Fishes of the Western North Atlantic
LA ELL a
This series of volumes, several of which are now available, presents authorita-
tive studies of the anadromous, estuarine, and marine fishes presently known
to frequent the western North Atlantic from Hudson Bay southward to the
Amazon. For many years to come, these studies will rank as primary references
for both amateur and professional persons interested in fishes and as significant
working tools for students of the sea.
Shortly after Part One was published, the eminent ichthyologist, Carl
Hubbs, said, “The first volume of Fisnes or THE WesTERN Nortu ATLAN-
ric sets a very high standard—perhaps so high a standard as to render diffi-
cult the completion of subsequent volumes by authors less well equipped than
Henry Bigelow and associates in courage, energy, time, meticulousness, ex-
perience, library facilities and willingness to sacrifice much else for this one
grand task. ... In several ways this volume has been successfully adapted,
in line with the policy set for the series, for the use and interest of sportsmen
and general naturalists as well as ichthyologists. Features that lead to this
desirable end ... include the excellent summaries of natural history informa-
tion, the limited treatment of internal and particularly microscopical anatomy,
the simplified keys, the complete coverage of the species in the clear-cut
illustrations.”
The over-all arrangement accords with our present knowledge of system-
atics, and each account is comprehensive in treatment, excellently illustrated,
fully documented, and completely indexed. In addition to the descriptive
details and discussion of orders, suborders, families, genera, and species, the
keys and synopses relate distinguishing characters that facilitate identification.
Details, aside from the description itself, include (when known) the color,
size, development, habitat, general habits, food, parasites, predators, sporting
qualities, range, abundance, and commercial importance. The accurate de-
tailed illustrations, which aid particularly in identification, have been pre-
pared by competent zoological artists, and for many species the indispensable
illustration of the adult is supplemented by figures of developmental stages
and by enlarged drawings of diagnostically important characters.
This series is especially useful as it brings together much information
usually found only in many inaccessible publications. The annotated References
direct the interested reader to more detailed information.
These volumes are published by the Sears Foundation at Yale University,
with an editorial board composed of prominent students of the sea. Contribu-
tors of the individual sections have been selected as recognized authorities on
their various groups.
iimuumny
MBL/WHO!I
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Fishes of the
Western North Atlantic
Part Three of Fisnzs oF THE WeEsTERN NorTH
ATLANTIC continues to provide the comprehensive
and informative accounts of fishes that distinguish
the first two volumes in this series.
This volume includes studies of nearly 100
species in 35 genera of “Bony Fishes.”
Each account combines a critical review of pub-
lished knowledge with original study and research,
_ resulting in distinctive contributions to ichthyologi-
cal literature not found elsewhere. These accounts,
following precedent, have been written with the
purpose of being useful to both amateur and pro-
fessional ichthyologist, to people casually or vitally
interested in the general phenomena of marine life,
to sportsmen whose objectives are pleasure and
relaxation, and to fishermen whose livelihood de-
pends upon a knowledge of fishes.
The contributions on the sturgeons and gars
bridge the gap between studies of the cartilaginous
fishes, treated in Parts One and Two, and studies
of the soft-rayed bony fishes that follow. To the
sportsman, the widely renowned gamefish—salmon,
trout, tarpon, ladyfish, and bonefish—will be of
particular interest. To the amateur and professional
fisherman, the shads, anchovies, menhaden and
other herrings, sardines, charrs, smelt, and capelin
will be of particular interest. To the ichthyologist
and other students of marine biology, all contribu-
tions in this and other volumes will be essential
adjuncts to future research and study.
The contributions in this volume were prepared
by the following well-known students of ichthyo-
logy: H.B. Bigelow, M.G. Bradbury, J. R. Dy-
mond, J. R. Greeley, the late S. F. Hildebrand, G.
W. Mead, R. R. Miller, L. R. Rivas, W. C. Schroe-
der, R. D. Suttkus, and V. D. Vladykov.
Published by the Sears Foundation for Marine
Research at the Bingham Oceanographic Labora-
tory, Yale University.
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Fishes oF THE WestTeRN Nortu Attantic. Lan-
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Isabel Pérez Farfante, and W. C. Schroeder. 576
pages, including 106 illustrations and 2 maps.
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FisHEs OF THE WESTERN NortH ATLANTIC. Saw-
fishes, Guitarfishes, Skates, Rays, and Chimaeroids,
by H. B. Bigelow and W. C. Schroeder. 588 pages,
including 134 illustrations and 2 maps.
$ 15.00
MEMOIR I, PART 3
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geons, Gars, Tarpon, Ladyfish, Bonefish, Salmon,
Anchovies, Herrings, Shads, Charrs, Smelt, Capelin,
et al. by numerous authors. 597 pages, including
134 illustrations and 2 maps.
$ 27.50
MEMOIR II
Tue ELEMENTARY CHEMICAL ComPosITION oF Ma-
RINE Orcanisms, by A. P. Vinogradov, translated
by Julia Efron and Jane K. Setlow, with complete
Bibliography by Virginia Odum. 647 pages, in-
cluding 327 tables,
$ 17.00
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MEMOIR
SEARS FOUNDATION FOR MARINE RESEARCH
Number I
Fishes of the
Western North Atlantic
eee
PART THREE
Soft-rayed Bony Fishes
Class Osteichthyes
Order Acipenseroidei, Order Lepisostet
Order Isospondyli, Suborder Elopoidea
Suborder Clupeoidea, Suborder Salmonoidea
NEW HAVEN 1963
SEARS FOUNDATION FOR MARINE RESEARCH, YALE UNIVERSITY
PRINTED IN DENMARK FOR
SEARS FOUNDATION FOR MARINE RESEARCH
BINGHAM OCEANOGRAPHIC LABORATORY
YALE UNIVERSITY
Yngve H. Olsen, Editor
Bianco Luno’s Printing, Copenhagen, Denmark
Fishes of the
Western North Atlantic
Authors
HENRY B. BIGELOW
Museum of Comparative Zoology
MARGARET G. BRADBURY
Hopkins Marine Station
JOHN R. DYMOND
University of Toronto
JOHN R. GREELEY
New York Conservation Department
SAMUEL F. HILDEBRAND
U.S. Fish and Wildlife Service
GILES W. MEAD
Museum of Comparative Zoology
ROBERT R. MILLER
The University of Michigan
LUIS R. RIVAS
University of Miami
WILLIAM C. SCHROEDER
Woods Hole Oceanographic Institution
ROYAL DY SUTTKUS
Tulane University
VADIM D. VLADYKOV
University of Ottawa
NEW HAVEN, 1963
ee eee Se Ss ee eS eee
SEARS FOUNDATION FOR MARINE RESEARCH, YALE UNIVERSITY
Editortal Board
Editor-in-Chief HENRY B. BIGELOW Museum of
Comparative Zoology, Harvard University
CHARLES M. BREDER
American Museum of Natural History
New York
DANIEL M. COHEN
U.S. Fish and Wildlife Service
Washington, D.C.
GILES W. MEAD
Museum of Comparative Zoology
Harvard University
DANIEL MERRIMAN
Bingham Oceanographic Laboratory
Yale University
vi
YNGVE H. OLSEN
Bingham Oceanographic Laboratory
Yale University
WILLIAM C. SCHROEDER
Museum of Comparative Zoology
Harvard University
LEONARD P.SCHULTEZ
United States National Museum
Washington, D.C.
JOHN TEE-VAN
New York Zoological Soctety
Bronx, N.Y.
Table of Contents
PREFACE Xill
INTRODUCTION XVII
Maps
BONY FISHES
Superclass, Class, Subclasses, and Orders, by H. B. Biczrtow I
Superclass Pisces 2
Class Osteichthyes 2
Key to Orders 16
Text and Footnote References 18
Names of Bones, by G. W. Mead and Margaret G. Bradbury 20
Order Acipenseroidei, by V. D. Vladykov and J. R. Greeley 24
Family Acipenseridae 26
Genus Acipenser 28
Acipenser brevirostris 36
Acipenser fulvescens 41
Acipenser oxyrhynchus oxyrhynchus 46
Acipenser oxyrhynchus desotoi 56
Text and Footnote References 59
Order Lepisostei, by R. D. Suttkus 61
Family Lepisosteidae 68
Genus Lepisosteus 69
Lepisosteus oculatus ah!
Lepisosteus osseus 75
Lepisosteus platyrhincus 80
Lepisosteus spatula 83
Text and Footnote References 87
Order Isospondyli, Composite Authorship
Characters and Keys to Suborders and Families, by H.B. Bigelow 89
Key to Suborders 95
Key to Families of Isospondyli, Iniomi, and
Giganturoidei 97
Text and Footnote References 105
Suborder Elopoidea, Composite Authorship
Characters and Key to Families, by H. B. Bigelow 107
Text and Footnote References 109
vil
Vill Table of Contents
Editorial Comments on the Hildebrand Manuscripts
Family Elopidae, by S. F. Hildebrand
Genus Tarpon
Tarpon atlanticus
Genus Elops
Elops saurus
Family Albulidae
Genus A/bula
Albula vulpes
Genus Dixonina
Dixonina nemoptera
Text and Footnote References
Suborder Clupeoidea, Composite Authorship
Characters and Key to Families, by H. B. Bigelow
Text and Footnote References
Family Engraulidae, by S. F. Hildebrand
Genus Axnchovia
Anchovia clupeoides
Anchovia nigra
Genus Anchoa
Anchoa spinifer
Anchoa argenteus
Anchoa duodecim
Anchoa trinitatis
Anchoa cayorum
Anchoa pectoralis
Anchoa mitchilli
Anchoa parva
Anchoa januaria
Anchoa choerostoma
Anchoa tricolor
Anchoa cubana
Anchoa ginsburgi
Anchoa lamprotaenia
Anchoa hepsetus
Anchoa lyolepis
Anchoa filifera
Genus Anchoviella
Anchoviella eurystole
Anchoviella estauquae
Anchoviella cayennensis
Anchoviella perfasciata
I1O
Tat
110)
113
123
124
132
133
134
143
143
146
148
IS1
152
ras
155
158
162
167
169
170
npn
173
174
176
181
183
185
186
188
190
192
194
200
202
204
208
Oh a0is
Oi)
213
Table of Contents ix
Anchoviella brevirostris 219
Anchoviella guianensis 217
Anchoviella alleni 219
Anchoviella nattereri 220
Anchoviella pallida 221
Anchoviella elongata 10'3
Anchoviella lepidentostole 224
Anchoviella blackburni 226
Anchoviella (Amplova) jamesi 225,
Genus Prerengraulis 228
Prerengraulis atherinoides 229
Genus Hildebrandichthys 230
Hildebrandichthys setiger i
Genus Lycengraulis 233
Lycengraulis grossidens 234
Lycengraulis limnichthys 237
Lycengraulis batesii 240
Lycengraulis abbotti 242
Lycengraulis barbouri 243
Genus Cetengraulis 24.5
Cetengraulis edentulus 245
Cetengraulis juruensis 247
Text and Footnote References 249
Family Alepocephalidae (Interim Account), by H. B. Brcrtow 250
Text and Footnote References 253
Family Searsiidae (Interim Account), by H. B. Bicztow 254
Text and Footnote References 256
Family Clupeidae, by S. F. Hitpesranpb; except
Genus Harengula by L. R. Rivas, and
Genus Dorosoma by R. R. MILteR 257
Genus Etrumeus 262
Etrumeus sadina 263
Genus Fenkinsia 267
Fenkinsia lamprotaenia 268
Fenkinsia viridis ie,
Genus Clupea 274
Clupea harengus 275
Genus Alosa 293
Alosa sapidissima 295
Alosa alabamae 308
Genus Pomolobus 312
Pomolobus chrysochloris 315
Table of Contents
Pomolobus mediocris
Pomolobus aestivalis
Pomolobus pseudoharengus
Genus Brevoortia
Brevoortia tyrannus
Brevoortia brevicaudata
Brevoortia patronus
Brevoortia smithi
Brevoortia gunteri
Genus Opisthonema
Opisthonema oglinum
Genus Harengula, by L. R. Rivas
Harengula humeralis
Harengula clupeola
Harengula pensacolae
Genus Sardinella
Sardinella pinnula
Sardinella anchovia
Sardinella brasiliensis
Genus Rhinosardinia
Rhinosardinia amazonica
Rhinosardinia serrata
Genus J/isha
Llisha altamazonica
llisha castelnaeana
Ilisha amazonica
llisha harroweri
Ilisha narragansetae
Genus Pristigaster
Pristigaster cayana
Genus Odontognathus
Odontognathus mucronatus
Odontognathus compressus
Genus Neoopisthopterus
Neoopisthopterus cubanus
Genus Chirocentrodon
Chirocentrodon bleekerianus
Genus Dorosoma, by R. R. Mitter
Dorosoma cepedianum
Dorosoma petenense
Text and Footnote References
319
324
332
342
346
363
365
372
376
380
381
386
387
390
393
397
399
401
407
411
412
413
415
417
419
421
423
426
427
428
430
431
433
435
436
438
439
443
444
448
452
Table of Contents x1
Suborder Salmonoidea, Composite Authorship
Characters and Key to Families, by H. B. Brcztow 455
Family Salmonidae, by J. R. Dymonp 457
Genus Salmo, by J. R. Dymonp 459
Salmo salar 460
Salmo trutta 498
Salmo gairdneri 499
Text and Footnote References 500
Genus Sa/velinus, by H. B. BrcrLow 503
Salvelinus alpinus 507
Salvelinus fontinalis pals
Genus Cristivomer, by H. B. BicELow $42
Cristivomer namaycush 543
Text and Footnote References 545
Family Coregonidae, by H. B. BicELow 547
Genus Coregonus 549
Coregonus clupeaformis 549
Text and Footnote References 552
Family Osmeridae, by H. B. Bicrtow and W.C. ScuroEDER = 553
Genus Osmerus 554
Osmerus eperlanus mordax 559
Genus Mallotus 573
Mallotus villosus villosus 576
Text and Footnote References 595
InpEx or Common Names 599
InDEx oF ScrENTIFIC NAMES 606
-*
oBa a ss gehes 502515122782
Preface
HE SYSTEMATIST has divided the Superclass Pisces into two major groups:
the elasmobranchs with an internal supporting skeleton of cartilage only, and bony
fishes with a skeleton consisting of true bone in greater or lesser amount, or some deri-
vation of it. Parts 1 and 2 of this series have already treated the sharks, skates, and rays,
which comprise the elasmobranchs. With publication of Parts 3—5, this series enters the
realm of bony fishes, by far the larger and more diverse of the two groups. The fishes
treated here, commonly referred to as “‘soft-rayed bony fishes,” include the anadromous
sturgeons and gars, the wide-ranging and numerous groups of herring-like isospondylids,
the little-known giganturoids, the broad assemblage of iniomes, and finally the strange-
looking lyomerids. The Introduction and Table of Contents for each volume provide
detailed information as to the subjects treated by the various authors.
The diverse shapes and characters of the forms detailed in these volumes, well
known to the ichthyologist and many students of oceanography, constitute a notable
display of the devious developments and modifications that have occurred in the course
of evolutionary changes throughout time. This is also quite apparent to the average
reader from a mere glance at the illustrations. Certainly none among nature’s multi-
plicity of creatures is more bizarre than the oddly shaped hatchet fish with its tubular
eyes and luminous organs, or the pelican fish with its long eel-like body and an enormous
mouth that allows it the unusual ability to devour prey larger than itself. Indeed, so
unusual and diverse are many of the groups treated here that even the trained system-
atist is put to it even to approximate their correct position on the ichthyological tree.
Parallel with their diverse development in form, we also find pronounced modifications
in their mode of life and habitat, as in the anadromous sturgeons and gars who have
assumed the capacity to live in both fresh and salt water, and the deep-sea creatures
who have adjusted to life at great depths under tremendous pressure where no sunlight
penetrates.
In Parts 3-5, as in the two previous volumes, our objectives have remained the
same—to provide for both professional and nonprofessional readers “critical reviews and
revisions of each group rather than perfunctory compilations or mere reprintings of
previously published works.” The extent to which these objectives are fulfilled in each
section will be determined by the usefulness of the account to both groups of readers.
Xili
X1V Preface
With multiauthorship, instead of dual authorship as in the case of Parts 1 and 2, and
with wide variation in both subject matter and available material, treatment and em-
phasis vary extensively according to authorship. All of the papers in these three Parts
have been edited by me to a greater or lesser extent, except the Family Sternoptychidae
in Part 4, which is published as it was submitted by the Editors-in-Chief.
All of these volumes should prove to be of considerable value in one way or an-
other to the ichthyologists and to other students of the sea. To the average reader, and
to sport and commercial fishermen in particular, Part 3 will be of special interest, for
here is contained a large accumulation of facts about the tarpon, ladyfish, bonefish, shad,
salmon, and trout, all of which are sportsmen’s favorites, and about the sturgeon, an-
chovy, herring, pilchard, sardine, smelt, menhaden, and shad, many of which are im-
portant to man either directly as table food or indirectly as food for the larger in-
habitants of the sea that are of economic value to man.
As in the case of Parts 1 and 2, the geographic area for these studies has been
arbitrarily defined as that extending from the vicinity of Hudson Bay, just below the
fringe of the Arctic, southward to the tropical latitudes of the Amazon, and seaward
more or less to the mid-Atlantic, including Bermuda and all of the Caribbean archipel-
ago. Although most of the accounts in these volumes treat fishes that spend at least
part of their life-span in waters ranging from brackish to maximum salinities, a few,
known at present from fresh water only, have been included when, in the author’s
opinion, they are likely to be taken in brackish or saline situations at some future date.
The format, general organization, and arrangement of the included material have
been modified from Parts 1 and 2 only insofar as the subject matter and the demands
of economy dictated. In Parts 1 and 2 there was excessive footnoting, and this has been
eliminated largely by the use of numbered references. Long museum and institutional
names have been replaced by the abbreviations listed in the Introduction for each Part.
It is regrettable that a number of deep-sea groups—particularly the isospondylous
Alepocephalidae and Searsiidae and the iniomous Myctophidae and Neoscopelidae—
are represented in their respective positions by only interim accounts, inasmuch as the
detailed studies could not be completed for these volumes. The genus Cyc/othone, though
treated in somewhat more detail than the groups just noted, also awaits further study.
It is hoped that these missing sections can be incorporated in the following Parts.
While these volumes were in preparation, a comprehensive study of the Searsiidae by
A.E. Parr has appeared in Dana Report No. 51, 1960, to which the reader is referred
in lieu of a detailed account here.
It is of interest to note that the contents of these three volumes were originally
intended for inclusion in Part 3 only, a commentary indicating quite clearly the impos-
sibility of predicting the ultimate total number of volumes for this series. Because of
the redistribution of the subject matter into three volumes, the sections on Bony Fishes
and Class Isospondyli in Part 3, prepared originally when only one volume was assumed,
now obtains for all three volumes. Of particular importance in these sections are the
Keys—to Orders in Bony Fishes, and to Suborders and Families in Class Isospondyli.
Preface XV
The Keys and other material in these two sections have not been repeated in Parts 4
and 5.
Despite the ever-increasing progress of ichthyology, and of oceanography in gen-
eral, and despite the impressiveness of the contents in these volumes, the ichthyologist,
and anyone else for that matter who becomes acquainted with the contents here, cannot
avoid being impressed with the great gaps in our knowledge of fishes, particularly with
regard to the deep-sea groups in Parts 4 and 5. For all the study, observation, and ex-
ploration throughout the centuries, our present knowledge is indeed meager when com-
pared to what is unknown. Taxonomic descriptions aside, our knowledge of the deep-
sea fishes is limited primarily to speculation—to what may be surmised from their shape
and color, stomach contents, or location of capture. Of many shallow-water forms we
know little more. And even of those that have been subjects of extensive investigation,
such as the salmon and some of the commercially important herring-like groups, much
of what is known appears to be contradictory or debatable.
Many species are as yet unknown, and of those now known, many are still re-
presented by only one or a few specimens, these often damaged. Not until larger and
more extensive collections are assembled will it be possible to clarify the inter-relation-
ships of many species and groups, to obtain a more complete knowledge of distribu-
tion in area and depth, and much other important information. For the great majority
of deep-sea fishes, the records of depth distribution are sparse or crude at best, a
lack that stems largely from the fact that most of the deep-sea collections have been
taken with open nets rather than closing nets. With the former device, one knows only
that a collected specimen frequented some level of the water column between the sur-
face and the estimated fishing depth of the net. Even with the closing net, exact depth
is questionable because of uncertainties in determining the wire angle. Add to this the
fact that the scientific expeditions that have explored the depths have been few and
widely dispersed. Considerable areas of all oceans, more so in some than in others, still
remain unexplored.
Even as the present volumes constitute a tribute to the expeditions and countless
investigators who have contributed to our present knowledge of fishes, so do they re-
emphasize, by what is not known, the need for earnest and continued expansion of
marine exploration and investigation, particularly at a time when exploration of the
“‘sea’’ beyond our atmosphere holds the stage. Especially notable in Parts 4 and 5,
dealing mainly with the deep-sea isospondylids and iniomes, are the contributions of the
Orecon, Sitver Bay, and Comsat, whose surveys, sponsored by the U.S. Fish and
Wildlife Service, have provided a wealth of study material. Similarly, the contributions
of specimens obtained from stomach contents during investigations of the tuna and
other commercially important fish have enhanced many accounts; in fact, a number of
species, presumably fast-swimming forms that are consistently successful in evading
the slow-moving nets, are known only from such a source. Hand in hand with more
extensive collecting must go technical development of gear such as closing nets, devices
for accurate measurement of wire angle, underwater luminescent and photographic
XV1 Preface
equipment, and a long list of other essentials. For experimental work at sea, fully
equipped floating laboratories are needed to study live specimens at the source. And
perhaps most important of all is the need for an increasing number of well-trained
marine biologists and technicians.
The Sears Foundation extends its gratitude to Henry Bigelow and Giles Mead for
their various contributions in bringing these three volumes to fruition, particularly for
harvesting the manuscripts. To those authors who have given their generous cooper-
ation in achieving uniformity in the presentation of the material and in supplying ad-
ditional information, I extend special thanks. Many of the authors have acknowledged
the help and cooperation of individuals and institutions; to them, and particularly to
the many who have been omitted, I add the gratitude of the Sears Foundation. It is also
a special privilege to note the continued interest and confidence of Henry Sears, who
has generously aided in the financing of these volumes, and to acknowledge financial
support of various author investigations by the National Science Foundation. In the past
two or three years, the Editorial Board has been expanded by the addition of Daniel
Cohen, Giles Mead, and Daniel Merriman to its roster; to them and to the older mem-
bers of the Editorial Board, the Sears Foundation is grateful for many and varied
services.
Finally, it is with no small regret that I note the discontinuance of Albert Parr’s
active participation in the functions of the Sears Foundation and in the publication of
these Memoirs and the Yournal of Marine Research, both of which were conceived and
brought into being as a result of his sincere and deep interest in all things marine and of
his keen perception of the need for vehicles to distribute the thoughts and scientific
observations of an expanding oceanographic community. The results seen in the publi-
cations themselves speak for his contribution far more significantly than mere words
Ynove H. O1sen
May 1963 Editor
Introduction
ART 3 orf ruis Series deals with the more primitive groups of soft-rayed bony
fishes. The volume commences with a characterization of the class Osteichthyes (bony
fishes) and comparison of this group with the class Chondrichthyes (elasmobranchs),
followed by a characterization of the subclass Actinopterygii and a key to the various
orders dealt with here and in Parts 4 and 5: Acipenseroidei, Lepisostei, Isospondyli,
Giganturoidei, Iniomi, and Lyomeri, all of which were originally to be included in this
volume. The characterization and discussion of the Isospondyli concludes with a short
key to the suborders and an extensive key to the external characters of all the families
of the orders noted above; this, then, also applies to Parts 4 and 5 as well as to this
volume. Because of the large number of groups in the Isospondyli, only part of the
families, mostly shallow-water, are included in this volume; the rest, mainly deep-sea,
are relegated to Part 4. Thus, in addition to the sturgeons and gars, this volume includes
accounts of the tarpon, ladyfish, bonefish, the numerous anchovies, the herring, men-
haden, shad, sardine, pilchard, gizzard shad, round herring, salmon, trout, and smelt.
The geographic area covered extends from Hudson Bay southward to the Amazon
and seaward more or less to the mid-Atlantic, with Bermuda and the Caribbean areas
included. For the most part, only fishes living in brackish water or salt water have
been included.
Unfortunately, the accounts of the Alepocephalidae and Searsiidae could not be
completed for inclusion in this volume. These are therefore represented by brief interim
accounts that contain enough information for general identification of specimens in this
category.
In these volumes, museum and institutional names have been abbreviated ac-
cording to the following list:
AM Amsterdam Museum, Holland
AMNH — American Museum of Natural History
ANSP — Academy of Natural Sciences of Philadelphia
BLBG — Biological Laboratory, U.S. Fish and Wildlife Service, Bureau
of Commercial Fisheries, Brunswick, Georgia
BMNH — British Museum (Natural History), London
BNM —_ Bergens Museum, Norway
XVli
XVill
BOC
BU
BrU
CAS
CF
CFG
CM
CNHM
(SLU)
FSM
IJ
IRSNB
LMNH
MCZ
MMF
MHNBA
MNHN
MOM
MRAC
MSNF
NHR
NYZS
POFI
ROMZ
SIO
SU
a
UCLA
UF
UI
UL
UMIM
UMML
UMMZ
USNM
UT,
UW
VOM
WHOI!I
ZMA
ZMC
Introduction
Bingham Oceanographic Collection, Yale University
Boston University
Brown University
California Academy of Sciences
Carlsberg Foundation, Copenhagen
California Division of Fish and Game
Carnegie Museum
Chicago Natural History Museum
Cornell University
Florida State Museum
Institute of Jamaica, B. W. I.
Institut Royal des Sciences Naturelles de Belge, Belgium
Leiden Museum of Natural History, Holland
Museum of Comparative Zoology, Harvard University
Museu Municipal do Funchal, Madeira Islands
Museo de Historia Natural de Buenos Aires, Argentina
Muséum National d’Histoire Naturelle, Paris
Musée Océanographique, Monaco
Musée Royal d’Afrique Central, Tervuren
Museo di Storia Naturale, Firenze
Naturhistoriska Riksmuseum, Stockholm
New York Zoological Society
Pacific Oceanic Fishery Investigation, U.S. Fish and Wildlife
Service (Bureau of Commercial Fisheries), Honolulu, Hawaii
Royal Ontario Museum of Zoology, Canada
Scripps Institution of Oceanography
Natural History Museum, Stanford University
Tulane University
University of California at Los Angeles
University of Florida
University of Indiana
University of Louisville
University of Miami Ichthyological Museum
University of Miami Marine Laboratory
University of Michigan Museum of Zoology
United States National Museum
University of Texas
University of Washington
Vanderbilt Oceanographic Museum
Woods Hole Oceanographic Institution
Zoological Museum, Amsterdam
Zoological Museum, Copenhagen
Introduction X1X
Of the other abbreviations used in this volume, the following require explanation
here:
SL —_ standard length
FL —- fork length
hl — head length
m.W. or m.w.o. — meters of wire or meters of wire out, used in lieu of depth of
capture when discussing the catches of nets fished at indeter-
minate depths and not equipped with closing devices.
Finally, acknowledgment with gratitude is extended to all of the authors who
contributed to this volume and to the institutions that supported their work; to the
Sears Foundation for Marine Research, Yale University, and to Mr. Henry Sears for
his assistance in financing publication; to the National Science Foundation for funds
to finance part of the cost of research (through grant NSF G 7123); and to the numer-
ous ichthyologists and many others who have contributed information for this volume.
Henry B. BicELow
March 1963 Editor-in-Chief
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PACIFIC
South America
Bony Fishes
Superclass, Class, Subclasses,
and Orders
HENRY B. BIGELOW
Museum of Comparative Zoology
Harvard University
ACKNOWLEDGMENTS
With grateful acknowledgment for many helpful criticisms and suggestions from
William W. Anderson, U.S. Fish and Wildlife Service ; Frederick H. Berry, U.S.
Fish and Wildlife Service; Rolf L. Bolin, Hopkins Marine Station; Daniel M.
Cohen, U.S. Fish and Wildlife Service; Myvanwy M. Dick, Museum of Compara-
tive Zoology, Harvard; Robert H. Gibbs, Boston University; “foseph Gregory,
Peabody Museum of Natural History, Yale; N.B. Marshall, British Museum
(Natural History); Giles W. Mead, Museum of Comparative Zoology, Harvard ;
‘fames E. Morrow, University of Alaska; George S. Myers, Natural History Mu-
seum, Stanford University; Yngve H. Olsen, Bingham Oceanographic Laboratory,
Yale; Alfred S. Romer, Museum of Comparative Zoology, Harvard ; Bobb Schaeffer,
American Museum of Natural History ; William C. Schroeder, Woods Hole Oceano-
graphic Institution; Leonard P. Schultz, U.S. National Museum; Royal D. Sutt-
kus, Tulane University; ‘fohn Tee-Van, New York Zoological Society; Ethelwynn
Trewavas, British Museum (Natural History); Vladimir Walters, University of
California; and Norman f. Wilimovsky, University of British Columbia.
I
2 Memoir Sears Foundation for Marine Research
Superclass PISCES
Both the cartilaginous fishes (8) and the bony fishes have been grouped together
under the name Pisces by Goodrich (45) as a Grade intermediate between Branch and
Subgrade, by Berg (4) as a Series intermediate between Superclass and Class, and by
Bertin and Arambourg (32) as a Superclass. This last arrangement, with Pisces ranked
as a Superclass, is accepted here to embrace both cartilaginous fishes (Chondrichthyes)
and bony fishes (Osteichthyes). The cyclostomes, not regarded here as fishes in the
usual sense, are considered a separate Branch (Agnatha) of the vertebrate Subphylum
Craniata.
Bertin and Arambourg (3: 1956) have expanded the Superclass Pisces to include
the fossil Placodermi (including the Acanthodii and the Athrodiri); this ancient group,
however, was only remotely related to either the living cartilaginous fishes or the
living bony fishes. See Romer (73: 38-59) for an account of the placoderms and
their evolutionary pattern.
Class OSTEICHTHYES
Characters of Living Members. The internal skeleton contains true bone in greater
or lesser amount, especially in the regions of the skull, the jaws, and the pectoral arch. In
most the jaws are well developed, but in a few they are greatly reduced. The palato-
pterygoid part of the upper jaw-complex is fused solidly with the lower surface of the
skull in the Dipnoi (lungfishes) but not in the others, with one exception (15: 273).
The teeth are usually embedded in the bone (but see p. 6 for exceptions). In a few there
is a spiracular opening between the jaws and the hyoid arch (polypteroids and some
acipenseroids). The external opening of each nostril is typically double but is single in
a few (p. 6); the nasal sac ends blindly in most but opens internally into the mouth in
some. There is only one external gill opening on each side, and the two openings are
often united across the throat; the common chamber into which the internal gill clefts
open is roofed over by a dermal opercular flap supported in most cases by a series of
opercular bones. A typical bony fish has four or five pairs of gill arches (the fifth the
smallest) and four to six pairs of internal gill clefts.
The majority has two sets of paired fins—pelvic and pectoral. The pectoral girdle
is well developed and is attached at its upper end to the temporal region of the skull
in the great majority (see p. 6 for exceptions). The endoskeletal support of the pectoral
and pelvic fins consists ot either one or two series of short basal elements;? in the typical
pectoral fin these elements are in parallel or fan-like arrangement (Actinopterygii in
general, p. 11), with the outer part of the fin supported by fin rays; but in some there
is a jointed midrib with either a series of short side branches on each side, as in the
Dipnoi, or a terminal fan of jointed rays (Fig. 2), as in the living coelacanth Latimeria
1. According to Tchernavin (84: 284), there are six gill clefts in Eurypharynx (gulper eel).
2. The basalia of the pectorals are fused in Dallia to form a single plate; in Lophius there are only two; and in
general they are much reduced in the pelvics.
Fishes of the Western North Atlantic 3
(31: 2567, 2570, fig. 1840). The marginal parts of both the single and paired fins
are supported, at least in part, by jointed (segmented) and usually branched rays which
are either bony or fibrous in composition; in some the rays are apparently single,*
but in the great majority each ray consists of a right-hand and left-hand half, the halves
closely pressed together (Fig. 1). In a great majority the dorsal, caudal, and anal fins
are separated, one from the next, but in some these fins are continuous around the tip
of the tail.
In a great majority the scales are wholly of mesodermic origin and are embedded
in the skin below the epidermis. The outer surface of the scales consists of a simple
calcified layer in most but of an enamel-like substance (ganoine) in a few. In most the
scales persist throughout life and grow as the fish grows, generally without increase
in number or modification except as they may be replaced when accidentally lost. How-
ever, in some of the goby-like fishes, additional scales are normally interpolated with
growth! in the region of the caudal peduncle.
The anal, genital, and urinary tracts open separately to the exterior in most but
into a common cloacal pouch in a few. Some of the members of most of the major
groups have a swim bladder developed as a diverticulum from the dorsal side of either
the oesophagus or the stomach; a few (Dipnoi, polypteroids) have a functional lung (or
lungs) developed as an outpocketing from the ventral side of the pharynx.®
The notochord is persistent and unconstricted from end to end in a few, but in
most it is sharply constricted in its passage through the vertebral centra, often to thread-
like proportions, if it is not obliterated altogether.
