laterals) much smaller, with only i definite
cusp, the outermost 4 to 6 teeth very low, rounded, without cusp ; 2 or 3 series functional in
center of upper jaw and i along its sides; i series functional in lower jaw.

Vertical fins small. Dorsal with rounded apex and weakly convex rear margin, free
rear tip broadly triangular, about half as long as base, its origin slightly behind cloaca,
the midpoint of its base about over origin of anal. Caudal about Vs of total length, with
well marked subterminal notch, its lower anterior corner expanded as a low rounded lobe
in newborn but not appreciably so in larger specimens (cf. Fig. 8 A and 9 A), its maximum
breadth a little more than Ys its length. Anal about as long at base as ist dorsal, rear margin
nearly straight, free rear tip short. Interspace between rear end of base of anal and origin
of caudal about Y2 as long as between dorsal and caudal. Pel vies with nearly straight mar-

1 1. The eye, as in many sharks, is relatively larger in newborn than in older specimens.

II. This median tooth is lacking in the small Mediterranean specimen listed under Study Material, p. So.

13. For further details, see discussion, p. 79.

Fishes of the Western North Atlantic 83

gins and rounded apices, their bases a little less than 1V2 times as long as base of dorsal,
their inner margins united for a very short distance posterior to cloaca in female, but
entirely separate in male, and partially enclosing the claspers, with their tips somewhat
elongate. Pectoral about i V3 times as long as broad, with broadly rounded apex, nearly
straight distal margin (moderately concave in smaller specimen) and very broad base, the
inner margin nearly or quite ^o as long as anterior margin.

Color. Fresh specimens are variously described as coffee-colored (darkest along the
midline of back) or as very dark gray above; at least some specimens with a pale streak
along the side; lower surface of a paler shade of the same hue, or whitish. Of the preserved
specimens we have seen, a s^/'o-foot Mediterranean specimen is dark chocolate brown above
and paler below; one newborn from Cuba is mouse gray above and paler below; a 4- foot
Puget Sound fish is very dark gray, hardly paler below than above.

Size. The fact that embryos of 650 mm. have been reported, with free-swimming
young as small as 429 to 700 mm., points to a length of 16 to 26 inches at birth. Maturity
is to be expected at a length of perhaps 6 to 6Y2 feet, large numbers of eggs having been
found in females of about 7 feet in Cuban waters. The recorded lengths of the larger
adults have ranged up to about 15V2 feet (4.82 m.). One of 26 feet 5 inches was reported
from Cornwall many years ago, a giant of its kind if its size was stated correctly." Re-
ported weights are about 220 lb. ( 100 kilo.) at 7 ft. 4 in. (2.25 m.) ; about 300-400 lb. at
about 9 ft.; about 528 lb. (240 kilo.) at about 13 ft. (4 m.); 1,085 lb. at 14 ft.; and 1,300
lb. at lift.'"

Developmental Stages. The litters are certainly large, for 47 embryos were counted
in a female of 4.8 meters,'^ while fishermen have reported as many as 108 in a specimen
slightly smaller (4.5 meters long)."

Habits. This shark is described as sluggish, hardly resisting when caught on hook and
line, but we have had no experience with it alive. Characteristically, it is a "ground" spe-
cies, usually living in at least moderately deep water. It is recorded from depths as great
as 800 to 1,875 meters off Portugal, at a little deeper than 100 fathoms off the Irish slope,
at about 90 to 560 fathoms in Scottish waters, including the Shetland-Faroe Channel, and
usually from 75 fathoms down to 300 fathoms off Cuba, where, in fact, few are caught
shoaler than 100 fathoms. And we have received photographs from Ollyandro del Valle of
three large ones (922, 1,400 and 1,682 pounds) taken in the deep water shark fishery
off the north coast of Cuba, said to have been hooked at 700 fathoms. On the other hand,
one of the earliest recognizable reports of it" was of a specimen from the coast of Holland
and therefore certainly from shoal water. Scattering specimens are caught in the North
Sea, in depths certainly no greater than 1 5 to 20 fathoms, and they have even been seen
swimming at the surface off Ireland. It has been suggested that this shark lies quiescent on

14. Day, Fish. Gt. Brit., 2, 1880-1884: 308; this size has been frequently quoted since then for this species.
14a. Cuban specimens, reported by Luis Howell-Rivero. 15. Bolivar, Bol. See. esp. Hist, nat., 7, 1907: 207.

16. Vaillant, Bull. Mus. Hist. nat. Paris, 1901 : 202.

17. Spallanzani, Viag. Sicil., 4, 1793: Chap. 31, pi. 2.

84 Memoir Sears Foundation for Marine Research

bottom by day, visiting the upper waters at night in search of food.'' High temperatures
probably act as a barrier to it toward the surface and inshore in the warmer part of its range,
as in the Mediterranean and around Cuba.

The Six-gilled Shark was long ago reported as mating in spring and autumn and pro-
ducing young at various seasons, but on how good evidence we cannot say.

Their food consists of fish and various crustaceans. In Spanish waters it feeds largely
on hake (Merluccius) ; an entire torpedo has also been found in one. Off Cuba, dolphins
(Coryphaena), small marlins (Makaira) and small swordfish (XipMas) are reported
from stomachs, as well as crabs, shrimps and parts bitten from other sharks that had been
hooked.'" They are described as coming to the surface on occasion to pick up fish thrown
overboard.

Relation to Man. This species is not sufficiently abundant in American waters to be of
any special importance, although such as are taken off Cuba are utilized for their oil. In
the North Sea, any that are caught are marketable in Germany, even though the flesh has
been credited with a purgative action. However, along the Iberian Peninsula, and in the
Mediterranean, where it is much more plentiful, it is of no commercial importance, except
as a nuisance to fishermen, since it drives away merchantable fishes.

Range. Continental waters on both sides of the Atlantic, including the Mediterra-
nean} also Pacific coast of North America from southern California to British Columbia}
Chile; Japan; Australia; southern Indian Ocean and South Africa.

Occurrence in the North Atlantic. On the eastern side of the Atlantic, although no-
where abundant, the center of population for this Shark appears to be in the Mediter-
ranean, where it is widespread, and thence northward along the Atlantic coasts of the
Iberian peninsula and France. It also enters the North Sea in numbers sufficient for fisher-
men to be familiar with it; it is taken from time to time on the south coast of England,
along the Irish Atlantic slope, off western Scotland to the Faroe-Shetland Channel, and
even as a stray off Iceland. To the southward it has been reported from Morocco to Mauri-
tania.

Occurrence in the Western North Atlantic. It has long been known that the Six-gilled
Shark occurs off the northern coast of Cuba, specimens being caught from time to time near
Matanzas and Havana, and since the recent development of a hook and line fishery at lOO
to 400 fathoms or deeper it has proved to be more plentiful there in deep water than was
formerly supposed, large specimens being taken daily."" However, for it to stray north-
ward must be a very rare event, the only record of its occurrence on the east coast of con-
tinental North America being a ten-foot two-inch specimen taken in March 1886 on the
coast of North Carolina near Currituck Lighthouse. Neither is there any evidence of its
presence anywhere in the Gulf of Mexico and Caribbean region, other than for Cuba.
But it is to be expected there, at appropriate depths, and along the coast of South America
generally, if a report of it from northern Argentina be well founded.''

18. Fraser-Brunner, Proc. R. Irish Acad., 42, B-9, 1935: 519.

19. Communication from Luis Howell-Rivero. zo. Communication from Luis Howell-Rivero.
21. Lahille, An. IVIus. nac. B. Aires, 24, 1913: 26, 32 (identified by the teeth).

Fishes of the Western North Atlantic 85

Synonyms and References:

I. Atlantic Ocean and South Africa:

Le Griset, Broussonet, Mem. math. phys. Acad. Sci. Paris, 1780: 663 (descr., Medit.).

Squalus griseus Bonnaterre, Tabl. Encyc. Meth. Ichthyol., I 788: 9 (descr., type loc. Medit.) ; Gmelin, in Lin-
naeus, Syst. Nat., 1789: 1495 (descr.); Bloch and Schneider, Syst. Ichthyol., 1801: 129 (Medit.); La-
treille, Nouv. Diet. Hist. Nat., 24, 1804: 72 (in table of contents); Risso, Ichthyol. Nice., 1810: ^7
(descr., Medit.); Nacarri, Ittiol. Adriat., 1822: 24 (Adriatic, not seen); Nardo, Oss. Agg. Adr. Itiol.,
Giorn. Fis. Nat. Pavia, 7, 1824: 261 (not seen); Martens, Reise Vened., 2, 1824: 408 (Medit., not
seen); Nardo, Prod. Ichthyol. Adriat., 1827: 9 (Adriatic); Trapani, Cat. Fish Malta, 1838: 16 (Malta,
not seen).

Squalo (not named), Spallanzani, Viag. Sicil., 4 (31), 1793: pi. 2 (jaws, Holland).

Le squale griset, Lacepede, Hist. Nat. Poiss., 4° ed., i, 1798: 167, 269, in BuflFon, Hist. Nat.; in Sonnini,
Hist. Nat. Poiss., 4, i 801-1 802: 96 (descr.).

Squdlus z'acca Bloch and Schneider, Syst. Ichthyol., 1801: 138.

Hexanchus gr'iseu! Rafinesque, Carratt. Gen. Nuov. Sicil., 1810: 14 (descr., Sicily) ; Indice Ittiol. Sicil., 1810:
47 (Sicily); Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1839: 316 (descr.); Muller and Henle,
Plagiost., I 841: 80 (descr., Medit., Atlant.) ; Heckel, Pesci Dalmat., in Carrara Opera, 1846: 91 (Medit.,
not seen) ; Busch, Selach. Ganoid. Encephalo., 1848: pi. 3, fig. 8 (brain) ; White, List Spec. Brit. Mus.,
Fish., 1851: 130 (Isle of Wight); Costa, Fauna Napol. Pesci Cat. Sist., 2-j, 1854-1857: 15, 16 (39, 40)
(Medit.); Canestrini, Cat. Pesci Genov., Arch. Zool. Anat. FisioL, /, 1861: 267 (Medit.); Guichenot,
Notes Isle Reunion, 2, 1 841 : 30 (I. Reunion, not seen) ; Dumeril, Hist. Nat. Poiss., r, 1865: 431, pi. 4,
fig. 9-12 (teeth); Poey, Repert. Fisico-Nat. Cuba, 2, 1 868: 454 (Cuba); Gray, Ann. Mag. nat. Hist.,
(4) I, 1868: 76 (Cornwall); Brito Capello, J. Sci. math. phys. nat. Lisboa, 2, 1870: 140 (Portugal);
Miklucho-Maclay, Beitr. Vergl. Neurol. Wirbelt., /, 1 870: 13, pi. 2, fig. 8-14 (brain) ; Ninni, An. Soc.
Nat. Modena, 5, 1 870: 66 (Venice) ; Costa, La Pesca., Napoli, 1 871 : 87 (Medit., not seen) ; Gegenbaur,
Unters. Vergl. Anat. Wirbelt., 5, 1872: pi. 3, fig. 6, 7 (skelet.) ; Poey, An. Soc. esp. Hist. Nat., 5, 1876:
398; also Sep., Enumerat. Pise. Cubens., 1 875: 202 (Cuba) ; Brito Capello, Cat. Peix. Portugal, 1880: 45
(Portugal); Doderlein, Man. Ittiol. Medit., 2, 1 881: 76^^ (Medit., distrib. in Atlant.); Moreau, Hist.
Nat. Poiss., France, /, 1881: 336 (descr., distrib., Medit. and Atlant.); Carus, Prod. Fauna Medit., 2,
1889— 1893: 499 (Medit.); Almeida and Roquete, Mammif. Peix. Costa e rios do Algarve, Inquerito
Industr. Lisboa, 1892: 372 (Portugal, not seen) ; Jordan and Evermann, Rep. U.S. Comm. Fish. (1895),
1896: 213 (distrib.) ; Bull. U.S. nat. Mus., 47 (1), 1896: 19 (descr., Cuba) ; Carruccio, Boll. Soc. Roma
Zool., 5, 1896: 165, I pi. (anat., Medit.) ; Vieira, Ann. Sci. nat. Porto, 4 (2), 1897: 67 (Portugal, size) ;
Sicher, Atti Accad. gioenia., (4) 11 (5), 1 898: 16 (Medit.) ; Jordan and Evermann, Bull. U.S. nat. Mus.,
47 (4), 1 900: pi. 2, fig. 8: Huber, Z. wiss. Zool., 70, 1901 : 600, pi. 27, fig. 1,2 (claspers) ; Vaillant, Bull.
Mus. Hist. nat. Paris, 7, 1901: 202 (Gulf of Gascony, embryos); de Braganza, Result. Invest. Sci.
"Amelia," Ichthyol., 2, 1904: 28 (Portugal) ; Regan, Ann. Mag. nat. Hist., (7) /<5, 1905: 571 (concludes
corinus equals griseus) ; Smith, Bull. N.C. geol. econ. Surv., 2, 1907: 30 (N. Carolina specimen) ; Fow-
ler, Proc. Acad. nat. Sci. Philad., 60, 1908: 52 (spec, no data) ; Diaz y Martinez, An. Acad. Habana, 46,
1910: 82 (Cuba); Laboissiere, Bull. Ass. Levall.-Perret, 5, 1910: 6 (Gulf of Gascony); Seabra,
Poiss. Port., 1911:195 (Portugal, not seen) ; Roule, Bull. Inst, oceanogr. Monaco, 243, 1912:3 (Medit.);
Garman, Mem. Harv. Mus. comp. Zool., j6, 1 91 3: 16 (descr.) ; Lahille, An. Mus. nac. B. Aires, 24, 1 91 3:
26, 32 (Argentina); Scharff, Irish Nat., 24, 1915: 99 (Ireland); Smith, J. Amer. Mus. nat. Hist., 16,
1916: 349, 351 (N. Carolina); Lahille, Physis B. Aires, 5, 1921: 63 (spec, recorded, Argentina); Nor-
man, Ann. Mag. nat. Hist., (9) p, 1922: 319 (Natal, S. Africa); Marelli, Elenc. Sist. Fauna, Mem.
Ministr. Obras Publ. B. Aires (1922-1923), 1924: 544 (Argentina); Barnard, Ann. S. Afr. Mus., 2/
(l), 1925: 22 (S. Africa); Rey, Fauna Iberica Feces, 7, 1928: 292 (descr., off Spain); Hickling, Ann.
Mag. nat. Hist., (10) 2, 1928: 199 (NE. Atlant.) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish.
(1928), 2, 1930: II (name, distrib.); Fowler, Proc. Pan-Pacif. sci. Congr. (1929), j, 1930: 484
(name); Sanchez-Roig, Revist. Agric. Feces Cubana, Commerc. Trabaj., j (9), 1 931: 18 (Cuba);
Baylis, Ann. Mag. nat. Hist., (10) 12, 1933: 322 (parasites); Belloc, Rev. des. trav. Peches Marit., 7,

XI. See Doderlein, 1881, for extensive list of Mediterranean records, some in publications not accessible to us.

86 Memoir Sears Foundation for Marine Research

Fasc. 2, 1934: 151 (ill., Morocco, Rio de Oro, Mauritania) ; Borri, Atti Soc. Tosc. Sci. nat., 44, 1934.: 94
(Medit.) ; Fraser-Brunner, Proc. R. Irish Acad., 42 (B), 1935: 319 (Irish Atlant. slope) ; Pozzi and Bor-

dale, An. Soc. cient. argent., 120, 1935: 149 (listed for Argentina) ; Nobre, Fauna Marinha Port. Vert.,
/, 1935: 413, pi. 61, fig. 189 (Portugal); Liibbert and Ehrenbaum, Handb. Seefisch. Nordeurop., 2,
1936: 272, pi. 19, fig. 239 (general; E. Atlant.) ; Howell-Rivero, Proc. Boston Soc. nat. Hist., 41, 1936:
42 (Cuba) ; Fowler, Bull. Amer. Mus. nat. Hist., yo (i), 1936: 26 (descr., distrib.) ; Norman and Eraser,
Giant Fishes, 1937: 5 (general); Desbrosses, Rev. des. Trav. Peches Marit., 11, Fasc. I, 1938: 53
(growth, migrations) ; Tortonese, Atti Soc. ital. Sci. nat., 77, 1938: 286 (Medit.) ; Howell-Rivero, Tor-
reya, 9, 1941: 4, pi. I, 2 (descr., Cuba); Fowler, Bull. U.S. nat. Mus., 1 00 {13), 1941: ii( part);
Holmgren, Acta Zool. Arg., 22, 1941: 9 (skelet.) ; Bigelow and Schroeder, Guide Comm. Shark Fish.,
Anglo Amer. Carib. Comm., Wash., 1945: 94, fig. 31 (descr., ill., habits, range).

Monopterhinus griseus Blainville, Bull. Soc. philom. Sci. Paris, 1816: 121.

Notidanus griseus Cuvier, Regne Anim., 2, 1817: 128 (general) ; 2nd Ed., 1829: 390; Bory de St. Vincent,
Diet. C1.1SS. Hist, nat., 75, 1829: 597 (general), Voigt, in Cuvier, Tierreich, 2, 1832: 509 (descr.);
Bonaparte, Fauna Ital. Pesc, 5 (2), 1835: pi. [55], fig. I (descr., Medit.); Agassiz, L., Poiss. Foss., 3,
1835-1838: 92, pi. E, fig. 2-4 (teeth); Lowe, Trans, zool. Soc. Lond., 2 (3), 1837: 194 (Madeira);
Bonaparte, Mem. Soc. neuchatel. Sci. nat., 2 (8), 1839: 9 (in synopsis); Cuvier, Regne Anim., Poiss.,
1843: 362, pi. 115, fig. I (teeth); Couch, Zoologist, 4, 1846: 1337 (Cornwall); Bonaparte, Cat. Pesc.
Europ., 1 846: 17 (name) ; Sassi, Cat. Pesci Legur., 1 846: I 3 1 (Medit., not seen) ; Nardo, Sinon. Modern.
Spec, descr. Pesci St. Chiereghin, 1847: ill (name) ; Owen, Cat. Osteol. Roy. Coll. Surg., i, 1853:91
(teeth) ; Machado, Feces Cadiz, 1857: 8 (near Cadiz); Nardo, Atti 1st. veneto., (3)5, 1 8 59-1 860: 787
(Medit.); Gemellaro, Atti Accad. gioenia, (2) ig, 1864: 122 (Medit., not seen); Bocage and Brito
Capello, Poiss. Plagiost., 1866: 15 (Portugal) ; de la Blanchere, Diet. Peches, 1868:371 (descr., Medit.) ;
Gunther, Cat. Fish. Brit. Mus., 8, 1870: 397 (descr., distrib.); Canestrini, in Cornalia, et al.. Fauna
d'ltal., 1872; 43 (Medit.); Lawley, Monog. Gen. Notidanus foss., 1875: 60, 65, pi. 4, fig. I (jaws);
Gervais and Boulart, Poiss., 3, 1877: 194, pi. 74, fig. 18 (descr.); Delfortrie, Act. Soc. linn. Bordeaux,
(4) 2, 1878: 253 (teeth) ; Doderlein, Prosp. Metod. Pesci SiciL, 1878: 30 (Sicily) ; Day, Fish. Gt. Brit.,
2, 1 880-1 884: 308, pi. 158, fig. 2 (descr., habits, Gt. Brit.); Stossich, Boll. Soc. adriat. Sci. nat., 5,
1880: 69 (Adriatic) ; Perugia, Elenc. Pesc. Adriat., 1881 : 55 (Adriatic) ; Graeffe, Arb. zool. Inst. Univ.
Wien, 7, 1886: 447 (Medit.) ; Gunther, Proc. roy. Soc. Edinb., 75, 1888: 207 (Orkneys and Shetlands) ;
Anderson, Ann. Scot. nat. Hist., 9, 1894: 182 (W. Scot.); Traquair, Ann. Scot. nat. Hist., 5, 1896: 159
(W. Scot.); Philippi, Ann. Univ. Chile, 109, 1902: 304; Bolivar, Bol. Soc. esp. Hist, nat., 7, 1907
(Spain); Gunther, Encycl. Brit., 24, 191 1 : 809 (general) ; Ehrenbaum, in Grimpe and Wagler, Tier-
welt N.- u. Ostsee, Lief 7 (r2*), 1927: 7 (general, North Sea).

Nolidanus vacca Cuvier, Regne Anim., 2, 1817: 28.

Squalus {Mono-pterhinus) griseus Blainville, in Vieillot, Faune Franc, 1825: 77 (descr., Medit.).

Notidanus monge Risso, Hist. nat. Europ. merid., 3, 1826: 129 (general, Medit.) ; Bory de St. Vincent, Diet.
Class. Hist, nat., i^, 1829: 597.

Grey Notidanus, Yarrell, Brit. Fish., 3rd Ed., 2, 1859: 515 (not seen).

Notidanus sf. dubia Poey, Memorias, 2, 1861: 359 (Cuba).

Six-gilled shark, Couch, Fish. Brit. Isles, /, I 867: 21, pi. 4 (England).

Notidanus (Hexanchus) griseus Werner, Zool. Jb., Syst. Abt., 2/, 1904: 286; Ehrenbaum, Faune Ichthyol.,
Cons, explor. Mer, 1930: plate not numbered (general).

2. Pacific:

Hexanchus corinus Jordan and Gilbert, Proc. U.S. nat. Mus., 3, 1880: 352 (descr., Monterey Bay and Puget
Sound) ; Bean, Proc. U.S. nat. Mus., 4, 1882: 267 (listed for Puget Sound) ; Jordan and Gilbert, Proc.
U.S. nat. Mus., 4, 1882: 30 (listed for Soquel, Calif., and Puget Sound) ; Jordan and Gilbert, Bull. U.S.
nat. Mus., 16, 1883: 62 (descr.); Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 792 (listed for Pa-
cific coast, U.S.) ; Jordan and Starks, Proc. Calif. Acad. Sci., (2) 5, 1895: 788 (listed for Puget Sound) ;
Jordan and Evermann, Bull. U.S. nat. Mus., 47 (l), 1896: 19 (descr., Monterey, California, to Puget
Sound); Rep. U.S. Comm. Fish. (1895), 1896: 213 (listed, Monterey Bay to Puget Sound); Starks,

Fishes of the Western North Atlantic 87

Ann. Carneg. Mus., 7, 191 1 : 163 (cf. with griseus; color; San Francisco, Puget Sound) ; Garman, Mem.
Harv. Mus. comp. Zool., 36, 191 3: 17 (descr.) ; Halkett, Check List Fish. Canad., 1913: 39 (listed Van-
couver I. and Puget Sound to Monterey) ; O'Donoghue, J. Anat., Lond., 61, 1 926: 41 (descr., meas., ill.,
teeth, denticles; anat.; photo of Nanaimo, British Columbia spec.) ; Jordan, Evermann and Clarke, Rep.
U.S. Comm. Fish. (1928), 2, 1930: 11 (listed Monterey Bay to Puget Sound); Townsend, Bull. N.Y.
zool. Soc, ^.^ (6), 1931: 169 (photograph, Washington) ; Walford, Fish Bull. Sacramento, 45, 1936: 24
(diagn., photo., San Diego to Puget Sound) ; Barnhart, Mar. Fish south. Calif., 1936: 7 (off San Diego).

'Notidonus vulgaris Perez Canto, Estud. Escual. Chile, 1886: 8 (descr., Chile) ; Philippi, An. Univ. Chile, 7/,
1887: 554, pi. 6, fig. I (descr., ill., Chile) ; Quijada, Bol. Mus. nac. Chile, 5, 1913: 112 (listed for San
Antonio, Chile).

Hexanchus griseus Regan, Ann. Mag. nat. Hist., (7) 16, 1905: 571 (Japan, Pacific U.S. cf. with Atlantic);
Starks, Calif. Fish Game, j, 1917: 146 (listed California to Puget Sound); Hubbs, J. Pan-Pacif. Res.
Instn., J, 1928: 1 1 (listed for Oregon and Washington) ; Desbrosses, Rev. des Trav. Peches Marit., it,
Fasc. I, 1938: 53 (migrations, rate of growth, breeding season) ; Fowler, Bull. U.S. nat. Mus., lOO {13),
1941: II (descr.. Pacific refs.) ; Phillipps, Dom. Mus. rec. Zool., / (2), 1946: 5 (descr., photo. New
Zealand).

Genus He-ptranchias Rafinesque, 1 8 lO

Heftranchias Rafinesque, Carratt. Gen. Spec. Sicil., 1810: 13; type species, H. cinereus Rafinesque, Sicily,
equals Squalus ferlo Bonnaterre, 1788.

Generic Synonyms:

Monofterhinus (in part) Blainville, Bull. Soc. philom. Paris, 1816: 121.
Cijrc^arjj/ (in part) Cloquet, Diet. Sci. Nat., 7, 1817:69.
Nolidanus (in part) Cuvier, Regne Anim., 2, 1829: 390.
Heftanchus (in part) Miiller and Henle, Plagiost., 1841: 81.

H eftranchus Gray, List Fish. Brit. Mus., Chondropt., 1851 : 68, equivalent to Heftranchias Rafinesque, 1810.
Heftrancus Costa, Fauna Napoli Pesci, 3, Chondropt., 1854-57: 5 (29), equivalent to Heftranchias Rafi-
nesque, 1 8 10.
Hexanchus (in part) Seabra, Bull. Soc. Portug. Sci. nat., 5, 191 1 : 195.

Generic Characters. Seven gill openings, decreasing in length from front to rear;
snout narrow, tapering; horizontal diameter of eye considerably greater than distance
between nostrils. Characters otherwise those of the suborder and family.

Range. Eastern and western North Atlantic, South Africa, Australia, Japan.

Species. Our own comparison of medium-aged and small specimens from Japan, with
others from Cuba and from the Mediterranean (see Study Material, p. 88) corroborates
GarmanV conclusion that the North Pacific representative of the genus is identical specifi-
cally with the Atlantic form; such differences in proportionate dimensions as exist between
them are no greater than might have resulted from the fact that the larger of the Japanese
examples had been dried. However, the Australian form may be distinct, as indicated in the
following key.''

1. Mem. Harv. Mus. comp. Zool., j6, 191 3: 23.

I. Heptranchias hasiuelti Ogilby (Proc. Linn. Soc. N. S. W., 22, 1897: 62), Nolidanus medinae and N. wolnicziyi
Philippi (An. Univ. Chile, 109, 1901 : 305, 307, Chile), and A', ferox Perez Canto (Estud. Escual. Chile, 1886:
7), which are included by Fowler (Bull. U.S. nat. Mus., too [/j], 1941 : 9) in the synonymy of Heftranchias
ferlo, all appear, from the original accounts, to fall in the genus N otorynchus.

Memoir Sears Foundation for Marine Research

Key to Species

I a. Origin of anal opposite rear base of dorsal,
lb. Origin of anal under middle of base of dorsal.

perlo Bonnaterre, p. 88.
dakini Whitley, 1931.

Australia.'

He-ptranchias ferlo (Bonnaterre), 1788

Seven-gilled Shark

Figures 10, 11

Study Material. Female, 932 mm. long, containing 9 embryos, and also an adult male
of 698 mm., both from Havana, Cuba (Harv. Mus. Comp. Zool., No. 36897) ; specimen
of 732 mm., from Nice, France (Harv. Mus. Comp. Zool., No. 945)} also specimens of
957, 980 and 255 mm. from Japan, the latter newborn with umbilical scar still faintly
visible (Harv. Mus. Comp. Zool., No. 35070, 1040, 1299).

Distinctive Characters. The presence of 7 gill slits combined with narrow head and
pointed snout separates ferlo from all other Atlantic sharks.

Figure 10. Heftranchias ferlo, male, about 689 mm. long, from the north coast of Cuba (Harv. Mus. Comp.
Zool., No. 35897) ; A Anterior part of head, about Y2 natural size. B Right-hand nostril, about 1.3 x. C Der-
mal den,ticles, about 37 x. Z) Lateral view of dermal denticle, about 64 x. E Apical view of dermal denticle,
about 64 X.

3. An additional diagnostic character, according to Whitley (Aust. Zool., 6, 1931: 310), is anal base as long as
dorsal base in ferlo, but shorter than the latter in dakini. Actually, however, no distinction can be drawn in this
respect, the anal being appreciably shorter than the dorsal in two of the three Atlantic specimens of ferlo that we
have seen.

Fishes of the Western North Atlantic

89

Description. Proportional dimensions in per cent of total length. Male, 698 mm.,
from Cuba (Harv. Mus. Comp. Zool., No. 35897). Female, 932 mm., from Cuba (Harv.
Mus. Comp. Zool., No. 35897).

Figure i i. Heftranchias ferlo, A upper and lower teeth of specimen pictured in Fig. lO, about 2.4 x. B An-
terior part of upper jaw to show arrangement of teeth as viewed from below, about 2.4 x. C Embryo from
Cuba with yolk sac attached (Harv. Mus. Comp. Zool., No. 35581), about 0.4 natural size.

Trunk at origin of fectoral: breadth 8.4, 9.1 } height 9.6, 10.7.

Snout length in front of: outer nostrils 2.0, 2.0; mouth 5.0, 4.8.

Eye: horizontal diameter 4.0, 3.6.

Mouth: breadth 8.0, 7.0 5 height 6.4, 7.1.

Nostrils: distance between inner ends 2.6, 2.1.

Gill ofening lengths: ist 5.7,7.2; 2nd 5.6,6.5; 3rd 4.7, 5.9; 4th 4.2, 5.3; 5th 3.6,

4.6; 6th 3.1,4.0; 7th 2.6, 3.1.

First dorsal fin: vertical height 4.3, 4.5; length of base 6.2, 6.6.

Anal fin: vertical height 2.2, 2.7; length of base 5.9, 5.9.

Caudal fin: upper margin 30.6, 30.4; lower anterior margin 9.0, 8.6.

Pectoral fin: outer margin 11.4, 11.4; inner margin 5.0, 5.5; distal margin, 9.6,

8.5.

Distance from snout to: ist dorsal 49.0, 48.3; upper caudal 69.4, 69.6; pectoral

20.9, 19. 1; pelvics40.o, 38.4; anal 54.8, 52.2.

go Memoir Sears Foundation for Marine Research

Interspace between: ist dorsal and caudal 14.2, 14.8; anal and caudal 9.0, 9.7.

Distance from origin to origin of: pectorals and pelvics 18.9, 22.0; pelvics and

anal 15.0, 15.0.
Trunk rather slender, compressed, its height at about midsection of body, where
highest, 1 1.6 to 1 1.8% of total length, the body sector shorter than tail sector by a distance
about equal to length of pectoral. Caudal peduncle about 75 to 80% as wide as deep; no
precaudal pits. Dermal denticles on sides of trunk closely overlapping, a little longer than
broad, each denticle with prominent median tooth, flanked by a pair of much smaller lat-
erals, a strong median ridge and upturned lateral edges, the blades so thin and transparent
that the pigment dots on the skin are visible through them; those along upper margin of
caudal ovoid, without lateral marginal teeth, but with 3 longitudinal ridges, the median
subdivided posteriorly, forming an ill-defined crest, much as in Hexanchus griseus (p.

81).

Head with dorsal profile slightly convex. Snout tapering, narrow, its tip slightly
rounded. Eye notably large (as in Hexanchus), oval, its anterior edge about opposite front
of mouth. Spiracle minute, about on level with upper edge of eye, its distance behind eye
about equal to horizontal diameter of latter. Gill openings extending down onto throat, the
1st about I V2 times as long as horizontal diameter of eye, the 2nd to 7th decreasing succes-
sively in length, the 7th only about Y2 as long as ist. Nostril about equidistant between
mouth and tip of snout, its anterior margin expanded as a broadly triangular, corrugated
lobe (Fig. 10 B). Mouth narrowly rounded in front, notably long, the length about equal
to breadth, with very extensive gape, lateral in position for most of its length, the margin
of upper lip extending rearward past corner of mouth for a distance equal to V2 to %
horizontal diameter of eye. An oblique labial furrow at angle of mouth, originating on
upper jaw and extending downward and forward for a short distance onto lower jaw.

Teeth ^zE^i ^^ grown specimens, ^'""j^" in young of 257 mm., unlike in the 2 jaws;
upper teeth fang-like, strongly oblique, the first 3 or 4 more or less sinuous in outline
with base as well as cusp smooth-edged, but subsequent upper teeth with i or 2 small
subsidiary cusps at base on inner side and i on the outer, the outermost upper tooth low,
without definite cusp; lower jaw with i symmetrical tooth at symphysis with large me-
dian cusp, and i or 2 smaller on either side, the lateral lower teeth very broad and low,
each with a series of 6 to 8 somewhat oblique triangular cusps in male, and 7 to 10 in
female, the 2nd or 3rd of which is much the largest, their edges perfectly smooth; 2 to 3
series of teeth functional in front of upper jaw and i along sides; i series functional in
lower jaw.

Origin of dorsal a little posterior to cloaca, its anterior margin straight or slightly con-
vex, its apex broadly rounded, its rear margin concave, its free rear corner prolonged a
distance equal to about V^ the horizontal diameter of eye, its vertical height about Vs ^s
great as length of pectoral. Interspace between dorsal and caudal about as long as between
axil of pectoral and origin of pelvics. Axis of caudal hardly raised, its upper margin mod-
erately convex, lower margin with well marked subterminal notch, rather strongly concave

Fishes of the Western North Atlantic 91

anteriorly, the lower anterior lobe about 30% as long as upper. Anal with nearly straight
margins and subacute corners, about as long as dorsal at base but only about % as high, its
origin about under rear end of base of dorsal. Pelvics a little higher than anal and almost
I Y2 times as long at base, prolonged rearward in male, and partially enclosing the claspers,
the inner margins entirely separate posterior to the cloaca in both sexes. Pectoral relatively
small, % to % as broad as long, with very broad base, the outer margin weakly convex,
distal margin moderately concave, apex narrowly rounded and inner corner more broadly
so.

Color. Fresh specimens from Cuba are described* as uniformly gray, sometimes
shaded with brownish, somewhat paler below than above; pectorals bordered with white;
pelvics and anal pale; dorsal black at apex, with two white spots, one midway of its an-
terior margin, the other near its rear base; caudal edged below with white, its apex with a
black spot, edged with white. After preservation, these same specimens (see Study Mate-
rial, p. 88) are dark mouse-gray above, grayish white below, with apex of dorsal and tip
of caudal dusky, the latter pale-edged.

Developmental Stages. Gravid females have been taken off Cuba with as many as 1 8
embryos, ranging in size up to 150 mm.^ Nine embryos, about lOO mm. long, taken from
the female listed above (p. 88), are of approximately adult form, the chief differences
being their much larger eyes, which is a common embryonic feature, relatively longer
caudals, less deeply emarginate dorsal and pectoral fins, and relatively shorter body cavi-
ties. The large oval yolk sac shows no signs of any attachment to the wall of the oviduct
of the mother.* Up to 20 embryos have been found in a female, in Cuban waters.'

Size. This Shark may be born at a length no greater than about 10 inches; males may
mature at 2 to 2Y2 feet, and females at about 3 feet, or perhaps while even smaller. The
few specimens for which sizes have previously been recorded in scientific literature have
ranged from about one foot, two inches (350 mm.)' to a maximum of seven feet (about
2.14 m.).^ Although the species has been credited repeatedly with reaching more than
three meters, or 10 feet, we find no definite proof of so large a size for it.

Habits. Very little is known of its mode of life. It seems to be a bottom dweller
chiefly, of coastal waters. Its depth range is wide, however, for on the one hand it is recorded
from 380 to 460 meters depth off Portugal and from deep water off Cuba, while on the
other hand it has been reported as common in the very shallow water of roadsteads and
lagoon-like situations along tropical West Africa." In Spanish waters it is classed as very
voracious, destroying great numbers of food fish, especially hake (Merluccius). No pre-
cise information is available as to its stomach contents. Nothing is known of its breeding
habits, other than as indicated above.

4. Howell-Rivero (Torreya, 9, 1941 : 8, and personal communication).

5. Personal communication from Luis Howell-Rivero.

6. See Lo Bianco (Mitt. zool. Sta. Neapel., ig, 1909: 667) for an account of the egg capsules; Ranzi (Pubbl. Sta.
zool. Napoli, 1$, 1934: 378, 417) for the structure of the uterine wall in the gravid female.

7. Personal communication from Luis Howell-Rivero. 8. Tortonese, Atti Soc. ital. Sci. nat., 77, 1938: 286.

9. Gunther, Cat. Fish. Brit. Mus., S, 1870: 398. 10. Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 46.

g2 Memoir Sears Foundation for Marine Research

Relation to Man. It is not sufficiently plentiful anywhere to be of commercial im-
portance.

Range. Atlantic, west and east, including Mediterranean; Cape of Good Hope;
Japan in the North Pacific; it is represented in Australian waters by a relative (dakini)
so close to perlo that it may finally prove identical (see discussion, p. 87). In the
eastern Atlantic its chief center of population is apparently the Mediterranean, where
it is widespread, although nowhere numerous. Its range extends thence southward to Sene-
gambia, where it is reported from many localities. It is also caught occasionally and in
small numbers to the northward along the Atlantic coasts of the Iberian Peninsula. The
most northerly records are on the Portuguese coast and in the Gulf of Gascony (off Bay-
onne). It is also known from Madeira.

Occurrence in the Western Atlantic. The only published record of its presence in the
western Atlantic is of the two specimens from Matanzas, Cuba, described above;" but
Howell-Rivero writes us that specimens are now being taken occasionally in Matanzas
Bay, including gravid females with embryos in all stages of development, suggesting that
it is now experiencing an upswing in abundance in Cuban waters.

Synonyms and References:

Le Perlon, Broussonet, Mem. Math. Phys. Acad. Sci. Paris, 1780; 668 (descr., Medit.).

Squalus ferlo Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: 10 (descr., type loc, Medit.).

Squalus cinereus Gmelin, in Linnaeus, Syst. Nat., 1789: 1497 (descr., Medit.); Bloch and Schneider, Syst.
Ichthyol., 1801: 133 (descr., Medit.); Bosc, Nouv. Diet. Hist. Nat., 21, 1803: 185 (diagn.) ; Latreille,
Nouv. Diet. Hist. Nat., 24, 1804: 72 (in table of contents) ; Risso, Ichthyol. Nice, 1810: 24 (near Nice,
size); Desvaux, Essai Ichthyol. France, 185 1 : 24 (France, not seen).

Le squale perlon, Lacepede, Hist. Nat. Poiss., 4° ed., i, 1798: 220 (general) ; Sonnini, Hist. Nat. Poiss., 4,
1801-1802: 14 (descr.).

Heftranchias cinereus Rafinesque, Carratt. Gen. Spec. Sicil., 1 810: 13; Indice Ittiol. Sicil., 1 8 10: 45 (Medit.) ;
Swainson, Nat. Hist. Fish. Amphib. Rept., 1839: 314 (general); Bonaparte, Mem. Soc. neuchatel. Sci.
Nat., 2 (8), 1839: 9 (in synopsis); Icon. Faun. Ital., j, 1841: I, index. No. 137 (name); Cat. Pesc.
Europ., 1846: 17 (Medit.) ; Costa, Fauna Napol. Pesci, j, 1854-1857: No. 4, 1 1 (Medit.) ; Gill, Ann.
N. Y. Lye, 7, 1861: 404 (class.); Costa, La Pesca, Napoli, 1871: 87 (Medit., not seen); Rey, Fauna
Iberica, Feces, /, 1928: 289 (descr., meas., Spain); de Buen, Faune Medit., Cons. Internat. Explor.
Medit., 1930: plate not numbered (Medit., Atlant.) ; Ranzi, Pubbl. Sta. zool. Napoli, 75, 1934: 378, 417
(struct, of uterine wall).

Monoperhinu! cinereus Blainville, Bull. Sci. Soc. philom. Paris, 1816: 121 (name).

Carcliarias cinereus Cloquet, Diet. Sci. Nat., 7, 1 81 7: 69 (general).

Squalus {Monofterhinus) cinereus Blainville, in Vieillot, Faune Franc, Poiss., 1825: 80 (descr.).

NoiiJafius cinereus Cuvier, Regne Anim., 2nd ed., 2, 1829: 390 (descr.); Lowe, Trans, zool. Soc. Lond., 2
(3), 1837: 194 (Madeira); Bonaparte, Icon. Faun. ItaL, j, 1835: plate not numbered. Index, No. 127
(descr., Medit.) ; Cuvier, Regne Anim., III. Poiss., 1843: 364 (descr.) ; Van der Hoeven, Handb. Dier-
kunde, 2nd ed., 2, 1855: 261 (general); Nardo, Atti 1st. veneto, (3) 5, 1859-1860: 787 (Medit.);
Fitzinger, Bild. Atlas Naturg. Fische, 1864: fig. 176 (good ill.); Giinther, Cat. Fish. Brit. Mus., 8,
1870: 398 (Medit., Atlant., Madeira); Gervais and Boulart, Poiss., 3, 1877: 195 (Medit. and
Atlant. coasts of France); Reguis, Ess. Hist. Nat. Provence, i, Fasc. I, 1877: 53 (Medit.); Heldreich,
Faune de Grece, 1879:91 (not seen) ; Giglioli, Elenc. Pesc. ItaL, 1880:52 (not seen) ; Perugia, Elenc.
Pesc. Adriat., i88i: 55 (Adriatic) ; Rochebrune, Act. Soc. linn. Bordeaux, (4) d, 1882: 46; Faune Sene-
gambie, /, Poiss., 1883-1885: 24 (Senegambia).

I I. Howell-Rivero, Torreya, 9, 1941 : 7.

Fishes of the Western North Atlantic 93

Sgualus {Notidattus) cinereus Voigt, in Cuvier, Tierreich, 2, 1832: 509 (descr.).

Heftanchus cinereus Miiller and Henle, Plagiost., 1 841: 81, pi. 35, fig. 3 (teeth, descr., Medit.) ; Dumeril,
Hist. Nat. Poiss., /, I 865: 437, pi. 4, fig. 1-4 (teeth, descr., Medit., C. of Good Hope f ? ] ) ; Bocage and
Brito Capello, Poiss. Plagiost., 1 866: 15 (no Portuguese record) ; Miklucho-Maclay, Beitr. Vergl. Neurol.
Wirbelt., /, 1870: 12, pi. 2, fig. 1-6 (brain); Ninni, Ann. Soc. Nat. Modena, 5, 1870: 66 (Medit.);
Canestrini, in Cornalia, etal.. Fauna d'ltal., Pesc, 5, 1871-1872: 43 (Medit.) ; Gegenbaur, Unters. Vergl.
Anat. Wirbelt., j, 1872: pi. 21, fig. 5, 6 (skclet., Medit.) ; Hertwig, Jena Z. Naturw., 8, 1 874: 349, pi.
12, fig. 7, 10, 13 (develop, of denticles, teeth); Doderlcin, Prosp. Metod. Pesci Sicil., 1 878-1 879: 30
(Medit.) ; Stossich, Boll. Soc. adriat. Sci. nat., 5, 1880: 69 (Adrlat.) ; Moreau, Hist. Nat. Poiss. France,
/, 1881: 339 (descr., Medit. and Atlant. coasts, France); Doderlein, Man. Ittiol. Medit., 2, 1881: 78
(Medit.) ;^= Graeffe, Arb. zool. Inst. Univ. Wien, 7, 1 886: 447 (Medit.) ; Carus, Prod. Fauna Medit., 2,
1889-1893: 500 (Medit.); Vieira, Ann. Sci. nat. Porto, 4, 1897: 67 (Portugal); Sicher, Atti Accad.
gioenia, (4) 11 (5), 1898: 16 (Medit.) ; Steinhard, Arch. Naturgesch., i (69), 1903:6, pi. I, fig. I-3
(oral and pharyngeal denticles) ; de Braganza, Result. Invest. Sci. "Amelia," Ichthyol., 2, 1904: 30, 102
(Portugal); Borri, Mem. Soc. tosc. Sci. nat., 44, 1934: 94 (Medit.); Belloc, Rev. des. Trav. Peches
Marit., 7, Fasc. 2, 1934: 152 (ill., Morocco to Senegal) ; Daniel, Elasmobranch Fishes, Univ. Calif. Press,
1934: 4, 24, 25, 51, 91 (scales, fin cartilages, musculature, thymus gland) ; Noronha and Sarmento, Peixes
Madeira, 1934: lOO (Madeira, not seen); Nobre, Fauna Marinha Port., Vert., r, 1935: 414 (Portugal,
size) ; Holmgren, Acta Zool. Intern. Tidskr. Zool., 22, 1941: 3 (skull).

Heftranchus cinereus Gi3.y, List Fish. Brit. Mus., Chondropt., 1851: 68 (Medit., ocean).

Heftrancus angio Costa, Fauna Napoli Pesci, 5, 1854-1857: No. 4, 5, pi. 13, 14, fig. 3 (descr., Medit.).

Heftanchus (H eptranchias) cinereus Brito Capello, J. Sci. math. phys. nat. Lisboa, 2, 1870: 141 (Portugal).

Notidanus {Heftanchus) cinereus, var. fristiurus (var. aetatis) Bellotti, Atti Soc. ital. Sci. nat., 20, 1877: 60
(Medit.).

Notidanus {Heftanchus) cinereus Lo Bianco, Mitt. zool. Sta. Neapel, 75, 1899: 542 (embryos) ; Imms, Proc.
zool. Soc. Lond., /, 1905: 44 (denticles).

H eftranchias deani Jordan and Starks, Proc. Calif. Acad. Sci., (3) 2, 1901 : 384 (type descr., Japan) ; Jordan
and Snyder, Annot. zool. jap., 5, 1901 : 128 (Japan) ; Jordan and Fowler, Proc. U.S. nat. Mus., 26, 1903:
595 (descr., Japan); Pietschmann, S. B. Akad. Wiss. Wien, Math.-Naturn. KL, 7/7 (i), 1908: 708
(descr., Japan).

Hexanchus cinereus Seabra, Bull. Soc. portug. Sci. nat., 5, 191 1 : 195 (Portugal).

Heftranchias ferlo Garman, Mem. Harv. Mus. comp. Zool., jd, 1913: 21 (descr.) ; Hussakoff, Copeia, 67,
1919: 9 (descr., Japan); Fovifler, Bull. Amer. Mus. nat. Hist., yo (1), 1936: 27 (descr., W. Africa);
Norman and Fraser, Giant Fishes, 1937: 5, fig. 8 (general) ; White, Bull. Amer. Mus. nat. Hist., ^4 (2),
1937: 40, pi. le, 4m, 17c, 23b, 29e, f (anat.); Tortonese, Atti Soc. ital. Sci. nat., -jy, 1938: 286
(Medit.) ; Fowler, Bull. U.S. nat. Mus., loo (75), 1941: 9 (Medit. and Pacif. refs.) ; Norris, Plagiost.
Hypophysis, 1941: 23, 37, pi. I, fig. 4; pi. 20, fig. 79 (brain); Howell-Rivero, Torreya, 9, 1941: 7,
pi. 3, 4 (descr., Cuba) ; Fowler, Revist. Chilen. Hist. Nat., anos 46-47, 1944: 1 13, fig. 5 (Chile) ; Bige-
low and Schroeder, Comm. Shark Fish., Anglo Amer. Carib. Comm., Wash., 1945: 92, fig. 30 (descr., ill.,
range).

Doubtful reference:

Monofterhinus ciliaris Blainville, Bull. Sci. Soc. philom. Paris, 1 8 16: 121 (name only).

Suborder CHLAMYDOSELACHOIDEA

Characters. Anal fin present; only I dorsal fin, without spine; 6 gill openings, all in
front of origins of pectorals; margins of ist gill openings continuous across throat; snout

12. See Doderlein (1881) and Carus (1889-1S93) for additional locality records for the Mediterranean in publi-
cations not accessible to us.

94- Memoir Sears Foundation for Marine Research

not beak-like, without lateral teeth or cirri; teeth alike in both jaws, those in front of
mouth essentially similar to those toward corners; trunk subcylindrical (shark-like) ; eyes
lateral; anterior margins of pectorals not expanded forward beyond ist gill opening;
mouth terminal, without distinct snout; nostril entirely separate from mouth, its anterior
margin without barbel; eye without nictitating membrane; spiracles present; vertebral
column only incompletely segmented, the notochord being somewhat constricted segmen-
tally for a short distance back from head, but of uniform diameter thence rearward; a few
of anterior vertebrae with primary calcifications (cyclospondylic), but the others not cal-
cified; upper jaw (palatoquadrate cartilage) with transverse process attached to orbital
region of cranium by a ligament (only attachment to cranium) ; also with ligamentary
attachment to the hyomandibular arch, which is well developed and provides the chief
suspension for both jaws; propterygial cartilage of pectoral fin bears no radial elements;
heart valves in 6 or 7 rows ; clasper of male not enclosed by margin of pelvic fin, its axial
cartilage attached to basipterygial cartilage of fin by i small element only, its tip with 3
movable accessory cartilages. Development ovoviviparous.^

Remarks. The majority of recent authors have placed Chlamydoselachus (sole
known representative of the group) among the notidanoids because of its large number of
gill openings and the incomplete segmentation of its vertebral column. We believe a sepa-
rate suborder is demanded for it," because it differs so widely from Hexanchus and
Heptranchias (representing the notidanoids) in the much less intimate attachment of its
upper jaw to the cranium, as well as in the facts that its much larger hyomandibular arch
affords the chief suspension for the jaws and that its notochord is of nearly uniform diame-
ter throughout most of its length.

Families, Genera, Species. Only one modern species is known, Chlamydoselachus
anguineus Garman; but teeth, apparently of this genus, have been described from the
Pliocene of Tuscany.

Range. Japan, also eastern Atlantic off southern France, off the Iberian Peninsula,
and near Madeira in moderately deep water; reported from New South Wales, but on
doubtful evidence.'

Fossil Teeth. From Miocene, West Indies; Pliocene, Europe.

1. See Garman (Bull. Mus. comp. Zool. Harv., :2 [i], 1885), Goodey (Proc. zool. Soc. Lond., 1910: 540), Allis
(Acta zool., 4, 1923: 122) and Smith (Dean Memor. Vol., Amer. Mus. nat. Hist., Art. 6, 1937) for detailed
accounts and illustrations of the skeleton and other anatomical features; Garman (Mem. Harv. Mus. comp.
Zool., 36, 1913; pi. 59, fig. 4, 5, pi. 61, fig. 7, 8) and especially Gudger (Dean Memor. Vol., Am. Mus. nat.
Hist., Art. 7, 1940) for excellent illustrations of the egg capsule and of embryos in different stages of develop-
ment.

2. Garman (Science, 5, 1884: 1 17) proposed for Chlamydoselachus a new order, Selachophichthyoidi, a name based
on the supposition that it "stands nearer the true fishes than do the sharks proper." Shortly afterward, however.
Gill (Science, 3, 1884: 346) united it with the fossil genus Didymodus (a pleuracanth) as the suborder Pterno-
donta, while Garman (Bull. Mus. comp. Zool. Harv., ii, 1885: 30) united it, as Cladodonti, with the fossil
Cladodui and its allies, of which he, by then, had come to consider it "the living representative." More recent
studies of the fossil genera in question, however, make it so unlikely that Chlamydoselachus can be properly
grouped with any pleuracanth or cladodont that we prefer to use for the suborder a name based on that of the
modern genus.

3. See Whitley (Fish. Aust., r, i94i: 70).

Fishes of the Western North Atlantic 95

Suborder HETERODONTOIDEA

Characters. Anal fin present 5 2 dorsal fins with well developed spines; only 5 gill
openings, the last 3 or 4 over base of pectoral j snout not beak-like, without lateral teeth
or cirri; teeth similar in both jaws, those toward center of mouth smaller, with 3 to 5
cusps,* but those along outer parts of jaws much larger, rounded (molar), without cusps,
several rows functional; midtrunk subcylindrical (shark-like), but head with snout
strongly flattened both above and below; tail sector flattened below; anterior margins of
pectorals not expanded forward beyond ist gill opening; nostrils connected with mouth
by a deep groove; eye without nictitating fold or membrane; spiracles present; inner mar-
gins of pelvics entirely separate, posterior to cloaca; vertebral column completely seg-
mented throughout its length, its axial canal much contracted in the region of the centra,
which are fully differentiated, and notochord greatly constricted segmentally in centra;
vertebral centra with internal calcareous lamellae radiating from a central ring; skull
without antorbital processes or separate antorbital bars; upper jaw (palatoquadrate carti-
lage) attached by a short ligament to hyomandibular arch as well as closely and much more
extensively to sides of preorbital region of cranium;" rostral cartilage lacking; neural
spines not attached to dorsals; propterygial cartilage of pectoral bears i radial element;
heart valves in only 2 rows; claspers of males projecting freely from pelvics, their axial
cartilages with 3 movable accessory cartilages at tip and attached to basipterygium of the
fin by 2 small connecting elements. Development oviparous; egg cases horny with spiral
flanges, but without tendrils.

Families and Genera. Only one modern family (Heterodontidae) and genus {Met-
er odontus Blainville, 1 8 1 6) with the characters of the suborder.*

Range. East Africa, East Indies, New Zealand, Australia, China, Japan and eastern
Pacific north as well as south; not known in Atlantic or Mediterranean.

Fossil Teeth. Upper Jurassic to Pliocene in Europe; Upper Cretaceous to Eocene in
Africa; Miocene in South America, New Zealand, Australia.

Suborder GALE OWE A

Characters. Anal fin present; 2 (rarely i) dorsal fins, without spines; only 5 gill
openings with rudimentary 6th arch; snout not beak-like, without lateral teeth or cirri;
teeth of essentially the same type in front of mouth as near corners; trunk subcylindrical,
not strongly depressed; eyes lateral; anterior edges of pectorals not extending forward
past 1st gill openings; nostril either connected with mouth or separate from it; nictitating
membrane and spiracles present or absent; inner margins of pelvics either separate posterior
to cloaca, or more or less united; vertebral column completely segmented throughout its
length, its axial canal much contracted or obsolete in regions of centra, the latter being fully
differentiated; notochord greatly constricted segmentally in centra, or even completely
obliterated there, but dilated in spaces between concave faces of adjoining vertebral centra;

4. Cusps may be entirely worn off in large specimens. 5. Firmly articulated there in fossil forms.

6. See Fowler (Bull. U.S. nat. Mus., 100 [x^], 1941 : 15) for list of generic synonyms.

96 Memoir Sears Foundation for Marine Research

vertebral centra with calcareous lamellae radiating from a central ring, or with latter alone
calcified (genera Galeus, Pseudotriakis) ; neural spines not attached to dorsals; skull with
antorbital processes more or less developed, but no separate antorbital bar; rostral carti-
lages 3 (united or separate at tip), i or none; upper jaw (palatoquadrate cartilage) not
articulated with cranium, but connected with ethmoid region by a longer or shorter liga-
ment;' its connection with hyomandibular arch also ligamentary only, at least in most
cases/ Propterygial cartilage of pectoral much smaller than mesopterygium, with I to
several radial elements; heart valves in 2 or 3 rows; claspers of male projecting freely
from pelvics; axial cartilages either single or double, usually with a group of movable
accessory cartilages at the tip when adult, and attached to basipterygium of fin by i small
connecting element only." Development oviparous, ovoviviparous, or viviparous.

Key to Families
la. Only i dorsal fin. Scyliorhinidae (part), p. 195.

lb. 2 dorsal fins.

2a. At least V2 of base of ist dorsal posterior to origin of pelvics.

3a. Caudal lunate, large; gill arches connected one with the next by masses of
spongy tissue, forming sieve-like structures. Rhincodontidae, p. 187.

3b. Caudal not lunate, not very large; gill arches not connected one with the next
by masses of spongy tissue.

4a. Nostril connected with mouth by a deep groove, its anterior margin with
a well developed barbel. Orectolobidae, p. 178.

4b. Nostril not connected with mouth by a deep groove, or, if so connected,
its anterior margin without a well developed barbel.

Scyliorhinidae, p. 195.
2b. Base of ist dorsal terminates over, or (usually) well anterior to, origin of pelvics.
5a. Head greatly expanded laterally. Sphyrnidae, p. 407.

5b. Head of normal shape, not expanded laterally.
6a. Caudal fin lunate, its axis steeply raised.

7a. Teeth large, few in number; gill arches without gill rakers.

Isuridae, p. 109.
7b. Teeth minute, very numerous; gill arches with well developed
rakers. Cetorhinidae, p. 146.

6b. Caudal fin not lunate, its axis raised only moderately at most.

8a. 1st dorsal fin longer at base than caudal. Pseudotriakidae, p. 228.
8b. ist dorsal fin much shorter at base than caudal.

9a. Caudal fin occupies nearly J/2 total length, or even more.

Alopiidae, p. 160.

7. This allows the jaws to be more or less protrusible in many cases.

8. Parker's (Trans, zool. Soc. Lond., lo, 1879: pi. 38, fig. 2) illustration of the skull of Scylliutn canicula, equals
Scyliorhinus caniculus (Linnaeus), 1758, which shows these ligamentary connections well, has been copied in many
subsequent textbooks of zoology.

9. For illustrations of the cartilages of the clasper in various galeoids, see especially Huber (Z. Wiss. Zool., 70,
1901 : pi. 27) i White (Bull. Amer. Mus. nat. Hist, 74, 1937: pi. 46-50).

Fishes of the Western North Atlantic

97

9b. Caudal fin occupies considerably less than y> total length.

lOa. 5th gill opening well in front of origin of pectoral; eye
without nictitating fold or membrane.
1 1 a. Jaws widely protrusible forward; snout greatly
elongate. Scapanorhynchidae, p. 109.

I lb. Jaws not widely protrusible ; snout not greatly elon-
gate. Carchariidae, p. 98.

Figure 12. A Eye of Sfhyrna diflana, about 1375
mm. long, to show nictitating membrane (U. S. Nat.
Mus., No. 108452), about 2 x natural size. B Eye of
Mustelus canis, about three feet long, to show sub-
ocular fold (Harv. Mus. Comp. Zool., No. 35245).

lOb. 5th gill opening over or behind origin of pectoral; eye

with a more or less strongly developed nictitating fold or

membrane.

1 2a. Upper edge of nictitating fold continuous with edge
of eyelid, or even arising outside latter posteriorly,
although enclosing it anteriorly; teeth low, rounded
or with 3 or more cusps, usually in mosaic arrange-
ment, several series functional simultaneously in
sides of jaws as well as in front.

Triakidae, p. 233.^°

12b. Upper edge of nictitating membrane arises far
within edge of eyelid posteriorly, as well as ante-
riorly; teeth blade-like with i cusp only, not in
mosaic arrangement, usually not more than i or 2
series functional in sides of jaws simultaneously.

Carcharhinidae, p. 262.'°

10. It may not be possible to draw a sharp line between Triakidae and Carcharhinidae with respect to the nictitating
membrane or the teeth. However, the definition given above will serve to place any genus yet known from the
Atlantic in the one family or in the other.

g8 Memoir Sears Foundation for Marine Research

Family CARCHARIIDAE
Sand Sharks

Characters. Two dorsal fins, the ist much shorter than caudal, the rear end of
its base over or anterior to origin of pelvics; caudal not more than Vs of total length, not
lunate, its lower anterior corner expanded as a distinct lobe, its axis raised but little j caudal
peduncle not greatly depressed or expanded laterally 5 a precaudal pit above but none
below; sides of trunk, anterior to anal, without longitudinal dermal ridges; inner margins
of pelvics more or less united posterior to cloaca in male, less so in female; jaws not
greatly protrusible; snout not greatly elongate; 5th gill opening anterior to origin of pec-
toral; gill arches without rakers, not interconnected by a sieve of modified denticles; nos-
trils entirely separate from mouth, their anterior margins without barbels; spiracles pres-
ent; lower eyelid without nictitating fold or membrane; teeth large, awl-shaped, with or
without lateral denticles and not very numerous (see counts, p. 102); skull of normal
form {i.e., not greatly expanded laterally); rostral cartilages united in one; mesoptery-
gium of pectoral with about ^2 as many radials as metapterygium, and nearly as large;
meso- and metapterygia not separated by foramen. Development ovoviviparous.

Genera. Only one modern genus, Carcharias, so far known.

Genus Carcharias Rafinesque, 18 10
Sand Sharks
Carcharias Rafinesque, Caratt. Gen. Nuov. Sicil., 1810: lO; type species, C. taurus Rafinesque. Sicily.''

Generic Synonyms:

Squalus (in part) Risso, Ichthyol. Nice, 1810: 38; for S. ferox; also subs, authors; not Squalus Linnaeus, 1758.

Galeorhinus (in part) Blainville, Bull. Soc. philom. Paris, 1 8 16: 121; for G. ferox.

Oiontasfis Agassiz, Poiss. Foss., 1838: 5, 87; type species, Carcharias ferox Risso, 1826, equals Squalus ferox

Risso, 1 810.
Triglockis Miiller and Henle, Arch. Naturg., 1837: 396; type species, Carcharias ferox Risso, 1826, equals

Squalus ferox Risso, i 8 1 0.
Eugotnfhodus Gill, Proc. Acad. nat. Sci. Phllad., Addend., 1 861: 60; type species, Carcharias griseus Storer,

1846, equals Carcharias taurus Rafinesque, i8lO; monotypic.
SynoJontasfns White, Vert. Faun. Engl. Eocene, 193 1 : 51 ; type species, Carcharias taurus Rafinesque, 1 8 10.
Paradontasfis White, Vert. Faun. Engl. Eocene, 1 931: 63; type species, Odontasfis flatensis Lahille, 1928.'^

Generic Characters. Caudal peduncle with a well marked pit above (none below)
and without lateral keels; dermal denticles with 3 broad longitudinal ridges; snout coni-
cal; jaws with or without labial furrows; anterior teeth in both jaws two-rooted, the pos-
terior teeth less obviously so; spiracle small; 2nd dorsal about as large as ist; caudal with
small but definitely outlined lower anterior lobe and well marked subterminal notch.
Characters otherwise those of the family.

11. Opinion 47 of the International Commission on Zoological Nomenclature (Smithson. Publ., 2060, 1912: 108;
Copeia, 29, 1916: 28) confirms Carc/mrias taurus Rafinesque, 1810, as the type of Carcharias Rafinesque, iSio; it
thus replaces Odontaspis Agassiz.

12. The fossil genus Oxytes Giebel, Fauna Vorwelt, Fische, 1847: 364, type species, O. obliqua Giebel (monotypic),
is referred to the synonymy of Carcharias by Fowler (Bull. U.S. nat. Mus., 100 [15], 1941 : 119).

Fishes of the Western North Atlantic 99

Range. Both sides of warm temperate and tropical North Atlantic, including the
Mediterranean; eastern South America south to northern Argentina; South Africa; India;
Australia; China; Japan.

Fossil Teeth. Lower Cretaceous to Pliocene, Europe; Upper Cretaceous to Miocene,
South America; Upper Cretaceous to Pliocene, North America, New Zealand; Upper Cre-
taceous, Asia; Paleocene to Pliocene, Africa; Miocene, Australia, West Indies.

Species. The members of this genus fall into two easily separable divisions, the one
represented by a single well defined species (ferox Risso), the other by a group of named
forms, so clearly allied one to another that it is still an open question how many of them
deserve separate specific names. While awaiting comparison of specimens from different
ocean areas, the accompanying key recognizes differences which may later prove merely
varietal.

Provisional Key to Species

I a. 1st upper tooth notably smaller than 2nd, each tooth usually with 2 denticles on each
side; 3rd upper tooth followed by 4 very much smaller teeth. ferox Risso, 1 8 lO.

Eastern Atlantic,
Mediterranean.

lb. 1st upper tooth only slightly smaller than 2nd, if so at all; each tooth usually with i
denticle only (rarely 2) on each side, or with none; 3rd upper tooth followed by 2 or
3 much smaller teeth at most.

2a. 3rd upper tooth followed by 2 or 3 much smaller teeth, no wide gap between these
and the next large (5th or 6th) tooth. flatensis Lahille, 1928.

Argentina.

arenarius Ogilby, 1 9 1 1 .

Australia.'*

2b. 3rd upper tooth followed by i much smaller tooth only, the latter separated from
the succeeding large tooth by a broad gap.

3a. Snout broadly rounded; inner margin of pectoral only ^/^ as long as outer;
no labial furrow at angle of mouth. tricuspidatus Day, 1888.

India, China."
3b. Snout pointed; inner margin of pectoral more than Ys as long as outer; well
marked labial furrows at corners of mouth.

4a. Lateral denticles lacking on most teeth, minute on others; length of
longest tooth less than Yi diameter of eye. owstoni Garman, 19 13.

Japan.

4b. Most or all of teeth with a well developed lateral denticle on each side;
length of longest tooth at least % diameter of eye.

/i2Mr«j Rafinesque, 1810, p. lOO.'^

13. Published descriptions are not suinciently detailed for critical comparison of flatensii with arenarius.

14. Fang and Wang, Contr. Biol. Lab. Sci. Soc. China, 8, 1932: 241.

15. It has been suggested recently that the American form {liUoralis) differs from the European taunts in having
no denticles on its first and fourth upper teeth (Giltay, Mem. Mus. Hist. nat. Belg., Hors Serie, 5, Fasc. 3, 1933 :
7). Our own examination of specimens of various sizes from southern New England and the vicinity of New
York shows that while the teeth in question are smooth in some small specimens (about three feet Ions), they have

lOO

Memoir Sears Foundation for Marine Research

Carcharias taurus Rafinesque, 1810

Sand Shark, Sand Tiger

Figures 13, 14

Study Material. Five Massachusetts specimens, male and female, 943 to 1,081 mm.
long (Harv. Mus. Comp. Zool.) 5 jaws of a large specimen from New Jersey, and also of a
female, 2,800 mm., from Engiewood, Florida} also many medium-sized specimens, fresh
caught at Woods Hole, Massachusetts.

\^'^S^^:^^S^:iS3iXPi^iax'^

Figure 13. Carcharias taurus, young male, loio mm. long, from Cape Cod, Massachusetts (Harv. Mus.
Comp. Zool., No. 351). A Anterior part of head of same from below. B Dermal denticles, general view, about
25 x; lateral and apical views, about 50 x. C Upper and lower teeth of larger specimen from New Jersey
(Harv. Mus. Comp. Zool., No. 351), about I x.

Distinctive Characters. The five gill openings in front of the pectorals, the second
dorsal about as large as the first, the position of the first dorsal entirely in front jf the
pelvics, the entire separation of the nostril from the mouth, and the highly characteristic
teeth (Fig. 13 C) are diagnostic among sharks of our province.

denticles on one or on both sides in others and denticles on both sides in the two larg-e specimens that we have exam-
ined. Neither do the supposed differences in the relative position of the rear end of the base of the first dorsal
fin, or ifl the origin of the pelvics, invoked by earlier authors as a specific character, prove any more significant,
for these vary considerably among American specimens (see p. 101).

Fishes of the Western North Atlantic

lOI

Descripion. Proportional dimensions in per cent of total length. Female, 982 mm.,
from Mass. (Harv. Mus. Comp. Zool., No. 436). Male, 1,081 mm., from Mass. (Harv.
Mus. Comp. Zool., No. 402).

A

Figure 14. Carcharias taurus, showing teeth from
jaws pictured in Fig. 1 3, about 2 x. A, B Third upper
tooth. C Seventh upper tooth. D Twelfth upper
tooth. E Seventeenth and eighteenth upper teeth.
F, G Third lower tooth. H Seventh lower tooth.
/ Tenth lower tooth. J Seventeenth and eighteenth
lower teeth.

Trunk at origin of -pectoral: hrezdih 10.7, 10.2; height 12.2, 10.7.

Snout length in front of: outer nostrils 3.4, 3.3; mouth 3.7, 3.9.

Eye: horizontal diameter 1.5, 1.2.

Mouth: breadth 8.1, 8.0} height 4.7, 5.2.

Labial furrow length: upper (visible part) 0.9, 0.9; lower 2.4, 2.3.

Nostrils: distance between inner ends 3.1, 3.2.

Gill opening lengths: ist S-Sj 5-i j 2nd 5.0, 5.1; 3rd 4.9, 4.6} 4th 4.9, 4.45 5th

3-8, 34-

First dorsal fin: vertical height 7.2, 6.6-, length of base 7.5, 8.4.
Second dorsal fin: vertical height 6.3, 6.1 ; length of base 7.0, 7.2.
Anal fin: vertical height 5.8, 5.95 length of base 7.1, 7.7.
Caudal fin: upper margin 30. i, 29.3 ; lower anterior margin 9.8, lO.o.
Pectoral fin: OMi^r margin 13.5, 14.2; inner margin 5.8, 6.0; distal margin 9.4, 9.5.
Distance from snout to: ist dorsal 40.8, 39.3; 2nd dorsal 56.8, 57.2; upper caudal
70.0, 70.5; pectoral 22.8, 22.8; pelvics 48.0, 49.5; anal 60.2, 61.7.
Interspace between: ist and 2nd dorsals 11.2, 10.8; 2nd dorsal and caudal 7.1,
6.2; anal and caudal 3.8, 2.8.

Distance from origin to origin of: pectoral and pelvics 29.9, 27.5; pelvics and
anal 14.4, 14.1.
Trunk moderately stout, its height abreast pectoral origin about /'g, opposite origin
of dorsal about Vtj of the total length. Caudal peduncle relatively high and laterally com-

I02 Memoir Sears Foundation for Marine Research

pressed. Dermal denticles about 0.4 mm. broad, by 0.45 mm. long in a specimen of about
100 cm. length, loosely spaced, their blades ovoid lanceolate, their anterior margins en-
tire or slightly indented between tips of the 3 ridges; axial ridge very prominent and
sharp-edged anteriorly but usually flat-topped and subdivided posteriorly.

Head moderately flattened above. Snout short, its length in front of mouth about V4
to Vs the length to ist gill slit, narrow ovoid, with rounded tip. Eye round and small, its
diameter only about % as long as distance between nostrils. Spiracle minute, about on a
level with upper margin of eye and behind latter by a distance about equal to length of
snout in front of mouth. Gill openings relatively large, 4th about as long as snout in front
of mouth, others slightly shorter, the 5th shortest. Nostril nearly transverse, its anterior
margin with a small rounded flap near inner end. Distance from inner angle of nostril to
mouth about equal to width of nostril. Mouth crescentic in front, about % to % as long as
broad; angle of mouth with well marked labial furrow on lower jaw and a less prominent
one on upper; upper furrow partially hidden when mouth is closed.

Teeth l\ |° H m specimens examined, ist to 6th or 7th teeth in each jaw either with
or without i or 2 small basal denticles on either side;'° ist upper tooth usually a little
smaller, but sometimes of the same size as 2nd or 3rd, the 4th much smaller than 3rd
or 5th, with a broad interspace between 4th and 5th} ist lower tooth much smaller than
2nd to 6th, the teeth posterior to 6th or 7th successively smaller in each jaw and broader
relative to length, with denticles successively larger relative to median cusp, the outermost
12 or 13 minute, close set, tricuspidate, about as broad as high; 3 or 4 series functional
toward corners of mouth, but only i or 2 series toward center."

Origin of ist dorsal about midway between axil of pectoral and origin of pelvics, its
base terminating a little anterior to latter, its apex subacute, its rear margin slightly concave,
the free corner about Vs as long as its base, its vertical height about V2 as great as length of
pectoral or about V4 as great as length of head. Second dorsal similar to ist and only
slightly smaller, its origin about midway between cloaca and origin of anal, Y^ to V2 of its
base overlapping base of latter. Caudal about 30% of total length, its axis only slightly
raised, the subterminal notch well marked, the posterior outline of terminal sector con-
cave, its lower anterior corner expanded as a definite lobe with rounded apex, its anterior
margin about Vs as long as upper caudal margin; re-entrant corner, included between the
2 lobes, broadly rounded. Anal a little larger than 2nd dorsal in area and a little longer
basally, its rear margin less deeply concave, its free rear tip about Vs as long as its base,
the interspace between anal and caudal only about V2 as long as base of anal. Pelvics origi-
nating a little posterior to rear end of base of ist dorsal, and about as large as latter, the
inner margins entirely separate posterior to cloaca in female, but connected for a short

16. On a large New Jersey specimen every tooth, from the first to the seventh, is flanked by one or two denticles
on each side; in another, from southern Massachusetts, the fourth upper tooth lacks a denticle, while on still
other specimens from the same general locality some of the teeth have a denticle on each side, some have a den-
ticle on one side only, and still others have no denticle on either side.

17. An account of the shedding of the teeth of specimens in an aquarium is given by Breder (Copeia, 1942: 42) ;
see also p. 65.

Fishes of the Western North Atlantic 103

distance in male. Pectoral a little naore than Vj as broad as long, with nearly straight distal
and outer margins, rounded corners, and wide base.

Color. Light gray-brown above, darkest along back, snout, and on upper sides of
pectorals, paling on the sides to grayish white on belly and on lower sides of fins 5 sides
of trunk rearward from pectorals as well as caudal and dorsals variously marked with
roundish to oval spots, varying in color from yellowish brown to ochre yellow. In a speci-
men 100 cm. in total length these spots vary from less than V2 cm. to more than iVs cm.
in diameter, numbering upwards of lOO. Posterior margins of fins edged with black on
some specimens but perhaps not on all.

Size. In the northern sector of their American range, from Delaware Bay to Cape
Cod, Sand Sharks are recorded from 3 feet to about 9 feet, the great majority of those
caught being immature, of perhaps 4 to 6 feet. Large adults (7 to 8 feet or more) are also
reported, not rarely, from widely scattered localities along the New Jersey coast, from the
vicinity of New York, from Clinton, Connecticut (8 feet 10 inches), and especially from
the vicinity of Nantucket, where commercial operations in the early nineteen-twenties are
said to have yielded "a wealth of eight and nine foot Sand sharks.'"' From North
Carolina southward, however, large ones alone have been reported, the recorded lengths
ranging from about 8 to 9 feet in the Beaufort-Cape Lookout region; 6V2. to 9^/2 feet for
Charleston, South Carolina; 9 feet 2 inches to 10 feet 5 inches for southwestern Florida
at Englewood, the last named being the greatest length yet positively recorded for Car-
charias taurus. The recorded weight of about 250 pounds for an 8-foot lO-inch specimen
from Clinton, Connecticut, shows how much lighter a fish this is, length for length, than
the Mackerel Shark, Mako or White Shark. We have no firsthand information to con-
tribute.

It appears, from the state of sexual development of the specimens we have seen, and
from the sizes of the few females so far reported as containing eggs or embryos, that this
Shark does not mature until it attains a length of perhaps seven feet or upward.

Developmental Stages. Females have been reported containing many eggs as well as
embryos.

Habits. In spite of its trim appearance and voracious appetite (see below) this is a
comparatively sluggish shark, living mostly on or close to bottom, being more active and
biting the hook more freely by night than by day. It is a coastwise species, as contrasted
with pelagic, most of those caught being taken in depths of not more than two to five
fathoms; and it is often encountered close in to the tide line in only two to six feet of water,
hence its frequent capture in pound nets. It has not been reported from the fishing banks
off Nantucket or at the mouth of the Gulf of Maine. To the southward, however, it may
not be so strictly confined, witness its presence on the North Carolina Banks.

Knowledge of its breeding habits is confined to the facts that a large female, taken at
Beaufort, North Carolina, in April contained many large eggs; also that specimens a little

18. Young and Mazet, Shark, Shark, 1933: 132.

I04 Memoir Sears Foundation for Marine Research

more than eight feet long, at Cape Lookout, North Carolina, contained many eggs and
embryos more than nine inches long in July; and that females with unripe eggs have been
reported at Woods Hole in the same month in different years. Since no embryos have been
found in large females in Florida, and since immatures three to five feet have been reported
so far only from the section north from Delaware Bay (where these constitute the majority
of the local stock, p. 103) this is probably the chief center for the production of young,
but information is still lacking as to the seasonal occurrence of gravid females there, or of
newborn young.

Proverbially voracious, the Sand Shark feeds chiefly on smaller fishes, for the
capture of which its slender raptorial teeth are admirably adapted. Large specimens have
been taken with as much as 100 pounds of fish in their stomachs, and by eyewitness accounts,
schools of them may surround other fish or even those imprisoned in fishermen's nets. On
the east coast of North America the recorded diet, depending on the geographical local-
ity, includes alewives {Pomolobus), black drum (Pogonias), bluefish {Pomatomus),
bonito (Sarda), butterfish (Poronotus), cunner {T autogolabrus) , eels (Anguilla), flat-
fishes, menhaden (Brevooriia), mullet (Mugil), scup {Stenotomus), sea bass {Ceniro-
fristis), sea robin (Prionotus), small sharks (species?), shark sucker (Echeneis), silver
hake (Merluccius), spadefish {Chaetodipterus), spot (Leiostomus), tautog (Tautoga)
and the weakfishes, spotted (Cynoscion nebulosus) and gray (C. regalis). No doubt a
complete list for any given locality would include practically all the local species that were
not too large. Squid have been found in their stomachs at Woods Hole, likewise crabs and
lobsters, although the latter are perhaps only exceptionally eaten, for they were not found
among the stomach contents of many more which were recently examined at Woods Hole
on different occasions. There is no reason to suppose that this species ever attacks large prey.

Relation to Man. Although plentiful, the Sand Shark is of little commercial impor-
tance at present. A few are included in the catch of the Florida shark-fishery; occasional
specimens are sold at a low price in fish markets. There were local fisheries for it for
leather in Nantucket Sound, in the first quarter of the present century, but these were
short-lived, reportedly because of exhaustion of the stock. However, it is of some interest
to sport anglers, considerable numbers being caught by them yearly, both as objects of
special pursuit or incidentally while surf-casting for other fish. But its resistance when
hooked is so much less vigorous for its size than that of the more active pelagic sharks, such
as the Mako or White Sharks (pp. 128, 139), that few would rate it as in the game class.

There is no record of attack by a Sand Shark on human beings in North American
waters, although bathers often come close to them, our own experience bearing this out.
Its relative (or relatives) in East Indian waters bears a sinister reputation, however.

Range. Mediterranean, tropical West Africa, Canaries and the Cape Verdes in the
eastern Atlantic; South Africa; western Atlantic from the Gulf of Maine to Florida and
southern Brazil; represented in Argentine waters and in the Indo-Pacific by close allies
(see Species, p. 99).

Fishes of the Western North Atlantic 105

Occurrence in the Western Atlantic. Next to the Smooth and Spiny Dogfishes (p.
466), the Sand Shark is probably the most abundant shark in season from Delaware Bay
northward to Cape Cod; in this region it is far more plentiful than it is anywhere in the
eastern Atlantic. Considerable numbers are caught all along the coast of New Jersey both
in the bays and outside; it is a common visitor yearly to the vicinity of New York, along
Long Island and presumably within Long Island Sound." It is common in summer in
Rhode Island waters, and it is fairly so around Block Island. So general is its occurrence
along the southern shores of Massachusetts, including Martha's Vineyard and Nantucket,
that every local fisherman knows it well. As an example of its local numbers we may cite
the fact that a catch of about 1,900 sharks, made by three boats on Horseshoe Shoal in
Nantucket Sound from June to September, 19 18, consisted chiefly of this species.^" Simi-
larly, a catch of 350 sharks, made near Nantucket in the early 1920's, consisted of this
species with few exceptions."' It is also taken in some numbers yearly along the outer
shores of Cape Cod. But this marks the eastern boundary of its center of abundance, for
while it is recorded at various localities around Massachusetts Bay, these are occasional
specimens only. Only as a stray does it wander north of Cape Ann; it was reported once
from Casco Bay and once from St. Andrews, New Brunswick, at the mouth of the Bay
of Fundy.

Our data are not adequate to describe its status from Delaware Bay southward.
It is reported from the Bay itself, both near the mouth at Bowers Beach and even from
the vicinity of Philadelphia at its head; likewise from the coast of Maryland, and
from Chincoteague and Smith Island in Virginia. However, these reports do not suggest
any great numbers. The survey of the fishes of Chesapeake Bay by the United States
Bureau of Fisheries" did not yield even a single record, although it has been reported
there more recently. Nor does it appear with any regularity along North Carolina, al-
though large schools appear at times off Cape Lookout, and it rarely enters the local
sounds." On the other hand, it is described as one of the commonest summer sharks on the
South Carolina coast, near Charleston, with as many as six large specimens recorded from a
single net haul. It is taken on the east coast of Florida at all seasons, as at Salerno, near
Jupiter Inlet, where it appears irregularly in considerable numbers. However, it apparently
reaches the west coast of Florida as a stray only, but two specimens being known from En-
glewood, where the shark stock has been the subject of special investigation.^* It has been
taken off the northern Bahamas." We find no published report of it anywhere else for the
Gulf of Mexico, the Bahamas, Cuba, the Antilles, or for the Caribbean region, although
it is so easily recognizable and usually comes so close inshore that it could hardly have been
overlooked if it occurred with any regularity within these general areas. However, its

19. The only published record of it in the Sound is for Clinton, Connecticut.

20. Identity established by excellent photographs by R. H. Bodman, who reported this catch to us.

21. Young and Mazet, Shark, Shark, 1933: 132.

22. Hildebrand and Schroeder, Bull. U.S. Bur. Fish., ^5, 1928.

23. Only two specimens, both large, reported from Beaufort, N.C.

24. Springer, Proc. Fla. Acad. Sci., 3, 1939: 34- ^S- Wise, Nat. Hist N.Y., 38, 1936: 322 (photo).

io6 Memoir Sears Foundation for Marine Research

presence, at least as a stray, has been proved recently at Bermuda by the capture of typical
specimens, in 1927 and 1942.^' It apparently has a second center of occurrence on the
coast of Brazil, since Sand Sharks, seemingly identical with the northern taurus, are plenti-
ful in October and November near Rio de Janeiro, where many are placed for sale in the
market} they are recorded as far south as the Rio Grande do Sul. But it is not yet possible to
define the boundaries of this southern population, owing to the uncertainty of identity
(whether taurus or plaiensis) of the nominal records from Brazil, Uruguay and Argentina.
Nor does any explanation suggest itself for the apparent discontinuity between the areas
of distribution of the North Atlantic and South Atlantic populations.

On the east coast of Florida C. taurus is taken irregularly at all seasons. From
South Carolina northward, however, it has been reported only during the warm half
of the year. Thus, at Charleston, South Carolina, it is reported for summer only; off
North Carolina it may appear from late April on through the spring and equally early in
the season at the mouth of Delaware Bay, as in 1921, when eight were taken at Cape May
on April 21. However, May 27 appears to be the earliest recorded date for it on Long
Island at Orient, with its season of maximum abundance extending from June into early
October all along the coast from New York to Cape Cod. It withdraws from the neigh-
borhood of New York in autumn, when the temperature of the water falls below about
67° F. (19-20° C), and departs from the coasts of southern New England and New
Jersey by November at the latest.

The winter home of the Sand Sharks that summer along the northeastern United
States is not known. No increase in their numbers in autumn or early winter has been noted
along North Carolina or Florida, coincident with their disappearance from the North.
Like various bony fishes, it is possible that they move offshore, and possibly southward,
to escape winter chilling.

Synonyms and References:

Carcharias taurus Rafinesque, Caratt. Gen. Nuov. Sicil., 1810: 10, pi. 14, fig. I ; Indice Ittiol. Sicil., 1810: 45
(type loc, Sicily) ; Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 25, pi. 6, fig. I-3, pi. 4.I, pi. 51,
fig. 7 (descr.) ; Radcliffe, Trans. Amer. Fish. Soc, 1914: 35; Bull. U.S. Bur. Fish., 34, 1916: 244, pi.
38, fig. I, 2 (N. Carolina, teeth) ; Nichols and Murphy, Brooklyn Mus. Sci. Bull., i (l), 1916: 21, pi. 2
(occur.) ; Fowler, Copeia, 30, 1916: 36; Copeia, 31, igi6: 41 (N. Jersey) ; Copeia, 35, 1916: 69 (Long
Island, N. York) ; Jordan, Copeia, 29, 1916: 281 (name) ; Smith, Amer. Mus. J., 16, 1916: 347 (gen-
eral) ; Thome, Copeia, 35, 1916: 69 (N. York); Fowler, Proc. Boston Soc. nat. Hist., 55, 191 7: no
(Nantucket, Woods Hole, Massachusetts) ; Proc. Acad. nat. Sci. Philad., 6g, 1 91 7: 1 08 (N. Jersey, size) ;
Occ. Pap. Mus. Zool. Univ. Mich., 56, 1918: 15 (Virginia); Copeia, 68, 1919: 13 (N. Jersey); Proc.
biol. Soc. Wash., 32, 1919: 72 (Delaware R.) ; Proc. Acad. nat. Sci. Philad., yi, 1919: 292 (N. Jersey);
Roule, Result. Camp. sci. Monaco, 52, 1919: 1 16 (St. Lucia L, C. Verde); Fowler, Proc. biol. Soc.
Wash., 55, 1920: 143 (N. Jersey) ; Sherwood, Copeia, loO, 1921 : 77 (Connecticut, large size) ; Fowler,
Proc. Acad. nat. Sci. Philad., ^2, 1921: 385 (N. Jersey, small size); Proc. Acad. nat. Sci. Philad., 74,
1922: 2, 5, 7 (N. Jersey, Delaware); Breder, Bull. N. Y. zool. Soc, 25, 1922: 137 (in aquarium);
Huntsman, Contr. Canad. Biol. (1921), j, 1922: 8 (St. Andrews, New Brunswick); Linton, Proc. U.S.
nat. Mus. 64, 1924: 13, 14 (parasites) ; Breder, Copeia, 127, 1924: 27 (Sandy Hook Bay, New York, and
New Jersey) ; Barnard, Ann. S. Afr. Mus., 21 (i), 1925:36 (Medit. and Cape Seas) ; Bigelow and Welsh,
Bull. U.S. Bur. Fish., 40 (l), 1925: 34 (descr., food. Gulf of Maine) ; Fowler, Copeia, 143, 1925: 41,

26. Personal communication from Louis Mowbray, director of the Bermuda Aquarium.

Fishes of the Western North Atlantic 107

42 (N. Jersey, Delaware Bay, size) ; Breder, Copeia, 153, 1926: 122 (near N. York) ; Fowler, Copeia,
156, 1926: 146 (N. Jersey) ; Chabanaud and Monod, Bull. Etud. Hist. Sci. Afr. Occid. Franc. (1926),
1927: 228 (tropical West Africa) ; Barnard, Ann. S. Afr. Mus., 2/ (2), 1927: pi. 2, fig. 2 (S. Africa) ;
Fowler, Copeia, 16;, 1927: 89 (Virginia) ; Rey, Fauna Iberica Feces, /, 1928: 394 (descr., Spain) ; Fow-
ler, Proc. Acad. nat. Sci. Philad., 80, 1929: 607 (N. Jersey, size); Truitt, Bean and Fowler, Bull. Md.
Conserv. Dept., 3, 1929: 30 (Chesapeake Bay) ; Uriarte and Mateu, Notas Inst. esp. Oceanogr., 53, 1931:
10 (Canary Is., size) ; Wilson, Bull. U.S. nat. Mus., 158, 1932: 464, 469 (parasites, Woods Hole) ; Giltay,
Mem. Mus. Roy., Hors Ser., 5, Fasc. 3, 1933: 8 (class.) ; Young and Mazet, Shark, Shark, 1933: 132, 271
(commercial catch, near Nantucket) ; Bigelow and Schroeder, Canad. Atlant. Fauna, biol. Bd. Canad.,
I2e, 1934: 12 (descr., distrib.) ; Budker, Bull. Mus. Hist. nat. Paris, (2) 7, 1935: 184 (Dakar, W.
Africa) ; Fowler, Bull. Am. Mus. nat. Hist., yo (i), 1936: 29 (W. Africa) ; Wise, Nat. Hist. N. Y., 2S,
1936: 322 (photo, Bahamas); Fowler, Proc. Acad. nat. Sci. Philad., 8g, 1937: 303 (N. Jersey); Wise,
Tigers of the Sea, 1937: 178 (general); Tortonese, Atti Soc. ital. Sci. nat., 77, 1938: 288 (Medit.) ;
Fowler, Arch. Zool. Estado Sao Paulo, 4, 1942: 127 (Brazil) ; Monogr. Acad. nat. Sci. Philad., 7, 1945:
262 (Florida) ; Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Carib. Comm., Wash.,
1945: 100, fig. 34 (descr., ill., habits, range).

Squalus americanus Mitchill, Trans. Lit. Phil. Soc. N. Y., 1815: 483 (descr., N. York), not 5. americanus
Gmelin, 1789; DeKay, Zool. N. Y., 4, 1842: 366 (descr., N. York) ; Gill, Proc. Acad. nat. Sci. Philad.,
Addend., 1 861 : 60 (named).

Squalus littoralis Lesueur, J. Acad. nat. Sci. Philad., /, 1817: 224 (descr., N. York) ; Mitchill, Amer. Month.
Mag. Crit. Rev., 2, 1 8 17: 328 (N. York, large size).

Squalus macrodous Mitchill, Amer. Month. Mag. Crit. Rev., 2, 1817: 328 (substitution for his earlier 5.
anuricanu!) .

Carcharias littoralis Bory de St. Vincent, Diet. Class Hist. Nat., 15, 1 829: 597 (ref.) ; DeKay, Zool. N. Y., 4,
1842: 351 (descr., N. York) ; Storer, Mem. Amer. Acad. Arts Sci., 2, 1846: 503; DeKay, Rep. State Cab.
Nat. Hist. N. Y. (1855), 8, 1858: 64 (in N. York list); Gill, Proc. Acad. nat. Sci. Philad., Addend.,
1861: 60 (name) and also subsequent eds.; Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 797 (name,
distrib.); Moore, Bull. U.S. Fish Comm. (1892), 12, 1894: 358 (N. Jersey); Linton, Bull. U.S. Fish
Comm. (1893), 13, 1894: 103 (parasites); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (l), 1896:
46 (descr., distrib.); Linton, Proc. U.S. nat. Mus., 20, 1897: 445, 450 (Woods Hole, parasites); Bean,
T. H., Bull. Amer. Mus. nat. Hist., p, 1897: 329, 371 (near N. York) ; Smith, Bull. U.S. Fish Comm.,
17, 1898: 89 (Woods Hole region) ; Jordan and Evermann, Bull. U.S. nat. Mus., 47 (3), 1898: 2747
(N. Carolina, large size); Parker and Davis, Proc. Boston Soc. nat. Hist., 29, 1899: 165, pi. I, fig. I;
pL 2, fig. 4; pi. 3, fig. 7, 10 (heart veins) ; Linton, Bull. U.S. Bur. Fish., rp, 1901: 273, 428 (Woods
Hole, food, parasites); Bean, Rep. For. Comm. N. Y., 1901: 379 (abundance, season, near N. York);
Gregg, Where to Catch Fish, 1902: 17 (named) ; Bean, T. H., Bull. N. Y. St. Mus., do, Zool. 9, 1903: 34
(general) ; Daniel, Elasmobranch Fishes, Univ. Calif. Press, 1934: 178 (veins) ; Sharp and Fowler, Proc.
Acad. nat. Sci. Philad., 56, 1904: 505, 506 (Nantucket) ; Wilson, Proc. U.S. nat. Mus., 28, 1905: 573
(parasites) ; Fowler, Rep. N. J. Mus. (1905), 1 906: 52 (N. Jersey) ; Tracy, Rep. R. I. Comm. inl. Fish.,
1906: 46 (Rhode Island) ; Fowler, Rep. N.J. Mus. (1906), 1 907: 254 (New Jersey) ; Wilson, Proc. U.S.
nat. Mus., 35, 1907: 414, 423, 430, 431 (Woods Hole, parasites) ; Smith, Bull. N. C. geol. econ. Surv., 2,
1907: 37 (C. Lookout) ; Field, Rep. U.S. Comm. Fish. (1906), 1907: 16 (food, Woods Hole) ; Sullivan,
Bull. U.S. Bur. Fish., 27, 1907: 10 (digest, syst.) ; Fowler, Proc. Acad. nat. Sci. Philad., 60, 1908: 54
(Nantucket and N. Jersey) ; Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 6 (New England rec-
ords) ; Fowler, Proc. Acad. nat. Sci. Philad., 61, 1909: 407 (C. May) ; Fowler, Rep. N. J. Mus. (1908),
1909: 352 (N. Jersey, food) ; Tracy, Rep. R. I. Comm. inl. Fish., 1910: 60 (Rhode Island) ; Linton, Bull.
U.S. Bur. Fish., 28 (2), 1910: l 200 (parasites) ; Sumner, Osbum and Cole, Bull. U.S. Bur. Fish., j/ (2),
1 91 3: 736 (Woods Hole, food); Gudger, Proc. biol. Soc. Wash., 26, 191 3: 98 (N. Carolina, sizes);
Nichols, Abstr. Linn. Soc. N. Y., 20-23, 1913:91 (season near N. York) ; Fowler, Copeia, 13, 191 4: not
numbered (N. Jersey) ; Coles, Proc. biol. Soc. Wash., 28, 1 91 5: 91 (C. Lookout, N. Carolina; eggs, em-
bryos, food) ; De Nyse, Bull. N. Y. zool. Soc, /p, 1916: 1425 (in aquarium) ; Latham, Copeia, 41, 1917:
17 (off N. York) ; Wilson, Proc. U.S. nat. Mus., 55, 191 7: 69 (N. Jersey, parasites); Latham, Copeia, 99,
1921: 72 (near N. York); Huntsman, Contr. Canad. Biol. (1921), 3, 1922: 56 (Passamaquoddy Bay);
Latham, Copeia, 118, 1923: 61 (Long Island, N. York) ; Nichols and Breder, Zoologica, N. Y., p, 1927:

io8 Memoir Sears Foundation for Marine Research

18 (N. York region); Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 20 (general); Wilson, Proc.
U.S. nat. Mus., 60 (5), 1922: 23, 24, 67 (Woods Hole, parasites); Breder, Copeia, 1932: 31 (Block
Island) ; Giltay, Mem. Mus. Hist. nat. Belg., Hors Ser., 5 (3), 1933: 8 (class.) ; Burton, Sci. Mon. N. Y.,
4°> 1935- 283 (in abundance near Charleston, S. Carolina); Smith, H. W., Biol. Rev., //, 1936: 52
(blood serum); Breder, Copeia, 194.2: 42 (shedding of teeth).

Triglochis taurus Muller and Henle, Arch. Naturg., (3) /, 1837: 396 (name); S. B. Akad. Wiss. Berlin,
1837: 114 (not seen); Gill, Ann. N. Y. Lye, 7, 1862: 398 (name); Canestrini, in Cornelia, et al.,
Fauna d'ltal., Pesci, 3, 1872: 44 (Medit.).

Odontasps taurus Miiller and Henle, Plagiost., 1 841: 73 (Medit., Atlant., C. Good Hope) ; Bonaparte, Cat.
Pesc. Europ., 1846: 17 (Medit.); Gray, List Fish. Brit. Mus., 1851: 63 (Cape seas); Dumeril, Hist,
Nat. Poiss., J, 1865: 417, pi. 7, fig. 3 (Medit., descr., teeth) ; Gervais and Boulart, Poiss., j^, 1877: 187
(Medit., Atlant.); Doderlein, Prosp. Metod. Pesci Sicil., 1878-1879: 30 (Sicily); Giglioli, Elenc.
Pesc. Ital., 1880: 52 (Medit., not seen) ; Doderlein, Man. Ittiol. Medit., 2, 1881: 58 (Medit.) ; Moreau,
Hist. Nat. Poiss. France, /, l88i: 291 (Algiers, descr.); Rochebrune, Act. Soc. linn. Bordeaux, (4) 6,
1882: 46; Faune Senegamb., Poiss., /, 1883-1885: 23 (Senegambia) ; Carus, Prod. Fauna Medit., 2,
1889-1893: 506 (Medit.); Carrucio, Boll. Soc. Zool. Ital., 77, 1910: 267-272 (Medit., not seen);
Roule, Bull. Inst, oceanogr. Monaco, 243, 1912: 9 (St. Lucia I., C. Verde); Metzelaar, Trop. Atlant.
Visschen, 1919: 189 (trop. W. Africa) ; Jordan, Copeia, 140, 1925: 20 (name) ; Copeia, 166, 1928: 4
(name) ; Belloc, Rev. des Trav. Peches Marit., 7 (2), 1934: 135 (Morocco, Senegal, Canaries) ; Norman
and Fraser, Giant Fishes, 1937: 8 (general) ; Fowler, Bull. U.S. nat. Mus., 100 {13), 1941: 120 (descr.,
synon. in part).

Carcharias griseus Ayers, Boston J. Nat. Hist., 4, 1843: 288, pi. 12, fig. 4 (Long Island, N. York, descr.);
Linsley, Amer. J. Sci., 47, 1844: 76 (same specimen as Ayers, 1843) ; Storer, Mem. Amer. Acad. Arts
Sci., 2, 1846: 504; Proc. Boston Soc. nat. Hist., 2, 1848: 256 (teeth) ; Mem. Amer. Acad. Arts Sci., 9,
1867: 219, pi. 36, fig. I; also Fishes Mass., 1867: 241, pi. 36, fig. 1 (descr., Massachusetts).

Odontasfis griseus Desor, Proc. Boston Soc. nat. Hist., 2, 1848: 264 (teeth).

C<wc/4i>r/ij/ /«-oa: Guichenot, Explor. Alger. Poiss., 1850: 124; not SfWtt/ /i;ro* Risso, 1810 (Algiers).^'

Odontasfis americanus Abbott, Proc. Acad. nat. Sci. Philad., 1 861 : 400 (N. Jersey, size) ; Dumeril, Hist. Nat.
Poiss., /, 1865: 419 (descr.) ; Abbott, in Cook, Geol. N. J., 1868: 828 (N. Jersey) ; Gunther, Cat. Fish.
Brit. Mus., 8, 1870: 392 (descr., synonyms in part) ; Ribero, Arch. Mus. nac. Rio de J., 14, 1907: 159,
202 (south. Brazil) ; Ribeiro, Fauna brasil. Peixes, 2 (l), Fasc. I, 1923: 15 (south. Brazil).

Eugomfhodus griseus Gill, Proc. Acad. nat. Sci. Philad., Addend., 1 86 1 : 60 (name) .

Eugomfhodus littoralis Gill, Proc. Acad. nat. Sci. Philad., 1863: 333 (name) ; Proc. Acad. nat. Sci. Philad.,
1864: 260 (name, syn.) ; Verrill and Smith, Rep. U.S. Comm. Fish. (1871-1872), 1873: 521 (Woods
Hole, food); Gill, Rep. U.S. Comm. Fish. (1871-1872), 1873: 813; Baird, Rep. U.S. Comm. Fish.
(1871-1872), 1873: 827 (Woods Hole); Bean, Proc. U.S. nat. Mus., 5, 1881: 115 (east coast, U.S.);
Wilson, Proc. U.S. nat. Mus., 5/, 1907: 711 (Woods Hole, parasites); Proc. U.S. nat. Mus., 60 (5),
1932: 67 (Woods Hole, parasites).

Carcharias americanus Jordan and Gilbert, Bull. U.S. Nat. Mus., 16, 1883: 27 (east coast, U.S.).

Odontasfis littoralis Jordan and Gilbert, Bull. U.S. nat. Mus., 1 6, 1 883 : 874 (name) ; Proc. U.S. nat. Mus., 5,
1883: 581 (S. Carolina) ; Stevenson, Trans. Vassar Bros. Inst., 2, 1884: 92, i pi. (Nantucket) ; Rathbun,
Proc. U.S. nat. Mus., 7, 1885: 490 (Vineyard Sound, parasites) ; Nelson, Rep. N. J. geol. Surv. (1888),
1890: 662 (N. Jersey record) ; Jordan, Guide to Study Fish., /, 1 905: 534 (general) ; Man. Vert. Anim.
NE. U.S., 1929: 12 (general) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930:

19 (distrib.) ; Norris, Plagiost. Hypophysis, 1 94 1 : pi. 1 8, fig. 69 (brain) ; Lunz, Bull. S. C. St. Planning
Bd., 14, 1944: 27 (S. Carolina, Florida).

Doubtful References:

Odontasfis taurus Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 46 (Senegambia; identity doubtful be-
cause color, as described, does not agree with that of taurus) .

27. Not seen; accordifig to Dumeril (Hist. Nat. Poiss., i , 1865 : 418) this specimen, preserved in Paris, was taurus,
not ferox.

Fishes of the Western North Atlantic 109

Odontasfis americanus Ihering, Rev. Mus. paul., 2, 1 897: 35 (listed, Rio Grande do Sul, Brazil) ; Thompson,
Mar. biol. Rep. Cape Town, 2, 1914: 146 (S. Africa, not seen) ; Lahille, Physis. B. Aires, 5, 1921: 63;
also Enum. Feces Cartilag. Argent., 1 921: 15 (Argentina); Devincenzi, An. Mus. Montevideo, (2) 2,
1926: 202 (size, weight, no descr., Uruguay).

Family SCAPANORHYNCHIDAE
Goblin Sharks

Characters. Two dorsal fins, the ist very much shorter than caudal, the rear end of
its base anterior to origin of pelvicsj caudal slightly more or less than Vs of total length,
its axis raised only very slightly, its lower anterior corner not expanded as a distinct lobe;
caudal peduncle not greatly depressed or expanded laterally; sides of trunk without longi-
tudinal dermal ridges; jaws greatly protrusible (much more so than in any other sharks)
and widely expansible; snout greatly elongate; 5th gill opening over or anterior to origin
of pectoral ; gill arches without rakers and not interconnected by a sieve of modified den-
ticles; nostrils entirely separate from mouth, their anterior margins without barbels;
spiracles present; lower eyelid without nictitating membrane or subocular fold; teeth simi-
lar in the two jaws, with thorn-like cusps, smooth-edged, with or without lateral denticles,
on broad bases; skull of normal shape {i.e.y not widely expanded laterally) ; rostral carti-
lages 3, united anteriorly as a long rod; radials of pectoral mostly borne on mesopterygium
and on metapterygium ; meso- and metapterygia not separated by a foramen ; heart valves
in 3 rows. Development not known, but probably ovoviviparous.

Genera. Only one genus, Scapanorhynchus Woodward, 1889, is known.

Range. Modern representatives of Scapanorhynchus^ are known from Japan, the
coast of Portugal, and perhaps from Australia/

Fossil remains of the genus, mostly from the Cretaceous, have been found at many
localities in Europe, North and South America, Asia, Africa and New Zealand.

Family ISURIDAE

Mackerel Sharks, Man-eater Sharks

Characters. Two dorsal fins, the ist much shorter at base than length of caudal, the
rear end of its base far in advance of origin of pelvics; 2nd dorsal and anal much smaller
than ist dorsal; caudal less than Vs of total length, lunate in form, its axis steeply raised;
caudal peduncle strongly depressed dorso-ventrally and widely expanded laterally, form-
ing a prominent keel on each side, extending well out on the caudal, with a less definite
longitudinal keel close below it on the anterior part of caudal in some species; sides of
trunk, anterior to anal, without longitudinal dermal ridges; upper and lower precaudal
pits well developed; inner margins of pelvics entirely separate, posterior to cloaca; snout
not very elongate and jaws not greatly protrusible; 5th gill opening in front of origin of

1. How many species these represent still remains unsettled.

2. According to Whitley (Fish. Aust, /, 1940: 136), the report of this Shark from Murray River (Zietz, Trans,
roy. Soc. S. Aust., 52, 1908 : 291) is open to doubt.

no Memoir Sears Foundation for Marine Research

pectoral ; gill arches without rakers and not interconnected by a sieve of modified denticles}
nostrils entirely separate from mouth, without barbels 5 spiracles present or absent} lower
eyelid without nictitating fold or membrane} both jaws with labial furrows at corners}
teeth large, few in number, awl- or blade-like, with i cusp} head of normal shape (not
widely expanded)} rostral cartilages 3, united at tip} metapterygium of pectoral with
about 3 times as many radials as mesopterygium, but the latter nearly as large as former}
meta- and mesopterygia not separated by a foramen } heart valves In 3 rows. Development
ovoviviparous.

Genera. One of the members of this family, set apart from all the others by its trian-
gular, serrate teeth, has long been considered as representing a well marked genus, Car-
charodon. The remaining isurids fall in two groups: (A) Very stout-bodied} first dorsal
originating over or anterior to inner corner of pectoral when latter is laid back} first two
teeth in each jaw similar in shape to subsequent teeth} caudal fin (so far as known) with a
secondary longitudinal keel on either side below the primary keel formed by the lateral
expansion of the caudal peduncle. (B) More slender-bodied} first dorsal originating defi-
nitely posterior to inner corner of pectoral} first two teeth in each jaw noticeably more
slender and more flexuous than the others} without secondary keels. It seems reasonable
to accept the difference between the two groups, and especially the presence or absence
of the secondary caudal keels, as sufficiently important for generic separation. This course
is followed here. Fortunately there has been no need to coin a new generic name in either
case.

Key to Genera

I a. Upper teeth broadly triangular with serrate edges.

Care Aar o<io« Agassiz, L., 1838, p. 133.

lb. Upper teeth slender, with smooth-edged cusps.

2a. First 2 teeth in each jaw similar in shape to the succeeding teeth} most or all of
teeth with lateral denticles in most species, and perhaps in all (lateral denticles in
young specimens may be so small as to be difficult to recognize} they may even be
lacking on some of the teeth) } origin of ist dorsal about over or anterior to inner
corner of pectoral when latter is laid back} anterior part of caudal fin with a sec-
ondary caudal keel on either side below the primary keel formed by the lateral
expansion of the caudal peduncle. Lamna Cuvier, 1 8 1 7, p. 1 1 1 .

2b. First 2 teeth in each jaw noticeably more slender and more flexuous than the
others} no lateral denticles on any of the teeth} origin of ist dorsal definitely
posterior to inner corner of pectoral when latter is laid back} caudal fin without
secondary keels, with only the primary keels formed by the lateral expansion of
the caudal peduncle. Isurus Rafinesque, 1 8 lO, p. 1 23.

Fishes of the Western North Atlantic iii

Genus Lamna Cuvier, 1 8 1 7

Lamna Cuvier, Regne Anim., 2, 1817; 126, 127; type species, Squalus cornuiicui Gmelin, 1789, equivalent
to Squalus nasus Bonnaterre, 1788.

Generic Synonyms:

Lamia Risso, Hist. Nat. Europe merid., 5, 1826: 123; type species, Squalus cornubicus Gmelin, 1789; not

Lamia Fabricius, 1775, for Coleoptera.
Selanonius Fleming, Hist. Brit. Anim., 1828: 169; type species, Selanonius walkeri Fleming, same as Squalus

nasus Bonnaterre, 1788.
Exoles Gistel, Naturg. Tier., 1848: 9; to replace Lamia Risso, 1826; preoccupied.

Generic Characters. Teeth slender, awl-shaped, smooth-edged, with lateral basal
denticles in most cases and perhaps in all, the first 2 teeth in each jaw similar in shape to
those succeeding, the anterior ones with two widely divergent roots, the third upper tooth
much smaller than second or fourth, but third lower tooth about same size as fourth j origin
of 1st dorsal over or anterior to inner corner of pectoral when latter is laid back; trunk
robust (Fig. 15); snout conical, pointed; caudal pits in the form of transverse furrows;
a less distinct secondary longitudinal keel, broadly triangular in cross section, on anterior
part of caudal on each side, close below the primary keel formed by the expanded caudal
peduncle, in all species so far known; upper jaw very slightly protrusible. Characters
otherwise those of the family.

Range. Widespread in boreal to warm temperate belts of the oceans in both hemi-
spheres; not known from tropical seas.

Sfecies. The genus Lamna is represented in the North Atlantic by the well known
Porbeagle (L. nasus, p. 112); in the North Pacific by a form that has usually been con-
sidered identical with nasus, but which has recently been found to be a distinct and well
marked species (L. ditropis Hubbs and FoUett, 1947);* in Australian-New Zealand
waters" and off Argentina* by close relatives whose precise relationships to nasus remain
to be determined; and in the eastern side of the South Pacific by a form (L. philippii Perez
Canto, 1886)^ resembling nasus in general appearance and in the position of the first
dorsal fin, but described and pictured as lacking lateral denticles on the teeth. Until it is
known whether this is actually the case, and whether or not philippii has the secondary
caudal keels (none are shown on the only published illustration of it), its status must re-
main problematical.

4. Hubbs and Follett, Copeia, 1947: 194.

5. L. luhitleyi Phillipps, N. Z. J. Sci. Tech., 16, 1935: 239, fig. 3; secondary caudal keels clearly shown in the
photograph.

6. Reported as nasus by Lahille (An. Mus. nac. B. Aires, ^4, 1928: 310, pi. 4) ; teeth described as with denticles;
secondary caudal keels clearly shown on the illustration.

7. Philippi, Anal. Univ. Chile, 71, 1887: 549, pi. 3, fig. 2.

112 Memoir Sears Foundation for Marine Research

Key to Species of the Northern Hemisphere'

I a. Distance from tip of snout to anterior edge of eye at least V2 as great as from posterior
edge of eye to ist gill opening, each measurement taken between perpendiculars;
lower surface plain-colored, without dark blotches.

nasus (Bonnaterre), 1788, p. 112.

lb. Distance from tip of snout to anterior edge of eye less than Vs as great as from pos-
terior edge of eye to ist gill opening; lower surface conspicuously marked with dark
blotches, except perhaps in very young specimens, diiropis Hubbs and Follett, 1 947.

Warm Temperate to Subboreal Belt, Both
Sides of North Pacific.

Lamna nasus (Bonnaterre), 1788

Mackerel Shark, Porbeagle; Blue Shark (in Gulf of Maine)

Figures 15, 16, 17

Study Material. Male, 935 mm., from Nahant, Mass. (Harv. Mus. Comp. Zool.,
No. 209) ; specimens of 660, 963 and 966 mm., from New England, North Atlantic and
Continental Slope off Sable I., Lat. 42° 37' N., Long. 60° 55' W. (U.S. Nat. Mus., No.
47528, 24288, 44057); head, from 70 miles SE. of Cape May (U.S. Nat. Mus., No.
125884) ; two embryos, no doubt of this species, about 180 mm. long, from Barnstable,
Mass., taken in October 1942 (Harv. Mus. Comp. Zool., No. 35901); also eight speci-
mens fresh caught in Gulf of Maine (not preserved) ; jaws from several of same; photo-
graph of i8o-mm. embryo from female caught off Portland, Maine, at "Mistaken
Ground" in January 1927 by Capt. D. C. Train.

Distinctive Characters. Easily separable from the Sharp-nosed Mackerel Shark by
its teeth (cf. Fig. 16 D with 19) ; from Carcharodon by the teeth (cf. Fig. 16 D with 20 B)
and by the relative positions of the second dorsal and anal fins.

Description. Proportional dimensions in per cent of total lengths. Male, 935 mm.,
from Nahant, Mass. (Harv. Mus. Comp. Zool., No. 209).

Trunk at origin of pectoral: hrea.dth 15.O; height 14.5.

Snout length in front of: outer nostrils 6.0; mouth 7.1.

Eye: horizontal diameter 2.4.

Mouth: breadth 8.1 ; height S-5-

Nostrils: distance between inner ends 3.6.

Labial furrow length: upper 2.2; lower 1.3.

Gill opening lengths : ist 7.5; 2nd 7.1 ; 3rd 6.5; 4th 6.$) 5th 6.5.

First dorsal fin: vertical height i i.O; length of base 9.1.

Second dorsal fin: vertical height 2.7 ; length of base 1.7.

Anal fin: vertical height 2.7; length of base 1.6.

8. It would be premature to expand the Key to the southern hemisphere for the reasons stated above.

Fishes of the Western North Atlantic

113

Caudal fin: upper margin 24.2; lower anterior margin 15.4.
Pectoral fin: outer margin 17.7; inner margin S-6\ distal margin 14.3.
Distance from snout to: ist dorsal 33.4; 2nd dorsal 66.3; upper caudal 76.O; pec-
toral 27.8 J pelvics 50.7} anal 67.0.

Inters-pace between: ist and 2nd dorsals 24.35 ^ri'i dorsal and caudal 9.2; anal and
caudal 9.2.
Distance from origin to origin of: pectoral and pelvics 26.3 ; pelvics and anal 1 6.4.

Trunk fusiform, much stouter than in Isurus oxyrinchus, its height opposite origin of
1st dorsal (where highest) about equal to distance from eye to 4th gill slit, or about 18%
of total length, tapering to a very slender caudal peduncle. Sides smooth, lateral line not
apparent. Caudal peduncle very strongly flattened dorsoventrally, widely expanded later-
ally and sharp-edged, with a less distinct longitudinal keel, broadly triangular in cross
section, on anterior part of caudal close below the rearward extension of the expanded
peduncle} this is more obvious in large specimens than in small j upper and lower precaudal

Figure 15. Lamna nasus, young male, 935 mm. long, from Nahant, Massachusetts (Harv. Mus. Comp.
Zool., No. 209). A Second dorsal and anal fins, about 0.4 x. B Caudal peduncle viewed from above, about
0.4 X. C Cross section of caudal peduncle at region indicated by transverse line in B. D Caudal peduncle
and base of tail, from the photograph of a fresh Gulf of Maine specimen about four feet long, to show
secondary caudal keel.

114 Memoir Sears Foundation for Marine Research

pits both strongly developed as transverse furrows, straight or only slightly curved, with
convexities directed rearward, the lower pit a little in advance of the upper. Dermal den-
ticles so small and flat that the skin is velvety to the touch, each a little broader than long,
with 3 teeth, the median a little longer than others, and a corresponding number of low,
sharp-topped ridges, separated by broad valleys 5 pedicels moderately long, on broad bases.
Head and snout conical, the length of head to pectoral a little less than V^ of total
length. Snout pointed, its length in front of mouth about Y^ of head to origin of pectorals.
Eye circular, its diameter about 30% as long as snout in front of mouth. Spiracles lacking
in specimens studied but described as sometimes present as minute pores behind eyes. First
gill opening slightly longest, about as long as snout in front of eye, the lower end of 5th
curving rearward and ventrad for a short distance around origin of pectoral, the space
between 4th and 5th only about V2 that between ist and 2nd, the 5th more oblique than
others. Nostrils approximately transverse, hardly Vs as long as distance between them,
their inner corners about ^/4 as far from mouth as from tip of snout, the anterior margin
with a low rounded lobe (Fig. 16 B). Mouth broadly rounded, about lYo times as broad
as high (thus somewhat shorter, relatively, than in oxyrinchus). Upper labial furrow
about 0.3 as long, the lower 0.2 as long, as distance to symphysis of the respective jaws, the
upper partly, and the lower almost entirely, concealed when mouth is closed.

^^V^^^^^ffW^

Figure 16. Lamna rtasus. A Lower view of head of young male from specimen shown in Fig. 15. B Right-hand
nostril, about 2.5 x. C Dermal denticles — -general view, about 40 x; apical view, about 80 x. D Upper and
lower teeth of a larger specimen from Platts Bank, Gulf of Maine, about 1.3 x.

Fishes of the Western North Atlantic 115

Teeth alike in the two jaws, j^2o '" specimens counted; no median tooth in either
jaw; all teeth except those next to corners of jaw slender, narrow, straight, sharp-pointed,
with a small sharp basal denticle on either side in adults, which, however, may be lacking
on some or all of teeth in young specimens, or at least so small that their detection is diffi-
cult; 1st and 2nd teeth in each jaw largest, 3rd upper tooth much smaller than 2nd or
4th and sometimes lacking, but 3rd lower tooth about equal to 4th ; 4th to 8th or 9th about
equal in size in each jaw, but lOth and subsequent teeth progressively smaller; lateral
teeth in both jaws, as well as anterior teeth in lower jaw, erect, but ist, 2nd and sometimes
3rd upper teeth directed sharply inward; i, or rarely 2, rows functional along sides of
mouth, but 2, or rarely 3, rows near the center.

Origin of ist dorsal over or very slightly posterior to axil of pectoral (thus relatively
much farther forward than in oxyrinchus) , its anterior margin slightly convex, its apex
broadly rounded, its rear margin straight toward apex but moderately concave toward base,
its free rear corner about Vs ^s long as its base, its vertical height nearly equal to distance
from eye to ist gill or about 60% as great as length of pectoral. Second dorsal about V4 as
high as 1st, its origin over origin of anal, its apex broadly rounded, rear margin deeply
concave, its free rear corner about i V2 times as long as base, but only moderately slender.
Lower lobe of caudal about 64 to 75% as long as upper, relatively shorter in young than
in older specimens (about 68% in Fig. 15), each measured from the respective precaudal
pit (thus somewhat shorter, relatively, than in oxyrinchus), the subterminal notch strongly
marked, the posterior outline subangular, with rounded corner. Anal slightly larger than
2nd dorsal, similar in shape. Pelvics with rounded corners and moderately concave outer
margins, their origins posterior to rear tip of ist dorsal by a distance about % to % as long
as from tip of snout to mouth. Pectoral nearly or quite as long as from posterior margin of
eye to 5 th gill opening, about Y2 as broad as long, the anterior margin moderately convex,
the tip and inner corner rounded, the distal margin only moderately concave (less so than
in oxyrinchus).

Color. Dark bluish-gray above, changing abruptly on the lower sides to the white
of the lower surface; pectorals dusky on outer half or third, the anal white or slightly
dusky.

Size. While nasus has repeatedly been reported to reach a length of 12 feet, a lO-foot
female from Monhegan, Maine," is the largest of which we find a definite record. How-
ever, at least one other of eight feet has been positively reported from the Gulf of Maine,
and a number from seven to nine feet (up to 2,800 mm.) at different times from northern
European waters. Very few, however, of those caught in the western side of the Atlantic
are more than six feet long, with four to five feet perhaps the commonest size. For exam-
ple, none of those that we have hooked has been longer than five feet, apart from one of
perhaps eight feet hooked and lost over Cashe's Ledge on September 30, 1927. At the
other extreme the smallest on record is 29 inches. Information as to the relationship be-

9. Hubbs, Copeia, 123, 1923: loi.

ii6

Memoir Sears Foundation for Marine Research

tween length and weight in nasus is scant. Reported weights of about 400 pounds at nine to
ten feet, and 305 pounds at eight feet three inches would suggest that this is a much lighter
fish than oxyrinchus. But since the stoutness of its trunk suggests rather the reverse, it
seems more probable that the few reported weighings have been of fish that had been
gutted, which, in the case of a shark with so large a liver, means the loss of a large part of
the total weight. Females may contain embryos at a length of five feet.

Figure 17. Lamna nasus, embryo, about 1 80 mm. long, from Barnstable, Massachusetts (Harv. Mus. Comp.
Zool., No. 35901), about 0.8 natural size.

Develo-pmental Stages. It has long been known that this is an ovoviviparous species,
the young lying free in the uterus without connection with the mother. It also seems estab-
lished that in nasus, contrary to the rule among most other ovoviviparous sharks, the yolk
sac is absorbed and the umbilical cord entirely obliterated while the embryo is still very
small (55 to 60 mm.) and still with well developed external gills. Thereafter, the embryo
nourishes itself by swallowing the unfertilized eggs which lie close to it in the uterus, the
result being that its stomach becomes enormously swollen by the masses of yolk so swal-
lowed, forming a so-called "yolk stomach"}" as the embryo grows this increases in relative

10. For more detailed accounts see Swenander (Zool. Stud. TuUberg, Uppsala, 1907 : 283, pi. 1) ; also, on the North
Pacific form, see Lohberger (Abh. bayer. Akad. Wiss., Suppl., f, Abt. 2, 1910).

Fishes of the Western North Atlantic 117

size from about 45 mm. in length (in one of 180 mm., Fig. 17) to about 235 mm. (in one
of 400 mm.") or to more than half the total length. The throat region of the embryo too
is enormously expanded, giving it a most grotesque appearance. Also, the caudal fin is at
first much more asymmetrical than in the adult, assuming the lunate form with growth
of the embryo, and the young are very large at birth, witness embryos of 19, 24 and 18
inches in a five- foot female.^^ A Gulf of Maine female of 10 feet contained a 20-pound
embryo.^' Corresponding to the large size of the embryo, gravid females normally contain
only one to four young (0-2 per oviduct), although five have been reported."

Habits. This has been described repeatedly as an active, strong-swimming species
when in pursuit of its prey. When hooked, however, it puts up only a very feeble resist-
ance, as we have experienced. We have never seen or heard of one jumping in its attempt
to escape, as the Mako does (p. 128). Nor is there any difficulty in landing specimens of
four to five feet on an ordinary hand-line j in fact, it is as proverbial for its sluggishness
under such circumstances as is the Mako for its activity.

Mackerel Sharks are often seen finning at the surface on calm days; on the other
hand, many have been caught on bottom with cod and halibut lines as well as at mid-depths
now and then in drift nets in northern European waters, while one is occasionally entangled
in a mackerel net. Evidently, then, their depth range is from the surface down to bottom;
on the cod fishing grounds that would be to some 70 to 80 fathoms at least; it is not known
how much deeper they descend.

In the waters of northern Europe gravid females have been taken from localities so
widely scattered as to show that the species produces young throughout its East Atlantic
range. Presumably this is true in the western Atlantic also, although embryos have actually
been recorded only from the vicinity of Monhegan Island, Maine, in August, from off
Portland, Maine, in November and in January (see p. 119) and from Barnstable, Massa-
chusetts, in October (see Study Material, p. 112). In Europe, females with embryos
have been reported for the winter months as well as for summer. But the fact that the
largest embryos have been found in summer indicates the latter as the chief season of
production.

Lamna nasus preys largely on schools of mackerel, herring and (in the eastern At-
lantic) pilchards; also on such ground fish as cod, hake, cusk, and other gadoids, flounders,
or any other fish that may be available, and on squid. In the eastern Atlantic its diet also
includes whiting (Gadus merlangus), spiny dogfish {Squalus acanthias) and John dory
{Zeus jaber). It also has the troublesome custom of foraging on cod, etc., that have been
hooked on long lines, biting off the snoods in the process.

Relation to Man. During the first quarter of the last century the liver oil of this
species, mixed with other fish oils, was in considerable demand (chiefly for tanning pur-

11. Nordgard, K. norske Vidensk-Selsk. Skr. Trondh. (1923— 1924), 1925: 38, fig. 22.

12. Shann, Rep. Fish. Bd. Scot., 1911: 73, pi. 9. 13. Hubbs, Copeia, 123, 1923: loi.

14. Lijbbert and Ehrenbaum, Handb. Seefisch. Nordeurop., j, 1936: 278. A shark with 10 embryos reported long
ago as this species (Wilder, Science, /, 1880: 236) probably was some other.

1 1 8 Memoir Sears Foundation for Marine Research

poses). Provincetown was the center of activity. However, this local demand for shark-
liver oil had almost entirely died before 1850. Of late years the only commercial impor-
tance of nasus in the western Atlantic (except as a nuisance to fishermen) has been its sal-
ability in the larger fish markets, for its range does not extend southward far enough to
bring it within the scope of the shark-leather industry that is now operating there. In
northern Europe, on the west coast of France, as well as in the Mediterranean, the flesh of
nasus is in much greater demand} this is especially so in Germany, where the local supply
was regularly augmented by considerable imports from Norway^° before the war. The
Norwegian catch has been made chiefly on long lines, the German catches chiefly in herring
trawls. The few that are caught in American waters are taken incidentally either on hand-
lines when fishing for cod, etc., or in mackerel nets. It is not game enough to be of inter-
est to sport-anglers.

Range. Continental waters of the northern North Atlantic, from the Mediterranean
and northwestern Africa to the North Sea, Scotland, Orkneys, and southern Scandinavia,
on the eastern sidej less common north to Iceland, northern Norway and the Murman
Coast} from the Newfoundland Banks and Gulf of St. Lawrence in the west south to New
Jersey, and perhaps to South Carolina." It is represented in the North Pacific from north-
ern California to southern Alaska, Kamchatka and Japan, as well as in the Australian-New
Zealand region," by forms very closely allied, but not identical.

Occurrence in the Western Atlantic. The area of regular occurrence for nasus is con-
fined to a much narrower latitudinal belt in the West than in the East, i.e.y from southern
New England to the outer coast of Nova Scotia and the Gulf of St. Lawrence, with the
chief center of population lying in the western side of the Gulf of Maine. Thus, while
there are but two records of it from the Newfoundland Banks,'* and one, except for vague
reports, from the Gulf of St. Lawrence, fishermen report it as the commonest large shark
in summer along the Atlantic coast of Nova Scotia, including Cape Breton. Apparently it
tends to shun the cold waters of the Bay of Fundy region, there being but one positive
record for it in Passamaquoddy Bay. Farther west, however, in the Gulf of Maine, it is so
numerous on occasion that there is record of incidental catches of 1 9 in one night by six
men working hand-lines, with about 1 50 taken on cod-lines by a crew of fishermen on a
three weeks' trip near Monhegan Island, Maine. During the cruises of the United States
Bureau of Fisheries vessels we have seen and caught them most often in the immediate
vicinity of Platts Bank oflF Cape Elizabeth. It is certainly the most often seen of the larger
sharks around the Isles of Shoals and near Cape Anne, while in Massachusetts Bay it has

15. Exact amounts are not available, because the landings of all species of sharks are combined in the published
statistics.

16. A shark reported under the name Lamia nasus from Argentina by Lahille (An. Mus. nac. B. Aires, 34, 1928:
310) appears actually to have been a Make (see p. 130).

17. See p. III.

18. This includes one of 7 feet loj^ inches, taken June 28, 1946, in Lat. 44° 27' N., Long. 50° 00' W., reported
by Dr. A. M. Ramalho of Lisbon, who sent us one tooth.

Fishes of the Western North Atlantic 119

been described repeatedly as "common."^" We have hooked or sighted on an average
about one per three or four days on the cod fishing grounds, generally in the western side
of the Gulf of Maine and on Nantucket Shoals during the summers of 1924 to 1930.^°
However, the fact that such large numbers have been caught in the past within brief periods
(see above) is sufficient evidence that their numbers vary widely from year to year, or over
a period of years, at least locally.

To the westward nasus is described (we have no first-hand information) as compara-
tively common in the vicinity of Woods Hole (more so in autumn) and it has been re-
ported on several occasions from Rhode Island coastal waters. However, it appears only
as a stray along Long Island, New York (one record), or along the New Jersey coast; the
only evidence of its presence farther south is one somewhat doubtful report of it off
Charleston, South Carolina. From this it appears that the isothermal belt of about 6s° F.
limits its normal range to the southward.

It seems equally certain that its on-and-off-shore range is similarly narrow, for we
find no record of it (nor report of it by fishermen), from the offshore fishing banks abreast
of the Gulf of Maine (Georges and Browns Banks) ; only one is reported from the Nova
Scotian slope (see Study Material, p. 112), and two from the Grand Banks. On the other
hand, few venture close enough to land to be picked up in the pound nets. There is, how-
ever, record of a Mackerel Shark, probably this species, which was entangled in the eel
grass (Zoslera) in Barnstable Harbor, Massachusetts, many years ago." In the western
Atlantic all published records of it, and those that we have observed, have been for the
warm half of the year, but its presence in the Gulf of Maine in winter is proved by our
receipt of a photograph of an embryo, certainly of this species, from a female caught off
Portland, Maine, in January of 1927. Similarly, it is taken in winter as well as in summer
off northern Europe, but less commonly. This, together with the absence of any evidence
of migration southward along the middle Atlantic coast of the United States, suggests
that in winter they simply descend into deeper water to avoid low surface temperatures,
apparently feeding little then, otherwise more of them would be picked up by the winter
fishery for hake.

Synonyms and References:

Porbeagle, Borlase, Nat. Hist. Cornwall, 1758: 265, pi. 26, fig. 4 (Cornwall) ; Pennant, Brit. Zool., 3, 1 769:

92 (descr., Cornwall) ; also later eds.
Squdus glaucus Gunnerus, K. norske Vidensk.-Selsk. Skr. Trondh., 1768: I, pi. I (descr., embryo, Nor-

19. Actually, no sharks, other than the Spiny Dogfish, are ever common off the northeastern coast of the United
States or Canada, in the sense in which that term can be applied to such fish as the cod, mackerel, etc., but only
as relative to other sharks of corresponding sizes.

20. Cod-tagging cruises of the U.S. Bureau of Fisheries.
»i. Goode, Fish. Fish. Industr. U.S., 1884: 670, footnote.

I20 Memoir Sears Foundation for Marine Research

way, not seen); Skr. Norg. Gesellsch. Wiss. Kbh. and Leipzig, 1770: I, pi. I, fig. 1-5 (translation of
foregoing); Ascanius, Icon. Rerum Natur. Kbh., 1777: pi. 31 ; Str0m, K. norske Vidensl«.-Selsk. Skr.,
N.S. 2, 1788: 340, pi. 2, figs. (Norway, not Squalus glaucus Linnaeus, 1758).

Beaumaris Shark, Pennant, Brit. Zool., 3, 1 776: 104, pi. 17 (descr., Wales).

Le Nez, Broussonet, Mem. Math. Phvs. Acad. Sci. Paris, 1780: 667; Sonnini, Hist. Nat. Poiss., 4, 1801-
1802: 5 (descr., Cornwall).

Touille-boeuf ou Loutre de Mer, Duhamel, Traite Gen. Peches, 4 (2), sect. 9, cap. 5, 1782: 298, pi. 20,
fig- 4 (general).

Squalus nasus Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: 10, pi. 85, fig. 350 (descr., type local., Wales,
by ref. to Pennant, 1776).

Squalus cornubicus Gmelin, in Linnaeus, Syst. Nat., 5, 1789: 1497 (descr., Cornwall) ; Goodenough, Trans.
Linn. Soc. Lond., Zool., 5, 1797: 80, pi. 15 (English Channel); Bloch and Schneider, Syst. Ichthyol.,
1801: 132; Sonnini, Hist. Nat. Poiss., 4, 1801-1802: 5 (descr.); Bosc, Nouv. Diet. Hist. Nat., 21,
1803: 185 (diagn.) ; Latreille, Nouv. Diet. Hist. Nat., 24, 1 804: 72 (in table of contents) ; Shaw, Gen.
Zool., 5 (2), 1804: 350 (general); Donovan, Nat. Hist. Brit. Fish., 5, 1807: pi. 108 (ill.); Turton,
Brit. Fauna, 1807: 113 (color); Risso, Ichthyol. Nice, 1810: 29 (Medit.) ; Neill, Mem. Werner. Soc.
Edinb., /, 1811: 549 (Scotland); Couch, Trans. Linn. Soc. Lond., Zool., 14, 1825: 91 (Cornwall);
Nilsson, Prod. Ichthyol. Scand., 1832: 116 (mention); Jenyns, Manual Brit. Vert. Anim., 1835: 500
(Gt. Brit., size) ; L. Agassiz, Poiss. Foss., j. Atlas, 1835: pi. G, fig. 3 (teeth) ; Wright, Fries and Ekstrom,
Skand. Fisk., 5, 6, 1838-1840: 135, pi. 30 (descr., Scandinavia); de la Blanchere, Diet. Peches, 1868:
747 (descr.); Gatcombe, Zoologist, (3) 5, 1881: 425 (English Channel).

Squalus fennanti Walbaum, P. Artedi Genera Pise. Emend. Ichthyol., j, 1792: 517 (descr., Atlant.).

Squale Long-nez, Lacepede, Hist. Nat. Poiss., 4° ed., j, 1798: 216, pi. 2, fig. 3 (descr.).

Beaumaris, Sonnini, Hist. Nat. Poiss., 4, 1801-1802: 8 (descr., Gt. Brit.).

Squalus monensis Shaw, Gen. Zool., 5 (2), 1804: 350 (general) ; Pennant, Brit. Zool., 5, 1812: 154, pi. 20
(descr., Wales) ; Cuvier, Regne Anim., 2, 1817: 127 (footnote).

Squalus cornubiensis Yennant, Brit. Zool., 5, 1 81 2: 152 (descr., Cornwall).

Squalus solanonus Leach, Mem. Werner. Soc. Edinb., 2, 1814; 64, pi. 2, fig. 2 (Scotland).

Carcharhinus cornubicus Blainville, Bull. Soc. philom. Paris, 1816: 121 (name).

Lamie nez., Cloquet, Diet. Sci. Nat., Atlas Poiss., 1816-1830: pi. 28.

Lamna cornubica Cuvier, Regne Anim., 2, 1817: 127; Cloquet, Diet. Sci. Nat., 25, 1822: 183 (general);
Fleming, Hist. Brit. Anim., 1828: 168; Ekstrom and Sundevall, Skand. Fauna, 1834: 67, pi. 30 (descr.,
Scandinavia, not seen) ; Bonaparte, Icon. Faun. Ital., i (2), 1835: plate not numbered (descr., Medit.) ;
Yarrell, Brit. Fish., 2, 1836: 384 (descr., Gt. Brit.); Kr0yer, Danmarks Fisk., j, 1st ed., 1838: 852
(around Denmark); Parnell, Mem. Werner. Soc. Edinb., 7, 1838: 413 (Gt. Brit.); L. Agassiz, Poiss.
Foss., 5, 1838: 86 (ref. to j, pi. A, should read pi. G, fig. 3, "Lamna") ; Bonaparte, Mem. Soc. neuchatel
Sci. Nat., 2 (8), 1839:9 (in synopsis) ; Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1839: 314 (general);
Owen, Odontogr., 1 840-1 845: 28 (teeth) ; Miiller and Henle, Plagiost., 1841:67 (descr.) ; Yarrell, Brit.
Fish., 2nd ed., 1841: 515 (descr., Gt. Brit.); Mayer, Neue Notiz. Natur. u. Heilk. (Froriep) Weimar,
2/ (446), 1842: 81; Agassiz, L., Poiss. Foss., j, 1835-1843: 287, pi. G, fig. 3 (teeth); Hamilton, Brit.
Fish., 2, 1843: 309 (general); also subsequent eds.; Bonaparte, Cat. Pesc. Europ., 1846: 18 (Medit.,
N. Europ.); Thompson, Zoologist, 8, 1850: 2970 (English Channel); Baikie, Zoologist, 11, 1853:
3846 (Orkneys); Kr0yer, Danmarks Fisk., 2nd ed., j, 1852-1853: 852 (Denmark); Owen, Cat. Osteol.
Roy. Coll. Surg., 1853: 95 (cranium, vertebrae); Nilsson, Skand. Fauna Fisk., 4, 1855: 718 (Scandina-
via) ; Thompson, Nat. Hist. Ireland, 4, 1856: 251 (Ireland) ; Machado, Feces Cadiz, 1857: 9 (Spain) ;
Jouan, Mem. Soc. Imp. Sci. Nat. Cherbourg, 7, :86o: 139 (English Channel) ; Nardo, Atti 1st. veneto,
(3) 5> 1 859-1 860: 787 (Medit.) ; Fitzinger, Bild. Atlas Naturg. Fische, 1864: fig. 172 (ill.) ; Perugia,
Cat. Pesc. Adriat., 1 866: 7 (Adriatic, not seen) ; Dumeril, Hist. Nat. Poiss., j, 1 865 : 405 (descr.) ; Bocage
and Brito Capello, Poiss. Plagiost., 1866: 12 (Portugal) ; Couch, Fish. Brit. Isles, /, 1867: 4i> pl- 8 (descr.,
Gt. Brit.) ; Brito Capello, J. Sci. math. phys. nat. Lisboa, 2 (7), 1869: 139 (Portugal) ; Gunther, Cat.
Fish. Brit. Mus., 8, 1870: 389 (descr., synon., distrib.) ; Van Beneden, Poiss. Cotes Belg. Parasit., 1870: 8
(parasites, Belgium) ; Ninni, An. Soc. Nat. Modena, 5, 1870: 66 (Medit.) ; Canestrini, in Cornalia, ct al..

Fishes of the Western North Atlantic I2i

Fauna d'ltal., 1872: 45 (Medit.) ; Gulliver, Quart. J. Micr. Sci., N. S. 12, 1872: 40 (size); Collett,
Norges Fisk., 1 875: 208 (Norway); Turner, J. Anat., Lond., 9, 1875: 301 (spiracle); Gervais and
Boulart, Poiss., 3, 1876: 1 80, pi. 68 (descr.) ; Clement, Bull. Soc. Sci. nat. Nimes, 4, 1876: 109 (not
seen); Reguis, Ess. Hist. Nat. Provence, (l) /, Ease. 1, 1877:48 (Medit.) ; Doderlein, Prosp. Metod. Pesci
Sicil., 1878: 30 (Medit.); Goode and Bean, Bull. Essex Inst. Salem, //, 1879: 29 (Gulf of Maine);
Campbell, Sci. Gossip, 16, 1880: 277 (Scotland, not seen); Brito Capello, Cat. Peixes Port., 1880: 44
(Portugal) ; Stossich, Boll. Soc. adriat. Sci. nat. Trieste, 5, 1880: 67 (Adriatic) ; Day, Fish. Gt. Brit., 2,
1880-1884: 297, pi. 156 (descr., Gt. Brit.); Buckland, Hist. Brit. Fish., 1881: 217 (descr.); Moreau,
Hist. Nat. Poiss. France, /, 1881: 296 (descr., distrib.) ; Doderlein, Man. Ittiol. Medit., 2, 1881: 60
(Medit.);" Hasse, Naturl. Syst. Elasm. besond. Theil., 1882: 215, pi. 28, fig. 1-6 (vertebrae); Jordan
and Gilbert, Bull. U.S. nat. Mus., :6, 1883: 30 (descr., N. Atlant., Woods Hole); Goode, Fish. Fish.
Industr. U.S., I, 1884: 670, pi. 249 (ill., habits, fishery) ; Storm, K. norske Vidensk-Selsk. Skr. Trondh.
(1883), 1884:45 (Trondhj em, Norway); Pilliet, Bull. Soc. zool. Fr., 10, 1885: 283 (anat.); Rathbun,
Proc. U.S. nat. Mus., 7, 1885: 488 (Vineyard Sound) ; Ehlers, Nachr. Ges. Wiss. Gottingen, /<?, 1886:
547 (Friesland, size); Whiteaves, Cat. Canad. Pinnep. Cetacea Fish., 1886: 5 (Gulf of St. Lawrence);
Jordan, Rep. U.S. Comm. Fish. (1885), 1887:797 (distrib.); Calderwood, 6th Ann. Rep. Fish. Bd. Scot.,
(3), 1888:263 (embryo, Scotland) ; Bean, T. H., Bull. U.S. Fish Comm., 7, 1889:256 (off N. Jersey) ;
Carus, Prod. Faun. Medit., I 889-1 893 : 505 (Medit.) ; Ewart, J. Anat., Lond., 24, 1 890: 227 (spiracle) ;
Bellotti, Atti Soc. ital. Sci. nat., 55, 1891 : 1 10 (Atlant. coast, France) ; Weber, Tijdschr. ned. dierk. Ver.,
(2) 4, 1893: IV (Holland); Duerden, Irish Nat., 5, 1894: 264 (Ireland); Dean, Fishes, living and
extinct, 1895: 90 (general); Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 218
(distrib.); Bull. U.S. nat. Mus., 47 (i), 1 896: 49 (descr., distrib.); Kermode, Zoologist, (4) /, 1897:
597 (Isle of Man); Knipovich, Annu. Mus. zool. Acad. St. Petersb., 2, 1897: 154 (Murman coast);
Jungersen, Danish "Ingolf" Exped., 2(1), 1899: 37, pi. 2, fig. 22, 23 (claspers) ; Yung, Arch. Zool. exp.
gen., (3) 7, 1899: 140 (physioL); Bassett-Smith, Proc. zool. Soc. Lond., 1899: 464, 469, 479 (para-
sites); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (4), 1900: pi. 6, fig. 22 (ill.); Warren, Irish
Nat., p, 1900: 293 (Ireland) ; Bean, T. H., Rep. For. Comm. N. Y., 1901 : 380 (Long Island, N. York) ;
Huber, Z. wiss. Zool., yo, 1901: 617, pi. 27, fig. 6 (claspers) ; Prince and Knight, Contr. Canad. Biol.,
1901: 55 (fins. New Brunswick); de Braganza, Result. Invest. Sci. "Amelia," 2, 1904: 54 (Portugal);
Bean, Bull. N. Y. St. Mus., 60, Zool., 9, 1903: 39 (New York); Drzewina, Arch. Zool. exper. gen.,
(4) 3, 1905: 226, 243 (anat.); Evans, Ann. Scot. nat. Hist., 1905: 56 (Scotland); Borca, Arch. Zool.
exp. gen., (4) 4, 1905: 204, 207 (France) ; Collett, Medd. Norges Fisk., SuppL, 3, 1905: 77 (size, food,
embryos, season, Norway) ; Fowler, Rep. N. J. Mus. (1905), 1906: 56 (New Jersey record) ; Swenander,
K. norsk. Vidensk.-Selsk. Skr. Trondh. (1905), 9, 1906: 73; Fowler, Science, N. S. 24, 1906: 596 (N.
Jersey) ; Rep. N. J. Mus. (1906), 1907: 254 (N. Jersey) ; Sullivan, Bull. U.S. Bur. Fish., 27, 1907: lO,
15, 23 (anat.); Swenander, Zool. Stud. Tullberg, Uppsala, 1907: 283, pi. I (embryo); Wilson,
Proc. U.S. nat. Mus., 33, 1907: 386, 410, 423 (parasites, Woods Hole region); Winther, Hansen
and Jensen, in Schi0dte and Hansen, Zool. Danica, 2, 1907: 296, pi. 27, fig. 2 (descr., distrib.); Ken-
dall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), igo8: 7 (off New England); Fowler, Proc. .■\cad. nat.
Sci. Philad., 60, 1908: 55 (Italy); Tracy, Rep. R. I. Comm. inl. Fish., 1910: 60 (Rhode Island);
Gunther, Encycl. Brit., nth ed., 24, 191 1 : 807 (general); Shann, Rep. Fish. Bd. Scot., 3, 191 1 : 73,
pi. 9 (embryos, Scotland) ; Arioh, Atti Soc. ligust. Sci. nat. geogr., 1913: 3 (Medit., not seen) ; Sumner,
Osburn and Cole, Bull. U.S. Bur. Fish. (191 1), j/ (2), 1913: 736 (Woods Hole region) ; Halkett, Check
List Fish. Canad., 1913: 40 (Canada) ; Le Danois, Ann. Inst, oceanogr. Monaco, 5 (5), 1913: 18 (Eng-
lish Channel) ; Leigh-Sharpe, J. Morph., 35, 1921: 367 (claspers) ; Shann, Proc. zool. Soc. Lond., 1923:
161 (embryos); Burne, Philos. Trans., (B) 2/2, 1923: 209 (blood system); Jenkins, Fish. Brit. Isles,
1925: 311 (Gt. Brit.); Elmhirst, Scot. Nat., 161, 1926: 151 (Scotland); Ehrenbaum, in Grimpe and
Wagler, Tierwelt N'.- u. Ostsee, Lief 7 (12"^), 1927: 14 (general); Nordgaard, K. norske \'idensk.-
Selsk. Skr. Trondh. (1925-1926), I, 1927: 38 (embryo); Ehrenbaum, Rapp. Cons. Explor. Mer.,
Fjune Ichthyol. N. Atlant., 1929: plate not numbered (descr., ill., distrib.); Nordgaard, K. norske

23. See Doderlein, 1881, for additional references for the Mediterranean in publications not accessible to us.

122 Memoir Sears Foundation for Marine Research

Vidensk.-Selsk. Skr. Forh., Trondh., /, 1929: 206 (embryos); Wilson, Proc. U.S. nat. Mus., 60 (5),
1932: 23, 59 (parasites) ; Belloc, Rev. des Trav. Peches Marit., 7, Fasc. 2, 1934: (Morocco to Senegal) ;
Borri, Mem. Soc. tosc. Sci. nat., 44, 1934: 92 (Medit.) ; Daniel, Elasmobranch Fishes, Univ. of Calif.
Press, 1934: 9 (teeth, body form); Nobre, Fauna Marinha Port., Vert., /, 1935: 430, pi. 58, fig. 182
(descr., Portugal) ; Norman and Fraser, Giant Fishes, 1937: 12 (general).

Lamia cornubica Risso, Hist. Nat. Europ. Merid., 5, 1826: 1 24 (Medit.); Bory de St. Vincent, Diet. Class.
Hist, nat., 75, 1829: 596.

Selanonius walkeri Fleming, Hist. Brit. Anim., 1828: 169 (Gt. Brit.).

Lamia monensis Bory de St. Vincent, Diet. Class. Hist, nat., 15, 1829: 596.

Squalus (Carcharias) cornubicus Blainville, in Vieillot, Faune Frang., 1825: 96, pi. 14,^* fig. 2 (descr.).

Lamna monemis YnTieW, Brit. Fish., 2, 1836: 387 (Wales); Bowerbank, Zoologist, (2) 8, 1873: 3617 (Eng-
lish Channel, size); Zoologist, (2) 9, 1874: 4202 (English Channel).

Lamna functata Storer, Mass. Zool. Bot. Surv., Rep. on Fish., 1839: 185, pi. 3, fig. 2 (ill., Mass. Bay) ; Boston
J. Nat. Hist., 2, 1839: 534, pi. 8, fig. 2 (descr., size, habits, utilization, Massachusetts) ; Linsley, Amer. J.
Sci., 4-j, 1844: 76 (Conn.) ; Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 504 (part, L. nasus and
/. oxyrinchus combined; refs. also incl. Squalus ptnctatus Mitchill, 1815, which was perhaps Scoliodon
terrae-novae) ; Mem. Amer. Acad. Arts Sci., N. S. 9 (i), 1867: 225, pi. 37, fig. I; also in Fishes
Mass., 1867: 249, pi. 37, fig. I (descr., ill., Massachusetts) ; Smith, S. I., Rep. U.S. Comm. Fish. (1871-
1872), 1873: 576 (parasites, Woods Hole region); Norman and Fraser, Giant Fishes, 1937: 12 (east
coast, U.S.).

Squalus {Lamna) cornubicus Cuvier, Regne Anim., Poiss., I 849: 362, pi. 114, fig. 3 (general).

Lamna fennanti Desvaux, Essai Ichthyol. France, 1851 : 23 (not seen).

Isurus cornubicus Gray, List. Fish. Brit. Mus., Chondropt., /, 1851: 58; Smitt, Hist. Scand. Fish., 2nd ed.,
2, 1895: I I 38, pi. 51, fig. I (descr., habits, Scandinavia); Mcintosh, Ann. Mag. nat. Hist., (7) ro,
1902: 254 (food, Scotland) ; Rey, Fauna Iberica Feces, /, 1928: 383 (descr.) ; Liibbert and Ehrenbaum,
Handb. Seefisch. Nordeurop., 2, 1936: 278 (general. North Sea).

Oxyrhina daekayi Gill, Proc. Acad. nat. Sci. Philad., Addend., 1 861 : 60.

Isuropis dekayi GiU, Rep. U.S. Comm. Fish. ( 1 871-72), 1873: 81 3 (part) ; Baird, Rep. U.S. Comm. Fish.
(1871-1872), 1873: 827 (Woods Hole).

Oxyrhina ptnctata Dumeril, Hist. Nat. Poiss., /, 1865: 409 (descr., N. Amer.).

Isurus functatus Garman, Bull. Mus. comp. Zool. Harv., ly, 1888: 79, pi. I (lateral line syst.) ; Mem. Harv.
Mus. comp. Zool., 56, 1913: 36, pi. 6, fig. 4-6, pi. 56, fig. 5, pi. 58, fig. 3, pi. 62, 63 (descr., anat., east
coast U.S.) ; Nichols and Murphy, Brooklyn Mus. Sci. Bull., 5(1), 1916: 23 (near N. York) ; Fowler,
Copeia, 30, 1916: 36 (mid-Atlant., U.S.); Smith, J. Amer. Mus. nat. Hist., 16, 1916: 342 (Atlant.
coast U.S.) ; Bigelow and Welsh, Bull. U.S. Bur. Fish., 40 (i), 1925: 36, 551 (habits, distrib.. Gulf of
Maine) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 19 (distrib.) ; Burton,
Sci. Mon. N. Y., 40, 1935: 283 (off Charlestown, S. Carolina, but ident. not certain); Anonymous,
Newfoundland Fish. Res. Lab. (1934), 2 (3), 1935: 79 (Grand Banks) ; Springer, Proc. Fla. Acad. Sci.,
5, 1939: 36 (no Florida record given).

Isurus dekayi Smith, Bull. U.S. Fish Comm., 27, 1898: 89 (Woods Hole region, ident. probable because of
local.); Tracy, Rep. R. \. Comm. inl. Fish., 1906: 46 (Narragansett Bay, ident. probable because of
local.) ; Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 7 (New England, ident. probable because
of loc.) ; Tracy, Rep. R. \. Comm. inl. Fish., 1910: 60 (Rhode Island, ident. probable because of loc.) ;
Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., 31 (2), 1913: 736 (Woods Hole region).

Lamna {Oxyrhina) cornubica Imms, Proc. zool. Soc. Lond., (i), 1905: 44 (pharyngeal denticles).

Isurus nasus Garman, Mem. Harv. Mus. comp. Zool., ji5, 1913; 34 (descr., Gt. Brit., Medit., W. Atlant.,
Japan); Fowler, Copeia, 30, 1916: 36 (mid-Atlant. U.S.); Smith, J. Amer. Mus. nat. Hist., 16, igi6:
342 (England) ; Fowler, Proc. biol. Soc. Wash., 55, 1920: 143 (off N. Jersey) ; Huntsman, Contr. Canad.
Biol. (1921), J, 1922: 56 (Passamaquoddy Bay); Bigelow and Welsh, Bull. U.S. Bur. Fish., 40 (l),
'925- 37 (European), 551 {functatus, identical with nasus); Bigelow and Schroedcr, Bull. Mus. comp.

24. The reference on p. 96 is to pi. "24" — actually the plate is No. 14.

Fishes of the Western North Atlantic 123

Zool. Harv., 68, 1927: 239 (discus., mcas., Gulf of Maine) ; Breder, Field Bk. Mar. Fish. Atlant. Coast,
1929: 21 (general) ; Schroeder, Bull. Boston Soc. nat. Hist., 58, 193 1 : 3 (Gulf of Maine) ; Bigelow and
Schroeder, Canad. Atlant. Fauna, biol. Bd. Canad., iz', 1934: 13 (descr., distrib.) ; Vladykov and Mc-
Kenzie, Proc. N. S. Inst. Sci., rp, 193;: 47 (Nova Scotia); Bigelow and Schroeder, Bull. U.S. Bur.
Fish., 48, 1936: 322 (Grand Banks of Newfoundland); Fowler, Bull. Amer. Mus. nat. Hist., 70 (i),
1936: 32 (Italy).

Lamna nasus Jordan, Copeia, 49, 1917: 87 (name); Hubbs, Copeia, 123, 1923: lOl (embryo. Gulf of
Maine); Jordan, Manual Vert. Anim. NE. U.S., 1929: 12 (general); Jordan, Evermann and Clark,
Rep. U.S. Comm. Fish. (1928), 2, 1930: 19 (distrib.) ; Piers, Proc. N. S. Inst. Sci., 18 (3), 1934: 202
(Nova Scotia).

Lamna%^.{\) Gudger, Sci. Mon., N. Y., _?^, 1932: fig. 412, 413 (teeth).

Isurus {Lamna) nasus Tonox\es&, Atti Soc. ital. Sci. nat., y^, 1938: 289 (small size, Medit.).

Doubtful Reference:

Lamna cornubica Lowe, Trans, zool. Soc. Lond., 2 (3), 1837: 194 (Madeira). ^^

Not Lamna functata DeKay, Zool. N. Y., 4, 1842: pi. 63, fig. 206, 207 (these ill. were evidently for /.
oxyrinchtis, see p. 131).

Not Lamna functata Wilder, Science, i, 1880: 236 (Wilder's report of 10 embryos, in this case, makes it
likely that the reference was actually to some genus other than Isurus).

Not Lamia nasus Lahille, An. Mus. nac. B. Aires, s4, 1928: 310 (the specimen in question seems in all prob-
ability to have been Isurus oxyrinchus, see p. I 30).

Not Lamia nasus Vozzi and Bordale, An. Soc. cient. argent., 120, 1935: 150 (this reference is evidently to the
same specimen reported by Lahille, see above ; no doubt /. oxyrinchus) .

Genus Isurus Rafinesque, 1 8 1 o

Isurus Rafinesque, Carratt. Gen. Nuov. Sicil., 1810: II; type species, /. oxyrinchus Rafinesque, Sicily.

Generic Synonyms:

Oxyrhina L. Agassiz, Poiss. Foss.,^' Feuill. Addit., 1835: 71, 86; type species, "Lamna oxyrhina Cuvier anJ

Valenciennes Mss.," equals Isurus oxyrinchus Rafinesque, I 8 10.
Oxyrrhina Bonaparte, Cat. Pesc. Europ., 1846: 17; type species, Oxyrhina gomphodon Miiller and Henle,

I 841 , equals Isurus oxyrinchus Rafinesque, I 8 1 0.
Plectrosoma Gistel, Naturg. Tier., 1848: lO; to replace Oxyrhina L. Agassiz, 1835.
Isuropsis Gill, Ann. Lye. Nat. Hist. N. Y., I, 1862: 397; type species, Oxyrhina glaucus Miiller and Henle,

1841.

Generic Characters. Teeth without lateral denticles; the first 2 in each jaw noticeably
more slender and more flexuous than the others; origin of ist dorsal definitely posterior
to inner corner of pectoral when latter is laid back; trunk slender (Fig. 18) ; caudal fin
without secondary caudal keels. Characters otherwise as in Lamna (p. 1 1 1 ).

Species. This genus includes: the "Sharp-nosed Mackerel Shark" or "Mako" of the
Atlantic (/. oxyrinchus, p. 124) ; the closely allied Pacific Mako (/. glaucus), with which
bideni-'' from South Africa and mako'^ from New Zealand and Australia appear to be

25. Being nominal only, and from a region where oxyrinchus is to be expected, there is no way of knowing to
which species this record actually referred.

26. For dates of publication of the separate parts of the "Poissons Fossiles," see Jeannet (Bull. Soc. neuchatel. Sci.
nat., 52, 1928: 102; 53, 1929; 197).

27. Phillipps, N. Z. J. Sci. Tech., 7j, 1932; 227. 28. Whitley, Rec. Aust. Mus., 77, 1929 : loi.

124 Memoir Sears Foundation for Marine Research

identical 5 also, if the original account be correct, a third Indian Ocean species {guntheri
Murray, 1884) that differs from glaucus in having about twice as many teeth and a much
more prominent lateral line.°°

Key to Species

I a. Teeth about 12-14 on each side on each jaw; lateral line not forming a prominent

ridge along the side.

2a. Height of ist dorsal about one-half as great as distance from eye to 4th gill open-
ing and a little greater than length of its base ; length of head to origin of pectoral
about as great as from axil of pectoral to rear ends of bases of pelvics (or distance
from axil of pectoral to origin of pelvics only about as long as from tip of snout
to 2nd gill opening) . oxyrinchus Rafinesque, 1 8 1 o, p. 124.

2b. Height of ist dorsal only about one-half as great as distance from eye to 2nd gill
opening and a little less than length of its base; length of head to origin of pec-
toral only about as great as distance from axil of pectoral to origin of pelvics (or
distance from axil of pectoral to origin of pelvics about as long as from tip of
snout to origin of pectorals). glaucus Miiller and Henle, 1841.'°

Indo-Pacific.
lb. Teeth 22-28 on a side in each jaw; lateral line forming a prominent ridge along the
side, rearward to caudal peduncle. guntheri Murray, 1884.

India."

Isurus oxyrinchus Rafinesque, 1810

Sharp-nosed Mackerel Shark, Mako

Figures 18, 19

Study Material. Skin, with head attached, of adult male, 7 feet 6 inches long, from
Bimini, Bahamas (Harv. Mus. Comp. Zool., No. 35367); young male (entire), 1,640
mm., from Ocean City, Maryland (Harv. Mus. Comp. Zool., No. 35899); skin, about
6 feet long, from Miami, Florida (Harv. Mus. Comp. Zool., No. ZSZ^^)'-! ^^so female
(mounted) from Miami, Florida, about 6 feet 8 inches long; jaws from medium-sized
and large specimens taken off New Jersey, New York and Cape Cod (U.S. Nat. Mus., No.
iio88i, Amer. Mus. Nat. Hist., No. 567 and 9220, Harv. Mus. Comp. Zool., No. 816).

Distinctive Characters. The Mako is separable from the common Mackerel Shark
by its teeth and more slender form; from Carcharodon by its teeth, its slender form and by
the relative position of the second dorsal and anal fins.

29. Only one specimen of this sort has ever been reported (Murray, Ann. Mag. nat. Hist., [5] 13, 1884: 349).

30. Including bideni PhiUipps (N. Z. J. Sci. Tech., 13, 1932: 227, fig. 2, S. Africa); mako Whitley (Rec. Aust.
Mus., ly, 1929: loi).

31. Only one specimen of this sort has ever been reported (Murray, Ann. Mag. nat. Hist., [5] ij, 1884: 349) ; the
validity of this species is very doubtful.

Fishes of the Western North Atlantic

125

Description. Proportional dimensions in per cent of total length. Male, 1,598 mm.,
from Ocean City, Maryland (Harv. Mus. Comp. Zool., No. 35899). Male, 2,337 mm.,
from Bahamas (Harv. Mus. Comp. Zool., No. 35367).

Trunk at origin of -pectoral: breadth 10.3, 12.4; height 1 1.3, 1 1.9.

Snout length in front of : o\ite.vnos\.v\\?, A,.l, j mouth 6.1,6.5.

Eye: horizontal diameter 1.8, 1.8.

Mouth: breadth 6.2, 6.1 ; height 5.0, S-^-

Nostrils: distance between inner ends 3.4, 3.8.

Labial furrow length: upper I . i , ; lower 0.6, .

Gill opening lengths: ist 6.9, 6.9; 2nd (}.()^ 5 3rd 6.4, -; 4th 6.3, 5

5th 6.8, 7.2.

F;W/<iorjfl/j^«; vertical height 9.2, lO.Oj length of base 8.8,9.1.

Second dorsal fin: vertical height 1.6, 2.1 ; length of base 1.6, i.i.

Anal fin: vertical height 1.9, 2.1 j length of base 1.5, 1.4.

Caudal fin: upper margin 20.6, 21.8; lower anterior margin 15.5, 16.7.

Pectoral fin: outer margin 17. i, ; inner margin 4.8, j distal margin

14.0, .

Distance from snout to: ist dorsal 36.6, 34.8; 2nd dorsal 69. o, 68.5; upper caudal

79.4, 78.2; pectoral 26.1, 25.05 pelvics 53.2, 50.O} anal 70.5, 68.5.

Figure 18. Imrus oxyrinchus, young male, 1,640 mm. long, taken off Ocean City, Maryland (Harv. Mus.
Comp. Zool., No. 35899). A Left nostril, about l.I x. 5 Caudal peduncle viewed from above, about Y^
natural size. C Cross section of caudal peduncle at point indicated by the transverse line on B. D Pelvic fins
and claspers. E Second dorsal and anal. F Dermal denticles, about 32 x. G Apical view of dermal denticle,
about 65 X.

126 Memoir Sears Foundation for Marine Research

Interspace between: ist and 2nd dorsals 24.8, 25.5; 2nd dorsal and caudal 8.7,

8.7} anal and caudal 8.0, 7.2.

Distance from origin to origin of: pectoral and pelvics 27.9, 28.3 ; pelvics and anal

16.9,17.2.

Figure 19. Isurus oxyrinchus. A Upper and lower teeth of specimen pictured in Fig. l8, about natural size.
B Side view of anterior part of jaws of a large Cape Cod, Massachusetts, specimen (Harv. Mus. Comp. Zool.,
No. 816), about J/2 X.

Trunk fusiform, considerably more slender than in Lamna nasus, its height at origin
of 1st dorsal (where highest) about equal to distance from eye to 2nd gill opening, or about
15% of total length, tapering both rearward and forward. Caudal peduncle very much
flattened dorso-ventrally, but broadly expanded laterally and sharp-edged as in other
Isuridae (Fig. 1 8 B), but without the secondary keel below it that is characteristic of nasus.
Sides smooth. Lateral line not prominent. Upper and lower caudal pits strongly developed
as deep furrows, nearly transverse to peduncle, or perhaps slightly arcuate (convexity
rearward) in some specimens. Denticles small, closely imbricate, with 3 to 5 ridges, and 3
marginal teeth, the median the longest but often worn down.

Fishes of the Western North Atlantic 127

Head conical but somewhat flattened dorsally. Snout pointed. Eyes round, their
diameters about Vs ^s long as snout in front of mouth. Nostrils nearly transverse, about
Vs as long as distance between them, their inner margins without definite lobe, the dis-
tance from inner corner of nostril to mouth between 33 and 50% as great as to tip of
snout. Spiracle a minute pore or slit, about at same level as upper margin of eye, and
situated behind the latter by a distance equal to about 3 times the eye's diameter. Gill open-
ings noticeably large, the i st to 4th about as long as snout in front of mouth, the 5th slightly
longest, the ist nearly straight, but lower outlines of others increasingly flexuous, that of
the 5th most strongly so, the 5th close in front of origin of pectoral and extending ven-
trally around the latter for a distance about ^o as long as diameter of eye, the distance
between 4th and 5th only about Y^, as great as between ist and 2nd. Mouth very broadly
rounded in front and notably long, about 1.15 times as broad as long. Upper labial fur-
row about 25% as long as distance (around the curve) from corner of mouth to symphy-
sis of upper jaw, ending about opposite 7th tooth, the lower furrow slightly more than Y2
as long as upper and entirely concealed when mouth is closed, ending opposite 6th tooth.

Teeth '^ °^ '^ — " ""^ '^ : alike in the 2 iaws; slender, somewhat flexuous in outline,

12 or 13 — 12 or 13-' J' ' '

smooth-edged, without lateral denticles} the ist 2 in each jaw much the largest, the ist
2 in each jaw recurved at base, but with curve reversed at tips, their outer faces flat but
inner faces rounded} subsequent teeth relatively broader and increasingly blade-like, their
outer margins varying from very strongly convex to very slightly concave, their inner
margins slightly more concave} 3rd upper tooth much smaller than 2nd or 4th to 7th, but
3rd lower tooth about as large as 4th to 6th} 9th to 13th teeth in each jaw successively
smaller and with cusps shorter relative to breadth of base, the 2 or 3 outermost minute }
I or 2 rows functional along sides, but 2 or 3 in front of jaws.

Origin of ist dorsal about over inner corner of pectoral when latter is laid back, or
perhaps slightly behind it in some specimens (thus relatively farther back than in nasus),
the midpoint of its base slightly nearer to anterior margin of eye than to origin of caudal,
its length at base about equal to V2 the distance from posterior margin of eye to 5th gill
opening or to about V2 the length of pectoral, its anterior margin slightly convex, its apex
moderately rounded, its rear margin rather strongly concave, its free rear corner broadly
triangular and about V4 as long as base. Origin of 2nd dorsal slightly but unmistakably
anterior to origin of anal, the rear end of its base about over midpoint of base of latter,
its length at base only about Yj to Vs as great as that of ist dorsal, its apex rounded, its
posterior margin deeply concave, its free rear corner about i ^,4 times as long as its base.
Lower lobe of caudal slightly more than 75% as long as upper (77 to 80% in 2 specimens
studied), each measured from its respective precaudal pit, the upper lobe about as long as
from front of mouth to origin of pectoral, or slightly longer than latter, the upper anterior
and lower anterior outlines of caudal only slightly convex, the tips subacute, posterior con-
tour deeply and nearly evenly concave, with well marked subterminal notch. Anal similar
to 2nd dorsal, but with relatively longer free rear corner (about twice as long as base), and

128 Memoir Sears Foundation for Marine Research

of about the same size. Pelvics originate posterior to rear tip of ist dorsal by a distance
about equal to distance from tip of snout to mouth, or slightly greater, their corners
rounded, their outer margins concave. Claspers of male long and slender, reaching about
% of the distance to origin of anal. Pectoral about as long as distance from posterior margin
of eye to 5th gill opening, or about twice as long as vertical height of i st dorsal, a little less
than V2> or about 45%, as broad as long, the outer margin slightly convex, the tip and
inner corner rounded, the rear margin moderately concave.

Color. Described as deep blue-gray above when fresh caught,"" but appearing cobalt
or ultramarine blue in the water j snow-white below; dark slate gray above after preser-
vation, and bluish white to pale dirty gray below, on head and body, and on lower surface
of pectoral, with gradual transition from one shade to the other along the middle of the
trunk.

Size. While the Mako is said to reach a length of 13 feet (4m.), the maximum length
reported for an actual specimen of this species is only about 1 2 feet.'"

The largest West Atlantic specimen of which we find definite record, taken ofiF St.
Petersburg, Florida, was about 10 feet 6 inches long; one nearly as large (10 feet, 2
inches) was taken off New York Harbor many years ago. Males are sexually mature, as
indicated by the claspers, at perhaps six feet, females perhaps not until somewhat larger.
Recorded weights at different lengths are about 135 pounds at 6 feet; 230 pounds at 7 feet
8 inches; about 300 pounds at 8 feet; 1,009 pounds at lO feet 6 inches. A weight of 700-
800 pounds may be expected at about 9 feet, depending on condition. The largest speci-
men so far caught on rod and reel was one of 786 pounds taken off Bimini, Bahamas, by
Ernest Hemingway in 1936. The largest Pacific Mako (glaucus) yet taken on rod and
reel, by E. White-Wickham off New Zealand, weighed 798 pounds.

Developmental Stages. Embryos, like those of other members of the family (p. 116),
are provided with a voluminous yolk stomach, and before birth they reach a very large size
relative to that of the mother. Presumably the number of young in a brood is correspond-
ingly small, but no definite information is at hand."*

Habits. This is one of the most active and strongest swimming of sharks, famous for
its habit of leaping clear of the water under natural conditions and when hooked. It ap-
pears to be typically a near-surface fish, often seen swimming on sunny days with the tips
of first dorsal and caudal fins above the water. Around the Canary Islands it is often
hooked at depths of from five to eight meters, but we have no definite information as to
how deep it may descend. Nothing is definitely known of its breeding habits, but presum-
ably it is similar in these to its more familiar relative, nasus (p. 117).

Very little is known of its diet other than that it is a fish-eater, preying upon the

32. Shown as dark slaty blue above and grayish white below in colored sketch of a fresh 8-foot 4-inch specimen,
by J. Henry Blake, Provincetown, Mass., October 1868.

33. 3.7 mm. calculated from the size of the jaws (Uriarte and Mateu, Notas Inst. esp. Oceanogr., 55, 1931 : 12) j
specimen from the Canaries.

34. Vaillant (Bull. Soc. philom. Paris, [8] x, 1889: 38) reports an embryo of this species from the Mediterranean,
50 cm. long, including caudal, with yolk stomach 23 to 24 cm. long; size of the mother is not known.

Fishes of the Western North Atlantic 129

schools of scombroids, clupeids, or other small fishes, of which it destroys great quan-
tities. Around Bermuda, for example, it is seen most often when in pursuit of scombroids;
off the coast of the United States it has at least the reputation of following schools of mack-
erel. It also feeds on much larger fish. A 120-pound swordfish (Xiphias gladius) nearly
intact with sword still attached was found in the stomach of a 730-pound specimen taken
near Bimini. Another Mako of about 800 pounds, harpooned off Montauk, Long Island,
had been seen attacking a swordfish and was found, when landed, to contain about 150
pounds of its flesh. These instances illustrate its capabilities, and one well known angler
described it as the only marine enemy of the broadbill swordfish.'^ But there is no reason to
suppose from the nature of its teeth, or from repute, that it attacks sea turtles. Probably,
like most other pelagic sharks, it feeds also on squids when opportunity offers itself.

Relation to Man. The flesh is sold in limited quantities, but the chief importance of
the species is as a game fish because of its famous habit of leaping when hooked, as men-
tioned above. In this respect, as well as in the fierceness of its resistance to capture, it falls
little or not at all behind its better known relative, glaucus, of New Zealand waters.

Range. An oceanic species of the tropical and warm-temperate Atlantic, north and
south; it is replaced in the Pacific (including New Zealand and Australian waters) by the
closely allied but easily distinguishable I. glaucus (p. 124). The fact that the ranges of
glaucus and oxyrinchus appear to be continuous around the Cape of Good Hope, although
widely separated off the southern part of South America, lends special interest to the iden-
tity of any specimens that may be caught off the Cape.

Occurrence in the Eastern Atlantic. In the eastern side of the Atlantic oxyrinchus is
known as far south as St. Helena and Ascension and northward to northern France; to
northern Scotland and southwestern Norway as a stray. Coastwise, however, its zone of
reasonably frequent occurrence appears to extend only from tropical West Africa to the
Iberian Peninsula, including the Mediterranean, whence it has been recorded repeatedly as
"common" or "abundant" from many localities. It is also known from the Azores, Madeira
and the Canaries, where it is said to be one of the commoner sharks, occasionally numerous
enough to be a great annoyance to net fishermen. This, together with its long known
presence in at least small numbers around Bermuda, shows that it is to be expected any-
where in the middle Atlantic.

Occurrence in the Western Atlantic. For the western Atlantic only a very fragmentary
picture of the occurrence of this offshore shark could be deri\-ed from the captures reliably
reported in scientific literature, since these^^ total not over 20 to 25 distributed as follows:
off the tip of Cape Cod; 10 miles N.E. of Nantucket Lightship, Mass.; Long Island, New
York; vicinity of New York Harbor; coast of New Jersey; off Cape Hatteras, North
Carolina; western and northwestern Florida; east coast of Florida; Santa Rosa Island near

35. See K. Farrington (Field and Stream, ^7, February 1943) for the instances mentioned above and for other
interesting notes on the Mako.

36. It seems certain that at least most of the nominal records for this species in the Gulf of Maine and for the vicinity
of Woods Hole actually referred to L. nasus (p. 1 30) .

I'lO

Memoir Sears Foundation for Marine Research

Pensacola, Florida; Cuba; Gulf of Mexico; Havana, Cuba; Rio de Janeiro, Brazil; Ber-
muda. A shark taken in 1927 off Mar del Plata, in northern Argentine waters," probably
belonged to this species also.

Fortunately, however, there is now available a much more extensive source of infor-
mation in the published and verbal reports of anglers, since oxyrinchus is a favorite game
fish. From these it is well known to be tolerably plentiful in the winter on the Bahaman
side of the Straits of Florida, where many are caught off Bimini, Cat Key and Nassau,
but it is less frequent on the Florida side, although it is a familiar fish there. Also, in the
summer considerable numbers journey northward along the continental shelf as far as the
offings of Maryland, New Jersey, New York and southern New England, although they
rarely, if ever, come close enough inshore to be picked up in the pound nets. Perhaps they
never penetrate far into inlets. However, to keep offshore is not an invariable part of its
behavior pattern, for on the tropical coast of West Africa it has been reported from sundry
estuarine situations. During the past few summers we have heard repeatedly of "Makos"
seen jumping, or occasionally hooked, near the tip of Cape Cod. Recently a large one
(about 9 feet long) was caught on the southern side of Massachusetts Bay a few miles
off Plymouth.'* It is thus evident that at least scattered individuals enter the south-
western part of the Gulf of Maine, probably in pursuit of the schools of mackerel, but
it appears that this is its extreme northerly outpost in inshore waters on this side of the
Atlantic. The sundry early reports that ostensibly referred to it farther north in the Gulf
of Maine all appear to have been based on its close relative, nasus (p. 118). Apparently
it rarely if ever occurs in water colder than about 60°. But it would not be astonishing if it
were encountered farther north, offshore, in the sweep of the Gulf Stream, although there
is as yet no positive record of it either from the Nova Scotian Banks or from the Banks of
Newfoundland.

Except for its presence in Bahaxnan waters, knowledge of it in the southern part of
its western Atlantic range is very scant, but the records for western and northwestern
Florida and Cuba,'" together with evidence from recently-received photographs of one
from southern Texas (Cameron County), is evidence that it ranges over the Gulf of
Mexico generally, and in all probability over the entire Caribbean region. But information
as to its occurrence off the South American seaboard is limited to the one positive record for
Rio de Janeiro, and one probable record for northern Argentina."

Synonyms and References:

Isurus ox-jrinchu! Rafinesque, Caratt. Gen. Nuov. Sicil., 1 8 10: 12, pi. 13, fig. I (type loc, Sicily); Fowler,
Bull. Amer. Mus. nat. Hist., yo (l), 1936: 33 (West Africa, descr.) ; Tortonese, Atti Soc. ital. Sci. nat.,

37. Pictured and described by Lahille (An. Mus. nac. B. Aires, ^4, 1929: 310) as "Lamia nasus." But his illustration
(p. 311, fig. 10) shows the origin of the first dorsal as being over the inner corner of the pectoral, the second
dorsal as slightly in advance of the anal, and the lower caudal lobe as only slightly shorter than the upper, as in
oxyrlnclius, while his statement that the teeth have lateral denticles appears to refer to the species nana as a whole,
rather than to the particular specimen.

38. Personal communication from W. J. Mixter in the late summer of 1941.

39. Luis Howell-Rivero writes us that it is always taken offshore there.

Fishes of the Western North Atlantic 131

77t '938: 290 (Medit.) ; Springer, Proc. Fla. Acad. Sci., 5, 1939: 35 (Florida) ; Bigelow and Schroeder,

Guide Coram. Shark Fish., Anglo Amer. Carib. Comm., Wash., 1945: 102, fig. 35 (descr., ill., habits,

range).
Isurus sfaUctnzanii Raiinesque, Indice Ittiol. Sicil., 1810: 45, 60 (Sicily) ; Jord.in and Gilbert, Bull. U.S. nat.

Mas., 16, 1883: 874 (name, descr., C. Cod, West Indies) ; Nobre, Fauna Marinha Port. Vert., /, 1935:

431, pi. 58, fig. 184 (descr., Portugal).
Lamia oxyrhincus Bory de St. Vincent, Diet. Class. Hist. Nat., 75, 1829: 596 (ref.).
Squalus cefedii Lesson, Voy. "Coquille," Zool., 2, 1830: 93 (equatorial Atlantic).
Lamna oxjrhina L. Agassiz, Poiss. Foss., 5, 1838: 86, ref. to pi. A, Lamna (teeth) ; Owen, Odontogr., 1840-

1845: 28, pi. 3, fig. I (teeth).
Isurus oxyrynchus Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1839: 313 (general).
Oxyrhina sfallanzanii Bonaparte, Icon. faun. Ital., j (2), I 839: pi. [136], fig. I ; Agassiz, L., Poiss. Foss., j,

1839: 276, ref. to pi. G, fig. 2, Lamna; Nardo, Atti 1st. veneto, (3) 5, 1859-1860: 787 (Medit.);

Dumeril, Hist. Nat. Poiss., /, 1865: 408, pi. 7, fig. 4 (descr., teeth); Miklucho-Maclay, Beitr. Vergl.

Neurol. Wirbelt., i, 1870: 26, pi. 5, fig. 3A, B (brain); Ninni, Ann. Soc. nat. Modena, 5, 1870: 66

(Medit.); Canestrini, in Cornalia, et al.. Fauna d'ltal., 1870-1872: 45 (Medit.); Poey, An. Soc. esp.

Hist, nat., 5, 1 876: 391, pi. 14, fig. I; Enumerat. Pise. Cubens., 1876: 185, pi. 9, fig. i (spec. 2,585

mm., ill., tooth, descr., discus., Cuba); Gervais and Boulart, Poiss., 5, 1877: 182, pi. 69 (descr.);

Stossich, Boll. Soc. adriat. Sci. Nat., 5, 1880: 68 (Adriatic) ; Moreau, Hist. Nat. Poiss. France, 7, 1881:

298 (descr., France) ; Doderlein, Man. Ittiol. Medit., 2, 1881 : 62 (Medit.) ; Vaillant, Bull. Soc. philom.

Paris, (8) 1, 1889: 38 (embryo); Huber, Z. Wiss. Zool., 70, 1901: 619 (claspers) ; Sicher, Atti Accad.

gioenia, (4) 77 (5), 1898; 16 (Medit.); Belloc, Rev. des. Trav. Peches Marit., 7 Fasc. 2, 1934: 137

(ill. after Bonaparte; Morocco, Senegal).
Oxyrrhina glauca Bonaparte, Mem. Soc. neuchatel. Sci. nat., 2 (8), 1839: 9 (in synopsis) ; Heckel, Peix. Dal-

maz. in Carrara, 1864:91 (Medit., not seen) ; not OA'yr^/wij ^/i3i«;3 Miiller and Henle, 1841.
0;«-yrA;»<»^OOT/i/^od^o«MullerandHenle, Plagiost., 1841:68, I9l,pl. 28 (descr.) ; Gray, List Fish. Brit. Mus.,

7, 1 851: 60; Bocage and Brito Capello, Poiss. Plagiost., 1866: 13, pi. 3, fig. 3 (Portugal) ; Brito Capello,

J. Sci. math. phys. nat. Lisboa, 2, 1869: 139; Hasse, Naturl. Syst. Elasm. besond. Theil, 1882: 230, pi. 31,

fig. 36-38, 41 (skelet.) ; Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 8; Faune Senegamb. Poiss.,

/, 1 883-1 885: 22 (Senegambia).
Lamna pinctata'DtKiy, Zool. N. Y., 4, 1842: 352, pi. 63, fig. 206 (descr., good ill., teeth, size, N. York) ; not

Lamna -punctata Stor&r, 1 839.
Lamna cornubicus Cuvier, Regne Anim. Poiss., ill. ed., 1843: pi. 114, fig. 3 (teeth).

Lamna punctata {^ ) Goode, Bull. U.S. nat. Mus., 5, iSj6: 7-} ; not La?;ma punctata Stoier, 1839 (Bermuda).
Oxyrrhina spalanzanii Bonaparte, Cat. Pesc. Europ., 1846: 17 (Medit.).
Isuropsis dekayi Gill, Ann. N. York Lye, 7, 1 862: 409; Poey, Repert. Fisico-Nat. Cuba, 2, 1 868: 446 (Cuba) ;

Yarrow, Proc. Acad. nat. Sci. Philad., 29, 1877: 217 (Ft. Macon, North Carolina). Not Oxyrhina

daekayi Gill, Proc. Acad. nat. Sci. Philad., Addend., 1 861: 60.
Lamna latro Owen, Cat. Osteol. Roy. Coll. Surg., 7, 1 853 : 96 (teeth, ident. by ref. to Owen, Odontogr., 1 840-

i845:pl. 5, fig. i).
Carcharias tigris Atwood, Proc. Boston Soc. nat. Hist., 70, 1865: 81; 72, 1869: 268 (descr., G. Mexico,

C. Cod).
Lamna spallanzanii Gunther, Cat. Fish. Brit. Mus., 8, I 870: 380 (descr., distrib., synon.) ; Reguis, Ess. Hist.

Nat. Provence, 7 (l), 1877: 49 (Medit.); Perugia, Elenc. Pesc. Adriat., 1.881: 53 (Adriatic); Graeffe,

Arb. zool. Inst. Univ. Wien, 1886: 446 (Medit.); Helgendorf, Arch. Naturgesh., S4 (0> '888: 213

(Azores); Carus, Prod. Fauna Medit., 2, 1 889-1 893: 505 (Medit.); Steindachner, S. B. Akad. Wiss.

Wien, 700 (l), 1891: 363 (Canaries); Taylor, Ann. Scot. nat. Hist., 1910: 250 (off Scot.); Gunther,

Encycl. Brit., nth ed., 24, 1911: 807 (general); Metzelaar, Trop. Atlant. Visschen, 1919: 189.
Lamna glauca Gunther, Cat. Fish. Brit. Mus., 8, 1870: 391 ; not Oxyrhina glauca Miiller and Henle, 1841.
Isuropsis %^. Goode, Amer. J. Sci., (3) 14, 1877: 293 (Bermuda).
Isurus glaucus Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 28 (Atlant. Oc, Cuba) ; Barnard, Ann. S.

Afr. Mus., 27 (l), 1925: 33 (part, S. Afr.).

132 Memoir Sears Foundation for Marine Research

Isurus tlekayi Jordan and Gilbert, Proc. U.S. nat. Mus., 5, l88z: 241 (descr., Santa Rosa 1., near Pensacola,
Florida, color); Goode and Bean, Proc. U.S. nat. Mus., 5, 1882: 240 (Gulf of Mexico); Jordan and
Gilbert, Bull. U.S. nat. Mus., 16, 1883: 874 (descr., C. Cod to W. Indies); Jordan, Rep. U.S. Comm.
Fish. (1885), 1887: 797 (distrib.); Nelson, Rep. N. J. Geol. Surv., 2 (2), 1890: 663 (N. Jcr=ey
record, feeding habits) ; Bean, Rep. For. Comm. N. Y., 1901 : 379 (near N. York, but perhaps confused
with L. nasus) ; Kendall, Occ. Pap. Boston Soc. nat. Hist., 7(8), I 908: 7 (part) ; Jordan and Evermann,
Bull. U.S. nat. Mus., 47 (4), 1900: pi. 6, fig. 21; Evermann and Kendall, Rep. U.S. Comm. Fish.
(1899), 1900: 49 (Florida) ; Gregg, Where to Catch Fish, 1902: 17 (Florida) ; Bean, Bull. N. Y. St.
Mus., 60, Zool. p, 1905: 38 (off N. York) ; Fowler, Rep. N. J. Mus. (1905), 1906: 54, pi. I (N. York) ;
Tracy, Rep. R. 1. Comm. inl. Fish., 1906: 46; Rep. R. I. Comm. inl. Fish., 1910: 60 (Rhode Island,
same as foregoing); Nichols, Abstr. Linn. Soc. N. Y., 20-23, 1913: 91 (off N. York); Nichols and
Murphy, Brooklyn Mus. Sci. Bull., 3 (l), 1916: 22 (descr., W. Indies-C. Cod).

Isurus {Isurus) oxyrhynchus ]oidzn and Evermann, Rep. U.S. Comm. Fish. (189;), 1896: 218 (distrib.);
Bull. U.S. nat. Mus., 47 (l), 1896: 48 (descr., distrib.).

Isurus (Isurofsis) liekayi J or d3.n and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 218 (distrib.); Bull.
U.S. nat. Mus., 47 (l), 1896: 48 (descr., C. Cod, W. Indies).

Isurus oxyrhynchus Schreiner and Ribeiro, Arch. Mus. nac. Rio de J., 12, 1903; 79 (Brazil); de Braganza,
Result. Invest. Sci. "Amelia," 2, 1904: 52 (Portugal) ; Collett, Norges Fisk., SuppL, j, 1905: 76 (Bergen,
Norway); Fowler, Proc. Acad. nat. Sci. Philad., 60, 1908: 55 (jaws); Garman, Mem. Harv. Mus.
comp. Zool., 3(5, 1913: 37 (descr.); Jordan, Copeia, 49, 1917: 87 (name); Ribeiro, Arch. Mus. nic.
Rio de J., Fauna Brasil., Peixes, 2 (l). Ease. I, 1923: 18, pi. 6 (Rio de Janeiro) ; Rey, Fauna Iberica
Feces, i, 1928: 376 (descr., habits, Spain) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928),
2, 1930: 19 (distrib.); Uriarte and Mateu, Notas Inst. esp. Oceanogr., (2) 55, 1 931: 12 (Canaries);
Bigelow and Schroeder, Canad. Atlant. Fauna, biol. Bd. Canada, 72^, 1934: 15 (general); Lubbert and
Ehrenbaum, Handb. Seefisch. Nordeurope, 2, 1936: 279 (Bergen, Norway; Medit.).

Isurus (probably iekayi) Bean, Field Mus. Publ. Zool., 7 (2), 1906: 30 (Bermuda).

Isurus ligris Garman, Mem. Harv. Mus. comp. Zool., ^(5, 1913: 36 (descr.); Fowler, Copeia, 30, 1916: 36
(off U.S.) ; Radcliffe, Bull. U.S. Bur. Fish., 5.;, 1916: 247 (teeth) ; Fowler, Proc. Acad. nat. Sci. Philad.,
69, 1917: 109 (N. Jersey); Jordan, Copeia, 49, 1917: 87 (name); Murphy, Copeia, 69, 1919: 32
(descr., photo., N. York) ; Bigelow and Welsh, Bull. U.S. Bur. Fish., 40 (l), 1925: 38 (descr.. Gulf of
Maine);*" Breder, Copeia, 153, 1926: 121 (Sandy Hook Bay, New York); Nichols and Breder, Zoo-
logica, N. Y., p, 1927: 19 (Long Island, N. York) ; Bigelow and Schroeder, BulL Mus. comp. Zool. Harv.,
68, 1927: 240 (discuss.) ; Jordan, Manual Vert. Anim. NE. U.S., 1929: i 2 (general) ; Breder, Field Bk.
Mar. Fish. Atlant. Coast, 1929: 22 (general) ;*^ Fowler, Proc. Acad. nat. Sci. Philad., So, I 929: 607, pi.
31 ; (N. Jersey) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 19 (distrib.) ;
Firth, Bull. Boston Soc. nat. Hist., 61, 1 931: 8 (off Nantucket) ; Pearson, Invest. Rep., U.S. Bur. Fish.,
(10) J, 1932: 18 (North Carolina); Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda, 1933: 23
(Bermuda) ;*- Bigelow and Schroeder, Canad. Atlant. Fauna, biol. Bd. Canad., 72^ 1934: 14 (general) ;
Bull. U.S. Bur. Fish., 48, 1936: 322 (near Nantucket); Fowler, Proc. Acad. nat. Sci. Philad., Sg, 1937:
303 (N. Jersey) ; Breder, Bull. N. Y. zool. Soc, 41, 1938: 28 (near N. York).

Isurus cepedii (in part) Fowler, Bull. Amer. Mus. nat. Hist., 70 (2), 1936: 34 (St. Helena, Ascension, but
glaucus also included).

Lamna oxyrhytichus Borri, Mem. Soc. tosc. Sci. nat., 44, 1934: 92 (Medit.); Norman and Eraser, Giant
Fishes, 1937: 12 (general).

Mako Shark {Lamna), Kaplan, Big Game Fisherman's Paradise, Dep. Agric. Fla., 1936: 104 (Florida).

Lamna tigris Norman and Eraser, Giant Fishes, 1937: 12 (general).

Isurus ox^rinchus Fowler, Arqu. Zool. Estado Sao Paulo, j, 1942: 127 (Brazil).

Isurus cefedii Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1945: 43, fig. 3, 4 (Bimini, Bahamas, and Worcester
Co., Maryland).

40. At least some of the Gulf of Maine records listed there probably referred to nasus, p. 118.

41. The illustrations of oxyrinc/ius {"tigris") and nasus are transposed.

42. Their illustration (p. 24) actually represents nasus.

Fishes of the Western North Atlantic 133

Doubtful References:

Squalus cefedii Lesson, Voy. "Coquille," ZooL, 2, 1 830: 93.^'

hurus dekayi Linton, BulL U.S. Bur. Fish., ig, 1901: 429 (Woods Hole); Sharp and Fowler, Proc. Acad.

nat. Sci. Philad., 56, 1 904: 506 (Nantucket).
Lamia nasus Lahille, An. Mus. nac. B. Aires, ^4, 1928: 310 (Mar de la Plata, Argentina).
hurus oxyrAyncAus Wilson, Proc. U.S. nat. Mus., 60 (5), 1932: 24 (parasites).
Isurus tigris WUson, Bull. U.S. nat. Mus., 158, 1932: 427, 446 (par.isitcs).
Not Isurus dekayi Smith, Bull. U.S. Bur. Fish., //, 1898: 89;" Kendall, Occ. Pap. Boston Soc. nat. Hist., 7

(8), 1908: 7;*° Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., 3/ (2), 1913: 736.*'

Genus Carcharodon Agassiz, 1838

Carcharodon Agassiz, L., Poiss. Foss., j, 1 838: 91 ; type species, verus Agassiz, L., equals Carcharias verus
Cloquet (Diet. Sci. Nat., 7, 1817: 69) and Squalus carcharias Linnaeus, 1758.^

Generic Synonyms:

Squalus (in part) Linnaeus, Syst. Nat., 1758: 233 and subsequent authors.^

Carcharias (in part) Cloquet, Diet. Sci. Nat., 7, 1817: 69; Cuvier, Regne Anim., 2, 1817: 125, and

subsequent authors; not Carcharias Rafinesque, 18 10 (see p. 98).
Carcharocles Jordan, Stanford Univ. Publ. Biol., j, 1923: 99; type species, Carcharias auriculatus Blainville,

fossil.
Carcharhinus Whitley, Mem. Qd. Mus., 10, 1934: 199; not Carcharhinus Blainville, 1816 (see p. 320).

Generic Characters. Teeth triangular, with slightly concave margins and coarsely ser-
rate edges, but without lateral denticles j lower teeth smaller and more slender than
uppers} 3rd upper tooth nearly as large as 2nd and 4th; snout conical, flattened above,
only moderately acute j anterior part of caudal without secondary longitudinal keel below
rearward extension of expanded caudal peduncle. Characters otherwise those of the family.

Range. Pelagic; cosmopolitan in tropical, subtropical and warm temperate seas, in-
cluding the Mediterranean.

Fossil Teeth. From Upper Cretaceous to Pleistocene, Europe; Eocene to Pliocene,

43. Tortonese (Atti Soc. ital. Sci. nat., 77, 1938 : 291) revives this name to replace glaucus Miiller and Henle for the
Indo-Pacific form. But Lesson (1830: 93) expressly states that his specimen was harpooned "dans la ocean
Atlantique," in Lat. 6° S., though the longitude as given, 27° E., is patently in error, which accords with the
general location of the ship on the stated date of capture, Sept. 28, 1822.

44. The account of occurrence near Woods Hole makes it highly probable that these citations actually referred to
tuisus.

45. From the authorities cited, from the widespread distribution and from the abundance credited to it in the Gulf
of Maine, this evidently referred to nasus.

1. The early history of the generic name Carcharodon is confused. Proposed in 1838 by Miiller and Henle (Charles-
worth Mag. nat. Hist., [2] 2: 37) with diagnosis but without mention of any particular species, its type species
was designated in tiie :anie ye.ir by L. Agassiz (Poiss. Foss., 5; 91) as "C. smithii Miiller and Henle." However,
since this is a nomen nudem, not used by Miiller and Henle, the genus must be credited to L. Agassiz, its type being
Carcharodon verus Agassiz (Poiss. Foss., 3, 1838: 91), the account of which, added to his illustration of its teeth
printed three years earlier simply as "Carcharias" (Poiss. Foss., 5, 1835: pi. F, fig. 3), leaves no doubt as to its
identity. We may point out that the specific name verus (equivalent to Squalus carcharias Linnaeus, 1758) actually
dates from Cloquet, 1 8 1 7, for as used earlier by Blainville (Bull. Soc. philom. Paris, 1 8 1 6 : 121) it was a no»:en
nudem also, since it lacked any indication as to identity.

2. See under References, Carcharodon carcharias, p. 142.

134 Memoir Sears Foundation for Marine Research

Africa; Eocene to Pleistocene, North America; Miocene, South America, West Indies;
Miocene to Pliocene, West Indies, New Zealand; Pliocene, eastern Asia.

Species. It is probable that all published accounts of this genus, whether Pacific or
Atlantic, belong to a single species.' Since final conclusion must await critical comparison
of specimens from the two oceans, or at least comparable measurements, the Pacific refer-
ences are segregated below (p. 144) from those for the Atlantic and Mediterranean.

Carcharodon carcharias (Linnaeus), 1758

White Shark, Man-eater

Figures 20, 21, 22

Study Material. Jaws from specimens of about 6 feet from Long Island, New York
(Amer. Mus. Nat. Hist., No. 14773), 8^2 feet from Woods Hole, Mass. (U.S. Nat.
Mus., No. 10899) ^"d of about 12 feet from an unknown locality (Harv. Mus. Comp.
Zool.) ; a mounted specimen about 6 feet long from Woods Hole (in New England Mus.
Nat. Hist.); two fresh caught specimens, about 9 and 10 feet long, from Massachusetts
Bay, but not preserved; good photographs of several fresh specimens, of about 5 to 10
feet long, taken off the tip of Cape Cod, off Rhode Island and off Sarasota, Florida.*

Distinctive Characters. The combination of strongly lunate caudal with very large
triangular and coarsely serrate teeth is diagnostic. The more rearward position of the anal
relative to the second dorsal and the blunter snout further separate it from its relatives of
the genera Isurus and Lamna.

Description. Proportional dimensions in per cent of total length. Female, immature,
4,700 mm. total length,^ from Florida.

Snout length in front of: mouth 6.3.

Mouth: height i.i.

Nostrils: distance between inner ends 3.8.

Gill opening lengths: ist 9.0; 5th 9.7.

First dorsal fin: anterior margin 12.8; length of base 9.7.

Second dorsal fin: anterior margin 2.8; length of base 1.4.

Anal fin: anterior margin 2.6; length of base 1.4.

Caudal fin: upper margin 20.0; lower margin 13.5.

Pectoral fin: outer margin 18.9; inner margin 4.6; distal margin 16.6.

Distance from snout to : ist dorsal 37.5; upper caudal 80.O; pectoral 27.7.

Interspace between: ist and 2nd dorsals 21.6; 2nd dorsal and caudal lO.i.

3. Whitley (Aust. Zool., 9 [3], 1939: 240) proposes the new name albimors for the Australian Carcliaroion earlier
described by McCoy (Prod. Zool. Victoria, Decade *, 1885: pi. 74) as Carcharodon rondelletii. But there is noth-
ing- apparent, either in McCoy's account or in his measurements, or in Whitley's subsequent illustrations (Fish.
Aust., /, 1940: 126) to set it apart from the Carcharodon of the Atlantic.

4. Contributed by J. W. Lowes, James Miller and Stewart Springer.

5. From measurements by Springer (Copeia, 2, 1939: 115).

Fishes of the Western North Atlantic

135

Trunk fusiform, moderately stout, broadest and highest opposite ist dorsal fin and
tapering to caudal peduncle, the girth In specimens 6 to 7 feet long about 58 to 62% of
total length. Caudal peduncle strongly depressed dorso-ventrally and widely expanded

(^ (A 1^^

-t} d^'^ t^ d:::!

■AAc^di

Figure 20. Carcharodon carcharias, young male, about seven feet long, after Garman, with some emendations
from photographs of fresh specimens. A Dermal denticles, after Garman. B Teeth of a Woods Hole, Massa-
chusetts, specimen about 8J/2 feet long (U.S. Nat. Mus., No. 10899), about 0.7 natural size. C Fourth upper
tooth. D Eighth upper tooth. E Fourth lower tooth. F Eighth lower tooth of same. C-F about 1.4 x.

Figure 2 1 . Carcharodon carcharias. Lower view of head
of specimen pictured in Fig. 20, after Garman.

136

Memoir Sears Foundation for Marine Research

laterally, as in Isurus, its breadth, including its lateral keel-like extensions, about 3 times
its depth, with a prominent transverse furrow above and below just in front of oria;in of
caudal. Dermal denticles minute, as in Isurus, 3-ridged, their free margins correspond-
ingly indented, the blades so nearly flat that the skin is hardly rough to the touch.

Head 25 to 30% of total length. Snout shorter than in Isurus, its length in front of
mouth a little less than % of length of head, obtusely conical, but somewhat flattened
dorsally, with blunt tip; but in large, heavy specimens, suspended or dragged up on the
beach by the front of the mouth, the head is often greatly distorted in appearance as seen
in photographs, since the upper jaw is slightly protrusible. Eye small, circular, its anterior
margin a little posterior to front of mouth. Spiracle lacking in fresh specimens seen by us,
pore-like if present, behind eye by a distance about equal to length of snout in front of
mouth. Gill openings as in Isurus, noticeably long, the 5th a little the longest, between
1.5 to 2 times as long as snout in front of mouth, the ist shortest, the spaces between them
successively narrower from front to rear, that between 4th and 5th being only about Yz as
great as that between ist and 2nd, the 5th close in front of origin of pectoral and curving
posteriorly around latter. Nostril narrow, transverse, near side of head, nearer to mouth
than to tip of snout, its anterior margin with very low subtriangular lobe. Mouth broadly
rounded, a little more than twice as broad as high. Labial furrows very short, the lower
concealed except when mouth is open.

^^^p^f9?^^

Figure 22. Carcharodon carcharias, about six feet long, from Long Island, New York (Amer. Mus. Nat. Hist.,
No. 14773). ^ Upper and lower teeth, right-hand side, about natural size. B Fourth upper tooth. C Seventh
upper tooth. D Third lower tooth. E Seventh lower tooth. B-D, about 2 x.

Fishes of the Western North Atlantic 137

Teeth ^^'^^^ in each side of mouth, large/ subtriangular, erect or very slightly
oblique, their edges coarsely and regularly serrate; uppers about as high as broad, ist with
inner margin nearly straight, but others with both margins usually slightly concave, the
outer edge the more so;' lowers narrower than uppers, their margins more concave; ist
and 2nd teeth the largest in each jaw, those toward corners of mouth successively smaller,
the outermost 2 or 3 minute; ist and 2nd lowers in small specimens (Fig. 22 A, D) with
basal serrations considerably the largest; i, or at most 2, series functional in each jaw. Gap
at symphysis wider in lower jaw than in upper.

First dorsal nearly an equilateral triangle, its apex rounded, its rear margin only
slightly concave, Its free rear tip only about V4 as long as its base, its origin opposite or a
little anterior to inner corner of pectoral.' Second dorsal only y^ to Ve as large in linear
dimensions as ist, Its apex rounded, its margins nearly straight, the rear end of Its base
over, or a litde anterior to, origin of anal. Upper anterior and lower anterior outlines of
caudal moderately convex, posterior outline lunate, with strongly marked subtermlnal
notch, the tips subacute, the lower anterior margin about % (76 to 92%) as long as upper
anterior margin, each measured from precaudal furrow. Anal similar In size and shape to
2nd dorsal, and wholly behind latter. Pelvics much larger than 2nd dorsal or anal, their
anterior margins about ^2 as long as anterior margin of ist dorsal, their distal margins
concave, their corners rounded. Pectoral noticeably larger in area than in Isurus, a little
less than Yi as long as from tip of snout to origin of caudal, and considerably less than V2
as broad as long, with convex anterior and concave posterior margins, subacute tip and
rounded inner corner.

Color. Specimens up to 1 2 to 1 5 feet long, including those seen by us, are slaty-brown,
dull slate-blue, leaden gray, or even almost black above, shading more or less abruptly
to dirty white on the lower surface with a black spot in the axil of the pectoral; the tips of
the pectorals also black, usually with some adjacent black spots; the dorsals and caudal
dark along rear edges, but the pelvics darkest (olive) along anterior edges, fading rear-
ward to white. Large specimens (perhaps some smaller ones also) are described as dun-
colored above, or even leaden-white. They may also lack the black axlllar spot."

Size. This shark has been credited repeatedly with reaching a length of 40 feet. Actu-
ally, however, the stated length of the Australian specimen on which the foregoing has
been based, the jaws of which are now In the British Museum, was 36V2 feet." The next
largest, the actual capture of which is authentically recorded, was reported as of about 30
feet, seemingly not measured." However, these appear to have been giants of their kind,
for while 20 to 25-footers have been reported as seen on several occasions, the three next

6. The larg^est teeth of a specimen 36/^ feet long were about two inches long.

7. Individual teeth vary in this regard, irrespective of their positions along the jaws.

8. Sometimes shown as a little behind inner corner of pectoral in photographs of specimens suspended by mouth, and
hence more or less distorted.

9. Personal communication from Stewart Springer, from his obser\'ations on about a dozen large Florida specimens.

10. Giinther, Cat. Fish. Brit. Mus., 8, 1870: 392; Guide to Study Fish., 1880: 321.
n. Jordan and Evermann, Bull. U.S. nat. Mus., 47 (i), 1896: 50.

138 Memoir Sears Foundation for Marine Research

largest actually measured have been 21 feet^" and 17 to 19 feet in length. We should
perhaps caution the reader that estimates of the size of the larger sharks are frequently
much too high; e.g., an Australian specimen, reported in the local newspapers as 16 feet
long, actually measured only eight feet six inches.'^ On the other hand, the smallest free-
living specimen of which we find record was about 5 feet long." Among 44 other speci-
mens from various localities, of which measurements are available, 15 were between 6 and
8 feet; 7 between 8 and 10 feet; 9 between 10 and 12 feet; 7 between 12 and 14 feet;
4 between 14 and 16 feet; 2 between 16 and 18 feet. The two gravid females on record
were 14 feet 9 inches and 18 feet 4 inches (5.7 m.) in length; similarly the gonads of a
male of 14 feet 6 inches, taken off Salerno, Florida, were much enlarged, but other males
of 12 to 12V2 feet showed no signs of approaching maturity.'^ The fact that females of
8 feet 6 inches and 1 5 feet 6 inches have been reported as containing neither embryos nor
even enlarged ova suggests that sexual maturity is not usually reached at a length less than
perhaps 13 to 14 feet. That so few adults are captured anywhere is no doubt due to their
large size, great strength and formidable nature.

Recorded weights of Atlantic specimens in relation to length are: 600 pounds at 8
feet 3 inches; 960 pounds at 9 feet 8 inches; 998 pounds at 12 feet; 940 pounds at 12 feet
2 inches; about 1,300 pounds at about 13 feet; and 7,100 pounds, with a liver of 1,005
pounds,^" at 21 feet (Cuban specimen mentioned above, see footnote 12, page 138);
also an estimated weight of 1,200 pounds for a specimen 12 feet 8 inches long. Weights of
Pacific specimens taken on the coast of the State of Washington are: 342 pounds at 8 feet
2 inches; between 800 and 1,000 pounds at about 12 feet; up to 2,000 to 2,400 pounds at
13 feet.^' A 5 foot 4 inch specimen from Catalina Island weighed 87 pounds.^* Australian
data'" show: 928 pounds at 1 1 feet 3 inches; 910 pounds at I2 feet 6 inches; 1,291 pounds
at 13 feet 6 inches; 1,334 pounds at 13 feet 5 inches; and 1,720 pounds at 15 feet 2 inches;
a South African specimen of only 13 feet 3 inches weighed 2,176 pounds.'" The variation
in weight at given lengths with differences in the condition of the individual specimens is
thus very wide, and increasingly so with growth.

Develo-pmental Stages. No account of the developmental stages has yet appeared.
The few embryos so far reported have ranged in length from about 20 to 61.6 cm. A
Mediterranean specimen, probably of this species, contained nine young, each about two
feet long."^

12. Taken recently off Havana, Cuba, and reported to us by Luis Howell-Rivero.

13. Madeay, Proc. Linn. See. N. S. W., ^, 1880: 459.

14. Doderlein (Man. IttioL Medit., 2, 1881 ; 66) reports a specimen of .63 in., or about 2 feet, but this may have
been an embryo.

15. Personal communication from Stewart Springer.

16. We have received a good photograph, apparently of this specimen, with weight stated at 7,302 pounds, from
OUyandro del Valle.

17. Bonham, Copeia, 1942: 264. 18. Personal communication from W. \. Follett.
19. Whitley, Fish. Aust., 1, 1940: 127. 20. London Illustr. News, July 14, 1928:53.

21. Norniaii and Fraser, Giant Fishes, 1937: 18; but the stated weights of these embryos (about 100 pounds at a
length of two feet) were evidently in error.

Fishes of the Western North Atlantic 139

Habits. This is an active, strong-swimming species, putting up a dogged and savage
resistance to capture. The reports of it attacking boats, when harpooned or hooked, are too
numerous and too circumstantial to be dismissed. However, it does not have the leaping
habit of the Mako. So few are seen that nothing is known of its life apart from the fore-
going and the fact that it is voracious. The great majority of records have been of speci-
mens taken at the surface or close to it. But it appears that they may descend to consider-
able depths, for a large one caught off the north coast of Cuba, of which we have a photo-
graph, was said to have been hooked at a depth of 700 fathoms. Nothing is known of its
breeding habits.

Characterization of this Shark by an earlier student as "the most voracious of
fish-like vertebrates,""' is no doubt well deserved. The frequency with which it captures
large prey, which it devours practically intact, is illustrated by the presence of other sharks
from four to seven feet long, as well as a young sea lion of 100 pounds, in the stomachs
of White Sharks} also seals, sturgeons and tuna have been found in specimens no larger
than eight to nine feet. Sea turtles are also described as a regular item in its diet in southern
waters. On the other hand, it also preys on a wide variety of smaller fishes and marine ani-
mals, including chimaeroids and squids. The mouth of a Massachusetts Bay specimen re-
cently examined by us was festooned with hooks and snoods from a long line, while its
stomach contained a spiny dogfish evidently torn off a hook. This, together with similar
reports by others, including the report of a large Florida specimen containing two Car-
charhinus milberti six to seven feet long which were evidently torn from hooks on the set-
line on which the Carcharodon itself was taken,^' shows that when White Sharks stray in on
the fishing grounds they find a convenient source of food.

It has been described also as a scavenger when occasion offers; for example, the
stomach of a shark said to be this species, caught in Sydney Harbor, New South Wales,
contained a variety of garbage, including horse meat, legs of mutton^ parts of a pig, a
dog, etc.

Relation to Man. This is perhaps the only shark against which the charge of un-
provoked attack on small boats is proved through identification of the teeth left imbedded
in the sides of the boat. It has borne an unsavory reputation from the earliest times as a
man-eater. It is so classed, for example, in Australia, where attacks by sharks on bathers,
especially near Sydney, are of such common occurrence that most of the bathing beaches
are protected by wire-netting enclosures."* It is not possible to tell whether men, reported
by earlier authors to have been found in the stomachs of White Sharks, were alive or dead
when eaten 5 but it is probable that a seven-foot specimen, taken a few days later in Sandy
Hook Bay at the mouth of New York Harbor, was responsible for four shark fatalities

2i. Jordan, Guide to Study Fish., 1905: 538.

23. Springer, Copeia, 2, 1939: 114.

24. See Coppleson (Med. J. Aust., April 15, 1933: 449) and Whitley (Fish. Aust., /, 1940: 259) for circumstan-
tial accounts (many of them recent) of shark fatalities in Australia. In most cases the species of shark responsible
was not determined.

140 Memoir Sears Foundation for Marine Research

that occurred on the bathing beaches of New Jersey from July 6 to 12, 1916.^' A Car-
charodon also may have been responsible for the fatal attack on a swimmer at Mattapoisett
on Buzzards Bay, Massachusetts, on July 25, 1936;"° in this case the shark was driven
away and not identified. However, these are the only recently recorded instances any-
where on the eastern seaboard of the United States in which Carcharodon is under suspi-
cion. Hence, while the possibility of attack by it on bathers is always present, since White
Sharks do occasionally come close inshore near populous sectors of the coast line, it is
exceedingly remote. The most recent report of an attack by this species (fatal in this in-
stance) was of a 6- to 7-foot specimen on a swimmer in Panama Bay, the species being
identified by a well known ichthyologist on the basis of fragments of its teeth taken from
wounds by the surgeon attending the victim."

In spite of its ferocity and its muscular power, the White Shark does not put up as
spectacular a resistance as the Mako when hooked (p. 129), not having the habit of
jumping. Nor does it seem to make as strong a fight, pound for pound, as the tuna or
the swordfish. For example, it is recorded that a 1,329-pound specimen was landed on rod
and reel by an angler after fifty-three minutes in Australia j"* another of 2,176 pounds
was landed in South Africa from the shore in five hours,"' the latter one of the largest, if
not the largest, fish ever landed on rod and reel.°°

Range. Oceanic; widespread in tropical, subtropical and warm temperate belts of all
the oceans, including the Mediterranean; exceedingly irregular in its occurrence; appar-
ently most numerous in Australian waters, but nowhere abundant.

Occurrence in the Atlantic. Although this shark has been so long known and so much
discussed because of its ill repute, very little detailed information is available as to its
geographic distribution anywhere. While repeatedly reported from the Mediterranean
and from many other localities, it certainly is not common there. It appears to be decidedly
scarce on the eastern side of the open Atlantic, it being positively recorded, so far as we can
learn, only from the Cape of Good Hope region, from Morocco, Rio de Oro, Mauritania,
Senegal, the Canaries, and from the coast of the Iberian Peninsula, with nominal records
from the vicinity of Teneriffe and Madeira.

The list of positively identified captures for the tropical-subtropical belt in the v/est
is limited to one record for Brazil (several times repeated by subsequent authors) ; one
from St. Lucia in the West Indies; one from the vicinity of Nassau in the Bahamas; four
from the west coast of Florida; and one or two from the east coast. Reputedly, however,

25. Nichols and Murphy, Brooklyn Mus. Quart., 4, 1916; 157.

26. The victim was taken to the hospital in New Bedford, where he died.

27. Reported in J. Amer. med. Ass., July 22, 1944, and in Science News Letter, July 29, 1944: 73. Identification by
J. T. Nichols.

28. Whitley, Fish. Aust., i, 1940: 126.

29. London lUustr. News, July 14, 1928: 53, photograph; recorded as a Mako, but identifiable by the teeth as a
Carcharoion.

30. For a graphic account of the capture of one 9 feet 2 inches long oflF Virginia by an angler, see Wise (Tigers of
the Sea, 1937: 61).

Fishes of the Western North Atlantic 141

it is considerably more plentiful among the West Indies than the paucity of the published
records would suggest; this is certainly true along the east coast of Florida, where one
correspondent (a well known student of sharks) reports the recent capture in the shark
fishery of about a dozen fair-sized ones." To the northward it is either more plentiful or at
least more often caught or reported. Thus, four were taken near Cape Lookout, North
Carolina, during the summer of 191 8, with others reported as seen in that and previous
summers; one is recorded off Smith Island, Virginia; three or four from the coast of
New Jersey, with others reported by sport fishermen.^^ Occasional specimens are encoun-
tered oif New York; a small one of about five feet was taken in a pound net at Sakonnet,
Rhode Island, May 30, 1939.'^ Nine or ten are definitely listed and several additional
ones are reported from the Woods Hole region and Nantucket, with two at the most,
however, in any one year. While it is generally considered a warm water species, reliable
reports of its presence have been received more often from the southwestern part of the
Gulf of Maine than from any other coastal sector of comparable length on the American
seaboard. In Massachusetts Bay alone at least nine were either actually captured or har-
pooned and lost during the period from 1935 to 1940, with stray specimens taken for
earlier years back to 1848, most of them in the vicinity of Cape Cod. Still farther north
there are scattered records for the vicinity of Portland, Maine (2),'* the most recent a 13-
foot specimen, taken in a gill net off Casco Bay in November 1931 ; from Eastport at the
mouth of the Bay of Fundy (i), and from Digby, Nova Scotia, within the Bay (i). It
may visit the outer coast of that Province more often than formerly supposed, there being
several reliable records for St. Margaret Bay, and perhaps for Halifax also. The most
northerly record for American waters is St. Pierre Bank, south of Newfoundland, where
one attacked a fisherman many years ago in a dory, leaving in the sides of the boat frag-
ments of its teeth, by means of which Dr. Garman was able to identify it."

