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

Fishes of the Western North Atlantic 



MEMOIR 

SEARS FOUNDATION FOR MARINE RESEARCH 

Number I 

Fishes of the 
Western North Atlantic 





PART TWO 

Sawfishes^ Guitarfishes^ 
Skates and Rays 

HENRY B. BIGELOW and WILLIAM C. SCHROEDER 

Museum of Comparative Zoology 

Chimaeroids 

HENRY B. BIGELOW and WILLIAM C. SCHROEDER 

NEW HAVEN 1953 
SEARS FOUNDATION FOR MARINE RESEARCH, YALE UNIVERSITY 



PRINTED IN DENMARK FOR 

SEARS FOUNDATION FOR MARINE RESEARCH 

BINGHAM OCEANOGRAPHIC LABORATORY 

YALE UNIVERSITY 

Albert E. Parr, Editor 

Yngve H. Olsen, Assistant Editor 

Bianco Luno's Printing, Copenhagen, Denmark 



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 

ALBERT E. PARR 

American Museum of Natural History 

WILLIAM C. SCHROEDER 

Museum of Comparative Zoology 

LEONARD P. SCHULTZ 

United States National Museum 

NEW HAVEN 1953 



SEARS FOUNDATION FOR MARINE RESEARCH, YALE UNIVERSITY 



Table of Contents 



INTRODUCTION XI 



Mai 



XIV 



SAWFISHES, GUITARFISHES, SKATES and RAYS, 

By Henry B. Bigelow and William C. Schroeder I 

Acknowledgments I 

General Discussion 2 

Subclass Elasmobranchii 4 

Order Batoidel 4 

Suborder Pristoidea 15 

Family Pristidae 18 

Genus Pristis 18 

Pristis pectinatus 23 

Pristis perotteti 34 

Suborder Rhinobatoidea 43 

Family Rhynchobatidae 44 

Family Rhinobatldae 46 

Genus Rhinobatos ^O 

Rhinobatos horkelii 56 

Rhinobatos lentiginosus 60 

Rhinobatos percellens 6 8 

Genus Rhinobatos Addendum 73 

Rhinobatos spinas us 73 

Genus Zapteryx 74 

Zapteryx brevirostris 75 

Suborder Torpedinoidea 80 

Family Torpedinidae 87 

Genus Torpedo 90 

Torpedo nobiliana 96 

Genus Narcine 107 

Narcine brasiliensis ill 

Genus Diplobatis 123 

Diplobatis pictus 1 24 

Genus Benthobatis 126 

Benthobatis marcida 128 

Suborder Rajoidea 132 

Family Rajidae 133 



viii Table of Contents 



Genus Raja 




138 


Raja ackleyi 




'55 


Raja bathyphila 




159 


Raja eglanteria 




165- 


Raja erinacea 




176- 


Addendum to Raja erinacea 


by D. Merriman, 




Y. H. Olsen, S. B. Wheatland, and 




L. H. Calhoun 




187 


Raja fyllae 




194 


Raja garmani 




200 


Raja hyperborea 




206 


Raja jenseni 




213 


Raja laevis 




217 


Raja lentiginosa 




228 


Raja lint e a 




232 


Raja mollis 




237 


Raja ocellata 




240 


Raja olseni 




251 


Raja radiata 




^55 


Raja senta 




264 


Raja spinicauda 




271 


Raja teevani 




276 


Raja texana 




279 


Genus Breviraja 




284 


Breviraja atripinna 




286 


Breviraja colesi 




289 


Breviraja cubensis 




293 


Breviraja plutonia 




297 


Breviraja sinus-mexicanus 




302 


Breviraja spinosa 




306 


Breviraja yucatanensis 




310 


Genus Cruriraja 




313 


Cruriraja atlantis 




3^5 


Cruriraja poeyi 




319 


Genus Dactylobatus 




323 


Dactylobatus armatus 




323 


Family Anacanthobatidae 




327 


Genus Springeria 




328 


Springeria folirostris 




328 


Suborder Myliobatoidea 




331 


Family Dasyatidae 




335 


Genus Dasyaiis 




340 



Table of Contents ix 

Dasyatis americana 34^ 

Dasyatis centroura 3^2 

Dasyatis geijskesi 363 

D asyatis guttata 365 

Dasyatis sabina 370 

Dasyatis say 378 

Genus Himantura 389 

Himantura schmardae 390 

Family Gymnuridae 395 

Genus Gymnura 396 

Gymnura altavela 399 

Gymnura micrura 408 

Family Urolophidae 416 

Genus Urolophus 418 

Urolophus jamaicensis 420 

Genus Urotrygon 427 

Urotrygon microphthalmum 428 

Urotrygon venezuelae 430 

Family Myliobatidae 433 

Genus Myliobatis 435 

Myliobatis freminvillii 438 

Myliobatis goodei 446 

Genus Aetobatus 451 

Aetobatus narinari 453 

Family Rhinopteridae 465 

Genus Rhinoptera 465 

Rhinoptera bonasus 469 

Rhinoptera brasiliensis 477 

Family Mobulidae 480 

Genus Mobula 486 

Mobula hypostoma 488 

Mobula mobular 495 

Genus Ceratobatis 497 

Ceratobatis robertsii 497 

Genus Manta 500 

Manta birostris 502 

CHIMAEROIDS By Henry B. Bigelow and William C. Schroeder 515 

General Discussion 515 

Subclass Holocephali 515 

Order Chimaerae 516 

Family Chimaeridae 523 



X Table of Contents 

Genus Chimaera 524 

Chimaera cubana _ 527 

Genus Hydrolagus 533 

Hydro! agus affinis 539 

Hydrolagus albertt 545 

Family Rhinochimaeridae 548 

Genus Harriotta 550 

Harriotta raleighana 551 

Family Callorhinchidae 558 

Genus Callorhinchus 560 

Index S^Z 



Introduction 



WITH THE PUBLICATION of the second part of Fishes of the Western 
North Atlantic it seems desirable to restate briefly the purposes and aims of 
this series, particularly for those who may not be familiar with Part One. In ich- 
thyology, as in other sciences, much of the early work was devoted to purely de- 
scriptive accounts which appeared in numerous publications here and abroad, and 
although new species are still described occasionally, a stage of relative stability has 
been achieved in our knowledge of the species which occur in the western North 
Atlantic. At this time, therefore, it seems particularly important to bring together 
and synthesize the vast amount of information which has been amassed in the past 
and thus make it available to both public and marine biologists. It is intended that 
each article in this series shall be a critical review and revision rather than a mere 
compilation of previously published works, and, as pointed out in the Preface to 
Part One, that it be "written on the premise that it should be useful to those 
in many walks of life — to those casually or vitally interested in the general phe- 
nomena of life in our waters, to the sportsman whose interests are closely associated 
with pleasure and relaxation, to the fisherman whose livelihood depends upon know- 
ledge of where fishes are gathered together, as well as to the amateur ichthyologist 
and the professional scientist." 

Part One, published in 1948, dealt with the Lancelets, Cyclostomes and Sharks 
of the western North Atlantic. The present publication, treating of Sawfishes, Gui- 
tarfishes. Skates, Rays and Chimaeroids, brings to completion a further effort in the 
over-all purpose of this series. In general the format and treatment of these groups 
are the same as in Part One. In conformance with general policy, these studies also 
are written for the layman as well as the specialist and are critical reviews rather 
than perfunctory compilations. The text, together with meticulously accurate illustra- 
tions, should leave no place for confusion in the reader's mind. As in Part One, 
under each species will be found not only a detailed description and the distinctive 
characters which set it apart from its nearest relatives, but notations on color, size, 
general habits, abundance, range, relation to man, and occurrence. The geographical 
range of the fishes described remains essentially the same — the western half of 
the North Atlantic, including the adjoining gulfs and seas, from Hudson Bay south- 
ward to the Amazon River. Numerous species found in other parts of the globe are 
often referred to, and occasional species from adjoining seas are described. 



xii Introduction 

Again we wish to acknowledge our gratitude to the Sears Foundation for Marine 
Research, which has financed the publication. We are also grateful to the Woods 
Hole Oceanographic Institution and the Museum of Comparative Zoology, which 
have contributed in large measure to these publications through salaries to the authors 
and through special grants for the preparation of illustrations and for travel to 
examine specimens. We are likewise indebted to those institutions which have gener- 
ously supported the work of the editors, particularly the Bingham Oceanographic 
Laboratory, the American Museum of Natural History and the New York Zoolog- 
ical Society. 

The Editorial Board again expresses its particular appreciation and thanks to 
Yngve H. Olsen, who has edited the manuscripts and seen them through press with 
untiring patience and enormous skill and to Henry Sears for his continued interest 
and support. 

Finally, we note with profound regret the death of the late Dr. Samuel F. Hilde- 
brand, who served the field of ichthyology so well and long and who gave so generously 
of his time and experience as a member of our Editorial Board. 

John Tee- Van, 
New York Zoological Society. 
February i, 1953 



lOCfTS* 50* 25* 




North America 



80* 75' 



60* 65* 



50* 45* 




45* 



70* 65' 60* 65* 50* 



South America 



CHAPTER ONE 

Sawfishes^ Guitarfishes, 
Skates and Rays^ 

BY 
HENRY B. BIGELOW and WILLIAM C. SCHROEDER 

ACKNOWLEDGMENTS 

Many persons have assisted us in the preparation of the following pages. Our thanks 
are due in particular to Leonard P. Schu/tz and to A. S. Romer for allowing us 
free access to the collections of the United States National Museum and of the Har- 
vard Museum of Comparative Zoology, which have been the chief sources of our 
Study Material; to the authorities of the British Museum (Natural History), the 
American Museum of Natural History, the Academy of Natural Sciences of Phila- 
delphia, the Boston Society of Natural History, the Chicago Museum of Natural 
History, the Charleston Museum, South Carolina, and the Museum of Zoology, 
University of Michigan, for the loan of much needed material; to the Woods Hole 
Oceanographic Institution for constant assistance ; to Stewart Springer who has con- 
tributed a large amount of valuable information and many specimens of Skates, Rays 
and Chimaeras from Florida and the Gulf of Mexico ; to Anton Fr. Bruun for the 
loan of rhinochimaeroid material collected by the Atlantide Expedition from deep 
water off equatorial West Africa ; to J. L. Baughman for Sawfishes, Guitarfishes, 
and other specimens from the Gulf of Mexico ; to Gordon Gunter for Guitarfishes 
from Texas ; to Michael Lerner and C. M. Breder for obtaining two specimens of 
M.zntzfor us at Bimini, Bahamas, and to Ivor Cornman for arranging to have parts 
of them shipped to Cambridge ; to William Royce and the scientific staff of the 

I. Contribution No. 570 from the Woods Hole Oceanographic Institution. 

I 

I 



2 Memoir Sears Foundation for Marine Research 

U. S. Fish ami Wildlife Service vessel Albatross III for desirable specimens and 
data on Skates and Rays jrom the North Carolina Coast taken during midwinter ; 
to Luis Ho%vell-Ii.ivero for Cuban Rays; to Captain R. Howard, A. U.S., for Tor- 
pedo and Sting Rays from the east coast of Florida ; to Marie Poland Fish for Rays 
from Rhode Island ; to Leslie W. Scatter good for information as to the season for 
Raja ocellata with egg cases off the coast of Maine ; to Paul M. Hansen for speci- 
mens o/'Raja radiatayrow Greenland and for other information ; to Captain C. W. 
Thomas, U. S. C. G., and Commander P. L. Stimpson, U. S. C. G.,for obtaining spe- 
cimens of Raja hyperboreayor us in West Greenland ; to Commander David C. Nutt 
and Richard H. Backus for records of Raja radiata from Labrador ; to W. Temple- 
man for a specimen of Raja spinicauda from northeastern Newfoundland, for spe- 
cimens oj R. radiata yro/w the Grand Banks, and for other information ; to V. D. 
V lady kov for specimens o/^Raja erinacea from the estuary of the St. Laivrence River; 
to Miss Ethelwyn Trewavas of the British Museum (Natural History ) for supplying 
the description of the type specimen of the Devil Ray Ceratobatis robertsii and for 
arranging to have drawings of it prepared by Hubert Williams ; to G. Palmer for 
drawings and measurements o/Diplobatis pictus; to Dr. George White, "J. Incar- 
done, and W. E. Kruger for the x-ray photographs reproduced in Fig. 62, and to Dr. 
Leonard D. Nathan for his assistance in obtaining x-rays ; to Sehora Helena Paes 
de Oliveira for photographs of Brazilian Skates; to John Tee-Van for contributing 
bibliographic citations without which the work could hardly have been undertaken ; 
to Ludloiv Griscom and 'James L. Peters for assistance in nomenclatural problems ; 
to H. W. Fowler for welcome information on various Skates and Rays ; to Frank 
Huber for notes on a large Manta taken off New Jersey ; to F. E. Firth, and Cap- 
tains Donald Campbell and Frank Janssen for interesting records of offshore cap- 
tures o/^Torpedo nobiliana; to Captains Henry W. Klimm, Jr. and Jared Vincent 
for furnishing information on the captures of various Skates on the offshore winter 
fishing grounds off southern New England; to J. C. Brew, Sir P. H. Buck and 
Captain E. H. Bryan for assembling information on the use of Sting Ray spines as 
weapons in various parts of the world, and to Airs. Myvanwy Dick for much assi- 
stance in the preparation of the manuscript. 

GENERAL DISCUSSION 

Scope of Study. The following pages give descriptions, illustrations, life histories, geo- 
graphic distribution, and references for all species of Sawfishes, Guitarfishes, Skates and 
Rays that are known in the western side of the North Atlantic. The characterizations 
of suborders, families, and genera, together with the corresponding Keys, cover the 
batoid fishes as a whole ; this also applies to Species Keys in cases where present know- 
ledge permits. 



Fishes of the IP^estern North Atlantic 3 

Descriptions. The descriptions are based on the Study Material listed under each 
species, except for Urotrygon microphthalmum and Mobula mobular, no specimens of which 
were available. The accounts of habits and distribution are based on published data, 
on information from correspondents whose assistance is acknowledged, and on our 
own observations. 




Figure i. Outlines of a typical Skate to illustrate terminology and methods of measurement. 



4 Memoir Sears Foundation for Marine Research 

Keys. The Keys are offered solely for purposes of identification. Therefore, we have 
selected for their construction such characters as are not only alternative but which are 
easily seen or measured. 

Arrangement. The larger groups, down to genera, are arranged in the sequence that 
seems to us to represent most nearly their probable relationships. The species within 
each genus are presented in alphabetical sequence as in our previous publication. Sharks.^ 

References. All citations were consulted in the original, except as noted, through 
the co-operation of the several libraries listed previously.^ 

Sources of Material. The collections in the Harvard Museum of Comparative 
Zoology and the United States National Museum have been the chief sources of our 
Study Material. We are much indebted also to the museums and persons listed (page i). 

Proportional Dimensions and Illustrations. The measurements from which the pro- 
portional dimensions of the several species have been calculated were taken on a hor- 
izontal line between perpendiculars at given points ; for example, the length of the snout 
in front of the orbits is on line BC in Fig. I, not AC ; the length of the disc is measured 
on line BC, not AC ; the anterior angle to the level of the spiracles is the angle DAE. 

The great majority of the drawings in this book were prepared by the well known 
zoological artist, E. N. Fischer, who executed most of the illustrations for Part I of 
this Memoir series and for Carman's Plagiostoma (19 13). Ceratobatis robertsii., the type 
specimen of which is in the British Museum (Natural History), was drawn by Hubert 
Williams, and original drawings of Diplabatis pictus were loaned to us by courtesy of 
G. Palmer. 

Subclass ELASMOBRANCHII 

Order BATOIDEI 

Sawfishes, Guitarfishes, Skates and Rays^ 

Characters. In the Batoidei the gill openings are wholly on the ventral surface. 
The anterior edges of the pectorals are united with the sides of the head forward past 
all five pairs of gill openings, about to the level of the mouth in some (Pristidae), to the 
level of the nostrils in others, to the tip of the snout in still others. The upper edges 
of the orbits are not free from the eyeballs, as they are in Sharks; that is, they do not 
have free upper eyelids. In these respects they differ from all modern Sharks. None 
have nictitating membranes, anal fin, or precaudal pits or furrows. 

The great majority of the batoids are easily recognizable by their shapes. Their 
trunks are strongly flattened dorsoventrally, with the pectorals widely expanded so that 
they are disc-like in shape. The tail sector is more or less distinct from the body sector, 
the eyes and spiracles are on the dorsal surface, and the mouth, as well as the entire 
lengths of the gill openings, is on the ventral surface. 

2. Bigelow and Schroeder, Mem. Sears Found. Mar. Res., I (i) Chap. 3, 194S. 

3. Mem. Sears Found. Mar. Res., I (i), 1948: 60. 

4. For characters of the order Selachii, see Mem. Sears Found. Mar. Res., I (i), 1948: 62-64. 



Fishes of the IV es tern North Atlantic 5 

However, one group, the Sawfishes (Pristoidea), are shark-like in general appear- 
ance, though they are grouped among the batoids because of the relationship of the 
pectorals to the gills and because of the absence of upper eyelids, as well as on skeletal 
grounds. And most of the Guitarfishes (families Rhynchobatidae and Rhinobatidae) 
are Intermediate in form, between shark-like and batoid-like. 

Some batoids have no dorsal fin, whereas others have one or two, in which case 
the first dorsal (when there are two) varies in position from over the pelvics to far back 
on the tail. Some have a distinct caudal fin, which others lack. In all fins of some species 
the radial cartilages are supplemented distally (as in Sharks) by much more numerous 
fine horny rays (ceratotrichia) in double series, their inner ends embracing the outer 
ends of the cartilaginous radials ; in others the unpaired fins have these horny rays 
whereas the pectoral and pelvic fins, in which the cartilaginous radials run out nearly 
or quite to the margin, do not ; in still others neither the unpaired nor paired fins have 
horny rays. The spiracles are larger than those of most Sharks and are situated on top 
of the head in all cases ; the rudimentary spiracular gill filaments are better developed 
than in Sharks, as is also the so-called spiracular valve, a stiff concentric fold of con- 
nective tissue on the anterior margin of the spiracle which is supported by a strong 
cartilage fixed at each end. The eyes are well developed in most, though degenerate 
in a few. All have five pairs of gill openings. 

The skins of some are naked while those of others are variously armed with thorns, 
tubercles or prickles ; the tails of some bear large saw-edged spines. The teeth vary 
from thorn-like to rounded or platelike (none have the blade-like dentition so charac- 
teristic of many Sharks), placed either in bands, in transverse rows, or in pavement 
or mosaic arrangement. The nostrils are connected with the mouth in some cases but 
are entirely separate from it in others. In one family, the Sawfishes, the snout is pro- 
duced as a long flat blade supported by the rostral cartilages in the form of five or more 
calcified tubes (described on p. 16) and is armed along either edge with a single series 
of strong tooth-like structures, much as in the Saw Sharks (Pristiophoroidea). The heart 
valves are in two to seven rows.^ 

Some members of the order have electric organs more or less well developed, 
but none have luminescent organs so far as is known. 

The vertebral column is completely segmented throughout its length in all species, 
with the centra fully differentiated and the axial canal so greatly constricted in its passage 
through them that the notochord is wholly or almost wholly obliterated there. Secondary 
calcifications of the vertebral centra in the form of radiating lamellae are described as 
growing inward toward the primary calcified double cone. 

The Batoidei in which the anatomy has been studied, including the Sawfishes, 
differ from all modern Sharks in the attachment of the pectorals to the sides of the 
head, in the lack of a free upper eyelid, in the better developed spiracles, in the ventral 

5. See Garman (Mem. Harv. Mus. comp. Zool., j6, 1913: pi. 56, figs. 7-10, pi. 57, figs. 1-6) for illustrations from 
original dissections of the heart valves of various batoids; see White (Bull. Amer. Mus. nat. Hist., ^4, 1937: 87, 91) 
for a general discussion of their significance in classification and for tabulations of their numbers in various Sharks 
and batoids. 



6 Memoir Sears Foundation for Marine Research 

positions of the gill openings, and in a number of skeletal characters, of which the follow- 
ing may be selected as the most obvious : 

The upper jaw cartilage is attached only loosely to the cranium, at most, by a 
ligament of considerable length. 

The cranium is firmly connected with the vertebral column by a definite articul- 
ation between its posterior face and the anterior vertebra by means of two condyles which 
are lacking in Sharks. 

The first few vertebrae, difi^ering in number in different families, are united together 
as a continuous rigid tube.* 

The ceratohyal cartilage is attached only to the lower end of the hyomandibular 
and thus plays no direct part in the support of the lower jaw, which is suspended from 
the hyomandibular alone. 

The shoulder girdle is directly and firmly attached to the vertebral column, either 
above the latter by a special scapular element (or elements) or to its sides. 

The propterygial cartilage of the pectoral bears at least as many radials as the 
metapterygial cartilage and many more than the mesopterygial, which is much smaller 
than either the propterygial or the metapterygial. 

The union of the two halves of the upper jaw at the symphysis is much more 
intimate in many batoids than in Sharks, but not in all.' On the other hand, the con- 
nections between the anterior parts of the upper jaw cartilages and the cranium, being 
by ligament only, are less intimate among the batoids than they are in many Sharks, 
though the two groups intergrade in this respect.^ 

It has long been known that the attachment of the pectorals to the sides of the 
head in batoids is a secondary development, for the early embryos of even the most 
highly specialized of them are slender-bodied like those of Sharks, with the rudiments 
of the pectoral fins at first wholly posterior to the gill openings. As development pro- 
ceeds, the pectorals expand rearward as well as forward past the gill openings in a 
form that has been variously described as blade-like or horn-like. The anterior horns 
of the pectorals then fuse with the sides of the head above the five posterior gill openings 
that are destined to persist as such but below the persistent portion of the first embryonic 
gill openings that are destined to form the spiracle of the adult. The sequence of events 
of this transformation of the pectorals has been observed among Skates (Rajidae), 
among Torpedoes (Torpedinidae),* and among Sting Rays. 

6. For excellent illustrations of the anterior part of the vertebral column showing this fusion, see Garman (Mem. Harv. 
Mus. comp. ZooL, j6, 1913: pi- 55, figs. 3-10). 

7. This is true of the Rajidae but not of the genus Gymnura (Dasyatidae), in which the two halves of each jaw cartilage 
are clearly separated. 

8. For further information on this point, see Bigelow and Schroeder (Mem. Sears Found. Mar. Res., I (i), 1948: 65). 

9. The sequence of stages leading to the fusion of the anterior parts of the pectorals with the sides of the head seems 
to have been observed first by Leuckart (Z. wiss. Zool., 2, 1850: 261, pi. 16, figs, i, 2) in the Mediterranean Torpedo 
marmorata more than a century ago. For other illustrations of embryo torpedinids with the anterior parts of the 
pectorals still wholly or partially free, see de Sanctis (Atti Accad. Sci. fis. mat. Napoli, 5, 1873 : pi. i, fig. 9), Goodrich 
(in Lankester, Treat. Zool., 9, 1909: 754, fig. 117), Prashad (Rec. Indian Mus., 19, 1920: pi. 7, figs. 7, 8), and espec- 
ially Waite (Rec. Canterbury [N. Z.] Mus., i [2], 1909: pi. 18, fig. 3); see also p. loi. Fig. 23. The process was de- 
scribed and pictured in some detail by Wyman (Mem. Amer. Acad. Arts Sci., [N. S.] 9, 1867 : 35, i pi.) in his classic 



Fishes of the Western Nort/i Atlantic 7 

Batoids in which the forward wing-like expansions of the pectorals have failed 
to unite with the head are caught occasionally, and monstrosities of this sort have 
served as the bases for new generic names (pp. 138, 139, 397).'" 

The absence of the anal fin among all batoids is equally a secondary character, 
for their embryos develop this fin, as do those of Sharks, but they lose it at an early stage 
in their development.'' 

Spiracular Breathing. It is common knowledge that the spiracles play a much more 
important role in respiration among most of the batoids than in any modern Shark. 
In many of the latter, water is taken in solely through the mouth in breathing and 
chiefly through the mouth even by such of them as inhale somewhat through the spirac- 
les, for example Squatina. Most of the batoids, however, take in water chiefly through 
the spiracles. Although it has been observed that Skates (genus Raja) which usually 
hold the lower surface of the head slightly elevated above the bottom do inhale some 
water through the mouth, it is likely that Sting Rays, when well buried in the sand, 
as they often are, take in water through the spiracles alone.'^ But the Devil Rays (Mo- 
bulidae) respire chiefly or wholly through the mouth, and their spiracles are correspond- 
ingly small (p. 484). 

The spiracles of a Skate or Ray, breathing undisturbed, open and close at roughly 
regular intervals, as one can easily see by watching one in an aquarium, closure being 
effected mainly by the contraction of their anterior margins that bear the rudimentary 
gill filaments and the spiracular valve. The water that is taken in through the spiracles 
when the pharyngeal cavity is expanded is prevented from passing out through the mouth 
by the presence of a broad transverse fold of the oral membrane on the roof of the 
mouth and by a narrower one on its floor ; the water is thus directed to the gill openings 
when the pharyngeal cavity is contracted. When a Sting Ray is buried in the sand, 
the periodic expulsion of water through the gills is made evident by "a regular geyser 
of sand grains arising to an inch or two in height at the anterior margin of the pectoral 
fin."" In one set of experiments,'* the rate of respiration in a Skate (Raja), easily mea- 
sured by timing the contractions and expansions of the spiracles, varied from about 
30 per minute at rest to 47.5 per minute after exercise. 

It is also well known and easily observed that the direction of flow through the 
spiracles is occasionally reversed in Skates (Raja), in Sting Rays (Urolophus), in Guitar- 
fishes (Rhinobatos), and also in Angel Sharks (Squatina). In a Skate at rest this was 
seen to happen at intervals of five to ten minutes. And it has been shown experimentally 
that this spouting can be brought about in various ways, i.e., by fatigue, by partial 

paper on the development of Raja. For accounts among Sting Rays, see Hill (Proc. Linn. Soc. N. S. W. [2] 10, 
1895: 208) and Daniel (Elasmobranch Fishes, 1934: 11, fig. 22, B. C.) for Urolophus. 

10. For a good general account with illustrations, see Gudger (Amer. Mus. Novit., 600, 1933)- 

11. For illustrations showing the anal fin in the embryo Skate, see Wyman (Mem. Amer. Acad. Arts Sci., [N. S.] 9, 
1867: pi. to face p. 44, figs. 6-9). 

12. See Rand (Amer. Nat., 41, 1907: 287-302), for observations and experiments on the spiracular breathing of Skates 
{Raja), and Daniel (Elasmobranch Fishes, 1934: 156-157), for similar observations on Sting Rays [Urolophus), 
Guitarfishes [Rhinohatos), and Angel Sharks [Squatina). 

13. Daniel, Elasmobranch Fishes, 1934: 157. 14. Rand, Amer. Nat., 41, 1907: 293. 



8 Memoir Sears Foundation for Marine Research 

asphyxiation from an excess of carbon dioxide in the water, or by mechanical stimul- 
ation. But seemingly the normal function of this reverse flow is to wash the eyes clean 
of sand grains that may fall upon them and to clear the spiracles of fragments of seaweed 
and other objects that may be drawn into them with the intake of water. 

Size. The members of the order range in size from a breadth of only a few inches 
(the smallest Rajidae and Torpedo Rays) to the enormous Devil Rays that sometimes 
grow to a breadth of 22—23 ^^^^ and to a weight of more than 3,000 pounds. The Saw- 
fishes (Pristidae) reach a length of at least 20 feet and exceptionally even more. 

Locomotion?^ The Sawfishes (family Pristidae) propel themselves chiefly with the 
posterior part of the trunk by lateral undulations which are effected by waves of muscular 
contraction that progress from front to rear and thus push against the water ; this 
action is combined with lateral strokes of the caudal fin, perhaps aided also by a sculling 
action of the latter and by undulating movements of the pectorals.^^ Locomotion is 
chiefly caudal also in the Guitarfishes (Rhinobatidae), though the pectorals may be of 
some aid in their swimming." The Torpedoes swim chiefly with the posterior part of 
the trunk, aided by the caudal fin, their discs not being flexible enough to be of much 
service in locomotion. The Skates (Rajidae) are driven ahead by the simultaneous 
passage of waves of undulation through the two pectorals from front to rear. Each 
undulation originates as an elevation of the anterior margin of the pectoral, which is 
lowered as the undulation passes rearward through the fin. The undulation increases in 
amplitude until it reaches the midlevel of the disc, posterior to which it decreases in 
amplitude as it continues to progress rearward. Then a new wave arises at the front of 
the fin just before the preceding wave has entirely died away at the posterior edge. 
In one species of Skate the time occupied by an individual undulation in its progress 
from the front of the pectoral to the rear was 0.6-0.8 seconds. The Sting Rays (Dasya- 
tidae and Urolophidae), as exemplified by Urolophus, use their pectorals in the same 
way." All the batoids that do this advance by a smooth gliding motion, often right 
along the bottom. But the Eagle, Cow-nosed, and Devil Rays (Myliobatidae, Rhino- 
pteridae, Mobulidae) have been described repeatedly by competent eye-witnesses as 
progressing by flapping their pectorals, more in the manner of birds. And some of them 
have the habit of leaping into the air or of planing at the surface, a spectacular sight 
when a school breaks the surface simultaneously, as sometimes happens (p. 485). 

The Sawfishes, Guitarfishes, and Torpedoes doubtless steer from side to side, as 
do Sharks and most other fishes, by a progressive curvature caused by a wave of muscular 
contraction from the anterior part of the body toward the tail, which is then swung 

15. For a general discussion of locomotion in fishes (including elasmobranchs), see Breder (Zoologica N. Y., 4, 1926: 
159-297); for a recent analysis of the mechanics of propulsion by undulations of the trunk in various fishes, including 
Sharks, illustrated by moving picture photographs, see J. Gray (J. exp. Biol. Lond., 10, 1933: 88-104). 

16. Breder (Zoologica N. Y., 4, 1926: 243) describes the pectorals of the Sawfishes [Pristis) as "fluttering" during swim- 
ming and suggests that they may have some propulsive effect. 

17. In Rhitiohatos the pectorals "may be put to considerable use other than in steering, as may be seen upon grasping 
the tail and attempting to pull the fish out of the water" (Daniel, Elasmobranch Fishes, 1934: 13). 

18. See Mcfrey (C. R. Acad. Sci. Paris, 116, 1893: 77, 2 figs.) for analysis by moving pictures of this pectoral locomotion 
in Raja, and Daniel (Elasmobranch Fishes, 1934: 13) for Urolopkus. 



Fishes of the Western North Atlantic 9 

toward the inner side of the curve.i* Skates also appear to employ the tail to some extent 
in turning. But the chief method by which they turn, and the only method for the 
members of the order in which the tail is either very slender or very short, is by inter- 
rupting the undulations of the pectoral on the one side while those on the other side 
are continued, or by varying the rates at which the waves of undulation pass along the 
two pectorals ; the Ray swings to the right if the undulation is the more rapid on the 
left-hand side thus driving the left-hand side ahead the more rapidly, or it swings to 
the left if it is the right-hand undulation that is the more rapid. 

Breeding and Development. Fertilization is internal in all the batoids and is effected 
in the same manner as among Sharks, i.e., by a pair of appendages, known as claspers, 
that develop in the male from the inner edges of the pelvic fins. Their presence makes 
the determination of the sex easy, for they are visible even in embryos shortly before 
birth. The inner margins of the claspers are deeply grooved, with the edges more or 
less overlapping for the transmission of sperm that is received at their inner ends via 
an opening known as the apopyle. The primary support of the clasper consists of a 
single basal cartilage connected with the basypterigial cartilage of the pelvic fin by 
2-4 short intermediate pieces. During growth this basal element elongates while other 
cartilages are formed secondarily from the surrounding connective tissue. Two of these 
secondary cartilages, more or less elongate, lie alongside the primary basal element, 
with which they fuse either wholly or partially to form a rigid axial rod. As maturity 
approaches a series of additional terminal cartilages develop in varying numbers (2-5 
or 6 in different groups), often in an exceedingly complex arrangement that exhibits 
a wide range of forms, blade-like and knife-sharp in some cases though covered by a 
thin layer of integument, or emerging from the skin as thorns in other cases. Some 
of these terminal elements may be more or less erected at right angles to the general 
axis of the clasper during copulation.^" 

The members of one family (Rajidae) are oviparous ; their eggs, enclosed in horny 
capsules (p. 141), are familiar objects on the seashore. All other members of the order, 
so far as is known (with one possible exception, p. 47, fn. 1 1), are ovoviviparous, the 
embryos developing within the oviducts of the mother until they are ready for inde- 
pendent existence. But there is no placental connection between young and parent. 

The embryos of the myliobatoid Rays as a group, and of some of the Electric 
Rays, are nourished by a secretion from the vascular villi that clothe the inner wall of 
that portion of the oviduct — the so-called uterus — where the embryo lies (pp. 91, 383, 
397). But it is not known whether this is true of those rhinobatoids in which the uterine 
wall bears only a series of longitudinal folds and no villi (p. 52) or of some Torpedo 
Rays in which the structure of the uterus is similar. There is no reason to suppose that 

19. For recent discussion, with moving picture photographs showing successive positions assumed by a flexible-bodied 
Shark [Scyliorhinus caniculus) in turning sharply, see J. Gray (Proc. roy. Soc. Lond., [B] 113, 1933: 118, pi. 4). 

20. For discussion and illustrations of the skeletal elements of the claspers in representative batoids, see especially Junger- 
sen (Danish "Ingolf" Exped., 2 [2], 1898: pis. 3, 4; Anat. Anz., 14, 1898: 498-513). For detailed accounts of the 
soft parts of the claspers, see Haber (Z. wiss Zool., yo, 1901: 627-634, pi. 28); for a more recent account (with 
complex terminology for the various parts), see Leigh-Sharpe (J. Morph., 34, 1920: 260; 36, 1921: 213-218, 236- 
243; 39' '9^: 560. 568-577; 42, 1926: 318-320). 



I o Memoir Sears Foundation for Marine Research 

the embryos of any of the batoids feed on unfertilized eggs that lie close to them in 
the maternal oviduct, as do the embryos of Mackerel Sharks {Lamna) and of the Sand 
Shark {Carcharias taurus) of the western North Atlantic.^! 

Luminescence. None of the batoid fishes have luminescent organs so far as is known. 

Habits and Food. Most of the batoids are comparatively sluggish, living on the 
bottom or close to it. Even the Sawfishes (Pristidae) keep close to the bottom, except 
when they may rise among a school of fishes ; although they are slow swimmers they 
are strong, as any angler will testify who has hooked or harpooned one. The Guitar- 
fishes (Rhinobatidae and Rhynchobatidae) either swim about slowly close to the bottom 
or lie half buried. The Torpedoes (Torpedinoidea) lie buried in the mud or sand most 
of the time and swim but feebly. The Skates (Rajidae) either lie flat on the bottom, 
often with their pectorals partly buried, or swim along slowly close to it. But they can 
dart ahead with astonishing velocity if disturbed or if in pursuit of prey (p. 143). The 
Sting Rays and their kin (Dasyatidae, Urolophidae) have much this same way of life, 
except that they are rather more likely to be wholly buried except for eyes and spiracles. 
The Butterfly Rays (Gymnuridae) also hold close to the bottom, though perhaps 
moving to and fro more actively with changes of the tide than most of the dasyatids 
(p. 413). However, the Eagle (Myliobatidae) and Cow-nosed Rays (Rhinopteridae) 
are decidedly more active, for while they feed right on the bottom (pp. 437, 466) they 
often swim actively at midlevels or near the surface, sometimes leaping clear of the 
water, as described elsewhere. The Devil Rays appear to have abandoned the bottom- 
living habit largely and spend most of their time swimming close to the surface (p. 485). 

The batoids, like the Sharks, subsist wholly on animal food, and the lists of stomach 
contents so far recorded are varied enough to show that practically all of the inverte- 
brate groups characteristic of sandy or muddy bottoms contribute to the diet of one 
or another Skate or Ray. The Eagle and Cow-nosed Rays as a group subsist chiefly 
on hard-shelled mollusks, but they are known to take Crustacea on occasion (p. 473). 
The dasyatid and Butterfly Rays (Dasyatidae, Gymnuridae) are rather more catholic 
in their tastes, consuming small fish as well as Crustacea and mollusks. Skates (Rajidae) 
as a group probably depend chiefly on whatever Crustacea may be available locally, 
but they also devour mollusks, polychaete worms, and cephalopods of the less active 
sorts, as well as small fishes, on which they may feed exclusively at times (p. 223). 

The Sawfishes (Pristidae) feed chiefly on fishes (p. 19) but to some extent on 
bottom-living invertebrates as well. Some of the Electric Rays (Torpedinoidea) are 
strictly fish-eaters, sometimes devouring fishes of considerable size relative to them- 
selves, whereas others subsist on small bottom-living invertebrates. 

Most interesting of all are the feeding habits of the Devil Rays (Mobulidae). In 
spite of the great size to which some of them grow, they feed on small Crustacea, small 
fishes, and other members of the animal plankton which are directed by the fin-like 
cephalic appendages into the mouth, there to be sifted out (from the water that is taken 
in at the same time) by the so-called prebranchial apparatus, as is described on p. 483. 

21. Springer, Copeia, 1948: 154. 



Fishes of the Western North. Atlantic 



1 1 



The largest of the batoids thus feed in essentially the same manner as do the Basking 
Shark and the Whale Shark. 

Relation to Man. The Skates (Rajidae) are of considerable value to the fishermen 
of Europe, as described on p. 145. But they are in little demand elsewhere. None of the 
other batoids are of any great commercial importance, for while Rays in considerable 
variety are brought in to fish markets in tropical ports in various parts of the world, 
the number involved is small. A minor use has been made of the spines of Sting Rays 
to tip spears or to arm the lashes of whips, as described later (p. 338). The large Devil 
Rays are pursued to some extent with the harpoon as objects of sport. Otherwise, the 
chief importance of Rays to fishermen is their nuisance value, some of them because 
they take the baits intended for better fishes (p. 145) and others because of the injuries 
they are capable of inflicting by their serrate spines on anyone who may handle them 
incautiously or who may tread on them while wading on flats in regions where they 
are plentiful (p. 336). 

Habitat and Range. The latitudinal range of the batoids as a group extends from 
the Equator to the subpolar belt in both hemispheres, and their distribution is equally 
wide in all three great oceans, Atlantic, Pacific, and Indian, including the tributary 
seas. Nor can the group as a whole be described as chiefly characteristic of any par- 
ticular latitudinal zone. While the most numerous group of all (Rajidae) has reached 
its maximum development (whether as to species or as to number of individuals) in 
the temperate-boreal belts of the two hemispheres, the Sting and Butterfly Rays and their 
kin (Dasyatidae, Gymnuridae, Urolophidae), the Eagle and Cow-nosed Rays (Mylio- 
batidae, Rhinopteridae), the Devil Rays (Mobulidae), the Sawfishes (Pristidae), and 
the Guitarfishes (Rhinobatidae, Rhynchobatidae) are more numerous in tropical and 
subtropical waters, with the Electric Rays as a group occupying an intermediate position. 

A similar ecological cleavage appears with regard to the range in depth between 
the Skates (Rajidae) and certain Electric Rays on the one hand and all other batoids 
on the other. The latter, so far as is known, are most abundant by far in shoal water. 
But the Electric Rays range from close to the tideline down to at least 500 fathoms, 
while the Skates not only extend down to 1,500 fathoms but are extremely abundant 
at depths (20-100 fath.) at which the great bottom fisheries of northern Europe and 
eastern North America are chiefly carried on (p. 144). It has been discovered-^ recently 
that the bottom along the north coast of Cuba, at depths of 200-500 fathoms, sup- 
ports an abundant community of Skates, several species of which have not been found 
elsewhere. Even more recently we have found that there exist in the Gulf of Mexico, 
in depths of 50-300 fathoms, some additional species which are new to science (pp. 
286, 314). 

Consequent on their wide distribution both in latitude and depth, the batoids 
cover nearly as broad a thermal range as do the bony fishes, i. e., from the highest tem- 
peratures to which the water warms over the shallows in tropical estuaries to polar 
waters that may be as cold as -1.5° C (about 29° F). However, only one species {Raja 

2z. Bigelow and Schroeder, J. Mar. Res., 7, 1948: 543. 



1 2 Memoir Sears Foundation for Marine Research 

hyperborea, p. 206) is known to find a congenial home regularly in water colder than 
0° C (32° F), and perhaps a second species (R. spinicauda, p. 271). 

As a whole the batoids constitute a salt-water group, but several species of Sting 
Rays (family Potamotrygonidae, p. 334) have colonized fresh water in the lower reaches 
of South American rivers that drain into the Atlantic (p. 334). Sawfishes are also found 
regularly in fresh water and are even landlocked. 

Geological History. The Guitarfishes (Rhinobatidae), which are intermediate in 
form between the more highly specialized groups of Rays and the Sharks, were in 
existence in the Upper Jurassic. Thus the oldest batoids were more recent in their 
geological appearance than the oldest of the groups of Sharks that exist today, ^^ and 
they were far more recent than some extinct groups of Sharks. The Sawfishes (Pristidae), 
the Skates (Rajidae), and the Sting and Eagle Rays (Dasyatidae, Myliobatidae) ap- 
peared in the Cretaceous, the Torpedo Rays in the Eocene, and teeth identifiable as 
those of a Devil Ray, closely allied to the modern Manta, are known from the Pliocene. 2* 

Classification. Scientific opinion is now tolerably crystallized as to the basic group- 
ing of the batoids, though the taxonomic systems that have been proposed by different 
students during the last half century have differed widely as to the number of sub- 
divisions recognized and as to the names employed for the groups above the grade of 
family. 25 

Beginning with the Electric Rays: these are sharply set apart from all other 
batoids by the possession of highly developed electric organs in the anterior part of 
the body, by the greatly expanded antorbital cartilages that support the anterior margin 
of the disc, by the peculiar branched rostral cartilage or cartilages, and by the rounded 
plate-like terminations of the slender branchial cartilages. We follow common usage 

23. The Port Jackson Sharks (Heterodontidae) date back to the Lower Jurassic, the Six-gilled Sharks (Hexanchidae) 
to the Middle Jurassic. 

24. See Romer (Vert. Palaeont., 2nd ed., 1945: 577) for a list of the geological horizons from which batoid remains 
of the various families and genera have been reported. 

25. This diversity of opinion may be illustrated by the following examples: Regan (Free. zool. Soc. Lond., 1906: 723- 
724); regarding the batoids only as a suborder (Hypotremata), proposed two "Divisions," the Narcobatoidei with 
one family for the Electric Rays, the Batoidei with two families for all other batoids. But later (Encyc. Brit., 24, 
191 1 : 596) he recognized seven families. Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 257, 258), who classed 
them as a "division" of the Chondropterygia, defined six "Groups of Families" (Rhinobatoidei, Narcoidei, Raioidei, 
Dasybatoidei, Myloidei, and Mobuloidei). Jordan (Class. Fish., Stanford Univ. Publ. Biol., 3 [2], 1923: 102) di- 
vided the order Batoidei into three suborders: Sarcura with five modern families for the Saw- and Guitar-fishes 
and for the Skates, Narcaciontes with one family for the Electric Rays, and Masticura with five modern families 
for the Sting, Butterfly, Eagle, Cow-nosed, and Devil Rays. Bertin (Bull. Inst, oceanogr. Monaco, 775, 1939: 19) 
includes all modern Elasmobranchs in a single order, Euselachii, distributing the Skates, Rays, and Sawfishes among 
four suborders (Squatinorajiformes, Rajiformes, Torpediformes, and Trygoniformes) with seven families, the 
modern Sharks among seven suborders. Fowler (Bull. U. S. nat. Mus., 100 [jj], 1941: 2S9-290) has classed the 
batoids as an order (Rajae) with four primary subdivisions (Rhinobatoidei, Torpedoidei, Rajoidei, and Mylio- 
batoidei). Beebe and Tee-Van (Zoologica N. Y., 26, 1941: 245) treat the batoids as a "super order" (Platosomeae) 
consisting of two orders: Narcobatea, with one family, for the Electric Rays; and Batea, with three "superfamilies," 
Rhinobatoidea, Rajoidea, and Dasybatoidea, each having two, one, and four families respectively. Fowler has 
recently followed them in ranking the batoids as a superorder, though he has given them the name Rajioidea (Notul. 
Natur. Acad. nat. Sci. Philad., 187, 1947: 13). Finally, Berg, in his Classification of Fishes (Trav. Inst. Zool. Acad. 
Sci. URSS, 5 [2], 1940, and lithoprinted ed. 1947: 139 [Russ.], 381 [Eng.]), classes the batoids as an order, Raji- 
formes, which is divided directly into eight modern families and one fossil family, without the intermediary of any 
suborders. 



Fishes of the Western North Atlantic 



13 



in regarding them as sole members of one of the primary divisions, ranked here as a 
suborder (Torpedinoidea, p. 80). The thirty-odd species concerned have been united 
commonly in a single family, but they are distributed here among three families (for 
further discussion, see p. 86). The presence of horny rays (ceratotrichia) as well as 
of cartilaginous radials in their unpaired fins (a shark-like character) relates them more 
nearly to the rhinobatoids than to any other batoids. 

The batoids that remain after the subtraction of the Electric Rays fall in four 
fairly well definable groups of species, classed here as suborders. 




Figure 2. Pristis perotteti, juvenile male, about 1,376 mm long, from Lake Nicaragua (U. S. Nat. Mus. 
No. 120468). 



A. The Sawfishes (Pristoidea). Snout prolonged as a narrow blade-like "saw," its 
either edge armed with a single series of large tooth-like structures; tail sector stout; 
two well developed dorsal fins, the first originating over or anterior to the pelvics; a 
large caudal fin; pectoral fins attached to head only a little past the gills; outer margins 
of pelvics straight or convex; the paired fins, as well as the unpaired, with horny rays 
(ceratotrichia) in addition to the cartilaginous radials. 

B. The Guitarfishes (Rhinobatoidea). Snout not blade- or saw-like; tail sector 
stout; two well developed dorsal fins, the first originating much nearer to tips of pelvics 
than to tip of tail ; a well developed caudal fin ; pelvics with the outer margins convex, 
or at least not notched ; their unpaired fins, like those of the Sawfishes and Torpedoes, 
have horny rays as well as cartilaginous radials, as do the unpaired fins of the Sharks, 
but not their paired fins. 

C. The Skates (Rajoidea). Tail more slender and without spine; one or two small 
fins, or none; fins with cartilaginous radials only (no horny rays); the origin of first 
dorsal (if any) much nearer to tip of tail than to tips of pelvics; caudal fin reduced to 
a low fold finely striate in structure; the outer margins of pelvic fins more or less 
deeply concave. This group is typified by the common Skates. 

D. The Rays (Myliobatoidea). Tail ranging from moderately to excessively slen- 
der; one dorsal fin or none; dorsal fin, if present, well forward on the tail, anterior to 
tips of pelvics or close behind them; pelvics with outer margins not concave; some 



1 4 Memoir Sears Foundation for Marine Research 

species with a well developed caudal fin (with cartilaginous radials) but others without 
one; fins without horny rays (ceratotrichia) ; in most species a strong serrated tail spine 
(or spines). This group includes the so-called Sting and Whip Rays, Butterfly Rays, 
Eagle Rays, and Devil Rays. 

The dividing lines that separate the torpedinoids, the rhinobatoids, and the 
pristoids from the other suborders are sharp. But the gap between the rajoids and 
myliobatoids is partially bridged by one group, Anacanthobatidae (p. 327), in which 
the margins of the pelvics are as deeply notched as in most of the Rajoidea but in 
which the tail is diagnostic in that it is whip-like, without trace of a dorsal fin. It is 
classed here among the Rajoidea, but as a separate family (p. 327) because of its pelvics. 

Number of Genera and Species. On the basis of present knowledge, the living 
representatives of the five suborders of batoids appear to be divisible into 16 families 
and about 47 genera. At present about 300—340 recognizable species have been de- 
scribed. A critical comparison of representatives of closely allied forms from different 
ocean areas doubtless will result in some condensation. But our own experience with 
the genera Breviraja (p. 284) and Cruriraja (p. 313) suggests that any reductions so 
caused will be more than counterbalanced by the discovery of new species from regions 
where the batoid fauna has been studied only casually, especially from the deeper levels 
along the continental slopes. 

Key to Suborders 

I a. Snout much prolonged as a flat narrow blade, its either edge armed with a single 

series of large tooth-like structures; radial cartilages of pectoral and pelvic fins 

supplemented distally by much more numerous fine horny rays, their inner ends 

embracing the outer ends of the cartilaginous radials. Pristoidea, p. 15. 

lb. Snout not prolonged as a blade-like structure, its edges without teeth. 

2 a. Cranial support of anterior margin of disc consisting of a preorbital cartilage 
on either side, expanded forward, variously branched or reticulate (easily felt 
even in large specimens, though not visible externally), with one or two 
rostral elements also branched in some; skin of disc as well as of tail wholly 
naked in most species;^* a highly developed electric organ on either side 
between head and forward extension of pectoral, often visible externally; tips 
of branchial rays expanded as rounded plates. Torpedinoidea, p. 80. 

2 b. A single rostral cartilage as a cranial support for anterior margin of head 
either present or not; electric organs, if any, rudimentary, on tail; skin in most 
species with scales, thorns, or spines; tips of branchial rays but little ex- 
panded, if at all. 

3 a. Tail sector so stout that it is not marked off definitely from body sec- 
tor; dorsal and caudal fins well developed, supported distally by horny 
rays, basally by the cartilaginous radials. Rhinobatoidea, p. 43. 

26. Described as having partly spinous papillae in one species (p. 92). 



Fishes of the Western North Atlantic i 5 

3 b. Tail sector slender, extremely so In many cases, and sharply marked 
off from body sector; dorsal and caudal fins, if any, not supported dis- 
tally by horny rays. 

4a. Pelvis with prepelvic spur at outer corners; outer margin of pelvics 
concave in most; spiracles with traces of gill folds. Rajoidea. p. 132. 
4b. Pelvis without spur at outer corners; outer margin of pelvics straight 
or convex; spiracles without trace of gill folds. 

Myliobatoidea, p. 331. 

Sequence of Presentation. The batoids as a whole constitute a highly specialized 
group, and the divergences between their major subdivisions represent still further 
specializations In one direction or another beyond their common divergence from the 
more generalized elasmobranch stem, as represented by the Sharks. Consequently It 
is not possible to represent the mutual interrelationships between the several suborders 
by any arrangement of them in linear series. Of the batoids, the Sawfishes (Pristoidea) 
are the most nearly shark-like in form of body; also, their fins, paired as well as un- 
paired, have horny rays (ceratotrichia) in addition to cartilaginous radlals, as do the fins 
of Sharks. Although the saw of the Sawfish represents extreme specialization in one 
direction, this is almost exactly paralleled in the Saw Sharks (Pristiophoroldea). We, 
therefore, follow common usage In commencing this account with the Pristoidea, 
leading to the Rhinobatoidea. Also, it is customary to place the Myliobatoidea after 
the Rajoidea since they represent the extreme development of batoid specialization In 
general, and nothing would be gained by doing otherwise. Although the Electric Rays 
have been placed between the rhinobatoids and the rajoids,^' between the rajoids and 
the myliobatoids,-^ or after the myliobatolds,^'' we place them immediately after the 
rhinobatoids, thinking their well developed unpaired fins, with horny rays in addition 
to the cartilaginous radlals, more significant from the standpoint of relationships than 
the extreme specialization of their electric organs or the great development of their 
antorbltal cartilages. 

Suborder PRISTOIDEA 
Sawfishes 

Characters. General form more shark-like than ray- or skate-like, but with trunk 
and head flattened ventrally; the head especially flattened, its anterior part so strongly 
depressed as to be very thin dorsoventrally. No evident demarkatlon between body 
and tail sectors of trunk (Fig. 2). Snout extended as a long narrow flattened blade 
(the so-called "saw") armed along either edge with a single series of transverse teeth 

27. See especially Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 257), Rey (Fauna Iberica, Feces, i, 1928: 515), 
and Fowler (Bull. U. S. nat. Mus., 100 [13], 1941: 332)- 

28. Jordan, Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 27; Benin, Bull. Inst, oceanogr. Monaco, 

775' 1939- 

29. Berg, Classification of Fishes, Trav. Inst. Zool. Acad. Sci. URSS, 5 (2), 1940, and lithoprinted ed. 1947: 139 (Russ.), 
381 (Eng.); Romer, Vert. Palaeont., 2nd ed., 1945: 577. 



1 6 Memoir Sears Foundation for Marine Research 

with persistently growing bases, each tooth representing a single enlarged dermal 
denticle;^" these teeth are homologous not only with the rostral teeth of the Saw Sharks 
(Pristiophoroidea) but with the dorsal fin spines of Heterodontus and of some squaloid 
Sharks as well ; rostral teeth deeply and firmly embedded in deep sockets of the hardened 
cartilage of the saw in all modern Sawfishes ; however, in some fossil forms (genera 
Propristis, Sclerorynchus) they were not socketed but were seemingly attached to the 
skin only. The saw of the Sawfishes, while resembling that of the Saw Sharks (Pristio- 
phoroidea) very closely in general appearance, does not bear the lateral cirri characteristic 
of the saw of the Saw Sharks. Anterior parts of pectorals fused with sides of head for- 
ward past all five gill openings and at a higher level, but not so far forward as the 
mouth; posterior corners of pectoral fins terminating considerably anterior to origin 
of pelvics. Origin of first dorsal varying from considerably anterior to origin of pelvics 
to a little posterior to rear ends of bases of latter. Second dorsal separated from first by 
a considerable interspace. Caudal well developed, either with or without definite lower 
lobe, its axis somewhat raised, with termination directed toward the rear margin of 
fin some distance below upper tip. 

Eye with a rounded velum; orbit outlined below by a deep semilunar fur- 
row, the skin of its lower half loose, described as acting like a nictitating (winking) 
membrane. Spiracles some distance posterior to eyes, oblique, their outer ends directed 
rearward (the reverse of what is usual among the batoids), their posterior margins 
without ridges or folds. Nostrils entirely separate from mouth and far removed from 
mouth, also from each other in the midzone; anterior margin expanded in a well marked 
lobe with rounded tip. Mouth transverse, nearly straight; skin at its corners wrinkled, 
but without well defined pits or furrows. Gill openings wholly on lower surface. Oral 
teeth small, rounded, close-set in quincunx arrangement and very numerous,'^ with 
several series in function simultaneously; tooth bands ovate to arcuate in cross section 
(Fig.4K). 

Dermal denticles over body as a whole minute, flat, ovoid, comparatively uniform, 
clothing the skin closely, including the saw; without larger thorns, tubercles, or spines. 

The saw is supported by an elongate rostral projection from the cranium. ^^ This 
cartilage encloses 3—5 longitudinal canals as well as other spaces toward its base, and 
it is strengthened by various calcifications, the entire outer layer immediately next to 
the integument being hardened in this way. Although the antorbital cartilages do not 
extend forward beyond the level of the nostrils, the anterior end of each is connected 
with the rostral cartilage by a ligamentous band.^^ Radial cartilages of paired as well 
as unpaired fins supplemented by much more numerous fine horny rays, the inner 

30. For a detailed account of the histology and development of the rostral teeth, with references to earlier literature, 
see especially Engel (Zool. Jb., Anat. Abt. 29, 1909: 51-100, pis. 3-6). 

31. 70-178 series above and nearly as many below in the different species and at different stages in growth. 

32. According to Gegenbaur (Unters. Vergl. Anat. Wirbelt., 3, 1872: 91, pi. 9, fig. 7, 8), who has given the most de- 
tailed account with which we are acquainted, this rostral extension involves a single cartilage corresponding to the 
single rostral cartilage of the Rhinobatoidea, not three as had been believed earlier. 

33. For a general account of the head skeleton and for a comparison with other groups of batoids, see Holmgren (Acta 
Zool. Stockh., 22, 1941: 53, 64). 



Fishes of the Western North Atlantic 1 7 

ends embracing the outer ends of the cartilaginous radials. Propterygial cartilages of 
pectorals as well developed as metapterygials. Pelvis convex in front but without pro- 
cesses directed forward from outer ends.^ Surfaces of gill arches smooth inward from 
gill filaments. 

Remarks. Attention has been called repeatedly to the shark-like features of the 
Sawfishes, i. e., to their elongated bodies, their powerful tails and caudal fins, and their 
comparatively small pectorals. Also, their mode of locomotion is shark-like, for they 
propel themselves through the water by powerful strokes of the rear part of the trunk 
and caudal fin, seemingly using the pectorals for steering and especially for directing 
their course upward toward the surface or downward toward the bottom. All their 
fins, like those of Sharks, have horny rays as well as cartilaginous radials. But their 
underlying structure is batoid — the fusion of their pectorals with sides of the head 
past the gill openings, the lack of free eyelids, the union of several of their anterior 
vertebrae which bear a wide lateral wing on either side in some, the presence of a 
scapular cartilage connecting the two ends of the pectoral girdle across the upper side 
of the vertebral column to which it is bound firmly by ligamentous tissue, the presence 
of many more radial cartilages on the propterygial cartilage of the pectoral fins than 
on the mesopterygial cartilage, and the fact that the lower jaw has no direct connec- 
tion with the ceratohyal cartilage.''^ 

The pristoids, in short, are true batoids, just as the squatinoids are true Sharks 
though batoid in general appearance. The saw of the Sawfish is a remarkable develop- 
ment, however, for its only counterpart in the entire vertebrate series is the similar 
structure developed by the Saw Sharks (Pristiophoroidea). 

Development. Ovoviviparous; in at least one species there are several embryos 
within a single egg capsule in each uterus.^* The rostral blade, at first more or less 
soft and flexible, is developed during the later embryonic stages; it has been known 
for more than two centuries that its lateral teeth do not project through the enclosing 
integument until after the young are set free,^' otherwise it would hardly be possible 
for the mother to give birth to them. But the saw becomes calcified and the teeth grow 
very rapidly after the young commence their independent existence. ^^ For example, 
in a 675 mm specimen of Pristis pectinatus (newborn to judge from its umbilical scar) 
the blade is already rigid, with the tips of the teeth slightly projecting, while in a P. 
perotteti of 865 mm (Fig. 6 A) the teeth are already about as long, relatively, as they 
are in the adult of that species. 

Families and Genera. The modern members of the suborder resemble one another 
so closely that they are all referred by common consent to the Pristidae. 

34. For further skeletal details, see Garman (Mem. Harv. Mus. comp. Zool., j6, 191 3: pi. 55, fig. 3; pi. 64> figs. 2-3)- 

35. For excellent illustrations of these skeletal characters, which we have verified by dissection, see Garman (Mem. 
Harv. Mus. comp. Zool., j6, 191 3: pi. 55, fig. 3, pi. 64, figs. 2, 3). 

36. In a specimen of Pristis cuspidatus each uterus had a single egg capsule divided into four compartments, each with 
one embryo. See Setna and Sarangdhar [Rec. Indian Mus. (194S), 46 (1-4), 1949: u, fig- 4]- 

37. "In the embryo state the sides of the snout are as smooth as the gums of a new-born infant" (Latham, Trans. Linn. 
Soc. Lend., Zool. 2, 1794: 274; transl. from Klein, Hist. Pise. Natural., Missus 3, 1743: 12). 

38. This seems to have been remarked first by Latham (Trans. Linn. Soc. Lond., Zool. 2, 1794: 274). 



I 8 Memoir Sears Foundation for Marine Research 

Family PRISTIDAE 
Sawfishes 

Characters. Those of the suborder. 

Genera. All modern Sawfishes appear to fall within a single genus, Pristis. At 
most, the species in which a lower caudal lobe is developed have been given a separate 
subgeneric name.^* The fossil pristids exhibit so wide a range of variation regarding 
calcification of the rostral blade and the implantation and other characters of the rostral 
teeth that a recent enumeration" lists not less than 12 genera. But it is likely that the 
number of these would be reduced considerably were material available for as critical 
a comparison of them as can be made for the modern Sawfishes. 



Genus Pristis Link 1790 

Sawfishes 

Pristis Link, Mag. Physik. Naturg. Gotha, 6 (3), 1790: 31 ; Latham, Trans. Linn. Soc. Lond., Zool. 2, 1794: 
276; type species, Squalus pristis Linnaeus 1758. 

Generic Synonyms.*^ 

Pristibatus Blainville, Bull. Soc. philom. Paris (181 6), 1816: 121; type species, Pristis antiquorum Latham 
1794, equals Squalus pristis Linnaeus 1758, designated by Fowler (Bull. geol. Surv. N. J., 4, 191 1 : 81). 

Pristibatys Blainville, Nouv. Diet. Hist, nat., ed. 2, 27, 1818: 385; emended spelling for Pristibatus Blain- 
ville 1 8 16. 

Pristibatis Blainville, in Vieillot, Faune Frani;., 1825: 49; emended spelling for Pristibatus Blainville 18 16. 

Myriosteon Gray, Proc. zool. Soc. Lend., 1864: 163, 164; type M./iigginsii Gray.*^ 

Pristiopsis Fowler, Proc. Acad. nat. Sci. Philad., 57, 1905: 459; type species, Pristis perotteti Miiller and 
Henle 1841;*^ subgenus for species with caudal having distinct lower lobe. 

Generic Characters. Rostral teeth deeply embedded in calcified sockets,** nearly 
straight or slightly recurved and strongly compressed dorsoventrally, their tips sharp in 
newborn specimens but slightly blunted in older ones; both edges (anterior and 
posterior) sharp in young specimens, but the posterior edge more or less flattened, 
evidently by wear, in large ones; neither surface channeled, though both surfaces 
(dorsal and ventral) may be faintly striate toward the base. Each side of tail with a 
low longitudinal ridge. Dorsal fins with free rear corners considerably extended, as 
in many Sharks; the pectorals likewise. Caudal with or without a definitely outlined 

39. Pristiopsis Fowler (Proc. Acad. nat. Sci. Philad., 57, 1905: 459). 

40. Romer, Vert. Palaeont., 2nd ed., 1945: 577. 

41. Fowler (Bull. U. S. nat. Mus., 100 [ij], 1941: 290) includes among the synonyms of Pristis the fossil Eopristis 
Stromer (Beitr. Palaont. Geol. Ost. Ung., 18, 1905: 52). 

42. Based on a fragment that Gray thought might "indicate a new group of radiated animals nearly allied to Asterias"; 
but later it was identified by Giinther (Cat. Fish. Brit. Mus., S, 1870: 436, footnote) as one of the lateral tubes of 
the saw of a Pristis that had become detached in some way. 

43. The date of publication of Fowler's Pristiopsis for a Sawfish was August 14, 1905. Thus it antedates Pristiopsis, 
proposed by Schmidt (Stettin, ento. Ztg., 66 Jahrg., heft 2, 1905: 332) for Coleoptera, the date of publication being 
November, as printed on p. 384 of the stated publication. 

44. This contrasts with their looser attachment to the blade in some fossil Sawfishes (p. 18). 



Fishes of the Western North Atlantic 19 

lower lobe. Dermal denticles covering integument of saw similar to those on trunk. 
Characters otherwise those of the family. 

Size. The ancients credited the Sawfishes with sizes equal to those of the Whale- 
bone Whales, with which they were sometimes confused — for what reason one can 
hardly guess. ^^ However, the ichthyologists of the eighteenth century were well aware 
that such stories had not the faintest justification. Sawfishes are among the larger of 
elasmobranchs, commonly growing to 15—16 feet in length; the larger species are 
reported as reaching 20-24 feet both in Indian and Australian waters''^ and in the 
Atlantic (p. 39), and we read that "individuals of 30 feet are sometimes encountered."*' 

When comparing the weights of Sawfishes with those of Sharks it is to be remem- 
bered that approximately V4~V3 of "^^e over-all length of the former consists of the 
saw, which adds but little to the weight, while the head as a whole is so flattened dorso- 
ventrally that it contributes much less to the weight than do the heads of most Sharks. 
On the other hand, the caudal fin occupies a smaller percentage of the total length than 
is the case in many Sharks. Thus, while a Sawfish probably would not weigh 600 pounds 
at less than about 14 feet, a Maneater {Carcharodon) might be expected to attain that 
weight at about eight feet, a Mako (Isurus oxyrinchus) at about nine feet, or a Green- 
land Shark (Somniosus) at about 1 1 feet. A Sawfish 1 7 feet 4V2 inches long weighed 
1,300 pounds.** The heaviest of which we have found record, a West Indian female 
whose length was not recorded, was estimated to weigh 5,300 pounds.** 

Habits and Food. Sawfishes are found in most warm seas and are plentiful locally. 
They live chiefly on bottom in shallow water where it is sandy or muddy, often close 
to the shore and perhaps seldom descending to a depth greater than five or six fathoms. 
They are most plentiful in sheltered bays or in estuarine situations, often in brackish 
water. They not only ascend far above tidal limits in large rivers in many parts of the 
world, but they are among the few elasmobranchs that are found regularly in fresh 
water, as in Lake Nicaragua where they appear to be landlocked (pp. 39, 40). 

Sawfishes subsist chiefly on whatever small schooling fishes may be abundant 
locally, such as mullets and the smaller members of the herring tribe; they also feed 
to some extent on Crustacea and other bottom-dwelling inhabitants. It is for the prosaic 
purpose of grubbing in the sand or mud in search of whatever prey they can uncover 
that the saw is chiefly used, the tips and posterior edges of its teeth often being more 
or less worn down in this way. They also use the saw to slash to and fro in schools of 
fishes, thus killing or stunning the victims, much as the swordfishes, sailfishes, and 
marlins use their swords. The sensational accounts of them as habitually attacking 

45. Thus Pliny, according to Latham (Trans. Linn. Soc. Lond., 1794: 275), speaks of "Pristes" of 200 cubits length 
in the Indian Ocean. 

46. Whitley, Aust. Mus. Mag., 3, 1927: 23; Fish. Aust., i, 1940: 178. 

47. Norman and Fraser (Giant Fishes, 1937: 60). An account has recently appeared of the capture in Panama Bay of 
a Sawfish said to have been 27 feet long (Mitchell-Hedges, Battles with monsters of the sea, 1937: 224-225). But 
its saw was described as being only a little more than five feet long, suggesting that the total length of the fish was 
overstated. 

48. Baughman (Copeia, 1943: 44), a Pristis perotteti from Texas. 

49. Norman and Fraser, Giant Fishes, 1937: 63. 



2 o Memoir Sears Foundation for Marine Research 

whales have no basis. ^^ Neither do we place any credence in reports that they customarily 
cut out and devour chunks of flesh from large fishes. 

However, a large Sawfish does have a powerful weapon of defense — and potentially 
of offense — at its disposal in its saw, for it can strike sideways with the latter with great 
power, either in the water, as when being hauled alongside a boat, or on land if drawn 
up on shore. Consequently, even small ones should be handled or approached with 
caution, as every fisherman knows who is familiar with them. Since they may strike 
in the same way when lying on the bottom if disturbed, stories of injuries inflicted by 
them on fishermen or on bathers, or even of fatalities, cannot be brushed aside as 
imaginary.^i But we know of no evidence of an unprovoked attack upon a bather in 
any part of the world by a Sawfish that had not been disturbed in some way. 

Relation to Man. Sawfish flesh is coarse and, so far as we can learn, no regular 
use is made of it along the western coasts of the Atlantic except that one is occasionally 
exposed for sale in some tropical fish market. Indeed, it has been listed as poisonous 
in Cuba.52 But small ones of three feet or so are described as "delicious as a breakfast 
pan fish. "^3 And we read that in India "the flesh is equally esteemed with that of sharks. 
The fins are prepared and sent to China; the oil is extracted from their livers, whilst 
the skins are useful for sword scabbards or for smoothing down wood."^* Sawfish saws 
are often offered for sale as curios in tropical ports, and they are still employed in 
religious ceremonies by the Australian natives. The skin is used to some extent for 
leather. But no attempt seems to have been made to obtain their skins or oil on a 
large scale, though a considerable yield of the latter per fish might be expected, 
judging from a recorded weight of 102^/4 pounds for a single liver (length of fish not 
recorded).^* 

Sawfishes are too sluggish to be held in any regard as game fish by anglers. But 
they take cut fish-bait rather readily if they detect it (no doubt by scent). Once hooked 
they swim so powerfully, though slowly, and are so enduring, that the capture of a 
large one entails a long and often wearisome struggle. Also, it is not unusual for a 
Sawfish to leap clear of the water under such circumstances, as happened on one occasion 
when it required more than two hours for one of us to subdue a 14-foot Sawfish on 
a handline from a small boat. 

Range. Tropical to subtropical coastlines, estuaries, and river mouths of all three 
great oceans, running up to fresh water in many localities. In the western Atlantic 
Ocean Sawfishes range northward regularly to the West Indies and Florida, seasonally 

50. According to Lacep^de (Hist. Nat. Poiss., 4° Edit., in Buffon, Hist. Nat., J, 1798: 292), there was in the Paris Museum 
part of a Sawfish saw that was said to have been taken from the side of a Whale. But there is no way of verifying 
this supposed origin. 

51. A report received by Day (Fish. India, 1878: 728) of a bather in India being cut in two by a large one is not beyond 
the bounds of credence. 

52. Hoffman, Rev. chil. Hist. Nat., 33, 1929: 29. 

53. Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 28. 54. Day, Fish. India, 1878: 729. 

55. See Hooper (Mem. Indian Mus., 2, 1909: 59-60), who also gives chemical analyses of the oil of Indian Sawfishes, 
mixed, however, with that of Sting Rays and Sharks; see Marcelet (Bull. Inst, oceanogr. Monaco, 817, 1942: 6) 
for analysis of liver oil from a Madagascar specimen. 



Fishes of the JVestern North Atlantic 2 i 

to the Carolinas, occasionally to Chesapeake Bay, and accidentally to New Jersey; in 
the eastern Atlantic they occasionally range to the Mediterranean and as strays to 
Portugal. To the southward they occur to southern Brazil and northern Argentina in 
one side of the ocean and along tropical West Africa to the Cape of Good Hope in 
the other.'^' Their range elsewhere is extensive also: in the eastern Pacific from northern 
Mexico to Ecuador; in the west from Indo-China to Queensland and south seasonally 
to New South Wales (Sydney), including the East Indies; the tropical coasts of the 
Indian Ocean as a whole, including the Arabian Sea, south to the Islands of Reunion 
and Madagascar and to Natal; and the Red Sea. It is probable that the failure of 
scientific literature to report their presence for any of the island groups of the western 
tropical Pacific chiefly reflects the imperfection of the published record. 

Species. The species of Pristis are separable into two groups according to whether 
the caudal has a distinct lower lobe or not. This distinction is used, in fact, as the basis 
for subgeneric separation in the most recent synopsis of the Sawfishes of the western 
Pacific-Indian Ocean. 5' The positions of the first dorsal relative to the pelvics and 
of the second dorsal relative to the caudal are characters of specific value. The num- 
ber of rostral teeth, a character that is fixed before birth, also differs in different 
species. 

The group in which the caudal fin has a lower lobe is represented in the western 
Atlantic by one species with 16—19 pairs of rostral teeth; this appears to be identical 
with the Sawfish described as Pristis perotteti from Senegal, West Africa, in 1841 by 
Miiller and Henle. P. zephyreus Jordan and Starks 1895 of the Pacific Coast of Central 
America appears not to be separable from it, except perhaps by a slightly larger average 
number of teeth (p. 41). And P. perotteti is closely allied to the form that has been 
reported repeatedly as P. microdon Latham 1794 from various localities in Australian 
waters, from Indo-China, from the East Indies, from the Indian Ocean, and recently 
from India as P. perotteti}^ The lower caudal lobe is still more prominent in P. cus- 
pidatus of the western Pacific and Indian oceans; and P . leichhardti Whitley 1945,^' 
recently described from Australia, also has a distinct lower caudal lobe. 

The group lacking a lower caudal lobe is similarly represented in the western 
side of the Atlantic (so far as is known) by only a single species. This has usually been 
referred to P . pectinatus Latham 1794 because of its numerous rostral teeth (25— 32 pairs) 
and it is so named here. A form resembling it closely in number of teeth and in relative 
positions of fins has been reported under this same name from the Pacific Coast of 
Central America, from South Africa, from the Indian Ocean and Red Sea, and from 
the Philippines. Here again the true relationship of the Atlantic to the Indian Ocean 
form remains to be established. 

It appears that the Atlantic Sawfish which Linnaeus, in 1758, named Squalus 
pristis (type species of the genus) also lacks a lower caudal lobe, for the illustration on 

56. Gunther (Cat. Fish. Brit. Mus., S, 1870: 438) lists one from the Cape. 

57. Fowler, Bull. U.S. nat. Mus., 100 (rj), 1941: 291. 

58. By Prater, J.Bombay nat. Hist. Soc, 41, 1939: 435. 59- Aust. Zool., 11 (i), 1945: 44- 



2 2 Memoir Sears Foundation for Marine Research 

which he based the species shows none."" But its saw is credited in the original account 
with having only 16—20 teeth. 

An adequate description of P. pristis is much to be desired, because this is the 
species, as type of the genus, with which all other members of Prhtis must be com- 
pared eventually. To date the only definite locality records of specimens positively 
referable to it are for Senegal, «! the Mediterranean (where it appears to have been con- 
fused with other Sawfishes), and perhaps Portugal. But watch should be kept for it 
in the American side of the tropical Atlantic. Two other members of the group without 
lower caudal lobe are known from the western Pacific-Indian Ocean area, P. zijsron 
Bleeker i 85 1 and P. clavata Garman 1 9 1 3 ; both of them are separable from P.pectinatus, 
from P. pristis, and from each other, by the characters listed in the accompanying Key. 

Key to Species 

I a. Caudal fin with a definite lower lobe. 

2 a. Origin of first dorsal posterior to origin of pelvics; subcaudal lobe prominent. 

cuspidatus Latham 1794. 

Southeastern Asia, East Indies, 

Ceylon, and India to Red Sea. 

2b. Origin of first dorsal considerably anterior to origin of pelvics; subcaudal 

lobe small (Fig. 5). 

3 a. Rear tip of second dorsal separated from origin of caudal by an inter- 
space about half as long as base of second dorsal. 

perotteti Miiller and Henle 1841, 

P-34- 

Also zephyreus Jordan and Starks 1895, 
from the Pacific Coast of Central America, 
and microdon Latham 1 794, from the West 
Pacific-Indian Ocean. *^ 

3 b. Rear tip of second dorsal reaching to origin of caudal. 

lekhhardti Whitley 1945. 
Queensland, Australia (in rivers). 
I b. Caudal fin without definite lower lobe. 

4 a. Origin of first dorsal over origin of pelvics, or anterior to latter. 
5a. 25-32 pairs of rostral teeth, pectinatus Latham 1794, p. 23. 
5b. 16-20 pairs of rostral teeth. pristis Linnaeus 1758. 

Eastern tropical Atlantic, ac- 
cidental in Mediterranean 
and perhaps to Portugal. 

4 b. Origin of first dorsal clearly posterior to origin of pelvics. 

6a. 25-32 pairs of rostral teeth; outer corners of pectorals 

60. LeCluse (Exoticorum libri decern . . . Animal. Plant . . . Petri Belloni, obs , 1605: 136); locality "in Oceano 

Occiduo." 

61. Rochebrune, Act. Soc. linn. Bordeaux, (2) 6, 1882: 49, as P. antiquorum Latham 1794. 

62. See discussion, p. 21. 



Fishes of the lVeste7~n North Atlantic 2 3 

rather broadly rounded; origin of first dorsal clearly pos- 
terior to midpoints of bases of pelvics; a large species, 
growing to upwards of 20 feet in length. 

zijsron Bleeker 1851. 
Cochin-China, Australia, East 
Indies, Ceylon, India, Gulf 
of Oman. 
6b. 18-21 pairs of rostral teeth; outer corners of pectorals sub- 
angular, about a right angle; origin of first dorsal anterior 
to midpoints of bases of pelvics ; said to be a small species, 
growing to about 4^/2 feet. clavata Garman 1906. 

Queensland, Australia; also 
reported from the Canaries.*^ 

Pristis pectinatus Latham 1794 

Common Sawfish 

Figures 3, 4 

Study Material. Immature male, 1,396 mm long, from off Galveston, Texas; six 
small specimens, male and female, 960—1,040 mm long, from the vicinity of Galveston, 
Texas; and one newborn male, 675 mm long, from Mobile, Alabama; also a saw from 
a 14-foot specimen (sex not recorded) from southern Florida, and a saw about three 
feet long from a female more than 12 feet long from Pascagoula Bay, Louisiana; all 
in the Museum of Comparative Zoology; female about 3 feet 7 inches long from the 
Indian River, Florida, in Chicago Museum of Natural History. 

Distinctive Characters. P. pectinatus is easily separable from the only other Sawfish 
{P . perottet'i) known from the western side of the Atlantic by the following facts: its 
first dorsal fin originates about over the origin of its pelvics (considerably in front of 
origin of pelvics in P. perottet'i) ; its caudal is much shorter, but broader relative to the 
length of the fin, with lower lobe only faintly indicated (cf. Fig. 3 with 5); its rostral 
teeth are more numerous, 24 or more on each side (not more than 1 9 or 20 in P. perotteti, 
but see p. 43, fn. 100); its saw is relatively somewhat shorter; its second dorsal has 
the posterior margin much less deeply concave; and its pectorals are smaller. It agrees 
with P. pristis of the eastern Atlantic in the shape of its caudal and in the position of 
the first dorsal fin relative to the pelvics, but it is separable from P. pristis by its more 
numerous rostral teeth (see Key, p. 22). 

Description. Proportional dimensions in per cent of total length. Female, 1,017 "^n^> 

63. Pristis clanjata closely resembles P. zijsron, but the differences in number of rostral teeth, in the shape of the pec- 
torals, and in the position of the first dorsal relative to the pelvics seem sufficient for specific recognition. We should 
point out, however, that Carman's (Bull. Mus. comp. Zool. Harv., 46, 1906: 20S; Mem. Harv. Mus. comp. Zool., 
36, 191 3: 264) characterization of its second dorsal as smaller than its first might be misleading, for the linear di- 
mensions of the two fins of the type specimen, which we have examined, are in the proportion of only about i.i to 
I. It has been reported from Australia by Whitley (Fish. Aust., i, 1940: 178) also; and from the Canaries by Bel- 
Ion and Mateu (Notas Inst. esp. Oceanogr., [2] 53, 193 1: 28) from a dried specimen with 21 saw teeth. 




Figure 3. Pristis pectinatus, juvenile male, about 1,396 mm long, from off Galveston, Texas (Harv. Mus. Comp. 
ZooL, No. 36659). A Cross section of caudal peduncle close in front of caudal fin, about 0.35 X. 



Fishes of the JVestern North Atlantic 25 

from Galveston, Texas (Ilarv. Mus. Comp. Zool., No. 36960). Male, 1,396 mm, from 
Galveston, Texas (Harv. Mus. Comp. Zool., No. 36659). 

Breadth: between outer corners of pectorals 26.5, 26.6. 

Snout length: in front of orbits 27.4, 25.4; in front of mouth 31.2, 28.9. 

Orbits: horizontal diameter 1.9, 1.8; distance between 4.9, 4.7. 

Spiracles: length 1.9, 1.6; distance between 4.0, 4.2. 

Mouth: breadth 5.4, 5.1. 

Nostrils: distance between inner ends 2.2, 2.0. 

Gill openings: lengths, ist 1.5, 1.6; 3rd 1.6, 1.7; 5th 1.2, i.i ; distance between 
inner ends, ist 9.8, lo.i; 5th 7.2, 7.4. 

First dorsal fin: vertical height 6.8, 6.8; length of base 7.0, 7.1. 

Second dorsal fin: vertical height 6.8, 7.6; length of base 6.0, 6.4. 

Caudal fin: upper anterior margin 15.0, 15.0; lower anterior margin 8.7, 8.7. 

Pelvics: anterior margin 7.2, 7.5. 

Distance from tip of snout to: ist dorsal 58.9, 55.8; pelvics 58.8, 55.3; center 

of cloaca 62.0, 59.6; from center of cloaca to lower caudal 24.9, 26.5. 

Trunk tapering nearly evenly rearward from level of axils of pectorals; flattened 
below; its height a little less than its width at origin of first dorsal where highest; its 
breadth at origin of pectorals between ^4 and Vs as great as its length from base of saw 
to origin of caudal ; a low fleshy ridge low down along either side from about opposite 
origin of second dorsal to opposite anterior part of caudal axis. Pectorals with nearly 
straight margins, narrowly rounded outer corners, and slightly blunted posterior cor- 
ners, their anterior margins about as long as distal margins and directed outward 
much less abruptly from sides of head than in P. perotteti (cf. Fig. 3 with 5). Extreme 
width across pectorals about equal to distance from mouth to level of rear corners of 
pectorals, therefore considerably less, relatively, than in P. perotteti (p. 36). Caudal 
peduncle, at upper origin of caudal, about 1.5 times as wide as deep. 

Minute dermal denticles closely covering skin everywhere (a shark-like character); 
those on upper surface of trunk and fins blunt ovate, varying in size one to the next, 
averaging smallest on outer parts of fins and on head anterior to eyes; blades nearly 
horizontal so that skin is only slightly rough to the touch; pedicels low. Denticles on 
lower surface smaller, varying considerably in size and in shape from roughly circular 
to blunt ovate or subpolygonal with rounded corners; without definite pedicels; so 
closely crowded that skin is visible only here and there; and lying so flat as to be 
smoother to the touch than those on upper surface. Denticles along midzone of saw of 
about same size as those on lower surface of trunk, but those along its edges con- 
siderably larger; extreme margin of saw naked in newborn specimens but completely 
covered with denticles by the time a length of about 1,400 mm or so is reached, if 
not sooner. 

Saw about V4 of total length in grown specimens; about Vt-Vs as wide at base 
as long, narrowing evenly forward to only a little more than half (about 0.6) as wide 



26 



Memoir Sears Foundation for Marine Research 



at tip as at base; weakly rounded above as well as below, and about Vs ^s thick as 
broad; its tip rounded-truncate. 

Rostral teeth 24—32 on either side, often one more tooth on one side than on the 



/UIJUMiiMAliliyjUXLL/UU/ULA 








Figure 4. Pristis pectinatus. A Saw from specimen about 14 feet long, from southern Florida (Harv. Mus. Comp. 
ZooL, No. 1220). 5 Two saw teeth from same, about 0.6 X- C Saw tooth from same, seen obliquely, about 1.2 X. 
D Margin of saw of same to show dermal denticles, about 5 X. £ Eye and spiracle of juvenile male, 1,396 mm 
long, from off Galveston, Texas (Harv. Mus. Comp. Zool., No. 36659), about 0.4X. F Lower surface of head 
of same, 0.4 x . G Dermal denticles of same from below first dorsal fin, about 28 x . // Dermal denticles from same 
in lateral and rear views, about 28 x . ^ Right-hand half of upper and lower jaws to show tooth bands, about 
1.8 X- K Transverse section of upper jaw to show transition from dermal denticles on upper lip (smaller) to 
teeth (larger), with lateral aspect of latter, about 7 X . i Teeth from central part of upper jaw, with dermal 
denticles on upper lip, about yx. 



Fishes of the IV es tern North Atlantic 27 

other j"^ final number of teeth established at birth or soon thereafter; saw teeth spaced 
c,—() times as far apart near base of saw as toward tip but with the decrease in spacing 
somewhat irregular ;8^ the basal pair separated from the apparent level of emergence 
of the saw by a distance about as great as length of orbit. Individual teeth shortest 
but broadest (about 1.5 times as long as broad) at base of saw, narrower and longer 
both relatively and absolutely along terminal half, where they are about four times as 
long as broad; the longest teeth a little less than half as long as distance between nostrils; 
teeth with longitudinal axes about transverse to axis of saw as a whole, their posterior 
outlines a little more convex toward tips which are sharp or variously blunted by wear; 
anterior edge sharp; posterior edge flat transversely, or nearly so, on basal teeth but 
more or less channeled longitudinally farther out on saw, perhaps as result of wear, 
thus giving each tooth two longitudinal cutting edges posteriorly. 

Head from base of saw to level of first gill opening between 1/5 and 1/4 (about 
23 "/o) of trunk from base of saw to origin of caudal; its dorsal profile weakly convex; 
distance from anterior margins of orbits to base of saw about half as great as distance 
between orbits, and distance from mouth to base of saw a little greater than breadth 
of mouth. Orbits and spiracles of about equal lengths and a little longer than nostrils. 
Gill openings with anterior outlines strongly convex; breadth between inner ends of 
fifth pair a little greater than length from mouth to base of saw. Nostrils with anterior 
margin expanded as a narrow lobe with rounded tip directed rearward across mid- 
sector of nasal aperture; the posterior margin also forming two shorter lobes. Mouth 
transverse, nearly straight, its breadth about twice as great as distance between nostrils, 
with irregular folds at corners allowing wide gape. 

Oral teeth I^Eivej minute, and dome-shaped anteriorly, with obtuse transverse 
cutting edge and base extended posteriorly in characteristic form (Fig. 4L); close-set 
in quincunx along each jaw, in a band nearly semicircular in cross section, with about 
10—12 rows in function simultaneously. 

The two dorsals similar in shape and of about equal size; subtriangular; anterior 
margins weakly convex toward apex, posterior margins moderately concave toward 
base, free rear corners about half as long as bases, vertical heights about equal to 
distance between outer ends of spiracles or a little greater; origin of first dorsal about 
over origin of pelvics. Interspace between first and second dorsals about 1.5 times as 
long as base of first dorsal. Interspace between second dorsal and origin of caudal a 
little shorter than base of second dorsal. Caudal subtriangular with narrowly rounded 
corners, its upper margin nearly straight and about as long as distance from origin 
of second dorsal to midpoint of base of first dorsal; lower margin weakly convex, 
posterior margin slightly sinuous, with lower posterior corner projecting only slightly. 

64. 25-26 in newborn male; 24-24, 24-25, and 25-25 on 30-inch specimens from Marco, Florida; 24-24 on male 
1,420 mm; 26-27 on 14-foot specimen from Key West, Florida; and 28—28 on saw of one more than 12 feet long, 
among those that we have seen. A specimen apparently of this species has been recorded as having only 22 teeth 
on one side but 25 on the other (Bloch, Naturg. ausland. Fische, i, 1785: 42). 

65. On two large saws, each about 640 mm long, the widest spacing (basal pair) is about 56 mm, the narrowest, a few 
pairs posterior to the tip, is only about 10 mm. 



2 8 Memoir Sears Foundation for Marine Research 

Pelvics about as long from origin to rear tip as interspace between dorsals; anterior 
margins nearly straight; corners abrupt. 

Color. Nearly uniform dark mouse gray to blackish brown above, paler along 
margins of fins. White to grayish white or pale yellow below. 

Size. The Common Sawfish is about two feet long at birth. It is known to reach 
a length of about i8 feet, perhaps even more.''' Most of the individuals that wander 
northward along the Atlantic Coast of the United States to North Carolina and beyond 
are large. Thus the smallest of nine specimens taken on Cape Lookout Shoals was 
12V2 feet long, while others as long as 16 feet, and none very small, have been re- 
ported from estuarine waters of North Carolina. One taken off Ocean City, Maryland, 
was 10 feet long; the two so far reported from New Jersey were 16 feet and 16 feet 

3 inches ; the most northerly record of all, from the vicinity of New York, was for one 
of I 5 feet. A female of 1 5 feet has been found to contain embryos, but the size at which 
they first breed is not known for either sex. One 12V2 feet long was reported as weighing 
425 pounds; another of about 16 feet, 700 pounds. 

Developmental Stages. Embryos, so young that they still bear the large yolk sac, 
already resemble their parents as regards relative position of fins and absence of lower 
caudal lobe; however, the posterior edges of the dorsals do not assume their characteristic 
concavity until near the time of birth. Their saws, like those of other embryonic Saw- 
fishes, are soft and leathery previous to birth, and the rostral teeth are also soft and 
entirely enclosed in the skin until birth, while a narrow band along each margin of the 
saw is naked, both below and above. But the teeth attain their full size proportionate 
to the size of the saw, and the margins of the latter become completely clothed with 
denticles soon after the young are set free. Gravid females have been found with 15— 
20 embryos. 

Habits. This species, like other Sawfishes, is almost exclusively restricted to the 
immediate vicinity of the land and to water only a few feet deep. Indeed, we once had 
the rather startling experience of striking a Sawfish of 12—14 feet with the bottom of 
our boat when rowing in water only about three feet deep near Key West, Florida. And 
we have seen another large one swimming there in water so shoal that its dorsal fins 
were above the surface. It is most often encountered in partially enclosed waters, lying 
in the deeper holes on bottoms of mud or muddy sand, and it frequents brackish water 
as often as water of oceanic salinity. It has long been known to run up into fresh water 
regularly, perhaps to remain there permanently, as in the lower reaches of the Amazon, 
in the Essequibo in British Guiana, in the Atrato and San Juan rivers of Colombia 
(tributary to the southwestern Caribbean), in the lower Mississippi, and in the St. Johns 
River, Florida, which they are said to ascend to Jacksonville.*' On the other hand, 
such specimens as wander northward in summer along the southeastern United States 

66. The maximum recorded length with which we are acquainted is i8 feet I'/j inches (vaguely rumored up to 20 ft.), 
while specimens of 12-16 feet have been reported repeatedly. 

67. A landlocked population of this species may exist in Lake Nicaragua, where the existence of two kinds of Sawfishes 
has been reported (Marden, Nat. geogr. Mag., 86, 1944: 184; and personal communication from Jose Arguillo 
Gomez). 



Fishes of the JVestern North Atlantic 29 

necessarily follow the outer coast for at least the major part of their journeys. To reach 
Bermuda, as a few certainly do from time to time, they must cross at least 600 miles 
of open ocean. To judge from the latitudinal limits within which they are year-round 
residents and from the summer-winter temperatures of the Carolinian waters that they 
visit during the warm half of the year, the lower thermal limit to their normal range is 
probably about 16— i8°C (60— 65° F); the upper limit may even be as high as 30° C 
(86° F).' 

P. pectinatus^ like its relatives, obtains its prey of various small animals chiefly by 
stirring the mud with its saw, and it may often be seen while so employed. The motion 
of the saw is described by an eyewitness as principally backward and forward.** But 
they are recorded also as playing havoc among schools of small mullets and clupeoids, 
slashing sidewise at them with their saws and devouring the wounded victims. They bite 
a hook freely if it is baited with fresh fish. 

The fact that young are abundant off Texas, with specimens of three feet or smaller 
being much more plentiful than larger adults both along the west coast of Florida 
and in the Indian River on the east coast, is good evidence that large numbers of young 
are born at least that far northward, as well as along the coasts of British and French 
Guiana*^ to the southward by similar evidence. The presence of small ones in greater 
numbers than adults up the St. Johns River makes it likely that young are not only 
produced in fresh water but that they thrive there. But there is no reason to suppose 
that any young that may be produced by stray females north of Florida would survive 
the cold of the succeeding winter. 

Gravid females, with embryos far advanced in development, have been taken in 
southern Florida waters in April and in July, and small free-living specimens have been 
caught there in January, suggesting that the young are set free in that region from 
late spring through the summer, and perhaps through the autumn. The presence of 
"young" specimens in abundance off southern Texas in May and June'" and in July 
near Galveston'^ is in line with this conclusion, for it is not likely that any born there 
in autumn would grow much during the first winter. Farther south, where the seasonal 
range of temperature is narrower and where the winter temperatures are considerablv 
higher, it is likely that young are produced throughout the year. 

It has been suggested that the period of gestation is about one year," but available 
information is not sufficient to establish this point. 

Numerical Abundance. The Common Sawfish is so plentiful in Florida waters that 
an eyewitness writes of seeing hundreds of them, "big and little,"'^ on the west coast 
of the peninsula, and one fisherman reports the accidental capture of 300 in his nets 
in the Indian River in a single season."* Many small ones are caught by fishermen 

68. Henshall, Bull. U.S. Fish Comm., 9, 1S91: 372. 

69. Miiller and Troschel, in Schomburgk, Reisen Brit. Guiana, 3, Fauna Flora Brit. Guiana, 1848: 642; Puvo, Bull. 
Soc. Hist. nat. Toulouse, 70, 1936: 89. 

70. Baughman, Copeia, 1943: 45. 71. See Study Material, p. 13. 

72. Nichols, Bull. Amer. Mus. nat. Hist., 37, 1917: 876. 73. Henshall, BuU. U. S. Fish Comm., 14, 1S95: 210. 

74. Evermann and Bean, Rep. U.S. Comm. Fish. (1896), 1898: 239. 



3 o Memoir Sears Foundation for Marine Research 

along the coast of French Guiana," and it is common along the south coast of Cuba.'* 
Definite information is lacking, however, as to its numerical status elsewhere. 

Relation to "Man. The Common Sawfish is of no commercial value anywhere in the 
western Atlantic, though small ones have been described as being delicious pan fish 
and the larger ones as yielding good steaks. But they are of considerable concern to 
fishermen as nuisances because of the damage they do to drift- and turtle-nets, to seines, 
and to shrimp trawls in which they often become entangled and because of the dif- 
ficulty of disentagling them without being injured by their saws (p. 20). 

A few are occasionally caught, incidentally, by anglers with hook and line, while 
some are harpooned. A large one puts up a powerful and dogged resistance, often 
towing a small boat for a considerable distance, as we know from experience (p. 20). 
But they are so slow-moving when hooked that few would rank them as worth special 
pursuit as game fish. 

Range. Tropical-subtropical Atlantic, north and south; equatorial West Africa to 
the Mediterranean in the east;" regularly in the west from mid-Brazil to the northern 
shores of the Gulf of Mexico and to northern Florida; to North Carolina as a summer 
visitor; less often to Chesapeake Bay; and as a rare straggler to New Jersey and the 
vicinity of New York. It is represented on the Pacific Coast of Central America (p. 2 i), 
and also in the corresponding latitudinal belt of the western Pacific and Indian oceans 
and in South African waters,'* by close relatives, but their precise relationship to P.pec- 
tinatus of the North Atlantic is still to be determined. 

Occurrence in the Western Atlantic. The localities of definite record for P.pectinatus 
are distributed widely enough to show that it is of general occurrence in estuarine 
situations and in the lower reaches of rivers, as well as along open coasts fronted by mud 
flats. Its range extends from middle Brazil northward along the South American Coast, 
throughout the Caribbean- West Indian region in general, around the western and 
northern shores of the Gulf of Mexico, and along both coasts of Florida,'* northward 
on the east coast to the Indian and St. Johns rivers. 

On the Atlantic Coast it is a year-round resident to northern Florida. Northward 
beyond that it is known only as a summer visitor. Small numbers appear yearly on the 

75. Puyo, Bull. Soc. Hist. nat. Toulouse, yo, 1936: 89. 76. Personal communication from Luis HoweU-Rivero. 

77. It has long been appreciated that old reports of Sawfishes frequenting high latitudes, and of P. pectinatus in partic- 
ular being near Spitzbergen (Bloch, Naturg. ausland. Fische, i, 1785: 42), had no basis in fact; we fancy that they 
lead back to confusion with the Narwhal. 

78. Sawfishes from South Africa have been reported recently by Barnard (Pict. Guide S. Afr. Fish., 1948: 22) and 
by Smith (Sea Fish. S. Afr., 1949: 63) under this name. 

79. Reported from Santos, Natal, and in fresh water from Para, in Brazil; from French and British Guiana; from fresh 
water in the Essequibo River; from the Island of Trinidad and Venezuela; from the Atrato and San Juan rivers 
in Colombia; from Martinique, Curasao, Puerto Rico, Haiti, Jamaica (personal communication from Luis HoweU- 
Rivero), and both coasts of Cuba; from the vicinity of Matamoros on the Gulf Coast of Mexico near the Texas 
border; along the Texas Coast in the Gulf of Mexico from oft" Laguna Madre in the south to the vicinity of Gal- 
veston in the north; from various passages and bayous along the Louisiana Coast; from Lake Ponchartrain and the 
lower Mississippi River, where it was said long ago to ascend to the Red River of Arkansas (Rafinesque, Ichthyol. 
Ohiensis, 1820: 86); from Bimini, Bahamas (personal communication from C. M. Breder, Jr.), and from many 
localities around the coasts of Florida, from Pensacola in the west to the Indian and St. Johns rivers in the east. 
It is also listed from Rio de Janeiro (doubtfully), and from northern Argentina, Lat. 38° S (Pozzi and Bordale, 
An. soc. cient. argent., 120, 1935: 152), but we have found no supporting evidence that it ranges that far south. 



Fishes of the Western North Atlantic 3 i 

shoals off Cape Lookout, and some have been said to visit the sounds and brackish 
rivers along the coast of North Carolina,*" though this perhaps calls for verification." 
A few even pass Cape Hatteras to enter the lower part of Chesapeake Bay, where one 
or two are sometimes taken in a year, but sometimes none. Stray specimens, all large, 
have been recorded also from Ocean City, Maryland, from the southern part of the 
New Jersey Coast, and one from the vicinity of New York more than a century and 
a half ago,'- this last being the most northerly record (backed by good evidence) for 
it on the Atlantic Coast of America. '^^ 

It is also known to occur occasionally around Bermuda, where a small one was 
found in the stomach of a dolphin; a second, with a saw 18 inches long, was taken 
near St. Davids; and a third (large) has been reported as seen.** But it is not likely 
that there is a locally maintained stock of Sawfishes in Bermudan waters, otherwise 
they would be reported more frequently there. 

Synonyms and Atlantic References:** 

Saegefisch, Bloch, Naturg. ausliind. Fische, j, 1785: 41, pi. 120 (descr., ill., embryo, saw, ident. by no. of teeth 
and position of first dorsal; Brazil, but ref. to Spitzbergen erroneous, see p. 30, fn. 77). 

Pes de Espada, Parra, Descr. Piez. Hist. Nat. Havana, 1787: 75, pi. 33 (descr., ill., Cuba). 

Le Squale Scie, Lacepede, Hist. Nat. Poiss., 4° Ed., in Buffon, Hist. Nat., I, 1798: 286, pi. 8, fig. 4 (descr., 
ident. by no. sawteeth and fins, coast of Africa, but other species perhaps included); and in Sonnini, Hist. 
Nat. Poiss., 4, 1801-1802: 128 (descr., size). 

Squalus prislis Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: 11, pi. 8, fig. 24 (diagn., ill., ident. by no. 
sawteeth, fins, size; Europ. Ocean); Schopf, Beschreib. nordamer. Fische, Schr. Ges. Naturf Freunde 
Berlin, 8, 1788: 185 (15-ft. specimen. New York, 1782, ident. by no. sawteeth). 

Pristis pectinatus Latham, Trans. Linn. Soc. Lond., 2, 1794: 278, pi. 26, fig. 2 (descr., ill. of saw, "Hab. in 
Oceano," doubtless Atlant.); Bloch and Schneider, Syst. Ichthyol., 1801: 351, pi. 70, fig. i (descr., ill. 
labelled "/'. serra" "in Oceano"); Lacepede, Hist. Nat. Poiss., 4° Ed., in Buffon, Hist. Nat., 4, 1801- 
1802: 680 (other species considered varieties); Risso, Ichthyol. Nice, 18 10: 22 (listed, Nice, Italy); 
Cuvier, Regne Anim., 2, 1 8 17: 131, ftn. 4 (ref to Latham, 1794); Blainville, in Vieillot, Faune Franj., 
1825: 51, pi. 12, fig. 3 (diagn., no. sawteeth, meas., color, France by ref to Risso, 1810; ill. not seen); 
Risso, Hist. Nat. Europ. Merid., 3, 1826; 141 (diagn., S. Europe); MuUer and Henle, Plagiost., 1841 : 
109 (Cayenne specimen ident. by no. sawteeth, but other species perhaps included); Bonaparte, Cat. 
Pesc. Europ., 1846: 15 (included with query); Miiller and Troschel, in Schomburgk (Robert), Reisen 
Brit. Guiana, J, Schomburgk (Rich.), Fauna Flora Brit. Guiana, 1848: 642 (size, Brit. Guiana); Gray, 
List Fish. Brit. Mus., i, 1851 : 90 (listed, Mexico, W. Indies, but refs. only in part and specimens from 
C. Good Hope and Calcutta perhaps not this species); Dumeril, Hist. Nat. Poiss., j, 1865: 475, pi. 9, 
fig. 3 (descr., ill. of saw, Haiti, but Red Sea ref. perhaps not this species); Poey, Repert. Fisico-Nat. Cuba, 
2, 1868: 456 (diagn., nos., Cuba); Gunther, Cat. Fish. Brit. Mus., 8, 1870: 437 (W. Indies, Mexico 
specimens, but refs. for Calcutta and C. Good Hope perhaps not this species); Brito-Capello, J. Sci. 
math. phys. nat. Lisboa, 4 (13), 1872: 88 (listed, Angola); Doderlein, Man. Ittiol. Medit., 2, 1881: 

80. Smith, Bull. N. C. geol. econ. Surv., 2, 1907: 39. 

81. According to Gudger, it had never been taken in Beaufort Harbor (Proc. bid. Soc. Wash., 25, 1912: 144). 

82. By Schopf, Beschreib. nordamer. Fische, Schr. Ges. Naturf. Freunde Berlin, S, 1788: 185. 

83. A Sawfish, seemingly this species because of the number of rostral teeth, was included by Denys (Hist. Nat. Amer. 
Septent., 1672, as quoted by Co.ic [Bull. nat. Hist. Soc. New Brunsw., 3 (13), 1896: 29]) in early Colonial days 
in an enumeration of fishes from the southern side of the Gulf of St. Lawrence. But the saw in question probably 
had been brought as a curiosity from some southern port. 

84. Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda, 1933: 30. 

85. See Beebe and Tee-Van (Zoologica N. Y., 26, 1941: 253-254) for references as P. peclinatus for the Pacific Coast 
of Central America; Fowler (Bull. U.S. nat. Mus., 100 [jj], 1941: 291) for the western Pacific-Indian Ocean 
region. 



3 2 Memoir Sears Foundation for Marine Research 

io8 (synonyms, descr., Medit. by ref. to Duhamel, 1782); Moreau, Hist. Nat. Poiss. France, j, 1881 : 
377 (descr., cf. other species, S. France, by ref. to Duhamel, 1782); Poey, in Gundlach, An. Soc. esp. 
Hist. Nat., JO, 1 881: 349 (listed, Puerto Rico, Mexico, Cuba, but ref. for Calcutta probably not this 
species); Goode and Bean, Proc. U. S. nat. Mus., 5, 1882: 240 (listed. Gulf of Mexico); Jordan and 
Gilbert, Bull. U. S. nat. Mus., 16, 1883: 875 (diagn., P.pectinatns replaces antiqmrum. Gulf of Mexico; 
ascending lower Mississippi R.); True, List V'ert. Anim. S. Carolina, in Handb. S. Carolina, 1883: 261 
(listed, S. Carolina); Goode, Fish. Fish. Industr. U. S., I, 1884: 668, pi. 248 (nos., sizes, Florida Coast, 
St. Johns R.; C. May, New Jersey); Jordan, Proc. U. S. nat. Mus., 7, 1884: 105 (Key West, Florida); 
Jordan and Swain, Proc. U. S. nat. Mus., 7, 1884: 230 (listed, Cedar Keys, Florida); Jenkins, Johns 
Hopk. Univ. Circ, 5 (43), 1885: 11 (listed, Beaufort, N. Carolina); Jordan, Rep. U. S. Coram. Fish. 
(1885), 1887: 798 (listed, W.Indies, S. Atlant. and Gulf States); Garraan, Bull. Mus. comp. Zool. 
Harv., ly, 1888: 88, pi. 23 (mucous canals); Carus, Prod. Fauna Medit., 2, 1889-1893: 515 (listed, 
Medit.); Henshall, Bull. U. S. Fish Comm., 9, 1891: 372 (sizes, habits, nos., W.Florida); Evermann 
and Kendall, Bull. U. S. Fish Comm., 12, 1894: 95 (listed, Galveston, Texas); Lonnberg, Oefvers. 
Svensk. Vet. Akad. Forh., 51 (3), 1894: in (listed Punta Gorda, Florida); Henshall, Bull. U. S. Fish 
Comm., J^, 1895: 210 (nos., size, feeding, Florida); Cox, Bull. nat. Hist. Soc. N. Brunsw., 5 (i3)> 
1896: 29 (ref. to S. shore Gulf St. Lawrence, by Denys, 1672, doubtless erroneous; see p. 31); Ever- 
mann and Bean, Rep. U. S. Comm. Fish. (1896), 1898: 239 (size, nos., season, embryos, Indian R., 
Florida); Jordan and Evermann, Bull. U. S. nat. Mus., ^7 (i), 1896: 60 (diagn., tropical, to W. Indies 
and Florida, Gulf of Mexico, lower Mississippi R.); Rep. U. S. Comm. Fish. (1895), 1896: 220 (listed, 
tropical, to W. Indies and Florida); Smith, Rep. U. S. Comm. Fish. (1895), 1896: 175 (listed, Biscayne 
Bay, Florida); Jordan and Evermann, Bull. U. S. nat. Mus., 4.^ (3), 1898: 2749 {P- zephyreus con- 
sidered distinct from pectinatus)\ Osorio, J. sci. math. phys. nat. Lisboa, (2) 5, 1898 : 201 (listed, Angola); 
Evermann and Kendall, Rep. U. S. Comm. Fish. (1899), 1900: 49 (W. and E. Florida); Jordan and 
Evermann, Bull. U. S. nat. Mus., 4"] (4), 1900: pi. 8, fig. 27 (ill., Pensacola, Florida); Wilson, Amer. 
Nat., 24-> 1900: 355 (listed, Beaufort, N. Carolina); Gregg, Where to Catch Fish, 1902: 23 (ill., listed, 
Florida); Evermann and Marsh, Bull. U. S. Fish Comm., 20 (i), 1902: 64 (size, nos., no. embryos, 
Indian R., Florida); Turner-Turner, Giant Fish. Florida, 1902: 190, 191 (habits, photo); Schreiner 
and Ribeiro, Arch. Mus. nac. Rio de J., 12, 1903: 80 (doubtfully listed, Rio de Janeiro); Werner, Zool. 
Jb., Syst. Abt., 21, 1904: 293 (no. sawteeth), 294 (in Key); Hargreaves, Fish. Brit. Guiana, 1904: 14 
(listed, Brit. Guiana); Jordan, Guide to Study Fish., J, 1905: 550 (in brackish water); Fowler, Proc. 
Acad. nat. Sci. Philad., ^8, 1906: 80 (listed, Florida Keys); Rep. N. J. Mus. (1905), 1908: 69 (listed. 
New Jersey, size); Ribeiro, Ann. Mus. nac. Rio de J., 14, 1907: 173 (diagn., Brazil), 206 (refs.); Smith, 
Bull. N. C. geol. econ. Surv., 2, 1907: 39 (diagn., ill., N. Carolina); Anon., Bull. N. Y. zool. Soc, 2q, 
1908: 424 (function of saw); Regan, Biol. Cent. Amer. Pisces, 1908: 183 (listed, Mexico, Central Ame- 
rica); Fowler, Proc. Acad. nat. Sci. Philad., 62, 1910: 469 (listed, Matamoros, Mexico, Essequibo R., 
Brit. Guiana); Vincent, Sea Fish. Trinidad, 1910: 52 (type of bottom, capture, Trinidad); Engelhardt, 
Zool. Anz., J9, 1912: 648 (listed. Para, Brazil, fresh water); Gudger, Proc. biol. Soc. Wash., 25, 1912: 
144 (no. sawteeth, size, capture, C. Lookout, N. Carolina); Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 
4 (3), 1913: loi (range); Garman, Mem. Harv. Mus. comp. Zool., j6, 1913: 262 (descr., refs. in 
part); Halkett, Check List Fish. Canad., 1913: 42 (erroneously listed by ref. to Denys, 1672; see p. 31); 
Starks, Leland Stanf. Univ. Pub., Univ. Ser., 191 3: 6 (listed, Natal, Brazil); Fowler, Copeia, 2, 1914: 
2 (no. sawteeth, size. Ocean City, Maryland); Coles, Proc. biol. Soc. Wash., 28, 191 5: 92 (nos., size, 
C. Lookout, N. Carolina); Fowler, Proc. Acad. nat. Sci. Philad., 6^, 191 5: 245 (listed. Ft. Pierce, 
Florida); Radcliffe, Bull. U. S. Bur. Fish., 24^ 1916: 268 (no. sawteeth, oral teeth, denticles, ills., C. 
Lookout, N.Carolina); Wilson, Ann. Carneg. Mus., 10, 1916: 58 (listed, Georgetown, Brit. Guiana, 
perhaps Atrato R. and San Juan R. in the north, Colombia); Nichols, Bull. Amer. Mus. nat. Hist., 37, 
1917: 876 (embryos, size, breeding season, S. Florida); Ribeiro, Rev. Mus. paul., 10, 1918: 708 (listed, 
Santos, Brazil); Metzelaar, Trop. Atlant. Visschen, 1919: 7 (listed, Cura9ao, W. Indies), 192 (listed, 
trop. W. Africa); Eigenmann, Indiana Univ. Stud., 46, 1920: 10 (listed, Atrato R., Colombia); Fowler, 
Proc. biol. Soc. Wash., JJ, 1920: 145 (listed, C. May County, New Jersey, New York by ref to Schopf, 
1788); Lahille, Physis B. Aires, 5, 1921: 63 (listed, Argentina); Eigenmann, Mem. Carneg. Mus., 9, 
1922: 25 (listed, Quito R., Atrato Basin, Colombia); Meek and Hildebrand, Field Mus. Publ. Zool., 
1^ (i), 1923: 66 (no Panama record); Ribeiro, Fauna brasil., Peixes, 2 (i), Fasc. i, 1923: 30, pi. 10 



Fishes of the Western North Atlantic 3 3 

(diagn., good ills., Brazil); Breder, Bull. Bingham oceanogr. Coll., I (i), 1927: 5 (listed, W.Indian 
region); Hildebrand and Schroeder, Bull. U. S. Bur. Fish., ^J, 1928: 55 (descr., color, size, Chesa- 
peake Bay); Rev, Fauna Iberica, Feces, J, 1928: 501 (ill.), 502 (in Key); Breder, Field Bk. Mar. Fish. 
Atlant. Coast, 1929: 27 (diagn., size, util., range); Jordan, Manual Vert. Anim. NE. U. S., 1929: 16 
(genl.); Truitt, Bean and Fowler, Bull. Md. Conserv. Dep., J, 1929: 27 (diagn., listed Worcester County, 
Maryland, Chesapeake Bay); Nichols, Sci. Surv. Porto Rico, Spec. Publ. N. Y. Acad. Sci., 10 (2), 1929: 
185 (listed, Porto Rico by ref. to Poey); Jordan, Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 
2, 1930: 23 (in check list); Uriarte and Mateu, Notas Inst. esp. Oceanogr., (2) 53, 1931: 27 (nos. 
sawteeth. Canaries, Lagos, W.Africa); Gowanloch, Bull. La. Conserv. Dep., 21, 1932: 88, 89 (diagn., 
size, edibility, Louisiana); Beebe and Tee- Van, Field Bk. Shore Fish. Bermuda, 1933: 30 (Bermuda); 
Gowanloch, Bull. La. Conserv. Dep., 23, 1933: 209 (feeding habits, Louisiana); Gudger, J. Elisha 
Mitchell sci. Soc, ^9, 1933: 59 (listed, C. Lookout, N. Carolina); Beltran, List. Feces Mexicanos, 1934: 
lo (listed, Mexico; not seen); Gunter, Copeia, 1935: 39 (Louisiana); Pozzi and Bordale, An. Soc. 
cient. argent., 120, 1935: 152 (listed, Argentina, Lat. 38° S); Fowler, Bull. Amer. Mus. nat. Hist., 
70 (i), 1936: 94 (diagn., size, ill., Mexico, Guiana, W. Indies; refs. for trop. W. Africa); Bull. Amer. 
Mus. nat. Hist., 70 (2), 1936: 1153 (listed. Canaries, Lagos, W.Africa by ref. to Uriarte and Mateu, 
193 1); Gunter, La. Conserv. Rev., 5 (4), 1936: 45 (Louisiana); Puyo, Bull. Soc. Hist. nat. Toulouse, 
"JO, 1936: 89 (sizes, nos., French Guiana); Smith, Biol. Rev. (Cambridge), II, 1936: 64 (listed, fresh 
water); Norman and Eraser, Giant Fishes, 1937: 60 (ill., range); Budker, Ann. Inst, oceanogr. Monaco, 
18, 1938 : 236 (scales, sensory pits, no local.); Gunter, Amer. Midi. Nat., 26, 1941 : 196 (Texas); Fowler, 
Arqu. Zool. Estad. Sao Paulo, j, 1942: 130 (listed, Santos and Natal, Brazil); Fish Culturist, 21 (9), 
1942: 66 (listed, Cuba); Gunter, Amer. Midi. Nat., 25, 1942: 312 (Texas); Baughraan, Copeia, 1943: 
43 (season, gravid females, young, Texas); Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1945: 264 
(listed, Florida); Schultz, Proc. U. S. nat. Mus., gg, 1949: 21 (no. saw teeth; Venezuela). 

Priuis granulosa Bloch and Schneider, Syst. Ichthyol., 1 801: 352 (no. sawteeth, by ref. to Parra, 1787). 

Pristis serra Bloch and Schneider, Syst. Ichthyol., 1801: pi. 70, fig. i (ill.). 

Pristis mississippiensis Rafinesque, Ichthyol. Ohiensis, 1820: 86 (descr., saw, young; listed, lower Mississippi 
to Red R., Arkansas, but rep. for Ohio R. probably erroneous). 

Pristis antiquorum Owen, Odontogr., 1840-184 5: pi. 8, fig. i (ill. of saw, sawteeth, oral teeth); DeKay, Zool. 
N. Y., 4, 1842: 365 (listed. New York, by ref to Schopf, 1788); Hall, 24th Ann. Rep. N. Y. St. Mus. 
(1870), 1872: 36 (listed New York by ref. to Schopf, 1788); Gill, Rep. U. S. Comm. Fish. (1871- 
1872), 1873: 811 (listed, C. Cod to Florida); Yarrow, Proc. Acad. nat. Sci. Philad., 1877: 217 (Beau- 
fort, N. Carolina); Uhler and Lugger, Rep. Comm. Fish. Md., ist ed., 1876: 190; 2nd ed., 1876: 160 
(Chesapeake Bay); Goode, Proc. U. S. nat. Mus., 2, 1879: 120 (listed, E.Florida); Jordan and Gil- 
bert, Bull. U. S. nat. Mus., 16, 1883: 37 (diagn., size, both shores Atlant.). 

Pristis acutirostris Dumeril, Hist. Nat. Poiss., i, 1865: 479 (descr., ident. by nos. sawteeth and position of 
fins, Antilles); Benin, Bull. Mus. Hist. nat. Paris, (2) 11, 1939: 78 (type specimen in Paris Mus., An- 
tilles). 

Pristis megalodon Dumeril, Hist. Nat. Poiss., i, 1865: 476, pi. 9, fig. 4 (descr., ill., saw, no local.); Bertin, 
Bull. Mus. Hist. nat. Paris, (2) 11, 1939: 78 (type in Paris Mus., Cayenne, French Guiana). 

Pristis, no specific name. Shields, Amer. Nat., 13, 1879: 262 (i6-ft., 700-pound specimen. Grassy Sound 
near C. May, New Jersey). 

Probable synonyms: 

Vivelle ou Poisson a Scie, Duhamel, Traite Peches, 3, Sect. 9, 1782: 331, pi. 25, figs. 3-5 (reported S. France, 

but specimen ill. without stated local.). 
Pristis occa Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 49; Faune Senegambie, J, Poiss., 1883: 26 

(trop. W.Africa, doubtful if ident. with P. occa Dumeril [Hist. Nat. Poiss., j, 1865: 479J, no local.). 
Pristis antiquorum Clarke, in Van Doren, Fish. Atlant. Coast, 1884: 156 (habits, size, Florida). 
Pristis woermanni Fischer, Jber. Wiss. Anstalt. Hamburg (1883), j, 1884: 39 (no. sawteeth and fins as in P. 

pectinatus, but sep. because anterior sawteeth not spaced closer than posterior; Cameroon); Monod, 

Faune Colon. Fran9., 1927: 648 (by ref. to Fischer, 1884; not seen). 
Pristis typica Hoflrnan, Rev. chil. Hist. Nat., 33, 1929: 29 (no descr., Cuba, ident. probable because of local., 

listed as poisonous). 



34 Memoir Sears Foundation for Marine Research 

Pristis cusfidatus Fowler, Arqu. Zool. Estad. Sao Paulo, J, 1942: 130 (listed, Bahia de Rio de Janeiro (?), 
and Natal, Brazil, by erroneous refs. to Schreiner and Ribeiro, 1903 and to Starics, 1913). 

Doubtful References: 

Pristis pectinatus Miles, Feces Rio Magdalena, Ministr. Econ. Nac. Secc. Piscicult., 1947: 36, 37 (fresh water, 

Magdalena R., Colombia, 600 km from sea, but ident. doubtful because only 21/21 sawteeth). 
Not Squalus pristis Linnaeus, Syst. Nat., j, 1758: 235 (E. Atlant.). 
Not Pristis antijuorum Latham, Trans. Linn. Soc. Lond., 2, 1794: 277, pi. 26, fig. i (equals Squalus pristis 

Linnaeus 1758). 
Not Pristis cuspidatus Latham, Trans. Linn. Soc. Lond., 2, 1794: 279, pi. 26, fig. 3. 



Pristis perotteti Miiller and Henle 1841 
Southern Sawfish 
Figures 2, 5, 6 

Study Material. Two immature males, 862—956 mm long, from Para, Brazil, and 
a female of about 8 10 mm, from Marajo Island at mouth of Amazon River (Harv. Mus. 
Comp. Zool.); immature male, 1,376 mm long, from Lake Nicaragua; female, about 
1,400 mm long, from Lake Yzabel, Guatemala, and two small saws, 301 and 309 mm 
long, from Lake Yzabel, Guatemala (U. S. Nat. Mus.); also a male, about 950 mm 
long, from Rio Tuyra, Panama, a female, about the same size, from Rio Culebra, 
Panama, tributary to the Pacific, and a saw, 560 mm long, from Rio Tilapa, Pacific 
Coast of Panama, these last three representing the closely allied P. zephyreus Gilbert 
and Starks 1895 (see discussion, p. 41, fn. 100). 

Distinctive Characters. P. perotteti is easily distinguishable from P. pectinatus by its 
fewer sawteeth (normally 19 or fewer, rarely 20 on a side, 24—32 in pectinatus),^^ by 
its first dorsal originating considerably in advance of the origin of the pelvics (about 
on a line with the origin of the pelvics in P. pectinatus)., by the much more deeply con- 
cave posterior margins of its dorsals, and by the fact that its caudal is not only relatively 
longer but has a well defined lower lobe {cf. Fig. 5 with 3). The position of its first 
dorsal relative to its pelvics and the presence of a lower caudal lobe similarly separate it 
from P. pristis (known only from the eastern Atlantic), which agrees with P. pectinatus 
in these respects but which has even fewer sawteeth than P. perotteti (see Key, p. 22). 
Description. Proportional dimensions in per cent of total length. Male, 862 mm, 
from Para, Brazil (Harv. Mus. Comp. Zool., No. 302). Male, 1,376 mm, from Lake 
Nicaragua (U. S.Nat. Mus., No. 120468). 

Breadth: between outer corners of pectorals 27.6, 31.4. 

Snout length: in front of orbits 29.3, 24.8; in front of mouth 33.3, 28.5. 

Orbits: horizontal diameter 1.7, 1.6; distance between 5.4, 5.2. 

Spiracles: length 1.4, 1.5; distance between 4.5, 4.6. 

Mouth: breadth 5.9, 5.7. 

86. This is the character that has been invoked most often as an alternative, doubtless because many of the published 
records of the two species have been based on detached saws alone. 




Figure 5. Pristis perotteti, juvenile male, about 1,376 mm long, from Lake Nicaragua (U. S. Nat. Mus., 
No. 120468). A Cross section of caudal peduncle close anterior to caudal fin, about 0.4 X. 



3 6 Memoir Sears Foundation for Marine Research 

Nostrils: distance between inner ends 2.4, 2.3. 

Gill openings: lengths, ist 1.5, 1.5; 3rd 1.7, 1.7; 5th i.o, 1.2; distance between 

inner ends, ist 9.4, 9.7; 5th 7.1, 7.0. 
First dorsal fin: vertical height 7.2, 7.9; length of base 7.5, 8.0. 
Second dorsal fin: vertical height 7.1, — ; length of base 6.0, 6.2. 
Caudal fin: upper anterior margin 16.5, 17.6; lower anterior margin 8.3, 9.2. 
Pe Ivies: anterior margin 6.2, 7.3. 
Distance: from tip of snout to ist dorsal 55.3, 51.7; to pelvics 60.8, 56.7; to 

center of cloaca 63.2, 60.5; from center of cloaca to lower caudal 2 1.2, 21.8. 
Interspace between: ist and 2nd dorsals 9.9, ii.6;2nd dorsal and caudal 5.6, 5.7. 

Trunk strongly flattened below, about 90 "/g as high as broad at origin of first 
dorsal, narrowing rather abruptly posterior to pectorals in small specimens but only 
gradually in larger ones, with a fleshy thickening or suppressed ridge low down along 
either side from a little posterior to rear tips of pelvics to base of caudal. Pectorals 
with nearly straight edges, directed outward much more abruptly from sides of head 
than in P.pectinatus (cf. Fig. 5 with 3); anterior edge nearly straight, distal edge about 
1. 1— 1.2 times as long as anterior edge and weakly concave; outer corners narrowly 
rounded; posterior corners subangular. Extreme breadth across pectorals about as 
great as distance from mouth to axils of pelvics (thus considerably greater relatively 
than in P.pectinatus^ p. 25). Caudal peduncle at rear end of base of second dorsal 
about 2.5 times or more as broad as thick, narrowing abruptly (as seen in dorsal view) 
to about 1.3 times as broad as thick at origin of caudal. 

Dermal denticles more widely spaced in general over upper surface than in P.pec- 
tinatus (cf. Fig. 6 F with 4 G) ; their blades rather strongly oblique, ovoid, with more 
or less definite median ridge and marginal thickening toward base on either side; the 
bases more or less definitely four-cornered, evident through the skin in very young 
specimens but more concealed in larger. Denticles on saw rounded to oval, so closely 
crowded as to conceal the skin entirely; those along the margins largest; those on lower 
surface similar in form to those on upper surface, but more closely crowded. 

Saw about as long as distance from front of eye to rear corner of pectoral and 1/5 
of total length or a little more (20-22 "/o)) thus somewhat shorter relatively than in 
P.pectinatus;^'' its width at base about ^/g as great as its length on small specimens, 
increasing relatively to nearly ^/g as wide as long on large; narrowing evenly forward 
to about 0.6 as wide at tip as at base on young and to 0.4-0.5 as wide at tip as at base 
on adults; Its tip rounded; its margin not definitely narrowed between each two suc- 
cessive teeth. 

Saw teeth 16—19, possibly 20 (17/18 on one of our specimens, 19/19 on others 
seen), sometimes with one more on one side than on the other, ^^ spaced only a little 
more closely toward tip than toward base (considerably more closely toward tip In 

87. For details as to saws and saw teeth in large specimens, see Baughman (Copeia, 1943: 44). 

88. 16/17, 18/18, 18/18, 18/19, '9/19 reported in a series from Texas (Baughman, Copeia, 1943: 43). 



Fishes of the Western North Atlantic 



37 



p. pectinatus)\ about 4.5-5.0 times as long as broad; their longitudinal axes nearly 
transverse to main axis of saw; the basal eight or nine teeth on each side dipping down- 
ward, about on a plane with slope of upper surface of saw on their side, but the more 
distal teeth extending about horizontally; their average lengths about Vs as great as width 
of saw, but varying considerably from tooth to tooth and without any regular gradation 



.liiiUXiW 




UUUL- 




Figure 6. Pristis perrotteti. ^Dried saw of specimen about 865 mm long (calculated length), from Lake Yzabel, 
Guatemala (U. S. Nat. Mus., No. 11 1423), about 0.4X. 5 Saw teeth from same, about 0.8 X. C Oblique view 
of saw tooth from same, about 1.2 X. D Eye and spiracle of juvenile male, about 1,376 mm long, from Lake 
Nicaragua (U. S. Nat. Mus., No. 120468), about 0.4 x. E Lower surface of anterior part of head of same, 
about 0.4 X. F Dermal denticles from below first dorsal fin of same, about 40 x. G Dermal denticles along 
margin of saw of same, about 5 X • ^ Dermal denticles on upper lip (smaller) and upper teeth (larger) from 
middle of jaw of same, about gX-J Tooth of same in lateral view, about 8 X ■ A' Cross section of upper lip and 
anterior part of upper jaw of very young male from Para, Brazil (Harv. Mus. Comp. Zool., No. 302), to show 
dermal denticles (smaller) and teeth (larger) in side view, about I2X. L Nostril of same, about 0.8 X. 



3 8 Memoir Sears Foundation for Marine Research 

between base of saw and its tip; length of longest tooth a little more than half as great 
as distance between nostrils ; anterior margins of saw teeth sharp, but posterior margins 
flattened transversely even on very small specimens; teeth longitudinally channeled 
posteriorly on larger specimens; their anterior outlines weakly convex, their tips sharp 
on young specimens but more or less blunted on large; basal pair of teeth separated 
from apparent level of emergence of saw by a distance a little greater than length 
of orbit. 

Head from origin of saw to level of first gill openings about 7* of trunk from 
base of saw to origin of caudal ; dorsal profile about as convex as in P. pectinatus, but 
lateral outlines narrowing more abruptly to base of saw (cf. Fig. 6 E with 4 F). Orbit 
about as long as spiracle; distance from anterior margins of orbits to base of saw, and 
length from mouth to base of saw, a little greater than width of mouth. Spiracles about 
0.8 as long as nostril. Gill openings with anterior outlines strongly convex; distance be- 
tween inner ends of fifth pair about equal to distance from mouth to base of saw. Nostrils 
with anterior margin expanded as a digitate lobe with rounded tip, extending rearward 
across nasal opening, the posterior margin following a continuous arc (bilobed in 
P. pectinatus, Fig. 4F). Mouth transverse, nearly straight, with irregular folds at corners, 
its breadth about 2.4 times as great as distance between nostrils. 

Teeth increasing in number of series from about ^ on newborn specimens to 
about 1^5^ at a length of 1,300 to 1,400 mm, and perhaps more on larger individuals; 
dome-shaped anteriorly, with obtuse cutting edge, relatively somewhat larger than in 
P. pectinatus, and with bases extending farther posteriorly; rather loosely set in quin- 
cunx; the oral bands as in P. pectinatus, with about 12 rows in function simultaneously 
in each jaw. 

The two dorsals of similar shape, with deeply concave posterior margins (much 
more so than in P.pectinatus)\ free lower margins less than half (about 40 %) as long 
as bases; apices narrowly rounded, their free rear tips subangular but blunted; anterior 
2/3 of base of first dorsal anterior to origin of pelvics, its origin opposite or a little 
anterior to level of rear corners of pectorals; base of second dorsal about 80 "/g as long 
as base of first dorsal. Interspace between first and second dorsals about 1.6 times as 
long as base of first dorsal. Interspace between second dorsal and caudal nearly or quite 
as long as base of second dorsal. Caudal subtriangular, with narrowly rounded corners, 
the lower expanded to form a low but definite lobe; upper margin nearly straight or 
weakly convex and nearly as long as distance from origin of second dorsal to origin 
of first dorsal (thus considerably longer relatively than in P. pectinatus)\ lower anterior 
margin weakly convex and about half as long as the upper; the caudal relatively nar- 
rower than in P. pectinatus^ its vertical height at level of lower corner only about half 
as great as length of upper margin. Caudal axis only slightly raised. Pelvics about as 
long from origin to tip as interspace between first and second dorsals; their shapes as 
in P. pectinatus, except with distal margins weakly concave. 

Color. Upper surface of trunk of salt-water specimens, when fresh-caught, either 
dark gray or golden brown, the base of the saw also golden brown in some cases. A 



Fishes of the IVestern North Atlantic 39 

specimen from the fresh water of Lake Nicaragua is pictured^" as being mouse gray 
on saw as well as over upper and lateral surfaces of anterior part of trunk; mouse gray 
shaded with reddish along midback posterior to first dorsal fin; reddish posterior to 
latter along lower part of sides; first dorsal pale yellow with reddish free rear corner; 
second dorsal, pelvics, caudal, and lower sides posterior to first dorsal dull brick red. 
But it is not clear whether the reddish tint was normal or was the result of suffusion 
with blood below the skin. 

The preserved specimens that we have seen are dark gray above and grayish 
white below. 

Size. The size at which P. pcrottcti first matures sexually is not known, but it 
has been suggested that it grows larger than P. pectinatus. The greatest lengths actually 
measured and recorded for P.perotteti are (in order of size) 17 feet 4 inches for one 
from the Ivory Coast, Equatorial West Africa;"" 17 feet 4 inches, about 18 feet (15 ft. 
without saw), and 1 8 feet 7 inches for three others from Texas. But P. perotteti^ like 
P. microdon^ is said to reach a length of 20 teet, and apparently this is no exaggeration, 
for a saw of P. perotteti four feet long has been reported from Natal, Brazil." And 
a length of about 21 feet 6 inches (6,500 mm) has been recorded'^ for its representa- 
tive in the Indian Ocean (whether specifically identical or not, see p. 21). 