Nomenclature.® The name Osteichthyes, proposed by Howes (34: 78, 87), has been
adopted by Goodrich (45: 210) and by numerous subsequent students, including Berg
(3: xv), Goodrich,? Romer (73: 75), Schultz and Stern (77: 227), Matsubara (47:
78, 154), and Bertin and Arambourg (31: 1978, 2068). It is equivalent to the Class
Pisces of Linnaeus (46), except that Linnaeus excluded the genera Acipenser (sturgeons)
and Lophius (anglerfishes); it is also equivalent to the Subclasses Lophobranchii plus
Potomabranchii of Bonaparte (zo: add. 13), with the addition of the lungfishes, which
were unknown to Bonaparte;§ it also equals: the combined Subclasses Dipnoi, Teleostei,
and Ganoidei of Miiller (53: 201-204); the Class Pisces as restricted by Jordan
(39: 108) and by Regan (71 [1929]: 305); the Teleostomi of Stensid;® the Classes
. The situation in this respect is not known for either the coelacanths or the Dipnoi.
4. Steindachner and Déderlein (8r: 269) have reported this for Cepola; Koumans (43: 267-279) has described in detail
and pictured clearly a similar situation for Oxyeleotris. Furthermore, J. R. Dymond (personal communication)
states that he has found in salmon and trout “....what appear to be rows or partial rows [of scales] that appear
to have been developed later than the rows above and below”; and Hubbs (36: 82) has reported an increase,
caused by parasites, in the number of scales in the genus Platygobius.
5- Millot and Anthony (3r: 2584) have reported a vestigial lung (“‘poumon degenere”) of large size in the living
coelacanth Latimeria. For a general survey of the relation between swim bladder and lung of living fishes, see Romer
(74: 344-345, 347-350):
- See Myers (54: 31-40) for nomenclature of terms that have been used for the higher categories of fishes.
. Ref. (30: xvm); considered a Subgrade of Class Pisces.
. Ref. (ro: 16); name Dipnoa used for Amphibia.
. Ref. (82: 111); taxonomic rank not stated.
we
oon n
4 Memoir Sears Foundation for Marine Research
Dipnoi and Teleostomii, combined, of Berg (4); the Teleostea of Fowler (27: 53);
and the Class Osteichthyes of Bertin and Arambourg (31: 1978).
The Class Osteichthyes as defined above includes all of the living groups of fishes
apart from the elasmobranchs (sharks and batoids) and chimaeroids. The cyclostomes
are not regarded here as “‘fishes”’ in the usual sense of the word.
Bony Fishes (Osteichthyes) Compared with Cartilaginous Fishes (Chondrichthyes). All
living bony fishes differ fundamentally from all living cartilaginous fishes in that the
internal skeleton of the former consists of true bone in greater or lesser amount, or of
some derivation of true bone, whereas the skeleton of living elasmobranchs and chimae-
roids contains no true bone. Bony fishes differ to an even greater extent from the cy-
clostomes in that the former have well-developed jaws and (typically) two olfactory
organs. The bone of the Osteichthyes represents in part the ossification of the primary
skeletal cartilages by the action of bone-forming cells from without (for details, see
Goodrich, 45: 65-67); but in part, especially in the regions of the skull and pectoral
arch, it consists of so-called membrane bone, which is not preformed in cartilage. In
neither case is it derived from calcifications of the kind that stiffen the cartilaginous
skeletons of elasmobranchs and chimaeroids.
Due to the presence of bone, the skull of all living bony fishes is marked by su-
tures,!° which is not so in the cranium or brain case of living cartilaginous fishes. How-
ever, this seems an appropriate place to note that in fossil placoderms, which are only
remotely related to the bony fishes, the internal skeleton was more or less bony;
many of them were more or less completely armored with bony plates, and their skulls
showed sutures corresponding to those in Osteichthyes. Also, many of the fossil Ag-
natha, of which the cyclostomes of today are commonly regarded as degenerate descen-
dants, were variously encased in bony armor, and at least in some cases their skulls
were marked with sutures.¥
Hardly less diagnostic for most of the living bony fishes, including the living
coelacanths,}* is the presence (typically) in some part of their fins of segmentally jointed
rays!® (mostly branched) which are either bony or fibrous. These are the soft rays known
to every student of fishes. Even among the living Dipnoi (lungfishes), some of the
fin rays (hair-fine in this group and of doubtful homology) are segmented, though most
of them continue unsegmented throughout life, thus resembling superficially the horny
rays (ceratotrichia) of sharks (for details, see Goodrich, 28: 480-482). In the fossil
Dipnoi, however, all of the fin rays were segmented.
No living elasmobranch or chimaeroid has in its fins any structures that correspond
structurally to the segmented fin rays of bony fishes. Thus the horny fin rays of sharks
are not jointed, and they grow inward from the outer margin of the fin, not outward
ro. The bony neurocranium of the extinct palaeoniscoids, a primitive group of bony fishes, is not marked with sutures
(Stensié, 83; Rayner, 65: 287).
11. For a readable account of these ancient groups, see Romer (73: 25, 38).
12. For the fin rays of Latimeria, see Millot and Anthony (3r: 2567, 2570, fig. 1840).
13. The fin rays are not jointed in the lyomerids, the trachypteroids, the stylophoroids, and perhaps in other
groups as well.
Fishes of the Western North Atlantic
5
from its base as the fin rays grow in actinopterygian fishes,'* and presumably in coela-
canths. Although the radial cartilages that support the wing-like pectoral and pelvic
fins of skates and rays recall in appearance the jointed fin rays of bony fishes, they are
a part of the internal skeleton; and they are single whereas the soft
rays of bony fishes are double (p. 3; Fig. 1).
Although the fin rays of the Dipnoi, which are intermediate
between elasmobranchs and actinopterygian bony fishes in this
respect, are jointed, they resemble the horny fin rays of elasmo-
branchs and chimaeroids in their growth inward from the outer edge
of the fin instead of outward from the base, in their arrangement in
two series, and in their seemingly single nature.
The contrast between bony and cartilaginous fishes is further
blunted by the presence of hair-thin unjointed horny rays like those
of the elasmobranchs in the so-called adipose dorsal fin of certain
bony fishes (salmons and catfishes)'* and in the free edges of the
rayed fins of other bony fishes.”
All of the other differences that have been proposed as alter-
native between bony and cartilaginous fishes are beset with excep-
tions. For example, it has been cited repeatedly that the dermal
fold, which roofs the common branchial cavity (into which the in-
ternal gill clefts open), is supported in bony fishes by a series of
opercular bones, whereas in the only living cartilaginous fishes that
have a corresponding dermal flap (chimaeroids) it is supported by a
series of cartilaginous rays only. The polyodontids (paddlefishes),
however, have only vestiges of the opercular bones; in the gigantu-
roids, in some stomiatoids, and in some eels they are ossified but
little; and in the lyomerids (an eel-like group of deep-sea fishes with
Ficure 1. Soft dor-
sal fin ray of Esox
Jucius (pike) with
its skeletal support;
Lert-lateral view;
Ricut-front view.
R ray; P basal seg-
ment, and m median
segment of support-
ing radial. After
Goodrich, emended
somewhat.
enormously distensible mouths) and in the lophobranchs they are lacking altogether.'8
Below the gill covers, the great majority of bony fishes has a series of parallel
branchiostegal rays, but these rays are lacking in the elasmobranchs, chimaeroids,
Dipnoi, acipenseroids, and deep-sea Lyomeri.
The scales of bony fishes are deeply implanted in the skin, are wholly of mesodermal
origin, and they usually persist throughout the life of the fish (for exceptions, see ftn. 4);
14. For the development of soft rays (leptotrichia) in the actinopterygians, see especially Pennant (63: 190-195) and
Bertin (31: 735)-
15. For the structure, growth, and homologies ‘of the fin rays of the different major groups of living fishes, see espe-
cially Goodrich (28: 480-491), Eaton (25: 195-200), and Bertin (31: 731-736).
16. According to Dr. George S. Myers (personal communication), what appear to be ossified spines are developed
in the adipose fin of some African catfishes, and with age in some characids of the genus Serrasalmus; also the lori-
cariid and calichthyid catfishes have the adipose fin supported by an initial bony spine.
17. For a review of early literature on this subject with accounts and illustrations of these rays in the adipose fin of
salmon and trout, see Valette St. George (88: 187-192, pl. 15); for a more recent review, see Goodrich (28:
473-477):
18. For a discussion of the evolution of the operculum in different groups of fishes, fossil and living, see Eaton
(24: 42-46).
6 Memoir Sears Foundation for Marine Research
in elasmobranchs and chimaeroids the so-called placoid scales are tooth-like in structure,
have an extensive pulp cavity, and are tipped with an enamel-like substance of ectodermic
origin. Furthermore, the scales of a shark are not only short-lived but increase in number
as it grows. Although the presence of bony-fish scales is positively diagnostic for those
fishes that have them, their absence is not negatively diagnostic, for it is common
knowledge that scales are wholly lacking in some bony fishes. Furthermore, certain
groups of the bony fishes have placoid denticles of the elasmobranch type on their
bony scales, e. g. the polypteroids, the lepisosteids (gars), and some ostariophysids (cat-
fishes); and some of this last group have placoid denticles on their fin spines as well.!
Denticles, believed to be placoid in nature (an interpretation calling for verification),
have also been described for the snouts of young Xiphias (swordfish) and young Jstio-
phorus (sailfish) (13: 321, pls. 1, 2); and Dr. Ethelwynn Trewavas informs us that they
are present on the sucking disc of remoras as well (personal communication).
It has been stated repeatedly that, whereas the teeth of elasmobranchs are simply
implanted in the gums, those of most bony fishes are set in the bone. The polyodontids
(paddlefishes), however, and certain Salariinae (blennies) parallel the elasmobranchs in
this respect, their teeth having no special connection with the jaw bones; and the teeth
in Plecoglossus (related to the salmonoids) are attached to the jaws by connective tissue
only (14: 437).
The nasal openings are single on either side among the elasmobranchs in general
and among the chimaeroids, but among bony fishes they are double (typically) on each
side. However, some bony fishes, e. g. pomocentrids and cichlids, have only one nasal
opening on each side. On the other hand, it has been discovered recently that each
nasal opening in one genus of torpedo rays (Dip/obatis) is divided into two (58: 482,
HO 27252, US OR aioe 7):
Various texts have emphasized the mode of suspension of the pectoral girdle as a
diagnostic feature: it is attached to the skull in typical bony fishes; it is not connected
at all to the axial skeleton in living sharks and chimaeroids; and it is attached to the
anterior part of the vertebral column in batoids. However, among bony fishes, the
pectoral girdle has lost its attachment to the skull in the deep-sea giganturoids and sac-
copharyngoids, in the amphipnoids (a group that is eel-like in appearance but differs
widely from the true eels in anatomy), and in the Apodes (eels) as a whole; and in some
Apodes (Muraenidae) the pectoral girdle is lacking altogether. On the other hand, the
girdle is attached by a ligament to the third or fourth vertebra in the Opisthomi.?°
The presence of either a swim bladder or a functional lung is diagnostic for such
of the bony fishes as have either, unless a small diverticulum from the dorsal side of
the oesophagus, observed in embryos of certain sharks by Miclucho-Maclay (49: 448,
pl. 5), represents the rudiment of a swim bladder. But the absence or presence of a
19. For illustrations conveniently available of placoid denticles among bony fishes, see especially Goodrich, in Lan-
kester (45: 231, fig. 201; 289, fig. 262; 291-292, figs. 264, 265). For a recent account, with references and excellent
illustrations of these denticles among living catfishes, see Orberg (56: 487, figs. 4C, 4D).
20. An Order of eel-shaped freshwater fishes of doubtful affinity, known from tropical Africa, the Euphrates, and
southern Asia north to Peking (68: 219).
Fishes of the Western North Atlantic 7
swim bladder is not a feature of much direct service to the systematist, for groups
that differ widely in other respects may agree in lacking the swim bladder, and
vice versa.
In some families, even where most of the genera have a swim bladder, others do
not. Familiar examples of such families are the Blenniidae (blennies), the Polynemidae
(threadfins), and the Scombridae (mackerels); in the last group the chub mackerel
(Pueumatophorus) has a swim bladder while the common mackerel of the North Atlantic
(Scomber) has none (Starks, 92: 223).
It has long been emphasized that, while the gill folds of elasmobranchs and chi-
maeroids are attached to the interbranchial septa outward nearly to their tips, those of
the higher groups of bony fishes are free from the septa for most of their lengths. These
extremes, however, are bridged on the one hand by the Dipnoi, where the gills reach
only slightly beyond the septa, and on the other hand by the polypteroids, acipenserids
(sturgeons), and lepisosteids (gars), where they extend only a little farther outward.
Furthermore, most of the bony fishes lack functional respiratory folds on the anterior
(hyoidean) wall of the first gill pouch, such as are characteristic of elasmobranchs and
chimaeroids; but the acipenserids and lepisosteids are exceptions to this rule, as is the
living coelacanth Latimeria (Millot and Anthony, 31: 2584).
The ventral or pleural ribs between the adjoining surfaces of the peritoneum and
body wall are diagnostic for such of those bony fishes as have them, for the ribs of
the living cartilaginous fishes*? that have any lie in the horizontal septum that separates
the musculature of the trunk into an upper and lower division.2? Many bony fishes,
however, lack ribs of any sort: the lophobranchs (26: 28-30), for example, the gi-
ganturoids (70: 57), the stylophorids (80: 21), and some trachypteroids (genus
Agrostichthyes) (information from Vladimir Walters). The tetraodontids (swellfishes),
the diodontids (porcupinefishes), and the ostraciodontids (trunkfishes), among the
plectognaths, lack ribs (76: 8, 10; 41: 325, pls. 15, 16), while their relatives,
the Balistidae (triggerfishes) and Aleuteridae (filefishes), have ventral ribs.*4 It is also
evident from a glance at the skeleton of Lophius (anglerfish) that it is ribless, as are
the Ogcocephalidae (batfishes), by personal observation, and Rhynchoceratias (60: 13),
among pediculate fishes.
So-called dorsal ribs are not peculiar to the elasmobranchs alone, for the po-
lypteroids, salmonids (salmons), some clupeids (herrings) (26: 209), Esocoidea (pikes)
(4: 242, 429), and balistids (triggerfishes) among the plectognaths (41: 304) have
both ventral ribs and ribs that occupy the dorsal position. However, it appears that
21. N.B. Marshall contributes the information (personal communication) that the major groups of bathypelagic
fishes, such as stomiatoids, deep-sea salmonoids, and Miripinnati, can be characterized by the structure of the
swim bladder.
22. The chimaeroids and some of the rays are ribless.
23. But these ribs are considered by Emelianoy (26: 244) to be homologous in their development with the pleural
ribs of bony fishes.
24. Regan (66: 285, 286, fig. 56) interpreted the rib-like bones of the Balistidae and Triacanthidae as intermuscular
in nature and “not bordering the abdominal cavity.” But it seems clear from Kashkaroff’s account (41: 303,
325, pls. 12-14) and from our own dissections that they are true ventral ribs and do embrace the body cavity.
8 Memor Sears Foundation for Marine Research
these dorsal ribs may not be homologous, embryologically, with those of the elasmo-
branchs (26: 215-217).
A spiral valve in the intestine, which is universal in elasmobranchs and chimae-
roids, is well developed in the living coelacanths (50: 426, 2582, Millot and Anthony),
in the Dipnoi, and in the polypteroids alone among bony fishes; but it is represented in
vestigial form in the acipenserids (sturgeons) and polyodontids (paddlefishes). A struc-
ture that more or less suggests the elasmobranch spiral valve has been reported also
as occurring in Chirocentrus, an Indo-Pacific genus falling among the clupeoids (z8:
160, pl. 565; 45: 116, fig. 77a, Goodrich), in Argentina silus (the common herring
smelt of the North Atlantic),*° in Salmo gairdneri (rainbow trout), Thymallus (grayling),
and the Coregonidae (whitefishes) among the salmonids (information from Daniel Co-
hen), in Macropinna among the Argentinoidea (15: 282), in the young of the characid
genus Curinata (information from G.S. Myers), and in Alepocephalus (18: 172, 176).
But the so-called spiral valve of these isospondylous fishes probably is not homologous
with the spiral valve of elasmobranchs and chimaeroids.
On the males of living cartilaginous fishes among which internal fertilization in
the female is universal, a copulatory organ commonly called a clasper is developed in
connection with each pelvic fin. But the male copulatory organ of the few bony fishes
whose eggs are fertilized internally is single (typically) and has no connection with the
pelvic fins, being developed either from the genital papilla on the urogenital orifice, or
in connection with the anal fin, or as a special structure on the chest. However, the
phallostethids (Indo-Australian freshwater fishes of the Order Percesoces) are an ex-
ception, for their “‘much modified pelvic fins and girdle” join in the formation of the
copulatory organ (2: 478).
Finally, such of the bony fishes as pass their lives in salt water maintain their body
fluids in osmotic balance with the surroundings in one way, elasmobranchs in another.
In each case it is a problem of maintaining the concentration of salts in the blood at a
level lower than that of the surrounding sea water; the osmotic pressure in the body
fluids of marine teleosts is only about 40°/, of that of sea water.?”? Elasmobranchs
solve this problem by retaining in their blood and tissues much of their nitrogenous
wastes in the form of urea; thus they raise the total concentration of materials in solution
to a level that is nearly as high as, or even higher than, that of the surrounding sea water
without increasing the salt concentration (74: 393). As a result of this mechanism, the
marine sharks and rays have no special need of conserving water. According to Black,
“They rarely drink salt water and obtain free water in their food” (12: 187). Marine
bony fishes, never having chanced on this device, prevent an accumulation of salt-ions
25. Kendall and Crawford (42: 10, fig. 1A) pictured Argentina as having a complete spiral valve, but Cohen writes
that while he has noted spiral thickenings in the intestine of various Argentinidae, including six specimens of Ar-
gentina silus (6: 121), in none of these last did he “observe a functional spiral valve as figured by Kendall and
Crawford.”
26. According to Jacobshagen (9: 611), the structure that has been called by this name among the Isospondyli actu-
ally represents a series of simple circular ridge-like thickenings of the inner surface of the intestine. For further
discussion, see Cohen (z6: 96, 98).
27. Estimated by Denton and Marshall (2r: 754) from data assembled by Krogh (44: 130 FF).
Fishes of the Western North Atlantic 9
in their body fluids by a complicated excretory process. Thus, via their gill membranes,
they excrete most of the salt from the sea water that is consumed, and at the same time
they reduce their excretion of water to the minimum that will provide the necessary
amount of urine. In some groups this reduction is effected by atrophy or loss of the
glomeruli,** but in other saltwater groups that still retain the glomeruli, including the
living coelacanths (37: 2588, Millot and Anthony), this goal appears to be reached by
constriction of the glomerular arterioles.”
We have yet to learn how bony fishes that spend part of their life in fresh water
and part in salt water or that suffer no apparent harm from rapid changes in the
salinity of the water manage to maintain themselves in osmotic balance with their sur-
roundings. This applies equally to such of the cyclostomes as spawn and pass through
their early stages in fresh water but make their growth in salt water. Those that pass
their entire life in salt water are isotonic to their surroundings, or very nearly so (see
especially McFarland and Munz, 48: 348).
Subclasses. Students of phylogeny have disagreed widely concerning the relation-
ships of the various groups that make up the ichthyological tree. In the first instance,
a decision on the best arrangement of the various groups of the Class Osteichthyes
into Subclasses depends on the relationship of the Dipnoi (lungfishes) to the other
bony fishes. Johannes Miiller (53), writing in 1844, rated the Dipnoi as a separate
Subclass of the Class Pisces, and this scheme was later adopted by Smith Woodward
(91: 234), by Goodrich (45: 230), by Bridge, and by Jordan (39). Berg’s (4) later
characterization of them as a class of the Series Pisces embodies this same point of view,
as does Bertin and Arambourg’s treatment of them as the Subclass Dipneusti of the
Class Osteichthyes (31: 2522). It is likely, indeed, that this scheme would still be
adopted generally were our knowledge of the Dipnoi limited to those of today, for these
are set apart from all other living groups of bony fishes by the solid fusion of the palato-
pterygoid-quadrate elements of the upper jaw-complex with the lower surface of the
skull, by the general morphology of their brains and hearts, and by their total lack of
both premaxillary and maxillary bones (a significant difference); in addition, the living
Dipnoi are set apart further from all other bony fishes except the coelacanths by the
presence of a jointed axial skeleton in their paired fins (Fig. 2G). None of the other
various features that have been cited as alternative between Dipnoi and other bony fishes
is strictly so. Palaeontological evidence, furthermore, is to the effect that the oldest
known fossil Dipnoi (Devonian) resemble in many characters the Rhipidistia (Middle
Devonian), whose nearest relatives among present-day fishes are the coelacanths
(73: 114).
Accordingly, the Dipnoi and the coelacanths among living fishes have been
united by several recent writers (55: 376; 72 [1937]: 3273 73: 5893 77: 2273 47:
28. Examples cited by Smith (79: 104) are Lophius (anglerfishes), Opsanus (toadfishes), Hippocampus (seahorses), and
Syngnathus (pipefishes).
29. For more extended discussions of this general subject, from which the foregoing summary has been drawn, see
Prosser et al. (64: 48-52), Smith (79: 62-64, 102-105), Romer (74: 392-393), Denison (20: 429-430, 439-441),
and Black (12: 182-199, refs. 199-206).
10 Memoir Sears Foundation for Marine Research
154) to form one of the Subclasses of the Class Osteichthyes, and this same course is
followed here.
In 1861 Huxley proposed for this Subclass the name Crossopterygidae (subse-
quently changed to Crossopterygii) as a Subordo of his Ordo Ganoidei (38: 23, 25). Other
names proposed later are: Amphiboidei by Hubbs in 1919 (35: 589); Choanichthyes
by Romer in 1937 (72: 56), with Crossopterygii retained as an Order to include fossil
as well as living Choanichthyes (73: 589); and Sarcopterygii by Romer in 1955 to
replace his earlier name Choanichthyes (75: 126).
Ficure 2. Skeleton and basal part of pectoral fin rays of: top, Latimeria, after Millot and Anthony, somewhat
emended; LowER LEFT, Neoceratodus (lungfish), after Giinther, somewhat emended; LowER RIGHT, Sa/mo trutta
(European trout), after Parker and Haswell, somewhat emended. c pectoral girdle; B basal segment, and p distal
segment of radialia; r fin rays.
None of these names seems wholly appropriate. While Crossopterygii, in a restric-
ted sense, has been widely employed in scientific literature,?° its continued use seems
likely to lead to confusion because it has long been associated with the polypteroids,
which were the basis for Huxley’s name but which have been excluded from the Cross-
opterygii, Goodrich (29: 91) having shown that the affinities of the polypteroids lie
with the actinopterygian fishes.
Amphiboidei does not seem a happy choice, for while the fishes concerned may
be “related to amphibian ancestry,” the living coelacanths are about ‘“‘as unamphibian
as a fish can be,” as Romer has aptly expressed it (75: 125, 126).
Choanichthyes is a misnomer if the coelacanths are included, for internal nares
neither were nor are present among fossil (75: 126) or living coelacanths, regardless
of the situation in their hypothetical ancestors; and it is doubtful whether the internal
30. For a recent defense of this course, see Trewavas, et al. (87: 126-127).
Fishes of the Western North Atlantic rea
nares of the Dipnoi are homologous with those of the tetrapod vertebrates. At any rate,
they are not used in respiration.
Although objection may be raised to the use of Sarcopterygii when older names
are available, this name does not carry with it any evolutionary implication that may
not be warranted; and furthermore, it has never been employed in more than one sense,
hence it is accepted here. This Subclass need not concern us further, for the Dipnoi are
strictly confined to fresh water while living representatives of the coelacanths have been
taken only off the southeastern coast of Africa and near Madagascar.
Actinopterygii has been chosen here as a Subclass name to represent the bony
fishes that remain after subtracting the Sarcopterygii, a course which accords with the
weight of contemporary opinion. An alternative scheme, proposed by Regan (69: 458;
7I [1937]: 312) and adopted by Norman (55: 376-377), would distribute them in two
Subclasses: Palaeopterygii for acipenserids, polyodontids, and polypteroids; and Neop-
terygii for the remaining groups. But this seems an unnatural arrangement; although
the polypteroids and acipenserids are similar in that the number of fin rays is greater
than the number of their basal skeletal supports and because there is an unmistakable
spiral valve in the intestine, the polypteroids differ widely from the acipenserids in the
nature of both caudal fin! and dorsal fin, in the segmentally constricted notochord,
and in the presence of a pair of bony gular plates between the branches of the lower
jaw. Indeed, the polypteroids differ so widely from all other living bony fishes that
Bertin and Arambourg’s separate Subclass for them, Brachyopterygii (31: 1981, 2500),
may represent a step forward in attempts to present more accurately the affinities of
the various groups of bony fishes.
Subclass ACTINOPTERYGII
Ray-finned Fishes
Characters of Living Members. The skull in its early stages is a primitive cartilaginous
cranium that is replaced during development by bone in varying degrees in different
groups; in most of the living bony fishes it is completely ossified in adults. Its outer
surface is invested with a covering sheath in an intricate pattern of so-called dermal or
membrane bones that develop as new structures ‘‘in the membranes of certain regions
of the skull” and that appear to represent such modified scales; thus the entire skull
is “‘so welded together to form a compact whole that in the adult fish it is often im-
possible to decide as to which category a particular element belongs” (55: 161).
Generally the upper jaw includes paired premaxillary and maxillary bones;%* the
premaxillaries, supplemented in many cases by the maxillaries, form the upper region
31. Symmetrical in the polypteroids, with the posterior part of the vertebral column continuing the axis of the body;
strongly asymmetrical in the acipenserids and polyodontids, with the posterior part of the vertebral column bent
upward (heterocercal).
32. Including the polypteroids.
33- Premaxillaries are lacking among acipenserids (sturgeons) and polyodontids (paddlefishes) (32: fig. 20); in Bathy-
saurus of the Iniomi and in Gigantura maxillaries are lacking; both premaxillaries and maxillaries are lacking
12 Memoir Sears Foundation jor Marine Research
of the mouth; the premaxillaries, and often the maxillaries, are attached to the skull
with varying degrees of firmness in different groups. The palatopterygoid-quadrate
elements of the upper jaw-complex are articulated anteriorly with the ethmoid (nasal)
region of the skull and are suspended posteriorly (usually via a symplectic bone) from
the hyomandibular arch; but they are not fused solidly anywhere with the lower surface
of the skull, except in one known case.*4 A bony gular plate is present in the chin region
between the two branches of the lower jaw in the Amiidae and Elopidae, and there is
a pair of such plates in the polypteroids. The cheek region of the skull is without
squamosal bone.
In most of the bony fishes, the jaw teeth, if present, are separate from each other,
and their substance is separate from that of the jaw except for those in which it is
cemented basally “by bony substance which is resorbed when the tooth is shed” (45:
272 — Goodrich).
A persistent spiracular opening is present in only a few (polypteroids, some aci-
penserids, and polyodontids). The nasal cavities end blindly in the great majority,
however, in Astroscopus (22: 9933 23: 348-365; I: 371) and in Uranoscopus (original
observation) of the Uranoscopidae they open both inwardly to the mouth and out-
wardly to the exterior, as they do in various members of the eel tribe as well (z: 371);
this is suggestive of similar conditions among the Dipnoi (lungfishes), but the two
are probably not homologous. Gill rakers, present in the great majority,®° vary widely
in number. Branchiostegal rays, though present in the great majority, are lacking in a
few (sturgeon and paddlefish tribes, polypteroids, and Lyomeri).
The endoskeletal support of the pectoral fin consists typically of one or two series
of short basal radialia, either in parallel or in more or less fan-like arrangement; these
radialia articulate with the pectoral girdle directly in most Actinopterygii but indirectly
in a few. The endoskeletal support of the pelvic fin is essentially similar, but here the
radialia are greatly reduced. The dorsal and anal fins are supported basally by a single
series of more or less rod-like radialia; in most, the rays of both dorsal and anal fins
are equal in number to the basal radialia, but in both acipenserids (sturgeons) and
polyodontids (paddlefishes) the rays of both fins are much more numerous than the
radialia; among the polypteroids, while the dorsal rays more or less equal the number
of radialia, the anal rays are much more numerous.
The outer surface of the scales (if any) consists of a simple calcified layer in most,
but is enamel-like in appearance (ganoine) in a few.
or are very small in some members of the Argentinoidea (86: 605-614, pl. 11; 62: 30-31, figs. 9-11); and the
Monognathidae (an aberrant family of lyomerids) have no upper jaw-complex at all (5: 533-540, figs. 1-4; 6: 1,
5, figs. 2, 3).
34. In the argentinoid genus Macropinna, according to Chapman (r5: 273), the palatoquadrate complex is “immoy-
ably synchronized” with the occipital portion of the skull. Among the Ophiocephalidae and Symbranchii, the
metapterygoid bone (part of upper jaw-complex but not forming part of gape) is in contact with the sphenotic
bone, which is part of the posterior rim of the orbital capsule of the skull. For references, see Berg (4: 306, 471).
For an explanation of the mode of suspension of the upper jaw that is typical of ray-finned fishes, see especially
Parker and Haswell (59: 243, fig. 207; 244).
35- Lacking in some members of the Cetomimidae (6r: 20-32), in the Istiophoridae, and perhaps in some others.
Fishes of the Western North Atlantic 13
Usually each of the genital, urinary, and intestinal tracts opens separately to the
exterior, but among the syngnathids (33: xxvu, ftn. 1) and lophotids (information
from Vladimir Walters) they empty into a common cloacal pouch. In many, a swim
bladder is developed as a single outpocketing from the dorsoanterior part of the di-
gestive tract, and an open pneumatic duct persists throughout life in some but not in
others; in the polypteroids a functional paired lung is developed as a ventral out-
pocketing from the pharynx.*
The brain, consisting of a large cerebellum, large optic lobes, and a feebly de-
veloped cerebral region, is roofed (typically) in its anterior part with simple nonnervous
epithelial tissue. The notochord is more or less constricted segmentally except in the
acipenserids, polyodontids, and deep-sea halosaurids. The jugal sensory canal (forward
extension of the lateral line) is represented in most by a horizontal line of sensory pits
that cross the cheek region (89: 378). The pericardium is thick-walled posteriorly; the
heart chambers show no indication of subdivision into arterial and venous channels.”
Nomenclature. The choice of a name for this Subclass lies between Actinopterygii
and Teleostomi.
In 1839 Bonaparte proposed the name Teleostomi* as a sectio to include his Ga-
noidei and certain other groups of bony fishes, but excluding the acipenserids (sturgeons)
and some others (r0: 15). Many years later Owen expanded Bonaparte’s Teleostomi
to include the bony fishes as a whole, excepting the Dipnoi, which were unknown to
Bonaparte (57: 7). Since 1868 Teleostomi has been used in this sense by numerous
authors.
The Subclass name Actinopteri,® changed subsequently to Actinopterygia and
Actinopterygii, was coined by Cope to include the acipenserids, lepisosteoids, amioids,
and all higher groups of bony fishes, except the polypteroids (17: 449). The Subclass
was employed in this same sense by Traquair (85: 505, 507), Goodrich (45), Jordan
(39: 111), Lehman, Bertin, and Bertin and Arambourg (31: 2070-2500). In 1928
it was expanded by Goodrich to include the polypteroids (29: 91), and it has been
employed with this modification by Stensié,4° Berg (4: 159, 392), Romer (73: 105,
$79), and Schultz and Stern (77: 228).
Superorders. Several students have distributed the Orders of Actinopterygii among
three Superorders, namely: Chondrostei (or Chondrosteoidea) for the acipenserids (stur-
geons) and polyodontids (paddlefishes); Holostei for the Lepisosteoidea (gars) and
Amioidea (bowfin); and Teleostei for the remaining bony fishes. It does not seem pos-
sible to draw a line of distinction between these Superorders if the fossil record be
taken into account, hence they are not recognized here.”
36. For discussion of the relationship of swim bladder to lungs, see especially Romer (74: 343-350, figs. 225, 227)-
37- Among the living Dipnoi, the pericardium is thin-walled, and there is an “incipient division of the heart into
a venous and an arterial channel” (45: 249 — Goodrich).
38. From the Greek words teleos (meaning perfected) and stoma (meaning mouth).
39. Based on the Greek words aétis, meaning a ray, and pferon, wing or fin. The spelling was later changed to
Actinopterygia by Cope and to Actinopterygii by subsequent writers.
40. In a series of papers (1932-1936).
41. For a discussion of this matter, see especially Berg (4: 160-163, or 392-395).
14 Memoir Sears Foundation for Marine Research
The name Teleosteit? has proved far too useful to be banished from nontechnical
literature, whatever its fate may be in technical writings; we urge that it be trans-
literated henceforth to fit the language in which an author may be writing; for example,
teleosts in English, téleostéens in French, Teleostier in German, etc.4? However, the
sooner “‘Ganoidea”’ and “‘ganoids” be relegated to oblivion the better, for the groups
of living or extinct fishes** that were united under this name by Miller (53: 203)
and that have been grouped similarly by subsequent authors do not represent a
natural assemblage.