The fact that all records of its presence off the northeast coast of the United States
and Canada are for the warm half of the year suggests that it is an oceanic visitor, but
nothing whatever is known of its status offshore in the western Atlantic, there being no
record of its presence around Bermuda.

Although typically an inhabitant of the high seas, it frequently comes inshore and
even into very shallow water, as in the following cases: one taken inside Sandy Hook Bay,
New York, in 19 16; a considerable number that have been picked up at different times
in the fish traps within a few yards of the beach in the vicinity of Woods Hole and on Cape
Cod; one harpooned in 10 feet of water in Provincetown Harbor many years ago; two
specimens caught close to Boston Harbor in 1 839; one harpooned about two miles off one
of the most popular bathing beaches at the mouth of Boston Harbor in 1937; another simi-

31. Personal communication from Stewart Springer.

31. A recently received photograph, supposedly of a Mako taken off New Jersey in October 1935, unmistakably

represents a Carcliaroion of 11 to 12 feet.
33. Photograph received from James Miller. 34. Received from Walter H. Rich.

35. Putnam, Bull. Es^ex Inst., d, 1874: 72.

142 Memoir Sears Foundation for Marine Research

larly harpooned within half a mile of the land off Cohasset, on the southern side of
Massachusetts Bay in August 1940/°

Synonyms and References:

I. Atlantic:''

Squalus carcharias Linnaeus, Syst. Nat., /, I 758: 235 (type loc, Europe) ; Brunnich, Ichthyol. Massil., 1768:
5 (Adriatic, food); Bloch, Naturg. Ausland. Fische, 4, Atlas, 1758: pi. 119; Gmelin, in Linnaeus, Syst.
Nat., 13th ed., 5, 1789: 1498 (part) ; Walbaum, P. Artedi Genera Pise. Emend. Ichthyol., 3, 1792: 514
(descr., Medit., Atlant.) ; Bloch and Schneider, Syst. Ichthyol., 1801: 132 (descr., ref.) ; Blumenbach,
Handb. Naturg., 7, 1803: 263 (ref.) ; Bosc, Nouv. Diet. Hist. Nat., 27, 1803: 185 (general) ; Latreille,
Nouv. Diet. Hist. Nat., 24, 1804: 72 (in table of contents) ; Nardo, Prod. Itiol. Adriat., 1827: 9 (Ad-
riatic) ; Voigt, in Cuvier, Tierreich, 2, 1832: 505 (descr.) ; Griffith, in Cuvier, Anim. Kingd. with Adds.,
70, 1834: 599 (general); Bonaparte, Mem. Soc. neuchatel Sci. nat., 2 (8), 1839: 10 (in synopsis);
Nardo, Atti 1st. veneto, (3) 5, 1859-1860: 787 (Medit.); Gemellaro, Sagg. Itiol. Catania, 1864: 120
(Medit., not seen) ; Vieria and Clavijo, Dice. Hist. Nat. Isl. Canaries, Real Soc. Econ. Las Palmas, 1866-
1869, Mss. of 1799 (Canaries, not seen) ; Buckland, Hist. Brit. Fish., 1881 : 211 (general).

White Shark, Brookes, Nat. Hist., 5, 1763: 28 (general); Pennant, Brit. Zool., 5, 1769: 82 (general); also
subsequent ed.; Pennant, Arctic Zool., Suppl., 1787: 105 (America).

Cane carcharia Cetti, Amfib. Pesci Sardegna, 1777: 69 (Sardinia).

Squale requin, Lacepede, Hist. Nat. Poiss., 7, 4° ed., 1798: 165, 169, in Buffon, Hist. Nat. (part, combined
with Carcharkinus), not pi. 8, fig. I, which is Carcharhinus; in Sonnini, Hist. Nat. Poiss., 5, I 802: 332
(part, combined with Carcharhinus, see p. 320), pi. 4, fig. 2, 3 (teeth), not pi. 4, fig. I, which is
Carcharhinus (see p. 3 2o) .

Carcharias lamia Rafinesque, Indice Ittiol. Sicil., 1810: 44 (substituted for Squalus carcharias Linnaeus, I 758,
Sicily); Fitzinger, Bild. Atlas Naturg. Fische, 1864: fig. 169 (ill.).

Squalus [Carcharias) carcharias Cuvier, Regne Anim., 2, I 81 7: I 26 (general, size).

Carcharias terus Cloquet, Diet. Sci. Nat., 7, 1822: 69 (general); Reguis, Ess. Hist. Nat. Provence, 7 (l),
1877: 46 (Medit.).

Squalus {Carcharhinus) carcharias Blainville, in Vieillot, Faune Franc, 1825: 89 (descr.).

Carcharias carcharias Bory de St. Vincent, Diet. Class. Hist, nat., 75, 1829: 596; Cuvier, Regne Anim., 111. ed.,
Poiss., 1842-1843: 360, pi. 114, fig. 2a (tooth); Jordan, Copeia, 140, 1925: 20 (name).

Carcharias rondeletti Bory de St. Vincent, Diet. Class. Hist, nat., 75, 1829: 596 (general); Reguis, Ess. Hist.
Nat. Provence, 7 (l), 1877: 47 (Medit.).

Squalus {Carcharias) vulgaris Richardson, Fauna Boreal. Amer., 5, 1836: 288 (ref.).

Carcharias (no specific name), Agassiz, L., Poiss. Foss., j, 1835: pi. F, fig. 3 (teeth, see footnote I, p. 133).

Carcharodon smithii Agassiz, L., Poiss. Foss., 5, 1838: 91 (name only) ;^* Bonaparte, Mem. Soc. neuchatel.
Sci. nat., 2 (8), 1839: 9 (in synopsis).

Carcharodon verus Agassiz, L., Poiss. Foss., j, 1838: 91 (teeth).

Carcharias vulgaris Hamilton, Brit. Fish., 2, 1843: 304, and subsequent eds. (Gt. Brit., general).

CfTrc/^arOi/ora/^wz/a Bonaparte, Icon. Faun. Ital., 5 (2), I 839: pi. [52] (descr., colored ill., general) ; Cat. Pesc.
Europ., 1846: 17 (Medit.); Sassi, Cat. Pcsci Liguri., 1846: 123 (Medit., not seen); Nardo, Atti 1st.
veneto, (3) 5, I 859-1 860: 787 (Medit.) ; Ninni, An. Soc. Nat. Modena, 5, 1870: 66 (Medit.) ; Morc-au,
Hist. Nat. Poiss. France, 7, 1881: 302 (descr., France); Bellotti, Atti Soc. ital. Sci. nat., jj, 1891: 111
(name) ; Belloc, Rev. des Trav. Peches Marit., 7, Fasc. 2, 1934: 138 (ill. after Bonaparte; Morocco, Rio
de Oro, Mauritania, Senegal).

Carcharodon rondeletti Miiller and Henle, Plagiost., 1841: 70 (Brazil, Australia, descr., meas.) ; Gray, List
Fish. Brit. Mus., 1851: 61; Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861: 60 (name); Dumeril,
Hist. Nat. Poiss., 7, 1865: 41 1 (descr., size, Medit., Algeria, Atl., C. Good Hope), pi. 7, fig. 7 (tooth) ;

36. We had the opportunity of examining two of these Massachusetts Bay specimens soon after they were landed,
and have received photographs of others.

37. For Indo-Pacific references, see p. 144. 38. Nomen nudem.

Fishes of the We sterol North Atlantic 143

Bocage and Brito Capello, J. Sci. math. phys. nat. Lisboa (l868-l86g), 2, 1870: 140 (Portugal);
Gunther, Cat. Fish. Brit. Mus., 8, 1870: 392, 518 (descr., Cape Seas, jaws of jSyi-h. spec,
Australia); Canestrlni, in Cornelia, et al., Fauna d'ltal., 1872: 45 (Medit.) ; Doderlein, Prosp.
Metod. Pesci Sicil., 1 878-1 879: 30 (Medit.) ; Stossich, Boll. Soc. adriat. Sci. nat., 5, 1 880: 68 (Adriatic) ;
Doderlein, Man. Ittiol. Medit., 2, 1 881: 66 (Medit.) ;^° Perugia, Elenc. Pesc. Adriat., 1881: 53 (sizes,
Adriatic spec); Hasse, Naturl. Syst. Elasm. besond. Theil, 1882: 224, pi. 30, fig. 24—33 (vertebrae,
dermal dent.); Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 46; Faune Senegambie, Poiss., i,
1883-1885: 23 (Senegambia) ; Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 30 (American
coast); Graeffe, Arb. zool. Inst. Univ. Wien, 7, 1886: 446 (Medit.); Carus, Prod. Fauna Medit., 2,
1889-1893: 506 (Medit.); Sicher, Atti Accad. gioenia, (4) 11, 1898: 16 (Medit.); Bridge, Camb.
nat. Hist., 7, 1904: 451 (general); Werner, Zool. Jb., Syst. Abt., 2j, 1904: 285 (embryo); Gunther,
Encyc. Brit., nth ed., 24, 191 1: 807 (general); Metzelaar, Trop. Atlant. Vischen, 1919: 190 (Sene-
gambia); Norman and Eraser, Giant Fishes, 1937: 14 (general, embryos).

CoTcharias atwooii Storer, Proc Boston Soc. nat. Hist., 5, 1848: 72 (Provincetown, Massachusetts); Gill,
Proc. Acad. nat. Sci. Philad., Addend., 1861 : 59 (listed) ; Storer, Mem. Amer. Acad. Arts Sci., 9, 1867:
222, pi. 36, fig. 4; also as Fishes of Mass., 1867: 246, pi. 36, fig. 4 (descr., ill., Massachusetts) ; Gill,
Rep. U.S. Comm. Fish. (1871-1872), 1873: 813 (Newfoundland to Florida); Smith, in Verrill, Rep.
U.S. Comm. Fish. (1871-1872), 1873: 576 (parasites. Vineyard Sound).

Carcharodon alzvoodi Lyman, Rep. Mass. Comm. inl. Fish., 1872: 53 (Massachusetts) ; Smith, S. J., Rep. U.S.
Comm. Fish. (1871-1872), 1873: 576 (Vineyard Sound) ; Baird, Rep. U.S. Comm. Fish. (1871-1872),
1873: 827 (Woods Hole); Goode, in Verrill, Rep. U.S. Comm. Fish. (1871-1872), 1873: 812 (listed
C. Cod to Florida) ; Goode and Bean, Bull. Essex Inst. Salem, rr, 1879: 38 (Massachusetts) ; Rathbun,
Proc U.S. nat. Mus., 7, 1885: 488, 489 (parasites).

Carcharodon (without specific name) Owen, Odontogr., 1840-1845: 30 (size of teeth in 37-ft. spec).

Carcharias (Prionodori) lamia Putnam, Bull. Essex Inst. Salem, 6, 1874: 72 (St. Pierre Bank, S. of Newfound-
land).

Ci»rc^aro<2fo« «rc/wria^ Perez-Areas, An. Soc. esp. Hist, nat., 7 (2), 1878: 15 (Valencia, Spain; history) ; Jordan
and Gilbert, Bull. U.S. nat. Mus., 16, 1883:875 (name) ; Stephenson, Trans. Vassar Bros. Inst., 2, 1884:
83, pi. 1,2 (meas., Nantucket) ; Amer. Nat., 18, 1884: 940, pi. 31 (same specimen as preceding) ; Goode,
Fish. Fish. Industr. U.S., i, 1884: 670 (habits, ferocity, distrib.) ; Jordan, Rep. U.S. Comm. Fish.
(1885), 1887: 797 (distrib.); Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 218
(distrib.); Bull. U.S. nat. Mus., 47 (l), 1896: 50 (descr., distrib., 30-ft. Calif, spec); Smith, H. M.,
Bull. U.S. Bur. Fish., ij, 1898: 89 (Woods Hole); Bean, Rep. For. Comm. N. Y., 1901: 380 (old
C. Cod records) ; Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 40; also in Rep. N. Y. St. Mus. (1902), 3 (5),
1904: 40 (general) ; Jordan, Guide to Study Fish., i, 1905: 538 (general) ; Ribeiro, Arch. Mus. nac Rio
de J., 14, 1907: 160, 203 (Brazil, synonyms); Wilson, Proc. U.S. nat. Mus., S3, 1907: 371, 414, 423
(Woods Hole, parasites) ; Kendall, Occ Pap. Boston Soc. nat. Hist., 7 (8), 1908: 7 (New England) ; Berg,
Faune Russie Poiss., /, 191 1: 55 (not seen); Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., 31 (2),
1913: 737 (Woods Hole region); Halkett, Check List Fish. Canad., 1913: 40; Garman, Mem. Harv.
Mus. comp. Zool., j(5, 191 3: 32, pi. 5, fig. 5-9 (descr., Massachusetts Bay spec.) ; RadclifTe, Trans. Amer.
Fish. Soc, 1914: 37 (teeth) ; Coles, Proc biol. Soc. Wash., 28, 191 5: 91 (N. Carolina) ; Smith, H. M., J.
Amer. Mus. nat. Hist., 16, 1916: 341, 343 (Martha's Vineyard, Mass.; Florida; danger to man) ; Fowler,
Copeia, 30, 1916: 36 (Middle Atlant., U.S.); Hussakoff, Copeia, 57, 1916: 86 (N. Jersey); Nichols,
Copeia, 37, 1916: 87 (N. Jersey) ; Nichols and Murphy, Brooklyn Mus. Quart., 3 (4), 1916: 145-157
(N. Jersey, shark fatalities of 1916) ; Nichols and Murphy, Brooklyn Mus. Sci. Bull., ^ (l), 1916: 24
(Woods Hole) ; Radcliffe, Bull. U.S. Bur. Fish., 34, 1916: 247 (N. Carolina, teeth. Woods Hole) ; Coles,
Copeia, 69, 1919: 34 (descr., habits, ill., N. Carolina records) ; Fowler, Proc. biol. Soc. Wash., jj, 1920:
144 (N. Jersey, fatalities) ; Proc. Acad. nat. Sci. Philad., y2, 1921 : 385 (N. Jersey) ; Huntsman, Contr.
Canad. Biol. (1921), 5, 1922: 56 (Bay of Fundy) ; Meek and Hildebrand, Field Mus. Publ. Zool., 15

39. For locality references from the Mediterranean in publications not accessible to us, see Doderlein.

144 Memoir Sears Foundation for Marine Research

(l), 1923: 63 (general); Ribeiro, Fauna Brasil., Peixes, Mus. nac. Rio de J., 2 (i) Fasc. i, 1923: 18
(Brazil); Wilson, Proc. U.S. nat. Mus., 64 (17), 1925: 12 (Woods Hole); Bigelow and Welsh, Bull.
U.S. Bur. Fish., 40 (1), 1925: 39 (Gulf of Maine) ; Nichols and Breder, Zoologica, 9, 1927: 19 (gen-
eral); Hildebrand and Schroeder, Bull. U.S. Bur. Fish., 43, 1928: 46 (general); Rey, F.iuna Iberica
Feces, i, 1928: 788 (Spain); Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 22 (general); Truitt,
Bean and Fowler, Bull. Md. Conserv. Dept., j, 1929: 29 (off Maryland) ; Jordan, Manual Vert. Anim.
NE. U.S., 1929: 12 (general); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930:
20 (distrib.) ; Townsend, Bull. N. Y. zool. Soc, 34 (6), 1931: 169 (St. Lucia, W. I., Australia) ; Bellon
and Mateu, Notas Inst. esp. Oceanogr., 2 (53), 193 i: 11 (Canaries); Wilson, Bull. U.S. nat. Mus., 20,
1932: 428, 432, 461, 464 (Woods Hole, parasites) ; Young and Mazet, Shark, Shark, 1933: 267 (gen-
eral) ; Piers, Proc. N.S. Inst. Sci., 18 (3), 1934: 192 (Nova Scotia, Bay of Fundy) ; Bigelow and Schroe-
der, Canad. Atlant. Fauna, biol. Bd. Canad., 12', 1934: 15 (descr., distrib.); Vladykov and McKenzie,
Proc. N. S. Inst. Sci., /p (i), 1935:47 (Nova Scotia) ; Fowler, Bull. Amer. Mus. nat. Hist., yo (i), 1936:
31 (Canaries, Madeira, Senegambia, N. Jersey); Breder, Bull. N. Y. zool. Soc, 59 (3), 1936: 161
(Bahamas, N. Jersey) ; Bull. N. Y. Zool. Soc, 59 (6), 1936: 243 (Nova Scotia) ; Bigelow and Schroeder,
Bull. U.S. Bur. Fish., 48, 1936: 322 (Gulf of Maine, Nova Scotia); Heilner, Schrenkeiscn and La
Monte, Field and Stream, Feb. 1936: 27 (size) ; White, Bull. Amer. Mus. nat. Hist., 74, 1937: 14, pi. 12,
fig. B; pi. 38, fig. F (anat.) ; Wise, Tigers of the Sea, 1937: 172 (Virginia, capture by angler); Tor-
tonese, Atti Soc. ital. Sci. nat., yy, 1938: 292 (Medit.) ; Springer, Proc Fla. Acad. Sci., j, 1939: 36
(Florida); Schroeder, Copeia, 1938: 46 (Massachusetts Bay); Copeia, 1939: 48 (Boston Harbor);
Springer, Copeia, 1939: 114 (Florida); Schroeder, Copeia, 1940: 139 (Massachusetts Bay); Anony-
mous, News Letter, Calif. Acad. Sci., June, 1940 (Nova Scotia); Hildebrand, Copeia, 1941: 222 (N.
Carolina); Norris, Plagiost. Hypophysis, 1941: pi. 19, fig. 73—75 (brain); Fowler, Arqu. Zool. Estado
Sao Paulo, j, 1942: 127 (Brazil); Lunz, Bull. S. C. St. Planning Bd., 14, 1944: 27 (Fla.); Fowler,
Mongr. Acad. nat. Sci. Philad., 7, 1945: 74 (Smith I., Virginia, 242 lb.), 159 (S. Carolina); Bigelow
and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Carib. Comm., Wash., 1945: 105, fig. 36 (descr.,
ill., habits, range).
Great White Shark, Allen, Bull. Boston Soc. nat. Hist., 24, 1921: 8 (Woods Hole).

2. Pacific Ocean, Indian Ocean, South Africa:

Carcharodon cafensis Smith, A., 111. Zool. S. Afr. Pisces, 1849: pi. 4 (S. Africa).

Squalus carcharias, Kitlitz, Denkwurd. Reise Micronesien, 2, 1 85 8: 188 (Bon in I., not seen) ; Bennett, Gather-
ings Nat. Australias, i860: 26 (Aust.) ; Jouan, Mem. Soc. Imp. Sci. Nat. Cherbourg, 8 SuppL, 1861: 5
(New Caledonia).

Carcharias leucas Bennet, Proc. zool. Soc. Lond., 2y, 1859: 223 (New South Wales) ; Jouan, Mem. Soc. Imp.
Sci. Nat. Cherbourg, 14, 1868: 229 (N. Zealand).

Carcharodon rondeletii Hector, Col. Mus. Govt. Surv. Dept. N. Z., 1872: 78 (N. Zealand, not seen) ; Klunt-
zinger, S. B. Akad. Wiss. Wien, 80 (i), 1880: 426 (Aust.); Macleay, Proc Linn. Soc. N. S. W., 4,
1880: 459 (Aust.) ; GiJnther, Introd. Stud. Fish., 1880: 320 (Aust., largest recorded specimen) ; Ramsay,
Proc Linn. Soc N. S. W., 5, 1880: 96 (Aust.); Macleay, Proc Linn. Soc N. S. W., 6, 1881: 358
(Aust.) ; Tcnison-Woods, Fish Fish. N. S. W., 1882: 25 (Aust.) ; McCoy, Prod. Zool. Victoria, Decade 8,
1883: pi. 24 (descr., Aust.); Haswell, Proc. Linn. Soc. N. S. W., 9, 1885: 83, pi. I, fig. 1-4 (skelet.);
Ogilby, Rep. Comm. Fish. N. S. W., 2A, 1887: 2 (Aust.) ; Parker, Proc. zool. Soc. Lond., 1887: 27, pi.
4-8 (anat., embryo, N. Zealand); Philippi, An. Univ. Chile, 7/, 1884: 550, pi. 4, fig. 4 (ill., tooth,
Chilean records); Etheridge, Proc Linn. Soc. N. S. W., (2) j, 1888: 159 (Aust.); Ogilby, Cat. Fish.
Aust. Mus., 14, 1888: 5 (Aust.); Proc Linn. Soc. N. S. W., (2) 3 (4), 1889: 1771 (Aust., abund.) ;
Lucas, Proc. roy. Soc. Vict., 2, 1890: 43 (Aust.) ; Gunther, J. Mus. Godeffroy, (5, Fish. Sudsee j, Heft 9,
1910: 485 (Pacific); Quijada, Bol. Mus. nac. Chile, 5, 1913: ill (listed for Chile).

Carcharodon carcharias Ogilby, Handbk. of Sydney, 1898 (Aust., not seen) ; Jordan and Snyder, Annot. zool.
jap., 5 (2-3), 1901: 127 (Japan); Jordan and Fowler, Proc. U.S. nat. Mus., 26, 1903: 624 (Japan);
Waite, Mem. N. S. W. Nat. Club, 2, 1907: 8 (.Aust., not seen) ; Jordan and Evermann, Bull. U.S. Bur.

Fishes of the Western North Atlantic 145

Fish., 25 (l), 1905: 44 (Hawaiian Is.) ; Stead, Fish. Aust., 1906: 223 (Aust.) ; Waite, Rec. Canterbury
[N. Zealand] Mus., i, 1907: 6 (N. Zealand) ; Zietz, Trans, roy. Soc. S. Aust., 52, 1908; 291 (Aust.) ;
Stead, The Lone Hand, Sydney, 1913: 35 (Aust., not seen); Ogilby, Mem. Qd. Mus., 5, 1916: 74
(Aust.) ; McCulloch, Aust. Zool., / (7), 1919; 223, pi. 17, fig. 23a (Aust.) ; Waite, Rec. S. Aust. Mus.,
2, 1921 : 21 (Aust.) ; Jordan and Jordan, Mem. Carneg. Mus., 10, 1922: 5 (Hawaiian Is.) ; McCulloch,
Roy. Soc. N. S. W., Check List Fish., 1922: 8, pi. 2, fig. 23a (Aust.); Waite, Fish S. Aust., 1923: 40
(Aust.) ; Phillipps, N. Z. J. Sci. Tech., 6, 1924: 269 (N. Zealand) ; Barnard, Ann. S. Afr. Mus., 27 (i),
1925: 33 (S. Afr.) ; McCulloch and Whitley, Mem. Qd. Mus., 8, 1925: 129 (Aust.) ; Jordan and Hubbs,
Mem. Carneg. Mus., 10, 1925: 102 (Japan); Fowler, Proc. Acad. nat. Sci. Philad., y/, 1926: 190 (E.
Afr.); Herre, Philipp. J. Sci., 2(5, 1925: 114 (Philippines); Barnard, Ann. S. Afr. Mus., 21, 1927: 2,
pi. I, fig. 7 (S. Afr.); Phillipps, Fish. Bull. Wellington, N. Z., /, 1927: 9 (synonyms, N. Zealand);
Whitley, Aust. Mus. Mag., 5, 1927: 13 (Aust.) ; McCulloch and Whitley, Fish. roy. soc. N. S. W., 2nd
ed., 1927: 8, pi. 2, fig. 23; also 3rd ed., 1934 (Aust.); Fowler, Mem. Bishop Mus., 10, 1928: 18
(Pacif.) ; Roughley, Aust. Mus. Mag., 3, 1928: 152 (Aust.); McCulloch, Mem. Aust. Mus., 5, 1929:
15 (Aust.); Chapman, Open Air Stud. Aust., 1929: 20, pi. opp. p. 68 (Aust., not seen); Soldatov and
Lindberg, Bull. Pacif. Fish. Res. Sta., 5, 1930: 12 (part);''" Fowler, Proc. Pan-Pacif. sci. Congr., 3
(4), 1930: 488 (Pacific) ; Tanaka, Jap. Fish. Life Colors, 20, 1933 (Jiipan, not seen) ; Coppleson, Med.
J. Aust., April 1932: II (reputation as man-eater); Marchand, Fish. M.ir. Biol. Surv. S. Afr., Fish
Bull. 2, 1935: 31 (Natal, S. Afr.); Walford, Fish. Bull. Sacramento, 45, 1935: 38 (California); Barn-
hart, Mar. Fish, south. Calif., 1936: 10 (off California); Grey, Amer. Angler in Aust., 1937: 48, pi.
34> 35 (Aust., not seen); Tubb, Proc. roy. Soc. Vict., 4^ (2), 1937: 422 (Aust.); Rox.is and Marten,
Dept. Agr. Comm. Manila Tech. Bull., 6, 1937: 12 (Philippines, not seen) ; Beebe and Tee-Van, Zoo-
logica, N. Y., 2(5, 1941: 98 (general) ; Fowler, Bull. U.S. nat. Mus., lOO (75), 1941: HO (Indo-Pacific
records); Bonham, Copeia, 1942: 264 (coast of Washington, weights).

Mako, Anonymous, Lond. Illustr. News, July 14, 1942: 53 (C. Province, S. Afr., caught by angler, size,
photo).

Carcharias maso Morris, Aust. Eng., 1898: 412 (not seen) ; not Squalus {Carcharias) maou Lesson, Voy. "Co-
quille," Zool., 2, 1830: 91, pi. I.

White Pointer, Stead, Giants and Pigmies of the Deep, 1933: 54 (Aust.).

Carcharhinus carcharias Whitley, Mem. Qd. Mus., 10 (4), 1939: 199 (N. Zealand).

Carcharodon albimors Whitley, Aust. Zool., 9 (3), 1939: 240 (Aust.) ; Fish. Aust., /, 1 940: 126 (Aust.).

Great White Shark, Kean, J. Amer. med. Ass., July 23, 1944: 846 (fatal attack, Panama Bay) ; Scheffer and
Slipp, Amer. Midi. Nat., 52, 1944: 399 (8-ft. 2-in. specimen with entire seal in stomach; coast of
Washington) .

Doubtful References:

Squalus carcharias Forskal, Descr. Anim., 1775: VIII, 20 (Red Sea, descr. insufficient); Forster, in Latham

and Davies, Faunula Indica, 1795: 13 (India, name only) ; Labillardiere, Rel. Voy. Rech. de La Perouse,

/, 1800: 40, 226, 399 (near Teneriffe, E. Indies, S. W. Aust., name only) ; Bowditch, Excurs. Gambia

(1823), 1825: 233 (Gambia; name only) ; Forster, Descr. Anim. Obs. Del. Cur. Lichtenstein, 1844: 256

(Tanna, New Hebrides, name only).
Carcharias vorax Owen, Cat. Osteol. Roy. Coll. Surg., i, 1853: 94 (brief notice of vertebrae, teeth, 23-ft.

spec).
Carcharodon rondeUtti Quijada, Bol. Mus. nac. Chile, 5, 191 3: I, 1 1 (Chile, name only).
Not Squalus carcharias Pallas, Zoogr., Rosso Asiat., 5, 1 831: 63''^ (the localities, Kamchatka and Bering Sea,

make it probable that these records actually refer to some other shark).
Not Carcharodon atzvoodi L'hler and Lugger, Rep. Comm. Fish. Maryland, 1st ed., 1876: 191; 2nd ed.,

1876: 161 (reported "common" in Chesapeake Bay, hence doubtless some other shark).

40. Includes report by Pallas, 1831, of Squalus carcharius from Kamchatka and Bering Sea; prob.^bly not this species.

41. First printing may have been 18 14: see Sherborn, Ibis, (13), 4. '934: >66-

146 Memoir Sears Foundation for Marine Research

Family CETORHINIDAE

Characters. Essentially those of Isurldae (p. 109), except that each gill arch bears
a great number of long, horny bristle-like rakers directed forward, analogous to those of
many bony fishes; the gill openings are very much larger; the teeth are minute, very nu-
merous, and conical with one cusp; the dorso-rostral cartilages are very slender, and the
ventro-rostral cartilage broad and blade-like^ (in the Isuridae all three of the rostral
cartilages are rod-like, and about equally stout) .

Remarks. The Cetorhinidae have usually been placed among the Isuridae, of which
they appear to be an offshoot. However, the presence of horny rakers on their gill arches,
a character which makes them unique among modern sharks, suggests to us that they be
classed as a distinct family.

Genera. Only one genus, Cetorhinus Blainville, 1 8 1 6.

Genus Cetorhinus Blainville, 1 8 1 6''
Basking Sharks

Cetorhinus Blainville, Bull. Soc. philom. Paris, 1816: 121 ; type species, Squalus gunnerianus Blainville, 1810,
equals Squalui maximus, Gunnerus, 1765.'

Generic Synonyms:

Squalus (in part) Gunnerus, K. norske Vidensk.-Selsk. Skr. Trondh., 1765: 33; and subsequent authors; not

iSyWtt/ Linnaeus, 1758.
Selache Cuvier, Regne Anim., 2, 1817: 129; type species, Selache maxima Cuvier, equals Squalus maximus

Gunnerus, 1765.
Selanche Jaroki, Zoologi, 4, 1822: 452 (not seen); type species, S. maximus Jaroki, equals Squalus maximus

Gunnerus, 1765.
Selachus Minding, Lehrb. Naturg., 1832: 52 (not seen); type species, Selachus maximus Minding, equals

Squalus maximus Gunnerus, 1 765.
Polyfrosofus Couch, Brit. Fishes, i, 1867: 68; type species, P. m.acer Couch; type locality English Channel.
Cethorhinus Escribano, An. Soc. esp. Hist, nat., p, 1909: 340; type species, C. maximus Escribano, equals

Squalus maximus Gunnerus, 1 765.

Doubtful Synonyms:

Halsydrus Fleming, Scots. Mag. Edinb. Misc., 1809: 6; type species, H. fontoffidani Fleming; type locality,

Orkney Islands.*
Tetraoras Rafinesque, Carratt. Gen. Nuov. Sicil., 1810: 46; type species, T. angiona Rafinesque.

1. The dorso-rostral cartilages have been pictured either as uniting some distance posterior to the point of union
between the resultant bar and ventral cartilage (Senna, Arch. ital. Anat. Embriol., 22, 1925: pi. 9, fig. 1, 2), or
as connected with each other by a pair of transverse bars which unite in the median line and extend thence forward
as a single member to the point of union with the ventral cartilage (Pavesi, Ann. Mus. Stor. nat. Genoa, 6, 1874:
pi. 2, fig. I, 2).

2. For reasons why Cetorhinus is retained for this genus rather than Halsydrus Fleming, 1809, see footnote 4,
p. 146.

3. Type designated by Jordan (Genera Fish., i, 1917: 95) as C. gunneri Blainville, 1816, which was a substitution
for Squalus gunnerianus Blainville, 1810.

4. Whitley (Mem. Qd. Mus., 10, 1934: 196), followed by Fowler (Bull. U.S. nat. Mus., 100 [t^], 1940: 112), has
replaced the generic name Cetorhinus with Halsydrus on the ground that the carcass of the Orkney animal, for
which the latter was proposed, was actually that of a very large Basking Shark, as is certainly suggested by pub-

Fishes of the Western North Atlantic 147

Generic Characters. Those of the family.

Range. Temperate belts of North and South Atlantic including the Mediterranean,
North and South Pacific and southern Indian Ocean.

Fossil Gill Rakers. Oligocene to Pliocene, Europe.

Species. Cetorhinus had long been thought to be monotypic, but Whitley" has re-
cently discussed its Australian representative under a name maccoyi Barrett," distinct
from that of its northern Atlantic representative niaxitnus. Comparison of Whitley's
photographs of an Australian specimen 25 feet long with a Massachusetts Bay specimen
of about the same size, and pictured below, suggests that a longer caudal and perhaps a
higher first dorsal may prove diagnostic for the former. If correct, this opens the whole
question of the specific relationship of the Basking Sharks of the western and eastern South
Atlantic' and of the northern and southeastern Pacific to the North Atlantic form. The
discontinuity of geographic distribution suggests that actually the genus may include
several species instead of one only. But definite decision must await critical comparison of
specimens from diflterent seas, or at least of comparable measurements and photographs.

Cetorhinus maximus (Gunnerus), 1765

Basking Shark, Bone Shark

Figures 23, 24

Study Material. Mounted specimens, about 26 feet 6 inches long (New Eng. Mus.
Nat. Hist.) and about 14 feet 6 inches, from New Jersey (Amer. Mus. Nat. Hist.) ; head
of a 12-foot specimen from Fire Island, New Yorkj gills and gill rakers of another from
same locality (Amer. Mus. Nat. Hist.)} excellent photographs of a specimen about 15
feet long taken off Jones Inlet, New York, June 20, 1941.*

Distinctive Characters. The combination of lunate caudal, enormously long gill
openings, long rakers on the gill arches, very many minute teeth, and nostrils widely sepa-
rated from mouth, sets Cetorhinus apart from all other sharks.

Description. Proportional dimensions in per cent of total length. Female, 4,400 mm.
(4,318 mm. between perpendiculars) from about 15 miles S by E of Long Branch, New
Jersey.^

lished illustrations of its cranium, vertebrae and pelvic skeleton (Barclay, Mem. Werner. Soc, i, 1811: 418).
But we agree with Norman ("Discovery" Rep., 16 [2], 1937: 7, footnote 2) that nothing would be gained by
abandoning a name as old and as generally used as Cetorhinus, at least until some modern student establishes, by
personal examination of the remains in question (if they are still in existence) , that they actually are those of a
Basking Shark and not of some other very large species.

5. Fish. Aust., /, 1940: 132. 6. Sun-Nature Book, Pt. IV, Water Life, 1933: 13.

7. Norman ("Discovery" Rep., 16, 1937: 7) had already suggested that the Falkland Island form may be distinct
from the northern. For a recent description of the South African form, see Barnard (Ann. S. Afr. Mus., j2 [2],

>937:43)-
i. Received from New York Herald Tribune.
9. This specimen is mounted in the American Museum of Natural History, New York, and the above proportions

are based on measurements made by E. W. Gudger at the time of its capture. Measurements of body, fins and

gills were made on the curvature.

148

Memoir Sears Foundation for Marine Research

Snout length: to angle of jaw, in straight line 1 6. i.

Eye: horizontal diameter i.O.

Gill opening lengths: ist 25.75 2nd 23. i; 3rd 20.0; 4th 17.1; 5th 14.6.

First dorsal fin: height 9.8 ; length of base 9.8.

Caudal fin: upper margin 22.3; lower anterior margin 13.9.

Pectoral fin: length 17.6.

Distance from snout to: ist dorsal 36.3; 2nd dorsal 65.3; upper caudal 77.6;

pectoral 27.7; pelvics 55.6.

Distance from origin to origin of: ist and 2nd dorsals 29.1 ; 2nd dorsal and caudal

13.0; pectoral and pelvics 28.5; pectoral and anal 45.O.

Trunk fusiform, stoutest from shoulders to ist dorsal, tapering rearward to moder-
ately stout caudal peduncle, the latter somewhat flattened dorso-ventrally, with strongly
developed lateral keel on either side originating opposite the tip of anal and extending
well out on caudal fin. Well developed precaudal pits both above and below, in the form of
lunate furrows. Dermal denticles small, but of various sizes, in patches or stripes with

Figure 23. Cetorhinus maximus. Drawing based on adult female, 26 feet 6 inches long, mounted in New
England Mus. Nat. Hist., and on other available information. A, B Head of 12-foot specimen from Fire
Island, New York (Amer. Mus. Nat. Hist.). C Sector of upper jaw of same, about 1.4 x. Z) Lateral view of
two teeth, about 4 x. £ Dermal denticles from back, above origins of pectorals, about 8 x. F Left-hand nostril,
about natural size.

Fishes of the Western North Atlantic

149

bare spaces between/" erect, close-set, thorn-like, with recurved tips having a median ridge
along the anterior face, their bases large and corrugated.

Figure 24. Cetorhinus maximus, from Long Island, New York (Amer. Mus. Nat. Hist.). A Gill folds and
gill rakers of one of the gill-arches, about V^ x. S Four of the gill rakers of same, with bases of the gill folds,
about 2 X.

10. Radcliffe (Bull. U.S. Bur. Fish., 34, 1916: 248) gives an excellent illustration from a North Pacific specimen.

150 Memoir Sears Foundation jar Marine Research

Head slightly compressed laterally opposite mouth (strongly so in small specimens).
Snout very short, subconical, with rounded tip in larger specimens but relatively much
longer in small ones, forming a subcylindrical proboscis, obliquely truncate in front,
terminating dorsally in a sharp point, with many circular pores on its dorsal surface; transi-
tion from the juvenile to the adult form takes place at lengths of 1 2 to 1 6 feet. Eyes nearly
circular without nictitating membrane or subocular folds, their diameters only about Vs
as great as distance between them opposite, or a little posterior to, front of mouth. Spiracles
described as minute, circular, a little posterior to angles of jaws or opposite latter. Gill
openings very large, extending from upper sides down onto lower surface of throat, the
1st longest, the 5th shortest, the ist pair separated below by 6 inches only, the 2nd pair
by 9 inches, the 4th pair by 21 inches and 5th pair by 27 inches in a specimen" 30 feet 3
inches long. Gill rakers very numerous (about 1,260 on gill studied), flattened basally on
the adjacent sides but bristle-like toward the tips, in a continuous series, and directed in-
ward; I series on the ist gill arch, 2 series on the 2nd, 3rd and 4th and only i series
described for the 5th." Nostrils wide apart at outer edges of snout, small, transverse, con-
siderably nearer to mouth than to tip of snout in young specimens, less so in adults because
of decrease in relative length of snout, their anterior margins slightly expanded in sub-
triangular outline. Mouth very large, occupying most of breadth of head, rounded in
adult but varying in young from nearly transverse, with corresponding lateral expansion
of sides of head behind the eyes, to broadly V-shaped; these variations probably associated
with wide distensibility of mouth and loose articulation at symphyses. A very short labial
furrow at corner of mouth on lower jaw, but none on upper.

Teeth minute, being only about 3 mm. high in specimen about 12 feet 9 inches (3,900
mm.) long and about 6 mm. in one of 30 feet; in 4 to 7 functional series, with lOO or more
teeth in each row on each side of jaw; those toward center of mouth low and triangular,
but those along the sides conical, slightly recurved, somewhat compressed laterally, with a
ridge on each side, the basal part striated ; a wide space with only scattered teeth at center
of upper jaw (106 mm. wide in 12-foot specimen) but not of lower jaw.

First dorsal fin an approximately equilateral triangle, its anterior margin nearly
straight, its posterior margin slightly concave or even slightly convex in some cases, its
apex subacute, its free rear corner extending only a slight distance beyond the rear termina-
tion of its base, the height along anterior margin varying from about 1 1 to 14% of total
length, its origin considerably behind the inner corner of the pectoral when latter is laid
back; the midpoint of its base about midway between tip of snout and fork of caudal.
Second dorsal's anterior margin only ^/4 to Vs as high as ist, with rounded apex, strongly
concave rear margin and free rear tip about as long as its base. Caudal ^^ to Vs of total
length, lunate, its axis steeply raised as in Isuridae, its posterior outline obtusely sub-
angular rather than rounded, with well marked subterminal notch, its lower anterior

11. Storer, Mem. Amer. Acad. Arts Sci., (2) p, 1867: 229.

12. See White (Bull. Amer. Mus. nat. Hist., 7^, 1937 : pis. 7, 8) for photographs of gill rakers in position.

Fishes of the Western North Atlantic 151

margin (lower lobe) about 60-65% ^s long as upper, each measured from the precaudal
pit, its tips subacute. Anal similar to 2nd dorsal, and about as large, its origin under rear
part of base of latter. Pelvics about % as high as ist dorsal along anterior margin. Claspers
described as about 3 feet 3 inches long in 30-foot male. Pectoral with straight or slightly
concave distal margin and blunt tip, but broadly rounded inner corners, relatively smaller
than in Isurus, the length along anterior margin being only about Vs of distance from
snout to origin of caudal.

Color. Grayish-brown to slaty gray, or nearly black above. The under parts may be
either uniformly of the same color as the back, of a paler shade of the same, or grading
into white, sometimes with a triangular white patch under the snout and with two pale
bands along the ventral surface on either side of the midline or otherwise marked with
white, there being a wide variation in this respect.

Size. The Basking Shark rivals, although it does not equal, the Whale Shark (p.
192) in size. It has been credited repeatedly with reaching a maximum length of 40 to
50 feet. For Basking Sharks to reach lengths of 35 to 40 feet is not exceptional, for one
of about 45 feet and three of about 40 feet, as well as smaller ones, were taken on the
Norwegian coast during the period 1884 to 1905.^^ The six next longest actually meas-
ured were 36 feetj 32 feet 2 inches; 32 feet; 31 feet; 30 feet 6 inches; and 30 feet 3
inches. The four largest, for which we find exact measurements for the western Atlantic,
were 32 feet 2 inches, 32 feet, 30 feet 3 inches, and 26 feet 6 inches, although others up to
40 feet have been reported without supporting evidence. Similarly, the longest of 21
Basking Sharks landed in Monterey, California, from November to February of 193 1, was
about 28 feet; the largest ever sold to the particular fishery firm in question was a few
inches less than 30 feet."

The smallest free-living specimens of which we find record were of 5 feet 5 inches,"
8 feet 4 inches,'" and about 8 feet 6 inches (2.6 m.)," which suggests that Basking Sharks
are as a rule at least 5 to 6 feet long at birth. Males mature at a length of perhaps 1 5 to 20
feet as indicated by the presence of small claspers in specimens up to about 1 1 feet, with
very large ones in specimens of 25 to 26 feet or longer.'* Similarly, most described speci-
mens of less than 1 1 to 13 feet have had the immature, proboscis-like form of snout. On
the other hand, a 14-foot 3-inch specimen taken recently near New York showed an inter-
mediate state,'^ and all specimens of 20 feet or upward, for which adequate information
is at hand, have been of adult conformation in this respect.

We have not succeeded in finding precise weights for the larger sizes in the Atlantic.
But since the two Monterey specimens just quoted actually weighed 6,580 pounds at 28
feet and 8,600 pounds at about 30 feet, this no doubt is a fair indication of the weight of

13. Collett, Norges Fiske, j, 1905: 83-86. 14. McGinitie, Science, N.S. 75, 1931 : 496.

15. Thompson, Nat. Hist. Ireland, 4, 1856: 253. 16. Pengelly, Zoologist, (3) 5, 1881: 337.

17. Nobre, Fauna Marinha Port. Vert., /, 1935: 441.

18. Pavesi, Ann. Mus. Stor. nat. Genoa, 12, 1878: 398, 406.

19. Gudger, J. Morph., 57, 1935 : 96.

152 Memoir Sears Foundation for Marine Research

the Atlantic specimens, there being no reason to suppose that Atlantic specimens would
differ very widely from those of the Pacific. Estimated weights of smaller specimens 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.^"

Developnental Stages. Developmental stages have not been described, except as
noted (p. 152).

Habits. Basking Sharks are sluggish and inoffensive fish. When in coastwise waters
they spend much time lying at the surface with backs awash, their dorsal fins standing high
above the water with tip of snout and caudal showing; or they swim slowly, with mouth
open gathering their diet of plankton. They are also described as sometimes lying on their
backs sunning their bellies. They are so little disturbed by boats that it is easy to approach
them closely; in fact, excellent moving pictures of them have been taken off Ireland.^'
However, on occasion they are reported as jumping, perhaps in an attempt to shake off
remoras or parasites. They often gather in schools of up to 60 or 100 individuals and
there are reports of two or three swimming tandem.

In the Gulf of Maine and off the middle Atlantic coast of the United States, as well
as in the northern part of their range in the opposite side of the Atlantic, Basking Sharks
appear almost exclusively during the warm half of the year,^^ and the early accounts sug-
gest some movement northward during the summer in northern European waters. The
winter habitat of the northern species is not known for either side of the Atlantic, although
lack of evidence of any increase in abundance to the southward suggests that they simply
retire in the fall and winter to deeper water."^ If so, the scarcity of animal plankton that
prevails generally in boreal seas during winter must result in very poor feeding for them,
suggesting that they are generally inactive at that time, perhaps lying on or close to
bottom.

The only definite information as to breeding habits is the report, more than a century
and a half old, that an embryo about one foot long was taken from the mother." It is not
known at what season the young are born, for while it has been stated that their habit of
schooling is associated with breeding, this seems more likely connected with their pursuit
of planktonic food. However, it seems certain that young are produced throughout their
entire range, for small ones have been reported both from the north (Ireland, Norway)
and from the south (Mediterranean).

The diet of the Basking Shark consists wholly of small planktonic organisms which
it sifts out of the water by means of its gill rakers, as do such plankton feeders as some
dupeoids, anchovies and whalebone whales. Usually the stomach contents are simply a

20. An estimated weight of about 3,000 pounds for one between 12 and 14 feet long was probably far too high.

21. In the widely popular film, "Men of Arran."

22. A skeleton found on the beach near Provincetown, Massachusetts, in January 1939, may have been there for
months.

23. On this, see Lijbbert and Ehrenbaum (Handb. Seefisch. Nordeurop., 2, 1936: 281).

24. Pennant, Brit. Zool., 5, 1776: loi.

Fishes of the Western North Atlantic 153

soupy or gelatinous mass. On several occasions, however, this has been found to consist
chiefly of minute Crustacea, this being true of the only western Atlantic specimen whose
stomach contents has been recorded.^'

Abundance. The published records show that there is much variation in the number
of Basking Sharks in the centers of abundance over periods of years. For example, great
schools were seen during the summer of 1776 and for a few succeeding summers off the
coast of Wales,^* but no comparable numbers have ever been reported there subsequently.
Similarly, along the Norwegian coast, where Basking Sharks formerly supported an inter-
mittent fishery, a paucity in the first half of the 1 8th century and again around 1 840 alter-
nated with a comparative abundance around 1800 and 1880; since then only occasional
specimens have been reported yearly from one Norwegian locality or another." Similar
fluctuations are also reported for the western Atlantic, but with less definite evidence (see

P- 155).

Basking Sharks Reported as Sea Serpents or Other Monsters. Without entering
into the controversy regarding the so-called "sea serpent," we may point out that the
Basking Shark has formed the demonstrable basis of sea serpent stories on several occa-
sions; "as the carcase 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, by far the bulkiest part of the
head skeleton, as well as the pectoral and pelvic fins, is soon washed away, leaving only the
backbone and the somewhat curiously shaped box-like cranium."'* As a recent instance of
this nature we may point to the wide publicity given by the press and radio to a supposed sea
serpent whose identity was based upon the skeleton of a Basking Shark about 25 feet long
that was stranded on the beach at the tip of Cape Cod near Provincetown, and which we
were able to examine.^" Also, it has been suggested repeatedly that the dorsal and caudal
fins of Basking Sharks, swimming in line at the surface, have been the basis for stories of
at least some of the reported sea serpents or other supposed monsters, especially in north-
ern Scandinavian waters.

Relation to Man. The livers of medium-sized to large Basking Sharks will yield any-
where from about 80 to 200 gallons of oil, and occasionally as much as 400 gallons, with
a maximum reported yield of 600 gallons.'" As the oil is considered nearly or quite equal
to sperm oil for use In lamps, it was readily saleable up to the time when animal oils were
replaced by petroleum products for lighting. For example, the oil of a specimen taken at
Provincetown in 1836 or 1837 yielded Its captor $ 103 ; even as far back as the last part of
the eighteenth century a large one in British waters was said to be worth the equivalent of

25. Hussakoff, Copeia, 21, 1915: 25. 26. Pennant, Brit. Zool., 5, 1776: 102.

27. Collett (Norg-es Fiske, 3, 1905: 83) lists about 25 records for the period 1S81-1905. See also Lijbbert and
Ehrenbaum (Handb. Seefisch. Nordeurop., 2, 1936: 281) for general summary of fluctuations.

28. Norman and Fraser, Giant Fishes, 1937: 21.

29. For detailed account, with photograph, see Schroeder (New Engl. Natural., 2, 1939: i).

30. Stevenson (Rep. U.S. Comm. Fish. [1902], 1904: 227); many of the older records of yield are expressed in
"barrels" of unknown volume.

154 Memoir Sears Foundation for Marine Research

20 pounds sterling." Their oil and sluggish nature made the Basking Shark the object of
intermittent small-boat fisheries with harpoon wherever and whenever they appeared in
any numbers, especially in Irish and Norwegian waters and around Iceland. Similarly, the
Pacific Basking Shark has supported, and probably still does, a local fishery of small boats,
each manned by six or eight men, off the coasts of Peru and Ecuador. Also, considerable
numbers are landed in California, where they are utilized for oil and fish meal.

The larger whaling vessels also pursued them in earlier days whenever encoun-
tered; for an instance of this in the Gulf of Maine, see p, 155. But it is now more than 100
years since Basking Sharks have been plentiful enough on the western side of the North
Atlantic for more than incidental capture. With its large yield one might wish that the liver
oil of the Basking Shark had a high vitamin content, but this appears not to be the case.

Range. Once thought to be an Arctic species, and often so characterized, the Basking
Shark is now known to be an inhabitant of temperate and boreal waters. In the North
Atlantic its range is bounded on the north by a line extending from the eastern side of the
Gulf of Maine and Newfoundland to the western and southern coasts of Iceland, the
Orkneys, the Faroes and northward along western Norway to the North Cape, with
occasional reports of it from the Murman Coast. In general, this line marks the zone of
transition from the region of influence of Atlantic waters to those of Arctic waters."

To the southward in the eastern side of the Atlantic it is reported occasionally from
the English Channel and the North Sea as far as the Skagerrak and Kattegat (never from
the Baltic) , along the coasts of France and the Iberian Peninsula, from Madeira, Morocco
and the Mediterranean. On the western side it is reported as far as North Carolina. At
present its chief centers of abundance appear to be west and south of Iceland, along western
Ireland, among the Orkneys, and off southwestern Norway. There is no evidence that it
occurs at all in the tropical Atlantic. However, it is represented on both sides of the South
Atlantic off South Africa, Argentina and the Falkland Islands, in the South Pacific off
Peru and Ecuador, off southern Australia and New Zealand, and in the northern Pacific
from California to British Columbia as well as in Japanese and Chinese waters, by a form
(or forms) whose precise relationship to the Basking Shark of the North Atlantic is still
to be determined (p. 147).

Occurrence in the Western Atlantic. There is no reason to suppose that the Basking
Shark ever occurred, other than as a stray, north of about 44° to 45° N. in the western
North Atlantic, there being only four positive records of it from the southern part of
Newfoundland: one from the outer coast of Nova Scotia, three from the Bay of Fundy
and a few from the vicinity of Eastport, Maine, at the mouth of that bay. In colonial
days the southern and western parts of the Gulf of Maine appear to have supported a con-
siderable population of them, however, for by old reports many were taken in Massachu-

31. Pennant, Brit. Zool., 5, 1776: 174. An estimate of 80 pounds sterling (Day, Fish. Gt. Brit., 2, 1880-188+:
306) seems too high.

32. There is no recent report of it for any Arctic locality; nor does Jensen (Mindskrif. Japetus Steenstrup, 2 [3],
1 9 14.) include it in his survey of the sharks of Greenland.

Fishes of the Western North Atlantic 155

setts waters, especially off the tip of Cape Cod, in the first half of the eighteenth century,
the oil being then in demand for illuminating purposes. But the local stock soon went the
way of the Atlantic Right Whale in these same waters, i.e., into the try-pot.

The only positive records of them north of Cape Cod since 1840, of which we have
learned, are as follows :

1840, a number seen, and several captured, by a whaler off Cape Elizabeth, Maine.
1847, one killed near Provincetown at the tip of Cape Cod.
1 85 1, a large one reported as about 40 feet long captured at Musquash Harbor, New

Brunswick, near the mouth of the Bay of Fundy.
1 864, one harpooned but lost in Massachusetts Bay.
1 868-1 870, several, 25 to 35 feet, killed near Eastport, Maine, at the mouth of the Bay

of Fundy.
1876, one stranded in Conception Bay, Newfoundland.

1908, one about 18 feet taken near Provincetown, Mass., in a weir.

1909, one about 22 feet in Provincetown Harbor.
19 1 3, one about 29 feet, Provincetown Harbor.
1925, one about 30 feet off Portland, Maine.'^
193 1, female, 12V2 feet, at York Harbor, Maine.

1934 0) three records from Newfoundland at Petty Harbor, the vicinity of St. John

and at Placentiaj the last one 32 feet long.
1936, two specimens ofiF Portland, Maine, the first a small one about 20 feet long and 550

pounds dressed, taken about May ist, the second a large specimen reported to have

been about 40 feet, taken August 2nd.

1938, one washed ashore near French Village, Halifax County, Nova Scotia, of which we
received a clearly recognizable photograph.

1939, January, a skeleton washed ashore near Provincetown and reported as a Sea Ser-
pent (see p. 153).

Unknown date, a 31 -foot specimen taken at Long Point, near Provincetown, Mass.

The hiatus in the foregoing list between 1876 and 1908 probably reflects the fact
that fishes generally, and especially sharks, in the Gulf of Maine received little scientific
attention during that period. But this large shark is probably no more plentiful now than
the paucity of the recent record suggests, for, so great has been the popular interest in
sharks of late, and so wide the newspaper publicity given to unusual captures, that any
large specimen is almost certain to be reported sooner or later — even if not captured — in
such frequented and hard-fished waters as those of the coastwise belt of the Gulf of Maine.

Near Woods Hole, a few miles west of Cape Cod, an incursion by Basking Sharks ap-
pears to have taken place in the summer of 1878, when at least twenty were found dead
in the local fish traps. However, only occasional specimens have been reported more re-
cently, e.g., one of 26 feet 6 inches (see Study Material, p. 147) taken at Martha's

33. Personal communication from Walter H. Rich.

156 Memoir Sears Foundation for Marine Research

Vineyard, June 24, 1920, and another of 20 feet 2 inches, stranded in the landlocked
waters of a small harbor (Hadley's) on Naushon Island in July 1937. There is nothing
in the published record to suggest that the zone of most frequent occurrence ever extended
much farther west or south than this along the North American coast, there being occa-
sional records only for Long Island," one or two near New York (one in New York Har-
bor many years ago) and about six for the coast of New Jersey, with one probable and one
positive record for North Carolina." The only reports of Basking Sharks farther south in
the western Atlantic are for northern Argentina*' and the Falkland Islands,*' which may be
distinct from those of the North Atlantic (p. 147).

Synonyms and References:

North and South Atlantic and South Africa:"

Squalus maximus Gunnerus, K. norske Vidensk.-Selsk. Skr. Trondh., 1765: 33, pi. 2 (type locality, Trond-
hjem, Norway); Drontheim Gesellsch. Schr. Leipzig, 5, 1767: 28, pi. 2 (German translation of the
foregoing); Linnaeus, Syst. Nat., 12th ed. 1766: 400 (descr.) ; Gunnerus, K. norske Gesellsch. Wiss.
Skr. Kbh. u. Leipzig, 4, 1770: 13, pi. 4 (descr., Norway) ; Olafsen and Povclsen, Reyse en Island, 1772:
988 (Iceland, not seen); Reise durch Island (German translation), 2, 1774: 204 (South Iceland); Pen-
nant, Brit. Zool., 5, 1776: lOl (Isle of Arran, embryo, fishery); Olavius, Oecon. Reyse Island, 1780:
80 (Iceland, not seen); Mohr, Fors0g. Island. Naturh., 1786: 60 (Iceland); Bonnaterre, Tabl. Encyc.
Meth. Ichthyol., 1788: 10 (descr.); Gmelin, Syst. Nat., i, 1789: 1498 (descr.); Bloch and Schneider,
Syst. Ichthyol., 1801: 134 (descr.); Olafsen and Povelsen, Voy. en Island, 5, 1802: 278 (Iceland);
Lehmann, Neue Schr. Naturf . Freunde, Berlin,.^, 1 803: 1 20; Bosc, Nouv. Diet. Hist, nat., 2/, 1 803: 185
(diagn.); Latreille, Nouv. Diet. Hist, nat., 24, 1 804: 72 (in table of contents); Turton, Brit. Fauna,
1807: 113 (Gt. Brit.); Home, Philos. Trans., 2, 1809: 206 (descr., anat.) ; J. Physique, 7/, 1810: 241
(descr., anat.); Philos. Trans., 2 (2), 1813: 227, pi. 6 (descr., anat.); Mitchill, Trans. Lit. Phil. Soc.
N. Y., I, 1815: 486 (fishery near Provincetown) ; Couch, Trans. Linn. Soc. Lond., Zool., 14, 1 825: 91
(Cornwall, size); Vrolik, Bijdr. Natuurk. Wetensch. Amsterdam, r, 1826: 305 (Holland); Fleming,
Hist. Brit. Anim., 1828: 164 (descr., Gt. Brit.); Vrolik, Z. Organ. Physik, 2, 1828: 490 (Holland);
Faber, Fische Islands, 1829: 20 (distrib., habits); Nilsson, Prod. Ichthyol. Scand., 1832: 114 (Scandi-
navia); Agassiz, L., Poiss. Foss., j, 1835-1843: 87, pi. F, fig. 8, 8"; Yarrell, Brit. Fish., 1836: 366
(descr., Gt. Brit.) ; Couch, Cornish Fauna, 1838: 51 (size) ; Bennett, Narr. Whaling Voy., 2, 1840: 240
(Gt. Brit.); Owen, Odontogr., 1 840-1845: 27 (teeth); Linsley, Amer. J. Sci., 47, 1844: 77 (Long
Island Sound) ; Gaimard, Voy. Islands et Groenland, Zool. Med., 1 851: 163 (Iceland) ; Schlegel, Natuurl.
Hist. Nederland Dieren, 1862: 191, pL 19, fig. I (size); Brito Capello, J. Sci. math. phys. nat.
Lisboa, 2, 1870: 233 (Portugal); Gervais and Gervais, C. R. Acad. Sci. Paris, ^2, 1876: 1237 (descr.,
Concarneau) ; J. Zool., 5, 1876: 319 (descr., ill. of head of juv., gills, rakers, teeth, vertebrae, Concar-
neau, France); Gervais and Boulart, Poiss., 5, 1877: 190, pi. 73 (descr.); Buckland, Hist. Brit. Fish.,
1881: 215 (Gt. Brit.); Pengelly, Zoologist, (3) 5, 1 891: 337 (Devonshire).

Basking Shark, Pennant, Brit. Zool., 5, 1769: 38, 342 (descr., distrib., embryo); Shaw, Gen. Zool., 5 (2),
1804: 327, pi. 149, 150 (general); Pennant, Brit. Zool., $, 1812: 134 (addit. record); Low, Fauna
Orcadensis, 1813: 173 (Orkneys, fishery); Brabazon, Deep Sea Coast Fish. Ireland, 1848: frontispiece,
48 (fishery); Couch, Trans, nat. Hist. Soc. Penzance, 1864: 234 (not seen); Fish. Brit. Isles, 1862:
60, pi. 14 (not seen); i, 1867: 60, pi. 14 (descr., habits, Gt. Brit.); Cornish, Zoologist, (2) 5, 1870:

34. Most recently a 12-foot specimen taken off Fire Island, July 1944 (see Study Material, p. 147).

35. Coles, Proc. biol. Soc. Wash., 78, 1915: 92; Brimley, J. Elisha Mitchell sci. Soc, 57, 1935: 311.

36. Lahille, An. Mus. nac. B. AireS; j^, 1928: 325; Pozzi and Bordale, An. Soc. cient. argent., /so, 1935: 150.

37. Norman, "Discovery" Rep., 16 (2), 1937: 143.

38. References for the South Atlantic and South Africa are included for convenience.

Fishes of the Western North Atlantic 157

2253 (descr., meas., Cornwall); Allman, Nature, Lond., 14, 1876: 368 (gill rakers, food); Harvey,
Nature, Lend., 75, 1877: 273 (Newfoundland) ; Stevenson, Rep. U.S. Comm. Fish. (1902), 1904: 227
(fishery, yield of oil) ; Grew, lllustr. London News, Aug. 1 1, 1945: 166 (general account).

Le Trcs Grand, Broussonet, Mem. Math. Phys. Acad. Sci. Paris, 1780: 669; Daubenton, Encyc. Method.
Hist. nat. Paris, 5, 1 787: 96 (teeth).

Le Squale Tres-Grand, Lacepede, Hist. nat. Poiss., 4° ed., /, 1798: 165, 2og, in Buffon, Hist. Nat. (descr.,
size) ; in Sonnini, Hist. Nat. Poiss., 5, 1801-1802: 400 (general).

Squalus gunnerianus Blainville, J. Physique, 1810: 256, pi. 2, fig. 3 (descr.).

Squalus homianus Blainville, J. Physique, 1810: 257, pi. 2, fig. I (descr.).

Squalus felegrinus Blainville, J. Physique, 18 10: 256, pi. 2, fig. 2 (descr.).

Squalus feregrinus Blainville, Bull. Soc. philom. Paris, 2, 181 1 : 365 (size); Ann. Mus. Hist. nat. Paris, 18,
181 1 : 132 (discuss.).

Squale pelerin Blainville, Ann. Mus. Hist. nat. Paris, 18, 1 811: 88, pi. 6 (detailed descr., ill. of 29-ft. 4-'m.
male, is the one most often copied).

Cetorhinus gunneri Blainville, Bull. Soc. philom. Paris, 1816: 121 (name).

Cetorhinus komianus Blainville, Bull. Soc. philom. Paris, 1816: 121 (name).

Cetorhinus feregrinus Blainville, Bull. Soc. philom. Paris, i8l6: 121 (name).

Cetorhinus shavianus Blainville, Bull. Soc. philom. Paris, 1816: 121 (name); J. Physique, 1816; 264 (not
seen).

SeUche maxima^" Cuvier, Regne Anim., 2, 1817: 129 (general) ; Regne Anim., 2nd ed., 2, 1829: 391 (gen-
eral) ; Cloquet, Diet. Sci. Nat., ^S, 1825: 308 (general) ; Bory de St. Vincent, Diet. Class. Hist. Nat., 15,
1829: 597 (general) ; Jenyns, Man. Brit. Vert. Anim., 1835: 503 (descr., Gt. Brit.) ; Richardson, Fauna
Boreal. Amer., 5, 1836: 29; Parnell, Mem. Werner. Soc. Edinb., 7, 1838: 418 (Scot.); Bonaparte,
Mem. Soc. neuchatel. Sci. Nat., 2 (8), 1839: 9 (in synopsis) ; Owen, Odontogr., 1840-1845: 27 (descr.
of teeth of 36-ft. spec.) ; Miiller and Henle, Plagiost., 1841 : 71 (descr., distrib.) ; Cuvier, Regne Anim.,
ill. ed. Poiss., 1843: 363, pi. 115, fig. 2 (good ill. of teeth); Owen, Cat. Osteol. Roy. Coll. Surg., i,
1853: 97 (brief descr., vertebrae, refs.) ; Traill, Proc. roy. Soc. Edinb., j, 1854: 210 (argues "animal
of Stronsa" not a Basking Shark) ; Van der Hocvcn, Handb. Dierkunde, 2nd ed., 2, 1855: 261 (general) ;
Nilsson, Skand. Fauna Fisk., 1855: 720 (Scandinavia); Thompson, Nat. Hist. Ireland, 4, 1856: 253
(Ireland, fishery); Nardo, Atti 1st. veneto, (3) 5, 1 859-1 860: 787 (Medit.) ; Fitzinger, Bild. Atlas
Naturg. Fische, 1864: fig. 173 (ill.); Malmgren, Arch. Naturgesch., jo (i), 1864: 346 (north. Nor-
way); Dumeril, Hist. Nat. Poiss., /, 1865: 413, pi. 3, fig. 18 (descr., size, ill., teeth); Bocage and Brito
Capello, Poiss. Plagiost., 1866: 14 (Portugal) ; Malmgren, Oefvers. Vetensk. Akad. Forh. Stockholm, 24,
1867: 264 (Norway) ; Brito Capello, J. Sci. math. phys. nat. Lisboa, 2, 1870: 140 (Portugal) ; Giinther,
Cat. Fish. Brit. Mus., 8, 1870: 394, 518 (descr., distrib., synonyms) ; Ninni, An. Soc. Nat. Modena, 5,
1870: 66 (Medit.) ; Van Beneden, Poiss. Cotes Belg. Parasit., 1870: 70 (parasites, Belgium), also in Mem.
Acad. R. Belg. CI. Sci., ^8 (4), 1871: 7; Canestrini, in Cornalia, et al., Fauna d'ltal., 1 871-1872: 44
(Medit.); Pavesi, Ann. Mus. Stor. nat. Geneva, 6, 1874: 36, pi. 1—3 (descr.; ill. juv.; anat., Medit.);
Collett, Norges Fisk., 1875: 209 (Norway); Trois, Atti 1st. veneto, (5) / (6), 1875: 612 (Medit.);
Wright, Nature, Lond., 14, 1876: 31 3 (gill rakers, diet, value of oil, Ireland) ; Giglioli, Nature, Lond.,
15, 1877: 273 (name) ; Hasse, Morph. Jb., Suppl. 4, 1878: 43, pi. 3, 4, fig. I (anat.) ; Pavesi, Ann. Mus.
Stor. nat. Genova, 12, 1878: 416, pi. 3 (meas., ill. of young male, anat., discus.) ; Doderlein, Atti Accad.
Palermo, 6, 1 878-1 879: 30 (Medit.); Stossich, Boll. Soc. adriat. Sci. nat., 5, 1880: 68 (Adriatic);
Turner, J. Anat., Lond., 14, 1 880: 273, pi. 12 (teeth, gill rakers) ; Day, Fish. Gt. Brit., 2, 1 880-1884:
303, pi. 158, fig. I (descr., Gt. Brit.); Doderlein, Man. Ittiol. Medit., 2, 1881: 70 (Medit.);*" Mo-
reau. Hist. Nat. Poiss. France, i, 1881 : 305 (general) ; Hasse, Naturl. Syst. Elasm. besond. Theil, 1882:
236, pi. 32, fig. 1-5 (vertebrae); Mela, Vert. Fennica, 1882: 365 (Murman coast, not seen); Vieira,
Ann. Sci. nat. Porto, 7, 1894: 137 (Portugal); Brandt, Biol. Zbl., 18, 1898: 257 (bristles described
on snout probably were dermal denticles); Saemundsson, Vidensk. Medd. naturh. Foren. Kbh., 1899:

39. Also variously spelled "maximus" or "maximum."

40. For additional Mediterranean citations in publications not accessible to us, see Doderlein (above) and Cascia
(Bull. 1st. zool. Palermo, 2, 1935: 173).

158 Memoir Sears Foundation for Marine Research

420 (Iceland); Facciola, Revist. Ital. Pesc. Aquic, 75 (4), 1900: 40 (Stiait of Messina, meas.) ; Sae-
mundsson, Vidensk. Medd. naturh. Foren. Kbh., 1 903: 51 (Iceland); Carazzi, Zool. Anz., 28, 1904:
161 (photo of head of juv., meas., Sardinia) ; Steuer, Verh. zool.-bot. Ges. Wien, §§, 1905: 275 (gill
rakers); Carruccio, Boll. Soc. zool. Ital., (2) 7, 1906: 191 (Medit., not seen); Southwell, Zoologist,
(4) 10, 1906: 355 (a large one, Loch Broom, Scot.); Hendricks, Z. wiss. Zool., 97, 1908: 427,
pi. 18, 19 (anat., histol., gill rakers); S. B. naturh. Ver. preuss. Rheinl. Westf., 1909: 31 (gill
rakers); Osorio, Mem. Mus. Bocage Lisbon, i, 1909: 49 (food, Portugal); Saemundsson, Skr.
Komm. Havunders. Kbh., 1909: 113 (Iceland); Giinther, Encycl. Brit., nth ed., 24, 1911: 808
(general); Seabra, Poiss. Port., 1911: 194 (Portugal, not seen); Mourgue, Bull. Soc. linn. Provence, 2,
'913' 53 (descr. of juv., Marseilles); Senna, Monit. zool. ital., 24, 1913: 229 (Medit., size); Monit.
zool. ital., 31, 1920: 35, pi. 3 (Medit., size, brain); Valle, Congr. Soc. Pesca Mar. Trieste, 28, 1922:
31 (ill., Adriatic, not seen) ; Vinciguerra, Ann. Mus. Stor. nat. Genova, 51, 1923: 133 (Medit., ill. head
of juv., clasper, stomach contents); Monit. zool. ital., 55, 1923: 36 (off Genoa); Jenkins, Fish. Brit.
Isles, 1925: 314 (general); Saemundsson, Vidensk. Medd. naturh. Foren. Kbh., 84, 1927: 183, pi. 5
(Iceland, photo, 27-ft. female) ; Ehrenbaum, in Grimpe and Wagler, Tierwelt N.- u. Ostsee, Lief. 7
12^ 1927: 16 (general); Faune Ichthyol. Nord Atlant., Cons. Explor. Mer, 1929: plate not numbered
(general); Gould, Case for Sea Serpent, 1930: 245, pi. 6 (hist, of "animal of Stronsa") ; Monterosso,
Atti Accad. gioenia, t8, 1931: 55, 4 pis. (proportions; photos of juv. and adult, Medit.); Belloc, Rev.
des Trav. Peches Marit., 7 (2), 1934: 140 (ill. after Day; Morocco); Noronha and Sarmento, Peixes
Madeira, 1934: no (Madeira, not seen) ; Nobre, Fauna Marinha Port. Vert., /, 1935: 440, pi. 58, fig.
183 (Portugal) ; Otto, Levende Nat., 41 (12), 1937: 373 (spiral valve, vertebrae, North Sea, not seen),

Squalus rostratus Macri, Atti Accad. Sci. fis. mat. Napoli, /, 1 819: 76, pi. 1, fig. 2 (Medit.).

Squalus hodus Macri, Atti Accad. Sci. fis. mat. Napoli, i, 1819: 76, pi. I, fig. I, pi. 2 (Medit.).

Zelanche maximus ]ixoY\,7jCi<^o%\, 4, 1822:452 (not seen).

Squalus elefhas Lesueur, J. Acad. nat. Sci. Philad., 2, 1822: 350, I pi. (off N. Jersey, size, descr.) ; Storer,
Mass. Zool. Bot. Surv., Rep. on Fish., Suppl., 1839: 407 (near Provincetown, Massachusetts); Boston J.
nat. Hist., j, 1 841: 270 (Provincetown).

Pelerin tres grand, Cloquet, Diet. Sci. Nat., Atlas Poiss., 1825: pi. 30 (ill.).

Squalus rhinoceros Mitchill, in 1828 newspaper, quoted by DeKay, Zool. N. Y., 4, 1842: 358 (coast of
Maine, not seen).

Selache elefhas Bory de St. Vincent, Diet. Class. Hist. Nat., 15, 1829: 597.

Selachus maximus Minding, Lehrb. Naturgesch. Fische, 1832: 52 (not seen); Swainson, Nat. Hist. Fish.
Amphib. Rept., 2, 1839: 314 (general); Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 506 (in
synopsis); Yarrell, Brit. Fish., 2nd ed., 1 841: 518 (general descr., Gt. Brit.); DeKay, Zool. N. Y., 4,
1 842 : 3 5 7, pi. 63, fig. 208 (name, N. York, descr., size) ; Hamilton, Brit. Fish., 2, 1 843 : 3 1 1 ; also sub-
sequent eds. (Gt. Brit.); Gosse, Nat. Hist. Fish., 1851: 306 (size); Baikie, Zoologist, 11, 1853: 3846
(Orkneys) ; Foulis, Proc. Boston Soc. nat. Hist., 4, 1853: 205 (descr., meas. 40-ft. spec, Bay of Fundy) ;
Storer, Proc. Boston Soc. nat. Hist., 4, 1853: 206 (name); DeKay, Rep. St. Cab. Nat. Hist. N. Y., 8,
1858: 64 (N. York) ; Storer, Mem. Amer. Acad. Arts Sci., N. S. 9 (i), 1867: 229, pi. 39, fig. 3; also as
Fishes Mass., 1867: 253, pi. 37, fig. 3 (descr., meas., ill., Massachusetts); Steenstrup, Overs, danske
Vidensk.-Selsk. Forh., 1873: 47, 65, pi. 2, and summary in French (descr., ill. of gill rakers); Lutken,
Vidensk. Medd. naturh. Foren. Kbh., 1880: 62 (Iceland); Cornish, Zoologist, (3) 9, 1885: 35 (Corn-
wall); Gatcombe, Zoologist, (3) 9, 1885: 266 (Cornwall); Carus, Prod. Faune Medit., 2, 1889-1893:
507*' (Medit.); Jungersen, Danish "Ingolf" exped., 2 (i), 1899: 39 (claspers, California); Damiani,
Monit. zool ital., 14, 1903: 351 (Medit.); Ariola, Atti Soc. ligust. Sci. nat. geogr., 24, 1913: 14
(Medit.).

Squalus (^Selache) maximus Voigt, in Cuvier, Tierreich, 2, 1832: 509 (descr.).

Squalus rashleighanus Couch, Cornish Fauna, i, 1838: 51 (Cornwall, not seen).

Cetorhinus maximus Gray, List. Fish. Brit. Mus. Chondropt., i, 1851: 61 ; Gill, Proc. Acad. nat. Sci. Philad.,
Addend., i86i: 60 (name); Abbott, in Cook, Geol. N. J., App. E, 1868: 828 (N. Jersey); Ver-

41. Spelled "maximum."

Fishes of the Western North Atlantic 159

rill, Bull. Essex Inst., 5, 1871: 6 (Eastport, Maine); Gill, Rep. U.S. Comm. Fish. (1871-72),
1873: 813; Gervais and Gervais, J. Zool., 5, 1876: pi. 13, fig. 1, 2 (ill., head of juv.); Goode and
Bean, Bull. Essex Inst. Salem, 11, 1879: 29 (Gulf of Maine); Jones, List Fish. N. S., 1879: 9 (Nova
Scotia); Proc. N. S. Inst. Sci., 1882: 95 (same .is Jones, 1879); Jordan and Gilbert, Bull. U.S.
nat. Mus., 16, 1883: 31, 875 (distrib.) ; Goode, Fish. Fish. Industr. U.S., I, 1884: 668, pi. 249 (gen-
eral); Jordan, Rep. U.S. Comm. Fish. (1885), 1887; 797 (distrib.); Nelson, Rep. State Geol. N. J.,
1890: 663 (off N. Jersey, teeth); Dean, Fishes Living and Extinct, 1895: 90 (general); Smitt,
Hist. Scand. Fish., 2, 1895: 1 143 (dcscr., habits, Norway); Jordan and Evermann, Rep. U.S. Comm.
Fish. (1895), 1896: 21 8 (distrib.) ; Bull. U.S. nat. Mus., 47 (l), 1896: 51 (descr., distrib.) ; Goode and
Bean, Smithson. Contr. Knowl., jo, 189;: 21 ; Mem. Harv. Mus. comp. Zool., 22, 1896: 21 (descr., dis-
trib.) ; Jordan and Evermann, Bull. U.S. nat. Mus., 47 (4), 1900: pi. 7, fig. 23 (ill.) ; Knipowich, Annu.
Mus. zool. Acad. St. Petersb., 2, 1897: I 54 (Murman coast) ; Bean, Rep. For. Comm. N. Y., 1901 : 380
(old N. Yori< record) ; Bull. N. Y. St. Mus., 60, Zool. 9, 1 903: 42 (general) ; de Braganza, Result. Invest.
Sci. "Amelia," 2, 1904: 56 (off Portugal) ; Collett, Norges Fisk., 5, 1 905: 79 (Norway, descr., meas., dis-
cus.); Jordan, Guide to Study Fish., j, 1905: 540 (general); Fowler, Rep. N. J. Mus. (190;), 1906:
57 (descr., N. Jersey) ; Proc. .Acad. nat. Sci. Philad., 60, 1908: 55 ; Kendall, Occ. Pap. Boston Soc. nat.
Hist., 7 (8), 1908: 8 (New England records); Berg, Faune Russle, Poiss., Mus. Zool. Acad. Imp. Sci.
St. Petersb., i, 1911: 56 (Russia); Roule, Bull. Inst, oceanogr. Monaco, 243, 191 2: 10 (Medit.) ; Gar-
man, Mem. Harv. Mus. comp. Zool., 36, 191 3: 39 (descr.); Halkett, Check List Fish. Canad., 191 3:
41 (distrib.); Nichols, Abstr. Linn. Soc. N. Y., 20-23, 1913: 91 (off N. York); Sumner, Osburn and
Cole, Bull. U.S. Bur. Fish., 5/ (2), 1913: 737 (Woods Hole) ; Coles, Proc. biol. Soc. Wash., 28, 1915:
92 (off C. Lookout); Gudger, Science, 42, 1915: 653 (southern hemisphere records); Hussakoff,
Copeia, 21, 1915: 25 (descr. of Long Island specimen) ; Scharff, Irish Nat., 24, 1915: 99 (abund. off
Ireland); Fowler, Copeia, 30, 1916: 36; Murphy and Nichols, Brooklyn Mus. Quart., 3, 1916: 156
(general) ; Nichols and Murphy, Brooklyn Mus. Sci. Bull., 3 {i), 1916: 28 (habits, food, fishery, occur.
near N. York) ; Radcliffe, Bull. U.S. Bur. Fish., 34, 191 6: 248 (denticles), pi. 38, fig. 4 (teeth) ; Smith,
J. Amer. Mus. nat. Hist., 16, 1916: 342 (size, habits); Fowler, Proc. biol. Soc. Wash., 33, 1920: 144
(off N. Jersey); Allen, Bull. Boston Soc. nat. Hist., 24, 1 921: 3 (occur, off New England, habits);
Fowler, Copeia, lOl, 1921: 89 (N. Jersey); Proc. Acad. nat. Sci. Philad., 74, 1922: 3 (N. Jersey);
Huntsman, Contr. Canad. Biol. (1921), 3, 1922: 56 (Bay of Fundy) ; Legendre, Bull. Soc. zool. Fr.,
48, 1923: 275 (north. France, size, food) ; Bull. Soc. zool. Fr., 4g, 1924: 322 (add. records, sizes, oil) ;
Barnard, Ann. S. Afr. Mus., 27 (l), 1925: 34 (descr., S. Afr.) ; Bigelow and Welsh, Bull. U.S. Bur.
Fish., 40 (l), 1925: 41 (descr., habits, Gulf of Maine); Senna, Arch. ital. Anat. Embriol., 22, 1925:
84, pi. 9, 10 (skull, branchial skelet.) ; Lahille, An. Mus. nac. B. Aires, 34, 1928: 325 (Argentina) ; Bar-
nard, Ann. S. Afr. Mus., 27 (2), 1927: pi. 2, fig. I, la (ill.) ; Nichols and Breder, Zoologica, N. Y., 9,
1927: 20 (Woods Hole, Martha's Vineyard, Long Island, descr.) ; Rey, Faune Iberica Feces, 7, 1928: 415
(descr., distrib.); Borri, Mem. Soc. tosc. Sci. nat. Proc. Verb., 38, 1929: 27 (notes, Medit., not seen);
Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 23 (general); Jordan, Manual of Vert. Anim. NE.
U.S., 1929: 13 (general) ; also earlier eds.; Navarro, Bol. Pesc. Madr., 14, 1929: 105 (Balearic Is., spelled
"Celhorinus," not seen); Desbott, Arch. Medit. Malta, 8, 1930: 5 (Medit., not seen); Fowler, Fish
Culturist, 9 (8), 1930: 115 (N. Jersey, not seen); Gudger, Science, 72, 1930: 341 (N. Jersey,
general); MacCoy, Bull. Boston Soc. nat. Hist., 61, 1933: 17 (Maine, meas., photos); Young and
Mazet, "Shark, Shark," 1933: 266 (general); Bertolini, R. C. Accad. Lincei., 18, 1933: 34; Boll. Soc.
Biol, sper., 9, 1934: 1270 (gill rakers) ; Bigelow and Schroeder, Canad. Atlant. Fauna, biol. Bd. Canad.,
72*, 1934: j6 (descr., distrib.); Borri, Mem. Soc. tosc. Sci. nat., 44, 1934: 93 (discus.); Petit, Terre
et La Vie Paris, 4, 1934: 337 (hist., not seen); Daniel, Elasmobranch Fishes, Univ. Calif. Press, 1934:
37> 38, 155, 268 (gill rakers, gills, eye); Piers, Proc. N. S. Inst. Sci., 18 (3), 1934: 203 (Bay of
Fundy) ; Brimley, J. Elisha Mitchell sci. Soc, 57 (2), 1935: 31 1 (N. Carolina) ; Cascia, Boll. 1st. zool.
Palermo, 2, 1935: 137 (descr., meas., Medit.) ;" Gudger, J. Morph., S7, I935'- 93 (anat.) ; Anonymous,

4Z. For additional citations for the Mediterranean in publications not accessible to us, see Cascia (1935) and Doder-
lein (Man. Ittiol. Medit., 2, i88i: 70).

i6o Memoir Sears Foundation for Marine Research

Newfoundland Dept. Nat. Res., Fish. Res. Lab., Ann. Rep., 2, 1935: 79, pi. 2 (Newfoundland, photos) ;
Pozzi and Bordale, An. Soc. cient. argent., 120, 1935: 150 (Argentina) ; Vladykov and McKenzie, Proc.
N. S. Inst. Sci., J 9 (l), 1935: 47 (general) ; Lubbert and Ehrenbaum, Handb. Seefisch. Nordeurop., 2,
1936: 280 (descr., distrib., fishery); Barnard, Ann. S. Afr. Mus., 32 (2), 1937: 43 (descr., meas., S.
Afr.); Norman, 'Discovery' Rep., 16 (2), 1937: 143 (Falkland Is.); Norman and Fraser, Giant Fishes,
1937: 20, fig. 14 (general) ; White, Bull. Amer. Mus. nat. Hist., 74, 1937: 64, 79, 1 14, pi. 4, fig. 5, pi.
7-9, pi. 29, fig. c, pi. 46, fig. b (anat., photo of gill rakers) ; Tortonese, Atti Soc. ital. Sci. nat., 77, 1938:
292 (Medit.) ; Babic, Zool. Anz., 127, 1939: 39 (anat. notes; ill. of denticles) ; Schroeder, New Engl.
Naturalist, 2, 1939: I (supposed sea serpent stranded on C. Cod, 1939, a Basking Shark); McKenzie,
Proc. N. S. Inst. Sci., 20 (2), 1940: 42 (Nova Scotian records; size; weights); Nybelin, Fauna Flora
Uppsala, 1940: 236 (Katteg.nt) ; Hildehrand, Copeia, 1941: 222 (N. Carolina, old records); Norris,
Plagiost. Hypophysis, 1941: 29, pi. 10, fig. 35-38 (brain).

Squalus cetaceous Gray, Cat. Fish. Coll. Descr. by L. T. Gronow, 1854: 6 (Norway).

Polyfrosofus rashleighanus GiU, Ann. N. Y. Lye, 7, 1862: 398 (name) ; Couch, Hist. Brit. Fish., t, 1867:
67, pi. 15, fig. I (descr., size, Cornwall) .

Polyfrosofus macer Couch, Hist. Brit. Fish., i, 1867: 68, pi. 15, fig. 2 (descr., young female, Plymouth,
England).

No name, Hannover, K. danske Vidensk.-Selsk. Skr., (5) 7, 1868: 491 (descr. and ill. of gill rakers in Copen-
hagen Mus., not identified).

Cetorhinus blairivillii Brito Capello, J. Sci. math. phys. nat., Lisboa, 2, 1870: 234, I pi. (descr., ill. of small
spec, Portugal).

Cetorhinus rostratus Cornish, Zoologist, 5, 1870: 2259 (Cornwall, descr., meas.).

Selacke rostrata Pavesi, Ann. Mus. Stor. nat. Genoa, 1874: 36 (descr. of young, anat., gill rakers, Medit.);
GIglioli, Nature, Lend., 15, 1877: 273 (discuss.); Doderlein, Atti Accad. Palermo, 6, 1878-1879: 30
(Medit.).

SelacAus fenrumti Coin'ish, Zoologist, (3) p, 1885: 351 (Cornwall).

Cetorhinus (Selache) maximus Bridge, Camb. nat. Hist., 7, 1904: 453 (general).

Cethorinus (Selache) maximus Escribano, Bol. Soc. esp. Hist, nat., 9, 1909: 340 (Medit., Morocco).

Halsydrus maximus Fowler, Bull. U.S. nat. Mus., lOO {13), 1941 : 1 1 3 (descr., synonymy, Australasian refs.) ;
Feces Peru, Mus. Hist. Nat. Javier Prado, 1 945: 12 (listed Peru).

Doubtful references:

Halsydrus fontoffidani Fleming, Scots. Mag. Edinb. Misc., 1809: 6; Edinb. Encycl., 1817: 713 (Stronsa,

Orkney Is.").
Tetroras angiova Rafinesque, Carrat. Gen. Nuov. Sicil., 1810: II (Sicily).
Not Cetorhinus maximus Poey, An. Soc. esp. Hist, nat., 5, 1876: 184 (Cuba, no doubt Rhincodon).

Family ALOPIIDAE

Characters. Two dorsal fins, the ist much shorter than caudal, the rear end of its
base anterior to origin of pelvicsj caudal nearly or quite V2 of total length, not lunate in
form, but its lower anterior corner expanded as a definite lobe, its axis raised but little/
inner margins of pelvics entirely separate, posterior to cloaca} caudal peduncle not de-
pressed dorso-ventrally, but moderately compressed laterally} a well marked precaudal
pit above, and sometimes below j sides of trunk anterior to anal without longitudinal

43. The animal of Stronsa, named Halsydrus fontoffidani by Fleming, seems to have been the partly decomposed
remains of a large Basking Shark; see footnote 4, p. 146.

I. The enormously elongate caudal is the most striking feature of the family.

Fishes of the Western North Atlantic i6i

dermal ridges; snout short, thick, fleshy, the jaws not greatly protrusible; 3rd to 5th
gill openings over origins of pectorals; gill arches without rakers and not interconnected
by a sieve of modified denticles; nostrils entirely separate from mouth, without barbels;
spiracles present; eyes without nictitating folds or membranes; each jaw with a labial fur-
row (or furrows) near corner; teeth small, blade-like, with i cusp; head and skull normal
in shape (not widely expanded); rostral cartilages 3, united at tip; radials of pectoral
nearly all borne on mesopterygium and on metapterygium. Development ovoviviparous;
the egg case, in early development, soft, thin, oval.'^
Genera. Only one genus, Alo-pias.

Genus ^/o/)Wj Rafinesque, 1810
Thresher Sharks

Alofias Rafinesque, Carratt. Gen. Nuov. Sicil., 1 8 10: 12; type species, A. macrourus Rafinesque, 18 10, Sicily,
equals Squalus vulfinus Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: g, pi. 85, fig. 349.

Generic Synonyms:

Squalus (in part) Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: 9; also subsequent authors; not Squalus

Linnaeus, 1758.
Ga/^tt/ (in part) Rafinesque, Indice Ittiol. Sicil., 1810:46.
Carcharhinus (in part) Blainville, Bull. Soc. philom. Paris, 1816: 121.'
Carcfutrias (in part) Cuvier, Regne Anim., 2, 1817: 126; and subsequent authors; not Carcharias Rafinesque,

1810.
Alopecias Muller and Henle, Arch. Naturg., (3) i, 1837: 397; type, Carcharias vulfes Cuvier, 1817, equals

Squalus vulfinus Bonnaterre, 1788.
Vulfecula Garman, Mem. Harv. Mus. comp. ZooL, 36, 1913: 30; type species, Vulfecula marina Valmont,

Diet. Hist. Nat., 3, 1768: 740.*
Alofes Vladykov and McKenzie, Proc. N. S. Inst. Sci., ig, 1935: 46, for A. vulfes (wrongly referred by them

to Bonnaterre, 1788, instead of to Gmelin, 1789); evident misspelling for Alopias.

Generic Characters. Those of the family.

Range. Cosmopolitan in low and mid latitudes of all oceans, including the Mediter-
ranean.

Fossil Teeth. Eocene, Africa; Oligocene to Miocene, Europe.

Species. The species of this genus fall in two sharply defined groups. In one the rear
tip of the first dorsal terminates far in front of the origin of the pelvics; in the other the
first dorsal overlaps the pelvics.

The first group includes: pelagicus Nakamura, which is set apart by its notched and
denticulate teeth; the well known vulfinus of the Atlantic and eastern Pacific; also two

2. Photograph of eggs from female taken off Florida, contributed by Stewart Springer.

3. See footnote la, p. 320.

4.. By ruling of the International Commission on Zoological Nomenclature (Smithson. misc. Coll., j} [3], 1925:
27) Carman's revival of the name Vulpecula Valmont is not acceptable, because such of the latter's names as were
binomial were only accidentally so.

1 62 Memoir Sears Foundation for Marine Research

other named species, cflWa/wj Phillipps (1932) znd greyi Whitley (1937). However, it is
doubtful whether either of these last two is actually separable from vtdf'mus. The char-
acters which are supposed to distinguish caudatus^ are: tail no longer than trunk, caudal
peduncle up to one-half as deep as trunk at first dorsal and the anterior margin of pectoral
straight instead of convex. But the first two of these characters apply equally to some
Atlantic specimens (see p. 171), perhaps leaving only the shape of the pectoral as diag-
nostic. The only characters supposedly diagnostic of greyi are eyes "modified for back-
ward vision, are situated much further forward" and greenish color." Its author has in
fact suggested recently that greyi may be merely a color variety of caudatus.^ Until Aus-
tralasian specimens are actually compared with those of the eastern Pacific and Atlantic,
the question whether or not they are specifically distinct must remain open.

The second group, in which the rear tip of the first dorsal reaches as far back as the
origins of the pelvics, or even overlaps the latter, includes two well marked species, super-
ciliosus Lowe, 1840, of the tropical Atlantic and -profundus Nakamura, 1935, so far
reported only from Formosa. These two differ further from the vulpinus group in the
enormous size of their eyes (cf. Fig. 25 with 27) ; this is, in fact, their most arresting fea-
ture apart from their elongate tails.

Key to Species

I a. Rear tip of ist dorsal terminates considerably anterior to origin of pelvics.

2a. Teeth with central cusp strongly oblique, the outer margins with i or 2 denticles.

/)<?/fl^/V«j Nakamura, 1935.
Formosa.
2b. Teeth with central cusp erect or only slightly oblique 5 no marginal denticles.
3a. Anterior margin of pectoral convex. vulpinus 'Qonnzttrvt^ 1788, p. 167.
3b. Anterior margin of pectoral nearly straight. caWa/wj Phillipps, 1932.'

New Zealand, Australia.
lb. Rear tip of ist dorsal extends at least as far as origin of pelvics, or even overlaps
the latter.

4a. Rear tip of 2nd dorsal terminates considerably anterior to origin of analj pelvics
a little higher vertically than ist dorsal and a little larger in area} anterior mar-
gin of 1st dorsal strongly convex j no lower precaudal pit.

superciliosus Lowe, 1840, p. 163.
4b. Rear tip of 2nd dorsal terminating over base of anal; pelvics less than ^^ as high
vertically as ist dorsal and much smaller in area; anterior margin of ist dorsal
only very weakly convex; a precaudal pit below as well as above.

profundus Nakamura (1935).
Formosa.

5. Phillipps, N. Z. J. Sci. Tech., /j, 1932: 226. 6. Whitley, Rec. Aust. Mus., 20, 1937: 5.

7. Whitley, Fish. Aust., i, 1940: 132 8. Including greyi Whitley, 1937.

Fishes of the Western North Atlantic 163

Alopias suferciliosus (Lowe), 1840

Big-eyed Thresher

Figures 25, 26

Study Material. Young male, 1,296 mm. in total lengthy two embryos, 64 mm. and
632 mm. long} jaws of 18-foot specimen (Harv. Mus. Comp. Zool.) 5 all from the north
coast of Cubaj° also photographs of embryos from Salerno, Florida."

Distinctive Characters. This species is set apart from the Common Thresher, the only
Atlantic Shark with which it might be confused, by the following features: its relatively
enormous eye, longer snout, the tip of its first dorsal fin overlapping the pelvics, its second
dorsal terminating considerably in advance of the anal and only 10 or 11 teeth on a side
in each of its jaws (about 20 in the Common Thresher).

Remarks. The original description of suferciliosus was limited to the statement that
it is "at once distinguished from the only other known species of the genus, Carcharias
vulpes, Cuv., by the enormous eye and its prominent brow."" But the size of the eye is
so striking a character that the specimens described here can safely be referred to Lowe's
old species.

Description. Proportional dimensions in per cent of total length. Male embryo, 632
mm., from Cuba (Harv. Mus. Comp. Zool., No. 36155). Male, 1,296 mm., from Cuba
(Harv. Mus. Comp. Zool., No. 36090).

Trunk at origin of pectoral: breadth 5.8, 7.4; height 8.2, 9.1.

Snout length in front of: outer nostrils 4.3, 4.6} mouth 7.4, 6.0.

Eye: horizontal diameter 4.1, 2.8.

Mouth: breadth 4.6, 4.4; height 3.0, 2.6.

Nostrils: distance between inner ends 1.9, 1.8.

Labial furrow length: upper part 1.9, 1.95 lower 0.7, 0.5.

Gill opening lengths: ist 2.1, 2.9; 2nd 2.5, 2.95 3rd 2.4, 2.8; 4th 2.1, 2.2; 5th

1.8, 1.9.

First dorsal fin: vertical height 5.2, 5.2; length of base S-Sy 6.3.

Second dorsal fin: vertical height 0.6, 0.7; length of base 0.8, 0.9.

Anal fin: vertical height 0.8, 0.9; length of base i.i, i.O.

Caudal fin: upper margin 48.7, 49.1 ; lower anterior margin 6.3, 6.3.

Pectoral fin: outer margin 20.4, 19.1; inner margin 4.3, 3.8; distal margin 17.2,

18.1.

Distance from snout to: ist dorsal 33.3, 31.2; 2nd dorsal 45.3, 46.O; upper caudal

51.3, 50.9; pectoral 20.1, 17.O; pelvics 37.2, 36.5; anal 48.2, 47.8.

Interspace between: ist and 2nd dorsals 8.9, 8.9; 2nd dorsal and caudal 5.4, 4.6;

anal and caudal 1.6, 1.9.

9. Contributed by Luis Howell-Rivero. lo. Contributed by Stewart Springer.

II. Lowe, Proc. zool. Soc. Lond., 8, 1840: 39.

164

Memoir Sears Foundation for Marine Research

Distance from origin to origin of: pectoral and pelvics 19.2, 20.8; pelvics and

anal ii.i, 11.2.
Trunk, opposite ist dorsal, a little less than V^ as high as the length to origin of
caudal, thus more slender than in vulpinus. Caudal peduncle compressed laterally, with-
out lateral keels or ridges. A well marked precaudal pit above, but none below. Dermal
denticles of two kinds, mostly minute, very widely spaced, lanceolate, but expanded ante-
riorly on either side and spinous rather than scale-like, the blades not definitely marked
oflF from the pedicels; interspersed among these small denticles are much larger ones, in
pairs, the one close behind the other, of shapes more easily illustrated than described
(Fig. 26 C).

Figure 25. Alofias sufercilioms, young male, 1,296 mm. long, from north coast of Cuba (Harv. Mus.
Comp. Zool., No. 36090). A Left nostril, about 2.5 x. B Caudal peduncle, to show precaudal pit as viewed
from above.

Fishes of the Western North Atlantic

165

Figure 26. Alo-pias suferciliosus. Same specimen as shown in Fig. 25. A Anterior part of head from below,
a little more than Y2 natural size. B Left-hand corner of mouth from below to show labial furrows, about
1.3 X. C Dermal denticles, large and small. D Lateral view of small dermal denticle. E Apical view of
same. F Lateral view of a pair of large dermal denticles. C-F, about 130 x. G Left-hand upper and lower
teeth, about 0.9 x natural size. H Second upper tooth. / Sixth upper tooth. J Tenth upper tooth. K Second
lower tooth. L Sixth lower tooth. M Ninth lower tooth. H-M about 1.8 x.

1 66 Memoir Sears Foundation for Marine Research

Head about Vs as long as trunk to origin of caudal. Snout blunt-conical, its length
in front of mouth about Va length of head to origin of pectorals. Distance between nostrils
about Vs as long as snout in front of mouth. Eye approximately spherical, or a little higher
than broad, much larger than in vulpmus, its diameter a little more than Y2 as long as
snout in front of mouth, or between Vs and Ve as long as head to origin of pectorals. Spira-
cle a minute pore in embryo, about level with middle of eye, and behind the latter by a
distance equal to about V2 the diameter of eye; also visible on one side on young male
but not on other, and probably obsolete in adult. First and and gill openings (a little the
longest) about as long as diameter of eye, the 5th between % and •% as long as 2nd, and
strongly oblique, the 3rd above origin of pectoral. Nostril approximately transverse,
about ^/5 as long as horizontal diameter of eye, its inner end only about ^^ as far from
front of mouth as from tip of snout, its anterior margin expanded as a low, subtriangular
lobe. Mouth broadly rounded, about % as high as broad. Two labial furrows above, the
outer originating at corners of mouth and overlapping the inner, which originates a little
farther forward and extends Vs to ^^4 of the way toward the symphysis; one labial furrow
below, extending forward a short distance along lower jaw and around corner of mouth.

Teeth -[5E15- '^^ specimen examined, with one subtriangular cusp; similar in the two
jaws, the ist erect, nearly symmetrical, about as broad basally as long, but the others in-
creasingly oblique toward corners of jaws, their outer edges more and more strongly
convex, their inner margins increasingly concave with even curvature; the ist lower tooth
a little shorter than 2nd, and the outer 3 (lower jaw) or 4 (upper jaw) progressively
smaller, the outermost of all much the smallest.

First dorsal originates about midway between perpendiculars at inner corner of
pectoral and at origin of pelvics, its vertical height about Ys as great as length of head to
origin of pectorals, its anterior margin strongly convex, its apex rounded, its posterior
margin weakly concave, its free rear corner only about Y^ to Ye as long as its base, its rear
tip overlapping the pelvics for a short distance. Second dorsal only about Yg as long at base
as 1st dorsal, its apex broadly rounded, its free rear tip very slender, a little longer than its
base, ending anterior to origin of anal by a distance nearly as long as base of 2nd dorsal.
Interspace between 2nd dorsal and caudal about % as long as base of ist dorsal. Caudal
a little less than Y2 or about 48% of total length, slender, scythe-shaped, much as in
vulpinus but with terminal sector somewhat broader and more sharply demarked, though
without definite notch, its lower anterior margin more strongly convex. Anal similar
to the 2nd dorsal in shape. Pelvics about as long at bases as high, about 1.2 as high verti-
cally as 1st dorsal, and a little larger In area, the anterior margins weakly convex, the
apices narrowly rounded, the posterior margins strongly and evenly concave. Pectoral
about I.I times as long as head and a little less than Y2 as broad as long, Its anterior margin
strongly convex toward tip, its apex moderately rounded, the inner margin weakly con-
cave toward tip but deeply so toward Inner corner, the latter narrowly rounded.

Color. Dark mouse gray above and hardly paler below, the posterior margins of the
1st dorsal, pectorals and pelvics more or less dusky.

Fishes of the Western North Atlantic 167

Size. That the Big-eyed Thresher grows as large as the Common Thresher is indi-.
cated by the following facts: one of our specimens was from a 12-foot mother, others have
been taken from a female of about the same size, and an 1 8-foot specimen has been taken
(teeth pictured in Fig. 26).

Developmental Stages. An embryo of 64 mm., taken from the horny egg capsule,
already showing the extremely elongate caudal so characteristic of the adult, still had
well developed external gills and a long yolk stalk. One of 632 mm. in total length is evi-
dently ready for birth, since a well marked scar is alone reminiscent of the yolk stalk; this
resembles the young male pictured in Fig. 25, except that its eyes are somewhat larger
relatively, which is a characteristic common to embryo sharks; the longest gill openings
are about % as long as the diameter of the eye, its snout is blunter, and its pectorals are
narrower toward their tips.

Habits. Nothing definite is yet known as to the habits of the Big-eyed Thresher of
the Atlantic. Its very large eyes and its coloration (nearly as dark below as above) suggest
that it is chiefly a deep-water species like its Formosan relative, profundus.^^ But it is not
exclusively so, for the specimen pictured in Fig. 25 was near the surface,^' and perhaps
others of the scanty list of captures likewise.

Abundance and Range. The Big-eyed Thresher was first reported from Madeira
more than a century ago. It was not seen again until August 1941 when a female of 1 1 to
12 feet, containing embryos, was taken oflF Englewood on the west coast of Florida."
Very recently, females containing embryos (the young male pictured in Fig. 25) and an
1 8-foot specimen have been taken oflF the north coast of Cuba (oflF Matanzas, and near
Havana). We have been informed'" that the Museum Poey in Havana has a large mounted
specimen from Miami, Florida. No doubt the species is widespread in the tropical and
subtropical Atlantic.

Synonyms and References:

Alopecias suJ>erciliosus Lowe, Proc. zool. Soc. Lond., 8, 1840: 39 (Madeira).

Alofias (no specific name), Springer, Copeia, /, 1943: 54 (off Englewood, Florida, brief descr., embryos,
comparison with vulpinus and with frojundus) .

Alofias vulfintis (Bonnaterre), 1788

Common Thresher

Figures 27, 28

Study Material. Three alcoholic specimens from Massachusetts, 1,225 to 1,315 mm.
long (Harv. Mus. Comp. Zool., No. 486, 706, 1166); mounted specimens, about 9 feet
long, from Massachusetts (Harv. Mus. Comp. Zool., No. 926), and 4 feet 5 inches long

12. For description of the latter, see Nakamura (Mem. Fac. Sd. Agric. Taihoku., i^ [i], 1935: i).

13. It was harpooned.

14. Springer, Copeia, 1943: 54. We have received photographs of one of the embryos.

15. By Luis Howell-Rivero.

1 68 Memoir Sears Foundation for Marine Research

from Miami, Florida, in the same collection j fresh specimens not preserved — an adult
male of 13 feet taken August i, 1941 (jaws saved), a female 7 feet i inch taken June 15,
1943, immature males of 5 feet 2 inches (1,577 mm.) and 6 feet 10 inches (2,083 nmi.);
and immature female, 4 feet 10 inches (1,478 mm.) from Woods Hole, Massachu-
setts} also two small specimens from San Francisco and one from the west coast of South
America (Harv. Mus. Comp. Zool., No. 345, 519, 705).

Distinctive Characters. The enormously elongate tail sets the Common Thresher
apart at a glance from all other Atlantic sharks, except for its close relative, the Big-eyed
Thresher. It is marked off from the latter by its much smaller eye, shorter snout, by the tip
of its first dorsal considerably anterior to the origin of its pelvics, by the tip of its second
dorsal overlapping the base of its anal, and in having about 20 teeth on each side in each
jaw (only about 10 in superciliosus) .

Description. Proportional dimensions in per cent of total length. Female, 1,225 mni.,
from Nahant, Mass. (Harv. Mus. Comp. Zool., No. 486). Male, 2,083 mm., from Buz-
zards Bay, Mass. (field specimen).

Trunk at origin of pectoral: breadth 8.3, 7.3 ; height 9.6, 9.3.
Snout length in front of: outer nostrils 2.5, 1.75 mouth 3.6, 3.8.

Figure 27. Alofias vulfhtus, female, about 1,300 mm. long, from Massachusetts Bay (Harv. Mus. Comp. Zool.,
No. 1 166). A Caudal peduncle from above to show precaudal pit. B. Right-hand corner of mouth, with lips
separated to show single upper labial furrow.

Fishes of the Western North Atlantic

169

Figure 28. Alofias vulfinus. A Dermal denticles of specimen pictured in Fig. 27, about 50 x. 5 Apical view
of dermal denticle, about lOO x. C Teeth of a 13-foot Woods Hole, Massachusetts, specimen (Harv, Mus.
Comp. Zool., No. 36089), about natural size. D Second upper tooth of same. E Third upper tooth. F Fifth
upper tooth. G Fifteenth upper tooth. H Second lower tooth. / Sixth lower tooth. / Fifteenth lower tooth.
D-J, about 2 X.

Eye: horizontal diameter 1,5, 1.2.
Mouth: breadth 4.4, 3.75 height 3.1, 2.1.
Nostrils: distance between inner ends 1.4, 1.2.

Labial furrow length: upper part, 1.7, j lower 0.8, 0.6.

Gill of ening lengths : ist 2.0, 1.7; 2nd 2.1, 1.9J 3rd 2.1, 2.0; 4th 1.9, 1.9; 5th

1.8,1.9.

First dorsal fin: vertical height 6.6, 6.9; length of base S-(>i 6.4.

Second dorsal fin: vertical height 0.6, 0.4; length of base 0.7, 0.8.

Anal fin: vertical height 0.8,0.7; length of base 0.7, 0.9.

Caudal fin: upper margin 53.O, 53.9; lower anterior margin 5.7, 6.7.

Pectoral fin: outer margin 15.1, 15.8; inner margin 3.3, 2.3; distal margin 11.5,

14.4.

Distance frotn snout to : ist dorsal 22.5, 21.3 ; 2nd dorsal 39.8, 40.5; upper caudal

47.0,46.0; pectoral 14.0, 13.2; pelvics32.i, 31.4; anal 40.7, 42.6.

Interspace between: ist and 2nd dorsals 12.2, 12.6; 2nd dorsal and caudal 6.1,

5.0; anal and caudal 2.7, 2.2.

Distance from origin to origin of: pectoral and pelvics 1 8.7, 18.5; pelvics and anal

9.1,11.2.

lyo Memoir Sears Foundation for Marine Research

Trunk stout, somewhat compressed laterally, its dorsal profile strongly convex ante-
rior to I St dorsal, its depth opposite the latter about V^ its length to origin of caudal.
Caudal peduncle strongly compressed laterally, about i V2 times as high as thick, without
longitudinal lateral ridges or keels. A well marke_d precaudal pit above (Fig. 27 A) but
none below. Dermal denticles closely overlapping and very small, being only about 0.2 X
0.21 mm. in a 15-foot specimen, blades horizontal, usually with 3, sometimes with 5, low
keels, and as many rather short marginal teeth, the median largest; moderately long
pedicels.

Head and snout together subconical, between Y^ and Y^ as long as trunk to origin
of caudal. Snout rounded at tip and very short, its length in front of mouth only about
Y4 to V5 the length of head. Eye circular, moderately large, its margin considerably ante-
rior to front of mouth, its diameter about Y2 as long as snout in front of mouth in small
specimens, but only about Ys that length in large. Spiracle pore-like, on same level as center
of eye and behind latter by a distance about Y2 as great as length of snout in front of mouth.
Gill openings terminating relatively high up on the sides of neck, noticeably short, the
longest only a little longer than diameter of eye in small specimens, but about twice as long
in large ones; ist to 4th evenly spaced, the lower ends of 4th and 5th close together over
origin of pectoral. Nostril transverse, considerably nearer to mouth than to tip of snout, its
anterior margin expanded in low, subtriangular contour. Mouth broadly rounded, about
twice as wide as high. Upper labial furrow reaching about Ys of distance to symphysis, the
lower furrow only about Y2 as long as upper and visible only when mouth is open.

Teeth l^Ezi i^i specimen counted; similar in the 2 jaws, blade-like, subtriangular,
with single sharp-pointed cusp and smooth edges, the ist to 3rd uppers and ist and 2nd
lowers nearly symmetrical, but successive teeth increasingly oblique, with their outer
margins increasingly deeply concave; the 3rd upper tooth only about % as high as ist
and 2nd, or as 4th to lOth; ist lower tooth also very small; lOth or nth and subsequent
teeth in each jaw decreasing successively in size toward corners of mouth, the outermost
minute in lower jaw; i and sometimes 2 rows functional in front of mouth, 2 rows toward
corners.^*"

Origin of ist dorsal only slightly behind inner corner of pectoral, its rear tip anterior
to origin of pelvics by a distance about as great as length of snout in front of mouth, its
anterior margin moderately convex, the apex rounded, the posterior margin only slightly
concave basally, its free rear tip only about V5 as long as its base, its vertical height less
than V2 as great as length of pectoral. Second dorsal only about Ys as long as ist dorsal
along anterior margin, its origin much nearer to origin of caudal than to rear end of base
of 1st dorsal, its apex rounded, its rear tip slender, elongate, nearly or quite twice as long
as its base, the rear end of its base about over origin of anal, or a little anterior to latter.

16. According to Moreau (Hist. Nat. Poiss. France, z, 1881 : 288) the first small tooth in the lower jaw is lost with
age; also, in the upper jaw there may be a minute first tooth in small specimens, which is similarly lost with
age. But the specimens we have seen lack this small median upper tooth, the first pair of large teeth being close
together at the upper symphysis.

Fishes of the Western North Atlantic 171

Caudal usually a little more than V2 the total length," its upper lobe narrow, scythe-
shaped, with a small rounded subterminal prominence but no definite subterminal notch,
its lower anterior corner produced as a small but definite subtriangular lobe, the lower
anterior margin about V^ to H as long as the upper margin, its axis raised only slightly.
Anal similar to 2nd dorsal in size and shape, its origin posterior to rear end of base of 2nd
dorsal by a distance as long as its own base, or a little longer. Pelvics about as large in area
as 1st dorsal, with weakly convex anterior margins, rounded apices, moderately concave
distal margins and subacute inner corners. Claspers of adult males about 4 times as long
as pelvic fins and very slender. Pectoral nearly or quite twice as long as height of ist dorsal
along anterior margin, falcate, with very broad base, the anterior margin rather strongly
convex in small specimens but tending to become less so in large,'' the apex broadly
rounded, the distal margin deeply and evenly concave, the inner corner subacute, the
inner margin only a little more than V2 as long as breadth of base.

Color. Back and upper sides varying between brown, blue slate, slate gray, blue gray
and dark lead, even nearly black, often with metallic luster; shading along sides to white
below, except that lower surface of snout in front of nostrils, as well as lower surfaces
of pectorals, may be of same hue as upper sides; white of lower surface reaching farthest
upward on sides from axil of pectoral to opposite rear part of ist dorsal and again rear-
ward from pelvics j the sides near pectorals, the lower surface from pelvics to caudal, and
sometimes the belly may be more or less mottled with gray; iris black or green.

Size. Maximum length 20 feet or more, with lengths of 13 to 16 feet common.
Judging from the sizes of females with embryos and of males with large claspers, sexual
maturity probably is not attained at a length less than 14 feet. A female of 4,410 mm. ( 14
feet 6 inches) was found to contain an embryo of 1,550 mm. (5 feet i inch)." On the other
hand, a free-living specimen as small as 46 inches has been reported, while many of 4 to 5
feet have been taken at Woods Hole. One of 149 cm. (about 4 feet, 10Y2 inches) taken
there is described as still showing the umbilical scar,^° but no trace of it is to be seen on
another slightly smaller specimen (4 feet 4 inches) which we have examined from the
same locality. The few recorded weights range from about 300 to 320 lbs. at lO feet, 375 to
400 lbs. at about 13 feet, and about 500 lbs. at 14 feet 5 inches, up to a maximum of per-
haps 1,000 lbs.

Develof mental Stages. No information is available about the embryo, except that
the caudal is about as long, relatively, as in the adult. But the decrease in relative size of
the eye with growth after birth suggests that it is even larger in the embryo, as is so com-
monly the case. Apparently the number of young in any one litter is much smaller than
in many other ovoviviparous species, for females have been reported as containing two

17. Ratio of caudal length to trunk length (snout to origin of caudal) ranges from i.i:i to 1.3:1 in specimens
examined from Massachusetts, San Francisco and the west coast of South America, but only 1.04:1 in a
Mediterranean example measured by Tortonese (Atti Soc. ital. Sci. nat., yy, 193S: 293).

18. Owing to the obliquity of their basal insertions, the pectorals often appear as straight-edged, or nearly so, in
photographs of large Threshers suspended by the caudal peduncle, as we have observed.

19. Poey, An. Soc. esp. Hist. Nat., 5, 1876: 383. 20. Springer, Copeia, 1943: 55-

1 72 Memoir Sears Foundation for Marine Research

or four only. But they are correspondingly large when born, for those in a mother of
about 15V2 feet (4,700 mm.) measured respectively 1,500 and 1,550 mm. (about 5
feet)." Others, perhaps from smaller mothers, are considerably smaller at birth, judging
from the small sizes of free-living specimens repeatedly recorded.

Habits. The Thresher is a typically pelagic species, most often seen at least a few
miles offshore, but often coming close in to pursue small fish. It is com.monly described as
usually keeping near the surface. We have seen Threshers jumping on one occasion, these
being easily identified by their long tails. But while it is from specimens seen at the surface,
or taken in nets set shoal, that the majority of records of its occurrence emanate, it is
equally certain that it may descend to moderate depths on occasion, since there is record
of at least one specimen captured on hook and line at 35 fathoms in British waters.^^

It feeds chiefly, if not solely, on whatever smaller schooling fishes may be avail-
able; in North American waters most commonly on mackerel, bluefish {Pomatomus)^
shad (Alosa) and menhaden (Brevoortia), of which it destroys great numbers; no doubt
it feeds on herring also, as well as on bonito and squid. In North European waters its diet
includes pilchards, garfish, etc. Twenty-seven mackerel have been recorded from a speci-
men 13V2 feet long, and one-half bushel of garfish (Belone) from another in Scottish
waters. The method by which it captures its prey is highly specialized; in general accounts
it has been described repeatedly as using "its whiplike tail to splash the water, while it
swims in narrowing circles round a school of fishes, which are thus kept crowded together
until the moment of slaughter . . . Sometimes a pair of threshers work together . . . "^°
That it also uses its tail on occasions to stun a prospective victim is proved by eyewitness
accounts of one in Irish waters rising and killing a wounded sea bird with a stroke of its
tail, then swallowing it,'* and of another at La Jolla, California,'"' injuring a single small
fish by lashing at it repeatedly with its tail. Perhaps it is hardly necessary at this late date
to remark that the time-honored tradition that the Thresher leagues with the swordfish
to attack whales, which was doubtless based on its being confused with the killer whale
(Orca), has long since been relegated to the category of myth.

Presumably, young are produced throughout its geographic range, very small speci-
mens having been taken off southern Florida on the one hand and in New England waters
on the other. No information is available as to season when the young are born, or when
mating takes place.

Abundance and. Relation to Man. Of late years the Thresher has not appeared in
sufficient abundance anywhere along the Atlantic coast of America to be of any commercial
importance one way or the other. However, when it gathers in any numbers in pursuit
of small fish, as is said to have happened at times in the past, it has been an annoyance
to mackerel fishermen by becoming entangled in their nets. This is a frequent occurrence
in British waters where the Thresher is a more familiar species. It is entirely harmless.

21. Vaillant, Bull. Soc. philom. Paris, (7) 10, 1885: 41. 22. Day, Fish. Gt. Brit., t, 1880-1884.: 302.

23. Nichols and Murphy, Brooklyn Mus. Sci. Bull., j, 1918: 21.

24. Blake-Knox, Zoologist, (2) i, 1866: 509. 25. Allen, Scierce, N. S. $&, 1923: 31.

Fishes of the Western North Atlantic 173

Range. Pelagic; in warm temperate and subtropical latitudes; north commonly in
the eastern Atlantic to southern Ireland, less regularly to the North Sea and inward as far
as the Danish coast and Kattegat; occasionally to the Orkneys and to the Norwegian coast
as far north as Lofoten and Trondhjem. Also widespread in the Mediterranean, and
recorded from Madeira and Cape of Good Hope. It is known in the west as far north as
Nova Scotia and the Gulf of St. Lawrence, south to Brazil and northern Argentina; also,
in the eastern Pacific, from Oregon to the Isthmus of Panama and Chile.^® It is also re-
ported from the Hawaiian Islands, Fanning Island and "Polynesia," Japan, Korea and
China, New Zealand and Australia, and from Ceylon, Arabia and Natal. But whether the
Thresher (or Threshers) of the vulpinus group of the western Pacific, New Zealand and
the Indian Ocean are identical with vulpinus of the eastern Pacific and Atlantic, or whether
more than one species of the group occurs in those regions, is still an open question (p.
162).

Occurrence in the Western Atlantic. The Thresher is reported more frequently and in
larger numbers from the offing of southern New England than from anywhere further
south along the east coast of the United States. Over the continental shelf off Block Island
it has been described as the commonest shark," appearing in May, being most plentiful
in June, and remaining until autumn. In the vicinity of Woods Hole, Vineyard Sound
and Buzzards Bay, Threshers are taken from time to time in the traps between April
and late autumn (see Study Material, p. 168). There is record, in fact, of three fish
of 16 feet in one trap in a single morning, and specimens up to 20 feet have been reported
locally. Although only two specimens have been recorded in print from Nantucket,
Threshers enter the Gulf of Maine in some numbers, at least during some years. Thus,
we saw several large ones leaping in Pollock Rip off the southern angle of Cape Cod on
August 4, 19 13; it is recorded from Provincetown at the tip of Cape Cod and repeatedly
from various localities in Massachusetts Bay {e.g., Boston Harbor, Nahant) as well as
from various localities along the coast of Maine, in Passamaquoddy Bay,^* and from the
cold waters of the Bay of Fundy (Basin of Minas).^° From time to time Threshers are
taken entangled in the nets off the outer coast of Nova Scotia; seemingly they are not
rare on the Scotian Fishing Banks and they have even been reported from the Bay of
Chaleur in the Gulf of St. Lawrence, this being the most northerly known record for them •
on the western side of the Atlantic.

Being a creature of at least moderately warm waters, it is surprising that the Thresher
has been reported more frequently and in larger numbers off southern New England than

26. Our own comparison of specimens from San Francisco, California, and the west coast of South America (p.
168) with a considerable series from Massachusetts fails to reveal any significant differences in proportionate
dimensions, shape and relative location of fins, length of tail relative to trunk, size of eye, or in shape and
number of teeth.

17. Nichols and Murphy, Brooklyn Mus. Sci. Bull., 3, 1916: 20.

28. McKenzie, Proc. N. S. Inst. Sci., 20 (:), 1939: 14.

29. But the old report of Threshers as common there (Knight, Cat. Fishes N. S., x866: 8) may not have been well
founded.

174 Memoir Sears Foundation for Marine Research

from anywhere farther south along the Atlantic coast of America. No doubt the fact that
there is no record of the Thresher for Georges Bank is accidental. But the paucity of re-
ported captures westward and southward from the Block Island-Woods Hole region can-
not be explained thus, for so striking is the Thresher in general aspect that any specimen
taken is likely to be reported in the daily press, if not in strictly scientific literature. Actu-
ally, we have found but one positive record of it for Rhode Island and Connecticut; four
for Long Island, New York; one near New York; three in recent years for New Jersey;
one for Maryland; two for Cape Lookout, North Carolina (from which it appears that
few come inshore close enough along this sector to be picked up in the pound nets) . While
Threshers have been described as rather numerous at times among the Florida Keys, there
are only three or four reports of it along the east coast of Florida, including a small one
from Miami in the Museum of Comparative Zoology. One has been reported to us from
Biloxi, Mississippi.'" Nor does it appear to be any more plentiful anywhere farther to the
south, where published captures total only three, one being for the Havana region, one for
Santa Catherina, Brazil, and one for northern Argentina (Lat. 38° S.). It has not been
reported from Bermuda.

In the northern sector of its range the Thresher appears only in spring, summer or
autumn, being wholly absent in winter. But nothing is known of its seasonal incidence any-
where else in the western Atlantic.

Synonyms and References:

I. Atlantic:

Sea Fox, Borlase, Nat. Hist. Cornwall, 1758: 265 (Cornwall) ; Brookes, Nat. Hist., 3, Fishes, Serpents, 1763:
31 (descr., Medit.) ; Pennant, Brit. Zool., 3, 1769: 86, pi. 4 (descr., Gt. Brit.) ; Couch, Fish. Brit. Isles,
I, 1867: 37, pi. 7 (descr., Gt. Brit.).

Long Tailed Shark, Pennant, Brit. Zool., 5, 1776: 1 10, pi. 14 (descr., Gt. Brit.).

Squalus vulfinus Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: g, pi. 85, fig. 349 (descr., Medit.).

Squalus vulfes GmtWn, in Linnaeus, Syst. Nat., i, 1789: 1496 (descr.); Bloch and Schneider, Syst. Ichthyol.,
1801: 127 (descr.); Turton, Brit. Fauna, i, 1807: I12 (Gt. Brit.); Risso, Ichthyol. Nice, 1810: 36
(descr., Medit.); Mitchill, Trans. Lit. Phil. Soc. N. Y., 1, 1815: 482 (descr., Long Island, N. York);
Nardo, Prod. Ichthyol. Adriat., 1827: 9 (Medit.); Voigt, in Cuvier, Tierreich, 2, 1832: 506 (descr.);
Jenyns, Manual Brit. Vert. Anim., 1835: 498 (descr.); Couch, Cornish Fauna, 1838: 50 (Cornwall);
Buckland, Hist. Brit. Fish., i88i: 218 (size, weight); Gatcombe, Zoologist, (3) 6, 1882: 434 (Devon-
shire, England); Zoologist, (3) 9, 1885: 352 (Sussex, England).

Squale renard, Lacepede, Hist. Nat. Poiss., 4° ed., /, 1798: 167, 267, in Buffon, Hist. Nat. (descr.) ; in Son-
nini. Hist. Nat. Poiss., 4, 1 801-1802: 89 (descr., meas.).

Alopas macrourus Rafinesque, Caratt. Gen. Nuov. Sicil., 1810: 12 (Sicily); Indice Ittiol. Sicil., 1810: 45
(Sicily) ; Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1839: 313.

Squalus (Carcharhinus) vulfes Blainville, Bull. Soc. philom. Paris, 1816: 121 (name); in Vieillot, Faune
Franc, Poiss., 1825: 94, pi. 14, fig. I (descr., meas., French seas).

Carcharias vulfes Cuvier, Regne Anim., 2, 1 8 1 7 : 126 (descr.) ; Cloquet, Diet. Sci. Nat., 7, 1 8 1 7 : 67 (general) ;
Risso, Hist. Nat. Europe Merid., 3, 1826: 120 (descr., Medit.) ; Fleming, Hist. Brit. Anim., 1828: 167
(descr., Gt. Brit.) ; Bory de St. Vincent, Diet. Class. Hist. Nat., 15, 1829: 597 (general) ; Yarrell, Brit.
Fish., 2, 1836: 379 (descr., Gt. Brit.); Templeton, Charlesworth's Mag. Nat. Hist., (2) i, 1837: 413
(Ireland); Storer, Rep. Fish, Rept. Birds Mass., 1839: 181 (Massachusetts); Boston J. nat. Hist., 2,

30. Personal communication from Stewart Springer.

Fishes of the Western North Atlantic 175

1839: 529 (descr., Massachusetts) ; Yarrcll, Brit. Fish., 2nd ed., 2, 1841 : 522 (descr., Gt. Brit.) ; DeKay,
Zool. N. Y., 4, 1842: 348, pi. 61, fig. 199 (descr., near N. York) ; Cuvicr, Rcgne Anim., 111. Ed., Poiss.,
1843: 361; Linsley, Amer. J. Sci., 47, 1844: 76 (Connecticut); Guichenot, Explor. Algier. Poiss.,
1850: 124 (Algeria); Gosse, Nat. Hist. Fish., 1851: 306 (general); Thompson, Nat. Hist. Ireland, 4,
l8;6: 250 (Ireland); DeKay, Rep. St. Cab. Nat. Hist. N. Y. (1855), 1858:64 (near N. York); Blake-
Knox, Zoologist, (2) /, 1866: 509 (Ireland) ; Storer, Mem. Amer. Acad. Arts Sci., N. S. 9, 1867: 221, pi.
36, fig. 3, also Fishes Mass.: 245, pi. 36, fig. 3 (descr., habits, Massachusetts) ; Buckland, Hist. Brit. Fish.,
1881: 218 (size, weight, Gt. Brit.); Clogg, Zoologist, (3) 5, 1881: 386 (Cornwall).

Alopecias vulfes Miiller and Henle, S. B. Akad. Wiss. Berlin, 1837: 114; Arch. Naturg., (3) i, 1837: 397
(name) ; Couch, Fish. Brit. Isles, /, 1867: 37, pi. 7 (descr., Gt. Brit.) ; Gunther, Cat. Fish. Brit. Mus.,
8, 1870: 393, 518 (descr., synonyms, distrib.) ; Collett, Norges Fisk., 1875: 208 (Norway); Giglioli,
Elenc. Pesc. Ital., 1880: 52 (Medit.) ; Parona, Atti Soc. Nat. Modena, (3) i, 1883 (Medit., not
seen); Carus, Prod. Faune Medit., 2, 1 889-1 893: 507 (Medit.); Oppel, Lehrb. Verg. Mikr. Anat.
Wirbelt., /, 1897: 51 (histology); Sicher, Atti Accad. gioenia, (4) 11 (5), i8g8: 16 (Medit.); Grieg,
Norsk Tidskr. Fiskeri, 21, 1902: 469 (Norway, not seen) ; Bridge, Camb. nat. Hist., 7, 1904: 452 (gen-
eral) ; Werner, Zool. Jb., Syst. Abt., 2/, 1904: 28; (Medit.) ; Southwell, Zoologist, (4) 10, 1906: 398
(Norfolk, England); Gunther, Encycl. Brit., nth ed., 24, 191 1: 808 (general); Metzelaar, Trop.
Atlant. Visschen, 1919: 190 (Madeira, C. of Good Hope); Borri, Mem. Soc. tosc. Sci. nat., 1934: 93
(Medit.).

Alofiasvulps Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1838-1839: 313 (general).

Alopias vulfes Bonaparte, Icon. Faun. Ital., 3 (2), 1835: plate not numbered (descr., ill.) ; Mem. Soc. neucha-
tel. Sci. nat., 2 (8), 1839: 9 (in synopsis); Miiller and Henle, Plagiost., 1841: 74, pi. 55 (descr., ill.);
Hamilton, Brit. Fish., 2, 1843: 313; also subsequent eds. (Gt. Brit.); Bonaparte, Cat. Pesc. Europ., 1 846:
18 (distrib.); Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 505 (in synopsis); Kneeland, Boston
J. nat. Hist., 5, i 847: 485 (anat.) ; Nardo, Sinon. Modern. Spec, descr. Pesci St. Chiereghin, 1847: 1 1 1
(name); Baikie, Zoologist, //, 1853: 3846 (Orkneys); Kr0yer, Danmarks Fisk., 3, 1853: 929, 937
(Denmark); Nardo, Atti 1st. veneto, (3) 5, 1859-1860: 787 (Medit.); Gill, Proc. Acad. nat. Sci.
Philad., Addend., 1 861: 60 (name); Dumeril, Hist. Nat. Poiss., /, 1 865: 421 (descr., distrib.);
Socage and Brlto Capello, Poiss. Plagiost., 1866: 14 (Portugal); Caruccio, Atti Soc. ital. Sci. nat., 12,
1869: 568 (Medit.) ; Brito Capello, J. Sci. math. phys. nat. Lisboa, 2, 1869: 1 40; also in separate, Cat.
Peixes Port., 1869: 10 (Portugal); Ninni, Ann. Soc. Nat. Modena, 5, 1870: 66 (Medit.); Canestrini,
in Cornelia, et al., Fauna d'ltal., 5, 1872: 46 (Medit.); Baird, Rep. U.S. Comm. Fish. (1871-1872),
1873: 827 (Woods Hole); Gill, Rep. U.S. Comm. Fish. (1871-1872), 1873: 813 (Cape Cod to
Florida); Poey, An. Soc. esp. Hist. Nat., 5, 1876: 383; Enumerat. Pise. Cubens., 1876: 187 (size, em-
bryo, Cuba) ; Gervais and Boulart, Poiss., 3, 1877: 188, pi. 72 (descr.) ; Jones, List Fish. N. S., 1879: 9
(Nova Scotia) ; Doderlein, Man. Ittiol. Medit., 2, 1 881 : 5 2 (Medit.);" Moreau, Hist. N.t. Poiss. France,
I, 1881: 287 (descr., France) ; Perugia, Elenc. Pesc. Adriat., 1881:54 (.Adriatic) ; Pascoe, Nature, Lond.,
23, 1881: 35; Hasse, Naturl. Syst. Elasm. besond. Theil, 1882: 221, pi. 29, fig. 14-21 (vertebrae,
dermal denticles) ; Jones, Proc. N. S. Inst. Sci., 5, 1882: 95 (same as Jones, 1879) ; Jordan and Gilbert,
Bull. U.S. nat. Mus., 16, 1883: 27 (distrib.); Day, Fish. Gt. Brit., 2, 1880-1884: 300, pi. 157 (descr.,
habits, Gt. Brit.); Goode, Fish. Fish. Industr. U.S., 1, 1884: 672 (general); Petersen, Vidensk. Medd.
naturh. Forcn. Kbh. (1884-1886), 1884: 160 (Kattegat); Vaillant, Bull. Soc. philom. Paris, (7) 10,
1885: 14 (embryos, sizes) ; Graeffe, Arb. zool. Inst. Univ. Wien, 7, 1886: 446 (Medit.) ; Jordan, Rep.
U.S. Comm. Fish. (1885), 1887: 797 (distrib.); Garman, Bull. Mus. comp. Zool. Harv., 17, 1888:
80, pi. 12, 13 (lateral line syst.); Nelson, Rep. St. Geol. N. J., 1890: 662 (common, N. Jersey);
Lilljeborg, Sverig. Norg. Fisk., 3 (2), 1 891 : 627 (Norway, Denmark) ; Dean, Fishes Living and Extinct,
1895: 89 (general); Smitt, Hist. Scand. Fish., 2, 1895: 11 36 (descr., Scand.) ; Jordan and Evermann,
Rep. U.S. Comm. Fish, (j 895), 1896: 217 (distrib.); Bull. U.S. nat. Mus., 47 (l), 1896: 45 (descr.,
distrib.); Smith, Bull. U.S. Bur. Fish., /;, 1898: 89 (Woods Hole); Jordan and Evermann, Bull. U.S.
nat. Mus., 47 (4), 1900: pi. 6, fig. 20 (ill.); Stirton, Ann. Scot. nat. Hist., 1900: 17 (Scotland); Bean,

31. See Doderlein, above, for additional references for the Mediterranean in publications not accessible to us.

1 76 Memoir Sears Foundation for Marine Research

Rep. For. Comm. N. Y. (1900), 1901 : 378 (abund. oflF Martha's Vineyard, and near N. York) ; Linton,
Bull. U.S. Bur. Fish., /p, 1901: 428 (parasites. Woods Hole) ; Popta, Ann. Sci. nat., (7) 12, 1901: 205
(gill arches, Concarneau, France); Mcintosh, Ann. Mag. nat. Hist., (7) 10, 1902: 254. (food, Scot-
land); de Braganza, Result Invest. Sci. "Amelia," 2, 1904: 46 (Portugal); Sharp and Fowler, Proc.
Acad. nat. Sci. Philad., 56, 1 904: 505 (Nantucket) ; Bean, Bull. N. Y. St. Mas., 60, Zool. 9, 1903: 33
(descr., N. York) ; Borca, Arch. Zool. exp. gen., (4) 4, 1905: 204, 207 (France) ; Collett, Norges Fisk., 5,
1905: 79 (Norway) ; Jordan, Guide to Study Fish., /, 1905: 536 (general) ; Fowler, Rep. N. J. Mus.
(1905), 1906: 53 (N. Jersey) ; Fowler, Proc. Acad. nat. Sci. Philad., $8, 1906: 79 (N. Jersey) ; Tracy,
Rep. R. I. Comra. inl. Fish., 1906: 45 (off Rhode Island) ; Kendall, Occ. Pap. Boston Soc. nat. Hist., 7
(8), 1908: 5 (New England) ; Fowler, Proc. Acad. nat. Sci. Philad., 60, 1908: 54 (Newport, Rhode Is-
land, size) ; Proc. Acad. nat. Sci. Philad., 61, 1 909: 407 (N. Jersey) ; Proc. Acad. nat. Sci. Philad., 62,
191 1 : 599 (N. Jersey) ; Tracy, Rep. R. I. Comm. inl. Fish., 1910: 60 (Rhode Island) ; Roule, Bull. Inst,
oceanogr. Monaco, 243, 1912: 9 (Medit.) ; Halkett, Check List Fish. Canad., 1913: 40 (Gulf of St.
Lawrence, Nova Scotia, Basin of Minas, Bay of Fundy) ; Le Danois, Ann. Inst, oceanogr. Monaco, 5(5),
1913: 16 (English Channel); Nichols, Abstr. Linn. Soc. N. Y., 20-23, 1913: 91 (season off N. York);
Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., jr (2), 1913: 736 (Woods Hole, Mass.); Fowler,
Proc. Acad. nat. Sci. Philad., 66, 1914: 342 (N. Jersey); Coles, Proc. biol. Soc. Wash., 2S, 1915: 91
(N. Carolina) ; Nichols and Murphy, Brooklyn Mus. Sci. Bull., 5 (i), 1 91 6: 20 (descr., habits, distrib.) ;
Lahille, Physis B. Aires, 5, 1 921: 63 (Argentina) ; Enum. Feces Cartilag. Argent., 1 921: 15 (north. Ar-
gent.) ; Townsend, Bull. N. Y. zool. Soc, 26, 1923: 78 (general) ; Ribeiro, Mus. nac. Rio de J., Fauna
Brasil. Peix., 2 (l), Fasc. I, 1923: 16, pi. 5 (descr., Brazil); Jenkins, Fish. Brit. Isles, 1925: 313 (Gt.
Brit.) ; Wilson, Proc. U.S. nat. Mus., 64 (17), 1925: 12 (parasites, Woods Hole) ; Barnard, Ann. S. Afr.
Mus., 21 (l), 1925: 34 (S. Afr.) ; Ehrenbaum, in Grimpe and Wagler, Tierwelt N.- u. Ostsee, 7 (l2'),
1927: 14 (general. North Sea) ; Nichols and Breder, Zoologica, N. Y., p, 1927: 18 (general) ; Lonnberg,
Fauna Flora Uppsala, 1927: 258 (Bohusland, Sweden, not seen) ; Rey, Fauna Iberica Feces, i, 1928: 41 1
(descr., Spain); Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 20 (general); Truitt, Bean and
Fowler, Bull. Md. Conserv. Dep., 1929: 29 (Maryland); Breder, Amer. Mus. Novit., 382, 1929: 4
(use of tail) ; Wilson, Proc. U.S. nat. Mus., 60 (15), 1932: 60 (ref.) ; Pearson, Invest. Rep. U.S. Bur.
Fish., (10) /, 1932: 18 (off C. Hatteras) ; Belloc, Rev. des Trav. Peches Marit., 7 Fasc. 2, 1934: 140
(Morocco, Madeira) ; Daniel, Elasmobranch Fishes, Univ. Calif. Press, 1 934: 38 (Stomodeal denticles) ;
Ehrenbaum, Fauna Ichthyol., Cons. Intern, explor. Mer., 1935: plate not numbered (general); Nobre,
Fauna Marinha Port. Vert., /, 1935: 439, pi. 63, fig. 198 (descr., Portugal); Pozzi and Bordale, An.
Soc. cient. argent., 120, 1935: 150 (north. Argent.) ; Lubbert and Ehrenbaum, Handb. Seefisch. Nordeu-
rop., 2, 1936: 279 (general, N. Europe) ; Norman and Eraser, Giant Fishes, 1937: 24 (general) ; Norris,
Plagiost. Hypophysis, 1941: 29, pi. II, fig. 41-43 (brain).

Squnlus alofecias Grxy, Cat. Fish Coll. Descr. by L. T. Gronow, in Brit. Mus., 1854: 7 (in list).

Alocefias vulfes Yung, Arch. Zool. exp. gen., (3) 7, 1899: 140 (name, evidently misprint for "Alopecias").

Vulfecula marina^- Garman, Mem. Harv. Mus. comp. Zool., jd, 1913: 30, pi. 7, fig. I (descr.) ; Radcliffe,
Trans. Amer. Fish. Soc, 1914: 37 (N. Carolina); Bull. U.S. Bur. Fish., 34, 1916: 246, pi. 38, fig.
3 (teeth, N. Carolina) ; Smith, Amer. Mus. J., 16, 1916: 347 (habits) ; Huntsman, Contr. Canad. Biol.,
(1921), J, 1923: 56 (Bay of Fundy) ; Meek and Hildebrand, Field Mus. Publ., 15, 1923: 62 (general) ;
Linton, Proc. U.S. nat. Mus., 64 (21), 1924: 13, 35, 56, 82 (parasites); Bigelow and Welsh, Bull.
U.S. Bur. Fish., 40 (l), 1925: 32 (descr., habits. Gulf of Maine) ; Wilson, Bull. U.S. nat. Mus., 158,
1932: 463, 464 (parasites. Woods Hole) ; Nigrelli, Amer. Mus. Novit., 996, 1938: 7 (parasites).

Alopias vulpinus Fowler, Copeia, 30, 1916: 36 (mid. Atlant., U.S.); Proc. Boston Soc. nat. Hist., 35, 1917:
1 10 (Newport, Rhode Island); Proc. biol. Soc Wash., 55, 1920: 143 (N. Jersey); Proc. Acad. nat.
Sci. Philad., ^4, 1922: 3 (size, N. Jersey); Copeia, 156, 1926: 146 (N. Jersey, small size); Jordan,
Manual Vert. Anim. NE. U.S., 1929: 11 (general); Jordan, Evermann and Clark, Rep. U.S. Comm.

32. According to the International Commission on Zoological Nomenclature (Smithson. Misc. Coll., 7j [3], 1925:
27), Carman's revival of the name Vulfecula marina Valmont is not acceptable because such of Valmont's
(1768) names as were binomial were only accidentally so.

Fishes of the Western North Atlantic 177

Fish. (1928), 2, 1930: 18 (distrib.) ; Bigelow and Schroeder, Canad. Atlant. Fauna, biol. Bd. Canad.,
12*, 1934: II (descr., Gulf of St. Lawrence, Nova Scotia); Marchand, Fish Mar. Biol. Surv. S. Afr.,
Fish. Bull. 2, 1935: 32 (S. Africa); Howell-Rivero, Proc. Boston Soc. nat. Hist., 41 (4), 1936: 45
(embryos, Cuba) ; Fowler, Bull. Amer. Mus. nat. Hist., yo (l), 1936: 43 (descr., distrib.) ; Bigelow and
Schroeder, Bull. U.S. Bur. Fish., 4S, 1936: 322 (name); Tortonese, Atti Soc. ital. Sci. nat., yy, 1938:
293 (Medit.) ; Springer, Proc. Fla. Acad. Sci., 5, 1939: 34 (Florida) ; McKenzie, Proc. N. S. Inst. Sci.,
2 0 (l), 1939: 14 (Bay of Fundy) ; Hildebrand, Copeia, 1941 : 221 (N. Carolina) ; Fowler, Arqu. Zool.
Estado Sao Paulo, j, 1942: i 28 (Brazil) ; Lunz, Bull. S. Carolina St. Planning Bd., 14, 1944: 27 (Flor-
ida) ; Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Carib. Coram., Wash., 194;: 107,
fig. 37 (descr., ill., habits, range) ; Fowler, Mongr. Acad. nat. Sci. Philad., 7, 1945: 93 (6oo-lb. spec, N.
Carolina).

Alofias vulfitius Bonnaterre, A. vulfes Gmelin, or Vulpecula marina Valmont; Jordan, Copeia, 49, 1 91 7: 89
(name).

Alofes vulfes Vladykov and McKenzie, Proc. N. S. Inst. Sci., ig, 1935: 46 (Nova Scotian Banks and Bay of
Fundy).

2. Pacific Coast of America:

Carcharias vulfes Gray, Hist. Chile, Zool., 2, 1858: 363 (Chile).

Alofias vulfes Ay ci%, Proc. Calif. Acad. Sci., 5, 1863: 66 (California) ; Eigenmann, Proc. U.S. nat. Mus., 1$,

1893: 134 (San Diego, California); Allen, Science, N. S. 58, 1923: 31 (use of tail, California);

Walford, Fish Bull. Sacramento, 45, 1935: 41 (California).
Alofecias vulfes Perez Canto, Estud. Escual. Chile, 1886: 5 (listed for Chile); Philippi, An. Univ. Chile,

ji, 1887: 551, pi. 5, fig. I (descr., ill., Chile); Delfin, Rev. chil. Hist, nat., 4, 1900: 71 (Chile);

Quijada, Boll. Mus. nac. Chile, 5, 191 3: ill (listed for Chile).
Alofecias barrae Perez Canto, Estud. Escual. Chile, 1886: 6 (descr., Chile) ; Philippi, An. Univ. Chile, 7/,

1887: 553, pi. 5, fig. 2 (descr., ill., Chile).
Alofecias chilensis Philippi, An. Univ. Chile, /op, 1901: 310 (Chile).
Alofecias longimana Philippi, An. Univ. Chile, xop, 1901: 308 (descr., Chile); Quijada, Boll. Mus. nac.

Chile, 5, 1913: III (listed for Chile).
Alofias vulfinus Barnhart, Mar. Fish, south. Calif., 1936: 10 (off California); Schultz, Univ. Wash. Publ.

Biol., 2 (4), 1936: 131 (Washington, Oregon); Beebe and Tee-Van, Zoologica, N. Y., 26, 1941: 97

(ref.).

3. Central and Western Pacific, Australasia, Indian Ocean:''

Alofecias vulfes Hutton and Hector, Colonial Mus. Geol. Surv. N. Z., 1872: 78 (N. Zealand); Macleay,
Proc. Linn. Soc. N. S. W., d, 1881 : 359 (descr., Aust.) ; McCoy, Prod. Zool. Victoria, i Decade 9, 1884:
pi. 88 (descr., meas., ill., Aust.); Boulenger, Proc. zool. Soc. Lond., 1889: 243 (Arabia); Lucas, Proc.
roy. Soc. Vict., N. S. 2, 1890:43 (Aust.) ; Sauvage, in Grandidier, Hist. Phys. Nat. Madagascar, /6, 1 891:
511 (Madagascar); Giinther, J. Mus. Godeffroy, 6 Heft 16, Fische der Sudsee, j Heft 8, 1910: 486
(distrib.) ; Zugmayer, Abh. bayer. Akad. Wiss., 26 (6), 1913: 8 (Oman).

Alofias vulfes Day, Fish. India, Suppl. 1888: 810 (Ceylon); in Blanford, Fauna Brit. India, i, 1889: 28
(Ceylon); Ogilby, Proc. Linn. Soc. N. S. W., (2) 5, 1889: 1772 (Aust., Tasmania); Gill, Mem. nat.
Acad. Sci. Wash., 6, 1893: ill (N. Zealand); Jordan and Snyder, Annot. zool. jap., 5, 1901: 127
(Japan) ; Jordan and Fowler, Proc. U.S. nat. Mus., 2(5, 1 903: 61 g (Japan) ; Jordan and Snyder, Proc. U.S.
nat. Mus., 21, 1904: 939 (Hawaiian Is.) ; Jordan and Evermann, Bull. U.S. Bur. Fish., 2j (l), 1905: 42
(Honolulu); Stead, Fish. Aust., 1906: 232 (Aust.); Zietz, Trans, roy. Soc. S. Aust., j2, 1908: 291
(Aust.); Thompson, Mar. biol. Rep. Cape Town, 2, 1914: 147 (S. Afr.); Jordan and Jordan, Mem.
Carneg. Mus., 10, 1922: 4 (Hawaiian Is.); Fish Soc. Japan, Ills. Japan. Aquat. Plants Anim., z, 1931:
pi. 7 (ill., Japan).

33. If A. caudatus Phillipps, including greyi Whitley, finally proves identical with vulfinus (see discussion, p.
162), the following references are to be included; if it is proved distinct then some of them may still refer
to vulfinus.

lyS Memoir Sears Foundation for Marine Research

Alofias vulfinus Ogilby, Mem. Qd. Mus., 5, 1916: 74 (Aust.); McCulloch, Biol. Result. Fish. F. I. S.
"Endeavour," .; (4), 1 91 6: 170 (Aust.) ; Waite, Rec. S. Aust. Mus., 2, 1921: 18 (Aust.) ; Phillipps, N. Z.
J. Sci. Tech., 6, 1924: 265 (N. Zealand) ; McCulloch and Whitley, Mem. Qd. Mus., * (2), 1925: 128
(Aust.); Jordan and Hubbs, Mem. Carneg. Mus., 10, 1925: 10 (Japan); McCulloch, Fish. N. S. W.,
1927: 8, pi. 2 (Aust., not seen); Phillipps, N. Z. Mar. Dept. Fish. Bull., I, 1927: 8 (N. Zealand) ; Mori,
J. Pan-Pacif. Res. Instn., 3, 1928: 3 (Korea) ; Fowler, Mem. Bishop Mus., 10, 1928: 18 (Hawaiian Is.,
Fanning I.); Proc. Pan-Pacif. sci. Congr. Java, (4) j, 1930: 488 (distrib.) ; Hongkong Nat., /, 1930:
87 (Hong Kong, China, not seen) ; Phillipps, N. Z. J. Sci. Tech., 13, 1932: 226 (comp. with coudatus) ;
Coppleson, Med. J. Aust., April 15, 1933: 11 (Aust., not dangerous) ; Fowler, Bull. U.S. nat. Mus., 100
(75), 1941: 25 (descr., Indo-Pacif. distrib.).

Alopas caudatus Phillipps, N. Z. J. Sci. Tech., 13, 1 932: 226 (descr., N. Zealand) ; Whitley, Fish. Aust., 7,
1940: 129 (N.Zealand, Aust.).

Alopas gre-ji'^Mx^ey, Rec. Aust. Mus., 20 (l), 1937: 5 (diagn., Aust.) ; Fish. Aust., /, 1 940: 132 (Aust.).

Family ORECTOLOBIDAE
Carpet Sharks, Nurse Sharks

Characters. Two dorsal fins, the ist much shorter than the caudal, its origin over or
posterior to the pelvicsj caudal much less than V2 the total length, not lunate in form, its
lower anterior corner not expanded as a definite lobe, its axis but little raised} inner mar-
gins of pelvics posterior to cloaca either separated, or united for only a very short distance ;
caudal peduncle not greatly flattened dorso-ventrally or expanded laterally, without pre-
caudal pitsj sides of trunk anterior to anal with or without longitudinal ridges } snout not
elongate, nor jaws widely protrusible; 4th and 5th gill openings over base of pectoral; gill
arches without rakers and not interconnected by a sieve of modified denticles; nostril
connected with mouth by a deep groove, its anterior margin with a well developed fleshy
barbel or cirrus;^ spiracles present; lower eyelid without nictitating fold or membrane,
but orbit, in some, with a longitudinal fleshy fold above and below, inside the eyelids, but
entirely free from latter; a labial furrow on each jaw near the corner; teeth small, with
several cusps; head of normal form (not widely expanded) ; rostral cartilages either none,
I, or 3, but separate at tip and very small; mesopterygium of pectoral nearly as large as
metapterygium and with nearly as many radials; mesopterygia and metapterygia sepa-
rated by a foramen; heart valves in 2 rows. Development ovoviviparous in some {Bra-
chaelurus, Orectolobus, Ginglymostoma) , but oviparous in others {Chiloscyllium, Hemi-
scyllium, NehrodeSy Stegostoma), the horny egg capsules of which are attached to algae
either by terminal tendrils or by fibrous extensions of the margin.^

Genera. Most of the members of this large family of warm-water sharks are inhabit-
ants of the western Pacific, Australian region, or Indian Ocean, including the Red Sea;
only one genus {Ginglymostoma) occurs in the Atlantic. Many of them live on bottom in
shallow water, are brilliantly marked, especially when young (hence the common name

1. This is the most striking characteristic of the family as a whole.

2. For illustrations of the egg case of Chiloscyllium, see Southwell and Prashad (Rec. Indian Mus., 16, 1919: 222,
pi. 19, fig. 5) and Whitley (Fish. Aust., 1, 1940: 39, fig. 28, 4.) ; for Stegostoma (Zebra Shark) and Nebrodes
(Tawny Shark), see Whitley (Fish. Aust., /, 1940: 39, fig. 28, 5, 6).

Fishes of the Western North Atlantic 179

Carpet or Zebra Sharks), or are ornamented with fleshy flaps on the head. The majority
are small, but a few grow to a considerable size. Their diet is chiefly bottom dwelling in-
vertebrates and fishes.

Key to Genera

la. Sides of head fringed with fleshy lobes.

2a. A continuous series of branching dermal lobes around lower jaw close to mouth.

Eucrossorhinus Regan, 1908.
East Indies.
2b. No dermal lobes on lower jaw, or at most only a few small ones below chin.

3a. Back smooth, without papillae or tubercles. Or^c/o/o^«j Bonaparte, 1834.

Eastern Pacific, Australia.

3b. Back with rows of papillae or tubercles. 5w/or^f/«j Whitley, 1940.

Australia.

lb. Sides of head without fleshy lobes.

4a. Second dorsal originating posterior to origin of anal.
5a. Throat with a pair of thread-like barbels.

Cirrhoscyllium Smith and Radcliffe, 19 13.

China Sea.'

5b. Throat without barbels. Parascyllium G\\\j i^Gl.

Australia, Tasmania.

4b. Second dorsal originating anterior to origin of anal.
6a. Spiracle minute.

7a. Teeth with central cusp largest and several rows functional.

Ginglymostoma Miiller and Henle, 1837, p. 180.
7b. Teeth with all cusps about equal, only i or 2 rows functional.

A^^^mw Riippell, 1835.

Australasia, Malay Peninsula,
Indian Ocean, Red Sea.*

6b. Spiracle nearly or quite as large as eye.

8a. Caudal more than Ys of total length j ist dorsal originating in front of
pelvics and terminating over latter.

Stegostoma Miiller and Henle, 1837.

Western Pacific, Australasia, Indian Ocean.

3. Whitley (Rec. Aust. Mus., 15, 1927: 289) proposes to substitute a new name, Zev, on the ground that Cirrho-
scyllium Smith and Radcliffe, 191 3, was preoccupied by Cirriscyll'tum Ogilby, 1908, which in turn is a synonym
of Brachaelurus Ogilby, 1907, as pointed out below (footnote 5, p. 180). According to the International Rules
on Zoological Nomenclature, however, this substitution is not required; see recommendation under Article 36
(Proc. bid. Soc. Wash., 39, 1926: 87).

4. Garman's (Mem. Harv. Mus. comp. Zool., 56, 191 3: 56) substitution of Nebrodes for Nebrius Riippell, because
of the earlier use of Nebria by Latreille (1802) for insects and of Nebris by Cuvier and Valenciennes (1830)
for bony fishes, was not necessary, according to the recommendations under Article 36, International Code of
Zoological Nomenclature (Proc. biol. Soc. Wash., jp, 1926: 87).

i8o Memoir Sears Foundation for Marine Research

8b. Caudal considerably less than Ys of total length; ist dorsal originating
over or behind pelvics and terminating behind latter.
9a. Cloaca much nearer to snout than to tip of caudal.

loa. Mouth closer to tip of snout than to front of eye; lower labial
furrow not crossing chin.

Hemiscyllium Andrew Smith, 1837.
Australasia, East Indies, India.

lOb. Mouth closer to front edge of eye than to tip of snout; lower
labial furrow continuous across chin.

Chiloscyllium Miiller and Henle, 1837.

Western Pacific, Australasia, Indian Ocean, Red
Sea, South Africa.

9b. Cloaca as near to tip of caudal as to tip of snout, or nearer.

1 1 a. Rear end of base of anal terminating close to caudal and its
rear tip overlapping latter. Brachaelurus Ogilby, 1907.

Australia. °

lib. Rear end of base of anal separated from caudal by a space twice
as long as the base. HeteroscylUum Regan, 1908.

Australia.'

Genus Ginglymostoma Miiller and Henle, 1837
Nurse Sharks

Ginglymostoma Miiller and Henle, S. B. Akad. Wiss. Berlin, 1837: 113; also Arch. Naturg., (3) i, 1837:
396 (no species given) ; type species, Squalus cirratus Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: 7,
American Seas, designated by Hay, U.S. Geol. Surv. Bull., 179, 1902: 310.

Generic Synonyms:

Squalus (in part) Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: 7; Gmelin, in Linnaeus, Syst. Nat., t,

1789: 1492; also subsequent authors; not Squalus Linnaeus, 1758.
Scyllium (in part) Griffith, in Cuvier, Regne Anim., 10, 1834: pi. 30; for S. cirrhosum Griffith, equals

Squalus cirratus Bonnaterre, 1788.
Ginglimostoma L. Agassiz, Poiss. Foss., 5, 1838: 85; equivalent to Ginglymostoma Muller and Henle, 1837.
Gingylostoma Springer, Proc. Fla. Acad. Sci., j, 1939: 13; evident misspelling for Ginglymostoma.

Generic Characters. Sides of head without fleshy lobes; back without papillae; no
longitudinal ridges on back or sides of trunk, and no longitudinal fold below eye ; eye very
small, the orbit with a longitudinal fleshy fold above and below, whoUy inside the eyelid,
and not connected with latter; spiracle minute, about on a level with eye; 4th and 5th gill

5. The new generic name Brachaelurus was proposed by Ogilby (Proc roy. Soc. Qd., so, 1907: 27) for Chiloscyl-
lium modestum Giinther, 1871 (equals Squalus •waddi Bloch and Schneider, 1801), in which the anal is close
to the caudal; but the name was transferred by him a year later (Proc. roy. Soc. Qd., a/, 1908: 3) to his new
species colcloughi, in which the anal is far from the caudal ; he proposed a new generic name, Cirriscyllium, for
modestum (Proc. roy. Soc. Qd., 21, 1908 : 4). Cirriscyllium Ogilby, 1908, is therefore a synonym of Brachaelurus
Ogilby, 1908, as pointed out by Regan (Ann. Mag. nat. Hist. [8] 3, 1908: 455), who proposed HeteroscylUum
for colcloughi to clarify this confusion.

Fishes of the Western North Atlantic i8i

openings very close together; anterior marginal expansions of nostrils reach to mouth, but
are wide apart; teeth with several cusps, the central much the largest, and several rows
functional; lower labial furrow not continuous across chin; ist dorsal originates over or
slightly posterior to origin of pelvics; 2nd dorsal originates anterior to origin of anal; rear
tip of 2nd dorsal not reaching to origin of caudal; caudal Y^ to Yi of total length; cloaca
about midway between tip of snout and tip of caudal. Development ovoviviparous. Char-
acters otherwise those of the family.

Range. Both sides of tropical and subtropical Atlantic; west coast of Mexico; western
tropical Pacific; Malaysia, Indian Ocean, Red Sea.

Fossil Teeth. Upper Cretaceous, West Indies; Upper Cretaceous to Eocene, Europe;
Eocene, Africa and North America.

Key to Species

la. Corners of fins angular. jerrugineum "Ltsson, i% 2,0-

Western tropical Pacific, Malay-
sia, Indian Ocean, Red Sea.

lb. Corners of fins rounded.

2a. Anal much smaller in area than 2nd dorsal ; nasal barbels reach to mouth.

«rra/«OT Bonnaterre, 1788, p. l8l.

2b. Anal nearly as large in area as 2nd dorsal; nasal barbels reach only about halfway

to mouth. brevicaudatum Gnnthtr, 1866.

Zanzibar, Seychelles.

Ginglymostoma cirratum (Bonnaterre), 1788

Nurse Shark

Figure 29

Study Material. 14 specimens, 275 to 650 mm. long, from Florida, Cuba, Jamaica,
Sombrero I., West Indies, and Panama Bay (Harv. Mus. Comp. Zool.); 2 eggs (about
125 mm. by 63 mm.) and an embryo ( 1 25 mm. long with traces of external gills visible)
from Key West, Florida (Harv. Mus. Comp. Zool., No. 783 and 819) ; female, 936 mm.
long, from Key West, Florida (U.S. Bur. Fish., No. 13927); several specimens fresh
caught, as well as others at liberty, from southern Florida.

Distinctive Characters. The "Nurse" is set apart from all other sharks of the western
Atlantic by the presence of a long barbel on the anterior margin of each nostril and of a
deep groove connecting the nostril with the mouth, together with the terminal position of
the latter. For characters distinguishing it from its several allies in the Indo-Pacific, see
the preceding Keys (pp. 179, 181).

Description. Proportional dimensions in per cent of total length. Female, 650 mm.,
from Cuba (Harv. Mus. Comp. Zool., No. 518).

1 82 Memoir Seats Foundation for Marine Research

Trunk at origin of -pectoral: breadth 15.7; height 10.9.

Snout length in front of: outer nostrils 1.4; mouth 2.5.

Eye: horizontal diameter i.i.

Mouth: breadth 6.S; height 0.6.

Nostrils: distance between inner ends 4.8.

Labial furrow length: upper part 2.6; lower 2.6.

Gill opening lengths: ist2.0; 2nd 2.5; 3rd 2.8; 4th 3. i; 5th 3.4.

First dorsal fin: vertical height 10.5; length of base lo.o.

Second dorsal fin: vertical height 7.1 ; length of base 8.6.

^««/^«; vertical height 4. 8; length of base 6.1.

Caudal fin: upper margin 30.7; lower anterior margin 9.2.

Pectoral fin: outer margin 16.9; inner margin 6.8; distal margin 13.7.

Distance from snout to: ist dorsal 42.4; 2nd dorsal 56.9; upper caudal 69.3;

pectoral 22.6; pelvics 42.7; anal 61.4.

Interspace between: ist and 2nd dorsals 5.4; 2nd dorsal and caudal 3.8; anal and

caudal 1.7.

Distance from origin to origin of: pectoral and pelvics 23.4; pelvics and anal 1 8.9.

Trunk very broad anteriorly, its breadth abreast origin of pectorals about Ve to Vr
of total length, tapering rearward. Caudal peduncle strongly compressed laterally, with-
out lateral ridges or precaudal pits. Dermal denticles closely spaced and more or less

Figure 29. Ginglymostonui cirratum, female, 650 mm. long, from Cuba (Harv. Mus. Comp. Zool., No. 518).
A Anterior part of head of same from below, about 0.4 x natural size. B Dermal denticles, about i 2 x. C Apical
view of dermal denticle, about 24 x. D Upper and lower teeth about 3.3 x. E Newborn male, 283 mm. long,
from Miami, Florida (Harv. Mus. Comp. Zool., No. 33393).

Fishes of the Western North Atlantic 183

overlapping," large (about 0.4 X 0.5 mm. in a specimen 650 mm. long), but varying
much in size, scale-like, their blades ovate, sharp-pointed, or blunted, usually with 3
ridges, the median longest, reaching about halfway to the apexj pedicels high and slender
on broad stellate bases.

Head flattened above (more so in males than in females), widest opposite ist gill
opening in males but opposite 5th gill opening in females. Snout broadly rounded and
very short, its length in front of mouth only about Vs to '/() as great as length of head to
origin of pectoral. Eye oval, about twice as broad as high, its horizontal diameter only
about i/'g as great as distance from eye to ist gill slit. Orbital folds as described above
for the genus (p. 180). Spiracle a minute slit or pore on a level with lower edge of eye
and behind the latter by a distance about i to i ^2 times the horizontal diameter of eye.
Gill openings high on sides and nearly straight, the 3rd over origin of pectoral, the ist to
4th widely spaced, but 4th and 5th very close together, the margin of the former some-
times overlapping and thus concealing the latter in large specimens, the 5th about 1.7 times
as long as the ist and about 3 times as long as the diameter of eye. Nostril nearly longi-
tudinal, its inner (posterior) end connected with front of mouth by a deep, open groove, its
anterior edge outwardly with a tapering, fleshy barbel reaching backward to mouth and
also expanded posteriorly as a subrectangular flap that is continuous across front of mouth
with that of the opposite nostril, and also with the upper lip. Mouth close to tip of snout,
notably small, its breadth a little less than ^3 as great as length of head, its corners with
very deep furrows which form the outline of thick fleshy labial folds on both jaws, the
upper extending inward to edge of nostril and the lower a little further.

Teeth ||-~|^ ; similar in the 2 jaws, with high triangular central cusp flanked on either
side by I to 3 smaller cusps (their number increasing with age of tooth) except when worn
away} cusps progressively smaller and curving outwardly more toward angles of jaw}
7 to 9 series functional in upper jaw and 8 to 1 2 series in lower, in medium-sized specimens.

Fins large, with rounded corners. Origin of ist dorsal over or a little behind origin
of pelvics, its vertical height about % as great as length of pectoral, its anterior margin
slightly convex, its rear margin nearly straight, its free rear tip ^o to % as long as its base
and reaching rearward considerably past tips of pelvics. Second dorsal similar to ist in
shape, and % to $5 as large in linear dimensions, its origin a little anterior to origin of
anal, the distance from its rear tip to origin of caudal about Y^, to % as long as its base.
Caudal a little less than Yz of total length, only a little narrower toward tip than toward
base, its axis very little raised, its tip rounded and slightly bilobed in some specimens but
scarcely so in others, its lower anterior and lower posterior margins nearly straight, except
for the deep subterminal notch, its lower anterior corner obtuse and not expanded as a lobe.
Anal less than half as large in area as 2nd dorsal, although nearly as long as at base, its
rear margin broadly rounded and slightly overlapping the caudal, its origin under or a
little posterior to midpoint of base of 2nd dorsal. Pelvics about % as large in area as ist

6. Evidently there is considerable variation in this respect, for Radcliffe (Bull. U.S. Bur. Fish., j.;, 1916: 249)
shows them as widely spaced.

184 Memoir Sears Foundation for Marine Research

dorsal, with broadly rounded corners. Pectoral about % as broad as long, with broad base,
its outer and inner margins moderately convex, its distal margin nearly straight or very
slightly concave, its corners broadly rounded} about lYo times as large in area in males as
in females.

Color. Rich yellowish to grayish-brown, darker above than below. Small specimens
are usually sparsely and variously marked with small dark spots below as well as above,
sometimes with brown crossbars across the snout and through the dorsals, ventrals and
analj adults may or may not retain these markings; also, some young specimens are plain-
colored.

Size. The Nurse Shark is small at birth, free-living specimens of only 270 to 290
mm.' being recorded, but it grows to a very considerable size, specimens of 7 to lO or 1 1
feet being commonly reported, with 11 to 12 feet not unusual. The maximum so far re-
ported is about 14 feet, but maturity may be attained at a comparatively small size, as in the
case of a female of only 5 feet that contained well developed embryos.* The weight is
given as about 330 to 370 pounds at about 8 V2 feet; 4V4 pounds at 2 feet 3V2 inches (692
mm.).

Develo-pmental Stages. Both ovaries may be functional,' or only one, with the other
atrophied. Mature eggs are very large (reported up to 130 X 180 mm.), blunt-ended,
with brownish-black, thin, horny shells. They remain in the hinder parts of the oviducts
until the shells break and the young are hatched into the uterus.'"

Later in development embryos have a short umbilical cord, with very large sub-
spherical or oval yolk sacs; the external gill filaments are retained up to a length of 130
to 140 mm. The length of the nasal barbel and the size of the eye decrease from embryo
to adult, but the fins increase in relative size. Females have been described as containing as
many as 28 large eggs; a West African specimen (2.43 meters long) has been reported as
giving birth to 26 young on capture."

Habits. In its centers of abundance, from Florida to the Caribbean region, this Shark
appears chiefly inshore, often in water as shallow as two to ten feet. It is frequently en-
countered in channels between the mangrove keys. Schools of one to three dozen are some-
times seen on sand flats and over rocky bottom where they are easily approached, the
sharks often lying motionless and close to one another, with dorsal fins out of water. Pro-
verbially sluggish in habit, the "Nurse" feeds chiefly on invertebrates — squids, shrimps,
crabs, spiny lobsters {Palinurus), sea urchins — and small fish. They bite readily on
almost any bait. It is common knowledge that they come into very shallow water to breed,
and here they are often seen mating. While in the act of copulation the male grasps the

7. See Study Material, p. iSi; also Beebe and Tee-Van, Zoologica, N. Y., lo, 19:8: 26.

8. Beebe, Zoologica, N. Y., 26, 1941 : 9. 9. Bell and Nichols, Copeia, 92, 1921 : 17.

10. For an account of eggs and early development, see Gudger (Yearb. Carnegie Instn., 1912: 11, 149; Copeia, 98,
1921: 57).

11. Budker (Bull. Mus. Hist. nat. Paris, [2] 7, 1935: 183) substantiates the general report that the number of
young in a litter is large.

Fishes of the Western North Atlantic 185

female with his mouth at the edge of her pectoral fin, these fins in females often being
much frayed in consequence." No information is available as to the duration of gestation
or the season when young are born.

Relation to Man. Nurse Shark hides are used to some extent for leather, having at
present a higher value in the Florida fishery than those of other sharks, but the fins are
not in demand. The yield of oil is relatively low." In the West Indies they are sold to
some extent in the fish markets, as are most other sharks. On the islands off the southern
coast of Brazil the liver oil is said to be in high repute, and the otoliths of this species, as
well as those of other sharks, are used by the local fishermen as a diuretic." This shark is
perfectly harmless to bathers, and is too sluggish when hooked to be of any interest to
sport anglers.

Range. Littoral, on both sides of the tropical and subtropical Atlantic; tropical West
Africa and the Cape Verde Islands in the east; southern Brazil to North Carolina and
accidentally to Rhode Island in the west; also west coast of America, from the Gulf of
California to Panama and Ecuador.

Occurrence in the Western Atlantic. The Nurse Shark is very generally distributed
throughout the Caribbean— West Indian region." It is common around Jamaica and
Cuba," and in southern Florida waters among the Keys; and it is a year-round resident
on the west coast north to Tampa and for some distance up the east coast. It is likewise
known from Bermuda. In the warm months it expands its range to the northern coast of the
Gulf of Mexico, and occasional Nurse Sharks are taken near Charleston, South Carolina;
schools of them sometimes appear in summer off Cape Lookout, North Carolina, and one
has been taken in the enclosed waters of Bogue Sound. But this is the boundary to regular
migrations in a northerly direction, the only records of them further north being one stray
individual for Chesapeake Bay, and one for Rhode Island.

To the southward the Nurse Shark probably occurs all along the northeastern coast
of South America, it being known from Maceio in northern Brazil, from Rio de Janeiro,
and from South Trinidad Island off southern Brazil (Lat. 20° 30' S., Long. 29° 22' W.).
There is no report of it farther south. Curiously enough there is but one record of it for
the western shores of the Gulf of Mexico-Caribbean region, that being from Colon,
Panama. But it is to be expected anywhere there, judging from the generality of its dis-
tribution throughout the West Indian region.

Synonyms and References:

Chien de Mer Barbillon, Broussonet, Mem. Math. Phys. Acad. Sci. Paris, 1 780: 656.

Gata, Parra, Hist. Nat., 1787: 86, pi. 34, fig. 2 (descr., Cuba).

Squalus cirratus Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: 7 (descr., American seas); Gmelin, in Lin-

12. See Gudger (Yearb. Carneg. Instn., //, 1912: 149) for a description.

13. Springer, Proc. Fla. Acad. Sci., 5, 1939: 14.

14. Nichols and Murphy, Bull. Amer. Mus. nat. Hist., 53, 1914: 262.

15. Recorded from French and British Guiana, Trinidad, St. Croix, Turks Island, St. Martins and Barbados in the
Lesser Antilles, Jamaica, Haiti, Porto Rico, the Bahamas and Cuba.

16. Personal communication from Luis Howell-Rivero.

1 86 Memoir Sears Foundation for Marine Research

nacus, Syst. Nat., i, 1789: 1492 (descr.) ; Bloch and Schneider, Syst. Ichthyol., l8oi: 128 (descr.,
spelled cirrhatus); Bosc. Nouv. Diet. Hist. Nat., 2/, 1803: 188 (diagn., Pacif.) ; Latreille, Nouv.
Diet. Hist. Nat., 24, 1804: 72 (in table of contents); Bennett, Zool. J. Lond., 5, 1835: 87 (refs.) ;
Bancroft, Zool. J. Lond., 5, 1835:416 (Jamaica).

Squale barbillon, Lacepede, Hist. Nat. Poiss., 4° ed., j, 1798: 166, 245 (diag., descr., Pacif. Amer.) ; in Son-
nini. Hist. Nat. Poiss., 4, 1801-1802: 54 (descr.).

Squalus pittctulatus Lacepede, Hist. Nat. Poiss., 8° ed., 5, 1 800: 153, pi. 4, fig. 3 (Cayenne, descr.) ; Bloch
and Schneider, Syst. Ichthyol., 1 801 : 549 (descr., ref. to Lacepede, 1 800) ; Lacepede, Hist. Nat. Poiss.,
4° ed., 2, 1800: 120, pi. 4, fig. 3, in Buffon, Hist. Nat.

Squdus pinctatus Bloch and Schneider, Syst. Ichthyol., 1801: pi. 134; Bennett, Zool. J. Lond., 5, 1835: 87
(ref.).

Scyllium cirrhosum Griffith, in Cuvier, Anim. Kingd., Lond., 1834: 10, pi. 30 (ill.).

Squalus argus Bancroft, Zool. J. Lond., 5, 1835: 82 (West Indies).

Scyllium cirratum Bennett, Zool. J. Lond., 5, 1835: 87.

Ginglymostoma cirrhosum Bonaparte, Mem. Soc. neuchatel. Sci. nat., 2 (8), 1839: 1 1 (in synopsis).

Ginglymostoma cirratum Miiller and Henle, Plagiost., 1 841 ; 23 (descr., Cayenne) ; Dumeril, Rep. Poiss. Afr.
Occid., 1861: 261 (trop. W. Afr.); Hist. Nat. Poiss., /, 1865: 334 (descr., distrib.) ; Poey, Repert.
Fisico-Nat. Cuba, 2, 1868: 455 (Cuba) ; Gunther, Cat. Fish. Brit. Mus., c?, 1870: 408 (distrib., descr.,
synonyms); Poey, Enumerat. Pise. Cubens., 1876: 204; An. Soc. esp. Hist. Nat., 5, 1876: 400
(Cuba); Goode, Bull. U.S. nat. Mus., 5, 1876: 73 (Bermuda); Lugger, Rep. Coram. Fish. Md.
(1877), 1877: 90 (Chesapeake Bay); Hasse, Naturl. Syst. Elasm. besond. Theil, 1882: 201, pi.
26, fig. 12-14, 16-20, 22-24 (vertebrae); Roehebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882:
47; Faune Senegambie, Poiss., .-, 1883-1885: 24 (Senegambia) ; Goode and Bean, Proc. U.S. nat.
Mus., 5, 1882: 240 (Gulf of Mexico); Jordan and Gilbert, Bull. U.S. nat. Mus., /6, 1883:
18 (Gulf of Mexico and W. Indies); Bull. U.S. Fish Comm., 2, 1883: 105 (Mazatlan, Mexico);
Proc. U.S. nat. Mus., 5, 1883: 371 (Colima, Mexico), 581 (S. Carolina), 620 (Panama); Jordan,
Proc. U.S. nat. Mus., 8, 1885: 363 (Mazatlan, Panama); Rep. U.S. Comm. Fish. (1881;), 1887:
794 (distrib.); Vaillant, Bull. Soc. philom. Paris, (8) 6, 1894: 70 (Gulf of Calif.); Jordan, Stanford
Univ. Pub., Univ. Ser., r, 1895: 381 (Gulf of California); Jordan and Evermann, Rep. U.S. Comm.
Fish. (1895), 1896: 214 (distrib.); Bull. U.S. nat. Mus., 47 (l), 1896: 26 (descr., distrib.);
Jordan and Rutter, Proc. Acad. nat. Sci. Philad., .^p, 1897: 91 (Jamaica) ; Osorio, J. Sci. math. phys. nat.
Lisboa, (2) 5 (19), 1898: 200 (Cape Verde); Boulenger, Boll. Mus. Zool. Anat. comp. Torino, /j
(329), 1898: I (Ecuador); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (4), 1900: pi. 4, fig. 13;
Gilbert, Proc. Wash. Acad. Sci., 2, 1900: 161 (Maceio, Brazil); Evermann and Kendall, Rep. U.S.
Comm. Fish., 1900: 48 (Florida); Gregg, Where to Catch Fish, 1902: 17, fig. 288 (Florida);
Evermann and Marsh, Bull. U.S. Fish Comm., 20, 1902: 60 (W. Indies) ; Werner, Zool. Jb., Syst. Abt.,
21, 1904: 288 (Jamaica) ; Gilbert and Starks, Mem. Calif. Acad. Sci., 4, 1904: 5 (Panama Bay) ; Har-
greaves, Fish. Brit. Guiana, 1904: 14 (Brit. Guiana) ; Jordan and Thompson, Bull. U.S. Bur. Fish., 24,
1905: 232 (Tortugas, Florida); Bean, B. A., in Shattuck, Bahamas Islands Fish., 1905: 296 (size, no. of
eggs, Bahamas) ; Fowler, Proc. Acad. nat. Sci. Philad., 5.?, 1906: 79 (Marquesas Keys, Florid.i, breeding,
large size, color) ; Garman, Bull. Mus. comp. Zool. Harv., 46, I906: 229 (Panama) ; Bean, T. H., Field
Mus. Publ., 7 (2), 1906: 29 (Bermuda) ; Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907: 166, 204 (descr.,
ref. Brazil); Regan, Proc. zool. Soc. Lond., 1908: 350 (descr., distrib., Jalisco, Mexico); Fowler, Proc.
Acad. nat. Sci. Philad., do, 1908: 53 (St. Martins, W. Indies); Vincent, Sea Fish. Trinidad, 1910: 53
(Trinidad) ; Rosen, Lund Univ. Arsskr., N. F. 7, 2 (5), 1 91 1 : 47; Fysiog. Sallsk. Handl., N. F. 22 (5),
191 i: 47 (Bahamas); Roule, Bull. Inst, oceanogr. Monaco, 243, 1912: 5 (C. Verde); Gudger, Yearb.
Carneg. Instn., 11, 1912: 148 (breeding habits, eggs, early devel., Tortugas, Florida); Garman, Mem.
Harv. Mus. comp. Zool., j6, 191 3: 54, pi. 7, fig. 4-6 (descr.) ; Gudger, J. Elisha Mitchell sci. Soc, 29,
1913: 8; Science, 40, 1914: 386 (Florida Keys, abund., habits); J. Elisha Mitchell sci. Soc, 30,
1914: 64; Nichols and Murphy, Bull. Amer. Mus. nat. Hist., jj, 1914: 262 (South Trinidad I., Lat. 20°
30' S., Long. 29° 22' W.) ; Fowler, Proc. Acad. nat. Sci. Philad., 6-j, 1916: 521 (Trinidad); Coles,
Proc. biol. Soc. Wash., 2S, 1915: 89 (C. Lookout, N. Carolina, size); Gudger, Yearb. Carneg. Instn.,

Fishes of the Western North Atlantic 187

14, 1916: 207 (Tortugas, Florida) ; Fowler, Copeia, 30, 1916: 36 (mid. Atlant., U.S.) ; Radclitfe, Buli.
U.S. Bur. Fish., 34, 191 6: 248, pi. 39, fig. l (teeth, Florida, dermal denticles, N. Carolina) ; Metzelaar,
Trop. Atlant. Visschen, 1919: 5 (W. Indies), 186 (w. coast Afr.) ; Fowler, Proc. Acad. nat. Sci. Philad.,
71, 1 91 9: 133 (St. Martins, W. Indies), 146 (Jamaica) ; Nichols, Bull. Amcr. Mus. nat. Hist., 44, 1921 :

22 (Turks I.) ; Nichols, J. Amer. Mus. nat. Hist., 21, 1921 : 275 (ill. adult and young, teeth, no. of eggs,
food); Bell and Nichols, Copeia, 92, 1 921: 17 (N. Carolina, food, size, weight, no. of eggs); Gudger,
Copeia, 98, 1921 : 57 (eggs, devcl., near Key West, Florida) ; Meek and Hildebrand, Field Mus. Pub!., 15
(i), 1923: 29 (descr.. Colon, Balboa, Panama) ; Fowler, Proc. biol. Soc. Wash., 36, 1923: 27 (Tortugas,
Florida) ; Ribeiro, Arch. Mus. nac. Rio de J., Fauna brasil. Pcixes, 2 (i) Fasc. i, 1923:23 (descr., Rio de
Janeiro) ; Fowler, Proc. Acad. nat. Sci. Philad., 78, 1926: 249 (south. Florida) ; Nichols and Breder, Zoo-
logica, N. Y., g, 1927: 1 1 (probably Rhode Island) ; Fowler, Proc. Acad. nat. Sci. Philad., 80, 1928: 451
(embryos, Key West, Florida) ; Hildebrand and Schroeder, Bull. U.S. Bur. Fish., 43, 1928: 45 (general,
Chesapeake Bay); Bccbe and Tee-Van, Zoologica, N. Y., 10 (l), 1928: 26 (size, weight, color, food,
Haiti) ; Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 1 1 (general) ; Jordan, Manual Vert. Anim. NE.
U.S., 1929: 9 (general) ; Nichols, Ann. N. Y. Acad. Sci., 10 (2), 1929: I 81 (Porto Rico) ; Truitt, Bean
and Fowler, Bull. Md. Conserv. Dep., 5, 1929: 29 (Chesapeake Bay) ; Jordan, Evermann and Clark, Rep.
U.S. Comm. Fish. (1928), 2, 1930: 12 (distrib.) ; Manter, Yearb. Carneg. Instn., 29, 1930: 34O; ibid.,
30, 1931:388 (parasites, Tortugas, Florida) ; Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda, 1933:

23 (Bermuda); Fowler, Proc. biol. Soc. Wash., }6, 1933: 27 (Tortugas, Florida); Borodin, Bull. Van-
derbilt Oceanogr. (Mar.) Mus., i (4), 1934: 107 (Miami, Florida) ; Brooks, Parasitology, 26 (2), 1934:
264 (parasites, Tortugas, Florida) ; Breder, Copeia, 1934: 27 (sexual dltfs.) ; Zoologica, N. Y., 18, 1934:
58-59 (Bahamas) ; Budker, Bull. Mus. Hist. nat. Paris, (2) 7, 1935: 183 (Dakar, W. Afr.) ; Burton, Sci.
Mon., N.Y., 40, 1935: 283 (S. Carolina); Fowler, Bull. Amer. Mus. nat. Hist., yo (l), 1936: 36
(descr., trop. Atlant.) ; Norman and Fraser, Giant Fishes, 1937: 28, fig. 16 (general) ; Butsch, J. Barbados
Mus. Hist. Soc, 7(1), 1939: 18 (common at Barbados, not seen) ; Beebe and Tee-Van, Zoologica, N. Y.,
26 (2), 1941: 96 (west coasts Mexico, Costa Rica, Panama); Beebe, Zoologica, N. Y., 26 (i), 1941: 9
(embryos, Bermuda); Hildebrand, Copeia, 1941: 221 (recent N. Carolina records); Longley and
Hildebrand, Yearb. Carneg. Instn., 34, 1 941: II (Tortugas, Florida); Norris, Plagiost. Hypophysis,
1941: 27, pi. 6, fig. 25 (brain)) Fowler, Fish Culturist, 21 (9), 1942: 66 (listed, Cuba); Lunz, Bull.
S. C. St. Planning Bd., 14, 1 944: 26 (S. Carolina, Florida).

GinglymostoTna fulvum Poey, Memorias, 2, 1861: 342, pi. 19, fig. 15, 16 (teeth, Havana, Cuba); Repert.

Fisico-Nat. Cuba, 2, 1868: 455, pi. 4, fig. 12-14 (teeth, Cuba); Enumerat. Pise. Cubens., 1876: 204;

An. Soc. esp. Hist. Nat., 5, 1876:400 (Cuba, Brazil) ; Stahl, Fauna Puerto Rico, 1883: 167 (Porto Rico) ;

Jordan, Proc. U.S. nat. Mus., 8, 1885: 363 (west coast Cent. Amer., doubtful species); Bigelow and

Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 98 (descr., ill.,

range).
GinglymostoTna cabozierdiauus Brito Capello, J. Sci. math. phys. nat. Lisboa, i (2), 1867: 167 (C. Verde).
Ginglymostoma cirrhatum Jordan, Guide to Study Fish., i, 1905: 533 (general).

GinglymostoTna cirrotunC-'' Gudger, Yearb. Carneg. Instn., 12, 1914: 176 (abund. near Key West, Florida).
Gingylostovu^^ cirratum Springer, Proc. Fla. Acad. Sci., 5, 1939: 13 (descr., Florida).
Nebrius cirratum Fowler, Arqu. Zool. Estado Sao Paulo, 3, 1942: 127 (Brazil) ; Mongr. Acad. nat. Sci. Philad.,

7, 1945: 94 (sizes, weights, N. Carolina).

Family RHINCODONTIDAE

Whale Sharks

Characters. Two dorsal fins, the ist much shorter than the caudal, its origin in front
of origin of pelvics, but its base overlapping the latter j 2nd dorsal and anal much smaller
than 1st dorsal} caudal much less than Y2. total length, lunate in form, its axis steeply

17. Evident misspelling for a'rra/am. 18. Evident misspelling for Ging/ymo^^oma.

1 88 Memoir Sears Foundation for Marine Research

raised; caudal peduncle not greatly expanded laterally, with a precaudal pit above but
none below; sides of trunk anterior to anal with longitudinal ridges; snout very short and
mouth not widely protrusible; gill openings very large, the 4th and 5th over base of pec-
toral; gill arches connected, one with the next, by numerous transverse cartilaginous bars
which support soft spongy masses of tissue developed from clumps of modified denticles,
the entire gill apparatus forming a sieve of innumerable minute meshes ( i to 2 X 2 to 3
mm. in specimen 31 ft. 6 in.) through which water is forced when the mouth is closed, the
planktonic food thus being retained and swallowed;' oesophagus lined with large papil-
lae, covered with denticles; nostril entirely separate from mouth,^ its anterior margin
without barbel; spiracles present; lower eyelid without nictitating fold or membrane; each
jaw with a labial fold near the corner; teeth minute, very numerous, many rows func-
tional; head of normal shape (not widely expanded); no rostral cartilages; no foramen
between mesopterygium and metapterygium of pectoral' (relative number of radials on
meta- and mesopterygia not known); heart valves in 2 rows. Development probably
ovoviviparous (see p. 192).

Genera. Rhincodon^ the only known representative of the family, has sometimes been
associated with the Orectolobidae (e.g., by Regan*) on the supposition that its nostrils are
connected with the mouth by oronasal grooves. Recently, however, it has been found that
this is not the case;^ and it is so widely separated from the Orectolobidae in other respects,
especially by its complex gill sieve and its lunate caudal with sharply raised axis, that it
clearly represents a distinct family.

Only one genus known, Rhtncodon.

Genus i?/2/'«coi^o« Smith, 1829
Whale Sharks

Rhlncodon Smith, Zool. J., 4, 1 829: 443; type species, Rhtncodon tyfus Smith. Table Bay, South Africa.

Generic Synonyms:^

Rineodon Muller and Henle, Charlesworth's Mag. Nat. Hist., 2, 1838: 37; no species mentioned/

Rhineodon Swainson, Nat. Hist. Fish. Amphib. Rept., r, 1838-1839: 142.

Rhiniodon Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1838-1839: 317.

Rhinodon Muller and Henle, Plagiost., 1841: 77; type species, "Rhinodon tyficus Smith"; evident misquo-
tation for Rhtncodon tyfus Smith, 1829.

Rhineodon Gray, List Fish. Brit. Mus., I, 1857: 66; Rhineodon tyficus Gray, equals Rhtncodon tyfus Smith,
1829.

Micristodus Gill, Proc. Acad. nat. Sci. Philad., 1865: 177; type species, M. functatus Gill. Gulf of California.

I. For photographs of this gill structure, unique among sharks, see White (Bull. Amer. Mus. nat. Hist., y^, 1937:

pi. 9, 10) and Gudger (J. Morph., 6S, 1941 : 91-95, fig. 6-8).
I. It has been stated repeatedly that the nostril is connected with the mouth by an oronasal groove (for example,

see Garman, Mem. Harv. Mus. comp. Zool., $6, 1913: 41) ; it has been found recently that such is not the case

(Barnard, Ann. S. Afr. Mus., jo, 1935: 647, pi. 25).

3. See White (Bull. Amer. Mus. nat. Hist., 61, 1931: 130) for account of skeletal characters.

4. Proc. zool. Soc. Lond., 1906: 745.

5. Most of the generic synonyms here listed are simply emended spellings of Rhtncodon Smith, 1829.

Fishes of the Western North Atlantic 189

Cetorhinus Poey, An. Soc. csp. Hist. n.it., 5, 1876: 184 (380) ; for C. maximus Poey (Cuba), which equals

Rhincodon tyfus Smith, 1829; not Cetorhinus Blainville, 1816.
5«/(j^/i« Thomas, Cannibals and Convicts West Pacif., 1887: 38; ior S . maxima Thoma.$ (New Guinea), which

equals Rhincodon tyfus Smith, 1829; not Selache Cuvier, 1817.

Generic Characters. Those of the family (p. 187).

Range. Tropical belts of all oceans.

Species. One species only, R. typus Smith, 1829.

Rhincodon typus Smith, 1829

Whale Shark

Figure 30

Study Material. Excellent mounted specimen, 17 feet 4^4 inches long, from Aca-
pulco, Mexico (Amer. Mus. Nat. Hist.) j dried skin and dental plates of 18-foot specimen
from Ormond, Florida (U.S.Nat. Mus., No. 27618)5 also photographs of a newly caught
Cuban specimen from Luis Howell-Rivero.

Distinctive Characters. Distinguished from all other sharks by its enormous size,
spotted color pattern, lunate tail, very wide gill openings, unique gill apparatus (see p.
188), and mouth at tip of snout.

Description. Proportional dimensions in per cent of total length from mounted speci-
men; female, 5,302 mm. (17 ft. 4% in.) from Acapulco, west coast of Mexico (Amer.
Mus. Nat. Hist., New York).

Trunk at origin of pectoral: breadth 18.5; height 14.2.

Snout length in front of: outer nostrils l.O ; mouth 0.5.

Eye: horizontal diameter 0.5.

Mouth :hr&'a.dth. 16.3; height O.

Nostrils: distance between inner ends 10.8.

Gill opening lengths: ist 8.8; 2nd 9.3; 3rd 8.9; 4th 8.0; 5th 6.6.

First dorsal fin: vertical height 6.9; length of base 8.5.

Second dorsal fin: vertical height 2.8 ; length of base 3.2.

Anal fin: vertical height 2.7; length of base 2.7.

Caudal fin: upper margin 26.4; lower anterior margin 1 1.8.

Pectoral fin: outer margin 15.9; inner margin 4.4; distal margin 14.4.

Distance from snout to: ist dorsal 39.7; 2nd dorsal 62.1 ; upper caudal 73.6; pec-
toral 18.O; pelvics 46. 1; anal 63.O.

Interspace between: ist and 2nd dorsals 13.9; 2nd dorsal and caudal 8.3; anal and

caudal lo.o.

Distance from origin to origin of: pectoral and pelvics 28.3 ; pelvics and anal 16.8.

Trunk moderately stout, each side with 2 prominent dermal ridges originating close
together high on the shoulders, the lower one extending backward the whole length of
trunk and as a keel out onto anterior part of caudal, the upper one dividing anterior to the

Fishes of the Western North Atlantic igi

origin of the ist dorsal, with both its branches terminating just in front of, under, or well
behind, the 2nd dorsal} also a ridge along midline of the back from the rear part of head
to 1st dorsal in some specimens, but seemingly not in others. Caudal peduncle with a pre-
caudal pit above but none below. Dermal denticles very small (less than i mm. broad in
specimen 31 ft. 5 in. long), slightly overlapping, scale-like, their blades varying in shape,
but usually with 3 to 5 ridges, the axial ridge very high, their margins with 3 to 7 teeth,
the median much the longest, on relatively high pedicels."

Head strongly flattened above, its dorsal profile concave anterior to gill openings
and broadly rounded in front. No distinct snout, the mouth being at the anterior margin
of the head. Eye minute, its horizontal diameter less than ^ 0 as great as length of nostril,
its center situated a little posterior to angle of mouth. Spiracle about as large as eye, a short
distance behind and above the latter. Gill openings notably large but high up on the sides
and widely separated ventrally, the 3rd-4th over origin of pectoral, the 2nd in front of
pectoral and extending below it, the 2nd and 3rd longest. Nostrils at front margin of head,
widely separated, the space between them about % as wide as mouth, their anterior margins
without barbels, but each expanded as a broad, quadrilateral lobe with rounded corners,
overlapping the upper lip, entirely separate from mouth but with outer end continuous
with upper labial furrow. Mouth very large, its width nearly as great as breadth of head,
transverse, hardly arched. Upper labial furrow extending so far that it joins outer end of
nostril,' the lower furrow hardly extending beyond corner of mouth.

Teeth similar in the 2 jaws, minute (averaging about 1.5 mm. long in an 18-foot
specimen, but about 4.5 mm. in a 31-foot fish), in about 310 rows, with 10 to 15 rows
(average about 12 to 13) functional all along the dental band, or a total count of around
3,600 teeth in an 18-foot specimen (Fig. 30) and perhaps still more in larger speci-
mens} the rows vertical toward center of mouth but somewhat oblique toward corners}
each tooth with a single sharp cusp curved backward.*

First dorsal nearly an equilateral triangle, of moderate size, its vertical height a little
less than Vs as great as length of head, its anterior margin nearly straight, posterior mar-
gin moderately concave, apex rounded, its free rear tip triangular, a little shorter than the
base, its origin considerably in front of origins of pelvics with the rear end of its base about
over rear ends of bases of pelvics. Second dorsal about V3 as large in area as ist and similar
in form, but with its free rear tip a little longer than its base} its origin considerably
anterior to that of anal. Caudal noticeably large, its posterior margin broadly concave in
subangular outline, its upper lobe about 22% of total length, its lower lobe about 45% as
long as upper, the tips of both lobes pointed, the upper without subterminal notch. Anal
nearly as large as 2nd dorsal and similar in shape but with shorter free rear tip, its origin

6. See White (Bull. Amer. Mus. nat. Hist., 6i, 193 1 : 144) for account of the variation in shape of the denticles.

7. This is shown very clearly in Barnard's (Ann. S. Afr. Mus., 30, 1935: pi. 25) photograph of a newly captured
specimen.

8. For excellent photographs of teeth, see Bean (Smithson. misc. Coll., ^8, 1905: pi. 36), and White (Bull. Amer.
Mus. nat. Hist, 61, 1931: pi. 10, 11); for additional counts, see Mowbray (Prelim. Rep. Sci. Cruise "Nour-
mahal," 1933: 2).

ig2 Memoir Sears Foundation for Marine Research

about under rear end of base of 2nd dorsal. Pelvics notably small, being only about as
large as anal. Claspers of male of usual galeoid type. Pectoral about Ve to Vr as long as
total length, its distal margin moderately concave, its apex subacute.

Color. Described as varying from dark gray to reddish or greenish brown on back and
sides, including upper surface of pectorals, and marked with round white or yellow spots
(2 to 3 inches in diameter in 38-foot specimen), these being smallest and most crowded
on the head, largest and most scattered rearward} also a variable number of narrow white
or yellow transverse stripes; lower parts plain white or yellow; lips, tongue and lining
of mouth whitish ; lining of oesophagus black.

Size. This is the largest of modern fish-like vertebrates ; specimens so far measured
have ranged between 6° and about 45°" feet in length, with 6-34 feet recorded for Cuban
specimens, 18-34 feet for Florida examples, and 31.5 feet for one taken at Fire Island,
New York. But the Whale Shark is creditably reported to reach lengths of 60 feet or even
more. The estimated weight of a 3 8 -foot Whale Shark, taken at Knight's Key, Florida,
in June 1912 was 26,594 pounds.'" The size at which sexual maturity is attained is not
known, nor is the size at birth.

Developmental Stages. Sixteen eggs have been counted in a specimen from Ceylon,
these being of the "same form as in dog fish.'"' Whether or not these hatch before birth
is not definitely known.

Habits. Notwithstanding the extensive literature regarding the Whale Shark that
has developed in the past few years, much of which is repetitious, very little is known of
its habits, other than that it gathers in schools, often basks at the surface and is so sluggish
and so little alert that specimens are rammed by steamers from time to time. It feeds by
gulping mouthfuls of small animals, as does the Basking Shark; it then drives out through
its branchial sieve the water that it takes with them into its mouth. It has been seen at the
surface with open mouth when so employed, swimming or even vertical in the water. Its
diet may be either small Crustacea, as in the case of a Galapagos specimen where 98 per
cent of the stomach contents consisted of such," or perhaps more often small fish. Cuban
fishermen, for example, describe it as gorging on schools of anchovies, sardines and
albacores, apparently standing vertical below the school while feeding. It is also known
to devour small squids when they are available. But the fact that a Philippine specimen
had in its stomach 47 buttons, 3 leather belts, 7 leggings and 9 shoes'' is evidence that it
is not very discriminating, if the individual in question actually was a Whale Shark.'*

Relation to Man. The Whale Shark has been the object of a fishery along the north-

9. Personal communication from Luis Howell-Rivero.

9a. Seychelles Is., Indian Ocean, reported by Wright (Spicil. Biol., Dublin, 1870: 64-65 [not seen]).

10. Field and Stream, Feb. 1936: 27. The length of this specimen was originally given as 45 feet (Garman, Mem.
Harv. Mus. comp. Zool., ^6, 1913: 456) but was later found to be only 38 feet (Gudger, Proc. Zool. Soc.
Lond., 1934: 881).

n, Southwell, Ceylon Adm. Rep. Mar. Biol., 1912: E. 44, E. 49.

12. Mowbray, Prelim. Rep. Sci. Cruise "Nourmahal," i, 1933: 2.

13. Jordan, Science, N. S. 36, 1915: 463.

14. Gudger (Amer. Nat., 75, 1941 : 550) suggests that it may have been some other shark.

Fishes of the Western North Atlantic 193

west coast of India, but elsewhere it is of no commercial importance. It is entirely harmless
to bathers or small boats, unless by accidental contact.

Range. Pelagic in tropical belts of all oceans. Reliable reports of it are from South
Africa (type locality). Red Sea and Straits of Bab-el-Mandeb, Seychelles, west coast of
India, Ceylon, Bay of Bengal, many localities in the Malaysian-Papuan region, Philip-
pines (a center of abundance), southeastern Australia, Indo-China, Gulf of Siam, Bonin
Islands, Japan, Paumotos, Gulf of California (especially numerous near Cape St. Lucas),
west coast of Mexico (taicen frequently at Acapulco), Panama-Galapagos region, coast of
Peru, in the Indo-Pacificj South Africa, Gulf of Guinea, Brazil, Caribbean-West Indian
region, Florida, and casually to New York in the Atlantic. Up to the present time, the most
northerly locality for it is about 42° North Latitude (near New York), the most southerly
locality 33° SS' South (Table Bay, South Africa).

Occurrence in the Western Atlantic. Records of the Whale Shark in the western At-
lantic are distributed as follows, from south to north : Abrolhas Island, Brazil, Lat. 17° 15'
S. (one) j western Caribbean, between Colon and Cartagena (one) ; Haiti (one) ; around
Cuba J '^ central part of the Gulf of Mexico (single individuals or schools reported on eight
occasions) j Bahamas (a school in Tongue of the Ocean, and one at Bimini) ; Gulf Stream
between Bahamas and Florida j southern and eastern Florida (five) ; mouth of Cape Fear
River, North Carolina (one) ; south shore of Long Island, near New York (one). From
the foregoing, the Caribbean-West Indian region is evidently the center of population
for it on this side of the Atlantic. Occasional captures on the coasts of North Carolina and
New York show merely that Whale Sharks, like other tropical animals, occasionally stray
far northward beyond their normal range in the warm months. There is one report of it
from Bermuda."^ The fact that it is sometimes reported in schools prevents estimation of
the numbers actually reported up to the present.

Synonyms and References:

Rhincoion fjpis Smith, Zool. J., 4, 1829: 443 (Table Bay, S. Afr.) ; Garman, Mem. Harv. Mus. comp. Zool.,
j(5, 1913: 42, 456 (descr., large Florida specimen); Herre, Philipp. J. Sci., 2(5, 1925: 116 (Philip-
pines); Fowler, Bull. U.S. n.it. Mus., lOO (z^), 1941: 116 (descr., synonyms, distrib.) ; Fowler, Bol.
Mus. Hist. N. Javier Prado, Lima, 5 (17), 1941: 220; Fowler, Fish Culturist, 21 (9), 1942: 66 (listed
Cuba); Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo .■\mer. Caribb. Comm., Wash., 194;:
96 (descr., ill., range).

Rhineodon (no specific name) Swainson, Nat. Hist. Fish. Amphib. Rept., /, 1838-1839: 142 (general); 2:
191, 314, 317^° (general); Gudger, Nat. Hist. N. Y., 22, 1922: 246 (a shark sucker, Remora, in its
mouth); Smith, Science, 62, 1925: 438 (Siam); Gudger, Nature, Lond., 131, 1933: 165 (Ceylon);
Denison, Bull. Amer. Mus. nat. Hist., 75, 1930: 477 (skelet.) ; Gudger, Nature, Lond., 143, 1939: 79
(feeding, Bahamas) ; Sci. Mon., N. Y., 48, 1939: 261 (Haiti, Gulf of Mexico).

Rhinodon fjficus Muller and Henle, Plagiost., 1841: 77, pi. 35, fig. 2 (descr.); Smith, 111. Zool. S. Afr.
Pisces, 1849: pi. 26 (descr., meas., color, ill. most often copied) ; Dumeril, Hist. Nat. Poiss., i, 1865: 428
(descr.) ; Gunther, Cat. Fish. Brit. Mus., 8, 1 870; 396 (descr.) ; Guide to Study Fish., I 880: 323 (gen-
era!) ; Haly, Ann. Mag. nat. Hist., (5) /2, 1883: 48 (Ceylon, meas.); Gunther, Nature, Lond., 30,

15. "Caught quite often in open waters, and seen many times feeding" (person.i! communication from Luis Howell-
Rivero) ; also five published records.

15a. Personal communication from John Tee-Van.

16. Spelling on page 317 is "R/iiniodon."

194 Memoir Sears Foundation for Marine Research

1884: 365 (Gulf of Panama) ; Chierchia, Coll. Sci. Nat. viaggio . . . Vett. Pisani (1882-1885), 1885: 68
(Bay of Panama) ; Day, Fish. India, suppl., 1888: 811 (Ceylon); Day, in Blandford, Fauna Brit. India,
Fishes, /, 1889: 29 (general) ; Thurston, Bull. Madras Govt. Mus., 1 894: 36-38, pi. 3 (descr., Ceylon,
Madras); Casto de Elera, Cat. Fauna Philipp., 1895: 615 (Philippines); Gill, Science, 15, 1902: 824
(history) ; Bean, B. A., Science, 75, 1902: 353 (Ormond, Florida) ; Bridge, Camb. Nat. Hist., 7, 1904:
287 (general) ; Bean, B. A., Smithson. misc. CoU., 48, 1 905: 139, pi. 34-36 (general, photos) ; Weber-
Van Bosse, Ein Jahr am "Siboga," Leipzig, 1905: 220 (Malaysia); Wilson, Proc. U.S. nat. Mus.,
33t 1907: 439 (parasites); Van Kampen, Natuurk. Tidschr. Ned.-Ind., 67, 1908: 124 (Batavia,
Java); Fountain, Rambles Aust. Nat., 1 907: 119 (Aust., not seen); Lloyd, Rec. Indian Mus., 2,
1908: 306 (Bay of Bengal); Regan, Proc. zool. Soc. Lond., igo8: 353 (general); Gunther, J. Mus.
Godeffroy, 6 Heft 17, Fische der Sudsee, 3 Heft 9, 1910: 486 (size); Smith, Proc. biol. Soc. Wash.,
25, 191 1 : 97 (Philippines, size) ; Gunther, Encycl. Brit., nth ed., 24, 1911 : 808 (general) ; Southwell,
Ceylon Adm. Rep. Mar. Biol., 1919-1923: E44, E49 (Ceylon, eggs); Weber, "Siboga" Exped., 57,
1 9' 3- 593 (Malaysia); Jordan, Science, N. S. 41, 1 91 5: 463 (Philippines, exceptional diet); Smith, J.
Amer. Mus. nat. Hist., 16, 1916: 352 (distrib., habits); Barnard, Ann. S. Afr. Mus., 2/ (1), 1925: 37
(descr., S. Afr.) ; Pillay, J. Bombay nat. Hist. Soc, 55, 1929: 351 (India, not seen) ; Tanaka, Jap. Fish.
Life Colors, 24, 1933 (Japan, not seen).

Great Basking Shark, Buist, Proc. zool. Soc. Lond., 18, 1850: lOO (fishery, Karachi, India).

Khineodon tyficus Gray, List Fish. Brit. Mus., 1851:67 (descr.) ; Jordan and Fowler, Proc. U.S. nat. Mus., 2<5,
1903: 626 (ref.) ; Jordan, Guide to Study Fish., /, 1905: 540 (general); Grey, Fishing Virgin Seas,
1925: 204—216 (Lower California, not seen); Jordan, Evermann and Clark, Rep. U.S. Coram. Fish.
(1928), 2, 1930: 18 (distrib.); Gowanloch, Bull. La. Conserv. Dep., 23, 1933: 223, 224, 229 (no
Louisiana record); Howell-Rivero, Proc. Boston Soc. nat. Hist., 41, 1936: 44 (Cuba); Springer, Proc.
Fla. Acad. Sci., 5, 1939: 33 (Florida); Hildebrand, Copeia, 1941: 222 (N. Carolina); Lunz, Bull. S.
Carolina St. Planning Bd., 14, 1944: 27 (Florida).

Micristodus functatus Gill, Proc. Acad. nat. Sci. Philad., 1865: 177 (Gulf of California) ; Jordan and Gil-
bert, Bull. U.S. nat. Mus., 16, 1883: 32 (Gulf of California) ; Jordan, Rep. U.S. Coram. Fish. (1885),
1887: 798 (in list) ; Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 218 (Gulf of Cali-
fornia) ; Bull. U.S. nat. Mus., 47 (i), 1896: 52 (descr.. Lower California); Gunther, Encycl. Brit.,
nth ed., 24, 1911: 808 (general); Mowbray, Prelim. Rep. Sci. Cruise "Nourmahal," 1933: 2 (descr.,
no. of teeth, color, size, stomach content).

Cetorhinus maximus Poey, Enumerat. Pise. Cubens., 1876: 184; Ann. Soc. esp. Hist, nat., 5, 1 876: 380
(Cuba).

Selache Tnaxima Thomas, Cannibals Convicts West. Pacif., 1887: 38 (New Guinea, not seen).

Rhitwdon typis Sauvage, in Grandidier, Hist. Phys. Nat. Madagascar, Poiss., 1 891: 511 (Madagascar);
Roxas and Martin, Dept. Agric. Comm. Manila, Tech. Bull. 6, 1937: 12 (Philippines, not seen).

Rhinodon fentaUneatus Kishinouye, Zool. Anz., 24, 1901: 694 (labial folds, descr., Japan); Zool. Mag.
Tokyo, i^ (172), 1903: 41 (Japan).

Rhlneodon typus Gill, Science, 21, 1905: 790 (Indian fishery, habits); Gudger, Science, 38, 1913: 270
(Florida); Fowler, Proc. Acad. nat. Sci. Philad., 6j, 1915: 245 (Knights Key, Florida); Gudger, Zoo-
logica, N. Y., /, 1915: 349 (Knights Key, Florida); Science, 48, 1918: 623; Science, 52, 1920: 192
(general) ; Science, 56, 1922: 251 (Florida record) ; Science, 58, 1923: 180 (Florida) ; Nat. Hist. N. Y.,
25, 1923: 62 (Brazil); Beebe, Arcturus Adv., 1926: 414 (Galapagos, not seen); Gudger, Zoologica,
N. Y., JO, 1927: 76 (trop. W. Afr.); Science, 65, 1927: 211 (Gulf of California); Science, 65,
1927: 545 (Galapagos); Gudger and Hoffmann, Araer. Mus. Novit., 318, 1928: I (Florida Straits);
Fowler, Mem. Bishop Mus., 10, 1928: 18 (ref.) ; Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 23
(general); Gudger and Mowbray, Nat. Hist. N. Y., 40, 1930: 182 (Florida); Gudger and Hoffmann,
Science, yo, 1930: 639 (near Havana, Cuba) ; White, Bull. Amer. Mus. nat. Hist., 61, 1930: 129 (anat.,
denticles, teeth, skelet.) ; Gudger, Bull. Amer. Mus. nat. Hist., 61, 193 1 : 613 (Florida, size); Gudger
and Hoffmann, Sci. Mon. N. Y., 3 2, 1 93 1 : 3 3 (near Havana) ; Gudger, Nature, Lend., 130, 1932: 1 69
(Seychelles); Science, y^, 1932: 412 (5th Florida record); Herre, Science, ys, 1932: 413 (N.

Fishes of the Western North Atlantic 195

Borneo); Pearson, Nature, Lend., 131, 1933: 729 (Ceylon); Gudger, Nature, Lond., 132, 1933:
569 (Galapagos); Bertolini, R. C. Accad. Lincei, (6) 18, 1933: 235 (teeth in relation to food); Boll.
Soc. Biol. Sper., 9, 1934: 1271 ; Barnard, Nature, Lond., 134, 1934: 66 (C. of Good Hope) ; Delsman,
Nature, Lond., 133, 1934: 176 (Str. Bab-eI-M,indeb) ; Gudger, Hydrogr. Bull. Wash., 2362, Dec. 12,
1934 (general); Barnard, Ann. S. Afr. Mus., 30, 1935: 647, pi. 23-25 (dcscr., meas., discuss., photos,
S. Afr.) ; Hcrre, Science, 81, 1935: 253 (N. Borneo) ; Brimley, J. Elisha Mitchell sci. Soc, 5/, 1935:
160 (C. Fear R., N. Carolina) ; Marchand, Fish. Mar. Biol. Surv. Union S. Afr., Fish. Bull. 2, 1935: 28
(S. Afr.) ; Gudger, Nat. Hist. N. Y., 36, 1935: 128 (general, Florida records) ; Proc. zool. Soc. Lend.,
1935: 863 (records up to end of 1934) ; Gudger and Smith, Bull. N. Y. zool. Soc, 38, 1935: 71 (sizes,
photos, Acapulco, Mexico) ; Gudger, Science, 84, 1936: 227 (Bimini, Bahamas) ; Nat. Hist. N. Y., 57,
1936: 159 (Fire Island, N. York) ; Sci. Mon. N. Y., 42, 1936: 84 (photos, near Havana, Cuba) ; Beebe,
Bull. N. Y. zool. Soc, 3g, 1936: 241-242 (photos, Gulf of California) ; Chevey, Note Inst. Oceanogr.
Indochine, 28, 1936: 3 (Indo-China) ; Suvatti, Index Fish. Siam., 1936: 2 (Gulf of Siam) ; Howell-
Rivero, Mem. Soc. cubana Hist, nat., 11 (2), 1937: 109, pi. 14, 15 (descr., meas., photos, Cuba); White,
Bull. Amer. Mus. nat. Hist., 74, 1937: 29, 41, 59, 61, 64, 73, 79, 90, 91, no, pi. 4, fig. 1, pi. 9 to 11,
pi. 12, fig. e, pi. 21, pi. 29, fig. m, pi. 36, pi. 37, pi. 46, fig. c (anat., photos of gills) ; Gudger, Science,
85, 1937: 314 (general) ; Nature, Lond., 159, 1937; 549 (Ceylon) ; Copeia, 1937: 60 (W. Caribbean) ;
Norman and Fraser, Giant Fishes, 1937: 30, pi. 2 (general); Gudger, Nature, Lond., 141, 1938: 516
(off Pacific Panama) ; Copeia, 1938: 172 (rammed by steamers. Red Sea, Gulf of Aden, Ceylon) ; Calif.
Fish Game, 24, 1938: 420 (Lower California); Beebe, Zaca Venture, N. Y., 1938: 162-170 (Gulf of
California, not seen) ; Gudger, Sci. Men., N. Y., 50, 1940: 225 (size, Fire I., N. York) ; Beebe and Tee-
Van, Zoologica, N. Y., 26, 1941 : 97, pi. l (photo. Gulf of California) ; Gudger, J. Morph., 68, 1 941:
81 (feeding, teeth, gill apparatus, photos) ; J. Elisha Mitchell sci. Soc, ^y, 1 941: 57 (N. Carolina speci-
men); Amer. Nat., y5, 1941: 550 (general, photo); Deraniyagala, J. Bombay nat. Hist. Soc, 44 (3),
1944: 427 (photos, Ceylon).

Family SCYLIORHINIDAE'

Cat Sharks

Characters. Two (rarely only one) dorsal fins, the ist much shorter than the caudal,
at least y-i of its base posterior to origin of pelvicsj caudal much less than V2 of total
length, not lunate in form, its lower anterior corner not expanded as a definite lobe, its
axis but little raised 5 caudal peduncle not greatly flattened dorso-ventrally or expanded
laterally; sides of trunk anterior to anal without longitudinal ridges; no precaudal pits,
at least in most species; inner margins of pelvics more or less united posterior to cloaca;
snout not greatly elongate or jaws widely protrusible; 5th, or 4th and 5th, gill open-
ings over origin of pectoral; gill arches without rakers and not interconnected by a sieve
of modified denticles; nostril not connected with mouth by a groove, or if so connected,
its anterior margin does not bear a well developed fleshy barbel; no nictitating membrane
within lower eyelid, but there may be a well developed longitudinal fold below the latter;
spiracles present; labial furrows more or less developed; teeth small, numerous, with
several cusps, and several rows functional; head of normal shape, not widely expanded
laterally; rostral cartilages 3, united at dp; radials of pectoral mostly on metapterygium ;

I. We include White's (Bull. Amer. Mus. nat. Hist., 74, 1937: 107, 108) Halaeluridae and Atelomycteridae under
the Sc^-liorhinidae, the differences in vertebral calcification on which they were based not seeming sharply enough
alternative to warrant the rank of families.