Estimated weights of about 1,300 pounds for a Texas specimen 17 feet 4 inches 
long, 1,200 pounds for another of 18 feet 7 inches, as well as 1,320 pounds (600 kilo) 
for a large one (length not stated) from French Guiana, '^ suggest that adults of this 
species are heavier, at equal lengths, than P . pectinatus (p. 28), and they may grow as 
large in fresh water as in salt, for the weights of four recently caught in Lake Nicaragua 
ranged from 354 pounds to more than 700 pounds."* 

Developmental Stages. Nothing is known of the course of development of this 
species to set it apart from P. pectinatus (p. 28). 

Habits. Compared with P. pectinatus, P. perotteti is perhaps even more strictly con- 
fined to shallow water in the immediate vicinity of the shore and to estuarine localities, 
partially enclosed lagoons and similar situations. Perhaps the 15—20 foot contour line 
would enclose practically the entire stock of the species throughout its geographic 
range in both sides of the Atlantic, and it has long been known to be as much at home 
in pure fresh water as it is in brackish or salt. In fact, the type specimen of the species 
was taken from fresh water. Also, present indications are that it tends to run farther up- 
stream in large rivers than P. pectinatus ordinarily does. Thus it has been taken some- 
thing like 450 miles up from the sea in the Amazon River (p. 41) and has long been 
known in Lake Nicaragua. While it may not be strictly landlocked there, in a topo- 
graphic sense, any more than it is up the Amazon, the fact that Sawfishes breed in the 

89. Marden, Nat. geogr. Mag., 86, 1944: 173 (colored photo). 

90. Pellegrin, Poiss. Eaux douces Afr. Occ, 1923: 31- 

91. By Starks, Leland Stanf. Univ. Pub., Univ. Ser., 1913: 6. A Sawfish taken in the Gulf of Paria, Trinidad, many- 
years ago was reported as 22 feet long by 8 feet wide, but the account (Wilson, Charlesworth Mag. nat. Hist., N. S. 
J, 1839: 519) does not state how the measurements were taken, nor was the specimen identified as to species. 

92. Annandale, Mem. Indian Mus., 2, 1909: 6. 93. Puyo, Bull. Soc. Hist. nat. Toulouse, 70, 1936: 89. 
94. Marden, Nat. geogr. Mag., S6, 1944: 184. 



40 Memoir Sears Foundation for Marine Research 

Lake'5 and are rather sluggish in habit (p. 20) makes it likely that most of the local 
inhabitants are permanent residents. In this connection, it would be interesting to know 
if any Sawfishes move back and forth indifferently between salt and fresh water in the 
tidal stretches at the mouths of the rivers they frequent or if individual specimens tend 
to remain constantly in one type of water or the other. 

Throughout most of its American range, P. perotteti inhabits temperatures higher 
than 20° C (68° F). While it may be subject to temperatures as low as about 18° C 
(65° F) along the northeast coast of Texas during periods of severe winter weather, 
this is for short periods only. At the opposite extreme, it certainly is at home in water 
as warm as 28—30° C (82—86° F), or even warmer, along the shores of the Guianas 
and northern Brazil. 

In the more strictly tropical part of its range, where it appears to be the most 
plentiful, as along the coast of French Guiana, small ones far outnumber the large 
(as is true also of P. fectinatus in its centers of abundance), which is evidence that 
while many young are produced their rate of survival is low. The fact that all spec- 
imens reported from the coast of Texas have been large, in contrast with the abun- 
dance of small ones farther south, also suggests that the production of young is confined 
chiefly to regions where the temperature of the water is at least as high as about 25— 
26° C (77—79° F) and that most of the large specimens taken to the northward in 
cooler water have spread from their tropical nursery, the journey perhaps occupying 
several years for any individual specimen, without return migration. 

Nothing else is known of its way of life to differentiate it from its relative P. pec- 
tinatus. But it is of interest that a specimen reported from Texas as 1 5 feet long from 
front of head to tip of tail had lost its saw so long previously that the wound was entirely 
healed over,** evidence that a Sawfish can in some way obtain sufficient food without 
the use of its saw for stirring the bottom. 

Numerical Abundance. No precise information is at hand as to the actual numbers 
of P. perotteti anywhere, but it has been characterized as common both along tropical 
West Africa and along French Guiana. The population inhabiting Lake Nicaragua is 
also so large that an angler reports catching four in one day. And in one summer seven 
large specimens were taken by one fisherman near Galveston, Texas.'' However, most 
of the other published records of it have been based on single specimens, while it is 
described as not common in the Amazon, though well known there.'* 

Range. Both sides of the Atlantic; tropical West Africa in the east,*' middle 
Brazil to northern Texas, and as a stray to southern Florida, in the west. It is repre- 
sented by a closely allied form (or forms) along the Pacific Coast of Central America, 
off northern Australia, off Indo-China, among the East Indies, and in the tropical- 

95. Females taken there dropped their young at the time of capture (Marden, Nat. geogr. Mag., 86, 1944: 184). 

96. Baughman, Copeia, 1943: 45. 97. By E. F. Reid (Baughman, Copeia, 1943: 43). 

98. Woodland, Proc. zool. Soc. Lond., 1934: 33. 

99. An illustration of P. perotteti, or at least of a Sawfish similar to it, is included by Costa in the Fauna Napoli (Pesci 
Cat. Sist., 3, 1854-1857: pis. 8, 9), but without mention of locality. There is no reason to suppose that P. perotteti 
ever strays as far north as the Mediterranean. A Sawfish is also reported under this same name from fresh water 
in South Africa (Barnard, Pict. Guide S. Afr. Fish., 1948: 22; not seen). 



Fishes of the IVestern North Atlantic 41 

subtropical belt of the Indian Ocean, but its exact relationship to P.perotteti of the 
Atlantic remains to be determined (p. 2i).i"'' 

Occurrence in the Western Atlantic. Knowledge of the presence of this particular 
species of Sawfish in the western side of the Atlantic is recent. In 1898, for example, 
leading ichthyologists wrote that it was "not authentically known, except from the 
rivers of Africa," ^oi and as recently as 1930 they wrote that it "probably does not occur 
in America." 1"^ Actually it had been reported long before from the coastal waters of 
tropical West Africa, from the Atlantic in general, and from the West Indies. "^ And 
a specimen in the Museum at Rio de Janeiro from Bahia de Batafogo, near Rio de 
Janeiro, was listed in 1903;"* its specific identity was established positively a few 
years later by a photograph. 1"^ It is now known that P.perotteti is widely distributed 
in suitable locations along the coasts of America within the latitudinal limits stated 
above. Localities of record for it include: Santos, the vicinity of Rio de Janeiro, Bahia 
(Sao Salvador), Natal, and Marajo Island on the coast of Brazil, as well as the Amazon 
up to Parintins, some 450 miles inland from the sea; Dutch, British and French Guiana; 
Venezuela; the San Juan River, Colombia, tributary to the Gulf of Darien;"^ Lake 
Yzabel and vicinity, Guatemala; Lake Nicaragua; from unspecified localities in the 
Gulf of Mexico and West Indies and the Texas Coast from the Mexican border 
(Brownsville) to the Louisiana line (Port Arthur), where it is said to be more common 
than P. pectinatus. This last seems to mark the usual limit to its range in that direction, 
for it could hardly have been overlooked if it occurred in any numbers along the northern 
shore of the Gulf of Mexico. The only records of it in Florida waters are of a saw at 
Key West"' and of a specimen taken at Salerno on the east coast. i"* Thus its latitudinal 
range is much less extensive than that of P. pectinatus on the American Coast, as is 
also the case in the opposite side of the Atlantic. And it has not been recorded for any 
locality on the Atlantic Coast north of Florida. 

Synonyms and Atlantic References:'^'" 

Pristis ferotteti^'^'^ Miiller and Henle, Plagiost., 1841 : 108 (descr., meas., no. teeth, Senegal, fresh water), 192 
(ref. to Valenciennes); Dumeril, Arch. Mus. Hist. nat. Paris, 10, 1861: 261 (listed, Senegal); Hist. 

100. The Sawfish that has been variously recorded from the Pacific Coast of Central America as P. zepkyreus Jordan and 
Starks 1895, as P. microdon Latham 1794, and as P.perotteti Miiller and Henle 1841 appears separable from P.pe- 
rotteti of the Atlantic only by the fact that it may have up to 23 sawteeth on a side (we have seen one with 
20/22) as compared with a recorded maximum of only 19 or 20 for the Adantic form. For references to it, see 
Beebe and Tee-Van (Zoologica N. Y., 26, 1941: 253). For references to Sawfishes recorded from the western 
tropical Pacific-Indian Ocean region as P. microdon Latham 1794 and as V.perotteti Miiller and Henle 1841, see 
Fowler (Bull. U. S. nat. Mus., 100 [/j], 1941: 295). 

loi. Jordan and Evermann, Bull. U. S. nat. Mus., 47 (3), 189S: 2749. 

102. Jordan, Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 23, footnote. 

103. Gunther, Cat. Fish. Brit. Mus., 8, 1870: 436. 

104. Schreiner and Ribeiro, Arch. Mus. nac. Rio de J., 12, 1903 : 80. 

105. Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907: 173, pi. 11. 

106. Specific identity of saws seen not altogether certain (Eigenmann, Mem. Carneg. Mus., 9, 1922: 25). 

107. Saw with only 17 teeth on one side and 18 on the other, hence almost certainly this species (Baughman, Copeia, 
1943: 46). 108. Reported to us by Stewart Springer. 

109. For references to P. zepkyreus, P. microdon and P. perotteti for the Indo-Pacific, see Beebe and Tee-Van (Zoologica 
N. Y., 26, 1941: 253) and Fowler (Bull. U.S. nat. Mus., 100 [13], 1941: 295). 

1 10. Sometimes spelled "perrotteti," or "perroteti." 



42 Memoir Sears Foujidation for Marine Research 

Nat. Poiss., J, 1865: 474, pi. 9, fig. 2 (descr., ill. of saw, rivers of Senegal); Giinther, Cat. Fish. Brit. 
Mus., 8, 1870: 436, in part (specimens from Atlant., W. Indies, but not from Chiapam, Guatemala, 
Zambesi and Bandjermassing); Steindachner, S. B. Akad. Wiss. Wien, 6/ (i), 1870: 577 (nos., Sene- 
gal); Rochebrune, Act. Soc. linn. Bordeaux, (4) 6, 1882: 48 (size, nos., Senegambia); Steindachner, 
Denkschr. Akad. Wiss. Wien, ^, 1882: 51 (listed, Senegal); Rochebrune, Faune Senegambie, j, Poiss., 
1883-1885: 25 (same as Rochebrune, 1882); Jordan and Evermann, Bull. U. S. nat. Mus., ^j (i), 
1896: 60 (descr.); Bull. U. S. nat. Mus., jf.j (3), 1898: 2749 (known only from Afr. rivers); Evermann 
and Marsh, Bull. U. S. Fish Comm., 20 (i), 1902: 64 (Senegal, said to occur in W. Indies); Schreiner 
and Ribeiro, Arch. Mus. nac. Rio de J., 12, 1903: 80 (listed, Bahia de Batafogo, Brazil); Hargreaves, 
Fish. Brit. Guiana, 1904: 14 (habits, ill., Brit. Guiana, but specimen with 23 sawteeth probably P. 
fectinatus)\ Ribeiro, Arch. Mus. nac. Rio de J., j.^, 1907: 173, 206, pi. 11 (diagn., size, refs., photo, 
Bahia de Batafogo, Brazil); Regan, Biol. Cent. Amer., Pisces, 1908: 183 (listed. Lake Nicaragua); Bou- 
lenger. Cat. Fr. Water Fish. Afr., j, 1909: 3 (listed for Gambia, but not refs. to Zambesi and Spire); 
Fowler, Proc. Acad. nat. Sci. Philad., 62, 1910: 469 (saws probably of this species, Bahia [Sao Salvador], 
Brazil, Gulf of Mexico and Surinam [Dutch Guiana]); Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 
^ (3), 191 3: loi (range); Starks, Leland Stanf. Univ. Pub., Univ. Ser., 191 3: 6 (saw. Natal, Brazil); 
Ribeiro, Rev. Mus. paul., lo., 191 8: 708 (listed, Santos, Brazil); Fowler, Proc. Acad. nat. Sci. Philad., 
yi, I9I9'. 129 (listed, Surinam [Dutch Guiana]); Metzelaar, Trop. Atlant. Visschen, 1919: 192 (listed 
both sides of Atlant.); Eigenmann, Mem. Carneg. Mus., 9, 1922: 25 (saws, San Juan R. in the north, 
Colombia); Pellegrin, Poiss. Eaux douces Afr. Occ, 1923: 32 (color, size, Sassandra, Ivory Coast, W. 
Afr.); Ribeiro, Fauna brasil., Peixes, 2 (i) Fasc. i, 1923: 30 (diagn., same as Ribeiro, 1907); Breder, 
Field Bk. Mar. Fish. Atlant. Coast, 1929: 28 (cf. P.fectinatus)\ Jordan, Evermann and Clark, Rep. U. 
S. Comm. Fish. (1928), 2, 1930: 23, footnote (Afr., prob. not Amer.); Gowanloch, Bull. La. Conserv. 
Dep., 21, 1932: 88 (not reported from Louisiana); Woodland, Proc. zool. Soc. Lond., 1934: 33 (Ama- 
zon R., near Parintins, Brazil); Puyo, Bull. Soc. Hist. nat. Toulouse, yo, 1936: 89 (size, nos., French 
Guiana); Smith, H. W., Biol. Rev. (Cambridge), 11, 1936: 64 (listed, fresh water); Berlin, Bull. Mus. 
Hist. nat. Paris, (2) 11, 1939: 77, 78 (one of orig. specimens in Paris Mus., Senegal R.); Gunter, Amer. 
Midi. Nat., 26, 1941: 196 (listed, Texas Coast); Baughman, Copeia, 1943: 43 (discuss., descr., photo 
of saw, cf. P. pectinatus, length, weight, Texas). 

Pristis antiquorum Giinther, Trans, zool. Soc. Lond., 6, 1869: 397, in part (Atlant., but not Chiapam, Gua- 
temala, and Pacif. ref.); Gill and Bransford, Proc. Acad. nat. Sci. Philad., 1877: 190 (saw. Lake Nica- 
ragua); Meek, Field Mus. Publ. Zool., 7 (4), 1907: 104 (saws. Lake Nicaragua). 

Pristis microdon Garman, Mem. Harv. Mus. comp. Zool., 56, 191 3: 265, in part (descr., synonyms, and 
refs. for Atlant.); Fowler, Bull. Amer. Mus. nat. Hist., 70 (i), 1936: 95 (W. Afr. refs., descr., Brazil, 
Surinam [Dutch Guiana], Gulf of Mexico); Bull. Amer. Mus. nat. Hist., 70 (2), 1936: 1153 (in Key); 
Rohl, Fauna Descript. \'enezuela, 1942: 366 (V'enezuela; not seen); Marden, Nat. geogr. Mag., 86, 
1944: 184, 186, pi. 5 (color photo, weights, nos.. Lake Nicaragua); Baughman, Copeia, 1947: 280 
(17—18 sawteeth, Port Isabel, Texas); Schultz, Proc. U. S. nat. Mus., 99, 1949: 21 (Maracaibo, Vene- 
zuela). 

Probable Synonyms: 

Pristis antiquorum Costa, Fauna Napol., Pesc. Cat. Sist., 3, 1854— 1857: pis. 8, 9 (ills., no local.); Metzelaar, 

Trop. Atlant. Visschen, 1919: 192, in part (listed both sides Atlant.). 
Pristis zep/iyreus Jordan and Starks, Proc. Calif Acad. Sci., (2) 5, 1895: 383 (descr., Rio Presidio, Gulf of 

California; for refs., see Beebe and Tee-Van [Zoologica N. Y ., 26, 1941: 253]). 
Pristis pristis Ribeiro, Fauna brasil., Peixes, 2 (i) Fasc. i, 1923: 30 (no. saw teeth, Brazil). 
Pristis microdon Rey, Fauna Iberica, Feces, I, 1928: 501, 505 (ill., after Costa, 1854-1857); Miles, Feces 

Rio Magdalena, Ministr. Econ. Nac. Secc. Piscicult., 1947: 36, 37 (listed, estuary of Magdalena R., 

Colombia; not seen). 
Not Squalus pristis Linnaeus, Syst. Nat., J, 1758: 235 (E. Atlant.). 
Not Pristis antiquorum Latham, Trans. Linn. Soc. Lond., 2, 1794: 277, pi. 26, fig. i (equals Squalus pristis 

Linnaeus 1758). 
Probably not Pristis microdon Latham, Trans. Linn. Soc. Lond., 2, 1794: 280, pi. 26, fig. 4 (no local., name 

commonly applied to Indian Ocean form; for refs., see Fowler [Bull. U. S. nat. Mus., 100 (13), 1941 : 

295])- 



Fishes of the IVestern North Atlantic 43 

Suborder RHINOBATOIDEA 
Guitarfishes 

Characters. Trunk anterior to cloaca ranging from shark-like to disc-like in form. 
Snout without blade-like extension, its edges without teeth. Tail sector much stouter 
than in any other batoids excepting the Sawfishes (p. 18) and Torpedo Rays (p. 80), 
sharply marked off from body sector in disc-shaped species but not in those with a more 
shark-like form. Attachment of anterior expansions of pectoral fins to sides of head 
extending only a little past mouth in some, for various distances thence forward in 
others, and nearly to midline of snout in extreme cases. Two well developed dorsal fins, 
the first originating much closer to tips of pelvics than to tip of tail. Caudal fin well 
developed. Pelvics with outer margins convexly rounded and wholly separate from 
pectorals. 

Eyes with an obscure fold of skin below. Nostrils entirely separate from mouth 
in most cases but connected with it by a broad shallow furrow^ in some. Corners of 
mouth without voluminous furrows extending rearward. 

Body and fins closely covered with small dermal denticles of various shapes, skin 
of upper and lower surfaces mostly concealed except for the interbranchial region, which 
is largely or wholly naked in some cases. Back more or less thorny along midbelt, on 
shoulders, around eyes and spiracles, and over rostral cartilage in some species. Tail 
without a serrate-edged spine. 

Anterior margin of cranium with a single unbranched rostral projection^ extending 
nearly or quite to tip of snout in most but falling considerably short of it in Platyrhina 
(p. 50), in which it is continued forward by soft ligamentous strands. In most species 
the separate antorbital cartilages do not take any part in supporting the anterior part 
of the head; in Platyrhina they extend forward beyond the orbital region but fall short 
of the anterior margin of the disc. Tips of branchial rays either cylindrical or some- 
what expanded, less so than in the Torpedo Rays. Dorsal and caudal fins supported 
basally by short cartilaginous radials and distally by much more numerous fine horny 
rays (ceratotrichia) in double series. Pectoral and pelvic fins without horny rays, the 
cartilaginous radials extending outward to margins of pectorals, nearly to margins of 
pelvics. Pelvis transverse, either with or without a small anterior process at either end. 
Anterior and posterior surfaces of gill arches inward from gill filaments with a series 
of low, firm, widely-spaced knobs. No electric organs. Development ovoviviparous in 
most species, perhaps oviparous in a few (p. 47, fn. 11).* 

Remarks. The rhinobatoids present nearly an unbroken series as regards shape of 
disc, length and shape of snout, degree of forward extension of pectorals, position of 

1. This connection between nostril and mouth is described by Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 
289) as "deep" for Platyrhina sinensis (Bloch and Schneider) 1801 ; actually we found that it is shallow though broad 
in an excellently preserved specimen of that species from Japan. 

2. Commonly termed "rostral cartilage," although it is continuous with the cranium. 

3. For an account of the embryonic stages of RJiyncfiobatos djiddensis, see Setna and Sarangdhar [Rec. Indian Mus. 
(1948), 46 (1-4), 1949: 18-23, figs. 7-8. pl- I. %-4]- 



44 Memoir Sears Foundation for Marine Research 

dorsal fins on tail, conformation of anterior margin of nostril, and relationship of nostrils 
to mouth. Some recent authors, in fact, unite all of them in a single family, Rhino- 
batidae.* They can be grouped in two families, however, according to the shape of 
the caudal, including the degree of elevation of its axis, and according to the position 
of the dorsal fins relative to the pelvics. 

Key to Families 

I a. Caudal fin conspicuously bilobed, more or less lunate in form, both lobes sharp- 
pointed; axis of caudal fin bent upward to a moderate degree; posterior edges of 
pectorals considerably anterior to origin of pelvics; origin of first dorsal over or 
a little anterior to bases of pelvics. Rhynchobatidae,* p. 44. 

lb. Caudal fin not bilobed, its axis bent upward only very little, if at all; posterior 
edges of pectorals extending rearward as far as origin of pelvics or farther; origin 
of first dorsal considerably posterior to posterior tips of pelvics. 

Rhinobatidae, p. 46. 

Family RHYNCHOBATIDAE 

Characters. General shape intermediate between shark-like and typical ray-like, 
only moderately flattened; tail sector not marked off from body sector. Snout not pro- 
duced as a blade, ranging in different genera from short and broadly rounded to wedge- 



FiGURE 7. Rhynchobatus djiddensis. Caudal fin of male, 
480 mm long, from Ceylon (Harv. Mus. Comp. ZooL, 
No. 806), about 0.8 X. 

shaped and elongate; no teeth on margins of snout. Posterior corners of pectorals 
considerably anterior to origin of pelvics. Origin of first dorsal ranging from about over 
midpoint of bases of pelvics to a little anterior to origins of latter. Caudal fin definitely 

4. Norman, Proc. zool. Soc. Lond., 1926: 941; Berlin, Bull. Inst, oceanogr. Monaco, 775, 1939: 19. 

5. Jordan (Class. Fish., Stanford Univ. Publ. Biol., j [2], 1923: 102) used the name Rhinidae for this family (derived 
from Rhina Bloch and Schneider 1801, see p. 45). But doing so is likely to lead to confusion, because Rhina has 
been used many times as the generic name of the squatinoid Sharks, though incorrectly so. 



Fishes of the Western North Atlantic 45 

bilobed (Fig. 7) with deeply concave posterior margin; both upper and lower lobes 
sharp-tipped; caudal axis moderately raised, its termination near posterior margin of 
fin some distance below tip of upper lobe. Eyeballs with low rounded velum above 
pupil, the orbits outlined below and anteriorly by a deep furrow. Spiracles large, about 
as broad as long, close to eyes and either transverse or only slightly oblique, with inner 
ends directed rearward, their posterior margins with or without transverse folds. Rostral 
projection from cranium extending to tip of snout. 

Genera. Two genera, Rhynchobatus and Rhina, are generally recognized. 

Key to Genera. 

I a. Snout narrow, pointed, its length in front of eyes nearly or quite as great as breadth 
of head at level of anterior margins of eyes; posterior margin of spiracles with 
two low vertical ridges; mouth only slightly undulate, with median projection of 
lower jaw fitting a corresponding depression of upper. 

Rhynchobatus^ Miiller and Henle 1837 
Tropical Indian Ocean, including Red Sea, south to 
Natal; tropical West Africa; Malay Peninsula; Phi- 
lippines; East Asia north to Japan; Australia.' 
lb. Snout broadly rounded, its length in front of eyes much less than breadth of 
head at eyes; posterior margin of spiracles without vertical ridges; mouth strongly 
undulate, with three forward projections of the lower jaw alternating with two 
rearward indentations of the upper. Rhina Bloch and Schneider 1801.* 

Tropical Indian Ocean in general, including 
the Red Sea and the Gulf of Oman; Malayan 
region; East Indies; Philippines; Australia; 
southeastern coasts of Asia northward to sou- 
thern China and Japan.* 

6. Rhinobatis Blainville 1825 (in Vieillot, Faune Franf., 1825: 47, 48) may have been an earlier name for the genus 
now known as Rhynchobatus (see p. 51, footnote), but we think it wiser to regard it as preoccupied by Rhinobatos 
Link 1790, since the use of names differing by only one letter for two different genera within the same suborder 
could only lead to confusion. 

7. The type species (the only species known until recendy) is Rhynchobatus djiddensis (Forskal) 1775 of the Indo- 
Pacific. A separate subspecific name, australiae, has been proposed by Whitley (Aust. Zool., 9, 1939: 245; Fish. 
Aust., I, 1940: 173, fig. 197) for Australian specimens whose form is wider toward the root of the tail and whose 
color pattern differs from the typical Rhynchobatus djiddensis. But a small specimen of the latter from Ceylon, 
examined by us, agrees so closely in both these respects with Whitley's accounts and illustration of his australiae 
that it seems unnecessary to retain that name. 

Three additional species have been described from tropical West Africa: Rhynchobatus liibberti Ehrenbaum 
(Fischerbote, 6, 1914: 303), with which R. atlanticus Regan (Ann. Mag. nat. Hist., [8] 4, 1915: 124) appears to 
be identical, Rhynchobatus albomaculatus (Norman) (Ann. Mag. nat. Hist., [10] 6, 1930: 226), and Rhynchobatus 
ir^inei (Norman) (Ann. Mag. nat. Hist., [10] 7, 1931: 352). The last two were originally placed in Rhinobatos 
but are clearly referable to Rhynchobatus by their bilobed caudals. 

8. The name Rhina was first used by Schaeffer (Epist., Stud. Ichthyol., 1760: 20) for the Shark genus properly known 
as Squatina, then by Klein (Neuer Schauplatz, 2, 1776: 587), by Walbaum (P. Artedi Genera Pise. Emend. Ichthyol., 
1792: 58), and by some more recent authors. However, Schaeffer did not include any species in his genus Rhina, 
and the International Commission on Zoological Nomenclature has ruled (Opin. Rend., Smithson. Publ., No. 
1938, 1910: 51; Smithson. miscell. Coll., 7J [3], 1925: 27) that neither Klein's names, nor Walbaum's revival of 
them, are to be taken into account. Consequently, Rhina as a generic name must date from Bloch and Schneider 
(Syst. Ichthyol., 1801: 352, pi. 72), who applied it to an Indian Ray with the specific name R. ancylostomus. 

9. For list of localides recorded for Rhina ancylostoma Bloch and Schneider 1801, see Fowler (Bull. U. S. nat. Mus., 
100 [jj], 1941: 299). 



46 



Memoir Sears Foundation for Marine Research 



Family RHINOBJTIDAE 

Characters.^'' Body sector of trunk ranging from moderately flattened, with wedge- 
shaped snout and tail sector not marked off from body, to strongly flattened and broadly 
rounded anteriorly in disc-like form, with tail sector more or less clearly marked off 
though moderately stout. Snout not produced as a blade, its edges without teeth. Anterior 
pectoral rays extending forward only a little past level of nostrils in some but to level 
of end of snout in others; posterior corners of pectorals extending rearward at least 
as far as origins of pelvics. Origin of first dorsal considerably posterior to rear tips of 




Figure 8. Rhinobatos Untiginosus, from off Useppa Island, 
Florida (Harv. Mus. Comp. ZooL, No. 35856). Gill arch 
(above) and side view of gill fold (below), about 2.8 x. 



pelvics. Caudal without definite lower lobe, its posterior contour straight or convex, 
its corners rounded; caudal axis not raised, or only very slightly so (Fig. 14 D; 18B). 
Eyes with or without rounded velum above pupil, the orbit outlined below and anteriorly 
by a deep groove in some species (as in the Rhynchobatidae) but only faintly so, if at 
all, in others. Spiracles immediately behind eyes, either transverse or slightly oblique, 
with inner ends directed rearward (as in Rhynchobatidae), their posterior margins with 
or without transverse folds. Rostral projection extending to tip of snout, or not. Anterior 
and posterior surfaces of gill arches, inward from gill filaments, each with a series of 
low, firm, widely-spaced knobs (Fig. 8). 

Development is ovoviviparous in the great majority, but perhaps oviparous in 

10. For account of the head skeleton and a comparison with other batoids, see Holmgren (Acta Zool. Stockh., 22, 
1941: 52, 64). 



Fishes of the Western Nortli Atlcuitk 47 

some." In certain ovoviviparous species, all of the embryos in each uterus are enclosed 
at first in a single pillow-shaped capsule, from which they are later set free into the 
uterus, the walls of which are thickly set with villi. ^^ It is not known whether multiple 
egg capsules of this sort occur in all ovoviviparous genera of the family. 

Genera. Two genera referred to this family by us have been made the basis of 
a separate family by some authors,!^ partly because the rostral projection falls con- 
siderably short of the tip of the snout in one of them {Platyrhind)., and probably in 
the other (^Zanobatus^*), whereas in all others it extends to the snout's extremity. In 
this respect, Platyrhina bears much the same relationship to the hard-nosed Guitar- 
fishes, as exemplified by Rhinobatos or Plalyrhinoidis, as do the soft-nosed Skates (Psam- 
mobatis^ Sympterygia, Brevirajd) to the hard-nosed Skates {Raja), with which they are 
united in a single family (Rajidae) by common consent. Therefore, we see no reason 
for their separation on this basis. ^^ 

Key to Genera 

I a. Snout wedge-shaped, ranging from obtusely so to slenderly prolonged; tips of 
anterior rays of pectorals falling considerably short of tip of snout. 
2 a. Lobe-like expansion of anterior margin of nostrils far apart from each other 
and from upper lip. 

3 a. Lobe-like expansions of anterior margin of nostrils covering only a narrow 
belt midway of nasal opening. 

4 a. Posterior margin of spiracles with one or two well marked ridges or 
folds; nostrils more or less oblique; inward extension of lobe-like 
expansion of anterior margin of nostrils ending some distance out- 
ward from inner corner of nostril. Rhinobatos Link 1790, p. 50. 
4b. Posterior margin of spiracles without folds; nostrils transverse; in- 
ward extension of lobe-like expansion of anterior margin of nostril 
crosses inner angle of nostril. Aptychotrema Norman 1926. 

Australia (two species).^* 

11. Muller (Denkschr. Akad. Wiss. Berl. [1840], 1842: 249, pi. 6, fig. 2; also separate, Ueber den flatten Hai, 1842: 
62, pi. 6, fig. 2) found in the uterus of a Zanobatus schoenUinii an egg with a horny capsule, much resembling the 
eggs of the scyliorhinid Sharks; its shape suggested that embryonic development takes place after the eggs are 
laid, since it was provided with two coiled tendrils at one end. 

12. This type of ovoviparity has been reported for the Australian Aptychotrema banksii (MiiUer and Henle) 1841, where 
each of the two capsules (one per uterus) contained 7-8 embryos (Haacke, Zool. Anz., 8, 1885: 48S, as R/iinobatus 
'vincentianus, n. sp.); also for Trygonorhina fasciata (likewise Australian), in which each capsule contained two 
or three embryos. For illustrations of the egg capsule and of the villous uterine wall in the former, see McCulloch 
(Biol. Result. Fish. F. I. S. 'Endeavour', 5, 1926: 158, figs. 1-4, as RAinobatus banksii), and Whitley (Fish. Aust., 
I, 1940: 172, fig. 196, as Aptychotrema 'vincentiana). For an account of the embryonic stages of Rhinobatos halai'i 
and Khynchobatos djiddensis, see Melouk (Publ. Mar. bid. Sta., Ghardaqa, Fouad I Univ., 7, 1949). 

13. Discobatidae (Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 258); Platyrhinidae (Fowler, BuU. U.S. nat. 
Mus., 100 [jj], 1941: 289). 

14. Norman (Proc. zool. Soc. Lond., 1926: 982) writes "as far as I am able to judge from a partial dissection of this 
very young example, the rostral cartilage does not reach the extremity of the snout." 

15. In this we follow Norman's Synopsis (Proc. zool. Soc. Lond., 1926: 941). 

i6. The characters used by Norman (Proc. zool. Soc. Lond., 1926: 977) in separating the genus Aptychotrema from 



48 Memoir Sears Foundation for Marine Research 

3 b. Lobe-like expansion of anterior margin of nostrils covering inner half of 
nasal opening almost entirely (Fig. 1 8 C). 

Zapteryx Jordan and Gilbert 1881, p. 74.^' 




Figure g. Trygoriorhina fasciata, female, 254 mm long, 
from Port Jackson, New South Wales (Harv. Mus. Comp. 
ZooL, No. 982). Ventral view of anterior part of head 
to show shape of nasal curtain, about 1.6 X- 




Figure 10. PlatyrkinoiJis triseriata, female, 452 mm long, 
from southern California (Harv. Mus. Comp. ZooL, No. 
36481). Nostrils and mouth, about ix. 



Khinobatos are slight, but the genus has been accepted by both Whitley (Fish. Aust., I, 1940: 169) and Fowler 
(Bull. U. S. nat. Mus., 100 [/j], 1941 : 322); this course has the practical advantage of subtracting a few from the 
large number of closely related species of the genus Khinobatos. 

The nomenclatural history of the generic name Zapteryx is confused. The earliest named representative of the 
genus, Khinobatos [Syrrhina) breuirostris Miiller and Henle 1841, was placed by its authors in Syrrhina, their new 
subgenus of the old genus Khinobatos. However, Muller and Henle failed to designate any one particular species 
as the type oi Syrrhina. In 18S1 Garman (Proc. U. S. nat. Mus., 3, 1881: 521) revived Syrrhina for the Californian 
Ray that had been described the year before by Jordan and Gilbert (Proc. U. S. nat. Mus., 3, 1880: 32) as Pla- 
tyrhina exasperata but by Garman (Bull. Mus. comp. Zool. Harv., 6, 1880: 169) as Trigonorhina alnjeata, which is 



Fishes of the JVestern North Atlantic 49 

2 b. Expansions of anterior margin of nostrils united to form a single broad quad- 
rangular curtain, the free edge close to upper lip, with which it is united at 
isthmus (Fig. 9). Trygonorhina}^ Miiller and Henle 1841. 

Australia. 
lb. Snout rounded; tips of anterior rays of pectorals reaching nearly or quite to tip 
of snout. 
5 a. Rostral cartilage extending nearly or quite to tip of snout. 

6a. Dorsal surface of tail with three rows of prominent thorns; origin of first 
dorsal much nearer to origin of second dorsal than to rear tips of pelvics; 



Figure ii. Xanobatus schoenleinii. Nostril 
with inner end to the left; after Norman. 



lobe-like expansion of anterior margin of nostril almost entirely covering 
inner end of nasal aperture; posterior margin of nostril only slightly 
expanded (Fig. 10). Flatyrhinoidis Garman 1881. 

Coast of California from Point Con- 
ception southward.^® 
6b. Dorsal surface of tail with only one median row of large thorns; origin 
of first dorsal about midway between rear tips of pelvics and origin of 
second dorsal; lobe-like expansion of anterior margin of nostril leaving 
inner part of nasal opening exposed; posterior margin of nostril expanded 
as a broad rounded flap (Fig. 11). Zanobatus Garman 1913. 

India and tropical West Africa. 2" 

clearly congeneric with Rhinobatos [Syrrhina] bre'-oirostris MuUer and Henle 1841. But, since this revival of Syr- 
rhina had been antedated by Jordan and Gilbert's (Proc. U. S. nat. Mus., 3, 1880: 53) proposal of the generic 
name Zapteryx for exasperata, we follow prevalent usage (Jordan and Evermann, Bull. U. S. nat. Mus., 47 [i], 
1896: 64; Jordan, Genera Fish., 2, 1919: 192; Norman, Proc. zool. Soc. Lond., 1926: 979; Beebe and Tee-Van, 
Zoologica N. Y., 26, 1941 : 252) in employing Zapteryx and in relegating Syrrhina to the synonymy of Rhinobatos. 
See also under generic synonyms of Rhinobatos, p. 50. 

18. One species, Trygonorhina fasciata MuUer and Henle 1841, with color varieties. These appear to be genetic rather 
than environmental, for specimens experimented upon showed only slight alterations in shade on white and on 
black backgrounds (Griffiths, Proc. Linn. Soc. N. S. \V., 61, 1936: 319). For an account of the skeleton of Try- 
gonorhina, see Haswell (Proc. Linn. Soc. N. S. W., g, 1884: 107, pi. 2, figs. 1-5). 

19. One species only, the Platyrhina triseriata of Jordan and Gilbert (Proc. U. S. nat. Mus., J, 1880: 36), for which 
Garman (Proc. U.S. nat. Mus., j, 1881: 522) proposed the genus Platyrhinoiiiis. 

lo. Illustrations of the adult of this genus (Muller and Henle, Plagiost., 1841 : pi. 45; Steindachner, Denkschr. Akad. 
Wiss. Wien, .^4, 1882: pi. 7) seem to picture the rostral cartilage as extending nearly or quite to the tip of the snout. 
But it appeared not to reach the extremity of the snout in a very young one examined by Norman (Proc. zool. Soc. 
Lond., 1926: 982). The West African form, originally reported and pictured by Steindachner as Platyrhina schoen- 
leinii Muller and Henle 1841, has been redescribed recently by Chabanaud (Bull. Soc. zool. Fr., 53, 1928: 419) 
as a separate species, Platyrhinoidis atlantica. 



50 Memoir Sears Foundation for Marine Research 

5 b. Rostral cartilage extending only a short distance beyond front of cranium 
and less than halfway to tip of snout. 

Platyrhina"^^ Miiller and Henle 1841. 
China and Japan. 



Genus Rhinobatos Link 1790 

Guitarfishes 

Rhinobatos Link, Mag. Physik Naturg., Gotha, 6 (3), 1790: 32; diagnosis, but no species mentioned; evidently 
for Raja rhinobatos Linnaeus 1758, which is generally accepted as the type species. 

Generic Synonyms:-- 

Rhinobatus Bloch and Schneider, Svst. Ichthyol., 1801: 353, and many subsequent authors; equivalent to 

Rhinobatos Link.^* 
Leiobatus Rafinesque, Caratt. Gen. Spec. Sicil., 1810: 16; type species, L. fanduratus Rafinesque, Sicily; ap- 
parently equals Squalus rhinobatos Linnaeus 1758.-* 
Rhinobates Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1839: 193; equivalent to Rhinobatos Link 1790 and 

Rhinobatus Bloch and Schneider 1801. 
Squatinoraja Nardo, Oss. Agg. Adr. Ittiol., 1824;^^ type species, S. colonna Nardo, designated by Jordan (Genera 

Fish., J, 1917: 116), equals Rhinobatus columnae Bonaparte 1835— 1836. 
Syrrhina (subgenus oi Rhinobatus) Miiller and Henle, Plagiost., 1841: 113, in part. For discussion, see p. 48, 

footnote 17. 
Glaucostegus Bonaparte, Cat. Pesc. Europ., 1846: 14; type species, Rhinobatus cemiculus GeofFroy St.-Hilaire 

1827; perhaps equals Raja rhinobatos Linnaeus 1758.^^ 
Squatiniraja L. Agassiz, Nom. Zool., Inde.x, 1846: 350; emended spelling for Squatinoraja Nardo 1824. 
Acroteriobatis (subgenus) Giltay, Ann. Soc. zool. Belg., ^g, 1928: 26; type species, Rhinobatus {Syrrhina) annu- 

latus Miiller and Henle 1841, designated by Whitley (Aust. Zool, 9, 1939: 244). 
Scobatus (subgenus) Whitley, Aust. Zool., 9, 1939: 244; type species, Rhinobatus granulatus Blainville i8r6, 

designated by Whitley. 

21. The name Discobatus was proposed by Garman (Proc. U. S. nat. Mus., J, 1881: 523) to replace Platyrhina, pre- 
occupied by Platyrhinus Schellenberg 1798, for Coleoptera; and another substitute name, Analithis, had been pro- 
posed earlier by Gistel (Naturg. Tier., 1848: x). But this substitution is not required by the rules of Zoological 
Nomenclature as generally accepted at present. Two species are known, Platyrhina sinensis (Bloch and Schneider) 
1801 and P . limbomhenyi Tang 1933 (Lingnan Sci., J., 12^ 1933: 561, pi- 42, figs, i, 2). For excellent illustra- 
tions o{ P. sinensis, including skeletal characters, see Garman (Mem. Harv. Mus. comp. Zool., j6, 1913: pi- 66). 

22. The fossil genera Jellopos Miinster 1836 (Neues Jb. Miner., 1836: 581), Euryarthra L. Agassiz 1843 (Poiss. Foss., 
J, 1843: 382) and Spathobatis ThioUi^re 1849 (Ann. Soc. Agric. Lyon, [2] i, 1849: 63) are included among the 
synonyms of Rhinobatos by Fowler (Bull. U.S. nat. Mus., 100 [xj], 1941: 305)- 

23. The generic name Rhinobatus had been employed previously by Klein (Neuer Schauplatz, 2, 1776: 592) and by 
Walbaum (P. Artedi Genera Pise. Emend. Ichthyol., 3, 1792: 58 1). But the International Commission on Zoological 
Nomenclature has ruled (Opin. 21, Smithson. Publ., No. 1938, 1910: 51; Opin. 89, Smithson. misc. Coll., yj [3], 
1925: 27) that neither Klein's names nor their revival by Walbaum (1792) are to be taken into consideration. 

24. Leiobatus had been proposed earlier by Klein (Neuer Schauplatz, i, 1775: 316) and republished by Walbaum 
(P. Artedi Genera Pise. Emend. Ichthyol., 2, 1792: 581), its type being designated as Raja oxyrincha Linnaeus 
1758 by Jordan (Genera Fish., i, 1917: 38). 

25. Squatinoraja is credited to Nardo, as above, by L. Agassiz (Nom. Zool. Pise, 1845: 61), by Schultze, Kiikenthal, 
Heider, and Hesse (Nom. Anima. Gen. Subgen., 5 Lief. 23, 1937: 3260), and by Neave (Nom. Zool., 4, 1940: 
271). We have not been able to verify the citation. 

26. While R. cemiculus St.-Hilaire (in Savigny, Zool. Egypte, i [i], 1817: 338, pi. 27, fig. 3) is retained as a distinct 
species by Norman (Proc. zool. Soc. Lond., 1926: 953) and by Rey (Fauna Iberica, Peces, i, 1928: 513, pi. 8, 
figs. 3, 4), it is referred to the synonymy of R. rhinobatos (Linnaeus) 1758 by Fowler (Bull. U. S. nat. Mus., 100 
[jj], 1941: 305). 



Fishes of the JVestern North Athmtic 5 i 

Platifornax (subgenus) Whitley, Aust. Zool., 9, 1939: 244; type species, Rhinobatus thouini Miiller and Henle 
1 841, designated by Whitley. 

Doubtful Synonym: 

Rhinobatis Blainville, in Vieillot, Faune Fran?., 1825: 47, 48; type species, Rhinobatis duhameli Blainville.^' 

Generic Characters. Trunk strongly flattened anteriorly; anterior parts of disc more 
or less wedge-shaped. Width across pectorals considerably less than distance from tip 
of snout to cloaca. Body sector not sharply marked off from tail sector. Snout with 
rounded tip. Tail rounded above, flattened below, with a conspicuous longitudinal 
dermal fold along each side, extending from opposite or a little anterior to origin of 
first dorsal to origin of caudal. Pectorals with anterior rays extending only a little beyond 
nostrils; outer margins evenly convex, without definite corners; posterior corners 
broadly rounded. Dorsals about alike in size and shape, with free rear corners, their 
posterior margins straight or weakly concave. Origin of first dorsal much nearer to 
rear tips of pelvics than to origin of second dorsal. Interspace between dorsals at least 
twice as long as base of first dorsal. Caudal without distinct lower lobe. Pelvics widely 
separated one from the other posteriorly, with nearly straight margins and subangular 
corners. Posterior margin of spiracles with one or two well marked ridges or folds. The 
young of Rhinobatos productus, and perhaps of others, with a fringed membranous flap 
above on the tip of the snout that is lost with growth. Nostrils more or less oblique, 
wholly separate from mouth and well separated one from the other. Anterior margin 
of nostrils expanded as a narrow lobe across middle of nasal aperture, the lobe's inward 
extension widely separated from inner end of nostril, leaving inner part of nasal aperture 
exposed; posterior margin of nostrils also more or less expanded. Mouth transverse, 
nearly straight. Teeth numerous (up to 60-65 series, and perhaps more in some species), 
low, rounded, without cusp in females, but with low blunt cusp in adult males; those 
in each jaw in a pavement-like band. Dermal denticles minute, except for larger tubercles 
or thorns along median line of back, on shoulder regions, around eyes and spiracles, 
and on tip of snout in some species; in the young of some species, a row of thorns 
along each rostral ridge that is lost with subsequent growth. 2* Rostral projection of 
cranium extending to tip of snout, with two longitudinal ridges separated by a median 
furrow (visible externally). Claspers of mature males slender, strongly flattened dor- 
soventrally, their tips simple, without projecting spines or blade. Characters otherwise 
those of the family. 

Size. Maximum length of largest species up to five or six feet. 

27. We see no way of determining whether the name Rhinobatis duhameli Blainville 1825 was based on a Rhinobatos 
from the Mediterranean or on some member of the genus now known as Rhynckobatus from an unknown locality. 
Favoring the last of these alternatives is the fact that Duhamel's (Traite Peches, 4 [2] Sect. 9, 1782: 292, pi. 15) 
illustrations, on which Blainville's account appears to have been based, show its caudal as having two lobes. On 
the other hand, Duhamel's characterization of it as being taken rather often in the Mediterranean, where Rhinobatos 
does occur but where Rhynckobatus does not, suggests that his illustration was of a Rhinobatos with the caudal in- 
correctly represented. 

28. This is true of the eastern Adantic Rhinobatos cemiculus as well as of the Californian-Mexican Rhinobatos productus, 
and perhaps of other species, the young stages of which are not known. 



5 2 Memoir Sears Foundation for Marine Research 

Developmental Stages. The Guitarfishes are ovoviviparous,^^ and it has long been 
known that the inner walls of the uterus in gravid females are thrown into a series of 
thin longitudinal folds. 

Habits. They swim slowly near the bottom, or they lie half buried in the sand 
or mud. We read that in Australian waters "they can be easily approached and picked 
up by the tail."^" When swimming, the muscular tail is used as the organ of propul- 
sion, the pectoral fins to raise or lower the body or to turn and bank. Their food seems 
to consist mainly of small fishes or of crustaceans, shellfish, and other ground-living 
animals, which they crush with their small rounded teeth. Some of the Indian Guitar- 
fishes are said to live in large schools and to do great damage to the pearl oyster beds 
of Ceylon and elsewhere. ^^ 

Relation to Man. In Peru, Guitarfishes are eaten extensively by the poorer people, 
mostly dried or salted, but they are regarded as mediocre in quality. ^^ A few are also 
exposed for sale in tropical fish markets in other parts of the world, some are taken 
to rendering plants in California, while in India the fins of a few are prepared for export 
to China, where they are used for soup, as Shark fins are. They are not of interest to 
anglers and are entirely harmless to bathers. 

Range. Tropical and warm temperate coastal waters of all oceans, running up into 
fresh water locally and even breeding there. ^^ Eastern Atlantic northward to Portugal 
and the Mediterranean; southward along northwestern and tropical western Africa, 
at least to southern Benguela (recorded from Mossamedes, about Lat. i6° S).^* Western 
Atlantic from North Carolina to Uruguay and northern Argentina.^^ Pacific Coast of 
the Americas from middle California (San Francisco) to Peru. Western Pacific from 
mid-China (Shanghai), Japan, and Korea to Queensland, Australia, in about Lat. 23— 
24° S; Philippines; Malayan region as a whole; south along West Australia to about 
Lat. 25° S (Sharks Bay); coasts of the Indian Ocean from Bay of Bengal, Arabian Sea, 
Gulf of Oman, Red Sea, and along the coast of Africa to Natal and to the Cape of Good 
Hope (Table Bay). 

Species. Twenty-six species oi Rhinobatos are recognized in the most recent synopsis 
of the genus^^ and four more have been described subsequently as new.'' But the 

29. See Southwell and Prashad (Rec. Indian Mus., j6, 1919: pi- 18, figs. 1-4) for illustrations of early embryos prior 
to the fusion of the anterior parts of the pectorals with the head; see Ranzi (Publ. Staz. Zool. Napoli, jj, 1934: 
365) for a recent account, with photograph, of the uterine wall. 

30. Whitley, Fish. Aust., 7, 1940: 169. 

31. Norman and Fraser, Giant Fishes, 1937: 59—60. 

32. Fiedler, Jarvis, and Lobell, Pesca y Industr. Pesqu. Peru, Lima, 1943: 285; Hildebrand, Bull. U.S. nat. Mus., 
189, 1946: 51. 

33. Ogilby (Mem. Qd. Mus., 5, 1916: 95, ftn.) describes Guitarfishes in Australia as "freely entering and even perma- 
nently residing and breeding in fresh water." 

34. We find no record of Rhinobatos from the West African Coast between about Lat. 16° S and Table Bay- 

35. Reported to Lat. 40° S (Pozzi and Bordale, An. Soc. cient. argent., 120, 1935: 153). 

36. Norman, Proc- zool. Soc. Lond., 1926: 945. 

37. Rhinobalus banksii McCulloch (Biol. Result. Fish. F. I. S. 'Endeavour', 5, 1926; 157), Australia; Rhinobatos congo- 
lensis Giltay (Ann. Soc. Zool. Belg., sg, 1928: 21), tropical West Africa; Rhinobatos petiti Chabanaud (Bull. Mus. 
Hist. nat. Paris, [2] i, 1929: 365), Madagascar; and Rhinobatos batillum Whitley (Aust. Zool., 9, 1939: 254), Australia. 
Two other recently named species, R. albomaculatus Norman (Ann. Mag. nat. Hist., [10] 6, 1930: 226) and R. ir- 



Fishes of the Western North Atlantic 53 

characters, supposedly specific,^' are so completely intergrading that the genus stands 
in urgent need of revision, which we are not in a position to attempt. 

The Guitarfishes of the western side of the Atlantic fall in the group in which the 
snouts are relatively long, in which the anterior nasal flaps do not extend inward onto 
the internarial space, and in which there are two folds on the posterior margin of the 
spiracles. The great majority of western Atlantic specimens that have been described, 
or that we have seen, are rather evidently in one or the other of the two following 
categories. 

A. Those with a cluster of prominent tubercles on the tip of the snout and with 
the rostral cartilage widening rather noticeably toward its tip. Specimens of this sort 
which have the upper surface of the trunk and tail densely freckled with a large number 
of small pale dots were the basis for the species lentiginosus Garman 1880, described 
from Florida. But the color pattern proves to be less dependable as a specific character 
than has been commonly assumed, for our Study Material includes both unmarked and 
sparsely spotted specimens from Texas that agree with typical lentiginosus in their mor- 
phological features. 

B. Those without prominent tubercles on the snout and with the rostral cartilage 
widening less toward its tip; plain-colored or with indistinct dark markings, or with 
pale spots fewer and larger than in Rhinobatos lentiginosus. The western Atlantic repre- 
sentatives of this category, known from Panama and the West Indies southward to 
northern Argentina, have been described as three species, R. percellens (Walbaum) 
1792, R. horkelii (Miiller and Henle) 1841, and R. stellio (Jordan and Rutter) 1897. 
The first two are separable by the characters listed on pp. 56, 58. But there appears 
to be no distinction between R. stellio and R. percellens (p. 68),^' and it seems doubtful 
whether a sharp line can be drawn between R. percellens and R. lentiginosus^ for we have 
seen one young Texan specimen which had the spatulate rostral cartilage ol percellens 
but which was plain-colored and had the rostral tubercles only faintly indicated (perhaps 
not yet developed). On the other hand, we have seen a Jamaican specimen which was 
typically R. percellens in coloration as well as in shape of rostral cartilage, but which had 
the tip of the snout armed with two rounded tubercles. Clarification of this puzzling situa- 
tion must await further study of the Guitarfishes from the coasts of the Gulf of Mexico. 

The group represented by the eastern Atlantic R. rhinobatos and its immediate 
allies, in which the anterior nasal flap extends inward across the inner margin of the 
nostril and so encroaches upon the internarial space (Fig. 12), has no known counter- 
part in the western Atlantic. 

■vinei Norman (Ann. Mag. nat. Hist., [lo] 7, 1931: 352) from West Africa, with caudals pictured as_ bilobed,_ fal 
properly in Rhinobatos according to Poll (Result. Sci. Exped. oceanogr. Beige Cot. Afr. Atlant. [1948-49], 1951: 

9'. 94)- 

38. Shape of snout and of rostral ridges; degree of inward extension of the flap-like expansions of the anterior margins 
of the nostrils; number (one or two) and prominence of the folds on the posterior margins of the spiracles. 

39. The statement in the original account (Jordan and Rutter, Proc. Acad. nat. Sci. Philad., 4r), 1897: 91) that the 
distance from the origin of the first dorsal to the origin of the second dorsal in K. stellio is equal to the distance from 
the axil of the pectoral to the origin of the first dorsal, was an evident slip, for re-examination of the original spe- 
cimen (for which we have to thank the Natural History Museum of Stanford University) shows that axil of the pelvic 
was intended. 



54 Memoir Sears Foundation for Marine Research 

Finally, one more species, R. spinosus Giinther 1870, must be mentioned because 
of the possibility that the one known specimen may have come from the Gulf of Mexico 
and hence may belong to the Ray fauna of the western Atlantic. It falls with R. rasus 
and with the young of R. cemiculus in the spininess of its rostral ridges. But we are not 
able to judge from the brief description (p. 73) how it is related to these species in 
other respects. 

Provisional Key to North Atlantic, Western South Atlantic, and Tropical 
West African Species 

I a. Anterior nasal flap extending across inner (anterior) margin of nostril as far as 
level of inner end of nostril (Fig. 12 A); rostral ridges separated by a considerable 
interspace throughout their lengths. rhinobatos Linnaeus 1758. 

Portugal, Mediterranean, West 
Africa south at least to Lat. 
16° S." 
lb. Anterior nasal flap extending but little, if at all, across inner margin of nostril 
(Fig. 12B-D); rostral ridges joined or close together anteriorly. 
2 a. Each rostral ridge with a more or less conspicuous row of small sharp thorns. 
3a. Thorns on shoulders arranged one anterior to the other; distance from 
tip of snout to anterior margin of eye a little shorter than distance from 
posterior margin of eye to axil of pectoral (adult). 

cemiculus (St.-Hilaire) 1 8 1 7 (young). 

Mediterranean to tropical West Africa. 

3b. Thorns on shoulder arranged one beside the other; distance from tip of 

snout to anterior margin of eye equal to that from posterior margin of 

eye to axil of pectoral. rasus Garman 1908. 

Tropical West Africa. *' 
2b. Rostral ridges without sharp thorns. 

4a. Rostral cartilage expanded toward tip, spatulate in form; tip of snout 
with a few enlarged tubercles, except perhaps on some small specimens 
(Fig. 14). lentiginosus Garman 1880, p. 60. 

4b. Rostral cartilage not expanded toward tip; tip of snout without enlarged 
tubercles, except on some small specimens (Fig. 1 3 A, C). 
5a. Nostril only i. 0—1.2 times as long as distance between nostrils, and 
little more than half as long as breadth of mouth. 

■percellens Walbaum 1792, p. 68. 
5 b. Nostril at least 1.4 times as long as distance between nostrils and 
about ^/4 as long as breadth of mouth. 

40. Apparently including albomaculatus Norman 1930 and irvinet Norman 1931. 

41. Norman (Proc. zool. See. Lend., 1926: 955) questions whether rasus is actually distinct from cemiculus, to which 
it certainly is closely allied, and Fowler (BuU. Amer. Mus. nat. Hist., yo [i], 1936: 100) united the two. R. spinosus 
Giinther 1870 would also fall under alternative za, if it is an Atlantic species (p. 73). But the only first-hand de- 
scription of it that has appeared is not detailed enough for us to be able to include it in this Key. 



Fishes of the Western N^orth Atlantic 



55 

6a. Distance from tip of snout to level of fronts of orbits about 
equal to distance from rear margins of orbits to axils of pec- 
torals, congolensis Giltay 1928." 

Belgian Congo Coast. 

6 b. Distance from tip of snout to level of fronts of orbits con- 
siderably less than distance from rear margins of orbits to axils 
of pectorals. 







Figure 12. Left-hand nostrils, A Rhinobatos rhinobatos, male, about 720 mm 
long, from Mediterranean (Acad. Nat. Sci. Philad.). B R. lentiginosus. C R. 
perceUens. D R. horkelii. 



7a. A cluster of 3-4 enlarged tubercles on each shoulder, with 
one over outer end of pectoral girdle ; snout in front of mouth 
more than three times as long as breadth of mouth; outer 
fold on posterior margin of spiracle conspicuously larger 
than inner fold. horkelii Muller and Henle 1841, p. ^d. 

7b. Only two tubercles on each shoulder and none over outer 
end of pectoral girdle; snout in front of mouth consider- 
ably less than three times as long as breadth of mouth; 
outer fold on posterior margin of spiracle not conspicuously 
larger than inner fold. 

cemiculus (St.-Hilaire) 18 17 (adult). 
Mediterranean to tropical West Africa. 



42. R. congolensis may prove to be identical with R. cemiculus. 



56 Memoir Sears Foundation for Marine Research 

Rhinobatos horkelii (Miiller and Henle) 1841 

Brazilian Guitarfish 

Figures 12D, 13A, B 

Study Material. Four specimens, male and female, 525 to 840 mm long, from Rio 
de Janeiro, Brazil, in Harvard Museum of Comparative Zoology. 

Distinctive Characters. R. horkelii closely resembles both R. lentiginosus and R. per- 
cellens in general form. However, its nostrils are longer relatively than in either of 
these; its crown is flat transversely or even slightly convex (definitely, if only slightly, 
concave in R. lentiginosus and in R. percellens); its tubercles in the median row are 
larger, more thorn-like, and more numerous; the origin of its first dorsal is posterior 
to the tips of Its pelvics by a distance about equal to the base of the first dorsal (by a 
distance about equal to the extreme length of the first dorsal from origin to rear corner 
in the others); and its teeth are fewer in number than in the other two species. Its plain 
coloration is a field mark which sets it off from most specimens of R. lentiginosus and 
R. fercellens. Its narrower rostral projection distinguishes it further from R. lentiginosus. 
And the anterior nasal flap differs in detail in these three species (Fig. 12). 

Description. Proportional dimensions in per cent of total length. Male, 705 mm, 
and female, 790 mm, from Rio de Janeiro (Harv. Mus. Comp. Zool., No. 544). 

Disc: extreme breadth 33.0, 34.7; length 38.6, 40.3. 

Snout length: in front of orbits 15.2, 16.8; in front of mouth 17.7, 19.2. 

Orbits: horizontal diameter 2.8, 2.7; distance between 3.9, 4.2. 

Spiracles: length 2.1, 1.9; distance between 5.1, ^.z^. 

Mouth: breadth 5.4, 5.8. 

Nostrils: length 3.5, 3.7; distance between inner ends 2.4, 2.5. 

Gill openings : \e.n^t\i?., ist 1.8, 1.6; 3rd 1.8, 1.8; 5th i.i, 1.5; distance between 
inner ends, ist 12.2, 13.0; 5th 8.9, 9.6. 

First dorsal fin: vertical height 6.4, 7.2; length of base 5.1, 4.8. 

Second dorsal fin: vertical height 6.0, 6.8; length of base 5.7, 5.6. 

Caudal fin: upper anterior margin 15.0, 15.6. 

Pelvics: origin to tip 14.5, 16.0. 

Distance: from tip of snout to center of cloaca 40.8, 42.7; from center of cloaca 
to tip of tail 59.2, 57.3; from tip of snout to ist dorsal 56.6, 58.8. 

Interspace between: ist and 2nd dorsals 10.5, 10.2; 2nd dorsal and caudal 
6.3, ^.6. 

Disc about ^e ^s broad as long to posterior limits of pectorals; wedge-shaped 
anteriorly but with well rounded tip, its angle in front of orbits about 50—60°; antero- 
lateral contours slightly concave anterior to level of eyes, posterior margins evenly and 
moderately convex, posterior corners well rounded. Tail from center of cloaca 1.3-1.4 
times as long as distance from snout to cloaca, nearly flat below, moderately rounded 
above, tapering evenly rearward, its breadth opposite axils of pelvics about twice as 



Fishes of the JVestern North Atlantic 



S7 




Figure 13. Rhinobatos horkelii, male, 700 mm long, from Rio de Janeiro, Brazil (Harv. Mus. Comp. Zool., No. 
544). A Anterior part of head in dorsal view. B Ventral view of same. C Anterior part of head, dorsal view, of 
R.percellens, 557 mm long, from Rio de Janeiro, Brazil (Harv. Mus. Comp. Zool., No. 435). D Anterior part 
of head of R. rhinobatos, from Italy (.'^cad. Nat. Sci. Philad.). All about 0.6 X. 



great as distance between spiracles; its lateral dermal folds originating about opposite 
or a little in advance of tips of pelvics and extending as far as lower origin of caudal, 
their width opposite interspace between dorsals (where widest) about Vs ^s great as 
horizontal length of eye. 



5 8 Memoir Sears Foundation for Marine Research 

Dermal denticles minute, close-set, mostly spear-shaped and rounded anteriorly 
over upper surface in general from nuchal region rearward, but some more ovate with 
skin more or less exposed between them; those on crown narrower, fluted anteriorly; 
those on snout flat, roundish to ovate; those on lower surface rounded to tetragonal 
with blunted corners, varying in size, so close-spaced that the skin is nearly or wholly 
concealed. Midline of back rearward from nuchal region with a more or less regular 
row of tubercles, each with sloping median ridge, more prominent than those oi R. lenti- 
ginosus and of R. percellens but irregularly interspersed with smaller; up to about 70 
tubercles to first dorsal fin, about 8 to 12 between dorsals, and a few smaller ones close 
beyond rear end of second dorsal; each shoulder with 2—4 somewhat lower tubercles, 
with one over each outer end of pectoral girdle*^ on small as well as on large specimens; 
4—7 larger and smaller and more conical ones close in front of orbit; 2-6 along inner 
margin of orbit, with one or two close to inner end of spiracle; tip of snout with 2—4 
small rounded tubercles on small and on some medium-sized specimens but without 
tubercles on large ones. 

Snout in front of orbits about 3.7 times as long as distance between orbits in 
young, about 4.0 times as long in large specimens of both sexes; its length in front of 
mouth about 3.0 times as great as width of mouth in young and about 3.3 times as 
great in larger specimens. Horizontal diameter of eye about half as long as distance 
between spiracles. Length of orbit plus spiracle about "Is—^U as great as distance between 
spiracles. Spiracles about V4 as long as orbit, nearly transverse, posterior margin with 
two well marked folds or ridges, the outer the longer. First to fourth gill openings of 
about equal lengths, about half as long as nostril, the fifth a little shorter; distance 
between inner ends of fifth gill openings about 1.6 times as great as breadth of mouth. 
Nostrils moderately oblique, their length about 1.5 times as great as distance between 
them and about ^/^ (63—66 "/o) as great as width of mouth; anterior margin expanded 
near the middle as a narrow lobe with rounded tip, curving outwardly and across nasal 
aperture and extending inward a short distance as a narrow fold that ends abruptly 
without encroaching on the internarial space; posterior (outer) margin of nostril ex- 
panded as three confluent flaps, two of them shorter and directed rearward, the third 
longer, digitate, originating about opposite anterior nasal flap and extending along 
nasal aperture about 40 '/o of the distance from its own origin toward inner corner of 
nostril. 

Teeth l^^l in specimens 515—790 mm long, those of immature specimens close- 
set in quincunx; low, oval, the longer axis transverse, with indistinct transverse cutting 
edge; the base extending a little rearward much as in R. Ientigiyiosus\ those of mature 
males not seen. 

First and second dorsals about equal in size, triangular, with abruptly rounded 
corners, their anterior margins straight or slightly convex, the posterior margins straight 
or slightly concave and approximately vertical, the free rear corner about V2— ^/s ^s long 

43. The type specimen had no tubercles over the outer ends of the pectoral girdle (Miiller and Henle, Plagiost., 1841 : 
122), but they are present on all that we have seen. 



Fishes of the Western North Atlantic 59 

as the base; origin of first dorsal posterior to tips of pelvics by a distance about equal 
to base of first dorsal or a little greater. Interspace between dorsals about twice as 
long as base of first dorsal. Interspace between rear end of base of second dorsal and 
upper origin of caudal about i. 0—1.2 times as long as base of second dorsal. Upper 
origin of caudal a little anterior to lower; upper margin weakly convex, about i — 1.5 
times as long as interspace between dorsals; lower posterior contour well rounded, 
without distinct lower lobe; height of caudal above extremity of axis about as great 
as its depth below latter. Caudal axis only slightly raised (perhaps horizontal in some 
cases). Pelvics with anterior, inner and distal margins nearly straight, the outer corners 
broadly rounded, forming an angle of about 1 30°, the posterior corners narrowly 
rounded; extreme length of pelvics, origin to rear tip, about 1.6-1.8 times as great 
as distance from origin of pelvic to axil; their axils separated by a distance about 
1. 2-1. 4 times as great as breadth of mouth. 

Rostral cartilage nearly uniform in breadth throughout most of its length, widening 
in somewhat rounded outline toward tip, its maximum breadth there about i.o— i.i 
times as great as distance between inner ends of nostrils; its ridges close together and 
approximated at tip. Anterior rays of pectorals extending anterior to nasal capsules of 
cranium by a distance about 1.3— 1.4 times as great as distance between nostrils. 

Color. Upper surface uniform olive gray or chocolate brown, without pale or dark 
markings. Lower surface either a pale shade of the same hue as upper surface or as 
dark as upper surface;" snout with an oval sooty patch in some but only faintly washed 
with sooty in others. *5 

Relation to Extralbnital Species. R. horkelii closely resembles R. cemiculus St.-Hilaire 
1827 of the eastern Atlantic but appears to be separable from it by a longer snout 
and by the facts that the outer of the two folds on the posterior margin of its spiracle 
is conspicuously larger than the inner and that there are 3—4 tubercles on each of its 
shoulders (only two in R. cemiculus)\ furthermore, horkelii seems to be separable from 
cemiculus by the presence of a larger number of tubercles (about 70; see p. 49) along 
the midline of its back from nuchal region to first dorsal fin (only about 18-20 in 
R. cemiculus).^^ 

R. horkelii is closely allied to R. productus of the Pacific Coast of Mexico and 
California, but when half-grown or larger it is separable from R. productus by the fact 
that the origin of its first dorsal fin is posterior to the tips of the pelvics by a distance 
about as great as that between the eyes or about as long as the base of the first dorsal 
(less than half as long as distance between eyes and only a little more than half as long 
as base of first dorsal in R. productus). The tubercles around the eyes and on the shoulders 
are more prominent in R. horkelii than in R. productus, those in the median dorsal row 
larger and more regularly arranged, especially in the interspace between the two dorsal 

44. Specimens with pale undersurfaces were classed as a color variety in the original account of the species. Those that 
we have seen are paler below than above, but evidently they have lost most of their original color. 

45. Our Study Material includes examples of each of these color phases. 

46. The original illustration of R. cemiculus (St.-Hilaire, in Savigny, Zool. Egypte, i [i], 1827: pi. 27, fig. 3), modified 
by Fowler (Bull. Amer. Mus. nat. Hist., yo [i], 1936: too, fig. 37), shows only 18. 



6o Memoir Sears Foundation for Marine Research 

fins. And the nostrils are up to 1.5 times as long as the distance between them in some 
R.horkelii but not more than 1.3 times that in R. productus. It is not possible to make 
a comparison of the younger stages for lack of information about R. horkelil. 

Size. Evidently this is the largest Guitarfish of the western Atlantic, for the 
claspers of a male 700 mm long still fall a little short of the tips of the pelvics. A female 
in our Study Material, 840 mm long, is the largest yet reported; the size at maturity 
is not known. 

Developmental Stages. The embryos have not been seen, nor have we seen new- 
born young. 

Habits. Nothing is known. 

Range. Coast of Brazil, perhaps north to the Lesser Antilles. Although R. horkelii 
has been known to science for more than half a century, and although it is easily dis- 
tinguishable from both of the other species of its genus that occur in the western Atlantic, 
all that is known about its geographic distribution is that a few were collected many 
years ago^' at Rio de Janeiro and at Bahia, Brazil, and that one small specimen has 
been reported more recently from the Lesser Antilles (St. Eustatius i'), but without 
sufficient information to establish its specific identity with certainty. 

Synonyms and References: 

Rhinobatus {Rhinobatus) horkelii Miiller and Henle, Plagiost., 1841 : 122, pi. 41 (descr., meas., color, ill., Brazil), 
192 (color); Dumeril, Hist. Nat. Poiss., I, 1865: 499 (descr., Bahia, Brazil). 

Rhinobatus horkelii'^^ Castelnau, Anim. Nuov. Rares Amer. Sud., Poiss., 1855: 100 (listed, Bahia, Brazil); 
Giinther, Cat. Fish. Brit. Mus., 8, 1870: 444 (cf. R. undulatus)\ Garman, Proc. U. S. nat. Mus., 3, 
1880: 518 (descr., color, Brazil); Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907: 175, 207 (diagn., refs., 
Rio de Janeiro and Bahia, Brazil); Engelhardt, Abh. bayer. .Akad. Wiss., Suppl. 4 (3), 191 3: loi (range); 
Garman, Mem. Harv. Mus. comp. Zool., j6, 1913: 280 (descr., cf. R. pcrcellens, Rio de Janeiro); 
Metzelaar, Trop. Atlant. Visschen, 1919: 7 (color, W.Indies, St. Eustatius.'); Ribeiro, Arch. Mus. 
nac. Rio de J., Fauna Brasil., Peixes, 2(1) Fasc. l, 1923: 31 (diagn., color, cf. R. percellens, Bahia, Brazil); 
Norman, Proc. Zool. Soc. Lond., 1926: 947 (in Key), 976 (descr., ill. head, Brazil); Jordan, Evermann 
and Clark, Rep. U. S. Comm. Fish. (1928), i, 1930: 23 (listed, Brazil to St. Eustatius, Windward Is.). 

Rhinohatos lentiginosus (Garman) 1880 

Spotted Guitarfish 

Figures 8, 12B, 14, 15 

Study Material. Thirteen specimens, male and female, 350—577 mm long, from 
Progresso, Yucatan; Freeport, Texas; Pensacola, Key West, Palmetto Key, Pine Island 
Sound, Captiva Pass, and New Smyrna Beach, Florida; Charleston, South Carolina; 
and Cape Lookout, North Carolina; embryo 180 mm long, nearly ready for birth, 
from Charleston, South Carolina; also 13 specimens from Texas which apparently 
represent a color variant of this species; all in the collections of Harvard Museum of 
Comparative Zoology, Museum of Zoology at University of Michigan, and U. S. 
National Museum. 

47. Those in our Study Material were collected between 1859 and 1872. 48. Also spelled horkeli. 




Figure 14. Rhinobatos lentiginosus. A Female, 577 mm long, from Captiva Pass, Florida (Harv. Mus. Comp. 
ZooL, No. 35857). B Upper surface of tip of snout of same to show tubercles, about i.6x. C Male, 508 mm 
long, from Captiva Pass (Harv. Mus. Comp. Zool., No. 35857). Z) Side view of tail of same. E Lower surface 
of tip of snout of female 518 mm long, from New Smyrna Beach, Florida (Harv. Mus. Comp. Zool., No. 36408). 



62 Memoir Sears Foundation for Marine Research 

Distinctive Characters. The peculiar shape of the disc and the long wedge-shaped 
snout, combined with the stout muscular tail and its two well developed dorsal fins, 
mark Rhinobatos lentiginosus off from all other batoids of the western Atlantic except 
its genus mates R. percellens and R. horkelii (see discussion, p. ^■;^'). The spatulate rostral 
cartilage of R. lentiginosus, its relatively shorter nostrils, and the fact that the tubercles 
along the midline of its back are smaller and less thorn-like in shape, set it apart from 
R. horkelii \ so too, the pale-spotted color pattern of specimens that are marked in that 
way. Its tubercle-tipped snout and more broadly spatulate rostral cartilage differentiate 
it from R. percellens, while the color pattern differs in detail between such specimens 
of lentiginosus and percellens as are marked with pale spots (p. 68). 

Description. Proportional dimensions in per cent of total length. Male, 508 mm, 
from Captiva Pass, Florida (Harv. Mus. Comp. Zool., No. 35857). Female, 577 mm, 
from Pine Island, Florida (Harv. Mus. Comp. Zool., No. 35856). 

Disc: extreme breadth 29.9, 33.0; length 39.0, 41.7. 

Snout length: in front of orbits 14.2, 14.0; in front of mouth 17.3, 17.3. 

Orbits: horizontal diameter 2,-Si 3*4' distance between 3.1, 2.9. 

Spiracles: length 2.0, 2.0; distance between 5.1, 5.2. 

Mouth: breadth 6.4, 6.2. 

Nostrils: length 3.8, 3.6; distance between inner ends 3.3, 3.1. 

Gill openings : lengths, ist 1.3, 1.2; 3rd 1.4, 1.5; 5th i.o, i.o; distance between 
inner ends, ist 12.4, 12.7; 5th 8.2, 9.0. 

First dorsal fin: vertical height 6.3, 6.4; length of base 4.8, 4.5. 

Second dorsal fin: vertical height 6.6, 6.5; length of base 5.3, 4.9. 

Caudal fin: upper anterior margin 14.9, 13.5. 

Pelvics: origin to tip 16.5, 15.2. 

Distance: from tip of snout to center of cloaca 41.3, 45.2; from center of cloaca 
to tip of tail 58.7, 54.8; from tip of snout to ist dorsal 59.3, 60.7. 

Interspace between: ist and 2nd dorsals 1 1.2, 10. o; 2nd dorsal and caudal 5.9, 5.9. 

Disc about V4~V6 ^^ broad as long to posterior limits of pectorals; wedge-shaped 
anteriorly but with well rounded tip, its angle in front of orbits about 50° ; posterior 
corners well rounded; anterolateral contours nearly straight in young specimens but 
slightly concave anterior to the level of eyes in large; posterior margins evenly and 
moderately convex; posterior corners of pectorals overlapping pelvics by a distance 
about as long as that between nostrils. Tail from center of cloaca 1.2— 1.4 times as 
long as distance from snout to cloaca, nearly flat below, moderately rounded above, 
tapering evenly rearward, its breadth opposite axils of pelvics about twice as great as 
distance between spiracles; its lateral dermal folds originating opposite tips of pelvics 
and extending to lower origin of caudal, their width opposite interspace between dorsals 
(where widest) about Ys ^s great as horizontal length of eye. 

Dermal denticles on upper surface minute and close-set, but skin exposed between 
them; low, flattish or slightly domed; those along midzone of back ovate posteriorly, 




Figure i 5. Rhinobatos lentiginosus. A Eye and spiracle of female, about 2X. B Nostril of male, about 2 X • C Cross 
section of posterior part of trunk of female at level of first dorsal fin, about 1 .0 X . D Dermal denticles along 
midline of back of female between nuchal region and pectoral girdle, anterior ends uppermost, about 10 X. 
E Dermal denticle of female from beside mid-dorsal line, in dorsal, anterior, and lateral views, about 38 X. 
F Dermal denticles of female from along upper margin of orbit, about lox. G Denticles on side of snout of 
male, about 24 X. ^ Denticles on lower side of pectoral fin of male, about 24 X. ^ Denticles on upper lip 
(smaller) and teeth (larger) from middle of jaw of female, about 14 X . A' Cross section of upper jaw of same to 
show denticles on upper lip (smaller) and teeth of successive rows (larger), about 14X. /. Denticles on upper 
lip (smaller) and teeth (larger) of mature male, about 14 X. A, C-F, J, K from female, 577 mm long, from 
off Useppa Island, Florida (Harv. Mus. Comp. Zool., No. 35856). B, G, H, L from mature male, 508 mm 
long, from Captiva Pass, Florida (Harv. Mus. Comp. Zool., No. 35857). 



64 Memoir Sears Foundation for Marine Research 

of various sizes, but irregularly fluted or arrowhead-shaped anteriorly; those on either 
side of rostral ridge spear-shaped and sharp-pointed with concave margins; those along 
sides of tail generally ovate and more or less pointed; those along outer margins of 
pectorals and of snout, around orbits, and along margins of spiracles, smaller, more 
rounded, and more closely crowded. Usually 33—37 larger tubercles along midline of 
back, from nuchal region to first dorsal; similar to smaller scales in shape, or more 
pointed posteriorly, but varying widely in prominence (apparently irrespective of age or 
sex); thorn-like on some but so slightly conspicuous on others as to be detected only 
by touch or by examination with a lens; eight or nine smaller tubercles in interspace 
between dorsal fins on some specimens ; some with one or two small tubercles on scapular 
region on one side or on both; anterior and posterior sectors of orbital ridge with 
enlarged flattish denticles in some cases. Tip of snout of typical specimens conspic- 
uously armed with blunt conical tubercles or thorns, varying in number from 2-7 
or perhaps even more in later stages of growth; see discussion (p. 53). Lower surface 
as a whole clothed with minute, smooth, flattish, or slightly rounded denticles irreg- 
ularly hexagonal or tetragonal in shape, and so closely crowded that the skin is entirely 
concealed. 

Snout in front of orbits about 4.0 times as long as distance between orbits in young, 
about 5.0 times in large females, and 4.5 times in large males; its length in front of 
mouth about 3.1 times as great as width of mouth in young and 2.7—2.9 times in larger 
specimens of both sexes. Horizontal diameter of eye about ^s as long as distance between 
spiracles, or a little longer. Length of eye plus spiracle about ^U—*U ^^ great as distance 
between spiracles. Spiracles about ^/j as long as orbit, nearly transverse, about -/j as 
long as eyes; posterior margin with two well marked folds or ridges, the outer the 
larger. First to fourth gill openings about equal in length, about 1/3 as long as nostril, 
the fifth a little shorter; distance between inner ends of fifth gill openings 1.3— 1.5 times 
as great as breadth of mouth. Nostrils moderately oblique, their length about as great 
as distance between them or a little greater, and somewhat more than half (57-58 "/o) 
as great as width of mouth; anterior margin expanded midway as a narrow lobe with 
rounded tip, curving outward and across nasal aperture and extending inward a short 
distance as a narrow fold that ends abruptly without encroaching on the internarial 
space; margins of nostril about as in R. horkelii (p. 58) except that digitate flap of 
posterior margin extends along nasal aperture about 70 "/o of distance toward inner 
corner of nostril. Mouth nearly straight with slight bow centrally, its corners with 
small wrinkles. Roof of mouth, close behind tooth-band, with a narrow transverse fold 
that widens as a low rounded lobe on either side and extends the whole breadth of 
mouth; its anterior surface sculptured with many low rounded prominences about 
equal in size to the teeth, its free edge conspicuously fringed. 

Teeth in quincunx, 51^!^ in specimens 390—580 mm long, alike in form in the 
two jaws and in the two sexes; closely crowded, rectangular to pentagonal basally, 
with rounded corners, the functional surface a little elevated in subangular form along 
the transverse axis; the younger rows with a low, blunt, conical cusp which is lacking 



Fishes of the Western North Atlantic 65 

on most of the older teeth, perhaps as the result of wear; eight to ten rows in function 
simultaneously in upper jaw and seven to nine in lower in large specimens. 

First and second dorsals approximately equal in size, triangular, with their cor- 
ners abruptly rounded, their anterior and posterior margins nearly straight and the 
latter approximately vertical, the free rear corners a little less than half as long as the 
base; origin of first dorsal posterior to tips of pelvics by a distance about equal to 
breadth of mouth or to extreme length of first dorsal from origin to rear corner. Inter- 
space between dorsals about twice as long as base of first dorsal. Interspace between 
rear end of base of second dorsal and upper origin of caudal about 1.3 times as long 
as base of second dorsal. Caudal with upper origin a little anterior to lower; its upper 
margin weakly convex, about 1.3 times as long as interspace between dorsals, its axis 
only slightly raised (perhaps horizontal in some cases); its lower posterior contour well 
rounded without distinct lower lobe; its height above extremity of axis about as great 
as its depth below latter. Pelvics with weakly convex or nearly straight anterior (outer) 
margins, nearly straight inner margins, and weakly convex distal margins; the outer 
corners moderately rounded, forming an angle of about 130-135°; the posterior 
corners abruptly rounded, the tips much more slender in adult males than in females 
or in young males; extreme length of pelvics from origin to rear tip about 1.9-2.0 
times as great as distance from origin of pelvics to axil ; their axils separated by a dis- 
tance about equal to breadth of mouth. 

Rostral cartilage nearly uniform in breadth throughout most of its length but widening 
noticeably in somewhat rounded outline toward tip, its maximum breadth there about 
1.3 times as great as distance between inner ends of nostrils, its ridges close together 
and approximated at tip. Anterior rays of pectorals extending anterior to nasal capsules 
of cranium by a distance about half as great as distance between nostrils. 

Color. Ashy gray to olive brown or chocolate brown above, the area on either side 
of rostral cartilage pale (probably translucent in life); edges of pectorals paler than 
general ground tint but with a slightly darker band inward from margin; pelvics pale 
or whitish-edged, distinctly so in young; dorsals and caudal not noticeably darker. 
Typically, the entire upper surface of the disc rearward from the orbits (including 
posterior half of rostral ridges) is thickly freckled with several hundred small whitish 
dots, as is the upper surface of the tail; there are a few dots on the pelvics but none 
on the dorsals or caudal. But we have seen one Texas specimen with only a few pale 
dots and 12 others without any pale markings though agreeing with R. lentiginosus in 
other aspects. Lower surface usually pale yellow or yellowish white or plain white, the 
fins slightly darker than trunk, a V-shaped dusky or dark gray area below snout per- 
sisting throughout growth in some specimens but becoming diffuse in others; some 
specimens almost as dark in general over lower surface as above.*' The color pattern 
of pale-spotted specimens is described as distinct in the young before birth. ^^ 

Relationship to Extralimital Species. R. lentiginosus is separable from all members 

49. Three of the unspotted Texas specimens we have seen are dark below, the others clear white. 

50. Jordan and Gilbert, Proc. U.S. nat. Mus., 5, 1883: 582. 



66 Memoir Sears Foiindatio?i for Marine Research 

of its genus known from the eastern side of tlie Atlantic by the characters summarized 
in the preceding Key. Among the species of the west coast of America, it differs sharply 
from R. flaniceps (Garman) 1880 in having two well developed folds on the posterior 
margin of each spiracle;*^ from R. glaucostigmus Jordan and Gilbert 1883 and from R. 
leucorhynchus Gunther 1866 in the fact that its rostral ridges are closely approximated 
along their anterior third to half. In this respect, R. lentiginosus more closely resembles 
R. productus Girard 1855, the common Guitarfish of southern California. But there is 
no likelihood of confusing the two, for the rostral cartilage is considerably narrower 
toward its tip in R. productus (maximum breadth only about as great as distance be- 
tween inner ends of nostrils), the first dorsal originates considerably closer to the 
pelvics,^^ and the tip of the snout lacks the large tubercles that are characteristic of 
R. lentiginosus. 

Remarks. All specimens of R. lentiginosus reported thus far from Florida and from 
northward on the east coast of the United States have shown the spotted pattern, as 
does one in our Study Material from Yucatan. On the other hand, all those from 
Texas that we have seen are plain-colored, except for one with a few pale dots. It may 
finally prove that the plain-colored and the spotted forms deserve recognition in nomen- 
clature, whether as color varieties or possibly as subspecies. But we think it premature 
to burden ichthyological literature with an additional name before it is known whether 
the ranges of the two forms are discontinuous, or whether they meet and perhaps inter- 
grade along the coast of Louisiana, whence one or the other of them, or both, are to 
be expected though not yet actually reported. 

Size. This Guitarfish is said to grow to a length of "several feet,"^^ but the largest 
size which we find actually recorded (two females) is 30 inches. Males of 1 9—20 inches 
already have the claspers well developed (Fig. 14 C). 

Developmental Stages. As noted above, the characteristic color pattern may be 
developed before birth. A gravid female has been recorded as containing six young. 

Habits. In Florida waters, the only region where more than occasional specimens 
of R. lentiginosus have been taken, they are often encountered in shallow water around 
the Florida Keys and along beaches. They have even been observed at Palm Beach, 
Florida, moving along at low tide with "dorsals and caudals clear of the water, and 
every few minutes they would poke their snouts up onto the little foot-high shelf that 
was just wet by the small advancing and receding billows" while feeding.^* But they 
are also reported as common off Florida in depths as great as 5—10 fathoms.^* They 
may produce young anywhere within their geographic range, for a female with well 
developed embryos has been taken as far north as Charleston, South Carolina. Nothing 
is known of their habits beyond what applies to the genus as a whole, nor of their 
diet, though it is probable that when they are searching along the intertidal zone on 

51. One spiracular fold only in R. planiceps, two in all other eastern Pacific members of the genus. 

52. Origin of first dorsal is posterior to tips of pelvics by a distance only about half as great as width of mouth or a 
little more than half as great as length of base of first dorsal in R. productus, but by a distance about as great as 
width of mouth or about as great as extreme length of first dorsal from origin to rear corner in R. lentiginosus. 

53. Henshall, Bull. U.S. Fish Comm., 14, 1895: 210. 54. Barbour, Copeia, 85, 1920: 71. 



Fishes of the IVestern North Atlantic 67 

sandy beaches, as described above, they are feeding on sand-dweUing Crustacea, such 
as gammarld amphipods, or Hippa. 

Relation to Man. This species is neither commercially important nor of interest 
to anglers. 

Range. Western Atlantic in coastwise waters from Yucatan to Cape Lookout, 
North Carolina. The great majority of published records for this species have been 
for captures from Florida, ^'^ where it occurs generally (not uncommonly by local report) 
along both coasts, presumably as a year-round resident. A few wander northward in 
spring and early summer along the Atlantic Coast, perhaps yearly, for it is "well- 
known to fishermen" at Charleston, South Carolina.^" Also, it has been taken occa- 
sionally on the North Carolina Coast. 5' But there is no reason to suppose that It ever 
passes Cape Hatteras, except perhaps as a stray. 

The northward distribution of R. lentiginosus along the Atlantic Coast, combined 
with the presence ot specimens from Texas that appear to represent a color variant of it 
(see Study Material, p. 60), makes its occurrence probable all along the northern 
and northwestern shores of the Gulf of Mexico. However, present indications are that 
its range is as sharply limited equatorward as it is northward in the Atlantic, for the 
only report of it from farther south than Florida and Texas is of two specimens from 
Progresso, Yucatan. ^^ And typical R. lentiginosus is so easily recognizable by its con- 
spicuous color pattern, thorny snout, and spatulate rostral cartilage that it is not likely 
to have been overlooked among the representatives of its genus that have come under 
observation from Atlantic Panama, from Jamaica, and from Brazil (pp. 60, 71). 

Synonyms and References: 

Rhinobatus lentiginosus Garman, Bull. Mus. comp. Zool. Harv., 6, 1880: 168 (descr., cf. other species, size, 
Florida); Proc. U. S. nat. Mus., J, 1881 : 519 (descr., cf. other species, Florida and S. Carolina); Goode 
and Bean, Proc. U. S. nat. Mus., 5, 1882: 240 (listed, Gulf of Mexico); Jordan and Gilbert, Proc. U. S. 
nat. Mus., 5, 1883: 582, 619 (descr., color, female with embryos, Charleston, S. Carolina); Bull. U. S. 
nat. Mus., 16, 1883: 65 (descr., Florida); True, List Vert. Anim. S. Carolina, in Handb. S. Carolina, 
1883: 261 (listed, S. Carolina); Jordan, Proc. U. S. nat. Mus., 7, 1884: 148, 149 (listed, Florida Keys); 
Rep. U. S. Comm. Fish. (1885), 1887: 798 (listed, W. Indian Fauna, including Florida Keys); Lonn- 
berg, Oefvers. Svensk. Vet. Akad. Forh., Arg. 51 (3), 1894: 11 1 (no. rostral spines, Florida Keys, Clear- 
water Bay); Henshall, Bull. U. S. Fish Comm., 14, 1895: 210 (size, Florida Keys, Tampa); Jordan 
and Evermann, Rep. U. S. Comm. Fish. (1895), 1896: 220 (listed, Charleston, S. Carolina southward); 
Bull. U. S. nat. Mus., 47 (i), 1896: 62 (descr., Charleston, S. Carolina, Florida); Evermann and Ken- 
dall, Rep. U. S. Comm. Fish. (1899), 1900: 49 (Florida); Jordan and Evermann, Bull. U. S. nat. Mus., 
47 (4), 1900: pi. 8, fig. 28, pi. 9, figs. 2 8a-28b (ill.); Evermann and Goldsborough, Bull. U. S. Fish 
Comm., 21, 1902: 139 (listed, Progresso, Yucatan); Jordan, Guide to Study Fish., j, 1905: 551, fig. 
343 (ill., Florida, S. Carolina); Smith, Bull. N. C. geol. econ. Surv., 2, 1907; 40 (descr., ill., N. Carolina); 
Gill, Smithson. misc. Coll., ^2 (2), 1908: 158 (body form, ill.); Coles, Bull. Amer. Mus. nat. Hist., 
J2, 1913: 33 (female with eggs, C. Lookout, N. Carolina); Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 

55. It has been reported from Tampa, and from Clearwater Bay nearby, Captiva Pass, Lemon Bay, and Punta Rassa 
on the west coast, from various localities in the region of the Keys, and from Palm Beach and New Smyrna Beach 
on the east coast. 

56. Jordan and Gilbert, Proc. U.S. nat. Mus., 5, 1883: 582. 

57. One definite report for Beaufort, North Carolina, one for Morehead City, and four specimens from Cape Lookout 
in July of 1912 and 1913. 

58. Evermann and Goldsborough, Bull. U.S. Fish Comm., 2/, 1902: 139; see also our Study Material. 



6 8 Memoir Sears Foundation for Marine Research 

4 (3), 1913: loi (range); Garman, Mem. Harv. Mus. comp. Zool., j6, 1913: 279, pi. 17 (descr., 
ill., Florida); Gudger, Proc. biol. Soc. Wash., 26, 191 3: 98 (listed, Beaufort, N. Carolina); Coles, Proc. 
biol. Soc. Wash., 28, 1915: 92 (nos., size, season, C. Lookout, N. Carolina); Radcliffe, Bull. U. S. Bur. 
Fish., 34, 1916: 269 (descr., ill., teeth, denticles, C. Lookout, N. Carolina); Barbour, Copeia, 85, 1920: 
71 (habits. Palm Beach, Florida); Meek and Hildebrand, Field Mus. Publ. Zool., 25 (i), 1923: 70 
(color, cf. R.percellens, not recorded for Atlant. Panama); Norman, Proc. zool. Soc. Lond., 1926: 947 
(in Key), 969 (descr., ill., Morehead City, N.Carolina); Breder, Field Bk. Mar. Fish. Atlant. Coast, 
1929: 28 (genl.); Jordan, Manual Vert. Anim. NE U. S., 1929: 16 (diagn., N. Carolina south); Jordan, 
Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 23 (listed, Charleston, S. Carolina- 
Yucatan); Bere, Amer. Midi. Nat., l"], 1936: 582 (parasites. Lemon Bay, Florida); Fowler, Monogr. 
Acad. nat. Sci. Philad., 7, 1945: 98, 160 (listed, N. and S. Carolina). 

Rhinobatos percellens (Walbaum) 1792 

Southern Guitarfish 

Figures 1 2 C, 16 

Study Material. Nineteen specimens, male and female, 210 to 591 mm long, in 
Harvard Museum of Comparative Zoology and U. S. National Museum, from Santos, 
Rio de Janeiro and Pernambuco, Brazil; Colon, Panama; and Jamaica. 

Distinctive Characters. Rhinobatos percellens closely resembles both R. lentiginosus 
and R. horkelii in general form and in most of its proportionate dimensions. It is dis- 
tinguishable from horkelii by its somewhat shorter nostrils; also by the facts that its 
crown is definitely, though only slightly, concave transversely, and that its tubercles 
along the midline of the back are smaller and less thorn-like in form. It differs from 
R. lentiginosus^^ chiefly in that there are no enlarged tubercles on the tip of the snout 
in typical specimens (several such tubercles in R. lentiginosus)\ on specimens oi per- 
cellens that are white-spotted, the spots are not only much less numerous than on most 
specimens of R. lentiginosus but are symmetrically arranged on either side of the median 
line; and the rostral cartilage is narrower in percellens than in lentiginosus., especially 
toward the tip. We have seen three specimens intermediate between the two species: 
one from Texas, which has tubercles on its snout (as in R. lentiginosus) but has no 
white spots (p. 65), and two from Jamaica, sparsely white-spotted (as in R. percellens) 
but with a pair of somewhat enlarged tubercles side by side on the tip of the snout 
(as in R. lentiginosus). 

Description. Proportional dimensions in per cent ot total length. Male, 557 mm 
(Harv. Mus. Comp. Zool., No. 435) and female, 591 mm (Harv. Mus. Comp. Zool., 
No. 542) from Rio de Janeiro, Brazil. 

Disc: extreme breadth 30.5, 32.3; length 38.8, 40.0. 

Snout length: in front of orbits 14.7, 14-7; in front of mouth 16.7, 17.1. 

Orbits: horizontal diameter 3.6, 3.4; distance between 3.2, 3.1. 

Spiracles: length 2.2, 2.3; distance between 5.0, 5.2. 

Mouth: breadth 6.3, 6.5. 

Nostrils: length 3.7, 3.2; distance between inner ends 3.1, 3.4. 

59. Perhaps the only significant differences that can be relied upon. 



Fishes of the Western North Atlantic 



69 




Figure 16. Rhinobatos percellens. A Female, from Rio de Janeiro, about 560mm long (U. S.Nat. Mus., No. 
79306). B Lower side of snout of another specimen of about same size (Harv. Mus. Comp. Zool., No. 435). 



yo Memoir Sears Foundation for Marine Research 

GUI openings: lengths, ist 1.4, 1.4; 3rd 1.6, 1.5; 5th i.i, 1.2; distance between 

inner ends, ist 11.6, 12.3; 5th 7.7, 8.6. 
First dorsal fin: vertical height 6.2, 6.8; length of base 4.7, 5.1. 
Second dorsal fin: vertical height 6.5, 6.7; length of base 4.9, 5.1. 
Caudal fin: upper anterior margin 15.1, 15.4. 
Pelvics: origin to tip 15.1, 15.4. 
Distance: from tip of snout to center of cloaca 41.8, 42.5; from center of cloaca 

to tip of tail 58.2, 57.5. 
Interspace between: ist and 2nd dorsals 11.3, 11.3; 2nd dorsal and caudal 

6.1, 5.8. 

R.percellens resembles R. lentiginosus so closely in appearance, in proportional 
dimensions generally, in shapes and locations of fins, and in the shapes of the dermal 
denticles on different parts of the body, that the points of difference alone need be 
noted. These are: Rostral cartilage narrower, less spatulate in form, its maximum breadth 
near tip of snout only about as great as distance between inner ends of nostrils, the two 
ridges close together and parallel for a little more than half their length but diverging 
a little rearward. Upper tip of snout without enlarged tubercles (but see p. 68). The 
outer of the two folds on the posterior margin of spiracle averaging a little longer re- 
latively. Nostril I— 1.2 times as long as distance between nostrils; the inward extension 
of anterior nasal flap ending more abruptly, perhaps averaging a little shorter relatively 
(Fig. 12). White dots on upper surface much less numerous (when present) than on 
spotted specimens of lentiginosus and arranged in symmetrical pattern on both sides of 
midline; also, the majority of specimens more or less definitely marked with dark 
blotches or incomplete crossbars. 

The enlarged tubercles low, either rounded transversely or with median ridge 
sloping forward; about 36-46 in midline of back from nuchal region to first dorsal 
in adults, the smaller ones irregularly interspersed with larger ones; about 3—7 in inter- 
space between first and second dorsal fins; 1-3 on each shoulder; -^-d close in front 
of orbit, with 2—7 of various sizes around inner margin of orbit to inner end of spiracle; 
none over outer end of pectoral girdle. 

Color. Olive gray, reddish brown, or chocolate brown above; an extensive pale 
translucent area on either side of rostral ridges; sides usually with darker brown spots 
or blotches, vaguely outlined and varying in number, often forming indistinct cross- 
bars on tail ; dorsals and caudal partly dusky, but outer margins of pectorals paler than 
general ground tone; most specimens with up to about 40—45 indistinct whitish spots 
on either side of midline, about as large as pupil, irregularly distributed, but arranged 
symmetrically on the two sides of trunk. Lower surface pale yellowish, greyish or dusky 
white, the tip of the snout with a more or less conspicuous sooty blotch, either solid 
or more or less interrupted; diamond-shaped in young specimens but usually spreading 
rearward with growth along margins of head as illustrated in Fig. 16B. 

Relation to Extralimital Species. Among eastern Atlantic species, R.percellens 



Fishes of the Western Nort/i Atlantic 7 1 

resembles R. cemiculus the most closely. But there is no danger of confusing young 
specimens of the two, for R. cemiculus has a row of sharp spines along each rostral 
ridge whereas R. ferccllens does not. Older R. percellens appear to be separable from 
R. cemiculus by a somewhat shorter nostril relatively, by a larger number of tubercles 
in the mid-dorsal row (only about 17—20 in R. cemiculus, see p. 50, footnote 26), by 
somewhat more prominent spiracular folds, by somewhat lower dorsals relative to their 
lengths, by less prominent tubercles on the shoulders, and usually by the pale-spotted 
and dark-clouded upper surface (R. cemiculus is plain-colored above). 

R. percellens, when fully grown, resembles R. productus of the Pacific Coast of 
Mexico and California, but it is separable from R. productus by the fact that the origin 
of its first dorsal is posterior to the tips of the pelvics by a distance about as great as 
the distance between the outer ends of its spiracles (less than half that great in R. pro- 
ductus). Less conspicuous differences are: tubercles in median row more regular in R. 
percellens, mouth a little less arched, and nostril a little shorter. Young of the two differ 
more noticeably; those of R. percellens have more numerous but much smaller mid- 
dorsal thorns and lack the fleshy fringe on the upper tip of the snout, a feature that 
is conspicuous in the young of R. productus. However, it is not until later in growth 
that the difference in the position of the first dorsal relative to the pelvics develops in 
the two. Also, R. productus is considerably the larger of the two, growing to a length 
of more than four feet. 

Size. The claspers of one male in our Study Material, 560 mm long (about 22 in.), 
appear to be fully developed or nearly so. The largest specimens reported were 31^" 
and about 39 inches^i long. 

Developmental Stages. A young female, 210 mm long, resembles the adult closely 
in general form and has 43 enlarged tubercles in the midline to the first dorsal, 10— i i 
between the first and second dorsals, 2—4 in front of each orbit, 3—5 around the inner 
orbital margin to the inner end of each spiracle, and 2—3 on each shoulder. The tubercles 
in the midline are more uniform in size and are rather more thorn-like in shape than 
in the adult, though not appreciably larger. The tip of the snout does not bear a fleshy 
fringe, such as that which characterizes the young of R. productus of the Pacific Coast 
of Mexico and California. 

Habits. In Uruguayan waters it is taken on sandy bottom throughout the year 
down to a depth of iio meters.*^ Nothing more is known of its habits. 

Range. Coastal waters of the western Atlantic from northern Argentina to the 
Caribbean; also reported off" tropical West Africa (Dahomey and mouth of the Congo).** 

Occurrence in the Western Atlantic. This Guitarfish is generally distributed from 
about Lat. 38° S to the Caribbean; it has been reported from: Puerto Quequen, Mar 
del Plata, and Buenos Aires, Argentina; from the vicinity of Montevideo and from 
Maldonadoj Uruguay; from Santos, Rio de Janeiro, Pernambuco, Bahia, Ilha S. 

60. Miiller and Henle, Plagiost., 1841: 416, as Rkinobatus undulatus. 

61. Kner, Novara Exped., Zool., i, Fische, 1865: 416, as Rkinobalus undulatus. 

62. Devincenzi, An. Mus. nac. Montevideo, (2) i (4), 1920: 126. 

63. For West African references, see Fowler (Bull. Amer. Mus. nat. Hist., yo [i], 1936: loi). 



72 Memoir Sears Foundation for Marine Research 

Sebastiao at the mouth of the Amazon, and Natal, Brazil; Trinidad; Atlantic Panama; 
Jamaica; and the Lesser Antilles (probably St. Eustatius). 

An interesting feature of its distribution is the fact that it ranges poleward nearly 
or quite to the 40th parallel of latitude in the southern hemisphere, where the max- 
imum temperature for the year is only about 20—21° C (68—70° F) and where the 
yearly minimum may fall as low as 9—10° C (48—50° F). However, in the northern 
hemisphere it appears to be confined to strictly tropical latitudes and to temperatures 
at least no lower than about 25—26° C (77—79° F). It is replaced farther north by 
R. Jentiginosus (p. 67). 

Synonyms and References: 

Raja percellens Walbaum, P. Artedi Genera Pise. Emend. Ichthyol., J, 1792; 525 (descr., by ref. to Marc- 
gravius [Hist.-Nat. Brazil, 1648]; Brazil). 

Rhinoiatus^* fercellens Jordan and Evermann, Bull. U. S. nat. Mus., 47 (i), 1896: 63 (descr., Brazil to Ja- 
maica); Rep. U. S. Comm. Fish. (1895), 1896: 220 (listed, W.Indies to S.Brazil); Schreiner and 
Ribeiro, Arch. Mus. nac. Rio de J., 12, 1903 : 80 (listed, Rio de Janeiro and Rio Grande do Sul, Brazil); 
Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907: 174, 207 (descr., refs.); Engelhardt, Abh. bayer. Akad. 
Wiss., 4 (3), Suppl., 1913: loi (range); Garman, Mem. Harv. Mus. comp. Zool., 56, 1913: 278, 
pi. 55, fig. 4, pi. 56, fig. 7, pi. 65, fig. 2 (descr., ills., vertebrae, heart, skel., Rio de Janeiro); Starks, Stan- 
ford Univ. Publ., Univ. Ser., 1913: 5 (descr., color, off Natal, Brazil); Fowler, Proc. Acad. nat. Sci. 
Philad., 68, 1916: 401 (listed, color); Ribeiro, Rev. Mus. pauL, 10, 1918: 708 (listed, Ilha S. Sebastiao 
and Santos, Brazil); Devincenzi, An. Mus. nac. Montevideo, (2) j (4), 1920: 126 (descr., Rio de la 
Plata, season); Lahille, Physis B. Aires, 5, 1921: 63 (listed, Argentina); Meek and Hildebrand, Field 
Mus. Publ. Zool., 15 (i), 1923: 69 (descr., color, refs., Atlant. Panama); Ribeiro, Fauna Brasil., Peixes, 
2 (i) Fasc. I, 1923: 31, pi. II (descr., photos, range); Marelli, Elenc. Sist. Fauna B. Aires, Mem. Ministr. 
Obras Publ. B. Aires (1922— 1923), Fishes, 1924: 547 (listed, Maldonado, Uruguay, Rio de la Plata 
and Mar del Plata, Argentina); Norman, Proc. Zool. Soc. Lond., 1926: 947 (in Key), 974 (descr., ill. 
head, Brazil); Devincenzi and Barattini, An. Mus. nac. Montevideo, (2) 2, Suppl. Alb. Ictiol., 1926: 
pi. 3, fig. 2 (ills.); Luderwalt, Rev. Mus. paul., 16, 1929: 40 (listed, Ilha S. Sebastiao, Brazil); Marini, 
Physis B. Aires, 9, 1929: 452 (listed, Puerto Quequen, Argentina); Jordan, Evermann and Clark, Rep. 
U. S. Comm. Fish. (1928), 2, 1930: 23 (listed, Jamaica to S. Brazil); Pozzi and Bordale, An. Soc. cient. 
argent., 120, 1935: 152 (Argentina, Lat. 35-40° S., depth); Fowler, Bull. Amer. Mus. nat. Hist., jo 
(i), 1936: loi (descr., color, nos. teeth, Trinidad and mouth of Congo); White, Bull. Amer. Mus. 
nat. Hist., 74, 1937: 91 (no. heart valves, after Garman, 1913); Fowler, Monogr. Acad. nat. Sci. Philad., 
6, 1944: 456 (listed, Jamaica; Panama); Paes de Oliveira, Ovogen. Peixes Brasil, i, in Bol. Minist. Agric. 
Brazil (1943), 1945: 5 (ovary), 10 (descr.), pi. not numbered (photo, Brazil); Lopez, An. Mus. Argent. 
Cienc. Nat., 42, 1947: 113 (kidney, Argentina). 

Rhinobatus electricus Bloch and Schneider, Syst. Ichthyol., 1801: 356 (descr. by ref. to Marcgravius [Hist.- 
Nat. Brazil, 1648: 151] Brazil). 

Rhinobatus glaucostictus Olfers, Die Gattung Torpedo, 1831: 22 {glaucoslictus substituted for electricus Bloch 
and Schneider 1801 because not electric, color). 

Rhinobatus undulatus Olfers, Die Gattung Torpedo, 1831 : 22 (color, Rio de Janeiro); Castelnau, Anim. Nouv. 
Rares Amer. Sud, Poiss., 1855: 100 (listed, Rio de Janeiro); Kner, Novara Exped., Zool., J, Fische, 
1865: 416 (descr., size, color, Rio de Janeiro); Giinther, Cat. Fish. Brit. Mus., 8, 1870: 444 (descr., 
Bahia, Brazil); Garman, Proc. U. S. nat. Mus., J, 1880: 518 (descr., color, Rio de Janeiro southward); 
Cockerell, Bull. Inst. Jamaica, i, 1892: 7 (listed, Jamaica); J. Inst. Jamaica, I (5), 1893: 178 (Jamaica; 
not seen); Berg, An. Mus. nac. B. Aires, (2) j, 189;: 10 (listed, Montevideo, Maldonado, Buenos Aires, 
Mar del Plata, nos.); Ihering, Rev. Mus. paul., 2, 1897: 35 (listed, off Rio Grande do Sul, Brazil); 
Metzelaar, Trop. Atlant. Visschen, 1919: 194 (listed, trop. Atlant.). 

Rhinobatus marcgravii Henle, Ueber Narcine, 1834: 34 {marcgravii substituted for electricus Bloch and Schnei- 
der x8or because not electric). 

64. Also spelled Rhinobatos. 



Fishes of the Western North Atlantic 73 

Rhinobatus {Rhinobatits) undulatus Miiller and Henle, Plagiost., 1841: 121, pi. 40 (descr., meas., color, ill., 
Bahia, Brazil); Dumeril, Hist. Nat. Poiss., j, 1865: 498 (descr., color, Brazil). 

Rhinobatus stellio Jordan and Evermann, Rep. U. S. Comm. Fish. (1895), 1896: 220 (listed, Jamaica, name 
only); Jordan and Rutter, Proc. Acad. nat. Sci. Philad., 1897: 91 (descr., color, Jamaica); Jordan and 
Evermann, Bull. U. S. nat. Mus., 47 (3), 1898: 2750 (descr. after Jordan and Rutter, 1897, Jamaica); 
Engelhardt, ."Vbh. bayer. Akad. Wiss., Suppl. ^ (3), 1913 : loi (range); Norman, Proc. Zool. Soc. Lond., 
1926: 947 (in Key), 972 (descr., ill. head, Jamaica); Jordan, Evermann and Clark, Rep. U. S. Comm. 
Fish. (1928), 2, 1930: 23 (listed, Jamaica). Fowler, .Acad. nat. Sci. Philad., 6, 1944: 456 (listed, Ja- 
maica). 

Rhinobatus pellucens Fowler, Proc. .Acad. nat. Sci. Philad., 6"], 1916: 521 (listed, Trinidad, /ii'//«c^Z!/ perhaps 
misspelling for perce/kns). 

Probable Synonym : 

Rhinobatus undulatus Osorio, J. Sci. math. phys. nat. Lisboa, (2) j, 1895: 253 (listed, Dahomey, \V. .Afr.). 

Genus Rhinohatos Addendum 

We include here one problematical species reported from Mexico without indi- 
cation as to whether it is from the Atlantic or the Pacific. 



Rhinohatos spinosus Giinther i 8 70 

Study Material. None. 

Knowledge of this Guitarfish is limited to the original account of a stuffed specimen 
13 inches long, now in the British Museum (Natural History). ^^ 

Compressed spines with dilated base along median line of back, on shoulder, 
and above eye and spiracle; entire upper surface rough. Snout much produced, the 
distance between outer angle of nostrils half of that between mouth and end of snout. 
Anterior nasal valve not dilated laterally. Mouth nearly straight. Rostral ridges con- 
fluent, very narrow, with a small and short groove at base and provided with spines 
in their entire length. Snout white. 

Miss Ethelwyn Trewavas contributes the additional information that the spines 
along the rostral ridges are thorn-like, pointing rearward, on ovoid bases, and in two 
irregular series, with smaller ones among and between them. 

According to this description, R. spinosus differs from all others of its genus in 
the Atlantic and eastern Pacific in the fact that it continues to be spiny along its rostral 
ridges until it reaches such a large size — unless R. rasus Garman 1908 ** is distinct from 
R. cemiculus and is similarly characterized. R. spinosus is known at present from the 
original specimen only. Further discussion of it is best postponed until other specimens, 
Atlantic or Pacific, come under observation. 

References : 

Rhinobatus spinosus Giinther, Cat. Fish. Brit. Mus., 8, 1870: 518 (descr., Mexico); Garman, Proc. U. S. nat. 
Mus., J, 1880: 518 (descr. after Gunther, 1870); Jordan and Evermann, Bull. U. S. nat. Mus., 47 

65. Gunther, Cat. Fish. Brit. Mus., 5, 1870: 578. 

66. So far known from one small specimen; see discussion, p. 54. 



74 Memoir Sears Foundation for Marine Research 

(i), l8g6: 63 (descr. after Giinther, 1870; Mexico, "probably the east coast"); Rep. U. S. Comm. 
Fish. (1895), 1896: 220 (listed, Mexico); Garman, Mem. Harv. Mus. comp. ZooL, j6, 1913: 282 
(descr. after Giinther, 1870). 

Genus Zapteryx Jordan and Gilbert 1880 

Zapleryx Jordan and Gilbert, Proc. U. S. nat. Mus., j, 1880: 53; type species, Platyrhina exasferata Jordan 
and Gilbert (Proc. U. S. nat. Mus., J, 1880: 32). California. 

Generic Synonyms: 

Syrriina (subgenus oi Rhinobatus) in part, Miiller and Henle, Plagiost., 1841: 113. 

Syrrhina Garman, Proc. U. S. nat. Mus., J, 1881: 521; type species, Platyrhina exasferala Jordan and Gil- 
bert 1880; see discussion, p. 48, footnote 17. 

Generic Characters. Disc wedge-shaped anteriorly and strongly flattened dorso- 
ventrally, outer pectoral margins rounded and tail stout, as in Rhinobatos^ but disc 






|?>.,5^|$1 



-•/ 










Figure 17. Zapleryx exasperata, female, 207 mm 
long, from San Diego, California (Harv. Mus. 
Comp. ZooL, No. 36022). Large tubercles and 
smaller denticles along mid-dorsal line of disc, 
about 10 X- 



relatively broader (cf. Fig. 18 with 14, 16), so that the general appearance is more 
skate- or ray-like. Nostrils entirely separate from mouth and approximately transverse. 
Flap-like expansions of anterior margins of nostrils almost wholly covering inner halves 
of nasal openings and extending inward considerably beyond inner corner of nostril 



Fishes of the Western Nortfi Atlantic 7 5 

(Fig. 18C) though widely separated one from the other in the median line; posterior 
margin of nostril also expanded as a blunt lobe. Dermal armature much as in Rhinobatos, 
except that minute dermal denticles are thickly interstrewn with larger conical ones 
on upper surface; a few on lower surface also. Generic characters otherwise as in 
Rhinobatos. 

Size. Maximum length about three feet. 

Habits. Similar to those of Rhinobatos, so far as is known. 

Range. Known only from Brazil and along the Pacific Coast of America from south- 
ern California and the Gulf of California southward to Panama. 

Species. Only two, the one Atlantic, the other Pacific. 

Key to Species 

I a. Dorsal fins with posterior margins longer than their bases; rostral ridges con- 
verging anteriorly; small denticles on dorsal surface posterior to nuchal region 
rising gently rearward or even nearly horizontal, their bases irregularly scalloped, 
not regularly stellate or striate (Fig. 19B). 

brevirostris Miiller and Henle 1841, p. 75. 

lb. Dorsal fins with posterior margins shorter than their bases; rostral ridges nearly 

parallel; small scales on dorsal surface conical, erect, their bases conspicuously 

striate or stellate (Fig. 17). exasperata Jordan and Gilbert 1880. 

Pacific Coast of America, southern California 
to Panama. 



Zapteryx brevirostris (Miiller and Henle) 1841 

Short-nosed Guitarfish 

Figures 18, 19 

Study Material. Four females, 445 to 480 mm long, and a juvenile male, 345 mm, 
from Brazil, in Harvard Museum of Comparative Zoology. 

Distinctive Characters. The heart-shaped disc of Zapteryx, combined with its stout 
muscular tail bearing two large dorsal fins, marks it off from all other batoids known 
from the western Atlantic except for the three local Guitarfishes, Rhinobatos horkelii, R. 
lentiginosus and R. percellens. It is separable from these by its much more obtuse snout 
and relatively broader disc, together with the fact that the flap-like expansions of the 
anterior margins of its nostrils almost wholly roof over the inner half of the nasal 
apertures. 

Description. Proportional dimensions in per cent of total length. Male, 345 mm, 
and female, 460 mm (Harv. Mus. Comp. Zool., No. 536) from Rio de Janeiro, Brazil. 
Disc: extreme breadth 47.3, 48.7; length 42.2, 41.7. 
Snout length: in front of orbits 9.0, 9.4; in front of mouth 1 1.9, 12.0. 
Orbits: horizontal diameter 4.4, 3.7; distance between 4.8, 4.1. 



76 



Memoir Sears Foundation for Marine Research 




Figure i8. Zapleryx brevirostris, immature male, about 345 mm long, from Rio de Janeiro, Brazil (Harv. Mus. 
Comp. Zool., No. 536). A Ventral view of pelvics. B Lateral view of tail. C Mouth and nostrils, about i X. 
D Eye and spiracle, about 1.5 X- E Upper teeth, about 18 X. 

Spiracles: length 2.8, 2.6; distance between 7.0, 6.5. 
Mouth: breadth 7.2, 7.2. 

Nostrils: distance between inner ends 4.1, 3.7. 

Gill openings: lengths, ist 1.7, 1.9; 3rd 1.6, 1.7; 5th 1.3, 1.3; distance between 
inner ends, ist 15.7, 15.2; 5th 12.2, 12.2. 



Fishes of the lVester?i Northi Atlantic 



11 



First dorsal fin: vertical height 7.6, 7.4; length of base 5.1, 5.0. 

Second dorsal fin: vertical height 7.3, 7.2; length of base 5.7, 5.6. 

Caudal fin: upper anterior margin 13.7, 13.1. 

Pelvics: origin to tip 17.4, 18.5. 

Distance: from tip of snout to center of cloaca 41.8, 44.3 ; from center of cloaca 

to tip of tail 58.2, 55.7. 
Interspace between: ist and 2nd dorsals 7.8, 8.0; 2nd dorsal and caudal 6.7, 5.4. 




Figure 19. Zapleryx brevirostris. A Large tubercle and smaller dermal denticles on right-hand shoulder of spe- 
cimen illustrated in Fig. 18, 17 x. B Dermal denticles from side of tail of female, 450 mm long, from Rio de 
Janeiro (Harv. Mus. Comp. ZooL, No. 429), about 30 X. C Dermal denticles from lower surface of abdomen 
of same, about 30 X. 



Disc a little broader (i.i) than long, heart-shaped, its anterior angle to level of 
fronts of orbits about 105—107°; anterior margins weakly convex opposite eyes, slightly 
concave opposite spiracles in young specimens and in adult females, but somewhat 
more deeply so in mature males;*' posterior corners broadly rounded, posterior margins 
moderately so; posterior corners of pectorals overlapping pelvics by a distance about 
as great as that between exposed nostrils. Tail from center of cloaca longer (1.2— 1.4) 
than distance from center of cloaca to tip of snout, about as broad opposite axils of 
pelvics as distance between outer ends of spiracles; tapering evenly rearward; flattened 
below and moderately rounded above, each side with a longitudinal dermal fold ex- 
tending from just posterior to tips of pelvics to a little beyond origin of caudal. 

Dermal denticles on dorsal surface close-set but with skin exposed between; those 

67. It is so shown in Ribeiro's (Fauna brasil., Peixes, 2 [i] Fasc. i, 1923 : pi. 12) photograph of a male with large claspers. 



78 Memoir Sears Foundation for Marine Research 

on head, on midzone of back, and on tail, low, conical to pyramidal, with irregularly 
fluted bases; those over pectorals as a whole more scale-like and raised but little from 
the skin, with sharp points and usually with two longitudinal ridges ; interspersed among 
these (most numerous over inner parts of pectorals and along sides of anterior part 
of tail) are considerably larger denticles, ranging in form from blunt-conical to blunt- 
pyramidal, their bases fluted anteriorly. Midline of back, from nuchal region to origin 
of first dorsal, with an irregular row of 21—23 domed prominences, fluted basally and 
covered with small rounded denticles, a larger rounded tubercle (similarly fluted) 
emerging from the tip; between first and second dorsals, two or three similar but 
smaller prominences, with tubercle at tip; two series of tubercles on each shoulder, 
the inner series of two, the outer series of two or three; eight to ten along posterior part 
of each orbital ridge (decreasing in size anteriorly), with a larger one on anterior margin 
of orbit; a row of four to eight pyramidal tubercles along outer margin of each pectoral 
a little anterior to axis of greatest breadth in some specimens.** Lower surface closely 
clothed with small flatfish denticles, irregularly quadrate or hexagonal, with rounded 
corners. 

Snout in front of orbits about 1.9—2.2 times as long as distance between orbits, 
its length in front of mouth about 1.6— 1.7 times as great as width of mouth and about 
3.0—3.2 times as great as distance between inner ends of nostrils. Crown concave 
transversely, the margins of orbits prominent. Horizontal diameter of eyes between 
^4 and ^/g (about 2 i "/o) as long as snout in front of orbits. Length of eye plus spiracle 
about half as great as distance between spiracles. Spiracles about ^\^ as long as orbits, 
moderately oblique, their Inner ends directed rearward, with one low ridge on posterior 
margin. First pair of gill openings (the longest) a little more than half as long as nostrils, 
the second to fifth pairs a little shorter in succession; distance between Inner ends of 
fifth pair about as long as snout in front of mouth. Nostrils transverse or slightly 
oblique; length about as great as distance between nostrils (95— 100 "/o) and a little 
less than half as great as width of mouth; anterior margin expanded as a broad rounded 
flap almost wholly covering Inner half of nasal aperture and continuing inward as a 
narrow fold with rounded lobular termination on Internarlal space, about halfway 
toward midline; posterior margin of nostril with two flap-like expansions directed 
rearward toward mouth, the outer narrower, the Inner broadly rounded; also a nar- 
rower and longer intervening lobe directed across nasal aperture. Mouth only slightly 
bowed, its corners wrinkled but without extensive pits or furrows. 

Teeth ^|Eifo) ^°w> those of females and of young males ranging from weakly 
rounded to nearly flat, without cusp, closely crowded in quincunx arrangement; about 
10 rows In function simultaneously In center of mouth In upper jaw and about 12 rows 
in lower; those of sexually mature males not seen. 

First and second dorsals about equal in size, triangular, with narrowly rounded 
apex and weakly convex anterior and posterior margins, the latter nearly vertical; free 
rear basal margins about half as long as base. Origin of first dorsal posterior to level 

68. A male and two females have these, but two other females lack them. 



Fishes of the lVester?i North Atlantic 79 

of axils of pelvics by a distance about half as great as distance from origin of first dorsal 
to origin of caudal and about as long as interspace between the two dorsals. Interspace 
between first and second dorsals about 1.5— 1.6 times as long as base of first dorsal. 
Interspace between base of second dorsal and origin of caudal about as long as base 
of second dorsal. Upper and lower origins of caudal about opposite one another; caudal 
axis horizontal or bent slightly upward; upper margin nearly straight and nearly as 
long as distance from origin of first dorsal to origin of second; lower posterior margin 
evenly rounded, without separate lower lobe; height of caudal above termination of 
axis about 1.3 times as great as depth below the latter. Pelvics broadly rounded out- 
wardly, decreasing to narrowly blunted tips, their inner margins weakly concave, their 
extreme length from origin to tip about twice as great as length of interspace between 
first and second dorsals; their axils separated by a distance about as great as breadth 
of mouth in females, but only about ^\^ that great in juvenile male. Claspers of mature 
male slender, as in Rhinobatos^ their tips somewhat swollen, extending back about to 
level of rear end of base of first dorsal. 

Rostral cartilage broad basally, tapering anteriorly to narrow tip close to ex- 
tremity of snout; its lateral ridges low, inconspicuous, far apart at base but converging 
toward tip. Anterior rays of pectorals extending for only a short distance anterior to 
level of front of orbital region of cranium, thus falling far short of extremity of snout. 

Color. Preserved specimens olive to brownish gray above, without definite markings; 
dorsals and caudal somewhat darker than general ground tint; margins of pectorals 
and pelvics paler; space on either side of rostral cartilage also pale, possibly translucent 
in life. Lower surface grayish or yellowish white, the posterior corners of pectorals and 
tips of pelvics more or less dusky. 

Size. The largest specimen recorded was about 2 i inches long (540 mm),*^ but 
it is not unlikely that Z. brevirostris may reach as great a size as Z. exasperata of the 
Pacific, a length of about three feet.'" 

Developfnental Stages. It is probable that development is ovovivi parous, but 
neither the embryos nor the newborn young have been reported. 

Habits. Nothing whatever is known of the habits of this Ray to distinguish it from 
the Guitarfishes of the genus Rhinobatos. 

Range. Known so far from Brazil alone, where it has been taken near Rio de 
Janeiro and near Bahia. Apparently it is not common though occasionally trawled in 
some numbers. 

Synonyms and References: 

Rhinobatus {Syrrhind) brevirostris MuUer and Henle, Plagiost., 1841: 114, 192, pi. 36 (descr., meas., ill., 

Brazil); Castelnau, Anim. Nouv. Rares Amer. Sud, Poiss., 1855: 100 (listed, Rio de Janeiro); Dumeril, 

Hist. Nat. Poiss., I, 1865: 489 (descr., size, Brazil). 
Platyrhina sinensis (in part) Gray, List Fish. Brit. Mus., j, 1851: 98 (specimen from Brazil, but not refs.). 
Rhinobatus brevirostris Giinther, Cat. Fish. Brit. Mus., 8, 1870: 447 (descr., size, Brazilian specimen pre- 

69. Dumeril, Hist. Nat. Poiss., i, 1865: 490. 

7c. Ribeiro (Fauna brasil., Peixes, 2 [i] Fasc. i, 1923: pi. 12) gives an excellent photograph of a male with fully devel- 
oped claspers but without stating the size. 



8o Memoir Sears Foundation for Marine Research 

viously listed by Gray, 1851, as P/atyrAina sinensis); Schreiner and Ribeiro, Arch. Mus. nac. Rio de J., 

12, 1903: 80 (listed, Rio de Janeiro); Ribeiro, Pescas 'Annie,' 1904: 18 (nos., Ilha Rasa, Brazil); Arch. 

Mus. nac. Rio de J., 14, 1907: 175, 207 (diagn., refs., Rio de Janeiro and Bahia, Brazil); Fauna Brasil., 

Peixes, 2 (l) Fasc. i, 1923: 31, pis. 12, 13 (diagn., photos, male and female, Rio de Janeiro and Bahia, 

Brazil). 
Syrrhina bnvirostris Garman, Bull. Mus. comp. Zool. Harv., ly, 1888: 89, pi. 25 (mucous canals); Mem. 

Harv. Mus. comp. Zool., j6, 1913: 285, pi. 65, fig. 3 (descr., ill. skel., Rio de Janeiro). 
Zapteryx brevirostris Norman, Proc. zool. Soc. Lond., 1926: 943 (in Key), 980, fig. lE (ill. nostril, size, refs. 

Brazilian specimen in Brit. Mus.). 



Suborder TORPEDINOIDEA 
Electric Rays, Torpedo Rays 

Characters. Trunk anterior to cloaca a depressed subcircular disc, fleshier toward 
its margins and thicker there than in most other disc-shaped batoids, the body softer. 
Tail rather sharply marked off from body sector, with or without lateral folds; broader 
basally than in any other batoids except the Sawfishes (Pristoidea) and Guitarfishes 
(Rhinobatoidea) ; as long as body in some, but shorter than body in most, and reduced 
in some to a mere rudiment bearing the fins.^ Attachment of anterior parts of pectorals 
to sides of head extending forward to or beyond level of eyes. One or two well developed 
dorsal fins, or none; the first (if there are two) partly over or close behind bases of 
pelvics. Caudal fin well developed, its axis raised but slightly if at all. Pelvics with 
outer margins continuously convex in most cases but somewhat emarginate in some, 
so deeply so in one genus that the anterior subdivision forms a separate limb-like 
structure arising from the lower surface of the disc some distance inward from the 
margin^ (as in some genera of Rajoidea also, pp. 314, 327). Inner posterior margins of 
pelvics free from sides of tail in some species but united with it in others. 

Eyes small, functional in most species, but rudimentary or even entirely obsolete 
in a few deep-water forms. Spiracles either close to eyes or separated from the latter 
by a narrow interspace, their margins either smooth or with larger or smaller knobs 
or papillae. Nostrils close to mouth but entirely separate from it, their anterior margins 
expanded and joined together as a single curtain-like flap with median attachment 
roofing over inner parts of nasal openings and extending rearward nearly or quite to 
the mouth. Mouth small to moderate in size; widely distensible in some genera, with 
the upper and lower jaw cartilages but loosely articulated together; more or less 
protractile in others, with the jaws not only firmly articulated but also bound together 
on either side by a labial cartilage of two elements with special muscles, thus limiting 
the gape (see also p. 108). Gill openings small. Teeth small, rounded, or with one to 
three more or less prominent cusps; in 12 to about 64 series in different species, 
arranged in bands that terminate some little distance short of either corner of mouth; 

1. Genus Hypnos, p. 87. 

2. This is the case in Typhlonarke, a genus of Narkidae (i. e., with one dorsal fin), founded by Waite (Rec. Canter- 
bury [N. Z.] Mus., I [2], 1909: 146) for an Electric Ray first described as Astrape aysoni Hamilton (Trans. N. Z. 
Inst., J4, 1902: 25, pis. 10-12). 



Fishes of the Western North Atlantic 8 1 

several rows in function simultaneously; the integument bearing the tooth bands firmly 
attached to jaw cartilages in some, only loosely so in others. 

Skin soft and entirely naked in most species, but with margins of pectorals de- 
scribed as having small papillae "partly spinous" in one.^ 

Rostral projection of cranium single or double and more or less branching in 
some; reaching to anterior margin of disc. Antorbital cartilages extending forward to 
support front of disc, their anterior margins dissected in complex fashion in some 
species but less so in others, and spreading laterally to anterior rays of pectorals or 
even overlapping latter.* Extremities of branchial rays widely expanded as plate-like 
discs. Dorsal and caudal fins supported basally by short cartilaginous radials, distally 
by much more numerous fine horny rays (ceratotrichia) in double series. Pectoral and 
pelvic fins without horny rays, the cartilaginous radials extending outward to margins. 
Pelvis (in species studied) bowed rearward, with a well developed lateral process ex- 
tending forward at either end. Margins of gill arches smooth inward from gill filaments. 
Electric organs well developed, between sides of head and forward extensions of pec- 
torals. 

Electric Organs. The anatomy and physiology of the electric organs of the Tor- 
pedo Rays, as well as of those of other fishes, and the characteristics of the electric cur- 
rents generated by them, have been the subject of much research, resulting in extensive 
literature.^ Each of the two organs of an Electric Ray occupies one side of the anterior 
part of the disc between the anterior extension of the pectoral and the head, extending 
forward about to the level of the eye and rearward past the gill region to the vicinity 
of the pectoral girdle. Together the two organs comprise about one-sixth of the total 
weight of the fish in each of the two western Atlantic species {Torpedo nohiliana and 
Narcine brasilioisis) for which this relationship has been determined. In most cases the 
outlines of the organs are visible externally on the ventral side and usually faintly so 
on the dorsal side as well, where, however, they are more hidden from view by the 
pigmentation of the skin. 

Numerous columnar structures, separated by loose fibrous tissue, make up each 
organ and occupy the entire thickness of the disc from the upper to the lower integu- 
ment, their lengths decreasing toward the periphery as the disc becomes thinner. These 
columns, often referred to as "prisms," have been described repeatedly and have been 
pictured more or less diagramatically as arranged like the cells of a honeycomb.* 

3. Torpedo mackayana Metzelaar (Trop. Atlant. Visschen, 1919: 197). 

4. For a recent discussion of the head skeleton and comparison with other batoids, see Holmgren (Acta Zool. Stockh., 
22, 1941 : 57, 65, 66). 

5. For general summaries of results and references, see Gotch (in Shafer, Text Bk. Physiol., 1900: 561-591) and espe- 
cially Garten (in Wintersheim, Handb. Vergl. Physiol., 3 [2], 1910-1914: i72fF). For more detailed accounts, see 
Gotch (Philos. Trans., [B] 17S, 1888: 487-537; [B] 179, 1889: 329-363), Schoenlein (Z. Biol., 31, N. S. 13, 1895: 
449-523), Fuji (J. CoU. Sci. Tokyo, J7, 1914: i), as well as Coates and Cox (Zoologica N. Y., 27, 1942: 25), and 
Cox and Breder (Zoologica N. Y., 28, 1943: 45) for observations on the discharges of Torpedo nohiliana and of 
Narcine brasiliensis respectively. 