Orders of the Subclass Actinopterygii. The various classifications that have been pro-
posed for the living ray-finned fishes since the appearance in 1844 of Miiller’s classic
“system” (53: 201-204) have been reviewed by Lénnberg (11: 2-62), by Berg (4),
and more recently by Matsubara (47: 1-54). Hence it seems sufficient to remark here
that the number of Orders that have been recognized within the Subclass Actinopter-
ygii during the past 50 years or so has ranged from nine by Goodrich (30: x—xvi1)
and 12 by Romer (73: 579-585) to 24 by Matsubara (47), 32 by Regan (67: 76-82),
36 by Schultz and Stern (77: 220-247), 43 by Jordan (39), 44 by Berg (4), and 35
by Bertin and Arambourg (32: 1978-1981). This clearly illustrates the diversity of
opinions and conclusions at which eminent students have arrived from consideration of
essentially the same facts.
Part 3 of this series of volumes deals with the Acipenseroidei (sturgeons) of the
western North Atlantic, with such of the Lepisostei (gars) as enter brackish or salt water,
and with part of the Isospondyli, namely the Elopoidea (Elopidae, Albulidae), Clupeoidea
(Engraulidae and Clupeidae, with only interim accounts of Alepocephalidae and Searsii-
dae), and Salmonoidea (Salmonidae, Coregonidae, and Osmeridae). Parts 4 and 5 deal
with the remainder of the Isospondyli, the Iniomi, the Giganturoidei, and the Lyomeri.
Order of Presentation. Since it is not possible to represent the true interrelation-
ships of different large groups of animals on the printed page, any sequence of pres-
entation that may be adopted (other than an alphabetical one) must necessarily be
artificial. This would be true even if the taxonomic units in question could be traced
back through the ages to their earliest known fossil ancestors and even if we had a
uniform yardstick by which we could measure the relative extent to which the dif-
ferent groups have diverged during their evolutionary histories. In any general account
of a group as large and as varied as the bony fishes, the most one can hope for is that
the sequence of treatment—Orders within Subclasses, and Families within Orders—
shall be consistent with animal affinities “insofar as is practical” (78: 13). The sequence
followed here represents a compromise between antiquity of ancestry, degree of special-
ization, and accepted precedent.
42. From the Greek #eleos (meaning perfected or completed) and osteon (meaning bone).
43. Ichthyostés, as used by Moreau (5r: 2; 52: 89), is an equivalent.
44. Among living fishes—the sturgeon family, the paddlefish family, the polypteroids, and the gars. For a list (per-
haps only partial) of the various extinct groups that have been joined together recently as “Ganoidei,” see
Jordan (39: 111-116).
45. For other places of publication, see Dean (rg: 174).
Fishes of the Western North Atlantic 15
Sere = An
os an eare)
ree
Bi: ToRLULSBrAN) YEA) ESAS REET ER ae
ETH ELEUOMTTFES EMA Asre rere e STE
SSS
Ficure 3. Caudal fin of: top, Acipenser oxyrhynchus (sturgeon), showing especially the extended scaly axis and
fulcral scales, drawn by E. N. Fischer; tower tert, Sa/velinus alpinus (Arctic charr), from eastern Greenland,
drawn by Jessie H. Sawyer. LowER RIGHT, Lepisosteus (longnose gar), after Kélliker, combined with drawing
of L. osseus by Jessie H. Sawyer, with fulcral scale from upper margin, about x 2.
The sturgeons head the list, with the gars in second place, for while the known
fossil record does not reach back beyond the upper Cretaceous for the living repre-
sentatives of either of these groups,** fishes that seemingly were ancestral to both of
them are known from as far back as the Upper Triassic. The Isospondyli and the
Iniomi come next in recorded antiquity; the Elopidae (tarpon and ladyfishes), Clupei-
dae (herrings), and Chanidae (milkfishes) are known from the Lower Cretaceous, with
the Albulidae (bonefishes) and even the Myctophidae (or some closely allied genera)
and Aulopidae known from the Upper Cretaceous. There is no known fossil record
46. Wilimovsky (90: 1205-1208, pl. 132) has described and figured an undoubted sturgeon from the Upper Creta-
ceous of Montana.
16 Memoir Sears Foundation Jor Marine Research
for either the Bathylaconoidea or the Lyomeri, curiously modified groups that are in-
cluded in Parts 4 and 5 because they appear to stem back either to isospondylic or to
eel-like ancestry; for further details, see Romer (73: 579-584).
Identification. The sturgeons, the gars, the lyomerids, and the giganturoids are set
apart by external features so obvious that no one at all acquainted with fishes would
be likely to mistake any one of them for any other kind of fish that is known from the
western North Atlantic. Although the Isospondyli, and probably the Iniomi as well,
represent several phyletic lines of descent, with their members being correspondingly
varied in appearance, they are easily separated as a group from the other Orders that
are dealt with in Parts 3-5.
The following Key to the Orders described in these three volumes is offered solely
as an aid to identification. The reader is referred to the sections dealing with the several
Orders for information as to their internal features.
Key to External Characters of Orders Represented in
the Western North Atlantic
and Described in Parts 3-5
1a. Separate rayed caudal fin clearly marked off from anal fin, from dorsal fin, or from
both anal and dorsal.
2a. Caudal fin with conspicuous fleshy axis (marking rearward extension of verte-
bral column) bending sharply upward at base of caudal and continuing out-
ward, close to upper margin of fin, nearly to its tip; upper margin of fin much
longer than lower margin (Fig. 3). Acipenseroidei, Part 3, p. 24.
2b. Caudal fin without conspicuous fleshy axis; upper margin of fin little if any
longer than lower margin.
3a. Trunk, rearward from gill openings, completely clothed below and above
with thick rhomboid scales in mosaic pattern, interlocking but hardly
overlapping, forming a flexible armor; rear boundary of fleshy base of
caudal fin sloping obliquely rearward-upward (Fig. 3); anterior part of
upper margin of caudal and nearly entire length of lower margin
edged with a series of large ‘‘fulcral” scales, shaped as in Fig. 3.
Lepisostei, Part 3, p. 61.
3b. Trunk, rearward from gill openings, not clothed in a continuous armor
of interlocking scales; rear boundary of fleshy base of caudal either nearly
vertical or symmetrically rounded; margins of caudal fin without fulcral
scales.
4a. Point of origin of pectoral fins higher on sides than upper end of
gill openings; fin rays not branched. Giganturoidei, Part 4.
Fishes of the Western North Atlantic 18
4b. Point of origin of pectoral fins at least no higher than upper end
of gill openings, and usually much lower; at least most of fin rays
branched.
sa. Upper edge of mouth formed by the maxillary bones as well
as by the premaxillaries. Isospondyli, Part 3, p. 89,
and Part 4.
5b. Upper edge of mouth formed by the premaxillary bones
alone.
Iniomi, Part 5.
1b. No distinct rayed caudal fin; extreme tip of tail either like a whiplash or narrow
band; its membrane, if any, without rays. Lyomeri, Part 5.
“9
SoS} ENSa
TEXT AND FOOTNOTE REFERENCES
Arz, Quart. Rev. Biol., 27, 1952.
Baney, J. Morpu., 59, 1936.
Berg, Freshwater Fish. Russia, 1923 (Russ.).
Berg, Class. Fish., Trav. Inst. zool. Acad. Sci.
URSS, 5 (2), 1940; also reprinting by J. W.
Edwards, Ann Arbor, Mich., 1947 (Russ.
and Eng.).
Bertin, Bull. Soc. zool. Fr., 67 (7), 1937-
Bertin, Dana Rep., 15, 1938.
Bigelow and Schroeder, J. Mar. Res., 7, 1948.
Bigelow, Schroeder, and Farfante; Bigelow and
Schroeder, Mem. Sears Found. Mar. Res., I
(1), 1948; I (2), 1953-
Boéhlk, ef a/., Handb. Vergl. Anat. Wirtbelt.,
33 1937-
Bonaparte, Mem. Soc. Neuchatel Sci. Nat., 2,
1839.
Bronn’s Klassen, 6 (1), 1901.
Brown, eé¢ a/., Physiol. Fish., 2, 1957.
Carrer, Proc. zool. Soc. Lond., 1919.
Chapman, J. Morph., 68, 1941.
Chapman, Ann. Mag. nat. Hist., (11) 9, 1942.
Cohen, Bull. Fla. St. Mus., 3, 1958.
Cope, Trans. Amer. philos. Soc., 74, 1871.
Cuvier and Valenciennes, Hist. Nat. Poiss., 79,
1846.
Daan, Bibliog. Fish. 2, 1917.
Denison, Fieldiana, Geol., rz (8), 1956.
Denton and Marshall, J. Mar. biol. Ass. U.K.,
37 (3), 1958.
Dahlgren, Science, N.S. 27, 1908.
Dahlgren, Sci. Mon., Lond., 24, 1927.
18
24.
25.
26.
278
28.
29.
30.
Syl
BA
33:
34-
35-
36.
37:
38.
39-
40.
43.
Eaton, Copeia, 1939.
Eaton, J. Morph., 76, 1945.
Emilianov, Zool. Jb., (Anat.) 60, 1935.
Fowrer, Fish. Red Sea, 2, 1957.
Goovricx, Quart. J. micr. Sci., 47, 1904.
Goodrich, Palaeobiologica, Jena, 7, 1928.
Goodrich, Stud. Struct. Devel. Vert., 1930.
Grassé, eft a/., Trait. Zool., 13, 1958.
Gregory, Trans. Amer. philos. Soc., 23 (11),
1933):
Guitel, Arch. Zool. exp. gén., 4 (2), 1908.
Howes, Sci. Progr., I, 1894.
Hubbs, Science, N.S., 49, 1919.
Hubbs, J. Parasit, 74, 1937.
Huot, Ann. Sci. nat. Zool., (8) 4, 1902.
Huxley, Mem. Geol. Surv. U.K., Dec. 10,
1861.
‘Teo Stanf. Univ. Publ. Biol., 3 (2), 1923.
Jordan, Evermann, and Clark, Rep. WSs
Comm. Fish. (1928), 2, 1930.
Kasuxarorr, Bull. Soc. imp. Nat. Moscow,
NS 27 (1913), 1914.
Kendall and Crawford, J. Wash. Acad. Sci., 12,
1922.
Koumans, Temminkia, I, 1936.
Krogh, Osmot. Reg. Aquat. Anim., 1939.
55:
56.
57-
58.
59-
60.
6r.
62.
63.
64.
65.
66.
67.
68.
Fishes of the Western North Atlantic 1g
Lanxesrer, Treat. Zool., 9, 1909.
Linnaeus, Syst. Nat., 1758.
Mazsusara, Fish. Morph. Hierarch., Pt. 1,
1955 (Japanese).
McFarland and Munz, Biol. Bull. Woods Hole,
T4 (3), 1958.
Miclucho-Maclay, Jena Z. naturw., 3, 1867.
Millot, Nature, London, 174, 1954.
Moreau, Hist. Nat. Poiss., 2, 1881.
Moreau, Man. Ichthyol. Fr., 1892.
Miller, Abh. Akad. Wiss. Berl., Phys. Mat. K1.,
1844.
Myers, Stanf. Ichthyol. Bull., 7 (3), 1958.
Norman, Hist. Fish., 1931, and subsequent
editions.
Qrserc, Ark. Zool., 10 (12), 1957.
Owen, Vert. Anat., r, 1868.
Parmer, Ann. Mag. nat. Hist., (12) 3, 1950.
Parker and Haswell, Textb. Zool., 2 (6th ed.),
1940; and earlier editions.
Parr, Occ. Pap. Bingham oceanogr. Coll., 3,
1930.
Parr, Bull. Bingham oceanogr. Coll., 4 (6),
1934.
Parr, Bull. Bingham oceanogr. Coll., 3 (7),
1937-
Pennant, Bull. Soc. zool. Fr., 62, 1937.
Prosser, e¢ a/., Comp. Anim. Physiol., 1950.
Rayner, Philos. Trans., 233 B (601), 1948.
Regan, Proc. zool. Soc. Lond., 2, 1902.
Regan, Ann. Mag. nat. Hist., (8) 3, 1909.
Regan, Ann. Mag. nat. Hist., (8) 9, 1912.
69.
70.
7I.
88.
S9.
go.
gr.
92.
Regan, Proc. zool. Soc. Lond., 1923.
Regan, Ann. Mag. nat. Hist., (9) 15, 1925.
Regan, Encyc. Britt., 9, 1929; 9 (4th ed.),
1937-
Romer, Bull. Mus. comp. Zool. Harv., 82,
1937.
Romer, Vert. Paleont., 2nd ed., 1945.
Romer, Vert. Body, 1955.
Romer, Nature, London, 776, 1955.
Rosen, Ark. Zool., ro (8), 1916.
Scuurrz and Stern, Ways of Fish., Pt. 1, 1948.
Simpson, Bull. Amer. Mus. nat. Hist., 85,
1945-
Smith, Fish to Philospher, 1953.
Starks, Bull. Mus. comp. Zool. Harv., 52,
1908.
See also g2 below.
Steindachner and Déderlein, Denkschr. Akad.
Wiss. Wien, Math-Nat. Kl. 53, 1887.
Stensié, Triass. Fish. Spitzbergen, Pt. 1, 1921.
Stensié, Medd. Grénland, 83 (3), 1932.
"Tcnerwavin, J. linn. Soc. (Zool.), 40 (279),
1947.
Traquair, Nature, London, 62, 1goo.
Trewavas, Proc. zool. Soc. Lond., 1933.
Trewavas, et a/., Nature, London, 176, 1955.
Varerre St. George, Arch. micr. Anat., 77,
1880.
Wisratt, J. Anat., Lond., 72, 1937.
Wilimovsky, J. Paleont., 30, 1956.
Woodward (Smith Woodward), Cat. Foss.
Fish. Brit. Mus., 2, 1891.
Starks, Science, N.S. 54, 1921.
Names of Bones
GILES W. MEAD!
Museum of Comparative Zoology, Harvard University
and
MARGARET G. BRADBURY
Hopkins Marine Station, Stanford University
The bones of the skull and pectoral girdle of the hickory shad, Pomolobus mediocris
(Mitchill), are depicted in Figs. 1-3. These figures do not represent an attempt at a
critical osteological study, and the names used were chosen with regard to those most
frequently used in the descriptive accounts that follow, as well as to questions of
homology. The species illustrated may serve as an adequate guide to the bones of the
more common and conservative of the Isospondyli, but it will be of limited value in
the study of grotesque forms such as the stomiatoids and the Lyomeri. Notes on the
osteology of these, or references to such studies, are included in the accounts of the
various species.
The fishes used were caught in Chesapeake Bay north of the town of Solomons,
Maryland, in November 1958. All were between 280 and 310mm SL. One dry
skeleton was prepared and two were cleared in potassium hydroxide and stained
with alizarine. These preparations and two additional whole specimens are now in the
United States National Museum (USNM 186078, 186090, and 186091).
1. This work was done while both authors were with the Ichthyological Laboratory, U.S. Fish and Wildlife Service.
20
Fishes of the Western North Atlantic 21
dermosphenotic
dorsal postorbital
ventral postorbital posttemporal
sclerotics
supraorbitals
nasal
premoxilla
maxilla
dentary : 2g
: a opercle
lacrimal | Pp
: "J ~~ subopercle
supramaxillae J |
: reopercle
jugal PIEOP
articular. interopercle
suborbital
ectopterygoid
angular \quadrate
hyomandibular
metapterygoid
opercle
subopercle
preopercle
interopercle
interhyal
Ficure 4. Superficial face bones and suspensorium of Pomolobus mediocris. a lateral view of left side of head;
underlying bones of the neurocranium not shown. B inner view of left suspensorium, lower jaw, and opercular
series. Drawn by Margaret G. Bradbury.
se)
bo
vomer (prevomer) au
SPs} ae
mesethmoid Tg Ra
\
prefrontal i
(lateral ethmoid) wy s
oi
ee F ie
sphenotic { =
pterotic ~“S id.
parietal 7 ie
supraoccipital |
opisthotic 5 By
epiotic i TS
exoccipital
parasphenoid
basioccipital
basioccipital
Memoir Sears Foundation for Marine Research
mesethmoid
vomer (prevomer)
prefrontal
(lateral ethmoid)
frontal
orbitosphenoid
pterosphenoid
(alisphenoid)
prootic
sphenotic
parasphenoid
opisthotic
epiotic
exoccipital
basioccipital
frontal
supraoccipital
parietal
Ea: a= epiotic
. | ee a
parasphenoid
hii
prefrontal
(lateral ethmoid ) exoccipital
opisthotic
mesethmoid basioccipital
vomer (prevomer) D prootic
orbitosphenoid parasphenoid
pterosphenoid (alisphenoid) basisphenoid
Ficure 5. Neurocranium of Pomolobus mediocris. « dorsal view; B ventral view; c posterior view; D lateral
view. Drawn by Margaret G. Bradbury.
Fishes of the Western North Atlantic 22
phatngotrenciel ——
epibranchial — es
. x2 = —S=
ceratobranchial___o Se interhyal
= NG =
hypobranchial__s- ——T
— ae as: epihyal
basibranchial : eee oe LE
first
arch
oe
3
i =
glossohyal ;
hypohyals branchiostegals
ceratohyal urohyal
first (or suspensory)
pharyngobranchial pharyngobranchials
basibranchials epibranchials
hypobranchials
ceratobranchials I
posttemporal posttemporal
supracleithrum supracleithrum
postcleithra cleithrum
scapula postcleithra
cleithrum actinosts mesocoracoid
coracoid
FicuRE 6. Hyoid arch, branchial basket, and pectoral girdle of Pomolobus mediocris. a lateral view of left hyoid
arch with associated branchiostegal rays and first branchial arch; B three-quarter view of branchial arches;
c outer view of left pectoral girdle; p inner view of left pectoral girdle. Drawn by Margaret G. Bradbury.
Order Acipenseroider’
VADIM D. VLADYKOVv?
and
JORWN 2 GREBE EN
Acknowledgments. We are indebted to the late Léon Bertin and to Miss Rolande
Esteve for their help and for their permission to examine Dumeéril’s types of Sturgeons
in the Paris Muséum National d Histoire Naturelle. We are also grateful to Henry
W. Fowler and Leonard P. Schultz for making available for study the Sturgeon
collections in the Academy of Natural Sciences at Philadelphia and of the United
States National Museum; to Loren P. Woods for specimens from the Chicago
Natural History Museum; to Carl E. Guthe and Ralph S. Palmer for their kind
offer of a rich collection of Acipenser brevirostris from the New York State
Museum; to the late Alexander D. Bajkov, R. E. Dimick, Ivan f. Donaldson, and
Charles E. Warren for collecting Sturgeons in Oregon; and to Henry B. Bigelow
and William C. Schroeder for their careful editing of this manuscript and for making
available their unpublished notes.
The illustrations were drawn by Paul I, Voevodine, artist of the Department
of Fisheries, Quebec.
Scope of Study. In the following text we give the characters of the Order Aci-
penseroidei and of the family Acipenseridae; also descriptions, life histories, and geo-
graphical distributions, as well as synonyms and references, for the following Stur-
geons: Acipenser brevirostris, A. fulvescens, A. oxyrhynchus oxyrhynchus, and A. oxyrhynchus
1. Contribution No. 53, Department of Fisheries, Quebec, P. Q., Canada.
2. Present address: Department of Biology, University of Ottawa, Ontario, Canada.
3- Present adress: Division of Fish and Game, Conservation Department, Albany, New York, U.S.A.
24
Fishes of the Western North Atlantic 25
desotoi; all of these occur in the western North Atlantic or Gulf of Mexico in fresh,
brackish, or salt water. Since 4. sturio of the eastern Atlantic and Mediterranean has
sometimes been considered synonymous with 4. oxyrhynchus, the relationship of these
two is discussed on p. §7.
Most of the material for our study consisted of collections made in New York
(35: 45-103) and Quebec (83: 143-204; 84: 129-154). Supplementary data were
obtained by examination of the material at the Paris Muséum National d’Histoire
Naturelle, the United States National Museum, and the Academy of Natural Sciences
of Philadelphia.
The Descriptions are based on the Study Material listed for each species. An
explanation of terms, measurements, and counts is given on p. 33. The accounts of the
habits and geographical distributions are based upon published records and personal
observations.
Many of the earlier authors who dealt with the taxonomy of Sturgeons based their
conclusions on differences associated with the age of the specimens, with the conse-
quences that they recognized several nonexistent species. Hence, among the refer-
ences, we use only those that we consider to be the most important from a taxonomic
point of view, and particularly those dealing with the life histories of the species.
Characters. Bovy subcylindrical. Snour greatly extended, blade-like and flexible
in some, in others only slightly extended, if at all, and rigid. Mourn on lower sur-
face of head, protractile or not. CHIN BARBELS prominent in some but lacking in others.
Dorsat fin situated far rearward; only one. Caupat fin more or less deeply forked,
with fleshy axis (enclosing notochord) extending rearward to tip of upper lobe, bending
upward beginning at base of fin; heterocercal, wider below than above the fleshy axis.
VentTRAL fins well developed, abdominal in position. Fin rays much more numerous
than their basal skeletal supports. Pecrorats with the first ray transformed in some
into an ossified ray, the fins otherwise without spines. Skin of trunk rearward from
eyes seemingly naked in some but actually with minute scattered Piarezers; others
with five longitudinal rows of Scures (large bony shields), the skin roughened else-
where with small scattered Dermat ossiFications.! A continuous series of rhombic to
lanceolate Prares, closely set, on axis of caudal; the upper margin of caudal with a
dorsal fringe of large v-shaped Futcra (spine-like scales), each representing the fusion
of a pair of opposing plates. TrerH minute, if present. BRaNCHIOSTEGAL Rays absent.
SKELETON cartilaginous for the most part, the primitive cartilaginous cranium
(chondrocranium) persisting and growing throughout life; ossification chiefly repre-
senting development of membrane bones (p. 4) in regions of skull, jaws, and pectoral
girdle. Upper jaw not articulating with skull either in the ethmoid region or in the
sphenoid. Premaxillary bone fused with maxillary. Pectoral girdle fused with clavicle.
4. There is no definite terminology for the plates or granules found in the skin between the regular rows of bony
shields; they have been variously called: stellate plates, stellate roughnesses, stellate ossifications, minute rough
spinules, small prickle-like plates (43: 104-106), dermal plates, and small spinous asperities (Ryder, The Sturgeons,
with an account of experiments bearing upon Sturgeon culture, 65: 231-328). In the present manuscript they are
called ‘dermal ossifications.”’
26 Memoir Sears Foundation jor Marine Research
Myodome absent. Interopercular bone absent. Radials in dorsal and anal fins cartilagi-
nous. Vertebral centra absent, the notochord entirely unconstricted throughout its
length. Air BLADDER well developed, oval, with smooth inner surface, opening into
oesophagus through a short, wide duct. Inrest1nE with well-developed spiral valve (see
Berg, T0387):
Habitat. Fresh water, or, if marine, entering fresh water to breed.
Families. Of the two living families, the Acipenseridae with four genera is much
more important throughout the northern hemisphere than Polyodontidae, which is rep-
resented now by only two freshwater genera, each with a single species; Polyodon
spathula (Walbaum), with a nonprotractile mouth, is found in eastern North America,
whereas Psephurus gladius (Martens), with a protractile mouth, inhabits China; one
family, Chondrosteidae, is extinct, being known only from fossils of the Lower Lias
to Lower Cretaceous periods.
Key to Modern Families
Ia. Snout in front of eyes little if at all longer than remainder of head, subconical,
rigid, more or less flattened in some but not blade-like in shape; a transverse
series of four fleshy and conspicuous barbels close in front of mouth; mouth on
lower surface of head, protrusible; jaws toothless in adult; five longitudinal rows
of large bony shields (scutes) on body, the lateral row extending onto base of
caudal, the skin elsewhere between them rough with small dermal ossifications;
bony plates separated by evident sutures covering head; anterior edge of pectoral
fins with a stout ossified ray; upper lobe of caudal much longer than lower.
Acipenseridae, below.
1b. Snout in front of eyes much longer than remainder of head, blade-like in form
and flexible in American species; no barbels in front of mouth; mouth lateral in
position, protrusible (in P. g/adius) or nonprotrusible (in P. spathula); jaws with
minute teeth; no bony shields on body, the skin everywhere on trunk smooth to
the touch; head not covered with bony plates; no ossified ray in pectoral fins;
lower lobe of caudal at least as long as upper lobe in American species.
Polyodontidae. Paddlefishes.
Fresh water of eastern North America
and China.
Family ACIPENSERIDAE
Sturgeons
Characters. Bovy elongate and fusiform. Scures or bony shields in five rows:
one dorsal, two lateral, and two ventral; all scutes very sharp and strongly developed
5. According to Dr. R. D. Suttkus (personal communication, July 9, 1957), a specimen of P. spathula, 481 mm FL
and weighing 354 g, was taken in the brackish water of Vermilion Bay at Cypremont Point, Louisiana, on May 1s,
1954, by Antoine Carrere while trawling for penaeid shrimp.
Fishes of the Western North Atlantic 27,
in young individuals, but becoming progressively blunter with age or even disappearing
through absorption. Skin between scutes with small ossifications. SNour protruding.
Mourn inferior, protractile. Teer absent in adults. Barges 4, in a crossrow in
front of mouth. Grits 4, and an accessory opercular gill. BrancurosTeGaLs absent.
Git rakers fewer than 50. Operce absent. Heap covered by bony plates separated
by sutures; particularly visible in younger specimens. Dermat sKELETON without
ganoine, Caupat fin with typical fulcra. Dorsat and anat fins behind ventrals. PEc-
roraL fin with first ray enlarged and ossified. Tam heterocercal. AIR BLADDER large,
simple. Sromacu with numerous pyloric appendages, forming a compact and rather
large gland. Recrum with spiral valve.
Anadromous and freshwater fishes of the northern hemisphere; Upper Cretaceous
to Recent.
Remarks. The cartilaginous skull has an opening under the frontal and parietal
plates, usually completely roofed over by these membrane bones. Hence there is a cay-
ity, evident in sagittal section of the head, for which we propose the name frontal sinus.
In young specimens of 4. oxyrhynchus of America and J. sturio of Europe, there
is a membranous, elongated, nonossified area on the top of the head which Ryder called
the “fenestra” or “fontanelle.”” In specimens less than 40 inches long, this can usually
be found by probing with a needle between the frontal and parietal plates; thus a probe
can be passed downward into the frontal sinus. Adults, and in rare instances young, have
the plates fully joined, closing the fontanelle.
The protractility of the mouth is made possible by movements of the jaws and
palatoquadrate bone (10: 409).
The pseudobranchs are rudimentary in Scaphirhynchus (31: 26) and completely
lacking in Pseudoscaphirhynchus (11: 104). The large element of the gill cover repre-
sents the subopercle, not the opercle (70: 48-50).
The appearance of the Sturgeon changes considerably with age; the snout
becomes shorter and blunter and the shields smoother while the lower caudal
lobe, not fully formed? in very small specimens, becomes longer. Apparently the
shields may be partially or completely absorbed in connection with ripening of the
gonads.
Genera. The family Acipenseridae is divided into two subfamilies. Acipenserini
(true Sturgeons), with spiracles present, is represented by two genera: Huso, Lower
Pliocene to Recent, with two species; and Acipenser, Upper Cretaceous to Recent,
with about 16 species. Scaphirhynchini (Shovelnose Sturgeons), without spiracles,
also includes two genera: Scaphirhynchus, with two species;§ and Pseudoscaphirhynchus,
with three species.
6. Ryder was probably the first American author to describe the presence of a fontanelle in 4. oxyrhynchus (65: 233)s
for that in 4d. sturio, see Stannius (68: 26) and Antoniu (4: 113, 114).
7. Grote, et al. (36: 215, fig. 158) and Roule (62: pl. 4, fig. 7) have presented drawings of young 4. sturio in which
the lower lobe is not well grown as yet.
8. Although one of the American species has been described as a separate genus, Parascaphirhynchus (31: 37-44),
recent studies fail to substantiate this (8: 174).
28 Memor Sears Foundation for Marine Research
Key to Genera®
1a. Spiracles present; snout subconical; tail not depressed or completely mailed; gill
rakers lanceolate.
2a. Gill membranes joined to isthmus. Acipenser Linnaeus 1758, below.
2b. Gill membranes free from isthmus and broadly connected between themselves.
Huso Brandt 1869.
Basins of the Adriatic, Black,
and Caspian seas, and the Far
East (Amur River).
1b. Spiracles absent; snout shovel-shaped, depressed above; tail depressed and mailed;
gill rakers fan-shaped.
3a. Caudal peduncle completely mailed, and long, with tip of anal fin remote
from caudal base; barbels fringed. Scaphirhynchus Heckel 1835.
Mississippi River Basin.!°
3b. Caudal peduncle incompletely mailed, and short, with tip of anal fin reaching
almost to caudal base; barbels smooth. Pseudoscaphirhynchus Nikolski 1900.
Turkistan (Basin of the Aral Sea).
Genus Acipenser Linnaeus 1758
Acipenser Linnaeus, Syst. Nat., ed. 10, 1758: 2373 type species, 4. sturio Linnaeus. Atlantic Ocean, along
European coast.
Generic Synonyms:
Ichthocolla Geoffroy, Descr. de 719 Plantes, etc., 1767: 399 (presumably 4. Auso L.).
Sterleta Giildenstidt, Nov. Com. Acad. Petropol., 16, 1772: 533 (4. ruthenus L.).
Szurio Rafinesque, Indice Ittiol. Sicil., 1810: 41 (Sturio vulgaris Rafinesque = 4. sturio L.).
Sterletus Rafinesque, Ichthyol. Ohiensis, 1820: 80 (4. serotinus Rafinesque = 4. fulvescens Rafinesque).
Helops Brandt and Ratzeburg, Medizinische Zool., 2, 1833: 3, 349 (4. ste//atus Pallas = A. helops Pallas).
Sterletus Brandt and Ratzeburg, Medizinische Zool., 2, 1833: 349 (4. ruthenus L.).
Sturio Miller, Vergleich. Anat. Myxinoiden, 1835: 77 (4. sturio L.).
Antaceus Heckel and Fitzinger, Ann. Wien. Mus., 2, 1836: 293 (4. schypa Eichwald).
Lioniscus Heckel and Fitzinger, Ann. Wien. Mus., Z, 1836: 370 (4. g/aébra Fitzinger).
Sterletus Bonaparte, Cat. Metod., 1845: 21 (4. vultharius L.).
Ellops, Gistel, Naturg. Thierreichs, 9, 1848 (4. Ae/ops Pallas).
Schipa Brandt, Bull. Acad. Sci. Petersb., 7, 1850, 113 (4. schypa Eichwald).
Characters. GILL MEMBRANES joined to isthmus. SprracLes present. SnouT sub-
conical. Tar depressed and completely mailed. Gitt Rakers lanceolate. Characters
otherwise those of the family.
g. Since Sturgeons change with age in such features as head proportions and characteristics of scutes, the preparation
of keys applicable to both young and adult stages has involved selection of those characters that are most constant
throughout life.
to, According to Dr. R. D. Suttkus (personal communication, July 9, 1957), a specimen of S. album, 532 mm FL
and weighing 553 g, was taken in the brackish water of Vermilion Bay at Cypremont Point, Louisiana, on May
10, 1954, by Lester Boudreaux while trawling for penaeid shrimp.
Fishes of the Western North Atlantic 29
Subgenera. There have been several attempts to subdivide the genus Acipenser as
defined by Linnaeus in 1758. The preferences of earlier authors may be judged from
the Generic Synonyms. Among contemporary authors, Berg (1: 66), for instance, favors
three subgenera: Lioniscus Heckel and Fitzinger for 4. nudiventris, Gladostomus Holly
for A. stellatus, and Acipenser s. sR. for the rest. On the other hand, Antoniu (5: 317,
318) has grouped Black-Sea Sturgeon in two subgenera: Ewacipenser Antoniu, comprising
A. ruthenus and A. nudiventris, and Acipenser Linnaeus, embracing 4. sturio, A. giilden-
stadti, and A. stellatus. We believe that in the future any subdivision of the genus
Acipenser should be based on meristic and anatomical characters; this is not possible
yet because the anatomy of the Pacific species is completely unknown and that of
Atlantic species has not been sufficiently studied.
Size. Among Acipenser species, the range in size is wide. The largest species
is A. transmontanus, with occasional individuals weighing up to 1,800 pounds (2r:
74), as in the case of one taken about 60 years ago at Mission, British Columbia,
the smallest is no doubt 4. brevirostris, about nine pounds (p. 38). The species of
Huso are much larger. Berg (rr: 61) has mentioned the capture of a female H. huso
in the Volga in 1924 that weighed 2,707 pounds (1,228 kg). A still larger specimen,
weighing 6,614 pounds (3,000 kg), was taken in 1827. Females grow much larger
than males.