196 Memoir Sears Foundation for Marine Research

mesopterygium much smaller, with few radials; meso- and metapterygia separated by a
foramen, or not ; vertebral calcifications widely variable in type j heart valves In 2 or 3 rows.
Development oviparous so far as known.

The family includes numerous species of small sharks In tropical and temperate lati-
tudes, from both shoal water and deep. Although it embraces two of the most common
and best known of the European sharks," the centers of abundance for both genera and
species are the western Pacific, Australasian region and Indian Ocean to South Africa. It is
represented in the western North Atlantic by only a few little known deep water species.

Genera. Opinions have differed widely as to the number of genera deserving recog-
nition In this family. At the one extreme Carman' recognizes eleven, a list to which no
less than seven more genera or subgenera have subsequently been added by Fowler* and
Whitley." At the other extreme Barnard" unites In a single genus the ten South African
representatives of the family, which would fall in some seven different genera under the
contrasting scheme. An intermediate view Is taken by Norman, who suggests the recogni-
tion of "some four natural groups as genera."^

Generic characters In so uniform a family must be based on definitely alternative
and easily discernible characters to be of any value to working Ichthyologists. For Instance,
one group of some nine recognizable species is set apart from all other members of the
family by the fact that the denticles along the dorsal margin of the anterior part of the
caudal are not only enlarged but modified In shape and directed laterally so as to form
a definite crest, which is outlined below by a narrow band of naked skin.' The mem-
bers of this group fall into two categories: one with the posterior margin of the nos-
tril widely expanded, the snout short and thick and the body cavity longer; the other
with the posterior margin of the nostril expanded little, if at all, the snout long and
thin and the body cavity shorter. These characters seem sufficiently alternative for the
retention of the genus Parmaturus Carman, 1906, for the first group, as distinct from
Galeus Rafinesque, 18 10, for the second. But Whitley's segregation of some members
of the latter into a separate subgenus {Figaro) because of the presence of a crest on both
lower and upper sides of the caudal peduncle seems to us an unnecessarily minute sub-
division. Among the other Scyllorhinidae Pentanchus profundicolus Smith and Radcliffe,
1912,° and another unnamed species^" are set apart from the rest and from all other galeoid
sharks by the fact that they have only one dorsal fin." Among the species that remain after
subtraction of the foregoing, the first dorsal of one, Catulus cephalus Cllbert, 1 891, orlgl-

2. Scyliorhinus caniculus Linnaeus and S. stellaris Linnaeus, the so-called Spotted Dogfishes.

3. Mem. Harv. Mus. comp. ZooL, jd, 1913 : 68 [Catulidae].

4. Proc. Acad. nat. Sci. Philad., Ss, 1934: 233- 5- Aust. Zool., 9, 1939: 227-

6. Ann. S. Afr. Mus., 27 (i), 1925; 39. 7. Nature, Lend., /.;«, 1941 : 7-

8. These fish are commonly called File Tails in California.

9. The type specimen, now in the U.S. National Museum, shows no sign of mutilation. It is further interesting for the
fact that its gill openings are of the character pictured in Fig. 38, 39 for Afristurus frofundorum and A. riveri.

10. A Japanese scyliorhinid with only one dorsal fin is briefly described, but without specific name, by Jordan and
Hubbs (Mem. Carneg. Mus., 10, 1925: 100).

11. Jordan and Hubbs (Mem. Carneg. Mus., /o, 1925: 100) also suggest that the doubtful genus Caninoa of

Fishes of the Western North Atlantic 197

nates considerably in front of the origin of the pelvics, whereas in all the others it origi-
nates over or considerably behind the latter.^^ In this respect, and in others also, it is so
aberrant that we have recently proposed the new genus Cephalurus for it," based on study
of a specimen from the original series.

Among the remaining scyliorhinids eight clearly distinct species from various parts
of the world are set apart by the facts that labial furrows, well developed on the lower
jaw, do not extend around the corner of the mouth or onto the upper jaw, and that the
upper lip is expanded to close over the lower near the corner of the mouth. It was for a
member of this group (caniculus Linnaeus, 1758) that the earliest scyliorhinid genus was
proposed {Scyliorhinus Blalnville, 18 16). It is true that in this instance generic diagnosis,
based primarily on the morphology of the labial furrows, runs counter to the grouping that
might be based on the details of the nostril, and on the relationship of the latter to the
mouth, for among the species with well developed lower labial furrows and no upper
furrow are some in which the anterior margins of the nostrils reach to the mouth but
others in which they fall short of the latter, and a similar range of variation, based on
whether or not the nostril is connected to the mouth by a shallow groove, exists among
them. But the varietal series are so continuous in these respects that nothing would be
gained by abandoning the labial furrows in favor of the nostrils as the primary character.
Therefore, it seems logical to use the labial furrow rather than the nostrils as the generic
criterion, except for two South African species, Poroderma fantherinum Muller and
Henle, 1841," and P. marleyi Fowler, 1934,'^ in which the anterior margin of the nos-
tril is extended as a long tapering barbel. For these a separate genus seems appropriate.
Unfortunately, however, the old name Poroderma is not available for them, because its
type species" lacks the barbel, and is in fact a typical Scyliorhinus. But there is no need to
coin a new name, Fowler^^ having proposed Conoporoderma as a subgenus for the species
with barbels.

In some of the members of the family still to be considered the labial furrows extend
from the lower jaw around the angle of the mouth onto the upper jaw, while in others they
are wholly lacking. The latter category includes the peculiar Swell Sharks, which are able

Nardo (Atti Riun. Sci. Ital., 1 841 : 312) from the Mediterranean may be Pentanchus-\\ke, and Fowler (Bull. U.S.
nat. Mus., 100 [/j], 1941 : 26) retains it provisionally among the Scyliorhinidae. But it does not seem likely that
the combination of characters credited to it by Nardo (single dorsal fin, but with only 5 gill openings and no
spiracle) actually applies to any existing shark. For a history of the case, with suggestions as to what the speci-
men in question may actually have been, see Fowler, 1941; also Doderlein (Man. Ittiol. Medit., 2, 1881: 82).
I 2. Another species with first dorsal far forward, classed in this family by Garman {Proscyllium habereri Hilgen-
dorf, 1904), is placed among the Triakidae by us.

13. See Bigelow and Schroeder (Copeia, 2, 1941 : 73) for discussion and detailed description.

14. Usually credited to Andrew Smith, 1837, but Smith (Proc. zool. Soc. Lond., 1837: 85) listed it only by name;
the earliest account of it was by Muller and Henle (Plagiost., 1841 : 13).

15. Fowler (Proc. Acad. nat. Sci. Philad., ^5, 1934: 234) has pointed out that it was actually a specimen of this
species that he pictured earlier (Proc. Acad. nat. Sci. Philad., 77, 1925 : 188) under the name Scyliorhtnus recant.

16. Porodertna africanum A. Smith (Proc. zool. Soc. Lond., 1837: 85) equals Squalus africanus Gmelin, 1789.

17. Proc. Acad. nat. Sci. Philad., 6$, 1934: 234.

198 Memoir Sears Foundation for Marine Research

to inflate themselves with air, and which have widely distensible jaws provided with ver-
tical "accordion" folds in the corners, as well as very broad, flat heads. The majority of
recent writers have grouped these in the genus CefhaloscylUum. Fowler^^ has also pro-
posed the subgenus H olohalaelurus for two other species" that agree with the Swell
Sharks in lacking labial furrows, but which differ from them in having no ability to in-
flate and in having less distensible mouths, more slender trunks, shorter body cavities, as
well as in different relative sizes and locations of the fins. We propose to raise this sub-
genus to generic rank.

The remaining species in which there is a well marked furrow around the corner of
the mouth reaching out onto both jaws are subdivisible by the relation of nostril to mouth,
size of the second dorsal relative to anal fin, and length of the interspace between anal and
caudal.

In one rather sharply defined category of some thirteen named species, all from deep
water, the nostril is widely separated from the mouth and wholly distinct from the latter,
the anal is more than two and one-half times as long as the second dorsal, the interspace
between the anal and the caudal is very short or even reduced to a mere notch, there are no
folds below the eyes, and the snout is long and fleshy with very prominent mucous pores.
Fowler'" has recently distributed these species among three subgenera, based on the pres-
ence or absence of cirri on one or both margins of the nostril. But according to pub-
lished accounts and to our own examination of three members of the group, there is too
much intergradation in this respect for sharp separation. We therefore refer all of them
to the genus Apristurus Garman, 19 13.

In a second category the anterior margins of the nostrils similarly fall considerably
short of the mouth and there is a labial furrow around the corner of the latter; but they
differ from Ap-is turns by having a much longer interspace between caudal and anal, a con-
siderably smaller anal relative to the second dorsal, and a well marked fold below the eye.
Although the twelve named members of this group {Halaelurus Gill, 1861) resemble
one another so closely that some reduction in the number of species is to be expected even-
tually. Fowler'^ divides them among two subgenera, Aulohalaelurus and HalaeluruSy
according to the lengths of the labial furrows, while Whitley has raised the former to
generic rank, besides proposing two new genera, Juncrus and Asymbolus^' But the differ-
ences between the several species of this group are so slight that we refer all of them to
the old genus Halaelurus.

There remain only those species which fall with Halaelurus in most respects, except
for the anterior margin of the nostril, which more or less overlaps the anterior part of the
mouth, and except for a shallow groove which extends either from the nostril to the mouth

18. Proc. Acad. nat. Sci. Philad., 8s, 1934: 235 ; Bull. U.S. nat. Mus., 100 (/j), 1941 : 4'-

19. ScyUiorhinus functatus Gilchrist (Mar. biol. Rep. Cape Town, 2, 1914: 129) and S. regani Gilchrist (Mar.
biol. Rep. Cape Town, 2 [3], 1923 : 45, 46).

20. Bull. U.S. nat. Mus., 100 (75), 1941 : 53. 21. Bull. U.S. nat. Mus., 100 i's), 1941 : 4'-

22. Aust. Zool., 9, 1939: 229; Juncrus for Scyllium mncenti Zietz, 1908, and Asytnbolus for Scyllium anale Ogilby,
1885, both from Australia.

Fishes of the Western North Atlantic 199

or part way to the latter. Although this group includes only three known species," our own
examination of specimens in the collection of the Museum of Comparative Zoology satis-
fies us that Carman's" reference of them to his two new genera, Haploblepharus and Ate-
lomycterus, was justified by the sharp differences summarized in the following key.

Key to Genera
la. Only one dorsal fin. Pentanchus Smith and Radcliffe, 1912.

Philippines, Japan.

lb. Two dorsal fins.

2a. Origin of ist dorsal considerably anterior to origin of pelvicsj rear contours of
dorsal fins straight or concave. Cephalurus Bigelow and Schroeder, 1 94 1 .

Gulf of California and Revillagigedo Islands, oflF
west coast of Mexico.
2b. Origin of ist dorsal over, or usually behind, origin of pelvics.

3a. Denticles along dorsal margin of anterior part of caudal enlarged and modi-
fied in shape, forming a distinct crest, outlined below by a narrow band of
naked skin.

4a. Nostrils far from mouth, the distance from their inner angles to corners
of latter about V2 as great as horizontal diameter of eyej posterior mar-
gin of nostril not lobed; snout long, thin, its mucous pores not conspicu-
ous. Galeus Rafinesque, 18 10, p. 214.
4b. Nostrils close to mouth, although entirely separate from latter j distance
from inner angle of nostril to corner of mouth not more than V4 as great
as horizontal diameter of eyej posterior margin of nostril with a well
developed lobe j snout short and thick, its mucous pores very prominent.

Parmaturus Garman, 1906.

California, Japan.

3b. Denticles along dorsal margin of anterior part of caudal similar to those
lower down, not forming a distinct crest.

5a. Anterior margin of nostril bilobed, the outer lobe in the form of a fleshy
barbel reaching to mouth (Fig. 31). Conoporoderma Fowler, 1934.

S. Africa, Natal, Mauritius.

5b. Anterior margin of nostril little or not at all bilobed j without well de-
veloped barbel.
6a. A well developed labial furrow on lower jaw, but not around corner

of mouth or on upper jaw. Scyliorhinus Blainville, 1 8 1 6, p. 202.
6b. Labial furrows either absent, or extending around corner of mouth

if present.

23. Scyllium edtuardsii Voigt (in Cuvier, Tierreich, s, 1832: 504), S. Afr.; S. marmoratum Bennett (Mem. Raffles,
1830: 693), Malaysia, India; and A telomycterus macleayi Whitley (Aust. Zool., 9, 1939: 230), Australia.

24. Mem. Harv. Mus. comp. Zool., $6, 1913: 100, 101.

200 Memoir Sears Foundation for Marine Research

Figure 31. i4, Ha-ploblefkarus edwardsii (Harv. Mus.
Comp. ZooL, No. 1028). Showing nasal flap and rela-
tionship of nostril to mouth, about l]^ x natural size. B,
Conoforoderma ■pantherinum{V{.zr\. Mus. Comp. Zool.,
No. 497). Right-hand nostril and part of upper jaw
showing the nasal barbel, about 3 x natural size.

Fishes of the Western North Atlantic 201

7a. No labial furrow on either jaw, or around corner of mouth.
8a. Mouth broadly distensible, with vertical folds at corners;
stomach inflatable with air; anal only about as long as and
dorsal, its origin under origin of latter; body sector of
trunk to cloaca considerably longer than tail sector.

Cephaloscyllium Gill, 1862.
Eastern Pacific from middle Cali-
fornia to Chile; Japan, Australia,
Tasmania and New Zealand re-
gion; South Africa.

8b. Mouth not distensible, without vertical folds at the cor-
ners; stomach not inflatable with air; anal more than 1V2
times as long as 2nd dorsal, its origin anterior to origin of
latter by a distance equal to at least V2 the length of its
base; body sector of trunk to cloaca considerably shorter
than tail sector. Holohalaelurus Fowler, 1934.

South Africa, Natal.

7b. A labial furrow around corner of mouth and extending for-
ward for a longer or shorter distance on each jaw.
9a. Anterior margin of nostril expanded as a flap, overlapping
front edge of mouth; no definitely outlined lower nasal
flap; a shallow groove extending at least part way from
nostril toward mouth (Fig. 31).

lOa. A groove extending from nostril to mouth; anterior
flaps of the 2 nostrils not separated by a definite
gap opposite symphysis of upper jaw, their outlines
nearly straight; anal larger than 2nd dorsal, its base
wholly anterior to base of latter; origin of ist dorsal
behind rear end of base of pelvics; fold below eye
hardly defined, if at all.

HaploHepharus Garman, 19 13.
South Africa.
lOb. Nasal grooves not extending to mouth; anterior nasal
flaps widely separated, their outlines, as well as out-
lines of intervening isthmus, forming 3 rounded
lobes; anal at least no larger than 2nd dorsal, the
rear end of its base under midpoint of latter; origin
of 1st dorsal in front of rear end of base of pelvics;
a strongly developed fold below eye.

A telomycterus Ga.rma.n, 19 13.
China, Indo-China, Siam, Malaysia,
Philippines, India.

202 Memoir Sears Foundation for Marine Research

9b. Anterior nasal flaps fall considerably short of mouth; a
posterior nasal flap is also present in most cases.
1 1 a. Interspace between anal and caudal at least as long
as base of anal; base of anal not more than twice as
long as base of 2nd dorsal; folds below eyes strongly
developed; mucous pores on snout not conspicuous.
Halaelurus Gill, 1862.
South Africa; tropical In-
dian Ocean and Arabian
Gulf; India; Australasia;
Philippines, China, Formosa,
Japan; Chile and Patagonia;
Argentina.
lib. Interspace between anal and caudal less than V5 as
long as base of anal ; base of anal more than twice as
long as base of 2nd dorsal; no fold below eye; mu-
cous pore system on lower surface of snout very con-
spicuous. Afristurus Garman, 1 9 1 3, p. 2 1 9.

Genus Scyliorhinus Blainville, 1 8 1 6

Scyliorhinus Blainville, Bull. Sec. philom. Paris, 18 16: 121; type species, S. caniculus Blainville,^" equals
Sgualus ctmicu/us hinnAeus, 1758.

Generic Synonyms:""

Calulus Valmont, Diet. Hist. Nat. Paris, 4, 1768:^' 51 ; type species, C. major vulgaris Valmont, equals Squalus

caniculus Linnaeus, 1758; Andrew Smith, Proc. zool. Soc. Lond., 1838: 85 (in part).
Galeus (in part) Rafinesque, Indice Ittiol. Sicil., 1810:46; for G. caniculus Rafinesque, equals Squalus caniculus

Linnaeus, 1758.
Scyllium Cuvier, Regne Anim., 2, 1817: 124; type species, Squalus caniculus Linnaeus, 1758, designated by

Jordan, Genera Fish., /, 1 91 7: 97.
Scylliorhinus Blainville, in VieiUot, Faune Franc, Poiss., 1825: 68; substitute for Scyliorhinus Blainville,

1816.
Poroderma A. Smith, Proc. zool. Soc. Lend., 1837: 85 ; type species, P. africanum Smith, equals Squalus ofri-

canus Gmelin, 1 789. South Africa.
Halaelurus Tzmki, Fish. Japan, i, 1911: 13, pi. 3, fig. 12; for H. r«<2f«Tanaka; not Halaelurus Gill, 1862.

Generic Characters. Two dorsal fins; origin of ist dorsal over or slightly anterior
to rear ends of bases of pel vies; denticles along dorsal margin of caudal similar to those
lower down, not forming a distinct crest; nasal barbels rudimentary or wholly lacking;

25. While Blainville gave no authorship for this or for any of the several other included species, his subsequent
diagnosis (in Vieillot, Faune Franc, 1825: 71) of canicula showed that it referred to Squalus caniculus Lin-
naeus, 1758, which was later designated as type of the genus by Gill (Ann. N.Y. Lye, 7, 1862: 407).

26. For list of fossil genera perhaps synonymous with Scyliorhinus, see Fowler (Bull, U.S. nat. Mus., joo [/j],
1941: 34).

27. Preoccupied (Kniphof, 1759) for insects and not available even otherwise for sharks; Valmont's names, when
binomial, were so only accidentally (see ruling by International Commission on Zoological Nomenclature,
Smithson. misc. Coll., 73 [3], 1925: 27) ; the name, as a shark, must therefore date from Andrew Smith, 1837.

Fishes of the Western North Atlantic 203

anterior nasal flaps may or may not reach mouth; nostril either entirely separate from
mouth or connected with latter by a very shallow groove only; a well developed labial
furrow on lower jaw, but none on upper; upper lip expanded to close over lower at corners
of mouth; eye with or without a longitudinal fold below lower eyelid; spiracle small,
close to corner of eye; anal considerably larger than 2nd dorsal, separated from caudal by
a considerable interspace; inner margins of pelvics united posterior to cloaca for a short
distance in females and for a longer distance in males; teeth with one large central, and
several small lateral, cusps, several series functional ; dermal denticles lanceolate, strongly
ridged. Egg cases horny, oblong, with long filamentous tendrils at the corners which wind
around sea weeds, etc. The eggs are said to be expelled two at a time, and the young to
hatch about six months after the eggs are laid.

Range. Both sides of North Atlantic; Mediterranean; South Africa; Natal; Japan;
Korea.

Fossil Teeth. Upper Cretaceous to Pliocene, Europe; Upper Cretaceous, western
Asia, North America; Eocene, North Africa.

Key to Species

I a. Anterior nasal flaps reach rearward nearly or quite to mouth.
2a. Anterior nasal flaps joined in the midline, or nearly so.

caniculus Linnaeus, 1758.

Eastern North Atlantic, Medi-
terranean.^*

2b. Anterior nasal flaps separated one from the other in the midline by a consider-
able interspace. j/(?//«m Linnaeus, 1758.

Eastern North Atlantic, Medi-
terranean.

lb. Anterior nasal flaps separated from mouth by a considerable space.

3a. Origin of ist dorsal as close to origin of anal as to rear ends of bases of pelvics;
2nd dorsal as large as ist. capensis Miiller and Henle, 1 841 .

South Africa, Natal, and perhaps India.^'

3b. Origin of ist dorsal over rear end of bases of pelvics, or at least much closer to
them than to origin of anal; 2nd dorsal smaller than ist.
4a. Origin of ist dorsal closer to tip of caudal than to tip of snout by a distance

equal to length of latter in front of mouth ; color pattern a dark network on

paler ground. retifer Garman, 1 8 8 1 , p. 207.

4b. Origin of ist dorsal as close to tip of snout as to tip of caudal, or a little

closer; color pattern spotted or blotched.

5a. Base of anal considerably longer than base of ist dorsal.

28. Including' duhamelii Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 73), the type specimens of which
(Harv. Mus. Comp. Zool., No. 60, 63) appear to represent a dwarf race of caniculus, or perhaps only a color
variety.

29. Day, Fish. India, 1878: 724, pi. 190, fig. 1.

204 Memoir Sears Foundation for Marine Research

6a. Caudal only about as long as from tip of snout to 5th gill opening}
color pattern white-spotted on dark ground tint.

/orm Howell-Rivero, 1936, p. 211.
6b. Caudal about as long as from tip of snout to axil of pectoral} color
pattern dusky or black-spotted on pale ground tint.

boa Goode and Bean, 1895, p. 204.
5b. Base of anal only as long as base of ist dorsal, or shorter.

torazame Ta.mka., 1908.
Japan.

Scyliorhinus boa Goode and Bean, 1895.

Figure 32

Study Material. Type specimen, a newly-hatched male, 1 5 1 mm. long, in poor con-
dition, from Barbados, in 200 fathoms (Harv. Mus. Comp. ZooL, No. 1335); newly-
hatched male, 87 mm. long, from north coast of Cuba, in 235 fathoms (Harv. Mus. Comp.
ZooL, No. 36156)} half-grown male, 316 mm. long, taken 25 to 30 miles ESE. from

Figure 32. Scyliorhinus boa, immature male, 316 mm. long, from near Rio de Janeiro, Brazil (Mus. Nac.
Rio de Janeiro). A Anterior part of head from below to show nostril and labial furrows, about I.4 x natural
size. B Side view of anterior part of head, almost 1.5 x natural size. C Dermal denticles, about 14 x. Z) Upper
teeth from side of jaw, enlarged.

Fishes of the Western North Atlantic 205

Ilha Rasa near Rio de Janeiro, Brazil, in 80 meters, the type of S. haeckelii (Ribeiro),
1907 (Mus. Nac. Rio de Janeiro, No. 494).

Distinctive Characters. Separable from 5. retifer by its obtusely rounded snout and
color pattern, and from S. torrei, which it closely resembles, by its relatively longer caudal
fin and by its color (see Key, p. 204).

Description. Proportional dimensions in per cent of total length. Male, 151 mm.,
from Barbados (Harv. Mus. Comp. Zool., type. No. 1335). Male, 316 mm., from Brazil
(Mus. Nac. Rio de J., type of S. haeckelii, No. 494).

Trunk at origin of -pectoral: breadth 10.6, i i.i ; height 7.3, 8.9.

Snout length in front of: outer nostrils 3.3, j mouth 4.0, 5.1.

Eye: horizontal diameter 3.3, 3.5.

Mouth: breadth 6.3, 6.3; height 2.6, 3.8.

Nostrils: distance between inner ends 2.5, 2.2.

Labial furrow length: lower 1.7, 1.3.

Gill opening lengths: ist 2.0, 1.6} 2nd 1.5, j 3rd 1.5, ; 4th 1.5, }

5th I.I, 1.2.

First dorsal fin: vertical height 4.6, 5.4; length of base 6.0, 6.3.

Second dorsal fin: vertical height 3.3, 3.2 ; length of base 4.3, 5. i.

Anal fin: vertical height 3.3, 3.2; length of base 9.4, 9.2.

Caudal fin: upper margin 26.8, 23.4; lower anterior margin 9.0, 8.5.

Pectoral fin: outer margin 10.2, 14.2 j inner margin 6.3, 6.3} distal margin 8.0, 9.8.

Distance from snout to: ist dorsal 43. 8, 48.4; 2nd dorsal 59.7, 67.15 upper caudal

73.2, 76.6} pectoral 17.2, 18.75 pelvics 37.9, 39.25 anal 53.0, 60.1.

Interspace between: ist and 2nd dorsals lO.O, 12.0 5 2nd dorsal and caudal 8.0,

6.O5 anal and caudal 10.6, 9.2.

Distance from origin to origin of: pectoral and pelvics 23.6, 20.9 5 pelvics and anal

15-3, I9-7-

Trunk slender, much compressed laterally rearward from pelvics. Dermal denticles
rather loosely spaced, much longer than broad, with 3-5 ridges and tridentate margins,
the median tooth considerably the largest, their blades erected at an angle of about 40°
over trunk as a whole, giving a very rough effect.

Head convex in dorsal profile but flattened below. Snout broadly rounded, its length
in front of mouth between Vs and ^4 of length of head. Eye narrow, oval, its horizontal
diameter nearly as long as snout in front of mouth, the fold below eye well marked when
eye is open but hardly distinguishable when it is closed. Spiracle round, very small, pos-
terior to rear corner of eye by a distance about 0.2 times as great as horizontal diameter of
latter. Gill openings moderately concave in outline anteriorly, the ist slightly the longest,
about V2 as long as horizontal diameter of eye, the 5th slightly the shortest, the interspaces
between them decreasing in breadth rearward, the interspace between 3rd and 4th over
origin of pectoral. Nostrils slightly oblique, entirely distinct from mouth and widely sepa-

2o6 Memoir Sears Foundation for Marine Research

rated from each other, the distance between them nearly Yo as great as length of snout in
front of mouth, the anterior margin expanded as a rather narrow subtriangular lobe with
rounded apex and well marked median crest but falling considerably short of the mouth,
the posterior margin also developed as a rounded flap (Fig. 32 A), much as in S. retifer
and S. torrei. Mouth obtusely ovate, about ^^ as high as broad. Lower labial furrow
slightly less than V3 as long as distance from corner of mouth to symphysis of lower jaw.

Teeth f^Efjj"* similar in the 2 jaws, usually with 5, occasionally with 3 (or even 7)
cusps, the median much the longest, narrow-triangular and sharp-pointed, curving slightly
toward corner of mouth in most cases, the anterior surfaces of teeth longitudinally striate;
5 rows (locally only 4) functional in each jaw.

First dorsal brush-shaped, its margins nearly straight, its corners narrowly rounded,
its origin slightly behind rear end of bases of pelvics, and a little nearer to snout than to
tip of caudal, the interspace between ist and 2nd dorsals about twice as long as base of ist
dorsal. Second dorsal similar in shape to ist, and nearly as long at base as latter, but only
about % as high vertically, its origin on a vertical line about halfway between midpoint
of base of anal and rear end of latter. Caudal a little less than Yi of total length, relatively
somewhat longer in small specimens than in large,'^ its upper contour nearly straight, its
terminal sector transversely truncate, with rounded corners, occupying about Vs total
length of the fin, lower anterior corner much more than a right angle. Anal a little
less than twice as long at base as 2nd dorsal, and longer than ist dorsal by a distance
about as long as horizontal diameter of eye, with nearly straight margins, rounded apex
and subacute free rear corner, about Vs as long as the base. Pelvics about as large as anal,
their anterior margins nearly straight, distal margins weakly concave, corners subangular,
their inner edges, in half -grown male, united behind cloaca for a little less than ^o their
lengths. Pectoral about 3 times as large as ist dorsal in area, about 70% as broad as long,
the outer and inner margins moderately convex, the distal margin nearly straight, apex
narrowly rounded, inner corner more broadly so.

Color. Back and sides pale yellowish brown, marked transversely with seven broad
but indistinct dark blotches, one midway of the caudal, one at caudal's origin, one oppo-
site each dorsal fin, and three equally spaced in front of the first dorsal, the most anterior
being opposite the origin of the pectorals; also a large number of small dark chocolate-
brown spots of varying sizes irregularly spaced, some nearly circular and some in the form
of rosettes; one much larger than the others below the first dorsal, with others opposite
the origin and rear part of anal; likewise a lunate blotch on each flank about midway be-
tween the rear corner of the pectoral and the origin of the pelvics ; lower surface very pale
yellowish brown, plain except that the lower side of head is faintly mottled; and there are a
few dark spots on the pectorals (about twice as many on the one as on the other in the half-

30. It is possible that there was one more series of teeth in each jaw, it being difficult to determine the precise number
in the available material.

31. A little longer than from snout to inner corner of pectoral in newly hatched specimen, but only about as long
as from snout to axil in a half-grown one.

Fishes of the Western North Atlantic 207

grown specimen). That the distribution of the dark spots is not the same on the two sides
of one specimen, and that there are many more on it and on a newly hatched specimen from
Cuba than on another from Barbados, show that their number is not a specific character.

Size. The state of sexual development of the larger specimen, as indicated by the
length of its claspers, suggests that this species becomes mature at a length of perhaps two
feet. S. boa is thus a considerably larger shark than S. torre't (p. 213). No females have
yet been seen.

Developmental stages. The egg cases have not been identified.

Habits. The depths of capture, listed above, make it likely that this is an inhabitant
of moderately deep waters and probably a bottom-dweller. Other than this nothing is
known of its habits.

Range. S. boa is positively known only from Brazil, from Cuban waters and from
the Barbados (see Study Material, p. 204).

Synonyms and References:

Scylliorhinus boa Goode and Bean, Smithson. Contr. KnowL, 30, 1895: 17; Mem. Harv. Mus. comp. Zool.,

22, 1896; Spec. Bull. U.S. nat. Mus., 2 (off Barbados) ; Howell-Rivero, Proc. Boston See. nat. Hist., 41,

1936: 44 (Cuba).
Scylliorhinus retifer Goode and Bean, Smithson. Contr. KnowL, 30, 1895: pi. 2, fig. 6; Mem. Harv. Mus.

comp. Zool., 22, 1896; Spec. Bull. U.S. nat. Mus., 2 (same); not Scyllium retiferum Garman, 1881.
Catulus retifer var. boa Ribeiro, Bol. Soc. nac. Agric. Brasil, 1 904: 17 (Brazil); not Scyllium retiferum

Garman, 1 88 1.
Catulus haeckelii Ribeiro, Mem. Mus. nac. Rio de J., 14, 1907: 163, pi. 8 (Ilha Rasa, near Rio de Janeiro,

Brazil, descr.) ; Fauna brasil. Peixes, 2 (i), Fasc. i, 1923: 21, pi. 7 (same as Ribeiro, 1 907); Fowler,

Arqu. Zool. Estado Sao Paulo, 3, 1942: 127 (Brazil).
Scyliorhinus retifer (in part) Regan, Ann. Mag. nat. Hist., (8) I, 1908: 457 (class.).
Catulus boa Garman, Mem. Harv. Mus. comp. Zool., 36, 191 3: 77 (descr.).

Scyliorhinus retifer (Garman), 1881

Chain Dogfish

Figure 33

Study Material. Type specimen, male, 307 mm. long, from off Virginia (Harv, Mus.
Comp. Zool., No. 825) ; a male, 428 mm, long, from off New Jersey (Harv, Mus. Comp,
Zool,, No, 33932); also two females, 300 and 370 mm., from offing of southern New
England, in 50-70 fathoms (Harv. Mus. Comp. Zool.),

Distinctive Characters. This species is most obviously separated from other local
species of the genus by its chain-like color pattern, by its wedge-shaped snout, and by the
fact that the origin of its first dorsal is closer to the tip of the caudal than to the snout.

Description. Proportional dimensions in per cent of total length. Male, 307 mm.,
from Lat, 38° N., Long. 73° W, (Harv, Mus, Comp, Zool,, type, No, 825). Male, 428
mm., from 1 10 miles SE. of Atlantic City, N. J, (Harv. Mus, Comp, Zool,, No, 33932),

2o8 Memoir Sears Foundation for Marine Research

Trunk at origin of ■pectoral: breadth 10.6, 11.7; height 9.2, 9.6.

Snout length in front of: outer nostrils 3.6, 3.3 ; mouth 5.2, 5.0.

Eye: horizontal diameter 3.6, 3.7.

Mouth: breadth 6.8, 8.2} height 3.6, 3.3.

Nostrils: distance between inner ends 2.1, 1.9.

Labial furrow length: lower 1.6, 1.6.

Gill opening lengths: ist 1.5, 2.2j 2nd 1.4, 1.95 3rd 1.4, 1.9; 4th 1.4, 1.9J 5th

1.1,1.3.

First dorsal fin: vertical height 6.0, 7.O5 length of base 6.$, 6.5.

Second dorsal fin: vertical height 3.7, 4.75 length of base 6.0, 5.1.

Anal fin: vertical height 4.1, 4.25 length of base 8.8, 8.2.

Caudal fin: upper margin 21.9, 19.3} lower anterior margin 12.3, 11.4.

Pectoral fin: outer margin 14.3, 13.3; inner margin 7.5, 7.0; distal margin 8.5,

II. 8.

Distance from snout to: ist dorsal 50.O, 53.O; 2nd dorsal 67.2, 69.8 j upper caudal

78.1, 80.7} pectoral 19.2, 21.75 pelvics 42.1, 44.35 anal 61.2, 62.0.

Interspace between: ist and 2nd dorsals lO.i, 11.8} 2nd dorsal and caudal 6.0,

7.0 } anal and caudal 9.1, 8.4.

Figure 33. Scyliorhinus retifer, immature male, about 428 mm. long, from off New Jersey (Harv. Mus.
Comp. Zool., No. 33932). A Anterior part of head from below. B Snout showing nosuils, natural size. C Pelvic
fins with claspers. D Dermal denticles, about ij x. E Apical view of dermal denticle, about 35 x. f Upper and
lower teeth from center of mouth, about 4 x. G Upper and lower teeth from sides of jaws near corners of
mouth, about 5 x.

Fishes of the Western North Atlantic 209

Distance from origin to origin of: pectoral and pelvics 24.8, 22.55 pelvics and anal
19.4, 19.8.

Trunk slender, its breadth at origin of pectorals only about Vs, its height Vio? of
total length, tapering rearward. Body sector to cloaca about as long as tail sector. Caudal
peduncle nearly as broad as deep, oval in cross-section. Dermal denticles narrow, lanceo-
late, with scute tips, their blades only slightly raised, 3-5 ridged, the axial ridge much the
strongest, their posterior margins entire on some denticles but notched between the ridges
on others.

Head flattened above. Snout wedge-shaped, but with blunt tip, its length in front
of mouth about V4 of length of head. Eye moderately narrow, oval, with horizontal diam-
eter twice or more the vertical, Its horizontal diameter about % as long as snout in front
of mouth, its anterior edge a little posterior to front of mouth, the longitudinal fold below
eye but weakly indicated. Spiracle an oblique slit, about Y'j as long as horizontal diameter
of eye, situated close behind, and a little below, latter. First gill opening the longest, about
V2 to % as long as horizontal diameter of eye, the 5th shortest, only about V2 as long as ist,
the 4th and 5th over anterior part of pectoral. Nostrils entirely distinct from mouth, nearly
transverse, their anterior margins expanded as subtriangular flaps, with strong transverse
median crests (no barbel), separated from mouth at nearest point by a distance about Vi
as great as horizontal diameter of eye, and separated one from the other in the midline
by a distance about V2 as great as from the median angle of the nostril to the mouth. Mouth
ovate, about V2 as long as wide; a strongly marked labial furrow at corner of lower jaw
extending inward about Vs the distance to the symphysis; no furrow on upper jaw but
upper lip somewhat expanded at corner of mouth, thus closing over the lower.

Teeth about |^=j=|i; alike in the 2 jaws, the triangular median cusp flanked near its
base on either side by i (rarely 2) smaller cusps, the median cusp larger, relative to the
laterals, in larger than in smaller specimens; lower jaw, but not upper, with a small
median tooth ; usually 3 or 4 rows functional.

First dorsal brush-shaped, its origin closer to tip of caudal than to tip of snout by a
distance about equal to length of latter in front of mouth, posterior to rear end of bases of
pelvics by a distance about % as long as horizontal diameter of eye, the rear end of its
base a little posterior to tips of pelvics; its anterior margin nearly straight, its posterior
margin slightly convex, its apex rounded, its free lower margin about as long as its base.
Second dorsal about V2 as large in area as i st, its rear margin weakly concave, its free rear
corner somewhat more slender than ist, its origin over rear part of base of anal. Caudal
only about V5 of total length, with well marked subterminal notch, its terminal sector ^3
to V2 the total length of the fin, brush-shaped, its tip either squarely truncate (Fig. 33)
or indented in the midline (type specimen), its lower anterior corner subangular and much
more obtuse than a right angle. Anal subtriangular, with nearly straight edges, broadly
rounded apex, and moderately acute rear corner, its base about Vs to V^ longer than that
of 2nd dorsal, its origin about midway between perpendiculars at rear end of base of ist

2IO Memoir Sears Foundation for Marine Research

dorsal and at origin of 2nd dorsal. Pelvics a little larger in area than ist dorsal, sub-
triangular, with rounded apices and moderately acute rear corners, the inner margins
united behind cloaca for a little more than V2 their lengths in immature male. Pectoral
about twice as large in area as ist dorsal and % to % as broad as long, with rounded cor-
ners, slightly convex outer margin and straight distal margin.

Color. The ground tint is dark reddish brown above, yellowish below, with a very
characteristic pattern of narrow, sooty black stripes in groups of two crossing the back
just behind the pectorals, at the first dorsal, between the first and second dorsals, at the
second dorsal, at the anterior end of the caudal, and midway out on the latter; these
branching over the sides and out onto the pectorals in a loose net of polygonal meshes
which are irregular in size and shape.

Size. The largest specimen so far measured was 17 inches (430 mm.) long, the maxi-
mum length probably not being more than 2 to 2 ^/'2 feet.

Developmental Stages. Horny egg cases, presumably of this species (the only ovipa-
rous shark common off the middle Atlantic United States), are 50 to 57 mm. long by
1 8 to 23 mm. broad, with a long tendril at each corner and brownish amber in color.

Habits. This little shark lives on or close to bottom on the outer part of the Con-
tinental Shelf, chiefly at least between about the 40 and 125 fathom contours, all definite
records of it having so far been from within this depth range. There is no reason to suppose
that it ever strays shoreward into shoal water. Eggs, one with an embryo nearly ready for
hatching and others less advanced, have been taken in February off Chesapeake Bay, evi-
dence that the young are produced in late winter or early spring. Nothing more is known
of its life history, and nothing of its diet.

Range. All recorded captures of S. retifer have been from between the ofiings of
Cape Lookout, North Carolina, and northern New Jersey. Fishermen also report small
sharks, probably this species, on the Tilefish grounds at the outer edge of the Continental
Shelf off New York. Within this short sector, however, it appears to be very generally
distributed in the appropriate depth zone. Its chief center of abundance appears to lie off
Virginia (type locality, Lat. 38° 23' N., Long. 73° 34' W.), especially in the general
offing of Chesapeake Bay, where considerable numbers are taken by the winter trawl fishery
from January to March, specimens being brought in daily at times. S. retifer has also been
reported by name from the Tortugas, Florida," and from some unspecified locality be-
tween southern Florida, the Bahamas and Honduras.'* Re-examination in the first case
shows that the shark in question was Galeus arae (p. 21 1) j and since the second of these
records is by name only, the same may be true of it also.

Synonyms and References:

ScylliujTi retiferum Garman, Bull. Mus. comp. Zool. Harv., 8, 1881: 233 (descr., Lat. 38° 23' N., Long.

73°34'W.).
Scyllior/iinus relifer Jordin and Gilbert, BuU. U.S. nat. Mus., 16, 1883: 869 (ref.) ; Bean, Rep. U.S. Comm.

Fish. (1882), 1884:343 (oflf Woods Hole) ; Jordan, Rep. U.S. Comm. Fish. (1885), 1887:733 (off SE.

33. Longley and Hildebrand, Pap. Tortugas Lab., 34, 1941 : i.

34. Breder, Bull. Bingham Oceanog. Coll., 1 (i), 1927: 5.

Fishes of the Western North Atlantic 2ii

U.S.) ; Goode and Bean, Smithson. Contr. Knowl., jo, 1895: 16, 508, pi. 4, fig. 14, 15; Mem. Harv.

Mus. comp. Zool., 22, 1896; Spec. Bull. U.S. nat. Mus., 2 (descr., off SE. U.S.) ; Jordan, Evei'mann and

Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 1 1 (distrib.).
Scyliorhinusretifer Regan, Ann. Mag. nat. Hist., (8) /, 1908: 457 (class.) ; Fowler, Copeia, 30, 1916: 36 (off

mid. Atlant. U.S.) ; Nichols, Copeia, 1931 : 38 (egg cases) ; Schroeder, Copeia, 1931: 42 (off N. Jersey) ;

Firth, Copeia, 1934: 45 (egg cases, season, off Chesapeake Bay).
Catulus relifer Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 214 (Gulf Stream); Bull. U.S.

nat. Mus., 47 (l), 1896: 25 (descr.. Gulf Stream in deep water off S. Atlant. coast) ; Smith, Bull. N. C.

geol. econ. Surv., 2, 1907: 31 (off N. Carolina); Gudger, Proc. biol. Soc. Wash., 25, 1912: 154 (egg

cases, perhaps this species, N. Carolina) ; Garman, Mem. Harv. Mus. comp. Zool., 56, 1 91 3: 76 (descr.);

Radcliffe, Bull. U.S. Bur. Fish., 34, 1916: 249 (teeth, denticles, off N. Carolina); Breder, Field Bk.

Mar. Fishes Atlant. Coast, 1929: 1 1 (general) ; Pearson, U.S. Bur. Fish. Invest. Rep., (10) i, 1932: 17

(winter trawl fishery, off N. Carolina).
(?) Catulus retifer Breder, Bull. Bingham oceanogr. Coll., i (l), 1927: 5 (no loc, see p. 210).
Not Scylliorhinus retifer Longley and Hildebrand, Pap. Tortugas Lab., 34, 1 941 : I (this is Galeus arae).

Scyliorhinus torrei Howell-Rivero, 1936

Figures 34, 35

Study Material. Type specimen, female, 250 mm. long, off Havana, Cuba (Harv.
Mus. Comp. Zool., No. 1457)} ^^so 14 others, male and female, 130 to 292 mm. long,
collected off the north coast of Cuba by the research ship "Atlantis" in March 1938 and
April 1939 at depths of 210 to 250 fathoms (Harv. Mus. Comp. Zool.).

Distinctive Characters. S. torrei is easily separable from S. retifer by its very broadly
rounded snout and by its color pattern} from S. boa, which it closely resembles, by the
fact that the caudal is about as long as the distance from the tip of the snout to the origin of
pectoral, and by its coloration.

Description. Proportional dimensions in per cent of total length. Female, 250 mm.,
from Cuba (Harv. Mus. Comp. Zool., type. No. 1457). Male, 292 mm., from Cuba
(Harv. Mus. Comp. Zool., No. 36093).

Trunk at origin of -pectoral: breadth 10.8, 10.6; height 9.9, 8.2.

Snout length in front of: outer nostrils 3.2, 2.4; mouth 4.4, 3.9.

Eye: horizontal diameter 3.3, 3.1.

Mouth: breadth 7.2, 6.$; height 3.2, 3.4.

Nostrils: distance between inner ends 2.4, 2.1.

Labial furrow length: \ovitr 1.6, 1.5.

Gill opening lengths: ist 1.8, 2.1; 2nd 1.6, 1.4; 3rd 1.6, 1.45 4th 1.4, 1.2; 5th

i.O, i.O.

First dorsal fin: vertical height 5.6, 5.5; length of base 6.8, 6.8.

Second dorsal fin: vertical height 2.6, 2.6; length of base 5.4, 4.3.

Anal fin: vertical height 3.4, 3.1 j length of base 9.2, 8.6.

Caudal fin: upper margin 20.O, 21.O; lower anterior margin 9.6, 8.2.

Pectoral fin: outer margin 12. r, 10.33 inner margin 6.8, 6.$; distal margin 8.8,

7-9.

212

Memoir Sears Foundation for Marine Research

Distance from snout to: ist dorsal 50.5, 49.45 2nd dorsal 68.4, 68.3 ; upper caudal

80.0, 79.O; pectoral 18.8, 18.5; pelvics 41.2, 39.3; anal 60.2, 60.0.

Interspace between: ist and 2nd dorsal 1 1.6, 12.3; 2nd dorsal and caudal 7.4, 7.7;

anal and caudal 9.6, 10.8.

Distance from origin to origin of: pectoral and pelvics 23.6, 2 1 .7 ; pelvics and anal

19.2, 20.9.

S. torrei resembles 5. boa very closely in body form, shape of snout, nostrils and nasal
flaps, shape, size and relative position of fins, and in the teeth and dermal denticles. The
significant points of difference are as follows: in torrei the mouth is slightly the lower-
arched, its height being only about 40 per cent of its breadth as against 50 per cent in boa;
in torrei the length of the snout in front of the mouth is slightly less, it being only a little
more than half as great as the breadth of the mouth as against about four-fifths in boa; the
pectorals of torrei are only a little larger in area than the first dorsal, whereas in boa they
are twice as large as the latter j and while the denticles rise steeply from the skin in newly
hatched specimens of both torrei and boa, in larger specimens of the former they lie nearly
flat and the surface texture of the skin is smoother compared with the pronounced rough-
ness of boa. Also, in the male torrei the inner edges of the pelvics are connected to one
another and to the ventral surface of the trunk more nearly to their tips than in either boa
or in retifer. However, the most striking difference between the species is in the color

Figure 34. Scyliorhinus torrei, female, 276 mm. long, from Havana, Cuba (Harv. Mus. Comp. 7,oo\.,
No. 34776). A Dermal denticles, about 30 x. B Apical view of dermal denticle, about 45 x. C Upper and
lower teeth from near center of mouth (Nos. 1—4). D Twelfth and thirteenth upper teeth. E Upper nine-
teenth tooth. F Lower tenth and eleventh teeth. G Lower eighteenth tooth. C-G, about 9 x.

Fishes of the Western North Atlantic

213

pattern. Although the pale brown back and upper sides of torrei are transversely marked
by a series of indistinct darker blotches, as in boa, two of these being on the caudal and one
opposite the origin of the pectoral, the finer markings of torrei consist of small oval whitish

Figure 35. Scyliorhinus torrei. A Anterior part of
head of specimen illustrated in Fig. 34, from below,
about 1.7 X. B Pelvic fins and claspers of male, about
292 mm. long(Harv. Mus. Comp. ZooL, No. 36093),
about 1.7 X.

spots (in contrast to the dark markings of boa) which are rather evenly distributed over
the whole back and upper sides. The lower surface is of a very pale shade of the same tint
as the upper sides, or nearly white, without evident markings either on the trunk or on
the fins.

Size. The male of torrei has claspers extending far beyond the tips of the pelvics
(suggesting maturity or approaching maturity) at a total length of only about 247 mm.,
showing that this is a much smaller species than boa, perhaps not growing much larger
than a maximum of 300 mm. or so.

Develof mental Stages. Neither the eggs nor the embryos of torrei have yet been
seen.

Habits. Nothing is known of its habits.

214 Memoir Sears Foundation for Marine Research

Range. S. torrei is so far known only off the northern coast of Cuba, but evidently
it is common there.

Synonyms and References:

Catulus boae Sanchez-Roig, Revist. Agric. Feces Cubana Commerc. Trabaj., 1931: 17 (Cuba, not seen); not
Scylliorhinus boa Goode and Bean, 1 895.

Scylliorhinus torrei Howell-Rivero, Proc. Boston Soc. nat. Hist., 41, 1936: 43, pi. 9 (descr., deep water off
Havana, Cuba) ;'' Fowler, Fish Guitarist, 21 (9), 1942: 66, fig. I (listed, Cuba) ; Bigelow and Schroeder,
Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 114 (ill.).

Genus Galeus Rafinesque, 1 8 10.

Galeus Rafinesque, Carratt. Gen. Nuov. Sicil., 1 8 10: 13; type species, G. melastomus Rafinesque, designated by
Fowler, Proc. Acad. nat. Sci. Philad., do, 1908: 53.^

Generic Synonyms:

Squalus (in part) Gunnerus, Trondh. Gesellsch. Schr. Leipzig, 2, 1766: 249; not Squdus Linnaeus, 1758.

Scyllium (in part) Risso, Ichthyol. Nice, 1 8 10: 30; not Scyllium Cuvier, 1 81 7.

Scylliorhinus (in part) Blainville, in Vieillot, Faune Franc, 1825: 68, 75.

Pristiurus^ Bonaparte, Icon. Faun. Ital., 5, 1834: 4th p. (not numbered) in description of "Scyllium caniculi^' ;

type species, P. meUmostomum Bonaparte; 1834, equals Galeus melastomus Rafinesque, 1 8 1 0.
Pristidurus^ Bonaparte, Mem. Soc. neuchatel. Sci. nat., 2 (8), 1839: 11 ; evident emendation of Pristiurus

Bonaparte, 1 834.
Figaro Whitley, Rec. Aust. Mus., t6, 1928: 238; type species, Figaro boardmani Whitley.

Generic Characters. Two dorsal fins, the ist originating over rear part of pelvics;
denticles along dorsal margin of anterior part of caudal enlarged and modified in shape,
forming a distinct crest, bounded below by a narrow band of naked skin on either side;
lower margin of caudal peduncle with or without a similar crest of enlarged denticles;
nostrils far from mouth and far apart, their anterior margins without barbels, their pos-
terior margins not expanded as flaps; snout long, thin, its mucous pores not conspicuous;
labial furrow extending from lower jaw around corner of mouth onto upper jaw, the upper
lip not closing outside lower at corner of mouth ; upper eyelid not closing outside lower
at corner of eye; a longitudinal fold or none below eye; 4th gill opening close in front
of pectoral, the 5th over pectoral; teeth alike in the 2 jaws, with long pointed median
cusp and i to 3 smaller cusps on each side, much as in Scyliorhinus, with several rows

35. Howell-Rivero states that the specimen on which he bases his new species, torrei, is the same one earlier referred
to by Sanchez-Roig as Catulus boae.

1. The name Galeus was first used by Klein, 1775 (Neuer Schauplatz) and by Valmont (Diet. Hist. Nat., /, 1798:
371) i but it must date from Rafinesque (Carrat. Gen. Nuov. Sicil., 18 10: 13), both Klein's and Valmont's names
having been ruled inapplicable by the International Committee on Zoological Nomenclature because such of them
as were binomial were so only accidentally (Smithson. misc. Coll., 75 [3], 1925: 27, Opinion 89). In his
account of the genus, Rafinesque mentioned only two species, melastomus Rafinesque and uyato Rafinesque, al-
though he expanded the genus to include seven species in his list of Sicilian fishes published later the same year
(Indice Ittiol. Sicil., 18 10).

2. The name Pristiurus has frequently been credited to Bonaparte, 1831 (Saggio Anim. Vert.: 121). But this first
mention of it was nominal only, without diagnosis or reference to any actual species, i.e., it was a nomen rmdem^.
For the actual dates of appearance of the individual plates and accompanying text of the Fauna Italica, see Sal-
vadori (Boll. Mus. Zool. Anat. comp. Torino, 5 [48], 1888).

3. The generic name Pristidurus was used a year earlier by L. Agassiz (Poiss. Foss., 3, 1838: 85) with a brief
account of the teeth, but without mention of any particular species.

Fishes of the Western North Atlantic 215

functional; tail sector of trunk considerably longer than body sector; caudal axis raised
but slightly, if at all; anal much longer than 2nd dorsal, separated from caudal by a
considerable interspace.

Remarks. Sharply diagnostic of this genus as contrasted with all other scyliorhinids
are the presence of the caudal crest and the wide separation of the nostrils from the mouth
and from each other in combination with the presence of a labial fold on each jaw, the
absence of a barbel and the absence or rudimentary state of the posterior nasal flap.

Range. Mediterranean; eastern North Atlantic northward to Norway; Iceland;
Madeira; Cuba and southern Florida in the western Atlantic; Japan; Formosa; Australia.

Key to Species

I a. Ventral margin of caudal peduncle, as well as anterior part of dorsal margin of caudal
fin, with a conspicuous crest of modified denticles. hoardmam Whitley, 1928.

Australia.

lb. Denticles along ventral margin of caudal peduncle not modified to form a crest.

2a. Tip of anal falls short of a vertical line at rear end of base of 2nd dorsal by a
distance about equal to that from eye to spiracle.
3a. Base of 2nd dorsal nearly twice as long as that of ist dorsal.

murinus Collett, 1905.

Iceland.

3b. Base of ist dorsal only about as long as that of 2nd dorsal.

eastmani Jordan and Snyder, 1904.

Japan.

2b. Tip of anal extends rearward nearly or quite as far as rear tip of 2nd dorsal.
4a. Trunk plain-colored.

5a. Interspace between anal and caudal about as long as snout in front of
mouth. jfl«/m* Jordan and Richardson, 1909.

Formosa.

5b. Interspace between anal and caudal less than V2 as long as snout in front
• of mouth. /<?»j^«iSaemundsson, 1922.

Iceland.
4b. Trunk marked with conspicuous dark stripes, spots or blotches.

6a. Interspace between anal and caudal at least ^/'o as long as base of anal;
base of anal only about twice as long as that of 2nd dorsal.

<zr«^ Nichols, 1927, p. 216.
6b. Interspace between anal and caudal only about Vi as long as base of anal;
base of anal about 3 times as long as that of 2nd dorsal.

W(9/tf.f/ow«J Rafinesque, 18 10.

Eastern North Atlantic, Mediterra-

4. Perhaps including hertijAgi Englehard, 191 2, Japan, the description of which is not sufficiently detailed for us to
locate it more precisely in this key.

2i6 Memoir Sears Foundation for Marine Research

Galeus arae (Nichols), 1927

Figures 36, 37

Study Material. 2i specimens, male and female, 138 to 329 mm. long, taken off the
north coast of Cuba, at "Atlantis" stations 2981, 2982, 2985, 2987, 3431, 3437, 3441 j
and near Tortugas, Florida, in 200 to 345 fathoms (Harv. Mus. Gjmp. Zool.)- Also a
specimen from Tortugas, Florida (U.S. Nat. Mus.).

Distinctive Characters. The presence of the caudal crest of large denticles marks
G. arae off from all other scyliorhinids yet known from the western Atlantic.

Descriftion. Proportional dimensions in per cent of total length. Female, 202 mm.,
from off Tortugas, Florida (Harv. Mus. Comp. Zool., No. 35250). Male, 324 mm.,
from Cuba (Harv. Mus. Comp. Zool., No. 361 18).

Trunk at origin of -pectoral: breadth 8.8, 9.25 height 7.4, 7.3.
Snout length in front of: outer nostrils 4.0, 3.45 mouth 7.6, 7. i .
Eye: horizontal diameter 4.2, 4.3.

Figure 36. Galeus arae, adult male, 324 mm. long, from off the north coast of Cuba (Harv. Mus. Comp.
Zool., No. 361 18). i4 Anterior part of head from below, about 1.4 x. 5 Pelvic fins and claspers, about 0.5 x natu-
ral size. C First to fourth upper teeth. D Twelfth and thirteenth upper teeth. E Thirty-fourth upper tooth.
F First to fifth lower teeeth. G Sixteenth and seventeenth lower teeth. H Twenty-third lower tooth. / Thirtieth
and thirty-first lower teeth. C-l, about 1 2 x. 7 Dermal denticles, about 60 x.

Fishes of the Western North Atlantic

217

Mouth: breadth 8.1, 7.7; height 3.5, 3.3.

Nostrils: distance between inner ends 2.8, 2.8.

Labial furrow length : upper 1.5, 1.75 lower 1.5, 1.8.

Gill Of ening lengths: ist 1.8, 1.4; 2nd 1.5, 1.35 3rd 1.2, 1.25 4th r.o, i.i; 5th

I.O, I.I.

First dorsal fin: vertical height 3.9, 4.2; length of base 6.4, 5.3.

Second dorsal fin: vertical height 3.7, 4.O} length of base 5.9, 5.2.

Anal fin: vertical height 3.5, 3.5; length of base 13.5, 11.4.

Caudal fin: upper margin 30.2, 29.2; lower anterior margin 11.5, 10.2.

Pectoral fin: outer ma.rgm 12.2, 10.8; inner margin 6.5, 5.9; distal margin 9.3, 9.0.

Distance from snout to: ist dorsal 43.3, 45.7; 2nd dorsal 59.7, 64.O; upper caudal

69.8,72.8; pectoral 16.8, 19. i; pelvics 37.7, 37.7; anal 51.5, 56.0.

Interspace between: ist and 2nd dorsals 12.2, 13.O; 2nd dorsal and caudal 3.2,

3.7; anal and caudal 4.0, 4.6.

Distance from origin to origin of : pectoral a.ndpelv\cs 19. i, 18.6; pelvics and anal

14.7, 19.4.

Figure 37. Galetis arae. Dermal denticles from dor-
sal margin of caudal. A From above. B From side,
about 17 X.

Trunk slender, its breadth opposite pectorals about ^Aoj and its height about Vi2> of
total length. Body sector to cloaca considerably shorter than tail sector. Dermal denticles

2 1 8 Memoir Sears Foundation for Marine Research

on trunk close-spaced, their blades only slightly raised, with 3 low ridges, their posterior
margins with 3 strong teeth, the median much the longest; the 2 or 3 rows along the dorsal
margin of the anterior half of the caudal larger, only weakly dentate and without ridges,
flanked on either hand by a single row of very much larger blade-like denticles, lanceolate
in shape, their inner margins with a deep notch, their tips directed outward and bounded
below by a narrow band of naked skin forming a noticeable crest, but grading rearward
into denticles of the usual size and shape.

Head strongly flattened above. Snout thin, broadly rounded in front and slightly
narrowed opposite nostrils, its length in front of mouth about Vs of length of head to origin
of pectoral. Eye narrow-oval, its horizontal diameter a little more than Yo as long as
snout in front of mouth, with a weakly marked longitudinal fold below it. Spiracle oval,
its diameter about Yi as great as that of eye, behind the latter by a distance about Yz as
great as the horizontal diameter of eye, and a little below it. Gill openings concave ante-
riorly in outline, the 4th and 5th closest together, the ist and 2nd (slightly the longest)
about Ys as long as horizontal diameter of eye, the 5th (shortest) about % as long as ist;
the 5th above or a little posterior to origin of pectoral. Nostrils oblique, separated one from
the other by a distance equal to about % the length of snout in front of mouth, and sepa-
rated from mouth by a distance about Y2 that great, the anterior margin expanded as a low,
subtriangular lobe with rounded tip, the posterior margin not expanded. Mouth obtusely
ovate, about V2 as long as broad, with labial furrows extending a short distance inward
along both jaws.

Teeth about ffEffj with slender median cusp, and a much smaller cusp on each side in
central part of mouth, but usually with 2, or even 3, lateral cusps on each side toward cor-
ners of mouth; 4 or 5 series functional in front of mouth, with 2 to 3 toward its corners
in upper jaw and 3 to 4 series in lower jaw.

Dorsals small, similar in size and shape, quadrate, with weakly convex anterior mar-
gins, straight distal margins and subrectangular corners, their bases about as long as snout
in front of eye or a little shorter, their free lower margins about Y2 as long as their bases
or a little less; origin of ist dorsal over rear Ys of bases of pelvics, origin of 2nd dorsal
about over midpoint of base of anal. Caudal about Y-i of total length, and noticeably
narrow, its axis only very slightly raised, its tip squarely truncate posteriorly, its lower
anterior corner much more obtuse than a right angle, the subterminal notch scarcely
marked. Interspace between caudal and anal varying from about Y2 as long as base of anal
to almost as long as latter. Anal about twice as long at base as 2nd dorsal, Its rear tip a little
anterior to rear tip of latter, with nearly straight margins and rounded apex, its free basal
margin very short. Pelvics with broadly rounded apices and tapering, blunted tips, widely
divergent in adults but less so in smaller specimens, their inner edges joined and attached
to ventral surface of trunk for about Y2 their lengths posterior to cloaca, both in males and
in females. Pectoral with very broad base, nearly straight margins and broadly rounded
corners, about as broad as long, and about 3 times as large in area as ist dorsal.

Color. Ground tint pale yellowish brown, strikingly marked along sides and back

Fishes of the Western North Atlantic 219

with rows of dark brown blotches and spots of various sizes, forming an especially intricate
pattern on top of headj a dark streak from snout to eyej a large and conspicuous blotch
extending up onto each dorsal fin, one on the upper half of the caudal near its anterior
end, one on its lower part and another across it abreast of the subterminal notch. The pre-
cise sizes, shapes and arrangements of the finer markings vary considerably, and they be-
come more or less confluent on the larger specimens, in which the upper surface, anterior
to the first dorsal fin, has a clouded rather than a spotted and striped appearance} roof of
mouth dusky or sooty, tongue and floor of mouth similar in some specimens, but pale in
others, perhaps faded in the preservative.

Size. The largest specimen yet seen is a male of 329 mm. (listed on p. 216). Since
the daspers fall considerably short of the tips of the pelvics in a specimen of about 295
mm., but extend far beyond them in another of about 317 mm., maturity is probably at-
tained at about 300 mm.

Develof mental Stages. Presumably oviparous, but the eggs have not been seen.

Habits. Knowledge of the habits of G. arae is confined to the fact that it is a deep-
water species; recorded depths of capture range from 200 down to 345 fathoms.

Range. So far known only off the north coast of Cuba where it is evidently common
at suitable depths, off the Tortugas, Florida, and off Miami, Florida (the type locality).

Synonyms and References:

Pristiurus arae Nichols, Amer. Mus. Novitates, 256, 1927: I, fig. I (descr., off Miami Beach, Florida) ; Jordan,

Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 12 (off Florida).
Scyliorhinus retifer Longley and Hildebrand, Pap. Tortugas Lab., ^4, 1 941: I (depth, color, Tortugas,

Florida).*

Genus Apristurus Garman, 19 13

Aprisiurus Garman, Mem. Harv. Mus. comp. Zool., j(5, 1 91 3: 96; type species, Scylliorhinus indicus, Brauer,
Wiss. Ergebn. 'Valdivia,' 75, 1908: 8, pi. 14, fig. I (Indian Ocean and Gulf of Aden), desig. by Jordan,
Genera Fish., 4, 1920: 548.

Generic Synonyms:

Catulus (in part) Gilbert, Proc. U.S. nat. Mus., 14, 1 891: 542; for C brunneus Gilbert (west coast of N.

America) ; not Catulus A. Smith, 1837.
Scylliorhinus (in part) Brauer, Wiss. Ergebn. 'Valdivia,' 15, 1908: 8; and subsequent authors; not Scyliorhinus

Blainville, 1 8 16.
Pristiurus (in part) Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 94; for Catulus sfongicefs Gilbert,

1905; not Pristiurus Bonaparte, 1834.
Scyllium Saemundsson, Vidensk. Medd. naturh. Foren. Kbh., 7./, 1922: 173; type species S. laurussonii Sae-

mundsson, monotypic; not Scyllium Cuvier, 1 81 7.
Ap-isturius Schulze, Kukenthal, et al., Nomencl. Anim., i (2), 1926: 244; evident misspelling for Afristurus

Garman, 191 3.
Pentanchus (in part) Fowler, Proc. Acad. nat. Sci. Philad., 85, 1934: 237; not Pentanchus Smith and Rad-

cliffe, Proc. U.S. nat. Mus., 41, 191 2: 490.
Pttrafristurus Fowler, Proc. Acad. nat. Sci. Philad., 8s, 1934: ^37; type species, Catulus sfongicep Gilbert,

1905.

5. We have examined and identified this specimen, now in the United States National Museum.

2 20 Memoir Sears Foundation for Marine Research

Generic Characters. Two dorsal fins, the origin of the ist considerably posterior to
origin of pelvicsj dermal denticles along dorsal margin of anterior part of caudal not
enlarged or modified as a distinct crest, or bounded below by a band of naked skin on either
side; nostrils entirely separate from mouth, their anterior margins without barbels and
falling considerably short of mouth j posterior as well as upper margin of nostril expanded
as a flap; labial furrow around corner of mouth and on each jaw; interspace between anal
and caudal less than Vs as long as base of anal; base of anal more than twice as long as base
of 2nd dorsal; no fold below eye; mucous pore system on lower surface of snout very con-
spicuous; gill openings either of the usual conformation, or so deeply concave anteriorly
that tips of gill filaments are exposed; 5th gill opening over or behind origin of pectoral;
teeth numerous, those in front of mouth with one chief cusp and one or more smaller cusps
on each side; several series of teeth functional.

Range. Both sides of North Atlantic, including Iceland; South Africa; west coast of
North America from Gulf of California to Puget Sound; Hawaiian Islands; Japan;
Philippines and East Indies; Indian Ocean and Gulf of Aden; west coast of South Africa.

Species. These are little known sharks of deep water, the majority of them so far
known from very few specimens. The named species of the genus, numbering 13 and
from widely separated seas, resemble one another very closely in general appearance, but
they appear to be separable by sufficiently precise differences. Fowler has even distributed
them among three subgenera,^ according to the degree of cirrus-like development on the
margins of the nostrils, anterior and posterior. According to published accounts, however,
and to our own study of three of the species, the differences in this respect are not sharp
enough to serve as a basis for generic separation. Nevertheless, the members of the genus
do fall into two sharply contrasting categories as regards the gill openings, for while
these are of the ordinary type in one group, typified by A. brunneus Gilbert from the
west coast of North America, they are close together above and below in other species,
but so deeply concave anteriorly at the midlevel that the tips of the gill filaments are
exposed (p. 227). It is astonishing that attention has not been directed to this earlier,
for the gills are clearly pictured thus mA. atlanticus Koefoed," as well as in ^ . microps Gil-
christ.' Furthermore, a re-examination of the specimens in the United States National
Museum shows gills of this same type in profundorum Goode and Bean, 1895 (p. 222),
verweyi Fowler, herklotsi Fowler, 1934,* spongiceps Gilbert, 1905, and platyrhynchus
Tanaka, 1909,° although no suggestion of the fact appears in the published accounts
or in the illustrations of these species. Under ordinary circumstances a diflterence so
striking would demand the institution of a new genus. In the present case, however, such
action does not seem advisable because neither the account nor the illustration of the type

1. Bull. U.S. nat. Mus., loo (15), 1941: 53; Parapristurus, Pentanchus and Afristurus.

2. Koefoed, Rep. Sars N. Atlantic Deep Sea Exped., 4 (i), 1932: 18, pi. 3.

3. Mar. biol. Rep. Cape Town, 2, 1922; 46, pi. 7, fig. i.
4.. Proc. Acad. nat. Sci. Philad., ^5, 1934: 237, 238.

5. There is a specimen of this species in the United States National Museum, although not the type.

Fishes of the Western North Atlantic 221

species of Afristurus'^ gives any Information as to its gill openings, i.e., there is no way of
knowing to which subdivision of the old genus Afristurus it belongs; nor are the specimens
available for study at present, being presumably in Berlin. Therefore, it seems wiser to
use Afristurus in the more inclusive sense for the time being/ Neither can a dependable
Key to Species be constructed for the genus as a whole until more complete information is
available in other respects regarding indicus Brauer, also one of the two supposedly distinct
species that have been named from Japan," and sibogae Weber" from the East Indies."

Key to Atlantic and South African Species

I a. Distance between ist and 2nd dorsal fins as great as from tip of snout to spiracle.

saldanha Barnard, 1925.
South Africa.
lb. Distance between ist and 2nd dorsal fins at least no greater than from tip of snout
to eye.

2a. Interspace between ist and 2nd dorsals less than ^/^ as long as from tip of snout
to eye; eye minute, its diameter only about y\ \ as long as from tip of snout to 5th
gill opening. wicro^j Gilchrist, 1922.

South Africa.
2b. Interspace between ist and 2nd dorsals nearly or quite as long as from tip of snout
to eye; eye larger, its diameter at least % as long as head to 5th gill slit.
3a. Second dorsal about twice as large in area as ist; ist to 3rd gill openings
nearly as long as distance between nostrils.

riveri Bigelow and Schroeder, 1944, p. 225.
3b. Second dorsal little if any larger in area than 1st; ist to 3rd gill openings
only about V2 as long as distance between nostrils.

4a. Horizontal diameter of eye slightly longer than distance between nos-
trils; rear ends of bases of pelvics slightly nearer to tip of snout than to
tip of caudal; caudal about Vs of total length, atlanticus Koefoed, 1932.

Eastern North Atlantic.

4b. Horizontal diameter of eye only about % as long as distance between
nostrils; rear ends of bases of pelvics nearer to tip of caudal than to tip
of snout; caudal only about ^/4 of total length.

frojundorum^^ Goode and Bean, 1895, p. 222.

6. indicus Brauer, Wiss. Ergebn. 'Valdivia,' 1$, 1908: 8, pi. 14., fig. i.

7. For further discussion, see Bigelow and Schroeder (Proc. New Engl. zool. CI., 35, 1944; ^O-
X. macrorhynchus Tanaka, J. Coll. Sci. Tokyo, 27, 1909; i.

9. Siboga Exped., 57, 1913: 595.

10. For comparable illustrations of spongicefs Gilbert, 1905, from the Hawaiian Islands, vervieyi Fowler, 1934.
from Borneo, herklotsi Fowler, 1934, of the Philippines, and flatyrkynchus Tanaka, 1909, from Japan, see
Fowler (Bull. U.S. nat. Mus., 100 [z^], 1941: Si-SSt Sj)-

11. Including /aurui^omi, Saemundsson, 1922, Iceland.

222 Memoir Sears Foundation for Marine Research

Afristurus frojundorum (Goode and Bean), 1 895
Figure 38

Study Material. Type specimen, mature male, 510 mm. long, taken off Delaware
Bay in 816 fathoms (U.S. Nat. Mus., No. 35646) j newly hatched male, 145.5 mm.., from
a nearby locality (U.S. Nat. Mus., No. 83894).

Distinctive Characters. The adult profundorum is separated from riveri by its con-
siderably smaller second dorsal relative to the first dorsal, shorter snout relative to length
of head, smaller eye, much shorter gill openings, relatively broader mouth and much
shorter caudal 5 from atlanticus by its relatively smaller eye, shorter caudal, and by the fact
that the tips of the pelvics are closer to the tip of the caudal than to the tip of the snout.
But it may be difficult to distinguish newly hatched specimens of the three species from
one another.

Figure 38. Afristurus frofumlorum, female, 510 mm. long, from off Delaware Bay (U. S. Nat. Mus.,
No. 35646, type). A Head of same from below. B Dermal denticles of same, about 12 x. C Newly hatched
male, 146 mm. long, from off Delaware Bay (U. S. Nat. Mus., No. 83894). D Head and pectorals of same
from below, about 1. 1 m. E Gill openings of same, about 4 x.

Description. Proportional dimensions in per cent of total length. Male, 510 mm.,
from Lat. 39° N., Long. 72° W. (U.S. Nat. Mus., type. No. 35646).
Trunk at origin of pectoral: breadth i i.O; height 9.2.
Snout length in front of: mouth 8.9.
Eye: horizontal diameter 2.7.
Mouth: breadth 8.45 height 2.9.
Nostrils: distance between inner ends 4. i .
Labial furrow lengths: upper 2.95 lower 3.5.
Gill opening lengths: ist 1.85 5th 1.3.

Fishes of the Western North Atlantic 223

First dorsal fin: vertical height 3.2; length of base 7.0.
Second dorsal fin: vertical height 3.3 ; length of base 6.9.
Anal fin: vertical height 4.35 length of base 13.9.
Caudal fin: upper margin 25.0.

Pectoral fin: outer margin 10.6; inner margin 6.4; distal margin 5.1.
Distance from snout to: ist dorsal 49.5; 2nd dorsal 62.5; upper caudal 75.O; pec-
toral 24.75 pelvics 43.3; anal 56.6.

Interspace between: ist and 2nd dorsals 8.2; 2nd dorsal and caudal about 3; anal
and caudal 0.0.
Distance from origin to origin of: pectoral and pelvics 19.6; pelvics and anal 12.5.

Trunk slender, highest opposite axil of pectoral, tapering evenly rearward, its
height at axil of pectoral (where highest) about Vr its length to origin of caudal. Body
sector to cloaca a little longer than tail sector. Dermal denticles with 3 ridges and 3 teeth,
as in riveri, but with the teeth shorter and overlapping more, so that the skin is more con-
cealed.

Head about Vi of total length, flattened above, and contracted laterally just anterior
to outer ends of nostrils. Snout broadly rounded, its length in front of mouth a little more
than Vs as great as length of head to origin of pectoral, with a median belt of conspicuous
mucous pores in 8 or 9 irregular rows on its ventral surface. Eye oval, its horizontal diame-
ter about % as great as distance between nostrils, its midpoint opposite corner of mouth.
Spiracle oval, its diameter about Yz as great as that of eye, and behind latter by a distance
about % as great as diameter of eye. Gill openings much smaller relatively than in riveri,
the 1st to 3rd (longest) a little less than V2 as long as distance between nostrils, or about
% as long as horizontal diameter of eye, the 5th (shortest) about % as long as ist, of the
same general type as in riveri (p. 227), their anterior outlines so deeply concave that the
tips of the gill filaments are exposed on all 5 of the interbranchial septa; the 4th and 5th
over origin of pectoral. Nostrils moderately oblique, at margins of head, their outer ends
about equidistant between tip of snout and center of mouth, the distance between them a
little less than Y-y as great as length of snout in front of mouth, the anterior margins more
broadly rounded than in riveri (the condition of the specimen is not good enough for de-
scription of the inward cirroid extensions of the nostril, if any). Mouth ovate, nearly 3
times as broad as high. Labial furrows very prominent, the upper extending about V2 the
distance toward the symphysis, and more nearly parallel with the jaw than in riveri, the
lower a little shorter than the upper.

Teeth about ||^; uppers with long, sharp median cusp, flanked on either side by
2 or 3 smaller cusps; lowers similar to uppers, except with the lateral cusps somewhat
larger relative to the median cusp, and more often 3 in number on one or both sides; no
median tooth in either jaw; several series functional.

Dorsals similar in form, brush-shaped, with rounded tips and weakly convex anterior
margins. Origin of ist about over midpoint of bases of pelvics, its base a little less than %

224 Memoir Sears Foundation for Marine Research

as long as snout in front of mouth, its rear tip about over origin of anal. Second dorsal
about as long at base as ist dorsal, and only a very little larger than the latter in area, if at
all so, its origin about over midpoint of base of anal. No definite interspace between 2nd
dorsal and caudal. Caudal about % of total length, with rounded tip and weakly marked
subterminal notch, its lower anterior corner subangular, its axis not appreciably raised
above main axis of trunk. No measurable interspace between lower origin of caudal and
rear end of base of anal. Anal a nearly equilateral and very obtuse triangle, with nearly
straight edges, slightly rounded corners and very short free tip, its origin about under tip
of I St dorsal, its base about twice as long as that of 2nd dorsal. Pelvics quadrate, with nearly
straight edges and blunted corners, apices broadly rounded, the rear corners more nar-
rowly so. Pectoral more than twice as large in area as ist dorsal, brush-shaped, about as
broad at base as at tip, with rounded corners, the outer margin nearly straight, but the
distal and inner margins moderately convex.

Color. Uniform grayish brown below as well as above after preservation in alcohol.

Size. The fact that the claspers of the type specimen (510 mm. long) are only mod-
erately developed suggests that this deep-sea shark does not mature until a length of
perhaps 550 to 600 mm. is reached.

Developmental Stages. Presumably A . frofundorum is oviparous, but its eggs have
not been identified, although Gudger^" suggests that certain egg cases found on the coast
of North Carolina might be of this parentage. If the very small specimen listed above and
illustrated in Fig. 38 C actually belongs to this species and not to riveri, as seems probable
(from the shortness of its gill openings, its small eyes, as well as from the locality of its
capture), frojundorum more closely resembles riveri wlien newly hatched than later
in growth, for the length of its caudal is then as great as in riveri (about V3 of total length)
and its second dorsal considerably larger than its first dorsal. Furthermore, the snout is
considerably longer, relatively, in newborn specimens than in adults of either of the two
possible parent species, since it occupies considerably more than one-third of the length
of the head, and the anal is actually confluent with the lower edge of the caudal. More
interesting still is the great breadth of the basal lines of attachment of the pectorals to the
lower sides of the trunk (Fig. 38 C).

Habits. Nothing is known positively of its habits, but the depth of capture listed
above and its uniformly dark coloration above and below suggest a deep-sea habitat.

Range. A.-profundorum is definitely known only from the continental slope off Dela-
ware Bay and from the specimen (or specimens) listed above (p. 222). However, if A.
laurussonii from Iceland is identical with it, as the only published account of laurussonii
suggests, it is no doubt wide-ranging around the slopes of the northern North Atlantic in
the appropriate latitudinal belt."

12. Proc. biol. Soc. Wash., 25, 19 12: 154.

13. Profunilorum has also been reported from British Columbia (Halkett, Check List Fish. Canad., 1913; ii?.)-
But probably the shark in question was actually A. brunneu; Gilbert, which is rath.er common along the Pacific
coast of North America from the Gulf of California northward, in deep water.

Fishes of the Western North Atlantic

225

Synonyms and References:

Scylliorhinus frojundorum Goode and Bean, Smithson. Contr. Knowl., 50, 1895: 17, pi. 5, fig. 16; Mem.
Harv. Mus. comp. Zool., 22; Spec. Bull. U.S. nat. Mus., 2, same date and p.igination (descr., off Dela-
ware Bay, Lat. 39° 9' N., Long. 72° 3' W., 816 fathoms); Jordan and Evermann, Rep. U.S. Comm.
Fish. (1895), 1896: 213 (n.ime, N. Atlant.) ; Bull. U.S. nat. Mus., 47 (i), 1896: 22 (descr.); Bull.
U.S. nat. Mus., 47 (4), 1900: pi. 3, fig. 11 j Gudger, Proc. bid. Soc. Wash., 2^, 1913: 154 (egg cases
perhaps this species).

Afristurus frojundorum Garman, Mem. Harv. Mus. comp. Zool., j(5, 191 3: 99 (descr.) ; Fowler, Copeia, 30,
1916: 36; Smith, J. Amer. Mus. nat. Hist., 7(5, 1916: 349 (ref. to type specimen) ; Jordan, Evermann
and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 12 (ref. to type specimen).

Not Scylliorhinus frojundorum Halkett, Check List Fish. Canad., 191 3: 1 17.

Probable synonym :

Scyllium laurussonii Saemundsson, Vidensk Medd. naturh. Foren. Kbh., 74, 1922:

[73, pi. 4, fig. I (Iceland).

Afristurus riveri Bigelow and Schroeder, 1944

Figure 39

Study Material. The type specimen, female, 407 mm. long, trawled at "Atlantis"
Sta. 2993, off north coast of Cuba, March 15, 1938, in 580 fathoms (Harv. Mus. Comp.
Zool., No. 36092).

Figure 39. Afristurus rk'eri, female, 407 mm. long, from off northern Cuba (Harv. Mus. Comp. Zool.,
No. 36092, type). A Head from below. B Gill openings, about 1.8 x. C Right-hand nostril, about 3 x. Z) Gen-
eral view of dermal denticles, about 22 x; lateral and apical views, about 43 x. E Upper and lower teeth from
near center of mouth. F Upper and lower teeth from outer parts of jaws, about 16 x.

226 Memoir Sears Foundation for Marine Research

Distinctive Characters. The adult is separated from A. frofundorum, A. laurussonii
and A. atlanticus^* by the considerably greater size of its second dorsal relative to the
first, and by the fact that its first to third gill openings are about as long as the distance
between the nostrils but only about one-half that relative length in the other three species.
It is further separated from frofundorum (the only local species with which it might be
confused) by its relatively larger eye, and by the fact that Its caudal occupies about one-
third of the total length as compared to only about one-quarter. Young specimens of the
diflFerent species may be difficult to separate.

Description. Proportional dimensions in per cent of total length. Female, 407 mm.,
from Cuba (Harv. Mus. Comp. Zool., type. No. 36092).

Trunk at origin of fectoral: breadth 10. i ; height 1 1.8.

Snout length in front of: outer nostrils 5.85 mouth 9.3.

Eye: horizontal diameter 2.7.

Mouth: breadth 6.1 ; height 2.2.

Nostrils: distance between inner ends 3.9.

Labial furrow lengths: upper 2.1 ; lower 2.5.

Gill opening lengths: ist 3.35 2nd 3.4; 3rd 3.2; 4th 3.O5 5th 2.1.

First dorsal fin: vertical height 2.9; length of base 4.4.

Second dorsal fin: vertical height 4.23 length of base 6.1.

Anal fin: vertical height 3.7; length of base 13.7.

Caudal fin: upper margin 33.O5 lower anterior margin lO. i.

Pectoral fin: outer margin lO.i ; inner margin 4.9; distal margin 7.7.

Distance from snout to: ist dorsal 47.85 2nd dorsal 57.7; upper caudal 67.O; pec-
toral 23.9; pelvics 40.2; anal 52.2.

Interspace between: ist and 2nd dorsals 7.O5 2nd dorsal and caudal Indefinite;

anal and caudal 0.0.

Distance from origin to origin of: pectoral and pelvics 18.5; pelvics and anal 1 2.5.

Trunk highest and broadest opposite axil of pectoral, narrowing evenly rearward.
Caudal peduncle strongly compressed laterally, about V2 as broad as deep. Body sector to
cloaca about as long as tail sector. Dermal denticles small in specimen examined, moder-
ately erect, leaf-like, with short pedicels, their blades with weak median crest but tridentate
free margin, the median tooth much the longest; the denticles slightly the largest, rela-
tively the narrowest, and the most closely spaced along upper sides of caudal.

Head strongly flattened anteriorly, and contracted laterally just anterior to the
outer ends of the nostrils, its dorsal surface noticeably concave, with a triangular belt of
about 1 10 very prominent pores along the midzone anterior to eyes. Snout thin, broadly
rounded at tip, its length in front of mouth slightly less than y^. as great as distance from
its tip to 5th gill opening, its lower surface with a belt of conspicuous pores in 4 rows along

14. The only other members of the genus yet known from the North Atlantic.

Fishes of the Western North Atlantic 227

the midzone. Eye oval, its horizontal diameter about twice its vertical height and nearly as
long as distance between nostrils, its midpoint about opposite corner of mouth. Spiracle
round, about ^'5 as long as horizontal diameter of eye, situated close behind latter. Gill
openings with their anterior margins so deeply concave in outline that the tips of the gill
filaments on the ist to 4th arches are exposed, but with the dorsal and ventral ends so close
together that the successive margins form an apparent frame around the gill area as a
whole; ist to 4th gill openings about as long as horizontal diameter of eye, or about 4
times as long as from posterior margin of eye to spiracle, the 5th considerably shortest, and
close in front of origin of pectoral. Nostrils moderately oblique and far apart, their inner
corners about 3 times as far from tip of snout as from symphysis of upper jaw, their outer
corners at outer edge of snout, their anterior margins obtusely triangular in outline, the
posterior margin of nostril also expanded, as shown in Fig. 39 C. Mouth obtusely ovate,
nearly 3 times as broad as high, occupying only about ^/'o the breadth of head. Labial
furrows very prominent, forming approximately a right angle at corner of mouth when
latter is closed, the upper extending forward about Vs the distance toward outer end of
nostril, the lower (slightly the longer) reaching only a short distance past corner of
mouth.

Teeth, about HeH j uppers with 3 cusps, the median erect and much the longest,
except that there are 2 small cusps on one or both sides toward corners of mouth, with
the median cusp curved outward; lowers similar to uppers in front of mouth, but usually
with 5 cusps along sides of mouth, the median only a little the longest; no tooth at sym-
physis of either jaw; a very small tooth next to the symphysis in lower; mostly 3 series
functional in upper jaw, but 3 to 4 in lower.

First dorsal very small, its base a little longer than horizontal diameter of eye, brush-
shaped, with convex anterior margin and rounded tip, its origin over rear ends of bases of
pelvics. Second dorsal similar to first but nearly i ^o times as long (at base) and i ^2 times
as high vertically (correspondingly larger in area), its origin about over midpoint of base of
anal. Caudal about Vs of total length, with brush-shaped tip and well marked subterminal
notch, its lower anterior corner more obtuse than a right angle, its axis only very slightly
raised, if at all. No measurable interspace between caudal and anal. Anal with rounded an-
terior corner and angular rear corner, its base slightly more than twice as long as base of
2nd dorsal. Pelvics a little less than V2 as long at base as anal, with rounded anterior and
angular rear corners, their inner margins very short and entirely separate one from the
other behind the cloaca in female. Interspace between pelvics and anal about % as long as
base of former. Pectoral a little more than '^/^ as long as head, nearly as broad at base as at
tip, with slightly convex outer margin, nearly straight distal and inner margins and
rounded corners.

Color. Uniform chocolate-brown above and below, in alcohol, the tongue and lining
of the mouth blackish.

228 Memoir Sears Foundation for Marine Research

Size. No information is available as to the length to which this species may grow.
The specimen at hand does not contain eggs.

Developmental Stages. Not known; see frojundorum, p. 224.

Range. Known only from the specimen recorded above, taken off the north coast of
Cuba. The depth of capture, combined with the uniformly dark coloration, suggests that
this is a deep-water species.

Synonym and Reference:

Afristurus riveri Bigelow and Schroeder, Proc. New Engl. zool. CL, 2j, 1944: 23, pi. 7 (descr, ill., type loc.
off the north coast of Cuba, "Atlantis" Sta. 2993, 580 fathoms).

Family PSEUDOTRIAKIDAE
False Cat Sharks

Characters. Two dorsal fins, the ist as long as caudal,^ or longer, the rear end
of its base over or a little anterior to origin of pelvics ; 2nd dorsal as high as ist or higher;
caudal less than ^/4 of total length, not lunate, its lower anterior corner not expanded as a
definite lobe, its axis but little raised; caudal peduncle not flattened dorso-ventrally or
expanded laterally, without precaudal pits above or below; sides of trunk without longi-
tudinal dermal ridges; snout not greatly elongate; jaws not widely protrusible; gill open-
ings very short, 5th over origin of pectoral; gill arches without rakers and not intercon-
nected by a sieve of modified denticles; nostril entirely separate from mouth, its anterior
margin without barbel; spiracles present; lower eyelid without nictitating membrane, but
with a well marked longitudinal fold; teeth small, numerous, with larger median and
smaller lateral cusps, 6 to 13 series functional; head of normal shape, not widely expanded
laterally; rostral cartilages 3, united terminally; radials of pectoral mostly on metapter-
ygium, those on meso- and propterygia fused; mesopterygium and propterygium much
smaller than metapterygium." Development ovoviviparous.

Genera. One genus only.

Genus Pseudotriakis Brito Capello, 1867

Pseudotriakis Brito Capello, J. Sci. math. phys. nat. Lisboa, i (2), 1867: 315, 321 ; type species, P. microdon
Brito Capello, Setubal, Portugal.

Generic Characters. Those of the family.

Range. Both sides of North Atlantic, in deep water; Japan.

Sfecies. Only two species known.

1. The very long first dorsal is the most striking character of this family.

2. For detailed account and illustrations of the skeleton, see Jacquet (Bull. Inst, oceanogr. Monaco, 3d, 1905).

Fishes of the Western North Atlantic 22g

Key to Species

I a. Caudal about V4 of total length} origin of anal only a little posterior to origin of ist
dorsal; distance from tip of snout to angle of mouth only about V4 as long as from
snout to 5th gill opening; length of snout in front of mouth about V3 as great as width
of mouth. acrages Jordan and Snyder, 1904.

Japan.'

lb. Caudal only about y^ of total length; origin of anal considerably posterior to origin
of 1st dorsal; distance from snout to angle of mouth Y2 as great as from snout to 5th
gill opening or greater; length of snout in front of mouth about V2 as great as width of
mouth. microdon Brito Capello, 1 867, p. 229.

Pseudotriakis microdon Brito Capello, 1867

Figure 40

Study Material. None.

Distinctive Characters. Separable from all other Atlantic sharks by the great length of
its first dorsal fin.

Description. Proportional dimensions in per cent of total length. Specimen, 2,950
mm., from Amagansett, N. Y. (after Goode and Bean).

Trunk at origin of ist dorsal: breadth 8.5; height 12.0.

Snout length in front of: mouth 3.0.

Eye: horizontal diameter 2.3.

Mouth: breadth 9.0.

Nostrils: distance between inner ends 4.2.

Gill ofening lengths: ist 2.6.

First dorsal fin: greatest height 3.2; length of base 22.7.

Second dorsal fin: greatest height 5.4; length of base 12.5.

Anal fin: greatest height 4.0; length of base 8.5.

Caudal fin: upper margin 18.O; lower anterior margin 7.7.

Pectoral fin: greatest length 1 1.2; greatest width 8.0.

Distance from snout to: ist dorsal 34.O; 2nd dorsal 67.O; upper caudal 82.O; pec-
toral 20.0; pelvics 5 6.0; anal 70.7.

Intersface between: ist and 2nd dorsals 10.5; 2nd dorsal and caudal 3.9.

Male embryo, ready for birth, 850 mm., from Iceland (after measurements by Saemunds-
son).

Snout length in front of: mouth 5.3.

First dorsal fin: height 4.1 ; length 21.2.

3. For these and other differences in proportionate measurements, see Jordan and Snyder (Smithson. misc. Coll., ^5,
1904: 233). The original spelling was "acrales," but Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 105)
has pointed out that this was a misprint.

230 Memoir Sears Foundation for Marine Research

Second dorsal fin: height 6.2 ; length 1 1 .7.
Anal fin: height 3.85 length 9.0.
Caudal fin: upper margin 19.4.
Pectoral fin: \tngth. 1 0.0.

Distance froin snout to: ist dorsal 32.4; 2nd dorsal 62.35 upper caudal 80.65 Pec-
toral 20.0; pelvics 52.93 anal 65.3.

"iilili

Figure 40. Pseudotriakis microdon, eastern Atlantic specimen, 930 mm. long, drawn by A. Fraser-Brunner,
from skin preserved in alcohol in the British Museum. A Head from below. B Upper and lower teeth to
show mosaic arrangement. C Front and side views of upper and lower tooth, enlarged. D Dermal denticles,
enlarged.

Trunk subcylindrical, its height at origin of ist dorsal about Y-, of length to origin
of caudal in large specimens, but a little less than Y\ 0 in small. Body cavity notably long,
the distance from origin of pectoral to cloaca being V2 the length of trunk, or a little more,
with origin of pelvics considerably closer to tip of caudal than to snout. Dermal denticles
lanceolate, with i to 5 low longitudinal ridges, raised steeply from the skin on short
pedicels.

Head about Vs of total length and somewhat flattened above. Snout in front of mouth

Fishes of the Western North Atlantic 231

a little less than ^6 to V? of length of head to 5th gill opening in large specimens, but rela-
tively somewhat longer in small (Fig. 40 A), ovate with rounded tip. Eye oval, with fold
below it. Spiracle oval, about as long as diameter of eye or longer, its long axis oblique,
situated close behind the eye. Second gill opening a little longer than diameter of eye in
large specimens, but a little shorter than eye in small. Nostrils far apart and much closer
to mouth than to tip of snout, only slightly oblique, the anterior margins with a low, sub-
triangular lobe. Mouth rounded in front with nearly straight sides, its height between %
and % its width. A well marked labial furrow on upper jaw extending Va to ^4 the dis-
tance toward symphysis, the lower furrow very short.

Teeth minute, extremely numerous, arranged in mosaic j smooth-edged; uppers with
3 to 5 triangular cusps, the median much the largest in front part of mouth, but the
laterals more nearly equalling it toward corners of mouth; lowers with 3 erect cusps in
front of mouth (the median longest) but usually with 4 along its sides, the cusps of the
outermost being of nearly equal lengths;' about 6 to 13 series functional, more being so
in lower jaw than in upper.

First dorsal very sloping, about Ve to Vr as high as the length at base, relatively some-
what higher in small specimens than in large (see proportional dimensions, p. 229),
convex in upper contours, but without definite apex, its free rear margin very short with
acute tip, its origin V2 to Vs as far from axil of pectoral as from origin of pelvics, its base
between % and Vs as long as the total length, and about i Vs times as long as caudal in
large specimens, but only about as long as caudal in small, the rear end of its base termi-
nating about over origin of pelvics or a little anterior to the latter. Second dorsal a little
higher than i st in small specimens and about 1.7 times that high in large, about twice as long
as high, subtriangular, with weakly convex anterior margin, very weakly concave or nearly
straight distal margin, broadly rounded apex and very short free rear corner, its origin a
little posterior to tips of pelvics, its tip over or a little posterior to tip of anal. Caudal only
between Y^ and % of total length, slightly less than V2 as broad as long, with well marked
subterminal notch, its posterior outline nearly straight, its lower anterior corner obtuse,
subangular. Anal similar to 2nd dorsal, but only about % as long at base and % as high,
its origin a little posterior to origin of 2nd dorsal. Pelvics Vs to ^i as long at base as ist
dorsal. Pectoral noticeably small, about 1.3 times as long as base of anal, a little more than
% as broad as long, with nearly straight distal margin, weakly convex outer margin,
rounded corners and broad base.

Color. Described as uniformly dark brownish gray, darkest on posterior margins of
pelvics, dorsals, anal and caudal ; the embryo is described as slate gray.^

Size. This is one of the larger deep-water sharks. Specimens so far measured have
ranged from 930 mm. to 2,950 mm. (9 feet 8 inches) in length. The length at which it
matures is not known.

4. For recent illustrations of the teeth of a large specimen, see Jacquet (Bull. Inst, oceanogr. Monaco, 36, 1905:
pi. 8).

5. Saemundsson (Vidensk. Medd. naturh. Foren. Kbh., y^, 1922: 197).

232 Memoir Sears Foundation for Marine Research

Developmental Stages. Knowledge of its development is limited to the fact that a
gravid female of nine feet contained two embryos, each about 850 mm. long.

Habits. The fact that most of the captures of specimens for which pertinent informa-
tion is available have been made at depths ranging between 300 and 1,477 rneters (164
and 807 fathoms) shows this to be a deep-water species, a habitat with which its uniformly
dark coloration accords. But the Long Island specimen mentioned below was washed
ashore on the beach and the New Jersey example was taken in a pound net, which is evi-
dence that it occasionally wanders into shallow water, as various other deep-water fishes do.
Nothing more is known of its habits.

Range. Both sides of the North Atlantic} also represented in Japanese waters by a
closely allied form (acrages Jordan and Snyder, 1904). Apparently this Shark is rare
everywhere, for positively identified specimens so far captured number only nine; these
are the type specimen and two others from the coast of Portugal; one from the Cape Verde
Islands; three from Iceland; one from an unspecified Atlantic locality; and one from
Amagansett, Long Island, New York. It is also reported nominally from Madeira, and it is
probable that an eight-foot shark taken in a pound net at Manasquam, New Jersey, in July
1936 also belonged to this species. All the numerous references to it in scientific literature
are based on the foregoing.

Synonyms and References:

Pseudotriakis microdon Brito Capello, J. Sci. Math. Phys. nat. Lisboa, / (2), 1867: 315, 321, pi. 5, fig. I (type
loc. Setubal, Portugal); ibid., 2, 1870: 139; Giinthcr, Cat. Fish. Brit. Mus., 8, 1870: 395 (Portugal);
Brito Capello, Cat. Peix. Portugal, 1880:44 (Portugal); Bean, T. H., Proc. U.S. nat. Mus., 6, 1883: 147
(Long Island, N. York); Baird, Bull. U.S. Fish Comm., 4, 1884: 1 77 (same specimen as Bean, 1883);
Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 794 (dlstrib.) ; Goode and Bean, Smithson. Contr. Knowl.,
50, 1895: 18, 508, pi. 5, fig. I 7; Mem. Harv. Mus. comp. Zool., 22, 1 896 (same ref.) ; Special Bull. U.S.
nat. Mus., 2, same pagination (descr., ill. of Long Island specimen, meas.) ; Jordan and Evermann, Rep.
U.S. Comm. Fish. (1895), 1896: 214 (distrib.) ; Bull. U.S. nat. Mus., 47 (l), 1896: 27; Bull. U.S.
nat. Mus., 47 (4), 1900: pi. 4, fig. 14 (dcscr., distrib.) ; Bean, T. H., Bull. N. Y. St. Mus., 60, Zool. 9,
1903: 18 (Long Island spec.) ; de Braganza, Result. Invest. Sci. "Amelia," 2, 1904: 28, 29 (off Portugal,
depths); Prince of Monaco, Bull. Inst, oceanogr. Monaco, 6, 1904: II (depth); Jacquet, Bull. Inst.
oceanogr. Monaco, 36, 1905: I, pi. 1-8 (anat., teeth, depth, C. Verdes) ; Regan, Ann. Mag. nat. Hist.,
(8) /, 1908: 464 (descr.j Atlant.) ; Richard, Bull. Inst, oceanogr. Monaco, 162, 1910: 152, fig. 109
(photo, C. Verdes); Roule, Bull. Inst, oceanogr. Monaco, 243, 191 2: 6 (C. Verdes); Carman, Mem.
Harv. Mus. comp. Zool., j(5, 191 3: 104 (dcscr.) ; Nichols and Murphy, Brooklyn Mus. Sci. Bull., (l) j,
1916: 6 (Long Island record); Fowler, Copeia, 30, 1916: 36; Roule, Result. Camp. sci. Monaco, 52,
1919: 114 (C. Verdes); Saemundsson, Vidensk. Medd. naturh. Foren. Kbh., 14, 1922: 197 (meas. em-
bryos, size, depth, Iceland) ; Noronha, Ann. Carneg. Mus., 16, \ 926: 385 (Madeira) ; Nichols and Breder,
Zoologica, N. Y., 9, 1927: II (the Long Island record) ; Rey, Fauna Iberica Feces, i, 1928: 320 (descr.,
distrib.) ; Breder, Field Bk. Mar. Fishes Atlant. Coast, 1 929: 12 (general) ; Jordan, Manual Vert. Anim.
NE. U.S., 1929: 8 (general); Fowler, Proc. Pan-Pacif. sci. Congr. (1929), Java, 1930: 489 (part);
Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 12 (distrib.); Frade, Cons. Ex-
plor. Mer., 1932: plate not numbered (general) ; Daniel, Elasmobranch Fishes, Univ. Calif. Press, 1934:
54, fig. 59 (skull); Nobre, Fauna Marinha Port. Vert., r, 1935: 442 (descr., Portugal); Fowler, Bull.
Amer. Mus. nat. Hist., yo (i), 1936: 42 (general); Proc. Acad. nat. Sci. Philad., 8g, ig^?'- 3°3' 3^4
(specimen in pound net, Manasquam, N. Jersey, July 1936, ident. probable, size) ; White, Bull. Amer.

Fishes of the Western North Atlantic 233

Mus. nat. Hist., 7^, 1937: 41, 119 (class.) ; Fowler, Bull. U.S. nat. Mus., 100 {13), 1941 : 1 18 (part,
class.).

Family TRIAKIDAE
Smooth Dogfishes

Characters. Two dorsal fins, the ist much shorter than the caudal, its base terminating
anterior to origin of pelvics; caudal less than ^4 of total length, not lunate, its lower ante-
rior corner expanded as a low lobe only, if at allj caudal peduncle not flattened dorso-
ventrally or expanded laterally, with or without a precaudal pit above, but none below j
sides of trunk, anterior to anal, without longitudinal ridges; inner margins of pelvics en-
tirely separate posterior to cloaca in both sexes; snout not greatly elongate, or jaws widely
protrusible; 5th gill opening posterior to origin of pectoral; gill arches without rakers and
not interconnected by a sieve of modified denticles; nostril either separate from mouth or
connected with latter by a groove, its anterior margin without barbel; spiracles present
or absent; lower eyelid with a longitudinal fold externally, but without internal nictitating
membrane; jaws with or without labial furrows; teeth small, rounded, or with 3 to 4
distinct cusps; several series functional along entire length of jaw; dermal denticles ovate,
lanceolate or tridentate; head of normal shape and not widely expanded laterally. Rostral
cartilages 3, united terminally; most of the radials of pectoral on metapterygium ; meso-
and propterygia much smaller; meso- and metapterygia separated by a foramen, at least in
some cases; heart valves in 3 rows. Development either ovoviviparous, or viviparous with
a well developed yolk-sac placenta.

Genera. These small sharks, of shoal or moderate depths, are widely distributed in
tropical and warm temperate belts of all oceans. They are perfectly harmless. The group
is very close to the Carcharhinidae where it is placed as a subfamily by Fowler,^ although
it more resembles the Orectolobidae and the Scyliorhinidae in its dentition. However,
the triakids seem sufiiciently different both from the typical carcharhinids (with respect
to the teeth and the absence of a true nictitating membrane), as well as from the orectolo-
bids and scyliorhinids, to be ranked as a distinct family for convenience, if for no better
reason.

The family, as defined above, corresponds to the Galeorhinidae as used by Garman
and by many subsequent authors. But the latter name is not appropriate in the present
connection, because Galeorhinus Blainville, 1 8 1 6, from which it is derived, is the correct
name of the so-called "Topes," a genus of carcharhinid sharks (see footnote 4, p. 264),
and not of the Smooth Dogfishes (Muslelus) to which Garman" and others have ap-
plied it. For this reason Bigelow and Schroeder^ substituted Mustelidae for the sharks
now in question, overlooking the fact that this name had been in common use for a family

I. Bull. U.S. nat. Mus., 100 (75), 1941 : i 27. 2. Mem. Harv. Mus. comp. Zool., 3d, 191 3 : 3, 169.

J. Canad. Atlant. Fauna, biol. Bd. Canad., 12^, 1934: 6.

2 34 Memoir Sears Foundation for Marine Research

of carnivorous mammals (weasels, etc.) for many years previous. To avoid the confusion
that would result from the use of this family name for sharks, as well as for mammals, it
seems wiser to replace it with "Triakidae" White,^" first used by that author in a more
restricted sense, but subsequently expanded by her.^"

Key to Genera
la. Nostril connected with mouth by a groove. Scylliogaleus Boulenger, 1902.

Natal, southeastern Africa.

lb. Nostril entirely separate from mouth.

2a. Teeth low, rounded, or with only slightly sinuous cutting edge.

Mustelus Link, 1790, p. 240.
2b. Teeth somewhat compressed, with 3 to 5 pointed cusps.

3a. No labial furrow at corner of mouth. Eridacnis H. M. Smith, 191 3.

Philippines.

3b. A labial furrow at corner of mouth.

4a. Anterior margin of nostril expanded as a well developed and conspicuous
barbel. Fwr Whitley, 1943.

Australia.''

4b. Anterior margin of nostril not expanded as a well developed barbel.
5a. Spiracle moderately large, easily seen.

Triakis Miiller and Henle, 1838, p. 235.'
5b. Spiracle minute or absent.

6a. Caudal peduncle with precaudal pit above; lower anterior cor-
ner of caudal expanded as a lobe.

Triaenodon Miiller and Henle, 1837.

Tropical Indian Ocean, Red and Arabian Seai;
India and Ceylon, Malaysia, Melanesia and
Polynesia, Hawaiian Islands, Cocos Island, and
Panama.
6b. Caudal peduncle without precaudal pit; lower anterior corner
of caudal not expanded as a lobe.

Leptocharias Miiller and Henle, 1838.
West Africa.

Remarks. The separation between the only two triakid genera that are known to occur
in the Atlantic, Triakis and Mustelus, is partly bridged by T. macidata of the west coast
of South America, the teeth of which are only a little more definitely cuspidate than those
of Mustelus dorsalis of the same geographic province."

3a. Amer. Mus. Novit., 879, 1936: 19. 3b. Bull. Anier. Mus. nat. Hist., 74, 1937: 121.

4. Whitley, Aust. Zool., 10 (2), 1943: 167; Rec. Aust. Mus., 7 (4), 1943: 397 > Aust. ZooL, to, 194+: 260, pi. 3.

5. Including R/iinotriacis Gill, 1862, Calliscylliutn Tanaka, 1912, and Hemhriakis Herre, 1923, all of which seem
clearly referable to the old genus Triakis (see p. 235).

6. For discussion, see Bigelow and Schroeder (Proc. Boston Soc. nat. Hist., 41 [8], 1940: 430-

Fishes of the Western North Atlantic 235

Genus Triakis Miiller and Henle, 1838

Triakts Miiller and Henle, Charlesworth's Mag. Nat. Hist., 2, 1838: 36; Arch. Naturg., Jahrg. 4, t, 1838: 84
(no species mentioned) ; Plagiost., 1841: 63 ; type species, T. scyllium Miiller and Henle. Japan.

Generic Synonyms:

Triads Giinther, Cat. Fish. Brit. Mus., 8, 1870: 384; emended spelling for Triakis Miiller and Henle, 1838.

Rhinotriacis G'lW, Proc. Acad. nat. Sci. Philad., 1862: 486; type species, R. henlei Gill. California.

Calliscyllium Tanaka, Fish. Japan, 10, 1912: 17; type species, C. venusta Tanaka. Japan.

Hemitriakis Herre, Philipp. J. Sci., 23, 1923: 70; type species, H. leucoferiftera Herre. Philippines.

Generic Characters. Nostrils entirely distinct from mouth, their anterior margins
without barbel} teeth compressed, with 2, 3 or 4 pointed cusps, the axial longest; a labial
furrow on each jaw at corner of mouth; spiracles moderately large, easily seen. Characters
otherwise those of the family.

Range. Until recently the genus was known only from the Indian Ocean (including
the Red and Arabian Seas), Malaysia, Melanesia and Polynesia, the western North Pa-
cific, and the eastern Pacific, north and south. Within the last few years it has been en-
countered in Cuban waters (p. 240).

Species. The members of this genus show a wide gradation in the number of dental
cusps and in. the relative length and acuteness of the median member;' the dermal den-
ticles, too, may be either simple-lanceolate, partly or weakly tri dentate, or strongly so;*
and the labial furrows are very prominent in some but very short and inconspicuous in
others.' The several species also differ in the relative positions of the dorsal and anal fins,
the size and shape of the caudal, and in the proximity of nostril to mouth, these being the
characters employed in the accompanying Key because they are the most conspicuous.

Key to Species

I a. Origin of 2nd dorsal about over origin of anal or behind it, its tip over or posterior to
tip of latter; labial furrows very short.

2a. Anterior margin of nostril close to mouth; rear tip of ist dorsal a little anterior
to origin of pelvics; tips of pelvics anterior to origin of 2nd dorsal by a distance
considerably longer than base of 2nd dorsal. venusta Tanaka, 1 9 1 2.

Japan.

2b. Anterior margin of nostril far from mouth; rear tip of ist dorsal posterior to
origin of pelvics; tips of pelvics anterior to origin of 2nd dorsal by a distance only
about y-i as long as base of 2nd dorsal.

barbouri Bigelow and Schroeder, 1943, p. 236.

7. Median cusp long and slender in scyllia, semifasciata and barbouri; only moderately so in venusta; short and
blunter in maculata and henlei.

8. Denticles simple-lanceolate in scyllia, semifasciata and /tenlei; weakly tridentate in venusta; partly so in maculata;
strongly tridentate in barbouri.

9. Labial furrows long and conspicuous in scyllia, semifasciata, henlei and maculata; very short in barbouri and
venusta.

236 Memoir Sears Foundation for Marine Research

lb. Origin of 2nd dorsal considerably anterior to origin of anal, its tip terminating anterior
to tip of latter; labial furrows long, conspicuous.
3a. Rear end of base of ist dorsal about over origin of pelvlcs.

maculata Kner and Steindachner, 1867.'°
Peru.
3b. Rear end of base of ist dorsal so far in advance of origin of pelvics that its tip is
over origin of latter.

4a. Lower anterior corner of caudal expanded as triangular lobe, with pointed
tip J pectoral also pointed, with deeply concave distal margin.

leucoperiptera Herre, 1923.
Philippines.

4b. Lower anterior corner of caudal rounded and only slightly expanded; pec-
toral rounded at tip, its distal margin only weakly concave, or nearly straight.
5a. Origin of ist dorsal anterior to inner corner of pectoral, its tip anterior
to origin of pelvics; plain colored. henlei Gill, 1862.

California.

5b. Origin of ist dorsal over or behind inner corner of pectoral, its tip about
over origin of pelvics; back and sides with dark bars, blotches or spots.
6a. Caudal about as long as from tip of snout to axil of pectoral; teeth
in sides of jaws similar to those in front, tricuspidate, the median
cusp straight although directed somewhat obliquely outward.

jcy///«"'''MullerandHenle, 1841.
Japan, Korea, Formosa, China. ^^
6b. Caudal shorter than from tip of snout to axil of pectoral by a dis-
tance about equal to that between nostrils; teeth in sides of jaws
noticeably unlike those in front, usually with the inner lateral cusp
lacking, and the median cusp strongly curved outward.

semifasciata Girard, 1854.

Oregon to Magdalena Bay, Lower
California.

Triakis barbouri Bigelow and Schroeder, 1944

Figure 4 1

Study Material. About lOO specimens, males and females, 225 to 338 mm. long,
including the type, collected off the north coast of Cuba in March 1938 and May 1939
by the research ship "Atlantis" (Harv. Mus. Comp. ZooL),

10. Including nigromactilata Evermann and Radcliffe, 1917 ; nigrotnacu'.ata has sometimes been referred to Mustelus,
but our own examination of Peruvian specimens inclines us to follow Garman (Mem. Harv. Mus. comp. ZooL,
j6, 1913 : 167) in retaining' it in Triakis where it was placed by its discoverers, its teeth being definitely cuspi-
date, though usually with only two cusps, rarely three (see Bigelow and Schroeder, Proc. Boston Soc. nat. Hist.,
41 [8], 1940: 42S, pi. 17, fi;:. M).

loa. The original spelling, scylliiim, is emended here to accord with the gender of the Greek noun (aKic) on
which the generic name Triakis was based.

11. Old reports of this species from South Australia probably were not correct; see Whitley (Fish. Aust., /, 1940:

• ■s).

Fishes of the Western North Atlantic

237

Distinctive Characters. The very short labial furrows of barbouri are enough to sepa-
rate it at a glance from all other members of the genus, except from venusta, from which
it differs in its strongly tridentate dermal denticles, in the fact that the anterior marginal
expansion of its nostril is far in advance of the front of the mouth, and in its teeth. It is not
likely to be confused with any other Atlantic shark, being the only member of its genus yet
known from this ocean.

Figure 41. Triakis barbouri, adult male, 283 mm. long, from north coast of Cuba (Harv. Mus. Comp. Zool.,
No. 36099, type). A Anterior part of head from below. B Pelvic fins with claspers, about I x. C Upper teeth
from midsection of jaw, about 15 x. Z) Enlarged tooth from C, about 30 x. E Lower teeth near center of
mouth. F Lower teeth near corner of mouth. G Four outermost series of lower teeth. E-G, about I 5 x. // Der-
mal denticles, about 25 x.

Description. Proportional dimensions in per cent of total length. Male, 283 mm.,
from Cuba (Harv. Mus. Comp. Zool., type. No. 36099). Female, 298 mm., from Cuba
(Harv. Mus. Comp. Zool., No. 36151).

Trunk at origin of pectoral: breadth 9.5, 9.55 height 8.8, 8.4.

Snout length in front of: outer nostrils 3.5, 3.5; mouth S-^^ 5-4-

Eye: horizontal diameter 3.9, 4.3.

Mouth: breadth 7.1, 7.2; height 3.5, 3.3.

Nostrils: distance between inner ends 2.9, 2.9.

Labial furrow lengths: upper 0.5, 0.6; lower 0.5, 0.7.

238 Memoir Sears Foundation for Marifie Research

Gill opening lengths: ist 1.8, 1.7; 2nd 1.8, 1.8; 3rd 1.8, 1.8; 4th 1.8, 1.7; 5th

1.4, 1.2.

First dorsal fin: vertical height 4.9, 5.4; length of base 6.9, 7.4.

Second dorsal fin: vertical height 4.2, 5.2; length of base 8.8, 8.7.

Anal fin: vertical height 2.9, 3.5; length of base 8.1, 9.1.

Caudal fin: upper margin 27.O, 27.8; lower anterior margin 9.9, 8.4.

Pectoral fin: outer margin 13.0, 13.8} inner margin 8.1, 8.5; distal margin 9.5,

10.7.

Distance from snout to: ist dorsal 31.4, 31.2; 2nd dorsal S^-S-> 5^-4-y upper caudal

73.0, 72.2; pectoral 20.4, 19.45 pelvics 39.2, 42.6; anal 57.8, 56.7.

Interspace between: ist and 2nd dorsals 16.9, 17. i; 2nd dorsal and caudal 7.6,

7.7; anal and caudal 5.0, 4.7.

Distance from origin to origin of: pectoral and pelvics 22.2, 24.35 pelvics and

anal 16.9, 14.6.

Trunk slender, its height abreast of origin of ist dorsal only about •'/'r as great as its
length to origin of caudal, its dorsal profile only slightly arched, its ventral profile nearly
straight except in gravid females. Tail sector about '^/q longer than body sector to cloaca.
Caudal peduncle laterally compressed, about twice as high as broad at origin of caudal,
without lateral keels or precaudal pit above or below. Dermal denticles close-spaced and
usually overlapping, with short pedicels and horizontal blades, the latter regularly tri-
dentate, the median tooth narrow and much the longest 5 3 ridges, the median ridge flat-
topped distally but rounded proximally.

Head to 5th gill opening about Vs of total length, flattened above, its dorsal profile
sloping steeply downward. Snout in front of mouth about Vs of length of head to origin
of pectoral, or a little less, its tip thin and broadly ovate. Eye oval, about 3 times as long
as high, and noticeably large, its horizontal diameter about % as long as snout in front of
mouth, the larger specimens with a well marked subocular fold which is separated from
the margin of the lower eyelid at both ends when the latter is drawn down, but this Is
only faintly indicated when the lower eyelid is drawn up over eyeball, or in small speci-
mens even when the eye is wide open. Spiracle oval, its longest diameter only about V^^ to
Ve as long as horizontal diameter of eye, situated slightly below level of center of latter
and behind it by a distance slightly less than V2 an eye's length. Gill openings deeply con-
cave anteriorly, the 4th to 5th over the origin of the pectoral, the ist, 2nd, 3rd, and 4th
about V2 as long as the horizontal diameter of the eye, the 5 th considerably shorter and
situated noticeably higher on the sides than the ist to 4th 5 the interspaces between gill
openings about V2 to % as wide as lengths of latter. Nostril moderately oblique, its inner
corner a little less than V3 as far from front of mouth as from tip of snout, its anterior mar-
gin expanded as a low, subquadrate lobe. Mouth broadly ovate, occupying about % of
breadth of head, and a little less than V2 as high as broad, with a very short labial furrow
on each jaw.

Fishes of the Western North Atlantic 239

Teeth "*°|| , closely crowded, with those of successive rows overlapping; 3 or 4
rows functional in front of upper jaw, but only 2 or 3 rows near corners; 3 to 4 rows
functional in front of lower jaw, and 5 to 6 rows toward the angles where the serial ar-
rangement is increasingly oblique. Upper teeth with 3 cusps, the axial erect or slightly
curved and much the longest, with irregular longitudinal striae basally, those toward
corners of mouth somewhat smaller, with relatively shorter median cusp and broader
base than those in front of mouth; lowers similar to uppers toward center of mouth, except
that the outer lateral cusp may be minute or lacking, the teeth along sides of jaw usually
with 2 or 3 basal cusps on the inner side, but none on the outer side, so that the cusp that
is primarily median is at the outer edge of the tooth.

Origin of ist dorsal a little behind inner corner of pectoral, its base about as long as
from center of eye to ist gill opening, its anterior margin very slightly convex, its apex
rounded, its posterior margin slightly concave, the free rear corner a little more than V2
as long as its base, the rear end of latter a little anterior to origin of pelvics. Second dorsal
similar to ist, and about as high, but about i V2 times as long at base, its origin about over
that of anal and slightly closer to origin of caudal than to rear tip of ist dorsal. Caudal
Vi to Yz of total length, noticeably narrow, obliquely truncate at tip, with well marked
subterminal notch, its rounded, lower anterior corner not expanded to form a distinct lobe.
Anal similar to 2nd dorsal in shape, and about as long at base, but only about % as high.
Interspace between anal and caudal a little shorter than base of anal, that between anal
and tip? of pelvics a little more than V2 as long as base of anal. Pelvics with nearly straight
anterior and distal margins but slightly concave inner margins, and broadly rounded outer
corners, the inner corners narrowly elongate in both sexes, with pointed tips. Pectoral be-
tween 2 and 3 times as large in area as ist dorsal, about % as broad as long, with weakly
convex outer and inner margins, very weakly concave distal margin and narrowly rounded
corners.

Color. Upper parts of trunk, caudal, dorsals and pectorals pale gray; lower surface
grayish white. At least in some specimens the anterior edge of the dorsals, as well as that
of the caudal midway of its length, is marked with faintly defined smoky blotches. The
embryo just before birth shows more extensive blotches in these same general regions,
in addition to similar blotches on the upper side of the caudal peduncle, on the back mid-
way between the two dorsals, and on the side just posterior to the pectoral; also, a sooty
area, irregular in outline, extending upward obliquely from the region of the gill open-
ings to the dorsal surface of the nape.

Size, Evidently this is one of the smallest of sharks. The fact that a female of only
303 mm. contained two large embryos, and that the claspers in a 338-mm. male are very
large and extend rearward as far as the origin of the anal, suggests a maximum length of
perhaps not more than about 350 to 400 mm., or about one foot two to four inches.

Developmental Stages. Two embryos, 90 to 100 mm. long, and nearly ready for
birth, contained in a female of 303 mm., resemble the parent, except for the facts that the

240 Memoir Sears Foundation for Marine Research

caudal Is relatively longer with slightly convex lower margin, that the origin of the ist
dorsal is relatively somewhat farther back, that the dorsals are more broadly rounded,
that the tips of the pelvics are relatively much shorter, and that the eyes are relatively
much larger (embryonic characters to be expected) ; the color pattern is not only more
pronounced but more extensive, as noted above. There is no placental attachment between
embryo and mother.

Habits. The recorded depths of capture of this little Shark range only between 235
and 335 fathoms, but since the trawls that yielded them were hauled up to the surface
open, the specimens may have been swimming at some mid-depth and not on bottom.
Nothing more is known of its habits.

Range. So far known only oflF the north coast of Cuba in the offing of the province
of Santa Clara, where it must be tolerably plentiful, judging from the number of speci-
mens taken.

Synonym and References:

Triakis barbouri Bigelow and Schroeder, Proc. New Engl. zool. CI., 23, 1944: 27, pi. 8 (descr., ill., Cuba) ;

Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 124

(descr., ill., range).

Genus Mustelus Link, 1790^
Smooth Dogfishes

Mustelus Link, Mag. Physik Naturg. Gotha, 6 (2), 1790: 31 ; type species, Squalus musttlus Linnaeus, 1 758.

Generic Synonyms:

Galeus (in part) Rafinesque, Indice Ittiol. Sicil., 1810: 46 (includes Squalus mustelus Linnaeus, 1758) ; Leach,
Mem. Werner. Soc. Edinb., 2, 1812: 62 (includes Squalus mustelus Linnaeus, 1758); not Galeus Rafi-
nesque, Carratt. Gen. Nuov. Sicil., 1 810: 13; see p. 214.

Galeorhinus (in part) Blainville, Bull. Soc. philom. Paris, 1816: 121.

Myrmillo Gistel, Naturg. Tier., 1 848: 4; substitution for Mustelus Link, 1 790; not seen.

Pleuracromylon Gill, Proc. Acad. nat. Sci. Philad., 1864: 148; type species, Mustelus laevis Risso, 1826.

Rhinotriakis (in part) Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 60; includes Mustelus laevis Risso,
1826.

Galeus Jordan and Evermann, 1896, Bull. U.S. nat. Mus., 47 (l), 1896: 29; for Mustelus dorsalis and M.
californicus Gill, 1864; not Galeus Rafinesque, Carratt. Gen. Nuov. Sicil., 1810: 13.

Cynias Gill, Proc. U.S. nat. Mus., 26, 1903: 960; type species, Squalus cants Mitchill, 1815.

Cynais Fowler, Proc. Acad. nat. Sci. Philad., 60, 1 908: 55 ; evident misprint for Cynias.

Galeorhinus Giiman, Mem. Harv. Mus. comp. Zool., 36, 1 91 3: 169; same as Mustelus Link, 1 790, substituted
because of the diversity of application of the latter; not Galeorhinus Blainville, 18 16, the type species of
which is Squalus galeus Linnaeus, 1 75 8, desig. by Gill, Ann. N. Y. Lye, 7, 1862: 402.

1. The International Commission on Zoological Nomenclature has ruled that Mustelus Link, 1790, is applicable
to sharks, in spite of the fact that Mustela was earlier used by Linnaeus, 1758, for weasels. See Opinion 93
(Smithson. misc. Coll., 75 [4], 1929: 8; and Science, N. S. 6$, 1927: 300).

Fishes of the Western North Atlantic 241

Generic Characters. Nostrils entirely separate from mouth j teeth numerous, alike in
the 2 jaws, low, rounded, or with somewhat sinuous cutting edge, in mosaic arrangement,
several rows functional; spiracles present, of at least moderate size; corners of mouth with
a strongly marked labial furrow on each jaw; caudal with well marked subterminal notch,
its lower anterior corner expanded as a short lobe in some species but not in others; dorsals,
anal and pelvics with their posterior corners considerably produced and their posterior
margins moderately concave; origin of anal below 2nd dorsal; rear end of base of ist
dorsal considerably anterior to origin of pelvics; in some species there is a placental attach-
ment between mother and embryo, but in others this is lacking. Characters otherwise those
of the family.

Remarks. The members of the genus are separated from others of this family by their
low rounded teeth, arranged in mosaic.

Range. Widely distributed in coastal waters in the warm and warm-temperate belts
of the Atlantic and Indo-Pacific, north and south; Gulf of Maine to northern Argentina
in the western Atlantic; Shetland Islands, North Sea and mouth of the Baltic to tropical
West Africa (Senegambia, Cameroons) in the eastern Atlantic, including the Mediter-
ranean, Madeira and the Canaries; also South Africa; western Indian Ocean (Natal);
Red and Arabian Seas; Australia, Tasmania and New Zealand; Indo-China, China, Korea
and Japan; west coasts of North, Central, and South America from northern California
to northern Chile.

Fossil Teeth. Oligocene to Pliocene, Europe.

Species. Although the members of this genus resemble one another very closely in
general appearance, certainly two, and probably three, recognizably distinct species occur
in the eastern Atlantic, four in the western Atlantic, four along the western coasts of
America; also at least two or three and perhaps more in the western Pacific, in the Aus-
tralian-New Zealand region and in the Indian Ocean with its tributary seas. We have
already published a comparison of the western Atlantic species with those of the eastern
Atlantic and eastern Pacific.^ The next step would be to compare these with their western
Pacific-Australian-Indian Ocean relatives. However, since we lack sufficient material for
this, the following key is limited to the Atlantic and to the eastern side of the Pacific.

Key to Atlantic and Eastern Pacific Species

la. Lower anterior corner of caudal expanded as a pointed lobe directed rearward (Fig.
43 D).

2a. Upper labial furrow as long as lower, or longer; denticles on shoulders loosely
spaced. norrisi Springer, 1939, p. 254.

2b. Upper labial fold shorter than lower; denticles on shoulders closely imbricate.

lunulatus Jordan and Gilbert, 1882.

Southern California to Colombia.
3. Bigelow and Schroeder, Bull. Boston Soc. nat. Hist., ^i (8), 1940: 417.

242 Memoir Seats Foundation for Marine Research

lb. Lower anterior corner of caudal broadly rounded.

3a. Midpoint of base of ist dorsal about midway between origin of pelvics and inner
corner of pectoral when latter is laid back.

4a. Teeth noticeably asymmetrical, their outer margins deeply concave, their tips
moderately pointed j dermal denticles strongly ridged, usually to their tips.

puncfulatus ^isso, 1826.

Mediterranean.

4b. Teeth nearly symmetrical, their outer margins only weakly concave, their
tips broadly rounded; dermal denticles ridged only at their bases, their tips
smooth, transparent. calif amicus Gill, 1864.

Northern California to Cape
San Lucas and Gulf of Cali-
fornia.

3b. Midpoint of base of ist dorsal much closer to inner corner of pectoral when this
is laid back than to origin of pelvics.
5a. Teeth symmetrical, evenly rounded in both jaws.

6a. Lower anterior corner of caudal not expanded as a lobe; snout in front
of mouth about 4 times as long as horizontal diameter of eye.

fasciaius Ga.rmzn, 19 13.
Southern Brazil, Uruguay,
northern Argentina.

6b. Lower anterior corner of caudal expanded as a low, rounded lobe ; snout
in front of mouth only 2 to 3 times as long as horizontal diameter of eye.
7a. Horizontal diameter of eye considerably longer than distance be-
tween nostrils, and about ^^ as long as snout in front of mouth;
denticles on shoulders strongly ridged, usually to their tips, and
opaque. asterias Cloquet, 1 8 1 9.

Eastern Atlantic, Mediter-
ranean.

7b. Horizontal diameter of eye slightly shorter than distance between

nostrils, and only Vs as long as snout in front of mouth ; denticles

on shoulders ridged only near their bases and so transparent that the

pedicels are visible from without, mento* Cope, 1877, p. 259.

5b. Teeth noticeably asymmetrical, their margins more or less concave.

8a. Sides in adults conspicuously marked with white spots.

asterias Cloquet, 1 8 1 9.

Eastern Atlantic, Mediter-
ranean.

8b. Sides in adults plain colored.

9a. Lower anterior corner of caudal not appreciably expanded even in
adult. schmitti Springer, 1938.

Southern Brazil, Uruguay and
northern Argentina.
4. Including ahbotti Evermann and RadcliflFe, 1917, and eiul't! Perez Canto, 1886.

Fishes of the Western North Atlantic 243

9b. Lower anterior corner of caudal noticeably expanded as a rounded
lobe from shortly after birth.

lOa. Teeth high-crowned, their outer margins deeply concave even
in adults, and sometimes the inner as well.

dorsalis Gill, 1864.

Pacific Panama, Peru.

lob. Teeth low-crowned, their tips bluntly rounded, their margins
only slightly concave in adults.

iia. Inner margin of pectoral 60 to 63% as long as outerj
distance between nostrils only about 1.3 times as long as
upper labial furrow j dermal denticles on shoulders
ridged only near their bases, their tips smooth.

mustelus Linnaeus, 1758.
Eastern Atlantic, Mediterra-
nean.
lib. Inner margin of pectoral only about 48 to 54% as long
as outer 5 distance between nostrils at least 1.3 to 1.7
times as long as upper labial furrow 5 denticles usually
strongly ridged to their tips.

cams Mitchill, 1815, p. 244.

Key to the Western Atlantic Species"

I a. Head to origin of pectoral considerably longer than interspace between ist and 2nd

dorsals. fasciatus GzTma.n, I9i3,p. 256.

lb. Head to origin of pectoral not longer than interspace between ist and 2nd dorsals.

2a. Lower anterior corner of caudal sharp-pointed and directed rearward (Fig.

43 D). norrisi Springer, 1939, p. 254.

2b. Lower anterior corner of caudal broadly rounded.

3a. Horizontal diameter of eye only about 60% as long as 3rd gill openings
teeth symmetrical, evenly convex in outline (Fig. 42 J).

mento Cope, 1877, p. 259.
3b. Horizontal diameter of eye about as long as 3rd gill opening; teeth some-
what asymmetrical, their cutting edges more or less concave.
4a. Lower anterior corner of caudal not appreciably expanded, even in adult
(Fig. 42 F) ; distance between nostrils only about '^/z as great as breadth
of mouth. schmitti Springer, 1939, p. 261.

4b. Lower anterior corner of caudal noticeably expanded as a rounded lobe
from shortly after birth (Fig. 42); distance between nostrils consider-
ably more than V2 as great as breadth of mouth.

canis Mitchill, 18 15, p. 244.

5. This simplified Key is offered for assistance in the identification of species from the American Atlantic coast.

244

Memoir Sears Foundation for Marine Research

Mustelus cams (Mitchill) ,1815

Smooth Dogfish

Figures 42 A-E, 43 C

Study Material. Numerous specimens from southern New England and New York,
newborn to adult, living, fresh-caught, and preserved} also preserved specimens from
South Carolina j Galveston, Texas; Havana, Cuba; Jamaica; Trinidad (embryo); Ber-
muda; and Brazil, including three embryos from Rio de Janeiro.

Distinctive Characters. M. canis is very closely allied to M. mustelus of the eastern
Atlantic, but is separable from the latter by the narrower pectoral fin, relatively greater
distance between the nostrils and the much more strongly sculptured denticles. Among
western Atlantic species of the genus it is distinguished from schmitti in that the lower
anterior corner of its caudal forms a lobe and that the distance between its nostrils is rela-
tively greater, that is, approximately half as great as the breadth of the mouth (only
about a third as great in schmitti) ; from norrisi in that its lower caudal lobe is rounded
rather than pointed and that the midpoint of its first dorsal is as close to the axil of the pec-

FicuRE 42. Mustelus canis, male, 78 1 mm. long, from Woods Hole, Massachusetts (Harv. Mus. Comp. Zool.,
No. 35245). A Head to pectorals. B Cross section of trunk, midway between the two dorsal fins, to show mid-
dorsal ridge. C Dentition of right-hand side of upper jaw, about 2 x. D Teeth of another specimen of about
the same size, about 7 x. E Dermal denticles of a female, about 678 mm. long, from Woods Hole, Massachu-
setts, about 22 X. F Caudal fin of Mustelus schmitti, 578 mm. long, from Muldonado, Uruguay (Harv. Mus.
Comp. Zool., No. 530), for comparison with M. canis. G Upper teeth of same, about j x. H Caudal fin of
M. tnento, 1,024 mm., from Payta, Peru. / Denticles of same, about 22 x. J Upper teeth of same, about 7 x.

Fishes of the Wester ti North Atlantic 245

toral as to the origin of the pelvics; from fasciatus by the shape of the teeth (cf. Fig. 42 D
with 43 A), by its lobed lower caudal, and by its plain coloration 5 from mento^ by a much
larger eye relative to the gill openings (the horizontal diameter of its eye being about as
long as the third gill opening but only about 60% as long in mento), by the distance be-
tween its nostrils which is at least approximately half as great as the width of the mouth
(considerably less than Y^ that great in mento), by a relatively smaller pectoral with inner
margin only about 1.3 times as long as snout (1.8 times as long in mento), by its more
strongly sculptured and more opaque denticles (cf. Fig. 42 E with 42 I) and by the shapes
of its teeth (cf. Fig. 42 D with 42 J). Adults of canis are plain colored also, whereas those
of mento often are white-spotted (p. 260).

Descriftion. Proportional dimensions in per cent of total length. Male, 781 mm.,
from Buzzards Bay, Mass. (Harv. Mus. Comp. Zool., No. 35245). Female, 1,231 mm.,
same locality.

Trunk at origin of -pectoral: breadth lO.O, 10.35 height 9.9, lO.i.

Snout length in front of: outer nostrils 3.8, 3.2; mouth 6.4, 5.4.

Eye: horizontal diameter 2.8, 2.3.

Mouth: breadth 5.5, S-5'i height 2.7, 2.3.

Nostrils: distance between inner ends 2.9, 2.6.

Labial furrow length: up-per 2.0, 1.9; lower 1.5, 1.4.

Gill opening lengths: ist 2.2, 2.75 2nd 2.4, 2.8; 3rd 2.6, 3.2; 4th 2.6, 3.2; 5th

2.0, 2.4.

First dorsal fin: venkzi height 9.5, 8.3; length of base ii.i, 12.3.

Second dorsal fin: vertical height 6.8, 6.7; length of base 8.9, 10.6.

Anal fin: vertical height 3.6, 4.2; length of base 5.4, 5.5.

Caudal fin: upper margin 21.2, 18.8; lower anterior margin 8.7, 7.0.

Pectoral fin: outer margin 13.6, 13.9; inner margin 7.4, 6.7; distal margin 12.4,

12.5.

Distance from snout to: ist dorsal 26.8, 28.6; 2nd dorsal 59.7, 61.7; upper caudal

78.8, 81. 2j pectoral 18. i, 19.1; pelvics 41.5, 47.6; anal 64.3, 68.0.

Interspace between: ist and 2nd dorsals 22.0, 22.0; 2nd dorsal and caudal 10.5,

9.5 ; anal and caudal 6.8, 6.7.

Distance frotn origin to origin of: pectoral and pelvics 24.3, 29.2; pelvics and

anal 22.4, 19.8.

Trunk slender and tapering rearward, its height at origin of ist dorsal about ^i its
length to origin of caudal, the midline of the back with a low but sharp-edged and unmis-
takable dermal ridge running rearward from about opposite the ist or 2nd gill opening
nearly to the origin of the caudal. Trunk sector to cloaca a little shorter than tail sector.
Caudal peduncle subcylindrical, without lateral ridges or precaudal pit either above or
below. Dermal denticles irregularly spaced, sometimes overlapping but sometimes with

6. Mento may occur in Argentine waters, see p. 260.

246 Memoir Sears Foundation for Marine Research

areas of skin visible between, the blades nearly horizontal, on short pedicels, lanceolate,
strongly sculptured with 2 to 6 longitudinal ridges, the median pair flanking the axis of the
blade and usually extending out to its extreme tip, the basal part so opaque that the pedicel
is not visible from without, even in fresh specimens, their margins usually entire but some-
times weakly notched between the tips of the ridges, and often irregularly worn.

Head flattened above, its dorsal outline nearly straight, sloping to thin-tipped snout.
Snout broadly ovate at tip, its length in front of mouth a little greater than Ys the length
of head to origin of pectoral. Eye oval, its horizontal diameter slightly shorter than dis-
tance between nostrils, the subocular fold separate at both ends from the margin of the
lower eyelid in small specimens, but joining the latter at the anterior corner of the eye
by the time a total length of about 700 mm. is reached 5 in very large specimens it becomes
continuous with the margin of the upper lid at both corners. Spiracle oval, about Ve to Yi
as long as horizontal diameter of eye, about on a level with center of latter and behind it
by a distance V2 as long as horizontal diameter of eye. Third and fourth gill openings
slightly the longest, about i Ys times as long as horizontal diameter of eye, the 5th con-
siderably the shortest, their outlines nearly straight or weakly concave anteriorly, the 5th
the most so, the 4th gill opening above origin of pectoral. Nostril about as long as hori-
zontal diameter of eye, oblique, its inner corner about Ys as far from front of mouth as
from tip of snout, its anterior margin expanded as a well developed subpentagonal lobe
with blunt tip. Mouth occupying between Y2 and % of breadth of head, ovate, about twice
as broad as high. A strongly marked labial furrow on each jaw, the upper usually consider-
ably the longer in northern specimens, but sometimes only about as long as the lower, or
even slightly shorter, in West Indian and South American races.^

Teeth ^ in specimen counted, usually 5 to 7 rows functional, the cutting edges with
bluntly rounded apices directed somewhat outward {i.e., asymmetrical), their margins
slightly concave (the outer usually the more deeply so) or sometimes even notched, espe-
cially in small specimens.

First and second dorsals similar in shape, with very slightly convex or nearly straight
anterior margins, narrowly rounded apices, deeply concave rear margins and acute rear
corners, the free lower edges about Ys as long as the bases. Origin of ist dorsal about over
midpoint of inner margin of pectoral, the midpoint of its base about as close to axil of pec-
toral as to origin of pelvics. Second dorsal nearly or quite as long as ist at base, but only
about % to % as high, its origin at a perpendicular about midway between tips of pelvics
and origin of anal, its rear tip a little anterior to rear tip of anal. Caudal about Yo of total
length, with truncate tip, the terminal sector noticeably large or a little more than Vs of
total length of caudal, with well marked subterminal notch, the lower anterior contour
expanded as a low but well marked lobe, with broadly rounded apex. Anal only about %
as long at base as 2nd dorsal, and about Y2 as high, with less deeply concave posterior mar-
gin and shorter free rear corner, its origin about under midpoint of base of 2nd dorsal.

7. See Bigelow and Schroeder, Proc. Boston Soc. nat. Hist,, ^i (8), 1940: 422.

Fishes of the Western North Atlantic 247

Pelvics with nearly straight anterior and weakly concave posterior margins, narrowly
rounded apices and subacute tips, their origins considerably closer to origin of anal than to
origin of pectoral. Pectoral % as broad as long, or a little more, and a little larger in area
than 1st dorsal, with moderately convex outer and inner margins, nearly straight distal
margin, narrowly rounded apex and broadly rounded inner corner.

Color. Adults in life are plain grayish olive, slaty-gray or brown above, without any
definite darker markings j yellowish or grayish white below with the margins of the fins
paler. In embryos, however, and in young specimens up to a length of 400 mm. or so, the
upper part of the first dorsal is more or less widely edged with dusky gray, the apex of the
second dorsal sooty-edged or tipped, its posterior margin white; the caudal with a sooty
blotch above at tip, white-edged below. The strength of these dark markings is variable,
however, and by the time a length of 600 to 700 mm. is reached they are only faintly dis-
cernible, or have wholly faded.

Color Changes. This species is capable of changing shade — to a degree unusual among
sharks — by expansion or contraction of its melanophores, from dark gray above on a dark
background to a pale, translucent pearly tint when lying or swimming over a pale sand
bottom. It has been found by experiment that it requires about two days* for it to attain the
maximum paleness.

Size. Smooth Dogfish range from about 340 mm. to about 390 mm. in length (aver-
age about 360 mm.) at birth. The majority of mature females with young are between
1,000 and 1,300 mm. (about 3 feet 3 inches to 4 feet 4 inches) long. The maximum length
is about five feet.

Remarks. There is some evidence that in the tropical part of its range, where the
stocks of cants appear to be resident rather than migratory as they are in the north, local
populations may differ slightly from the typical form in their proportionate dimensions,
especially in the relative lengths of the labial folds, in the outlines of the fins and in the
sculpturing of the denticles.

Developmental Stages. It has long been known that this species, like its close relative
M. mustelus of the eastern Atlantic, is truly viviparous, the embryo deriving its nourish-
ment from its mother by a highly organized yolk-sac placenta." The number of young in a
litter usually ranges between 10 and 20 (average about 16 in a large number of gravid
females recently examined at Woods Hole) , but litters as small as four have been reported.

Habits. This is an inhabitant of the continental shelf and is not pelagic. During its
summer stay on the coast in the northeastern part of its range (see p. 249), it is most
commonly taken in comparatively shoal water of 10 fathoms or less, often coming into
enclosed bays and harbors, or even into fresh water on occasion.^" Large numbers of them

8. For accounts of these experiments, some of which we have witnessed, see Parker and Porter (Biol. Bull. Wood's
Hole, 66, 1934: 30-37) and Parker, G. H. (Color Changes of Animals, Univ. Penn. Press, Sect. 2, 1936: 12-20).

9. For account of this structure, see Fowler (Science, 50, 1909: 815) and Ranzi (Pubbl. Staz. zool. Napoli, /j,
1934: 387) for the European M. mustelus with list of earlier references.

10. Definitely reported in fresh water in the North East River, Maryland, by Gunter (Amer. Midi. Nat., 28, 1942:
316).

248 Memoir Sears Foundation for Marine Research

are taken in pound nets, as well as on hook and line close to bottom or actually on it. How-
ever, we are told by a well informed fisherman that a few are also caught along the outer
edge of the continental shelf on the so-called "Tilefish Grounds," in the offings of New
York and of southern New England, in depths of 80 to 90 fathoms during June and
September; and the species is described to us as a midwater form around Cuba.^^

Its food consists chiefly of the larger Crustacea, with crabs of one species or another
ranking first in most localities. It also preys largely on lobsters, of which it is perhaps the
most destructive enemy ofF the southern New England coast, where lobsters have been
found to form up to 1 6 per cent of its food. Smooth Dogfish also feed on squid, especially
in spring, and on whatever small fish may be available, such as menhaden (Brevoortia),
tautog (Tautoga), puffers (Spheroides), sticklebacks (Gasierosteus), scup (Stenotomus)
and sculpins (Myoxocephalus) . It has been estimated that in Buzzards Bay 10,000
Smooth Dogfish might annually devour over 60,000 lobsters, and perhaps V5 million
crabs, as well as possibly 70,000 fish of one kind or another. They also feed on mollusks to
a lesser extent, both univalve and bivalve, as well as on worms (Nereis). And they swallow
considerable quantities of eel grass (Zosiera), although probably only incidental to the
capture of their animal food.^^ They are also scavengers when occasion offers; off Havana,
for example, they have often been taken with garbage (chicken-heads, etc.) in their stom-
achs." Experiments have shown that food is found chiefly by the sense of smell,^* although
they also have fairly keen vision for nearby objects. A crab, for instance, is found as quickly
when hidden as if lying in the open. In captivity, and no doubt at liberty also, they con-
stantly search the bottom for food. When a crab is found it is shaken to and fro and de-
voured. It has also been observed in the aquarium that they never molest active fish, but
soon devour any sick or injured specimens, suggesting that it is only the smaller fishes that
they normally capture in any considerable numbers.

The fact that every female of i,000 mm. or longer taken at Woods Hole in the first
half of July during a recent investigation had either ovulated or was in the process of so
doing shows this to be the mating season. Corresponding to this, many kept in captivity
became pregnant during the late summer. The period of gestation appears to be about ten
months, i.e., the Smooth Dogfish carries her young during the winter migration.^" Off
southern New England the young are born between early May and mid- July, when new-
born specimens are often caught in the pound nets.

Such facts as are known regarding its winter and summer ranges show that the north
and south migrations of the northern stock of this species are chiefly thermal in nature. Thus
the temperature of its winter home ranges from about 6° to 7° C. (43°-45° F.) up to

11. Personal communication from Luis Howell-Rivero.

12. For lists of stomach contents and estimates of destructiveness, see Field (Rep. U.S. Comm. Fish. [1906], Spec.
Pap. 6, 1907: 12, 15).

13. Personal communication from Luis Howell-Rivero.

1 4. Sheldon, J. exp. Zool., 10, 1 9 1 1 : 51; Parker and Sheldon, Bull. U.S. Bur. Fish., 32, 1913: 33; Parker, Bull. U.S.
Bur, Fish., 33, 1914: 61.

15. Unpublished studies by F. L. Hisaw and A. Abramowitz.

Fishes of the Western North Atlantic 249

about 10° to 15° C. (50°-59° F.) ; it does not appear on the coast of the Middle Atlantic
United States or southern New England until the bottom water has warmed to at least
6° to 7° C. or higher} it departs in autumn when the temperature falls below about 10° to
12° C. At the opposite extreme, specimens kept in the tanks at Woods Hole show no ill
effects at the highest summer readings which are usually up to about 22° to 23° C. or 72°
to 73° F., but there is some evidence of withdrawal locally when the water warms above
about 70° F. The Caribbean populations of the species are inhabitants of tropical tempera-
tures.

Between North Carolina and Cape Cod the Smooth Dogfish moves north and south
regularly with the seasons, wintering chiefly within the sector between the southern half
of North Carolina and the offing of Chesapeake Bay. In the southern part of this range
many are caught in beach seines at times during the cold months. We have found odd
specimens on the flats in Pamlico Sound in early winter, chilled to death by the sudden
onset of freezing weather. Further north, however, along the southern Virginia coast,
where water temperatures are lower, winter records of them are chiefly from the offshore
fishing banks, in depths of 30 to 60 fathoms, where they are abundant enough to be a
nuisance.

The northward migration is progressive from early spring on, considerable numbers
lingering in North Carolina waters until June, with occasional specimens present there into
July, although none are seen thereafter. They appear by May along the peninsula that
separates Chesapeake Bay from the Atlantic," and they arrive nearly simultaneously at
about this same season all along the coasts of New Jersey, New York and southern New
England, the average date of their vernal arrival at the entrance to Long Island Sound
for a fifteen-year period being May 10, the earliest record being May 2. As a rule they
also appear in appreciable numbers at Woods Hole some time in May. However, the date
of their vernal arrival varies locally from year to year. Near New York, for example,
they usually are not plentiful until June. They summer all along the coast from Delaware
Bay to Cape Cod in such numbers that every fisherman is familiar with them, and they are
also present to some extent along the outer part of the continental shelf off southern New
England (p. 248). But the return movement of "Smooth Dogs" southward from south-
ern New England may commence as early as July, a decrease in their numbers often having
been reported there after June, with a corresponding increase in their numbers from sum-
mer to September near New York and offshore on the outer edge of the continental shelf.
But no mass movement southward takes place until late October or November, when they
withdraw almost simultaneously from the coast line as a whole to as far south as Chesa-
peake Bay, though stragglers (most often small specimens) may linger in the vicinity of
New York, and no doubt elsewhere, as late as the second week in December during some
years." Presumably they also withdraw from the outer edge of the continental shelf off

16. For some reason, odd specimens only have been taken within the Bay.

17. Latest date for the mouth of Long Island Sound, Dec. 13.

250 Memoir Sears Foundation for Marine Research

New York at about the same time, for none are taken there in winter according to reports
of fishermen J in fact, there is no record of their capture between mid-December and early
May anywhere to the northward of the offing of Chesapeake Bay.

Numerical A bundance. Along southern New England and on the mid-Atlantic coast
of the United States the Smooth Dogfish is the second most numerous shark, although fall-
ing far short of the Spiny Dogfish {Squalus acanthius, p. 466). Old accounts report occa-
sional catches of as many as 100 at a time in pound nets during their periods of abundance,
with 10 to 20 on a hand line not exceptional in a few hours' fishing, though catches of 5 or 6
in this way are more usual. More precise information is that 373 specimens were brought
in to the Laboratory of the U. S. Bureau of Fisheries at Woods Hole during an investiga-
tion of the food of the species, in the summer of 1 903, from pound net catches varying from
I to 41, and averaging about 7. It is also described as the most common local shark in
Uruguayan waters at the opposite extreme of its geographic range (p. 251). While the
populations of the intervening regions (Caribbean and Gulf of Mexico) have attracted
very little attention, they may be numerous there also, for Smooth Dogfish are common
around Cuba'* and have been so described around Bermuda."

Relation to Man. In spite of its abundance the Smooth Dogfish is of no commercial
importance except for classroom study in schools, for which purpose considerable numbers
are preserved yearly. Many are caught incidentally by anglers, for they bite freely when
fish or squid are used for bait, and they usually take the hook more freely by night than by
day, as so many sharks do. But few anglers consider them game fish.

Range. Western Atlantic; abundant northward to Cape Cod during part of year,
occasionally to Massachusetts Bay, and as a stray to Passamaquoddy Bay at the mouth of
the Bay of Fundy; southward to Texas, Cuba, the Caribbean region, central Brazil (Rio
de Janeiro) and Uruguay; also Bermuda. Present indications are that several more or less
isolated populations of Smooth Dogfish exist, their areas of distribution being separated
one from the next by wide gaps between which little or no intermigration occurs. The best
known of these populations is found along the coasts of the Middle Atlantic United States.
To the northward the Smooth Dogfish occurs regularly as far as Cape Cod, but only
as a stray in the southwestern part of the Gulf of Maine (odd specimens reported from
time to time for different localities in Massachusetts Bay), while only a single speci-
men has been reported from farther north, i.e., from St. Andrews on Passamaquoddy Bay
at the mouth of the Bay of Fundy. What the barrier may be to a more general dispersal
of them into the Gulf of Maine is not clear. Since they may appear on the coast of southern
New England when the temperature has not yet risen above 7° to 8° C, and since they are
most plentiful there during June when the water is still only 13° to 15° C, it is unlikely
that their failure to pass Cape Cod more regularly or to reach Georges Bank at all is
the result of temperature. Nor is there any other obvious explanation, for it seems hardly

18. Personal communication from Luis Howell-Rivero. 19. Goode, Bull. U.S. nat. Mus., 5, 1876: 73.

Fishes of the Western North Atlantic 251

likely that the change in the specific composition of the crab fauna (on which they largely
subsist) from the waters west of Cape Cod to those to the east can be responsible.

Present indications also are that this particular population is bounded equally sharply
to the southward, for while "Smooth Dogs" are common in season off the coasts of south-
ern Virginia'" and North Carolina, at least as far as Cape Lookout, there are only three
reports of the species for South Carolina (including one specimen in the collection of the
Harvard Museum of Comparative Zoology), and no positive record or rumor of its
presence on the east coast of Florida."

In contrast with the considerable amount of information that has accumulated about
this species along the east coast of the United States, little is known of it in the Gulf of
Mexico and the Caribbean, except that it occurs on the coast of Texas,''^ around Cuba and
Jamaica in some numbers (p. 250), at Curagao,^' and at Trinidad.^* Whether the Cuban
center of population receives any recruits in winter from the northern stock is not known.
Positive knowledge of the distribution farther south is even more scant, for while it has
been reported repeatedly by name from southern Brazil, Uruguay and northern Argen-
tina, there is no knowing how many of these records actually may have referred to the
newly described schmitti (p. 261 ).^° But M. canis does occur in southern Brazil, as proved
by the fact that the collection in the Harvard Museum of Comparative Zoology includes
several small specimens from Rio de Janeiro, as well as 3 1 embryos and the head of a large
female from an indeterminate Brazilian locality that we cannot distinguish from canis.
And we judge from an excellent illustration"" that a Mustelus, said to be the most common
Uruguayan shark," is likewise identical with the North American canis. Comparison also
of a specimen from Bermuda with extensive series from North America, West Indies and
Brazil failed to reveal differences sufficient to warrant specific separation."* The coastwise
nature of this species makes it likely that the Bermudian population has long been entirely
isolated.

Synonyms and References:"'

S^Wtt/ <ra«!V Mitchill, Trans. Lit. Phil. Soc. N. Y., /, 1815:486 (type loc, N. York).

Mustelus canis DeKay, Zool. N. Y., 4, 1842: 355, pi. 64, fig. 209 (descr., N. York) ; Linsley, Amer. J. Sci.,
4y, 1844: 77 (Connecticut); Storer, Mem. Amer. Acad. Arts Sci., 2, 1846: 505 (in synopsis); Baird,

20. Only occasional specimens are taken in Chesapeake Bay.

21. Canis has been recorded from southern Florida, but it is probable that these reports actually referred to the
newly discovered M. norrisi, for which that is the center of abundance (p. 255).

22. See Study Material, p. 244. It has also been reported as common in fresh water in Louisiana (Fowler, Proc.
biol. Soc. Wash., ^6, 1933 : 57), but perhaps not on good evidence.

23. Measurements given by Metzelaar (Trop. Atlant. Visschen, 1919: 5) identify this specimen as canis, not norrisi.

24. We have examined the embryo reported by Fowler (Proc. Acad. nat. Sci. Philad., 6y, 1916: 521) from Trinidad.

25. For discussion of this species, see Bigelow and Schroeder (Proc. Boston Soc. nat. Hist., ^i [s], 1940: 423).

26. Devincenzi and Barattini, An. Mus. Hist. nat. Montevideo, Suppl. Atlas, (2) 2, 1926: pi. 2, fig. 2.

27. Devincenzi, An. Mus. Hist. nat. Montevideo, (2) /, 1920: 12.

28. Bigelow and Schroeder, Proc. Boston Soc. nat. Hist., 41, 1940: 417.

29. The many studies of the physiology of Mustelus canis are omitted here, unless pertinent to knowledge either
of itf geographic distribution or of its habits.

252 Memoir Sears Foundation for Marine Research

Rep. Smithson. Instn. (1854), 1855: 337 extra (abund., N. Jersey); DeKay, Rep. St. Cab. nat. Hist.
N. Y. (185s), 8, 1858: 64 (N. York); Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861: 59 (ref .) ;
Proc. Acad. nat. Sci. Philad., 1863: 333 (Massachusetts); Putnam, Bull. Mus. comp. Zool. Harv., /,
1863: 10 (ref.); Gill, Proc. Acad. nat. Sci. Philad., 1864: 263 (class.); Dumeril, Hist. Nat. Poiss., i,
1865: 402 (ref., N. York) ; Storer, Mem. Amer. Acad. Arts Sci., 9, 1867: 227, pi. 37, fig. 2; also Fishes
Mass., 1867: 251, pi. 37, fig. 2 (descr., Massachusetts); Poey, Repert. Fisico.-Nat. Cuba, 2, 1868: 453
(Cuba) ; Glinther, Cat. Fish. Brit. Mus., 8, 1870: 386, 518 (ref.) ; Gill, Rep. U.S. Comm. Fish. (1871-
72), 1873:813 (C. Cod toC. Hatteras); Verrill, Rep. U.S. Comm. Fish. (1871-72), 1873: 521 (Mas-
sachusetts, food) ; Goode, Bull. U.S. nat. Mus., 5, 1876: 73 (common, Bermuda) ; Poey, Enumerat. Pise.
Cubens., 1876: 201 ; An. Soc. esp. Hist, nat., 5, 1876: 397 (Cuba); Goode, Amer. J. Sci., 14, 1877:
293 (Bermuda); Goode and Bean, Bull. Essex Inst. Salem, //, 1879: 30 (Massachusetts Bay); Bean,
T. H., Proc. U.S. nat. Mus., j, 1881: 115 (N. Jersey to Massachusetts) ; Jordan and Gilbert, Proc. U.S.
nat. Mus., 5, 1882: 581 (S. Carolina) ; Bull. U.S. nat. Mus., 16, 1883: 870 (comp. with Europ. species) ;
Bean, T. H., Rep. U.S. Comm. Fish. (1882), 1884: 343 (off Woods Hole); Bull. U.S. Fish Comm.
(1887), 7, 1889: 152 (N.Jersey) ; Nelson, Rep. St. Geol. N. J., 2 (2), 1890: 660 (common, N. Jersey) ;
Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 214, part (C. Cod, Cuba) ; Bull. U.S. nat.
Mus., 47 (i), 1896: 29, part (C. Cod, Cuba) ; Bean, T. H., Bull. Amer. Mus. nat. Hist., p, 1897: 329
(near N. York) ; Linton, Proc. U.S. nat. Mus., 20, 1897: 423, 433 (parasites, Woods Hole) ; Smith, BulL
U.S. Fish Comm., 77, 1898: 88 (food, abund. near Woods Hole) ; Linton, Bull. U.S. Bur. Fish., rp, 1901:
270 (parasites. Woods Hole) ; Bean, Rep. For. Comm. N. Y., 1901 : 376 (abund., season, near N. York) ;
Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 23 (genera!) ; Sharp and Fowler, Proc. Acad. nat. Sci. Philad.,
56, 1904: 505 (Nantucket); Bean, Field Mus. Publ., Zool., (7) 2, 1906: 29 (Bermuda); Tracy, Rep.
R. L Comm. inl. Fish., 1906: 49 (Rhode Island) ; Field, Rep. U.S. Comm. Fish. (1906), Spec. Pap. 6,
1907: 10 (food. Woods Hole) ; Linton, Bull. U.S. Bur. Fish., 27, 1907: 125, 410-414, 425 (parasites,
Woods Hole) ; Wilson, Proc. U.S. nat. Mus., j/, 1907: 707, 711 (parasites) ; Proc. U.S. nat. Mus., 33,
1907: 360, 422, 423 (parasites) ; Smith, Bull. N. C. geol. econ. Surv., 2, 1 907: 32 (N. Carolina) ; Ken-
dall, Occ. Pap. Boston Soc. nat. Hist., 7(8), 1908: 3 (New England) ; Parker, Bull. U.S. Bur. Fish., 29,
1910: 43 (senses, movements); Linton, Bull. U.S. Bur. Fish., 28, 1910: 120 (parasites. Woods Hole);
Tracy, Rep. R. I. Comm. inl. Fish., 1910: 58 (Rhode Island); Sheldon, J. exp. Zool., 10, 1911: 54
(sense of smell, feeding) ; Parker and Sheldon, Bull. U.S. Bur. Fish., 52, 1913: 33-36 (food found by
smell) ; Fowler, Proc. Acad. nat. Sci. Philad., 65, 1 91 3: 62 (Virginia) ; Nichols, Abstr. Linn. Soc. N. Y.,
20-23, 191 3: 91 (season off N. York) ; Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., 31, 1913: 734
(Woods Hole); Parker, Bull. U.S. Bur. Fish., 33, 1914: 61 (food, found by smell); Fowler,
Proc. Acad. nat. Sci. Philad., 66, 1914: 353 (N. Jersey); Coles, Proc. biol. Soc. Wash., 28, 1915:
89 (N. Carolina); Fowler, Proc. Acad. nat. Sci. Philad., 67, 1916: 521 (Trinidad, W. Indies);
Wiegmann and Nichols, Copeia, 23, 1915: 43 (near N. York); Nichols and Murphy, Brooklyn
Mus. Sci. Bull., 3 (i), 1916: 7 (Long Island, N. York); Fowler, Copeia, 27, 1916: 12 (N.
Jersey); Jordan, Copeia, 49, 1 91 7: 87 (name); Latham, Copeia, 41, 1 91 7: 17 (mouth Long
Island Sd.) ; Copeia, 57, 1918: 53 (no. of embryos); Copeia, 71, 1919: 53 (dates of arrival and
departure, mouth Long Island Sd.) ; Fowler, Copeia, 68, 1919: 13 (N. Jersey); Wilson, Proc. U.S.
nat. Mus., 55, 1920: 596 (parasites, N. Carolina); Fowler, Proc. biol. Soc. Wash., 33, 1920: 144 (N.
Jersey); Latham, Copeia, 87, 1920: 91 (mouth Long Island Sd.) ; Devincenzi, An. Mus. Hist. nat.
Montevideo, (2) /, 1 920: 12 (Uruguay); Breder, Copeia, 1 01, 1 921: 85 (food); Fowler, Proc. Acad,
nat. Sci. Philad., 74, 1922: 3 (N. Jersey); Breder, Copeia, 127, 1924: 26 (near N. York); Mowbray,
Bull. N. Y. zool. Soc, 28, 1925: 75 (near N. York); Fowler, Copeia, 143, 1925: 41, 42 (N. Jersey,
Delaware Bay); Devincenzi and Barattini, An. Mus. Hist. nat. Montevideo, (2) 2, 1926: Suppl. Atlas,
pi. 2, fig. 2 (Uruguay); Coles, Copeia, 151, 1926: 105 (N. Carolina) ; Fowler, Copeia, 156, 1926: 146
(N. Jersey); Copeia, 165, 1927: 89, 90 (Maryland, Virginia, Delaware); Nichols and Breder, Zoo-
logica, N. Y., 9, 1927: 13 (general); Truitt, Bean and Fowler, Bull. Md. Conserv. Dep., 3, 1929:
29 (Maryland, Chesapeake Bay); Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 14 (general);
Fowler, Proc. Acad. nat. Sci. Philad., 80, 1929: 607 (N. Jersey); McCallum, Proc. U.S. nat. Mus.,
79 (26), 1931: 2 (parasites); Breder, Copeia, 1932: 31 (Block Island); Lundstrom and Bard, Biol. Bull.

Fishes of the Western North Atlantic 253

Wood's Hole, 62, 1932: I (color changes); Pearson, Invest. Rep., U.S. Bur. Fish., (lo) /, 1932: 19
(winter trawl fishery off North Carolina) ; Fowler, Proc. biol. Soc. Wash., 46, 1933: 57 (Calcasieu R.,
Louisiana, fresh water, but we question identification) ; Bigelow and Schroeder, Canad. Atlant. Fauna,
biol. Bd. Canad., 12', 1934: 6 (descr., distrib.) ; Burton, Sci. Mon. N. Y., 40, 1935: 283 (S. Carolina) ;
Parker and Porter, Biol. Bull. Wood's Hole, 66, 1934: 30 (color changes) ; Fowler, Bull. Amer. Mus. nat.
Hist., 70 (l), 1936: 68 (in part); Parker, Biol. Bull. Wood's Hole, yo, 1936: 6, pi. I (color changes
of young); Color Changes of Animals, Univ. Penn. Press, 1936: 12 (color changes); Smith, H. W.,
Biol. Rev., II, 1936: 64 (ref. to report of it in fresh water by Fowler, 1933) ; Fowler, Proc. Acad. nat.
Sci. Philad., <?9, 1937: 303 (N. Jersey); Hubbs, Occ. Pap. Mus. Zool. Univ. Mich., 374, 1938: I, 13
(correct name); Parker, Proc. Amer. Acad. Arts Sci., 72 (7), 1938: 269 (color changes in young);
Tortonese, Atti Soc. ital. Sci. nat., yy, 1938: 303 (part, Brazil) ; Springer, Proc. Fla. Acad. Sci., 5, 1939:
14 (had no Florida record) ; Breder, Bull. N. Y. zool. Soc, 41, 1938: 28 (N. York Harbor) ; Springer,
Proc. U.S. nat. Mus., 86, 1939: 461 (comp. with other species); Bigelow and Schroeder, Proc. Boston
Soc. Nat. Hist., 41, 1940: 417, pi. 14, fig. A, B, pi. 15, fig. A, B, C, pi. 17, fig. A, B (compar. with other
species); Gunter, Amer. Midi. Nat., 28, 1942: 316 (in fresh water, Maryland); Fowler, Arqu. Zool.
Estado Sao Paulo, 3, 1942: 129 (Brazil) ; Fish Culturist, 21 (9), 1942: 66 (listed) ; Lunz, Bull. S. C.
St. Planning Bd., 14, 1944: 26 (S. Carolina, Florida); Merriman and Warfel, Trans. N. Amer. Wild
Life Conf., 1944: 234 (Block Island, Rhode Island) ; Bigelow and Schroeder, Guide Comm. Shark Fish.,
Anglo Amer. Caribb. Comm., Wash., 1945: 1 13 (descr., ill., range).

Mustelus laevis Gunther, Cat. Fish. Brit. Mus., 8, 1 870: 385 (in part); Metzelaar, Trop. Atlant. Visschen,
1 91 9: 5 (meas., Curasao, W. Indies) ; not M. laevis Risso, 1826.

Mustelus hinnulus Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 19 (part, C. Cod) ; not Squalus (Galeo-
rhinus) hinnulus Blainville, in Vieillot, Faune Franc, 1 825: 83.

Gdeus cams ]or Am, Rep. U.S. Comm. Fish. (1885), 1887: 795; Moore, Bull. U.S. Fish Comm., 12, 1894:
358 (abund., N. Jersey) ; not Gd/<r«.r ca««V Muller and Henle, 1841.

Cynias canis Gill, Proc. U.S. nat. Mus., 26, 1 903: 960 (name) ; Jordan, Manual Vert. Anim. NE. U.S., 1929:
9 (general); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 13 (part); Beebe
and Tee- Van, Field Bk. Shore Fish. Bermuda, 1933: 24 (Bermuda).

Cynais canis Fowler, Rep. N. J. Mus. (1905), 1906: 60 (N. Jersey) ; Fowler, Rep. N. J. Mus. (1906), 1907:
254 (N. Jersey) ; Proc. Acad. nat. Sci. Philad., 60, 1908: 55 (N. Jersey, Rhode Island, Nantucket) ; Proc.
Acad. nat. Sci. Philad., 61, 1909: 407 (N. Jersey) .°°

Mustelus mustelus Fowler, Science, 50, 1909: 815 (embryo with placenta, N. Jersey); Proc. Acad. nat.
Sci. Philad., 64, 1912: 57 (Virginia) ; Copeia, 30, 1916: 36 (Mid. Atlant. Coast) ; Copeia, 31, 1916: 41
(N. Jersey) ; Proc. Acad. nat. Sci. Philad., 69, 1917: 109 (N. Jersey) ; Proc. Boston Soc. nat. Hist., 35,
1917: no (Rhode Island, Nantucket); Copeia, 39, 1917: 4 (Argentina, name); Occ. Pap. Mus. Zool.
Univ. Mich., 56, 1918: 15, pi. 2 (ill., descr., embryos with placenta, N. Jersey); Proc Acad. nat. Sci.
Philad., //, 1920: 292 (N. Jersey); Hildebrand and Schroeder, Bull. U.S. Bur. Fish., 43, 1928: 47
(Chesapeake Bay); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 13 (part);
Beebe and Tee-Van, Zoologica, N. Y., 13, 1932: 1 19 (Bermuda); Field Bk. Shore Fish. Bermuda,
1933: 24 (Bermuda); Bigelow and Schroeder, Bull. U.S. Bur. Fish., 48, 1936: 321 (off N. Carolina,
winter).

Galeorhinus laevis Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 176 (part) pi. 4, fig. 6-9;'^ Radcliffe,
Trans. Amer. Fish. Soc, 1914: 38 (N. Carolina); Bull. U.S. Bur. Fish., 34, 1916: 267, pi. 42, fig. 3
(denticles, teeth, N. Carolina) ; Huntsman, Contr. Canad. Biol. (1921), 3, 1922; 56 (St. Andrews, Passa-
maquoddy Bay); Meek and Hildebrand, Field Mus. Publ., Zool., 75, 1923: 34 (part); Bigelow and
Welsh, Bull. U.S. Bur. Fish., 40, 1925: 25 (descr.. Gulf of Maine) ; White, Bull. Amer. Mus. nat. Hist.,
■J4, 1937: 64, 92, 12^, pi. 6, fig. C, pi. 13, fig. P, pi. 41, fig. A (class., anat.) ; not Squalus {Galeorhinus')
/iiefw Blainville, 1825.

30. The locality suggests that a nominal record of C. canis from the Florida Keys (Fowler, Proc. Acad. nat. Sci.
Philad., 5*, 1906: 79) probably referred to the newly described Mustelus norrisi; see Synonyms, p. 256.

31. Carman's description was based on a Mediterranean specimen of the European Af. mustelus, but his illustrations
on a canis from Long Island, New York.

254 Memoir Sears Foundation for Marine Research

"Mustelus asterias (Valmont) or Cynias cams Mitchill," Jordan, Copeia, 49, 1917: 87 (name).

Doubtful South American References:

Galeus cani?"^ Berg, An. Mus. nac. B. Aires, (2) 4, 1895: 7 (Argentina, might equally refer to schmitti);

Lahille, Rev. Mus. paul., 6, 1895: 276 (Argentina, might equally refer to schmitti).
Mustelus vulgaris Bassett-Smith, Proc. zool. Soc. Lond., 1899: 468 (Rio de Janeiro, might refer equally to

schmitti) .
Mustelus canis Eigenmann, Rep. Princeton Exped. Patagonia (i 896-1899), 3 (2), 1910: 377 (La Plata R.,

might equally refer to schmitti) ; Gilbert, Proc. Wash. Acad. Sci., 2, 1 900: 161 (Maceio, Brazil, might

equally refer to schmitti); Schreiner and Ribeiro, Arch. Mus. nac. Rio de J., 12, 1903: 78 (Rio de

Janeiro, might equally apply to schmitti); Evermann and Kendall, Proc. U.S. nat. Mus., 31, 1906: 68

(Rio de la Plata, might equally refer to schmitti) ; Puyo, Bull. Soc. Hist. nat. Toulouse, yo, 1936: 89

(French Guiana, name only; might equally refer to norrisi).
Mustelus laevis Schreiner and Ribeiro, Arch. Mus. nac, Rio de J., 12, 1 903 : 78 (Rio de Janeiro, might equally

refer to schmitti).
Cj/mW cawjV Ribeiro, Arch. Mus. nac, Rio de J., 14, 1907: 161, 203 (Brazil, might equally apply to jcA»j»«») ;

Fauna brasil. Peixes, Mus. nac. Rio de J., 2 (i), Fasc. I, 1923: 19, pi. I (Brazil, might equally refer to

schmitti) .
Mustelus mustelus Fowler, Copeia, 39, 1917: 4 (Argentina, might equally apply to schmitti) ; Proc. Acad. nat.

Sci. Philad., y8, 1926: 261 (Argentina, might equally refer to schmitti).
Mustelus asterias Lahille, Enum. Feces Cartilag. Argent., 1921: 13 (Argentina, name only); probably not

asterias Cloquet, 1 819, of the eastern North Atlantic.

Mustelus norrisi Springer, 1939

Florida Dogfish

Figure 43 D-F

Study Material. Three males, 677 to 825 mm, long, taken near Englewood, western
Florida (Harv. Mus. Comp. Zool., No. 35222, 35223, 35224) ; one male, 623 mm., from
the Florida Keys.

Distinctive Characters. Among Atlantic members of the genus, the most distinctive
feature of M. norrisi is the expansion of the lower anterior corner of its caudal as a sharp-
pointed lobe directed rearward (Fig. 43 D).

Description. Proportional dimensions in per cent of total length. Male, 671 mm.,
from Englewood, Florida (Harv. Mus. Comp. Zool., No. 35223). Male, 753 mm., from
same locality (Harv. Mus. Comp. Zool., paratype. No. 35222).

Trunk at origin of -pectoral: hrezdth 9.1, 8.55 height 8.2, 8.0.

Snout length in front of: outer nostrils 3.4, 3.25 mouth 6.1, 5.9.

Eye: horizontal diameter 2.6, 2.6.

Mouth: breadth 4.4, 4.25 height 3.1, 3.2.

Nostrils: distance between inner ends 2.S,i.s.

Labial furrow length: upper 1.3, i.2j lower i.i, i.i.

Gill opening lengths: ist 1.9, 1.65 2nd 2.1, 1.75 3rd 2.1, 1.9J 4th 2.1, 1.95 5th

1.7,1.5.

32. Galeus canis Lahille (Physis, B. Aires, 5, 1921: 63), Marini (Physis, B. Aires, 9, 1929: 452), and Pozzi and
Bordale (An. Soc. cient. argent., 120, 1935: 151) reported for Argentina appears to refer not to a Mustelus
but to the genus Galeorhinus ; see p. 264.

Fishes of the Western North Atlantic 255

First dorsal fin: vertical height 7.9, 8.1 5 length of base 10.4, 10.9.

Second dorsal fin: vtT{\cz\\\&\g\\t G.iy 5.8; length of base 8.2,7.9.

Anal fin: vertical height 3.0, 3.55 length of base 5.9, 5.4.

Caudal fin: upper margin 1 8.4, 17.8 ; lower anterior margin 8.6, 8.2.

Pectoral fin: outer margin 12.5, 12.65 inner margin 6.1, 5.8; distal margin 8.5,

9.1.

Distance from snout to: ist dorsal 28.6, 27.5; 2nd dorsal 61.2, 63.25 upper caudal

81.6, 82.2} pectoral 17.9, 17.25 pelvics 41.9, 42.4; anal 65.2, 67.7.

Intersface between: ist and 2nd dorsals 23.4, 25.2; 2nd dorsal and caudal 12.8,

11.1} anal and caudal 8.3, 8.0.

Distance from origin to origin of: pectoral and pelvics 24.4, 26. i } pelvics and anal

23.8,25.6.

M. norrisi resembles canis so closely that the points of difference alone need be noted.
These are: lower anterior corner of caudal acute instead of rounded, directed rearward,
forming a distinct but short lobe (Fig. 43 D)} midpoint of base of ist dorsal nearer to
origin of pelvics than to axil of pectoral by a distance about equal to horizontal diameter
of eye, instead of as near to axil of pectoral as to origin of pelvics, or nearer} body cavity
relatively shorter, with origin of pelvics about midway between origins of pectorals and of
anal, and under rear tip of ist dorsal, instead of being considerably nearer to origin of anal
than to origin of pectorals and behind rear tip of ist dorsal} pectorals with relatively shorter
inner and distal margins, the latter more deeply concave} mouth narrower} fins generally
smaller} labial folds not only shorter but the upper and lower folds of more nearly equal
length than is usual in typical canis, in which the upper is in most cases considerably the
longer} space between nostrils relatively narrower} trunk more slender, with its dorsal
outline less highly arched. The teeth also are higher-crowned in general, their margins
more deeply concave or even notched (Fig. 43 F)} dermal denticles, however, not dis-
tinguishable from those of typical canis.

Color. No information is available as to its color in life} preserved specimens are gray
above, paler gray or dirty white below and without definite markings.

Size. Norrisi reaches a corresponding stage in development at a size somewhat smaller
than does canis, the subocular fold being continuous anteriorly with the edge of the upper
eyelid, and the claspers of large size in males as small as about 620 mm. in length. And
females also may mature at a size no greater than this, one of 825 mm, having been found
to contain embryos nearly ready for birth.

Develofmental Stages. It is not yet known whether or not the embryo develops a
placental connection with the mother, as in canis and in mustelus (p. 247).

Habits. Nothing is known of its diet or of its breeding habits.

Range. This Dogfish is known up to the present time only from the Florida Keys
and from the west coast of southern Florida (Englewood and Naples, where large num-
bers have been taken in mackerel nets). It has been taken only in winter, suggesting that

256 Memoir Sears Foundation for Marine Research

it has a center of abundance in moderate depths and comes into shallow water only when
the temperature of the latter is near the seasonal minimum."

Synonyms and References:

Musielus norrid Springer, Proc. U.S. nat. Mus., 86, 1939: 462 (descr., type loc. off Englewood, Florida, also
vicinity Key West, Florida) ; Proc. Fla. Acad. Sci., 3, 1939: 15 (same loc. as the preceding, comp. with
other species) ; Bigelow and Schroeder, Proc. Boston Soc. nat. Hist., 41, 1940: 417, pi. 14, fig. A, pi. 15,
fig. F, pi. 17, fig. C (meas., comp. with other species); Lunz, Bull. S. Carolina St. Planning Bd., 14,
1944: 26 (Florida); Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Coram.,
Wash., 1945: I 10 (descr., ill., range).

\ Mustelus cams Y.vermzr\r\ and Kendall, Rep. U.S. Comm. Fish. (1899), 1 900: 48 (Key West, Florida)."

^Cynais canis Fowler, Proc. Acad. nat. Sci. Philad., 58, 1906: 79 (Key West, Florida)."

Genus Mustelus, Addendum

Under this heading we include accounts of two more species of the genus that occur
in the coastal waters of Uruguay and southern Brazil 5 likewise of a third that has been
reported from northern Argentina.

Mustelus fasciatus (Garman), 191 3

Striped Dogfish

Figure 43 Upper, A, B, C

Study Material. Female and male, 484 and 613 mm. long (the types), Rio Grande
do Sul, Brazil (Harv. Mus. Comp. Zool., No. 154 and 315).

Distinctive Characters. Fresh specimens of fasciatus are separable at a glance from
canis, norrisi and schmitti by their dark striped color pattern, and further, by a much longer
snout relative to the size of the eye,'° by the teeth, which are more nearly symmetrical,
their cutting edges evenly convex (Fig. 43 A), and by the distal margin of the pectoral,
which is considerably longer relative to the other margins of the fin. A further distinction
between fasciatus on the one hand and canis and norrisi on the other is that the lower ante-
rior corner of its caudal is not expanded as a definite lobe. The most obvious distinctions
between it and mento, which may also be dark-striped when young, are that the head (to
pectoral) is considerably longer than the interspace between the first and second dorsals in
fasciatus but shorter than the interspace in mento, and that the caudal of fasciatus lacks a
definite lower lobe (cf. Fig. 43 upper with 42 H).

Description. Proportional dimensions in per cent of total length. Female, 484 mm.,
from Rio Grande do Sul, Brazil (Harv. Mus. Comp. Zool., No. 315). Male, 613 mm.,
same locality (Harv. Mus. Comp. Zool., No. 154).

33. Springer, Proc. Fla. Acad. Sci., 5, 1939: 5.

34. These nominal records are referred tentatively to this species because of the locality.

35. Snout in front of mouth about 4.5 as long as horizontal diameter of eye in fasciatus, but only about 2.5 to 3
as long in either of these other three species.

Fishes of the Western North Atlantic

^S1

Trunk at origin of fectoral: breadth 1 1.6, I2.0; height 8.1, 8.5.

Snout length in front of: outer nostrils 5.7, 5.95 mouth 8.2, 8.0.

Eye: horizontal diameter 2.1, 2.4.

Mouth: breadth 7.2, 7.4; height 3.3, 3.7.

Nostrils: distance between inner ends 3.1, 3.3.

Labial furrow lengths: upper 2.3, 2.1 ; lower 2.0, 1.8.

Gill opening lengths: ist 3.0, 1.$; 2nd 3.0, 2.5; 3rd 3.0, 2.5; 4th 2.9, 2.35 5th

2.5, 2.0.

First dorsal fin: vertical height 8.5, 8.6 j length of base 14.6, 14.O.

Second dorsal fin: vertical height 6.0, 6.95 length of base 10.7, 10.4.

Anal fin: vertical height 3.0, 3.6; length of base 6.4, 6.8.

Caudal fin: upper margin 21. i, 21.6; lower anterior margin 7.9, 9.3.

Pectoral fin: outer margin 15. i, 14.7J inner margin 8.7, 9.0; distal margin 1 1.4,

12.9.

Distance from snout to: ist dorsal 28.5, 28.55 2nd dorsal 59.8, 59.55 upper caudal

Figure 43. Mustelus fasciatus, male, 607 mm. long, from Rio Grande do Sul, Brazil (Harv. Mus. Comp.
Zool., No. 154, type). A Upper teeth of same, about 6 x. 5 Dermal denticles of same, about 24 x. C Tracings
of pectoral fins adjusted to equal lengths along outer margin: dotted line, M. cants, 768 mm. long, from Woods
Hole, Massachusetts; solid line, M. schmitti, 578 mm. long, from Muldonado, Uruguay; broken line, M. fas-
ciatus, same specimen 3&\n A, B. D Mustelus norrisi, adult male, 643 mm. long, from Florida Keys (Harv.
Mus. Comp. Zool., No. 442). E Head of same, from below. F Upper teeth of same, about 7 x.

258 Memoir Sears Foundation for Marine Research

78.9, 78.4; pectoral 22.7, 22.O5 pelvics 49.0, 45. 4j anal 68.5, 65.7.

Intersface between: ist and 2nd dorsals 17.7, i7-i; 2nd dorsal and caudal 8.7,

8.4; anal and caudal S-(>y 4-6.

Distance from origin to origin of: pectoral and pelvics 26.9, 24.5 ; pelvics and anal

19.2, 20.4.

General form much as in canis. Dermal denticles more loosely spaced, with notice-
ably weaker sculpture, usually with only 2 ridges, and these as a rule confined to the ante-
rior basal half of the blades, the latter so transparent that the outlines of the pedicels are
visible from outside.

Head more flattened above than in canis and relatively much longer, its length to
origin of pectoral about equal to distance from rear base of ist dorsal to midbase of 2nd
dorsal. Snout in front of mouth a little more than V2 as long as pectoral (considerably
shorter than that in other western Atlantic species), more narrowly ovate than in canis.
Eye relatively small, its horizontal diameter only about V4 to Vs as long as snout in front
of mouth. Spiracle a little less than Y2 as long as horizontal diameter of eye. Third gill
opening a very little longer than horizontal diameter of eye, the 4th about over origin of
pectoral. Anterior margin of nostril with a well marked rounded lobe. Mouth a little less
than V2> or about 45%, as high as broad. Upper labial furrow about 70% as long as dis-
tance between nostrils, the lower furrow about % as long as upper.

Teeth nearly symmetrical, with evenly convex cutting edges.

First dorsal about as long at base as along anterior margin, the posterior margin only
very weakly concave (much less so than in canis or norrisi), its origin a little posterior to
axil of pectoral, the midpoint of base about midway between origin of pelvics and axil
of pectoral. Interspace between ist and 2nd dorsals about as long as from snout to ist gill
opening. Second dorsal between % and % as long as ist at base and about % as high, its
origin about midway between tips of pelvics and origin of anal. Caudal with lower anterior
corner considerably more obtuse than a right angle, not expanded as a definite lobe. Anal
about % as long at base as 2nd dorsal, its origin about under midpoint of base of latter.
Pectoral a little less than %, or about 60%, as long as head, with nearly straight distal
margin and very broadly rounded inner corner, the distal margin only a little shorter
(about 75-90%) than outer margin.

Color. Described^" as "back brown with narrow transverse bands of darker, separated
by spaces of about equal width: one or a pair crossing the orbits, one across the spiracles,
one on the nape, four between the nape and the dorsal, four or five on the base of the
dorsal, six between the dorsals, three on the base of the second dorsal, and two or three
between it and the caudal. A yellow spot in front of each eye above each orbit. Fins dark
with narrow edgings lighter. Lower surfaces whitish. On a larger individual pairs of bands
are more or less confluent and all are more indefinite, indicating a probable loss of the

36. Garman (Mem. Harv. Mus. comp. Zool., 515, 1913 : 173), perhaps from field notes from the "Thayer" Expedi-
tion on which the specimens were collected.

Fishes of the Western North Atlantic 259

bands later in life." After many years in alcohol the dark bands have wholly faded, leav-
ing the specimens, which were collected in 1858 or 1859, mouse-gray above and a paler
shade of the same tint below.

Size. The fact that the claspers are pnly about half as long as the inner margins of the
pelvics in a male of 613 mm. suggests that maturity is not reached until at a length of per-
haps 900 to 1,000 mm., or at about the same size as in cams (p. 247).

Developnental Stages. Not known.

Habits. Nothing known.

Range. M. fasciatus is so far known only from Rio Grande do Sul, southern Brazil
(the type locality), and from the vicinity of Montevideo, Uruguay, and from Argentina,
Lat. 35° 30' S."

Synonyms and References:

Galeorhinus fasciatus GitTam, Mem. Harv. Mus. comp. Zool., 36, 1913: 172 (type descr., south. Brazil).

Mustelus siriatus Devincenzi, An. Mus. Hist. nat. Montevideo, (2) i, 1920: 122, pi. 12 (descr., good photos;
vicinity of Montevideo, Uruguay); Devincenzi and Barattini, An. Mus. Hist. nat. Montevideo, (2) 2,
Suppl. Album Ictiol., 1926: pi. 2, fig. 3 (the dark cross bars indicated on ill.); Pozzi and Bordale, An.
Soc. cient. argent., 120, 1935: 151 (Lat. 35° 30' S.).

MusCelus fasciatus Tortonese, Atti Soc. ital. Sci. nat., yy, 1938: 305 (descr., meas., ill., Rio Grande do Sul,
Brazil); Springer, Proc. U.S. nat. Mus., 86, 1939: 467 (in Key to species of Mustelus) ; Bigelow and
Schroeder, Proc. Boston Soc. nat. Hist., 41 (8), 1940: 417, 418, Table column J; pi. 14, fig. B, pi. 15,
fig. D, pi. 17, fig. E, pi. 18, fig. E (meas., discus., proport. dimensions, ill. of fins, teeth, denticles);
Fowler, Arqn. Zool. Estado Sao Paulo, 3, 1942: 129 (listed for Brazil).

Mustelus mento Cope, 1877

Figure 42 H-J

Study Material. Female, 1,024 "irn- lo^^gj from Payta, Peru (Harv. Mus. G)mp.
Zool., No. 35246).

Distinctive Characters. See following Description.

Descri-ption. Proportional dimensions in per cent of total length. Female, 1,024 mm-j
from Payta, Peru (Harv. Mus. Comp. Zool., No. 35246).

Trunk at origin of fectoral: breadth 10.5; height 9.5.

Snout length in front of: outer nostrils 3.8 j mouth 5.6.

Eye: horizontal diameter 2.0.

Mouth: breadth 6.0 5 height 2.5.

Nostrils: distance between inner ends 2.5.

Labial furrow lengths: upper 2.45 lower 2.0.

Gill opening lengths: ist 2.95 2nd 2.95 3rd 2.9; 4th 2.95 5th 2.2.

First dorsal fin: vertical height 10.2 ; length of base 1 3.7.

Second dorsal fin: vertical height 7.05 length of base 10.7.

Anal fin: vertical height 4.OJ length of base 6.8.

37. Pozzi and Bordale, An. Soc. cient. argent., 120, 1935: 151.

26o Memoir Sears Foundation for Marine Research

Caudal fin: upper margin 1 8.95 lower anterior margin 8.6.

Pectoral fin: outer margin 16.O; inner margin 11.4; distal margin 14.2.

Distance from snout to: ist dorsal 27. i; and dorsal 61.O; upper caudal 81. i;

pectoral 17.9; pelvics45.i; anal 66.8.

Inters face between: ist and 2nd dorsals 2 1 . i ; 2nd dorsal and caudal 9.5 ; anal and

caudal 6.0.

Distance from origin to origin of: pectoral and pelvics 27.8 ; pelvics and anal 22.0.

Since it is doubtful (see below) whether this species actually occurs in the Atlantic,
its most distinctive features alone need be mentioned as an aid toward its identification. It
falls with canis, norrisi and schmitti among the western Atlantic species in the relative
shortness of its head} but it is separable at a glance from norrisi by the shape of its caudal
(cf. Fig. 42 H with 43 D) ; from schmitti by a much smaller eye relative to the gill open-
ings (only about 60% as long as the 3rd gill opening in mento, but about as long as the
3rd gill opening in canis and schmitti), and by the shape of its caudal (cf. Fig. 42 F with
42 H) } from canis similarly by a small eye, by the distance between its nostrils which is
considerably less than V2 as great as the breadth of its mouth (approximately one-half
as great as that in canis) and by a relatively larger pectoral (inner margin about 1.8 times
as long as snout in mento and only about 1.3 times that long in canis). It differs from
schmitti, norrisi and canis in the fact that its teeth are symmetrical and with evenly convex
cutting edges. The most striking differences between mento and fasciatus are its much
shorter head and a caudal that has a well defined lower lobe (cf . Fig, 42 H with 43 upper) .
At least some of the adults of mento resemble asterias of the eastern Atlantic and Mediter-
ranean in being conspicuously marked with many small white spots, but mento differs
from asterias in the features stated in the Key, p. 242.

Color. Adults are often conspicuously marked with many small white spots, but
sometimes they are plain colored,^' whereas young specimens may be marked with dark
colored bars.

Range. Coasts of Peru and Chile; perhaps Argentina.

Remarks. This species is included because a white-spotted Mustelus,'' said to be com-
mon in northern Argentina, seems more likely (on geographic grounds) to be mento than
the European asterias, under which name it was reported.

Synonyms and References:

Muite/us mento Cope, Proc. Amer. phil. Soc, 77, 1877: 47 (descr., Peru) Bigelow and Schroeder, Proc.

Boston Soc. Nat. Hist., 41, 1940: 429, Table column K, pi. 14, fig. C, pi. 16, fig. D, pi. 17, fig. I, pi. 19,

fig. C (discus.; meas.; ills, fins, teeth, denticles); Fowler, Feces Peru, Mus. Hist. Jav. Prado, 1945: 12

(listed, Callao, Peru).
Mustelus edulis Perez Canto, Estud. Escual. Chile, 1886: 4 (descr., Chile); Philippi, An. Univ. Chile, 77,

1887: 547, pi. 6, fig. 5 (descr., meas., color, ill., Chile).

38. If our reference of edulis Perez Canto, 1886, and abbotti Evermann and Radcliffe, 1917, to the synonymy of
mento be correct. For discussion, see Bigelow and Schroeder (Proc. Boston Soc. nat. Hist., 41 [s], 1940: 429).

39. Lahille, An. Mus. nac. B. Aires, 34, 1928: 310.

Fishes of the Western North Atlantic 261

Galeus mento, Quijada, Bol. Mus. nac. Chile, 5, 1913: 107 (listed for Chile).

Mustelus abbotti Evermann and Radcliffe, Bull. U.S. nat. Mus., 95, 1917: 6 (descr., meas., color, ill., Peru).

Possible South Atlantic References:

Mustelus asterias Lahille, Physis, B. Aires, 5, 1 921: 63 (name only, in list for Argentina) ; Enum. Peces Car-
tilag. Argent., 1921: 13 (brief account of teeth, Argentina); An. Mus. nac. B. Aires, 5^, 1928: 310
(Argentina, said to be common) ; Pozzi and Bordale, An. Soc. cient. argent., 120, 1935: 151 (Argentina,
Lat. 35°-42° S.; name only).

Mustelus schmitti Springer, 1939

Figures 42 F, Gj 43 C

Study Material. Four specimens, 542 to 742 mm., from Rio Grande do Sul, Brazil,
and from Uruguay (U.S. Nat. Mus. and Harv. Mus. Comp. Zool.).
Distinctive Characters. See following Description.

Description. Proportional dimensions in per cent of total length. Male, 571 mm.,
from Rio Grande do Sul, Brazil (Harv. Mus. Comp. Zool., No. 35316). Female, 596
mm., from Maldonado, Uruguay (Harv. Mus. Comp. Zool., No. 530).

Trunk at origin of -pectoral: breadth lO.0, 9.7; height 9.5, 8.4.

Snout length in front of: outer nostrils 4.6, 3.55 mouth 6.$, 5.4.

Eye: horizontal diameter 3.0, 2.5.

Mouth: breadth 5.6, 6.4; height 3.3, 2.8.

Nostrils: distance between inner ends 2.8, 2.2.

Labial furrow lengths: upper 1.9, 1.95 lower 1.2, 1.5.

Gill opening lengths: ist 2.1, 1.8; 2nd 2.3, 1.9; 3rd 2.3, 2.0; 4th 2.4, 2.0; 5th

2.0, 1.9.

First dorsal fin: vertical height 8.2, j length of base 12.6, 12.9.

Second dorsal fin: vertical height 5.8, 6.45 length of base 10.2, 10.9.

Anal fin: vertical height 3.5, 3.45 length of base 7.0, 7.4.

Caudal fin: upper margin 19. i, 19. 5j lower anterior margin 8.9, 9.1.

Pectoral fin: outer margin 14.9, 14.95 i'^^'' margin 8.1, 8.9; distal margin ii.i,

1 1.7.

Distance from snout to: ist dorsal 28.4, 28.55 2nd dorsal 61.5, 58.85 upper caudal

80.9, 80.55 pectoral 20.2, 16.85 pelvics, 44.6, 42.85 anal 65.7, 65.1.

Interspace between: ist and 2nd dorsals 21.9, 21.75 2nd dorsal and caudal 10.3,

1 1.75 anal and caudal 6.3, 6.1.

Distance from origin to origin of: pectoral and pelvics 24.2, 26.75 pelvics and anal

21.2,21.2.
M. schmitti so closely resembles canis in general appearance, in the size, shape, and
location of fins, and in the teeth and denticles, that the points of difference alone need be
mentioned.*" The most striking difference between schmitti and canis lies in the outline of

40. For discussion, see Bigelow and Schroeder (Free. Boston Soc. nat. Hist., ^i, 1940: 420).

262 Memoir Sears Foundation for Marine Research

the caudal fin; in schmitti this lacks any definitely outlined lower lobe, which (added to a
proportionately longer terminal sector) gives the fin an aspect quite different from that of
cams (Fig. 42). The distance between the nostrils is only about V2-V3 as great as the
breadth of the mouth in schmitu, but averaging more than one-half that great in canis. The
inner margin of the pectoral is longer relatively in schmitti (Fig. 43 C). The upper labial
furrow averages somewhat longer (see Proportional Dimensions) as does the interspace
between the pelvics and the anal. The denticles, which are otherwise similar, are so trans-
parent in schmitti that the pedicels are visible from the outside, which is seldom the case
in canis. The teeth are not distinguishable from those of canis (cf. Fig. 42 G with 42 D).

Color. Preserved specimens are mouse-gray above and of a pale shade of the same
tint below (much as in preserved canis) without any conspicuous markings.

Size. Seemingly this is a somewhat smaller species than canis, for males may mature
at a length of only 600 mm.

Develof mental Stages. Not known.

Habits. No information is available.

Range. So far known only from Rio Grande do Sul, in southern Brazil, from Uru-
guay (including Maldonado), and from Buenos Aires in northern Argentina.

Synonyms and References:

Mustelus schmitti Springer, Proc. U.S. nat. Mus., 86, 1939: 465 (descr. of type, meas., size at maturity);

Uruguay, and Buenos Aires, Argentina.
Mustelus scAmidti*^ Bigelow and Schroeder, Proc. Boston Soc. nat. Hist., 41 (8), 1940: 420, column I; pi.

14, fig. B, 15, fig. E, 17, fig. D (meas., discus., ill., of pectoral, caudal and teeth). For other doubtful

references, see under M. canis, p. 254.

Family CARCHARHINIDAE

Characters. Two dorsal fins, the ist much shorter than the caudal, its base wholly
anterior to origin of pelvics; caudal much less than V2 of total length, with well marked
subterminal notch, not lunate, but its lower anterior corner expanded as a definite lobe,
its axis raised but little; caudal peduncle not strongly flattened dorso-ventrally or widely
expanded laterally; precaudal pits more or less strongly marked; sides of trunk, anterior
to anal, without longitudinal ridges; inner margins of pelvics entirely separate, posterior
to cloaca, in both sexes; jaws not widely protrusible; 5th gill opening over or posterior to
origin of pectoral; gill arches without rakers, not interconnected by a sieve of modified
denticles; nostrils entirely separate from mouth, their anterior margins without barbel;
spiracles present or absent; eyes with well developed nictitating membrane within lower
lid, its upper edge continuous with edge of lower eyelid anteriorly, but enclosed far within
the latter posteriorly; labial furrows more or less developed, on one or both jaws; teeth
blade-like, with only i cusp, only i row functional along sides of jaws, or 2, depending on
the stage in their replacement; head of normal shape, not widely expanded laterally;

41. Misprint for "schmitti."

Fishes of the Western North Atlantic 263

rostral cartilages 3, united anteriorly^ radials of pectoral mostly on metapterygium;
meso- and propterygia much smaller} meso- and metapterygia separated by a foramen, at
least in some cases; heart valves in 3 rows. Development either ovoviviparous, or vivipa-
rous with well developed yolk-sac placenta.

Genera. This is not only the largest family of sharks, but the majority of modern
sharks fall within it. All its western Atlantic members are inhabitants of the tropical or
warm temperate belts, only entering the boreal zone in summer with the vernal expan-
sion of high water temperatures, if at all. Some of them are cosmopolitan in the appropriate
thermal belt, but others are confined to comparatively narrow areas of distribution. The
majority of species are harmless, but a few bear evil reputations as dangerous to bathers.

Key to Genera

la. Anal nearly 4 times as long at base as 2nd dorsal. PAyjoiow Miiller and Henle, 1841.

India, China, Australia.
lb. Anal at base less than 3 times as long as 2nd dorsal.
2a. Spiracles present, from large to minute.

3a. Second dorsal originates behind rear end of base of anal.

Loxoiow Muller and Henle, 1841.
Red Sea, Mauritius.
3b. Second dorsal originates over or in front of midpoint of base of anal.

4a. Midpoint of base of ist dorsal considerably nearer to origin of pelvics
than to axil of pectoral. Thalassorhmus Miiller and Henle, 1 841 .'

Eastern North Atlantic, Mediterranean.
4b. Midpoint of base of ist dorsal at least as near to axil of pectoral as to
origin of pelvics, or nearer.

5a. Caudal peduncle with a low longitudinal dermal ridge on each sidej
upper labial furrow as long as snout in front of mouth.

Galeocerdo Miiller and Henle, 1837, p. 265.

5b. Caudal peduncle without longitudinal dermal ridges; upper labial

furrow not more than ^^ as long as snout in front of mouth.

6a. Inner margins of upper teeth regularly serrate nearly to tips,

but without basal denticles; their bases very deeply incised in

the midline. HemiprisUs L. Agassiz, 1 843.

Red Sea.=

1. The genus seems not to have been reported since i 88 i, when Moreau (Hist. Nat. Poiss. France, / .- 319) described a
specimen from Cette on the French coast of the Mediterranean. Watch should be kept for it, for it is likely to be
confused with Prionace, from which it differs chiefly in having- spiracles.

2. We follow several previous authors in referring the Dhr/iizodon elongatus of Klutzinger (Verh. zool.-bot. Ges.
Wien, 11, 1871 : 664) to Hemipristis on the strength of Probst's (Wurt. Jahresh., 5.;, 1878: 141) statement that
the teeth of the single known specimen resemble very closely the fossil shark's teeth that have been described under
that name from the Upper Cretaceous to Miocene of North America, Upper Cretaceous to Pliocene of Europe,
Eocene and Miocene of Africa, Miocene of Asia and South America, and Tertiary of the West Indies.

264 Memoir Sears Foundation for Marine Research

6b. Inner margins of upper teeth either smooth, or with i to several
basal denticles (but not serrate) ; their bases not deeply incised
in the midline.

7a. Precaudal pits well developed, both- above and below.
8a. Lower teeth erect, smooth-edged all along jaw.

Negogaleus^ Whitley, 1931.

India, Philippines, Australia, East
Indies, Indo-China.

8b. Lower teeth along sides of jaw strongly oblique, their
outer margins notched and denticulate.

Paragaleus Budker, 1935, p. 275.
7b. No precaudal pit above or below.

Galeorhinus Blainville, 1 8 1 6.

Eastern Atlantic, including Mediter-
ranean ; southern Scandinavia to tropi-
cal West Africa; South Africa; Uru-
guay and Argentina; west coast of
America (Chile, Peru, Lower Cali-
fornia, California); Central Pacific;
Japan, Formosa, East Indies, Aus-
tralia, Tasmania, New Zealand.*

2b. Spiracles lacking.

9a. Midpoint of base of ist dorsal considerably nearer to origin of pel vies than
to axil of pectoral. Prtonace Cantor, 1849, P- ^^O-

9b. Midpoint of base of ist dorsal at least as near to axil of pectoral as to origin
of pelvics, or nearer.
lOa. Cusps of upper teeth smooth-edged, as well as those of lower.

1 1 a. Second dorsal at least % as long at base as ist, its posterior margin
deeply concave. Negafrion Whitley, 1939, p. 308.

3. Proposed by Whitley (Aust. Zool., 6, 1931 : 334) to replace Hemigaleus Bleeker, 1S52, the latter being preoccu-
pied by Jourdain, 1837, for mammals.

4. Recorded nominally from Argentina and from Uruguay as Galeorhinus galeus (Berg, An. Mus. nac. B. Aires,
[2] /, 1895: 7; Devincenzi, An. Mus. Hist. nat. Montevideo, [2] /, 1920: 119), and as Galeus cams (Lahille,
Physis B. Aires, 5, 1921: 63; Enum. Peces Cartilag. Argent., 1921: 13; An. Mus. nac. B. Aires, S4> '9^8:
310; Marini, Physis B. Aires, 10, 1929: 452; Pozzi and Bordale, An. Soc. cient. argent., 120, 1935: 150).

None of these citations include any description of the South American specimens. But the illustration of one
from Uruguay by Devincenzi and Barattini (An. Mus. Hist. nat. Montevideo, Suppl. Album Ictiol., 1926: pi. 1,
fig. 3) resembles the common Tope of Europe (G. galeus) so closely in general appearance, especially in the
very characteristic shape of the caudal, that the Uruguayan form must be regarded as identical with it, at least
until specimens can be compared critically. This appears to be true also of the Oil Shark of the eastern side of the
Pacific, described as Galeorhinus zyofterus Jordan and Gilbert, from California (Bull. U.S. nat. Mus., 16, 1883:
871) ; also as Galeus chiUnsis Perez Canto (Estud. Escual., Chile, 1886: 3), and as G. ?no/<'»<»f Philippi (An. Univ.
Chile, 7/, 1887: 544, pi. 4, fig. 1) from Chile.

That this species does not occur on the western side of the North Atlantic is one of the puzzling features in
the geographic distribution of sharks.

Fishes of the Western North Atlantic 265

lib. Second dorsal less than ^2 as long at base as ist and much smaller
in area, its posterior margin only weakly concave, or nearly
straight.

1 2a. Bases of upper teeth, as well as of lowers, smooth-edged.
13a. Teeth slender, symmetrical, erect in both jaws; long-
est gill opening nearly or quite V2 as long as base of
1st dorsal.' Aprionodon Gill, 1861, p. 303.

13b. Teeth in sides of jaws oblique, their outer edges
notched; longest gill opening only about Y^ as long as
base of ist dorsal.
14a. Teeth with swollen rounded bases.

Protozygaena Whitley, 1 940.

Australia.

14b. Teeth not swollen at base.

ScoUodon Muller and Henle, 1837, p. 292.
1 2b. Bases of upper teeth with serrate or denticulate edges.

Hypo-prion Muller and Henle, 1841, p. 315.
lOb. Margins of cusps of upper teeth regularly serrate; lowers either ser-
rate or smooth. Carcharhinus Blainville, 18 16, p. 320.

Genus Galeocerdo Muller and Henle, 1837

Galeocerio Muller and Henle, Arch. Naturg., (3) i, 1837: 397; type species, Squalus arcticus Faber, 1829,
Iceland, equals Squalus cuvier Lesueur, 1 822, Australia.

Generic Synonyms:

Squalus (in part) Lesueur, J. Acad. nat. Sci. Philad., 2, 1822: 351; Faber, Fische Islands, 1829: 17; not

Squalus Linnaeus, -1758.
GaUus L. Agassiz, Poiss. Foss., 5, 1835: pi. E, fig. 5, 6; Poiss. Foss., 5, 1838: 91 ; type species, G. cefedianus

L. Agassiz, East Indies; not Galeus Rafinesque, 18 10.
Prionodon (subgenus in part) Bleeker, Verh. bata"ia. Genoot., 24, Plagiost., 1852: 37; for P. jasciatus, Java;

not Prionodon Muller and Henle, 1 841.
Boreogaleus Gill, Ann. N. Y. Lye, 7, 1 861 : 402, 41 1 ; type species, Squalus arcticus Faber, 1829.
/f^ray Townsend, Bull. N. Y. zool. Soc, 5^ (6), 1931: 168, photograph of Galeocerdo mislabelled "Isurus

tigrinus," evidently by mistake.

Generic Characters. Anal only a little longer at base than 2nd dorsal; spiracles pres-
ent, small; 2nd dorsal originates over or a little in front of origin of anal; 2nd dorsal only
a little more than V2 as long at base as ist dorsal, and considerably less than Yo as large in
area; midpoint of base of ist dorsal only about Yz as far from axil of pectoral as from
origin of pelvics; caudal peduncle with a low longitudinal dermal ridge on each side;

5. The gill openings are as long as this in the most recent illustration of A. brevipinna Muller and Henle, 1841
(Whitley, Fish. Aust., /, 1940: 108, as "Longmania brevi-pinna"), although pictured as considerably shorter in
the original illustration of that species (Muller and Henle, Plagiost., 1 84 1 : pi. 9) .

266 Memoir Sears Foundation for Marine Research

upper labial furrow about as long as snout in front of mouth; a well marked precaudal
pit below as well as above; a low dermal ridge along midline of back between dorsal fins;
caudal with pointed tip and lower lobe; teeth alike in the 2 jaws, large, few in number,
with coarsely serrate edges, convex inwardly, their outer margins deeply notched; longest
gill opening about Ys as long as base of i st dorsal, the 4th over origin of pectoral. Develop-
ment ovoviviparous. Characters otherwise those of the family.

Range. Cosmopolitan in tropical and subtropical seas.

Fossil Teeth. Upper Cretaceous to Miocene, North America; Eocene to Miocene,
Africa; Eocene to Pliocene, Europe; Miocene, Asia, South America, West Indies.

S-pecies. It is probable that all published references to the genus, from all parts of
the world, concern one or another race of a single wide-ranging species, the common
Tiger Shark of tropical seas; although the Australasian Galeocerdo has been considered
a distinct species by some authors,* there is nothing in the published accounts to suggest
any clear distinction between it and the Galeocerdo of the tropical Atlantic.

Galeocerdo cuv'ter (Lesueur), 1822

Tiger Shark, Leopard Shark

Figure 44

Study Material. Two young females, 1,368 and 1,380 mm. (about 4 feet 6 inches)
long, and a young male of 1,245 m"^- (about 4 feet i inch), from southern New England;
very small female, 585 mm. long (about 23 inches), with well marked umbilical scar,
hence either late embryo or newborn, from Cuba; also jaws of larger specimens from vari-
ous localities.

Distinctive Characters. There is no danger of confusing the "Tiger" with any other
shark, so diagnostic are its teeth, combined with the very short snout, very long upper
labial furrows and sharp-pointed tail.

Description. Proportional dimensions in per cent of total length. Male, 1,245 n^m-j
from Rhode Island (Harv. Mus. Comp. Zool., No. 35145).

Trunk at origin of pectoral: breadth lO. i ; height 10.9.

Snout length in front of: outer nostrils 1.9; mouth 4.2.

Eye: horizontal diameter 2.1.

Mouth: breadth 8.4; height 5.0.

Nostrils: distance between inner ends 4.3.

Labial furrow lengths: upper 4.3, lower 1.9.

Gill opening lengths: ist 2.4; 2nd 2.5; 3rd 2.5; 4th 2.6; 5th 2.3.

First dorsal fin: vertical height 8.3 ; length of base 8.4.

6. Most recently by Whitley (Fish. Aust., /, 1940: 113) as G. rayneri MacDonald and Barron, 1868. If the Aus-
tralian form should finally prove to be distinct from the Atlantic, its correct name is cuvier Lesueur, 1822, type
locality "New Holland," the name then in use for eastern Australia.

Fishes of the Western North Atlantic

267

Second dorsal fin: vertical height 2.75 length of base 4.7.

Anal fin: vertical height 3.45 length of base 5.0.

Caudal fin: upper margin 30.2; lower anterior margin 12.6.

Pectoral fin: outer margin 14.2; inner margin 5.4; distal margin 12.1.

Distance from snout to: ist dorsal 26.2; 2nd dorsal 56.85 upper caudal 69.7;

pectoral 19.85 pelvics 44.1 ; anal 57.5.

Interspace between: ist and 2nd dorsals 23.O5 2nd dorsal and caudal 9.65 anal and

caudal 7.3.

Distance from origin to origin of: pectoral and pelvics 26.05 pelvics and anal 1 2.4.

Figure 44. Galeocerdo cuvier, young male, 1,245 nim. long, from Newport, Rhode Island (Harv. Mus. Comp.
Zool., No. 35145). A Anterior part of head of same. B Cross section of upper part of trunk opposite origin of
pelvics showing mid-dorsal ridge. C Cross section of caudal peduncle to show lateral ridges. D General view
of dermal denticles, about 28 x; lateral and apical views, about 56 x. E Teeth of larger specimen (U.S. Nat.
Mus., No. 1 10900), about 0.2 natural size. F Third lower tooth, enlarged. G Fifth upper tooth of same, en-
larged.

Trunk stoutest opposite ist dorsal and tapering evenly rearward, the midline of
back with a low dermal ridge extending rearward from a short way anterior to rear tip of
1st dorsal about % of distance to 2nd dorsal, where it gives place to a narrow, ill-defined
furrow that reaches to the latter.' Caudal peduncle slender, oval in cross-section, with a low
rounded ridge along each side at the midlevel, from opposite rear end of 2nd dorsal to a
little beyond origin of caudal. Precaudal pits in the form of obtusely subangular furrows,
the upper considerably the more distinct and larger. Dermal denticles large, variously

7. The combination of ridge and furrow in this region appears to be unique.

268 Memoir Sears Foundation for Marine Research

spaced but usually not overlapping, their blades nearly horizontal, longer than broad,
usually with 3 ridges, the median very high and divided anteriorly, the lateral margins up-
turned, the posterior margin with a short, broad median tooth, usually flanked by a pair of
very small teeth, but sometimes by one only; pedicels very short; basal plates very broad,
distinctly 4-rayed.

Head flattened above, nearly or quite as broad opposite front of mouth as at origin of
pectorals. Snout very broadly rounded, noticeably short, its length in front of mouth a little
less than % of length of head to origin of pectorals. Eye broadly oval, its horizontal
diameter about V2 as long as snout in front of mouth, its center a little anterior to mid-
height of mouth. Spiracle a narrow but easily visible slit, Vs to Yi as long as horizontal
diameter of eye and behind the latter by a distance a little greater than Yo the diameter
of eye. Gill openings about evenly spaced, the 2nd, 3rd, and 4th (slightly the longest)
a little more than V2 as long as snout in front of mouth, or about 1.2 times as long as diame-
ter of eye, the ist and 5th the shortest, the 4th over origin of pectoral. Nostril nearly trans-
verse, its inner corner about equidistant between tip of snout and front of mouth. Its ante-
rior margin expanded at the Inner end as a low triangular lobe with rounded apex. Mouth
broadly ovate, a little more than V2 as high as broad, occupying between % and % of
breadth of head, lateral (not inferior) for most of Its length. Upper labial furrow as
long as snout in front of mouth or a little longer, extending forward to a point about oppo-
site anterior edge of eye. Lower labial furrow, a little less than Yo as long as upper,
approximately parallel to lower jaw.

Teeth ^ to Ij-EJEif ii^ specimens examined; very large In front and sides of jaws
(up to an Inch or more high In large specimens) but decreasing In size toward corners
with the outermost very small, similar in the 2 jaws, about V2 to % as high as broad,
their inner outlines convex, their tips directed obliquely outward, their outer margins with
a deep primary notch; both edges serrated, very finely so near the tip which may be worn
smooth, but more coarsely so toward the base, especially on the outer margin basal to the
notch, where the primary serrations are themselves finely serrate secondarily along their
edges. There may or may not be a small symmetrical tooth at the symphysis of either jaw
(Fig. 44 E) ; If this Is lacking the tooth next to the symphysis In the upper jaw Is usually
considerably smaller than the 2nd and subsequent teeth, on either one side or on both,
though similar to them in shape.

Anterior margin of ist dorsal about Y2 as long as from snout to axil of pectoral, Its
origin over or a little posterior to the latter, Its anterior edge very slightly convex, its
posterior margin deeply concave, its apex narrowly rounded, the free rear tip slender,
about % as long as base. Second dorsal about V2 as long as ist at base, but a little less than
Ys as high vertically, its anterior edge more sloping. Its free rear tip more narrowly
acuminate and relatively longer (about as long as base). Its origin a little anterior to origin
of anal. Caudal a little less than Ys of total length, with very slender pointed tip and deep
subterminal notch, its lower anterior corner expanded as a narrow sharp-pointed lobe.

Fishes of the Western North Atlantic 269

a little more than Vs as long on its anterior margin as the upper lobe. Anal about as long
as 2nd dorsal at base, and slightly higher vertically, but with posterior margin much more
deeply concave and free rear tip a little shorter relatively, its rear tip a little posterior to
rear tip of 2nd dorsal. Pelvics with nearly straight edges and narrowly rounded corners.
Pectoral about M; ^s long as head, or a little longer than ist dorsal and a little larger in
area, about Vi; ^s broad as long, with moderately convex outer margin, moderately and
evenly concave inner margin, and narrowly rounded corners.