6. For semidiagrammatic illustrations of the arrangement of the columns in various species, see especially Fritsch 
(Elektr. Fische, 2, 1890: pis. 3-15). For a more realistic illustration of their actual appearance in Torpedo when 
exposed to view, see Jobert (Appar. Electr. Poissons, 1858: pi. i, figs. 1-3). 



8 2 Memoir Sears Foundation for Marine Research 

Actually, however, it is a matter of common knowledge that they are more loosely 
aggregated than the foregoing would suggest (Fig. 25F) and that they vary in cross 
section from hexagonal with rounded corners to roughly circular. The number of 
columns in a single organ ranges from as few as 140—150 in some genera (Temera and 
Discopyge) up to an average of 1,025—1,083 for Torpedo nobiliana.'' 

Each column is divided by transverse partitions into a large number of so-called 
electric discs, each consisting of a clear jelly-like mass that includes a number of large 
nuclei and each being separated from its neighbors by a connective tissue layer through 
which run the nerve fibers and blood vessels. Since there may be 375 or more discs 
per column, their total number in the larger species* is in the hundreds of thousands 
for each organ. It has been stated that both the final number of columns in the organ 
and the numbers of electric plates per column are established before birth. But a slight 
increase In the number of columns takes place with growth, 267, 276, and 315 having 
been counted for each organ on three embryos of Narcine brasiliensis from Florida, 
250, 382, and 409 for three adults. Also, It has been found that the number of plates 
per column varies according to the length of the column ; in one embryo of Narcine 
brasiliensis the average counts were 305 near the Inner and thicker edge of the organ 
but only 179 per column near the outer and thinner edge; the counts were 482 and 288 
respectively In the organ of another.^ 

The nerve supply to the electric organs Is highly developed, each organ receiving 
one branch of the trigeminal nerve and four branches of the vagus nerve; the former 
and the three anterior branches of the latter are as thick as the spinal cord and all arise 
together from a special lobe of the brain known as the "lobus electricus." Distally, 
these nerves branch again and again and terminate in a cluster of fine fibrils on the 
ventral connective tissue wall of each electric plate. It has been found that the ventral 
sides of the plates are negative (electrically speaking) to the dorsal sides, and that the 
lower side of each electric organ as a whole is negative, thus representing the plus or 
anode pole of what appears to be a form of multiple concentration cell. In other words, 
the discharge produced by the organ as a whole passes through the latter from the 
ventral side toward the dorsal, whereas in an Electric Eel {Electrophorus) it runs through 
the organ from the tail toward the head, in an Electric Catfish {Malopterurus) from 
front to rear. All parts of a single organ, and the two organs as well, discharge 
almost simultaneously, and it has been observed repeatedly that the discharge Is ac- 
companied by slight muscular contractions of the disc, cupping the margins of the 
latter upward. 

The ability of Electric Rays to produce shocks was "well known to the ancient 
Greeks and Romans,"!" ^iX^di it has been known for many years that the discharge has 

7. For a table of average numbers of columns in different species, see Fritsch (Elektr. Fische, 2, 1890: 97; nobiliana 
listed as "hebetans"); also Ballowitz and Schmidt (in Bolk and others, Handb. Vergl. Anat. Wirbelt., 5, 1938: 669). 

8. The number per organ for a specimen of the European Torpedo marmorata was 179,625, as calculated by Fritsch 
(Elektr. Fische, 2, 1890: 102). It may reach 500,000 in large specimens of other species of the genus. 

9. Cox and Breder, Zoologica N. Y., 28, 1943: 46. 

10. For further details, see Norman and Eraser (Giant Fishes, 1937: 66). 



Fishes of the Wester?! North Atlantic 8 3 

all the properties of electricity, being capable of producing a spark, of deflecting or 
magnetizing a needle, of electrolyzing chemical compounds (e. g., of dissociating iodide 
of potassium), and so forth; also, it is audible on a telephone suitably connected. 

It is commonly said that the production of the shock is under the voluntary 
control of the Torpedo, both as to time of discharge and as to its strength. Electric 
Rays, lying undisturbed on the bottom of an aquarium, have been observed (by tele- 
phone) to give oft' a succession of discharges spontaneously; i. e., without any apparent 
stimulation." However, the regularity with which a discharge follows when the skin is 
touched or otherwise stimulated suggests that it is chiefly a simple reflex action induced 
by tactile stimulation. It has long been known that the fish lies quiet for some little 
time without further discharge after delivering a shock, and the delivery of several 
discharges at rather brief intervals leaves it in an exhausted condition, after which it 
requires a considerable rest period before it can discharge again. We may cite the case 
of an Australian specimen of Hypnarce that delivered 50 successive shocks within about 
ten minutes, intense at first but weakening until hardly discernible. ^^ And it has been 
observed in laboratory experiments that the voltage and power decline rapidly as the 
fish becomes increasingly fatigued. 

The individual discharges or pulses, each lasting perhaps 0.03 second, follow 
one another in rapid succession" in trains. In Torpedo nobiUana of the western Atlantic, 
the number of pulses per train appears to average about 1 2 but may go as high as 
100, exceptionally more than that. Either one train, or a few trains following one 
another in rapid succession, constitute what is commonly termed a "shock." The 
electromotive force of the discharge varies not only from species to species and from 
specimen to specimen of a given species but with the condition of the fish, while dif- 
ferences in recorded results are partly to be explained as due to differences in the sen- 
sitivity of the apparatus employed. Measured voltages range from as low as 8—17^* 
to as high as 70— 80^* for the European Torpedo marmorata, with estimates going as 
high as 200.1^ Seven to 37 volts, with various resistances, have been recorded for 
Narcine brasiliensis\ 25 volts was recorded for one specimen of Torpedo nohiliana and 
220 volts for another of about the same size, a divergence which illustrates the dif- 
ferences in the electric capabilities of different specimens of the same species." This 
last voltage (220), which probably approximates the maximum to be expected from any 
Electric Ray, is about equal to that of the Electric Catfish {Malopterurus) of North 
African rivers but falls considerably below that of the so-called Electric Eel (Electro- 
phorus) of South America. 

The literature on the Electric Rays includes almost endless references, from 
classic times down to the present, pertaining to the effects of the shocks received by 

II. Schonlein, Z.Biol., 31 (N. S. 13), 1S95: 451. 12. McCuUoch, Aust. Mus. Mag., i (3), 1921: 89. 

13. Rates reported as being up to 150 per second. 14. d'Arsonval, C. R. Acad. Sci. Paris, I2i, 1S95: 145. 

15. Cremer, Verh.Gesell. dtsch. Naturf., Art. 80, samml. Cologne, 1909 (2), 2 half. 2 grup.: 523. 

16. Gotch and Burch, Proc. roy. Soc. Lend., 65, 1900: 442. 

17. See Coates and Cox (Zoologica N. Y., 27, 1942: 28) and Cox and Breder (Zoologica N. Y., 28, 1943 : 49) for tabul- 
ations of voltage, current, and power for Torpedo nobiliana and Narcine brasiliensis. 



84 Memoir Sears Foundation for Marine Research 

persons who have come in contact with them in one way or another. The following is an 
interesting first-hand account of such an experience in Australia. 

Whilst wading at Gunnamatta Bay I trod on an electric ray and it was as if a large hand clutched 
my foot and ankle. I netted the specimen and put it on a sand bank. Here I noticed a black leech between 
its eyes, and in trying to knock this off with my net got a more severe shock, which so suddenly contracted 
the muscles of my arms and legs that I leapt a foot in the air.^* 

Indeed, the shock from a large one in rested condition is strong enough to knock 
down and temporarily disable a full-grown man (p. 102). So Electric Rays might even 
be dangerous to bathers who accidentally step on them as they lie buried in the sand 
in the shallow water of regions where the larger sizes are common enough for this to 
be a likely occurrence. 

It has been stated that Torpedoes are immune to their own shocks. But it seems 
that they are not wholly so, for it has been observed that the discharge is accompanied 
by a slight and brief muscular contraction, as noted before (p. 82), this apparently 
resulting from the reception of its own discharge. 1^ It has been argued that their ap- 
parent immunity results, in reality, from the weakening of the electric discharge as 
it passes through the water. ^^ 

The requirement that the recipient of a shock must complete the circuit by making 
contact with the fish at two points, whether directly or indirectly, is fulfilled in the 
normal life of the Ray by means of the surrounding sea water. This has been illustrated 
experimentally with the recording of the shock by telephone, one pole of which is in 
metallic contact with the Torpedo, the other pole (also metallic) in the water at a dis- 
tance not greater than 20 cm (8 in.) from the fish." It is only if an Electric Ray is out 
of water, as when lying on the dry planks of a dock, that the mechanical two-point 
contact is requisite. And even in such cases it has been found that "quite a powerful 
sensation of numbness can be produced through the medium of a stream of water, that 
is to say, by pouring water onto a living fish."^^ 

The disabling effect that shocks by Torpedoes may be expected to have on small 
animals that may come in contact with them has given rise to a general belief that 
electric organs of the Torpedo normally serve a defensive purpose. Although it has 
been questioned recently whether the organs are of much importance in this respect, ^^ 
it has been observed in the Naples Aquarium that crayfish, crabs, and cephalopods are 
greeted by discharges if they chance to touch a Torpedo marmorata lying on the bottom. 
Indeed, one was seen to drive off a large octopus in this way.^* However this may be, 
it seems probable that the shocks do serve more or less to stun the prey, for wholly 
uninjured fishes, so large that it seems hardly conceivable that they could have been 
swallowed unless they had been rendered helpless beforehand, have been found in the 

iS. Whitley, Fish. Aust., j, 1940: 165. 

19. Jobert and Jolyet, Trav. Lab. Soc. sci. Arcachon (1895), 1896: 57. 

20. Schonlein, Z.Biol., 31 (N. S. ij), 1895: 512. 21. Schiinlein, Z.Biol., 31 (N. S. Jj), 1895: 450. 

22. Norman and Fraser, Giant Fishes, 1937: 65. 

23. Liibbert and Ehrenbaum, Handb. Seefisch. Nordeurop., 2, 1936: 317. 

24. Schonlein, Z.Biol., 31 (N. S. 13), 1895: 450, 453. 



Fishes of the Western North Atlantic 85 

stomachs of Torpedoes. On the other hand, it has been noted that small fishes (Gobies) 
and invertebrates placed on the head and fins of a large Australian species, Hypnos 
monopterygium (Shaw and Nodder) 1795, which is capable of delivering a shock of 
considerable intensity, "seemed to suffer no ill effects, though they must have received 
shocks." 2* Even if some species of Torpedo Rays do numb their prey to some extent 
by electric discharges, it is not likely that they are limited in their feeding to victims 
that have been partially incapacitated in this way, for at least some fishes are so resistant 
to currents passing through sea water that the Ray would actually have to come in 
contact with them for its discharges to have any disabling effect. Invertebrates, it seems, 
are more susceptible to the effect of the discharge. 

Size. The largest members of the genus Torpedo grow to a length of five or six 
feet and, according to reports, may reach a weight of 200 pounds (p. loi); the blind 
genus Typhlonarke (p. 80) probably reaches a length of at least four feet. At the op- 
posite extreme, the greatest length attained by some members of the genus Narke may 
be less than one foot. 

Development. They are ovoviviparous ; the European Torpedo marmorata is one of 
the Rays on which the development of the pectorals (leading to their attachment to the 
sides of the head) has been traced (p. 10 1). 2* 

Habits. The Electric Rays prey more or less indiscriminately on various crustaceans, 
mollusks, worms, and other invertebrates, as well as fishes (p. 102), some of the latter 
so large that it is a puzzle how the Rays manage to swallow them. A specimen of the 
Mediterranean Torpedo marmorata^ lying o" the sand in an aquarium, has been seen to 
raise the front of its disc while keeping the margins pressed down, thus forming a 
cavity into which small fish were swept by the inrush of water. In one instance a small 
shark emerged again undamaged, but in another a mullet was not seen again, probably 
having been captured." 

Sluggish in habit and feeble swimmers, they lie on the bottom most of the time, 
partially buried in sand or mud. The group as a whole covers a wide bathymetric range, 
for while most of its members occur in rather shallow water and some even survive 
stranding for some hours in the intertidal zone,^* others (genus Benthohatis) are confined 
to at least moderately great depths (p. 127). 

Range. The geographic range of one Electric Ray or another extends to all the 
oceans, including the Mediterranean, and covers the temperate, subtropical, and 
tropical belts. In the Atlantic they occur northward regularly to southern New Eng- 
land in the west and to northern Scotland (Moray Firth and Wick Bay) in the east, 
southward to southern Argentina in the west,-^ and to the southern extremity of the 

25. Whidey, Fish. Aust., i, 1940: 165. 

26. For illustrations of successive stages in the embryonic development of the European Torpedo marmorata, see especially 
Fritsch (Elektr. Fische, 2, 1890: pi. 15, figs. 38-43) and Garman (Mem. Harv. Mus. comp. Zool., j6, 1913: pi. 61, 
figs. 1-3). 27. Schonlein, Z.Biol., 3/ (N. S. jj), 1895: 453. 

28. The Australian genus Hypnos has this ability (Whitley, Fish. Aust., j, 1941: 166), although its vertical range also 
extends down to at least 120 fathoms. 

29. The most southerly record of any Electric Ray on the Argentine Coast with which we are acquainted is from Lat. 
46° 18' S (Norman, 'Discovery' Rep., 16, 1937: 11, 12). 



86 Memoir Sears Foundation for Mari?ie Research 

African continent (Table Bay) in the east. In the eastern side of the Pacific they are 
known from the State of Washington in the north to middle Chile in the south and 
in the western side from Japan southward to New Zealand, southern Australia and 
Tasmania. They are also distributed generally through the East Indies and all around 
the coasts of the Indian Ocean, including the Arabian Sea, the Persian Gulf, and the 
Red Sea, southward to southern Africa. In view of this wide distribution and of the 
evident tolerance of the group as a whole to a wide range of temperature and depth, 
the apparent failure of any of the Electric Rays to have colonized any of the oceanic 
island groups of the tropical Pacific^" presents something of a puzzle. 

Relation to Man. None of the Electric Rays is of any commercial value at present, 
nor are they ever likely to be, for their flesh is soft and flabby and it is reported to be 
tasteless. But years ago, before the use of kerosene oil, the liver oil of Torpedo nobiliana 
of the western Atlantic was considered equal to the best of sperm oil for illuminating 
purposes. According to classical writers, they were used to some extent as food by the 
ancient Greeks and Romans ; also their shocks were remedies for disease of the spleen, 
for chronic headaches, and for gout; their brains, mixed with alum, were used as a 
depilatory, and the presence of a Torpedo within the room was considered a sure stim- 
ulus to easy delivery for a pregnant woman. ^^ 

Families. The Electric Rays commonly have been united in a single family fol- 
lowing a precedent established more than a century ago.'^ But by 1865 those with 
two dorsal fins, those with one, and those with none had already been distributed among 
three corresponding (unnamed) groups,^^ which have been treated recently as the sub- 
families Torpedininae, Narkinae, and Temerinae respectively.^* This arrangement has 
not only the disadvantage of separating genera that resemble each other closely in other 
respects but of running counter to a dichotomous grouping based on the firmness of 
articulation of the upper and lower jaws and the presence or absence of labial cartilages, 
characters which are probably of greater importance phylogenetically than the number 
of dorsal fins. However, in a general work such as this it would be premature to adopt 
the structure of the jaws as the chief distinguishing character for families until con- 
ditions in this respect are known for a larger proportion of the genera involved.^® The 
scheme based on the number of dorsal fins is followed here as a matter of convenience, 
though we expect that it will be abandoned eventually in favor of the alternate scheme 
based on the jaws. 

Key to Families 
I a. Two dorsal fins. Torpedinidae, p. 87. 

30. Or at least the failure of ichthyologists to have encountered them there. 

31. For a readable account of early knowledge of the Electric Rays, and for classical references to them, see Couch 
(Fish. Brit. Isles, j, 1867: 121-124) and RadclifFe (Fishing from Earliest Times, 1921: iSi, 281, 282). 

32. Miiller and Henle, Plagiost., 1841. 33. Dumeril, Hist. Nat. Poiss., i, 1865: 504. 

34. Fowler, Bull. U.S. nat. Mus., 100 (/j), 1941: 332. 

35. Out of a total of nine known genera of Electric Rays, the presence of labial cartilages that connect the two jaws 
has been established definitely for five genera {Benthobaiis, Narcine, Diplobatis, Discopyge, and Narke), and their 
absence has been determined for two {Torpedo and Hypnos). 



Fishes of the IV ester ?i Nort/i Atlantic 



87 



lb. One dorsal fin or none. 
2 a. One dorsal fin. 



2 b. No dorsal fin. 



Narkidae. 

Indian Ocean in general, South Africa and 
Natal to India, Malaysia, East Indies, southern 
China to Japan, and New Zealand. 

Temeridae. 

Malay Peninsula and Cochin-China. 



Family TORPEDINIDAE 

Characters. Disc ranging from subcircular to elongate. Tail sector as long as body 
sector or shorter, with or without lateral folds. Two dorsal fins, both of them well 
developed. Tips of pelvics either separate from sides of tail or united to latter. Eyes 
either well developed, with evident pupil, or obsolete. Nostrils either simple or sub- 
divided by a cross bridge; separate from mouth, but connected by a groove with fur- 
row surrounding mouth in some species. Mouth protractile as a short tube and with 
labial cartilages in some genera, but widely distensible and without labial cartilages in 
others. Characters otherwise those of the suborder. 

Remarks. The members of this family vary widely in general form, in length of 
tail relative to that of disc, in the condition of the eyes (i. e., whether comparatively 
large and fully functional or obsolete or nearly so), and in the extent to which the inner 
margins of the pelvic fins are united with the sides of the tail. They also cover the 
entire distributional range of the suborder, bathymetric as well as geographic. 



Key to Genera 

la. Mouth widely distensible (Fig. 20) but only slightly protractile; upper and lower 

jaw cartilages not bound together at corners of mouth by labial cartilages, their 

lateral articulation loose; tooth bands firmly connected with jaw cartilages. 

2 a. Tail from tips of pelvics about as long as breadth of mouth when latter is 

closed, its length from center of cloaca to termination only about 1/3 as great 

as distance from cloaca to snout; teeth with two or three cusps. 

Hypnos Dumeril 1852.'^ 
New South Wales, Queensland, 
and West Australia. 
2 b. Tail from tips of pelvics more than three times as long as breadth of mouth 

36. Waite (Rec. Aust. Mus., 4, 1902: 180) proposed the name Hypnarce to replace Hypnos, on the ground that that 
name had been preoccupied by Hypna Hilbner 18 18, for Lepidoptera. But this substitution appears not to be re- 
quired under the International Rules of Zoological Nomenclature, as now accepted. Hypnarea Sharp (Zool. Rec, 
J9, Index, 1903: 9) appears to have been a misprint, the reference being to Hypnarce Waite 1922. Whitley (Fish. 
Aust., J, 1940: 167) points out that the illustration by Shaw and Nodder (Naturalist Misc., 6, 1795: pis. 202, 203) 
of their problematical Lophius monopterygius actually represented a "beach bloated specimen" of the Australian 
Electric Ray later described by Dumeril (Rev. Mag. Zool., (2) 4, 1859: 279, pi. 12) as Hypnos subnigrum; hence, 
the correct name of the species is Lophius monopterygius Shaw and Nodder. We point out in passing that our refe- 
rence of Hypnos to the subdivision of the family that lacks labial cartilages and in which the mouth is widely 
distensible is based on our own observations (Fig. 20). 



8 8 Memoir Sears Foundation for Marine Research 

when latter is closed, its length from center of cloaca more than half as great 
as distance from cloaca to snout; teeth with one cusp only. 

Torpedo Houttuyn 1764, p. 90. 
lb. Mouth narrow, not widely distensible, protractile as a short tube; upper and 
lower jaw cartilages bound together at each corner of mouth by a labial cartilage 
of two triangular elements (p. 108)," the cartilages rigidly articulated one with the 
other; integument bearing the tooth bands only loosely attached to jaw cartilages. 
3 a. Nostril not divided into two separate apertures by a cross bridge midway of 
its length; teeth extending well out onto upper and lower lips and largely 




Figure 20. Hypnos subnigrunt, female, about 185 mm 
long, from Port Jackson, New South Wales (Harv. 
Mus. Comp. Zool., No. 984). Nostrils and mouth, 
about 1.5 X. 

exposed when mouth is closed; lateral terminations of tooth bands not marked 
by deep cross furrows. 

4a. Eye minute, sometimes entirely concealed by overlying integument, 
hence doubtless blind; sides of tail without longitudinal membranous 
fold, at most with a low fleshy ridge. 

Benthobatis Alcock 1898, p. 126. 
4b. Eyes normally developed and functional; each side of tail with a thin 
ribband-like longitudinal fold. 
5 a. Nasal curtain much broader than long. 

6a. Posterior margins of pelvics not joined across base of tail in a 
continuous arc by a membranous connection. 

Narcine Henle 1834, including 
Narcinops Whitley 1940, p. 107.^^ 

37. The presence of these labial cartilages is easily detected by touch in specimens not hardened too much in preservative. 

38. See p. 107, footnote io8. 



Fishes of the JVestern North Atlantic 



89 




Figure 2 1 . Discopyge tschudii. Left, Pelvic fins of female, 264 mm long, from off Argentina, Lat. 46°! 5' S, Long. 
65°02' W (British Museum [Natural History]). Right, Same of male, about 400 mm long, from Argentina (U. S. 
Nat. Mus., No. 53439). 



6 b. Posterior margins of pelvics joined across base of tail by a mem- 
branous connection, either continuous or narrowly indented in 
the midline (Fig. 21). Discopyge Heckel 1845.^* 

Peru, Chile, and the western 
South Atlantic from southern 
Argentina to Rio de la Plata. 
5 b. Joint nasal curtain only slightly broader than long. 

Heteronarce Regan 1921.^° 
Arabian Sea and Indian Ocean south 
to East London, about Lat. 33° S. 
3 b. Nostril divided into two separate apertures by a cross bridge about midway 
of its length; teeth entirely enclosed within the mouth when latter is closed; 

39. It has long been known that females of the type species of Discopyge {Torpedo tschudii Heckel 1845) differ in a strik- 
ing way from those of Narcine in that the pelvics are united by a membranous connection across the base of the 
tail. This was emphasized in the original account of the species by Heckel (in Tschudi, Fauna Peruana, Ichthyol., 
1845: 32, pi. 6) and was verified subsequently by Berg (An. Mus. nac. B.Aires, 4, 1895: 12). While the tips of 
the pelvics of the male project a little farther rearward than in the female, they are similarly joined across the tail 
in an Argentine specimen that we have examined (Fig. 21). The only previous illustration of the male (Stein- 
dachner, Zool. Jb., Suppl. 4, 1898: pi. 21, fig. 14 b) shows the membrane as narrowly interrupted in the midline. 

40. Heteronarce is so close to Narcine that its generic validity is doubtful. According to its author (Regan, Ann. Mag. 
nat. Hist., [9] 7, 192 1 : 414), it differs from Narcine only in the greater length (relative to breadth) of its nasal curtain. 
Presumably, therefore, its type species [Heteronarce garmani Regan) has a protractile mouth with labial cartilages. 
And Annandale's illustration (Mem. Indian Mus., 2, 1909: pi. 3 A, fig. 3) suggests that this is also the case in Nar- 
cine mollis Lloyd (Rec. Indian Mus., I, 1907: 8), which has subsequendy been referred to Heteronarce (Fowler, 
Bull. U.S. nat. Mus., 100 [jj], 1941: 338). 



90 Memoir Sears Foundation for Marine Research 

median sectors of lips, to which the tooth bands extend, marked at either 
end by a deep transverse groove. 

Diplobatis Bigelow and Schroeder 1948, p. 123. 



Genus Torpedo Houttuyn 1764 

Torpedo Houttuyn, Nat. Hist. Dieren, Planten en Mineral., 6, 1764: 453; type species, generally accepted as 
Raja torpedo Linnaeus 1758. Mediterranean. 

Generic Synonyms: 

Narcobatui Blainville, Bull. Soc. philom. Paris, 18 16: 121 ; type species. Raja torpedo Linnaeus 1758; designated 

by Jordan (Genera Fish., I, l<^\1\ 95)-^^ 
Narcobatis Blainville, in Vieillot, Fauna Fran^., Poiss., 1825: 43; emended spelling for Narcobatus Blainville 

1816. 
Narcacion Gill, Ann. N. Y. Lye, 7, 1862: 387; diagnosis, no species mentioned.*- A revival oi Narcacion Klein 

1777- ^ 
Tetronarce Gill, Ann. N. Y. Lye, 7, 1862: 387; type species. Torpedo occidentalis Storer 1843. 
Gymnotorpedo (subgenus) Fritsch, Arch. Anat. Physiol., Leipzig (1886), Physiol. Abt., 1886: 365; type species. 

Torpedo occidentalis Storer 1843, designated by Jordan (Genera Fish., /f., 1920: 435). 
Fimbriotorpedo (subgenus) Fritsch, Arch. Anat. Physiol., Leipzig (1886), Physiol. Abt., 1886: 365; type species, 

Torpedo marmorata Risso 1 8 10. Mediterranean. 
Tetranarce '\oxdi3Xi and Evermann, Rep. U. S. Comm. Fish. (1895), 1896: 222; Jordan, Genera Fish., J, 1919: 

307; emended spelling for Tetronarce Gill 1861. 
Tetronarcine Tanaka, J. Coll. Sci. Tokyo, 2J, igio: 2; type species, T. tokionis Tanaka. Japan. 
Eunarce (subgenus) Fowler, Proc. Acad. nat. Sci. Philad., 62, 1910: 472; type species, Torpedo narke Risso 

1 8 10, Meditterranean, probably equals Raja torpedo Linnaeus 1758. 
Notostrape Whitley, Rec. Aust. Mus., 18, 1932: 372; type species, N.macneilli Whitley, New South Wales; 

probably not distinguishable from Torpedo fairchildi Hector (Colonial Mus. Geol. Surv. N. Z., 1872: 

83, pi. 12, fig. 134). Not seen. 

Generic Characters. Disc broader than long and arcuate laterally, its anterior mar- 
gin either slightly convex, nearly straight, or even slightly concave along median 
sector; length in front of orbits not greater than distance between outer ends of spiracles 
or between outer edges of eyes. Snout soft to the touch. Tail with a low fold along each 
side; its length from center of cloaca to termination considerably less than distance 
from cloaca to snout; its length from tips of pelvics more than twice as great as apparent 
breadth of mouth. Base of first dorsal wholly or partly above bases of pelvics. Second 
dorsal noticeably smaller than first dorsal. Caudal subtriangular, about as large in 
extent below its axis as above, its axis slightly raised. Pelvics with at least their tips free 
from sides of tail. Eyes well developed, pigmented and functional, though small. 

41. The list of species mentioned by Blainville 18 16 in his original diagnosis of Narcobatus did not include Raja torpedo, 
but it did include R. unimaculatus, presumably of Risso 1810, which equals Raja torpedo Linnaeus 1758. The de- 
signation of the latter as type of the genus was therefore justified. 

42. The earliest post-Linnaean use of Narcacion was by Klein (Neuer Schauplatz, 4, 1777: 726); type species Raja 
torpedo Linnaeus 1758. But the International Commission on Zoological Nomenclature has ruled (Opin. Rend., 
Smithson. Publ., No. 1938, 1910: 51; Smithson. misc. Coll., 7 [3], 1925: 27) that neither Klein's generic names, 
nor the revivals of them by Walbaum (P. Artedi Genera Pise. Emend. Ichthyol., 1792) are permissible. Hence, 
Narcacion must date from Gill, who seems to have been the first to define the genus subsequently. 



Fishes of the Western North Atlantic 9 1 

Spiracles separated from eyes by a definite interspace, their posterior margins either 
smooth or with papillae. Midline of back a little posterior to level of spiracles with 
either a pair of conspicuous mucous pores side by side or a group of 5-9. Nostrils 
transverse to slightly oblique, much closer to mouth than to tip of snout, longer than 
space between them. Nasal curtain subquadrangular, considerably broader than long, 
concealing all but outer ends of nasal apertures and extending nearly or quite to upper 
jaw; free posterior margin of nasal curtain smooth, more or less sinuous, or with the 
midsector projecting somewhat. Mouth only slightly protractile, if at all, without labial 
cartilages and therefore widely distensible; the lateral articulations between upper and 
lower jaw cartilages small in area and hardly interlocking; skin at corners of mouth 
loose, with a well marked furrow extending rearward for some distance;*' roof of mouth 
without digitate lobes anterior to the main transverse fold, the latter smooth-edged, 
of nearly uniform breadth throughout its length. Tooth bands not reaching to apparent 
corners of mouth when latter is closed, the integument bearing them rather firmly 
attached to the narrow jaw cartilages. Teeth with one sharp conical cusp, arranged in 
quincunx close together; up to about 64 series in each jaw, with 3 or 4 to 6 or 7 rows 
in function simultaneously. Two rostral cartilages, each a flexible unbranched vertical 
plate, the pair well separated next to the cranium but approximating each other and 
flattened dorsoventrally toward tip of snout, where they and the neighboring antorbital 
cartilages are interconnected by sheets of firm fibrous tissue.** Characters otherwise 
those of the family. 

Size. This genus includes the largest of Atlantic Electric Rays (p. 85). 

Developmental Stages. It has been known for more than a century that the inner 
wall of the uterus in gravid females of some species is set with vascular villi, while in 
that of others it bears a series of longitudinal folds only.*' The later embryonic stages 
of Torpedo are of special interest because the anterior limits of the pectorals in some 
are marked by a well marked notch on either margin of the disc until the young are 
nearly ready for birth (Fig. 23). These notches are described and figured as persisting 
for a time after birth in three species.*" 

Habitat and Range. In the Atlantic the genus Torpedo occurs in water of only a 

43. The degree to which the jaws may protrude and the width to which the mouth may gape when spread to its widest 
(not yet known), can be determined only from observation on living specimens when they are feeding. 

44. Carman's illustrations (Mem. Harv. Mus. comp. Zool., 36, 191 3: pi- 67, figs, i, 2) show the rostral cartilages of 
Torpedo marmorata as rounded bars branched at the tips. But those on his original dissection are as described above. 
Also, the margin of the antorbital cartilages are not as complexly dissected as they are represented in his illu- 
strations. 

45. Davy, Philos. Trans., 1834: 547; Muller, Ober den glatten Hai (1839-1840), 1842: 55. For a recent account of 
the uterine wall of Torpedo torpedo (Linnaeus) 1758, with photographs, see Ranzi (Publ. Staz. zool. Napoli, Jj, 
1934: 359); also see Needham (Biochem. Morphogen., 1942: pi. i, fig. 7). 

46. Small European specimens of Torpedo nobiliana are so pictured by Bonaparte (Icon. Faun. Ital., J, 1835: pi. not 
numbered), by Rey (Fauna Iberica, Feces, r, 1928: 524, fig. 16S), and by Fowler (Bull. Amer. Mus. nat. Hist., 
JO [i], 1936: 120, fig. 48), as are the New Zealand T.fairchildi (as Torpedo fusca by Parker, Trans. Proc. N. Z. 
Inst., 16, 1884: 283, pi. 22, fig. i) and T. nobiliana of the western Atlantic by Carman (Mem. Harv. Mus. comp. 
Zool., 36, 1913: pi. 61, figs. 4-5). Kaempfer's early illustrations (Amoenitatum Exoticorum, 1712: pi. to face 
p. 511, figs. A, B) of T. sinus-persicus Olfers 1831 of the Persian Culf, Red Sea, and Indian Ocean, though crude, 
also show the notched pectorals. 



9 2 Memoir Sears Foundation for Marine Research 

few feet deep down to 50—60 fathoms, but its depth range is known to reach at least 
130 and perhaps 300 fathoms in the Great Australian Bight. ^' 

The geographical range of the genus is known to extend from northern Scot- 
land" and the North Sea (inward as a stray to the Skagerrak and Kattegat)*' to tropical 
West Africa (Senegal) in the eastern Atlantic, including the Mediterranean; from 
Nova Scotia to Cuba in the west. Torpedo is also known from: northern Argentina; 
southern British Columbia to southern California on the Pacific Coast of North America, 
and perhaps from Chile;*" Japan; southeastern and southern Australia and New Zea- 
land; the Indian Ocean region in general, including the Persian Gulf and Red Sea, 
southward to Madagascar, Natal, and the Cape of Good Hope. 

Species. One species, Torpedo mackayana Metzelaar 1919,^1 that falls within the 
limits of the genus in other respects is set apart from all of its generic relations — for 
that matter, from all other members of the suborder — by the presence of small papillae 
along the margins of its pectorals, described as "partly spinous." Indeed, it is so aber- 
rant in this respect that a separate genus may prove requisite for it when it is better 
known. The species that remain, after the subtraction of T. mackayana., have been 
divided by some authors into two genera or subgenera, or even into three, depending 
on whether the margins of the spiracles are smooth or are rimmed with short ten- 
tacular structures or papillae.*- But it is not possible to draw any sharp line even between 
those species in which the spiracles are the most evidently papillate and those in which 
they are perfectly smooth. The gap between them is bridged by Torpedo fuscomaculata 
of the Indian Ocean, in which the spiracular papillae are conspicuous in young spe- 
cimens but decrease in relative size with growth, and by T. torpedo (Linnaeus) 1758, in 
which the papillae are represented in the young by only low knobs that dwindle with 
growth until they are probably wholly obliterated by maturity in some specimens. 

47. McCulloch (Biol. Result. Fish. F. I. S. 'Endeavour', 5 [4], 1926: 159) reports Torpedo f aire hiUi from trawl hauls 
made at 100-220 fathoms in Bass Strait, at 80-120 fathoms and at 130-320 fathoms in the Great Australian Bight. 

48. Sim (Vert. Fauna Dee, 1903: 274) reports (seemingly on good evidence) the capture of several Torpedo nobiliana 
from Wick Bay and its offing, and from Moray Firth for the years 1890-1894, one specimen being 3 feet 9 inches 
long. 

49. Lubbert and Ehrenbaum, Handb. Seefisch. Nordeuropas, 2, 1936: 317; Andersson, Fiskar och Fisk i Norden, 
1942: 275. 

50. It is not clear from Guichenot's (Fauna Chilensis, 2, 1848: 368) brief account whether his Torpedo chilensis actually 
belonged to Torpedo or to Discopyge. 

51. Trop. Atlant. Visschen, 1919: 197, fig. 57, Senegal; known from a single specimen only. 

52. The genus Tetronarce was proposed by GiU (Ann. N. Y. Lye, 7, 1862 : 387) for species with spiracles having smooth 
borders, those with "dentated" spiracles being referred by him to Narcacion, which is equivalent to Torpedo. Fritsch 
(Arch. Anat. Physiol., Leipzig [1886], Physiol. Abt., 18S6: 355) divided Torpedo into two new subgenera, Gymno- 
torpedo for species with smooth spiracles, Fimbriotorpedo for those with papillate ones. Whitley (Rec. Aust. Mus., 
18, 1932: 327) has instituted a new genus Notostrape for a smooth-spiracled species [Notostrape macneilli Whitley 
1932, Australia, apparently equals Torpedo fairchildi Hutton 1872, New Zealand) without mentioning any differ- 
ences to set it apart from the older subdivisions (generic or subgeneric) that had been proposed previously for species 
belonging to this category. Fowler (BuU. U. S. nat. Mus., 100 [xj], 1941: 342) used the old names Torpedo for 
the smooth-spiracled category and Tetronarce for the papillate, having suggested three subgenera earlier (Proc. 
Acad. nat. Sci. Philad., 62, 1910: 472), Narcobatus with papillate spiracles, Tetronarce with smooth, and a new sub- 
genus Eunarce with "spiracle fringes as rudimentary papillae." And Fraser-Brunner (Ann. Mag. nat. Hist., [2] 
2, 1949: 943) recognizes two subgenera, Torpedo for the species with papillate spiracles, Tetronarce for those with 
smooth ones. 



Fishes of the JVester?i North Atlantic 93 

A character more useful as the primary alternative for specific identification is 
whether the rear end of the first dorsal base is considerably posterior to the rear ends 
of the pelvic fin bases or is level with or even anterior to the latter.^^ 

All of the four species of Torpedo known from the Atlantic (7". torpedo Linnaeus 
1758; T.marmorata Risso 18 10; T. nobiliana Bonaparte 1835; ^"^ T.puelcha Lahille 
1928) fall in the group in which the first dorsal base extends rearward well beyond 
the pelvic bases.^* Among these, T. marmorata of the eastern Atlantic and JVIediter- 
ranean is recognizable by the prominent papillae that surround its spiracles; T. torpedo, 
also of the eastern Atlantic and Mediterranean, has a conspicuous color pattern with 
a few large blue-centered ocelli •j'^^ T. nobiliana, from both the eastern and western North 
Atlantic, is recognizable by its plain coloration and smooth spiracles and by an inter- 
space between its second dorsal and caudal that is little longer than the base of the 
first dorsal; T.puelcha from Argentina is identifiable by the long interspace between 
its second dorsal and caudal fins (Key, p. 94), also by plain coloration and smooth 
spiracles. 

Two more Torpedoes are known in which the first dorsal extends far rearward; 
sinus-persicus Olfers 1831, Indian Ocean, with papillate spiracles, and a smooth-spir- 
acled form of problematic identity, from Agulhas Bank (South Africa).^" 

The subdivision of the genus in which the base of the first dorsal terminates at 
a level with the rear ends of the pelvic fin bases, or anterior thereto, known only from 
the western Pacific and Indian oceans so far, includes one group of two species (T.fair- 
childi Hutton 1872 and T. macneilli Whitley 1932) in which the edges of the spiracles 
are smooth and another group of named forms ^' in which the spiracles bear more or 
less obvious papillae. All of these have been united recently with T. marmorata of the 
eastern Atlantic and Mediterranean. ^^ But they all differ from T. marmorata in the 
position of the first dorsal, which has the rear end of its base at least as far anterior as 
the rear ends of the bases of the pelvics (well posterior to the rear ends of the pelvic 
bases in marmorata). They are regarded here as representing two species, T. panthera 
Olfers 1 83 1 and T.fuscomaculata Peters 1855.^" 

53. The number of columns in each electric organ was proposed as the primary specific character by DuBois Reymond 
(Arch. Anat. Physiol., Leipzig [1882], Physiol. Abt., 1882: 400; Rep. Brit. Ass., 1882: 542), for it differs in differ- 
ent Torpedoes and increases but little from birth onward. 

54. The illustration of T. marmorata by Rey (Fauna Iberica, Peces, i, 1928: pi. 9, fig. 2) shows the rear end of the 
base of the first dorsal about even with the rear ends of the bases of the pelvics. Actually it is considerably posterior 
thereto in the specimens we have examined, and it has been so described repeatedly. 

55. For identification of occasional specimens of T. torpedo that may be plain-colored, see Key to Species, alternatives 
8b (p. 94) and lib (p. 95). 

56. Reported first by Thompson (Mar. bid. Rep. Cape Town, 2, 1914: 159) as Torpedo hcbetatu and subsequently 
by Barnard (Ann. S. Afr. Mus., 21 [i], 1925: 89) as T. nobiliana. 

57. Torpedo panthera Olfers 1831; T.fuscomaculata Peters 1855; T.polleni (Bleeker) 1866; T.suessi Steindachner 1S98; 
probably also T.zugmayeri Engelhardt 1912. 

58. By Fowler (Bull. U. S. nat. Mus., 100 [/j], 1941 : 343). The Raja maculata and K. bicolor of Shaw (Gen. Zool., 
5, 1804: 316) from India, based on Russell's illustrations (Fish. Coromandel, i, 1803: pis. i, 2) of his "Temeree" 
and of his "Nalla Temeree," are also listed by Fowler as synonyms of Torpedo marmorata. But he refers the first 
of these to Narcine on another page (BuU. U. S. nat. Mus., 100 [zj], 1941: 333), no doubt correctly; Shaw's Raja 
bicolor was evidently a Narcine also. 

59. We follow Fraser-Brunner (Ann. Mag. nat. Hist., [12] 2, 1949: 943). 



94 Memoir Sears Foundation for Marine Research 

Key to Species^" 

I a. Margins of rear parts of pectorals with two rows of small papillae, partly spinous. 

mackayana Metzelaar 191 9. 
Tropical West Africa. 
lb. Margins of rear parts of pectorals without spinous papillae, the skin naked there, 
as elsewhere. 

2 a. Rear end of base of first dorsal considerably posterior to rear ends of pelvic 
fin bases. 

3 a. Dorsal surface of disc with five (occasionally one, three, or seven) large 
and conspicuous blue-centered and pale-ringed ocelli, symmetrically 
arranged. /or/)efrt'o (Linnaeus) 1758. 

Eastern Atlantic and Mediter- 
ranean. 
3 b. Dorsal surface of disc without conspicuous ocelli though sometimes 
otherwise blotched or spotted. 
4a. Margins of spiracles smooth. 

5 a. Disc about as broad as long. tokionis (Tanaka) 1908. 

Japan. 
5 b. Disc appreciably shorter (less than 80 "/q) than its breadth. 
6 a. Height of caudal fin not greater than distance from its own 
upper origin to origin of first dorsal. 
7a. Interspace between second dorsal and caudal little 
longer than base of first dorsal. 

8 a. Caudal, from upper origin to midpoint of rear 
margin, nearly or quite as long as distance from 
its own origin to origin of first dorsal. 

nohiliana Bonaparte 1835, P- 9^- 
8 b. Caudal, from upper origin to midpoint of rear 
margin, only about as long as distance from its 
own origin to rear end of base of first dorsal. 
torpedo (Linnaeus) 1758. 
Eastern Atlantic, Mediterranean. ^^ 
7b. Interspace between second dorsal and caudal about 1.5 
times as long as base of first dorsal. 

puelcha Lahille 1928. 

Argentina.*^ 

6 b. Caudal higher than distance from its own upper origin to 

origin of first dorsal by an amount about equal to length 

of base of first dorsal. 

9a. Distance between inner ends of nostrils about half as 

60. Based in part on Key by Fraser-Brunner (Ann. Mag. nat. Hist., [12] 2, 1949: 944) to species with papillate spiracles. 

61. Occasional adult specimens may be plain-colored and the spiracular knobs may be visible no longer. 

62. P'or full account with measurements and illustrations, see Lahille (An. Mus. nac. B.Aires, 34, 1928: 332, pi. 5). 



Fishes of the Western North Atlantic 95 

long as distance from tip of snout to mouth; head an- 
terior to spiracles about 2.5 times as long as distance 
between inner ends of spiracles. 

macneiUi (Whitley) 1932. 

Southern Australia.'^ 

9 b. Distance between inner ends of nostrils only about 

Vs as long as distance from tip of snout to mouth; 

head anterior to spiracles only about 1.8 times as long 

as distance between inner ends of spiracles. 

calif or nica Ayres 1855. 
Pacific Coast of North America, 
southern British Columbia to 
southern California. 
4 b. Margins of spiracles with a row of papillae or knobs. 
10 a. Margin of spiracle with seven papillae. 

1 1 a. Spiracular papillae so long that their tips come close 
together in center of spiracular opening; distance from 
midpoint of posterior margin of caudal to origin of first 
dorsal only about half as long as distance from origin of 
first dorsal to tip of snout; nuchal region with a group 
of 5—7, or more, conspicuous mucous pores irregularly 
arranged.^* marmorata Risso 18 10. 

Eastern Atlantic and Mediter- 
ranean, 
lib. Spiracular papillae low, inconspicuous, knob-like; dis- 
tance from midpoint of posterior margin of caudal to 
origin of first dorsal about ^3 as long as distance from 
origin of first dorsal to tip of snout; nuchal region with 
only one pair of conspicuous mucous pores, side by side. 

/or^^i/o (Linnaeus) 1758. 
Eastern Atlantic and Mediter- 
ranean.^^ 
lob. Margin of spiracle with 9 or 10 papillae. 

sinus-persicus Olfers 1 8 3 1 . 

Persian Gulf, Indian Ocean south 
to Natal, Red Sea.6« 

63. Torpedo macneilli from New Zealand has been united with T. fairchildi Hutton 1872 by Fowler (Bull. U. S. 
nat. Mus., 100 [13], 194: : 345)- But the rear end of the base of its first dorsal is considerably posterior to the rear 
ends of the pelvic fin bases, not in a line with the latter, as appears to be the case in the New Zealand T. fairchildi 
(Hutton, Colonial Mus. Geol. Surv. N. Z., 1872: 83, pi. 12, fig. 134). For a detailed account of the Australian 
T. macneiUi, see McCuUoch (Rec. Aust. Mus., 12, 1919: 171, pi- 25, as Torpedo fairchildi). 

64. In all of the other species of the genus that we have examined there is only a pair of these pores, side by side, in 
the nuchal region. 

65. Occasional specimens lack the ocellar spots that are usually the most conspicuous feature of this species. 

66. This characterization of T. sinus-persicus follows Fraser-Brunner (Ann. Mag. nat. Hist., [/a] 2, 1949: 943); it in- 
cludes T. suessi Steindachner 1808, Red Sea. 



96 Memoir Sears Foundation for Marine Research 

2 b. Rear end of base of first dorsal level with rear ends of pelvic fin bases, or 
anterior to them. 
12a. Edges of spiracles smooth. fairchildi Hutton 1872. 

New Zealand.^' 
12 b. Edges of spiracles with a row of papillae or knobs. 

13 a. Eyes about midway between anterior margin of snout and spir- 
acles, or very little nearer the latter; interspace between first and 
second dorsals only as long as distance between second dorsal and 
caudal. panthera Olfers 1831. 

Red Sea; also reported from western Indian 
Ocean from Gulf of Oman to Reunion, 
Mauritius, Madagascar, Mozambique Chan- 
nel, and the Natal Coast,** but perhaps 
not correctly. 
13b. Eyes much nearer to spiracles than to anterior margin of snout; 
interspace between first and second dorsals longer than distance 
between second dorsal and caudal. fuscomaculata Peters 1855. 

Zanzibar, Seychelles, Mauritius, 
Madagascar, and Mozambique.*' 



Torpedo nohiliana Bonaparte 1835 

Electric Ray, Torpedo, Numbfish, Crampfish 

Figures 22, 23 

Study Material. Male, 835 mm long, taken 12-15 n^il^s off:' Plymouth, Mass., in 
about 30 fathoms; two females, one about 1065 mm long from Provincetown, another 
720 mm long from Woods Hole; four specimens, males and females, 800—1,245 mm 
long, from 50—67 fathoms, 60—90 miles southwest of Gay Head, Marthas Vineyard 
Island, Mass.; jaws of a large specimen, and an embryo about 16 cm long from Woods 
Hole; all in the Harvard Museum of Comparative Zoology. Also a male, 6^^ mm long, 
with large claspers, from the North Sea (loaned by British Museum [Natural History]). 

Distinctive Characters. The broad subcircular disc, short snout anterior to the eyes, 
perfectly smooth skin, short thick tail, two well developed dorsal fins, and broad caudal 
fin are enough to mark T. nohiliana off at a glance from all other batoids of the western 

67. See p. 95, footnote 63. 

68. The n^mt panthera was proposed originally (Olfers, Die Gattung Torpedo, 1S31: 15, 16) as a color variety of 
T. marmorata, and Fowler (BuU. U. S. nat. Mus., 100 [rj], 1941 : 343) has relegated it to the synonymy of that 

[species. But Riippel's illustration (Neue Wirbelt. Abyssinia, Fische, 1835: pi. 19, fig. lA) of what seems to be 
the same Red Sea form represents it as differing from marmorata in that the first dorsal fin is whoUy anterior to 
the rear ends of the bases of the pelvics. Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 307) so describes it 
for a specimen in the Harvard Museum of Comparative Zoology that we have re-examined. T. polleni Bleeker 
1866 appears to be a synonym of T. panthera, as does T. zugmayeri Engelhardt 191 2, so far as the published account 
(Zool. Anz., J9, 1912: 617) of it goes, though the latter fails to include any information as to the position of the 
first dorsal relative to the bases of the pelvics. The specimen described by Garman (Mem. Harv. Mus. comp. Zool., 
36, 1913: 309) as Narcacion sinus-persici (Olfers) 1831, on re-examination, also proves to be referable X.0 panthera. 

69. Characterization and range according to Fraser-Brunner (Ann. Mag. nat. Hist., [12] 2, 1949: 945). 



Fishes of the Western North Atlantic 



97 




Figure 22. Torpedo nobiliana, male, 835 mm long, from ofF Plymouth, Massachusetts (Harv. Mus. Comp. Zool., 
No. 36040). A Pelvics of same. B Lateral view of tail of same. C Eye and spiracle of same, about I.JX-D Nostrils 
and mouth of same, about 0.7 X . E Upper teeth from center of jaw of same, about 4.9 x . F Upper teeth near 
right-hand cornerof mouth of same, about 5 X. G Side viewof tail of female, about 1,065 mm long, from Province- 
town, Mass. (Harv. Mus. Comp. Zool., No. 653). 

Atlantic except for the local representatives of the genera Benthobatis^ Narcine, and 
Diplobatis, from which it is easily distinguishable by the wide distensibility of its mouth 
(gape not limited at corners by labial cartilages) and by the rather firm attachment to 
the upper and lower jaw cartilages of the integument that bears the tooth bands. 

Further distinctions, perhaps more convenient as field marks, that set it apart 
from Benthobath are its well developed and functional eyes, relatively much shorter 
snout, and wider disc. Obvious features that differentiate it from Narcine brasiliensis 
are a disc that is considerably wider than long, the anterior margin of which is nearly 
straight or even slightly emarginate rather than convexly rounded, and a first dorsal 
fin that considerably overlaps the bases of the pelvics. The fact that at least the tips of 



9 8 Memoir Sears Foundation for Marine Research 

the margins of its pelvics are free from the sides of the tail separates it (and its Argentine 
representative T.puelcha Lahille 1928) from Discopyge., the western Atlantic range of 
which is confined in any case to midlatitudes in the southern hemisphere. 

Description. Proportional dimensions in per cent of total length. Female, 720 mm 
long, from Buzzards Bay, Massachusetts (Harv. Mus. Comp. Zool., No. 36621). 
Male, 835 mm long, from off Plymouth, Massachusetts (Harv. Mus. Comp. Zool., 
No. 36040). 

Disc: extreme breadth 66.(1, 65.0; length 56,0, 56.0. 

Snout length: in front of orbits 7.1, 7.5; in front of mouth 9.3, 9.1. 

Orbits: horizontal diameter 2.9, 3.0; distance between 3.9, 3.8. 

Spiracles: length 2.8, 2.6; distance between 4.9, 5.4. 

Mouth: breadth 6.7, 6.4. 

Exposed nostrils: distance between inner ends 4.3, 4.4. 

Gill openings : lengths, ist 2.5, 2.0; 3rd 3.0, 2.4; 5th 1.9, 1.8; distance between 
inner ends, ist 15.7, 18.0; 5th 14.6, 15. i. 

First dorsal fin: vertical height 6.0, 7.2; length of base 5.8, 6.2. 

Second dorsal fin: vertical height 3.1, 4.0; length of base 3.8, 1,.^. 

Caudal fin: upper anterior margin 17.6, 18.3. 

Distance: from tip of snout to center of cloaca 58.3, 57.4; from center of cloaca 
to tip of tail 41.7, 42.6. 

Interspace between: ist and 2nd dorsals 3.9, 4.0; 2nd dorsal and caudal 6.8, 6.6. 

Disc about 1.2 times as broad as long, its edge fleshy and thick in front but thin- 
ning progressively rearward to posterior parts of pectorals, where it is exceedingly thin; 
the median sector of its anterior contour nearly straight, or even slightly emarginate 
there in some specimens; its outer margins broadly and evenly rounded, with the 
embryonic pectoral notches persisting for a short time after birth in some European 
specimens, hence probably in American examples also.'"' Axis of greatest breadth about 
55—60 "/o of distance rearward from snout toward axils of pectorals. Tail from center 
of cloaca about 67— 74 "/o as long as distance from cloaca to snout; noticeably stout 
anteriorly, about twice as broad as deep at rear bases of pelvics, tapering thence rear- 
ward to caudal; moderately rounded below as well as above; each side of tail a little 
below the midlevel, with a low cutaneous fold extending from about opposite origin 
of second dorsal rearward for a short distance beyond origin of caudal. 

Skin wholly naked above as well as below. Nuchal region with a pair of con- 
spicuous mucous pores side by side. 

Snout in front of eyes about as long as distance between outer margins of eyes; 

70. This was the case in specimens of about 244 mm (9.66 in.) and 2S1 mm (11.06 in.) long, figured respectively by 
Bonaparte (Icon. Faun. Ital., 3, 1835: pi. not numbered) and Rey (Fauna Iberica, Feces, i, 1928: 524, fig. i68). 
An Irish specimen also has been described and pictured by McCoy (Ann. Mag. nat. Hist., 6, 1841: 407, 408, as 
T. emarginata) as having notches still visible at a length of 2 feet 8 inches. But such notches have not been reported for 
any American specimen subsequent to birth. We should point out that the indentation in the left-hand margin of 
the disc of the specimen pictured in Fig. 22 is far posterior to the anterior limits of the pectoral, and hence it is 
the result of deformation or injury rather than the persistence of one of the embryonic pectoral notches. 



Fishes of the Western North Atlantic ()() 

its length in front of moutli about twice as great as distance between outer ends of 
exposed nostrils. Eyes noticeably small but fully developed; horizontal diameter about 
1/3— i/g as long as distance between orbits, or a little less. Spiracles moderately oblique, 
sloping rearward inwardly, about as long as orbit and about 1.5 times as long as 
horizontal diameter of eye; posterior to eyes by a distance a little longer than eye; 
their margins perfectly smooth. Second to fourth gill openings (about equal in length) 
a little more than Va as long as breadth of mouth; first about 85 "/o and fifth about 
75 */o as long as third or fourth (the longest); distance between inner ends of fifth gill 
openings about 2.0-2.3 times as great as breadth of mouth, and 82—92 "/o as great 
as distance between inner ends of openings of first pair. Nostrils moderately oblique, 
close to mouth but entirely separate from it; distance between their inner ends (con- 
cealed by the nasal curtain) about half as great as distance between orbits and about 
1/3 as great as breadth of mouth; joint nasal curtain subrectangular, about three times 
as broad as long, roofing over the inner half to third of each nasal aperture; its free 
rear margin somewhat sinuous, sometimes with a shallow median notch; posterior 
margin of nostril with two semilunar expansions, the free margin of the outer (anterior) 
directed toward the nasal aperture, that of the inner (posterior) away from it and toward 
the mouth. Mouth moderately arched, its apparent breadth when closed about 1.7 
times as great as distance between orbits, the furrow from each corner extending rear- 
ward for a distance slightly more than 1/3 as great as breadth of mouth when mouth 
is closed, perhaps entirely obliterated when mouth is open. 

Teeth increasing in number of series with growth from H at a length of 655 mm 
(North Sea specimen) to || at 835 mm and to |j at 1,040 mm (American specimens); 
cusps sharp or slightly blunted, about as long as breadth of bases in both sexes, directed 
into mouth, those on each side of jaws curving slightly toward corner of mouth; up to 
seven rows in function simultaneously in medium and large specimens, these succeeded 
by several additional replacement rows that are already of the final shape though still 
soft and covered over by folds of skin from roof and floor of mouth. 

First dorsal about as high vertically as long; subtriangular with rounded corners; 
anterior margin convex toward apex, distal margin nearly straight, free lower posterior 
margin about half as long as base; its length along anterior margin about equal to 
distance from posterior margin of spiracle to tip of snout; its origin anterior to axils 
of pelvics by a distance ^Is—^U ^s long as its base in female, about ^/g as long as its base 
in mature male. Second dorsal only about V2— Vs ^^ large as first dorsal in linear dimen- 
sions and relatively lower, its anterior margin more sloping, its lower posterior free 
margin relatively longer. Interspace between first and second dorsals about ^/j as long 
as base of first dorsal. Interspace between second dorsal and caudal about as long as 
base of first dorsal or a little longer. Caudal approximately an equilateral triangle; its 
upper and lower margins weakly convex, its posterior margin nearly vertical and weakly 
emarginate; corners broadly rounded; its axis slightly raised; its length from origin to 
midpoint of rear margin about as great as distance from origin of first dorsal to rear 
tip of second dorsal; its upper origin about opposite its lower origin. Pelvics with 



lOO 



Memoir Sears Foundation for Marine Research 



broadly convex outer margins'^ but with abruptly rounded or subangular posterior 
corners, their inner posterior margins free from sides of tail for a distance about 1/4 
as great as distance from origin of pelvic to its tip; their length (origin to tip) about 
as great as distance from origin of first dorsal to origin of caudal. 

Each electric organ may have as many as 1,000—1,100 columns. 

Color. Dark chocolate to purplish brown above, or even nearly black, either uniform 
or with a few obscure darker spots. Generally white below, but with edges of disc and 
pelvics of same hue as upper surface; tail with irregular dark margins. 

Size. At birth the length of Torpedo nobiltana is probably between 200 and 250 mm, 
for the embryonic pectoral notches still showed on Mediterranean specimens of 244 
and 281 mm (see p. 91) and the umbilical scar was still evident on an Algerian 
specimen 261 mm long.'^ The smallest free-living American specimen recorded so far 
had already grown to a length of 610 mm. It is not known at just what length this 
species matures sexually, but it is likely that all specimens that have grown to two 
feet or more are close to sexual maturity, if they have not already reached it. One, 
netted off Woods Hole, Massachusetts, was 56^/4 inches long," and the largest for 
which the dimensions are definitely recorded, taken at Cape Lookout, North Carolina 
in February, measured 60^/2 inches in length by 41 inches in breadth. But larger ones 
are caught in the Woods Hole region,'* while Mediterranean specimens 63 inches 
(1.6 m)'* and 5 feet 11 inches (1.8 m)'^ lo"g have been reported. 

The average weight of specimens taken at Woods Hole is reported to be about 
30 pounds, and most of those taken anywhere on the coast weigh less than 75 pounds. 
But the relationship between length and weight varies considerably, depending on the 
fatness of the fish. This is illustrated by the fact that a specimen only 47.5 inches long 
from Chesapeake Bay, examined by us, weighed about 100 pounds, whereas one of 
48.5 inches from Cape Cod Bay weighed 81 pounds," and another of 52 inches from 
the southern part of Georges Bank weighed only 78 pounds. The North Carolina 
specimen mentioned in the preceding paragraph (60.5 in. long) weighed 125 pounds; 
one of about 134 pounds (61 kg) has been reported from Cape Lookout, and many 
years ago another of 144 pounds was brought from Nantucket to the station of the 
U. S. Bureau of Fisheries at Woods Hole, Massachusetts. '^ The heaviest ones taken 

71. The pelvics of the specimen pictured in Fig. 22 show an interesting divergence on the two sides of the body, the 
outer contour being the more evenly rounded on the right-hand fin. 

72. The original account of this specimen (Guichenot, Explor. Alger., Poiss., 5, 1850: 131, pi. 8, as Torpedo nigra) 
did not mention its length, which was given subsequently by Dumeril (Hist. Nat. Poiss., i, 1865: 512). 

73. Reported to us by F. E. Firth, U. S. Bureau of Fisheries. 

74. Captain Donald Campbell, a fisherman of many year's experience, informs us that he has seen them up to five or 
six feet across near Woods Hole, Massachusetts. 

75. Dumeril, Hist. Nat. Poiss., j, 1S65: 511, by ref. to Bonaparte (probably Obs. sur la torpille, Actes i Congr. Pise, 
1840: 18; not seen). 

76. Gibert, Bull. Inst. Catalon. Hist, nat., 1911: 102; Rey, Fauna Iberica, Peces, j, 1928: 526. 

77. Reported by Dorothy Snyder. 

78. Recorded weights of a few British specimens of different sizes are as follows: 13 pounds at 28 inches, 25 pounds at 
33 inches, 42 pounds at about 40 inches, 82 pounds at 46 inches, and no pounds for one between 60 and 64 inches 
long, the latter the largest that had been reported from British waters up to 1925 (Jenkins, Fish. Brit. Isles, 1925: 
329)- 



Fishes of the Western North Atlantic 



lOI 



near Provincetown, at the tip of Cape Cod, Massachusetts, were estimated long ago 
by a fisherman of keen observation as reaching 170 to 200 pounds.'^ 

Developmental Stages. The number of young has not been recorded for any 
American specimen.^" The disc of an embryo about 140 mm long, taken from a female 
that was landed at Woods Hole, Massachusetts, is deeply notched on both sides about 
opposite the mouth, marking the anterior limits of the pectorals. At this stage, still 




Figure 23. Torpedo nobiliana, embryo, 159 mm long, 
from Woods Hole, Mass. (Harv. Mus. Corap. ZooL, 
No. 1016), about 0.9 natural size. 



considerably previous to birth, the posterior margins of the pectorals merge insensibly 
with the sides of the trunk, the pelvics are still small, and the curtain-like expansion of 
the anterior margins of the nostrils has not developed and hence the entire lengths of 
the nasal openings are freely exposed. But the caudal is already of approximately adult 
form, the relative sizes of the dorsals are already established, the eyes and spiracles 
are approximately of adult proportions, and the older series of teeth already have cusps. 
The embryo lies in the uterus with the tail bent forward ventrally and the caudal fin 
flat against the left-hand pectoral (Fig. 23). 

79. This estimate by Captain N. E. Atwood, first quoted by Storer (Amer. J. Sci., 4$, 1843: 167), has been the basis 
for many subsequent statements that this Torpedo reaches a weight of 200 pounds. 

80. A European female has been described as having 60 embryos (Dumeril, Hist. Nat. Poiss., i, 1865: 512, by ref. 
to Bonaparte [probably Obs. sur la torpille, Actes i Congr. Pise, 1840; 18; not seen]). However, this number is 
much larger than that reported for other Torpedoes. 



I02 Memoir Sears Foundation for Marine Research 

Habits. Practically nothing is known of the way of life of T. nobiliana. Probably 
it spends most of its time on the bottom partially buried in sand or mud, awaiting 
whatever prey may come within its reach. But the rather frequent captures of Tor- 
pedoes in pound nets near Woods Hole, Massachusetts, and the netting of at least one 
in a seine, shows that they move about more or less actively and may swim up from the 
sea floor, probably in pursuit of small fish. 

A few of the specimens that have been examined at Woods Hole have contained 
fish: a summer flounder {Paralichthys dentatus) about 14V2 inches long (37 cm) was 
taken from one. A two-pound eel and a one-pound flounder were found in one from 
British waters, a salmon weighing between four and five pounds was found in another, 
a red mullet (Mullus surmuletus) and a plaice {Phuronectes platessa) were recovered in 
a third, and the remains of a small Spotted Dogfish (genus Scyliorhinus) were found 
hanging from the mouth of still another British specimen when caught. ^^ Evidently 
the wide distensibility of the jaws enables it to swallow fishes much larger than might 
seem possible from the apparent breadth of its mouth when closed, and its sharp 
recurved teeth serve not only to prevent its victim from escaping but as an aid in 
swallowing. It is believed generally that it stuns the fishes on which it preys by means 
of its electric discharges. Otherwise it is difficult to conceive how a fish as sluggish 
as this could capture other active fishes. However, the actual process has not been 
observed. Also, it has been reported repeatedly as caught by hook and line on dead 
bait of one kind or another. 

Its scarcity on the grounds where otter trawlers normally operate, though within 
its known range, suggests that it is most apt to be found on flat sandy or muddy bottom, 
as in the case of its representative in British waters.'^ 

Of its breeding habits in American waters we know only that a female containing 
one embryo was taken near New York in June, that another gravid female was cap- 
tured at Woods Hole in summer,^^ and that the eggs of this species are said to be 
nearly ripe late in June,^* which suggests that the period of gestation is nearly a year. 
It seems that the young are born offshore, for the smallest individual recorded from 
American inshore waters, a New Jersey specimen, was 610 mm long.^* 

The statement has long been current that the shock from a large Torpedo of this 
species in rested condition may be strong enough to throw a grown man to the ground,** 
and it is stated also that a shock may be received through the wet handle of a gaff- 
hook or harpoon. Antedating the scientific naming of the New England Torpedo is 
the anecdote*' of a dog, who, being in the habit of wading in shoal water on a Cape 

81. Day, Fish. Gt. Brit., 2, 1880-1884: 331. 82. Day, Fish. Gt. Brit., 2, 1880-1884: 331. 

83. Our Study Material includes one embryo from this female, but no record was kept of the total number that she 
contained. 

84. Sumner, Osburn and Cole, Bull. U.S. Bur. Fish., 31 (2), 1913: 739. 

85. Fowler, Proc. Acad. nat. Sci. Philad., 7.^, 1922: 3. 

86. It seems that most of the published statements to this effect for Torpedo nobiliana in American waters lead back to 
Atwood's report that he had "received many powerful shocks" from Torpedoes taken near Provincetown, Mas- 
sachusetts, "which have thrown me upon the ground as quick as if I had been knocked down with an ax" (see 
Storer, Amer. J. Sci., 45, 1843: 167). 87. Storer, Rep. Fish. Rept. Birds Mass., 1839: 201. 



Fishes of the IV es tern North Atlantic 103 

Cod beach to catch flounders, was so shocked by a Torpedo that it ran away howling 
and could never be persuaded to go fishing again. 

However, shocks of a strength even approaching what is suggested by such reports 
are to be expected only from Torpedoes of the largest size, and then only if they 
are in a rested condition while in the water or soon after capture. But it is likely 
that the voltages of 170-220 that have been recorded recently for T. nobiliana after 
several weeks in a live well are considerably lower than the voltage that these same 
specimens might have delivered had they been at liberty and had they fed recently.*' 
A slight benumbing or tingling sensation is the most that we have felt from medium- 
sized specimens lying on the dock at Woods Hole. 

The great majority of American records for T. nobiliana are of specimens that 
have been either seined or taken in pound nets in a few feet of water or stranded on the 
beach. It has even been taken in sounds behind barrier beaches, as near Manteo on 
Roanoke Island, North Carolina, and on the coast of New Jersey. But the number of 
such captures has been so small as to make it likely that they represent merely the 
fringe of a population that centers farther out in depths of perhaps 10—40 fathoms, 
as is said to be true of it in British waters.*' Our Study Material includes one trawled 
in Massachusetts Bay at 30 fathoms and another taken by Captain Jared Vincent 
(with 12 others, all large) in February 1949 about 60 miles off Marthas Vineyard in 
50 fathoms. Another large specimen, about four feet wide, was taken in January 1949°" 
with three smaller ones in 50—60 fathoms in the Hudson Gorge off New York. In 
June 1 931" a large one was netted in 17 fathoms some 30 miles from the nearest land 
off Woods Hole, Massachusetts. And specimens have been taken on La Have Bank 
and on the southwest part of Georges Bank in water at least 30—50 fathoms deep. 

Except for a single specimen taken at Cape Lookout, North Carolina, in February, 
the records of its capture close inshore have been between spring and November. It 
appears that the few individuals that do stray close inshore to the northward during 
the warmer half of the year may do so as early in the season at one point along the 
coast as at another. October and November have been described as the months of 
most frequent occurrence for them off southern Massachusetts and along Cape Cod. 
The paucity of records for it south of Cape Lookout suggests that it cannot survive 
subtropical temperatures for any length of time. 

Numerical Abundance. The coastwise sector from the tip of Cape Cod to the offing 
of Rhode Island is the only one in the western Atlantic where reports of its presence 
rest on more than an occasional capture, and even there the numbers taken are small 
as compared with some of the Skates (genus Raja). At most, 60—80 are said to have 
been taken yearly along Cape Cod when they have been most plentiful there (p. 104), 
and perhaps 15-20 during the spring on the south shore of Marthas Vineyard. As 
many as 12 have been taken in one lift of a fish trap near Woods Hole and in Vineyard 

88. See Coates and Cox (Zoologica N. Y., 2y, 1942: 28) for voltages, current, and power delivered by adult T. nobiliana 
with various resistances. 

89. Day, Fish. Gt. Brit., 2, 1880-1884: 331. 90. Reported to us by Captain Frank Janssen. 
91. Personal communication from F. E. Firth, U. S. Bureau of Fisheries. 



1 04 Memoir Sears Foundation for Marine Research 

Sound, but more commonly only 1-4, if any, are captured. "^ All reports from farther 
west and south have been based on only one or two individuals there, at long intervals. 
Numbers such as these are not large enough to suggest the presence of a self-supporting 
local stock. For this reason we believe that the center of population for T. nobUiana 
lies farther out on the continental shelf (p. 103). 

It has been known for a long time that Torpedoes are encountered much more 
frequently in some years than in others along the northeastern seaboard of the United 
States. *3 Thus, it is said that Torpedoes were unusually abundant near Provincetown 
during the period from 18 19 through the succeeding four or five years, with 60-80 
taken there yearly. But according to reports, the catch for Cape Cod during the ten- 
year period preceding i 845 did not exceed 30 in all.^* The next incursion of Torpedoes 
near Provincetown appears to have taken place in 1845, with a dozen or so reported. 
Any fluctuations that may have taken place from year to year thereafter seem to have 
attracted no attention until the summer of 1896, when several were collected along 
the coast of Maine. Within a short period during the summer of 1941, several were 
taken at Cape Lookout, North Carolina, where only three specimens had been reported 
in scientific literature previously (p. 105). It is not known whether these fluctuations 
are reflections of corresponding variations in the strength of the offshore population 
or are merely an indication of the chance distribution of individuals that come close 
inshore from year to year. 

Relation to Man. At present the Torpedoes that are taken along the eastern sea- 
board of the United States are of no commercial value. It is rumored that local fishermen 
at an earlier date considered their oil a useful remedy for muscular cramps when applied 
externally and for cramps of the stomach when taken internally.^^ We are informed^" 
that small amounts were also used for lubricating farm machinery. The few specimens 
that are landed at Woods Hole attract the attention of workers at the Marine Biological 
Laboratory because of the electric organs and their discharges. 

Range. Both sides of the North Atlantic, from northern Scotland to the Mediter- 
ranean, Azores, Madeira, and tropical West Africa" in the east; from southern Nova 
Scotia, La Have Bank, mouth of the Bay of Fundy and Georges Bank to North Carolina 
in the west; also reported from the Florida Keys and from Cuba.^* 

92. Captain Donald Campbell advises us that catches of 1-3 Torpedoes per day were made on 20 out of 50 days (33 
fish) between June 16 and September 17, 1947 off the southeastern shore of Vineyard Sound in 30 feet of water 
in a fish trap set with runner extending in toward shore to the 20-foot line. 

93. The most instructive information in this respect dates back many years. 

94. Storer, Mem. Amer. Acad. Arts Sci., N. S. 9, 1867: 245; also Fishes Mass., 1867: 271. 

95. Reports by Captain N. E. Atwood, in Storer (Amer. J. Sci., 45, 1843: 167). 

96. By Captain Donald Campbell. 

97. For tropical West African references, see Fowler (Bull. Amer. Mus. nat. Hist., jo [i], 1936: 121; -jo [2], 1936: 
1157) and Poll (Result. Sci. Exped. oceanogr. Beige Cot. Afr. Atlant. [1848-49], 1951: 79). The relationship of 
T. nobiliana to the Torpedo reported under that specific name from the vicinity of the Cape of Good Hope (Bar- 
nard, Ann. S. Afr. Mus., 2/, 1925: 89, by ref. to Thompson [Mar. biol. Rep. Cape Town, 2, 1914: 159, as Tor- 
pedo hebetans]; Smith, Sea Fish. S. Afr., 1949: 75) remains to be determined. 

98. It has been suggested (Dubois Reymond, Arch. Anat. Physiol., Leipzig [1882], Physiol. Abt., 1882: 400; Rep. 
Brit. Ass., 1882: 594) that the large Torpedoes that are taken from time to time in British waters may be strays 
that have drifted across from America in the Gulf Stream. We think it far more likely that the populations of 
T. nobiliana in the two sides of the Atlantic are entirely independent of each other. 



Fishes of the IVestern North Atlantic 105 

Occurrence in the Western Atlantic. The most northeasterly records for T. nobiliana 
off the American Coast are: one from Eastport, Maine, at the mouth of the Bay of 
Fundy, another, "presumably of this species,""^ taken in St. Margaret Bay (Nova 
Scotia) some 30 years ago; and a third caught on a long line set for cod on La Have 
Bank in 1890, reported by an observer v?ho was undoubtedly well acquainted with 
T. nobiliana at Woods Hole."" It has also been taken near Seguin Island, Maine, at 
the mouth of Casco Bay; in the vicinity of Cape Ann, Massachusetts; in the southern 
side of Massachusetts Bay near Provincetown and along the outer coast of Cape Cod 
(see Study Material, p. 96). Localities so distributed show that it is to be expected 
anvwhere along the coast of the Gulf of Maine. Also, a 58-inch specimen was taken 
on the southwest part of Georges Bank in December 1930. Torpedoes are caught 
yearly in Buzzards Bay and in Vineyard Sound, while records of long standing indicate 
its presence off Nantucket Island and the southern shore of Marthas Vineyard. It is 
reported commonly off Rhode Island and in the vicinity of Block Island, and it has 
been reported from the vicinity of New York and at localities thence southward, show- 
ing that odd specimens are to be expected anywhere along New Jersey and Maryland 
and within the mouth of Chesapeake Bay. 

The status of T. nobiliana southward from the region of Cape Lookout is doubt- 
ful. "i An "electric fish" reported from the vicinity of Charleston, South Carolina"^ 
actually may have been a Narcine brasiliensis, as seems to have been true of some of 
the early records that ostensibly refer to Torpedo on the North Carolina Coast. ^"^ The 
only mention of its occurrence in Florida waters known to us"* fails to state whether 
any specimens were actually examined. It has been mentioned repeatedly as ranging 
to Cuba, but all these reports appear to emanate from an early reference to a small 
Electric Ray (360 mm long) of doubtful identity taken near Havana many years ago."' 
And the recent inclusion of it as a member of the fish fauna of the northern Gulf of 
Mexico"^ also appears to lead back to this doubtful Cuban record or to the Florida 
Keys report mentioned above; at least we find no mention of it in the several published 
lists of fishes for the west coast of Florida or for Texas. 

Synonyms and Western Atlantic References: 

Raja torpedo MitchiU, Trans. Lit. philos. Soc. N. Y., J, 1815: 476 (reported, Georges Bank and near Block 
Island, by fishermen). 

Torpedo nobiliana Bonaparte, Icon. Faun. Ital., J, 1835: pi. not numbered (descr., ill. of small specimen, Italy); 
Storer, Amer. J. Sci., ^, 1843: 214 (4 ft. 2 in. specimen, WeUfleet, Mass.); Ann. Mag. nat. Hist., 11, 
1843: 326 (same as preceding); Proc. Boston Soc. nat. Hist., j, 1844: 94 (same specimen as Storer, 
1843); Fowler, Proc. biol. Soc. Wash., JJ, 1920: 145 (listed. New Jersey); Proc. .'^cad. nat. Sci. Philad., 

99. Vladykov and McKenzie, Proc. N. S. Inst. Sci., 19, 1935: 52. 
100. By G. F. O. Hansen, then second mate and later master of the U. S. Bureau of Fisheries schooner Grampus 

(Smith and Kendall, Rep. U.S. Comm. Fish. [1896], 1897: 169). 
loi. Two were taken in June of 1941, and several more during the next few weeks, at Cape Lookout, North Carolina. 

102. Jordan and Gilbert, Proc. U.S. nat. Mus., 5, 1883: 584. 

103. Radcliffe, Bull. U. S. Bur. Fish., 24, 1916: 270. 

104. "Found occasionally among the Keys," Fowler (Proc. Acad. nat. Sci. Philad., 5S, 1906: 80). 

105. By Poey (Repert. Fisico-Nat. Cuba, 2, 1868: 458) as Tetronarce occidentalis ; identification doubtful. 

106. Gowanloch, Bull. La. Conserv. Dept., 21, 1932: 86; Bull. La. Conserv. Dept., 23, 1933: 238. 



1 06 Memoir Sears Foundation for Marine Research 

y4, 1922: 3 (specimen 610 mm, New Jersey); Hildebrand and Schroeder, Bull. U. S. Bur. Fish., 
4J, 1928: 62 (descr., meas., weight, Chesapeake Bay); Truitt, Bean and Fowler, Bull. Md. Conserv. 
Dep., 3, 1929: 31 (diagn., listed, Maryland); Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1945: 99 
(listed, Manteo, N. Carolina). 

Cramp Fish, Storer, Rep. Fish. Rept. Birds Mass., 1839: 201 (Wellfleet, Mass., electric shocks). 

Torpedo zvahhii Thompson, Ann. Mag. nat. Hist., 5, 1840: 292 (descr., Ireland). 

Torpedo emarginata McCoy, Ann. Mag. nat. Hist., 6, 1 841 : 407 (descr. of specimen 2 ft. 8 in. long with em- 
bryonic pectoral notches; Ireland). 

Numb Fish or Cramp Fish, DeKay, Zool. N. Y., 4, 1842; 378 (inch by ref. to Mitchill, 1815). 

Torpedo occidentalis Storer, Amer. J. Sci., //5, 1843: 165, pi. 3 (descr., ill., size, habits, nos., electric shocks, 
Provincetown, Mass.); Linsley, Amer. J. Sci., ^7, 1844: 79, footnote (listed, E. side Cape Cod); Storer, 
Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 516 (in synopsis); Proc. Boston Soc. nat. Hist., 2, 1848: 
71 (living specimen), 170 (nos., south shore of Marthas Vineyard); Dumeril, Rev. Mag. Zool., 4, 1852: 
245 (descr., after Storer, 1843); Hist. Nat. Poiss., j, 1865 : 513 (descr., color, after Storer); Storer, Mem. 
Amer. Acad. Arts Sci., (2) 9, 1867: 247, pi. 39, fig. 5; also Fishes Mass., 1867: 271, pi. 39, fig. 5 (descr., 
size, ill., habits, electric shocks, nos., season, util., C. Cod, Marthas Vineyard); Giinther, Cat. Fish. Brit. 
Mus., 8, 1870: 448, footnote i (listed, U. S.); Baird, Rep. U. S. Comm. Fish. (1871-1872), 1873: 
826 (listed, Woods Hole); Gill, Rep. U. S. Comm. Fish. (1871-1872), 1873: 812 (listed, C. Cod to 
Florida); Uhler and Lugger, Rep. Comm. Fish. Maryland, ed. r, 1876: 188; ed. 2, 1876: 159 (listed, 
off Chesapeake Bay); Goode and Bean, Bull. Essex Inst., II, 1879: 28 (listed, near C. Ann, Mass.); 
du Bois Raymond, Arch. Anat. Physiol., Leipzig (1882), Physiol. Abt., 1882: 400; Rep. Brit. Ass., 
1882: 592, 594 (no. of columns in electric organ as spec, char., suggests specimens carried to England 
by Gulf Stream); Jordan and Gilbert, Bull. U. S. nat. Mus., 16, 1883 : 39 (diagn., listed, Atlant. Coast); 
Jordan, Rep. U. S. Comm. Fish. (1885), 1887: 799 (listed); Wilson, Proc. U. S. nat. Mus., JJ, 1907: 
373 (copepod parasites, season, Provincetown, Mass.); Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 4 
(3), 1913: loi (range); Norman and Eraser, Giant Fishes, 1937: 64 (listed, Atlant. Coast U. S.); Coates 
and Cox, Zoologica N. Y., 2J, 1942: 25 (electric discharges, nos., season, weight, C. Lookout, N. Caro- 
lina). 

Torpedo nigra Guichenot, Explor. Alger. Poiss., 5, 1850: 131, pi. 8 (ill. Algeria; for size, see Dumeril [Hist, 
nat. Poiss., I, 1865: 512]). 

Narcacion occidental Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861: 61 (listed, E. Coast N. America). 

Tetronarce occidentalis Gill, Ann. N. Y. Lye, 7, 1862: 387 (type of new genus Tetronarce)\ Jordan and Ever- 
mann. Bull. U. S. nat. Mus., 47 (i), 1896: 77 (descr., C. Cod to Cuba); Smith and Kendall, Rep. U. S. 
Comm. Fish. (1896), 1898: i6g (Maine, LaHave Bank); Jordan and Evermann, Bull. U. S. nat. Mus., 
47 (4), 1900: pi. II, fig. 33 (ill.); Linton, Bull. U. S. Fish Comm., ig, igoi: 274, 432 (stom. con- 
tents, nos., season, female with young, cestode parasites. Woods Hole); Bean, Bull. N. Y. St. Mus., 60, 
Zool., 9, 1903: 51 (descr., color, habits, range); Jordan, Guide to Study Fish., J, 1905: 554 (size, cf. 
European T. nobiliana and T. calif ornicd)\ Fowler, Proc. Acad. nat. Sci. Philad., ^8, 1906: 80 (listed, 
Florida Keys); Rep. N. J. Mus. (1906), 1907: 260 (listed. New Jersey); Smith, N. C. geol. econ. Surv., 
2, 1907: 43 (descr., range, size, but previous reports from N. Carolina probably Narcine)\ Sullivan, Bull. 
U. S. Bur. Fish., 2'], 1907: 10, 13, 15-16, 22 (morphol. of intestine, physioL); Kendall, Occ. Pap. Bos- 
ton Soc. nat. Hist., 7 (8), 1908: 13 (New England locals.); Nichols, Abstr. Proc. Linn. Soc. N. Y., 
20-23, 1913- 92 (listed, near New York); Sumner, Osburn and Cole, Bull. U. S. Bur. Fish., JJ, 1913: 
739 (Woods Hole region and Nantucket, weight); Coles, Copeia, 32, 1916: 46 (listed, C. Lookout, 
N. Carolina); Copeia, 151, 1926: 105 (listed, size, C. Lookout, N. Carolina); Nichols and Breder, Zoo- 
logica N. Y., 9, 1927: 30 (New York and south. New England, size); Jordan, Manual Vert. Anim. NE 
U. S., 1929: 17 (diagn., Maine southward); Townsend, Guide N. Y. Aquarium, 1937: 41 (near New 
York). 

Torpedo (no scientific name) Couch, Fish. Brit. Isles, J, 1867: 119, pi. 39 (habits, electric shocks, ill., Gt. 
Britain). 

Torpedo hebetans (in part) Giinther, Cat. Fish. Brit. Mus., 8, 1870: 449 {T. nobiliana Bonaparte 1835 included 
in synonymy of T. hebetans Lowe 1841). 

Torpedo [Gymnotorpedo) occidentalis Fritsch, Arch. Anat. Physiol., Leipzig (1886), Physiol. Abt., 1886: 362, 
364 (no. columns in electric organ); Elektr. Fische, 2, 1890: 14, pis. i, 3 (cf. other species, no. electric 
columns, nerve syst., habits, range by ref. to Storer, suggests carried to England by Gulf Stream). 



Fishes of the IVestern North Atlantic 107 

Tetranarce occidentalis Jordan and Evermann, Rep. U. S. Comm. Fish. (1895), 1896: 222 (listed, C. Cod 
to Cuba); Smith, Bull. U. S. Fish Comm., IJ, 1898: 89 (nos., season. Woods Hole region); Tracy, 
36 Rep. R. I. Comm. inl. Fish., 1906: 47 (off Rhode Island); 40 Rep. R. I. Comm. inl. Fish., 1910: 
62 (same as Tracy, 1906); Jordan, Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 27 
(listed, C. Cod to Cuba); Wilson, Bull. U. S. nat. Mus., 158, 1932: 429 (copepod parasites. Woods 
Hole and Provincetown, Mass.); Gowanloch, Bull. La. Dep. Conscrv., 23, 1933: 238 (electric organs, 
size, listed Gulf of Mexico). 

Narcacion nobilianus Garman, Mem. Harv. Mus. comp. ZooL, j6, 1913: 310, pi. 25, fig. 2, pi. 61, figs. 4, 5 
(descr., ill. adult and embryo, skel.; American occidentalis considered ident. with European nobilianus); 
Radcliffe, Bull. U. S. Bur. Fish., 24^ 1916: 270 (early N. Carolina report probably was Narcine); Linton, 
Proc. U. S. nat. Mus., 64 (21), 1924: 24, 25, 64, 84 (season, cestode parasites, Woods Hole); Bigelow 
and Welsh, Bull. U. S. Bur. Fish., 40 (j), 1925: 68 (genl.. Gulf of Maine); Breder, Field Bk. Mar. Fish. 
Atlant. Coast, 1929: 33 (genl.); Gowanloch, Bull. La. Conserv. Dep., 1932: 86 (electric shocks, size, 
listed Gulf of Mexico); Bigelow and Schroeder, Canad. Atlant. Fauna, i2e, 1934: 35 (descr., ill. after 
Garman, range); Vladykov and McKenzie, Proc. N. S. Inst. Sci., ig, 1935: 52 (St. Margaret Bay, 
Nova Scotia); Bigelow and Schroeder, Bull. U. S. Bur. Fish., 48, 1936: 325 (season, size, Georges Bank, 
Provincetown, Mass.). 

Narcobatus nobilianus Fowler, Copeia, 31, 191 6: 41 (listed, Sandy Hook Bay, New York). 

Torpedo Ray, Wilson, Bull. U. S. nat. Mus., 58, 1932: 403 (copepod parasites, Woods Hole). 

Doubtful American References: 

Tetronarce occidentalis Poey, Repert. Fisico-Nat. Cuba, 2, 1868: 458; also An. Soc. esp. Hist. Nat., 5, 1876: 
402 (Havana specimen referred doubtfully to T. occidentalis'); Enumerat. Pise. Cubens., 1875-1876: 
206 (listed, Cuba, by ref. to Poey, 1868). 

Torpedo occidentalis Jordan and Gilbert, Proc. U. S. nat. Mus., 5, 1883: 583 (electric fish reported by fisher- 
men, Charleston, S. Carolina, but perhaps Narcine; see p. 121); True, List Vert. Anim. S. Carolina, in 
Handb. S. Carolina, 1883: 261 (listed, S. Carolina). 

Not Raja torpedo Linnaeus, Syst. Nat., j, 1758: 231 (E. Atlant.). 

Not Torpedo occidentalis Yarrow, Proc. Acad. nat. Sci. Philad. (1877), 1877: 216 (listed, Fort Macon, N. Ca- 
rolina).!"' 

Genus Narcine Henle 1834 

Narcine Henle, Ueber Narcine, 1834: 2; type species, Torpedo brasiliensis Olfers 1831. Brazil. 

Generic Synonyms: 

Raja (in part) Bloch and Schneider, Syst. IchthyoL, 1801: 359; includes Raja timlei Bloch and Schneider, 
Tranquebar. 

Narcobatus (in part) Blainville, Bull. Soc. philom. Paris, 18 16: 121; Narcobatus maculatus Blainville included, 
doubtless equals Raja maculata Shaw (Gen. ZooL, 5 [2J, 1804: 316). Indian seas, based on the 
Temeree of Russell (Fish. Coromandel, j, 1803: i; Coromandel Coast). 

Torpedo van Hasselt, Bull. Sci. Nat. Ferussac, 2, 1824: 90; for Torpedo timlei van Hasselt, equals Raja timlei 
Bloch and Schneider 1801. 

Syrraxis Jourdan, in Bonaparte, Icon. Faun. Ital., J, Pesci, 1841 : 6th p. of descr. of Torpedo narke (no pagina- 
tion); type species, Narcine indica Henle 1834. 

Cyclonarce Gill, Ann. N. Y. Lye, 7, 1862: 387; type species, Narcine timlei Henle 1834, equals Raja timlei 
Bloch and Schneider 1 801. 

Gonionarce Gill, Ann. N. Y. Lye, 7, 1862: 387; type species, Narcine indica Henle 1834, equals Raja macu- 
lata Shaw 1804. Indian seas. 

Narcina Jordan and Seale, Proc. Davenport Acad. Sci., 10, 1905: pi. i; evident misprint for Narcine. 

Narcinops Whitley, Fish. Aust., I, 1940: 164; type species, Narcine tasmaniensis Richardson 1840. Tasmania. i"* 

Not Torpedo Houttuyn, Nat. Hist. Dieren, Planten en Mineral., 6, 1764: 453. 

107. It has been pointed out already by Radcliffe (Bull. U. S. Bur. Fish., J.^, 1916: 271) that the fish reported to Yarrow 
by fishermen was "not a ray but the Electric Stargazer {Astroscopus y-graecum)." 

108. Narcinops was separated from Narcine as differing from the type species of the latter [Torpedo brasiliensis Olfers 



I o8 Memoir Sears Foundation for Marine Research 

Generic Characters. Disc ranging from a little broader than long to a little longer 
than broad, its anterior margin broadly rounded; outer margin from continuously 
rounded to weakly concave a little posterior to level of spiracles; posterior contours 
of pectorals either curving inward and forward to junction with trunk or merging insen- 
sibly rearward with sides of latter. Snout in front of eyes rigid to the touch nearly to 
anterior margin. Tail with a ribband-like membranous fold along each side, its length 
from tips of pelvics more than six times as great as breadth of mouth, its length from 
center of cloaca greater than distance from center of cloaca to mouth. Margins of 
pectorals without papillae, spinous or otherwise. Origin of first dorsal over or slightly 
posterior to rear ends of bases of pelvics. Pelvics with outer corners ranging from 
subangular to broadly rounded, their distal margins from weakly convex to more or 
less deeply concave,"' their inner posterior margins either united with sides of tail to 
their tips or free from sides of tail for a shorter or longer distance in both sexes ;ii'' males 
with a considerable area of loose flabby skin in the axil of each pelvic fin. Eye fully 
developed, as large as spiracle in some species, smaller in others. Spiracles either close 
behind eyes or separated from them by a moderate interspace, their length at least Vs as 
great as distance between spiracles, their margins either smooth or more or less cor- 
rugated or with papillae. Nuchal region with a pair of large mucous pores, close together 
side by side. Nostrils slightly oblique, apparently minute on cursory appearance because 
they are roofed inwardly by the joint nasal curtain, but actually they are about as long 
as the breadth of the isthmus between them, or longer. Nasal curtain much broader 
than long, extending to mouth; posterior margin of nostril also somewhat expanded 
outwardly as a low flap, directed toward the mouth. Mouth narrow, transverse, ap- 
proximately straight, and protractile as a short tube, flattened dorsoventrally.^^^ Teeth 
with one sharp cusp. Tooth bands falling considerably short of corners of mouth; 
largely exposed when mouth is closed in some species but entirely enclosed within 
the closed mouth in at least one (p. 123). Integument bearing teeth only loosely 
attached to jaws. Upper and lower jaw cartilages broad, their lateral articulations ex- 
tensive in area and very firm; a special labial cartilage of two subtriangular elements 
on either side of mouth; proximal end of upper element attached to upper jaw car- 
tilage, that of lower element to lower jaw cartilage, about midway between symphysis 

1 831) in "form of body, margin of nasal valves, in having a wider skull and different cartilages" (Whitley, Fish. 
Aust., J, 1940: 164). But the first two differences are not greater than is to be expected between species of a genus 
with a geographical range as wide as that of Narcine. Nor does a somewhat wider skull seem important. Also, the 
absence of small intermediate cartilaginous elements between rostral and antorbital cartilages and of a transverse 
foramen in the anterior part of the rostral cartilage in A^. tasmaniensis, compared with their presence in N. brasiliensis, 
seems more appropriate as a specific than as a generic character. 

109. The original illustration of N. tasmaniensis (Richardson, Trans, zool. Soc. Lond., J, 1849: pi. 2, fig. 2) shows the 
distal margins of the pelvics as so deeply concave that the posterior part of each fin appears as a distinct lobe. But 
seemingly this was an exaggeration, for they are represented as only weakly concave in a recent picture of this 
same species (Whitley, Fish. Aust., i, 1940: 165, fig. 186). 

no. The pelvics in some are adnata to their tips (e. g., Narcine timlei, p. 112), in others (e. g., TV. indica) the inner pelvic 
margins are free for only a short distance, in others (e. g., N. brasiliensis) the pelvics are free for a somewhat greater 
distance. 

III. In a specimen of N. timlei, about 34 cm long, the mouth protruded to a distance of about 35 mm (Annandale, 
Mem. Indian Mus., 2, 1909: 44). 



Fishes of the Western North Atlantic 109 

and outer articulation; the two elements of each labial cartilage flexibly joined. The 
presence of these cartilages, combined with the firmness with which the two jaws are 
articulated, limits the gape of the mouth. ^'^ Rostral cartilage single but bifid at tip; 
trough- or shovel-shaped, with a deeper or shallower median depression bounded on 
either side by a more or less sharply defined ridge; rigid next to cranium but soft and 
flexible toward its tip; its proximal portion with or without a transverse foramen; 
anterior margins of antorbital cartilages more or less dissected, with or without one 
or two isolated cartilages on either side between each of them and rostral cartilage. "=* 
Each electric organ with an average of 146—428 columns in the species in which they 
have been counted."* Characters otherwise those of the family. 

Sixe. The maximum length definitely reported for Narcine appears to be about 
30 inches (762 mm),"^ the second longest 18 inches,^i^ sizes far below the length 
reached by species of the genus Torpedo (p. 85). 

Developmental Stages. The inner walls of the uterus are thickly set with villi (up 
to 200 per sq. in. of uterine surface), as is generally the case in ovovlviparous batoids. 
These villi are spatulate in form and are provided with a dense net of capillaries. Ap- 
parently they secrete the yellowish milky solution that fills the uterus and probably 
nourishes the embryo. In embryos at a stage when the external gill filaments still per- 
sist, the anterior limits of the pectorals may still be marked by deep notches in the 
margins of the disc, or the notches may have disappeared already."' In either case, 
the pectoral notches have disappeared before birth in all species of the genus, so far 
as is known. 

Habits. The various species of Narcine are most commonly encountered close to 
land, in water so shoal that they can be taken in seines hauled from shore, or they can 
be seen lying on bottom. And 60 fathoms appears to be the maximum depth recorded 
for the genus. They have been described as sluggish in habit. The fact that the pro- 
tractile mouth cannot be gaped widely would suggest that they subsist on less active 
and smaller prey than members of the genus Torpedo (p. 102). In fact, the stomach 
contents of the few specimens examined have consisted solely of polychaete worms, 

112. These cartilages with their attached muscles were described and accurately pictured for Narcine brasilietisis more 
than a century ago by Henle (Ueber Narcine, 1834: 12, pi. 4, fig. 2); they have been illustrated more recently for 
Narke japonica by Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: pi. 67, fig. 3). 

113. For a general account of the head skeleton with illustrations, see Holmgren (Acta Zool. Stockh., 22, 1941 : 57-59, 
figs. 50-55)- 

114. The number of columns in the Indo-Pacific species [Narcine tasmaniensis, N. lingula, N.timlei, and N.indica) 
average from 146-27S per organ, those in adult .V. brasiliensis (Fritsch, Elektr. Fische, 2, 1890: 98; Cox and Breder, 
Zoologica N. Y., 2S, 1943: 46) from as few as 380 to as many as 428. 

115. Breder (Bull. Bingham oceanogr. Coll., 2 (i), 1928: 5) for a specimen of Narcine entemedor; jaws only. 

116. Reported for N. brasiliensis (p. 118); also for N. tasmaniensis by Whitley (Fish. Aust., j, 1940: 164). A two-foot 
specimen from the west coast of Mexico, reported by Kumada and Hyama (Marine Fish Pac. Coast Mexico, 1937: 
21) as a Narcine, may not have belonged to that genus, for they describe its snout as "much angular." 

117. See Prashad (Rec. Indian Mus., ig, 1920: 104, pi. 7, figs. 4-9) for an account of the uterine wall in Narcine indica 
Henle 1834 and for illustrations of embryos with the anterior parts of the pectorals still separate from the sides of 
the head. For illustrations of an embryo of N. limlei (which we have examined) at about the same stage in devel- 
opment but with the pectorals fully united with the sides of the head, see Garman (Mem. Harv. Mus. comp. Zool., 
36, 1913; pi. 61, fig. 6). 



1 1 o Memoir Sears Foundation for Marine Research 

ascldians, amphipods, small shrimps, and other Crustacea. i" Their feeding habits have 
not been observed, but their known diet and the structure of their jaws and lips suggest 
that they suck into their protruded mouths whatever soft-bodied animals they may 
encounter on bottom (or close to it), rather than that they pursue more actively swim- 
ming prey. 

The electric shocks of Narcine are much weaker than those of Torpedo, the highest 
recorded voltage being about 37, and apparently a more active stimulus is required to 
induce the former to discharge."^ 

Relation to Man. These little Electric Rays, though locally common, have never 
been of any commercial importance. 

Range. Continental waters of the western Atlantic from N. Carolina and Texas 
to southern Argentina; Pacific Coast of America from the Gulf of California to Pa- 
nama; Indo-China, China and Japan; Malaysia and East Indies; southeastern and 
southern Australia (Queensland and Victoria) and Tasmania; India. There is no re- 
liable report of the presence of Narcine anywhere in the eastern side of the Atlantic, 
in the Red Sea, or along the African Coast of the Indian Ocean,!-" unless the limits 
of the genus be expanded to include the species that are classed here (p. 89) as 
Heteronarce.^'^^ 

Species. Examination of representatives of five of the species concerned ^-^ from 
widely separated localities leads to the conclusion that the most useful characters for 
purposes of specific identification are: (a) distance between eyes and spiracles; (b) shape 
of caudal; (c) margins of spiracles smooth or corrugated; (d) shape of joint anterior 
nasal curtain; (e) shape of pelvics; and (f) color in the case of the more conspicuously 
marked species. The position of the origin of the first dorsal relative to the axils of 
the pelvics is useful only if the question of sex be taken into account, since males and 
females differ in this respect. But final diagnoses must await first-hand comparisons 
between adequate series of the Malaysian species, including a final decision as to 
whether the Raja maculata of Shaw i 804^^^ was an earlier name for Narcine indica Henle 
1834 or a later name for N. timlei Bloch and Schneider 1801. 

118. Stomach contents have been recorded by Beebe and Tee-Van (Zoologica N.Y., 26, 1941: zi^j) iov N. enlemedor, 
and by Richardson (Trans, zool. Soc. Lond., 3, 1844: 182) for N. iasmaniensis. For stomach contents oi N. brasili- 
ensis, see p. 119. 

119. This reluctance, reported by Annandale for the Indian' A^. timlei (Mem. Indian Mus., 2, 1909; 45), has been empha- 
sized recently by Cox and Breder (Zoologica N. Y., 28, 1943: 47) for N. brasiliensis. 

120. Giinther's (Cat. Fish. Brit. Mus., 8, 1870: 453) report oi Narcine brasiliensis from the Cape of Good Hope probably 
was an error, for Narcine has not been found during recent surveys of the fish fauna of South African waters (von 
Bonde and Swart, Fish. Mar. biol. Surv. S. Afr., Rep. 3 [1922], 1924, Spec. Rep. i ; Barnard, Ann. S. Afr. Mus., 
21 [i], 1925: 91). 

121. The type species of Heteronarce, H. garmani Regan (Ann. Mag. nat. hist., [9] 7, 1921 : 414), is reported so far only 
from the coast of Natal. Narcine mollis Lloyd (Rec. Indian Mus., i, 1907: 5), also referred to Heteronarce by Re- 
gan, is known only from the Gulf of Aden. 

122. Narcine brasiliensis (p. 112); A^. indica from Penang; N.Ungida from Penang; N. timlei from China; N. mermiculata, 
and the form previously called N. entemedor which probably is not separable from N. brasiliensis, from the west 
coast of Central America. 

123. Gen. Zool., 5(2), 1804: 316. 



Fishes of the Western North Atlantic 



III 



Tentative Key to Species >^* 

la. Spiracles separated from eyes by a distance as long as horizontal diameter of orbit. 

tasmaniensis Richardson 1840. 
Tasmania (type locality), Victoria, and 
New South Wales. 
lb. Spiracles close behind eyes. 

2 a. Caudal fin fan-shaped, with more or less definite lower and upper corners, 
its posterior margin approximately vertical, its lower margin only weakly 
convex at most. 

3 a. Disc conspicuously marked above with white vermiculations on a dark 
ground. vermiculatus Breder 1928. 

Pacific Coast of Mexico. 
3 b. Disc either plain-colored above or with dark markings on paler ground. 
4a. Outer (posterior) margins of pelvics weakly but continously convex 
or straight. 

5 a. Margins of spiracles conspicuously corrugate or tuberculate. 
brasiliensis (Olfers) 1831, 

including entemedor Jordan and Starks 1895, p. 120. 
5 b. Margins of spiracles only faintly tuberculate, if at all so. 

Hngula Richardson 1846. 
Indo-China, China, Japan. 
4 b. Outer (posterior) margins of pelvics conspicuously concave. 

schmitti Hildebrand 1948.1^^ 
Gulf of California. 
2 b. Caudal fin ovoid, its posterior margin convex and strongly oblique, its lower 
margin continuously rounded without definite lower corner. 
6a. Posterior parts of pectorals slightly overlapping anterior parts of pelvics; 
posterior outlines of pectorals definitely though slightly recurved; inner 
margins of pelvics separate from sides of tail for a short distance anterior 
to rear tips. indica Henle 1834.^^^ 

India, Malaysia, East Indies. 
6b. Posterior parts of pectorals not overlapping anterior parts of pelvics; 
posterior outlines of pectorals not recurved at junction with sides of 
trunk; inner margins of pelvics united with sides of tail to their tips. 

124. Narcine mollis Lloyd (Rec. Indian Mus., i, 1907 : 8) falls in Heteronarce, according to the generic definitions adopted 
here. 

125. Smithson. misc. Coll., 7/0(9), '94^= '• 

126. Fowler (Bull. U. S. nat. Mus., 100 [13], 1941: 333) identifies N. indica Henle 1834 with Raja maculata Shaw 1S04 
(Gen. Zool., 5 [2], 316), which was based on the Electric Ray pictured by Russell (Fish. Coromandel, i, 1803: 
pi. i) under the vernacular name "Temeree." But Russell's account and illustration give so little detail that we 
think it premature to accept this identification until some student is able to re-examine the whole question of the 
inter-relationships of the Indian species of Narcine, especially since Rudolphi (Grundriss Physiol., i, 1S21 : 199) 
long ago suggested that the Temeree might equal Narcine timlei (Bloch and Schneider) i8oi, not N. indica Henle 
1834. 



112 Memoir Sears Foundation for Marine Research 

7a. Upper surface of disc conspicuously marked with dark spots; nasal 
curtain projecting but little, if at all, in the midline. 

tinilei (Bloch and Schneider) 1801. 

India, Malaysia, East Indies, Indo-China to 

China and Japan. i^' 

7b. Upper surface plain-colored; nasal curtain with a distinct projection 

in the midline. brunnea Annandale 1909. 

India and Ceylon. ^^^ 



Narcine brasiliensis (Olfers) 18311^^ 

Lesser Electric Ray 

Figures 24, 25 

Study Material. Thirty-five specimens of both sexes, from iio mm (one embryo 
with scar, and one newborn) to 450 mm in total length, from Rio de Janeiro and Rio 
Parahyba, Brazil; from St. Vincent, West Indies; from the Gulf of Campeche; and 
from New Smyrna Beach and the St. Johns River, Florida (including type of var. coral- 
Una Garman 188 1), in Harvard Museum of Comparative Zoology; 19 specimens, 
including eight embryos 69-81 mm long, from Palmetto Key, Florida, in American 
Museum of Natural History; 14 embryos, each about 80 mm long, from Long Key, 
Florida, in U. S. National Museum;i^" and one from off Cape Lookout, North Carolina, 
from U. S. Fish and Wildlife Service; also three specimens (one male, two females) 
of the Pacific form that has been known as Narcine entemedor Jordan and Starks 1895, 
375 to 405 mm long, one from Magdalena Bay, and two from Inez Bay, Gulf of Cali- 
fornia, in the American Museum of Natural History. 

Distinctive Characters. Torpedo nobiliana, Diplobatis pictus, and Benthobatis marcida 
are the only other western North Atlantic Rays with which Narcine brasiliensis might 
be confused. It is easily separable from Torpedo by the much more strongly convex 

127. Prashad (Rec. Indian Mus., ig, 1920: 99), following Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 300), 
concluded that timlei "is not an Indian form, being confined to the East Indies and Japan." However, the type 
locality of the species is Tranquebar, southeastern India ("Habitat Tranquebariam . ." Bloch and Schneider, 
Syst. Ichthyol., 1801: 359). 

128. Day (Fish. India, 1875-1878: 733) reported some specimens of N. timlei as spotted and others as plain -colored, 
and it was to the latter that Annandale (Mem. Indian Mus., 2, 1909: 45) gave the name A'', brunnea. However, 
the color patterns of various species of Narcine are so variable (p. 117) that it would not be astonishing should the 
extremes in this respect, represented by .V. timlei and N. brunnea, prove to intergrade. It is also questionable whether 
or not the degree to which the free edge of the nasal curtain projects in the midline is a dependable specific character. 
Annandale (Mem. Indian Mus., 2, 1909: 44, pi- 3 A, figs. 1,2) stated, as an additional specific character of N. brun- 
nea, that each of the ridges on the floor and roof of its mouth behind the teeth bears a pair of vertical processes. 
But the difference in this respect, as illustrated between timlei and brunnea, is not clear cut. Narcine firma Garman 
(Mem. Harv. Mus. comp. Zool., j6, 191 3 : 30), described from a single Ceylon specimen that seems to be in existence 
no longer, has already been relegated by Fowler to the synonymy of A', brunnea, and justly so in our opinion. 

129. The earhest account of this species, with unmistakable illustrations, was by Gronow (Zoophyl., I [37], No. 158, 
1763 : pi. 9, fig. 3) as Kaja lae'vis. But Gronow's names have been declared unavailable by the International Com- 
mission on Zoological Nomenclature (Opin. 89, Smithson. misc. Coll., 73 [3], 1925: 27). 

130. The U. S. National Museum also has specimens, which we have seen, from: Jamaica; Colon; Cozumel; Texas; 
Pensacola, Key West and Cape Canaveral, Florida; and Cape Lookout, North Carolina. 



Fishes of the Western North Atlantic 



113 




Figure 24. Narcine brasiliensis, illustrating variations in color pattern. A Male, 396 mm long, from Rio Parahyba, 
Brazil (Harv. Mus. Comp. Zool., No. 308). B Female, 238 mm long, from St. Vincent Island, West Indies 
(Harv. Mus. Comp. Zool., No. 752). C Male, 197 mm long, from Key West, Florida (Harv. Mus. Comp. Zool., 
No. 44). D Male, 152 mm long, from New Smyrna Beach, Florida (Harv. Mus. Comp. Zool., No. 36398). 
E Male, no mm long, from New Smyrna Beach, Florida (Harv. Mus. Comp. Zool., No. 36398). F Male, 
114 mm long, from Desterio, Brazil (Harv. Mus. Comp. Zool., No. 232). 



114 Memoir Sears Foundation for Marine Research 

anterior outline of its disc, by the close proximity of its spiracles to its eyes, by its 
much narrower mouth with labial cartilages at the corners (this may be felt readily 
in the larger sizes), and by its much more rigid snout. From the local species of Ben- 
thobatis it is distinguishable by the tuberculate margins of its spiracles, by the shape of 
its caudal (cf. Fig. 25 A with 26B), by its larger and fully formed eyes, and by the 
fact that the inner margins of its pelvics are free from the sides of the tail for some 
distance anterior to their tips. 

Description. Proportional dimensions in per cent of total length. Female, 239 mm 
long, from New Smyrna Beach, Florida (Harv. Mus. Comp. Zool., No. 36398). Male, 
274 m long, from Rio de Janeiro (Harv. Mus. Comp. Zool., No. 643). 

Disc: extreme breadth 54.8, 51.3; length 51.0, 47.8. 

Snout length: in front of orbits 13.0, 11.3; in front of mouth 15.0, 13.0. 

Orbits: horizontal diameter 4.5, 4.0; distance between 5.9, 5.8. 

Spiracles: length 2.5, 2.2; distance between 7.7, 8.0. 

Mouth: breadth 4.5, 5.1. 

Exposed nostrils: distance between inner ends 6.1, 6.6. 

Gill openings: lengths, ist 1.5, 2.2; 3rd 2.1, 2.4, 5th 1.5, 1.8; distance between 
inner ends, ist 13.2, 13.0; 5th 7.1, 7.5. 

First dorsal fin: vertical height 9.2, 9.9; length of base 7.5, 7.3. 

Second dorsal fin: vertical height 8.6, 9.9; length of base 7.5, 8.4. 

Pelvics: anterior margin 12.5, 15.0. 

Distance: from tip of snout to center of cloaca 52.3, 50.0; from center of cloaca 
to 1st dorsal 26.5, 24.9; to tip of tail 47.7, 50.0. 

Interspace between: ist and 2nd dorsals 2.7, 4.7; 2nd dorsal and caudal 3.5, 4.7. 

Disc subcircular, its anterior contour varying from evenly arcuate to moderately 
ovoid, its breadth usually a little greater than its length ^^^ and 49—55 "/o as great as total 
length; posterior margins of pectorals moderately recurved in broadly rounded contour. 
Tail from center of cloaca somewhat shorter to slightly longer (84—101 "/o) than dis- 
tance from center of cloaca to tip of snout; rounded above and moderately flattened 
below, each side with a narrow longitudinal cutaneous fold extending from about 
opposite or a little anterior to midpoint of base of first dorsal to origin of caudal fin. 

Length of snout in front of orbits about 1.6—2.2 times as great as distance between 
eyes, its length in front of mouth about twice as great as distance between outer ends 
of nostrils; the two lateral ridges of the trough-shaped rostral cartilage firm to the touch 
nearly to the anterior margin of the head. Orbits moderately prominent (this may not 
be the case in preserved specimens). Eyes fully developed and about ^/g as long as 
spiracles, oval, their horizontal diameter about 42—78 °/o as long as distance between 
orbits. Spiracles close behind eyes, transverse or slightly oblique, squarish in shape 
when expanded, the margin ringed with a single series of 12 to 20 low rounded papillae 

131. Among specimens in our Study Material, the ratios of breadth of disc to its length are from 1.01-1.08; in one 
Brazilian specimen, probably of this species, this ratio was 0.99. See discussion of Narcine brachypkura Ribeiro 
1923 (p. 123). 




Figure 25. Narcine brastliensis. A Side view of tail of specimen illustrated in Fig. 24A. B Ventral view of pelvics 
of same. C Ventral view of pelvics of female, 450 mm long, from Rio de Janeiro, Brazil (Harv. Mus. Comp. 
ZooL, No. 643). D Side view of tail of male, 270 mm long, from Rio de Janeiro, Brazil (Harv. Mus. Comp. 
ZooL, No. 643). .ff Nostrils and mouth of female, 450 mm long (Harv. Mus. Comp. ZooL, No. 643), about 1.5 X. 
F Lower surface of disc of female, 258 mm long, from New Smyrna Beach, Florida (Harv. Mus. Comp. ZooL, 
No. 36398), with skin dissected away from left-hand side to show electric organ somewhat diagramatically. 
G Eye and spiracle of another female of about same size and from same locality, about 1.2 X. // Upper teeth of 
female, 450 mm long, from Rio de Janeiro, Brazil (Harv. Mus. Comp. ZooL, No. 643), about i 5 X . y Upper 
teeth of male, 396 mm long, from Rio Parah}ba, Brazil (Harv. Mus. Comp. ZooL, No. 308), about IJX- 



1 1 6 Memoir Sears Foundation for Marine Research 

or knobs that vary in distinctness from specimen to specimen. Gill openings noticeably 
small; first to fourth a little less than half as long as breadth of closed mouth, fifth about 
75 "/o as long as first to fourth; distance between inner ends of gill openings of fifth 
pair a little smaller than distance between spiracles. Nostrils approximately transverse, 
their inner ends reaching the deep furrow that surrounds mouth, thus approaching 
closely to latter though separate from it; all of nostril, except for a small circular aper- 
ture at outer end, rooted over by expansion of anterior margin; joint nasal curtain sub- 
rectangular with rounded corners, about three times as broad as long, extending rear- 
ward to slightly overlap anterior tooth band; free posterior margin of nasal curtain 
smooth In large specimens, weakly zigzag in small, or perhaps even irregularly and 
narrowly fringed, with a low median projection in some preserved specimens; outer 
half of posterior margin of nostril also expanded as a short rounded flap, directed 
inward and rearward toward the mouth, its free edge either smooth or irregularly 
zigzag.1^2 Mouth transverse, straight, its breadth when closed 77—92 "/q as great as 
distance between orbits and about half as great as distance from anterior margin of 
upper tooth band to tip of snout. Lips conspicuously fleshy at corners, with many short 
wrinkles. Tooth bands occupying about 60 % of breadth of mouth when closed, the 
extreme length from front to rear of exposed portion of upper band about half as great 
as its transverse width from one end to the other; length of lower band from front to 
rear only about Va as great as Its transverse width. 

Teeth Increasing in number of series with growth from about J^ in specimen 
239 mm long to about 33 in adults; with single conical cusp that is rather sharp; closely 
crowded; posterior teeth considerably longer than anterior ones, a difi:erence probably 
resulting from wear; seven to eight rows in function simultaneously on both upper 
and lower tooth bands. 

First and second dorsals approximately equal in size and similar in shape on any 
given individual; outlines characteristic of young (Fig. 25D) persisting to maturity 
with little If any alteration in some though changing with growth in others to more 
narrowly ovoid, with loss of basal recurvature (Fig. 2 5 A); anterior margins a little 
less than twice as long as bases. Origin of first dorsal posterior to posterior ends of 
bases of pelvlcs by a distance a little more than half as great as length of its own base. 
Interspace between first and second dorsals about 36—74 % as long as base of first 
dorsal. Distance between second dorsal and caudal 43—65 % as long as base of second 
dorsal. Caudal subtriangular, nearly equilateral; its posterior margin approximately 
vertical, moderately and continuously convex in small specimens but becoming less 
so with growth until nearly straight in adults ; upper posterior corner abrupt, the lower 
corner somewhat more rounded; axis little raised; depth of caudal below tip of axis 
about equal to height above latter; Its upper margin about as long as distance from 
origin of first dorsal to origin of second dorsal, or a little longer. Pelvics originating 
below axils of pectorals, which slightly overlap them; their anterior margins In both 
sexes about 50—55 "/o as long as distance from origin of pelvics to rear tip and varying 

132. We have seen one specimen in which it is smooth on one side, zigzag on the other. 



Fishes of the Western Nortli Atlantic 117 

from nearly straight when fins are spread to weakly and continuously convex; rear tips 
reaching about as far as origin of first dorsal in females and young but as far as mid- 
point of first dorsal base in adult males; inner margins of pelvics anterior to rear tips 
free from sides of tail for a distance about as long as base of first dorsal in females, but 
about 1.3 times that long in adult males. 

Proximal edge of upper element of each labial cartilage with a distinct notch; 
one or two small cartilages intervening between rostral cartilage and antorbital cartilage 
on each side, or none."^ 

Color. Upper surface dark brown, grayish brown, or leaden to orange or reddish; 
uniform in some cases but more often irregularly clouded with vaguely defined darker 
bars or blotches. Most specimens have one band extending across the head anterior 
to the eyes, but others have another across the posterior part of the head, one or two 
across the posterior part of the disc extending onto pelvics and tail, and one on the tail 
extending onto the dorsals; or they may have any or all of these in combination. Late 
embryos and newborn specimens usually have a varying number of dark rings, ovals, or 
loops, or darker blotches with pale centers. These markings persist undimmed to 
maturity in some cases, but more often they become disassociated into separate dots 
or irregular larger spots that recall the original pattern only faintly at maturity."* In 
fact, they may finally fade out altogether with growth. "^ Lower surface white, either 
pure or of a yellowish or greenish cast, with the posterior parts of pectorals and outer 
margins of pelvics dusky-edged. Some specimens show irregular black blotches ex- 
tending inward and forward from the region of the axils of the pectorals, and some 
have scattered sooty spots on the posterior parts of the latter or elsewhere on the lower 
side of the disc. 

Remarks. More than a century ago three color varieties were defined but not 
named."* The name Narcine coraUina has been proposed, first as varietal,"' next as a 
full species,"* and subsequently as a subspecies,"^ for specimens from the Florida Keys 
in which the ground tint of the upper surface was orange or red. Similarly, West Indian 
specimens with the dark blotches replaced by loops of small dark spots have been named 
Narcine punctata^'^^ but subsequently Narcine hrasiliensis var. bancrofti.^'^^ Still others from 
Key West, Florida, with dark cloudings but without a superimposed pattern have been 
made the basis of a separate species, Narcine umhrosa Jordan 1884.^*^ However, these 
differences in hue and pattern, while striking enough between the extremes (Fig. 24), 

133. Two such intervening cartilages are pictured for Narcine brasiliensis by Henle (Ueber Narcine, 1S34: pi. 4, figs. 
2, 3). However, only one was found on one side and none on the other in a specimen that we dissected. 

134. For photographs of a female that falls in this category, and her 15 embryos, see Bean and Weed (Proc. U. S. nat. 
Mus., 40, 191 1 : pis. 10, 11). 

135. All of these variations are to be seen among specimens that we have observed from the west and east coasts of Florida. 
Also Brazilian specimens may have either the clouded variety of coloration (Fig. 24) or may retain the juvenile pat- 
tern more or less distinctly (Ribeiro, An. Mus. nac. Rio de J., 14, 1907: pi. 17). 

136. MuUer and Henle, Plagiost., 1841: 129. 137. Garman, Bull. Mus. comp. Zool. Harv., 8, 1881: 234. 
138. Jordan, Proc. U.S. nat. Mus., 7, 1884: 105. 139. Radcliffe, Bull. U. S. Bur. Fish., 34, 1916: 270. 

140. Garman, Bull. Mus. comp. Zool. Harv., 11, 1881: 233. 

141. Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 298. 

142. Proc. U. S. nat. Mus., 7, 1884: 105. 



1 1 8 Memoir Sears Foundation for Marine Research 

intergrade to such a degree that they cannot even be regarded as appropriate bases for 
separation to subspecies. Indeed, it seems more likely that the variation in color is 
associated with small local populations, perhaps only temporarily, rather than with 
any continuing correlation between coloration and geographical distribution. 

Relation to Extralimital Species. It has been thought that Narcine entemedor Jordan 
and Starks 1895 of the Pacific Coast of Central America differs from N. brasiliensis 
of the Atlantic in the dimensions and shape of its pelvic fins"' and in having eyes much 
smaller than the spiracles. However, in specimens from the Pacific Coast of Central 
America that we have examined (see Study Material, p. 112) the eyes are actually 
about as large as the spiracles. Nor does there appear to be any consistent difference 
between Atlantic and Pacific specimens in shape of pelvic fins or in proportional dimen- 
sions in general. While none seen so far from the Pacific had as definite a color pat- 
tern as is often shown by iV. brasiliensis, this is a highly variable character in the latter, 
as already noted (p. 1 1 7). 

Size. The average length at birth is about i 10—120 mm. Males mature at a length 
of about 225—250 mm (about 9—10 in.), to judge from the lengths of their claspers 
(see Study Material, p. 112) and from the sizes of male specimens taken in West 
Florida waters in company with females containing embryos. Gravid females reported 
thus far have ranged from 271— 320 mm in length. The maximum recorded length 
(sex not stated) is about 450 mm (17V4 in.), though the species is credited with reaching 
two feet."^ Specimens from the west coast of Florida weighed 230— 270 g (about 8 — 
10 oz.) at lengths of 240-250 mm, 280-300 g (about lo-ii oz.) at 270-280 mm, 
and 450— 650 g (about 16—23 oz.) at 320— 330 mm. "^ 

Developmental Stages. Embryos, when nearly ready for birth, have lost all trace 
of the marginal notches that mark the anterior limits of the pectorals in early stages, 
and in general form they resemble the adult with the interesting exception that the 
caudal fins are oval in form, not fan-shaped as are those of their parents. But their color 
pattern is even more conspicuous than that of the most strongly marked adults. 

Four to 1 5 embryos have been reported in gravid females, and there appears to 
be a tendency for the majority of the young of a given female to be of one sex."'' 

Habits. This little Electric Ray is found for the most part in water so shallow 
that it is often taken in beach seines or in trammel nets; barefoot fishermen have been 
known to tread upon them, and they have been observed buried in the sand with only 
the eyes and parts of the head exposed."' But they sometimes move out into somewhat 
deeper water, for there is a record of one taken in Argentine waters at a depth of 10 
fathoms, another trawled off Corpus Christi, Texas in 14 fathoms, of several taken 

143. Meek and Hildebrand, Field Mus. Publ. Zool., 15 {i), 1923: 74- 

144. Jordan and Evermann, Bull. U.S. nat. Mus., 47(1), 1896: 78. 

145. Cox and Breder, Zoologica N. Y., 2S, 1943: 50. 

146. For earlier illustrations and accounts of embryos, see Bean and Weed (Proc. U. S. nat. Mus., 40, 191 1 : 231-232) 
and Cox and Breder (Zoologica N. Y.,28, 1943 : 46); for the relative numbers of the two sexes, see Breder and Sprin- 
ger (Zoologica N. Y., 25, 1940: 451). 

147. Coles, Bull. Amer. Mus. nat. Hist., 2S, 1910: 337. 



Fishes of the Western North Atlantic 119 

off Venezuela in 20 fathoms,''"' and one off Cape Lookout, North Carolina in 20 fathoms. 
There is no reason to suppose, however, that their depth range extends much below this. 

They are at home in the highest equatorial temperatures, but they can survive 
considerable chilling, for some of the Texas captures were made in water as cold as 
15.4° C (59-7° F). The few definite determinations of salinity for them, again off 
Texas, have ranged from 30.6—36.5 "/oo ^"d the fact that they have not been found 
"in Texas inside waters, except occasionally in the Laguna Madre, which is a peculiar 
environment with very high salinity at times,"i** is an additional reason for questioning 
the correctness of the older reports of them in fresh water. 1^" 

Of its diet we know only that the stomach of one specimen, taken near Cape 
Lookout, North Carolina, was full of red annelid worms, '^' and that of another from 
New Smyrna Beach, Florida, opened by us, contained a single good-sized annelid. 

Reports of gravid females have been confined to June and July, whether for 
Florida or for North Carolina; this suggests that the young are born during the warm 
months in latitudes, where there is a considerable difference in temperature between 
winter and summer; in tropical latitudes, however, they may be produced throughout 
the year. 

Electric Capabilities. The number of columns In each of the electric organs, which 
make up about Ye of ^^he weight of the fish, ranges from as high as 428 (average of 
three specimens) '^^ to as low as 382 for adults and 286 for embryos ;i*^ the number of 
electric plates (or electro-plaxes) averages from 179—288 per column in the outer part 
of the organ (where the columns are shortest) to 288—482 in the inner part (where 
longest). 

An observer, familiar with A^. brasiliensis in North Carolina waters, writes that 
he has been knocked down by their shocks, as have others who have chanced to tread 
on them while wading barefoot in shallow water. It seems, however, that it is the sud- 
denness rather than the intensity that is responsible in such cases, for the peak voltage 
for 12 specimens of both sexes in good condition was only from 14—37 volts."* To 
induce a discharge, it has been found necessary "to prod, lightly pinch, or otherwise 
annoy them .... and it finally developed that a new stimulus was more effective than 
any one kind often repeated.""-' From this it seems doubtful whether discharges of 
this particular Electric Ray have any great efficiency as a protective device. This doubt 
is strengthened by the further observation that specimens kept in a live car did not 
protect themselves from attacks upon their eyes by small pinfish {hagodon rhomboides), 

148. Delsman, Mem. Mus. Hist. nat. Belg., [2] Fasc. 21, 1941: 65. 

149. Gunter, Publ. Inst. mar. Sci. Texas, /, 1945: 22. 

150. In the U. S. National Museum there is a specimen labelled "St. Johns River, Florida" (see Study Material, p. 112), 
but there is no knowing how far upstream it was actually taken, i. e., whether in fresh water or in salt or brackish 
water near the mouth of the river. 

151. Gudger, J. Elisha Mitchell sci. Soc, 28, 1913: 160. 152. Fritsch, Elektr. Fische, 2, 1890: 98. 

153. Cox and Breder (Zoologica N. Y.,28, 1943 : 46) give the most complete account of the electric organs of this species. 

154. See Cox and Breder (Zoologica N. Y., 28, 1943: 50) for voltage, current, and power at peak of discharge with 
various resistances; recorded by oscillograph for two specimens. 

155. Cox and Breder, Zoologica N. Y., 25, 1943: 48. 



1 2 O Memoir Sears Foundation for Marine Research 

a species which "has a tendency to pick out the eyes of fish which lie prone on the bot- 
tom. "^^* Nor does the little that is known of its diet (p. 119) or of that of other mem- 
bers of its genus (p. 109) suggest that the electric organs are likely to be of as much 
service to them in feeding as they may be to other Electric Rays that prey on actively 
swimming fishes (p. 102). 

Numerical Abundance. No exact information is available as to the numbers of 
N. brasiliensis. But reports that it is taken frequently in trammel nets on the west coast 
of Florida, plus the fact that considerable numbers are contained in the collections 
of museums and that they have been reported with considerable frequency from one 
place or another, suggest that the population within its year-round range is consider- 
ably more abundant than is that of its larger relative. Torpedo nobiliana. It appears, 
however, that few take part in the summer migration northward, for the total numbers 
reported at Cape Lookout in years when watch was kept for them were small : only two 
in 1909, II plus about a dozen more by fishermen in 19 10, four in 191 1, and 16 in 
1912.1*' Also, it seems that few reach the opposite boundary of the species' range, 
for the number of recorded specimens is only four from Argentine waters. ^^^ 

Relation to Man. The Lesser Electric Ray is of no commercial importance; in fact, 
we have no evidence that it is brought into tropical fish markets. But a correspondent^^' 
who has tried them on the table and who has supplied the specimens from the east 
coast of Florida (listed, p. 112) informs us that they are excellent eating. 

Range. Inshore waters of the western Atlantic, from southern Brazil to Florida 
and Texas; south in small numbers to northern Argentina in about Lat. 39—40° S, 
and north to North Carolina. It is represented on the Pacific Coast of Central America, 
Gulf of California to Panama, by a form {Narcine entemedor Jordan and Starks 1895) 
so closely allied that it will probably prove indistinguishable when more fully studied 
(see discussion, p. i 18). N. brasiliensis has been reported also from the Cape of Good 
Hope, but probably not correctly.^^" 

Occurrence in the Western Atlantic. This Electric Ray has been reported from 
localities so widely distributed,"^ and it is so well represented in the larger museums 
of both America and Europe, that it is to be expected anywhere along the American 
littoral, provided the type of bottom and depth be suitable, from the coast of southern 
Brazil at about Lat. 28° S northward along the South American Coast, throughout 

156. Cox and Breder, Zoologica N. Y., 28, 1943: 47. 

157. Coles, Bull. Amer. Mus. nat. Hist., 28, 1910: 337; Gudger, Proc. bid. Soc. Wash., 26, 1913: 99. 

158. Lahille, An. Mus. nac. B.Aires, 34, 1928: 338. 159. Captain R.A.Howard, A. U. S. 

160. Reports of N. brasiliensis in South African waters appear to lead back to Dumeril's (Hist. Nat. Poiss., J, 1865: 
515) suggestion that the Torpedo ocellata of Quoy and Gaimard (Voy. 'Uranie', Zool., 1824: 199) might be identical 
with N. brasiliensis. But its large mouth (so described by Quoy and Gaimard) seems to locate it in the genus Torpedo. 
In any case, the governmental marine surveys of South African waters have not taken any Narcine (von Bonde and 
Swart, Fish. Mar. biol. Surv. S. Afr., Rep. 3 [1922], Spec. Rep 5, 1924: 14). 

161. Santos, Rio de Janeiro and general region, Sao Salvador (Bahia), Rio Parahyba, and Para in Brazil; Venezuela; 
Colon; Yucatan; Guadeloupe, Martinique, St. Vincent, Jamaica, and Cuba among the West Indies; the Florida 
Keys; various Florida localities along the east coast northward to New Smyrna and the St. Johns River, and along 
the west coast to Pensacola; Corpus Christi and the offing of Aransas Bay in Texas; also listed as a member of the 
fish fauna of Louisiana. 



Fishes of the Western North Atlantic i 2 i 

the Caribbean region in general, to northern Florida on both coasts, to Texas, and 
perhaps to Louisiana (not yet reported there). 

To the northward it has been reported once from Charleston, South Carolina (a 
newborn specimen), as well as from Southport, Beaufort, and Cape Lookout, North 
Carolina. The Bight immediately south of the Cape appears to act as a sort of cul de sac 
for such Narcine as wander that far north, for 13 out of 23 specimens reported near 
Cape Lookout during the summers of 1909, 19 10, and 191 2 were taken in this Bight. ^^^ 
This marks the extreme northern limit of its range, so far as known. Southward from 
southern Brazil it has been reported only from Bahia Samboronbon at the southern 
side of the entrance to the Rio de la Plata, and from Bahia Blanca on the northern 
Argentina Coast in about Lat. 39°3o' S. 

