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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 (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 ^ 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\ 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