Span of Life. The species of Acipenser and Huso may have a longer life span than
most fishes. For instance, through study of cross sections of the pectoral bony ray,
an A. sturio 230 cm long was found to be 25 years old (20: 61). The age of a Huso
dauricus, 500 cm long and weighing 1,600 pounds (655 kg), was estimated to be be-
tween 50 and 55 years (60: 199). Further information is given in the accounts of the
western North Atlantic species.
Age in Sturgeons has been studied by two methods, one employing cross sections
of the pectoral ossified ray, the other otoliths.”
Breeding Habits and Sex. Sturgeons breed for the first time at a much more ad-
vanced age than most other fishes, and apparently the small species are more precocious
than the large. For instance, the relatively small 4. ruthenus from the Volga may mature
at four years (Ir: 75) and J. brevirostris from the Hudson River at five years (35:
82-83). On the other hand, such a large species as 4. oxyrhynchus reaches maturity
in 10-12 years. According to Harkness," 4. fu/vescens from Lake Nipissing, Ontario,
spawns for the first time at 22 years of age.™
Apart from the period when they are sexually mature, there is practically no
exterior distinction between the sexes. The cow Sturgeons, when full of ripe eggs,
have a greatly swollen abdomen with rather thin walls, and the mature males have a
more elongated appearance, with the walls of the abdominal cavity much thicker.
11. For schematic drawings of otoliths of 4. oxyrhynchus and brevirostris, see Greeley (35: 81); and for fulwv-e
scens, see Harkness (38: 16-19). Harkness (38: 13-42) and Greeley (35) used otoliths for determination of age
while Classen (20) and other European investigators based their findings on cross sections of the pectoral
bony ray.
12. For a review of the rate of growth and of the age at spawning of different species, see Classen (20).
30 Memoir Sears Foundation for Marine Research
Among the large species, accurate determination of the sex in specimens smaller than
30 inches is difficult and often requires the use of a magnifier.
All of the species are oviparous and spawn in fresh water, and there is no adap-
tation for the care of eggs or young. The eggs are large, at least 2.5 mm in diameter,
demersal and adhesive (prior to water-hardening), and their number varies with the
species and with the size of fish; in large specimens they exceed several million, as
in the case of the 2,707-pound female reported by Berg (above), which yielded an
estimated 7.7 million eggs weighing 542 pounds. More details are given in the ac-
counts of the respective species.
In all Acipenseridae!® the ripening gonads'* have a characteristic appearance,
being thick and elongate and extending along each side of the air bladder near the
dorsal surface of the body cavity. The immature gonads in both sexes are covered on
the underside by a large mass of yellow fat which apparently plays a role of energy
storage for the ripening gonads; in those close to spawning it is almost entirely absorbed.
The fully mature ovaries are very large and make up 10—20°/, of the total weight of
the fish. As the ripe eggs become dark, the whole ovary darkens. Mature testes
are pale yellow to whitish.
It is a common observation that, even in the spring during the spawning season,
large individuals with immature ovaries are found among fully mature females. This
may be explained by the fact that the fish, after the first spawning, may spawn only
at intervals of two or even three years.
Studies of gonad maturation in 4. nudiventris from the Aral Sea found that
there are two distinct “waves” in the ripening process (73: 581-584). Fish ascend
the Syr Darya River for spawning around the end of July, when they have rather
well developed gonads. Soon the advanced sex cells are absorbed. After about two
weeks, during the second “wave,” the maturation of gonads starts again and is nearly
complete by mid-October. During the winter the maturing fish hibernate in the river
and spawn the following April or May. However, the material at Trusov’s disposal
was rather limited (160 gonads) and was collected during four different years (1941,
1946-1948).
Habits and Food. Sturgeons are sluggish but strong. During the summer they often
leap from four to six feet above the water. There are many known cases of Sturgeons
jumping into a passing boat. About 70 years ago Ryder mentioned such a case for
A. oxyrhynchus near Delaware City, and on August 14, 1951, the Quebec newspaper
Le Soleil (p. 20) reported that a 35-pound 4. fulvescens jumped into a small rowboat
on the Ottawa River, near Pembroke, Ontario.
The small protractile mouth preceded by four sensitive barbels in the middle of
the lower surface of the head may be considered an adaptation for bottom feeding.
13. The same phenomenon has been reported for Polyodon as well (46: 116-124).
14. For further details on the ripening of eastern American Sturgeons, see Ryder (65: 268-273).
15. Nikitin (56: 76) gives practical suggestions for the recognition of eggs from different species of Sturgeon made
into caviar. The eggs may be distinguished by size, color, and particularly by the tint of their germinal disc
(“eye” of the egg).
Fishes of the Western North Atlantic ai
However, it has been observed that Sturgeon may rise to the surface to seize swimming
objects (65: 265).
The main Sturgeon food consists of bottom organisms such as oligochaete worms,
insects, mollusks, and crustaceans; occasionally plant material is found in their stomachs
as well. Sturgeons may feed on their own eggs, at least in the case of 4. fulvescens from
Lake St. Peter, Quebec (unpublished data from Messrs. L. Philippe and J.-P. Cuerrier),
and on different species of fish and birds. The gigantic Huso huso of the Caspian Sea
may even feed on newborn seals (57: 435).
Habitat. While the majority lives in both fresh and salt water and makes most of
the growth in the sea, at least two, fu/vescens and ruthenus, are found typically in
fresh water, although the former has been reported as occurring in brackish water.
Range. The species of <Acipenser are spread throughout the whole northern
hemisphere, being particularly numerous in the Caspian and Black Sea basins.
Commercial Importance. The fisheries for Sturgeon in North America have declined
greatly in the last hundred years, and their maintenance on a sustained-yield basis
constitutes a challenge to conservationists. Toward the end of the 19th century, catches
for certain years were over 5,000,000 pounds for all species combined, but in 1956
only 392,000 pounds were taken in Canada and 719,000 pounds in the United States
(combined value about $ 386,000).!° The Lake Sturgeon (4. fu/vescens) in Canada and
the White Sturgeon (4. ¢ransmontanus) in the western United States yield the greater
part of the commercial catch.
Most of the catch is consumed in the form of smoked Sturgeon and as caviar
made by curing the eggs with salt. Over the years, production of caviar has dropped”
so that in 1951 only 2,000 pounds, made from eggs of 4. fulvescens, were produced
in Canada, worth about $10,000 to the fishermen.
In Europe, particularly in Russia, Sturgeon fisheries are extensive. The fish are
valuable not only for the flesh and eggs but for several other products; the commercial
product from the spinal cord is called ‘“‘vyaziga,” and the inner lining of the swim
bladder is used for the preparation of gelatine and glue of the best quality.1s The
head of a Sturgeon, rich in gelatinous substances, is a valuable food product; in Russia
it is sold separately from the flesh under the name “‘golovizna.’”’ In the Province
of Quebec, older people use oil made from Sturgeon liver as an ointment against
rheumatism.
Although angling for Sturgeon by hook-and-line is not common, it is practiced
in some places, as around Montreal, Canada; there also is an important spear fishery
for sport for 4. fulvescens in several northern waters of the United States.1* The
16. A comprehensive summary of former catches of Sturgeon in North America is given by Rostlund (6r: 10-12).
The most recent available data for the whole continent are in “Fishery Statistics of the United States, 1956”
(74) and in “Fisheries Statistics of Canada, 1956” (26).
17. According to Ryder, in 1888 fishermen from the Delaware River produced for the German market about 50
tons of caviar from eggs of 4. oxyrhynchus (65: 278).
18. For the preparation of isinglass, see Ryder (65: 278) and Classen (19: 426-428).
1g. Some information on Sturgeon spearing, for instance in Lake Winnebago, has been reported (86: 29).
gz Memoir Sears Foundation for Marine Research
occurrence of Sturgeon scutes in materials excavated from prehistoric American Indian
villages indicates that these large fish were utilized in aboriginal times (61: 10-12).
Hybridization. Hybrids are found in nature quite frequently, not only between
different species of Acipenser but between Acipenser and Huso. Antipa described at
least eight different types of Sturgeon hybrids (3: 270-273), and recently Antoniu
added two new findings among Roumanian Sturgeon (6: 308-313). In some areas, for
instance the estuary of the Lena River, Siberia, the hybrid 4. baeri x A. ruthenus is
much more common than either of the parent species (14: 141).
Species. At present at least 16 species are recognized, five of which are found in
North America. In eastern North America there are three: 4. oxyrhynchus Muitchill
1814, A. brevirostris LeSueur 1818, and J. fulvescens Rafinesque 1817. The two found
in the Pacific coastal waters of America are 4. medirostris Ayres 1854 and AZ. trans-
montanus Richardson 1836 (p. 36).
In Europe there are five species of Acipenser: A. ruthenus Linnaeus 1758, a fresh-
water species inhabiting the rivers of the Black Sea and Caspian regions and the Arctic
basin from the Ob to the Kolyma; 4. xudiventris Lavetsky 1828, of the Black, Caspian,
and Aral seas and their tributaries; 4. gii/denstddti Brandt 1833 and Z. ste//atus Pallas
1771, found principally in the Caspian and Black sea regions, but occasionally entering
the Mediterranean; and 4. sturio Linnaeus 1758, inhabiting both the Atlantic and
Mediterranean areas. Furthermore, in his list of 4cipenser species, Magnin recognized
A. nacarii Bonaparte from the Adriatic Sea as a distinct species (48: 280). Chalikov has
described a new species from the Caspian Sea under the name of 4. primigenius
(17: 47-50), but Berg considered it a hybrid 4. ruthenus x A. giildenstadti (II: 77).
According to Professor Tamezo Mori (personal communication, December 20,
1954), seven species of <Acipenser are represented in the waters of eastern Asia:
A. schrencki Brandt 1869 is limited to the Amur River and Okhotsk Sea; 4. kikuchit
Jordan and Snyder rgo01 and 4. multiscutatus Tanaka 1908 are restricted to Japan;
A. dabryanus Duméril 1868 has been reported from only the Yellow Sea basin in
China and western Korea; and Z. sinensis Gray 1834 is known from China, Korea,
and Japan; the sixth, 4. medirostris Ayres, which is found on our Pacific coast, also
frequents the western Pacific coast; the seventh Asiatic species, 4. baeri Brandt 1869,
inhabits the Arctic Ocean basin along the coast of Siberia, from the River Ob to
Kolyma. Some information on the taxonomic characters and distribution of Asiatic
species has been given (53: 6-10; 54: 1-228; 18: 69-71; 55: 15-173 71: 65-68).
Relationship between Species. The total number of gill rakers increases in American
species in the following order: medirostris (18-20), oxyrhynchus (17-27), brevirostris
(22-29), fulvescens (25-40), and sransmontanus (34-36). The American species may
be classified in order of increasing development of their bony shields in all the rows
(using specimens of corresponding sizes) as follows: brevirostris, fulvescens, transmon-
tanus, oxyrhynchus, and medirostris. Several small plates, with or without a weak crest,
are present immediately behind the dorsal fin and in front of the anal; these plates are
typically paired in the case of oxyrhynchus but are single in the case of the two other
Fishes of the Western North Atlantic OMe)
western North Atlantic species. In American species, the development of the modified
predorsal shield increases in the following order: transmontanus (not only rudimentary,
but sometimes absent), brevirostris, fulvescens, oxyrhynchus, and medirostris. In the case
of the enlarged fulcrum along the lower lobe of the caudal and of the dermal ossifica-
tions, the order is brevirostris, fulvescens, transmontanus, oxyrhynchus, and medirostris.
Explanation of Terms, Measurements, and Counts. In order to facilitate the use of
the Key and Descriptions, a brief explanation of certain terms and methods used in
measuring and counting follows.
REGULAR ROWS OF BONY SHIELDS: The dorsal row and the lateral row on each side
are the most important from a taxonomic standpoint. The first shield in the dorsal row
corresponds to the occipital plate or nuchale, and the last is just in front of the modified
shield (see below) at the beginning of the dorsal fin. The first shield of the lateral
row, Called the supracleithrum, borders the posterior margin of the branchial cavity;
although its size may vary, it is typically larger than the next posterior shield; its
spine is usually less developed than the spines on the shields posterior to it. The last
shield of the lateral row (a small one) is situated at the posterior extremity of the
caudal peduncle, just anterior to the shields that continue on from the lateral row
along the upper lobe of the caudal.
The elongated fulcrum at the base of the lower caudal lobe (difficult to see on
large fish) is of considerable taxonomic importance when compared to the length of
the anal fin base.
There are also modified shields at the beginning of both dorsal and anal fins;
these, because of their respective positions, can be called predorsal and preanal. The
one in front of the dorsal fin is more important from the taxonomic point of view;
this modified shield is oval in shape, has no median crest or spine, and its posterior end
is somewhat bent upwards to embrace the first dorsal ray; the anterior end is not forked
but does have a slight indentation. There are two or more preanal shields; the nearest
one to the fin is very similar in shape and structure to the predorsal shield; those farther
in front, according to the species, are present in pairs or in a single file; the disposition
of these preanal shields is the same as that for postdorsal shields (see below).
Dermal ossiFicaTions: These small bony structures (ftn. 4), which vary in
size and shape with species and with age of fish, lie in the skin between the regular rows
of bony shields; they are best developed on the sides between the dorsal and lateral
rows. The degree of development, according to species, corresponds closely to that
of the regular bony shields.
ToraL LencTH (TL) is taken along the median line from the tip of the snout to
the posterior of the upper lobe of the tail when the fish is lying in a natural position.
Fork LENGTH (FL) is measured from the tip of the snout to the posterior end of the
median caudal rays—that is, to the fork of the tail. [In specimens less than five inches
(125 mm) FL, the lower caudal rays are still not fully developed and are therefore
short. The absence of the lower caudal lobe is not only typical of young oxyrhynchus
but of other Sturgeons as well. We observed this in fu/vescens; and Grote, et al. (36:
3
Memoir Sears Foundation for Marine Research
34
“g °SIyy Ul UMOYSs ‘s21750412019 “Pv
£6 °31y ur umoys ‘swassaaznf{-p Sor “Stq ur uMOYs ‘suyrudyshxo “p :yYBII 0} Yoy ‘spray Jo sMarA [eUaA *L aNNnoIy
Fishes of the Western North Atlantic eas
215, fig. 158) and Roule (62: pl. 4, fig. 7) have presented drawings of young J. sturio
in which the lower lobe is not well grown. ]
Heap LENGTH is the distance from the anterior tip of the snout to the rearmost
point of the opercle, opercular membrane excluded. SNour LENGTH is from the ante-
rior tip of the snout to the anterior margin of the orbit, the membranous rim included.
Movurtu wiprTu is the greatest transverse distance across the mouth slit, with the thick
lips excluded and the mouth closed. Posrorsirat pisTANce is that from the posterior
edge of the orbit, membranous rim included, to the rearmost point of the opercle,
membrane excluded. INTERORBITAL WIDTH is the minimum distance across the top of
the head between the bony edges of the orbits. GrLL RAKERs are counted on the outer
surface of the first branchial arch, including the rudimentary ones; they are usually
counted on the left side of the fish; in specimens smaller than eight inches (20 cm),
some rakers on the upper limb are not yet developed; therefore the counts for young
individuals are not given (see also 84: 149-154).
Bask oF ANAL FIN is the length from the structural base of the first ray to the
point where the membrane behind the last ray contacts the body. Fin ray counts
for the dorsal and anal are sometimes given, but it is difficult to count these accurately,
even after staining with alizarin and clearing in glycerin (83: 3-9); the fin membrane
is very thick, and the rays are numerous, close together, and show little bifurcation;
and in young individuals several rays are detached from the basal elements (radials).
Since the counting of rays is not only tedious but uncertain, we omit the number of
fin rays from the taxonomic characters.
Remarks on Identification, Within the same species, various characters change con-
siderably with growth; young have much longer snouts than adults, and their scutes
(shields) are sharper and closer together. Because of these natural changes it is difficult
to prepare a universal key for the specific identification of young specimens as well as
adults. There are, it is true, several fairly constant interspecific differences in anatomical
features, such as the intestinal tract, gill arches, grinding ridges within buccal cavity,
and otoliths, to mention a few. But to present these in abbreviated form seems likely
to prove confusing rather than helpful, hence they are not included in the following
Key to Species.
Key to American Atlantic and Pacific Species of Acipenser
Ia. Species of the Atlantic Seaboard.
2a. Mouth width less than 55°/, of interorbital; average difference between TL
and FL 14°/, of FL; gill rakers 17-27 (av. 21.6); postdorsal and preanal
shields in pairs; viscera pale, unpigmented.
3a. In young specimens 50-70 cm long, head length 26-28 °/, of FL; bony
shields of dorsal row oval in shape, their longitudinal length being greater
than their transverse width; carina on dorsal shields low, without a
pronounced hook; dermal ossifications between dorsal and lateral rows
“foc
36 Memor Sears Foundation for Marine Research
of shields only weakly developed; spleen short, not reaching farther back
than middle loop of small intestine.
oxyrhynchus oxyrhynchus Mitchill 1814, p. 46.
3b. In young specimens 50—70 cm long, head length 30—34 °/, of FL; bony
shields of dorsal rows rather square in shape, their longitudinal length
much shorter than their transverse width; carina on dorsal shields high,
typically with two strong hooks; dermal ossifications between dorsal and
lateral rows of shields strongly developed; spleen long, reaching much
farther back than middle loop of small intestine.
oxyrhynchus desotoi Viadykov 1955, p. 56.
2b. Mouth width over 62°/, of interorbital; average difference between TL and
FL less than 12°/, of FL; gill rakers 22-40; postdorsal and preanal shields
in a single row; viscera blackish, heavily pigmented.
4a. Gill rakers 22-29 (av. 25.4); interorbital width 34—40°/) (av. 37°/o)
of hl; dorsal shields 8—13 (av. 10); lateral shields 25—32 (av. 28.3);
dorsal and lateral shields pale, contrasting with dark background;
length, so far as known, not over 100 cm.
brevirostris LeSueur 1818, below.
4b. Gill rakers 25-40 (av. 33.1); interorbital width 29-35°/, (av. 32°/9)
of hl; dorsal shields 9-17 (av. 13.4); lateral shields 29-42 (av. 35.4);
dorsal and lateral shields brownish, of same color as background;
length commonly over roo cm. /fu/vescens Rafinesque 1817, p. 41.
1b. Species of the Pacific Seaboard.
sa. Caudal peduncle short; tip of anal fin reaching or even extending beyond base
of lower lobe of caudal; modified shield at beginning of dorsal fin rudimentary
or absent; 11-14 dorsal shields; 38-48 lateral shields; gill rakers 34-36;
viscera blackish, heavily pigmented. — tramsmontanus Richardson 1836.
Southern Alaska to California; enters salt water.
5b. Caudal peduncle long; tip of anal fin not reaching base of lower lobe of caudal;
a large modified shield at beginning of dorsal; 8-11 dorsal shields; 23-31
lateral shields; gill rakers 18-20; viscera pale, practically without pigment.
medirostris Ayres 1854.
Southern Alaska to California; enters salt water.
Acipenser brevirostris LeSueur 1818
Shortnose Sturgeon
Figures 7, 8
Study Material. A total of 109 specimens: 95, 450-900 mm TL, from the Hudson
River, New York;2° 1, presumably LeSueur’s type, ANSP 16953;% 1, from Dela-
20. For age and localities of these specimens, see Greeley (35: 89, 90).
21. This specimen, originally No. 84, was described by Ryder (65: 236) and later by Fowler (33: 604, pl. 38, fig. 1).
On June 23, 1949, the specimen (stuffed) measured 746 mm FL.
Fishes of the Western North Atlantic 37
ware River at Bay-at-Green Creek, Cape May County,
New Jersey, in salt water, May 5, 1907, ANSP 25471;
4, from Delaware River at Torresdale, Philadelphia
County, Pennsylvania, in fresh water, November
1911, ANSP 44298-44301; 3, 470-580 mm FL,
from Delaware River, March 29-31, 1913, USNM
125816-125818;22 1 young, 185mm FL, Salmon
Creek, North Carolina, USNM 64330; 1 of head,
fins, and dried strips of skin with plates, from Poto-
mac River, USNM 26273; 3 stuffed specimens in
Paris Museum.?3
Distinctive Characters. A. brevirostris is distin-
guished from J. oxyrhynchus by blackish viscera, a wider
mouth (see Key), the absence of a fontanelle, almost
complete absence of the postdorsal shields, and by
preanal shields arranged in a single row. Also, indi-
viduals of 4. brevirostris assume adult proportions
when they are only about two feet TL; 4. oxyrhynchus
retains juvenile characteristics even up to four feet. The
most conspicuous difference between drevirostris and
fulvescens is in the lateral scutes, which are much
paler than the background in the former but of the
same shade as the background in the latter.
Description. Scutes in all five main rows not
closely set and rather weakly developed in adults in
comparison with other eastern American species, but
sharp and set close together in a young specimen
7.75 inches FL (USNM 64330); in 12 specimens,
8-13 (av. 10) dorsal shields, 25-32 (av. 28) lateral
shields, and 7-11 (av. 8) ventral shields; shields be-
hind dorsal fin either in a single row or paired; elon-
gated fulcrum at base of lower caudal lobe shorter
than base of anal fin.
Heap averaging 22°/, of FL in adult, 28 °/, in
a young specimen 7.75 inches FL. Fonrane.ie ab-
22. According to the tag, these specimens were submitted as “young
Sturgeon” (presumably as 4. oxyrhynchus) for identification.
23. Details on these specimens are given by Bertin (r2: 252). Two of
these fish were sent to the Paris Museum by LeSueur, apparently
as paratypes.
Ficure 8. Acipenser brevirostris. Lateral view of spawning female,
580 mm long, from the Hudson River near Kingston, New York.
38 Memor Sears Foundation for Marine Research
sent on top of head. Snour in 22 grown specimens (390-700 mm FL), much shorter
than postorbital distance (av. 70°/) of the latter), but in a young specimen, longer than
postorbital distance (183°/)). Posrorpirar distance in adults 51—61°/y (av. 55°/,) of
hl, but 33°/) in a young specimen. INTERORBITAL width 34—40°/) (av. 37°/,) of hl, and
Mourn width (excluding lips) 69-81°/, (av. 74°/,) of interorbital width, these two
characters not differing with age.
GiLL RaKeERs rather long, triangular, in 13 specimens 22-29 (av. 25.5) on first
branchial arch; in number of gill rakers, 4. brevirostris occupies an intermediate
position between 4. oxyrhynchus with 17-27 (av. 21.5) and A. fulvescens with 25-40
(av. 33).
Cavupat fin with short upper but rather long lower lobe; average difference be-
tween TL and FL, 11°/, of FL; no notch at tip of upper caudal lobe.
Color. In preserved specimens, general color similar to that of 4. oxyrhynchus
(p. 46). Top of head and back dark, becoming paler on sides; entire lower surface
white. Central part of dorsal shields whitish, contrasting with darkly pigmented back
and black skin between shields. Lateral shields pale, clearly distinguished from sur-
rounding skin. Ventral shields of the same whitish color as lower surface of fish. Intes-
tinal tract, air bladder, and fat surrounding gonads darkly pigmented, nearly black;
the peritoneum only slightly pigmented.
In an excellent photograph of a live brevirostris, the similarity in color to oxyrhyn-
chus is particularly striking (44: 14, 15): in addition to the details observed on preserved
specimens, the white edging on the paired fins is similar; but the anal fin in drevirostris
is pigmented while that in 4. oxyrhynchus remains whitish. Small live specimens from
the Hudson River show black pigment blotches similar to those of young 4. fulvescens.
Size: This is the smallest species of all the Sturgeons. Apparently it never grows
to more than about three feet; the largest specimen recorded thus far is a female
39.5 inches TL (899 mm FL), from the Connecticut River. The measurements sub-
mitted by Professor Andrews and two specimens sent to us (p. 40) confirm his deter-
minations. The largest sizes encountered during the present studies were: a female
34-8 inches TL, weighing eight pounds 15 ounces, in its 13th year as determined by
its otoliths; and a male 25.5 inches, weighing four pounds five ounces; both were taken
from the Hudson River (35: 68—70).
Development and Growth. The early life history is unknown. Few small specimens
have been recorded; probably the smallest fish is one of 7.3 inches (about 185 mm),
from North Carolina (USNM 64330). The smallest specimens taken in the Hudson
River were two females; one was 17 inches (about 433 mm) and weighed only 15
ounces; the other was a little less than 18 inches (about 600 mm) and weighed 19
ounces. The sizes of five specimens from the Delaware River observed by Ryder ranged
between 18-23 inches. Age determinations, based on otolith readings (35: 68-70),
have shown that 4. brevirostris is a very slow-growing species. Specimens of brevirostris
from the Hudson River that measured 17-35 inches (about 430-890 mm) TL were
4-15 years old; on the other hand, 4. oxyrhynchus 74-100 long were 11-13 years of
Fishes of the Western North Atlantic 39
age. The growth rate of the two species, however, is very similar in early life, which
suggests that the subsequent divergence is due to more rapid growth by oxyrhynchus
while at sea.
Spawning. Males may mature when they are only about 20 inches TL, and most of
them do so by the time they pass 21 inches; most of the females mature at about 24
inches. The ripe eggs are dark brown. Their number is not known. Spawning takes place
in rivers early in the spring. For Hudson River fish, Greeley (35: 90) stated that:
The spawning season evidently included late April during 1936. Several fish approaching spawning
condition were examined from Rhinecliff during early April. A female, weighing five pounds, from Highland
April 21 contained eggs which appeared ripe, while a three pound male taken at the same time was spent. Roe
fish from Highland, May 1, were spent (2 pounds, 13*/, ounces; 5 pounds, 8 ounces). Several others, also
taken during early May had spawned.
General Habits. On account of its small size, 4. brevirostris has attracted little atten-
tion except when taken in nets in fresh, brackish, or salt water. It is found most often
in tidal rivers. But the place of capture of Gulf of Maine specimens shows that some
certainly go out into the open sea and wander for some distance from the parent
stream.
Food. Studies of stomach contents from Hudson River specimens showed that 4.
brevirostris feeds upon the bottom, eating small animals and plants intermingled with
mud (72: 141-144, table 4). The organisms consumed were sludgeworms, chironomid
larvae, small crustaceans, etc., such as are found in the stomachs of young Atlantic
Sturgeon (p. 52).
Abundance and Relation to Man. Because of its scarcity and small size, 4. brevirostris
is of slight interest to commercial fishermen, hence there is little information about
its abundance.
Ryder (65: 232), writing of the fishery for oxyrhynchus in the Delaware River,
stated, ‘It was my good fortune to secure no less than five specimens of the 4. brevirostris
of LeSueur, which has, so far as I can learn, not been certainly recognized since that
Naturalist’s time,” and our own Study Material (p. 36) marks it as at least not uncom-
mon in the Hudson today. But while LeSueur in 1811 stated that the Shortnose
Sturgeon is “more sought after and commands a higher price than the large common
species” in the Delaware River (75: 383-394), it is of virtually no economic importance
at present due to its small size and apparent scarcity. Its flesh, however, is of good
quality, and its eggs are suitable for caviar, but they are not very numerous. A few
fishermen prior to 1937 set gill nets for brevirostris in the Hudson River just before
the yearly run of American shad (/osa sapidissima). Some have also been taken in the
Hudson River with hook-and-line by anglers.
Range. Atlantic seaboard of North America, from New Brunswick, Canada (47:
504, and later comments), to Florida, mostly in rivers, including the Connecticut, Hud-
son, Delaware, and Potomac. This range is based upon records for a period of years,
and the species may now occupy a more restricted area. :
Until recently the southernmost definite record was for Charleston, South Carolina
40 Memor Sears Foundation for Marine Research
(43: 106). Smith stated, “while it doubtless ascends all suitable streams in North
Carolina, actual records of its occurrence are rare” (67: 58). However, thanks to Dr.
Daniel M. Cohen, we had an opportunity to examine a female 4. brevirostris (UF 5714)
735 mm TL taken on May 11, 1949, in Big Lake George of the St. Johns River
drainage, Putnam County, Florida. This is a proven record of the occurrence of the
Shortnose Sturgeon in Florida. Thus the geographic range of 4. brevirostris is almost
as extensive as that of 4. oxyrhynchus.
It has been reported as being taken as far north as the St. Lawrence River (32:
703 59: 226; 37: 44), but these records seem to be based chiefly on 4. fulvescens,
in part on 4. oxyrhynchus. The most northerly proven record of its occurrence is for
the St. John River, New Brunswick (47: 504). We also obtained some specimens from
this same area, thanks to the kindness of the late Dr. Leim. Collections taken at Gage-
town include three heads of adults caught in 1959 (collected by J. C. Medcof, Biolog-
ical Station, St. Andrews, New Brunswick); also, seven adult females (700-870 mm
TL) and three adult males (740-800 mm TL) were taken in May and early June 1960
(obtained by Fishery Officer J. O. Jenkins). One female and one male of the latter
sample were almost ready to spawn. In the Connecticut River, near Hadley, Massa-
chusetts, four adults were caught in October 1951 and September 1952 (obtained by
Professor T. J. Andrews, University of Massachusetts).
The breeding range of 4. brevirostris is not clearly defined, but it is known to
include the Hudson River, where the spawning areas appear to be very restricted.
The Delaware River may still maintain a small local population, and it seems likely
that the St. John River, N.B., has a spawning population, judging by the near-
spawning condition of the above-noted male and female taken at Gagetown. If,
through increased pollution or habitat changes, the population is no longer able to
persist in these northern rivers, the species may become dangerously reduced.
The Shortnose Sturgeon has no close relative in Europe.
Synonyms and References:*4
Acipenser brevirostris LeSueur, Trans. Amer. philos. Soc., Z, 1818: 390 (descr., Delaware R.); DeKay, Zool.
N.Y., 4, 1842: 345 (descr., ident. from Virginia not certain); Ryder, Bull. U.S. Fish Comm. (1888),
8, 1890: 231 (descr., Delaware R.); Jordan and Evermann,* Bull. U.S. nat. Mus., 47 (1), 1896: 106
(descr., synon., Cape Cod to Florida, specimen from Charleston, S. Carolina); Smith,* N.C. geol. econ.
Surv., 2, 1907: 57 (genl. acct., doubtless occur. N. Carolina); Kendall,* Occ. Pap. Boston Soc. nat. Hist.,
7 (8), 1908: 17 (New England); Fowler, Proc. Acad. nat. Sci. Philad., 1910: 604; Halkett,* Check List
Fish. Canad., 1913: 44 (genl. range of distr.); Nichols and Breder,* Zoologica, N.Y., 9, 1927: 31 (genl.
acct.); Hildebrand and Schroeder,* Bull. U.S. Bur. Fish., 43, 1928: 76 (Provincetown, Massachusetts) ;
Breder,* Field Bk. Mar. Fish. Atl. Cst., 1929: 43 (genl. acct.); Truitt, Bean, and Fowler, Bull. Md.
Conserv. Dep., 3, 1929: 33 (genl. acct.); Jordan,* Manual Vert. Anim. NE U.S., 1929: 33 (distr., key,
and brief characteristics); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 34
(in checklist); Jordan and Evermann, Amer. Food Game Fish., 1937: 12 (genl. acct., key); Greeley,
N.Y. Biol. Surv. Lower Hudson Watershed (1936), 11, 1937: 90 (size ranges, age determ., Hudson R.);
Schrenkeisen, Field Bk. Freshw. Fish. N. Amer., 1938: 14 (genl. acct.); Vladykov and Beaulieu, Nat.
24. We follow the lead of Jordan, et al. (45: 34) in calling this species brevirostris. The authors who used the spelling
brevirostrum are marked by an asterisk (*) following their names.
Fishes of the Western North Atlantic 41
Canad., 73, 1946: 43 (detail. acct. of shields and gill rakers, size); Nat. Canad., 78, 1951: 199 (detail.
acct. of gill rakers for three w. Atlantic species).
Acipenser (Huso) lesueurii Valenciennes (Ms.) Duméril, Hist. Nat. Poiss. 2, 1870: 166 (descr., New York).?°
Acipenser (Huso) rostellum Duméril, Hist. Nat. Poiss., 2, 1870: 173 (descr., New York?).7°
Acipenser (Huso) sinus Valenciennes (ms.) Dumeéril, Hist. Nat. Poiss., 2, 1870: 175 (descr., New York?).?°
Acipenser fulvescens Rafinesque 1817
Lake Sturgeon,® Smoothback
Figures 7, 9
Study Material. Over 800 specimens, 6—229 cm or 2.5—g0 inches TL (5.7—
211 cm or 2.3—83 in. FL), from different sections of the St. Lawrence River, Quebec;
also odd specimens from Lake Ontario on both the Canadian and New York sides.
Specimens from a large number of interior waters not included. For details, see Vla-
dykov and Beaulieu (83: 18).
Distinctive Characters. A. fulvescens, like A. brevirostris, is distinguishable from
A. oxyrhynchus by: the arrangement of its preanal shields in a single row, a mouth
not less than 66°/, as wide as the interorbital width, the absence of a soft ‘area
(fontanelle, p. 27) on the top of the head, and black viscera. From drevirostris it
differs in that its lateral ‘shields are of the same color as the background.