Captures of Narcine brasiliensis off the Texas Coast in the months of September, 
November, and March show that it winters that far north and probably does likewise 
at least along the southern part of Florida. However, to the northward along the 
Atlantic Coast of the LTnited States all the records of it, except one, have been in summer. 
Here its visits appear also to be brief, for while it reaches Cape Lookout in many sum- 
mers, if not yearly, it has been found inshore there only between June 27 and July 8.^** 
But a few must winter offshore as far north as this, for one was trawled in 20 fathoms 
off the Cape in February 1950,^"* and two other Torpedo Rays, probably of this species, 
were reported as taken in 34—35 fathoms nearby in January of that same year. 

Synonyms and References: 

Torpedo brasiliensis Olfers, die Gattung Torpedo, 1831: 19, pi. 2, fig. 4 (descr., color, ill., Rio de Janeiro, 
Brazil). 

Narcine brasiliensis Henle, Ueber Narcine, 1834: 31, pi. i, figs, i, 2, pi. 4, figs. 1-4, 8 (descr., meas., ills., 
cranium with attached cartilages, teeth; Rio de Janeiro); Miiller and Henle, Plagiost., 1841 : 129 (descr. 
meas., color varieties, Rio de Janeiro and W. Indies); Dumeril, Rev. Mag. Zool., (2) 4, 1852: 272 
(descr., color, Brazil, Martinique, Guadeloupe) ;i** Hist. Nat. Poiss., j, 1865: 514, pi. 11, fig. 3 (descr. 
color, ills, mouth, teeth; Antilles and Brazil); Kner, 'Novara' Exped., Zool., I, Fische, 5, 1865: 418 
(descr., color variations, size; Rio de Janeiro); Giinther, Cat. Fish. Brit. Mus., 8, 1870: 453 (diagn.. 
Para, Caribbean, Cuba, but ref. to C. of Good Hope probably erroneous; see p. 120, footnote 160); Goode 
and Bean, Proc. U. S. nat. Mus., 5, 1882: 240 (listed. Gulf of Mexico); Jordan, Proc. U. S. nat. Mus., 
7, 1884: 105 (cf. N.umirosa Jordan, n.sp.); Fritsch, Arch. Anat. Physiol., Leipzig (1886), Physiol. 
Abt., 1886: 353 (av. no. electric columns); Jordan, Rep. U. S. Comm. Fish. (1885), 1887: 799 (listed, 
W. Indies and Florida Keys); Garman, Bull. Mus. comp. Zool. Harv., ly, 1888: 93, pi. 33 (mucous 
canal syst.); Bean, Bull. U. S. Fish Comm., 8, 1890: 206 (listed, Cozumel, Yucatan); Fritsch, Elektr. 
Fische, 2, i8go: 41, 98, pi. 9, figs. 17, 18, pi. 10, fig. 20 (descr., varieties, av.no. of columns in electric 
organ, ills.); Jordan and Evermann, Bull. U. S. nat. Mus., 4y (i), 1896; 78 (descr., W. Indies and Brazil 
to Key West and Pensacola, Florida); Rep. U. S. Comm. Fish. (1895), 1896: 222 (listed, W. Indies 
and Brazil to Key West and Pensacola, Florida); Jordan and Rutter, Proc. Acad. nat. Sci. Philad., 4g, 
1897: 92 (listed, Jamaica); Duerden, J. Inst. Jamaica, 2, 1899: 614 (listed, Jamaica); Evermann and 
Kendall, Rep. U. S. Comm. Fish. (1899), 1900: 49 (listed, Florida Keys, Key West and Pensacola); 

162. Gudger, Proc. bid. Soc. Wash., 26, 1913: 99. 

163. Coles, Bull. Amer. Mus. nat. Hist., 32, 1913: 33. 

164. Specimen 250 mm long, examined by us, trawled by the U. S. Fish and Wildlife Service steamer Albatross III, 
Lat. 34°oi' N, Long, ^6°2■i' W. 

165. Credited by Dumeril (1852, 1865) to the Cape of Good Hope by reference to the Torpedo ocellata of Quoy and 
Gaimard (Voy. 'Uranie', Zool., 1824: 199); but the latter probably was not a Narcine (see p. 120). 



12 2 Memoir Sears Foundation for Marine Research 

Jordan and Evermann, Bull. U. S. nat. Mus., ^7 (4), igoo: pi. 13, figs. 35, 35a (ills.); Schreiner and 
Ribeiro, Arch. Mus. nac. Rio de J., 12, 1903: 81 (listed, S. Salvador de Bahia, Brazil); Werner, Zool. 
Jb., Syst. Abt. 21, 1904: 296 (listed, Jamaica); Ribeiro, Arch. Mus. nac. Rio de J., l^, 1907: 180, 208, 
pi. 17 (diagn., refs., photo, Rio de Janeiro to Pensacola, Florida); Aller, Science, N. S. J7, 1910: 712 
(listed, C. Lookout, N. Carolina); Coles, Bull. Amer. Mus. nat. Hist., 28, 1910: 337 (habits, electric 
shocks, nos., 1909, 1 9 10; C. Lookout, N. Carolina); Eigenmann, Rep. Princeton Exped. Patagonia, j 
(4), 1910: 377 (entering rivers); Fowler, Proc. Acad. nat. Sci. Philad., 62, 1910: 473 (listed, Key West, 
Florida); Bean and Weed, Proc. U. S. nat. Mus., ^o, 191 1: 231, pis. 10, 11 (colors, sizes, photos, em- 
bryos and adult; Long Key, Florida); Coles, Bull. Amer. Mus. nat. Hist., J2, 1913: 33 (abundance, 
season, no. embryos; C. Lookout, N. Carolina); Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 4. (3), 
1913: loi (range); Garman, Mem. Harv. Mus. comp. Zool., j6, 1913: 297, pi. 26, fig. i (descr., 
ill., Brazil); Gudger, J. Elisha Mitchell sci. Soc, 28, 191 3: 157, 158 (stom. contents, nos., C. Lookout, 
N. Carolina); Proc. biol. Soc. Wash., 26, 1913: 99 (nos., C. Lookout, N. Carolina); Coles, Proc. biol. 
Soc. Wash., 2<5, 191 5: 93 (season, C. Lookout, N. Carolina); Copeia, 32, 1916: 46 (doubts taxonomic 
value of color); Ribeiro, Rev. Mus. paul., JO, 1918: 708 (listed, Santos, Ilha San Sebastiao and Uba- 
tuba, Brazil); Lahille, Physis B. Aires, 5, 1921: 63 (listed, Argentina); Fowler, Proc. biol. Soc. Wash., 
j6, 1923: 28 (listed, Key West, Florida); Meek and Hildebrand, Field Mus. Publ. Zool., 75 (i), 1923: 
73, pi. 3 (descr., color, ill., Col6n); Ribeiro, Fauna brasil., Peixes, 2 (i) Fasc. i, 1923: 37, pi. 18 (diagn., 
cf. Narcine brach'sfleura Ribeiro, n. sp., photo, Pensacola, Florida to Rio de Janeiro); Borodin, Bull. 
Vandcrbilt Oceanogr. (Mar.) Mus., I (i), 1928: 5 (listed, no local.); Lahille, An. Mus. nac. B. Aires, 
24, 1928: 338, text fig. 27 (meas., discuss., ill., Bahia Samborombon and Isla Verde, Bahia Blanca, 
Argentina); Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 33 (genl.); Jordan, Manual Vert. Anim. 
NE U. S., 1929: 17 (genl.); Luderwaldt, Rev. Mus. paul., 16, 1929: 40 (listed, Ilha Sao Sebastiao, 
Brazil); Jordan, Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 27 (listed, Brazil to 
Key West and N. Carolina); Gowanloch, Bull. La. Conserv. Dep., 21, 1932: 86 (included in Gulf 
Mexico fish fauna); Bull. La. Conserv. Dep., 23, 1933: 239 (ill. after Meek and Hildebrand, 1923; 
listed, Louisiana); Beltran, List. Feces Mexicanos, 1934: 12 (listed, Mexico; not seen); Pozzi and Bor- 
odale. An. Soc. cient. argent., 120, 1935: 154 (listed, Argentina, Lat. 39°3o' S); Bere, Amer. Midi. Nat., 
Ij, 1936: 582 (copepod parasites. Lemon Bay, W.Florida); Smith, H. W., Biol. Rev. (Cambridge), 
II, 1936: 65 (in fresh water by ref. to Eigenmann, 1909); Breder and Springer, Zoologica N. Y., 2^, 
1940: 431 (electric discharges, sex ratios of embryos. Palmetto Key and Punta Gorda, Florida); Dels- 
man, Mem. Mus. Hist. nat. Belg., (2) Fasc. 21, 1941 : 65 (depth, Venezuela); Fowler, Proc. Acad, 
nat. Sci. Philad., 95, 1941 : 82 (listed, Sanibel, Florida); Hildebrand, Copeia, 1941 : 222 (listed, C. 
Lookout, N. Carolina); Rohl, Fauna Descript. Venezuela, 1942: 370 (listed, Venezuela; not seen); Cox 
and Breder, Zoologica N. Y., 28, 1943: 45 (meas. adults, embryo, color pattern, electric organs, meas. 
of discharges by oscillograph; Palmetto Key, Florida); Fowler, Monogr. Acad. nat. Sci. Philad., 6, 1944: 
456 (listed, Jamaica; Panama); Monogr. Acad. nat. Sci. Philad., 7, 1945: 98, 160, 264, 371 (listed, 
Southport, N. Carolina; Charleston, S. Carolina; Boca Chica and Key West, Florida; Texas); Gunter, 
Publ. Inst. mar. Sci. Texas, J, 1945 : 21 (colors of young, temp., salinities, season; Laguna Madre, Texas, 
and Gulf of Mexico off Aransas Bay); Schultz, Proc. U. S. nat. Mus., gg, 1949: 33 (listed, Venezuela). 

Torpedo bancrofti Griffith, in Cuvier, Anim. Kingd., 10, 1834: pi. 34 (colored ill., no local.). 

Torpedo pictus Gray, Cat. Fish. Coll. and Descr. by L. T. Gronow, in Brit. Mus., 1854: 13 (diagn., Antilles). 

Narcine brasiliensis var. punctatus Garman, Bull. Mus. comp. Zool. Harv., 8, 1881: 233 (color, cf. N. brasi- 
liensis; St. Vincent, W. Indies). 

Narcine brasiliensis var. corallina Garman, Bull. Mus. comp. Zool. Harv., 8, 1881: 234 (color. Key West, 
Florida); Mem. Harv. Mus. comp. Zool., j6, 1913: 298, pi. 26, fig. 3 (color, ill., Florida Keys). 

Narcine corallina Jordan, Proc. U. S. nat. Mus., 7, 1884: 105 (var. corallina Garman 1881, recognized as 
variety of species). 

Narcine umbrosa Jordan, Proc. U. S. nat. Mus., 7, 1884: 105 (descr.. Key West, Florida, based on dark-blotched 
specimens). 

Narcine, no spec, name. Coles, Bull. Amer. Mus. nat. Hist., J2, 1913: 33 (embryos). 

Narcine brasiliensis var. bancrojti Garman, Mem. Harv. Mus. comp. Zool., j6, 191 3 : 298, pi. 26, fig. 3 (color, 
ill., Jamaica to St. Vincent). 

Narcine brasiliensis corallina Radcliffe, Bull. U. S. Bur. Fish., J^, 1916: 270 (descr., ill. teeth, rep. by Jenkins, 



Fishes of the Western Nortli Atlantic 123 

1887, of Torpedo occidentalis was Narcinc brasiliensii; C. Lookout, N. Carolina); Coles, Copeia, 32, 
1916: 46 (specific status). 

Probable Synonyms: 

tTorpedo occidentalis Jenkins, Johns Hopkins Univ. Stud. BioL, 4, 1887: 84 (listed, Beaufort, N. Carolina). 

Narcine entemedor ]orda.n and Starks, Proc. Calif Acad. Sci., (2) 5, 1895: 387 (descr., nos.. La Paz, Mazatlan, 
Mexico); Gilbert and Starks, Mem. Calif Acad. Sci., 4, 1904: 15, 207 (descr., Panama Bay); Osburn 
and Nichols, Bull. Amer. Mus. nat. Hist., J5, 1916; 144 (Pacif Coast, Mexico); Meek and Hildebrand, 
Field Mus. Publ. ZooL, 75 (i), 1923: 74 (cf N. brasi/iensis); Breder, Bull. Bingham oceanogr. Coll., 
2 (i), 1928: 5, fig. 2 (color, photos, Pacif Coast, Mexico); Beebe and Tee-Van, Zoologica N. Y., 28, 
1941: 247 (color, ill.. Gulf of California). 

Tetronarce occidentalis Smith, N. C. geol. econ. Surv., 2, 1907: 43 (at least in part, C. Lookout, N. Carolina). 

Narcine brachypleura Ribeiro, Fauna brasil., Peixes, 2 (i) Fasc. i, 1923: 36, pis. 16, 17 (diagn., photos, adult 
male, Brazil, supposed to differ from A'', brasiliensis in relatively narrower disc, but probably only an 
extreme variant). 

Doubtful Synonyms: 

Narcine nigra Dumeril, Rev. Mag. Zool., (2) 4, 1852: 276 (cf N. brasiliensis, Brazil); Hist. Nat. Poiss., I, 
1865: 515, pi. II, figs. 4, 4a (descr., color, ill. mouth, teeth, supposed to differ from N. brasiliensis in 
shape of fins, rounded teeth and smaller eye; only one specimen seen); Bertin, Bull. Mus. Hist. nat. 
Paris, (2) II, 1939: 82 (type specimen in Paris). 

Genus Diplobatis Bigelow and Schroeder 1948 

Diplobatis Bigelow and Schroeder, J. Mar. Res., 7 (3), 1948: 562; type species, Discopygc omrnata Jordan and 
Gilbert 1890. Pacific Ocean, off coast of Columbia, 33 fathoms. 

Generic Characters. Nostril subdivided about midway of its length into two separate 
apertures by a cross bridge of stiff tissue. Teeth entirely concealed within mouth 
when latter is retracted and closed. ^^^ Median sector of each lip marked off at either 
end by a deep transverse groove when the mouth is retracted. Characters otherwise 
those of Narcine (p. 108). 

Size. The largest specimen reported was only 198 mm long. 

Range. Known only from the coast of British Guiana in the Atlantic and from the 
Gulf of California to Panama in the Pacific. 

Species. Two species have been described, D. pictus from the Atlantic and Z). om- 
rnata from the Pacific. D. pictus is distinguished from £). omrnata by angular pelvics 
which are evenly rounded along their outer margins in the latter. Z). pictus has a disc 
no wider than long and the tips of its dorsal and caudal fins are pointed, whereas in 
D. ommata the disc is wider than long and the tips of the dorsal and caudal fins are 
rounded. A conspicuous circular marking on the center of the disc, present on Z). om- 
rnata^ is lacking on D. pictus. 

Key to Species 

I a. Disc scarcely as wide as long; tips of dorsals and caudal pointed; anterior margin 
of pelvics not concealed by pectorals. pictus Palmer 1950, p. 124. 

166. In all other genera of the family in which the mouth is protractile [Narcine, Benthobatis, Discopyge, and probably 
Heteronarce), the tooth bands extend so far forward that several of the anterior rows of teeth are fully exposed even 
when the mouth is tightly closed. 



124 Memoir Sears Foundation for Marine Research 

lb. Disc wider than long; tips of dorsals and caudal rounded; anterior margin of 
pelvics concealed by pectorals. ommata (Jordan and Gilbert) 1890. 

Eastern Pacific from Gulf of California to 
Panama. 

Diplobatis pictus Palmer 1950 

Figure 25 a 
Study Material. None. 

Distinctive Characters. The closest relatives of D. pictus in the western Atlantic are 
Benthobatis marcida. Torpedo nobiliana, and Narcine brasiliensis. It may be separated from 
B. marcida by its relatively large eyes, from T. nobiliana by the strongly convex anterior 
outline of its disc and from N. brasiliensis^ which it most closely resembles, by its divided 
nostrils and concealed teeth (p. 123). 

Description. Proportional dimensions in per cent of total length. Female, 137 mm 
long, holotype, from off Georgetown, British Guiana (British Museum [Natural 
History]). 16' 

Disc: extreme breadth 46.7; length 47.5. 

Snout length: in front of orbits 10.9; in front of mouth 13.1. 

Orbits: horizontal diameter 3.6; distance between 6.6. 

Spiracles: length 3.3; distance between 6.6. 

Mouth: breadth 6.6. 

Nostrils: distance between inner ends 5.1. 

Gill openings: lengths, ist 1.5; 3rd 1.8; 5th 1.5; distance between inner ends, 

ist 13.9; 5th 9.5. 
First dorsal fin: vertical height 8.0; length of base 6.6. 
Second dorsal fin: vertical height 8.7; length of base 6.2. 
Caudal fin: upper anterior margin 13.9; lower anterior margin 13.2. 
Pelvics: anterior margin 14.6. 
Distance: from tip of snout to center of cloaca 51.8; from center of cloaca to 

tip of tail 48.2. 
Interspace between: ist and 2nd dorsals 3.6; 2nd dorsal and caudal 5.8. 

"Disc obtusely rounded, spade-shaped, its greatest width being opposite the last 
gill-cleft. Margin of snout elliptical. Length of disc equal to its greatest width, con- 
tained 2.1 times in total length and 1.9 times in length to caudal notch. Interorbital 
width twice in length of snout (preocular). Spiracles, greatest length of which is con- 
tained twice in interspiracular width, situated Immediately behind the orbits. Posterior 
margin of each spiracle fringed with 7-8 small papillae. Interspiracular width con- 
tained twice in pre-oral length of snout plus nasal curtain. Anterior aperture of nostril 
exposed, separated by a bridge of tissue from the concealed posterior aperture. Nasal 
curtain much wider than long, Its free edge slightly crenulated. Mouth narrow, pro- 

167. These proportions were furnished by G. Palmer, Dept. of Zoology, British Museum (Natural History). 



Fishes of the Weste?'n North Atlantic 



125 




Figure 25 a. Diplobatis pictus, holotype, female, 137 mm long. A Right-hand side of nasal curtain raised to 
show the transverse division of the nasal aperture, 3 X . 5 Ventral view of pelvic fins, i .5 X ■ From G. Palmer. 



tractile. Teeth diamond-shaped, in bands, each tooth with a small pointed denticle 
at its posterior angle directed inwards. Teeth concealed by lips when mouth is fully 
retracted and closed. The first dorsal commences slightly in advance of a vertical from 
the posterior edge of the pelvics. The base of the first dorsal a little longer than that 
of the second dorsal. Tail compressed dorso-ventrally, its depth 1.3 in its width im- 
mediately before origin of second dorsal. Two lateral dermal folds, commencing below 
posterior end of first dorsal base and terminating below caudal notch. Length of tail, 



12 6 Mej?ioir Sears Foundation fo?' Marine Research 

measured from posterior end of slit-like vent to caudal notch, 1.3 in length of disc. 
Caudal fin broad, almost as deep as long. The dorsal and ventral edges nearly straight, 
posterior margin rounded. Upper lobe a little longer and more acute than lower. Skin 
smooth. Pelvic fins separate one from the other and free from the tail at their extreme 
tips. Their outer anterior margins not concealed by posterior end of pectorals. 

Colour in Spirits. Dorsal surface light brown, overlaid with a series of darker 
brown blotches and white and brown spots. There is a series of irregularly rounded 
dark brown blotches arranged symmetrically along the outer margins of the pectorals 
and pelvics, with another pair immediately in front of the eyes. Behind the spiracles 
and covering the centre of the disc and anterior part of the tail is a series of scattered 
dark brown spots, interspersed with some white spots and larger dark brown blotches. 
Dorsals and caudal light brown. At the base of both dorsal fins is a dark brown blotch 
which extends a short way on to the fins. Near the tip of each dorsal is a dark brown 
band. The caudal fin is crossed by two brown bands, one near the base, the other near 
the tip. A narrow band of white along the tree margins of pectorals and pelvics. Ventral 
surface white."'®* 

Si%e. The single specimen known was 137 mm in total length and it is probable 
that the species does not reach a large size. 

Developmental Stages. Nothing is known. The one specimen described was a 
female, probably immature. 

Habits. There is no information as to the breeding habits or the diet of this species. 

Range. Known only from off Georgetown, British Guiana. 

Reference : 

Diplobatis pictus Palmer, Ann. Mag. nat. Hist., (12) J, 1950: 480 (descr. holotype). 



Genus Benthobatis Alcock 1898 

Benthobatis Alcock, Ann. Mag. nat. Hist., (7) 2, 1898: 144; type species, B. moresbyi Alcock. India, ofF Tra- 
vancore Coast, 430 fathoms. 

Generic Characters. Benthobatis agrees with Narcine in its relative length of tail, 
shapes of fins, wholly naked skin, proportions of anterior nasal flap, protractile mouth 
with labial cartilages, and single cusp on teeth. But it diflfers from Narcine in that the 
eyes are either entirely rudimentary or at least minute and seemingly blind, the cutaneous 
folds along the tail are absent or replaced at most by a low fleshy ridge, and the spir- 
acles are relatively smaller. In known species the caudal is oval, its lower posterior 
contour continuously rounded. Pelvics attached to sides of tail to their tips in some 
species, perhaps in alL^"' Skin in fresh specimens soft and loose, the body limp and 
flaccid."" The rostral cartilage is like that in Narcine, but with its lateral ridges softer 

168. From Palmer, Ann. Mag. nat. Hist., (12) j, 1950: 480. 

169. This is the case in Benthobatis marcida (p. 131, Fig. 26). But the account and illustration of 5. moresbyi {AXcoA, 
Ann. Mag. nat. Hist., [7] z, 1898: 145; 111. Zool. 'Investigator', 4, 1899, pi. 26, fig. i) fail to cover this point. 

170. This was the case in the considerable series of specimens from Cuban waters listed on p. 12S, and the skin was so 



Fishes of the Western North Atlantic 127 

and moderately divergent anteriorly; the antorbital cartilages are directed almost trans- 
versely outward, their tips with several branches. 

Size. The maximum length recorded for any member of the genus is 490 mm 

(197,6 in-, P- 13 0- 

Range. Benthobatis has been found only on the continental slope off South Carolina 
and northern Florida in 353 to 504 fathoms, along the northern coast of Cuba from 
150 to 351 fathoms, off Travancore in India, and in the eastern side of the Arabian 
Sea at 430 and 585 fathoms. 

The depths of capture, its apparent blindness, and the flabbiness of its body, 
suggest that Benthobatis is confined to at least moderately deep water. Beyond this, 
nothing whatever is known of the habits of the genus. Its embryos have not been 
seen. The capture of the considerable series (listed, p. 128) by the Atlantis in 1938 
and 1939 shows that it is more plentiful in suitable locations, latitudes and depths 
than previous information might have suggested. 

Species. Three species have been named: B.moresbyi Alcock 1898,^" from the 
coast of India and Arabian Sea; B. marcida and B. cervina Bean and Weed 1909,1'^ 
from the continental slope of the northeastern United States. B. moresbyi is distinguished 
by its uniformly velvet black hue from its Atlantic relative (or relatives) which are 
fawn-colored with white markings above and white below. Also, the disc of B. moresbyi 
is pictured 1" as being considerably narrower relatively than that in B. marcida. But 
the only account of B. moresbyi is so brief that proportional comparisons in other respects 
must await re-examination of it. 

Examination of the type specimens^'* of the two forms that have been named from 
the Atlantic has convinced us that they represent a single species. Indeed, in the original 
accounts the only comparison made between them states that the eyes of B. cervina 
are less reduced than those of B. marcida. 



Key to Species 

I a. Fawn-colored above, with white markings; white below. 

marcida Bean and Weed 1909, p. 128. 

lb. Purplish-black above and perhaps below.^" 

moresbyi Alcock 1898. 

Off Travancore Coast, southern India, 

and eastern side of Arabian Gulf.''^ 

described in the original account of the type species of the genus. However, other specimens that we have seen are 
firmer and the skins much stiffer, doubtless due to preservation in strong alcohol. 
171. Ann. Mag. nat. Hist., (7) 2, 1S98: 145. 172. Proc. U.S. nat. Mus., j6, 1909: 677, 679. 

173. Alcock, 111. Zool. 'Investigator', .;, 1899: pi. 26, fig. i. 

174. Now in the U. S. National Museum. 

175. It is not clear from the original account of 5. moresbyi (Alcock, Ann. Mag. nat. Hist., [7] 2, 1898: 145) whether 
the purplish-black color is confined to the upper surface or whether it extends to the lower surface as well. 

176. For list of references, see Fowler (Bull. U.S. nat. Mus., 100 [/j], 1941: 339)- 



12 8 Memoir Sears Foundation for Marine Research 

Benthohatis marcida Bean and Weed 1 909 

Deep-sea Electric Ray 

Figure 26 

Study Material. Five specimens, 167 to 490 mm long, including the types of 
Benthohatis marcida and B. cervina Bean and Weed 1 909, i" trawled on the continental 
slope off South Carolina and northern and middle Florida in 353-504 fathoms, in 
U. S. National Museum; also 60 specimens, 81 to 207 mm long, trawled in 1938 and 
1939 by the Atlantis in 150—351 fathoms at 26 localities scattered along the northern 
coast of Cuba."* 

Distinctive Characters. The only Rays in the western North Atlantic with which 
Benthohatis marcida might be confused are Torpedo, Narcine, and Diplobatis. It is 
easily separable from T. nobiliana by its much narrower protractile mouth and by the 
minuteness of its eyes. It is distinguishable from Narcine hrasiliensis (which it resembles 
in nature of mouth) by its degenerate eyes, by the oval shape of its caudal, by the 
relatively much greater length of its head anterior to the spiracles, by the union of the 
inner margins of its pelvics to their tips with the sides of the tail, by the replacement 
of the lateral membranous folds along the tail (characteristic of the genus Narcine) 
with low fleshy ridges, and by the softness and limpness of its body and skin (unless 
hardened in alcohol). Its nostrils separate it from Diplohatis. 

Description. Proportional dimensions in per cent of total length. Male, 184 mm, 
from off Matanzas, Cuba (Harv. Mus. Comp. Zool., No. 36984). Female, 430 mm, 
off S. Carolina, Lat. 32=39' N, Long. 77°oi' W (U. S. Nat. Mus., No. 37886). 

Disc: extreme breadth 38.7, 40.5; length 42.3, 42.6. 

Snout length: in front of eyes 13.6, 13.5; in front of mouth 12.5, 14.4. 

Eyes: horizontal diameter 0.0, 0.3; distance between — , 7.5. 

Spiracles: length 2.2, 2.1; distance between 7.6, 7.3. 

Mouth: breadth 4.5, 4.9. 

Exposed nostrils: distance between inner ends 7.4, 6.3. 

Gill openings: lengths, ist 1.9, 2.2; 3rd 1.9, 2.3; 5th 1.3, 2.0; distance between 
inner ends, ist 12.2, 11.4; 5th 7.9, 8.1. 

First dorsal fin: vertical height 4.6, 5.1; length of base 6.0, 7.7. 

Second dorsal fin: vertical height 6.0, 5.8; length of base 6.0, 7.7. 

Caudal fin: upper anterior margin 18.0, 19.3. 

Pelvics: anterior margin 9.8, 8.4. 

Distance: from tip of snout to center of cloaca 48.3, 45.7 ; from center of cloaca 
to tip of tail 51.7, 54-3; from snout to origin of ist dorsal — , 53.7. 

Interspace between: ist and 2nd dorsals 6.0, 6.1 ; 2nd dorsal and caudal ^.^, 5.8. 

177. Albatross Sta. 2626, Lat. 32°28' N, Long. 77°2i' W, 353 fath.; Sta. 2660, Lat. 28°4o' N, Long. yi°^6' W, 504 
fath.; Sta. 2664, Lat. 29°4i' N, Long. 79°55' W, 373 fath.; Sta. 2676, Lat. 32°39' N, Long. 77°oi' W, 407 fath. 

178. Atlantis Sta. 2938, 2980, 2981, 2982, 2983, 2984, 2999, 3305, 3387, 3388, 3391, 3392, 3410. 34i2) 34i3> 34i4) 
3416, 3417, 3418, 3421, 3422, 3428, 3436, 3437, 3478, 3482; for precise localities and depths, see Chace (Contrib. 
No. 274, Woods Hole oceanogr. Inst., 1940). 



Fishes of the IVestern NortJi Atlantic 



I 29 




Figure 26. Benthobatis marcida, female, 430 mm long, from off Charleston, S. Carolina, Lat. 32°39' N, Long. 
77°oi' W (U. S. Nat. Mus., No. 37886). A Ventral view with electric organs visible through the skin. B Side 
view of tail. C Eye and spiracle, about z."] x. D Nostril and mouth, about 1.2 X. ^ Cross section of tail between 
second dorsal and caudal fins, about 1.4 X. F Upper and lower teeth from center of mouth, about 6.8 X. 



Disc rounded, about as long as broad or a little longer (maximum ratio, length 
to breadth, about 1.2), the posterior margins of pectorals merging gradually with sides 



130 Memoir Sears Foundation for Marine Research 

of trunk (not appreciably recurved). Tail from center of cloaca about as long as dis- 
tance from center of cloaca to tip of snout on small specimens but a little longer rela- 
tively in adults; rounded above, moderately flattened below; sides of tail evenly rounded 
in small specimens (at least in most cases), but each with a low dermal ridge in adults 
from about opposite origin of first dorsal to origin of caudal. 

Snout anterior to spiracles about twice as long as breadth between spiracles, its 
length in front of mouth about 2.5 times as great as distance between outer ends of 
nostrils. Eyes separated from spiracles by a distance about equal to length of latter; 
minute, apparently nonfunctional, entirely covered over with skin and thus largely 
concealed in small specimens and in some mature ones, but exposed as dark dots up 
to one mm in diameter on other large specimens. Spiracles transverse or slightly oblique, 
about ^li—^U ^s long as distance between them, rounded posteriorly when expanded, 
their margins perfectly smooth. A pair of small mucous pores on nuchal region visible 
under a lens. All gill openings about as long as spiracle; distance between inner ends 
of first pair about 1.6 times as long as distance between spiracles; distance between 
inner ends of those of fifth pair about 65—72 "/o as great as that between first pair. 
Exposed portions of nasal apertures minute (even smaller than in Narcine) ; the isthmus 
separating their inner ends (visible only when nasal curtain is lifted) about half as broad 
as mouth when latter is retracted and closed (thus broader relatively than in N. bra- 
siliensis)\ nasal curtain thick, fleshy, about V4 ^s long as broad, its free margin with 
three low rounded expansions of about equal lengths; outer part of rear margin of 
nostril expanded as a low, rounded, fleshy lobe directed inward toward isthmus between 
both nostrils. Mouth with thick, wrinkled lips and surrounded by a deep moat; about 
'V3— V4 as wide as distance between spiracles when retracted and closed; forming a short 
tube (8 mm long in a 220-mm specimen) when protracted and opened. Tooth bands 
occupying ^U—^U of breadth of mouth. 

Teeth j| to ^; closely crowded in quincunx arrangement, the bases subquadran- 
gular; those of the younger series with a single slender cusp blunted at tip, directed 
rearward into the mouth; those of older series (exposed when mouth is closed) with 
cusps partly or wholly worn down; about seven rows in function simultaneously along 
centers of tooth bands, four to five rows toward outer extremities. 

Dorsals approximately equal in size and similar in shape; anterior margins strongly 
sloping; posterior margins weakly convex or nearly straight, recurving a little basally 
in young specimens but hardly so in adults; apex rounded narrowly; base a little less 
than */« (about 70 "/o) as long as anterior margin; vertical height a little less than length 
of base. Origin of first dorsal anterior to rear tips of pelvics by a distance about 7$ 
as long as base of first dorsal in young and in mature females; its position relative to tips 
of pelvics not known for mature males. Interspace between dorsals about Vs as long 
as base of first dorsal in adults but only about half as long in young. Interspace between 
second dorsal and caudal varying from Vs ^^ about as long as base of second dorsal. 
Caudal ovoid, its lower-posterior margin forming a continuous curve without inter- 
rupting corner; upper margin less strongly convex; tip well rounded in small specimens 



Fishes of the Western North Atlantic 131 

but somewhat more pointed in large; axis slightly raised; depth below tip of axis a 
little less than height above it; its length nearly as great as distance from origin of first 
dorsal to rear end of base of second dorsal. Pelvics conspicuously thick and fleshy, 
their origin either about opposite termination of pectorals or a little anterior; anterior 
margins nearly straight; outer corners broadly and evenly arcuate; outer margins 
ranging from nearly straight to slightly concave when fins are fully spread, but more 
or less wrinkled or even folded transversely when relaxed in what appears to be the more 
natural position ; inner margins of pelvics attached to sides of tail to extreme tips in 
both sexes by loose skin posterior to last radial cartilage; extreme length of pelvic, 
origin to tip, about half as great as breadth of disc, or a little less. Claspers of male 
originating a little anterior to posterior limit of pelvics and a little inward from outer 
pelvic margin. 

Electric organs more or less apparent externally (when viewed from below), the 
honeycomb structure showing through overlying skin; 200—220 columns in each 
organ of specimen 430 mm long. 

Color. Upper surface a delicate fawn color, clouded more or less with darker and 
paler and sometimes with a few small white spots irregularly distributed, shading to 
paler fawn or whitish along margins of disc, around spiracles, on sides of anterior 
part of tail and on upper parts of dorsals. Lower surface pure white or with pale yel- 
lowish or brownish tinge. After many years of preservation, a specimen from the con- 
tinental slope is light ochre brown and but little paler below. 

Size. The length at birth is less than 81—87 "^^n (smallest trawled by us off Cuba), 
the maximum recorded length (type specimen) 490 mm (19 Vie i^i-)- The size at which 
the two sexes mature is not known. 

Developmental Stages. Embryos have not been seen. 

Habits. This species is known only from depths ranging from 1 50 fathoms (young) 
down to 504 fathoms (adults) and at temperatures from as low as 5.5-8° C (42-46° F) 
on the continental slope of the southeastern United States to higher than i i°C (52° F) 
off Cuba. Presumably it lives on or near the bottom. Part of the leg of a small crustacean 
is the only recognizable object that was found among the partially digested stomach 
contents of two Cuban specimens. 

Range. Continental slope off South Carolina and Florida at 350—503 fathoms; 
off the north coast of Cuba, where young specimens appear to be generally distributed 
along the zone between the 150-200 fathom and 350 fathom contours (taken at 24 
stations out of a total of 166).'" 

Synonyms and References: 

Benthobatis marcida Bean and Weed, Proc. U. S. nat. Mus., j6, 1909: 677 (descr., meas., ill., color, size. 

Albatross Sta. on continental slope off mid-Florida, 504 fath.); Garman, Mem. Harv. Mus. comp. 

Zool., j6, 1913: 295 (descr. after Bean and Weed, 1909). 

179. 103 trawl hauls off northern Cuba at depths greater than 351 fathoms failed to yield any specimens, large or 
small, though adults are known to descend to depths a little greater than 500 fathoms off the southeastern United 
States. 



132 Memoir Sears Foundation for Marine Research 

Benthobat'ts cervina Bean and Weed, Proc. U. S. nat. Mus., j6, 1909: 679 (descr., color, meas., size, cf. B. 
moresiyi Alcock; continental slope off N. Florida, 373 fath.); Garman, Mem. Harv. Mus. comp. ZooL, 
36, 191 3: 295 (descr. after Bean and Weed, 1909). 

Suborder RAJOIDEA 
Skates and Their Immediate Relatives 

Characters. Head and body anterior to axils of pectorals strongly flattened dorso- 
ventrally. Snout neither produced as a long flat blade nor with lateral teeth; anterior 
sides of head without separate fin-like expansions of pectorals. Tail sharply marked off 
from body sector; moderately slender in most cases but extremely so exceptionally; 
with or without dermal folds along its sides. Pectorals united with sides of head nearly 
or quite to tip of snout, the whole forming a thin flat disc ranging in shape from sub- 
circular to more or less rhomboidal. Two dorsal fins, with cartilaginous rays,^ excep- 
tionally one or none; first much closer to tip of tail than to tips of pelvics. Caudal 
reduced to a small membranous fold, sometimes lacking in adults. Outer margins of 
pelvics more or less concave or notched, weakly so in some but so deeply so in others 
that anterior division of fin forms a separate limb-like structure; inner margins of 
pelvics either free or attached to sides of tail to their tips. Teeth numerous, rounded, 
or with conical cusp. Pelvis approximately straight, a prepelvic spur at each corner. 

Upper surface of disc more or less rough with prickles or thorns in most cases, 
but perfectly smooth in some. 

Front of cranium with a single unbranched rostral process, or none; antorbital 
cartilages not extending forward to help support anterior part of disc; tips of branchial 
rays not expanded as rounded plates. Pectoral, pelvic and dorsal fins without horny 
rays (ceratotrichia), the cartilaginous radials extending outward to margins; caudal 
membrane without radial supports. Surfaces of gill arches, inward from gill filaments, 
either smooth or with minute fleshy knobs. Spiracle with vestiges of gill folds. 

Families. The great majority of the members of this extensive suborder so closely 
resemble one another that they are united by common consent in a single family, the 
Rajidae, typical Skates. However, three genera depart from the typically rajid con- 
formation. Two of these, Anacanthohatis and Springeria, differ in that they have: a per- 
fectly smooth upper surface, no dorsal fins, pelvics attached to sides of tail nearly or 
quite to their tips, whip-like tail with membranous caudal, and snout produced at the 
tip as a soft filament ; the other, Arhynchobatis, differs in having only a single dorsal fin 
and a caudal that is larger than that of any other Skate.^ These features seem important 
enough to warrant the institution of the family Anacanthobatidae for the reception of 
Anacanthohatis and Springeria and of the family Arhynchobatidae for Arhynchobatis, 
but with the reservation that the need for the latter would vanish should it prove that 
the single known specimen of its type genus was abnormal as regards the number of 
dorsal fins. 

1. The type specimen of Raja garmani has three dorsals, an interesting reduplicative abnormality (p. 204). 

2. See Waite (Rec. Canterbury [N. Z.] Mus., i [2], 1909: 150, pi. 20) and Whitley (Fish. Aust., /, 1940: 192, fig. 221) 
for description and photograph of the unique specimen of Arynchobatis asperrimus Waite. 



Fishes of the IVestern North Atlantic 133 

Key to Families 

1 a. Two dorsal fins. Rajidae, p. 133. 
lb. Only one dorsal fin, or none. 

2a. One dorsal fin; disc prickly; inner margins of pelvics separate from tail. 

Arhynchobatidae. 

New Zealand.' 
2b. No dorsal fin; disc without prickles or thorns; inner margins of pelvics fused 
with tail nearly or quite to their tips. Anacanthobatidae, p. 327. 

Family RAJIDAE 

Characters. Tail only moderately slender, at most not more than about 2.0 times 
as long as body sector; rounded above, flattened below, with a narrow dermal fold 
along each side. Two dorsal fins. Pelvics with outer margins ranging from only weakly 
concave to so deeply so that anterior division of fin forms a limb-like structure entirely 
detached from the posterior part, three-jointed and (as it seems) separately movable. 
Orbits prominent, rising considerably above general level of head. Eye in some with 
expanded velum above the pupil. Spiracle close behind eye. Nostrils more or less 
oblique, their inner ends actually separate from mouth though superficially connected 
with outer corners of latter by shallow furrows; their expanded inner (anterior) margins 
joined across a broad isthmus in front of mouth and expanded rearward on either side 
as an extensive rounded curtain, smooth or more or less deeply fringed, roofing all but 
outer (anterior) ends of nasal apertures and reaching back to either corner of mouth; 
posterior margin of outer (exposed) part of nostril also expanded as a narrower flap, 
smooth-edged in some species but more or less deeply fringed in others, capable of 
assuming a more or less ring-like or tubular form, depending on the state of con- 
centration. Mouth moderately broad, transverse, its median sector more or less bowed. 
Teeth numerous, flattish to rounded or with one sharp cusp; arranged either in quin- 
cunx or in looser transverse series; several rows in function simultaneously. Skin of 
upper surface of disc and tail more or less rough with small prickles, larger thorn-like 
denticles, or both, but without serrate tail spines; lower surface either smooth or with 
prickly areas, in some cases with a few thorns. Pectoral girdle with a scapular element 
fused with the upper side of vertebral column. Pelvis nearly straight transversely, with 
a longer or shorter process extending forward from the outer corner at each end (Fig. 

2 7 A). Front of cranium with or without a rostral projection (rostral cartilage), the 
midline of snout stiff^er or softer accordingly.* Anterior rays of pectorals either close 
together at tip of snout (when rostral cartilage is short or lacking) or separated there 
by rostral cartilage. Characters otherwise those of the suborder (p. 132). 

3. Arhynchobatis Waite 1909 is placed by Fowler (Bull. U. S. nat. Mus., 100 [/j], 1941: 333) as a subfamily (Arhyn- 
chobatinae) of Platyrhinidae, which we refer here to the Rhinobatidae (p. 47). It is described (Waite, Rec. Canter- 
bury [N. Z.] Mus., I [2], 1909: 150, pi. 20) as not having a rostral cartilage and its resemblance in this respect to 
Psammobalis among Rajidae is pointed out. 

4. For a general account of the head skeleton and comparison with other batoids, see Holmgren (Acta Zool. Stockh., 
22, 1941 : 52, 64, 65). 



134 Memoir Sears Foundation for Marine Research 

Sex Characters. On the dorsal side of the outer part of each pectoral, adult males 
have from one to five rows of claw-like "alar" spines, retractile in grooves of the skin. 
It is supposed that they play some role in holding the female during copulation, but 
just how is not known. Also, in some species, either in female alone or in both sexes 
when mature, there may be a belt of large, nonretractile, so-called "malar" thorns on 
the outer anterior parts of the disc, between the anterior ends of the pectoral rays and 
the level of the spiracles. Furthermore, large thorns or bucklers are usually much 




Figure 27. A Pelvis oi Raja ocellata, female, about 790 mm long, from Waquoit, Massachusetts (Harv. Mus. 
Comp. ZooL, No. 359); dorsal view on right; about 0.5 x. B Gill arch (above) and lateral view of gill fold 
(below) of Raja laevis, from off Nantucket Island (Harv. Mus. Comp. ZooL, No. 36691), about 0.8 X. 



more highly developed or more numerous in adult females than in adult males; the 
latter are the smoother in general. The degree to which the anterior margins of the 
disc are concave in outline also differs in the two sexes in some species. 

Development is oviparous in all rajids in which it is known and doubtless in all 
other members of the family as well. The egg cases are quadrate with thick horny 
shells, their corners prolonged as long flexible horns (see also p. 141). 

The claspers' are cylindrical, narrowing gradually from base toward tip in early 
stages of growth but becoming more or less dilated terminally with the approach of 
sexual maturity, at which time they extend considerably beyond the tips of the pelvic 
fins. The seminal groove along the outer face of the clasper is wide open for a con- 

5. For recent accounts of the claspers in various European Skates, see Jungersen (Danish Ingolf Exped., 2 [2], 1898: 
56, pi. 4, figs. 45-57; pi. 6, figs. 67, 68), Huber (Zool. Anz., 32, 1901: 717), and Leigh-Sharpe (J. Morph., 34, 
1920: 260; 36, 1922: 236, 242, fig. 19), who shows the claspers of Kaja blanda when relaxed and distended; he also 
gives brief descriptions of various other European species (J. Morph., J9, 1924: 567-577)- 



Fishes of the IV es tern Nortfi Atlantic 135 

siderable distance at the anterior end*^ for the reception of the sperm and at the terminal 
end for its emergence.' Along the midsector of the groove, however, one of the edges 
overlaps the other, the two being held so closely in contact by the scroll-like form of the 
supporting cartilages that the channel is to all intents a closed tube although its edges 
are not actually fused. The supporting structures consist of: (a) two or more short basal 
joints that connect with the tip of the metapterygial cartilage of the pelvic fin; (b) an 
axial cartilage extending throughout most of the length of the clasper; (c) two narrow 
lateral cartilages (right and left) that develop along the outer face of the axial cartilage, 
to enclose the seminal groove as maturity approaches; and (d) three or more terminal 
cartilages, more or less movable, which vary in number and shape from species to 
species and which are often extremely complex; one or more of them may have spine- 
like or blade-like tips or edges. These terminal cartilages, which model the overlying 
tissues, often throw open the terminal section of the seminal channel, thus exposing 
the inner walls and a complex pattern of ridges and pockets; undoubtedly this is a func- 
tion associated with copulation. The function of the cutting structures has been the 
subject of speculation into which we need not enter here. 

Genera. One of the species that belongs here is set apart from all others, indeed 
from all other known batoids, by the conspicuous fact that several of the middle rays 
of each of its pectorals are elongated so as to form a narrow spatulate lobe (Fig. 77). 
It formed the basis of the genus Dactylobatus Bean and Weed 1909, which commonly 
has been accepted. The remaining members of the family, i. e., those of normal skate- 
like outlines, are divisible into two rather sharply defined groups according to the 
structure of the pelvics. 

A. The margins of the pelvics are so deeply notched as to give the anterior division 
the form of a slender subcylindrical limb which is entirely separate externally from the 
posterior division of the fin and which arises independently from the lower surface of 
the disc some distance in from the edge of the latter. This specialization is also accom- 
panied by interesting skeletal modifications (described on p. 314). Four species are 
known within this group, all of them referable to the genus Cruriraja Bigelow and 
Schroeder 1948 (p. 313). 

B. The anterior and posterior portions of the pelvics are continuous externally 
one with the other, as is usual among batoids, though the outer margin may be so 
deeply concave that the fin is definitely bilobed. This group includes all of the numerous 
members of the family that remain. Its representatives can be distributed among three 
subdivisions: 

1. Those in which the rostral projection from the front of the cranium (rostral 
cartilage) extends at least as far as the tips of the anterior rays of the pectoral fins and 
nearly to the extreme tip of the snout. 

2. Those in which the rostral projection is so reduced that its tip falls short of 
the tips of the anterior rays of the pectoral fins, and shorter still of the tip of the snout. 

3. Those in which there is no cartilaginous rostral projection from the cranium, 

6. The "apopyle" of Leigh-Sharpe. 7. The "hypopyle" of Leigh-Sharpe. 



136 Memoir Sears Foundation for Marine Research 

the snout being supported solely by the anterior pectoral rays, assisted more or less 
by strands or sheets of ligamentous tissue that extend forward from the anterior face 
of the cranium.* 

The members of Group i (i. e., those with long rostral cartilage and with pelvics 
and pectorals of the usual rajid shapes) are so many, and those from different seas so 
closely resemble one another in many cases, that it would be a boon to the student of 
fishes if this group could be subdivided generically by any criteria that would be easy 
to see or feel and still be sharply alternative. Various suggestions have been put for- 
ward, as is reflected in the considerable number of generic synonyms listed on p. 138 
under the genus Raja. But all the characters that have been tested (such as length and 
shape of snout, dermal armature, relative position of dorsal fins on tail, presence or 
absence of caudal fin-membrane, and structure of the claspers of adult males) inter- 
grade so completely that there seems no escape from the necessity of uniting all in 
the old genus Raja Linnaeus 1758. 

The members of Group 2, possessing a well developed rostral cartilage which 
falls short of the tips of the inner radial cartilages of the pectorals, are grouped together 
as the genus Breviraja Bigelow and Schroeder 1948. 

The members of Group 3 (i. e., those in which the rostral projection from the 
cranium is altogether lacking or only faintly indicated) have at one time or another 
been distributed among four genera, Sympterygia Miiller and Henle 1841,' Psam- 
mobatis Giinther 1870,^" Malacorhina Garman 1877, ^^ and Irolita Whitley 1931.12 It 
has been pointed out already that Malacorhina is not separable from Psammobatis," 
which appears to apply equally to Irolita. 

Our examination of available material," combined with published accounts and 
illustrations, lead to the conclusion that Psammobatis,^^ which has the outer margins 
of the pelvics deeply concave and hence definitely bilobed (Fig. 2 8 A), probably can 

8. Garman (Mem. Harv. Mus. comp. ZooL, 36, 1913: 369, pi. 68, fig. 3) characterizes the sheets of tissue that extend 
forward from the cranium between the pectoral rays of the two sides in the oldest named member of the genus 
{Sympterygia Miiller and Henle 1841) as "semi-cartilaginous." But it is more correctly described as "ligamentous" 
in the specimen he dissected, since it consists of a series of longitudinal bands, as is true of other representatives 
of the group that we have examined. 

9. Plagiost., 1 841: 155. 

10. Cat. Fish. Brit. Mus., 8, 1870: 470. Whitley (Rec. Aust. Mus., 18, 1931: 97) has pointed out that Psammobatis 
Giinther 1870 is antedated by Psammobates, proposed by Fitzinger (Ann. Wiener Mus., i [i], 1835: 113) for rep- 
tiles. But the use of it is permissible for Rays, according to the International Code of Zoological Nomenclature, 
just as the use of Mustelus Link 1 790 as a generic name for Sharks is permissible, although it was long antedated 
by Mustela Linnaeus 1758, for mammals. 

11. Proc. Boston Soc. nat. Hist., ig, 1877: 203. 12. Rec. Aust. Mus., 18, 1931; 97. 

13. Regan (Rep. Brit. Antarc. Exped., Zool., i [i], 1941: 22); and Norman (Discovery Rep., 16, 1937: 33). 

14. Three females, 222-720 mm long, seemingly referable to Sympterygia bonaparti Miiller and Henle 1841, from 
Uruguay and northern Argentina; one male, 505 mm long, and one female, 530 mm, of S. acuta Garman 1877, 
from Argentina; two females, about 475 and 560 mm long, of Raja microps Giinther 18S0, from Argentina; and 
one small male, about 190 mm long, of R. scobina Philippi 1857, from the vicinity of Montevideo; all the fore- 
going in U. S. National Museum; likewise the dissected specimens on which Garman based his illustrations of 
Sympterygia acuta Garman (Mem. Harv. Mus. comp. Zool., 36, 191 3: pi- 68, figs. 3,4) and of Malacorhina mira Gar- 
man (Mem. Harv. Mus. comp. Zool., j6, 1913: pi. 69, figs, i, 2). 

15. Type species Psammobatis rudis Giinther equals Raja scobina Philippi 1857. 



Fishes of the IVestern North Atlantic 



137 



be retained as generically distinct from Sympterygia,^^ in which these margins are so 
slightly concave that the fins, when spread, can hardly be characterized as bilobed 
(Fig. 28 B). This course is followed in the accompanying Key (p. 138), but with the 
reservation that future study, especially a more critical comparison of species that 
have been described recently from Peru, may prove that a complete gradation exists 
between these extremes.^' 





A \ 



Figure z?,. A Pelvic fins ol Psammobatis scobina, female, from Uruguay (U. S. Nat. Mus., No. 86721). B Pelvic 
Rns of Sympierygia microps, kma.\e, from Buenos Aires, Argentina (U. S. Nat. Mus., No. 55579). Each about 0.5 x. 



Key to Genera 

I a. Middle rays of each pectoral prolonged as a narrow spatulate process (Fig. 77). 

Dactylobatus Bean and Weed 1909, p. 323. 
lb. Middle rays of each pectora not prolonged as a narrow spatulate process. 

2 a. Anterior subdivision of pelvic fin forming a slender limb-like structure, en- 
tirely separate externally from posterior subdivision of fin, arising from lower 
surface of disc some little distance inward from edge of latter. 

Cruriraja Bigelow and Schroeder 1948, p. 313. 
2 b. Anterior subdivision of pelvic fin continuous externally with posterior sub- 
division, not a separate limb. 

3 a. Front of cranium with a definite conical rostral projection (rostral car- 
tilage), longer or shorter. 

4 a. Tip of rostral cartilage extending forward beyond extremities of pectoral 
rays to tip of snout, or nearly so. Raja Linnaeus 1758, p. 138. 

4 b. Tip of rostral cartilage falling short of extremities of pectoral rays 
and of tip of snout. 

Breviraja Bigelow and Schroeder 1948, p. 284. 

i6. Type species Sympterygia bonaparti Miiller and Henle 1841. 

17. See Norman (Discovery Rep., 16, 1937: 28) for a general synopsis of species referred by him to Psammobatts, and 

Hildebrand (Bull. U. S. nat. Mus., 189, 1946: 52) for a Key to Peruvian species, with accounts of four described 

as new. 



138 Memoir Sears Foundation for Marine Research 

3 b. Rostral projection from front of cranium lacking or only faintly indicated; 
anterior part of disc supported chiefly by anterior rays of pectorals more 
or less assisted by ligamentous bands extending forward from front of 
cranium. 

5 a. Outer margins of pelvics deeply concave or notched, the fins de- 
finitely bilobed, even when spread wide (Fig. 28 A). 
Psammobatis Giinther 1 8 70. 
Western South Atlantic from Cape Frio (Lat. 22^56' S) 
near Rio de Janeiro southward to Straits of Magellan; 
Chile, Peru, and vicinity of Cocos I. (Lat. 4°5o' N);i* 
also southern Australia. 
5 b. Outer margins of pelvics weakly concave, the fins not definitely 
bilobed when spread wide (Fig. 28 B). 

Sympterygia Miiller and Henle 1841. 
Argentina, Chile, Peru; perhaps Ecuador also.*^ 

Genus Raja Linnaeus 1758 

Raja Linnaeus, Syst. Nat., J, 1758: 231; tj'pe species, R. clavata Linnaeus 1758. European seas, designated 
by Jordan and Gilbert (Proc. U. S. nat. Mus., 5, 1882: 36). 

Generic Synonyms: 

Rata Scopoli, Introd. Hist. Nat., 1777: 464; emended spelling for Raja Linnaeus 1758. 

Dipturus Rafinesque, Caratt. Gen. Spec. Sicil., 1810: 16; type species, Raja batis Linnaeus 1758. European seas. 

Cefhaleutherus Rafinesque, Indice Ittiol. Sicil., 1810: 48, 61; type species, C.maculatus Rafinesque. Sicily. 2* 

Platopterus Rafinesque, An. Nature, 181 5: 93; proposed to replace Raja Linnaeus 1758. 

Dasybatus Blainville, Bull. Soc. philom. Paris, 1816: 112; type species. Raja clavata Linnaeus 1758.^1 

Proplerygia Otto, Nova Acta Acad. Leop.-CaroL, 10, 1821: 112; type species, P. hyposticla Otto.-- Scotland 

near Edinburgh. 
Dasy batis Blainville, in Vieillot, Faune Fran?., Poiss., 1825: 12; type species. Raja batis Linnaeus 1758. 

European seas; designated by Jordan (Genera Fish., I, 191 7: 134).^' 
Uraptera Miiller and Henle, S. B. Akad. Wiss. Berlin, 1837: 117; also Arch. Naturg., (Jahrg. 3) j, 1837: 

400; generic diagnosis but no species named; type species, I] . agassizii Miiller and Henle (Plagiost., 

1 841: 155, pi. 50). Brazil. 
Laevirajia Bonaparte, Nuov. Ann. Sci. Nat. Bologna, 2, 1838: 203; type species. Raja oxyrincha Linnaeus 

1758. Mediterranean and European seas. 

18. Psammobatis spinosissimus Beebe and Tee-Van 1941 (Zoologica N. Y., 26, 1941: 259), described from vicinity of 
Cocos Island. 

19. Reported by Tortonese (Boll. Mus. Zool. Anat. comp. Torino, [3] 4^ [89], 1939: 48), but the description is not 
sufficient for judgement as to specific identity. The type species Sympterygia bonaparti Miiller and Henle 184: was 
without locality. 

20. Based on an abnormal Skate in which the anterior e-xtensions of the pectorals had remained separate from the sides 
of the head after birth. Similar monstrosities have been described subsequently under other names, as noted below. 

21. /?<2;'a A<2/!j Linnaeus 1758 was designated as type by Fowler (Bull. U. S. nat. Mus., 100 [jj], 1941: 355). But ifl/w was 
not among Blainville's list of 22 species, so we propose that K. clavata replace it, since it was among his species. 

22. As Miiller and Henle (Charlesworth Mag. Nat. Hist., 2, 1838: 90, footnote) pointed out long ago, the specimen 
described and pictured by Otto was a monstrosity of the same sort that had been named Cephaleutherus earlier by 
Rafinesque 18 10, the anterior part of each of its pectorals forming a narrow ear-like lobe that suggests the cephalic 
fins of the Devil Rays (Mobulidae). Similar malformations have been reported from time to time. Couch (Fish. 
Brit. Isles, j, 1867: 96) pictures one with a secondary pectoral lobe on the one side but not on the other. 

23. Jordan credits this genus to De Serville. However, the section on fishes in Faune Francaise seems to have been 
by Blainville. For dates of publication, see Sherborn and Woodward (Ann. Mag. nat. Hist., [7] 8, 1901: 493). 



Fishes of the Westerfi Nortfi Atlantic 139 

Propleygia Miiller and Hcnlc, Charlcsworlh Mag. Nat. Hist., 2, 1838: 90, footnote; equals Proplerygia Otto 

1821. 
Laeviraja Bonaparte, Icon. Fauna ItaL, j Fasc. 23, 1839: in descr. oi L. oxyrychus, pi. not numbered; equi- 
valent to Laevirajia Bonaparte 1838. 
Balis Bonaparte, Mem. Soc. Neuchatel Sci. Nat., 2, 1839: 7, extra; type species. Raja radnia Delaroche 1809. 

Balearic Isles, Mediterranean. Not Batis Boie 1833, for birds. 
Hicroptera Fleming, New philos. J. Edinb., JJ, 1841: 236, pi. 4; t}pe species, H. abrcdonensis Fleming. Scot- 

land.2« 
Actinobatis Agassiz, L., Poiss. Foss., j, 1843: 372; type species, Raia ornata .'\gassiz, a fossil. 
Eleutherocephalus Agassiz, L., Nom. Zool. Pise, 1846: 71, Index, 136; substituted for Cephaleutherus Rafi- 

nesque 18 10. 
Perioptera Gistel, Naturg. Tier., 1848: X; nomen nudum \ perhaps a misspelling for Hieroptera Fleming 1841. 

Not seen. 
Peropera Gistel, Naturg. Tier., 1848: X; same as Perioptera Gistel 1848. Not seen. 
Amblyraja Malm, Goteborgs och Bohuslans Fauna, 1877: 607; type species, Raia radiata Donovan 1807. 

Northern Great Britain. 
Leucoraja Malm, Goteborgs och Bohuslans Fauna, \%1l: 609; type species. Raja fullonica Linnaeus 1758. 

European seas. 
Alpharaia Leigh-Sharpe, J. Morph., jp, 1924: 567, 568; type species, "Raja circularis, the Cuckoo Ray."^^ 
Betaraia Leigh-Sharpe, J. Morph., J9, 1924: 568; type species. Raja clavata Linnaeus 1758. European seas. 
Gammaraia Leigh-Sharpe, J. Morph., J9, 1924: 567, 571 ; type species. Raja batis Linnaeus 1758. European seas. 
Deitaraia Leigh-Sharpe, J. Morph., jp, 1924: 567, 573; type species, Raia radiata Donovan 1807. Northern 

Great Britain. 
Epsi/onraia Leigh-Sharpe, J. Morph., J9, 1924: 567, 574; type species. Raja platana Giinther 1880. Estuary 

of Rio de la Plata. 
Zetaraia Leigh-Sharpe, J. Morph., jy, 1924: 567, 575; type species. Raja brachyura Giinther 1880. Straits 

of Magellan and westward. 
Etaraia Leigh-Sharpe, J. Morph., J9, 1924: 567, 576; type species. Raja murrayi Giinther. Kerguelen I. 
Thetaraia Leigh-Sharpe, J. Morph., J9, 1924: 567, 577; type species. Raja eatonii Giinther 1876. Kerguelen I. 
lotaraia Leigh-Sharpe, J. Morph., J9, [924: 567, 577; type species. Raja marginata Lacepede 1802— 1803. 

France and England. 
Spiniraja (subgenus) Whitley, .'Vust. Zool., 9, 1939: 251; type species, Raja [Spiniraja) ogilbyi Whitley. Au- 
stralia.^* 
Zearaja Whitley, Aust. Zool., 9, 1939: 254; type species. Raja nasuta Miiller and Henle 1841. New Zealand. 
Pavoraja Whitley, Aust. Zool, 9, 1939: 254; type species, Raja nitida Ciiinther 1880. New South Wales. 
Rioraja Whitley, Aust. Zool., 9, 1939: 254; proposed in substitution for Vraptera Miiller and Henle 1837, 

preoccupied by Billberg, 1820, for Lepidoptera. 
Dentiraja (subgenus) Whitley, Fish, .'^ust., i, 1940: 184; type species. Raja dentata Klunzinger 1872. 
Argoraja (subgenus) Whitley, Fish. Aust., i, 1940: 190; type species. Raja polyommata Ogilby 191 o. 

Generic Characters. Disc subquadrangular to rhomboid, the snout ranging from 
short and obtuse to long and more pointed. Tail with a narrow ribband-like longitudinal 

24. A monstrosity similar to those described earlier under the generic names Cephaleutherus Rafinesque 1810 and Pro- 
pterygia Otto 182 1. 

25. "Cuckoo" is the common name for Raja naei'us MuUer and Henle 1S41, not for Raja circularis Couch 1838. There- 
fore it is doubtful which of these two species Leigh-Sharpe had in mind. For the history of R. circularis, see Clark 
(Rep. Fish. Bd. Scot. [1926], i, 1926: 37). Leigh-Sharpe proposed the names Alpharaia, Betaraia, Gammaraia, 
Deitaraia, Epsilonraia, Zetaraia, Etaraia, Thetaraia and lotaraia as "Pseudogenera"; but this term has no nomen 
clatural standing. 

26. Spiniraja is characterized as differing "from true Raja in being very spiny above and below." But no spines are 
indicated on Whitley's subsequent illustration (Fish. Aust., i, 1940: 186) of its lower surface; hence we cannot 
judge whether it is actually any rougher below than are some species of typical Raja. Nor do we see any characters 
to separate Whitley's genera Zearaja and Panjoraja, proposed without generic diagnoses (Aust. Zool., 9, 1939: 
254), or his subgenera Dentiraja and Argoraja (Fish. Aust., i, 1940: 184, 190). 



140 Me^noir Sears Foundation for Marine Research 

fold along either side. Pectorals of ordinary form, without spatula-like lateral process; 
outer posterior corners more or less broadly rounded, their posterior outlines definitely 
recurved before their junction with the sides of the trunk, the fin thus having a distinct 
inner posterior margin; pectorals overlapping pelvics somewhat in most cases. Two 
dorsal fins." Caudal reduced to a narrow finfold, sometimes lacking in adults. 
Outer margins of pelvics more or less concave but not cut deeply enough to separate 
anterior from posterior section of fin. Eyes with upper part of pupil covered by a dark 
fleshy veil or velum, with crenulate margin that can be expanded or contracted (con- 
trols amount of light entering pupil). Nostrils strongly oblique. Teeth either closely 
crowded in quincunx arrangement or more loosely spaced in transverse series; rounded 
in most females, cuspidate in mature males, and both sexes of some species^* with 
sharp cusp. Upper surface of disc with prickles or larger thorns, or both. Anterior 
margin of cranium with a distinct rostral projection (rostral cartilage) reaching nearly 
or quite to tip of snout, extending at least as far as extremities of pectoral rays or far- 
ther (see discussion p. 135). Anterior lobe of pelvic supported by three radials artic- 
ulated along posterior bar of pelvic arch, followed without intervening gap by first 
2—4 radials borne along anterior part of basipterygial cartilage. 

Young Skates at hatching are already so much like the adult in form that they 
are recognizable as to species, at least In most cases (for an exception, see p. 148; Key 
alternative 7 b). However, they may differ more or less from their parents In form of 
disc. In details of spination, and in possessing a somewhat longer tail. There is only 
one juvenile character that calls for special mention here, and that is the persistence for 
some little time after hatching of a greater or lesser part of the long caudal filament 
that is characteristic of embryo Skates in general during late embryonic stages. Dif- 
ferent species, and possibly different individuals even within a given species, appear 
to differ in this respect. Consequently, the length of the tail to the origin of one or the 
other of the two dorsal fins Is a more dependable character for diagnostic purposes than 
is the tail's total length. 

The electric organs, of species for which they are known, are slender spindle- 
shaped masses along the sides of the tail, extending nearly the entire length of the 
latter in some species but shorter In others. They also vary in thickness from bulky, 
occupying practically the entire space between the skin and the vertebral column, to 
much narrower and separated from the vertebral column by a thick muscular layer. 
In some species the electric elements consist of columns of discs at right angles to the 
longitudinal axis of the tail, whereas in others they are small club- or cup-shaped bodies 
with a longer or shorter tail-like prolongation. They have a rich blood supply and are 
Innervated with Innumerable fibrils which are derived from spinal rather than cranial 

27. Abnormalities in this respect have been described; see under R.garmani, p. 204, footnote 14. 

28. The alteration that takes place in the shape of the teeth in males of many species at sexual maturity results from the 
replacement of the older rows by younger ones, not from any alteration in the shape of the teeth present earlier. 
Examination of specimens approaching maturity also shows that this replacement takes place a little earlier in the 
upper jaw than in the lower. 



Fishes of the Western Nortli Atla7ttic 141 

nerves as in the Electric Rays. Their embryonic derivation is from the caudolateral 
musculature.''' 

In one set of observations, shocks of about one-half volt were recorded instrumen- 
tally per centimeter of length of organ. ^" But we find no recorded instance of a fisher- 
man reporting a shock from any Skates of this genus, though many thousands of them 
are handled yearly; nor have we ever felt the least sensation from grasping the tail of 
a Skate other than the rather unpleasant one of being wounded by the thorns. 

Sixe. The various species of the genus Raja of the western North Atlantic range 
in length from about iVa to about 5 feet, perhaps to as much as 6 feet, and in weight 
from about a pound to perhaps 50-60 pounds (p. 223). The females, at least in most 
cases, are larger than the males, often by as much as one-third. 

Developmental Stages. In the commercial catches in British waters the females 
considerably outnumber the males. This disparity appears to result from the fact that 
the two sexes are more or less segregated, the gravid females apparently congregating 
In more compact bodies or schools than the males. ^^ 

Skates are described as mating ventral side to ventral side, and pairs so engaged 
are sometimes hauled up on hook and line. It has been observed*^ that the males and 
females of one of the larger European species {R. batis) hold their discs flat while mat- 
ing; but the female of the smaller R. asterias curves her pectorals ventrally, while the 
male, rolling the outer corners of his pectorals out of the way ventrally, then bends the 
fins inward around her back, which brings his alar spines in position to fasten to her. 

At least for some of the larger species it is reported that only one clasper is intro- 
duced into the cloaca of the female at a time, but for other species it is said that both 
are introduced simultaneously.'* 

Just after copulation the claspers are much swollen and reddened by suffused 
blood.34 

Experiments*^ have proved that the egg is fertilized just before its capsule is formed 
around it. The process by which the sperm is introduced into the oviduct has not 
been traced for any Skate or Ray, but according to Clark the female apparently can 
lay a series of eggs fertilized by sperm from one act of coition. Completion of the egg 
capsules and extrusion of the egg appears to follow shortly after fertilization. 

Egg capsules of Skates are horny or leathery in texture, ranging in color from 
light auburn to dark brown or even black. The shell, composed of a substance resem- 

29. For a detailed account of the electric elements and their development in various Skates, see especially Ewart (Philos. 
Trans., [B] J79, 1888: 399-416, pis. 66-68; 1889: 539-552, pis. 79-80). 

30. See Sanderson and Gotch (J. Physiol., 9, 1888 : 141) for a detailed account of these experiments and of the general 
nature of the electric discharges. 

31. For summary of available evidence on this point, see Steven (J. Mar. biol. Ass. U. K., iS, 1933: 611). 

32. See Le Danois (Vie et Moeurs Poiss., Payot, Paris, 1949: 199, text figs. 55, 56) for an eye-witness account and 
illustrations of mating pairs of Raja batis and of R. asterias. 

33. Fowler (Rep. N. J. Mus. [1905], 1906: 730) quotes an eye-witness account of a pair of Raja eglanteria harpooned 
while in the act of coition, the male with both claspers inserted. 

34. See Leigh-Sharpe (J. Morph., j6, 1922: 242, fig. 10) for illustration of clasper of Raja blanda, relaxed and distended. 

35. Clark, J. Mar. biol. Ass. U.K., 12, 1922: 584. 



142 Memoir Sears Foundation for Marine Research 

bling keratin'* and constructed of several fibrous layers, becomes more brittle on ex- 
posure to sea water. The body of the capsule is quadrate in shape and the four corners 
are drawn out in long tubular horns, which taper to filamentous tips in some species 
but which are blunter in others, the pair at one end being considerably longer than 
those at the other. As a rule each horn has a slit-like opening on the outer side which 
is closed in early stages either by a plug of albumen or by a delicate membrane. Later 
the albumen is absorbed, leaving the slits open. Also, in some species the lateral margins 
of the capsule are keeled or fringed, and in most species (perhaps all) the margins, 
sometimes the sides as well, bear series of silky and sticky filaments; these mat into 
felt-like masses that serve to anchor the egg to algae, pebbles or other objects on the 
bottom. Before they are laid, the egg capsules lie with the longer pair of horns directed 
toward the cloaca and with the more convex face of the capsule toward the dorsal side 
of the mother. 

It appears that osmosis, which takes place through the shell and by which equi- 
librium is maintained with the surrounding sea water" provides the necessary oxygen 
for the embryo at first. The means by which the aeration of the embryo is accomplished 
after the albumen has been absorbed has been the subject of controversy. However, 
recent experiments seem to have confirmed the old view that sea water passes in and 
out through the slits on the horns of the capsule after they have opened, thus sup- 
plementing the effects of osmosis through the shell in providing the necessary intake 
of oxygen and outgo of carbon dioxide.'* 

Observations in aquaria have shown that the eggs of at least some species are 
laid in pairs, one soon after the other, with a rest period of one to five days before the 
deposition of another pair. A single female may produce eggs during a period of several 
months, in spring, summer, or fall. For example, one female of the European Raja 
brachyura Lafont 1873 ^^^^ 25 capsules at irregular intervals during a 49-day period 
(April 12 to May 31) in the aquarium at Plymouth, England. It has been observed 
also that the egg capsules in aquaria are buried in the sand. In most cases the capsules 
that have been dredged from the sea bottom have been firmly fastened to masses of 
broken shells, fragments of algae, gravel, sand, etc., and Skate eggs in the Plymouth 
Aquarium have been seen to adhere to objects of this sort immediately after they were 
laid. Under aquarium conditions, the incubation period for six common European 
species averaged from 4V2 to about 14V4 months. 

Normally, the embryo lies within the capsule with its head pointing diagonally 
toward the longer pair of horns, its pectorals folded over onto the dorsal surface and 
its tail curled forward over its back. At hatching it emerges through a transverse slit 
between the two longer horns,'^ and the empty capsules, popularly called mermaids' 
or sailors' purses, are familiar objects along the seashore. 

36. Its chemical composition was determined by Hussak and Welker (J. biol. Chem., 4, 1908: XLIV-XLV). 

37. This maintenance of equilibrium was demonstrated by Peyrega (Bull. Soc. zool. Fr., Jp, 1914: 211). 

38. See Clark (J. Mar. biol. Ass. U. K., 12, 1922: 5S4) for an account of these experiments, with summary of older 
observations on the aeration of the Skate embryo. 

39. See Clark (J. Mar. biol. Ass. U. K., 12, 1922: 582-591) for an interesting account of the egg capsule, egg laying, 



Fishes of the JVestern North Atlantic 143 

The embryonic Skate is more shark-like than batoid in its general appearance 
during the early stages of development; it is slender, with its pectorals and pelvics 
still represented only by a pair of short rounded lobes on either side and with one or 
two anal fin-folds besides the dorsals. The external branchial filaments, developed on 
the second to sixth gill arches, are a prominent feature of its general aspect at this 
stage, and the tip of its tail is drawn out as an attenuated filament, longer or shorter. 
The enormous development of the pectorals that finally leads to the "Skate" form com- 
mences while the external branchial filaments are still persistent and continues until 
after they have disappeared. The fusion of the pectorals with the sides of the head is 
normally completed some time prior to birth; however, abnormal specimens are some- 
times encountered in which the anterior part of one pectoral or the other, or both, 
continues separate from the head after birth and probably throughout life, as described 
elsewhere (p. 261). The yolk is entirely drawn within the abdomen some little time 
previous to hatching, and the abdomens of late embryos are commonly much swollen 
with it. Newly hatched Skates resemble their parents in general shape, except as noted 
(p. 140), and they hold as strictly to the bottom as the older ones. 

Habits and Food. Typically, Skates are ground fish, often lying partially buried. 
When alarmed, a Skate will usually press down against the bottom instead of dashing 
away, and the colors of their backs blend so closely with the background that they are 
as difficult to detect as any member of the flounder tribe. But they often rise some 
distance above the bottom in pursuit of prey, and occasionally a Skate may be seen 
at the surface, seemingly basking in the sun. 

They are most plentiful on smooth, sandy, gravelly bottoms, or on mixtures of 
sand and broken shells; they are found much less frequently on soft mud (see also p. 262). 
To find a Skate among rocks or on ledges is an unusual event, though we have occas- 
ionally seen them there. 

Observations of Skates in aquaria*" show that they are mostly inactive by day, 
lying quietly on bottom, often so much buried in sand up to the eyes, spiracles, and 
ridge of the back that their outlines are not to be seen. But they swim actively by day 
if disturbed, and after swimming they return to the bottom either by swimming or simply 
by sinking. To bury themselves they stir the sand by movements of their pectorals, thus 
allowing it to fall back upon them. 

They advance by gently undulating the margins of the wing-like pectorals as 
described previously (p. 8); the tail, if used at all, serves only as a rudder and often 
drags on bottom, leaving a trail behind it. They swim rather slowly unless violently 
disturbed, when they can advance with astonishing speed. 

Skates are strictly carnivorous and feed mostly at night. The diet for such species 
as have been studied consists chiefly of crabs and shrimps, lobsters, smaller Crustacea 
such as amphipods, isopods, mysids, and various polychaete worms. They also eat 

fertilization, duration of incubation, and other aspects of the early life history of several of the more common British 
Skates. 
40. Schmitlein, Mitt. Zool. Stat. Neapel., j, 1878: 9. 



1 44 Memoir Sears Foundation for Marine Research 

bivalve mollusks, small fish of various kinds, and cephalopods of the less active sorts. ^^ 
Occasionally pieces of eel grass {Zosterd) or algae, even pebbles, are found in the stomach 
of a Skate, no doubt taken accidentally with its living prey. 

A Skate cannot capture active animals by direct attack because of the ventral 
position of its mouth, so it swims over its victim by a sudden dart forward and settles 
down upon it, thus preventing its escape. In fact, in the English Channel, Skates some- 
times feed almost exclusively on fishes, especially herring. ^^ 

In the case of at least some of the European species (e. g., Raja clavata\ the 
young are hatched in shallow water, after which they gradually disperse to greater depths 
as they grow. Some of them also carry out feeding migrations on a considerable scale,** 
but no precise information is available in this respect for any of the western Atlantic 
Skates. 

Numerical Abundance. Skates may be extremely numerous locally on suitable bot- 
toms in temperate and boreal latitudes. In northern Europe, where they are valued for 
human food, their abundance is reflected in the considerable landings in various coun- 
tries," for example, about 92,100,000 pounds in 1937, and 92,400,000 pounds in 
1936, which is equivalent to about 18-46 million individuals, assuming the average 
weight per Skate to be about 2—5 pounds. 

Statistics of the commercial catch in North America do not afford comparable 
information for the western Atlantic, for most of the Skates are thrown back. In 1 940 
the total landings reported for the entire Atlantic seaboard of the United States amounted 
to only 373,000 pounds, in 1944 to 501,300 pounds. But Skates are far more plentiful 
than the foregoing might suggest. Average catches of about one Skate to 33 fish of 
all kinds on long lines have been reported at various localities in the Gulf of Maine.*^ 
And an average of perhaps 120—150 Skates per square mile in 37 hauls of an otter 
trawl on Georges Bank (equivalent to about one Skate per four acres of bottom fished) 
probably far understates the actual numbers present, for the trawl, equipped with large 
wooden rollers, doubtless passed over many Skates that were lying on the bottom, while 
others probably escaped the nets as the latter approached them. 

Skates are also abundant from southern New England to New Jersey on sandy 
bottoms. Old records for a pound net on the shore of Nantucket Sound,** operated 
during the season of 1871 chiefly for clupeoids and therefore not particularly well 
placed for ground fish, show that more Skates (2,923) than flatfishes of all kinds com- 
bined (2,714) were taken. A more recent report states that not less than 10,000 pounds 
of Skates, weighing 1-5 pounds, were discarded from one lift of a pound net at Bradleys 

41. On the northeastern coast of the United States, as at Woods Hole, Massachusetts, pieces of squid have often been 
found in Skates' stomachs, although these were probably captured by the latter while in the pound nets or in fish 
traps in which the Skates were taken. According to information furnished by the Bingham Oceanographic Laboratory, 
whole squid have been found in Skates caught in otter trawls. 

42. Steven, J. Mar. biol. Ass. U.K., i8, 1932: 23-24. 43. Steven, J. Mar. bid. Ass. U.K., 18, 1932: 18. 

44. England, Scotland, Eire, France, Germany, Holland, Norway, Sweden, Poland, Iceland. 

45. Bigelow and Welsh, Bull. U. S. Bur. Fish., 40 (i), 1925: 57; Bigelow and Schroeder, Bull. U. S. Bur. Fish., .#<?, 
1936: 324. 

46. At Waquoit, Mass. (Lyman, 6th Rep. Mass. Comm. inl. Fish., 1872: table i). 



Fishes of the Western North Atlantic 145 

Beach, New Jersey." On the other hand, they are so much less abundant on soft bot- 
tom that on one occasion in the Gulf of Maine we took only one Skate for every 250— 
300 fish of other kinds." Unfortunately, there are no statistics available as to the num- 
bers of Skates taken along our Middle Atlantic Coast southward from New Jersey or 
in their southern center of occurrence off southern Brazil, Uruguay, and Argentina. 

Relation to Man. In northern Europe during 1936 and 1937 the average market 
price to the fisherman was about 4-5 cents per pound, the total value of Skates sold 
being about ;^ 1,050,000 and ;^ 92 6,000 or $5,100,000 and $4,500,000, respectively. 
But in North American markets the demand for Skates is very small; according to the 
reported total landings for 1944, only about 501,000 pounds, worth about % 18,000, 
were sold along the Atlantic Coast of the United States (including Chesapeake Bay), 
about 310,000 pounds, worth about $3,000,*" along the Pacific Coast. Small numbers 
have been used from time to time as fertilizer, as fresh manure, and as bait for lobster 
traps along the New England Coast. But the vast majority of those caught along the 
Atlantic coasts of Canada and the United States are thrown back. 

Depth Range. Skates are most abundant in depths less than 100 fathoms, many 
of them occurring plentifully in shallow water, a few right up to the tideline. Stray 
Skates have even been taken in rivers so far in from the mouth that they had doubtless 
been in fresh water for a longer or shorter time.^" But there is no conclusive evidence 
that any member of the genus exists permanently in fresh water. On the other hand, 
several of the species that are common in 30-60 fathoms also range down the slopes 
to depths of 200—300 fathoms or deeper. There is a considerable list of species that 
are known only from depths greater than 100 fathoms, some of these being from depths 
greater than 200 fathoms while others have been recorded from depths greater than 
800-1,500 fathoms. 51 However, we doubt whether any Skate can be classed as a typical 
member of the abyssal fauna, although two {R. abyssicola Gilbert 1893, from the offing 
of British Columbia, and R. badia Garman 1899, from the Gulf of Panama) have been 
reported from depths greater than 1,200 fathoms, knowledge of each being confined 

47. On July 8, 1892 (Smith, Bull. U.S. Fish Comm., 12, 1894: 368). 

48. For details of catches totaling about 11,900 fish of all species, see Bigelow and Schroeder (Biol. Bull. Woods Hole, 
76. 1939: 3^2)- 

49. Statist. Dig. U.S. Fish Wildl. Serv., 16, Fish. Statist. U.S. (1944), 1948. 

50. A large Skate was taken many years ago in the River Ouse near Bedford, England, some 60-70 miles from the 
sea (Pascoe, Zoologist, [3] 7, 1883: 506). More recently one has been reported from the Yangtze in China (Tchang, 
Science Shanghai, 14, 1929: 39S-407). A Skate has been reported also from fresh water of the Rio de la Plata 
(Eigenmann, Rep. Princeton Exped. Patagonia, Zool., 3 [2], 1909: 377; Smith, H. W., Biol. Rev. [Cambridge], 
II, 1936: 11). But the specimen in question [Raja platana Gunther, Challenger Rep., Zool., i, i88o: 12, Chal- 
lenger Sta. 321) was actually taken off Montevideo; i. e., in salt water where the dredge also yielded a rich catch 
of various marine invertebrates. We might point out in passing that Eigenmann's reference of Raja microps Giin- 
ther iSSo (actually referable to the closely related genus Psammobatis) to fresh water rests on the capture of a 
specimen at that same station. 

51. Raja badia Garman 1899, from 1,270 fathoms. Pacific off Panama; R. abyssicola Gilbert 1893, 1,588 fathoms, off 
British Columbia; R.fyllae Lutken 1887, down to 983 fathoms, northeastern Atlantic; R. hyperborea Collett 1878, 
down to 1,309 fathoms, N. Atlantic; R. re-Tiersa Lloyd 1906, 820 fathoms, northern part of Arabian Sea; R. trachura 
Gilbert 1892, S22 fathoms, off southern California; R.j^nst'ni Bigelov/ and Schroeder 1949, 991 and 1,043 fathoms, 
off southern New England; R. bat/typhila Holt and Byrne 1908, 835-1,188 fathoms, continental slope between 
offings of Chesapeake Bay and Nova Scotia; and R. mollis Bigelow and Schroeder 1949, 85S fathoms, off southern 
Nova Scotia. 



146 Memoir Sears Foundation for Marine Research 

to the original capture; it is possible, therefore, that both of them may occur in lesser 
depths as well. 

Range. Skates are most numerous, both as to species and individuals, in warm 
temperate and boreal latitudes along the two sides of the Atlantic and Pacific oceans, 
both north and south, and they occur in some variety in the subarctic belts of both 
hemispheres. A number of species are known also around Cuba, from the warm waters 
of the northern part of the Indian Ocean (including the Gulf of Aden and Arabian 
Sea), the Philippines and East Indies, Formosa, Queensland, the west coast of Central 
America and northern South America. But none seem to have been reported from the 
Micronesian, Polynesian, or Hawaiian Islands in the Pacific or from the American 
side of the tropical Atlantic between Yucatan and mid-Brazil. From the equatorial 
coast of West Africa, anywhere between Cape Verde and Walfish Bay,^^ we find but 
a single record of the genus. And it is hardly conceivable that they would not have been 
reported frequently in scientific literature if they occurred within these extensive areas 
in numbers at all approaching the Skate populations that exist off the coasts of Europe, 
off the eastern United States and Canada, or from southern Brazil southward. 

An interesting aspect of distribution is found in at least four pairs of species; the 
members of each pair are so closely allied that they may prove indistinguishable, yet 
each pair has one member confined to the temperate-boreal latitudes in the northern 
hemisphere while the other is found in the corresponding latitudinal belt in the southern 
hemisphere. These pairs are: R. batis Linnaeus 1758 of the northeastern Atlantic and 
a Skate that has been reported as R. batis from South Africa;^^ R. alba Lacepede 1803 
from the northeastern Atlantic and a form that has been reported from South Africa 
under that same name; R. radiata Donovan 1807 of the northern North Atlantic and 
R. doello-juradoi Pozzi 1935 from Argentina and the Patagonian-Falkland Islands 
region; and R. spinicauda Jensen 19 14 of the subarctic North Atlantic and R. griseo- 
cauda Norman 1937 of the Patagonian-Falkland Islands region. Similar cases of bitem- 
perate, biboreal, or bipolar distribution are known among Sharks of the genera Car- 
charias (Sand Sharks), Lamna (Mackerel Sharks), Cetorhinus (Basking Sharks), Squalus 
(Spiny Dogfishes), and Somniosus (Greenland Shark and a subantarctic counterpart). 

Species. Raja includes a greater number of species by far than any other genus 
of elasmobranchs. The named forms from various parts of the world that are recognized 
in recent synopses of the genus approximate 95—100. While critical comparisons may 
be expected to result in the union of some species that are now regarded as separate,"* 
reductions in the total number from this cause are likely to be more than counter- 
balanced by the ultimate discovery of new species, especially down the slopes of the 
continents. 

The numerical distribution of Raja species as known at the present time is ap- 
proximately as follows: Common to the two sides of the North Atlantic in high latitudes, 

52. /?. maderensis Lowe 1837, reported from Dahomey by Osorio (J. Sci. math. phys. nat. Lisboa, [2] j, 1895: 253). 

53. See Norman (Discovery Rep., J2, 1935: 37) for a recent revision of South African species of Raja. 

54. A comparison of the Skates from the two sides of the northern North Pacific is especially needed. 



Fishes of the IVestern North Atlantic 147 

4; peculiar to the western North Atlantic, south to Yucatan, 16; peculiar to the eastern 
North Atlantic and Mediterranean south to the Canaries, Morocco, and equatorial 
South Africa, i 8 ; common to the western North Atlantic and western South Atlantic, 
poleward from about Lat. 40° S, 2 ; common to the eastern North Atlantic and South 
Africa, 2 ;^^ peculiar to the western South Atlantic, from mid-Brazil southward to Falk- 
land Islands and Straits of Magellan, and southern Chile, 15; South Georgia, i ; South 
African region, y;'^" common to east and west sides of northern North Pacific in high 
latitudes, including Bering Sea, 5 ; Alaska to Lower California, 8 ; Pacific Coast of 
Central America to Ecuador, 3; western North Pacific, Sea of Okhotsk to Formosa 
and southern China, 12; Australia, Tasmania, and New Zealand, 8; East Indies, 
northern Indian Ocean, Arabian Sea, and Gulf of Aden, 6; Kerguelen Island and 
subantarctic Indian Ocean, 2.'^' 

Generally, the specific identification of Skates has been regarded as difficult for 
anyone not thoroughly conversant with the group, primarily because some of the 
characters that have been stressed in published accounts (because the most obvious)^* 
do not lend themselves easily to precise definition or are subject to variation with age 
and sex. Also, a character that is constant in one species may show considerable variation 
in another. But our examination of considerable collections to which we have had access 
has convinced us that the various species differ so definitely from each other — though 
often in less obvious ways — that specific identification calls only for precise observation 
of characters readily seen, felt, measured, or counted. 

Lack of adequate study material from other parts of the world has prevented 
us from extending the accompanying Key to Species (p. 147) beyond the confines of 
the western North Atlantic. On the other hand, it has seemed wise to expand it to 
include local species of Breviraja as well as Raja, because generic identification of one 
or the other is difficult in some cases without x-ray photographs to show the level at 
which the tip of the rostral cartilage terminates relative to the anterior rays of the 
pectorals. A Provisional Key to western South Atlantic Species of Raja is included 
(p. 153) as a convenience. 

Key to Western North Atlantic Species of Raja and Breviraj a^^^ 

I a. Mucous pores on lower surface of disc conspicuously marked by black dots or 
lines (Fig. 47 B). Raja laevis Mitchill 1817, p. 217. 

lb. Mucous pores on lower surface of disc not marked by black dots or lines. 

2a. No conspicuous thorns on tail or on disc posterior to pectoral girdle; rostral 
projection from cranium soft and flexible from base to tip. 

Raja mollis Bigelow and Schroeder 1950, p. 237. 

55. But see p. 146. 56. From recent synopsis by Norman (Discovery Rep., 12, 1935: 38). 

57. R. eatonii Giinther 1S76 and R. murrayi Gunther 1S80. See Norman (Rep. Brit. Antarct. Exped. [1929-1931], 
[B] I [2], 1937: 67; and Discovery Rep., iS, 1938: 100). 

58. Examples are the precise outlines and degree of concavity of the anterior margins of the pectorals, the anterior 
angle of the disc, the length of the tail relative to the body, and the degree of spininess of the disc. 

58 a. Based on specimens examined. 



148 Memoir Sears Foundation for Marine Research 

2 b. Tail or disc posterior to pectoral girdle, or both, with few or many conspicuous 
thorns; rostral projection from cranium firm, whether short or long. 
3 a. Distance from origin of second dorsal to level of axils of pelvics longer 
than distance from level of axils of pelvics to tip of snout by an amount 
at least as great as distance between nostrils, or greater. 
4 a. Either the tail with dark crossbars or the dorsal fins conspicuously 
blackish, or both. 

5a. Upper surface of disc plain-colored; tail without dark crossbars. 
Breviraja atripinna Bigelow and Schroeder 1950, p. 286. 
5b. Upper surface of disc with dark markings; tail with dark 
crossbars. 

6 a. Thorns on shoulders (2—4), along midline of disc, and on 
anterior part of tail, large and conspicuous; anterior margin 
of anterior lobe of pelvics as long as distance from origin 
of pelvics to rear tip, or longer. 

Breviraja plutonia (Garman) 188 i, p. 297. 
6 b. Thorns on shoulders (0—2), along midline of disc, and on 
tail, small and inconspicuous; anterior margin of anterior 
lobe of pelvics shorter than distance from origin of pelvics 
to rear tip. 
Breviraja cubensis Bigelow and Schroeder 1950, p. 293. 
4 b. Neither the tail with definite dark crossbars nor the dorsal fins 
blackish. 
7 a. Midbelt of tail with 3—4 rows of conspicuous thorns. 

Breviraja sinus-mexicanus Bigelow and Schroeder 1950, p. 302. 
7 b. Midbelt of tail with only a single median row of thorns. 

Raja fyl/ae^^ hiitken 1887; newborn, p. 194. 

Raja sema^^ Ga.rma.n 1885; newborn, p. 264. 

3 b. Distance from origin of second dorsal to level of axils of pelvics little if 

any longer than distance from level of axils of pelvics to tip of snout, 

and in most cases considerably shorter. 

8 a. Tip of firm rostral cartilage terminating considerably posterior to tip 
of snout (Fig. 62).''" 

9a. Midzone of disc posterior to scapular region with only one or 
two inconspicuous thorns, if any, in midline. 

Breviraja colesi Bigelow and Schroeder 1948, p. 289. 
9 b. Midzone of disc posterior to scapular region with two to several 
rows of thorns. 
10 a. Snout anterior to orbits not more than about twice as 

59. It may be difficult in some cases to distinguish newborn specimens of K.fyllae and K. senta from each other. 
See under Distinctive Characters, pp. 194-266. 

60. The point at which the rostral cartilage ends can be felt, if not seen, for all the species included under Sa. 



Fishes of the JVestern North Atlantic 149 

long as distance between orbits; first and second dorsal 
fins confluent, without interspace; dorsal surface of disc 
not freckled with many small dark dots. 

Breviraja spinosa Bigelow and Schroeder 1950, p. 306. 
lob. Snout anterior to orbits about ^-Ti times as long as dis- 
tance between orbits; first and second dorsal fins separated 
by a definite (though short) interspace with one thorn; 
upper surface of disc closely freckled with small dark 
brown dots. 

Breviraja yucatanensis Bigelow and Schroeder 1950, p. 310. 
8 b. Firm rostral cartilage reaching nearly to extreme tip of snout. 
1 1 a. No large thorns on posterior V4~V3 of t^'l- 

Raja senta Garman 1885, p. 264.*' 
I lb. One or more rows of thorns along posterior part of tail as well 
as farther forward along it. 

12 a. No large thorns anywhere on disc between nuchal region 
and level of axils of pectorals, except for alar spines of 
sexually mature males. 

1 3 a. Upper surface of pectorals smooth, without prickles. 
14 a. First and second dorsals confluent. 

Raja teevani Bigelow and Schroeder 1951, 

p. 276. 
14 b. First and second dorsals separated by a 
definite interspace. 

15 a. Lower surface with a band of prickles 
along anterior edge of disc and along 
rostral cartilage; distance between dor- 
sals about as long as base of first dorsal ; 
outer margin of nostril lobe fringed. 
Raja olseni Bigelow and Schroeder 
1951, p. 251. 
15b. Lower surface smooth everywhere; dis- 
tance between dorsals less than half as 
long as base of first dorsal ; outer mar- 
gin of nostril lobe smooth. 

Raja laevis Mitchill I 8 1 7 ; 
newly hatched, p. 217. 
1 3 b. Upper surface of pectorals rough with close-set 
prickles except along outer margins. 

Raja spinicauda Jensen 19 14, p. 271. 

61. Newly born R. senta may have thorns on the posterior part of the tail, and thus fail under alternative 7b; see p. 266, 
footnote 138. 



150 Memoir Sears Foundation for Marine Research 

12 b. Disc between nuchal region and level of axils of pectorals 
with more or fewer large thorns in addition to alar spines 
of mature males. 

1 6 a. Distance from origin of first dorsal to axils of pel- 
vies at least as long as distance from axils of pelvics 
to fronts of orbits, or longer. 
17 a. Lower surface of disc and tail uniform brown, 
darker than upper surface. 
Raja hathyphila Holt and Byrne 1908, p. 159. 
17 b. Lower surface of disc and of tail paler than 
upper surface, either uniformly whitish, gray, 
or fawn-colored, or with dusky markings. 
I 8 a. First and second dorsals confluent. 

Raja fyllae Liitken i 887; half-grown 

and adults, p. 194. 

1 8 b. First and second dorsals separated by 

a short but definite interspace. 

19 a. Upper surface of disc densely 

though irregularly strewn with 

small dark freckles. 

Raja lentiginosa Bigelow and 
Schroeder 1951, p. 228. 
19 b. Upper surface of disc with dark 
freckles mostly concentrated in a 
few conspicuous rosettes. 
Raja garmani^\i\\\^'S[ 1939, p. 200. 
1 6 b. Distance from origin of first dorsal to axils of pelvics 
hardly longer than distance from axils of pelvics to 
rear margins of orbits, and in most cases consider- 
ably shorter. *- 

20a. Only one row of large thorns along tail; if 
more than one, those of median row much 
larger and more conspicuous than the others. 
2 1 a. Not more than 1 9 large thorns along 
midline of back and tail between nuchal 
region and first dorsal fin. 
Raja radiata Donovan 1807, p. 255. 
2 lb. At least 24 large thorns along midline 
of back and tail between nuchal region 
and first dorsal fin. 

62. On some very young specimens of Raja erinacea the distance from the axils of the pelvics to the origin of the first 
dorsal fin is a little longer than the distance from the axils of the pelvics to the rear margins of the orbits (Fig. 37). 



Fishes of the JVestern North Atlantic 151 

22 a. Tail posterior to tips of pelvics 
with a lateral row of smaller 
thorns on either side in addition 
to the larger median row; at least 
34 large thorns in young and up 
to 50 or more in adults in median 
row from nuchal region to first 
dorsal fin. 

Raja Untea Fries 1838, p. 232. 
22 b. Tail with only one (the median) 
row of large thorns; not more 
than 26-31 thorns in median 
row from nuchal region to first 
dorsal fin. 

23 a. Upper teeth in only about 
44 series; adults with large 
thorns on posterior parts of 
pectorals and on outer an- 
terior parts of disc. 

Raja hyperborea Collett 
1878, p. 206. 
23 b. Upper teeth in at least c,c^ 
series; adults without large 
thorns on posterior parts of 
pectorals or on outer anter- 
ior parts of disc. 

Raja jenseni Bigelow and 

Schroeder 1950, p. 213. 

20b. More than one row of large thorns along tail ; 

no one row much more conspicuous than the 

others. 

24a. Three or more rows of thorns along 
midbelt of disc between shoulders and 
level of axils of pectorals; first and 
second dorsal fins confluent or only nar- 
rowly separated, without intervening 
thorns. 

25a. Upper teeth in at least 72*^^ series, 
most often 90-100; does not 

63. Bigelow and Schroeder (Bull. U. S. Bur. Fish., 48, 1936: 324) gave as few as 70. While re-examination of the par- 
ticular specimens on which this count was based showed that the number should have been stated as 80, W. Temple- 
man reports to us the capture of one from the Grand Bank of Newfoundland with only !jj teeth. 



152 Memoir Sears Foundation for Marine Research. 



mature sexually until at least 26 
inches long. 

Raja ocellata Mitchill 18 15, 
p. 240. 

25 b. Upper teeth in not more than 66 

series, usually less than 54 ; ma- 
tures when only 18—20 inches 
long. 

Raja erinacea Mitchill 1825, 
p. 176. 
24b. Only one row of thorns along midbelt 
of disc between shoulders and level of 
axils of pectorals; first and second dor- 
sal fins separated by a definite inter- 
space or by one or more thorns. 

26 a. Upper surface of each pectoral 

marked with a conspicuous dark- 
centered ocellar spot, much larger 
than any other of the dark mark- 
ings. 

27 a. Lateral margins of snout 
rather deeply concave (Fig. 
60); horizontal diameter of 
eye only about Vs as long as 
snout anterior to orbits. 
Raja texana Chandler 1921, 
p. 279. 
27 b. Lateral margins of snout 
hardly concave; horizontal 
diameter of eye about V3 as 
long as snout anterior to 
orbits. 
Raja ackleyi Garman i 8 8 i , 

P- 155- 
26 b. Upper surface of each pectoral not 
marked with a large and prom- 
inent ocellar spot; surface of disc 
as a whole with many irregular 
dark dots and narrow bars, trans- 
verse or oblique. 
Raja eglanteria Lacepede 1802, 
p. 165. 



Fishes of the JV ester ?i North Atlantic 153 

Provisional Key to Western South Atlantic Species of Rcija^'^ 

I a. Distance from origin of first dorsal fin to tip of tail as long as distance from origin 
of first dorsal to tips of pelvic fins, or longer. 

agassizii Miiller and Henle 1841. 
Rio de Janeiro. 
I b. Distance from origin of first dorsal fin to tip of tail considerably shorter than 
distance from origin of first dorsal to tips of pelvic fins. 

2 a. Lower surface of disc with many conspicuous black dots and streaks marking 
the mucous pores. 

3 a. Inner part of each pectoral fin marked with one large and conspicuous 
pale-centered ocellar spot. cyclophora Regan 1903. 

Rio de Janeiro. 
3 b. Each pectoral not marked with a large ocellar spot, or, if present, in- 
distinct. 

4 a. Distance from axils of pelvics to rear end of second dorsal fin only 
about Vs as long as distance from axils of pelvics to rear margins of 
orbits; distance from origin of first dorsal to tip of tail considerably 
shorter than that from level of rear margins of orbits to tip of snout; 
mucous pores on upper surface of anterior part of disc not marked 
by black dots or streaks. flavirostris Philippi 1892. 

Patagonian-Falkland Islands re- 
gion, Straits of Magellan, and 
Chile. 
4 b. Distance from axils of pelvics to rear end of second dorsal as long 
as distance from axils of pelvics to rear margins of spiracles, or 
longer; distance from origin of first dorsal to tip of tail about as long 
as distance from level of rear margins of orbits to tip of snout; 
upper surface of anterior part of disc with black spots and streaks, 
marking mucous pores. 

5a. Tail with three rows of thorns; margins of orbits and scapular 
regions also with thorns; snout anterior to orbits only about as 
long as distance between outer margins of orbits. 

castelnaui Ribeiro 1904. 
Vicinity of Rio de Janeiro. 
5b. Tail with only one row of thorns; no thorns around orbits or 
on scapular regions; snout anterior to orbits about 1.5 times as 
long as distance between outer margins of orbits. 

64. See Ribeiro (Arch. Mus. nac. Rio de J., 14, 1907: 177-179; Fauna brasil., Peixes, 2 [i] Fasc. i, 1923: 33-36) 
for descriptions and illustrations of Brazilian species and Norman (Discovery Rep., 16, 1937: 12-47) for those of 
the Patagonian-Falkland Islands region and Straits of Magellan. The Brazihan Skate reported by Ribeiro, first 
as Kaja erinacea Mitchill (Arch. Mus. Nac. Rio de J., 14, 1907: 176) and subsequently as R. cirrifera Regan 
(Fauna brasil. Peixes, 2 [i], Fasc. i, 1923: 33), and by Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 356) 
as a new species, R. extenia, is a Psammobatis (see Norman, Discovery Rep., 16, 1936: 28). 



154 Mejnoir Sears Foundation for Marine Research 

■platana Giinther 1880. 
Rio de Janeiro'^ and mouth 
of La Plata River. 
2 b. Lower surface of disc not marked with black dots or streaks. 

6 a. No large thorns anywhere on disc anterior to level of axils of pectorals. 
7 a. Distance from cloaca to tip of tail shorter than that from cloaca to 
tip of snout; outer posterior parts of pectorals mostly smooth. 

scaphiops Norman 1937. 
North of Falkland Islands. 

7 b. Distance from cloaca to tip of tail longer than distance from cloaca 

to tip of snout; outer posterior parts of pectorals with extensive 
prickly areas. griseocauda Norman 1937. 

Patagonian-Falklands region. 
6 b. Disc anterior to level of axils of pectorals with conspicuous thorns either 
along midline, around orbits, on scapular regions, or on all three parts. 

8 a. No thorns on scapular regions. 

9 a. Tail with only one row of thorns. 

loa. Snout in front of orbits at least four times as long as 
distance between orbits; anterior margins of disc deeply 
concave. echinorhyncha Ribeiro 1923. 

Rio de Janeiro, 
lob. Snout in front of orbits not more than 2.5 times as long 
as distance between orbits; anterior margins of disc not 
deeply concave. 

1 1 a. Disc with many small rounded white spots; thorns 
on tail noticeably larger than those along mid-dorsal 
line on disc. albomaculata Norman 1937. 

Patagonian-Falklands region. 
lib. Disc not marked with many small white spots; 
thorns on tail not larger than those on midline of disc. 
brachyurops Fowler 19 10. 
Northern Argentina (.'); Pata- 
gonian-Falklands region. 
9 b. Tail with two lateral rows of thorns along each side in addition 
to median row. brasiliensis M tiller and Henle 1841. 

Brazil. 
8 b. Each scapular region with 1—3 thorns. 

12 a. Only 12 — 15 thorns along midline of back and tail trom nuchal 
region to first dorsal fin, all large. 

doello-juradoi Pozzi 1935. 
Northern Argentina to Patago- 
nian-Falklands region. 

65. On re-examination, the specimen from Rio de Janeiro described by Garman (Mem. Harv. Mus. comp. Zool., 
36, 191 3: 361) under the name R. castelnaui Ribeiro 1904 proves not to be that species; it agrees closely with Gunther's 
account and illustration (Challenger Rep., Zool., i [6], 1880: 11, pi. 3) of R. platana. 



Fishes of the Western Nort/i Atlantic 155 

12 b. 25—42 moderate-sized thorns along midline of back, and tail 
from nuchal region to first dorsal. 

13a. No large thorns on margins of orbits; about 42 thorns 
in mid-dorsal row. muhispinis Norman 1937. 

Northwest of Falkland Islands. 
13 b. One thorn at anterior margin and one at posterior margin 
of each orbit; not more than about 30 thorns in mid- 
dorsal row. 

14a. Inner parts of pectorals roughened with small 
scattered prickles; only one thorn on each scapular 
region. macloviana'Hovvm.n I93 7" 

Patagonian-Falklands region.'® 
14b. Inner parts of pectorals mostly smooth; each scap- 
ular region usually with two thorns. 

magellanica Steindachner 1903. 
Patagonian-Falklands region, and Straits 
of Magellan. 



Raja ackleyi Garman i 8 8 i 

Ackley's Skate 

Figure 29 

Study Material. Two specimens, a male 410 mm long, the type (Harv. Mus. Comp. 
Zool., No. 748) and a female 231 mm long (U. S. Nat. Mus., No. 148290), both taken 
off Yucatan. 

Distinctive Characters. R. ackleyi appears to be separated from R. texana in that 
the outer margins of its disc are broadly rounded (rather abrupt in R. texand)\ also, on 
the average the former has a narrower disc, i.i — 1.2 times as broad as long (1.2— 1.35 
times for texand)\ and its disc is sprinkled with small light and dark spots with the 
ocellar spots*' oval (disc plain brownish with round ocellar spots on R. texana).^^ The 
color pattern in itself is enough to set R. ackleyi (and R. texana") apart from all other 
Skates of the western North Atlantic, except for specimens of R. ocellata on which 
there is a single large ocellar spot on each side and on which the other dark markings 
are not conspicuous. The disc and tail of i?. ocellata are so much thornier and the anterior 
contour is so much more obtusely rounded than those of R. ackleyi that the two could 
hardly be confused. R. cyclophora Regan 1903,*' from Brazil, closely parallels R. ackleyi 
(and R. texana) in the presence of a single conspicuous ocellar spot on the posterior 

66. R. maclo'viana Norman 1937 seems so closely allied to R. magellanica Steindachner 1903 that it may prove to re- 
present a variety of the latter. 

67. In R. ackleyi the distance from the center of the ocellus to the center of the orbit is about i. 0—1.3 times the distance 
between the centers of the ocelli; in four specimens of R. texana, this distance was 0.83—1.0 times. 

68. If the two species are finally united, the name ackleyi has priority. 

69. For an excellent photograph of R. cyclophora, see Ribeiro (Arch. Mus. nac. Rio de J., 14, 1907: pi. 16). 




Figure 29. Rtija ackleyi, adult male, 410 mm long, from Yucatdn Bank (Harv. Mus. Comp. Zool., No. 748, 
type). A Ventral view of anterior part of head. B Posterior margin of nasal curtain, about 3.4X . C Outer posterior 
margin of nostril, about 3-4 X. D Upper teeth, near middle of mouth, about 12 X- 



Fishes of the Western North Atlantic 157 

part of each pectoral. But the pectorals of R. cyclophora are angular, hence they con- 
trast sharply with the rounded outer contours of R. ackleyi. 

Description. Proportional dimensions in per cent of total length. Female, 231 mm, 
from off Yucatan (U. S. Nat. Mus., No. 148290). Male, 410 mm, from Yucatan Bank 
(Harv. Mus. Comp. Zool., type, No. 748). 

Disc: extreme breadth 63.0, 53.7; length ^2)-Si 48.0. 

Snout length: in front of orbits 15. i, 11.6; in front of mouth 17.7, 12.7. 

Orbits: horizontal diameter 4.3, 3.7; distance between 4.6, 3.9. 

Spiracles: length 2.6, 2.4; distance between 7.0, 5.8. 

Mouth: breadth 7.8, 7.5. 

Nostrils: distance between inner ends 8.7, 7.5. 

Gill openings: lengths, ist 1.9, 1.7; 3rd 1.9, 1.8; 5th 1.3, 1.2; distance between 
inner ends, ist 15.4, 13.0; 5th 8.2, 7.3. 

First dorsal fin: vertical height 3.9, 2.2; length of base 7.7, 4.6. 

Pelvics: anterior margin 9.5, 10.4. 

Distance: from tip of snout to center of cloaca 48.2, 45.8 ; from center of cloaca 
to 1st dorsal 31.6, 40.7; to tip of tail 51.8, 54.2. 

Interspace between: ist and 2nd dorsals 4.1, 2.3. 

Disc about 1.1-1.2 times as broad as long; maximum anterior angle in front of 
spiracles 97° on male, 106" on female; anterior rays of pectorals extending about 60- 
70 "/o of distance from level of front of orbits toward tip of snout, the latter not ap- 
preciably projecting; anterior contour only weakly concave rearward from tip of snout 
and then a little more deeply so opposite spiracles on male, but less so on female; outer 
and posterior corners broadly rounded; posterior margins rather strongly convex. Tail 
with a narrow fold beginning about an eye's length posterior to axil of pelvic on each 
side and extending to tip; length of tail from center of cloaca to origin of first dorsal 
0.7—0.9 times, to origin of second dorsal 0.9—1.0 times, and to tip i.i — 1.2 times as 
great as distance from center of cloaca to tip of snout. 

A cluster of small thorns on tip of snout; an irregular patch in front of spiracle 
with three or four along its inner edge; one thorn on each shoulder on larger (male) 
specimen (not yet developed on smaller female); one or two rows of small thorns over 
anterior part of rostral cartilage; inner edge of orbit with three thorns on smaller 
specimen, the number increasing to about eight on mature male; midline of back 
from nuchal region to first dorsal of smaller specimen with 27 thorns, the row inter- 
rupted at first between scapular region and level of axils of pectorals but becoming 
continuous later; thorns of midline increasing to 70—72 at maturity (about 20 anterior 
to axils of pectorals), the larger roughly alternating with the smaller, decreasing generally 
in size rearward; on partly grown specimens, each side of tail with one additional 
irregular row of thorns of various sizes extending rearward from near axil of pelvic 
to opposite second dorsal; large specimens with two such rows on each side of tail, 
the uppermost row (developed the latest) extending from a little anterior to axils of 



158 Memoir Sears Foundation for Marine Research 

pectorals back to first dorsal and followed by prickles as far as second dorsal; skin of 
disc and tail naked otherwise except for a few minute prickles opposite orbits and 
spiracles and along edge of disc on male abreast of alar spines. Adult male with two rows 
of alar spines;'" also a patch of about 18 thorns on either side in malar region abreast 
of eye and'spiracle. Lower surface of larger specimen (male) prickly from tip of snout 
rearward to a little beyond midlevel of nostrils and along margins of disc to level of 
mouth, but otherwise naked on disc and tail; smaller specimen (female) smooth every- 
where below. 

Snout in front of orbits about 3.0-3.5 times as long as distance between orbits; 
about 1.6—2.0 times as long in front of mouth as distance between exposed nostrils. 
Orbit about 1.5 times as long as spiracle; distance between orbits a little greater than 
length of orbit. Length of first gill openings about V4 as great as width of mouth, the 
fifth gill openings about 70 "/o as long as the first; distance between first gills 1.5-1.8 
times as long as distance between exposed nostrils; distance between fifth gills about 
as great as distance between nostrils. Both nasal curtain and expanded posterior (outer) 
margin of nostril fringed. Mouth a little bowed in adult male, less so in female. 

Teeth ^ in adult male, in transverse series, with thorn-like and somewhat recurved 
cusp on circular or oval base, the uppers and lowers alike; ^ series in female, arranged 
in quincunx, the median teeth with low cusp, the outer teeth without cusp. 

Dorsal fins about equal in size, with convex anterior margin and well rounded 
apex; base of first dorsal about 87 "/o as long as distance between spiracles. Interspace 
between first and second dorsals about V2-V3 (50-66 "/o) as long as base of first dorsal. 
Pelvlcs moderately concave outwardly, weakly scalloped; anterior margin about ^/g as 
long as distance from pelvic origin to rear tip; anterior lobe narrow, outer margin of 
posterior lobe weakly convex, rear tip well rounded even In adult male, reaching back 
about V3 the distance from origin of pelvic to tip of tail. Claspers of adult male reaching 
a little more than halfway from tip of pelvic toward first dorsal fin. 

Anterior rays of pectorals reaching about ^/j the distance from level of fronts of 
orbits toward tip of snout. 

Color. Upper surface pale yellowish brown, sprinkled with small indefinitely out- 
lined darker and lighter spots and pale patches; a dark oval ocellar spot with pale 
margin on inner part of each pectoral a little posterior to axis of greatest breadth. Lower 
surface white without dark markings. 

Size. The type specimen, a male 410 mm long, appears to be mature sexually. 

Developmental Stages. The egg cases have not been seen. 

Habits. Nothing is known. 

Range. So far known with certainty only from the Yucatan Bank and near there 
in Lat. 22°o8' N, Long. 86°53' W, and from southern Florida." Skates have also 

70. On this particular specimen there are five alar spines in the outer row on the left-hand side but only two in the 
outer row on the right-hand side. 

71. A third specimen of R. ackleyi was sent to us for identification by Stewart Springer. It was a female, 404 mm 
long, caught off the Dry Tortugas in Lat. 25°03' N, Long. 82^56' W, in 25 fathoms, January 19, ig^i, 
Oregon St. 233. 



Fishes of the JVeste7^?i North Atlantic 159 

been reported under this same name from tropical West Africa" and from the Azo- 
res,'^^ but they differed from the typical ackleyi in dermal armature and in plain colora- 
tion without ocellar spots. 

Synonyms and References: 

Raja''^ ackleyi Garman, Bull. Mus. comp. Zool. Harv., 8, 1881: 234 (descr., Yucatan Bank); Goode and 
Bean, Smithson. Contr. KnowL, jo, 1895: 25; also Mem. Harv. Mus. comp. Zool., 22, 1896: 25 (descr., 
after Garman, 1881; Yucatan Bank); Jordan and Evermann, Bull. U. S. nat. Mus., 4J (i), 1896: 70 
(descr., Yucatan Bank); Rep. U. S. Comm. Fish. (1895), 1896: 221 (listed, Yucatan Bank); Engel- 
hardt, Abh. bayer. Akad. Wiss., Suppl. 4 (3), 1913: 102 (range); Garman, Mem. Harv. Mus. comp. 
Zool., j6, 1913: 336, pi. 19 (descr., ill., Yucatan Bank); Jordan, Evermann and Clark, Rep. U. S. 
Comm. Fish. (1928), 2, 1930: 25 (listed, Yucatan Bank); Beltran, List. Feces Mexicanos, 1934: 5 
(listed, Mexico; not seen). 

Raja ackleyi (in part) Fowler, Bull. Amer. Mus. nat. Hist., 70(1), 1936: 107 (descr., after Garman; but ill. 
after Goode and Bean and ref. to Azores from Roule, 191 2, probably are not for ackleyi^. 

Doubtful References: 

Raia''* ackleyi Goode and Bean, Smithson. Contr. Knowl., jr, 1895: pi. 7, fig. 23; also Mem. Harv. Mus. 
comp. Zool., 22, 1896: pi. 7, fig. 23 (ident. doubtful); Jordan and Evermann, Bull. U. S. nat. Mus., 
^7 (4), 1900: pi. 10, fig. 31 (same ill. as Goode and Bean, 1895: pi. 7, fig. 23); Roule, Bull. Inst, oceanogr. 
Monaco, 254, 1912: 20 (probably not R. ackleyi because dermal armature diif. and color plain; Azores). 



Raja bathyphila Holt and Byrne 1908 
Abyssal Skate 
Figures 30, 31 

Study Material. Two females and three males, i 17—467 mm long, from the lower 
part of the continental slope between the offings of Chesapeake Bay and southern Nova 
Scotia, at 835-1,188 fathoms, in U. S. National Museum." 

Distinctive Characters. The most convenient field mark for this little known species, 
and one that separates it from all other Skates known from the North Atlantic except 
R. olseni, is the dark brown color of the lower surface of the disc, which is darker 
than the upper surface. R. bathyphila has been classed (incorrectly in our opinion) 
as a synonym of the young of R. lintea (p. 165), but there is no danger of confusing 
halfgrown or larger specimens of the two species." The anterior contour of the disc 
is noticeably diff"erent (cf. Figs. 30, 31 with 49); the tail oi R. bathyphila is much longer 
relatively, as are the anterior margins of its pelvic fins; the upper surface of its disc is 
much more prickly generally; and there are only about six large mid-dorsal thorns 
from pectoral girdle to pelvic girdle on R. bathyphila (about twice that many on R. lintea). 
Also, the tail of R. bathyphila is thornier, and while it is the sides of the tail low down 

72. Poll (Result. Sci. Exped. oceanogr. Beige Cot. Afr. Atlant. [1948-49], 1951: 144-149. 

72a. Roule, Bull. Inst, oceanogr. Monaco, 243, 1912: 20. 73. Also spelled Rata. 74. Also spelled Raja. 

75. One specimen each at the following Albatross Sts.: 2206, Lat. 39°35' N, Long. 7i°25' W, 1,043 fathoms; 2210, 
Lat. 39°38' N, Long. 7i°i9' W, 991 fathoms; 2691, Lat. 39°37' N, Long. 7i°o8' W, 835 fathoms; 2706, Lat. 4i°29' N, 
Long. 65°36' W, 1,188 fathoms; and 2728, Lat. 36°3o' N, Long. 74°33' W, 859 fathoms. 

76. Very young R. lintea of proven parentage have not been seen, so far as we can learn. 




Figure 30. Raja batfiyphi/a, female, 463 mm long, from lower part of continental slope off southern New Eng- 
land, Lat. 3g°38' N, Long. 7i°i9' W, 991 fathoms (U. S. Nat. Mus., No. 35591). ^ Lower surface of ante- 
rior part of head with nostrils and mouth. B Ventral view of pelvics and base of tail. C Posterior margin of nasal 
curtain, about 3.5 X. D Left nostril, about 3.5 X. E Upper teeth, about 6.5 X. 



Fishes of the Western North Atlantic 1 6 1 

that are prickly (in addition to the large thorns) on R. bathyphila^ it is the upper surface 
of the tail that is prickly on R. lintea. The lower surface of R. lintea, though it often 
exhibits dark markings, seems never to be uniformly dark chocolate as on R. bathy- 
phihiy except perhaps on some very small specimens. 

Actually, R. bathyphila more nearly resembles R.fyllae, apart from color, but its 
snout is more acute and relatively longer than that of R./yl/iie; it has fewer large thorns 
along the midzone of its disc rearward from the pectoral girdle; and the bands of prickles 
along the sides of its tail are much denser and more conspicuous. 

Among Skates of the Pacific Coast of America, R. bathyphila parallels R. badia 
Garman 1899 from the Gulf of Panama, R. trachura Gilbert 1892 from California to 
Alaska, and R. abyssicola Gilbert 1895 from off British Columbia in the uniformly dark 
color of its lower surface and in its abyssal habitat. But the shape of its disc marks it 
off from R. badia (outer corners angular in badia and axis of greatest breadth far rear- 
ward), and the presence of conspicuous thorns around its eyes and on its scapular 
regions separates it just as sharply from both R. trachura and R. abyssicola. R. bathy- 
phila has been characterized as "allied to Rata isotrachys^ Giinther 1877," which was 
taken off" Japan by the Challenger in 365 fathoms and which is brownish black below. 
But the original illustration of the unique specimen of R. isotrachys pictures it as re- 
sembling R. badia more nearly than R. bathyphila in shape and as lacking large thorns 
on the disc.'* Among the other deep-water Skates of the genus Raja from the western 
Pacific-Indian ocean regions that are wholly or largely dark-colored below," R. bathy- 
phila seems closest to R. mamillidens Alcock 1889 from the Gulf of Manaar in having 
a small number of teeth, a generally prickly disc, and large thorns around the orbits, 
on the scapular regions, and along the midline of the back from the nuchal region 
posteriorly. But the snout of R. bathyphila is relatively much longer and the disc rela- 
tively wider than these characters in R. mamillidens,^'^ while the teeth of the latter are 
cuspidate even on small females (rounded in R. bathyphihi) and its color uniformly 
black below as well as above. 

Description. Proportional dimensions in per cent of total length. Male, 328 mm 
long, from Lat. 4i°28' N, Long. 65°3o' W (U. S. Nat. Mus., No. 148276). Female, 
467 mm long, from Lat. 39°38' N, Long. 7i°i9' W (U. S. Nat. Mus., No. 35591). 

Disc: extreme breadth 49.4, 52.0; length 43.3, 44.9. 

Snout length: in front of orbits 11.3, 12.2; in front of mouth 12.8, 12.4. 

Orbits: horizontal diameter 3.6, ^t-^'-, distance between 3.3, 3.2. 

Spiracles: length 2.4, 2.6; distance between 6.4, 6.3. 

Mouth: breadth 4.9, 5.9. 

Nostrils: distance between inner ends 6.4, 6.1. 

77. By Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 329. 

78. Gunther, Challenger Rep., 22, 1877: pi. 3. 

79. For a recent survey of Indo-Pacific rajids, see Fowler (Bull. U. S. nat. Mus., 100 [/j], 1941 : 357). 

80. Length of snout in front of nostrils described as being only about as great as distance between nostrils for R. mamil- 
lidens, the disc only about as broad as long (Alcock, Ann. Mag. nat. Hist., [6] 4, 18S9: 380; 111. Zool. Investigator, 
Fishes, 1892: pi. 8, fig. i). 



1 6 2 Memoir Sears Foundatio?2 for Marine Research 




Figure 3 1 . Raja bathyphila. A Immature male, about 328 mm long, from off southern Nova Scotia, Lat. 4i°29'N, 
Long. 65°36' W, 1,188 fathoms (U. S.Nat. Mus., No. 148276). B Newly hatched male, about 117 mm long, 
from ofFNew Jersey, Lat. 39°37' N, Long. 7i°o8' W, 835 fathoms (U. S. Nat. Mus., No. 148269). 

Gill openings: lengths, ist i.i, 1.3; 3rd 1.2, 1.5; 5th 1.2, 1.3; distance between 
inner ends, ist 12.8, 13.2; 5th 8.2, 8.4. 



Fishes of tJie JVestern North Atlantic 163 

First dorsal fin: vertical height 2.3, 2.4; length of base 5.2, 4.9. 

Second dorsal fin: vertical height 2.1, 2.1; length of base 5.8, 4.6. 

Pelvics: anterior margin ii.o, 12.2. 

Distance: from tip of snout to center of cloaca 41.5, 42.2 ; from center of cloaca 

to 1st dorsal 44.6, 45.0; to tip of tail 58.5, 57.8; from rear end of 2nd 

dorsal base to tip of caudal 3.3, 3.4. 
Interspace between: ist and 2nd dorsals 0.0, 0.0. 

Disc about i.i times as broad as long; maximum anterior angle in front of spir- 
acles about 90° on smaller specimens but about 105° on larger; tip of snout rounded, 
not noticeably projecting; anterior margins nearly straight or slightly sinuous, the outer 
and posterior corners broadly rounded, the posterior margin weakly convex. Axis of 
greatest breadth about 77— 78''/o of distance rearward from tip of snout toward axils 
of pectorals. Tail with narrow lateral folds along its posterior two-fifths, widest oppo- 
site pelvics; its length from center of cloaca about i.i — 1.2 times as great to first dorsal 
and 1. 4-1. 5 times as great to tip of tail as distance from center of cloaca to tip of snout. 

Dorsal surface of disc on larger specimens with 2,-6 medium to large thorns 
on radiate bases close behind tip of snout; several smaller ones along rostral cartilage; 
a row of 5—6 large thorns around inner margin of orbit, preceded by a few smaller 
ones close in front of eye; 1—2 larger thorns inward from inner end of spiracle; a 
group of up to 12 — 14 o" nuchal region, small to large and irregularly distributed; 
three large thorns on each shoulder region, arranged like the mathematical symbol for 
the word therefore (.•.), with a few smaller ones between these and the median line; 
also a median row of 30—35 from nuchal region to first dorsal (six from pectoral 
girdle to level of axils of pectorals), widely spaced anterior to level of axils of pectorals 
but closer together and decreasing a little in size toward first dorsal; tail with one to 
two irregular rows along either side in addition to the median row, the row lowest 
down on either side continuing about halfway along base of second dorsal. In addition 
to the larger thorns, the entire upper surface of the disc, including the skin above the 
eyes, is strewn with numerous smaller prickles on small specimens, except for naked 
areas along the posterior margins, and on the tip of the snout which continue so up 
to a length of 350 mm or more. With continued growth these marginal naked bands 
widen and additional naked areas develop on either side posterior to the scapular region 
as well as on either side of the rostral ridge, while the skin above the eyes loses most 
of its prickles; also, the prickles are relatively smaller and more closely spaced than on 
smaller specimens (illustrated by female 463 mm long, Fig. 30). Each side of the tail 
below the outermost row of thorns has a narrow band of close-set prickles along the 
anterior half, but thence rearward it is only sparsely prickly. And the prickles in this 
band are relatively larger and more thorn-like on younger Skates than on older ones. 
The dorsals of the smallest specimens are smooth, their upper parts are prickly on 
somewhat larger ones (316 and 350 mm); but on the largest ones seen there are only 
a few prickles. Caudal membrane smooth. The pelvics of some small specimens are 
prickly on the central part of the posterior lobe, but they are smooth on most, whether 



1 64 Memoir Sears Foundation for Marine Research 

larger or smaller.*' Lower surface smooth, except along anterior part of tail where the 
lateral bands of prickles encroach somewhat on the lower surface. 

A comparison of larger with smaller specimens shows that the total number of 
large thorns (28-35) '" ^^^^ median row is established early, there being six from the 
pectoral girdle to the level of the axils of the pectorals at the smallest size. There is 
only one row along the tail on the smallest seen, but the lateral rows develop along 
the posterior third when a length of about 300 mm is reached. These rows have ex- 
tended forward to the level of the tips of the pelvics at a length of about 350 mm. 
The thorns on the snout (lacking on smallest seen) are present at a length of 3 i 6 mm; 
at first there are only two orbital thorns and one scapular on each side.*^ 

Snout in front of orbits about 3.0-3.15 times as long as orbit, its length in front 
of mouth about twice as great as distance between exposed nostrils. Orbit about as 
long as distance between orbits and about i .4 times as long as spiracle. First gill open- 
ings about 22-23 °/o as long as breadth of mouth; distance between inner ends of 
first gills about 2.0-2.2 times as long as distance between exposed nostrils, about 
1.3— 1.4 times between inner ends of fifth. Nasal curtain and expanded posterior 
(outer) margin of nostrils deeply fringed. Mouth, on females and immature males, a 
little arched forward centrally; its shape on mature males not known. 

Teeth in 34-42 series in upper jaw, those of females close-set in quincunx, with 
moderately high conical cusp; those of mature males not seen. 

Dorsals alike in shape and about equal in size, their bases confluent without 
intervening thorn. Caudal membrane, posterior to second dorsal, about 80 "/„ as long 
as base of second dorsal. Pelvics deeply concave outwardly, their anterior margin about 
63-70 "/o as long as distance from origin of pelvic to rear tip; anterior lobe slender, 
with three radials besides the first stout one and with rounded tip and deeply scalloped 
rear margin; posterior lobe strongly and evenly convex, scalloped anteriorly but only 
faintly wavy posteriorly; rear tip abrupt, extending about V4 of the distance from level 
of axils of pectorals toward first dorsal. 

Anterior rays of pectorals extending forward about -\^ of distance from level of 
fronts of orbits toward tip of snout. 

Color. Upper surface uniform grayish brown with greenish tinge (after many years 
in alcohol), without definite markings; outer edges of pectorals and pelvics shading to 
darker. Lower surface of disc and pelvics uniform chocolate brown, except for paleness 
on jaws; lower surface darker than upper. Smaller specimens similar except for paleness 
close over pectoral girdle; lower surface of tail paler chocolate brown. *^ 

81. The pelvics on a female about 357 mm long are prickly as above but they are smooth on all the others we have seen. 

82. There appears to be considerable variation, whether individual or geographic, in the stage at which the additional 
large orbital and scapular thorns first appear. A male of 117 mm has two oculars and one scapular on each side; 
an eastern Atlantic specimen of 184 mm (Holt and Byrne, Fish. Ireland Sci. Invest. [1906], 5, 1908: 57) had three 
oculars and two scapulars on each side; a male, 316 mm, two oculars and two scapulars, one smaller, on each side; 
a female about 350 mm (tip of tail damaged), three oculars on each side, two scapulars on one side and three on the 
other; a female 463 mm (largest seen), five and six oculars, and three scapulars on each side. 

83. The lower surface of the type specimen from the Irish Atlantic slope is described as "brown, except the front of 
the snout, mouth parts and belly" (Holt and Byrne, Fish. Ireland Sci. Invest. [1906], 5, 1908: 53). 



Fishes of the If'^estern North Atlantic 165 

Remarks. The smaller specimens in our Study Material agree so closely (bodily 
proportions, dermal armature of disc and tail, and color) with the description** of the 
only specimen of R. bathyphila recorded previously that there seems to be no justifica- 
tion for separating them from that species unless future study should show that sub- 
sequent development follows a different course in the eastern Atlantic form from that 
illustrated by successive stages in the growth of the western Atlantic form. Thus we 
conclude that R. bathyphila is a well marked species rather than a young stage o( R. lintea 
as has been suggested.*^ 

Si-ze. The young are hatched at a length less than i 1 7 mm (our smallest specimen). 
The largest specimen seen (a female) is 463 mm long. But we have no clue as to the 
size to which this Skate may grow, for the claspers on the largest male (3 1 6 mm) still 
reach only about halfway rearward along the inner margins of the pelvics. 

Developmental Stages. The egg cases have not been seen. 

Habits. The great depths at which this Skate has been taken (835-1,188 fath. 
in the western side of the Atlantic and somewhere between 673 and 893 fath. in the 
eastern side) point to a strictly deep-sea habitat, as does the uniformly dark coloration. 
Nothing else is known of its habits. 

Range. Both sides of the North Atlantic in deep water; Irish Atlantic Slope (type 
locality) in the east, a single young specimen ; lower part of the continental slope in 
the west, from the offing of Chesapeake Bay to the offing of southern Nova Scotia. 
This is one of the few Skates that occur on both slopes of the North Atlantic basin. 

Occurrence in the Western Atlantic. Five specimens have been taken so far in the 
western side of the Atlantic, all of them by the Albatross in the summers of 1884 
and 1886 at the localities and depths given (p. 159, footnote 75). 

Synonyms and References: 

Rata bathyphila Holt and Byrne, Fish. Ireland Sci. Invest. (1906), 5, 1908: 51 (descr., meas., Irish Atlant. 

slope between 893 and 673 fath.); Garman, Mem. Harv. Mus. comp. Zool., j6, 1913: 329 (descr., 

after Holt and Byrne, 1908). 
Rata lintea (in part) Clark, Rep. Fish. Bd. Scot. (1926), i, 1926: 45 (considered synonym of R. lintea Fries 

1838); Clark, Faune Ichthyol. Cons. int. Explor. Mer, 1930: pi. not numbered (considered synonym 

of R. lintea Fries 1838). 
Raja lintea (in part) Jensen, Skr. zool. Mus. Kbh., 9, 1938: 46 (considered synonym of ^. lintea Fries 1838). 