Color. Gray or grayish brown, darker above than on sides and belly; small specimens
up to about five or six feet long are more or less prominently marked on back with darker
brown spots, often fusing irregularly into oblique or transverse bars on the sides and fins,
sometimes surrounded with pale reticulations; but these markings fade with growth,
leaving the larger specimens only faintly marked on the caudal peduncle, or even plain-
colored in some cases.

Size. This is one of the sharks to which a gigantic size (up to 30 feet in length) has
been accredited. However, the majority of specimens that are taken in its centers of
abundance are less than 12 to 13 feet long.'" The longest of which we find positive record
within recent years in the western Atlantic have been a Cuban specimen of about 1 8 feet,*
and one of 1 5 feet 2 inches from South Carolina.^ The weight at different lengths varies
with fatness, and with the stage of development of the embryos in gravid females. Aus-
tralian specimens are reported as weighing 7 10 to 825 pounds at 1 1 to 12 feet, 850 to 1,324
pounds at 12 to 13 feet, and 1,028 to 1,395 pounds at 13 to 14 feet;" recorded weights
from the Pacific coast of Central America are 37 pounds at 5 feet 4 inches (1,625 mm.),
366 pounds at 10 feet i inch (3,073 mm.), 505 pounds at 10 feet 6 inches (3,200 mm.),
and 780 pounds at 12 feet 9 inches. One 1,368 mm. long from Woods Hole, Mass.,
weighed 25% pounds, fresh. And there is no reason to suppose that the weights of larger
Atlantic specimens would be different at equal lengths from Pacific examples, although
they have been previously estimated as somewhat less.^^

Although they may grow very large, Tiger Sharks are comparatively small at birth
corresponding to the large numbers produced at one time, free-living specimens having
been recorded as small as 1 8 to 19 inches.

Developmental Stages. Development is ovo viviparous; the embryos have no placen-
tal connection with the mother. The broods are very large, gravid females having been re-
ported repeatedly as containing as many as 30 to 50 embryos, some more nearly ready for
birth than others; recently we have received an account of an 18-foot Cuban specimen

7a. Stewart Springer informs us that none of the many measured by him in Florida waters were as long as 14 feet.

8. Personal communication from Luis Howell-Rivero.

9. Burton, Copeia, 1941:40. 10. Whitley, Fish. Aust., /, 1940: 113.

II. 450 to 636 pounds at 11 to 12 feet (Bell and Nichols, Copeia, 92, 1921: 17; Nichols and Breder, Zoologica,
N.Y., 9, 1927: 15). The following weights are also mentioned, without locality; 58.8 pounds at 5 feet 2 inches,
168.4 pounds at 6 feet, 718.3 pounds at lo feet 8 inches (Schultz, J. Mammal., tg, 1938: 484).

270 Memoir Sears Foundation for Marine Research

containing 82 young.^° On the other hand, litters as small as lO to 14 have been recorded."
Habits. The "Tiger" is found indiflFerently far out on the high seas and in coastwise
waters. In tropical and subtropical seas they have been seen pursuing sting-rays on the
flats in only a few feet of water and even in harbors; they are caught from the shore; and
it is not unusual for them to enter enclosed sounds and river mouths in Florida and North
Carolina. Most of the few records of them further north are from pound nets set out
from the land in a few fathoms of water only. Except when aroused by the scent of food or
other stimuli, the "Tiger" is rather sluggish ; when stimulated, however, it is one of the
most vigorous and strong swimming of sharks. In Florida waters, and presumably
throughout its normal range, its young may be born at any time of year.

Although this is perhaps the commonest large shark in the tropics, little more is
known about its life history, except for its diet, it being proverbially one of the most
voracious of sharks. It is also one of the most omnivorous, for its diet ranges from objects
as small as crabs and the smaller migrating land birds that have fallen into the sea to
others as big as the larger sea turtles, other sharks, and sea lions." "The large, coarsely
serrated teeth are extremely efficient cutting instruments. . . . Bites on large objects are
made by a rolling motion with both jaws cutting much in the manner of a saw";"^ and a
Tiger Shark has no difficulty in cutting through the shell of a sea turtle. The recorded list
of its stomach contents includes crabs (half a bushel of them were taken from a 13-foot
specimen in Florida), gastropods (Buccinum, Lunatia), spiny lobsters (Palinurus), horse-
shoe crabs (Limulus), squid, a wide variety of fishes, among them sharks smaller than
themselves (a case in point is a specimen taken off Morehead City, North Carolina, which
contained a Carcharhinus limbatus) , skates, and even sting-rays, which they devour regard
less of the poisonous spines, these often being imbedded in their jaws or elsewhere in their
bodies. It is a common habit of this species to bite great chunks from other sharks, often of
its own kind or of any other species which may be entangled in nets. The stomach contents
of 34 specimens, netted off North Carolina, contained crabs, Limulus, sharks (small and
large, entire and in pieces), large amounts of mackerel and unidentified small fish, sea
turtles (entire and in pieces), bones and feathers of sea fowl, pieces of shark and porpoise
that had seemingly been bitten from the nets, and garbage (sheep-bones, etc.)." As further
evidence of its voracity we may quote an instance in which a large one, rearing head
out of water, tore out the throat and belly from another shark that had been hoisted up to a
boom." The "Tiger" is also known as a scavenger, feeding on any kind of carrion, for
example, parts of sheep, dead dogs, beef bones, remains of poultry, and even on such
unappetizing objects as lumps of coal, tin cans or empty sacks. There is a recorded case in

12. Personal communication from Luis Howell-Rivero.

13. Whitley, Fish. Aust., /, 1940: 1 13. For an account of early embryonic development, see Sarangdhar (J. Bombay
nat. Hist. Soc, ^^ [i], 1943: 105).

14. See Beebe (Galapagos Worlds End, 1924: 201) for an eye-witness account of a Tiger Shark devouring a young
sea lion.

14a. Springer, Proc. Fla. Acad. Sci., J, 1939: 16. 15. Bell and Nichols, Copeia, 92, 1921 : 18-19.

16. Radcliffe, Bull. U.S. Bur. Fish., 34, 1916: 263.

Fishes of the Western North Atlantic 271

Australia in which one, after capture, vomited the entire arm of a man who had been mur-
dered at sea and his body dismembered." "Tigers" also join the company of various other
sharks that are soon attracted to the carcasses of dead horses or cattle in tropical harbors
in the vicinity of slaughter houses.

Relation to Man. The "Tiger" is of considerable commercial value wherever a
shark fishery is actively carried on in warm seas, as in southern Florida at present and until
recently among the Virgin Islands, for it not only probably forms the largest single item in
the catches, but yields excellent leather which is used for many purposes. Its yield of liver
oil is also higher than that of many other tropical sharks. It likewise affords some sport to
anglers, for it bites readily, provided the bait is large and strong-smelling. On the other
hand, "these sharks are very destructive to gill nets, biting out great holes to take a single
fish, and swimming back and forth through the nets as they feed on the gilled fish."'*
Worse yet, Tiger Sharks, when they come into shallow water, may be a danger to
bathers; in the West Indies they are said to be considered the most dangerous of sharks.
Some of the many shark fatalities that are well attested in medical journals for Australian
waters are also credited to this species, although perhaps not on conclusive evidence. A
recent instance is recorded from Malwan, south of Bombay, India."

Range. Widespread in the tropical and subtropical belts of all the oceans, inshore and
offshore alike.

Occurrence in the Eastern Atlantic. In the Eastern Atlantic, positive records for the
Tiger Shark are comparatively few in number, i.e., for the Canaries, tropical West Africa
(Senegambia), western South Africa, and accidentally for Iceland.^"

Early writers repeatedly credited it to northern Scandinavian waters and to the
Faroes, an error springing from the fact that Faber's account of his "arcticus'' was based on
a combination of the latter with Isurus nasus, the common Porbeagle of boreal waters.
Actually there is no positive record of the Tiger Shark for North Europe, other than the
one for Iceland. It has never been reported from the Mediterranean, but no doubt it is
much more plentiful along the tropical coast of West Africa and around the off-lying
islands than the paucity of published records would suggest.

Occurrence in the Western Atlantic. This is one of the more numerous, if not the most
abundant, of the larger sharks in the appropriate thermal zone of the western Atlantic.
As with various other tropical species, its center of abundance appears to be the Carib-
bean-West Indian-South Florida region. Among the West Indies there is a published
record of it at Trinidad, Porto Rico, the Virgin Islands, Cuba, between Turks Island and
the Barbados, and near Nassau in the Bahamas, where it is so plentiful that 3 1 "Tigers"

17. For account of this happening and the subsequent investigations, see Whitley (Fish. Aust., /, 1940: 34).
x8. Springer, Proc. Fla. Acad. Sci., 5, 1939: 16.

19. Sarangdhar, J. Bombay nat. Hist. See, 44 (i), 1943: i04.

20. The identity of this specimen is attested by the account of its teeth by Faber (Fische Islands, 1829: 17) and more
recently by Kr0yer (Danmarks Fiske, 3, 1852-1853: 933).

272 Memoir Sears Foundation for Marine Research

up to 16 feet in length have recently been reported among one catch of 51 sharks of all
kinds."' No doubt it is equally common among the Antilles generally, and around Cuba.
In southern Florida waters it is present among the Keys and on both the Atlantic and the
Gulf of Mexico coasts throughout the year. Curiously enough, we have found no records
of it for the Atlantic coasts of Central America and only one vague report for the inner
Gulf of Mexico. But it has been encountered recently in July in the northern side of the
Gulf off Biloxi, Mississippi.'" And the poverty of the printed record, rather than any local
scarcity, probably explains the lack of reports of it along Central America.

The Tiger Shark is only a summer visitor to the Atlantic coast of the United States
north of Florida. Although there is only one definite record of it for South Carolina,
considerable numbers must pass by there, at least in some years, for they have been reported
repeatedly along North Carolina, sometimes in schools, even entering the enclosed
sounds and river mouths at times. Only odd specimens have been reported from the sector
thence northward past New York, i.e., in Chesapeake Bay (once), Delaware Bay (once),
New Jersey (about four times), Long Island, New York (once), and Newport, Rhode
Island (once. Fig. 44). But like many other tropical fishes, Tiger Sharks appear more
often in the Woods Hole region, where one to three are taken in the pound nets almost
every summer, more often small but sometimes large. However, this is the northeastern
limit to their occurrence inshore,^' though odd specimens may be expected to stray much
farther in this direction offshore in the tropical waters of the Gulf Stream j the often quoted
Icelandic specimen may well have journeyed by that route.

To the southward the Tiger Shark is known from southern Brazil and Uruguay.
Probably it occurs commonly all along the northeastern and northern coasts of South
America, although it is not yet recorded there in scientific literature. It is also taken or
seen from time to time around Bermuda.

Synonyms and References:

I. Atlantic:

Cants carcharias^* Duhamel, Traite Gen. Peches, 4 (2) Sect. 9, 1782: 297 (in part), pi. 19, fig. 3 (teeth, not

fig- 1-3)-
Squale (no spec, name) Lacepede, Hist. Nat. Poiss., 4° ed., i, 1798: pi. 8, fig. 2, in Buffon, Hist. Nat. (jaws).
S^ua/us cuvier Le^near, ]. Acad. nat. Sci. Philad., 2, 1822: 351 (Aust.).
Carchorhinus lamia Blainville, in Vieillot, Faune Franc, 1825: 88 (teeth, ident. by ref. to Duhamel, Traite

Gen. Peches, 4 [2] Sect. 9, 1782: 298, pi. 19).
Squalus arctiais Faber, Fisches Islands, 1829: 17 (teeth, Iceland, confused with Isurus nasus) ; Nilsson, Prod.

Ichthyol. Skand., 1832: 115 (Iceland).
Galeus (ho spec, name) Agassiz, L., Poiss. Foss., j, 1835: pi. E, fig. 5, 6 (teeth); Owen, Odontogr., 1840-

1845: pi. 28, fig. 9 (teeth; shows a sting-ray's spine imbedded in jaw).
Galeus maculatus Ranzani, Nov. Comment. Acad. Sci. Inst. Bonon., 1840: 7, pi. I (descr., Brazil).
Galeocerdo arcticus Miiller and Henle, Arch. Naturg., (3) i, 1837: 398 (name); Plagiost., 1841: 60,

pi. 64 (descr., distrib. probably confused with that of Lamna nasus) ; Bonaparte, Mem. See. neu-

21. Wise, Nat. Hist. N. Y., 38, 1936: 311. 22. Personal communication from Stewart Springer.

23. Doubtfully reported from Provincetown at the tip of Cape Cod.

24. Duhamel's names, if binomial, are only accidentally so.

Fishes of the Western North Atlantic 273

chatel. Sci. nat., 2 (8), 1839: lO (in synopsis); Cat. Pesc. Europ., 1846: 19 (Boreal Ocean); Gray,
List Fish. Brit. Mus., 1851 : 54 (north, seas) ; Kr0yer, Danmarks Fiske, 3, 1 852: 933 (Iceland) ; Nilsson,
Skand. Fauna, 4, Fisk., 1855: 717 (ref.) ; Dumeril, Hist. N.it. Poiss., i, 1865: 394 (descr., spec, with-
out loc.) ; Gunther, Cat. Fish. Brit. Mus., 8, 1870: 377 (descr., "Arctic seas" by ref.); Hasse, Naturl.
Syst. Elasm. besond. Theil, 1882: 259, pi. 37, fig. 19-24, 27 (vertebrae) ; Gunther, Encj'cl. Brit., 11th
ed., 24, 191 1 : 807 (general); Garman, Mem. Harv. Mus. comp. Zool., j(5, 1913: 148 (descr.); Rad-
cliffe. Trans. Amer. Fish. Soc, 1914: 39 (N. Carolina) ; Fowler, Proc. Acad. nat. Sci. Philad., 6y, 1916:
521 (Trinidad, W. I.); Fowler, Copeia, 30, 1916: 36 (mid-Atlant. coast U.S.); Radcliffe, Bull. U.S.
Bur. Fish., ^4, 1916: 261, pi. 42, fig. i, 2 (teeth, denticles, N. Cirolina) ; Smith, J. Amer. Mus. nat.
Hist., 16, 1916: 348 (Atlant. coast U.S.) ; Meek and Hildebrand, Field Mus. Publ., Zool., 15, 1923: 56
(general); Bigelow and Welsh, Bull. U.S. Bur. Fish., 40 (i), 1925: 27 (New England); Fowler, Proc.
Acad. nat. Sci. Philad., So, 1929: 608 (C. May, N. Jersey) ; Breder, Field Bk. Mar. Fish. Atlant. Coast,
1929: 15 (general); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 14
(distrib.) ; Bellon and Mateu, Notas Inst. esp. Oceanogr., (2) 55, 1931: 17 (descr., meas., teeth,
Canaries); Beebe and Tee-Van, Zoologica, N. Y., 13, 1932: II9 (Bermuda); Wilson, Bull. U.S.
nat. Mus., 158, 1932: 454, 464 (parasites, Woods Hole); Beebe and Tee-Van, Shore Fish. Bermuda,
1933: 25 (Bermuda); Gowanloch, Bull. La. Conserv. Dept., 23, 1933^ 222 (Gulf of Mexico); von
Bonde, J. comp. Neurol., 193 3: pi. 2, fig. 2, 3 (brain) ; Bigelow and Schroeder, Canad. Atlant. Fauna, bid.
Bd. Canad., 12', 1934: 9 (Woods Hole region); Budker, Bull. Mus. Hist. nat. Paris, (2) 7, 193;: 186
(Dakar, W. Afr.) ; Marchand, Mar. biol. Rep. Cape Town, 2, 1935: 38 (west. S. Afr., Natal) ; Fowler,
Bull. Amer. Mus. nat. Hist., yo (1), 1936: 56 (W. Afr.) ; Cadenat, Rev. des. Trav. Peches Marit., 10
(4), 1937: 429 (off Dakar); Norman and Eraser, Giant Fishes, 1937: 42 (general); White, Bull.
Amer. Mus. nat. Hist., 7^, 1937: 124 (in Key) ; Nigrelli, Amer. Mus. Novit., 996, 1938: 10 (parasites) ;
Devincenzi, Ann. Mus. Hist. nat. Montevideo, (2) 4, 1939: 3 (Uruguay); Springer, Proc. Fla. Acad.
Sci., 3, 1939: 16 (Florida); Hildebrand, Copeia, 1941: 221 (N. Carolina); Norris, Plagiost. Hypophy-
sis, 1941: pi. I, fig. I (brain) ; Fowler, Arqu. Zool. Estado Sao Paulo, 3, 1942: 129 (Brazil) ; Fish Cul-
turist, 21 (9), 1942: 66 (sizes, Cuba); Lunz, Bull. S. C. St. Planning Bd., 14, 1944: 26 (S. Carolina,
Florida); Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1945: 263 (Long Key, Florida).

Squalus carcharias Blake, Dent. Format Struct., Baltimore, 1848: tab. 6, fig. 5 (teeth, not seen) ; not S. corcharias
Linnaeus, 1758.

Boreogaleus arcticus Gill, Ann. N. Y. Lye, 7, 1861 : 402, 41 1 (name).

GaUocerdo tigrinus Gill, Proc. Acad. nat. Sci. Philad., 1864: 263 (jaws, east coast U.S., W. Indies, Calif.,
W. Pacif.); Gunther, Cat. Fish. Brit. Mus., 8, 1870: 378 (descr., Japan, E. Indies, Atlant.); Verrill,
Rep. U.S. Comm. Fish. (1871-72), 1873: 521 (Massachusetts, food); Lutken, Vidensk. Medd. naturh.
Foren. Kbh., 1875: 37 (with a shark sucker, Echeneis remora) ; Goode, Proc. U.S. nat. Mus., 2, 1879:
121 (Florida); Hasse, Naturl. Syst. Elasm. besond. Theil., 1882: 259; Rochebrune, Act. Soc. linn. Bor-
deaux, (4) 6, 1882: 43; Faune Senegambie, Poiss., r, 1883-1885: 20 (Senegambia) ; Jordan and Gil-
bert, Bull. U.S. nat. Mus., 16, 1883: 21 (C. Cod to Indian Oc.) ; Nelson, Rep. St. Geol. N.J., 2 (2),
l8go: 660 (N. Jersey); Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 215 (distrib.);
Bull. U.S. nat. Mus., 47 (i), 1896: 32 (descr., distrib.); Linton, Proc. U.S. nat. Mus., 20, 1897: 424,
448 (parasites. Woods Hole) ; Smith, Bull. U.S. Fish Comm., 77, 1898: 88 (Woods Hole region) ; Ever-
mann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 48 (St. John's and Indian Rivers, Florida);
Bean, Rep. For. Comm. N. Y., 1901: 377 (Vineyard Sound); Linton, Bull. U.S. Bur. Fish., rp, 1901:
270, 425 (food. Woods Hole) ; Bean, Bull. N. Y. St. Mus., do, Zool. p, 1903: 24 (Martha's Vineyard,
size); Fowler, Science, N. S. 24, 1906: 596 (N. Jersey); Proc. Acad. nat. Sci. Philad., 5*, 1906: 79
(Florida Keys) ; Rep. N. J. Mus. (1906), 1907: 255, pi. 71 (ill., N. Jersey records) ; Sullivan, Bull. U.S.
Bur. Fish., 27, 1908: 14 (digestion); Kendall, Occ. Pap. Boston Soc. Nat. Hist., 7 (8), 1908: 3 (New
Engl. Iocs.); Gudger, Science, N.S. 57, 1 91 3: 993 (Key West, Florida) ; Yearb. Carneg. Instn., 12, 191 3:
177 (Tortugas, Florida); Sumner, Osbum and Cole, Bull. U.S. Bur. Fish., 31 (2), 1913: 735 (Woods
Hole region); Coles, Proc. biol. Soc. Wash., 28, 191 5: 89 (N. Carolina); Gudger, Science, N. S. 41,
1915: 437 (Key West, Florida); Yearb. Carneg. Instn., jj, 1915: 203 (Tortugas, Florida); Nichols,
Bull. Amer. Mus. nat. Hist., j^, 1915: 141 (Porto Rico) ; Wiegmann and Nichols, Copeia, 23, 1915: 43

2 74 Memoir Sears Foundation for Marine Research

(Long Island, N. York) ; Gudger, Yearb. Cameg. Instn., 14, 1916: 207 (Tortugas, Florida) ; Nichols and
Murphy, Brooklyn Mus. Sci. Bull., j (i), 1916: 8 (Long Island, N. York, Woods Hole region, food);
Coles, Copeia, 69, 1919: 38 (size, meas., food, C. Lookout, N. Carolina) ; Metzelaar, Trop. Atlant. Vis-
schcn, 1919: 188 (trop. W. Afr.) ; Bell and Nichols, Copeia, 92, 1921: 18 (N. Carolina, size, food);
Nichols, Nat. Hist. N. Y., 21, 1921 : 273 (food) ; Gudger, Nat. Hist. N. Y., 22, 1922: 245 (ref.) ; Nich-
ols and Breder, Zoologica, N. Y., 9, 1927: 14 (general) ; Jordan, Manual Vert. Anim. NE. U.S., 1929: 9
(general) ; Truitt, Bean and Fowler, Bull. Md. Conserv. Dept., 3, 1929: 28 (Chesapeake Bay) ; Nichols,
Sci. Surv. Porto Rico, N. Y. Acad. Sci., zo (2), 1929: 182 (food, Porto Rico) ; Gudger, Sci. Mon. N. Y.,
34, 1932:411 (sting-rays eaten) ; Young and Mazet, "Shark, Shark," 1933:268 (general) ; Wise, Tigers
of the Sea, 1937:175 (general) ; Gudger, Bull. Amer. Mus. nat. Hist., 75, 1937: 273 (tooth abnormali-
ties) ; Burton, Sci. Mon. N. Y., 40, 1935: 283 (S. Carolina) ; Devincenzi, An. Mus. Hist. nat. Monte-
video, (2) 4, 1939: 3 (jaws, Uruguay); Longley and Hildebrand, Pap. Tortugas Lab., ^4, 1942: 2
(color, Tortugas, Florida),

Galeocerio maculatus Poey, Repert. Fisico-Nat. Cuba, 2, 1868: 453 (Cuba) ; An. Soc. esp. Hist, nat., 5, 1876:
397, pi. 14, fig. 7; Enumerat. Pise. Cubens., 1876: 201, pi. 9, fig. 7 (teeth, Cuba); Jordan, Rep. U.S.
Comm. Fish. (1885), 1887: 795 (distrib.) ; Henshall, Bull. U.S. Fish Comm., 9, 1891: 383 (Florida);
Linton, Bull. U.S. Fish Comm., 15, 1894: 103 (parasite) ; Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907;
149, 199, pi. 2 (Brazil, refs.) ; Fauna brasil. Peixes, 2 (i) Fasc. I, 1923: 7, pi. I (Brazil).

Galeus tigrinus Fowler, Rep. N. J. Mus. (1905), 1 906: 61 (N. Jersey).

Isurus tigrinus Townsend, Bull. N. Y. zool. See, 34, 1931 : 168 (photo, Porto Rico).

Galeocerdo cuvier Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash.,
1945: 118 (descr., ill., range).

2. West Coast of North and South America:

Galeocerdo tigrinus Gilbert, Bull. U.S. Fish Comm., 2, 1883: 112 (Pacif. coast Guatemala); Jordan and
Gilbert, Bull. U.S. Fish Comm., 2, 1883: 105 (Mazatlan, Mexico) ; Jordan, Stanf. Univ. Pub., /, 1895:
382 (W. coast Mexico and Guatemala) ; Pellegrin, Bull. Mus. Hist. nat. Paris, 7, 1901 : 161, 166 (dan-
ger to man. Gulf of Calif.) ; Gilbert and Starks, Mem. Calif. Acad. Sci., 4, 1904: 9 (Pacif., Panama) ;
Snodgrass and Heller, Proc. Wash. Acad. Sci., 6, 1905: 342 (Galapagos) ; Fowler, Proc. Acad. Nat. Sci.
Philad., 60, 1908: 61 (Gulf of Calif.).

Galeocerdo arcticus Meek and Hildebrand, Field Mus. Publ. Zool., 75 (l), 1923: 56 (Pacif. coast, Panama) ;
Beebe, Arcturus Adv., 1926: 247, 435 (Cocos Is., not seen) ; Breder, Bull. Bingham Oceanogr. CoU., 2
(l), 1928: 3 (W. coast Cent. Amer.) ; Walford, Fish Bull. Sacramento, 4$, 1935: 35 (Calif.) ; Barnhart,
Mar. Fishes south. Calif., 1936: 9 (off Calif.); Schmitt, List Fish. President. Cruise, 1938: privately
printed (Cocos, Clipperton, Galapagos Is.); Beebe and Tee-Van, Zoologica, N. Y., 2(5 (2), 1941: 113
(food, coast Central Amer., Galapagos, Cocos I.).

Galeocerdo maculatus Jordan and Bollman, Proc. U.S. nat. Mus., 12, 1889: 179 (Panama).

Galeocerdo royneri Philippi, An. Univ. Chile, log, 1 901: 304 (Ecuador).

3. Central and Western Pacific, Australasian Region, Indian Ocean:
Squalus cuvier Lesueur, J. Acad. nat. Sci. Philad., 2, 1822: 351 (descr., Aust.).

Galeus cefedianus L. Agassiz, Poiss. Foss., 5, 1835: pi. E, fig. 5, 6; 1838: 91 (teeth, East Indies).

Galeocerdo tigrinus Miiller and Henle, Plagiost., 1841: 59, pi. 23 (small size, India) ; Gray, List. Fish. Brit.
Mus., 1851: 54 (Indian seas) ; Bleeker, Verh. batavia. Genoot., 25, 1853: 80 (ref.) ; Blyth, J. Asiat. Soc.
Beng., 29, i860: 36 (India) ; Gill, Ann. N. Y. Lye, 7, 1862: 402 (class.) ; Proc. Acad. nat. Sci. Philad.,
1864: 263 (jaws, E. coast. U.S., West Indies, Calif., W. Pacific); Dumeril, Hist. Nat. Poiss., /, 1865:
393 (descr., India); Gunther, Cat. Fish. Brit. Mus., 8, 1870: 378 (descr., Japan, E. Indies, Atlant.);
Klunzinger, Verh. zool.-bot. Ges. Wien, 21, 1871: 663 (Red Sea); Day, Fish. India, 1878: 718 (Red
Sea, India, Japan) ; Brusina, Glasnik Naravosi Drutzva, j, 1888: 224 (Red Sea) ; Ogilby, Cat. Fish. Aust.
Mus., I, 1888: 3 (Solomon Is.) ; Day, in Blandford, Fauna Brit. India, i, 1889: 21 (India) ; Boulenger,
Proc. zool. Soc. Lond., 1889: 243 (Muscat, Arabia); Sauvage, in Grandidier, Hist. Phys. Nat. Mada-
gascar, Poiss., 1 891 : 510 (ref.) ; Jordan and Snyder, Annot. zool. Jap., 3, 1 901 : 39, 128 (Japan) ; Jordan
and Fowler, Proc. U.S. nat. Mus., 26, 1903: 612 (Japan); Fowler, J. Acad. nat. Sci. Philad., (2) 12,

Fishes of the Western North Atlantic 275

1904: 499 (Padang, Sumatra) ; Jordan and Snyder, Proc. U.S. nat. Mus., 27, 1 904: 940 (Hawaiian Is.) ;
Jordan and Evermann, Bull. U.S. Bur. Fish., 25 (l), 1 905: 36 (Hawaiian Is.) ; Gunther, J. Mus. Godef-
froy, 6, 17, Fische der Sudsee 5, 1910: 483 (Hawaii) ; Southwell, Ceylon Adm. Rep. Mar. Biol., 1912—
1913: E 46, 49 (Ceylon) ; Zugmayer, Abh. bayer. Akad. Wiss., Math. Phys. Kl., 26 (6), 1913: 8 (Ara-
bian Sea); Pearson, Ceylon Adm. Rep. Mar. Biol., 1915-1918: f 12 (Ceylon); Jordan and Jordan,
Mem. Carneg. Mus., 10, 1927: 4 (Hawaiian Is.) ; Sarangdhar, J. Bombay nat. Hist. Soc, 44 (l), 1943:
103, 3 pis. (food, breeding, embryos, attacks on man, utilization, Bombay coast).

Carcharias {Prionodon) jasciatus Bleeker, Verh. batavia. Genoot., 24, 1852: 37 (Batavia) ; Gunther, Cat. Fish.
Brit. Mus., 8, 1870: 368 (Java).

Galeocerdo rayneri MacDonald and Barron, Proc. zool. Soc. Lond., 1868: 368, pi. 32 (excellent ill., Aust.,
Lord Howe I.) ; Gunther, Cat. Fish. Brit. Mus., 8, 1870: 377 (India) ; Day, Fish. India, 4, 1878: 718,
pi. 187, fig. 3 (India, Aust.) ; Ramsay, Proc. Linn. Soc. N. S. W., 5, 1 880: 95 (Aust.) ; Macleay, Proc.
Linn. Soc. N. S. W., 6, 1881: 353 (Aust.); Ogilby, Catal. Fish. N. S. W., 1886: i (Solomon Is.);
Proc. Linn. Soc. N. S. W., (z) j, l 889: 1768 (Aust., Solomon Is.) ; Day, in Blandford, Fauna Brit. India,
J, 1889: 20 (India); Kent, Naturalist in Aust., 1897: 193 (Aust.); Ogilby, Handbk. of Sydney, 1898:
117 (Aust.); Stead, Fish. Aust., 1906: 232 (Aust.); Zeitz, Trans, roy. Soc. S. Aust., j2, 1908: 290
(Aust.) ; Weber, "Siboga" Exped., 5-7, Fische, 1913: 591 (E. Indies) ; Robinson, Natal Fish. Rep., 1919:
50 (Natal) ; Pillay, J. Bombay nat. Hist. Soc, 33, 1929: 350 (India) ; Whitley, Mem. Qd. Mus., to (4),
1934: 199 (ref.); Aust. Zool., 8, 1937: 216 (Aust.); Fish. Aust., i, 1940: 113 (descr., size, weight,
Aust.).

Galeocerdo obtusus Klunzinger, Verh. zool.-bot. Ges. Wien, 2/, 1871: 664 (Red Sea).

Galeocerdo jasciatus van Kampen, Bull. Dep. Agric. Ind. neerl., 8, 1 907: 9 (East Indies).

Galeocerdo hemp-ichii Klunzinger, Verh. zool.-bot. Ges. Wien, 21, 1871: 663 (name only, and ref. to un-
published [?] ill. by Ehrenberg) ; Hilgendorf, in Hemprich and Ehrenberg, Symbol Physic, 1899: 8,
pi. 5 (ill.. Red Sea, not seen).

Galeocerdo arcticus Ogilby, Mem. Qd. Mus., 5, 1916: 78 (Aust.) ; Barnard, Ann. S. Afr. Mus., 21 (i), 1925:
27 (Natal) ; McCulloch and Whitley, Mem. Qd. Mus., 8 (2), 1925: 128 (ref.); Fowler and Ball, Bull.
Bishop Mus., 2(5, 1925: 4 (N. central Pacif. Is.) ; Griffin, Trans. Proc. N. Z. Inst., 5*, 1927: 137 (N.
Zealand) ; McCulloch, Fish. N. S. W., 2nd ed., 1927: 6, pi. I, fig. lia (Aust.) ; Fowler, Mem. Bishop
Mus., 10, 1928: 19 (Honolulu, Laysan, Nihoa) ; Phillipps, N. Z. J. Sci. Tech., 10, 1928: 226 (N. Zea-
land) ; Tirant, Serv. Oceanogr. Peches Indo-Chine, Note 6, 1929: 62 (Indo-China) ; Fowler, Proc. Pan.-
Pacif. sci. Congr., Java^ (4) 3, 1930: 490 (Indo-Pac. distrib.) ; Mem. Bishop Mus., 11 (5), 1931: 314
(ref.) ; Chevey, Inst. Oceanogr. Indo-chine, Note 19, 1 93 2: 6 (Indo-China) ; Giltay, Mem. Mus. Hist,
nat. Belg., Hors Ser., 5 (3), 1933: 12 (East Indies); Fowler, Mem. Bishop Mus., 11 (6), 1934: 385
(Solomon Is.) ; Marchand, Mar. biol. Rep., Cape Town, 2, 1935: 38 (west. S. Afr., Natal) ; Fowler, List
Fish. Malaya, 1938: 10 (ref.).

Galeocerdo cuvier Whitley, Mem. Qd. Mus., 10 (4), 1934: 199 (ref., Aust.) ; Fowler, Bull. U.S. nat. Mus.,
100 (/j), 1941: 186 (refs., loc, W. Pacif., Indian Oc).

Ge.n\xs Paragaleus'&udker, 1935

Paragaleus Budker, Bull. Mus. Hist. nat. Paris, (2) 7, 1935: 107; type species, P. gruveli Budker, 1935;
tropical West Africa, near Dakar.

Generic Synonyms:

Hemigaleus Garman, Bull. Mus. comp. Zool. Harv., 46, 1896: 203; for H. fectoralis Garman, southern New

England.
Hemigaleus (in part) Garman, Mem. Harv. Mus. comp. Zool., 36, 191 3: 149 (in part); not Hemigaleus

Bleeker, Verh. batavia. Genoot., 24, Plagiost., 1852: 45; not Negogaleus^ Whidey, Aust. Zool., 6, 1931:

334-

I. Proposed by Whitley to replace Hemigaleus Bleeker, 1851, the latter being preoccupied; see footnote 3, p. 164.

276 Memoir Sears Foundation for Marine Research

Generic Characters. Anal a little shorter at base than 2nd dorsal; spiracle present and
easily detected though small; 2nd dorsal originates over or a little anterior to origin of
anal; midpoint of base of ist dorsal nearer to axil of pectoral than to origin of pelvics;
caudal peduncle without lateral ridges; well marked precaudal pits, below as well as
above; a well marked labial furrow around corner of mouth and on each jaw, the upper
less than Yz as long as snout in front of mouth; teeth with smooth-edged cusps, the uppers
in sides of jaws oblique, notched outwardly, with 3 to 5 strong denticles toward the base;
lower teeth slender, erect, without basal denticles in front of jaw, but increasingly oblique
toward its corners, and with 3 to 5 strong denticles on the outer side toward the base, as in
the uppers; anterior margin of nostrils expanded as a narrow triangular lobe; gill open-
ings of moderate length, the 4th over origin of pectoral; axis of caudal only very slightly
raised, its lower anterior corner expanded as a definite lobe with pointed tip. Characters
otherwise those of the family.

Remarks. This genus is separated from Negogaleus Whitley by the fact that the
lower teeth in the sides of the jaw are oblique, notched, with their bases strongly denticu-
late on the outer sides. See Key, p. 264.

Range. So far known only from tropical West Africa, and from the coast of southern
New England.

Species. Two species known,^ very closely allied to each other but apparently sepa-
rable by the shapes of the snout and mouth."

Key to Species

la. Snout broadly rounded (Fig. 45 A) ; mouth about 2% times as broad as high.

feet oralis Garman, 1913, p. 276.

I b. Snout subrectangular, with narrowly rounded tip ; mouth only about 2 V3 times as broad

as high. ^rwui?// Budker, 1935.

Tropical West Africa.

Paragaleus fectoralis (Garman), 19 13
Figures 45, 46

Study Material. The type specimen, a female, 651 mm. long (Harv. Mus. Comp.
Zool.jNo. 847).

Distinctive Characters. This species is characterized, among carcharhinids having
spiracles, by the shortness of the anal fin relative to the second dorsal, by the position of
the first dorsal far forward, and by the comparatively long snout and very characteristic
teeth.

2. Only one specimen of each yet seen.

3. Budker's (Bull. Mus. Hist. nat. Paris, [2] 7, 1935: 110) measurements of the mouth do not agree with his illus-
tration i the present Key is based on the former.

Fishes of the Western North Atlantic

277

Figure 45. Paragaleus fectoralis, female, 651 mm. long, from oflF southern New England (Harv. Mus.
Comp. Zool., No. 847). A Anterior part of head from below, about 0.5 x. B Left-hand nostril, about 2.2 x.
C Upper and lower teeth, about 4.5 x. Z) Third upper tooth. E Tenth upper tooth. F Third lower tooth.
G Seventh lower tooth. D-G, about 9 x.

FiGtJRE 46. Paragaleus fectordis, pictured in Fig. 45.
A Dermal denticles, about 42 x. B Apical view of der-
mal denticle, about 84 x.

278 Memoir Sears Foundation for Marine Research

Description. Proportional dimensions in per cent of total length. Female, 651 mm.,
from the New England coast (Harv. Mus. Comp. Zool., type, No. 847).
Trunk at origin of fectoral: breadth 9.1; height 10.8.
Snout length in front of: outer nostrils 3.4; mouth 7.7.
Eye: horizontal diameter 2.3.
Mouth: breadth 6.5; height 2.5.
Nostrils: distance between inner ends 3.5.
Labial furrow length: upper 2.2 j lower 1.6.
Gill of ening lengths : ist 2.2; 2nd 2.2; 3rd 2.2; 4th 2.3; 5th 2.3.
First dorsal fin: vertical height 9. i ; length of base 9.8.
Second dorsal fin: vertical height 5.0 ; length of base 7.2.
Anal fin: vertical height 3.8; length of base 5.8.
Caudal fin: upper margin 23.5; lower anterior margin 10.2.
Pectoral fin: outer margin 14.8; inner margin 5.7; distal margin 11.7.
Distance from snout to: ist dorsal 27.6; 2nd dorsal 57.7; upper caudal 76.5;
pectoral 19.2; pelvics 47.2; anal 61.O.

Interspace between: ist and 2nd dorsals 21.5; 2nd dorsal and caudal 10. i; anal
and caudal 7.6.
Distance from origin to origin of: pectoral and pelvics 26.9; pelvics and anal 14.8.

Trunk slender, without mid-dorsal ridge, its height at origin of ist dorsal (where
highest) a little more than V7 of its length to origin of caudal, tapering evenly rearward.
Body sector from snout to cloaca about as long as tail sector. Caudal peduncle slender,
without lateral ridges, but with a well marked precaudal pit below as well as above.
Dermal denticles moderately closely spaced, partially overlapping, their blades on short
pedicels, nearly horizontal, with 5 longitudinal ridges, the margins with as many rather
blunt teeth, of moderate length, the median a little longest, and the outermost very small.

Head about Vs of total length, its dorsal profile moderately arched posteriorly but
flattened above anteriorly. Snout moderately thick, broadly rounded, its length in front of
mouth about ^3 length of head. Eye broad-oval, its midpoint about opposite front of
mouth, its horizontal diameter a little less than ^73 as long as snout in front of mouth.
Spiracle a small horizontal slit, about Vr as long as horizontal diameter of eye, on a level
with center of latter and behind it by a distance equal to about % the horizontal diameter
of eye. Gill openings all about equal in length, about as long as horizontal diameter of
eye, the 4th over origin of pectoral. Nostril strongly oblique, its inner end a little nearer
to front of mouth than to tip of snout, its anterior margin expanded as a prominent subtri-
angular lobe with sinuous inner margin and blunt tip. Mouth obtusely ovate, about 2V2
times as broad as high, occupying about % of breadth of head. Labial furrows strongly
developed, around corners of mouth, the upper extending about halfway toward the
symphysis, the lower about V2 as long as upper.

Teeth j^ °^ IgZaZu or is ' '^°* serrate; uppers subtriangular with broad bases, the 3 at

Fishes of the Western North Atlantic 279

symphysis small, symmetrical, erect, the next 9 to 12 increasingly oblique with nearly
straight inner margins but outer margins deeply notched, with 3 to 4 strong denticles
near basej those toward corners of upper jaw decreasing successively in size, broader
relative to height, and with cusps and denticles less prominent, the outermost 2 or 3
low, evenly rounded, and the outermost of all minute j first 6 lower teeth erect, with
slender cusps and broad bases, without denticles, the next 5 to 6 increasingly oblique,
their bases denticulate on the outer side as in upper teeth, the cusps decreasing in rela-
tive length in successive teeth, the outermost 5 lower teeth low and evenly rounded, the
outermost of all hemispherical, minute j i to 3 rows functional in upper jaw at symphysis,
I row along sides of jaw, and 2 to 3 rows at corners of mouth; 2 to 4 rows functional at
symphysis of lower jaw, i to 2 rows along sides of jaw, and 2 to 3 rows near corners.

Origin of ist dorsal about opposite inner corner of pectoral, the midpoint of its
base only about % as far from axil of pectoral as from origin of pelvics, its anterior
margin only very slightly convex, apex subacute, rear margin deeply concave, free
rear tip slender and about Vs as long as the base, its vertical height about as great as
distance from eye to and gill slit. Second dorsal similar to ist, but only a little more
than % as long at base and not more than Yo as large in area, its origin a little anterior to
origin of anal. Caudal with narrowly rounded tip, well marked subterminal notch, its
terminal sector nearly V3 the length of fin, the lower anterior corner forming an arcuate
sharp-tipped lobe directed rearward, about 40% as long as upper lobe, each measured from
its respective precaudal pit. Anal similar to 2nd dorsal, but only a little more than %
as long at base. Pelvics (in female) a little smaller than anal in area, with nearly straight
anterior margins, moderately concave distal margins, narrowly rounded apices and
subacute tips. Pectoral about 70% as long as head, only about as large in area as ist dorsal,
and very characteristic in shape,* with moderately convex outer margin which is increas-
ingly so toward tip, deeply concave distal margin, nearly straight inner margin, and nar-
rowly pointed tip.

Color. Described as grayish brown in life, paler below, the fins dark with pale hinder
margins; after many years in alcohol the type is mouse-gray above and of a paler shade of
the same below.

Size. The fact that a female of the closely allied West African species, 1,380 mm.
(about 54 inches) long, contained embryos, suggests that this Shark does not reach a
large size.

Developmental Stages. It is not known whether or not a placental connection is de-
veloped between embryo and mother; the embryos have not been described.

Habits. Nothing is known of its habits, but its teeth suggest a diet of fish or squid.

Range. So far known orJy from the type specimen taken oflF the coast of southern
New England. All that is known of its origin is that Garman° obtained it, apparently in a

4. Hence the specific name fectoralis.

5. Garman, Bull. Mus. comp. Zool. Harv., ^6, 1906: 203.

2 8o Memoir Sears Foundation for Marine Research

fresh condition, from a public aquarium known as the "Aquarial Gardens," the exhibits
for which came from "off the coasts of Massachusetts and Rhode Island.""

Remarks. We refer this species to Paragaleus Budker, 1935, rather than to Hemi-
galeus Bleeker, 1852 (in which genus Garman placed it), because of the conformation
of its lower teeth j Carman's description of the latter as having "erect narrow cusps on
broad bases, without denticles"" applies only to those in the front of the mouth, and not
to those along the sides of the jaw as noted above.

Synonyms and References:

Hemigaleus ■pecloralis Garman, Bull. Mus. comp. Zool. Harv., 46, 1906: 203 (descr., oif the coast of Massa-
chusetts or Rhode Island) ; Mem. Harv. Mus. comp. Zool., 36, 191 3: 1 50, pi. 4, fig. I-5, pi. 50, fig. 9,
pi. 52, fig. 2, pi. 56, fig. 4 (descr., ill. of type spec.) ; White, Bull. Amer. Mus. nat. Hist., y4, 1937:
124, pi. 13, fig. g (class., tooth).

Genus Prionace Cantor, 1849

Prionace Cantor, J. Asiat. Soc. Beng., t8, 1849: 1381; proposed in substitution for Prionodon Muller and
Henle, 1 841, preoccupied by Horsfield, 1823, for fossil mammals; type species, Squalus glaucus Linnaeus,
1758, designated by Jordan, Genera Fish., 2, 1919: 242.

Generic Synonyms:

Squalus (in part) Linnaeus, Syst. Nat., t, 1 75 8: 235.

Galeus Valmont, Diet. Hist. Nat., /, 1768: 371 ;^ type species, G. glaucus Valmont.

Carcharias (in part) Rafinesque, Indice Ittiol. Sicil., 1810: 45; Cuvier, Regne Anim., 2, 1817: 126.

Carcharhinus (in part) Blainville, Bull. Soc. philom. Paris, 1 81 6: 121 ; and many subsequent authors.

Prionodon (in part) Muller and Henle, Plagiost., 1841: 35.^

Cynocefhalus Gill, An. N. Y. Lye, 7, 1862: 401 ;' type species, Squalus glaucus Linnaeus, 1758.

Galeus Garman, Mem. Harv. Mus. comp. Zool., 56, 1913: 145 ;* not Galeus Rafinesque, 1810 (see discussion,

footnote I, p. 214).
Glyfkis Fowler, Mem. Bishop Mus., 10, 1928: 19; type species, Squalus glaucus Linnaeus, 1 75 8; Bull. U.S.

nat. Mus., lOO (/j), 1941: 178; not Glyfhis L. Agassiz, Poiss. Foss., 5, 1843: 243.°
Carcharhinus Whitley, Fish. Aust., i, 1940: 106, 107 (restricted to Squalus glaucus Linnaeus, 1758, and to

Prionace mackei Phillipps, 1 93 5, which appears to be identical with the latter).

6. Mem. Harv. Mus. comp. Zool., ^6, 191 3: 150, 151.

1. By Opinion 89 of the International Commission on Zoological Nomenclature (Smithson. Misc. Coll., 75 [3],
1925: 27), Valmont's names are not available, because they were not properly binomial.

2. Preoccupied by Horsfield, 1823, for fossil mammals.

3. This pre-Linnaean name, first proposed by Klein (Pise. Natural., Gedoni, 1742), was revived by Gill to replace
Prionace Cantor, 1849.

4. Carman's revival of Galeus Valmont, 1768, is not acceptable, according to the International Commission on Zoo-
logical Nomenclature; see footnote i, p. 214.

5. The fossil shark's teeth, to which L. Agassiz gave the name Glyfhis, and which he illustrated (Poiss. Foss., j, 1838:
pi. 36, fig. 10-13), are not at all suggestive of the corresponding teeth of Prionace, being cylindrical near the
base and with cutting edge confined to the lanceolate, laterally expanded tip. But they do resemble closely the
anterior lower teeth of Carcharias {Prionodon) glyf/iis Muller and Henle, 1841 (see footnote 4, p. 321).

Fishes of the Western North Atlantic 281

Generic Characters. Base of anal only about as long as base of 2nd dorsal} midpoint
of 1st dorsal considerably nearer to origin of pelvics than to axil of pectoral; 2nd dorsal
only about V2 as long at base as ist dorsal and much smaller in area; spiracles lacking;
caudal peduncle without longitudinal ridges, but with well marked precaudal pits both
above and below; midline of back, between dorsal fins, smooth, without longitudinal
ridge; a very short labial furrow at corner of mouth and on upper jaw, but none on lower;
upper teeth subtriangular, oblique, with inner margins strongly convex and outer margins
deeply concave; lower teeth more slender, erect; uppers with finely serrate margins,
lowers serrate or smooth. Development viviparous, with yolk-sac placenta. Characters
otherwise those of the family.

Remarks. Prionace is very closely allied to Carcharhinus but separable from it by the
location of the first dorsal fin relative to the pelvics and pectorals.

Range. Cosmopolitan in tropical and warm temperate latitudes of all oceans, includ-
ing the Mediterranean.

Species. The representatives of this genus generally had been considered as belonging
to a single wide ranging species until recently, when Phillipps" separated its New Zealand
representative as a new species mackei. According to Phillipps, mackei is distinguishable
from the Atlantic glauca by a shorter head (20% of total length as against 2S%), by a
shorter snout relative to its head, by pelvics larger than the anal, and by a straight instead
of concave distal margin of the pectoral. But these supposed differences are not consistent
when tested against Atlantic specimens. Among seven fish from Massachusetts Bay, for
example, ranging in length from about three to ten feet, the length of head from snout to
pectoral origin ranged from 20 to 24 per cent of the total length, it being 22 per cent in a
Japanese example of about 5 feet 6 inches (1,675 mm.).' The pelvics are also somewhat
larger in area than the anal in five Massachusetts Bay specimens of which we have meas-
urements, just as is the case in the New Zealand form. Neither does a comparison of the
outline of the pectorals of the Massachusetts Bay, Japanese and Australian specimens
reveal any consistent difference. The teeth, also, of an Australian specimen, as pictured
by Whitley,* are indistinguishable from those of the Japanese and Atlantic specimens that
we have examined (Fig. 47, 48). In short, we find no justification for retaining mackei as a
distinct species."

6. N. Z. J. Sci. Tech., z6, 1935: 238.

7. This is the specimen on which Garman (Mem. Harv. Mus. comp. Zool., ^6, 1913 : 145) based his description of

glaucm.

8. Fish. Aust., /, 1940: 95, fig. 88, 8.

9. Phillipps (N. Z. J. Sci. Tech., t6, 1935 : 238) further states that the origin of the first dorsal in his mackei is mid-
way between tip of snout and rear tip of second dorsal. But his photograph of the type specimen shows it as midway
between tip of snout and midlength of caudal peduncle, as it is in one of the larger Massachusetts Bay specimens,
as well as in our Japanese example; on the other hand Whitley (Fish. Aust., i, 194.0: fig. 104) pictures it as about
midway between snout and origin of caudal both for Australian and for New Zealand specimens. It is evident,
then, that no geographic separation exists in this respect either.

282 Memoir Sears Foundation for Marine Research

Prionace glauca (Linnaeus), 1758

Great Blue Shark

Figures 47, 48

Study Material. Twenty freshly caught specimens (2 females and 18 males), about
5 to 1 1 feet long, from various localities in the Gulf of Maine, Georges Bank and from
the offing of southern New England (jaws preserved) ; four preserved specimens, 539 to
2,160 mm. long, from Georges Bank and southern New England; Japanese specimen
1,675 mm. (about 5 feet 6 inches) ; also several other large specimens caught ofiF the New
England coast but not measured, and many seen at liberty.

Figure 47. Prionace glauca, male, about 2,175 ™™- l°ng) from Martha's Vineyard, Massachusetts (Harv. Mus.
Comp. Zool., No. 36035). A Head from below, about % natural size. B Left nostril, about natural size. C Der-
mal denticles, about 25 x. Z) Lateral and apical views of dermal denticle, about 25 x. £ Left-hand upper and
lower teeth, about % natural size. F Third upper tooth. G Ninth upper tooth. H Third lower tooth. / Eighth
lower tooth. F-I, about 1.5 x.

Distinctive Characters. The Blue Shark is easily distinguished from other West
Atlantic Sharks of its family by the combination of very long pointed snout, long falcate
pectorals, first dorsal fin set far back, teeth, and brilliant blue upper parts.

Description. Proportional dimensions in per cent of total length. Male, 2,175 mm.,
from Martha's Vineyard, Mass. (Harv. Mus. Comp. Zool., No. 36035). Male, 2,442
mm., from oflF Nauset, Cape Cod, Mass. (field specimen).

Fishes of the Western North Atlantic

283

Trunk at origin of pectoral: breadth 9.1, 10.8} height 8.8, 9.8.

Snout length in front of: outer nostrils 3.8, ; mouth 8.0, 7.8.

Eye: horizontal diameter 1.4, 1.4.
Mouth: breadth 5.4, 5.4; height 3.7, 3.6.
Nostrils: distance between inner ends 3.5, 3.3.

Labial furrow lengths: upper 0.3, .

Gill opening lengths: ist 1.9, 2.4J 2nd 2.0, 2.95 3rd 2.3, 3.15 4th 2.3, 2.95 5th

1.8,2.3.

First dorsal fin: vertical height 7.2, 7.2 j length of base 7.3, 7.7.

Second dorsal fin: wcrticzlhti^t 3.1, 2.7; length of base 4.4, 4.2.

Anal fin: vertical height 3.5, 3.1 ; length of base 3.6, 4.0.

Caudal fin: upper margin 25.5, 25.8; lower anterior margin 12.8, 12. i.

Pectoral fin: outer margin 23.4, 21.8 j inner margin 4.1, 4.1 ; distal margin 20.2,

18,0.

Distance from snout to: ist dorsal 35.5, 35.7; 2nd dorsal 63.7, 62.75 upper caudal

l^yiyi^ \

Figure 48. Prionace glauca. Teeth of a 6-foot 8-inch specimen from Platts Bank, Gulf of Maine. A Right-
hand upper and lower teeth, about 1.3 x. B Third upper tooth. C Seventh upper tooth. D Eleventh upper
tooth. E Third lower tooth. F Sixth lower tooth. G Tenth lower tooth. B-G, about 3 x.

2 84 Memoir Sears Foundation for Marine Research

74.5, 74.2; pectoral 21.5, 20.4; pelvics 49.5, 49-7; anal 63.7, 63.0.

Interspace between: ist and 2nd dorsals 21.4, 20.O; 2nd dorsal and caudal 6.8,

7.3 ; anal and caudal 6.2, 7.9.

Distance from origin to origin of: pectoral and pelvics 28.2, 29.2; pelvics and anal

14.4, 13.2.

Trunk very slender, its height at origin of ist dorsal (where highest) only about Ve
to y^ of its length to origin of caudal, without mid-dorsal ridge. Body sector from snout
to cloaca a little longer than tail sector. Caudal peduncle a little deeper than thick, without
lateral ridges but slightly rhomboid in cross-section. Precaudal pits subrectangular.
Dermal denticles close-spaced, usually overlapping, and so small that the skin is smooth to
the touch, their blades horizontal, as broad as long or broader, usually with 3, and occa-
sionally with 4 or 5, ridges, the apical margins weakly toothed to correspond} pedicels
short and stout.

Head noticeably long, its length to origin of pectoral averaging about Vs of total
length. Snout conical, with narrowly rounded tip, noticeably long, its length in front of
mouth about % of length of head to origin of pectoral in large specimens and relatively
a little longer in small specimens. Eye broadly oval or nearly circular, with well de-
veloped nictitating membrane, its midpoint about opposite front of mouth, its hori-
zontal diameter between ^4 and % as long as snout in front of mouth. Gill openings
noticeably short, the 3rd longest, a little shorter than horizontal diameter of eye in
small specimens, but about twice as long as eye in large ones, the ist a little longer than 5th,
the 4th over or very slightly posterior to origin of pectorals." Nostril oblique, its inner end
a little nearer to front of mouth than to tip of snout, its anterior margin only slightly ex-
panded as a low, inconspicuous subtriangular lobe with rounded tip. Mouth evenly
rounded, a little less than % as high as broad," occupying about % of breadth of lower
surface of head. A deep pit at corner of jaw, concealed when mouth is closed, but sub-
triangular when open, and extending for a very short distance at approximately a right
angle onto the upper jaw but not onto the lower.^^

Teeth itZi °o ^n to 15 '"^ J^^^ examined; uppers so closely spaced that successive

teeth overlap laasally, subtriangular, slightly longer than broad, oblique, their outer mar-
gins deeply concave and inner margins convex, with edges serrate; usually one tooth at
symphysis much smaller than those flanking it, but similar in form, its point directed
toward the right in some specimens, toward the left in others (this tooth is lacking occa-
sionally) ; next 4 or 5 teeth largest and about equal, the subsequent teeth successively
smaller toward the angle of the mouth, with the outermost very small; lower teeth erect,
much more slender than uppers toward center of mouth, but increasing in relative breadth
and decreasing in length toward angles of mouth, their margins usually very finely serrate,

10. In a Japanese specimen, the third is similarly longest, and the fifth is slightly shorter than the first.

11. 56 to 68 per cent in specimens examined. 12. Much as in Carc/uirhinus.

Fishes of the Western North Atlantic 285

but an occasional tooth smooth-edged and others partially so; an irregular group of 2 to 4
teeth at the lower symphysis, much smaller than those on either side and with relatively
narrower bases; i to 2 rows of teeth functional in front and i in the sides of mouth in upper
jaw; I to 3 rows functional in front and i row laterally in lower jaw.

Anterior margin of first dorsal about as long as snout in front of mouth, its origin
posterior to inner corner of pectoral by a distance about % as long as its anterior margin
and about midway between tip of snout and precaudal pit, the midpoint of its base at a
vertical a little less than % (about 70%) as far from origin of pelvics as from axil of
pectoral ; its anterior margin nearly straight, apex rounded, its posterior margin deeply con-
cave toward the base, its free rear corner moderately acute, sharp-tipped, about % as long
as its base. Second dorsal about Yi as long at base as ist, and less than Vs as large in area,
its posterior margin less deeply concave, but its free rear corner more slenderly pointed
and about as long as the base, its origin about over or very little posterior to origin of anal.
Caudal about V4 of total length or a little more, its axis moderately raised, its terminal
sector slender with pointed tip, the subterminal notch strongly marked, its lower anterior
corner expanded as a blunt-tipped lobe, about V2 as long as the upper margin of fin. Anal a
little larger than 2nd dorsal, its anterior margin convex, apex rounded, its posterior margin
very deeply concave, its free rear corner acutely pointed and about % as long as its base.
Pelvics only about as large in area as anal, or a little larger, with nearly straight anterior and
inner margins, slightly concave distal margins and narrowly rounded corners. Pectoral
noticeably long, being about as long as head to 5th gill opening in medium-sized and large
specimens, but relatively somewhat shorter in small ones,'"^ only a little more than ^/3
as broad as long, tapering toward tip, its anterior margin moderately convex (more
strongly so in small specimens), the inner margin moderately concave proximally, the
apex very narrowly rounded, the inner corner more broadly so.

Color. Living and freshly caught specimens are dark indigo blue along the back,
shading to a clear bright blue" along the sides, and to snow white below; the tips of the
pectorals are usually dusky and the anal partly so. But the beautiful blue of the back and
sides darkens to a slaty or sooty gray soon after death.

Size. The Blue Shark is reputed to reach a length of 20 feet and commonly 1 5 feet.
Actually, however, about 12 feet 7 inches (3.83 m.) is the longest of which we have found
positive record; an i i-foot specimen is the longest we have handled. Embryos as large as
350 to 450 mm. have been recorded, and free living specimens as small as 2 1 to 36 inches
(539 mm.; 661 mm.; 910 mm.; see Study Material, p. 282). The sizes of the females in
which young have been found suggest that this Shark does not mature until a length of at
least seven to eight feet is reached. Corresponding to their slender build. Blue Sharks
are less heavy, length for length, than the more stout-bodied species; probably the follow-
ing weights at different lengths, collected from various sources, are representative : 6 to 7

12a. 15.4 to 17.1 per cent of total length in three specimens, 539 to 910 mm. long.
13. "Sailor Blue" in Ridgway, Color Standards and Color Nomenclature, 1912: pi. 21.

286 Memoir Sears Foundation for Marine Research

feet, 6s to 70 pounds; 7 to 8 feet, 100 to 114 pounds; about 9 feet, 164 pounds." Al-
though we have handled many, we have weighed none.

Developnental Stages. The Blue Shark is viviparous, its embryo having a well de-
veloped yolk-sac placenta attached to the uterine wall of the mother.*" The number of
young in a litter is large, 2 8 to 54 having been reported in the Mediterranean from females
of 8 feet 3 inches to 9 feet 4 inches."

Habits. This is a pelagic species, encoimtered indifferently far out at sea and in con-
tinental waters, its wanderings no doubt directed chiefly by the search for food, although
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. Many are seen in coastal waters
as well as offshore, and in some regions, near Woods Hole for example, it often comes
close enough to the land to be caught in pound nets, as many other sharks often are.
In our experience it is rather sluggish when not disturbed, but it swims powerfully and
swiftly when in pursuit of prey. Normally it feeds on the smaller fishes that may be
available locally, and on a variety of cephalopods. In northern waters herring and mack-
erel, and in European seas sardines, appear to be the chief items in its diet, as well as
Spiny Dogfish (Squalus acanthias). No doubt it also consumes large quantities of bottom
fish on the fishing banks. For example, we have repeatedly had Blue Sharks pick up cod,
haddock and American pollock (Pollachius virens) that had b?en returned to the water
on Georges Bank during the cod-tagging cruises of the United States Bureau of Fisheries.

In warmer seas they are also known to feed on anchovies and flyingfish, and occa-
sionally on a sea bird that is resting on the water. We find no record of their preying on
larger animals while the latter are alive. They sometimes follow sailing vessels in warm
seas for days or even weeks picking up offal. And their habit of gathering when a Sperm
Whale has been killed, probably by tracing the blood-scent, has long been proverbial
among whalemen, one often struggling up on the carcass to "cling there until a descending
blubber-spade had put an end to all its ambitions," to quote from an eye-witness account.
"If the cutting in of the whale was at any time deferred . . . the sharks . . . would then
attack the carcass, and, thrusting their heads partly above the surface, would bite large
mouthfuls out of the blubber. ... A blue shark horribly mutilated by repeated thrusts
of a whaleman's blubber-spade, was seen to return immediately to the whale on which it
had been feeding and to continue ravenously. . . . "" A recent report of one that came to
eat scraps thrown to it from a boat, even after it had been transfixed by a harpoon, similarly
illustrates its indifference to injury.**

14. From Roule, Result. Camp. sci. Monaco, 52, 1919: 1145 Holcombe, Modern Sea Angling, 1921: 144; Sdiultz
(J. Mammal., ip, 1938: 484, "Prionace") gives a weight of 433.6 kg. (about 950 pounds), but this is so far
out of line with other recorded weights that some other stouter-bodied shark was doubtless intended.

15. For a recent anatomical account of the placenta, with references, see Calzoni (Pubbl. Staz. zool. Napoli, 75,
1936: 109).

16. For numbers and sizes of embryos, see Lo Bianco (Mitt. zool. Sta. Neapel, 19, 1909: 666).

17. Nichols and Murphy, Brooklyn Mus. Sd. Bull., 5 (1), 1916: 11.

18. Piers, Proc. N. S. Inst. Sci., 18, 1934: 202.

Fishes of the Western North Atlantic 287

It is not known whether there is a circumscribed breeding season or whether young
are produced at all times of the year, which seems more likely, this being a warm-water
species. Available information as to its young stages is summarized under Developmental
Stages (p. 286).

Relation to Man. The Blue Shark is of no commercial value, nor has it been in the
past, but it takes a large bait readily, and a few are caught for sport by anglers." Our own
experience, often repeated, has been that a "Blue" puts up little resistance when hooked on
a heavy hand line until drawn in nearly to the ship's side, but then it threshes about vio-
lently as it is being hoisted aboard. But by anglers' accounts a large one hooked on rod and
reel may resist strongly, making long rushes for a considerable time. While most often
hooked on natural bait, it will sometimes take an artificial lure, as in the case of one five
feet long recently caught on a feather jig tipped with pork rind, oflF Boone Island, Maine.
In spite of its razor-sharp teeth the Blue Shark has always been held in contempt by whale-
men who are the most familiar with it. There is no well authenticated record of its attack-
ing swimmers, notwithstanding sailors' yarns to the contrary.

Range. Cosmopolitan, in the tropical, subtropical and warm-temperate belts of all the
oceans (including the Mediterranean).

Occurrence in the Atlantic. This is no doubt the most plentiful of the larger oceanic
sharks of the Atlantic''" and it is the one with which we are the most familiar} around it
most of the sailors' superstitions about sharks have centered. In the eastern side of the
Atlantic it has been reported for so many localities and has been described so often as com-
mon that there is adequate evidence that it is practically universal off the coasts of west
tropical Africa (Senegambia, Morocco), around the off-lying island groups (Cape Verdes,
Canaries, Azores), and throughout the Mediterranean. It is also common, at least in
summer, offshore along the Atlantic coasts of the Iberian Peninsula and France, although
not often coming close to land. During the warm months it appears regularly off the
south and west coasts of England north to Scotland in numbers sufficient for fishermen
to be familiar with it, although it is seen less often on the French coast of the Channel,
where we find only two records, both for Cherbourg. It penetrates the North Sea eastward
to the Skagerrak, occasionally entering the western Baltic, and stray specimens are met with
as far north as the Orkneys and southern Norway. Southward, in the eastern Atlantic, it is
recorded for the west coast of South Africa.

Old time reports by sperm whalers, who were very familiar with the Blue Shark
for reasons given above, show that it is generally, although very irregularly, distributed
over the midbelt of the Atlantic. Its latitudinal range is as wide in the western side as it is
in the eastern, i.e., from the offing of the Rio de La Plata in the south to Nova Scotian
waters (regularly) and to the Banks of Newfoundland (occasionally) in the north. Its

19. For readable accounts of rod and line fishing for Blue Sharks, see Wise (Tigers of the Sea, 1937: 67) and
Holcombe (Modern Sea Angling, 1921: 152).

20. Recent authors (Nichols and Murphy, Brooklyn Mus. Sci. Bull., 3 [i], 1916: 10) write of seeing "hundreds,
even thousands" of them during a sperm-whaling voyage in the tropical Atlantic

288 Memoir Sears Foundation for Marine Research

coastwise distribution in the west is in strong contrast to that in the eastern side of the
Atlantic, for while In the latter area it is most often encountered In the tropical-subtropical
belt, in the former there are but two published inshore records of It for the entire West
Indian region (St. Thomas and Cuba), with one for Florida (Miami) and none for the
Gulf of Mexico or Caribbean littoral. But it occurs more commonly there, offshore, than
this meagre record would suggest, for It is occasionally caught and often seen out in the
open sea around Cuba," while recently (September 1945) one about 12 feet long was taken
600 miles ESE. of Bermuda by the research vessel "Atlantis."

Neither is there any record for It on the coast between southern Florida and Chesa-
peake Bay; and stray specimens only have been reported from the coast of Maryland,
New Jersey (two records), or from the vicinity of New York (two records) ; this is suffi-
cient evidence that Blue Sharks rarely come Inshore anywhere along this extensive sector
of the coast. But they are much more common as summer visitors farther to the east and
north. For example, 28 were counted 4 to 10 miles off Block Island on August 22, 1943,
in an hour's run, with the number seen during the day estimated as 150 to 200.^^ There
are many records of specimens taken in the traps close to land at Woods Hole, and it is a
well known shark at Nantucket and on the off-lying shoals. Blue Sharks swimming at the
surface are a familiar sight to fishermen In summer on Georges Bank, as we can bear wit-
ness. It was formerly regarded as a stray only, but it Is now known to be a rather regular
summer visitor in the Gulf of Maine, where it appears occasionally in July but more com-
monly In August and September, at least as far northward as Platts Bank, where three were
caught and others were In sight of the vessel at nearly all times during the day on Septem-
ber 3, 1925. Two have been reported recently to us as taken on the Maine coast a few miles
east of Casco Bay.^' Many have been seen also within Cape Cod and Massachusetts Bay,
and to our own knowledge several have been taken there in recent summers, even close to
Boston Harbor,^*

Still farther to the northward the Blue Shark is quite common In some summers along
the Nova Scotian coast as far as Cape Breton, both inshore and on the offshore banks.
For example, near Halifax in 1920 It was first reported on August 15, was most plentiful
during the last week of that month, and was last reported on October lOth. It has been
recorded also as a stray on the Grand Banks of Newfoundland. However, It is strictly a
summer visitor to the coasts of the northeastern United States and Canada 5 none have been
reported there later than mid-October. The great majority of those taken or seen there are
of medium or large size. Moreover, it appears that few, if any, females take part in this
yearly incursion, for all except two regarding which we have pertinent information have
been males.

21. Personal communication from Luis Howell-Rivero.

22. Three were harpooned and one, about 21 inches long, was brought into Woods Hole and identified.

23. Personal communication from Walter H. Rich.

24. Eighteen were reported to us from Massachusetts during the summer of 1935; measurements and photographs
of several of them were contributed by J. R. Lowes, Jr., a shark angler of wide experience.

Fishes of the Western North Atlantic 289

Information as to the occurrence of the Blue Shark in coastal waters in the south-
western Atlantic is limited to records for Brazil and the offing of the Rio de La Plata.

Synonyms:

References for Atlantic, South Africa, West Coast of America:^"

Squalus gkucus Linnaeus, Syst. Nat., /, 1758: 235 (Europ. Oc.) ; Syst. Nat., 12th ed., 1766: 401; Watson,
Philos. Trans., 68 (i), 1779: 789, pi. 12, 2 fig. (descr., ill., Devonshire, Eng.) ; Bloch, Naturg.
Fisch. Dtsch., 3, 1784: 78, pi. 86 (North Sea, Baltic); Bonnaterre, Tabl. Encyc. Meth. Ichthyol.,
1788: 9, pi. 7, fig. 22 (general); Gmelin, in Linnaeus, Syst. Nat., /, 1789: 1496 (descr.); Walbaum,
P. Artedi Genera Pise. Emend. Ichthyol., 1792: 513 (descr.); Bloch and Schneider, Syst. Ichthyol.,
1 801: 131 (descr.); Latreille, Nouv. Diet. Hist. Nat., 24, 1804: 72 (in table of contents); Turton,
Gen. Syst. Nature, i, 1806: 919 (general); Risso, Ichthyol. Nice, 1810: 26 (descr., Medit.) ; Retzius,
Obs. Anat. Chondropt. Lundae, 1819: 3, pi., fig. I (anat.) ; Martens, Reise Vened., 2, 1824: 408
(Medit., not seen); Nardo, Prod. Ittiol. Adriat., 1827: 9 (Adriatic); Voigt, in Cuvier, Tierreich, 2,
1832: 506 (descr.); Couch, Cornish Fauna, 1838: 50 (Cornwall); Swainson, Nat. Hist. Fish. Amphib.
Rept., 2, 1839: 312 (general) ; Bonaparte, Icon. Faun. Ital., 3, Fasc. 37: 1840: pi. not numbered, fig. I
(descr., Medit.); Cat. Pesc. Europ., 1846: 18 (Medit.); Nardo, Pesc. Publ. com. Nuovo Venez., 1853:
15 (Medit.); Machado, Feces Cadiz, 1857: 9 (near Cadiz); Gemmellaro, Atti Acad. Gioenia, (2) 19
(3), 1864: 123 (Medit.) ; Ninni, Ann. Soc. Nat. Modena, 5, 1870: 66 (Medit.) ; Buckland, Hist. Brit.
Fish., 1 881: 212 (general, Gt. Brit.); Tarel, Act. Soc. linn. Bordeaux, 86, Proc. Verb., 1934: 113
(Arcachon, France).

Squalus adscensionis Osbeck, Voy. China E. Indies, 1771 : 78 (not seen, quoted from Bloch, 1784).

Le Squale Glauque, Lacepede, Hist. Nat. Poiss., i, 4° ed., 1 798: 165, 213, pi. 9, fig. I, in Buffon, Hist. Nat.
(general) ; in Sonnini, Hist. Nat. Poiss., 3, 1802: 405 (color, general).

Blue Shark, Yarrell, Brit. Fish., 2, 1836: 381 (Brit. loc. records).

Carcharias glaucus Rafinesque, Indice Ittiol. Sicil., 1810: 45 (Sicily); Cloquet, Diet. Sci. Nat., 7, 1817: 68
(general); Cuvier, Regne Anim., 2, 1817: 126 (general); Fleming, J., Hist. Brit. Anim., 1828: 167
(Cornwall); Bory de St. Vincent, Diet. Class. Hist. Nat., 15, 1829: 597 (general); Agassiz, L., Poiss.
Foss., 3, 1835-1837: 12, pi. F, fig. I (teeth); Owen, Odontogr., 1840-1845: 27 (teeth); Cuvier,
Regne Anim., ill. ed., Poiss., 1843: 361 ; Hamilton, Brit. Fish., 2, 1843: 303 (Gt. Brit.); Busch,
Selach. Ganoid Encephalo, 1848: pi. 3, fig. 4-6 (brain) ; Gosse, Nat. Hist. Fish., 1851 : 306 (general) ;
Baikie, Zoologist, //, 1853: 3846 (Orkney I.); Owen, Cat. Osteol. Roy. Coll. Surg., /, 1853: 93, 94
(vertebrae); Gray, Hist. Chile Zool., 2, 1854: 364 (Chile, not seen); Van der Hoeven, Handb. Dier-
kunde, (2) 2, 1855: 262 (general); Thompson, W., Nat. Hist. Ireland, 4, 1856: 250 (Ireland); Brito
Capello, J. Sci. math. phys. nat. Lisboa, 2, 1870: 142 (Portugal) ; Miklucho-Maclay, Beitr. Vergl. Neurol.
Wirbelt., I, 1 870: 23, pi. 4, fig. IO-12 (brain); Mcintosh, Ann. Mag. nat. Hist., (4) 14, 1874: 424
(w. coast of Scot.); Reguis, Ess. Hist. Nat. Provence, I (i), 1877: 47 (Medit.); Gunther, Guide to
Study Fish., 1880: 317, fig. 112 (general); Brito Capello, Cat. Peix. Portugal, 1880: 46 (Portugal);
Moreau, Hist. Nat. Poiss. France, i, 1881: 329 (France, Medit.) ; Perugia, Elenc. Pesc. Adriat., 1881:
52 (Adriatic) ; Day, Fish. Gt. Brit., 2, 1 880-1 884: 289, pi. 152 (descr., Gt. Brit.) ; Jordan and Gilbert,
Bull. U.S. nat. Mus., 16, 1883: 872 (name) ; Jouan, Mem. Soc. Imp. Sci. nat. Cherbourg, 24, 1884: 313
(Cherbourg, France); Gunther, "Challenger" Rep., 31, 1 889: 5 (S. Atlant., Japan); Bellotti, Atti
Soc. ital. Sci. nat., 33, 1891: 112; Carus, Prod. Fauna Medit., 2, 1 889-1 893: 512 (Medit., Iocs,
and refs.); Smitt, Hist. Scand. Fish., 2, 1895: 1 1 30, pi. 50, fig. 3 (descr., refs., Seand. Iocs.);
Murray, "Challenger" Rep., Summary, 2, 1 895: 892, 893, 1 207 (S. of Japan; west. S. Atlant.
off Rio de la Plata); Nobre, Ann. Sci. nat. Porto., 2, 1895: 225 (Portugal); Osorio, J. Sci. math,
phys. nat. Lisboa, (2) 5, 1898: 200 (St. Thome I.); Evans, Ann. Scot. nat. Hist., 7, 1898: 239
(Firth of Forth, Scot.); Sicher, Atti Accad. gioenia, (4) ri, 1898: 14 (Medit.); Jouan, Mem.
Soc. Sci. nat. Cherbourg, 31, 1900: 222 (Cherbourg); Steinhardt, Arch. Naturg., Jahrg. 69 (x),

25. For references for the central and western Pacific, Australasian region and Indian Ocean, see Fowler (Bull. U.S.
nat. Mus., 100 [x^], 1941 : 178).

290 Memoir Sears Foundation for Marine Research

1903: pi. I, fig. 13-15 (oral and pharyngeal denticles); Richard, Bull. Inst, oceanogr. Monaco, ip,
1904: 29 (off Portugal); Borca, Arch. Zool. exp. gen., (4) 4, 1905: 204, 208 (Roscoff, France);
Imms, Proc. zool. See. Lond., 1905: 43 (denticles) ; Lo Bianco, Mitt. zool. Sta. Neapel, jp, 1909: 666
(embryos, food, Medit.) ; Le Danois, Ann. Inst, oceanogr. Monaco, 5 (5), 191 3: 20 (Eng. Channel);
Jenkins, Fish. Brit. Isles, 1925: 307, pi. 118 (Gt. Brit.); Ehrenbaum, in Grimpe and Wagler, Tier-
welt N.- u. Ostsee, Lief 7 (12'), 1927: 12 (general. North Sea, Baltic) ; Wilson, Proc. U.S. nat. Mus., do
(5), 1932: 60, 440 (parasites); Ehrenbaum, Cons, explor. Mer Faune Ichthyol. N. Atlant., 1933:
plate not numbered (N. Europ. distrib.) ; Belloc, Rev. des Trav. Peches Marit., 7 (2), 1934: 131
(Morocco, Rio de Oro, Senegal, Madeira, Azores, Canaries, Gorringe Bank) ; Ranzi, Pubbl. Staz. Zool.
Napoli, 13, 1934: 351, 398 (uterine fluid and refs. to placenta); Nobre, Fauna Marinha Port.
Vert., 7, 1935: 421 (off Portugal); Calzoni, Pubbl. Staz. Zool. Napoli, 15, 1936: 169 (develop,
placenta, Medit.); Needham, Biochem. Morphogen., 1942: 41 (embryonic weight).

Squalus {Carcharhinui) caeruleus Blainville, in VieiUot, Faune Franc, 1825: 90, pi. 23, fig. l^' (descr.,
Medit.).

Squalus {Carcharhinus) glaucus Blainville, 1 825 : in Vieillot, Faune Franc, 1 830: 92, pi. 23, fig. 2'° (descr.) ;
Gray, List. Fish. Brit. Mus., 1 851: 44 (English coast).

Squalus (Carckarias) glaucus Jenyns, Manual Brit. Vert. Anim., 1835: 499 (Gt. Brit.).

Carcharias (Prionodon) glaucus Muller and Henle, Plagiost., 1841: 36, pi. 11 (descr., Medit., Atlant.);
Dumeril, Hist. Nat. Poiss., 1865: 353 (descr., distrib.); Giinther, Cat. Fish. Brit. Mus., 8, 1870: 364
(descr., East Atlant., Medit., St. Helena, India); Van Beneden, Poiss. Cotes Belg. Parasit., 1870: 4
(parasites, Belgium); Doderlein, Man. Ittiol. Medit., 2, 1881: 42 (Medit.) ;''^ Rochebrune, Act. Soc.
linn. Bordeaux, (4) 6, 1882: 41; Faune Senegambie, Poiss., i, 1882-1885: 18 (color, Senegambia) ;
Werner, Zool. Jb., Syst. Abt., 21, 1904: 283 (Medit.); Metzelaar, Trop. Atlant. Visschen, 1919: 187
(St. Thomas, W. I.); Lampe, Dtsch. Sudpol. Exped., (15) Zool., 7 (2), 1914: 213 (Simonstown, S.
Afr.).

Corcharias (PrionoJon) hirundinaceus Valenciennes, in Muller and Henle, Plagiost., 1 841 : 37; Dumeril, Hist.
Nat. Poiss., I, 1865: 354 (descr., Brazil).

Prionodon glaucus Socage and Brito Capello, Poiss. Plagiost. Port., 1 866: 1 7 (chars., Portugal) ; Canestrini, in
Cornalia, et al., Fauna d'ltal., tg, 1 872: 47 (Medit.); Gegenbaur, Unters. Vergl. Anat. Wirbelt., 3,
1872: pi. 2, fig. 4, pi. 7, fig. 4, pi. 18, fig. 5 (skull, skelet.) ; Graeffe, Arb. zool. Inst. Univ. Wien, 28,
1886: 446 (Medit.) ; de Braganza, Result. Invest. Sci. "Amelia," 2, 1904: 44, 45 (off Portugal) ; Borri,
Mem. Soc. tosc Sci. nat., 44, 1934: 89 (Medit.).

Carc/iarinus glaucus Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1 883: 22, 60 (descr., range); Jordan, Rep.
U.S. Comm. Fish. (1885), 1887: 795 (distrib.); Eigenmann, Proc U.S. nat. Mus., 15, 1893: 133 (S.
California) ; Barnard, Ann. S. Afr. Mus., 21 (l), 1925: 26 (Agulhas Bank) ; Norman and Fraser, Giant
Fishes, 1937: 36 (general).

Corcharias fugae Perez Canto, Estud. Escual., Chile, 1886: 2 (Chile); Philippi, An. Univ. Chile, yi, 1887:
541, pi. I, fig. 2 (Valparaiso).

Prionace glauca Jordiji and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 215 (distrib.) ; Bull. U.S. nat.
Mus., 47 (i), 1896: 33 (descr., California) ; Linton, Proc. U.S. nat. Mus., 20, 1897:423,424, 439, 443,
447, 449 (parasites. Woods Hole record); Smith, H. M., Bull. U.S. Fish Comm., 17, 1898: 88
(Woods Hole record) ; Bean, Rep. For. Comm. N. Y., 1901: 377 (Woods Hole) ; Bull. N. Y. St. Mus.,
60, Zool. 9, 1903: 25 (descr.); Sharp and Fowler, Proc. Acad. nat. Sci. Philad., 5(5, 1904: 505 (Nan-
tucket); Collett, Norges Fiske, 5, 1905: 76 (south. Norway); Cornish, Contr. Canad. Biol. (1902—
1905), 1907: 81 (off C. Breton, Nova Scotia); Ribeiro, Arch. Mus. nat. Rio de J., 14, 1907: 51,
199 (Brazil); Starks and Morris, Univ. Calif. Publ. Zool., 5, 1907: 167 (San Diego, California);
Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 4 (Woods Hole, Nantucket); Fowler, Proc.
Acad. nat. Sci. Philad., 60, 1908: 62 (Italy); Roule, Bull. Inst, oceanogr. Monaco, 243, 1912: 6 (loc.
records, E. Atlant. and Medit.); Halkett, Check List Fish. Canad., 1913: 40 (Banks of Newfound-

26. We cannot find whether or not this plate was ever published.

27. See Doderlein, 1881, for additional references for the Mediterranean in publications not accessible to us.

Fishes of the Western North Atlantic 291

land, California); Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., 5/ (2), 1913: 735 (Wood*
Hole region) ; Nichols and Murphy, Brooklyn Mus. Sci. Bull., 3 (i), 191 6: 9 (habits, N. York and south.
New Engl.) ; Osburn and Nichols, Bull. Amer. Mus. nat. Hist., 35, 1916: 141 (lower California) ; Ribe-
iro. Fauna brasil. Peixes, 2(1) Fasc. I, Mus. nac. Rio de J., 1923: 9, pi. I (descr., Brazil) ; Nichols and
Breder, Zoologica, N. Y., 9, 1927: 15 (near N. York); Rey, Fauna Iberica Feces, /, 1928: 336 (descr.,
Spain) ; Jordan, Manual Vert. Anim. NE. U.S., 1 929: 9 (general) ; Breder, Field Bk. Mar. Fish. Atlant.
Coast, 1929: 15 (general); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 17
(distrib.) ; Schroeder, Bull. Boston Soc. nat. Hist., 58, 1 931: 3 (Gulf of Maine, Georges Bank);
Bellon and Mateu, Notas Inst. esp. Oceanogr., (2) 53, 1931: 16 (Canaries, Iberian Penin.) ; Gudger, Sci.
Mon. N. Y., 24, 1932: 417 (food); Beebe and Tee-Van, Zoologica, N. Y., 15, 1932: 119 (Ber-
muda); Field Bk. Shore Fish. Bermuda, 1933: 26 (Bermuda); Gowanloch, Bull. La. Conserv. Dep.,
23, 1933: 222 (name only); Young and Mazet, "Shark Shark," 1933: 267 (general); Bigelow and
Schroeder, Canad. Atlant. Fauna, biol. Bd. Canad., 12*, 1934: 8 (descr., distrib.) ; Piers, Proc. N. S. Inst.
Sci., 18, 1934: 202 (off Nova Scotia) ; Nichols and La Monte, Amer. Mus. Novit., 901, 1935: 6 (Nova
Scotia) ; Vladykov, Proc. N. S. Inst. Sci., ig, 1935: 7 (Nova Scotia) ; Vladykov and McKenzie, Proc. N. S.
Inst. Sci., jp, 1935: 45 (Nova Scotia) ; Walford, Fish Bull. Sacramento, 45, 1935: 34 (Pacif. coast U.S.,
Gulf of California to Washington); Barnhart, Mar. Fish, south. Calif., 1936: 9 (off California);
Bigelow and Schroeder, Bull. U.S. Bur. Fish., 48, 1936: 32 (small sizes, Gulf of Maine, Grand
Banks); Schultz, Publ. Biol., Univ. Wash., Seattle, 2 (4), 1936: 131 (Washington, Oregon); Howell-
Rivero, Mem. Soc. cubana Hist. Nat., //, 1937: 285 (Cuba); Wise, Tigers of the Sea, 1937: 68, 176
(general) ; Tortonese, Atti Soc. ital. Sci. nat., yy, 1938: 300 (Medit., Japan) ; Springer, Proc. Fla. Acad.
Sci., 5, 1939: 1 7 (Florida) ; Beebe and Tee-Van, Zoologica, N. Y., 26, 1 941 : 104 (Gulf of California) ;
Norris, Plagiost. Hypophysis, 1941 : 30, pi. 62, fig. 30-32 (brain) ; Lunz, Bull. S. Carolina St. Planning
Bd., 14, 1944: 26 (Florida); Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb.
Comm., Wash., 1945: 121, fig. 43 (descr., habits, range, ill.). Not Prionace glauca Townsend, Bull. N. Y.
zool. Soc, 24 (6), 1931 : upper fig. on p. 170, which is an isurid.

Galeus glaucus^^ Garman, Mem. Harv. Mus. comp. Zool., jtf, 1 91 3: 145, pi. 3, fig. 1-3 (descr., Mass., Japan);
Nichols, Abstr. Linn. Soc. N. Y., 20-23, 1913: 91 (off N. York) ; Roule, Result. Camp. sci. Monaco, 52,
1919: 114 (E. Atlant.) ; Linton, Proc. U.S. nat. Mus., 64, 1924: (21), 7, 38, 39, 44 (parasites. Woods
Hole) ; Bigelow and Welsh, Bull. U.S. Bur. Fish., 40 (i), 1925: 28 (Gulf of Maine, Nova Scotia) ; Bige-
low and Schroeder, Bull. Mus. comp. Zool. Harv., 68, 1927: 244 (Gulf of Maine) ; Firth, BuU. Boston
Soc. nat. Hist., 61, 1931: 9 (off C. Cod); Wilson, Bull. U.S. nat. Mus., 152, 1932: 440, 453 (parasites.
Woods Hole) ; Wise, Tigers of the Sea, 1937: 176 (general) ; White, Bull. Amer. Mus. nat. Hist., 74,
1937: 125, pi. 2, fig. g; pi. 6, fig. h; pi. 13, fig. b; pi. 19, fig. a; pi. 26, fig. a; pi. 38, fig. d, e; pi. 50,
fig. a (class., anat.).

Glyfhis glaucus Fowler, Copeia, 30, 1916: 36 (mid. Atlant. coast); Proc. Acad. nat. Sci. Philad., y8, 1926:
276 (Chile) ; Mem. Bishop Mus., jo, 1928: 19 (Oceania) ; Proc. Acad. nat. Sci. Philad., 80, 1929: 608
(N. Jersey); Bull. Amer. Mus. nat. Hist., yo (i), 1936: 56 (descr., W. Afr.) ; Bull. U.S. nat. Mns.,
100 {13), 1941: 178 (descr., refs. for west. Pacif., Australasia, Indian Ocean, Atlant.); Arqu. Zool.
Estado Sao Paulo, 3, 1942: 128 (listed, Brazil).

Eulomia glaucus Marchand, Mar. biol. Rep. Cape Town, 2, 1935: 36 (S. Afr.).

Prionace mackei Phillipps, N. Z. J. Sci. Tech., 16, 1 93 5: 238 (descr.. New Zealand).***

Glyfhisnuickei Fowler, Bull. U.S. nat. Mus., 100 (15), 1941: 181 (based on Phillipps, 1935).

Carcharhinus mackei Whitley, Fish. Aust., /, 1940: 106 (descr.. New Zealand, Tasmania, Australia).

28. The name Galeus glaucus was first proposed by Valmont (Diet. Hist. Nat., i, 1768: 371) and next by Duhamel
(Traite Gen. Peches, 3 [9], 1777: 298, pi. 19, fig. 6), but by ruling of the International Commission on Zoo-
logical Nomenclature neither Valmont's nor Duhamel's names are to be taken into consideration (see footnote
I, p. 214), for if binomial they are so only accidentally.

28a. See footnote 25, p. 289.

292 Memoir Sears Foundation for Marine Research

Probable References:

Carc/iarias obscurus Storer, Proc. Boston. Soc. nat. Hist., 2, l8jg: 256 (teeth, Massachusetts Bay) ; Mass. Zool.
Bot. Surv., Rept., Fish., 1839: 184 (same as preceding); Mem. Amer. Acad. Arts Sci., N. S. 9, 1867:
219, pi. 36, fig. 2; Fishes Mass., 1867: 243, pi. 36, fig. 2 (descr., ill., Provincetown, Massachusetts, not
Squalus obscurus Lesueur, 1 81 8).

Carcharias gracilis Philippi, An. Univ. Chile, 71, 1 887: 539, pi. 2, fig. 1 (Chile)."

Genus Scoliodon Miiller and Henle, 1837*

Scoliodon Muller and Henle, S. B. Akad. Wiss. Berlin, 1837: 114; Arch. Naturg., (3) i, 1837: 397, without
tjpe or included species; Plagiost. 1 841: 28; type species, Carcharias {Scoliodon) laticaudus Muller and
Henle, India, designated by Gill, Ann. N. Y. Lye, 7, 1862: 401 ; equals Carcharias sorrakowah Cuvier,
1817.

Generic Synonyms:

Squalus (in part) Richardson, Fauna Boreal. Amer., 5, 1836: 289; and subsequent authors; not Squalus

Linnaeus, 1758.
Carcharias (in part) Cuvier, Regne Anim., 2, 1817: 388; and subsequent authors; not Carcharias Rafinesque,

1810.
Cynocefhalus Bleeker, Verh. Akad. Wet. Amst., 18, 1879: 2; type species, Carcharias {Scoliodon) macro-

rhynchus Bleeker, 1858, E. Indies; not Cynocefhalus Gill, 1861, which equals Prionace (see footnote

3, p. 280).
Carcharinus (in part) Adams and Kendall, Bull. U.S. Bur. Fish, p, 1891 : 292, 308; and subsequent authors; not

Carcharhinus Blainville, 1 816.
Rhizofrion Ogilby, Mem. Qd. Mus., 3, 1915: 132; type species Carcharias {Scoliodon) crenidens Klun-

zinger, 1 880, Australia, equals Carcharias falasorrah Cuvier, Regne Anim., 2, 1829: 338; preoccupied

by Jourdan, 1861, for fossil mammals (Cetacea).
Rhizofrionodon Whitley, Aust. Zool., 5, 1929: 354; substitution for Rkizofrion Ogilby, 1915, preoccupied

by Jourdan, 1 86 1, for fossil mammals (Cetacea).

Doubtful Synonym :

Lamna (in part) Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1 846: 504 (Storer's L. pinctata appears to have
been a combination of Lamna nasus and Squalus functatus Mitchill, 1 81 5, which has sometimes been
thought to have been Scoliodon terrae-novae Richardson, 1836; see Synonyms, p. 303); not Lamna
Cuvier, 1817.

Generic Characters. No spiracles; anal not more than about twice as long at base as 2nd
dorsal j 2nd dorsal not more than 40% as long at base as ist dorsal; midpoint of base of
1st dorsal about equidistant between origin of pelvics and axil of pectoral (sometimes a
little nearer one than the other) ; labial furrow either confined to corner of mouth or ex-
tending out onto one jaw or both; caudal peduncle with a triangular precaudal pit below as
well as above, but without lateral longitudinal ridges; gill openings short, the length of
the longest only about equal to the diameter of eye; anterior margin of nostril with a

29. The forward position of the first doisal fin in Phllippi's (1S87) illustration of his gracilis makes it doubtful
whether or not it is identical with glaucus, which it otherwise resembles.

I. The fossil genus Alofiofsis Lioy (Atti Soc. ital. Sci. nat., 8, 1865: 398, pi. 4) is included in the synonymy of
Scoliodon by Fowler (Bull. U.S. nat. Mus., 100 [13], 1941 : 131). But its teeth, as pictured, are quite difierent
from those of Scoliodon.

Fishes of the Western North Atlantic 293

small lobe; teeth alike in the 2 jaws, erect, narrow-cusped in front, but broad and strongly
oblique along sides of jaws, their outer margins deeply notched, their edges smooth or
slightly wavy at base, their bases not swollen. Characters otherwise those of the family.

Remarks. These are small, warm-water sharks, seldom if ever encountered far from
land. In some localities they are the most abundant sharks. All are fish-eaters, so far as
known, and are entirely harmless.

Range. Oaastal waters in tropical and warm-temperate seas; Morocco to Cameroon;
North Carolina (accidentally to Bay of Fundy) to Uruguay in the Atlantic; Mexico to
Panama in the eastern Pacific; China and Japan to Australia in the western Pacific; Indian
Ocean (including Red Sea and Arabian Gulf) south to Natal.

Fossil Teeth. Eocene to Miocene, Europe.

Species. Of the dozen or so named forms that fall in Scoliodon, as defined here,
all but one (possibly two) are Indo-Pacific and represent not more than eight good species
at most. And it is likely that critical comparison of collections from different seas would
result in a further reduction, because terrae-novae of the Atlantic, the only species of which
a large series has been examined, shows considerable variation (see discussion, p. 299);
hence, others may also. Furthermore, the differences that now seem to be diagnostic are
so inconspicuous, and the several supposed species all resemble one another so closely in
general appearance, that identification of individual specimens calls for close examination
if they happen to be from regions where more than one kind is to be expected. For this
reason it is not yet possible to define the ranges of any of the Indo-Pacific species in detail.

Tentative Key to Species*

I a. Origin of 2nd dorsal posterior to base of anal; labial furrow confined to comer of
mouth, not extending inward along either jaw. dumerilii Bleeker, 1 8 56.

East Indies, southern China.

lb. Origin of 2nd dorsal over rear part of base of anal ; labial furrow extends inward from
comer of mouth for some distance along one jaw or both.

2a. Lower labial furrow considerably longer than upper, which is very short; base of
anal about twice as long as base of 2nd dorsal. sorrakowah Cuvier, 1 829.

India, Malaysian region, China,
Japan.
2b. Upper labial furrow at least as long as lower, if latter is present; base of anal less
than twice as long as base of 2nd dorsal.
3a. Origin of 2nd dorsal definitely anterior to rear end of base of anal.

4a. A short labial furrow on upper jaw directed outward at right angles to
the jaw; none on lower jaw. jordani Ogilby, 1908.

Australia.

2. Carcharias falasorrah of Cuvier (Regne Anim., 2, 1829: 388), commonly referred to Scoliodon, falls in
Hyfofrion 3» here defined, its teeth being conspicuously serrate or denticulate at the base on the outer side.

294 Memoir Sears Foundation for Marine Research

4b. A labial furrow on lower jaw as well as on upper.

5a. Origin of ist dorsal over inner corner of pectoral when latter is laid
back, or a little anterior to it.

6a. Lower labial furrow nearly or quite as long as upper 5 distance
from tip of 2nd dorsal to upper precaudal pit only % to % as
long as horizontal diameter of eye. vagatus Garman, 1 9 1 3.

Zanzibar.

6b. Upper labial furrow considerably longer than lower; distance
from tip of 2nd dorsal to upper precaudal pit longer than hori-
zontal diameter of eye.

7a. Upper labial furrow only about V2 to % as long as hori-
zontal diameter of eye.

terrae-novae Richardson, 1836, p. 295.°
7b. Upper labial furrow as long as horizontal diameter of eye,
or a little longer.

8a. Distance from tip of 2nd dorsal to upper precaudal pit
only about as long as horizontal diameter of eye ; upper
labial furrow about 1V3 times as long as horizontal
diameter of eyej lower furrow only about V2 as long
as upper. longurio Jordan and Gilbert, 1882.

Eastern Pacific, Mexico to Panama.
8b. Distance from tip of 2nd dorsal to upper precaudal pit
about I V2 times as long as horizontal diameter of eye;
upper labial furrow only about as long as diameter of
eye; lower furrow about % as long as upper.

intermedius Ga.rma.n, 191 3.
Philippines, East Indies.*
5b. Origin of ist dorsal a little posterior to inner corner of pectoral when
latter is laid back. longmani Ogilby, 1 9 1 2.

Australia.*

3b. Origin of 2nd dorsal over rear end of base of anal.

walbeehmi^lt&k&T, 1856.
Indian Ocean south to Natal,
Malaysian region, southern Chi-
na, Formosa, Japan.*

5. Including lalandii Muller and Henle, 1841.

4. Intermedius, longmani and <walbeehmi resemble one another so closely that it is doubtful whether they actually
represent more than one rather variable species. We have studied the type specimens of intermedius. The speci-
mens recorded by Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 112) as tualbeehmi show the diagnostic
characters of sorrakowah, with a specimen of which (identified by Garman) we have compared them. We have
also examined the type of vagatus, specimens of longurio and the extensive series of terrae-novae listed on p. 295.

Fishes of the Western North Atlantic 295

Scoliodon terrae-novae (Richardson), 1836

Sharp-nosed Shark

Figures 49, 50

Study Material. 115 specimens, 175 to 930 mm. long, from Uruguay, Brazil, Vene-
zuela, Cuba, the Bahamas, Texas, Alabama, Florida, South and North Carolina, and one
from Grand Manan Island at the mouth of the Bay of Fundy (Harv. Mus. Comp. Zool.,
U.S. Nat. Mus. and Bingham Oceanogr. Coll.); also two, about 316 mm. long, from
Ashantee, tropical West Africa (U.S. Nat. Mus., No. 42212, 42247).

Distinctive Characters. S. terrae-novae is easily separable from such of the other
West Atlantic members of its family as lack spiracles and have the ist dorsal far forward,
by the presence of well marked labial furrows around the corners of the mouth and inward
along both jaws, and by its teeth.

Description. Broad snout type. Proportional dimensions in per cent of total length.
Male, 522 mm., from Rio de Janeiro, Brazil (Harv. Mus. Comp. Zool., No. 160).
Female, 783 mm., from the Bahamas (Harv. Mus. Comp. Zool., No. 1144).

Trunk at origin of -pectoral: breadth 9.4, 10.6; height 9.0, i i.i.

Snout length in front of: outer nostrils 4.8, 4.0 j mouth 7.7, 7.5.

Eye: horizontal diameter 2.4, 2.2.

Mouth: breadth 7.3, 7.2; height 4.4, 5.3.

Nostrils: distance between inner ends 4.9, 5.4.

Labial furrow lengths : upper 1.8, 1.8; lower 1.6, 1.4.

Gill opening lengths : ist 1,9, 1.9; 2nd 2.3, 2.3; 3rd 2.5, 2.4; 4th 2.3, 2.4; 5th

1.9, 1.8.

First dorsal fin: vertical height 8.0, 9.1 ; length of base 8.6, 9.0.

Second dorsal fin: vevticzWiQight l.i, 2.6; length of base 2.9, 3.5.

Anal fin: vertical height 2.5, 3.2; length of base 4.5, 5.3.

Caudal fin: upper margin 27.8, 25.6; lower anterior margin 10.7, 11.2.

Pectoral fin: outer margin 13.O, 14.0; inner margin 4.8, 5.1; distal margin 9.1,

1 1.4.

Distance from snout to: ist dorsal 29.8, 31.7; 2nd dorsal 59.8, 63.3; upper

caudal 72.2, 74.4; pectoral 20.5, 20.6; pelvics 44.2, 46.3; anal 57.0, 60.5.

Interspace between: ist and 2nd dorsals 21.7, 24.9; 2nd dorsal and caudal 8.1,

7.8; anal and caudal 8.6,9.1.

Distance from origin to origin of: pectoral and pelvics 24.4, 24.7; pelvics and

anal 13.4, 13.3.
Narrow snout type. Proportional dimensions in per cent of total length. Male, 581 mm.,
from Rio de Janeiro, Brazil (Harv. Mus. Comp. Zool., No. 412). Female, 608 mm., from
Rio de Janeiro, Brazil (Harv. Mus, Comp. Zool., No. 91).

Trunk at origin of pectoral: breadth 9.2, 10.8; height 9.6, 10.3.

Snout length in front of: outer nostrils 4.8, S-S'^ mouth 7.8, 8.6.

Eye: horizontal diameter 2.2, 2.3.

296 Memoir Sears Foundation for Marine Research

Mouth: breadth 6.2, 6.75 height 5.2, 5.1.

Nostrils: distance between inner ends 4.6, 4.6.

Labial furrow lengths: upper 1.8, 2.2; lower 1.5, 1.5.

Gill opening lengths: ist 1.9, 2.1; 2nd 2.1, 2.2; 3rd 2.2, 2.3; 4th 2.2, 2.3; 5th

1.9,2.1.

First dorsal fin: vertical height 7.8, 8.4; length of base 9.1, 8.1.

Second dorsal fin: vertical height 2.1, 2.2; length of base 2.7, 3.0.

Anal fin: vertical height 2.4, 2.8; length of base 4.7, 5.0.

Caudal fin: upper margin 24.4, 25.5; lower anterior margin 9.7, ii.O.

Pectoral fin: outer margin 12.6, 13.5; irmer margin 5.0, 5.7; distal margin 9.3,

1 0.0.

Distance from snout to: ist dorsal 31.2, 32.7; 2nd dorsal 64.9, 64.O; upper caudal

75.6, 74.5; pectoral 22.2, 22.0; pelvics 46.5, 47.2; anal 61.5, 61.O.

Inters-pace between: 1st and 2nd dorsal 24.6, 25.8 ; 2nd dorsal and caudal 7.7, 8.4;

anal and caudal 8.1, 8.8.

Distance from origin to origin of: pectoral and pelvics 25.2, 28.3 ; pelvics and anal

15.1,15.8.

Figure 49. Scoliodon terrae-novoc. A Female, about 783 mm. long, from the Bahamas (Harv. Mus. Comp.
Zcol., No. 1 144). B Anterior part of head of same from below, about 0.4 x. C Left-hand nostril, about 2.2 x.
D Head of another specimen with relatively longer snout, 6o8 mm. long, from Rio de Janeiro, Brazil, about
0.55 X (Harv. Mus. Comp. Zool., No. 91). See discussion p. 299.

Fishes of the Western North Atlantic

297

Trunk rather slender, its height at origin of ist dorsal (where highest) about V&
of length to origin of caudal. No mid-dorsal ridge. Body sector to cloaca about as long as
tail sector. Caudal peduncle about % to % as thick as deep. Upper and lower precaudal
pits well developed as triangular furrows, the upper the larger. Dermal denticles very
small (aver. 0.17 x 0.17 mm. in specimen 610 mm. long), close-spaced, usually overlap-
ping, their blades about as broad as long, usually with 5, but sometimes with only 3, low
keels, their posterior margins with as many teeth, the median somewhat the longest, on
short pedicels.

Head (to 5th gill opening) a little less than V4 of total length, its dorsal profile only
slightly convex. Snout flattened above and rather thin toward tip, varying in shape from
broadly to more narrowly ovate, its length, in front of a line connecting inner corners of
nostrils, also varying from a little shorter than the distance between the inner ends of the
latter to nearly i V2 times that long." Eye nearly circular, its anterior edge a little posterior
to front of mouth, or nearly opposite latter, its diameter nearly or quite V2 as long as dis-
tance between inner corners of nostrils. Gill openings evenly spaced, the 3rd (slightly
longest) a very little longer than diameter of eye, the 5th slightly the shortest, the 4th
above origin of pectoral. Nostril strongly oblique, its inner corner varying with length of
snout from a little less to a little more than V2 as far from front of mouth as from tip

Figure 50. ScoUodon terrae-novae, illustrated in Fig. 49. A Dermal denticles, about 22 x. B Apical view
of dermal denticle, about 70 x. C Upper and lower teeth, left-hand side, about 3 x. Z) Fourth upper tooth.
E Tenth upper tooth. F Fourth lower tooth. G Eighth lower tooth. D-G, about 6 x.

5. For further comments on this variation,. as regards the relationship of the supposedly long, narrow-mouthed form
Ulandii to the shorter, broader-snouted terrae-novae, see remarks, p. 199.

298 Memoir Sears Foundation for Marine Research

of snout, its anterior margin with a short, blunt-tipped, finger-like lobe near the inner
corner. Mouth ovate, about % to % (60 to 83%) as high as broad, occupying about %
of breadth of head. Labial furrow extending around corner of mouth and onto each jaw,
the upper furrow averaging a little more than % as long as the diameter of eye,° the lower
averaging about % as long as upper.

Teeth usually i^=^=^, similar in the 2 jaws, except that the lowers are a little smaller
than the uppers, with triangular cusp; median upper tooth and ist lower tooth usually
erect, symmetrical and smaller than those on either side, but those along sides of jaws
increasingly oblique toward corners of mouth ; inner margins slightly concave, outer mar-
gins deeply notched about midway toward base, with the basal sector strongly convex, the
edges smooth, or at most slightly wavy, basally, on the outer side} the lOth to 12th succes-
sively smaller than 2nd to 9th.

First dorsal originates about over inner corner of pectoral when latter is laid back or a
little anterior to it, the midpoint of its base varying from a little nearer to axil of pec-
toral than to origin of pelvics to a little nearer to the latter than to the former, its ante-
rior margin only weakly convex, apex subacute, its posterior margin moderately concave
basally, its free rear corner slender and a little more than V3 as long as its base, its rear tip
over, or a little anterior to, origin of pelvics. Second dorsal on an average only about Vs
as long at base as ist, relatively lower, its apex broadly rounded, its free rear tip much more
slender and elongate, being about i V2 times as long as its base, its origin about over, or a
little posterior to, midpoint of base of analj the distance from rear tip of 2nd dorsal to
precaudal pit about i V4 to i Vs times as long as diameter of eye. Caudal about V4 of total
length, with moderately raised axis, narrowly rounded tip and deep subterminal notch, its
terminal sector about V3 the length of fin, its lower anterior corner forming a definite tri-
angular lobe with subacute tip, a little more than Vs as long as the upper, each measured
from the respective precaudal pit. Anal similar to 2nd dorsal in shape, but averaging about
I V2 times as long at base, hence considerably larger in area. Pelvics a little longer at base
than anal, with nearly straight anterior margins, weakly concave distal margins, rounded
apices and subacute tips. Pectoral only about as long as length of ist dorsal along its outer
margin and smaller than the latter in area,' a little more than V2 as broad as long, its
anterior margin weakly convex, distal margin nearly straight toward tip but moderately
concave toward inner corner, the latter subacute, the apex more rounded.

Color. Brownish to olive-gray above, white below and along rear margins of pec-
torals; dorsals and caudals more or less dark-edged, the 2nd dorsal and lower lobe of
caudal the most widely so, especially in small specimens.

Size. Commonly these sharks are about 26 to 30 inches long when mature, rarely
growing much larger than 36 inches, the greatest length definitely recorded for a West

6. Average 82 per cent; extremes 64 and 1 10 per cent.

7. In most of the Carcharhinidae the pectoral is considerably longer than the ist dorsal, and larger in area.

Fishes of the Western North Atlantic 299

Atlantic specimen being only about 36V2 inches or 930 mm. (Harv. Mus. Comp. Zool.,
No. 702, from Rio de Janeiro, a male with large claspers).'

Developmental Stages. The eggs, in early development, are enclosed in thin yellow
shells with pointed ends and are imbedded in crypt-like depressions in the walls of the
maternal uteri. A preliminary account" suggests that a placental connection later develops
between yolk-sac and mother, i.e., that the shark is truly viviparous, as are its close rela-
tives, S. sorrakowah and S. walbeehmi of the Indian Ocean."

Size. Newborn specimens are usually about 275 to 400 mm. long." It appears that
some males may mature when only perhaps 600 mm. long, for we have seen one of 650 mm.
with claspers 61 mm. long (Rio de Janeiro); but in another of 660 mm. from the same
locality they were only 52 mm. long, while in two others of about the same size (642 and
650 mm.) from Florida they were 33 mm. and 30 mm. long respectively. A 20-inch speci-
men from Haiti weighed three pounds.

Remarks. Opinions have differed as to whether or not the form with the longer and
more pointed snout deserves recognition as a distinct species (Jalandii Muller and Henle,
1841; see Synonyms, p. 301). Examination of the extensive series above (p. 295) shows
that an unbroken gradation occurs from those with longer, narrower snouts to those with
shorter and broader snouts. Since we have not been able to draw any sharp line between
them in this or in any other respect, the two extremes are included here under the one
specific name. But the situation still remains somewhat obscure, for while the broader-
snouted specimens appear to be the more common throughout the latitudinal range of the
combined species, north to south, the range of the narrower-snouted members appears to
be definitely restricted to warmer waters, being recorded only for the southern part of the
Gulf of Mexico, West Indies (Martinique, Guadeloupe), Pernambuco and Rio de Janeiro.
This raises the interesting question whether the two forms may not represent two species
which were originally distinct and with distinct ranges, but which have so hybridized (their
ranges having overlapped) that it is not possible to distinguish between them now."

8. A reputed len^h of 2,135 mm. (Fowler, Bull. Amer. Mus. nat. Hist., 70 [1], 1936: 45) is so much larger than
the usual run of adults as to suggest an error.

9. Gudger, Science, N. S. ^/, 1915 : 4.39.

10. For accounts of the placental cord in these, see Southwell and Prashad (Rec. Indian Mus., 16, 1919: 223, pi. 17,
fig. I, 2, 4, 7, 8 [lualbeehmt], and 225, pi. 17, fig. 6, 9, 10 [sorrakoiuahY) 5 see also Thillayampalam (Indian
zool. Memoir 2, Lucknow, Scoliodon, 1928: 107, fig. 93 \^sorrakowahY) .

11. A series of eleven newly bom specimens from Texas, with traces of the umbilical scar still visible, range from
280 to 407 mm. in length.

12. In twelve specimens with the broadly-rounded snout, 410 to 930 mm. in total length, measurements are: distance
from tip of snout to outer corner of nostril 67 to 76% (average 71%) of the distance between outer nostrils;
width of head at outermost part of nostril 95 to 103^ (average 99%) of length of snout in front of mouthj
shortest distance from inner end of nostril to mouth 42 to 52% (average 45%) of length of snout in front of
mouth; distance between nostrils 65 to 73% (average 69%) of length of snout in front of mouth.

In seven specimens with narrowly rounded snout, 544 to 660 mm. in total length, the measurements are:
distance from tip of snout to outer end of nostril 77 to 87% (average 83%) of distance between outer ends
of nostrils; width of head to outer end of nostril 78 to <)ofo (average 84%) of length of snout in front of
mouth; shortest distance from inner end of nostril to mouth 33 to 38% (average 35%) of length of snout
in front of mouth; distance between nostrils 51 to 59% (average s^fo) of length of snout in front of mouth.

300 Memoir Sears Foundation for Marine Research

Habits. This little shark is often taken along the beach, even in the surf, as well as in
harbors and partially enclosed sounds and estuaries. In fact, so far as we are aware it has
never been reported more than a mile or two out from the land or from water more
than a few fathoms deep. It occurs in brackish water in Mississippi (Pascagoula River) and
even in tidal fresh water elsewhere.

It feeds chiefly on small fish that may be available locally; in North Carolina waters,
for example, its stomach is often full of menhaden {Brevoortia) ; parrotfish have been
found in its stomach in Haitian waters. It is also known to eat shrimps and moUusks, and
it bites readily on almost any bait.

It is probable that young are born chiefly in late spring and summer in the northern
sector of its range, for newborn specimens still showing traces of the umbilical scar have
been reported from Florida in July, when they are common in Texan waters also;" they
are abundant off the mouth of the Mississippi in August, and in June and July in North
Carolina waters, where gravid females containing both eggs and late-term embryos are
reported in August. All that is known of its breeding in more tropical waters is that newly
born specimens have been reported from Haiti in early April, and that pregnant females
with as many as twelve embryos are taken around Cuba.^*

Relation to Other Species. It closely resembles S. longurio Jordan and Gilbert of the
Pacific coasts of Mexico and Panama, but it is separable from the latter by the facts that
its upper labial furrow is definitely shorter than the diameter of the eye (as long, or longer
in longurio) and that it has only 25 rows of teeth in the upper jaw (27 to 29 rows in
longurio').

Relation to Man. The only commercial value of this little Shark is that some are
sold in fish markets in the West Indies and perhaps in South America. On the other hand,
its habit of taking the bait intended for better fish makes it a great nuisance to the fisher-
men at times and places where it is numerous.

Range. Both sides of the tropical and subtropical Atlantic; Morocco to Cameroon and
the Cape Verde Islands in the east; Uruguay to North Carolina in the west, and north
accidentally to the mouth of the Bay of Fundy.

Occurrence in the Western Atlantic. The chief center of abundance of this Shark
appears to lie in the West Indian-Caribbean region and in the Gulf of Mexico, whence it
has been recorded at many localities" as plentiful. For example, considerable numbers are
caught by the Louisiana shrimp-trawlers, and it is present throughout the year around
southwestern Florida and among the Keys. However, to the northward it is chiefly a sum-
mer visitor only, present in abundance off the mouth of the Mississippi from June until
September, and the commonest summer shark along South Carolina and the southern part

13. One small collection of sharks taken in the vicinity of Galveston, Texas, in July included eleven S. terrae-novae,
ranging in size from 280 to 407 mm., all with a more or less conspicuous umbilical scar.

14. Personal communication from Luis Howell-Rivero.

15. Venezuela, Yucatan, Colon, Curasao, Trinidad, Martinique, Guadeloupe, Puerto Cabello, Saba, St. Croix,
Jamaica, Cuba, Haiti, Bahamas, Florida Keys, western and northwestern Florida, Mississippi, Louisiana, Texas.

Fishes of the Western North Atlantic 301

of the coast of North Carolina. In some years {e.g., in 1891) it is taken in some numbers
even in winter as far north as Cape Lookout. But it has been recorded only once at the
mouth of Chesapeake Bay and not at all within the Bay; and it reaches New Jersey and the
vicinity of New York only rarely, four specimens being reliably recorded. Occasional
specimens do wander even farther to the northward at rare intervals, for several were taken
near Woods Hole in the summer of 1 9 1 6, while the collection of the Harvard Museum
of Comparative Zoology contains a specimen taken at Grand Manan Island at the mouth
of the Bay of Fundy in 1857 by A. E. Verrill. Early reports of it from Newfoundland
are unfounded."

To the southward it occurs in abundance along the coast of Brazil as far as Rio de
Janeiro and Rio Grande do Sul, and the collection of the Harvard Museum of Compara-
tive Zoology contains one taken many years ago at Maldonado, Uruguay. But apparently
the estuary of the Rio de La Plata marks the southern limit to its usual range in that direc-
tion, for it has not been recorded from Argentina. Neither is it known at Bermuda.

Synonyms and References:

Squalus (CarcAarias) terraf-novae Richardson, Fauna Boreal. Amer., j, 1836: 289 (locality given as "New-
foundland" for specimen received from Audubon, but probably either Florida or South or North
Carolina; see footnote 16, p. 301).

Carcharias (Scoliodon) lalandii Muller and Henle, Plagiost., 1 841: 30 (descr., Rio de Janeiro, Martinique,
Guadeloupe); Dumeril, Hist. Nat. Poiss., 1865: 346 (descr., Brazil).

Lamna terrae-novae (in part)"^' Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 504 (in synopsis).

ScoUodon lalandii Castelnau, Anim. Nouv. Rares Amer. Sud, 1855: 100 (Brazil); Hasse, Naturl. Syst.
Elasm. besond. Theil, 1882: 268, pi. 39, fig. 1-4 (vertebrae); Garman, Mem. Harv. Mus. comp.
Zool., 36, 1913: 113 (descr.); Fowler, Proc. Acad. nat. Sci. Philad., 69, 1917: 128 (Colon); Meek
and Hildebrand, Field Mus. Publ. Zool., 75, 1 923: 53, pi. 2, fig. 2 (descr., discus., Colon) ; Jordan, Ever-
mann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 15 (distrib.) ; White, Bull. Amer. Mus. nat.
Hist., 7^, 1937: 129 (in Key) ; Fowler, Arqu. Zool. Estado Sao Paulo, j, 1942: 128 (Brazil).

ScoUodon terrae-novae Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861: 59 (class., name); Rep. U.S.
Comm. Fish. (1871-1872), 1873: 813 (range); Jordan, Proc. U.S. nat. Mus., 5, 1882: 245 (Pensa-
cola, Florida); Goode and Bean, Proc. U.S. nat. Mus., 5, 1883: 240 (Gulf of Mexico); Jordan and
Gilbert, Bull. U.S. nat. Mus., 16, 1883: 24 (descr., distrib.) ; Proc. U.S. nat. Mus., 5, 1883: 581 (abund.,
S. Carolina); Nelson, Rep. St. Geol. N. Jersey, 2 (2), 1890: 66l (diagn., N. Jersey distrib.); Bean,

B. A., Proc. U.S. nat. Mus., 14, 1891: 94 (C. Charles, Virginia); Jordan and Evermann, Rep. U.S.
Comm. Fish. (1895), 1896: 217 (C. Cod to Brazil); Bull. U.S. nat. Mus., 47 (i), 1896: 43 (descr.,

C. Cod to Brazil); Jordan and Rutter, Proc. Acad. nat. Sci. Philad., 1897: 91 (Jamaica); Evermann
and Bean, Rep. U.S. Comm. Fish. (1896), 1898: 239 (Indian R., Florida) ; Gilbert, Proc. Wash. Acad.
Sci., 2, 1900: 161 (Brazil); Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 48
(Florida) ; Bean, T. H., Rep. For. Comm. N. Y., 1901 : 378 (distrib.) ; Rep. N. York St. Mus., do, Zool.
9, 1903: 29 (descr., general); Jordan and Thompson, Bull. U.S. Bur. Fish., 24, 1905: 232 (Tortugas,
Florida); Linton, Bull. U.S. Bur. Fish., 24, 1905: 342 (food, parasites); Cole and Barbour, Bull.
Mus. comp. Zool. Harv., $0, 1906: 155 (Yucatan) ; Fowler, Rep. N. Jersey Mus. (1905), 1906: 63 (N.
Jersey) ; Rep. N. J. Mus. (1906), 1907: pi. 74 (ill.) ; Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907: 155,

16. "This species, with others belonging to the Florida fauna, is said by Richardson to have ijeen brought from New-
foundland by Audubon. They doubtless came from some locality in Florida or Carolina" (Jordan and Ever-
mann, Bull. U.S. nat. Mus., 4.7 [i], 1896: 43; footnote).

17. This was a combination of Lamna caudata Dekay (equals Carchar/Umu milberti, see p. 376) with ScoUodon
terrae-novat Richardson.

302 Memoir Sears Foundation for Marine Research

200 (descr., refs., Brazil); Linton, Bull. U.S. Bur. Fish., 26, 1907: 122, 125 (parasites); Smith, Bull.
N.C. geol. econ. Surv., 2, 1907: 34 (N. Carolina); Wilson, Proc. U.S. nat. Mus., 55, 1908: 340, 360,
423, 431, 626 (parasites) ; Fowler, Proc. Acad. nat. Sci. Philad., 60, 1908: 66 (Florida) ; Gudger, Amer.
Nat., 44, 1910: 399 (N. Carolina); Linton, Bull. U.S. Bur. Fish., 28, 1910: 1201, pi. 2 (parasites);
Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 115, pi. 2, fig. 1-4 (descr.); Radcliffe, Copeia,
26, 1916: 2 (Woods Hole); Coles, Proc. biol. Soc. Wash., 28, 191 5: 90 (N. Carolina); Gudger,
Science, N. S. 41, 1915: 439 (eggs, uterine attachment of young, Tortugas, Florida); Nichols and
Murphy, Brooklyn Mus. Sci. Bull., 3 (l), 1916: 18 (distrib.) ; Radcliffe, Bull. U.S. Bur. Fish., 34,
1916: 250, pi. 39, fig. 2 (denticles, teeth, N. Carolina) ; Fowler, Copeia, 30, 1916: 36 (E. coast U.S.) ;
Proc. Acad. nat. Sci. Philad., 7/, 1919: 146 (Jamaica); Wilson, Proc. U.S. nat. Mus., 55, 1920: 592
(parasites); Fowler, Proc. biol. .Soc. Wash., 33, 1920: 144 (N. Jersey); Proc. Acad. nat. Sci. Philad.,
72, 1921: 394 (Pensacola, Florida); Wilson, Proc. U.S. nat. Mus., 60 (5), 1922: 60 (parasites); Meek
and Hildebrand, Field Mus. Publ. Zool., ig, 1 923 : 5 5, pi. 2, fig. 3 (descr., discus.. Colon) ; Ribeiro, Fauna
brasil. Peixes, Mus. nac. Riode J., 2(1), Fasc. I, 1923: 12, pi. 4 (Brazil) ; Linton, Proc. U.S. nat. Mus.,
64 (21), 1924: 7, 35, 36, 46, 47, 48 (parasites. Woods Hole, N. Carolina) ; Chabanaud and Monod, Bull.
Etud. Hist. Sci. Afr. Occid. Franc. (1926), 1927: 229 (trop. W. Afr., not seen) ; Monod, Faune Colon-
Franc, 1927: 646 (not seen) ; Beebe and Tee-Van, Zoologica, N. Y., ro, 1928: 27 (Haiti) ; Fowler, Proc.
Acad. nat. Sci. Philad., 80, 1928: 456 (Haiti) ; Hildebrand and Schroeder, Bull. U.S. Bur. Fish., 43 (l),
1928: 49 (descr., C. Charles, Virginia) ; Breder, Field Bk. Mar. Fish. Atlant. Coast, 1 929: 18 (general) ;
Jordan, Manual Vert. Anim. NE. U.S., 1929: 10 (general); Jordan, Evermann and Clark, Rep. U.S.
Comm. Fish. (1928), 2, 1930: 14 (in check list); Wilson, Bull. U.S. nat. Mus., 158, 1932: 439
(parasites); Fowler, Proc. biol. Soc. Wash., 36, 1933: 27 (Bayport, Louisiana); Borri, Atti Soc. tosc.
Sci. nat., 44, 1934: 90 (Bahia, Brazil) ; Burton, Sci. Mon. N. Y., 40, 1935: 283 (abund., S. Carolina) ;
Budker, Bull. Mus. Hist. nat. Paris, (2) 7, 1935: 184 (Dakar, W. Afr.) ; Fowler, Bull. Amer. Mus. nat.
Hist., 70 (i), 1936: 45 (trop. W. Afr.); Bere, Amer. Midi. Nat., 17, 1936: 604 (Florida); White,
Bull. Amer. Mus. nat. Hist., 74, 1937: 129, pi. 13, fig. C (anat., in Key) ; Springer, Proc. Fla. Acad.
Sci., 5, 1939: 18 (Florida) ; Tortonese, Atti Soc. ital. Sci. nat., 77, 1938: 298 (Rio de Janeiro) ; Longley
and Hildebrand, Pap. Tortugas Lab., 34, 1941: 2 (Tortugas, Florida); Gunter, Amer. Midi. Nat.,
28, 1942: 316 (in brackish and tidal fresh water, Mississippi and Maryland); Fowler, Arqu. Zool.
Estad. Sao Paulo, 3, 1942: 128 (listed for Brazil); Lunz, Bull. S. C. St. Planning Bd., 14, 1944: 26
(S. Carolina, Florida) ; Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm.,
Wash., 1945: 128, fig. 46 (descr., habits, range, ill.) ; Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1 945:
94, 263 (Beaufort, N. Carolina, south. Florida Iocs.), 159 (Charleston, S. Carolina).

Carcharias (Scoliodon) terrae-novae Dumeril, Hist. Nat. Poiss., 1 865: 346 (descr., Newfoundland, after
Richardson, 1836); Gunther, Cat. Fish. Brit. Mus., 8, 1870: 360 (W. Indies, Bahia, Brazil); Werner,
Zool. Jb., Syst. Abt., 21, 1904: 283 (Jamaica [? ], N. Carolina) ; Lampe, Dtsch. Sudpol. Exped., 75, Zool.
7, 1 91 4: 204, 213 (C. Verde Is.) ; Metzelaar, Trop. Atlant. Visschen, 1 91 9: 186 (trop. W. Afr.).

Carcharias (Scoliodon) acutus Steindachner, S. B. Akad. Wiss. Wien, 61 (l), 1870: 575 (Senegambia) ;"
Denkschr. Akad. Wiss. Wien, 44, 1882: 51 (Senegambia) ; Osorio, Mem. Mus. Bocage Lisbon, I, 1909:
77 (C. Verde Is.); Metzelaar, Trop. Atlant. Visschen, 1919: 186 (Senegambia); not Carcharias acutus
Ruppell, 1835, which is a synonym of Hyfop'ion falasorrah (Cuvier) ; see footnote 2, p. 293.

Carcharhinus {Scoliodon) terrae-novae Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 796 (distrib.).

Carcharhinus terrae-novae Adams and Kendall, Bull. U.S. Fish Comm., p, 1 891: 292, 308 (W, Florida);
Henshall, Bull. U.S. Fish Comm., p, 1891 : 384 (SW. Florida) ; Lonnberg, Ofvers. Forhand. Vet Akad.
Stockholm, s^ (3)> 1894: m (Florida Keys); Wilson, Amer. Nat., 34, 1900: 355 (listed, North
Carolina).

Carcharias terrae-novae Jordan and Gilbert, Proc. U. S. nat. Mus., 5, 1883: 245 (Gulf of Mexico) ; Dnerden,
J. Inst. Jamaica, 6, 1899: 614 (Jamaica) ; Schreiner and Ribeiro, Arch. Mus. nac. Rio de J., 12, 1903: 79
(Brazil) ; von Ihering, Rev. Mus. paul., 2, 1897: 34 (Brazil).

Carcharias eumeces Pietschmann, Jb. nassau. Ver. Naturk., 1 91 3: 172 (descr., Cameroon, trop. W. Afr.).

18. Steindachner's excellent and detailed description leaves no doubt that his West African specimens were actually
terrat-novae.

Fishes of the Western North Atlantic 303

Scoliodon eumeces Fowler, Proc. U.S. nat. Mus., 56, 1919: 249 (trop. W. Afr.).

Scoliodon acutus Fowler, Bull. Amer. Mus. nat. Hist., 70 (i), 1936: 46; not Carcharias acutus Ruppell, 1835.

Probable Synonyms and References:

Squalus forosus Yoty, Memorias, 2, i860: 339, 452, pi. 19, fig. 1 1, i 2 (teeth, Cuba).'"

Scoliodon forosus Poey, Repert. Fisico-Nat. Cuba, 2, 1868, 452; An. Soc. esp. Hist, nat., 5, 1876: 396; Enum-
erat. Pise. Cubens., 1876: 200 (Cuba); Fowler, Fish Culturist, 21 (9), 1942: 66 (size of mature fe-
male, and embryos, Cuba) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1 930: 15
(Cuba); Fowler, Proc. Acad. nat. Sci. Philad., 55, 1931: 391 (Trinidad).

Doubtful References:

Squalus pinctatui Mitchill, Trans. Lit. Phil. Soc. N.Y., 1815: 483 (near New York); not Squalus functatus
Bloch and Schneider, 1801.

Lamna functata (in part) Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 504 (this appears to be a com-
bination of Squalus functatus Mitchill with Lamna nasus) .

Lamna pinctata Llnslcy, Amer. J. Sci., 47, I 844: 76 (name only, Connecticut).

Ap-ionodon functatus Gill, Proc. Acad. nat. Sci. Philad., Addend., 1 861 : 59 (in synopsis).

Carcharias forosus Goeldi, Bol. Mus. Paraense, 2, 1898: 488 (Brazil).

Carcharias (Scoliodon) walbenii Osorio, J. Sci. math. phys. nat. Lisboa, (2) 5, 1 898: 200 (C. Verde, St.
Thome); Metzelaar, Trop. Atlant. Visschen, 1919: 186 (C. Verde); not Carcharias (Scoliodon)
walieehmi Bleekcr, 1856.

Carcharias longurio Engelhardt, Zool. Anz., 39, 1912: 648 (St. Thomas, W. Indies); not Carcharias (Scolio-
don) longurio Jordan and Gilbert, I 888.

Probably not Scoliodon forosus Garman, Mem. Harv. Mus. comp. Zool., 36, 1 91 3: 112 (descr., Cuba).^"*

Genus Aprionodon Gill, 1861

AfrionodonG\U,Tioc. Acad. nat. Sci. Philad., Addend., 1861: 59; Ann. N. Y. Lye, 7, 1862: 400, 401, 41 1;
substitution for Afrion Miiller and Henle, 1841, preoccupied by Cuvier and Valenciennes, 1 830, for
bony fishes, and by Audinet-Serville 1839, for insects; type species, Afrionodon functatus Gill, equals
Carcharias (Afrion) isodon Miiller and Henle, 1841.'

Generic Synonyms:

Carcharias (in part) Ruppell, Neue Wirbelt. Abyssinia, Fische, 1835: 65; and subsequent authors; not Car-
charias Rafinesque, I 8 1 0.

Afrion Muller and Henle, Plagiost., 1841: 31; type species, Carcharias (Afrion) brevifinna Muller and
Henle, Java; not Afrion Cuvier and Valenciennes, 1 8 30, for bony fishes.

Squalus (in part) Gray, List Fish. Brit. Mus., I 85 1 : 41 ; not Squalus Linnaeus, 1758.

Carcharhinus (in part) Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 796; not Carcharhinus Blainville, 1816.

Longmania Whitley, Aust. Zool., p, 1939: 231; type species, Carcharias (Afrion) brevifinna Muller and
Henle, 1841.

19. Porosus is classed as a probable synonym of terrae-novae on the strength of Poey's descriptions of it and
his illustrations of its teeth. Although a pliotograph of an unpublished drawing by him of the lower side of its
head fails to show any labial furrows on the lower jaw, this may have been an oversight.
19a. Probably a Carcharhinus, not a Scoliodon, because described as with weakly serrated teeth and wider gill open-
ings. The specimen on which he based his account is no longer in existence.
I. Of the two species included by Gill (1861), Carcharias (Afrion) isodon Muller and Henle must necessarily be
taken as the type and not Squalus functatus Mitchill, 1815, which was designated by Jordan (Genera Fish., 3,
1919: 303), because Afrionodon was obviously a substitution for Afrion. Furthermore, the name Squalus
functatus, having been long antedated by Bloch and Schneider (1801), would not be available in the present
connection.

304 Memoir Sears Foundation for Marine Research

Generic Characters. Carcharhinidae with anal less than twice as long at base as 2nd
dorsal; without spiracles; c?.udal peduncle without lateral ridges, but with a precaudal
pit below as well as above; midpoint of base of ist dorsal as near to axil of pectoral as to
origin of pelvics, or nearer; and dorsal only about ^o as long at base as ist and much
smaller in area; teeth slender and symmetrical in both jaws, their bases as well as their
cusps with smooth edges; gill openings notably large, the longest nearly or quite V2 as
long as base of ist dorsal and more than twice as long as horizontal diameter of eye (see
footnote 5, p. 265), the 5th being over origin of pectoral; anterior margin of nostril
only slightly expanded; labial furrow around corner of mouth but extending inward for
only a very <5hort distance onto either jaw, if at all. Characters otherwise those of the
family.

Range. Tropical and subtropical. Senegambia in the eastern Atlantic; North Carolina
(perhaps New York) soutn to Cuba and Texas in the western Atlantic; tropical Indian
Ocean; Red Sea and Arabian Gulf; India; East Indies and Australia, Indo-China, Japan,
Micronesia.

Species. This genus of small sharks, about which little is known, is represented by one
species in the Atlantic and by one in the Indo-Pacific.*

Key to Species*

la. Snout in front of mouth only about % (about 29%) as long as head, or % as long as
from eye to ist gill opening; distance between nostrils a little more than % as long as
snout; pectoral about V2 as long as head. isodon Muller and Henle, 1841, p. 304.

lb. Snout in front of mouth about % (40%) as long as head, and about as long as from
eye to ist gill opening; distance between nostrils ^ as long as snout; pectoral about %
as long as head. brevifinna Muller and Henle, 1 841 .

Arabia, East Indies, Australia, Japan.

Afrionodon isodon (Muller and Henle), 1841
Figure 5 1

Study Material. Two females, 460 and 504 mm., from off Biloxi, Mississippi, and
from Texas (U.S. Nat. Mus.) ; 4 young males, 500 to 567 mm., from Texas (Harv. Mus.
Comp. Zool.).*

Distinctive Characters. Easily recognizable among local Carcharhinidae by its slender,
symmetrical, smooth-edged teeth, very long gill openings and a 2nd dorsal fin that is much
smaller than its ist dorsal.

2. Another Indo-Pacific species, Carcharias acutidens Riippell, 1835, has usually been referred to this genus, but is
placed in Negaprion (p. 309) according to generic definitions adopted here.

3. Carcharias jronlo Jordan and Gilbert (Proc. U.S. nat. Mus., 5, 1882: 102) is also referred to this genus by Beebe
and Tee-Van (Zoologica, N. Y., 28, 1941: 105), but it is placed here in Negafrion because of the large size
of its second dorsal fin; for discussion, see footnote i, p. 300.

4. Contributed by J. L. Baughman.

Fishes of the Western North Atlantic

305

Description. Proportional dimensions in per cent of total length. Female, 504 mm.,
from Galveston, Texas (U.S. Nat. Mus., No. 1 18457). Male, 560 mm., same locality
(Harv. Mus. Comp. Zool., No. 35831).

Trunk at origin of pectoral: breadth 11.3, 10.7; height 11.3, 12.1.

Snout length in front of: outer nostrils 4.1, 3.8; mouth 6.8, 7.1.

Eye: horizontal diameter 1.9, 1.8.

Mouth: breadth 9.1, 8.75 height 5.2, 5.6.

Nostrils: distance between inner ends 5.5, 5.3.

Labial furrow lengths: upper 0.9, 0.8 ; lower 0.8, 0.7.

Gill opening lengths: ist 4.4, 5.4} 2nd 4.7, 5.7; 3rd 4.8, 5.75 4th 4.8, S-^'')

5th 4.3, 4.7.

First dorsal fin: vertical height 9.4, 9.0; length of base lO.o, 9.6.

Second dorsal fin: vertical height 2.8, 2.9 j length of base 4.8, 4.8.

Anal fin: vertical height 3.4, 3.4; length of base 5.2, 5.6.

Caudal fin: upper margin 28.1, 28. 2^ lower anterior margin 1 1.3, 1 1.8.

^L^J^^AAM^^S^^

Figure 51. Afrionoion isodon, female, 504. mm. long, from Texas (U. S. Nat. Mus., No. 1 18457). ^ An-
terior part of head from below. B Left-hand nostril, about 2.5 x. C Dermal denticles, about 34 x. D Apical
view of dermal denticle, about 68 x. E Left-hand upper and lower teeth, about twice natural size; this figure
is inverted by error. F Fourth upper tooth. G Tenth upper tooth. H, I Fourth lower tooth. J Sixth lower tooth.
F-J, about 4 X.

3o6 Memoir Sears Foundation for Marine Research

Pectoral fin: outer margin 14.0, 15.5; Inner margin 4.9, 5.3; distal margin 10.7,

1 1.4.

Distance from snout to: ist dorsal 30. 3, 30. 7 } 2nd dorsal 60. 7, 60. 7 ; upper caudal

71.9,71.8; pectoral 25.0, 24.O; pelvics48.o, 46.6; anal 59.5, 59.7.

Interspace between: ist and 2nd dorsals 19.3, 19.6; 2nd dorsal and caudal 6.6,

j.S; anal and caudal 6.0, 6.4.

Distance from origin to origin of: pectoral and pelvics 23.4, 24. i ; pelvics and anal

12.5,13.4.

Trunk moderately slender, its height at ist dorsal a little less than Vs its length to
caudal pit. No mid-dorsal ridge. Caudal peduncle only slightly compressed, without
lateral ridges. Precaudal pits subtriangular. Body sector to cloaca a little longer than tail
sector. Dermal denticles small, closely overlapping, a little broader than long, their blades
horizontal, broadly oval, with 3 low ridges and as many short teeth, the median only a little
the longest; pedicels slender.

Head about V4 of total length, moderately flattened above and a little broader oppo-
site corners of mouth than in region of gill openings. Snout wedge-shaped, its tip narrowly
rounded, its length in front of mouth about V4 of length of head. Eye approximately cir-
cular, its anterior edge about opposite front of mouth, its horizontal diameter about ^4 as
long as snout in front of mouth. Gill openings 3 and 4 (slightly the longest) about % as
long as snout in front of mouth, 2^/4 times as long as diameter of eye and about as
long as distance between nostrils, the ist and 5th slightly the shortest; the spaces between
successive gill openings about equally broad at upper ends, but those between 3rd and
4th, and 4th and 5th much narrower at lower end, the 5th opening moderately oblique and
over origin of pectoral, the 4th close in front of latter. Nostril strongly oblique, its outer
corner at margin of snout, its inner corner nearer to mouth than to tip of snout by a dis-
tance about equal to horizontal diameter of eye, its anterior margin only very slightly
expanded in subtriangular outline. Mouth broadly rounded in front, about V2 as high as
broad, occupying about % of breadth of head. A well marked labial furrow around corner
of mouth and extending inward for a very short distance on each jaw, the lower usually
concealed when mouth is closed.

Teeth '^ |° '^3^^ [° \\ j similar in the 2 jaws, smooth-edged, symmetrical, with sharp,
slender, erect median cusp without lateral denticles, on broad bases; i small tooth at sym-
physis in upper jaw and 3 in lower, the median minute, as are the outermost 2 teeth in
each jaw.

Origin of ist dorsal a little posterior to axil of pectoral, its anterior margin slightly
convex, its posterior margin deeply concave, its apex rounded, its free rear corner mod-
erately slender and a little less than ^/^ as long as its base, its anterior margin about V2 as
long as head. Second dorsal about % as long at base as ist dorsal, but only V4 as high
vertically, its free rear corner nearly'as long as its base, its origin a little posterior to origin
of anal. Caudal between Vs and % (about 28%) of total length, its axis raised at an angle

Fishes of the Western North Atlantic 307

of about 30°, its upper margin nearly straight, its tip slender and narrowly rounded, its
terminal sector only about \^i the length of fin, the subterminal notch well marked, its
lower anterior corner a narrow-tipped lobe, about 40% as long as upper and with convex
anterior margin. Anal a little longer at base than 2nd dorsal, but about as large in area and
similar in shape except that its posterior margin is much more deeply concave. Pelvics
about as long at base as 2nd dorsal, with nearly straight edges, their apices broadly rounded
and their tips narrowly so, their origin posterior to rear tip of ist dorsal by a distance about
equal to diameter of eye. Pectoral only a little more than Yz (about 56%) as long as
head, and little, if any, longer than anterior margin of ist dorsal, a little more than ^o as
broad as long, the outer margin moderately convex, distal margin moderately and evenly
concave, apex and inner corner narrowly rounded, or subacute.

Color. Slate-blue above and on upper surface of pectorals, shading through grayish
white on lower sides to pure white below j pelvics and anal white.

Size. The few specimens reported so far have ranged between 500 and 747 mm. (20
to 30 inches) in length for the western Atlantic, but up to 1.2 meters (about 4 feet) off
West Africa. The maximum size may be considerably greater, for a male of 747 mm. was
immature.

Developmental Stages. Not known.

Habits. The teeth suggest that this is a fish-eater, like others of its family. All re-
corded specimens have been taken close inshore. Nothing definite is known of its habits or
diet.

Range. Both sides of the Atlantic; Senegambia, West Africa, in the east; Cuba,
Texas, off Biloxi, Mississippi, Southwest Florida, South and North Carolina, Virginia
and New York in the west.** It is described as common in Senegambian waters, and several
have been reported from southwestern Florida, from Biloxi on the north shore of the
Gulf of Mexico and from Texas (see Study Material, p. 304). However, the more
northerly records are for single individuals only. The above facts suggest that it is a
tropical species which occasionally strays northward along the east coast of the United
States in summer, as do so many other fishes of warm-water origin.

Synonyms and References:

Carcharias {A-prion) isoion Miiller and Henle, Plagiost., 1841: 32 (descr., no locality given for type speci-
men in Paris Museum; but received from Milbert, hence probably New York).

Afrionodon functatus Gill, Proc. Acad. nat. Sci. Philad., Addend., 1 861: 55; and later eds. (name); Ann.
N. Y. Lye, 7, 1862: 401 (name); Proc. Acad. nat. Sci. Philad., 1864: 262 (descr., probably N. York
because type spec, from Milbert); Jordan and Gilbert, Bull. U.S. nat. Mas., 16, 1883: 24 (Atlant.) ;
not Squalu! ■punctatus MitchiU, 181 5, which was probably Scoliodon terrae-novae i see p. 292.

Afrionodon isodon Gill, Ann. N. Y. Lye, 7, 1862: 41 1 (name) ; Poey, An. Soc. esp. Hist, nat., 5, 1876: 396;
Enumerat. Pise. Cubens., 1876: 200 (teeth, Cuba) ; Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883:
874 (discus.); Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 217 (Atlant.); Bull. U.S.
nat. Mus., 47 (i), 1896: 42 (descr., N. York, Virginia, Cuba) ; Bean, T. H., Rep. For. Comm. N. Y.,

5. No locality was given by Miiller and Henle, 1841, for the type specimen, which is in the Paris Museum, but
Dumeril (Hist. Nat. Poiss., 1865: 349) states that it was from the coast of New York state.

3o8 Memoir Sears Foundation for Marine Research

1901: 377; Bull. N. Y. St. Mus., 60, Zool. p, 1903: 28 (descr., N. York, Virginia, Cuba); Garman,
Mem. Harv. Mus. comp. Zool., 36, 1913: 119 (descr.); Fowler, Copeia, 30, 1916: 36 (mid-Atlant.,
U.S.) ; Radcliffe, Bull. U.S. Bur. Fish., ^4, 1916: 252, pi. 39, fig. 3 (meas., teeth, N. Carolina) ; Nichols
and Murphy, Brooklyn Mus. Sci. Bull., 5 (l), 1916: 17 (N. York, Virginia, Cuba); Breder, Field Bk.
Mar. Fish. Atlant. Coast, 1929: 18 (general); Jordan, Manual Vert. Anim. NE. U.S., 1929: 10 (gen-
eral); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 15 (N. York, Virginia,
Cuba) ; Fowler, BuU. Amer. Mus. nat. Hist., 70 (i), 1936: 46 (W. Afr.) ; Bere, Amer. Midi. Nat., 17,
1936: 604 (W. Florida) ; White, Bull. Amer. Mus. nat. Hist., 74, 1937: 129 (in Key) ; Springer, Proc.
Fla. Acad. Sci., 3, 1939: 29 (Florida, off Mississippi); Whitley, Aust. Zool., 9 (3), 1939: 231 (com-
parison with brevifinna Miiller and Henle, 1841); Burton, Copeia, 1940: 140 (S. Carolina, size);
Hildebrand, Copeia, 1941: 221 (N. Carolina, old record); Norris, Plagiost. Hypophysis, 1941: pi. 18,
fig. 72 (brain) ; Lunz, Bull. S. Carolina St. Planning Bd., 14, 1944: 27 (Florida) ; Bigelow and Schroe-
der, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 126, fig. 45 (descr., range,
ill.) ; Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1945: 159 (Morris I., S. Carolina).

Carcharias {Afrionodon) isodon Dumeril, Hist. Nat. Poiss., i, 1865: 349 (descr. of type specimen, coast of
New York); Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 41 ; Faune Senegambie, Poiss., /,
188 3- 1885: 18 (Senegambia, size); Metzelaar, Trop. Atlant. Visschen, 1 91 9: 187 (Senegambia).

Carcharias {Afrionodon) functatus Giinther, Cat. Fish. Brit. Mus., 8, 1870: 361 (N. York, not Squalus
functatus Mitchill, 1 81 5, which probably was Scoliodon terrae-novae ; see p. 292).

Carchorkinus {Afrionodon) isodon Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 796 (W. Indies, accidental
on U.S. coast) .

Genus Negaprion Whitley, 1 940

Negafrion Whitley, Fish. Aust., i, 1940: iii; type species, Afrionodon acutidens queenslandicus Whitley,
Aust. Zool., 9, 1939: 233, Queensland.

Generic Synonyms:

Hyfoprion (in part) Poey, Repert. Fisico-Nat. Cuba, 1868: 451 ; and subsequent authors; not Hyfofrion

Muller and Henle, 1 841 .
Carcharias (in part) Jordan, Bull. U.S. Fish Comm., 4, 1884: 79; Proc. U.S. nat. Mus., 7, 1887: 104; not

Carcharias Rafinesque, 1 8 1 0.
Carcharinus (in part) Henshall, Bull. U.S. Fish Comm., 9, 1891: 383; not Carcharhinus Blainville, 1816.
Afriortodon Whitley, Aust. Zool., 9, 1939: 233, for A. acutidens queenslandicus Whitley; not Afrionodon

Gill, 1 861.

Generic Characters. Carcharhinidae with anal not longer at base than 2nd dorsal;
without spiracles j midpoint of base of ist dorsal at least as near to axil of pectoral as to
origin of pelvics; 2nd dorsal at least % as long at base as ist dorsal; caudal peduncle
without lateral ridges; a precaudal pit above but none below; back without mid-dorsal
ridge; gill openings relatively large, the longest at least ^/4 as long as snout in front of
mouth and more than Yz as long as base of ist dorsal; a labial furrow at comer of mouth
and extending outward a very short distance on upper jaw, but none on lower; teeth erect,
symmetrical in front of mouth but increasingly oblique toward corners of latter, their
cusps smooth-edged, their bases smooth, wavy, or even indistinctly serrate. Characters
otherwise those of the family.

Range. Western Atlantic in tropical and subtropical belt; tropical Indian Ocean j

Fishes of the Wester ti North Atlantic 309

Red Sea and Gulf of Arabia; India; Indo-China; North Australia and Queensland;
Micronesia.

Species. Medium-sized tropical sharks of littoral waters; one species so far known
from the Atlantic; four from the Indo-Pacific.

Key to Species

I a. Snout obtusely wedge-shaped.

2a. Posterior margins of pectorals and pelvics deeply concave.

queenslandicits Whhlty, 1939.
Queensland, Australia.

2b. Posterior margins of pectorals and pelvics only very weakly concave.

fronto Jordan and Gilbert, 1882.
Pacific coasts of Mexico and Costa Rica.^

lb. Snout broadly and evenly rounded.

3a. Bases of teeth, as well as cusps, smooth-edged. odontas f is Yo'^l&v, 1908.

Indian Ocean. ^

3b. Edges of bases of upper teeth at least wavy, irregularly serrate, or denticulate.

4a. Bases of upper teeth with wavy or irregularly serrate edges, those of lower

teeth smooth; distance between outer ends of nostrils only about % as great

as breadth of mouth. brevirostris Poey, 1868, p. 3 lO.

4b. Bases of some of the teeth, upper or lower, with one strong denticle on the

outer side;' distance between outer ends of nostrils equal to breadth of mouth.

acutidens'RxvpT^tW, 1835.

Tropical Indian Ocean, in-
cluding Red Sea (type local-
ity) and Arabian Gulf, India
and Indo-China, Torres Strait,
Micronesia; perhaps Philip-
pines.*

1. Beebe and Tee-Van (Zoologica, N. Y., 26, 1941 : 105) have pointed out that the two specimens on which Jordan
and Gilbert's (Proc. U.S. nat. Mus., 5, 1882: 102) original account of fronto was based represented two different
species: one with narrow-cusped, broad-based teeth and with the second dorsal nearly as large as the first; the
other with small second dorsal and serrate teeth. The second of these was obviously a Carc/iarhinus, perhaps
azureus Gilbert and Starks, 1904, but the first, designated by Beebe and Tee-Van as the type of the species fronto,
falls in Negafrion as defined here, for Beebe and Tee-Van (Zoologica, N. Y., 26, 1941 : 106) found the teeth
of another specimen to be smooth-edged, except where "nicked by some external agency," this last explaining
Jordan and Gilbert's original account of them as appearing minutely serrulated under a lens.

2. Fowler (Bull. U.S. nat. Mus., 100 [13], 1941 : 194) recently has relegated this species to the synonymy of Triae-
nodon obesus Ruppell, 1835. But in his original account of it (Proc. Acad. nat. Sci. Philad., 60, 1908: 63) he stated
that the teeth are not only slender, erect and smooth-edged, but without basal cusps, and he so pictures them,
whereas those of Triaenodon have one or two lateral cusps on each side of the longer median cusp, this being a
family characteristic.

3. According to Mijller and Henle (Plagiost., 1841 : 33) it was the lower teeth that were denticulate at the base in
the specimen (probably the type) that they examined and for which they gave measurements. However, if the
Afrionodon silakaiensis of Herre, 1934 (Herre, Philippine Exped. Fish., 193 i : 11), is identical with acutidens,
as it appears to be, the upper teeth may be so armed.

4. That is, if silakaiensis Herre, 1934, is identical with acutidens; see footnote 3, p. 309.

3IO

Memoir Sears Foundation for Marine Research
Negaprion ifrevirostris (Poey), 1868

Lemon Shark

Figures 52, 53

Study Material. Six specimens, male and female, 610 to 893 mm. (about 24 to 35
inches), from coasts of Texas, Louisiana and Florida (U.S. Nat. Mus. and Amer. Mus.
Nat. Hist.) ; also jaws of a 6-foot lO-inch female from Bay of Florida.

Distinctive Characters. Made easily recognizable among western Atlantic Carcha-
rhinidae by a second dorsal that is nearly as large as the first, with a very broadly rounded
snout, and by its characteristic teeth.

Description. Proportional dimensions in per cent of total length. Female, 610 mm.,

Figure 52. Negafrion irevirostris, immature male, 782 mm. long, from Florida (Amer. Mus. Nat. Hist.).
A Dennal denticles of same, about 26 x. B Apical view of dermal denticles, about 52 x. C Left-hand upper
and lower teeth of a 6-foot lo-inch female from the Bay of Florida (Amer. Mus. Nat. Hist., No. 5735),
about i.i X. Z) Fourth upper tooth. E Tenth upper tooth. F Second lower tooth. G Tenth lower tooth.
D-G, about 2.2 z.

Fishes of the Western North Atlantic

311

from Florida (U.S. Nat. Mus., No. 104332). Male, 782 mm., from Florida (Amer.
Mus. Nat. Hist.).

Trunk at origin of -pectoral: breadth 11.5, 12.8; height 10.8, i i.o.

Snout length in front of: outer nostrils 2.9, 2.9; mouth 5.7, 5.3.

Eye: horizontal diameter 2.1, 1.8.

Mouth: breadth 8.5, 9.3; height 5.1, 5.0.

Nostrils: distance between inner ends 5.6, 5.6.

Gill opening lengths: ist 3.1, 4.OJ 2nd 3.2, 4.1 5 3rd 3.4, 4.1 ; 4th 3.4, 4.O; 5th

3-3, 3-7-

First dorsal fin: vertical height 6.6, 6.9; length of base lO.O, 10.2.

Second dorsal fin: vertical height 5.4, 6.0; length of base 8.2, 7.8.

Anal fin: vertical height 4.3, 5.4; length of base 6.6., 6.4.

Caudal fin: upper margin 23.6, 24.O; lower anterior margin 12.0, 12.0.

Pectoral fin: outer margin 15.7, 16. 2; inner margin 6.6, 6.6:, distal margin 11.5,

I3-4-

Distance from snout to: ist dorsal 35.1, 34.1; 2nd dorsal 61.5, 61.O; upper

caudal 76.4, 76.0; pectoral 21.9, 21.5; pelvics 50.3, 48.O; anal 61.3, 61.O.

Interspace between: ist and 2nd dorsals 16.7, 18.1; 2nd dorsal and caudal 6.9,

7.OJ anal and caudal 6.6, 6.4.

Distance from origin to origin of: pectoral and pelvics 27.0, 27.0 j pelvics and anal

13.4, 14.4.

Trunk moderately stout, tapering only slightly rearward, without mid-dorsal ridge.
Body sector from snout to cloaca about i V4 times as long as tail sector. Caudal peduncle

rrfp^

Figure 53. Negafrion brevirostris. A Head of specimen pictured in Fig. 52, from below, about % natural
size. B Left-hand nostril, about 4.5 x.

312 Memoir Sears Foundation for Marine Research

only slightly compressed laterally, about % as high as thick. Upper precaudal pit strongly
marked as a subtriangular depression, its concavity rearward, but no lower precaudal pit.
Dermal denticles comparatively large (average about 0.4 x 0.6 mm. in 2,490-mm. speci-
men), mostly overlapping, with 3 or 5 ridges, the median ridge and the pair next to it high,
sharp-topped and separated by deep furrows, the posterior margins with prominent teeth
opposite the 3 primary ridges, the median a little the longest, with or without small teeth
opposite the outermost pair of ridges on such of the denticles as have the latter; pedicels
moderately broad, as are the basal plates.

Head moderately flattened above, its length to 5th gill opening a little less than V4
of total length, its breadth opposite corners of mouth a little less than % its length. Snout
very broadly and evenly rounded, its length in front of a line connecting outer ends of
nostrils only about Yo as great as distance between inner ends of latter, its length in front
of mouth equal to distance between nostrils, or to about V4 the length of the head. Eye
oval, noticeably small, its horizontal diameter a little less than V3 as long as distance be-
tween nostrils, and only about V2 to % as long as ist gill opening. Gill openings all of
very nearly equal lengths, about i V2 to 2 times as long as diameter of eye, evenly spaced,
the 1st about perpendicular, but the 3rd to 5th increasingly oblique, the 4th above origin of
pectoral. Nostril moderately oblique, its inner end a little nearer to front of mouth than
to tip of snout, its anterior margin expanded as a triangular lobe about as long as broad, the
distance between inner ends of nostrils about V2 to % as great as breadth of mouth and
about % as great between their outer ends. Mouth broadly rounded and moderately
arched, its height slightly more than V2 as great as its breadth (51 to 60% in specimens
examined), occupying about % of breadth of head. Upper labial furrow extending out-
ward nearly at right angles to upper jaw as a deep groove for a distance about V3 as long
as horizontal diameter of eye; no furrow on lower jaw.

Teeth i3o'/j7^'3°i."^" 14' ^^'■^ ^^T sharp cutting edges; upper teeth with narrow trian-
gular cusps and broad bases, symmetrical and erect in central part of jaw but increas-
ingly oblique toward its corners, the outer margins increasingly notched with the outer-
most deeply so, the edges of cusps smooth but edges of basal sectors with moderately
fine, irregularly rounded serrations, except for the smooth small teeth at the sym-
physis and near the corners of the jaw; lower teeth similar in general to uppers, except
somewhat more slender and more erect, the bases, as well as cusps, smooth-edged, except
near the corners of the jaw where they are somewhat wavy; i to 3 minute, smooth-edged
teeth at symphysis in upper jaw and 3 in lower; outermost 3 or 4 teeth in each jaw very
small; i row functional, or 2 rows in places.

First dorsal low relative to its length, its anterior margin about 1.25 times as long as its
base, its vertical height slightly less than V2 as great as length of pectoral, its origin a little
posterior to corner of pectoral, its anterior margin nearly straight, apex narrowly rounded,
posterior margin nearly straight toward apex but moderately concave basally, its free rear
corner about ^o as long as the base. Second dorsal similar to ist dorsal in shape, nearly as

Fishes of the Western North Atlantic 313

high, and about % to % as long at base, its origin over or slightly anterior to origin of
anal, its free rear corner a little longer than V2 the base {SS%)' Caudal a little less than
V4 of total length, its axis raised at an angle of about 15 to 18°, its upper margin weakly
convex, tip subacute, the terminal sector a little less than Yi the length of fin, the lower
anterior corner expanded as a definite lobe about V2 as long as the upper, with moderately
convex anterior edge narrowing to a subacute tip. Anal slightly but evidently smaller than
2nd dorsal, its anterior margin a little more convex, its apex more broadly rounded and its
posterior margin much more deeply concave, its free rear corner a little more than % as
long as base (69 to 70%). Pel vies with weakly concave rear margins, narrowly rounded
distal corners and subacute tips, their anterior margins about as long as anterior margin of
anal. Pectoral a little less than % (about 7 1 % ) as long as head, about % as broad as long,
with noticeably long base (as long as, or a little longer than, inner margin), the outer
margin moderately convex, distal margin moderately concave and corners very narrowly
rounded.

Color. Usually yellowish brown above, but sometimes dark brown or dark bluish
gray; lower sides more or less tinged with yellow, or with greenish olive; lower surface
either white, pale yellowish or in some cases grayish olive, like the back; anal usually yel-
lowish, edged with gray; other fins grayish, either with or without dark edges; margins of
gill openings white, shading to dark gray; inside of mouth white, at least in some speci-
mens.

Size. Matures at about 7 to 7 V2 feet and grows to a maximum length of about 1 1 feet.
One of 9 feet 6 inches is said to have weighed only 265 pounds.

Developmental Stages. It is not known whether or not the embryo develops a yolk-
sac placenta.

Habits. Enough information has now accumulated to show that this, like Ginglymos-
toma, is strictly an inshore species, common around docks {^e.g.y at Key West, Florida), in
salt-water creeks {^e.g., around southern Florida) and in enclosed sounds as along the
coast of North Carolina. It has even been reported from within the mouth of the Amazon
River and from fresh water elsewhere in Brazil. But it is not known from Bermuda, nor is
there any positive record that it appears elsewhere more than a very short distance out
from land. Around southwestern Florida it evidently breeds in spring and summer, for
newborn specimens with umbilical scars still open (624 to 630 mm.) are taken in shallow
inlets from May to September. The only direct information available as to its diet is that
cowfish (^Lacto-phrys) were found in the stomach of one, and a sting-ray's spine was
imbedded in the jaw of another. But this, with the fact that it readily takes a hook baited
with fish, makes it likely that it feeds indiscriminately on whatever fish may be available
locally, as its teeth would suggest.

Relation to Man. Around southern Florida it has some value commercially, its hides,
fins and liver oil being of good quality. On the other hand, it has been suspected of attacks
on bathers in South Carolina waters, whether justly or not.

314 Memoir Sears Foundation for Marine Research

Range. Littoral, in the western Atlantic, northern Brazil to North Carolina, and acci-
dentally to New Jersey 5 also reported from tropical West Africa."

Occurrence in the Western Atlantic. This is one of the more plentiful of the larger
sharks along the Florida Keys and on the southern and southwestern coast of Florida,
where it constitutes a considerable portion of the catches of the shark fishery. And it is com-
mon along the west coast of Florida, at least as far north as Tampa and Pensacola. In all
probability its center of abundance covers the West Indian— Caribbean region as a whole,
and the southern part of the Gulf of Mexico, although definite records of it there are con-
fined to the Bahamas,' Cuba, Jamaica,' and the Atlantic coast of Panama.

To the northward its presence has been established recently off Mississippi in July,*
and it ranges in the summer not uncommonly as far as South Carolina and the southern
part of North Carolina. But it appears that few pass the latitude of Cape Hatteras, unless
perhaps they enter the warm enclosed waters of Pamlico Sound, for the only record of it
further north is of a single large specimen from Beach Haven, New Jersey, in July 1919.
Present indications are that its range is equally circumscribed in the opposite direction, the
only South American records for it being of a very small specimen from Para in northern
Brazil, and another from fresh water of some unspecified Brazilian locality.

SynonjTns and References:

Hyfofrion brevirostris Poey, Repert. Fisico-Nat. Cuba, 2, 1868: 451, pi. 4, fig. 5, 6, 20 (descr., teeth,
Cuba) ; An. Soc. esp. Hist, nat., 5, 1876: 394; Enumerat. Pise. Cubens., 1876: 198 (Cuba) ; Goode and
Bean, Proc. U.S. nat. Mus., 2, 1879: 156 (W. Florida); Jordan and Gilbert, Proc. U.S. nat. Mus.,
5, 1882: 581 (descr., color, S. Carolina); Bull. U.S. nat. Mus., 16, 1883: 61 (W. Indies, Gulf coast
of U.S.); Goode and Bean, Proc. U.S. nat. Mus., 5, 1883: 240 (Gulf of Mexico); Jordan and
Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 217 (W. Indies to S. Carolina); Bull. U.S.
nat. Mus., 47 (i), 1896: 41 (descr., W. Indies to S. Carolina); Bull. U.S. nat. Mus., 47 (4),
1900: pi. 5, fig. 18 (ill.); Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 48
(Florida); Bean, B. A., in Shattuck, Bahama Islands, Fish., 1 905: 296 (Bahamas); Rosen, Lunds Univ,
Arsberatt., (7) 2 (5), 191 1: 47 (Watlings I.); Garman, Mem. Mus. comp. Zool. Harv., 37, 1913: 120
(descr., Pensacola, Florida) ; Gudger, Science, N. S. 57, 1 91 3 : 993 (Key West, Florida) ; Starks, Stanford
Univ. Publ., Univ. Ser., 1913: 5 (Para, Brazil); Gudger, Yearb. Carneg. Instn. (1913), 12, 1914: 177
(Tortugas, Florida); Yearb. Carneg. Instn. (1914), 13, 1915: 204 (Tortugas, Florida); Science, N. S.
41, 1915: 437 (anat.. Key West, Florida); Coles, Proc. biol. Soc. Wash., 28, 1915: 90 (N. Carolina);
Radcliffe, Bull. U.S. Bur. Fish., 34, 1916: 253, pi. 39, fig. I (meas., descr., N. Carolina) ; Nichols, Bull.
Amer. Mus. nat. Hist., 57, 1917: 875 (Florida); Fowler, Proc. Acad. nat. Sci. Philad., 72, 1921: 386
(size, weight, Beach Haven, N. Jersey); Meek and Hildebrand, Field Mus. Publ. Zool., 75, 1923: 50
(descr., Atlant.) ; Borodin, Bull. Vanderbilt Oceanogr. (Mar.) Mus., i (l), 1928: 5 (Florida); Breder,
Field Bk. Mar. Fish. Atlant. Coast, 1929: 18 (general); Gudger, Publ. Carneg. Instn., 391, 1929: 200
(food, Tortugas, Florida) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 17
(distrib.) ; Gudger, Sci. Mon. N. Y., 34, 1932: 409 (sting-ray spine in jaw, west coast, Florida) ; Breder,
Zoologica, N. Y., 18, 1934: 59 (W. Indies, Bahamas); Brooks, Parasitology, 26, 1934: 260 (Tortugas,

5. Budker, Bull. Mus. Hist. nat. Paris, (2) 7, 1935: 185. The collection of the Harvard Museum of Comparative
Zoology also contains a female of 872 mm. received in 1864 and catalogued as from the Kingsmill Islands.
But we hesitate to include the West Pacific in the range of the species, in view of the possibility that the speci-
men may not have come from the stated locality.

6. There is a specimen from Watlings Island in the United States National Museum (No. 38497).

7. Personal communication from Luis Howell-Rivero. 8. Personal communication from Stewart Springer.

Fishes of the Western North Atlantic 315

Florida, parasites) ; Budker, Bull. Mus. Hist. nat. Paris, (2) 7, 1935: 185 (Dakar, W. Afr.) ; Burton, Sci.
Mon. N. Y., 40, 1935: 279 (perhaps attacks on bathers, S. Carolina); White, Bull. Amer. Mus. nat.
Hist., 7^, 1937: I 28 (in Key) ; Springer, Proc. Fla. Acad. Sci., 5, 1939: 28 (breeding season, Florida) ;
Hildebrand, Copeia, 1941:221 (N. Carolina) ; Longley and Hildebrand, Pap. Tortugas Lab., 5.;, 1941:
2 (Tortugas, Florida); Bomkamp, Contr. biol. Lab. Cath. Univ. Amer., 44., 1942: 4 (chemistry of
liver, Salerno, Florida); Fowler, Arqu. Zool. Estad. Sao Paulo, 3, 1942: 128 (listed for Brazil); Fish
Culturist, 21 (9), 1942: 66 (listed Cuba, but not p. 67, fig. 56); Boos, Contr. biol. Lab. Cath. Univ.
Amer., 45, 1943: 10 (chemistry of pancreas, Salerno, Florida), Lunz, Bull. S. C. St. Planning Bd., 14,
1944: 27 (S. Carolina, Florida); Gunter, Publ. Inst. Mar. Sci. Univ. Texas, 1945: 20 (temp., breeding,
Texas) ; Bigelow and Schroeder, Guide Coram. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945:
116, fig. 41 (descr., habits, range, ill.); Fowler, Monogr. Acad. nat. Sci., 7, 1945: 96 (Beaufort, N.
Carolina), 160 (S. Carolina), 263 (Bahamas).

Carcharias {Hypofrion) brevirostris Gunther, Cat. Fish. Brit. Mus., 8, 1870: 362 (descr., Cuba).

Carcharias brevirostris Jordan and Gilbert, Proc. U.S. nat. Mus., 5, 1883: 245 (Gulf of Mexico); Jordan,
Bull. U.S. Fish Comm., 4, 1884: 79 (Key West, Florida); Proc. U.S. nat. Mus., 7, 1887: 104 (Key
West, Florida) ; Englehardt, Zool. Anz., 59, 1912: 648 (Brazil, in fresh water).

Carcharhinui {Hy-pofrion) brevirostris Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 796 (W. Indies).

Carcharhinus brevirostris Henshall, Bull. U.S. Fish. Comm., 9, 1891: 383 (Florida, teeth).

Genus Hyfop'ion Miiller and Henle, 1 841

Hyfofrion Miiller and Henle, Plagiost., 1841 : 34; type species, Carcharias {Hyfoprion) mactoti Muller and
Henle, 1841, designated by Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 61.

Generic Synonyms:

Squalus (in part) Gray, List. Fish. Brit. Mus., 1851: 43; not Squalus Linnaeus, 1758.

HyfofrionoJon Gill, Ann. N. Y. Lye, 7, 1862: 399, 401, 409; type species, Carcharias {Hyfofrion)

hemiodon Muller and Henle, 1 841.
Carcharias (in part) Gunther, Cat. Fish. Brit. Mus., 8, 1 870: 362 ; not Carcharias Rafinesque, 1 810.

Generic Characters. Carcharhinidae with anal less than i ^o times as long at base as
2nd dorsal; without spiracles; midpoint of ist dorsal at least as near to axil of pectoral as
to origin of pelvics; 2nd dorsal only about V2 as long at base as ist dorsal; caudal peduncle
without lateral ridges, but with a precaudal pit below as well as above; back with a low
mid-dorsal ridge, at least in some species; a labial furrow at corner of mouth and extend-
ing for a short distance on upper jaw, but not onto lower; upper teeth strongly oblique and
notched outwardly, or erect, their cusps smooth-edged, but their bases with several coarse
marginal serrations or low denticles on the outer side, more or less wavy or indistinctly
serrate on the inner; the lowers slender, erect, both bases and cusps smooth-edged. Char-
acters otherwise those of the family.

Range. Western Atlantic; Chile; China and Indo-China; East Indies; Philippines;
New Guinea; India and tropical Indian Ocean, including Red Sea and Gulf of Arabia.^

Species. One species is known in the warm belt of the western Atlantic, and two or per-
haps three in the Pacific and Indian Oceans.^

1. Also reported for Australia, but incorrectly, according to Whitley (Fish. Aust., /, 1940: 107).

2. An additional species, H. brevirostris Poey, 1868, has been referred to this genus previously, but it falls in
Negafrion, according to the generic definitions adopted here.

3i6 Memoir Sears Foundation for Marine Research

Key to Species

I a. Snout in front of mouth considerably longer than breadth of mouth.

2a. Tip of 1st dorsal terminates anterior to origin of pelvics by a distance at least as

long as base of latter j origin of 2nd dorsal opposite or only a very little posterior

to origin of anal. signatus Poey, 1868, p. 316.

2b. Tip of 1st dorsal reaches nearly to a perpendicular at origin of pelvics; origin of

2nd dorsal over or posterior to midbase of anal.

macloti Muller and Henle,' 1841.
New South Wales, East Indies, India; also
probably Chile.*

lb. Snout in front of mouth not longer than breadth of mouth.

3a. Base of 2nd dorsal only % as long as base of anal; posterior margin of 2nd dorsal
not concave, but that of anal deeply so j upper teeth oblique, lowers erect.

hemiodon Muller and Henle, 1 841.

Arabian Gulf, India, Indo-China, East In-
dies, Philippines.'

3b. Base of 2nd dorsal as long as base of analj 2nd dorsal of same shape as anal;
upper teeth erect, like lowers. ■playfairii Gunther, 1 870.

Madagascar.

Hypofrion signatus Poey, 1868

Night Shark

Figure 54

Study Material. Immature female, 935 mm. long, from offing of South Carolina,
Lat. 33° 37' 30" N., Long. 77° 36'3o" W., in 14 fathoms on October 20, 1885 (U.S.Nat.
Mus., No. 38508) ; three embryos (two females, one male), 385 to 407 mm. long (Harv.
Mus. Comp. ZooL, No. 36091); also photograph of adult female taken oflf north coast
of Cuba.*

Distinctive Characters. Easily separable from all other local carcharhinids by the
combination of very long, pointed snout with smooth-cusped teeth, but the uppers strongly
serrate at the base, and with the presence of a low but unmistakable mid-dorsal ridge.

Description. Proportional dimensions in per cent of total length. Female, 935 mm.,
from Lat. 33° 37' 30" N., Long. 77° 36' 30" W. (U.S. Nat. Mus., No. 38508).

3. Proportional dimensions calculated from measurements given by Muller and Henle (Plagiost., 1841 : 34) for the
type specimen.

4. Reported as Hyfofrion ? {Hemigaleus i ) /leterodus and Hyfoprion J (Hemigaleus ? ) isodus by Philippi (An.
Univ. Chile, yt, 1887: 541, 542).

5. According to Whitley (Fish. Aust., i, 1940: 107), a report of this species from South Australia by Zietz probably
was not correct.

6. Contributed by Luis Howell-Rivero.

Fishes of the Western North Atlantic

317

Trunk at origin of pectoral: hrea.dih 11.6; height 11.6.
Snout length in front of: outer nostrils 6.0 ; mouth lO.O.
Eye: horizontal diameter 2.2.
Mouth: breadth 7.4; height 5.2.
Nostrils: distance between inner ends 5.2.

Gill opening lengths: ist 2.2} 2nd 2.35 3rd 2.55 4th 2.3; 5th 1.8.
First dorsal fin: vertical height 7.83 length of base 9.7.
Second dorsal fin: vertical height 2.0; length of base 3.5.
Anal fin: vertical height 2.7; length of base 4.0.
Caudal fin: upper margin 26.9 ; lower anterior margin 1 1.4.
Pectoral fin: outer margin 16.8 j inner margin 5.4; distal margin 13.2.
Distance from snout to: ist dorsal 34.7; 2nd dorsal 64.3; upper caudal 73.1 j pec-
toral 25.2} pelvics 51.5; anal 63.0.

Interspace between: ist and 2nd dorsals 23.45 2nd dorsal and caudal 5.8 j anal and
caudal 6.3.
Distance from origin to origin of: pectoral and pelvics 29.2; pelvics and anal 12.1.

Figure 54.. Hyfofrion signatus, female, 935 mm. long, from off South Carolina (U.S. Nat. Mus., No.
38508). A Anterior part of head from below, about 0.3 x natural size. B Left-hand nostril, about 2.6 x.
C Left-hand upper and lower teeth, about 1.5 x. D Sixth upper tooth. E Twelfth upper tooth. F Fourth
lower tooth. G Eleventh lower tooth. D—G, about 3.0 x. H Dermal denticles, about 30 x. / Dermal denticle,
apical view, about 60 x.

3 1 8 Memoir Sears Foundation for Marine Research

Trunk comparatively stout, about Vs as high at ist dorsal (where highest) as length
to origin of caudal. Caudal peduncle moderately slender, the lower as well as upper pre-
caudal pit strongly marked, subrectangular. Dermal ridge low but unmistakable along
midline of back from close behind rear end of base of ist dorsal to origin of 2nd dorsal/
Dorsal profile sloping sharply, in convex contour, from ist dorsal forward. Dermal den-
ticles rather loosely spaced and overlapping but little, their blades nearly horizontal, about
as broad as long, usually with 3 low ridges, the posterior margins usually with 3 teeth, but
occasionally 5, the median considerably the largest; pedicel rather slender.

Head noticeably long, forming about V3 of trunk to origin of caudal. Snout narrow,
ovate at tip and very long, its length in front of mouth a little more than Vs of length of
head, or about 1.25 times as great as breadth of mouth, and its length anterior to outer ends
of nostrils a little more than V2 as great as length in front of mouth. Eye nearly circular, its
diameter about V2 as great as distance between nostrils. Gill openings noticeably small,
the 1st to 3rd (slightly the longest) about as long as diameter of eye, the spaces between
1st and 2nd and between 2nd and 3rd of about equal breadth, but those between 3rd and
4th and between 4th and 5th a little narrower, the 4th gill opening over origin of pectoral.
Nostril strongly oblique, its inner corner nearer to mouth than to tip of snout by a dis-
tance about V2 as great as that between them, the anterior margin with a pronounced narrow
triangular lobe near the inner end; the distance between nostrils a little greater than %
breadth of mouth and about ^^ length of snout. Mouth broadly ovate, about i Vs times as
broad as high. Upper labial fold about % as long as nostril.

Teeth '^~^°''_^'^ ; those at symphysis small, triangular, the base with or without
a blunt denticle on each side; uppers otherwise acute-triangular, increasingly oblique
toward corners of mouth, the inner margins nearly straight and smooth-edged on
cusps but more or less wavy or irregularly serrate basally, the outer margins strongly
notched, smooth toward tips, but with 2 to 4 very prominent serrations or low denticles on
basal sector, the distal serration considerably the largest; lowers symmetrical, more slender
than uppers, nearly erect, bases as well as cusps with smooth edges.

First dorsal comparatively small, its anterior margin only about as long as snout in
front of mouth, and about % as long as pectoral, its origin about over inner corner of pec-
toral,* its anterior margin weakly convex, its posterior margin strongly concave basally, its
apex rounded, its free corner a little less than Yo as long as its base. Second dorsal a little
less than V2 as long as ist dorsal at base, relatively much lower, and only about V5 to %
as great in area, its apex broadly rounded, posterior margin only very weakly concave, its
free rear tip very slender and nearly as long as its base, its origin very little posterior to
origin of anal. Caudal about V4 of total length, with bluntly rounded tip, its terminal
sector about V4 the length of fin, the lower anterior lobe about 40% as long as upper mar-
gin, with rather broadly rounded tip. Interspace between caudal and anal about i V2 times
as long as base of anal. Anal about as high and long as 2nd dorsal and with similarly slender

7. In the preserved state this ridge lies at the bottom of a groove of muscular contraction.
S. Its precise point of origin is difficult to determine in the preserved specimen.

Fishes of the Western North Atlantic 319

free rear tip, but with much more deeply concave posterior margin. Pelvics with nearly
straight edges and narrowly rounded corners, about as long as anal along anterior margins,
about I V2 times as large in area, their origin considerably nearer to rear end of base of ist
dorsal than to origin of anal. Distance from cloaca to caudal about % as great as from cloaca
to inner corner of pectoral in female, but perhaps somewhat longer, relatively, in male
(not seen). Pectoral about % (about 66%) as long as head, and about V2 as broad as long,
with strongly convex outer margin, weakly and evenly concave distal margin, and rather
narrowly rounded corners.

Color. Adults in life are bluish gray above, grayish white below, with small black
spots scattered over the body ; embryos are silvery gray above, dirty white below.'

Size. The lengths of recorded specimens, for which the identity is established, are 955
mm. (immature, see above), 2,766 and 2,270 mm. (male and female); thus this Shark
grows to at least a moderately large size.

Developmental Stages. The embryos bear long yolk stalks and well developed yolk-
sac placentae, spongy in texture, showing that development is viviparous. Females have
been taken with as many as 1 2 embryos.^"

Habits. The only available information as to its habits is that Cuban fishermen report
it as caught well oflFshore only, on set lines at depths greater than 1 50 fathoms, and only at
night (hence its local name "Tiburon de Noche'').

Range. The Night Shark is known only off the north coast of Cuba, where it is so
common in the Gulf Stream that 36 specimens have been counted at one time at the shark
fishery station at Cojimar," and from the offing of South Carolina (a single specimen,
see Study Material, p. 3 1 6) . It has been recorded also by name from Georgetown, British
Guiana, as well as from Key West and the Tortugas, Florida, but without any supporting
evidence as to the actual identity of the specimens concerned. The fact that only one speci-
men has been reported from the east coast of the United States, although signatus is easily
recognizable, suggests that it wanders northward from its tropical home only as a stray.

Synonyms and References:

H-jfofrion signatusYovf, Repert. Fisico.-Nat. Cuba, 1868: 452, pi. 4, fig. 7, 8 (descr., teeth, Cuba); An. Soc.
esp. Hist, nat., 5, 1876: 395 ; Enumerat. Pise. Cubens., 1875: 199 (Cuba) ; Jordan and Evermann, Bull.
U.S. nat. Mus., 47 (l), 1896: 41 (descr., Cuba); Blosser, Ann. Carneg. Mus., 6, 1909: 295 (Brit.
Guiana, ident. queried) ; Garman, Mem. Harv. Mus. comp. Zool., 56, 1913: 122 (descr.) ; Gudger, Sci-
ence, N. S. ^/, 1915: 437 (name only, Tortugas, Florida) ; Yearb. Carneg. Instn. (1914), 15, 1915= 203
(name only. Key West, Florida) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930:
17 (Cuba) ; White, Bull. Amer. Mus. nat. Hist., T4, 1937: 128 (in Key) ; Bigelow and Schroeder, Guide
Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1 945: 130, fig. 47 (descr., range, ill.).

Carcharias (^Hyfofrion) signatus Gunther, Cat. Fish. Brit. Mus., 1870: 362.

Hypofrionlongirostris Poey, An. Soc. esp. Hist, nat., 5, 1876: 198, pi. 9, fig. 8, 9.^'

9. Information contributed by Luis Howell-Rivero. 10. Information contributed by Luis Howell-Rivero.

11. Personal communication from Luis Howell-Rivero.

12. We agree with Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 123) that longirostris, based on a 2,266 mm.
specimen, was a synonym of signatus (based on jaws only), there being no essential diflferences in the teeth; a
photograph of unpublished drawings by Poey, showing the anterior part of the head of "longirostris," agrees
closely with the specimen illustrated here (Fig. 54) in shape and length of snout relative to breadth of mouth.

320 Memoir Sears Foundation for Marine Research

Genus Carcharhinus^ Blainville, 1 8 1 6

Carcharhinus Blainville, Bull. Soc. philom. Paris, 1816: 121 ; J. Phys. Chem. Hist, nat., S^, 1816: 264; in
Vieillot, Faune Franc, 1825: 88; type species, C. commersonii Blainville, 1816, 1825.^*

Generic Synonyms:

Carcharinu! Cloquet, Diet. Sci. Nat., 7, 18 17: 77; and subsequent authors; emended spelling for Carcharhinus

Blainville, 1 81 6.
Carcharias (in part) Cuvier, Regne Anim., 2, 1817: 125; and subsequent authors; not Carcharias Rafinesque,

1810.
Frionodon (in part) Muller and Henle, Plagiost., 184.1: 35 (preoccupied by Horsfield, 1823, for mammals).
Galeolamna Owen, Cat. Osteol. Roy. Coll. Surg. London, /, 1853: 96, no. 427; type, G. greyi Owen.

South Austialia.^
Eulamia Gill, Ann. N. Y. Lye, 7, 1862: 401 (name), 410 (diagn.) ; type species, Carcharias {Prionodon)

milberti Muller and Henle, 1841.'

1. Often spelled Carcharinus.

la. The generic name Carcharhinus has long been a "football" in Elasmobranch nomenclature. Briefly, its history is as
follows: Blainville, in his original diagnosis of the genus, listed the following species by name only: commersonii,
lamia, li'vidus, ustus, heterodon, verus, broussonetii, glaucus, caeruleus, megalofs, heterobranchialis, cormibicus,
monensis and vulfes. This assemblage includes representatives of Lamna Cuvier, 1817, as now understood
(cornubicus, monensis) ; of Alofias Rafinesque, 1810 {yulfes) ; of Prionace Cantor, 1849 {glaucus, caeruleus) \
and of Carcharodon L. Agassiz, 1838 (^carc/mrias) ; also one (lamia) that by subsequent evidence apparently
represented the combination of some unidentifiable member of the genus here named Carcharhinus, with the teeth
of Galeocerdo. The remaining members of the list were nominal only, and have so remained except for commer-
sonii which was later stated by Blainville (in Vieillot, Faune Franc, 1825 : 90) to have been based by him on the
shark pictured by Laccpede, "T. t, pag. 169, pi. 5, fig. i." Unfortunately, this reference was erroneous, for pi. 5,
fig. 1 pictures a skate and not a shark. However, if it was pi. 8, fig. i, that was intended (as seems almost certain),
commersonii seems to have been a member of the genus now under discussion, though neither the illustration in
question nor the accompanying measurements of a "requin" sufiice for specific identification.

The specific name commersonii seems then to have lain in abeyance until 1913, when Garman (Mem. Harv.
Mus. comp. ZooL, 36, 1913 : 140) revived it. However, his account appears to have been based on a combination
of longimanus Poey, 1861, with leucas Muller and Henle, 1841, and various authors following Garman have
subsequently used the name commersonii for one or the other of these two species, although there is no apparent
reason for identifying leucas with Lacepede's illustration, while longimanus certainly cannot be so identified.

Bosc (Nouv. Diet. Hist. Nat., 5, 1816: 277) was the first to designate a type species for Carcharhinus in the
words "Squalus carcharias lui sert de type." But this designation is not valid, because the original assemblage of
species listed in the genus by Blainville (Bull. Soc. philom. Paris, 1816: 121 and J. Phys. Chem. Hist, nat., 1816:
264) did not include a "carcharias," although he did describe a Carc/iarhinus under that specific name subsequently
(in Vieillot, Faune Franc, 1825: 89). The next designation of a type was commersonii Blainville, 1816, by
Jordan and Gilbert (Bull. U.S. nat. Mus., 16, 1883 : 22) ; and this selection appears to be valid, for while Fowler
(Proc Acad. nat. Sci. Philad., 60, 1908 : 62) has preferred vulfes as the type on the ground that commersonii was
a nomen nudem, thus reducing Carcharhinus to a synonym of Alofias Rafinesque, 1810, commersonii cannot prop-
erly be discarded, for, as pointed out above, the illustration on which it was based is identifiable with reasonable
certainty at least as to genus if not to species.

2. According to Whitley (Aust. Zool., 9, 1939: 230), the type, which he saw in London, is the jaws of the com-
mon "Whaler Shark" of Australia (Carcharias brachyurus of Giinther, 1870, and C macrurus of Ramsay and
Ogilby, 1887) which seems clearly referable to Carcharhinus Blainville 18 16, as here recognized.

3. In his first mention of the genus, Gill (Ann. N. Y. Lye, 7, 1S62 : 401) included only one species, "Eulamia lamia
Gill," which he stated to be the type, and Jordan and Gilbert (Bull. U.S. nat. Mus., 16, 1883 : 60), and later Jordan
(Genera Fish., 3, 1919: 306), accepted this designation, changing the authorship of lamia, however, from "Gill"
to "Risso" in the one case and to "Rafinesque" in the other. But this alteration was not warranted, for "lamia Gill"
was a nomen nudem, no account having ever been published by Gill himself of any shark under that name, or
even any indication as to whether he referred to the "latnia" of Rafinesque, 1810 (a name substituted for Squalus
carcliarias Linnaeus, 1758, and therefore equivalent to Carcharodon), of Risso, 1826, or of Mijller and Henle,
1 84 1. Therefore, it is fortunate that the only species included by Gill under his diagnosis of Eulamia on a later

Fishes of the Western North Atlantic 321

Platyfodon Gi!I, Ann. N. Y. Lye, 7, 1862: 4.01 ; type, Carcharias {Prionodon) menisorrah Miiller and Henle,

1841.
hogomfhodon Gill, Ann. N. Y. Lye, 7, 1862: 401 ; type, Carcharias (Prionodon) oxyrhynchus Muller and

Henle, 1 841.
Lamiofsit Gill, Ann. N. Y. Lye, 7, 1862: 401 ; type, Carcharias (Prionodon) temmincki Muller and Henle,

1841.
Isoflagiodon Gill, Ann. N. Y. Lye, 7, 1862: 401; type, Carcharias {Prionodon) sorra Muller and Henle,

1841.
Gymnorhinus Hilgendorf, in Hemprich and Ehrenberg, Symbol. Phys. Icon. Ined. Pisces, 1899: 8; type,

Carcharias (Prionodon) menisorrah Muller and Henle, 1 84 1 (not seen); preoccupied by Maximillian,

1 841, for birds.
MafoUtmia Whitley, Mem. Qd. Mus., 10 (4), 1934: 185, 188; type, Carcharias melano-pterus Quoy and

Gaimard, 1 824.
Gillisqualus Whitley, Mem. Qd. Mus., 10 (4), 1934: 185; type, Carcharias (Prionodon) amilyrhynchus

Bleeker, 1856.
Gdeolamnoides Whitley, Mem. Qd. Mus., 10 (4), 1934: 185, 191 ; type, Carcharias macrurus Ramsay

andOgilby, 1887.
Gaholamna Whitley, Aust. Zool., p, J939: 23O; revives GaUolamna Owen, 1853, vyhich see above.
OgUamia Whitley, Aust. Zool., 9, 1939: 231, subgenus; type, Carcharias sUvensi Ogilby, 191 1. Australia.
Uranga Whitley, Proc. Linn. Soe N. S. W., 68, 1943: 115; type, U. nasuta Whitley. Australia.
Uranganop Whitley, Proc. Linn. Soe N. S. W., 68, 1943: 117, subgenus; type, GaUolamna (Uranganofs)

fitzroyensis Whitley. Australia.
Lamnarius Whitley, Proc. Linn. Soe. N. S. W., 68, 1943: 119, subgenus; type, Carcharias sfenceri Ogilby,

1 910. Australia.
Bogimia Whitley, Proc. Linn. Soe N. S. W., 68, 1943: 123, subgenus; type, GaUolamna (Bogimia)

bogimba Whitley. Australia.
Longmania (in part) Whitley, Aust. Zool., 10 (3), 1944: 257; for L. calamaria Whitley, Australia; not

Longtnania Whitley, 1939, which is a synonym of Afrionodon (see p. 303).

Doubtful synonym:

Glyfhis L. Agassiz, Polss. Foss., 3, 1838: pi. 36, fig. 10-13; Poiss. Foss., 3, 1843: 243; type species, G.
has talis h. Agassiz, 1838.*

Generic Characters. Carcharhinidae with anal fin little if any longer at base than 2nd
dorsal J without spiracles; midpoint of ist dorsal nearer to axil of pectoral than to origin
of pelvics; cusps of upper teeth regularly serrate, those of lowers serrate or smooth 5 back
with or without mid-dorsal ridge; caudal peduncle without lateral ridges, but with well
developed upper precaudal pit, the lower varying from well developed to hardly discern-
ible; 1st dorsal subtriangular, its posterior margin more or less deeply concave, its lower
posterior angle more or less prolonged as a free corner; 2nd dorsal and anal of approxi-
mately equal size and much smaller than ist dorsal; pelvics quadrilateral, their inner

page of the same publication (Ann. N. Y. Lye, 7, 1862: 410) was the well knovni Carcharias (Prionodon)
milberti Muller and Henle, 184.1. This must necessarily stand as the type of the genus. See also footnote 82,
P- 368.
4.. The fossil shark's teeth to which L. Agassiz gave this name resemble the anterior lower teeth of Carcharias (Priono-
don) glyf/iis Muller and Henle, 1841, which falls in Carcharhinus as here defined; they are slender, erect, cylin-
drical near the base, and with cutting edge confined to the lanceolate, laterally-expanded tips. But we doubt the
propriety of reviving the name Glyfhis for any modern shark on the evidence of these two teeth alone. See also
footnote 5, p. 280.

322 Memoir Sears Foundation for Marine Research

corners not elongate; dermal denticles either overlapping or not, their blades with 3 or
more ridges, their apical margins toothed or not; axis of caudal raised only moderately;
upper labial furrow very short, at an obtuse angle with the jaw; no lower labial furrow.
Development viviparous, with well developed yolk-sac placenta in the few cases where it is
known (see pp. 359, 394). Characters otherwise those of the family.

Range. Tropical and warm-temperate belts of all oceans, including the Mediter-
ranean, both inshore and on the high seas; also landlocked in Lake Nicaragua in fresh
water.

Fossil Teeth, closely resembling those of Carcharhinus (perhaps including Hypofrion
and AprionoJon), have been described under various names from: Eocene, Africa ; Eocene
to Pliocene, Europe and North America; Oligocene to Miocene, South America; and Mio-
cene, West Indies.

Species. Carcharhinus mdudes a much larger number of species than any other genus
of modern sharks and many of the most familiar of the larger sharks of warm seas. Its
members cover a wide range as regards teeth, the relative sizes and shapes of fins, and to a
lesser degree the relative positions of the latter. But the extremes are connected by such a
continuous series of intermediate stages in all these respects that attempts to subdivide the
genus have not been easy.^ In fact, the only alternative character which might form a sharp-
cut basis for such subdivision, from the standpoint of specific identification, is the presence
or absence of a mid-dorsal ridge. But the use of this would entail the generic separation of
species that closely resemble one another in other respects, and the union of others that do
not, which seems too high a price to pay for reducing the length of the generic Key, which
would be the only advantage gained.

The genus has received much less attention than it deserves, no doubt due to the fact
that most of the species are rather large, with consequent paucity of specimens in collec-
tions. Many of the species resemble one another so closely in general appearance that little
or no dependence can be placed on published reports of occurrence unless accompanied by
some indication as to fins, teeth, etc. Therefore, we are very fortunate in having been able
to study specimens of all 1 3 species now known to occur in the western Atlantic ; finding
that while some of them look much alike on cursory examination, they are separable by
characters so precise and so little variable that specific identification is not difficult, although
attention to detail is required.

The genus is as universally distributed in the warmer belt of the Pacific and Indian
Oceans as it is in the Atlantic, and some of its Indo-Pacific representatives are evidently
very close to some of the Atlantic species, if not identical with them. However, to attempt
to revise the genus as a whole would be idle without access to adequate material of at least
a majority of the supposed Indo-Pacific species, which we have not had. The following
Key is therefore limited to the western Atlantic.

5. Whitley, in a series of papers, has recently broken the genus down into no less than nine genera and subgenera (see
Generic Synonyms, p. 321). But the characters on which these are based seem to us more appropriate for the
definition of species within the genus.

Fishes of the Western North Atlantic 323

Key to Western Atlantic Species

I a. Length of snout, anterior to a line connecting outer corners of nostrils, about 1.5
times as great as distance between nostrils.

oxyrhynchus Miiller and Henle, 1841, p. 391.
lb. Length of snout anterior to a line connecting outer corners of nostrils less than
distance between nostrils.
2a. Origin of 2nd dorsal over or behind midpoint of base of anal,

■porosus Ranzani, 1839, p. 394.
2b. Origin of 2nd dorsal over origin of anal, or anterior to it.

3a. Midline of baclc between ist and 2nd dorsal fins with a low but distinct
dermal ridge.

4a. Free rear corner of 2nd dorsal more than twice as long as vertical
height of the fin and notably slender (Figs, s^ B, 59 B).
5a. Anterior margin of pectoral nearly as long (about 92%) as
from tip of snout to origin of pectorals, and longer than from
snout to I St gill opening.

fioridanus Bigelow and Schroeder, 1943, p. 333.

5b. Anterior margin of pectoral not more than % as long as from

tip of snout to origin of pectorals, and shorter than from snout

to I St gill opening by a distance as great as that between nostrils.

falciformis Miiller and Henle, 1841, p. 329.

4b. Free rear corner of 2nd dorsal considerably less than twice as long

as vertical height of fin, and not notably slender.

6a. Apex of ist dorsal very broadly rounded} tip of anal reaches

nearly to origin of caudal. longimanus Poey, 1 8 6 1 , p. 3 54.

6b. Apex of ist dorsal subangular, or very narrowly rounded j tip

of anal separated from origin of caudal by a distance at least as

long as diameter of eye.

7a. Origin of ist dorsal about over inner corner of pectoral, its
vertical height less than distance from eye to ist gill open-
ing} dermal denticles regularly overlapping, with strongly
marked marginal teeth.

8a. Distance from tips of pelvics to origin of anal longer
than base of anal} 5th gill opening at least 1.5 times as
long as horizontal diameter of eye } vertical height of
1st dorsal only a little more than length of snout in
front of mouth } anterior margin of nostril not lobed.
obscurus Lesueur, 18 18, p. 382.
8b. Distance from tips of pelvics to origin of anal less than
% as long as base of anal} 5th gill opening only about
as long as horizontal diameter of eye} vertical height

324 Memoir Sears Foundation for Marine Research

of 1st dorsal about i Vs times as long as snout in front
of mouth J anterior margin of nostril with low, sub-
triangular lobe.

springeri Bigelow and Schroeder, 1 944, p. 404.

7b. Origin of ist dorsal over axil of pectoral, its vertical height

(after birth) at least as great as distance from eye to 3rd

gill opening} dermal denticles loosely spaced, without well

marked teeth, milberti Miiller and Henle, 1 841, p. 368.

3b. Midline of back between dorsals smooth, without dermal ridge.

9a. Apex of 1st dorsal very broadly rounded j tip of anal reaches nearly

to origin of caudal. longimanus Poey, 1 8 6 1 , p. 3 54.°'

9b. Apex of 1st dorsal subangular or narrowly rounded; tip of anal

separated from origin of caudal by a distance at least as long as

diameter of eye.

loa. Snout, in front of line connecting outer ends of nostrils, less
than ^ as long as distance between inner ends of nostrils.
I la. Anterior margin of eye opposite or a little anterior to
front of mouth; ist gill opening not more than ^2 as
long as distance between nostrils.

leucas Miiller and Henle, 1841, p. 337.
lib. Anterior margin of eye a little posterior to front of
mouth; ist gill opening nearly % as long as distance
between nostrils.

nicaraguensis Gill and Bransford, 1877, p. 378.
lob. Snout, in front of a line connecting outer ends of nostrils, at
least % as long as distance between inner ends of nostrils.
1 2a. Upper teeth strongly asymmetrical, their outer margins
deeply concave in subangular contour (notched).

acronotus Poey, i860, p. 325.
12b. Upper teeth along inner half of jaw erect, nearly sym-
metrical, their outer margins not much more concave
than the inner, if at all.

13a. First gill opening not more than 1.5 times as long
as horizontal diameter of eye; lower precaudal pit
only faintly indicated; fins without conspicuous
black markings; anterior margin of nostril con-
spicuously lobed (Fig. 76 F).

remotus Dumeril, 1865, P- 400.

5a. Longimanus is included under alternative 3b, as well as under 3a, because of uncertainty as to whether or not the
mid-dorsal ridge, evident in embryos, persists after birth.

Fishes of the Western North Atlantic 325

1 3b. First gill opening nearly twice as long as horizontal
diameter of eye, or longer} lower precaudal pit
strongly marked; fins conspicuously tipped with
black; anterior margin of nostril only slightly ex-
panded (Figs. 63 G, 67 B).
14a. Origin of ist dorsal about over midpoint of
inner margin of pectoral; ist gill opening less
than 2.5 times as long as horizontal diameter
of eye ; horizontal diameter about V5 as long
as snout in front of mouth; edges of lower
teeth regularly though very finely serrate.
Umbatus Muller and Henle, 1841, p. 346.
14b. Origin of ist dorsal over or a little posterior
to inner corner of pectoral; ist gill opening
more than 4 times as long as horizontal diam-
eter of eye; horizontal diameter of eye only
about Vt' to Vs as long as snout in front of
mouth; margins of lower teeth smooth.

maculipinnis Poey, 1866, p. 364.

Carcharhinus acronotus (Poey), 1861

Black-nosed Shark

Figure 55

Study Material. Two males, about 485 mm. long, either embryo or newborn with
umbilical scar still showing, from Rio de Janeiro; 2 male embryos, 358 and 371 mm.,
from Cuba; male embryo, 480 mm., about ready for birth, from Brazil (Harv. Mus.
Comp. Zool., No. 723); immature female, 637 mm., from Englewood, Florida (U.S.
Nat. Mus., No. 10433 ^ ) > also photographs of Poey's unpublished drawings of this species.
Distinctive Characters. Easily recognized, among those of the smooth-backed mem-
bers of the genus in which the second dorsal originates above the origin of the anal, by the
relatively long snout, combined with the fact that the upper teeth are noticeably asym-
metrical with deeply notched outer edges and much more coarsely serrate bases than tips.
Description. Proportional dimensions in per cent of total length. Male, 485 mm.,
from Rio de Janeiro (Harv. Mus. Comp. Zool., No. 701). Female, 637 mm., from Engle-
wood, Florida (U.S. Nat. Mus., No. 10433 1).

Trunk at origin of pectoral: breadth 10.7, li.O; height 1 1.5, i i.O.

Snout length in front of: outer nostrils 3.7, 3.6; mouth 8.0, 8.0.

Eye: horizontal diameter 2.5, 1.9.

Mouth: breadth 7.2, 7.2; height 4.5, 4.2.

Nostrils: distance between-inner ends 4.7, 4.7.

326

Memoir Sears Foundation for Marine Research

Labial furrow length: upper 0.8, 0.7.

Gill ofening lengths: ist 2.5, 2.9; 2nd 2.7, 3.O5 3rd 2.7, 3.2; 4th 2.5, 2.9; 5th

2.3,2.5.

First dorsal fin: vertical height 8.2, 9.9; length of base 9.5, 9.7.

Second dorsal fin: vertical height 3.0, 2.9; length of base 3.7, 4.1.

^««/^«; vertical height 3.9, 3.85 length of base 4.5, 5.0.

Caudal fin: upper margin 28.3, 27.4 j lower anterior margin 1 1.7, 1 1.6.

Pectoral fin: outer margin 15.6, 14.85 inner margin 6.0, 5.6; distal margin 11. 1,

12. 1.

Distance from snout to: ist dorsal 29.7, 32.4; 2nd dorsal 60.8, 61.35 upper caudal

71.7, 72.6; pectoral 21.4, 21.9; pelvics 45.6, 47.25 anal 60.2, 60.6.

Interspace between: ist and 2nd dorsals 22.4, 21. 1 5 2nd dorsal and caudal 7.2,

7.45 anal and caudal 6.5, 7.2.

Distance from origin to origin of: pectoral and pelvics 24.5, 27.0 5 pelvics and anal

15.1,13.8.

l^A^cLAAAA^^^^^

Figure 55. Carcharhinus acronotus, immature female, 637 mm. long, from Englewood, Florida (U.S. Nat.
Mus., No. 104331). A Anterior part of head from below, about 0.4. x. B Dermal denticles, about 23 x. C Api-
cal view of dermal denticle, about 46 x. D Left-hand upper and lower teeth, about 1.5 x. £ Third upper tooth.
F Eighth upper tooth. G Third lower tooth. H Eighth lower tooth. E-H, about ^ x. I Left-hand nostril,
about 2 X.

Trunk moderately stout, its height at origin of ist dorsal (where highest) about Vs
its length to origin of caudal, back smooth, without mid-dorsal ridge. Body sector to cloaca
a little shorter than tail sector. Caudal peduncle only slightly compressed laterally. Lower
precaudal pit strongly marked, similar to upper. Dermal denticles mostly overlapping.

Fishes of the Western North Atlantic 327

their leaf-like blades nearly horizontal with usually 3 (sometimes 5) low keels separated
by shallow valleys, their edges with 3 or 5 sharp teeth, the median considerably the
longest.

Head a little more than y^ of total length. Snout rather thin-tipped, narrow-ovoid,
its length in front of a line connecting outer ends of nostrils % to % as great as distance
between inner ends of latter, its length in front of mouth about i V^ times as great as from
front of latter to origin of pectoral. Eye approximately circular, its diameter about % as
great as length of snout in front of mouth, or about % as long as ist gill opening, its mid-
point about opposite front of mouth. First gill opening about % times as long as diameter
of eye, the 3rd slightly longest, 5th a little the shortest, evenly spaced, the 4th above origin
of pectoral. Nostrils strongly oblique, the inner corners nearer to front of mouth than to
tip of snout by a distance equal to about V2 the diameter of eye, the anterior margins
slightly sinuous and expanded near inner end as a prominent triangular lobe with narrowly
rounded tip. Mouth broad-ovoid, about 1.7 times as broad as high in young specimens, but
1.5 times in larger.

Teeth '^or u— n>r2— i2ori3. yppers broadly triangular, their edges serrate, most coarsely
so on outer side of base, their inner margins straight near center of mouth but weakly con-
vex along outer part of jaw, the outer margins deeply notched and increasingly oblique
along the jaw, the outermost teeth very strongly so; lowers with slender cusps and broad
bases, erect in sides of jaw as well as in front, more finely serrate than uppers; one or two
very small erect teeth at symphysis in upper jaw, and one at symphysis in lower; outer-
most 2 or 3 teeth also very small in upper jaw, and with very short cusps in lower.

Origin of ist dorsal over or slightly behind inner corner of pectoral, its anterior mar-
gin moderately convex in young but only slightly so in larger specimens, the rear margin
nearly straight toward apex but concave toward base, the apex narrowly rounded in young
but subacute in adult, the free rear corner only moderately slender, about Vs as long as base;
its vertical height a little greater than length of snout in front of mouth and about 57%
as great as length of pectoral. Second dorsal a little more than V3 as long at base as ist
dorsal, its origin about opposite that of anal, its anterior margin convex in young but nearly
straight in adult, its posterior margin weakly concave, apex broadly rounded in young but
more narrowly so in adult, its free rear corner about as long as base. Caudal a little more
than V4 (27 to 28%) of total length, with slender, subacute tip, the lower anterior lobe
(expanded lower anterior corner) 40 to 45% as long as upper margin, each measured
from its respective precaudal pit, with moderately convex lower anterior margin (more
so in young than in adult), and subacute tip (but rounded in newborn). Anal a little longer
at base than 2nd dorsal, its anterior margin more convex, its posterior margin much more
deeply concave, its apex subacute, its free rear corner about as long as its base, with tip
extending slightly farther rearward than tip of 2nd dorsal and anterior to lower precaudal
pit by a distance % as long as base of anal or about twice as long as diameter of eye. Dis-
tance between origin of anal and tips of pelvics a little longer than base of anal. Pelvics

328 Memoir Sears Foundation for Marine Research

about 1.3 times as long at base as anal, their margins nearly straight. Pectoral about %
(about 67% ) as long as head, and about 1.7 times as long as broad, its outer margin strongly
convex in young but decreasingly so with growth, the distal margin concave for its entire
length, most deeply so in young, the inner margin weakly convex, tip subacute, inner
corner rounded.

Color. Described as variable in lifej some specimens cream-colored or yellowish gray
above, and of a paler shade of the same, or white, below; others uniform brown below as
well as above; the fins are without markings but the tip of the snout is dusky, this nose-
spot being strongest in the young, becoming diffuse or even obscure in adults.

Size. This is one of the smaller members of the genus, maturing while still no longer
than three feet four inches to four feet six inches, and perhaps seldom exceeding a length
of five to six feet. To judge from our Study Material (p. 325) the usual length at birth
is not far from 450 mm.

Developmental Stages. It is not known whether or not there is a yolk-sac placenta in
this species.

Habits. Nothing definite is known of its diet or habits beyond the fact that females,
with embryos (usually three to six) nearly ready for birth, have been taken oflF south-
western Florida from January to April. It is often found in the stomachs of larger sharks.

Relation to Man. It is not nvimerous enough anywhere to be of any commercial im-
portance, or of interest to anglers.

Range. Western tropical and subtropical Atlantic, Rio de Janeiro to North Carolina.
The southwestern coast of Florida is the only region where this species has been reported
in any numbers. It is also known from Rio de Janeiro, from Cuba, from the north coast
of the Gulf of Mexico (Biloxi, Mississippi, one specimen), and from North Carolina,
where stray specimens have been taken from time to time. This suggests that its center of
abundance lies in the Florida-West Indies region, probably including the southern part
of the Caribbean generally and northern coasts of South America to Brazil.

Synonyms and References:

Squalus acronotus Poey, Memorias, 2, i860: 335, pi. 19, fig. 3, 4 (descr., teeth, Cuba).

Platyfodon acronotus Poey, Repert. Fisico.-Nat. Cuba, 2, 1868: 450 (Cuba); An. Soc. esp. Hist, nat., 5,

1876: 389; Enumerat. Pise. Cubens., 1876: 193 (Cuba).
Carcharias {Prionodon) acronotus Gunther, Cat. Fish. Brit. Mus., 8, 1 870: 369 (Cuba).
Carcharhinus {Pktyfodon) acronotus Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1 896: 216

(Cuba); Bull. U.S. nat. Mus., 47 (i), 1896: 36 (descr., Cuba).
Carcharinus acronotus Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 136 (descr., Cuba); Gudger,

J. Elisha Mitchell sci. Soc, 28, 1913: 158 (N. Carolina); Coles, Proc. biol. Soc. Wash., 28, 1915:

go (N. Carolina) ; Radcliffe, Bull. U.S. Bur. Fish., 5^, 1916: 259, pi. 41, fig. I, 2 (denticles, teeth, N.

Carolina); Smith, J. Amer. Mus. nat. Hist., t6, 1916: 348 (Cuba, N. Carolina); Nichols, Bull. Amer.

Mus. nat. Hist., 37, 1917: 875, pi. Ill, fig. 2 (south. Florida); Breder, Field Bk. Mar. Fish. Atlant.

Coast, 1929: 16 (Florida, W. Indies) ; White, Bull. Amer. Mus. nat. Hist., ■J4, 1937: 127, pi. 50, fig. d

(in Key, cartilages of claspers) ; Springer, Proc. Fla. Acad. Sci., 5, 1 939: 21 (Florida, embryos, color);

Hildebrand, Copeia, 1941: 221 (N. Carolina); Lunz, Bull. S. Carolina St. Planning Bd., 14, 1944: 27

(Florida).

Fishes of the Western North Atlantic 329

Carcharias acronotus Jordan, Manual Vert. Anim. NE. U.S., 1929: 10 (general); Jordan, Evermann and

Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 16 (Cuba).
Eulamia acronotus Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1945: 95 (N. Carolina, but 8-ft. specimen of

250 pounds is perhaps not this species).

Carcharhinus falciformis (Miiller and Henle), 1841

Figures 56, 57

Study Material. Five immature specimens, males and females, 782 to 1,065 "^^n-
long, from offing of Delaware Bay and of Cape Fear, North Carolina, and from the Baha-
mas (U.S. Nat. Mus. and Harv. Mus. Comp. Zool.) }* jaws, measurements of embryos,
and photographs of latter from Florida Keysj' a skin, without locality (Harv. Mus. Comp,
Zool., No. 1384); also photographs of Poey's unpublished drawings of this species.

Distinctive Characters. Among the ridge-backed members of the genus, falciformis
resembles floridanus in the very elongate free tips of the second dorsal and anal, and in
the teeth, but it has a much shorter pectoral, a larger eye and a more broadly ovate snout.

Description. Proportional dimensions in per cent of total length. Female, 785 mm.,
fromLat. 33° 38' N., Long. 77° 36' W. (U.S. Nat. Mus., No. 38510). Male, 848 mm.,
from Lat. 38° 37' N., Long. 73° 1 1' W. (U.S. Nat. Mus., No. 35643)-

Trunk at origin of -pectoral: breadth 1 1.2, i i.O; height 1 1.4, 9.6.

Snout length in front of: outer nostrils 4.5, 4.4; mouth 8.7, 8.0.

Eye: horizontal diameter 1.9, 1.8.

Mouth: breadth 7.9, 7.65 height 5.4, 5.1.

Nostrils: distance between inner ends 5.8, 5.7.

Labial furrow lengths: upper 0.6, 0.5.

Gill o-pening lengths: ist 2.2, 2.1 j 2nd 2.3, 2.4; 3rd 2.3, 2.5; 4th 2.2, 2.4; 5th

1.8, 1.8.

First dorsal fin: vertical height, 7.3, 7.55 length of base 8.5, 8.6.

Second dorsal fin: vertical height 2.0, 1.95 length of base 3.3, 3.4.

Anal fin: vertical height 3.0, 2.9 ; length of base 3.5, 3.5.

Caudal fin: upper margin 27.7, 28.2 j lower anterior margin 10.9, 1 1.6.

Pectoral fin: outer margin 15.3, 14.7; inner margin 4.2, 4.25 distal margin 12.2,

II. 8.

Distance from snout to : ist dorsal 33.9, 32.75 2nd dorsal 63.2, 60.35 upper caudal

72.3, 71.85 pectoral 24.5, 23.O5 pelvics 50.3, 47.75 anal 63.0, 60.7.
Inters-pace between: ist and 2nd dorsals 22.5, 21.65 ^nd dorsal and caudal 6.6.,
7.O5 anal and caudal 6.2, 6.1.

Distance from origin to origin of: pectoral and pelvics 26.8, 26.25 pelvics and anal,

13.4, 14.8.

6. Lat. 38° N., Long. 73° 11' W., Sept. 12, 1884, off Delaware Bay, male (U.S. Nat. Mus., No. 35643) i Lat. 33°
38' N., Long. 77° 36' W., off Cape Fear, North Carolina, Oct. 20, 1885, "Albatross" Sta. 2617 and 2623, 1
male, 2 females (U.S. Nat. Mus., No. 38509, 38510); female, Bahamas.

7. Contributed by Stewart Springer.

330

Memoir Sears Foundation for Marine Research

Figure 56. Carcharhinus falciformis, immature male, 848 mm. long, off Delaware Bay (U. S. Nat. Mus.,
No. 35643). A Head from below, about Y2 natural size. B Second dorsal fin, about I x. C Dermal denticles,
about 36 X. D Apical view of dermal denticle, about 72 x. E Left-hand nostril, about 2.5 x.

Figure 57. Carcharhinus jalcijormis, illustrated in Fig. 56. j4 Right-hand upper and lower teeth, about
2.5 X. B Fifth upper tooth. C Ninth upper tooth. D Second lower tooth. E Tenth lower tooth. B-E, about
7.5 X.

Fishes of the Western North Atlantic 331

Trunk slender, its height at origin of ist dorsal about Vfi its length to origin of
caudal. Midline of back with a low but unmistakable dermal ridge from close behind tip
of first dorsal to a point about as far in front of origin of 2nd dorsal as length of base of
latter." Upper precaudal pit subrectangular in outline, the lower pit more obtusely angular
and only faintly marked. Dermal denticles small, regularly overlapping, with 3 or some-
times 5 low ridges, the apical margin with a corresponding number of teeth, the axial only
a little larger than the pairs flanking it.

Head a little less than % (22 to 24%) of total length, nearly as wide at eyes as at
origin of pectorals, and about % as wide opposite outer ends of nostrils as at eyes. Snout
rather thin-tipped, broadly rounded, its length in front of a line connecting outer ends of
nostrils about 70% as great as distance between inner ends of latter, its length in front of
mouth about V3 of length of head. Eye nearly circular, its diameter about Vs as long as dis-
tance between inner ends of nostrils. First gill opening about i % times as long as diameter
of eye, the 3rd slightly longest, the 5th about % as long as ist, the 4th and 5th more nar-
rowly spaced than ist to 4th, the space between 3rd and 4th above origin of pectoral. Nos-
tril strongly oblique, its inner end nearer to mouth than to tip of snout by a distance about
equal to diameter of eye, the anterior margin only very slightly expanded toward inner
end, without definite lobe. Mouth ovate, about 70% as high as broad.