Description. ScutTEs in young specimens (up to 12 in. FL), in all rows, developed
even more strongly than in 4. oxyrhynchus of corresponding sizes; progressively ab-
sorbed with age, beginning with the ventral rows, until they may be completely so, near
spawning time, especially in the ventral rows (hence the commercial name Smooth-
back); in 516 Quebec specimens, 9-17 (av. 13.4) dorsal shields, and in 242 specimens,
29-42 (av. 35.4) lateral shields; shields behind dorsal fin in a single row; elongated
fulcrum at base of lower caudal lobe never longer than base of anal fin. Dermat
OSSIFICATIONS minute, but the skin much tougher than in 4. oxyrhynchus.
Heap longer in young than in larger individuals, on the average 24°/, of FL in
specimens 40—70 cm FL. Snour in specimens less than 50 cm FL longer than post-
orbital distance, but shorter in older individuals. Inrerorsirat width rather large,
varying; in 79 young specimens 8—36 cm FL, 21.6-31.8°/, (av. 28°/,) of hl; in 143
older specimens 40-211 cm FL, 29-40°/, (av. 34°/,). Mourn large, its width (ex-
cluding lips) 66—93°/, (av. 77°/9) of interorbital width.
GrLt RAKeERs rather short and blunt, but numerous; 350 Quebec specimens above
8 inches FL with 25-40 (av. 33.1) rakers.
25. The Duméril holotypes of these three species are kept mounted in good condition at the Paris Muséum National
d’Histoire Naturelle. They were examined in 1950 and identified as 4. brevirostris by Vladykov. A similar identi-
fication was made by Bertin (r2: 252). Jordan, ef al. (45: 34) quoted two more names given by Duméril
(28: 164-168), based on locality only: microrhynchus (New York) and dekayi (New York).
26. In Quebec, the French-speaking fishermen call the adults Esturgeon jaune or Camus, that is, short and flat
snout; the latter name was already in use at the end of the last century (52: 188). The young specimens are usually
named Escargot, or in certain sections (Nicolet) Charbonnier, on account of the black blotches (83: 32). On the
New York market, this species is often called the Smoothback.
42 Memor Sears Foundation for Marine Research
CaupaL fin with rather short upper but long lower
lobe; the average difference between TL and FL 9°/, of
FL; no notch at tip of upper lobe.
Color. There is a pronounced difference between the
young and the adult. In young below 300 mm (12 in.) FL,
two pairs of large black blotches present on anterior half
of upper surface of snout and on back between dorsal and
lateral rows of shields, the anterior blotch the larger and
immediately behind the base of pectoral fin, the smaller
posterior one below the dorsal fin;”” in addition, several
small, irregular, black spots on top of head, on back and
sides, and on lower surface of caudal peduncle. Intestinal
tract darkly pigmented, nearly black. Air bladder and fat
surrounding gonads, dark gray. Peritoneum pale or only
slightly pigmented. In general, the dark pigmentation of the
intestinal tract is similar to that of 4. brevirostris but very dis-
tinct from that of 4. oxyrhynchus. In half-grown specimens
up to about 24 inches or 610mm FL, the large blotches
have disappeared, but black spots may persist. Older in-
dividuals above 40 inches FL are uniformly dark brownish
or grayish on top of head and on back and sides. In fish of
all ages, dorsal and lateral shields of the same color as back-
ground; shields of ventral rows sometimes pigmented on
upper part but always whitish on lower part. Lower surface
of young and adults uniformly whitish. In young below 24
inches FL or about 610 mm, preserved in 4—5°/, formalin,
lower surface and sides of head and body clear green.
At all ages, fins dark brown or gray, typically without
white edging.
Size and Growth. Among western North Atlantic spe-
cies, 4. fulvescens reaches a size exceeded only by 4. oxyrhyn-
chus. The largest recorded is a female Lake Sturgeon, of
approximately 275 pounds, taken in Lake Winnipeg, Mani-
toba, in the summer of 1941 (40: 17). The largest male,
caught in Lake Erie in 1948, weighed 220 pounds (66: 7).
The specimens from Quebec listed in Study Material, 2.5—90
inches TL, weighed 0.032 ounces—212 pounds (0.9 g—96
27. For a good illustration of these black blotches in young A. fulwescens, see
Forbes and Richardson (3r: 36); for color illustrations of an adult, see Vla-
dykoy (82: 6).
Ficure 9. Acipenser fulvescens. Lateral view of immature female, 655 mm
long, from the St. Lawrence River near Neuville, Quebec, P. Q., Canada.
Fishes of the Western North Atlantic 43
kilo).28 Data for the smallest 11 specimens from the St. Lawrence River, taken from
August to November in péches located near Quebec City, are tabulated in Table 1.
Judging by their long snouts, dorsal scutes in a continuous row without separation,
general coloration, and small size, we consider all of these specimens to be less than
one year old. During the first year, juveniles of 4. fulvescens (Table 1) and of 4. oxy-
rhynchus grow at about the same rate.
Table I. Growth Rate of 4. fulvescens.
Date Total Length Fork Length Weight Place
(mm) (inches) (mm) (inches) (g) (0z.)
Sept. 19, 1952... 63 2.48 57 2.24 0.9 0.032 St. Nicolas
Octr1o3) 1953-1 84 3.31 82 3.23 2.0 0.071 St. Nicolas
Aug. 8, 1947... 87 3-43 79 Ziti 2.5 0.088 Neuville
Nov. 19, 1947... 99 3.90 go 3-54 3.2 0.113 Neuville
Oct, 1os2--- 106 4.17 98 3.86 4.4 0.155 Neuville
Sept. 12, 1944... 109 4.29 96 3.78 4.3 0.152 Neuville
Aug. 20, 1952... 117 4.61 107 4.21 4.8 0.169 Neuville
Sept. 30, 1947... I2I 4.76 109 4.29 4.7 0.166 Neuville
Aug. 29, 1947... 121 4.76 III 4.37 Tel: 0.271 St. Vallier
Auge 255 LO5 2s 4 123 4.84 I12 4.41 6.5 0.229 Neuville
Sept. 30, 1947... 123 4.84 112 4.41 6.2 0.219 Neuville
AVERAGE ........ 104.8 4.13 95-8 3-77 4.29 0.151
Although the ages of Lake Sturgeon in our material from the St. Lawrence River
have not as yet been determined, probably their range is up to 30 years, or even more.
According to Cuerrier and Roussow, who studied the growth of fu/vescens from the
upper St. Lawrence River (Lake St. Francis), fish 3-22 years old averaged 16.5—37.5
inches FL (24: 8-14). The oldest fish in their material, a female 23 years of age,
was 50.5 inches FL and weighed 42 pounds.
Spawning and Reproduction.2® The principal spawning season for 4. fu/vescens in
Quebec extends from about the beginning of May to the end of June. The smallest
male ready to breed, taken at St. Vallier on May 5, 1944, was 38 inches TL (35 in. FL)
and weighed 11 pounds 4 ounces. The smallest mature female, taken at the same place
on June 13, 1949, was 45 inches TL (41 in. FL) and weighed 17 pounds. The Lake
Sturgeon from Lake Nipigon, Ontario, does not become sexually mature until it is
about 22 years of age, when it is approximately three feet TL (38: 15-25).
Roussow, in studying cross sections of the first pectoral ray of 4. fulvescens,
found variations in the broad and narrow zones on this bone (64: 553-572). He con-
cluded that “‘the growth retardations caused by the ripening of the gonads and the
28. For the length-weight relationship of St. Lawrence specimens from Quebec, see Vladykov (78: 57). Roussow
has given some information on the rate of growth and movements of 4. fulvescens from the St. Lawrence
and Ottawa rivers (63: 1-124). Results of extensive tagging at Quebec of 1,507 4. fulvescens and 2,995 A. oxy-
rhynchus have been summarized by Magnin and Beaulieu (50: 237-252).
29. Additional information on spawning habits and experimental hatching of Lake Sturgeon has been given by
Stone (69: 118-123) and Carter (16: 60-63).
44 Memoir Sears Foundation for Marine Research
losses at the consecutive spawnings are visible in the ray sections in the form of ‘belts’
of 2 to 7 narrower zones (or annuli). These indicate that the interval between spawning
periods can vary from 4 to 7 years and more.” It is highly desirable to corroborate these
findings by observing the frequency of spawning by tagged specimens.
The principal spawning rivers in Quebec are the St. Francis for Lake St. Peter,
and the Batiscan and Chaudiére for the respective sections of the St. Lawrence from
Three Rivers to Montmagny.
The diameter of mature but unfertilized eggs preserved in 4—5°/, formalin varied
between 2.7—3.1 mm. Thus the ripe eggs of 4. fulvescens are somewhat larger than
those of 4. oxyrhynchus. Table 11 gives data on ovaries and eggs from mature females
of 4. fulvescens from the St. Lawrence River near Quebec.
Table II. Data on Ovaries and Eggs of 4. fulvescens.
TL ee FL Weight Wet. of Ovaries ea
Locality Date (cm) (cm) (in.) (kg) (Ib.) (kg) (Ib.) (mm)
Stavalliermmpunenss Odd rticte eioraler= 136 127 50.0 i7au ity hae 6.9 2.7
Stavallicnmm Miaysncs ol QS O-nerctrersierel = 144 136 53-5 26.3 58 4.7 10.4 2.8
steele IMENTS) SO\Fec none bhooeooc 142 133 52.4 19.1 42 2.9 6.4 2.9
St Valier el Une 75) LOA Ollila 145 135 53-1 22.2 49 — _— 3.0
St VallieraVlayetgsetO4 Omer r 175 170 6=—-_ 66.9 50.3* 100* — — 3.0
St. Nicolas May 8, 1949............. 170 163 64.2 37-0 81 II.2 24.6 3.1
* Approximate.
Food. In the St. Lawrence, 4. fulvescens subsists on about the same type of food
as A. oxyrhynchus (p. §2). In 29 stomachs of fu/vescens we found that larvae of the bur-
rowing May fly (Hexagenia) were present in 76°/, of the samples, amphipods in 73°/o,
gastropod mollusks in 48°/,, bivalve mollusks in 35°/,, and fish in 24°/,. Indeed,
A. fulvescens of different sizes are easily taken by hook-and-line, and the best bait is
young shad, or, as second choice, banded killifish (Fundulus diaphanus). Harkness®
found that 4. fulvescens in Lake Nipigon, Ontario, fed principally on May fly and
chironomid larvae and on mollusks.
Abundance and Commercial Importance. According to official statistics for 1951, the
catches of Lake Sturgeon were (in pounds): 9,000 for Saskatchewan, 164,000 for
Ontario, 194,000 for Quebec, and 13,000 for the U.S. Great Lakes; total, 380,000
pounds, with a combined monetary value of $300,000. Late nineteenth-century yields
were several times greater. From Lake Michigan alone, in 1880, the catch was 3,839,600
pounds (32: 26). Lake of the Woods (Minnesota and Canada) was formerly called
“the greatest sturgeon pond in the world”; the catch there in 1893 was 1.6 million
pounds (29: 121-136).
In the St. Lawrence River, from the Ontario border to Cap St. Ignace at Quebec,
30. Harkness (38: 28, 29) has given references to other authors who studied food requirements of 4. fulvescens from
various localities.
Fishes of the Western North Atlantic 45
there is a regular fishery for this species; at Quebec, although present catches are less
abundant than formerly, 4. fu/vescens is still taken in fair quantity; and it is reported
as being still very plentiful in the Hudson Bay area. Large 4. fu/vescens are taken early
in the spring or late in the fall. During the summer months, in the vicinity of Quebec
City from St. Nicolas to Montmagny, 4. oxyrhynchus is more plentiful than fulvescens;
however, both species are taken together quite often at the beginning and end of the
fishing season. The fish are caught with either gill nets (8—10 in. stretched mesh) or
weirs, and a few are taken on set lines baited with fish, or in New York on unbaited
set lines (“trap lines’’).
Relation to Man. Considering its entire range, the Lake Sturgeon is at present of
much greater commercial value than both the American Atlantic Sturgeon and the
Shortnose Sturgeon combined. The principal product is the meat for “smoked
Sturgeon.” In 1951, in Canada, 2,000 pounds of caviar were made from the eggs of
this species.
Range and Occurrence in the Western North Atlantic. Larger rivers and lakes, from
Hudson Bay to the Mississippi Valley; according to Hubbs and Lagler (47: 30),
“from the Red River of the North, the Saskatchewan River in Alberta, and the Hudson
Bay, St. Lawrence and Lake Champlain drainages of Canada (and possibly from Labra-
dor), southward, west of the Appalachian Mountains, to the Tennessee River of Ala-
bama, to Missouri and to eastern Nebraska.’’ On several occasions it has been reported
as being taken on the shores of Hudson Bay (75: 30 [42]).*1 In the St. Lawrence River,
where a general line of demarcation between fresh and brackish water can be drawn
from Cap Brulé on the north shore to St. Jean-Port-Joli on the south shore, it is taken
in commercial quantities from Lauzon to Cap St. Ignace, together with oxyrhynchus.
Occasionally Lake Sturgeon venture even farther into brackish water. For instance, on
October 10, 1954, an 4. fulvescens weighing five pounds was taken in a péche at St. Roch-
des-Aulnaies; information from Mr. Alfred Pelletier, proprietor. For further details
see Vladykov (79: 54-57). It should be emphasized, however, that a sea-resident
stage, characterized by a rapid growth rate, has not been demonstrated for this species.
Synonyms and References:
Acipenser fulvescens Rafinesque, Amer. Mon. Mag., 1, August, 1817: 288 (descr., Great Lakes); Hubbs,
Copeia, 1917: 48 (synon.); Hubbs, Misc. Publ. Mus. Zool. Univ. Mich., 75, 1926: 8 (in checklist);
Dymond, Univ. Toronto Stud. Biol., 27, 1926: 33 (descr., Lake Nipigon, Ontario); Dymond, Hart,
and Pritchard, Univ. Toronto Stud. Biol., 33, 1929: 13 (statist. of catches); Jordan, Manual Vert.
Anim. NE U.S., 1929: 33 (distr., key, and brief characteristics); Jordan, Evermann, and Clark, Rept.
U.S. Comm. Fish. (1928), 2, 1930: 35 (in checklist); Greeley, N.Y. Biol. Surv. Champlain Watershed
(1929), 4, 1930: 73 (Lake Champlain); Bajkov, Trans. Amer. Fish. Soc. (1930), 60, 1931: 11 (statist.
of catches, Manitoba); Greeley, N.Y. Biol. Surv. St. Lawrence Watershed (1930), 5, 1931: 81 (St.
Lawrence R., New York); Bajkov, Trans. Amer. Fish. Soc. (1930), 60, 1931: 11 (Manitoba, statist.
of catches); Gowanloch, Bull. La. Conserv. Dep., 23, 1933: 412 (genl. acct.); Vladykov, Contr. Canad.
Biol., 8 (29), 1933: 30 (42) (Hudson Bay region); Schrenkeisen, Field Bk. Freshw. Fish. N. Amer.,
1938: 13 (genl. acct.); Kuhne, Guide Fish. Tennessee, 1939: 17 (genl. acct.); Dymond, Contr. R.
31. A young specimen of A. fulvescens about 107 mm FL (USNM 27784) was taken by Walton Hayden at Moose
Factory, James Bay (a southern extension of Hudson Bay), no doubt in brackish water.
46 Memoir Sears Foundation for Marine Research
Ontario Mus. Zool., 15, 1939: 8 (statist. of catches, Ottawa R.); Hinks, Fish. Manitoba, 1943: 15
(Manitoba); Eddy and Surber, North. Fish. Upper Mississippi Valley, 1943: 57 (size, artif. propag.,
Mississippi R.); Vladykov and Beaulieu, Nat. Canad., 73, 1946: 27 (detail acct. of shields and gill rakers,
size, Quebec); Hubbs and Lagler, Bull. Cranbrook Inst. Sci., 26, 1947: 30 (distr.); Dymond, Misc.
Publ. R. Ontario Mus. Zool., 1, 1947: 4 (genl. distr., range); Vladykov and Beaulieu, Nat. Canad., 78,
1951: 129 (detail. acct. of gill rakers for three w. Atlantic species); Scott, Freshw. Fish. East. Canada,
1O54: 7
Acipenser rubicundus LeSueur, ‘Trans. Amer. philos. Soc., 7, 1818: 38 (descr., Lakes Ontario, Erie and all
upper lakes); DeKay, Zool. N.Y., 4, 1842: 344 (descr.); Jordan and Evermann, Bull. U.S. nat. Mus.,
47 (1), 1896: 106 (descr., synon., distr.); Carter, Trans. Amer. Fish. Soc., 34, 1904: 60 (artif. propag.
Vermont); Evermann and Goldsborough, Proc. biol. Soc. Wash., 20, 1907 (Canada); Kendall, Occ. Pap.
Boston Soc. nat. Hist., 7 (8), 1908: 16 (Lake Champlain, Vermont); Nash, Vertebr. Ontario, 1908:
14 (genl. acct.); Kendall, Proc. Portland Soc. nat. Hist., 2, 1909: 219 (checklist, Labrador); Halkett,
Check List Fish. Canad., 1913: 44 (Canada); Forbes and Richardson, Fish. Illinois, 1920: 24 (descr.,
statist. of catches); Harkness, Univ. Toronto Stud. Biol., 24, 1923: 13 (age, food); Kendall, Contr.
Canad. Biol. (1923), 23, 1924: 21 (439) (between Lake Winnipeg and Hudson Bay); Breder, Field
Bk. Mar. Fish. Atl. Cst., 1929: 43 (genl. acct.); Jordan and Evermann, Amer. Food Game Fish., 1937:
10 (genl. acct., key).
Acipenser maculosus LeSueur, Trans Amer. philos. Soc., I, 1818: 393 (descr., young from Ohio R.).
Acipenser serotinus Rafinesque, Ichthyol. Ohiensis, 1820: 80 (Ohio R.).
Acipenser ohiensis Rafinesque, Ichthyol. Ohiensis, 1820: 81 (Ohio R.).
Acipenser macrostomus Rafinesque, Ichthyol. Ohiensis, 1820: 81 (Ohio R.).
Dinectus truncatus Rafinesque, Ichthyol. Ohiensis, 1820: 82 (erroneous, based on drawing by Audubon).
Acipenser rupertianus Richardson, Fauna Boreal. Amer., 1836: 311 (specimen from Albany R., Rupert
Land).
Acipenser laevis Agassiz, Lake Superior, 1850: 267 (Lake Superior).
Acipenser carbonarius Agassiz, Lake Superior, 1850: 271 (Lake Superior).
Acipenser rhynchaeus Agassiz, Lake Superior, 1850: 276 (Lake Superior).
Acipenser liopeltis Giinther, Cat. Fish. Brit. Mus., 8, 1870: 341 (Mississippi).
Nominal specific names, given by Duméril (32: 105, 220) to specimens of 4. fu/vescens, according to locality
only: copei (supposedly upper Missouri), rauchi (Osage R., Missouri), richardsoni (upper Mississippi),
anasimos (Missouri R.), parazasimos (Huntsville, Alabama), azthracinus (Lake Erie), megalaspis (Lake
Champlain), /emarii (supposedly Lake Erie), ate/aspis (Saskatchewan R.), rafinesquii (Ohio R.), rosarium
(supposedly Lake Erie), p/atyrhinus (supposedly upper Mississippi), Airt/andi (Lake Erie), mertinianus
(Michigan), cincinnati (Ohio R.), and duffalo (Lake Erie).
Acipenser oxyrhynchus oxyrhynchus Mitchill 1814
Atlantic Sturgeon, Sea Sturgeon, Common Sturgeon*®
Rigures 3, 75080
Study Material. Over 3,000 specimens: 1, ca. 485 mm (19 in.) FL, New Brit-
ain, Florida, Jan. 6, 1900, CNHM 35376 (received through kindness of Loren P.
Woods); 42, 200-2,490 mm TL, Hudson River, New York**; the remainder, 65—
32. According to the List of Common and Scientific Names (rz: 7), oxyrhynchus should be called the Atlantic Stur-
geon, but the name Sea Sturgeon is more generally used. In the Hudson River adults are known as the Big
Sturgeon and small ones as Sharpnose Sturgeon or Pelican (35: 89). In Quebec, French-speaking fishermen
commonly call the larger ones Esturgeon noir, or Esturgeon d’eau salée, while the young are known as Escargots
or Ecailles (83: 43).
33- For age and size of these specimens, see Greeley (35: 68, 139).
Fishes of the Western North Atlantic 47
2,670 mm TL (60-2,500 mm FL), St. Lawrence River,
Quebec, between Three Rivers and Riviére-aux-Outardes.*4
Distinctive Characters. A. oxyrhynchus oxyrhynchus is
readily distinguishable from both Jdrevirostris (Shortnose
Sturgeon) and fulvescens (Lake Sturgeon) by the arrange-
ment of its preanal shields in a double row, by its pale un-
pigmented viscera, by a mouth less than 55°/, as wide as
the space between the orbits, and by the presence of a soft
area (fontanelle) on the top of its head. It differs from /. oxy-
rhynchus desotoi (p. §6) in that its bony shields are oval and
longer than broad, and that the carina on its dorsal shields
do not have a conspicuous hook. Among western Atlantic
species, furthermore, 4. oxyrhynchus has much the longest
upper caudal lobe but a rather short lower one. In 4. oxy-
rhynchus the average difference between TL and FL is 14°/,
of FL, in 2. dbrevirostris 11°/,, and in A. fulvescens 9°|,.
For a discussion of the differences between oxyrhynchus and
sturio of the eastern Atlantic, see p. 57.
Description.** Scutss strongly developed in all longitu-
dinal rows; in 1,814 Quebec specimens, 7—13 (av. 9.8) dorsal
shields, and in 692 specimens, 24-35 (av. 28.7) lateral
shields; shields behind dorsal fin in pairs; elongated fulcrum
at base of lower caudal lobe decidedly longer than base of
anal fin. Dermat ossiFIcaTIoNs also well developed on skin
between dorsal and lateral rows.
Heap elongate. A membranous FonTaNELLe between
frontal and parietal plates in young individuals, and a sinus
at all ages. SNout longer than postorbital distance in indi-
viduals up to 95 cm, but in older specimens shorter than
postorbital distance. INrERoRBITAL width rather narrow, var-
ying; in 239 young specimens (6-49 cm), 22.4-28.4°/,
(av. 26°/,) of hl, in 439 older specimens (s0—231 cm), 25—
35.6%) (av. 29°/,). Mourn narrow, its width (excluding
lips) less than 55°/, of interorbital width.
GILL RAKERS among specimens from the St. Lawrence
River, Quebec (studied by Vladykov and Beaulieu, 83: 43-
34. For origin, size, and method of fishing, see Vladykov and Beaulieu (83: 43-47).
35- In part from Vladykoy and Beaulieu (83: 43-47), but mainly from an unpub-
lished manuscript by the same authors on the morphometric studies on Quebec
Sturgeons.
Ficure 10. Acipenser oxyrhynchus. Lateral view of immature male, 581 mm
long, from the St. Lawrence River near St. Vallier, Quebec, P. Q., Canada.
we
/
Cie
48 Memoir Sears Foundation for Marine Research
47): total number on first branchial arch in 70 young below eight inches, 15-24 (av.
20.4); in 800 large individuals 8-91 inches, 16-27 (av. 21.5). This difference was
due to the fact that in young specimens not all rakers were developed as yet on the
upper limb. Therefore, for purposes of comparison of gill raker counts, only individuals
above eight inches FL should be used. In 70 large specimens of 4. oxyrhynchus, 9-14
(av. 11.5) rakers on lower limb and 7-13 (av. 9.8) on upper limb.
Caupalt fin with average difference between TL and FL, 14°/, of FL; a notch
at tip of upper caudal lobe in undamaged specimens.
Color. There is no significant difference in coloration between young specimens
and adults.%* Back and top of the head bluish-black, becoming progressively paler
down the sides; entire lower surface whitish. Median carina and spine on dorsal shields
(in dorsal row) whitish, contrasting strongly with darkly pigmented back, particularly
with the jet-black skin covering the extremities of each shield, the dorsal row thus
appearing as a series of whitish spots on a dark background. Upper half of lateral
shields of same color as background, but central and lower part of these shields whitish,
contrasting well with pigmented sides of fish, but not as sharply as in the case of the
dorsal shields. Sides of head and body, including ventral row of shields, white. Iris
pale golden. Anterior edge of pectorals, ventrals, and lower caudal lobe white, clearly
defined against remaining dark grey part of these fins; anal fin entirely white; dorsal
fin and upper caudal lobe dark grey or blue-black in the central portion, their broad
posterior margins whitish. Intestinal tract and peritoneum nearly white, never darkly
pigmented.
The coloration of the three species that occur on the Atlantic coast of America,
when preserved in 4—5°/, formalin, is characteristic. In 4. oxyrhynchus, the ventral sur-
face and sides of the fish, even after a long period in perservative, are whitish or faintly
pinkish. The skin of the two other species (brevirostris and fulvescens), however, soon
acquires and permanently retains a green color that is particularly pronounced on the
unpigmented parts of the body, such as the ventral surface, sides of the head, etc.
Size. Ryder found that females averaged about 8 feet TL, and sometimes
reached 10 feet, whereas males ranged between 6—7 feet (65: 268). Weights and total
lengths of two specimens taken in the Hudson River in 1936 and recorded by the
New York State Conservation Department were as follows: unspent male 7 feet 4
inches, 176 pounds; a spent male 8 feet 4 inches, 199 pounds 15 ounces. For the Gulf
of Maine, Bigelow and Schroeder (73: 82) stated that “‘about 12 feet is perhaps the
greatest length to be expected today. But 18 feet, reported for New England many
years ago, may not have been an exaggeration. ... The heaviest Gulf of Maine Sturgeon
reliably reported (to our knowledge), was one of 600 pounds, landed in Portland by
the steam trawler Fabia from Georges Bank, December 21, 1932.” A 14-foot female,
weighing 811 pounds, was caught at Middle Island Maguerville, about 64 miles off
the estuary of the St. John River, New Brunswick, in July 1924.%7
36. For color illustrations of an adult Atlantic Sturgeon, see Vladykov (82: 7).
37. Caught by Mr. John Neilson of Moncton. This information was kindly furnished by Mr. J. Raymond Tippett,
Moncton, New Brunswick, a Sturgeon fisherman himself.
Fishes of the Western North Atlantic 49
The St. Lawrence River specimens noted in the Study Material, 65-2,670 mm
TL, weighed 0.001 5-352 pounds (0.7—160 kg). Every year in the St. Lawrence River,
Quebec, some Atlantic Sturgeons weighing between 200 and 300 pounds are taken
by local fishermen. The two largest specimens at our disposal were taken in the
brackish section of the St. Lawrence River at Ile-aux-Patins in a fascine weir.** A spent
female®® 8 feet 3 inches TL (7 ft. 7 in. FL) and weighing 203 pounds was taken
July 5, 1946. Another female, almost ready to spawn, 8 feet 9 inches TL (8 ft. 2.5
in. FL) and weighing 352 pounds, was caught June 23, 1954.
Development and Growth. Eggs hatch in about one week at a temperature of 17.8°C
(64°F). In one of the New York hatcheries where some fry were kept until they were
about an inch long, the yolksac apparently lasted for about six days.‘ Unfortunately, the
temperatures were not recorded. The newly hatched fry are about 11 mm (0.4 in.)
long. The later growth of young oxyrhynchus has not been followed, but in Europe
A. sturio reach a length of 4—5.5 inches in two months.
Little is known about the early stages of oxyrhynchus in nature, but it appears
that yearlings grow rather rapidly, for in freshwater sections of the St. Lawrence River
in Quebec we collected some young specimens ranging between 60-96 mm FL
that we consider to be less than one year of age (see Table 111). The smallest of
Table III. Data for the Five Smallest Specimens from Fresh Water,
St. Lawrence River.
Date Place TL FL Weight
(mm) (mm) (g)
Aug. 8, 1945 Berthier-en= bass treshiwatene tei elise sielet= selec 65 60 9:7
Aug. 24, 1950 Sie Walllisy, inailn WEl@egnen chanaondaocns sdadacpuee 96 87 3-4
Sept. 26, 1944 Berthier-en-Bas, fresh waters. ..00).0<0csicc es cecees 103 89 4.3
Sept. 26, 1944 Berthier-en-basytreshy water eierieie eisie se erei-l= 1 -0- 102 gi 4.0
Sept. 22, 1944 Riviere! @ wells seus cicraietsyoussoyties sv sretscorsietane-s cs sisisie alors I10 96 4.2
these was tangled in one corner of a net with 8-inch stretched mesh, set in 60 feet of
water. The others were taken in péches, that is, weirs made from chicken wire or
fascines. Berthier-en-Bas and St. Vallier are on the freshwater section of the St. Law-
rence River. Riviére Ouelle is situated at the estuary of the river of the same name,
which empties into the brackish section of the St. Lawrence. Additional information
about the growth of young fish that we consider to be yearlings is given in Table trv.
Determinations of the apparent growth rates of 4. oxyrhynchus tagged in the
St. Lawrence are made difficult because of damage to their tails.41 However, instances
38. Taken by Mr. Maurice Ouellet, Kamouraska, Quebec, a Sturgeon fisherman.
39- The ovaries of this fish weighed 30 pounds.
40. Eighth Annu. Rep. Comm. Fish., New York (1875), 1876: 12-14.
41. In Quebec, fishermen who catch Sturgeon in a péche built in salt water at St. André de Kamouraska carry them
alive in a horse-drawn cart (“tombereau”’ locally) for about 30 minutes and dump them in a freshwater brook.
But if one of them is a tagged fish, it is measured and weighed and returned to the sea by horse-cart after a
sojourn of a few hours to one or two days in the brook. On a hot summer day, fish in the cart thrash so
4
50 Memoir Sears Foundation for Marine Research
Table IV. Sizes and Weights of Young Taken during 1944 in a Péche at
Ste. Famille Ile d’Orléans, near Quebec City.
No. of Fork Length Weight
Date Specimens (mm) (g)
Range Average Range Average
INVERSE Bk oocacsncoosanDDoLeUs 25 130-144 122.8 6.8-17.4 10.7
SijsiG G=Btsotele cooda45060000 43 100-205 150.6 5.6-35.1 21.1
Oct: TI=26 44. havactesctee oe oetae 40 125-201 168.5 9-9-47.7 28.0
of good growth are afforded in four specimens, 27.8—33.3 inches FL and weighing
4.8-8.8 pounds; the indicated annual increases were 6.3-14.4°/) in length, corre-
sponding to 28.8—47.0°/, in weight (for further details, see Vladykov, 78).
Sturgeon 11-34 inches long from the Hudson River were 2—8 winters old, as
indicated by their otoliths (35: 68, tab. 10). Otolith studies, though based on few
specimens, also indicate a greatly accelerated growth following the seaward migration of
the immature Sturgeon, if estimated ages of 11 years for a 75-inch fish and of 12 years
for two others of 88 and 100 inches are approximately correct. The maximum age, as
indicated by the winter marks on the otoliths, is at least 18 winters in the case of a
female 8 feet 8.5 inches and weighing about 225 pounds, taken from the Hudson
River, Malden, N.Y., April 30, 1937.42 Vladykov (78) has given a graph for the
length-weight relationship for a series of 1,592 Atlantic Sturgeon from the lower St.
Lawrence River, Quebec.
Spawning and Migrations. The Atlantic Sturgeon is an anadromous species, in-
variably spawning in fresh water but making its growth in salt water. Adults migrate
from the sea to fresh water in advance of the spawning season. The spawning migra-
tion begins during February in the St. Marys River, Georgia, typically during April
in Chesapeake Bay (39: 73-76), at the end of April and in May in the Hudson River,
and during May and June in the Gulf of Maine (13: 82). The exact movements of
mature Sturgeon in the St. Lawrence River have not been reported, but they prob-
ably begin to ascend the current in May (fishing during this month is prohibited)
and continue to run upstream through June and sometimes in early July.
In the Delaware River in 1925, the spawning migration started on April 23;
a few fish were taken during the last of April and the first part of May, with the
largest catches between May 12 and 22 inclusive (75: 184). During the last week of
May and the first ten days of June the catch was scarcely one fish per day, and on
June 10 the Sturgeon fishing ceased. The gill nets (13-in. stretched mesh) were set
in a narrow arm of the river between Pea-Patch Island and the New Jersey shore
opposite Delaware City, where the channel is about 2.5 miles long, 6—7 fms. deep,
and the current strong. The Sturgeon gathered for spawning in the upper part of the
river near the eastern shore, where the bottom is hard clay. Spawning took place
hard with their tails that the posterior ends of the caudal rays are broken, and only by slow regeneration do
they regain their original length. Thus some fish, after repeated recaptures within the same season, seem to have
grown somewhat shorter.
42. Age determination by Greeley, not previously published.
Fishes of the Western North Atlantic iat
at water temperatures ranging between 56°-64°F. Among 63 adult females, three
were “‘pasters’’; 1. e. with the roe in the form of a slimy, grayish white paste consisting
of squeezed eggs, white yolks, and ruptured membranes. Between the roth and 22nd
of May, seven females with running roe and three males with ripe milt were taken
during the act of spawning.
Little is known about the breeding grounds or spawning behavior along the
rest of the Atlantic coast. Apparently the sexually mature individuals ascend the St.