Raja eglanteria Bosc 1802 
Brier Skate, Clear-nosed Skate, Summer Skate 
Figures 32, 33, 34, 35 

Study Material. Thirty-one specimens, male and female, 135-785 mm long, 
from: Woods Hole and 80-90 miles southward off Marthas Vineyard, Massachusetts; 
Middle Atlantic Coast; Chesapeake Bay; off Cape Hatteras (Lat. 35°26' N, Long. 

84. Holt and Byrne, Fish. Ireland Sci. Invest. (1906), 5, 1908: 51. 

85. Clark, Rep. Fish. Bd. Scot. (1926), i, 1926: 45. 



66 



Memoir Sears Foundation for Marine Research 




Figure 32. Raja eglanteria. A Female, 745 mm long, from Woods Hole, Massachusetts (Harv. Mus. Comp. 
Zool., No. 36234). B Male, 704 mm long, from Woods Hole, Massachusetts (Harv. Mus. Comp. Zool., 
No. 36233). C Posterior margin of nasal curtain of same, about 2.1 X. i? Margin of nostril of same, about 2.5 X . 

75°26' W) and off Southport (Cape Fear), North Carolina; Charleston, South Carolina; 
Brunswick, Georgia; Pass-a-Grille and New Smyrna Beach, Florida; all in the Harvard 
Museum of Comparative Zoology, the Museum of Zoology, University of Michigan, 
and the U. S. National Museum; two egg cases, with young hatched from them, taken 
at Woods Hole, in Harvard Museum of Comparative Zoology; an embryo with large 
yolk sac and egg case fragments, from Florida, in the American Museum of Natural 
History. 

Distinaive Characters. The presence of only a single row of thorns along the 
midridge of the back on the disc separates R. eglanteria from R. erinacea, R. ocellata, 
and R. garmani; the midrow of thorns extending forward to the region of the shoulders 
on eglanteria distinguishes it from laevis in which the midrow is confined to the tail; 
it is separated from R. senta in that the entire tail is thorny; the facts that the thorns 
of the midrow along the tail are not much larger than those of the lateral rows and 
that they number upwards of 1 8 separate it from R. radiata. The dark bars on the 
upper surface of its disc are also distinctive. Should its range prove to extend farther 



Fishes of the JVestern Nortfi Atlantic 



167 




Figure 33. Raja eglanteria. A Female, 640 mm long, and B Male, 568 mm long, both from Charleston, South 
Carolina (Harv. Mus. Comp. ZooL, Nos. 122 and 81). 



southward than present records indicate, its color pattern of many small dark spots 
and bars without ocellar spots would serve to separate it from both R. ackleyi and 
R. texana. 

R. eglanteria is not closely paralleled in shape of disc, distribution of thorns, and 
color pattern by any Skate so far known from the eastern North Atlantic, South Atlantic, 
or Pacific coasts of America. 

Description. Proportional dimensions in per cent of total length. Male, 704 mm, 
and female, 745 mm, from Massachusetts (Harv. Mus. Comp. ZooL, Nos. 36233 and 
36234, respectively). 



i68 



Memoir Sears Foundation for Marine Research 




Figure 34. Raja eglanterla. A Female, 156 mm long, from New Smyrna Beach, Florida (Harv. Mus. Comp. 
ZooL, No. 36397), to illustrate juvenile color pattern and dermal armature. B Ventral view of anterior part of 
head of nearly mature male illustrated in Fig. 32, about 0.4 X. C Lateral view of posterior part of tail of same, 
about 0.5 X. D Upper teeth of same, about 5.5 X. E Upper and lower teeth of female illustrated in Fig. 32, 
about 4X . 



T)isc: extreme breadth 66.8, 65.7; length 51.3, 53-7- 

Snout length: in front of orbits 14.3, 15.3; in front of mouth 14.6, 16.8. 

Orbits: horizontal diameter 3.8, 4.1; distance between 4.7, 5.0. 

Spiracles: length 3.0, 3.0; distance between 6.5, 6.8. 

Mouth: breadth 7.5, 8.7. 

Nostrils: distance between inner ends 7.5, 8.0. 

Gill openings: lengths, ist 1.7, 1.9; 3rd 1.9, 2.1 ; 5th 1.3, 1.3; distance between 

inner ends, ist 15.4, 16.0; 5th 8.8, 9.7. 
First dorsal fin: vertical height 2.7, 2.7; length of base 6.1, 5.4. 
Second dorsal fin: vertical height 2.2, 2.5; length of base 6.4, 6.4. 
Pelvics: anterior margin 10. o, 12.2. 
Distance: from tip of snout to center of cloaca 47.8, 51.5; from center of cloaca 



Fishes of the JVester'n Northr Atlantic 169 

to 1st dorsal 37.0, 32.3; to tip of tail 52.2, 48.5; from rear end of 2nd 
dorsal base to tip of caudal 2.0, 2.8. 
Interspace between: ist and 2nd dorsals 0.9, 1.6. 

Disc about 1.2 — 1.3 times as broad as long; anterior contour about a right angle, 
a little more obtuse in southern specimens than in northern;*' anterior margins weakly 
concave posterior to snout, weakly convex abreast of spiracles in females and only 
slightly more so in adult males; posterior corners broadly rounded, the outer corners 
more narrowly so. Axis of greatest breadth 73-76 "/o of distance back from tip of 
snout toward axils of pectorals. Tail with moderately wide lateral folds, its length from 
center of cloaca to origin of first dorsal about 0.7—0.8 as great, and from center of 
cloaca to origin of second dorsal about 0.8-0.9 ^s great, as distance from center of 
cloaca to tip of snout, on both large and small specimens. 

Upper surface of disc with 1—5 thorns on each shoulder region;*' 2-4 thorns 
at inner end of spiracle; 3-4 close behind orbit and as many close in front of it; also 
an irregular row around inner margin of orbit; a single continuous midrow, extending 
rearward along disc from nuchal region, the thorns varying in size from place to place 
(perhaps with age) and increasing in number with growth from 14-16 in small speci- 
mens (185—267 mm long) to 33—39 in those longer than 400 mm. 

On southern specimens (Figs. t,i,, 34)/* in addition to thorns, the upper surface 
of the disc, the base of the tail, and the central part of the pelvics is generally beset with 
small prickles from the time of hatching; females either continue so until maturity or 
may develop small naked areas outward and rearward from the shoulders. But mature 
males continue to be prickly only on the outer anterior parts of the pectorals, on the 
head outward from the eyes, in front of the eyes, and between the orbits. Northern 
examples*" seem less rough; the upper surface of the disc and the pelvics is naked at 
hatching (Fig. 35) except for the characteristic thorns; later the females become prickly 
along the margins of the disc rearward from the level of the orbits (perhaps more gener- 
ally so over pectorals in some cases), in a band along either side of the mid-dorsal ridge 
from the nuchal region rearward, over a patch in front of each orbit, along the edges 
of the rostral ridge, on the tip of the snout, and sometimes in the space between the 
orbits (Fig. 32). Mature northern males are prickly only along the anterior margins 
of the pectorals posterior to the level of the spiracles, over patches in front of the eyes, 
on the rostral ridge and tip of snout, and hence they are considerably smoother than 
adult females. 

The tail, from the axils of the pectorals to the first dorsal, has a median row of 
about 16-19 moderately large thorns on newborn specimens which increase to 26—32 

S6. Maximum anterior angle in front of spiracles 90° in two specimens from Woods Hole; 100-105^ in two from North 

Carolina and 92-105° in three from South Carolina. 
87. One or two on large specimens of both sexes from Buzzards Bay, Massachusetts and on a female from Charleston, 

South Carolina; 3-5 on a smaller but sexually mature male from the latter locality. Sometimes these thorns are 

lacking. 
XS. Florida and South Carolina. 89. Woods Hole, Massachusetts. 



I 70 Memoir Sears Foundatmi for Mari?ie Research 

on large; also, low down along either side of the tail of northern specimens there 
are 1—2 rows, 2—4 on southern. The dorsal fins are uniformly prickly at all sizes 
and in both sexes, the interspace between them having 1—3 thorns, larger or smaller. 
Sexually mature males have about 25—40 large thorns on the marginal region on 
each side about at the level of the eyes; the alar spines are arranged in 4—7 rows on 
the outer parts of the pectorals, there being about 28 spines in the innermost (longest) 
row. The lower surface on smaller specimens is smooth except for a prickly snout tip, 
but with growth the head becomes roughened with prickles rearward in a narrow band 
along the margins of the disc about to the level of the mouth on males, and a short 
distance farther on some females. Females also develop a prickly patch close in front 
of the axil of the pectoral. The lower surfaces of the pelvics are smooth on males but 
more or less prickly on large females. 

Snout in front of orbits 2.7—3.1 times as long as distance between orbits; its 
length in front of mouth 1.7—2.2 times as great as distance between exposed nostrils. 
Orbit 1. 2-1. 4 times as long as spiracle; distance between orbits about 1.2 times as 
great as length of orbit. 

Distance between first gill openings about 1.9— 2.1 times as long as distance 
between exposed nostrils; fifth gill openings about 75 "/o as long as first; distance be- 
tween fifth pair of gills i. 0—1.2 times as long as distance between exposed nostrils 
and about 0.9—1.2 times as long as breadth of mouth. 

Nasal curtain fringed; expanded posterior (outer) margin of nostril also fringed. 
Mouth nearly straight in females, only a little more arched in adult males. 

Teeth 48Z49 (four specimens) in transverse series, close-set, with low conical cusp, 
blunter on older rows and sharper on younger; sharper also on mature males than on 
females, especially along the younger (posterior) rows. 

First and second dorsals similar in size and shape; interspace between them 
14-30 % as long as base of first dorsal. Caudal membrane from rear end of base of 
second dorsal 32—53 "/o as long as base of first dorsal. Pelvics only moderately con- 
cave outwardly, more or less scalloped, most strongly so at base of marginal concavity; 
anterior margin about half as long as distance from origin of pelvic to rear tip; anterior 
lobe fleshy with rounded tip, including 4—5 slender radial cartilages as well as the 
first and stoutest; outer margin of posterior lobe nearly straight anteriorly, weakly 
convex toward narrowly rounded rear tip. 

Anterior rays of pectorals extending about 55-65 "/o of distance from level of 
fronts of orbits toward tip of snout. 

Color. A living specimen nearly three feet long, observed in an aquarium at Woods 
Hole, Massachusetts, was dark and light brown above, with many roundish spots as 
well as more elongate bars of the darker shade, and with irregular spots of the lighter 
shade about ^U—\ inch across; these markings appear on the tail as wide dark brown 
crossbars separated by light brown areas ; the disc, pelvics, lateral edges of tail, and dor- 
sal outlined with a narrow band of white. There is a translucent space on either side 
of the rostral ridge. The lower surface is white without dark markings. Other specimens 



Fis/ies of the Western North Atlantic 



171 





Figure 35. Raja eglanterta, from Woods Hole, Massachusetts. Egg case, and young skate hatched from it, 
about 0.9 X . 

have been described as brown or gray above, but all appear to have the characteristic 
dark spots and elongate bars. 

Remarks. Attention has been called earlier to the greater prickliness of southern 
specimens (p. 169) and to the tendency for southern specimens to mature at a smaller 
size than the northern ones (p. 172). The series that we have studied is not sufficiently 
extensive or the geographic coverage sufficiently complete for us to judge whether or 
not these apparent differences indicate the existence of two separate races or subspecies 
of eglanteria — a southern (typical) form, and a northern form for which the name 
americana DeKay 1842 would appear to be available. 

Size. At hatching, Woods Hole specimens are 135— 144 mm long to the rear 
end of the second dorsal fin, our smallest Florida specimens 1 30 mm. In the northern 
part of its range some specimens of R. eglanteria may mature at a length of about 
650 mm, but others probably not until 725 mm or longer."" Also, maturity may be 



90. In the Block Island, Rhode Island, region males were found to mature at 750—770 mm (one at 620 mm) and females 
at 760-780 mm (from unpublished data at the Bingham Oceanographic Laboratory). 



17-2 Memoir Sears Foundation for Marine Research 

reached at a smaller size in the south, for the tips of the claspers of two males 568 mm 
long from Charleston, South Carolina, and 535 mm long from Pass-a-Grille (near 
Tampa), Florida, appear to have reached their full development. A specimen 30 inches 
long weighed 6 pounds and one of 31 inches 6V2 pounds."' A length of 37V4 inches 
is the greatest recorded. '^ 

Developmental Stages. Eggs are probably laid in the spring, at least in the northern 
part of its range. A female 785 mm long, taken off southern Massachusetts'^ in January, 
had well developed ovaries with eggs 5—15 mm in diameter. 

The egg cases range in size from about 2—372 inches long (without the horns) 
and i^l2—l^U inches wide,'* those produced by Florida specimens being considerably 
smaller than those from New York'^ and Massachusetts specimens. The cases are of 
the usual Skate shape, the transverse margin being nearly straight and more or less 
ragged at the end with the longer pair of horns, but moderately concave at the end 
with the shorter pair. The horns are shorter than the egg case proper. At the end 
of the case with the longer horns, newly laid eggs in the aquarium had a thin, trans- ■ 
parent, delicate area which subsequently sloughed away, thus forming a slit which 
probably serves the same respiratory function as do the slits on the horns in Raja 
ocellata (p. 246). But this was not visible in preserved specimens. Incubation occupies 
a period of at least three months'^ but it is not known how much longer. Newborn 
specimens already show dark markings somewhat similar to the adult pattern. 

Habits. R. eglanteria breeds while inshore. A pair in coitus has been seen (and 
harpooned) off the coast of Virginia in the spring, "The male grasping the anterior 
part of the pectoral of the female from above with his teeth, and with claspers directed 
forward and well inserted."" A female brought in to the New York Aquarium from 
Sandy Hook in June laid two eggs the next day, and two more three days later. Egg 
cases, probably of this species, have also been found on beaches in August. 

The stomachs of R. eglanteria that have been examined so far have contained 
fish, squid,'* crabs and shrimps." 

R. eglanteria often comes so close to the land that it is taken regularly on hook 
and line from piers and jetties; however, it moves offshore for the cold months and has 
been taken in depths as great as 65 fathoms in January and February. 

91. Average weights in the Block Island region were found to be: males 24.4 inches long, 2.61 pounds; 27.7 inches, 
4.28 pounds; 29.25 inches, 4.64 pounds; females 24.5 inches, 3.18 pounds; 27.6 inches, 3.96 pounds; 32.7 inches, 
7.65 pounds (from unpublished data at the Bingham Oceanographic Laboratory). 

92. Schroeder, Bull. Boston Soc. nat. Hist., 66, 1933: 5. 

93. Caught by the dragger Eugene H., January 31, 1950 in 60 fathoms at about Long. 70° 30' W. 

94. Two egg cases from females taken off Marthas Vineyard, Massachusetts, were 3'/$ by z'/j and 3 by z'/g inches. 

95. For description and photographs of eggs laid in the New York Aquarium by a female taken near Rye, New York, 
see Breder and Nichols (Copeia, 1937: 181-183). 

96. Eggs laid on June i;-i8 in the New York Aquarium had died by September 13 (Breder and Nichols, Copeia, 
1937: 182). 97. Fowler, Rep. N.J. Mus. (1905), 1906: 72. 

98. In the Block Island, Rhode Island, region, the stomachs of R. eglanteria commonly contain squid throughout early 
summer. Butterfish {Poronotus) and scup [Stenotomus) are a dominant food during September and October (from 
unpublished data at the Bingham Oceanographic Laboratory). 

99. For stomach contents of Chesapeake Bay specimens, see Hildebrand and Schroeder (Bull. U. S. Bur. Fish., 43, 
1928: 59). 



Fishes of tJie Western NortJi Atlantic 173 

In the northern part of its range, R. eglanteria is strictly a warm-season visitor 
in the shore waters. It appears in April in the sector between Chesapeake and Delaware 
bays and is present along New Jersey and near New York from mid-May (occasionally 
late April) through the summer into October (sometimes until November), but it is 
taken only from July until September off southern Massachusetts. It withdraws from 
the immediate vicinity of the coast during the hottest season in the more southern 
part of its range.'"" Thus, it has not been reported in July or August for Chesapeake 
Bay though common there in April, May, June, as well as September and October; 
and odd specimens only have been reported for July at Cape Lookout, North Carolina, 
where it is plentiful in April and May.'"' 

The stock that summers to the northward appears to migrate southward and off"- 
shore in autumn, for specimens have been trawled in winter and early spring off southern 
Massachusetts,'"- New Jersey, Virginia and North Carolina. '"^ However, it is doubtful 
whether the stock inhabiting the warmer waters from South Carolina south to Florida 
leave the coast at all during the winter. 

The relationship of i?. eglanteria to temperature is puzzling. Although its autumnal 
withdrawal from the immediate vicinity of the coast in the northern part of its range 
occurs when the upper few fathoms have chilled to about 13—16° C (about 55—61° F,)'"* 
all winter records for it have been from water as cold as 6.3—1 1° C (43.3—51.8° F);'"^ 
also, it reappears along the shores of Virginia, New Jersey and New York when the 
water has warmed only a little from the winter minimum. The direction of temperature 
change may be the governing factor in its thermal migrations. It is possible also that 
this Skate is more sensitive to low temperatures in the southern than in the northern 
part of its range, for it is reported as appearing in abundance on the coast at Cape 
Lookout, North Carolina only after the water has warmed to about 12—17° C (54— 
63° F). In the northern part of its range'"^ it has not been reported regularly anywhere 
in water warmer than perhaps about 21-22° C (70-72° F). Its presence along the 
New Jersey beaches throughout the summer might seem to contradict this statement, 
for the mean surface temperature there usually rises to at least about 23° C (73—74° F), 
and may reach 25° C (77° F). But those taken in pound nets at a depth of 20-30 feet 

100. It is also said that it leaves New York Harbor during the hottest months, but reports conflict in this respect. 
loi. Unfortunately no information is available in this regard for any areas farther south. 

102. We examined five specimens taken in 44 hauls while aboard the dragger Eugene H., fishing 80-90 miles to the 
southward of Marthas Vineyard, Massachusetts, in 47-65 fathoms, January 27-February 3, 1950. These included 
females 297, 600, 640 and 785 mm long and a mature male with large claspers 760 mm long. 

103. The U.S. Fish and Wildlife Service vessel Albatross III trawled six specimens about 35 miles southeast of Cape 
Fear, January 31, 1950, in Lat. 33° 38' N, Lung. 77° 28' W, in 13 fathoms; also two specimens, about a mile off 
Southport on February 7, 1950. 

104. According to Breder (Copeia, 127, 1924: 27) it is not to be found in Sandy Hook Bay after the water cools to 14.4° C 
(58° F). 

105. Specimens reported off northern New Jersey in 6-17 fathoms in January (Fowler, Copeia, 31, 1916: 41) may have 
been in water as cool as 5.5-6° C (42-43° F). 

106. Taken only occasionally at Cape Lookout after the water has warmed to more than 21.1° C or 70° F (mid- 
May); absent from Chesapeake Bay during the period (July-August) when the bottom water is much warmer 
than that. 



174 Memoir Sears Foundation for Marine Research 

may actually be in water of considerably lower temperature due to inshore movements 
of cooler bottom water. i"' 

In the southern part of its range (North Carolina to Florida) "^ ^. eglanteria may 
inhabit somewhat higher temperatures than those in the north, but definite conclusions 
in this regard would be premature. 

The highest salinity in which R. eglanteria has been taken is about 34 "/oo, while 
those that enter Chesapeake Bay meet with salinities as low as 27—31 °/oo near the 
entrance and lower still if they venture up the Bay, as some do.i"^ However, there is 
no evidence that R. eglanteria ever runs up into fresh water. 

Numerical Abundance. R. eglanteria has been characterized repeatedly as "abundant" 
or as the "most numerous Skate" at various coastwise points between New Jersey 
and Virginia. There is an early report of not less than 10,000 pounds of them (weighing 
1—5 pounds each) being discarded from one pound net on the New Jersey Coast on 
a single July day,ii" but this represented an exceptional concentration, even allowing 
for exaggeration. Ten specimens is the largest catch that has been reported from any 
one trawl haul on the wintering grounds offshore. 

Relation to Man. Most of those caught by commercial fishermen and by anglers 
are discarded; however, a small percentage of the catch is utilized as food. 

Range, Western North Atlantic from Massachusetts Bay to Florida, mostly in 
shoal water from close in to the coast out to about 65 fathoms. 

Details of Occurrence. This Skate occurs regularly and plentifully at one season 
or another along the whole coastline of the United States from the northern part of 
Florida on both coasts to Long Island, New York."^ To the northward and eastward, 
a few are taken yearly off southern New England and in the vicinity of Woods Hole; 
we caught two specimens in 10—15 fathoms on a hand line on Nantucket Shoals 
in July and September 1926."^ Also, there are odd records for it from Province- 
town at the tip of Cape Cod and from Gloucester at the northern boundary of Massa- 
chusetts Bay. To the southward, it has long been known on the South Carolina Coast 
— in fact, Charleston is the type locality; it has been taken off Brunswick, Georgia; 
many are caught off northern Florida;"* and we have specimens from New Smyrna 
Beach farther to the southward. It has been recorded from the west coast of Florida 
but not from the northern shore of the Gulf of Mexico, but perhaps it is to be expected 
there. It is not known from any localities farther south than middle Florida. 

107. Information is scant. On August lo, 1916 the surface close to Cape May was about 21.9 ° C (71° F), but the bottom 
at about 50 feet was only about 15.8° C (60° F). 

108. Near Charleston, South Carolina the surface chills to about 10-11° C (50—52° F) in winter, warms to about 21.1° C 
(70° F) by mid-May, and is warmer than 24° C (75° F) from about mid-June to the end of September. Off the 
mouth of the St. Johns River, Florida the summer maximum is about the same and the winter minimum only 
a little higher; i. e., about 12.2-13.0° C (54-55° F) (Parr, Bull. Bingham oceanogr. Coll., 4 [3], 1933 : 68, fig. 22). 

109. Newcombe and Lang, Proc. Amer. phil. Soc, 81, 1939: 399. 
no. Smith, Bull. U.S. Fish Comm., 12, 1894: 368. 

111. Recorded repeatedly in scientific literature from many localities in New York, New Jersey, Delaware, Maryland, 
Virginia, lower Chesapeake Bay, and North Carolina. 

112. Bigelow and Schroeder, Bull. U.S. Bur. Fish., 48, 1936: 325. 

113. Personal communication from Stewart Springer. 



Fishes of the Western NortJi Atlantic 175 

Synonyms and References: 

Raja}-^^ eglanleria Bosc, in Lacep^de, Hist. Nat. Poiss., 4° ed., in Buffon, Hist. Nat., 2, 1802: 104, 109, pi. 4, 
fig. 2 (descr., ill., Charleston, S. Carolina); Giinther, Cat. Fish. Brit. Mus., 8, 1870: 462 (descr., New 
York); Garman, Proc. Boston Soc. nat. Hist., ij, 1874: 179 (descr., Peni!<ese I., Mass.); Uhler and 
Lugger, Rep. Coram. Fish. Md., ist ed., 1876: 188; 2nd ed., 1876: 159 (Chesapeake Bay); Yarrow, 
Proc. Acad. nat. Sci. Philad., [29], 1877: 217 (listed, near Beaufort, N.Carolina); Goode and Bean, 
Bull. Essex Inst., II, 1879: 28 (Essex County, Mass.); Bean, Proc. U. S. nat. Mus., J, 1880: 115 (Mass. 
and Conn.); \^errill, Amer. J. Sci., [3] 22, 1881 : 297 ("sparingly" in 130-160 f., at Gay Head); Jordan 
and Gilbert, Bull. U. S. nat. Mus., 16, 1883: 41 (descr., Atlant. Coast U. S.); True, List Vert. Anim. 
S.Carolina, in Handb. S.Carolina, 1883: 261 (listed, S.Carolina); Bean, Rep. U. S. Coram. Fish. 
(1882), 1884: 343 (listed, Woods Hole); Jordan, Rep. U. S. Coram. Fish. (1885), 1887: 799 (listed, 
N. Atlant., U. S.); Bean, Bull. U. S. Fish Corara., 7, 1888: 151 (New Jersey); Moore, Bull. U. S. Fish 
Comm., 12, 1894: 357 (New Jersey); Smith, Bull. U. S. Fish Coram., 12, 1894: 368 (nos., size. New 
Jersey); Jordan and Everraann, Rep. U. S. Coram. Fish. (1895), 1896: 221 (listed, C. Cod to Florida); 
Bull. U. S. nat. Mus., 47 (i), 1896: 71 (descr., C. Cod to Florida); Smith, Bull. U. S. Fish Coram., 
Ij, 1898: 89 (nos.. Woods Hole, Mass.); Bean, 6th Rep. For. Coram. N. Y., 1900: 383 (nos.. Long 
Island, New York and Virginia); Evermann and Kendall, Rep. U. S. Comm. Fish. (1899), igoo: 49 
(Florida, by ref. to R. desmarestia Lesueur 1824); Howe, Bull. U. S. Fish Corara., ig, 1901 : 240 (south. 
New England); Linton, Bull. U. S. Fish Comm., ig, 1901 : 431 (nematode parasites); Bean, Bull. N. Y. 
St. Mus., 60, Zool. 9, 1903: 49 (descr., cf. other species, season, near New York); Werner, Zool. Jb., 
Syst. Abt., 21, 1904: 298 (C. Cod); Fowler, Rep. N. J. Mus. (1905), 1906: 72 (descr., nos., season, 
pair in coition; New Jersey, Delaware Bay, Virginia); Rep. N. J. Mus. (1906), 1907: 259 (New Jersey); 
Smith, N. Carolina geol. econ. Surv., 2, 1907: 42 (descr., iU., C. Lookout, N. Carolina); Fowler, Rep. 
N. J. Mus. (1907), 1908: 131 (season. New Jersey); KendaU, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 
1908: 12 (New England locals.); Fowler, Proc. Acad. nat. Sci. Philad., 61, 1909: 407 (Delaware Bay); 
Rep. N. J. Mus. (igo8), 1909: 352 (New Jersey); Proc. .Acad. nat. Sci. Philad., 62, 1910: 470 (New 
Jersey); Gudger, Amer. Nat., 44, 1910: 398 (near Beaufort, N.Carolina); Fowler, Proc. Acad. nat. 
Sci. Philad., 63, 191 1: 5 (egg cases. New Jersey); Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 4 (3), 
1913: 102 (range); Fowler, Proc. Acad. nat. Sci. Philad., 65, 1913: 62 (nos., season, Maryland, Vir- 
ginia); Garman, Mem. Harv. Mus. comp. Zool, j6, 1913: 341, pi. 23 (descr., ill., Charleston, S. Caro- 
lina); Gudger, J. Elisha Mitchell sci. Soc, 28, 1913: 166 (season, nos., C. Lookout, Beaufort, N. Caro- 
lina); Nichols, Abstr. Linn. Soc. N. Y., 1913: 91 (season, near New York); Suraner, Osburn and Cole, 
Bull. U. S. Bur. Fish., jr (2), 1913: 738 (parasites. Woods Hole); Fowler, Copeia, 1914: 2 (listed, 
Maryland); Proc. Acad. nat. Sci. Philad., 65, 1914: 62 (nos., season, Maryland); 66, 1914: 353 (nos., 
season, Delaware); Coles, Proc. biol. Soc. Wash., 28, 191 5: 92 (nos., season, C. Lookout, N. Carolina); 
Fowler, Copeia, 27, 1916: 12 (New Jersey); Copeia, 31, 1916: 41 (winter, off New Jersey); Nichols, 
Copeia, 27, 1916: 9 (season, nos., near New York); Radcliffe, Bull. U. S. Bur. Fish., 34, 1916: 272, 
pi. 45, fig. I (descr., ill., teeth, derraal arraature, season, N. Carolina); Fowler, Proc. Acad. nat. Sci. 
Philad., 6g, 1917: 109 (New Jersey); Occ. Pap. Mus. Zool. Univ. Mich., No. 56, 1918: 2, 16 (New 
Jersey, Virginia); Copeia, 68, 1919: 13 (New Jersey); Lathara, Copeia, 71, 1919: 54 (nos., weight, 
season. Long Island, New York); Fowler, Proc. Acad. nat. Sci. Philad., yi, 1920: 293 (New Jersey); 
Proc. biol. Soc. Wash., JJ, 1920: 145 (New Jersey); Lathara, Copeia, 87, 1920: 91 (season, Long Is- 
land, New York); Copeia, 99, 1921 : 72 (listed. Long Island, New York); Townsend, N. Y. zool. Soc, 
25th Annu. Rep. (1920), 1921 : 126 (listed. New York Bay); Townsend and Nichols, Copeia, 91, 1921 : 
10 (season. New York Harbor); Latham, Copeia, 112, 1922: 81 (season. Long Island, New York); 
Fowler, Proc. Acad. nat. Sci. Philad., 7./, 1923: 3, 6 (New Jersey); Breder, Copeia, 127, 1924: 27 
(temp.. New York); Linton, Proc. U. S. nat. Mus., 64 (21), 1924: 7, 9, 10, 29, 66 (cestode parasites. 
Woods Hole); Bigelow and Welsh, BuD. U. S. Bur. Fish., ./o (i), 1925 : 64 (descr., iU., Gulf of Maine); 
Fowler, Copeia, 143, 1925: 41, 43 (Delaware and New Jersey); Copeia, 156, 1926: 146 (New Jersey); 
Breder, Zoologica N. Y., 4 (5), 1926: 243 (locoraotion) ; Fowler, Copeia, 165, 1927: 89 (season, Dela- 
ware, Maryland); Nichols and Breder, Zoologica N. Y., 9, 1927: 25 (ill.. New York and south. New 
England); Fish Culturist, 7, 1928: 225 (listed. New Jersey); Hildebrand and Schroeder, BuU. U. S. 
Bur. Fish., 43, 1928: 58 (descr., iU., food, season, nos., Chesapeake Bay); Breder, Field Bk. Mar. Fish. 

114. Also spelled Rata. 



I 76 Memoir Sears Foundatio?i for Marine Research 

Atlant. Coast, 1929: 31 (genl); Fowler, Proc. Acad. nat. Sci. Philad., 80, 1929: 608 (New Jersey); 
Truitt, Bean and Fowler, Bull. Md. Conserv. Dep., 3, 1929: 30 (descr., Maryland); Jordan, Evermann 
and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 25 (listed, C. Cod to Florida); Breder, Copeia, 
1932: 31 (season. Block Island); Wilson, Bull. U. S. nat. Mus., 158, 1932: 403 (copepod parasite. 
Woods Hole); Schroeder, Bull. Boston Soc. nat. Hist., 66, 1933: 5-6 (max. size, south. Mass.); Bige- 
low and Schroeder, Canad. Atlant. Fauna, I2e, 1934: 32 (descr., ill., C. Ann to Florida); Bere, Amer. 
Midi. Nat., 77 (3), 1936: 583 (parasites, season, Englewood, Florida); Bigelow and Schroeder, Bull. 
U. S. Bur. Fish., 48, 1936: 325 (max. size, depth, south. Mass.); Breder and Nichols, Copeia, 1937: 
181 (egg cases, New York, Florida; cf. other species); Fowler, Proc. Acad. nat. Sci. Philad., i'y, 1937: 
304 (season. New Jersey); Breder, Bull. N. Y. zool. Soc, 41, 1938: 28 (New York Harbor); Delsman, 
Mem. Mus. Hist. nat. Belg., (2) Fasc. 21, 1941 : 65 (depth, off Jacksonville, Florida); Fowler, Monogr. 
.'\cad. nat. Sci. Philad., 7, 1945: 160 (listed, S.Carolina). 

Raja''^'-' diaphanes Mitchill, Trans. Lit. philos. Soc. N. Y., r, 181 5: 478 (descr., shows it equal to R. eglanteria 
Bosc 1802, not R. ocellata Mitchill 181 5, to which the majority of recent authors have referred it; New 
York); Ayers, Boston J. nat. Hist., 4, 1843: 289, pi. 12, fig. 2 (ident. by ill.. Long Island, New York); 
Linsley, Amer. J. Sci., 4-], 1844: 77 (ident. by ref. to Mitchill, 181 5; Conn.); Baird, Rep. U. S. Comm. 
Fish. (1871-1872), 1873: 826 (ident. probable by name of "Summer Skate," Woods Hole). 

Raia^'^'' desmarestia Lesueur, J. Acad. nat. Sci. Philad., 4, 1824: 100, pi. 4 (descr., ill., Florida); Miiller and 
Henle, Plagiost., 1841 : 1 54 (diagn. by ref. to Lesueur, 1824); DeKay, Zool. N. Y., 4, 1842: 372 (diagn. 
after Lesueur, 1824, Florida); Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 511 (diagn., after 
Lesueur, 1824, Florida); Tuomey, Rep. Geol. S. C, 1848: app. XIII (listed, S. Carohna); Dumeril, 
Hist. Nat. Poiss., J, 1865: 551 (descr., Charleston, S.Carolina); Giinther, Cat. Fish. Brit. Mus., 8, 
1870: 455, footnote a (ref to Lesueur, 1824); Goode, Proc. U. S. nat. Mus., 2, 1879: 120 (Florida, 
probably by ref. to Lesueur, 1824). 

Rata eglanteria (in part) DeKay, Zool. N. Y., 4, 1842: 373 (but R. eglantiera Lesueur 1824, which equals 
R.erinacea Mitchill 181 5, also included); Storer, Mem. Amer. Acad. .%ts Sci., N. S., 2, 1846: 512 
(but R. eglantiera Lesueur 1824, which equals R.erinacea Mitchill 181 5, also included); Gill, Rep. 
U. S. Comm. Fish. (1871-1872), 1873: 812 (listed, Nova Scotia to Florida, but includes R. eglantiera 
Lesueur 1824, which equals R.erinacea Mitchill 181 5). 

Raia diaphanes (in part) Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 510 (descr. apparently also refers 
to R. ocellata Mitchill 181 5); Lyman, 6th Rep. Mass. Comm. inl. Fish., 1872: 54, 57 (probably includes 
R. ocellata Mitchill 181 5 because of descr. as abundant, south. Mass., April, Ma)--). 

Probable References: 

Raia americana DeKay, Zool. N. Y., 4, 1842: 368, pi. 66, fig. 215 (ident. probable from ill., New York); 

Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861: 61 (hsted). 
Raia ocellata DeKay, Zool. N. Y., 4, 1842: Fishes, pi. 65, fig. 212 (ill. apparently R. eglanteria, but perhaps 

not descr. on p. 369). 
Raja eglantiera Tuomey, Rep. Geol. S. C, 1848: Append. XIII (listed, S.Carolina; probably R. eglanteria 

Bosc 1802, not R. eglantiera Lesueur 1824, which equals R.erinacea Mitchill 1815). 
Not Raia eglantiera Lesueur, J. Acad. nat. Sci. Philad., 4, 1824: 103 (equals R.erinacea Mitchill 181 5: see 

reference, p. 185). 
Not Raia eglanteria Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861 : 61 (ref to R. eglantiera Lesueur 1824, 

equals R.erinacea Mitchill 181 5). 

Raja erinacea Mitchill 1825 

Little Skate, Hedgehog Skate 

Figures 36, 37, 38 

Study Material. Sixty-two specimens, from embryos to adults of both sexes, from 
many localities between St. Andrews, New Brunswick and the offing of Delaware 

115. Also spelled Kaia. ii6- Also spelled Kaja. 



Fishes of the JVestern North Atlantic 



177 




Figure 36. Jiaja erinacea. A Male, 507 mm long, from Boston Harbor, Massachusetts (Harv. Mus. Comp. Zool., 
No. 1021). B Female, 443 mm long, from Mystic, Connecticut (Harv. Mus. Comp. Zool., No. 1273). C Side 
view of posterior part of tail of same, about 0.7 X. 



(Lat. 38°32' N), in the Harvard Museum of Comparative Zoology; three specimens 
from Kamouraskai" (lower St. Lawrence River) and others in the U. S. National 
Museum; also a considerable number, identified by us but not preserved, trawled by 
the U. S. Bureau of Fisheries steamer Albatross II and the U. S. Fish and Wildlife 
Service vessel Albatross III at stations scattered along the continental shelf off southern 
New England and the mid-Atlantic United States. 

Distinctive Characters. Females of R. erinacea., and males in which the claspers 
are still small, resemble specimens of R. ocellata so closely that the only sure means 
of identification is by counting the teeth; R. erinacea never has more than 66 series, 

117. From v. D. Vladykov. 



178 Memoir Sears Foundation for Marine Research 

usually less than 54, in the upper jaw, while R. ocellata always has at least 72 series 
and usually more than 80. i" In the case of sexually mature males, size is an equally 
dependable specific character, for R. erinacea grows only to a length of about 2 i inches, 
whereas R. ocellata does not mature sexually until it has reached a length of at least 
26—28 inches. Adults of R. erinacea resemble those of R.fyllae of West Greenland 
and the northeastern Atlantic in arrangement of thorns, in proportional dimensions, 
in the shape of snout, and in the fact that the two dorsals are confluent. But they are 
easily distinguishable from R.fyllae by their shorter tails and by the fact that the upper 
surface of R. erinacea between the thorns is smooth (partly spinulose in R.fyllae') and 
the margin of the disc less deeply concave opposite the spiracles. There is no dangre 
of confusing young specimens of the two; in R.fyllae the upper surface is uniformly 
prickly with the large thorns on the disc limited to the midline of the back, to the 
shoulders and to the head, whereas in the young of R. erinacea the disc as a whole is 
thorny with the skin smooth between the thorns. Neither R. erinacea., R. ocellata, nor 
R.fyllae is likely to be confused with any other Skate of the western North Atlantic, 
so characteristic are the thorn pattern, shape of disc, and proportionate dimensions 
of each. 

R. erinacea falls closest to R. naevus Miiller and Henle 1841 among European 
Skates in shape of disc, relative length of tail, distribution of thorns, and number of 
teeth; also, in the fact that the mid-dorsal row of thorns is lost with growth. However, 
the midbelt of the disc and the anterior part of the tail are thornier in R. erinacea than 
in R. naevus, whereas the upper surface of the disc is prickly generally between the 
thorns on R. naevus but naked between them on R. erinacea. The inner part of each 
pectoral, which is marked ordinarily with a conspicuous ocellar spot in R. naevus, 
is not so marked in R. erinacea. R. erinacea does not closely resemble any Skate known 
from the South Atlantic or from the Pacific Coast of America. 

Description. Proportional dimensions in per cent of total length. Male, 441 mm, 
and female, 449 mm, from Massachusetts (Harv. Mus. Comp. Zool., Nos. 547 and 
708 respectively). 

Disc: extreme breadth 57.2, 57.7; length 46.5, 46.0. 

Snout length: in front of orbits 9.9, 10.2; in front of mouth 9.5, i 1.3. 

Orbits: horizontal diameter 2,-^^ 3-8; distance between 4.8, 4.9. 

Spiracles: length 2.9, 2.9; distance between 6.8, 6.9. 

Mouth: breadth 8.0, 8.2. 

Nostrils: distance between inner ends 6.8, 6.2. 

Gill openings :\tr\gXh?,, ist 1.7, 1.8; 3rd 2.0, 1.9; 5th 1.6, 1.6; distance between 
inner ends, ist 14. i, 15. i; 5th 7.3, 8.9. 

First dorsal fin: vertical height 2.8, 3.1; length of base 6.6, 6.7. 

Second dorsal fin: vertical height 2.5, 2.9; length of base 5.9, 6.2. 

Pelvics: anterior margin 13.6, 11.6. 

Distance: from tip of snout to center of cloaca 44.3, 45.4; from center of cloaca 

118. See p. 242, footnote 74. 



Fishes of the Western Nort/i Atlantic 
A. 



179 




Figure 37. Raja erinacea. A Juvenile male, 180 mm long, from offNew Jersey, Lat. 39°! i' N, Long. 73°4o' W, 
23 fathoms (Harv. Mus. Comp. Zool., No. 34399), showing dermal armature. B Lower surface of anterior 
part of head of female, 460 mm long, from Mystic, Connecticut (Harv. Mus. Comp. Zool., No. 1270). C Pos- 
terior margin of nasal curtain of same, about 4.1 X. D Outer posterior margin of nostril of same, about 4.1 X. 
E Upper teeth near middle of jaw of female, 445 mm long, from off Woods Hole, Massachusetts (Harv. Mus. 
Comp. Zool., No. 36228), about loX. /" Upper teeth near middle of jaw of mature male, 440 mm long, from 
off Woods Hole, Massachusetts (Harv. Mus. Comp. Zool., No. 36230), about lOX- 

to 1st dorsal 41.5, 39-5; to tip of tail 55.7, 54-6; from rear end of 2nd 
dorsal base to tip of caudal 1.8, 2.2. 
Interspace between: ist and 2nd dorsals 0.0, 0.0. 

Disc about 1.2 times as broad as long, extremely obtuse in front, the maximum 
anterior angle in front of spiracles about 1 10—130°; anterior margins weakly concave 
for a short distance behind tip of snout, thence evenly and weakly convex rearward 
to outer corners of disc in newly hatched specimen ; weakly concave opposite spiracles 
in medium-sized specimen and adult females, but much more deeply so In adult males ; 
outer and posterior corners broadly rounded, posterior margins evenly and moderately 



I 8o Memoir Sears Foundation for Marine Research 

convex; inner margins convex to axils. Axis of greatest breadtli about 70—72 "/o of 
distance back from tip of snout toward axils of pectorals. Tail with narrow lateral 
folds low down along posterior two-thirds, its length from center of cloaca to origin 
of first dorsal about i.i times as great as distance from center of cloaca to tip of snout 
in newly hatched specimen, decreasing relatively to about 0.85-0.95 that great in 
adults; distance from center of cloaca to tip of tail 1.2 -1.3 5 times as great as distance 
from cloaca to snout after embryonic prolongation is lost (4 specimens, 441-452 mm 
long)."9 

On newly hatched specimens, upper surface conspicuously rough; inner and an- 
terior parts of pectorals closely set with thorns, the largest posteriorly; 6—10 thorns 
on each shoulder region; 2-3 behind each eye, with two or more in front of it; 2-3 
along inner margin of orbits, and a few in space between orbits; a row along either 
edge of rostral cartilage, with a group of thorns of various sizes on tip of snout; also a 
regular mid-dorsal row of about 5 or 6 thorns on disc rearward from pectoral girdle and 
15—17 on tail, flanked by a lateral row on either side extending equally far rearward; 
another less regular row lower down along anterior half of tail. With growth the thorns 
increase in number to: 2—4 irregular rows along either side of midline of disc and of 
anterior part of tail; 1—2 rows near tip of tail; an irregular triangular group of about 
30-60 covering nuchal and shoulder region; 3—8 behind eye and 8—15 in front 
of it; 9—12 along inner margin of orbit and several between orbits; and an irregular 
double row along each edge of rostral cartilage, with a cluster at tip of snout. On the 
other hand, some of the large thorns are lost progressively from the inner parts of 
the pectorals, their anterior margins continuing rough with large thorns. Specimens 
from the Gulf of Maine and southward also lose the median row of thorns posterior 
to the nuchal region by the time a length of 300—350 mm is reached, but mature 
individuals (to at least 469 mm) that we have seen from the lower St. Lawrence River 
still bear this median row of thorns. Skin of disc and tail smooth between thorns in all 
specimens examined. Both dorsals prickly posteriorly in small specimens and prickly 
toward margins on larger. Pelvics prickly on small specimens; naked or with only a 
few small thorns on larger males and on partly grown females, but with a thorny 
area developing on mature females. 

At sexual maturity the outer posterior parts of the pectorals become increasingly 
thorny in females, but maturing males lose most of the thorns from the inner parts 
of their pectorals and some from the mid-dorsal ridge, so that they are noticeably 
smoother than mature females, but there is wide variation from specimen to specimen. 
Alar spines of mature males in 2—3 rows on outer two-thirds of pectoral; 13-14 
spines in each of inner two rows, 5—7 in third row. 

Lower surface wholly naked in newly hatched specimens, but snout soon becoming 
prickly, the rough area expanding as a narrow band along margin of disc about to level 
of nostrils in half-grown specimens, about to level of mouth in adults. 

119. In three specimens, 346-469 mm long, from the St. Lawrence River, the distance from cloaca to tip of tail was 
only 0.93-1. 18 as great as the distance from snout to cloaca. 



Fishes of the Western North Atlantic i 8 i 

Snout in front of orbits 1.9—2.4 times as long as distance between orbits, its 
length in front of mouth 1.5—2.0 times as great as distance between exposed nostrils. 
Orbit about i. 0-1.4 times as long as spiracle; distance between orbits 1.1-1.7 times 
as long as length of orbit. 

Distance between first gill openings 2.0—2.4 times as long as distance between 
exposed nostrils, between fifth gill openings i.i — 1.4 times; first gill openings i.o- 
1.6 times as long as fifth and about 18— 26''/o as long as breadth of mouth. 

Nasal curtain fringed; expanded posterior (outer) margin of nostril fringed finely 
or coarsely. Mouth broadly bowed, the lower jaw much more narrowly arched in adult 
males than in females, the upper jaw somewhat more so to accomodate it. 

Teeth ggZ^; low in young of both sexes and of females to maturity, rounded with 
faintly marked transverse cutting edges, in quincunx arrangement; those of mature 
males about as high as broad, with sharp or only slightly blunt points, more widely 
spaced in transverse series. 

First and second dorsals similar in size and shape, confluent at base. Caudal 
membrane from rear end of base of second dorsal about 27—35 "/o ^^ ^o^g ^^ base of 
first dorsal. ^^^ Pelvics moderately concave outwardly, scalloped around indentation; 
anterior margin about 51—64 % as long as distance from origin of pelvic to rear tip; 
anterior lobe broad with well rounded tip, including about five slender radial cartilages 
besides first stout one; posterior lobe with weakly and evenly convex outer margin and 
rather narrowly rounded tip. 

Claspers of mature males reaching about midway from axils of pelvics toward 
tip of tail. One of the terminal cartilages'^' is a broad scimitar-shaped blade with a 
sharp cutting edge, covered with thin skin, its recurved point directed forward along 
the outer ventral side of the clasper channel. This blade, which is entirely enclosed 
within the leaf-like walls of the clasper, is exposed after the latter has been inserted in 
the female. 

Anterior rays of pectorals reaching about 68—76 "/o of distance from level of 
fronts of orbits to tip of snout. 

Color. Upper surface grayish to dark brown or clouded with light and dark brown, 
usually with small round to oval darker spots varying in size and arrangement on 
different parts of disc and on different specimens. Lower surface usually white or pale 
gray without markings; disc and tail occasionally with irregular dusky blotches, or 
entire lower surface of tail dark gray.'^^ 

Size. Length at hatching is about 75—85 mm, the usual length at maturity about 
450-500 mm (18-20 in.) in both sexes; the largest specimen recorded was about 
530 mm long (21 in.).'" One 18 inches long weighs about 1.3 pounds. 

Developmental Stages. The eggs are often taken in considerable numbers in nets 

120. 8—20 "/„ as long in three specimens examined from the St. Lawrence River. 

121. Cartilage T', according to the terminology adopted by Jungersen (Danish 'Ingolf Exped., 2 [2], 1898: 10); often 
termed the thorn or spine. 

122. All these variations are illustrated on specimens in our Study Material. 

123. A specimen in the Harvard Museum of Comparative Zoology. 



I»2 



Memoir Sears Foundation for Marine Research 



or dredges in a few fathoms of water, and the empty cases cast up on the beach are 
familiar objects. They are amber or golden yellow when first laid but are almost black 
when found empty. The average dimensions are 55—63 mm long (exclusive of the 
horns) by 34-45 rnm broad. *2* The transverse margin at the end bearing the longer 
pair of horns 1^^ is straight or somewhat ragged, the margin at the end with the shorter 
horns either weakly convex, straight, or more or less concave; the two lateral faces are 
convex so long as the case contains its embryo. The horns taper to slender tips, and 
they are so fragile that ordinarily only the basal parts are intact on empty cases. The 
longer horns are nearly straight, the shorter pair more or less curved, one toward the 



i'^^^ 




Figure 38. Raja erlnacea, egg case, from Buzzards Bay, Massachusetts, about i.iX- 



other.12^ The lateral margins are fringed with thin hair-like elastic and adhesive fibers 
so tangled together as to form a sort of false membrane that accumulates sand or broken 
shells and serves to anchor the case. Respiratory slits do not exist at the time the egg 
is laid but are present before hatching (p. 142). 

In embryos nearly ready for release, the tail posterior to the second dorsal fin 
may be as long as the disc from snout to axil of pectoral; and in some young specimens 
the terminal prolongation may still be as long as the distance from the origin of the 
first dorsal to the rear end of the base of the second dorsal. But in others more recently 
hatched, as indicated by their smaller size, the tail may have shrunk nearly to its adult 
proportions. 1" Newly hatched specimens differ noticeably from aduks in their dermal 
armature (see p. 180). 

Habits. This Skate, like most others, is usually found on sandy or gravelly bottom, 
less often on mud and rarely on submarine rocks or ledges. Also, it is more plentiful in 

124. Daniel Merriman and Yngve H. Olsen advise us that in Long Island Sound the median length is as small as 46 mm 
and the median width as small as 27 mm. 

125. It is from this end that the young Skate is released. 

126. For photographs of egg cases, see Vladykov (Nat. canad., [3] 7, 1936: 213, 216) and Breder and Nichols (Copeia, 
1937: 182). 

127. For an account of the cleavage of the egg, see Ryder (Bull. U. S. Fish Comm. [1886], 6, 18S7: 8). 



Fishes of the Western North Atlantic 183 

water shoaler than 15—18 fathoms than in deeper water, while many come so close in 
shoreward that they are often found stranded on the beach, especially after rough 
weather. The greatest depths from which positively identified specimens have been 
recorded are 31-44 fathoms off the coasts of Virginia, Delaware, New Jersey, and Long 
Island, New York;i-* 71-80 fathoms off southern Massachusetts ;i2^ 30 fathoms on the 
southwestern part of Georges Bank;^^'' 25 fathoms near Seguin Island, Maine; and 50 
fathoms off the Bay of Fundy.'^' It tolerates a wide range of temperature, from about 
20-21° C (68-70° F)"2 in summer to as cold as about 3-4° C (37-39° F) in winter.'^^ 
Specimens have survived until heated to 29.1-30.2° C (84.4-86.4° F), even though 
taken from the cool water of Passamaquoddy Bay.^^* 

The maximum salinity recorded on bottom within the zone regularly inhabited 
by R. erinacca is 33.8 "/oo, the usual upper limit a little less than 33 "/oo- The great 
majority of the population lives in salinities lower than 32.5 %o the year round, but 
29-30 "/oo is about the lower limit of its optimum range, except in Long Island Sound, 
where it is found in water with salinity of only about 27 "/po. It has never been reported 
from fresh water, and specimens placed in the latter showed signs of great physiological 
disturbance. "5 

Stomach contents listed for R. erinacea include hermit and other crabs {Cancer^ 
Ovalipes, Panopaeus, Pagurus), shrimps, amphipods, copepods, annelid worms (chiefly 
Nereis), ascidians, bivalves including razor shells (Ense//a), and fragments of squids. 
Crabs are an important item, for 29 "/o of the Skates opened in one series of examin- 
ations at Woods Hole contained them, 15 "/o contained bottom-living shrimps of one 
kind or another, and 6 % had eaten squid. In Long Island Sound, amphipods were 
found to be the chief food, followed by crabs and shrimp (see above). It also preys 
on small fish, including the sand launce (Jmrnodytes), alewives (Pomolobus), herring 
(Clupea), cunners (Tautogolabrus), silversides (Menidid), tomcod (Microgadus), flounders 
{Paralichthys), and silver hake (Merluccius). 

It appears that R. erinacea produces eggs throughout its latitudinal range, except 
perhaps toward its southern limits, for egg cases have been found as far north as Halifax 
Harbor, Nova Scotia. "« For an account of fertilization and egg laying, see p. 188. 

R. erinacea probably does not carry out any extensive migrations along the coast 
with the change of seasons. It has been described repeatedly as coming up into shallower 
water for the summer and moving out into deeper water in autumn or early winter. 

128. Specimens collected by the U. S. Bureau of Fisheries steamer Albatross II and examined by us. 

129. Specimens trawled on the Albatross III. 

130. Ten specimens trawled on September 5, 1926, during one of the cod tagging cruises of the U. S. Bureau of Fisheries. 
It was not represented among the Skates taken in 37 trawl hauls at 40 fathoms or deeper on the northern part of 
the Bank in September 1929. For details of these catches, see Bigelow and Schroeder (Bull. U. S. Bur. Fish., 48, 
1936: 324). 

131. Huntsman, Contr. Canad. Biol. (1921), 3, 1922: 57. 132. Off New Jersey and southern New England. 

133. Shoal water off the eastern end of Long Island, New York, and off Connecticut; also southern side of Gulf of St. 
Lawrence. 

134. Huntsman and Sparks, Contr. Canad. Biol., N. S. 2, 1924: 102. 

135. Chaisson (Contr. Canad. Biol., Fisher., N. S. 5, 1930: 479). 

136. Vladykov, Nat. canad., 6j [(3)/], 1936: 213, 215. 



184 Memoir Sears Foundation for Marine Research 

But this seasonal shift in depth appears to be less than has sometimes been stated, for 
Little Skates are taken in winter in fish traps near New York in depths not greater 
than 20—30 feet, are trawled regularly throughout the winter off the Connecticut shore 
and off southern Massachusetts in 15—20 fathoms, and specimens tagged in June near 
Block Island, Rhode Island, were recaught nearby the following winter and spring.'" 
But it is also present in winter and spring in depths as great as 70—80 fathoms off 
southern New England and on southern Georges Bank where it probably is a year- 
round resident."* It has also been reported off North Carolina in winter in depths 
greater than 30 fathoms. ^^^ 

Numerical Abundance. This is not only the commonest Skate along the coasts of 
New England but the most familiar because of its habit of coming into shoal water. 
More exact information as to its abundance is found in the report that during one 
series of observations a trawler took an average of 98.8 pounds of them per hour in 
Long Island Sound, August 1943 to January 1944;"" during a second series, 1944- 
1946, 2i(> hauls yielded an average of around 200 specimens per haul;"^ and several 
hundred a day are sometimes landed at Woods Hole by draggers during the winter 
for preservation as teaching material for students of zoology. 

Relation to Man. This Skate is seldom used as food, but small numbers are utilized 
for baiting eel and lobster traps and some are used as zoological specimens. 

Range. Western North Atlantic, North Carolina to Nova Scotia and southern 
side of Gulf of St. Lawrence; from near shore out to about 80 fathoms. 

Details of Occurrence. The lower St. Lawrence River and southern shore of the 
Gulf of St. Lawrence,'*^ including the Bay of Chaleur, appear to be its most northerly 
outposts, for it has not been reported anywhere else within the Gulf, from Newfound- 
land (coastwise or offshore) or from Labrador. But it is described as "very common""^ 
all along the Atlantic Coast of Nova Scotia. It is plentiful along both shores of the Bay 
of Fundy and is generally distributed in suitable situations all around the northern and 
western coastline of the Gulf of Maine, including its larger bays (e. g., Massachusetts 
Bay) and interinsular passages. It is plentiful also along Cape Cod, on southwestern 
Georges Bank, off the southern coast of Massachusetts, around the islands of Nan- 
tucket and Marthas Vineyard, in Rhode Island waters, along the coast of Connecticut, 
on both shores of Long Island, near New York Harbor, and along New Jersey. It 

137. Reported by Daniel Merriman as a result of tagging experiments conducted in June 1946 by the Bingham Ocean- 
ographic Laboratory. 

138. We trawled one January 27, 1950 in 55 fathoms south of Block Island on the dragger Eugene H. On a cruise of the 
Albatross III, May 11-17, 1950, we took the following specimens in trawl hauls between Hudson Canyon and 
69° 10' W: 47 in 6 hauls, 20-30 fath.; 77 in 8 hauls, 31-40 fath.; 55 in 15 hauls, 41-50 fath.; 7 in 1 1 hauls, 51-60 
fath.; 8 in 6 hauls, 61-70 fath.; 18 in 9 hauls, 71-80 fath.; o in 8 hauls, 101-250 fath. 

139. Specimens reported by the Albatross III as taken in January 1950 in the vicinity of Cape Lookout, North 
Carolina. 

140. Merriman and Warfel, Trans. 9th N. Amer. Wildl. Conf. (1944): 234. 

141. Merriman and Warfel, Bull. Bingham oceanogr. Coll., II (4), 1948: 148, 157. 

142. Halkett, Check List Fish. Canada, 1913 : 42, "Gaspe Bay"; Cox, Bull. nat. Hist. Soc. New Brunsw. 13, 1S95: 63. 
There is also a specimen from Prince Edward Island in the American Museum of Natural History. 

143. Vladykov and McKenzie, Proc. N. S. Inst. Sci., J9, 1935: 51. 



Fishes of the Western North Atlantic 185 

is known from Delaware Bay, the coast of Maryland and northern Virginia, and it 
has been reported from deep water off Cape Lookout, North Carolina."* We have 
examined specimens "» that were trawled at stations scattered along the continental 
shelf southward to the offing of Chesapeake Bay in depths of 14—44 fathoms. In 
Chesapeake Bay it has been reported from Tangier Sound in the lower third of the 
Bay and at Hampton and Cape Charles, Virginia, near the mouth. 

Owing to its preference for shallow water (p. 183), the majority of the population 
of i?. erinacea is confined to a narrow coastline belt perhaps not more than 20—25 miles 
wide between the offings of Delaware Bay and New York, 10-12 miles wide along 
the southern Long Island shore, 40-60 miles wide off southern Massachusetts (in- 
cluding Nantucket Shoals) and widening to 60-80 miles or more to include the shoaler 
parts of Georges Bank (p. 184), and about 15-30 miles wide (measured from the 
nearest land) around the shores of the Gulf of Maine in general."' 

No information is available as to its presence or absence on the offshore Nova 
Scotian Banks. However, there are considerable areas shoaler than 20—30 fathoms on 
Sable Island Bank, and similar smaller areas exist on Browns and Banquereau Banks 
which may offer a suitable habitat for it unless the temperature there falls too low for 
its welfare during part of the year (see discussion, p. 183). 

Synonyms and References: 

Raid eglantiera^'^'' Lesueur, J. Acad. nat. Sci. Philad., 4 (i), 1824: 103 (descr. of Philad. specimen ident. as 
R. erinacea by dermal armature and small size of male with large claspers). 

Raja erinaceus'^'^^ Mitchill, Amer. J. Sci., 9, 1825: 290 (descr., ill., New Jersey; type specimen sent to Paris); 
DeKay, Zool. N. Y., 4, 1842: 372, pi. 78, fig. 246 (ill., New Jersey); Storer, Mem. Amer. Acad. Arts 
Sci., N. S. 2, 1846: 512; Synopsis Fish. N. Amer., 1846: 259, 260 (diagn., size. New Jersey); Perley, 
Cat. Fish. New Brunsw. and Nova Scotia, isted., 1851: 39 (Grand Manan I.); Rep. Fish. Bay of 
Fundy, 1851: 155 (Grand Manan I.); Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861: 61 (listed, 
Atlant. Coast N. Amer.); Lyman, 6th Rep. Mass. Comm. inl. Fish., 1872: 54, 57 (south. Mass., Conn.); 
Baird, Rep. U. S. Comm. Fish. (1871-1872), j, 1873: 826 (listed. Woods Hole); Gill, Rep. U. S. 
Comm. Fish. (1871-1872), J, 1873: 812 (listed, C. Cod to Florida); Garman, Proc. Boston Soc. nat. 
Hist., 77, 1874: 176 (descr., Mass. Bay and mouth of Connecticut R.); Goode and Bean, Bull. Essex 
Inst., II, 1879: 28 (Mass., but R.ocellata also included); Bean, Proc. U. S. nat. Mus., J, 1880: 115 
(Mass. and Conn.); Jordan and Gilbert, Bull. U. S. nat. Mus., 16, 1883: 40 (descr., size, nos., range); 
Bean, Rep. U. S. Comm. Fish. (1882), 10, 1884: 343 (listed. Woods Hole); Rathbun, Proc. U. S. nat. 
Mus., 7, 1885: 485 (parasites. Woods Hole); Jordan, Rep. U. S. Comm. Fish. (1885), 13, Append. E, 
1887: 799(listed,N. Atlant. states); Ryder, Bull. U. S. Comm. Fish. (1886), 6, 1887: 8 (early embryoL); 
Linton, Rep. U. S. Comm. Fish. (1886), 1889: 460, 462 (parasites); Rep. U. S. Comm. Fish. (1887), 
1891: 766, 840 (parasites); Goode and Bean, Smithson. Contr. KnowL, JO (Publ. 981), 1895: 28; 

144. It was credited to South Carolina by True (List Vert. Anim. S. Carolina, in Handb. S. Carolina, 1883: 261), but 
seemingly this harks back to the fact that it was confused with K. eglanteria by Lesueur (J. Acad. nat. Sci. Philad., 
4, 1824: 103, 105). Nor do we find any factual basis for Gill's (Rep. LT. S. Comm. Fish. [1871-1872], 7, 1873 : 812) 
report of it as occurring "to Florida." 

145. Some in the U. S. National Museum, others collected during the cruises of the U. S. Bureau of Fisheries steamer 
Albatross IL 

146. Twenty-two trawl hauls, made by the Atlantis in August 1936 in the deeper troughs farther out in the Gulf of 
Maine, did not yield a single K. erinacea, though three other species {K. senta, R. radiata, and R. lacvis) were taken. 

147. Obvious misspelling for eglanteria. 

148. The spelling was corrected to "erinacea" by Storer (1846) who, with some authors, has preferred the spelling Rata 
to Raja. 



86 Memoir Sears Foundation for Marine Research 

Spec. Bull. U. S. nat. Mus., No. 2, 1895: 28; Mem. Harv. Mus. comp. Zool., 22, 1896: 28 (New 
England); Cox, BuU. nat. Hist. Soc. New Brunsw., JJ, 1895: 63 (Grand Manan I., and S. Gulf St. 
Lawrence); Jordan and Evermann, Rep. U. S. Comm. Fish. (1895), 1896: 221 (listed, Virginia to 
Maine); Bull. U. S. nat. Mus., 47 (i), 1896: 68 (descr., Virginia to Maine); Bean, Bull. Amer. Mus. 
nat. Hist., 9, 1897: 331 (egg cases. New York); Linton, Proc. U. S. nat. Mus., 20, 1898: 423, 440, 
450 (parasites. Woods Hole); Bumpus, Science, N. S. 8, l8g8: 851 (season with eggs. Woods Hole); 
Smith, Bull. LT. S. Fish Comm. (1897), ij, 1898: 89 (nos.. Woods Hole); Jordan and Evermann, Bull. 
U. S. nat. Mus., 47 (4), 1900: pi. 9, fig. 29 (ill.); Bean, 6th Rep. For. Comm. N. Y., 1901 : 382 (season, 
range); Gorham, Bull. U. S. Fish Comm., jp, 1901: 34 (physiol.. Woods Hole); Linton, Bull. U. S. 
Fish Comm. (1899), ig, 1901: 274 (stom. contents); Bull. U. S. Fish Comm. (1899), ig, 1901 : 430 
(food, parasites); Bean, Bull. N. Y. St. Mus., 60, Zool. 9, 1903 : 47 (descr., habits, egg cases. New York); 
Sharp and Fowler, Proc. Acad. nat. Sci. Philad., ^6 (2), 1904: 506 (Nantucket); Parker, Bull. U. S. Bur. 
Fish. (1904), 24, 1905: 190, 196, 197, 202 (physiol. lateral line syst.); Wilson, Proc. U. S. nat. Mus., 
28, 1905: 571, 573, 575, 631 (parasites); Bull. U. S. Bur. Fish. (1904), 24, 1905: 131 (parasites); 
Fowler, Rep. N. J. Mus. (1905), Pt. II, 1906: 70 (descr.. New Jersey); Tracy, 36th Rep. R. I. Comm. 
inl. Fish., 1906: 47 (descr., food, breeding season, Rhode Island); Field, Rep. U. S. Comm. Fish. (1906), 
Spec. Pap. [6], Doc. 622, 1907: 23 (descr., food. Woods Hole); Fowler, Rep. N. J. Mus. (1906), Pt. 
Ill, 1907: 257 (descr.. New Jersey); Rand, Amer. Nat., 4.1, 1907: 287 (spiracular breathing); Sullivan, 
Bull. U. S. Bur. Fish. (1907), 2^, 1908: 13 (physiol. digestive syst.); Fowler, Rep. N. J. Mus. (1907), 
Pt. Ill, 1908: 128 (color, size, Delaware Bay); Gill, Smithson. misc. CoU., ^2, 1910: 159, fig. 46 (ill.); 
Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 10 (New England locals.); Smallwood, Science, 
N. S. 28, 1908: 930 (abnormality); Sumner, Bull. U. S. Bur. Fish. (1908), 28 (2), 1910: 1246 (distrib. 
off Marthas Vineyard); Fowler, Proc. Acad. nat. Sci. Philad. (1909), 61 (2), 1909: 407 (Delaware 
Bay); Proc. Acad. nat. Sci. Philad., 62 (2), 1910: 470 (Maine, Mass., and New jersey); Tracy, 40th 
Rep. R. I. Comm. inl. Fish., 1910: 61 (breeding season, nos., Rhode Island); Fowler, Proc. Acad. nat. 
Sci. Philad., 64 (1), 1912: 57 (Virginia); Denis, J. biol. Chem., 16, 1913-14: 390 (blood chemistry); 
Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 4 (3), 191 3: 102 (range); Garman, Mem. Harv. Mus. 
comp. Zool., j6, 1913: 337, pi- 20, pi. 55, fig. 5, pi. 68, fig. I (descr., anat., Halifax to Carolinas); 
Halkett, Check List Fish. Canada, 1913: 12, 42 (listed, Gulf St. Lawrence to Virginia); Nichols, 
Abstr. Linn. Soc. N. Y., 1913: 91 (New York); Sumner, Osburn and Cole, Bull. U. S. Bur. Fish., 
JJ (i), 1913'. 74, 162, 168, 417; JT (3), 1913: 737 (breeding season, food. Woods Hole); Ken- 
dall, Proc. Portland Soc. nat. Hist., J (i), 1914: 12 (Maine locals.); Welsh, in Bigelow, Bull. Mus. 
comp. Zool. Harv., ^8, 1914: in (Seguin I., Maine, 25 fath.); Fowler, Copeia, 13, 1914: 2 (New 
jersey); Proc. Acad. nat. Sci. Philad., 66 (2), 1914: 353 (Delaware); Copeia, 27, 1916: 12 (New Jersey); 
Copeia, 31, 1 9 16: 41 (New Jersey); Proc. Boston Soc. nat. Hist., J5, 191 7: in (Maine and Woods 
Hole); Latham, Copeia, 41, 1917: 17 (season. Long Island, New York); Huntsman, Trans, roy. Soc. 
Canad., (3) 12 (4), 1918: 63 (temp., Gulf St. Lawrence); Fowler, Occ. Pap. Mus. Zool. Univ. Mich., 
56, 1918: 2 (New Jersey); Copeia, 68, 1919: 13 (season. New Jersey); Fowler, Proc. Acad. nat. Sci. 
Philad., "JI, 1920: 293 (New Jersey); Proc. biol. Soc. Wash., JJ, 1920: 145 (New Jersey); Townsend, 
N. Y. zool. Soc, 25th Annu. Rep. (1920), 1921 : 126 (New York Bay); Huntsman, Contr. Canad. Biol. 
(1921), 3, 1922: 57 (Bay of Fundy and Passamaquoddy Bay); Breder, Copeia, 127, 1924: 28 (Sandy 
Hook Bay); Huntsman and Sparks, Contr. Canad. Biol., N. S. 2, 1924: 102 (lethal temp.); Linton, 
Proc. U. S. nat. Mus., 64 (21), 1924: 9, 61, 64, 66, 83 (parasites); Scott, Contr. Canad. Biol., N. S. 
2, 1924: 131 (pathol. thyroid); Bigelow and Welsh, Bull. U. S. Bur. Fish. (1924), 40 (i), 1925: 58 
(descr., ill., habits. Gulf of Maine); Battle, Trans, roy. Soc. Canad., (3) 20 (5), 1926: 127, 132 (effects 
of high temp.); Huntsman, Trans, roy. Soc. Canad., (3) 20 (5), 1926: 202, 203 (lethal temp.); Breder, 
Zoologica N. Y., 4 (5), 1926: 244 (listed); Manter, Illinois biol. Monogr., 10 (2), 1926: 8, no (tre- 
matode parasites); Nichols and Breder, Zoologica N. Y., 9, 1927: 23 (descr., habits, breeding season, 
New York and south. New England); Borodin, Bull. Vanderbilt oceanogr. (Mar.) Mus., I (i), 1928: 
5 (listed, no local.); Fowler, Fish Culturist, 7, 1928: 225 (New Jersey); Hildebrand and Schroeder, 
Bull. U. S. Bur. Fish., 42 (i), 1928: 60 (descr., Hampton and C. Charles, Virginia); Burwash, Contr. 
Canad. Biol., N. S. 4, 1929: 117 (iodine in thyroid); Battle, Contr. Canad. Biol., N. S. 4, 1929: 497 
(reflexes at high temp.); Contr. Canad. Biol., N. S. 4, 1929: 503 (physiol.); Breder, Field Bk. Mar. 
Fish. Atlant. Coast, 1929: 29 (descr., Carolinas to Halifax, Nova Scotia); Jordan, Manual Vert. Anim. 



Fishes of the Western North Atlantic 187 

NE U. S., 1929: 16 (diagn.); Truitt, Bean and Fowler, Bull. Md. Conserv. Dep., 3, 1929: 30 (descr., 
ill., Atlant. Coast Maryland, Chesapeake Bay); Bere, Contr. Canad. Biol., N. S. 5, 1930: 426 (copepod 
parasites, Passamaquoddy Bay); Chaisson, Contr. Canad. Biol., N. S. 5, 1930: 477 (effects of fresh water, 
Passamaquoddy Bay); Jordan, Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 25 (com- 
mon names, listed, Virginia to Maine); Lutz, Amer. J. Physiol., g4, 1930: 136 (effects of adrenalin 
on auricle); Marshall, Amer. J. Physiol., 9./, 1930: 7 (excretion); Huntsman, Contr. Canad. BioL, 7 (4), 
1931 : (4) 34 (heart physiol.); Lutz, Biol. Bull. Woods Hole, 61, 1931 : 94 (nerves, action of adrenalin); 
Smith, Amer. J. Physiol., g8, 1931: 285 (ref. to Marshall, 1930); Breder, Copeia, 1932: 31 (Block 
Island); NichoUs, Contr. Canad. Biol., N. S. 7, 1933: 449 (gastric mobility); Contr. Canad. Biol., N. S. 
8, 1933: 139 (temp, effects); Bigelow and Schroeder, Canad. Atlant. Fauna, I2e, 1934: 21 (descr., 
ill., range); Daniel, Elasmobranch Fishes, 1934: 9 (external form); Parker, Annu. Rep. Woods Hole 
oceanogr. Inst. (1933), 1934: 27 (color changes); Gudger, J. Morph., =)J, 1935 : 99 (maxillary respiratory 
curtains, by ref to Rand, 1907); Vladykov and McKenzie, Proc. N. S. Inst. Sci., jp, 1935: 50, 51 
(diagn., ill.. Nova Scotia); Bigelow and Schroeder, Bull. U. S. Bur. Fish., ^5, 1936: 324 (in Key); Smith, 
Biol. Rev., II, 1936: 52, 62, 67 (osmotic pressure); Vladykov, Nat. canad., 6j [(3) 7], 1936: 213 (descr., 
ill., egg case); Breder and Nichols, Copeia, 1937: 182 (meas. egg case); Fowler, Proc. Acad. nat. Sci. 
Philad. (1937), 8g, 1938: 304 (New Jersey); Breder, Bull. N. Y. zool. Soc, 41, 1938: 28 (season, New 
York Harbor); Merriman and Warfel, Trans. 9th N. Amer. Wildl. Conf., 1944: 234 (nos. Long Island 
Sound); Bull. Bingham oceanogr. Coll., II (3), 1948: 134, 148, 155, 157 (common names, sizes, nos., 
season, off Connecticut and Rhode Island). 

Raia^'^^ eglanteria (in part) DeKay, Zool. N. Y., 4, 1842: 373 {R. eglantiera Lesueur 1824 included, equals 
R. erinacea Mitchill 181 5); Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 512 (R. eglantiera 
Lesueur 1824 included, equals R. erinacea Mitchill 181 5); Gill, Proc. Acad. nat. Sci. Philad. (1861), 
Addend., 1862: 61 {R. eglantiera Lesueur 1824 included, equals R. erinacea Mitchill 181 5); Dumeril, 
Hist. Nat. Poiss., j, 1865: 532 (R. erinacea Mitchill 181 5 included, also R. eglantiera Lesueur 1824, 
equals R. erinacea Mitchill 181 5); Giinther, Cat. Fish. Brit. Mus., 8, 1870: 462 {R. erinacea Mitchill 
1 81 5 included). 

Common Skate, Wilson, Bull. U. S. nat. Mus., 158, 1932: 403 (copepod parasite, Woods Hole). 

Summer Skate, Wilson, Bull. U. S. nat. Mus., 158, 1932: 412 (copepod parasite. Woods Hole). 

Probable Synonyms: 

Raia diaphanes (in part) Storer, Mem. Amer. Acad. Arts Sci., N. S. 9, 1867: 240, pi. 39, fig. i; Fishes Mass., 

1867: 264, pi. 39, fig. I {R. erinacea probably included). 
Not Raja eglanteria Bosc 1802, in Lacepede, Hist. Nat. Poiss., 4° Ed., in Buffon, Hist. Nat., 2, 1802: 104, 

log, pi. 4, fig. 2. 
Not Raja diaphanes Mitchill, Trans. Lit. philos. Soc. N. Y., J, 181 5: 478 (equals R. eglanteria Bosc 1802; 

see p. 176). 
Not Raia diaphanes DeKay, Zool. N. Y., 4, 1842: 366, pi. 67, fig. 218 (equals R. ocellata Mitchill 181 5: see 

p. 251). 
Not Raja erinacea Ribeiro, Arch. Mus. nac. Rio de J., 14, 1907-. 176, pi. 13 (a Psammobatis, renamed extenta 

by Garman [Mem. Harv. Mus. comp. Zool., j6, 1913: 356], SE coast S. Amer.). 

ADDENDUM TO RAJA ERINACEA 
By 

Daniel Merriman, Yngve H. Olsen, Sarah B. Wheatland and 

Louva H. Calhoun 

Bingham Oceanographic Laboratory^ Tale University 

At the time this volume went to press we were conducting an intensive study of 
Raja erinacea. Since the results of our work were not sufficiently advanced for full 

149. Also spelled Kaja. 



I 8 8 Memoir Sears Foundation for Marine Research 

publication in the Bulletin of the Bingham Oceanographic Collection and hence were 
not available to the authors of this volume, it was suggested that a summary of our 
findings to date, in the form of an Addendum, would be useful. Although our studies 
have been carried on intermittently since 1943 and intensively in 1950, it is apparent 
that much of the life history of the Little Skate still rests on highly tentative ground, 
and therefore some of the conclusions put forth below remain to be confirmed, modified, 
or disproved by further observation and study. 

Our results have been derived chiefly from a study of collections obtained monthly, 
or even more frequently, on a year-round basis from commercial draggers working 
off the southern New England Coast, particularly in Block Island and Long Island 
Sounds; in 1950 over 15,000 specimens were examined in detail. We have also been 
fortunate in having large outdoor tanks and laboratory facilities available to us at the 
U. S. Fish and Wildlife Service Station at Milford, Conn., through the generous 
cooperation of Victor L. Loosanoff. 

In adult specimens, mating takes place throughout the year. This is clearly in- 
dicated both by the sperm plugs in the uterus and by fresh wounds at the fundus of 
the uterus, the latter being inflicted during copulation by the scimiter-shaped cartilage 
or "sentinel" of the clasper. In our samples to date, not less than 30 "/q of the adult 
females have shown wounds of this sort and a figure as high as 70 "/o has been recorded 
in one sample; in general, then, at any time during the year a rough average of 50 % 
of the adult females show fresh and often severe uterine lacerations — incontrovertible 
evidence of recent mating. Since copulation is probably effected at times without so 
wounding the female, it is a fair assumption that mating takes place frequently throughout 
the year. Sperm are stored in large numbers in the convolutions of the shell gland, as 
demonstrated by sectioning; we have no idea of the viability of sperm under these con- 
ditions, but it would not seem to be a matter of importance in the reproduction of this 
species in view of the frequency of mating. Indeed, perhaps there should be a revision 
of the general conception that sperm are "stored" in the shell glands of other species 
of Skates for the purpose of future fertilization; at least in erinacea, the whole female 
genital tract apparently contains sperm so consistently that it is no wonder that some 
find their way into the convolutions of the shell gland, a level which they must attain 
to accomplish fertilization. 

Fertilization in adult females, as well as the production and laying of egg cases, 
takes place the year around. However, there are two periods when egg-laying is at a 
maximum, as judged by the percentage of pregnant fish in our samples. From late 
October to early January this figure averages between 30 and 50 "/o and has run as 
high as 59 % in one sample of Block Island Sound fish. Again in June and July, 25— 
30 % of the adult females from these waters are pregnant. In other months of the year 
less than 25 "/o are pregnant, a rough average for these "off" months for Block Island 
Sound fish being 10 "/q, with the lowest level of production coming in August and 
September. The same general pattern obtains in Long Island Sound samples, with 
peaks of 10—15 "/o pregnancy in the November— January and June— July periods and 



Fis/ies of the JVester?i North Atlantic 189 

considerably lower percentages of pregnancy in other months of the year; in these 
latter waters the percentage of pregnancy is always much lower than that in Block 
Island Sound fish. There is considerable variation from sample to sample, but in general 
the above pattern is remarkably consistent. We believe that the seasonal nature of 
breeding as outlined is the true situation, and we have much evidence to show that 
this picture does not arise from the vagaries of sampling by commercial fishing vessels — 
i. e., the tendency to fish possible spawning grounds of Skates in certain seasons (June- 
July and November— January) and to avoid those grounds in other months. Incidentally, 
in this regard the percentage of females with uterine wounds shows no striking cor- 
relation with the seasonal nature of productivity, although there may be a slight increase 
in the' percentage two to four weeks before the peaks of productivity. 

Ovulation takes place in both ovaries at approximately the same time, and the 
formation of the egg case by each of the two shell glands begins before either of the 
eggs has reached that level in its passage down its respective oviduct. By the time the 
cases are approximately V4— Vs formed, single eggs enter each after being much elongated 
in passing through the shell gland. Following the completion of egg-case formation, 
activity in the right arm of the genital tract invariably precedes that in the left arm. 
Thus, while eggs are always prepared for laying in pairs, the right egg descends and 
reaches the uterus first and is laid slightly in advance of the left egg (two exceptions 
out of thousands examined); we have detected no anatomical difference to account for 
this phenomenon. At one end the egg case always has two long straight tendrils, at 
the other end two shorter tendrils whose terminal portions are likely to be recurved; 
the end with the longer tendrils is laid down first by the shell gland. As a result of 
aquarium experiments and palpating techniques, we find that egg cases are formed 
and laid in a surprisingly short period of time, the entire process in June and July 
occurring in 48—72 hours. In the winter months the process is undoubtedly slower, 
and a female may retain fully formed egg cases for as much as three weeks. It appears 
that a few eggs in each ovary become ripe at approximately the same time and that 
these are laid in pairs at intervals from as little as five days to several weeks in the early 
summer production peak. Thus one of our aquarium fish laid four pairs of egg cases 
between June 7th and July 1 2th, i 950. Whether a mating is requisite for the fertilization 
of each successive pair of eggs or whether one copulation may serve to effect fertilization 
in several successive pairs of eggs is not known, since we had both sexes in our ex- 
perimental tanks. Following the period of egg-laying, it appears likely that the whole 
genital tract undergoes somewhat of a regression, probably prior to building up the 
ovaries to a condition where they once again contain ripe eggs {vide infra). On the 
other hand, adult males carry sperm continuously at all times of year from the onset 
of maturity to death. 

In our tank experiments, 12 females produced 65 egg cases between May 30th 
and July 12th, 1950, an average of over three pairs per fish. Of these 65 egg cases, 
only 10 (15 %) were found to contain living embryos; the others were either infertile 
or failed to survive past an early stage of development. This low percentage may have 



190 Memoir Sears Foundation for Marine Research 

been due to tank conditions, but it is interesting that it compares almost exactly with 
the results obtained by Clark^ with R. brachyura kept in aquaria at Plymouth. These 
data suggest the possibility that in nature not more than 10—25 % of the eggs laid 
are fertile or develop beyond an early stage. In this connection, exceptions to the rule 
that each egg case contains only a single egg or embryo in this species are rare. 
From the many specimens we have examined we have found two eggs in a single egg 
case twice, and four eggs in one case once; all three anomalies were detected in pregnant 
fish, and in each instance the matching egg case was empty. 

Hatching of eggs laid in the May— July period and maintained under aquarium 
conditions took place from November 29th, 1950 to January ist, 1951, that is, five to 
six months after they were laid. This is probably the shortest time in which hatching 
takes place in nature, since the water temperatures in the aquaria were somewhat 
above those of the normal environment over the entire period and since eggs laid in 
June and July would be subject to optimum temperatures for the fastest development 
in these waters. Those laid in the November— January period probably take consider- 
ably longer to reach hatching. It seems probable that the eggs of R. erinacea take a 
minimum of six months and up to nine months or more after laying to arrive at the 
hatching stage in nature. 

At hatching, Skates from Long Island Sound waters are 9.5-10.0 cm in total 
length and the yolk in the abdomen is still apparent. As this time the tip of the tail 
has a fine, distinctively pigmented, whip-lash elongation which disappears after a few 
days; thus the distance from the base of the second dorsal to the end of the tail is 
1. 4-1. 5 cm at hatching and only 0.925-0.950 cm several weeks later when this fil- 
amentous piece has sloughed off. It seems quite possible that this 5 mm elongation is 
directly concerned with hatching; in short, it may be comparable in function to the 
egg-tooth in birds and similar devices in some Amphibia. Within the egg case before 
hatching the Skate is oriented so that its head points toward the end of the case pos- 
sessing the long straight tendrils. The transverse margin of the case at this end is 
straight and sometimes frayed, and the two broadly convex surfaces of the case meet 
approximately 5 mm from the margin to form a flat area at that end. It is via the separ- 
ation of the two surfaces of the egg case in this area that the Skate escapes, and after 
hatching this marginal area resembles an envelope which has been opened by a sharp 
paper cutter. Indeed, the opening through which the fish has emerged would not be 
detected by casual observation. Before hatching, the tail is curled around in a broad 
arc so that its tip is above the head and extends into the region which later becomes 
slit for the emergence of the young fish. It seems probable from observation and 
inference that the fine extension of the tail acts like a flagellum and eventually wears 
the case open in the slit-like manner described above. Apertures in the tendrils, through 
which the water presumably circulates freely, are present before hatching; apparently 
these "respiratory slits" do not exist at the time the egg is laid and we do not know 
at precisely what stage of development they appear. 

I. J. Mar. biol. Ass. U. K., N. S. 12, 1920: 577-643. 



Fishes of the Western North Atla?it'tc 191 

Determination of growth rates and ages at different lengths after hatching pre- 
sents a major problem. Obviously the analysis of length-frequency curves is fraught 
with difficulty in a fish which breeds on a year-round basis, even though the species 
has peaks of productivity in June— July and November-December. Study of the hard 
parts by sectioning and clearing techniques is also unrewarding, although examination 
of whole vertebrae after subjecting them to various treatments shows considerable 
possibilities for age determination. The return of tagged fish has been too limited to 
be of use in checking growth estimates, and aquarium fish have not provided any data. 
Furthermore, we have taken relatively few small individuals in our collections. Piecing 
together all the fragmentary information at our disposal, it seems probable that this 
species grows at an average rate of 8-10 cm a year for the first three years following 
hatching and more slowly thereafter. Thus a 20 cm Skate might be i — 1.5 years old, 
a 30 cm individual 2—3 years of age, a 40 cm Skate 3-4 years, a 45 cm fish 4-5 and 
possibly 6 years, and a 50 cm individual 6—7 and perhaps 8 years old. Specimens 
above 50 cm are comparatively rare and the great majority are males; the largest we 
have recorded is 54 cm. In monthly samples of Block Island Sound fish the length- 
frequency curves show the peak of abundance to be between 43 and 46 cm; above this 
point the curve falls away very steeply, thus indicating a high mortality rate after the 
fish are approximately five years old. 

In general, female specimens are adolescent in the range between 32 and 43 cm; 
we have found fish up to 42.5 cm which were still immature, as judged by the shell 
gland, size of the uterus and condition of the ovaries, and we have found fully mature 
individuals as small as 36 cm. Males tend to reach adolescence at a somewhat greater 
size, the general range being from 36 to 45 cm; we have found immature males as large 
as 44 cm and fully mature males at 37 cm. 

With respect to the average length and weight of adolescent and fully mature 
males and females, there appear to be significant differences between the Long Island 
and Block Island Sound populations: 

Adolescent Adolescent Mature Mature 

females males females males 

Long Island 38.4 cm 40.0 cm 42.4 cm 43.1 cm 
344 g 338 g 475 g 455 g 

Block Island 42.0 cm 42.2 cm 45.8 cm 46.4 cm 
462 g 422 g 596 g 587 g 

These figures represent yearly averages, and it is abundantly clear from them that 
either the Long Island Sound fish mature earlier than those from Block Island Sound 
or they grow more slowly and perhaps mature at approximately the same age as their 
counterparts to the east. We suspect that the latter alternative is the true situation. 
Continuing our comparison of samples of trawl-net catches from the two areas, 



192 Memoir Sears Foundation for Marine Research 

we find that in Long Island Sound the majority of Skates range from 39-44 cm, with 
a few up to 47 cm in length. In the Block Island Sound samples the majority of fish 
fall between 42 and 48 cm with a few from 49—54 cm. In short, on an average the 
Long Island Sound erinacea of both sexes are adolescent at a lesser length and mature 
at a lesser length than their Block Island Sound counterparts; further, the peaks of 
length-frequency curves of fish from Long Island Sound are consistently a good 3 cm 
less than the peaks of those from Block Island Sound samples, and the Long Island 
Sound fish never attain the uppermost size limit of the largest ones from Block Island 
Sound, i. e., up to 48—54 cm. Considering this, and the previously mentioned fact 
that Long Island Sound erinacea are much less prolific, plus the fact that there is ap- 
parently no substantial intermingling between the two areas, it is rather clearly indicated 
that these are two essentially separate populations. The one from Long Island Sound 
we think of as poor, slow-growing, and relatively unproductive; the other, from Block 
Island Sound and seaward, we think of as the more normal, healthy, faster-growing, 
and comparatively productive population. Further evidence to support this contention 
is found in the fact that at all times of year comparable trawl hauls in Block Island and 
Long Island Sounds yield many more fish in the former area than in the latter; the 
former appears to be a flourishing population, the latter a sparse and rather unsuccessful 
one. Comparing the Block Island and Long Island Sound populations further, the sex 
ratios show peculiar differences. In Block Island Sound, males dominated the adult 
population (60— 80 "/o) throughout 1950 (except for March-April), while in Long 
Island Sound, females dominated the adult population (70 "/q) throughout 1950 (except 
in March— April). In both areas the females dominated the immature population (at 
least 70 "/o) with few exceptions. Morphometric comparisons between the two popul- 
ations will almost certainly show significant differences, and apart from occasional 
strays and a minor amount of intermingling at the adjacent boundaries of the two areas, 
the Long Island and Block Island Sound erinacea may well be considered two separate 
races. 

In this connection, if this species should be fished hard commercially for fish 
meal or other purposes (Vitamin A is low, approximately 100 units per gram of liver 
oil), we have every reason to believe that the catch-per-unit-of-effort would decline 
rapidly; in short, the Little Skate would not be capable of yielding a steady supply 
over long periods. Leaving the Long Island Sound population out of consideration 
because of its paucity, the Block Island Sound fish command our attention because 
they are probably more representative of this species over its whole range. From what 
we have indicated previously of growth rates, sharp mortality above 46 cm, age at 
maturity, percentage of fertilization, survival to hatching, etc., it would appear that this 
species has no great capacity to increase the present size of the total population ; it 
is clearly not a fish which is subject to marked fluctuations in abundance from natural 
causes. Correspondingly, a reduction in the numbers of sexually mature fish by in- 
creased fishing effort would shortly lead to a reduction in the number of young that 
would be added to the stock each year. If only one out of every four to ten egg cases 



Fishes of the Western North Atlantic 193 

reaches a hatching stage {vide supra), it is clear that in nature each female would have 
to lay more than the maximum observed by us in aquarium experiments (eight) to 
maintain the stock. If the stock is to be held at the same level, each female must pro- 
duce two offspring which attain sexual maturity. We suspect that mortality after 
hatching is relatively low, but if our rough estimate of the percentage of eggs which 
reach hatching is of the right order of magnitude, this would mean that each female 
would have to lay a general minimum of 10-20 eggs; actually, each one probably lays 
20—30 eggs under average conditions. Whether this occurs consecutively, once ma- 
turity is reached, or whether there is a resting period between successive ovulations, 
we do not know. However, we strongly favor the latter alternative. We find many 
large-sized females with reduced shell glands and ovaries, the majority of which are 
probably in a resting stage prior to another ovulation. Also, the ripe ovary never con- 
tains at one time more than about three eggs that have reached or are approaching the 
laying size. We suspect, therefore, that a typical female produces on the order of five 
to six eggs over a laying period, say the June-July peak, and that the metabolic strain 
of producing these large-yolked eggs is such that the female requires a number of 
months to build up to a condition where she can lay another comparable batch of 
eggs. According to our estimates, each female would have to have at least two or three 
such laying periods in order to maintain the population, and some may have as many 
as four. From our knowledge of the age at maturity, the rate of growth of these older 
fish (roughly 4—5 cm a year as judged from length-frequency and other data), the 
sharp mortality rate at greater sizes, and from our estimate that there are a number 
of months between laying periods, we derive a time schedule which would allow for 
three or four such laying periods in the life of the average female erinacea. Because 
of the nature of this reproductive pattern, we do not believe that the stock could stand 
intensive fishing without showing a sharp reduction in the size of the population in a 
relatively short period and a consequent drop in the catch-per-unit-effort, a situation 
which has occurred in other elasmobranch fisheries. 

Marking experiments have yielded little information due to the low percentage 
of return, which we attribute chiefly to the fact that the V2" disc tags we used were not 
obvious enough to catch the attention of the fishermen as the Skates were shovelled 
overboard as trash or barrelled in bulk for fish meal or other uses. Out of approximately 
600 erinacea tagged in June 1946 in the Block Island Sound area, only five fish were 
returned: one in July 1946; two in August 1946; another in February 1947; and the 
last in April 1947. All of these recaptures were within five miles or less of the original 
point of release except the last, which was taken approximately 50 miles to the west, 
off Herod Point on the north shore of Long Island. If we can judge by such meager 
returns, it would seem that this species undertakes no extensive migrations in this 
region, although the specimen captured in Long Island Sound would indicate occasional 
straying between the Block Island and Long Island Sound populations. However, there 
does appear to be some evidence of definite seasonal offshore and onshore movements 
in the Block Island Sound region involving distances of '^—c, miles and upward. This 



194 Memoir Sears Foundation for Marine Research 

evidence, from conversations with local fishermen and from trawl catch samples, 
indicates that in general the bulk of the Skates move inshore in the spring, offshore in 
the middle or late summer, inshore again in the late fall and offshore again in mid- 
winter. Our data on relative abundance in different seasons and at different localities, 
as judged by trawl catch samples, correlates well with this apparent movement. Note, 
too, that the inshore movements fit well with the spawning peaks in the late spring 
and early winter mentioned above. From this and other evidence we think it probable 
that this species of Skate customarily spawns in relatively shallow water (15 fathoms 
and less) on a hard sandy bottom. Observations on aquarium fish indicate that the eggs 
are usually at least partly buried in the sand. 

Turning to the matter of food eaten by Skates in this region, the amphipod, 
Leptocheirus pinguis, is the dominant element, constituting anywhere from a third to 
a half of the volume of the stomach contents at all times of year; Cancer irroratus follows 
at one-fifth, and Crago septemspinosus at a tenth. 

Although there is considerable variation in the weight at any given length, in 
general Block Island Sound fish weigh 0.5 lb. when they are 12^/4-13 in. long, 0.75 lb. 
at 15 in., i.o lb. at 16V4 in., 1.25 lb. at 18 in., and 1.5 lb. at 19V4 in. The largest 
fish, about 21^4 in., weighed roughly 2.0 lbs. 

Since hermaphroditism appears to be rare in elasmobranch fishes, we record here- 
with the capture of a specimen of R. erinacea which shows this phenomenon in rather 
extreme form. This fish, taken by a commercial dragger on July 18, 1950, about three 
miles south of Fisher's Island, N.Y., was 41.5 cm in length and 408 g in weight. On 
the left side this individual was a fully developed male with large testis, vas deferens, 
and a completely adult and apparently functional clasper with the typical prominent 
clasper gland at its base. On the right side there was a tiny and abortive clasper, barely 
recognizable as such, an ovary which was characteristic of an adolescent female, a quite 
well developed shell gland and an oviduct which was characteristic of mature female 
erinacea. 

The work here reported would not have been possible without the generous 
cooperation of many individuals, and we make grateful acknowledgment to the follow- 
ing: Captains Ellery F.Thompson, Joseph Roderick and Harold McLaughlin trawling 
out of Stonington; Andrew, Bonaventura, and Gabriel Gargano out of New Haven; 
and the entire staff of the Bingham Oceanographic Laboratory. 

Raja fyllae Liitken 1887 
Figures 39, 40 

Study Material. Female, 452 mm long, trawled off southwestern Ireland (Lat. 
5i°37' N, Long. 1 1°56' W) between 610 and 640 fathoms (British Museum [Natural 
History]); adult male, 438 mm long, taken from southern slope of Georges Bank, 
420-480 fathoms, July 12, 1952, by Woods Hole Oceanographic Institution. 

Distinctive Characters. R. fyllae resembles R. erinacea and R. ocellata in that its 



Fishes of the Western North Atlantic 



195 




Figure 39. Raja fyllae, female, 452 mm long, from off SW Ireland (British Museum [Natural History]). 
A Ventral view of pelvics. B Nostrils and mouth, about 1.3 X. C Dorsal view of posterior part of tail, 
about 1.3 X. 

'3' 



196 Memoir Sears Foundation for Mari?ie Research 

snout is short and obtuse and the midbelt of its disc and the upper surface of its tail 
are rough with rather large thorns in several irregular rows, the thorns not much larger 
in any one row than in any of the others. But R. fyllae differs noticeably from the 
other two species in its longer tail. Young specimens of R. fyllae^ in which there is 
still only a single row of thorns along the tail, are separable by their longer tails from 
all other western North Atlantic Skates of similar thorn pattern, with the exceptions 
of R. bathyphila, R. garmani and young specimens of R. senta. The more obtuse snout 
oi R. fyllae with its pale lower surface sets it apart from i?. hathyphila, and there is no 
danger of confusing R. fyllae with R. garmani at any stage in growth, so characteristic 
is the color pattern of the latter. But the only character we have found to separate 
newly hatched R. fyllae from newly hatched R. senta (now that the former has been 
encountered within the range of the latter) is the color of the tail, which is plain- 
colored in young fyllae but pale cross-barred in young senta. 

Description. Proportional dimensions in per cent of total length. Female, 452 mm 
long, from Lat. 5i°37' N, Long. 1 1°56' W (British Museum [Natural History]). 

Disc: extreme breadth 49.3; length 40.8. 

Snout length: in front of orbits 9.5; in front of mouth 10.5. 

Orbits: horizontal diameter 4.0; distance between 3.3. 

Spiracles: length 3.1; distance between 6.4. 

Mouth: breadth 5.8. 

Nostrils: distance between inner ends ^.2,. 

Gill openings: lengths, ist 1.3; 3rd 1.3; 5th 1.2; distance between inner ends, 
1st 12.6; 5th 6.7. 

First dorsal fin: vertical height 2.2; length of base 5.1. 

Second dorsal fin: vertical height 2.0; length of base 6.2. 

Pelvics: anterior margin 10.6. 

Distance: from tip of snout to center of cloaca 39.8; from center of cloaca to 
1st dorsal 45.7; to tip of tail 60.2; from rear end of 2nd dorsal base to 
tip of caudal 3.1. 

Interspace between: ist and 2nd dorsals 0.0. 

Disc 1.2 times as broad as long, conspicuously obtuse in front but decreasingly 
so with growth, the maximum anterior angle in front of spiracles 130—140° in small 
specimens, 115—125° in adults; tip of snout marked by a low prominence. Anterior 
margins of pectorals nearly evenly convex in young specimens but weakly concave in 
larger females and considerably more so in adult males at level of spiracles, some being 
definitely indented;^ outer corners broadly rounded, posterior margins evenly and 
moderately convex, posterior corners broadly rounded with curvature extending to axils. 
Axis of greatest breadth about 70 '/o of distance rearward from tip of snout toward 
axils of pectorals. Tail with lateral folds confined to posterior one-third and widening 

2. A male of this shape, from Lat. 63° 30' N, Long. 54° 25' W, is pictured by Clark (Rep. Fish. Bd. Scot. [1926], 
I, 1926: pi. 22, fig. a). 



Fishes of the JVestern North Atlantic 



197 



toward tip; its length from center of cloaca to first dorsal greater than distance from 
center of cloaca to tip of snout by a distance 1.5-3.0 times as long as horizontal dia- 
meter of orbit in very young to medium-sized specimens and at least as long as distance 
from center of cloaca to snout in adults; total length of tail from center of cloaca to 
tip about 1.3— 1.6 times as great as distance from cloaca to snout in large specimens; 
juveniles with tail perhaps averaging a little longer.^ 

Small specimens up to 90—100 mm long with a single median row of 30—40 
large thorns on upper surface from nuchal region to first dorsal; 1—3 thorns on each 
shoulder; 1—2 close behind orbit, with as many immediately anterior to it; a few smaller 




Figure 40. Raja fy/lae, same specimen as in Fig. 39. 
A Posterior margin of nasal curtain, about 2.8 X. B 
Nostril, about 3.6 X. C Upper teeth near middle of 
jaw, about 7.2 X. 



ones near tip of rostral ridge; mid-dorsal row less regular with growth, 2—3 additional 
rows developing irregularly along either side of it (no single row more conspicuous 
than others); thus, typical adults are thorny along whole mid-dorsal belt of disc as well 
as along upper surface of tail (Fig. 39), but some have midback thornless or nearly 
so immediately posterior to shoulder region;* additional large thorns also developing 
with growth over shoulder region as a whole, around orbits, and in 2—3 irregular rows 
along rostral ridge; also a few developing in space between orbits in some specimens. 
The outer posterior margins of the pectorals as well as areas of varying extent on either 
side of the mid-dorsal belt continue thornless, but the anterior marginal regions become 
conspicuously thorny by maturity in males, as is also the case in some females but not 
in others (var. Upacanthd). A patch of large thorns also develops on the inner part of 

3. The relative length of the tail appears to vary considerably in half-grown and adult K.fyllae, to judge from published 
photographs (Clark, Rep. Fish. Bd. Scot. [1926], i, 1926: cf. pi. 22, fig. a with pi. 24, figs, a, b). 

4. Described as a separate variety, Hpacantha, p. 198. See Clark (Rep. Fish. Bd. Scotl. [1926], i, 1926: pi. 24, figs. 
a, b) for photographs of an adult male and female with thorn pattern of this sort. 



198 Memoir Sears Foundation for Marine Research 

each pectoral on some females but not on males. The skin of the disc and tail is also 
rough with small prickles on small specimens, but extensive areas on the median and 
posterior parts of the disc and on the upper surface of the tail are largely smooth in 
adults, apart from the thorns. The pelvics are naked except for prickly patches on the 
central parts of the posterior lobes, the dorsals are sparsely prickly. Alar spines of 
mature males in two to three rows. Lower surface naked in adults of both sexes, but 
described as prickly along margins of tail in newly hatched specimens.* 

Snout anterior to orbits about 2.4 times as long as distance between orbits, its 
length in front of mouth about 2.0 times as great as distance between exposed nostrils. 
Diameter of orbit about 1.3 times as long as spiracle, distance between orbits about 
as great as length of orbit or a little less. Distance between first gill openings about 
2.4 times as long as distance between exposed nostrils, distance between fifth gill open- 
ings 1.3 times; first gill openings slightly longer than fifth and about V4 the breadth 
of mouth. Nasal curtain deeply fringed with simple, bifid, or trifid lobelets; expanded 
outer (posterior) margin of nostrils also fringed. Mouth moderately arched centrally, 
the outer ends more or less concealed when closed. 

Teeth in 30—38 series, closely crowded in quincunx; those of young, and of 
females to maturity with a circular base and low, blunt, conical cusp; those of adult 
males with sharp cusp. 

First and second dorsal fins confluent, the second a little the larger (in specimen 
examined) and continuous with caudal membrane, the latter about half as long as second 
dorsal. Pelvics moderately concave, the marginal excavation having one rather notice- 
able subtriangular projection; anterior margin about 55—60 "/o ^s long as distance from 
its own origin to rear tip of pelvic; posterior lobe strongly convex outwardly and weakly 
scalloped; rear tips broadly rounded. Claspers of maturing males extending about half 
of distance from axil of pelvics toward first dorsal. 

Anterior rays of pectorals reaching about 80 "/o of distance from level of fronts 
of orbits toward tip of snout. 

Color. Upper surface ash gray to chocolate brown, either uniform or somewhat 
clouded with paler and darker. Young with more or less pronounced darker brown spots, 
pale-ringed in var. Upacantha (see discussion below). Lower surface white, grayish 
white, pale gray or light fawn color, uniform or with sooty patches on pelvics and on 
axils of pectorals in roughly symmetrical pattern. 

Remarks. A separate varietal name, Upacantha, has been proposed* for represen- 
tatives of R.fyllae from the Skagerrak, where, according to descriptions, the young 
specimens are more finely prickly than those of the typical form; adults have a sparser 
development of thorns along the midbelt of the back, and the anterior margin is defin- 
itely notched in the adult male.'' R. fyllae of the western Atlantic are the typical 
form, and it is these that come within the scope of the present account. 

5. Clark, J. Mar. biol. Ass. U.K., 12, 1922: 626. 

6. By Jensen, Vidensk. Medd. naturh. Foren. Kbh., 1905: 233. 

7. Clark, Rep. Fish. Bd. Scot. (1926), j, 1926: cf. pi. 24, fig. a with pi. 22, fig. a. 



Fishes of the Western North Atlantic 199 

Si-ze. The smallest specimens seen were 72.5 and 80 mm long, suggesting a 
length (apart from the embryonic caudal filament) of about 70 mm at hatching. Some 
males, and presumably females also, may mature at a length no greater than 450— 
460 mm;* others, however, do not mature until they are somewhat larger, for the 
claspers on a male 483 mm long are described as not fully formed." The greatest length 
that we have found recorded for R.fyllae is 543 mm, but the maximum size attained 
is probably somewhat greater. 

Developmental Stages. An egg capsule, probably of this species, taken in the Folden 
Fjord on the west coast of Norway at 250—285 meters, was smooth-shelled and black, 
40 mm long (exclusive of horns) by 24 mm broad, with the longer horns 32 mm and 
the shorter pair 2 i mm long." 

Newly hatched specimens" differ chiefly from half-grown and adults in having less 
thorny discs and the thorns along the midbelt of the back in a single regular row (p. 197). 

Habits. Of the habits of R.fyllae we know only: that the stomach of a small speci- 
men contained small crustaceans (copepods, amphipods, mysids); that nearly all speci- 
mens taken thus far have been from 148-983 fathoms (mostly 230-500 fath.), the 
greatest depth recorded for it being 1,121 fathoms (2,050 m);^^ and that they came 
from localities where the bottom temperature was between 3 and ^.^° C (37.4 and 
41.9° F). 

Range. Barents Sea, Bear Island, western Spitzbergen, Murman Coast, northern 
Norway, Denmark Strait west of Iceland, and Davis Strait off West Greenland; south 
in the east to the southern part of the Norwegian Basin, inward along the Norwegian 
trough to the Skagerrak between Norway and Denmark, Faroe Channel, the Atlantic 
Ocean west and south of Iceland and the slope westward from the Straits of Gibraltar; 
in the west to the southern slope of Georges Bank. The range of R.fyllae appears to 
be defined by its preference for a narrow temperature range of a few degrees above 
the freezing point of salt water (see above) and by restriction to at least moderately 
deep water. 

Occurrence in the Western North Atlantic. Records of its capture in the western 
Atlantic have been confined to four stations in the Greenland side of Davis Strait, 
Lats. 63°3o' to 66°49' N and to one station on the southern slope of Georges Bank. 
Since it appears to be confined in its northward dispersal to water of Atlantic influence, 
there is no reason to expect it north of the Davis Strait Ridge. But a path would 
appear to be open for it, so far as temperature is concerned, around the upper part of 
the southern slope of the Ridge at depths of 400-600 fathoms. Watch should be 
kept for it on the slope off Labrador and Newfoundland and southward at appro- 
priate depths. 

8. See Clark (Rep. Fish. Bd. Scot. [1926], /, 1926: pi. 24, fig. a) for photograph of an adult male of this size as cal- 
culated from its breadth. 

9. Jensen, Vidensk. Medd. naturh. Foren. Kbh., 1905: 234. 

10. Nordgaard, Tromso Mus. Skr., j {4), 1925: 9. 

11. See Clark (J. Mar. bid. Ass. U. K., 12, 1922: 624) for description and photographs of a very small specimen. 

12. For depth and temperature records for R.fyllae, see especially Hofsten (K. svenska VetenskAkad. Handl., 5.^ ['°]> 
1919: 83) and Clark (Rep. Fish. Bd. Scot. [1926], i, 1926: 42). 



2 00 Memoir Sears Foundation for Marine Research 

Synonyms and References: 

Raja circularis Collett, Nyt Mag. Naturv., 29, 1885: iig (listed fishing banks off Jaederen, W. Norway, also 
Skagerrak off Arendal, S. Norway, 200 and 370 fath.); Rep. Norweg. Fish. Invest., 2 Pt. 2 (3), 1905: 
5 (meas., Finmark, also Arendal, S. Norway). 

Raja fyllae^^ Liitken, Vidensk. Medd. naturh. Foren. Kbh., 1887: i, pi. i (descr., meas., cf. other species, 
ill., Davis Str., 80 fath.); Vidensk. Medd. naturh. Foren. Kbh., 1891: 32 (descr., meas., Davis Str., 
235 and 289 fath., Denmark Str., 426 fath.); Jordan and Evermann, Bull. U. S. nat. Mus., 4y (i), 
1896: 69 (descr. after Liitken, Davis Str.); Liitken, Danish Ingolf Exped., 2 (i), i8g8: 4, pi. 2 (descr., 
ill., depth, temp., Denmark Str. and W. Greenland); Garman, Mem. Harv. Mus. comp. ZooL, 24, 
1899: 380 (depth, range); Knipowitsch, .Annu. Mus. zool. Acad. St. Petersb., 5, 1900: 245 (listed, 
Murman Coast); Lonnberg, Swed. Polar Exped., Fishes, in Rev. int. Peche, Spb., 2 (4), 1900: 11 (listed, 
Spitzbergen; not seen); Knipowitsch, Annu. Mus. zool. Acad. St. Petersb., 6, 1901: 82 (depth, temp., 
Spitzbergen and Murman Sea); Ehrenbaum, in Romer and Schaudin, Fauna Arctica, 2 (i), 1902: 142 
(depths, Davis Str., Murman Sea, N. of Bear L, Spitzbergen); Knipowitsch, Exped. Wiss. Prakt. Unters. 
Murman Kiiste, I, 1902: 592 (depths, Murman Coast; in Russian, with German abstr.); Collett, Arch. 
Math. Naturv., 25 (2), 1903: 3 (refs., hist., descr., depth, range, oif N. Norway); Knipowitsch, Annu. 
Mus. zool. Acad. St. Petersb., 8 (2), 1903: 155 (listed, W. Spitzbergen); Collett, Forh. Vidensk-Selsk. 
Krist., 7, 1905: 114 {R. circularis Collett 1885, 1905 and R. falsavela Smitt 1895 considered ident. 
with R. fyllae, range, depth, temp.); Ehrenbaum, Wiss. Meeresuntersuch., Abt. Helgoland, N. S. 7, 
1905: 65 (size, thorn pattern, depth, NW. of Bear L, 400 m); Jensen, Vidensk. Medd. naturh. Foren. 
Kbh., 1905: 227 (depth, SW. from Faroes; see also under R. fyllae var. lipacant/ia); in Schiodte and 
Hansen, Zool. Danica, 2, Fisk., 12, 1907: 329, pi. 32, fig. i (descr., ill., range); Berg, Faune Russie, 
Poiss., I, 191 1 : 104 (refs., descr., meas., Barents Sea and Spitzbergen; in Russian); Murray and Hjort, 
Depths of Ocean, 1912: 433 (depths, S. of Faroes and W. from Gibraltar); Engelhardt, Abh. bayer. 
Akad. Wiss., Suppl. 4 (3), 1913: 102 (range); Garman, Mem. Harv. Mus. comp. Zool., j6, 1913: 
322 (descr.); Jensen, Mindeskr. Steenstrup Fods. Kbh., 2 (30), 1914: 27, 38 (refs., depths, temp., 
W. Greenland, chart of distrib.); Hofsten, K. svenska VetenskAkad. Handl., 54 (10), 1919: 83 (locals., 
depths, temp.); Clark, J. Mar. biol. Ass. U. K., 12 (4), 1922: 624 (descr., meas., color, stom. contents, 
depth, NW. of Scotland); Nordgaard, Tromso Mus. Skr., I (4), 1925: 9 (photo egg case, Folden Fjord, 
Norway); Clark, Rep. Fish. Bd. Scot. (1926), i, 1926: 41, pi. 22, pi. 23, fig. 2 (descr., size, discuss., 
range, photos, juv. and adult); Koeford, Rep. Sars N. Atlantic Deep Sea Exped., 1910, 4, 1927: 27 
(depths, temps., W. of Ireland and W. of Gibraltar); Clark, Fauna Ichthyol. Cons. int. Explor. Mer, 
1931 : pi. not numbered (descr., size, range, depth, ills., egg case, juv., adult); Jensen, Spol. Zool. Mus. 
Hauniensis, 9, 1948: 43 (meas., egg capsules, distr., Greenland). 

Raja falsavela Smitt, Hist. Scand. Fish., 2, 1895: 1112, fig. 319 (ill., juv., off Arendal, Norway, 370 fath., 
but not descr. or refs.). 

Raja fyllae var. lifacantha Jensen, Vidensk. Medd. naturh. Foren. Kbh., 1905: 233 (descr., color, size, depth, 
Skagerrak); Berg, Faune Russie, Poiss., j, 1911: 108, footnote (Russian); Clark, Rep. Fish. Bd. Scot. 
(1926), I, 1926: 42, pi. 23, fig. b, pi. 24 (descr., color, range, depth, photos, discuss.); Faune Ichthyol. 
Cons. int. Explor. Mer, 193 1: pi. not numbered (descr., range, photos, Skagerrak). 

Doubtful Synonym: 

Raja circularis Giinther, Challenger Rep., 22, 1887: 8 (Faroe Channel, ident. doubtful because dark below). 
Not Raia circularis Couch, Cornish Fauna, 1838: 53. 

Not Raja falsavela Bonaparte, Icon. Faun. Ital., 1841: pi. not numbered; equals R. circularis Couch 1838; 
see Clark (Rep. Fish. Bd. Scot. [1926], i, 1926: 34). 

Raja garmani Whitley 1939 

Resetted Skate, Leopard Skate 

Figures 41, 42 

Study material. One hundred and twenty specimens, male and female, 85 to 
432 mm long, from various localities between the offings of southern New England 

13. Also spelled Kaia. 



Fishes of the Western North Atlantic 



20I 




Figure 41. Raja garmani. Left, Male, 398 rnm long, from off North Carolina, Lat. 36°46' N, Long. 74°4i' W, 
57-97 fathoms (Harv. Mus. Comp. Zool., No. 34762). Right, Female, 401 mm long, from off Montauk Point, 
Long Island, New York (Harv. Mus. Comp. ZooL, No. 36315). 



and Palm Beach, Florida, in U. S. National Museum and Harvard Museum of Com- 
parative Zoology (including type). 

Distinctive Characters. The conspicuous dark rosettes on the upper surface provide 
the readiest field mark for specimens of this species from shortly after hatching, since 
no other Skate of the western Atlantic is marked in this way. Among Skates of the eastern 
Atlantic, R. garmani falls nearest to R. naevus Muller and Henle 1841 in shape of disc, 
length of tail, and general distribution of thorns; but it differs from R. naevus in 
that its back is thornier (especially when adult), the skin of the disc is naked apart 
from the thorns (largely prickly in R. naevus), and the color pattern is different. It has 
no close counterpart among Skates of the Pacific Coast of America. 

Description. Proportional dimensions in per cent of total length. Male, 398 mm 



2 02 Memoir Sears Foundation for Marine Research 

long, from Lat. 36°46' N, Long. 74°4i' W (Harv. Mus. Comp. Zool., No. 34762). 
Female, 401 mm, from 70 miles SSW of Montauk Point, Long Island, New York 
(Harv. Mus. Comp. Zool., No. 36315). 

Disc: extreme breadth 55.3, 54-6; length 44.2, 44.0. 

Snout length: in front of orbits 9.5, 9.0; in front of mouth 10.3, ii.o. 

Orbits: horizontal diameter 3.6, 4.2; distance between 3.3, 3.6. 

Spiracles: length 2.9, 3.2; distance between 6.0, 6.2. 

Mouth: breadth 5.9, 5.7. 

Nostrils: distance between inner ends 6.3, 6.0. 

Gill openings: lengths, ist 1.8, 2.1 ; 3rd 1.8, 2.1 ; 5th i.o, 1.7; distance between 
inner ends, ist 12.3, 13.0; 5th 5.4, 7.2. 

First dorsal fin: vertical height 2.9, 1.7; length of base 6.0, 5.1. 

Second dorsal fin: vertical height 2.4, 1.5; length of base 6.1, 4.9. 

Pelvics: anterior margin 11.3, 11.2. 

Distance: from tip of snout to center of cloaca 40.5, 40.7 ; from center of cloaca 
to 1st dorsal 43.0, 45.7; to tip of tail 59.5, 59-3; from rear end of 2nd 
dorsal base to tip of caudal 2,-S^ 3-°- 

Interspace between: ist and 2nd dorsals 0.9, 0.8. 

Disc 1.2 — 1.3 times as broad as long, noticeably obtuse in front, the maximum 
anterior angle in front of spiracles about 112—115° ^" mature specimens; anterior 
margins weakly concave just posterior to tip of snout and again opposite spiracles; 
otherwise rather strongly convex, the contour about the same in males as in females; 
outer corners broadly rounded, posterior corners less so; posterior margins strongly 
and evenly convex. Axis of greatest breadth about 70 "/o of distance back from tip of 
snout toward axils of pectorals. Tail with narrow dermal folds along posterior 2/3 to 
Vs; its length from center of cloaca to origin of first dorsal about 1.2 times as great as, 
to tip of tail about 1.5 times as great as, distance from snout to cloaca both in adults 
and in young after loss of filamentous prolongation. 

Smallest specimens (85—125 mm long) with a few small thorns on rostral ridge; 
three larger ones in front of each orbit, a few along inner margin of orbit, and 2—3 
close behind latter; 1—3 on each shoulder; a row of about 33—35 thorns along midline 
of disc and tail to first dorsal, and 1—2 thorns between first and second dorsals. Larger 
specimens have a row of thorns along each margin of the rostral cartilage (none on tip 
of snout), 2—5 in the space between the orbits, a row around the inner margin of the 
orbit, several in front of it and 1-3 behind, a patch of about 3—6 thorns on each 
shoulder, 1—3 irregular rows along the midbelt of back and tail on either side, but 
the median row is partially lost with growth in some cases. On small specimens the 
upper surface of the disc and tail is largely prickly in addition to the thorns, but on 
large ones the upper surface is naked apart from the thorns except for a prickly patch 
that persists beside each eye and sometimes a few prickles on the outer parts of the 
pectorals. Pelvics prickly in young; posterior parts of pelvics also prickly in females 



Fishes of the Western North Atlantic 



203 




Figure 42. Raja garmani. A Female, 173 mm long, from off southern New England, Lat. 40°o8' N, Long. 
7l°36' W (Harv. Mus. Comp. ZooL, No. 34767), to illustrate juvenile color pattern and dermal armature. 
B Ventral view of pelvics of female illustrated in Fig. 41. C Ventral view of anterior part of head of male illus- 
trated in Fig. 41, about 0.6 X. D Margin of nasal curtain of same, about 4.2 X. E Outer posterior margin of 
nostril of same, about 5.2 X- ^ Left-hand upper teeth of same near middle of jaw, about 14X. G Side view of 
posterior part of tail of female illustrated in Fig. 41, about 0.6 X . H Upper teeth of same from center of jaw, 
about 14 X. 



2 04 Memoir Sears Foundatiofi for Marine Research 

but smooth or nearly so in mature males. Alar spines of maturing males in two rows, 
perhaps more in some cases. Lower surface smooth. 

Snout in front of orbits 2.4-2.9 times as long as distance between orbits, its 
length in front of mouth about 1.6-1.7 times as great as distance between exposed 
nostrils. Distance between orbits about as long as horizontal diameter of orbit in young 
but only about 70-80 "/o as long in adults; orbit about 1.3 times as long as spiracle. 
Distance between first gill openings 2.0-2.2 times as long as distance between ex- 
posed nostrils, the distance between fifth gill openings 0.86-1.2 times as long; first 
gill openings 1.2-1.7 times as long as fifth and about 30-37 "/o as long as breadth 
of mouth. Nasal curtain and posterior (outer) margin of nostrils deeply fringed. Mouth 
considerably more arched centrally in adult males than in females or young. 

Teeth 4^50; those of females and of young males closely set, in quincunx, the 
older rows evenly rounded (as a result of wear), the younger rows with faintly indicated 
cusp; teeth of adult males spaced more loosely in more nearly transverse series, with 
blunt-tipped conical cusp. 

Interspace between first and second dorsals about 15 % as long as base of first 
dorsal, with i-2 thorns." Caudal membrane posterior to base of second dorsal about 
2/3 as long as base of first dorsal. Pelvics rather deeply concave outwardly, strongly 
scalloped around the concavity; anterior margin about 48-56 % as long as distance 
from its own origin to rear tip of pelvic; anterior lobe narrow with rounded tip, including 
only 1-2 radials besides the first stout one; posterior lobe with weakly convex outer 
margin and narrow tip. 

Anterior rays of pectorals reaching about ^4 the distance from level of fronts 
of orbits toward tip of snout. 

Color. Upper surface pale buff or brown freckled with small spots, darker or paler, 
and conspicuously marked with dark rosettes, each consisting of a group of six or more 
dark brown or black spots surrounding a central one; rosettes arranged in symmetrical 
pattern on disc and along tail, with one at origin of each dorsal fin. On specimens only 
85 mm long, one rosette is already visible on either side of the shoulder region, one 
opposite and a little behind each eye, and one or two on the tail. Lower surface white 
or pale yellow. 

Size. This is one of the smaller members of the genus, males maturing at a length 
of only about 400 mm,!^ no doubt females as well. A specimen of 85 mm is the 
smallest seen. 

Developmental Stages. Neither the egg cases nor its newly hatched young have 
been seen. 

Habits. Little is known of its habits. It appears to be restricted to depths greater 

14. The type specimen (Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: pi. 18. fig. 2) has three distinct dorsal 
fins on the tail. See Schnakenbeck (Zool. Anz., 140, 1942: 88-90) for description and photo of a specimen of the 
European R. cla'vata with an accessory dorsal fin close behind the pectoral girdle and for references to earlier ac- 
counts of similar abnormalities. 

15. The claspers of a male 395 mm long in our Study Material already extend more than halfway from the level of 
the axils of the pelvics toward the tip of the tail; they appear to be fuUy developed or nearly so. 



Fishes of the Western North. Atlantic 205 

than 35—40 fathoms. Temperatures recorded for captures oi R. garmani have fallen 
between 5.3° C (41.6° F) and 9.8-10° C (49-50° F) in the northern part of its 
range to as high as i 5° C (59.1° F) in the southern part. The known salinities at which 
it has been taken have also fallen within a narrow range: 33.8 "/oo off Block Island to 
35.3 */oo off Chesapeake Bay. 

Range. Western North Atlantic from southern Florida to the offing of southern 
New England in Long. 69°22' W, mostly on the outer part of the continental shelf 
and the upper part of the continental slope. 

Details of Occurrence. R. garmani has been taken between the 30 and 290 fathom 
contours 1" in the vicinity of the Tortugas; off the Florida Keys and Palm Beach, Florida; 
off North and South Carolina; off Chesapeake and Delaware bays, and off southern 
New England, a total of about 20 locality records. The fact that it has never been 
reported among other Skates that are taken in abundance from the slopes of Georges or 
the Scotian banks locates the eastern boundary of its regular range near the longitude 
of Cape Cod or about 70° W. In the opposite direction, it appears that it does not cross 
the Straits of Florida, for it was not represented among the extensive series of Skates 
collected in Cuban waters by the Atlantis in 1 938-1 939. 

Along southern New England it is one of the most abundant of the Skates found 
offshore. Thus we observed it in 2 i out of 44 hauls made in 47-67 fathoms between 
Long. 71° 1 5' and 7i°54' W," in 12 out of 49 hauls in 41—240 fathoms between 
Long. 67°io' and 72°2o' W." 

Synonyms and References: 

Riija^^ ornata Garman, Bull. Mus. conip. Zool. Harv., 8, 1881 : 235 (descr., depth, off Alligator Key, Florida, 
and ofFS. Carolina; considered possibly a variety of ^. ackleyi Garman i88r); Jordan and Gilbert, Bull. 
U. S. nat. Mus., 16, 1882: 877 (descr. after Garman, 1881; off S. Carolina and Florida); True, List 
Vert. Anim. S.Carolina, in Handb. S.Carolina, 1883: 261 (listed, S.Carolina); Jordan, Rep. U. S. 
Coram. Fish. (1885), 1887: 799 (listed, off Atlant. Coast N. Amer.); Goode and Bean, Smithson. Contr. 
Knowl., JO, 1895 : 26 (descr., but not ill., pi. 7, fig. 24 in vol. Jl); also, Mem. Harv. Mus. comp. Zool., 
22, 1896 (depth, off Alligator Key, Florida, and off S. Carolina); Jordan and Evermann, Bull. U. S. 
nat. Mus., 47 (i), i8g6: 70 (descr., off S. Carolina and Florida); Rep. U. S. Comm. Fish. (1895), 1896: 
221 (listed, off S.Carolina and Florida); Garman, Mem. Harv. Mus. comp. Zool., 24, 1899: 380 
(depth, range); Evermann and Kendall, Rep. U. S. Comm. Fish. (1899), 1900: 49 (listed, Alligator 
Key, Florida); Roule, Bull. Inst, oceanogr. Monaco, 243, 1912: 20 (cf. R. ackleyi); Engelhardt, Abh. 
bayer. Akad. Wiss., Suppl. -/ (3), 191 3: 102 (range); Garman, Mem. Harv. Mus. comp. Zool., j6, 
1913: 336, pi. 18, fig. 2 (descr., depth, off Florida); Radcliffe, Bull. U. S. Bur. Fish., 24^ 1916: 271 
(descr., color, size, depth, off C. Lookout, N. Carolina); Jordan, Evermann and Clark, Rep. U. S. Comm. 
Fish. (1928), 2, 1930: 26 (listed, N.Carolina and Florida); Schroeder, Bull. Boston Soc. nat. Hist., 
5(?, 193 1 : 3 (depth, off Block Island); Fowler, Proc. Acad. nat. Sci. Philad., Sg, 1937: 297 (off C. May, 

16. Depths of capture (where recorded) are: oft' Nantucket and Marthas Vineyard, 44-150 fath.; off Block Island, 
50 fath.; off New York, 70 fath.; off Chesapeake Bay, 93 fath.; oft' northern North Carolina, about 30-70 fath.; 
Lat. 36° 46' N, Long. 74° 41' W, 57-90 fath.; of}" Cape Lookout, loo-i 1 1 fath.; off South Carolina, 60-142 fath.; 
off southern Florida and the Tortugas, 127-290 fath. 

17. Hauls by the dragger Eugene H., January 27 to February 3, 1950. In each of 11 hauls only one specimen was 
caught, but in other hauls as many as 20 to 50. 

18. Hauls by the Albatross III. Only one specimen was taken in the 26 hauls made east of Long. 70° \V (in 69° 22') 
and none were caught in 14 hauls made in depths shoaler than 41 fathoms. 

19. Spelled Kaia by some authors. 



2o6 Memoir Sears Foundation for Mari?ie Research 

New Jersey); Longley and Hildebrand, Pap. Tortugas Lab., 535, 1941 : 4 (depth, ofFTortugas, Florida); 

Hildebrand, Copeia, 1941 ". 222 (listed, N. Carolina). 
Raia ackleyi ornata Goode and Bean, Smithson. Contr. Knowl., JJ, 1895 : pi. 7, fig. 24; also Mem. Harv. Mus. 

comp. Zool., 22, 1896: pi. 7, fig- 24 (ill.). 
Raja garmani Whitley, Aust. Zool., 9, 1939: 248 (^garmani proposed to replace ornata Garman, preoccupied 

by ornata Agassiz, L. [Poiss. Foss., J, 1843: 372, pi. 37, fig. 34] for a fossil Skate). 

Raja hyperborea Collett 1878 

Arctic Skate 

Figures 43, 44 

Study Material. Males, 103 and 445 mm wide, and female, 138 mm wide, from 
Faroe Channel, 607 fathoms (British Museum [Natural History]); six specimens, 
male and female, 600— 850 mm long, including two with fully formed claspers, from 
Ivigtut Fjord (about Lat. 61° N) near Julianehaab, Southwest Greenland (Harv. Mus. 
Comp. Zool., Nos. 36548 to 36553). 

Distinctive Characters. R. hyperborea resembles R.jenseni (p. 2 1 6), R. lintea (p. 232), 
and R. radiata (p. 255) in its noticeably short tail and in its mid-dorsal row of stout 
thorns which extends from the nuchal region to the first dorsal fin, the thorns close 
posterior to the pelvic girdle being much larger than any other thorns farther rearward 
along the tail. But it differs from R. radiata by its more numerous (22-31) mid-dorsal 
thorns; from R. lintea by its fewer thorns (47—51 in R. lintea) and by its shorter and 
more obtuse snout (cf. Fig. 43 with 49); and from R.jenseni by its fewer teeth (about 
38-44 series in upper jaw in R. hyperborea., 58-66 series in R.jenseni) and by the 
development with growth of large thorns on the posterior parts of the pectorals and on 
the outer anterior parts of the disc (prickly only in R.jenseni). Young specimens resemble 
young R.fyllae in general appearance, but they are separable from them by their shorter 
tails, more pointed snouts and lozenge-shaped discs. The subarctic range of R. hyper- 
borea in the Atlantic lends special interest to the fact that it seems clearly separable 
from all of the Skates that have been reported from the Bering Sea-Northeast Siberian 
region. Among these it appears closest to R. rosispinis Gill and Townsend 1897 in num- 
ber and arrangement of large thorns. But the skin of the disc (mostly smooth in R. hy- 
perborea apart from thorns) is described as largely prickly in R. rosispinis.^" Nor does 
R. hyperborea have any close counterpart in boreal-subantarctic latitudes of the South 
Atlantic, as have R. radiata and R. spinicauda. 

Description. Proportional dimensions in per cent of total length. Female, 8 10 mm 
(Harv. Mus. Comp. Zool., No. 36549). Male, 850 mm (Harv. Mus. Comp. Zool., 
No. 36553), from Ivigtut Fjord, Southwest Greenland. 

Disc: extreme breadth 76.5, 75.0; length 55.6, 54.3. 

Snout length: in front of orbits 14.3, 13.5; in front of mouth 13. i, 14.6. 

Orbits: horizontal diameter 3.1, 3.0; distance between 6.0, 6.5. 

20. The original account of R. rosispinis (Gill and Townsend, Proc. biol. Soc. Wash., 11, 1897; 231) is brief and 
without illustration. 



Fishes of the JVestern North Atlantic 



207 




Figure 43. Raja hyperborea. A Male, 850 mm long, from Tunugdliarfik Fjord, West Greenland (Harv. Mus. 
Comp. ZooL, No. 36552). B Lateral view of posterior part of tail of same, about 0.5 x . C Dorsal view of tail 
of same, about midway of its length, about 0.5 x. D Female, 785 mm long, from Tunugdliarfik Fjord, West 
Greenland (Harv. Mus. Comp. Zool., No. 36551). 



2 o 8 Memoir Sears Foundation for Marine Research 

Spiracles: length 2.3, 2.8; distance between 10. i, 10. i. 

Mouth: breadth 12.5, 12.3. 