Teeth ''^"^"' ;° upper teeth broadly triangular, the ist tooth erect and nearly
symmetrical, with both margins slightly concave and strongly serrate, especially toward
base, the 2nd and successive teeth increasingly oblique with inner margins slightly
convex and outer margins notched more and more deeply toward angles of jaws, their
inner edges serrate, their outer margins similarly so on cusps and even more coarsely
so on basal sectors where the ist one or two serrations are the most prominent} lower
teeth with narrow, lanceolate cusps on broad bases, symmetrical and erect along entire
jaw, their edges smooth, or at most slightly wavy or irregularly serrate toward tips;
I or 2 very small symmetrical teeth with notched margins at symphysis in upper jaw, and
I minute, slender, symmetrical tooth in lower; outermost teeth in each jaw also very small.

Vertical height of ist dorsal about as great as length of snout in front of mouth, or
about % length of pectoral, its origin a little posterior to inner corner of pectoral, its
anterior margin weakly convex, the apex moderately to broadly rounded, the posterior
margin convex distally but very deeply concave proximally, the free rear corner only mod-
erately slender, a little less than ^2 as long as base. Second dorsal only about V3 as long at
base as ist dorsal, its origin about over or a little anterior to that of anal," its anterior mar-
gin nearly straight, posterior margin moderately concave, its apex rounded, the free rear
corner very slender and noticeably elongate, being more than twice as long as the vertical
height, and a little longer than the base, extending nearly % of the distance from rear end
of base toward precaudal pit. Caudal between V4 and V3 of total length, its upper margin

8. In preserved specimens this ridge may lie along a deep longitudinal furrow of muscular contraction.

9. Fourteen teeth could be counted in the side of each jaw in one specimen, with perhaps another very small one at
the corner.

10. It is difficult to determine its point of origin exactly in the specimens seen.

332 Memoir Sears Foundation for Marine Research

moderately convex, its terminal sector only about Vs of total length of fin, slender, with
rounded tip and weakly concave lower margin, its lower lobe (expanded anterior corner)
about % (42%) as long as upper lobe with convex lower anterior margin and narrowly
rounded tip ; general posterior re-entrant contour, included by the two lobes, more broadly
rounded than in C. fioridanus (cf. Fig. 56 with 58). Anal about as long at base as 2nd dor-
sal, or a little longer, with similarly slender free rear tip, about i V2 times as long as the
vertical height and about as long as the base, but with much more deeply incised rear mar-
gins and more broadly rounded apex. Distance from origin of anal to tips of pelvics about
2V2 times as long as base of anal. Pelvics a little longer at base than anal, with nearly
straight edges, their origins about midway between origins of ist and 2nd dorsals. Pectoral
only about % as long as head (about as long as head in fioridanus) and only about twice
as long as vertical height of ist dorsal, a little more than twice as long as broad, the outer
margin weakly convex toward tip, distal margin moderately concave, inner margin only
weakly convex, inner corner and apex narrowly rounded.

Color. Described as dark gray above, grayish white below j those we have seen after
preservation are mouse gray above and a paler shade of the same tint below.

Size. The claspers of the males listed above have not yet reached the tips of the pelvic
fins, and a female of 7% feet has been found to contain embryos.^^ These facts suggest that
falciformis matures at a length of perhaps six feet, but it is said to attain a length of about
10 feet (3,050 mm.).'^

Developmental Stages. It is not yet known whether this is an ovoviviparous or a vi-
viparous species; its embryos have been reported only once."

Habits. While several falciformis have been taken on set lines along the reef off
Metacumbe Key in southeastern Florida during summer in about 60 feet of water, none
have ever been reported in the passages between the Keys, in spite of the great amount of
angling that is done from the bridges that span the latter; similarly, one of the three
records of it farther to the north was from the outer edge of the continental shelf, the
other two being from about 1 7 to 20 miles out from the nearest land, although taken in
shoal water of 14 to 15 fathoms. Thus it appears that this is an oflFshore species, not to be
expected close to the beach unless as a stray. Nothing further is known of its habits, and
nothing of its diet."

Relation to Man. Falciformis is not caught anywhere in numbers large enough to
make it of commercial importance.

Range. Both sides of the Atlantic, in waters of high temperature; Goree, West Africa,
on the one side, West Indies to the offing of Delaware Bay on the other. So far known in
the western Atlantic from: Trinidad (nominal record only); Porto Rico; Haiti; Ber-
muda; Cuba; Bahamas; east coast of Florida (Salerno), where half -grown specimens as

11. Personal communication from Stewart Springer.

12. 3,050 mm. (Fowler, Bull. Amer. Mus. nat. Hist., 70 (i), 1936; 49).

13. Personal communication from Stewart Springer.

14. The head of a menhaden (Brevoortia), found in the stomach of one of the present series taken off Cape Fear,
North Carolina, may have been a bait taken from a hook before this Shark was caught.

Fishes of the Western North Atlantic 333

well as adults with embryos have been taken often enough in summer to mark it as common
outside the reefs; two stations 17 to 20 miles off Cape Fear, North Carolina, in October
1885; and the outer edge of the continental shelf off Delaware Bay, in September 1884.
The geographic distribution of these localities, together with the fact that the more north-
erly captures in September and October were at stations where the temperature at the sur-
face (where it is probable that the specimens were caught) was 74° to 75° F., shows this to
be a tropical-subtropical species, occasionally straying northward along the coast of the
United States during the late summer and early autumn. It is to be expected throughout
the Caribbean region generally, and at least as far to the south as northern Brazil, if not
farther.

Synonyms and References:

Carchoruis {Prionodon) fdciformis Miiller and Henle, Plagiost., 1 841: 47 (descr., Cuba); Dumeril, Hist.
Nat. Poiss., I, 1865: 374 (descr. of type, Cuba); Gunther, Cat. Fish. Brit. Mus., 8, 1870: 363, foot-
note (C?uba); Steindachner, Denkschr. Aicad. Wiss. Wien., 45 (l), 1 882: 14 (Goree, W. Africa);
Metzelaar, Trop. Atlant. Visschen, 1919: 187 (both coasts of Atlantic).

Prionodon falciformis Guichenot, in Sagra, Hist. Cuba, 1855: 248, pi. 5, fig. 3 (ill., Cuba, not seen);
Poey, Repert. Fisico.-Nat. Cuba, 2, 1868: 172 (discus.).

Squalus tiburo Poey, Meraorias, 2, i860: 331, 334 (descr., Cuba); Repert. Fisico.-Nat. Cuba, 2, 1868: 172^'
(Cuba) ; not Squalus tiburo Linnaeus, 1758.

Prionodon tiburo Poey, Memorias, 2, i860: pi. 19, fig. I, 2 (teeth, Cuba).

Platyfodon tiburo Poey, Repert. Fisico.-Nat. Cuba, 2, 1 868: 448, pi. 4, fig. 18 (teeth, Cuba).

Plotyfodon falciformis Poey, Repert. Fisico.-Nat. Cuba, 2, 1868: 449; Ann. Soc. esp. Hist, nat., 5, 1876: 387;
Enumerat. Pise. Cubens., 1876: 191 (discus., refs. to Muller and Henle, 1841, and to Dumeril, 1865).

Carcharhinus {Platyfodon) falciformis Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 216
(Cuba); Bull. U.S. nat. Mus., 47 (i), 1896: 36 (descr., Cuba).

Carcharhinus falciformis Evermann and Marsh, Bull. U.S. Bur. Fish., 20 (i), 1902: 62 (meas., Porto Rico;
one of two specimens, the other being C. floridanus Bigelow, Schroeder and Springer, 1 943: 71) ; Vincent,
Sea Fish. Trinidad, 1910: 53 (name only, Trinidad) ; Garman, Mem. Harv. Mus. comp. Zool., 36, 1913:
129 (descr., Cuba, West Indies); Beebe and Tee-Van, Zoologica, N. Y., 10 (6), 1928: 28 (Haiti,
meas.) ; Nichols, Sci. Surv. Porto Rico, N. Y. Acad. Sci., 10 (2), 1929: 183 (Porto Rico) ; White, Bull.
Amer. Mus. nat. Hist., 74, 1937: 125 (in Key).

Carcharias falciformis Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 16 (Cuba
and neighboring w/aters) ; Beebe and Tee-Van, Zoologica, N. Y., 13, 1932: 119 (Bermuda); Field Bk.
Shore Fish. Bermuda, 1933: 27 (Bermuda).

Eulamia falciformis Fowler, Bull. Amer. Mus. nat. Hist., 70 (l), 1936: 49 (trop. W. Afr., max. size) ; Bige-
low and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 78, fig. 23
(descr., range, ill.).

Carcharhinus floridanus Bigelow, Schroeder and Springer, 1943

Silky Shark

Figures 58, 59

Study Material. Type, female about 8 feet (2,328 mm.) long, taken off Fort
Pierce, Florida, in about 100 feet of water on November 2, 1942 (Harv. Mus. Comp.

15. Poey concluded that tiburo was a synonym of falciformis, but later (ibid., p. 448) questioned this. However,
there appears to be nothing to difierentiate it from falciformis.

334 Memoir Sears Foundation for Marine Research

Zool., No. 35807); also jaws, fins, photographs and measurements of a female about 8
feet long caught at the surface from the "Atlantis" in Cochinas Bay, Cuba, on April 5,
1939 (Harv. Mus. Comp. Zool., No. 35515).

Distinctive Characters. Among the ridge-backed members of the genus, with which
it falls, floridanus most closely resembles falciformis in the elongate free rear tips of the

"W^^^^^^^^^^^^^

^^

A'6

Figure 58. Carcharhinus floridanus, adult female, about eight feet long, from Fort Pierce, Florida (Harv.
Mus. Comp. Zool., No. 35807). A Right-hand nostril, about l x. 5 Dermal denticles, about 26 x.
C Left-hand teeth, upper and lower, about Y3 natural size. D Fourth upper tooth. E Sixth upper tooth.
F Fourth lower tooth. G Eleventh lower tooth. D—G, about 1.3 x.

Figure 59. Carcharhinus floridanus, pictured in Fig. 58. j1 Anterior part of head from below, about % x.
B Second dorsal and anal fins, about Yi natural size.

Fishes of the Western North Atlantic 335

second dorsal and anal, and in its teeth. But it is easily separable from falciformis by its
much longer pectorals, much smaller eye, and the more narrowly ovate snout (cf. Fig.
56 with 58).

Descriftion. Proportional dimensions in per cent of total length. Female, 2,328 mm.,
from Fort Pierce, Florida (Harv. Mus. Comp. ZooL, type. No. 35807).

Snout length in front of: outer nostrils 4.0; mouth 7.2.

Eye: horizontal diameter 1.2.

Mouth: breadth 8.35 height 4.4.

Nostrils: distance between inner ends 5.5.

Labial furrow length: upper 0.5.

Gill opening lengths: ist 2.9; 2nd 3.O} 3rd 3.35 4th 3.15 5th 2.6.

First dorsal fin: vertical height 8.6; length of base 8.7.

Second dorsal fin: vertical height 2.2; length of base 2.8.

Anal fin: vertical height 3.6; length of base 3.2.

Caudal fin: upper margin 26.5; lower anterior margin 13.6.

Pectoral fin: outer margin 20. 7; inner margin 5.O; distal margin 16.7.

Distance from snout to: ist dorsal 30.O; 2nd dorsal 63.7; upper caudal 73.5; pec-
toral 20.7; pelvics 48.5; anal 63.7.

Interspace between: ist and 2nd dorsals 25.8; 2nd dorsal and caudal 7.1; anal and

caudal 5.8.

Distance from origin to origin of: pectoral and pelvics 27.8 ; pelvics and anal 15.7.

Trunk rather slender and tapering evenly both anteriorly and posteriorly, its height
at origin of ist dorsal only about Ve its length to origin of caudal. Body sector to cloaca
considerably longer than tail sector. A low but unmistakable dermal ridge along midline
of back extending part way between ist and 2nd dorsal fins. Lower precaudal pit, as well as
upper, well marked. Dermal denticles so small and flat that the skin feels silky, close
set, regularly overlapping, those on trunk usually with 7 ridges but occasionally with only
6, the marginal teeth short and rather blunt, those on head minute, less strongly sculp-
tured and broadly oval.

Head about Vs of total length. Snout flattened above, its tip narrow-ovate, its
length in front of a line between outer ends of nostrils a little less than % (71%) as
great as distance between inner ends of latter, the length in front of mouth about Vs
of length of head. Eyes round and noticeably small, the horizontal diameter only about
V4 to /'5 as long as distance between nostrils, or Ve the length of snout in front of
mouth. Gill openings evenly spaced, of medium length, the first a little more than
twice as long as diameter of eye, the 3rd a little the longest and 5th a little the shortest,
the 3rd, or space between 3rd and 4th, above origin of pectoral. Nostril oblique, its
inner end nearer to upper jaw than to tip of snout by a distance aboit twice as great as
diameter of eye, the anterior margin without definite lobe but only somewhat sinuous
in outline. Mouth broad-ovate, its length slightly less than V2 its breadth.

336 Memoir Sears Foundation for Marine Research

Teeth ^^-^jEjotI^is"^^"' "PP^'"S triangular, increasingly oblique toward corners of
mouth, their outer edges more or less notched, with some toward center of jaw notched
on both sides, the edges serrate, very finely toward tips and somewhat more coarsely
toward bases, but very much worn;'* lower teeth smaller, more slender and more nearly
erect, symmetrical, with narrow cusps and broad bases, their margins smooth toward bases
and slightly wavy toward tips but not definitely serrate; 2 or 3 minute teeth at symphysis
in both jaws.

Origin of ist dorsal over, or slightly posterior to, inner corner of pectoral, its vertical
height about Vs as great as length of head to 5th gill opening, its apex rounded, its rear
margin deeply concave basally, its free rear tip very slender and about % is long as its base.
Origin of 2nd dorsal about over origin of anal, its vertical height only about V4 to Vs as
great as that of ist dorsal, its posterior margin only weakly concave, its free rear tip very
slender and greatly elongate, it being about 2 V2 times as long as the vertical height of the
fin, or I V2 times as long as its base, and extending about % the distance from rear end of
base toward precaudal pit. Caudal a little more than V4 of total length, its terminal sector a
little less than Vs total length of fin, slender, with concave lower margin and narrowly
rounded tip, the lower lobe (expanded lower anterior corner) about V2 as long as upper,
with subacute tip and moderately convex anterior margin, the re-entrant corner included
between the two lobes subrectangular (rounded in falclformis). Anal a little longer at
base than 2nd dorsal, with much more deeply incised posterior and more broadly rounded
posterior margins, its free rear tip slender, a little longer than its base, about i V2 times its
vertical height, and terminating about under tip of 2nd dorsal. Distance from origin of
anal to tips of pelvics nearly or quite twice as long as base of anal. Pelvics about i ^2 times as
long at base as anal, with weakly concave distal margins, their origin about midway
between origins of ist and 2nd dorsals. Pectoral nearly or quite as long as head or about
2V2 times as long as vertical height of ist dorsal, a little less than V2 as broad as long, the
outer margin moderately convex, distal margin deeply concave basally, the apex and
inner corner narrowly rounded.

Color. The type specimen was shiny black above when fresh caught, dirty white
below; the tips of the pectorals somewhat dusky below; the Cuban specimen was dark
gray above in life.

Size. This is one of the larger members of the genus; all the specimens so far
reported have been between about eight and ten feet long.

Developmental Stages. Embryos of this species have been reported but not described.

Habits. Little is known of the habits of this newly described species except that ofiF
Salerno, Florida, it is usually taken only where lines are set at a depth of 100 feet or
more. While one large individual was taken on the reef near Metacumbe Key in about
45 feet of water, it was so thin as to suggest that its excursion into relatively shoal water
was abnormal.

16. Those of the Cochinos Bay specimen are more definitely notched than those of the type specimen, and with
coarser basal serrations.

Fishes of the Western North Atlantic 337

Relation to Man. Such specimens as are caught in the Florida fishery (see below) are
used for leather, etc., like other large sharks.

Range. It is known to the present time from the south coast of Cuba (see Study
Material, p. 333) and the north coast near Havana," Porto Rico, and southeastern
Florida. Hence it probably occurs generally throughout the tropical belt of the western
Atlantic. Evidently it is plentiful locally, for it is taken so frequently in the shark fishery
that is now carried on from Salerno, Florida, that it has been given the vernacular name
"Silky Shark," appropriate because of the small size of its dermal denticles. As many
as 60 adults of nine to ten feet were caught there in a single day during the winter of
1943, making it dominant in the total catch of sharks of all sorts for the time being. It is
less numerous there in summer.^' That a shark so common, so large and so easily recognized
should have continued unknown for so long casts an unflattering light on scientific knowl-
edge of the group to which it belongs.

Synonvins and References:

Carcharkinus jalcijormis (in part) Evermann and Marsh, Bull. U.S. Fish Comm., 20 (l), 1902: 6?. (meas.,

Porto Rico, one of two specimens, the other being correctly identified as C. falciformis Muller and Henle,

1841).
Carcharinus foridanus Bigelow, Schroeder and Springer, Proc. New Engl. zool. CL, 22, 194.3: 71, pi. 13 (ill.,

descr., Salerno, Florida, type loc, and south coast of Cuba).
Eulamia fioridanus Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo ."^mer. Caribb. Comm., Wash.,

1945: 76, fig. 22 (descr., range, ill.),

Carcharhinus leucas (Muller and Henle), 1841"

Cub Shark, Bull Shark, Ground Shark

Figures 60, 61

Study Material. Female, 924 mm. long, from Florida (Amer. Mus. Nat. Hist.)j
skin of a female, about 1,137 mm-j at Miraflores Locks, Panama Canal (U.S. Bur. Fish.,
No. 1 3961); headof specimen, about 6 /'o feet, from Bahamas (from Daniel Merriman);
embryo, 435 mm. long, from Cuba (Harv. Mus. Comp. Zool., No. 722)} tracings of
fins of an adult male, 2,310 mm., from Metacumbe, Florida (from Stewart Springer);
male embryos, 490 mm. (Harv. Mus. Comp. Zool.) and 625 mm. (U.S. Nat. Mus., No.
108,456), from Englewood, Florida; female, 692 mm. long, either embryo or newborn to
judge from the umbilical scar, also female, 920 mm., from Lake Yzabal, Guatemala (U.S.
Nat. Mus.).

Distinctive Characters. Leucas is separable from all other Atlantic carcharhinids
except Negaprion brevirostris, Carcharhinus longimanus and C. nicaraguensis by its ex-

17. Personal communication from Luis Howell-Rivero. i8. Personal communication from Stewart Springer.

19. The account of leucas by Miiller and Henle (Plagiost., 1S41: 42) agrees in detail with the present species,
except that the dried specimens on whicli it was based were described as white above as well as below. But their
paleness (probably from fading) was perhaps exaggerated, for the same specimens (presumably) were described
later as whitish gray (Dumeril, Hist. Nat. Poiss. /, 1S65; 35S).

338

Memoir Sears Foundation for Marine Research

tremely short, very broadly rounded snout and smooth back (without dorsal ridge).
Its second dorsal is much smaller than the first and its teeth are regularly serrate on the
cusps, features which mark it off from A^. brevirostrisi from C. longimanus it is separated
by the shape of its dorsal with subacute apex, by its relatively shorter and broader pectorals,
and by the long interspace between the tip of its anal and its caudal, but relatively much

,c£^k^k3.

Figure 6o. Carcharhinus leucas, immature female, about 924 mm. long, from southern Florida (Amer. Mus.
Nat. Hist.). A Anterior part of head, about 0.3 x. B Left-hand upper and lower teeth, about 1.3 x. C Third
upper tooth. D Ninth upper tooth. E Second lower tooth. F Eighth lower tooth. C-F, about 2.6 x. G Right
nostril, about 2 x.

Figure 6 1 . Dermal denticles of Carcharhinus leucas
pictured in Fig. 60, about 45 x.

Fishes of the Western North Atlantic 339

shorter distance from origin of anal to tips of pelvicsj C. nkaraguensis appears to be a
landlocked variant of it (see discussion, p. 381).

Description. Proportional dimensions in per cent of total length. Female, 924 mm.,
from Florida (Amer. Mus. Nat. Hist.).

Trunk ai origin of pectoral: hrea.dth. 12.8} height 12.9.

Snout length in front of: outer nostrils 2.0} mouth 6.6.

Eye: horizontal diameter 1.5.

Mouth: breadth 9.85 height 4.9.

Nostrils: distance between inner ends 6.7.

Gill opening lengths : ist 2.95 2nd 3.25 3rd 3.2; 4th 2.9} 5th 2.4.

First dorsal fin: vertical height 9.6; length of base 13.4.

Second dorsal fin: vertical height 4.1 j length of base S-S-

Anal fin: vertical height 4.9; length of base 5.3.

Caudal fin: upper margin 28.35 lower anterior margin 12.3.

Pectoral fin: outer margin 20.6; inner margin 6.6:, distal margin 17.6.

Distance from snout to: ist dorsal 27.O; 2nd dorsal 60.O; upper caudal 71.7;

pectoral 21.4; pelvics 48.3; anal 60.0.

Interspace between: ist and 2nd dorsal 21.O; 2nd dorsal and caudal 7.0; anal and

caudal $.$•

Distance from origin to origin of: pectoral and pelvics 28.4; pelvics and anal 1 1.3.

Trunk moderately stout, its height at origin of ist dorsal about Vs of its length to
precaudal pits, its breadth at origin of pectorals about equally great, its dorsal profile
more convex than the ventral. Midline of back smooth, without dermal ridge, either in em-
bryo or in adult. Anterior outline of upper caudal pit obtusely subangular in smaller
specimens, the lower caudal pit less well marked than upper but similar in outline. Dermal
denticles moderately raised so that skin feels slightly rough when stroked toward head,
loosely spaced and overlapping but little, as broad as long, or a little broader, usually with
3 low ridges in smaller specimens, but perhaps more often 5 in larger, with 3 prominent
teeth (the axial considerably the largest) separated by rounded notches; if there are 5
teeth, the outermost pair are very small; pedicels very short and broad.

Head very wide, its breadth as great opposite corners of mouth as at origin of
pectoral or slightly greater and only a little narrower at eyes. Snout thick, very broadly
rounded and notably short, its length in front of a line connecting outer ends of nostrils
only between y^ and V2 (30 to 42%) as great as distance between inner ends of latter
in specimens seen, its length in front of mouth a little less than Va of length of head to
origin of pectoral. Eye approximately circular, small, its diameter a little less than ^
as great as distance between inner ends of nostrils. Longest gill openings (2nd and 3rd) a
little more than twice as long as diameter of eye or about half as long as distance between
inner ends of nostrils, the 5th only a little more than % that long; all about evenly
spaced, the 4th about over origin of pectoral. Nostril strongly oblique, its inner end

340 Memoir Sears Foundation for Marine Research

nearly or quite as close to tip of snout as to mouth and about equidistant between rear
edge of eye and tip of snout j its anterior margin slightly expanded in obtusely triangular
outline with rounded corner. Mouth broadly ovate, its height about ^ its breadth. Upper
labial furrow very short even for this genus, and at approximately a right angle with the
jaw.

^^^^^ ia or 13—1^12071!' uppsfs broadly triangular, both edges regularly and moder-
ately coarsely serrate from base to tip, the first 2 teeth erect and nearly symmetrical, with
both margins slightly concave, but 3rd and subsequent teeth with only very slightly con-
cave or nearly straight inner margins, and outer margins increasingly concave toward cor-
ners of mouth, the outermost 3 or 4 teeth rather definitely notched j lower teeth with
narrow triangular cusps on broad bases, nearly erect in front and sides of mouth and only
slightly oblique toward its corners, the cusps successively shorter from 8th or 9th to 12th
or 13th, both edges serrate, but more finely so than on uppers. One small symmetrical
tooth at symphysis in each jaw.

First dorsal noticeably large, its origin about over or a little anterior to axil of
pectoral, its vertical height about % (40-45%) as great as distance from tip of snout to
origin of pectorals, its base nearly as long as its anterior margin, its anterior margin
nearly straight, its apex subacute or very narrowly rounded, its posterior outline moder-
ately concave (a little the more so basally), its free rear corner relatively obtuse and only
about V4 as long as its base, the midpoint of latter only a little more than Ys as far from
axil of pectoral as from origin of pelvics. Second dorsal about % (41%) as long as ist
at base, relatively lower, its origin over or a little anterior to that of anal,^" its anterior
outline nearly straight, apex rounded, posterior outline only weakly concave, its free rear
corner broad and about V2-% as long as its base, its rear tip definitely though only a little
anterior to that of anal and separated from origin of caudal by a distance about % as long
as its base. Caudal with weakly convex upper margin and subacute tip, the narrow-tri-
angular terminal sector about Y^ the total length of fin, the lower lobe (expanded lower
anterior corner) a little less than Y2 (about 44%) as long as upper, with moderately con-
vex anterior outline, nearly straight posterior outline and narrowly rounded tipj the
re-entrant corner, included by the 2 lobes, broadly rounded. Distance from origin of
caudal to tip of anal about V2 as long as base of anal (a convenient field mark separating
this species from longhnanus) . Anal with base and free rear corner about as long as those
of 2nd dorsal, but anterior margin about i Vs times as long and more convex, and posterior
edge much more deeply incised. Distance from origin of anal to tips of pelvics only about
Y2 to % as long as base of former (longer than base of anal in longimanus) . Pelvics with
nearly straight anterior margins and slightly concave distal margins, their bases a little
longer than base of anal. Pectoral nearly as long (about 87%) as head to origin of
pectoral, about Y2 as broad as long, the outer margin nearly straight toward base but

io. The origin of the second dorsal is about over that of the anal in the female studied, but a little anterior to it in
male embryos and in a tracing of an adult male contributed by Stewart Springer.

Fishes of the Western North Atlantic 341

moderately convex toward tip, the distal margin only weakly concave, the inner corner
moderately rounded, the tip more narrowly so.

Color. Described in life as white below, gray above, varying from very pale to much
darker, apparently as the result of environmental conditions, for those living over white
sand bottom may be very pale. Adults show no conspicuous fin markings. But in embryos
the tip and lower edge of the caudal and the margin of the second dorsal are sooty, and
perhaps the tips of the other fins likewise, in some cases, and these fin markings may per-
sist for a considerable time after birth, at least in some instances, for a 32-inch specimen,
apparently of this species, has been described as with second dorsal and anal dusky-tipped,
and the caudal wholly so."

Size. Mature at a length of about seven feet, this species certainly grows to 10 feet
and perhaps somewhat longer, but reports of specimens longer than 12 feet may have
referred to C. longhnanus, with which it has often been confused. Specimens of 8 to 8^/4
feet have been reported as weighing 250 to 375 pounds; the weight of a male of 10 feet
caught off North Carolina is given as about 400 pounds." It has been suggested that the
three-foot Florida specimen illustrated in Fig. 60, taken in winter, was a yearling,"
and it is probable that the young are born there in spring. Also, the advanced stage of
development of the embryos listed above indicates a length of perhaps 650 to 700 mm. as
usual at birth.

Developmental Stages. Presumably development is viviparous, but the presence of
a yolk-sac placenta has not been definitely recorded for this species, so far as we are
aware. Embryos have relatively stouter bodies and blunter heads than their parents, but
they do not differ nearly so much from the adults in the shapes and relative sizes of the
fins as do those of longimanus ; like the adults, they are smooth-backed. Five or six
appear to be the usual number of young in a litter.

Habits. This is a heavy, slow-swimming species, most common inshore in shoal
water, perhaps never very far from land except by accident. They are most often caught
around docks, at the entrances to the passages between islands, in estuaries and in harbors.
They often run up rivers for considerable distances, and it seems that they do not hesitate
to enter fresh water. Thus the series we have studied includes one from the Panama Canal
at Miraflores Locks, besides others from Lake Yzabal, Guatemala, a body of water that is
said to vary between fresh and brackish, and a 55-pound specimen has been reported, at
least by name, as having been caught in the Atchafalaya River, Louisiana, 1 60 miles from
the sea." We have also received a photograph of a shark four or five feet long that appears
to be of this species (unless possibly of the landlocked form nicaraguensis, which cannot be
determined from the photograph), taken 180 miles up the Patuca River, northeastern
Honduras." C. leucas is, in fact, the only Shark that is known to have permanently adapted

21. Jordan and Gilbert, Proc. U.S. nat. Mus., 5, 1882: 243. 22. Bell and Nichols, Copeia, 92, 1921 : 17.

23. Nichols, Bull. Amer. Mus. nat. Hist., 57, 1917: 874. 24. Gunter, Copeia, 1938: 69.

25. This capture has already been reported, without attempt at specific identification (Strong, Explor. Smithson.
Instn. [1933], 1934: 46> 47i fig- 5s).

342 Memoir Sears Foundation for Marine Research

itself to fresh water and developed a local race (see under C. nicaraguensis, p. 381).
On the other hand, it rarely shows itself at the surface, as the more pelagic members of
the genus commonly do, unless lured up by the scent of food, such as floating offal. We
have never heard of one jumping, whether at liberty or after being hooked.

Under natural conditions its diet is perhaps no less varied than that of the Tiger
Shark (p. 270). Thus the stomach contents of a series taken off North Carolina in
summer have been reported as including crabs, smaller sharks that had been taken in
the same net, the fin of a porpoise which was probably obtained in the same way, shad
(Alosa) and mackerel. That they had eaten the last-named proves that they have a
greater ability to capture fast-swimming fishes than their rather sluggish habits would
suggest. Also found in the stomach were large pieces of devil-rays {Mobula), which,
when fitted together, showed the victim to have been bitten into five parts. It also attacks
sting-rays in Florida waters, as evidenced by a spine imbedded in one's mouth. It is
notorious also as a scavenger, devouring any kind of offal, and it bites readily on almost
any large bait of fish or meat.

Females with embryos nearly ready for birth have been taken in Florida in October,
January and February, which suggests that the young are born there in late winter and
early spring, but beyond this nothing is known of the breeding habits.

Relation to Man. The only commercial importance of this species is that it con-
tributes something to the general catch of sharks around Florida. But no data are available
to show its importance relative to other species. Like various other sharks it has been
named a man-eater, but we think it unlikely that this reputation is deserved, for other-
wise shark fatalities probably would be far more frequent than they actually are in
Florida and the West Indies, where it is one of the more common of the larger sharks.

Range. In the western Atlantic from southern Brazil to North Carolina and occa-
sionally north to the vicinity of New York. It is not yet certain whether leucas occurs at
all in the eastern Atlantic. It is true that sharks have been recorded under that name from
tropical West Africa and from Algeria, but the first of these reports was by name only,"
while the brief description accompanying the second" would apply equally well to any
other Carcharhinus with moderately blunt snout, for example, longimanus, milberti or
obscurus. The commersonii described by Rey^^ from the coast of Spain was doubtless an
obscuruSy for it had a mid-dorsal ridge and agreed with that species in other respects as
well. Nor is any Carcharhinus other than longimanus included in the most recent survey
of the fishes of Portugal.^*

Occurrence in the Western Atlantic. Published accounts are in line with word-of-
mouth reports to the effect that leucas is a tropical species, expanding its range northward
during the warm months and perhaps southward as well. But it is not yet possible to

26. Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 42; Metzelaar, Trop. Atlant. Visschen, 1919: 187.

27. Guichenot, Explor. Alger., .?, Poiss., 1850: 124; a specimen sent to Paris (Dumeril, Hist. Nat. Poiss., j, 1865:

358).
»8. Fauna Iberica Peces, /, 1928: 342, 343. 29. Nobre, Fauna Marinha Portugal, t, 1935.

Fishes of the Western North Atlantic 343

write of it in more than the most general terms because of the uncertainty in many cases
as to whether published records referred to it, to longimanus or to obscurus, or to a com-
bination of these three.'" Locality records for it that can be accepted as reasonably sup-
ported by description or other information locate its center of abundance in the West In-
dian-Caribbean region, no doubt including the southern part of the Gulf of Mexico; it is
reported from Cayenne (probably), French Guiana, the Antilles" (see Study Material,
P- 337)j Cuba, Porto Rico (probably), the Bahamas, Colon (probably), from the Pan-
ama Canal, from Lake Yzabal, Guatemala; and from many localities on both coasts of
southern Florida, where It is certainly one of the more common of the larger sharks inshore
and one with which commercial fishermen and anglers are far more familiar than might
be expected from the meager scientific records. In fact, we would hazard the guess that
more of this species are caught there from wharves, etc., than of any other large shark.

To the southward it is no doubt distributed generally along the South American
coast as far as southern Brazil, being positively known from Para and Rio de Janeiro. Avail-
able information also suggests that this is the most numerous shark in shoal water around
Bermuda. In southern Florida it is resident throughout the year, but perhaps not north
of Cape Romaine, for it is not seen at Englewood (Lat. about 27° N.) in December, Janu-
ary or February although well known there during the warmer months. To the north-
ward in the Gulf of Mexico it has been described similarly as the commonest large shark
in summer on the Texas coast near Galveston ; it has been encountered off Biloxi, on the
north shore of the Gulf in July," and has even been reported in fresh water in Louisiana,
as noted above (p. 341). Also, it has been described, nominally at least, as moderately
common near Charleston, South Carolina. But seemingly it does not range much farther
north than this with any frequency, for while eleven large ones were reported as caught
near Morehead City in July and August of 1 930, only three specimens had been definitely
reported for the North Carolina coast up to 1916,^^ although a rather productive shark
fishery had been carried on there for some years previously. Moreover, some or all of
these may have been longimanus, milberti or obscurus. Evidently it occurs only as a stray
along the sector thence northward as far as New York, where the only report ostensibly
referring to it is of one New Jersey specimen." Perhaps it visits the stretch of coast east of
New York oftener, for it has been described as uncommon along Long Island,'' but there
is no way of knowing how many of the captures on which this characterization was based
were leucas, or obscurus, while the single published report of its presence farther east is
based only on a photograph of a specimen about five feet long, supposedly of this species,
taken at Woods Hole."

30. On this, see p. 361 under longimanus, and p. 38S under obscurus.

31. The Eulamia flatyodon reported froni Jamaica by Fowler (Proc. Acad. nat. Sci. Philad., 7/, 1919: 146)
appears to have been some other species, for its teeth were described as narrow and the lowers as "entire."

32. Personal communication from Stewart Springer. 33. Radcliffe, Bull. U.S. Bur. Fish., 3^, 1916: 261.

34. Fowler (Proc. biol. Soc. Wash., 55, 1920: 144, footnote 2) states that a shark earlier reported from the Dela-
ware River by him as obscurus (Rep. N. J. Mus. [1907], 190S: 12) was actually commersonii, i.e., leucas.

35. Nichols, Bull. Amer. Mus. nat. Hist., 37, 1917: 873; Copeia, 140, 1925: 21.

36. Nichols and Breder, Zoologica, N. Y., 9, 1927: 16.

344 Memoir Sears Foundation for Marine Research

Synonyms and References:^'

Carcharias {Prionodon) leucas Muller and Henle, Pkgiost., 1841 : 42 (descr., Antilles) ; Dumeril, Hist. Nat.
Poiss., /, 1865: 358 (descr., Antilles, La Trinite, S. Coast Cuba; but spec, from Algeria probably not this
species; see discuss., p. 342) ;'' Gunther, Cat. Fish. Brit. Mus., 8, 1870: 368 (discuss.).

Carcharia! {Prionodon) leucos (in part) Guichenot, Explor. Sci. Alger., j, Poiss., 1 850: 124 (ref. to Cayenne
and Porto Rico probably leucas, but not his Algerian spec; see footnote 27, p. 342.

Squalus flatyodonVoey, Memorias, 2, 1861: 336 (descr., Cuba).

Prionodon flatyodon Poey, Memorias, 2, 1861 : pi. 19, fig. 5, 6 (teeth, Cuba).

Sfua/us obtusus Poey, Memorias, 2, 1861 : 337 (descr., Cuba).

Prionodon obtusus Poey, Memorias, 2, 1 861: pi. 19, fig. 7, 8 (teeth, Cuba).

Eulamia obtusa Poey, Repert. Fisico-nat. Cuba, 2, 1868: 447, pi. 4, fig. 16 (discuss., dermal denticles, Cuba);
An. Soc. esp. Hist, nat., 5, 1876: 385; Enumerat. Pise. Cubens., 1876: 189 (Cuba).''

Carcharias flatyodon Jordan and Gilbert, Proc. U.S. nat. Mus., 5, 1882: 243 (descr., abund., Galveston,
Texas); Bull. U.S. nat. Mus., 16, 1883: 872 (descr., Cuba to Texas); Evermann and Kendall, Bull.
U.S. Fish Comm., 12, 1894: 95 (in list for Texas); Jordan and Dickerson, Proc. U.S. nat. Mus., 34,
1908: 1 1 (Tampico, Mexico) ; Gunter, Amer. Midi. Nat., 28, 1927: 316 (Atchafalaya R., Louisiana, in
fresh water); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 15 (Cuba to
Texas) ; Beebe and Tee- Van, Field Bk. Shore Fish. Bermuda, 1933: 26 (descr., abund., Bermuda).

Carcharias lamia Jordan, Proc. U.S. nat. Mus., 7, 1 884: 104 (ident. seemingly established by ill. of embryo
by Jordan and Evermann, Bull. U.S. nat. Mus. 47 [4], 1900: pi. 5, fig. 17; abund., embryos, S. Florida).

Carcharhinus {Eulamia) flatyodon Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 795 (ident. by ref., W.
Indies, S. Atlant. and Gulf coasts of U.S.).

Carcharhinus {Carcharhinus) flatyodon Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 216
(ident. by ref., Cuba to Texas) ; Bull. U.S. nat. Mus., 47 (i), 1896: 39 (descr., size, Cuba to Texas).

Carcharhinus lamia Jordan and Evermann, Bull. U.S. nat. Mus., 47 (4), 1 900: pi. 5, fig. 17 (embryo, probably
Florida); Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 48 (ident. by included refs.,
S. Florida); Smith, J. Amer. Mus. nat. Hist., 16, 1916: 346 (ident. probable from context, southeast
coast U.S.); Nichols and Murphy, Brooklyn Mus. Quart., 3, 1916: 152 (name only, but ident. prob-
able because of loc. Key West, Florida) ; not Carcharias {Prionodon) lamia, Muller and Henle, 1 841.

Eulamia longimanus Fowler, Proc. Acad. nat. Sci. Philad., 60, 1908: 65 (spec, no data, probably leucas be-
cause of meas. ; j aws, Florida) .

Carcharhinus ohscurus Fowler, Rep. N. J. Mus. (1907), 1908: 124 (spec. Delaware R. brackish water; later
stated by Fowler, Proc. biol. Soc. Wash., jj, 1920: 144, to have been "commersonii," i.e., leucas).

Carcharhinus flatyodon VerriU, Trans. Acad. Arts Sci., //, 1903: 55 (Bermuda, ident. by Garman) ; Barbour,
Bull. Mus. comp. Zool. Harv., 46, 1 905 : 1 1 1 (Bermuda, ident. by Garman) ; Garman, Mem. Harv. Mus.
comp. Zool., 36, 1913: 126, in part (synonymy, but embryos described are longimanus; see p. 354);
Starks, Stanford Univ. Publ., Univ. Ser., 1913: 4 (descr., 29-inch spec. Para, Brazil) ; Breder, Field Bk.
Mar. Fish. Atlant. Coast, 1929: 17 (general); White, Bull. Amer. Mus. nat. Hist., y4, 1934: 126 (in
Key) ; Gunter, Copeia, 1938: 69 (fresh water in Louisiana) ; Springer, Proc Fla. Acad. Sci., 5, 1939: 19
(descr., size, discuss., south. Florida); Bomkamp, Contr. biol. Lab. Cath. Univ. Amer., 44, 1942: 4
(chemistry of liver, Florida); Boos, Contr. biol. Lab. Cath. Univ. Amer., 45, 1943: lO (chemistry of
pancreas, Florida); Lunz, Bull. S. Carolina St. Planning Bd., 14, 1944: 26 (Florida).

Carcharinus commersonii (in part) Garman, Mem. Harv. Mus. comp. Zool., 36, 1 91 3: 140 {leucas Muller and
Henle incl. in synonymy, but refers otherwise to longimanus) ; Radcliffe, Bull. U.S. Bur. Fish., 34, 1916:
260, pi. 41, fig. 3, 4 (denticles, teeth, N. Carolina, but meas. are of a longimanus taken north of the

37. This species has so often been confused with others that we confine the following list to such citations as include
reasonably conclusive evidence as to specific identity.

38. Spelled "leucos."

39. Photographs of unpublished drawings by Poey of his obtusa appear clearly referable to C. leucas.

Fishes of the Western North Atlantic 34.5

Bahamas) ; Meek and Hildebrand, Field Mus. Publ. ZooL, 75 (i), 1923: 43 {Uucas MUller and Henle,
1 84 1, inch in synonymy, but specs, dcscr. from Colon are apparently limbatus) ; Fowler, Bull. Amer. Mus.
nat. Hist., -JO (1), 1936: 53 (dcscr. of Florida specimen but not the synonymy).

Carcharhinus commersonii Nichols, Bull. Amer. Mus. nat. Hist., 57, 1917: 873, pi. 91, fig. I, 3, 4 (descr.,
photo of spec. ill. here, in Fig. 60; size, no. and size of embryos, winter, S. Florida) ; Nat. Hist. N. Y., 21,
1 921: 275 (food); Bell and Nichols, Copeia, 92, 1 921: 17 (food, size, number, N. Carolina); Nichols,
Copeia, 140, 1925: 21 (dcscr., compar. with milberti and obscurus, a straggler, Long Island, N. York and
Woods Hole); Nichols and Breder, Zoologica, N. Y., p, 1927: 16 (uncommon. Long Island and near
Woods Hole, but evidence perhaps not conclusive, see p. 343); Breder, Field Bk. Mar. Fish. Atlant.
Coast, 1929: 17 (general) ; Townsend, Bull. N. Y. zool. Soc, ^4 (6), 193 i: 172 (Florida, good photo) ;
Gudger, Sci. Mon. N. Y., ^4, 1932: 417 (ident. by included ref. to Bell and Nichols, 1921, food);
White, Bull. Amer. Mus. nat. Hist., y4, 1934: 127, pi. 42E (in Key, ill. of heart valves, ident. not estab-
lished) ; Lunz, Bull. S. C. St. Planning Bd., 14, 1944: 27 (S. Carolina).

Eulamia commersonii Fowler, Proc. biol. Soc. Wash., ^5, 1920: 144 (footnote' 2 states that his earlier N.
Jersey record was commersonii); Proc. biol. Soc. Wash., ^6, 1923: 27 (name, Florida).

Carcharias melanoperus Ribeiro, Fauna brasil., Peixes, 2(1) Fasc. I, 1923: 12, pi. 2 (descr., photo of 785 mm.
spec, evidently leucas, not meUnopterus Quoy and Gaimard, 1824, of the Indo-Pacilic).

Carcharias commersonii (in part) Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 15
(leucas combined with longimanus in synon.) ; Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda,
1933: 28 (ill., apparently leucas, but text and photograph of snout appear to be longimanus').

Eulamia flatyodon Fowler, Arqu. Zool. Estado Sao Paulo, 3, 1942: 128 (listed, Brazil) ; Fish Culturist, 21
(9), 1942: 66 (listed, Cuba).

Eulamia melanoftera Fowler, Arqu. Zool. Estado Sao Paulo, 5, 1942: 128 (listed, Brazil).

Eulamia leucas Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945'.
86, fig. 27 (descr., habits, range, ill.).

References, Presumably to leucas, But Not Definitely Identifiable:

Carcharhinus flatyodon Henshall, Bull. U.S. Fish Comm., 9, 1891 : 383 (S. Florida) ; Evermann and Kendall,

Rep. U.S. Comm. Fish. (1899), 1900: 48 (Florida); Bean, Field Mus. Publ. Zool., 7 (2), 1906: 29

(Bermuda) ; Linton, Proc. U.S. nat. Mus., 33, 1907: 86; Bull. U.S. Bur. Fish., 26, 1907: 122; Engel-

hardt, Zool. Anz., 59, 1913: 648 (S. Atlantic); Linton, Proc. U.S. nat. Mus., 64 (21), 1924: 5, 48

(parasites) ; von Bonde, J. comp. Neurol., 1933: pi. 2, fig. 2, 3 (brain) ; Beebe, Proc. Wash. Acad. Sci.,

/p, 1933: 184 (Bermuda).
Carcharhinus obscurus (probably in part) Thome, Copeia, 35, 1916: 69; Nichols, Copeia, 35, 1916: 72-73;

Thome, Bull. N. Y. zool. Soc, 31, 1928: 114 {leucas likely represented among obscurus reported from

Long Island, N. York).
Carcharinus lamia W\c)\oh,Co^t\3, 53, 191 8: 13 (abund., Florida).
Carcharhinus commersonii Linton, Proc. U.S. nat. Mus., 64 (21), 1924: 6, 7, 12, 33, 38, 49, 54, 84, 87

(parasites) ; Burton, Sci. Mon. N. Y., 40, 1935: 283 (Charleston, S. Carolina).
Carcharias commersonii Marini, Physis B. Aires, p, 1929: 452 (Argent.) ; Bere, Amer. Midi. Nat., 17, 1936:

593 (parasites).
Carcharias carcharias (in part) Jordan, Manual Vert. Anim. NE. U.S., 1929: 10 (account perhaps partly

based on leucas).

Eastern Atlantic References, Nominally to leucas, But Probably Not This Species (see footnote 37, p. 344):
Carcharias (Prionodon) leucas Guichenot, Explor. Alger., 5, Poiss., 1 850: 124 (spec, from Algeria) ; Dumeril,

Hist. Nat. Poiss., 1, 1865: 358 (ref. to Algerian spec, from Guichenot) ; Doderlein, Man. Ittiol. Medit.,

2, 1 881: 44 (credited to Medit., on basis of Algerian spec, reported as leucos by Guichenot, 1850);

Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: ig; Faune Senegambie, Poiss., /, 1883-1885: 19

(name only, C. Verde, Senegambia).
\ Carcharias leucos Cams, Prod. Fauna Medit., 2, 1 889-1 893: 513 (name, Algiers).

346 Memoir Sears Foundation for Marine Research

Red Sea Reference, Probably Not leucas:

Carcharkinus commersonii Gudger, Bull. Amer. Mus. nat. Hist., 75, 1937: 264 (photos of teeth, prob. from
Gulf of Aden).

Not Carcharias leucas Bennett, Proc. zool. Soc. Lond., 1859: 223 (N. Zealand).*"

Not Carcharinus flatyodon Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: pi. 3, fig. 1-4 (this is a
milberti, mislabelled).

Not Eulamia flatyodon Fowler, Proc. Acad. nat. Sci. Philad., yi, 1919: 146 (a Jamaica spec, not leucas be-
cause teeth narrow and the lowers entire ; perhaps macuUpnnis) .

Not Carcharkinus commersonii Rey, Fauna Iberica Feces, j, 1928: 342 (this is obscurus, see p. 342).

Carcharkinus limbatus (Miiller and Henle), 1 841

Small Black-tipped Shark

Figures 62, 63

Study Material. Female, 990 mm. (about 3 feet 3 inches) long, from Florida (Amer.
Mus. Nat. Hist.) ; also 14 specimens, 460 to 621 mm. long, from Galveston, Texasj" one,
585 mm., from Pine Island, Floridaj 3 embryos, 351 to 398 mm., from Captiva Key,
Florida (Harv. Mus. Qjmp, Zool.).

Distinctive Characters. Among smooth-backed members of the genus in the Atlantic,
limbatus falls with maculifinnis (p. 364) in the conspicuously black-tipped fins. But it is
easily distinguishable from the latter by its relatively larger eyes, its shorter gill openings
and upper labial furrow, and by the more obtuse free rear tip of its second dorsal j also, the
edges of its lower teeth, as well as those of the uppers, are regularly, though very finely,
serrate, and its first dorsal usually originates somewhat farther forward relative to the
pectorals.

Description. Proportional dimensions in per cent of total length, Male, 621 mm.,
from Galveston, Texas (Harv. Mus. Comp. Zool., No. 35842). Female, 990 mm., from
Florida (Amer. Mus. Nat. Hist.).

Trunk at origin of pectoral: hrezdth 11.4, 10.4; height 12.6, 12.3.

Snout length in front of: outer nostrils 3.5, 3.3; mouth 7.9, 7.9.

Eye: horizontal diameter 1.5, 1.5.

Mouth: breadth 9.0, 8.95 height 4.8, 4.7.

Nostrils: distance between inner ends 5.4, 5.2.

Labial furrow length: upper 0.8, 0.8.

Gill opening lengths: ist 4.3, 3.65 2nd 4.5, 3.95 3rd 4.7, 4.1 j 4th 4.2, 3.8} 5th

3.6, 2.9.

40. As leucas was preoccupied, the name brachyurus was substituted by Giinther (Cat. Fish. Brit. Mus., 8, 1870: 369)
for this Indo-Pacific species, which differs from the Atlantic leucas in a more pointed snout, and in that its first
dorsal originates considerably farther back. It is classed as a synonym of lamia, i.e., longimanus, by Fowler
(Bull. U.S. nat. Mus., 100 [/j], 1941: 171)) but differs equally sharply from the latter in its angular first
dorsal, in its much shorter pectoral, and in that the tip of its anal is far in advance of the origin of its caudal.
For photograph, see Whitley (Fish. Aust., i, 1941: 102, fig. 97).

4.1. Contributed by J. L. Baughman.

Fishes of the Western North Atlantic

347

First dorsal fin: vtT{\C2.\h.G.\g\\t 10.6, 11.3; length of base 11.3, ir.6.

Second dorsal fin: vertical height 3.4, 3.2; length of base 4.5, 5.0.

Anal fin: vertical height 3.7, 3.7; length of base 5.3, 5.4.

Caudal fin: upper margin 28.3, 27.6; lower anterior margin 12.7, 12.5.

Pectoral fin: outer margin 17.7, i7-7; inner margin 5.8, 5.4; distal margin 13.4,

14.8.

Distance from snout to: ist dorsal 30.9, 31.6; 2nd dorsal 61. i, 62.05 upper

caudal 71.7, 72.4; pectoral 22.1, 25.4; pelvics 48.2, 49-0} anal 59.2, 61. i.

Interspace between: ist and 2nd dorsals 20.5, 20.9; 2nd dorsal and caudal 6.S,

6.3 } anal and caudal 5.9, 6. i .

Distance from origin to origin of: pectoral and pelvics 26.1, 25.05 pelvics and

anal 11.6, 11.9.

Trunk moderately slender, its dorsal profile only slightly arched, its height at origin
of 1st dorsal about V^-Vg its length to origin of caudal. No mid-dorsal ridge. Upper pre-
caudal pit rounded, the lower a subrectangular furrow and less strongly marked than
upper. Caudal peduncle about i Vs times as thick as deep. Dermal denticles closely spaced
and usually overlapping, the skin visible only here and there between them, the smaller

Figure 62. Carcharhinus limbatus, female, about 990 mm. long, from southern Florida (Amer. Mus. Nat.
Hist.). A, B Caudal peduncle, from above and below respectively, to show precaudal pits. C Left-hand
upper and lower teeth, about 1.8 x. D Dermal denticles, about 30 x. E Apical view of dermal denticle,
about 60 X.

348

Memoir Sears Foundation for Marine Research

ones with 3 to 5 low ridges and the larger with 5 or even 7, the marginal teeth relatively
low and broad, mostly worn down in specimens seen, except in the cases of newly formed
denticles} pedicels moderately slender, on large basal plates.

Head about V4 of total length. Snout moderately thick, its lateral outlines con-
verging gradually from opposite front of mouth to opposite ends of nostrils and then

Figure 63. Carcharhinus limbatus, pictured in Fig. 62. A Lower view of head. B Pelvic fins. C Fourth
upper tooth. D Tenth upper tooth. E Fourth lower tooth. F Ninth lower tooth. C-F, about 5.4 x. G Right-
hand nostril, about 3 x.

more abruptly j its tip rounded (relatively somewhat shorter and broader anteriorly
than in maculipinnis) , its length in front of a line connecting outer ends of nostrils nearly
% (63-65%) as great as distance between inner ends of latter, and its length in front of
mouth a little more than i V2 times as great as that, or about Vs as great as length of head.
Eye approximately circular, its vertical diameter about 0.9 the horizontal diameter,
the latter about Vs ( 1 7 to 20% ) as long as snout in front of mouth, or a little less in large
specimens. Gill openings relatively somewhat shorter than in maculipinnis, the ist only
about % as long as distance between inner ends of nostrils and 1.9 to 2.8 times as long as
horizontal diameter of eye, the 3rd a little the longest, the 5th oblique and a little more than
% as long as ist, the 4th over origin of pectoral. Nostril strongly oblique, its inner end

Fishes of the Western North Atlantic 349

closer to mouth than to tip of snout by a distance about equal to diameter of eye, the
anterior margin slightly expanded near the inner end as an inconspicuous, broadly-tri-
angular lobe with rounded tip. Mouth ovate, moderately arched, its breadth 1.6 to 2 times
its height. Upper labial furrow only about half as long as nostril (thus shorter, rela-
tively, than in C. maculifinnh)-, at an angle of about 45° with the jaw.

Teeth ^^[|~?°2i-n°oJ5 > uppers nearly symmetrical with narrow triangular cusps on
broad bases, with edges finely serrate on cusps and more coarsely so on bases, those toward
center of mouth erect, their margins nearly straight, but those along sides slightly oblique;
lowers with slender erect cusps, recurved slightly forward near tips, and broad bases, the
cusps very finely serrate, but bases less clearly so; i to 3 minute teeth at symphysis in
upper jaw, and i or 2 in lower; outermost 2 or 3 teeth in each jaw also very small.

Origin of ist dorsal about over midpoint of inner margin of pectoral, its position
somewhat variable, its anterior margin moderately convex in small specimens but only
very slightly so in larger, its apex very narrowly rounded, the posterior margin nearly
straight toward apex but deeply concave toward base, its free rear tip noticeably slender
but only about Vs as long as base, its vertical height about equal to distance from corner
of mouth to 5th gill slit or slightly more than Y2 (62 to 64%) the length of pectoral, the
midpoint of its base only about V2 as far from axil of pectoral as from origin of pelvics.
Second dorsal between V3 and Y2 ( 3 8 to 4 1 % ) as long at base as i st, its origin over or ver}'
slightly behind origin of anal, its anterior margin nearly straight, its rear margin slightly
and evenly concave, its apex narrowly rounded or subacute, its free rear tip relatively
broad and only about as long as the base or slightly shorter. Caudal between V4 and Vs
(26 to 29%) of total length, the terminal sector between l^ and ^7-5 the length of the
upper margin, narrow-triangular with slightly sinuous lower contour and narrowly
rounded tip, the lower lobe (expanded lower anterior corner) a little less than V2 as long
as upper, its anterior margin moderately convex, its tip narrowly rounded or subacute, the
re-entrant corner (included by the two lobes) subrectangular and only slightly rounded.
Anal about as long at base as 2nd dorsal or a little longer, but its anterior margin some-
what more convex, and its rear margin much more deeply concave, its apex subacute, its
free rear tip slightly but definitely shorter than its base. Distance from origin of anal to tip
of pelvics a little shorter than base of anal. Pelvics with weakly concave distal margins,
their anterior margins slightly convex in small specimens but nearly straight in larger
ones, with bases about Yi as long as base of ist dorsal. Pectoral a little less than % (70
to 72%) as long as head, about V2 as broad as long but with base only about % as long as
outer margin, distal margin nearly straight toward tip but deeply concave toward base,
apex narrowly rounded and inner corner only a little more broadly so.

Color. Dark gray, dusky bronze, or ashy blue above; pure white or yellowish white
below, with a more or less pronounced band of the dark upper tint extending rearward
along each side to about over the origin of the pelvic, including above it a forward exten-
sion of the pale or white of the lower side, which narrows forward. This dark band varies

350 Memoir Sears Foundation for Marine Research

widely in strength from specimen to specimen and gradually fades after preservation in
formalin or alcohol. The pectorals are conspicuously tipped with black; the dorsals, anal
and lower lobe of caudal are also black-tipped in young, but fade more or less with growth.

Size. This Shark may mature at a length no greater than about four to five feet;
adults perhaps average 5^/2 to 6V2 feet, and few grow longer than seven or possibly eight
feet in length. It is, however, relatively much larger at birth than many other sharks.
Embryos 21 to 23 inches (nearly ready for birth) have been taken from females only four
to five feet long, and embryos 23 to 26 inches (580 to 660 mm.) from a Pacific speci-
men of about 6 feet 9 inches (2,070 mm.). A specimen of about 28 inches (705 mm.)
weighed about 4.2 pounds (1.9 kilo); one of about 47.7 inches (1,210 mm.) about 19.6
pounds (8.9 kilo); one of about 54.2 inches (1,375 mm.) about 32 pounds (14.5 kilo);
one of about 61.3 inches about 42.2 pounds (19. i kilo); and one of about 65.8 inches
about 68.3 pounds (31 kilo)."

Developmental Stages. Embryos of this species have been reported repeatedly, but
we find no statement as to whether it is viviparous or ovoviviparous, although the large
size which the embryos reach before birth suggests the former; nor do the specimens we
have studied (see Study Material, p. 346) clarify the matter. Three to nine embryos have
been recorded, but the usual number is four to six, with the sexes in approximately equal
numbers.

Habits. This is a very active, swift-swimming shark, often seen in schools at the
surface, frequently leaping clear of the water, sometimes revolving as much as three
times around the horizontal axis in the air before falling back into the sea.*' It is en-
countered indifferently near shore and out at sea. It feeds on fishes, its teeth suggesting
chiefly the smaller species, and probably on squid; menhaden (Brevoortia) and butterfish
(Poronotus triacanthus) have been found in the stomachs of North Carolina specimens,
and 24 California sardines (Sardinops caerulea) in an eastern Pacific example. Like sun-
dry other sharks, it devours sting-rays on occasion and their spines have been found
imbedded in its jaws. On the other hand, smallish Black-tipped Sharks have been found
in the stomachs of larger sharks of other species.

Embryos in gravid females taken oflF Biloxi, Mississippi, in midsummer were small;**
but larger embryos nearly ready for birth were taken around southern Florida in April,
suggesting that the period of gestation is nearly a year, and that the young are born chiefly
in late spring. Other than the foregoing, nothing is recorded of its habits, notwithstanding
that it has been known to science for more than a century.

Relation to Man. Such of the larger specimens as are taken in shark fisheries in
Florida or elsewhere are put to the same uses as other species. Thanks to the readiness
with which it takes a hook baited with fish (stiU or trolled) many are so caught; in fact,
it is often caught by anglers trolling for tarpon in Florida waters. But reports differ as to

42. Florida specimens measured and weighed by Stewart Springer.

43. Recent observation off Biloxi, Mississippi, by Stewart Springer.

44. About 125 mm. J personal communication from Stewart Springer.

Fishes of the Western North Atlantic 351

its game qualities; we have caught none on light tackle. Some anglers describe it as
putting up a fast fight when hooked and often jumping clear of the water, but according
to others it is more stubborn and "mulish" than active in its resistance.

Range. Pelagic in tropical and subtropical seas; southern Brazil to North Carolina
and occasionally to New York and southern New England in the western Atlantic; Ma-
deira, Cape Verde Islands and tropical West Africa (Dakar, and Kribi in Cameroon) in
the eastern Atlantic; also eastern tropical Pacific, from Lower California to Peru.*" A
shark (or sharks) is also reported under this same name from Cochin China, India, Red
Sea, Seychelles, Madagascar and Natal. But its actual relationship to limbatus of the
eastern Pacific and Atlantic cannot be determined until specimens from the different
ocean areas have been compared critically.

Occurrence in the Western Atlantic. This can be outlined only within broad limits,
partly because of the uncertainty in some cases as to whether published records actually
refer to Ihnbatus or to maculipnnu, and partly because reliable records for it are very
irregularly distributed. It is certainly one of the commoner, if not the commonest, pelagic
shark around the Bahamas and southern Florida; also along the coasts of Mississippi and
Louisiana, where many are caught by shrimp fishermen, and of Texas" at least in the
warm season. In all probability it is equally widespread and locally common through-
out the West Indian-Caribbean region in general and in the southern part of the Gulf
of Mexico; but published records of it there are confined to Haiti, Porto Rico (where
it is said to be one of the commoner sharks), Turks Island, Martinique, Antilles in
general and Surinam. Southward It Is common along the coast of Brazil at least as far as
Rio de Janeiro (reported also from Bahia and from Ilha de Victoria) ; but It has not been
reported farther south. No doubt it is present In the truly tropical belt throughout the
year, and it is common around southern Florida throughout spring, summer and autumn."
During the warm months many visit the coast of South Carolina, and a few are taken in
some summers along the southern half of North Carolina. To the north of Cape Hatteras,
however. It occurs on the coast only as a stray, there being only about six reliable reports of
it for the vicinity of New York and Long Island. But at least twenty small ones were taken
In pound nets on the eastern shore of Buzzards Bay, near Woods Hole, during the summer
of 1878, and one other in the summer of 19 16, which shows that it reaches the southern
New England coast in unusual numbers at rare intervals. Nor Is It unusual to see Black-
tipped Sharks In the warm oceanic waters off this sector of the continental shelf in summer,
drifting north in the Gulf Stream, probably never to return to their tropical home.

45. There appears to be nothing in the accounts of aethlorus Jordan and Gilbert, 1882, from the west coast of
Mexico and Lower California to separate it from limbatus as Garman (Mem. Harv. Mus. comp. Zool., 56, 1913:
127) has already concluded; also Meek and Hildebrand (Field Mus. Publ. Zool., 75 [i], 19^3: 43). after
examining the type specimen. Neither has our own examination of the type specimen of natator Meek and Hilde-
brand, 1923, from Panama revealed significant differences from the Atlantic specimens of limbatus (listed
above) whether in shape of snout, shape and relative positions of fins, teeth, or color.

46. Personal communication from J. L. Baughman; see Study Material, p. 346.

47. Not reported there in December, January or February.

352 Memoir Sears Foundation for Marine Research

Synonyms and References:

Atlantic, Eastern Pacific:'"

Carcharias {Prionodon) limbatm Muller and Henle, Plagiost., 1 841: 49, pi. 19, fig. 9 (descr., no loc.) ;
Dumeril, Hist. Nat. Poiss., 1865: 375 (descr., Martinique [type], Bahia) ; Gunther, Cat. Fish. Brit.
Mus., 8, 1870: 373 (descr., C. Verdes, Seychelles, Indian Oc, Guatemala) ; Rochebrune, Act. Soc. linn.
Bordeaux, (4) (5, 1882: 42; Faune Senegambie, Poiss., 1883-1885: 19 (C. Verde); Metzelaar, Trop.
Atlant. Visschen, 1 91 9: 188 (nominal, both sides of Atlantic).

Prionodon cucuri Castelnau, Anim. Nouv. Rares Amer. Sud., 1855: 99'" (Bahia, Brazil).

Lamiopis limialus G'\\\, Ann. N. Y. Lye, 7, 1862: 410 (name only).

Carcharias {Prionodon) miil / eri Ste'indichner, S. B. Akad. Wiss. Wien, 56, 1867: 356 (Antilles).

Carcharias maculifinna Gunther, Trans, zool. Soc. Lond., 6, 1 868 : 490 (teeth, meas., Chiapam, Pacific Guate-
mala) ; not Isogomfhodon maculifinnis Poey, 1865 ; see p. 368.

Platyfodon ferezii Poey, An. Soc. esp. Hist, nat., 5, 1876: 390, pi. 14, fig. 2, 3; Enumerat. Plsc. Cubens.,
1876: 194, pi. 9, fig. 2, 3 (descr., color, size, teeth, Cuba).'"

Carcharias aethlorus Jordan and Gilbert, Proc. U.S. nat. Mus., 5, 1882: 104 (descr., Mazatlan, Mexico) ; Jor-
dan and Gilbert, Bull. U.S. Fish Comm., 2, 1882: 105, 109 (Mazatlan, Panama); Gilbert and Starks,
Mem. Calif. Acad. Sci., 4, 1904: 9, 207 (descr., meas., Panama, Gulf of California) ; Jordan, Evermann
and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 17 (Mazatlan to Panama).

Isogomfhodon limbatus ]ordL3in and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 23 (tropical seas and Woods Hole
region) ; Nelson, Rep. St. Geol. N. J., 2 (2), 1890: 661 (in N. Jersey list) ; Springer, Proc. Fla. Acad.
Sci., 5, 1939: 26 (descr., habits, Florida) ; Norris, Plagiost. Hypophysis, 1941 : 20, 28, pi. 3, fig. 10, loa
(brain); Lunz, Bull. S. C. St. Planning Bd., 14, 1944: 27 (S. Carolina, Florida); Gunter, Publ. Inst.
Mar. Sci. Univ. Texas, i, 1945: 20 (temp., breeding, Texas).

Isogomfhodon maculifinnis Goode, Fish. Fish. Industr. U.S., i, l884:j673 (Woods Hole, Massachusetts) ; not
Isogomfhodon maculifinnis Poey, 1865 ; see p. 368.

Carcharhinus aethlorus jorian, Proc. U.S. nat. Mus., 8, 1885: 363 (refs.) ; Rep. U.S. Comm. Fish. (1885),
1887: 795 (east. trop. Pacif.) ; Proc. Calif. Acad. Sci., (2) 5, 1895: 383 (discuss.) ; Garman, Bull. Mus.
comp. Zool. Harv., 46, 1906: 229 (Panama).

Eulamia limbata Henshall, Bull. U.S. Fish Comm., 9, 1 891: 372 (color, west coast Florida); Fowler, Bull.
Amer. Mus. nat. Hist., jo (l), 1936: 48 (descr.); Arqu. Zool. Estado Sao Paulo, 3, 1942: 128; Fish
Culturist, 21 (9), 1942: 66 (listed, Cuba); Feces Peru, Mus. Hist. Nat. Javier Prado, 1945: 12 (listed,
Peru); Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945-
88, fig. 28 (descr., habits, range, ill.).

Carcharhinus {Isogomfhodon) aethlorus Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 216
(Mazatlan to Panama) ; Bull. U.S. nat. Mus., 47 (i), 1896: 40 (descr., Mazatlan to Panama).

Carcharhinus {Isogomfhodon") limbatus Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 216
(distrib.) ; Bull. U.S. nat. Mus., 47 (l), 1896: 40 (descr., distrib.).

Carcharhinus {Platyfodoti) ferezi Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 216 (name,
Cuba) ; Bull. U.S. nat. Mus., 47 (l), 1896: 36 (descr., Cuba).

Carcharhinus limbatus Smith, Bull. U.S. Fish Comm., 77, 1898: 88 (Woods Hole region); Evermann and
Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 48 (Florida) ; Pellegrin, Bull. Mus. Hist. nat. Paris, 7,

48. References for "limbatus" for the western Pacific and Indian Oceans are omitted because of uncertainty whether
the shark (or sharks) to which they refer are identical with limbatus of the Atlantic and eastern Pacific.

49. The name cucuri dates back to Marcgrave (in Piso de Medicina brasiliense, et G. Marcgravius Hist, rerum
natur. Brasiliae, 1648: 164). But Castelnau's revival of it for the shark in question to replace limbatus, Muller
and Henle, was not a fortunate one, even apart from nomenclatural grounds, for Marcgrave's original account
"pinnae omnes cinereae, exceptis parvulibus in inferiori corpore, quae albae" does not suggest this species.

50. Perezii was classed by Garman (Mem. Harv. Mus. comp. Zool., 36, 1913 : 138) as a synonym of remotus. Actu-
ally, however, its color as well as the original illustrations of its teeth more nearly resemble limbatus, its pec-
torals being described as with black tips and its sides with a dark band. A photograph of Poey's unpublished
drawing of an adult female shows nothing to separate it from limbatus.

Fishes of the Western North Atlantic 353

1901: 161 (Gulf of California); Evermann and Marsh, Bull. U.S. Fish Comm., 20, 1902: 62 (dcscr.,
Porto Rico) ; Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 5 (Woods Hole region) ; Garman,
Mem. Harv. Mus. comp. Zool., $6, 1 91 3: 127 (dcscr., Rio de Janeiro) ; Gudgcr, J. Elisha Mitchell Sci.
Soc, 2*, 191 3: 158 (N. C.irolina); Nichols, Abstr. Proc. Linn. Soc. N. Y., 20-23, 1913: 91 (N.York);
Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., jr (2), 1913: 736 (Woods Hole region); Coles,
Proc. biol. Soc. Wash., 28, 1915: 90 (N. Carolina); Nichols and Murphy, Brooklyn Mus. Sci. Bull., 3
(i), 1916: 17 (trop. seas, Atlantic coast to Buzzards Bay, Massachusetts); Nichols, Copeia, 35, 1916: 73
(name) ; Thorne, Copeia, 35, 1916: 69 (Long Island, N. York); Helmuth, Copeia, 36, 1916: 80 (Long
Island, N. York); Radcliffe, Bull. U.S. Bur. Fish., 34, 1916: 255, pi. 40, fig. 3, 4 (teeth, denticles,
food, N. Carolina) ; Smith, J. Amer. Mus. nat. Hist., 16, 1916: 348 (Woods Hole spec.) ; Nichols, Bull.
Amer. Mus. nat. Hist., 57, 1917: 874 (size, embryos, Florida) ; Nichols and Mowbr.iy, Copeia, 48, 1917:
78 (Porto Rico) ; Nichols, Copei.i, 53, 1 91 8: 13 (Long Island, N. York) ; Bell and Nichols, Copeia, 92,
1 921: 18 (N. Carolina); Nichols, Bull. Amer. Mus. nat. Hist., 44, 1 921: 22 (Turks I.); Nichols and
Murphy, BulL Amer. Mus. nat. Hist., 46, 1922: 504 (jaw, Peru); Meek and Hildebrand, Field Mus.
Publ. Zool., 15, 1923: 41 (descr., Pacif. Panama) ; Linton, Proc. U.S. nat. Mus., 64 (21), 1924: 12, 33,
46, 49, 87 (parasites. Woods Hole); Breder, Bull. N. Y. zool. Soc, 28, 1925: 187 (C. Hatteras, em-
bryos) ; Nichols and Breder, Zoologica, N. Y., 9, 1927: 17 (general) ; Thorne, Bull. N. Y. zool. Soc, 5/,
1928: 114 (Great South Bay, N. York) ; Beebe and Tee-Van, Zoologica, N. Y., jo, 1928: 28 (Haiti) ;
Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 17 (general); Nichols, Sci. Surv. Porto Rico, N. Y.
Acad. Sci., 10 (2), 1929: 183 (Porto Rico); Gowanloch, Bull. La. Conserv. Dep., 21, 1932: 89 (Louisi-
ana) ; Budker, BuU. Mus. Hist. nat. Paris, (2) 7, 1935: 183 (Dakar, W. Afr.) ; Burton, Sci. Mon. N. Y.,
40, 1935: 283 (Charleston, S. Carolina); Cadenat, Rev. des Trav. PSchcs Marit., 10 (4), 1937: 43°
(Dakar) ; White, Bull. Amer. Mus. nat. Hist., 7./, 1937: 127, pi. 6, fig. g, pi. 13, fig. a (in Key, denticles,
teeth) ; Breder, Bull. N. Y. Zool. Soc, 41, 1938: 28 (N. York Harbor) ; Hildebrand, Copeia, 1941: 221
(N. Carolina).

Carcharias limbatus Jordan and Gilbert, Proc U.S. nat. Mus., 5, 1883: 245 (Gulf of Mexico); Schreiner
and Ribeiro, Arch. Mus. nac Rio de J., 12, 1 903: 79 (Rio de Janeiro) ; Ribeiro, Arch. Mus. nac Rio de
J., 14, 1907: 153, 200, pi. 3 (descr., photo, W. Atl. refs.) ; Rev. Mus. paul., 10, 1918: 707 (Ilha de
Victoria) ; Fauna brasil. Peixes, 2 (2) Fasc i, 1923: 10 (same as Ribeiro, 1907) ; Jordan, Manual Vert.
Anim. NE. U.S., 1929: 10 (general); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2,
1930: 16 (distrib.) ; Gowanloch, Bull. La. Conserv. Dep., 25, 1933: 83, 220, 236 (Louisiana); Bere,
Amer. Midi. Nat., 17, 1936: 589, 593, 604 (Florida, ident.?); Gunter, La. Conserv. Rev., 5 (4), 1936:
45 (Louisiana).

Carcharhinus natator Meek and Hildebrand, Field Mus. Publ. Zool., 15 (l), 1923: 41, pi. I, fig. I (descr.,
ill., Panama City, Panama).

Carcharias natator Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 17 (Panama
City).

Prionodon limbatus Borri, Mem. Soc. tosc Sci. nat., 44, 1934: 89 (no loc).

Eulamia aethlorus Beebe and Tee-Van, Zoologica, N. Y., 26, 1941: 106 (descr., size, weight, parasites, food,
embryos. Pacific coasts, Mexico, Guatemala, Panama).

Probable References:

Carcharias {Prionodon) melanofterus Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 42 (ident. probable

because of fins descr. as conspicuously black-tipped; trop. W. Africa); not C. melanofterus Quoy and

Gaimard, 1824.
Carcharhinus commersonii Meek and Hildebrand, Field Mus. Publ. Zool., r5 (0. '9^3: 43 (synonymy, refs.

to longimanus and leucas, but Colon specs, apparently limbatus because of meas. and color).

Doubtful Reference:

Carcharias microp Lowe, Proc. zool. Soc. Lond., 1 840: 38; Trans, zool. Soc. Lond., 3, 1843: 18 (not idcntif.
by brief notice, Madeira).

354 Memoir Sears Foundation for Marine Research

Carcharhinus longimaivus (Poey), 1861"

White-tipped Shark

Figures 64, 6$

Study Material. Adult male, about 6 feet 9 inches long, taken ofiF Guantanamo,
Cuba, in April 1939; adult female of about the same length, taken off Santa Clara
Province on the north coast of Cuba in May 1939, the latter specimen with 6 female em-
bryos, 550 to 580 mm. long and nearly ready for birth 5" jaws from Guadeloupe and
Santa Cruz, West Indies, specimens} jaws from six specimens, with pieces of skin from
two of these, up to 6V2 to 7 feet long by calculation, caught from the research ship "At-
lantis" in the Caribbean (precise localities not recorded) in February 1934; also the fol-
lowing embryos: a male, said to have been one of 9 embryos, from north of the Bahamas
in Lat. 28° 30' N., Long. 77° 35' W. (Harv. Mus. Comp. ZooL, No. 35249) ; two males,
395 and 460 mm., from Guadeloupe, West Indies (Harv. Mus. Comp. ZooL, No. 756"') ;
female of about 580 mm., from off Havana, Cuba (Harv. Mus. Comp. ZooL, No. 33439) }
four others, female and male, about 515 to 525 mm. long, from north of the Bahamas
(U.S. Nat. Mus., No. 1 18548, 11 8549)} also photographs of unpublished drawings by
Poey.

Distinctive Characters. C. longimanus is set apart from all other Atlantic members
of the genus by the very broadly rounded apex of its first dorsal fin, the convexity of the
posterior outline of the lower caudal lobe, its very short snout in front of the nostrils
{leucas alone resembles it in this respect) and by the fact that the rear tip of the anal reaches
nearly to the lower precaudal pit (see also comparison with C. leucas, p. 338).

Description. Proportional dimensions in per cent of total length. Female, 2,070

51. In early shark literature this species was almost inextricably entangled with the man-eater now universally
known as Carcharodon, while more recently it has been confused frequently with Carcharhinus leucas. Thus it
appears to have been combined with Carcharodon by Linnaeus, 1758, under the name Squalus carcharias, and
clearly was so combined by Risso (Ichthyol. Nice, 1810: 25). But this specific name is not available for it,
because S. carcharias Linnaeus is now universally accepted as the type of Carcharodon. In 1826 Risso (Hist. Nat.
Europ. Merid., j: 119) redescribed it as Carcharias lamia, this time omitting: such of the features as obviously
referred to Carcharodon in his earlier account. It was as Carcharias (Prionodon) lamia that Muller and
Henle (Plagiost., 1841 : 37, pi. 12) published what has continued to be the best account and illustration of it up
to the present time. But the name Carcharias lamia had been used previously by Rafinesque (Indice Ittiol.
Sicil., 18 10: 44) as a substitute for Squalus carc/mrias Linnaeus, the White Shark; hence it is a synonym of the
latter, according to the rules of zoological nomenclature as generally accepted, and cannot be used for any other
shark. To replace it for the species here under discussion we must therefore turn to the next oldest name under
which the latter has been cited, «.«., to longimanus Poey, 1861. Fortunately there can be no doubt as to the iden-
tity of the shark so named by him; his account and the photographs of his unpublished drawings specify the
short, broad snout, the rounded first dorsal, the very long pectorals, the close proximity of the tip of the anal to
the origin of the caudal, and the teeth of the specimen pictured here reproduce almost exactly the one figured
by him.

52. Measurements and photographs were taken of the latter; the jaws and fins, with the embryos, are in the Harvard
Museum of Comparative Zoology.

52a. Basis of Carman's (191 3) account of C. flatyodon.

Fishes of the Western North Atlantic 355

mm., from Cuba (Harv. Mus. Comp. Zool., No. 35516). Male, 2,075 mm., from Cuba
(Harv. Mus. Comp. Zool., No. 35517).

Snout length in front of: outer nostrils 2.5, 2.5; mouth 6.$, 6.1.

Eye: horizontal diameter i.i, 1.2.

Mouth: breadth 9.8, 9.3; height 4.6, 5.0.

Nostrils: distance between inner ends 5.8, 5.8.

Gill opening lengths: 1st 4.0, 3.25 5th 3.5, 2.8.

First dorsal fin: vertical height 12.2, 13.2; length of base i i.o, 10.4.

Second dorsal fin: vertical height 4.3, 4.0; length of base 3.7, 3.7.

Anal fin: vertical height 5.4, S-5; length of base 3.7, 4.0.

CaW^/ ^«.- upper margin 28.2, 28.3 j lower anterior margin 13.5, 14.1.

Pectoral fin: outer margin 23.4, 22.0j inner margin 5.3, 5.25 distal margin 19.9,

19.6.

Distance from snout to: ist dorsal 30.8, 31.55 2nd dorsal 63.0, 62.5; upper caudal

71.8,71.7} pectoral 22.8,23.3; pelvics 54.7,49.6.

Interspace between: ist and 2nd dorsals 2 1.4, 20.5 j base of 2nd dorsal and caudal

5.8, 6.O3 anal and caudal 4.1, 4.3.

Distance from origin to origin of: pelvics to caudal pit 21.2, 21.4.

Figure 64. A, Carcharhinus longimanus, adult female, about six feet nine inches long, off north coast of
Cuba, from photographs, measurements and preserved fins (Harv. Mus. Comp. Zool., No. 35516). B Em-
bryo about S7S mm. long taken from the above. C Anterior part of head of this same embryo.

356

Memoir Sears Foundation for Marine Research

Proportional dimensions in per cent of total length. Female embryo, 580 mm.
(Harv. Mus. Comp. Zool., No. 35518, taken from No. 35^16).
Trunk at origin of pectoral: breadth 1 1.7; height 12.9.
Snout length in front of: outer nostrils 2.7; mouth 7.3.
Eye: horizontal diameter 2.2.
Mouth: breadth 8.6; height 5.9.
Nostrils: distance between inner ends 5.9.

Gill opening lengths: ist 2.9; 2nd 3.4; 3rd 3.5; 4th 3.4; 5th 2.6.
First dorsal fin: vertical height 12.O; length of base 12.5.

\sr

Figure 65. Carefiarhinus longimanus, adult illustrated in Fig. 64. A Upper and
lower teeth, left-hand side, about natural size. B Sixth upper tooth, about 2 x. C Sec-
ond lower tooth, about 2 x. D Eighth lower tooth, about z x. E Dermal denticles,
about 35 X. F Apical view of dermal denticle.

Fishes of the Western North Atlantic 357

Second dorsal fin: vertical height 3. i ; length of base 4.2.
Anal fin: vertical height 3.65 length of base 4.6.
Caudal fin: upper margin 30.3; lower anterior margin 12.4.
Pectoral fin: outer margin 23.85 inner margin 6.4; distal margin 17.6.
Distance from snout to: ist dorsal 33.O; 2nd dorsal 61.O; upper caudal 69.7 j pec-
toral 22.O; pelvics 49.2; anal 62.0.

Interspace between: ist and 2nd dorsals 17.25 2nd dorsal and caudal 5.3; base of
anal and caudal 3.6.
Distance from origin to origin of: pectoral and pelvics 26.75 pelvics and anal 12.4.

Trunk moderately stout, its height at origin of ist dorsal a little more than % its
length to origin of caudal. Midline of back with a low dermal ridge occupying the middle
% of the space between ist and 2nd dorsals in embryos 5°' whether or not this ridge persists
throughout life, or how regularly, is not yet known.'* Caudal peduncle moderately flat-
tened, upper precaudal pit well marked, subrectangular, the lower only weakly defined.
Dermal denticles so nearly flat that skin is smooth to the touch, overlapping only very
little, the skin exposed here and there, their blades broader than long, usually with 5
(occasionally 6 or 7) low, sharp-edged ridges, the posterior margins very broadly ovate,
usually with 5 very short teeth (the median only a little the largest), or in some cases
merely sinuous in the corresponding radii, depending on the position on the body and on
the degree of wear; pedicels rather slender.

Head about Y^ of total length, its dorsal profile moderately and evenly convex,
about % as wide at outer ends of nostrils as at eyes. Snout thick-tipped, very broadly
rounded in front and noticeably short, its length in front of a line connecting outer ends
of nostrils a little less than '^/o as great as distance between inner ends of latter," its length
in front of mouth between V3 and ^^ as long as head to origin of pectoral (about 26 to
28%) in adult, but relatively somewhat longer in embryo. Eye approximately circular, its
anterior edge about opposite front of mouth, noticeably small and increasingly so with
growth, its horizontal width decreasing from about 40% as great as distance between
inner ends of nostrils in embryos to only about 20% as great as that in large speci-
mens. Gill openings evenly spaced, the ist 2 to 3 times as long as diameter of eye in
adult, but only a little longer than eye in embryo, the 5th slightly the shortest, the 3rd
slightly the longest, the 4th above origin of pectoral. Nostril moderately oblique, its inner
end nearer to mouth than to tip of snout by a distance a little shorter than diameter of eye,
its inner margin slightly expanded in obtusely subangular outline near inner end. Mouth
about twice as broad as high.

Teeth ' '' " ti^H— T- u'^H r^ > uppers broadly triangular, the ist and 2nd nearly sym-
metrical, but subsequent teeth increasingly oblique, the inner margins weakly convex

53. All the embryos listed above show this ridge more or less clearly.

54. We did not appreciate the importance of this character at the time when we examined the adults listed above in
the fresh state.

55. About 50 per cent that great as calculated in adults, 42 to 43 per cent by direct measurement in embryos.

358 Memoir Sears Foundation for Marine Research

toward tip but weakly concave toward base, the outer margins moderately concave (most
so toward base), the outermost 3 or 4 teeth being definitely notched outwardly; both
margins strongly and evenly serrate from base nearly to tip; lower teeth erect on broad
bases with lanceolate cusps narrowing rather abruptly toward the tip, the apical part of
cusps very finely serrate but bases smooth except toward corners of mouth, where occa-
sional teeth show more or less serration along the basal expansions as well as on the cusp ;
I or 2 minute teeth at symphysis in upper jaw, i in lower; outermost teeth in each jaw
very small.

First dorsal noticeably large, its vertical height a little more than ^2 as great as dis-
tance from tip of snout to 5th gill opening, its origin opposite inner corner of pectoral in
embryo but slightly posterior to it in adult specimens, its anterior margin weakly convex
in adult but strongly so in embiyo, its apex very broadly rounded, its posterior margin
convex near apex but deeply concave toward base (much more so in young specimens), its
free rear corner about V3 as long as the base, the midpoint of its base considerably nearer
to axil of pectoral than to origin of pelvics. Second dorsal about Vs as long at base as ist and
slightly less than V3 as high, but with rear corner much more elongate, relatively, and a
little longer than the base, its origin over or slightly before origin of anal. Caudal a little
less than Vs (28%) of total length, its terminal sector about V4 the length of fin, moder-
ately slender, the tip rounded in embryo but subacute in adult, the lower lobe (expanded
lower anterior corner) nearly or quite V2 as long as upper (relatively somewhat shorter
in embryo), its tip very broadly rounded in embryo but altering to subacute with growth,
its posterior margin evenly convex, the re-entrant corner (included between the 2 lobes)
narrowly rounded in adult but broadly so in embryo. Distance from lower precaudal pit
to tip of anal only about V4 as long as base of anal. Anal about as long at base as 2nd dorsal,
with broadly rounded apex, but about i .3 times as high vertically, and with much more
deeply incised rear outline, its free rear tip about as long as its base. Distance from origin
of anal to tips of pelvics only about as long as base of anal. Pelvics about as long as anal
along anterior margins. Pectoral as long as, or a little longer than, head, or slightly less
than ^/4 the total length,°° about 2.3 times as long as broad, its anterior margin moderately
convex (increasingly so toward tip) in adult and very strongly so in embryo, its distal
margin moderately and increasingly concave toward inner corner, the inner corner moder-
ately rounded, the tip similarly rounded in adult, but much more broadly so in embryo.

Color. Varying from light gray or pale brown to slaty-blue above and yellowish or
dirty white below. In the two fresh adults we have seen, the pelvics and the lower surfaces
of the pectorals were spotted with gray, the tips of the dorsals being grayish white and
similarly spotted. But in some cases these fins, as well as the caudal lobes and the pectorals,

56. Pectoral a little longer relatively in one of the two adults measured and a little shorter in the other. Among
seven embryos (five of them from one brood), the ratio of length of pectoral to length of head (snout to pec-
toral origin) is i :i.2 at 395 mm. (male) ; i :i at 460 mm. (male) ; and from about i :o.9 to about 1:1 in 5
females of 555 to 580 mm.; this range of variation shows a small increase in the relative length of the fin with
growth. In a Mediterranean embryo of 410 mm., reported by Moreau (Poiss. France, Suppl., 1891: 7), the
ratio was about i :i.5.

Fishes of the Western North Atlantic 359

are white tipped. In embryos, however, the dorsals, pectorals, pelvics and lower lobe of
caudal are more or less conspicuously tipped with sooty gray or black.

Size. The stage of development of the embryos listed above (up to about 580 mm.
long) suggests a length of perhaps 650 to 700 mm. at birth. Maturity probably is not
reached at less than about six feet. The longest for which we find exact measurement was
3.5 meters (about 1 1 V2 feet). The maximum size is said to be 12 to 13 feet, but we think
it likely that at least some may grow considerably longer, for larger sharks, apparently of
this species because of the rounded shape of the first dorsal fins, have been described re-
cently to us as seen at the surface over the continental slope in the offing of Woods Hole
from the research vessel "Atlantis."

Developmental Stages. Development is viviparous j the embryos which we have col-
lected" have a long umbilical cord, about 410 mm. in length, terminating in a well devel-
oped yolk-sac placenta by which they were attached to the uterine wall of the mother,
C. longimanus is also peculiar among carcharhinids for the very considerable changes in
the shape of its fins with growth, as illustrated by drawings of the embryo and mother
shown in Fig. 64, and emphasized above in the description. It is interesting that in one
case all members of a litter of embryos were of the same sex (female), whereas in another
case both sexes were represented (see Study Material, p. 354).

Remarks. C. longimanus very commonly has been confused with leucas, which it re-
sembles in general form of trunk, very broad head, very short and broadly rounded snout,
low-arched mouth, relative positions of fins, and teeth; consequently the synonymy of the
two species is almost hopelessly confused. Actually, however, the two species are separable
at a glance by the shape of the first dorsal fin (strongly rounded in longimanus, but sub-
angular in leucas) ; also the tip of the anal reaches nearly to the precaudal pit in longimanus
but falls considerably short of it in leucas, and the pectoral is much longer, relatively, in
the former than in the latter. The outline of the lower lobe of the caudal is also distinctive,
being convex posteriorly and nearly straight anteriorly in adult longimanus with the re-
verse in leucas, while the margins of the dermal denticles are much more strongly toothed
in leucas than in longimanus. The differences in the shapes of the fins, especially the pec-
toral, are even more striking in the case of embryos than of adults.

Habits. Astonishingly little is known of the habits of longimanus, considering that it
is one of the members of its genus that has been recognized the longest. Apparently it is
more strictly pelagic than any other members of Carcharhinus in the western Atlantic, and
more strictly tropical there than most of them are. We have not found a single report of
one caught from the beach or taken in a pound net anywhere along the coast of the United
States that can be referred with certainty to this particular species.'* And while "Carcharias

57. See Study Material, p. 354.

58. Jordan, it is true (Bull. U.S. nat. Mus., 7, 1884: 104), has characterized a large shark under the name Car-
charias lamia as very common around wharves and off the Keys of southern Florida, adding that one of 5 or 6
embryos which were taken from a yyi-ioot female was kept. But the subsequent illustration, probably of this
specimen (Jordan and Evermann, Bull. U.S. nat. Mus., 47 [4], 1900: pi. 5, fig. 17), is not of a longimanus but
apparently of a leucas.

360 Memoir Sears Foundation for Marine Research

(Prionodon) lamia" has been said to run up into fresh water in Senegal," there is no
way of knowing whether this report (by name only) actually referred to longimanus or to
some other carcharhinid, the latter being more likely. In the Mediterranean it is often
mentioned as entering the tuna nets close to land and as being common offshore.

It is not possible as yet to relate its geographical distribution to physical factors in a
satisfactory way. Although it has never been reported reliably in the western Atlantic in
temperatures lower than about 21 to 22° C. or 70 to 71.5° F. (Bahamas, March 19 14),
it is described in the Mediterranean around Sicily as caught most often in winter,*" «.<?.,
when the water is only about 13 to 15° C. (about 55.5 to 59° F.). Hence, its failure to visit
the coasts of the eastern United States in summer (see below) is less likely to be due to
unfavorable temperature than to low salinity, for it occurs chiefly where the water is more
saline than 35.5 per mille, or even more than 36 per mille. If such is the case, it is excep-
tional among sharks.

In the Mediterranean it is said to destroy large numbers of fish 5 we know from per-
sonal experience that it takes a large bait readily, and it is so well armed that it would not
be astonishing if it preyed on large as well as small fishes, or on sea turtles. But no precise
information is available as to its diet.

Of its breeding habits it is only known that a female, caught off the north coast of
Cuba in May (see Study Material, p. 354), contained six embryos.

Relation to Man. This species has never been of commercial importance anywhere.
On the other hand, it has been accused vaguely of being a man-eater, but we do not know
on what evidence.

Range. Tropical and subtropical Atlantic. In the east it is well known in the Mediter-
ranean and along the Iberian peninsula, and is reported by name from off Cape Verde,"
where it is certainly to be expected, and from Senegalj*' but it seems likely that its re-
ported occurrence in the latter region refers in part to some other shark (see discussion of
its reputed presence in fresh water there, p. 360). In the west its normal zone of occur-
rence is from Uruguay and southern Brazil to the more northerly waters of the West
Indies, and thence northward in the Gulf Stream, perhaps to the ofiing of southern New
England. The species "lamia" has also been reported from various localities in the Pacific
and Indian Oceans, Australia and Red Sea.°^ All but one of these reports are by name only,
however, hence they afford no clue regarding the actual species of Carcharhinus ; and even
that one°' seems actually to have referred to C. brachyurus (Giinther), 1 870, the "Whaler"

59. Steindachner, S. B. Akad. Wiss. Wien, 61 (i), 1870: 576.

60. Doderlein, Man. Ittiol. Medit., 2, 1881:41.

61. Steindachner, S. B. Akad. Wiss. Wien, 61 (i), 1870: 576 (Senegal, in fresh water); Denkschr. Akad. Wiss.
Wien, 44, 1882: 51 (Senegambia) J Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 19 (Cape Verde,
Senegal) .

62. For a list of these records, see Fowler (Bull. U.S. nat. Mus., 100 [/j], 1941: 169) in synonymy of Eulamia
lamia.

63. Phillipps, N. Z. J. Sci. Tech., 6, 1924: 260, fig. 3 ; we may point out that his illustration (fig. 3) is apparently

Fishes of the Western North Atlantic 361

of New Zealand seas, a species only remotely resembling the Atlantic longimanus. Twelve
out of 1 3 other western Pacific-Indian Ocean species included by Fowler" in the synonymy
of his Eulamia lamia equally fail to show the combination of characters most distinctive of
longimanus, at least if the published accounts of them are to be relied upon. The several
species of Carcharhinus that occur along the Pacific coast of Central America are also clearly
separable from longimanus by one character or another."* Although insularum Snyder,
1904,°° from the Hawaiian Islands, does resemble longimanus in the roundness of its first
dorsal, in the close proximity of the tip of its anal to the origin of the caudal, and in its
teeth (particularly in the serration of the lowers), its pectoral appears to be definitely
much shorter than that of longimanus."

Occurrence in the Western Atlantic. Definite information as to the actual frequency
of occurrence of this species in the western Atlantic is astonishingly scant, partly because
it is so seldom encountered in continental waters there, but equally because it has been
confused so often with C. leucas, and perhaps with C. obscurus also. The only reports of it
there that include evidence as to their actual identity are for: Uruguay; the Island of South
Trinidad (Lat. 20° 30' S., Long. 29° 23' W.) off southern Brazil; northern Brazil
south to Rio de Janeiro; Santa Cruz, Dominica and Guadeloupe, West Indies; off the
north and south coasts of Cuba; three stations off Florida north of the Bahama Bank («.<?.,
seaward of the Gulf Stream) ; and the Caribbean in general. But the wide distribution
of these localities proves it to be generally distributed in the western side of the tropical
Atlantic. And the fact that we counted 28 and caught one on an occasion from the research
ship "Atlantis" off the north coast of Cuba in May of 1 939 is in line with earlier characteri-
zations of it as abundant in the Caribbean-West Indian region, and with reports to us of
"White-finned" sharks being seen there very often. The scanty information available
suggests that it is also common offshore around Bermuda but not inshore.

We find no reliable record of it for the coast of Florida"* or for anywhere else on the
east coast of the United States farther north, which is in agreement with its oceanic nature.
But a school"" of large sharks, apparently of this species, was encountered at the surface
over the continental slope by the research vessel "Atlantis" on one occasion in June 1941
in the offing of southern New England (about Lat. 39° 30' N., Long. 70° 30' W., see
P- 3 59) J suggesting that the transition-band between oceanic and continental waters is its
normal boundary in the western Atlantic north of tropical latitudes.

copied from the one by Jordan and Evermann (Bull. U.S. nat. Mus., 47 [4], 1900: pi. 5, fig. 17) of an em-
bryo, probably of C. leucas i see also footnote 58, p. 359.

64. Bull. U.S. nat. Mus., 100 (xj), 1941 : 169.

65. For the most recent survey of these, see Beebe and Tee-Van, Zoologica, N. Y., 26, 1941 : 106.

66. Bull. U.S. Bur. Fish, s2, 1904: 513, pi. i, fig. i, Carcharias insularum.

67. Pectoral only 80 per cent of length of head to origin of pectoral in adult insularum, and 83 per cent of length
of head in embryo, whereas in longimanus it is about as long as the head or longer.

68. It is not included in the most recent survey of the sharks of Florida (Springer, Proc. Fla. Acad. Sci., 5, 1939:
9-41) ; an earlier characterization of it by Jordan (Bull. U.S. nat. Mus., 7, 1884: 104) as plentiful there seems
actually to have referred to C. leucas; see footnote 58, p. 359; jaws from two Florida localities were re-
ported by Fowler (Proc. Acad. nat. Sci. Philad., 60, 1908: 65) as only "probably" this species.

69. Described as "several hundred."

362 Memoir Sears Foundation for Marine Research

Synonyms and References:'"

Squalus carcharias (in part) Linnaeus, Syst. Nat., t, 1758: 235 ; i, 1776: 400 (this appears to be a combination
of the present species with Carcharodon, of which 5. carcharias Linnaeus is now universally considered
the type; for discussion, see Jordan, Copeia, 166, 1928: 4) ; Walbaum, P. Artedi Genera Pise. Emend.
Ichthyol., 5, 1792: 514 (after Linnaeus); Bloch and Schneider, Syst. Ichthyol., 1801: 132 (after
Linnaeus); Risso, Ichthyol. Nice, 1810: 25 (descr., a combination of this species with Carcharodon,
Medit.); Gray, Cat. Fish. Brit. Mus. descr. by L. T. Gronow, 1854: 5 (combined in descr. with Car-
charodon, Medit., Atlant.).

Squalus sp. (in part) Gronow, Zoophyl., i,'l763: 32 (incl. in synon., not seen).

Le Requin, Cloquet, Diet. Sci. Nat., Atlas. Poiss., 181 6-1 830: pi. 26 (clearly recognizable ill.) ; not Le Requin,
Cloquet (Diet. Sci. Nat., 7, 181 7: 69, which is Carcharodon).

Carcharias lamia Risso, Hist. Nat. Europ. Merid., j, 1826: 119 (descr., Medit.); Bory de St. Vincent,
Diet. Class. Hist. Nat., 15, 1829: 598 (name only, by ref. to Risso, 1826); Jordan and Gilbert, Bull.
U.S. nat. Mus., 16, 1883: 873 (by ref., coasts of Europe, not Carcharinus lamia, p. 23, which was the
tooth of a Carcharodon) ; Carus, Prod. Fauna Medit., 2, 1 889-93 : 5 1 2 (by refs., Medit.) ; Ribeiro, Arch.
Mus. nac. Rio de J., 14, 1907: 154, pi. 4 (descr., photo of jaws, north. Brazil south to Rio de Janeiro) ;
Fauna brazil. Peixes, 2 (l) Fasc. I, 1923: 11 (same as preceding) ; Nobre, Fauna Marinha Port. Vert., i,
1935: 422 (descr., off Portugal).

Carcharias {Prionodon) lamia Miiller and Henle, Plagiost., 1841: 37, pi. 12 (descr., excellent ill., common in
Medit. and Ocean) ; Dumeril, Hist. Nat. Poiss., j, 1865: 356, in part (good descr., but specs, from India,
Aust. and C. of Good Hope probably not this species); Giinther, Cat. Fish. Brit. Mus., 8, 1 870: 372
(descr., Medit., Atlant.) ; Doderlein, Man. Ittiol. Medit., 2, 1 881: 40 (good descr., season, voracity, dam-
age to nets, Medit.).

Squalus Ion gimanus Poey, Memorias, 2, 1861: 338 (excellent descr., Cuba).

Prionodon longimanus Poey, Memorias, 2, 1861 ; pi. 19, fig. 9, 10 (excellent ill. of teeth, Cuba).

Eulamia lamia Gill, Ann. N. Y. Lye, 7, 1862: 401 (name only, with Gill as authority, see footnote 3,
p. 320) ; Poey, An. Soc. esp. Hist, nat., 5, 1876: 384; Enumerat. Pise. Cubens., 1876: 188 (size, ref. to
earlier descr., Cuba) ; Fowler, Bull. U.S. nat. Mus., 1 00 {13), 1941: 170 (Atlant. refs. in synonymy at
least in part, but not Pacif. refs.) ; Arqu. Zool. Estado Sao Paulo, 5, 1942: 128 (Brazil).

Prionodon lamia Bocage and Brito Capello, Poiss. Plagiost. Port., 1866: 18 (brief descr., size, off Portugal).

Eulamia longimana Poey, Repert. Fisico-nat. Cuba, 2, 1 868 : 448 (descr., one 3.5 meters long, Cuba) ; Bigelow
and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 80, fig. 24 (descr.,
habits, range, ill.).'"^

Prionodon lamia Canestrini, in Cornelia, et al., Fauna d'ltal., 3, Pesci, 1 871-1872: 48 (ident. by refs., size,
Medit.).

Carcharias {Prionodon) obtusus Garman, Bull. Harv. Mus. comp. Zool., 8, 1881: 232, at least in part (ident.
by descr. of long pectorals, rounded dorsal, white-tipped fins, number of embryos; Cuba, Santa Cruz,
Guadeloupe, Dominica).

Carcharias obtusirostris MoTczu, Hist. Nat. Poiss. France, /, 1 881: 332 (descr. at least in part, but perhaps com-
bined with some other Carcharhinus, Medit.) ; Poiss. Franc, Suppl., 1 891: 7 (meas. of embryo 420 mm.,
Medit.).

Carcharhinus {Eulamia) lamia Jordun, Rep. U.S. Comm. Fish. (1885), 1887: 796 (name by ref., W. Europ.).

Carcharhinus (Carcharhinus) lamia Jordun and Evermann, Bull. U.S. nat. Mus., 47 (l), 1896: 38 (descr.,
probably in part, and by refs., but apparently confused with C. leucas).

Carcharinus commersonii (in part) Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 140 (descr., Atlant.
refs. in part, but not Pacif.) ; Radcliffe, Bull. U.S. Bur. Fish., 34, 1916: 260 (meas. of specs, taken north
of Bahama Bank, but not fig. of denticles, or N. Carolina records which probably were leucas) ; Meek and
Hildebrand, Field Mus. Publ. Zool., 15 (l), 1923: 43 (Atlant. refs., in part only; not descr., which is of

70. This species has been confused so commonly with others that we limit the following list to such citations as con-
tain evidence that they did in fact refer to this particular species, at least in part, and not to some other. For a list
of references to Carcharhinus that cannot be allocated with certainty to any particular species, see p. 363.

70a. Spelled longimanus.

Fishes of the Western North Atlantic 363

a Careharhinus with short pectorals) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2,

1930: 15 (synonymy in part); White, Bull. Amer. Mus. nat. Hist., 7^, 1934: 127, but not pi. 42E

(included in synonymy, but ill. of heart valves probably leucas).
Carcharinus flatyodon (in part) Garman, Mem. Harv. Mus. comp. Zool., 36, 1 91 3: 126 (descr. of embryo,

in Harv. Mus. Comp. Zool., Guadeloupe, W. I., see Study Material, p. 354; but not synonymy, and not

pi. 3, fig. 4-6, which is a mislabeled milberti) ; Devincenzi, An. Mus. Hist. nat. Montevideo, (2) 4, 1939:

5 (jaws, teeth, apparently this species, Uruguay).
Careharhinus commenonii Jordan, Copeia, 49, 1917: 87 (name only, substituted for lamia); Norman and

Fraser, Giant Fishes, 1937: 36 (distrib., based on old accounts).
Careharhinus sp. Nichols and Murphy, Bull. Amer. Mus. nat. Hist., jj, 1914: 262 (descr. of embryo. South

Trinidad I., S. Atlant., Lat. 20° 30' S., Long. 29° 22' W.).
Coreharias commersonii (in part) Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda, 1933: 28 (includes

both this species and leucas, Bermuda).

Doubtful References:

Squale requin Lacepede, Hist. Nat. Poiss., 4° ed., 7, 1798: 169, but not pi. 8, fig. I j^"^ in Soninni, Hist. Nat.

Poiss., ^, 1802: 332 (a rewrite of Lacepede's account, but the illustrations, pi. 6, are of the teeth and jaws

of Carcharodon, combined with a general view so poor that it might be interpreted as representing either

the latter or one of the larger carcharhinids) .
Careharhinus lamia Tonontit, Atti Soc. ital. Sci. nat., jy, 1828: 298 (discus., Medit.).
Coreharias coreharias (in part) Jordan, Manual Vert. Anim. NE. U.S., 1929: 10 (class., descr., apparently

based partly on longimanus and partly on leucas, S. Atlant., U.S.) .

References, Ostensibly longimanus, But Which Cannot be Definitely Identified as Any Particular Species of the

Genus:

Coreharias {Prionodon) lamia Steindachner, S. B. Akad. Wiss. Wien, 61 (l), 1870: 576; Denkschr. Akad.

Wiss. Wien, 44 (i), 1882: 51 (Senegambia, name only).
Corcharinus lamia Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 48 (south. Florida);

Bean, Field Mus. Publ. Zool., 7 (2), 1906: 29 (Bermuda); Fowler, Proc. Acad. nat. Sci. Philad., $S,

1906: 80 (Florida Keys); Gudger, Proc. biol. Soc. Wash., 26, 1913: 97 (C. Lookout, N. Carolina);

Coles, Proc. biol. Soc. Wash., 2S, 191 5: 90 (Carolina); Nichols, Copeia, 53, 1918: 13 (Florida); Borri,

Mem. Soc. tosc. Sci. nat., 44, 1934: 89 (Medit.) ; Norman and Fraser, Giant Fishes, 1937: 36 (Medit.,

trop. Atlant.).
Prionodon lamia de Braganza, Result. Invest. Sci. "Amelia" Ichthyol., 2, 1904: 44, 45 (Portugal); Richard,

Bull. Inst, oceanogr. Monaco, rp, 1904: 1 1 (Canaries).
Coreharias lamia Jordan and Thompson, Bull. U.S. Bur. Fish., 24, 1905: 232 (Tortugas, Florida) ; Ribeiro,

Rev. Mus. paul., 10, 1918: 707 (Santos, Brazil); Marini, Physis B. Aires, 9, 1929: 452 (Argentina);

Pozzi and Bordale, An. Soc. cient. argent., 120, 1935: 150 (Argentina).
Eulamia longimanus Fowler, Proc Acad. nat. Sci. Philad., 60, 1908: 65 (Florida).
Eulomia commersonii Fowler, Proc. Acad. nat. Sci. Philad., 69, 1917: 127 [Colon, provisional ident. by ref.

to Muller and Henle's (Plagiost., 1 841: 37, pi. 12) account of Coreharias (Prionodon') lamia],
Eulamia commersonii (in part) Fowler, Bull. Amer. Mus. nat. Hist., yo, 1 936: 53, pi. i [trop. W. Afr. by

ref. to Coreharias {Prionodon) lamia Steindachner, 1870, 1882, but descr. i$ of a Florida spec, probably

leucas] .
Eulamia lamia Fowler, Monogr. Acad. nat. Sci. Philad., 1945: 95, 263 (size, Marco, Florida, off N. Carolina).
Not Coreharias lamia Rafinesque, 1810; see footnote 51, p. 354.
Not Careharhinus lamia Blainville, in Vieillot, Faune Franc, 1825: 88, pi. 22, fig. I^^ (jaws, ident. by ref. to

Duhamel, Traite gen. Peches, 4 [9], pi. 19, as Galeocerdo).

71. Much of the confusion in shark nomenclature comes from uncertainty as to actual identity of Lacepede's Squale
requin. The measurements on pag'e 184 might apply to longimanus ; but the account of the first dorsal, and its
representation in the illustration as only a little rounded at apex, do not. Most of the description is devoted to a
general account of the larger carcharhinids, in general, perhaps combined with Carcharodon.

72. We cannot find whether or not this plate was ever published.

364 Memoir Sears Foundation for Marine Research

Not Carcharias (Prionodon) lamia Putnam, Bull. Essex Inst., 6, 1874: 72 (this was a Carcharodofi).

Not Prionodon lamia Goode and Bean, Bull. Essex Inst., 11, 1879: 30 (the specimen of Carcharodon reported

by Putnam, 1874).
Not Eulamia lamia Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 60 (descr. of San Diego, Calif.,

spec, actually tomidla, Jordan and Gilbert, 1882).
Not Carcharias lamia Jordan, Proc. U.S. nat. Mus., 7, 1884: 104 (almost certainly refers to leucas; see

p. 344).
Not Carcharhinus lamia Jordan and Evermann, Bull. U.S. nat. Mus., 47 (4), 1900: pi. 5, fig. 17 (embryo,

evidently of leucas, see p. 344).
Not Carcharias lamia Lahille, An. Mus. nac. B. Aires, 34, 1928: 305 (appears to be remotus, see p. 403).

Carcharhinus macuUfinnis (Poey), 1865

Large Black-tipped Shark

Figures 66., 61

Study Material. Female, about 6 feet 6 inches (1,975 mm.) long, from Salerno,
Florida (Harv. Mus. Comp. Zool., No. 35772) and jaws of two males, about 6 feet
4 inches (1,915 mm.) long, from Englewood, Florida (U.S. Nat. Mus., No. 109957,
I 10306).

Distinctive Characters. Among the smooth-backed Carcharhinus of the Atlantic,
maculipinnis is most like limbatus in general appearance, fins, teeth and color. But it is
separated from limbatus by its noticeably smaller eyes, its relatively longer upper labial
furrows and gill openings, the noticeably more slender free rear tip of its second dorsal and
the perfectly smooth edges of its lower teeth (finely serrate in limbatus^. Also, its first
dorsal usually originates somewhat farther rearward, although there may be some varia-
tion in this respect. Its dermal denticles are distinctive, also (cf. Fig. 66 A with 62 D).

Description. Proportional dimensions in per cent of total length. Female, 1,975 mm.,
from Salerno, Florida (Harv. Mus. Comp. Zool., No. 35772).

Snout length in front of: outer nostrils 4.85 mouth 8.0.

Eye: horizontal diameter 0.9.

Mouth: breadth 9.1 j height 4.9.

Nostrils: distance between inner ends 5.3.

Labial furrow length /upper i . i .

Gill Of ening lengths : ist4.ij 2nd4.4j 3rd 4.85 4th 4.7; 5th 3.5.

First dorsal fin: vertical height 9.0 ; length of base 9.7.

Second dorsal fin: vertical height 2.65 length of base 3.7.

Anal fin: vertical height 2.75 length of base 4.2.

Caudal fin: upper margin 26.5 } lower anterior margin 12.2.

Pectoral fin: outer margin 17.2; inner margin 4.85 distal margin 15.0.

Distance from snout to: ist dorsal 32.5; 2nd dorsal 65.2; upper caudal 73.55

pectoral 23.95 pelvics 52.05 anal 65.2.

Fishes of the Western North Atlantic

365

Intersface between: ist and 2nd dorsals 21.7; 2nd dorsal and caudal 5.3; anal and

caudal 5.4.

Distance from origin to origin of: pectoral and pelvics 27.7; pelvics and anal 13.5.

Trunk moderately slender, its height at ist dorsal (where highest) between Vs and
Yq its length to precaudal pits, tapering both anteriorly and posteriorly, the midline of

Figure 66. Carcharhittus maculifinnis, female, about six feet six inches (1,975 mni.) long, from Salerno,
Florida (Harv. Mus. Comp. Zool., No. 35772). A Dermal denticles, about 18 x. B Side and apical views of
dermal denticle, about 1 8 x. C Upper and lower teeth, right-hand side, about % natural size. D Fourth upper
tooth. E Twelfth upper tooth. F Fourth lower tooth. G Twelfth lower tooth. D—G, about 2 x.

Figure 67. Carckarhinui maculifinnis, illustrated in
Fig. 66. A Anterior part of head from below. B Left-
hand nostril, about 1.2 x.

366 Memoir Sears Foundation for Marine Research

back evenly rounded, without dermal ridge. Precaudal pits semilunar in outline, the
upper the deeper, but the lower also well marked. Dermal denticles so close-spaced and
evenly overlapping that the skin is wholly concealed, but their blades raised enough from
skin to feel slightly rough when stroked from rear to front, broader than long, with
mostly 7 (rarely 5) low ridges, their posterior margins broadly ovate or subangular, with
7 very short teeth (shorter than in limhatus) or even entire in some cases; pedicels very
short.

Head a little less than V4 of total length and flattened above anterior to gill region.
Snout rather thin-tipped, ovate, with narrowly rounded tip (relatively somewhat longer
and more pointed than in limiatus), its length in front of a line connecting outer ends of
nostrils about % as great as distance between inner ends of latter, and length in front of
mouth about Vs as great as length of head. Eye approximately circular, and noticeably
small, its diameter only about Vo (11%) as great as length of snout in front of mouth.
Gill openings noticeably large, the 3rd (longest) nearly as long as distance between nos-
trils or about 5 times as long as diameter of eye (only about 2.5 times as long as diameter of
eye in limbatus), the 5th a little less than % (73%) as long as 3rd, the 4th over origin of
pectoral. Nostril strongly oblique, its inner end nearer to mouth than to tip of snout by a
distance a little more than twice as great as diameter of eye, its anterior margin sinuous,
with low, rounded expansion near inner end. Mouth ovate, about Yo as high as broad,
occupying only about % of breadth of head. Upper labial furrow about as long as nostril,
thus considerably longer relatively than in limbatus.

Teeth ^^ "^ Hzl^il'^ti" ^^ '"^ specimen examined; uppers with narrow triangular cusps
on broad bases, smaller and relatively narrower than in limbatus (cf. Fig. 66 C-E with
62 C and 63 C, D), the first 3 nearly symmetrical and erect, but 4th and subsequent teeth
slightly oblique, with outer margin more deeply concave than the inner toward base, and
decreasing in length toward corner of jaw, the outermost 3 very low, the edges regularly
though finely serrate, except that the tips are smooth-edged; lowers with much more
slender cusps than uppers, on very broad bases, their tips not recurved forward as they are
in limbatus, very slightly oblique along whole length of jaw, the ist smaller than 3rd,
and 1 2th and 1 3th to 1 6th again successively smaller, the edges perfectly smooth on base as
well as on cusp; 2 minute teeth at symphysis in upper jaw and i in lower.

Origin of ist dorsal a little posterior to inner corner of pectoral, its apex rather
narrowly rounded, its free rear corner relatively obtuse and only about Yz as long as its
base, the base about % as long as anterior margin, its vertical height a little less than Y2 as
long as head or about V2 as long as pectoral, the midpoint of base about % as far from axil
of pectoral as from origin of pelvics. Second dorsal a little less than Yz as long at base as
1st dorsal, its origin about over origin of anal, its free rear tip about as long as the base,
and noticeably more slender than that of ist. Caudal about Yi of total length, the ter-
minal sector a little less than Yi of the fin, slender, with narrowly rounded tip, the
lower lobe (expanded lower anterior corner) a little less than V2 as long as upper, with
moderately convex lower anterior margin and very narrowly rounded or subacute tip, the

Fishes of the Western North Atlantic 367

re-entrant corner between the two lobes rather broadly rounded/* Anal slightly longer at
base than 2nd dorsal, its outline presumably similar to that of limbatus but damaged in the
specimen we have seen, its free rear corner more obtuse and only a little more than V2 as
long as the base. Distance from origin of anal to tips of pelvics about i V2 times as long as
base of anal. Pelvics a little longer than 2nd dorsal along anterior margin. Pectoral a little
less than % (72%) as long as head, and about V2 as broad as long, similar in form to that
of limbatus, with very narrowly rounded or subacute apex and inner corner.

Color. Varying shades of gray above, the colors said to be more intense in life than in
limbatus; white or whitish below, resembling limbatus in having a narrowing band of the
darker tint of the upper parts extending rearward to about over the origin of the pelvics;
above this there is a corresponding extension forward of a narrowing band of white (or
whitish) from the lower sides to a point % of the distance forward from the origin of
the pelvics toward the axil of the pectoral j" lower lobe of caudal, apex of 2nd dorsal and
lower surfaces of tips of pectorals broadly and conspicuously tipped with black, with apex
of ist dorsal narrowly so, much as in limbatus.

Size. The few specimens of both sexes for which the sizes have been recorded have
ranged from five feet eight inches to about eight feet in length; it apparently grows
larger than limbatus.'"

Developnental Stages. We have a photograph of a female six feet three inches from
Florida with ten young."

Habits. This shark has been seen in schools and leaping at the surface, in which
habit it resembles limbatus, but nothing definite is known of its diet, other than that in
Florida waters it follows shrimp trawlers to pick up the discarded fish; nothing is known
of its life in other respects.

Range. So far reported only from Cuba (the type locality), from Porto Rico by name
only, and from both coasts of southern Florida. Although more than three-fourths of a
century has passed since m^culifinnis was first described, it generally has been confused
with limbatus and until very recently" reported under that name, if at all; consequently
the published records afford no information as to the details of its occurrence, other than
that it is common in winter off southeastern Florida, also off northeastern Florida (Jack-
sonville) in the spring," and that the local shark fishermen are familiar enough with it to
have recognized it as distinct from limbatus. But whether or not it ranges northward in
summer like limbatus is not yet known.

Relation to Man. This, like various other species, forms part of the catch of the

74. Slight apparent differences in the shape of the caudal between maculifinms and limbatus may represent individual
variation.

75. These contrasting- lighter and darker bands are still visible in the preserved specimen, although the latter is now
much discolored in the preservative.

76. Females of eight feet were recently reported to us off Salerno, Florida, by Stewart Springer.

77. From Stewart Springer.

78. Springer (Proc. Fla. Acad. Sci., j, 1939; 27) was the first to show that maculipinnis is not a synonym of
limbatus and that it is a distinct and easily recognizable species.

79. Personal communication from Stewart Springer.

368 Memoir Sears Foundation for Marine Research

Florida shark fishery. But no information is available as to its percentage in the total.
Recently an attack was reported on a bather at Mayport near Jacksonville by a SV2- to
6V2-foot shark, which, judging by circumstantial evidence, seems to have been maculi-
finnis.^"

Synonyms and References:

Isogomfhoion maculifinnis Poey, Repert. Fisico-nat. Cuba, 7, 1865: 191, pi. 4, fig. 2, 3:2, 1866-

1868: 245, 450, pi. 2, fig. 1-3 (descr., teeth, Cuba); Synop. Pise. Cubens., 1868: 450 (diagn., Cuba);

Springer, Proc. Fla. Acad. Sci., 5, 1939: 27 (descr., size, color, Florida); Lunz, Bull. S. Carolina St.

Planning Bd., 14, 1944: 27 (Florida).
Carcharias limbatus (in part) Gunther, Cat. Fish. Brit. Mus., <?, 1 870: 373 (incl. in synon.) ; Ribeiro, Arch.

Mus. nac. Rio de J., 14, 1907: 200; Jordan, Evermann and Clark, Rep. U.S. Coram. Fish. (1928), 2,

1930: 16 (included in synonymy of limbatus).
Platyfodon (?) maculipnnis Poey, Enumerat. Pise. Cubens., 1876: 197, pi. 9, fig. 6 (ill., denticles, Cuba);

An. Soc. esp. Hist, nat., 5, 1376: 393, pi. 14, fig. 6 (Cuba).
Carcharhinus limbatus (in part) Jordan and Evermann, Bull. U.S. nat. Mus., 47 (l), 1896: 40 ; Garman,

Mem. Harv. Mus. comp. Zool., 56, 1913: 127; Meek and Hildebrand, Field Mus. Publ. Zool., 75, 1923:

41 (included in synonymy of limbatus').
Eulamia maculifinnis Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash.,

1945: 90, fig. 29 (descr., range, ill.).

Doubtful References:

Platyfodon maculifinnis Stahl, Fauna Puerto Rico, 1883: 167 (Porto Rico, name only).

Eulamia flatyodon Fowler, Proc. Acad. nat. Sci. Philad., ji, 191 9: 146 (Jamaica spec, perhaps maculifinnis

because teeth narrow and the lowers "entire").
Not Carcharias maculifinnis Gunther, Trans. Zool. Soc. Lond., 6, 1867: 490.'^

Carcharhinus milberti (Miiller and Henle), 1841"

Brown Shark, Sand-bar Shark

Figures 68, 69

Study Material. Male, 1,496 mm., taken at Woods Hole in August 1941 (Harv.
Mus. Comp. Zool., No. 35370) ; male, 1,400 mm., taken at Woods Hole in August 1944
(not preserved) J male, 2,000 mm. (about 6 feet 7 inches), from Vineyard Sound near
Woods Hole, taken Sept. 18, 1943} jaws (Harv. Mus. Comp. Zool., No. 36032)5 head
and skin of male, about 800 mm., from Somer's Point, New Jersey (Harv. Mus. Comp.
Zool., No. 147)} newborn specimens, 562 mm. and 580 mm. (female and male), from
Woods Hole (U.S. Bur. Fish.) ; young male, 760 mm., from Chesapeake Bay, 540-mm.
specimen, from off Grand Terre, Texas, and one young male, 747 mm., from Virginia

80. Stewart Springer has supplied us with details of this occurrence, mentioned on page 70 also.

81. See p. 35Z.

82. If it is finally proved that the Mediterranean form is identical with the American, the name flumieus Nardo,
1827, must be used for the combined species in place of milberti; see discussion, p. 374.

Fishes of the Western North Atlantic

369

Beach, Virginia (U.S. Nat. Mus., No. 104969, 13540, 119698)5 male embryo, 402 mm.
long, from Englewood, Florida (Harv. Mus. Comp. Zool., No. 35361); head and skin
of female embryo, about 600 mm., from Cuba (Harv. Mus. Comp. Zool., No. 715);

Figure 68. Carcharhinus milberti, female, 1,496 mm. long, from Woods Hole, Massachusetts (Harv. Mus.
Comp. Zool., No. 35370). A Cross section of upper part of trunk, midway between the two dorsal fins, to
show the mid-dorsal ridge. B Left-hand upper and lower teeth, about I x. C Fourth upper tooth. D Ninth
upper tooth. E Third lower tooth. F Eighth lower tooth. C-F, about 1.8 x.

Figure 69. Carcharhinus milberti, pictured in Fig. 68. A Anterior part of head from below. B Left-hand
nostril, about I.4 x. C Dermal denticles, about 36 x.

370 Memoir Sears Foundation for Marine Research

measurements by Stewart Springer of an 8o-inch specimen taken at Woods Hole in Aug.
1942, and of 3 females, 2,125 to 2,220 mm., from Englewood, Florida; also a full-term
embryo, 540 mm., taken in upper Chesapeake Bay (U.S. Nat. Mus., No. 83597).

Distinctive Characters. Among the ridge-backed subdivision of the genus, milberti
after birth is made easily recognizable by the large size of the first dorsal and by its posi-
tion far forward (originating over the axil of the pectoral), and by the free rear corner
of the second dorsal which is only about as long as the base. The wide spacing of the dermal
denticles and their free edges without definite teeth are diagnostic also.

Descriftion. Proportional dimensions in per cent of total length. Male, 747 mm.,
from Virginia Beach (U.S. Nat. Mus., No. 104969). Male, 1,496 mm., from Buzzards
Bay, Mass. (Harv. Mus. Comp. Zool., No. 35370).

Trunk at origin of fectoral: breadth 12.O, — ; height 12.9, — .

Snout length in front of: outer nostrils 4.1, 3.3; mouth 8.2, 7.0.

Eye: horizontal diameter 2.1, 1.5.

Mouth: breadth 9.5, 8.5; height 4.7, 4.1.

Nostrils: distance between inner ends 5.9, 5.5.

Labial furrow length: upper — , 0.5.

Gill opening lengths: ist 3.0, 2.7; 2nd — , 3.0; 3rd — , 3.2; 4th — , 3.0; 5th 2.5,

2.4.

First dorsal fin: vertical height 10.9, 12.4; length of base 13.3, 1 1.3.

Second dorsal fin: vertical height 3.1, 2.9; length of base 6.0, 4.1.

Anal fin: vertical height 3.5, 3.9; length of base 5.7, 5.0.

Caudal fin: upper margin 26.7, 27.8; lower anterior margin 12.O, 12.1.

Pectoral fin: outer margin 18.1, 20.4; inner margin 6.3, 6.$; distal margin 14.4,

17.7.

Distance from snout to: ist dorsal 28.3, 28. 7j 2nd dorsal 60.3, 61.3; upper

caudal 73.3, 72.25 pectoral 24.7, 20.7; pelvics 49.1, 47.1; anal 61.5, 61.0.

Interspace between: ist and 2nd dorsals 20.7, 22.5; 2nd dorsal and caudal 7.4,

7.1 ; anal and caudal 7.4, 7.4.

Distance from origin to origin of: pectorals and pelvics 27.4, 27. i ; pelvics and anal

13-3, I4-4-

Trunk comparatively stout, its height at ist dorsal a little more than Vs as great as
length to origin of caudal, with dorsal profile rather strongly arched. Midline of back
with a low but unmistakable dermal ridge from about under rear tip of ist dorsal to a
point about as far in front of origin of 2nd dorsal as length of base of latter; this also
discernible in embryos, though less prominent. Caudal peduncle % to % as thick as deep.
Upper precaudal pit strongly marked, obtuse-ovate in outline, the lower pit much smaller
than upper and hardly visible in small specimens. Dermal denticles widely spaced, seldom
if ever overlapping, the skin exposed between them, blades thick, strongly convex antero-
posteriorly, usually with 5 high, moderately sharp ridges separated by round-bottomed

Fishes of the Western North Atlantic 371

valleys, their apical margins not definitely toothed but at most slightly sinuous opposite
the ridges; pedicels very short."

Head about V4 of total length or a little less, and very broad forward, its breadth
being nearly as great at eyes as at ist gill opening. Snout broad-ovate, relatively more ob-
tuse in large specimens than in small, its length in front of a line connecting outer ends of
nostrils a little more than Yi ^s great as distance between inner ends of latter, and length in
front of mouth about V3 of length of head to 5th gill opening. Eye approximately
circular, noticeably small in adult, its diameter varying in medium-sized and large speci-
mens from a little more than Vs to a little more than V4 as great as distance between oos-
trils, but relatively larger in young, its diameter being about as long as the 3rd gill open-
ing in late-term embryos, about V2 to % that long in newborn and in adults; its anterior
edge about opposite front of mouth. Gill openings nearly evenly spaced, their outlines
nearly straight or slightly sinuous, the 3rd (very slightly the longest) a little more than
V2 as long as distance between nostrils, the 5th slightly the shortest, the 4th above origin of
pectoral. Nostril strongly oblique, its inner end nearer to mouth than to tip of snout by a
distance a little greater than its own length, its anterior margin expanded in obtusely angu-
lar outline, subacute at apex, and weakly crested opposite latter. Mouth very broadly ovate
or nearly arcuate, its height approximately V2 its breadth (this proportion varies somewhat
from specimen to specimen). Upper labial furrow about ^4 as long as diameter of eye.

Teeth jj'" 1^iH}2 Io 15 j" uppers broadly triangular, their margins regularly but finely
serrate from tip to base, the first 2 erect and nearly symmetrical, but subsequent teeth in-
creasingly oblique, with inner margins slightly convex and outer margins either evenly
concave or slightly subangular in contour, the 9th or lOth and subsequent teeth decreasing
successively in size and height relative to breadth j lower teeth erect, symmetrical, with
narrow triangular cusps on broadly expanded bases, the edges of cusps more finely serrate
than those of uppers, the bases smooth-edged, or at most slightly irregular.

First dorsal origin about over axil of pectoral, its vertical height increasing relatively
with growth from a little greater than distance from eye to ist gill opening in embryo to
about as great as distance from eye to 2nd gill opening when newborn and to about as
great as from eye to 3rd gill opening in specimens of medium size; apex also more broadly
rounded in embryo but very narrowly so in adult, the free rear corner a little more than %
as long as base, the midpoint of base only about % (40%) as far from axil of pectoral
as from origin of pelvics. Second dorsal about V3 as long at base as ist and relatively much
lower, its origin about over origin of anal, its posterior margin only weakly concave, its
free rear corner moderately tapering and only about as long as the base. Caudal a little
more than V4 of total length, the terminal sector between V4 and Vs the length of fin, the

83. Previous accounts (Radcliffe, Bull. U.S. Bur. Fish., 3^, 1916: 257; Springer, Proc. Fla. Acad. Sci., 3, 1939: 13)
locate this loose spacing only along the upper sides; but it extends over the trunk generally, below as well as
above, in the specimens we have examined.

84. In one Woods Hole specimen (Harv. Mus. Comp. Zool., No. 36031) the tooth count isitzL^ilzll, but the
spacing of the lower teeth near the corners of the jaws is irregular, suggesting some abnormality.

372 Memoir Sears Foundation for Marine Research

tip narrowly rounded in adults but more broadly rounded in young, the lower lobe
(expanded lower anterior corner) about % (40 to 44%) as long as upper and with sub-
acute tip, the re-entrant corner, included by the two lobes, well rounded. Distance from
origin of caudal to tip of anal nearly or quite as long as base of anal. Anal about as long at
base as 2nd dorsal, and about as large in area, but with subacute apex and much more
deeply concave posterior margin, its free rear corner about % as long as base. Distance
from origin of anal to tips of pelvics a little longer than base of anal. Pelvics a little longer
at base than anal. Pectoral nearly as long (87 to 97%) as head to origin of pectoral in large
specimens but relatively somewhat shorter (about 73%) in small, a little less than twice
as long as broad, the outer margin only weakly convex, distal margin only weakly concave,
the corners very narrowly rounded in adult but somewhat more broadly so in young.

Color. Varying from slate-gray to brownish gray or brown above, perhaps depending
on color of the environment; a pale tint of the same hue, or whitish, below; fins without
any conspicuous markings. When the shark is newly caught some of the dermal denticles
may be brilliant blue, at least in Florida specimens.

Size. The usual size at birth is said to be about 22 inches (weight about 2V2 pounds).
Maturity is attained at about six feet and it appears that very few reach as great a length
as eight feet; 7 feet lO inches is the longest of which we find unquestionable record.*" If
one about three feet long, taken near New York on June 9, was born the previous sum-
mer or early autumn, as seems probable,*" milberti may be expected to grow by about lO
inches during the first winter, for the newborn young of 25 to 26 inches have been taken
in that general region in September. But nothing whatever is known of the subsequent
rate of growth.

Usual weights of adults are about 100 pounds at six feet, 125 to 130 pounds at about
seven feet, 200 pounds at about 7 feet 8 inches.

Develofmental Stages. Presumably milberti is viviparous, but whether the young
develop placental connection with the mother has not yet been definitely established. Em-
bryos resemble the adults in general; 6 to 13 are recorded in a litter, the usual number
being 8 to 1 2, with the two sexes about equally represented.

Habits. Although this is undoubtedly the most abundant member of its genus in
season along the middle Atlantic coast of the United States, and the one most often seen,
knowledge of its habits is scant. Certainly it is littoral rather than pelagic, for considerable
numbers enter shallow bays, harbors and river mouths; hence, they are often harpooned or
taken in pound nets, and it is said to be the only large shark that regularly visits the small
bays on the north shore of Long Island, New York, as it and others also do the much more

85. Two specimens of milberti of 8 feet 6 inches have been reported from New Jersey, but by name only (Fowler,
Proc. Acad. Nat. Sci. Philad., 72, 1921 : 386) ; an old report (Baird, Rep. Smithson. Instn. [1854], 1855: 352)
that the largest in New Jersey waters are of about nine feet may not have been based on actual measurements;
and the report of a North Carolina milberti of 9 feet 2 inches (Smith, Bull. N. C. geol. econ. Surv., 2, 1907:
35) actually may have referred to some other shark, so far as the brief account goes; perhaps ta the larger-
growing obscurus.

86. Nichols and Breder, Zoologica, N. Y., 9, 1917: 16.

Fishes of the Western North Atlantic 373

extensive enclosed waters along the south shore of Long Island. Similarly, its representa-
tive in the eastern Atlantic is said to enter the larger of the Venetian canals and the neigh-
boring lagoons adjacent to the Mediterranean.'' We have looked down from a beach side
bluff on a middle-sized one swimming so close to the tide line that its dorsal fin was neces-
sarily exposed. Correspondingly, we have never heard of one taken more than a short
distance out from the land. But it appears never to enter fresh water. Although it is an in-
shore species, as a rule it is only when crossing some shoal that the Brown Shark shows
itself at the surface.

It feeds chiefly on fish and on mollusks and Crustacea also, its diet depending on what
may be available for it locally. Near New York, for example, flounders {Pseudopleu-
ronectes) are reported as predominant in its stomach contents, with an occasional eel and
crab; others taken at Woods Hole have been found to have fed on amphipod Crustacea, as
well as on the bivalve moUusk Yoldia, which is plentiful in 2 to lO fathoms in the general
vicinity. Still others off the east coast of Florida had eaten Octopus chiefly, and also small
fish and crabs." Skates and Dogfish are listed for it, and it is able to capture fast-swimming
fishes also, for bonito (Sarda), weakfish (Cynoscion), mackerel, menhaden (Brevoortia)
and pinfish (Lagodon) are included in its known diet. But there is no reason to suppose
it ever attacks larger prey.

In the bays of Long Island, New York, its young are born from June until August.
It seems to be chiefly for that purpose that it enters those very shoal waters, for a great
majority of the adults taken there are females, males being very few in number and re-
ported only for August. A large proportion of the adult females that visit the shallow
bays of Long Island carried embryos nearly ready for birth. Newborn specimens have often
been taken there in summer and early autumn, as well as in Chesapeake Bay in September,
but on the other hand, no young ones have been reported from Florida, although some of
the large females taken there carry embryos. These facts are evidence that the young are
produced chiefly in the northern part of its range. It seems that this applies equally to such
milberti as visit the Gulf of Mexico, for the only Texas record is for a newborn specimen
taken in early August.

Relation to Extralimital Species. Milberti is so closely allied to azureus Gilbert and
Starks, 1903, of Ecuador and the Pacific coast of Central America that the latter has been
classed by some as probably identical with it.*' But the most recent illustration of azureus^"
shows it to be easily separated from milberti, the origin of its second dorsal being consider-
ably anterior to that of the anal, instead of about over the latter, to mention only the most
obvious difference between the two.

Relation to Man. Milberti is reported as the rnost abundant of the commercially valu-
able sharks taken off southeastern Florida. Some are also caught by anglers, and a few

87. Nardo, Sopra due Sp. Pesci Nuov. Venet., 1853: 15. 88. Personal communication from Stewart Springer.

89. Garman, Mem. Harv. Mus. comp. Zool., j6, 1913 : 133 ; Meek and Hildebrand, Field Mus. Publ. Zool., 15 (i),
1923: 38.

90. Beebe and Tee-Van, Zoologica, N. Y., 3d, 19^1 : 109, fig. 18.

374 Memoir Sears Foundation for Marine Research

sportsmen have thought it worth special pursuit with the harpoon." We have never heard
any rumor of its molesting bathers, nor would this be likely, judging from the nature of
its prey.

Range. Western Atlantic, from southern New England to southern Florida, Louisi-
ana and southern Brazil. It has been reported also from various localities in the Mediter-
ranean, southern coast of Spain (but whether inside or outside the Straits of Gibraltar is
not stated), the Canaries and Cape Verdes. But we still await actual comparison of speci-
mens from the two sides of the ocean."

Occurrence in the Western Atlantic. Owing to a long-standing tendency to call any
Carcharhinus encountered on the middle Atlantic coast of the United States a Dusky Shark,
milberti has been reported so often as obscurus that little dependence can be placed on pub-
lished locality records for it unless these are accompanied by some evidence of actual iden-
tity more convincing than the mere name. However, information is at hand to show that
it is a regular seasonal migrant along the east coast of the United States. Thus it is present
along both coasts of southern Florida from December to March, being perhaps the most
abundant of the species caught commercially at Salerno on the east coast at that season," and
it visits the coasts of New Jersey and New York regularly only in summer though in such
numbers that it has been repeatedly described as "abundant" and is to be seen most any
summer day in the bays of New Jersey." Recent reports of 305 sharks being harpooned in
Great South Bay, Long Island, during the summers of 191 1 to 1927 (almost all being
milberti) J of 46 being taken in one summer, and of 14 being harpooned there in one day
(August 1 1, 1906)," give a more precise indication of the actual numbers concerned. It is
rather common off Rhode Island also during the warm months, occasionally entering
Narragansett Bay. Likewise it visits the Buzzards Bay-Vineyard Sound-Nantucket Sound
region yearly, but so much less abundantly that the number taken near Woods Hole m an
average summer probably does not exceed six or seven. And Cape Cod so sharply marks
the usual limit to its northerly dispersal that there is no reliable record of it for the Gulf
of Maine,"' for the fishing banks off its mouth or for Nova Scotia. In the vicinity of New
York its season of maximum abundance (mostly females as noted above) is from mid-
June to mid-September, the latest for Sandy Hook Bay being October 195 extremes re-
ported for it at Block Island are May and November.

91. For a vivid account of harpooning milberti in the bays of Long Island, New York, see Nichols and Murphy
(Brooklyn Mus. Sci. Bull., 3 [i], 1916: 16).

92. The view is generally held that the milberti of the two sides of the Atlantic are one species. Although one from
southern Spain, described by Rey (Fauna Iberica, Feces, /, 1928: 346), agreed with American specimens as to
fins and teeth, the account does not state whether or not it had a mid-dorsal ridge; its denticles, too, were more
closely spaced, and their margins more definitely dentate than in those we have examined.

93. Knowledge that milberti occurs around southern Florida dates only from the recent development of the local
fishery; the only previous Florida record for it, and that by name only, was for the Indian River (Goode, Proc.
U.S. nat. Mus., j, 1879: 121).

94. Baird, Rep. Smithson. Inst. (1854), p, 1855: 352.

95. Rockwell, Brooklyn Mus. Quart., 3, 1916: 160-167; Thome, Bull. N. Y. zool. Soc, 5/, 1928: 114.

96. An early statement that it ranges as far as New Hampshire (DeKay, Zool. N. Y., 4, 1 842 : 350) teems not to have
had any factual basis.

Fishes of the Western North Atlantic 375

Apparently its migrations between its southern wintering and northern summering
grounds on the Atlantic coast of the United States are comparatively direct, for it has
never been reported in so great abundance anywhere along the intervening sector, although
it is occasionally taken on the coast of Delaware, and in some numbers in autumn in Chesa-
peake Bay." Although our Study Material includes one October specimen from the Vir-
ginia coast a few miles south of the entrance to the Bay,"' only occasional specimens, whose
identities are well attested, have been taken off North Carolina. Although it must pass and
repass the South Carolina coast twice yearly in its seasonal migrations, the only report
of it there is by name only."'

The status of milberti in the Gulf of Mexico and to the southward continues doubtful.
The only record of it for the Gulf is a Texan specimen (see Study Material, p. 368) ; so
far as we know the only records of it from the Caribbean or West Indies are represented
by a pair of jaws from the coast of Nicaragua,"" the head and skin of a Cuban specimen
listed above, and a nominal report from Yucatan, with no further clue to identity."^ Nor is
it likely that a shark frequenting shoal inshore waters so regularly would have been over-
looked throughout the whole of such an extensive area if it occurred there In numbers at all
rivaling those that visit the Atlantic coasts of the United States. Hence, a very small (650
mm.) specimen taken near Rio de Janeiro"^ seems more likely to have been a stray visitor
from the north"^* than a representative of a local southern Brazilian center of population}
but it is still possible that it was a representative of such a population. It Is not known from
Bermuda.

Synonyms and References:""

I. Western Atlantic:

Carcharias (Priofwdon) milberti Miiller and Henle, Plagiost., 1841: 38, pi. 19, fig. not numbered (descr.,
ill., teeth, N. York); Dumeril, Hist. Nat. Poiss., i, 1865: 360 (descr., N. York, Medit.) ; Gunther, Cat.
Fish. Brit. Mus., 8, 1870: 363, footnote 5 (ref. only).

Carcharias ceruleus DeKay, Zool. N. Y., 4., 1842: 349, pi. 61, fig. 200 (descr., ill., N. York to N. Hamp-
shire); Baird, Rep. Smithson. Instn. (1854), 9, 1855: 352, extra (N. Jersey, ident. probable because
of local abund.) ; Ann. Rep. St. Cab. nat. Hist. N. Y. (1855), 1858:64 (N. York) ; Jordan and Gilbert,
Bull. U.S. nat. Mus., 16, 1883: 873 (ident. probable from brief descr., New England to Florida);
Proc. U.S. nat. Mus., 5, 1883: 245 (Gulf of Mexico); Nelson, Rep. St. Geol. N. J., 2 (2), 1890:
660 (N. Jersey loc.).

Lamna caudato DeKay, Zool. N. Y., 4, 1842: 354, pi. 62, fig. 205 (descr., recognizable ill., Rhode Island) ;
Linsley, Amer. J. Sci., 47, 1844: 77 (Rhode IsUnd, N. York) ; DeKay, Rep. St. Cab. nat. Hist. N. Y.
(1855), 1858: 64 (name only).

97. For nuirbers caught there, see Hildebrand and Schroeder (Bull. U.S. Bur. Fish., 4$ [1], 1928: 48).

98. An earlier record of "milberti" for Virginia (Linton, Bull. U.S. Bur. Fish., 34, 1905: 341) was based on so
large a specimen (longer than nine feet) that it may have referred to some other Carcharhinus.

99. Reported by Gunter (Amer. Midi. Nat., 2* [2], 1942; 28) as entering the Cooper's River and other rivers near
Charleston, South Carolina.

100. Fowler, Proc. Acad. nat. Sci. Philad., y^, 1923: 24. 101. Bean, Bull. U.S. Bur. Fish., 8, 1890: 206.

102. Ribeiro, Fauna brasil. Peixes, 2 (i) Fasc. i, 1923: 51, pi. 3; identity established by the excellent illustrations.
102a. Or its mother may have been the stray visitor, since it cannot have been born long previous to its capture.

103. Citations for the western and eastern Atlantic are listed separately, since it is still an open question as to whether
the Mediterranean form is identical with the American.

376 Memoir Sears Foundation for Marine Research

Squalus (Carcharifius) milberti (in part) Gray, List. Fish. Brit. Mus., 1851: 45 (refs., but specimen from
India some other species) .

Squalus {Carcharinus) caeruleus Gray, List. Fish. Brit. Mus., /, 1851: 44 (ref.).

Squalus {Carcharinus) caudataGny, List. Fish. Brit. Mus., i, 1 851: 44 (ref.).

Squalus coeruleus Gill, Proc. Acad. nat. Sci. Philad., Addend., 1 861 : 59 (incl. in list) ; Abbott, Rep. N. J. geol.
econ. Surv., 1868: 828 (N. Jersey, not seen).

Squalus milberti Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861 : 59 (in list), and subsequent eds.

Eulamia milberti Gill, Ann. Lye. nat. Hist. N. Y., 7, 1862: 410 (name); Proc. Acad. Nat. Sci. Philad.,
1864: 262 (name, discuss.); Rep. U.S. Comm. Fish. (1871-72), 1873: 813; Fowler, Proc. Acad. nat.
Sci. Philad., do, 1908: 62 (N. Jersey) ; Science, N. S. 50, 1909: 81 5 (N. Jersey) ; Proc. Acad. nat. Sci.
Philad., (5j, 1911: 5 (Maryland); Copeia, 30, 1916: 36 (in list); Copeia, 31, 1916: 41 (N. Jersey);
Proc. Acad. nat. Sci. Philad., 69, 1917: 122 (Northeast R., Maryland); Proc. Acad. nat. Sci. Philad.,
71, 1920: 292 (N. Jersey); Proc. bio!. Soc. Wash., jz, 1919: 72 (Delaware); Proc. biol. Soc. Wash.,
jj, 1920: 144 (N. Jersey); Proc. Acad. nat. Sci. Philad., -ji, 1921: 386 (meas., weights, ident. perhaps
doubtful because of large size, N. Jersey); Proc. Acad. nat. Sci. Philad., 7^, 1922: 3 (N. Jersey);
Proc. Acad. nat. Sci. Philad., -j^, 1923: 24 (jaws, Caribbean coast of Nicaragua) ; Monogr. Acad. nat. Sci.
Philad., 7, 1945: 95, 263 (Beaufort, N. Carolina; Florida); Bigelow and Schroeder, Guide Comm.
Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 84, fig. 26 (descr., habits, range, ill.).

Eulamia coerulea Gill, Proc. Acad. nat. Sci. Philad., 1863: 333 (Massachusetts).

Carcharias milberti Ribeiro, Fauna brasil. Peixes, 2(1) Fasc. I, 1923: Append., 50a, pi. 3 (descr., excellent ill.,
Rio de Janeiro); Jordan, Manual Vert. Anim. NE. U.S., 1929: 10 (general); Jordan, Evermann and
Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 16 (C. Cod to Florida, Medit.).

Carcharias {Eulamia) coeruleus Jordan and Gilbert, Proc. U.S. nat. Mus., 5, 1883: 245 (ref., no loc).

Carcharinus^''^'' milberti Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 22 (ref., C. Cod to Medit.);
Smith, Bull. U.S. Fish Comm., 17, 1898: 88 (Woods Hole); Bean, Rep. For. Comm. N. Y.,
1902: 377 (Long Island, N. York); Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 26 (descr., near
N. York); Fowler, Rep. N. J. Mus. (1905), 1906: 62 (descr., N. Jersey); Tracy, Rep. R. I.
Comm. inl. Fish., 1906: 45 (ident. probable because of loc, Rhode Island); Smith, N. C. geol.
econ. Surv., 2, 1907: 34 (N. Carolina, but doubtful if 9 ft. 2 in. spec, was this species because
so large); Fowler, Rep. N. J. Mus. (1906), 1907: pi. 73 (ill.); Kendall, Occ. Pap. Boston Soc.
nat. Hist., 7 (8), 1908: 4 (south. New Eng.) ; Tracy, Rep. R. I. Comm. inl. Fish., 1910: 59
(ident. probable because of local abund., Rhode Island) ; Garman, Mem. Harv. Mus. comp. Zool.,
36, 1913: 133 (descr.); Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., 31 (2), 1913: 735 (Woods
Hole); Fowler, Copeia, 2, 1914: 2 (in list, Maryland); Copeia, 13, 1914: [2] (N. Jersey);
Nichols, Copeia, 35, 1916: 72-73 (lengths, weights, Long Island, N. York); Nichols and Murphy,
Brooklyn Mus. Sci. Bull., 5 (i), 1916: 14, pi. I (descr., breeding, habits, food, season, abund.. Long
Island, N. York); Brooklyn Mus. Quart., 3, 1916: 151 (N. York); Radcliffe, Bull. U.S. Bur. Fish.,
34, 1916: 257, pi. 39, fig. 4 (descr., denticles, teeth, N. Carolina); Rockwell, Brooklyn Mus. Quart.,
3, 1916: 162 (abund., season, harpoon fishery. Long Island, N. York); Smith, Nat. Hist. N. Y., 16,
1916: 348 (Atlant. coast, U.S.); Thorne, Copeia, 35, 1916: 69—71 (abund., season, size, weight, sex,
embryos, near N. York); Nichols, Bull. Amer. Mus. nat. Hist., 57, 1917: 874 (Woods Hole specs.);
Copeia, 53, 1918: 13 (states that majority of the sharks taken around Long Island, New York, are
female milberti) ; Meek and Hildebrand, Field Mus. Publ. Zool., 75 (l), 1923: 38 (descr., discuss., con-
cludes the Pacific azureus Jordan and Gilbert, 1882, probably identical); Breder, Copeia, 138, 1925:
2 (Sandy Hook Bay, N. York); Nichols, Copeia, 140, J925: 21 (Woods Hole); Breder, Copeia, 153,
1926: 122 (Sandy Hook Bay) ; Nichols and Breder, Zoologica, N. Y., 9, 1927: 15 (local dist., life hist.) ;
Thorne, BuU. N. Y. zool. Soc, 31, 1928: 1 14 (Long Island, N. York) ; Hildebrand and Schroeder, Bull.
U.S. Bur. Fish., 43 (i), 1928: 48 (descr., weights, lengths, numbers, Chesapeake Bay); Breder, Field
Bk. Mar. Fish. Atlant. Coast, 1929: 16 (general, size at birth); Truitt, Bean and Fowler, Bull.
Md. Conserv. Dep., 3, 1929: 29 (Maryland, from previous records); White, Bull. Amer. Mus. nat.
Hist., 14^ 1937: 126 (in Key), pi. 31, fig. a, pi. 42, fig. a, pi. 50, fig. b (spiral valve, heart valves, cartilages

103a. Also often spelled "Car char hinus."

Fishes of the Western North Atlantic 377

of claspers, New York spec.) ; Breder, Bull. N. Y. zool. Soc, 41, 1938: 28 (N. York Harbor, in list);

Springer, Proc. Fla. Acad. Sci., j, 1939: 22 (descr. denticles, abund., season, S. Florida) ; Lucey, Contr.

biol. Lab. Cath. Univ. Amer., 43, 1942: 9 (chemistry of pancreas, Florida) ; Bomkamp, Contr. biol. Lab.

Cath. Univ. Amer., 44, 1942: 4 (chemistry of liver, Florida) ; Boos, Contr. biol. Lab. Cath. Univ. Amer.,

45) '943- 10 (chemistry of pancreas, Florida); Lunz, Bull. S. C. St. Planning Bd., 14, 1944: 27 (S.

Carolina, Florida); Springer, Copeia, 1946: 1 74 (food, off Salerno, Florida).
CarcAarias flunibeus Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 872 (near N. York and Medit., by

ref.).
Carcharhinus {Eu/amia) caudatus Jordan, Rep. U.S. Coram. Fish. (1885), 1887: 796 (name, northeast. U.S.).
Carcharhinus {Carcharhinus) mMerti Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 216 (C.

Cod to Florida) ; Bull. U.S. nat. Mus., 47 (l), 1896: 37 (compar. with other sp., C. Cod to Florida).
Carcharhinu! obscurus (in part) Bean, Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 25 (abund., N. Jersey Bays,

probably refers to milberti, but not the descr.) .
Carcharhinus flatyodon Garman, Mem. Harv. Mus. comp. Zool., 5 (5, 1 91 3: pi. 3, fig. 4-6 (good ill., mis-
labeled) ; not Squalus flatyodon Poey, 1861.
Eulamia flumbeus Fowler, Bull. Amer. Mus. nat. Hist., yo (l), 1936: 51 (descr., ill. of American specs.;

states flumbeus Nardo has priority over milberti); Proc. Acad. nat. Sci. Philad., 8g, 1937: 304 (size,

N. Jersey) ; Fish Culturist, 21 (9), 1942: 66 (listed, Cuba) ; Arqu. Zool. Estado Sao Paulo, 3, 1942: 128

(listed, Brazil).

Doubtful references:

Eulamia milberti Vernll and Smith, Rep. U.S. Comm. Fish. (1871-1872), 1873: 521 (Woods Hole) ; Goode,
Proc. U.S. nat. Mus., 2, 1879: 121 (name only, Indian River, Florida, no clue to actual identity).

Carcharias obscurus Bean, T. H., Bull. U.S. Fish Comm., 7, 1888: 152 (probably milberti, because of re-
ported abund., N. Jersey).

Carcharias coeruleus Bean, T. H., Bull. U.S. Fish Comm., 8, 1 890: 206 (name only, Yucatan).

Carcharhinus milberti Yverminn and Bean, Rep. U.S. Comm. Fish. (1896), 1898: 239 (nominal, Indian R.,
Florida, by ref. to earlier report by Goode, 1879); Evermann and Kendall, Rep. U.S. Comm. Fish.
(1898), 1900: 48 (nominal, Indian R., Florida); Linton, Bull. U.S. Bur. Fish., rp, 1901: 426 (name
only, food, parasites. Woods Hole); Bull. U.S. Bur. Fish., 24, 1905: 341 (ident. doubtful because of
large size, Virginia) ; Wilson, Proc. U.S. nat. Mus., ^j, 1907: 360 (name only, parasites. Woods Hole) ;
Linton, Bull. U.S. Fish Comm., 28, 1910: l 200 (name only, parasites, no loc.) ; Proc. U.S. nat. Mus., 64
(21), 1924: 3, 5, 6, 12, 33, 36, 38, 40, 46, 56, 87, go, 92 (name only, parasites. Woods Hole) ; Wilson,
Proc. U.S. nat. Mus., 64 (17), 1924: 11, 12 (name only, parasites. Woods Hole); MacCallum, Proc.
U.S. nat. Mus., /p (26), 1931: 7 (name only, parasites, no loc); Wilson, Bull. U.S. nat. Mus., 158,
1932: 427, 434, 440, 453, 464, 524 (name only, parasites. Woods Hole region) ; Nigrelli, Amer. Mus.
Novit., 996, 1938: 10 (name only, parasites, no loc); Gunter, Amer. Midi. Nat., 28 (2), 1942: 316
(name only, rivers of S. Carolina but not into fresh water).

Carcharinus milberti Norns, Plagiost. Hypophysis, 1 941: 28.

Carcharinus %'p. (probably C. milberti), Norris, Plagiost. Hypophysis, 1941: pi. 9, fig. 33, 34 (brain).

2. Eastern Atlantic:

Squalus flumbeus Nardo, in Oken's Isis, 20 (6), 1827: 483; also sep., as Prod. Ittiol. Adriat., 1827: 9 (brief

diagn., Adriatic) ;'^°* Sinon. Modern. Spec, descr. Pesci. St. Chiereghin, 1847: 1 1 1 (name, Medit.).
Squalus caecchia'iizTdo., Prod. Faun. Mar. Venet. Estuario, 1847: 38 (near Venice, not seen).
Squalus, sp. n., Molin, Revist. I. R. Acad. Padova, Trim. 3-4, 1853: 381 (ref. to Squalus milberti by Nardo,

1853; not seen).

104. Nardo's statement that his flumbeus "convenit perfect] Squalus glaucus Bloch si colorem excipetur et forman
rostri quae in exemplari nostro rotunda est" would apply equally to any round-snouted Carcharhinus. But the
fact that Nardo (Pesc. Publ. com. Nuov. Venez., 1853: 15) later referred his flumbeus to milberti justifies
Fowler's (Bull. Amer. Mus. nat. Hist., jo [1], 1936: 50) substitution of flumbeus for milberti as the correct
specific name, at least of the European form and of the American as well if the two prove to be identical.

378 Memoir Sears Foundation for Marine Research

Squalus milberti Bonaparte, Cat. Pesc. Europ., 1846: 18 (incl. in list) ; Nardo, Pesc. Publ. com. Nuov. Venez.,

1853: 15 (brief diagn., in lagoons and canals, Venice); Ninni, Ann. Soc. Nat. Modena, 5, 1870: 66

(nominal, Medit. near Naples).
Prionodon milberti Canestrini, in Cornalia, et at.. Fauna d'ltal., 1872: 48 (descr., ref., Adriatic); Doder-

lein, Prosp. Metod. Pesc. Sicil., 7; Atti Acad. Palermo, N. S. 6, 1878-1879: 30 (in Adriatic list);

Graeffe, Arb. zool. Inst. Univ. Wien, 28, 1886: 446 (Medit.).
Carcharias caecchia Ninni, Revist. Critica Pesc. Adriat. descr. St. Chiereghin, 1872: 6—7 (name, Adriatic,

not seen).
Carcharias (Prionodon) milberti Doderlein, Man. Ittiol. Medit., 2, 1 881: 44 (descr., refs., Medit.);'"'

Bellotti, Atti Soc. ital. Sci. nat., 55, 1891: 112 (discuss.).
Carcharias milberti Moreau, Hist. Nat. Poiss. France, /, 1881 : 335 (ref., Medit.) ; Perugia, Elenc. Pesc. Adriat.,

1881: 52 (in list); Carus, Prod. Faune Medit., 3, J889-1893: 512 (Iocs., Medit.); Moreau, Poiss.

France, Suppl., 1 891 : 2 (descr. of types from N. York, and meas. Medit. embryo, but ill. of teeth of some

other species, if correctly drawn); Sicher, Atti Accad. gioenia, (4) ti (5), 1898: 14 (Medit.);

Pietschmann, Ann. naturh. (Mus.) Hofmus. Wien, 2t, 1906: 133 (descr., discuss., south. Medit.).
Prionace milberti Roule, Bull. Inst. Oceanogr. Monaco, 243, 1912: 7 (nominal, C. Verde).
Carcharhinus milberti Rey, Fauna Iberica Peces, r, 1928: 346 (descr., S. coast of Spain) ; Bellon and Mateu,

Notas Inst. esp. Oceanogr., (2) ^^, 1931: 15 (nominal, Canaries); Borri, Mem. Soc. tosc. Sci. nat., 44,

1934: 88 (Medit.).
Carcharhinus -plumbeus Tortonese, Atti Soc. ital. Sci. nat., yj, 1938: 299 (discuss, specs, from Medit.).

Carcharhinus nicaraguensis (Gill and Bransford), 1877

Lake Nicaragua Shark

Figure 70

Study Material. Immature male, about 1,511 mm. long, from San Carlos, Lake
Nicaragua, caught by Maj. C. M. Duke, U.S.A. (Harv. Mus. Comp. Zool., No. 35896) }
photographs of two females, fresh-caught, by President Don Anastasio Somoza of Nicara-
gua and Capt.W. B. Bunker 5 three males of about 1,330 mm., 1,568 mm. and 1,710 mm.,
from Lake Nicaragua (U.S. Nat. Mus., No. 1 20371, 120372, 120373).

Distinctive Characters. Nicaraguensis very closely resembles leucas, of which it is a
landlocked representative, but is perhaps separable from leucas by the following charac-
ters. The anterior margin of the eye is posterior to the front of the mouth by a distance
equal to half its own diameter in nicaraguensis (a little anterior to front of mouth in
leucas) ; the gill openings are relatively somewhat longer in nicaraguensis, the third being
nearly % as long as the distance between the nostrils (in leucas the third is a little less than
half that long) ; the free tip of the second dorsal is about two-thirds as long as its base in
nicaraguensis (only about half that long in leucas).

Description. Proportional dimensions in per cent of total length. Male, 1,511 mm.,
from Lake Nicaragua (Harv. Mus. Comp. Zool., No. 35896). Male, 1,568 mm., from
Lake Nicaragua (U.S. Nat. Mus., No. 120372).

Trunk at origin of -pectoral: breadth 14.0, 15.3; height 14.O, 15.3.
Snout length in front of: outer nostrils 2.3, 3.1 ; mouth 6.0, 6.9.
Eye: horizontal diameter i.i, i.o.

105. See Doderlein for additional Mediterranean citations in publications not accessible to us.

Fishes of the Western North Atlantic

379

Mouth: breadth 9.5, 10.8} height 5.0, 4.9.

Nostrils: distance between inner ends 6.3, 6.9.

Labial furrow length: upper 0.3, 0.3.

Gill opening lengths: ist 4.1, 3.3} 2nd 4.2, 3.6; 3rd 3.8, 4.0; 4th 3.7, 3.7; 5th

2.7,3.1.

First dorsal fin: vertical height 12.7, 12.3; length of base 13.2, 12.2.

Second dorsal fin: vertical height 4.9, 4.7; length of base 5.9, 6.4.

Anal fin: vertical height 5.3, 5.4; length of base 6.0, 5.3.

Caudal fin: upper margin 27.1, 29.6} lower anterior margin 13.2, 13.6.

Pectoral fin: outer margin 21.9, 23.6; inner margin 6.4, 6.S'-, distal margin 19.6,

20.4.

Distance from snout to : ist dorsal 27.6, 27.95 2nd dorsal 61.0, 56.6; upper caudal

72.9, 70.4} pectoral 22.3, 20.4; pelvics 47.7, 44-9} anal 61.4, 58.1.

Inters-pace between: ist and 2nd dorsals 21.8, 19.95 2nd dorsal and caudal 7.0,

7.9} anal and caudal 5.8, 5.9.

Distance from origin to origin of: pectoral and pelvics 28.6, 28.O; pelvics and anal

14.1, 12.2.

UM>AAAAA=^yyu.

Figure 70. Carcharhinus nicaraguensis, immature male, 1,511 mm. long (Harv. Mus. Comp. Zool., No.
35896). A Anterior part of head from below, about Yj natural size. B Right-hand nostril, about 0.8 x.
C Dermal denticles, about ij x. D Apical view of dermal denticle, about 1 7 x. £ Upper and lower teeth, left-
hand side, about 0.6 x natural size. F Third upper tooth. G Ninth upper tooth. H Second lower tooth. / Eighth
lower tooth. F—I, about 1.2 x.

380 Memoir Sears Foundation for Marine Research

Trunk moderately stout, as in leucas, its height at ist dorsal about Vr> its length to
origin of caudal. Back without trace of median dermal ridge. Upper precaudal pit strongly
marked, semi-lunar, the lower smaller but still apparent. Dermal denticles overlapping
in varying degrees, with skin more or less exposed between their blades, thick, strongly
arched and rising steeply, about as broad as long, usually with 3-5 high, sharp-topped
ridges, 3, 4, or 5 blunt teeth, the median much the longest 5 pedicels verv broad and stout
as in leucas.

Head strongly flattened above and very wide anteriorly, its breadth about % as great
at eyes as at origin of pectoral. Snout very broadly rounded and shorter than in any other
local Carcharhinus, its length in front of a line connecting outer ends of nostrils only a
little more than Vz as great as distance between inner ends of latter, the length in front of
mouth about V4 to V3 (27 to 33%) that of head to origin of pectoral. Eye noticeably
small as in leucas, its diameter about '^/o as great as distance between nostrils. Nostril
strongly oblique, its inner end only a little more than V2 as far from mouth as from tip of
snout, its anterior margin expanded toward inner end in subrectangular outline, much as in
leucas. Gill openings about evenly spaced, the ist to 3rd between V2 and % as long as dis-
tance between nostrils or about 3V2 times as long as diameter of eye, the 5th about % as
long as 1st, the 3rd over origin of pectoral. Mouth broadly rounded, its height about V2
its breadth. Upper labial fold so short as to be easily overlooked.

Teeth '^ "'' '^^Y— 12'^ ""^ '^ > shaped much as in leucas; uppers broadly triangular, their
edges moderately serrate from tip to base, the ist and 2nd teeth symmetrical, erect, their
edges slightly concave, but successive teeth increasingly oblique along the jaw with outer
edges increasingly concave in rounded or subangular contour, the 9th to 13th with inner
edge convex, and successively smaller, the outermost very small ; lower teeth erect, sym-
metrical, with narrow triangular cusps on broadly expanded bases, the cusps finely serrate
and bases partially so, the 9th to 12th successively smaller, with relatively lower cuspj a
very small symmetrical tooth at symphysis on lower jaw, i present or not in upper."*

First dorsal of about same size relatively as in leucas, with a similar subacute apex,
but somewhat more erect in specimens seen, and with a relatively somewhat longer free
rear corner (about Vs as long as the base), its origin a little anterior to axil of pectoral.
Second dorsal about Yi as long at base as ist, as in leucas, and of the same general form
except that the free rear corner is about 73 as long as base (only about V2 that long in
leucas), and the rear margin somewhat more deeply concave, its origin slightly but defi-
nitely anterior to origin of anal. Caudal about 28% of total length, of same shape as in
leucas. Distance from lower origin of caudal to tip of anal about V2 as long as base of latter.
Anal about as long at base as 2nd dorsal, and nearly i V2 times as long on anterior margin,
its shape as in leucas. Distance from origin of anal to tips of pelvics about V2 as long as base
of former. Pelvics a little longer along anterior margins than 2nd dorsal and a little shorter

106. This tooth is recorded for the type specimen (Gill and Bransford, Proc. Acad. nat. Sci. Philad., 1877: 190) but
is lacking in one which we have studied.

Fishes of the Western North Atlantic 381

than anal, as in leucas. Pectoral very nearly as long as head and thus a little longer than
in leucas, but of same general shape, a little more than Yi as broad as long, the outer
margin nearly straight except near tip, the distal margin a little more deeply concave
toward base than in leucas.

Color. Dark mouse-gray above after preservation in salt, grayish to yellowish white
below, with lower surfaces of pectorals dusky at tipsj photographs at time of capture show
the lower surface a clearer white, the lower pectoral tips and tip of lower caudal dusky or
nearly black. It is said that the bellies of large specimens may have a reddish bronze tinge.

Size. The fact that the specimens we have seen are immature, although up to 5
feet long, is in line with information reaching us from Nicaragua that the average size of
those caught around San Carlos is 6 to 6V2 feet. We have a definite report of one of 8
feet, and they are rumored to reach 10 feet. One of 4 feet is reported as weighing about
50 pounds. Specimens of 62 and 67 inches (1,568 mm. and 1,710 mm.) in the United
States National Museum weighed 73 pounds and 98 pounds respectively when caught.

Develof mental Stages. Embryos have not been seen.

Remarks. Nicaraguensis was classed by Garman,^"' and more recently by Meek and
Hildebrand,'"' as a synonym of milherti, in spite of the fact that firsthand accounts had
credited it with a much shorter snout. Actually it is so close to leucas that it is undoubtedly
an offshoot of the latter. But the several small differences, enumerated above, seem suffi-
cient for retention of a separate name for it, especially since it is the only shark that is
known to have adapted itself permanently to life in fresh water. Nor is it astonishing that
this should have happened, for leucas has been reported in fresh water far up rivers else-
where (p. 341).

Habits. Very little is known of the habits of this fresh-water shark, except that it
comes commonly into very shoal water although it is seldom actually seen at the surface
and that it bites very readily on bait of meat or fish. Presumably it feeds on fish, but no
precise information is available as to its diet."° Nothing is known about its breeding habits.

Relation to Man. The fins are valued locally for food, and the livers are sold for
their vitamin content.

It is reputedly a danger to bathers, as well as to any dog that may venture into the
lake. And published accounts of its ferocity appear to be well founded, for a correspondent
in whom we have full confidence"" reports that he has not only seen an attack on a youth
swimming at San Carlos but has heard of actual fatalities at other localities around the lake.
Very recently the press has reported attacks on bathers and fishermen at Granada, where
one of the victims lost an arm, while another lost his right leg and had his left leg injured."^

Range. Known definitely only from Lake Nicaragua, its tributaries and outlet."^

107. Mem. Harv. Mus. comp. ZooL, jd, 1913: 133. 108. Field Mus. Publ. Zool., 15 (i), 1923: 38.

109. For an account of the fishes of Lake Nicaragua, see Meek (Field Mus. Publ. Zool., 7 [4], 1907: 95-132).

1 10. Capt. W. B. Bunker.

111. In the Diario Nuevo, San Salvador, for April 24, 1944.

112. Jordan, Evermann and Clark (Rep. U.S. Comm. Fish. [1928], 2, 1930: 16) include "Bay of Panama" in its

382 Memoir Sears Foundation for Marine Research

It would be interesting to know whether a shark of this general type, and of which
we received a photograph, recently taken in the Patuca River, Honduras, in fresh water
was this same landlocked species, or whether it was simply a leucas that had run far up-
stream, which we think more likely (see discussion, p. 341).

Recent contributors who have caught sharks in Lake Nicaragua'" report them as plen-
tiful there as well as in the upper reaches of the San Juan River, by which the lake dis-
charges into the Caribbean. It is not known how far downstream its range extends in the
San Juan, but since there are heavy rapids in the river it seems likely that the sharks from
the lake and upper river are entirely landlocked. According to local reports it also runs
at least 40 miles up the Rio Frio, which empties into the lake close to where the latter
discharges via the San Juan River. Rumors of its presence in Lake Managua appear to lack
foundation. The fact that the specimen shown in Fig. 70 was caught from the pier after
only 1 5 minutes fishing is some indication of the abundance of these sharks, at San Carlos
at any rate."*

Synonyms and References:

Eulamia nicaraguensis Gill and Bransford, Proc. Acad. nat. Sci. Philad., 1877: 190 (descr., Lake Nicaragua) ;
Eigenmann, Proc. U.S. nat. Mus., 16, 1893: 54 (name only, San Juan R., Nicaragua) ; Marden, Nat.
Geogr. Mag., S6, 1944: 183 (fatalities, capture of specs., excellent photos, L. Nicaragua).

Corcharias nicaraguensis Liitken, Vidensk. Medd. naturh. Foren. Kbh., 1879-1880: 65 (ill., meas., descr.,
San Juan R., Nicaragua); Regan, Biol. Cent. Amer. Pisces, 1906-1908: 183 (size. Lake Nicaragua
and San Juan R.) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 16 (Lake
Nicaragua, San Juan R., Panama Bay, but see footnote 112, p. 380).

Carcharhinus nicaraguensis Jordan, Proc. U.S. nat. Mus., p, 1887: 556 (name only in list. Cent. Amer.);
Meek, Field Mus. Publ. Zool., 7, 1907: 103 (local abund., Lake Nicaragua and San Juan R.) ; Eigen-
mann, Rep. Princeton Exped. Patagonia (1896— 1899), 5 (4), 1910:377 (listed) ; Smith, H. W., Biol.
Rev., //, 1936: 64 (fresh water) ; Norman and Fraser, Giant Fishes, 1937: 36 (confined to fresh water).

Carcharhinus {Carcharhinus) nicaraguensis Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 216
(Lake Nicaragua and San Juan R.) ; Bull. U.S. nat. Mus., 47 (l), 1896: 39 (descr., Lake Nicaragua
and San Juan R.).

Carcharinus milberti (in part) Garman, Mem. Harv. Mus. comp. Zool., ^6, 1913: 133 (referred to synonymy
of milberti); Meek and Hildebrand, Field Mus. Publ. Zool., 75, 1 923: 38 (referred to synonymy of
milberti); not Carcharias {Prionodon) milberti Miiller and Henle, I 84 1.

Carcharhinus obscurus Lesueur, 1 8 1 8

Dusky Shark, Shovelnose

Figures 71, 72

Study Material. One female, 996 mm., and 5 males, 993 to 1,560 mm., from the
vicinity of Woods Hole (Harv. Mus. Comp. Zool.); 8 males, 970 to 1,500 mm. long,

range, but seemingly not on any factual basis; nor would it be expected there even as a stray, for Lake Nica-
ragua drains into the Atlantic and not into the Pacific.

1 13. We have received several personal communications on this subject.

114. Smith's (Science, 22, 1893: 166) statement that unnamed sharks are abundant in Lake Nicaragua no doubt
refers to this species.

Fishes of the Western North Atlantic

383

from the vicinity of Woods Hole, not preserved; also measurements of 5 adult females,
3,1 15 to 3,465 mm., from Englewood, Florida,"' one of which was gravid; and a new-
born specimen, 848 mm., taken off Bay Chaland, Louisiana, in August 1930 (U.S. Bur.
Fish., No. 1360, in U.S. Nat. Mus.).

! B

Figure 71. Carcharhinui obscurus, female, 996 mm. long, from Woods Hole, Massachusetts (Harv. Mus.
Comp. Zool., No. 35312). A Right-hand pectoral fin. B Dermal denticles, about 16 x. C Upper and lower
teeth, left-hand side, 1.2 x. Z) Third upper tooth. E Ninth upper tooth. F Fourth lower tooth. G Tenth
lower tooth. D-G, about 2.6 x.

Figure 72. Corckarhinus obscurus, pictured in Fig. 71.
A Anterior part of head from below, about 0.4 x natural
size. B Left-hand nostril, about 2.3 x.

1 15. Contributed by Stewart Springer.

384 Memoir Sears Foundation for Marine Research

Distinctive Characters. Among ridge-backed members of the genus, obscurus has
frequently been confused with milberti. But It is easily separable from the latter by the
following characters: the first dorsal is somewhat less in vertical height than the distance
from the eye to the first gill opening and it originates over or a little posterior to the inner
corner of the pectoral (much larger and originating over the axil in milberti) ; the free
rear corner of its second dorsal is about 1.5 times as long as the vertical height (only about
as long as the vertical height in milberti) ; its dermal denticles are regularly overlapping
and with toothed margins. It is easily separable from the other ridge-backed species jalcp-
formis and floridanus by the shape of the second dorsal and anal fins (see Key, p. 323) j
from longim.inus by the shape of the first dorsal and by the remoteness of the tip of its
anal from the caudal.

It most closely resembles the newly described springeri, Bigelow and Schroeder, 1944
(p. 404), but differs from it in a number of respects, most noticeably: (a) in a consider-
ably smaller eye relative to the lengths of the gill openings; (b) that the anterior margin
of its nostril is not definitely lobed; (c) the distance from the tips of its pelvics to the origin
of its anal is relatively longer (about 1.3 times as long as the base of the anal in obscurus,
but only about 0.7 that long in springeri) ; (d) the outer corner of its pelvic is considerably
more obtuse than a right angle, or about 115° (only about a right angle in springeri) ; and
(e) its denticles usually have fewer teeth than those of springeri at corresponding stages
in growth. The teeth of the two species also differ somewhat in detail (cf. Fig. 71 C-G
with78E-I).

Description. Proportional dimensions in per cent of total length. Male, 993 mm.,
from Buzzards Bay, Mass. (Harv. Mus. Comp. ZooL, No. 35371). Female, 996 mm.,
same locality (Harv. Mus. Comp. ZooL, No. 35312).

Trunk at origin of pectoral: hrea.dth. 11.9, 13.7; height 13.3, 12.9.

Snout length in front of: outer nostrils 3.5, 4.O; mouth 7.9, 7.9.

Eye: horizontal diameter 1.9, 1.7.

Mouth: breadth 7.9, 9.2; height 4.7, 4.9.

Nostrils: distance between inner ends 6.2, 6.3.

Gill opening lengths: ist 2.6, 2.93 2nd 2.8, 3.4; 3rd 3.2, 3.5; 4th 3.2, 3.5; 5th

2.7,2.9.

First dorsal fin: vertical height 7.6, 8.6; length of base 9.8, 9.6.

Second dorsal fin: vertical height 2.3, 2.6; length of base 4.0, 3.5.

Anal fin: vertical height 3.1, 3.4; length of base 3.9, 4.5.

Caw^a/^w.-upper margin 27.4, 28.8; lower anterior margin 11. 2, 11.4.

Pectoral fin: outer margin 17.4, 18.5; inner margin 5.0, 5.2; distal margin 13.O,

14.I.

Distance from snout to: 1st dorsal 30.7, 32.O; 2nd dorsal 62.2, 61. 2; upper caudal

72.6, 71.2; pectoral 22.6, 23.7; pelvics 50.2, 48.5; anal 62.6, 61.2.

Interspace between: ist and 2nd dorsal 22.7, 21.2; 2nd dorsal and caudal 6.4, 6.1 ;

anal and caudal 6.1, 5.9.

Fishes of the Western North Atlantic 385

Distance from origin to origin of: pectoral and pelvics 27.8, 26. i ; pelvics and anal

12.7, 12.8.
Proportional dimensions in per cent of total length. Averages of 3 females, 3,195 to
3,465 mm., from Englewood, Florida (calculated from measurements by Stewart
Springer) .

Snout length in front of: mouth 6.3.

Eye: horizontal diameter i.O.

Mouth: breadth 10.6; height 4.7.

Nostrils: distance between inner ends 6.2.

Gill opening lengths: ist 3.4; 5th 2.7.

First dorsal fin: length of base 9.2.

Second dorsal fin: length of base 2.8.

Anal fin: length of base 3.9.

Caudal fin: upper margin 26.55 lower anterior margin 12.0.

Pectoral fin: outer margin 20.6} inner margin 5.4; distal margin 18.6.

Distance from snout to: ist dorsal 32.95 upper caudal 73.55 pectoral 23.6.

Interspace between: ist and 2nd dorsal 23 95 2nd dorsal and caudal 6.1.

Trunk more slender than in milherti, its height at ist dorsal about y^ its length to
origin of caudal. A low but unmistakable dermal ridge along midline of back between the
2 dorsal fins. Body sector to cloaca about 1.2 times as long as tail sector. Upper precaudal
pit deep, subrectangular, the lower less deeply marked than upper. Dermal denticles over-
lapping so regularly that skin is entirely concealed, their blades but little arched longitu-
dinally, 3 to 5 low ridges in small specimens but perhaps most often 7 in adults, with a
corresponding number of strong pointed marginal teeth, the median considerably the
largest and the outermost pair very small when there are more than 5.

Head to 5th gill opening about V4 of total length, its width nearly or quite 80% as
great at eyes as at origin of pectorals. Snout moderately thick anteriorly, broadly rounded,
its length in front of a line connecting outer ends of nostrils about % as great as distance
between inner ends of latter, and length in front of mouth about Vs as great as that of head.
Eye circular, its diameter only a little more than V2 (50 to 60%) as long as 3rd gill open-
ing, or a little more than V4 (27%) as great as distance between nostrils in young speci-
mens and still smaller relatively in adults; its anterior margin about opposite front of
mouth. Gill openings about evenly spaced, slightly oblique and weakly concave in outline,
the 2nd to 4th (longest) a little less than V2 (about 46%) as long as distance between
nostrils or about twice as long as diameter of eye, the ist and 5th about 0.8 to 0.9 as long
as 2nd to 4th, the 4th over margin of pectoral. Nostril strongly oblique, its inner end about
% as far from mouth as from tip of snout, its anterior margin slightly sinuous only without
definite expansion. Mouth broadly ovate, about twice as broad as high, occupying a little
less than % (about 72%) of breadth of head. Upper labial furrow only about 0.3 times as
long as diameter of eye and almost or entirely concealed when mouth is closed.

386 Memoir Sears Foundation for Marine Research

Teeth '■*°'- j^'^|°^— j^^'^ ^ upper teeth broadly triangular, their edges serrate, most
coarsely so toward base, the ist two erect, symmetrical, with nearly straight or very slightly
concave edges, but subsequent teeth weakly oblique, their outer margins considerably
more deeply concave in subangular contour, the ist to 9th or loth about equal in size, the
lOth to 14th or 15th successively smaller and more deeply notched, the outermost i or 2
very short; lower teeth erect, symmetrical, with narrow triangular cusps on broadly ex-
panded bases, serrate from tip to base but more finely so than uppers, the 2nd to 6th or 7th
a little the largest, the outermost 2 or 3 much the smallest; i to 3 small teeth at symphysis
in each jaw.

First dorsal much smaller than in milberti, leucas or longimanus, its base appreciably
shorter than from eye to ist gill opening, its origin about over inner corner of pectoral or a
little posterior to latter, its anterior margin moderately convex toward tip, its posterior
margin much more deeply concave than in milberti or in springeri, its apex subacute, its
free rear corner slender and nearly V2 as long as base, the midpoint of its base about 1.6
times as far from origin of pelvics as from axils of pectorals. Second dorsal only about V3
as long at base as ist and relatively much lower, its free rear tip rather slender and about as
long as base or about i V2 times as long as the vertical height, its extreme length from origin
to tip about 2.5 its vertical height and therefore considerably greater relatively than in
s-pringeri; its origin over origin of anal or a little anterior to it. Caudal between Yz and V4
of total length (27 to 29%), its terminal sector a little less than % the length of the fin
and noticeably slender with narrowly rounded tip and moderately concave lower posterior
outline, the lower lobe (expanded lower anterior corner) about Vs to % (about 40%) as
long as upper and thus somewhat shorter relatively than in springeri, with moderately and
evenly convex anterior margin, nearly straight posterior margin and very narrowly
rounded or subacute tipj the re-entrant corner (included by the 2 lobes) moderately
rounded. Distance from origin of caudal to tip of anal about % as long as base of latter,
or about as long as its free rear corner. Anal about 1.2 times as long at base as 2nd dorsal
and almost 1.5 times as long on anterior margin but only a little higher vertically, its pos-
terior margin much more deeply incised, its free rear corner between % and % as long as
base, its tip a little anterior to tip of 2nd dorsal. Distance from origin of anal to tips of pel-
vics about 1.3 times as long as base of anal. Pelvics a little longer at base than anal, their
outer corners considerably more obtuse than a right angle (about 115°). Pectoral about
%o (78 to 91%) as long as head, a little less than V2 as broad as long, with subacute or
very narrowly rounded corners, usually with only weakly convex outer margin and
weakly concave distal margin, as is the right-hand fin on specimen illustrated, but some-
times with much more strongly convex outer edge and concave distal edge, as is the left-
hand fin on this same specimen (Fig. 71)."'

Color. Back and upper sides, including upper surfaces of pectorals, bluish or leaden

116. A striking example of variability in this genus, illustrating the danger of basing specific distinctions on small
diflEerences in the shape of a given fin.

Fishes of the Western North Atlantic 387

gray in the fresh-caught specimens we have seen, but also described as dirty gray, or very
pale, perhaps as a result of living over a white sand bottom; lower parts white; lower sur-
faces of pectorals grayish and sooty toward tips; pelvics and anal grayish white.

Size. The fact that embryos up to 965 mm. have been reported, as well as a free-
living specimen hardly larger (see Study Material, p. 382), suggests that this is about the
usual size at birth. Although it is generally recognized that this is a considerably larger
shark than milberti, the only exact length records of large adult specimens, identified be-
yond question as this species, are nine females from southern Florida that ranged from lO
feet 4 inches to 1 1 feet 8 inches'" and one specimen from Georges Bank 1 1 feet long."*
It is reputed to reach 14 feet, although perhaps not from any exact evidence.

Developnental Stages. Embryos have not been described; ten have been recorded in
one Florida female and embryos up to 965 mm. long in another.

Habits. Although obscurus has been known to science since 1818 and is common
enough to be caught occasionally close to Woods Hole and reputedly more often near
New York, our only information regarding its diet is that it is a fish-eater; off the east
coast of Florida portions of other sharks have been found in its stomach as well as various
reef fishes, such as groupers, lizardfish {T rachinocefhalus) , flatfishes {Citharichthys)^
red goatfish {Mullus) and cusk eels (Ophidian). The wide distribution of the locali-
ties where positively identified specimens of obscurus have been taken show it to be much
more pelagic in habit than are either milberti (p. 372) or leucas (p. 341). On the other
hand, the record of captures proves that it comes closer inshore, even into very shoal water,
than does longimanus (p. 359). All captures in the northern part of its range have been
for the warm months, whereas it is present the year round along eastern Florida but only
in the winter off southwestern Florida, which is evidence that some obscurus migrate north-
ward along the United States coast in spring or early summer, as do various other warm-
water sharks.

All that is definitely known of its breeding habits, beyond what is stated above, is that
adult females containing embryos of 575 to 965 mm. are reliably reported off southwestern
Florida, and that free-living specimens, so small that they had evidently been born only
shortly previous, have been taken off southeastern Florida in late winter,"* and off Louisi-
ana and near Woods Hole in August (see Study Material, p. 382). It seems, therefore,
that obscurus may produce young anywhere within its geographic range and over a long
season.

Relation to Man. Obscurus is not taken in large enough numbers to be of any commer-
cial importance anywhere, although such as are caught in the shark fishery in southern
Florida or in the West Indian region are utilized for leather, etc., as in the case of other
large sharks.""

117. Springer, Proc. Fla. Acad. Sci., 5, 1939: 25.

118. Firth, Bull. Boston Soc nat. Hist., 61, 1931 : 9; Bigelow and Schroeder, Bull. U.S. Bur. Fish., 48, 1936: 321.

119. Personal communication from Stewart Springer.

120. It seems more likely from the context that a recent account of the actions of a Dusky Shark, when hooked on
rod and reel (Wise, Tigers of the Sea, 1937: 262), may have referred to milberti.

388 Memoir Sears Foundation for Marine Research

Range. Tropical and warm-temperate waters on both sides of the Atlantic. In the
west obscurus is reliably recorded from southern Massachusetts and Georges Bank to
southern Florida, Louisiana, and the Bahamas; and as far south as southern Brazil by name.
In the eastern Atlantic reports apparently referable to obscurus include the Mediterranean
coast of Spain,"* Madeira, Senegal, the Canaries and Cape Verdes, as well as the vicinity
of Sierra Leone, Ascension Island, St. Helena, and Table Bay, South Africa.*" But final
decision as to whether or not the "obscurus" of the two sides of the Atlantic are identical
must await comparison of specimens from the two ocean areas.

Occurrence in the Western Atlantic. Obscurus has been characterized repeatedly in
the past as common or even very common along the coast of the United States from
New Jersey to Cape Cod. However, it has been proved recently that most of these reports
were actually based upon C. milberti (p. 374) or on leucas in some instances, and that
North Carolina records of obscurus similarly refer either to limbatus or to Negafrion
brevirostris.^^" In fact, it is only around southern Florida that positively identified speci-
mens of obscurus have been taken recently in any numbers. Along southwestern Florida
it is present in winter, while off the southeastern coast it is common throughout the year.
Also, it has been taken off the coast of Louisiana (one specimen, see Study Material, p.
383), but is not reported otherwise from the western waters of the Gulf of Mexico. To
the northward its distribution presents a puzzling picture, for we find no reliable record of
it anywhere along the coast between Florida and Delaware Bay. But it has been taken off
the mouth of Delaware Bay and repeatedly on the coast of New Jersey, at Long Island,
New York and at Woods Hole, where twelve specimens have come into our hands in recent
summers, six of them during August 1944 (see Study Material, p. 383) in addition to
others reported in earlier years. There is at least one record for Nantucket and another for
Georges Bank, which, while by name only, seem referable to this species and not to milberti
because of the large sizes of the specimens in question (11 to 12 feet). However, these
last appear to be the most northerly and easterly of the reliable records of the species on
this side of the Atlantic, for while obscurus has been reported by name at three localities
in the Gulf of Maine,"* at least one of these records"" was almost certainly based on
Prionace glauca (p. 282), and the others probably were the same. Also, reliable reports
from New Jersey northward rest on odd specimens only, showing that the numbers of in-
dividuals that visit any part of the coast north of Florida are very small as compared with
milberti, although printed references to "obscurus" for Long Island and for southern New
England would suggest the reverse. Its recorded appearances in the northern part of its
range are limited to the warm months, chiefly August and September.

Information on its occurrence south of Florida is even more scanty, i.e., nominal

121. Carcharhtnus commersonii Rey (Fauna Iberica Feces, r, 1928 : 342), identifiable as obscurus by the presence of a
mid-dorsal ridge, position of first dorsal and teeth.

122. A stuffed specimen in South African Museum (Barnard, Ann. S. Air. Mus., 31 [i], 1925: 26).

123. Radcliffe, Bull. U.S. Bur. Fish., 5.^, 1916: 255, 257.

124. Bigelow and Welsh, Bull. U.S. Bur. Fish., ^o (i), 1925: 30.

125. Storer, Mem. Amer. Acad. Arts Sci., N. S. 9, 1867: 219, pi. 36, fig. 2.

Fishes of the Western North Atlantic 389

records for the Bahamas (hence perhaps not actually based on that species at all), Trinidad,
British Guiana and Brazil. It is also reported as taken well offshore around Bermuda, as
is to be expected from Its pelagic nature.

Synonyms and References:*"

Squalus obscurus Lesueur, J. Acad. nat. Sci. Philad., i, 1818: 223, pi. g (descr., good ill., N. Amer. but no

precise loc.) ; Gill, Proc. Acad. nat. Sci. Philad., Addend., 1 861 : 59 (ref. to Lesueur, in list, N. Amer.).
Carcharias jalci-pinnis Lowe, Proc. zool. See. Lond., 7, 1839: 90; Proc. zool. Soc. Lond., 11, 1848:

93; Trans, zool. Soc. Lond., j, 1849: 18 (specs, later ident. by Gunther, 1870, as obscurus, Madeira).
Carcharias {Prionodon) obscurus Muller and Henle, Plagiost., 1 841: 46 (ref. to Lesueur, N. Amer.); Du-

meril, Hist. Nat. Poiss., r, 1865: 371 (descr., from Lesueur, i8i8, N. Amer.); Gunther, Cat.

Fish. Brit. Mus., 8, 1870: 366 (descr., specs., Madeira, St. Helena) ; Metzelaar, Trop. Atlant. Visschen,

1919: 187 (refs., both coasts N. Atlant.).
Carcharias obscurus Bory de St. Vincent, Diet. Class. Hist. Nat., 75, 1829: 597 (ref. to Lesueur, 1 8 1 8) ; Storer,

Mass. Zool. Bot. Surv., Rep. on Fish., 1839: 184 (ref. to Lesueur only) ; Boston J. nat. Hist., 2, 1839:

533 (descr. after Lesueur, but teeth from Mass. Bay more likely Prionace glauca) ; DeKay, Zool. N. Y.,

4, 1842: 350, pi. 61, fig. 201 (descr., ill. after Lesueur, no loc.) ; Rep. St. Cab. nat. Hist. N. Y. (1855),
8, 1858: 64 (listed for N. York); Mellis, "St. Helena Pisces," 1875 (not seen); Gunther, "Chal-
lenger" Rep., Zool., I (6), 1880: 5 (name only, near St. Helena); Jordan and Gilbert, Bull. U.S.
nat. Mus., 16, 1883: 872 (ref. to Lesueur); Proc. U.S. nat. Mus., 5, 1883: 244 (coast of U.S.);
Bean, T. H., Rep. U.S. Comm. Fish. (1882), 1884: 343 (Woods Hole) ; Gunther, "Challenger" Rep.,
Zool., 31 (2), 1889: 5 (name only, near Sierra Leone and Ascension L) ; Nelson, Rep. St. Geol. N. J.,
2 (2), 1890: 660 (N. Jersey, not common); Vincinguerra, Atti Soc. ital. Sci. nat., 34, 1892: 30
(ref. to obvclatus Valenciennes, 1 844, Canaries); Jordan, Manual Vert. Anim. NE. U.S., 1929: 10
(general); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 10 (N. and mid.
Atlant.); Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda, 1933: 28 (descr., Bermuda); Belloc,
Rev. des Trav. Peches Marit., 7 Fasc. 2, 1934: 132 (ill. after Lesueur; Canaries, Madeira).

Prionodon obvtlatus Valenciennes, in Webb and Berthelot, Hist. Nat. Canaries, 1844: 103, pi. 26 (descr.
applicable to obscurus, but ill. not recognizable; Canaries spec).

Squalus (Carcharinus) obscurus Gray, List. Fish. Brit. Mus., 1 851: 47 (N. Amer.).

Ptatyfodon obscurus Gill, Proc. Acad. nat. Sci. Philad., 1864: 262 (east. Amer.).

Carcharias {Prionodon) obvclatus Dumeril, Hist. Nat. Poiss., I, 1865: 376 (redescr., applicable to obscurus
except perhaps teeth; Canaries).

Carcharhinus (Eulamia) obscurus Joidixi, Rep. U.S. Comm. Fish. (1885), 1887: 795 (name, N. Amer.).

Carcharinus obscurus (at least in part) Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 22 (descr.
after Lesueur, N. Atlant.) ; Bean, T. H., Rep. For. Comm. N. Y., 1901: 377; Bull. N. Y. St. Mus.,
60, Zool. 9, 1903: 25 (descr., perhaps confused with other species; reported abund., N. Jersey,
no doubt milberti); Fowler, Rep. N. J. Mus. (1906), 1907: pi. 72 (ill., after Lesueur); Sharp
and Fowler, Proc. Acad. nat. Sci. Philad., 56, 1907: 505 (Nantucket spec, ident. accepted be-
cause of large size, II ft. 6 in.) ; Garman, Mem. Harv. Mus. comp. Zool., 36, 191 3: 130 (descr.. Buz-
zards Bay spec.) ; Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., 31 (2), 1913: 735 (probably partly
obscurus because of loc, Woods Hole, but refers no doubt to milberti chiefly) ; Nichols, Bull. Amer.
Mus. nat. Hist., 57, 1917: 873 (discuss.); Copeia, 140, 1922: 21 (compares with milberti, from
photographs, but mid-dorsal ridge said to be absent, Woods Hole) ; Meek and Hildebrand, Field
Mus. Publ. Zool., J 5 (i), 1923: 46 (descr., N. and mid. Atlant., no definite loc); Bigelow
and Welsh, Bull. U.S. Bur. Fish., 40 (i), 1925: 29 (descr. and ill., but Gulf of Maine records
probably were Prionace glauca); Barnard, Ann. S. Afr. Mus., 21 (i), 1925: 25 (spec, Table Bay,

5. Afr.); Nichols, Copeia, 140, 1925: 21 (comparison with other species); Thorne, Bull. N. Y. zool.

126. Owing to the fact that other species, especially C. milberti, have frequently been reported as obscurus, the
following list is limited to such references as can be referred to the latter with some confidence from included
evidence.

3 go Memoir Sears Foundation for Marine Research

Soc, 31, 1928: 114 (two reported harpooned near Long Island, N. York, ident. acceptable because
distinguished from milberti similarly taken); Nichols and Breder, Zoologica, N. Y., 9, 1929: 15
(states confused with milberti, but young near N. York Sept.— Nov.) ; Breder, Field Bk. Mar. Fish.
Atlant. Coast, 1929: 16 (ill., but distrib. evidently confused with that of milberti); Bellon and
Mateu, Notas Inst. esp. Oceanogr., (2) 53, 1 931: 15 (ref. to obvelatus Valenciennes, 1844, Cana-
ries); Firth, Bull. Boston Soc. nat. Hist., 61, 1931: 9 (li-ft. spec. Georges Bank, ident. prob-
able because of large size); Bigelow and Schroeder, Canad. Atlant. Fauna, 12', 1934: 7 (ill.,
Georges Bank, but C. Elizabeth record probably for Prionace glauca) ; Norman, "Discovery" Rep.,
/2, 1935: 56 (name, listed for St. Helena, W. Indies, Brazil, Canaries, C. Verde I., W. Afr., S. Afr.) ;
Bigelow and Schroeder, Bull. U.S. Bur. Fish., 48^ 1936: 321 (Georges Bank) ; Cadenat, Rev. des Trav.
Peches Marit., 10 (4), 1937: 430 (C. Verde; Dakar); Norman and Fraser, Giant Fishes, 1937: 36
(name, N. and mid. Atlant.); Springer, Proc. Fla. Acad. Sci., 5, 1939: 20, 23-24 (descr., ill., size,
season, embryos, Florida); Wise, Nat. Hist. N. Y., 21?, 1938: 323 (recognizable photo, Bahamas);
Boos, Contr. biol. Lab. Cath. Univ. Amer., 45, 1943: 10 (chemistry of pancreas, Florida); Springer,
Copeia, 1946: 174 (food, off Salerno, Florida).

Corcharhinus (Platyfodon) obscurus Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 215
(name, N. Atlant.); Bull. U.S. nat. Mus., 47 (l), 1896: 35 (descr., refs., N. and Mid. Atlant.).

Eutamia obscura Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 60 (name only); Fowler, Rep. N. J.
Mus. (1905), 1906: 61 (l2-ft. spec, off C. Henlopen, Delaware, ident. probably correct because of
large size) ; Bull. Amer. Mus. nat. Hist., yo (l), 1936: 49 (list of East Atlant. loc, descr. and ill. of
N. Jersey specs.) ;^^^ Bigelow and Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm.,
Wash., 1945: 82, fig. 25 (descr., habits, range, ill.).

References Probably Based on Some Other Shark:

Carcharias obscurus Storer, Boston J. nat. Hist., 2, 1839: 533 (teeth) ; Mass. Zool. Bot. Surv., Rep. on Fish.,
1839: 184 (same as preceding); Mem. Amer. Acad. Arts Sci., N. S. p, 1867: 219, pi. 36, fig. 2; also
Fishes Mass., 1867: 243, pi. 36, fig. 2 (this probably was Prionace glauca; see p. 292) ; Bean, Bull. U.S.
Fish Comm., 7, 1889: 132, 152 (this probably milberti because of reported abund. in bays of N. Jersey).

Corcharhinus obscurus Smith, Bull. U.S. Bur. Fish., 17, 1898: 88 (Woods Hole, more likely milberti because
descr. as "very common"); Linton, Bull. U.S. Bur. Fish., 24, 1905: 339 (N. Carolina, but apparently
Negafrion brevirostris from his descr.); Tracy, Rep. R. I. Comm. inl. Fish., 1906: 45 (probably
milberti, from context); Fowler, Rep. N. J. Mus. (1906), 1907: 256 (N. Jersey, "commersonii,"
i.e., leucas Muller and Henle, 1841, according to Fowler, Proc. biol. Soc. Wash., 33, 1920: 144);
Smith, Bull. N. C. geol. econ. Surv., 2, 1907: 33 (N. Carolina, probably Negafrion brevirostris, see
Linton, 1905); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 4 (reps, for Gulf of Maine
probably Prionace glauca; those for Connecticut and Rhode Island Carcharhinus milberti") ; Tracy,
Rep. R. I. Comm. in!. Fish., 1910: 59 (probably milberti, from context); Thome, Copeia, 35, 1916:
69 (name only, Long Island, N. York; probably milberti); Nichols, Copeia, 35, 1916: 72, 73 (Long
Island, N. York spec, actually "commersonii," i.e., leucas Miiller and Henle, 1841 ; see correction by
Nichols, Bull. Amer. Mus. nat. Hist., 37, 1917: 873); Latham, Copeia, 71, 1919: 53 (20 to 22-in.
young. Long Island, N. York, probably not obscurus because smaller than recorded embryos of latter, see
p. 387) ; Truitt, Bean and Fowler, Bull. Md. Conserv. Dep., 3, 1929: 30 (descr. is of Prionace glauca).

Carcharhinus (Plat-jodon) obscurus Pietschmann, Ann. naturh. (Mus.) Hofmus. Wien., 21, 1906: 99 (Mo-
rocco, not obscurus because origin of 1st dorsal described as over middle of pectoral base; perhaps
flumbeus'^Sirdo, 1827).

Western Atlantic Records by Name Only, Without Other Indication as to Identity:

Carcharias obscurus Linsley, Amer. J. Sci., 47, 1844: 76 (Connecticut); Goode, Bull. U.S. nat. Mus., 5,
1876: 76 (Bermuda); Amer. J. Sci., (3) 14, 1877: 293 (Bermuda); Goode, Fish. Fish. Indust. U.S.,
I, 1884: 672 (Woods Hole); Wilson, Proc. U.S. nat. Mus., 33, 1908: 624 (N. Carolina); Nichols,
Abstr. Linn. Soc. N. Y., 20-23, 1913: 91 (off N. York); Wilson, Proc. U.S. nat. Mus., 64 (17),

127. S^tWti "obscurus."

Fishes of the Western North Atlantic 391

1925: 12 (Woods Hole); Berc, Amer. Midi. Nat., ly, 1936: 593, 604 (Englewood, Florida); Longley
and Hildebrand, Pap. Tortugas Lab., 34, 1941: 2 (Tortugas, Florida).

Eulamia ohscurus Verrill and Smith, Rep. U.S. Comm. Fish. (1871-1872), 1873: 520 (Woods Hole);
Bean, Proc. U.S. nat. Mus., j, 1881: 115 (Woods Hole); Fowler, Copeia, 30, 1916: 36 (in list);
Copeia, 31, 1916: 41 (N.Jersey).

Carcharinus obscurus Rathbun, Proc. U,.S. nat. Mus., 7, 1885: 488, 489 (off N. York, Lat. 39° 30' N.,
Long. 72° W.); Moore, Bull. U.S. Fish Comm., 12, 1894: 358 (N. Jersey); Linton, Bull. U.S. Bur.
Fish., /5, 1894: 104 (parasites); Proc. U.S. nat. Mus., 20, 1897: 424, 452 (no loc.) ; Means, Bull.
Amer. Mus. Nat. Hist., 10, 1898: 311 (Hudson R., N. York); Linton, Bull. U.S. Bur. Fish., ig, igoi:
272, 426 (food, parasites, Woods Hole); Hargreaves, Fish. Brit. Guiana, 1904: 14, app. 7 (Brit.
Guiana, abundance, attacks on man); Bean, B. A., in Shattuck, Bahama Islands, 1 905: 296 (Baha-
mas); Linton, Bull. U.S. Bur. Fish., 26, 1907: 122, 123 (parasites, N. Carolina); Sullivan, Bull.
U.S. Bur. Fish., 27, 1907: 13 (name only); Wilson, Proc. U.S. nat. Mus., 55, 1907: 326, 360, 409,
414, 423, 431, 629 (parasites, Woods Hole region) ; Linton, Bull. U.S. Bur. Fish., 28 (2), 1910: 1200
(parasites); Vincent, Sea Fish. Trinidad, 1910: 53 (Trinidad, W. Indies, abund.) ; Rosen, Lunds
Univ. Arsberatt., N. S. 7 (5), 1911: 47 (Bahamas); Gudger, Yearb. Carneg. Instn., 12, 1913: 177
(Tortugas, Florida); Nichols, Copeia, 36, 1916: 81; Rockwell, Brooklyn Mus. Quart., 5, 1916: 162
(Long Island, N. York) ; Latham, Copeia, 99, 1921:72 (Long Island, N. York) ; Breder, Copeia, 127,
1924: 25 (Sandy Hook B.ay, N. York); Linton, Proc. U.S. nat. Mus., 64 (21), 1924: 5, 7, 12, 30, 34,
38, 47, 48, 49, 65, 80, 87, 90 (parasites. Woods Hole and N. Carolina) ; Wilson, Bull. U.S. nat. Mus.,
158, 1932: 463, 464, 524 (parasites. Woods Hole region) ; Nigrelli, Amer. Mus. Novit., 996, 1938: 10
(parasites); Norris, Plagiost. Hypophysis, 1 941: 28 (brain); Lunz, Bull. S. Carolina St. Planning Bd.,
14, 1944: 27 (Florida).

Carcharias {Prionodon) obscurus Werner, Zool. Jb., Syst. Abt., 2/, 1904: 283 ("Carteret County, New York,"
N. Carolina perhaps intended).

Eulamia obscura Bean, T. H., Field Mus. Publ. Zool., 7 (2), 1906: 29 (Bermuda).

Carcharinus sp. (probably C. obscurus) Norris, Plagiost. Hypophysis, 1 941 : pi. I, fig. 2 (brain).

Carcharhinus oxyrhynchus Miiller and Henle, 1841
Figure 73

Study Material. Stuffed dried skin of a female, about 443 mm. long, from Suri-
nam (.?) (Acad. Nat. Sci. Philad., No. 34635).'='

Distinctive Characters. This species is sharply marked off from all other Atlantic
members of the genus by the large number of teeth and by the great length of the very
narrow snout anterior to a line connecting the outer ends of nostrils, this being about i V2
times as long as the distance between the inner ends of nostrils, both in the original illustra-
tion of the species and in the specimen listed above. In this respect, and in the narrowness
of its teeth, it seems to be most nearly related to C. velox Gilbert, 1 898, of the west coast
of Central America, but it is separable from velox by its relatively much shorter nostrils
and by the fact that the outer margins of the upper teeth toward the corner of the mouth
are not notched.

Description. Proportional dimensions are not available.'^^

Trunk moderately slender, the back without trace of mid-dorsal ridge between

128. Specimen recorded by Fowler (Proc. Acad. nat. Sci. Philad., 60, 1908: 65).

129. The specimen listed above was obviously so greatly lengthened in the process of stuffing that measurements
taken from it would be only roughly approximate.

392 Memoir Sears Foundation for Marine Research

1st and 2nd dorsal fins. Upper precaudal pit well marked, subangular, a little less than a
right angle, the lower pit only faintly indicated. Dermal denticles overlapping only
partly, with the skin visible between them here and there, their blades broader than long,
usually with 3 (rarely 5) strong longitudinal ridges and as many moderately strong
teeth, the median a little the longest j pedicels very short.

c

aAAAAAAAAAA AAAAAa^a,<aa<^^

Figure 73. Carcharhhms oxyrhynchus. A, B Immature male, about i8 inches long, from Surinam, somewhat
emended after Muller and Henle. C Upper and lower teeth, after Muller and Henle. D An upper and a
lower tooth, much enlarged, after Muller and Henle. E Dermal denticles from a female, about 443 mm.
long, from Surinam, much enlarged (see Study Material, p. 391).

Head with snout relatively much longer than in other local species of CarcharhinuSy
its length to origin of pectoral about 40% of length of trunk to origin of caudal in the
original illustration. Snout narrow, wedge-shaped, contracted anterior to nostrils, with
narrowly rounded tip, its length in front of a line connecting outer ends of nostrils
about I V2 times as great as distance between inner ends of latter, its length in front of
mouth about twice as great as distance between nostrils or about Y^ ^s great as length of
head to origin of pectoral."" Anterior margin of eye very little posterior to front of
mouth. Gill openings about twice as long as diameter of eye, the space between 4th and
5th over origin of pectoral.^^' Nostril only slightly oblique, its inner end only about V2
as far from front of mouth as from tip of snout, its anterior margin only slightly expanded,

130. This is according to the original illustration; in the dried skin the head has obviously been lengthened in the
process of stuffing.

131. In the stuffed skin the 5th is a little anterior to the origin of the pectoral, but this is probably due to distortion.

Fishes of the Western North Atlantic 393

in obtusely subangular outline. Mouth narrow-ovate, about 75% as high as broad. Upper
labial furrow about Vs as long as diameter of eye.""

Teeth in type specimen about |^2^|; similar in the 2 jaws, symmetrical, erect,
with slender cusps on broadly expanded bases, the edges of cusps coarsely serrate in
uppers but very finely so in lowers,'^^ the edges of bases smooth; 4 minute teeth at sym-
physis in upper jaw and 2 in lower, the 3 or 4 outermost teeth in each jaw also very small.

Origin of ist dorsal a little anterior to axil of pectoral in specimen seen, but over mid-
base of pectoral in original illustration, its base a little more than Vs as long as head to
origin of pectoral by original illustration, its anterior margin moderately convex and
increasingly so toward apex, the posterior margin moderately concave, apex subacute or
very narrowly rounded, the free rear tip broad, about y^ to y^ as long as the base, its
vertical height a little less than M: as great as length of pectoral, the midpoint of base less
than V2 as far from axil of pectoral as from origin of pelvics. Second dorsal a little more
than Yi as long at base as ist, its origin about over origin of anal,"^ its apex narrowly
rounded, rear margin weakly concave, its free rear tip moderately slender and about V2 as
long as base. Caudal about ^/4 of total length, its upper margin moderately convex,
its tip very narrowly rounded or subacute, the terminal sector about ^^ the length
of fin in original illustration (about y^ in dried skin), the lower lobe a little more than
Vs as long as upper (about 36% in original illustration but about 33% in dried skin), its
anterior margin strongly convex toward apex, the re-entrant corner (included by the 2
lobes) moderately rounded. Distance from origin of caudal to tip of anal % (original
illustration) to % (dried skin) as long as base of latter. Anal a little shorter at base than
2nd dorsal but about equally high, its anterior margin only slightly more convex and rear
margin more concave than in 2nd dorsal, apex narrowly rounded, free rear corner only
about V2 as long as base. Distance from origin of anal to tips of pelvics about V2 (in
original illustration) to % (dried skin) as long as base of anal. Pelvics with nearly
straight margins, about as long at base as anal. Pectoral about % as long as head in original
illustration but only about V2 that long in dried skin, about V2 as broad as long, with
moderately convex outer margin but only very slightly concave distal margin and nearly
straight inner margin, rounded inner corner and subacute tip.

Color. Described as yellow-gray above, white below.

Size. The greatest length so far definitely reported for it is about five feet (1,520
mm.), although it has been said to attain six to eight feet. The fact that embryos of 14
inches have been reported, and free-living specimens 17 to 18 inches, suggests that a length
of about 1 5 to 16 inches is usual at birth.

132. GiJnther (Cat. Fish. Brit. Mus., 1870: 375) credits it with a short labial furrow on the lower jaw as well as
on the upper. Actually, however, the dried skin shows that the upper alone is present, as in other members of
the genus.

133. Miiller and Henle (Plagiost., 1841 : 41) state that the upper teeth are serrate only toward their tips, the lowers
smooth. But their illustration (pi. 13) shows the uppers as serrate from tip to base, and the lowers as very finely
serrate toward their tips.

I 34. A little anterior to latter in original illustration, but a little posterior to it in dried skin.

394 Memoir Sears Foundation for Marijie Research

Developmental Stages. Development is viviparous, the (four) embryos having been
described as attached to the mother by a placenta.""'

Habits. All that is known of its habits is that it has been described (if identified cor-
rectly) as often entering estuaries and river mouths and feeding on small fish, ravaging
schools of clupeids and sciaenids in particular."" Apparently it is a littoral species.

Relation to Man. While not considered very desirable as food, some are sold in the
markets of Trinidad and no doubt in the Guianas as well.

Range. Western tropical Atlantic. The few records of this species are from: Surinam,
Dutch Guiana J Demerara, British Guiana; French Guiana, where it has been described
as rather common; Trinidad."'

Synonyms and References:

Carcharias (Prionoiori) oxyrhynchus Muller and Henle, Plagiost., 1841: 41, pi. 15 (descr., ill., Surinam);
MiJller and Troschel, in Schomburgk, R. H., Reisen Brit. Guiana, 1840-44,5, 1848:642 (Brit. Guiana,
said to grow to 6 to 8 ft.) ; Dumeril, Hist. Nat. Poiss., /, 1865; 356 (descr., Cayenne, Surinam) ; Gun-
ther. Cat. Fish. Brit. Mus., 8, 1 870: 375 (descr., no loc).

hogomfhodon oxyrhynchus Gill, Ann. N. Y. Lye, 7, 1862: 410 (name).

Carcharhinus {Isogomfhodon) oxyrhynchus Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896:
216 (name, Surinam); Bull. U.S. nat. Mus., 47 (i), 1896: 40 (descr., Surinam).

Carcharinus oxyrhynchus Hargreaves, Fish. Brit. Guiana, 1904: 14, and append. 8 (name only, Brit. Guiana) ;
Vincent, Sea Fish. Trinidad, 1910: 53 (name only, Trinidad) ; Garman, Mem. Harv. Mus. comp. ZooL,
5<5, 191 3: 143 (descr., east. S. Amer.) ; Puyo, Bull. Soc. Hist. nat. Toulouse, yo, 1936: 85, 87,'**
250 (ill., but labels transposed, habits, food, vernacular name, French Guiana); White, Bull. Amer.
Mus. nat. Hist., ■j4, 1937: 128 (in Key).

Euhmia oxyrhynchus Fowler, Proc. Acad. nat. Sci. Philad., 60, igo6: 65 (meas. of spec, listed above in Study
Material, Surinam?); Proc. Acad. nat. Sci. Philad., 67, 1916: 521^'' (female with embryos, in market,
Trinidad); Proc. Acad. nat. Sci. Philad., 7/, 1919: 129 (listed for Surinam).

Carcharhinus forosus RanzanI, 1839
Figures 74, 75

Study Material. Seven small specimens, 330 to 395 mm. long, from Surinam,
Pernambuco and Bahia (Harv. Mus. Comp. ZooL, No. 307, 526, 721, 1403, 1404);
female, 485 mm., and male, 500 mm., from Colon (U.S. Nat. Mus., No. 79317, 79316);
male, 831 mm.. Pacific Panama (U.S. Nat. Mus., No. 79293); also two females, from
Peru (Harv. Mus. Comp. ZooL, No. 692).

Distinctive Characters. Porosus differs from all other western Atlantic members of
the genus in that the second dorsal originates about over the midpoint of the base of the
anal, that the terminal sector of the caudal is relatively smaller and that the outermost four
or five lower teeth are strongly asymmetrical with deeply notched outer margins like the
uppers.