Lawrence River through deep channels that are kept free of nets because of navigation,
and it is probable that spawning takes place in pools below waterfalls of certain St.
Lawrence tributaries, for instance the Batiscan River along the southern shore.
Moreover, there are several rivers along the northern shore (Riviére-aux-Outardes, for
instance) where 4. oxyrhynchus spawn. It is probable that the spent fish return gradu-
ally to salt water.
The eggs when laid are light to dark brown. The outside membrane of ripe eggs
readily imbibes water and becomes glutinous and sticky. Being demersal and adhesive,
they become attached to weeds, stones, and so forth, and it is believed that the eggs
are scattered over a wide area. There is no evidence of prenatal care, such as prepara-
tion of a nest area (65: 267-268; 39: 73).
The diameter of ripe eggs is 2.5 or 2.6 mm; using this size as a basis for esti-
mates, there would then be 168,000 eggs per gallon (U.S.) and 800,000—2,400,000
eggs per fish (65: 268; 15: 186).
The largest ripe female examined by us, caught on June 23, 1954, in the St.
Lawrence River at Ile-aux-Patins, Quebec, was 8 feet 9 inches TL (8 ft. 2.5 in. FL),
weighed 352 pounds, and would have spawned in about a week; its ovary weighed g1
pounds and contained an estimated total of 3,755,745 eggs. No female 4. oxyrhynchus
have been reported as being ready to spawn before reaching at least 1 50 pounds and an
age of about 10 years. The testes of the smallest ripe male encountered during this
study, a fish of 69 inches TL (60 in. FL) and 70 pounds, taken in the St. Lawrence
River at St. André de Kamouraska on June 2, 1949, weighed 3 pounds 13 ounces.
General Habits. Sturgeon are bottom fish and are seldom seen except when taken
in nets or when jumping. It is of interest that this relatively sluggish species is capable
of making powerful jumps (p. 30). In the opinion of fishermen, at least in Quebec,
Sturgeon jump to rid themselves of attached lampreys.
Very little appears to be known about the behavior of the Sturgeon in salt water.
It is astonishing how easily these fish can adapt to a sudden change from salt to fresh
water, or vice versa. Some tagged specimens were forced to abruptly change habitats
at least twice during the same season, apparently without harmful results, because
they were recaptured again alive.
Thus far the actual movements of 4. oxyrhynchus have been studied by means of
tagging in Quebec only.*8 During five years, 1945-1949, a total of 1,948 was liberated
43- For details, including a photograph of a tagged Sturgeon, see Vladykov (80: 65-69; 76: 47). Recently in Oregon
several White Sturgeon (4. transmontanus) were marked with Peterson modified tags (58: 4-5).
4°
52 Memor Sears Foundation for Marine Research
at different localities in the St. Lawrence River, in both fresh and salt water. Up to
December 31, 1952, a total of 47 fish was recovered. Several of them were recaptured
as many as four and five times each and were subsequently reliberated. The majority of
the recaptures showed definite mass movements toward fresh water in spring (May—
June) and back to salt water in the fall (Sept.—Nov.). There were four recaptures of
tagged Atlantic Sturgeon of unusual interest: three liberated at Kamouraska and one
at Ile aux Coudres, Quebec. After periods varying between 307—705 days, three of
them were recaptured not far from Halifax, Nova Scotia, having traveled a minimum
of 900 miles. The fourth fish was retaken near the Strait of Canso. The weights of these
fish when recaptured, according to the fishermen, ranged between 6-24 pounds.
Food and Feeding. The large Sturgeon feeds on mollusks and other bottom organ-
isms. The fish roots in the sand or mud with its snout, like a pig (the barbels serving as
organs of touch), as it noses up the worms and mollusks on which it feeds and which
it sucks into its toothless mouth with considerable amounts of mud (65: 265). The
Sturgeon also eats small fishes, particularly launce (Ammodytes) (13: 83). The mature
Sturgeon, like the salmon, eats little or nothing while it travels up the river to spawn.
The digestive tracts of 26 young oxyrhynchus weighing 1-7 pounds, from the
Hudson River, contained bottom mud along with plant and animal matter, including
sludgeworms (Limnodrilus), chironomid larvae, isopods, amphipods, and small bivalve
mollusks (Pisidium) (72: 141-144, tab. 5). The food of 4. oxyrhynchus varies with the
type of habitat, as in the St. Lawrence River, Quebec. In 27 half-grown Sturgeon
taken in salt water, polychaete worms (Nereis virens) were found—265 on the average;
the maximum number in a single stomach was 1,221. In addition, the Sturgeon fed
on marine gastropods, shrimps (Crago), amphipods, and isopods, in that order. In fresh
water, the bulk of the food consisted of aquatic insects, amphipods, and oligochaete
worms; in 88°/, of 178 Sturgeon examined, larvae of the burrowing May fly (Hexagenia)
were present.*#
Abundance and Commercial Importance. The present Sturgeon fishery along the
western Atlantic coast is very small. In 1956, according to statistics (in pounds), the
following catches of 4. oxyrhynchus were made: Quebec 57,000, New Brunswick 800,
Nova Scotia 1,000, New England 9,000, Middle Atlantic, U.S., 15,000, Chesapeake
Bay 23,000, South Atlantic, U.S., 129,000, and Gulf of Mexico 15,000; the total was
249,800 pounds.
Former catches were manyfold greater: the catch in North Carolina in 1880
was 437,000 pounds (67: 56), in Chesapeake Bay in 1890 over 900,000 pounds
(39: 75), and in the Delaware River that year, 5,000,000 pounds (22: 372). Seven
years later, however, the Delaware catch was only 2,428,616 pounds, about half as
large as in 1880. By 1920 the catch had declined to 22,886 pounds in Chesapeake
Bay, by 1950 to about 1,200 pounds in and at the mouth of Delaware Bay, and to
18,900 pounds in Chesapeake Bay (see also Details of Occurrence, p. 54).*°
44. For further details on the food of 4. oxyrhynchus in Quebec, see Vladykov (77: 53-55).
45. For the abundance of Sturgeon in New England waters in Colonial days, see Bigelow and Schroeder (z3: 83).
Fishes of the Western North Atlantic 52
At the turn of the century, when Sturgeon were caught in very large numbers
and when mature individuals were the mainstay, fishing was so intensive that very
few fish were able to reach the upper waters to spawn. This was probably one of
the reasons for their great decrease in subsequent years. Also, the building of dams in
many important rivers (e. g. the Susquehanna, Maryland) deprived Atlantic Sturgeon
of their favorite spawning areas, and pollution by wastes of all kinds from the factories
and by municipal sewage from the towns and cities along the Atlantic shore aided
in reducing still further the local populations.
Laws prohibiting the capture of Atlantic Sturgeon below a minimum size or
during certain periods of fishing (39: 76) were helpful in protecting and maintaining
the species. However, even at its lowest ebb the Atlantic Sturgeon in North America
never was close to complete extermination, as some authors*® were inclined to believe
(in 1950), There are indications, along the Atlantic seaboard, that this Sturgeon is
coming back. The small number of large fish taken during recent years is partially
due to a reduction in the quantity of appropriate fishing gear. In New York, for
instance, the ro-inch mesh nets, legally specified for taking this species, are not used
in the Hudson River because fishermen do not consider it likely that enough fish
could be taken to justify the expense of gear and license (35: 139).
In Quebec, the Atlantic Sturgeon is perhaps as plentiful now as it used to be
in bygone years. For instance, at our request a fisherman of St. Vallier counted all the
young 4. oxyrhynchus that he liberated alive from his péche; during the night of Oc-
tober 2, 1944, he freed 1,000 young 6-15 inches long.
The principal Quebec catches are made with weirs situated along the southern
shore of the St. Lawrence River, from Riviére Ouelle to St. Nicolas. Some fish
are also taken regularly with gill nets in the same area. Along the northern shore of
the St. Lawrence, some large fish are caught in either weirs or in salmon nets, par-
ticularly in the estuaries of Riviére-aux-Outardes and Riviére Manicouagan, and
occasionally large fish are taken at Sept Iles.
Relation to Man. The Atlantic Sturgeon is a good quality foodfish, but less so
than the Lake Sturgeon (fu/vescens), which reaches maturity at around 20 pounds and
is more generally appreciated for “smoking” purposes. The flesh of the young or half-
grown Atlantic Sturgeon, which make up the great bulk of the catch of this species,
is Coarse, contains many ligaments, and has only a small amount of fat; it is only the
large adult that furnishes a tender and rich flesh. Some years ago the price paid to Quebec
fishermen for a fish weighing 30 pounds when dressed was 35 cents a pound for an
Atlantic Sturgeon as compared with 80 cents or even one dollar per pound for a Lake
Sturgeon of the same dressed weight; the male fish brings a better price than the female.
Moreover, the Atlantic Sturgeon carries heavier scutes, thus there is a greater loss of
weight during preparation of the meat. Dressing for the United States market consists
46. Darlet and Prioux (25: 6), in their review of the Atlantic Sturgeon supply in different countries, said that there
were no more left in America (“aux Ameriques il n’y en a plus”).
54 Memor Sears Foundation for Marine Research
of removing the head, collar bone (cleithrum), tail, all fins, and the viscera.4? Thus the
weight of prepared meat, which varies with size of fish, sex, stage of maturity, and
condition of fatness, is only about 50 °/, or even 40°/, of the entire Sturgeon. The roe
from Atlantic Sturgeon is as good for making caviar as that from Lake Sturgeon, and
being mature at a larger size, the Atlantic Sturgeon furnishes a larger quantity of eggs.
In early Colonial days, around 1709 in North Carolina, the rough Atlantic Sturgeon
scutes were considered “‘good nutmeg graters’’ (67: 56).
The value of the Atlantic Sturgeon for angling purposes is negligible. Neverthe-
less, occasional captures of large ones with hook-and-line stir public interest. It is worth-
while to mention the capture of a 6-foot Sturgeon, foul-hooked, by a skilled woman
angler at Wasque Point, Martha’s Vineyard, Nantucket Sound (z3: 84).
General Range. The regular range of 4. oxyrhynchus is limited to the Atlantic
coast of North America and the Gulf of Mexico. The northern subspecies, 7. oxy-
rhynchus oxyrhynchus, occurs from Hamilton Inlet on the Atlantic coast of Labrador
(7: 290) and the Gulf of St. Lawrence to eastern Florida. In the Gulf of Mexico is
found the southern subspecies 4. oxyrhynchus desotoi (p. 56) (34: 408, 409, 411). Oc-
casional captures of Atlantic Sturgeon have been reported for Bermuda‘ and even for
French Guiana,** but these southern stragglers probably are of the subspecies desotoi.
And a large Sturgeon, taken near the mouth of the Mississippi River, the photograph
of which has been published (34), can be identified as desotoi by its broad and strongly
developed scutes.
There are early reports of the presence of 4. sturio (equals 4. oxyrhynchus) in
Hudson Bay, but from the descriptions it is evident that the specimens in question
were 4. fulvescens (see also 75: 30 [42]).
Details of Occurrence. Hamilton Inlet on the Atlantic coast of Labrador is the
most northerly point where 4. oxyrhynchus has been reported (7: 290). Blanc Sablon,
on the Quebec side of the Strait of Belle Isle, is the next most northerly point (some
specimens from this locality are in our collections). Atlantic Sturgeon are found reg-
ularly throughout the Gulf of St. Lawrence and in the St. Lawrence River up to Three
Rivers, and odd specimens are taken even in Lake St. Peter, near Sorel, Quebec (23:
19). They are found also in small numbers on the Newfoundland side of the Gulf of
St. Lawrence, are well known in Nova Scotian waters, especially near estuaries (85:
58), and are caught regularly in the St. John River, New Brunswick, as well as at the
head of the Bay of Fundy (42: [10] 58).
To the south, Atlantic Sturgeon are (or were) well known in the Penobscot,
Kennebec, and Merrimack rivers; indeed, they entered nearly every stream of any size
47. Ryder (65: 276) and Hildebrand and Schroeder (39: 75) have given details on the methods of preparing Stur-
geon for the market.
48. According to Beebe and Tee-Van, two records of 4. oxyrhynchus were mentioned in early Bermuda lists, one
for 1876 and the other for 1887 (9: 32).
49. In 1867 Dumeéril (27: 161) described a new Sturgeon, as 4. cayennensis, from French Guiana “‘de la riviere
Oyapock dite la Cayenne.” Bertin (12: 248-249), after examining the holotype (No. 3493, 740 mm, stuffed,
in good condition) in the Paris Museum, considered it as a possible hybrid 4. sturio x A. brevirostris, but Vladykov,
who re-examined it in 1950, identified it as 4. oxyrhynchus.
Fishes of the Western North Atlantic 55
that empties into the Gulf of Maine. There are also definite records of Sturgeon taken
off the open coast from the Bay of Fundy southward to Cape Cod (r3: 83), and along
the coasts of southern New York. Along the Middle and South Atlantic seaboard
of the United States there are several rivers that formerly maintained important
fisheries: the Hudson, Delaware, Susquehanna, Potomac, York, James, St. Marys
(Georgia), and St. Johns (Florida).*°
At Quebec, it seems that young 4. oxyrhynchus pass their entire growth in the
fresh and brackish water of the St. Lawrence River, for Sturgeons of all sizes are taken
there—from a few inches long up to nine feet or longer (78: 53-56). Why it was that
some half-grown tagged fish were recaptured several hundred miles from the place of
release is not known (pp. 51, 52). But the small number of these stragglers does not ap-
preciably change the impression that the majority of the Atlantic Sturgeon produced in
the St. Lawrence River does not venture far from the estuary, for they are taken there
in some numbers throughout the entire fishing season.
In the Hudson River, where gill nets used for shad and other species accidently
capture Sturgeon 11-34 inches long and 2-8 winters old (age determinations based
on otoliths), there is abundant evidence from both winter and summer catches that
these immature Sturgeon inhabit the freshwater areas of the river throughout the
year. But there is a wide gap in the Hudson River collections between these and the
adults; since immature Sturgeon are frequently encountered in salt water in the
general vicinity, it seems likely that all of them go to sea sooner or later to complete
their growth. This rule seems to apply also to the rivers of the Gulf of Maine. However,
some of the Delaware River Sturgeon seeking salt water may remain within the lower
reaches of Delaware Bay while others go to sea, as proven by catch records of fish
taken in the pound nets near Ocean City, Maryland. In Chesapeake Bay, fish of all
sizes are taken in both the upper and lower parts of the Bay throughout that portion
of the year when the pound nets are in operation (39: 73). But it is not known how
many Sturgeon stray beyond the outer headlands, from Chesapeake Bay, the sounds
of North Carolina, or the rivers farther southward.
Published reports of Sturgeon taken at sea are mostly for waters close to shore,
but landings of 3,700 pounds in 1950 and 3,900 pounds in 1952 in New York
and New Jersey by otter trawlers show that some wander out on the Continental
Shelf; and to the east, some even stray to the offshore fishing banks. Thirty
Sturgeon, for example, were landed in Boston and Portland from Brown Bank,
Georges Bank, the South Channel, and Nantucket Shoals by otter trawlers during
the years 1927-1934; and landings of 5,300 pounds in 1940 and 6,600 pounds
in 1947 in New England ports from these offshore grounds correspond to about
50 and 70 fish, respectively, if an average weight no greater than 100 pounds is
assumed.*1
50. We believe that the specimens from South Carolina and Florida, referred to as 4. brevirostris by Jordan and Ever-
mann (43: 106) and by several subsequent authors, were oxyrhynchus.
51. From data collected by the late Walter H. Rich, U.S. Bureau of Fisheries, and from notices in the daily press.
56 Memoir Sears Foundation for Marine Research
Synonyms and American References:
Acipenser oxyrhynchus Mitchill, Trans. Lit. philos. Soc. N.Y., 1, 1814: 462 (descr., New York); DeKay, Zool.
N.Y., 4, 1842: 346 (descr.); Provancher, Nat. Canad., 8, 1876: 226 (genl. acct.); Smith, N.C. geol.
econ. Surv., 2, 1907: 57 (Albemarle Sd.); Hildebrand and Schroeder, Bull. U.S. Bur. Fish., 43, 1928:
72 (Chesapeake Bay); Breder, Field Bk. Mar. Fish. Atl. Cst., 1929: 42 (genl. acct.); Jordan, Manual
Vert. Anim. NE U.S., 1929: 32 (distr., key, brief descr.); Jordan, Evermann, and Clark, Rep. U.S.
Comm. Fish. (1928), 2, 1930: 34 (in checklist); Gowanloch, Fish. Fishing Louisiana, Bull. La. Conserv.
Dept., 23, 1933: 411 (mouth of Mississippi R.); Schrenkeisen, Field Bk. Freshw. Fish. N. Amer.,
1938: 12 (genl. acct.); Vladykoy and Beaulieu, Nat. Canad., 73, 1946: 43 (detail. acct. of shields and gill
rakers, size, Quebec); Vladykov and Beaulieu, Nat. Canad., 78, 1951: 129 (detail. acct. of gill rakers
for three w. Atlantic species); Rostlund, Univ. Calif. Publ. Geogr., 9, 1952: 248 (import. to native In-
dians).
Acipenser sturio Linnaeus,°* Syst. Nat., ed. 10, 1758, 237 (descr., European seas); Ryder, Bull. U.S. Fish
Comm. (1888), 8, 1890: 232 (descr., Atlant. cst.); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1),
1896: 105 (descr., Atlant. cst.); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 16 (New England);
Kendall, Proc. Portland Soc. nat. Hist., 3 (1), 1914: 13 (Maine); Huntsman, Contr. Canad. Biol. (1921),
3, 1922: 10 (58) (St. John R., head of Bay of Fundy); Bigelow and Welsh, Bull. U.S. Bur. Fish., 40
(1), 1925: 74 (descr., habits, Gulf of Maine); Borodin, Trans. Amer. Fish. Soc., 55, 1925: 184 (biol.
observ.); Nichols and Breder, Zoologica, N.Y., 9, 1927: 30 (size, distr. New England, New York);
Truitt, Bean, and Fowler, Bull. Md. Conserv. Dep., 3, 1929: 33 (Chesapeake Bay); Beebe and Tee-
Van, Field Bk. Shore Fish. Bermuda, 1933: 32 (2 recs. for Bermuda cited); Vladykov and McKen-
zie, Proc. N.S. Inst. Sci., 79 (1), 1935: 53 (genl.acct.); Jordan and Evermann, Amer. Food Game
Fish., 1937: 8 (genl. acct., key); Bigelow and Schroeder, Fish. Bull. (74) U.S. Fish Wildl. Serv., 53,
1953: 81 (descr., habits, Gulf of Maine).
Acipenser sturio oxyrhynchus, Smith, Bull. U.S. Fish Comm. (1891), 1893: 190 (N. Carolina rivers: Pasquo-
tank, Edenton, and Roanoke); Halkett, Check List Fish. Canad., 1913: 44 (distr.); Greeley, N.Y. Biol.
Surv. Lower Hudson Watershed (1936), 11, 1937: 89 (age determ., size ranges, Hudson R.); Bailey,
Biol. Surv. Merrimack Watershed, New Hampshire Fish Game Dept., 1938: 155 (former abund.
Merrimack R.).
Acipenser cayennensis, Duméril, Nouv. Arch. Mus. Hist. nat. Paris, 3, 1867: 161 (descr., River Oyapock
known as Cayenne, French Guiana; see ftn. 49).
To the above Synonyms should be added several nominal specific names, which were given by Duméril (27:
161-177; 28: 116-228) to specimens of 4. oxyrhynchus according to locality only: mitchil/i (New
York), kennicotti (James R.), girardi (Maryland), macrohinus (New York), milberti (New York), bairdi
(Maryland), storeri (Boston), Ao/brooki (Charleston), and /econtei (New York).
Acipenser oxyrhynchus desotoi Viadykov 1955
Gulf of Mexico Sturgeon, Common Sturgeon
Figure 3
Study Material. Two specimens, 50 and 60cm FL, obtained by T. Dawson be-
tween Twin and Rabbit islands at the mouth of Singing River, off Gautier, Missis-
sippi, November 30, 1953, CNHM 59803, 59804);*! 3 specimens (seen by V.D.V.),
from western coast of Florida (two from Cedar Keys, the other from Suwannee River),
UF collections.
525
53"
54-
Among the references, we have omitted some early authors whose incomplete or erroneous descriptions of the
species are of little scientific value.
European authors in general, such as d’Ancona (2), Berg (rr: 94), and others, consider the American form identi-
cal to the European 4. sturio.
Kindness of Loren P. Woods. These specimens served as the basis for the original description (81: 2, pls. 1, 3, 5, 7):
Fishes of the Western North Atlantic Gel
Distinctive Characters and Description. The subspecies desotoi is distinguishable
from typical oxyrhynchus of comparable size by the following differences, the most
striking and characteristic difference being the length of the spleen. Head, pectoral,
and spleen given in per cent of fork length. For other details, see Vladykov (81).
Character desotot oxyrhynchus
Scutes — dorsal row... squarish, with length much _ oval, with length greater than
shorter than width; carina width;* carina (keel) low,
(keel) typically with two without a pronounced hook
strong hooks
Pleads ec met aes 30.9-33-6°/, 26.5—-27.6°/,
Pectoral! fins) .9.<5.-\! 15.5-16.3°/, Ties 0%/,
Spleenmencthyes i ya2 16-19°/, 3-9°/o
* Specimens from eastern Florida and Quebec. { Two specimens.
Habits. The subspecies desotoi is anadromous, as is the northern subspecies oxy-
rhynchus. Further than this there is no detailed information on its habits, except that
Louisiana fishermen occasionally capture large Sturgeon around the mouth of the Mis-
sissippi River and in the Sound, where they have been reported as ““Common Stur-
geon” (34: 411-412). Adults probably visit several rivers on the northern and eastern
shore of the Gulf of Mexico for spawning purposes.
Range. Apparently its occurrence is limited to the Gulf of Mexico, the northern
coast of South America, and Bermuda (p. 54).
Synonyms and References:
Acipenser oxyrhynchus de sotoi Vladykov, J. Fish. Res. Bd. Canada, 12 (5), 1955: 2 (orig. descr.).
Acipenser oxyrhynchus Gowanloch, Bull. La. Conserv. Dep., 1933: 411 (mouth of Mississippi R.).
Acipenser sturio Rivas, Bull. U.S. Fish Wildl. Serv., 55 (89), 1954: 504 (n. and e. shore, Gulf of Mexico).
Acipenser (oxyrhynchus) de sotoi Springer and Bullis, Spec. Sci. Rep., U.S. Bur. Comm. Fish., 196, 1956: 46
(Mississippi Sd.).
Relationship of 4. oxyrhynchus to A. sturio
Some authors (see Synonyms and References, p. 56), following Ryder (65: 234-
238), have considered the Atlantic Sturgeon of America to be the same species as
the Common Sturgeon of Europe,
A. sturio Linnaeus; the two species Table V. Comparative Counts of 4. sturio and
resemble each other very closely A. oxyrhynchus.
in certain features, as is shown in Species ers - 210-7. A. sturio A. oxyrhynchus
Table v. Authority ...... Berg Vladykov and Beaulieu
D’Ancona, in a summary of Elric: (II: 93°95) (83: 4457)
the characters of 4. sturio, presum- eee sue te
ably from the Mediterranean Sea, Gill rakers ..... 18-25 17-27
58 Memoir Sears Foundation for Marine Research
has quoted somewhat higher figures than those given in Table v for the shields: to-14
in the dorsal row (excluding the nuchale) and 25-36 in the lateral row (2). Antoniu,
for A. sturio from the Black Sea, mentioned the presence of pairs of postdorsal shields
and of a frontal sinus in the cartilaginous skull; both of these characters are found
in A. oxyrhynchus (4: 109-116).
Of two specimens obtained by Cope nearly a century ago in Adige (Verona), Italy,
and labeled 4. sturio, one is probably 4. gii/denstadti (ANSP 646) and the other is
definitely an 4. sturio (ANSP 645), 15 inches long (380 mm) FL. The latter has a
fontanelle, 10 shields (including nuchale) in the dorsal row, and 30 shields (including
supracleithrum) in the lateral rows. On the first branchial arch we counted 20 gill
rakers. The body proportions are also very similar to those of 4. oxyrhynchus of cor-
responding size, and its color, or what remains of it, corresponds very closely to that
of American specimens.
In a drawing of 4. sturio by d’Ancona, the coloration is also very similar to
that of the western Atlantic Sturgeon (2: fig. 1); on the other hand, the mouth of
d’Ancona’s 4. sturio is larger than that of 4. oxyrhynchus (2: fig. 2), the shape of its
lower lip is very different, and there are other differences as well. According to Berg,
the adult of 4. sturio has several (up to 10-12) dense, oblique series of rhombic plates
(dermal ossifications) between the dorsal and lateral rows of shields (rr: 97); in western
Atlantic specimens these ossifications are much less developed and are rather irregular
in shape. Moreover, Classen (20: 88-102) has mentioned several ripe females of
sturio from the Gulf of Cadiz, Spain, 63 inches TL (145 cm FL) and weighing as
little as 62 pounds (28 kg); oxyrhynchus apparently attains sexual maturity at a larger
size than sturio, the males of oxyrhynchus being approximately six feet TL and 70
pounds in weight, the females not less than 150 pounds (p. 51).
In Europe, probably because of a general scarcity of 4. sturio (25: 5-13), little
work has been done to elucidate the problem of local races. Nevertheless, there are
observations by Marti that 4. stwrio from the Black Sea are similar to those from the
Mediterranean region but differ from those of the Baltic (52: 435-442, Russ.);
the Black Sea specimens have on the average higher numbers of dorsal and lateral
scutes and dorsal rays. Magnin, by studying cross sections of the pectoral ray, found
that the rate of growth of the European Sturgeon varies according to geographical
region (49: 152-159). The growth observed was fastest in Italy (Tiber and Po rivers),
intermediate in Spain (Guadalquivir River), and slowest in France (Gironde River).
In conclusion, until an adequate comparison of anatomical and meristic characters
of these two species is made, it is preferable to retain separate specific rank for 4.
sturio and oxyrhynchus.
Ont
Io.
Tek
72s
73!
T4.
TS
16.
sigh
Is.
19.
20.
Dit.
EXD AND IEOOTRNOT EsREFERENCES
American Fisheries Society, List of Common
and Scientific Names... Fish...U.S. and
Canada, Publ. 1, 1948.
d’Ancona iz Faune Ichthyol. d’Atl. nord, 2,
1929.
Antipa, Fauna Ichthyol. Romaniei (Buch.),
1909.
Antoniu, Ann. Sci. Univ. Jassy, 23 (2), 1937.
Antoniu, Ann. Sci. Univ. Jassy, 28 (2), 1942.
Antoniu, Bull. Acad. Roumaine, 29 (5), 1946.
aexus! Copeia, 1951.
Bailey, Pap. Mich. Acad. Sci., 39, 1954.
Beebe and Tee-Van, Field Bk. Shore Fish.
Bermuda, 1933.
Berg, Class. Fish., Trav. Inst. zool. Acad. Sci.
URSS, 5 (2), 1940; reprinted by J. W. Ed-
wards, Ann Arbor, Mich., 1947.
Berg, Poiss. des eaux douces... U.R.S.S., 7,
1948.
Bertin, Bull. Mus. Hist. nat. Paris, (2) r2 (6),
1940.
Bigelow and Schroeder, Fish. Bull. (74) U.S.
Fish Wild. Serv., 53, 1953-
Borisov, Trav. Comm. Rép. Jakoute, 9, 1928.
Borodin, Trans. Amer. Fish. Soc., 55, 1925.
Cxcm Trans. Amer. Fish. Soc. (1904),
1905.
Chalikov, Zool. J., 23, 1944.
Cheng, Fukien Univ. Sci. J., r, 1938.
Classen, Tech. Fish. Prod., 1925 (Russ.).
Classen, Trab. Inst. esp. Oceanogr., 19, 1944.
Clemens and Wilby, Bull. Fish. Res. Bd.
Canad., 48, 1946.
59
22.
23.
37:
38.
Cobb, Rep. U.S. Comm. Fish. (1899), 1g00.
Cuerrier, Fry, and Prefontaine, Nat. canad., 73,
1946.
Cuerrier and Roussow, Canad. Fish Cult., ro,
Ig5I.
Danzer and Prioux, Bull. franc. Piscic., 758,
1950.
Dominion Bureau of Statistics, 1956.
Duméril, Nouv. Arch. Mus. Hist. nat., Paris,
3, 1867.
Duméril, Hist. Nat. Poiss., 2, 1870.
Evermann and Latimer, Proc. U.S. nat. Mus.,
39, I9TI.
Forges and Richardson, Bull. Ill. Lab. nat.
Hist., 7, 1905.
Forbes and Richardson, Bull. Ill. nat. Hist.
Surv., ed. 2, 3 (3), 1920.
Fortin, Ann. Rep. Fish. (1863), Append., 1864.
Fowler, Proc. Acad. nat. Sci. Philad., rgro.
Gowantocn: Bull. La. Conserv. Dep., 23,
1933-
Greeley, 26th Rep. N.Y. St. Conserv. Dep.,
Suppl., 1937.
Grote, Vogt, and Hofer, Siisswasserfische von
Mittel-Europa, 1 (text), 1 (Atlas), 1909,
1908.
Hazkert, Check List Fish. Canad., 1913.
Harkness, Publ. Ont. Fish. Res. Lab., Biol.,
24 (18), 1923.
39:
40.
Memoir Sears Foundation for Marine Research
Hildebrand and Schroeder, Bull. U.S. Bur.
Fish., 43, 1928.
Hinks, Fish. Manitoba, 1943.
Hubbs and Lagler, Bull. Cranbrook Inst. Sci.,
26, 1947.
Huntsman, Contr. Canad. Biol. (1921), 3,
1922.
: (Pars and Evermann, Bull. U.S. nat. Mus.,
47 (1), 1896.
Jordan and Evermann, Amer. Food Game
Fish., 2nd ed., 1937.
Jordan, Evermann, and Clark, Rep. U.S.Comm.
Fish. (1928), 2, 1930.
: Loarimore, Copeia, 2, 1950.
Leim and Day, J. Fish. Res. Bd. Canad., 16
(4), 1959-
Mien: Rev. Trav. Inst. Péches marit., 23,
1959:
Magnin, Bull. franc. Piscic., 193, 1959.
Magnin and Beaulieu, Nat. canad., 87, 1960.
Marti, Zool. J., 78, 1939.
Montpetit, Poiss. d’eau douce Canad., 1897.
Mori, J. Chosen nat. Hist. Soc., 16, 1933.
Mori, Mem. Hyogo Univ. Agr., I (3), 1952.
: Nicuots, Nat. Hist. Centr. Asia, 9, 1943.
Nikitin, Fish Products, 1949 (Russ.).
Nikolski, Rept. Amph., Fish., 1906 (Russ.).
Onzcon State Game Commission, Bull., 1950.
Provancuer, Nat. canad., 8, 1876.
Putchkoy, Phys. Fish., 1941 (Russ.).
4 Rosrzunp, Univ. Calif. Publ. Geogr., No. 9,
1952.
62.
63.
56.
Roule, Poiss. des eaux douces Fr., 1925.
Roussow, Off. Biol. Min., Chasse Péches,
Quebec, 1955.
Roussow, J. Fish. Res. Bd. Canada, r4, 1957.
Ryder, Bull. U.S. Fish Comm. (1888), 8, 1890.
Scorr, Freshw. Fish. east. Canad., 1954.
Smith, N.C. geol. econ. Surv., 2, 1907.
Stannius, Handb. Zool. Fische, ed. 2, 1854.
Stone, Trans. Amer. Fish. Soc., 29, 1900.
Tararxo, Trav. Mus. zool. Acad. Sci. Ukr.,
IO, 1936.
Ting, Copeia, 1949.
Townes, 26th Rep. N.Y. St. Conserv. Dep.,
Suppl., 1937.
Trusov, Doklady Acad. Sci. USSR, 67 (3),
Oa:
Us. Fish Wildl. Serv., Statist. Digest., 1956
Viapyxov, Contr. Canad. Biol., 8 (29), 1933.
Vladykov, Rapp. Gén. Dép. Péch. Quebec
(1945-1946), 1946.
Vladykov, Rapp. Gén. Dép. Péch. Quebec
(1947-1948), 1948.
Vladykov, Rapp. Gén. Dép. Péch. Quebec
(1948-1949), 1950.
Vladykov, Rapp. Gén. Dép. Péch. Quebec
(1950-1951), 1951.
Vladykov, Rapp. Gén. Dép. Péch. Quebec
(1952-1953), 1953-
Vladykov, J. Fish. Res. Bd. Can., 12 (5), 1955.
Vladykov, Fish. Quebec, Alb. 5, 1955.
Vladykov and Beaulieu, Nat. canad., 73,
1946.
Vladykov and Beaulieu, Nat. canad., 78,
1951.
Vladykov and McKenzie, Proc. N.S. Inst. Sci.,
19 (1), 1935-
Wisconsin Conserv. Bull., August 1954.
Order Lepisostei
ROYAL DS UrykuUSs
Tulane University
Acknowledgments. My thanks are due the following persons for the loan of specimens
or for help in making specimens available for study: Leonard P. Schultz, Ernest A.