Nostrils: distance between inner ends 10.4, ii.o. 

Gill openings: lengths, ist 1.2, 1.2; 3rd 1.4, 1.5; 5th 1.2, 1.3; distance between 

inner ends, ist 19.7, 18.8; 5th 13.8, 13.6. 
First dorsal fin: vertical height 2.0, 2.1; length of base 3.8, 3.9. 
Second dorsal fin: vertical height 1.6, 1.8; length of base 3.7, 4.0. 
Pelvics: anterior margin 12.0, 12.0. 
Distance: from tip of snout to center of cloaca 57.5, 57.1 ; from center of cloaca 

to 1st dorsal 31.3, 32.5; to tip of tail 42.5, 42.9; from rear end of 2nd 

dorsal base to tip of tail 2.8, 2.2. 
Interspace between: ist and 2nd dorsals 0.9, 1.4. 

Disc lozenge-shaped, about 1.3-1.4 times as broad as long; maximum anterior 
angle in front of spiracles about 90-105° in females and in young males but about 
110° in mature males; tip of snout not projecting noticeably; anterior margins of pec- 
torals weakly convex close anterior to level of eyes in females, considerably more strongly 
convex in mature males; outer corners abruptly rounded, posterior corners somewhat 
more broadly rounded, especially in males; posterior margins nearly straight along for- 
ward two-thirds and gently convex along rearward third. Axis of greatest breadth 66- 
7 1 "/o of distance rearward from snout toward level of axils of pectorals. Tail with mod- 
erately broad lateral folds extending its entire length ; its length from center of cloaca 
to first dorsal about 77-82 7o (small specimens) and about 54-5? % (large) as great 
as distance from center of cloaca to tip of snout; on adults, total length of tail from 
center of cloaca 74-75 % as great as distance from center of cloaca to tip of snout. 

Specimens of all sizes usually with 2-4 large thorns on each shoulder, one close 
to inner end of spiracle and another close in front of orbit; one on posterior part of 
orbital ridge and 1-3 smaller ones on some specimens; a varying number of small to 
medium thorns on anterior part of rostral ridge; a single mid-dorsal row of conspicuous 
thorns on strongly radiate bases extending from nuchal region nearly to first dorsal 
fin, the largest on disc and the smallest along posterior part of tail, numbering about 
30-31 on small specimens (i 1-15 anterior to axils of pelvics) but decreasing to about 
24-27 on half-grown specimens and larger (10-12 anterior to axils of pelvics); also 
1-3 irregular rows of thornlets developing along either side of anterior part of tail; 
median areas and anterior margins of pectorals also becoming more or less thorny; 
interspace between the two dorsals with 1-2 thorns on small specimens but not on 
large ones. In small specimens, skin of disc largely prickly, the region between and 
anterior to eyes continuing so to maturity, but half-grown specimens and larger with 
extensive naked areas on either side of mid-dorsal ridge, abreast of spiracles and eyes, 
over inner parts of pectorals, and along outer posterior margins; pelvics smooth in small 
and half-grown specimens, but central parts of posterior lobes of adults with a few small 
thorns and prickles, more conspicuous in some males than in females; dorsals either 



Fishes of the IVestern North Atlantic 



209 




Figure 44. Raja kyperiorea. A Male illustrated in Fig. 43. B Female illustrated in Fig. 43. C Nasal curtain 
of same female, about 1.5 X. D Upper teeth of same female near center of jaw, about 3.9 X. £ Nostril of male, 
illustrated in Fig. 43, about 2.9 X . 



smooth or with a few minute prickles; sides of tail, from base to tip, with a conspicuous 
belt of closely crowded prickles increasing in density rearward and outlined above by 
small thorns in some specimens but separated from mid-dorsal row of large thorns by 



2 I o Memoir Sears Foundation for Marine Research 

a more or less naked band. Alar spines of adult males in two rows, about 15 in each 
row. Lower surface entirely smooth. 

Snout in front of orbits about 4.4—4.6 times as long as distance between orbits, 
its length in front of mouth about 1.3 times as great as distance between nostrils. Orbits 
in young about 80 "/o as long as distance between them and about 1.8 times as long as 
spiracle; in adults, about 47-51 ",'0 as long as distance between them and about i.i— 1.3 
times as long as spiracle. Nasal curtain deeply fringed; expanded posterior (outer) 
margin of nostril either smooth or slightly jagged. Mouth slightly more bowed in adult 
males than in females. 

Teeth 3!"^, alike in both sexes, long and needle-pointed on broad bases and 
loosely spaced in transverse series. 

Dorsal fins close to tip of tail, about equal in size, the apex of first dorsal pointed 
(a characteristic feature), that of second dorsal more rounded; the two dorsals either 
confluent or separated by a short interspace, sometimes with 1-2 intervening thorns 
in small specimens but not in adults. Caudal membrane so low as to be distinguished 
only with difficulty. Pelvics deeply concave outwardly, the indentation scalloped in some 
specimens but not in others; anterior margin about 45-55 "/o as long as distance from 
its own origin to rear tip of pelvic; anterior lobe slender, with rounded tip, including 
only two radials besides the first stout one; posterior lobe moderately convex, its tip 
narrowly rounded or subangular. 

Anterior rays of pectorals extending forward about 55-70 "/o of distance from 
level of fronts of orbits toward tip of snout. 

Color. Upper surface dark mouse gray, dark blue gray, or dark brown, either uni- 
form or irregularly marked with small indefinite rounded spots, light or dark; outer 
posterior margins of pectorals dusky, margins of pelvics narrowly so. Lower surface of 
medium-sized and large specimens variegated white and sooty in variable patterns, some 
chiefly white below with more or less symmetrical dark markings, others chiefly dark 
with white areas confined to head, and still others intermediate. For typical patterns, 
see Fig. 44. Small specimens either lack dark markings at first ^i or may already show 
various small blotches on disc, pectorals and tail.^^ 

Relationship to Other Species. It has been suggested that R. badia Garman 1899, 
taken at 1,270 fathoms in the Gulf of Panama (Lat. 7°05' N, Long. 79°4o' W) by the 
Albatross, may be identical with R. hyperborea.^^ However, our own examination of the 
type (and unique) specimen of R. badia shows that, while it agrees closely in its dermal 
armature with R. hyperborea^ its axis of greatest breadth is much farther rearward (only 
a little anterior to the level of the axils of its pectorals) than in R. hyperborea, and the 

21. Jensen, Mindeskr. Steenstr. Fods. Kbh., 2 (30), 19 14: 21. 

22. The two small specimens we have seen are plain-colored below, but they are so evidently stained that we cannot 
say whether they were white or uniformly gray originally. One 180 mm long from the Faroe-Shetland Channel 
was yellow below, the pelvics and cloaca tinged with violet (Koefoed, Rep. Sars N. Atlantic Deep Sea Exped., 
4 [i], Zool., 1927: 24), and one 160 mm long from the Barents Sea is described as yellowish-white below, marked 
with dark spots behind the mouth, along the margins of the trunk, on the posterior portions of the pectorals, and 
on the tail (Thielemann, Wiss. Meeresuntersuch., Abt. Helgoland, N. F. 13, Heft 2, 1922: 197). 

23. Berg (Faune Russie, Poiss., J, 191 1 : 103) includes R. badia with query in the synonymy of R. hyperborea. 



Fishes of the Western North Atlantic 211 

inner edge of each of its nostrils bears a narrow barbel-like lobe directed outward and 
rearward, a character that is not duplicated in any other Skate that we have examined. 2* 

Size. The length shortly after hatching (irrespective of whatever traces may persist 
of the embryonic caudal filament) appears to average about 160-180 mm. ^^^ A male 
598 mm long is still immature, its claspers extending only a little beyond the tips of 
its pelvics; but one 850 mm long is in breeding condition, judging from the size of the 
claspers, which reach about halfway along the tail. The largest female recorded was 
860 mm long.^* 

Developmental Stages. The egg cases average about 1.5 times as long as broad, 
exclusive of the horns; the egg cases measured have ranged from 81 — 125 mm in 
length (apart from horns) by 56—80 mm in breadth." In general appearance they 
resemble those of R. radiata and their margins are similarly flanged. However, their 
blackish brown to golden yellow shells lack the rough transverse wrinkles with which 
those of R. radiata are usually sculptured, but they are described as covered with silk- 
like hairs when newly laid. 

Habits. R. hyperborea is at home in polar temperatures from hatching to maturity 
and its eggs are incubated successfully and regularly in water as cold as 0° C (32° F) 
or colder; so far it has not been taken in water warmer than 1.5° C.^* Furthermore, it 
appears to be confined to depths greater than 100 fathoms (shoalest recorded capture 
120 fath.) even in the subpolar part of its range; it occurs chiefly in depths greater than 
150-200 fathoms, and it has been taken at a depth of 1,309 fathoms.-^ In the Nor- 
wegian Basin, where the icy bottom water of polar origin is overlaid by a much warmer 
layer, the upper boundary for R. hyperborea falls about at the transition between the 
two water masses, i. e., at about 300 fathoms. But its failure to come into shoal water 
in subpolar situations, such as the Greenland fjords, cannot be explained on the basis 
of temperature. 

The nature of its teeth suggests that R. hyperborea feeds on active prey, which is 
confirmed by the fact that the stomach of a specimen taken west of Spitzbergen con- 
tained 50 large pelagic amphipods (Euthemisto libellula\ fragments of an Arctic prawn 

24. These barbels are pictured by Garman (Mem. Harv. Mus. comp. Zool., 24, i S99 : pi. 6, fig. 2) though not mentioned 
by him. 

25. The smallest free-living specimen recorded measured 160 mm (Thielemann, VViss. Meeresuntersuch., Abt. Helgo- 
land, N. F. 13, Heft 2, 1922: 195), but another of 180 mm still showed traces of the yolk sac (Koefoed, Rep. Sars 
N.Atlantic Deep Sea Exped., 4 [i], Zool., 1927: 24). 

26. Vanhotfen, in Drygalski, Gronl. Exped. Gesellsch. Erdkunde Berlin, 2 (i), 1897: 127, as R. radiata; Jensen, Minde- 
skr. Steenstr. Fods. Kbh., 2 (30), 1914: 23. 

27. For more detailed accounts of the egg cases, some positively identified because they were taken from the mother 
and another by the embryo it contained, see Jensen (Mindeskr. Steenstr. Fods. Kbh., 2 [30], 1914: 25-26; Spol. 
Zool. Mus. Hauniensis, 9, 1948: 41-43), and Thielemann (Wiss. Meeresuntersuch., Abt. Helgoland, N. F. /j, 
Heft 2, 1922: 194, 197). 

28. Our specimens from Southwest Greenland (p. 206), and one reported earlier from Ilua Fjord, near Cape Farewell, 
were from localities where the bottom temperature at the depth in question is probably below 1° C throughout 
the year, to judge from the readings that have been taken at Julianehaab and in two adjacent fjords (Krummel, 
Handb. Ozeanogr., i, 1907: 455; Jensen, Mindeskr. Steenstr. Fods. Kbh., 2 [30], 1914: 39). See Hansen (Rapp. 
Cons. Explor. Mer, 123, 1949: 8—12) for secular variations in temperature off West Greenland. 

29. 142 fathoms (Barents Sea) to 1,309 fathoms (south of Jan Mayen) in the northeastern part of its range; 120 fathoms 
down to 260 fathoms in Greenland waters. 



2 12 Memoir Sears Foundation for Mari?te Research 

(Hymenodora glacialis), and three fishes, one of them more than a third as long (185 mm) 
as its captor (518 mm). Cephalopod fragments also have been found in its stomach. 

Its egg cases have been collected in West Greenland waters, in the Barents Sea, 
and in the Norwegian Basin from midsummer to early autumn in temperatures ranging 
from 0.5-1.1° C and through the general depth range where the species occurs. 

Range. Arctic waters tributary to the northern North Atlantic, south to the Scot- 
land-Faroe-Iceland Ridge in the east and to southern Greenland in the west. 

Details of Occurrence. Thus far, recorded captures of R. hyperborea have been for 
the eastern part of the Barents Sea (three stations), from northwest of Spitzbergen (one 
station), from south of Jan Mayen (one station), from the western, southern, and south- 
western slopes of the Norwegian Sea, including the Faroe-Shetland Channel (about a 
dozen stations), and from the deep fjords along the west coast of Greenland from the 
southern extremity to as far north as Lat. 7 8° 14' N.^" But it seems that it does not occur 
along West Greenland between about Lat. 61° and 69° 13' N or on the offshore fishing 
banks along this sector. It has not been reported from the American side of Davis 
Strait, but there appears to be no thermal barrier to its dispersal westward along the 
Baffin's Bay ridge or southward, perhaps even to the northeastern slopes of the Grand 
Banks. Therefore, watch should be kept for it along the shores of outer Labrador and 
Newfoundland, both in the deeper fjord-like bays and on the slope in the ice-chilled 
Labrador Current. It is not likely to penetrate the Gulf of St. Lawrence except as a stray, 
for the temperature of the deep trough of the latter below 125-150 fathoms is too 
high for it (4-5° C), while the overlaying layer of icy cold water (colder than 1-2° C) 
does not ordinarily extend down deep enough to more than touch the uppermost bound- 
ary of the ordinary depth range of this particular species. *i 

Synonyms and References: 

Raja^''- hyperborea Collett, Vidensk. Selsk. Forh. Christiania (1878), 14, 1879: 7 (descr., meas., stom. contents, 
off NW. Spitzbergen, 459 fath.); Norske Nordhaus Exped., J, Fisk., 1880: 9, pi. i, figs, i, 2 (descr., 
ill., same specimen as Collett, 1879, off NW. Spitzbergen); Gunther, Challenger Rep., 22, 1887: 8, 
pi. 4 (descr., ill., between Scotland and Faroes, 400-600 fath.); Lilljeborg, Sverig. Norg. Fisk., J (2), 
1 891: 604 {R. hyperborea Collett 1879, cf. R. hyperborea Gunther 1887, descr., food, depth, range); 
Goode and Bean, Smithson. Contr. Knowl., 50, 31, 1895: 28, pi. 9, fig. 28; also Mem. Harv. Mus. 
comp. ZooL, 22, 1896: 28, pi. 9, fig. 28 (descr. after Collett and Gunther; ill. after Collett); Smitt, 
Hist. Scand. Fish., 2nd ed., 2, 1 895 : 1 1 1 1 (descr., ill., doubts if distinct from R. radiata) ; Liitken, Danish 
Ingolf Esped., 2 (i), 1898: 2 (descr., size, S. of Jan Mayen, 1,309 fath.; N. of Faroes); Garman, Mem. 
Harv. Mus. comp. Zool., 24, 1899: 24 {R. hyperborea Collett 1879 cf. R. hyperborea Giinther 1887); 
Lonnberg, Bih. svensk. VetenskAkad. Handl., 24 (4) 9, 1899: 34 (listed, Faroe Channel); Ehrenbaum, 
in Romer and Schaudin, Fauna Arctica, 2, 1902: 142 (listed, off NW. Spitzbergen and Faroe Channel); 
Knipowitsch, Exped. Wiss. Prakt. Unters. Murman Kiiste, i, 1902: 592 (listed, Murman Coast; in 
Russian, with German abstr.); Collett, Arch. Math. Naturv., 24 (2), 1903: 7 (refs., descr., meas., stom. 
contents, range, depth, off Vesteraalen, Norway); Knipowitsch, Annu. Mus. zool. Acad. St. Petersb., 
8 (2), 1903: 150, 151, 153, 155 (depth, W. of north. Spitzbergen and Barents Sea); Collett, Rep. Norweg. 
Fish. Invest., 2 (3), 1905: 10 (descr., meas., stom. contents, depth, refs., W. of Romsdalen, Norway, 

30. See Jensen (Spol. Zool. Mus. Hauniensis, 9, 1948: 38) for a list of localities along the west coast of Greenland. 

3 1 . For the distribution of temperatures in the deeper strata of the Gulf of St. Lawrence, see especially Bjerkan (Canad. 
Fish. Exped. [1914-1915], 7, 1919). 

32. Also spelled Rata. 



Fishes of the Western North Atlantic 213 

640 m); Medd. Norges Fisk. (1884-1901), J, 1905: 121 (Andenaes, V'esteraalen, Norway, 914 m); 
Pietschmann, Ann. naturh. (Mus.) Hofmus. Wien, 22 (4), 1907-1908: 296 (listed, NW. Greenland); 
Saemundsson, Skr. Komm. Havunders. Kbh., 5, 1909: 126 (E. of Langancs, Iceland, 455 fath.); Berg, 
Faune Russie, Poiss., I, 191 1: 103, pi. 2 (refs., descr., ill., depth, Barents Sea; in Russian); Murray 
and Hjort, Depths of Ocean, 1912: 436, 437 (depths, temps., Norwegian Sea, Faroe-Shetland Channel); 
Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 4 (3), 1913: 102 (range); Garman, Mem. Harv. Mus. 
comp. Zool., 56, 1913: 330 (descr. after Collett); Jensen, Mindeskr. Steenstr. Fods. Kbh., 2 (30), 1914: 
20 (descr., meas., egg cases, S. and NW. Greenland), 37 (range), 38, fig. 12 (chart of distrib.); Thiele- 
mann, Wiss. Meeresuntersuch., Abt. Helgoland, N. S. JJ, Heft 2, 1922: 193 (descr., meas., photo of 
egg case and embryo, depths, temps., Barents Sea); Clark, Rep. Fish. Bd. Scot. (1926), i, 1926: 30 
(descr., egg case, refs., range); Koeford, Rep. Bars N. Atlantic Deep Sea Exped. (19 10), 4 (i), 1927: 
24 (descr., meas., Faroe Channel, i,og8 fath.); Clark, Faune Ichthyol. Cons. int. Explor. Mer, 1930: 
pi. not numbered (descr., ills., range); Bigelow and Schroeder, Canad. Atlant. Fauna, I2e, 1934: 31 
(descr., ill., after Collett, range); Jensen, Spol. Zool. Mus. Hauniensis, 9, 1948: 31 (refs., descr., W. 
Greenland records). 

Raja radiata Vanhoffen, in Drygalski, Gronl. Exped. Gesellsch. Erdkunde Berlin, 2 (l), 1897: 127 (descr., 
NW. Greenland). 

Raja borea Garman, Mem. Harv. Mus. comp. Zool., 24, 1899: 24 (/?. borea, n. sp. proposed for R. hyferborea 
Giinther 1887, as distinct from R. hyferborea Collett 1879); Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 
4 (3)> 1913= i°2 (range); Garman, Mem. Harv. Mus. comp. Zool., 56, 1913: 330 (descr. after Gunther, 
188733). 

Not Raja radiata Donovan, Nat. Hist. Brit. Fish., 5, 1807: pi. 114 (see p. 255). 

Raja j ens eni Bigelow and Schroeder 1950 
Jensen's Skate 
Figures 45, 46 

Study Material. Male, 223 mm long, from the eastern slope of Georges Bank, 
Lat. 4i°09' N, Long. 66°o2' W, in 1,255 fathoms; female, 541 mm long, from the 
continental slope off southern New England, Lat. 39°35' N, Long. 7i°2i' W, in 
1,043 fathoms; female, about 850 mm long, from the continental slope off Halifax, 
Nova Scotia, in 200 fathoms ;3* all in U. S. National Museum and all collected many 
years ago. 

Distinctive Characters. R.jenseni resembles R. hyperborea closely in general appear- 
ance as well as in the arrangement of thorns on the head and shoulders and on the 
midline of the disc and tail. But it is quite sharply set apart from R. hyperborea at all 
ages in that it has 56-66 series of teeth in each jaw (38-44 in R. hyperborea); it lacks 
large thorns on the anterior margins and midposterior parts of the pectorals and on 
either side of the mid-dorsal ridge on the disc rearward from the shoulder region; also, 
thorns are lacking on the rostral ridge of adults. 

Description. Proportional dimensions in per cent of total length. Male, 223 mm, 
from Lat. 4i°o9' N, Long. 66°02' W (U. S. Nat. Mus., No. 33457)- Female, 541 mm, 
from Lat. 39°35' N, Long. 7o°2i' W (U. S. Nat. Mus., No. 35592, type). 

33. Spelled Raia. 

34. Caught from the fishing schooner AUGUSTA H. JOHNSTON, Captain G. A. Johnston. It is catalogued as being from 
Lat. 42° 37' N, Long. 62' 55' W, but the depth at which it was taken actually locates it as being taken a few miles 
farther north and closer inshore. 



214 



Mejnoir Sears Foundation for Marine Research 




Figure 45. Rajajenseni, female, 541 mm long, from continental slope off southern New England, Lat. 39°35' N, 
Long. 7i°2i' W, 1,043 fathoms (U. S. Nat. Mus., No. 35592, type). A Upper surface of midsector of tail of 
same, about i.zx. B Right-hand nostril and nasal curtain of young specimen illustrated in Fig. 46, about 2.3 X . 



Disc: extreme breadth 70.2, 81.3; length 53.7, 60.0 
Snout length: in front of orbits 14.4, 15.5; in front of mouth 16.9, 16.3. 
Orbits: horizontal diameter 4.4, 4.8; distance between 6.2, 6.6. 
Spiracles: length 2.5, 2.8; distance between 10.6, 12.0 
Mouth: breadth 12.5, 12.5. 
Nostrils: distance between inner ends 12.8, — . 

Gill openings: lengths, ist 1.3, 1.3; 3rd 1.6, 1.5; 5th i.o, i.o; distance between 
inner ends, ist 18.9, 21.6; 5th 14.0, 18.1. 



Fishes of the Western Nort/i Atlantic 



215 




Figure 46. Raja jenseni. A Juvenile male, 223 mm long, from eastern slope of Georges Bank, 1,255 fathoms 
(U. S. Nat. Mus., No. 33457). B Thorn from head posterior to e}es (above) and from tail (below) of same, 
enlarged. C Posterior part of tail of larger female illustrated in Fig. 45, about 1.2 X- D Upper teeth of same, 
about 12 X. £ Apical and lateral views of tooth, about 12 X. 

First dorsal fin: vertical height 1.7, 1.8; length of base 5.0, 5.4. 

Second dorsal fin: vertical height 1.5, 2.0; length of base 5.2, 5.2. 

Pelvics: anterior margin 10.7, 11.5. 

Distance: from tip of snout to center of cloaca 51.5, 52.8 ; from center of cloaca 

to 1st dorsal 33.6, 32.5; to tip of tail 48.5, 47.2; from rear end of 2nd 

dorsal base to tip of caudal 3.9, 2.6. 
Interspace between: ist and 2nd dorsals 0.8, 0.9. 



2 1 6 Memoir Sears Foundation for Marine Research 

R.jenseni agrees so closely with R. hyperhorea in proportional dimensions and in 
shape of disc and fins that detailed description in these respects would only be repeti- 
tious. There is a median row of 24—3 1 large thorns on conspicuously radiate bases 
from the nuchal region to the first dorsal fin, these becoming successively smaller and 
increasingly crowded rearward along the tail, the total number decreasing somewhat 
with growth ;^^ four large thorns on each shoulder on smaller specimen, 2—3 on larger; 
one close to inner end of spiracle, one just in front of each orbit, and one close behind 
it; a row of several small thorns around inner anterior margin of orbit in young but 
not in older specimens; also on young, 2—3 rows of thorns of various sizes along 
rostral ridge which are shed progressively with growth; disc also generally rough with 
small prickles (In addition to the thorns) on small specimens except around posterior 
margins of pectorals; the snout, the region between orbits, the anterior margins of disc, 
a band across each pectoral, and a belt along either side of posterior part of mid-dorsal 
ridge continuing prickly with growth; pelvics prickly on inner part but without thorns 
(more or less thorny on R. hyperhorea); first and second dorsals with a few small prickles; 
sides of tail with a dense prickly band from base to tip. Lower surface smooth.^* Teeth 
56—66 above and below on specimens counted, loosely spaced in transverse series, 
sharp-pointed in both sexes, as in R. hyperhorea (cf. Fig. 46 D with 44 D). 

Color. Upper surface (after many years' preservation in alcohol) plain light brown, 
either grayish or of chocolate hue, darker along margins of fins. Ground tint of lower 
surface brownish gray, grayish white, or perhaps pure white, but areas around cloaca 
and over inner parts of pelvics of small specimens dusky, these dark areas expanding 
over the abdomen with growth; outer margins of pectorals also brown in large speci- 
mens. Conditions in the related species, R. hyperhorea (p. 210), suggest that the dark 
markings on the lower surface may be widely variable in R.jenseni also. 

Remarks. R.jenseni was first reported and discussed as Raja granulata Goode and 
Bean, 3' and our earlier accounts of it^^ were under that same name. But it seems prac- 
tically certain that the Skate for which the specific name granulata was proposed orig- 
inally^^ and the specimen that was figured subsequently under that name by its de- 
scribers*" actually was a Raja laevis (p. 228). 

Si-ze. The largest specimen known (a female) was about 850 mm long (see Study 
Material), suggesting that its maximum size is about the same as that for R. hyperhorea. 

35. Thirty-one thorns in the median row in the smallest R.jenseni seen but only 24 or 25 in the half-grown specimen; 
a complete count could not be made for the largest specimen. 

36. The largest of the three listed specimens (see Study Material, p. 213) was described earlier by us (Bull. Mus. comp. 
Zool. Harv., 68, 1927: 246; Canad. Atlant. Fauna, 12% 1934: 29) under the name R. granulata Goode and Bean 
1879 as having minute rounded tubercles sparsely sprinkled over the lower surface of the disc. Re-examination 
has shown, however, that Jensen's (Mindeskr. Steenstr. Fods. Kbh., 2 [30], 1914: 22, footnote) characterization 
of it as perfectly smooth below was correct; doubtless the supposed tubercles (no longer to be seen or felt) were 
merely grains of sand imbedded in the mucous. The specimen was in a good enough state to afford a detailed de- 
scription when we first saw it, but it is now fragmentary. 

37. Jensen, Mindeskr. Steenstr. Fods. Kbh., 2 [30], 1914: 31. 

38. Bull. Mus. comp. Zool. Harv., 68, 1927: 246; Canad. Atlant. Fauna, 12*^, 1934; 29. 

39. Goode and Bean, Bull. Essex Inst., 11, 1879: 28. 

40. Goode and Bean, Smithson. Contr. Knowl., 31, 1895: pi. 9, fig. 30; also, Mem. Harv. Mus. comp. Zool., 22, 1896: 
pi. 9, fig. 30. 



Fishes of the JVestern North Atlantic 217 

Developmental Stages. Neither the egg cases nor the newly hatched young of 
R.jenseni have been seen. 

Habits. The localities and depths of capture indicate that R.jenseni lives at rather 
higher temperatures (3.6-3.9° C, or 38-39° F) than does R. hyperborea; that it is 
strictly a deep-water species, finding its upper limit at or below 200 fathoms, otherwise 
fishermen would almost certainly pick it up along the upper slopes of the Banks; and 
that its range almost certainly extends down to a depth of at least 1,200-1,300 fathoms.*^ 
The nature of its teeth suggests that it feeds chiefly on active prey such as small fishes 
and free-swimming Crustacea, as its relative R. hyperborea is known to do. 

Range. R.jenseni is known only from the offing of Halifax, Nova Scotia in 200 
fathoms and from the lower part of the continental slope ofi-" Georges Bank and southern 
New England in 1,043-1,255 fathoms, at the localities listed (p. 213). But its range, 
if governed by distribution of water temperature (3-4° C, 37-39° F) at the consider- 
able depths at which they have been taken, may well extend much farther south, for 
the bottom water is of about that temperature all along the lower part of the conti- 
nental slope from the southeastern corner of the Newfoundland Banks to the offing of 
northern Argentina. 

Synonyms and References: 

Raja jenseni Bigelow and Schroeder, Bull. Mus. comp. Zool. Harv., JOJ, 1950: 385 (descr., ill.). 

Raja granulata Jensen, Mindeskr. Steenstr. Fods. Kbh., 2 (30), 1914: 31 (off Halifax, N.Scotia, 42°37' N, 
62°55' W, 200 fath., cf. R. spinicauda); Bigelow and Schroeder, Bull. Mus. comp. Zool. Harv., 68, 
1927: 246 (descr., meas., same spec, as Jensen, 1914, off Halifax, Nova Scotia); Canad. Atlant. Fauna, 
126, 1934: 29 (descr., ill. of same specimen as in Jensen, 1914, and Bigelow and Schroeder, 1927, off 
Halifax, Nova Scotia; but LaHave Bank ref. is to R. granulata Goode and Bean 1879, equals R. laevis 
Mitchill 1 8 17); Bull. U. S. Bur. Fish., 48, 1936: 323, footnote (ref. to old record off Halifax, Nova 
Scotia); Vladykov, Nat. canad., 6j, 1936: 212, 228 (suggests R. granulata ident. with R. hyperborea). 

Raja granulata (in part) Vladykov and McKenzie, Proc. N. S. Inst. Sci., ig, 1935: 51 (off Halifax, but ref. 
for LaHave Bank is to R. granulata Goode and Bean 1879, equals R. laevis Mitchill 18 17). 

Raja laevis Mitchill 1817*2 

Barndoor Skate, Sharp-nosed Skate 

Figures 47, 48 

Study Material. Thirty-one specimens, male and female, 7^/5 to 52 inches long, 
from Nahant and Woods Hole, Massachusetts, Georges Bank, Nantucket Shoals, the 
continental shelf 70-80 miles southward from Marthas Vineyard, 50 miles east-south- 
east of Chesapeake Bay, and the offing of Charleston, South Carolina, in the collections 
of the Harvard Museum of Comparative Zoology and the U. S. National Museum; 

41. It seems safe to assume that the two specimens for which depths of 1,043 ^"d 1,255 fathoms are recorded were in 
fact picked up by the trawl on the bottom or close to it. 

42. Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 341, 342) substituted the name stabuliforis for this Skate to re- 
place lae'vis on the ground that the latter had been preoccupied by Gronow (1763), Valmont (1768), and Duhamel 
(1782) for other species. But Valmont's and Gronow 's names have been ruled out by the International Commission 
on Zoological Nomenclature (Opin. 89, Smithson. misc. Coll., 73 [3], 1925: 27), and Duhamel's names cannot 
be taken into account because, when binominal, they were so only incidentally. 



2 I 8 Memoir Sears Foundation for Marine Research 

also many specimens up to about five feet long examined by us in the field between 
Chesapeake Bay and eastern Maine. 

Distinctive Characters. The midbelt of the disc of R. laevis, from the level of the 
axils of the pectorals to the vicinity of the spiracles, lacks large thorns, which sets 
it apart from all other Skates of the genus Raja of the western North Atlantic, except 
for R. mollis and R. spinicauda. And its thorny tail differentiates it from mollis. It is 
easily separable from spinicauda (at least from the time it is one-third grown) in that 
its tail is armed with three rows of thorns (only a single median row in R. spinicauda), 
and that the mucous pores on its lower surface are marked by conspicuous black dots 
and dashes (not so in R. spinicauda or in any other Raja of the western North Atlantic). 
Newly hatched specimens of R. laevis, on which the mucous pores are not yet pig- 
mented (p. 222) and on which only one row of thorns has developed along the tail, 
may prove difficult to separate from R. spinicauda at a corresponding stage. The young 
of the latter have not been seen. R. laevis is so closely allied to the well known Gray 
Skate (R. batis) of the northeastern Atlantic, which also has the mucous pores of the 
lower surface marked by black dots and dashes, that future study may show that the 
two represent extremes of one varietal series. However, present indications are that 
the teeth average fewer in R. laevis (30-40) than in R. batis (44-55), those of females 
being relatively lower and more rounded; the tail may average a little longer in R. 
laevis; the lateral rows of thorns along the tail are more regular (often largely lacking 
in R. batis). On the other hand, the lower surface of the disc of some specimens is 
more generally prickly on R. batis.*^ 

The only Skates known in the western Atlantic, other than R. laevis, which have 
pigmented mucous pores on the lower surface are R. castelnaui Ribeiro 1907" and 
some specimens of i?. agassizii Miiller and Henle 1841, both from Brazil;*^ R.platana 
Giinther from Rio de Janeiro to the mouth of the Rio de La Plata; and R. flavirostris 
Philippi 1892 from the Patagonian-Falklands region and Straits of Magellan to Chile. 
The anterior contour of the disc is so much more obtuse in R. castelnaui, and the dorsal 
fins are so much farther removed from the tip of the tail in R. agassizii, that R. laevis 
is not likely to be confused with either of these species. R.platana and R. flavirostris 
resemble R. laevis closely in general form. But R. flavirostris is characterized by the 
presence of 2-3 strong thorns around each orbit and one large nuchal thorn, which 
are lacking on R. laevis. And in R.platana the median thorns on the tail (18 on spe- 
cimen studied) are not only fewer and much larger than in R. laevis but are successively 
larger rearward; there are no side rows on the tail; the space between the eyes and 
also the mid-dorsal belt in the nuchal region (smooth in R. laevis) are close-set with 
small, low, rounded tubercles; and the caudal extension, posterior to the second dorsal 
fin, is considerably longer relatively than in laevis. 

4j. We have for comparison two large R. batis from Sweden. 

44. Arch. Mus. nac. Rio de J., 14, 1907: 177, pi. 15. 

45. Some of the specimens of R. agassizii that we have examined have the mucous pores pigmented on the rostrum, 
around the mouth, and in the area between the gill openings, but others are immaculate below except for a sooty- 
blotch under the tip of the snout. 



Fishes of the JVestern North Atlantic 



219 




Figure 47. Raja laevis. A Dorsal view of female, 1,187 mm long, from off Massachusetts (Harv. Mus. Comp. 
Zool., No. 368). B Ventral view of another female, 676 mm long, from Nantucket Shoals (Harv. Mus. Comp. 
ZooL, No. 36302), to show pigmented mucous pores. C Nasal curtain of same, about 2.6 X . T> Nostril of same, 
about 2.6 X. E Upper teeth from center of jaw of female, 1,265 mm long, from Nantucket Shoals, about 3.5 X. 
F Teeth from center of jaw of male, 1,320 mm long, from Nantucket Shoals (Harv. Mus. Comp. ZooL, 
No. 36264), about 3.5 X; symphysis marked by dotted line. 



Among Skates of the Pacific Coast of North America, the closest counterparts to 
R, laevis are R.rhina Jordan and Gilbert 1880, also with pigmented mucous pores, 
and R. binoculata Girard 1854, both of which range from southern California to Alaska. 
However, the anterior margins of R. rhina are much more deeply concave than are 
those of R. laevis^ and the snout of the former is narrower and considerably longer 
relatively (snout in front of mouth about 4 times as long as distance between orbits 
in R. rhina\ only about 2.5-3 times in R. laevis)^ to mention only the more obvious dif- 
ferences. R. binoculata has two large orbital thorns and one large mid-dorsal thorn on 
the nuchal region when young but only one row of thorns (in addition to prickles) 
along the tail when adult; the mucous pores on its lower surface are not pigmented; 



2 2 o Memoir Sears Foundation for Marine Research 

and in many cases the upper surface of each pectoral is marked with a large and con- 
spicuous ocellar spot. 

Description. Proportional dimensions in per cent of total length. Male, 1,064 mm, 
and female, 1,187 "^"^> from off Massachusetts (Harv. Mus. Comp. Zool., No. 368). 
Disc: extreme breadth 74.3, 72.0; length 53,9, 51.8. 
Snout length: in front of orbits 15.2, 15.7; in front of mouth 16.8, 15.6. 
Orbits: horizontal diameter 3.8, 2-S'-< distance between 5.4, 5.5. 
Spiracles: length 3.0, 2.5; distance between 7.2, 7.0. 
Mouth: breadth 9.8, 9.2. 

Nostrils: distance between inner ends 8.8, 8.0. 
Gill openings: lengths, ist 1.8, 2.0; 3rd 2.1, 2.1 ; 5th 1.5, 1.4; distance between 

inner ends, ist 16.5, 17.0; 5th 10.4, 11.4. 
First dorsal fin: vertical height 3.1, 3.4; length of base 5.7, 6.0. 
Second dorsal fin: vertical height 2.4, 2.9; length of base 6.1, 4.9. 
Pelvics: anterior margin lo.o, 10. i. 

Distance: from tip of snout to center of cloaca 51.9, 50.8 ; from center of cloaca 
to 1st dorsal 31.7, 33-8; to tip of tail 48.1, 49.2; from rear end of 2nd 
dorsal base to tip of tail 3.8, 2.8. 
Interspace between: ist and 2nd dorsals 0.8, 1.8. 

Disc about i .4 times as broad as long, the maximum anterior angle in front of 
spiracles about 82-90°, a little more acute on newborn specimens than on larger; 
anterior margins weakly concave just posterior to tip of snout, weakly convex opposite 
eyes and spiracles (no more so in males than in females), weakly concave thence rear- 
ward; outer corners narrowly rounded, posterior corners more broadly so, posterior 
margins nearly straight forward but gently convex rearward. Axis of greatest breadth 
about 70 o/o of distance back from tip of snout toward axils of pectorals. Tail with 
lateral folds beginning posterior to axils of pelvics by a distance about ^s as long as 
eye and extending nearly to tip; its length from center of cloaca to origin of first 
dorsal fin about 76 "/o as great as distance from center of cloaca to tip of snout in 
newly hatched specimens, about 70 "/o on half-grown ones, and about 66-68 "/o on 
large; total length of tail about 1.3 times as great as distance from center of cloaca to 
tip of snout at hatching, decreasing relatively to about as long as body sector, or a 
little shorter, at maturity. 

Newly hatched specimens with 1-2 thorns in front of orbit and one behind, a 
row of about 14-17 along midline of tail from a little posterior to axils of pectorals to 
first dorsal fin, and one in interspace between first and second dorsals; otherwise smooth 
both above and below. Larger specimens with the pre- and post-ocular thorns replaced 
by patches of much smaller ones; also thorns in mid-dorsal row on tail increasing to 
23-40 in both sexes, often irregularly spaced, the most anterior one a little behind 
axils of pectorals in some specimens but a little behind axils of pelvics in others; an 
additional continuous row of equally large thorns developing (irregularly at first) low 



Fishes of the Western North Atlantic 



221 



down along either side of tail from about opposite tips of pelvics about to level of 
second dorsal; a second thorn developing on some in interspace between first and 
second dorsals, the two members of the resultant pair usually standing side by side; 
small prickles also developing irregularly over restricted areas on snout, around and 
between eyes, along outer anterior margins of disc, on outer parts of dorsal fins, and 
along margin of caudal membrane. Maturing females becoming more or less prickly 




Figure 48. Raja laevis. A Ventral view of pelvics of mature male, 1,206 mm long, from off Woods Hole, Massa- 
chusetts (Harv. Mus. Comp. ZooL, No. 36691), about 0.23 X . B Oblique view of clasper of same. C Opened 
mouth of same, with right-hand nasal curtain turned up to show nostril, transverse curtain on roof of mouth, 
and fleshy knobs on floor of mouth, about 0.45 X. 

over upper surface of tail, on posterior part of mid-dorsal belt of disc, on shoulders, 
and outward from eyes; also a few strong thorns, directed forward, developing on 
rostral ridge, with smaller thorns scattered along anterior margins of pectorals. Mature 
males smoother, the rostral ridge, region between orbits, and the central area of disc 
posterior to eyes with only a few sparsely scattered prickles. Tail also prickly apart from 
its thorns. Alar spines of sexually mature males In three rows. Lower surface glossy 
smooth on small specimens, likewise on large males except for a few prickles along 
margins of snout; but large females are rough with small spines and prickles along a 
narrow marginal belt from tip of snout to level of nostrils, and more or less prickly on 
spaces between first and third gill openings of each side and on anterior parts of pelvics. 



2 2 2 Memoir Sears Foundation for Marine Research 

Snout in front of orbits about 3—4 times as long as orbit in small specimens, 4—5 
times in adults, its length in front of mouth 1.8—2.2 times as great as distance between 
exposed nostrils in young and old alike. Distance between orbits about as great as 
length of orbit at hatching, 1.4- 1.7 times when full grown; orbits about 1.8 times as 
long as spiracles on young but about 1.3 times on adults. Distance between first gill 
openings 1.8— 2.1 times as long as distance between exposed nostrils, between fifth 
gill openings 1.2— 1.4 times; first gill openings 1.2— 1.4 times as long as fifth and about 
18—21 "/o as long as breadth of mouth. Nasal curtain deeply fringed, the individual 
lobes simple or bifid; expanded posterior (outer) margin of nostrils smooth. Upper 
jaw nearly straight, lower weakly arched in both sexes. 

Teeth 23^? usually one or two more series in upper jaw than in lower, some- 
times the reverse; those of females and of immature males close-set in quincunx, their 
bases nearly circular in outer series but quadrate with rounded corners in central series; 
the oldest (anterior) four or five rows worn smooth, the youngest (posterior) two or three 
rows with low conical median cusp; those of mature males more loosely spaced in trans- 
verse series, with conical cusp, considerably longest and sharpest in the younger rows. 

First and second dorsals similar in size and shape. Interspace between dorsals 
14—30 "/o as long as base of first dorsal, independent of size or sex. Caudal from 
rear end of base of second dorsal about 1.5 times as long as base of first dorsal on 
newly hatched specimens, about as long as base of first dorsal on half-grown spe- 
cimens, but only about half as long on large. Pelvics noticeably small in area relative to 
disc, rather deeply concave, strongly scalloped around anterior side of the excavation 
but only weakly so rearward; anterior margin about V2— ^/d as long as distance from its 
own origin to rear tip of pelvic; anterior lobe broad, including 5-6 radial cartilages 
besides the first stout one; posterior lobe strongly convex outwardly; rear tips abruptly 
rounded, extending rearward about 1/4 the distance from axils of pelvics toward first 
dorsal. Claspers of sexually mature males conspicuously massive, reaching rearward 
about 3/4 the distance from axils of pelvics toward first dorsal. 

Anterior rays of pectorals extending about -/g the distance from level of front of 
orbits toward tip of snout. 

Color. On fresh specimens, the upper surface brownish, with many scattered small 
darker spots of various sizes, the largest usually about the size of eye, more numerous 
on some specimens than on others; usually a larger and irregularly elongate spot or 
blotch on inner part of each pectoral about level with outer corner. Mucous pores on 
nuchal region and anterior to eyes with black dots (Fig. 47). Lower surface white, 
frequently blotched irregularly with gray, especially on large individuals; specimens 
one-third grown and larger always with small black or dusky dots or short streaks 
marking the orifices of mucous pores, in continuous arcs around midbelts of pectorals, 
across midzone of disc, and on pelvic regions (Fig. 47 B); also scattered less regularly 
elsewhere. Very small ones may lack these black markings.** 

46. The mucous pores on the smallest specimens we have seen (about 190 mm long) are only faintly visible after many 
years in alcohol, but they may have been more conspicuously marked when the specimens were fresh. 



Fishes of the JVestern North Atlantic 223 

Size. The length at hatching probably averages about 180-190 mm.*' It has been 
said that the Barndoor grows to a length as great as six feet; but the maximum that 
we find definitely reported is about five feet; the largest we have seen measured 58 
inches. Specimens taken off southern New England weigh about i^U-i'U pounds 
when 20 Inches long, about 4—6 pounds at 28—30 inches, about 10— 11 pounds at 2^ 
inches, about 20—25 pounds at 45—46 inches, and 30-38 pounds at 50-56 inches." 

Developmental Stages. The yellowish or greenish brown egg cases, much larger 
than those of any other shoal-water Skate occupying the same geographic range, are 
124— 132 mm long by 68— 72 mm broad, exclusive of the terminal horns, smooth- 
shelled, with slightly bulging sides, the horns short (only 13— 19 mm long), stout 
basally but tapering to slender tips." Fine filaments are attached to the margins of the 
case, but they are not united into a membrane. Normally the newly hatched Skates 
differ so little from their parents that they are easily identifiable. 

Abnormalities. Two specimens, 19 and 20^2 inches long, in which the anterior 
parts of the pectorals continued separate from the sides of the head,^" have been reported 
from Woods Hole. 

Habits. The Barndoor Skate feeds chiefly on the larger Crustacea, such as lobsters, 
crabs, spider crabs, and shrimps. But isopods have been found in the stomachs also, 
as have bivalves {7'oldia, and the razor clam Ensella)^ large gastropods (Buccinum, 
Lunatia), squid, and various worms. They are also more destructive to fishes than are 
other local Skates, their diet including spiny dogfish (Squalus), herring (Clupea), alewives 
(^Pomolobus), menhaden (Brevoortia), sand launce (Ammodytes), cunners [Tautogolabrus), 
tautog (Tautoga), butterfish (Poronotus), sculpins (Myoxocep/ialus), whiting (Merluccius), 
hake (^Urophycis), and various flatfishes. No doubt cod, haddock, etc., also suffer to 
some extent from them on the offshore fishing banks, for their European relative, 
R. batis, is a well known enemy of the cod. They bite readily on almost any bait. 

The young are produced throughout the latitudinal range of the species. Females 
containing eggs have been taken off outer Nova Scotia as well as in Kennebecasis Bay, 
tributary to the New Brunswick shore of the Bay of Fundy, and recently hatched 
specimens have been captured on Nantucket Shoals and off Chesapeake Bay. It is 
probable that eggs are deposited from close to tide line down to the greatest depth at 
which this !Skate ordinarily occurs. ^^ Females containing fully formed capsules have 
been taken in December and January, ^^^ evidence that the eggs are laid in winter. How- 
ever, it seems that the young are not hatched until late spring or early summer, for we 

47. Specimens of about 190 mm, listed under Study Material, appear to have been newly hatched, to judge from the 
considerable lengths of their tails posterior to the second dorsal fin. 

48. Information contributed by Daniel Merriman. Reported weights of 60 pounds (at a length of 3 ft.) and 200 pounds 
(Perley, Rep. Fish. Bay of Fundy, 1851: 154; Storer, Boston J. nat. Hist., 2, 1839: 545) were evidently much 
too high. 

49. Account based on description and photograph by Vladykov (Nat. canad., 63, 1936: 216, 219) of egg cases taken 
from females off Nova Scotia and in Kennebecasis Bay, New Brunswick. 

50. Radcliffe, Nat. Hist. N. Y., 28, 192S: 58; Gudger, Amer. Mus. Novit., 600, 1933: i. 

51. Eggs have been found in females caught in 15-25 fathoms and the empty cases have been picked up on the beach. 

52. Vladykov, Nat. canad., 6j, 1936: 216. 



2 24 Memoir Sears Foundation for Marine Research 

have a specimen, taken on Nantucket Shoals in mid-July, that is so small (193 mm 
long) that it could not have been set free long before its capture. 

Barndoors are usually taken on sandy or gravelly bottom, but individuals that 
live in the deepest water are often on muddy ground. In the Gulf of Maine, for example, 
we have trawled them in the deep mud-floored troughs in company with R. senta and 
R. radiata. 

The depth range for R. laevis covers the whole breadth of the continental shelf, 
from tide-mark down to 235 fathoms ;^^ the 5-fathoni and the 70— 80-fathom contours 
approximately enclose the zone of greatest abundance and of most regular occurrence. 

It has been reported off northern Nova Scotia and on the southwestern part of 
the Newfoundland Banks in water as cold as 1.2 and 2.2° C (about 34 and 36° F). 
The facts that many more are taken in autumn than in summer in shoal water along 
southern New England and that they are reported as not entering Sandy Hook Bay 
near New York City until the water has cooled to about 17° C (63° F), suggest that this 
is near the upper limit of the optimum range of the species. However, the individuals 
that live closest inshore (i. e., shoalest) along southern New England and southward 
may be in water as warm as 18—20° C (about 64—68° F) in summer, if not warmer. 
Especially interesting in this connection is the report of small R. laevis being taken 
in 18 fathoms off Cape Lookout, North Carolina in mid-August^* when the bottom 
temperature there was probably not less than 22-24° C (72—76° F). 

R. laevis occurs in salinities that range from about 35 "/oo along the continental 
edge to 31—31.5 "/oo inshore along the open coast and to as low as 21—24 "/oo within 
the mouth of Chesapeake Bay. One specimen has even been reported from so far up 
the Delaware River (above Philadelphia) as to suggest that R. laevis may occasionally 
run up into water that is only slightly saline or even fresh. 

The fact that R. laevis is taken much more regularly and in larger numbers 
close inshore along southern New England, New York, and New Jersey in autumn and 
spring than in summer suggests that the inshore fringe of the population tends to move 
out into slightly deeper water during the warmest season and to move shoreward again 
as the water cools with the onset of autumn. It is likely also that such individuals as 
come into water less than a fathom or so in depth during autumn tend to withdraw 
again during the winter as the water chills. But there is no reason to suppose that these 
thermal migrations involve such of the stock as live deeper than 5—10 fathoms (includ- 
ing the great majority of the population), for Barndoors are taken regularly on the 
offshore fishing banks in winter as well as summer.^^ 

53. The records of greatest depth are 120 fathoms on the southwest slope of the Grand Banks of Newfoundland; 235 
fathoms along the continental edge off Nantucket, Massachusetts; 105 fathoms in the western side of the Gulf of 
Maine; no fathoms in the central basin of the Gulf; and 159 fathoms off Charleston, South Carolina. 

54. Radcliffe, Bull. U. S. Bur. Fish., j.#, 1916: 273. 

55. It is taken throughout the year in the vicinity of Nantucket Lightship where it is especially plentiful in winter in 
depths of about 30-50 fathoms and in summer in about 20 fathoms (reported by Captains Jared Vincent and H. W. 
Klimm, catches from 1945-1950). And we saw many trawled in midwinter south of Rhode Island in 47-67 fathoms 
(by the Eugene H., January 27 to February 3, 1950), as well as along the southern New England Coast in May 
(by the Albatross III). 



Fishes of the JVestern Nort/i Atlantic 225 

Nor is it likely that any north-south migration takes place with the seasons, 
R. laevis having been taken off North Carolina both in August and in December- 
January. 

Relation to Man. This species (with a few R. oceUatd) provides the greater part of 
the small landings of Skates along the Atlantic Coast of the United States that are 
used for food. The fleshy basal parts of the pectorals, known as "skate saddles," are 
the only portions marketed. 

Numerical Abundance. The average catch per hour, made by one trawler with an 
80-foot trawl, between August 1943 and October 1944 at the eastern end of Long 
Island Sound was 25—73 pounds, or about two specimens per hour if the average weight 
were 12—15 pounds. ^^ This corresponds to a population of about 120 specimens per 
square (sea) mile, assuming that the strip swept by the trawl was 50 feet wide, that 
the distance covered per hour was two miles, and that the trawl picked up every Skate 
that lay in its path, which certainly was not the case. Also, 37 hauls made with an 
otter trawl on Georges Bank during September 1929 yielded 42 R. laevis from a total 
of 495 Skates of all kinds. Off southern New England 44 midwinter hauls in 47—67 
fathoms and 63 hauls in May at 22—235 fathoms yielded 441 R. laevis from a total 
of 748 Skates of all kinds. And as many as three dozen have been taken from one of 
the fish traps in Narragansett Bay in one lift, according to an old report. 

Range. Atlantic shelf of North America, from the Grand Banks of Newfound- 
land and southern side of the Gulf of St. Lawrence to North Carolina; also doubtfully 
reported from Florida. 

Details of Occurrence. The most northerly records of R. laevis are for the south- 
western parts of the Newfoundland Banks (reported from three stations) and from 
the southern side of the Gulf of St. Lawrence (Cape Breton and Magdalen Islands), 
where large ones have been described as fairly common. It occurs all along the outer 
coast of Nova Scotia, both inshore and on the offshore fishing banks; on both sides of 
the Bay of Fundy, including the larger tributary bays such as St. Mary, Kennebecasis, 
and Passamaquoddy; along the coast of Maine; in Massachusetts Bay where we have 
seen many; also on Georges Bank and on Nantucket Shoals where it is taken in some 
numbers by trawlers. In fact, any very large Skate reported from the Gulf of Maine 
or from the offshore fishing grounds that front the latter is almost certain to have been 
a Barndoor. It is present throughout the year on the continental shelf off southern New 
England and, in season, it is equally universal all along that coast, off New York, New 
Jersey and Maryland, and doubtless off the northern coast of Virginia," for it is fre- 
quently taken in pound nets in the lower part of Chesapeake Bay. There is a record of 
it between Cape Charles and Cape Hatteras;^^ odd individuals have been taken near 
Cape Lookout, North Carolina, and our Study Material includes a small specimen 

56. Merriman and Warfel, 9th N. Amer. Wildl. Conf., 1944: 234. 

57. Reported thence from two localities only. 

58. One specimen, trawled on February 11, 1930 by the U. S. Fisheries steamer Albatross II off Bodie Island, North 
Carolina, in 16 fathoms. 



2 2 6 Memoir Sears Foundation for Marine Research 

from the offing of Charleston, South Carolina. But it is questionable whether it ranges 
regularly any farther south than this, although it has been credited to Florida. ^^ 

Synonyms and References: 

Raja^" laevis Mitchill, Amer. Mon. Mag., 2, 1817: 327 (descr., New York); DeKay, Zool. N. Y ., 4, 1842: 
370 (descr., New York); Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 259 (in synopsis); Perley, 
Rep. Fish. Bay of Fundy, 1851: 154; also Cat. Fish. N. Brunsw. and Nova Scotia, 1854: 38 (food, size. 
Bay of Fundy); Wynian, Proc. Boston Soc. nat. Hist., 6, 1859: 44 (electric organs); Gill, Proc. Acad, 
nat. Sci. Philad., Addend., 1861: 61 (listed, Atlant. Coast N. Amer.); Knight, Descr. Cat. Fish. Nova 
Scotia, 1866: 8 (descr.. Nova Scotia); Storer, Mem. Amer. Acad. Arts Sci., N. S. 9, 1867: 242; also 
Fishes Mass., 1867: 266 (descr. in part but not ill., pi. 39, fig. 2, which is R. radiatd)\ Abbott, in Cook, 
Geol. N. J., 1868: 829 (util, Delaware R.); Gunther, Cat. Fish. Brit. Mus., 8, 1870: 455 (footnote); 
Lyman, 6th Rep. Mass. Comm. inl. Fish., 1872: 53 (nos., size, south. Mass.); Baird, Rep. U. S. Comm. 
Fish. (1871-1872), 1873: 826 (listed, Woods Hole); Gill, Rep. U. S. Comm. Fish. (1871-1872), 
1873: 812 (listed, Nova Scotia to Florida); V'errill and Smith, Rep. U. S. Comm. Fish. (1871-1872), 
1873: 521 (food. Woods Hole); Garman, Proc. Boston Soc. nat. Hist., IJ, 1874: 180 (descr., color, 
size, Massachusetts Bay); Uhler and Lugger, Rep. Comm. Fish. Md., ed. I, 1876: 189; ed. 2, 1876: 
160 (Maryland, Chesapeake Bay); Yarrow, Proc. Acad. nat. Sci. Philad., 29, 1877: 217 (listed, near 
Beaufort, N.Carolina); Goode and Bean, Bull. Essex Inst. Salem, II, 1879: 28 (N. Massachusetts); 
Jones, List Fish. Nova Scotia, 1879: 11 (Bay of Fundy); Verrill, Amer. J. Sci., 22 (3), 1881 : 297 (43-202 
fath., off Gay Head); Jones, Proc. N. S. Inst. Sci., 5 (l), 1882: append. 97 (Bay of Fundy); Jordan and 
Gilbert, Bull. U. S. nat. Mus., 16, 1883: 42 (descr., Virginia); True, List Vert. Anim. S.Carolina, 
in Handb. S.Carolina, 1883: 261 (listed, S.Carolina); Bean, Rep. U. S. Comm. Fish. (1881), 1884: 
343 (listed, Woods Hole); Goode, Fish. Fish. Industr. U. S., i, 1884: 667 (util); Rathbun, Proc. U. S. 
nat. Mus., 7, 1884: 486 (copepod parasites, depth, off Woods Hole); Garman, Bull. Mus. comp. Zool. 
Harv., 17, 1888: 90, pis. 27-29 (mucous canal syst.); Jordan, Rep. U. S. Comm. Fish. (1885), 1887: 
799 (listed, N. Atlant. states, U. S.); Bean, Bull. U. S. Fish Comm., 7, 1889: 151 (New Jersey); Proc. 
U. S. nat. Mus., 14, 1891 : 94 (Chesapeake Bay); Moore, Bull. U. S. Fish Comm., 12, 1894: 358 (New 
Jersey); Smith, Bull. U. S. Fish Comm., 12, 1894: 368 (nos., util., weight, New Jersey); Goode and 
Bean, Smithson. Contr. Knowl., JO, 1895: 28; jr, 1895: pi. 9, fig. 29; also Spec. Bull. U. S. nat. Mus., 
1895; and Mem. Harv. Mus. comp. Zool., 22, 1896: 28, pi. 9, fig. 29 (ill., depth, continental slope 
ofFNew England); Smith, Bull. U. S. Fish Comm., 14, 1895: 345 (common names, mid-Atlant. U. S.); 
Cox, Bull. nat. Hist. Soc. New Brunsw., 13, 1896; 63 (listed, N. Brunswick); Jordan and Evermann, 
Bull. U. S. nat. Mus., ^.j (i), 1896: 71 (descr.. New England to Florida); Rep. U. S. Comm. Fish. 
(1895), 1896: 221 (listed. New England to Florida); Bean, Bull. Amer. Mus. nat. Hist., 9, 1897: 423, 
441 (cestode parasites. Woods Hole); Proc. U. S. nat. Mus., 20, 1898: 507, 522 (trematode parasites); 
Smith, Bull. U. S. Fish Comm., 27, 1898: 89 (season. Woods Hole); Garman, Mem. Harv. Mus. comp. 
Zool., 24, 1899: 380 (depth, range); Bean, 6th Rep. For. Comm. N. Y., 1900: 383 (New York and 
C. Cod); Linton, Bull. U. S. Fish Comm., ig, 1901 : 431 (parasites, food, depth, off Woods Hole); Bean, 
Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 50 (descr., season, near New York and Woods Hole); Sharp 
and Fowler, Proc. Acad. nat. Sci. Philad., ^6, 1904: 506 (listed, Nantucket); Linton, Bull. U. S. Bur. 
Fish., 24, 1905: 346 (food, cestode parasites, C. Lookout, N. Carolina); Wilson, Proc. U. S. nat. Mus., 
28, 1905: 573, 582 (copepod parasites); Fowler, Rep. N. J. Mus. (1905), 1906: 73, pi. 3 (descr., ill.. 
New Jersey); Tracy, 36th Rep. R. I. Comm. inl. Fish., 1906: 47 (season, food, size, Rhode Island); 
Fowler, Rep. N. J. Mus. (1906), 1907: 259 (season. New Jersey); Smith, N. C. geol. econ. Surv., 2, 
1907: 41 (food, near C. Lookout, N. Carolina); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 
13 (New England locals.); Fowler, Proc. Acad. nat. sci. Philad., 61, 1909: 407 (New Jersey); Rep. 
N. J. Mus. (1908), 1909: 353 (Delaware R., by ref to Abbott, 1868); Tracy, 40th Rep. R. I. Comm. 
inl. Fish., 1910: 62 (Rhode Island); Cornish, Contr. Canad. Biol. (1906-1910), 1912: 79 (nos., Prince 
Edward I.); Gudger, Proc. biol. Soc. Wash., 26, 1913: 98 (season, N.Carolina); Engelhardt, Abh. 

59. Egg cases with embryos, washed ashore near Charlotte Harbor many years ago, were doubtfully referred to 
this species (Henshall, Bull. U.S. Fish Comm., 2, 1891:372). 

60. Sometimes spelled Raia. 



Fis/ies of the JVestern North Atlantic 22 j 

bayer. Akad. Wiss., Suppl. 4 (3), 191 3: 102 (range); Halkett, Check List Fish. Canad., 191 3: 42 (listed, 
Maritime Provinces to Florida); Nichols, Abstr. Linn. Soc. N. Y., 20—23, 1913: 91 (nos., season, near 
New York); Sumner, Osburn and Cole, Bull. U. S. Bur. Fish., JJ, 1913: 738 (food, parasites, nos., 
season, near Woods Hole); Fowler, Copeia, 13, 1914: [2] (listed. New Jersey); Proc. Acad. nat. Sci. 
Philad., 66, 1914: 353 (season, New Jersey); Coles, Proc. biol. Soc. Wash., 28, 1915: 92 (season, off 
C. Lookout, N.Carolina); Wilson, Proc. U. S. nat. Mus., ^7, 1915: 641 (copepod parasites, Maine); 
Fowler, Copeia, 31, 1916: 41 (depth. New Jersey); Latham, Copeia, 41, 1917: 18 (season, size. Long 
Island, New York); Fowler, Proc. Acad. nat. Sci. Philad., yi, 1920: 293 (New Jersey); Proc. biol. Soc. 
Wash., 33, 1920: 145 (New Jersey); Cox, Contr. Canad. Biol. (1918-1920), 1921: 109 (S. side Gulf 
St. Lawrence); Townsend, N. Y. zool. Soc, 25th Annu. Rep. (1920), 1921: 126 (listed. New York 
Bay); Scott, Contr. Canad. Biol., N. S. 2 (8), 1924: 131 (pathol. thyroid gland); Trans, roy. Soc. Canad., 
(3) 20 Sect. 5 (2), 1926: 229 (histol., physioL, thyroid gland); Burwash, Contr. Canad. Biol., 4, 1929: 
117, 118 (iodine content, thyroid gland); Fowler, Proc. Acad. nat. Sci. Philad., 80, 1929: 608 (weight, 
season, New Jersey); Jordan, Manual Vert. Anim. NE U. S., 1929: 17 (diagn., size, name); Truitt, 
Bean and Fowler, Bull. Md. Conserv. Dep., 3, 1929: 31 (descr., nos., Atlant. Coast Maryland, and 
Chesapeake Bay); Bere, Contr. Canad. Biol., 5, 1930: 423, 426 (copepod parasites, Passamaquoddy 
Bay); Jordan, Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 25 (listed, NE coast 
N. Amer.); Vladykov and McKenzie, Proc. N. S. Inst. Sci., ig, 1935: 51 (ill., depth. Nova Scotia); 
V'ladykov, Nat. canad., 63, 1936: 216 (descr., photo egg cases, breeding season, depth, temp, off N. 
Scotia and Kennebecasis Bay, New Brunswick); Breder and Nichols, Copeia, 1937: 182 (egg cases); 
Fowler, Proc. Acad. nat. Sci. Philad., 8g, 1937 : 304 (New Jsersey); Merriman and Warfel, Bull. Bingham 
oceanogr. Coll., II (4), 1948: 134 (listed, off Connecticut). 

Rata chantenay Lesueur, J. Acad. nat. Sci. Philad., 4, 1824: 106, pi. 5 (descr., ill., no local.); Storer, Mem. 
Amer. Acad. Arts Sci., N. S. 2, 1846: 512 (after Lesueur, 1824). 

Raia batis Storer, Boston J. nat. Hist., 2, 1839: 545 (descr., size, Mass.); Rep. Fish. Rept. Birds Mass., 1839: 
193 (same as preceding); Linsley, .^mer. J. Sci., 4^, 1844: 77 (Connecticut). 

Barn Door Skate, Wilson, Proc. U. S. nat. Mus., 31, 1907: 711 (copepod parasites); Bull. U. S. nat. Mus., 
158, 1932: 399, 403, 419 (copepod parasites. Woods Hole). 

Raia^^ stabuHforis Garman, Mem. Harv. Mus. comp. Zool., 36, 1913: 341, pi. 22, fig. 2, pi. 44, figs. 4-6 
(descr., ills., name R. laevis considered preoccupied); Kendall, Proc. Portland Soc. nat. Hist., 3 (i), 
1914: 13 (Maine locals.); Radcliffe, Trans. Amer. Fisher. Soc, 44, 19 14: 38 (teeth); Bull. U. S. Bur. 
Fish., 34, 1916: 272, pi. 45 (descr., photos teeth, C. Lookout, N. Carolina); Jordan, Copeia, 49, 1917: 
88 (R. StabuHforis replaces R. laevis, preoccupied); Huntsman, Contr. Canad. Biol. (1921), 1922: 57 
(Passamaquoddy Bay and Bay of Fundy); Breder, Copeia, 127, 1924: 28 (season, temp., near New 
York); Linton, Proc. U. S. nat. Mus., 64 (21), 1924: 9, 11, 20, 24, 28, 66 (cestode parasites); Bigelow 
and Welsh, Bull. U. S. Bur. Fish., 40 (i), 1925: 66 (descr., ill, habits. Gulf of Maine); Manter, Illinois 
biol. Monogr., 10 (2), 1926: 8, 9, 16, 43 (parasites); Nichols and Breder, Zoologica N. Y., 9, 1927: 
25 (ill., habits, near New York and south. New England); Hildebrand and Schroeder, Bull. U. S. Bur. 
Fish., 43 (i), 1928: 59 (descr., Chesapeake Bay); Radcliffe, Nat. Hist. N. Y., 28, 1928: 58 (abnormality. 
Woods Hole); Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 32 (genl.); Marshall, Contr. Canad. 
Biol., 7 (1-7), I93I'- 19 (heart physioL); Anon., Rep. Newfoundland Fish. Res. Comm., I (4), 1932: 
107 (depth, SW. part Grand Banks); Gudger, J. Elisha Mitchell sci. Soc, 4g, 1933: 63 (abnormal 
dentition); Amer. Mus. Novit., 600, 1933: i (ill., abnormality. Woods Hole); Anon., Rep. Newfound- 
land Fish. Res. Comm., 2 (i), 1933: 125 (Grand Banks, Banquereau Bank off Cape Breton); Bigelow 
and Schroeder, Canad. Atlant. Fauna, I2e, 1934: 35 (descr., ill., depth. Gulf of St. Lawrence, Grand 
Banks to Florida); Anon., Rep. Newfoundland Fish. Res. Comm., 2 (2), 1934: 115 (Banquereau and 
C. Breton, Nova Scotia); Roth, Arch. int. Pharmacodyn., 4g (3), 1935: 355 (cardiac regulative nerves; 
not seen); Bigelow and Schroeder, Bull. U. S. Bur. Fish., 48, 1936: 325 (nos., size, depth, Georges 
Bank and Nantucket Shoals); Smith, H. W., Biol. Rev. (Cambridge), 11, 1936: 67 (blood serum, urine); 
Norman and Fraser, Giant Fishes, 1937: 68 (.'\tlant. Coast N. Amer.); Breder, Bull. N. Y. zool. Soc, 
41, 1938: 28 (listed, New York Harbor); Bigelow and Schroeder, Biol. Bull. Woods Hole, 76, 1939: 
309 (Gulf of Maine); Merriman and Warfel, Trans. 9th N. Amer. Wildl. Conf., 1944: 234 (mean 
catch per hour. Long Island Sound). 

6 1 . Sometimes spelled Kaja. 

>5* 



2 2 8 Memoir Sears Foundation for Marine Research 

Probable Synonym: 

Raia granulata Goode and Bean, Bull. Essex Inst., ll, iSj(): 28; Smithson. Contr. Know!., JJ, 1895: pi. 9, 
fig. 30; Mem. Mus. comp. Zool. Harv., 22, 1896: pi. 9, fig. 30 (see discussion, p. 216). 

Doubtful Reference: 

Raia laevis {}) Henshall, Bull. U. S. Fish Comm., 9, 1891 : 372 (embryos, doubtfully ident.. Stump Pass, SW. 
Florida); Evermann and Kendall, Rep. U. S. Comm. Fish. (1899), 1900: 49 (SW. Florida, by ref to 
Henshall, 1891). 

Not Raja laevis Gronovius, Zoophyl., J, 1763: 37, no. 158, pi. 9, fig. 3 (equals Narcine brasiliensis Olfers 
1 831; see p. 112, footnote 129); Valmont, Diet. Hist. Nat., J, 1768: 706 (equals Raja oxyrincha Lin- 
naeus 1758). 

Not Raia laevis alba Duhamel, Traite Peches, (2) J Sect. 9, 1782 : 284, pi. 1 1 (equals R. talis Linnaeus 1758). 

Not Raia laevis major Duhamel, Traite Peches, (2) J Sect. 9, 1782: 285 (equals R. marginata Lacepede 1803). 

Not Raia laevis Seal, Aquarium, 2 (10), 1914: 105 (probably not R. laevis, because female only 15 in. wide 
with eggs; perhaps R. ocellata, see p. 251). 

Not Raia stabuliforis von Bonde and Swart, Fish. Mar. biol. Surv. S. Afr., Rep. 3 (1922), Spec. Rep. 5, 1924: 
12 (a South African form subsequently ref to the synonomy of ^. marginata Lacepede 1803 by Barnard 
[Ann. S. Afr. Mus., 21 (i), 1925: 65] and to that of the South African representative of ^. batis Lin- 
naeus 1758 by Norman ['Discovery' Rep., 12, 1935: 39]). 

Not Raia stabuliforis Marini, Physis B. Aires, 9, 1928: 138 (equals R.favirostris Philippi 1892). 



Raja lentiginosa Bigelow and Schroeder 1951 
Figure 48 a 

Study Material. Male, 402 mm in total length, and female, 302 mm, from Lat. 
22°32' N, Long. 88°47' W, 29 fathoms, Oregon St. 222; male, 323 mm, from Lat. 
28°io'N, Long. 85°oo' W, 85 fathoms, Oregon St. 256; a male and two females, 
151-234 mm, from Lat. 2 8°4i' N, Long. 86°03' W, 165 fathoms, Oregon St. 257; 
a male, 395 mm, from Lat. 2 8°49' N, Long. 85°45' W, 1 12 fathoms, Oregon St. 278 ; 
a male, 415 mm, from Lat. 29°! i' N, Long. 86°52' W, 305 fathoms, Oregon St. 279; 
and a female, 375 mm, from Lat. i(fl']' N, Long. 87°26'W, 104 fathoms, Oregon 
St. 318; and a male and two females, 330-402 mm, from Lat. 27°5o' N, Long. 
91°! I ' W, 205 fathoms, Oregon St. 500; all in the collections of the U. S. National 
Museum and the Harvard Museum of Comparative Zoology. 

Distinctive Characters. Raja lentiginosa closely resembles R. garmani, but it may 
be distinguished from the latter by the color pattern of its upper surface, which is densely 
freckled with small dark, light brown and whitish spots (sparse in garmani and grouped 
mostly in a distinct rosette pattern). 

Description. Proportional dimensions in per cent of total length. Male, 402 mm 
(U. S. Nat. Mus., type, No. 153552), and female, 302 mm (Harv. Mus. Comp. Zool., 
No. 37188), from Lat. 22°32' N, Long. 88°47' W, 29 fathoms. 
Disc: extreme breadth 60.0, 52.3; length 46.5, 44.3. 
Snout length: in front of orbits 9.2, 9.7; in front of mouth 1 1.2, 10.3. 
Orbits: horizontal diameter 4.2, 4.7; distance between 2.9, 2.8. 
Spiracles: length 2.1, 2.5; distance between 6.2, 6.3. 
Mouth: breadth 6.^, 7.2. 



-M^t<^i'\^T^^2^!^ji^?:rflll>— 




Figure 48a. Raja lentiginosa, male, 402 mm long, from Campeche Bank, Lat. 22°32'N, Long. 88°47' W 
(U. S.Nat. Mus., No. 153552, type). A Upper teeth, about lox. B Thorns in midrow on tail, about 2X. 
C Mouth and nostrils, about 1.5 X. D Posterior part of tail, about iX- E Section of upper surface to show 
color pattern. 



230 Memoir Sears Foundation for Marine Research 

Nostrils: distance between inner ends 6.0, 6.2. 

Gill openings: length, ist 2.0, 1.5; 3rd 1.9, 1.8; 5th 1.2, 1.5; distance between 

inner ends, ist 13.9, 14.3; 5th 7.0, 7.8. 
First dorsal fin: vertical height 3.0, 2.0; length of base 5.6, ^.■t,. 
Second dorsal fin: vertical height 3.0, 2.0; length of base f,.^, 4.3. 
Pelvics: anterior margin 12.2, ii.o. 
Distance: from tip of snout to center of cloaca 39.2, 41.3 ; from center of cloaca 

to 1st dorsal 45.3, 44.0; to tip of tail 60.8, 58.7; from rear end of 2nd 

dorsal base to tip of caudal 2.9, 2.3. 
Interspace between: ist and 2nd dorsals 1.2, 2.0. 

Disc about 1.2 — 1.3 times as broad as long, the maximum angle in front of spiracles 
about 120°; anterior margins weakly convex from just posterior to tip of snout to 
opposite orbits, thence gently concave between spiracles and outer corners, which are 
broadly rounded; posterior margins and corners and inner margins all rounded. Axis 
of greatest breadth about 68 "/o of distance back from tip of snout to axils of pectorals. 
Tail with lateral fold low down on each side beginning abruptly posterior to axils of 
pelvics by a distance about equal to space between spiracles and continuing almost to 
extreme tip; width of fold about the same throughout its length; length of tail from 
center of cloaca to origin of first dorsal fin i.i— 1.2 times as great as, its length to tip 
1.4— 1.5 times as great as, distance from center of cloaca to tip of snout. 

Four to six prominent thorns along anterior edge of orbit, none or one opposite 
its inner central margin, and two to five along its posterior margin, the last opposite 
spiracle; an additional thorn inward from this last one; a row of thorns over margins 
of rostral process; prickles and small thorns present over anterior third of disc in 
advance of nuchal region, extending along outer margin rearward about to axils of 
pectorals; a triangular patch of about 15—17 thorns on shoulder region, 4 or 5 of which 
extend along median line from nuchal to scapular region ; a naked area behind these 
for a distance about equal to length of distance between spiracles on males (not on 
females); band of 3—5 rows of thorns along median zone of back and tail from nuchal 
region to first dorsal, the thorns rather regular on some specimens but irregular on 
others, the lowermost row reaching nearly to tip of tail, the median row consisting of 
about 45—53 thorns in most cases, large and small alternating on males but all of about 
equal size on females; most of the thorns on tail with sharp points, directed rearward; 
1—4 prominent thorns in space between dorsals; prickles on dorsal fins and caudal; 
skin over eye naked. Males with a band of alar spines in one to three rows on outer 
part of each pectoral, the longest row with 19 spines. Female of 302 mm with small 
thorns on posterior part of pectorals and a few on pelvics. Lower surface naked except 
for a small median patch of spines at extreme tip of snout, lacking on our specimens 
of 232 mm and smaller. 

Snout in front of orbits 2.0—2.2 times as long as orbit, its length in front of mouth 
about 1.7— 1.8 times as great as distance between exposed nostrils. Orbit about twice 



Fishes of the JVestern North Atlantic 231 

as long as spiracle; distance between orbits about 0.6—0.7 ^^ great as length of orbits. 
Distance between first gill openings 2.3 times as great as distance between exposed 
nostrils; between fifth openings 1.2— 1.3 times; first gill openings i.o*-— 1.6 times as 
long as fifth and 0.2-0.3 as long as breadth of mouth. Nasal curtain fringed; expanded 
posterior (outer) margins of nostrils fringed. Upper and lower jaws rather strongly 
arched centrally. 

Teeth ^gZl^, close-set; those in nearly mature males mostly in straight series (not 
in quincunx), with small base, circular or oval, those in median sector of mouth with 
slender sharp cusp pointing toward symphysis or inward toward throat, those in outer 
sector with triangular cusp pointing toward corner of mouth, the one series of teeth at 
symphysis in upper jaw pointing straight downward; those in female in quincunx, 
ovate, with low triangular cusp not pointing inward or outward; no symphysis tooth. 

First and second dorsals similar in size and shape. Interspace between dorsals 0.22 — 
0.38 as long as base of first dorsal. Caudal membrane from rear end of base of second 
dorsal about half as long as base of first dorsal. Pelvics deeply concave, strongly scalloped 
along anterior side of excavation but only weakly so rearward; anterior margin 0.52 — 
0.55 as long as distance from its own origin to rear tip of pelvic; anterior lobe slender, 
including four radial cartilages besides the first stout one; posterior lobe moderately 
convex along its forward half, thence nearly straight to its narrowly rounded tip, ex- 
tending a little more than 1/5 of the distance from axil of pelvics toward first dorsal; 
inner margin straight. Claspers on 402-mm male reaching beyond tips of pelvics by 
a distance about equal to diameter of orbit. 

Rostral cartilage firm and extending nearly to tip of snout. Anterior pectoral rays 
reaching about '/lo of the distance from front of orbits toward tip of snout. 

Color. Upper surface sprinkled everywhere with small light to dark brownish and 
whitish spots, including the tail, pelvics and claspers; many groups of about 30—50 
dark spots scattered everywhere, the most prominent marking being the group of spots 
at axil of pectoral ; some of spots on tail grouped in form of about five or six prom- 
inent bars; light and dark spots also present on anterior part of each dorsal fin and on 
caudal. Whitish below; a group of grayish blotches, mostly fused, on each pectoral, 
and an elongate blotch of the ground color of upper surface along inner part of 
claspers, anteriorly, on nearly mature males. Younger specimens of both sexes with- 
out these blotches. 

Si-ze. On a 232-mm male the claspers extend a little beyond the tips of the 
pelvics, while on a 285-mm specimen of R. garmani they fail to reach the tips of the 
pelvics by a distance as great as the diameter of the orbit. This suggests that lentiginosa 
matures at a length a little below 400 mm (see garmani., p. 204). 

Occurrence. This newly discovered species is apparently widespread in the Gulf 
of Mexico, for it has been trawled in depths of 29, 85, 104, 112, 165, 205 and 305 
fathoms and in bottom temperatures ranging from 48.5° F (305 fath.) to 73.4° F 

62. It is exceptional to find a Raja with the fifth pair of gill openings as long as the first pair, but such was the case 
on the female 302 mm long. 



232 Memoir Sears Foundation for Marine Research 

(29 fath.). It is known from the northern part of the Gulf in the offings of Pensacola 
and Cape San Bias, Florida and in the southern part on Campeche Bank. 

Reference : 

Raja kntiginosa Bigelow and Schroeder, J. Wash. Acad. Sci.,^ (12), 195 1 : 383 (descr., ill., Gulf of Mexico). 

Raja lintea Fries 1838 
Figures 49, 50 

Study Material. Female, 1,050 mm long, and adult male, 970 mm, from Iceland 
(British Museum [Natural History]). 

Distinctive Characters. R. lintea falls in the group in which there is a single row of 
large thorns along the median belt of the disc and in which the thorns of the median 
row along the tail are much larger than those of any other rows. Among Skates of the 
western North Atlantic, it is closest to R. hyperborea and R.jenseni in general appearance 
and in the arrangement of thorns. But it is sharply marked off from both of these by 
a considerably larger number (47—51) of thorns in the mid-dorsal row from nuchal 
region to first dorsal fin and by its longer and narrower snout, the distance from its tip 
to the level of the front of the orbits being considerably greater than the distance be- 
tween the outer edges of the spiracles (shorter than that in R. hyperborea and in R.jenseni). 
R. lintea does not bear a close resemblance to any Skate so far described either from 
boreal latitudes in the western South Atlantic or from the Bering Sea-Northeast Siberian 
region of the Pacific. 

Description. Proportional dimensions in per cent of total length. Male, 970 mm, 
and female 1,050 mm, from Iceland (specimens from British Museum [Natural History]). 

Disc: extreme breadth 65.7, 64.0; length 54.3, 54.3. 

Snout length: in front of orbits 14.8, 16.5; in front of mouth 16.3, 18.2. 

Orbits: horizontal diameter 3.6, 4.0; distance between 5.1, 4.1. 

Spiracles: length 2.6, 2.6; distance between 8.0, 8.1. 

Mouth: breadth 10.4, lo.o. 

Nostrils: distance between inner ends 8.6, 9.3. 

Gill openings: lengths, ist 1.3, 1.2; 3rd 1.4, 1.5; 5th 0.9, 0.9; distance between 
inner ends, ist 15.5, 15.3; 5th 9.7, 9.7. 

First dorsal jin: vertical height 3.3, 2.6; length of base 4.9, -. 

Second dorsal jin: vertical height 2.4, 2.4; length of base 4.6, 4.3. 

Pelvics: anterior margin 9.0, 9.5. 

Distance: from tip of snout to center of cloaca 5 1 .0, 53.2 ; from center of cloaca 
to ist dorsal 38.8, 37.2; to tip of tail 49.0, 46.8; from rear end of 2nd 
dorsal base to tip of caudal 0.5, 0.4. 

Interspace between: ist and 2nd dorsals 2.0, 0.0. 

Disc about 1.2 times as broad as long, the maximum anterior angle in front of 
spiracles about 85-90°; tip of snout rounded, rather markedly protruding in male but 



Fishes of the Western North Atlantic 



233 




Figure 49. Raja lintea, male, 970 mm long, from Iceland (British Museum [Natural History]). A Side view 
of posterior part of tail of same, about 0.5 X . B Upper teeth of same from near center of jaw, about 5.5 X. 
C Upper teeth of female, 1,050 mm (British Museum [Natural History]). 

less so in female; anterior margins of pectorals in adult male deeply concave just posterior 
to snout and again at level of spiracles, but only weakly incurved posterior to level of 
spiracles in adult females; otherwise nearly straight; outer and posterior corners broadly 
rounded, the latter continuously so to axils of pectorals; posterior margins nearly straight 
or weakly convex. Axis of greatest breadth about 70 «/o of distance rearward from tip 
of snout toward level of axils of pectorals. Tail with narrow lateral folds along posterior 



2 34 Memoir Sears Foundation for Marine Research 




Figure 50. Raja Hntea. A Lower side of male pictured in Fig. 49. B Margin of nasal curtain, about 2.1 x. 
C Nostril of same, about 2.5 X. D Outline of snout of female of about the same size (British Museum [Natural 
History]). E Ventral view of pelvics of same, about 0.2 X. 

half to third; its length from center of cloaca to first dorsal about 0.7 as great, to tip 

of tail about 0.9 as great, as distance from center of cloaca to snout on large specimens. 

Upper surface of half-grown and larger specimens with a single median row of 



Fishes of the Western North Atlantic 235 

large thorns; about four anterior to pectoral girdle, about 11 -12 from pectoral girdle 
to level of axils of pectorals, and about 35—36 along tail to first dorsal (no thorn, however, 
between first and second dorsals when these are separated); each side of tail with 1—3 
irregular rows of smaller thorns anteriorly, one row posteriorly, extending to level of 
origin of second dorsal ; disc also with two large thorns (sometimes worn flat) on each 
shoulder in small specimens, three in larger, these increasing in relative size with growth; 
3—5 smaller thorns inward from posterior part of spiracle, 1—2 larger ones close in front 
of orbit, one large one just behind orbit and 2—3 around its inner edge; rostral ridge 
with an irregular band of small thorns, but tip of snout bare; outer anterior margins of 
pectorals more or less prickly along a narrow band; regions opposite eyes (sparsely 
prickly on partly-grown specimens) with a patch of thorns, large to small, in adults of 
both sexes; pelvics smooth, but upper portions of dorsals prickly. Alar spines of mature 
males in 2—3 rows. Lower surface smooth in young, in mature males and in some adult 
females, but other mature females are described as having a patch of 2-3 medium-sized 
thorns a little back from tip of snout. *^ 

Snout anterior to orbits about 2.8—4.0 times as long as distance between orbits, its 
length in front of mouth about two times as great as distance between exposed nostrils. 
Orbit about 80 "/o as long as distance between orbits and about 1.5 times as long as 
spiracle. Distance between first gill openings 1.6— 1.8 times, between fifth gill openings 
i.o— I.I times, as long as distance between exposed nostrils; first gill openings 1.3— 1.4 
times as long as fifth and about 1 2 "/g as long as breadth of mouth. Nasal curtain con- 
spicuously fringed, but expanded posterior (outer) margin of nostrils smooth. Mouth 
only weakly arched centrally, even in adult male. 

Teeth ^g (in two specimens), rather widely spaced in transverse series, with conical 
cusp, only a little longer and sharper in adult males than in females; the bases ovoid 
transversely. 

First and second dorsals either confluent at base or separated by a short interspace 
without thorns; second dorsal extending nearly to tip of tail in specimens seen, with 
only faint indication of separate caudal membrane. Pelvics rather strongly scalloped 
around the concavity; anterior margin 51 "/, as long as distance from its own origin to 
rear tip of pelvic on one female, 43 "/o on a male; anterior lobe broadly subtriangular; 
posterior lobe weakly convex and only faintly scalloped; rear tip narrowly rounded in 
both sexes. 

Anterior rays of pectorals extending about 55—60 "/o of distance from level of fronts 
of orbits toward tip of snout. 

Color. Described as having upper surface plain fawn color, slate-gray or clay-gray 
when fresh (changing to chocolate brown in formalin). Lower surface uniformly white 
in some,** but others with a gray longitudinal band along tail and a gray blotch on either 
side of cloaca, or with outer posterior parts of pectorals and posterior lobes of pelvics 

63. Clark, Rep. Fish. Bd. Scot. (1926), i, 1926: 46. 

64. So described bv Clark (Rep. Fish. Bd. Scot. [1926], i, 1926: 46) on specimens from between Denmark and southern 
Norway. 



236 Memoir Sears Foundation for Marine Research 

broadly edged with dark gray, a dark gray spot near tips of anterior pelvic lobes, and 
irregular dark markings below tip of snout. 

Size. This is one of the larger Skates of the region, males maturing sexually when 
they are about 970 mm long (about 38Y2 in.). The maximum recorded length is about 
1,117 "^"^ (44 i'^O' '^^t t^^ ^'-^^ ^^ hatching is not known. 

developmental Stages. The egg case is about 107 mm long by 77 mm wide, exclu- 
sive of the horns; its walls have longitudinal striae and cross-hatching much as in R. ra- 
diata\ the lateral flanges are margined with fine threads; the tip of each of the longer 
pair of horns is produced into a fine point. *^ 

Habits. R. lintea is confined to water of at least moderate depth, all recorded cap- 
tures having been from 80 fathoms or deeper. Available evidence fails to establish the 
lower limits of its usual range, but it has been taken most often in water shoaler than 
350 fathoms. 

The bottom temperatures recorded for stations where R. lintea has been taken have 
ranged from 3.3° C (38° F) ofi-" West Greenland to about 6° C. Thus its thermal range 
appears to average a little higher than that oi R. fyllae (p. 199) and considerably higher 
than that of R. hyperborea (p. 211). 

The nature of its teeth suggests that R. lintea, like R. hyperborea., feeds chiefly on 
active prey, such as small fishes and Crustacea, but no examination appears to have been 
made of its stomach contents. 

Range. Fishing banks ofl^ southwestern Norway, Faroe-Shetland Bank, Skagerrak 
between southern Norway and Denmark, slope west of Ireland, and Iceland in the eastern 
North Atlantic;^' also on the West Greenland side of Davis Strait. 

Occurrence in the Western Atlantic. The only reason for mentioning R. lintea here is 
that its range extends to West Greenland, where three specimens have been taken at 
depths of 210— 318 fathoms in bottom temperatures of 3.3—3.9° C.^' Its restriction to 
bottom waters warmer than 3° C, and its apparent preference for depths less than 400- 
500 fathoms, seem likely to prevent its dispersal to the American slope. 

Synonyms and References: 

Raja^^ lintea Fries, K. svensk. VetenskAkad. Handl. (1837), 1838: 154 (descr., length, color, but not syno- 
nyms; off Bohuslan, Sweden); Miiller and Henle, Plagiost., 1841: 147 (descr., after Fries, 1838); Ek- 
strom, Goteborg VetenskSamh. Handl., (3) J, 1850: 41 (listed, Kattegat, but probably Skagerrak in 
reality); Kroyer, Danmark's Fisk., J (2), 1853: 1005 (descr., meas., ill., but not synonyms; off Stavangar, 
Norway, but Kattegat ref. probably Skagerrak in reality); Nilsson, Skand. Fauna, Fish., 4, 1855: 738 
(descr., but not synonyms, off Bohuslan, Sweden); Malm, Ofvers. Vet. Akad. Forh. Stockh., 1857: 
193 (size off Jaederen, Norway); Dumeril, Hist. Nat. Poiss., i, 1865: 557 (descr., size, but not syno- 
nyms, Sweden); Giinther, Cat. Fish. Brit. Mus., 8, 1870: 466 (descr., off Sweden); Olsson, .Acta Univ. 

65. For description and photograph of an e.%% case taken from a fish from Iceland, see Clark (Rep. Fish. Bd. Scot. 
[1926], I, 1926: 45, pi. 27, fig. b). 

66. K. lintea has been reported from South Africa also (Barnard, Ann. S. Afr. Mus., 21 [i], 1925: 72). But the Skate 
in question is clearly a different species, namely K. leoparda von Bonde and Swart 1924 (see Norman, Discovery 
Rep., 12, 1935: 44). 

67. The locahties are: Lat. 66°35' N, Long. 56°38' VV; Lat. 66°45' N, Long. 56°39' W; and Lat. 66°53' N, Long. 
56°i7'W. For details, see Jensen (Mindeskr. Steenslr. Fods. Kbh., 2 [30], 1914: 28). 

68. Also spelled Kaia. 



Fishes of the Western North Atlantic 237 

lund. (1871), 7, 1871-1872: 12 (listed); Collett, Forh. VidenskSelsk. Krist. (1882), 29, 1883: 4 (depth, 
Jaederens-Rev, near Stavanger, Norway); Lilljeborg, Sverig. Norg. Fisk., (7) J (2), i8gi: 580 (descr., 
but not synonyms, off Bohuslan, and Norwegian fishing banks; but ref. to Kattegat probably Skagerrak 
in reality); Collett, Forh. VidenskSelsk. Krist., 1905: [7], 122 (diagn., depth, banks offjaederen, Norway, 
Skagerrak); Smitt, Hist. Scand. Fish., 2nd ed., 1895: 1117 (descr., ill., but not synonyms, otFS. Norway, 
but ref. to Bohuslan probably Skagerrak); Jensen, in Schiodte and Hansen, Zool. Danica, 2 (12), 1907: 
338, pi. 32, fig. 2 (descr., ill., size, range, depth); Engelhardt, Abh. bayer. Akad. Wiss., Suppl. ^ (3), 
1913: 102 (range); Garman, Mem. Harv. Mus. comp. Zool., j6, 1913: 329 (descr., but color probably 
not correct); Jensen, Mindeskr. Steenstr. Fods. Kbh., 2 (30), 1914: 26, 38, fig. 12 (refs., descr., color, 
ills, teeth, depths, temps., chart of distrib., W. Greenland); Clark, Rep. Fish. Bd. Scot. (1926), i, 1926: 
45, pis. 25—27 (descr., ills, egg case and adults, of. other species, range); Faune Ichthyol. Cons. int. Explor. 
Mer, 1930: pi. not numbered (descr., ill., range); Jensen, Spol. Zool. Mus. Hauniensis, 9, 1948: 46 
(W. Greenland). 

Leucoraia lintea Malm, Goteborgs och Bohuslans Fauna, 1877: 611 (descr., depth, banks off S. Norway). 

Raja ingolfiana Lutken, Danish Ingolf Exped., 2 (i), 1898: 3, pi. I (descr., ill., depth, temp, off Holstenborg, 
W. Greenland); Garman, Mem. Harv. Mus. comp. Zool., j6, 1913: 327 (descr., local., after Lutken, 
i898).«9 

Not Raia lintea Barnard, Ann. S. Afr. Mus., 21 (i), 1925: 72 (equals Rata leofarda von Bonde and Swart 
[Fish. Mar. biol. Surv. S. Afr., Rep. 3 (1922), Spec. Rep. 5, 1924: 7, pi. 20, fig. 2] S. Afr.; see Norman 
[Discovery Rep., 12, 1935: 44]). 

Raja mollis Bigelow and Schroeder 1950 
Figure 51 

Study Material. Juvenile male, 262 mm long (type), from lower part of continental 
slope, south of southern Nova Scotia, Lat. 4i°53' N, Long. 6 5° 3 5' W,'" 858 fathoms, 
in U. S. National Museum. 

Distinctive Characters. R. mollis is characterized by the following features: a com- 
paratively short tai^i a wedge-shaped anterior contour, no conspicuous thorns anywhere 
on its disc posterior to the scapular region or on its tail, and rostral projection from its 
cranium soft from base to tip. In general form and in prickliness of the upper and lower 
surfaces of the tail and of the upper surface of its disc, it resembles R. spinicauda, as well 
as females and juvenile males oi R. senta. But it differs noticeably from R. senta in lacking 
large thorns anywhere on its disc posterior to the scapular region at a size at which the 
shoulder and mid-dorsal thorns are a conspicuous feature of i?. senta. Equally convenient 
external field marks for R. mollis., as contrasted with R. spinicauda, are the considerably 
greater length of its tail, the prickliness of the entire breadth of the lower surface of its 
tail (not only the outer edges), the lack of large mid-dorsal thorns on its tail, and the 
fact that its two dorsal fins are confluent. In the softness of the rostral projection from 
its cranium it appears to be unique among the members of the genera Raja and Breviraja 
known from the western Atlantic. Raja abyssicola Gilbert 1895, known from one speci- 
men from 1,588 fathoms off the coast of British Columbia, and R. stellulata Jordan and 
Gilbert 1880, from southern California to Alaska, appear to parallel R. mollis in this 
last respect. But each of them has a mid-dorsal row of thorns on the disc and on the tail. 

69. Spelled Raia. 70. Albatross Station 2072, September 2, 1883. 

7 1 . Not longer from axils of pelvics to first dorsal fin than from axils of pelvics to front of orbits. 



238 Memoir Sears Foundation for Marine Research 




Figure 51. Raja mollis, juvenile male, 262 mm long, from lower part of continental slope off southern Nova 
Scotia, Lat. 4i°53'N, Long. 65°35' W, in 858 fathoms (U. S. Nat. Mus., No. 33385, type), y/ Pelvic fins, ven- 
tral view. B Central part of anterior part of head, ventral view. C Nasal curtain and exposed nostril, about 3.6 X . 



Description. Proportional dimensions in per cent of total length. Juvenile male, 
262 mm long, from Lat. 4i°53' N, Long, (yfzs' ^ (U- S. Nat. Mus., type. No. 233S5). 
Disc: extreme breadth 61.0; length 49.7. 



Fishes of the Western North Atlantic 239 

Snout length: in front of orbits 14.9; in front of mouth 15.7. 

Orbits: horizontal diameter 3.8; distance between 3.8. 

Spiracles: length 2.1 ; distance between 7.1. 

Mouth: breadth 8.0. 

Exposed nostrils: distance between inner ends 8.6. 

Gill openings: lengths, ist 1.5; 3rd 1.5; 5th i.o; distance between inner ends, 

ist 13.2; 5th 8.8. 
First dorsal fin: vertical height 2.3; length of base 4.4. 
Second dorsal fin: vertical height 2.3; length of base 5.0. 
Pelvics: anterior margin 10.3. 

Distance: from tip of snout to center of cloaca 42.8; from center of cloaca to 
1st dorsal 44.4; to tip of tail 57.2; from rear end ot 2nd dorsal to tip of 
tail 3.4. 
Interspace between: ist and 2nd dorsals 0.0. 

Disc about 1.2 times as broad as long, subangular in front; tip of snout slightly 
blunted, projecting a little; maximum anterior angle in front of spiracles about 115°; 
anterior margins of pectorals from close behind tip of snout nearly straight or weakly 
concave abreast of eye and spiracle;''^ outer corners broadly rounded; posterior margins 
strongly convex; posterior corners well rounded with curvature continuous to axils of 
pectorals. Tail with narrow lateral folds confined to posterior third; distance from center 
of cloaca to first dorsal fin about equal to distance from center of cloaca to tip of snout, 
and distance to tip of tail about 1.3 times that great. Distance from axils of pelvics to 
first dorsal a little greater (i.i) than distance from axils of pelvics to front of orbits. 

Upper surface of disc (including skin above eyes) rugose with prickles or minute 
thornlets except close along posterior margins, some of them low and conical, others 
higher but straight, and others still higher, their tips very fine and curved rearward in 
varying degree; one or two somewhat larger thorns close in front of eye on juveniles," 
but perhaps more numerous on adults; one blunt and slightly larger thorn inward from 
each spiracle, one on nuchal region, one on mid-dorsal line over pectoral girdle, and a 
smaller one on each shoulder (perhaps more on adults); other than these no sizeable 
thorns on disc; upper surface of tail rough with prickles slightly larger than those on 
disc, its median line also with an irregular row of somewhat larger thornlets scattered 
from opposite tips of pelvics for about a third of the distance toward the first dorsal, but 
without conspicuous thorns; upper surface of pelvics prickly on inner parts of posterior 
lobe but otherwise smooth; dorsal fins prickly; caudal membrane with a few prickles. 
Lower surface smooth on disc, pelvics, and root of tail ; but remainder of tail close-set 
with small thornlets rearward to level of posterior end of second dorsal, similar to upper 
surface. 

Snout in front of orbits about 3.9 times as long as distance between orbits, its 

72. The two sides of our specimen differ slightly in this respect. 

73. One such thorn on one side of our specimen, two on the other. 



240 Memoir Sears Foundation for Marine Research 

length in front of mouth about 1.8 times as great as distance between exposed nostrils. 
Orbit about as long as distance between orbits and about 1.8 times as long as spiracle. 
Distance between inner ends of first gill openings about 1.6 times as long as distance 
between exposed nostrils, that between fifth gills about 2/3 (67 "/o) as long as distance 
between first gills; first gills about 1.5 times as long as fifth gills, about 19 "/o as long as 
breadth of mouth. Nasal curtain fringed, but edge of expanded posterior (outer) margin 
of nostril smooth or nearly so. Mouth nearly straight on juvenile male, probably so on 
females also, but perhaps more arched on mature males. 

Teeth j^; with low conical cusp; bases about as long anteroposteriorly as they are 
broad transversely, closely crowded in quincunx. Teeth of mature specimens not seen. 

First and second dorsal fins about equal in size, confluent at base without definite 
interspace, their shape as shown on Fig. 51. Caudal membrane posterior to second dorsal 
about ^/j (77 "/o) as long as base of first dorsal. Pelvics deeply concave outwardly, strongly 
scalloped around concavity; anterior margin nearly as long (90 "/o) as distance from 
pelvic origin to rear tip; anterior lobe spatulate, rounded at tip; posterior lobe moder- 
ately rounded outwardly, its inner margin straight; tip well rounded, extending rear- 
ward about Ys (21 "/o) °^ ^^^ distance from axil of pelvic toward first dorsal. 

Rostral cartilage extending nearly to tip of snout ; so soft and flexible that it can 
hardly be felt, but visible against a strong light. Tips of anterior rays of pectorals falling 
a little short of level of tip of rostral cartilage. 

Color. Upper surface of disc, tail and pelvics light grayish brown (after many years 
in alcohol), but either side of rostral ridge pale yellowish, perhaps translucent in life; 
no dark markings. Lower surface pale yellowish without markings. 

Size. It is not known how large this Skate may grow; the only specimen seen is 
a juvenile male, its claspers reaching only ^/j of the distance rearward along the inner 
margins of the pelvics. 

Habits. The great depth at which this one known specimen was trawled (858 fath.), 
plus its absence from the catches of Skates that are made along the slopes of the fishing 
banks off Nova Scotia and the Gulf of Maine, mark this as a deep-water species, prob- 
ably confined to depths greater than 200—300 fathoms. 

Range. Known only from the lower part of the continental slope off" southern 
Nova Scotia. 

Reference : 

Raja mollis Bigelow and Schroeder, Bull. Mus. comp. Zool. Harv., 103, i9?o: 3^8 (descr., ill., continental 
slope off S. Nova Scotia). 

Raja ocellata Mitchill i 8 1 5 

Big Skate, Eyed Skate, Winter Skate 

Figures 52, ^':>,, 54 

Study Material. Thirty-two specimens, from about one-fourth grown to mature, of 
both sexes, from the Massachusetts Coast and Georges Bank, in Harvard Museum of 



Fishes of the JVester?! North AtUmtic 



241 




Figure 52. Raja ocellata, male, 809 mm long, from Nahant, Massachusetts (Harv. Mus. Comp. Zool., No. 352) 
A Side view of posterior part of tail of same, about 0.6 x . B Nostril of same, about 5.7 X. C Margin of left nasa 
curtain of same, about 5.7 X. 



242 Memoir Sears Foundation for Marine Research 

Comparative Zoology; also two eggs with embryos, one of them still with small yolk 
sac, the other about ready to have hatched. 

Distinctive Characters. Among the Skates of the geographic province where R. ocel- 
lata occurs, the ocellar spots usually present on the upper surface of its disc (p. 246) 
are a reliable means of identification for specimens that have them. However, specimens 
that lack these spots resemble R. erinacea so closely when small that the only certain 
means of distinguishing one from the other is by the number of series of teeth, R. ocellata 
having at least 80'* series in the upper jaw (usually 90-100), R. erinacea never more 
than 66 (usually less than 54). Adults of the two species can also be separated by size, 
R. ocellata not maturing until it reaches a length of at least 25-26 inches whereas R. eri- 
nacea matures at 18-20 inches, its maximum length being only about 21 inches. What 
is said on p. 178 as to the differences between R. erinacea and R. fyllae applies equally 
to R. ocellata. 

R. ocellata., like R. erinacea, falls closest to R. naevus Muller and Henle 1 84 1 among 
European Skates in shape of disc, relative length of tail, distribution of thorns, loss of 
the mid-dorsal row of thorns with growth, and the frequent presence of a large ocellar 
spot on the upper inner part of each pectoral. But the disc and anterior part of the tail 
are thornier in R. ocellata., especially on females, and the skin of its disc is smooth be- 
tween the thorns, whereas on R. naevus it is generally prickly between the thorns. R. ocel- 
lata does not bear a close resemblance to any Skate described from the South Atlantic 
or from the Pacific Coast of North, Central, or South America. 

Description. Proportional dimensions in per cent of total length. Female, 806 mm 
(Harv. Mus. Comp. Zool., No. 359). Male, 809 mm, (Harv. Mus. Comp. Zool.,No. 
352) from Massachusetts. 

Disc: extreme breadth 60.6, 64.8; length 46.6, 49.3. 

Snout length: in front of orbits i 1.6, 10.6; in front of mouth i 1.5, i i.i. 

Orbits: horizontal diameter 3.5, 4.1 ; distance between 5.3, 5.9. 

Spiracles: length 3.6, 3.5; distance between 7.8, 8.0. 

Mouth: breadth 9.9, lo.i. 

Nostrils: distance between inner ends 7.6, 6.9. 

Gill openings : lengths, ist 2.0, 2.0; 3rd 2.3, 2.3; 5th 1.7, 1.7; distance between 
inner ends, ist 16.0, 17.4; 5th 8.8, 8.4. 

First dorsal fin: vertical height 3.2, 3.3; length of base 7.6, 7.5. 

Second dorsal fin: vertical height 3.0, 2.6; length of base 6.7, 6.7. 

Pe/wVj; anterior margin 11.7, 14.8. 

Distance: from tip of snout to center of cloaca 50.6, 49.2 ; from center of cloaca 
to 1st dorsal 33.7, 34-7; to tip of tail 49.4, 50.8; from rear end of 2nd 
dorsal base to tip of caudal 1.5, 1.9. 

Interspace between: ist and 2nd dorsals 0.0, 0.0. 

74. Bigelow and Schroeder (Bull. U. S. Bur. Fish., 48, 1936: 324) give as few as 70, but re-examination of the specimens 
on which this count was based showed that the number should have been stated as 80. 



Fishes of the Western Northi Atlantic 



243 




Figure 53. Raja ocellata. A Female, 806 mm long, from southern Massachusetts (Harv. Mus. Comp. Zool., 
No. 359). B Lower surface of anterior part of head of male pictured in Fig. 52. C Upper teeth (front rows) 
of another adult male from Nahant, Massachusetts (Harv. Mus. Comp. Zool., No. 1 1 36), about 7 X. D Upper 
teeth from rearmost rows of same, about JX- E Upper teeth (front rows) of adult female from Waquoit, Massa- 
chusetts (Harv. Mus. Comp. Zool., No. 548), about 7 X . /" Upper teeth of lothand I ith rows of same, about 7X. 



Disc about 1.2— 1.3 times as broad as long, obtuse in front (averaging a little more 
so than in R. erinacea, p. 179); maximum anterior angle in front of spiracles 135—150°, 
irrespective of size; anterior margins weakly concave for a short distance close behind 

1 6* 



244 Memoir Sears Foundation for Marine Research 

tip of snout, then weakly" convex rearward in small specimens, weakly concave opposite 
spiracles in partly grown of both sexes and in adult females, and a little more deeply 
so in adult males; outer and posterior corners broadly rounded, posterior margins gently 
convex. Axis of greatest breadth on adult males about 75 "/o of distance back from snout 
toward axes of pectorals, about 70 '/o on adult females. Tail with narrow lateral folds 
beginning posterior to axils of pelvics by a distance equal to about twice diameter of eye 
and extending to extreme tip; its length from center of cloaca to origin of first dorsal 
about 80 »/o as great as distance from center of cloaca to tip of snout on small and 
medium-sized specimens, about 70 "/o on large; extreme length of tail from center of 
cloaca about i. 0-1.2 times as great as distance from cloaca to snout. 

Young specimens with 3-4 thorns on each shoulder and with others between orbits, 
surrounding eyes, and along margins of rostral ridge to tip of snout; a row of 12-16 
thorns, about uniform in size, along midline of disc from nuchal region; 16-18 on tail 
to first dorsal fin, this row flanked by a second less regular row of slightly smaller thorns 
from close behind pectoral girdle and by a third irregular row of smaller thorns low down 
on either side of midsector of tail. Outer anterior parts of pectorals loosely strewn with 
small thorns, but posterior corners and considerable areas along either side of mid-dorsal 
ridge bare. 

With growth, the thorns of the mid-dorsal row are lost from both the disc and tail,'* 
and those between the orbits decrease in number and relative size, while the bare areas 
expand over the inner parts of the pectorals and abreast of the spiracles, eyes, and rostral 
ridge; however, the thorns on the shoulders increase in number and a thorny area 
develops on the central part of the posterior lobe of each pelvic. 

Specimens that are V2-V3 grown have a patch of thorns on the tip of the snout, a 
regular or irregular series on each edge of the rostral ridge, a patch of small thorns in 
front of the orbit, a row around its inner margin, several larger ones (one usually notice- 
ably larger) just behind the orbit, and a few small ones between the orbits. The pectorals 
are rough with small thorns and prickles along the anterior margins nearly to the poste- 
rior corners, with larger thorns scattered abreast of the spiracles and eyes and on the 
posterior parts. Each shoulder bears a roughly triangular patch of medium-sized thorns 
(variable in number), the outermost 2-3 usually considerably largest; also, there are 2-3 
irregular rows of medium-sized thorns with radiate bases along either side of the mid- 
dorsal line from the pectoral girdle rearward past the first dorsal fin,'« these thorns de- 
creasing in size and in regularity rearward, not much larger in any one of these rows than 
in the other rows, and those lowest down on the sides of the tail smallest. The central 
parts of the posterior lobes of the pelvics are prickly at first but later become thorny. 
The first and second dorsals are prickly. Otherwise the skin is smooth, with extensive 
bare areas along either side of the mid-dorsal zone, alongside the spiracles and eyes, and 
between the rostral ridge and the anterior radial cartilages of the pectorals. 

75. Twenty inches is the greatest length at which a definite midrow of thorns still persists in any specimen that we 
have seen; they are usually lost by the time specimens are half-grown. 

76. Sometimes four rows locally on one side of the tail or the other. 



Fishes of the IVestern North Atlantic 245 

By maturity the thorns decrease in number on the shoulder regions, around the 
eyes, and on the rostral ridge and the bare areas expand over most of the inner parts of 
the pectorals, but mature females develop large thorns, widely spread, over the posterior 
corners of the pectorals, the thorny areas expanding over the posterior lobes of their 
pelvics; a few thorns also appear on the anterior lobes. Mature males develop 4—6 ir- 
regular rows of conspicuous thorns over a narrow triangular area from the tips of the 
pectoral cartilages back about to the level of the eyes. Alar spines of mature males are 
ordinarily in three rows, these appearing when the claspers extend about a third of the 
way back from the tips of the pelvics toward the end of the tail; 19—2 i spines in each 
of the two inner rows, 10—12 in the outermost. The lower surface is smooth in small 
specimens, but the tip of the snout becomes prickly with growth, as does a narrow band 
along the anterior margins reaching rearward to the level of the mouth in half-grown 
specimens and for some distance posterior to the mouth in adults. 

Snout in front of orbits about 2.0—2.3 times as long as distance between orbits; 
its length in front of mouth 1.4-1.7 times as great as distance between exposed nostrils. 
Orbit about 65—70 "/„ as long as distance between orbits, and i. 0—1.2 times as long as 
spiracle. Distance between first gill openings 2.1—2.5 times as long as distance between 
exposed nostrils, between fifth gill openings i. 0—1.3 times; first gill openings i. 0—1.2 
times as long as fifth and about 18—22 "/q as long as breadth of mouth. Nasal curtain 
fringed, expanded posterior (outer) margin of nostril with very short fringe, sometimes 
irregular and easily overlooked. Mouth but little arched, hardly more so in males than 
in females." 

Teeth yfZn^; those of young and of females to maturity rounded, or sometimes 
with blunt cusp faintly indicated, and close-set in quincunx; those of mature males with 
high conical cusp blunted at tip, more widely spaced in transverse series.'* 

First and second dorsals similar in size and shape, the two confluent at base. Caudal 
membrane posterior to second dorsal about Vi—Vs ^^ '°"§ ^® h'i.^^ of first dorsal. Pelvics 
moderately concave outwardly, weakly scalloped around the indentation ; anterior margin 
about 59-66 o/o as long as distance from its own origin to rear tip of pelvic; anterior 
lobe broad, including 4-5 radial cartilages besides the first and stoutest; posterior lobe 
weakly convex outwardly, its rear corner narrowly rounded. Claspers of sexually mature 
males extend about halfway from axils of pelvics toward tip of tail. 

Anterior rays of pectorals extending forward about 80 to 90 "/o of distance from 
level of fronts of orbits toward tip of snout in most specimens, reaching almost opposite 
tip of snout in others. 

Color. Light brown above, with varying numbers of rounded blackish spots; on 
half-grown specimens and larger, the spots usually about 3-6 mm in diameter and scat- 
tered 8-12 mm apart on disc, pelvics, tail, and claspers of mature males; also, outer 

77. In R. erinacea at maturity the lower jaw is considerably more arched in males than in females. 

7S. In maturing males the replacement of rounded teeth by conical ones takes place when the claspers are about half- 
grown; in a specimen of about Soo mm the lower rows of teeth are still of the immature shape, whereas in the 
upper jaw the teeth in successive rows are progressively more and more conical from front to rear; i. e., from the 
older rows of teeth to those more recently formed. 



246 Memoir Sears Foundation for Marine Research 

or posterior part of each pectoral often with 1-4 black or dark brown ocellar spots 
edged with white and var\nng from round to oval; sometimes 1—2 smaller ones nearby; 
size, arrangement and number of these spots varying from specimen to specimen; some- 
times each pelvic with a similar ocellar spot. Each side of snout in front of eyes with a 
whitish translucent area. Lower surface usually white but marked with irregular pale 
brownish blotches of various sizes'" on posterior part of disc and along tail of some 
specimens. 

Size. Specimens so recently hatched that their abdomens are still more or less swol- 
len with yolk range from about 1 12-127 rn^n in length, apart from whatever remnant 
of the embryonic caudal filament may still persist. In a male of 665 mm the claspers 
reach about to the tips of the pelvics; in one of 715 mm they extend about one-third of 
the way from the axils of the pelvics toward the tip of the tail ; and halfway in one 8 i o mm 
long. All of the males are sexually mature by the time they have reached a length of 
about 750 mm, some by 620 mm.*" The longest female we have seen measured 806 mm, 
the longest male 834 mm ; the maximum recorded length is about 43 inches ( i ,090 mm).*^ 
On the average, males about 20 inches long weigh 2 pounds; 30 inches, 7 pounds; 
32 inches, 9 pounds. Females about 20 inches long weigh 2 pounds; 25 inches, 4V2 
pounds; 28 inches, 6^2 pounds. ^^ 

Developmental Stages. The greenish brown or brownish olive egg cases are about 
55-86 mm in length by 35-52 mm in breadth, excluding the horns, and northern spe- 
cimens average larger than southern ones.*^ The end with the shorter pair of horns is 
weakly concave, that with the longer horns straight or weakly convex and more or less 
ragged. The horns are strongly flattened and each is described as having a slit on its 
outer edge near its tip.** The longer pair of horns on egg cases that have been laid is 
usually about 1.5 times as long as the shorter pair and about 1.5 times as long as the 
egg case, but the extreme tips are likely to have been broken off. The outer margin, 
close to each of the shorter pair of horns of newly laid cases, bears a short slender spur, 
the edge of which, like the remainder of the lateral border, is fringed with a series of 
silky filaments that are tangled and matted together. But these as well as the spurs are 
so easily torn off that only tufts remain here and there on cases that have been deposited 
for more than a brief period. No doubt the filaments serve as anchors.*^ 

Embryos ready for hatching already show the distinctive characters of the species 
so clearly that they are easy to identify. 

79. 15 by 15 mm to 3; by 74 mm in one specimen. 

80. This is the case in the Block Island region, according to the Bingham Oceanographic Laboratory. 

8 1. A maximum length of six feet credited to this species by Hildebrand and Schroeder (Bull. U. S. Bur. Fish., 43 
[i], 1928: 58) actually referred to R.lae'vis. 

82. We are indebted to the Bingham Oceanographic Laboratory for these data on maximum recorded lengths and 
on length-weights. 

83. 75-86x44-52 mm from Nova Scotia (Vladykov, Nat. canad., 63, 1936: 216); 66-70X45-45 mm for two from 
Provincetown, Massachusetts; and 64-70X 35-42 mm from New York (Breder, Copeia, 1937: 182). 

84. These slits, recorded by Wyman (Mem. Amer. Acad. Arts Sci., N. S. 9 [2], 1864: 31) are not visible on the pre- 
served specimens we have seen. 

85. For an account, with exellent illustrations, of the lateral spurs and filaments and of the embryonic development, 
see Wyman (Mem. Amer. Acad. Arts Sci., N. S. 9 [2], 1864: 32, 33, fig. i, 2-10). 



Fishes of the Western Nortli Atlantic 



247 



Habits. The Eyed Skate has much the same diet as R. erinacea (p. 183), crabs 
(Cancer) being the chief item where available; but they also take annelids, shrimps, 
bivalves, and whatever small fish they can capture.'' A specimen opened by us recently 
contained the body of an ascidian {Boltenia), the bristles from an annelid (apparently 
Aphrodite)., vertebrae of small fishes, and two sizeable pebbles. Another, 3 i inches long, 
taken off Massachusetts, was reported to have been full of beach fleas {Talitrus)\ one 
caught oft" northern Nova Scotia had eaten butterfish, cunners, and squid.*' 




Figure 54. Raja ocellata, egg case, opened to show enclosed embryo, from Woods Hole, Massachusetts, about iX. 



This Skate is closely confined to sandy or gravelly bottoms. No doubt this prefe- 
rence for hard bottom is the explanation for its reputed absence from Buzzards Bay 
as contrasted with its regular occurrence in season at depths no greater in Vineyard 
Sound nearby. 

Its depth range is wide, for while it is often speared from the wharves in the north- 
ernmost part of its range** and is taken in pound nets farther south, it is taken down to 
50—60 fathoms on the offshore fishing banks northward and eastward of Nantucket. But 
most of those we have seen from many otter trawl hauls off the southern New England 
Coast were taken shoaler than 40 fathoms.** The upper limit to its optimum thermal 

86. The list at Woods Hole includes smaller skates, eels, herring [Clupea), alewives [Pomolobus pseudoharengus), blue- 
backs [Pomolobus aesti'valis), menhaden (Bre'voortia), smelt (Osmerus), launce [Ammodytes), chub mackerel [Pneuma- 
tophorus), butterfish [Poronolus), cunners [Tautogolabrus), sculpins {Myoxocephalus), silver hake [Merluccius), tomcod 
{Microgadus), and hake {Urophycis). 

87. Cornish, Contr. Canad. Biol. (1902-1905), 1907: 81. 

88. At Prince Edward Island (Cornish, Contr. Canad. Biol. [1906— 1910], 1912: 79). 

89. No specimens of R. ocellata were taken in 44 hauls made by the El'CENE H. in 47-67 fathoms between Long. 
7i°54' and 7i°i5'W, January 27-February 3, 1950, and only one was taken deeper than 40 fathoms by the AL- 
BATROSS III in 63 hauls down to 240 fathoms between Long. 67°io' and 72°2o' W, May 11— 18, 1950. 



248 Memoir Sears Foundation for Marine Research 

range is probably near 18-19° C (about 64-66° F), for it tends to withdraw from very 
shallow water along southern New England in early summer when the temperature has 
risen to about that value and to reappear there and near New York in early autumn'" 
when the water has cooled to about that same temperature. At the opposite extreme it 
is taken frequently on the Nova Scotian Banks in water colder than 5° C (41° F)." The 
majority of the population inhabits salinities ranging from about 34.4 to 32 "/oo- Some 
individuals may even be exposed to salinities as low as 29 «/oo in the southern side of the 
Gulf of St. Lawrence, if any remain through the critical season (April-June) and in 
shoal enough water, around the shores of the Gulf of Maine, and possibly in the imme- 
diate vicinity of New York and at the mouth of Chesapeake Bay. One has even been 
reported from the Delaware River opposite Philadelphia. 

The stock inhabiting depths of 10 fathoms or more appears to be resident the 
year-round there. But it has been known for a long time that the coastwise fringe of 
population tends to work shoreward with the autumnal decrease in temperature in the 
southern part of its range and to withdraw again into deeper (i. e., somewhat cooler) 
water as bottom temperatures again approach the summer maximum. Thus all the dated 
records of its presence inshore between Chesapeake Bay and New Jersey have been be- 
tween December and early June.'^ Similarly, it appears in shallow water from Cape 
Cod to New York in the first part of October, and larger numbers are said to be taken 
in Massachusetts Bay in winter than in summer. But It is present inshore regularly 
during summer in Passamaquoddy Bay at the entrance to the Bay of Fundy, in Nova 
Scotian waters, and around Prince Edward Island in the south side of the Gulf of 
St. Lawrence (p. 249). 

It is now known that some of the population inhabiting the shore waters along 
southern New England carry out extensive coastwise journeys. Thus three tagged a 
few miles off Block Island, Rhode Island, in June if^^d^"^ were recaptured 80-120 
miles to the westward.** 

This Skate breeds throughout its geographic range.^^ In Nova Scotian waters it 
deposits its eggs from summer into autumn,'^ apparently at the same season in the Gulf 
of Maine; off southern New England it appears that eggs are deposited throughout 

90. Occasional specimens have been reported as summering in shoal water off the eastern end of Long Island, where 
the bottom temperature may rise to 18-19° C (about 64-66° F) at its seasonal maximum. And we have caught 
them in partially enclosed waters along outer Cape Cod in August. 

91. K. ocellata has been reported from the Newfoundland Banks in temperatures as low as -0.4 to -1.1° C (31-30° F). 
But the identity of the specimens has been questioned. 

92. Only a few of the published records for it along this sector have been dated. 

93. Four were recaptured out of a total of 400 fish tagged. We are indebted to Daniel Merriman for these records. 

94. One off Fire Island, New York, October 1946, one in New York Bay (Staten Island) in November 194S, and 
one in Sandy Hook Bay, New Jersey, in December 1946. The fourth recapture was in June 1947, near Watch 
Hill, Rhode Island, about 15 miles distant from the locality where tagged. 

95- Egg^ have been taken either from the mother or by dredge or have been found on the shore near Sable Island off 

Nova Scotia, in Hahfax Harbor, off the coast of Maine, at the tip of Cape Cod, in Vineyard Sound, southern coast 

of Massachusetts, off New York Harbor, and on the Delaware Coast. 
96. Eggs have been found in September in females trawled near Sable Island and caught off the coast of Maine; empty 

cases have been found in Halifax Harbor as early as the end of October. An egg case with advanced embryo, taken 

in Vineyard Sound late in August, 1936, is included in our Study Material. 



Fishes of the JVestern North Atlantic 249 

the year, for they have been found in ocellata collected in Block. Island Sound in April, 
May, August, November and February. A female, probably of this species, that was 
brought to Woods Hole laid an egg in the aquarium on January 16 and laid six more 
within a few days; the embryos were well advanced by May 12, but none had hatched 
by June 16, after which they were lost."' 

Numerical Abundance. R. ocellata has been characterized as common or as abun- 
dant about as often for one sector of its range as for another. But the only precise infor- 
mation regarding its numerical abundance with which we are acquainted is that one 
trawler, in 37 hauls, took 71 R. ocellata out of a total of 495 Skates of all kinds on 
Georges Bank in September 1929,"^ and that another trawler obtained an average 
hourly catch of 22.4 pounds with an 80-foot trawl in Long Island Sound from August 
1943 to January 1944,®" or about five specimens per hour, if the average weight were 
about five pounds. This would correspond to about 300 specimens per square mile if 
the bottom strip swept by the trawl were about 50 feet broad, if the distance covered 
were two sea miles per hour, and if the trawl caught every Skate that lay in its path, 
which it certainly did not. 

Range. Continental waters of the western North Atlantic from the offing of 
northern North Carolina to northern Nova Scotia, southern side of the Gulf of St. Law- 
rence, and Newfoundland Banks. 

Details of Occurrence. R. ocellata is generally distributed, if not very common, 
along the southern side of the Gulf of St. Lawrence from the Cape Breton shore to 
the Magdalen Islands, including Prince Edward Island, and it has been described as 
very common at Canso. It has been reported also by name for various localities along 
the periphery of the Banks of Newfoundland and has been identified from the southern 
part recently.'"" To the southward it has been taken at Banquereau Bank, Sable Island 
Bank, and Halifax, Nova Scotia, and fishermen are familiar with it throughout this 
general region to the mouth of the Gulf of Maine. It is well known in the Bay of Fundy, 
including the Passamaquoddy Bay region, and it has been taken at nearly all localities 
around the northern and western parts of the Gulf of Maine, in Massachusetts Bay, 
and off Cape Cod where information is available as to the composition of the local 
fish fauna. On Georges Bank,!"' as well as on Nantucket Shoals, it forms an appreciable 
proportion of the Skate population. It occurs regularly in season (p. 248) along the coast 
of southern New England, in the vicinity of New York (Sandy Hook and lower New 
York bays), along New Jersey,"* Delaware, and the Atlantic Coast of Virginia, and 
within the mouth of Chesapeake Bay; it also occurs on the offshore fishing grounds off 
New York and New Jersey, and it has been recorded in 1 6 fathoms off Bodie Island, North 
Carolina. But we find no report and have seen no specimen of it from farther south. 

97. This specimen was reported as R. lae'vh (Seal, Aquarium, 2 [lo], 1914: 105); but it seems likely that it was an 
ocellata because of its size (15 in. wide) and because of the season at which it deposited its eggs. 

98. Bigelow and Schroeder, Bull. U. S. Bur. Fish., 48, 1936: 324. 

99. Merriman and Warfel, Trans. 9th N. Amer. Wildl. Conf., 1944: 234. 
100. Information from Wilfred Templeman. 

loi. 14 "/o °f the Skates caught on one trip to Georges Bank in September 1929 were K. ocellata. 
102. Reported from many localities, including Delaware Bay. 



250 Memoir Sears Foundation for Marine Research 

Synonyms and References: 

Raja^"^ ocellata Mitchill, Trans. Lit. philos. Soc. N. Y., 7, 181 5: 477 (descr., New York); Storer, Boston J. 
nat. Hist., 2, 1839: 543 (descr., food, Mass.); Rep. Fish. Rept. Birds Mass., 1839: 191 (Mass.); Linsley, 
Amer. J. Sci., 4"^^ 1844: 77 (listed, Connecticut); Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 
510 (diagn., New York and Conn.); Gill, Proc. Acad. nat. Sci. Philad., Addend., 1861: 61 (listed, 
Atlant. Coast N. Amer.); Giinther, Cat. Fish. Brit. Mus., 8, 1870: 455 footnote (ref); Lyman, 6th 
Rep. Mass. Comm. inl. Fish., 1872: 53 (Mass.); Goode and Bean, Bull. Essex Inst., 11, 1879: 29 
(Maine); Bean, Proc. U. S. nat. Mus., J, 1880: 115 (Mass. locals.); Jordan and Gilbert, Bull. U. S. 
nat. Mus., 16, 1883: 40 (descr., size, egg cases); Bean, Rep. U. S. Comm. Fish (1882), 1884: 343 
(listed. Woods Hole, Mass.); Jordan, Rep. U. S. Comm. Fish. (1885), 1887: 799 (listed, N. Atlant. 
U. S.); Garman, Bull. Mus. comp. Zool. Harv., ly, 1888: gi, pi. 30 (mucous canal syst.); Jordan and 
Evermann, Rep. U. S. Comm. Fish. (1895), 1896: 221 (listed, Mass. northward); Bull. U. S. nat. Mus., 
47 (i), 1896: 68 (descr., egg cases, New York and northward); Bumpus, Science, N. S. 8, 1898: 58 
(Woods Hole); Smith, Bull. U. S. Fish Comm., ij, 1898: 89 (season. Woods Hole); Jordan and Ever- 
mann, Bull. U. S. nat. Mus., 47 (4), 1900: pi. 10, fig. 30 (ill.); Bean, 6th Rep. For. Comm. N. Y., 1901 : 
382 (season, New York); Linton, Bull. U. S. Fish Comm., 79, 1901 : 274, 431 (food, parasites, Vineyard 
Sound); Bean, Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 49 (descr., season, food. New York); Fowler, 
Rep. N. J. Mus. (1905), igo6: 71, pi. 3 (size, cf. R.erinacea, ill.. New Jersey); Tracy, 36th Rep. R. 
\. Comm. inl. Fish., 1906: 47 (season, Narragansett Bay;) Cornish, Contr. Canad. Biol. (1902-1905), 
1907: 82 (nos., food, Canso, Nova Scotia); Fowler, Rep. N. J. Mus. (1907), 1908: 129 (descr., nos., 
season. New Jersey, Delaware Bay); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 11 (New 
England locals.); Fowler, Proc. Acad. nat. Sci. Philad., 62, 1910: 470 (Maine, Mass., New Jersey); 
Linton, Bull. U. S. Bur. Fish., 28, 1910: 1201 (cestode parasite, Woods Hole); Tracy, 40th Rep. R. L 
Comm. inl. Fish., 1910: 62 (season, food, Narragansett Bay); Fowler, Proc. Acad. nat. Sci. Philad., 
6j, 191 1 : 5 (egg cases, Delaware); Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 4 (3), 191 3 : 102 (range); 
Cornish, Contr. Canad. Biol. (1906-1910), 1912: 79 (nos., Prince Edward L); Fowler, Proc. Acad, 
nat. Sci. Philad., 6^, 191 3: 62 (listed, Virginia); Halkett, Check List Fish. Canad., 191 3: 42 (listed. 
Maritime Provinces to New York); Nichols, Abstr. Linn. Soc. N. Y., 20-23, '91 3- 9i (season, near 
New York); Sumner, Osburn and Cole, Bull. U. S. Bur. Fish., 31 (2), 1913: 738 (season, depth, food, 
parasites. Woods Hole); Fowler, Proc. Acad. nat. Sci. Philad., 66, 19 14: 353 (nos., Delaware); Wieg- 
man and Nichols, Copeia, 3, 19 14: 2 (season, female with egg, New York); Fowler, Copeia, 13, 19 14: 
2 (New Jersey); Proc. Acad. nat. Sci. Philad., 6"], 1915: 517 (Nova Scotia); Wiegman and Nichols, 
Copeia, 23, 1915: 43 (New Jersey); Fowler, Copeia, 27, 1916: 12 (New Jersey); Copeia, 31, 1916: 
41 (New Jersey); Proc. Boston Soc. nat. Hist., J5, 1917: 11 1 (color, thorn pattern, Maine and Mass.); 
Jordan, Copeia, 49, 1917: 87 {diaphanes replaces oceUatd)\ Huntsman, Trans, roy. Soc. Canad., (3) 
12 (4-5), 1918: 63 (temp.. Gulf St. Lawrence); Fowler, Proc. biol. Soc. Wash., J2, 1919: 72 (Dela- 
ware R.); Proc. Acad. nat. Sci. Philad., JI, 1920: 293 (New Jersey); Proc. biol. Soc. Wash., JJ, 1920: 
145 (New Jersey); Cox, Contr. Canad. Biol. (1918-1920), 1921: 109 (S. side Gulf St. Lawrence); 
Fowler, Fish Culturist, 6, 1927: 115 (listed. New Jersey); Dill, Edwards and Florkin, Biol. Bull. Woods 
Hole, 62, 1932: 23 (blood);!""* Fowler, Proc. Acad. nat. Sci. Philad.. 8g, ^^37'- 304 (season, New Jersey). 
Raia^"^ diaphanes DeKay, Zool. N. Y., 4, 1842: 366, pi. 67, fig. 218 (descr., ill.. New York); Storer, Mem. 
Amer. Acad. Arts Sci., N. S. 9, 1867: 240, pi. 39, fig. i; also Fishes Mass., 1867: 264, pi. 39, fig. i 
(descr., ill., size, nos., Mass.); Garman, Mem. Harv. Mus. comp. Zool., j6, 191 3: 339, pi- 22, fig. i 
(descr., ill., cf R. erinacea); Huntsman, Contr. Canad. Biol. (1921), 1922: 57 (depth, season, Passama- 
quoddy Bay and Bay of Fundy); Breder, Copeia, 127, 1924: 28 (season, temp., near New York); Hunts- 
man and Sparks, Contr. Canad. Biol., N. S. 2, 1924: 102 (lethal temp.); Linton, Proc. U. S. nat. Mus., 
64 (21), 1924: 8, 61, 65 (cestode parasites); Bigelow and Welsh, Bull. U. S. Bur. Fish., 40 (2), 1925: 
60 (descr., ill., habits, Gulf of Maine); Manter, Illinois biol. Monogr., 10 (2), 1926: 8 (parasites, Maine); 
Breder and Nichols, Zoologica N. Y., 9, 1927: 23 (ill., season, food, near New York and south. New 
England); Hildebrand and Schroeder, Bull. U. S. Bur. Fish., 42 (i), 1928: 56 (descr., ill., season, Chesa- 
peake Bay); Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 30 (descr., egg cases, depth, season, Vir- 
ginia to Gulf of St. Lawrence); Jordan, Manual Vert. Anim. NE U. S., 1929: 16 (diagn., New Jersey 
north); Jordan, Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 25 (listed, E. coast 

103. Sometimes spelled Kaia. 104. Spelled ocillata. 105. Also spelled Raja. 



Fishes of the JV ester ft North Atlantic 251 

U. S.); Babkin and Komarov, Contr. Canad. Biol., 7 (1-7), 1931 : 13 (gastric juice, St. Andrews, New 
Brunswick); Huntsman, Contr. Canad. Biol., 7 (1-7), 1931 : 4 (physiol., St. Andrews, New Brunswick); 
MacKay, Contr. Canad. Biol., 7 (1-7), 1931 : 19 (physiol.); Smith, H. W., Amer. J. Physiol., ()8, 1931 : 
279—295 (excretion); Breder, Copeia, 1932: 31 (season, nos.. Block Island); Anon., Rep. Newfound- 
land Fish. Res. Lab. (1931), 1932: 107 (listed, SW. side of Grand Bank); Dawson, Biol. Bull. Woods 
Hole, 64, 1933"- 33 (blood); Nichols, Contr. Canad. Biol., 7, 1933: 34, 449 (physiol.); Anon., Rep. 
Newfoundland Fish. Res. Lab. (1932), 1933: 125 (depth, temp., sal., Newfoundland and Nova Scotian 
Banks); Bigclow and Schroeder, Canad. Atlant. Fauna, I2e, 1934: 21, 22 (descr., ill., range); Anon., 
Rep. Newfoundland Fish. Res. Lab. (1934), 1935: 77 (Banquereau, off Nova Scotia); Vladykov and 
McKenzie, Proc. N. S. Inst. Sci., 79, 1935: 51 (Nova Scotia); Bigelow and Schroeder, Bull. U. S. Bur. 
Fish., 48, 1936: 324 (cf. R. erinacea, nos., Georges Bank); Smith, H. W., Biol. Rev. (Cambridge), J7, 
1936: 67 (blood serum); Vladykov, Nat. canad., 6j, 1936: 217 (descr., photos, season, egg cases. Nova 
Scotia, cf. other species); Nichols and Breder, Copeia, 1937: 182 (egg cases, cf. other species); Breder, 
Bull. N. Y. zool. Soc, 41, 1938: 28 (listed. New York Harbor); Merriman and Warfel, Trans. 9th 
N. Amer. Wildl. Conf, 1944: 234 (nos.. Long Island Sound); Bull. Bingham oceanogr. Coll., 11 (4), 
1948: 134 (listed, off Rhode Island and Conn.). 