Description. Proportional dimensions in per cent of total length. Female, 485 mm.,

135. Fowler, Proc. Acad. nat. Sci. Philad., 6y, 1916: 521.

136. Puyo, Bull. Soc. Hist. nat. Toulouse, jo, 1936: 87.

137. Specimens are in the British Museum and in the museums of Leyden and Paris.

138. Spelled "Carcharhynus." 139. Spelled oxyrhinchus.

Fishes of the Western North Atlantic

395

from Colon, Panama (U.S. Nat. Mus., No. 79317). Male, 500 mm., same locality (U.S.
Nat. Mus., No. 79316).

Trunk at origin of pectoral: breadth 12.4, 1 1.45 height 1 1.9, 1 1.8.

Snout length in front of: outer nostrils 4.5, 4.35 mouth 8.9, 8.7.

Eye: horizontal diameter 2.3, 2.1.

Mouth: breadth 8.2, 7.9; height 5.4, 5.3.

Figure 74. Carcharhinus forosus, immature male, about 377 mm. long, from Pernambuco, Brazil (Harv.
Mus. Comp. ZooL, No. 526). A Upper and lower teeth, left-hand side, about 4 x. 5 Fifth upper tooth.
C Ninth upper tooth. D Fourth lower tooth. E Seventh lower tooth. F Tenth lower tooth. C-F, about 7.6 x.
G Eighth to thirteenth lower teeth of a female, 485 mm. long, from Colon, Atlantic Panama (U. S. Nat. Mus.,
No. 793 16), about 3 X. // Seventh lower tooth. / Tenth lower tooth of same, about 7.6 x.

Figure 75. Carcharhinus forosus, pictured in Fig. 74. A Anterior part of head from below, about % natural
size. B Dermal denticles, about 50 x. C Apical view of dermal denticle, about 100 x. Z) Left nostril, about 6 x.

396 Memoir Sears Foundation for Marine Research

Nostrils: distance between inner ends 5.8, 5.8.

Gill opening lengths: ist2.3,2.4; 2nd 2.5, 2.6; 3rd 2.7, 2.8; 4th 2.7, 2.8; 5th 2.5,

2.2.

First dorsal fin: vertical height 9.1, 9.4; length of base ii.i, 10.8.

Second dorsal fin: vertical height 2.5, 2.9; length of base 3.4, 3.4.

Anal fin: vertical height 3.4, 3.2; length of base 4.6, 4.6.

Caudal fin: upper margin 26.3, 26.6; lower anterior margin 1 1.7, 12.4.

Pectoral fin: outer margin 15.5, 15.7; inner margin 5.9, 6.0; distal margin 12.2,

12.4.

Distance from snout to: ist dorsal 33.0, 32.2; 2nd dorsal 64.3, 62.6; upper

caudal 73.7, 73.4; pectoral 23.5, 23.8; pelvics 47.O, 48.6; anal 60.7, 60.4.

Interspace between: ist and 2nd dorsals 20.7, 20.6; 2nd dorsal and caudal 7.8,

7.0; anal and caudal 7.6, 6.8.

Distance from origin to origin of: pectoral and pelvics 25.2, 26.O; pelvics and anal

13.4,13.8.

Proportional dimensions in per cent of total length. Male, 831 mm., from Pacific
Panama (U.S. Nat. Mus., No. 79293).

Trunk at origin of pectoral: breadth 1 1.2; height 12.0.

Snout length in front of: outer nostrils 4. i ; mouth 7.5.

Eye: horizontal diameter 1.6.

Mouth: breadth 8.2; height 5.1.

Nostrils: distance between inner ends 5.2.

Gill opening lengths: 1st 3.1 ; 2nd 3.5; 3rd 3.4; 4th 3.4; 5th 3.0.

First dorsal fin: vertical height 9.5; length of base 12.2.

Second dorsal fin: vertical height 2.9; length of base 4.6.

Anal fin: vertical height 3.8 ; length of base 4.7.

Caudal fin: upper margin 24.7; lower anterior margin 1 1.9.

Pectoral fin: outer margin 16.0; inner margin 5.8; distal margin 14.2.

Distance from snout to: ist dorsal 33.3; 2nd dorsal 64.7; upper caudal 75.3;

pectoral 23.6; pelvics 48.2; anal 62.5.

Interspace between: ist and 2nd dorsals 21.9; 2nd dorsal and caudal 7.4; anal

and caudal 7.1.

Distance from origin to origin of: pectoral and pelvics 25.3; pelvics and anal 13.5.

Height of trunk at ist dorsal about y^ its length to origin of caudal. Midline of back
smooth, with no trace of mid-dorsal ridge. Caudal peduncle about % as thick as high.
Upper precaudal pit strongly marked, semilunar to subangular, the lower similar but
less strongly marked. Dermal denticles overlapping so little that the skin is regularly or
partially exposed between them, their blades rising rather steeply and varying in size, only
a little broader than long, moderately arched longitudinally, with 3 ridges in small speci-
mens but 5 in larger, the median tooth considerably the largest; pedicels very short.

Fishes of the Western North Atlantic 'i^c^'j

Head about V4 of total length, its dorsal profile very weakly convex, about as. broad
at eyes as at origin of pectorals. Snout ovoid with rather broadly rounded tip, its length in
front of a line connecting outer ends of nostrils very nearly equal to distance between
inner ends of latter on smallest specimens but only % that in larger, the length in front of
mouth about 1.4 to 1.5 times as great as distance between nostrils and a little less than %
(37 to 38%) as great as length of head to origin of pectorals. Eye approximately circular
and relatively large, its diameter a little less than V2 as long as distance between inner
ends of nostrils in smallest specimens, Vs that long in larger, its anterior edge opposite
front of mouth or a little anterior to latter. Gill openings very slightly oblique with sinu-
ously concave contours, the ist about as long as the horizontal diameter of the eye in
smallest specimens but increasing relatively to about i .2 times as long as eye in specimens of
500 mm. and twice the eye at a length of 800 to 900 mm., the 3rd very little longer than
1st, the 5th about as long as ist, the 4th about over origin of pectoral. Nostrils moderately
oblique, about V^ as long as the distance between their inner ends, which are nearer to
mouth than to tip of snout by a distance about V2 as long as between nostrils, the anterior
margins somewhat sinuous and expanded near inner end as a short digitate lobe with
rounded tip."" Mouth broad-ovoid, its height relative to its breadth somewhat greater in
smaller specimens (60 to 66%) than in the larger (about 54%), occupying about % of
breadth of head.

Teeth i2ori3^o'rT— i2ori3 > ^PP^''^ broadly triangular, their edges serrate, most coarsely
so basally except for the 2 or 3 outermost which are only slightly irregular, the ist nearly
symmetrical with nearly straight edges, but 2nd and subsequent teeth increasingly oblique,
their inner margins slightly sinuous or concave, their outer margins notched more and
more deeply toward corners of mouth, the lOth to 13th successively lower, the outermost
tooth very low, its cusp hardly discernible, the median upper tooth small and symmetrical;
lower teeth with much narrower cusps than uppers, on expanded bases, the ist to 8th or
9th serrate from tip to base, although somewhat less coarsely so than uppers, but subse-
quent teeth irregularly wavy at most, the ist much smaller than 2nd to 6th, those toward
center of mouth erect, nearly symmetrical, the outermost 4 or 5 in smallest specimens, but
outermost 1-3, only, in larger examples, very oblique, deeply notched outwardly and
with very low cusps.

Origin of ist dorsal about over midpoint of inner margin of pectoral, its anterior
margin about V2 as long as head, its vertical height about as great as length of snout in
front of mouth in small specimens but relatively somewhat greater in larger, its anterior
margin only very slightly convex toward apex, its posterior margin moderately and evenly
concave, its apex narrowly rounded, its free rear corner about V2 as long as base, the mid-
point of base about % as far from axil of pectoral as from origin of pelvics. Second dorsal
about Vs (30 to 37% ) as long at base as ist, about V3 to V4 as high vertically and relatively
much lower in form, its origin about over midpoint of base of anal, the rear end of its

140. This is clearly shown in Ranzani's illustration (Nov. Comment. Acad. Sci. Inst. Bonon [Bologna], ^, 1839:
pi. 2).

398 Memoir Sears Foundatioft for Marine Research

base a little posterior to rear end of base of anal, its tip a little posterior to tip of anal,
its apex only very narrowly rounded, its free rear corner slender and about as long as
base. Caudal about Yi of total length, its upper margin slightly to moderately convex,
its terminal sector tapering and a little less than ^,4 the length of fin, with narrowly
rounded tip and weakly concave lower posterior margin, the lower lobe a Jittle less than Y2
as long as upper with its tip narrowly rounded, the re-entrant corner (enclosed by the 2
lobes) more obtuse than a right angle and broadly rounded. Distance from caudal to tip
of anal about as long as base of anal in smallest specimens but only about 73 that long in
larger. Anal about i.o to 1.3 times as long at base as 2nd dorsal, its anterior margin much
more strongly convex and posterior margin much more deeply concave, its apex more
rounded, its free corner slender and about as long as base or slightly shorter. Distance from
origin of anal to tips of pelvics a little longer than base of anal. Pelvics with nearly straight
anterior and posterior margins, about as long at base as anal. Pectoral noticeably small,
about % as long as head or only a little longer than anterior margin of ist dorsal and a
little more than Y^ as broad as long, the outer margin only very weakly convex, the distal
margin weakly and evenly concave, the tip and inner corner very narrowly rounded.

Color. Described as leaden or bluish gray above, the sides sometimes tinged with red-
dish, the lower surface pale 5 the pelvics sometimes with reddish tinge toward their bases j
edges of lower fins and hind edge of lower caudal lobe white.

Size. The largest specimen so far recorded was 1,235 mm. (49 inches) long."^ The
fact that the claspers in an 831-mm. Pacific specimen are twice as long as the pelvic fins
suggests that this shark does not reach a length much greater than perhaps four feet.

Develo-pmental Stages. Embryos have not been described as yet.

Remarks. The specimens listed above can be referred to forosus without hesitation,
so clearly diagnostic are the original account and illustrations of that species."^

Habits. The localities of capture, listed below, show this to be a strictly subtropical-
tropical species and probably littoral. But nothing more is known of its habits.

Relation to Man. Saleable for human food in the markets of Colon and Panama.

Range. Western tropical Atlantic; northern Brazil to north shore of Gulf of Mexico;
also eastern tropical Pacific, Peru to Panama;"' represented on the Atlantic coast of North
Africa by a form that may finally prove to be identical."*

141. Herre, Field Mus. Publ. Zool., 21, 1936: 22; a Pacific specimen.

142. Ranzani, Nov. Comment. Acad. Sci. Inst. Bonon (Bologna), ./, 1839; 8, pi. 2.

143. Our comparison of specimens from Payta, Peru and Panama with others from the Atlantic corroborates Meek
and Hildebrand's (Field Mus. Publ. Zool., 75 [i], 1923: 49) conclusion that examples from the two sides
of the Isthmus of Panama represent only a single species, which they recorded and described as cerdale Gilbert,
1898. But we find nothing to separate the latter from the original account and illustrations of forosus Ran-
zani, 1839. And we should perhaps point out that Carman's (Mem. Harv. Mus. comp. Zool., 36, 1913: 131)
account of forosus as with broadly rounded snout and with nostril as far from end of snout as from eye, charac-
ters used by Meek and Hildebrand in their Key as alternative between forosus and cerdale, does not fit the West
Indian and Brazilian specimens in the Museum of Comparative Zoology, from which Carman's description
appears to have been taken.

144. Carckarias fissidens Bennett (Proc. zool. Soc. Lond., 1830-1831: 148), in which the origin of the second
dorsal is described as over the middle of the anal and the outer edges of the teeth as deeply notched. But the
account of it is not detailed enough for decision.

Fishes of the Western North Atlantic 399

Occurrence in the Western A tlantic. Positive records for -porosus are from Bahia, Per-
nambuco, Marajo Island at the mouth of the Amazon; British, Dutch and French Guiana;
Trinidad; Colon; and from the north shore of the Gulf of Mexico."' Evidently it ranges
generally throughout the Gulf of Mexico, Caribbean region, and southward as far as
central Brazil.

Synonyms and References:

Carcharias -porosus Ranzani, Nov. Comment. Acad. Sci. Inst. Bonon (Bologna), 4, 1839: 8, pi. 2 (descr.,
color, excellent ill., Brazil); Boulenger, Ann. Mag. nat. Hist., (6) 20, 1897: 298 (Marajo I., Brazil);
Goeldi, Bol. Mus. Paraense, 2, 1898: 488 (Marajo I., Brazil) ; Ribeiro, Ann. Mus. nac. Rio de J., 14,
1907: 153, 200 (descr., Brazil, refs.) ; Pellegrin, Poiss. Guyana Franc, Rev. Colon., 67, 1908: II
(French Guiana); Ribeiro, Fauna brasil. Peixes, 2 (1) Fasc. I, 1923: 11 (descr., Brazil).

Carcharias {Prionodon) henlei Miiller and Henle, Plagiost., 1841: 46,"° pi. 19, fig. 6 (descr., ill. of
teeth, Cayenne); Miiller and Troschel, in Schomburgk, Reisen Brit. Guiana (1840-1844), 3, 1848:
641 (Brit. Guiana) ; Dumeril, Hist. Nat. Poiss., i, 1865: 372 (descr., Cayenne).

Carcharias (Prionodon) -porosus Dumeril, Hist. Nat. Poiss., i, 1865: 373 (Brazil, Cayenne); Gunther, Cat.
Fish. Brit. Mus., 8, I 870: 365 (Guiana).

Carcharhinus {Platypodon) henlei Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 216 (Brazil
to Guiana) ; Bull. U.S. Nat. Mus., 47 (l), 1896: 37 (descr., Brazil to Guiana).

Carcharhinus cerdale Gilbert, in Jordan and Evermann, Bull. U.S. nat. Mus., 47 (3), 1898: 2746 (descr.,
Pacif. Panama); Wilson, Ann. Carneg. Mus., to, 1916: 58 (Colombia, Ecuador); Meek and Hilde-
brand. Field Mus. Publ. Zool., 15 (l), 1923: 47 (descr., size, embryos, abund., Atlant. and Pacif. Pan-
ama) ; Herre, Field Mus. Publ. Zool., 21, 1936: 22 (refs., descr., max. size, Galapagos and at sea).

Carcharias cerdale Gilbert and Starks, Mem. Calif. Acad. Sci., 4, 1904: 10, pi. 2, fig. 4 (Panama Bay);
Starks, Proc. U.S. nat. Mus., 30, 1 906: 762 (Ecuador) ; Jordan, Evermann and Clark, Rep. U.S. Comm.
Fish. (1928), 2, 1930: 16 (Panama).

Carcharinus tnenisorrah (in part) Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 135 {cerdale incl. In
synonymy) ; not Carcharias {Prionodon) menisorrah Muller and Henle, 1 841.

Carcharinus porosus Garman, Mem. Harv. Mus. corap. Zool., 36, 191 3: 131 (descr., W. Indies, Brazil);
Meek and Hildebrand, Field Mus. Publ. Zool., 15 (0> 1923: 49 (descr.) ; White, Bull. Amer. Mus. nat.
Hist., -^4, 1937: 126 (in Key).

Eulamia porosus Fowler, Proc. Acad. nat. Sci. Philad., dp, 1917: 127 (Colon); Proc. Acad. nat. Sci. Philad.,
^J) '931- 391 (Trinidad); Arqu. Zool. Estado Sao Paulo, 3, 1942: 128 (listed, Brazil).

Carcharias henlei Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 16.

Carcharhinus lamiella Kumada and Hiyama, Mar. Fish. Pacif. Mexico, 1937: 16, pi. 48 (ident. by ill.); not
Carcharias lamiella Jordan and Gilbert, 1882.

Eulamia cerdale Fowler, Monog. Acad. nat. Sci. Philad., 2, 1938: 249 (listed for Galapagos); Beebe and
Tee-Van, Zoologica, N. Y., 26, 1941: 109 (descr., ill., refs., distrib. in Pacif.).

Doubtful References:

Carcharias fissidens Bennett, Proc. zool. Soc. Lond., 1830-31: 148 (Atlant. coast N. Afr., see footnote 144,

P- 398).
Carcharhinus henleiY\xyo, Bull. Soc. Hist. nat. Toulouse, yo, 1 936: 85, 89, 250 (descr., meas., French Guiana;
ident. doubtful since 2nd dorsal is described as "inseree sur le meme plan et au meme niveau que la
nageoire anale.").

145. Stewart Springer, in a personal communication, reports "a nice series of 'cerdale' " taken oflF Biloxi, Mississippi,
by himself in August 1943.

146. If this page was actually distributed in 1838, as it is stated in Jordan and Evermann (Bull. U.S. nat. Mus.,
47 [ill "896: 37) and in Jordan, Evermann and Clark (Rep. U.S. Comm. Fish. [1928], 2, 1930: 16) the
name henlei would have priority over porosus. Not being in a position to verify this, we credit Muller and
Henle's classic work with the date (1841) with which the title page is inscribed.

400 Memoir Sears Foundation for Marine Research

Carcharhinus remotus (Dumeril), 1865
Figures 76, 77

Study Material. Seven specimens from Rio de Janeiro, about 650 to 695 mm. long
(Harv. Mus. Comp. Zool., No. 703).'"

Distinctive Characters. Among the smooth-backed subdivision of the genus, with
which it falls, remotus is closest to limbatus and maculifinnis as regards teeth, but it is
separated from both of these by its much shorter gill openings (see Key, p. 324), as well
as by its lower precaudal pit, which is only faintly indicated, and by its fins which are
without conspicuous black markings, at least after preservation.

yyyyyyy^^

Figure 76. Carcharhinus remotus, female, about 690 mm. long, from Rio de Janeiro, Brazil (Harv. Mus.
Comp. Zool., No. 703). A Upper and lower teeth, left-hand side, about 2.6 x. B Third upper tooth.
C Eleventh upper tooth. D Fourth lower tooth. E Ninth lower tooth. B-E, about 5.2 x. F Left-hand nostril,
about 3.2 X.

Description. Proportional dimensions in per cent of total length. Male, 678 mm.,
from Rio de Janeiro (Harv. Mus. Comp. Zool., No. 703). Female, 693 mm., same local-
ity and number.

147. While provisional acceptance of Carman's (Mem. Harv. Mus. comp. Zool., 36, 1913: 138) reference of these
specimens to this species seems justified, the original account of it was so brief that the correctness of this identifi-
cation can be tested only by re-examination of the type specimen, novc or formerly in the Paris Museum.

Fishes of the Western North Atlantic

4-OI

Trunk at origin of -pectoral: breadth 9.3, 9.4; height 9.6, 9.5.

Snout length in front of: outer nostrils 3.8, 4.0; mouth 7.2, 7.1.

Eye: horizontal diameter 2.0, 2.0.

Mo«/A.' breadth 7.5, 7.4; height 5.2, 5.1.

Nostrils: distance between inner ends S-S^ S-S-

Gill opening lengths: ist 2.8, 2.7; 2nd 3.0, 2.8; 3rd 2.9, 2.7; 4th 2.5, 2.5; 5th

2.4, 2.0.

First dorsal fin: vertical height 8.5, 8.5; length of base 8.9, 9.0.

Second dorsal fin: vertical height 2.5, 2.5 ; length of base 4.0, 3.9.

Anal fin: vertical height 3.7, 3.6; length of base 4.4, 4.6.

Caudal fin: upper margin 27.O, 27.5; lower anterior margin 1 1.8, 11. 8.

Pectoral fin: outer margin 16.8, 17.O; inner margin 5.3, 5.5; distal margin 13.3,

13-6.

Distance from snout to: ist dorsal 33.6, 33.4; 2nd dorsal 64.4, 62.7; upper caudal

73.0, 72.5; pectoral 24.2, 23.5; pelvics 50.2, 48.7; anal 62.5, 61.5.

Interspace between: ist and 2nd dorsals 21.8, 21.1 ; 2nd dorsal and caudal 6.8,

6. 8 ; anal and caudal 6.2, 6. i .

Distance from origin to origin of: pectoral and pelvics 26.9, 26.4; pelvics and anal

12.5, I2.I,

Figure 77. Carcharhinus remotus, pictured in Fig. 76. A Anterior part of head from below. B Dermal
denticles: general view, about 45 x; apical view, about 90 x.

Trunk slender, its height at ist dorsal only about Vf. its length to origin of caudal.
Back smooth, without mid-dorsal ridge. Body sector to cloaca about l.i times as long as
tail sector. Caudal peduncle about % as thick as deep. Upper precaudal pit strongly devel-

402 Memoir Sears Foundation for Marine Research

oped, subangular and a little less than a right angle in outline, the lower pit only weakly
marked. Dermal denticles overlapping so regularly that the skin is exposed only here and
there, the blades rising rather steeply, a little broader than high, with 3 to 5 very low
ridges and 3 to 5 short, broad teeth, the median considerably the largest and the outermost
pair very small when there are 5; pedicels very short.

Head a little less than Vs length of trunk to origin of caudal, its dorsal profile sloping
evenly forward. Snout moderately thin-tipped, ovate, with broadly rounded tip, its length
anterior to a line connecting outer ends of nostrils a little more than % (about 69 to 73%)
of distance between inner ends of latter, its length in front of mouth about 1.3 times dis-
tance between nostrils and a little less than Vs (about 30%) as great as length of head to
origin of pectoral. Eye approximately circular, its anterior margin opposite or very slightly
behind front of mouth, its diameter a little more than V3 (about 35%) as great as distance
between nostrils. Gill openings about evenly spaced, their margins weakly concave, the
I St about V2 (50%) as long as distance between nostrils and a little less than 1.5 times as
long as horizontal diameter of eye, the 2nd and 3rd very slightly the longest, the 5th about
% (74 to 85%) as long as ist, the space between 4th and 5th over origin of pectoral.
Nostrils strongly oblique, about % as long as distance between their inner ends, which are
a little less than twice as far from tip of snout as from mouth, the anterior margin moder-
ately expanded as a low subangular lobe with rounded apex. Mouth ovate, about %
(67%) as high as broad, occupying about % of breadth of head. Upper labial fold between
Ys and V4 as long as diameter of eye.

Teeth '^ to i5--ij)r2— ujo 15 . uppers with narrow triangular cusps on moderately ex-
panded bases, nearly symmetrical toward center of mouth but slightly oblique toward its
corners, both margins concave, the outer margins increasingly so toward corners of mouth
in subangular contour, the edges regularly and rather finely serrate from tip to base except
for the outermost i or 2 teeth, which are only irregular or wavy, the ist to lOth or nth
teeth of about equal lengths, but subsequent teeth successively shorter, the outermost very
short J lower teeth with considerably narrower cusps than uppers, on more broadly ex-
panded bases, erect and symmetrical all along jaw, the edges much more finely serrate
than uppers from tip to base, the 2nd to nth longest and the outermost very short} i or 2
small teeth at symphysis in upper jaw and i in lower.

Origin of ist dorsal about over inner corner of pectoral, its vertical height a little
more than Vs as great as length of head, its anterior margin slightly convex toward apex,
its posterior margin moderately concave toward base, its apex moderately rounded, its
free rear corner only about Vs as long as the base, the midpoint of base only a little nearer
to axil of pectoral than to origin of pelvics. Second dorsal slightly less than V2 as long at
base as ist and a little less than Vs as high vertically, its apex broadly rounded, its rear
margin only very slightly concave, its free rear corner about as long as its base, its origin a
little posterior to origin of anal but considerably anterior to midpoint of latter. Caudal a
little more than V4 (27 to 28%) of total length, its upper margin only slightly convex

Fishes of the Western North Atlantic 403

with terminal sector between '^ and Vs (28 to 29%) the length of fin, slender, with nar-
rowly rounded tip, the lower lobe about 40% as long as upper, its tip narrowly rounded,
the re-entrant contour (included by the 2 lobes) well rounded. Distance from origin of
caudal to tip of anal about % as long as base of anal. Anal about i.i times as long at base
as 2nd dorsal, with slightly more convex anterior and much more deeply concave posterior
margins and rounded apex, its free rear corner only about % as long as base, its tip a little
anterior to that of 2nd dorsal. Distance from origin of anal to tips of pelvics a little shorter
than base of anal. Pelvics about as long at base as anal, and only about as large as latter in
area. Pectoral about 0.7 as long as head, and a little less than V2 as broad as long, the outer
margin weakly and evenly convex, the distal margin moderately concave, the apex and
inner corner both very narrowly rounded.

Color. Preserved specimens are mouse gray or brownish-gray above, paler below, the
fins with darker edges, but without conspicuous black markings. The color of fresh-caught
specimens has not been recorded.

She. The fact that two of the present series still show traces of the umbilical
scar suggests a length of about 600 to 650 mm. at birth. But any statement as to the size to
which remotus grows would be pure speculation, the only pertinent information being that
the type specimen was 1,200 mm. (about 47 inches), and that an immature male from
northern Argentina, probably of this species, was 1,030 mm. (about 41 inches) long.

Developmental Stages. Not known.

Habits. Nothing whatever is known of the habits of this species.

Range. Western tropical and subtropical Atlantic. The few records that can be re-
ferred to remotus with confidence are for the Antilles (type specimen"'), Rio de Janeiro
(see Study Material, p. 400) and probably northern Argentina.

Synonyms and References:

Carcharias {Prionodon) remotus Dumeril, Hist. Nat. Poiss., j, 1865: 374 (descr., 1.2-m. spec, Antilles);
Gunther, Cat. Fish. Brit. Mus., 8, 1870: 363 (footnote ref. to Dumeril, 1865).

Carcharhinus {Platyfodon) remotus ]oidd,n and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 216 (Mar-
tinique); Bull. U.S. nat. Mus., 47 (l), 1896: 37 (descr., Martinique).

Carcharinus rem.otus Garman, Mem. Harv. Mus. comp. Zool., 56, 1913: 138 (descr. of specs, listed in
Study Material, p. 400, Rio de Janeiro) ; Meek and Hildebrand, Field Mus. Publ. Zool., 75 (l), 1923:
47 (descr., after Garman) ; White, Bull. Amer. Mus. nat. Hist., 7^, 1937: 127 (in Key).

Carcharias remotus Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 16 (W. Indies
and Brazil).

Eulamia remota Fowler, Arqu. Zool. Estado Sao Paulo, 3, 1943, 128 (listed, Brazil).

Probable Reference:

Carcharias lamia Lahille, An. Mus. nac. B. Aires, 34, 1929: 305, pi. 3, lower fig. (north. Argentina, probably
referable to remotus by proportional dimensions of 1,030-mm. male, by ill. of ventral side of head
and absence of black fin markings; but upper teeth, as illustrated, are intermediate between that species
and limbatus) ; Pozzi and Bordale, An. Soc. cient. argent., 120, 1935: 150 (name only, apparently refers
to Lahille's reference) ; not C. lamia Risso, 1826, or Muller and Henle, 1 841.

148. Martinique, according to Jordan and Evermann (Bull. U.S. nat. Mus., 47 [1], 1896: 37).

404 Memoir Sears Foundation for Marine Research

Carcharhinus sfringeri (Bigelow and Schroeder), 1944
Figure 78

Study Material. Female, 805 mm. in total length, from Cozumel, Mexico (U.S.
Nat. Mus., type. No. 37141) ; a somewhat shrivelled skin and head of a female, probably
this species, about 1,390 mm. in total length, from Englewood, Florida (Harv. Mus.
Comp. Zool., No. 35900). These are the only specimens of the species yet seen.

Distinctive Characters. C. springeri most nearly resembles C. obscurus, with which it
agrees generally in the relative size and position of fins and the presence of a mid-dorsal
ridge. But it differs in a number of features from specimens of obscurus of approximately
the same size with which we have compared it; (a) its eye is considerably larger relative to

Figure 78. Carcharhinus sfringeri, female, 805 mm. long, from Cozumel, Mexico (U. S. Nat. Mus.,
No. 37141, type). A Anterior part of head from below, about 0.3 x. B Cross-section of back, midway be-
tween the two dorsal fins, to show the mid-dorsal ridge, slightly enlarged. C Left-hand nostril, about 1.6 x.
D Dermal denticles, about 18 x. £ Upper and lower teeth from right-hand side, about 1.4 x. F Fifth upper
tooth. G Seventh upper tooth. H Second lower tooth. / Ninth upper tooth. F-I, about 2.6 x.

the lengths of the gill openings; (b) the anterior margin of its nostril is expanded as a low,
triangular lobe (not lobed in obscurus) ; (c) its first dorsal is relatively larger and more
erect, but with the free rear corner relatively shorter and the posterior margin less deeply
concave; (d) its second dorsal is larger in area but shorter (from origin to rear tip) rela-
tive to its vertical height; (e) the distance from the tips of the pelvics to the origin of the
anal is considerably shorter, i.e., about 0.7 of the anal base (1.3 times the anal base in obscu-

Fishes of the Western North Atlantic 405

rus); (f) the outer corners of its pelvics are only about at a right angle (about 115° in
obscurus) ; (g) its dermal denticles have a larger number of marginal teeth and ridges in
specimens of equal size; and (h) its upper teeth are more strongly oblique, more deeply
incised outwardly and more coarsely serrate basally than those of obscurus.

Among carcharhinids of the west coast of America, springeri resembles most nearly
platyrhynchus (Gilbert), 1 89 1, in its teeth and fins. But the length of its snout in front of
the mouth is considerably less than the breadth of the mouth (a little longer than breadth of
mouth in platyrhynchus), and its fins show no trace of the white edgings that are so con-
spicuous in platyrhynchus. In combination these differences seem sufficient to demand rec-
ognition in nomenclature, especially in view of the geographic discontinuity between the
areas of occurrence of the two sharks.

Springeri is similar to galapagoensis Snodgrass and Heller, 1905, in the teeth and
snout, but separated from it by the shapes of the first dorsal and pectoral fins, much larger
eye relative to the lengths of the gill openings, and by the fact that its second dorsal is only
very little smaller than its anal in area.

Description. Proportional dimensions in per cent of total length. Female, 805 mm.,
from Cozumel, Mexico (U.S. Nat. Mus., type, No. 37140-

Trunk at origin of pectoral: breadth 12.O; height 1 1.4.

Snout length in front of: outer nostrils 3.1 ; mouth 6.6.

Eye: horizontal diameter 2.1.

Mouth: breadth 8.9; height 5.7.

Nostrils: distance between inner ends 6.$-

Labial furrow length: upper 0.6.

Gill opening lengths: ist 2.3; 2nd 2.6; 3rd 2.7; 4th 2.6; 5th 2.2.

First dorsal fin: vertical height 10.7; length of base 9.9.

Second dorsal fin: vertical height 3.3; length of base 4.5.

Anal fin: vertical height 3.9; length of base 4.9.

Caudal fin: upper mzrgm 29.85 lower anterior margin 14.2.

Pectoral fin: outer margin 19.9; inner margin 5.2; distal margin 15.6.

Distance from snout to: ist dorsal 30.73 2nd dorsal 58.8; upper caudal 70.2; pec-
toral 20.5; pelvics 47.9; anal 58.8.

Interspace between: ist and 2nd dorsals 20.7; 2nd dorsal and caudal 7.7; anal and

caudal 6. i .

Distance from origin to origin of: pectoral and pelvics 27.2; pelvics and anal i i.o.

General form moderately stout. Trunk height at origin of ist dorsal (where highest)
about Vs of length to origin of caudal. Body sector to cloaca somewhat longer than tail
sector. Midline of back with a low but unmistakable dermal ridge between dorsals. In the
preserved state there is also a similar ridge between 2nd dorsal and caudal, but since this
lies along the bottom of a groove of muscular contraction we question whether it is a normal
feature. Upper precaudal pit strongly marked, subangular in outline, the lower pit lunate,
less distinct than upper. Dermal denticles closely and regularly overlapping, broader

4o6 Memoir Sears Foundation for Marine Research

than long, with 5 (sometimes 7) short, marginal teeth and an equal number of low ridges:
in the larger specimen, probably of this species, the denticles usually have 7 ridges and
teeth.

Head to origin of pectoral a little less than V3 of length (29%) to origin of caudal,
moderately flattened above. Snout broadly rounded, its length anterior to a line connecting
outer ends of nostrils a little less than V2 as great as length in front of mouth, the length
in front of mouth nearly % (74%) as great as breadth of mouth or about V3 of length of
head to origin of pectorals. Eye approximately circular, its diameter a little less than Vs
as long as distance between nostrils and about as long as ist gill opening. Third gill opening
(a little the longest) a little more than % (42%) as long as distance between nostrils, the
5th about % as long as 3rd and more oblique than the others, the 3rd gill opening above
origin of pectoral. Nostril strongly oblique, its inner corner nearer to mouth than to the tip
of snout by a distance a little longer than its own length or diameter of eye, its anterior
margin expanded toward inner end as a distinct but low subtriangular lobe. Mouth broad-
ovate, a little less than % (about 62 to 64%) as high as wide.

Teeth -jI^I^yI- ; uppers triangular, ist and 2nd erect and nearly symmetrical, with
concave margins and expanded bases, but subsequent teeth increasingly oblique toward
corners of mouth, their inner margins nearly straight, but their outer margins more and
more deeply concave in subangular contour, the outermost strongly so, the margins regu-
larly serrate from tip to base, most strongly so outwardly on basal sector; lower teeth
nearly erect, except for the outermost 3 or 4, which are moderately oblique, their cusps
much more slender than those of uppers, on broadly expanded bases, their margins much
more finely serrate than those of uppers; one small symmetrical tooth at symphysis, and
outermost 3 to 5 successively smaller in each jaw.

Origin of ist dorsal a little anterior to inner corner of pectoral, its base nearly as long
as from posterior edge of eye to ist gill opening, its vertical height about equal to its base,
its anterior margin only weakly convex toward the apex, the posterior margin only weakly
concave (less so than in obscurus), apex narrowly rounded, the free rear corner only about
Vs to V4 as long as the base (thus relatively shorter than in obscurus), the midpoint of
base about 1.7 times as far from origin of pel vies as from axil of pectoral. Second dorsal
between Yo and V3 as long at base as ist, relatively lower, its vertical height about % as
great as its length at base, its free rear corner a little shorter than its base, its posterior
margin weakly concave, its extreme length from origin to rear tip about 2.5 times as great
as the vertical height (considerably shorter relative to its height than in obscurus) , its origin
about opposite to that of anal. Caudal between Vs and V4 (29.8%) of total length, its
terminal sector about V4 of total length of fin, slender, with narrowly rounded tip and
weakly concave lower posterior outline; the lower lobe (expanded lower anterior corner)
only a little less than V2 (about 47%) as long as upper (somewhat longer, relatively, than
in obscurus), with weakly convex anterior margin, nearly straight posterior margin, and
narrowly rounded tip, the re-entrant corner (included by the 2 lobes) well rounded. Dis-

Fishes of the Western North Atlantic 407

tance from origin of caudal to tip of anal about % as long as base of latter. Anal a little
longer at base than 2nd dorsal and a very little higher vertically, its posterior margin much
more deeply concave, its free rear corner about % as long as base. Distance from origin of
anal to tips of pelvics only about % as long as base of anal. Pelvics about as long at base as
base of anal, or a little longer, their outer corners approximately a right angle (less obtuse
than in obscurus). Pectoral about % as long as head, a little less than V2 as broad as long,
with narrowly rounded tip and inner corner, weakly and evenly convex outer margin, and
distal margin deeply concave proximally.

Color. After preservation the type specimen is olive gray above, and of a paler shade
of yellowish olive below, without any conspicuous fin markings, dark or light.

Size. The fact that the type specimen still shows the umbilical scar, although it is 805
mm. long, suggests that this is one of the larger members of its genus.

Develof mental Stages. Embryos have not been seen as yet.

Habits. Nothing is known of the life history of this newly described species.

Range. C. springeri is known only from Cozumel, east coast of Yucatan, and (prob-
ably) off the west coast of Florida j see Study Material, p. 404.

Synonyms and References:

Eulamia sfringeri Bigelow and Schroeder, Proc. New Engl. zool. CI., 23, 1944: 30, pi. 9, lO (descr., ill.,
Cozumel).

Family SPHYRNIDAE
Hammerhead Sharks

Characters. In general the characters are those of the Carcharhinidae (p. 262), except
that the anterior portion of the head is much flattened dorso-ventrally and very widely ex-
panded laterally in "hammer" or "bonnet" form, with the eyes at its outer edges j and the
skull is modified accordingly, its anterior portion with the olfactory capsules and orbital
region being very widely expanded, and the three rostral bars transversely and broadly
truncate in front at their union. Development viviparous, with yolk-sac placenta in some
species,' but perhaps ovoviviparous in others.

Genera. Two, as indicated in the following Key.

Key to Genera

I a. Nostrils closer to midline of snout than to eyes. Eusphyra Gill, 1862.

Tropical Indian Ocean,
Malaysian region, Indo-
China and northern Aus-
tralia.

lb. Nostrils much closer to eyes than to midline of snout.

Sphyrna Rafinesque, 18 10, p. 408.

I. For the placenta in Eusphyra blochil, see Alcock (J. Asiat. Soc. Beng., 59 [2], 1890: 52).

4o8 Memoir Sears Foundation for Marine Research

Genus Sfhyrna Rafinesque, 1 8 lO

Sfhyrna Rafinesque, Indice Ittiol. Sicil., 1810: 46, 60; type species, Squalus zygaena Linnaeus, 1758,
designated by Jordan and Gilbert, Bui]. U.S. nat. Mus., 16, 1883: 26.

Generic Synonyms:

Cestracion Klein, Pise. Natural. Gedoni, j, 1742: 12; Neuer Schauplatz Natur., 3, 1776: 523 (not seen);

Walbaum, P. Artedi Genera Pise. Emend. Ichthyol., 1792: 580; type species, Squalus zygaena Linnaeus,

1758, designated by Gill, Ann. N. Y. Lye, 7, 1862: 403.^
Sphyrnias Rafinesque, An. Nature, 1 81 5: 93; substituted for Sfhyrna Rafinesque, 1810.
Cestrorhinus Blainville, Bull. Soc. philom. Paris, 1816: 121; type species, Squalus zygaena Linnaeus, 1758,

designated by Fowler, Bull. geol. Surv. N. Jersey, 4, 191 1 : 77.
Zygaena Cuvier, Regne Anim., 2, 1817: 27; type species, Squalus zygaena Linnaeus, 1758, preoccupied by

Zygaena Fabricius, 1775, for Lepidoptera.
Zygoena Risso, Hist. Nat. Europe Merid., 5, 1826: 125; emended spelling for Zygaena Cuvier, 1817.
Sphyrnichthys Thienemann, Lehrb. ZooL, 1828: 408; substituted for Sfhyrna Rafinesque, 1810.
Zygana Swainson, Nat. Hist. Fish. Amphib. Rcpt., 2, 1839: 318; emended spelling for Zygaena Cuvier, 1 81 7.
Platysqualus Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1839: 318; type species, Squalus tiburo Linnaeus,

1758.
S-phyra Van der Hoeven, Handb. Dierkunde, 2, 1858: 68; emended spelling for Sfhyrna Rafinesque, 1810.
Reniceps Gill, Ann. N. Y. Lye, 7, 1862: 403, 412 ; type species, Squalus tiburo Linnaeus, 1758.

Generic Characters. Nostrils much closer to eyes than to midline of snoutj jaws with
or without labial furrows; anterior margin of nostril expanded at inner end as a stiff,
triangular flap, hollowed on lower side and overhanging the orifice; ist dorsal narrow-
triangular; 2nd dorsal much smaller than ist dorsal; caudal with well marked subterminal
notch, its lower anterior corner expanded as a definite lobe. Characters otherwise those of
the family.

Range. Tropical to warm temperate zones of all oceans, including the Mediter-
ranean.

Fossil Teeth. Upper Cretaceous to Miocene, North America; Miocene, Africa; Mio-
cene to Pliocene, Europe.

A ttacks on Bathers. The larger Hammerheads have long borne an unsavory reputa-
tion as "man-eaters," partly on the basis of unverifiable rumor, and partly because of the
fact that a large specimen taken many years ago oflF Long Island, New York (hence prob-
ably zygaena)^ contained portions of a man in its stomach."" Positive evidence is now at
hand that this reputation is deserved, for on Sept. 21, 19 13, a Hammerhead about eight
feet long (species not determined) attacked and so seriously injured a bather at West
Palm Beach, Florida, that the lacerations required some 200 stitches. However, the victim
recovered after seven weeks in the hospital.*'' Attacks on bathers by Hammerheads (nomi-
nally, at least, zygaena) have also been reported from British Guiana. Hammerheads
(probably 5. lewini) are also considered very dangerous in Australian waters,^ where
shark fatalities are of much more frequent occurrence than anywhere in the western
Atlantic.

2. By Ruling 89 of the International Commission on Zoological Nomenclature (Smithson. Misc. Coll., 75, 1925 : 27)

Klein's names are not to be taken into account.
2a. Mitcliill, Trans. Lit. Phil. Soc. N. Y., /, 1815: 482.
lb. Gudger, Bull. Amer. Mus. nat. Hist., 40, 1937: 417. 3. Coppleson, Med. J. Aust., April 15, 1933: 59.

Fishes of the Western North Atlantic 409

Sfecies. It was long thought that this was a very monotonous genus Including some
four or five species at most the world over. However, recent studies have shov/n that
the western Atlantic alone actually supports at least five well marked representatives,
separated by well defined and easily detectable characters, but which are so overshadowed
by the bizarre appearance of the head that they were largely overlooked in most of the
early accounts of the genus. Three new species have been described recently from the
eastern tropical Pacific also,* while the remaining sphyrnids of the Indo-Pacific region
as a whole stand in urgent need of critical revision. Unfortunately many of the older
descriptions, other than those of S. tiburo, which is the most easily recognizable member
of the genus in the Atlantic, omit precisely those characters that have recently been found
to be specific. Hence there is no knowing to which species, as now recognized, they actually
referred. The case is still further complicated by the fact that opportunity has not yet
been offered for a sufficiently extensive comparison of the species now known to exist in the
Atlantic with those of the Pacific and Indian Oceans. Consequently, the accompanying
Key is restricted to Atlantic species. Fortunately it is clear to which of the Atlantic forms
the Linnaean name zygaena (type species of the genus) actually referred, because
WillughbyV illustration of the lower surface of his zygaena, to which Linnaeus refers as
one of the bases of the species, is an excellent representation of a Hammerhead with head
rounded in front, with eyes close to corners of oculo-nasal prominences, and with long
caudal peduncle.

Key to Atlantic Species

I a. Anterior contour of midsector of head evenly rounded or nearly straight; not in-
dented or scalloped in median line (Figs. 82 B, 86 A).

2a. Contour of head only slightly concave opposite nostrils, if at all (Fig. 82 B) 3
groove from nostril, if any, shorter than horizontal diameter of eye; free tip of
2nd dorsal not longer than its anterior margin; posterior margin of anal only
weakly concave; teeth near corners of mouth rounded, without cusps.

tiburo Linnaeus, 1758, p. 420.
2b. Contour of head deeply scalloped opposite nostrils (Fig. 86 A); grooves from
nostrils more than twice as long as horizontal diameter of eye; free tip of 2nd
dorsal considerably longer than its anterior margin; posterior margin of anal
deeply concave; teeth near corners of mouth like those further forward, with
cusps. zygaena Linnaeus, 1758, p. 436.

lb. Anterior contour of head unmistakably indented or scalloped in midline.

3a. Free tip of 2nd dorsal only about as long as its vertical height, and considerably
shorter than its anterior margin; teeth serrate on cusps as well as basally.

/«i^j Valenciennes, 1822, p. 428.

4. Vesfertlna, media and corona Springer (Stanford Ichth. Bull., i [5], 1940: 161-169).

5. Hist. Pise, 1686: pi. B, I.

4IO

Memoir Sears Foundation for Marine Research

3b. Free tip of 2nd dorsal considerably longer than its vertical height, and at least

as long as its anterior margin; teeth with smooth-edged cusps, serrated only on

basal expansions, if at all.

4a. Center of eye opposite or posterior to front of mouth; corner of mouth an-
terior to outer posterior corner of head (hammer) ; posterior margin of anal
fin deeply concave. diplana Springer, 1941, p. 415.

4b. Center of eye considerably anterior to front of mouth ; corner of mouth con-
siderably posterior to outer posterior corner of head (hammer) ; rear mar-
gin of anal only weakly concave. bigelowi Springer, 1944, p. 410.

Sfhyrna bigelowi Springer, 1 944
Figures 79, 80

Study Material. Immature male, about 886 mm. long, from Rio de Janeiro, Brazil
(Harv. Mus. Comp. Zool., No. 463) ; young male, about 385 mm., from Uruguay (U.S.
Nat. Mus., type, No. 87682).

Distinctive Characters. This newly discovered species falls with diplana and tudes

Figure 79. Sfhyrna bigelowi, immature male, about 886 mm. long, from Rio de Janeiro, Brazil (Harv. Mus.
Comp. Zool., No. 463). A Dermal denticles, about 22 x. 5 Dermal denticle, side view, about 44 x. C Dermal
denticle, apical view, about 44 x. D Upper and lower teeth, left-hand side, about 2 x. E Fifth upper tooth.
F Twelfth upper tooth. G Fifth lower tooth. H Twelfth lower tooth. E-H, about 4 x.

Fishes of the Western North Atlantic

411

in that the anterior contour of the head is unmistakably indented in the midline. But it
differs from tudes in the much more definitely marked prenarial groove, more erect first
dorsal, and less deeply concave second dorsal, as well as in the fact that the teeth are
smooth-edged (serrate in tudes) with the lowers noticeably more slender and more erect
than the uppers. The most obvious diflFerences from diplana are: eyes relatively much
smaller, the free rear tip of the second dorsal not longer than the anterior margin of
latter (twice that long in diplana), origin of pelvics almost under rear tip of first dorsal
(considerably behind it in diplana) and the anal less deeply concave and considerably
longer than the second dorsal. It is separated from zygaena by: the relatively long,
narrow hammer which is scalloped anteriorly in the midline, the very short free rear
corner of the second dorsal, the lower teeth which are much more slender and oblique than
the uppers, and the anal fin which is much longer than the second dorsal. The shape of the
head separates it from tiburo.

Figure 80. Sphyrna bigelouii, illustrated in Fig. 79. Head from below, about 0.6 X.

412 Memoir Sears Foundation for Marine Research

Description. Proportional dimensions in per cent of total length. Male, 886 mm.,
from Rio de Janeiro (Harv. Mus. Comp. Zool., No. 463).

Trunk at origin of pectoral: breadth 7.9; height 10.7.

Snout length in front of: outer nostrils 3.3; mouth 7.1.

Eye: horizontal diameter i.O.

Mouth: breadth 6.3; height 3.5.

Nostrils: distance between inner ends 18.1.

Gill opening lengths: ist 3.2; 2nd 3.45 3rd 3.7; 4th 3.5; 5th 2.9.

First dorsal fin: vertical height 14.7; length of base 9.8.

Second dorsal fin: vertical height 4.1; length of base 4.1.

Anal fin: vertical height 5.15 length of base 8.5.

Caudal fin: upper margin 29.5; lower anterior margin i i.i.

Pectoral fin: outer margin 13.9; inner margin $.6-j distal margin 14.0.

Distance from snout to: ist dorsal 27.5; 2nd dorsal 59.5} upper caudal 70.55

pectoral 21.3; pelvics42.5; anal 55.3.

Interspace between: ist and 2nd dorsals 21.2; 2nd dorsal and caudal 7.6; anal and

caudal 5.8.

Distance from origin to origin of: pectoral and pelvics 20.3; pelvics and anal 14.6.

Trunk about Vs as high at origin of ist dorsal as its length to origin of caudal j more
strongly compressed than in other local species. No mid-dorsal ridge. Caudal peduncle
about % as wide as deep, the upper precaudal pit strongly developed as a narrow, trans-
verse, lunate furrow, the lower pit similar but much smaller. Dermal denticles evenly
and closely spaced, overlapping but little, the blades thick and rather strongly arched,
broader than long, strongly sculptured, usually with 5 (occasionally only 3) high, sharp-
topped ridges separated by V-bottomed valleys, the marginal teeth broad and short, the
axial only very little the longest; pedicels long, rather slender, the four corners of bases
short.

Head in front of pectoral a little less than Vs of length of trunk to origin of caudal,
its dorsal profile sloping steeply, very thin anterior to eyes, hammer-shaped, the outer
posterior margins nearly straight and transverse or even sloping slightly forward (much as
in tudes) ; the breadth of head at eyes about 1.2 times its length to origin of pectorals, and
about 3.3 times its length at oculo-narial prominence; anterior margin of hammer
scalloped with a deep, rounded indentation in midline (as in diplana\ a shallower
indentation opposite each nostril (much shallower than in diplana or zygaena) and a
still more shallow sinuosity between the two others; a well marked groove running inward
from corner of nostril along anterior margin of head about Vs of distance toward mid-
line (much more strongly marked than in tudes). Distance from anterior margin of eye
to anterior corner of oculo-narial prominence about equal to diameter of eye; a line con-
necting the inner ends of nostrils passes about midway between front of mouth and an-
terior margin of head, one through center of eyes passes anterior to mouth by a dis-

Fishes of the Western North Atlantic 413

tance about twice as long as diameter of eye, and one connecting the corners of the
hammer passes a little posterior to the corners of the mouth in very small specimens but
a little anterior to them in larger, an alteration that results from an increase in the breadth
of the hammer relative to its length with growth, such as takes place in tudes also.
Mucous pores on lower side of head in midline cover a trapezoidal area (Fig. 80). Eye
a little broader than high, much smaller than in the diplana-zygaena group, its hori-
zontal diameter only about ^^ as long as head in front of mouth. Gill openings extending
ventrally and spaced nearly equally," the first Vs to V2 (41%) as long as head in front of
mouth and about 3.2 times as long as diameter of eye, the 3rd hardly longer than ist, and
5th very little shorter and about over origin of pectoral. Nostril transverse. Mouth
strongly arched, its breadth about 1.8 times its length, without definite labial furrow on
either jaw.

Teeth |^^~|^ in specimen counted, smooth-edged and narrow-triangular on ex-
panded bases J uppers oblique, deeply notched outwardly and increasingly so toward
corners of mouth, the 4th or 5th to nth largest, the outermost 2 very low but still
with definite cuspj lower teeth with much narrower cusps, erect toward center of mouth
but somewhat oblique toward corners, the basal expansion somewhat swollen outwardly as
a rounded boss or obscure denticle on each side on median tooth and on a few teeth next
to the latter}' the 2nd to 7th or 8th longest, the outermost 2 very short, rounded, without
definite cusp (as in tiburo) ; 2 small symmetrical teeth at symphysis on upper jaw and
I on lower; i or 2 series of teeth functional in alternating rows along sides of upper jaw
and 2 to 3 rows along sides of lower.

First dorsal at base a little less than V2 as long as head in front of origin of pec-
torals, its vertical height about i V2 times the base, erect, a perpendicular from its apex
falling through rear end of base or a little anterior to latter; its origin about over mid-
point of inner margin of pectoral, its anterior margin only slightly convex, rear margin
moderately concave toward base, apex subacute, its free rear corner about Y2 as long as
base, the midpoint of base only a little nearer to axil of pectoral than to origin of pelvics
with the rear tip about over the latter. Second dorsal at base a little more than Vs as long
as 1st and a little less than Vs as high vertically, its origin about over midpoint of base of
anal, its posterior margin moderately concave (much less deeply so than in tudes), and
apex narrowly rounded, its free rear corner only moderately slender, about i Yo times as
long as the base. Caudal a little less than Y3 (29%) of total length, the terminal sector
about Yi the length of fin and broadly triangular (considerably larger than in di-plana,
zygaena or iudes), with nearly straight lower posterior margin and subacute tip; the
lower lobe a little more than Vs (37%) as long as upper, with weakly convex anterior
margin, nearly straight posterior margin and subacute tip; the re-entrant contour (enclosed

6. The g86-mm. specimen is asymmetrical as re^rds its gill slits, the 4.th and 5th coming close together at the lower
ends on one side of the head but evenly separated from end to end on the other.

7. On the type specimen the first to fourth lower teeth on each side have these denticles more or less developed; in
the 8g6-mm. specimen the first right-hand and second left-hand lower teeth have one on each side, end the first
left tooth has one on one side only.

414- Memoir Sears Foundation for Marine Research

by the 2 lobes) well rounded and a little more than a right angle. Distance from origin of
caudal to tip of anal a little less than V3 (30%) as long as base of latter. Anal a little more
than twice as long as 2nd dorsal at base but only about as high vertically, its anterior margin
weakly convex, its posterior margin deeply concave near apex but nearly straight thence to
tip, its apex subacute, its free rear corner about ^/^ as long as the base. Distance from origin
of anal to tips of pelvics a little less than % (61%) as long as base of anal. Pelvlcs with
nearly straight edges, rather broadly rounded apices and narrowly rounded tips, their
origin about under rear tip of ist dorsal. Pectoral larger than in diplana, tudes or zygaena
{bigelowi more nearly resembles tiburo in this respect), its length about % (65%) that
of head or about equal to vertical height of ist dorsal, about % as broad as long, its outer
margin nearly straight basally but convex toward apex, the distal margin only very weakly
concave or perhaps nearly straight, its apex subacute, the Inner corner very narrowly
rounded.

Color. The preserved specimens we have studied are grayish brown above and of a
paler tint of the same hue below, the fins without conspicuous markings. In the type speci-
men the anterior margin of the hammer is bordered with yellowish. No information is
at hand as to the color of this Hammerhead in life.

Size. The state of sexual development of the 886-mm. specimen, on which the
claspers extend a little beyond the tips of the pelvlcs, suggests that this Is a rather small
species, perhaps becoming mature when only four to five feet long.

Developnental Stages. Embryos have not been seen.

Habits. Nothing Is known of its habits.

Range. So far S. btgelowi is known only from Uruguay (type locality) and from
Rio de Janeiro, Brazil. Probably It is a tropical species and therefore watch should be kept
for it in the West Indian-Caribbean region, in the Gulf of Mexico, and around southern
Florida, as well as along the northern and northeastern coasts of South America.

Remarks. This little known species seems closest to S. corona Springer, 1940, and S.
media Springer, 1940, of the Pacific coasts of Central and South America.'^ It is separated
from corona by its more broadly rounded mouth, its longer head in front of the mouth
relative to the distance between the nostrils (head length in front of mouth about 40 per
cent of distance between nostrils in bigelowiy but S5 per cent in corona), the much more
deeply indented anterior outline of its head in the midline, and its much more strongly
developed prenarial grooves. It diflFers from media, with which it shares the broadly
rounded mouth, in the anterior outline of its head (rounded in media), in its more erect
lower teeth, and in the fact that the distance between eye and nostril is relatively greater.

Synonyms and References:

Sfhyrna tudes (in part) Garman, Mem. Harv. Mus. comp. Zool., j(5, 1913: 159 (Carman's description was

based in part on the spec, descr. and pictured here) ; not S. tudes Valenciennes, 1 822.
Sfhyrna bigelowi Springer, J. Wash. Acad. Sci., 5^, 1944: 274 (descr., ill., Uruguay).

7a. For accounts of these, with illustrations of their heads, see Springer (Stanford Ichthyol. Bull., i [5], 1940:
162, 163, fig. 3, 4).

Fishes of the Western North Atlantic 415

S-phyrna di-plana Springer, 1 94 1

Hammerhead

Figures 1 2 A, 81

Study Material. Eight males and females, 470 to 639 mm. long, and the head of
another, about 1,200 mm. (calculated from size of head), from Rio de Janeiro and Rio
Grande do Sul, Brazil, and from the vicinity of Galveston, Texas, and male, about
1,340 mm., labelled "Europe" (Harv. Mus. Comp. Zool.). Male, 445 mm., and fe-
male, 504 mm., from Galveston, Texas; four specimens, two to three feet, from Colon,
and from Charleston, S. Carolina; head of one, about 1,375 mm. (calculated from breadth
of head), from Englewood, Florida; jaws of 1,850 mm. and 2,500 mm. specimens from
that same locality (U.S. Nat. Mus.). Also, photographs of head, region of caudal pe-
duncle, and second dorsal and anal fins, of a large Florida specimen (from Stewart
Springer).

Figure 8i. Sfhyrna diflana, female, 639 mm. long, from Rio de Janeiro, Brazil (Harv. Mus. Comp. Zool.,
No. 462). A Dermal denticles, about 55 x. S Left-hand upper and lower teeth, about 0.7 x natural size.
C Fourth upper tooth. D Twelfth upper tooth. E Third lower tooth. F Eleventh lower tooth. C-F, about 1.4 x.
G Head from below.

Distinctive Characters. Diflana has long been confused with zygaena, but it is easily
distinguished from the latter by the facts that its head is scalloped in front at the midline
(rounded in zygaena), that its eyes are farther from its nostrils, and by the much shorter
interspace between the rear tip of its second dorsal and the origin of its caudal (cf. Fig.

41 6 Memoir Sears Foundation for Marine Research

8 1 with 86 B). It shares a median indentation in the anterior outline of the head with both
tudes and bigelowi, but it is separable from them at a glance by the facts that the corners
of its mouth are considerably anterior to the outer rear corners of the head (hammer) and
that the free rear tip of its second dorsal is considerably longer than the anterior margin of
the fin J it is further separated from tudes by its smooth-edged teeth. The hammer shape of
its head obviates any danger of confusing it with tiburo.

Description. Proportional dimensions in per cent of total length. Female, 554 mm.,
from Galveston, Texas (Harv. Mus. Comp. Zool., No. 35826). Female, 639 mm., from
Rio de Janeiro (Harv. Mus. Comp. Zool., No. 462).

Trunk at origin of -pectoral: breadth 9.4, 8.55 height 1 1.9, 1 1.6.

Snout length in front of: outer nostrils 4.4, 4. i ; mouth 7.4, 7. i .

Eye: horizontal diameter 2.3, 2.4.

Mouth: breadth 6.8, 7.0 ; height 3.8, 3.8.

Nostrils: distance between inner ends 20.8, 20.2.

Gill opening lengths: ist 3.6, 3.7; 2nd 3.6, 3.7J 3rd 3.5, 3.7; 4th 3.2, 3.5; 5th

2.5,2.8.

First dorsal fin: vertical height 13.4, 12.7 j length of base 12.4, i i.o.

Second dorsal fin: vertical height 2.9, 3.35 length of base 4.1, 3.1.

Anal fin: vertical height 3.4, 3.3 ; length of base G.t^, 5.3.

Ca«^a/^«; upper margin 33.4, 3 1.0 J lower anterior margin 11. 7, 12.0.

Pectoral fin: outer margin 13.2, 13.OJ inner margin 4.9, 5.OJ distal margin, lO.O,

11.3.

Distance from snout to : ist dorsal 26.4, 28.2; 2nd dorsal 56.O, 6o.2} upper caudal

66.6., 69.O} pectoral 22.4, 22.9; pelvics 41.3, 46.6; anal 55.O, 57.2.

Interspace between: ist and 2nd dorsals 20.6, 23.3; 2nd dorsal and caudal 7.4,

6.7} anal and caudal 6.3, S-(>-

Distance from origin to origin of: pectoral and pelvics 20.6, 22.7; pelvics and anal

12.8, II. 7.

Trunk strongly compressed, its height at ist dorsal about Y^ its length to origin of
caudal, with moderately arched dorsal profile. Back without mid-dorsal ridge. Body sec-
tor to cloaca, a little longer than tail sector. Caudal peduncle about % as wide as deep, the
upper precaudal pit strongly marked, subrectangular in outline, the lower pit only weakly
indicated as a short, semilunar furrow. Dermal denticles much as in zygaena, partly over-
lapping and with skin partly exposed, the blades thin, moderately arched, small specimens
usually with 3, but large with 4 or 5, sharp-topped ridges running back about half the
length of the blade, the marginal teeth much as in zygaena, the axial a little the longest;
pedicels very short, moderately slender.

Head about V3 of length to origin of caudal, sloping evenly forward to anterior mar-
gin, very broadly expanded in hammer form but with posterior edges (outward from
the neck) sloping only a little rearward; the breadth at eyes about 1.2 to 1.3 times as great

Fishes of the Western North Atlantic 417

as its length to origin of pectorals, and a little more than 3 times its length at oculo-narial
prominence} anterior margin of head scalloped with a deep rounded depression opposite
nostril, a somewhat shallower indentation in the median line, and a still shallower sinuosity
midway between these two, also with a well marked groove running from the nostril in-
ward along anterior margin of head for about 40% of the distance toward the midline (a
little farther than this in zygaena). Distance from anterior corner of oculo-narial promi-
nence to anterior margin of eye about as long as diameter of eye (only about V2 to Vl 0 that
long in zygaena) ; a line connecting outer ends of nostrils passes anterior to front of mouth
by a distance about as long as horizontal diameter of eye, one connecting centers of eyes
passes about through front of mouth, and one through the outer posterior corners of ham-
mer passes a little posterior to corners of mouth. Head (snout) in front of mouth a little
less than ^ as long as length to origin of pectorals. Rostral cartilage with a median oval
hole, the wings of the preorbital processes with an inwardly directed point on anterior
margin (this hole and point usually lacking in zygaena). Mucous pores in median sector
of oral side of head near its anterior margin cover a subrectangular or dumbbell-shaped
area (a subtriangular area in zygaena). Eye approximately circular, its diameter about
% as long as head in front of mouth. Gill openings noticeably longer than in zygaena, the
1st a little more than lYz ( i.6 to 1.7) times as long as diameter of eye, the 5th about 80%
as long as ist, their outlines evenly and moderately concave, the space between 4th and 5th
over origin of pectoral. Nostril nearly transverse. Mouth strongly arched, about V2 (49
to 56%) ashighasbroad. Labial furrows on lower jaw, about '^A to ^5 as long as horizontal
diameter of eye and concealed when mouth is closed, but none on upper jaw.

Teeth '^ °' '^~° '° ^~'^ " 'I , triangular on expanded bases, the cusps smooth-edged, but

15 or 16 — ^1 or 2 — 15 orlo'" r ' i o»

the bases more or less wavy or fluted on some of the teeth; ist 3 upper teeth nearly sym-
metrical and erect, but subsequent teeth increasingly oblique toward corners of mouth,
their inner margins more nearly straight, the outer margins more and more deeply
notched; the ist tooth smaller than 2nd, the 15th and i6th very small; lower teeth with
somewhat narrower cusps than uppers, similarly oblique and notched outwardly in em-
bryos' and in small specimens generally, but with successive series tending to become more
erect and their cusps relatively narrower with growth, although there is considerable
variation in this respect."

First dorsal erect, a perpendicular from the apex passing close behind rear end of
base, its vertical height about % (57 to 66%) as great as length of head to origin of pec-
torals with its length at base only a little less, its origin about over midpoint of inner
margin of pectoral, its anterior margin rather strongly convex toward apex, posterior mar-

8. Personal communication from Stewart Springer.

9. In the head of a Florida specimen, about 1,375 "^"i- lo^S (U.S. Nat. Mus., No. 1 10296, length calculated from
the breadth of the head), the first to eighth lower teeth are nearly erect with narrow triangular cusps, and the
lower teeth of the type specimen appear to have been similar. In another head of about the same size from Braz.l,
however, which is otherwise indistinguishable (Harv. Mus. Comp. Zool., No. 8+5), only the first and second lower
teeth are of this type, the third and subsequent teeth being increasingly oblique, and intermediate stages are shown
by other specimens. It is not yet known whether the variations in this respect are individual or racial.

41 8 Memoir Sears Foundation for Marine Research

gin moderately concave, the free rear tip not very slender, about V3 as long as the base; the
midpoint of base only about Yo as far from axil of pectoral as from origin of pelvics and
the rear tip anterior to latter by a distance about Vs as long as the base (this distance is con-
siderably greater relatively in zygaena). Second dorsal about Vs as long as ist at base but
only about V5 as high, its origin about over midpoint of base of anal, with narrowly rounded
apex and only weakly concave posterior outline, its noticeably slender free rear corner
nearly twice as long as base and about twice as long as the vertical height. Distance from
tip of 2nd dorsal to precaudal pit a little less than Vs as long as base of 2nd dorsal (nearly
or quite as long as base of 2nd dorsal in zygaena). Caudal a little less than V3 (about 3 1 to
33%) of total length, its upper margin only very slightly convex, the terminal sector about
Vs the length of fin, narrow-triangular, with subacute tip and rather deeply concave lower
posterior margin; the lower lobe a little more than Ys as long as upper with weakly convex
anterior and posterior margins and subacute tip; the re-entrant contour (included by the
two lobes) with rather abruptly rounded corner and a little more than a right angle. Dis-
tance from origin of caudal to tip of anal a little less than V2 as long as base of anal. Anal
about 1 .4 to 1 .7 times as long at base as 2nd dorsal, about i .0 to i .4 times as high vertically,
its anterior margin much more convex, posterior margin more deeply concave, apex acute,
free rear corner less slender than that of 2nd dorsal, about % as long as the base, its tip
considerably anterior to that of 2nd dorsal. Distance from origin of anal to tips of pelvics
only about V2 as long as base of anal. Pelvics with nearly straight margins and narrowly
rounded corners, and about as long at base as anal. Pectoral a little less than % (about 57
to 61%) as long as head, a little more than V2 (about 56%) as broad as long, the outer
margin moderately convex toward tip, distal margin weakly and uniformly concave, the
apex and inner corner very narrowly rounded.

Color. Light gray above shading to white below, the pectorals tipped on their ventral
surfaces with black in life.

Size. The Study Material (p. 415) suggests that diplana may be born at a length
no greater than 400 to 450 mm. Males mature at about 1,800 mm. (about 6 feet), and
grow to at least lO feet; it is not known how much larger.

Developmental Stages. Embryos have not been described as yet, except for the teeth
(see p. 417), nor have we seen any.

Habits. Recognition of the fact that diplana is distinct from zygaena is so recent^" that
no attention has yet been devoted to its life history as contrasted with that of zygaena.
Neither do the few records of Hammerheads that can be positively referred to it con-
tain any pertinent information. It is probable, however, that the account of the habits, food,
etc., of zygaena (p. 441) applies equally to diplana, at least in a general way. Gravid
females have been taken in southeastern Florida waters."

Range. Tropical and warm-temperate Atlantic, probably including the Mediter-

10. It was unmistakably pictured by Valenciennes more than a century ago (Mem. Mus. Hist. nat. Paris, 9, 1822:
pi. II, fig. i) as Zygaena malleus.

11. Personal communication from Stewart Springer.

Fishes of the Western North Atlantic 419

ranean. It is represented in the tropical-subtropical waters of the eastern and western Indo-
Pacific by a form {S. lewini Griffith, 1834) closely resembling diplana in form of head
(including arrangement of mucous pores) and in the shape and relative position of the fins.
We have not been able to find any significant differences between small specimens from
Panama, southern California, Hawaiian Islands or Celebes^" and the Atlantic series of
comparable sizes listed above (p. 415). However, since Whitley^' describes the teeth of
the Australian lewini as becoming "finely denticulated" with growth, which is not the case
in the Atlantic diplana, it seems wise to retain both specific names, awaiting a comparison
of adult specimens.

Occurrence in the Atlantic. Diplana was separated from zygaena so recently that very
few reports of it have yet appeared under its own name. Information as to its occurrence
in the eastern Atlantic is confined to the facts that Valenciennes^* described his malleus^
which his illustration shows to be the head of diplana, as Mediterranean and Atlantic;
that Springer" reports a head of diplana from tropical West Africa (Gold G)ast) ; that the
collection of the British Museum contains specimens apparently of this species in addition
to zygaena from the Mediterranean ; and that there is a specimen of it labelled "Europe"
in the collection of the Harvard Museum of Comparative Zoology. Locality records that
can be referred with certainty to diplana in the western Atlantic are Rio Grande do Sul,
Rio de Janeiro and probably Pernambuco, Brazil, Colon, both coasts of Florida, South
and North Carolina, and a station about 90 miles off Cape May, New Jersey. These are
enough to show that its range closely parallels that of zygaena. It is so common off south-
eastern Florida that we have recently received a report of 1 9 adult males taken there in a
single day. It appears not to range as far north along the United States coast during sum-
mer as zygaena does. Similarly it is possible that its range may not extend as far to the
south in the southern hemisphere, although information of its presence is so scant for the
South American coast as to preclude any definite statement in this regard. Neither is any
information available as to its abundance relative to that of zygaena anywhere off the
American Coast.

Synonyms and References:

Shark, no name, Marcgrave, Hist. Nat. Brazil, 1648: frontispiece (this is the earliest illustration that we have

found of a Hammerhead with head of the diflana shape).
Le Marteau (in part), Duhamel, Traite Gen. Peches, (2) j (9), 1777: 303, pi. 21, fig. 3—7 (ill., apparently

this species) ; Broussonet, Mem. Acad. Roy. (1780), 1784: 66l (by ref. to Duhamel, 1 777, as above).
Zygaena malleus (in part) Valenciennes, Mem. Mus. Hist. nat. Paris, 9, 1822: 223, pi. 11, fig. I (descr., ill.

of head, Medit., Atlant., but apparently confused with his own tudes, because teeth descr. as denticulate) ;

Risso, Hist. Nat. Europ. merid., j, 1826: 125 (part, by ref. to Valenciennes, 1822); Storer, Mem. Amer.

Acad. Arts Sci., N. S. 2, 1846: 508 {malleus Valenciennes, 1822, incl. in synon.) ; Giinther, Cat. Fish.

Brit. Mus., 8, 1870: 381 (OTa//e«/x Valenciennes, incl. in synon.); Day, Fish. Gt. Brit., 2, 1880-1884:

294 (ot<j//^«x Valenciennes, incl. in synon.), pi. 154 (ident. by shape of head, no loc.) ; Doderlein, Man.

12. Specimens in the collection of Harvard Museum of Comparative Zoology, 480 to 740 mm. long.

13. Fish. Aust., /, 1941: 121. 14. Mem. Mus. Hist. nat. Paris, p, 1822: 213.
15. Proc. Fla. Acad. Sci., 5, 1941 : 49.

42 o Memoir Sears Foundation for Marine Research

Ittiol. Medit., 2, 1881: 46 {malleus Valenciennes, incl. in synon.) ; not Squdus malleus Shaw and
Nodder, 1796.

Sfhyritias zygaena (in part) Gray, List Fish. Brit. Mus., 1 851: 48 {malleus Valenciennes, incl. in synon.) ;
White, List Spec. Brit. Mus., Fish., 8, 1851: 126 {malleus Valenciennes, incl. in synon.).

Sfhyrna zygaena (in part) Ribeiro, Arch. Mus. nac. Rio de J., 14, 1 907: 202 {malleus Valenciennes, incl. in
synon.); Nichols and Murphy, Brooklyn Mus. Sci. Bull., 3 (l), 1916: 19 (ill. shows diflana shape of
head, no loc, but discussion refers to zygaena) ; Springer, Proc. Fla. Acad. Sci., 5, 1939: 31 (combined
with zygaena in descr., habits, ident. by shape of head in ill., Florida) ; Fowler, Bull. U.S. nat. Mus., lOO
{23), 1941: 217 (OTa//i!a/ Valenciennes, incl. in synon.).

Cestracion zygaena (in part) Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 157 {malleus Valenciennes,
incl. in synon.) ; Radcliffe, Bull. U.S. Bur. Fish., 34, 1916: 263, pi. 43, fig. I (N. Carolina spec, ident.
by ill. of head) ; Meek and Hildebrand, Field Mus. Publ. Zool., 15 (1), 1923: 58 {malleus Valenciennes,
incl. in synon.; ident. of spec, from Colon, whether diflana or zygaena, not certain from descr.); not
Squalus zygaena \Jmniitxis, 1758.

Hammerhead Shark, Smith, J. Amer. Mus. nat. Hist., 16, igi6: 348 (photo of lo-foot spec, descr. as Cestra-
cion zygaena by Welsh, 1 91 6).

Cestracion zygaena Welsh, Copeia, 38, 1 916: 94 (meas. of spec, shown in photo by Smith, 1 916, as above;
90 miles off C. May, N. Jersey) ; not Squalus zygaena Linnaeus, 1758.

Sfhyrna diflana Springer, Proc. Fla. Acad. Sci., 5, 1 941: 46 (descr., ill., discuss., both coasts of Florida,
Mississippi, Louisiana, Texas, trop. W. Afr., and probably Medit.) ; Bigelow and Schroeder, Guide
Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 136, fig. 50 (descr., range, ill.).

Doubtful References:

Sfhyrna tudes Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907: 157, pi. 5 (ident. probable because of shape of

head in photo, Maria Farinha, near Pernambuco, Brazil); Fauna brasil. Peixes, 2 (i) Fasc. I, 1923: 14

(same as foregoing) ; not Zygaena tudes Valenciennes, 1822.
Sfhyrna tiburo (in part) Puyo, Bull. Soc. Hist. nat. Toulouse, yo, 1936: 82 (head ill. of diflana shape, but

discuss, apparently of tiburo); not Squalus tiburo Linnaeus, 1 75 8.

Sfhyrna tiburo (Linnaeus), 1758

Bonnet Shark, Shovel Head

Figure 82

Study Material. Seventy-one specimens, embryos to adults, 217 to 1,090 mm. long,
including one female containing eight young nearly ready for birth and another with nine,
from: Hampton Roads, Virginia} Charleston, South Carolina; Tortugas, Florida; Barra-
taria Bay and Bayou Fifi, Louisiana; Galveston, Corpus Christi and Harbor Island, Texas;
Cuba; Bahia, Pernambuco, Rio Parahyba and Rio de Janeiro, Brazil; also, from Pacific
coast of Panama and San Diego, California (Harv. Mus. Comp. Zool. and U.S. Nat.
Mus.). Also, paratypes of 5. ves-pertina Springer, Pacific Panama, loaned by the Carnegie
Museum of Pittsburgh and Stanford University, California.

Distinctive Characters. Tiburo is most obviously marked oflF from diflana, bigelowi,
tudes and zygaena in having a shovel- (not hammer-) shaped head not indented margin-
ally opposite the nostrils; also, the outermost 4 or 5 teeth in the lower jaw are evenly
rounded, without cusps or a definite cutting edge. It is separated from bigeloim, diflana

Fishes of the Western North Atlantic

421

and tudes by the evenly rounded anterior outline of its head, and further from tudes by its
smooth-edged teeth.

Description. Proportional dimensions in per cent of total length. Female, 621 mm.,
from Galveston, Texas (Harv. Mus. Comp. Zool., No. 36157). Male, 812 mm., from
Tortugas, Florida (Harv. Mus. Comp. Zool., No. 848).

Trunk at origin of pectoral: breadth 8.7, 8.9; height lO.O, 8.4.

Snout length in front of: outer nostrils 4.0, 4.1 5 mouth 8.0, 6.6.

Eye: horizontal diameter 1.8, 1.5.

Mouth: breadth 7.2, 6.O5 height 3.8, 3.5.

Nostrils: distance between inner ends 12.2, 9.9.

Labial furrow length: lower 0.7, 0.5.

Gill opening lengths: ist 2.7, 2.2; 2nd 3.2, 2.5; 3rd 3.5, 2.6; 4th 3.2, 2.5; 5th

2.8,2.1.

First dorsal fin: vertical height 1 1.6, 1 1.4; length of base 9.6, 9.2.

Second dorsal fin: vertical height 4.2, 3.9 ; length of base $-5^ 5-2.

Anal fin: vertical height 3.5, 3.4; length of base 7.6, 6.7.

Caudal fin: upper margin 26.7, 25.7; lower anterior margin lO.i, lO.i.

Pectoral fin: outer margin 13.8, 15.O; inner margin 5.8, 5.8; distal margin 11. 6,

12.7.

Figure 82. Sfhyrna tiburo. A Female, 367 mm. long, from Rio de Janeiro, Brazil (Harv. Mus. Comp.
Zool., No. go). B Head of same, from below. C Dermal denticles of same, about 18 x. Z) Lateral and apical
views of dermal denticle, about 36 x. £ Left-hand upper and lower teeth of male, 812 mm. long, from
Tortugas, Florida (Harv. Mus. Comp. Zool., No. 848), about 2 x. F Fourth upper tooth. G Tenth upper
tooth. H Second lower tooth. / Seventh lower tooth. J Tenth lower tooth. F-J, about 4 x.

422 Memoir Sears Foundation for Marine Research

Distance from snout to: 1st dorsal 28.9, 29.1 ; 2nd dorsal 59.4,60.7; upper caudal

73-3>74-3j pectoral 21.6, 20.7; pelvics 46.7, 42.7; anal 58.1, 59.2.

Intersface between: ist and 2nd dorsals 21.6, 22.9; 2nd dorsal and caudal 9.2,

9.O; anal and caudal 6.9, 7.8.

Distance from origin to origin of: pectoral and pelvics 24.O, 23.7; pelvlcs and anal

12.2, 16.5.
Trunk at origin of ist dorsal between y^ and ^ as high as its length to origin of
caudal, moderately compressed. No mid-dorsal ridge. Caudal peduncle about % as wide
as deep, the upper precaudal pit strongly developed with its anterior outline oval or sub-
angular, the lower pit only about V2 as wide, but of similar form. Dermal denticles
relatively somewhat larger than in zygaena, varying from closely overlapping to loosely
spaced, the blades mostly rather steeply raised, usually with 5 strong ridges and as many
sharp-pointed marginal teeth (longer than in other local species), the axial slightly the
largest; short, slender pedicels.

Head a little less than Vs (average about 28 to 29%) of trunk to origin of caudal,
convex dorsally to opposite the angles of mouth but concave thence forward, very thin an-
terior to the eyes, its anterior part shovel- and not hammer-shaped, the outer posterior
margins of the shovel sloping a little rearward with broadly rounded outer corners;
the breadth of shovel at eyes about 2.5 times as great as its length in front of mouth
in small specimens but nearly 3 times that great in large, 1.3 to 1.6 times as great as
its length at outer ends of nostrils" and a little shorter than head to origin of pectorals;
anterior contour of head an uninterrupted curve from eye to eye, without definite depres-
sions opposite nostrils and merely a slight concavity or sinuosity median to the latter in
some cases only, the midsector a little more ovate in adult males where length in front of
mouth is almost % as great as distance between nostrils, while in adult females or in young
of both sexes it averages less than % (almost 57%) ; a faintly marked groove runs for-
ward from nostril along anterior margin of head in some small specimens for a distance
about as great as distance from nostril to eye, but this is hardly visible or wholly obsolete
in larger specimens; distance from outer end of nostril to anterior edge of eye a little
greater than diameter of latter; a line connecting inner ends of nostrils passes anterior to
mouth by a distance about 1V2 times the diameter of eye, one connecting the centers
of eyes passes about through front of mouth, while one connecting the outer posterior
corners of shovel passes posterior to corners of mouth by a distance about equal to diameter
of eye. Rostral cartilage without median hole, but preorbital process with a strongly marked
inwardly directed spur on anterior margin. Large and small mucous pores on lower surface
of anterior part of head distributed in diffuse pattern. Diameter of eye ^/i to Vs as long as
length of head in front of mouth. Gill openings extending ventrally and almost evenly
spaced, the ist almost 1.5 times as long as diameter of eye or a little longer, the 5th almost

16. The shovel of tiburo may average a little narrower relative to its length in northern specimens than in those
of the southern Atlantic. But the probability of local or racial variation is so great that recognition of this
difference does not seem essential in nomenclature.

Fishes of the Western North Atlantic 423

as long as ist, the 3rd only very little longer, the 5th over origin of pectoral. Nostril ap-
proximately transverse, a little shorter than diameter of eye. Mouth strongly arched,
about ^/-o as high as broad. A weakly marked labial furrow around corner of mouth extend-
ing a very short distance along lower jaw (concealed when mouth is closed), but none on
upper jaw.

Teeth '^ '° '''^Y— 7^'^ *° '^ > smooth-edged, except somewhat wavy on outer basej ist
upper tooth erect and symmetrical on expanded base, but subsequent upper teeth strongly
and increasingly oblique, their inner margins straight or slightly curved, their outer mar-
gins deeply notched in angular outline on 2nd to nth or 12th, the outermost 2 or 3 very
low without definite cusps, the 3rd to loth or i ith considerably the largest; lower teeth
shorter than uppers, with narrower cusps and relatively broader bases, the ist to 3rd erect,
symmetrical, the 4th to 7th or 8th slightly oblique with increasingly incised outer margin,
relatively broader bases and shorter cusps, the 8th and subsequent teeth without cusps,
low, oval, evenly rounded and without definite cutting edge; 3 series functional in front
of upper jaw and i to 2 along sides; 4 to 6 series functional in front of lower jaw, usually
2 along sides and 3 toward corners of mouth; i small symmetrical tooth at symphysis
on upper jaw, or none, and i on lower.

First dorsal moderately sloping with a perpendicular from its apex falling almost
at its rear tip, its origin a little anterior to inner corner of pectoral (or over it), its vertical
height a little less than % (about 6o%) and base a little less than V2 (about 44%) of the
length of head to origin of pectoral, its apex narrowly rounded, free rear corner about V2
as long as base, the midpoint of base approximately midway between axil of pectoral and
origin of pelvics, its rear tip anterior to latter by a distance about as great as diameter of
eye. Second dorsal a little less than V2 as long at base as ist in large specimens (slightly
longer in small) and almost ^3 as high, its anterior margin moderately convex, apex sub-
angular, posterior margin deeply concave, its free rear corner very slender, about 1.2 times
as long as the base or about 1.3 as long as the vertical height, its origin a little anterior to
midpoint of base of anal. Caudal between V4 and Vs (26 to 29%) of total length, the
upper margin nearly straight, its terminal sector about % of the fin, with narrowly
rounded apex and concave lower posterior margin; the lower lobe about Vs (average 37
to 38%) as long as upper, with weakly convex anterior margin, nearly straight posterior
margin, and narrowly rounded or subacute tip; the re-entrant contour (included by the 2
lobes) approximately a right angle with abruptly rounded corner. Distance from origin
of caudal to tip of anal about Yz as long as base of latter. Anal about 1.2 to 1.5 times as long
at base as 2nd dorsal but a little lower vertically, its anterior margin weakly convex, apex
subangular, posterior margin weakly concave toward apex, but nearly straight toward rear
tip, its free rear corner moderately slender, a little more than V2 as long as base in large
specimens or a little less in small. Distance from origin of anal to tips of pelvics about %
as long as base of anal. Pelvics about as long at base as anal, with weakly convex anterior
margins, weakly concave posterior margins, and narrowly rounded corners, their origins

424 Memoir Sears Foundation for Marine Research

about as near to inner corners of pectorals as to origin of anal. Pectoral a little more than
% (average about 68%) as long as head, or a little longer than vertical height of ist
dorsal and about % as broad as long, its outer margin moderately convex, distal margin
nearly straight or very slightly concave, apex and inner corner very narrowly rounded.

Color. Gray or grayish-brown above and a paler shade of the same hue below, some
specimens with a few small round dark spots on the sides j no conspicuous fin markings.

Size. A female of 1,076 mm. in the Harvard Museum of Comparative Zoology
contains eight young of about 300 mm. nearly ready for birth. Sexual maturity is attained
at a length of 3 V2 to 4 feet j it is said to reach 6 feet, but few grow longer than 5 feet.

Developmental Stages. Development is viviparous. The egg, with its developing
embryo, is enclosed at first in a tough but elastic shell, iridescent in appearance, the ends
of which are curiously plaited and folded, allowing for the growth of the embryo by their
expansion. After the embryo is set free from the eggshell and the yolk has been absorbed,
the empty yolk sac becomes attached to the uterine walls of the mother, forming a so-called
yolk-sac placenta richly supplied with blood vessels. The embryos are described as lying
with their heads toward the anterior end of the uterus. The umbilical cord may be about
as long as the embryo and is closely set along its whole length with large villi, some of
which are simple but others branched. The number of embryos (6 to 9) is much smaller
than in -zygaena, corresponding to their type of development, with males and females in
about equal numbers,"

Habits. This species occurs chiefly in shallow water, close inshore, often in bays and
estuaries, sometimes coming right up to wharves. It is said to be more sluggish than other
Hammerheads. It feeds largely on whatever crabs may be available locally and on other
Crustacea such as mantis shrimps (Squilla), shrimps, isopods and even barnacles. But its
recorded diet also includes bivalve mollusks, cephalapods (Octopus), small fish, and
even seaweed, the latter no doubt taken incidentally with crabs, etc. And it has been de-
scribed as burrowing under coral masses in search of small fish and Crustacea in southern
Florida waters. It takes a hook readily on almost any kind of bait and is often said to follow
fishing boats to pick up any fish or other scraps that may be discarded.

Relation to Man. It is of no commercial value except for a few that may be sold in the
fish markets. It is entirely harmless.

Range. Tropical to warm-temperate belt of the Atlantic from southern Brazil north-
ward regularly to the southern part of North Carolina and as a stray to southern New
England and Massachusetts Bay in the west} it apparently occurs also in tropical West

17. For an excellent illustration of the embryo with yolk stalk and large yolk sac before the disappearance of the
external gills, see Lcuckart (Unters. Auss. Kiemen Rochen Hayen, Stuttgart, 1836: 22, pi. 3) ; for further ac-
counts of the early development of tiburo, see Gudger (Science, N. S. 55, 1912: 466; Proc. biol. Soc. Wash., 25,
1912: 143), Radcliffe (Bull. U.S. Bur. Fish., 40, 1916: 266) and Longley and Hildebrand (Pap. Tortugas
Lab., 31, 1941: 3).

Fishes of the Western North Atlantic 4.25

Africa in the east (probably not the Mediterranean^") as well as on Pacific coasts of Amer-
ica from southern California to Ecuador."" Tiburo has also been reported by name' from
China"' and from the Philippines/' but without supporting evidence as to the identity of
the particular specimens in question; and since it is not included in any of the descriptive
surveys of the sharks of China, Australia, the Philippines or India that have appeared so
far, although it is not likely to be overlooked if at all common, we think its presence un-
likely in eastern Pacific or Indian waters.

Occurrence in the Western Atlantic. This shark is known from so many localities from
southern Brazil to Florida, including the South American coastline, the West Indian-
Caribbean region as a whole, and the Gulf of Mexico," and so often reported as common
as to prove it not only generally distributed throughout this entire area but in fact one of
the more plentiful of littoral sharks there. No doubt it is also a year-round resident
throughout this belt, as it certainly is around southern Florida, except perhaps for mid-
summer. To the north of Florida, however, it occurs on the Atlantic coast only as a summer
visitor, common in the warm months between June and October on the coast of South
Carolina, where it has been characterized as one of the more abundant of summer sharks,
and as far as Beaufort, North Carolina. But few pass Cape Hatteras; it is taken only occa-
sionally in the pound nets in Chesapeake Bay, and it occurs only as a stray farther north,
being recorded once or twice in New Jersey waters, once at Newport, Rhode Island, and
once in Massachusetts Bay. We have also received a reliable report that about six speci-
mens, about four feet long, were taken in Nantucket Sound among the other species caught
in the shark fishery that was carried on there during the summer of 191 8."'

Synonyms and References:

Squalus tiburo Linnaeus, Syst. Nat., I, 1 758: 234 (descr., refs., Amer.) ; Syst. Nat., i, 1766: 399 (same
as foregoing); Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1 788: 9 (descr., S. Amer.); Gmelin, in
Linnaeus, Syst. Nat., / (3), 1 779: 1495 (same as Linnaeus, 1758, Amer. seas); Walbaum, P. Artedi
Genera Pise. Emend. Ichthyol., 3, 1792: 516 (diagn., refs.); Shaw and Nodder, Naturalist Misc., 7,
1795: pi. 229 (good ill., S. Amer. seas) ; Bloch and Schneider, Syst. Ichthyol., 1 801: 131 ; Bosc, Nouv.
Diet. Hist. Nat., 21, 1803: 189 (diagn.); Latreille, Nouv. Diet. Hist. Nat., 24, 1 804: 72 (in table of

19. The brief description by Rochebrune (Act. Soc. linn. Bordeaux, [4] 6, 1882: 44) of the head of the Sene^am-
bian shark, reported by him as Zygaena leeuivenii, suggests that he was actually dealing with tiburo. But we think
it improbable that it occurs in the Mediterranean, for while it is credited tentatively to the local fish fauna by
Doderlein (Man. Ittiol. Medit., 2, 18S1: 50), the few reports of it there or for southern Spain lack positive
evidence as to the specific identity of the specimens concerned.

20. The eastern Pacific form has recently been made a separate species, vesfertina Springer (Stanford Ichthyol. Bull.,
' [sli 1940: 161) on the basis of a supposedly wider shovel than in tiburo relative to its length, longer rela-
tive distance between nostrils, higher first dorsal, shorter pectoral and longer caudal. But comparative measure-
ments show that there is no discontinuity in any of these respects between specimens from San Diego, California
or Panama and the extensive Atlantic series with which we have compared them.

21. Gunther, Cat. Fish. Brit. Mus., 8, 1870: 3S2; Elera, Cat. Fauna Filipp., i, 1895: 615.

22. Localities of definite record from south to north are: Santos, Rio de Janeiro, Bahia, Pernambuco, Rio Parahyba
and Natal in Brazil; French Guiana; Trinidad; Colon; Progreso, Yucatan; Belize, British Honduras; Turks
Island; Cuba; the Bahamas; many localities along both coasts of Florida; several along the coasts of Louisiana
and Texas.

13. Personal communication by R. H. Bodman, who operated the fishery.

4-26 Memoir Sears Foundation for Marine Research

contents) ; Cuvier, Regne Anim., 2, 1817: 127, footnote (diagn.) ; Nardo, Prod. Ittio!. Adriat., 1827: 9
(name only, Medit.) ; Gray, Cat. Fish. Coll. Descr. by L. T. Gronow in Brit. Mus., 1854: 6 (diagn.,
Amer. seas).

Le Pantouflier, Broussonet, Mem. Math. Phys. Acad. Sci. Paris, 1780: 662 (not Squale Pantouflier, Lacepede,
1798, in Sonnini, 1802, which is tudes; see p. 434).

Round Headed Zygacna, Shaw and Nodder, Naturalist Misc., 7, 1795: pi. 229 (good ill., S. Amer. seas).

Squale carolinien, Bosc, Nouv. Diet. Hist. Nat., 27, 1803: 190 (diagn., Carolina).

Heart Headed Shark, Shaw, Gen. Zool., 5, 1804: 355, pi. 154, upper fig. (good ill., S. Amer. seas).

Sfhyrna tiburo Rafinesque, Indice Ittiol. Sicil., 1810: 47 (name, for Squalus tiburo Linnaeus, 1758);''*
Muller and Henle, Plagiost., 1841: 53 (descr., Brazil); Jordan, Proc. U.S. nat. Mus., 5, 1882: 345
(Pensacola, Florida, abundant); Jordan and Gilbert, Proc. U.S. nat. Mus., 5, 1882: 581 (commonest
summer shark, Charleston, S. Carolina); Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 796 (name,
S. Atlant. and Gulf States of U.S.) ; Nelson, Rep. St. Geol. N. J., 2 (2), 1890: 661 (N. Jersey, size) ;
Carus, Prod. Fauna Medit., 2, 1 889-1 893: 514 (occur, in Medit. questioned) ; Henshall, Bull. U.S. Fish
Comm., 9, 1891: 384 (Florida) ; Lonnberg, Overs. Vet. Akad. Forhandl., $1, 1894: ill (S. Florida);
Henshall, Bull. U.S. Bur. Fish., 14, 1895: 210 (size, abund., Florida Keys and west coast); Evermann
and Bean, Rep. U.S. Comm. Fish. (1896), 1898: 239 (reported Indian R., Florida); Jordan and
Evermann, Bull. U.S. nat. Mus., 47 (3), 1898: 2748 (Mazatlan, Mexico); Evermann and Kendall,
Rep. U.S. Comm. Fish. (1899), 1900; 44 (W. coast Florida); Jordan and Evermann, Bull. U.S. nat.
Mus., 47 (4), 1900: pi. 5, fig. 19 (ill-); Wilson, Amer. Nat., ^4, 1900: 355 (common, N. Carolina);
Bean, T. H., Rep. For. Comm. N. Y., 1901: 378 (occasional, N. York); Gregg, Where to Catch Fish,
1902: 16 (ill., descr., size, E. Florida) ; Bean, T. H., Bull. N. Y. St. Mus., do, Zool. 9, 1903: 30 (descr.,
N. Jersey) ; Gilbert and Starks, Mem. Calif. Acad. Sci., 4, 1904: 13 (comp. with other species, Mazatlan,
Mexico) ; Bean, B. A., in Shattuck, Bahama Islands, 1 905 : 296 (off Abaco, Bahamas) ; Jordan and Thomp-
son, Bull. U.S. Bur. Fish., 24, 1905 : 232 (habits, food, Tortugas, Florida) ; Linton, Bull. U.S. Bur. Fish.,

24, 1905: 344 (abund., embryos, food, parasites, N. Carolina) ; Cole and Barbour, Bull. Mus. comp. Zool.
Harv., 50, 1906: 155 (Yucatan) ; Schreiner and Ribeiro, Arch. Mus. nac. Rio de J., 12, 1906: 79 (Bahia,
Rio de J.) ; Starks, Proc. U.S. nat. Mus., 50, 1906: 763 (Guayaquil, Ecuador) ; Gudger, Science, N. S.

25, 1907: 1006 (food) ; Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907: 156, 201 (descr., Brazil, refs.) ;
Smith, Bull. N. C. geol. econ. Surv., 2, 1907: 35 (abund., embryos, food, N. Carolina); Wilson, Proc.
U.S. nat. Mus., 55, 1907: 431 (parasites, N. Carolina) ; Fowler, Proc. Acad. nat. Sci. Philad., 60, 1908:
66 (Newport, Rhode Island, Florida specs.); Wilson, Proc. U.S. nat. Mus., 55, 1908: 626 (parasites,
N. Carolina); Rosen, Lund Univ. Arsberiitt., N. S. (2) 7 (5), 191 1: 47; Kongl. Fysiogr. Sallsk. Hand.,
N. S. 22 (5), 191 1 : 47 (Bahamas) ; Wilson, Proc. U.S. nat. Mus., 59, 191 1 : 390 (parasites, N. Carolina) ;
Gudger, Science, N. S. 55, 191 2: 466 (young embryo, yolk stalk, N. Carolina) ; Proc. biol. Soc. Wash.,
25, 1 91 2: 143 (size, embryos, N. Carolina); Starks, Stanford Univ. Publ. Biol., 1913: 5 (Natal, Brazil);
Coles, Proc. biol. Soc. Wash., 21?, 1 91 5: 90 (abund., season, N. Carolina); Fowler, Proc. Acad. nat. Sci.
Philad., 6y, 1916: 521 (Trinidad); Copeia, 30, 1916: 36 (listed, mid. Atlant. U.S.); Nichols and
Murphy, Brooklyn Mus. Sci. Bull., 5 (i), 1916: 18 (diagn., reported near N. York, but no definite
Long Island record); Smith, J. Amer. Mus. nat. Hist., 16, 1916: 347 (Atlant. Coast, U.S.); Wilson,
Ann. Carneg. Mus., jo, 1916: 58 (Guayaquil, Ecuador) ; Fowler, Proc. Boston Soc. nat. Hist., 55, 191 7:
1 10 (Newport, Rhode Island); Proc. Acad. nat. Sci. Philad., dp, 1917: 127 (Colon); Ribeiro, Rev.
Mus. paul., 10, 1918: 707 (Santos, Brazil); Fowler, Proc. biol. Soc. Wash., jj, 1920: 144 (N.
Jersey, size); Nichols, BuU. Amer. Mus. nat. Hist., 44, 1921: 22 (Turks Island); Fowler, Proc. biol.
Soc. Wash., 5<5, 1923: 28 (Florida S. Coast); Ribeiro, Fauna brasil. Peixes, 2 (i) Fasc. I, 1923: 13
(same as Ribeiro, 1 907) ; Borodin, Bull. Vanderbilt Oceanogr. (Mar.) Mus., \ (l), 1928: 5 (no loc.) ;
Hildebrand and Schroeder, Bull. U.S. Bur. Fish., 4^, 1928: 51 (descr., Chesapeake Bay); Rey, Fauna
Iberica Feces, i, 1928: 370 (descr., occur, in Medit. questioned) ; Breder, Field Bk. Mar. Fish. Atlant.
Coast, 1929: 19 (general); Fowler, Copeia, 1931: 46 (Corpus Christi, Texas); von Bonde, J. Comp.
Neurol., 5,?, 1933: 381 (ill. head), pi. 2, fig. 4 (ill. brain, no loc); Borri, Atti Soc. tosc. Sci. nat.,
44, 1934: 91 (Bahia, Brazil); Burton, Sci. Mon. N. Y., 40, 1935: 283 (abund., season, S. Carolina);

24. Spelled "liburo," an evident misprint.

Fishes of the Western North Atlantic 427

Brooks, Parasitology, 26, 1936: 266 (parasites, Tortugas, Florida); Norman and Fraser, Giant Fishes,
1937: 47 (mentioned); Tortonese, Atti Soc. ital. Sci. nat., yj, 1938: 303 (Rio de J.); Springer, Proc.
Fla. Acad. Sci., ^, 1939: 30 (descr., ill. head, teeth, size, food, abund., Florida W. coast); Stan-
ford Ichthyol. Bull., / (5), 1940: 162, 164-166 (comp. with vesfertina, key, meas., ill. of head);
Fowler, Bull. U.S. nat. Mus., 100 {13), 1941: 212 (ref. for Atlant., Rhode Island record, but probably not
the refs. for China, Philippines); Longley and Hildebrand, Pap. Tortugas Lab., 34, 1941: 3 (embryos,
descr. yolk stalk, Tortugas, Florida); Springer, Proc. Fla. Acad. Sci., 5, 1941: 5 (comp. with other
species, ill. of skull); Norris, Plagiost. Hypophysis, 1941: pi. n, fig. 40 (brain); Fowler, Arqu. Zool.
Estado Sao Paulo, 3, 1942: 129 (Brazil) ; Fish Culturist, 21 (g), 1942: 66 (listed, Cuba) ; Lunz, Bull.

5. C. St. Planning Bd., 14, 1944: 27 (S. Carolina, Florida) ; Bigelow and Schroeder, Guide Comm. Shark
Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 132, fig. 48 (descr., range, ill.); Fowler, Monogr.
Acad. nat. Sci. Philad., 7, 1945: 96 (N. Carolina), 160 (S. Carolina); Gunter, Publ. Inst. Mar. Sci.
Univ. Texas, x, 1945: 20 (temp., breeding, Texas).

Cestrorhinus tiburo Blainville, Bull. Soc. philom. Paris, 1816: 121 (name only).

Zygaeim tiburo Valenciennes, Mem. Mus. Hist. nat. Paris, p, 1822: 226, pi. 12, fig. 2*, 2" (ill. of head,
Brazil); Bory de St. Vincent, Diet. Class. Hist. Nat., 15, 1829: 597 (general); Cuvier, Regne Anim.,
2, 1829: 394, footnote (diagn., refs.); Cloquet, Diet. Sci. Nat., 60, 1830: 623 (general); Voigt, in
Cuvier, Tierreich, 2, 1832: 514 (diagn., Brazil); Yarrell, Brit. Fish., Suppl., 1839: 64, 65 (synonyms,
ill. of head, Brazil); Hist. Brit. Fish., 2, 1 841: 507, 508 (same as Yarrell, 1 839); Cuvier, Regne Anim.,
ill. ed., 1843: 309, footnote; Baird, Rep. Smithson. Instn. (1854), 1855: 337 (N. Jersey record);
Castelnau, Anim. Nouv. RaresAmer. Sud., 2, 1855: 99 (Bahia, Brazil) ; Gunther, Trans, zool. Soc. Lond.,

6, 1869: 396 (Atlant. coast. Cent. Amer.); Cat. Fish. Brit. Mus., 8, 1870: 382 (descr., refs., Bahia,
Belize, but spec, from China probably not this species); Doderlein, Man. Ittiol. Medit., 2, 1 881: 50
(occur, in Medit. questioned); Garman, Bull. Mus. comp. Zool. Harv., 8, 1881: 232 (Florida Keys);
Werner, Zool. Jb., Syst. Abt., 21, 1904: 284 (Atlant. Oc.) ; Huber, Z. wiss. Zool., yo, 1901 : 626 (anat.
of claspers).

Zygaena (no specific name) Agassiz, L., Poiss. Foss., 3, 1835: pi. E, fig. 9 (teeth).

Zy gaena (Sgua/us) tiburo Leuckart, Unters. Auss. Kiemen Rochen Hayen, Stuttgart, 1836: 22 (ill., early
embryo, yolk stalk, yolk).

Platysqualus tiburo Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1839: 318 (descr.).

Sfhyrnias tiburo Gray, List. Fish. Brit. Mus., /, 1 851: 50.

Cestracion-^ tiburo Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861: 59 (in N. Amer. cat.), and later eds.;
Dumeril, Hist. Nat. Poiss., /, 1865: 385 (descr., east coast, N. and S. Amer.); Abbott, Geol. Surv.
N. J., Geol. N. J., Append. E, 1868: 829 (N. Jersey) ; Fowler, Rep. N. J. Mus. (1905), 1906: 65 (N.
Jersey); Proc. Acad. nat. Sci. Philad., 5,?, 1906: 80 (Florida Keys, abund.); Rep. N. J. Mus. (1906),
1907: 257 (N. Jersey record); Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 160, pi. I, fig. 4, 5
(descr., ill., Massachusetts Bay spec.) ; Raddiffe, Bull. U.S. Bur. Fish., 34, 1 91 6: 265, pi. 44, fig. I, 2,
but not fig. 3, which appears to be either diplana or zygaena (denticles, teeth, food, N. Carolina) ; Meek
and Hildebrand, Field Mus. Publ. Zool., 75 (l), 1923: 60 (descr.. Colon, Panama) ; Breder, Bull. Bing-
ham oceanogr. Coll., 1 (l), 1 92 7: 5 (Florida) ; Bigelow and Schroeder, Canad. Atlant. Fauna, biol. Bd.
Canad., I 2^ 1934: 10 (descr., distrib.) ; Bull. U.S. Bur. Fish., 48, 1936: 322 (ref. to Massachusetts Bay
spec, descr. by Garman, 1 91 3).

Renicefs tiburo Gill, Ann. N. Y. Lye, 7, 1862: 412 (name); Rep. U.S. Comm. Fish. (1871-72), 1873:
813 (Cape Cod to Florida); Poey, An. Soc. esp. Hist. Nat., 5, 1876: 384; Enumerat. Pise. Cubens.,
1876: 188 (listed, Cuba); Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 25, 874 (descr., size,
Atlant. Oc, but rep. for China probably not this species) ; Bean, T. H., Bull. U.S. Fish Comm., 7, 1888:
152 (old N. Jersey record); Wilson, Proc. U.S. nat. Mus., 33, 1907: 360 (parasites, St. Mary's R.,
Florida); Jordan, Manual Vert. Anim. NE. U.S., 1929: 11 (general); Jordan, Evermann and Clark,
Rep. U.S. Coram. Fish. (1928), 2, 1930: 17 (Atlant. Oc, Mazatlan, Panama, but rep. for China prob-

25. This name was first used by Klein (Neuer Schauplatz, 3, 1776: 526), but must date from Gill, 1861, because
Klein's names are not applicable by Opinion 89 of the International Commission on Zoological Nomenclature;
see footnote 2, p. 408.

4-2 8 Memoir Sears Foundation for Marine Research

ably not this species) ; Gowanloch, Bull. La. Conserv. Dep., 23, 1933: 219, 220 (abund. Gulf coast U.S.) ;

Bere, Amer. Midi. Nat., 17 (3), 1936: 583, 605 (parasites, Florida west coast); Whitley, Aust. Zool.,

8, 1936: 190 {Squalus carolinianus Latreille, 1804, a synonym of tiburo).
Zygaena leeuwenn Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 44; Fauna Senegambie, Poiss., 1,

1883-1885: 21 (ident. probable because of descr. of head; abund. Senegambia; not Zygaena letuini

Griffith, in Cuvier, Anim. Kingd., ro, 1834: 640, pi. 50) ; Metzelaar, Trop. Atlant. Visschen, 1919: 189

(Senegambia by ref. to Rochebrune, 1882).
Renicefs (no spec, name) Adams and Kendall, Bull. U.S. Fish Comm., 9, 1 891 : 295 (Florida west coast).
Sfhyrna {Renicefs) tiburo Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 217 (Long Island

southward and Mazatlan, Mexico, but rep. for China probably not this species) ; Bull. U.S. nat. Mus.,

47 (l), 1896: 44 (descr., Long Island southward, and Mazatlan, Mexico, but rep. for China probably

not this species).
Cestracion zygaena Radcliffe, Bull. U.S. Bur. Fish., 34, 1916: pi. 43, fig. 2 (photo, of jaws, evidently mis-
labelled by accident).
Sfhyrna tiburo (perhaps in part) Puyo, Bull. Soc. Hist. nat. Toulouse, yo, 1930: 83 (ill. of head is difhna,

but discuss, perhaps covers tiburo also).
Sfhyrna vesfertina Springer, Stanford Ichthyol. Bull., i (5), 1940: 161, 164-166 (type descr., comp. with

other species, meas., ill. of head, Panama, Guayaquil, Ecuador) ; Beebe and Tee-Van, Zoologica, N. Y.,

2(5, 1941: 116 (diagn., Panama, Ecuador).
Not Zygaena tiburo Reguis, Ess. Hist. Nat. Provence, Fasc. I, 1 877: 59 (either diflana or tudes because head

descr. as emarginate in midline, Medit.).

Sfhyrna tudes (Valenciennes), 1822

Great Hammerhead

Figures 83, 84

Study Material. Male embryo, about 585 mm., a newborn male of 673 mm., another
small specimen, and the head of a specimen of about 1,660 mm. as calculated from breadth
of head, all from Englewood, Florida (Harv. Mus. Comp. Zool. and U.S. Nat. Mus.) j
jaws from specimens of 13V2 feet and 7 feet 10 inches from Englewood, Florida (U.S.
Nat. Mus., No. 1 10299, 110300)5 excellent cast of a male, a little more than lO feet
long, from Miami, Florida^* (Harv. Mus. Comp. Zool.). Measurements of six speci-
mens, male and female, 700 to 3,155 mm., from Englewood, Florida, as well as photo-
graphs of a large one on the beach.^^

Distinctive Characters. The hammer (not shovel) shape of its head places tudes with
zygaena, diflana and bigelowi; and it falls with the last two because the anterior margin
of its head is definitely indented in the midline. But it is easily distinguishable from all
three by the edges of both its lower and upper teeth, which are regularly serrate from tip
to base (smooth or weakly serrate in zygaena), and by its first dorsal fin which is less erect.
It is further marked off from zygaena and diflana by the much shorter free rear corners of
its second dorsal and anal fins and by its differently shaped hammer head (cf. Figs. 83, 84
with 81 G, 86 A). Also, its pelvics are much more strongly convex anteriorly and concave
posteriorly than those of any other Atlantic member of the genus.

26. Prepared by AI. Pflueger. 27. Contributed by Stewart Springer.

Fishes of the Western North Atlantic

429

Description. Proportional dimensions in per cent of total length. Male, 673 mm.,
from Englewood, Florida (U.S. Nat. Mus., No. 106543).

Trunk at origin of pectoral: breadth 10.4; height 10.4.
Snout length in front of: outer nostrils 3.1 5 mouth 7.0.
Eye: horizontal diameter 2.4.
Mouth: breadth 6.0; height 3.3.
Nostrils: distance between inner ends 16.9.

Gill opening lengths: ist 2.8; 2nd 2.9; 3rd 3.0; 4th 2.8; 5th 2.4.
First dorsal fin: vertical height 1 1-9; length of base lO.i.
Second dorsal fin: vertical height 4.4; length of base $•(>•
Anal fin: vertical height 4.0; length of base 6.7.
Caudal fin: upper margin 30.O; lower anterior margin 10.4.
Pectoral fin: onttr mzTgin 12.3; inner margin 4.4; distal margin 9.2.
Distance from snout to: ist dorsal 25.4; 2nd dorsal 57.3; upper caudal 70.O; pec-
toral 20.2; pel vies 43.7; anal 55.9.

Figure 83. Sfhyrna tudes, about l,66o mm. long, from Englewood, Florida (U. S. Nat. Mus., No. 108453).
A Anterior part of head from below. B Left-hand upper and lower teeth of same, about 1.3 x. C Fifth upper
tooth. D Twelfth upper tooth. E Fifth lower tooth. F Eleventh lower tooth. C-F, about 2 x.

43 o Memoir Sears Foundation for Marine Research

Interspace between: ist and 2nd dorsals 20.8 ; 2nd dorsal and caudal 7. i ; anal and

caudal 6.8.

Distance from origin to origin of: pectoral and pelvics 2 1 . i ; pelvics and anal 12.9.

Figure 84. Sfhyrna tudes, new-born male, 673 mm. long, from Englewood, Florida (U. S. Nat. Mus.,
No. 106543). A Anterior part of head from below, about 0.4 x natural size. B Dermal denticles, about 65 x.

Two females, 1,745 and 3,155 mm., from Englewood, Florida (calculated from
measurements by Stewart Springer).

Snout length in front of: mouth 5.4, 4.6.

Eye: horizontal diameter 1.4, 0.9.

Mouth: breadth S.6, 7.1; height 2.9, 4.3.

Nostrils: distance between inner ends 16.3, 19.0.

Gill opening lengths: ist 3.3, 4.0 j 5th 2.3, 2.8.

First dorsal fin: length of base lO.o, lO.i.

Second dorsal fin: length of base 4.9, 4.4.

Anal fin: length of base 6.3, 6.0.

Caudal fin: upper margin 31.0, 29.5; lower anterior margin 12.O, 11.7.

Pectoral fin: outer margin 14.1, 15.85 inner margin 3.9, 3.85 distal margin 11.4,

14.4.

Distance from snout to: ist dorsal 27.8, 28.55 upper caudal 69.0, 70.5; pectoral

20.6, 21.8.

Interspace between: ist and 2nd dorsals 2 1 .7, 1 9.8 j 2nd dorsal and caudal 8.3, 7.6.

Fishes of the Western North Atlantic 431

Trunk moderately compressed, as usual in this genus, but somewhat stouter when
adult than in the diflana-zy gaena group, its dorsal profile moderately arched. No trace
of mid-dorsal ridge. Caudal peduncle about % as thick as high, the upper precaudal pit
strongly developed as a broadly triangular depression but no definite lower pit. Dermal
denticles close-spaced and overlapping regularly (laterally as well as longitudinally) but
with skin visible here and there, the blades but little arched longitudinally and not much
raised, very thin, about as broad as long, smooth toward base, but with 3 to 5 low rounded
ridges toward free margin in small specimen (perhaps 5 to 6 in large specimens), the mar-
ginal teeth rather short and broad, the median very little the longest; pedicels short and
thick; basal plates with 4 rather long rays.

Head to origin of pectoral about Vs of length of trunk to origin of caudal or a little
less, its dorsal profile evenly convex posterior to eyes but concave forward to a rather thin
tip, hammer-shaped, the outer posterior margins slightly concave and nearly transverse
(sloping more strongly rearward in diplana and zygaena) ; width of head at eyes about
1.2 times its length to origin of pectoral in small specimens, the width of hammer rela-
tive to its length at oculo-narial prominences increasing from about 3 : i in very small speci-
mens to about 4:1 in medium-sized and large; anterior margin of head scalloped with a
depression opposite each nostril (not as deep as in the diflana-zy gaena group), a shallow
but unmistakable indentation in the midline, and a shallow sinuosity between these two;
the groove running from the nostril inward along anterior margin of head only very
faintly marked, but visible upon careful examination nearly halfway to the midline. Dis-
tance from anterior margin of eye to anterior corner of oculo-narial prominence about as
long as diameter of eye; a line connecting inner ends of nostrils passes anterior to front of
mouth by a distance nearly or quite twice as long as diameter of eye (not longer than
diameter of eye in the diplana-zy gaena group), one connecting centers of eyes passes
in front of mouth by a distance only about ^2 as long as diameter of eye in newborn speci-
mens but about I V2 times that long in larger (it passes close in front of mouth in diflana,
and a little posterior to it in zygaena) and one connecting outer posterior corners of
hammer passes a little posterior to corners of mouth in newborn specimens but anterior
to it in large by a distance at least V2 as long as diameter of eye (in adults of the
diplana-zy gaena group this line passes posterior to corners of mouth). Head in front of
mouth about Ys as long as to pectorals in small specimens but decreasing in relative length
with growth to only about Vs that long in large adults. Rostral cartilage with a median
oval or triangular hole, but wings of preorbital processes without inwardly directed points
on anterior margin (intermediate in this respect between diplana and zygaena). Mucous
pores in median belt on oral side of front of head covering a subtriangular area, much as in
zygaena. Eye a little broader than high, its horizontal diameter about Vs as long as head
in front of mouth in young but increasing so little in size with later growth as to be only
about Vs as long as head in front of mouth in large specimens (much smaller than in the
diplana-zy gaena group ).^* Gill openings extending ventrally, the ist to 4th nearly straight

28. Diameter of eye i6 to 18 mm. in specimens of 700 to 72J mm. but only 25 mm. at a length of 1,735 to 1,745
mm., 26 mm. in a specimen of 2,370 mm., and 29 mm. in one of 3,155 mm.

4-32 Memoir Sears Foundation for Marine Research

but the 5th slightly concave anteriorly, their relative lengths increasing with growth, the
I St being only a little more than Mi as long as head in front of mouth or about 1.3 times
as long as diameter of eye in young, nearly % as long as head in front of mouth or 2.2 to
3.3 times as long as diameter of eye in medium-sized specimens, and nearly 90% as long
as head in front of mouth or 4.3 times as long as diameter of eye in largest measured (about
10 ft. 4 in. long); the 3rd gill opening very little longer than ist, the 5th only a little
shorter (about 84%) than ist, the two spaces between ist and 3rd about equally broad,
but those between 3rd and 5th much narrower, the 4th gill opening about over origin of
pectoral. Nostril approximately transverse. Mouth about ^o as long as broad. No trace of
lower or upper labial furrows, although there may be wrinkles at corners of mouth when
the latter is closed.

Xeeth ■ 17— 2 or 3— 17 — (.j^gjj. edges reg;ularly and moderately coarsely serrate from

16 or 17 — 1 to 3 — 16 or 17' o o ^ ^J J

tip to base, except for the outermost i or 2, which are nearly or quite smooth ; upper teeth
triangular, on expanded bases, the ist tooth erect, symmetrical, but the subsequent upper
teeth increasingly oblique toward corners of mouth, their outer margins increasingly
notched, inner margins concave toward base except along outer Vs of jaw where they are
evenly and rather strongly convex; 2nd to lOth or nth upper teeth considerably the larg-
est and the outermost 2 to 3 very short; lower teeth similar to uppers, except perhaps a
little narrower and with inner edges along outer Ms of jaw nearly straight instead of con-
vex; 2 or 3 minute teeth at symphysis in upper jaw, and i to 3 in lower.

First dorsal narrower toward apex and more sloping than in diplana or zygaena, a
perpendicular from its apex falling posterior to its rear tip in young specimens but a little
anterior to its tip in large, its origin about over midpoint of inner margin of pectoral, its
vertical height about V2 as great as length of head, the anterior margin moderately convex
toward apex but perhaps less so in adult than in young, the posterior margin deeply con-
cave in young but less so in adult, its free rear corner comparatively broad and only about
1/4 as long as base both in young and old (more than Y^ as long as base in diplana) ; the mid-
point of base about 1.8 times as far from origin of pelvics as from axil of pectoral in young.
Second dorsal at base averaging a little less than V2 as long as ist and about Y^ as high, its
origin a little anterior to midpoint of base of anal, its anterior margin moderately convex,
posterior margin very deeply concave in subangular outline (much more deeply so than
in either diplana or zygaena), apex rounded, its free rear corner about as long as the base
(much longer, relatively, in the diplana-zy gaena group). Caudal a little less than Yz (29
to 31%) of total length, its upper margin weakly convex in young and even less so in
adult, the terminal sector about % of the fin, with narrow tip and rather deeply concave
lower posterior margin, the lower lobe a little more than Vs (35 to 42%) as long as upper,
slender, with very narrowly rounded apex, its lower anterior margin rather strongly con-
vex in young but weakly so in adult, its posterior margin nearly straight; the re-entrant
contour (included between the 2 lobes) with rounded corner, a little less than a right angle.
Distance from origin of caudal to tip of anal a little more than Yi as long as base of latter.

Fishes of the Western North Atlantic 433

Anal about i.i to 1.3 times as long at base as 2nd dorsal, generally similar in shape, deeply
incised near the apex much as in diplana and in zygaena. Distance from origin of anal to tips
of pelvics between % and % as long as base of anal. Pelvics only about % as long at base
as anal, their anterior margins more strongly convex and the posterior margins more
deeply concave than in any other Atlantic member of the genus. Pectoral noticeably small,
its length a little more than Y2 as great as that of head in young but nearly % (72 to 73%)
that length in adults or about as long as vertical height of ist dorsal, a little more than Y>
as broad as long, its outer margin strongly convex toward apex in young but less so in
adult, its distal margin only moderately concave, apex subacute and inner corner narrowly
rounded.

Color. Small specimens are brownish gray above and a paler shade of the same tint
below; the dorsals, lower and upper caudal lobes, upper surfaces of the pectorals and
lower edge of the caudal are dusky toward the tips. A cast of a large one, colored from the
fresh-caught specimen, is dark olive above and pale olive below, without any conspicuous
fin markings.

Size. Available information suggests a length slightly less than 700 mm. at birth
which corresponds to the large number in a litter. This is the largest of Atlantic Hammer-
heads, apparently not maturing at a length less than about 10 feet and commonly growing
to 13 to 14 feet, with individuals of 1 5 feet reliably reported.

Developmental Stages. It is not known whether tudes is ovoviviparous or viviparous;
30 to 38 embryos have been found in females off southern Florida. Embryos ready for
birth differ from adults in the relatively greater length of the hammer relative to its
breadth, much larger eyes, longer head in front of mouth, shorter gill openings, more
rounded pectoral and caudal, and more oblique and more rounded first dorsal.

Habits. Nothing is known of the habits or diet of tudes to set it apart from the
diflana-zy gaena group. Around southern Florida females have been taken in June.

Relation to Man. The Great Hammerheads that are taken in the shark fisheries of
Florida and the West Indies are utilized with others for leather, etc. Now that shark liver
oil is in demand for its vitamin content, large specimens of tudes may prove very valuable,
for some (but not all) are extremely rich in Vitamin A. As an example we may cite a 13-
foot 1 0-inch Florida specimen which was recently caught and whose liver oil had a
potency of 357,000 units of Vitamin A per gram with a sale value of about $500 at
current prices. But this was exceptional. Another of about the same size, caught at the same
locality a few days later, had a Vitamin A potency of only about 55,000 units per gram,
although it yielded about three times as much oil, its market value thus being only about
$150."

Range. Tropical and subtropical Atlantic. Hammerheads have also been reported as
tudes from the west coast of Central America, the Hawaiian Islands, Australia, Indo-
China, the East Indies, the Philippines, India and the Gulf of Arabia.^" But whether or

29. Personal communication from Stewart Springer.

30. See Fowler (Bull. U.S. nat. Mus., loo ['j], 1941 : 213) for a list of Australian and East Indian citations.

434 Memoir Sears Foundation for Marine Research

not the Indo-Pacific form is actually identical with that of the Atlantic remains uncertain,
awaiting critical comparison of specimens from the respective seas."

Occurrence in the Atlantic. It is not yet possible to describe the distribution of tudes
in any detail for either side of the Atlantic, because in very few instances have reports of
its presence been accompanied by evidence sufficient to establish the actual identity of the
particular specimens on which they were based. On the eastern side of the Atlantic its
presence is so far positively established only for the Mediterranean, where it has been
taken widely, although never in any numbers. It is also reported off the Atlantic coast of
Spain, from the Cape Verde and Canary Islands, and from tropical West Africa (Morocco,
Senegambia, Gambia). Positive locality records for the western North Atlantic are: French
and Dutch Guiana (Cayenne, Surinam); Cuba; western and southeastern Florida; and
North Carolina (Cape Lookout and Beaufort, one specimen each). We have no reason to
suppose that it accompanies the other Hammerheads in their yearly migration farther
north offshore in the Gulf Stream (p. 442). In the opposite direction there is report of it
from Brazil (Santos, Pernambuco), from Uruguay, and from northern Argentina. But
the only Brazilian record of tudes that is accompanied by an illustration" appears actually
to have been based on a specimen of S. diflana (p. 420), while reports of its presence
farther south lack any supporting evidence.

The foregoing, together with the fact that tudes is taken in some numbers off both
coasts of southern Florida in summer, although large ones at least are unknown there in
winter, marks it as characteristically a tropical species.

Synonyms and References :''

Le Marteau, Duhamel, Traite Gen. Peches, (2) 4 (9), 1777: 303 (in part), pi. 21, fig. 8 (ident. by shape of
head) ; Broussonet, Mem. Math. Phys. Acad. Sci. Paris (1780), 1784: 661 (in part, by ref. to Duhamel,

1777)-

No species name, Parra, Desc. Piez. Hist. Nat. Havana, 1787: 71, pi. 32 (ident. probable from ill.).

Squale pantouflier, Lacepede, Hist. Nat. Poiss., 4° ed., 7, 1798: 167, 261, pi. 7, fig. 3 (ident. from ill., Cay-
enne) ; in Sonnini, Hist. Nat. Poiss., 4, 1802: 81 (from Lacepede, 1798).

Squale Marteau, Lacepede, Hist. Nat. Poiss., 4° ed., i, 1798: 257 (part, because teeth descr. as serrate, but not
pi. 8, fig. 3, which hzygaena).

Hammer Headed Shark, Shaw, Gen. Zool., 5 (2), 1 804: 354, pi. 154 (ident. from descr. of teeth, and ill.,
Medit., but report for Indian seas and Tahiti perhaps not this species).

Squalus tiburo Risso, Ichthyol. Nice, 1810: 35 (ident. by descr. of head and ref. to Lacepede, Squale pantou-
flier, as above, Nice, very rare) ; not Squalus tiburo Linnaeus, 1758.

Zygaena tudes Valenciennes, Mem. Mus. Hist. nat. Paris, 9, 1822: 225, pi. 12, fig. i (type descr., ill. of
head, Medit., Cayenne, but Coramandel record perhaps not this species) ; Risso, Hist. Nat. Europ.
Merid., 5, 1826: 126 (ident. by ref. to Valenciennes, 1822); Bory de St. Vincent, Diet. Class. Hist.

31. The Zygaena tudes pictured by Philippi (An. Univ. Chile, 7/, 1887: pi. 2, fig. 4.) from Chile vpas actually
of the diflana group, to judge from the shape of its head.

32. Ribeiro, Arch. Mus. nac. Riu di; J., i^, 1907: pi. 5.

33. Because of the uncertainty as to whether or not sharks reported from the Pacific and Indian Oceans as tudes are
actually identical with the Atlantic Hammerhead of that name, the following is limited to citations for the
Atlantic. For Indo-Pacific citations of tudes, see Fowler (Bull. U.S. nat. Mus., 100 ['iL 1941 : ^n) ^"d Beebe
and Tee-Van (Zoologica, N. Y., 26 [2], 1941: 115).

Fishes of the Western North Atlantic 435

Nat., IS, 1829: 596 (ref. to Valenciennes, 1822); Voigt, in Cuvier, Tierreich, 2, 1832: 514 (ref. to
Valenciennes, 1822); Agassiz, L., Poiss. Foss., 3, 1835-1843: 91, pi. E, fig. 8 (teeth); Yarrell, Brit.
Fish., 2, 1841 : 507, 508, fig. I (ill. after Valenciennes, 1822) ; Gunther, Cat. Fish. Brit. Mus., 8, 1870:
382 (refs. in synon., but ident. of descr. specs, doubtful because of long prcnarial groove) ; Moreau, Hist.
Nat. Poiss. France, /, 1881: 327 (ident. by descr., Nice, France); Doderlein, Man. Ittiol. Medit., 2,
l88i: 50 (ident. by refs., Medit., occur.);'* Perugia, Elenc. Pesc. Adriat., 1881: 53 (name only, Adri-
atic) ; Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 4; ; Faune Senegambie, Poiss., /, 1 883-188 5:
22 (refs. in synon., but ident. of W. Afr. specs, doubtful) ; Belloc, Rev. des Trav. Peches Marit., 7(2),
1934: 133 (ill. head), 134 (Morocco to Senegal; Canaries).

Sfhyrnichthys zygaena Thiencmann, Lehrb. Zool., 5, 1828: 408 (general, ident. probable because teeth
descr. as "gezahnelte").

Sfhyrna malleus L. Agassiz, Poiss. Foss., 3, 1835-1843: 12, 13, pi. E, fig. 7, pi. N, fig. 8 (ident. probable
because teeth serrate).

Sfhyma tudes Miiller and Henle, Plagiost., 1841: 53 (ident. from ref. to Valenciennes, 1822, Cayenne, but
Coramandel record perhaps some other species); Bonaparte, Cat. Pesc. Europ., 1846: 18 (name only,
ident. from included refs.); Canestrini, in Cornalia, et a!., F.iuna d'ltal., 5, 1871-1872: 47 (ident. by
inference, Nice) ; Doderlein, Atti Accad. Palermo, N. S. 6, 1878: 30 (name only, Sicily) ; Jordan, Rep.
U.S. Comm. Fish. (1885), 1887: 797 (name only, Europ., W. Indies, but trop. Pacif. ref. perhaps not
this species); Carus, Prod. Fauna Medit., 2, 1 889-1 893: 544 (name only, Medit.) ;*' Ribeiro, Arch.
Mus. nac. Rio de J., /.;, 1907: 201 (synon. in part, but not S. tudes, p. 157, pi. 5, which appears to
be diflana, see p. 41;); Rey, Fauna Iberica Peces, 1, 1928: 365, fig. 2 (head, after Valenciennes,
1822, Spain) ; Borri, Atti Soc. tosc. Sci. nat., 44, 1934: 91 (name only, Lagos, W. Afr.) ; Fowler, Bull.
Amer. Mus. nat. Hist., yo (l), 1936: 64 (ident. from descr. of young, Surinam); Puyo, Bull. Soc.
Hist. nat. Toulouse, 70, 1936: 84 (name only, French Guiana); Norman and Eraser, Giant Fishes,
1937: 47 (general) ; Springer, Proc. Fla. Acad. Sci., 3, 1939: 32 (first adequate descr. since Valenciennes,
1822, size, no. of embryos, occur. SW. Florida) ; Stanford Ichthyol. Bull., 1(5), 1940: 162, 164 (comp.
with other species, changes in relative proportions with growth); Proc. Fla. Acad. Sci., 5, I94i- 49
(discuss., ill. of sicull) ; Fowler, Bull. U.S. nat. Mus., \oo {13), 1 941 : 213 (Atlant. citations in synon., but
list of citations of tudes for Indo-Pacific and ident. of described Pacific specs, doubtful) ; Arqu. Zool.
Estado Sao Paulo, 3, 1942: 129 (listed, Brazil) ; Lunz, Bull. S. Carolina St. Planning Bd., 14, 1944: 27
(listed, Florida) ; Bigelow and Schroeder, Guide Comm. Sharic Fish., Anglo Amer. Caribb. Comm.,
Wash., 1945: 138, fig. 51 (descr., range, ill.); Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1945: 97
(Beaufort, N. Carolina).

Sfhyrna chiereghini Nardo, Sinon. Modern. Spec, descr. Pesci St. Chiereghin, 1847: III (name); Atti 1st.
veneto., (3) 4, i860: 787 (name only, Medit.); Perugia, Cat. Pesc. Adriat., 1866: 7 (name only,
Adriatic).

Sfhyrnias tudes Gray, List Fish. Brit. Mus., 1851: 50.

Cestracion {Zygaena) tudes Dumeril, Hist. Nat. Poiss., i, 1865: 384 (descr. from Valenciennes, 1822, Nice,
Algiers, Cayenne, Demerara? ).

Squalus zygaena de la Blanchcre, Diet. Peches, 1868: 499 (at least part, because teeth descr. as "dentelees").

Cestracion tudes Ninni, Revist. Critica Pesc. Adriat. descr. Chiereghini, 1872: 10 (not seen) ; Garman, Mem.
Mus. comp. Zool. Harv., 36, 1913: 1 59 (synonymy, but Brazilian spec, descr. is S. bigelotvi, see p. 414).

Sfhyrna [Platysqualus) tudes Joidm and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 217 (name, W.
Indies, Medit., but ref. to Gulf of Calif, and Indian Ocean perhaps not this species) ; Bull. U.S. nat.
Mus., 47 (1), 1896: 44 (refs., descr., Medit., W. Indies, but ref. to Gulf of Calif, and Indian Ocean
perhaps not this species) .

Cestracion zygaena Radcliffe, Bull. U.S. Bur. Fish., 34, 1916: 263 in part, pi. 3, fig. 2 (ident. by descr. of
teeth of ill. spec, as serrate, 12-ft. 6-in. spec, Beaufort, N. Carolina).

Sfhyrna zygaena Coles, Copeia, 69, 1919: 41 in part, pi. 3, fig. 2 (ident. by shape of head in photo, of i 3-ft.
lO-inch female, C. Lookout, N. Carolina and meas. of same) ; not Sjualus zygaena Linnaeus, 1758.

34. See Doderlein for additional Mediterranean locality records in publications not accessible to us.

35. See Carus for additional Mediterranean records in publications not accessible to us.

4.36 Memoir Sears Foundation for Marine Research

Platysqualus tudes Howell-Rivero, Proc. Boston Soc. nat. Hist., 41 (4), I 936: 44 (ident. probable because of
large size, I 2 to 14 ft., Cuba).

Doubtful References:

Zy gaena tudes Cuvier, Regne Anim., ill. ed., 1843: pi. 1 17, fig. I ; Osorio, J. Sci. math. phys. nat. Lisboa, (2)
J, 1894: 182 (name only, C. Verde Is.); J. Sci. math. phys. nat. Lisboa, (2) 5, 1898: 200 (name only,
C. Verde Is.) ; Metzelaar, Trop. Atlant. Visschen, 191 9: 189 (by ref. to Rochebrune and Osorio as above,
trop. W. Afr.).

Sphynia tudes Miillcr and Troschel, in Schomburgic, Reisen Brit. Guiana, 1840—44, 5, I 848: 642 (one re-
ported as seen, Demerara) ; Machado, Feces Cadiz, 1 857: 9 (Spanish coast, name only) ; Berg, Ann. Mus.
nac. B. Aires, (2) i (4), 1895: 8 (Montevideo, Muldonado, no evidence for ident.); Fowler, Proc.
Acad. nat. Sci. Philad., 60, igo8: 66 (meas., 8-in. spec, not diagnostic, and ident. doubtful because of
small size, Surinam) ; Ribeiro, Rev. Mus. paul., 1918: 107 (name only, Santos, Brazil) ; Devincenzi, An.
Mus. Hist. nat. Montevideo, (2) / (4), 1920: 119 (name only, Uruguay) ; An. Mus. Hist. nat. Monte-
video, (2) (6), 1925: 323 (name only); Marelli, Elcnc. Sist. Fauna B. Aires (1922—23), 1924: 546
(name only, Uruguay, Argentina); Pozzi and Bordale, An. Soc. cient. argent., /20, 1935: 151 (name
only, north. Argentina).

Cestracion tudes Meek and Hildebrand, Field Mus. Publ. ZooL, 15 (l), 1923: 59 (descr. appears to be pardy
tiburo).

Not S-pkyr?ia tudes Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907: 157, pi. 5 (photo of head appears to be
dif.ana, see p. 420, Maria Farinha, Pernambuco, Brazil) ; Arch. Mus. nac. Rio de J., Fauna brasil. Peixes,
2(1) Fasc. I, 1923: 14 (same record as foregoing).

Sf hyrna zy gaena (Linnaeus), 1758

Common Hammerhead

Figures 85, 86

Study Material. Eleven specimens of both sexes, from 510 to about 1,780 mm. long,
from: Muldonado, Uruguay; Rio de Janeiro, Brazil; New York; off Nahant in Massa-
chusetts Bay; Cape Cod; and off Woods Hole, Massachusetts (Harv. Mus. Comp. ZooL).
Others from Virginia Beach, Virginia, and Buzzards Bay, Massachusetts (U.S. Nat.
Mus.). Specimens from Woods Hole, Massachusetts (U.S. Bur. Fish.). Several others
caught in the offing over the continental shelf by members of the Woods Hole Oceano-
graphic Institution and identified by us, and ten small specimens from Peru, Panama, the
Galapagos, Lower California and Japan (Harv. Mus. Comp. ZooL, No. 382, 383, 421,
441, 515, 1042, and U.S. Nat. Mus., No. 51289, 51291, 71774, 77711).

Distinctive Characters. The very differently shaped head is enough to separate
zygaena at a glance from tiburo (cf. Fig. 86 A with 82 B). It is easily distinguished from
diplana, bigelowi and tudes by the evenly convex anterior outline of the head in the mid-
line (not indented) ; further from diplana by the additional differences summarized under
that species (p. 415); further from bigelowi and tudes by the much more slenderly acu-
minate tip of its second dorsal fin, and further from tudes by its much more erect first
dorsal fin.

Description. Proportional dimensions in per cent of total length. Male, 524 mm.,

Fishes of the Westerfi North Atlantic 4.37

from New York (Harv. Mus. Comp. Zool., No. 136). Female, 687 mm., from Nahant,
Mass. (Harv. Mus. Comp. Zool, No. 1159).

Trunk at origin of pectoral: breadth 8.2, 8.9; height 9.0, 10.2.

Snout length in front of: outer nostrils 5.9, 5.45 mouth 6.9, 6.1.

Eye: horizontal diameter 2.4, 2.2.

Mouth: breadth 7.6, 7.1 j height 4.2, 4.4.

Nostrils: distance between inner ends 19.7, 20.6.

Labial furrow length: lower 0.5, 0.6.

Gill opening lengths: ist 2.2, 2.5; 2nd 2.4, 2.5; 3rd 2.5, 1.6; 4th 2.5, 2.5; 5th

2.0, 2.1.

First dorsal fin: vertical height ii.i, 12.4; length of base 9.7, 9.5.

Second dorsal fin: vertical height 2.4, 2.35 length of base 3.2, 3.3.

Anal fin: vertical height 3.7, 3.5; length of base S-2ij 4-6.

Caudal fin: upper margin 30.2, 30.6 j lower anterior margin 1 1.6, 13.2.

Pectoral fin: outQv mzrgm 12.O, 12.55 inner margin 4.4, 4. 1 5 distal margin 9.2, 9.5.

Distance frotn snout to: ist dorsal 27.3, 28.35 2nd dorsal 60.2, 60.55 upper caudal

69.8, 69.45 pectoral 22.1, 21.35 pelvics 47.5, 46.55 anal 57.8, 58.0.

Figure 85. Sfhyrna zygaena, female, 687 mm. long, from Nahant, Massachusetts (Harv. Mus. Comp.
Zool., No. 1 1 59). A Upper and lower teeth, left-hand side, about 2.5 x. B Second upper tooth. C Ninth
upper tooth. D Third lower tooth. E Ninth lower tooth. B-E, about 5 x.

438

Memoir Sears Foundation for Marine Research

Figure 86. Sfhyrna zygaena, illustrated in Fig. 85. A Head from below, about 0.6 x natural size. B Second
dorsal and anal fins. C Dermal denticles, about 60 x. D Fourth upper tooth. E Fifth lower tooth. D-E, about
12 z.

Fishes of the Wester ?t North Atlantic 439

Interspace between: ist and 2nd dorsals 23.6, 25.3; 2nd dorsal and caudal 7.8,
7.0; anal and caudal 7.8, 7.0.

Distance from origin to origin of: pectoral and pelvics 26.5, 27.O; pelvics and anal
12.0, 12.8.

Trunk strongly compressed, its height at ist dorsal about V^ (18 to 19%) of its
length to origin of caudal. Back smooth, without mid-dorsal ridge. Caudal peduncle
about % as wide as high, the upper precaudal pit strongly developed and subtriangular,
lower lacking in smallest specimens examined but weakly indicated in larger. Dermal den-
ticles so close-spaced and overlapping that skin is mostly concealed, the blades thin and
moderately arched, about as broad as long, small specimens usually with 3 ridges but large
ones with 5 and sometimes 7 ridges extending about halfway back from the anterior mar-
gin, 3 to 5 marginal teeth, the median considerably the longest; pedicels very short.

Head about Vs (37%) of length of trunk to origin of caudal, its dorsal profile
slightly and evenly convex from origin of ist dorsal to eyes, but concave thence forward,
its anterior edge very thin, very broadly expanded laterally in hammer form, with
outer posterior margins, outward from the neck, directed slightly toward the rear;
breadth at eyes about 1.2 to 1.4 times its length to origin of pectoral, and its length at
oculo-narial prominence between V3 and V4 its breadth, much as in diplana; anterior mar-
gin of head scalloped with a deep depression opposite each nostril, and with a shallow con-
cavity midway between latter and the median line but with the midsector evenly and
rather strongly convex (indented in diplana, p. 415) ; a well marked groove from nostril
inv/ard along anterior margin of head a little more than halfway toward the midline; dis-
tance from anterior corner of oculo-narial prominence to anterior edge of eye only about
V2 to % 0 as great as diameter of latter (about as great as diameter of eye in diplana) ; a
line connecting outer ends of nostrils passes anterior to mouth by a distance about V2 as
great as diameter of eye, one connecting centers of eyes passes a little behind front of upper
jaw (through front of upper jaw in diplana), and one connecting outer posterior corners
of hammer passes posterior to corners of mouth by a distance about V2-% as great as diam-
eter of eye. Head (snout) in front of mouth a little less than Ys (about 26 to 3 1 %) as long
as head to origin of pectoral. Rostral cartilage usually without median hole, and wings of
preorbital processes without inwardly directed point on anterior margin (there is a median
hole and such a point in diplana). Mucous pores in median sector on oral side of head cover
a subtriangular or irregular area near its anterior margin (a subrectangular or dumbbell-
shaped area in diplana) . Eye a little broader than high and much larger relatively than in
tudes or tiburo, its horizontal diameter about Vs as long as head in front of mouth. Gill
openings rather strongly concave in outline and about evenly spaced, the ist about equal
to diameter of eye in small specimens (a little more than i V2 times as long as diameter of
eye in diplana of equal size) but about i V2 times in large, the 3rd only a little longer than
1st, 5th equal to it or a little shorter, the space between 4th and 5th over origin of pectoral.
Nostril sloping a little forward from inner end to outer. Mouth strongly arched, a little

440 Memoir Sears Foundation for Marine Research

more than ' j as high as broad, with a very short labial furrow on lower jaw from corner
of mouth, but none on upper jaw.

Teeth "i2°t'o^H— °i— il'o 'V^ '"■ specimens counted, uppers triangular, the cusps smooth-
edged in young, but tending to become weakly serrate with growth,^^" the bases more or
less fluted or wavy on outer side of some teeth but not on others j the ist upper tooth nearly
symmetrical and erect, but subsequent upper teeth strongly oblique, their inner margins
straight or slightly convex, their outer margins deeply incised, even the outermost with
cusps well developed though very small j lower teeth similar to uppers but a little smaller,
smooth or very weakly serrate, the 4 next to the center of mouth with narrower cusps and
less strongly oblique; i small symmetrical tooth at symphysis of lower jaw, one or none in
upper in specimens examined; i (in places 2) series functional along sides of mouth and 2
to 3 series toward center in upper jaw; 2 to 3 series all along lower jaw.

First dorsal perhaps averaging a little more sloping than in diflana, a perpendicular
from its apex falling a little anterior to its tip, its origin a little posterior to axil of pectoral,
its vertical height a little more than Y2 (50 to 58%) as great as length of head, its
anterior margin only slightly convex toward apex, posterior margin moderately concave
toward base, apex very narrowly rounded, its free rear corner only moderately slender,
a little less to a little more than ^/"s as long as the base, the midpoint of its base about
2.5 times as far from origin of pelvics as from axil of pectoral. Second dorsal about Va ^s
long at base as ist, its origin about over midpoint of base of anal, its apex abruptly rounded,
rear margin moderately concave, its free rear corner very slender, a little longer than the
base. Caudal nearly Yi of total length, its upper margin varying from very weakly con-
vex to a little more strongly so toward base and tip; the terminal sector a little more than
Viy the length of fin, slender, with narrowly rounded tip, its lower posterior margin only
weakly concave (less deeply so in specimens seen than in diflana), the lower lobe a little
more than y^ as long as upper with weakly convex anterior margin, nearly straight pos-
terior margin, and narrowly rounded tip, the re-entrant contour, enclosed by the 2 lobes,
approximately a right angle, with rather broadly rounded corner. Anal about i Vs to i V2
times ( 1 .3 to 1.6) as long at base as 2nd dorsal and a little higher, its anterior margin more
convex, apex acute, rear margin m.uch more deeply incised in angular contour, its free rear
corner about as long as the base or a little longer. Distance from origin of anal to tips of
pelvics I '/'s to I V2 times as long as base of former. Pelvics about as long at base as anal.
Pectoral a little less than % (59 to 64%) as long as head, about as long as vertical height
of ist dorsal or a little longer, a little smaller than ist dorsal in area, a little more than V2
as broad as long, the outer margin nearly straight, distal margin only slightly concave, the
apex and inner corner narrowly rounded.

35a. Stewart Springer informs us that in large zygaena the upper teeth may have noticeable serrations while the
lowers may have extremely fine serrations or none. In a specimen about six feet long (1,780 mm.), caught in
August 191.6 off Woods Hole, Massachusetts (Harv. Mus. Comp. Zool., No. 36425), the teeth, both uppers
and lowers, vary from smooth to very weakly serrate or slightly irregular.

Fishes of the Western North Atlantic 441

Color. Deep olive leaden or brownish gray above, paler on sides, shading into pure or
grayish white below; fins of same color as back or sides, with tips or margins more or less
dusky; pectorals black-tipped in some specimens, but not in others.

Size. Reports of embryos of 450 to 460 mm. and of free-living specimens of only
510 to 590 mm. suggest a length of about 500 mm. at birth. Adults seemingly do not
mature at less than 7 to 8 feet, are often taken at 9 to 1 1 feet in length, and occasionally
12 to 13 feet. It seems likely that the still larger Hammerheads that are reported from
time to time are in reality the Great Hammerhead (ludes). The following weights of
Hammerheads of different lengths probably refer to zygaena: 1,651 mm., 57 pounds
(26 kilo); 3,610 mm., 836 pounds (380 kilo); and 3,810 mm., 900 pounds (409 kilo).'"

Developmental Stages. Presumably development is ovoviviparous; at least the pres-
ence of a placenta has never been reported. As many as 29 to 37 embryos have been re-
ported repeatedly as being found in a single female.

Habits. This is a strong-swimming shark, often seen at the surface with the tips of its
first dorsal and caudal fin exposed. But we have never seen or heard of one jumping clear
of the water. They are to be met with indifferently far out at sea, or so close in to the shore
that considerable numbers are often taken in beach seines or in pound nets. On the West
African coast they (or they and diflana) are common in salt-water lagoons; they are also
reported in similar situations {e.g., Indian River) in Florida. There is even one report of a
Hammerhead of some sort in tidal fresh water in Maryland."

The diet of the zy gaena-diflana group consists chiefly of fish. Inshore, in the southern
part of their range, they feed largely on sting-rays, which they are often seen chasing and
which have been recorded frequently from their stomachs. Sting-ray spines are also found
embedded in the jaws of Hammerheads, as are the spines of the gaff-topsail catfish
{Felichthys). Often the stomach contents of the larger ones contain parts of other sharks,
or entire small ones, including their own kind. Where net fisheries for sharks are carried
on in warm waters it is their common habit to devour the sharks that are entangled in
the nets. In the northern part of their range, skates are a major item in their diet; a
Hammerhead of 1,780 mm. caught off Woods Hole in August 1944 had in its stomach one
seven-inch scup {Stenotomus) ; they are also known to prey on herring and bass in waters
of North Europe, and on Spanish mackerel {Scomberomorus maculatus) and menhaden
(Brevoortia) in North America; no doubt they also feed on any other fishes that may be
available locally. Their recorded diet also includes shrimp, crabs, barnacles, and Crustacea
generally, as well as squid.

Relation to Man. The considerable number taken in the shark fisheries of southern
Florida and the West Indies are utilized for leather, fish meal and liver oil. They also
afford good sport to any angler who chances to hook one, for they bite freely and have
been described (we have never taken one on light tackle) as so lively that one has been
known to die of exhaustion when hooked.

36. Schultz, J. Mammal., /p, 1938: 484. 37. Gunter, Amer. Midi. Nat., 28, 1942: 316.

442 Memoir Sears Foundation for Marine Research

Range. Tropical to warm-temperate belts of the Atlantic; north regularly to Portu-
gal and occasionally to the English Channel, Welsh coast and Scotland, in the east;
Mediterranean; Azores; Madeira; TenerifFe; Canaries; Cape Verde Islands; Morocco;
Dakar; tropical West Africa and South Africa. In the west, north commonly to southern
New England and as a stray to Massachusetts Bay and Nova Scotia; south to Uruguay and
(nominally) to northern Argentina. It is also widespread in the tropical and warm-tem-
perate Pacific*' and probably in the Indian Ocean as well. But a more precise statement as
to the Indo-Pacific range of zygaena must await critical study of the Hammerheads as a
whole in that region.

Occurrence in the Western Atlantic. It is not possible to present a satisfactory picture
of the distribution of S. zygaena in the western Atlantic from existing literature be-
cause of the recent discovery that many of the older accounts that ostensibly referred to
it may also have covered its companion species, d'l-plana (p. 419). In fact, the only western
Atlantic localities where the presence of zygaena (not including diplana or tudes) is posi-
tively established by pertinent information, either verbal or pictorial, as to shape of head,
teeth, relative proximity of the tip of second dorsal fin to caudal, or shape of skull, are:
Nahant in Massachusetts Bay; Cape Cod; southern Massachusetts in the general vicinity
of Woods Hole*" and the continental shelf in its offing; vicinity of New York; several
localities along the New Jersey coast; near Beaufort and Cape Lookout, North Carolina;
southern Florida on both coasts; the Virgin Islands; southern Brazil; Uruguay; and
(nominally) northern Argentina. But this is enough to prove it widespread all along the
American seaboard in low and midlatitudes.

No doubt it is also responsible, at least in part, for the frequent reports of Hammer-
heads for the West Indian-Caribbean region, i.e., Porto Rico, Haiti, Jamaica, Cuba, Turks
Island, Trinidad, Dutch, French and British Guianas, Venezuela, the Atlantic coast of
Panama and likewise for Bermuda. Although definite information is lacking for the south-
ern part of the Gulf of Mexico, it is to be expected there generally, and no doubt it visits
the northern coast of the Gulf, at least in small numbers, for there are a few records of
Hammerheads for Louisiana.

The most spectacular aspect of the occurrence of Hammerheads is their migration
northward (often in schools) in summer along the Atlantic seaboard, both inshore and out
along the continental shelf. Zygaena is chiefly responsible for this seasonal movement, the
most northerly records for diplana being from the offing of Delaware Bay (well offshore),
and for tudes from North Carolina (p. 434), whereas many zygaena have been posi-
tively identified from New Jersey to southern New England. For example. Hammerheads
of one sort or another are moderately common during the summer months near Charles-

41. Specimens that we have examined from Peru, Panama, the Galapagos, Lower California and Japan do not differ
in any significant respect from the Atlantic specimens with which we have compared them in regard to teeth, pro-
portionate dimensions, shape of head, or fins. Hussakoff (Copeia, 34, 19 16: 63) had already reached the same
conclusion for Japanese specimens.

42. A nine-foot specimen, taken in a fish trap in Buzzards Bay on August 6, 1934, and identified by a good photo-
graph published in the Boston Globe, is one of several well attested records for the region.

Fishes of the Western North Atlantic 443

ton, South Carolina 5 they are among the more plentiful of summer sharks in the vicinity
of Cape Lookout, where there is record of as many as 65 taken in a single haul of a seine 5
and numbers of them enter the shoal North Carolina sounds in some summers but only
occasionally in others. Although they appear as strays only in Chesapeake Bay, Hammer-
heads are common summer visitors to the Atlantic coasts of Maryland, New Jersey and
New York, where they are present yearly from July until October in varying numbers,
not only offshore but in the shallow coastal bays as well. They even enter New York Har-
bor occasionally} in fact one of the largest Hammerheads on record north of Cape Hatteras
(about 1 1 feet) was taken many years ago at the head of Peconic Bay, Long Island. Farther
east along southern New England they appear less often inshore, although there are a few
records for Connecticut and Rhode Island, as well as many reports of them at Woods
Hole and Nantucket for summer and early autumn (July to October) ; we obtained a 6-
foot female from a trap near Woods Hole in August 1944. But Hammerheads basking at
the surface are familiar objects a few miles offshore along this sector, as we can bear
witness, and they are brought in from time to time by tuna and other fishermen. There
are also a few records for the tip of Cape Cod, one for the inner part of Massachusetts Bay,
and for Casco Bay, Maine, where the capture of two small ones, no doubt zygaena, has
been reported to us." But the longitude of Cape Cod so sharply bounds their coastwise
dispersal in this direction that there are only two records of Hammerheads farther east
on the continental shelf, one for Halifax, Nova Scotia, the other (a 12-foot specimen
caught in August 1928) between Georges and Browns Banks. Outside the edge of the con-
tinental shelf, in the sweep of the Gulf Stream, Hammerheads (probably both zygaena and
diplana) are to be expected much farther to the east and north, perhaps even past the New-
foundland Banks.

The great majority of individuals sharing in the summer movement northward are
small (less than 6 feet). In fact, many of them are so small as to suggest that they were
born only a short time previously. Dozens of little ones, of about 2V2 feet, have been
seined on the outer shore of Long Island in August. This has led to the suggestion that
they are born in northern coastal waters, but on the other hand large ones are seldom taken
near shore along our northern coast,** making it more likely that whatever production of
young there may be in the northern sector of the range takes place well offshore.

Off New York and to the eastward Hammerheads usually disappear when the tem-
perature of the water falls below 67° F., i.e., by late September or early October. Occa-
sionally, however, one lingers into November, and there is even one record for February
in Long Island Sound. It is not known if the Hammerheads that reach the shores of the
northeastern United States in the summer migrate south again in the autumn, or if they
merely move offshore to escape falling temperatures and are then picked up by the sweep
of the Gulf Stream and so lost to the parent population.

Information as to the status of zygaena coastwise in South American waters south-

+3. W. H. Rich saw these in the fish market at Portland, Maine.

44. Exceptions to this rule are specimens 1 1 feet i inch, from North Carolina, of 1 1 feet from Long Island, New
York, and 9 feet 10 inches from Rhode Island.

4-44 Memoir Sears Foundation for Marine Research

ward from the Caribbean is much less extensive than for the United States coast. But since
zygaena is known positively from as far south as Uruguay, it is undoubtedly responsible
at least in part*^ for the nominal reports of the species from Santos, Pernambuco and
Maceio; i.e., it occurs generally along the coast of Brazil. But whether or not it expands
its range yearly to the southward during the warm season in the southern sector of its range,
as it does to the northward in the northern sector, is not yet known.

Synonyms and References:

Atlantic:"

Squalus zygaena Linnaeus, Syst. Nat., 1758: 234 (by ref. to ill. by Willoughby, 1686; see above, p. 409;
type loc. Europe, Amer.) ; Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: 9, pi. 6, fig. 15 (ident. by
ref. to Linnaeus, 1758; descr. and ill. would apply equally to diftana; Atlant. and Medit.) ; Gmelin, Syst.
Nat., I, I 789: 1494 (ident. by ref. to Linnaeus, 1758) ; Bloch, Naturg. Ausland. Fische, /, 1785 : 29, pi.
117, upper fig. (ill. apparently this species, but lower fig. is Eusphyra blochii; N. Europ., Antilles,
Medit.); Bloch and Schneider, Syst. Ichthyol., 1801: 131 (ident. by ref. to Linnaeus, 1758); Bosc,
Nouv. Diet. Hist. Nat., 21,1 803: 189 (diagn., ident. from included refs.) ; Risso, Ichthyol. Nice, 1810:
34 (ident. by ref. to Linnaeus, 1758; general, Medit.); Mitchill, Trans. Lit. Phil. Soc. N. Y., i,
1815: 482 (li-ft. spec, its stomach containing parts of a man; ident. probable from loc, Riverhead,
N. York) ; Yarrell, Brit. Fish., Suppl., 2, 1839: 62 (ident. by ill. of head, Norfolk, Eng.) ; Gray, Cat.
Fish. Coll. Descr. by L. T. Gronow in Brit. Mus., 1854: 6 (at least in part, ident. by included refs.).

Balance Fish, Brookes, Nat. Hist., j, I 763: 31, pi. 28, fig. 3 I ; also 3rd ed., 1772: I 8, pi. 19, fig. 18 (ident.
by ill., general).

Le Marteau, Broussonet, Mem. Acad. Math. Phys. Sci. Paris (1780), 1784: 661 (at least in part, by included
refs.).

Squalus malleus Shaw and Nodder, Naturalist Misc., 8, 1796: 375, pi. 267 (ident. by ill., Medit.).

Le squale marteau, Lacepede, Hist. Nat. Poiss., 4° ed., /, I 798: 156, 257, pi. 8, fig. 3, in Buffon, Hist. Nat.
(in part, ident. by ill. of head, but also includes tudes, see p. 434, general) ; in Sonnini, Hist. Nat. Poiss.,
4, 1802: 74, pi. 8, fig. I, 2 (after Lacepede, 1798).

Sfhyrna zygaena Rafinesque, Indice Ittiol. Sicil., 1810: 46 (ident. by included refs., Sicily) ; Bonaparte, Icon.
Faun. Ital., j, 1840: pi. not numbered, fig. 1, and index (descr., ident. by ill., arrangement of pores on
head, Italy) ; Miiller and Henle, Plagiost., I 841 : 51 (descr., ident. by proportional meas., Medit., France,
Brazil, India); Cuvier, Regne Anim., ill. ed., 2, 1843: 368, pi. 1 1 7, fig. I (ident. by ill., but ref. in
text is to pi. I 16, fig. I, which is a torpedo, Narcacion) ; Bonaparte, Cat. Pesc. Europ., I 846: 18 (ident. by
included refs.); Fitzinger, Bild. Atlas Naturg. Fische, 1854: fig. 170 (ill., ident. by shape of head);
Nardo, .Atti 1st. veneto, 5, i860: 787 (at least in part from refs. and loc, near Venice) ; Brito Capello, J.
Sci. math. phys. nat., Lisboa, (1868-1869), 2, 1869: 141 (at least in part, ident. by inference from incl.
refs., Portugal) ; Canestrini, in Cornalia, et at., Fauna d'ltal., I 870— 1874: 47 (at least in part, ident. by
inference from included ref.) ; Gegenbaur, Unters. Vergl. Anat. Wirbelt., 5, 1872: 23, pi. 9, fig. I (ident.
by ill. of skull) ; Perugia, Elcnc. Pesc. Adriat., I 88 l (at least in part, ident. by included refs.) ; Jordan and
Gilbert, Bull. U.S. nat. Mus., 16, 1883: 26 (at least in part, ident. from range, C. Cod, southward);
Kingsley, Stand. Nat. Hist., 5, 1885: 81 (general, ident. from ill.); Jordan, Rep. U.S. Comm. Fish.
(1885), 1887: 797 (at least in part from range, N. Atlant. states to W. Indies, and California southward) ;
Bean, Bull. U.S. Fish Comm., 7, 1888: 151 (at least in part, because of descr. as abund., N. Jersey);
Carus, Prod. Fauna Medit., 2, I 889-1 893: 513 (at least in part, because of included refs., Medit.) ; Nel-

+5. Probably diflana also.

46. The following list is limited to citations that include information, verbal or pictorial, sufficient to make
it reasonably certain that they were based at least in part on zygaena and not on diflana, or that refer to localities
where zygaena is known to be common but not diflana. For a list of citations, nominally to zygaena, but which
have referred to diflana in reality, see p. 420. For an extensive list of Indo-Pacific citations that certainly or
possibly refer to this species, see Fowler (Bull. U.S. nat. Mus., 100 [r^], 1941 : 217).

Fishes of the Wester 7i North Atlantic 445

son, Rep. St. Geol. N. J., 2 (2), 1890: 661 (occasional, N. Jersey, idcnt. probable because of loc.) ;
Almeida and Roquete, Mammif. Peix. costa e rios do Algarve Inquerito Indust. Lisboa (1889), 2,
1892: 374 (at least in part because of loc, Portugal, not seen) ; Jordan and Evermann, Rep. U.S. Comm.
Fish. (1895), 1896: 217 (at least in part from range, C. Cod and C. Conception southward); Bull.
U.S. nat. Mus., 47 (i), 1896: 45 (descr. not diagnostic, but zygaena at least in part because of range,
C. Cod and C. Conception southward); Bean, Bull. Amer. Mus. nat. Hist., 9, 1897: 329 (at least
in part from loc. near N. York, Aug. and Sept.); Mearns, Bull. Amer. Mus. nat. Hist., 10, 1898: 311
(ident. probable because of loc, Hudson River); Smith, Bull. U.S. Bur. Fish., ly, 1898: 88 (ident.
probable because of loc, Woods Hole); Evermann and Kendall, Rep. U.S. Comm. Fish. (1899),
1900: 49 (at least in part from loc, Indian R. to Key West, Florida) ; Sharp and Fowler, Proc Acad,
nat. Sci. Philad., 56, 1901: 505 (ident. probable because of loc, Nantucket); Linton, Bull. U.S. Bur.
Fish., rp, 1 901: 272 (food, parasites, ident. probable because of loc, Woods Hole) ; Bean, T. H., Bull.
N. Y. St. Mus., 60, Zool. 9, 1903: 31 (ident. probable because of loc. near N. York) ; de Braganza, Result.
Invest. Sci. "Amelia," 2, 1904: 46, 47 (name only, Portugal, ident. probable because of loc); Tracy,
Rep. R. I. Comm. inl. Fish., 1906: 45 (ident. probable because of loc, Rhode Island); Pietschmann,
Ann. naturh. (Mus.) Hofmus. Wien, 21, igo6: 99 (descr. not diagnostic, but ident. probable, in part
at least, because of loc, abund., Morocco); Linton, Bull. U.S. Bur. Fish., 26, 1907: 122 (ident. prob-
able because of loc. Woods Hole, August) ; Smith, Bull. N. C. geol. econ. Surv., 2, 1907: 36 (ident. by ill.,
N.Carolina) ; Ribeiro, Arch. Mus. nac Rio de J., 14, 1907: 157, 201 (at least in part, descr. and loc. north,
and south. Brazil, equally applicable to diflana, synonyms) ; Kendall, Occ Pap. Boston Soc nat. Hist.,
7 (8), 1908: 5 (ident. probable from loc, south. New England); Fowler, Rep. N. J. Mus. (1907),
1908: 127 (ident. probable because at many N. Jersey loc, also Delaware Bay); Proc. Acad. nat. Sci.
Philad., 60, 1908: 66 (ident. probable because of loc, N. Jersey, Nantucket by name, also for Italy,
Surinam, Padang) ; Tracy, Rep. R. I. Comm. inl. Fish., 1910: 59 (Rhode Island and Connecticut,
ident. because of loc); Giinther, Encyc. Brit., I ith ed., 24, 191 1 : 807 (at least in part, general);
Nichols, Abstr. Linn. Soc. N. Y., 20—23, 1913: 91 (ident. probable because of loc. near N. Y.) ;
Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., 31 (2), 1913: 736 (ident. because of loc. Woods
Hole); Murphy and Harper, Copeia, 23, 1915: 41 (Montauk, Long Island, ident. probable because of
loc.) ; Coles, Proc. biol. Soc. Wash., 28, 191 5: 90 (at least in part, abund. off C. Lookout, N. Carolina,
but probably covers tudes also); Hussakoff, Copeia, 34, 1916: 63 (Woods Hole spec, compared with
Japanese); Nichols and Murphy, Brooklyn Mus. Sci. Bull., j (i), 1916: 19 (occur, near N. York,
size, food, frequency, season, ident. by loc, but ill. on p. ig apparently of difland); Brooklyn Mus.
Quart., 5 (4), 1916: 151 (fig.), 153 (general, ident. by ill.); Thome, Copeia, 35, 1916: 69 (ident.
probable because of loc. Long Island, N. York, small size); Latham, Copeia, 43, 1917: 37 (Long
Island, N. York, ident. probable because of loc); Fowler, Proc. Acad. nat. Sci. Philad., dp, 1917: 109
(Atlantic City, N. Jersey, ident. probable because of loc); Proc. Boston Soc. nat. Hist., 55, 1917: iio
(Nantucket, ident. probable because of loc); Nichols and Mowbray, Copeia, 48, 1917: 78 (at least
in part because of loc, "tropical seas"); Latham, Copeia, 57, 1918: 53 (Long Island, N. York, ident.
probable because of loc); Coles, Copeia, 69, 1919: 41 (C. Lookout, N. Carolina, food, meas. of
adult, ident. by photo of head in fig. 3, but fig. 2 is of tudes^ see p. 435); Fowler, Proc biol. Soc.
Wash., a, 1920: 144 (N. Jersey records, ident. because of loc.) ; Latham, Copeia, 87, 1920: gi (Long
Island, N. York, November, ident. probable because of loc.) ; Fowler, Proc. Acad. nat. Sci. Philad.,
•J4, 1922: 3, 5, 7 (abund., size, season, N. Jersey and Delaware, ident. probable because of loc.) ; Nichols,
Copeia, 116, ig23: 52 (Long Island, N. York, ident. probable because of loc); Breder, Copeia, 127,
ig24: 27 (seasonal occur, near N. York, relation to temperature, ident. probable because of loc.) ; Wilson,
Proc. U.S. nat. Mus., 64 (17), 1924: 8, 12 (parasites, Woods Hole, ident. probable because of loc);
Barnard, Ann. S. Afr. Mus., 27 (l), 1925: 32 (S. Afr., probably in part, but descr. not diagnostic);
Nichols and Breder, Zoologica, N. Y., p, ig27: 17 (occur., season, size, south. New Engl., ident. probable
because of loc); Chcvey, Cons, explor. Mcr. Fauna Medit., 1927: plate not numbered (Medit., ident.
by ill.); Hildebrand and Schroeder, Bull. U.S. Bur. Fish., 43, 1928: 50 (ident. probable by ill. from
Garman, 1913, Chesapeake Bay); Rey, Fauna Iberica Feces, /, ig28: 364 (Spain, Portugal, ident. by
ill.); Breder, Field Bk. Mar. Fish. Atlant. Coast, ig2g: 19 (general, ident. by ill.); Truitt, Bean and

44^ Memoir Sears Foundation for Marine Research

Fowler, Bull. Md. Conserv. Dep., 5, 1929: 28 (young abundant, coast of Maryland and Chesapeake
Bay, summer, ident. probable because of loc.) ; Jordan, Manual Vert. Anim. NE. U.S., 1929: 11
(general, at least in part) ; Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 18
(in check list); Chevey, Faune Ichthyol. Cons, explor. Mer., 1930: plate not numbered (distrib. E.
Atlantic, ident. by ill.); Townsend, Bull. N. Y. zool. Soc., 34 (6), 1931: 167 (Virgin Is., ident. by
photo); Wilson, Bull. U.S. nat. Mus., 158, 1932: 423, 439, 440, 454, 456 (parasites, Woods Hole,
ident. probable because of loc); Breder, Copeia, 1932; 31 (abundance off Block I., ident. probable
because of loc); Gowanloch, Bull. La. Conserv. Dep., 25, 1933: 219, lower ill. (ident. probable from
ill., listed for Louisiana); Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda, 1933: 29 (ident.
probable, at least in part, because of loc, but descr. not diagnostic and ill. not original, Bermuda) ;
Vladykov, Proc. N. S. Inst. Sci., rp, 1935: 8 (ident. probable because of loc, Halifax, Nova Scotia) ;
Burton, Sci. Mon. N. Y., 40, 1935: 283 (occur, near Charleston, S. Carolina, but perhaps also
incl. other species as well); Vladykov and McKenzie, Proc. N. S. Inst. Sci., ig, 1935: 45 (Halifax
Harbor, Nova Scotia, same specimen as Vladykov, 1935); Marchand, Mar. biol. Rep. Cape Town, 2,
1935: 42 (S. Afr., ident. by photo) ; Fowler, Bull. Amer. Mus. nat. Hist., 70 (i), 1936: 64 (W. Afr.,
but may also cover dip/ana, for ill. after Carman's of Mass. spec, and descr., not diagnostic) ; Lubbert
and Ehrenbaum, Handb. Seefisch. Nordeurop., 2, 1936: 276 (occur., N. Europ., ident. by ill.);
Sigalas and Budker, Bull. Soc. Sci. Arcachon, 34, 1937: 59 (Arcachon, north. France, newborn,
ident. probable because of loc); Wise, Tigers of the Sea, 1937: 179 (descr., may include other
species) ; Norman and Eraser, Giant Fishes, 1937: 41, fig. 21 (general, may also include other species) ;
Breder, Bull. N. Y. zool. Soc, 41, 1938: 28 (N. York Harbor, ident. probable because of loc); Tor-
tonese, Atti Soc. ital. Sci. nat., 77, 1938: 302 (occur. Medit., may cover other species also); Springer,
Proc Fla. Acad. Sci., 5, 1939: 31 (in part, combined with diflana, Florida, see p. 420) ; Fowler, Bull.
U.S. nat. Mus., 100 {13), 1941: 217 (extensive synonymy also incl. diplana) ; Springer, Proc. Fla. Acad.
Sci., 5, 1941: 48 (comp. with other species, skull, N. York, New England, N. Jersey, Italian, Azores
specs.); Springer, Stanford Ichthyol. Bull., 1 (5), 1941: 168, fig. 5 (ill., comp. with other species);
Gunter, Amer. Midi. Nat., 28, 1942: 316 (tidal fresh water in Maryland, ident. probable because of
loc); Fowler, Arqu. Zool. Estado Sao Paulo, 5, 1942: 129 (Brazil); Bigelow and Schroeder, Guide
Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 134, fig. 49 (descr., habits, range, ill.),
Boos, Contr. biol. Lab. Cath. Univ. Amer., 45, 1943: 10 (chemistry of pancreas, Florida); Fowler,
Monogr. Acad. nat. Sci. Philad., 7, 1945: 97 (Beaufort, N. Carolina), 160 (Charleston, S. Carolina).

Zygaena zygaena Cuvier, Regne Anim., 2, 1817: 127 (general, ident. by refs.) ; Seabra, Bull. Soc. portug.
Sci. nat., 5, 191 1 : 192 (ident. by refs., Portugal); Noronha and Sarmento, Peixes Madeira, 1934: 107
(by ref., not seen).

Zygaena Tnalleus Risso, Hist. nat. Europ. Merid., 5, 1826: 125 (in part, but also incl. difla-rm by ref. to
malleus Valenciennes, 1822, general, Medit.); Voigt, in Cuvier, Tierreich, 2, 1832: 513 (at least in
part, ident. by refs.) ; Bory de St. Vincent, Diet. Class. Hist. Nat., I 5, 1829: 597 (at least in part, ident.
by refs.); Jenyns, Manual Brit. Vert. Anim., 1835: 507 (Yarmouth, England, ident. probable because
of loc); Lowe, Trans, zool. Soc. Lond., 2 (3), 1837: 195 (Azores, ident. because of subsequent ill.,
see Lowe, 1 843-1 860); Yarrell, Brit. Fish., 2, 1836: 406 (Yarmouth, England, ident. by ill.); also
2nd ed., 2, 1841 : 504 (Brit, records, number of young, ident. as above) ; Couch, Cornish Fauna, 1838:
51 (Cornwall); DeKay, Zool. N. Y., 4, 1842: 362, pi. 62, fig. 204 (near N. York, ident. by ill.);
Storer, Boston J. nat. Hist., 4, 1842: 185 (near Woods Hole, ident. by ill.); Lowe, Fish. Madeira, /,
1843— 1860: 83, pi. 12 (Madeira, ident. by descr. and ill.); Hamilton, Naturalist Libr., Ichthyol., 6;
Brit. Fish., 2, 1843: 306, pi. 24, fig. I, also subsequent eds. (Eng. loc, ident. by ill.) ; Linsley, .Amer. J
Sci., 4y, 1844: 77 (Long Island Sound, N. York, ident. probable because of loc.) ; Storer, Mem. Amer.
Acad. Arts Sci., N. S. 2, 1846: 508 (Massachusetts, N. York, Caribbean, Brazil; in part, ident. by loc, and
refs., diflana also incl. by ref. to Valenciennes, 1822) ; DeKay, Rep. St. Cab. nat. Hist. N. Y., 8, 1855:
64 (listed for N. York, ident. by a ref. to DeKay, 1842) ; Storer, Mem. Amer. Acad. Arts Sci., N. S.
9, 1867: 238, pi. 38, fig. 3; also Sep., Fishes Mass., 1867: 262, pi. 38, fig. 3 (occur, south. Mass.ichusetts,
ident. by ill.); Gunther, Cat. Fish. Brit. Mus., 8, 1870: 381, 518 (descr., distrib., refs., but also covers
diflana, see p. 419) ; Gervais and Boulart, Poiss., j, 1 877: 1 74, pi. 66 (Medit. and Adant., ident. by ill.) ;

Fishes of the Western North Atlantic 447

Doderlein, Man. Ittiol. Medit., 2, 1881: 46 (occur, in Medit., ident. by refs., but probably covers
iiflana also);*' Moreau, Poiss. France, r, 1881: 324 (occur. French coasts; ident. by ill.); Day,'Fish.
Gt. Brit., 2, 1880-1884: 294 (occur. Gt. Brit., ident. by local, but ill., pi. 154, is diflana); Vieira,
Ann. Sci. nat. Porto, 4, 1897: 66 (Portugal, ident. by inference from included refs.); Lloyd and
Sheppard, Proc. zool. Soc. Lond., 1922: 971 (Wales, neuro-anatomy, skull, ident. by ill. of skull);
Jenkins, Fish. Brit. Isles, 1925: 308 (ident. probable because of loc.) ; Joubin and Le Danois, Mem.
Peches Marit. France, 2, 1925: 27 (coast of France, Tunis, ident. at least in part from refs., but may
also cover diflana) ; Ehrcnli.ium, in Grimpe and Wagler, Tierwelt N.- u. Ostsee, Ijef 7 (12"), 1927: 13
(North Sea, ident. by loc); Lloyd and Sheppard, Zool. Anz., 80, 1929: 65 (skelet., same spec, as
Lloyd and Sheppard, 1922) ; Belloc. Rev. des Trav. Peches Marit., 7, Fasc. 2, 1934: 133 (ill. of head;
Morocco; trop. W. Afr. in general; C. Verde Is., Canaries); Noronha and Sarmento, Peixes Madeira,
1934: 107 (not seen).
Zygaev-a fu! garis C\oc\\i£t, Diet. Sci. Nat., 60, 1 8 30: 621 (at least in part, general).
Zygaejta (no spec, name) Storer, Rep. Fish. Rept. Birds Mass., 1839: 200 (south. Massachusetts, ident.

probable because of loc).
Zygaena nib'jrcnata Storer, Proc. Boston Soc. nat. Hist., 5, 1848: 70 (ident. by descr. of 2-ft. spec. Province-
town, C. Cod).
S-phymias zygaena Gray, List Fish. Brit. Mus., 1851: 48 (in part; ident. by refs., but also incl. diflana,
see p. 420); White, List. Spec. Brit. Mus., Fish., 8, 1851: 126 (part; ident. by refs., but also incl.
diflana, see p. 420).
Sfhyrna malleus Van der Hoeven, Handb. Dierkunde, 2nd ed., 2, 1855: 262 (general, Medit., ident. by

refs. to Linnaeus) ; Handb. Zool., Eng. transl., 2, 1858: 68.
Cestracion subarcuata Gill, Proc. Acad. nat. Sci. Philad., Addend., 1 861: 59 (name only, evidently based on
Zygaena subarcuata Storer, 1848) ; Abbott, Bull. geol. Surv. N. J., Geol. N. J., Append. E, 1868: 829
(occasional N. Jersey, ident. probable because of loc).
Cestracion zygaena Gill, Ann. N. Y. Lye, 7, 1862: 403 (name only, ident. by ref. to Linnaeus, 1758);
Proc. Acad. nat. Sci. Philad., 1863: 333 (Massachusetts, ident. by refs.); Dumeril, Hist. Nat. Poiss.,
J, 1865: 382 (general, ident. by refs., but also incl. diflana by ref. to malleus Valenciennes, 1822);
Fowler, Rep. N. J. Mus. (1905), 1906: 65 (occur. N. Jersey, ident. probable because of loc) ; Fowler,
Rep. N. J. Mus. (1906), 1907: 257, pi. 76 (N. Jersey record, ident. by ill.); Garman, Mem. Harv.
Mus. comp Zool., 56, 1913: 157, pi. l, fig. 1-3 (in part, N. York spec, in Harv. Mus. Comp. Zool.,
ident. from ill., and by personal examination; but refs. also include diflana, see p. 420); Radcliffe,
Bull. U.S. Bur. Fish., 5^, 1916: 263 (in part), pi. 43, fig. 3 (N. Carolina; ident. by ill., but pi. 43, fig. I,
clearly diflana [see p. 420I ; fig. 2, tiburo; fig. 4 probably tudes because of teeth described as serrate [see
p. 43;]); Linton, Proc. U.S. nat. Mus., 64 (21), I924: 5, 7, 21, 31, 37, 38, 40, 41, 42, 44, 96
(parasites. Woods Hole, ident. probable because of loc.) ; Bigelow and Welsh, Bull. U.S. Bur. Fish., 40
(l), 1925: 31 (Gulf of Maine records); Bigelow and Schroeder, Bull. U.S. Bur. Fish., 48, 1936: 322
(Halifax, Nova Scotia, record, from Vladykov, 1935 ; also between Browns and Georges Banks, ident. by
loc); Chabanaud and Monod, Bull. Etud. Hist. Sci. Afric Occid. Franc, 1926: 229 (at least in part,
Morocco; not seen); Uriarte and Mateu, Notas Inst. esp. Oceanogr., (2) 53, 1931: 51 (Canaries,
Teneriffe, n.ime only, but ident. probable at least in part); Cadenat, Rev. des Trav. Peches Marit.,
70 (4), 1937: 430 (Dakar, Canakry, W. Africa).
Hammerhead, Couch, Hist. Brit. Fish., /, 1867: 70, pi. 16 (Gt. Brit., ident. by ill.).
Sfhyrna {Cestracion) zygaena von Bonde, J. comp. Neurol., 58, 1933: 377, pi. i, 3> 4 (S. Afr., skull and

nerves, ident. by photo of head).
Zygaena zigaena Nobre, Fauna Marinha Port., Vert., i, 1935: 425 (Portugal, ident. by descr. of head).

47. For additional Mediterranean records in publications not accessible to us, see Doderlein, 1881.

448 Memoir Sears Foundation for Marine Research

Atlantic, by Name Only:'"

Squalus zygaena Briinnich, Ichthyol. Massil., 1768: 4 (Medlt.) ; Forskal, Descr. Anim., 1775: xviii (no
local.); Bowditch, Excurs. Madeira, 1825: 75 (Porto Santo); Nardo, Prod. Ittiol. Adriat., i, 1826: 9
(Adriatic).

Zygene marteau, Cloquet, Diet. Sci. Nat., Atlas. Poiss., 1816-1830: pi. 32 (not diagnostic).

Zygaena malleus Busch, Selach. Ganoid. Encephal., 1848: 22, pi. 2, fig. 5 (brain, no loc.) ; Hasse, Naturl.
Syst. Elasm. besond. Theil., 1882: 273, pi. 39, fig. 21—25 (denticles, vertebrae); Rochebrune, Acte.
Soc. linn. Bordeaux, (4) 6, 1882: 43; Faune Senegambie, Poiss., /, 1883-1885: 20 (Senegambia) ;
Graeffe, Arb. zool. Inst. Univ. Wien, 28, 1886: 446 (Medit.) ; Vinciguerra, Atti Soc. ital. Sci. nat., $4,
1892: 300 (Teneriffe, name only) ; Sicher, Atti Accad. gioenia, (4) 11 (5), l8g8: 15 (Medit.) ; Wilson,
Proc. U.S. nat. Mus., 5^, 1 907: 396 (Cape Verde Is., name only) ; Brian, Bull. Inst, oceanogr. Monaco,
no, 1908: 3 (C. Verde, Azores, name only); Metzelaar, Trop. Atlant. Visschen, 1919: 189 (W. Afr.,
name only).

Sphyrna zygaena Nardo, Sinon. Modern. Spec, descr. Pesci St. Chiereghin, 1847: ill (name only);
Machado, Peces Cadiz., 1 857: 9 (name only); Canestrini, Arch. Zool. Anat. Fisiol., j, 1 861: 266
(Genoa, name only); Bocage and Brito Capello, Peixes Plagiost. Portugal, i, 1866: 17 (Portugal, name
only) ; Ninni, Ann. Soc. Hist. nat. Modena, 5, 1870: 66 (Venice, name only) ; Miklucho-Maclay, Beitr.
Vergl. Neurol. Wirbelt., 1870: 25, pi. 5, fig. 2 (brain) ; Goode, Bull. U.S. nat. Mus., 5, 1876: 73 (Ber-
muda, name only) ; Goode, Amer. J. Sci., 14, 1877: 293 (Bermuda, name only) ; Proc. U.S. nat. Mus., 2,
1879: 121 (Florida, name only); Yarrow, Proc. Acad. nat. Sci. Philad., 29, 1877: 217 (N. Carolina);
Bean, T. H., Proc. U.S. nat. Mus., 5, 1881: 115 (East Coast, U.S.) ; Jordan and Gilbert, Proc. U.S. nat.
Mus., 5, 1882: 581 (name only); von Ihering, Rev. Mus. pauL, 2, 1897: 34 (Muldonado, Monte-
video, name only); Hilgendorf, Arch. Naturgesch., 54 (l), 1888: 213 (Azores, name only);
Carus, Prod. Fauna Medit., 2, 1889-93: 513 (Medit., refs. probably include both species); Hen-
shall, Bull. U.S. Fish Comm., 9, 1891: 384 (Florida, name only); Smith, Rep. U.S. Comm.
Fish. (1895), 1896: 175 (Florida, name only); Jordan and Rutter, Proc. Acad nat. Sci. Philad.,
1897: 91 (Jamaica, name only); Berg, Commun. Ittiol. Mus. nac. B. Aires, i (l), 1898: 9 (Mon-
tevideo, not diagnostic); Evermann and Bean, Rep. U.S. Comm. Fish. (1896), 1898: 239 (Flor-
ida, name only); Gilbert, Proc. Wash. Acad. Sci., 2, 1900: 161 (Brazil, name only); Gregg, Where
to Catch Fish, 1902: 16 (not diagnostic); Evermann and Marsh, Bull. U.S. Fish Comm., 20 (l),
1902: 63 (Porto Rico, name only); Schreiner and Ribeiro, Arch. Mus. nac. Rio de J., 72, 1903: 79
(Pernambuco, name only) ; Hargreaves, Fish. Brit. Guiana, 1904: 14, and Append., 8 (Brit. Guiana, at-
tacks on man, name only) ; Devincenzi, An. Mus. Hist. nat. Montevideo, (2) i, 1 904: 120 (Rio de La
Plata, name only) ; Jordan and Thompson, Bull. U.S. Bur. Fish., 24, 1905: 232 (Tortugas, Florida, name
only) ; Bean, Field Mus. Publ. Zool., 7 (2), 1906: 30 (Bermuda, name only) ; Wilson, Proc. U.S. nat.
Mus., J J, 1907: 416, 431, 626 (C. Verde Is., N. Carolina, parasites, name only) ; Gudger, Science, N. S.
2$, 1907: 1005 (N. Carolina, I2j^-ft. spec, food, name only); Fowler, Prcc. Acad. nat. Sci. Philad.,
61, 1909: 407 (C. May, N. Jersey, name only) ; Rep. N. J. Mus. (1908-1910), 1910: 352 (N. Jersey
record, name only) ; Proc. Acad. nat. Sci. Philad., 63, 191 1 : 5 (Delaware, name only) ; Roule, Bull. Inst.
oceanogr. Monaco, 243, 1912: 10 (Medit., name only); Fowler, Copeia, 2, 1913: 2 (Maryland, name
only) ; Pietschmann, Jb. nassau. Ver. Naturh., 66, 1913: 176 (Cameroons, name only) ; Fowler, Copeia,
13, 1 91 4: 2 (N. Jersey, name only) ; Proc. Acad. nat. Sci. Philad., 6y, 191 5: 245, 521 (Florida, Trinidad,
names only); Copeia, 31, 1916: 41 (N. Jersey, name only); Copeia, 27, 1916: 11 (N. Jersey, name
only); Copeia, 30, 1916: 36 (name only); Proc. Acad. nat. Sci. Philad., dp, 191 7: 127 (Colon, name
only); Copeia, 43, 1917; 39 (Florida, name only); Nichols, Bull. Amer. Mus. nat. Hist., jy, 1917:
875 (Florida, name only); Ribeiro, Rev. Mus. paul., 10, 1918: 708 (Santos, Brazil, name only);
Fowler, Proc. Acad. nat. Sci. Philad., yi, 1919: 128, 129, 146, 293 (Panama, Surinam, Jamaica, N.
Jersey, name only); Nichols, Bull. Amer. Mus nat. Hist., 44, 1921: 22 (Turks I., name only);
Lahille, Physis, B. Aires, 5, 1 921: 63 (Argentina, name only); Enumer. Peces Cartilag. Argent.,
1921: 14 (Argentina, name only); Fowler, Proc. Acad. nat. Sci. Philad., y2, 1 921: 386 (N. Jersey,

48. References ostensibly to Sfhyrna zygaena, but by name only, either without locality, or for localities where
diplana may also be expected to occur, hence which are as likely to have been based on the latter as on the former,
or on the two combined.

Fishes of the Western North Atlantic 44.9

name only); Ribeiro, Arch. Mus. nac. Riode J., Fauna brasil. Peixcs, 2(1) Fasc. 1, 1923: 14 (Brazil north
and south, dcscr. not diagnostic) ; Fowler, Copeia, 143, 1 92 5: 41 (Delaware, name only), 43 (N. Jersey,
name only) ; Proc. Acad. nat. Sci. Philad., 78, 1926: 249 (Florida, food, embryos, name only) ; Copeia,
156, 1926: 146 (N. Jersey, name only); Copeia, 165, 1927: 90 (Delaware B.iy, name only); Bcebe
and Tee-Van, Zoologica, N. Y., 10, 1928: 29 (Haiti, name only) ; Nichols, Sci. Surv. Porto Rico, N. Y.
Acad. Sci., 10 (2), 1929: 184 (Porto Rico, not diagnostic) ; Fowler, Proc. Acad. nat. Sci. Philad., 80,
1929:608 (N.Jersey records, names only ■) ; MacCallum, Proc. U.S. nat. Mus., 79 (26), 1931 :6 (par.)sitcs,
name only); Fowler, Proc. Acad. nat. Sci. Philad., 83, 1931: 391 (Trinidad, name only); Breder, Pap.
Tortugas Lab., 2,?, 1932: 16 (Tortugas, name only) ; Pearson, Invest. Rep., U.S. Bur. Fish., (10) /, 1932:
18 (listed, N.Carolina) ; Fowler, Proc. biol. Soc. Wash., 46, 1933: 57 (Louisiana, name only) ; Young and
Mazct, "Shark, Shark," 1933: 268 (not diagnostic); Brooks, Parasitology, 26, 1934: 265 (N. Carolina,
parasites); Pozzi and Bordnle, An. Soc. cient. argent., r2o, 193;: i;i (N. Argentina, name only);
Fowler, Bull. Amer. Mus. nat. Hist., 70 (2), 1936: 1 152 (Azores, name only); Smith, H. W., Biol. Rev.,
ir, 1936: 64 (Louisiana, name only); Fowler, Proc. .Acad. nat. Sci. Philad., 8^, 1937: 304 (7; miles
off N. Jersey coast, name only) ; Longley and Hlldebrand, Pap. Tortugas Lab., 34, 1 941: 3 (Tortugas,
Florida) ; Lunz, Bull. S. C. St. Planning Bd., 14, 1944: 27 (S. Carolina, Florida).

Sfhyrna leunrri Dumeril, Rep. Poiss. Afr. Occid., 1861: 261 (Gorree, probably not Zygaena lezc-ini Griffith,
Cuvier, Regne Anim., 10, 1834: 640, pi. 50, Aust.).

Cestracion (Sf/iyrna) zygaena Steindachner, S. B. Akad. Wiss. Wien, 61 (i), 1870: 576 (Senegal, name only) ;
Denkschr. Akad. Wiss. Wien, ^^, 1882:51 (Senegambia, name only).

Cestracion zygaena Poey, .^n. Soc. esp. Hist. Nat., 5, 1876: 382; F.numerat. Pise. Cubens., 1876: 187 (Cuba) ;
Fowler, Proc. Acad. nat. Sci. Philad., 58, 1906: 80 (Florida, name only); Radcliffe, Trans. Amer.
Fish. Soc, 44, 1914: 38 (N. Carolina, name only); Roule, Result. Camp. sci. Monaco, 52, 1919: 117
(Morocco, Azores, name only); Monod, Faune Colon. Fr.inc., 1927: 647 (Lokundje, not seen);
Nigrelli, Amer. Mus. Novit., 996, 1938: 10 (parasites, name only).

Zygaena vu/garis Reguh, Ess. Hist. nat. Provence, i (l), 1877: 58 (Medit.).

Cestracy on zygaena Poey, An. Soc. esp. Hist, nat., 70, 1 881: 348 (Porto Rico, Guiana, Medit., E. Indies, name
only); Stahl, Fauna Puerto Rico, 1883:81, 167 (Porto Rico, not seen).

Sfhyrna {Zygaena) zygaena Imms, Proc. zool. Soc. Lond., /, I 905: 43 (pharyngeal denticles, name only).

Hammerhead, Gudger, Amer. Mus. nat. Hist., .^o, 1937:417-418 (Palm Beach, Florida, attack on girl).

Sfhyrnasp. {-prohahly zygaena) Norris, Plagiost. Hypophysis, 1 941: pi. i, fig. 3 (brain).

Suborder SQUALOIDEA

Characters. No anal fin; 2 dorsal fins, with or without spines; only 5 gill openings, all
anterior to pectorals; snout not beak-like, without lateral teeth or cirri; teeth in front of
mouth essentially similar to those toward corners; general form subcylindrical (shark-
like) ; eyes lateral; anterior margins of pectorals not expanded forward past ist gill open-
ing; inner margins of pelvics entirely separate, posterior to cloaca; nostrils entirely sepa-
rate from mouth; spiracles present; eyes without nictitating fold or membrane; vertebral
column completely segmented throughout its length, its axial canal much contracted in the
regions of the well differentiated centra, the notochord greatly constricted segmentally, or
even obliterated in the centra, but dilated in the spaces between the concave surfaces of
adjoining vertebrae; vertebral centra with calcareous lamellae in a ring around central
axis; neural spines not attached to dorsals; cranium with antorbital processes more or less
developed, but without separate antorbital bar; upper jaw (palatoquadrate cartilage)
attached to cranium by a transverse process at one point only, in the ethmoid region, as
well as to hyomandibular arch; rostral cartilage single; propterygial cartilage of pectoral

450 Memoir Sears Foundation for Marine Research

with I to several radial elements} pelvics transverse; heart valves in 2 to 4 rows. Develop-
ment usually ovoviviparous, but probably oviparous in some cases.

Key to Families

la. Each dorsal fin preceded by a long or short spine with tip exposed or concealed.^

Squalidae, p. 450.
lb. Second dorsal fin, and usually the ist, without a spine.^

2a. Teeth with only i cuspj uppers and lowers unlike, the former narrow, raptorial,
the latter expanded widely laterally as a cutting edge (sectorial).

Dalatiidae, p. 499.
2b. Teeth with several cusps, uppers and lowers similar, sectorial.

Echinorhinidae, p. 526.

Family SQUALIDAE

Characters. Squaloidea with a spine in each dorsal fin, long in some cases but so short
in others as to be easily overlooked; teeth with i or several cusps, alike or unlike in the 2
jaws. Characters otherwise those of the suborder.

Genera. Generic distribution of the various members of this family is still in some
confusion owing to the fact that all the characters that have been regarded as generically
diagnostic by one author or another are intergrading, not strictly alternative. Conse-
quently, the accompanying Key is necessarily tentative.

Tentative Key to Genera

I a. Fin spines originating about at midpoint of bases of dorsal fins and running forward
to emerge from anterior margins of latter; trunk very stout, sub triangular, with longi-
tudinal dermal ridges anterior to pelvic fins. Oxynotus^ Rafinesque, 1 8 lO.

Eastern Atlantic, Mediterranean,
Australian region.
lb. Fin spines originating at origins of dorsal fins and lying along anterior margins of
latter; trunk slender, subcylindrical, with dermal longitudinal ridges (if any) con-
fined to sector posterior to pelvic fins.
2a. Upper teeth with several cusps.
3a. Teeth similar in the 2 jaws.

Centroscyllium Miiller and Henle, 1841, p. 480.
3b. Upper and lower teeth noticeably unlike, the lower with only i cusp.

Etmofterus Rafinesque, 18 10, p. 487.

1. Even in the genera in which the spines are shortest (e.g., Centroscymnus, p. 493) they are easily detected by
touch.

2. In Euprotomicrus the first dorsal may or may not have a spine.

3. Classed as a separate family (Oxynotidae) by some authors.

Fishes of the Western North Atlantic 451

2b. Upper teeth with only i cusp.

4a. Snout in front of mouth considerably longer than from center of mouth to
origin of pectorals j dermal denticles pitchfork-shaped, on tall slender pedi-
cels. Deania* Jordan and Snyder, 1902."

Eastern Atlantic, South Africa, Japan,
Philippines, Australia, New Zealand.
4b. Snout in front of mouth considerably shorter than from center of mouth to
origin of pectorals j dermal denticles at most only moderately dentate, on
short broad pedicels, or sessile.
5a. Teeth similar in the 2 jaws.

6a. Anterior margin of nostril without long barbel.

Squalus Linnaeus, 1758, p. 452.

6b. Anterior margin of nostril with a barbel reaching past corner of

mouth. Cirrhigaleus Ta-ndk-di^ 19 12.'

Japan.

5b. Teeth noticeably dissimilar in the 2 jaws.

7a. Inner corner of pectoral broadly rounded.

8a. Blades of dermal denticles on trunk behind ist dorsal smooth,
with rounded margins} ridged or striate denticles confined to
more anterior part of body.

Centroscymnus Bocage and Brito Capello, 1 864, p. 493.
8b. Blades of dermal denticles with 3 to several ridges 5 with mar-
ginal teeth on posterior as well as on anterior parts of trunk.

Scymnodon Bocage and Brito Capello, 1864.'

Eastern Atlantic, Straits of Magellan, Japan, New Zea-
land, Philippines, India.

7b. Inner corner of pectoral angular and more or less produced.

Centrofhorus Miiller and Henle, 1837.'

Eastern Atlantic, Mediterranean, South Africa,
Japan, New Zealand, Australia.

4. We agree with Fowler (Bull. U.S. nat. Mus., loo [z^], 1941 : 237) that this name should take precedence over
A canth'idium as used by Garman (Mem. Harv. Mus. comp. Zool., jiS, 1913 : 215), which is properly a synonym of
Etmofterns (see also footnote i, p. 487).

5. Including Deaniofs Whitley, 1932. 6. Including P/taenofogon Herre, 1935.

7. Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 206) has proposed a separate genus, Centroselachus, for spe-
cies in which the denticles have numerous ridges, as contrasted with those in which they have only three. But this
does not seem to us a sufficiently important difference to be regarded as generic.

8. Including Lefidorhinus Bonaparte, 1838, and Entoxychirus Gill, 1862. Fowler (Bull. U.S. nat. Mus., 100 [z^],
1941 : 223) redivides this group of species between Centrofhorus and En'z.xych'irus. But the differences on which
this division is based, i.e., the relative degrees to which the mner corners of the pectorals are produced and the
shapes of the dermal denticles, do not seem to us sufficient for generic separation. We may note also that three of the
species included by him in Centrofhorus {rossi Alcock, 1898; waitei Thompson, 1930; and foliaceus Giinther,
1877) appear to us to belong to Scymnodon. The genus Lefidorhinus of Bonaparte (Nuov. Ann. Sci. nat.
Bologna, 2, 1838: 207) has also been revived by Garman (Mem. Harv. Mus. comp. Zool., j<5, 1913: 211). But
its type species, the European Squalui squamoius of Bonnaterre (Tabl. Encyc. Meth. Ichthyol., 1788: 12), falls in
Centrofhorus as defined here, the inner corners of its pectorals t>eing angular and at least slightly produced.

452 Memoir Sears Foundation for Marine Research

Genus Squalus Linnaeus, 1758'

Squalus Linnaeus, Syst. Nat., /, 1758: 233; type species, S. acanthias Linnaeus, European Ocean; designated
by Gill, Ann. N. Y. Lye, 7, 1862: 405.

Generic Synonyms:

Squaltus Scopoli, Intrcd. Nat. Hist., 1777: 464; type species, Squalus acanthias Linnaeus, 1758 (emend, spell-
ing)-

Acanthorhinus Blainville, Bull. See. philom. Paris, 1 8 16: 121 ; type species, Squalus acanthias Linnaeus, 1758,
designated by Jordan, Genera Fish., I, 191 7: 95.

Sfinax (in part) Cuvier, Regne Anim., 2, 1817: 129.

Acanthias Risso, Hist. Nat. Europe Merid., ^, 1826: 131 ; type species, A. vulgaris Risso, equals Squalus acan-
thias Umnaeus, 1758.

Carcharias Gistel, Naturg. Thier., 1848: 8; to replace Acanthias Risso, 1826: not Carcharias Rafinesque, 1810.

Flakeus Whitley, Aust. Zool., 9, 1939: 242; type species, Squalus megalop Macleay, 1881.'"'

Koinga'Whhlzy, Aust. Zool., p, 1939: 242; type species, Squalus griffini Phillipps, 1931.'°

Generic Characters. Squalidae with well developed dorsal spines, without lateral
grooves, originating at origins of fins and lying along anterior margins of latter, at least
their terminal V3 to V4 free} trunk slender, rounded, with longitudinal ridges confined
to caudal peduncle j caudal peduncle with a precaudal pit above but none below; a labial
furrow on each jaw and a voluminous pit at corner of mouth ; upper and lower teeth alike,
with I cusp, deeply notched outwardly, and so oblique that their inner margins form a
nearly continuous cutting edge; snout in front of mouth considerably shorter than from
center of mouth to origin of pectorals; dermal denticles very small, lanceolate, heart-
shaped or tridentate, with sharp tip, but varying in shape on different parts of body; eyes
and spiracles large; nostrils far from mouth, without barbels; both dorsals with concave
posterior margins and elongate free rear corners, the ist triangular, the 2nd considerably
smaller than ist, the origin of ist dorsal over inner margin of pectoral or a little posterior
to inner corner of latter, the origin of 2nd dorsal behind bases of pelvics; caudal without
subterminal notch, its lower anterior corner expanded as a definite lobe, but much shorter
than the upper lobe and smaller in area; luminous organs lacking. Characters otherwise
those of the family.

Range. Widely distributed in the North Atlantic, west and east, including the Medi-
terranean; western South Atlantic (Uruguay and Argentina) ; South Africa; both sides of
the Pacific, from southern Alaska, the Aleutians, Kamchatka, Japan, Korea and China in
the north to New Zealand, southern Australia, Tasmania and Chile in the south, including
the Hawaiian Islands and Philippines; also southern Indian Ocean (Mauritius) ; in tropi-
cal to subarctic and subantarctic latitudes.

Fossil Teeth. Upper Cretaceous, western Asia; Upper Cretaceous to Pliocene, Eu-
rope; Eocene, South Africa; Oligocene, South America; Miocene, North America.

Sfecies. The named representatives of Squalus fall into three groups, as defined by
the relative locations of the fins and by the shape of the anterior margin of the nostril.

9. Fowler (Bull. U.S. nat. Mus., 100 [15], 1941 : 255) includes in the synonymy of Squalus the fossil genus Cen-
trofhorides Davis (Trans, roy. See. Dublin, [2] 5, 1887: 478).

10. Whitley's proposed subdivision of the genus Squalus into two subgenera (^Flakeus and Koinga) according to the
position of the first dorsal fin, size of eye and coloration, does not appear to us acceptable.

Fishes of the Western North Atlantic 453

A. The acanthias group: ist dorsal spine over or behind the inner corner of the pec-
toral; midpoint of bases of pelvics much closer to 2nd dorsal than to ist; distal margin of
pectoral at least moderately concave; anterior margin of nostril simple (not bilobed);
at least most of the members of the acanthias group are white-spotted, while the others
are not.

B. The blainville-jernandinus group: ist dorsal spine almost over midpoint of
inner margin of pectoral ; midpoint of bases of pelvics about midway between the two dor-
sal fins; inner margin of pectoral nearly straight; anterior margin of nostril bilobed (Fig.
87E,F).

C. The brevirostris-cubensis group resembles the blainville-jernandinus group in
relative position of fins and in bilobed nasal margin, but is set apart by the distal margin of
the pectoral, which is deeply concave with its inner angle noticeably acute.

Group A is represented in the North Atlantic by the familiar Spiny or Piked Dog-
fish {S. acanthias) of temperate and boreal latitudes. A close relative in the North Pacific
is usually regarded as specifically distinct {suckleyi Girard) but recently has been listed
as acanthias. ^^ According to recent Keys'" the chief alternative character supposedly sepa-
rating suckleyi from acanthias is the position of the first dorsal spine, opposite or a little
behind the inner corner of the pectoral in the former, behind and remote from it in
the latter. Actually, however, our Study Material shows that this criterion is not tenable;
not only is the variation considerable in this respect among both Atlantic and Pacific speci-
mens, but in some of the latter the first dorsal spine stands as far behind the pectoral as it
does in any of the Atlantic series.'' It is even doubtful whether there is any average dif-
ference between the two populations in this respect. Nor have we been able to find any
other difference to separate them, whether in position of fins, in proportionate dimensions,
or in teeth. In short, the North Pacific and North Atlantic populations of the acanthias
group have not differentiated themselves specifically during the period since their ranges
became discontinuous.

It is doubtful whether this group occurs in the equatorial Atlantic, unless accidentally.
But it is as widespread in the temperate and boreal belts of the southern as of the northern
hemisphere. Thus the Spiny Dogs of the Straits of Magellan," Australia and New Zea-
land,"* with those reported from Uruguay and northern Argentina as Squalus acanthias'''

11. Soldatov and Lindberg, Bull. Pacif. Fish. Res. Sta., 5, 1930: 16; eastern Asia.

12. Garman, Mem. Harv. Mus. comp. Zool., 56, 1913: 192; Fowler, Bull. U.S. nat. Mus., 100 (/j), 1941: 257.

13. Especially significant is the fact that among three embryos of a single brood from San Francisco the first dorsal
spine occupies the same position relative to the pectoral as is commonly true in Atlantic specimens; in another
one it is as usually stated for the Pacific form.

14. A. lebruiii Vaillant, Miss. Sci. Cape Horn (1882-83), 1885: 13, pi. i.

15. Fernandinus Waite (Rec. Canterbury [N. Z.] Mus., i, 1901: 14.2, pi. 16, fig. i) appears to be identical with
kirki Phillipps (N. Z. J. Sci. Tech., 12, 1931 : 361) from New Zealand, and with whitleyi Phillipps (N. Zealand
J. Sci. Tech., 12, 193 i : 361) from South Australia. It is true that the illustration (McCoy, Prod. Zool. Victoria,
1886: pi. 75, fig. lb) on which w/titUyi was based fails to show any labial furrows, but this was probably an
oversight, for McCoy not only includes labial furrows in the generic diagnosis but states that he %vas unable to
detect any difference between Australian and British specimens.

16. Berg, An. Mus. nac. B. .\ircs, (2) /, 1895: 6; Devincenzi, An. Mus. Hist. nat. Montevideo, (2) /, 1920: 123;
Lahille, Physis B. Aires, 5, 1921 : 63.

454 Memoir Sears Foundation for Marine Research

or as Acanthias vulgaris" all fall in the acanthtas group, so far as the fins are concerned,
but may perhaps be set apart by the teeth^^ when adult. A form at least very close to acan-
thias is also known under that name from South Africa and from the Island of Reunion in
the southern Indian Ocean. The relationship of these southern hemisphere forms to one
another and to the northern acanthias is uncertain.

In the North Atlantic group B is confined to the Mediterranean [hlainville Risso,
1826). But it is widely represented in the southern hemisphere (Mauritius, South Africa,
Argentina, Tasmania) and in Philippine, Japanese and Korean waters by forms so closely
resembling one another, that while they have formed the basis for at least seven supposed
species, most of these have recently been united by Fowler" under the oldest name, jernan-
^iM«j Molina, 1782."

Our own comparison of specimens of this group from the Mediterranean with others
from the west coast of South America, Juan Fernandez and Tasmania" confirms this union
in so far as concerns the fins, shape of head, and snout and margin of nostril. But the
dorsal fin spines (second as well as first) are considerably shorter in the eastern Pacific
and Tasmanian specimens, and also (by published accounts) in the South African"^ and
Argentine"*" representatives of this group than in the Mediterranean representative or in
the Japanese as pictured. Hence, since the length of the spines is fairly constant in the only
member of the genus {^acanthias') of which a large series has been examined, it seems wise
to retain the name hlainville for the Mediterranean form, at least for the present, post-
poning decision as to how many species the fernandinus group includes in the southern
hemisphere and in the Pacific until adequate series can be compared from representative
localities.

Group C includes four named forms: cubensis Howell-Rivero, 1936, from Cuba 5
brevirostris Tanaka, 19 17, from Japan; me galops Macleay, 1881, from southern Aus-
tralia and Tasmania;''^ and grifmi Phillipps, 1931, from New Zealand. Here again deci-
sion as to whether or not these are all distinct species, and if so by what alternative charac-

xj. Perugia, Ann. Mus. Stor. nat. Genova, (2) to (30), 1891: 608.

18. See Devincenzi (An. Mus. Hist. nat. Montevideo, [2] 4 [14]) '939: 4) for a recent discussion.

19. Bull. U.S. nat. Mus., 100 (/j), 1941 : 260-

20. Exceptions are tasman'tensis Rivero, 1936, which Fowler retains as distinct, and mitsukurii Jordan and Fowler,
1903, which he relegates to the synonymy of suckleyi Girard, 1854, i.e., the North Pacific representative of the
acanthias group. Actually, however, mitsukurii was a compound species, a fact which has resulted in much
confusion in the nomenclature of Japanese sharks of this genus. Thus the form pictured under that name by its
authors, Jordan and Fowler (Proc. U.S. nat. Mus., 26, 1903; 630, fig. 3), and subsequently by Tanaka (Fish.
Japan, 26, 1917: pi. 130, fig. 36S-370), was clearly acant/rias-Wke, but the specimen described by Jordan and
Fowler on the preceding page (which is therefore the type of the species) was of the fernandinus group, as
pointed out by Jordan and Hubbs (Mem. Carneg. Mus., 10, 1925: 105, 106). And this is also true of jafonicus
Ishikawa (Proc. Acad. nat. Sci. Philad., 1908: 71); Tanaka (Fish. Japan, 26, 1917: pi. 130, fig. 365-3*7)-
Jafonicus is therefore a synonym of mitsukurii, and the latter in turn probably a synonym of fernandinus.

21. The latter is the type specimen of tasmaniensis Howell-Rivero, 1936.
2ia. Acutipinnis Regan, Ann. Natal Mus., 7, 1908: 248, pi. 37.

2ib. Fernandinus Lahille, An. Mus. nac. B. Aires, }j, 19:9: 327, fig. 17.

22. For illustration of megalofs and griffini, see Whitley (Fish. Aust., /, 1941 : 138).

Fishes of the Western North Atlantic 455

ters they are separated, must await comparison of specimens from the three geographic
regions.

Because of these uncertainties the following Key is limited to the western Atlantic
representatives of the genus, with the Mediterranean and eastern Atlantic blainville in-
cluded to facilitate comparison.

Key to Western Atlantic Species

I a. First dorsal spine over or posterior to inner corner of pectoral; midpoint of bases of
pelvics much nearer to origin of 2nd dorsal than to rear end of base of ist dorsal;
anterior margin of nostril expanded as a simple lobe.

acanthias Linnaeus, 1758, p. 455.
lb. First dorsal spine about over midpoint of inner margin of pectoral; midpoint of
bases of pelvics about midway between rear end of base of ist dorsal and origin of
2nd dorsal; anterior margin of nostril usually with a small secondary lobe. Fig. 87 F,
89 B.
2a. Inner margin of pectoral deeply concave, its inner corner acutely pointed.

cubensis Howell-Rivero, 1936, p. 473.
2b. Inner margin of pectoral only very weakly concave, its inner corner approxi-
mately a right angle.

3a. Second dorsal spine reaches to apex of fin (Fig. 87 I) ; horizontal diameter
of eye longer than distance between nostrils, and nearly % (60%) as long as
snout in front of mouth. blainville RIsso, 1826.

Mediterranean, Canaries,
Portugal, Black Sea.
3b. Second dorsal spine reaches only about % the way to apex of fin (Fig. 87 E) ;
horizontal diameter of eye a little shorter than distance between nostrils, and
less than V2 (40%) as long as snout in front of mouth.

jernandinus Molina, 1782, p. 478.

Squalus acanthias L,innztus, 1758

Spiny Dogfish, Piked Dogfish

Figures 87 A-D, 88

Study Material. About 1 20 preserved specimens, of all sizes, from embryos and new-
born to large adults, from various localities along the New England coast and Newport
News, Virginia, in the western Atlantic," from Bohuslan, Sweden, North Sea and Mediter-
ranean in the eastern Atlantic, and from Siberia, Gulf of Georgia, Puget Sound and Cali-
fornia in the North Pacific (Harv. Mus. Comp. Zool.); also many specimens, fresh-

23. Including a female, 272 mm. long, from Cuba, with umbilical scar still visible, the type specimen of S. barbouri
Howell-Rivero, 1936.

456

Memoir Sears Foundation for Marine Research

cauo-ht, from the Gulf of Maine and the vicinity of Woods Hole, Virginia and North Caro-
lina.

Distinctive Characters. S. acanthias is easily separated from other members of its
genus in the North Atlantic by the facts that its first dorsal spine is over or posterior to the
inner corner of the pectoral (about over the midpoint of the inner margin of the pectoral

Figure 87. A, Squalus acanthias, female, about 815 mm. long, from Woods Hole, Massachusetts (Harv. Mus.
Comp. Zool., No. 35862). B Head of same from below. C Right-hand nostril of same, about z x. D Second
dorsal fin of adult male from the same locality to show the length of the spine. E, Squalus jernandinus, female,
about 914 mm. long, from Island of Juan Fernandez (Harv. Mus. Comp. Zool., No. 841). F Right-hand
nostril of same, about 2 x. G Dermal denticles of same, about 20 x. H Apical view of denticle. / Second dorsal
fin of Squalus ilainville, female, 570 mm. long, from Italy (? ) (U. S. Nat. Mus., No. 28473) to show length
of spine.

in blainville and cubensis), that the midpoint of the bases of its pelvics is much nearer
to the second dorsal than to the first (about midway between the two in blainville and
cubensis) ^hy the simple anterior margin of the nostril (bilobed in blainville and cubensis) j
by the rounded inner corner of the pectoral (angular in cubensis), and by its shorter dorsal
spines and white-spotted coloration. It is further separated from blainville by the deeply
concave distal margin of its pectoral.

Description. Proportional dimensions in per cent of total length. Male, 705 mm.,
from Buzzards Bay, Mass. (Harv. Mus. Comp. Zool., No. 35864). Female, 814 mm.,
same locality (Harv. Mus. Comp. Zool., No. 35863).

Fishes of the Western North Atlantic

457

Trunk at origin of -pectoral: breadth 10.9, 1 1. 1 j height 9.2, 8.8.
Snout length in front of : outer nostrils 4.1, 4.0 j mouth 8.9, 8.6.
Eye: horizontal diameter 3.5, 3.3.
Mouth: breadth 6.9, (>.(>\ height 1,1, i.o.
Nostrils: distance between inner ends 3.4, 3.3.

Labial furrow length from angle of mouth: upper 2.4, 2.2; lower i.i, i.i.
Gill Of ening lengths: ist 1.7, 1.7; 2nd 1.6, 1.5; 3rd 1.6, 1.5; 4th 1.7, 1.7; 5th
2.1, 2.1.

First dorsal fin: vertical height 6.0, 5.7; length of base 7.1, 7.4.
Second dorsal fin: vertical height 3.8, 3.9 ; length of base 5.4, 6. i .
Caudal fin: upper margin 21.3, 20.3; lower anterior margin 10.2, ii.i.
Pectoral fin: outer margin 15.6, 15.8; inner margin 7.1, 6.8; distal margin 10. i,
II.I.

Distance from snout to: ist dorsal 33.4, 32.7; 2nd dorsal 63.8, 62.3; upper caudal
78.7, 79.7; pectoral 19.2, 18. 2; pelvics 50.7, 51.3.

Interspace between: ist and 2nd dorsals 23.0, 23.1 ; 2nd dorsal and caudal 10.7,
I1.7.

Distance from origin to origin of: pectoral and pelvics 3 1 .2, 32.7 ; pelvics and cau-
dal 28.8, 28.3.

Figure 88. Squalus acanlhias, illustrated in Fig. 87. A Upper and lower teeth, left-hand side, about 3.5 x.
B Third upper and lower teeth, about 4 x. C Dermal denticles of another Massachusetts specimen (Harv. Mus.
Comp. Zool., No. 842), about 34 x.