Lachner, William Ralph Taylor, Robert Kanazawa, and “fames Tyler of the United
States National Museum; ‘fames E. Bohlke of the Academy of Natural Sciences at
Philadelphia; Henry B. Bigelow and Mrs. Myvanwy Dick of the Museum of
Comparative Zoology at Harvard University; Reeve M. Bailey, Robert R. Miller,
and Carter Gilbert of the Museum of Zoology, University of Michigan; Clark
Hubbs of the University of Texas; Charles M. Breder and Vladimir Walters of
the American Museum of Natural History ; Luis Rivas of the University of Miami;
William M. Clay and Louis A. Krumholz of the University of Louisville ; Loren P.
Woods, Robert F. Inger, and Pearl Sonoda of the Chicago Natural History Museum,
and Shelby D. Gerking of the University of Indiana. Thanks are also due Edward
C. Raney and Henry B. Bigelow for reading the manuscript and for making helpful
suggestions for improvement ; to Bill Komodore for delineation of Figs. II, 14, 15, 16;
to Rudolph *f. Miller for Figs. 12,13, 18; and to Miss Patricia Hale for Fig. 17.
Preparation of the illustrations and examination of many specimens at various
museums and institutions were made possible by financial aid from the Sears Founda-
tion through a grant-in-aid from the National Science Foundation (N.S. F. No.
G 7123), and from a National Science Foundation grant to the author (N.S. F.
No. G 9026).
Scope of Study. This account gives detailed descriptions of the Order Lepisostei
and family Lepisosteidae, of the genus Lepésosteus, and of four species—ocu/atus,
osseus, platyrhincus, and spatula. The ‘Key to Species of the Western North Atlantic”
encompasses subgenera as well as species and gives the alternate characters that are
61
62 Memoir Sears Foundation for Marine Research
readily recognizable. The Study Material 1s listed for each species; in most cases this
includes specimens from the many institutions listed. For easy and quick identification,
the Distinctive Characters, as in the Key, compare the easily recognizable features of
one species with the others. Many specimens were examined in preparing the taxonomic
accounts, and the methods of measuring and counting the characters are detailed on
p. 68.
The Descriptions are broken down into three major categories: (1) Proportional
measurements in per cent of length, (2) Proportional measurements in per cent of head
length, and (3) Meristic counts. Thus measurements for most of the head characters
(snout, lower jaw, etc.) are given in per cent of head length as well as in per cent of
length of fish. Following submission of this account for publication, a small specimen
of spatula, 49.5 mm (57.3mm TL), came to hand; the measurements for this individual
have been incorporated in the Description with figures enclosed in brackets.
What is known of the life history of Gars in general and of the species in particular
has been culled from the literature and gained by direct observation and personal com-
munication with field workers. The Synonyms and References at the end of each species
account include references to only those taken in salt water; these are not numerous,
because Gars for the most part frequent fresh water, and less commonly brackish or
salt water.
Characters. Lepisostei are slender ray-finned fishes with well-ossified skeletons,
as in the teleosteans, but with ganoid scales (see below), an archaic character shared by
the polypteroids alone among living fishes. The arterial cone of the heart has eight trans-
verse rows of 4—8 valves each,! a number of rows greater than in any other living ray-
finned fish; this character also puts the Gars apart from the polypteroids. The caudal
fin has no prolonged fleshy axis, but all of the hypural bones that bear the fin rays are
supported by the upturned end of the vertebral column (caudal “abbreviate hetero-
cercal,”’ Fig. 11), and the rear boundary of the fleshy caudal peduncle slopes obliquely
dorsoposteriorly, the latter character shared by Amia alone among living fishes.
The vertebrae are completely ossified and opisthocoelus, i.e. posterior face con-
cave and anterior face convex, a conformation unique among living fishes though paral-
leled in some tailed Amphibia and in the thoracic region of penguins, gulls, and plovers
among birds. In the adult the pleural ribs extend from the vertebrae to the skin (Balfour
and Parker, 7: 387, pl. 28 fig. 72; Emelianov, 20: 176-180, figs. 8-10).
The elongated snout, with nasal openings and olfactory sacs at its end, is the result
of a lengthening of the ethmoid region, and olfactory nerves course through long canals
in the ethmoid cartilage (Regan, 58: 447, fig. 3). The preorbital (lacrimal or maxillary
of some authors) is subdivided into a row of 6-8 bones that bear small, medium,
and large teeth (Holmgren and Stensié, ro: 474, fig. 363; Hammarberg, 29: figs. 41,
43; Berg, 8: 211, 414); the larger teeth are radially grooved at the base. The
infraorbital sensory canal is a prominent surface feature of the preorbitals in the early
1. For an excellent illustration and for a table giving the numbers of valves in various other fishes, see Bertin
(25: 1402, fig. 1005B, 1404).
Fishes of the Western North Atlantic 63
developmental stages (Collinge, 75: 265, pl. 4, fig.2; 24: 511, 512; Landacre and
Conger, 42: 593, 594; Hammarberg, 29: 309, figs. 39-43). A small sliver-shaped
bone is present, at least in the young, at the angle of the mouth; this bone is continuous
with the preorbitals in its dentition but is not traversed by the infraorbital canal; thus
it is considered to represent the maxillary. The vomer is paired. There is no opisthotic
See
Ficure 11. Lepisosteus spatula. Caudal fin skeleton; original drawing (natural size) of a specimen from Lake
Pontchartrain, Louisiana.
(Mayhew, 47: 327), supraoccipital, gular plate, or myodone in the Garfishes. The lower
jaw, a complex of six dermal bones—dentary, articular, angular, preangular, prearticu-
lar, and coronoid (Arambourg and Bertin, 25: 2185, fig. 1546)—articulates in front
of the eye. The sacculus and lagena form a common sac, the largest otolith being in
the former. Three pairs of branchiostegals support the gill membrane. The opercular
gills are especially well developed in the Garfishes, an upper horizontal “arm” and a
bisegmented lower one being noticeable (Wright, 76: 483).
The dorsal fin is situated posteriorly, above the anal fin. The fins are without
spines, and the rays of the dorsal and anal fins are equal in number to the supporting
radialia. There is no adipose fin.
The body of the adult is completely encased in an armor of rhombic ganoid scales,
which may be denticulated and sculptured on the exposed surface.? Most of the scales
2. The degree of sculpturing is of some taxonomic value when specimens of comparable size are used.
64 Memotr Sears Foundation for Marine Research
are hinged by a peg- and groove-type articulation. Although the bony scales form nearly
an impenetrable covering, the articulations, the fibrous connections, and the arrangement
in diagonal rows allow some flexibility of the body. The lepisosteid ganoid scale is com-
posed of two layers, ganoine above and isopedine below, both of which are irregularly
penetrated by vascular canals.3 In addition, both layers, especially the latter, are per-
forated by a system of small tubes; these tubules penetrate the scale from all sides and
are dendritic at their inner terminations. A similar layer of ganoine is present on the
dermal bones about the head. Both margins of the caudal fin as well as the leading
margins of the other median and paired fins are fortified with a biserial row of fulcra.
The cheeks are covered by numerous irregular plates.
The swim bladder,‘ cellular and highly vascularized, is connected to the pharynx
by the enlarged pneumatic duct (Wiedersheim, 70: 1-16, pls. 1-3). The gonads are
staggered (Miiller, 49: pl. 6, fig. 2), 1.e. the right one is mostly anterior to the pelvic
fins and the left one mostly posterior to them. The ovaries are closed, and the right
and left oviducts join the respective urinary ducts. A short distance posterior to their
junction, a common chamber is formed by the union of right and left parts; thus
the products from both systems exit from the body through a single urinogenital
sinus. No vestigial Miillerian ducts exist in the male (Pfeiffer, 52: 465). The conus
arteriosus is a contractile vessel with several rows of valves (49: pl. 5, fig. 2).
Remnants of the spiral valve occur in the posterior part of the gut. There is no lumi-
nescent organ.
Taxonomic Rank. In 1844, Johannes Miiller combined his family Lepidosteini
(evidently to include the amiids) with his family Polypterini to constitute his Order
Holostei (49: 201-204), and they have been similarly ranked as one of the constituent
divisions of a more widely inclusive Order by some subsequent authors. Thus they were
associated: with the Amiidae and Polypteridae by Ginther (28: 328) and with the Amii-
dae (among living fishes) by Bridge (12: 495, 502) as the Order Holostei; with the
Amiidae (among living fishes) by Arambourg and Bertin (25: 2181-2194) in 1958 as
the Order Amiiformes.
They have been regarded, however, as the sole living representatives of a separate
Order by the majority of recent authors: Ginglymodi by Cope (16: 452, 453), Rhom-
boganoidea by Jordan and Evermann (37: 108), Lepidosteoidei by Goodrich (23: 340),
Holostei by Jordan (36: 115), Ginglymodi by Regan (59: 313), Holostei by Jordan
et al. (38: 36), Lepidosteiformes by Berg (8: 211, 414), Semionotoidea by Romer
(62: 580), Lepisosteida by Matsubara (46: 170), and Ginglymodi by Norman (51:
61). Here also they are regarded as representing a separate Order.
Name of the Order. The choice here lies between Ginglymodi and some derivative
3. For accounts of the scales, see especially L. Agassiz (2: 74, 75, 77, tab. G, figs. 8-10), Williamson (73: 435-447,
470, pl. 40, figs. 1, 2; 74: 651, 658, 687, 699), Reissner (6r: 254-268, pl. 5, figs. 1-6), Nickerson (50: 115-139,
4 pls., 31 figs.), Scupin (63: 166, 167, pl. 10, fig. 1), Goodrich (23, 758, 759, fig. 199; 43: 218, 219, fig. 192),
Kerr (40: 63-66, figs. 3, 4).
4. Apparently the swim bladder functions as a breathing organ (Potter, 54: 63) in addition to the gills. When
removed from water, the Garfishes regularly breathe air into and out of the swim bladder.
Fishes of the Western North Atlantic 65
of Lepisosteus.’ Lepisostei is chosen here because of the connotation carried by this
name.
Relationships. It is customary in general works on living fishes to place the Lepi-
sostei next to the amiids, with which they agree: in the abbreviate-heterocercal nature
of their caudal fin skeleton (made evident externally by the dorsoposterior slope of the
rear boundary of the fleshy caudal peduncle); in their flattened, overlapping pelvic
bones; in their chambered swim bladder that serves as an accessory respiratory organ;
and in the presence of a rudimentary spiral valve in their intestine. However, they dif-
fer widely from the amiids in many respects. The body scales, for example, which are
thick and interlocking but not overlapping in the lepisosteids, are thin and widely over-
lapping in the amiids, but the fulcral scales that arm the margins of the fins in the
former are rudimentary in the latter. The long snout, the segmented “maxillary” bones,
a lower jaw articulating anterior to the eye, and the three rod-like branchiostegals of
the lepisosteids are replaced in the amiids by a short snout, unsegmented maxillaries,
a lower jaw articulating posterior to the eyes, and by 10-12 broad plate-like branchio-
stegals; and the free margin of the branchiostegal membrane, which crosses the isthmus
in a continuous arc of long radius in the lepisosteids, is deeply incised there in Amia,
one side overlapping the other. In lepisosteids the preopercular bone does not extend
as far forward as that in the amiids, the cheek region is covered by irregular plates
in lepisosteids but not in amiids, there is no bony gular plate in the chin region be-
tween the branches of the lower jaw as in Amia, and the dorsal is much shorter in
lepisosteids than in amiids. The following internal differences deserve mention: the
vertebral centra, which are concave posteriorly but convex anteriorly in the lepisosteids,
are concave anteriorly as well as posteriorly in 4mia; the vertebral column in the tail
region is diplospondylous; and the arterial cone of the heart, which has eight trans-
verse rows of valves in the lepisosteids, has only two rows in the amiids.
Among fossil groups, lepisosteids are usually associated with the Semionotidae,
some of which are known from as far back as the Jurassic.
Spawning and Development. Spawning takes place entirely in fresh water during a
rather brief period beginning about mid-May and lasting until mid-June, in the latitude
of New York (z [1879]: 65, 67). When Garfish move into shallows to spawn, each
female is accompanied by one to four males and there is a great amount of thrashing
during the breeding activities. Fertilization is external. The large numbers of individ-
uals that concentrate in the shoal areas suitable for spawning disperse rather suddenly
to other areas afterwards.
The eggs are adhesive and adhere to the substratum. No parental care is given to
the eggs or young. On the ventral surface at the end of the snout the larvae have peculiar
adhesive suckers that are used for attachment to objects on or above the bottom. The
larvae are relatively inactive until the yolk mass is absorbed, but then they become very
active predators.
A characteristic caudal filament that develops at the termination of the upturned
5. Many authors have preferred the spelling Lepidosteus.
66 Memoir Sears Foundation for Marine Research
vertebral column is generally kept in constant rapid motion, as are the fringed, fleshy
pectoral fins. The caudal filament atrophies at different times in the various species.
Fin rays develop much sooner in the pelvic than in the pectoral fins; hence the pelvics
are actually well-defined fins while the pectorals are still delicately fringed, fleshy stumps.®
The lateral-line sensory system appears early in life, before any scales are formed.
The lateral-line scales develop in a posterior-to-anterior direction, as do the other body
scales. Usually by the time the anterior lateral-line scales are developed, lateral scales
have formed on the caudal peduncle; on the lateral areas, scales develop ahead of those
on the mid-dorsal and midventral regions. Moreover, those of the mid-dorsal area de-
velop ahead of the ones on the midventral section, so the breast is the last to become
scaled. The most posterior fringe of scales that is superimposed on the bases of the
caudal rays does not appear until after the scales on the whole anterior part of the lateral
line and on the body up to the region of the pelvic fins have developed. This terminal
fringe of scales usually contributes one to three scales to the lateral-line count. Fulcral
scales appear after the fins are otherwise well developed. The caudal fin is somewhat
of an exception in that one to three rays form in the superior region following the caudal
filament. The last of these upper rays may not be fully developed until the early juvenile
stage is reached.
Differential growth of structures is pronounced in the Garfishes; although it be-
comes less marked with age, it does not terminate with maturity. The diameter of the
orbit continually changes in proportion to other parts of the body—particularly to
snout length, interorbital distance, and postorbital distance. Primarily because workers
have failed to consider such changes properly, the systematics of these fishes has been
in a chaotic condition for many years.’? Males not only mature at a smaller size but
grow to a smaller size than females, and sexual dimorphism exists in some structures,
e.g. length of snout in the Spotted Gars.
Habits and Food. The Garfishes are sluggish creatures most of the time. During
the summer they frequent the surface waters and appear more or less motionless for
minutes at a time. In streams below barriers, such as dams or falls, frequently large
numbers of Longnose Gar can be seen near the surface on bright sunny days. In this
quiet state, the caudal fin is usually undulated slowly while they are at rest or moving
slowly forward or backward. But they are capable of swimming rapidly for short dis-
tances by rapid undulations of the body. When disturbed, they immediately ‘‘sound.”
In the winter they are found in the deeper waters; the shrimp trawlers frequently
catch Alligator and Longnose Gars in their trawl nets from the deep holes of estuaries
and bayous in the salt marshes of Louisiana.
During the summer months the periodic surfacing of Garfishes is a common sight,
and there is some debate about their successfulness in gulping air during these surface
6. A young specimen of L. osseus without scales and with the peculiar fleshy pectoral fins induced Rafinesque to
describe the form as a new genus, Sarchirus (56: 418).
7. L. Agassiz (4: 360) recognized the remarkable changes that take place in the ontogeny of Garfishes, and
possibly this was the reason he never followed through with the description of 22 supposed new species, most
of which were discovered during a journey in the eastern United States (L. Agassiz, 3: 136).
Fishes of the Western North Atlantic 67
excursions. However, considering their anatomy, results of experimentation, observa-
tions on individuals in their natural habitat, and the regular breathing actions of in-
dividuals removed from water, there is little doubt that the swim bladder and associated
structures leading to it serve as a supplementary respiratory mechanism.
Gars feed primarily on other fishes, most of which are forage species (Bonham, zr:
358-360). There are very few published accounts of cannibalism, but Gars with por-
tions of their tails missing are frequently captured, indicating that they have been
snapped at by other individuals; this has been observed in streams and canals of penin-
sular Florida, where the Florida Gar (L. platyrhincus) is very abundant. In estuarine
waters of Louisiana, Alligator Gar frequently consume the blue crab, Ca/linectes sapidus,
as well as garbage (Weed, 69: 9, 10; Gudger, 26: 120, 121) where refuse is regularly
thrown into the water.
Relation to Man. Although Alligator Gar, L. spatula, attain a large size, no authen-
ticated records of an attack on man are available (69: 9, 10; 26: 120, 121). The usual
response by Gar when disturbed by a bather or fisherman is a dash for deeper water.
While Gars are considered to be a nuisance and are detrimental to game fishes, sport
fishing for Alligator Gars has become popular in recent years in Louisiana, Arkansas,
Mississippi, and other states in the Mississippi Valley.
Alligator Gar are sold in the French Market in New Orleans at the present time,
and small numbers are consumed elsewhere in the United States. Dr. Robert R. Miller
contributes the information that L. sropicus is of considerable importance as a food item
on the Pacific side of southern México and Guatemala. And personal conversation with
Nicaraguans reveals that L. tropicus is used for food in the Lago de Nicaragua area
and lower Rio San Juan. The North and Central American Indians used the ganoid
scales and bones for arrow points, ritual instruments, and ornaments. The late Mr.
Percy Viosca, Jr., of New Orleans, at one time made an array of ornaments and jewelry
from their scales.
Habitat and Range. Their range extends from Quebec in the northeast, west to
the upper Mississippi Valley and Great Lakes region, and south to Costa Rica in
Central America and southwestern Cuba, including the Isle of Pines. Although they
are primarily inhabitants of freshwater streams and lakes, several species enter brackish
and marine waters. L. spatula is a common inhabitant of brackish water in Louisiana,
and several specimens have been captured from the Gulf of Mexico, on the Gulf side
of Breton Island and Grand Isle, Louisiana, and at Destin, Florida. The Longnose Gar,
L. osseus, is somewhat less tolerant of marine waters and is seldom captured outside
of the brackish estuarine areas. The Spotted Gar, L. ocu/atus, is a common summer
inhabitant of the fresher parts of the estuaries in Louisiana.
One species, L. sinensis (Bleeker, 9: 148, 154), supposedly occurs in China (Wag-
ner, 68: 738—741). Most standard references fail to mention an extant Asiatic form.
Geological History. Garfishes are known from the Upper Cretaceous to Lower
Miocene time; in the Eocene of India and in North America from the Middle Eocene
to Recent period (Berg, 8: 214, 415; Romer, 62: 548).
5°
68 Memoir Sears Foundation for Marine Research
Measurements and Counts.
LencrH: measurement from tip of snout to posterior margin of last lateral-line
scale at base of caudal fin. Unless stated otherwise, Lencru is used.
STANDARD LENGTH (SL): nearly equivalent to length.
Tora tenctu (TL): measurement from tip of snout to longest rays of caudal fin.
Caupat pase: defined as a point at end of lateral-line row of scales, located
approximately at center of caudal fin in a vertical direction; this point is used in:
Lencru; CauDaL peDuUNCLE: length; and Caupat Fin: length.
CaupAL PEDUNCLE: length—measurement from caudal base to posterior tip.
Heap: mid-dorsal length—measurement from tip of snout to occiput.
Heap: length—distance from tip of snout to posterior margin of bony opercle.
Heap: depth—measurement at occiput.
Heap: width—measurement at occiput.
Snout: length—measurement from tip of snout to anterior rim of bony orbit.
Snour: least width—measurement immediately behind perforations usually formed
by the large anterior teeth of lower jaw; these tooth perforations are immediately
posterior to the nares and should not be confused.
Snout: width—measurement at posteriormost teeth.
Lower jaws: width—measurement at articulation with skull.
Lower jaws: least width—measurement immediately posterior to symphysis of
dentaries.
PosrorBITAL DISTANCE To: bony margin—measurement from posterior bony rim
of orbit to posterior margin of bony opercle.
PosrorBITAL DISTANCE To: fleshy margin—distance from posterior rim of orbit to
posterior margin of fleshy opercular valve.
Posrsnout: distance—measurement from anterior rim of orbit to free margin of
bony opercle.
FronTaL Bones: least width—measurement at central constriction.
Scass: lateral line—includes the small scales at base of caudal fin, whether pores
are discernible or not, so long as they are within the lateral-line scale row.
Scates: predorsal—includes all median dorsal scales anterior to the paired struc-
tures (fulcra) on the anterior margin and at the origin of dorsal fin.
ScaLes: transverse rows—includes all those in an anteriorly slanted diagonal row
from the single median anal plate to the mid-dorsal scale, including the anal plate and
mid-dorsal scale.
GILL RAKERsS: total—number on left outer arch.
Family LEPISOSTEIDAE
Garfishes
Characters. Those of the Order.
Genera and Subgenera. All modern Garfishes fall within a single genus, Lepisosteus.
Rafinesque (55: 69, 71, 76) placed the Gar or Gar-like fishes in three genera: Sarchirus,
Fishes of the Western North Atlantic 69
Lepisosteus, and Litholepis, the first being based on a young individual of L. osseus and
the last on inaccurate information communicated personally by J. J. Audubon. The
sketch and description (long dorsal and anal, bilobed tail) of the fish in Rafinesque’s
field notes and the description in “Ichthyologia Ohiensis” clearly indicate a fictitious
fish that should not be recognized with a valid name. Rafinesque subdivided the genus
Lepisosteus into two subgenera, Cy/indrosteus and Atractosteus.
Fowler recognized two genera, Lepisosteus and Cylindrosteus (22: 604, 605). Holly
divided the genus Lepisosteus into three subgenera: Litholepis, Lepisosteus, and Cylindros-
teus (65: 49). More recently, Moore (67: 54) and Eddy (1g: 39-41) have recognized
a single genus, Lepisosteus. When Fowler split the Gars into two genera on the basis of
presence or absence of enlarged teeth on the palatine (22: 604), he did not find a second
row of enlarged teeth on the palatine surface in the smallest specimens of L. osseus;
however, small examples of the same form in the Tulane University collection reveal
an inner row of enlarged teeth on the upper jaw.
Two subgenera, Lepisosteus and Atractosteus, are recognized here. The nomen-
clature of fossil Gar material has been confused because of chaotic conditions that
have existed for many years with regard to the recent forms. Perhaps additional sub-
genera will have to be recognized for some of the fossil forms.
Genus Lepisosteus Lacépéde 1803
Lepisosteus Lacépéde, Hist. Nat. Poiss., 5, 1803: 331; type species, Lepisosteus gavialis Lacépéde 1803 equals
Esox osseus Linnaeus.
Generic Synonyms:
Esox Linnaeus (in part), Syst. Nat., roth ed., 1758: 3133 type species, Esox osseus Linnaeus 1758.
Litholepis Rafinesque, Amer. Mon. Mag., 3, 1818: 447; type species, Litholepis adamantinus Rafinesque 1818;
based on a fictitious description personally communicated by J. J. Audubon.
Sarchirus Rafinesque, J. Acad. nat. Sci. Philad., r (2), 1818: 418; type species, Sarchirus vittatus Rafinesque 1818;
young of Lepisosteus osseus.
Cylindrosteus Rafinesque, Ichthyol. Ohiensis, 1820: 72; type species, Lepisosteus platostomus Rafinesque 1820.
Atractosteus Rafinesque, Ichthyol. Ohiensis, 1820: 75; type species, Lepisosteus ferox Rafinesque 1820.
Lepidosteus Koenig, Icon. Foss., 1825: t.12, emended spelling for Lepisosteus Lacépéde 1803; Lepidosteus
Agassiz, Rech. Poiss. Foss., 2 (2), 1843: _Z, emended spelling for Lepisosteus Lacépéde.
Doubtful Synonym:
Psallisostomus Walbaum, P. Artedi Genera Pisc., Emend. Ichthyol., 1792, 581; Psa/lisostomus Fowler (after
Walbaum), Fish. New Jersey iz Rep. N. J. Mus., Pt. 2, 1905: 89; Suppl. Acct. Fish. N. J., Pt. 3, 1906:
263, 387, pl. 83 (after Agassiz, Rech. Poiss. Foss., 1843).
Characters. TEETH numerous, of various sizes on both jaws and on most bones
lining roof of oral and pharyngeal cavities; one row of enlarged teeth with radially
grooved base on lower jaws, and two rows on upper jaws, at least during younger
stages; outer margin of both upper and lower jaws fortified with a closely set row of
needle-shaped teeth; a pair of large teeth at anterior end of lower jaws fitting into
depressions in upper jaws and, in many individuals, perforating anteriormost pre-
70 Memoir Sears Foundation for Marine Research
orbital (maxillary auctorum). A peculiar wedge-shaped Biinp pocxer in skin of gular
region. ToncuE bifurcate-spatulate. Three Mopirrep scares bordering anus; median
scale, anterior to anus, a large shield-shaped plate; lateral plates elongate and slightly
curved. Other characters those of the Order.
Species. Currently seven species are recognized in North American waters, but
detailed studies that are in progress may show that two of these forms are only
subspecifically distinct. There is some indication of hybridization, but verification of
this must await further study. L. oculatus, osseus, platyrhincus, and spatula are treated
in the following account.
Key to Species of the Western North Atlantic
1a. Total gill raker count on left outside arch, 59-81 (70 specimens); palatines of
adult with a series of enlarged teeth. Subgenus 4rractosteus Rafinesque 1820.
2a. Lateral-line scales, 58-62 (25 specimens); diagonal rows between insertion
of pelvic and origin of dorsal fin, 34-383; predorsal scales, 48-54.
3a. Total gill rakers on left outside arch, 59-66 (20 specimens); females
maturing at a larger size, probably not less than 600 mm; anterior body
scales of 400-mm specimens finely serrated on posterior border.
spatula Lacépéde 1803, p. 83.
3b. Total gill rakers, 67-81 (5 specimens); females maturing at a smaller
size, 470 mm; anterior body scales of 400-mm specimens strongly ser-
rated on posterior border. sristoechus (Bloch and Schneider) 1801.
Fresh waters of southwestern Cuba and Isle of Pines; not
yet recorded for salt water but likely to be found there.
2b. Lateral-line scales, 51-56 (60 specimens); diagonal rows between insertion
of pelvic and origin of dorsal fin, 28-32; predorsal scales, 43-48 (usually
44-47). tropicus (Gill) 1863.
Rio Usumacinta of Guatemala and
México and tributaries of Lake
Nicaragua and Rio San Juan in
Costa Rica on the Atlantic slope, and
from Pacific drainage of Chiapas,
México (Miller, 48: 230, 231).8
Atractosteus bocourti Duméril 1870;
mouth of (Rio) Nagualate, 14°N.
1b. Total gill rakers, 14-33 (148 specimens); palatines of adult without enlarged
teeth. Subgenus Lepisosteus Lacépéde 1803.
4a. Snout’s least width 13-25.5 times in its length (specimens more than somm
long). osseus (Linnaeus) 1758, p. 75.
4b. Snout’s least width 4.5—-11 times in its length (specimens more than somm
long).
8. There are no records from salt water on the Atlantic side but L. ¢ropicus is common in the tidal zones of the
Pacific drainages.
Fishes of the Western North Atlantic a
sa. Lateral-line scales, 59-65, usually 60-63 (47 specimens); predorsal scales
50-55, usually 52 or 53; anterior part of body and head without dark
spots. platostomus Rafinesque 1820.
Larger rivers of the Mississippi River
drainage.
5b. Lateral-line scales, 53-59, usually 54-58 (249 specimens); predorsal
scales 45—54, usually 47-50, rarely more than 51; anterior part of body
and head with dark spots and blotches.
6a. Adults with bony plates on ventral surface of isthmus, under gill
membrane (177 specimens); snout and lower jaw longer and nar-
rower. oculatus Winchell 1864, below.
6b. Adults without bony plates on ventral surface of isthmus (76 speci-
mens); snout and lower jaw shorter and broader.
platyrhincus DeKay 1842, p. 81.
Lepisosteus oculatus Winchell 1864
Spotted Gar
Figures 12, 14
Study Material. A total of 177 specimens, 32-819 mm TL, including the type
of L. oculatus Winchell 1864 (dry mount with fins broken; length 700 mm, estimated
caudal length 119 mm, total length 819 mm, UMMZ 55062), from Michigan, In-
diana, and Ohio south to the Gulf of Mexico, and from western Florida to central
Texas along the Gulf coast, in TU, UMMZ, ANSP, USNM, AMNH, and UT
collections.
Distinctive Characters. L. oculatus is readily distinguished from L. osseus and L.
spatula by the profusion of dark spots on the body, head, and fins, and from L. p/aty-
rhincus by the plates on the ventral surface of the isthmus. The Spotted Gar has a head
of medium width, approximately intermediate between the narrow head of L. osseus
and the very broad head of L. spatula.
Description. Proportional dimensions in per cent of length, based on 32 specimens:
from Lake Pontchartrain near New Orleans, Louisiana (5), Sabine R., Texas (3),
Mermentau R., Louisiana (4), Biloxi R., Mississippi (1), Pearl R., Louisiana and
Mississippi (19); specimens 180-512 mm length.
Body: depth 9.2—12.8; width 8.6—12.6. length 27.1-36.2; depth 6.6-8.3;
Caudal peduncle: length 10.8—13.4; least width 7.6—10.0.
depth 5.0—7.3. Snout: length 15.9-23.3; least width
Head: mid-dorsal length 24.5-33.9; 2.3—-3.23 width 3.8—5.3.
g. All of the types listed under Study Material were examined by me.
72
Memoir Sears Foundation for Marine Research
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Ficure 12. Top. Lepisosteus oculatus, 534 mm TL, from Lake Pontchartrain at Goose Point, St. Tammany Parish, Louisiana, TU 6454.
Ficure 13. Borrom. Lepisosteus osseus, 554. mm TL, from Pearl River, Washington Parish, Louisiana, six miles east of Varnado, TU 1119.
Fishes of the Western North Atlantic 73
Orbit: length 2.3-3.3.
Interorbital: least bony distance 4.8—5.9.
Postorbital distance to: bony margin 7.8—
9.8; fleshy margin 8.9—I1.0.
Frontal bones: least width 2.0-3.1.
Lower jaws: least width 1.7—3.0.
Mandible: length 13.6—20.8; posterior
width 4.8—6.9.
Dorsal fin: depressed length 13.7—17.6.
Anal fin: depressed length 12.0-18.7.
Caudal fin: length 13.5-24.3.
Pectoral fin: length 8.2-12.8.
Pelvic fin: length 10.2—1 5.0.
Distance from tip of snout to: dorsal origin
83.2-87.6; pelvic insertion 53.4—
61.2.
Distance from pelvic insertion to: anal or-
igin 23.8—29.9; pectoral insertion
23.5—29.3; lateral line 4.6-6.8.
Distance from dorsal origin to: occiput
52.4—60.2; caudal base 14.0—-17.2;
lateral line 5.2—7.6.
Distance from anal origin to: caudal base
16.8—20.6.
Proportional dimensions in per cent of head length for 31 specimens, 180-700 mm
length, with head lengths 65.1-196.0 mm.
Snout: length 57.0-65.0; least width
6.5—10.6.
Orbit: length 8.2—-11.0.
Interorbital: least bony distance 15.4—
BiTaik«
Postorbital distance to: bony margin 26.4—
Belk.
Lower jaws: least width 5.0—10.8.
Mandible: length 50.0-57.5.
Proportional dimensions in per cent of snout length (sl) and mandible length (ml).
Snout: least width 5.1-9.9 times in sl.
Lower jaw: least width 4.8-11.6 times
in ml,
Statistics of meristic characters: first number—number of specimens; second and
third numbers—extremes in range of variation; fourth number—mean figure.
Scales: transverse rows between pelvic
and dorsal origins 172, 27-32,
30.3; transverse rows 170, 18-24,
20.6; predorsal 173, 45-54, 48.23
lateral line 177, 53-59, 56.1.
Gill rakers: total 32, 15-24, 19.3.
Fin rays: dorsal 33, 6-9, 7.4; anal 33,
7=9, 7.05 .candale 32... 12and 13,
12.5; pectoral on left 32, 9-13;
10.6, and on right 32, 9-13, 10.75
pelvic on left 33, all with 6 rays,
and on right 33, all with 6 rays
except I with 5.
Sexual Dimorphism. It is apparent in this species (Hubbs and Lagler, 33: 76, 77)
and occurs in at least one proportion. The female proportionally has a longer snout
than the male, but this difference is not easily demonstrated because the snout-length/
head-length ratio changes with size of individual.
Color. General coloration darker than in L. osseus; intervening spaces between
numerous dark spots also dark in some specimens, making such individuals appear
74 Memoir Sears Foundation for Marine Research
black; contrast between dark spots and background coloration usually greater in spec-
imens living in clear water. Seemingly the intensity of pigmentation and the area of
body pigmented are somewhat correlated with the color of the water, i.e. how darkly
stained; streams and bayous that course through pinelands and cypress swamps have
light brown to nearly black water and their fish inhabitants tend to be darkened also,
even to the extent of a darkened ventral surface.