Raja ocellata (in part) Dumeril, Hist. Nat. Poiss., J, 1865: 539 (descr., but R. eglanteria perhaps included). 

Rata diaphanes (in part) Lyman, 6th Rep. Mass. Comm. inl. Fish., 1872: 54 {R. ocellata probably included, 
because abundant April and May, south. Mass.). 

Raja ocellata var. ocellata Garman, Proc. Boston Soc. nat. Hist., 77, 1874: 177 (descr., size, Mass.). 

Raja ocellata var. diaphana Garman, Proc. Boston Soc. nat. Hist., ly, 1874: 177 (color variety). 

Big Skate, Wilson, Bull. U. S. nat. Mus., 158, 1932: 403 (copepod parasite). 

Raia scabrata Garman, Mem. Harv. Mus. comp. Zool., j6, 1913: pi. 21, fig. I (ill., mislabelled).^"^ 

Probable Sj'nonyms : 

Raia batis Wyman, Mem. Amer. Acad. Arts Sci., N. S. 9, 1864: 39, i pi. (descr., ill., egg cases, embryos and 
newly hatched); Ann. Mag. nat. Hist., (3) 14, 1864: 399 (abstr. of Wyman, 1864); Amer. J. Sci., (2) 
j5, 1864: 29 (abstr. of Wyman, 1864); Proc. Boston Soc. nat. Hist., 9, 1865: 334 (abstr. of Wyman, 
1864); Putnam, Amer. Nat., J, 1869: 621 (after Wyman, 1864). 

Raja laevis Seal, Aquarium, N. Y., 2 (10), 1914: 105 (female with eggs. Woods Hole, winter, probably R. ocel- 
lata because of small size, see p. 249, footnote 97; not seen). 

Doubtful References: 

Raia diaphanes Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1846: 510 (doubtful whether R. ocellata or R. 

eglanteria). 
Not Raja batis Linnaeus, Syst. Nat., z, 1758: 231 (E. Atlant.). 
Not Raja ocellata, capite magna . . . Russell, Fish. Coromandel, r, 1 803 : 5, pi. 8 (a Sting Ray of the genus Jeto- 

batus). 
Not Raja diaphanes Mitchill, Trans. Lit. philos. Soc. N. Y., j, 181 5: 478 (equals R. eglanteria Bosc 1802; 

see p. 187); Ayers, Boston J. nat. Hist., 4, 1843: 289, pi. 12, fig. 2 (season, near New York, ill. appears 

to be R. eglanteria, see p. 176). 
Not Raia batis Storer, Rep. Fish. Rept. Birds Mass., 1839: 193 (equals R. laevis Mitchill 1817). 
Not Raia ocellata DeKay, Zool. N. Y., 4, 1842: 369, pi. 65, fig. 212 (descr., ill., equals R. eglanteria Bosc 

1802, see p. 176). 

Raja olseni Bigelow and Schroeder 1951 

Figure 54 a 

Study Material. Immature male, 280 mm long, type, from the Gulf of Mexico, 
Lat. 27°25' N, Long. 96° 13' W, 76 fathoms, Oregon St. 157, in U. S. National Mu- 

106. This illustration was actually of a specimen of R. ocellata in the collection of the Harvard Museum of Comparative 
Zoology which we have examined. 



252 Memoir Sears Foundation for Marine Research 

seum; an immature male, 282 mm long, from Lat. 27°27' N, Long. 96°i7' W, 65 
fathoms, Oregon St. 158; 2 males and 2 females, 230— 310mm long, Lat. 30°02'N, 
Long. 86°53' W, 60 fathoms, Oregon St. 332, and two males each of 310 mm, from 
Lat. 30°02'N, Long. 86°55' W, 64 fathoms, Oregon St. '},(^'t^. 

Distinctive Characters. Raja olseni closely resembles R. laevis in general appearance 
and in lacking thorns along the midbelt of the disc from the level of the axils of the 
pectorals to the vicinity of the spiracles. But it differs from laevis in having an inter- 
space between the dorsal fins nearly or quite as long as the base of the first dorsal (only 
0.1—0.3 that long in laevis) and by having a fringe on the expanded outer margin of 
the nostril (smooth in laevis); also, the lateral folds along its tail extend only to the 
anterior third of the caudal fin, whereas in laevis they extend almost or quite to the 
extreme tip of the tail; the lower sides of its disc are prickly along the anterior edges 
and over the rostral cartilage (smooth on laevis of equal size), and its mucous pores 
are not marked with black as they are in laevis. It differs from R. spinicauda in its 
fringed nostril lobe, in its shorter tailfolds, in the interspace between its dorsals, and 
in having three rows of thorns on the tail (only one row on spinicauda). 

Description. Proportional dimensions in per cent of total length. Male, 280 mm 
(U. S. Nat. Mus., type, No. 153556), and male, 282 mm (Harv. Mus. Comp. ZooL, 
No. 37176), from the northwestern part of the Gulf of Mexico. 

Disc: extreme breadth 69.0, 72.0; length 54.0, 58.2. 

Snout length: in front of orbits 15.6, 17.4; in front of mouth 17.7, i9-5. 

Orbits: horizontal diameter 4.4, 4.3; distance between 3.6, 4.3. 

Spiracles: length 2.0, 2.3; distance between 6.3, 6.7. 

Mouth: breadth 7.9, 8.2. 

Nostrils: distance between inner ends 7.9, 8.3. 

Gill openings: lengths, ist 2.0, 2.0; 3rd 2.1, 2.1 ; 5th 1.6, 1.6; distance between 
inner ends, ist 13.6, 14.2; 5th 7.3, 8.2. 

First dorsal fin: vertical height 2.5, 2.5; length of base 4.3, 4.4. 

Second dorsal fin: vertical height 2.4, 2.3; length of base 4.6, 4.4. 

P^/w'c5; anterior margin 13.2, 13.5. 

Distance: from tip of snout to center of cloaca 47.5, 49.3 ; from center of cloaca 
to 1st dorsal 30.4, 30.2; to tip of tail 52.5, 50.7; from rear end of 2nd 
dorsal base to tip of tail 9.0, 7.8. 

Interspace between: ist and 2nd dorsals 4.3, 3.9. 

Disc about 1.3 times as broad as long, the maximum anterior angle in front of 
spiracles about 90° ; anterior margins concave just posterior to tip of snout, weakly 
convex opposite eyes and spiracles, thence about straight rearward; outer corners nar- 
rowly rounded; posterior corners more broadly so; posterior margins gently convex. 
Axis of greatest breadth about 67 °/o of distance rearward from tip of snout towards 
axils of pectorals. Tail with a lateral fold low down on each side beginning posterior 
to axils of pelvics by a distance about */& ^s long as eye and ending opposite anterior 



Fishes of the Western North Atlantic 



253 




Figure 54a. Raja oheni, male, 280 mm long, from Gulf of Mexico, Lat. 2'j°zc,' N, Long. 96°! 3' W (U. S. 
Nat. Mus. No. 153556, type). J Posterior part of tail, about 1.5 x. 5 Mouth and nostrils, about 2X. C Upper 
teeth, about 1 2 x . 



third of caudal fin; length of tail from center of cloaca to origin of first dorsal fin 0.61 — 
0.67 as great as, to its tip i.o— i.i times as great as, distance from center of cloaca to 
tip of snout. 



2 54 Memoir Sears Foundation for Marine Research 

One to three small thorns immediately in front of orbit, one being on the inner 
and one or two on the outer margin, also one or two on inner rear margin of orbit, and 
sometimes one or two opposite middle of orbit; these are the only thorns or prickles 
on the disc; i 3 to 16 thorns along midline of tail from a little in advance of axils of 
pelvics to first dorsal fin; three in interspace between first and second dorsals; also an 
additional row of thorns, widely and unevenly spaced, on each side of median row, 
beginning about opposite tip of pelvic and extending to opposite beginning of caudal 
fin; dorsals and caudal smooth. Lower surface with a narrow band of small prickles 
along anterior margin of disc from level of nostrils to tip of snout and also along rostral 
cartilage. 

Snout in front of orbits 3.5-4.0 times as long as orbit, its length in front of mouth 
2.2-2.3 tinges as great as distance between exposed nostrils. Orbit 1.9-2.2 times as 
long as spiracle; distance between orbits i. 0-1.2 times as great as length of orbit. 
Distance between first gill openings 1.7 times as great as distance between exposed 
nostrils, between fifth gill openings i.o-i.i times; first gill openings 1.25 times as 
long as fifth and about 0.25 as long as breadth of mouth. Nasal curtain fringed; ex- 
panded posterior (outer) margins of nostrils fringed. Upper and lower jaws moderately 
arched centrally. 

Teeth |^40) close-set in quincunx, ovate, with a triangular cusp. 

First and second dorsals similar in size and shape. Interspace between dorsals 
0.9-1.0 as long as base of first dorsal. Caudal membrane from rear end of base of 
second dorsal about twice as long as base of first dorsal. Pelvics deeply concave, strongly 
scalloped along anterior side of excavation but only weakly so rearward; anterior margin 
about as long as distance from its own origin to rear tip of pelvic; anterior lobe moder- 
ately slender, including five radial cartilages besides the first stout one; posterior lobe 
moderately convex outwardly; rear tips abruptly rounded, extending about 2/7 of the 
distance from axils of pelvics toward first dorsal; inner margin straight. Claspers falling 
well short of tips of pelvics on specimens seen. 

Rostral cartilage firm, extending to tip of snout. Anterior pectoral rays reaching 
about halfway from level of front of orbits toward tip of snout. 

Color. In life, upper surface dark olive brown with many small roundish obscure 
spots of darker brown on disc; a small dark spot, smaller than pupil, on each side 
of disc inward near its center; series of small whitish pores extending in 3 or 4 rows 
along midzone of back from region of pectoral girdle to axils of pelvics and onto tail 
in I or 2 rows; 2 rows extending rearward and outward on each side of disc posterior 
to scapular region; whitish pores also present opposite and in front of orbits, some 
extending toward outer margin of disc in sinuous rows. Below, jet black everywhere, 
but this pigment tending to diminish in intensity or disappear in preservative. 

Si-ze and Habits. Since all the males have very small claspers, it is probable that 
the species attains at least a moderately large size, possibly two feet or more in total 
length. Nothing is known of its habits except that it is apparently a moderately deep- 
water species. 



Fishes of tlie Western North. Atlantic 255 

Occurrence. It is known only from the northern and northwestern part of the Gulf 
of Mexico, as listed in Study Material. 

Reference: 

Raja o/seni Bigelow and Sclirocder, J. Wash. Acad. Sci., 41 (12), 195 1 : 386 (descr., ill., Gulf of Mexico). 

Raja radiata Donovan 1807 

Thorny Skate, Starry Skate 

Figures c,K,, 56 

Study Material. Fifty-seven embryos and adults of both sexes, 126 to 935 mm 
long, from: Norway; Egemonde and Tunugdliarfik Fjord, West Greenland; Grand 
Banks of Newfoundland; Halifax, Nova Scotia; Emerald Bank, Georges Bank and 
various localities within the Gulf of Maine, including the Bay of Fundy; off the con- 
tinental slope from southwest of Georges Bank to New Jersey in 118-260 fathoms 
and off South Carolina in 74 fathoms; in the U. S. National Museum and Harvard 
Museum of Comparative Zoology. 

Distinctive Characters. R. radiata is marked off from all other Skates of the North 
Atlantic by the large size of the thorns along the midline of the back from the nape 
to the first dorsal fin, plus the facts that not more than 10 large thorns are present 
on the tail posterior to the axils of the pelvics and that the anterior contour of the disc 
is very obtuse (anterior angle at least 110°). It is so closely allied to R. doello-juradoi 
of the Argentinian-Patagonian-Falklands region in the South Atlantic that no reliable 
criteria have been found to distinguish the one from the other. 

Description. Proportional dimensions in per cent of total length. Male, 530 mm 
(Harv. Mus. Comp. Zool., No. 249). Female, 800 mm (Harv. Mus. Comp. Zool., 
No. 1 1 39), from Nahant, Massachusetts. 

Disc: extreme breadth 71.7, 74-5; length 52.8, 51.3. 

Snout length: in front of orbits 13.0, 12.9; in front of mouth 14.0, 13.9. 

Orbits: horizontal diameter 4.3, 3.8; distance between 5.1, 4.8. 

Spiracles: length 3.0, 3.3; distance between 8.1, 8.0. 

Mouth: breadth 10. i, 10.7. 

Nostrils: distance between inner ends 9.5, 9.8. 

Gill openings :\en^th?,, ist 1.9, 1.9; 3rd 2.1, 2.0; 5th 1.7, 1.9; distance between 
inner ends, ist 17.0, 19.6; 5th 10.7, 12.2. 

First dorsal fin: vertical height 3.0, 3.2; length of base 5.2, 5.1. 

Second dorsal fin: vertical height 2.8, 2.8; length of base 5.9, 6.6. 

Pelvics: anterior margin 13.2, 11.2. 

Distance: from tip of snout to center of cloaca 49.4, 49.2 ; from center of cloaca 
to 1st dorsal 36.5, 35.1; to tip of tail 50.6, 50.8; from rear end of 2nd 
dorsal base to tip of tail 3.0, 2.8. 

Interspace between: ist and 2nd dorsals 0.5, 1.3. 



256 Memoir Sears Foundation Jhr Marine Research 

A 




Figure 55. Raja rad'iata. A Female, 800 mm long, from Massachusetts Bay (Harv. Mus. Comp. Zool., No. 1 1 39). 
B Pelvics of same. C Nostril of female, 576 mm long, from Nahant, Massachusetts (Harv. Mus. Comp. Zool., 
No. 254), about 2.3 X. Ti Outer posterior margin of right-hand nostril (above) and of left-hand nostril (below) 
of male 378 mm long, from Emerald Bank, Nova Scotia, 55 fathoms (Harv. Mus. Comp. Zool., No. 33920), 
to show variation, about 2.3 x. E Juvenile female, 127 mm long (Harv. Mus. Comp. Zool., No. 34755). 



Disc about 1.3— 1.4 times as broad as long; maximum anterior angle in front of 
spiracles about 110—140°, small specimens averaging more obtuse than large; an- 



Fishes of the Western North Atlantic 257 

terior margins slightly concave close behind tip of snout, weakly and evenly convex 
thence rearward to outer corner of pectorals on newly-hatched specimens but becom- 
ing weakly concave with growth at level of eyes and spiracles and about equally so in 
both sexes to maturity ji"' posterior corners much more broadly rounded than outer 
corners. Axis of greatest breadth about 70 % of distance back from snout toward axils 
of pectorals. Tail with narrow lateral folds low down and beginning close behind axils 
of pelvics; its length from center of cloaca to origin of first dorsal about 80 "/o as great 
as distance from center of cloaca to snout in young, 60-70 "/o at maturity; extreme 
length of tail from center of cloaca about i.i — 1.2 times as great as distance from cloaca 
to snout in young, but about 0.9— i.i times in half-grown specimens and adults. 

Upper surface with a median row of i i — 19 large conspicuous thorns on radiate 
bases from nuchal region back nearly to first dorsal fin, usually 1-2 more on adults 
than on young specimens ;!"* 6-9 of these anterior to axils of pectorals and about an equal 
number along tail, the thorn nearest to the first dorsal sometimes very small ; midbelt 
of back with 1—3 irregular rows of smaller thorns on either side of the midrow, these 
confined to disc on very small specimens but extending along tail to first dorsal fin 
on adults, decreasing in size rearward; anterior and median parts of pectorals with 
scattered thorns, most closely spaced from level of spiracles forward; 2-6 near tip of 
snout, sometimes one pair conspicuously largest; one prominent thorn close in front of 
eye, one on inner posterior margin of orbit, and one close to inner end of spiracle; 
also 2—3 on each shoulder, the most posterior of these usually largest; skin of disc 
usually without prickles around eyes, behind spiracles, behind shoulders, and on outer 
posterior parts of pectorals. Skin also without prickles elsewhere between the larger 
thorns on some specimens but more or less prickly on others, especially on the snout, 
between the orbits and on the shoulders; skin above eyes either naked or prickly; 
tail with a thick-set belt of very small prickles low down along either side from axils 
of pectorals to below dorsals, these prickles being especially conspicuous on small 
specimens; dorsals prickly; space between dorsals with one small thorn, or none, if the 
fins are separated; caudal membrane sometimes with a few minute prickles, but usually 
smooth; pelvics sometimes naked, but more often with central area more or less prickly, 
sometimes with 1—2 thorns. Sexually maturing females become increasingly thorny 
over the midzone of the disc in general, on the shoulders, and over the outer anterior 
parts of the pectorals ; they also become increasingly prickly between the thorns. But 
maturing males lose most of the thorns from the pectorals except along the anterior 
margins. Alar spines of mature males in 'i^—^ rows, each row about as long as distance 
between outer margins of orbits. Lower surface smooth on small specimens, but prickles 
usually develop with growth on the midzone of the snout about halfway back toward 
the mouth, along the extreme outer margins of the disc to the level of the spiracles in 

107. We have a male 540 mm long with well developed claspers, caught oft" South Carolina, on which the anterior margin 
of disc is sharply indented opposite level of spiracles. 

loS. From the Gulf of Maine, three out of five small specimens, 126-153 mm long, had 12 mid-dorsal thorns, one had 
13, and one had 14; among 31 larger specimens, one had 11, two had 12, eleven had 13, nine had 14, four had 15, 
and four had 16; West Greenland specimens had 13-19 (Jensen, Spol. Zool. Mus. Hauniensis, 9, 1948: 28). 



258 Memoir Sears Foundation for Marine Research 




Figure 56. Raja radiata. A Mature male, 530 mm long, from Massachusetts Bay (Harv. Mus. Comp. ZooL, 
No. 249). B Mouth and nostril of female, 576 mm long, from Nahant, Massachusetts (Harv. Mus. Comp. Zool., 
No. 254). C Margin of nasal curtain of same, about 2.3 X . i) Upper teeth of female, from Grand Manan Island, 
New Brunswick (Harv. Mus. Comp. Zool., No. 1 3 2), about 4.6 X • £ Upper teeth of mature male, from Ivigtut 
Fjord, West Greenland (Harv. Mus. Comp. ZooL, No. 36557), about 4.6X. 



Fishes of the Western North Atlantic 259 

females by maturity, and over the lower surface of the snout as a whole anterior to the 
nostril in mature males. ^'''' 

Snout in front of orbits about 2.4-2.5 times as long as distance between orbits 
in adult males, a little longer relatively in adult females; its length in front of mouth 
about 1.3— 1.7 times as long as distance between nostrils. Orbit about 75-100 "/q as 
long as distance between orbits in small specimens, relatively a little shorter in large, 
and about 1.2— 1.4 times as long as spiracle. Distance between first gill openings 1.8 — 
2.1 times as long as distance between exposed nostrils, 1.4— 1.7 times between fifth 
gill openings; first gill openings i. 0—1.3 times as long as fifth and about 15-20 "/o 
as long as breadth of mouth. Nasal curtain fringed, the expanded posterior (outer) 
margin of nostril either smooth or coarsely or finely fringed, not always alike on the 
two sides. Mouth only a little more arched in adult males than in females and young. 

Teeth 35^46) with round bases and low cusps, worn nearly smooth in oldest 
rows but conical in youngest, close-set in quincunx in young specimens but teeth more 
loosely spaced and the series more nearly transverse in older ones; those of adult males 
only slightly sharper and spaced only a little more widely than in females. 

First and second dorsals similar in size and shape, either confluent at base or 
separated by an interspace up to nearly half as long as base of first dorsal. Caudal mem- 
brane posterior to base of second dorsal about 1/4 as long as base of first dorsal sub- 
sequent to loss of filamentous embryonic prolongation. Pelvics moderately concave 
and scalloped from base of excavation rearward; anterior margin about 55—72 "/q as 
long as distance from its own origin to rear tip of pelvic in large specimens, about 
78-86 "/o in small of both sexes; anterior lobe broad, including about five radials; 
outer margin of posterior lobe moderately convex, rear corner narrowly rounded. 
Claspers of sexually mature male conspicuously robust, extending rearward about ^/g 
the distance from axils of pelvics toward first dorsal. 

Anterior rays of pectorals extending about ^Is—^U of the distance forward from 
level of front of orbits toward tip of snout. 

Color. Upper surface brown, either uniform or partially clouded or spotted with 
darker; small specimens often spotted more definitely than older ones; sometimes a 
white spot beside each eye, one on each side opposite the nuchal region, and another 
on each side on the posterior part of the disc. Lower surface white, sometimes with 
irregular sooty or brownish blotches. 

Remarks. The North American form, R. scabrata, has been separated from R. radiata 
of the eastern Atlantic because it is "larger, less rough, and rather more angular" and 
because the small prickles are more sparsely scattered or absent from considerable 
areas. 1^" But our own examination of the series listed under Study Material (p. 255) 
has failed to show any consistent differences in any of these respects between those 
from American, West Greenland, or Norwegian waters. "^ 

109. These areas are smooth on a mature male 540 mm long from South Carolina, 
no. Garman, Mem. Harv. Mus. comp. Zool., j6, 191 3: 340. 

III. Some specimens from Norway that we have seen are densely prickly over the upper surface as a whole, except 
for the anterior lobes of the pelvics; but another specimen from the same region has only a few scattered prickles, 

17* 



2 6o Memoir Sears Foundation for Marine Research 

Size. In American waters, R. radiata is hatched at a length of about loo mm from 
snout to origin of first dorsal fin. But those of northern European seas, where the egg 
cases average somewhat smaller (see p. 260), may hatch at a correspondingly smaller 
size.i^^ It may mature through a wide range of sizes. Thus, a male only 21V4 inches 
long (540 mm) was fully mature, whereas another about 33^/2 inches (845 mm), both 
seen by us, has the claspers extending only slightly beyond the tips of the pelvics. On 
another of about the same size as the latter (830 mm, or about 33 in.), the claspers 
reach nearly halfway along the tail and appear to be fully developed; they are surely 
so on another of 935 mm. Females as long as 1,020 mm (Nova Scotia) and as short 
as 450 mm (North Sea and Norway) have been reported as containing eggs. The 
Icelandic and American stocks tend to grow larger than those of northern Europe or 
West Greenland. Thus, the maximum recorded length is only 450 mm for the North 
Sea, about 600 mm for Norwegian waters,^" and 590 mm for West Greenland;"* but 
the maximum size is 990 mm for Iceland, ^^^ 1,020 mm for the Scotian Banks,"* about 
895 mm (35 V4 in-) for Georges Bank,"' 800 mm for Massachusetts Bay, and 935 mm 
for the offing of New Jersey. 

Developmental Stages. The egg cases, flat on one side but strongly convex on the 
other, are rough with narrow longitudinal ridges. A membrane-like mass of delicate 
fibrils, matted together, extends along each lateral margin (over the surface as a whole 
at deposition); each horn terminates in a slender fibril. i" The cases vary widely in size, 
probably corresponding to the size of the parent fish. In the North Sea they average 
about 48 by 34 mm (exclusive of horns), in Trondhjem Fjord 50-65 by 40—48 mm, 
off West Greenland"' 45—68 by 26—51 mm, on the outer Nova Scotian Banks 77— 
90 mm long from mothers 840—1,020 mm long, and in the estuary of the St. Lawrence 
River (size of parents not recorded) 66—68 mm long.^-" The single egg case recorded 
from Georges Bank, taken from a fish about 8 10 mm long (32 in.), measured 76 by 
57 mm (3 by 27^ in.). 

At hatching the young already resemble their parents so closely (apart from 
whatever vestiges of the embryonic caudal filament may persist) that their identification 
presents no difficulty. This is one of the Skates for which monstrosities, such as those 

as is often true of the Gulf of Maine specimens. Some from Southwest Greenland and from the Newfoundland 
Banks are more or less prickly above the eyes and have a few prickles scattered among the thorns, whereas others 
are smooth above the eyes and between the thorns, as is commonly the case on New England specimens. However, 
it may develop that there is some geographic variation in the average number of mid-dorsal thorns, for the maximum 
number seen on any Gulf of Maine specimen has been i6, whereas there are i6 each on two out of six from West 
Greenland and on two out of three from Norway. 

112. An embryo 84 mm long to origin of first dorsal (120 mm long to tip of tail, but with the latter still filamentous) 
bears a yolk sac of considerable size, but the thorns have not yet been formed though their pattern is already evident. 
The smallest free-living specimen we have seen, a newly hatched one from Georges Bank which had already lost 
its caudal filament, was 100 mm long to first dorsal and 126 mm in total length. 

113. Smitt, Hist. Scand. Fish., 2nd ed., 1895: 1108. 114. In our Study Material. 

115. Saemundsson, Skr. Komm. Havunders. Kbh., 5, 1909: 125. 116. Vladykov, Nat. canad., 63, 1936: 225. 

117. Bigelow and Schroeder, Bull. U.S. Bur. Fish., 48, 1936: 325. 

118. For description of egg cases and of early stages in embryonic development, see Nordgaard (K. norske vidensk. 
Selsk. Skr. [1915], Heft 2 [9], 1916: 24; and Vladykov, Nat. canad., 63, 1936: 223). 

119. Jensen, Spol. Zool. Mus. Hauniensis, 9, 1948: 30. 120. Vladykov, Nat. canad., 63, 1936: 225. 



Fishes of the IVestern North Atlantic 261 

described on p. 143, have been reported, with the anterior parts of the pectorals separate 
from the sides of the head.^^^ 

Habits. Regarding the diet of R. radiata off the American Coast, we only know 
that several specimens caught by us on Georges Bank in September had been feeding 
on shrimps, spider crabs, anemones, hydroids, worms, and fishes of undetermined 
species. In Icelandic waters R. radiata is reported as feeding on small Crustacea, especially 
gammarids and spider crabs {Hyas), and on small fishes {Ammodytes) \ off Norway they 
feed on Crustacea, fishes, and worms,!^^ and off West Greenland on Crustacea. 

In the western side of the Atlantic R. radiata evidently produces young through- 
out its geographic range, for females containing well formed eggs, or egg cases con- 
taining embryos, have been taken all along the coast of West Greenland from Disko 
Bay (Lat. 69°26' N) southward, on the southwest slope of the Newfoundland Banks, 
off the outer coast of Nova Scotia, and on Georges Bank fronting the Gulf of Maine. 
Also, it appears that this species may deposit its eggs at any time of year in those parts 
of its range where winter chilling is not too severe, for gravid females, or deposited eggs 
in various stages of incubation, have been found in January and February in Trond- 
hjem Fjord, Norway; from February to June, and again in October, in the North Sea 
off Aberdeen, Scotland; from June to August around Iceland ;i^^ in the summer off 
West Greenland; in July on the southwest slope of the Grand Banks of Newfoundland 
(see p. 263); in April, June, July, and September off Nova Scotia; and in September 
on Georges Bank. The period of incubation is not known. 

In general R. radiata is restricted to depths greater than about 10 fathoms even 
in the northern part of its range. In the Gulf of St. Lawrence, for example, it is de- 
scribed as having been caught "at considerable depths" only.^^* Recorded captures 
from the Newfoundland and Scotian Banks have been 20 fathoms and deeper; in the Bay 
of Fundy-Passamaquoddy region it is taken only in water deeper than 10 fathoms; of 
the many specimens taken by us in the Gulf of Maine, the shoalestwas at 14 fathoms. i^s 
At the other extreme of its depth range, it is taken regularly down to 100 fathoms 
throughout its latitudinal range (many as deep as 120 fath.), while it has been trawled 
between 160 and 336 fathoms at numerous localities along the upper part of the con- 
tinental slope in the offing of southern New England; down to 218 fathoms south of 
Newfoundland; to 250 fathoms in Trondhjem Fjord; to 330 fathoms off West Green- 
land; and to 459 fathoms near Spitzbergen. There is nothing in the available record 
to suggest that this species carries out any regular seasonal migrations in American 
waters, whether inshore and offshore or north and south. 

The Starry Skate is more catholic in respect to its choice of bottom habitat than 
are some other Skates, for while it is most plentiful on good fishing grounds of sand 

121. Reported for a specimen off Iceland by Saemundsson (Skr. Komm. Havunders. Kbh., 5, 1909: 126). 

122. Saemundsson, Skr. Komm. Havunders. Kbh., 5, 1909: 126; Smitt, Hist. Scand. Fish., 2nd ed., 1895: iiio. 

123. Saemundsson, Skr. Komm. Havunders. Kbh., 5, 1909: 126. 

124. Cox, Contr. Canad. Biol. (1918-1920), 1921: 109. 

125. The report of it being taken in fish traps at Woods Hole (Sumner, Osburn and Cole, Bull. U. S. Bur. Fish., 31, 
1913: 738) lacks supporting evidence as to identity. 



262 Memoir Sears Foundation for Marine Research 

and broken shells, sand and gravel, or gravel and pebbles, we have also taken it at many- 
stations in the Gulf of Maine where soft mud floors the deeper troughs. It does not occur 
regularly at any locality where the bottom temperature rises above 9-10° C (48-50° F) 
for more than short intervals, though it can survive brief exposure to temperatures 
much higher than those to which it is subjected in any part of its range.i^s At the other 
extreme, many have been taken in Newfoundland waters where the temperatures go 
down to 1.1° C (34° F), or nearly as low as temperatures are in the open sea off the 
eastern American seaboard. Nor does it show any apparent preference for either the 
upper half of its thermal range or for the lower. However, the polar temperature of 
the ice-chilled Labrador Current may be responsible for its scarcity along the north- 
eastern coast of Newfoundland north of the offing of Conception Bay and along Atlantic 
Labrador as contrasted with its abundance along southwestern Greenland, where the 
temperature of the bottom water at depths where it occurs are upwards of 2-3° C 
(36-37° F) for the most part during late summer and early autumn. 1-' 

R. radiata is seldom (if ever) encountered in water less saline than 3 1.2-3 1.5 °/oo 
(shoaler parts of Gulf of St. Lawrence ; also Passamaquoddy Bay) ; the recorded max- 
imum for it is about 35.3 "/oo, and the great majority of the American population 
throughout its latitudinal range exists in salinities ranging from about 32-34.5 "/oo- 

Numerical Abundance. In September 1929, 37 trawl hauls on Georges Bank yielded 
a catch of 325 R. radiata, and trawling on the Newfoundland Banks yielded a maximum 
catch of 54 per hour. As further evidence of its abundance we may quote catches of 
12 R. radiata in 30 minutes in the western side of the Gulf of Maine with a beam 
trawl that was only eight feet across at the mouth, and of i-ioo specimens in 26 trawl 
hauls between Mt. Desert Island and Massachusetts Bay.^^s Oflf the coast of southern 
New England, 21 were taken in 17 half-hour trawl hauls in 71-260 fathoms, but 
none were taken in 46 hauls made in 22-69 fathoms on the same cruise. ^^^ 

Relation to Man. No commercial use is made of this particular Skate in American 
waters, but a few are dried and salted in Iceland."" 

Range. Continental waters in both sides of the northern North Atlantic in boreal 
and subarctic latitudes. On the east, from the White and Barents seas, Bear Island, 
Spitzbergen, Iceland, Norway, and the western part of the Baltic to the Dutch Coast, 
and reported doubtfully from Belgium and the Bay of Biscay ;i3i in the west from Labra- 

126. The lethal upper temperature for it is 26.5-26.9° C, according to experiments by Huntsman and Sparks (Contr. 
Canad. Biol., N. S. 2, 1924: 102). 

127. The bottom water at depths of 25-50 fathoms is colder than 0° C (32° F) for the most part from July into Septem- 
ber along northeastern Newfoundland and Atlantic Labrador. See Iselin (Proc. Amer. Acad. Arts Sci., 66, 1930: 
3-25) and Smith, Soule and Mosby (Sci. Res. Marion-Gen. Greene Exped., U. S. Cst. Guard, 2, 1937: fig. 56) 
for bottom temperatures along northeastern Newfoundland and Atlantic Labrador; for the Grand Banks region, 
see tables and charts in Rep. Newfoundland Fish. Res. Comm. [i [4], 1932; 2 [i], 1933; 2 [2], 1934; 2 [3], 
1935); for the West Greenland fishing grounds, see Beauge (Rev. Trav. Off". Sci. Techn. Peches marit., 4 [i], 
1931: 111-112; 5 [i], 1932: 59-68; 6 [i], 1933: 97-100) and Smith, Soule and Mosby (Sci. Res. Marion-Gen. 
Greene Exped., U. S. Cst. Guard, 2, 1937: figs. 13, 15, 20). 

128. July-November, 1927-1930. 129. Albatross III, May ii-iS, 1950, between Long. 67°io' and 72°2o'W. 

130. Saemundsson, Skr. Komm.' Havunders. Kbh., 5, 1909: 126. 

131. For a recent summary (with references) of the distribution oi K. radiata in the eastern Atlantic and tributary seas, 
see Clark (Rep. Fish. Bd. Scot. [1926], i, 1926: 29). 



Fishes of the Western North Atlantic 263 

dor, West Greenland, Hudson Bay, Grand Banks region, and Gulf of St. Lawrence 
to South Carolina. 

Occurrence in the Western North Atlantic. R. radiata is common in depths of 30- 
330 fathoms all along the west coast of Greenland from Disko Bay (about Lat. 69°26'N) 
to the vicinity of Cape Farewell, both offshore and in the fjords; many have also been 
taken on Fylla Bank and in the central part of Davis Strait. "^ Knowledge of it off the 
coasts of subarctic America is confined to one report for the southeastern part of Hud- 
son Bay,"* a second from Lake Melville, Labrador"* and another from off the outer 
coast of Labrador in the offing of Hamilton Inlet. "^ However, it is common on the 
fishing grounds all around the marginal belt of the Grand Banks area as a whole (south- 
ward from the latitude of Conception Bay) and at the mouths of the larger bays (Pla- 
centia, St. Mary) on the southern coast of Newfoundland. It is widespread throughout 
the southern side of the Gulf of St. Lawrence as far inward as the estuary of the St. Law- 
rence River, and doubtless it is present along the eastern (Newfoundland) side of the 
Gulf as well, to judge from its reported occurrence in the southern side of the Strait of 
Belle Isle. It occurs also along the outer Nova Scotian shelf and fishing banks, through- 
out the Gulf of Maine, and over Georges Bank. However, most of the positive records 
for it westward from about the longitude of Cape Cod (70° W) have been from a nar- 
row belt that extends along the upper part of the continental slope between the 1 50 
and 300 fathom contours, about as far south as the so-called Hudson Trough (about 
Long. 72° W) that cuts across the shelf off New York. But we have seen one specimen 
taken 25 miles off the eastern end of Long Island, New York at a depth of only 32 
fathoms; and we have another that was caught off Charleston, South Carolina, in 74 
fathoms."* 

Synonyms and References: 

Raja^^'' radiata Donovan, Nat. Hist. Brit. Fish., 5, 1807: pi. 1 14 (ill., Gt. Britain); Gill, Proc. Acad. nat. Sci. 
Philad., Addend., 1861: 61 (listed, Atlant. Coast N. Amer.); Garman, Proc. Boston Soc. nat. Hist. 
I'], 1874: 178 (descr., cf. other species, Mass.); Goode and Bean, Bull. Essex Inst., 11, 1879: 28 (Mass.); 
Jones, List Fish. Nova Scotia, 1879: 11 (listed, Nova Scotia); Bean, Proc. U. S. nat. Mus., J, 1880: 
1 1 5 (Nova Scotia and Mass. locals.); Verrill, Amer. J. Sci., (3) 22, 1881 : 297 (small nos. 160-241 fath. 
off Gay Head); Jones, Proc. N. S. Inst. Sci., 5 (i), 1883: 97 (same as Jones, 1879); Jordan and Gilbert, 
BuD. U. S. nat. Mus., 16, 1883: 41 (descr.); Bean, Rep. U. S. Comm. Fish. (1882), 1884: 343 (listed. 
Woods Hole); Jordan, Rep. U. S. Comm. Fish. (1885), 1887: 799 (listed, north. U. S., Europe); Goode 
and Bean, Smithson. Contr. Knowl., JO, JJ, 1895; also, Mem. Harv. Mus. comp. ZooL, 22, 1896: 
25, pi. 9, fig. 27 (ill., depth off New England Coast); Cox, Bull. nat. Hist. Soc. N. Brunsw., 13, 1896: 
75 (Bay of Fundy); Jordan and Evermann, Bull. U. S. nat. Mus., 47 (i), 1896: 69 (descr., N. Atlant., 
Amer., Europe); Rep. U. S. Comm. Fish. (1895), 1896: 221 (listed, N. Atlant., Amer., Europe); Gar- 

132. For a general survey of its occurrence in West Greenland waters, see Jensen (Spol. Zool. Mus. Hauniensis, 9, 1948 : 
28). 

133. Vladykov, Contr. Canad. Biol., N. S. S, 1933: 17- 

134. Reported by David C. Nutt and identified by us from a photograph. Caught by the Blue Dolphin in 20 fathoms, 
with otter trawl, July 25, 1950. Length about 275 mm. For additional Labrador records, just pubhshed, see 
Backus (Copeia, 1951: 289). 

135. Rep. Newfoundland Fish. Res. Comm., 2 (i), 1933: 125; Lat. 54° 15' N, Long. 55°05' W. 

136. Taken by the Albatross III February i, 1950 in Lat. 33°io' N, Long. 77°25' W (Harv. Mus. Comp. Zool., No. 
37061). 

137. Also spelled Kaia. 



264 Memoir Sears Foundation for Marine Research 

man, Mem. Harv. Mus. comp. ZooL, 24, 1899: 380 (depth, range); Smith, Bull. U. S. Fish Comm., 
I-j, 1898: 89 (listed. Woods Hole); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 12 (New 
England locals., but ref. for New London (.'), Conn, based on R. americana DeKay 1842, which equals 
R.eglanteria Bosc 1802); Roule, Bull. Inst, oceanogr. Monaco, 243, 1912: 20 (depth. Grand Banks); 
Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 4 (3), 191 3: 102 (range); Halkett, Check List Fish. Canad., 
191 3: 42 (listed, Maritime Provinces and southward); Sumner, Osburn and Cole, Bull. U. S. Bur. Fish., 
JJ, 1913: 738 (listed, Woods Hole); Bigelow, Bull. Mus. comp. Zool. Harv., 5<§, 1914: 100, iii 
(depth, Gulf of Maine); Huntsman, Trans, roy. Soc. Canad., (3) 12 (4-5), 1918: 63 (depth, temp., 
Gulf St. Lawrence); Cox, Contr. Canad. Biol. (1918-1920), 1921: 109 (depth, S. side Gulf St. Law- 
rence); Huntsman, Contr. Canad. Biol. (1921), 3, 1922: 6, 9 (Passamaquoddy Bay and Bay of Fundy); 
Huntsman and Sparks, Contr. Canad. Biol., N. S. 2, 1924: 102 (upper lethal temp.); Battle, Trans, 
roy. Soc. Canad., Sect. 5, (3) 20 (i), 1926: 127, 132 (effects of high temp.); Huntsman, Trans, roy. 
Soc. Canad., Sect. 5 (3) 20 (2), 1926: 203 (upper lethal temp.); Nichols and Breder, Zoologica N. Y., 
9, 1927: 24 (ill., old Woods Hole record); Vladykov, Contr. Canad. Biol., N. S. 8, 1933: 17 (SE shore, 
Hudson Bay); Nat. canad., 6j, 1936: 123, 125 (meas., photos egg cases, estuary St. Lawrence R.; R. 
radiata considered distinct from R. scabrata); Contrib. 17, Dept. Pech. Quebec, 1946: 26 (in stomach 
of White Whale, estuary St. Lawrence R.); Jensen, Spol. Zool. Mus. Hauniensis, 9, 1948: 27 (descr., 
Greenland). 

Raja clavata or R.radtata Storer, Rep. Fish. Rept. Birds Mass., 1839: 201 (Mass.); Boston J. nat. Hist., 2, 1839: 
556 (same as Storer, 1839). 

Rata laevis (in part) Storer, Mem. Amer. Acad. Arts Sci., N. S. 2, 1867: 242, pi. 39, fig. 2; also Fishes Mass., 
1867: 266, pi. 39, fig. 2 (ill., but descr. partly based on R.laevh, Mass. Bay). 

Raia scabrata Garman, Mem. Harv. Mus. comp. Zool., 56, 191 3: 340, pi. 21, fig. 2, pi. 44, figs. 1-3 (descr., 
ills., Mass. Bay, but pi. 21, fig. i, labelled R. scabrata, is actually R. ocellata); Jordan, Copeia, 49, 191 7: 
87 [scabrata replaces radiata for American form); Bigelow and Welsh, Bull. U. S. Bur. Fish., 40 (i), 
1925: 62 (descr., ill. Gulf of Maine); Breder, Field Bk. Mar. Fish. Atlant. Coast, 1929: 30 (genl.); 
Jordan, Manual Vert. Anim. NE U. S., 1929: 17 (diagn.); Jordan, Evermann and Clark, Rep. U. S. 
Comm. Fish. (1928), 2, 1930: 25 (Mass. Bay, considered distinct from European R. radiata); Lutz, 
Amer. J. Physiol., 94, 1930: 135 (effects of adrenalin) ; Marshall, Contr. Canad. Biol., N. S. 7 (1-7), 
1931: 19 (heart physioL); Schroeder, Bull. Boston Soc. nat. Hist., 58, 1931: 3 (nos., depth. Gulf of 
Maine and Georges Bank); Anon., Rep. Newfoundland Fish. Res. Comm., I (4), 1932: 117 (Newfound- 
land Coast, Newfoundland and Nova Scotian banks); Rep. Newfoundland Fish. Res. Comm., 2 (i), 
1933: 125 (depth, temp., off Nova Scotia, Grand Banks, and outer Labrador); Prefontaine, Trans, roy. 
Soc. Canad., (3) 27 (5), 1933: 257 (depth, estuary of St. Lawrence R.); Bigelow and Schroeder, Canad. 
Atlant. Fauna, I2e, 1934: 23 (descr., ill., Hudson Bay to Woods Hole); Anon., Rep. Newfoundland 
Fish. Res. Comm. (1933), 2 (2), 1934: 115 (Newfoundland, east. Nova Scotian Banks and Gulf of 
St. Lawrence); Vladykov and McKenzie, Proc. N. S. Inst. Sci., ig, 1935: 51 (off Nova Scotia); Bigelow 
and Schroeder, Bull. U. S. Bur. Fish., 48, 1936: 324 (descr., size, female with eggs, egg cases, food, 
nos., Georges Bank); Vladykov, Nat. canad., 63, 1936: 223 (descr., size, photos egg cases, depth, temp., 
season, off Nova Scotia); Norman, Discovery Rep., 16, 1937: 16, footnote {R. scabrata Garman 1913 
probably ident. with European R. radiata); Bigelow and Schroeder, Biol. Bull. Woods Hole, y6, 1939: 
309 (deep basins, Gulf of Maine). 

Doubtful References: 

Raja radiata Fowler, Proc. Acad. nat. Sci. Philad., 62, 1910: 470 (egg cases with embryos "perhaps" this species. 

New Jersey). 
Raja scabrata Fowler, Proc. biol. Soc. Wash., JJ, 1920: 145 (same as R. radiata Fowler 1910). 

Raja senta Garman 1885 

Figures 57, 58 

Study Material. Sixty specimens, 83 mm (newly hatched) to 547 mm long, from: 
Emerald Bank off Halifax, Nova Scotia; St. Andrews, New Brunswick; various localities 



Fishes of the IV es tern North Atlantic 

A 



265 




Figure 57. Raja senta. A Female, 424 mm long, from Emerald Bank, Nova Scotia (Harv. Mus. Comp. Zool., 
No. 33919)- B Lower surface of anterior part of head of same, about 0.4X . C Lower side of anterior part of tail 
of same, about 0.6 X . D Right-hand nostril of same, about 3 .3 X - £ Upper teeth of same, 2nd-6th rows, about 
6.5 X . f Juvenile female, 1 59 mm long, from Cashes Ledge, Gulf of Maine, Lat. 42°53' N, Long. 68°58' W 
(Harv. Mus. Comp. Zool, No. 34768). 

in the Gulf of Maine; the offings of southern New England and New Jersey; also one 
specimen from off Charleston, South Carolina; all in the collections of the Harvard 
Museum of Comparative Zoology and the U. S. National Museum. 



2 66 Memoir Sears Foundation for Marine Research 

Distinctive Characters. R. senta Is unique among hard-nosed Skates {Raja) of the 
North and South Atlantic from the time it has grown to one-fourth its mature size^'* 
in that the thorns on the tail dwindle in size rearward until those along the posterior 
half to third are no longer distinguishable from the small prickles with which the tail 
is generally clothed. Newly hatched specimens, in which this character is not yet 
established, are separable from other hard-nosed Skates of the western North Atlantic 
by the color pattern of the tail, which has two pale crossbars, each outlined anteriorly 
and posteriorly by a dark band or blotch. They are set apart from the barred-tailed species 
of Breviraja by the long rostral cartilage (reaching to tip of snout). 

Description. Proportional dimensions in per cent of total length. Female, 424 mm, 
and male, 520 mm, from Emerald Bank, off Nova Scotia (Harv. Mus. Comp. Zool., 

No. 33919)- 

Disc: extreme breadth 62.0, 63.5; length 51.2, 50.5. 

Snout length: in front of orbits 15. i, 12.5; in front of mouth 15.3, 12.8. 

Orbits: horizontal diameter 4.2, 3.7; distance between 3.8, 4.0. 

Spiracles: length 2.6, 2.5; distance between 6.6, 7.1. 

Mouth: breadth 7.3, 8.1. 

Nostrils: distance between inner ends 7.3, 7.1. 

Gill openings: lengths, ist 1.9, 1.8; 3rd 1.8, 1.8; 5th 1.2, 1.3; distance between 

inner ends, ist 13.7, 13.7; 5th 7.5, 6.0. 
First dorsal fin: vertical height 2.2, 2.6; length of base 5.4, 5.2. 
Second dorsal fin: vertical height 2.4, 2.5; length of base 5.0, 6.1. 
Pij/wVj; anterior margin 14.6, 13.6. 
Distance: from tip of snout to center of cloaca 47.5, 46.5 ; from center of cloaca 

to 1st dorsal 38.7, 39.0; to tip of tail 52.5, ^2)-S'-> from rear end of 2nd 

dorsal base to tip of tail 3.3, 3.1. 
Interspace between: ist and 2nd dorsals 0.0, 0.0. 

Disc about 1.2— 1.3 times as broad as long; maximum anterior angle in front of 
spiracles about 110°; anterior margins nearly straight rearward from tip of snout in 
adult females and in young of both sexes, but bulging moderately in front of level of 
eyes in adult males; outer and posterior corners broadly rounded, posterior margins 
strongly convex. Axis of greatest breadth about 70 "/o of distance back from tip of snout 
toward axils of pectorals. Tail with lateral folds confined to posterior two-thirds and so 
narrow that they are likely to be overlooked; its length from center of cloaca to first 
dorsal about as great as distance from center of cloaca to tip of snout in small specimens, 
decreasing in relative length with growth to about 80—90 "/o as great in adults; extreme 
length of tail from center of cloaca about i.i times as great as length from cloaca to 
snout in one-third-grown specimens and larger. 

Small specimens closely and uniformly prickly above on entire disc, on inner parts 
of pelvics and on tail; 1—2 large thorns in front of orbit and as many behind it; 4—5 

138. It is also distinguishable from hard-nosed Skates of the Pacific Coast of America by the same character. 



Fishes of the Western North Atlantic 



267 




Figure 58. Raja senta. A Male, 520 mm long, from Emerald Bank, Nova Scotia (Harv. Mus. Comp. Zool., 
No. 33919). B Margin of right-hand nasal curtain of same, about 3.7X. C Upper teeth of same, about 7.4X. 
D Side view of posterior part of tail of female pictured in Fig. 57, about 0.9 X. 



268 Memoir Sears Foundation for Marine Research 

around inner margin of orbit; one on each shoulder; a row of thorns along midline of 
back, lo— 12 large ones anterior to axils of pelvics, followed by successively smaller ones 
along tail until no longer distinguishable from the prickles a short distance anterior to 
first dorsal. Large specimens of both sexes with 2—3 preocular thorns, 1-2 postoculars, 
about eight along inner margin of orbit, 3—5 on each shoulder, and 16 or more larger 
and smaller thorns spaced irregularly along midline of disc; tail usually with 20—30 
thorns in midrow along anterior one-half to two-thirds, these of different sizes, progres- 
sively smaller rearward; each side of tail low down with 1—3 irregular rows of thorns 
along anterior part, grading down to minute prickles above, below, and rearward; pos- 
terior one-third to one-half of tail without large thorns — the most distinctive character 
of the species. Females remain prickly on the upper surface as a whole to maturity (in 
addition to thorns), densely so on the tail, but irregular bare areas develop in the general 
region of the shoulders and around the outer parts of the pelvics. Maturing males lose 
the prickles from the central portion of the disc as a whole, but they develop a few 
medium-sized thorns on the anterior part of the rostral ridge; also smaller thorns over 
the anterior parts of the pectorals (only prickly in females) and large thorns over a roughly 
triangular area on either side abreast of the eye and anterior to it. Alar spines of mature 
males in two rows on either side, 13—14 in each row. Lower surface of disc smooth in 
small specimens, but prickles develop later along a narrow marginal band from snout 
back about halfway toward level of nostrils; lower surface of tail either densely prickly 
throughout on young of both sexes and on females to maturity, or at most with a narrow 
naked median band, but becoming generally smooth in males by maturity. 

Snout in front of orbits about 3.1—4.0 times as long as distance between orbits, 
its length in front of mouth about 1.8— 2.1 times as long as distance between exposed 
nostrils. Orbit about 1.5 times as long as spiracle. Distance between first gill openings 
1.9 times as long as distance between exposed nostrils, 0.9—1.0 times between fifth 
gill openings; first gill openings 1.4— 1.6 times as long as fifth and about 26 % as 
long as breadth of mouth. Nasal curtain as well as expanded posterior (outer) margin 
of nostril fringed. Mouth moderately arched in adult females, more strongly so in 
adult males. 

Teeth ||Ez|g) evenly rounded in anterior rows but with faintly indicated cusps in 
posterior rows and closely crowded in quincunx in young specimens and in females to 
maturity; but teeth of mature males with high, recurved, sharp-pointed cusp and spaced 
more loosely in transverse series. 

First and second dorsals similar in size and shape, confluent at base. Caudal mem- 
brane posterior to base of second dorsal about half as long as base of first dorsal. Pelvics 
deeply concave outwardly and strongly scalloped around the indentation ; anterior margin 
about 75—85 "/o as long as distance from its own origin to rear tip of pelvic in half-grown 
and large specimens, as much as 90 "/o in young; anterior lobe narrow, including only 
two radial cartilages besides the first and stoutest; posterior lobe with moderately convex 
outer margin and well rounded posterior corner. Claspers of mature male extend back 
about 1/2 to '/5 the distance from axils of pelvics toward first dorsal. 



Fishes of the Western North Atlantic 269 

Anterior rays of pectorals extend forward about 80-85 "/q of distance from level 
of fronts of orbits toward tip of snout. 

Color. Upper surface pale brown with numerous obscure darker spots, sometimes 
with irregular pale markings; tail similar to disc in adults, but newly hatched specimens 
with two transverse pale bars, each outlined by a dark crossbar in front of it and another 
behind. These bars are lost with growth, so that specimens of 180-200 mm may show 
either one or two; specimens of 280-300 mm have either one or none. Lower surface 
of disc and pelvics either plain white or with a few small dusky spots on outer parts of 
pectorals, or one at tip of snout; the tail either white below, variously dark-blotched, or 
uniformly dark along the posterior third. 

Size. The three smallest specimens we have seen — apparently hatched not long 
previously but with the filamentous tip of the caudal already lost — were 83-87 mm long. 
A male of 515 mm is shown to be mature or nearly so by the size of its claspers. The 
maximum length reported is 24 inches (610 mm). 

Developmental Stages. Egg cases, probably of this species, are dark brown or almost 
black, squarish and smooth-shelled, 56-59 mm long (exclusive of horns) by 35-39 mm 
broad, one side nearly flat but the other strongly convex. The horns are thick and at 
least as long as the capsule (tips broken off in all specimens seen)."* 

Habits. Nothing is known of the diet of this Skate. It has been taken, apparently 
with equal regularity, on the soft mud and clay bottoms of the deeper troughs and basins 
as well as on the sand, sand and shells, gravel, and pebbles of the offshore fishing banks. 
Egg cases, some empty and others containing embryos, apparently of this species,"" have 
been trawled in the estuary of the St. Lawrence River at i 50-300 meters (82-164 fath.) 
in July and August. The fact that an empty egg capsule has been found on the shore 
near Halifax, Nova Scotia, makes it probable that R. senta breeds throughout its geo- 
graphic range. 

The shoalest captures of R. senta that have been definitely recorded were at 25 
and 44 fathoms in the southwestern part of the Gulf of Maine, off Cape Cod, and the 
great majority of records range from 50-250 fathoms off southern New England "^ and 
210 fathoms off New Jersey to 478 fathoms (one specimen) oft' South Carolina. It is 
recorded from 82-178 fathoms in the Gulf of St. Lawrence, and from 50-100 fathoms 
on LaHave and Emerald banks off outer Nova Scotia. 

The lowest temperature from which R. senta has been definitely recorded, in the 
estuary of the St. Lawrence River, was about 2-4° C (36-39° F). The local populations 
may also be exposed to temperatures equally low on the banks off southeastern Nova 
Scotia into the summer when the ice-cold outdraught from the Gulf of St. Lawrence ex- 
tends coastwise in that direction ;i*2 similar exposure to cold waters is likely on Georges 

139. For description and photographs, see Vladykov (Nat. canad., 6j, 1936: 220-221). 

140. Separable by their small size from those of K. radiata, the only other Skate that is known to occur in the region 
where they were collected. 

141. K. senta was taken in eight hauls deeper than 100 fathoms by Albatross III between Long. 67°io' and 72°2o' W, 
but not in 55 hauls shoaler than that. 

142. The bottom temperature on July 29, 1914 was about 2.8° C (37° F) on La Have Bank at 95 fathoms, but 5.6° C 
(42° F) at 80 fathoms on Emerald Bank off Halifax a week later. 



270 Memoir Sears Foundation for Marine Research 

Bank during March when the water is coldest, and likewise in the inner parts of the 
Gulf of Maine, if any R. senta remain there in shallow water of 30—40 fathoms through 
the late winter and spring. 

The fact that R. senta has never been reported from water colder than about 2° C 
(36° F) suggests that low temperature is the barrier that bars it from regularly populating 
the more easterly sections of the Nova Scotian (Banquereau) or the Newfoundland banks, 
for these bottoms are flooded yearly with water as cold as 0° C (or even a little colder) 
at seasons when the icy outflow from the Gulf of St. Lawrence (in the first case) or the 
southerly expansion of the Labrador Current (in the second) is at its maximum. At the 
other extreme, some of the specimens that have been taken along the continental edge 
off New England and farther south may have been from water about as warm as 1 1° C 
(52° F); however, most of them have been taken from water with temperatures between 
about 6° C (42 or 43° F) and about 9-10° C (49 or 50° F).i" 

Generally R. senta has been taken from water at least as saline as 32.6 "/oo, most 
of them from water with salinities higher than 32.8 "/oo; undoubtedly specimens along 
the continental edge live in salinities as high as 34.7 "/oo and probably as high as 35.0- 
35.4 "/oo- There is nothing in the available records to suggest that R. senta carries out 
any regular seasonal migrations. 

Numerical Abundance. Most of the records for this particular Skate report the cap- 
ture of odd individuals only. However, 57 specimens of i?. senta^ out of a total catch of 
495 Skates, were taken in 37 trawl hauls on Georges Bank in September 1929, thus 
giving an average of one to two per haul and indicating a population density similar to 
that of R. ocellata (about two per haul) or R. laevis although much smaller than that of 
R. radiata (about eight to nine per haul). Off^ southern New England and New Jersey, 
13 R. senta out of a total catch of 77 Skates were taken in eight trawl hauls deeper than 
100 fathoms (about one to two per haul).^** 

Range. Atlantic shelf of North America from the latitude of Charleston, South 
Carolina to the Nova Scotian Banks and Gulf of St. Lawrence; as a stray to the southern 
part of the Newfoundland Bank. 

Occurrence in the Western North Atlantic. Raja senta, once considered a rare species, 
is now known to occur generally in water of sufficient depth throughout the western 
side of the Gulf of Maine 1*^ and along the Nova Scotian side of the Bay of Fundy; this 
distribution suggests that it will prove as widespread at appropriate depths in the eastern 
side of the Gulf as it is on Georges Bank. It is known also from the southeast slope of 
Browns Bank,"* from LaHave Bank off southeastern Nova Scotia, and from Emerald 
Bank off Halifax, as well as from the estuary of the St. Lawrence River. It has been 

143. To judge from the depths and localities of capture; one specimen from the offing of South Carolina was taken 
in water of 4° C (39.3° F). 

144. By Albatross III between Long. 6 7° 10' and 72°2o' W, May ii-iS, 1950. 

145. We have taken it on several occasions in the trough west of Jeffreys Ledge, in the west central part of the Gulf 
near Cashes Ledge, and at three stations east and southeast from Cape Cod; it has been reported from the Bay 
of Fundy and off Provincetown at the tip of the Cape. 

146. Trawled by Caryn of the Woods Hole Oceanographic Institution, June 18, 1949, at Lat. 42°! 8' N, Long. 65°oi' W, 
in 380-420 fathoms. 



Fishes of the Western North Atlantic 271 

reported doubtfully from the northern side of the inner part of the Gulf of St. Lawrence, 
but it has never been reported from the Nova Scotian shelf east of the offing of Halifax, 
and it is so rare on the Newfoundland Banks that only a single specimen was taken among 
the many Skates of other kinds during the trawling experiments of the Newfoundland 
Fisheries Research Committee during the years 193 i — 1934. "' To the westward and 
southward it occurs regularly, if in small numbers (p. 270), along the upper part of the 
continental slope in the offing of southern New England, of New York, and of New 
Jersey"^ within the appropriate depth-zone. One has also been taken at the edge of 
the continental shelf off northern Virginia,^" one off Chesapeake Bayi^" at 104 fathoms, 
and one off Charleston, South Carolina, in 478 fathoms. "' 

Synonyms and References: 

Raja^^^ senta Garman, Proc. U. S. nat. Mus., 8, 1885: 43 (descr., Provincetown, Mass., and LaHave Bank, 
Nova Scotia); Goode and Bean, Smithson. Contr. Knowl., jo, 1895: 508; also, Mem. Harv. Mus. 
comp. Zool., 22, 1896: 508 (LaHave Bank, by ref. to Garman, 1885); Jordan and Evermann, Bull. 
U. S. nat. Mus., 47 (i), 1896: 71 (descr., Newfoundland Banks to C. Cod); Rep. U. S. Comm. Fish. 
(1895), 1896: 221 (listed, Newfoundland Banks to C. Cod); Garman, Mem. Harv. Mus. comp. Zool., 
24, 1899: 380 (depth, local.); Schmitt, Monogr. d'Anticosti, 1904: 283 (doubtful rep. N. shore Gulf 
of St. Lawrence near Anticosti); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 12 (C. Cod 
Bay by ref. to Garman, 1885); Proc. Portland Soc. nat. Hist., 2, 1909: 207 (Labrador [.'] by ref. to 
Schmitt, 1904); Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 4 (3), 1913: 102 (range); Garman, Mem. 
Harv. Mus. comp. Zool., 36, 1913: 328, pi. 25, fig. i (descr., ill., off New England); Halkett, Check 
List Fish. Canad., 1913: 42 (listed, Labrador [.^], Newfoundland Banks to C. Cod); Bigelow and Welsh, 
Bull. U. S. Bur. Fish., 40, 1925: 65 (descr., ill., old records); Breder, Field Bk. Mar. Fish. Atlant. Coast, 
1929: 31 (ill., Newfoundland to C. Cod); Jordan, Manual Vert. Anim. NE U. S., 1929: 16 (diagn.); 
Jordan, Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 25 (listed, Newfoundland Banks 
to C. Cod); Schroeder, Bull. Boston Soc. nat. Hist., 58, 1931: 2 (depth, temp., season, nos.. Gulf of 
Maine and Georges Bank); Prefontaine, Trans, roy. Soc. Canad., Sect. 5 (3) 2y, 1933: 257 (depth, 
type of bottom, estuary of St. Lawrence R.); Anon., Rep. Newfoundland Fish. Res. Comm. (1933), 2 
(2), 1934: 215 (listed, SW. part of Newfoundland Bank); Bigelow and Schroeder, Canad. Atlant. Fauna, 
I2e, 1934: 33 (descr., LaHave Bank, Gulf of Maine); Vladykov and McKenzie, Proc. N. S. Inst. Sci., 
ig, 1935: 51 (depth, LaHave and Emerald banks); Bigelow and Schroeder, Bull. U. S. Bur. Fish., 48, 
1936: 324, 325 (Gulf of Maine locals., depths); Vladykov, Nat. canad., 6j, 1936: 220, 221 (egg cases, 
season, depth, estuary of St. Lawrence R. and Bay of Fundy) ; Bigelow and Schroeder, Biol. Bull. Woods 
Hole, j6, 1939: 309 (Gulf of Maine) ; Vladykov, Contrib. 17, Dept. Pech. Quebec, 1946: 26 (in stom- 
achs of White Whale, estuary St. Lawrence R.). 

Raja species Vladykov and Tremblay, Nat. canad., 62, 1935 : 79 (apparently same specimens rep. by Vladykov, 
1936). 

Raja spinicauda Jensen 19 14 
Figure 59 

Study Material. Male, 1,236 mm long, nearing sexual maturity, taken off Bonavista 
Bay on the east coast of Newfoundland at Lat. 48°47' N, Long. 52°47' W, from be- 

147. Earlier characterizations of it as ranging to the Newfoundland Banks appear to reflect a failure to realize that LaHave 
Bank (one of the type localities of the species) actually lies off southeastern Nova Scotia. 

148. Taken by Caryn in three hauls near Lat. 39^55' N, Long. 7o°4o' W, June and October 1948; in five hauls by 
Albatross III between Long. 67°io' and 72°2o' VV in the 100-240 fathom zone, May 1950; also one specimen 
from off New York, in U. S. National Museum; and other specimens seen from offshore trawling stations. 

149. Lat. 37°36' N, Long. 74°i5' W. 150. Lat. 37°03' N, Long. 74°3i' W. 
151. Albatross Sta. 2677, Lat. 32°39' N, Long. 76°5o' W. 152. Also spelled Rata. 



272 Memoir Sears Foundation for Marine Research 

tween 91 and 127 fathoms and at a temperature between 0.65 and -1.5° C, in collec- 
tion of Newfoundland Department of Natural Resources. Also two egg-cases, appar- 
ently of this species (p. 275), one with embryo from Banquereau Bank off Nova Scotia, 
Lat. 44° 1 5' N, Long. 58°03' W (source unknown), the other empty, from the conti- 
nental slope off southwestern Georges Bank, Lat. 3 9° 5 3' N, Long. 69°oo' W, from 
between 260 and 350 fathoms, trawled by the research vessel Caryn of the Woods Hole 
Oceanographic Institution, September 11, 1949. 

Distinctive Characters. Although the upper surface of its disc and tail are more or 
less prickly (see p. 274), R. spinicauda is set apart from all other members of its genus 
in the western North Atlantic (Greenland to the Amazon) by the fact that its only large 
thorns are in a single row of 2 1-26 extending from the level of the axils of the pelvics 
rearward along the midline of the tail. In these respects it falls closest to R. laevis. How- 
ever, adult specimens of the two species are easily separable, for the tail of R. spinicauda 
bears only the single median row of thorns whereas that of R. laevis bears three rows, 
and while the mucous pores on the lower surface of R. laevis are marked with black pig- 
ment dots or streaks, those of R. spinicauda are not so marked. 

Description. Proportional dimensions in per cent of total length. Male, 1,236 mm, 
from Lat. 48°47' N, Long. 52°47' W (specimen from Newfoundland Department Nat- 
ural Resources). 

Disc: extreme breadth 68.0; length 56.6. 

Snout length: in front of orbits 17.8 ; in front of mouth 17.8. 

Orbits: horizontal diameter 2.8; distance between 4.9. 

Spiracles: length 3.0; distance between 7.3. 

Mouth: breadth 8.1. 

Nostrils: distance between inner ends 7.4. 

Gill openings: lengths, ist 1.4; 3rd 1.5; 5th i.i; distance between inner ends, 

1st 15.7; 5th 12.4. 

First dorsal fin: vertical height 2.0; length of base 3.3. 

Second dorsal fin: vertical height 2.0; length of base 3.0. 

Pelvics: anterior margin 8.6. 

Distance: from tip of snout to center of cloaca 52.7; from center of cloaca to 
1st dorsal 37.1 ; to tip of tail 47.3; from rear end of 2nd dorsal base to tip 
of tail 2.9. 

Interspace between: ist and 2nd dorsals 1.2. 

Disc about 1.2 times as broad as long; maximum anterior angle in front of spiracles 
about 95°; tip of snout subangular or slightly blunted; anterior outlines nearly straight 
rearward to level of spiracles, then bulging a little outwardly and following a sinuous 
outline that is weakly concave and next weakly convex toward outer corners; outer 
corners rather abruptly rounded, posterior corners much more broadly so; posterior 
margins weakly convex. Tail with narrow lateral folds extending nearly its entire length, 
at line of transition from rounded upper surface to flat lower surface; its length from 



Fishes of the Western North Atlantic 



273 







Figure 59. Raja spinkauda, male, 1,236 mm long, from off east coast of Newfoundland (Newfoundland Dept. 
Nat. Resources). A Side view of posterior part of tail, about 0.4 x. B Dorsal view of tail at about midlength, 
about 0.8 X. C Right-hand nostril and nasal curtain, about 0.6 X. D Upper teeth, about 2.7 X. 



2 74 Memoir Sears Foundation for Marine Research 

center of cloaca to first dorsal fin about 70 "/o as great as distance from center of cloaca 
to tip of snout, its length to tip nearly as great as distance from center of cloaca to 
snout in halfgrown specimens, relatively a little less in adults. 

Disc smooth along a narrow marginal band rearward from tip of snout in males 
to maturity and probably in females as well; rough elsewhere with large prickles or 
small spines, closest together abreast of eyes, on outer parts of pectorals, and over 
rump region; sparser between orbits and outward from shoulders; males may develop 
scattered naked areas at maturity on central parts of pectorals; tail thickly and 
uniformly prickly above, sparsely so along lateral folds; disc without thorns (except for 
alar spines of males), but tail with a median row of 21— 26 large and conspicuous thorns 
on strongly striate bases from a little anterior to axils of pectorals to first dorsal fin; 
also one large thorn in interspace between first and second dorsals; inner parts of pelvics 
prickly, outer portions smooth; dorsals prickly; caudal membrane smooth or with a few 
minute prickles. Alar spines of sexually mature males in 3—4 rows. Lower surface of 
disc smooth, but tail with a narrow prickly band along either margin rearward from 
axils of pelvics. 

Snout anterior to orbits about 3.6 times as long as distance between orbits, its 
length in front of mouth about 2.5 times as great as distance between exposed nostrils. 
Orbit a little more than half as long as distance between orbits, and about as long as 
spiracle. Distance between first gill openings 2.1 times as long as distance between ex- 
posed nostrils, between fifth gill openings 1.7 times; first gill openings 1.2 times as 
long as fifth and about 1 7 "/o as long as breadth of mouth. Nasal curtain fringed with 
about 10 short rounded lobelets; expanded posterior (outer) margin of nostril smooth. 
Mouth nearly straight in females, a little arched in adult males. 

Teeth l^jf, with sharp thorn-like cusp on rounded base in both sexes, rather loosely 
spaced in regular transverse series, much as in R. hyperborea and R. lintea. 

First and second dorsals about equal in size, brush-shaped, with jagged posterior 
margin on specimen seen. Interspace between dorsals a little less than half as long as 
base of first dorsal. Caudal membrane beyond second dorsal a little shorter than base 
of first dorsal. Pelvics deeply concave outwardly; anterior margin about 50 "/o as long 
as distance from its own origin to rear tip of pelvic; anterior lobe broadly rounded at 
tip and strongly scalloped along posterior edge, thus marking the positions of radial 
cartilages ; posterior lobe with moderately convex and somewhat wavy outer margin, the 
rear tips subangular, reaching back about Vs of the way from axils of pelvics toward 
first dorsal. 

Anterior rays of pectorals extending forward about 85 % of distance from level 
of fronts of orbits toward tip of snout. 

Color. Upper surface uniformly pale brown or bluish gray without dark markings, 
paler or whitish around posterior edges of pectorals ; lateral folds on tail either of same 
color as upper surface or white. Lower surface of disc white with small sooty blotches 
irregularly distributed around margins of pectorals; pelvics white below, sometimes with 
sooty margins ; lower surface of tail white anteriorly but pale brown rearward from a 



Fishes of the Western North Atlantic 275 

little posterior to axils of pelvics, crossed on posterior quarter by two irregular yellowish 
white bands. 1^' 

Relationship to Extralimital Species. In the southern hemisphere R. spinicauda has 
a close counterpart in R. griseocauda Norman 1937"* of the Patagonian-Falkland Islands 
region. The resemblance between these two Skates is so close, in fact, that only a com- 
parison of specimens from the two regions can show whether there are significant dif- 
ferences between them. The relationship between the members of this pair thus parallels 
that between R. radiata of the northern North Atlantic and R. doello-juradoil?ozx\ 1 933'^* 
of Argentina, Patagonia, and the Falkland Islands (p. 255). Among Skates of the north- 
eastern Siberian-Bering Sea region, R.violacea Uvarov 1935^^^ resembles R. spinicauda 
most closely. But the former has only about half as many thorns along the tail as R. spini- 
cauda^ and there is an extensive area free from prickles on either side of the mid-dorsal 
belt of its disc. 

Size. Specimens measured (about nine males and females) have ranged from 740— 
1,720 mm in length, showing this to be one of the larger Skates of northern seas. A 
male 910 mm wide (hence about 1,340 mm long) with slender claspers was sexually 
mature. 

Developmental Stages. Egg capsules, apparently of this species, in our Study Mate- 
rial (p. 272), measure 137 by 90 mm and 142 by 95 mm. They are of the usual quadrate 
shape, with nearly straight transverse outline at the end bearing the longer pair of horns, 
concave at the end with the shorter pair. The lateral flanges are narrow, continue out 
along the basal parts of the horns, and bear masses of fine filaments matted together and 
more or less interconnected by a delicate membrane, as is true of various other Skate 
eggs. In the preserved state these masses of filaments are irregularly scattered along the 
flanges (chiefly near their extremities); but they may have been continuous along the 
flanges when the eggs were first laid. On the larger specimen, one of the horns of the 
shorter pair is about 115 mm long, with filamentous tip; all of the horns on both of the 
specimens have lost their extremities. The most interesting feature of these capsules, 
and one that sets them apart from those of any other Atlantic Skates whose eggs are 
known, is that their entire surface, apart from the extremities of the horns and the 
lateral flanges, bears a great number of low narrow longitudinal ridges, each close-set 
with a single series of numerous stiff rod-like structures about two mm long, with com- 
plexly dissected tips. On one of our capsules there are between 140—150 ridges on each 
side, each with 200—220 or more rods, or a total upwards of 30,000 on each side, giving 
the capsule a velvety appearance. 

Similar egg capsules, taken from shark's stomachs off Southwest Greenland, have 
already been credited to R. spinicauda because they were certainly not referable to any 
other Skate known from Greenland. i" And their identity as egg cases of R. spinicauda 
is now made highly probable by the wedge-shaped anterior contour of the embryo con- 

153. On specimen seen. 154. Discovery Rep., 16, 1937: 26. 

155. Physis B.Aires, 11, 1935: 491; see also Norman, Discovery Rep., 16, 1937: 15. 

156. Bull. Acad. Sci. Leningrad [1935], 1935: 3, 431, 432, fig. i- 

157. Jensen, Mindeskr. Steenstr. Fods. Kbh., 2 [30], 1914: 33, figs. 10, 11; Spol. Zool. Mus. Hauniensis, 9, 1948: 56. 



276 Memoir Sears Foundation for Marine Research 

tained in one of those in our Study Material, combined witli the presence of a single 
series of translucent dots along the mid-dorsal line of the tail, seemingly presaging the 
later development of caudal thorns, with no indication of future thorns anywhere on 
the disc.i^* 

Habits. Nothing whatever is known of the way of life of R. spinkauda beyond the 
fact that the few specimens taken so far have come from depths of: 120-440 fathoms, 
where bottom temperatures ranged between 0.5 and 3.8° C (about 32 and 39° F) in 
West Greenland waters; 66-88 fathoms (120-160 m) off Iceland; 90-140 fathoms in 
Newfoundland waters, where the temperature of the bottom water was between + 0.65 
and -1.5° C; and, if our identification of two egg capsules in our Study Material 
(p. 275) is correct, 260-350 fathoms off southern Nova Scotia and on the continental 
slope off the southern part of Georges Bank. Several taken off Southwest Greenland 
had fed on capelin (Mallotus) and on the Starry Skate {Raja radiata). 

Range. R. spinkauda is known definitely only from: Barents Sea; off the south- 
east coast of Iceland; from the Greenland side of Davis Strait, Lat. 65° 14' N, Long. 
55° SS'^'-> from Tunugdliarfik Fjord, Skovfjord, and Julianehaab,!^' Southwest Green- 
land; from the continental slope (91-127 fath.) off Cape Bonavista, east coast of New- 
foundland; and from the deep trough of Hermitage Bay (90-140 fath.) on the south 
coast of Newfoundland (two specimens).i«'' To judge from the distribution of these local- 
ities, it is to be expected also in other bays along the coasts of eastern Labrador and of 
eastern and southern Newfoundland, wherever the water is deep enough and the bottom 
temperature lower than 2-3° C (35-37° F); also around the slopes of the Grand Banks 
at suitable depths. And, if the identification of egg capsules in our Study Material is 
correct (p. 275), R. spinicauda ranges westward along the continental slope at least as 
far as the offing of southwestern Georges Bank. 

Synonyms and References: 

Raja spinicauda Jensen, Mindeskr. Steenstr. Fods. Kbh., 2 (30), 1914: 30, 33, 38, fig. 12, pi. not numbered 
(descr., color, meas., size, ills, adult and egg case, depths, temps., Davis Str. and fjords of SW. Green- 
land); Clark, Rep. Fish. Bd. Scot. (1926), i, 1926: 49 (diagn., range, after Jensen, 1914); Faune Ichthyol. 
Cons. int. Explor. Mer, 193 1: pi. not numbered (descr., size, ills., range, depth, egg cases, W. Green- 
land, after Jensen, 1914; and Western Horn, Iceland); Lundbeck, Mitt, dtsch. SeefischVer., 4g, 1933: 
8 (Barents Sea); Jensen, Spol. Zool. Mus. Hauniensis, 9, 1948: 49 (W. Greenland). 

Raja teevani Bigelow and Schroeder 1951 
Figure 59a 

Study Material. Immature male, 558 mm in total length, type, in U. S. National 
Museum, and another immature male, 302 mm in total length, in Harvard Museum of 
Comparative Zoology, both from Lat. 29°ii'N, Long. 86°52' W, 305 fathoms, 
Oregon St. 279. 

158. Unfortunately the embryo is in poor condition. 159. Personal communication from Paul M. Hansen. 

160. These Newfoundland captures, made during experimental fishing for halibut by the research vessel of the New- 
foundland Department of Natural Resources, were reported to us by letter from W. Templeman, Director, to 
whom we are also indebted for the specimen pictured in Fig. 59. 



Fishes of the W ester ?i North Atlantic 277 

Distinctive Characters. Raja teevani differs from all other raj ids in the western North 
Atlantic in the shape of its tail which widens rearward toward the dorsal fins (in all 
other rajids it narrows rearward). It resembles R. olseni and R. laevis in general appear- 
ance and in lacking thorns along the midbelt of the disc from the level of the axils of 
the pectorals to the vicinity of the spiracles. But it differs from olseni in having no inter- 
space between the bases of the dorsal fins. From laevis of comparable size it may be 
separated by its longer snout (distance from tip of snout to eye about V4-V3 the width 
of disc in teevani, but only about Vs in laevis) and by the fact that the anterior margin 
of the pelvic fin is longer than the distance from its own origin to the rear tip of the 
pelvic (shorter in laevis). 

Description. Proportional dimensions in per cent of total length. Male, 558 mm 
(U. S. Nat. Mus., No. 153557), and male, 302 mm (Harv. Mus. Comp. Zool., No. 
37189), from the Gulf of Mexico. 

Disc: extreme breadth 72.7, 73.5; length 58.2, 54.4. 

Snout length: in front of orbits 22.2, 19.2; in front of mouth 24.2, 20.9. 

Orbits: horizontal diameter 3.2, 3.5; distance between 4.1, 2,-S- 

Spiracles: length 2.0, 1.8; distance between 6.1, 6.3. 

Mouth: breadth 7.7, 7.0. 

Exposed nostrils: distance between inner ends 8.7, 8.3. 

Gill openings: length, ist 1.7, 1.2; 3rd 1.8, 1.3; 5th 1.2, i.o; distance between 
inner ends, ist 13.8, 14.4; 5th 8.4, 8.6. 

First dorsal fin: vertical height 2.7, 2.3; length of base 4.3, 4.6. 

Second dorsal fin: vertical height 2.7, 2.3; length of base 4.0, 4.6. 

Pelvics: anterior margin 14.7, 14.0. 

Distance: from tip of snout to center of cloaca 51.7, 47.0; from center of cloaca 
to 1st dorsal 34.3, 34-8; to tip of tail 48.3, 53.0; from rear end of 2nd 
dorsal base to tip of tail 5.4, 9.3. 

Interspace between: ist and 2nd dorsals 0.0, 0.0. 

Disc 1.25— 1.35 times as broad as long, the maximum anterior angle in front of 
spiracles about 70° ; anterior margins sinuous from snout to outer corners, convex op- 
posite a little in front of orbits; outer corners sharply rounded; posterior margins gently 
convex; posterior corners broadly rounded. Axis of greatest breadth about 77 % of 
distance from tip of snout toward axils of pectorals. Tail with a lateral fold low down 
on each side beginning almost imperceptibly about opposite tips of pelvics, widening 
rearward and ending opposite middle of caudal fin; its length from center of cloaca to 
origin of first dorsal fin 0.66—0.74 ^^ great as, to its tip 0.93— i.i times as great as, 
distance from center of cloaca to tip of snout. 

One or two small thorns along inner anterior margin of orbit and another one on 
posterior margin; minute prickles scattered over interorbital area and over entire disc 
anterior to orbits; tail with a median row of 15 thorns pointing backward, beginning 
about an eye's diameter posterior to axils of pelvics and ending a little in front of first 



278 Memoir Sears Foundation for Marine Research 



0/^M>^ A 




Figure 59a. Raja teevani, male, 558 mm long, from offing of Pensacola, Florida, Lat. 29°! i' N, Long. 86°52' W 
(U. S. Nat. Mus., No. 153557, type). J Upper teeth, about 6x. B Margin of left nasal curtain, about 2.5 X. 
C Posterior part of tail, about i X • -O Mouth and nostrils, about i X ■ 

dorsal, the thorns somewhat more closely spaced and larger rearward than near row's 
origin ; on tail, minute prickles from about tips of pelvics nearly to tip ; dorsals and caudal 
with a few minute prickles. Lower surface with a band of prickles along anterior margins 
of disc from a little posterior to level of mouth to tip of snout, and also along anterior 
half of rostral cartilage. 



Fishes of the JVestern North Atlantic 279 

Snout in front of orbits 5.5-6.9 times as long as orbit; its length in front of mouth 
2.5—3.6 times as great as distance between exposed nostrils. Orbit 1.6-1.8 times as 
long as spiracle; distance between orbits r. 0—1.3 times as great as orbit. Distance be- 
tween first gill openings 1.6— 1.7 times as great as distance between exposed nostrils, 
between fifth gill openings about i.o times; first gill openings 1.2— 1.4 times as long as 
fifth and 0.17—0.22 as long as breadth of mouth. Nasal curtain fringed; expanded pos- 
terior (outer) margins of nostrils smooth except for a few fringes on extreme outer angle. 
Upper and lower jaws moderately arched centrally. 

Teeth j^eII, rather widely spaced, in quincunx, triangular or ovate, with smooth 
rounded apex on outer margin and a low triangular cusp, pointing inward, on inner 
margin. 

First and second dorsals similar in size and shape. No interspace between dorsals. 
Caudal membrane from rear end of base of second dorsal 1.25—2.0 times as long as 
base of first dorsal. Pelvics deeply concave, strongly scalloped along anterior side of 
excavation but only weakly so rearward; anterior margin i.i times as long as distance 
from its own origin to rear tips of pelvics; anterior lobe moderately slender, including 
five radial cartilages besides the first stout one; posterior lobe convex outwardly; rear 
tips abruptly rounded, extending about V's of the distance from axils of pelvics toward 
first dorsal; inner margin straight. 

Rostral cartilage firm, narrow, extending to tip of snout. Anterior pectoral rays 
reaching only about 2/5 of distance from level of front of orbits toward tip of snout. 
Translucent area in front of orbits and on either side of rostral cartilage very thin and 
membranous. 

Color. Upper surface of larger specimen pale brown, somewhat darker along pos- 
terior margins of disc, on pelvics, and on tail. Dorsal fins and caudal black. Below, creamy 
on disc except somewhat dusky along outer margins from outer angle rearward and on 
pelvics. Tail blackish. On the smaller specimen there is a narrow black margin along the 
posterior edge of the disc above, and the lower surface is distinctly margined with black 
rearward from the level of the mouth ; the rear parts of the pelvics are blackish also. 

Size and Habits. The small claspers on the larger specimen indicate that this species 
attains at least a moderately large size. Nothing is known of its habits beyond the fact 
that our two specimens were taken at a depth of 305 fathoms. 

Occurrence. It is known only from the offing of Pensacola, Florida. 

Reference : 

Raja teevani Bigelow and Schroeder, J. Wash. Acad. Sci., ^J (12), 1951: 388 (descr., ill., off Pensacola, Fla.). 

Raja texana Chandler 1921 

Texas Skate 

Figures 60, 61 

Study Material. Thirteen specimens, 105—550 mm long, from Barataria and Chau- 
vin, Louisiana and from Austin, Texas, in the collections of the U. S. National Museum, 



28o 



Me;noir Sears Foundation for Marine Research 




Figure 6o. Raja texana, male, 468 mm long, from Barataria, Louisiana (U. S. Nat. Mus., No. 127294) and 
female, 387 mm long, from Chauvin, Louisiana (Amer. Mus. Nat. Hist., No. 16351). 



American Museum of Natural History, and Harvard Museum of Comparative Zoology, 
including a 468-mm male, close to maturity; also an embryo 87 mm long, with large 
yolk sac, probably of this species, from Cortez, Florida. Other specimens from Galve- 
ston, Aransas Pass, and Corpus Christi, Texas, in the U. S. National Museum were 
also examined. 

Distinctive Characters. The most striking feature of R. texana (and of R. ackleyi as 
well) is its color pattern with a single conspicuous dark-centered ocellar spot on the 



Fishes of the Western Nortfi Atlantic 281 

inner part of each pectoral. This in itself is enough to distinguish it (and ackleyi) from 
all other Skates of the western North Atlantic, except for specimens oi R. ocellata which 
are marked with one large ocellar spot on each side (p. 246). And the disc and tail of 
R. ocellata are so much thornier than those of R. texana, and its anterior contour is so 
much more obtusely rounded, that the two could hardly be confused. The only Skate 
of the western South Atlantic that closely parallels R. texana (and R. ackleyi) in color 
pattern is R. cyclophora Regan 19031*1 from Brazil, but the outer corners of the pectorals 
of R. cyclophora are angular as contrasted with the rounded disc of R. texana. R. texana 
is distinctive also in that the thorns on the mid-dorsal row are much smaller i*^ between 
the pectoral and pelvic arches than they are forward to the nose and rearward along the 
tail ; and, while the inner margins of the orbits are thorny, there are no thorns on the 
shoulder regions. R. texana is closely allied to R. ackleyi (p. 155), but it appears to be 
distinguishable from the latter by its relatively wider disc with abruptly rounded outer 
corners (broadly rounded on R. ackleyi), by its color, the upper surface of its disc lacking 
the small light and dark spots that mark R. ackleyi, and by the shape of the ocellar spots, 
which are round in R. texana but oval in R. ackleyi. 

Description. Proportional dimensions in per cent of total length. Male, 293 mm, 
from Freeport, Texas (Harv. Mus. Comp. Zool., No. 35845). Female, 387 mm, from 
near Chauvin, Louisiana (Amer. Mus. Nat. Hist., No. 16351). 

Disc: extreme breadth 62.3, 64.2; length 46.1, 50.8. 

Snout length: in front of orbits 13.5, 14.5; in front of mouth 15.0, 15.5. 

Orbits: horizontal diameter 3.1, 3.4; distance between 4.8, 4.2. 

Spiracles: length 2.6, 2.3; distance between 6. 2, 6.1. 

Mouth: breadth 7.5, 7.5. 

Nostrils: distance between inner ends 8.0, 7.7. 

Gill openings: lengths, ist 1.2, 1.4; 3rd 1.5, 1.7; 5th 1.2, 1.2; distance between 
inner ends, ist 14.7, 14.5; 5th 8.5, 8.0. 

First dorsal fin: vertical height 2.0, 1.8; length of base 5.8, 5.7. 

Second dorsal fin: vertical height 1.7, 1.6; length of base 6.1, 5.2. 

Pelvics: anterior margin 9.6, 10.3. 

Distance: from tip of snout to center of cloaca 44.7, 46.5 ; from center of cloaca 
to 1st dorsal 37.5, 34.4; to tip of tail 55.3, 53-5; from rear end of 2nd 
dorsal base to tip of tail 3.8, 4.7. 

Interspace between: ist and 2nd dorsals 2.2, 3.6. 

Disc about 1.2-1.3 times as broad as long; maximum anterior angle in front of 
spiracles about 106° in smaller specimens, about 95° in half-grown, about 90° in adult 
males, and about 85° in adult females; snout rather noticeably projecting, rounded at 
tip; anterior margins weakly concave just posterior to tip of snout, then bulging slightly 
and becoming weakly concave again opposite spiracles in both sexes; outer corners 

161. See Ribeiro (Arch. Mus. nac. Rio de J., 7, 1914: pi. 16) for photograph of R. cyclophora. 

162. On some specimens the thorns in the midrow may be absent for a short space about in hne with the two ocellar 
spots. 



282 



Memoir Sears Foundation for Marine Research 



rather abrupt, posterior corners broadly rounded, posterior margins weakly convex. 
Axis of greatest breadth about 70 "/o of distance back from snout toward axils of pec- 
torals. Tail with lateral folds extending nearly its entire length but so narrow that they 
are likely to be overlooked; distance from center of cloaca to origin of first dorsal 70- 
80 "/o as great as distance from snout to cloaca, the variation depending upon the 
position of first dorsal relative to second dorsal rather than on size or sex; extreme 
length of tail, from center of cloaca to tip, about 1.1-1.2 times as great as distance 
from center of cloaca to snout. 




Figure 61. Raja texana. A Lower side of anterior part of head of male pictured in Fig. 60. B Ventral view of 
pelvics of same. C Margin of right-hand nasal curtain of same, about 4X. -iO Left-hand nostril of same, about 
\y^.E Side view of posterior part of tail of female pictured in Fig. 60. F Upper teeth of another female, 550 mm 
long, from Galveston, Texas (Harv. Mus. Comp. Zool., No. 36389), about 7 X . G Upper teeth of male, 41 8 mm 
long, from Freeport, Texas (Harv. Mus. Comp. Zool., No. 36393), about 7X. 



Small specimens, up to 1 50-200 mm long, with rostral ridge either smooth or 
bearing 2-3 small thorns; 2-3 larger thorns in front of orbit and one close behind it; 
usually three in midline of back from nuchal region to shoulder girdle, these followed 
by a gap and then by a continuous row of 16-17 thorns from pelvic girdle to first 
dorsal fin; 1-3 thorns between first and second dorsals; also one irregular row low 
down either side of tail from a little posterior to axils of pectorals nearly to tip ; first 
and second dorsals with a few prickles; upper surface smooth otherwise except for 
thorns. Half-grown specimens and larger with the midrow of thorns continuous from 
nuchal region rearward to first dorsal, about eight large ones anterior to pectoral girdle, 
several small ones between pectoral girdle and pelvic girdle, 42-47 larger ones on 
tail, and 1-3 up to 6-7 smaller ones in interspace between first and second dorsals; 



Fishes of the Western North Atlantic 283 

also two rather regular rows of thorns a little larger than those of the median row"* 
on each side of tail and extending nearly to tip; 9—10 thorns around orbit, and a patch 
of smaller thorns inward from spiracle. Tip of snout and rostral ridge, space between 
spiracles, midbelt of disc back to pelvic girdle, and outer margins abreast of eyes 
(smooth on small specimens) developing a few scattered prickles with growth. Maturing 
males developing 12—13 large thorns near margins of disc opposite eyes but losing the 
prickles from midbelt of disc. Alar spines of mature males in several irregular rows, 
covering a roughly triangular area that extends forward along margins of pectorals past 
level of posterior edges of spiracles, the innermost row longest. Lower surface entirely 
smooth on small specimens; prickles, interspersed with small thorns, developing on 
tip of snout with growth; also, prickles developing along outer anterior margins of 
disc finally roughen entire lower surface of head rearward to about the mouth on males 
and along the median belt about as far as fifth gill openings in females; lower side of 
extremity of tail prickly on females but smooth on males. 

Snout in front of orbits about 3.3—4.0 times as long as orbit in young of both 
sexes and in males to maturity, about 4.5—5.0 times in mature females; its length in 
front of mouth about 1.6— 1.8 times as long as distance between nostrils in young and 
in mature males, about 2.0—2.2 times in large females. Distance between first gill 
openings 1.8—2.0 times as long as distance between exposed nostrils, about equal 
between fifth gill openings; first gill openings i. 0—1.2 times as long as fifth and about 
16-27 "/o as long as breadth of mouth. Nasal curtain and expanded posterior (outer) 
margin of nostril each with shallow fringe. Mouth nearly straight in females, lower 
jaw a little more arched in mature males. 

Teeth |^8' those of small and half-grown specimens close-set in quincunx, with 
nearly circular base and stout conical cusp on all except those of the oldest row; those 
of adults of both sexes spaced more loosely in transverse series with well developed 
cusp; uppers of males a little more slender than those of females, the lowers con- 
spicuously so. 

Dorsals similar in size and shape, posterior margin more or less re-entrant. Inter- 
space between first and second dorsals varying from only about 30 "/o as long, to about 
as long, as base of first dorsal. Caudal membrane posterior to second dorsal about as 
long as base of first dorsal. Pelvics moderately concave outwardly, strongly scalloped 
around the indentation; anterior margin about 67—72 "/o as long as distance from its 
own origin to rear tip of pelvic; anterior lobe noticeably small but including three or 
four slender radials besides the first stout one; posterior lobe with weakly convex 
outer margin; rear tip broadly rounded in female but narrowly so in mature male, 
extending back about 10—16 "/o of distance from axils of pelvics toward tip of tail in 
young and adult female, about 20 % iri adult male. Claspers of sexually mature male 
reaching back a little more than halfway from axils of pelvics toward first dorsal. 

Anterior rays of pectorals reaching about 51— 56*/o of distance from level of 
fronts of orbits toward tip of snout. 

163. This contrast in size is the reverse of what is usual among Skates of the western North Atlantic. 



284 Memoir Sears Foundation for Marine Research 

Color. Upper surface rich chocolate or coffee brown except for a translucent area 
on either side of rostral cartilage, with indistinct pale spots and blotches irregularly- 
distributed, most conspicuous on small specimens but perhaps entirely lost on some 
adults; also a single conspicuous dark brown or black ocellar spot bordered with yellow 
on inner part of each pectoral a little posterior to axis of greatest breadth. Lower surface 
plain white. 

Size. A male 468 mm long is close to sexual maturity, as indicated by the sexual 
spines and claspers. The largest specimen is 537 mm long (see Study Material, p. 280). 

Habits. Nothing whatever is known of the habits of this species, except that most 
of them have been taken from shallow water, the deepest being 37 fathoms for a new- 
born specimen trawled off Aransas Pass, Texas. 

Range. Known only from the west coast of Florida (Cortez and Englewood), from 
the coast of Mississippi, where it is reported to us as fairly common ;i^^ from Louisiana 
(Barataria, Chauvin); and from Texas (Aransas Pass, Corpus Christi, Austin, Galveston, 
Houston); hence it is probably restricted to the Gulf of Mexico. 

Synonyms and References: 

Rata texana Chandler, Proc. U. S. nat. Mus., ^g, 1921: 657 (descr., Houston, Texas); Baughman, Copeia, 

1943: 133 {R. texana Chandler 1921 antedates R. texana Leriche 1940, a fossil; stellata proposed as 

substitute for latter). 
Raja texana Springer, Copeia, 1939: 237 (descr. egg case, Englewood, Florida); Woods, Copeia, 1942: 191 

(meas., near Corpus Christi, Texas); Gunter, Publ. Inst. mar. Sci. Texas, I, 1945: 21 (size, temp., sal., 

Texas). 

Genus Breviraja Bigelow and Schroeder 1948 

Breviraja Bigelow and Schroeder, J. Mar. Res., 7 (3), 1948: 548, 558; type species, B.coksi Bigelow and 
Schroeder 1948. 

Generic Synonyms: 

Raja (in part) Garman, Bull. Mus. comp. Zool. Harv., 8, 1881: 236; Mem. Harv. Mus. comp. Zool., 36, 

191 3: 335; for R.plutonia Garman 1881. 
Malacorhtna (in part) Garman, Bull. Mus. comp. Zool. Harv., 8, 1881: 237 (suggests R.plutonia may be a 

Malacorhina because of short rostral cartilage.) 

Generic Characters. Pelvics not completely divided, although their margins may 
be so deeply concave that the fin is definitely bilobed. Tip of rostral projection of 
cranium falling short of level of anterior extremities of pectoral rays and falling shorter 
still of tip of snout; anterior rays of pectorals reaching nearly to level of tip of snout. 
Characters otherwise as in Raja. 

Size. Some of the species are small, males maturing at lengths no greater than 
200-300 mm, as indicated by the condition of the claspers, but others grow larger. 

Developmental Stages. Presumably oviparous, but the eggs have not been seen. 

Habits and Range. They appear to be confined to moderately deep water, with 
the recorded depths of capture ranging from 200 to 500 fathoms. Beyond this nothing 

164. Personal communication from Stewart Springer. 



Fis/ies of the Western North Atlantic 



285 









Figure 6z. X-ray photographs, slightly retouched, to show rostral cartilage and foward extension of pectoral 
radials in Breviraja: Upper left, B atripinna ; upper right, B.co/esi; middle left, B.cuiensis; middle right, 
B.flutonia; lower left, B.sfinosa; lower right, B. sinus-mexicanus. 



2 86 Memoir Sears Foundation for Marine Research 

is known of their habits. So far the genus is known only along the north and south 
coasts of Cuba, in the northern part of the Gulf of Mexico off Pensacola, Florida, and 
northward along the continental slope of the southeastern United States to the offing 
of South Carolina, the most northerly record (for a specimen of B.plutonia) being 
Lat. 32°43' N, Long. 77°2i' W. But it may prove that some short-nosed Skates from 
other parts of the world that are now referred to Raja actually belong in this genus. 
Species. Seven species of this genus have been discovered, six of them recently; 
the seventh (j>lutonia Garman 1881) was usually referred to Raja until Breviraja was 
recently separated from it, though the suggestion had been made in the original account 
of the species that the rostral cartilage might remain undeveloped. For a Key to Species, 
see p. 147.^ 

Breviraja atripinna Bigelow and Schroeder 1950 

Black-fin Skate 

Figures 62 (upper left), 63 

Study Material. One male and two females, 187-278 mm long, trawled by the 
research vessel Atlantis off Santa Clara Province on the north central coast of Cuba 
in 250-500 fathoms, in the Harvard Museum of Comparative Zoology.- 

Distinctive Characters. B. atripinna falls with B. sinus-mexicanus and with the B. 
cui>ensis-a.nd-p/utoma group in that its tail is of a great length. But it is separable from 
all three of these species by its relatively longer rostral cartilage. Halfgrown and larger 
specimens also differ conspicuously from B. sinus-mexicanus in having a relatively much 
longer interspace between the dorsal fins. Very young B. atripinna more nearly re- 
semble B. sinus-mexicanus in this respect, but the tail of B. atripinna is so much less 
thorny than that of the latter (cf. Fig. 63 with 69, 70) that there is little danger of 
confusing the two. The plain coloration of its disc, without dark markings, is a con- 
venient field mark for distinguishing atripinna from cuhensis and from plutonia. 

Description. Proportional dimensions in per cent of total length. Male, 225 mm, 
and female, 278 mm, from Cuba (Harv. Mus. Comp. Zool., Nos. 36367 and 36370, 
respectively). 

Disc: extreme breadth 44.5, 45-3; length 37.3, 38.5. 

Snout length: in front of orbits 8.9, 9.7; in front of mouth 12.0, 12.6. 

Orbits: horizontal diameter 4.0, 5.0; distance between 2.7, 2.7. 

Spiracles: length 2.2, 1.9; distance between 6.0, 6.1. 

Mouth: breadth 5.3, 5.0. 

Nostrils: distance between inner ends 5.8, 5.9. 

Gill openings : lengths, ist 1.3, 1.3; 3rd 1.3, 1.3; 5th 1.2, 1.2; distance between 
inner ends, ist 9.8, 10. o; 5th 5.8, 6.1. 

First dorsal fn: vertical height 1.5, 1.3; length of base 4.2, 4.7. 

I. Bre-viraja combined with Raja. 

z. Atlantis Sts. 2985, 3443, 3459 (Harv. Mus. Comp. Zool., Nos. 36367, 36370 type, 36371). 




Figure 63. Breviraja atripinna, female, 278 mm long, from off Santa Clara Province, north coast of Cuba, 
Lat. 23°22' N, Long. 79°53' W, Atlantis St. 3443 (Harv. Mus. Comp. ZooL, No. 36370, ty-pe). A Ventral 
view of pelvics. B Lower surface of anterior part of head to show nostril and mouth. C Side view of posterior 
part of tail, about 2.1 X. D Right-hand nostril, about 7.9 X. £ Margin of left-hand nasal curtain, about 5.2 X. 
F Upper teeth. 



2 88 Memoir Sears Foundation for Marine Research 

Second dorsal fin: vertical height 1.5, 1.4; length of base 4.0, 3.6. 

Pelvics: anterior margin 10.7, 11.8. 

Distance: from tip of snout to center of cloaca 35.6, 35.7; from center of cloaca 

to 1st dorsal 48.0, 47.0; to tip of tail 64.4, 64.3. 

Interspace between: ist and 2nd dorsals 4.5, 5.0; 2nd dorsal and tip of tail 3.5, 4.0. 

Disc about 1.2 times as broad as long, snout projecting only slightly on newborn 
specimens but more conspicuously on larger ones, tip narrowly blunted; maximum 
anterior angle in front of spiracles about 1 15° on small specimens, about 100° on larger; 
anterior margins weakly convex anterior to orbits, thence nearly straight to the broadly 
rounded outer corners; posterior margins evenly and moderately convex, posterior 
corners broadly rounded, inner margins straight toward axil. Axis of greatest breadth 
about 75 "/o of distance back from tip of snout toward axils of pectorals. Tail slender, its 
lateral folds narrow and confined to posterior two-thirds, widening rearward to opposite 
second dorsal; its length to first dorsal about 1.4 times as great as distance from center 
of cloaca to tip of snout and about 1.8 times as great to tip in large specimens; a little 
longer relatively in smaller specimens. 

Upper surface around posterior margins of pectorals and on tip of snout smooth, 
disc and tail rough elsewhere with small and sharp close-set prickles, many of them 
covered over with pigmented skin; inner margin of orbit to inner end of spiracle with 
a single series of small thorns, 7—8 on small and 12—15 ^^ older specimens; 2—3 thorns 
inward from each spiracle; two on each shoulder of young increasing to 2,—^ on older 
ones; midline of back with a row of thorns — about six from nuchal region to pectoral 
girdle, then a gap followed by about 60 smaller thorns along posterior part of disc 
and on tail to first dorsal fin; thorns in midline paired here and there side by side and 
more widely spaced rearward, decreasing in regularity with growth; an additional 
irregular band of small thorns on either side of posterior part of mid-dorsal ridge from 
about opposite axils of pectorals rearward, losing its identity among prickles of tail; 
also a row of small thorns along lower edge of tail on either side to a little in advance 
of first dorsal; anterior two-thirds of dorsals as well as interspace between them prickly, 
but caudal membrane smooth; pelvics prickly except at margins. Alar spines of adult 
male not seen. Lower surface of disc and pelvics smooth; tail also smooth below on 
very young specimens but more or less roughened with prickles on larger specimens. 

Snout in front of orbits about three times as long as orbit on young, about two 
times on older specimens; its length in front of mouth 2.0—2.3 times as great as distance 
between exposed nostrils. Orbit about 1.6—2.6 times as long as spiracle; about 1.2— 
1.5 times as long as distance between orbits on young specimens, about two times on 
largest specimen seen. Distance between first gill openings 1.7 times as long as distance 
between exposed nostrils, between fifth gill openings i.o times; first gill openings 
about as long as fifth and about 25 "/o as long as breadth of mouth. Nasal curtain smooth- 
edged, posterior (outer) margin of nostril with a few marginal lobelets about midway 
of its length. Mouth slightly arched centrally. 



Fishes of the lVester?2 North Atlantic 289 

Teeth about 4^; close-set in transverse series in both sexes, with low conical 
cusp on females and immature males. 

First and second dorsals similar in shape, the base' of first about i.i — 1.3 times 
as long as base of second. Interspace between first and second dorsals increasing in 
relative length from about 40 % ^s long as base of first dorsal on very young specimens 
to about as long as base of first dorsal on large. Caudal membrane about as long as base 
of second dorsal. Pelvics deeply concave outwardly and scalloped around the inden- 
tation; anterior margin slightly shorter to slightly longer than distance from its own 
origin to rear tip of pelvic; anterior lobe narrowing to a pointed tip, with 2-3 radials 
besides the first and stoutest; outer margin of posterior lobe rather strongly convex 
and weakly scalloped, rear tip well rounded, reaching back about Vi the distance from 
level of axils of pectorals toward first dorsal. Claspers of mature males not seen. 

Rostral cartilage narrow, extending about 75 "/o of distance from front of cranium 
toward tip of snout; anterior rays of pectorals about 85—90 "/o of that distance. 

Color. Upper surface of disc, pelvics, and tail pale pinkish brown without dis- 
tinctive markings; dorsals and caudal membrane pale and translucent on newborn 
specimens but soon turning brownish black and thus becoming conspicuous; lateral 
folds on tail transparent anteriorly but brownish black posteriorly to second dorsal. 
Lower surface plain whitish on small specimens, later becoming clouded with chocolate 
brown; darkest on abdomen and anterior lobes of pelvics while remaining whitish on 
median region of head from snout to first gill openings; anterior part of tail opposite 
pelvics with a chocolate blotch on large specimens. 

Size. The claspers of a male 225 mm long still fall short of the tips of the pelvics, 
making it likely that this Skate does not mature until larger than other known species 
of its genus; the size at maturity is not known. 

Developmental Stages. The eggs have not been seen. 

Habits. The facts that the three specimens seen so far were trawled by the Atlantis 
at 250—500 fathoms and that none were taken in the many shoaler hauls around Cuba 
show that it is a deep-water species. 

Range. Known only off the north central coast of Cuba. 

Reference : 

Breviraja atrtpinna Bigelow and Schroeder, Bull. Mus. comp. Zool. Harv., JOJ (7), 1950: 390 (descr., ills., 
depth, ofF N. Central Cuba). 



Breviraja colesi Bigelow and Schroeder 1948 
Figures 62 (upper right), 64, 65 

Study Material. Four males and four females, 81— 333 mm long, including the 
type,^ trawled by the research vessel Atlantis off the north and south coasts of Cuba, 
in the Harvard Museum of Comparative Zoology. 

3. Type specimen, a female 333 mm long (Harv. Mus. Comp. Zool., No. 36374). 

'9 



290 Memoir Sears Foundation for Marine Research 

Distinctive Characters. B. colesi is marked off from all other known members of its 
genus in the western Atlantic by a tail that is at least no longer from the axils of the 
pelvics to the origin of the second dorsal than from the axils of the pelvics to the tip 
of the snout, by the fact that the midzone of the disc between the pectoral and pelvic 
girdles bears only one or two conspicuous thorns, if any, and by the dark markings on 
its upper surface. The shortness of its rostral cartilage separates it from the hard-nosed 
Skates of the genus Raja. 

Description. Proportional dimensions in per cent of total length. Male, 294 mm 
(Harv. Mus. Comp. Zool., No. 36497), female, 333 mm (Harv. Mus. Comp. Zool., 
type, No. 36374), from Cuba. 

Disc: extreme breadth 49.3, 48.0; length 41.5, 45.9. 

Snout length: in front of orbits 8.3, 9.8; in front of mouth 8.8, 1 1.2. 

Orbits: horizontal diameter 4.8, 4.5; distance between 2.5, 3.0. 

Spiracles: length 2.9, 2.8; distance between 5.8, 5.7. 

Mouth: breadth 7.1, 6.3. 

Nostrils: distance between inner ends 6.^^ 6.6. 

Gill openings : lengths, ist 1.4, 1.8; 3rd 1.4, 1.8; 5th 1.3, 1.5; distance between 
inner ends, ist 12.2, 13.6; 5th 5.8, 6.6. 

First dorsal fin: vertical height 1.4, 1.5; length of base 4.8, 3.6. 

Second dorsal fin: vertical height i.o, 1.5; length of base 4.1, ^t-^)- 

Pelvics: anterior margin 10.5, 10.2. 

Distance: from tip of snout to center of cloaca 38.4, 41.4; from center of cloaca 
to 1st dorsal 51.0, 50.0; to tip of tail 61.6, 58.6; from rear end of 2nd 
dorsal base to tip of caudal 1.4, 1.5. 

Interspace between: ist and 2nd dorsals 0.3, 0.0. 

Disc about i. 0—1.2 times as broad as long, very obtuse in front; tip of snout 
marked by a small rounded prominence; maximum anterior angle in front of spiracles 
about 135°; anterior margins moderately and evenly convex on young and on females 
to maturity but weakly concave opposite and posterior to spiracles on adult males; 
outer corners well rounded, more broadly so on adult males than on females; posterior 
corners broadly rounded, merging into convex inner margins. Axis of greatest breadth 
about 70 "/o of distance back from tip of snout toward axils of pectorals. Tail with 
narrow lateral folds for entire length from a little behind tips of pelvics, widening 
posteriorly; its length from center of cloaca to origin of first dorsal about 1.2 — 1.3 times 
as great as distance from center of cloaca to snout, about 1.3— 1.5 times as great to 
origin of second dorsal, and about 1.4— 1.6 times as great to tip. 

Disc rough with small prickles on adults of both sexes except close along anterior 
and posterior edges; a narrow band of thorns along anterior margins from level of 
spiracles forward, very small on females, larger on adult males ; a cluster of small thorns 
on rostral ridge about halfway between level of fronts of orbits and tip of snout; lo-i i 
thorns around inner margin of orbit, the row interrupted midway; two thorns inward 



Fishes of the JVestern North Atlantic 



291 




Figure 64. Breviraja colesi, male, 320 mm long, from off Santa Clara Province, north coast of Cuba, Lat.22°5o'N, 
Long. 79°o8' W, Atlantis St. 3423, 245 fathoms (Harv. Mus. Comp. Zool., No. 36447) and female, 333 mm 
long, from same general region, Lat. 23°i2' N, Long. 8i°23' W, Atlantis St. 3483, 285 fathoms (Harv. Mus. 
Comp. Zool., No. 36374, type). 



292 Memoir Sears Foundation for Marine Research 

from inner end of each spiracle; 4-5 on each shoulder; a median row of about six 
larger thorns from nuchal region to pectoral girdle, followed by 5-6 smaller ones more 
widely spaced to level of axils of pectorals, then by about 40—44 larger ones more closely 
spaced along tail; also, each side of tail with 2—3 irregular rows, the thorns successively 
a little larger and more closely spaced rearward to first dorsal, followed by smaller 




Figure 65. Breviraja colesi. A Mouth and nostrils of female illustrated in Fig. 64. B Margin of right-hand nasal 
curtain of same, about 5.6 X. C Left-hand nostril of same, about 5.6 X. D Upper teeth of same, near center of 
jaw, about 17X. £ Upper teeth, near center of jaw, of male illustrated in Fig. 64, about 17X. f Dorsal view 
of tip of clasper of another mature male, from off north coast of Cuba, about 0.7 X- 



ones below dorsals; pelvics of females with a few small prickles on central part of 
posterior lobe, those of adult males smooth; dorsals prickly along upper parts; caudal 
membrane sparsely prickled. Very small specimens with fewer thorns around orbits, 
only 1—2 on each shoulder, 3-4 in median row from nuchal region to pectoral girdle; 
thorns on tail in more regular rows than on adults. Alar spines of sexually mature males 
in 2—3 rows near outer corners of pectorals. Lower surface smooth. 

Snout in front of orbits about 1.7-2.2 times as long as orbit, its length in front 
of mouth about 1.4- 1.7 times as great as distance between exposed nostrils. Orbit 
about 1.5— 1.9 times as long as distance between orbits and about 1.6 times as long 
as spiracle. Distance between first gill openings 1.9— 2.1 times as long as distance be- 
tween exposed nostrils, i.o-i.i times between fifth gill openings; first gill openings 
about as long as fifth and about 20-30 "/o as long as breadth of mouth. Nasal curtain 



Fishes of the Western North Atlantic 293 

deeply fringed, posterior (outer) margin of nostril finely so. Mouth moderately 
arched centrally. 

Teeth about 42^50, close-set in quincunx in females to maturity, with low con- 
ical cusp on ovate base; mature males with longer and sharper cusp. 

First and second dorsals similar in size and shape, confluent at base or with 
short interspace; caudal membrane posterior to second dorsal only about Vs as long as 
base of second dorsal. Pelvics deeply and sharply concave outwardly, weakly scalloped; 
anterior margin about 57—62 "/o ^s long as distance from its own origin to rear tip of 
pelvic; anterior lobe with narrowly rounded tip, including only two radials besides the 
stoutest; posterior lobe moderately convex on females, less so on adult males, its rear 
corner narrowly rounded, reaching back about Vs the distance from axils of pectorals 
toward first dorsal. Claspers of sexually mature males reaching back about 1/3 the 
distance from axils of pelvics toward first dorsal, their tips with blade-like expansion 
and complex arrangement of hooked processes. 

Rostral projection from cranium triangular, forming an angle of about 40°, ex- 
tending rigidly for about 60-75 "/o of the distance from front of cranium toward tip 
of snout. 

Color. Upper surface of disc pale brown, mottled with irregular spots and blotches 
of darker brown, more numerous on some specimens than on others; outer part of 
disc with a variable number of larger rounded spots with pale centers and brown mar- 
gins; also one on center of posterior lobe of each pelvic; tail with various brown mark- 
ings, some forming crossbars that may number 7—8, more definitely outlined on young 
specimens than on adults, its lateral folds whitish; upper anterior part of each dorsal 
with a bronze blotch; another at anterior end of base of first dorsal on some specimens, 
not on all. Lower surface of disc and of tail pale yellowish or whitish. 

Size. Apparently this species does not reach a length of much more than 400 mm, 
for the claspers of a male 320 mm long (Fig. 64) appear to be fully formed. 

Developmental Stages. Nothing is known except that a young specimen with disc 
only 43 mm wide bears a close resemblance to the adults. 

Habits. All specimens taken have been trawled in depths ranging from 200—285 
fathoms. 

Range. Known only from Cochinas Bay on the south coast of Cuba and off Matan- 
zas and Santa Clara provinces on the north coast. 

Reference : 

Breviraja colesi Bigelow and Schroeder, J. Mar. Res., 7 (3), 1948 : 559 (descr., ills., ofFN. and S. coasts of Cuba). 



Breviraja cubensis Bigelow and Schroeder 1950 
Figures 62 (middle left), 66 

Study Material. Seventy specimens, male and female, 68—221 mm long, includ- 
ing the type, trawled by the research vessel Atlantis off North Central Cuba at 235- 



2 94 Memoir Sears Foundation for Marine Research 

405 fathoms,* in the Harvard Museum of Comparative Zoology; two specimens 
trawled off St. Augustine, Florida, at 373 fathoms,^ in U. S. National Museum. 

Distinctive Characters. B. cuhensis falls with B. sinus-mexicanus, B. atripinna, and 
B. plutonia because of the great length of its tail. But it is easily separable from B. sinus- 
mexicanus by the facts that its tail posterior to the tips of the pelvics is not conspicuously 
thorny, that it has only prickles and no thorns along its rostral ridge, and that the 
prickles on its disc and tail are not densely pigmented. The dark markings on its disc 
and the dark bars on its tail serve to distinguish it from B. atrifinna, which is plain- 
colored. It resembles B. plutonia closely in general form, and some specimens have 
much the same color pattern ; but the anterior lobes of the pelvics are relatively smaller 
in B. cubensis than in B. plutonia (see Key, p. 148), the thorns on its disc are relatively 
smaller, and usually there are only 1—2 thorns on either shoulder; the first and second 
dorsals (confluent in B. plutonia) are usually separated by a definite interspace in B. 
cubensis \ the teeth average a little more numerous in B. cubensis than in B. plutonia, 
and the rostral cartilage is conspicuously longer, though intermediates may occur in 
this respect. 

Description. Proportional dimensions in per cent of total length. Female, 203 mm 
(Harv. Mus. Comp. Zool., No. 36445), and male, 210 mm (Harv. Mus. Comp. Zool., 
type. No. 36443), from Cuba. 

Disc: extreme breadth 46.2, 49.7; length 37.1, 39-0. 

Snout length: in front of orbits 8.4, lo.o; in front of mouth 10.8, ii.o. 

Orbits: horizontal diameter 4.4, 4.8; distance between 2.8, 2.9. 

Spiracles: length 2.2, 2.4; distance between 6.7, 7.1. 

Mouth: breadth 5.7, 6.2. 

Nostrils: distance between inner ends 4.4, 5.0. 

Gill openings: lengths, ist i.o, 1.2; 3rd i.o, 1.2; 5th 0.7, i.o; distance between 
inner ends, ist 10.3, ii.o; 5th 6.9, 6.7. 

First dorsal fin: vertical height 1.0; 1.2; length of base 4.9, 5.2. 

Second dorsal fin: vertical height 1.0, 1.7; length of base 4.2, 3.8. 

P^/wVj; anterior margin 10.8, 11.2. 

Distance: from tip of snout to center of cloaca 35.0, 38.1 ; from center of cloaca 
to ist dorsal 50.8, 50.0; to tip of tail 65.0, 61.9; from rear end of 2nd 
dorsal base to tip of caudal 2.5, 1.4. 

Interspace between: ist and 2nd dorsals 3.0, 1.4. 

Disc about 1.2-1.3 times as broad as long, obtusely rounded in front; tip of snout 
marked by a low projection; maximum anterior angle in front of spiracles 115—130°; 
anterior margin of disc rather strongly convex anterior to level of orbits and concave 
opposite spiracles in most cases, more strongly so in males but more nearly straight 
in some females; outer and posterior corners broadly rounded, posterior and inner 

4. Atlantis Sts. 2961, 2983, 2984, 2986, 2987, 2988, 2999, 3432, 3438, 3449, 3451, 3457, 3483> ^nd 34^5; fo'' P''^- 
cise localities, see Chace (Contrib. No. 274, Woods Hole oceanogr. Inst., 1940). 

5. Albatross Sta. 2664. 



Fishes of the Western North Atlantic 



295 




Figure 66. Breviraja cubensts. A Male, 210 mm long, from off Santa Clara Province, north coast of Cuba, 
Lat. 23°5o' N, Long. 79°5i' W, Atlantis St. 3451, 405 fathoms (Harv. Mus. Comp. Zool., No. 36443, type). 
B Lower surface of anterior part of head of same, to show mouth and nostrils. C Nostril of same, about 6.3 X. 
D Upper teeth of same from near center of jaw, about 1 3 X . £ Female, 202 mm long, from off Santa Clara 
Province, north coast of Cuba, Lat. 23°i i' N, Long. 79°o8' W, 235-260 fathoms, Atlantis St. 2983 (Harv. 
Mus. Comp. Zool., No. 36364). F Upper teeth of same from near center of jaw, about 1 3 X . 



296 Memoir Sears Foundation for Marine Research 

margins both strongly convex. Axis of greatest breadth about 75 "/o of distance back 
from snout toward axils of pectorals. Tail slender, its lateral folds narrow, confined 
to posterior two-thirds, broadening a little rearward; its length from center of cloaca 
to first dorsal about 1.3— 1.4 times as great as distance from center of cloaca to snout, 
about 1.6— 1.8 times as great to tip. 

Upper surface of disc (except for tip of snout) and tail rough with small prickles 
on half-grown specimens and on females to maturity, but mature males smoother along 
a narrow marginal band and over areas of varying extent on inner parts of pectorals; 
both sexes with 9—15 small thorns around inner margin of orbit, the series sometimes 
interrupted midway; 1—2 small thorns inward from inner end of spiracle, and 1—2 on 
each shoulder close to midline; also a median row of 50—100 or more thorns of various 
sizes grading down to prickles from nuchal region to first dorsal fin, the row con- 
tinuous on some specimens but more or less interrupted on others between pectoral 
and pelvic girdles and much more prominent on some than on others; tail posterior to 
axils of pelvics with 1—2 additional rows of small thorns or large prickles along either 
side; anterior parts of dorsals and interspace between first and second dorsals prickly; 
pelvics either smooth or more or less prickly along inner parts. Sexually mature males 
with a patch of small thorns on outer part of disc opposite orbits, their alar spines in 
2—4 rows opposite outer corners of pectorals, 3—6 spines in each row. Lower surface 
smooth. 

Snout in front of orbits 1.6— 2.1 times as long as orbit, its length in front of mouth 
2.1—2.6 times as great as distance between exposed nostrils. Orbit about 1.5 times as 
long as distance between orbits and about two times as long as spiracle. Distance 
between first gill openings 2.1—2.7 times as long as distance between exposed nostrils, 
1.3— 1.6 times between fifth gill openings; first gill openings i.i— 1.3 times as long as 
fifth and about 18—26 "/o ^s long as breadth of mouth. Nasal curtain and posterior 
(outer) margin of nostril both smooth-edged. Mouth weakly arched centrally. 

Teeth jgZ^g; close-set in quincunx, with obscure transverse cutting edge and 
faintly indicated cusp in young and in females to maturity, but with long and sharp 
recurved cusp in mature males. 

Dorsals similar in shape, the base of the first equal to or slightly longer than that 
of the second, usually separated by an interspace 0.25—1.0 times as long as base of first 
dorsal but sometimes confluent. Caudal membrane posterior to second dorsal about half 
as long as base of second dorsal. Pelvics deeply concave outwardly in subtriangular con- 
tour, scalloped around the indentation; anterior margin about 90— 100 "/o as long as 
distance from its own origin to rear tip of pelvic; anterior lobe slender, tapering to a 
narrowly-pointed tip, of two slender radials in addition to the first stout one; posterior 
lobe with moderately convex and more or less wavy outer margin, extending nearly 1/4 
the distance rearward from level of axils of pectorals toward first dorsal fin, the rear 
tip narrowly rounded on young of both sexes and on adult females but more pointed 
on adult males. Claspers of sexually mature males slender, reaching about Va the distance 
from axils of pelvics toward first dorsal, their tips simple. 



Fishes of the Western North Atlantic 297 

Rostral cartilage triangular, extending about 70—75 "/q of the distance from front 
of cranium toward tip of snout. 

Color. Upper surface of disc pale brown, with darker brown spots and blotches 
vaguely outlined and varying widely in size, number and distribution, most conspicuous 
on small specimens; very young ones usually with a dark curved bar extending from 
behind orbit to shoulder region and with a narrow crescentic darker band (concavity 
forward) crossing base of tail; outer posterior part of each pectoral of some adults with 
one dark blotch much larger than the others; anterior lobes of pelvics whitish above, 
the posterior lobe of same brownish hue as disc, sometimes with a few irregular dark 
spots; tail pale brown, usually with 5—6 crossbars vaguely outlined, more conspicuous 
on large specimens than on small, the two more posterior ones crossing the two dorsal 
fins; anterior part of caudal membrane blackish. Lower surface pale yellowish without 
dark markings. 

Size. The smallest specimens seen (about 68 mm long) apparently were newly 
hatched. Males mature (as shown by their claspers) at a length of 180-210 mm. This, 
with the fact that the largest specimen (a female) among 70 was only 221 mm long, 
indicates that this Skate does not grow to a length greater than about 300 mm. 

Developmental Stages. Egg cases have not been seen. Newly hatched specimens 
closely resemble their parents. 

Habits. We know only that this is a deep-water species, so far recorded only from 
235-405 fathoms. 

Range. Known only off the north central coast of Cuba. 

Reference : 

Breviraja cubensis Bigelow and Schroeder, Bull. Mus. comp. Zool. Harv., JOJ (7), 1950: 394 (descr., ills., 
depth, off N. Central Cuba). 

Breviraja plutonia (Garman) 1 8 8 i 
Figures 62 (middle right), 67, 68 

Study Material. Forty-two specimens, males and females, 132—253 mm long; one 
(the type) from the offing of Savannah, Georgia, Lat. 3i°57' N, Long. 78° 19' W, in 
333 fathoms; the remainder trawled off Jacksonville, Florida, Lat. 30°2i'N, Long. 
19° SS' ^1 ^^ 230—250 fathoms, Feb. 24, 1940 (Atlantis Station 3779); all in the 
Harvard Museum of Comparative Zoology. 

Distinctive Characters. B. plutonia falls with B. sinus-mexicanus, B. atripinna and 
B. cubensis in the long-tailed division of the genus. But it is easily separable from B. sinus- 
mexicanus by the anterior contour of its disc (cf. Fig. 67 with 69, 70). It is separated 
from B. atripinna by this same feature as well as by the fact that its first and second 
dorsals are confluent, and by the variegated color pattern on the upper surface of its 
disc. It resembles B. cubensis closely in shape of disc and in dermal armature, but the 
anterior lobes of its pelvics are considerably longer relatively; the thorns on the anterior 
part of its tail and on its disc are larger and more conspicuous, usually with 2—4 on 



298 Memoir Sears Foundation for Marine Research 




.■••.^v%. 

mm 






Figure 67. Breviraja flutonia, male, 230 mm long, and female, 238 mm long, from off northern Florida, Lat. 
30°2i' N, Long. 79°5 5' W, 230-250 fathoms, Atlantis St. 3779 (Harv. Mus. Comp. Zool., No. 36493). 



Fishes of the Western North Atlantic 299 

each shoulder; its dorsal fins are confluent (usually more or less separated in cubensis)\ 
its teeth average slightly fewer than in cubensis\ and its rostral cartilage is considerably 
shorter relatively than is usual in cubensis (see Fig. 62). 

Description. Proportional dimensions in per cent of total length. Female, 216 mm, 
and male, 230 mm (Harv. Mus. Comp. Zool., No. 36493), from off Jacksonville, 
Florida. 

Disc: extreme breadth 43.5, 44.3; length 35.7, 38.2. 

Snout length: in front of orbits 7.9, 6.7; in front of mouth 8.8, 8.7. 

Orbits: horizontal diameter 4.4, 5.2; distance between 3.0, 3.5. 

Spiracles: length 2.3, 2.4; distance between 7.0, 7.0. 

Mouth: breadth 5.1, 5.6. 

Nostrils: distance between inner ends 3.9, 4.8. 

Gill openings: lengths, ist 1.3, 1.3; 3rd 1.3, 1.3; 5th 1.2, i.i ; distance between 
inner ends, ist 1 1.3, i i.i ; 5th 7.6, 5.6. 

First dorsal fin: vertical height 1.4, 1.5; length of base 3.7, 4.8. 

Second dorsal fin: vertical height 1.6, 1.4; length of base 3.5, 4.8. 

Pelvics: anterior margin 9.3, 1 1.7. 

Distance: from tip of snout to center of cloaca 33.3, 36. i ; from center of cloaca 
to ist dorsal 57.0, 51.8; to tip of tail 66.7, 63.9; from rear end of 2nd 
dorsal base to tip of caudal 2.5, 2.6. 

Interspace between: ist and 2nd dorsals 0.0, 0.0. 

Disc about 1.2 times as wide as long, broadly rounded in front; tip of snout marked 
by a low, blunt subtriangular projection; maximum anterior angle in front of spiracles 
about 130-135°; anterior margins concave abreast of spiracles, more deeply so on adult 
males than on females or on young; both outer and posterior corners broadly rounded; 
posterior margins moderately and evenly convex, inner margins weakly so. Tail with 
narrow lateral fold low down on each side along posterior two-fifths extending to tip; 
its length to first dorsal about 1.4-1.7 times as great as distance from center of cloaca 
to tip of snout and about 1.8-2.0 times as great to tip. 

Upper surface smooth on tip of snout and around rear corners of pectorals but 
prickly elsewhere on disc and on tail, most densely so above eyes; orbit with two small 
thorns on posterior margin on small specimens and up to 12 in a continuous row around 
inner edge on adults; one larger thorn inward from inner end of spiracle; 2—3 on each 
shoulder on young, 3—4 in a transverse row on adults, these being larger and more 
conspicuous on females than on mature males and preceded on some by one thorn on 
either side in advance of shoulder; midline of back with a continuous row of larger and 
smaller thorns (varying considerably in number on specimens of the same size), about 
45-70 from nuchal region back to about midlength of tail, followed rearward to first 
dorsal by thorns less regularly arranged, often interrupted and usually not in exact mid- 
line; an additional row of irregularly arranged smaller thorns on either side of the median 
row from pectoral girdle rearward along anterior part of tail, succeeded by 1—3 irregular 
rows to first dorsal and by smaller thorns or prickles to tip of tail; pelvics naked in both 



300 Memoir Sears Foundation for Marine Research 

sexes; anterior part of dorsals prickly; caudal membrane also with a few prickles. Matur- 
ing males developing additional small thorns along marginal belts abreast of eyes; their 
alar spines in 3—4 irregular rows. Lower surface smooth, except sometimes prickly along 
margins of anterior part of tail. 