458 Memoir Sears Foundation for Marine Research

Trunk very slender, its height at ist dorsal only about Vr its length to origin of
caudal, its dorsal profile sloping forward from ist dorsal. Body sector to cloaca longer than
tail sector by a distance about % as long as head, and without mid-dorsal ridge. Caudal
peduncle flattened below but rounded above, with a low rounded longitudinal dermal ridge
along each side a little below the midlevel, which extends from a little behind base of
2nd dorsal to beyond origin of caudal. Upper precaudal pit subrectangular, more or less
strongly developed, although some specimens appear to lack it 5 no lower pit. Dermal denti-
cles Idosely spaced and exposing the skin, rising steeply over the trunk as a whole but lying
flat and in close contact (overlapping only a little if any) on top of snout and along edges
of fins J essentially spine-like but somewhat expanded laterally, the terminal portion
mostly a little longer than broad with a strong flat-topped median ridge, but varying con-
siderably in shape on difi"erent parts of the body; those on sides and back posterior to ist
dorsal fin as well as on upper surfaces of pectorals more or less definitely tridentate (me-
dian tooth much the largest), but interspaced here and there with faintly tridentate forms;
those on top of head less strongly tridentate, interspersed with broad-lanceolate; those on
lower surface weakly tridentate anteriorly, but mostly very narrow-lanceolate posteriorly,
without definite lateral teeth; those on sides of ist and 2nd dorsals broad-lanceolate and
but faintly ridged; those on top of snout and along anterior margins of fins broad-oval and
smooth, or very faintly ridged.'^

Head about V4 of trunk to origin of caudal, moderately flattened above. Snout rather
thick, ovate, with rounded tip, its length in front of nostrils a little more than Vs its length
in front of mouth, its length in front of mouth a little less than ^ (about 43 to 44%) of
length of head. Eye a little longer than high, its upper outline less convex than its lower,
its horizontal diameter a little more than V3 as long as snout in front of mouth or about
the same as distance between nostrils; relatively a little larger in newborn specimens than
in adults. Spiracle close behind eye, about Vs as long as horizontal diameter of eye, its
lower margin about level with upper margin of latter or a little above it. Gill openings
low down on sides, the ist to 4th about evenly spaced, but the 5th closer to 4th; the ist to
4th about same length, the 5th the longest, the latter about % as long as horizontal diam-
eter of eye or a little less than % as long as snout in front of mouth and i Vs times as long
as 1st; the 5th close in front of origin of pectoral. Nostril approximately transverse, its
inner end a little nearer to tip of snout than to symphysis of upper jaw, its anterior margin
expanded as a simple, subtriangular lobe. Mouth only very slightly arched. Upper labial
furrow extending inward and forward for a distance about V2 as long as diameter of eye,
the lower furrow V2 to % as long as upper.

'^^^'•'^ iior 12— 0— iiori2 ^'^ specimcns counted, essentially similar in the two jaws,
smooth-edged, with single sharp-pointed cusp, deeply notched outwardly and so strongly
oblique that the inner margins form a nearly continuous cutting edge from one corner
of the mouth to the other; the lowers somewhat larger than uppers, and considerably

24. For a more detailed account and discussion of the denticles, see Sayles and Hershlowitz (Biol. Bull. Wood's Hole,
73, 1937: 5)-

Fishes of the Western North Atlantic 459

widest near corners of mouth; either i or 2 series in function all along each jaw, or per-
haps even 3, depending on their stage in the process of replacement.'"'

Length of ist dorsal from origin to rear tip nearly 2.3 times its vertical height, which
is only a little more than V4 of length of head, its origin varying in Woods Hole specimens
from about over the inner corner of pectoral to posterior to the latter by a distance about
as long as horizontal diameter of eye; the spine a little longer in males than in females
(cf. Fig. 87 A with 87 D) and reaching at most to the midpoint of anterior margin of
fin, the apex rounded, posterior margin moderately concave, free rear corner a little shorter
than base, the midpoint of latter nearer to axil of pectoral than to origin of pelvics.
Interspace between ist and 2nd dorsals as long as, or somewhat longer than, head in adult,
but only about as long as from tip of snout to 2nd or 3rd gill opening in newborn speci-
mens. Second dorsal nearly as long at base as ist but only about % as high vertically, its
posterior margin more deeply concave, its free rear corner a little shorter than base, its
origin about over tips of pelvics, its spine reaching about % the way to the apex in females
but nearly to the apex in males. Interspace between 2nd dorsal and caudal about twice as
long as base of 2nd dorsal. Caudal only about ^ the total length, without subterminal
notch, its axis only very little raised, its upper margin nearly straight, apex moderately
rounded and lower margin somewhat sinuous, the lower lobe about V2 as long as upper,
but considerably less than Y2 as large in area with narrowly rounded tip ; the re-entrant
contour between the 2 lobes subrectangular with well rounded corner. Pelvics about as
long at base as 2nd dorsal, their anterior margins straight or very slightly convex, pos-
terior margins moderately concave, tips tapering, subangular, midpoint of base nearer
to origin of 2nd dorsal than to rear end of base of ist dorsal by a distance nearly or quite
as long as base of latter. Clasper of adult male subdivided at tip into 2 short rounded lobes,
the outer soft, the inner cartilaginous, with a sharp recurved hook in its inner edge. Pectoral
about tf) as long as head and a little more than V2 as broad as long, the outer margin
moderately convex toward apex, distal margin moderately and evenly concave, apex and
inner corner rather narrowly rounded.

Color. Usually slate-colored above, although sometimes tinged with brown; pale
gray, grayish white, or pure white below; a row of small white spots irregularly arranged
on each side from above the pectorals to abreast of the pelvics, with a few others in front
of and behind the first dorsal, as well as close in front of the second dorsal and scattered
on the upper sides of the anterior part of the trunk. These spots are most conspicuous in
young specimens up to 12 to 14 inches long, fading with growth, and sometimes entirely
lacking in large adults. The upper distal margins of caudal and of first and second dorsals
are dusky in at least some newborn specimens, but fade with growth.

Size. Although an occasional Spiny Dog may be born at a length no greater than
165-179 mm. (6V2-7 in.), the majority are about 220 to 330 mm. (8%-i3 in.) at
birth. It has also been observed that the smaller parent fishes contained smaller embryos

25. See discussion, p. 65.

4.60 Memoir Sears Foundation for Marine Research

than the larger parent fishes.^" Males mature" at about 600 to 800 mm. and females at 700
to 1,000 mm. in different localities. Most of the adult males are from two feet to slightly
. less than three feet in length, the females from a little less than lYz feet to almost 3 V2 feet,
averaging about 7-1 0 pounds in weight; the maximum length attained is about four feet;
occasional very large fat females may weigh 1 5 pounds, and a weight of 20 pounds has
been reported.

Developmental Stages. It has been known since Aristotle's day that the Spiny Dog
is ovoviviparous. The eggs are large, with much yolk. During the early stages of develop-
ment those in each oviduct, one to four or more in number, are enclosed in a thin, amber-
colored, horny capsule (known as a "candle") which tapers to a fine tip anteriorly but
to a blunter end posteriorly; this capsule later breaks down to leave the embryos free
in the oviduct. The young have no placental attachment to the uterine wall of the mother,
but the latter is complexly folded and has numerous highly vascular papillae. It has been
reported that the developing embryo gains about 40 per cent in weight over that of the
ripe egg, presumably by absorption of water through the yolk sac, or at least chiefly so.^'
While the embryos are developing, a fresh set of ovarian eggs are growing to take their
place. At Woods Hole the number of young in a litter is commonly four to six, sometimes
as many as eight to eleven, or as few as two.

According to recent studies the period of gestation is about 1 8 to 22 months, whether
in the Black Sea,^^ the English ChanneF" or the western Atlantic."^ Accordingly, the adult
females taken in summer and autumn in the Gulf of Maine contain either very small em-
bryos that grow to an average length of about 17 mm. by September, or much larger ones
ranging from 7 to 1 1 inches by that month, i.e., nearly full term.

Habits. Spiny Dogs are neither swift swimmers nor very active, putting up little
resistance when hooked. They may be either scattered or in schools, and in the latter case
it appears that they continue to stay together as they grow. As a rule a given school con-
sists either of small immatures of both sexes in almost equal numbers, of medium-sized
mature males and immature females, or of large mature females. It is common knowledge
that they are constantly on the move, their appearances and disappearances being so erratic
that where there may have been good fishing for cod one day there may be only Dogfish
the next, and nothing at all the day after, they having departed in pursuit of the better
fish they had driven away. They use their spines for defense, curling around in a bow to
strike, and it is probable that the spines are slightly poisonous, the general report to this

:6. See Templeman (Res. Bull. Dep. Nat. Resources Newfoundland, 15, 1944: 44) for a detailed account of the

life history of the Spiny Dog in Newfoundland waters.
27. Eng'lish Channel data.
i8. See Scammon and Minot (in Keibel, Noimaltafeln Entwick. Wirbelt., 12, 191 1) for excellent description and

illustrations of embryonic development. For recent accounts of the uterine wall and of the gain in weight of

the developing embryo, see Widakowitch (Z. wiss. Zool., SS, 1907: 499, pi. 30, 31), Ranzi (Pubb. Staz. zool.

Napoli, 75 [3], 1934: 372) and Templeman (Res. Bull. Dep. Nat. Resources Newfoundland, 15, 1944: 45).

29. Popovici, in Grig. Antipa. Homm. Ocuvre, Bucharest, 1938: 44S-

30. Ford, J. Mar. bid. Ass. U. K., N. S. 12, 1921:481.

31. Hisaw and Albert, Biol. Bull. Wood's Hole, 92 (3), 1947: 187.

Fishes of the Western North Atlantic 461

effect being corroborated by the fact that their concave surfaces are sheathed with a glan-
dular tissue resembling the poison glands of the venomous European weaver {Trachinus
draco) J'^

They may be anywhere between the surface and bottom to depths as great as 90 to 100
fathoms, and perhaps even deeper. But they have never been found to be pelagic in the
ocean basin. On the other hand, while their ability to survive in brackish water has been
proved by experiment, and although they have been reported in at least one river" in
Denmark, they do not normally enter fresh water, and specimens placed in fresh water
died within a few hours."

Relationship to Temperature. Spiny Dogfish do not appear on the United States coast
in spring until the temperature of the water has risen to about 6° C, and most of them dis-
appear from the inshore belt west and south of Cape Cod by the time the surface has
warmed to about 15°, either moving into deeper cooler water nearby or northward to
colder seas. Similarly, during their summer stay farther north they are seldom taken in
water warmer than about 1 5° or colder than 6 or 7°, unless temperatures within this range
are to be found a few fathoms shallower or deeper. Their northward advance along
the coasts of Newfoundland and Labrador clearly appears to follow the vernal warning
of the coastal waters, and their autumnal reappearance west and south of Cape Cod coin-
cides roughly with the date when the surface has cooled to 12 to 15° i few if any are seen
anywhere along the coast, north or south, after the surface has chilled below about 6° or 7° ;
and the winter temperature ranges between 6° and 1 1° on the bottom along the offshore
belt, where the majority of the stock is now known to spend the cold months.

The foregoing, added to similar data for European waters, shows that the seasonal
migrations north and south, and between shoal water and deep, are chiefly thermal in
character, i.e., to avoid extremes of temperature, either lower than about 7° or 8 ° or higher
than about 1 2° to 15°. The thermal relationship of the Spiny Dog is thus analogous with
that of the Mackerel, except that its optimum thermal range is slightly lower.

Breeding Habits. Females, with young nearly ready for birth, are taken in New-
foundland waters in early autumn (October), in the Gulf of Maine in late summer and
autumn, near Woods Hole and New York in autumn, oflF Virginia and North Carolina
in January to February. This, with the fact that very young specimens are rarely seen
anywhere on the coast, suggests that most of the young are born from late autumn through
the winter on the oflFshore wintering grounds. However, the season of production may
extend through the spring in some years, as evidenced by recent captures of young Dogfish
with the umbilical scar still discernible, near Woods Hole in June, in the Gulf of Maine
in summer, and in schools at the entrance to Long Island Sound in July. It may even
extend sporadically into summer, which is proved by a catch of 74 adult females which

32. For details, see Evans (Philos. Trans., [B] 212, 1923: 27).

33. The fact that they were reported there with cod and Merlucciui (Feddersen, Naturh. Tidt^kr., [3] 72, 1879:
68, 69, footnote i; Bean, Amer. Nat., 14, 1880: 525-526) indicates that the bottom water in the stream io
question was salt, or at least brackish and not fresh.

34. For accounts of experiments on the survival of Spiny Dogfish in brackish and fresh water, see Scott (Ann. N. Y.
Acad. Sci., 33, 1913: 30, 60).

4-62 Memoir Sears Foundation for Marine Research

gave birth to young in July on capture off Gloucester, Massachusetts.*" Young are
produced throughout the year in the Mediterranean, during autumn in the Black Sea,
while in more northern European waters there is wide regional variation, i.e., late summer
through autumn into winter in the English Channel, and late April through the summer
in the North Sea and in Scandinavian waters.'^ It is probable that pairing takes place shortly
after the young are born, but no definite information is available.

Food. The Spiny Dog is as voracious as any fish of its size, and its wanderings on
the coast are no doubt chiefly in pursuit of food. Its recorded diet in the western Atlantic
includes capelin, herring, menhaden {Brevoortia), mackerel, scup {Stenotomus), silver
hake (Merluccius), cod, haddock, pollock, blennies and croakers (Micropogon). No
doubt it preys on practically any species of fish smaller than itself. Even when newly
born they have been seen attacking herring much larger than themselves, as adults do cod
and haddock. Fishermen have often described them as harrying schools of mackerel and
herring even in the seines, as well as destroying large numbers of cod and haddock in
addition to driving them away, and they often bite these and other ground fish from the
hooks of long lines. They also prey on squid and to some extent on worms, shrimps, prawns,
crabs and amphipods. They occasionally feed on gastropods and jellyfish (Aurelia), and
even red, brown and green algae have been found in their stomachs.^*" When they first
arrive near Woods Hole in spring they are often full of ctenophores. Probably they feed
very little during the winter, for fishermen describe them as thin when they reappear on
the coast in spring.

Relation to Man. In northern Europe the Spiny Dogfish is of considerable commer-
cial value as a food fish; in 193 1, for example, it fetched the fishermen about 5 cents per
pound in the markets of Germany, and the landings for that year came to perhaps 14,000,-
000 pounds, as estimated from the total catch of sharks of all sorts." Similarly, the British
landings for 1923 were 9,597,900 pounds, worth £49,980.'* However, on the American
coast the Spiny Dogfish has never been in demand for the table, although many years ago
they were of some value for oil. During more recent periods when they were in great
abundance, various efforts were made in America to utilize Dogfish on a large scale both
as fertilizer and as a source of oiP^ to combine with cod liver oil (it compares favorably
with the cod for Vitamin A, although its Vitamin D content is much lower) ; it has also been
canned for human consumption. And fresh, this is a better food fish than is generally appre-
ciated. On the coasts of the eastern United States and Canada, however, these attempts
have been short-lived. Of late years Spiny Dogfish have been so little considered that there
is no way of knowing how great a proportion of the total landings of sharks of all kinds

35. Mclntire, in Rep. Comm. Fish. Game Mass. (1905), 1906: 108.

36. For details and authorities, see Ford (J. Mar. biol. Ass. U. K., N. S. /;, 1921 : 481, 482).

36a. For a recent list of stomach contents, see Templeman (Res. Bull. Dep. Nat. Resources Newfoundland, 1944: 49).

37. For details, see Lijbbert and Ehrenbaum (Handb. Seefisch. Nordeurop., 2, 1936: 285).

38. Jenkins, Fish. Brit. Isles, 1925: 321.

39. Liver oil in the amount of 176,200 gallons was produced from this species in Canada in .936 (Hampton,
Newfoundl. Fish. Res. Inst. Serv. Bull., 5, 1938: 5).

Fishes of the Western North Atlantic 463

from the Gulf of Maine to North Carolina may have consisted of them. The United
States Bureau of Fisheries reported about 1,250,000 pounds for 1938.

From a practical aspect the Spiny Dog in the western Atlantic is chiefly important
because it is undoubtedly more destructive to gear and interferes more with fishing opera-
tions than does any other fish — shark or teleost. Its habit of taking the bait is proverbial.
In fact, when Dogfish are plentiful, hook and line fishing for cod, haddock and other
ground fish is often actually prevented unless cockles (Lunatia) are used for bait. Still
more serious is the damage they do by tearing and biting nets, biting snoods off long lines,
attacking netted or hooked fish and by driving away better fish. It has been estimated that
in these ways they do some $400,000 worth of damage annually off the coast of Massachu-
setts alone, and perhaps much more in their periods of abundance.*"

Range. Both sides of the North Atlantic, chiefly in temperate to subarctic latitudes;
also both sides of the northern Pacific south to California, Japan, northern China and the
Hawaiian Islands (as pointed out on page 453, suckleyi appears to be indistinguishable
from acanthias). It is represented in the corresponding thermal belt of the southern
hemisphere (South Atlantic, Pacific, Indian Oceans, South Africa) by relatives so close
that it is still an open question whether or not any valid specific distinctions can be drawn."

Occurrence in the Eastern Atlantic. The chief center of abundance for the Spiny
Dog is from the Atlantic coast of France north to Ireland, Scotland and southern Scan-
dinavia, including the English Channel and the North Sea in general, and as far eastward
as the Kattegat. But it rarely enters the Baltic. The Spiny Dogfish is plentiful around the
Orkneys, Faroes, and south and east of Iceland in season, but less so to the north and west;
it occurs regularly ofi^ Norway and as far north and east as the Murman coast. It is also
generally distributed in the Mediterranean and in the Black Sea. To the southward it
occurs commonly off Morocco and is reported from the Canaries, Madeira and Senegal.

Occurrence in the Western Atlantic. Fishermen are familiar with it in season all along
the coast from North Carolina to Nova Scotia and on the southern side of the Gulf of
St. Lawrence, as well as offshore on Nantucket Shoals, on Georges and on Browns Banks
and on the Nova Scotian and Newfoundland Banks. It is common northward along both
coasts of Newfoundland and is known past the Straits of Belle Isle to southeastern Labra-
dor. It is also recorded on the north shore of the Gulf of St. Lawrence from Red Bay. And
specimens have been taken on the west coast of Greenland at Sukkertoppen and Holstein-
borg, no doubt these being visitors with the summer drift of Atlantic water. But there is no
record of it on the east coast of continental North America to the north of Hamilton Inlet.

Southward it is a yearly visitor as far as Cape Lookout, North Carolina. But it is
doubtful whether it occurs coastwise any further in that direction in numbers, for while
it has been described repeatedly as plentiful in East Florida waters, and around Cuba and
Trinidad, the former report seems likely to have referred to some other fish,*" the latter

40. For detailed records and discussion, see Rep. Comm. Fish. Game Mass. (1905), 1906: 97; (1906), 1907: 20.

41. For discussion of this question, see p. 453.

42. Evermann and Bean (Rep. U.S. Comm. Fish. [1896], 1898: 239) describe it as "probably the most abundant

464 Memoir Sears Foundation for Marine Research

chiefly to the newly described S. cubensis (p. 473). However, at least a few stray as far as
southern Florida and Cuba." Offshore its range reaches to the outer edge of the continental
shelf j inshore, into the outer reaches of Chesapeake, Delaware, Narragansett and Passa-
maquoddy Bays and the larger harbors. But it rarely, if ever, enters river mouths, at least
on the American coast (but see footnote 33, p. 461).

Seasonal Migrations. The Spiny Dog is a spring and autumn migrant in the southern
coastwise section of its range from North Carolina to New Yorlc, and mostly so along the
southern coast of New England, but it is chiefly a summer visitor to the Gulf of Maine (in-
cluding Georges Bank) and more northerly waters. South of New York Spiny Dogs are apt
to "strike in" nearly simultaneously all along the coast j there are records for New Jersey
(March 6, 13, or even earlier) and Chesapeake Bay (March) as early in the season as for
North Carolina (April and early May). But the date of their arrival varies considerably
from year to year. They depart entirely from Chesapeake Bay and the coast south of it by
early May in some years and by late May at the latest j but they do not leave New Jersey
waters and the immediate vicinity of New York until early or middle June. In the Long
Island and southern New England areas they usually do not appear before late April or
early May," and the majority have departed by the end of that month, or by the close of
June at the latest. But even in July and August considerable numbers of adults are taken
at the mouth of Long Island Sound in deeper water ( 1 7 to 24 fathoms) while schools of
young are taken inshore j and odd specimens are caught near Woods Hole throughout the
summer in some years. On Georges Bank, in the only year of record, a few were taken in
late March and April, but not until late June did their numbers sharply increase, the peak
of abundance continuing through August. In the western side of the Gulf of Maine they
may appear as early as mid-May, as in 1903, or not until well into June, as in 1905 and
1 913, when the first big run struck near Cape Ann about the middle of that month. But
there may be wide variation in this respect from place to place, as in 1903, when they did
not appear at the tip of Cape Cod until early July, although they were numerous a month
earlier in Massachusetts Bay, near Cape Ann, and off Penobscot Bay. In most years they
have also appeared by June in the eastern part of the Gulf of Maine in general, although
not until July in the cold waters of Passamaquoddy Bay, tributary to the Bay of Fundy.
Within Massachusetts Bay, where the surface warms to about 1 8 ° C. in summer, few are
taken between June and September, and the diminution recorded on Georges Bank after
July similarly suggests a movement thence into the Gulf of Maine as the water warms.

Shark in the Indian River," Florida, and as a permanent resident there. But it is proba.ble that this record actually
referred to some other small shark, or perhaps even to Amia (known locally as Dogfish), for the Spiny Dog
has never been definitely reported subsequently for the east coast of Florida, either in scientific literature or by
fishermen.

43. We have examined the embryos recorded from the Tortugas by Longley and Hildebrand (Pap. Tortugas Lab.,
34, 1942: 3) and we have found nothing to separate the type of S. barbouri Howell-Rivero, taken off Havana,
from young free-swimming acanthias of the same size from New England waters.

44. May 17 is the earliest date recorded for the eastern end of Long Island; in 1940 they were first taken at Woods
Hole on May 9, or six days after the last were taken south of Delaware Bay (Carolina Biol. Supply Co., Caro-
lina Tips, Elon Coll., N. Carolina, 3 [7], 1940: 25)-

Fishes of the Western North Atlantic 465

Along outer Cape Cod, however, they are present in varying abundance all summer, at
least in some years. And this is the case generally thence eastward and northward as far as
the species occurs. Ordinarily they appear all along the outer coast of Nova Scotia about as
early as in the eastern side of the Gulf of Maine, and a little later (third week in June)
along the southeastern coast of Newfoundland, but it is well into July before they are
encountered in any numbers in the inner parts of the Gulf of St. Lawrence. In 1942 (the
one year of record) they had advanced to the Straits of Belle Isle along both the St. Law-
rence and Atlantic coasts of Newfoundland by the second week in July. But they were not
reported in southeastern Labrador until the beginning of September.

In general the autumnal withdrawal takes place as early from the western side of the
Gulf of Maine as from Labrador, Newfoundland, Nova Scotian waters, Cape Breton or the
Gulf of St. Lawrence, the majority ordinarily departing during October, with few caught
in November, and as a rule they depart even earlier from the smaller bodies of water {e.g.,
Passamaquoddy and Massachusetts Bays) than off the outer coast. In some years, however,
they may be present in abundance well into November, as in 1903 and again in 1942, or
even into December, as in 1913, when large catches were made between Cape Ann and
Cape Elizabeth from the fifth to the twelfth,**'' and again in 1 942 when they were reported
along the eastern, southeastern and western shores of Newfoundland. Of especial interest
is the definite record of a number of Dogfish washed ashore on January 1 1, 1939, in St.
Marys Bay, Newfoundland, after a severe galej"" the implication of this report is dis-
cussed on page 466 in relation to their winter home.

Corresponding to this withdrawal from the north they reappear in autumn all along
the coast from southern New England to North Carolina, their appearance being succes-
sively later from northeast to southwest. And the fact that catches on Georges Bank do not
show any marked peak at that time indicates that the autumnal migration route is mostly
along shore at first. In the Woods Hole region, near Nantucket and on Nantucket Shoals,
they reappear in October, ordinarily disappearing again in November. At the mouth
of Long Island Sound the recorded dates of their autumnal arrival in numbers have varied
between October 5 and November 7, their subsequent disappearance between Novem-
ber 24 and December 12. Near New York, where they remain into December, they seldom
appear in any numbers until November, and similarly along the New Jersey coast, where
they may arrive late in October and remain plentiful into the winter.*' At Cape Charles,
at the mouth of Chesapeake Bay, the earliest autumnal record is for November 1 5. How-
ever, they have been reported as early as November 7*° from North Carolina, and we have
ourselves seen them in great numbers, dead on the beach (discarded by seiners) here and
there on the coasts of southern Virginia and northern North Carolina late in that month

44a. For this record, for recorded dates of arrival on the Newfoundland coast, and for months when Dogfish were
present, see Templeman (Res. Bull. Dep. Nat. Resources Newfoundland, 15, 1944: 56-66, fig. 13-16).

44b. Schools were reported as seen at the surface off Portsmouth, New Hampshire, on February 10, 1882 (Collins,
1883), but there is no proof of identity.

45. Precise information is scanty. 46. Carolina Tips, Elon Coll., N. C, 3 (7), 1940: 26.

466 Memoir Sears Foundation for Marine Research

and early in December. It is certain that at least some of the Dogfish that summer as far
north as Newfoundland journey southward in autumn past the Gulf of Maine, for one
tagged near St. John on July 14, 1942, was recaptured on the 23rd of the following No-
vember, off Thatcher's Island, Cape Ann, Massachusetts, having travelled a distance of at
least 1,000 miles during the interval of 132 days or at an average of about 7.6 miles per
dayj actually, it probably travelled much farther and faster, for it is not likely to have
followed a straight line. An equally interesting case is that of another fish tagged near St.
Johns, Newfoundland, in April 1942 which was recaptured in September of the following
year at the mouth of the Bay of Chaleur, within the Gulf of St. Lawrence;" it is evident
that Dogfish that summer off one part of the coast during one year may do so off some
other coast many miles distant during another. Additional information in these respects is
much to be desired.

Wintering Grounds. It now seems certain that the Spiny Dogfish winter chiefly on
bottom in deeper water offshore, from the ofiing of New York southward, for while none
are reported from Georges Bank in February (though a few in January, however), con-
siderable numbers have been trawled on the outer part of the shelf off New York in late
November and in January and likewise in depths of 16 to 70 fathoms*' between the
offings of Delaware Bay, of northern Virginia and of Cape Hatteras in February. The
fact that Spiny Dogfish have been washed ashore in some numbers on the southwest
coast of Newfoundland in mid-January (p. 465) also opens the very interesting possi-
bility that some of those that summer in that general region may survive the winter in the
trough of the Gulf of St. Lawrence in temperatures (4°-5° C; or 39°-4i° F.) consider-
ably colder than those that prevail on the southern wintering grounds.

Numerical Abundance. During its periods of abundance this is by far the most nu-
merous local shark; in fact, it is the only one that even remotely rivals the commercially
important food fishes in abundance. It has been described repeatedly as "in great abund-
ance," in "schools of thousands," or as being caught as fast as fishermen can haul them
in. Unfortunately the statistics of commercial landings do not afford any Information
in this respect about the Dogfish in American waters. But the foregoing is no over-
statement, judging from such records as the following: 690 caught on a 700 hook line at
Cape Breton; a Dogfish on nearly every one of 1,500 hooks in the Gulf of Maine; three
wagon loads from a single lift of two pound-nets on Long Island; 1,800 pounds in one
day in pound nets in North Carolina} more than two tons preserved for use in biological
laboratories in a three- weeks' period; or an average trawl catch of 6,000 to 8,000 per trip
on Georges Bank during the peak of abundance in 1913. At the time of the 1904-1905
peak of abundance it was estimated, from reports of fishermen's catches, that at least
27,000,000 were taken yearly off the coast of Massachusetts alone." There is, in short, no

47. For these and other tagging records, see Templeman (Res. Bull. Dep. Nat. Resources Newfoundland, 15, 1944:
67, fig. 18).

48. For details, see Bigelow and Schroeder (Bull. U.S. Bur. Fish., ^S, 1936: 323).

49. Rep. Comm. Fish. Game Mass. (1906), 1907: 20.

Fishes of the Western North Atlantic 467

reason to doubt that the concentrations may be as great in the western side of the Atlantic
as in the eastern, where the record catch of 20,000 in a single haul was made many years
ago on the Cornwall coast. The foregoing instances also show that Dogfish may be as
plentiful, on occasion, oflF one sector of the coast line as off another from North Carolina
to Cape Breton. But in general they are much less plentiful in the bays that they pene-
trate {e.g., Chesapeake, Delaware, Passamaquoddy) than off the open coast.

The Spiny Dog fluctuates very widely in abundance over periods of years, but there
is so much irregularity on different parts of the coast, and the peaks of maximum abund-
ance fall so far apart, that no consistent picture of its ups and downs can yet be offered.
Around Newfoundland there seems to have been a period of scarcity for five or six years
prior to 1871,''° but on the other hand a period of abundance in southern New England
waters about 1875 to 1880. Available information points next to a pronounced peak about
1904 or 1905. While reports of local fluctuations reflect in part the movements of great
schools that may visit one locality in one year and another the next, less complaint has been
made of them since about 1 9 1 3 than previously.

Synonyms and References:"'

I. North Atlantic:

Squalus acanthias Linnaeus, Syst. Nat., /, 1758: 233 (descr., refs., Europ. Oc.) ; i, 1766: 397; Olafsen
and Povelsen, Reyse en Island, 1772: 359 (Iceland) ; Miiller, Prod. Fauna Danica, 1776: 3 (Denmark) ;
Olavius, Oecon. Reyse Island, 1 780: 80 (Iceland, not seen); Bloch, Fische Deutsch., 3, 1784: 74,
pi. 85 (general, early devel., size at birth); Mohr, Fors0g Island Naturh., 1786: 57 (Iceland);
Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: II, pi. 5, fig. 12 (descr., ill.); Blumenbach, Handb.
Naturg., 1788: 294 (general) and subsequent eds.; Walbaum, P. Artedi Genera Pise. Emend. Ichthyol.,
1792: 505, pi. 3 (refs.) ; Forster, in Latham and Davis, Faunula Indica, 1795: 13 (ref. to Bloch, 1784) ;
Vandelli, Mem. R. Acad. Lisboa, /, 1797: 70 (not seen); Bloch and Schneider, Syst. Ichthyol., 1801:
135 (descr., refs.); Bosc, Nouv. Diet. Hist. Nat., 2/, 1803: 191 (diagn.) ; Donovan, Nat. Hist. Brit.
Fish., 4, 1805: 82, pi. 82 (text not seen); Turton, Brit. Fauna, 1807: 114 (descr., Gt. Brit.); de la
Roche, Ann. Mus. Hist. nat. Paris, 75, 1809: 314 (Ivica, Balearic Is. in market); Risso, Ichthyol. Nice,
1810: 40 (general, Medit.) ; Faber, Fische Islands, 1829: 79 (descr., no. of embryos, Iceland);
Nilsson, Prod. Ichthyol. Skand., 1832: 117; Johnston, Hist. Berwicksh. Nat. CI., /, 1834: 17 (Gt. Brit.,
not seen); Jenyns, Manual Brit. Vert. Anim., 1835: 505 (Brit, coast); Couch, Cornish Fauna, 1838:
51 (Cornwall, abund.) ; Fries, Ekstrom and Sundevall, Skand. Fisk., 1 845: 187, pi. 46 (descr., Scand.) ;
Gaimard, R. Voy. Islande et Greenland, Vol. Zool. Med., 1851: 163 (Iceland); Gray, Cat. Fish.
Coll. Descr. by L. T. Gronovir in Brit. Mus., 1854: 8 (listed); Knight, Descr. Cat. Fish. N. S.,
1866: 8 (Nova Scotia) ; Jones, List Fish. Nova Scotia, 1879: 9 (common. Nova Scotia) ; Bean, T. H.,
Proc. U.S. nat. Mus., j, 1881: 1 16 (New England Iocs.); Jones, Proc. N. S. Inst. Sci., 1882: 95
(same as Jones, 1879); Jordan and Gilbert, BuU. U.S. nat. Mus., 16, 1883: 16 (descr., value for oil,
N. and Mid. Atlantic, U.S.); Bean, T. H., Rep. U.S. Comm. Fish. (1882), 1884: 344 (Woods Hole);
Goode, Fish. Fish. Indust. U.S., I, 1884: 673 (distrib., habits, abund.); Kingsley, Stand. Nat. Hist., 5,
1885: 76 (general); Rathbun, Proc. U.S. nat. Mus., 7, 1885: 490 (parasites); Whiteaves, Cat. Canad.
Pinnep. Cetacea, Fish., 1886: 5 (Gulf of St. Lawrence); Jordan, Rep. U.S. Comm. Fish. (1885), 1887:
793 (distrib.); Nelson, Final Rep. St. Geol. N. J., 2 (2), 1890: 659 (common, N. Jersey);

50. Saxby, Zoologist, (2) d, 1871 : 1554.

51. Many studies have been made of the anatomy of Squalus acanthias, and it has served as the subject of numerous
directions for laboratory dissection for the use of students. We have not thought it necessary to burden the present
list with citations of these sorts.

468 Memoir Sears Foundation for Marine Research

Smitt, Hist. Scand. Fish., 2, 1895: 1159, pi. 52, fig. l (descr., habits, fishery); Jordan and Everraann,
Rep. U.S. Comm. Fish. (1895), l8g6: 219 (Atlant. coast, around Cuba) ; Bull. U.S. nat. Mus., 47 (l),
1896: 54 (descr., n. and mid. U.S., Cuba); Bean, T. H., Bull. Amer. Mus. nat. Hist., 9, I 897: 330
(off N. Jersey); Knipowich, Annu. Mus. zool. Acad. St. Petersb., 2, 1897: 154 (Murman coast);
Bumpus, Science, N. S. 8, 1898: 850 (breeding season, size of embryos, C. Cod) ; Smith, Bull. U.S. Fish
Comm., ly, 1898: 89 (abund., season, commercial use. Woods Hole) ; Jordan and Evcrmann, Bull. U.S.
nat. Mus., 47 (4), 1900: pi. 7, fig. 24, 24a (ill.) ; Herring, Rapp. Fisk. Faroerne og Island Fiskberet. Kbh.,
1901 : 207 (Faroes, Iceland, not seen) ; Bean, Rep. For. Comm. N. Y., 1901 : 381 (season, abund., near
N. York); Linton, Bull. U.S. Bur. Fish., rp, 1901: 274 (parasites. Woods Hole), 430 (stomach con-
tents); Gregg, Where to Catch Fish., 1902: 23 (descr., but no record of occur.) ; Johansen, Beret. Fisk
"Diana" Islands (1898-1900), Fisk Beret. Kbh. (1901-1902), 1903: 250 (Iceland, not seen); Bean,
T. H., Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 43 (descr., season, util., N. York and New England) ; de
Braganza, Result. Invest. Sci. "Amelia," 2, 1904: 56 (Portugal) ; Sharp and Fowler, Proc. .\c^id. nat. Sci.
Philad., 56, 1904: 506 (season, abund., damage to nets, Nantucket); Collet, Norges Fisk., 5, 1905: 96
(reproduction, Norway) ; Genz0e, Fiskberet. Kbh. ( 1 903-1 904), 1905: 290 (Iceland, not seen) ; Jordan,
Guide to Study Fish., /, 1 905: 545 (general) ; Wilson, Proc. U.S. nat. Mus., 28, 1 905: 573 (parasites) ;
Fowler, Rep. N. J. Mus. (1905), 1906: 67 (abund., season, N. Jersey); Pictschmann, Ann. naturh.
(Mus.) Hofmus. Wien, 21, 1906: 74 (abund., Iceland); Tracy, Rep. R. I. Comm. inl. Fish., 1906:
46 (abund., season, Rhode Island); Cornish, Contr. Canad. Biol. (1902-1905), 1907: 81 (abund.,
north. Nova Scotia) ; Field, Rep. U.S. Comm. Fish. (1906), Spec. Pap. 6, 1907: 18, pi. I, fig. 2 (.ibund.
NW. .Atlant., food, possible util.); Jensen, in Schi0dte and Hansen, Zool. Danica, 72, 1907: 308,
pi. 28, fig. 3 (descr., depth, abund., food, devel., Denmark) ; Widakowitch, Z. wiss. Zool., 88, 1907: 499,
pi. 30, 31 (struct, of uterine wall, gain in weight by embryo) ; Wilson, Proc. U.S. nat. Mus., 57, 1907:
711 (parasites); Proc. U.S. nat. Mus., ^j, 1907: 452 (parasites, Woods Hole); Fowler, Proc. Acad,
nat. Sci. Philad., 60, 1908: 68 (New England, N. Jersey); Rep. N. J. Mus. (1907), 1908: 128
(season. Iocs., N. Jersey); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 8 (New England
Iocs.); Regan, Ann. Mag. nat. Hist., (8) 2, 1908: 46 (class., refs., distrib., size); Fowler, Proc. Acad,
nat. Sci. Philad., 61, 1909: 407 (N. Jersey); Saemundsson, Skr. Komm. Havunders. Kbh., 5, 1909:
115 (Iceland); Gill, Smithson. Misc. Coll., 52, 1909: 158 (ill.); Tracy, Rep. R. I. Comm. inl. Fish.,
1910: 61 (food, size, Rhode Island); Scammon and Minot, in Keibel, Normal Tafeln Entwick.
Wirbelt., 191 1: 140 pp., 5 pis. (descr., ill., embryonic develop.); Gudger, Proc. biol. Soc. Wash.,
2$, 1912: 143 (N. Carolina); Roule, Bull. Inst, oceanogr. Monaco, 243, 1912: 11 (Brittany, Morocco,
Medit.) ; Garman, Mem. Harv. Mus. comp. Zool., j(5, 1913: 192, pi. 14, fig. 1-4, pi. 43, fig. 9-10, pi.
59, fig. 1-2 (refs., descr., ill.); Gudger, Proc. biol. Soc. Wash., 26, 1 91 3: 98 (N. Carolina);
Halkett, Check List Fish. Canad., 1913: 41 (Delaware to Davis Strait); Le Danois, Ann. Inst,
oceanogr. Monaco, 5 (5), 1913: 21 (Engl. Channel); Nichols, Abstr. Linn. Soc. N. Y., 20-23, 1913:
91 (near N. York); Scott, Ann. N. Y. Acad. Sci., 25, 1913: 30, 60 (physiol., in fresh and brackish
water, season near N. York); Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., 31 (2), 1913: 737
(abund., season, embryos. Woods Hole) ; Jensen, Mindeskr. Jap. Steenstrup, F0ds., Kbh., (2) 30, 1914: 7
(W. Greenland, Iceland, Murman Coast, Faroes); Coles, Proc. biol. Soc. Wash., 21?, 191 5: 94 (season,
N. Carolina); Fowler, Proc. Acad. nat. Sci. Philad., 67, 1 91 6: 517 (Nova Scotia); Wilson, Proc. U.S.
nat. Mus., 47, 1915: 656 (Maine); Fowler, Copeia, 30, 1916: 36; Copeia, 31, 1916: 41 (in list,
N. Jersey); Nichols and Murphy, Brooklyn Mus. Quart., 5 (4), 1916: 150 (abund., season, near
N. York) ; Nichols and Murphy, Brooklyn Mus. Sci. BuU., 3 (l), 1916: 31, pi. 2, fig. I (ill., abund.,
migration, season, devel., util., near N. York); De Nyse, Bull. N. Y. zool. Soc, jp, 1916: 1424 (as a
food fish); Radcliffe, Bull. U.S. Bur. Fish., 34, 1916: 267 (teeth, denticles, abund., season, N.
Carolina); Fowler, Proc. Acad. nat. Sci. Philad., <5p, 1917: 109 (N. Jersey); Proc. Boston Soc. nat.
Hist., 35, 1917: 110 (Gulf of Maine Iocs.); Latham, Copeia, 41, 1917: 17 (abund., season. Long
Island, N. York); Copeia, 57, 1 91 8: 53 (season, young in summer, Long Island, N. York); Hunts-
man, Trans, roy. Soc. Canada, (3) 12 (4), 1 91 8: 63 (Gulf of St. Lawrence); Fowler, Proc.
Acad. nat. Sci. Philad., yi, 1920; 292 (season); Latham, Copeia, 71, 1919: 54 (abund., season,
young. Long Island, N. York); Roule, Result. Camp. sci. Monaco, 1919: 117 (Norway, Gulf of

Fishes of the Western North Atlantic 469

Gascony, Morocco, food); Fowler, Proc. biol. Soc. Wash., 33, 1920: 145 (N. Jersey loc.) ; Latham,
Copeia, 87, 1920: 91 (young, abund.. Long Island, N. York in summer); Ford, J. Mar. b!ol. Ass.
U. K., N. S. 12, 1921: 468 (abund., season, commercial util., breeding, habits, Plymouth, Engl.);
Latham, Copeia, 99, 1921: 72 (occur.. Long Island Sound, N. York); Mavor, Contr. Canad. Biol.
(191 8-1 920), 1 921: 129, 130 (occur., Passamaquoddy region, util.); Fowler, Proc. Acad. nat. Sci.
Philad., 74, 1922: 3, 5 (N. Jersey); Huntsman, Contr. Canad. Biol. (1921), 3, 1922: 56 (abund.,
B.iy of Fundy); Drensky, Trav. Soc. bulg. Sci. nat., 79, 1923: 56, pi. 58, fig. I (descr., abund.. Black
Sea) ; Hubbs, Copeia, 123, 1923: loi (Maine) ; Meek and Hildebrand, Field Mus. Publ. Zool., 15 (l),
1923: 65 (in key); Breder, Copeia, 127, 1924: 26 (season); Linton, Proc. U.S. nat. Mus., 64 (21),
1924: 4, 20, 26, 52, 57, 63, 64, 92 (parasites. Woods Hole); Bigelow and Welsh, Bull. U.S. Bur.
Fish., 40 (l), 1925: 44, 551 (habits, abund.. Gulf of Maine); Fowler, Copeia, 143, 1925: 43 (N.
Jersey); Proc. Acad. nat. Sci. Philad., y8, 1927: 276 (comp. with Chilean spec); Manter, Illinois
biol. Monogr., 10 (2), 1926: 8 (parasites, Mt. Desert, Maine); Nichols and Breder, Zoologica, N. Y.,
9, 1927; 21 (abund., season, food, also as food fish, N. York to south. New England); Borodin, Bull.
Vanderbilt Oceanogr. (Mar.) Mus., i (i), 1928: 5 (Newfoundland); Hildebrand and Schroeder,
Bull. U.S. Bur. Fish., 43 (i), 1928: 52 (descr., devel., migrations, season in Chesapeake Bay); Rey,
Fauna Iberica Feces, /, 1928: 424, pi. 4, fig. i (descr., ill., Spanish waters); Breder, Field Bk. Mar.
Fish. Atlant. Coast, 1929: 24 (general); Jordan, Manual Vert. Anim. NE. U.S., 1929: 14 (general);
Truitt, Bean and Fowler, Bull. Md. Conserv. Dep., 5, 1929: 28 (Maryland); Jordan, Evermann and
Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 21 (both sides N. and mid. Atlant. U.S., Cuba) ; Bere,
Contr. Canad. Biol., N. S. 5 (13), 1930: 427 (parasites, Passamaquoddy Bay); Schroeder, Bull. U.S.
Bur. Fish., 46, 1930: 11, 72 (enemy of cod, migrations, season, Nantucket Shoals) ; MacCallum, Proc.
U.S. nat. Mus., 79 (26), 1931: 3 (parasites); Schroeder, Copeia, 1931: 41, 42 (winter, depth, off Dela-
ware Bay) ; Breder, Copeia, 1932: 31 (off Block Island) ; Anonymous, Newfoundland Fish. Res. Comm.
Rep., I (4), 1931: 107 (Nova Scotian and Newfoundland Banks, Nfd. coast); Wilson, Bull. U.S. nat.
Mus., 158, 1932: 511 (parasites. Woods Hole); Anonymous, Newfoundland Fish. Res. Comm. Rep.,
2 (l), 1932: 125 (Nova Scotian Banks, off NE. Newfoundland, Gulf of St. Lawrence); Rep., 2 (2),
'934: 115 (Nova Scotian Banks, Straits of Belle Isle); Belloc, Rev. des Trav. Peches Marit., 7 (2),
1934: 140 (Morocco to Senegal, Canaries, Madeira); Bigelow and Schroeder, Canad. Atlant. Fauna,
biol. Bd. Canad., 12', 1934: 17 (general, east. Canad., W. Greenland); Vladykov and McKenzie,
Proc. N. S. Inst. Sci., rp, 1935: 48 (abund., season, util.. Nova Scotia); Bigelow and Schroeder, Bull.
U.S. Bur. Fish., 48, 1936: 322 (breeding, winter habitat) ; Fowler, Bull. Amer. Mus. nat. Hist., yo (l),
1936: 69 (refs., descr., ill., embryo, adult, E. Atlant. to Morocco); Lubbert and Ehrenbaum, Handb.
Seefisch. Nordeurop., 2, 1936: 286, pi. 21, fig. 251 (ill., adult, embryos in capsule, general, N. Europe);
Howell-Rivero, Proc. Boston Soc. nat. Hist., 41, 1936: 47, 48 (comp. with S. cubensis and barbouri);
Smith, H. W., Biol. Rev., //, 1936: 64 (in fresh water in Scand.) ; Cadenat, Rev. des Trav. Peches Marit.,
^o (4). 1937:431 (C. Blanc) ; Fowler, Proc. Acad. nat. Sci. Philad., Sg, \^S7- ^97 (N. Jersey) ; Gudger,
Sci. Mon. N. Y., ^5, '937: 5 10 (off Chesapeake Bay, Gulf of Maine) ; Sayles and Hershlowitz, Biol. Bull.
Wood's Hole, 73, 1937: 51, i pi. (dermal denticles); Raven and LaMonte, Amer. Mus. Novit., 902,
1937: I (eaten by swordfish) ; Breder, Bull. N. Y. zool. Soc, 41, 1938: 28 (N. York Harbor); Hampton,
Newfoundland Fish. Res. Inst. Serv. Bull., 5, 1938: 3 (possible util., production of liver oil in Canada) ;
Popovici, in Grig. Antipa, Homm. Oeuvre Bucharest, 1938: 445 (breeding habits. Black Sea) ; Tortonese,
Atti Soc. ital. Sci. nat., 77, 1938: 308 (Medit., Canaries, Massachusetts) ; Springer, Proc. Fla. Acad. Sci.,
3> 1939: 37 (no definite Florida record given); Popovici, Bull. Acad. Roumaine, 22 (2), 1939: I
(develpp., abund. in Black Sea) ; Fowler, Bull. U.S. nat. Mus., 100 (13), 1941: 257 (N. Atlant. refs.) ;
Hildebrand, Copeia, 1941 : 222 (N. Carolina, April, May) ; Holmgren, Acta zool.. Int. Tidskr. zool., 1-3,
1941: 29> 55 (skull); Longley and Hildebrand, Pap. Tortugas Lab., 34, 1941: 3 (embryos, Tortugas,
Florida); Fowler, Fish Culturist, 21 (9), 1942: 66 (listed, Cuba); Lunz, Bull. S. Carolina St. Planning
Bd., 14, 1944: 27 (Florida); Merriman and Warfel, Trans. N. Amer. Wild Life Conf., 1944: 234
(season, east. Long Island Sd.) ; Templeman, Res. Bull. Dep. Nat. Resources Newfoundland, 15, 1944: I
(life hist., season, migrations, Newfoundland, Labrador) ; Bigelow and Schroeder, Guide Comm. Shark
Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 140, fig. 52 (descr., iU., habits, range).

47 o Memoir Sears Foundation for Marine Research

Piked Dog, Brookes, Nat. Hist., 3, 1763: 32, pi., fig. 32 (general); 1772: 18, pi., fig. 18; Pennant, Brit.
ZooL, 3, 1769: 77 (descr., weight); Yarrell, Brit. Fish., 2, 1836: 400 (abund., habits, food, defensive
use of spines, Gt. Brit., general) ; 2nd ed., 2, 1841 : 524; Couch, Hist. Brit. Fish., i, I 867: 49, pi. 1 1
(size, abund., record catch of 20,000, nuisance to fishermen, Gt. Brit.).

Spinax, Duhamel, Traite Gen. Pcches, (2) 4 (9), 1769: 299, pi. 20, fig. 5, 6 (general; ill. as "Aiguillat").

L'Aiguillat, Broussonet, Mem. Acad. Sci. Paris, 1780: 673 (abund., Medit., Atlant. to Scotland).

Squalu! sphtax Olavius, Oecon. Reyse Island, 1780: 562, pi. 7 (Iceland, not seen).

Squale aiguillat, Lacepede, Hist. Nat. Poiss., 4° ed., i, 1798: 270; in Sonnini, Hist. Nat. Poiss., 4, 1802: 100
(descr.).

Squalus achantias Latreille, Nouv. Diet. Hist. Nat., 24, 1 804: 72 (evident misspelling for "acanthias" in table
of contents).

Acanthorhinus acanthias Blainville, Bull. Soc. philom. Paris, 18 16: 121 (name only).

Sfinax acanthias Cloquet, Diet. Sci. Nat., /, SuppL, 1 8 16:^^ 91 (abund., defensive spines, fishery, N. Europe) ;
Cuvier, Rcgne Anim., 2, 1817: 130 (diagn., abund.); Bory de St. Vincent, Diet. Class. Hist. Nat., 15,
1829: 598; Bonaparte, Icon. Faun. Ital., 5 (2), 1834: plate not numbered (descr., ill., Italy); Agassiz,
L., Poiss. Foss., J, 1837: pi. B, fig. 3 (spines) ;"* Templeton, Charlesworth Mag. nat. Hist., (2) /, 1837:
413 (Ireland); Parnell, Mem. Werner. Soc. Edinb., 7, 1838: 420 (abund., food, habits, Gt. Brit.);
Swainson, Nat. Hist. Fish. Amphib. Kept., 2, 1839: 315 (general) ; Storer, Rep. Fishes Rept. Birds Mass.,
1839: 187 (abund., season, fishery, Massachusetts); Boston J. nat. Hist., 2, 1839: 538; Nordman, in
Demidoff, Voy. Russ. Merid., 5, 1840: 548 (common. Black Sea) ; Ayres, Boston J. nat. Hist., 4, 1S42:
288 (near N. York) ; DeKay, Zool. N. Y., 4, 1842: 359, pi. 64, fig. 210 (descr., ill., abund., season, fish-
ery, C. Cod); Selys-Longchamps, Fauna Belg., I, 1842: 233 (Belgium); Cuvier, Regne Anim., ill. ed.,
1 843 : 366, pi. 115, fig. 4 (food, ill.) ; Linsley, Amer. J. Sci., 4y, 1 844: 77 (Connecticut) ; Gosse, Nat.
Hist. Fish., 1 851: 308 (abund., Brit, waters); Desvaux, Essai Ichthyol. France, 1851: 19 (not seen);
Owen, Cat. Osteol. Roy. Coll. Surg., r, 1858: 91 (vert., jaws) ; Van der Hoeven, Handb. Dierkunde, 2,
1855: 261 (general) ; Thompson, W., Nat. Hist. Ireland, 4, 1856: 254 (Ireland) ; DeKay, Rep. St. Cab.
nat. Hist. N. Y. (1855), 8, i8;8: 64 (N. York); Doumet-Adanson, Rev. Zool., (2) 12, i860: 446
(Medit.) ; Schlegel, Naturl. Hist. Nederl. Dieren, 1862: 192, pi. 18, fig. 3 (descr., Netherlands) ; Gill,
Proc. Acad. nat. Sci. Philad., 1863: 333 (Massachusetts) ; Knight, Descr. Cat. Fish. N. S., 1866: 8 (Nova
Scotia); de la Blanchere, Diet. Peches, 1868: II (descr., breeding season); Van Beneden, Poiss. Cotes
Belg., 1870: 9; Mem. Acad. R. Belg. (1870), 1871: 9 (food, parasites, Belgium); Saxby, Zoologist,
1871: 2554 (Newfoundland); Reguis, Ess. Hist. Nat. Provence, i (l), 1877: 54 (Medit.); Cocco,
Nat. Sicil., 5, 1884: 176 (Messina, not seen) ; Grondal, Islenzka Fisk. (Pisces Island) Reykj., 1890-1891
(Iceland, not seen).

Acanthias vulgaris Risso, Hist. Nat. Europe Merid., j, 1826: 131 (Medit.); Miiller and Henle, Plagiost.,
1 841: 83 (descr., Medit., North Sea, Atlant.); Hamilton, Brit. Fish., 2, 1843: 313 (Gt. Brit.), also
later eds. ; Bonaparte, Cat. Pesc. Europ., 1846: 15; Nardo, Sinon. Modern. Spec, descr. Pesci St. Chiere-
ghini, 1847: III (name only); Gray, List Fish. Brit. Mus., 1 851: 70 (in part); White, List Spec. Brit.
Mus., Fish., S, 1851: 131; Baikie, Zoologist, 11, 1 853: 3846 (Orkneys); Kr0yer, Danmarks Fisk., 5,
1853: 12, 868 (Denmark) ; Nilsson, Skand. Fauna, 4, Fisk, 1855: 731 (Scand.) ; Machado, Feces Cadiz.,
1859: 8 (off Cadiz) ; Jouan, Mem. Soc. Imp. Sci. nat. Cherbourg, 7, 1859 (not seen) ; Kessler, Bull. Soc.
Nat. Moscou, 52 (4), 1859: 473 (abund.. Black Sea); Nardo, Atti 1st. veneto, (3) 5, 1859-1860: 787
(Medit., near Venice); Malmgren, Arch. Naturg., 50 (l), 1864: 346 (N. Norway, west of North
Cape) ; Fitzinger, Bild. Atlas Naturg. Fische, 1864: fig. 177 (ill.) ; Dumeril, Hist. Nat. Poiss., i, I 865:
437 (refs., descr., general); Bocage and Brito Capello, Poiss. Plagiost. Portugal, 1866: 21 (Portugal);
Giinther, Trans, zool. Soc. Lond., d, 1869: 396 (mention, but report for Cent. Amer. perhaps not this
species) ; Ninni, Ann. Soc. nat. Modena, 5, 1870: 66 (near Venice) ; Brito Capello, J. Sci. math. phys. nat.
Lisboa ( 1 868-1 869), 2, 1870: 143 (Portugal); Gunther, Cat. Fish. Brit. Mus., S, 1 870: 418 (refs.,

53. This date has frequently been quoted as 1821, but the copy in the library of the Harvard Museum of Compara-
tive Zoology is dated i8i6.

54. This plate, including other species also, is labeled simply sfinax. But fig. 3 is identified as S. acanthias by the
Explic. des Planches which appeared in 1843.

Fishes of the Western North Atlantic 471

descr., includes south, hemisphere records, probably not this species) ; Miklucho-Maclay, Beitr.Vergl. Neu-
rol. Wirbelt., i, 1 870: 10, fig. 2, A-C (bmin) ; Saxby, Zoologist, (2) 6, 1 871: 2554 (Newfoundland);
Canestrini, in Cornalia, et al., Faun, d'ltal., 5, Pesc, 1872: 39 (Medit.) ; Hertwig, Jena Z. Naturw.,
8, 1874: 349, pi. 12, fig. 4, 8, II, 12, 14, pi. 13, fig. 10, 13, 16 (develop, of dermal denticles, teeth);
Mcintosh, Ann. Mag. nat. Hist., (4) 14, 1874:424 (off W. Scotland) ; Collett, Norges Fisk., 1875: 21 1
(Norway) ; Gervais and Boulart, Poiss., 5, 1877: 206, 207, pi. 79 (descr., ill., France) ; Doderlein, Prosp.
Metod. Pesci Sicil., 1878; 30 (Medit.); Feddersen, Naturh. Tidsskr., (3) 12, 1879: 70, footnote (in a
Danish stream) ; Jones, Proc. N. S. Inst. Sci., 5(1), Append., 1879: 95 (Nova Scotia) ; Winther, Prod.
Ichthyol. Dan. Marin, in Naturh. Tidsskr., I 879-1 880: 58 (Denmark) ; Bean, Amer. Nat., 14, 1880: 526
(in a stream in Denmark) ; Giinther, Guide to Study Fish., 1 880: 33 I (general) ; Stossich, Boll. Soc. adriat.
Sci. nat., 5, 1880: 70 (Adriatic) ; Day, Fish. Gt. Brit., 2, 1S80-1884: 3 15, pi. 160, fig. 2 (descr., ill., hab-
its, abund.. Iocs., food, Gt. Brit.); Doderlein, Man. Ittiol. Medit., 2, 1 881: 86 (Medit.) ;°'' Moreau, Poiss.
France, i, 1881 : 342 (descr., France) ; Perugia, Elenc. Pesc. Adriat., 1881 : 56 (Adriatic) ; Hasse, Naturl.
Syst. Elasm. besond. Theil, 1882: pi. 12, fig. 20-28 (denticles, vertebrae) ; Marion, Ann. Mus. Hist. nat.
Marseille, /, 1883: 100 (off Marseilles, not seen) ; Storm, K. norske Vidensk.-Selsk. Skr. (1883), 1884:
45 (Trondhjera Fjord) ; Graeffe, Arb. zool. Inst. Univ. Wien, 7, 1886: 3, pi. I (29), fig. I, 2 (develop.,
descr. of embryos, breeding season, abund., Trieste) ; Kolombatovic, Cat. Vert. Dalmat., 1888: 28 (abund.,
Adriatic); Carus, Prod. Fauna Medit., 2, 1889-1893: 503 (Medit.);" Bellotti, Atti Soc. ital. Sci.
nst., 5j, 1891: no (no. of embryos) ; Garstang, J. Mar. biol. Ass. U. K., N. S. j, 1894: 229 (breed-
ing season, Plymouth, England) ; Sicher, Atti Accad. gioenia, (4) 11 (5), 1898: 17 (Medit.); Jungerson,
Danish "Ingolf" Exped., 2 (l), 1899: 27, pi. I, fig. 10, II (claspers) ; Lo Bianco, Mitt. zool. Sta.
Neapel, ij, 1899: 538 (breeding season, Naples); Huber, Z. Wiss. Zool., 70, 1901: pi. 28, fig. 11
(claspers); Steinhardt, Arch. Naturgesch., (69) i, 1903: 13, pi. I, fig. 24-31 (oral and pharyngeal
denticles); Schmidt, Fiskeriunders. Island og Faer0erne (1903), 1904: 22 (not seen); Werner, Zool.
Jb., Syst. Abt., 21, 1904: 291 (Medit.) ; Borcea, Arch. Zool. exp. gen., (4) 4, 1905: 203, 205 (breeding
season, Roscoff) ; Imms, Proc. zool. Soc. Lond., i, 1905: 45 (pharyngeal denticles); Bridge, Camb. nat.
Hist., 1910: 45; (general); Giinther, Encyc. Brit., nth ed., 24, 191 1 : 804 (general); Murray and
Hjort, Depths of Ocean, 191 2: 391 (off Morocco, Faroes) ; Le Danois, Ann. Inst, oceanogr. Monaco, 5
(5), 1913: 21, 134 (English Channel); Leigh-Sharpe, J. Morph., 34, 1920: 259 (claspers); Evans,
Philos. Trans., (B) 212, 1923: 27 (poisoning by spines) ; Jenkins, Fish. Brit. Isles, 1925: 321 (general,
commercial catch, Gt. Brit.) ; Ehrenbaum, in Grimpe and Wagler, Tierwelt N.- u. Ostsee, Lief 7 (12'),
1927: 17 (descr.. North Sea, Baltic); Hickling, Rep. Brit. Ass. Adv. Sci., 1927: 331 (bathymetric segre-
gation of sexes, Faroe Shetland Channel) ; Ehrenbaum, Faune Ichthyol., Cons, explor. Mer., 1929: plate
not numbered (general) ; Borri, Atti Soc. tosc. Sci. nat., 44, 1934: 99 (Medit.) ; Ranzi, Pubbl. Staz. zool.
Napoli, /J, 1934: 343, 372 (uterine fluid and wall); Needham, Biochem. Morphogen., 1942: 41 (em-
bryonic weight).

Squalus (Acantherinus) acanthias Blainville, in Vieillot, Faune Franc, 1825: 57 (descr., France).

Squalus [Sfinax) acanthias Voigt, in Cuvier, Tierreich, 2, 1832: 510 (descr.); Richardson, Fauna Boreal.
Amer., j, 1836: 291 (Nova Scotia) ; van der Hoeven, Handb. Dierkunde, 2, 1855: 260.

Acanthias acanthias Bonaparte, Mem. Soc. neuchatel. Sci. nat., 2 (8), 1839: 8 (in synopsis) ; Poey, An. Soc.
esp. Hist. Nat., 5, 1876: 398; Enumerat. Pise. Cubens., 1876: 202, in part (discus., refs.) ; Nobre, Fauna
Marinha Port. Vert., j, 1935: 445 (descr., Portugal).

Sfinax (no spec, name) Owen, Odontogr., 1 840-1 845: 29, pi. 3, fig. 3 (teeth).

Acanthias americanus Storer, Mem. Amer. Acad. Arts. Sci., N. S. 2, 1846: 506 (N. York to Labrador and be-
yond) ; Boston J. Nat. Hist., 6, 1857: 270 (Nova Scotia, Labrador) ; Gill, Proc. Acad. nat. Sci. Philad.,
Addend., 1 861: 60 (name); Leidy, Proc. Acad. nat. Sci. Philad., 1865: 175 (Isles of Shoals, N. Hamp-
shire) ; Storer, Mem. Amer. Acad. Arts Sci., N. S. p, 1867: 232, pi. 38, fig. I ; also as Fishes Mass., 1867:
256, pi. 38, fig. I (descr., ill., abund., season, fishery for oil, Massachusetts) ; Hertwig, Jena. Z. Naturw.,
S, 1874: 349, pi. 12, fig. 5, 9, pi. 13, fig. 15 (devel. denticles, teeth).

55. See Doderlein for additional Mediterranean citations in publications not accessible to us.

56. See Carus for additional Mediterranean citations in publications not accessible to us.

472 Memoir Sears Foundation for Marine Research

Sfinax vulgaris Guichenot, Explor. Alger. Poiss., 5, 1 850: 125 (Algeria).

Squalus americanus Gill, Proc. Acad. nat. Sci. Philad., 1 864: 263 (east coast, U.S.); Abbott, Rep. N. J.
geol. Surv., App. E, 1868: 824 (common, N. Jersey).

Acanlhias linnet Malm, Goteborgs och Bohuslans Fauna, 1877: 624 (commercial catch, Sweden).

Dog Fish, Collins, Bull. U.S. Fish Comm., 2, 1883: 8 (in winter off N. Hampshire); Gallup, Bull. U.S.
Fish Comm., 2, 1883: 246 (util., Maine); Collins, Bull. U.S. Fish Comm., 4, 1884: 248 (preying on
mackerel); Stevenson, Rep. U.S. Comm. Fish. (1902), 1904: 228 (size, util.) ; Evermann, Rep. U.S.
Comm. Fish. (1903), 1905: 114 (abund., no. of embryos, destroying other fish, Gulf of Maine);
Atkins, Bull. U.S. Bur. Fish., 2S, 1910: 253 (possible util.) ; Cornish, Contr. Canad. Biol. (1906-1910),
1 91 2: 79 (abund., season, Nova Scotia, Prince Edward I.) ; Beatty, Rep. Fish. Res. Bd. Canad. (1942),
1943: 17 (season, abund.. Vitamin A content, Bay of Fundy, Nova Scotia, Gulf of St. Lawrence).

Squalus acanthias or Acanthias vulgaris Prince, Rep. Mar. Fish. Canada (1903), 1904: XLV (abund., de-
structiveness).

Horned Dog Fish, Field, Bull. U.S. Bur. Fish., 28, 1910: 247 (as a food fish).

Crayfish, Moore, Bull. U.S. Bur. Fish., Econ. Circ, 22, 1916: (as a food fish, abund.).

Squalus Nichols and LaMonte, Amer. Mus. Novit., 901, 1913: 4 (eaten by swordfish).

Squalus barbouri Howell-Rivero, Proc. Boston Soc. nat. Hist., 41, 1936: 47, pi. 12 (descr., ill., Cuba, ident.
by exam, of type spec).

Doubtful References:

Sfinox acanthias Bowditch, Excurs. Madeira, 1825: 74 (Porto Santo, nominal only, may equally have

referred to S. blainville) .
Acanthias vulgaris Vincent, Sea Fish. Trinidad, 1910: 53 (common, Trinidad, nominal only, more likely

to have referred to the newly discovered S. cubensis, p. 473).
Probably not Squalus acanthias Evermann and Bean, Rep. U.S. Comm. Fish. (1896), 1 898: 239 (abund. Indian

R., Florida) ; Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1 900: 49 (Indian R., Florida)."

2. North Pacific:

Acanthias vulgaris Bleeker, Verh. batavia. Genoot., 25 (7), 1853: 21 (name only, Japan); Verh. batavia.
Genoot., 26 (4), 1857: 44 (name only, Japan); Gunther, Cat. Fish. Brit. Mus., 8, 1870: 418;
Nystrom, Bih. Svensk. Vet. Acad. Handl., Stockholm, 13 (4), 1887: 50 (name only, Japan); Ishikawa
and Matsuura, Prel. Cat. Fish. Mus. Tokyo, 1897: 61 (name only, Japan) ; not Acanthias vulgaris Tem-
minck and Schlegel, in Siebold, et al., Fauna Japan Pisces, 4 (15), 1850: 304, pi. 135 (this is probably
jernandinus; see discussion, p. 454).

S-pinax {Acanthias) suckleyi Girard, Proc. Acad. nat. Sci. Philad., 7, 1854: 196 (descr., Puget Sound).

Acanthias sucklii Girard, Pacif. R. R. Surv. Fish., 10, 1 858: 368 (descr., color, Puget Sound).

Acanthias suckleyi Suckley, Pacif. R. R. Survey, 12 (2), i860: 367 (descr., size, embryos, abund., util., Puget
Sound).

Squalus suckleyi (or sucklii) " Gill, Proc. Acad. nat. Sci. Philad. ( 1 862) , 1 863 : 499 (refs.) ; Jordan and Gilbert,
Bull. U.S. nat. Mus., 16, 1883: 17 (Pacific coast, U.S.) ; Jordan and Starks, Proc. Calif. Acad. Sci., (2)
5, 1895: 789 (abund., Puget Sound) ; Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 219
(name only, Aleutian Is. to Santa Barbara, California) ; Bull. U.S. nat. Mus., 47 (l), 1896: 54 (descr.,
abund., util., Aleutian Is. to Santa Barbara, California) ; Jordan and Gilbert, in Jordan, et al.. Fur Seals and
Fur Seal Industr. N. Pacif. Ocean, 3, 1 899: 434, pi. 3 (Bering I.) ; Evermann and Goldsborough, Bull.
U.S. Bur. Fish., 26, 1907: 228 (SE. Alaska) ; Starks and Morris, Univ. Calif. Publ. Zool., 3, 1907: 168
(off San Diego, California, in deep water) ; Regan, Ann. Mag. nat. Hist., (8) 2, 1908: 46 (in synopsis) ;
Starks, Ann. Cameg. Mus., 7, 191 1 : 207 (name only, Puget Sound); Halkett, Check List Fish. Canad.,
1913: 41 (Brit. Columbia, Aleutians to California); Thompson, Proc. U.S. nat. Mus., 50, 1916: 420
(name only, San Diego, California) ; Hubbs, Copeia, 43, 1 91 7: 37 (stomach contents, California); Starks,
Fish Bull. Sacramento, 3, 1917: 152 (ill., abund. northward, size, depth, California to Alaska);
Jordan and Hubbs, Mem. Cameg. Mus., 10, 1925: 105 (complete list refs.; they point out that ill. by

57. See footnote 42, p. 463. 58. Spelled sucklii by some authors, suckleyi by others.

Fishes of the Wester 7i North Atlantic 473

Jordan and Fowler, 1 903, of their mitsukurii was actually suckleyi) ; Hubbs, J. Pan-Pacif. Res. Instn.,
3 (3), 1928: II (listed for Oregon, Washington); Mori, J. Pan-Pacif. Res. Instn., j (3), 1928: 3
(listed for Korea); Quigley, Biol. Bull. Wood's Hole, ^o, 1928: 439 (weight in rel. to length, per-
centages of sexes; breeding. Straits of Georgia); Fowler, Hongkong Nat., /, 1930: 88 (not seen);
Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 21 (in check list; Aleutians
to Santa Barbara, California; Kamchatka); Fang and Wang, Contr. biol. Lab. Sci. See. China, 8, 1932:
246 (dcscr., ill., Chefoo, China); Tanaka, Jap. Fish Life Colors, 22, 1933: (not seen); Daniel,
Elasmobranch Fishes, Univ. Calif. Press., 1934: 37, 154, fig. 147 (gill arches and rakers); Walford,
Fish Bull. Sacramento, ^5, 1935: 42 (ill., California); Barnhart, Mar. Fish, south. Calif., 1936: 10
(distrib., California).

Squalus acanthias Jordan and Gilbert, Proc. U.S. nat. Mus., j, 1881: 458 (listed, Puget Sound to Santa
Barbara, California); Bean, Proc. U.S. nat. Mus., 4, 1882: 261, 269, 272, 274 (records for Brit.
Columbia, Alaska); Jordan and Gilbert, Proc. U.S. nat. Mus., 4, 1882: 33 (habits, food, abund.,
breeding season) ; Bean, Proc. U.S. nat. Mus., <5, 1884: 361 (Brit. Columbia); Eigenmann, Proc. U.S.
nat. Mus., J5, 1893: 129, 132 (near San Diego, California); Bean and Bean, Proc. U.S. nat. Mus.,
/p, 1897: 237 (name only. Commander L) ; Schmidt, Pisces Mar. Orient. Rossici, 1904: 287 (descr.,
meas., discuss., E. Siberia, in Russian); Pavlenko, Kazani Turdy Obschestro E. Stest., 42 (2), 1910: II
(Peter the Great Bay; in Russian); Berg, Faune Russie Poiss., /, 1911: 71 (California to Vladivos-
tok; in Russian); Soldatov and Lindberg, Bull. Pacif. Sci. Fish. Inst. Vladivostok, 1930: 16 (Peter the
Great and Olga Bays, Japanese Sea, in Russian).

Squalus mitsukurii (in part) Jordan and Fowler, Proc. U.S. nat. Mus., 29, 1 903: 630, fig. 3 (Japan; but as
pointed out by Jordan and Hubbs in Mem. Carneg. Mus., 10, 1925: 103, the descr. was of a member
of the jernandirius group; see discuss., p. 454); Jordan, Tanaka and Snyder, J. Coll. Sci. Tokyo, 33,
1913: 18 (ill. and refs. in part).

Sgualus mitsukurii Jordan and Evermann, Bull. U.S. Bur. Fish., 25, 1 905: 45, fig. 6 (ill. only; wrongly
stated to be "from the type"); Tanaka, Annot. zool. jap., 6, 1908: 236 (Sakhalin I.); Smith, Proc.
U.S. nat. Mus., 41, 1912: 679 (Japan); Jordan and Metz, Mem. Carneg. Mus., 6, 1913: 4, fig. 2
(ill., Korea); Tanaka, Fish. Japan, 26, 1917: 471, pi. 130, fig. 368-370 (descr., ill., Japan); not
Squalus mitsukurii Jordan and Fowler, Proc. U.S. nat. Mus., 26, 1903: 629 (descr. of type spec).

Squalus suckleyi (in part) Garman, Mem. Harv. Mus. comp. Zool., 36, 191 3: 194 (descr., NE. Pacif., but
not his var. mitsukurii, for mitsukurii Jordan and Fowler, 1 903: 629, is probably one of the jernaniiinus
group, see discussion, p. 454); Fowler, Proc. Pan-Pacif. sci. Congr., 4, 1930: 495 (California, Brit.
Columbia, Japan, Formosa; but refs. for Hawaii probably fernandinus) ; Fowler, Bull. U.S. nat. Mus.,
100 (rj), 1 941: 258 (descr., but refs. only in part) ; not Squalus suckleyi Fowler, Mem. Bishop Mus., 10,
1928: 23 (Hawaiian Is., probably fernandinus).

Squalus zvakiyae Tanaka, Fish. Japan, 27, 1918: 475 (to replace mitsukurii Tanaka, 1917, because the latter
is preoccupied by Jordan and Fowler, 1 903, 629).

Squalus sucklii suhsp. mitsukurii Schmidt, Trans. Pacif. Comm., Leningr., 2, 1931: 7 (doubts if Japanese and
Black Sea specs, are separable).

Squalus cubensis Howell-Rivero, 1936

Cuban Dogfish

Figures 89, 90

Study Material. Male, 524 mm. (Harv. Mus. Comp. Zool., type. No. 1458); fe-
male, 672 mm. long (Harv. Mus. Comp. Zool., No. 1461)5 very young male and female,
each about 280 mm., with umbilical scars still evident, and male embryo about 203 mm.
long, with yolk sac attached, all from Havana, Cuba (Harv. Mus. Comp. Zool., No.
1459, 1460, 1462).

474

Memoir Sears Foundation for Marine Research

Distinctive Characters. S. cubensis differs most sharply from acanthias and blainville
in having the inner corner of its pectoral acutely angular (broadly rounded in the other two
species) } it differs further from acanthias in that its first dorsal originates only a little
posterior to the axil of its pectoral (over or behind the inner corner of the pectoral in
acanthias) , the midpoint of its pelvics are only very little nearer to its second dorsal than

Figure 89. Squalus cubensis, female, about 672 mm. long, from near Havana, Cuba (Harv. Mus. Comp.
Zool., No. 1461). A Anterior part of head from below, Yz natural size. B Left-hand nostril, about 4 x.
C Second to fourth upper and lower teeth, about 8.5 x.

to its first (much nearer to the second dorsal than to the first in acanthias)., it has longer
fin spines and its coloration is plain; it differs further from blainville in having the distal
margin of its pectoral deeply concave.

Description. Proportional dimensions in per cent of total length. Male, 524 mm.,
from Havana, Cuba (Harv. Mus. Comp. Zool., holotype. No. 1458). Female, 672 mm.,
same locality (Harv. Mus. Comp. Zool., paratype. No. 1461).

Trunk at origin of fectoral: breadth 12.4, 12.9; height 9.4, 9.5.

Snout length in front of: outer nostrils 3.0, 3.7; mouth 9.5, 9.5.

Eye: horizontal diameter 4.8, 4.8.

Mo«/A; breadth 6.7, 7.I} height 1.3, 1.2.

Nostrils: distance between inner ends 4.0, 4.6.

Labial furrow length from angle of mouth: upper 2.7, 2.2; lower 1.5, 1.6.

Fishes of the Western North Atlantic

475

Gill o-pening lengths: ist 1.3, 1.9; 2nd i.i, 1.6; 3rd i.i, 1.6; 4th 1.2, 2.0; 5th

1.9,2.4.

First dorsal fin: vertical height 7.8, 8.0; length of base 8.6, 9.2.

Second dorsal fin: vertical height 5.3, 4.9; length of base 6.9, 6.0.

CflW<?/ ^»; uppermargin 21.8, 21.4; lower anterior margin 12.0, 12.0.

Pectoral fin: outer mzr gin 14.7, 15.9; inner margin 1 1.2, 11.6; distal margin 12.4,

10.4.

Distance from snout to: ist dorsal 29.6, 27.2; 2nd dorsal 61.3, 64. i ; upper caudal

78.2, 78.6; pectoral 21.7, 21.4; pelvics 46.0, 46.8.

Interspace between: ist and 2nd dorsals 24.0, 27.5; 2nd dorsal and caudal 10.7,

9-8.

Distance from origin to origin of: pectoral and pelvics 25.3, 27. i ; pelvics and cau-
dal 31.3, 30.0.

Trunk slender, as in other members of the genus, its height at ist dorsal only about
^/7 its length to origin of caudal. Body sector only a little longer than tail sector (much
longer than tail sector in acanthias), without mid-dorsal ridge. Caudal peduncle strongly
flattened below, with an obscure longitudinal ridge low down on each side (much as in
acanthias). Upper precaudal pit strongly marked, but no lower pit. Dermal denticles close-

FiGURE go. A Dermal denticles of Squalus cubensis, illustrated in Fig. 89, about 80 x. B Dermal denticles of
adult female, from Cuba (Harv. Mus. Corap. Zool., No. 1458), about 70 x. C Embryo, about 203 mm. long,
from near Havana, Cuba (Harv. Mus. Comp. Zool., No. 1459), about 0.6 x.

476 Memoir Sears Foundation for Marine Research

spaced, but not overlapping, varying considerably in shape on different parts of the body
and at different agesj in the adult, those on upper sides along midsector of trunk lanceo-
late, sharp-tipped, with very high median crest dividing anteriorly into 2 or 3 definite
ridges, and with a broad wing-like expansion either side opposite the median axis of the
pedicel ; farther forward, on the trunk, and also rearward and downward, these give place
to flatter forms without lateral wings, those on top of snout being ovoid, those on anterior
margins of pectoral and dorsal fins either ovoid or lanceolate, with the primary median
ridge dividing anteriorly into 2 or 3; those on sides of caudal peduncle either evenly
ovoid or weakly tridentate, with 3 separate longitudinal ridges; those on lower surface
generally lanceolate, the median ridge either simple or dividing into 2 anteriorly.

Head a little more than V4 (about 28%) of trunk to origin of caudal, its dorsal out-
line weakly convex to eye, but flat or slightly concave thence forward. Snout thicker at tip,
more broadly ovate and relatively shorter than in acanthias, its length in front of nostrils
about Vs its length in front of mouth, and the length in front of mouth only a little more
than V3 (about 39 to 40%) of length of head. Eye oval, its outline more convex below
than above, its horizontal diameter about V2 as long as snout in front of mouth (only about
Vs that long in acanthias) or about as long as distance between nostrils. Spiracle about V4
as long as horizontal diameter of eye, about on a level with upper margin of eye and behind
latter by a distance about Vs as long as horizontal diameter of eye. Gill openings low down
on the sides, the 5th slightly the longest, about V2 as long as horizontal diameter of eye
and 1.3 times as long as ist. Nostril very slightly oblique, a little more than V2 as long as
horizontal diameter of eye, its inner end nearer to tip of snout than to mouth by a distance
% to % as long as horizontal diameter of eye, its inner margin expanded as a broad, sub-
triangular lobe, which may or may not have a small subsidiary lobe."' Mouth only very
slightly arched, occupying about % of breadth of head. Upper labial furrow about V2 as
long as diameter of eye and running obliquely forward, the lower furrow less than %
as long as upper.

Teeth }|^Er|) so closely resembling those of acanthias that the illustration (Fig.
89 C) is sufficient, i or 2 series functional, depending on the stage in the process of replace-
ment.

First dorsal somewhat less sloping than in acanthias, its extreme length from origin
to rear tip about twice its vertical height (about 2.3 times in acanthias), its vertical height
about Vs as great as length of head, its origin a little anterior to midpoint of inner margins
of pectorals, its spine reaching nearly to apex in male and apparently in female also, its an-
terior margin weakly convex, posterior margin only slightly concave, its apex broadly
rounded, the free rear corner about as long as base or a little shorter, the midpoint of base
only about ^z as fatirom axil of pectoral as from origin of pelvics. Interspace between ist
and anddorsHsaboutaslong as from tip of snout to axil of pectoral. Second dorsal about

59. In the adult male specimen (the type), the left-hand nostril lacks this lobe, but it is present on both nostrils in
the female specirhen~(Fig. 89 B).

Fishes of the Western North Atlantic 477

% as long at base as ist and about % to % as high vertically, its posterior margin much
more deeply concave, free rear corner about as long as base, the spine extending nearly or
quite to apex in female as well as in male. Interspace between 2nd dorsal and caudal about
twice as long as base of 2nd dorsal. Caudal about Vs of total length (as in acanMas), its tip
perhaps averaging a little narrower than in acanthias, its lower lobe a little less than %
(about 60%) as long as upper, the general posterior contour, as included by the 2 lobes,
much as in acanthias (cf. Fig. 87 with 89). Pelvics about as long at base as 2nd dorsal, the
posterior margins more nearly straight and posterior corners perhaps averaging a little less
slenderly pointed than in acanthias, the midpoint of base in female nearer to origin of 2nd
dorsal than to rear end of base of ist dorsal by a distance only about V2 as long as horizontal
diameter of eye and about midway in male (much nearer 2nd dorsal than ist in both sexes
of acanthias). Claspers of adult male not subdivided at tip (subdivided in acanthias, p.
459). Pectoral a little more than % (69 to 70%) as long as head (about % in acanthias
or even a little more), its outer margin rather strongly convex toward apex, distal mar-
gin deeply and evenly concave (more deeply so than in acanthias) , the apex rounded, but
inner corner considerably produced and acutely angular, this being the most distinctive
feature of the species.

Color. Dark gray above, paler gray below, the young paler than adult; described,
when fresh-caught, as having upper lobes of both dorsals black, the caudal, pelvics and
pectorals edged with white and the iris green; after preservation the fin markings are much
stronger in the young than in the adults.

Size. The only available information is that the male listed above has large claspers,
suggesting that maturity is reached at a length not much greater than 500 mm., i.e., some-
what smaller than in the case of acanthias, although it may be as large as the latter at birth.

Developmental Stages. Probably the early stages in development (not yet described)
correspond to those in acanthias. Embryos with a yolk sac already show the bilobed nostril,
the shape of the pectoral, the relative positions of dorsal and pelvic fins and the long fin
spines characteristic of the adult.

Remarks. This Shark was first described and unmistakably pictured as long ago as
1787 by Parra; although this Dogfish is very different from the northern S. acanthias, it
was overlooked until 1936, when it was named by Howell-Rivero (see Synonyms, p. 478).

Habits. The fact that all specimens so far taken (about 20 in number) have been from
depths greater than 75 fathoms suggests that it is a deep-water species. Nothing further
is known of its habits, its breeding season or its diet.

Range. S. cubensis has been reported under this name only from the vicinity of
Havana and Matanzas, Cuba. But no doubt most of the Cuban reports of acanthias
actually refer to it,°° for specimens of all sizes, including pregnant females, are caught
commonly at mid-depths off the north coast of Cuba.'^ While none have yet been reported
from the south coast, this is probably due merely to the failure of local fishermen to

60. See p. 4.63 on this point. 6i. Personal communication by Luis Howell-Rivero.

478 Memoir Sears Foundation for Marine Research

report them, for they appear to be plentiful around Trinidad also.°^ It is to be expected
along the northeast coasts of South America generally, for the Spiny Dogfish reported by
Ribeiro for Rio de Janeiro appears to have been of this species." No information is at hand
regarding its distribution in the Gulf of Mexico, if it occurs there at all.

Synonyms and References:

Galludo, Parra, Desc. Piez. Hist. Nat. Havana, 1787: 84, pi. 34, fig. I.

Squalus spec, dubia, Poey, Synop. Pise. Cubans., Havana, 1868: 454 (spec, preserved in Havana and identified

as cubensis by Howell-Rivero).
Acanthias acanthias Poey, Enumerat. Pise. Cubens., 1876: 202; An. Soc. esp. Hist. Nat., 5, 1876: 398, in part

(Cuban spec, as above).
Squalus blainvillei Ribeiro, Ann. Mus. nac. Rio de J., 14, 1 907: 168; Fauna brasil. Peixes, 2(1) Fasc. i, 1923:

25, pi. 8 (descr. good ill.; ident. as cubensis by shape of pectoral, Rio de Janeiro).
Squalus acanthias Sanchez-Roig, Revist. Agric. Peces Cubana 9, Commerc. Trabaj., 193 1 : 18, part (Cuba,

based on Poey, 1876, not seen).
Squalus cubensis Howell-Rivero, Proc. Boston Soc. nat. Hist., 41 (4), 1936: 45, pi. 10, 11 (type descr., ill.,

adult and embryo, Cuba) ; Fowler, Fish Culturist, 21 (9), 1942: 66, fig. 9 (listed, Cuba) ; Bigclow and

Schroeder, Guide Comm. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 142, fig. 53 (descr.,

ill., range).

Probable References:

Squalus acanthias Jordan and Evermann, Bull. U.S. nat. Mus., 47 (l), 1 896: 54 (probably in part, because

reported from Cuba).
Squalus blainvillei Schreiner and Ribeiro, Ann. Mus. nac. Rio de J., 72, 1 903: 79 (name only, Rio de Janeiro,

but probably same specs, described under this name by Ribeiro, 1 907, 1923) ; Ribeiro, Pescas "Annie,"

Bol. Soc. Nat. Agric, Rio de J., 1904: 18 (near Ilha Rassa, Brazil).
Acanthias vulgaris Vincent, Sea Fish. Trinidad, 1910: 53 (common, mud banks, Trinidad).

Genus Squalus, Addendum

We include a brief account of S. jernandinus, present in southern Argentine waters,
but which seems not to range farther north than that in the western South Atlantic.

Squalus jernandinus Molina, 1782

Figure 87 E-H

Study Material. Female, 914 mm. long, from Island of Juan Fernandez (Harv.
Mus. Comp. ZooL, No. 841).

Distinctive Characters. See Description.

Description. Proportional dimensions in per cent of total length. Female, 914 mm.,
from Juan Fernandez (Harv. Mus. Comp. ZooL, No. 841).
Trunk at origin of pectoral: breadth 12.95 height 10.9.
Snout length in front of: outer nostrils 4.1 ; mouth 8.9.

62. Acanthias vulgaris reported by Vincent (Sea Fish. Trinidad, 1910: 53).

63. Reported by Ribeiro (Ann. Mus. nac. Rio de J., 14, 1907: 168; Fauna brasil. Peixes, 2 [1] Fasc. i, 1923: 15,
pi. 8) as S. blainville, but probably cubensis, its pectoral being pictured as with acute inner corner and concave
distal margin.

Fishes of the Western North Atlantic /yjc)

Eye: horizontal diameter 4.3.
Mouth: breadth 6.S--, height 0.7.
Nostrils: distance between inner ends 4.6.
Labial furrow length: upper 2.6; lower 1.9.
Gill ofening lengths: ist 1.7; 2nd 2.1 ; 3rd 2.1 ; 4th 2.1 ; 5th 2.4.
First dorsal fin: vertical height 7.7; length of base 8.1.
Second dorsal fin: vertical height 4.6; length of base 6.S-
Caudal fin: upper margin 21.2; lower anterior margin 12.6.
Pectoral fin: outer margin 17.7; inner margin lO.O; distal margin 14.3.
Distance from snout to: ist dorsal 29.8; 2nd dorsal 62.8; upper caudal 78.7; pec-
toral 20.2; pelvics 48.2.
Interspace between: ist and 2nd dorsals 26.O; 2nd dorsal and caudal 10.4.

S. fernandinus is sharply marked off from the acanthias group by the positions of the
first dorsal spine about over the midpoint of the inner margin of the pectoral (over or
posterior to the inner corner of the pectoral in acanthias^ and of the midpoint of the bases of
the pelvics about midway between the two dorsals (much nearer to the origin of the second
dorsal than to the rear end of the base of the first dorsal in acanthias) ; and by the more or
less noticeably bilobed anterior margin of the nostril (cf. Fig. 87 F with 87 C).

It falls with cubensis in the relative positions of the fins and in the contour of the
anterior margin of the nostril. But it differs very obviously from cubensis in that the distal
margin of its pectoral is only very slightly concave (deeply concave in cubensis) and its
inner corner rounded (acute in cubensis). Its teeth are indistinguishable from those of
acanthiasy but its dermal denticles differ noticeably.

S. blainville of the Mediterranean resembles !*■ so closely that it has sometimes been
considered a synonym of fernandinus ; but it appears to be distinct for the reasons given
on p. 454.

Color. Described as dark grayish or brown above, paler below, with the tips of the
dorsals more or less dusky, the caudal blackish medianly, its lobes pale yellowish gray.
Fernandinus lacks the white spots so characteristic of the acanthias group.

Range. Circumpolar and very widely distributed in boreal and cool temperate lati-
tudes of the southern hemisphere (for localities, see p. 454) ; it is also represented in the
Philippines and Japan by forms so closely allied that it is a question whether they are
separable from it (see discussion, p. 454) .

Occurrence in the Western Atlantic. S. fernandinus is listed for Argentina, from Lat.
56° S. to Lat. 35° S.,°* but the only positive record for Atlantic Argentine waters with
which we are acquainted is that of a small specimen 370 mm. long taken from the stomach
of an albatross at Lat. 34° 44' S., Long. 53° W.°*

64.. Pozzi and Bordale, Ann. Soc. cient. argent., 120, 1935: 151.

65. Lahille, An. Mus. nac. B. Aires, j^, 1928: 327; identification is made positive by the excellent illustrations that
show in particular a very long second dorsal spine.

480 Memoir Sears Foundation for Marine Research

Synonyms :°°

Squalus fernandinus Molina, Saggio Stor. Nat. Chili, 1782: 229 (Chile).

Squaltis acanthias var. fernandinus Walbaum, P. Artcdi Genera Pise. Emend. Ichthyol., 1792: 505 (Chile).

Acanthorhinus fernandinus Blainville, Bull. Soc. philom. Paris, 1816: 121 (name only).

Acanthias blaim>illii Gray, List. Fish. Brit. Mus., 1851: 70 (Cape Seas, S. Afr.) ; not Acanthias blainviiU

Risso, 1826 (see discussion, p. 454).
Spinax fernandezianus Guichcnot, in Gay, Hist. Chile, Zool., 2, 1848: 365; Perez Canto, Estud. Escual.

Chile, 1886: 10 (descr., Juan Fernandez).
Acanthias blainvillei Bleaker, Natuurk. Tijdschr. Ned. Ind., 27, i860: 50, 58, 80 (Cape of Good Hope);

not Acanthias blainville Risso, 1826 (see discussion, p. 454).
Acanthias fernandezianus Philippi, Tiburones Chile, 1887: 27, pi. 4, fig. 3 ; An. Univ. Chile, yi, 1887: 559,

pi. 4, fig. 3 (descr., ill., Juan Fernandez).
Squalus blainvillei Delfin, Rev. Chil. Hist. Nat., 4, 1900: 1 10 (Chile) ; not S. blainville Ribeiro, Fauna brasil.,

Peixes, Mus. nac. Rio de J., 2 (l) Fasc. I, 1923: 25, pi. 8 (Brazil); equals Squalus cubensis Howell-

Rivero, see p. 478.
Squalus fernandezianus Delfin, Cat. Feces Chile, igoi : 21 (refs., Juan Fernandez).
Squalus acutifinnis Regan, Ann. Natal Mus., 2, 1908: 248, pi. 37 (descr., ill., Mauritius, S. Afr.).
Squalus tasmaniensis Howell-Rivero, Occ. Pap. Boston Soc. nat. Hist., 8, 1936: 267, pi. 10, fig. a-e (descr.,
ill., Tasmania).

Doubtful Synonyms:

Squalus fhilifpnus Smith and Radcliffe, Proc. U.S. nat. Mus., 41, 1912: 677, pi. 51, fig. I (descr., ill.,

Philippines) ; not S. 'philiffinus Shaw, which is a Heterodont.
Squalus mitsukurii Jordan and Fowler, Proc. U.S. nat. Mus., 26, 1903: 630 (descr., but not the ill., fig. 3,

Japan; see discuss., footnote 20, p. 454).
Squalus jafonicush\\\\:iVi:i,VTOC. kc^A. nat. Sci. Philad., 60, 1908:71 ; Tanaka, Fish. Japan, 2(5, 1 91 7: pi. 130,

fig. 365-367 (Japan).
Squalus montalbani Whitley, Aust. Zool., 6, 1931: 310 (substitution for S. fhiliffinus Smith and Radcliffe,

1912, preoccupied by Shaw, 1804, for a Heterodont).

References for Western South Atlantic:

Squalus fernandinus Lahille, Enum. Feces Cartilag. Argent., 1921 : 16 (listed for Argentina) ; Physis B. Aires,

5, 1921 : 63 (listed for Argentina) ; Ann. Mus. nac. B. Aires, ^4, 1928: 327 (descr. and good ill. of spec.

from stomach of an albatross, Lat. 34° 44' S., Long. 53° W.) ; Pozzi and Bordale, Ann. Soc. cient.

argent., 120, 1935: 151 (listed for Argent., Lat. 35° S. to 56° S.) ; not Squalus fernandinus Fowler,

Arch. Zool. Estado Sao Paulo, J, 1941: 128 (Brazil, by ref. to S. Wj««£'i7/« Schreiner and Ribeiro, 1903,

which appears to have been cubensis; see discussion, p. 478).

Probable Reference:

Squalus blainvillei Lahille, Enum. Feces Cartilag. Argent., 1 921 : 16 (listed for Argentina).

Genus Centroscyllium Muller and Henle, 1841

Centroscy Ilium Muller and Henle, Plagiost., 1841: 191: type species, Sfinax fabricii Reinhardt, 1825, mono-
typic. West Greenland.

66. Squalus megalofs Macleay (Proc. Linn. Soc. N. S. W., 6, 1881 : 367 and subsequent authors, Aust.) is referred
by Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 195) and by Fowler (Bull. U.S. nat. Mus., 100 [/j],
1941 : 261) to the synonymy of fernandinus, but it appears to belong to another division of the genus; see dis-
cussion, p. 454.

Fishes of the Western North Atlantic 481

Generic Synonym:

ParacentroscyHium Alcock, Ann. Mag. nat. Hist., (6) 4, 1889: 379; type species, P. ornatum Alcock, mono-
typic. Bay of Bengal.

Generic Characters. Squalidae with well developed dorsal spines, their anterior edge
sharp, posterior flattened, each side with two shallow longitudinal furrows, originat-
ing at origins of fins and lying along anterior margins of latter, their tips well ex-
posed; trunk slender, subcylindrical; caudal peduncle with or without faintly marked
longitudinal dermal ridges and without precaudal pits; teeth (most distinctive generic
character) similar in the 2 jaws, with 3 to 5 cusps, the median much the largest; snout in
front of mouth shorter than from center of mouth to origin of pectorals; dermal denticles
thorn-like on stellate bases, or granular; eyes and spiracles large; nostrils oblique, their
anterior margins expanded as triangular lobes; mouth but little arched, a voluminous tri-
angular pit at corner of mouth extending as a short labial furrow on each jaw; posterior
margins of dorsals weakly concave, if at all so, the free rear corners not slenderly elongate;
origin of ist dorsal behind axil of pectoral; 2nd dorsal at least as large as ist, its origin
over or behind bases of pel vies; caudal with only weakly defined lower lobe, its axis raised
only a little; pectoral brush-shaped, its inner corner rounded and not at all produced;
some species, at least, with dermal thickenings that are presumably luminous.

Range. North Atlantic; Falkland Islands; eastern tropical Pacific; vicinity of Ha-
waiian Islands; Japan; Indian Ocean, including Bay of Bengal and Arabian Gulf.

Species. The few species so far known are deep-water forms from widely distributed
localities which closely resemble one another in general appearance and in their black or
very dark brown coloration. While closely allied, they appear sufficiently differentiated
from each other to deserve separate names.

Key to Species'

la. First dorsal spine over or in front of tips of pectorals when latter are laid back.

2a. Tip of caudal truncate, with definite subterminal notch, nigrum Garman, 1899.

Eastern Pacific off Panama;
near Galapagos Islands; near
Cocos Island and off Ha-
waiian Islands.^

2b. Tip of caudal pointed, without definite subterminal notch.

* ornatum Klcock, 1889.

Bay of Bengal and Arabian
Gulf.»

1. Species marked * have not been seen by us.

2. For list of references and localities of captures, see Fowler (Bull. U.S. nat. Mus., 100 [/j], 194.1 : 252) and Beebe
and Tee-Van (Zoologica, N. Y., 36, 1941: 120).

3. It is probable that Burckhardt's (Ann. Mag. nat. Hist., [7] 6, 1900: 567, fig. 7) figure more nearly represents
the normal shape of the caudal than does Alcock's (111. Zool. "Investigator," 1894: pi. 8, fig. 3) original illus-
tration of it. For list of references, see Fowler (Bull. U.S. nat. Mus., 100 [z^], 1941 : 254).

482 Memoir Sears Foundation for Marine Research

lb. First dorsal spine behind tips of pectorals when latter are laid back.

3a. Interspace between 2nd dorsal and caudal as long as from eye to origin of pectoral,
or longer.

4a. Second dorsal spine over midbase of pelvics, and extending considerably
beyond apex of 2nd dorsal fin; interspace between 2nd dorsal and caudal
about as long as from eye to axil of pectoral. ^granulosum Gunther, 1 8 80.

Falkland Islands.*

4b. Second dorsal spine posterior to rear ends of bases of pelvics, and extending
only V2 to % the distance toward apex of 2nd dorsal; interspace between
2nd dorsal and caudal only about as long as from eye to origin of pectoral.

ritteri Jordan and Fowler, 1903.

Japan.

3b. Interspace between 2nd dorsal and caudal only about as long as from eye to ist
gill opening. fabrkii Reinhardt, 1825, p. 482.

Centroscyllium fabricii (Reinhardt), 1825

Black Dogfish

Figure 91

Study Material. Two females, 640 and 727 mm. long, one from the continental slope
off Browns Bank, the other from off Nova Scotia in 300 fathoms (Harv. Mus. Comp.
Zool., No. 35702 and 743).

Distinctive Characters. Among West Atlantic members of its family this species falls
with Squalus acanthiasy S. cubensis and Etmofterus hillianus in its long and very prominent
fin spines. But it is easily separable from the first two by its teeth, which have several cusps,
by its dermal denticles, and by its color. It differs most obviously from Etmofterus hil-
lianus in that its lower teeth have several cusps like the uppers, and that its second dorsal
originates over the bases of the pelvics.

Description. Proportional dimensions in per cent of total length. Female 640 mm.,
from 240 miles ESE. of Boston Lightship (Harv. Mus. Comp. Zool., No. 35702). Fe-
male, 727 mm., from off Nova Scotia (Harv. Mus. Comp. Zool., No. 743).

Trunk at origin of fectoral: breadth 1 1.8, 1 1.8 ; height 7.8, 8.2.

Snout length in front of: outer nostrils 1.4, 1.8 ; mouth 7.5, 7.4.

Eye: horizontal diameter 4.7, 4.3.

Mouth: breadth 8.1, 8.2; height 3.3, 3.7.

Nostrils: distance between inner ends 3.9, 4.2.

Labial furrow length from angle of mouth: upper 1.9, 2.2; lower 1.9, 1.9.

Gill opening lengths: ist 3.1, 3.O; 2nd 3.1, 3.0; 3rd 3.1, 3.0; 4th 3.1, 3.0; 5th

2.7, 2.6.

4. Diagnosis based on Burckhardt's (Ann. Mag. nat. Hist., [7] 6, 1900: 567, fig. 8) illustration of the type speci-
men.

Fishes of the Western North Atlantic

483

First dorsal fin: vertical height 3.9, 4.O5 length of base 9.8, 12. i.

Second dorsal fin: vertical height 5.3, S-S'y length of base 1 1.2, i i.o.

Caudal fin: upper margin 24.2, 20.9; lower anterior margin 13. i, 1 1.8.

Pectoral fin: extreme length 9.7, 10.6 j extreme breadth 5.5, 5.8.

Distance from snout to: ist dorsal 30.5, 27.2; 2nd dorsal 58.6, 58.O; upper caudal

75.8, 79.1; pectoral 18.8, 21.3; pelvics 54.7, 55.3.

Interspace between: ist and 2nd dorsals 1 8.0, 19.5; 2nd dorsal and caudal 7.8, 8.8.

Distance from origin to origin of: pectoral and pelvics 35.2, 35.7; pelvics and

caudal 18.3, 22.0.

J

Figure 91. Centroscyllium jabricii, female, 640 mm. long, from 240 miles ESE. of Boston Lightship (Harv.
Mus. Comp. Zool., No. 35702). A Head from below, about 0.3 x natural size. B First to fifth upper teeth,
right-hand side. C Twentieth to twenty-second upper teeth. D First to fifth lower teeth. E Sixteenth to
eighteenth lower teeth. F Twenty-sixth lower tooth. B-F, about 5.4 x. G Dermal denticles, about 10 x.
H Lateral and apical oblique views of denticle, about 20 x.

Trunk slender, its height at ist dorsal only a little more than Ve its length to origin
of caudal. Body sector to cloaca about 1.8 times as long as tail sector, without mid-dorsal
ridge. Caudal peduncle without longitudinal ridges or precaudal pits either above or below.
Dermal denticles minute and very widely spaced, leaving most of the skin bare, on irregu-
larly stellate bases, thorn-like and more or less recurved on the trunk generally, somewhat
widest on the tail, those on top of head conical. Skin along upper sides and on top of head

484 Memoir Sears Foundation for Marine Research

with widely scattered and deeply pigmented epidermal thickenings or dots, presumably
luminous (p. 485).

Head between V3 and ^/4 (about 28%) of length to origin of caudal, its dorsal
profile weakly and evenly convex. Snout thick, fleshy, broadly-ovate, its length in front of
nostrils only about Yi of length in front of mouth, the distance between nostrils about Yo
as long as snout in front of mouth, and length in front of mouth a little more than Ys
(about 35 to 36%) of length of head. Eye oval, its lower outline the more convex, its
horizontal diameter about % as long as snout in front of mouth or a little longer than
distance between nostrils. Spiracle about Vs to Y-i as long as horizontal diameter of eye,
behind latter by a distance V3 to V2 as long as diameter of eye, the lower edge a little above
center of eye. Nostril near anterior contour of snout, moderately oblique and about Y2
as long as diameter of eye, its anterior margin produced as a well developed, narrow-tri-
angular lobe, its inner corner about midway between mouth and tip of snout. Mouth about
% as high as broad, nearly evenly arcuate. Pit at corner of mouth extending in deep fur-
rows about V4 of the distance toward symphysis on upper jaw and nearly as far on lower,
but only a very short distance rearward.

Teeth about ■~5j2=li in specimen examined (counting difficult), the uppers and lowers
similar, with 3 (sometimes 4 or 5) sharp cusps, the median much the longest; 2 to 3
series functional in each jaw.

First dorsal brush-shaped, noticeably small, its vertical height only Ve to Yl as great
as length of head, its anterior margin very sloping, apex very broadly rounded, posterior
margin nearly straight, its free rear corner only a little more than V2 as long as base, its
origin a little posterior to tips of pectorals (when latter are laid back), the ist dorsal spine
a little more than Va as far from axils of pectorals as from origin of pelvics, with the
exposed portion about Y2 as long as horizontal diameter of eye and reaching about Vs the
way along anterior margin of fin. Second dorsal about 1.2 times as long at base as ist and
about twice as large in area, its general contour subrectangular (rounded in ist dorsal),
with nearly straight anterior margin, slightly concave posterior margin and very narrowly
rounded apex, its free rear corner a little less than Y2 as long as base and only a little more
acute than a right angle, its spine a little anterior to midpoint of bases of pelvics, free for
about V2 its length, and reaching about midway along anterior margin of fin. Interspace be-
tween 2nd dorsal and caudal only about % as long as base of 2nd dorsal. Caudal a little less
than V4 of total length, its upper margin convex, apex broadly truncate, and lower posterior
margin with obtuse subterminal notch, thus marking off the terminal sector, its lower an-
terior corner rounded, only very slightly expanded (not a definite lobe), its anterior mar-
gin approximately Y2 as long as upper margin of fin. Pelvics about as large as 2nd dorsal,
their anterior margins slightly convex, apices broadly rounded, posterior margins nearly
straight, tips pointed, their origin a little anterior to origin of 2nd dorsal. Pectoral small,
a little less than Y2 as long as head, brush-shaped, with transversely truncate tip, rounded
corners, the inner not at all produced.

Fishes of the Western North Atlantic 485

Color. Deep chocolate brown, darkest (almost black) below and on fins generally;
inner edge of anterior part of upper eyelid densely pigmented with dark brown or blackish.

Size. Reported by fishermen as growing to about ^V-i feet, but adults average only
about 2 to 2V2 feet, the maximum length of which we find definite record being only about
33 inches (829 mm.). Females average larger than males, at least in Greenland waters.

Developmental Stages. Presumably ovoviviparous like Squalus, but the early devel-
opmental stages have not been described, although embryos have been reported repeatedly.

Habits. Captures of the Black Dogfish in West Greenland waters range from close
to the surface through the ice in winter down to 900 meters at least. Along the Nova
Scotian Banks they are most often taken at 200 to 300 fathoms and seldom shoaler than
150 fathoms, often in company with Centroscymnus coelolepis (p. 498). Although no
regular hook and line fishery is operated there at a greater depth, there is no reason to doubt
that they occur as deep in those waters as oflf Greenland (see above). Records for Icelandic
waters (perhaps the chief center of abundance) are mainly from about 500 to about 900
meters; they are recorded down to 1,100 meters oflF the Faroes, to 1,495 nieters off north-
west Africa, if the reports of the captures there are well founded (see footnote 5, p. 485).

This, in short, is a deep-water species, normally approaching the surface only in
Arctic latitudes and at the coldest, or perhaps darkest, season. The fact that a trawl haul
on the south slope of the Davis Strait ridge (in water of 3.12° C.) yielded 42 specimens,
while another just north of the ridge took none (from water of 2.47°) is suggestive evi-
dence that it is an inhabitant of Atlantic and not of truly polar waters.

Nothing is known of its life history other than that it bites the hook freely. Cephalo-
pods, pelagic crustaceans, and medusae (Atolla) have been found in its stomach, and
females containing embryos up to 124 mm. in length have been taken in West Greenland
waters in February. The fact that their skins are provided with minute, deeply pigmented
papillae, resembling the luminous organs of the brightly luminescent Isistius brasiliensis
(p. 512), suggests that C. fabricii also emits light, although it has not actually been seen to
do this so far as we know.

Relation to Man. The Black Dogfish is of no commercial value, hence most of those
caught are thrown back.

Range. Both sides of the North Atlantic, chiefly in depths greater than 150 fathoms;
Iceland (where most plentiful); Faroe-Shetland Channel and Faroe Bank in the east;
also reported from Arguin Bank oflF Cape Blanco, Northwest Africa;* Davis Strait; W^est
Greenland slopes and outer parts of the offshore fishing grounds in the west, from the
Grand Banks of Newfoundland to Georges Bank.

Occurrence in the Western Atlantic. To the north the Black Dogfish is at least toler-
ably common in southwestern Greenland waters, both in the fjords and on the offshore
fishing banks; in the southern part of Davis Strait, and northward to the ridge that con-

5. Doubt has been expressed as to the actual identity of the specimen from this locality reported by Vaillant (Exped.
Sci. "TravaiUeur" and "Talisman," Poiss., 1888: 72), since it was not only very small (175 mm.) but in a bad
state of preservation.

486 Memoir Sears Foundation for Marine Research

nects Greenland with Baffin Land. But it has not been found on the northern (polar) slope
of the ridge.

It is also taken often by halibut fishermen on long lines all along the edge of the North
American continent from the south slope of the Grand Bank southward along the Nova
Scotian Banks (Banquereau, Sable Island, Lahave, Browns) to the eastern extremity of
Georges Bank. But neither the total number of recorded captures nor the personal reports
we have received from fishermen suggest more than a comparatively sparse though wide-
spread population. How far to the westward and southward it may occur regularly is not
yet known. Garman" credited it with ranging as far as New York, but we find no support-
ing evidence for this, and while Goode and Bean' characterized a young specimen taken in
the northern side of the Gulf of Mexico, off the coast of Alabama, in 1885, as "probably
this species," the bottom temperature of the water there was so high (67° F., or 19.4° C.
at 210 fathoms) as to suggest that some other small shark actually was in hand.

Synonyms and References:

Squalus acanthia! Fabricius, Fauna Groenl., 1780: I 26 (W. Greenland) ; not 5. acanihias Linnaeus, I 758.

Sfinax fabricii Reinhardt, Overs, danske Vidensk-Selsk. Forh. (i 824-1 825), 1825: 3 (W. Greenland, not
seen); Overs, danske Vidensk-Selsk. Forh., 3, 1828: vi (descr., West Greenland, not seen); K. danske
Selsk. Naturvid. Math. Afhand., 7, 1838: 116, 132 (W. Greenland).

Sfinax acanthias (?) Reinhardt, K. danske Selsk. Naturvid. Math. Afhand., 7, 1838: 116 (W. Greenland,
suggests Squalus acanthias Fabricius, 1780, equals Sfinax fabricii Reinhardt, 1828).

Cenlroscyllium fabricii Muller and Henle, Flagiost., 1 841: 191 (descr., Greenland); Kr0yer, Danmarks
Fisk., 3, 1852-1853: 888 (note); Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861: 60 and later
eds. (name); Dumeril, Hist. Nat. Poiss., i, 1865: 449, pi. 5, fig. 10 (embryo reported, descr.,
ill. of scales, Greenland); Giinther, Cat. Fish. Brit. Mus., 8, 1 8 70: 425 (descr., refs., Greenland);
Goode and Bean, Bull. Essex Inst. Salem, 11, 1879: 30 (Banks off Massachusetts); Jones, List Fish.
Nova Scotia, 1879: 9 (meas.. Nova Scotia Banks); Bean, Proc. U.S. nat. Mus., j, 1 881: 1 16 (Iocs,
of capture, abund.. Grand Banks and Nova Scotia Banks) ; Hasse, Naturl. Syst. Elasm. besond. Theil,
1882: pi. 10 (vertebrae) ; Jones, Proc. N. S. Inst. Sci., 5, 1882: 95 (same as Jones, 1879) ; Jordan and
Gilbert, Bull. U.S. nat. Mus., 16, 1883: 16 (descr., off Gloucester, Massachusetts); Bean, Rep. U.S.
Comm. Fish. (1882), 1884: 344 (off Woods Hole); Gunther, "Challenger" Rep., Zool., 22, 1887:
XL, 6 (depths of capture); Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 793 (distrib.) ; Vaillant,
Exped. Sci. "Travailleur" and "Talisman," Poiss., 1888: 72 (off Arguin Bank, Morocco, Lat. about
20° 44' N.) ; Goode and Bean, Smithson. Contr. Knowl., 50, 1895; Mem. Harv. Mus. comp.
Zool., 22, 1896: II, pi. 2, fig. 7 (descr., iU., offshore banks, and perhaps Gulf of Mexico); Jordan
and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 219 (name. Nova Scotia Banks); Bull. U.S.
nat. Mus., 47 (l), 1896: 56 (descr., off Gloucester and Nova Scotia Banks) ; ibid., 47 (4), 1900: pi. 8,
fig. 26 (ill.) ; Burckhardt, Ann. Mag. nat. Hist., (7) 6, 1900: 568 (luminescent organs) ; Werner, Zool.
Jb., Syst. Abt., 21, 1904: 291 (color, embryo, Greenland); Collett, Rep. Norweg. Fish. Invest., 2 (2),
1905: 25 (sizes, embryos, off the Faroes) ; Fowler, Proc. Acad. nat. Sci. Philad., do, 1908: 69 (Georges
Bank) ; Kendall, Occ. Pap. Boston Soc. nat. Hist., 7(8), 1 908: 9 (off Gloucester, Massachusetts) ; Regan,
Ann. Mag. nat. Hist., (8) 2, 1908: 40 (taxonomic discus., cf. with other species); Murray and Hjort,
Depths of Ocean, 1912: 392 (Faroe-Shedand Channel and Faroe Bank); Garman, Mem. Harv. Mus.
comp. Zool., 56, 1913: 231, pi. 10, fig. 5-8 (descr., iU., Greenland to N. York); Halkett, Check
List. Fish. Canad., 1913: 41 (off Nova Scotia); Jensen, Mindskr. Jap. Steenstrup, 2 (30), 1914: 4
(discuss., history, size, embryos, food, depth, south. Davis Strait and SW. Greenland) ; Fowler, Copeia,

6. Mem. Harv. Mus. comp. Zool., 36, 1913: 232.

7. Smithson. Contr. Knowl., 30, 1895 : 1 1 j Albatross Station 2377.

Fishes of the Western North Atlantic 487

30, 1916: 36 (in list); Saemundsson, Vidensk. Medd. naturh. Foren. Kbh., 7^, 1922: 167 (depth,
occur. Iceland) ; Bigelow and Welsh, Bull. U.S. Bur. Fish., 40 (i), 1925: 52, 551 (diagn., ill., general,
in W. Atlant.) ; Jensen, Rapp. Cons, explor. Mer., 59, 1926: 98, loi (south side of Davis Strait ridge);
Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 25 (general); Jordan, Evermann and Clark, Rep.
U.S. Comm. Fish. (1928), 2, 1930: 21 (listed, Greenland seas, Gloucester, Nova Scotia Banks) ; MacCoy,
Bull. Boston Soc. nat. Hist., 69, 1933: 8 (240 miles ESE. from Boston Lightship) ; Bigelow and Schroe-
der, Canad. Atlant. Fauna, biol. Bd. Canad., 12^, 1934: 18 (general); Vladykov and McKenzie, Proc.
N. S. Inst. Sci., rp, 193;: 49 (off Nova Scotia) ; Fowler, Bull. Amer. Mus. nat. Hist., yo (i), 1936: 83
(descr., ill. of Georges Bank spec, discuss, of ident. of Morocco spec. rep. by Vaillant, 1888) ; Lubbert
and Ehrenbaum, Handb. Seefisch. Nordeurop., 2, 1936: 287 (Iceland).
Not Centroscyllium fahricii Lahille, Enum. Feces Cartilag. Argent., 1 921: 16 (Falkland Is.); Pozzi and
Bordale, An. Soc. cient. argent., 120, 1935: 10 (Argentina, Lat. 52° S.) ; these doubtless refer to the
type specimen of C. granulosum Giinther, 1 880.

Genus £/wo^;(?rMj Rafinesque, 18 10

Etmopterus Rafinesque, Carratt. Gen. Nuov. Sicil., 1810: 14; type species, E. aculeatus Rafinesque, 1810,
equals Squalus sfinax Linnaeus, 1758.

Generic Synonyms:

Ac<mthorhinus (in part) Blainville, Bull. Soc. philom. Paris, 1 8 16: 121.

Sfinax Cuvier, Regne Anim., 2, 1817: 129; type species, Squalus sfintue Linnaeus, 1758.

Acanthias (in part) Risso, Hist. nat. Europ. Merid., j, 1826: 132.

Centring (in part) Lowe, Proc. zool. Soc. Lond., 1833: 144; type species, C. nigra Lowe, equals Squalus sfinax

Linnaeus, 1758; not Centrina C\x\\tT, 1817.
Acanthiiium Lowe, Proc. zool. Soc. Lond., 1839: 91 ; type species, A. ptsillum Lowe, 1839, designated by

Goode and Bean (Smithson. Contr. Knowl., 30, 1895: lO; Mem. Harv. Mus. comp. Zool., 22, 1896:

10^).
Acanthidim Sollas, Zool. Rec, 43, 1906: 58 (obvious misprint for Acanthiiium') .

Generic Characters. Squalidae with dorsal spines largely exposed, arising at origins
of fins and lying along anterior margins of latter; trunk slender, subcylindrical, the pe-
duncle without lateral ridges or precaudal pits; snout in front of mouth is somewhat
shorter than from front of mouth to origin of pectorals; upper and lower teeth unlike, the
former with several cusps, the latter with only i cusp, deeply notched outwardly and so
oblique as to form a continuous cutting edge; dermal denticles ranging from bristle-like
to scale-like; eyes and spiracles large; anterior margin of nostrils with a long, narrow-
triangular lobe; a voluminous triangular pit at corner of mouth and a labial furrow on each
jaw; dorsals triangular, their free rear corners elongate but not very slender; origin of ist
dorsal posterior to tips of pectorals; 2nd dorsal larger than ist, its origin over or posterior
to base of pelvics; caudal truncate, with well marked subterminal notch, but without defi-
nite lower anterior lobe, its axis only very slightly raised; pectorals brush-shaped, their
inner corners rounded and not at all produced; inner edge of anterior part of upper eyelid
deeply pigmented; some of the species, perhaps aU of them, with luminous organs.^

1. We agree with Fowler (Bull. U.S. nat. Mus., 100 [r^], 1941: 246) that Acanthiiium Lowe is a synonym of
Etmofterus Rafinesque, and that the species grouped under A canthidium by Garman (Mem. Harv. Mus. comp.
Zool., 36, 1913: 215) fall properly in Deania Jordan and Snyder, 1902. See also footnote 4, p. 451.

2. For a general account of these organs, see Daniel (Elasmobranch Fishes, Univ. Calif. Press, 1934: 29).

4-88 Alemoir Sears Foundation for Marine Research

Range. Both sides of North Atlantic, Mediterranean, South Africa, Straits of Magel-
lan, southwest coast of South America, East Indies, Philippines, Japan and Hawaiian
Islands.

Fossil Teeth. Upper Cretaceous (?) and Miocene, Europe.

Species. The ten or eleven supposed species of these small, deeply pigmented deep
water sharks that have been named^ resemble one another so very closely that a drastic
reduction in the number of recognizable species is to be anticipated. But since we lack
adequate material from other ocean areas for comparison we limit the accompanying Key
to the North Atlantic representatives of the genus.

Key to North Atlantic Species

la. Interspace between rear end of bases of pelvics and origin of caudal as long as distance

from origin of pelvics to tips of pectorals, or longer. hillianiis Poey, i86r, p. 488.

lb. Interspace between rear end of bases of pelvics and origin of caudal considerably

shorter than distance from origin of pelvics to tips of pectorals.

2a. Dermal denticles bristle- or thorn-like; caudal a little longer than from tip of

snout to origin of pectorals. j/)i«^.v Linnaeus, 1758.*

Eastern Atlantic, Mediterra-
nean, South Africa.
2b. Dermal denticles scale-like 5 caudal considerably shorter than from tip of snout
to origin of pectorals. fusillus Lowe, 1839.

Eastern Atlantic, also Ja-
pan, or represented there
by a very close ally."

Etmopterus hillianus (Poey), 1861

Figures 92, 93

Study Material. Type specimen, 269 mm. long, from Cuba (Harv. Mus. Comp.
ZooL, No. 1025); males, 251 and 270 mm. long, from off St. Kitts, West Indies in 208

3. Brac/iyurus Smith and Radcliffe, 1912, Philippines; front'unaculatus Pietschmann, 1907, Japan; granulosus Gun-
ther, 1880, southwest coast of South America; /nltianus Poey, 1861, western North Atlantic, Florida region;
lucifer Jordan and Snyder, 1902, Japan, Philippines, East Indies, Natal; tnolleri Whitley, 1939, Australia; -paess-
leri Lonnberg, 1907, Straits of Magellan, Argentina; frinceps Collett, 1904, vicinity of the Faroes (almost cer-
tainly a synonym of sfhutx) ; fusillus Lowe, 1839, eastern Atlantic; villosus Gilbert, 1905, Hawaiian Islands;
and sfinax Linnaeus, 1758, eastern Atlantic, Mediterranean, South Africa.

4. Including frlnceps Collett, 1904. This was thought by Collett (Forh. Vidensk.-Selsk. Krist., 9, 1904: 3) to be
separable from sfinax because of its somewhat stouter and more thorn-like denticles; but we doubt the validity of
this supposed species, based on poorly preserved material.

5. The forms described under this name by Tanaka (Fish. Japan, 5, 1912: pi. 22; 6, 1912: 88) and as E. frontima-
culatus by Pietschmann (Anz. Akad. Wiss. Wien, 44, 1907: 395; S. B. Akad. Wiss. Wien, 117, 1908: 654, pi. 1,
fig. 2, pi. 2, fig. 2), both from Japan, agree with the East Atlantic fusillus in the form of the denticles. Whether
or not they are actually identical with the latter, as classed by Garman (Mem. Harv. Mus. comp. Zool., 36, 1913 :
228) and Fowler (liuU. U.S. nat. Mus., 100 [rj], 1941: 249), can be determined only by comparison of speci-
mens from the respective ocean areas.

Fishes of the Western North Atlantic

489

fathoms, and off northwestern Cuba at Lat. 23° 12' N., Long. 81° 23' W. in 375 fathoms
(Harv. Mus. Comp. ZooL, No. 1024, 1025)} also 16 specimens, 225 to 295 mm. long,
taken off the north central and northeastern coast of Cuba by the "Atlantis" in March and
April 1938, including a female (295 mm.) containing four embryos about 90 mm. long

.#^

Figure 92. Etmofterus hillianus, female, 292 mm. long, from oflF northwestern Cuba (Harv. Mus. Comp.
ZooL, No. 361 12). A Anterior part of head from below, about 1.5 x. B Dermal denticles, about 30 x. C Tip
of left-hand clasper of an adult male, 270 mm. long, from dorsal side (Harv. Mus. Comp. Zool., No. 36104).

^537^^

w ■■'■(■■ w ) V 4 W • f^' w^' m V-

Figure 93. Etmofterus hillianus, pictured in Fig. 92. A Upper and lower teeth at center of mouth. B Eighth
to twelfth (outermost) upper teeth, and sixteenth to eighteenth (outermost) lower teeth, about 10 x.

490 Memoir Sears Foundation for Marine Research

apparently ready for birth, and a male, 270 mm. long, with the trifid tips of the claspers
fully differentiated (Harv. Mus. Comp. Zool., No. 36 104 to 36 11 6); a specimen 214 mm.
from the offing of Chesapeake Bay, Lat. 37° 24' N., Long. 74° 17' W. in 300 fathoms
(U.S. Nat. Mus., No. 26740).

Distinctive Characters. Among northwestern Atlantic members pf the family this
species falls with Squalus acanthias, S. cubensis and Centroscyllium in its long and con-
spicuous fin spines. But it is easily distinguishable from all these by the striking dissimilarity
of the teeth in its two jaws. It has sometimes been confused with E. fusillus of the eastern
Atlantic, but it is distinguishable from the latter at a glance by the fact that the interspace
b«*een its pelvics and its caudal is at least as long as the distance from the origins of the
former to the tips of pectorals (considerably shorter than this in -pusillus and spinax). Its
relatively slender caudal further separates it from spinax.

Description. Proportional dimensions in per cent of total length. Male, 225 mm.,
from north coast of Cuba (Harv. Mus. Comp. Zool., No. 36105). Female, 230 mm., same
locality (Harv. Mus. Comp. Zool., No. 36104).

Trunk at origin of pectoral: breadth 9.8, 10.4; height 7.1, 7.8.

Snout length in front of: outer nostrils 2.7, 2.2j mouth lO.O, lO.O.

Eye: horizontal diameter S-(>i 6.1.

Mouth: breadth 7.6, 7.65 height 1.3, i.i.

Nostrils: distance between inner ends 3.5, 3.0.

Labial furrow length frotn angle of mouth: upper 1.8, 1.4; lower 1.8, 1.2.

Gill opening lengths: ist 1.3, 1.3; 2nd 1.3, 1.3; 3rd 1.3, 1.35 4th 1.3, 1.35 5th

1-3, I-3-

First dorsal fin: vertical height 3.8, 2.8 ; length of base 5.8, 5.2.

Second dorsal fin: vertical height 6.0, 5.2; length of base 8.9, 7.4.

C««i«/^«; upper margin 21.3, 21.3; lower anterior margin 10.2, 9.6.

Pectoral fin: extreme length 10.2, 9.6} extreme breadth 5.8, 4.8.

Distance from snout to: ist dorsal 34.7, 34.85 2nd dorsal 58. 0, 60.O; upper caudal

78.7, 78.7; pectoral 24.9, 23. i; pelvics 49.3, 50.O.

Interspace between: ist and 2nd dorsals 19. i, 20.95 2nd dorsal and caudal 13.3,

I3-9-

Distance from origin to origin of: pectoral and pelvics 24.0, 26.9; pelvics and

.caudal 28.8, 28.2.

Trunk subcylindrical, moderately slender, its height at ist dorsal about Ve its length
to origin of caudal. Body sector to cloaca about i .3 to i .4 times as long as tail sector, without
mid-dorsal ridge. Dermal denticles minute, close-set, similar over body as a whole, thorn-
like, slender, moderately curved, tapering, their bases more or less stellate but concealed in
the skin.

Head a little less than V3 (28 to 29%) of length to origin of caudal, flattened above.
Snout thick and fleshy at tip, its sides slightly concave at eyes, its anterior outline only

Fishes of the Western North Atlantic 491

slightly rounded, its lower surface with large mucous pores arranged in a prominent pat-
tern, its length in front of nostrils only about Ve of length in front of mouth, but length
in front of mouth only a little less than V2 length of head. Distance between nostrils aver-
aging a little less than Vs (28 to 35% ) of length in front of mouth. Eye oval, its lower out-
line much more convex than upper, its horizontal diameter about I/4 as long as head. Spi-
racle about y^ as long as eye, a little above upper margin of latter and behind it by a dis-
tance about Va as long as diameter of eye. Gill openings about evenly spaced, all of about
equal lengths and very short, about Vs to V4 as long as diameter of eye, the 5th close
in front of the pectoral. Nostril close to anterior margin of snout, about V2 as long as hori-
zontal diameter of eye, moderately oblique, its anterior margin with a long narrow lobe
near its outer end. Mouth very little arched, in somewhat sinuous contour, about % as
broad as length of snout in front of mouth. Upper and lower labial furrows each a little
less than ^/3 as long as to the respective symphysis, the rearward prolongation of pit at
corner of mouth a little more than V2 as long as horizontal diameter of eye.

Teeth HEiI in specimen illustrated; upper teeth usually with 5 (rarely 3 or 7)
cusps, the median cusp longest and the outermost pair very short, except on the outermost
2 teeth, which are much lower and lack definite cusps j lower teeth subquadrate, a little
longer than broad and with i sharp cusp so oblique that the inner margins are approxi-
mately parallel with the jaw, each overlapping the next outermost to form a continuous
cutting edge, the outermost tooth of all more broadly expanded than the others basally on
outer side; usually 3 series functional all along upper jaw, i or 2 on lower, depending on
their stage in replacement.

First dorsal with base about V4 as long as head, its margins nearly straight, its
apex rounded, its free rear corner about as long as base, its origin a little posterior to inner
corner of pectoral, its spine exposed for more than ^o its length, reaching about % the way
along the fin, the midpoint of its base about % as far from axil of pectoral as from origin
of pelvics. Second dorsal similar to ist in shape, but nearly twice as high vertically and
I V2 times as long at base, its origin a little posterior to rear ends of bases of pelvics, its spine
exposed for about % its length, reaching nearly to apex of fin (thus much longer than ist
dorsal spine, relatively). Interspace between 2nd dorsal and caudal about 1.5 to 1.8 times
as long as base of 2nd dorsal. Caudal a little less than % of total length or about as long as
from tip of snout to 3rd gill opening, transversely rounded at tip, with obtuse subterminal
notch, the terminal sector about V5 of fin, its lower anterior corner rounded, a little more
than a right angle, the lower anterior margin between V3 and V2 as long as upper margin.
Interspace between caudal and rear ends of bases of pelvics about as long as from origins
of pelvics to tips of pectorals in female and considerably longer in male. Pelvics about as
long at base as 2nd dorsal, with nearly straight edges and tapering subacute tips. Pectoral
a little less than Vs as long as head, about % as broad as long, brush-shaped with broadly
truncate tip and rounded corners, the inner not at all produced.

Color. After preservation the specimens are dark grayish or chocolate brown above,

492 Memoir Sears Foundation for Marine Research

very pale along midzone of back, with a pale spot on top of head and another above pos-
terior part of each eyej lower surface black. In some specimens the gradation from the
paler upper parts is gradual, but in others the black of the ventral surface extends in narrow
triangular zones forward above the bases of pelvics and backward onto the caudal, but
interrupted midway of the peduncle by a pale belt; the posterior portions of the dorsal
fins and caudal are pale, the tip of the latter dusky or blackish; the inner surface of the
anterior part of the upper eyelid is dark brown and densely pigmented. All the specimens
examined also show more or less clearly defined black dots sparsely scattered on the top of
head and rearward in a single row along the midline of the back to the origin of the caudal,
flanked by others in a scattered belt; also two to four lines of short, very narrow black
dashes lower down on each side, one line following the lateral line out onto the caudal.
Presumably these are luminous organs, and conditions in the closely allied E. lucijer from
Japan suggest that in life their centers are of a pearly luster.'

Size. Length at birth is a little more than 90 mm.; females mature at a little less than
300 mm., and males by the time they have reached 250 mm. (see Study Material, p. 489).
This, with the fact that the maximum length yet reported for it is 3 1 5 mm. without caudal
fin,' shows E. hillianus to be one of the smallest of the sharks.

Develof mental Stages. Development is ovoviviparous, and females have been taken
with as many as five embryos. One in our Study Material contains four young* about 80 to
85 mm. long with small yolk sac; these young already show the characters of the adult, in-
cluding the coloration; two on one side lie with heads forward, the two on other with heads
rearward.

Habits. Apparently this is strictly a deep-water species, the recorded depths of capture
ranging from 208 fathoms down to 392 fathoms. Nothing else is known of its habits. It
is not known positively whether or not it is luminous, as are one of the Japanese representa-
tives of the genus and E. spinax, although its coloration suggests that such is the case, for
the fine black dots on its back and sides (presumably indicating glandular areas) resemble
the luminous spots of other luminescent sharks."

Range. West Indian region and southern Florida to the offing of Chesapeake Bay;
probably Bermuda. This little shark is so far known only from Cuban waters, where
it is taken quite often on hook and line from deep water^" (here the "Atlantis" took
five specimens on one collecting cruise, Matanzas Bay, vicinity of Havana and off the
northwest coast) ; from near the Island of St. Kitts; from the Tortugas, Florida," and from

6. See Oshima (J. Coll. Sci. Tokyo, 27 [15], 1911: x-25) for histology of these organs in E. lucifer.

7. Longley and Hildebrand, Pap. Tortugas Lab., 24> >94': 3-

8. Not in very good condition.

9. For accounts of the luminous organs and phosphorescence of E. sfinax, see Johann (Z. wiss. Zool., 66, 1899: 158)
and Burckhardt (Ann. Mag. nat. Hist., [7] 6, 1900: 559) ; for the Japanese E. fusillus and E. frontimaculatus,
see Oshima (J. Coll. Sci. Tokyo, 27 [15], 1911) and Schmidt (Proc. Pan-Pacif. sci. Congr., [4.] 5, 1929: 461 ;
Trans. Pacif. Comm., Leningr., 2, 193 i ; 9).

10. Personal communication from Luis Hovvell-Rivero.

11. Longley and Hildebrand, Pap. Tortugas Lab., }^, 1941 : 3.

Fishes of the Western North Atlantic 493

one station in the offing of Chesapeake Bay (Lat. 37° 24' N., Long. 74° 17' W., 300 fath-
oms) ; probably also from Bermuda."

Synonyms and References:

Sfinax hillianus Poey, Memorias, 2, 1861: 340, pi. 19, fig. 13-14 (descr., teeth, Cuba) ; Repert. Fisico.-nat.

Cuba, 2, i868: 454 (Cuba) ; Regan, Ann. Mag. nat. Hist., (8) 2, 1908: 44 (St. Kitts, W. Indies, class.,

depth).
Sfinax fusillus (in part) Giinther, Cat. Fish. Brit. Mus., 8, 1870: 425 {hillianus incl. in synon.) ; Jordan

and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 219 (ref. for Cuba); Metzelaar, Trop.

Atlant. Visschen, 1919: 190, part (included in synon.); Fowler, Bull. Amer. Mus. nat. Hist., yo (i),

1936: 81 (ref. for W. Indies, ill. of W. Indian spec).
Spnax sfinax Poey, An. Soc. esp. Hist, nat., 5, 1876: 399; Enumerat. Pise. Cubens., 1876: 203 (Cuban spec).
Etmofterus fusillus Goode and Bean, Smithson. Contr. KnowL, 50, 1895: 10, pi. 2, fig. 2; Mem. Harv. Mus.

comp. Zool., 22, 1896: 10, pi. 2, fig. 2 (descr., ill., St. Kitts, W. Indies) ; Jordan and Evermann, Bull.

U.S. nat. Mus., 47 (l), 1896: 55 (descr., St. Kitts, W. Indies) ; Beebe and Tee-Van, Zoologica, N. Y.,

15, 1933: 157 (Bermuda, old record) ; not Etmofterus fusillus Lowe, 1939.
Etmofterus sfinax Carman, Mem. Harv. Mus. comp. Zool., 2^, 1899: 27 (thinks type spec, ident. with

sfinax') ; not Etmofterus sfinax L,\nr\3e\is, 1758.
Etmofterus hillianus Carman, Mem. Harv. Mus. comp. Zool., 56, 1913: 224, pi. 10, fig. 1-4 (descr., ill.,

off St. Kitts, W. Indies); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 21

(Cuba, St. Kitts) ; Howell-Rivero, Proc Boston Soc. nat. Hist., 41, 1 936: 49 (recent Cuban specs.) ; Bull.

Mus. comp. Zool. Harv., S2, 1938: 170 (tvpe spec, in Harv. Mus. Comp. Zool.) ; Longley and Hilde-

brand. Pap. Tortugas Lab., ^4, 1941: 3 (Tortugas, Florida, 357 to 392 fathoms, descr.); Fowler, Fish

Culturist, 2/ (9), 1942: 66 (listed, Cuba); Bigelow and Schroeder, Cuide Comm. Shark Fish., Anglo

Amer. Caribb. Comm., Wash., 1 945: 144, fig. 54 (descr., ill., habits, range).

Genus Centroscytnnus Bocage and Brito Capello, 1864

Centroscymnus Bocage and Brito Capello, Proc. zool. Soc. Lond., 1864: 263 ; Mem. R. Acad. Sci. Math. Phys.
Lisboa, 5 (2), 1865: 3, extra; type species, C. coelolefis Bocage and Brito Capello, 1864, Portugal.

Generic Synonym:

Centrofhorus (in part) Giinther, Cat. Fish. Brit. Mus., 8, 1870: 423; Seabra, Bull. Soc. portug. Sci. nat.,
5, 1912: 198; not Centrofhorus Muller and Henle, 1837.

Generic Characters. Squalidae with dorsal spines arising at origins of fins and lying
along anterior margins of latter, their tips either exposed or concealed; trunk subcylindri-
cal, without lateral longitudinal ridges or precaudal pits; snout in front of mouth much
shorter than from mouth to origin of pectorals; teeth unlike in the 2 jaws, the uppers
with one slender, lanceolate cusp, the lowers approximately quadrate, their outer margins
deeply notched and so oblique that the inner margins are nearly parallel to the jaw, form-
ing a continuous cutting edge; dermal denticles scale-like, closely overlapping, with flat
or concave blades, smooth or weakly ridged, their margins not toothed, on short, broad
pedicels; eyes and spiracles moderate to large; anterior margin of nostrils expanded as a
low triangular lobe only; a voluminous triangular pit at corner of mouth; labial furrow on
each jaw; origin of ist dorsal considerably posterior to tips of pectorals; origin of 2nd dor-
sal over bases of pelvics; caudal with subterminal notch, its lower anterior corner expanded

II. Reported as E. fusillus by Beebe and Tee-Van (Zoologica, N. Y., 15, 1933: 157).

4-94 Memoir Sears Foundation for Marine Research

as a weakly defined lobe, its axis raised at an angle of about 30° to 40° ; inner corner of
pectoral broadly rounded, not at all produced; luminous organs lacking.
Range. Both sides of North Atlandc; South Africa; Japan.

Key to Species*

I a. Length of snout in front of mouth considerably less than distance from eye to ist gill
opening. coelole-pis Bocage and Brito Capello, 1864, p. 494.

lb. Length of snout in front of mouth at least as great as distance from eye to ist gill open-
ing.

2a. Length of snout about as great as distance from eye to ist gill opening; ist dorsal
about as large as 2nd dorsal. juscus Gilchrist and von Bonde, 1924.

South Africa.^
2b. Length of snout definitely greater than distance from eye to ist gill opening; ist
dorsal smaller than 2nd dorsal.

3a. Tips of dorsal spines exposed; tip of 2nd dorsal extends back considerably
beyond tips of pelvics. owstoni Garman, 1906.

Japan.

3b. Tips of dorsal spines concealed by skin; tip of 2nd dorsal extends only as far
back as tips of pelvics. cryftacanthus Regan, 1 906.

Madeira.

Centroscymnus coelolefis Bocage and Brito Capello, 1864

Portuguese Shark

Figures 94, 95

Study Material. Two adult females, 1,1 17 and 1,080 mm. long, taken off Banquereau
Bank in 200 to 270 fathoms (Harv. Mus. Comp. ZooL, No. 35144? 35237)} also very
young male, about 328 mm. long, from the continental edge south of Nantucket, Lat. 39°
51' N., Long. 70° 17' W. (U.S. Nat. Mus., No. 118396).

Distinctive Characters. Among the local members of the suborder, Centroscymnus is
marked off from Squalus acanthias, S. cubensis, Centroscyllium and Etmopterus by the
following: its fin spines protrude so little from the skin that they are apt to be overlooked
(cf. Fig. 94 with 87, 89, 91, 92) ; while its teeth have only one cusp in each jaw, the lowers
and uppers are strikingly unlike. Owing to the inconspicuous nature of its spines it might
perhaps be confused with small specimens of Somniosus, which it resembles in the general

1. Centrofhorus crepidaler Bocage and Brito Capello, 1864, from Portuguese waters, was referred by Garman
(Mem. Harv. Mus. comp. Zool., 36, 1913: 207) to his new genus Centroselachus because of the pluricarinate
scales, but by Rey (Fauna Iberica Feces, /, 1928: 449) to Centroscymnus, but it falls in Scymnodon as here de-
fined, likewise Centroscymnus macracanthus Regan, 1906, from the Straits of Magellan and Argentina.

2. Known only from the type specimen.

Fishes of the Western North Atlantic 495

shapes and positions of the fins, as well as in the shape of the teeth, or with Dalatias. But
even apart from the fin spines, its overlapping, scale-like dermal denticles differ widely
in appearance from the thorns with which the skin of Somniosus is beset, and its lower
teeth differ markedly from those of Dalatias.

Figure 94. Ceniroscymnus coelole-pis, female, about 1,080 mm. long, from off Banquereau Bank, Nova
Scotia (Harv. Mus. Comp. Zool., No. 35237). A Dermal denticles, about 5 x. 5 Upper teeth from center
of jaw. C Upper teeth from side of jaw. D Outermost three rows of upper teeth. E Lower teeth from center
of jaw. F Lower teeth from side of jaw. G Outermost rows of lower teeth. B—G, about ^ x. H Front and
lateral views of upper teeth from side of jaw, about 6 x.

Figure 95. Head of Ceniroscymnus coelolefis, pic-
tured in Fig. 94, from below.

496 Memoir Seats Foundation for Marine Research

Description. Proportional dimensions in per cent of total length. Male, 328 mm.,
fromLat. 39° 51' N., Long. 70° 17' W. (U.S. Nat. Mus., No. 1 18396). Female, 1,080
mm., from Banquereau Bank (Harv. Mus. Comp. ZooL, No. 35237).

Trunkal origin of pectoral: breadth 12.2, 12.6; height 10.7, 11.2.

Snout length in front of: outer nostrils 2.4, 1.95 mouth 8.8, 6.2.

Eye: horizontal diameter 4.9, 3.4.

Mo«//^.- breadth 7.6, 8.1 j height i.i, i.i.

Nostrils: distance between inner ends 4.3, 3.1.

Labial furrow length from angle of mouth: upper 3.7, 2.4; lower 2.6, 1.4.

Gill opening lengths : ist 1.2, 1.8; 2nd 1.2, 1.7; 3rd i.r, 1.5; 4th 1.2, 1.2; 5th

1.7, 1.6.

First dorsal fin: vertical height 3.4, 3.7; length of base 5.2, 4.6.

Second dorsal fin: vertical height 3.8, 4.4; length of base 5.8, 5.7.

Caudal fin: upper margin 25.3, 20.4; lower anterior margin 15.4, 12.8.

Pectoral fin: extreme length 13.4, 13.2; extreme breadth 7.9, 6.3.

Distance from snout to: ist dorsal 34.7, 36.8; 2nd dorsal 61.3, 67.7; upper caudal

74.7, 79.6; pectoral 23.8, 18.8; pelvics 57.3, 60.7.

Interspace between: ist and 2nd dorsals 22.0, 24.2 ; 2nd dorsal and caudal 7.0, 8.0.

Distance from origin to origin of: pectoral and pelvics 35.7, 43-5; pelvics and

caudal 15.8, 17.5.

Trunk subcylindrical, moderately stout, its height at ist dorsal between ^q and Vs ( 17
to 18%) its length to origin of caudal, without mid-dorsal ridge. Caudal peduncle with-
out lateral ridges or precaudal pits. Dermal denticles very large, except on fins and lower
side of head anterior to gills, and so closely overlapping as to form a continuous armor,'
the peduncles short and stout, the blades smooth, ovoid, flat or concave, with rounded
margins posterior to level of gills, but weakly sculptured with 3 or 5 ridges further forward
on head.

Head about ^4 of trunk to origin of caudal, its dorsal profile weakly and evenly
convex, somewhat flattened anteriorly. Snout thick-tipped, very broadly ovate, notice-
ably short, its length in front of nostrils about ^/i to Vs as great as length In front of mouth,
its length in front of mouth a little less than % of length of head. Distance between nos-
trils about V2 as great as length in front of mouth. Eye oval, its outline about as
convex above as below, much smaller than in Centroscyllium, its horizontal diameter
about y2 as long as snout in front of mouth, its center about opposite front of mouth.
Spiracle about ^3 to V4 as long as diameter of eye, about level with upper margin of
latter. Gill openings much smaller than in Centroscyllium , about V2 as long as diameter of
eye, all of about equal length and evenly spaced, the 5th close in front of pectoral. Nostril

3. The edges of the denticles are so sharp that one must handle Cenlroscymnus carefully, lest one's hands be cut.
The closely overlapping denticles are one of the most distinctive features of the genus.

Fishes of the Western North Atlantic 497

moderately oblique, a little less than V2 as long as horizontal diameter of eye, its inner
margin expanded midway of its length as a short, triangular lobe with blunted tip, its inner
corner about equidistant from tip of snout and from front of mouth. Mouth only very
slightly arched, its breadth about twice as great as distance between nostrils and a little
greater than length of snout in front of mouth. Upper labial furrow extending nearly V2
of the way, the lower about V3 of the way, toward the respective symphyses. Pit at corner
of mouth very voluminous, allowing for considerable expansion when mouth is opened,
and prolonged as a narrowing furrow rearward, nearly Y> the way back toward ist gill
opening.

Teeth about ^ on specimen counted ( - also reported) 5 widely unlilce in the
two jaws; upper teeth with i slender, erect, lanceolate cusp on bifid base,* considerably
broader toward corner of jaw than toward center, their tips slightly curved outward, the
outer margins notched near corner of mouth in some cases, the successive series rather
widely spaced; lower teeth quadrate, each overlapping the next on the outer side, their
outer margins deeply notched, the i broad sharp cusp so strongly oblique that the inner
margins form a nearly continuous cutting edge parallel to the jaw; those near center of
mouth about twice as high as broad, but the 3 or 4 next to the corner of mouth successively
broader, the outermost of all widely expanded basally on outer side, with cusp but weakly
outlined; 2 or 3 series regularly functional in upper jaw, i or 2 in lower, depending on
the stage in replacement.

First dorsal noticeably small, with broadly rounded apex, its length at base only about
V4 to Vs as great as length of head, its vertical height a little less than length of base, its
rear margin nearly straight, free rear tip a little longer than base, its origin posterior to
inner corner of pectoral by a distance about twice as long as horizontal diameter of eye,
the midpoint of its base only about Y2 as far from axil of pectoral as from origin of pelvics,
its spine exposed at tip but so short as to be easily overlooked. Second dorsal similar to ist
in shape, but about 1.2 times as long at base, its origin a little posterior to midpoint of base
of pelvics, its rear margin weakly concave, its spine exposed at tip like that of ist dorsal, but
so short as to be apt to escape notice. Interspace between 2nd dorsal and caudal about i^^
times as long as base of 2nd dorsal. Caudal Y^ to ^^ (about 22 to 23%) of total length,
noticeably wide, its extreme breadth being about % its length, truncate posteriorly with
deep subterminal notch, its lower posterior contour weakly concave, the lower anterior
margin a little more tlian Y-j, as long as upper margin. Pelvics a little longer at base than
2nd dorsal, with nearly straight margins, rounded apices and pointed tips. Pectoral a little
less than -/w as long as head, a little more than ^2 as broad as long, with nearly straight
outer and distal margins and moderately rounded corners.

Color. Dark chocolate brown below as well as above.

She. The smallest recorded specimen is about nine inches long (230 mm.); adults

4.. The upper teeth resemble those of Somniosus in general appearance, but may be recognized by the fact that their
cusps do not taper uniformly from base to tip, but are lanceolate.

498 Memoir Sears Foundation for Marine Research

average 3 to 2Y2 feet in length, the largest for which actual measurements are available
being about 44 inches long (1,117 mm.) ; see Study Material, p. 494. Twelve kilograms
(about 26^/^ pounds) is the only weight of which we find record.

Developmental Stages. It is no doubt ovoviviparous. All that is known of its early
development is that gravid females have been taken with 1 3 to 16 embryos.

Habits. This is strictly a deep-water shark, as noted below. Apparently it is also a
very sluggish one, for those caught in the Portuguese deep-water fishery have been de-
scribed as falling into the boat entirely inert. But this may be the result of the change in
pressure or in temperature to which they are subjected while being hauled in. Off the
American copst, this shark occurs mostly at temperatures of 5° to 6° C; between about
4° and 10° or 11° off Portugal; and at 12° to 13° in the Mediterranean. All that is known
of its food is that an argentine {Argentina silus) was found in one, suggesting a fish diet.
Nothing is known of its breeding habits, except for the number of embryos (see Develop-
mental Stages, p. 498).

Relation to Man. This shark is considered worthless in American waters, but it has
been the object of a local deep-water fishery with long lines off Portugal in the past.

Range. Both sides of the North Atlantic, chiefly in depths greater than 200 fathoms,
and recorded down to 1,487 fathoms (2,718 meters); taken off Cape Verde, Morocco,
Azores, Madeira, Portugal, Faroe Bank, and Iceland in the east, as well as in the western
part of the Mediterranean; offing of Nantucket to slopes of the Grand Banks in the west.

Occurrence in the Western Atlantic. Positive records of this deep-water shark in the
western Atlantic are from the continental edge off Nantucket, from the deeper slopes of
Georges and the Nova Scotian Banks, and from the Grand Banks; a total of perhaps 15 to
20 specimens are recorded at depths ranging from 1 80 fathoms, which is the shoalest cap-
ture of it anywhere, down to 250 fathoms. No doubt, however, an old characterization of it
as abundant on the offshore banks at 200 fathoms^ or deeper presents its status much more
correctly than does the meager printed record, for fishermen, long-lining for halibut, take
odd specimens all over the halibut grounds in the deep gullies between the offshore banks,
usually at least one or two per trip. Since this is the only local type of fishery that is carried
on at a depth great enough to take them at all, it would not be astonishing if experimental
hook and line fishing on the still deeper slopes, down to 300 or 400 fathoms, were to yield
them as plentifully as was the case formerly off Portugal, where there is record of five or
six hauled in on a long line with 30 to 40 hooks (baited with fish) after a set of only two
hours.' As it is of no commercial value only an odd one is brought in as a curiosity.

Synonyms and References:

Centroscymnus coelolefis Bocage and Brito Capello, Proc. zool. Soc. Lond., 1864: 263, fig. 4; Diag. Famil.
Squalidae, 1864: 3; also same title in Mem. R. Acad. Lisboa, j, 1865: 3 (descr., Portugal); Poiss. Pla-
giost., 1866: 30, pi. 2, fig. 3 (Portugal, Madeira); Wright, Ann. Mag. nat. Hist., (4) 2, 1868: 426

5. Goode and Bean, Bull. Essex. Inst. Salem, 11, 1879: 30.

6. Wright, Ann. Mag. nat. Hist., (4) 2, 1868: 426.

Fishes of the Western North Atlantic 499

(Portugal, fishery); Brito Capello, J. Sci. m.ith. phys. nat. Lisboa, 4 (13), 1872: 88 (Madeira);
Goode and Bean, Bull. Essex Inst. Salem, //, 1879: 30 (abund., Banks off New England); Jones,
List Fish. Nova Scotia, 1879: 10 (meas., off Nova Scotia); Bean, Proc. U.S. nat. Mus., 3, 1881:
116 (list of Iocs., off New England, Nova Scotia, Grand Banks); Jones, Proc. N. S. Iiist. Sci., 5,
882: 96 (same as Jones, 1879); Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 17 (descr.,
Portugal, near Gloucester, Massachusetts); Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 793
(distrib.) ; Vaillant, Exped. Sci. "Travailleur" and "Talisman," Poiss., 1888: 63, pi. 2, fig. I (ill.,
weight, no. of embryos, temp., off Portugal); Bellotti, Atti Soc. ital. Sci. nat., jj, 1 891: 113
(Medit.) ; Moreau, Poiss. France, Suppl., l8gi: 9 (small no. of embryos, Nice, France); Carus, Prod.
Fauna Medit., 2, 1889— i 893: 503 (Medit.) ; Goode and Bean, Smithson. Contr. Knowl., 30, i 895: 14,
508, pi. 4, fig. 13; Mem. Harv. Mus. comp. Zool., 22, 1896: 14, 508, pi. 4, fig. 13 (descr., ill., Iocs, off
New England) ; Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 219 (off Portug.il, Massa-
chusetts, Nova Scotia); Bull. U.S. nat. Mus., 47 (1), 1S96: 55 (descr., off Portugal, Massachusetts,
Nova Scotia) ; Bull. U.S. nat. Mus., 47 (4), 1 900: pi. 8, fig. 25 (ill.) ; de Braganza, Result. Invest. Sci.
"Amelia," 2, 1904: 86, 106 (abund., depth, off Portugal); Collett, Rep. Norweg. Fish. Invest.
2 (2), 1905: 24 (size, off Faroes) ; Regan, Ann. Mag. nat. Hist., (7) 18, 1906: 437 (in Key) ; Kendall,
Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 9 (off Gloucester); Regan, Ann. Mag. nat. Hist., (8)
2, 1908: 49 (size, distrib.); Murray and Hjort, Depths of Ocean, 191 2: 392 (depth, Faroe Bank);
Roule, Bull. Inst, oceanogr. Monaco, 243, 1912: 13 (Medit. and E. Atlant., list of specs.); Garman,
Mem. Harv. Mus. comp. Zool., 36, 1913: 204, pi. 14, fig. 5-8 (descr., ill., off New England) ; Halkett,
Check List Fish. Canad., 1913: 41 (off Nova Scotia and Massachusetts); Roule, Result. Camp. sci.
Monaco, 52, 1919: 120 (list of E. Atlant. and Medit. specs.); Saemundsson, Vidensk. Medd. naturh.
Foren. Kbh., 74, 1922: 177 (Iceland, Faroe Ridge) ; Bigelow and Welsh, Bull. U.S. Bur. Fish., 40 (l),
1925: 51 (Gulf of Maine and Nova Scotia, general) ; Hickling, Ann. Mag. nat. Hist., (10) 2, 1928: 199
(Atlant. slope off Scotland); Rey, Fauna Iberica, Feces, /, 1928: 451 (descr., ill., Portugal); Breder,
Field Bk. Mar. Fish. Atlant. Coast, 1929: 25 (off New England); Jordan, Manual Vert. Anim. NE.
U.S., 1929: 14 (off C. Ann and fishing banks); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish.
(1928), 2, 1930: 21 (Gloucester and Nova Scotia Banks); MacCoy, Bull. Boston Soc. nat. Hist., 69,
1933: 8 (240 miles ESE. of Boston Lightship) ; Belloc. Rev. des Trav. P£ches Marit., 7 (2), 1934: 144
(ill. after Vaillant; Morocco, Madeira) ; Bigelow and Schrocder, Canad. Atlant. Fauna, biol. Bd. Canad.,
12^, 19H- I 8 (descr., distrib.) ; Noronha and Sarmento, Peixes Madeira, 1934: I 34 (Madeira, not seen) ;
Nobre, Fauna Marinha, Port. Vert., 7, 1935: 455, pi. 63 (depths, off Portugal) ; Vladykov and McKenzie,
Proc. N. S. Inst. Sci., rp, 1 93 5: 49 (off Nova Scotia) ; Fowler, Bull. Amer. Mus. nat. Hist., jo (1), 1936:
74, 75 (descr., no. of embryos, distrib. in E. Atlant., Madeira specs.).
Centrofhorus coeloUfu Gunther, Cat. Fish. Brit. Mus., 8, 1 870: 423 (at least in part, descr., Portugal, Ma-
deira);' "Challenger" Rep., Zool., 22, 1887: XL, 5 (depth of capture); Seabra, Bull. Soc. portug. Sci.
nat., 5, 191 2: 198 (Portugal).

Family DALATIIDAE

Characters. Squaloidea with 2nd dorsal, and in most cases the ist also, lacking a spine;
teeth with i cusp only, but widely unlike in the 2 jaws, the uppers slender and conical, the
lowers broad and blade-like, each overlapping the next outermost, their edges serrate or
smooth.

7. Fowler (Bull. Amer. Mus. nat. Hist., 70 [i], 1936: 75) includes this citation in the synonymy of Centroscymnus
cryptacanthus Regan, 1906. But Gijnther's brief account with the cited references and localities no doubt covers
both that species and coelolepis.

500 Memoir Sears Foundation for Marine Research

Key to Genera

I a. Rear end of base of ist dorsal over or posterior to origins of pelvicsj interspace between
1st and 2nd dorsals shorter than between 2nd dorsal and caudal.

Isistius Gill, 1864, p. 508.
lb. Rear end of base of ist dorsal considerably anterior to origins of pelvics; interspace
between ist and 2nd dorsals considerably longer than between 2nd dorsal and caudal.
2a. Lower teeth erect, triangular, nearly symmetrical, serrate.

Dalatias Rafinesque, 1810, p. 500.^
2b. Lower teeth strongly asymmetrical with cusps directed outward, not serrate, their
outer margins notched.
3a. First dorsal larger than 2nd, its apex angular 5 tip of pectoral angular.

H eteroscymnus Tanakz, 1912.

Japan.

3b. First dorsal at least no larger than 2nd, its apex rounded; tip of pectoral
rounded.

4a. Second dorsal only about as large as ist; snout in front of mouth about
V2 as long as from eye to origin of pectoral.

Somniosus Lesueur, 1 8 1 8, p. 514.
4b. Second dorsal considerably larger than ist; snout in front of mouth
nearly or quite as long as from eye to origin of pectoral.
5a. Second dorsal twice as long at base as ist dorsal, or more; ist dorsal
with a spine either partly free or entirely hidden in the skin.

Euprotomicrus G'dl, 1864.^
Indian Ocean; Philippines; New
Zealand; Pacific between Hawaii
and California; Madeira.

5b. Second dorsal not more than 1V2 times as long at base as ist; ist
dorsal without spine. H eteroscymnoides Fowler, 1934.

Natal, South Africa.

Genus Dalatias Rafinesque, 1 8 10

Dalatias Rafinesque, Carratt. Gen. Nuov. Sicil., l8lo: lO; type species, D. sfarofhagus Rafinesque, 1810,
equals Squalus licha Bonnaterre, 1788, designated by Jordan, Tanaka and Snyder, 1913.^

1. Including Pseudoscymnus Herre, 1935. The differences in the denticles, cited by Herre (Copeia, 1935: 1^4), do
not seem sufficient for generic, and perhaps not even for specific, separation; see p. 501.

2. Including Squaliolus Smith and Radcliffe, 1912.

3. Jordan, Tanaka and Snyder's (J. Coll. Sci. Tokyo, jj, 1913; 22) designation of sparof/iagus Rafinesque, 1810,
as the type of Dalatias is not invalidated by Swainson's (Nat. Hist. Fish. Amphib. Rcpt., t, 1838: 160) acciden-
tal (?) limitation of that genus to D. nocturnus alone in one connection, for on an earlier page (129) in the same
publication Swainson included D. sfarofhagus Rafinesque in it as well. Therefore, Jordan and Evermann's
(Genera Fish., 1, 191 7: 77) subsequent designation of nocturnus Rafinesque, 18 10, as the type species is not
tenable. For the rather confused history of the case, see Gill (Proc. U.S. nat. Mus., 18, 1896: 191), who arrived
at the conclusion that Dalatias is a synonym of Squalus, hence that the correct generic name for the shark now
under consideration is Scymnorhinus Bonaparte, 1846,

Fishes of the Western North Atlantic 501

Generic Synonyms:

Scymnus Cuvicr, Regne Anim., 2, I 81 7: 130; type species, Squalus americantis Gmelin, 1789, equals Sq-udus

Hcha Bonnaterre, 1788, but preoccupied for insects by Kugelman, 1794-
Scymnhim Cuvicr, Regne Anim., ill. ed., Poiss., I 838-1 843: pi. I 15; type species, Squalus jiicaense Cuvier,

equ.ils Squalus licha Bonnaterre, I 788.
Dalatius L. Agassiz, Nomcncl. Zool. Index, i 845 : 21 ; emended spelling for Dalalias.
Scymnorhinus Bonaparte, Cat. Pesc. Europ., 1846: 16; type species, Squalus americanus Gmelin, 1789, equals

Squalus licha Bonnaterre, 1788.
Barborodes Giste!, Naturg. Tierreich, 1848: X; proposed to replace Scymnorhinus Bonaparte, 1846.
Pseudoscymnus Herre, Copeia, 1935: 124; type species P. boshueiisis Herre.

Generic Characters. Dalatiidae without dorsal spines; snout very short; caudal
peduncle without lateral ridges or precaudal pits; a labial fold on each jaw and a
voluminous pit at corner of mouth; upper teeth slender, thorn-like, in several functional
series; lower teeth broad-triangular with regularly serrate edges; dermal denticles low,
ridged, their margins more or less definitely toothed; 2nd dorsal somewhat larger, and
pelvics much larger, than ist dorsal; ist dorsal far anterior to pelvics; rear end of base of
2nd dorsal considerably posterior to origin of pelvics; interspace between ist and 2nd
dorsals considerably longer than between 2nd dorsal and caudal; caudal noticeably large,
with rounded corners, its terminal sector sharply marked off, but lower anterior corner not
expanded as a definite lobe; skin without luminous organs. Characters otherwise those
of the family.

Range. Both sides of North Atlantic; Mediterranean; South Africa; Japan; Aus-
tralia; New Zealand.

Fossil Teeth. Upper Cretaceous, western Asia and North America; Eocene, North
Africa; Eocene to Pliocene, Europe; and Miocene, North America.

Species. The Australian-New Zealand and South African representatives of the
genus have recently been separated'' from the well known D. licha of the North Atlantic
and Mediterranean. But we find nothing in the several accounts or illustrations of the
Australian form (see Synonyms, p. 508) to separate it from licha. It is equally doubt-
ful whether the supposed differences noted by the author of brevipinnis, i.e., lower teeth
more oblique in adult, smaller fins and less pronounced lower caudal lobe, will prove suffi-
cient for specific separation when critically tested.

New generic and specific names {Pseudoscymnus boshuensis) have also been pro-
posed by Herre" for a Japanese form, no doubt the same as one earlier reported as Scy>nnus
lichia^ as Dalatias americanus^ and as D. licha^ the separation being based on its denticles,
which differ in shape between the lower side of the snout and the trunk in general, and its

5. As f /lillipf si Whkley (Aust. Zool., <5, 1931 : 310) and as brevifinnis Smith (Trans, roy. See. S. Afr., 2^, 1936: i)
respectively.

6. Copeia, 1935 : 124.

T. Ishikawa and Matsuura, Prel. Cat. Fish. Mus. Tokyo, 1897: 61.

8. Jordan and Snyder, Annot. zool. jap., 3, 1901: 129.

9. Jordan and Fowler, Proc. U.S. nat. Mus., 26, 1903: 637; also Izuka and Matsuura, Cat. Zool. Tokyo Mus. Vert.,
19Z0: 188.

502 Memoir Sears Foundation for Marine Research

lower teeth, which are serrate. But it has long been known that all this applies to the Atlan-
tic form, and our own comparison of a Japanese specimen with one from the Atlantic coast
of the United States shows no significant differences in proportional dimensions, shape or
position of fins, teeth, or denticles.

Dalatias licha (Bonnaterre), 1788
Figures 96, 97

Study Material. Female, 1,470 mm. long, from Georges Bank (Amer. Mus. Nat.
Hist., No. 14056") j 4 specimens of about 367 to 1,080 mm., from Nice, France, and an
embryo of 245 mm. from the same locality (Harv. Mus. Comp. Zool.) ; also immature
male, 1,114 n^"^- long, from Japan (Harv, Mus. Comp. Zool., No. 11 16).

Distinctive Characters. The serrate margins and triangular shape of its lower teeth
mark D. licha off from all other North Atlantic members of its suborder. It is further
separated very obviously from the species of Squalus, Centroscy Ilium and Etmopterus by
its lack of fin spines, and from Isistius by the position of its first dorsal fin farther forward.

Figure 96. Dalatias licha, female, 1,470 mm. long, from Georges Bank, Gulf of Maine (Amer. Mus. Nat.
Hist., No. 14056). A Head from below. B Left-hand corner of mouth to show labial furrows, about 0.4
natural size. C Right-hand nostril, about 1 .2 x. Z) Dermal denticles from side, below first dorsal fin, about 1 2 x.
E Dermal denticles from ventral surface of snout, about () n. F First to seventh upper teeth, and median
and first to fifth lower teeth from left-hand side, about 1.2 x. G Fourth upper tooth. H Median lower tooth.
G-H, about 2.4 X.

10. Reported five feet one inch (approximately 1,550 mm.) long (Nichols and Firth, Proc. bid. See. Wash., 52,
'939= 85) > ''"t low only '147° """• by the system of measurement here employed (p. 61).

Fishes of the Western North Atlantic

503

Description. Proportional dimensions in per cent of total length. Female, 531 mm.,
from Nice, France (Harv. Mus. Comp. Zool., No. 948). Female, 1,470 mm., from
Georges Bank (Amer. Mus. Nat. Hist., No. 14056).

Trunk at origin of pectoral: breadth 1 1.3, 1 1.7; height 10.9, 9.0.

Snout length in front of: outer nostrils i.i, 0.6; mouth 5.4, 4.6.

Eye: horizontal diameter 3.9, 2.1.

Mouth: breadth 4.5, 5.0; height 0.8, 1.3.

J

'\ /\ A A A A AAAvA

mh-.

V B ^/ \0 M

Figure 97. Dalatias licha, female (dried skin), about 1,080 mm. long, from Europe (Harv. Mus. Comp.
Zool., No. 664). A Left-hand lower teeth, viewed from without. B Dentition of right-hand lower teeth
viewed from within the mouth to show the one series of teeth in function, with five replacement series still
occupying the reversed position with their points directed downward and inward, about 2.3 x natural size.

504 Memoir Sears Foundation for Marine Research

Nostrils: distance between inner ends 3.2, 2.6.

Labial furrow length from corner of mouth: upper 1.8, 1.8; lower 2.0, 1.6.

Gill opening lengths: ist r.6, 1.8} 2nd 1.6, 1.85 3rd 1.6, 1.9; 4th 1.9, 1.9; 5th

2.1, 2.1.

First dorsal -fin: vertical height 5.3, 4.4; length of base 4.9, 5.2.

Second dorsal fin: vtTt\zz\he.\ght. 6. 1, 5.9; length of base 6.2, 6.8.

Cfl«ii<z/^«; upper margin 25.2, 2 1.8 J lower anterior margin I2.l, 11. 3.

Pectoral fin: extreme length 14.1, 13.O; extreme breadth 6.2, 6.3.

Distance from snout to: ist dorsal 35.3, 34.2; 2nd dorsal 60.2, 63.OJ upper

caudal 74.8, 78.25 pectoral 22.1, 19.7; pelvics 55.4, 57.6.

Interspace between: ist and 2nd dorsals 20.6, 23.5; 2nd dorsal and caudal lO.o,

9'5-

Distance from origin to origin of: pectoral and pelvics 36.8, 39.2; pelvics and

caudal 17.9, 18.2.

Trunk slender, subcylindrical, its height at ist dorsal a little less than Yq (15 to
16%) its length to origin of caudal. Body sector to cloaca a little less than twice as long
as tail sector, the back without mid-dorsal ridge. Caudal peduncle without lateral ridges
or precaudal pits. Dermal denticles, over trunk as a whole, small, loose-spaced, scale-like,
their blades close to the skin, thick, quadrate, with 3 weak ridges uniting posteriorly at the
margin in a tooth that varies in length and in acuteness from denticle to denticle 5 pedicels
thick and short. Denticles on lower side of snout overlapping, ovate, without marginal
teeth, but usually with 3 weakly marked longitudinal ridges.

Head about % (24 to 25%) of length to origin of caudal, strongly flattened above.
Snout thick, fleshy, broadly rounded or slightly ovate anteriorly, very short, its length in
front of mouth being only V4 to Vs (about 22%) as great as that of head. Eye oval, its
horizontal diameter % to nearly % as long as snout in front of mouth in late embryos and
newly born specimens, but decreasing in relative size with growth to only about V2 as long
as snout in front of mouth and thus only about % as long as head in adult, its midpoint a
little anterior to front of mouth. Spiracle on dorsal side of head a little above level of eyes,
transverse, about ^2 as long as horizontal diameter of eye. Gill openings small, the longest
about V2 as long as horizontal diameter of eye in newborn, but about as long as eye in adult,
low on the sides, the 5th close in front of pectoral. Nostril close to anterior margin of snout,
oblique, about V2 as long as horizontal diameter of eye, its anterior margin with a low,
triangular lobe, rounded "at the tip. Mouth only very slightly arched, its breadth about as
great as length of snout in front of mouth. Lips noticeably thick and fleshy, but without
special cartilaginous supports near corner of mouth, the lower lip free, but the upper joined
to gum along central Vs of jaw. Upper labial fold extending about Vs of distance toward
symphysis, but lower less than V2 that far. Pit at corner of mouth extremely voluminous,
but its rearward prolongation extending only about Vs or Ve of the distance toward the ist
gill opening.

Fishes of the Western North Atlantic 505

Teeth ^\°l^^l^^^g-, widely unlike in the 2 jaws; upper teeth thorn-like, on broad
bifid bases, curved rearward, erect toward center of mouth but moderately oblique toward
corners, the ist tooth small; lower teeth blade-like, with quadrate bases and broad-
triangular cusps, the latter with regularly serrate edges, erect toward center of mouth,
but oblique and decreasing in size toward corners to a degree apparently depending on
age and perhaps on individual variation; the median lower tooth as large as others,
symmetrical, weakly notched on both edges at junction of cusp with base and overlapping
its neighbor basally on either hand, the lateral lower teeth notched only on outer side and
each overlapping the next outermost tooth; 3 or 4 series functional in upper jaw, and i or
2 series functional in lower jaw, depending on the stage in replacement.

First dorsal only about % as long at base as head, brush-shaped with broadly
rounded apex, its posterior margin nearly straight and perpendicular, its free rear corner,
or free lower margin, about as long as base, Its origin posterior to tips of pectorals by a
distance about as long as horizontal diameter of eye when pectorals are laid back. Second
dorsal a little larger than ist, its origin about over middle of bases of pelvics, its distal
margin concave and rear corner acute, thus differing from ist dorsal, its free rear tip about
as long as its base. Interspace between 2nd dorsal and caudal a little more than i V2 times as
long as base of 2nd dorsal. Caudal a little more than Vs of total length, obliquely truncate
terminally with broadly rounded apex, its lower margin deeply incised subterminally in
rectangular outline and thus sharply marking off the terminal sector, its lower anterior
corner rounded, about a right angle, the lower anterior margin a little less than V2 as long
as upper margin. Pelvics about i Y^ times as long at base as 2nd dorsal, with nearly straight
margins, broadly rounded apices and tapering rear corners. Pectoral about % as long as
head, paddle-shaped, with very broadly rounded tip, weakly convex outer margin and
more strongly convex distal margin, the transition from distal to inner margin gradual,
there being no definite inner corner.

Color. After preservation, uniformly dark chocolate or cinnamon brown below as
well as above; also described as sometimes violet brown with poorly defined blackish spots
in life, the fins with pale or whitish margins and caudal black-tipped.

Size. The young are born at a length of approximately 300 mm., but most of those
caught are between 1,000 and 1,500 mm. (40 to 60 inches) long; the longest of which we
have found definite measurement was 1,820 mm. (72 inches)." A specimen of about five
feet weighed about 23^/'2 pounds gutted."^ Females are larger than males, as is commonly
the case among sharks.

Developmental Stages. Development is ovoviviparous. Gravid females are reported
as containing 10 to 16 young. An embryo of 270 mm., still with the large yolk sac, already
shows all the diagnostic characters of the adult except for the teeth, denticles and rela-
tively larger eyes (horizontal diameter about % as long as snout in front of mouth). ^^

11. Dumeril, Hist. Nat. Poiss., /, 1865: 452.

iia. Nichols and Firth, Proc. biol. Soc. Wash., 52, 1939: 85.

12. For account of the uterine wall of a gravid female, see Ranzi (Publ. Staz. zool. Napoli, /j, 1934: 366).

5o6 Memoir Sears Foundation for Marine Research

Habits. In its centers of abundance in the eastern Atlantic this shark is taken most
often in at least moderately deep water. OS Nice, on the Mediterranean coast of France,
it was long ago described as commonly caught at i,ooo meters depth; other depth records
are from 300 to 600 meters and many have been taken on the Irish Atlantic slope between
200 and 350 fathoms (366 to 640 m.)/' But since the Georges Bank specimen was taken in
only 50 fathoms, with report of at least one other on the beach at Madeira," it is apparent
that it is not confined exclusively to deep water, and its New Zealand representative is also
occasionally washed ashore. Gravid females are taken throughout the year in the Medi-
terranean. Nothing whatever is known of its feeding habits, although the nature of its
teeth, and the fact that Mediterranean ones have been commonly caught on hook and
line, suggest a fish diet.

Relation to Man. It is of relatively little commercial importance at present, but
in the Azores it is the object of a special fishery for leather; formerly its skin was prized
as an abrasive by cabinet makers and jewelers.

Range. In the eastern Atlantic from Rio de Oro, the Canaries, Madeira, Morocco,
Azores and western Mediterranean north to the Irish Atlantic slope; plentiful locally
(perhaps periodically) off the Mediterranean coasts of France and Portugal, as well as on
the fishing grounds west of Ireland. There is but one record for the western Atlantic (see
below). If is represented off South Africa, in the New Zealand-Australian region and in
Japanese waters by allies so close that they appear to be identical with the Atlantic form
(p. 501),

Occurrence in the Western Atlantic. The only record of the capture of this shark in
the western Atlantic is the female here pictured (Figs. 96, 97), about five feet long, taken
on the northern edge of Georges Bank in 50 fathoms on August 19, 1937."

Synonyms and References:

1. North Atlantic:

La liche, Broussonet, Mem. Math. Phys. Acad. Sci. Paris, 1780: 677 (C. Breton, France); Duhamel, Traite

Gen. Pcches, 4, 1782: 301, 328 (descr.).
Squalus licha Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: 12 (descr., size).
Squalus americanus Gmelin, in Linnaeus, Syst. Nat., J (3), I 789: 1503 (descr., named on incorrect assumption

that Cap du Breton specimen of Broussonet, 1780, was taken at the Nova Scotian loc. of that name);

Walbaum, P. Artedi Genera Pise. Emend. Ichthyol., j, 1792: 521 (descr.) ; Bloch and Schneider, Syst.

Ichthyol., 1801: 136 (descr.); Bosc, Nouv. Diet. Hist. Nat., 21, 1803: 192 (diagn.) ; Latreille, Nouv.

Diet. Hist. Nat., 24, 1804: 72 (in contents); Shaw, Gen. Zool., 5, 1804: 347.
Le squale liche, Lacepede, Hist. Nat. Poiss., 4° ed., i, 1 798: 279, pi. 10, fig. 3 (descr., ill., C. Breton), also

other eds.; in Sonnini, Hist. Nat. Poiss., 4, 1802: 117 (same as Lacepede, 1798).
Squalus nicaetisis Risso, Ichthyol. Nice, 1 8 10: 43, pi. 4, fig. 6 (descr., occur, near Nice, France).
Dalatias ifarofhagus Rafinesque, Carrat. Gen. Nuov. Sicil., 1810: 10, pi. 13, fig. 2 (descr. and ill. recogniz-
able, though spiracle said to be lacking) ; Indice Ittiol. Sicil., 1810:44 (Sicily) .
Scymnus americanu! Cuvier, Regne Anim., 2, 1817: 130 (genl., notes earlier error regarding type loc, C. du

13. Fraser-Brunner, Proc. R. Irish Acad., 42, 1935: 320. 14. Bowditch, Excurs. Madeira, 1825: 74..

15. Nichols and Firth, Proc. biol. Soc. Wash., 52, 1939: 85.

Fishes of the Wester?i North Atlantic 507

Breton, not in America), also later eds. and translations; Bory de St. Vincent, Diet. Class. Hist. Nat., 75,
1829: 598; Bonaparte, Mem. Soc. ncuchatcl Sci. nat., 2 (8), I 839: 9 (in synopsis).

Scymnus vulgaris Cloquet, Diet. Sci. Nat., 25, 1822: 433 (general); Reguis, Ess. Hist. Nat. Provence, / (i),
1877: 56 (descr., Medit.).

Scymnus licha Bowditch, T. E., Excurs. Madeira, 1825: 74 (Madeira); Roulc, Result. Camp. sci. Monaco,
52, 1917: 123 (off Lisbon).

Scymnus jiicaensis Risso, Hist. Nat. Europe Merid., 3, 1826: 137, pi. 2, fig. 4 (descr., size, color, Medit.);
Lowe, Trans, zool. Soc. Lond., 2 (3), 1837: 194 (Madeira); Cuvier, Regne Anim., ill. ed., 1843: pi.
115, fig. 5 (jaws).

Squalus {Acanthorhinus) americanus Blalnville, in Vieillot, Faune Franc. Poiss., 1 825 : 63, pi. I 5, fig. 2 (descr.,
ill., France).

Squalus scymnus Voigt, in Cuvier, Tierreich, 2, 1832: 512.

Scymnus lichia Bonaparte, Icon. Faun. Ital., 5, 1835-1836: plate not numbered (descr., ill., Italy); Agassiz,
L., Poiss. Foss., 5, 1843: 367, pi. 40B, fig. 5 (vertebrae); Busch, W., Selach. Ganoid. Encephal., 1848:
2 2, pi. 2, fig. 3, 4 (brain) ; Van der Hoeven, Handb. Dierkunde, (2) 2, 1855: 260 (general) ; Canestrini,
Arch. Zool. Anat. Fisiol. Bologna, i, 1861: 267 (off Genoa) ; Fitzinger, Bild. Atlas Nnturg. Fische, 1864:
fig- 179 (good '11-)! Bocage and Brito Capello, Poiss. Plagiost., 1866: 34 (Portugal); Brito Capello, J.
Sci. math. phys. nat. Lisboa, 2, 1869: 145 (Portugal); Giinther, Cat. Fish. Brit. Mus., 8, 1870: 425
(descr., refs., Medit., Madeira) ; Miklucho-Macklay, Beitr. Vergl. Neurol. Wirbelt., /, 1870: 11, pi. I,
fig. 3, 4 (brain); Canestrini, in Cornalia, el al., Fauna d'ltal., 1872: 41 (Medit.); Gegenbaur, Unters.
Vergl. Anat. Wirbelt., j, 1872: 23, 24, pi. I, fig. 3, pi. 7, fig. 3, pi. II, fig. 1, pi. 17, fig. 4> ?'• I9> fig- 2,
pi. 22, fig. 5-7 (anat.); Hertwig, Jena. Z. Naturw., 8, 1874: 349, pi. 12, fig. 10, 13 (develop, of
denticles, teeth); Gervais and Boulart, Poiss., 3, 1876: 210, pi. 8l (Medit., Portugal); Doderlein,
Prosp. Metod. Pesci Sicil., 1 878-1 879: 30 (near Sicily); Man. Ittiol. Medit., 2, 1881: 100 (west.
Medit.) ;^' Moreau, Poiss. France, j, 1881: 358 (descr., ill., French coasts); Hasse, Naturl. Syst. Elasm.
besond. Theil, 1882: 65, pi. 9 (skelet., dermal denticles) ; Carus, Prod. Fauna Medit., 1889-1893: 501
(W. Medit.) ;" Collett, Bull. Soc. zool. Fr., 15, 1 890: 219 (Madeira) ; Almeida and Roquete, Mammif.
Peix. Costa e rias do Algarve Inquerit. Indust. Lisboa (1889), 1892: 374 (Portugal, not seen); Vieira,
Ann. Sci. nat. Porto, 4, 1897: 138 (Portugal); Sicher, Atti Accad. gioenia, (4) // (5), 1898: 18
(Medit.); Jungerson, Danish "Ingolf" Exped., 2 (l), 1899: 31 (claspers) ; Huber, Z. wiss. Zool., 70,
1901: 604, pi. 27, fig. 3, 39 (claspers); Bridge, Camb. nat. Hist., 8, 1904: 455, and subsequent eds.
(general, Medit., neighboring Atlantic) ; de Braganza, Result. Invest. Sci. "Amelia," 2, 1904: 90, 106
(Portugal); Helbing, Nova Acta Leop. Carol., 82 (4), 1904: 523 (comp. with Somniosus, skelet.);
Seabra, Cat. Poiss. Port., 1911: 199 (Portugal); Roule, Bull. Inst, oceanogr. Monaco, 243, 1912: 17
(west. Medit., off Morocco, Madeira); Jenkins, Fish. Brit. Isles, 1925: 326 (descr.); Leigh-Sharpe,
J. Morph., 42, 1926: 314 (claspers); Ranzi, Pubbl. Staz. zool. Napoli, 13 (3), 1934: 340, 366
(uterine liquid and wall) ; Needham, Biochem. Morphogen., 1 942 : 4 1 (embryonic weight) .

Scymnus (no specific name) Agassiz, L., Poiss. Foss., 5, 183.5-1837: 12, pi. f, fig. 7 (teeth).

Scymnium niciense Cuvier, Regne Anim., ill. ed., 1843: pi. 115, fig. 5 (ill., jaws).

Scymnus {Scymnus) lichia Miiller and Henle, Plagiost., 1 841: 92 (descr., Medit., French coast); Dumeril,
Hist. Nat. Poiss., /, 1865:452 (descr., distrib.).

Scymnorhinus lichia Bonaparte, Cat. Pesc. Europ., 1846: 16 (in list); Goode and Bean, Smithson. Contr.
Knowl., 30, 1895: 7, pi. 2, fig. 4; Mem. Harv. Mus. comp. Zool., 22, 1896: 7, pi. 2, fig. 4 (descr., ill.,
W. Medit., Madeira) ; Regan, Ann. Mag. nat. Hist., (8) 2, 1908: 57 (specs, in Brit. Mus.) ; Belloc, Rev.
des Trav. Peches Marit., 7 (2), 1934: 148 (ill. after Lozano Rey; Morocco, Rio de Oro, Canaries);
Borri, Atti Soc. tosc. Sci. nat., 44, 1934: 88 (Medit.) ; Lubbert and Ehrenbaum, Handb. Seefisch. Nord-
europ., 2, 1936: 290 (W. Medit., Portugal); Nichols and Firth, Proc. biol. Soc. Wash., 52, 1939: 85
(Georges Bank spec, length, weight).

Dalatias licha Gray, List. Fish. Brit. Mus., r, I 851: 75 (Madeira); Fowler, Proc. Pan-Pacif. sci. Congr.,

16. See Doderlein, 1881, and Carus, 1889-1893, for additional records for the Mediterranean in publications not
accessible to us.

5o8 Memoir Sears Foundation for Marine Research

4, 1930: 497 (Atlant.) ; Bull. Amer. Mus. nat. Hist., yo (i), 1936: 86 (descr., ill., Medit. spec);

Cadcnat, Rev. des Trav. Peches Marit., 10 (4), 1937: 432 (Azores; special fishery for leather); Fowler,

Bull. U.S. nat. Mus., loo {13), 1941: 267 (descr., refs., Atlant. and Pacif.).
Squalus (Srymnus) lichia Van der Hoeven, Handb. Dierkundc, 2, 1855: 260.
Scymnor/iiuu! licha Garman, Mem. Harv. Mus. comp. Zool., ^6, 1913: 237 (descr.); Rey, Fauna Iberica

Feces, 1, 1928: 474, pi. 5, fig. I (Iberian coasts) ; Uriarte and Mateu, Notas Inst. esp. Oceanogr., (2) 5j,

1931: 26 (Canaries); Fraser-Brunner, Proc. R. Irish Acad., 42, 1935: 320 (Irish Atlant. slope, color);

Nobre, Fauna Marinha Port. Vert., i, 1935: 460, pi. 64, fig. 200 (descr., ofl[ Portugal) ; Tortonese, Atti

Soc. ital. Sci. nat., yy, 1938: 311 (Medit.).
Scymnorhinus licki {Scymnus lichia) Holmgren, Acta zool., 22 (1-3), 1941: 24 (skull).

2. South .Africa and Pacific; apparently referrable to D. licha-y

Scymnus lichia Parker, Trans. Proc. N. Z. Inst., /f, 1883: 223, pi. 31, 32 (descr., ill., N. Zealand) ; Ishikawa

and Matsuura, Prel. Cat. Fish. Mus. Tokyo, 1897: 61 (Japan).
Dalatias americanus Jordan and Snyder, Annot. zool. jap., j, 1901 : 129 (Japan).
Dalalias licha Jordan and Fowler, Proc. U.S. nat. Mus., 2(5, 1 903: 637 (Japan) ; Izuka and Matsuura, Cat. zool.

Mus. Tokyo, 1920: 188 (Japan) ; Fowler, Proc. Pan-Pacif. sci. Congr., 4, 1 930: 497 (Aust., N. Zealand,

Japan, Atlant.); Bull. U.S. nat. Mus., 100 {13)-, 1941: 267 (descr., listed Japan, also Atlantic and

Medit.).
Scymnorhinus licha McCulloch, Res. "Endeavour," 2 (3), 1914: 81, pi. 14 (ill., Aust.) ; Waite, Trans. Proc.

N. Z. Inst., 46, 1914: 128, pi. 4, fig. I (ill., N. Zealand); Rec. S. Aust. Mus., 2, 1921: 24, fig. 33;

Handb. Brit. Sci. Guild, Fish. S. Aust., 1923: 43; Phillipps, N. Z. J. Sci. Tech., 10, 1 928: 224 (N.

Zealand).
Scymnorhinus fhillifpsiWhltley, Aust. Zool., 6, 1 931 : 3 10; Fish. Aust., i, 1940: 150 (ill., Aust., N. Zealand).
Pseudoscymnus bosliuensis Herre, Copeia, 1935: 124 (descr., Japan) ; Fowler, Bull. U.S. nat. Mus., 100 (z^),

1941: 275 (descr., Japan).
Scymnorhinus hrevipinnis Smith, Trans, roy. Soc. S. Afr., 24, 1936: I (descr., S. Afr.).
Dalatias fhilliffsi Fowler, Bull. U.S. nat. Mus., 100 (rj), 1941: 268 (Aust., N. Zealand).
Dalatias brevifinnis Fowler, Bull. U.S. nat. Mus., lOO (rj), 1941 : 268 (S. Afr.).

Genus Isistius Gill, 1864

Isistius Gill, Proc. Acad. nat. Sci. Philad., 1864: 264; type species, Scymnus brasiliensis Quoy and Gaimard,
1824, monotypic.

Generic Synonyms:

Scymnus (in part) Quoy and Gaimard, Voy. "Uranie," Zool., 1824: 197; not Scymnus Cuvier, 1817.

Dalatias (in part) Gray, List. Brit. Mus., /, 1851 : 76; not Dalalias Rafinesque, I 8 10.

SyW«j Johann, Z. Wiss. Zool., 66, 1891 : 152; for S. {Scymnus) Jul gens Bennett, 1 840; not Squalus Linnaeus,

1758.
Leius Kner, Denkschr. Akad. Wiss. Wien., 24, 1865: 9; type species, L. ferox Kner, equals ScyTnnus brasiliensis

Quoy and Gaimard, 1824.

Generic Characters. Dalatiidae without dorsal fin spines; snout in front of mouth
much shorter than from front of mouth to origin of pectorals; caudal peduncle without
lateral ridges or precaudal pits; pit at corner of mouth prolonged below upper lip and
rearward as a narrow furrow; expanded lips at corners of mouth with special cartilaginous
supports; teeth widely unlike in the 1 jaws, the uppers slender, thorn-like, widely spaced,
the lowers with triangular smooth-edged or partly serrate cusp and quadrate base, each

17. See page 501.

Fishes of the Wester ti North Atlantic

509

overlapping the next outermost; dermal denticles low, with depression in the crown; rear
end of base of ist dorsal about over origin of pelvics; 2nd dorsal and pelvics only a little
larger than ist dorsal; interspace between ist and 2nd dorsal much shorter than between
pelvics and caudal; caudal with axis approximately in continuation of main axis of trunk,
very broad relative to its length, with shallow subterminal notch, its lower anterior corner
expanded as a well defined lobe; pectoral small, paddle-shaped; skin sprinkled with
strongly luminescent, glandular points. Characters otherwise those of the family.

Range. Tropical and subtropical belts of Atlantic, Pacific and Indian Oceans.

Fossil Teeth. Upper Cretaceous to Eocene, Africa; Eocene to Miocene, Europe.

Species. Only one species is known.

Isistius brasiliensis (Quoy and Gaimard), 1824

Figures 98, 99

Study Material. Immature male, 383 mm. long, taken by the Research Vessel
"Atlantis," Station 2947, north of the Bahamas at Lat. 25° 11' N., Long. 77° 19' W.,
where the depth was 1,000 fathoms^ (Harv. Mus. Comp. ZooL, No. 36039); two fe-

FicuRE 98. Isistius brasiliensis, immature male, about 383 mm. long, from north of the Bahamas (Harv.
Mus. Comp. Zool., No. 36039). A Anterior part of head from above to show spiracles, about 1. 1 x. S Head
from below, about I.I x. C Dermal denticles, about 4.5 x. D Lateral oblique view of dermal denticle, about
45 X.

I. But not necessarily from so great a depth, as the nets were brought up open to the surface.

5IO

Memoir Sears Foundation for Marine Research

males, 465 and 501 mm. long, from Japan (Harv. Mus. Comp. ZooL, No. 1368, 1245);
female about 485 mm. long, from the vicinity of Albemarle Island, Galapagos, Lat. 2°
34' N., Long. 92° 06' W., in a trawl haul from 1,360 fathoms. Albatross Sta. 3413
(Harv. Mus. Comp. Zool., No. 1005).

Distinctive Characters. Isistius is separated from all other northwestern Atlantic
sharks of its suborder except Dalatias by the triangular shape of its lower teeth j the posi-

FiGURE 99. Isistius brasiliensis, pictured in Fig. 98. Upper and lower teeth, left-hand side, about 4.4 x.

tion of its first dorsal fin far rearward marks it off at a glance from Dalatias. It shares this
last character with Echinorhinus, but there is no danger of confusing it even with new-
born specimens of the latter, for the shape of its caudal, its teeth and its dermal denticles
are very different and its gill openings much smaller.

Description. Proportional dimensions in per cent of total length. Male, 383 mm.,
from Lat. 25° 1 1' N., Long. 77° 19' W. (Harv. Mus. Comp. ZooL, No. 36039). Female,
501 mm., from Tokyo, Japan (Harv. Mus. Comp. Zool., No. 1245).

Trunk at origin of -pectoral: breadth 10.2, ii.O; height 8.4, ii.o.

Snout length in front of: outer nostrils 0.8, 0.8; mouth 7.0, 6.6.

Eye: horizontal diameter 3.4, 3.2.

Mouth: breadth 4.7, 4.2; height O, o.

Nostrils: distance between inner ends 1.6, 1.4.

Labial furrow length from angle of jaw: upper 4.2, S-^-

Gill opening lengths: ist 0.8, 0.8; 2nd 0.8, 0.8; 3rd 0.8, 0.8; 4th 0.8, 0.8; 5th

0.8,0.8.

First dorsal fin: vertical height 3.4, 2.6; length of base 3.1, 3.2.

Second dorsal fin: vertical height 2.9, 2.4; length of base 3.9, 3.8.

Caudal fin: upper margin 14.6, 15.3; lower anterior margin 1 1.2, 10.2.

Pectoral fin: outer margin 7.8, 7.3; inner margin 4.4, S-6; distal margin 4.2, 4.2.

Distance from snout to: ist dorsal 59.5, 59-3; 2nd dorsal 70.5, 71.8; upper caudal

85.4, 84.7; pectoral 19. i, 17.9; pelvics 60.1, 62.8.

Fishes of the Wester 71 North Atlantic 511

Interspace between: ist and 2nd dorsals 8.5, 9.3; 2nd dorsal and caudal 10.7, 9.6.
Distance from origin to origin of: pectoral and pelvics 43.3, 44.1 5 pelvics and
caudal 22.9, 21.9.

Trunk subcylindrical and very slender, its greatest height only about % its length
to origin of caudal, the dorsal profile only weakly arched and the ventral profile nearly
straight. Body sector to cloaca more than twice as long as tail sector. Caudal peduncle with-
out lateral ridges or precaudal pits, and without mid-dorsal ridge. Dermal denticles small,
closely spaced, but with skin exposed between them, highly diagnostic in shape, being very
low with no distinction between pedicel and blade, transversely truncate apically, quadri-
lateral in outline, but occasionally polygonal, with concave margins and rounded corners,
the crown with a quadrate depression.

Head a little less than y^ (22 to 23%) of length to origin of caudal. Snout thick,
fleshy, broadly-ovate and very short, its length in front of mouth a little less than Vs of
length of head. Eye oval, its horizontal diameter almost V2 as great as length of snout
in front of mouth. Spiracles on top of head, slightly oblique, oval, about V2 as long
as horizontal diameter of eye and posterior to latter by a distance a little greater than their
own length. Gill openings very small, only about Ys to ^/4 as long as horizontal diameter
of eye, rather high on the sides, the 5th close in front of origin of pectoral. Nostril close to
anterior margin of snout, about Vs as long as horizontal diameter of eye, oblique, its ante-
rior margin expanded as a short broadly-triangular lobe rounded at the tip. Mouth very
little arched. Lips fleshy, adnate to gum along midsector of upper jaw but elsewhere free
and widely distensible, with rounded wing-like expansions at corners of mouth, enclosing
an extensive funnel-like cavity that extends inward along upper jaw on either side; upper
lip overlaps lower as a thin skin fold rearward from corner of mouth for about V2 of the
distance to 2nd gill opening.

Teeth Jy^is^^^T^TfS^' ^^owers increasing in number with growth ; upper teeth slender,
thorn-like, increasingly curved outward toward corners of mouth; lowers much larger,
erect, symmetrical, their bases subquadrate with a shoulder on each side at point of transi-
tion to the triangular, sharp-pointed cusp, their edges mostly smooth, partly wavy, or even
showing faint indication of serrations, the central lower tooth overlapping its neighbor
on either side basally, with each subsequent tooth correspondingly overlapping the next
outermost; the outermost lower tooth widely expanded basally on the outer side; 3 to 4
series functional in upper jaw, one in lower.

First dorsal very small, its base between Vs and % as long as head, sloping, with
broadly rounded apex, its posterior outline very weakly concave, its free rear tip slender
and about as long as the base, the rear end of base about over origin of pelvics. Second
dorsal similar to ist, but a little longer at base and larger in area, its origin about over
rear tips of pelvics. Interspace between ist and 2nd dorsals about as long as between 2nd

512 Memoir Sears Foundation for Marine Research

dorsal and caudal. Caudal only about Vr the total length, about % as broad as long, the tip
broadly rounded, its posterior outline deeply concave in angular contour, marking off the
terminal sector, the lower lobe broadly triangular with narrowly rounded tip, its anterior
margin about % as long as upper margin of fin. Pectoral about % (40-45%) as long as
head to 5th gill opening.

Color. Dark brown above, paler brown or brownish white below, except for a con-
spicuous dark collar around the neck in the region of the gill openings; fins brown, the
pectorals, dorsals and pelvics with pale distal margins, the upper and lower lobes of the
caudal with dusky or darker brown tips; the inner side of upper eyelid not pigmented. Ex-
cept in the region of the dark collar the lower surface is closely, and the sides sparsely,
sprinkled with black dots, presumably luminous ;'' these also occur in patches on the sides
of the head, on the dorsal and caudal fins, and on the basal parts of the pectorals, with a few
along the back.

Size. Recorded specimens have ranged in length from about 140 mm. (5% inches)
to about 495 mm. (19V2 inches); females are mature at a length of 18 inches.

Developmental Stages. Presumably development is ovoviviparous, but the only
available definite information is that a female has been reported as containing six large
eggs.'

Habits. This is a pelagic species, the majority of recorded specimens having been
taken either from small depths or at the surface at night. And while a few have been re-
corded from deep hauls,* it is likely that they were picked up by the net on its way down
or up. Nothing is known of its diet, nor of its breeding habits.

This is the most brilliantly luminescent of sharks. According to an eyewitness
account'' the entire lower surface of its trunk, with the exception of the dark collar, its
paired fins and its caudal, emits a vivid greenish light. While the luminescence apparently
is not under nervous control, since it is not affected by handling, it is not a constant charac-
teristic of the species, for one specimen taken alive failed to show any trace of it."

Range. The localities of capture include the Gulf of Guinea, the offings of Sierra
Leone and Cape Verde in the eastern Atlantic, as well as Brazil, the Bahamas, and north
of the Bahamas in the western Atlantic; also the vicinity of the Galapagos, Hawaiian Is-
lands, Japan, Fiji, central equatorial Pacific west of Christmas Island, equatorial belt
north of New Guinea, Lord Howe Island off New South Wales, Australia, Mauritius,
and between Java and western Australia. These localities are dispersed widely enough to
prove this shark cosmopolitan in the tropical and subtropical belts of all three oceans. Rec-
ords for the western Atlantic are: off Rio de Janeiro (one specimen), among the Bahamas

z. The distribution of these has been described in detail by Burckhardt (Ann. Mag. nat. Hist., [7] 6, 1900:565, 566).

3. Bennett, Narr. Whaling Voy., j, 184.0: 255.

4. See Study IVIaterial, p. 509; also Garman (Mem. Harv. Mus. comp. Zool., 34, 1899; 40) and Parr (Bull.
Bingham oceanogr. Coll., 3 [7], 1937: i).

5. F. D. Bennett's (Narr. Whaling Voy., 2, 1840: 255) account has been quoted repeatedly.

6. Duncker and Mohr, Mitt. zool. Stinst. Hamburg, ^j, 1929: 84.

Fishes of the Western North Atlantic 513

(one specimen), and about 160 miles north of the Bahamas (one specimen). See Study
Material, p. 509.

Synonyms and References:

Scymnus brasiliensis Quoy and Gaimard, Voy. "Uranie," Zool., 1824: 198 (brief descr., Brazil).

Squalus (Scymnus) fulgeus Bennett, F. D., Narr. Whaling Voy., 2, I 840: 255 (dcscr., size, eggs, luminescence,
trop. Pacif. near Christmas I.') ; Bennett, G., Gatherings Nat. Australia, i860: 66 (luminescence, a sec-
ond trop. Pacif. spec).

Scymnus {Scymnui) brasiliensis (inch var. torqualus and var. unicolor) Muller and Henle, Plagiost., 1 841 : 92
(dcscr., Mauritius, C. Verde at St. Jago, Rio de Janeiro) ; Dumeril, Hist. Nat. Poiss., i, 1865: 4.53 (refs.,
dcscr., Mauritius).

Dalatias brasiliensis Gray, List. Fish. Brit. Mus., j, 1 85 1 : 76 (Isle of France, St. Jago, Rio de Janeiro).

Scymnus torqualus Dumeril, Arch. Mus. Hist. nat. P.iris, ro, 1861: 261 (name only, St. Jago, C. Verde).

Isislius brasiliensis Gill, Proc. Acad. nat. Sci. Philad., 1864: 264, footnote (name) ; Giinther, Cat. Fish. Brit.
Mus., S, 1870: 429 (refs., descr., S. Pacif. and Gulf of Guinea specs.) ; Peters, W. C. H., Monatschr.
Berlin Akad., 1876: 853 (Indian Oc, Lat. 14° 23' S., Long. 118° 16' E.) ; Rochebrune, Act. Soc. linn.
Bordeaux, (4) 6, 1 882: 48; Faune Senegambie, Poiss., /, I 883-1 885: 26 (C. Verde) ; Sauvage, in Gran-
didier, Hist. Phys. Nat. Madagascar, 16, 1 891 : 5, 511 (spec, in Paris Mus.) ; Garman, Mem. Harv. Mus.
comp. Zool., 24, 1899: 34, pi. I, fig. I, pi. 2-3, pi. 69, fig. 2 (descr., size, no. of teeth, anat., near Gala-
pagos) ; Johann, Z. wiss. Zool., 66, 1899: 152 (luminescence) ; Burckhardt, Ann. Mag. nat. Hist., (7) 6,
1900: 56; (luminescence, luminous organs) ; Waite, Rec. Aust. Mus., j, 1900: 195, fig. I, 2 (no. teeth,
largest recorded spec, Lord Howe I.); Mangold, Pflug. Arch. Gcs. Physiol., rrp, 1907: 583 (lumines-
cence); Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907: 169, 205 (descr., refs., Brazil); Brauer, Wiss.
Ergebn. 'Valdivia,' 15, 1908: I 33, pi. 2 (luminescence) ; Regan, Ann. Mag. nat. Hist., (8) 2, 1908: 55
(class., size); Garman, Mem. Harv. Mus. comp. Zool., 56, 191 3: 238 (descr.); Jordan, Tanaka and
Snyder, J. Coll. Sci. Tokyo, 55, 1913: 23 (old loc records, also Fiji, Japan) ; Lampe, Deutsche sudpolar
Exped., 75, Zool., 7, 1914: 214, pi. 2 (no. of lower teeth, sizes, 3 spec, off Sierra Leone, W. Afr.) ;
Metzelaar, Trop. Atlant. Visschen, 191 9: 191 (refs.) ; Ribeiro, Fauna brasil., Peixes, Mus. nac. Rio de J.,
2 (i) Fasc. I, 1923: 26 (same as Ribeiro, 1907); Fowler and Ball, Bull. Bishop Mus., 2(5, 1926: 5
(footnote that Echinorhinus brucus Fowler, 1923, from Hawaii, was actually /. brasiliensis) ; Whitley,
J. Pan-Pacif. Res. Instn., 2 (i), 1927: 3 (Fiji); Fowler, Mem. Bishop Mus., 10, 1928: 23 (descr.,
refs., Honolulu) ; Duncker and Mohr, Mitt. zool. StInst. Hamburg, 44, 1929: 84 (size, depth of capture.
Equatorial Pacif. near New Guinea, Lat. 00° N., Long. 146° 5' E.) ; Fowler, Proc. Pan-Pacif. sci.
Congr., (4) J, 1930: 497 (distrib.) ; Mem. Bishop Mus., //, 1931: 314 (refs.) ; Bull. Amer. Mus. nat.
Hist., -JO (1), 1936: 87 (refs., descr.); Parr, Bull. Bingham oceanogr. Coll., 5 (7), 1937: I (north of
Bahamas); Beebe and Tee-Van, Zoologica, N. Y., 26, 1 941: 121 (ref.) ; Fowler, Bull. U.S. nat. Mus.,
100 ( /j), I 941 : 270 (refs., descr., distrib., luminescence) ; Arqu. Zool. Estado S.io Paulo, 5, 1942: I 29
(Brazil) ; Bigelow and Schroeder, Guide Coram. Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945:
146, fig. 55 (descr., ill., habits, range).

Leius ferox Kner, Denkschr. Acad. math. Natur. Wien, 24, 186;: 10, pi. 4, fig. 2 (descr., ill., Australia);
Schmeltz, Cat. Mus. GodeflFroy, 5, 1866: 13 (South Seas, not seen); Johann, Z. wiss. Zool., 66, 1899:
152 (luminescence); Whitley, Fish. Aust., /, 1940: 149 (descr., ill., luminosity, Aust.).

Isislius ferox Schmeltz, Cat. Mus. Godeffroy, 2, I 865: 10 (South Scis, not seen).

Isislius braziliensis Macleay, Proc. linn. Soc. N. S. W., 6, 1881 : 368 (ref., Aust.).

Sgualus fulgens Johann, Z. wiss. Zool., 66, 1899: 152 (luminescence).

Leius brasiliensis Giinther, J. Mus. Godeffroy, 6 (17), 1910: 490 (refs., depth, distrib.).

Afristurus sfongicefs Jordan and Jordan, Mem. Carneg. Mus., 10, 1922: 3 (Hawaii) ; not Catulus sfongicep
Gilbert, 1905.*

7. A second specimen reported from Lat. 55° N., Long. 110° W.; either the latitude or long^itude is evidently an
error, for this would place it far inland in Canada.

8. A cast in Bishop Museum, later identified by Fowler (Mem. Bishop Mus., 10. 1928: 23) as /. brasiliensis.

514 Memoir Sears Foundation for Marine Research

Echinorhinus brucus Fowler, Occ. Pap. Bishop Mus., 8 (7), 1923: 375 (Honolulu); not Squalus brucus
Bonnaterre, 1788.'

Genus Somniosus Lesueur, 1 8 1 8

Somniosus Lesueur, J. Acad. nat. Sci. Philad., i, 1818: 222; type species, S. brevipjnna Lesueur, Massachusetts,
monotypic.

Generic Synonyms:

Squalus Gunnerus, Drontheim Gesellsch. Schr. Leipzig, 2, 1776: 299; for S. carcharias Gunnerus; not Squalus

Linnaeus, 1758.
Acanthorhinus (in part) Blainville, Bull. Soc. philom. Paris, 1816: 121: for 5. nortvegianus Blainville.^
Scymnus (in part) Fleming, Hist. Brit. Anim., 1828: 166, for S. borealis Fleming, equals Squalus borealis

Scoresby, 18 20; not Scymnus Cuvier, 181 7.
Laemargus (subgenus) Muller and Henle, Plagiost., 1 841: 93; type species, 5. borealis Fleming, 1828, equals

Squalus borealis Scoresby, I 8 20, Spitzbergen.
Leiodon Wood, Proc. Boston Soc. nat. Hist., 2, 1846: 174; type species, S. echtnatum Wood, monotypic.
Dalatias (Somniosus) Gray, List Fish. Brit. Mus., 7, 1851 : 76, for Scymnus (Laemargus) borealis Muller and

Henle, 1841, equals Squalus borealis Scoresby, 1820; not Dalatias Rafinesque, 1810.
Scimnus Vzn Beneden, Bull. Acad. Sci. Roy. Bruxelles, 20 (2), 1853: 258; emended spelling for Scymnus.
Rhinoscymnus Gill, Proc. Acad. nat. Sci. Philad., 1864: 264, footnote 5; type species, Scymnus rostratus Risso,

1826, monotypic.

Generic Characters. Dalatiidae without dorsal fin spines 5 snout in front of mouth
much shorter than from front of mouth to origin of pectorals; midline of back with a
faint dermal ridge; caudal peduncle with faint lateral ridges, at least in some cases, but
without precaudal pits; labial furrows and a pit prolonged rearward from corner of mouth ;
teeth widely dissimilar in the two jaws; the uppers slender, conical, widely spaced; the
lowers quadrate, each overlapping the next outermost, the cusps smooth edged and so
oblique that their inner margins form a continuous cutting edge, much as in Squalus and in
Centroscymnus; dermal denticles conical to thorn-like, curved rearward; rear end of base
of 1st dorsal far anterior to origin of pelvics; 2nd dorsal over or a little posterior to rear
end of bases of pelvics; 2nd dorsal only about as large as ist dorsal, but pelvics consider-
ably larger; interspace between ist and 2nd dorsals longer than between pelvics and
caudal; caudal very wide relative to its length, its lower anterior corner forming a more
or less definite lobe, its subterminal margin notched; pectorals with broadly rounded cor-
ners; with or without functional luminous organs (see p. 516). Development ovovi-
viparous in one species, perhaps oviparous in another. Characters otherwise those of the
family.

Range. Arctic Atlantic (including White Sea) south to North Sea, Portugal, Mediter-
ranean and Cape Cod; Bering Sea, in North Pacific, south to Japan, southeastern Alaska,
and occasionally southern California; also Antarctic (Maquarie Island).

9. Fowler and Ball (Bull. Bishop Mus., i6, 1926: 5, footnote) point out that specimens in the Bishop Museum on
which this record was based are actually /. brasiliensis.

I. This is a nomen nudem, see footnote 37, p. 523.

Fishes of the Western North Atlantic 515

Species. It seems sufficiently established that the Greenland Sharks of the sub-
Arctic on the two sides of the North Atlantic (including the White Sea) and of neighbor-
ing parts of the Arctic seas belong to a single species, described first by Gunnerus in 1766 as
Sgualus carcharias, but which, under the rules of nomenclature, must be called micro-
cephalusy Bloch and Schneider, 1801, the name Sgualus carcharias having been used previ-
ously by Linnaeus, 1758, for a very different shark (p. 142). It is still an open question
what the relationship is between microcephalus and the very much smaller form that has
been reported repeatedly from the Mediterranean and from the coast of Portugal as
rostratus. The difficulty, as is so often the case, is that the older portrayals differ widely
as regards relative locations of the fins and shape of the head. It is even possible that more
than one species may be included among the supposed rostratus.' But if the more recent
illustrations of rostratus can be accepted as reliable, it diflFers from microcephalus in a rela-
tively much shorter interspace between the tip of second dorsal and caudal, relatively
larger fins, smaller denticles, strongly developed luminous organs, and in various skeletal
characters;' it also attains maturity when much smaller, and, still more important, it is
ovoviviparous.*

The North Pacific representative of the genus has usually been considered identical
with the North Atlantic form. However, the first dorsal stands considerably farther rear-
ward in a Japanese specimen that we have examined' (as shown by Tanaka also") than is
ordinarily the case in Atlantic specimens, and its pectoral and caudal fins are larger; the
lower anterior and upper posterior margins of its caudal are much more strongly convex,
the distance from the tip of its second dorsal to the origin of its caudal shorter relatively,
its upper teeth are considerably broader, and the basal outlines of its lower teeth more
deeply incised. These differences seem sufficient to mark it off as a separate species. We
have therefore proposed the name pacificus for it.''

In all probability the Greenland Shark of Bering Sea and Alaska belongs to this
species. However, no detailed account of it has yet appeared, nor have we adequate mate-
rial for comparison.

The Antarctic representative of the genus, known from a single specimen only, has
also been made the basis of a separate species, antarcticus Whitley, 1939. Its first dorsal
appears to stand even farther forward than in microcephalus, and critical examination may
reveal additional differences.*

2. Brito Capello's measurements and illustration (J. Sci. math. phys. nat. Lisboa, 2, 1870: 141, pi. 9, fig. 2) repre-
sent both the snout and the caudal peduncle as much longer than in any other Somniosus.

3. Burckhardt, Ann. Mag. nat. Hist., (7) 6, 1900; 559; Helbing, Nova Acta Leop. Carol., Si, 1904: 347 ff., pi. 8, 9.

4. For a recent report of its embryo, see Borri (Atti Soc. tosc. Sci. nat., 44, 1934: loi).

5. This is the specimen illustrated by Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: pi. 15, fig. 1-3) as brevi-
finna.

6. Fish. Japan, 5, 191 1 : pi. 13, fig. 32-36.

7. Bigelow and Schroeder, Proc. New Engl. zool. CI., jj, 1944: 35.

8. See Whitley (Fish. Aust., /, 1940: 152) for illustration of the unique specimen.

5 1 6 Memoir Sears Foundation for Marine Research

Tentative Key to Northern Hemisphere Species

la. Origin of ist dorsal much nearer to tip of snout than to tip of caudal; interspace be-
tween 1st and 2nd dorsals at least as long as from snout to ist or 2nd gill opening.
2a. Distance from tip of 2nd dorsal to origin of caudal nearly or quite as long as
base of 2nd dorsal; base of ist dorsal only Yi to Y^ as long as distance from tip
of snout to origin of pectorals; no evident luminescent organs.

mkrocephalus Bloch and Schneider, i8or, p. 516.

2b. Distance from tip of 2nd dorsal to origin of caudal only about V2 as long as base

of 2nd dorsal; base of ist dorsal nearly or quite Yz as long as from tip of snout to

origin of pectoral; adult with luminescent dots. rostratus Risso, 1 826.

Mediterranean; probably
Portugal.

lb. Origin of ist dorsal almost as near to tip of caudal as to tip of snout; interspace between
1st and 2nd dorsals only about % as long as from tip of snout to 2nd gill opening.

pacificus Bigelowand Schroeder, 1944.

Japan, probably also Bering Sea and Alaska, south
to Puget Sound and occasionally to southern
California.

Somniosus mkrocephalus (Bloch and Schneider), 1801

Greenland Shark, Gurry Shark

Figures lOO, lOi

Study Material. Excellent mounted specimen, 10 feet long, from Upernavik, West
Greenland (Amer. Mus. Nat. Hist.) and old mount of a specimen 1,740 mm. (Harv.
Mus. Comp. Zool.) ; jaws of Gulf of Maine specimen, about 1 1 feet (Harv. Mus. Comp.
Zool.) ; lower jaw, from a Grand Banks specimen, taken at Lat. 43° 11' N., Long. 51° 22'
W. (U.S. Nat. Mus., No. 26270) ; excellent photographs of an i i-foot specimen;" also a
fresh-caught specimen of 10 to 11 feet from Cape Cod Bay examined by us in the flesh
some years ago.'"

Distinctive Characters. The adult Greenland Shark is so large that it could not be
confused with any other Atlantic squalid. Its lack of dorsal fin spines further marks it off
from the members of Squalus, Centra scyllium and the pigmy genus Etmopterus among
local genera. Its quadrate lower teeth with strongly oblique cusps separate it sharply from
Dalatias and Isistius, and its dermal armature sets it off from Centroscymnus (cf. Fig.
lOi B with 94 A). The forward position of its first dorsal, the shape of its teeth in general,
its dermal denticles, the shape of its caudal and its much smaller gill openings separate it
from Echinorhinus.

9. Contributed by F. E. Firth.

10. We have examined the Japanese specimen mentioned above, the lower jaw of one found on the beach near St.
Michaels, Alaska, and the head of another from California loaned by the U.S. National Museum,

Fishes of the Western North Atlantic 517

Description. Proportional dimensions in per cent of total length. Specimen, 2,990
mm. (mounted, Amer. Mus. Nat. Hist., New York).