Snout in front of orbit about 1.3— 1.8 times as long as orbit, its length in front of 
mouth about twice as great as distance between exposed nostrils. Orbit about 1.5 times 




Figure 68. Bremraja flutonia. A Ventral view of pelvics of female pictured in Fig. 67, about ly.. B Upper 
teeth of same, about 17 X ■ C Lower side of anterior part of head of male pictured in Fig. 60, about IX- D Right- 
hand nostril and nasal curtain of same, about 5X. £ Upper teeth of same, about 17 X. 



as long as distance between orbits and about 2.0 times as long as spiracle. Distance 
between first gill openings 2.3—2.9 times as long as distance between exposed nostrils, 
1.2 — 1.9 times between fifth gill openings; first gill openings i.i— 1.2 times as long as 
fifth and about 25 "/o as long as breadth of mouth. Nasal curtain smooth or slightly ir- 
regular; posterior (outer) margin of nostrils smooth. Mouth only slightly arched cen- 
trally. 

Teeth Ij^; those of young and of adult females close-set in quincunx, with obscure 
transverse cutting edge and faintly indicated cusp; those of mature males loosely spaced 
in transverse series, with narrow conical cusps directed outward toward corners of mouth. 

First and second dorsals similar in size and shape, conflluent at base; caudal mem- 
brane posterior to second dorsal only about half as long as base of second dorsal. Pelvics 
deeply concave outwardly; anterior margin about as long as distance from its own origin 



Fishes of the Western North Atlantic 301 

to rear tip of pelvic or a little longer; anterior lobe fleshy, narrow, with abruptly rounded 
tip, including four radials besides the first stout one; outer margin of posterior lobe 
scalloped and moderately convex; rear corner subangular, extending about 20—25 "/o 
of distance trom level of axils of pectorals toward first dorsal. Claspers of mature males 
slender, extending rearward a little less than '/a the distance from axils of pelvics toward 
tip of tail; tip of clasper with prominent blade, but otherwise simple. 

Rostral cartilage narrow, blunt, extending about 60 "/o of distance from front of 
cranium toward tip of snout. 

Color. Upper surface varying from pale yellowish brown to darker greyish brown, 
purplish brown, or mouse gray; skin at bases of prickles and at rear sides of larger 
thorns densely pigmented, thus producing a finely freckled appearance; disc irreg- 
ularly and more or less strongly marked with dark spots or blotches, vaguely outlined 
and varying from specimen to specimen in size, shape, number, and arrangement; some 
individuals variegated in addition with a greater or lesser number of small roundish 
white spots irregularly distributed. Tail with 5—7 irregular dark crossbars, either con- 
tinuous across upper surface or interrupted there, the two more posterior crossbars 
usually nearly black and crossing the first and second dorsal fins; tail on some individuals 
also with whitish lateral blotches, either confined to sides or meeting above; caudal 
membrane wholly or partly black. Lower surface yellowish white, sometimes with pos- 
terior corners of pectorals more or less dusky and with dark crossbars encroaching 
somewhat onto lower surface from sides of tail. 

Size. Males may mature sexually at a length no greater than about 230 mm to 
judge from the length of claspers (Fig. 67). The largest specimen seen (a female, the 
type) is 253 mm long. 

Developmental Stages. The egg cases have not been seen. 

Habits. The depths of capture (230—333 fath.) mark this as a deep-water species; 
nothing else is known of its habits. 

Range. Thus far B. plutonia has been taken only on the continental slope in the 
offings of Georgia and South Carolina, where the original specimens were trawled by 
the Blake in i 880,* off northern North Carolina, where it has been taken by the winter 
trawl fishery, and in the offing of Jacksonville, Florida (see Study Material, p. 297).' 

Synonyms and References: 

Raja^ plutonia Garman, Bull. Mus. comp. Zool. Harv., 8, 1881: 236 (descr., meas., depth, off S. Carolina 
and Georgia); Goode and Bean, Bull. Mus. comp. Zool. Harv., 10, 1883: 225 (listed after Garman, 
1881); Jordan and Gilbert, Bull. U. S. nat. Mus., 16, 1883: 878 (descr., after Garman, 1881); True, 

6. Blake Sts. 316, 317, 321; Lat. 3i°57' N, Long. 78°i9' VV; Lat. 32°07' N, Long. 78°38'W; Lat. 32°43' N, Long. 
77°2i' W. 

7. Kaia plutonia Barnard 1925 (Ann. S. Afr. Mus., 21 [i], 1925: 68) from South Africa, earlier described as K. al- 
balinea by von Bonde and Swart (Fish. Mar. biol. Surv. S. Afr., Rep. 3 [1922], Spec. Rep. 5, 1924: 6, pi. 20, fig. i), 
differs noticeably from Brei'iraja plutonia of the western Atlantic in its considerably shorter tail, in larger and 
fewer thorns around the eyes, and in its pale dorsals. It has been identified provisionally by Norman (Discovery 
Rep., 12, 1935: 46) with Raia spinacidermis Barnard 1923 (Ann. S. Afr. Mus., 13 [8]: 440), which appears to be 
closest to the European Raja fullonica Linnaeus 1758. 

8. Sometimes spelled Raia. 



302 Memoir Sears Foundation for Marine Research 

List Vert. Anim. S. Carolina, in Handb. S. Carolina, 1883: 261 (listed, S. Carolina); Jordan, Rep. U. S. 
Comm. Fish. (1885), 1887: 799 (listed: Deep Sea Fauna); Giinther, Challenger Rep., 22, 1887: 10 
(descr., locals., after Garman, 1881); Goode and Bean, Smithson. Contr. Knowl., 50, JJ (Atlas), 1895: 
27, pi. 8, fig. 26; also, Mem. Harv. Mus. comp. Zool., 22, 1896: 27, pi. 8, fig. 26 (descr., locals., after 
Garman, 1881, ill.); Jordan and Evermann, Bull. U. S. nat. Mus., 47 (i), 1896: 69 (descr., off S. Caro- 
lina); Rep. U. S. Comm. Fish. (1895), 1896: 221 (listed off S. Carolina); Garman, Mem. Harv. Mus. 
comp. Zool., 24, 1899: 380 (range, depth); Engelhardt, Abh. bayer. Akad. Wiss., Suppl. 4 (3), 1913: 
102 (range); Garman, Mem. Harv. Mus. comp. Zool., j6, 191 3: 335, pi. 18, fig. i (refs., descr., ill., 
depth, off S.Carolina); Jordan, Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 26 
(listed, off S. Carolina); Pearson, Invest. Rep. U. S. Bur. Fish., 10 (i), 1932: 18 (off north. N. Caro- 
lina, winter). 
Malacorhina flutonia Garman, Bull. Mus. comp. Zool. Harv., 8, 1881: 237 (subgen. Malacorhina proposed). 

Probable Synonym: 

Raja acanthiderma Fowler, Fish Culturist, 26, 1947: 73; Notul. Nat. Acad. nat. Sci. Philad., 199, 1947: 14, 
ftn. (descr., ill.. Gulf Stream off Florida). 



Breviraja sinus-mexicanus Bigelow and Schroeder 1950 
Figures 62 (lower right), 69, 70 

Study Material. Ninety-four specimens,* males and females (including type), 117 
to 355 mm long, from the northeastern part of the Gulf of Mexico between the offings 
of Pensacola and of the Mississippi River Delta, in 170 to 347 fathoms, all in the U. S. 
National Museum and the Harvard Museum of Comparative Zoology. 

Distinctive Characters. The shortness of its rostral cartilage separates B. sinus-mexi- 
canus from all Skates of the genus Raja. Within its own genus it falls with B. atripinna 
and the B. cubensis-^nd-plutonia group in the great length of its tail. But it is distinguish- 
able from young B. atripinna by the thorniness of its tail and from older ones by the 
shortness of the interspace between its two dorsal fins (cf. Figs. 69, 70 with S'^) and by 
the pale coloration of the dorsals. Its tail is so much thornier than that of the B. cubensis- 
und-p/utonia group that there is no danger of confusing it with either of these, and its 
anterior angle is less obtuse. 

Description. Proportional dimensions in per cent of total length. Female, 228 mm 
long, the type (U. S. Nat. Mus., No. 103376), and male, 321 mm long (U. S. Nat. 
Mus., No. 148275), from the Gulf of Mexico. 

Disc: extreme breadth 44.7, 48.0; length 39.4, 40.0. 

Snout length: in front of orbits 9.4, 10. o; in front of mouth 1 1.4, 1 1.6. 

Orbits: horizontal diameter ^i-S') 3-9 i distance between 2.6, 3.0. 

Spiracles: length 2.2, 2.5; distance between 5.9, 6.^. 

Mouth: breadth 4.6, 5.0. 

Nostrils: distance between inner ends 4.6, 4.8. 

9. Albatross Sta. 2396 in Lat. 28°34' N, Long. 86°48'W; Sta. 2395 in Lat. 28^36' N, Long. 86°5o' VV; Sta. 2398 
in Lat. 28°45' N, Long. 86°26' W; Sta. 2377 in Lat. 29°o8' N, Long. 88°o8' W; Oregon Sta. 307 in Lat. 29°oo' 
N, Long. 88°35' W and Sta. 319 in Lat. 29^20' N, Long. 87°25' W, and Oregon Sts. 62, 351, 472, 476, 481, 489, 
490, and 500 in the same general region. 



Fishes of the Western North Atlantic 



303 




Figure 69. Breviraja sinus-mexicanus, female, 228 mm long, from off Pensacola, Florida (U. S. Nat. Mus., 
No. 103376, type). A Lower side of anterior part of head. B Ventral view of pelvics and base of tail. C Left-hand 
nostril, about \\y,. D Margin of right-hand nasal curtain, about 9.2 X. E Upper teeth, about 23 x. 

Gill Openings: lengths, ist 0.9, 1.2; 3rd 0.9, 1.2; 5th 0.8, 0.6; distance between 

inner ends, ist ii.o, 10. o; 5th 6.2, 5.0. 
First dorsal fin: vertical height 1.3, 1.6; length of base 3.5, 3.7. 
Second dorsal fin: vertical height 1.3, 1.6; length of base 2.6, 3.7. 
Pelvics: anterior margin 10.5, 10.3. 
Distance: from tip of snout to center of cloaca 32.9, 2S-S'^ from center of cloaca 



3 04 Memoir Sears Foundation for Marine Research 

to ist dorsal 54.8, 51.5; to tip of tail 67.1, 64.5; from rear end of 2nd 
dorsal base to tip of caudal — , 4.1. 
Inters-pace between: ist and 2nd dorsals 0.9, 0.6. 

Disc about i.i — 1.2 times as broad as long; snout angular, blunted at tip; maximum 
anterior angle in front of spiracles about 100—110°; anterior margins nearly straight 
anterior to level of orbits, weakly convex thence outward on females but rather conspi- 
cuously concave at general level of spiracles on mature males, and tip of snout more 
prominent; outer and posterior corners broadly rounded; posterior margin evenly and 
rather strongly convex and inner margin weakly convex to axil. Axis of greatest breadth 
about 72 "/o of distance rearward from snout toward axils of pectorals. Tail from center 
of cloaca to first dorsal about 1.5— 1.7 times as long as distance from center of cloaca 
to snout and about twice as long to tip; its lateral folds narrow, extending entire length 
from close posterior to tips of pelvics and widening a little rearward. 

Upper surface of disc roughened with small, sharp, movable spines except on pos- 
terior margins and along mid-dorsal belt, most of them covered with skin ; skin above 
eyes finely prickly; two small thorns over tip of rostral cartilage on youngest, three 
small and one large on half-grown specimens; none on tip on adults, but there is a 
cluster a little rearward; about 7—1 1 of various sizes around inner margin of orbit on 
adults, either in continuous row or interrupted midway; one large thorn inward from 
inner end of spiracle; a patch of 8 — 12 scattered outward from each eye on half-grown 
specimens, increasing in number and size and spreading by maturity over the marginal 
belt as a whole anterior to level of spiracles; at first two, later 3—5, large thorns on each 
shoulder; midline of back with a continuous row of 1 1— 14 thorns from nuchal region 
to pelvic girdle, larger ones alternating with smaller at first but later all about equal in 
size, followed by about 45-50 along tail to first dorsal, decreasing in size rearward 
and in regularity of arrangement ;i*' the midrow flanked on either side by one irregular 
additional row from a little behind pectoral girdle to base of tail and by 1—2 such rows 
thence rearward past dorsals; central and posterior parts of pelvics with a few small 
prickles, also upper parts of dorsals; caudal membrane smooth. Mature males with 
thorns of mid-dorsal belt of disc somewhat fewer, smaller, and in less regular linear 
arrangement than on females; also outer corners of pectorals strewn with small thorns 
which are not present on females; alar spines about 12-20, directed rearward-inward, 
irregularly arranged, not conspicuous. Lower surface smooth except sometimes a few 
small spinelets along outer edges of tail close behind tips of pelvics, and a few small 
scattered prickles along its anterior one-sixth. 

Snout in front of orbits about 2.6 times as long as orbit, its length in front of 
mouth about 2.1 times as great as distance between exposed nostrils. Orbit about 1.3 
times as long as distance between orbits and about 1.6 times as long as spiracle. Dis- 
tance between first gill openings 2.1—2.4 tinges as long as distance between exposed 
nostrils, 1.3— 1.4 times between fifth gill openings; first gill openings 1. 1—2.0 times as 

10. Counting is difficult along the tail. 



Fishes of the JVestern North Atlantic 



305 




Figure 70. Breviraja sinus-mexicanus, male, about 321 mm long, from Gulf of Mexico off Mississippi River 
delta (U. S. Nat. Mus., No. 148275). 

long as fifth and about V5 as long as breadth of mouth. Nasal curtain more or less jagged, 
posterior (outer) margin of nostril more or less definitely fringed. Mouth nearly straight 
in females, more arched centrally in mature males. 



3 o6 Memoir Sears Foundation for Marine Research 

Teeth |^2' dose-set in quincunx in females, mostly with low blunt cusp, but 
cusp lacking on some near corners of mouth; those of adult males in transverse series 
and with sharp conical cusp. 

Dorsals similar in shape; either confluent or with interspace up to about Va as long 
as base of first dorsal ; interspace with one or two thorns ; base of first dorsal about i .o- 
1.3 times as long as base of second. Caudal membrane posterior to second dorsal about 
1.3— 1.4 times as long as base of second dorsal. Pelvics deeply concave outwardly, scal- 
loped around indentation; anterior margin about 88 "/o as long as distance from pelvic 
origin to rear tip; anterior lobe narrowing to slightly blunted tip, with four slender 
radials besides first stout one; outer margin of posterior lobe strongly convex and weakly 
scalloped; rear corner abrupt, reaching back about ^s of distance from level of axils 
of pectorals toward first dorsal. Claspers of adult male slender, widening a little toward 
tips, extending rearward about Vs of distance from axils of pelvics toward first dorsal ; 
no terminal spines exposed when not in function. 

Rostral cartilage narrowly triangular, its tip reaching a little more than halfway 
from front of cranium toward tip of snout. Anterior rays of pectorals reaching about 
halfway between level of tip of rostral cartilage and tip of snout. 

Color. Upper surface brownish purple, palest on either side of rostral ridge, along 
edges of pectorals, and on pelvics; either plain-colored ^^ or irregularly marked with 
small slightly darker blotches, indistinctly outlined, the most conspicuous being one 
to four on each side, close to base of tail^^ and midway out on disc anterior to pelvic 
girdle." Skin over prickles of small females and juvenile males much more deeply 
pigmented, thus producing a dark speckled appearance; only scattered prickles so 
pigmented on mature specimens. Lower surface plain yellowish white, but darker hue 
of upper surface showing through along outer edges of pectorals. 

Size. Males are mature at a length of 310—335 mm, and the presence of large 
thorns on the malar regions of the head of a female 325 mm long suggests that this 
species does not grow to a length greater than about 375 mm. 

Developmental Stages. The egg cases have not been seen. 

Habits. Seemingly this is a deep-water species. The stomach of one contained the 
remains of fish. 

Range. Known only from the northern and northeastern part of the Gulf of Mexico, 
at the localities and depths listed under Study Material (p. 302). 

Reference : 

Breviraja sinus-mexlcanus Bigelow and Schroeder, Bull. Mus. comp. Zool. Harv., lOj {j), 1950: 396 (descr., 
iUs., Gulf of Mexico, oiF Pensacola, Florida). 

Breviraja spinosa Bigelow and Schroeder 1950 

Figures 62 (lower left), 71 

Study Material. Female, 280 mm long, from off mid-Florida, Lat. 29°4i' N, Long. 
79°55' W, in 373 fathoms, taken at Albatross St. 2664, in U. S. National Museum; 

II. One adult female is plain-colored. 12. This is the case on a half-grown female. 

13. An adult male shows this pattern. 



Fishes of the Western North Atlantic 307 

two females, 187 and 288 mm long (including the type), from offing of Jacksonville, 
Florida, Lat. 30°2i' N, Long. 79°55' W, and Lat. 30°58' N, Long. 79°34' W, in 290 
fathoms, collected by the Atlantis, in Harvard Museum of Comparative Zoology; also 
four females, probably this species, the largest about 220 mm long, taken on the con- 
tinental slope in the offings of North Carolina and Delaware Bay, in 727 and 258 
fathoms, in U. S. National Museum.'* 

Distinctive Characters. The presence of three or more irregular rows of large and 
conspicuous thorns along the midbelt of its disc marks this species off from all other 
western Atlantic soft-nosed Skates (genera Breviraja^ Psammohatis) except E. yucatanen- 
sis. It is separable from the latter in that it has more conspicuous thorns, its snout anterior 
to the orbits is not more than about twice as long as the distance between the orbits, its 
first and second dorsal fins are confluent at the base, and the upper surface of its disc 
is not closely freckled with small dark dots. 

Description. Proportional dimensions in per cent of total length. Females, 187 and 
288 mm long (type), from the offing of Jacksonville, Florida (Harv. Mus. Comp. Zool., 
Nos. 36369, 36373). 

Disc: extreme breadth 47.7, 50.8; length 41.2, 42.8. 

Snout length: in front of orbits 8.0, 8.0; in front of mouth 9.1, 10.2. 

Orbits: horizontal diameter C1.6, 4.8; distance between 3.2, 3.6. 

Spiracles: length 3.7, 3.4; distance between 6.9, 7.0. 

Mouth: breadth 6.4, 7.8. 

Exposed Nostrils: distance between inner ends 5.3, 6.0. 

Gill openings: lengths, ist 1.6, 1.4; 3rd 1.6, 1.4; 5th 1.5, 1.2; distance between 
inner ends, ist 14.2, 15.9; 5th 8.3, 7.9. 

First dorsal fin: vertical height 1.9, 1.4; length of base 5.4, 5.8. 

Second dorsal fin: vertical height 1.6, 1.5; length of base 5.4, 5.5. 

Pelvics: anterior margin 10.7, 11.6. 

Distance: from tip of snout to center of cloaca 39.0, 42.3; from center of cloaca 
to 1st dorsal 48.1, 44.4; to tip of tail 61.0, 57.7; from rear end of 2nd 
dorsal base to tip of caudal 2.1, 2.0. 

Interspace between: ist and 2nd dorsals 0.0, 0.0. 

Disc about 1.2-1.3 times as broad as long, obtusely rounded in front; tip of snout 
blunt at apex and projecting little if at all; maximum anterior angle in front of spiracles 
about 135-140°; anterior margins convex to level of orbit, thence nearly straight to 
broadly rounded outer corners, the contour continuing in an unbroken arc around the 
convex posterior margin and broadly rounded posterior corners to axils of pectorals. 
Axis of greatest breadth 70-75 "/o of distance back from tip of snout toward axils of 
pectorals. Tail moderately robust, decreasing in relative length with growth; distance 
from center of cloaca to first dorsal about 1.3 times as great as distance from center of 
cloaca to tip of snout on young, about i.i times on larger specimens; extreme length 

14. Albatross Sts. 2624 and 2730; these specimens are in poor condition. 

20* 



3o8 Memoir Sears Foundation for Marine Research 




Figure 71. Breviraja spinosa, female, 288 mm long, from off northern Florida, Lat. 3o°58' N, Long. 79°34' W, 
Atlantis St. 3781, 250-290 fathoms (Harv. Mus. Comp. ZooL, No. 36373, type). A Lower surface of mid- 
anterior part of head. B Ventral view of pelvics. C Side view of posterior part of tail, about 1.2 X. D Left-hand 
nostril, about 6.4 x. E Margin of right-hand nasal curtain, about 6.4 X. F Upper teeth, about 12 X. 

of tail from center of cloaca about 1.6 times as great as distance from cloaca to snout 
on young, about 1.4 times on larger; the lateral folds narrow, confined to posterior third 
of tail. 



Fishes of the Western North Atlantic 309 

Upper surface of disc conspicuously rough; medium-sized thorns scattered along 
rostral ridge; about 10 around anterior and inner margins of orbit on largest specimen, 
fewer on smaller, sometimes one large thorn close in front of orbit ;i^ one to two large 
thorns inward from spiracle; a few small thornlets in space between eyes on small speci- 
mens; a patch of 12-15 conspicuous thorns near outer margin abreast of eye and spir- 
acle; 1—3 conspicuous thorns on midline in nuchal region; inner parts of pectorals with 
many moderate-sized thorns irregularly distributed, progressively smaller outward, 
posterior margins of pectorals smooth; 2—3 large thorns on each shoulder; also a band 
of thorns of various sizes along midbelt of back from nuchal region rearward, irregularly 
scattered to anterior part of tail but arranged partially in 3—5 rows back along tail to 
first dorsal, and successively smaller along rear half of tail, followed by 1—2 rows 
of smaller ones along either side of tail to tip; thorns on disc and some of those on tail 
large and their bases conspicuously radiate. Some small specimens lacking the pair of 
thorns between the spiracles, but with thorns more numerous along midbelt of disc and 
more nearly linear in arrangement, including a definite median row of about 25 along 
tail, besides 1-2 less regular rows along either side of tail. Disc as a whole (except 
above body cavity and around posterior margins of pectorals) further roughened by 
prickles of various sizes, these more numerous on small specimens than on large; skin 
above eyes with a few small prickles on large specimens, more prickly on smaller ones ; 
each side of tail with a band of close-set prickles extending along anterior two-thirds on 
young but confined to anterior third of tail on larger specimens, followed rearward by 
scattered small thornlets; space between orbits more prickly on young than on adults; 
anterior parts of dorsals prickly, decreasingly so with growth; caudal membrane with 
a few scattered prickles; pelvics smooth. Alar thorns of adult male not seen. Lower 
surface smooth, except sometimes with a few small spines along edges of tail abreast 
of posterior lobes of pelvics.^* 

Snout in front of orbits about 1.5— 1.7 times as long as orbit, its length in front 
of mouth about 1.5— 1.7 times as great as distance between exposed nostrils. Orbit about 
1.3— 1.7 times as long as distance between orbits and about 1.5 times as long as spiracle. 
Distance between first gill openings 2.3—2.7 times as long as distance between exposed 
nostrils; between fifth gill openings 1.3 — 1.5 times; first gill openings i. 0—1.5 times as 
long as fifth and about 18—27 % ^s long as breadth of mouth. Nasal curtain fringed, 
posterior (outer) margin of nostril either fringed or only somewhat irregular. Mouth 
weakly arched centrally. 

Teeth j^lllf on specimens seen, perhaps more numerous on adults; those of 
females close-set in quincunx, the older rows worn smooth, the younger with low 
triangular cusp; teeth of adult male not seen. 

First and second dorsals similar in size and shape, confluent at base, their upper 
margins noticeably sloping, their posterior margins slightly recurved. Caudal mem- 
brane posterior to second dorsal a little less than half as long as base of second dorsal. 

15. This is true of one of the fragmentary specimens from the offing of Delaware Bay; see Study Material, p. 307. 

16. On our larger specimen one side of the tail but not the other has these spines. 



3 I o Memoir Sears Foundation for Marine Research 

Pelvics deeply concave outwardly in subtriangular contour, weakly scalloped around 
the indentation; anterior margin about 65—70 "/o as long as distance from pelvic origins 
to rear tips of pelvics; anterior lobe small and fleshy with well rounded tip and three 
radials besides the first stout one; posterior lobe rather strongly convex outwardly, 
its rear corners abruptly rounded and reaching back a little less than Vs of distance 
from level of axils of pelvics toward first dorsal. Claspers of mature males not seen. 

Rostral cartilage triangular, extending about half the distance from front of cranium 
toward tip of snout. Anterior rays of pectorals reaching close to tip of snout, not far 
apart there. 

Color. Upper surface light brown, the disc either plain or indistinctly marked 
with a few whitish blotches, larger and smaller, arranged symmetrically on either side 
of median line ; anterior parts of dorsals and tip of caudal membrane brownish ; some 
specimens with lower surface either plain whitish (including one of the larger speci- 
mens) or pale brownish (after many years in alcohol), but others with dark brown 
blotches developing inward from each nostril; central part of disc and anterior parts 
of pelvics of some also becoming more or less washed or blotched with sooty brown. 

Remarks. The only Albatross specimen that is still in adequate condition differs 
from the type specimen (taken off Jacksonville, Florida) in having one large thorn 
close in front of each orbit, fewer small thorns along the inner margins of the orbits, 
the mid-dorsal thorns somewhat more conspicous, and the rostral cartilage perhaps 
a little longer and narrower relatively. But these divergences do not seem wider than 
can be explained on the basis of individual or local variation, to judge from conditions 
among better known Skates. 

Size. The size at maturity is not known. 

Developmental Stages. The egg cases have not been seen. 

Habits. Seemingly this is a deep-water species, the few specimens taken thus far 
having been trawled at 250, 258, 373 and 727 fathoms. 

Range. Known only from off middle and northern Florida, off North Carolina, 
and off Delaware Bay, a total of seven specimens (see Study Material, p. 306). 

Reference : 

Breviraja spinosa Bigelow and Schroeder, Bull. Mus. comp. Zool. Harv., 103 (7), 1950: 400 (descr., ill., off 
Jacksonville, Florida). 

Breviraja yucatanensis Bigelow and Schroeder 1950 
Figure 72 

Study Material. Juvenile male (type), 2 1 5 mm long, from notheastern slope of 
Yucatan, in 231 fathoms, in U. S. National Museum. 

Distinctive Characters. This newly discovered species is marked off from all other 
known members of its genus in the western Atlantic, except for B. spinosa., by the 
presence of two or more irregular rows of thorns along the median belt of its disc 
rearward from the scapular region. It differs from B. spinosa in having less conspicuous 



Fishes of the Western North Atlantic 



311 



thorns, snout anterior to the orbits about 3.5 times as long as the distance between 
the orbits (not more than 2 times in B. spinosa\ first and second dorsal fins separated 
by a definite (though short) interspace with one or two thorns, and in having the upper 
surface of its disc closely freckled with small dark brown dots (not freckled on B. 
spina so). 




Figure 72. Breviraja yucatanensis, juvenile male, 215 mm long, type, from 
northeast slope of Yucatan (U. S. Nat. Mus., No. 148273). J Dorsal view to 
show thorns and prickles. B Same to show color pattern, without thorns and 
prickles. C Pelvic fins, ventral view. D Ventral view of midanterior part of 
head. E Part of same to show fringed nasal curtain and margin of nostril. 
A-D, about 0.9 X; E about 2.7 X. 

Description. Proportional dimensions in per cent of total length. Male, 2 1 5 mm 
long, from Lat. 20° 19' N, Long. 87°04' W (U. S. Nat. Mus., type. No. 148273)- 
Disc: extreme breadth 55.0; length 45.2. 
Snout length: in front of orbits 9.8; in front of mouth 11.6. 
Orbits: horizontal diameter 4.9; distance between 2.8. 



312 Memoir Sears Foundation for Marine Research 

Spiracles: length 2.6; distance between 6.5. 

Mouth: breadth 6.7. 

Exposed nostrils: distance between inner ends 7.0. 

Gill openings: lengths, ist 1.4; 3rd 1.9; 5th 1.3; distance between inner ends, 

ist 13.2; 5th 8.1. 
First dorsal fin: vertical height 2.3; length of base 5.8. 
Second dorsal fin: vertical height 2.1; length of base 4.9. 
Pelvics: anterior margin 12.6. 
Distance: from tip of snout to center of cloaca 39.5; from center of cloaca to 

1st dorsal 44.7; to tip of tail 60.5; from rear end of 2nd dorsal base to 

tip of tail 3.7. 
Interspace between: ist and 2nd dorsals 1.4. 

Disc about 1.2 times as broad as long, moderately obtuse in front, tip of snout 
hardly protruding; maximum anterior angle in front of spiracles about 128°; anterior 
margins weakly convex from tip of snout to level of eyes, slightly concave at level of 
spiracles and straight thence outwardly; outer corners broadly rounded; posterior mar- 
gins weakly and evenly convex; posterior corners broadly rounded, their curvature 
continuous to axils of pectorals. Lateral folds on tail extremely narrow, extending along 
posterior four-fifths nearly to tip. Tail from center of cloaca to first dorsal about i.i 
times as long as distance from center of cloaca to tip of snout, and about 1.5 times as 
long to tip. Distance from axils of pelvics to first dorsal about as long (i.i) as distance 
from axils of pelvics to fronts of orbits, about 90 "/o as long as distance from center of 
cloaca to tip of snout. 

A few small thorns on anterior part of rostral ridge; 5-7 along inner anterior 
margin of orbit;i' 1-2 on inner margin of orbit and a group of about four between 
inner posterior edge of orbit and inner end of spiracle; about three on nuchal region; 
2-3 on each shoulder; two rows along mid-dorsal belt of disc from a little behind 
pectoral girdle rearward, with an occasional thorn between them in the midline; outer 
anterior parts of pectorals sparsely strewn with small thorns grading down to prickles; 
inner parts of pectorals as well as nuchal region roughened generally with minute 
prickles; a few prickles here and there on anterior part of head, between thorns on 
shoulders and along mid-dorsal belt; skin over eyes prickly; posterior parts of pectorals 
smooth. Upper surface of tail with 3-4 irregular rows of thorns along anterior part, 
these about as large as those on disc, decreasing to 2-3 rows rearward to first dorsal 
fin; one thorn in interspace between first and second dorsals; each side of tail (from 
axil of pelvic) with a band of close-set prickles along anterior half, decreasing in number 
posteriorly to 1—2 rows interspaced with small thornlets past second dorsal fin; dorsal 
fins and upper surface of pelvics smooth. Lower surface smooth everywhere. 

Snout in front of orbits about 3.5 times as long as distance between orbits, its 
length in front of mouth about 1.6 times as great as distance between exposed nostrils. 

17. Five on one side and seven on the other on specimen examined. 



Fishes of the Western North Atlantic 313 

Orbit about 1.7 times as long as distance between orbits and 2.2 times as long as 
spiracle. Distance between first pair of gill openings about 1.9 times as great as distance 
between exposed nostrils, about 1.2 times between fifth gills; first gills about 1.3 times 
as long as fifth gills and 21 "/o as long as breadth of mouth. Nasal curtain, as well as 
expanded outer (posterior) margin of nostril, fringed. Mouth arched only a little for- 
ward in juveniles and probably in females, perhaps more strongly so in adult males. 

Teeth 46 in upper jaw; those of juvenile males (probably those of females also) 
oval, rounded, without cusp, close-set in quincunx; those of mature males not seen. 

First and second dorsals about alike in size and shape, the anterior margins slop- 
ing and weakly convex, the posterior margins slightly recurved; dorsal fins separated 
by a short but definite interspace with one thorn. Caudal membrane posterior to second 
dorsal about 30 "/o as long as base of first dorsal. Pelvics deeply concave outwardly, 
with three conspicuous scallops at base of concavity; anterior margin about 60 "/q as 
long as distance from pelvic origin to rear tip; anterior lobe fleshy with rounded tip; 
posterior lobe weakly and evenly convex outwardly, rear tip narrowly blunted, reaching 
about Vs of distance from axils of pelvics toward first dorsal fin. Claspers of mature 
males not seen. 

Rostral cartilage extends a little more than 2/3 (69-70 %) of distance from front 
of cranium toward tip of snout. Anterior radials of pectorals extend nearly to level of 
tip of snout. 

Color. Upper surface (after many years in alcohol) brownish gray; disc and an- 
terior part of tail thickly freckled with darker brown dots; posterior part of tail with 
similar dots aggregated in two indistinct crossbars; also an additional more definite 
dark bar crossing tail and anterior part of each dorsal fin; caudal membrane sooty. 
Lower surface uniformly pale yellowish. 

Size. A male (only specimen known), 215 mm long, is juvenile, its claspers reach- 
ing only about halfway along inner margins of pelvics. Size at maturity not known. 

Habits. The depth of capture (231 fath.) of the only recorded specimen suggests 
that this is a deep-water species, but nothing definite is known of its habits. 

Range. So far known only from the slope off northeastern Yucatan, in 23 1 fathoms. 1* 

Reference : 

Breviraja yucatanensis Bigelow and Schroeder, Bull. Mus. comp. Zool. Harv., 103 (7), 1950: 402 (descr., ill., 
ofF Yucatin). 

Genus Cruriraja Bigelow and Schroeder 1948 

Cruriraja Bigelow and Schroeder, J. Mar. Res., 7 (3), 1948: 549; type species, C. atlantis Bigelow and Schroe- 
der. OfF north coast of Cuba. 

Generic Synonyms: 

Raia (in part) von Bonde and Swart, Fish. Mar. biol. Surv. S. Afr., Rep. 3 (1922), Spec. Rep. 5, 1924: 9, 11, 

pi. 21, fig. 2, pi. 22, fig. I, for R. parcomaculata and R. durbanensis von Bonde and Swart 1924. South 

Africa. Not Raja Linnaeus 1758. 

18. Albatross Sta. 2359, Lat. 2o°i9' N, Long. 87^04' W, January 29, 1885. 



314 Memoir Sears Foundation for Marine Research 

Generic Characters. Rajidae with pectorals of ordinary form, without spatula-like 
lateral processes. The outer margins of the pelvics are so deeply notched that the 
anterior division, which arises independently from the lower surface of the disc some 
distance inward from the edge of the latter, is entirely cut off from the remainder of 
the fin as a separate limb-like structure. The two or three radial cartilages that are 
ordinarily borne near the anterior end of the basipterygial cartilage are lacking, and 
the gap resulting from their absence corresponds in position to the gap seen externally 
between the anterior and posterior subdivision of the pelvic fin. Anterior subdivision 
of pelvics slender, subcylindrical, tapering toward tip, stiff proximally but softer distally, 
consisting of three articulated segments and hence flexible at two or three points as 
well as at its base (stoutness of musculature toward its base suggests considerable 
powers of movement); its support is maintained by the first stout radial and two or 
three slender radials which are articulated directly to the outer extremity of the pelvis. 
Posterior lobe of pelvics finlike, supported by about 1 8 radials. Rostral cartilage rigid, 
narrow, extending nearly or quite to tip of snout. Tips of anterior pectoral rays falling 
well short of tip of rostral cartilage. Characters otherwise as in Raja. 

Remarks. The limb-like modification of the anterior subdivisions of the pelvics 
of Cruriraja is the outstanding feature of the genus. In this respect it appears to be 
unique among batoids so far as is known, except for the little-known genus Anacantho- 
hatis von Bonde and Swart 1924,18 from Natal, in which the anterior subdivisions of 
the pelvics are described as "leg-like" and "segmented" (as in Springeria, p. 328), and 
except for one genus of Electric Rays (Typhlonarke), family Narkidae. The function 
of the pelvic limbs is not known ; they may be tactile, as are the feeler-like ventrals 
of the hakes (Urophycis), or they may assist in the progress of their owners over 
the bottom. 

Size. The two known species from the Atlantic are small, males having large and 
fully developed claspers at lengths no greater than 300-325 mm. But the maximum 
size of the Indian Ocean representatives is not known, for no mature specimens were 
included among the few that have been seen. 

Habits and Range. Nothing whatever is known of the life histories of the members 
of the genus, except that they are confined to moderately deep water, the recorded depths 
of capture ranging from 280-425 fathoms for the Atlantic (p. 319) and from 298- 
470 fathoms off the Natal Coast. 

Key to Species 2" 

I a. Interspace between first and second dorsals about 1.5 times as long as base of 
first dorsal. atlantis Bigelow and Schroeder 1948, p. 315. 

I b. Interspace between first and second dorsals not more than about half as long as 
base of first dorsal. 

19. Fish. Mar. biol. Surv. S. Afr., Rep. 3 (1922), Spec. Rep. 5, 1924: 18, pi. 23. 

20. Kaja andamanka Lloyd 1909 (Mem. Indian Mus., 2 [3], 1909: 140; 111. Zool. Investigator, Fishes, 10, 1909: pi. 46, 
fig. 2) may prove to be a Cruriraja also, when the nature of its pelvic fins is known definitely. 



Fishes of the Western NortJi Atlantic 315 

2 a. No large thorns along midline of back between levels of spiracles and of outer 
corners of pectorals, or on shoulder regions; upper surface without small 
prickles except close along outer anterior margins. 

poeyi Bigelow and Schroeder 1948, p. 319. 
2 b. Midline of back with an unbroken line of large thorns from nuchal region 
back onto tail; upper surface partly or entirely roughened with smaller prickles 
in addition to the larger thorns. 

3 a. A group of thorns on tip of snout and others along anterior half of rostral 
ridge; two thorns in interspace between dorsal fins. 

parcomaculata (von Bonde and Swart) 1924.^' 
Off Natal Coast, South Africa. 
3 b. No thorns on tip of snout, along anterior half of rostral ridge, or in inter- 
space between dorsal fins. durbanensis (von Bonde and Swart) 1924.22 

Off Natal Coast, South Africa. 



Cruriraja atlantis Bigelow and Schroeder 1948 
Figures 73, 74 

Study Material. Thirteen specimens, male and female, 100—332 mm long, includ- 
ing the type (No. 36320), trawled by the Atlantis off Havana and the north central 
coast of Cuba, in the Harvard Museum of Comparative Zoology. 

Distinctive Characters. Except for its genus mates C. poeyi, C. parcomaculata and 
C. durbanensis, C. atlantis is separable from all other known members of the family 
Rajidae by the complete subdivision of its pelvic fins. And the great width of the inter- 
space between its first and second dorsal fins marks it off from its three genus mates. 
The fact that the upper surface of its disc Is about uniformly covered with small prickles 
further separates it from C. poeyi; the rounded outer corners of its disc distinguish it 
from C. parcomaculata, and the several rows of enlarged thorns on its tail separate it 
from C. durbanensis, which has a single row from nuchal region to first dorsal when 
young and a single row falling short of the first dorsal when older. 

Description. Proportional dimensions in per cent of total length. Male, 297 mm 
(Harv. Mus. Comp. Zool., No. 36321). Female, 332 mm (Harv. Mus. Comp. Zool., 
type. No. 36320), from Cuba. 

Disc: extreme breadth 54.2, 52.7; length 41.1, 40.7. 
Snout length: in front of orbits 10. i, 9.5; in front of mouth 12.4, 12.3. 
Orbits: horizontal diameter 4.7, 5.0; distance between 2.7, 2.6. 
Spiracles: length 2.5, 2.6; distance between 6.1, 6.3. 

21. Von Bonde and Swart (Fish. Mar. biol. Surv. S. Afr., Rep. 3 [1922], Spec. Rep. 5, 1924: 9, pi. 21, fig. 2). This 
species was referred by Barnard (Ann. S. Afr. Mus., 21 [i], 1925: 68) to the synonymy of Raja miraletU4 Linnaeus 
1758, previously known from the Mediterranean and the northwest coast of Africa. But we find nothing in the 
published accounts or illustrations of R. miralelus (not seen) to suggest that its pelvics are divided, as they are in 
Cruriraja parcomaculata. 

22. Fish. Mar. biol. Surv. S. Afr., Rep. 3 (1922), Spec. Rep. 5, 1924: 11, pi. 22, fig. i. 



3 1 6 Memoir Sears Foundation for Marine Research 




Figure 73. Cruriraja atlanth, male, 297 mm long (Harv. Mus. Comp. Zool., No. 36321) and female, 332 mm 
long (Harv. Mus. Comp. Zool., No. 36320, type), trawled by Atlantis off Santa Clara Province, north coast 
of Cuba, Lat. 23°22' N, Long. 79°53' VV, 325 fathoms, and Lat. 23°i2' N, Long. 8i°23' W, 375 fathoms. 



Mouth: breadth 6.2, c^.d. 

Nostrils: distance between inner ends 5.2, 5.1. 

Gill openings: lengths, ist 1.2, i.i; 3rd 1.2, 1.3; 5th 0.9, i.o; distance between 
inner ends, ist 10.8, 11.9; 5th 5.7, 6.6. 



Fishes of the Western North Atlantic 317 

First dorsal fin: vertical height 2.4, 2.4; length of base 4.4, 3.6. 

Second dorsal fin: vertical height 2.4, 1.8; length of base 3.7, 3.8. 

Pelvics: anterior margin 10.4, 9.0. 

Distance: from tip of snout to center of cloaca 37.3, 38.0; from center of cloaca 

to 1st dorsal 39.8, 41.2; to tip of tail 62.7, 62.0; from rear end of 2nd 

dorsal base to tip of caudal 3.7, 4.4. 
Interspace between: ist and 2nd dorsals ii.i, 9.0. 

Disc about 1.3 times as broad as long, angular in front, tip of snout narrowly 
rounded, maximum anterior angle in front of spiracles about 80 to 95°; anterior mar- 
gins weakly concave close behind tip of snout, otherwise nearly straight in females but 
noticeably concave again abreast of spiracles in mature males; outer corners narrowly 
rounded, posterior corners more broadly so; posterior and inner margins weakly con- 
vex. Axis of greatest breadth about 70 "/o of distance rearward from tip of snout toward 
axils of pectorals. Tail noticeably slender, its lateral folds originating a little before 
first dorsal fin, narrow anteriorly but a little wider from opposite origin of second 
dorsal to tip of tail; its length from center of cloaca to origin of first dorsal about i.i 
times as great as distance from center of cloaca to tip of snout, about 1.4— 1.5 times as 
great to origin of second dorsal, and about 1.6— 1.7 times as great to tip. 

Half-grown specimens and larger with upper surface of disc (including skin over 
eyes) thickly and evenly roughened with small prickles except on tip of snout, around 
posterior margins of pectorals and over naked areas of varying extent on shoulder 
region and rearward along either side of dorsal ridge. Adults of both sexes with 2-3 
rows of small thorns along anterior part of rostral ridge; 1 1 — 12 somewhat larger thorns 
around inner margin of orbit and 1—2 inward from inner end of spiracle; 2—4 on each 
shoulder; a median row of about seven thorns flanked on either side by 4—5 lower 
and more rounded tubercles, from nuchal region to pectoral girdle, these followed 
after a short gap by 4—5 small thorns and then by 5—6 much larger ones to level of 
axils of pectorals; the marginal regions abreast of eyes and spiracles thickly strewn 
with small thorns on radiate bases grading down to prickles. Anterior part of tail with 
5—7 irregular rows of thorns in adult females, these decreasing in number to 2—3 rows 
nearing first dorsal; tails of adult males with about three irregular rows interspersed 
with prickles; interspace between first and second dorsals with 1—2 rows of small 
thorns. Pelvics smooth in both sexes, the dorsals with a few prickles. Small specimens 
(100 mm) with upper surface of disc largely bare of prickles except along median zone 
from shoulder girdle rearward; only 4—5 orbital thorns, 2—3 thorns along mid-dorsal 
line between nuchal region and level of axils of pectorals, and 1—2 rows along tail. 
Alar spines of sexually mature males in 4—5 rows. Lower surface smooth on both 
disc and tail. 

Snout in front of orbit about i. 9-2.1 times as long as orbit, its length in front 
of mouth about 2.4 times as great as distance between exposed nostrils. Orbit about 
twice as long as width between orbits and about twice as long as spiracle. Distance 



3 I 8 Memoir Sears Foundation for Marine Research 

between first gill openings 2.1—2.3 times as long as distance between exposed nostrils, 
I.I — 1.3 times between fifth gill openings; first gill openings i.i— 1.3 times as long as 
fifth and about Vs as long as breadth of mouth. Nasal curtain with narrow, irregular 
fringe, posterior (outer) margin of nostril smooth. Mouth nearly straight in females, 
considerably more arched centrally in adult males. 




Figure 74. Cruriraja atlantis. A Ventral view of pelvics of male pictured in Fig. 73. B Lower side of anterior 
part of head of same. C Margin of right-hand nasal curtain of same, about 4 X . i) Left-hand nostril of same, 
about 6.5 X. E Side view of posterior part of tail of same, about 1.3 X. F Upper teeth of same, about 20 X. 
G Upper teeth of female pictured in Fig. 73, about 20 X. 



Teeth in ^^ series ; those of young and of females to maturity with low blunt 
cusp, close-set in quincunx, mostly with oval base; those of adult males with long sharp 
cusp at and near center of mouth, near its corners cusps triangular and bases more oval. 

First and second dorsals similar in shape and about equal in size. Interspace 
between first and second dorsals about 2.5 times as long as base of first dorsal. Distance 
from rear end of base of second dorsal to origin of caudal membrane about half as long 
as base of second dorsal. Caudal membrane about half as long as base of second dorsal. 
Anterior division of pelvic about equal in length to posterior division; outer margin 
of posterior division moderately convex, more so in females than in males, and slightly 
wavy; rear tip angular, reaching back about 1/4 the distance from axils of pectorals 



Fishes of the Western North Atlantic 319 

toward first dorsal. Claspers of mature males slender, their tips simple, reaching back 
halfway from axils of pelvics toward first dorsal. 

Anterior rays of pectorals extending only about 42—50 "/o of distance from level 
of fronts of orbits toward tip of snout. 

Color. Upper surface pale brown, the disc without definite markings but the tail 
with a dusky cross-band at the region of each dorsal fin ; first and second dorsals black 
(a conspicuous feature) as well as caudal membrane. Lower surface whitish or yellowish; 
plain in young but variously blotched and washed with brown on abdomen, on posterior 
parts of pectorals, and on posterior lobes of pelvics of larger specimens ; anterior divisions 
of pelvics darker brown below in males; tail pale dusky below. 

Size. Judging from the claspers, a male 297 mm long is evidently close to ma- 
turity, hence it is likely that our largest specimen, a female 332 mm long, is close to the 
maximum size for this species. 

Developmental Stages. The egg cases have not been seen. 

Habits. We know only that it is a deep-water species, all specimens so far seen 
having been taken in depths of 280—425 fathoms. 

Range. Known only off the north coast of Cuba. 

Reference : 

Cruriraja atlantis Bigelow and Schroeder, J. Mar. Res., 7 (3), 1948: 550 (descr., ill., off N. Central Cuba). 



Cruriraja poeyi Bigelow and Schroeder 1948 
Figures 75, 76 

Study Material. Six males and nine females, 83 to 328 mm long, including the 
type, trawled by the Atlantis off Cuba, in the Harvard Museum of Comparative 
Zoology and U. S. National Museum; and one trawled by the U. S. Bureau of Fisheries 
vessel Albatross off St. Augustine, Florida (U. S. Nat. Mus., No. 1236 18). 

Distinctive Characters. C. poeyi is separable at a glance from all other Rajidae of 
the western Atlantic, except C. atlantis, by its pelvic fins. It differs from the latter in 
the facts that the upper surface of its disc is largely smooth except around the eyes 
and snout and along the mid-dorsal zone posterior to the scapular region, and that its 
two dorsal fins are close together. The smoothness of its disc also separates it from 
C. parcomaculata and C. durbanensis from South Africa, these being more or less covered 
with small spines in addition to the enlarged median and orbital thorns. 

Description. Proportional dimensions in per cent of total length. Male, 327 mm 
(Harv. Mus. Comp. Zool., No. 36322). Female, 328 mm (Harv. Mus. Comp. Zool., 
type. No. 36324), from Cuba. 

Disc: extreme breadth 60.3, d"},.^:, length 48.9, 48.2. 
Snout length: in front of orbits 11.3, 12.2; in front of mouth 13.7, 14.0. 
Orbits: horizontal diameter 4.9, 5.2; distance between 3.1, 3.3. 
Spiracles: length 2.8, 2.7; distance between 6.7, 7.9. 



320 



Memoir Sears Foundation for Marine Research 




Figure 75. Cruriraja poeyi, male and female, each about 328 mm long, trawled by Atlantis off Santa Clara 
Province, north coast of Cuba, Lat. 22°5o'N, Long. 79°o8' W, 245 fathoms, and Lat. 22°48' N, Long. 78°5o'W, 
210 fathoms (Harv. Mus. Comp. Zool., Nos. 36322 and 36324, type). 

Mouth: breadth 7.0, 7.0. 

Nostrils: distance between inner ends 6.4, 6.4. 

Gill openings : length?,^ ist 1.4, 1.4; 3rd 1.5, 1.5; 5th i.i, i.o; distance between 

inner ends, ist 11.3, 13.7; 5th 6.0, 7.6. 
First dorsal fin: vertical height 2.4, 1.8; length of base 4.1, 4.0. 



Fishes of the IVestern North Atlantic 321 

Second dorsal fin: vertical height 2.6, 1.8; length of base 4.3, 4.6. 

Pelvics: anterior margin 13.5, 12.5. 

Distance: from tip of snout to center of cloaca 41.7, 42.7; from center of cloaca 

to 1st dorsal 44.8, 43.2; to tip of tail 58.3, 57.3; from rear end of 2nd 

dorsal base to tip of caudal 4.0, 4.0. 
Interspace between: ist and 2nd dorsals i.i, 1.2. 

Disc about 1.2-1.3 times as broad as long, angular in front, tip of snout only 
a little blunted; maximum anterior angle in front of spiracles about 100° in females 
and about 85° in males; anterior margins weakly concave close behind tip of snout 
and again at level of spiracles; outer corners narrowly rounded but posterior corners 
broadly so; posterior margins weakly convex; inner margins nearly straight. Axis of 
greatest breadth about 65-70 "/o of distance back from snout toward axils of pec- 
torals. Tail with lateral folds extending from about midway between tips of pelvics 
and origin of first dorsal nearly to tip of tail, widest opposite dorsals; distance from 
center of cloaca to first dorsal about as great as distance from center of cloaca to tip 
of snout, about 1.1-1.2 times as great to second dorsal, and 1.3-1.4 times as great to 
tip of tail. 

Upper surface of disc of adults prickly only along mid-dorsal ridge posterior to 
pectoral girdle and narrowly along anterior margins from level of eyes rearward; 1-3 
large thorns and a few smaller ones on anterior part of rostral ridge; 7-8 in malar 
regions between orbits and outer margins in females but up to 20 or more there on 
males; 7-8 thorns around inner margin of orbit, the series usually interrupted mid- 
way, with 1-2 inward from spiracles; likewise a median row of about 7-9 from about 
halfway between pectoral and pelvic girdles to level of axils of pectorals; tail with 4-5 
irregular rows along anterior part, about four rows along posterior part to first dorsal, 
the thorns spaced progressively wider rearward, no one row exactly in the midline; 
interspace between first and second dorsals with two pairs of thorns, the members of 
each pair side by side; the dorsals prickly; caudal membrane and pelvics smooth. Alar 
thorns of mature males covering a triangular area, broadest anteriorly, based a little 
in advance of outer corners of pectorals. Young specimens with fewer orbital thorns, 
only about three rows on tail and none outward from orbits. Lower surface smooth 
both on disc and on tail. 

Snout in front of orbits about 2.3 times as long as orbit, its length in front of 
mouth 2.1-2.2 times as great as breadth between exposed nostrils. Orbit about 1.6 
times as long as distance between orbits and 1.8 times as long as spiracle. Distance 
between first gill openings i. 8-2.1 times as long as distance between exposed nostrils, 
0.9-1.2 times between fifth gill openings; first gill openings 1.3-1.4 times as long as 
fifth and about V5 as long as breadth of mouth. Nasal curtain more or less conspicuously 
fringed with short rounded lobelets;^^ posterior (outer) margin of nostrils smooth. 
Mouth nearly straight. 

23. On one specimen the nasal curtain is fringed on one side and smooth on the other. 



32 2 Memoir Sears Foundation for Marine Research 

Teeth 4^43; those of females close-set in quincunx, circular to oval at base and 
with low triangular cusp except near corners of mouth where base is more oval and 
cusps are usually lacking, the anterior rows much worn and nearly smooth; those of 
adult males with long sharp cusps, pointing obliquely outward near center of mouth 
beside the symphysis, those toward outer corners of mouth with low triangular cusps. 

First and second dorsals similar in size and shape, the posterior margin some- 
what re-entrant. Interspace between dorsals about 1/3 as long as base of first dorsal. 




Figure 76. Cruriraja poeyi. A Lower side of anterior part of head of male pictured in Fig. 75-5 Margins of 
right-hand (above) and left-hand (below) nasal curtains of same, about 5.8 X. C Side view of posterior part of 
tail of same, about i.zx. D Upper teeth of same, from near center of jaw, about 1 2 x . jff Ventral view of pelvics 
of female pictured in Fig. 75. /" Left-hand nostril of same, about 6.9 x . G Upper teeth of same from near center 
of jaw, about 14 X. 



Caudal membrane posterior to second dorsal about as long as base of first dorsal. 
Anterior division of pelvics about as long as posterior division, the latter strongly con- 
vex outwardly and slightly wavy in females and in young males, only weakly convex 
in adult males; rear tips subangular, reaching back about V4 the distance from axils 
of pectorals toward first dorsal. Claspers of mature males slender with simple tips, 
reaching back a little less than halfway from axils of pelvics toward first dorsal. 

Anterior rays of pectorals extending about 49—55 "/o of distance from level of 
fronts of orbits toward tip of snout. 

Color. Upper surface pale coffee brown except for a translucent area either side 
of rostral cartilage, usually with roundish dark spots, about half as long as orbit, scat- 
tered over disc posterior to spiracles except along outer posterior margins; anterior 



Fishes of the Western North Atlantic 323 

parts of dorsals and tip of caudal membrane black. Lower surface of disc of about 
same shade as upper surface, somewhat mottled paler and darker; pale areas around 
mouth and gill openings; tail paler below than disc. 

Si7.e. The largest seen is 328 mm long, and a male of nearly this size has well 
developed claspers, suggesting that the maximum length for this species is probably 
not much greater than 400 mm. 

Developmental Stages. The egg cases have not been seen. 

Habits. All the specimens taken thus far have been trawled on hard sandy bottom 
in depths of 210-475 fathoms. 

Range. Known only off the north central and southwest-central coasts of Cuba and 
off St. Augustine, Florida. 

Reference : 

Cruriraja poeyl Bigelow and Schroeder, J. Mar. Res., 7 (3), 1948: 555 (descr., ill., off N. coast of Cuba). 

Genus Dactylobatus Bean and Weed 1909 

Dactylobatus Bean and Weed, Proc. U. S. nat. Mus., j6, 1909: 459; type species, D. armatus Bean and Weed. 
From offing of South Carolina, 258 and about 270 fathoms. 2* 

Generic Characters. Rajidae with the six to eight median rays of each pectoral 
elongated to form a narrow spatula-shaped lobe. Outer margins of pelvics so weakly 
concave that fins are not definitely bilobed. Cranium with rostral cartilage extending 
nearly to tip of snout (as in Raja). Tips of anterior rays of pectorals reaching almost 
to level of tip of rostral cartilage. Characters otherwise those of the family. 

Species. D. armatus Bean and Weed 1 909 is the only species of the genus known. 

Dactylobatus armatus Bean and Weed 1909 
Figures 77, 78 

Study Material. Immature male, 278 mm long, with small claspers, from offing 
of South Carolina, Lat. 32°36' N, Long. 77^29' W, in 258 fathoms, and a female, 
264 mm, from Lat. 31° N, Long. 80° W, in about 270 fathoms, in U. S. National 
Museum. 

Distinctive Characters. The presence of a narrow spatula-shaped lobe projecting 
from the margin of each pectoral at about its midlength gives to Dactylobatus armatus 
an outline that has no counterpart among other batoids. 

Description. Proportional dimensions in per cent of total length. Male, 278 mm, 
from Lat. 32°36' N, Long. 77°29' W (U. S. Nat. Mus., cotype. No. 62914). 
Disc: extreme breadth 72.5; length 51.1. 

Snout length: in front of orbits 11.5; in front of mouth 13.3. 
Orbits: horizontal diameter 3.8; distance between 4.0. 

24. Collected by the Albatross on October 21, 1885 and on May 5, 1886. 




Figure 77. Dactylobatus armatus, male, 278 mm long, from off South Carolina, Lat. 32°36' N, Long. JJ°2()' W, 
258 fathoms. Albatross St. 2624 (U. S. Nat. Mus., No. 62914). A Side view of posterior part of tail, about 1.4 X. 
B Margin of left-hand nasal curtain, about 8 x . C Hooked thorn from lower surface of outer corner of pectoral 
fin, about 14 x. 



Fishes of the IVestern North Atlantic 325 

Spiracles: length 2.9; distance between 7.4. 

Mouth: breadth 9.5. 

Nostrils: distance between inner ends 5.6. 

Gill openings: lengths, ist 1.4; 3rd 1.6; 5th 1.3; distance between inner ends, 

ist 14.7; 5th 10.4. 
First dorsal fin: vertical height 2.7; length of base 6.1. 
Second dorsal fin: vertical height 2.3; length of base 5.7. 
Pelvics: anterior margin i 1.9. 
Distance: from tip of snout to center of cloaca 48.2; from center of cloaca to ist 

dorsal 36.7; to tip of tail 51.8; from base of 2nd dorsal to tip of caudal 3.2. 
Interspace between: ist and 2nd dorsals 0.0. 

Disc including lateral lobes about 1.4 times as broad as long, or about i.i times 
as broad as long disregarding the lobes ; its anterior contour, back about to level of shoul- 
ders, nearly an even semicircle; position of tip of snout marked by a low rounded 
prominence; posterior corners so broadly rounded that outer-posterior and inner- 
posterior margins merge without evident transition; posterior corners slightly over- 
lapping pelvics; median 6—8 rays of each pectoral prolonged, forming a narrow spatula- 
shaped lobe that tapers somewhat to a rounded tip and extends out from the general 
contour line of the fin by a distance about equal to that between spiracles. Axis of 
greatest breadth (disregarding lobes) about 65 "/o of distance from snout toward rear 
margins of disc. Tail with narrow lateral folds beginning about an eye's diameter 
posterior to tips of pelvics and extending almost to tip, widest posteriorly; length of 
tail from center of cloaca to first dorsal about 0.76 as great as distance from center of 
cloaca to tip of snout, 0.89 as great to second dorsal, and 1.07 times as great to tip. 

Upper surface of disc sparsely strewn with minute and low conical thorns, their 
tips only breaking the skin; four or five larger curved thorns outlining each orbital 
ridge, with a smaller pair inward from each spiracle; one large, and one or two smaller 
ones on each shoulder; one small thorn midway between scapular thorns and spiracle; 
a row of larger and smaller thorns along midline of back from nuchal region to first 
dorsal fin, the largest ones on anterior part of tail, about 17—18 in number anterior 
to axils of pectorals, about 22 thence to first dorsal; this median row of thorns flanked 
on either side of disc by occasional smaller thorns and along tail by a more continuous 
though irregular series, with a few still smaller thorns low down on its sides; skin 
above eyes prickly; pelvics smooth; dorsals and caudal membrane with a few small 
sharp prickles. Lower surface edged with an irregular double row of sharp thorns from 
tip of snout rearward around head and anterior margins of disc to pectoral lobes, their 
points curved inward toward median line of disc; three irregular rows along anterior 
edge of each pectoral lobe; otherwise smooth below. 

Snout in front of orbits about three times as long as orbit, its length in front of 
mouth about 2.4 times as great as distance between exposed nostrils. Orbit about as 
long as distance between orbits and 1.3 times as long as spiracle. Exposed nostril small, 



326 Memoir Sears Foundation for Marine Research 

about halfway between mouth and tip of snout, its expanded posterior (outer) margin 
a little jagged. First to third gill openings (longest) about half as long as eye; distance 
between inner ends of fifth gills about twice as long as distance between nostrils. Free 
margin of nasal curtain shorter than in most rajids, deeply fringed with narrow rounded 
lobelets. Mouth a little arched centrally in young specimens, perhaps more so in males 
at maturity. 2' 

Teeth ^ in specimen counted, moderately close-set in quincunx, thorn-like with 
needle-sharp cusp curved rearward (into the mouth) on longitudinally oval base; only 
about three rows in function simultaneously in small specimens but perhaps more in 
larger ones. 




Figure 78. Dactylobatus armatus, same as in Fig. 77. A Mouth and nostrils, about 1.8 X. B Lower surface of 
outer corner of left-hand pectoral fin, about 1.8 X. C Upper teeth from near center of jaw, about 17 X. 



Dorsal fins rounded, close to tip of tail, confluent at base or nearly so, without 
intervening thorn; base of first dorsal about as long as distance between orbits, the 
second dorsal a little smaller. Caudal membrane about 37 "/o as long as base of first 
dorsal. Pelvics only weakly concave outwardly but continuously so nearly to rear tips 
and rather strongly scalloped along midsector; anterior margin about 65 % as long 
as distance from its own origin to rear tip of pelvic; tip of anterior lobe rather narrowly 
rounded, rear corners narrowly so, reaching back a little more than 1/3 the distance 
from axils of pectorals toward first dorsal. Claspers of mature male not seen. 

Anterior rays of pectorals reaching nearly to level of tip of snout. 

Color. Upper surface ashy gray after many years in alcohol; irregularly marked 
with sooty or blackish spots and blotches of difi^erent sizes, round to long-oval in shape; 
upper margins of first and second dorsals sooty. Lower surface yellowish white, vaguely 
clouded with mouse gray but described as appearing more or less pinkish with brown 
spots after removal of the mucous that is coagulated by the preservative; posterior 
margins of pectorals and pelvics smoky. 

Size. The claspers of a male 278 mm long are so small (Fig. 77) as to suggest 

25. Sexually mature specimens have not been seen. 



Fishes of the Western Nortli Atlantic 327 

that Dactylobatus is one of the medium-sized if not larger members of its family, but 
its size at maturity is not known. 

Developmental Stages. Nothing known. 

Habits. The depths of capture, 258 and about 270 fathoms, suggest at least a 
moderately deep-water habitat. One of the two specimens taken was from a temperature 
of about 12° C (54° F). Nothing more is known of its habits. 

Range. Known only from the upper part of the continental slope in the offing of 
South Carolina, at the localities and depths listed (Study Material, p. 323). 

References : 

Dactylobatus armatus Bean and Weed, Proc. U. S. nat. Mus., j6, 1909: 459, pi. 38 (descr., ill., upper con- 
tinental slope in offing of S. Carolina); Garman, Mem. Harv. Mus. comp. ZooL, j6, 191 3: 368 (descr., 
after Bean and Weed, 1909); Jordan, Evermann and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 
27 (listed, S. Carolina). 



Family ANACANTHO BATIDAE 

Characters. Rajoidea with tip of snout extended as a filament. Tail without lateral 
folds. No dorsal fins, but with caudal membrane. Pelvics so deeply notched outwardly 
that anterior subdivisions of fins are leg-like, of two segments, resembling pelvics of 
the rajid genus Cruriraja (p. 314) and Typhlonarke among Electric Rays (p. 80); 
inner margins of posterior lobes attached to sides of tail nearly or quite to tips. Skin 
smooth everywhere, without denticles of any sort either on disc or on tail. 

Remarks. The type genus of this family, Anacanthohatis von Bonde and Swart 
1924, is placed in the family Dasyatidae by Fowler^" and by Smith. 2' But the nature 
of its pelvics refers it, rather, to the Rajoidea. 

Genera. Two genera are known, Anacanthohatis von Bonde and Swart 1924, from 
the Natal Coast, South Africa, and Springeria Bigelow and Schroeder 1951 from the 
Gulf of Mexico. 

Key to Genera 

I a. Outer margins of posterior lobes of pelvic fins not united to inner margins of 
pectorals; snout prolonged as a simple filament. 

Anacanthohatis^^ von Bonde and Swart 1924. 

Natal Coast, South Africa. 
I b. Outer margins of posterior lobes of pelvics united along first half to two-thirds of 

26. Bull. U.S. nat. Mus., 100 [13], 1941: 397. 27. Sea Fish. S. Afr., 1949: 71. 

28. The type and only known species was described and figured (von Bonde and Swart, Mar. biol. Surv. S. Afr., Rep. 
3 [1922], Spec. Rep. 5, 1924: 18, pi. 23) as Leiobatis marmoratus, and Anacanthohatis was proposed as a substitute 
in the attached "errata slip" because Leiobatis was preoccupied. The original illustration was of a female; Smith 
(Sea Fish. S. Afr., 1949: 71, fig. 84) has recently pictured a male. A second species, dubius, included by von Bonde 
and Swart in Anacanthohatis (Mar. biol. Surv. S. Afr., Rep. 3 [1922], Spec. Rep. 5, 1924: 19) is referred provisionally 
to Springeria because the outer margins of its pelvics are described as fused along their anterior half with the inner 
margins of the pectorals. But its status will remain somewhat doubtful until it is known whether it has dorsal fins 
or not. 



328 Memoir Sears Foundation for Marine Research 

their length to inner margins of pectorals; snout either expanded terminally in 
form shown in Fig. 78 a or prolonged as a simple filament. 

Springeria Bigelow and Schroeder 1951, p. 328. 



Genus Springeria Bigelow and Schroeder 1951 

Figure 78 a 

Springeria Bigelow and Schroeder 195 1 : J. Wash. Acad. Sci., 41 (3), 1951 : in; type species, S.folirostris Bige- 
low and Schroeder; Gulf of Mexico off mouth of Mississippi River, Lat. 2<foz' N, Long. 88°34' W., 
232-258 fathoms. 

Generic Characters. Snout either prolonged as a simple filament or expanded 
terminally as shown in Fig. 78 a. Lower side of tail as well as upper with caudal mem- 
brane. Outer margins of posterior lobes of pelvics united along first ^/g of their length 
with inner margins of pectorals; inner margins of posterior pelvic lobes attached nearly 
to tips to sides of tail. No radial cartilages along anterior half of basipterygial cartilages 
of pelvic fins. Pelvis transverse, its anterior profile slightly concave rearward, a long 
slender process at either end directed forward. Firm rostral cartilage extending to base 
of filament. Other characters those of the family (p. 327). 

Species. Two species known, S.folirostris Bigelow and Schroeder 1951, from Gulf 
of Mexico, probably also dubia (von Bonde and Swart) 1924, South Africa (see p. 327, 
footnote 28). 

Key to Species 

I a. Snout prolonged as a simple filament. dui>ia von Bonde and Swart 1924. 

South Africa.29 
I b. Snout expanded terminally in leaf-like form (Fig. 78 a). 

foUrostris Bigelow and Schroeder 1951, p. 328. 



Springeria foHrostris Bigelow and Schroeder 1951 
Figure 78 a 

Study Material. Immature male, type, 400 mm long to base of terminal filament, 
from northern Gulf of Mexico off the Mississippi River, Lat. 29°o2' N, Long 88°34' W, 
in 232-258 fathoms (U. S. Nat. Mus., No. 152546); very young male, 125 mm long, 
from same general locality, Lat. 2 9°oi' N, Long. 8 8° 30' W, in Harvard Museum of 
Comparative Zoology. Also 1 1 others, of both sexes, 220—620 mm long, from the 
north central and northeastern parts of the Gulf of Mexico, collected by U. S. Fish 
and Wildlife Service vessel Oregon in 185-254 fathoms, September 1951, at stations 
476, 481, and 489. 

Distinctive Characters. S.folirostris differs from all other known batoids in the 

29. Status doubtful; see p. 327, footnote 28. 



Fishes of the Western North Atlantic 



329 



peculiar leaf-like expansion of the end of its snout. Specimens with this, even though 
the tail were damaged, would still be easily separable from all other rajoids of the 
Atlantic by their perfectly naked skins, and from all dasyatid and myliobatid Rays by 
the nature of their pelvic fins. 




Figure 78a. Springeria folirostris, holotype, male, 422 mm long, from nor- 
thern Gulf of Mexico, Lat. 2g°02' N, Long. 88°34' W, 232-258 fathoms 
(U.S. Nat. Mus., No. 152546). y/ Tip of tail, about 2.3 X. B Nasal curtain 
and mouth, about 2.3 X. C Teeth from upper jaw, about gx. 

Description. Proportional dimensions in per cent of total length. Male, 400 mm 
to base of terminal filament (U. S. Nat. Mus., type. No. i 52546), and female, 620 mm, 
both from the Gulf of Mexico. 



3 3 o Memoir Sears Foundation for Marine Research 

Disc: extreme breadth 51.6, 53.3; length 55.2, 57--- 

Snoui length: in front of orbits 21.8, 22.4; in front of mouth 24.3, 24.3. 

Orbits: horizontal diameter 2.9, 3.3; distance between 2.6, 2.6. 

Spiracles: length i.o, 1.5; distance between 5.1, 5.0. 

Mouth: breadth 4.5, 4.8. 

Nostrils: distance between inner ends 3.8, 4.0. 

Gill openings: lengths, ist 0.75, i.o; 3rd 0.75, i.o; 5th 0.5, 0.6; distance be- 
tween inner ends, ist 9.2, 10.5; 5th 4.8, 5.6. 

Caudal fin: length of base, upper 6.0, 3.7; lower 5.0, 4.4. 

Pelvics: anterior margin 12.7, 12.2. 

Distance: from tip of snout'" to center of cloaca 47.6, 50.0; from center of 
cloaca to tip of tail 52.4, 50.0. 

Disc from base of terminal filament about i.i times as long as broad; maximum 
anterior angle from level of base of terminal expansion of snout to level of spiracles 
about 85°; end of snout expanded in leaf-like form as shown in Fig. 78 a, terminating 
in a slender filament a little longer than distance between spiracles; margins of disc 
rearward from terminal expansion weakly concave or straight about to level of spiracles, 
then altering to continuously and strongly convex around to short inner margins, with- 
out definite outer or posterior corners. Tail slender, compressed dorsoventrally, increa- 
singly so rearward; its width at axils of pelvic fins (where thickest) about 72—^4 ^s 
great as length of eye; its length from center of cloaca to tip about 1.0- i.i times as 
great as distance from cloaca to base of terminal filament of snout. 

Skin completely naked everywhere, without dermal denticles of any sort. 

Snout in front of eyes about 8.4—8.7 times as long to base of terminal filament as 
distance between orbits, its length in front of mouth about 6.0—6.5 times as great as 
distance between exposed nostrils. Orbit about i.i times as long as distance between 
orbits, and 2.2—2.9 times as long as spiracle, which is noticeably small. Gill openings 
minute; first about Ve as long as breadth of mouth; fifth about -/s as long as first; 
distance between inner ends of first gills about 2.5 times as long as distance between 
exposed nostrils, between fifth gills about 1.3 times. Nasal curtain conspicuously 
fringed, each side with 10— 1 1 lobelets; outer margin of nostril only slightly expanded 
with irregular edge; exposed nostril noticeably minute. Mouth on immature males and 
on females a little arched forward, its shape not known for mature males. 

Teeth ^|^ on young males, low, with obscure cutting edge but no cusp, but H 
on largest female, with low, broad triangular cusp, arranged in quincunx. Teeth of 
mature males not seen. 

No dorsal fins. Base of upper caudal fin-membrane about 1.0— 1.6 times as long as 
distance between exposed nostrils, of shape illustrated in Fig. 78 a, its maximum width 
about Vio (about 9 "jo) as great as length of its base; lower caudal membrane about 
half (55 %) as wide as upper, its origin a little posterior to origin of upper; the two 

30. Exclusive of rostral filament, which is 22 mm long. 



Fishes of the IVestern North Atlantic 331 

lobes discontinuous at tip of tail. Anterior leg-like subdivision of pelvics nearly as 
long (95 "/o) ^s distance from pelvic origin to rear corners, broader than thick, fleshy, 
with one articulation about midway of its length, inner edge of the terminal segment 
scalloped, corresponding to tips of the three radial cartilages; posterior lobe of pelvics 
with narrowly rounded rear corner reaching rearward only about as far as rear limits 
of disc; outer margin joined for about ^3 its length to margin of pectoral, inner edge 
joined to side of tail nearly to tip. 

Anterior rays of pectorals extending forward to a little posterior to base of ter- 
minal expansion of snout; firm rostral cartilage reaching about to base of terminal 
filament. 

Color. Ash gray above, except for unpigmented and translucent spaces between 
rostral ridge and anterior rays of pectorals; orbits dusky; terminal expansion of snout 
narrowly and irregularly margined with black; posterior part of back with a sooty 
blotch on one side near midline on largest male, several such blotches, haphazardly 
located, on largest female. Lower surface pale grayish white, the outer posterior belt 
of pectorals sooty gray, terminal expansion of snout narrowly and irregularly edged 
with black; tail sooty at base. 

Size. How large this Skate may grow is not known, for the largest male seen 
so far, 400 mm long to base of terminal filament, is immature, its claspers not yet 
reaching as far as the tips of its pelvics. 

Developmental Stages. Presumably Springeria is oviparous like other rajids, but its 
eggs have not been seen. 

Habits. All the specimens seen were trawled at 185—258 fathoms; this, with the 
improbability that this Skate would have been overlooked if it occurred in shallow 
water, suggests that it is confined to depths greater than about 1 50 fathoms. Nothing 
else is known of its habits. 

Range. So far known only in the northern side of the Gulf of Mexico at the 
localities listed on page 328 under Study Material. 

Reference : 

Springeria folirostris Bigelow and Schroeder, J. Wash. Acad. Sci., ^l (3), 195 1 : 112 (descr., ill., Gulf of Mexico). 



Suborder MYLIO BATOIDEA 

Sting Rays, River Rays, Butterfly Rays, Eagle Rays, Cow-nosed Rays, 
Devil Rays, Mantas 

Characters. Head and trunk anterior to axils of pectorals strongly flattened dorso- 
ventrally, forming a disc with the pectorals, essentially as in Rajoidea; disc ranging 
from oval longitudinally to much broader than long. Tail sharply marked off from body 
sector, ranging from shorter than disc to much longer, in many cases like a whiplash 
toward tip. Pectorals either continuous along sides of head or with anterior portions 
separated along head as independent rostral lobe (or lobes) or as fin-like structures. 



332 Memoir Sears Foundation for Marine Research 




Figure 79. Pelvis of Myliobatoidea, ventral view (left) and dorsal view (right). A Dasyatis cenlroura, after Gar- 
man. B Potamotrygon circularis, after Garman. C Gymnura altavela, after Garman. D Myliobatis freminvillii, 
after Garman. E Mobula hypostoma, about 1,070 mm wide, from Cape Lookout, North Carolina (Harv. Mus. 
Comp. ZooL, No. 1378), about 0.6 X. 



Fishes of the Western North Atlantic 333 

A single dorsal fin situated far forward on tail, or none. A well developed caudal fin 
with radial supports in some but not in others. Outer margins of pelvics either straight 
or convex.^ Spiracles with no trace of gill folds. 

Eyes and spiracles either on dorsal surface of head or on its sides. Apparent inner 
(morphologically anterior) margins of nostrils united and expanded rearward as a 
curtain, either simple or bilobed, the free posterior edge either smooth or fringed, 
concealing all but outer end of nasal aperture. Teeth ranging from small and numerous 
to large and few in number and compacted as a single grinding plate in each jaw. 

Skin of upper surface either naked or variously armed with tubercles or thorns, 
the lower surface either naked or nearly so. Upper surface of tail with a large and saw- 
edged poisonous spine (or spines) in many species but not in all. 

Anterior part of head supported either solely by anterior radial cartilages of pec- 
toral fins or, in the case of the Devil Rays (Mobulidae), by the anterior contour of the 
cranium, which is nearly straight transversely.^ Cranium without rostral projection; 
antorbital cartilages not supporting part of disc. Fins without horny rays (ceratotrichia), 
the cartilaginous radials extending out to margins of pectorals and pelvics, as well as 
to margins of dorsal if present. Pelvis (Fig. 79) more or less strongly arched forward, 
without prepelvic processes at outer ends and with or without a median process directed 
forward. Surfaces of gill arches inward from gill filaments either smooth, or with papillae, 
or with knobs or transverse folds or ridges (Fig. 80). 

Size. The members of this suborder range from only a few inches up to 22 or 
23 feet in breadth and reach a weight of more than 3,000 pounds in the case of the 
Giant Devil Rays {Mobula and Mantd), which are by far the largest of all batoids and 
indeed among the largest of fishes. 

Developmental Stages. Ovoviviparous in all species in which development is known. 

Range. As a whole, the suborder is tropical-subtropical, only a few species ranging 
regularly to warm-temperate latitudes. For the most part they are confined to small 
and moderate depths and are most common in shallow water, where they are plentiful 
in suitable situations around tropical shores. Many of them enter brackish and fresh 
water freely, and one family of the order is confined to fresh water. 

Families. As seen in the list of names at the beginning of this section, the sub- 
order includes a varied assemblage, all members of which agree in the characters 
stated above. 

Key to Families 

I a. A well developed caudal fin with cartilaginous radial supports. 

, -, , 1 r Urolophidae.. p. 416. 

I b. No caudal fin. -^ ^ 

2 a. Outer anterior margins of pectorals continuous along sides of head, without 

separate cephalic fins or rostral lobes; eyes and spiracles on top of head. 

1. This contrasts with their usual concavity in the Rajoidea, as does the position of the dorsal fin. 

2. For a general account of the head skeleton in the families Dasyatidae and Myliobatidae, see Holmgren (.'\cta Zool. 
Stockh., 22, 1941: 62, 66). 



334 Me?notr Sears Foundation for Marine Research 

3a. Disc not more than 1.3 times as broad as long; tail from center of cloaca 
to tip longer than breadth of disc; no distinct dorsal fin; transverse cur- 
tain on roof of mouth with fringed margin ; floor of mouth with several 
fleshy papillae. 

4a. Pelvis not bearing a slender median process directed forward; salt- 
and brackish- water species. Dasyatidae, p. 335. 





Figure 80. Gill arch (above) and lateral view of gill fold (below) of Myliobatoidea; left, Myliobatis goodei, 
990 mm wide, from Rio de Janeiro, Brazil (Harv. Mus. Comp. Zool., No. 994), 0.65 X; right. Rhinoptera 
bonasus, 11 \ mm wide, from Rio de Janeiro, Brazil (Harv. Mus. Comp. Zool., No. 374), about 1.3 X. 



4b. Pelvis bearing a long slender median process, directed forward; 
fresh-water species exclusively. Potamotrygonidae. 

Rivers of Atlantic and 
Caribbean watersheds in 
tropical and warm-tem- 
perate South America. 
3 b. Disc more than 1.5 times as broad as long; tail from center of cloaca to tip 
considerably shorter than breadth of disc; with or without a small dorsal 
fin near midlength of tail; transverse curtain on roof of mouth smooth- 
edged; no papillae on floor of mouth. Gymnuridae, p. 395. 
2 b. Margins of pectorals deeply indented or entirely interrupted j'ust posterior to 
eyes, the anterior part of head thus sharply marked off from remainder of 
disc; anterior subdivisions of pectorals forming a separate lobe (or lobes), 
or fins; eyes and spiracles on sides of head. 
5 a. Anterior subdivisions of pectorals in the form of two thin and narrow 



Fishes of the Western North Atlantic 335 

tin-like projections widely separated (Figs. 113, 116); teeth minute, in 
many series. Mobulidae, p. 480. 

5 b. Anterior subdivisions of pectorals forming either one soft fleshy lobe 
extending forward below front of head, or two such lobes joined together 
basally; teeth large, in few series. 

6a. A single subrostral lobe or fin (Fig. 102). Myliobatidae, p. 433. 
6 b. A pair of subrostral lobes or fins (Fig. 107). Rhinopteridae, p. 465. 



Family DASTATIDAE 
Sting or Whip Rays 

Characters. Myliobatoidea with anterior portions of pectorals continuous along 
sides of head, not separated off as subrostral lobes or fins; anterior part of head not 
marked off from more posterior part of disc; crown only slightly elevated; disc not 
more than 1.3 times as broad as long. Tail slender, tapering, much longer than disc 
in many species; some species with a longitudinal membranous fold (or folds) above 
or below, or both. Upper surface of tail armed in most species with one or more long 
and sharp-pointed poisonous spines with serrate edges, flattened dorsoventrally and 
attached rigidly to skin; tail spines directed rearward with tip lifted only slightly above 
general contour of tail, the one close behind the other if there are two or more; mar- 
ginal teeth of spine directed toward its base, their outer edges forming cutting edges 
interrupted only by short interspaces between one tooth and the next. No rayed dorsal 
or caudal fins. Eyes without expanded velum above pupil; eyes and spiracles on dorsal 
surface. Nostrils separate from mouth, their anterior margins widely expanded and 
confluent across a narrow isthmus, forming a single quadrate curtain with more or less 
deeply fringed margin reaching rearward as far as front of mouth; outer posterior 
margin of nostril expanded as a rounded lobe directed forward. Mouth straight or 
slightly arched, on ventral surface considerably posterior to snout, the lower jaw more 
or less indented centrally with the upper jaw curved rearward to correspond;^ floor 
of mouth with a transverse row of several fleshy papillae of various lengths, their tips 
often more or less swollen. Teeth small, in numerous series, closely crowded in bands 
along jaws, rounded or with one or more cusps, often with ridges or tubercles. Upper 
surface of disc and tail (apart from tail spine) smooth or variously roughened with 
tubercles, thorns or prickles. Front of cranium weakly concave in most, perhaps straight 
in some. Pelvis moderately arched forward in rounded contour but without median 
process.* Surfaces of gill arches smooth inward from gill filaments. 

Remarks. The tail spine is a striking anatomical feature of all but one of the known 

3. This direction of curvature is opposite to that usual among Rajidae. 

4. In the freshwater genera Potamotrygon and Disceus the pelvis bears a long median process extending forward (Fig. 
79B), and it is because of this that they were grouped bv Garman (Mem. Harv. Mus. comp. Zool., 36, 191 3: 415) 
in the separate family Potamotrygonidae, which is accepted here for this same reason. In all other respects (other 
than their freshwater habitat) they agree with the typical Dasyatidae. 



336 Memoir Sears Foundation for Marine Research 

genera of the family. It has been described as renewed yearly, but we find no definite 
evidence that its replacement follows any regular periodic schedule, nor does this seem 
likely to us. While the majority of specimens are armed with a single spine, others have 
two, rarely three or even four, showing that new spines may develop before the old 
one is dropped. On the other hand, it is not unusual to find specimens with tails from 
which the spine has been dropped (leaving only a scar) before any trace is to be seen 
of the spine that should succeed it. At least in some species the replacement spine 
may develop either close anterior to the pre-existing one or close posterior to it.^ How- 
ever, in other species it may be that the replacement spine invariably develops either 
anterior to the older one (Figs. 83 and 90, Dasyalis centroura and Z). say) or posterior 
to it (Dasyatis sabina. Fig. 88). 

The upper (anterior) surface of the spine varies from low-half-oval to nearly semi- 
circular in cross section; its lower (posterior) surface has a rounded median ridge 
flanked on either hand by a deep furrow extending nearly to the tip; the rows of mar- 
ginal teeth mark the transition from upper surface to lower. Although the spine itself 
Is not mobile, the tail is so flexible that it swings far away from the terminal part of 
the spine when lashed about, thus bringing the spine into a position favorable for 
penetrating whatever It may strike. The tails of large specimens are so powerful that 
It Is not exceptional for their spines to drive through a heavy boot and several layers 
of clothing, to penetrate full length into an arm or leg, or to pierce the side of a wooden 
boat deeply. Comparative anatomists and ichthyologists of the twentieth century 
commonly regarded the effects of Sting Ray wounds (p. 333) as due to mechanical 
Injury and to infection by the mucus of the Ray's skin rather than to any specific 
poison. But as long ago as 1848^ it was pointed out that the symptoms following 
these wounds are similar to those resulting from bites by poisonous snakes. It seems 
well established now that the spines do secrete a poison from cells lying In the lateral 
grooves along the side that faces the tail.' While the evidence to this effect is circum- 
stantial, the pain and swelling follow far sooner after the wound is inflicted than might 
be expected In the case of an ordinary Infection. The effects resemble so closely the 
effects of wounds by the European Weaver Fishes, Trachinus (proved venomous by 
clinical experiments of various kinds), and the secretory organ of the Sting Ray resem- 
bles that of Trachinus so closely In histological structure, as to leave no reasonable 
doubt of the venomous nature of the spine.* And the final proof has recently come 
from laboratory experiments on the properties of their venom ^ and on the effects of 

5. In Dasyatis imbricala (Bloch and Schneider) iSoi from the East Indies, the replacement spine is posterior in two 
out of 17 double-spined specimens that we have examined, anterior in the others. 

6. By R. Schomburgk; see p. 339, footnote 24. 

7. See Evans (Philos. Trans., [B] 212, 1923: 1-33) for the histology of the spine organ, Jorg (Nuov. Reun. See. Ar- 
gent. Patol. Reg., 1939: 1599-1616) for the closely allied Potamotrygonidae, and Evans (Sting Fish and Sea Farer, 
1943) for a popular account of venomous fishes. 

8. For an extended historical account of the effects of Sting Ray wounds, see Gudger (Bull. Hist. Med., 14, 1944: 
467-504). 

9. Vellard (C. R. Acad. Sci. Paris, 792, 1931: 1279; Mem. Soc. zool. Fr., 29, 1932: 514-532); see also full summary 
and discussion by Gudger (Amer. Nat., 81, 1947: 297). We might point out also that Jorg (Nuov. Reun. Argent. 
Patol. Reg., J, 1939: 1603), who arrived at a contrary conclusion for Potamotrygon, did not study the histology 



Fishes of the Western North Atlantic 337 

injecting the poisonous tissue from the spines of two species of Brazilian River Rays 
{Potamotrygon) into various batrachians, birds and mammals. 

Size. The members of the family vary widely in size, the smallest being only one 
to two feet across the disc at maturity, the largest'" six to seven feet and perhaps more. 

So far as we can learn, none of the largest ones have ever been weighed, but from 
the relationship between size and weight for Dasyatis centroura up to five feet broad 
(p. 357), we suspect that estimated weights reported for large Rays are likely to be 
considerably too high. 

No information is at hand as to the rate of growth of any of the Sting Rays or 
the number of years required by the larger species to reach full size. But at least some 
of them are long-lived, for one is known to have "lived for 16 years in captivity in 
Berlin."" 

Developmental Stages. The dasyatid Rays are ovoviviparous, the embryos lying 
loose in the uterus without any physical connection with the mother. The yolk sac 
persists throughout the greater part of embryonic life, and the yolk is taken directly 
from it into the alimentary canal. But the chief source of nutriment for the embryo 
is a creamy albuminous fluid secreted by vascular filaments which clothe the uterine 
wall of the mother so closely that they appear like coarse fur. Seemingly this secretion, 
squeezed out by the superficial musculature of the filaments, is taken in through the 
embryonic spiracles, into which the maternal filaments have been found inserted in 
some cases; it is absorbed along the digestive tract, and clots of it have been found within 
the spiral valve of the embryonic intestine. '^ 

Habits. Sting Rays are seen (or caught) most commonly lying on the bottom on 
the flats of bays, shoal lagoons and river mouths or on patches of sand between coral 
heads, etc. Often they are partially buried in the mud or sand with only a portion of 
the tail, eyes, and spiracles exposed. It is chiefly by excavating the bottom with their 
pectoral fins that they obtain the worms, mollusks or crustaceans on which they feed; 
such hollows in the sand are familiar spectacles when they are exposed at low tide in 
regions of the tropics where Sting Rays abound, as in Australian waters. '^ They also 
succeed in capturing small fish in greater or lesser numbers. On the other hand, they 
often fall prey to large Sharks, and it is not unusual to find a Sting Ray spine imbedded 
in a Shark's mouth. It has been observed that an Indian species {Dasyatis kuhlii)., while 
buried in the sand, utilizes the inner posterior margin of the spiracle to form a pro- 
of the terminal portion of the spine where the venomous tissue is situated, but only that of the basal part of the 
spine, and of the tail itself beneath the free portion of the spine. 

10. Dasyatis centroura of the western Atlantic (p. 352), T>. aspera of the eastern Atlantic, and D. bre'vicaudata of New 
Zealand and Australia. 

11. Whitley, Fish. Aust., i, 1940: 201. 

12. Knowledge of the embryonic nutrition of Sting Rays is due chiefly to a series of observations by Alcock (J. Asiat. 
Soc. Beng., sg [2], 1890: 51; Ann. Mag. nat. Hist., [6] 9, 1892: 417); and by Alcock and Wood-Mason (Proc. 
roy. Soc. Lond., 50, 1891: 202). See also Gill (Smithson. misc. Coll., 52, 1909: 173); and Southwell and Prashad 
(Rec. Indian Mus., z6, 1919: 240), and Ranzi (Pubbl. Staz. zool. Napoli, 13, 1934: 39S) for a general survey, with 
references, and for a photograph of the inner uterine wall of the Mediterranean Dasyatis 'violacea. 

13. For an excellent photograph showing the flats dotted with these hollows in North Queensland, see Whitley (Fish. 
Aust., I, 1940: 198, fig. 224). 



338 Memoir Sears Foundation for Marine Research 

jecting fold which is expanded when the water is agitated, thus protecting the spiracular 
opening from sand, etc. Normally, however, the spiracles are open in respiration for 
the intake of water, which is expelled via the gill openings, as in the case of Skates." 

Though lying quiescent for a large proportion of the time. Sting Rays are active 
swimmers on occasion, progressing rapidly by undulating motions of the margins of 
the pectorals much as Skates do. When on migration, or at other times for reasons 
not evident, they swim at the surface or plane along the latter. 

Numerical Abundance. We have found no precise information as to the numerical 
abundance of Sting Rays anywhere. It is common knowledge, however, that in suitable 
localities in tropical coastwise waters they occur in such great plenty that it may seem 
as though the bottom were almost paved with them. 

Relation to Man. Sting Rays of one sort or another are often offered for sale in 
the fish markets of tropical ports, the thicker parts of their discs alone being utilized. 
Their fins are used to some extent for gelatine and their liver-oil "is scarcely distin- 
guishable in appearance or composition from cod-liver oil,"i^ but we are not aware 
that its vitamin content has been measured. 

The spines of Sting Rays are such effective weapons that their use for tipping 
spears, either singly or several bundled together, was once fairly common practice in 
the Malay-Siam region, in New Zealand, among the island groups of the tropical and 
subtropical Pacific, and in Central and South America; in fact, they are still used by 
the aborigines of northern Australia.^* In the Carolines they were used on daggers." 
They have been used also as needles or as awls. In Central America, at localities well 
inland, they have been found in burial sites in numbers so great as to suggest that they 
were a regular article of trade." In Malaya, too, they have been employed as a poison," 
and along the Congo in tropical West Africa whips were made from the thorny tails 
of an African Sting Ray, and perhaps they still are.^" Pieces of the spiny skins of Rays 
(genus Urogymnus), sewed with coconut fibre around sticks of wood 6-10 inches long, 
were used by the natives of the Gilbert group "so to shape the boards of their canoes 
that when sewed together they were watertight;"-' however, this practice has now been 
replaced by the use of steel rasps. The skins of other smoother Rays have been em- 
ployed for drum heads. On the debit side, Rays do more or less damage to cultivated 
oysters in Cavite Bay, Luzon, Philippines; at high tide they swim over the bamboo 
stakes that surround the beds and crack "open the oysters so that only halves of oyster 
shells cemented to the stakes remain. "^2 And the inroads that they make on the local 
shellfish beds elsewhere must be great in regions where they are abundant. Also, they 

14. Raj, Rec. Indian Mus., jo, 1914: 317. 15- Donovan, Trans. Proc. N. Z. Inst., 52, 1920: 29. 

16. See Whitley (Fish. Aust., i, 1940: 193, fig. 222) for distribution of Australian Sting Ray spears. 

17. The late Sir P. H. Buck informed us that there is a dagger from the Carolines, with two Sting Ray spines, in the 
Bishop Museum, Honolulu, T. H. 

18. Lothrop, Peabody Mus. Mem., 7, 1937: 97-98; Merwin and Vaillant, Peabody Mus. Mem., 3, 1932: 90; Kidder, 
Amer. Antiquities, 2 (2), 1945: 68, footnote. 

19. Gimlette, Malay Poisons, ed. 1923: 117. 20. Whitley, Fish. Aust., i, 1940: 194. 

21. See Walcott (Occ. Pap. Bishop Mus. Honolulu, i [2], 1900; 32) for description and photographs. 

22. Villadolid and Villaluz, Philipp. J. Sci., 67, 1938: 394. 



Fishes of the IVestern North Atlantic 330 

are objects of concern to fishermen and to anyone else wading in the shallow water in 
regions where they are likely to be encountered, because wounds by their tail spines are 
followed almost immediately by excruciating pain,^^ by severe swelling, by violent 
muscular cramps, and often by subsequent inflammation and gangrene. Even fatalities 
have been reliably recorded. ^^ Ordinarily recovery follows without any serious com- 
plications, but in any case the mechanical damage to tissues incident on the extraction 
of the spine is likely to be considerable because of its many-barbed structure. 

Range and Bathymetric Distribution. The Whip-tailed Sting Rays are primarily a 
warmwater group; many of them are confined strictly to tropical-subtropical latitudes 
the year round; some are resident along warm-temperate coasts, and others carry out 
more or less regular and extensive migrations to higher latitudes with the vernal rise 
in sea temperatures, withdrawing again to warmer waters with autumnal cooling. Most 
of the Sting Rays are confined to shallow coastwise waters; in many parts of the world 
they commonly follow the rising tide onto mud- and sand-flats to retreat again on the 
ebb. Some occur commonly in the shoal channels and lagoons among the mangroves 
on tropical coasts, and some run up into fresh water. But others are known to occur 
as deep as 20-60 fathoms, possibly deeper. -^ 

Genera. One member of the family is set apart so sharply from all others by its 
lack of tail spines that recent students generally refer it to a separate genus, Urogymnus 
Muller and Henle 1837. A second group, consisting of a few species that are charac- 
terized by the possession of a broad fold extending along the lower surface of the tail 
to the tip, constitutes a second well marked genus, Taeniura Muller and Henle 1837. 
A separate genus, Urolophoides Soldatov and Lindberg 1930, seems also needed pro- 
visionally for an East Asian Ray that has a tail spine, an upper caudal fold but seemingly 
no lower caudal fold, and a tail much shorter than the disc; unless, indeed, the unique 
specimen that served as the basis for the description had been mutilated (see p. 340, 
footnote 29). Opinions have diff"ered, however, as to whether the 30 or more remaining 
members of the family, with long whiplash tails armed with spines and with longi- 
tudinal caudal folds (if any) terminating far short of the tip of the tail, are all referable 
to a single genus or whether the differences in degree of development of caudal folds 
and ridges deserve generic or subgeneric recognition. ^^ On the basis of our own ex- 

23. See Bassler (Science, ()6, 1942: 274) for a recent eye-witness account of the agonizing effects of a wound inflicted 
on a man's foot by the tail spine of one of the closely allied River Rays [Potamotrygonidae) of the Amazon. 

24. Thus Schomburgk (Reisen Brit. Guiana, 1840-1844, 2, 1848: 37-38) wrote that during his stay in British Guiana 
a laborer who was struck by a Sting Ray died in convulsions; Crevaux (Arch. Med. nav., 1882: 37), quoted by 
Gudger (Bull. Hist. Med., 14, 1943: 478), reported that a companion on the Orinoco died in 1881 from a wound 
by one of the river Rays; Vellard (Mem. Soc. zool. Fr., 29, 1932: 514-532) speaks of fatalities as sometimes occurring 
from wounds by Sting Rays in Brazil. And Whitley (Fish. Aust., i, 1940: 198) quotes the case of a bather's death 
following the direct penetration of the heart, almost certainly by a Sting Ray spine. 

25. Whitley (Fish. Aust., i, 1940: 201) states that the .-Australian Dasyatis bre'vicaudata is taken most often in 20-60 
fathoms. 

26. Barnard (Ann. S. Afr. Mus., jj [i], 1925: 75) and Rey (Fauna Iberica, Feces, i, 1928: 620) refer all members 
of this group to a single genus. On the other hand, Garman (Mem. Harv. Mus. comp. Zool., j6, 1913: 375) distribu- 
ted them among four named "groups" within the genus Dasybatus, which Fowler (Bull. U. S. nat. Mus., 100 
[/j], 1941: 403) ranks as subgenera of the genus Dasyatis. However, Jordan, Evermann and Clark (Rep. U. S. 
Comm. Fish. [1928], 2, 1930: 21) dignify the three groups that they mention [Paslinachus, Dasyatis, Amphotistius) 



340 Memoir Sears Foundation for Marine Research 

amination of a considerable series, we conclude that the most workable subdivision 
of the group of species in question is as follows: (a) those in which the tail bears neither 
membranous folds nor well marked longitudinal ridges either above or below (genus 
Himanturd), and (b) those in which there is a membranous fold on the lower surface 
of the tail, with or without a fold or ridge above (genus Dasyatis). 

Key to Genera 

I a. No spine on tail. Urogymnus Miiller and Henle 1837." 

Australia, East Indies, Indian Ocean, Red Sea. 
I b. Upper surface of tail with a large serrated spine (or spines). 

2a. Lower tailfold extending to tip of tail. Taeniura Miiller and Henle 1837. 

Red Sea, Indian Ocean south on African Coast 
to Delagoa £37,^' East Indies, Philippines, 
Australia, Polynesia; also Mediterranean and 
Cape Verde Islands. 
2 b. Lower tailfold either lacking or terminating far short of tip of tail. 

3a. Tail shorter than disc. Urolophoides Soldatov and Lindberg 1930. 

Peter the Great Bay, Northeast Asia; doubtful genus. ^^ 
3 b. Tail longer (usually much longer) than disc. 

4 a. Lower surface of tail, posterior to spine, without membranous fold, 
either rounded or at most with a low cutaneous ridge. 

Himantura^" Miiller and Henle 1837, p. 389. 

4 b. Lower surface of tail posterior to origin of spine with a longitudinal 

membranous fold. Dasyatis Rafinesque 18 10, p. 340. 

Genus Dasyatis Rafinesque i 8 i o 

Dasyatis Rafinesque, Carratt. Gen. Spec. SiciL, 1810: 16; type species, D.ujo Rafinesque, Sicily, equivalent 
to Raja pastlnaca Linnaeus 1758.^1 

with the rank of full genera, and Whitley (Fish. Aust., i, 1940: 200-208) recognizes four genera of long-tailed 
Rays with tail spines but without definitions. 

27. Urogymnus MuUer and Henle 1837 is antedated by Anacanthus Ehrenberg 1833 (in Van der Hoeven, Handb. Dier- 
kunde, 2, 1833: 179), but this name was preoccupied by Gray 1831 for teleost fishes and by Audinet-ServiUe 1832 
for Coleoptera. To replace ^wacawMa/ for the genus of Rays in question. Cantor (Malay. Fish., 1849: 1404) proposed 
RJiachinotus, which was accepted by Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 373) in spite of the fact 
that the older name Urogvmnus Muller and Henle 1837 (Arch. Naturg., J, 1837: 437) was expressly based on the 
same species {Raja asperrima Bloch and Schneider 1801) that was later christened Rhachinotus afrkanus by Cantor. 
The name Urogymnus was revived by Ogilby (Mem. Qd. Mus., 5, 1916: 88) and has been used by subsequent 
writers generally. 

28. Reported from Delagoa Bay by Smith (Sea Fish. S. Afr., 1949: 70). 

29. The original and only account of this Ray (Soldatov and Lindberg, Bull. Pacif. Sci. Fish. Inst., 5, 1930: 24), from 
a mounted specimen, records an upper tailfold but fails to state whether or not there was a lower tailfold. The ac- 
count of the tail as stout with blunt tip leaves open the possibility that its shortness (only about Vs ^s long as the 
disc) may have been due to mutilation, followed by subsequent healing of the wound; specimens with tails in this 
state are often seen among other species of Sting Rays. 

30. Himantura was proposed by MuUer and Henle (Arch. Naturg., 3 [i], 1837: 400) with diagnosis but without mention 
of any particular species. Dumeril (Hist. Nat. Poiss., j, 1865: 585), however, referred a considerable list of species 
to it, including Raja uarnak Forskal 1775, which was later designated as the type species by Garman (Mem. Harv. 
Mus. comp. Zool., j6, 1913: 375). 

31. Besides the synonyms listed here. Fowler (Bull. U.S. nat. Mus., 100 [jj], 1941: 402) includes the fossil genera 



Fishes of the Western North Atlantic 341 

Generic Synonyms: 

Raja (in part) Linnaeus, Syst. Nat., J, 1758: 232; for R.fastinaca Linnaeus. Europe. 

Uroxis Rafinesque, Indice Ittiol. Sicil., 18 10: 48; type species, Dasyatis ujo Rafinesque, Sicily, equivalent to 
Raja pastinaca Linnaeus 1758. 

Trygonoiattts (in part) Blainville, Bull. Soc. philom. Paris, 1816: 112; includes Raja sephen Forskal 1775. Red 
Sea.32 

Trygon Cuvier, Regne Anim., 2, 181 7: 136; type species, Raja pastinaca Linnaeus 1758, designated by Jordan 
(Genera Fish., i, 1917: 98). 

Trygonoialis Blainville, in Vieillot, Faune Fran^., Poiss., 1825: 35; type and only described species. Raja pa- 
stinaca Linneaeus 1758. East Atlantic. 

Pastinachus Riippell, Atlas Reise nordl. Afr., 2, Fische Rothen Meeres, 1828: 51, footnote; type species, Raja 
sephen Forskal 1775, Red Sea, designated by Jordan (Genera Fish., I, 1917: 122).^' 

Trigon (in part) Riippell, Atlas Reise nordl. Afr., Fische Rothen Meeres, Abt. i, 1828: 51; includes Raja sephen 
Forskal 1775. 

Hypolophus (subgenus) Miiller and Henle, Arch. Naturg., I, 1837: 400; no species named; but Raja sephen 
Forskal 1775 included as example by Bonaparte (Nuov. Ann. Sci. Nat. Bologna, 2, 1838: 202, and Mem. 
Soc. Neuchatel Sci. Nat., 2 [7], 1839: 6) and designated as type species by Jordan (Genera Fish., 2, 
1919: 193). 

Pastinaca Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1838: 192, 319; type species, P. olivacea Swainson, 
equals Raja pastinaca Linnaeus 1758. 

Hemitrygon (subgenus) Miiller and Henle, Charlesworth Mag. Nat. Hist., 2, 1838: 90; no species mentioned; 
type species, Trygon bennetti Miiller and Henle 1841, designated by Jordan (Genera Fish., 2, 1919: 190). 

Dasibatus L. Agassiz, Nom. Zool. Pise, 1845: 21; emended spelling for Dasybatus Klein 1775; ^ee p. 341, 
footnote 34. 

Dasybatis Jordan and Gilbert, Proc. U. S. nat. Mus., J, 1880: 31 ; t)'pe species, D. dipterunis Jordan and Gil- 
bert. San Diego, California. 

Dasybatus Jordan and Gilbert, Proc. LT. S. nat. Mus., 4, r88i: 35; for Dasybatis dipterunis Jordan and Gil- 
bert 1880;'* Garman, Mem. Harv. Mus. comp. Zool., j6, 1913: 375; by ref to Dasybatus Klein 1775 
and Walbaum 1792. 

Dasibatis Garman, Bull. U. S. nat. Mus., 16, 1883: 68; type species. Raja pastinaca Linnaeus 1758; alternative 
spelling for Dasybatis Jordan and Gilbert 1880. 

Taeniura Macleay, Proc. Linn. Soc. N. S. W., 7, 1883: 598; for T. atra Macleay, equals Raja sephen Forskal 

1775- 
Dasabatis Evermann and Kendall, Bull. U. S. Fish Comm., 12, 1894: 95; type species. Raja sa-^ Lesueur 1817; 

emended spelling for Dasibatis Garman 1883 or for Dasybatis Jordan and Gilbert 1880. 
Pterop/atytr\gon (subgenus) Fowler, Proc. Acad. nat. Sci. Philad., 62, 1910: 474; type species, Trygon violacea 

Bonaparte 1832. 
Amphotistius (subgroup) Garman, Mem. Harv. Mus. comp. Zool., j6, 1913'. 37;, 392; Upe species, Trygon 

sinensis Steindachner 1892, designated by Jordan, Evermann and Clark (Rep. U. S. Comm. Fish. [1928J, 

2, 1930: 28). 

Heliobatis Marsh (Amer. J. Sci., [3] 14, 1877: 256) and Xipholrygon Cope (Amer. Nat., 13, 1879: 333). But while 
the latter certainly (and the former probably) falls among the long-tailed Sting Rays, it is doubtful whether either 
of them was actually congeneric with the modern Dasyatis. 

32. This is the only identifiable species among the nine that were included by Blainville in the subdivision of his genus 
Pastinachus with a fold on the lower surface of the tail. Jordan's (Genera Fish., i, 191 7: 94) designation of Raja 
pastinaca Linnaeus 1758 as the type cannot be accepted because this name was not included in Blainville 's list of 
species for either subdivision of his Trygonobatus. 

33. The term "Pastinachae," applied by Nardo (Prod. Ittiol. Adriat., Giorn. Fisica Pavia, i, 1827: 8) to his "Sect. I" 
of Raja and for which Jordan (Genera Fish., j, 1917: 121) designated a type species (Raja pastinaca Linnaeus 1758), 
does not come into consideration as a generic name since it is not binomial. 

34. The name Dasybatus had been employed for Sting Rays previously by Klein (Neuer Schauplatz, 1775: 991) and 
by Walbaum (P. Artedi Genera Pise. Emend. Ichthyol., j, 1792: 581). But the International Commission on 
Zoological Nomenclature has ruled that neither Klein's generic names nor Walbaum's republication of them are 
available. Dasybatus Blainville 1816 and Dasybatis Blainville 1825 (see Synonyms, p. 138) are equivalents of Raja 
Linnaeus 1758 and not of Dasybatus Klein 1775. 



342 Memoir Sears Foundation for Marine Research 

Pastinachus (subgroup) Garman, Mem. Harv. Mus. comp. Zool., j6, 191 3: 375; type species, Raja fastinaca 

Linnaeus 1758. 
Bathytoshia Whitley, Rec. Aust. Mus., Jp, 1933: 61; type species, Dasyatis thettdis Waite 1899, New South 

Wales, probably equals Trygon lata Garman (Bull. Mus. comp. Zool. Harv., 6, 1880: 170).^* 
Toshia Whitley, Rec. Aust. Mus., J9, 1933: 60; type species, Dasyatis fliwiorum Ogilby 1908. Australia. 
Neotrygon Whitley, Fish. Aust., j, 1940: 208; type and only included species, Trygon kuhlii Miiller and Henle 

1 841. East Indies and India, also Philippines, South China, Australia, and western tropical Pacific. 

Probable Synonym: 

Neotngon Castelnau, Proc. Zool. Acclim. Soc. Victoria, 2, 1873: 122; type species, N. trigonoides Castelnau.^* 
Not Dasybatus Blainville, Bull. Soc. philom. Paris, 1816: 112; equals Raja Linnaeus 1758, which see, p. 138. 
Not Dasyatis Blainville, in Vieillot, Faune Fran?., Poiss., 1825: 12; equals Raja Linnaeus 1758, which see, 

p. 138. 
Not Dasibatis L. Agassiz, Norn. Zool. Pise, 1845: 21; emended spelling for Dasybatis Blainville 1825. 
Not Pastinachus Riippel, Neue Wirbelt. Abyssinia, 4, Fische, 1835: 69; equals Himantura Miiller and Henle 

1 841, which see, p. 389. 

Generic Characters. Tail much longer than disc, with a large serrate-edged spine 
(or spines) above; its lower surface, close posterior to spine, with a well developed 
longitudinal dermal fold, its upper surface either with a corresponding fold (usually- 
lower), or with a recognizable ridge," or merely rounded. Teeth small and numerous, 
closely crowded in quincunx in both sexes; those of females and of young males low, 
rounded, or nearly flat, often worn in more or less distinct ridges; those of sexually 
mature males with conical cusp in some species, perhaps in all.^* Upper tooth band 
strongly convex in cross section, the lower weakly so; both bands moderately broad 
centrally, narrowing toward corners; 6 or 7 to 10 or 12 rows of teeth exposed and in 
function simultaneously along different parts of jaws. Pelvis without median process, 
so far as known. Characters otherwise those of the family. 

Remarks. The genus, as here defined, corresponds to the subgenera Pastinachus., 
Dasyatis., and Amphotistius combined, of Garman and of Fowler.^" 

Size. This genus includes the largest known Sting Rays, namely Dasyatis brevi- 
caudata (Hutton) 1875 of New Zealand and Australia, D. aspera (Cuvier) 18 17 of the 
Mediterranean, Madeira, and tropical West Africa, and D. centroura (Mitchill) 18 15 
of the western North Atlantic (p. 352). 

Range. Tropical-subtropical coastlines in general. In the Atlantic, representatives 
of Dasyatis occur from tropical West Africa to the North Sea, Skagerrak, and western 
Baltic in the east; from Uruguay to southern New England in the west. 

35. The specimen, originally in the Harvard Museum of Comparative Zoology and from which Garman described 
this species, is no longer to be found. 

36. Castelnau states that his trigonoides from New Caledonia, for which he proposed the genus Neotrygon, has tail folds 
above and below ("dorsal" and "anal" fins) but no tail spine. But Fowler (Bull. U. S. nat. Mus., 100 [if\, 1941: 
426) has relegated trigonoides to the synonymy of Dasyatis kuhlii (Muller and Henle, 1841), which has well devel- 
oped spines. If this allocation be correct, as appears to be the case, Neotrygon Castelnau 1873 is clearly a synonym 
of Dasyatis Rafinesque 1810, as defined here, as is Neotrygon Whitley 1940. 

37. In preserved specimens this ridge sometimes becomes so obscured, perhaps through muscular contraction, that 
the upper surface of the tail behind the spine is merely arched or simply rounded. 

38. The transition from the flat or rounded teeth in older (anterior) rows to cuspidate teeth in younger (more posterior) 
rows may often be seen on males nearing sexual maturity. 

39. Garman (Mem. Harv. Mus. comp. Zool., 36, 1913: 373) termed them subgroups; Fowler (Bull. U. S. nat. Mus., 
100 [/j], 1941: 404), subgenera. 



Fishes of the IVestern North Atlantic 343 

Species. Recent surveys of the genus, as here defined, for various parts of the 
world recognize a total of some 30 species." But it is likely that a general revision 
would result in a considerable reduction in this number. Six species are known from 
the western Atlantic (see Key, p. 343). 

The representatives oi Dasyatis and oi Himantura in the western Atlantic are more 
readily identifiable by the proportionate dimensions of disc and tail and by the shape 
of the disc than they are by the degree of development of the longitudinal ridges or 
folds on the tail, though these last characters are more significant from the generic 
standpoint. The local members of the two genera are therefore grouped together in 
the following Key for convenience in identification. 

Key to North Atlantic, Western South Atlantic and Tropical West African 
Species of Dasyatis and Himantura*^ 

I a. Anterior contour of median sector of disc either a uniform arc of long radius or 
nearly straight. 

2 a. Distance from center of cloaca to origin of tail spine nearly as long as dis- 
tance from center of cloaca to tip of snout; entire upper surface of disc closely 
clothed with small tubercles; tail without folds above or below. 

Himantura schmardae (Werner) 1904, p. 390. 

2 b. Distance from center of cloaca to origin of tail spine much shorter than dis- 

tance from cloaca to snout; upper surface of disc mostly naked; lower surface 
of tail with well developed longitudinal cutaneous fold. 

Dasyatis violacea (Bonaparte) 1832. 
Mediterranean. 
I b. Anterior contour of median sector of disc subangular, with tip of snout forming 
the apex. 

3 a. Entire upper surface of disc rough with minute dense asperities. 

Dasyatis rudis (Giinther) 1 8 70. 
Old Calabar, Equatorial West Africa.*'^ 

3 b. Upper surface of disc mostly smooth except along median belt, and here or 
there elsewhere. 

4 a. Distance from level of axils of pectorals to origin of tail spine about as 
long as distance from level of axils of pectorals to rear margins of orbits; 
center of back with a single large white tubercle. 

Dasyatis margarita (Giinther) i 8 70. 
Tropical West Africa. 

40. Garman, Mem. Harv. Mus. comp. Zool., j6, 1913: 381, 388, 392; Fowler, Bull. Amer. Mus. nat. Hist., 70 (i), 
1936: 125; Bull. U.S. nat. Mus., 100 (/j), 1941: 413, 420, 421. 

41. China and Trinidad are given as localities for Dasyatis bennetti by Muller and Henle (Plagiost., 1841: 160), its 
describers. But recent authors have restricted the name to the Pacific form, hence it is not included in the Key. 
For references, see Fowler (Bull. U.S. nat. Mus., loo [/j], 1941: 413). 

42. Giinther, Cat. Fish. Brit. Mus., 8, 1870: 479. Thus far D. rudis appears to be known from the original account 
only, which was based on a single large specimen 6^/2 feet broad. A more detailed account of this species is much 
to be desired, especially as regards its tailfolds, if any. 



3 44 Memoir Sears Foundation for Marine Research 

4 b. Distance from level of axils of pectorals to origin of tail spine consider- 
ably shorter than distance from level of axils of pectorals to rear margins 
of orbits; center of back without a single large white tubercle. 
5 a. Outer corners of disc broadly and evenly rounded. 

6 a. Pelvic fins with narrowly pointed outer corners, their anterior 
margins about 2.5 times as long as distance between outer 
margins of orbits. Dasyatis geijskesi Boeseman 1948, p. 363. 
6 b. Pelvic fins with broadly rounded outer corners, their anterior 
margins little, if any, longer than distance between outer mar- 
gins of orbits. 

7 a. Snout anterior to orbits considerably longer than distance 
between spiracles; anterior contour of disc concave on 
either side of tip of snout. 

Dasyatis sahitia (Leseuer) 1824, p. 370. 
7 b. Snout anterior to orbits shorter than distance between 
spiracles; anterior outlines of disc weakly convex on either 
side of tip of snout. Dasyatis say (Leseuer) 18 17, p. 378. 
5 b. Outer corners of disc only narrowly rounded or abruptly sub- 
angular. 

8 a Anterior margins of disc continuously though weakly concave 
anterior to level of posterior margins of spiracles; tip of snout 
projecting from general contour in subtriangular outline 
(Fig. 86). 

9 a. Posterior parts of pelvics extending rearward beyond poste- 
rior limits of pectorals for a distance as great as longitu- 
dinal diameter of spiracle; disc naked except for tubercles 
along midline of back and on shoulders of large specimens. 
Dasyatis pastinaca (Linnaeus) 1758. 
Eastern North Atlantic to North Sea, Skagerrak 
and western Baltic; Mediterranean; south to 
South Africa; also reported from India and 
East Africa south to Agulhas Bank.'*^ 
9 b. Posterior parts of pelvics extending only very little rear- 
ward beyond level of posterior limits of pectorals; mid- 
belt of disc, rearward from interocular region, densely 
clothed with small rounded tubercles on half-grown and 
larger specimens. 

Dasyatis guttata (Bloch and Schneider) 1801, p. 365. 
8 b. Anterior margins of disc nearly straight, or at most weakly 
sinuous, anterior to level of posterior edges of spiracles; tip of 
snout not projecting conspicuously from general anterior con- 
tour (Figs. 81, 83). 

43. It is not yet certain whether the Ray reported under this name from the Indian Ocean is identical with D. pastinaca 
of the Mediterranean and eastern Atlantic. 



Fishes of the Western North Atlantic 345 

loa. A longitudinal cutaneous fold along lower side of tail 
about as wide as height of tail ; upper surface of tail with 
a low longitudinal ridge close posterior to spine, or at 
least arched in cross section; sides of tail without con- 
spicuous tubercles. 

Dasyatis americana Hildebrand and Schroeder 
1928, p. 345. 
lob. A longitudinal cutaneous fold along lower side of tail only 
about half as wide as height of tail ; upper surface of tail 
posterior to spine rounded, without longitudinal ridge; 
sides of tail, in half-grown specimens and larger, rough 
with conspicuous tubercles or thorns. 

Dasyatis centroura (Mitchill) 18 15, p. 352, and 
Dasyatis aspera (Cuvier) 18 17. 
Eastern Atlantic, tropical West Africa, Madeira, and Med- 
iterranean.''* 



Dasyatis americana Hildebrand and Schroeder 1928 
Figures 81, 82 

Study Material. Fifteen specimens, 126—525 mm wide, including embryos, from 
Brazil, Trinidad, Puerto Rico, Cuba, Bahamas, Gulf of Campeche, Yucatan, Texas, 
Florida, and Chesapeake Bay, in Museum of Comparative Zoology and U. S. National 
Museum. 

Distinctive Characters. D. americana, regardless of size, is separable at a glance 
from D. sabina, D. guttata and Himantura schmardae by the shape of the anterior part 
of its disc (cf. Fig. 81 with 86, 88, 93). It is separable from H. schmardae also by the 
position of its spine well forward on the tail (cf. Fig. 81 with 93). It is distinguished 
further from D. guttata by the fact that the armature on the dorsal crest of its disc on 
either side of the midline of tubercles consists only of minute prickles. If the tail is 
intact, D. americana has only a low ridge on its upper surface, whereas D. say has a 
well developed fold there. Specimens of D. americana that have lost their tails are 
separable from D. say by the fact that both the outer and the posterior corners of 
their discs are much more abrupt than those of the latter (cf. Fig. 8 i with 90). The 
relatively great breadth of the lower tailfold of D. americana (cf. Fig. 8 i A with 83 A), 
plus the presence of a ridge or keel on the upper surface of the tail posterior to the 
spine, marks it off from D. centroura. Apart from the size, a tailless D. americana, prior 
to the appearance of the mid-dorsal thorns, resembles a young D. centroura so closely 
that we have been unable to find any dependable criteria by which to distinguish the 
one from the other. Even so, the danger of confusing them is small, for D. centroura 

44. It is still an open question whether or not D. centroura of the western Atlantic and D. aspera of the eastern are 
separable; for discussion, see p. 357. 



34^ Memoir Sears Foundation for Marine Research 




Figure 8i. Dasfatis americana, male, 505 mm %vide, from Molasses Key, Florida (Han-. Mus. Comp. ZooL, 
No. 367). A Side view of midsector of tail to slightly larger scale. B Tip of tail spine, about 3.9 X - C Midsector 
of tail spine, about 3.9 X. T> Cross section of tail spine about mid%vay of its length, about 3.9 X. 

Still is smooth-skinned to a size considerably greater than that at which £). americana 
commences to show its characteristic mid-dorsal thorns. 



Fishes of the Western North Atla?itic 347 

Description. Proportional dimensions in per cent of extreme breadth of disc. Male, 
364mm broad, from Key West, Florida (U.S.Nat. Mus., No. 125797). Female, 
527 mm broad, from Pernambuco, Brazil (Harv. Mus. Comp. Zool., No. 246). 
Bisc: vertical length 86.2, 85.5. 

Snout length: in front of orbits 18.4, 17.1; in front of mouth 20.3, 18.6. 
Orbits: horizontal diameter 8.0, 5.7; distance between 9.3, 8.0. 
Spiracles: length 7.1, c^.'^\ distance between 17.0, 14.2. 
Mouth: breadth 8.2, 8.9. 

Exposed nostrils: distance between inner ends 10.7, 10.4. 
Gill openings: lengths, ist 3.0, 2.7; 3rd 3.0, 2.7; 5th 2.3, 1.9; distance between 

inner ends, ist 18.6, 19.4; 5th 11. 4, 11. 9. 
Pf/i'/V^; anterior margin 17.0, 15.6. 

Distance: from tip of snout to center of cloaca 78.8, 74.1 ; from center of cloaca 
to origin of caudal spine 37.1, 33.8. 

Disc about 1.2 times as broad as long (specimens i 5 to 60 in. wide), rhomboid; 
tip of snout subangular, projecting at most only a little or not at all; maximum anterior 
angle in front of spiracles about 135°; anterior margins varying from nearly straight 
to slightly concave or weakly convex anterior to level of eyes, becoming increasingly 
convex toward outer corners, the latter being either abruptly rounded or subangular; 
posterior margins nearly straight anteriorly but becoming weakly convex posteriorly; 
posterior corners abrupt, much as in D. centroura\ inner posterior margins straight or 
weakly convex. Axis of greatest breadth about 38— 4070 of distance rearward from 
tip of snout toward rear limits of disc. Tail moderately depressed dorsoventrally and 
evenly rounded laterally anterior to spine; its upper surface posterior to spine with a 
low fleshy keel extending rearward nearly or quite as far as rear termination of lower 
tailfold on most specimens, but rounded on some;*^ lower surface of tail with a more 
or less conspicuous longitudinal fold originating about under origin of tail spine and 
extending rearward for a distance about 1.2— 1.4 times as long as distance from axils 
of pectorals to spine; nearly uniformly wide throughout most of its length but nar- 
rowing at either end to merge with general contour of tail; its maximum width at 
spine about equal to or greater than height of tail on some large specimens, but only 
about half as high as tail on others and on embryos. Tail from center of cloaca 1.9 (small 
specimen) to 2.3 times as long (large specimen) as distance from center of cloaca to 
tip of snout, if not damaged; distance from origin of tail spine to center of cloaca about 
45 "/o as long as distance from cloaca to snout. 

Length of free portion of tail spine (one only on specimens seen) about as great 
as distance between outer margins of orbits; total length of spine from origin about 
1.3 times that great; its lateral teeth 52 and 80 on each side in two specimens counted,^* 
often covered with skin to tip even on large specimens. 

45. We have examined one such specimen. 

46. These spines were not damaged. The specimen whose tail spines had 52 teeth was 36 inches wide, the one with 
80 teeth was 60 inches wide. 



348 



Memoir Sears Foundation for Marine Research 



Midline of back with a row of ridge-like tubercles, truncate posteriorly, on smooth 
oval bases; partly-grown specimens (375—425 mm wide) with about eight of these 
from nuchal region rearward, followed next by a considerable gap, then by two or three 
tubercles close in front of tail or on its base; mature specimens with the median row 
numbering 30 or more,*' continuous or nearly so though irregularly spaced; each 
shoulder also developing a longitudinal series of smaller tubercles, from 2—3 at first 




Figure 82. Dasyatis americana. A Nostrils, nasal curtain and mouth of male pictured in Fig. Si. B Detail of 
margin of nasal curtain from right-hand side, about 1.3 X. C Detail of margin of nasal curtain from left-hand 
side, about 1.3 X. D Left-hand half of upper tooth band, about 3.3 X. E Right-hand half of upper tooth band 
of female, 448 mm long, from Rio de Janeiro, Brazil (Harv. Mus. Comp. ZooL, No. 571), about 3-3 X. 

to 10—12 at maturity; both medians and scapulars larger and more conspicuous on 
females than on males; areas between orbits and between spiracles sparsely prickly on 
males by maturity, more densely so on females, also prickly on nuchal and scapular 
regions; a belt along either side of the median row of tubercles more or less prickly 
on medium-sized specimens (about 40 in. wide), the largest (about 60 in. wide) with 
prickles covering entire disc; upper and lateral surfaces of tail close in front of spine 
and posterior to it rough with prickles on large specimens, but smooth farther forward; 
pelvics smooth in both sexes. Lower surface smooth on disc and on tail anterior to spine. 
Snout in front of orbits about as long as distance between outer edges of orbits, 
its length in front of mouth 1.8-2.0 times as great as distance between exposed nostrils 

47. Sixty have been reported for a Cuban specimen (1,058 mm wide) that may have been of this species, by Fowler 
(Fish Culturist, 21, 1942: 66 as "Dasyatis scabrata"); see also p. 352, footnote 64. 



Fis/ies of the Western N^orth Atlantic 349 

and about equal to distance between inner ends of first gill openings. Eye about 75- 
85 7o ^s long as distance between eyes and about equal to length of spiracle. Distance 
between inner ends of first gills about 1.7— 1.9 times as long as distance between ex- 
posed nostrils; distance between inner ends of fifth gills about 62 "/o ^s great as that 
between inner ends of first gills. Free posterior margin of nasal curtain weakly con- 
cave, fringed, the individual lobes simple or variously divided, those of the two sides 
separated in midline by a narrow gap. Mouth with upper jaw moderately projecting 
centrally and lower jaw moderately recessed, a little more so on mature males than on 
females. Floor of mouth, centrally, with a transverse series of three stout papillae, 
sometimes with another more slender papilla on one side or on both sides. 

Teeth 3' to 5*,^* about equally large all along each tooth band except for some- 
what smaller ones near outer corners; those of females and immature males tetragonal 
with rounded corners, a little broader (transversely) than long (anteroposteriorly), the 
functional surfaces weakly rounded or flat, often scored ; those of sexually mature males 
with low conical cusp; 6—8 rows in function simultaneously in upper jaw and about 
12 rows in lower jaw near center but only 8—9 rows near outer corners. 

Anterior margins of pelvics nearly straight on small specimens, usually more 
convex on larger; outer corners moderately rounded, the posterior corners usually 
more broadly rounded but varying widely in degree of curvature among large spe- 
cimens; anterior margin of pelvic about as long as distance from its own origin to rear 
tip of longest ray of pelvic. 

Color.*^ Upper surface of disc gray, dark or olivaceaous brown, or olive green, 
depending on the color of the bottom on which the Ray is lying; sometimes darker 
toward outer margins and narrowly edged with white; a gray or whitish spot on median 
line of snout close in front of eyes; sides of anterior part of tail grayish or whitish, its 
dorsal keel, ventral fold, and terminal portion black or brownish black. Lower surface 
of disc white or whitish with an edging of gray or brown, either solid or broken into 
irregular spots; inner posterior portions of pelvics so marked also. Tail spine described 
as dark on fresh specimens. 

Size. Embryos nearly ready for birth are about 126 (see Study Material) to 170— 
180 mm broad, suggesting a width not far from 200 mm as usual at birth. It is probable 
that females may mature at a breadth of 750—800 mm, possibly smaller, for a North 
Carolina specimen 828 mm wide (33 in.) gave birth to three young on capture. Males 
may mature at a somewhat smaller size, for one about 510 mm wide (20 in.) has been 
reported as adult ; furthermore, the claspers of another of about that same size are so 
large as to suggest the imminence of sexual activity. The largest specimens of D. ameri- 
cana positively recorded have been: 40, 42, 44, and 60 inches in width, all of these 
recently examined by us at Bimini, Bahamas. One about 37 inches wide from the 
Tortugas, Florida weighed 58 pounds. =" 

48. The smallest specimen which we counted, with disc 14 inches wide, had 39 series in each jaw; the largest, 60 inches 

wide, had 56. 
4.Q. We have no color notes from life. 50. Gudger, Aust. Mus. Mag., 6 (6), 1937: 208. 



350 Memoir Sears Foundation for Mari?ie Research 

Developmental Stages. Embryos nearly ready for birth closely resemble their parents 
except that the lower tailfold is relatively narrower. A female gave birth to three young 
on capture. ^^ Other females that gave birth to four or five, one of them also containing 
15 eggs of various sizes, likewise have been referred to this species. ^'^ Nothing further 
is known of its breeding habits. 

Habits. All recorded captures of this Ray have been from partially enclosed waters, 
or at least from close inshore. The fact that all dated records of it from North Carolina 
to New Jersey have been during the months of July, August and early September 
shows that it is a summer visitor to the northern part of its range. But reports of its 
northerly presence have not been numerous enough for us to establish the approximate 
dates of its vernal arrival or of its autumnal departure, or to afford a correlation between 
its migrations and the seasonal cycle of temperature. 

The stomachs of Florida specimens ^^ and of others collected near Beaufort, North 
Carolina, contained clams, blue crabs {Callinectes), shrimps, worms, and small bony 
fishes. Specimens which we opened at Bimini, Bahamas, contained stomatopods, 
shrimps, crabs, worms, and fish. Further information as to its way of life is confined 
to an observation among the Florida Keys that they are usually seen gliding in pairs 
close to the sandy bottom from which it is difficult to distinguish them, and that they 
swim away rapidly upon the approach of a small boat.^* They are reported as being 
sometimes taken on hook and line as well as in seines on the bottom around Trinidad. 
And the fact that a specimen, referred to this species, towed a 22-foot launch for 15 
minutes '^^ illustrates the power with which the larger Sting Rays drive ahead when 
disturbed. 

Numerical Abundance. No precise information is available as to its actual abundance 
anywhere except that it is common on the coasts of Mississippi and western Florida,^^ 
that 20 were taken in the vicinity of Beaufort, North Carolina, in 1936 between July 30 
and August 16,^' and that we saw five specimens brought in at Bimini by one fishing 
boat on one day in January 1949. 

Relation to Man. Adult individuals, being even greater in size than Dasyatis say., 
are doubtless as great a menace as the latter to anyone haneiling or treading upon them, 
but no specific reports have been received of injuries inflicted by this particular Ra