Figure ioo. Somniosus microcefhdus. Side view of a female from a mounted specimen 1,740 mm. long
(Harv. Mus. Comp. Zool.), from photographs of a fresh ii-foot specimen and from other available in-
formation. A Upper and lower teeth from right-hand side, at center of mouth, from midway along the
jaws and from the corner of the mouth of a specimen about 1 1 feet long from the Gulf of Maine (Harv.
Mus. Comp. Zool., No. 361 19), about 1.8 x. B Front and side views of second upper tooth, about 3.5 x.
C Eleventh upper tooth, about 3.5 x.

Figure ioi. Somniosus microcefhalus. A Anterior part of head from below, obtained from available informa-
tion. B Dermal denticles from a mounted specimen, 1,740 mm. long (Harv. Mus. Comp. Zool.), about 1 5 x.

5 1 8 Memoir Sears Foundation for Marine Research

Trunk at origin of -pectoral: breadth 13.7; height 13.4.
Snout length in front of: outer nostrils 2. i ■■, mouth 8.3.
Eye: horizontal diameter 1.3.
Mouth: breadth 11.7; height 3.5.
Nostrils: distance between inner ends 6.3.
Labial furrow length from corner of mouth: upper 1.3.
Gill opening lengths: ist 2.2; 2nd 2.8; 3rd 2.8; 4th 2.8; fth 3.1.
First dorsal fin: vertical height 3.5; length of base 6.6.
Second dorsal fin: vertical height 2.5; length of base 4.9.
Caudal fin: upper margin 19.3; lower anterior margin 13.8.
Pectoral fin: outer margin 12. i; inner margin 6.S'-, distal margin 6.2.
Distance from snout to: ist dorsal 40. 7; 2nd dorsal 66.6; upper caudal 80.7; pec-
toral 23.7; pelvics 63.2.

Interspace between: ist and 2nd dorsals 19.5; 2nd dorsal and caudal 8.5.
Distance from, origin to origin of: pectoral and pelvics 39.5; pelvics and caudal
18.9.

Trunk subcylindrical anteriorly, but tapering posteriorly, its greatest height about
% of its length to origin of caudal, somewhat compressed, about % as thick as high oppo-
site the pectorals and 70 to 80% as thick as high at caudal peduncle, but so soft and flabby
that one lying on the dock bears little resemblance to its form when in the water." Back
smooth between dorsals, but with a faintly indicated dermal ridge along midline ex-
tending rearward from opposite pectorals about to origin of ist dorsal." Caudal pe-
duncle described as with an indistinct lateral longitudinal ridge, much as in Squalus, but
this is not visible in the mounted specimens we have examined. Dermal denticles conical,
curved rearward, longitudinally fluted, with high axial and lower lateral crests, their
bases quadrilateral, moderately closely spaced and of essentially the same form over the
trunk as a whole.

Snout thick, fleshy, broadly rounded in front, somewhat contracted between nostrils
and eyes, its length in front of mouth about Vs of length of head. Eye approximately
circular and very small, its diameter only about V5 to % of length of snout, its center about
opposite front of mouth. Spiracle a little above level of eye and behind latter by a distance
about 1V2 times the diameter of eye. Gill openings low on sides of neck, about evenly
spaced, very small and all about twice as long as diameter of eye, the 5th, slightly the
longest, close in front of origin of pectoral. Nostril nearly transverse and much closer to tip
of snout than to mouth, its anterior margin only slightly expanded. Mouth transverse,
very little arched, its breadth about equal to length of snout in front of mouth. Labial
furrows extending rearward for a considerable distance from corner of mouth and part
way along upper and lower jaws:"

11. For this reason many of the earlier representations of it are no better than caricatures, for they picture it as
enormously stout of body, which is not the case.

12. The exact extent cannot be stated for the Atlantic specimens studied because of their condition.

13. On the specimens examined, the lower labial furrow has been entirely obliterated in the process of mounting,
the upper and posterior folds mostly so as well.

Fishes of the JVestej'n Noyth Atlantic 519

Teeth f^ in specimen counted (up to j^ reported), widely unlike in the 2 jaws;
upper teeth thorn-like, tapering gradually to tip,'* moderately stout, very slightly
recurved and flattened anteriorly (thus subtriangular in cross-section); lower teeth sub-
quadrate, about Yi as broad as high, except that the outermost 3 or 4 are relatively much
broader and lower, each overlapping the next outermost, their cusps smooth-edged, deeply
notched outwardlv and so strongly oblique that inner margins are nearly parallel with the
jaw, forming a continuous saw-like cutting edge (much as in Centroscymnus) , the basal
sectors with a strong longitudinal ridge; about 3 series functional as a rule in upper jaw,
I or 2 series functional in lower, depending on the stage in their replacement."

Pectorals, dorsals and pelvics all very small. First dorsal brush-shaped, about V4 as
long at base as head, and about 3 to 4 times as long as high, its anterior edge very sloping,
apex rounded, posterior margin nearly straight or weakly concave, its free lower posterior
corner nearly or quite as long as base, its origin on mounted specimens about midway
between tip of snout and origin of caudaP" or posterior to axil of pectoral by a distance
I to 2 times as long as its own base. Interspace between ist and 2nd dorsals about as long
as from snout to ist or 2nd gill opening. Second dorsal nearly as long as ist at base, but only
about % as high vertically, with more broadly rounded apex, its origin over rear end of
bases of pelvics. Distance from tip of 2nd dorsal to origin of caudal about as long as base
of 2nd dorsal, or a little longer. Caudal Y^ to % of total length, its posterior margin
notched subterminally, about opposite end of caudal axis; its lower lobe about % as long
as upper lobe along upper margin of fin, the re-entrant contour included between the
two lobes considerably more obtuse than a right angle. Pelvics about as long at base as 2nd
dorsal or a little longer and about as high vertically, with broadly rounded apices and
tapering tips. Pectoral between Yz and Y2 as long as head, with nearly straight outer mar-
gin, moderately rounded tip, and much more broadly rounded inner corner.

Color. Described as varying in life between coffee brown or black and ashy-gray,
purplish-gray, or slaty-gray, below as well as above, changing to bluish-gray if the epider-
mis is rubbed off, as often happens after capture; the sides are sometimes tinged with
violet; the back and sides are crossed by numerous indistinct dark bands, at least in some
specimens, or with numerous indistinct whitish spots. Vestigial luminous dots, not func-
tional, are reported'' as scattered on the top and sides of the head, in a band along the
lateral line, and on the caudal.

Size. This is one of the larger sharks, and by far the largest of typically Arctic fishes.
It has been said to reach a length of 24 feet, but few, if any, actually grow to so great a
size, for the longest of which we find definite record'* was 21 feet, with specimens of 16 to
1 8 feet unusual, although occasionally taken. The largest western Atlantic specimens whose

14. They closely resemble those of Centroscytnnus in general appearance but are recognizable as Somnlosus by their
progressive taper; in Centroscymnus they are definitely lanceolate (cf. Fig. loo A-C with 94. B, C, H).

15. For discussion of this process in squalids with lower teeth of this type, see p. 65.

16. Garman (Mem. Harv. Mus. comp. Zool., j(5, 1913: pi. 15, fig. 4) so pictures it also for a Massachusetts speci-
men newly caught.

17. Burckhardt, Ann. Mag. nat. Hist., (4) 6, 1900: 562. 18. Jenkins, Fish. Brit. Isles, 1923= I^S-

520 Memoir Sears Foundation for Marine Research

lengths have been published are one of 16V2 feet from the Grand Banks in 1934, one of
about 16 feet off Portland, Maine, in 1846, another of about 15 feet off Cape Cod in 1849,
a fourth of about that same size caught in an otter trawl north of Cape Ann in February
193 1. Perhaps 8 to 14 feet is a fair average for adults, few among the hundreds annually
caught around Iceland and Greenland exceeding this size. Females average larger than
males, the maximum among 120 specimens brought into Aberdeen, Scotland, being il
feet 3 inches for the latter, and 15 feet 6 inches for the former. The British specimen of 21
feet, mentioned above, is said to have weighed about 2,250 pounds; an ii-foot specimen
from the Gulf of Maine, which we inspected, weighed about 650 pounds; another, of 11
feet 6 inches, taken off Cape Ann in January 1939, weighed about 600 pounds; but one
of 12Y2 feet, found alive on the beach in the estuary of the River Seine many years ago,
was only between 300 and 400 pounds. Males of about six feet are still immature; smaller
ones of either sex are seldom caught, but there is one record of a free-swimming specimen
of only one foot six inches (445 mm.)."

Developmental Stages. Adult females have been found repeatedly containing great
numbers of soft eggs without horny capsules, with up to as many as i Yo barrels of them
in large specimens, these eggs ranging in size up to that of a goose egg. This, combined
with the fact that none of the many examined have ever been found with embryos, supports
the general belief that this shark, unlike other squalids, is oviparous. If so, it seems likely
that the eggs are deposited on the bottom in mud, but eggs naturally laid have not been
found as yet.'° On the contrary, the Mediterranean Somniosus rostratus is ovoviviparous,
its embryos having been seen by several students.

Habits. Eyewitnesses agree that this is one of the most sluggish of sharks, offer-
ing no resistance whatever when hooked or even when drawn up out of the water. An
observer of long experience writes that he had driven a boat hook into one larger than
himself as it lay basking at the surface and had drawn it easily onto the ice." In view of this
passivity it is somewhat astonishing that it is able to capture prey as active as herring,
halibut, salmon, and seals which are said to become very scarce when these sharks gather.
Experience in the Iceland and Greenland fisheries indicates that they usually lie close to
bottom in summer but often swim toward the surface for prey, even in the warm season,
and in the winter fishery through the ice of West Greenland they are often lured to the
surface by a light.

The diet of the Greenland Shark includes a wide variety of fishes, both large and
small, such as skates, herring, salmon, capelin (Mallotus), rosefish (Sebastes), sculpin
(Myoxocephalus), lumpfish (Cyclopterus), saithe or American pollock {Pollachius
'i!irens),Ymg (Molva), cod, haddock, wolffish (Anarrhichas) , znd various flatfishes, among
them halibut (Hippoglossus) and the Greenland halibut (Reinhardtius). Seals are a

19. Winther, Prod. Ichthyol. Dan. Mar., 1879: 59.

20. For a summary of evidence on this subject, see Liitken (Vidensk. Medd. naturh. Foren. Kbh. [1S79], 'SSo: 56).

21. Grenfell, Labrador, 1910: 351.

Fishes of the Western North Atlantic 521

favorite food as well as small cetaceans, the latter perhaps dead when eaten. Sometimes
sea birds are captured, and squids, crabs, large snails, and even medusae are devoured.
Objects as large as an entire reindeer (without horns), a whole seal, a four-foot ling
(Moha), a three-foot cod, and a 39-inch salmon have been found in stomachs of the
Greenland Shark, which gives some measure of its appetite. They also greedily devour
any carrion, such as whale meat, blubber, etc., from whaling operations, and their habit
of gathering around whaling stations for this purpose, or when there has been a big killing
of narwhals in Greenland waters, is proverbial. Similarly, large numbers are described as
haunting the ice fields in spring off the Labrador coast, where sealers have left the carcasses
of young seals. But there appears to be no basis for the old story that they attack living
whales.

Its depth range is wide. In its centers of abundance it tends to approach the surface
in winter, coming right up to the ice o£F Greenland and along the Labrador coast. In
summer, however, it is most often caught at 100 to 300 fathoms, and has been recorded
as deep as 660 fathoms. Although it usually lies close to the bottom during the warm
season on the Labrador coast, it often becomes entangled in seal nets even then. Its habit
of gathering when whales are being cut up was well known during the days of the Arctic
Right Whale Fishery. The frequency with which the remains of seals and sea birds are
found in its stomach is further evidence of its readiness to swim upward in pursuit of prey.
The considerable number that are taken in the North Sea are all caught shoaler than lOO
fathoms, irrespective of the season, which applies equally to most of the Gulf of Maine
records."

It has been taken in water as cold as minus 0.6° C," and it is the only shark regularly
inhabiting polar temperatures. At the other extreme it is able not only to survive but to
feed actively in water at least as warm as 10° to 1 2° C, as indicated by the repeated capture
of specimens in the northern part of the North Sea, their stomachs full of recently eaten
fish. In the Gulf of Maine, too, it has been taken in water as warm as about 10° C."* But
most of the local records have been based on specimens taken when the water temperatures
were between 2° and about 7° at the bottom.

Relation to Man. In North American waters the Greenland Shark is of no commer-
cial value. Off northern Norway, however, around Iceland and in West Greenland waters
it has long been sought regularly. By the middle eighteen-hundreds the catch off West
Greenland was 2,000 to 3,000 sharks yearly, which had risen to 11,000 to 15,000 by
the eighteen-nineties, and to upwards of 30,000 by the first decade of the present cen-
tury. The catch is obtained by hand lines, or on long lines, for the most part in depths of
100 to 200 fathoms, except along the northern part of the West Greenland coast, where

22. Stray specimens have been taken in a few feet of water near the southern boundary of its range, or found
stranded on the beach.

23. Murray and Hjort, Depths of Ocean, 1912: 436.

24. Taken in a weir during summer or early autumn in Passamaquoddy Bay.

522 Memoir Sears Foundation for Marine Research

they are caught chiefly in winter through holes in the ice, close to the surface, either on
hook and line, or sometimes even with short-handled gaffs."

They are utilized chiefly for their liver oil, a large specimen sometimes yielding as
much as 30 gallons or more (up to about 50% of the volume of the liver). In Greenland
the flesh is also dried for dogfood, and in Iceland small amounts have been consumed for
human food in the past. But it produces a sort of intoxicant poisoning if eaten fresh,
whether by man or by dogs," although it is wholesome when dried. Fishermen regard
them as harmless; old tales that they attack Greenlanders in their kayaks appear to be
mythical.

Range. White Sea, Spitzbergen, Bear Island, Norwegian Sea, East and West Green-
land, south regularly to the North Sea and Kattegat, less commonly to the southern part
of the North Sea, accidentally to the mouth of the Seine (one specimen) and perhaps to
Portugal, in the eastern Atlantic; regularly to the northern part of the Gulf of St. Law-
rence, in the western Atlantic, and less commonly to the Gulf of Maine. Whether its range
extends along the Arctic coasts of Eurasia, or to the coasts of Arctic North America west-
ward from Baffins Bay, is not known. It is represented in the North Pacific by a form
which, while close, appears to be distinct (for discussion, see p. 515).

Occurrence in the Western Atlantic. This shark is so plentiful along the Greenland
side of Davis Strait and in Bafiins Bay that in the first decade of the present century the
yearly catch there was around 32,000." Similarly, during fishing experiments with long
lines off West Greenland by the "Ingolf" Expedition at least 20 per cent of the hooks
were bitten off by them. They have been reported from Hudson Strait"* and are plentiful
along the east coast of Labrador, where specimens are often entangled in seal nets and
where a recent author also reports catching five through one hole in the ice."

No doubt its normal range includes the outer coast of Newfoundland, although we
find no definite statement as to its numbers, there being only one published record for the
Grand Banks.^° It is sufficiently numerous to be a nuisance to fishermen in the Straits of
Belle Isle, at least in some years, and inward along the north shore of the Gulf of St. Law-
rence;^' specimens have even been reported from the Saguenay River^' and from the
lower reaches of the St. Lawrence.'^ But while it undoubtedly occurs on the southern side
of the Gulf as well, it is at least not numerous enough there for its presence to have occa-

25. For a description of the West Greenland fishery, see especially Jensen (Mindeskr. Jap. Steensfrup. Ffids., Kbh.,

J [30], 1914: 15).
z6. For accounts, see Jensen (Mindeskr. Jap. Steenstrup. F0ds., Kbh., 2 [30], 1914: 12) and Clark (Science, N. S.

i', 1915: 795)-

27. Jensen, Mindeskr. Jap. Steenstrup. F0ds., Kbh., 2 (30), 191+: 9.

28. Vladykov, Contr. Canad. Biol., N. S. 8 (2), 1933: 5.

29. Grenfell, Labrador, 19 10: 351.

30. Rep. Newfoundland Div. Fisher. Res., Fisher Res. Lab. (1934), 1935; 79.

31. Stearns, Proc. U.S. nat. Mus., 6, 1883 : 123-1 ;,.

32. Whiteaves, Cat. Canad. Pinnep. Cetacea, Fish., i886: 4.

33. At Metis; Dawson, Canad. Rec. Sci., 4, 1S91 : 304.

Fishes of the Western North Atlantic 523

sioned any printed comment; we find but two records from the coast of Nova Scotia, one
being from Cape Breton and the other near Halifax."

In view of its apparent scarcity in Nova Scotian waters, it is somewhat astonishing
that there is published record of about 27 specimens in the Gulf of Maine up to 1938, with
several more reported subsequently by local fishermen. The localities include Passama-
quoddy Bay (tributary to the Bay of Fundy), off Eastport, Portland and Cape Elizabeth,
Maine; Jeffrey's Ledge,'^ inner part of Massachusetts Bay, Cape Cod Bay, tip of Cape
Cod, and the southwestern part of the Gulf of Maine basin. This distribution is wide
enough to show that odd specimens are to be expected anywhere in the western side of the
Gulf at any time of year. In fact, it is rumored that in early colonial times, when Atlantic
Right Whales were still being killed in numbers off the Massachusetts coast, the Green-
land Sharks were more abundant there than they have been at any time during the last
hundred years. None have been reported either from the Nova Scotian side of the Gulf
on the one hand, although this lies in their route from the north, or westward from Cape
Cod on the other; but recorded captures in the Gulf of Maine include both small and
large specimens (five small ones, from only 39 inches up to four to five feet long, off
Portland between 1925 and 1933) ; furthermore, they have been recorded for November,
January, February, March, April, June and August; these facts suggest that the partial
enclosure of the comparatively deep western waters of the Gulf by the shoaler banks to
the south forms a sort of cul-de-sac for any that may stray that far. Once arrived, they
may survive there for years.

Synonyms and References:

Haa-Skierding, Gunnerus, Trondh. Selsk. Skrift., 2, 1763: 330, pi. 10, II (size, food, descr., Norway).

Squdus carcharias Gunnerus, Drontheim Gesellsch. Schr., 2, 1766: 299, pi. 10, 11 (size, food, descr., Nor-
way) ; Muller, Prod. Fauna Danica, 1776: 38 (Denmark) ; Fabricius, Fauna GrocnI., 1780: 127 (general
account, food, abund. W. Greenland) ; not Squalus carcharias Linnaeus, I 758.

Haa-Kiacrringen, Rosted, K. norske \'idcnsk.-Selsk. Skr., N. S. 2, 1788: 203, I pi. (fishery, Norway).

Squalus mkrocefhalus Bloch and Schneider, Syst. Ichthyol., 1801: 135 (refs., descr., Arctic Seas); Blain-
ville, in Vieillot, Faune Franc, 1825: 66 (ref. to Bloch and Schneider, 1801).

Squalus squatina Pallas, Zoogr. Rosso .^siat., 5, 1814:^^ 64 (White and Arctic Seas) ; not Squalus squalina Lin-
naeus, 1758.

Acanthorhinus TtorzL-egianus Blainville, Bull. Soc. philom. Paris, 8, I 816: 12 1.^'

Squalus brevifinna Lesueur, J. Acad. nat. Sci. Philad., /, 1818: plate facing p. 222 (ill., spec, from Massa-
chusetts).

Somniosus brez-ipnna Lesueur, J. Acad. nat. Sci. Philad., i, I 81 8: 222 (descr. of spec. ill. as Squalus brevi-
pmia, Marblehead, Massachusetts); Bory de St. Vincent, Diet. Class. Hist. Nat., 75, 1829: 597 (ref.,
Massachusetts); Storer, Rep. Fish. Rept. Birds Mass., 1839: 189 (Massachusetts); Boston J. nat. Hist.,
2| 1839: 541 (descr., Massachusetts); Gill, Proc. Acad. nat. Sci. Philad., 1863: 333 (.Massachusetts);

34. Storer, Boston J. Nat. Hist., 6, 1857: 270; Jones, Proc. N. S. Inst. Sci., 5 (i), 1882: 96.

35. Six specimens, which ranged in length from 39 inches upward and were taken in the months of January, Febru-
ary', April, June and .•\ugust, have been reported to us by W. W. Rich for this general region since 1925.

36. It is generally accepted that 18 14 is the date of publication of the part of volume 5 in question; see Cat. Library
Brit. Mus., page 1505.

37. Name only, but identification probable by Inference.

524 Memoir Sears Foundation for Marine Research

Garman, Mem. Harv. Mus. comp. ZooL, 56, 1913: 240, in part (Atlant. refs., but descr. and il!., pi. 15,
fig. 1-3, are of a Japanese spec.) ; Rey, Fauna Iberica Feces, /, 1928: 479 (in Key) ; Jordan, Manual
Vert. Anim. NE. U.S., 1929: 14 (part) ; Tortonese, Atti Soc. ital. Sci. nat., 77, 1938: 312 (concludes that
brevi-pititia is distinct from microcefhalui).

Squalus horealis Scoresby, Arctic Regions, /, 1820: 538, pi. 15, fig. 3, 4, 5 (descr., habits, size, Spitzbergen
Sea); Cordcau.x, Zoologist, (2) i, 1866: 230 (Dogger Bank, North Sea, stomach contents); Southwell,
Zoologist, (2) 10, 1875: 4424 (England).

Squalus norvegiatius Blainvillc, in Vieillot, Fnune Franc, 1825: 61 (refs.).

Scymnus borealis Fleming, Hist. Brit. Anim., 1828: 166 (descr.) ; Jenyns, Manual Brit. Vert. Anim., 1835:
506 (Scotland) ; Swainson, Nat. Hist. Fish. Amphib. Rept., /, 1839: 145; 2, 1839: 315 (general) ; Ham-
ilton, Brit. Fish., 2, 1843: 315, also subsequent eds. (Gt. Brit.) ; Busch, Selach. Ganoid. Encephal., 1848:
pi. 3, fig. 7 (brain); Smith, Zoologist, 9, 1851: 3058 (Scotland); Baikle, Zoologist, //, 1853: 3846
(Orkneys); Van der Hoeven, Handb. Dierkunde, (2) 2, 1855: 26 1 (general); Nilsson, Skand. Fauna,

4, Fisk., 1855: 724 (Greenland, Iceland, Scand., Spitzbergen); Thompson, Nat. Hist. Ireland, ^,
l8;6: 255 (Ireland); Brown, Zoologist, 18, i860: 6861 (Scotland); Fiedler, Tidskr. Fiskeri, Jahrg.
1866: 26-33 (Denmark, not seen); Saxby, Zoologist, (2) (5, 1871: 2553 (Newfoundland); Trail,
Scot. Nat., 7, 1872: 48 (numbers, size, Scotland).

ScymNU! gU7i7ieri Thienemann, Lehrb. ZooL, j, 1828: 409.

Scymnus glacialis Faber, Fische Islands, 1829: 23 (habits, eggs, food, fishery, Iceland, Polar Sea); Van Bene-

dcn, Sur Sciaena aqu'tla et Scymnus ghcialiSy 1852 (not seen).
Scymnus microfterus Valenciennes, Ann. Mus. Hist. nat. Paris, r, 1832: 458, pi. 20 (descr., weight, 13-foot

spec, estuary of the Seine).
Squalus glacialis Nilsson, Prod. Ichthyol. Skand., 1 832: 115 (Norway); Lilljeborg, Vet. Akad. Handl.

Stockholm, 2, 1850: 334 (not seen).
Greenland Shark, Yarrell, Brit. Fish., 2, 1836: 403 (descr., Scotland); Couch, Fish. Brit. Isles, i, 1867: 57,

pi. 13 (descr., habits, ill., Gt. Brit.); Buckland, Hist. Brit. Fish., 1881: 213 (Scotland, Iceland, habits,

fishery) ; Norman and Eraser, Giant Fishes, 1937: 38 (general, habits).
Squalus {Scymnus) gunneri Richardson, Fauna Boreal. Amer., j, 1836: 313 (Greenland Seas).
Laemargus carcharias Bonaparte, Mem. Soc. neuchatel. Sci. nat., 2 (8), 1839: 9 (in synopsis).
Scymnus (Laemargus) borealis Miiller and Henle, Plagiost., 1841: 93 (descr., refs.); Gaimard, Voy. Islande

et Groenland (1835-1836), Atlas 5, Zool. Med., 1851: pi. 22 (good ill.); Dumeril, Hist. Nat.

Polss., I, 1865: 456, pi. ;, fig. 3, 4 (refs., descr., comp. with brevifinna, ill. of teeth).
Scymnus brevifinna DeKay, Zool. N. Y., 4, 1842: 361, pi. 61, fig. 202 (descr., ill. after Lesueur, 1818,

Massachusetts); Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 507 (in synopsis); Kneeland,

Boston J. nat. Hist., 5, 1847: 479 (anat., Massachusetts); Storer, Boston J. nat. Hist., 6, 1857: 270

(jaw, C. Breton); DeKay, Rep. St. Cab. nat. Hist. N. Y. (1835), S, 1858: 64; Storer, Mem. Amer.

Acad. Arts Sci., N. S. 9, 1867: 235, pi. 38, fig. 2; Fishes Mass., 1867: 259, pi. 38, fig. 2 (descr., orig.

ill., Massachusetts records).
Laemargus borealis Bonaparte, Cat. Pesc. Europ., 1846: 16 (in Europ. list); Giinther, Cat. Fish. Brit. Mus.,

5, 1870: 426 (refs., descr., Brit, spec); Turner, J. Anat. Lond., 7, 1873: 233 (size, anat., Scotland,
North Sea); J. Anat. Lond., 8, 1874: 285 (anat., stomach contents, meas.) ; Gervais and Boulart, Poiss.,
5, 1877: 212, pi. 82 (descr., ill.); Turner, J. Anat. Lond., 12, 1878: 604 (corrects earlier statements
that oviducts were lacking); Jones, List Fish. Nova Scotia, 1879: 96 (Nova Scotia record, meas.);
Hassc, Naturl. Syst. Elasm. besond. Theil, j, 1882: 55, pi. 8, fig. 1-6 (vert., dermal denticles); Jones,
Proc. N. S. Inst. Sci., 5 (i), 1882: 96 (same as Jones, 1879); Turner, J. Anat. Lond., ig, 1885: 221
(anat., stomach contents, Scottish spec.) ; Dawson, Canad. Rec. Sci., ^, 1 891: 304, pi. 4 (ill., meas., size,
lower St. Lawrence R.) ; Dean, Fishes Living and Extinct, 1895: 91 (general) ; Burckhardt, Anru Mag.
nat. Hist., (7) 6, 1900: 562 (vestigial luminous organs); Helbing, Nova Acta Leop. Carol., 82, 1904:
351, 364, 377, 394, 421, 441, 450, 461, 475, 491, 511, pi. 8, fig. 2B, pi. 9, fig. B (skeleton);
Giinther, Encyc. Brit., llth ed., 24, 191 1: 808 (general); Ehrenbaum, in Grimpe and Wagler, Tier-
welt N.- u. Ostsee, Lief 7 (12^), 1927: 19 (general. North Sea).

Leiodon echinatum Wood, Proc. Boston Soc. nat. Hist., 2, 1846: 174 (descr. of 16-foot spec, off Portland,
Maine).

Fishes of the Western North Atlantic 525

Datatias (Somniosus) borealis Gny, List. Fish. Brit. Mus., /, 1 851: 76.

Dalatias microre^halus W'Kwi^, List Spec. Brit. Mus., Fish., 8, 1851: 132 (Scotland, England).

Scimnus g/ac-alis Vsn Benedcn, Bull. Acad. Sci. R. Bruxelles, 20 (2), 1853: 258 (parasites, Belgium) ; Bull.
Acad. Sci. R. Bruxelles, (2) I, 1857: 226 (parasites).

Scymnus mkrocefhalus Krcivcr, Danmnrlcs Flsk., 5 (2), 18,3: 914 (descr., meas., Denmark); Malmgren,
Oev. SvensL Vet. Akad. Forhand. Stockholm (1864), 186;: 536 (N. of Norway); Collett, Forh.
Vidensk-Selsk. Krist., 1874: 212 (fishery, depths, Norwegian Iocs.) ; Norges Fisk., 1875: 208 (Norway) ;
Malm, Goteborgs och Bohuslans Fauna, 1877: 626 (Kattegat); Lutken, Dyreriget (Laer'oog Zool.),
/, 1881-1882: 379 (Iceland, Greenland, Spitzbergen, fishery); Storm, K. norske Vidensk.-Selsk. Skr.,
1884: 44 (Trondhjem Fjord).

Squolus nortegicus Gray, Cat. Fish. Coll. Descr. by L. T. Gronow, in Brit. Mus., 1854: 8.

Laemargus microccfhalus Malmgren, Finland Fisk. Fauna, 1, 1863: 71 (N. Norway, not seen); Arch.
Naturgesch., jo (1), 1864: 34; (transl. of foregoing by Frisch; N. Norway, Greenland, Iceland) ; Win-
ther, Naturh. Tidskr. Copenhagen, 72, 1879: 59 (Danish Iocs., Kattegat, size); Day, Fish. Gt. Brit., 2,
1880-1884: 320, pi. 162, fig. 1 (genl. account, Gt. Brit.); Calderwood, Rep. Fish. Bd. Scot., .App. F,
7(5, 1886: 228, pi. 10 (Scotland, size, eggs, stomach contents, numbers) ; Mobius, S. B. Ges. naturf. Fr.
Berlin, 1890: 11 (color, size, Helgoland); White, P. J., Trans, roy. Soc. Edinb., 57 (2), 1893: 287,
2 pis. (skull and visceral skelet.) ; Jenkins, Fish. Brit. Isles, T925: 32;, pi. 12; (general, size, Gt. Brit.);
Beaufort, Tijdschr. ned. dierk. Ver., 191 1, (2) 2, iginLXVIII (Holland).

Somniosus m-.cTocefha!us Gill. Proc. Acid. nat. Sci. Philad., .Addend., 1861 : 60 (listed) ; Proc. Acad. nat. Sci.
Philad., 1864: 264 (name); Rink, Danish Greenland, 1877: 131 (abund., habits, toxicity of flesh, Green-
land); Goode and Bean, Bull. Essex Inst. Salem, 11, 1879: 31 (off Massachusetts); Lutken, \'iJensk.
Medd. naturh. Foren. Kbh., 1 880: 56 (eggs, reproduction probably oviparous) ; Jordan and Gilbert, Bull.
U.S. nat. Mus., 16, 1883: 15 (descr., C. Cod to Arctic seas) ; Stearns, Proc. U.S. nat. Mus., <5, 1883: 125
(Gulf of St. Lawrence, Labrador) ; Petersen, Vidensk. Mcdd. naturh. Foren. Kbh. (1884-1886), 1887:
160 (Denmark); Collett, Nyt Mag. Naturv., 29, 188;: 117 (Banks off Norway); Kingsley, Stand.
Nat. Hist., 5, 1885: 76 (general); Whiteaves, Cat. Canad. Pinnep. Cetacea, Fish., 1886: 4 (weight,
Saguenay R. spec); Jordan, Rep. U.S. Comm. Fish. (188;), 1887: 793 (distrib.) ; Holm, Medd.
GroenLind, 2, 1888: 81 (E. Greenland); Packard, L.ibrador Coast, 1891: 397 (Gulf of St. Law-
rence, Labrador); Goode and Bean, Smithson. Contr. Knowl., 50, 1895: 7, 507, pi. 3, fig. 8; Mem.
Harv. Mus. comp. Zool., 22, l8g6 (descr., ill., distrib.); Jordan and Evermann, Rep. U.S. Comm.
Fish. (1895), 1896: 219 (distrib.); Bull. U.S. nat. Mus., 47 (1), 1896: 51 (refs., descr., distrib.);
Vanhoffen, in Drygalski, Griinl. Exped. Gesellsch. Erkundc, Berlin, 2 (l), 1S9-: 124 (N. Green-
land) ; Jungersen, Danish "Ingolf" Exped., 2, 1899: I, pi. I, fig. I-9 (anat., off W. Greenland) ; Jensen,
Medd. Greenland, 29, 1904: 276 (abund., E. Greenland); Collett, Vidensk. Selsk. Foren. Christiania,
7, 1905: 100 (Norway) ; Medd. Norges Fiske, 5, 1905: 100 (food, size, eggs, Norway) ; Jordan, Guide
to Study Fish., 1905: 547 (in part, but incl. Pacif. form also); Kendall, Occ. Pap. Boston Soc. nat.
Hist., 7 (8), 1908: 9 (N. Engl, records); Regan, Ann. Mag. nat. Hist., (8) 2, 1908: 54 (class.,
distrib.); Saemundsson, Skr. Komm. Havunders. Kbh., ;. IQ09: 119 (eggs, sizes, depths, numbers,
Iceland) ; Grenfell, Labrador, 1910: 350 (abund., captures, yield of oil, Labrador) ; Berg, Faune Russie
(Russian), /, 1911: 78 (Murman Co.ast, White Sea, but Bering I. ref. probably for facificus) ; Murray
and Hjort, Depths of Ocean, 191 2: 436 (Norwegian Sea) ; Halkett, Check List Fish. Canad., 1913: 44
(Labrador) ; Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 241, pi. 13, fig. 4-6 (refs., descr., ill.,
Massachusetts); Jensen, Mindeskr. Jap. Steenstrup. F0ds., Kbh., 2 (30), 1914: 8 (abund., food, devel.
fishery, Greenland); Clark, Science, N. S. 41, 1915: 795 (toxicity of flesh); Smith, J. .Amer. Mus.
nat. Hist., 16, 1916: 349 (general); Huntsman, Contr. Canad. Biol. (1921), 1922: 56 (Bay of
Fundy, Passamaquoddy Bay); Wollebaek, Norges Fisk., 1924: 24 (Norway, not seen); Bigelow and
Welsh, Bull. U.S. Bur. Fish., 40 (i), 1925: 53, 551 (descr., habits. Gulf of Maine); Causev, Para-
sitolog)', 18, 1926: 195 (parasites); Raven, Nat. Hist. N. Y., 26 (6), 1926: 654 (color, food, cap-
tures, W. Greenland); Hickling, Ann. Mag. nat. Hist., (lo) 2, 1928: 197, 199 (Faroe-Shetland
Channel) ; Rey, Fauna Iberica Feces, /, 1928: 480 (descr. after Garman, 191 3, habits, comp. with other
species); Breder, Field Bk. Mar. Fish. AtJant. Coast, 1929: 25 (general); Jordan, Manual Vert.

526 Memoir Sears Foundation for Marine Research

Anim. NE. U.S., 1929: 14 (general); Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928),
2, 1930: 22 (in check list for N. Amer., N. Pacif. form also included) ; ^'ladykov, Contr. Canad. Biol.,
N. S. S (2), 1933: 5 (Port Burwell, Hudson Strait); Proc. Pan-Pacif. sci. Congr., 5, 1934: 3787, 3788
(Hudson Bay region); Bigelow and Schroeder, Canad. Atlant. Fauna, bid. Bd. Canad., 12', 1934: 19
(dcscr., range); Brooks, Parasitology, 26, 1934: 259 (parasites); Piers, Proc. N. S. Inst. Sci., 18, 1934:
203 (ref. to old Nova Scotia record by Jones, 1882); Vladykov and McKenzie, Proc. N. S. Inst. Sci.,
'9> '935- 49 (°1'1 Nova Scotia record); Rep. Newfoundland Fish. Res. Lab. (1934), 1935: 79
(165/2-ft. spec, Grand Banks); Bigelow and Schroeder, Bull. U.S. Bur. Fish., 48, 1936: 323 (addit.
Gulf of Maine records); Liibbert and Ehrenbaum, Handb. Seefisch. Nordeurop., 2, 1936: 288 (size,
depths, fishery, N. Europ.) ; White, Bull. Amer. Mus. nat. Hist., 7^, 1937: 43 (ill. after Garman,
191 3); MacCoy, New Eng. Nat., 2, March 1939: 25 (photos, stomach contents of ll-ft. spec. Gulf
of Maine) ; Fowler, Bull. U.S. nat. Mus., loo (13), 1941 : 276 (refs., descr., but N. Pacif. form also in-
cluded) ; Holmgren, Acta Zool., Haft. 1-3, 1941: 28 (skull).

Scymnus (Laemargus) brevifinna Dumeril, Hist. Nat. Poiss., i, 1865: 456 (descr., ref. to old record for
R. Seine estuary).

Scimrius borealis Van Beneden, P. J., Poiss. Cotes Belg. Parasit., 1870: 10 (parasites, Belgium).

Laemargus brevifinna Moreau, Nat. Hist. Poiss. France, j, I 881 : 361 (descr., ref. to old record for estuary of
R. Seine).

Somniosus carcharias Garman, Bull. Mus. comp. Zool. Harv., 77, 1888: 85, pi. 20 (mucous canals, no loc,
but probably Massachusetts).

Laemargus (no specific name) Ewart, Trans, roy. Soc. Edinb., 57 (i), 1893: 59, pi. 1, 2 (sensory canals).

Acanthorhinus carcharias Smitt, Hist. Skand. Fish., 2, 1895: 1 1 67, pi. 52, fig. 3 (descr., habits, fishery,
Skand.); Knipowich, Annu. Mus. zool. Acad. St. Petersb., 2, 1897: 154 (Murman Coast, White
Sea); Ehrenbaum, Faune IchthyoL, Cons. Explor. Mer., 1930: plate not numbered (descr., ill., distrib.
in E. Atlant.).

Family ECHINORHINIDAE

Characters. Squaloidea without dorsal fin spines, the teeth with several cusps and
similar in the two jaws.

Genera. One genus, Echinorhinus. It is doubtful whether the characters forming the
basis for the subgenus Rubusqualus Whitley, 1931,' are even of specific value.

Genus Echinorhinus Blainville, 18 16

Echinorhinus Blainville, Bull. Soc. philom. Paris, 1 8 16: 121 ; type species, E. sfinosus (no authority quoted,
but doubtless Gmelin, 1789), equals Squalus brucus Bonnaterre, I 788.

Generic Synonyms:

Scymnus (in part) Cuvier, Regne Anim., 2, 1817: 131, footnote.

Goniodus L. Agassiz, Poiss. Foss., ^, 1838: 94; same as Echinorhinus Blainville, 1816.

Centrofhorus (in part) Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1839: 315 (incl. sfinosus") ; not Centro-

fhorus Muller and Henle, 1837.
Rubusqualus (subgenus) Whitley, Aust. Zool., d, 1 931: 31 1; type species, Echinorhinus sfinosus McCoy,

Australia.

Generic Characters. Teeth with 3 to 7 cusps, the median much the largest and so
strongly oblique that the inner margins form a continuous cutting edge along the jaw 5 der-
mal denticles in the form of tubercles or shields with small central spines, scattered singly

I. Whitley, Aust. Zool., (i, 1931: 311; eye over anterior mouth, heavier tail, dorsal fins closer together and first
dorsal originating over the anterior part of the "anal" (sic), instead of over the middle of that fin.

Fishes of the Western North Atlantic 527

or in groups and varying greatly in size ; a dermal ridge above and one below the lateral
line, posterior to gill openings; caudal without precaudal pits; spiracles minute; a labial
furrow on each jaw and voluminous pit at each corner of mouth; origin of ist dorsal over
bases of pelvics, and far behind midlength of trunk; caudal without subterminal notch, its
lower anterior corner not expanded as a definite lobe; gill openings much larger than is
usual in the suborder.

Range. Eastern Atlantic, from Ireland to tropical South Africa, including the Medi-
terranean; Argentina; California; Japan; Hawaiian Islands; New Zealand; Australia
and Tasmania; Arabia; accidental in western North Atlantic.

Fossil. Miocene, North America; Pliocene, Europe.

Species. The South African representatives of the genus, as well as the Australian-
New Zealand and Hawaiian representatives, have all been given separate names as sup-
posedly distinct from E. brucus of the North Atlantic. By common consent, however, the
first of them {E. obesus Smith, 1 849) has been relegated to the synonymy of brucus. Simi-
larly, it has been held recently^ that the Hawaiian cookei Pietschmann, 1928, is merely a
variant of brucus. A Californian specimen, recently taken, proved to be a typical brucus ^"^
and it is at least questionable whether the features that are believed by its author to dis-
tinguish the Australian-New Zealand mccoyi Whitley, 193 1, from E. brucus, represent
anything more than individual variations.' Final conclusions must await critical compari-
son of adequate series of specimens, however. References for the several geographic re-
gions are therefore segregated in the accompanying synonymy (p. 530).

Echinorhinus brucus (Bonnaterre), 1788
Spiny Shark

Figure 102

Study Material. None.

Distinctive Characters. The following combination makes this Shark easily recog-
nizable among local Squaloidea, should one be taken in the western side of the North
Atlantic: dorsal fins without spines, teeth with several cusps in each jaw but so oblique
as to form a nearly continuous cutting edge, very large gill openings and peculiar, shield-
like dermal denticles.

Description.* Trunk subcylindrical, moderately stout. Lateral line lying in a well
marked furrow, rearward from opposite 5th gill opening, flanked above and below by a
pair of thin, palisade-like dermal ridges, sparsely fringed with small fleshy papillae. Der-
mal denticles in the form of flat shields, varying greatly in diameter, each with a more or
less strongly developed conical, sharp-pointed spine in the center, sometimes 2 spines, from
which numerous furrows radiate outward to the margin so that the latter is more or less

2. Fowler, Bull. U.S. nat. Mus., loo (/j), 1941: 278; Hubbs and Clark, Calif. Fish Game, j/, 1945 : 65.
2a. Hubbs and Clark, Calif. Fish Game, J 7, 1945: 65. 3. Whitley, Aust. Zool., fi, 1931 : 31 1.

4. Based on published accounts and illustrations.

528

Memoir Sears Foundation for Marine Research

denticulate (the larger scales resemble more the bucklers of certain skates than those of
other sharks), irregularly distributed, either singly and wide-spaced or in groups of 3 to 5,
in which case they may be so closely crowded that their circular outlines are more or less
lost, or the adjoining denticles may even be more or less fused.

Figure 102. A, Echinorhinus briicus, eastern Atlantic specimen, about three feet long, in British Museum.
B Head of same from below. C Dermal denticles. D Upper and lower teeth a little longer than natural size.
E Third upper tooth, about z x. F Upper and lower teeth, after L. Agassiz.

Head flattened above. Snout ovate, tapering from eyes. Eye opposite front of mouth,
approximately circular, its horizontal diameter Yz to V2 as long as snout in front of mouth.
Spiracle posterior to eye by a distance a little longer than diameter of latter. Gill openings
slightly oblique, the 5th about twice as long as ist or about as long as snout in front of
mouth and more than twice as long as diameter of eye (thus much larger than in any other
member of the suborder). Nostril about midway between tip of snout and corner of mouth,
its anterior margin with a pointed lobe. Mouth crescentic, about V2 as high as broad (thus
more strongly arched than in other local Squaloidea). Labial furrows confined to corners
of mouth.

Teeth ^°|°^g, alike in the 2 jaws, each with a pointed median cusp usually flanked by
I small cusp on the inner side and 2 on the outer though described as sometimes*' with-

4a. Rey, Fauna Iberica, Feces, i, 1928: 485.

Fishes of the Western North Atlantic 529

out denticles, and so strongly oblique that their inner margins form a cutting edge, but
with the teeth separated by distinct interspaces.

Dorsal fins very small, brush-shaped, with broadly rounded corners, the origin of ist
over or slightly anterior to anterior third of bases of pelvics. Second dorsal a little smaller
than ist, its origin pictured as varying from over or a little posterior to rear ends of bases
of pelvics to over their rear tips. Interspace between ist and 2nd dorsals pictured for Euro-
pean specimens as varying from as long as base of ist dorsal to so short as to be hardly
discernible. Caudal extremely characteristic, being broadly scythe-shaped with tapering
tip, its posterior contour evenly concave without definite subterminal notch, its lower in-
ferior corner broadly rounded. Pelvics much larger than dorsals, their bases nearly or quite
twice as long as bases of latter, with rounded apices and tapering rear corners. Pectoral a
little less than V2 as long as head, brush-shaped, with weakly convex outer margin, broadly
rounded corners and notably broad base.

Color. This is variously described as dark gray, dull olive or brown above, with
reflections of violet, silver, gold or coppery yellow, and with or without obscure darker
blackish or reddish blotches 5 paler brown or gray to white below. The scales have iaeen
described as luminescent,' but there are no special luminous organs.*

Size. The smallest European specimens on record were about three feet in length, the
largest about nine feet} the majority of measurements available have ranged between
approximately 5 feet and 8 V2 feet. A specimen of eight feet four inches weighed about
300 pounds, and an eight-foot five-inch example of the New Zealand form about 350
pounds. Females appear to average larger than males.

Developmental Stages. It is not known whether the development is viviparous or
ovoviviparous, the latter being much more likely.

Habits. This is described as a ground shark, caught in E.^ropean waters most often
on hook and line. OflF the coast of Portugal and in the Gulf of Gascony it is most numerous
at depths of about 400 to 900 meters (about 220 to 500 fathoms). But there is also a long
list of recorded captures from the shallow waters of the English Channel and North Sea,
proving that its choice of depth is not narrow.

Its recorded diet includes smaller sharks {Squalus acanthias), other fish, and crabs.
Nothing is known of its breeding habits.

Range. Eastern Atlantic, from tropical West Africa to Ireland and the North Sea,
including the Mediterranean; apparently it is most numerous in the southern part of the
Bay of Biscay and oflF the coast of Portugal; it is known also from Morocco to Senegal.
It is accidental in the western Atlantic. It is also represented off South Africa, Argentina,
California,' in the Hawaiian, Japanese and Australo-New Zealand regions, and in Arabian

5. Cornish, Zoologist, (2) 10, i875:4SOi.

6. Burckhardt, Ann. Mag. nat. Hist, (7) 6, 1900: 568.

7. Hubbs and Clark (Calif. Fish Game, 31 [i], 1945: 64) report the recent capture of a six-foot specimen, off
Los Angeles County, California, indistinguishable from the eastern Atlantic brucus.

530 Memoir Sears Foundatioti for Marine Research

waters by a form (or forms) so close that It probably cannot be distinguished specifically
(see discussion, p. 527).

Occurrence in the Western Atlantic. The only reports of it in the western Atlantic
are of one that drifted ashore at Provincetown, Massachusetts, at the tip of Cape Cod, in
December, 1878,* and of a second, 2V2 meters long, taken near Buenos Aires more re-
cently."

Synonyms and References:

I. North Atlantic and Mediterranean:

Le boucle, Broussonet, Mem. Math. Phys. Acad. Sci. Paris, 1780: 672 (descr.).

Squalus brucus Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: II (descr., size, N. Atlant.).

Squale boucle, Lacepede, Hist. Nat. Poiss., 4° ed., j, 1798: 167, 283, pi. 3, fig. 2 (descr., ill.) ; in Sonnini,
Hist. Nat. Poiss., 4, 1802: 123 (from Lacepede, 1798).

Squalus spnosus Gmelin, in Linnaeus, Syst. Nat., / (3), 1788: 1500 (by ref. to Broussonet, 1 780) ; Walbaum,
P. Artedi Genera Pise. Emond. Ichthyol., 5, 1792: 519 (refs.) ; Bloch and Schneider, Syst. Ichthyol.,
1801: 136 (refs.); Bosc, Nouv. Diet. Hist. Nat., 21, 1803: 192 (diagn.) ; Latreille, Nouv. Diet. Hist.
Nat., 24, 1804: 72 (in table of contents); Risso, Ichthyol. Nice, 1810: 42 (descr., size, Medit.) ;
Strickland, Ann. Mag. nat. Hist., 4, 1840: 315 (Yorkshire, England); also, Calcutta J. nat. Hist.,
I, 1840: 285; Gatcombe, Zoologist, (3) i, 1877: 108 (size, stomach contents, off Plymouth, England).

Echinorhinus sfinosus Blainville, Bull. Soc. philom. Paris, 1816: 121 (name); in Vieillot, Faune Franc,
1825: 66, pi. 16, fig. I, 2'° (descr.) ; Bonaparte, Fauna Ital. Pesc, 5, 1835: plate not numbered (descr.,
iU., Medit.) ; Mem. Soc. neuchatel. Sci. nat., 2 (8), 1839: 9 (spelled Echinorrhinus in synopsis) ; YarreU,
Brit. Fish., SuppL, 2, 1839: 54 (descr., ill., teeth, denticles, Brit, record) ; Muller and Henle, Plagiost,
1841: 96 (descr.), pi. 60 (dermal denticles); Yarrell, Brit. Fish., 2, 1841: 532 (descr., Brit, records);
Hamilton, Brit. Fish., 2, 1842: 317, pi. 28 (descr., Gt. Brit.), also subsequent eds.; Bonaparte, Cat.
Pesc. Europ., 1846: 16 (in list); Machado, Feces Cadiz, 1847: 8 (off Cadiz); Cocks, Ann. Mag. nat.
Hist., (2) 5, 1850: 71 (south coast, England) ; Gray, List Fish. Brit. Mus., 7, 1 85 1 : 78 (refs., Medit.,
S. Afr.); Smith, Zoologist, 9, 1851: 3057 (Scotland) ; White, List. Spec. Brit. Mus., Fish., 8, 1 851: 132;
Costa, Fauna Napol. Pesci Cat. Sist. Condrott., in pt. 3, 1854-1857: 6 pp., pi. 16, pi. 17, fig. I, 2 (descr.,
best ill., near Naples) ; Dumeril, Hist. Nat. Poiss., r, 1865: 459, pi. 12, fig. 16-20 (denticles, descr.; see
Pacif. refs., p. 531) ; Bocage and Brito Capello, Poiss. Plagiost., 1866: 35 (Portugal) ; Gray, Ann. Mag. nat.
Hist., (4) I, 1868: 76 (Cornwall); Brito Capello, J. Sci. math. phys. nat. Lisboa, 2, 1870: 148 (Portu-
gal) ; Giinther, Cat. Fish. Brit. Mus., 8, 1 870: 428 (refs., descr., Cornwall, Medit., see also S. Afr. refs.,
p. 531) ; Canestrini, in Cornalia, et al.. Fauna d'ltal., 5, 1871— 1872: 42 (Medit.) ; Jackson and Clarke,
J. Anat. Lond., 10, 1876: 75 (brain and spinal nerves, Cornwall); Trois, Atti 1st. veneto, (5) 5,
1876-1877: 1 179 (Adriatic) ; Gervais and Boulart, Poiss., j, 1877: 214, pi. 83 (descr., ill.) ; Doderlein,
Prosp. Metod. Pesci Sicil., 1878: 30 (Sicily); Goode and Bean, Bull. Essex Inst. Salem, 11, 1879: 31
(Provincetown, Massachusetts, record); Stossich, Boll. Soc. adriat. Sci. nat., 5, 1 880: 70 (Adriatic);
Day, Fish. Gt. Brit., 2, 1880-1884: 323, pi. 162, fig. 2 (descr., ill., Brit, records); Doderlein, Man.
Ittiol. Medit., 2, 1881: 104 (records from Medit.);'' Moreau, Hist. Nat. Poiss. France, i, 1881: 365
(descr., ill., teeth, denticles, coasts of France) ; Perugia, Elenc. Pesc. Adriat., 1881 : 54 (Adriatic) ; More,
Zoologist, (3) 6, 1882: 434 (Ireland); Rochebrune, Act. Soc. linn. Bordeaux, (4) <5, 1882: 48; Faune
Senegambie, Poiss., J, 1883-1885:25 (Senegambia) ; Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883:
14 (old C. Cod record); Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 793 (distrib.) ; Carus, Prod.
Fauna Medit., 2, 1889-1893: 501 (Medit.); Goode and Bean, Smithson. Contr. Knowl., jo, 1895: 8,
pi. 3, fig. 9; also Mem. Harv. Mus. comp. Zool., 22, 1896 (descr., ill., old C. Cod record) ; Nobre, Ann.
Sci. nat. Porto, 2, 1 895: 225 (Portugal) ; Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896:

8. Goode and Bean, Bull. Essex Inst. Salem, /r, 1879: 31.

9. Berg, Com. Ictiol., Comm. Mus. nac. B. Aires, i (i), 1898 : 10. 10. We have not been able to see this plate.

II. See Doderlein for additional Mediterranean records in publications not accessible to us.

Fishes of the Western North Atlantic 531

219 (range) ; Bull. U.S. nat. Mus., 47 (l), 1896: 58 (refs., descr., old C. Cod record) ; Vieira, Ann. Sci.

nat. Porto, 4, i8g8: 138 (Portugal); Ridcwood, Anat. Anz., xs, 1899: 346 (skeleton, Bay of Biscay);

Popta, Ann. Sci. nat. (Zoo].) Paris, (8) 12, 1901: 205 (gill arches, France) ; dc Braganza, Result. Invest.

Sci. "Amelia," 2, 1904: 94, J06 (off Portugal) ; Ninni, Neptunia Venice, rp (20), 1904: 20 (Adriatic) ;

Borca, Arch. Zool. exp. gen., (4) 4, 1905: 203, 205 (France) ; Kendall, Occ. Pap. Boston Soc. nat. Hist.,

7 (8), 1908: 10 (old C. Cod record); Regan, Ann. Mag. nat. Hist., (8), 2, 1908: 42 (class., also

S. Afr. and Pacif. refs.); Giinther, Encyc. Brit., nth ed., 24, 191 1: 808 (general); Seabra, Bull.

Soc. portug. Sci. nat., 6, 191 1 : 200 (not seen); Roule, Bull. Inst, oceanogr. Monaco, 243, 1912: 18

(Gulf of Gascony) ; Le Danois, Ann. Inst, oceanogr. Monaco, 5 (5), 1913: 2i (Roscoff, France);

Metzelaar, Trop. Atlant. Visschen, 1919: 191 (old records); Jenkins, Fish. Brit. Isles, 1925: 327, pi.

125 (ill., Gt. Brit.); Leigh-Sharpe, J. Morph., 42, 1926: 313 (claspers) ; Ehrenbaum, in Grimpe and

Wagler, Tierwelt N.- u. Ostsee, Lief 7 (12^), 1927: 20 (general, North Sea) ; Rey, Fauna Iberica Feces,

/, 1928: 485 (descr., ill. of teeth, Spanish coasts); Gudger, Sci. Mon. N. Y., ^4, 1932: 417 (food);

Belloc, Rev. des Trav. Peches Marit., 7 (2), 1934: 149 (ill. after Bonaparte, Morocco to Senegal) ; Borri,

Atti Soc. tosc. Sci. nat., 44, 1934: lOO (Medit.) ; Nobre, Fauna Marinha Port. Vert., /, 1935: 403, pi.

64, fig. 201 (descr., ill., Portugal); Lubbert and Ehrenbaum, Handb. Seefisch. Nordeurop., 2, 1936:

290 (North Sea); Norman and Eraser, Giant Fishes, 1937: 53 (general).
Scymnus sfinosus Cuvier, Regne Anim., 2, 1817: 131, footnote; Cloquet, Diet. Sci. Nat., 25, 1822: 434

(general); Risso, Hist. Nat. Europe merid., 3, 1826: 136 (descr., Medit.); Bory de St. Vincent, Diet.

Class. Hist. Nat., 15, 1829: 598 (general); Cuvier, Regne Anim., 2, 1829: 368; Reguis, Ess. Hist. Nat.

Provence, i, Fasc. 1, 1877: 57 (Medit.).
Leiche boucle, Cloquet, Diet. Sci. Nat., Atlas Poiss., 1816-1830: pi. 28, fig. 2.
Squalus {Scymnus) sfinosus Vo'xff., in Cuvier, Tierreich, 2, 1832: 513 (descr.).
Goniodus sfinosus hg^%%\z.,'L.,Yo\%i.'Fo%s., 3, 1835-1838: 94, pi. E, fig. 13 (teeth) ; Owen, Odontogr., 1840-

1845: 31, pi. 4, fig. 4 (teeth). 1=
Spinous Shark, Cornish, Zoologist, i, 1866: 102 (Cornwall); Couch, Fish. Brit. Isles, i, 1867: 54, pi. 12

(ill., Brit, records); Cornish, Zoologist, (2) 5, 1870: 2347 (Cornwall); Zoologist, (2) 10, 1875: 4501

(luminescence, Cornwall) ; Zoologist, (3) (5, 1882: 22 (Cornwall).
Echinorhinus (no specific name) Hasse, Naturl. Syst. Elasm. besond. Theil, 1882: 73, pi. 9 (vertebrae,

dermal denticles); Burckhardt, Ann. Mag. nat. Hist., (7) d, 1900: 568 (luminescent organs lacking).
Echinorhinus brucus Garman, Mem. Harv. Mus. comp. Zool., jd, 1913: 243 (descr.); Roule, Result. Camp.

sci. Monaco, c,2 , 1 91 9: 123 (off Brittany) ; Bigelow and Welsh, Bull. U.S. Bur. Fish., 40 (l), 1 92 5: 55

(ill., old C. Cod record); Jordan, Manual Vert. Anim. NE. U.S., 1929: 14 (general, C. Cod);

Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 26 (old C. Cod record); Jordan, Evermann and

Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 22 (ref. to old C. Cod record); de Buen, Faune Ich-

thyol.. Cons, explor. Mer., 1931 : plate not numbered; Bigelow and Schroeder, Canad. Atlant. Fauna, biol.

Bd. Canad., 12', 1934: 20 (descr., ill., old C. Cod record) ; Fowler, Bull. Amer. Mus. nat. Hist., jo (i),

1936: 88 (descr., ill., W. Afr.).

2. South Africa and South Atlantic:

Echinorhinus obesus Smith, A., 111. Zool. S. Afr., 1849: pi. i (descr., ill., S. Afr.).

Echinorhinus sfinosus Giinther, Cat. Fish. Brit. Mus., 8, 1 870: 428 (Cape of Good Hope); Berg, Com.
Ictiol. Comm. Mus. nac. B. Aires, i (i), 1898: 10 (near B. Aires) ; Lahille, Physis B. Aires, 5, 1921 : 63 ;
Enum. Feces Cartilag. Argent., 1921: 17 (Argentina); Marelli, Elenc. Sist. Fauna B. Aires (1922-
1923), 1924: 546 (Argentina) ; Barnard, Ann. S. Afr. Mus., 21 (l), 1925: 46; also, 21 (2), 1927: pi. 2,
fig. 6 (S. Afr.) ; Pozzi and Bordale, An. Soc. cient. argent., 120, 1935: 152 (listed, Argentina).

3. Pacific:

Echinorhinus sfinosus Dumeril, Hist. Nat. Poiss., j, 1865: 459 (refs. for Aust., Tasmania); Parker, Trans.
Proc. N. Z. Inst., 16, 1884: 28 (N. Zealand) ; McCoy, Prod. Zool. Victoria, Decade 75, 1887: pi. 144

12. Spelled Gonjaiaj.

532 Memoir Sears Foundation for Marine Research

(dcscr., meas., good ill., Aust.) ; Lucas, Proc. roy. Soc. Vict., N. S. 2, 1890: 44 (Aust.) ; Jordan, Tanaka

and Snyder, J. Coll. Sci. Tokyo, jj, 1913: 23 (Japan).
Echinorhinus brucus Waite, Rec. Canterbury [N. Z.] Mus., 2 (l), 1913: 17 (size, weight, N. Zealand);

Phillipps, N. Z. J. Sci. Tech., 10, 1928: 221 (N. Zealand); Fowler, Proc. Pan-Pacif. sci. Congr., (4)

5, 1930: 497; Bull. U.S. nat. Mus., 100 (r^), 1941: 277 (Pacif. refs.) ; Hubbs and Clark, Calif. Fish

Game, 57 (2), 1945: 64, fig. 16, 17 (descr., photos, California).
Echmorhinus cookei Pietschmann, Anz. .Akad. Wiss. Wien, 27, 1928: 297 (descr., Hawaiian Is.) ; Bull. Bishop

Mus., 7j, 1930: 3 (descr., Hawaiian Is.).
Echmorhinus (Rubusqualus) mccoyi Whitley, Aust. Zool., 6, 1 931: 311 (descr., Victoria); Mem. Qd. Mns.,

10 (4), 1934: 200; Fish. Aust., i, 1940: I 5 I (descr., ill., Aust.).

Suborder PRISTIOPHOROIDEA

Characters. No anal fin; 2 dorsal fins without spines; either 5 or 6 gill openings, all
of them anterior to origin of pectoral; snout greatly elongate, blade- or beak-like; each
edge of snout and of head, anterior to mouth, armed with a row of sharp transverse, tooth-
like structures and with a long fleshy barbel; oral teeth small, numerous, with i cusp, simi-
lar in front and sides of mouth, with several rows functional; trunk subcylindrlcal, except
that head and snout are flattened dorso-ventrally ; anterior margins of pectorals not ex-
panded forward past ist gill opening; nostrils entirely separate from mouth; eye without
nictitating fold or membrane; spiracles present; inner margins of pelvics entirely separate
posterior to cloaca ; vertebral column completely segmented, with well developed centra,
the notochord segmentally constricted correspondingly; skull with a separate antorbital
bar of spongy cartilage extending rearward from nasal capsule, past orbit, as far as corner
of mouth ;^ upper jaw (palatoquadrate cartilage) attached to hyomandibular, and also
firmly articulated by a short, narrow transverse process to lower side of cranium in post-
orbital region and by a ligament to the antorbital bar;" rostral cartilages united as a single
elongate, blade-like bar, occupying the entire breadth of the snout to its tip; propterygial
cartilage of pectoral bears i radial element only; pelvic transverse; heart valves in 3 rows.
Development ovoviviparous.

Remarks. The sharks of this group have usually been placed among the Squaloidea,
with which they agree in lacking an anal fin. But we believe they should rank as a distinct
suborder (see p. 77), for they differ not only as regards their beak-like snout with its lateral
teeth, in which they are unique among modern sharks, but in the presence of a separate
antorbital bar, from which the upper jaw is suspended by a broad ligament in addition to
the articulation to the cranium.

The saw-like beak makes them resemble superficially the sawfishes (Pristidae)
among the Batoidei. But they are true sharks because of their free upper eyelids, their
pectorals not united with the sides of the head, and their lateral, not ventral, gill openings.

Range. Indo-Pacific; South Africa; Tasmania; Australia; Philippines; Korea;
Japan. Fossil pristiophorids are known from the Cretaceous, Miocene, and Tertiary.

Families. One family only, Pristiophoridae, with characters of the suborder.

I. This bar, like the rostrum proper, is armed with lateral teeth. 2. Description based on original dissection.

Fishes of the Western North Atlantic 533

Key to Genera
la. Five gill openings. Pm//opAor«i- Miiller and Henle, 1837.

South Africa, Australia, Tasmania, Philippines,
Japan, Korea.
lb. Six gill openings. Pliotrema Reg^n, i()o6.

South Africa.
Species. Pristiophorus includes three known species, Pliotrema only one.'

Suborder SQUATINOIDEA

Characters. No anal fin; 2 dorsal fins v/ithout spines; only 5 gill openings, all ante-
rior to origin of pectorals; snout not beak-like, without lateral teeth or cirri; teeth in front
of mouth essentially similar to those toward its corners; general form skate-like rather
than shark-like, with the trunk very much flattened dorso-ventrally and expanded later-
ally anterior to cloaca, but tapering thence rearward; eyes dorsal; anterior margins of
pectorals extending forward past gill openings* and partly concealing them; pelvics also
very broad, wing-like, their inner margins entirely separate posterior to cloaca; nostrils
separate from mouth ; spiracles present, dorsal ; eyes without nictitating fold or membrane;
vertebral column completely segmented throughout its length, its axial canal much con-
tracted in the region of the centra, which are well differentiated, with the notochord greatly
constricted segmentally in its passage through them; vertebral centra with 2 or more series
of concentric calcified lamellae; neural spines attached to dorsals;' rostral cartilage single,
very short; skull without antorbital processes or separate antorbital bars; upper jaw (pala-
toquadrate cartilage) articulated to hyomandibular bar, with a long transverse process that
is attached to the ethmoid region of the cranium by a ligament;" propterygial cartilage of
pectoral elongate, directed anteriorly, corresponding to forward expansion of the fin, but
bearing much fewer radials than the metapterygium;' pelvis curved rearward, corre-
sponding to the expanded pelvic fins; heart valves in 6 to 7 rows (an exceptionally large
number for sharks, but characteristic of skates and rays) .' Development ovoviviparous.

Remarks. Although very skate-like in appearance, in number of heart valves, and in
some skeletal .characters, the squatinoids are usually classed with the sharks because they
have free eyelids, pectorals with anterior margins not attached to the sides of the head,
and gill openings that are not confined to the lower surface but extend up onto the sides
of the neck as well.

Their method of swimming, also shark-like, is by a sculling motion of the tail, little

3. For descriptions of the species of the two genera, with lists of references, see Fowler (Bull. U.S. nat. Mus., loo

['ill '94' ■ i8o, 283).
4.. But not attached to sides of head. 5. Mivart, Trans, zool. Soc. Lend., to, 1879: pi. 77, fig. 5-

6. For account and excellent illustration of the skull, see Gegenbaur (Unters Vergl. Anat. Wirbelt., 5, 1872: 190,
pi. II, fig. 2, pi. 12, fig. 4).

7. Gegenbaur, Unters. Vergl. Anat. Wirbelt., 3, 1865: pi. 9, fig. 10.

8. For account of the vertebrae and dermal denticles, see Hasse (Naturl. Syst. Elasm. besond. Theil, 1882: 126,
pi. 17, fig. i-io) } for heart valves, see Marples (Trans, roy. Soc. Edinb., 5* [3], 1936: 817).

534 Memoir Sears Foundation for Marine Research

use being made of the wing-like pectorals, which, on the contrary, are the effective swim
ming organs of skates and rays.*
Only one family is known.

Family SQ UA TINIDAE

Characters. Snout very broad and short; eyes dorsal, without nictitating membranes:
spiracles large; nostrils terminal, entirely separate from mouth, their anterior margin;
with barbels which are variously lobed; gill openings lateral, but extending onto lowei
surface; mouth protrusible at corners with well developed labial cartilages, nearly ter-
minal but separated from front of snout by a deep transverse furrow which is edged by £
thin, variously-lobed fold of skin; lower jaw with deep labial furrows near corners; teetl
numerous, similar in the 2 jaws, with single thorn-like cusp on broad base, 3 or 4 series
functional; caudal axis not raised at all above main axis of trunk; lower lobe of cauda.
longer than upper. Development ovoviviparous ; embryo with very large yolk sac."

Genera. Only one genus is known.

Genus Squatina Risso, 1 8 10

Squatina Risso, Ichthyol. Nice, 1810: 45; type species, Squatina vulgaris Risso, equals Squalus squatim
Linnaeus, 1758.^'

Generic synonyms:

Rhino SchaefiFer, Epist. Stud. Ichthyol., 1760: 20; Klein, Neuer Schauplatz, 2, 1776: 587; Walbaum, P
Artedi Gen. Pise. Emend. Ichthyol., 1792: 580, not available;'^ not Rhina Bloch and Schneider, Syst. Ich-
thyol., 1 801: 352; Cuvier, Regne Anim., 2, 1 81 7: 133; Muller and Henle, Plagiost., 1 841: lio; a ra)
which was subsequently named Rhamfhobatis by Gill, Ann. N. Y. Lye, 7, 1 862: 408.

Generic Characters. Head broadly rounded, with wing-like lateral expansions;
spiracles behind eyes and at the same level as the latter; lips with well developed support-
ing cartilages, widely protrusible at corners of mouth but not centrally; anterior margins
of pectorals expanded anteriorly as narrow shoulder-like extensions, lying below the
lateral expansions of the sides of the head, partly concealed by the latter, and more or less
overlapping the pel vies rearward; pel vies originating anterior to posterior corners of pec-
torals; dorsals much smaller than pectorals or pelvics; caudal triangular, its posterior con-

9. For a recent discussion of the affinities of the suborder, based chiefly on the skull, see Holmgren (Acta Zool., 22.
1941 : 79). They are classed (as Angeliformes) among the skates and rays by Le Danois (Rev. des Trav. Peches
Marit., t^, 1945: 67) because of the nature of their vertebral calcifications.

10. Garman, Mem. Harv. Mus. comp. Zool., 56, 1913 : pi. 61, fig. 9— 1 1.

11. The name Squatina was first proposed by Valmont (Diet. Rais. Univ. Hist. Nat., /, 1768: 117). But by ruling
of the International Commission on Zoological Nomenclature (Smithson. Misc. Coll., 75 [3], 1925: 27, Opinion
89) his names are not taken into consideration. It was next mentioned by A. M. C. Dumeril (Zool. Anal., 1806:
102), but without inclusion of any particular species, so that, as a generic name, it must date from Risso, 1810.

I 2. Schaeffer did not include any species in his Rhina; and the generic names proposed by Klein and republished by
Walbaum are not to be taken into account, according to Opinions 21 and 89 of the International Commission
on Zoological Nomenclature (Opinions Rendered, Smithson. Publ., No. 1938, 1910: 51; Smithson. Misc.
Coll., 7 [3], 1925: 27).

Fishes of the Wester ti North Atlantic

535

tour truncate or concave, its lower anterior margin longer than the upper anterior margin
(a relationship the reverse of that which obtains in all other sharlcs) ; caudal peduncle with
a faintly defined longitudinal ridge along each side; claspers of males extend only a little
beyond pelvics, even in adults, and are attached to inner margins of fins nearly to tips of
latter;" dermal denticles on dorsal side conical, on broad bases, those on ventral side flat,
with scale-like blades. Characters otherwise those of family and suborder.

Range. Continental waters on both sides of the Atlantic, including the Mediterra-
nean, north to the Shetlands and Cape Cod and south to northern Argentina; South Africa
(Natal); west coast of America from Chile to southern Alaska; Japan, Korea, Australia
and Tasmania.

Fossjl. From Upper Jurassic to Pliocene, Europe; Upper Cretaceous, western Asia;
Upper Cretaceous to Miocene, North America; Eocene, Africa.

Figure 103. Margins of right-hand nostrils of different species of Squatina: A, australis from Australia
(Harv. Mus. Comp. Zool., No. 659), about 3 x. B, Left-hand nostril of same, to show asymmetry. C, squatina
from the Irish Sea (Harv. Mus. Comp. Zool., No. 846), about natural size. D, argentina from Uruguay
(U. S. Nat. Mus., No. 87684), about 1.5 x. E, japonica from Japan (Harv. Mus. Comp. Zool., No. 1 112),
about 2 X. F, californica from California (Harv. Mus. Comp. Zool., No. 952), about 2.5 x. G, dumeril
from off New York (U. S. Nat. Mus., No. 11 8461), about 1.5 x. H, armata from Mejillones Island, Peru
(Harv. Mus. Comp. Zool., No. 531), about 1.5 x.

13. For account, see Leigh-Sharpe (J. Morph., 55, 1921 : 373).

536 Memoir Sears Foundation for Marine Research

Species. Recent estimates of the number of species actually represented by the named
representatives of this curious genus range from only one" to eight or nine," a list to
which two more have subsequently been added." Our own examination of specimens from
widely separated seas" has convinced us that the latter opinion is more nearly correct., i.e.,
that most of the supposed species are separable from one another by characters that seem
precise enough to be accepted as specific, although they all resemble one another so closely
in general apearance that identification requires close inspection. The conformation of the
nasal margins and barbels (Fig. 103) proves a reliable diagnostic character, but the degree
to which the mid-dorsal line of denticles is enlarged is far less so, since this may vary widely
with age.

Key to Species

I a. Inner nasal barbel strongly ramose at tip 5 margin of nostril between barbels deeply
fringed (Fig. 103 A, B).

2a. Outer nasal lobe strongly fringed. oculata Bonaparte, 1 840.

Eastern Atlantic, Mediterra-

nean.

2b. Outer nasal lobe smooth or only weakly fringed (Fig. 103 A, B).

3a. Outer corner of pectoral little if any more obtuse than a right angle, its inner
corner subangular j upper surface with small white and gray spots only.

australis^^ Regan, 1906.

Southern Australia, Tasmania.

3b. Outer corner of pectoral much more obtuse than a right angle, its inner corner

broadly rounded} upper surface with conspicuous brown ocelli as well as

small spots. tergocellata^'' McCuUoch, 19 14.

Western and southern Australia.
lb. Inner nasal barbel simple, or at least not strongly ramose j margin of nostril between
barbels smooth, or at most feebly fringed (Fig. 103 C-H).
4a. Fold along front of head with 2 lobes opposite and in front of corner of mouth.

nebulosa Regan, 1906.

Japan.^°
4b. Fold in front of head with i lobe only, or none.

5a. Fold along front of head expanded as a noticeable triangular lobe outside
corner of mouth (Fig. 103 C). squatina Linnaeus, 1758.

Eastern North Atlantic; Medi-
terranean.

14. Ribeiro, Fauna brasil. Peixes, 2 (i) Fasc. i, 1923: 27.

15. Regan, Ann. Natal Mus., r, 1908 : 2+8; Garman, Mem. Harv. Mus. comp. Zool., 36, 1913 : 251.

16. Tergocellata from Australia (McCulloch, Biol. Res. "Endeavor," 2, 1914: 84) and argentina from Argentina
(Marini, Pliysis B. Aires, ,0, 1930: 0.

17. From off the east coast of United States; Mediterranean; coast of northwestern Europe; Uruguay; S. Peruj
California; Japan; Australia.

18. We question whether tergocellata is actually distinct from australis.

19. This form is known only from the original description (without illustration) of the type specimen; it may prove
identical with jafonica.

Fishes of the Western North Atlantic 537

fb. Fold along front of head either nearly straight or at most obtusely rounded
opposite corner of mouth (Fig. 103 D-H).
6a. Inner nasal barbel broadly spatulate (Fig. 103 D-F).

7a. Distance from eye to spiracle nearly or quite twice as long as hori-
zontal diameter of eye. argentina^" Marini, 1930, p. 544.
7b. Distance from eye to spiracle little if any longer than horizontal
diameter of eye.

8a. Distal margin of pectoral nearly straight, its inner corner
broadly rounded, its inner margin strongly convex.

jafonica Bleeker, 1857.

Japan, Korea.

8b. Distal margin of pectoral weakly concave, its inner corner sub-
angular, its inner margin only slightly convex.

calif ornka Ayres, 1859.

Mexico to southern Alaska.

6b. Inner nasal barbel narrow, tapering (Fig. 103 G, H).

9a. Distal margin of pectoral marked off from inner margin by a defi-
nite, subangular corner. dumeril Lesueur, 1818, p. 538.
9b. Distal margin of pectoral not marked off from inner margin by a
subangular corner.

lOa. Distance from anterior corner of pectoral to rear end of its
base equals % or more of its length ; posterior margin of lower
lobe of caudal, as well as upper, vertically truncate.

arwtf/fl Philippi, 1887.
Chile, Peru."

lOb. Distance from anterior corner of pectoral to rear end of its
base equals only about % of its length j posterior margin of
lower lobe of caudal oblique. africana Regan, 1908.

Natal, South Africa.-*

20. A specimen from Uruguay which we have examined in the U.S. National Museum is clearly referable to this
species, and probably this also applies to the form reported from Rio de Janeiro by Ribeiro (Arch. Mus. nac. Rio
de J., 14, 1907: pi. 10; Fauna brasil. Peixes, 2 [i] Fasc. i, 1923: pi. 9) as Squatina squatina. Although the
inner corners of its pectorals are shown as angular or subangular (rounded in argentine) , the conformation of the
margin of the nostrils agrees with that of argentina, in addition to the fact that the outer corners of the pectorals
are much more obtuse than a right angle, and that the eyes and spiracles are far apart.

21. The original account of this species is not sufficiently detailed to locate it positively in the key. The characters
given above are from a specimen from Mejillones Island, Peru (Harv. Mus. Comp. Zool., No. 531), presumably
the type of p/iilifpi Garman, 1913, which we believe to be a synonym of armata.

22. Records under this name from Lagos and Goree in tropical West Africa (Gilchrist and Thompson, Ann. Durban
Mus., /, 1 91 6: 284; Metzelaar, Trop. Atlant. Visschen, 1919: 191), and from Cape Blanco, Morocco (Cha-
baoaud and Monod, Bull. £tud. Hist. Sci. Afr. Occid. Franc, 1927) do not include evidence sufficient for
specific identification.

538 Memoir Sears Foundation for Marine Research

Squatina dtimeril (Lesueur), 1 8 1 8

Angel Shark

Figures 103 G, 104, 105

Study Material. Female, 382 mm. long, from the continental slope off New York,
Lat. 39° 42' N., Long. 71° 17' W., taken Sept. 19, 1887 (U.S.Nat. Mus., No. 118461);
male, 1,080 mm. (42 ^^ inches) long, from lower Chesapeake Bay at Lynnhaven Roads,
Virginia; mounted specimen, 43 inches, from Martha's Vineyard, Massachusetts (New
Eng. Mus. Nat. Hist.).

Distinctive Characters. Its skate-like appearance separates the Angel Shark at a
glance from any other shark except for some other members of its own genus. For specific
characters within the genus, see the preceding Key (p. 536).

Description. Proportional dimensions in per cent of total length. Female, 382 mm.,
from Lat. 39° 42' N., Long. 71° 17' W. (U.S.Nat. Mus., No. 118461).

Extreme breadth at outer extremity of pectorals: 60.8.

Trunk at origin of pectoral: height 9.0.

Snout length in front of: outer nostrils O; mouth O.

Eye: horizontal diameter 2.0; distance between eyes 8.4.

Spiracles: distance between lO.O; from spiracle opening to eye 2.2.

Mouth: breadth 13.6; height 2.1.

Nostrils: distance between inner ends 5.8.

Gill opening lengths : ist 7.0; 2nd 6.3; 3rd 5.8; 4th S-S) 5th 5.0.

First dorsal fin: vertical height ^.S; length of base 3.3.

Second dorsal fin: vertical height 5.O; length of base 3.1.

Caudal fin: upper margin 13.6; lower margin 16.7.

Pectoral fin: oxxttr mzvgm i%.0; inner margin 15.7; distal margin 14.9.

Distance from snout to: ist dorsal 65.7; 2nd dorsal 76.5; upper caudal 86.4;

pectoral 17.5; pelvics 37.2.

Interspace between: ist and 2nd dorsals 7.5; 2nd dorsal and caudal 7.3.

Distance from origin to origin of: pectoral and pelvics 22.2; pelvics and caudal

45.6.

General form of trunk as described for family and genus. Caudal peduncle ex-
panded laterally as a low ridge on either side posterior to 2nd dorsal. Dermal denticles
on dorsal surface loosely spaced but generally distributed, conical, the spinous portion
weakly recurved, usually with 4 or more longitudinal ridges; bases broad, their outlines
more or less radiate; larger denticles in clusters of 5 or 6 beside inner anterior and inner pos-
terior edges of eyes, with a row extending from near inner side of nostril toward eye; young
specimens also with a single row of denticles, 3 to 4 times as large as the others, along mid-
line of back from opposite anterior ends of bases of pectorals to origin of ist dorsal, but in
large specimens these are little, if any, larger than the denticles that flank them; adult

Fishes of the Western North Atlantic

539

Figure 104. Squatina dumeril, female, 382 mm. long, from off New York (U. S. Nat. Mus., No. 1 18461).
A Dorsal view. B Anterior view. C Dermal denticles, about 7 x.

Figure 105. j4 Ventral view of Squatina dumeril pictured in Fig. 104. B Side view of posterior part of
trunk with caudal fin. C Upper and lower teeth from center of mouth. D Eighth and ninth lower teeth,
about 3.6 X. E Fifth upper tooth, about 7.2 x. F Fifth lower tooth, about 7.2 x.

540 Memoir Sears Foundation for Marine Research

males with larger denticles along anterior margins of pectorals and near their outer cor-
ners;" denticles on lower surface with flat ovoid blades on very short pedicels, close-spaced
on outer parts of paired fins and on lower edge of tail sector of trunk, but abdomen and
inner parts of paired fins naked except for patches here and there.

Head broadly rounded anteriorly, but its postero-lateral margin nearly straight, its
greatest breadth about 3 times as great as distance between spiracles or V3 as great as dis-
tance from snout to rear end of base of 2nd dorsal. Horizontal diameter of eye about
equal to maximum diameter of spiracle, the distance from eye to spiracle about as long as
horizontal diameter of eye. Distance between spiracles longer than between eyes by a
distance about equal to vertical diameter of eye. Distance from eye to nostril a little longer
than from eye to spiracle. Nostril terminal, its inner anterior margin with 2 barbels, the
outer barbel triangular with broad base and slender tip, the inner barbel narrow, widening
slightly outward, then tapering to pointed tip and rising from the base of the outer without
an intervening lobe. Both barbels smooth-edged or nearly so, the outer anterior margin
of nostril expanded as a smooth-edged, subtriangular lobe; the posterior margin of nostril
smooth."* Fold at front of head only slightly expanded in obtusely rounded contours out-
side corners of mouth. Mouth terminal, its breadth equal to about % of length of head.
Lower labial furrow extending about % of the distance toward center of mouth, but no
upper furrow.

Teeth '^^° in specimen counted, alike in the 2 jaws, with erect, conical cusp on broadly
expanded base, the outermost teeth slightly the smallest; 3 rows functional in each jaw;
a broad gap at symphysis in each jaw.

Dorsals similar in form, brush-shaped, with broadly rounded apex. First dorsal about
V5 as long at base as head, its vertical height a little greater than length of base, its origin
posterior to tips of pelvics by a distance about V2 as great as distance between eyes. Inter-
space between ist and 2nd dorsals about as long as between eyes. Second dorsal a little
smaller than ist. Interspace between 2nd dorsal and caudal about as long as between ist
and 2nd dorsals. Caudal between % and Yj of total length, its upper margin only about
% as long as its lower margin, both its corners rounded but the lower more broadly so, the
posterior margin moderately concave with a shallow obscure notch opposite termination of
caudal axis. Pelvics about 73 as long as exlreme length of pectorals, the anterior margins
weakly convex and distal margins nearly straight, their outer corners broadly rounded,
their posterior corners tapering with acute tip (about 40°). Transition from distal margin
to inner margin of pectoral marked by a definite subangular corner, its narrowly rounded
outer corner approximately a right angle, its outer margin nearly straight, distal margin
weakly concave, inner posterior margin moderately rounded and definitely notched at
axil; distance from anterior corner of pectoral to rear end of its base about 73 as great as
length along outer anterior margin, as is also the distance from outer corner to rear end of

13. Clearly shown in a photograph of a 42j^-inch male from Chesapeake Bay (Hildebrand and Schroeder, Bull.
U.S. Bur. Fish., ^5 [i], 1928: fig. 30) and mentioned by Dumcril (Hist. Nat. Poiss., /, 1S65: 467, footnote).
24. More or less fringed in calif ornica.

Fishes of the Western North Atlantic 541

base} its anterior corner posterior to a transverse line at rear edges of eyes by a distance
about V2 as great as distance between eyesj posterior parts of pectorals overlap anterior
parts of pelvics by a distance a little less than distance between nostrils.

Color. Fresh specimens are described as bluish-gray or ashy-gray above, tinted with
red on head and margins of finsj white below, with a reddish spot on the throat, a second
on the abdomen and a third extending from behind the cloaca to the caudal j the pelvics are
bordered below with irregular reddish bands. After preservation in alcohol the upper sur-
face of specimen described is chocolate brown with pale mottlings, the upper surface of
anterior corner of pectorals brownish white, the lower surface of the trunk grayish white
anterior to the vent, but pale reddish brown on the tail sector, and the paired fins broadly
edged with pale reddish brown.

Size. The fact that the claspers were well developed in a male of 42 V2 inches suggests
that maturity is reached at a length of 3 to 3 V2 feet ; the maximum length so far definitely
reported is four to five feet. One of four feet weighed about 60 pounds. It is not known
whether the American 5. dumeril ever grows as large as its European relative, S. squatina,
which often reaches a length of six feet (maximum reported, eight feet) with a weight of
1 60 to 1 70 pounds.

Developmental Stages. Embryos of S. dumeril have not been seen, but conditions in
allied species make it likely that the yolk sac is very large, and that gravid females may
contain as many as 13 to 25 embryos.

Remarks. This species has frequently been considered identical with 5. squatina of
the eastern Atlantic. Comparison of American with European specimens has shown, how-
ever, that they are easily distinguishable by the following characters: the shapes of the
head folds (cf. Fig. 103 G with 103 C), the conformation of the nasal flaps and barbels,
the innermargin of the pectoral notched at the axil in dumeril but not in squatina, the hori-
zontal diameter of eye almost as great as the maximum diameter of the spiracle in dumeril
but definitely smaller than the spiracle in squatina, the maximum breadth of the head only
about % as great as the length of the pectoral in dumeril but nearly or quite as great as the
length of the pectoral in squatina, and (in small specimens) the mid-dorsal denticles larger
in dumeril.

Habits. Most of the specimens so far reported have been taken in depths of only a
few feet. However, one of the specimens listed above (p. 538) was taken in September
over the continental edge 80 miles from the coast, where the depth was 705 fathoms,"'
while several others were trawled by the United States Fisheries Steamer "Albatross"
about 75 miles off Long Island, New York, in 109 fathoms in February 1 920, showing that
it may stray far offshore and that it may also occur at considerable depths irrespective of sea-
son. Fragments of fish and of bivalve moUusks were found in the stomach of one taken in
North Carolina, where it is described as "often troublesome, getting snarled in the nets or
eating other fish caught therein j it also bites the fishermen if they are not wary."'^* No

15. "Albatross" Dredging Station 2749. 25a. Smith, Bull. N. Carolina geol. econ. Surv., », 1907: 38.

542 Memoir Sears Foundation for Marine Research

other firsthand observations appear to ha\'e been made on the habits of the North American
species. Analogy, however, with its close relative 5. sqitatina of the eastern Atlantic sug-
gests that it lives on or close to bottom, often burying itself partially in the sand or mud, as
do rays and flatfishes (pleuronectids), and that it feeds on a variety of fish, perhaps chiefly
on flounders and skates, as well as on crustaceans and gastropod mollusks.'" Probably the
young are born in summer, when the adults are close inshore.

Relation to Man. Squattna is not plentiful enough in American waters to be of any
commercial importance. In Europe a certain number are marketed for fried-fish shops.
Formerly its skin was in some demand for polishing wood and ivory, and in earlier days its
dried flesh was "prescribed as a sovereign remedy for the itch.""'

Range. East coast of the United States from southern New England to North
Carolina and southern Florida, north coast of the Gulf of Mexico, and reported recently
from Jamaica."^

This is a summer visitor to the mid-Atlantic coast of the United States. In most years
it appears at Cape Lookout, North Carolina, in late March or April to remain until about
the first of May. To the northward it has been reported between May and October from
Chesapeake and Delaware Bays, and from various localities along the coast to the vicinity
of New York, as well as in the bays along the southern shore of Long Island. Most of the
records for it have been based on odd specimens only. But it has been reported as some-
times common on the outer coasts of Virginia and Maryland, as well as Delaware, suggest-
ing that this section may be a center of abundance for it. Three specimens have been re-
ported from Rhode Island, two from the vicinity of Woods Hole,"^ this last being the
most easterly and northerly record for it. Positive knowledge of it southward from North
Carolina is limited to reports that it is occasionally taken in summer among the Florida
Keys,^° and that it is caught occasionally in shrimp trawls off the mouth of the Mississippi."
There is one record for Corpus Christi, Texas, and a 30-inch specimen has been taken in
the harbor of Port Royal, Jamaica.^' But it has not been reported at all anywhere between
Jamaica and Rio de Janeiro, nor has any other member of its genus for that matter, sug-
gesting that its normal range does not extend to the equatorial belt. Although described as
quite abundant for a short period in spring off the North Carolina coast,^^ present indica-
tions are that it is far less plentiful than is its European relative S. squattna, as many as 26
of which have been reported as lying on the beach on the west coast of England at one
time." We have seen only one fresh-caught specimen.'"

26. We wonder whether an old and oft-quoted account of one seen to come to the surface and to seize a living cor-
morant may not actually have referred to an ang'ler {Lof/iiiis) which commonly captures sea fowl in this way.

27. Norman and Fraser, Giant Fishes, 1937: 55. 28. Personal cuiiimunication from Luis Howell-Rivero.

29. Both of them from Menemsha Bight, Martha's Vineyard Island, the one in 1S73, 'he other in September 1921.

30. Fowler (Proc. Acad. nat. Sci. Philad., 5S, 1906: 80). It has also been conjectured that at least one of the speci-
mens on which the species was founded was from Florida. But the wording of Lesueur's original account (P'oc.
Acad. nat. Sci. Philad., /, 1818: 226) suggests, rather, that both of his specimens were studied by him in a fresh
condition, i.e., that they were collected not far from Philadelphia.

31. Personal communication from Stewart Springer. 32. Personal communication from Luis Howell-Rivero.
33. Coles, Proc. bid. Soc. Wash., 2S, 1915: 92. 34. Day, Fish. Gt. Brit., 2, 1S80— 1884: 32s.

35. In Chesapeake Bay.

Fishes of the Western North Atlantic 543

The wintering ground of the sparse population of both young and adults that visit
the east coast of the United States in summer is not known. It seems more likely that they
move out into deeper water than that they migrate southward along the shore, for Angel
Sharks are scarce around Florida, while one specimen was trawled about 75 miles off Long
Island, New York, in February 1920, and another about 25 miles off Bodie Island, North
Carolina, in about 40 fathoms in February 1931.'°

Synonyms and References:

Sqtmtina duineril Lesueur, J. Acad. nat. Sci. Philad., /, 1818: 225, pi. lO (descr., no loc.) ; Bory de St.
Vincent, Diet. Class. Hist. Nat., 75, 1829: 598; DeKay, Zool. N. Y., 4, 1842: 363, pi. 62, fig. 203
(descr., ill., after Lesueur, 1818, N. York); Rep. St. Cab. nat. Hist. N. Y. (1855), S, 1858: 64 (listed
for N. York); Fowler, Copeia, 30, 1916: 36 (mid-Atlantic states); Proc. biol. Soc. Wash., a, 1920:
145 (N. Jersey) ; Proc. Acad. nat. Sci. Philad., 7^, 1922: 3, 6 (N. Jersey) ; Hildebrand and Schroeder,
Bull. U.S. Bur. Fish., ^5, 1928: 54 (descr., photo, Chesapeake Bay) ; Jordan, Manual Vert. Anim. NE.
U.S., 1929: 15 (general); Fowler, Proc. Acad. nat. Sci. Philad., 80, 1929: 608 (N. Jersey); Truitt,
Bean and Fowler, Bull. Md. Conserv. Dep., 5, 1929: 28 (Maryland) ; Jordan, Evermann and Clark, Rep.
U.S. Comm. Fish. (1928), 2, 1930: 22 (Newport, Rhode Island); Fowler, Monogr. Acad. nat. Sci.
Philad., 7, 1945: 97 (one 42 inches, Beaufort, N. Carolina).

Sjuatina dumerilii Wils, Observ. Squatina laevi, Inaug. Dissert. Lugduni-Batavorum, 1 844: 5 (America).

Squatina dumerili Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, I 846: 256 (in synopsis) ; Griffith, Proc. Acad.
nat. Sci. Philad., 5, 1847: 246 (Delaware Bay); Leidy, Proc. Acad. nat. Sci. Philad., 3, 1847: 247 (no
loc); Baird, Rep. U.S. Comm. Fish. (1871-72), 1873: 827 (Woods Hole, Massachusetts); Gill, Rep.
U.S. Comm. Fish. (1871-72), 1873: 813 (listed, C. Cod to Florida); Smith, Copeia, 106, 1922: 33
(Martha's Vineyard).

Squatina angelus (in part) Valenciennes, Hist. Nat. Canaries, 2, 1837-44: 102 (N. York to Brazil); Jordan
and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 35 (Atlant. coast, N. Amer.); Gaskill, Forest and Stream,
36, 1891 : 68 (C. May, N. Jersey, not seen).

Rhina dumerili Gill, Proc. Acad. nat. Sci. Philad., Addend., 1 861 : 61 (name) ; Dumeril, Hist. Nat. Poiss., /,
1865: 467 (descr., spec, from N. York).'"

Rhina dumeril Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 252 (descr., Newport, Rhode Island);
Radcliffe, Bull. U.S. Bur. Fish., 34, 1916: 268 (N. Carolina); Bigelow and Schroeder, Guide Comm.
Shark Fish., Anglo Amer. Caribb. Comm., Wash., 1945: 148, fig. 56 (descr., ill., discuss.).

Rhina squatina (in part) GUnther, Cat. Fish. Brit. Mus., 8, 1 870: 430 {dumerilii included in synonymy);
Day, Fish. Gt. Brit., 2, 1880-1884: 326 {dumerilii incl. in synonymy).

Squatina squatina (in part) Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 1896: 220 (Atlant. coast
U.S.); Bull. U.S. nat. Mus., 47 (l), 1896: 58 (descr., C. Cod southward; considered ident. with 'Emto-
pezn squatina) ; Bean, T. H., Bull. Amer. Mus. nat. Hist., p, 1897: 331 (near N. York) ; Smith, Bull. U.S.
Fish Comm., 17, 1898: 89 (Martha's Vineyard); Evermann and Kendall, Rep. U.S. Comm. Fish.
(1899), 1900: 49 (probably Florida) ; Bean, Rep. For. Comm. N. Y., 1901: 381 (old Martha's Vineyard
record) ; Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 45 (descr., near N. York) ; Smith, Bull. N. C. geol. econ.
Surv., 2, 1907: 38 (diagn., food, N. Carolina); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908:
10 (Woods Hole region); Tracy, Rep. R. I. Comm. inl. Fish., 1910: 61 (Newport and West Passage,
Rhode Island) ; Fowler, Proc. Acad. nat. Sci. Philad., 62, 191 1 : 599 (Delaware Bay and N. Jersey) ; Proc.
Acad. nat. Sci. Philad., 63, 191 1 : 5 (Virginia, Maryland, Delaware); Proc. Acad. nat. Sci. Philad., 64,
191 2: 35 (N.Jersey); Gudger, J. Elisha Mitchell sci. Soc, 28, 191 3: 166 (N. Carolina); Nichols, Abstr.
Linn. Soc. N. Y., 20-25, '9' 3= 9' (near N. York) ; Sumner, Osburn and Cole, Bull. U.S. Bur. Fish.,
31 (2), 1913: 737 (Martha's Vineyard); Coles, Proc. biol. Soc. Wash., 28, 1915: 92 (season, abund.,
N. Carolina); Nichols and Murphy, Brooklyn Mus. Sci. Bull., 3 (l), 1916: 33 (near New York);
Nichols and Breder, Zoologlca, N. Y., p, 1927: 22 (vicinity of N. York and Woods Hole) ; Breder, Field

36. This latter was taken by the United States Fisheries Steamer "Albatross 11."
36a. Spelled dumerilii.

544

Memoir Sears Foundation for Marine Research

Bk. Mar. Fish. Atlant. Coast, 1929: 26 (general) ; Jordan, Evermann and Clark, Rep. U.S.-Comm. Fish.
(1928), 2, 1930: 22 (N. to Cape Cod, but incl. European squatina) ; Breder, Bull. N. Y. zool. Soc, 41,
1938: 25 (Sandy Hook Bay, near N. York) ; Parks, Tech. Bull. Stephen Austin State Teachers' Coll., I
(4), 1939: I (Texas; not seen) ; Gunter, Amer. Midi. Nat., 26 (l), 1941: 197 (Corpus Christi, Texas) ;
not Squalus squalina Linnaeus, 1 75 8.

Rhina squatiiia Fowler, Proc. Acad. nat. Sci. Philad., 58, 1906: 80 (reported for Florida Keys).

Not Squatina squatina Fowler, Arqu. Zool. Estado Sao Paulo, 2, 1 942: 129 (listed for Rio de Janeiro, Brazil;
probablv argentina Marini, 193O; see p. 546).

Genus Squatina, Addendum

The common Squatina of the temperate coasts of the western South Atlantic seems
clearly separable from all other members of the genus. A short notice of it therefore
follows.

Squatina argentina Marini, 19 30
Figure 106

Study Material. Female, 714 mm. long, from Uruguay (U.S. Nat. Mus., No.
87684).

Distinctive Characters. See Description.

Description. Proportional dimensions in per cent of total length. Female, 714 mm.,
from Uruguay (U.S. Nat. Mus., No. 87684).

Figure 106. Squatina argentina, female, 714 mm. long, from Uruguay (U. S. Nat. Mus., No. 87684).

Fishes of the Western North Atlantic 545

Extreme breadth at outer extremity of -pectorals: 54.O.
Trunk at origin of pectoral: height 8.0.
Snout length in front of: outer nostrils Oj mouth o.
Eye: horizontal diameter 1.55 distance between eyes 8.4.
Spiracles: distance between 8.1 ; from spiracle opening to eye 2.7.
Mouth: breadth 11.8} height 3.2.
Nostrils: distance between inner ends 4.8.

Gill opening lengths: ist7.l} 2nd 6.7; 3rd 6.6; 4th 6.2; 5th 6.0.
First dorsal fin : vertical height 4.9 ; length of base 4. i .
Second dorsal Jin: vertical height 4.5 ; length of base 4.0.
Caudal fin: upper margin 12.O; lower margin 14.3.
Pectoral fin: outer margin 26.5; inner margin 13.8; distal margin 14.0.
Distance from snout to: ist dorsal 65.6; 2nd dorsal 76.O; upper caudal 88.0; pec-
toral 16.8; pelvics 37.8.

Interspace between: ist and and dorsals 7.O; 2nd dorsal and caudal 8.5.
Distance from origin to origin of: pectoral and pelvics 20.9; pelvics and caudal
49.8.

S. argentina resembles S. dumeril so closely in general proportions and in the shape
of the pectoral fin" that we have found no clear distinction between the two species in
most of their proportional dimensions, and the teeth are of the same number ( ^^^ ) and
shape. However, the distance from the spiracle to the eye is only a little more than V2
(55% ) as long as the diameter of the eye in argentina, but nearly as long as the eye (91%)
in dumeril. The inner nasal barbel of argentina is spatulate (tapering in dumeril), while
there is a broad quadrate lobe between the two barbels, with a second lobe-lilce expansion
just basal to the outer margin of the outer barbel on the outer side in argentina, which is
not the case in dumeril (cf. Fig. 103 D with 103 G). A further difference if minor is that
the dermal fold along the front of the head is not expanded at all opposite the corner of the
mouth in argentina, but is slightly expanded there in dumeril (although much less so than
in the European squatina; Fig. 103 C).

S. argentina differs from armata of the west coast of South America in the contour
of the margin of the nostril (cf. Fig. 103 D with 103 H) ; it differs further from armata
in that the distal margin of its pectoral is marked off from the inner margin by a definite,
narrowly rounded corner, whereas in armata the one margin grades insensibly into the
other in an even curve.

Color. Described as gray-brown above, marked with many small oval cinnamon-
colored spots; white below."

Size. Said to grow to a length of 1.7 meters (5V2 feet).

37. In the original illustration of argentina (Marini, Physis B. Aires, lo, 1930: 6) the outer corner of the pectoral
is shown as considerably more obtuse than a right angle. But it is only a little more than a right angle in the
specimen we have studied, and Ribeiro's (Arch. Mus. nac. Rio de J., r^, 1907 : pi. 10) photograph of a Brazilian
specimen shows about a right angle.

j8. The only specimen we have seen has not only lost the color pattern, but is now stained red with iron rust.

546 Memoir Sears Foundation for Marine Research

Range. Temperate latitudes in the western side of the South Atlantic j northern
Argentina (the type locality), and apparently common along the coasts of Uruguay and
southern Brazil as far north as Rio de Janeiro/"

Synonyms and References:

Squatina squatina Schreiner and Ribeiro, Arch. Mus. nac. Rio de J., 12, 1903: 80 (near Rio de Janeiro);

Ribeiro, Feces "Annie," 1904: 18 (off Ilha Rassa, Brazil); Arch. Mus. nac. Rio de J., 14, 1907: 170,

pi. 10 (descr., ill., Rio de Janeiro); Devincenzi, An. Mus. Hist. nat. Montevideo, (2) 1, 1920: 124

(abund., Uruguay); Ribeiro, Fauna brasil. Peixes, 2 (l) Fasc. I, 1923; 27, pi. 9 (descr., ill., Rio de

Janeiro) ; Devincenzi and Barattini, An. Mus. Hist. nat. Montevideo, Suppl. Album Ictiol., 1926: pi. 3,

fig. I (ill., Uruguay); Fowler, Arqu. Zool. Estado Sao Paulo, j, 1942: 129 (listed for Bay of Rio de

Janeiro, Biazil) ; not Squdus squatina Linnaeus, 1758.
Squatina angelus Lahille, Physis B. Aires, 5, 1 92 1 : 63 (listed for Argentina) ; Enum. Feces. Cartilag. Argent.,

1921: 17 (listed for Argentina); Pozzi and Bordale, An. Soc. cient. argent., 120, 1935: 152 (listed

for Argentina, Lat. 3 5 "-47° S.).
Rhina angelus Marini, Physis B. Aires, 9 (34), 1929: 422 (listed for north. Argentina) ; not Squatina angelus

Blainville, 1816 (sometimes referred, but incorrectly, to Dumeril, A. M. C, Zool. Anallt, 1806: 102);

same as Squalus squatina Linnaeus, 1758.
Rhina argentina Marini, Physis B. Aires, 10 (35), 1930: 5, I fig. (descr., good ill., meas., Argentina, Lat.

39° S.) ; Pozzi and Bordale, An. Soc. cient. argent., 120, 1935: 15: (listed for Argentina, Lat. 39° S.).
Squatina annata Norman, "Discovery" Rep., 16, 1937: 10 (descr., comp. with other sp., Argentina); Hart,

"Discovery" Rep., 23, 1946: 260 (Argentina by ref. to Norman, 1937); not Rhino armata Philippi,

1887.

39. If our reference to it of the several Uruguayan and Brazilian records for angelus and squatina be correct.

INDEX OF COMMON NAMES

(Principal references in boldface)

A.ICUILLAT, squale, 470.

Albacore, 192.

Albatross, 480.

Alewives, 104.

Algae, 462.

American pollock, 49, 54, 286, 520.

Ammocoete, 43.

Amphipods, 462.

Anadromous herring, 49, 55.

Anchovies, 152, 192, 286.

Angel shark, 538.

Angler, 542 note 26.

Argentine, 498.

Atlantic Right Whale, 155, 523.

Aveugle, gastrobranche, 42.

Jdalance fish, 444.
Barbillon, squale, 1 86.

chien de mer, 185.
Barnacles, 424, 441.
Barracuda, 69 note 12.

Basking shark, 49, 68, 70 note 17, 74, 146, 147,
156, 160 note 43, 192.

great, 194.
Bass, 441.

sea, 104.
Beaumaris shark, 120.
Big-eyed thresher, 163, i68.
Birds, 172, 270, 286, 521.
Black dogfish, 482.
Black drum, 104.
Black-nosed shark, 325.
Black-tipped shark, large, 364.

small, 346.
Blennies, 462.
Bluefish, 104, 172.
Blue shark, 69, 70 note 17, 112, 289.

great, 282.
Bone shark, 147.
Benito, 104, 172, 373.
Bonnet shark, 420.
Borer, 42.

Boucle, 530.

leiche, 531.

squale, 530.
Brown shark, 70 note 17, 368.
Bull shark, 337.
Butterfish, 104, 350.

VvANE carcharia, 142.

Capelin, 462, 520.

Carcharhinids, 65, 68, 233, 276, 316, 337, 359,

360, 363,405.
Carolinien, squale, 426.
Carpet shark, 178.
Catfish, gaflF-topsail, 441.
Cat shark, 195.

false, 228.
Cephalopods, 286, 424, 485.
Cetaceans, 521.
Chain dogfish, 207.
Chien de mer barbillon, 185.
Chimaeroids, 2, 62, 139.
Cladodont, 94.
Clupeids, 129, 394.
Clupeoids, 152.
Cockles, 463.
Cod, 39, 40, 49, 57, 117, 119 note 19, 286, 460,

461 note 33, 462, 463, 520, 521.
Common hammerhead, 436.
Common mackerel shark, 70 note 17.
Common thresher, 163, 167.
Cormorant, 542 note 26.
Cowfish, 313.
Cow shark, 78, 80.
Crabs, 68, 84, 104, 184, 248, 251, 270, 342, 373,

424, 441, 462, 521, 529.
horseshoe, 270.
Croakers, 462.
Crustacea, 4, 68, 153, 248, 373, 424, 441, 485,

542.
Ctenophores, 462.
Cuban dogfish, 473.
Cub shark, 337.

547

548

Index of Common Names

Cunner, 104.
Cusk, 1 17.
Cusk eels, 387.
Cyclostomcs, 2, 29, 62.

Uesmids, 4.

Devil-ray, 342.

Diatoms, 4.

Dogfish, 192, 373,470,472.

black, 482.

chain, 207.

Cuban, 473.

Florida, 254.

horned, 472.

piked, 453,455.470.

smooth, 66, 68, 105, 233, 240, 244.

spiny, 66 note 5, 73, 105, 1 17, 119 note 19,
139, 250, 286,453,455,478.

spotted, 196 note 2.

striped, 256.
Dolphins, 84.
Drum, black, 1 04.
Dusky shark, 70 note 17, 374, 382.

tLcTHNODERMS, 4.

Eel grass, 1 19, 248.
Eel, lamper-, 57.
Eels, 49, 104, 373.

cusk, 387.
Elasmobranchs, 2, 62, 75.

modern, 64.
European weaver, 461.

False cat sharks, 228.

File tails, 1 96 note 8.

Fishes, 270, 373, 381, 387, 394, 424. 441. 463-

FLitfishes, 104, 387, 520, 542.

Florida dogfish, 254.

Flounders, 117,373, 542.

Flyingfish, 286.

Fowl, sea, 270.

Fox, sea, 174.

(jADOiDS, 39, 117.
Galeoid shark, 65.
Galeoids, 75, 76, 96 note 9.
Galluda, 478.
Garfish, 172.

Gastrobranche avcugle, 42.
Gastrobranchus, glutinous, 42.
Gastropods, 270, 462, 542.

Gata, 18;.

Glauque, squale, 289.

Glutinous gastrobranchus, 42.

Glutinous hag, 42.

Goatfish, red, 387.

Goblin shark, 109.

Grayfish, 73, 472.

Great basking shark, 194.

Great blue shark, 282.

Great hammerhead, 428, 441.

Great white shark, 144, 145.

Greenland halibut, 520.

Greenland shark, 59, 66, 67, 71, 72, 74, 515, 516,

524.
Grey notidanus, 86.
Grey shark, 80.
Griset, 85.
Griset, squale, 85.
Ground shark, 337-
Groupers, 387.
Gurry shark, 516.

tlAA-KlAERRINCEN, 523.

Haa-Skierding, 523.

Haddock, 37, 39, 49, 286, 462, 463, 520.

Hagfish, 34.

Hag, glutinous, 42.

Hags, 31,32.

Hake, 39, 49, 84, 91, 117, 119.

silver, 104, 462.
Halibut, 520.

Greenland, 520.
Hammerhead, 65, 69, 72, 74, 415, 447, 449.
common, 436.
great, 428, 44 1. .
Hammerhead shark, 407, 420.
Hammer headed shark, 434.
Heart headed shark, 426.
Herring, 39, 117, 172, 286, 441, 462, 520.

anadromous, 49, 55.
Heterodontoids, 76.
Hcterodonts, 68, 75.
Hexanchids, 75.
Horned dogfish, 472.
Horseshoe crabs, 270.
Hybodoids, 75.
Hybodonts, 64, 480.

sopoDS, 424.

Jellyfish, 462.
John Dory, 117.

Index of Common Names

549

K-iLLERwhale, 172.

1_/AKE lamprey, 46, 57.
Lake Nicaragua shark, 378.
Lamie nez, 120.
Lamper-eel, 57.
Lamprey, 43, 45, 57.

lake, 46, 57.

sea, 46, 56.
Lancelets, I.

Large black-tipped shark, 364.
Leiche boucle, 531.
Lemon shark, 69, 74, 310.
Leopard shark, 266.
Liche, 506.

squale, 506.
Ling, 39, 520, 521.
Lizard fish, 387.
Lobster, spiny, 184, 270.
Lobsters, 68, 104, 248.
Long nez, squale, 120.
Long tailed shark, 174.
Loutre de mer, 1 20.
Lumpfish, 520.
Lung fishes, 62.

JVIackerel, 39, 49, 55, 117, 119 note 19, 129,
130, 172, 270, 286, 342, 373, 461, 462, 472.

Spanish, 68, 441.
Mackerel shark, 39, 66, 103, 109, 112.

common, 70 note 17.

sharp-nosed, 112, 123, 124.
Mako, i9, 69, 103, 104, 117, 118 note 16, 123,
124, 139, 140, 14;.

Pacific, 123, 128.
Mako shark, I 3 2.
Man-eater, 134.
Man-eater shark, lOg.
Mantis shrimps, 424.
Marlin, 84.
Marteau, 419, 434, 444.

squale, 434, 444.

zygene, 448.
Medusae, 485, 52 1.
Menhaden, 104, 172, 248, 300, 332 note 14, 350,

373. 441. 462.
Modern elasmobranchs, 64.
Modern sharks, 64, 532.

Mollusks, 68, 248, 300, 373, 424, 541, 542.
Mud shark, 78, 80.
Mullet, 104.

JNarwhals, 521.

Nez, lamie, 1 20.

Night shark, 316.

Notidanoids, 76, 94.

Notidanus, grey, 86.

Nurse shark, 74, 178, 180, 181.

Oil shark, 72, 73, 264 note 4.
Orectolobids, 75, 233.

Xacific mako, 123, 128.
Pantouflier, 426.

squale, 426, 434.
Parrotfish, 300.
Pelerin, squale, I 57.
Pelerin tres grand, 158.
Perlon, 92.

squale, 92.
Piked dogfish, 453, 455, 470.
Pilchards, 117, 172.
Pinfish, 373.
Pleuracanth, 94.
Pleuronectids, 542.
Pointer, white, 145.
Pollock, 462.

American, 49, 54, 286, 520.
Polychaete worms, 37.
Porbeagle, 70 note 17, 112, 119, 271.
Porpoise, 270, 342.
Portuguese shark, 66, 494.
Prawns, 462.
Pristiophorids, 532.
Pristiophoroids, 76.
Protozoa, 5.
Puffers, 248.

K.AYS, 62, 03, 64, 65, 77, 542.

devil-, 342.
Red goatfish, 387.
Reindeer, 521.
Remoras, 152.
Renard, squale, I 74.
Requin, 362.

squale, 142, 363.
Rosefish, 520.
Round headed zygaena, 426.

o AiTHE, 520.
Salmon, 49, 520, 521.
Sand-bar shark, 368.

550

Index of Common Names

Sand shark, 70 note 17, 74, 98, 100.

Sand tiger, lOO.

Sardines, 192, 286, 350.

Sawfishes, 77 note 36, 532.

Saw sharks, 77 note 36.

Sciaenids, 394.

Scombroids, 129.

Sculpins, 248, 520.

Scup, 104, 248, 441, 462.

Scyliorhinids, 67, 196 note 10, 197, 215, 216,

233.
Sea bass, 104.
Sea fowl, 270.
Sea fox, 174.
Sea lamprey, 46, 56.
Sea lion, 139, 270.
Sea robin, 104.
Sea serpents, 153.

Sea turtles, 68, 69, 129, 139, 270, 360.
Sea urchins, I 84.
Seals, 68, 70, 139, 520, 521.
Seaweed, 424.
Serpents, sea, I 53.
Seven-gilled shark, 88.
Shad, 49, 55, 172, 342.

Shark or Sharks, 30, 59, 63, 270, 328, 342, 419,
441,443,463,492, 529.

angel, 538.

basking, 49, 68, 70 note 17, 74, 146, 147, 156,
160 note 43, 192.

Beaumaris, 120.

black-nosed, 325.

blue, 69, 70 note 17, 112, 289.

bone, 147.

bonnet, 420.

brown, 70 note 17, 368.

bull, 337.

carpet, 178.

cat, 195.

cow, 78, 80.

cub, 337.

dusky, 70 note 17, 374, 382.

false cat, 228.

galeoid, 65.

goblin, 109.

great basking, 194.

great blue, 282.

great white, 144, 145.

Greenland, 59, 66, 67, 71, 72, 74, 515, 516,
524.

grey, 80.

ground, 337.

gurry, 516.

hammerhead, 407, 420.

hammer headed, 434.

heart headed, 426.

Lake Nicaragua, 378.

large black-tipped, 364.

lemon, 69, 74, 310.

leopard, 266.

long tailed, 174.

mackerel, 39, 66, 103, 109, 1 12.

mackerel, common, 70 note 1 7.

mako, 132.

man-eater, 109.

modern, 64, 532.

mud, 78, 80.

night, 316.

nurse, 74, 178, 180, 181.

oil, 72, 73, 264 note 4.

Portuguese, 66, 494.

sand, 70 note I 7, 74, 98, 100.

sand-bar, 368.

saw, 77 note 36.

seven-gilled, 88.

sharp-nosed, 295.

sharp-nosed mackerel, 112, 123, 124.

silky, 333.

six-gilled, 78, 80, 86.

small black-tipped, 346.

soupfin, 72, 73.

spinous, 531.

spiny, 527.

swell, 197, 198.

tawny, 178 note 2.

thresher, 161.

tiger, 69, 74, 266, 342.

whale, 67, 68, 74, 151, 187, 188, 189.

whaler, 320 note 2.

white, 65, 69, 103, 104, 134, 142, 354 note 51.

white-tipped, 354.

zebra, 178 note 2, 179.
Shark sucker, 104, 193, 273.
Sharp-nosed mackerel shark, 112, 123, 1 24.
Sharp-nosed shark, 295.
Shovel head, 420.
Shovelnose, 382.
Shrimps, 84, 184, 300, 424, 441, 462.

mantis, 424.
Silky shark, 333.
Silver hake, 104, 462.
Six-gilled shark, 78, 80, 86.

Skates, 63, 64, 65, 77 note 36, 270, 320 note la,
373,441, 520, 542.

Index of Common Names

551

Sleep marken, 42.

Small black-tipped shark, 346.

Smooth dogfish, 66, 68, 105, 233, 240, 244.

Snails, 521.

Soupfin shark, 72, 73.

Spadefish, 104.

Spanish mackerel, 68, 441.

Sperm whale, 286.

Sphyrnids, 409.

Spinax, 470.

Spinous shark, 531.

Spiny dogfish, 66 note J, 73, 105, 117, 119 note

19, 139, 250, 286,453,455,478.
Spiny lobster, i 84, 270.
Spiny shark, 527.
Spot, 104.

Spotted dogfish, 196 note 2.
Squale, 272.
Squale aiguillat, 470.
Squale barbillon, 186.
Squale boucle, 530.
Squale chien de mer, 185.
Squale carolinien, 426.
Squale glauque, 289.
Squale griset, 85.
Squale leiche, 231.
Squale liche, 506.
Squale long nez, 1 20.
Squale marteau, 434, 444.
Squale pantouflier, 426, 434.
Squale pelerin, 157.
Squale perlon, 92.
Squale glauque, 289.
Squale renard, 1 74.
Squale requin, 142, 363.
Squale tres grand, I 5 7.
Squalid, 516, 519 note 15.
Squalo, 85.
Squaloids, 75, 76.
Squatinoids, 75, 76, 533.
Squids, 68, 104, 117, 129, 139, 172, 184, 192,

248, 250, 270, 271, 274, 279, 350, 441,

462, 521.
Sticklebacks, 248.

Sting-rays, 270, 313, 314, 342, 350, 441.
Stone sucker, 46.
Striped dogfish, 256.
Sturgeon, 39, 49, 139.

Sucker, shark, 104, 193, 273.

stone, 46.
Swell shark, 197, 198.
Swordfish, 49, 52, 57, 84, 129, 140, 172, 472.

Iautoc, 104, 248.
Tawny shark, 178 note 2.
Teleost, 463.
Thresher, big-eyed, 163, 168.

common, 163, 167.
Thresher shark, 161.
Tiburon de noche, 319.
Tiger, sand, lOO.
Tiger shark, 69, 74, 266, 342.
Tope, 70 note 17, 264 note 4.
"Topes," 233.
Torpedo, 84, 444.
Touille-boeuf, 120.
Tres grand, 157.

pelerin, 158.

squale, 157.
Triakids, 67, 233, 234.
Tuna, 139, 140.
Tunicates, 4.

Weakfish, 104, 373.

Weaver, European, 461.

Whale shark, 67, 68, 74, 151, 187, 188, 189.

Whale, 68, 172, 521.

Atlantic Right, 155, 523.

killer, 172.

sperm, 286.

whalebone, 152.
Whalebone whale, 152.
Whaler shark, 320 note 2.
White pointer, 145.

White shark, 65, 69, 103, 104, 134, 142, 354
note 5 1 .

great, 144, 145.
White-tipped shark, 354.
Whiting, 117.
Wolffish, 520.
Worms, 248, 462.

polychaete, 3 7.

Z/EBRA shark, 178 note 2, 179.
Zygaena, round headed, 426.
Zygene marteau, 448.

INDEX OF SCIENTIFIC NAMES

(Principal references in boldface)

Abbotti, Mustelus, 242 note 4, 260 note 38,

261.
Acanthias, 4.52, 487.

acanthias, 471, 478.

americanus, 471.

blainville, 480.

blainvillei, 480.

blainvillii, 480.

fernandezianus, 480.

lebruni, 453 note 14.

linnei, 472.

suckleyi, 472.

sucklii, 472.

vulgaris; 452, 454, 470, 472, 478.
acanthias, Acanthias, 471, 478.

Acanthorhinus, 470.

Spinax, 470, 472, 486.

Squalus, 65 note 4, 66, 73, 117, 250, 286, 452,
453. 454, 455> 467, 47° note 54, 472, 473,
474. 475, 476, 477, 478, 479, 482, 486, 490,
494. 529-

Squalus (Acantherinus), 471.

Squalus (Spinax), 471.

var. fernandinus, Squalus, 480.
Acanthidim, 487.
Acanthidium, 451 note 4, 487.

pusillum, 487.
Acanthorhinus, 452, 487, 5 14.

acanthias, 470.

carcharias, 526.

fernandinus, 480.

norwegianus, 523.
achantias, Squalus, 470.
acrages, Pseudotriakis, 229, 232.
acrales, Pseudotriakis, 229 note 3.
acronotus, Carcharhinus, 324, S^S-

Carcharhinus (Platypodon), 328.

Carcharias, 329.

Carcharias (Prionodon), 328.

Carcharinus, 328.

Eulamia, 329.

Platypodon, 328.

Squalus, 328.
aculeatus, Etmopterus, 487.

Carcharias, 304 note 2.
acutidens, Carcharias, 304 note 2.

Negaprion, 309.

queenslandicus, Aprionodon, 308.
acutipinnis, Squalus, 454 note 2ia, 480.
acutus, Carcharias, 302, 303.

Carcharias (Scoliodon), 302.

Scoliodon, 303.
adscensionis, Squalus, 289.
(aetatis), Notidanus (Heptanchus) cinereus, var.

pristiurus, 93.
aethiops, Carcharias, 291.
aethlorus, Carcharhinus, 352.

Carcharhinus (Isogomphodon), 352.

Carcharias, 351 note 45, 352.

Eulamia, 353.
affinis, Myxine, 33, 38, 42.
africae, Branchiostoma, 9, 10, II, 14, 16.
africana, Squatina, 537.
africanum, Poroderma, 197 note 16, 202.
africanus, Squalus, 197 note 16, 202.
Agnatha, 30.

albiraors, Carcharodon, 134 note 3, 145.
Alocepias vulpes, 176.
Alopecias, 161.

barrae, 177.

chilensis, 177.

longiraana, 177.

superciliosus, 167.

vulpes, 175, 177.
alopecias, Squalus, 1 76.
Alopes, 161.

vulpes, 161, 177.
Alopias, 161, 167, 320 note la.

caudatus, 162, 177 note 33, 178.

greyi, 162, 177 note 33, 178.

macrourus, 1 6 1, 174.

pelagicus, 161, 162.

profundus, 162, 167.

superciliosus, 162, 163, 168.

55i

Index of Scientific Names

553

vulpes, 175, 177, 320 note la.

vulpinus, 161, 162, 164, 166, 167, 176, 177,
178.

vulpis, 175.
Alopiidae, 96, 160.
Alopiopsis, 292.
Alosa, 4.9, 172, 34.2.

amblyrhynchus, Carcharias (Prionodon), 321.
americanus, Acanthias, 471.

Carcharias, 108.

Dalatias, 501, 508.

Odontaspis, 108, 109.

Petromyzon, 56.

Scymnus, 506.

Squalus, 100, 472, 501, 506.

Squalus (Acanthorhinus), 507.
(americanus), Petromyzon marinus, 57.
Amia, 463 note 42.
Ammocaetes, 45.
Ammocites, 45.
Ammocoetes, 45.

appendix, 58.

bicolor, 57.

branchialis, 56.

concolor, 57.

fluviatllis, 57.

unicolor, 57.
Ammocoetus, 45.
Amphioxi, 3.
Amphioxides, 7, 22, 23, 24, 26, 27.

"Larvae," 7, 23, 24, 25, 27.

pelagicus, 22, 23, 24, 25, 26, 27, 28.

stenurus, 24, 25, 27.

valdiviae, 20, 24, 25, 26, 27, 28.
Amphioxididae, 7.
Amphioxos, 18.
Amphioxus, 8, 13, 16.

caribaeus, 13, 16.

lanceolatus, 16.

miilleri, 17.
Amphipleurichthys, 8.

minucauda, 8.
anale, Scyllium, 198 note 22.
Anarrhichas, 520.
Angel i formes, 534 note 9.
angelus, Rhina, 546.

Squatina, 543, 546.
angio, Heptrancus, 93.
angiona, Tetraoras, 146, 160.
angiova, Tetrorus, 160.
Anguilla, 104.
anguineus, Chlamydoselachus, 94.

Antacea, 76.

antarcticus, Somniosus, 5 i 5.

appendix, Ammocoetes, 58.

Aprion, 303.

Aprionodon, 265, 303, 308, 321, 322.

acutidens quecnslandicus, 308.

brevipinna, 205 note 5, 304, 308.

isodon, 304, 307.

punctatus, 303, 307.

sitakaiensis, 309 note 3.
Aprist.urius, 219.
Apristurus, 198, 202, 219.

atlanticus, 220, 221, 222, 226.

brunneus, 220, 224 note 13.

herklotsi, 220, 221 note 10.

indicus, 221.

laurussonii, 221 note 11, 224, 226.

macrorhynchus, 221 note 8.

microps, 220, 221.

platyrhynchus, 220, 221 note 10.

profundorum, 196 note 9, 220, 221, 222, 225,
226, 228.

riveri, ig6 note 9, 221, 222, 223, 224, 225,
228.

saldanha, 221.

sibogae, 221.

spongiceps, 220, 221 note 10, 513.

verweyi, 220, 221 note 10.
aquiUa, Sciaena, 524.
arae, Galeus, 210, 211, 215, 216.

Pristiurus, 219.
arcticus, Boreogaleus, 273.

Galeocerdo, 272, 274, 275.

Squalus, 265, 271, 272.
arenarius, Carcharias, 99.
Argentina silus, 498.
argentina, Rhina, 546.

Squatina, 535, 536 note 16, 537, 544.
argus, Squalus, I 86.
armata, Rhina, 546.

Squatina, 535, 537, 545, 546.
asterias, Mustelus, 242, 254, 260, 261.
Asterospondyli, 76.
Asymbolus, 198.
Asymmetron, 4, 18, 23, 24.

caudatum, 19, 22.

lucayanum, 18, 19, 22, 25, 26 note 58.

macricaudatum, 19, 21 note 40, 22.

orientale, ig, 23.

pelagicum, 26.
Asymmetrontidae, 18 note 34.
Atelomycteridae, 195 note I.

554

Index of Scientific Names

Atelomycterus, 199, zoi.

macleayi, 199 note 23.
atlantica, Myxine, 38, 43.
atlanticus, Apristurus, 220, 221, 222, 226.
Atolla, 485.
atwoodi, Carcharias, 1 43.

Carcharodon, 143, 145.
Aulohalaelurus, 198.
Aurelia, 462.

auriculatus, Carcharias, 133.
australis, Myxine, 33 note 8, 38, 42.

Squatina, 535, 536.
azureus, Carcharhinus, 309 note I, 373, 376.

iJAiRDii, Bathymyzon, 45, 46, 57.

Petromyzon (Bathymyzon), 57.
Barborodes, 501.
barbouri, Squalus, 455 note 23, 464, 469, 472.

Triakis, 235, 236, 240.
barrae, Alopecias, 177.
Bathyamphioxus, 18 note 36.
Bathymyzon, 45, 47 note 57, 55.

bairdii, 45, 46, 57.
Batoidei, 64, 76, 77 note 36, 532.
bazarutense, Branchiostoma, 9.
Bdellostoma, 32 note 5.
belcheri, Branchiostoma, 4, 6, 10, 16.
Belone, 172.

bermudae, Branchiostoma, 9, 10, 11, 13, 14, 15, 17.
bicolor, Ammocoetes, 57.
bideni, Isurus, 123, 124 note 30.
bigelowi, Sphyrna, 410, 414, 416, 420, 428, 435,

436.
blainville, Acanthias, 480.

Squalus, 453, 454, 455, 456, 472, 474, 478 note
63,479,480.
blainvillei, Acanthias, 480.

Squalus, 478, 480.
blainvillii, Acanthias, 480.

Cetorhinus, 160.
blochii, Eusphyra, 407 note I, 444.
boa, Catulus, 207.

Catulus retifer, var., 207.

Scyliorhinus, 204, 211, 212, 213.

Scylliorhinus, 207, 214.
boae, Catulus, 2 1 4.
boardmani, Figaro, 214.

Galeus, 215.
Bogimba, 321.

bogimba, Galeolamna (Bogimba), 321.
borealis, Dalatias (Somniosus), 525.

Laemargus, 524.

Scimnus, 526.

Scymnus, 514, 524.

Scymnus (Laemargus), 514, 524.

Squalus, 514, 524.
Boreogaleus, 265.

arcticus, 273.
boshuensis, Pseudoscymnus, 501, 508.
Brachaelurus, 178, 1 79 note 3, 180.
brachyurus, Carcharhinus, 360.

Carcharias, 320 note 2, 346 note 40.

Etmopterus, 488 note 3.
branchialis, Ammocoetes, 56.

Petromyzon, 45, 56.
Branchiostoma, 4, 8, 18, 23.

africae, 9, 10, 11, 14, 16.

bazarutense, 9.

belcheri, 4, 6, 10, 16.

bermudae, 9, 10, II, 13, 14, 15, 17.

californiense, 10.

capense, 9.

caribaeum, 6 note 10, 9, 10, 11, 12, 13, 15, 16,

17-

caribbaeum, 13, 15 note 30, 16.

carribaeum, 13, 17 note 33.

elongatum, 10.

floridae, 9, 10 note 20, 16.

gravelyi, 10 note 19.

haeckeli, 9.

lanceolata, 16.

lanceolatum, 4, 5, 9, 10, II, 13, 14, 15, 16, 24.

lubricum, 8, 13, 16.

lucayanum, 22.

minucauda, 9.

pelagicum, 23, 24, 26, 28.

platae, 9, 10, II, 12, 14, 15, 16, 17, 18.

tattersalli, 10.

virginiae, 9, 10 note 20, 16.
Branchiostomidae, 7.
brasiliensis, Dalatias, 513.

Isistius, 485, 509, 513, 514 note 9.

Leius, 513.

Scymnus, 508, 513.

Scymnus (Scymnus), 513.

var. torquatus, Scymnus (Scymnus), 513.

var. unicolor, Scymnus (Scymnus), 513.
brevicaudatum, Ginglymostoma, 181.
brevipinna, Aprionodon, 265 note 5, 304, 308.

Carcharias (Aprion), 303.

Laemargus, 526.

Longmania, 265 note 5.

Scymnus, 524.

Scymnus (Laemargus), 526.

Index of Scientific Names

555

Somniosus, 514, 515 note 5, 523, 524.

Squalus, 515 note 5, 523.
brevipinnis, D.ilalias, 50 1, 508.

Scymnorhinus, 508.
brevirostris, Carcharhinus, 315.

Carcharhinus (Hypoprion), 315.

Carcharias, 315.

Carcharias (Hypoprion), 315.

Hypoprion, 314, 315 note 2.

Negaprion, 69, 72, 309, 310, 337, 338, 388,

390-

Squalus, 453, 454.
Brevoortia, 104, 172, 248, 300, 332 note 14, 350,

373,441,462.
broussonetii, Carcharhinus, 320 note la.
brucus, Echinorhinus, 60, 51 3, 5 14, 527, 53 i, 532.

Squalus, 514, 526, 530.
brunneus, Apristurus, 220, 224 note 13.

Catulus, 219.
Buccinum, 270.

Cjaboverdianus, Ginglymostoma, 187.
caecchia, Carcharias, 378.

Squalus, 377.
caecus, Gastrobranchus, 42.
caerulea, Sardinops, 350.
caeruleus, Carcharhinus, 320 note la.

Prionace, 320 note la.

Squalus (Carcharhinus), 290.

Squalus (Carcharinus), 376.
calamaria, Longmania, 321.
californica, Squatina, 535, 537, 540 note 24.
californicus, Mustelus, 240, 242.
californiense, Branchiostoma, 10.
Calliscyllium, 234 note 5, 235.

venusta, 235.
canicula, Scyllium, 96 note 8, 214.

Scyliorhinus, 202 note 25.
caniculus, Galeus, 202.

Scyliorhinus, 67, 96 note 8, 196 note 2, 197,
202, 203.

Squalus, 202.
Caninoa, ig6 note 1 1.
Canis carcharias, 272.
canis, Cynais, 253, 256.

Cynias, 253, 254.

Galeus, 253, 254, 264 note 4.

Mustelus, 68, 97 fig. 12, 243, 244, 251, 254,
255, 256, 257 fig. 43, 258, 259, 260, 261,
262.

Squalus, 240, 2; I.
capense, Branchiostoma, 9.

capensis, Carcharodon, 144.

Myxine, 33 note 8, 38.

Scyliorhinus, 203.
Carcharhinidae, 97, 233, 262, 298 note 7, 304,

308, 310, 315,407-
Carcharhinus, 69, 74, 133, 142, 161, 265, 280,
281, 284 note 12, 292, 303, 308, 309 note I,
320, 342, 359, 360, 361, 362, 363, 364, 374,
375 note 98, 376 note 103a, 377 note 104,
380, 392.

acronotus, 324, 325.

(Platypodon) acronotus, 328.

aethlorus, 352.

(Isogomphodon) aethlorus, 352.

azureus, 309 note I, 373, 376.

brachyurus, 360.

brevirostris, 31 5.

(Hypoprion) brevirostris, 315.

broussonetii, 320 note la.

caeruleus, 320 note la.

carcharias, 145.

(Eulainia) caudatus, 377.

cerdale, 398 note 143, 399.

commersonii, 320, 342, 343 note 34, 344, 345,
346, 353. 363. 388 note 121, 390.

cornubicus, 1 20, 320 note I a.

falciformis, 323, 329, 333, 334, 335, 336, 337,

384-
(Platypodon) falciformis, 333.
floridanus, 323, 329, 332, 333, 337, 384.
galapagoensis, 405.
glaucus, 320 note I a.
henlei, 399.

(Platypodon) henlei, 399.
heterobranchialis, 320 note la.
heterodon, 320 note la.
(Aprionodon) isodon, 308.
lamia, 272, 320 note la, 344, 363, 364.
(Carcharhinus) lamia, 362.
(Eulamia) lamia, 362.
lamiella, 399.
leucas, 72, 74, 320 note la, 324, 337, 344, 345,

346, 353> 354 note 51, 359, 361, 362, 363,

364, 378, 380, 381, 382, 386, 387, 388, 390.
limbatus, 72, 270, 325, 345, 346, 352, 353, 364,

366, 367, 368, 388, 400, 403.
(Isogomphodon) limbatus, 352.
lividus, 320 note la.
longimanus, 320 note la, 323, 324, 337, 338,

340, 341, 342, 343, 344, 345, 353, 354, 363

note 71, 384, 386, 387.
mackei, 291.

5S6

Index of Scientific Names

maculipinnis, 70, 325, 346, 348, 349, 351, 364,

400.
megalops, 3 20 note i a.
milberti, 70 note 17, 72, 139, 301 note 17,

324, 342, 343, 345, 346, 368, 377, 378,

381, 382, 384, 385, 386, 387, 388, 389,

390.
(Carcharhinus) milberti, 377.
monensis, 320 note I a.
natator, 351 note 45, 353.
nicaraguensis, 324, 337, 339, 341, 342, 378,

382.
(Carcharhinus) nicaraguensis, 382.
obscurus, 70 note 17, 72, 323, 342, 343, 344,

345. 346, 361, 372 note 85, 374, 377, 382,

389, 390, 391, 404, 405, 406, 407.
(Eulamia) obscurus, 389.
(Platyodon) obscurus, 390.
(Platypodon) obscurus, 390.
oxyrhynchus, 323, 391.
(Isogomphodon) oxyrhynchus, 394.
(Platypodon) perezi, 352.
platyodon, 344, 345, 354 note 52a, 377.
(Carcharhinus) platyodon, 344.
(Eulamia) platyodon, 344.
platyrhynchus, 405.
plumbeus, 368 note 82, 378, 390.
porosus, 323, 394.

remotus, 324, 352 note 50, 364, 400, 403.
(Platypodon) remotus, 403.
springeri, 324, 384, 386, 404.
terrae-novae, 302.
(Scoliodon) terrae-novae, 302.
ustus, 32c note la.
velox, 391.
verus, 320 note la.
vulpes, 320 note I a.
Carcharhynus, 394 note 138.

Carcharias, 65, 66 note 5, 74, 87, 98, 133, 142,
161, 280, 292, 303, 308, 315, 320, 452.
acronotus, 329.
(Prionodon) acronotus, 328.
acutidens, 304 note 2.
acutus, 302, 303.
(Scoliodon) acutus, 302.
aethiops, 291.

aethlorus, 351 note 45, 352.
(Prionodon) amblyrhynchus, 321.
americanus, 108.
arenarius, 99.
atwoodi, 143.
auriculatus, 133.

brachyurus, 320 note 2, 346 note 40.

(Aprion) brevipinna, 303.

brevirostris, 315.

(Hypoprion) brevirostris, 315.

caecchia, 378.

carcharias, 142, 345, 363.

cerdale, 399.

ceruleus, 375.

cinereus, 92.

coeruleus, 377.

(Eulamia) coeruleus, 376.

commersonii, 345, 363.

(Scoliodon) crenidens, 292.

eumeces, 302.

falciformis, 333.

(Prionodon) falciformis, 333.

falcipinnis, 389.

(Prionodon) fasciatus, 275.

ferox, 98, 99, 108.

fissidens, 398 note 144, 399.

fronto, 304 note 3.

glaucus, 289, 292 note 29.

(Prionodon) glaucus, 290.

(Prionodon) glyphis, 280 note 5, 321 note 4..

gracilis, 292.

griseus, 98, 108.

(Hypoprion) hemiodon, 315.

henlei, 399.

(Prionodon) henlei, 399.

(Prionodon) hirundinaceus, 290.

insularum, 361.

(Aprion) isodon, 303, 307.

(Aprionodon) isodon, 308.

(Scoliodon) lalandii, 301.

lamia, 142, 344, 346 note 40, 354 note 51, 359

note 58, 362, 363, 364, 403.
(Prionodon) lamia, 143, 344, 354 note 51, 359,

362, 363, 364.
lamiella, 399.

(Scoliodon) laticaudus, 292.
leucas, 144, 346, 363.
(Prionodon) leucas, 344.
leucos, 345.

(Prionodon) leucos, 344, 345.
limbatus, 353, 368.
(Prionodon) limbatus, 352.
littoralis, 99 note 15, 107.
longimanus, 346 note 40.
longurio, 303.
(Scoliodon) longurio, 303.
(Hypoprion) macloti, 315.
(Scoliodon) macrorhynchus, 292.

Index of Scientific Names

557

macrurus, 320 note 2, 321.

maculipinna, 352.

maculipinnis, 368.

maso, 145.

melanopterus, 321, 345, 353.

(Prionodon) melanopterus, 353.

(Prionodon) menisorrah, 321, 399.

microps, 353.

milberti, 376, 378.

(Prionodon) milberti, 320, 321 note 3, 375,

378,382.
(Prionodon) mulleri, 352.
natator, 353.
nicaraguensis, 382.
obscurus, 292, 377, 389, 390.
(Prionodon) obscurus, 389, 391.
obtusirostris, 362.
(Prionodon) obtusus, 362.
(Prionodon) obvelatus, 389.
owstoni, 99.

(Prionodon) oxyrhynchua, 321, 394.
palasorrah, 292, 293 note 2.
platensis, 99, 106.
platyodon, 344.
plumbeus, 377.
porosus, 303, 399.
(Prionodon) porosus, 399.
pugae, 290.

(Aprionodon) punctatus, 308.
remotus, 403.

(Prionodon) remotus, 403.
rondeletti, 142.
(Hypoprion) signatus, 319.
(Prionodon) sorra, 321.
sorrakovvah, 292.
spenceri, 321.
stevensi, 321.

taurus, 70 note 17, 98, 99, 100, 106.
(Prionodon) temmincki, 321.
terrae-novae, 302.
(Scoliodon) terrae-novae, 302.
tigris, 131.
tricuspidatus, 99.
verus, 133, 142.
vorax, 145.
vulgaris, 142.

vulpes, 161, 163, 174, 177.
(Scoliodon) walbeehmi, 303.
(Scoliodon) walbenii, 303.
carcharias, Acanthorhinus, 526.
Canis, 272.
Carcharhinus, 145.

Carcharias, 142, 345, 363.

Carcharodon, 69, 72, 133 note 2, 134, 143,
144, 320 note la.

Laemargus, 524.

Somniosus, 526.

Squalus, 133, 142, 144, 145, 273, 320 note la,
354 note 51,362, 514, 515, 523.

Squalus (Carcharhinus), 142.

Squalus (Carcharias), 1 42.
Carchariidae, 97, 98.
Carcharinida, 76.
Carcharinus, 292, 308, 320, 377, 391.

acronotus, 328.

commersonii, 344, 362.

glaucus, 290.

lamia, 345, 362, 363.

menisorrah, 399.

milberti, 363, 376, 377, 382.

(probably milberti), 377.

obscurus, 389, 391.

(probably obscurus), 391.

oxyrhynchus, 394.

platyodon, 346, 354 note 52a, 363.

porosus, 399.

remotus, 403.
Carcharocles, 133.

Carcharodon, 59, 65, 69, 70, 1 10, 1 1 2, 124, 133,
139, 140, 141 note 32, 143, 320 note la, 354
note 51, 362, 363, 364.

albimors, 1 34 note 3, 145.

atwoodi, 143, 145.

capensis, 144.

carcharias, 69, 72, 133 note 2, 134, 143, 144,
320 note la.

lamia, 142, 320 note la.

rondeletii, 134 note 3, 142, 144, 145.

smithii, 133 note I, 142.

verus, I 33, 142.
caribaeum, Branchiostoma, 6 note 10, 9, 10, 11, 12,

13, 15, 16, 17.
caribaeus, Amphioxus, 13, 15 note 30, 16.
caribbaeum, Branchiostoma, 13, 16.
carolinianus, Squalus, 428.
carribaeum, Branchiostoma, 13, 17 note 33.
Caspiomyzon, 44, 58.
castaneous, Petromyzon, 54 note 98a.
Catulidae, 196 note 3.
Catulus, 202, 219.

boa, 207.

boae, 214.

brunneus, 2ig.

cephalus, 196.

558

Index of Scientific Names

haeckelii, 207.

retifer, 21 1.

retifer var. boa, 207.

spongiceps, 219, 513.

major vulgaris, 202.
caudata, Lamna, 301 note 17, 375.

Squalus (Carcharinus), 376.
caudatum, Asymmetron, 19, 22.

Notasymmetron, 23.
caudatus, Alopias, 162, 177 note 33, 178.

Carcharhinus (Eulamia), 377.

Epigonichthys, 23.
Centrina, 487.

nigra, 487.
Centrophorides, 452 note 9.
Centrophorus, 451, 493, 526.

coelolepis, 499.

crepidater, 494 note I.

foliaceus, 45 1 note 8.

rossi, 45 I note 8.

spinosus, 526.

waitei, 45 I note 8.
Centropristis, 104.

Centroscyllium, 450, 480, 490, 494, 496, 502,
516.

fabricii, 482, 486, 487.

granulosum, 482, 487.

nigrum, 481.

ornatum, 481.

ritteri, 482.
Centroscymnus, 65 note 4, 66, 450 note I, 451,
493. 494. 5 H, 516, 519.

coelolepis, 485, 493, 494, 498.

cryptacanthus, 494, 499 note 7.

fuscus, 494.

macracanthus, 494 note I .

owstoni, 494.
Centroselachus, 451 note 7, 494 note I.
cepedianus, Galeus, 265, 274.
cepedii, Isurus, 132.

Squalus, 131, 133.
Cephalochordata, i.
Cephaloscyllium, 198, 201.
Cephalurus, 197, 199.
cephalus, Catulus, 1 96.
cerdale, Carcharhinus, 398 note 143, 399.

Carcharias, 399.

Eulamia, 399.
ceruleus, Carcharias, 375.
Cestracion, 408.

subarcuata, 447.

tiburo, 427.

tudes, 435, 436.
(Zygaena) tudes, 435.
zygaena, 420, 428, 435, 447, 449.
(Sphyrna) zygaena, 449.
Cestracyon zygaena, 449.
Cestrorhinus, 408.

tiburo, 427.
cetaceous, Squalus, 1 60.
Cethorhinus, 146, 159.
maximus, 146.
(Selache) maximus, 160.
Cetorhinidae, 96, 146.
Cetorhinus, 146, 189.
blainvillii, 160.
gunneri, 146 note 3, 157.
homianus, 157.
maccoyi, 147.
maximus, 70 note 17, 146, 147, 158, 160, 189,

194.
(Selache) maximus, 160.
peregrinus, 157.
rostratus, 160.
shavianus, I 57.
Chaetodipterus, 1 04..
chiereghini, Sphyrna, 435.
chilensis. Alopecias, 177.

Galeus, 264 note 4.
Chiloscyllium, 178, 180.
colcloughi, 180 note 5.
modestum, 180 note 5.
Chlamydoselachoidea, 77, 93.
Chlamydoselachus, 75, 76, 77, 94.

anguineus, 94.
Chondrichthyes, 43, 62.
Chordata, I.

ciliaris, Monopterhinus, 93.
cinereus, Carcharias, 92.
Heptanchus, 93.

Heptanchus (Heptranchias), 93.
Heptranchias, 87, 92.
Heptranchus, 93.
Hexanchus, 93.
Monopterhinus, 92.
Notidanus, 92.

Notidanus (Heptanchus), 93.
var. pristiurus (var. aetatis), Notidanus (Heptan-
chus), 93.
Squalus, 92.

Squalus (Monopterhinus), 92.
Squalus (Notidanus), 93.
circifrons, Myxine, 33.

Index of Scientific Names

cirratum, Ginglymostoma, 72, 181, 186.
Gingylostcma, 187.
Nebrius, 187.
Scyllium, 186.
cirratus, Squalus, I 80, I 8;.
cirrhatum, Ginglymostoma, 187.
cirrhatus, Squalus, 186.
Cirrhigaleus, 45 I.
Cirrhoscyilium, 179.
cirrhosum, Ginglymostoma, 186.

Scyllium, 180, 186.
Cirriscyllium, 179 note 3, 180 note 5.
cirrotum, Ginglymostoma, 187.
Citharichthys, 387.
Cladocera, 4.
Cladodonti, 94 note 2.
Cladodus, 94 note 2.
clavata, Raja, 65 note 2.
coeca, Myxina, 42.
coecus, Gasteobranchus, 42.

Gastrobranchus, 32, 42.
coelolepis, Centrophorus, 499.

Centroscymnus, 485, 493, 494, 498.
coerulea, Eulamia, 376.
coeruleus, Carcharias, 377.
Carcharias (Eulamia), 376.
Squalus, 376.
colcloughi, Chiloscyllium, 180 note 5.
commersonii, Carcharhinus, 320, 342, 343 note 34,
344. 345. 346, 353. 363. 388 note 121, 390.
Carcharias, 345, 363.
Carcharinus, 344, 362.
Eulamia, 345, 363.
concolor, Ammocoetes, 57.

Petromyzon, 57.
Conoporoderma, 197, 199.

pantherinum, 200.
cookei, Echinorhinus, 527, 532.
corinus, Hexanchus, 79, 85, 86.
cornubica. Lamia, 122.
Lamna, 120, 123.
Lamna (Oxyrhina), 122.
cornubicus, Carcharhinus, 120, 320 note la.
Isurus, 122.

Lamna, 131, 320 note la.
Squalus, 1 1 1, i 20.
Squalus (Carcharias), 122.
Squalus (Lamna), I 22.
cornubiensis, Squalus, 120.
corona, Sphyrna, 409 note 4, 414.
Coryphaena, 84.
crenidens, Carcharias (Scoliodon), 292.

crepidator, Centrophorus, 494 note I.
cryptacanthus, Centroscymnus, 494, 499
cubensls, Squalus, 453, 454, 455, 456,
472. 473. 478, 479. 480, 482, 490,
cucuri, Prionodon, 352.
cuvier, Galeocerdo, 72, 266, 274, 275.

Squalus, 265, 272, 274.
Cyclopterus, 520.
Cyclospondyli, 76.
Cyclostomata, 30.
Cynais, 240.

canis, 253, 256.
Cynias, 240.

canis, 253, 254.
Cynocephalus, 280, 292.
Cynoscion, 373.

nebulosus, 104.

regalis, 1 04.

Uaekayi, Oxyrhina, 122, 131.

dakini, Heptranchias, 88, 92.

Dalatias, 65 note 4, 495, 500, 508, 510,

(Somniosus), 514.

americanus, 501, 508.

(Somniosus) boreal is, 525.

brasiliensis, 513.

brevipinnis, 501, 508.

licha, 501, 502, 507, 508.

microcephalus, 525.

nocturnus, 500 note 3.

phillippsi, 501 note 5, 508.

sparophagus, 500, 506.
Dalatiidae, 450,499, 501, 508, 514.
Dalatius, 501.
deani, Heptranchias, 93.
Deania, 451, 487 note I.
Deaniops, 4; i note 5.
dekayi, Isuropsis, 122, 131.

Isurus, 122, 132, 133.

Isurus (Isuropsis), 132.
Didymodus, 94 note 2.
diplana, Sphyrna, 72, 97, 410, 411, 412,
415, 420, 427, 428, 431, 432, 433,

436, 439. 440, 441, 442. 443. 444.

447, 448 note 48.
Dipnoi, 62.

Dirrhizodon elongatus, 263 note 2.
ditropis, Lamna, 1 1 1, 112.
Dolichorhynchus, 8.
indicus, 7 note 16.
dombei, Eptatretus, 32 note 5.

559

note 7.
464, 469,
.494-

514. 5'6.

413.414,
434,435.
445, 446,

56o

Index of Scientific Names

dorsalis, Mustelus, 234, 240, 243.
dorsatus, Pctromyzon, 57.

Petromyzon marinus, 46, 57.

Pctromyzon marinus var. 57.
draco, Trachinus, 461.
dubia, Notidanus sp., 86.
duhamclii, Scyliorhinus, 203 note 28.
dumcril, Rhina, 543.

Squatina, 535, 537, 538, 543, 545.
dumcrili, Rhina, 543.

Squatina, 543.
dumerilii, Rhina, 543.

Scoliodon, 293.

Squatina, 543.

JiASTMANi, Galeus, 215.
Echeneis, 1 04.

remora, 273.
echinatum, Leidon, 524.

Somniosus, 514.
Echinorhinidae, 450, 526.
Echinorhinus, 59, 61, 510, 516, 526, 531

brucus, 60, 513, 514, 527, 53 1> 532-

cooliei, 527, 532.

mccoyi, 527.

(Rubusqualus) mccoyi, 532.

obesus, 527, 531.

spinosus, 526, 530, 531.
Echinorrhinus, 530.
edulis, Hexanchus, 79.

Mustelus, 242 note 4, 260.
edwardsii, Haploblepharus, 200.

Scyllium, 199 note 23.
Elasmobranchii, 63.
elephas, Selache, 158.

Squalus, I 58.
elongatum, Branchiostoma, 10.
elongatus, Dirrhizodon, 263 note 2.
Entosphenus, 44.
Entoxychirus, 45 I note 8.
Epigonichthyidae, 7, 18, 24.
Epigonichthys, 18, 24, 27.

caudatus, 23.

leucayanum, 22.
Eptatretus, 32.

dombei, 32 note 5.

stouti, 31.
Eridacnis, 234.

Etmopterus, 450, 451 note 4, 487, 494, 502, 5i(
aculeatus, 487.
brachyurus, 488 note 3.
frontimaculatus, 488 note 3, 492 note 9.

granulosus, 488 note 3.

hillianus, 482, 488, 493.

lucifcr, 488 note 3, 492.

molleri, 488 note 3.

paessleri, 488 note 3.

princcps, 488 note 3.

pusillus, 488, 490, 492 note 9, 493.

spinax, 488, 490, 492, 493.

villosus, 488 note 3.
Euchordata, I.
Eucrossorhinus, 179.
Eudontomyzon, 44.
Eugomphodus, 98.

griseus, 108.

littoralis, 108.
Eulamia, 320.

acronotus, 329.

aethlorus, 353.

cerdale, 399.

coerulea, 376.

commersonii, 345, 363.

falciformis, 333.

floridanus, 337.

glaucus, 291.

lamia, 320 note 3, 360, 361, 362, 363, 364.

lamiella, 364.

leucas, 345.

limbata, 352.

longimana, 362.

longimanus, 344, 363.

maculipinnis, 368.

melanoptera, 345.

milberti, 376, 377.

nicaraguensis, 382.

obscura, 390, 391.

obscurus, 391.

obtusa, 344.

oxyrhinchus, 394 note 139.

oxyrhynchus, 394.

platyodon, 343 note 31, 345, 346, 368.

plumbeus, 377.

porosus, 399.

remota, 403.

springer!, 407.
eumeces, Carcharias, 302.

Scoliodon, 303.
Euprotomicrus, 450 note 2, 500.
Euselachii, 76.
Eusphyra, 407.

blochii, 407 note I, 444.
Exoles, III.

Index of Scientific Names

561

rUnER, Zeus, 1 17.

fabricii, Centroscyllium, 482, 486, 487.

Spinax, 480, 486.
falciformis, Carcharhinus, 323, 329, 333, 334,

335.336,337.384-

Carcharhinus (Platypodon), 333.

Carcharias, 333.

Prionodon, 333.

Eulamia, 333.

Pla-typodon, 333.

Prionodon, 333.
falcipinnis, Carcharias, 389.
fasciatus, Carcharias (Prionodon), 275.

Galeocerdo, 275.

Galeorhinus, 259.

Mustelus, 242, 243, 245, 256, 259, 260.

Prionodon, 265.
Felichthys, 44 1,
fernandezianus, Acanthias, 480.

Spinax, 480.

Squalus, 480.
fernandinus, Acanthorhinus, 480.

Squalus, 453, 454, 455, 456, 472, 473, 478,
480.

Squalus acanthias, var., 480.
ferox, Carcharias, 98, 99, 1 08.

Galeorhinus, 98.

Isistius, 513.

Leius, 508, 5 13.

Notidanus, 87 note 2.

Squalus, 98, 108.
ferrugineum, Ginglymostoma, 181.
Figaro, 196, 214.

boardmani, 214.
fissidens, Carcharias, 398 note 144, 399.
fitzroyensis, Galeolamna (Uranganops), 321.
Flakeus, 452.

floridae, Branchiostoma, 9, 10 note 20, 16.
floridanus, Carcharhinus, 323, 329, 332, 333, 337,
384.

Eulamia, 337.
fluviatilis, Ammocoetes, 57.

Lampetra, 45, 56.

Petromyzon, 57.
foliaceus, Centrophorus, 451 note 8.
Foraminifera, 57.
frontimaculatus, Etmopterus, 488 note 3, 492

note 9.
fronto, Carcharias, 304 note 3.

Negaprion, 309.
fulgens, Squalus, 513.

Squalus (Scymnus), 508, 513.

fulvum, Ginglymostoma, 187.

Fur, 234.

fuscus, Centroscymnus, 494.

VJADus mcrlangus, 1 1 7.

galapagocnsis, Carcharhinus, 405.

Galea, 76.

Galeocerdo, 69, 74, 263, 265, 320 note la, 363.

arcticus, 272, 274, 275.

cuvier, 72, 266, 274, 275.

fasciatus, 275.

hemprichii, 275.

maculatus, 274.

obtusus, 275.

rayneri, 266 note 6, 274, 275.

tigrinus, 273, 274.
Galeoidea, 76, 77, 95.
Galeolamna, 320, 321.

(Bogimba) bogimba, 321.

(Uranganops) fitzroyensis, 321.

greyi, 320.
Galeolamnoides, 321.
Galeorhinidae, 233.
Galeorhinus, 72, 98, 233, 240, 254 note 32, 264.

fasciatus, 259.

ferox, 98.

galeus, 70 note 17, 72, 73, 264 note 4.

laevis, 253.

zyopterus, 264 note 4.
Galeus, 96, 161, 196, 199, 202, 214, 240, 265,
272, 280.

arae, 210, 211, 215, 216.

boardmani, 21 5.

caniculus, 202.

canis, 253, 254, 264 note 4.

cepedianus, 265, 274.

chilensis, 264 note 4.

eastmani, 21 5.

glaucus, 280, 291.

hertwigi, 2 1 5 note 4.

jenseni, 215.

maculatus, 272.

melastomus, 214, 215.

mento, 261.

molinae, 264 note 4.

murinus, 215.

sauteri, 2 1 5.

tigrinus, 274.

uyato, 2 1 4 note i .
galeus, Galeorhinus, 70 note 17, 72, 73, 264 note 4.

Squalus, 240.
garmani, Myxine, 33.

562

Index of Scientific Names

Gasteobranchus, 32.

coecus, 42.
Gasterobranchus, 8.
Gasterosteus, 248.
Gastrobranchus, 32.

caecus, 42.

coecus, 32, 42.
Gillisqualus, 321.
Ginglimostoma, 1 80.
Ginglymostoma, 74, 178, 179, 180, 187 note 18,

313-

brevicaudatum, 18 1.

caboverdianus, 187.

cirratum, 72, 181, 186.

cirrhatum, 187.

cirrhosum, 186.

cirrotum, 187.

ferrugineum, 181.

fulvum, 187.
Gingylostoma, 180.

cirratum, I 87.
glacialis, Scimnus, 525.

Scymnus, 524.

Squalus, 524.
gladius, Xiphias, I 29.
glauca, Lamna, 131.

Oxyrhina, 131.

Oxyrrhina, 131.

Prionace, 69, 70 note 17, 281, 282, 290, 291,
388, 389,390.
glaucus, Carcharhinus, 320 note la.

Carcharias, 289, 292 note 29.

Carcharias (Prionodon), 290.

Carcharinus, 290.

Eulamia, 291.

Galeus, 280, 291.

Glyphis, 291.

Isurus, 69, 123, 124, 128, 129, 131, 132, 133
note 43.

Oxyrhina, 123.

Prionace, 320 note I a.

Prionodon, 290.

Squalus, 119, 120, 280, 289, 377 note 104.

Squalus (Carcharhinus), 290.

Squalus (Carcharias), 290.
glutinosa, Myxine, 32, 33, 34, 40, 42.
Glyphis, 280, 321.

glaucus, 291.

hastalis, 321.

mackei, 291.
glyphis, Carcharias (Prionodon), 280 note 5, 321
note 4.

gomphodon, Oxyrhina, 123, 131.
Goniadus, 531 note 12.
Goniodus, 526.

spinosus, 531.
gracilis, Carcharias, 292.
granulosum, Centroscyllium, 482, 487.
granulosus, Etinopterus, 488 note 3.
gravelyi, Branchiostoma, 10 note 19.
greyi, Alopias, 162, 177 note 33, 178.

Galeolamna, 320.
griffini, Squalus, 452, 454.
griseus, Carcharias, 98, 108.

Eugomphodus, 108.

Hexanchus, 78, 79, 80, 85, 87, 90.

Monopterhinus, 86.

Notidanus, 86.

Notidanus (Hexanchus), 86.

Odontaspis, 1 08.

Squalus, 78, 79, 85.

Squalus (Monopterhinus), 86.
gruveli, Paragaleus, 275, 276.
gunneri, Cetorhinus, 146 note 3, 157.

Scymnus, 524.

Squalus (Scymnus), 524.
gunnerianus, Squalus, 146, 157.
guntheri, Isurus, 124.
Gymnorhinus, 321.

tiAECKELi, Branchiostoma, 9.

Scyliorhinus, 205.
haeckelii, Catulus, 207.

Scyliorhinus, 59.
Halaeluridae, 195 note I.
Halaelurus, 198, 202.

rudis, 202.
Halsydrus, 146.

maximus, 160.

pontoppidani, 146, 160.
Haploblepharus, 199, 201.

edwardsli, 200.
harbereri, Proscyllium, 197 note 12.
Harriota, 63 note 5.
hastalis, Glyphis, 321.
haswelli, Hepiranchias, 87 note 2.
Hemigaleus, 264 note 3, 275, 280.

pectoralis, 275, 280.
hemiodon, Carcharias (Hypoprion), 315.

Hypoprion, 316.
Hcmlpristis, 263.
Hemiscyllium, 178, 180.
Hemitriakis, 234 note 5, 235.

Index of Scientific Names

563

leucoperiptera, 235.
hemprichii, Galcocerdo, 275.
henlei, Carcharhinus, 399.

Carcharhinus (Platypodon), 399.

Carcharias, 399.

Carcharias (Prionodon), 399.

Rhinotriacis, 235.

Trlakis, 235 note 7, 236.
Heptanchus, 87.

cinereus, 93.

(Heptranchias) cinereus, 93.
Heptatremus, 32 note 5.
Heptranchias, 78, 87, 94.

cinereus, 87, 92.

dakini, 88, 92.

deani, 93.

haswelli, 87 note 2.

perlo, 87 note 2, 88, 93.
Heptranchus, 87.

cinereus, 93.
Heptrancus, 87.

angio, 93.
herklotsi, Apristurus, 220, 221 note 10.
hertwigi, Galeus, 215 note 4.
heterobranchialis, Carcharliinus, 320 note la.
heterodon, Carcharhinus, 320 note la.
Heterodontea, 76.
Heterodontidae, 65, 67, 95.
Heterodontoidea, 76, 77, 95.
Heterodontus, 95.

heterodus, Hypoprion? (Hemigaleus? ), 316 note 4.
Heteropleuron (Asymmetron) lucayanum, 23.
Heteroscyllium, 1 80.
Heteroscymnoides, 500.
Heteroscymnus, 500.
Hexanchea, 76.
Hexanchidae, 75, 78.
Hexanchus, 78, 87, 90, 94.

cinereus, 93.

corinus, 79, 85, 86.

edulis, 79.

griseus, 78, 79, 80, 85, 87, 90.
Hexancus, 79.
hillianus, Etmopterus, 482, 488, 493.

Spinax, 493.
hinnulus, Mustelus, 253.

Squalus (Galeorhinus), 253.
Hippoglossus, 520.

hirundinaceus, Carcharias (Prionodon), 290.
Holocephali, 63, 76.
Holohalaelurus, 198, 201.
Homea, 32 note 5.

homianus, Cetorhinus, 157.

Squalus, 1 57.
Hybodus, 75.
Hypoprion, 265, 293 note 2, 308, 315, 322.

brevirostris, 314, 315 note 2.

hemiodon, 316.

longirostris, 319.

macloti, 316.

palasorrah, 302.

playfairii, 316.

signatus, 316, 3 19.
Hypoprion? (Hemigaleus?) heterodus, 316 note 4.

(Hemigaleus?) isodus, 316 note 4.
Hypoprionodon, 315.

J CHTHY0MY70N, 44, 52 note 83, 54 Hote 98a, 57.
indicus, Apristurus, 221.

Dolichorhynchus, 7 note 16.

Scylliorhinus, 219.
Infusoria, 4.

insularum, Carcharias, 361.
intermedius, Scoliodon, 294.
Isistius, 65 note 4, 500, 502, 508, 516.

brasiliensis, 485, 509, 513, 514 note 9.

ferox, 513.
isodon, Aprionodon, 304, 307.

Carcharhinus (Aprionodon), 308.

Carcharias (Aprion), 303, 307.

Carcharias (Aprionodon), 308.
isodus, Hypoprion? (Hemigaleus?), 316 note 4.

Squalus, 158.
Isogomphodon, 321.

limbatus, 352.

maculipinnis, 352, 368.

oxyrhynchus, 394.
Isoplagiodon, 321.
Isurida, 76.

Isuridae, 65, 66, 96, 109, 126, 146, 150.
Isuropsis, 123, 131.

dekayi, 122, 131.
Isurus, 39, no, 123, 132, 134, 136, 137, 151,
256.

bideni, 123, 124 note 30.

cepedii, 132.

cornubicus, 122.

dekayi, 122, 132, 133.

(Isuropsis) dekayi, 132.

glaucus, 69, 123, 124, 128, 129, 131, 132, 133
note 43.

guntheri, 124.

mako, 123, 124 note 30.

564

Index of Scientific Names

nasus, 122, 271, 272.

(Lamna) nasus, 123.

oxjrhynchus, 132, 133.

(Isurus) oxyrhynchus, 132.

oxyrinchus, 69, 1 13, 1 14, 1 1 5, 116, 122, 123,

124, 130, 132.
oxyrynchus, 131.
punctatus, i 22.
spallanzanii, I 31.
tigrinus, 265, 274.
tigris, 132, 133.

J APONicA, Squatina, 535, 536 note 19, 537.
japonicus, Squalus, 454 note 20, 480.
jenseni, Galcus, 215.
jordani, Scoliodon, 293.
Juncrus, 198.

l^iRKi, Squalus, 453 note 15.
Koinga, 452.

I-/ACTOPHRYS, 313.
Laemargus, 514, 526.

borealis, 524.

brevipinna, 526.

carcharias, 524.

microccphalus, 525.
laevi, Squatina, 543.
laevis, Galeorhinus, 253.

Mustelus, 240, 253, 254.

Squalus (Galeorhinus), 253.
Lagodon, 373.
lalandii, Carcharias (Scoliodon), 301.

Scoliodon, 294 note 3, 297 note 5, 299, 301.
Lamia, 1 1 1.

cornubica, 122.

monensis, 122.

nasus, 1 18 note l6, 123, 130 note 37, 133.

oxyrhincus, 131.
lamia, Carcharhinus, 272, 320 note la, 344, 363,
364.

Carcharhinus (Carcharhinus), 362.

Carcharhinus (Eulamia), 362.

Carcharias, 142, 344, 346 note 40, 354 note 51,
359 note 58, 362, 363, 364, 403.

Carcharias (Prionodon), 143, 344, 354 note 51,
359, 362, 363,364.

Carcharinus, 345, 362, 363.

Carcharodon, 142, 320 note la.

Eulamia, 320 note 3, 360, 361, 362, 363, 364.

Prionodon, 362, 363, 364.
lamiella, Carcharhinus, 399.

Carcharias, 399.

Eulamia, 364.
Lamiopsis, 321.

limbatus, 352.
Lamna, 1 10, III, 123, 132, 134, 292, 320 note la.

caudata, 301 note 17, 375.

cornubica, 120, 123.

(Oxyrhina) cornubica, 122.

cornubicus, 131, 320 note I a.

ditropis, III, 1 1 2.

glauca, 131.

latro, 131.

monensis, 122, 320 note I a.

nasus, 70 note 17, 1 1 1, 112, 122, 123, 126, 127,
128, 129 note 36, 130, 132, 133 notes 44,
45, 272, 292, 303.

oxyrhina, 123, 131.

oxyrhynchus, 132.

pennanti, 122.

philippii, III.

punctata, 122, 123, 131, 292, 303.

spallanzanii, 131.

terrae-novae, 301.

tigris, 132.

whitleyi, 1 1 1 note 5.
Lamnarius, 321.
Lamniformes, 76.
lamotteni, Petromyzon, 58.
Lampetra, 44, 58.

fluviatilus, 45, 56.

marinus, 57.
lampetra, Petromyzon, 58.
lanceolata, Branchiostoma, 16.
lanceolatum, Branchiostoma, 4, 5, 9, 10, 11, 13, 14,

15, 16, 24.
lanceolatus, Amphioxus, 16.

Limax, 8, 16.
"Larvae," Amphioxides, 7, 23, 24, 25, 27.
laticaudus, Carcharias (Scoliodon), 292.
latro, Lamna, 131.
laurussonii, Apristurus, 221 note 11, 224, 226.

Scyllium, 219, 224, 225.
lebruni, Acanthias, 353 note 14.
leeuwenii, Zygaena, 425 note 19, 428.
Leiodon, 514.

echinatum, 524.
Leiostomus, 104.
Leius, 508.

brasiliensis, 513.

ferox, 508, 513.

Index of Scientific Names

565

Lepidorhinus, 451 note 8.
Leptocardii, I.
Leptoch.irias, 234.

leucas, Carcharhinus, 72, 74, 320 note la, 324, 337,
344, 345, 346, 353, 354. 359. 361, 362, 363.
364, 378, 380, 381, 382, 386, 387, 388, 390.

Carcharias, 144, 346, 363.

Carcharias (Prionodon), 344.

Eulamia, 345.
leucayanum, Epigonichthys, 22.
leucoperiptera, Hemitriakis, 235.

Triakis, 236.
leucos, Carcharias (Prionodon), 344, 345.
leucos?, Carcharias, 345.
lewini, Sphyrna, 408, 419, 449.

Zygaena, 428, 449.
liburo, Squalus, 426 note 24.
licha, Dalatias, 501, 502, 507, 508.

Scytnnorhinus, 508.

Scymnus, 507.

Squalus, 500, 501, 506.
lichi (Scymnus lichia), Scymnorhinus, 508.
lichia, Scymnorhinus, 507.

Scymnus, 501, 507, 508.

Scymnus (Scymnus), 507.

Squalus (Scymnus), 508.
Limax, 8.

lanceolatus, 8, 16.
limbata, Eulamia, 352.

limbatus, Carcharhinus, 72, 270, 325, 345, 346,
352. 353> 364. 366, 367, 368, 388, 400,
403.

Carcharhinus (Isogomphodon), 352.

Carcharias, 353, 368.

Carcharias (Prionodon), 352.

Isogomphodon, 352.

Lamiopsis, 352.

Prionodon, 353.
limosa, Myxine, 38, 42.
Limulus, 270.
linnei, Acanthias, 472.
littoralis, Carcharias, 99 note 15, 107.

Eugomphodus, 108.

Odontaspis, 108.

Squalus, 107.
lividus, Carcharhinus, 320 note la.
longimana. Alopecias, 177.

Eulamia, 362.
longimanus, Carcharhinus, 320 note la, 323, 324,
337. 338. 340, 341. 342. 343. 344. 345. 353,
354, 363 note 71, 384, ?86, 387.

Carcharias, 346 note 40.

Eulamia, 344, 363.

Prionodon, 362.

Squalus, 362.
longirostrls, Hypoprion, 319.
longmani, Scoliodon, 294.
Longmania, 303, 321.

brevipinna, 265 note 5.

calamaria, 321.
longurio, Carcharias, 303.

Carcharias (Scoliodon), 303.

Scoliodon, 294, 300.
Lophius, 542 note 26.
Lota maculosa, 53.
Loxodon, 263.

lubricum, Branchiostoma, 8, 13, 16.
lucayanum, Asymmetron, 18, 19, 22, 25, 26 note
58.

Branchiostoma, 22.

Heteropleuron (Asymmetron), 23.
lucifer, Etmopterus, 488 note 3, 492.
Lunatia, 270, 463.
lunulatus, Mustelus, 241.

JMaccoyi, Cetorhinus, 147.
macer, Polyprosopus, 146, 160.
mackei, Carcharhinus, 291.

Glyphis, 291.

Prionace, 280, 281, 291.
macleayi, Atelomycterus, 199 note 23.
macloti, Carcharias (Hypoprion), 315.

Hypoprion, 316.
macracanthus, Centroscymnus, 494 note I.
macricaudatum, Asymmetron, 19, 21 note 40, 22.
macrodous, Squalus, 1 07.
macrorhynchus, Apristurus, 221 note 8.

Carcharias (Scoliodon), 292.
macrourus, Alopias, 161, 174.
macrurus, Carcharias, 320 note 2, 321.
maculata, Triakis, 234, 235 note 7, 236.
maculatus, Galeocerdo, 274.

Galeus, 272.

Scomberomorus, 441.
maculipinna, Carcharias, 352.

maculipinnis, Carcharhinus, 70, 325, 346, 348,
349,351,364,400.

Carcharias, 368.

Eulamia, 368.

Isogomphodon, 352, 368.

Platypodon, 368.
maculosa. Lota, 53.
maculosus, Petromyzon, 57.
major vulgaris, Catulus, 202.

566

Index of Scientific Names

Makaira, 84.

mako, Isurus, 123, 1 24 note 30.

malleus, Sphyrna, 435, 447.

Squalus, 420, 444.

Zygaena, 418 note 10, 419, 420, 446, 448.
Mallotus, 520.

maou, Squalus (Carcharias), 1 45.
Mapolamia, 32 1.

marina, Vulpecula, 161, 176, 177.
marinus, Lampetra, 57.

Petromyzon, 45, 46, 54, 56, 57, 58.

(amcricanus), Petromyzon, 57.

dorsatus, Petromyzon, 57.

var. dorsatus, Petromyzon, 57.

unicolor, Petromyzon, 57.
marleyi, Poroderma, 197.
marmoratum, Scyllium, 199 note 23.
maso, Carcharias, 1 45.
maxima, Selache, 146, 157, 189, 194.
maximum, Selache, 157 note 39.
maximus, Cethorhinus, 146.

Cethorhinus (Selache), 160.

Cetorhinus, 70 note 17, 146, 147, 158, 160,
189, 194.

Cetorhinus (Selache), 160.

Halsydrus, 160.

Selache, 157 note 39.

Selachus, 146, I 58.

Selanche, 146, 158.

Squalus, 146, I 56.

Squalus (Selache), 158.
mccoyi, Echinorhinus, 527.

Echinorhinus (Rubusqualus), 532.
media, Sphyrna, 409 note 4, 414.
medinae, Notidanus, 87 note 2.
megalops, Carcharinus, 320 note la.

Squalus, 452, 454, 480 note 66.
melanoptera, Eulamia, 345.
melanopterus, Carcharias, 32 I, 345, 353.

Carcharias (Prionodon), 353.
melanostomum, Pristiurus, 214.
melastomus, Galeus, 214, 215.
menisorrah, Carcharias (Prionodon), 321, 399.

Carcharinus, 399.
mento, Galeus, 261.

Mustelus, 242, 243, 244 fig. 42, 245, 251, 256,
259, 260, 261.
merlangus, Gadus, 117.
Merluccius, 84, 91, 1 04, 461 note 33, 462.
Merscalpellus, 18 note 36.
Micristodus, 188.

punctatus, 188, 194.

microcephalus, Dalatias, 525.

Laemargus, 525.

Scymnus, 525.

Somniosus, 516, 525.

Squalus, 515, 523.
microJon, Pseudotriakis, 60, 228, 229, 232.
Micropogon, 462.
microps, Apristurus, 220, 221.

Carcharias, 353.
micropterus, Scymnus, 524.

milberti, Carcharhinus, 70 note 17, 72, 139, 301
note 17, 324, 342, 343, 345, 346, 368, 377,
378, 381, 382, 384, 385, 386, 387, 388,
389, 390.

Carcharhinus (Carcharhinus), 377.

Carcharias, 376, 378.

Carcharias (Prionodon), 320, 321 note 3, 375,
378,382.

Carcharinus, 363, 376, 377, 382.

Eulamia, 376, 377.

Prionace, 378.

Prionodon, 378.

Squalus, 376, 377, 378.

Squ.ilus (Carcharinus), 376.
minucauda, Amphipleurichthys, 8.

Branchiostoma, 9.
mitsukurii, Squalus, 454 note 20, 480.

Squalus sucklii, var., 473.
Mobula, 342.

modestum, Chiloscyllium, 180 note 5.
molinae, Galeus, 264 note 4.
molleri, Etmopterus, 488 note 3.
Molva, 39, 520, 521.
monensis, Carcharhinus, 320 note la.

Lamia, 122.

Lamna, 122, 320 note la.

Squalus, I 20.
monge, Notidanus, 86.
Monopterhinus, 78, 87.

ciliaris, 93.

cinereus, 92.

griseus, 86.
montalbani, Squalus, 480.
Mordaciidae, 43.
Mugil, 104.
miilleri, Amphioxus, 17.

Carcharias (Prionodon), 352.
Mullus, 387.
Muraenoblenna, 32.

olivacea, 32.
murinus, Galeus, 215.
Mustek, 240 note I.

Index of Scientific Names

567

Mustelidae, 233.
Musteliformcs, 76.

Mustelus, 66, 68, 233, 234, 236 note 10, 240, 254
note 32, 256, 259, 260.
abbotti, 242 note 4, 260 note 38, 261.
asterias, 242, 254, 260, 261.
californicus, 240, 242.
canis, 68, 97, 243, 244, 251, 254, 255, 256,

257 fig. 43, 258, 259, 260, 261, 262.
dorsalis, 234, 240, 243.
edulis, 242 note 4, 260.
fasciatiis, 242, 243, 245, 256, 259, 260.
hinnulus, 253.
laevis, 240, 253, 254.
lunulatus, 241.
mento, 242, 243, 244 fig. 42, 245, 251, 256,

259, 260, 261.
mustelus, 243, 244, 247, 253, 254, 255.
norrisi, 241, 243, 244, 251 notes 21, 23, 253
note 30, 254, 256, 257 fig. 43, 258, 260.
punctulatus, 242.
schmidti, 262.
schmitti, 242, 243, 244, 251, 254, 256, 257

fig. 43, 260, 261, 262.
striatus, 259.
vulgaris, 254.
mustelus, Mustelus, 243, 244, 247, 253, 254, 255.

Squalus, 240.
Myoxocephalus, 248, 520.
Myrmillo, 240.
Myxina coeca, 42.
My.xine, 8, 31, 32, 39, 40, 41, 43.
affinis, 33, 38, 42.
atlantica, 38, 43.
australis, 33 note 8, 38, 42,
capensis, 33 note 8, 38.
circifrons, 33.
garmani, 33.

glutinosa, 32, 33, 34, 40, 42.
limosa, 38, 42.
paucidens, 33.
tridentiger, 33.
myxine, Petromyzon, 42.
Myxinidae, 31.
Myxinoidea, 30, 31.

N ARCAciON, 444.

nasus, Isurus, 122, 271, 272.

Isurus (Lamna), 123.

Lamia, 1 18 note 16, 123, 130 note 37, 133.

Lamna, 70 note 17, ill, 112, 122, 123, 126,

127, 128, 129 note 36, 130, 132, i33notes
44,45,272,292, 303.
Squalus, III, 120.
nasuta, Uranga, 3 2 1.
natator, Carcharhinus, 351 note 45, 353.

Carcharias, 353.
Nebria, 179 note 4.
Nebris, 1 79 note 4.
Nebrius, 179.

cirratum, 187.
Nebrodes, 178, 179 note 4.
nebulosa, Squatina, 536.
nebulosus, Cynoscion, 104.

Negaprion, 74, 264, 304 notes 2, 3, 308, 315
note 2.
acutidens, 309.
brevirostris, 69, 72, 309, 310, 337, 338, 388,

390.
fronto, 309.
odontaspis, 309.
queenslandicus, 309.
Negogaleus, 264, 275, 276.
Nereis, 248.
nicaense, Squalus, 501.
nicaensis, Scymnus, 507.

Squalus, 506.
nicaraguensis, Carcharhinus, 324, 337, 339, 341,
342,378,382.
Carcharhinus (Carcharhinus), 382.
Carcharias, 382.
Eulamia, 382.
niciense, Scymnium, 507.
nigra, Centrina, 487.
nigricans, Petromyzon, 46 note 55, 56.
nigromaculata, Triakis, 236 note 10.
nigrum, Centroscyllium, 481.
nocturnus, Dalatias, 500 note 3.
norrisi, Mustelus, 241, 243, 244, 251 notes 21, 23,
253 note 30, 254, 256, 257 fig. 43, 258,
260.
norvegianus, Squalus, 524.
norvegicus, Squalus, 525.
norwegianus, Acanthorhinus, 523.

Somniosus, 514.
Notasymmetron, 18 note 35, 19 notes 38 and 39,
23.
caudatum, 23.
Notidamus, 79.
Notidanoidea, 77.
Notidanus, 79, 87.
cinereus, 92.
(Heptanchus) cinereus, 93.

568

Index of Scientific Names

(Heptanchus) cinereus, var. pristiurus (var. aeta-

tis), 93.
ferox, 87 note 2.
griseus, 86.

(Hexanchus) griseus, 86.
medinae, 87 note 2.
monge, 86.
sp. dubia, 86.
vacca, 86.
vulgaris, 87.
wolniczkyi, 87 note 2.
Notorynchus, 78, 87.

Obesus, Echinorhinus, 527, 531.

Triaenodon, 309 note 2.
obliqua, Oxytes, 98 note 1 2.
obscura, Eulamia, 390, 391.

obscurus, Carcharhinus, 70 note 17, 72, 323, 342,
343, 344. 345- 346, 361, 372 note 85, 374.
377, 382, 389, 390, 391, 404. 405. 406, 407-

Carcharhinus (Eulamia), 389.

Carcharhinus (Platyodon), 390.

Carcharhinus (Platypodon), 390.

Carcharias, 292, 377, 389, 390.

Carcharias (Prionodon), 389, 391.

Carcharinus, 389, 391.

Eulamia, 391.

Platypodon, 389.

Squalus, 292, 389.

Squalus (Carcharinus), 389.
obtusa, Eulamia, 344.
obtusirostris, Carcharias, 362.
obtusus, Carcharias (Prionodon), 362.

Galeocerda, 275.

Prionodon, 344.

Squalus, 344.
obvelatus, Carcharias (Prionodon), 389.

Prionodon, 389, 390.
Oceanomyzon, 45, 47 "°t^ 57-

wilsoni, 45, 46, 57.
Octopus, 373, 424.
oculata, Squatina, 536.
Odontaspis, 98.

americanus, 108, 109.

griseus, 108.

littoralis, 108.

platensis, 98.

taurus, 108.
odontaspis, Negaprion, 309.
Ogilamia, 32 1.
olivacea, Muraenoblenna, 32.
Ophidion, 387.

Orca, 172.

Orectolobidae, 96, 178, 188, 233.
Orectolobus, 1 78, 1 79.
orientale, Asymmetron, 19, 23.
ornatum, Centroscyllium, 481.

Paracentroscyllium, 481.
owstoni, Carcharias, 99.
Centroscymnus, 494.
Oxynotidae, 450 note 3.
Oxynotus, 450.
Oxyrhina, 123.
daekayi, 122, 131.
glauca, 131.
glaucus, J 23.
gomphodon, 123, 1 3 1.
punctata, 122.
spallanzanii, 1 3 1,
oxyrhina, Lamna, 123, 131.
oxyrhinchus, Eulamia, 394 note 139.
oxyrhincus. Lamia, 131.
oxyrhynchus, Carcharhinus, 323, 391.
Carcharhinus (Isogomphodon), 394.
Carcharias (Prionodon), 321, 394.
Carcharinus, 394.
Eulamia, 394.
Isogomphodon, 394.
Isurus, 132, 133.
Isurus (Isurus), 132.
Larana, 132.
oxyrinchus, Isurus, 69, 1 1 3, 1 1 4, 1 1 5, 116, 122,

123, 124, 130, 132.
Oxyrrhina, 123.
glauca, 131.
spalanzanii, 1 3 1.
oxyrynchus, Isurus, 131.
Oxytes, 98 note 12.
obliqua, 98 note 12.

Pacificus, Somniosus, 515, 516, 525.
paessleri, Etmopterus, 488 note 3.
palasorrah, Carcharias, 292, 293 note 2.

Hypoprion, 302.
Palinurus, 184, 270.
pantherinum, Conoporoderma, 200.

Poroderma, 197.
Paracentroscyllium, 481.

ornatum, 481.
Paradontaspis, 98.
Paragaleus, 264, 275, 280.

gruveli, 275, 276.

pectoralis, 276.
Paramphioxus, 18 note 36.

Index of Scientific Names

569

Paramyxine, 31, 32.
Parapristurus, 2ig, 220 note I.
Parascyllium, 179.
Parmaturus, 196, 199.
paucidcns, Myxine, 33.
pectoralis, Hemigaleus, 275, 280.

Paragaleus, 276.
pelagicum, Asymmetron, 26.

Branchiostoma, 23, 24, 26, 28.
pelagicus, Alopias, 161, 162.

Amphioxides, 22, 23, 24, 25, 26, 27, 28.
pelegrinus, Squalus, I 5 7.
pennanti, Lamna, 122.

Selachus, 160.

Squalus, 120.
pentalineatus, Rhinodon, 194.
Pentanchus, 197 note 11, 199, 219, 220 note I.

profundicolus, ig6.
peregrinus, Cetorhinus, 157.

Squalus, 157.
perezi, Carcharhinus (Platypodon), 352.
perezii, Platypodon, 352.
pcrlo, Heptranchias, 87 note 2, 88, 93.

Squalus, 87, 92.
Petromyzon, 32, 40, 44, 45, 57.

americanus, 56.

(Bathymyzon) bairdii, 57.

branchialis, 45, 56.

castaneous, 54 note g8a.

concolor, 57,

dorsatus, 57.

fluviatilis, 57.

lamotteni, 58.

lampetra, 58.

maculosus, 57.

marinus, 45, 46, 54, 56, 57, 58.

marinus (americanus), 57.

marinus dorsatus, 46, 57.

marinus var. dorsatus, 57.

marinus unicolor, 57.

myxine, 42.

nigricans, 46 note 55, ;6.

unicolor, 57.
Petromyzonida, 30, 43.
Petromyzon idae, 43.
Phaenopogon, 45 1 note 6.
philippi, Squatina, 537.
philippii, Lamna, iii.
philippinus, Squalus, 480.
phillippsi, Dalatias, 501 note 5, 508.

Scymnorhinus, 508.
Pholis, 32, 39 note 35, 40.

Physodon, 263.

platae, Branchiostoma, 9, 10, 11, 12, 14, 15, 16,

17, 18.
platensis, Carcharias, 99, 106.

Odontnspis, g8.
platyodon, Carcharhinus, 344, 345, 354 riote 52a,
377-

Carcharhinus (Carcharhinus), 344.

Carcharhinus (Eulamia), 344.

Carcharias, 344.

Carcharinus, 346, 363.

Eulamia, 343 note 31, 345, 346, 368.

Prionodon, 344.

Squalus, 344, 377.
Platypodon, 321.

acronotus, 328.

falciformis, 333.

maculipinnis, 368.

obscurus, 389.

perezii, 352.

tiburo, 333.
platyrhynchus, Apristurus, 220, 221 note 10.

Carcharhinus, 405.
Platysqualus, 408.

tiburo, 427.

tudes, 436.
playfairii, Hypoprion, 316.
Plectrosoma, 123.
Pleuracromylon, 240.
Pliotrema, 533.
plumbeus, Carcharhinus, 368 note 82, 378, 390.

Carcharias, 377.

Eulamia, 377.

Squalus, 377.
Pogonias, 104, 107.
Pollachius, 49.

vircns, 286, 520.
Polyprosopus, 146.

macer, 146, 160.

rashleighanus, 160.
Pomatomus, 104, 172.
Pomolobus, 104.

pontoppidani, Halsydrus, 146, 160.
Poroderma, ig7, 202.

africanum, ig7 note 16, 202.

marlcyi, ig7.

pantherinum, ig7.
Poronotus, 104.

triacanthus, 350.
porosus, Carcharhinus, 323, 394.

Carcharias, 303, 3gg.

Carcharias (Prionodon), 3g9.

570

Index of Scientific Names

Carcharinus, 399.

Eulamia, 399.

Scoliodon, 303.

Squalus, 303.
princeps, Etmopterus, 488 note 3.
Prionace, 263 note I, 264, 280, 286 note 14, 292,
320 note I a.

caeruleus, 320 note I a.

glauca, 69, 70 note 17, 281, 282, 290, 291, 388,
389, 390.

glaucus, 320 note la.

mackei, 280, 281, 291.

milberti, 378.
Prionodon, 265, 280, 320.

cucuri, 352.

falciformis, 333.

fasciatus, 265.

glaucus, 290.

lamia, 362, 363, 364.

limbatus, 353.

longimanus, 362.

milberti, 378.

obtusus, 344.

obvelatus, 389, 390.

platyodon, 344.

tiburo, 333.
Prionotus, 104.
Pristidae, 77 note 36, 532.
Pristidurus, 214.
Pristiophoridae, 77, 532.
Pristiophoroidea, 77, 532.
Pristiophorus, 533.
Pristiurus, 214, 219.

arae, 219.

melanostomum, 214.
pristiurus (var. aetatis), Notidanus (Heptanchus)

cinereus, var., 93.
profundicolus, Pentanchus, 196.
profundorum, Apristurus, 196 note 9, 220, 221,
222, 225, 226, 228.

Scylliorhinus, 225.
profundus, Alopias, 162, 167.
Proscyllium harbereri, 197 note 12.
Protozygaena, 265.
Pseudopleuronectes, 373.
Pseudoscymnus, 500 note I, 50 1.

boshuensis, 501, 508.
Pseudotriakidae, 96, 228.
Pseudotriakis, 59, 61, 96, 228.

acrages, 229, 232.

acrales, 229 note 3.

microdon, 60, 228, 22g, 232.

Pternodont.1, 94 note 2.

pugae, Carcharias, 290.

punctata, Lamna, 122, 123, 131, 292, 303.

Oxyrhina, 122.
punctatus, Aprionodon, 303, 307.

Carcharias (Aprionodon), 308.

Isurus, 122.

Micristodus, 188, 194.

Scylliorhinus, 198 note 19.

Squalus, 122, 186, 292, 303, 307, 308.
punctulatus, Mustelus, 242.

Squalus, I 86.
pusillum, Acanthidium, 487.
pusillus, Etmopterus, 488, 490, 492 note 9, 493.

Spinax, 493.

(ciuEENSLANDicus, Aprionodon acutidens, 308.
Negaprion, 309.

K-ADIOLARIA, 4.

Raja clavata, 65 note 4.
rashleighanus, Polyprosopus, 160.

Squalus, 158.
rayneri, Galeocerda, 266 note 6, 274, 275.
regalis, Cynoscion, 104.
regani, Scyliorhinus, 197 note 15.

Scylliorhinus, 198 note 19.
Reinhardtius, 520.
Remora, 193.
remora, Echeneis, 273.
remota, Eulamia, 403.

remotus, Carcharhinus, 324, 352 note 50, 364,
400, 403.

Carcharhinus (Platypodon), 403.

Carcharias, 403.

Carcharias (Prionodon), 403.

Carcharinus, 403.
Reniceps, 408, 428.

tiburo, 427.
retifer, Catulus, 211.

Scyliorhinus, 203, 205, 206, 207, 211, 212,
219.

Scylliorhinus, 207, 210, 211.

var. boa, Catulus, 207.
retiferum, Scyllium, 207, 210.
Rhamphobatis, 534.
Rhina, 534.

angelus, 546.

argentina, 546.

armata, 546.

dumeril, 543.

Index of Scientific Names

571

dumerili, 543.

dumerilii, 543.

squatina, 543, 544.
Rhincodon, 61, 67, 160, 188.

typus, 188, 189, 193.
Rhincodontidae, 96, 187.
Rhineodon, 188, 193.

typicus, 188, 194.

typiis, 194.
Rhinida, 76.

Rhiniodon, 188, 193 note 16.
riiinoceras, Squalus, I 5 8.
Rhinodon, i 88.

pentalineatus, 194.

typicus, 188, 193.

typus, 194.
Rhinoscymnus, 514.
Rhinotriacis, 235 note 5.

hcnlei, 235.
Rhinotriakis, 240.
Rhizoprion, 292.
Rhizoprionodon, 292.
Rineodon, 188.
rittcri, Centroscyllium, 482.
riveri, Apristurus, 196 note 9, 221, 222, 223,

224, 225, 228.
rondeletti, Carcharias, 142.
rondelettii, Carcharodon, 134 note 3, 142, 144,

145.
rossi, Centrophorus, 45 1 note 8.
rostrata, Selache, 1 60.
rostratus, Cetorhinus, 160.

Seym n us, 5 14.

Somniosus, 516, 520.

Squalus, 1 58, 515.
Rubusqualus, 526.
rudis, Halaelurus, 202.

OALDANHA, Apristurus, 221.

Sarda, 104, 373.

Sardlnops caerulea, 350.

sauteri, Galeus, 215.

Scapanorhynchidae, 97, 109.

Scapanorhynchus, 109.

schmidti, Mustelus, 262.

schmitti, Mustelus, 242, 243, 244, 251, 254, 256,

257 fig. 43, 260, 261, 262.
Sciaena aquilla, 524.
Scimnus, 5 14.

borealis, 526.

glacialis, 525.

Scoliodon, 265, 292, 299, 303.

acutus, 303.

dumerilii, 293.

eumcces, 303.

intermedius, 294.

jordani, 293.

lalandii, 294 note 3, 297 note 5, 299, 301.

longmani, 294.

longurio, 294, 300.

porosus, 303.

scrrakowah, 293, 294 note 4, 299.

terrae-novae, 122, 292, 293, 294, 295, 301,
303 note 19, 307, 308.

vagatus, 294.

walbeehmi, 294, 299.
Scomberomorus maculatus, 441.
Scyliorhinidae, 96, 195, 233.
Scyliorhinus, 197, 199, 202, 214, 219.

boa, 204, 211, 212, 213.

canicula, 202 note 25.

caniculus, 67, 96 note 8, 196 note 2, 197, 202,
203.

capensis, 203.

duhamelii, 203 note 28.

haeckeli, 205.

haeckelii, 59.

regani, 197 note 1 5.

retifer, 203, 205, 206, 207, 211, 212, 219.

stellaris, 196 note 2, 203.

torazame, 204.

torrei, 204, 205, 206, 207, 211.
scyllia, Triakis, 235 note 7, 236.
Scylliformes, 76.
Scylliogaleus, 234.
Scylliorhinus, 202, 214, 219.

boa, 207, 214.

indicus, 219.

profundorum, 225.

punctatus, 198 note 19.

regani, 198 note 19.

retifer, 207, 210, 211.

torrei, 214.
Scyllium, 180, 202, 214, 219.

anale, 198 note 22.

canicula, 96 note 8, 214.

cirratum, 186.

cirrhosum, 180, 186.

edwardsi, 199 note 23.

laurussonii, 219, 224, 225.

marmoratum, 199 note 23.

retiferum, 207, 210.

vincenti, 198 note 22.

572

Index of Scientific Names

scyllium, Triakis, 235, 236 note loa.
Scymnium, 501.

niciense, 507.
Scymnodon, 451, 494 note I.
Scymnorhinus, 500 note 3, 501.

brevipinnis, 508.

licha, 508.

lichi (Scymnus lichia), 508.

lichia, 507.

phillippsi, 508.
Scymnus, 501, 507, 508, 514, 526.

americanus, 506.

borealis, 514, ;24.

(Laemargus) borealis, 514, 524.

brasiliensis, 508, 513.

(Scymnus) brasiliensis, 513.

(Scymnus) brasiliensis var. torquatus, 513.

(Scymnus) brasiliensis var. unicolor, 513.

brevipinna, 524.

(Laemargus) brevipinna, 526.

glacialis, 524.

gunneri, 524.

licha, 507.

lichia, 501, 507, 508.

(Scymnus) lichia, 507.

microcephalus, 525.

micropterus, 524.

nicaensis, 507.

rostratus, 5 1 4.

spinosus, 531.

torquatus, 513.

vulgaris, 507.
scymnus, Squalus, 507.
Sebastes, 520.
Selache, 146, 189.

elephas, 158.

maxima, 146, 157, 189, 194.

maximum, 157 note 39.

maximus, I 57 note 39.

rostrata, 160.
Selachii, 64, 76.
Selachophichthyoidi, 94 note 2.
Selachus, 146.

maximus, 146, 158.

pennanti, 160.
Selanche, 146.

maximus, 146, 158.
Selanonius, ill.

walkeri, III, 122.
semifasciata, Triakis, 235 note 7, 236.
shavianus, Cetorhinus, 157.
sibogae, Apristurus, 221.

signatus, Carcharias (Hypoprion), 319.

Hypoprion, 316, 319.
silus, Argentina, 498.
sitakaiensis, Apriondon, 309 note 3.
smithii, Carcharodon, 133 note I, 142.
solanonus, Squalus, 120.

Somniosus, 65 note 4, 66, 67, 71, 75, 494, 495,
497 note 4, 500, 507, 514, 519 note 14.

antarcticus, 515.

brevipinna, 514, 515 note 5, 523, 524.

carcharias, 526.

echinatum, 5 14.

microcephalus, 516, 525.

norwegianus, 514.

pacificus, 515, 516, 525.

rostratus, 5 16, 520.
sorra, Carcharias (Prionodon), 32:.
sorrakowah, Carcharias, 292.

Scoliodon, 293, 294 note 4, 299.
spalanzanii, Oxyrrhina, I 31.
spallanzanii, Isurus, 131.

Lamna, 131,

Oxyrhina, 131.
sparophagus, Dalatias, 500, 506.
spenceri, Carcharias, 321.
Spheroides, 248.
Sphyra, 408.
Sphyraena, 69.
Sphyrna, 407, 408, 449.

bigelovi'i, 410, 414, 416, 420, 428, 435, 436.

chiereghini, 435.

corona, 409 note 4, 414.

diplana, 72, 97, 410, 411, 412, 413, 414, 415,
420, 427, 428, 431, 432, 433, 434, 435,
436. 439. 4+0, 441, 442, 443. 444. 445.
446, 447, 448 note 48.

lewini, 408, 419, 449-

malleus, 435, 447.

media, 409 note 4, 414.

tiburo, 409, 411,41 3, 414, 416, 420, 426, 428,

436.439.447-
(Reniceps) tiburo, 428.
tudes, 72, 409, 410, 411, 412, 413, 414, 416,

419, 420, 421, 426, 428, 435, 436, 439, 441,

442, 444, 445, 447.
(Platysqualus) tudes, 435.
vespertina, 409 note 4, 420, 425 note 20, 427,

428.
zygaena, 408, 409, 411, 412, 413, 414, 415,

416, 417, 418, 419, 420, 422, 424, 427,

428, 431, 432, 433, 434, 435, 436, 444,

448, 449.

Index of Scientific Names

573

(Cestracion) zygaena, 447.

(Zyg.iena) zygaena, 449.
Sphyrnias, 408.

tiburo, 427.

tudes, 435.

zygaena, 420, 447.
Sphyrnichthys, 408.

zygaena, 435.
Sphyrnidne, 96, 407.
Spinax, 452, 470, 471, 487.

acanthias, 470, 472, 486.

fabric!!, 480, 486.

fernandez!anus, 480.

hilHanus, 493.

pus!llus, 493.

sp!nax, 493.

(Acanth!as) suckleyi, 472.

vulgaris, 472.
spinax, Etmoptcrus, 488, 490, 492, 493.

Spinax, 493.

Squalus, 470, 487.
spinosus, Centrophorus, 526.

Echinorhinus, 526, 530, 53 1.

Goniodus, 53 I.

Scymnus, 531.

Squalus, 530.

Squalus (Scymnus), 531.
spongiceps, Apristurus, 20, 221 note 10, 513.

Catulus, 2ig, 513.
springer!, Carcharhinus, 324, 384, 386, 404.

Eulamia, 407.
Squalida, 76.

Squalidae, 450, 452, 481, 487, 493.
Squaliolus, 500 note 2.
Squallus, 452.
Squalo, 85.

Squaloidea, 77, 449, 326, 527, 532.
Squalus, 98, 133, 146, 161, 180, 214, 265, 280,
292, 303, 315, 362, 377, 451, 452, 472,
478, 485, 500 note 3, 502, 508, 514, 516,
518.

acanthias, 65 note 4, 66, 73, 1 17, 250, 2S6, 452,
453. 454. 455, 467. 470 note 54, 472, 473,
474. 475. 476, 477. 478, 479, 482, 486,
490,494, 529.

(Acantherinus) acanthias, 471.

(Spinax) acanthias, 471.

acanthias var. fernandinus, 480.

achantias, 470.

acronatus, 328.

acutipinnis, 454 note 21a, 480.

adscensionis, 289.

afric.inus, 1 97 note 16, 202.

alopecias, 1 76.

americanus, 107, 472, 501, 506.

(Acanthorhinus) americanus, 507.

arcticus, 265, 271, 272.

argus, I 86.

barbouri, 455 note 23, 464, 469, 472.

blainvillc, 453, 454, 455, 456, 472, 474, 478

note 63, 479, 480.
blainvillc!, 478, 480.
borealis, 5 14, 524.
brcvipinna, 515 note 5, 523.
brevirostris, 453, 454.
brucus, 5 14, 526, 530.
caccchia, 377.

(Carcharhinus) cacruleus, 290.
(Carcharinus) caeruleus, 376.
caniculus, 202.
canis, 240, 251.
carcharias, 133, 142, 144, 145, 273, 320 note la,

354 note 51, 362, 514, 515, 523.
(Carcharhinus) carcharias, 142.
(Carcharias) carcharias, 142.
carolinianus, 428.
(Carcharinus) caudata, 376.
cepedii, 131, 133.
cetaceous, 160.
cinercus, 92.

(Monopterhinus) cinereus, 92.
(Notidanus) cinercus, 93.
cirratus, 180, 185.
cirrhatus, 186.
coeruleus, 376.
cornubicus, 1 1 1, i 20.
(Carcharias) cornubicus, 122.
(Lamna) cornubicus, 122.
cornubiensis, 120.
cubcnsis, 453, 454, 455, 456, 464, 469, 472,

473, 478, 479, 480, 482, 490, 494.
cuvier, 265, 272, 274.
elcphas, 158.
fernandezianus, 480.
fernandinus, 453, 454, 455, 456, 472, 473,

478, 480.
ferox, 98, 108.
fulgens, 5 I 3.

(Scymnus) fulgens, 50S, 513.
galeus, 240.
glacialis, 524.

glaucus, 1 19, 120, 280, 289, 377 note 104.
(Carcharhinus) glaucus, 290.
(Carcharias) glaucus, 290.

574

Index of Scientific Names

griffini, 452, 454.

griscus, 78, 79, 85.

(Monopterhinus) griseus, 86.

(Scymnus) gunneri, 524.

gunnerianus, 146, 157.

(Galeorhinus) hinnulus, 253.

homianus, I 57.

isodus, 158.

japonicus, 454 note 20, 480.

kirki, 453 note 15.

(Galeorhinus) laevis, 253.

liburo, 426 note 24.

licha, 500, 501, 506.

(Scymnus) lichia, 508.

littoralis, 107.

longimanus, 362.

macrodous, 1 07.

malleus, 420, 444.

(Carcharias) maou, 145.

maximus, 146, 156.

(Selache) maximus, 158.

megalops, 452, 454, 480 note 66.

microcephalus, 515, 523.

milberti, 376, 377, 378.

(Carcharinus) milberti, 376.

mitsukurii, 454 note 20, 473, 480.

monensis, 120.

montalbani, 480.

mustelus, 240.

nasus. III, 120.

nicaense, 501.

nicaensis, 506.

norvegianus, 524.

norvegicus, 525.

obscurus, 292, 389.

(Carcharinus) obscurus, 389.

obtusus, 344.

pelegrinus, 157.

pennanti, 1 20.

peregrinus, 157.

perlo, 87, 92.

phillppinus, 480.

platyodon, 344, 377.

plumbeus, 377.

porosus, 303.

punctatus, 122, 186, 292, 303, 307, 308.

punctulatus, 186.

rashleighanus, 158.

rhinoceras, 158.

rostratus, 158, 515.

scymnus, 507.

solanonus, 120.

spinax, 470, 487.

spinosus, 530.

(Scymnus) spinosus, 531.

squamosus, 45 1 note 8.

squatina, 523, 534, 544, 546.

sucklcyi, 453, 454 note 20, 463, 472, 473.

sucklii, 472.

sucklii, var. mitsukurii, 473.

tasmaniensis, 454 note 20, 480.

(Carcharias) terrae-novae, 301.

tiburo, 333, 408, 420, 425, 426, 434.

vacca, 85.

(Carcharias) vulgaris, 142.

vulpes, 1 74.

(Carcharhinus) vulpes, 174.

vulpinus, 161, 174.

waddi, 180 note 5.

wakiyae, 473.

whitleyi, 453 note 15.

zygaena, 408, 420, 435, 444, 448.
squamosus, Squalus, 45 1 note 8.
Squatina, 534, 544.

africana, 537.

angelus, 543, 546.

argentina, 535, 536 note 16, 537, 544.

armata, 535, 537, 545, 546.

australis, 535, 536.

californica, 535, 537, 540 note 24.

dumeril, 535, 537, 538, 543, 545.

dumerili, 543.

dumerilii, 543.

japonica, 535, 536 note 19, 537.

laevi, 543.

nebulosa, 536.

oculata, 536.

philippi, 537.

squatina, 535, 536, 537 note 20, 541, 542, 543>
544. 545. 546.

tergocellata, 536.

vulgaris, 534.
squatina, Rhina, 543, 544.

Squalus, 523, 534, 544, 546.

Squatina, 535, 536, 537 note 20, 541, 54^. 543.
544. 545. 546.
Squatinidae, 534-
Squatinoidea, 64, 77, 533.
Squilla, 424.
Stegostoma, 178, 179.
stellaris, Scyliorhinus, 196 note 2, 203.
Stenotomus, 104, 248, 441, 462.
stenurus, Amphioxides, 24, 25, 27.
stevensi, Carcharias, 321.

Index of Scientific Names

SIS

stouti, Eptatretus, 31.
striatus, Mustelus, 259.
subarcuata, Ccstracion, 447.

Zygaena, 447.
suckleyi, Acanthias, 472.

Spinax (Acanthias), 472.

Squalus, 453, 454 note 20, 463, 472, 473.
sucklli, Acanthias, 472.

Squalus, 472.

var. mitsukurii, Squalus, 473.
superciliosus, Alopecias, 167.

Alopias, 162, 163, 168.
Sutorectus, 179.
Synodontaspis, 98.

lASMANiENSis, Squalus, 454 note 20, 480.
tattersalli, Branchiostoma, 10.
taurus, Carcharias, 70 note 17, 98, 99, 1 00, 106.

Odontaspis, 1 08.

Triglochls, 108.
Tautoga, 104, 248.
Tautogolabrus, 1 04.
Tectospond}-li, 76 note 27.
temmlncki, Carcharias (Prionodon), 321.
tergocellata, Squatina, 536.
terrae-novae, Carcharhinus, 302.

Carcharhinus (Scoliodon), 302.

Carcharias, 302.

Carcharias (Scoliodon), 302.

Lamna, 301.

Scoliodon, 122, 292, 293, 294, 295, 301, 303
note 19, 307, 308.

Squalus (Carcharias), 301.
Tetraoras, 146.

angiona, 146, 160.
Tetrcrus angiova, 160.
Thalassorhinus, 263.
tiburo, Cestracion, 427.

Cestrorhinus, 427.

Platypodon, 333.

Platysqualus, 427.

Prionodon, 333.

Reniceps, 427.

Sphyrna, 409, 411, 413, 414, 416, 420, 426,
428,436,439,447.

Sphyrna (Reniceps), 428.

Sphyrnias, 427.

Squalus, 333, 408, 420, 425, 426, 434.

Zygaena, 427, 428.

Zygaena (Squalus), 427.

tigrinus, Galeocerda, 273, 274.

Galeus, 274.

Isurus, 265, 274.
tigris, Carcharias, 131.

Isurus, 132, I 33.

Lamna, 132.
torazame, Scyliorhinus, 204.
torquatus, Scymnus, 513.

Scymnus (Scymnus) brasiliensis, var., 513.
torrei, Scyliorhinus, 204, 205, 206, 207, 211.

Scylliorhinus, 2 1 4.
Trachinocephalus, 387.
Trachinus draco, 461.
triacanthus, Poronotiis, 350.
Triacis, 235.
Triaenodon, 234, 309 note 2.

obesus, 309 note 2.
Triakidae, 97, 197 note 12, 233.
Triakis, 234, 235.

barbouri, 235, 236, 240.

henlei, 235, 236.

leucoperiptera, 236.

maculata, 234, 235 note 7, 236.

nigromaculata, 236 note 10.

scyllia, 235 note 7, 236.

scyllium, 235, 236 note loa.

semifasciata, 235 note 7, 236.

venusta, 23;, 237.
tricuspidatus, Carcharias, gg.
tridentiger, Myxine, 33.
Triglochis, 98.

taurus, 108.
tudes, Cestracion, 435, 436.

Cestracion (Zygaena), 435.

Platysqualus, 436.

Sphyrna, 72, 409, 4 10, 41 1, 41 2, 41 3, 41 4, 4 1 6,
419, 420, 421, 426, 428, 435, 436, 439, 441,
442,444,445,447.

Sphyrna (Platysqualus), 435.

Sphyrnias, 435.

Zygaena, 420, 434, 436.
typicus, Rhineodon, 188, 194.

Rhinodon, 188, 193.
typus, Rhineodon, 188, 189, 193.

Rhineodon, I 94.

Rhinodon, 194.

Unicolor, Ammocoetes, 57.
Petromyzon, 57.
Pctromyzon marinus, 57.
Scymnus (Scymnus) brasiliensis, var., 513.

576

Index of Scientific Names

Uranga, 321.

nasuta, 321.
Uranganops, 321.
Urophycis, 39, 49.
nstus, Carcharhinus, 320 note I a.
uyato, Galeus, 214 note i.

Vacca, Notidanus, 86.

Squalus, 85.
vagatus, Scoliodon, 294.

yaldiviae, Amphioxides, 20, 24, 25, 26, 27, 28.
velox, Carcharhinus, 391.
venusta, Calliscyllium, 235.

Triakis, 235, 237.
Vertebrata, i.
verus, Carcharhinus, 320 note la.

Carcharias, 133, 142.

Carcharodon, 133, 142.
verweyi, Apristurus, 220, 221 note 10.
vespertina, Sphyrna, 409 note 4, 420, 425 note 20,

427, 428.
villosus, Etmopterus, 488 note 3.
vincenti, Scyllium, 198 note 22.
virens, Pollachius, 286, 520.
virginiae, Branchiostoma, 9, 10 note 20, 16.
vorax, Carcharias, 145.
vulgaris, Acanthias, 452, 454, 470, 472, 478.

Carcharias, 142.

Catulus major, 202.

Mustelus, 254.

Notidanus, 87.

Scymnus, 507.

Spinax, 472.

Squalus (Carcharias), 142.

Squatina, 534.

Zygaena, 447, 449.
Vulpecula, 1 6 1.

marina, 161, 176, 177.
vulpes, Alocepias, 1 76.
vulpes. Alopecias, 175, 177.

Alopes, 161, 177.

Alopias, 175, 177, 320 note la.

Carcharhinus, 320 note la.

Carcharias, 161, 163, 174, 177.

Squalus, I 74.

Squalus (Carcharhinus), 174.
vulpinus, Alopias, l6l, 162, 164, 166, 167, 176,
177, 178.

Squalus, 161, 174.
vulpis, Alopias, 175.

Waddi, Squalus, 180 note 5.
Waitei, Centrophcrus, 451 note 8.
wakiyae, Squalus, 473.
walbeehmi, Carcharias (Scoliodon), 303.

Scoliodon, 294, 299.
walbenii, Carcharias (Scoliodon), 303.
walkeri, Selanonius, ill, 122.
whitleyi, Lamna, 1 1 1 note 5.

Squalus, 453 note 15.
wilsoni, Oceanomyzon, 45, 46, 57.
wolniczkyi, Notidanus, 87 note 2.

A.IPHIAS, 84.
gladius, 129.

YoLDiA, 373.

Z/EVS faber, 1 17.
Zev, 179 note 3.
zigaena, Zygaena, 447.
Zostera, 1 1 9, 248.
Zygaena, 408, 427, 447.

leeuwenii, 425 note 19, 428.

lewini, 428, 449.

malleus, 418 note 10, 419, 420, 446, 448.

subarcuata, 447.

tiburo, 427, 428.

(Squalus) tiburo, 427.

tudes, 420, 434, 436.

vulgaris, 447, 449.

zigaena, 447.

zygaena, 446.
zygaena, Cestracion, 420, 428, 435, 447, 449.

Cestracion (Sphyrna), 449.

Cestracyon, 449.

Sphyrna, 408, 409, 411, 412, 413, 414, 415,
416, 417, 418, 419, 420, 422, 424, 427,
428, 431, 432, 433, 434, 435, 436, 444,
448, 449.

Sphyrna (Cestracion), 447.

Sphyrna (Zygaena), 449.

Sphyrnias, 420, 447.

Sphyrnichthys, 435.

Squalus, 408, 420, 435, 444, 448.

Zygaena, 446.
Zygana, 408.
Zygoena, 408.
zyopterus, Galeorhinus, 264 note 4.

Fishes of the

Western North Atlantic

Part I

COMPOSED AND PRINTED BY E. L. HILDRETH & COMPANY

IN BRATTLEBORO, VERMONT

IN AN EDITION OF 25OO COPIES ON FIFTY PER CENT RAG TEXT PAPER

SPECIALLY MADE BY THE CURTIS PAPER COMPANY

DESIGNED BY LEWIS F. WHITE

REPRINTED BY BIANCO LUNGS BOGTRYKKERI A-S

COPENHAGEN, DENMARK

IN AN EDITION OF I OOO COPIES ON FIFTY PER CENT RAG TEXT PAPER

SPECIALLY MADE BY DE FORENEDE PAPIRFABRIKKER A-S

COPENHAGEN, DENMARK

BOUND IN BOARDS AND FULL CLOTH,

EUROPEAN LINEN FINISH; STAMPED IN GOLD