The young are brightly colored as in L. osseus (p. 78), but a few differences in
color and pattern are apparent. Median dorsal stripe very broad and dark brown in
contrast to narrow reddish or cinnamon stripe in L. osseus. Dark lateral band nearly
straight on its upper margin, and narrow reddish-brown stripe above usually separated
from lateral band. Ventral surface usually a chocolate color as in L. osseus. Mid-dorsal
stripe and lateral band each breaking up into a single row of spots. Mid-dorsal spots
developing first and forming in a posterior-to-anterior direction. These spots begin
to form when the fish is between 100 and 150 mm length. The lateral spots appear
in the same sequence but do not start to form until the individuals have reached an
approximate length of 170 mm.
Size. The 26 oculatus collected from the brackish waters of Lake Pontchartrain
from July 1953 to February 1955 were 297-690 mm TL. On March 25, 1959,
four ocu/atus were seined in one haul from a borrow pit in the Bonne Carre Spillway,
near Norco, Louisiana. The total length for the largest individual, a female, was
757mm; for the three males, 522, 524, and 575 mm. This was probably a spawning
group, but no spawning activity was observed prior to capture. The female was greatly
distended at the time of capture because of the enlarged ovaries, but no ova were present
in the oviducts.
Development. The scales develop in a posterior-to-anterior manner as described in
Development (p. 66), but the Spotted Gar develops scales and other structures at a small
size. Specimens 100 mm long have a fully developed lateral line as well as lateral body
scales forward to the region of the pelvic fins at the same time as the pectoral fin rays
are just starting to develop. Specimens of 130 mm have scales on the body except for the
anterior belly and breast, and the pectoral fins may have half to all of their rays formed.
The body scalation is complete in specimens 140-150 mm. Plates begin to form on the
ventral surface of the isthmus in individuals of about 200 mm. The enlarged teeth in
the row on the palatines remain as prominent structures until about 200 mm, and some
retain them until they reach 300 mm; however, most individuals have only relatively
small teeth at that size. The caudal appendage may be atrophied in individuals as small
as 160 mm but may not be completely atrophied until a length of 225 mm.
Habits. The feeding habits of this species are similar to those described for L.
osseus (p. 78). In the Lake Pontchartrain area, the diet of the Spotted Gar is composed
of fishes and crustacea, the crustacean most often eaten being the blue crab, Ca/linectes
sapidus.
Relation to Man. The Spotted Gar is usually considered an obnoxious fish.
Range. Lepisosteus oculatus ranges from the Great Lakes south in the Mississippi
Fishes of the Western North Atlantic 75
Valley to the Gulf of Mexico, and from western Florida to central Texas along the
Gulf coast. It does not invade brackish or marine waters as much as either the Longnose
or Alligator Gars. In the Lake Pontchartrain area, L. ocu/atus is found most often
along the marshy shore, and seldom very far out in the lake.
Specimens in the Tulane University collection were obtained from the tidewater
section of Deer River on the west side of Mobile Bay, Alabama, and from Choctawhat-
chee River in Florida. Bailey, et a/. reported the Spotted Gar for the tidewater sec-
tion of the Escambia River, Florida (5: 117). Gunter reported the capture of two from
Copano Bay, Texas (27: 23), and Reid gave records of it for East Bay, Texas (60:
[1955]: 431). All records cited above indicate that, of the four Gar considered, this spe-
cies and the Florida Gar (L. p/atyrhincus) are the least tolerant of saline conditions.
Synonyms, with References to Occurrence in Brackish or Salt Water:
Lepisosteus oculatus Winchell, Proc. Acad. nat. Sci. Philad., 76, 1864: 183 (Huron R., Michigan); Gunter,
Publ. Inst. mar. Sci. Texas, I (1), 1945: 23 (Copano Bay, Texas); Bailey, Winn, and Smith, Proc. Acad.
nat. Sci. Philad., 106, 1954: 117 (in tidewater, Florida); Reid, Texas Sci., 7 (4), 1955: 431 (East Bay,
Texas).
Cylindrosteus productus Cope, Proc. Acad. nat. Sci. Philad., 77, 1865: 86 (San Antonio, Texas).
Cylindrosteus agassizii Duméril, Hist. Nat. Poiss., 2, 1870: 347-348, 351 (St. Louis, Missouri).
Cylindrosteus bartonii Duméril, Hist. Nat. Poiss., 2, 1870: 347-348, 356 (New Orleans, Louisiana).
Lepisosteus osseus (Linnaeus) 1758
Longnose Gar
Figures 13, 15
Study Material. Many specimens, including 42, 139-779 mm TL, used for the
Description; from tributaries of the Gulf of Mexico from Florida to México; from the
Mississippi River drainage, Louisiana to Indiana; in USNM, CNHM, UMMZ,
UI, UL, UT, and TU collections; also, the head only of the type of Lepidosteus lepto-
rhynchus Girard 1858, USNM 1002.
Distinctive Characters. Lepisosteus osseus is distinguishable from L. spatula, L.
oculatus, and L. platyrhincus at a glance by its long narrow snout.
Description. Proportional dimensions in per cent of length, based on 42 speci-
mens, 114—695 mm length, as listed under Study Material. Although the variability of
most dimensions for a limited size range is not very great, the following figures never-
theless include allometric variation.
Body: depth 7.0-10; width 5.9—-9.5. Snout: length 21.4-28.9; least width
Caudal peduncle: length 9.4—13.0; least I.1-2.1; width 2.6-3.7; very nar-
depth 3.7—5.1. row.
Head: mid-dorsal length 29.4-38.8; Orbit: length 2.2—3.7.
length 31.8—41.2; depth 5.4—7.2; Interorbital: least bony distance 3.8—
width 5.3—7.6. Gee
Memoir Sears Foundation for Marine Research
76
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Fishes of the Western North Atlantic a
Postorbital distance to: bony margin 6.6—
8.9; fleshy margin 7.5—10.0.
Frontal bones: least width 1.7—2.5.
Lower jaws: width 3.6—5.2; least width
O.7—1.3:
Mandible: length 20.4-26.4.
Dorsal fin: depressed length 10.2-14.8.
Anal fin: depressed length 11.1-15.9.
Caudal fin: length 12.0-22.3.
Pectoral fin: length 2.6—10.3.
Pelvic fin: length 8.1-13.2.
Distance from tip of snout to: dorsal origin
84.8-89.3; pelvic insertion 56.1—
62.1.
Distance from pelvic insertion to: anal or-
igin 20.4—26.13; pectoral insertion
18.7—24.43 lateral line 3.3—4.7.
Distance from dorsal origin to: occiput
48.2—56.9; caudal base 10.6-15.2;
lateral line 4.2—5.4.
Distance from anal origin to: caudal base
1 5.3-19.2.
Proportional dimensions in per cent of head length for 39 specimens, 114.1-
695 mm length, with head lengths 81.8-272 mm, including head of type (see Study
Materia!).
Snout: length 67.4—73.8; least width
3.1-4.5.
Orbit: length 6.3-9.1.
Interorbital: least bony distance 10.6—
13.4.
Postorbital distance to: bony margin 17.7—
PY op
Lower jaws: least width 1.8—3.6.
Mandible: length 64.3-68.4.
Proportional dimensions in per cent of snout length.
Snout: least width 12.9—-25.7.
Postsnout: distance 2.1—3.0.
Statistics of meristic characters: first number—number of specimens; second and
third numbers—extremes in range of variation; fourth number—mean figure.
Scales: transverse rows between pelvic
and dorsal origins 35, 31-35, 32-53
transverse rows 37, 19-24, 21.2;
predorsal 38, 47-55, 50.8; lateral
line 40, 57-63, 60.1.
Gill rakers: total 38, 14-31, 22.6.
Fin rays: dorsal 40, 6-9, 7.4; anal 40,
8—10, 8.8; caudal 40, 11-14, 12.8;
pectoral on left 39, 10-13, I1.1,
and on right 39, 10-13, 11.2; pel-
vic on left 40 and on right 40, all
with 6 rays.
Color. In general, olivaceous brown above and white below, with specimens from
clear water showing more contrast in coloration; their backs usually more greenish,
those from turbid waters more brownish. Dark spots on median fins and on body,
more clearly defined on specimens from clear water; darkened areas on body frequently
appearing as blackened margins of scales (as if ink were dropped on the body and then
wiped off, leaving traces in grooves between the scales); old individuals sometimes
lacking dark spots on body or fins. Catesby has described and figured ‘“‘acus maxima
squamosa viridis” from Virginia as having a pink belly and pink fins (13: 30, pl. 30).
78 Memoir Sears Foundation for Marine Research
This peculiar coloration may be indicative of a hemorrhagic condition that is easily
produced by exposure and rough handling before preservation.
Young individuals, colorful with various shades of brown to black, and white to
cream. Broad dusky or dark brown band on the side of body, extending from snout
through eye to base of caudal fin; upper margin of band scalloped, the elevated portions
fusing into an interrupted stripe of reddish brown or cinnamon. A narrow mid-dorsal
stripe of similar color. Ventral surface of small young chocolate colored on midbelly
and bordered above on either side by a milky to cream stripe, the dark coloration dis-
appearing in larger young and leaving only a narrow ventrolateral stripe. All fins more
or less spotted or blotched with dark brown, the dark color of fins usually in the form
of bars on pelvic, dorsal, and anal.
Size. The maximum size of osseus is about 1,500 mm TL. Abbott gave a length
of five feet for a specimen speared in Crossweelssen Creek, New Jersey (31: 270). The
28 specimens collected from July 1953 to February 1955 from Lake Pontchartrain
were $41-1,180 mm TL.
Spawning and Development. The early life history of L. osseus is quite well known
(A. Agassiz, r[1878]; Mark, 45). The more extensive studies were made by Wilder
(7I: 155-163; 72: I-12, 192, 195, 10 figs.), Balfour (6: chap. 6), Balfour and Par-
ker (7), and Eycleshymer (21). Agassiz has described the spawning activities and early
development of the Longnose Gar of Black Lake, New York (zr [1879]: 65-75,
5 pls.), and many illustrations represent the developmental stages of the fins and ad-
hesive organs. The adhesive organs on the ventral surface at the tip of the snout on
newly hatched fry are used for attachment to objects above the bottom silt. Although
larvae are relatively inactive until the yolk supply is absorbed, they are capable of very
rapid movements when disturbed. At three weeks of age the yolk has been used, and
the adhesive organs are reduced to a small swelling.
The following data give an approximation of the size of individuals at the time
of atrophy of the fleshy caudal appendage projecting from the upturned end of the
vertebral column. A caudal appendage has been observed at the following lengths: 191,
219, 233, 245, 260, and 271 mm.; but none was seen on specimens of these lengths:
266, 268, 271, 274, 284, and 286 mm. Wilder’s observation of a 300-mm specimen
with the caudal appendage nearly atrophied is in line with measurements given above.
Our largest specimen (271 mm) is actually 323 mm TL when the caudal fin length
is added. Apparently no one has yet determined the maximum age attained by Gar-
fish, but numerous specimens have been kept in aquariums for a number of years.
Mark, who kept L. osseus for nearly 31/. years in aquariums at Cambridge, obtained
the material from Black Lake, New York, when it was in early stages of segmen-
tation (45: 5).
Food and Feeding. The Longnose Gar usually waits for its prey to get within “easy
reach” and then, with a sudden thrust, grasps the victim (Abbott, 37: 269, 270).
The prey is often held crosswise in the mouth for several minutes before it is swal-
lowed. In the brackish waters of Lake Pontchartrain, this Gar feeds on various fishes
Fishes of the Western North Atlantic 79
and crustacea; largescale menhaden (Brevoortia patronus), bay anchovy (Anchoa mit-
chilli), sea catfish (Galeichthys felis), and the blue crab (Callinectes sapidus) were found
in the stomachs by Dr. Rezneat M. Darnell.
Relation to Man. In most areas the Longnose Gar is considered not only an obnoxi-
ous predatory fish but a destructive one as well, gill and trawl nets often being damaged
by it in the brackish waters of Louisiana, and probably elsewhere. In some parts of the
country it is used as food, although most fishermen throw it out on the banks, or
mutilate it before release back into the water. Smith (64: 59) reported (after Earl, 18:
485) that in 1880 this was one of the principal foodfishes in the New Bern market.
The roe, being poisonous, is not used in caviar production as is that of the sturgeons
and the paddlefishes.
Range. It frequents waters from Quebec to Florida along the Atlantic except for
the eastern part of the New England states, and in the west it ranges from the Great
Lakes region south to northern México.
The adult is frequently found in the brackish waters of Louisiana; Smith has
stated that the Longnose Gar sometimes enters salt water and is not rare in Albemarle
Sound (64: 59). In winter it frequents the deeper waters of Lake Pontchartrain, where,
during a biological survey, several specimens were collected with trawl nets from dredge
holes 30 feet in depth. Uhler and Lugger reported it as common in brackish water of
the Potomac and Patapsco rivers (66: 154), but Hildebrand and Schroeder recorded
it as not common in Chesapeake Bay (32: 77). Joseph and Yerger reported three
specimens, 1,000-1,070 mm TL, for Alligator Harbor, Florida (39: 120).
The type of Lepidosteus crassus Cope 1865 was collected from brackish water at
Bombay Hook, near the mouth of the Delaware River.
Synonyms, with References to Occurrence in Salt Water:
Esox osseus (in part) Linnaeus, Syst. Nat., roth ed., 1758: 313; after Artedi, based on “Acus maxima squa-
mosa viridis,” Catesby, 1738: 30, pl. 30 (Virginia).
Esox virdis Gmelin iz Linnaeus, Syst. Nat., 13th ed., Z (3), 1789: 1389.
Lepisosteus gavialis Lacépéde, Hist. Nat. Poiss., 5, 1803: 333 (lakes, rivers of both Indies).
Sarchirus vittatus Rafinesque, J. Acad. nat. Sci. Philad., r (2), 1818: 419, pl. 17, fig. 2 (based on juv.).
Lepisosteus oxyurus Rafinesque, Ichthyol. Ohiensis, 1820: 73 (Ohio R.).
Lepisosteus longirostris Rafinesque, Ichthyol. Ohiensis, 1820: 74 (Muskingum R., Ohio, descr. based on head).
Lepisosteus huronensis Richardson, Fauna Boreal. Amer., 3, 1836: 237 (Penitanguishene, Lake Huron).
Lepidosteus rostratus Cuvier in Richardson, Fauna Boreal. Amer., 3, 1836: 238 (Penitanguishene, Lake Huron).
Lepidosteus semiradiatus Agassiz, Rech. Poiss. Foss., 2, 1836: 2, pl. 2.
Lepidosteus gracilis Agassiz, Rech. Poiss. Foss., 2, 1836: 3.
Lepidosteus bison DeKay in Zool. N.Y., 1842: 271, pl. 43, fig. 139 (Lake Erie, Buffalo, New York).
Lepisosteus lineatus Thompson, Hist. Vermont, 1842: 145 (ill., Winooski R., Burlington, Vermont; based on
juv.).
Macrognathus loricatus Gronow in Gray, Cat. Fish. Coll. and Descr. by L. T. Gronow, in Brit. Mus., 1854:
148 (after Linnaeus).
Lepidosteus leptorhynchus Girard in Pacif. R. R. Surv., Fish., To (4), 1858: 351 (Devil R., Texas; head only,
USNM 1oc0o2).
Lepidosteus otarius Cope, Proc. Acad. nat. Sci. Philad., 77, 1865: 86 (Platte R. near Fort Riley).
Lepidosteus crassus Cope, Proc. Acad. nat. Sci. Philad., 77, 1865: 86 (type from brackish water at Bombay Hook,
near mouth of Delaware R.).
80
Memoir Sears Foundation for Marine Research
ee
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U.S. Hwy. 41, TU 16920.
from Tamiami Canal, Collier County, Florida, 2.7 miles W. of Collier and Dade County line,
488.8 mm TL, from Lake Pontchartrain, Tangipahoa Parish, Louisiana, at mouth of Tangipahoa River, TU 8619.
Ficure 17. Top. Lepisosteus platyrhincus, 516 mm ae;
Ficure 18. Borrom. Lepisosteus spatula,
Fishes of the Western North Atlantic 81
Lepidosteus treculi Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 327 (Mississippi R.).
Lepidosteus milberti Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 328 (New York).
Lepidosteus harlani Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 329, Atlas, pl. 21, fig. 1, 1a, 1b (Wabash R.).
Lepidosteus smithi Dumeéril, Hist. Nat. Poiss., 2, 1870: 323-324, 330 (upper Mississippi R.).
Lepidosteus ayresii Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 331 (Wabash R.).
Lepidosteus copei Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 332 (n. North America).
Lepidosteus lesueurii Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 335 (Wabash R.).
Lepidosteus elisabeth Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 336 (no local.).
Lepidosteus lamarii Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 337 (n. North America).
Lepidosteus clintonii Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 338 (no local.).
Lepidosteus troostii Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 339 (U.S.A).
Lepodosteus piqguotianus Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 340 (Lake Erie).
Lepidosteus horatii Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 341 (n. North America).
Lepidosteus thompsoni Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 342 (upper Mississippi R.).
Lepidosteus louisianensis Duméril, Hist. Nat. Poiss., 2, 1870: 323-324, 343, Atlas, pl. 22, fig. 3 (New Orleans,
Louisiana).
Lepidosteus osseus Giinther, Cat. Fish. Brit. Mus., 8, 1870: 330.
Lepidosteus osseus Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 109; Uhler and Lugger, Rep.
Comm. Fish. Md., 1876: 154 (brackish water, Chesapeake Bay region); Smith, N.C. geol. econ. Surv.,
2, 1907: 59 (brackish and salt water, N. Carolina); Hildebrand and Schroeder, Bull. U.S. Bur. Fisher.,
43, 1928: 77 (Chesapeake Bay); Joseph and Yerger, Pap. oceanogr. Inst. Fla. St. Univ. Stud., 22, 1956:
120 (brackish or salt water, Florida).
Doubtful Synonyms:
Lepisosteus stenorhynchus Rafinesque, Amer. Mon. Mag., 3, 1818: 447; no description (Ohio, Wabash, Green
rivers).
Sarchirus argenteus Rafinesque, Ichthyol. Ohiensis, 1820: 86 (Licking R., Slate Creek; brief descr. based on
communication from Mr. Owings).
Lepisosteus platyrhincus DeKay 1842
Florida Gar, Florida Spotted Gar
Figure 17
Study Material. A total of 76 specimens, 187-516 mm TL; and type of Cy/indro-
steus megalops Fowler 1910, 369.4 mm length, 419.0mm TL, ANSP 25371 (in alco-
hol); from peninsular Florida, in TU, ANSP, and AMNH collections.
Distinctive Characters. Like L. oculatus, L. platyrhincus differs from L. osseus and
L. spatula by the presence of numerous dark spots on the anterior part of its body
and head, but it is distinguished from L. ocu/atus primarily by its lack of plates on the
ventral surface of the isthmus, by its wider snout, and by the wider lower jaws.
Description. Proportional dimensions in per cent of length, based on 30 speci-
mens, 157.0—416.0 mm length; from peninsular Florida.
Body: depth 10.5—14.3; width 9.2—12.0. length 27.4—36.2; depth 7.2-8.5;
Caudal peduncle: length 9.8—13.0; least width 8.1-9.8.
depth 5.7—7.5. Snout: length 15.8-22.5; least width
Head: mid-dorsal length 24.5—-33.1; 2.6—3.4; width 4.1-5.4.
82 Memoir Sears Foundation for Marine Research
Orbit: length 2.6—3.5. Pelvic fin: length 11.4-13.8.
Interorbital: \east bony distance 5.0—5.9. Distance from tip of snout to: dorsal
Postorbital distance to: bony margin 8.3— origin 81.0-88.5; pelvic insertion
10.1; fleshy margin 9.5-I1.5. §2.6—61.3.
Frontal bones: least width 2.2—2.7. Distance from pelvic insertion to: anal ori-
Lower jaws: width 5.3-6.8; least width gin 24.9—32.0; pectoral insertion
2.3-3-5. 23.5—28.1; lateral line 4.5-7.5.
Mandible: length 13.7-19.6. Distance from dorsal origin to: occiput
Dorsal fin: depressed length 13.9—-16.4. 53.8-61.3; caudal base 14.2—16.3;
Anal fin: depressed length 14.3-16.5. lateral line 5.4—7.5.
Caudal fin: length 15.1-19.3. Distance from anal origin to: caudal base
Pectoral fin: length 8.8-11.8. 16.0-19.8.
Proportional dimensions in per cent of head length for 30 specimens, 157-416 mm
length, with head lengths 61.5—118.3 mm.
Snout: length 57.3-62.3; least width Postorbital distance to: bony margin 28.0—
7.5-12.0. B20.
Orbit: length 9.0-11.2. Lower jaws: least width 6.3-12.6.
Interorbital: least bony distance 15.5— Mandible: length 49.3-54.8.
Dao.
Proportional dimensions in per cent of snout length (sl) and mandible length (ml).
Snout: least width 4.8—8.2 in sl. Lower jaws: least width 4.0-8.5 in ml.
Postsnout: distance 1.3—1.6 in sl.
Statistics of meristic characters: first number—number of specimens; second and
third numbers—extremes in range of variation; fourth number—mean figure.
Scales: transverse rows between pelvic Fin rays: dorsal 30, 7 or 8, 7.3; anal
and dorsal origins 76, 30-33, 30,97) Of) 8s 17-65 ‘caudal 30,01 240r
31.7; transverse rows 76, 21-25, 13y112-7\, pectoral. on lett (30;
22.3; predorsal 76, 47-51, 48.6; g-11, 9.9 and on right 30, 9-11,
lateral line 76, 54-59, 56.5. 9.7; pelvic 30, all with 6 rays on
Gill rakers: 30, 19-33, 24.9. both sides.
Sexual Dimorphism. As in the case of L. ocu/atus, sexual dimorphism is apparent
in this species also, i.e. females attain a larger size and proportionally have a longer
snout than males. In length of snout, the females of p/atyrhincus overlap the males of
oculatus.
Color. General coloration on back darker than in L. ocu/atus; coloration and pattern
on ventral surface highly variable, with specimens taken from the same place and at
the same time exhibiting variation from immaculate to solid black. Many specimens
Fishes of the Western North Atlantic 83
with a pattern of two or four or more dark stripes on ventral surface. Two other color
phases also, a brown and an orange (Phillips, 53: 331).
Young, similar in coloration to color given for L. oculatus (pp. 73, 74):
Size. Kilby reported observations on specimens taken off Bayport, Florida, a mile
from shore in the Gulf of Mexico (41: 91); and three specimens, 169, 445, and 495 mm,
were collected from the coastal marsh near Bayport. Hammett and Hammett reported
a maximum of 1,330 mm TL in a sample of 225 specimens (30: 197).
Development. \n this species the development is probably similar to that in LZ. ocu-
/atus (p. 74).
Habits. The diet of this species in brackish and marine waters is unknown.
Relation to Man. Kilby reported that this species is a nuisance to fishermen
(41: 191).
Range. L.platyrhincus ranges from the southern tip of peninsular Florida north-
ward into the lowlands of Georgia. The paucity of records for salt water seems to in-
dicate little tolerance of this species for saline conditions, but there may be a lack of
collections from the areas of its occurrence.
Synonyms, with Reference to Occurrence in Salt Water:
Lepisosteus platyrhincus DeKay, Zool. N.Y., I, 1842: 273, p- 43, fig. 137 (Florida); Kilby, Tulane Stud. Zool.,
2 (8), 1955: 151 (salt water, off Florida).
Cylindrosteus castelnaudi Duméril, Hist. Nat. Poiss., 2, 1870: 347-348, 355, Atlas, pl. 21, figs. 2, 2a, 2b
(Lake Lafayette, Florida).
Cylindrosteus megalops Fowler, Proc. Acad. nat. Sci. Philad., 62, 1910: 609, pl. 38, figs. 15, 16 (Bayport, Florida).
Lepisosteus spatula Lacépéde 1803
Alligator Gar
Figures 16, 18
Study Material. A total of 29 specimens: 28, 187-2,159 mm TL'® (dry mounts
or preserved in alcohol), from western Florida along the Gulf coast to México; in-
cluding the type of Lepidosteus berlandieri Girard 1858, from Matamoros, Tamaulipas,
México, USNM 1003; 1, 49.5 mm length, 57.3mm TL, from Hildebrandt Bayou
near Port Acres, Jefferson County, Texas, TU 22288; all in TU, UMMZ, USNM,
ANSP, AMNH, CNHM, and UT collections.
Distinctive Characters. Lepisosteus spatula is separable from L. osseus and L. ocu-
Jatus by its large size and broad, short snout. The young of spatula are distinguish-
able from the young of oculatus, osseus, platyrhincus, and platostomus by the light
dorsal stripe.
Description. Proportional dimensions in per cent of length, based on 25 speci-
mens, 156.5—1,760.0 mm length; also the 49.5-mm specimen [with measurements
enclosed in brackets].
10. The largest specimen is now at Tulane University (TU Osteol. Coll. 360).
6*
84 Memotr Sears Foundation for Marine Research
Body: depth 9.8-14.2 [12.5]; width
g.1-14.0 [9.0].
Caudal peduncle: length 11.7-14.2
[10.9]; least depth 5.8—7.5 [6.2].
Head: mid-dorsal length 21.5-32.4
[31.5]; length 28.4-36.5 [35.7];
depth 8.2-10.6 [9.4]; width 9.4-
12.9 [9.4].
Snout: length 15.7-21.4 [19.5]; least
width 3.9—-5.1 [5.0]; width 5.3-6.7
[6.8].
Orbit: length 2.5—4.1 [5.6].
Interorbital: least bony distance 7.2—9.0
[7.2].
Postorbital distance to: bony margin 9.7—
11.1 [10.5]; fleshy margin 10.5-
208 (Ltt).
Frontal bones: least width 3.7-4.4 [4.8].
Lower jaws: width 6.1-9.8 [7.8]; least
width 4.5—5.5 [4.6].
Mandible: length 13.7-18.5 [15.9].
Dorsal fin: depressed length 13.4-18.4
uel e
Anal fin: depressed length 13.0-17.9
[14.5].
Caudal fin: length 13.6-19.0 [16.7];
filament [19.3].
Pectoral fin: length 9.1-13.3 [5.0].
Pelvic fin: length 11.1-15.0 [8.0].
Distance from tip of snout to: dorsal ori-
gin 82.8—86.0 [82.0]; pelvic inser-
tion 53.5-59.4 [56.7].
Distance from pelvic insertion to: anal ori-
gin 23.2-30.7 [25.2]; pectoral
insertion 21.0—32.9 [21.4]; lateral
line 5.3-8.2.
Distance from dorsal origin to: occiput
51.9-59.8 [50.9]; caudal base
14.5-17.5 [16.9]; lateral line 5.8—
ips
Distance from anal origin to: caudal base
17.0—20.8 [18.1].
Proportional dimensions in per cent of head length for 27 specimens, 156.5—
1,760.0 mm [49.5], including 2,159-mm specimen, TU Osteol. Coll. 360 (see ftn. 10),
with head lengths 54.6—562.0 mm [17.7].
Snout: length 54.1-59.7 [54.7]; least
width 13.0-15.3 [14.1].
Orbit: length 5.4-12.1 [15.8].
Interorbital: bony distance 23.0—30.5
fizo.3))
Postorbital distance to: bony margin 28.6—
36.5 [29.3].
Lower jaws: least width 13.5-19.2
fir2%o}]:
Mandible: \ength 46.7-52.4 [44.6].
Proportional dimensions in per cent of snout length (sl) and mandible length (ml).
Snout: least width 3.7-4.5 [3.9] times
in sl.
Postsnout: distance 1.2-1.5 [1.2] in sl.
Lower jaws: least width 2.7-3.8 [3.4]
times in ml.
Statistics of meristic characters: first number—number of specimens; second and
third numbers—extremes in range of variation; fourth number—mean figure.
Scales: transverse rows between pelvic
and dorsal origins 24, 34-38, 35.33
transverse rows 27, 23-32, 26.3;
pectoral 27, 49-54, 51.13 lateral
line 27, 58-62, 60.3.
Gill rakers: total 20, 59-66, 62.7.
Fishes of the Western North Atlantic 85
Fin rays: dorsal 22, 7—10, 7.7; anal 22, and 22, 8-16, 12.6 on right; pel-
7-10, 7.8; caudal 22, 12-14, 12.9; vic 22, all with 6 rays except 1
pectoral 22, 11-15, 12.7 on left with 5 on left.
Color. In general, dark olivaceous brown above and white to yellowish beneath,
but some nearly black dorsally; this coloration not unusual for specimens kept in aqua-
riums. Two small specimens (156.5 and 170.5 mm length) dark brown except for light
flesh-color on belly, breast, and throat regions. Skin between mandibles finely speckled
with brown; numerous dark spots on sides—only below lateral line anteriorly, but
above, below, and along lateral line on peduncle posteriorly. Rays of all fins dark brown,
the dorsal, anal, and caudal having noticeably darker spots. Individuals of about 500mm
have few dark spots on sides, and then usually on peduncle; although only a few dark
spots are present, there are numerous places where the groove between the scales is
markedly darker than the surface of the scale; a few dark spots on dorsal, anal, and
caudal fins; some large individuals from Lake Pontchartrain devoid of spots on fins as
well as on body.
Color of 49.5-mm specimen. Light median dorsal stripe extending from tip of
snout to origin of dorsal, and from posterior insertion of dorsal to upper base of caudal
fin; this light stripe bordered on either side by a broad dark brown area extending about
a third of the way down the sides and reaching to dorsal rim of orbit. Dark lateral band
from anterior tip of mandible to anterior rim of orbit, thence posteriorly from posterior
rim of orbit to base of caudal. Dark lateral band extending posteriorly along mandible
from its anterior tip to anterior rim of orbit, and from posterior orbit to base of caudal;
this dark band on body composed of close-set mottlings, forming irregular borders
dorsally and ventrally. Venter light except for dark strip along inside of rami of lower
jaws, thus leaving a light midventral band. All fins with dark brown blotches.
Size. The smallest Alligator Gar seen is the 49.5-mm specimen described above.
The largest one on record, a female taken from Belle Island Lake, Vermilion Parish,
Louisiana, was 9 feet 8.5 inches long and weighed 302 pounds; statistics on other large
individuals are given by Gudger (26: 118-120), Weed (69: 5, 6), and Hussakof (34:
2). During a recent biological survey of Lake Pontchartrain and adjoining brackish
waters, 21 specimens, 410—1,472 mm TL,!! were collected. Additional specimens were
taken from the Gulf of Mexico at Grand Isle, Jefferson Parish, Louisiana.
Spawning and Development. Practically nothing is known of the spawning and
development of the Alligator Gar, but according to the unpublished writings of George
Powers Dunbar (Wortman, 75: 385), it spawns during December and January. The
egg is enveloped in gelatinous material, and the egg strands are draped on snags
and vegetation; by the end of August the young fish has reached 14 inches in length.
These notes by Dunbar need verification. [Recent observations prove the foregoing to
be incorrect. The Alligator Gar spawns in April, May, and June in the Louisiana
11. The remains of several taken from the Gulf side of Breton Island as well as the head of a specimen (ca. 6 ft.) taken
from the Gulf of Mexico at Destin, Florida, are in the Tulane University osteology collection.
86 Memoir Sears Foundation for Marine Research
area. The description of the eggs probably was of some amphibian, as suggested by
Dr. Edward C. Raney (personal communication).
The 49.5-mm specimen (see Description and Color above) has neither scales nor
lateral line developed, the pelvics and pectorals are relatively much shorter than in
older fish, and in the head region, the lower jaws, mandible, and interorbital dimen-
sion are relatively shorter while the orbit is much larger, as is common in the young
of many species. In coloration it is distinguished by the light median stripe from the
snout to the upper caudal except for its interruption by the dorsal.
Specimens 156.5 and 170.5 mm length have the caudal appendage but lack scales
in the mid-dorsal region and on the midventral surface except for a partially developed
anal plate. The rays of the pectoral as well as those of the other fins are fully developed.
Food and Feeding. L. spatula is credited with eating large numbers of game fishes
in fresh water (Gudger, 26: 120), but little has been published about its diet in brackish
and salt water. Raney reported that it ate ducks and water turkey (4uhinga anhinga)
in Cuartez, Resaca, Texas (57: 50), and Gunter found mullet (Muwgi/) in the gut tract
of 12 out of 24 specimens (27: 24). No other food items were mentioned by these
authors. Those examined during the Lake Pontchartrain studies contained striped
mullet (Mugil cephalus) and blue crabs (Callinectes sapidus).
Jordan (35: 313), Weed (69: 9), and Gunter (27: 24) have given evidence that
the Alligator Gar is a scavenger.
Relation to Man. It is sold at present for human consumption in the French Market
in New Orleans, and its scales were used by the Indians (Gowanloch, 24: 389-392)
and by the late Mr. Percy Viosca in his jewelry industry. Many unauthenticated ac-
counts of Gar attacks on humans have appeared in both popular and semipopular liter-
ature, but swimmers probably need have very little fear of them (24: 389-392; 26:
120, 121). With the recent development of underwater spear-fishing, all of the Gars
are desirable targets. Gar fishing rodeos are common annual events in Louisiana.
Range. The Alligator Gar is a frequent invader of brackish and marine waters, and of
the four species it is the most tolerant to higher salinities. It occurs in fresh water in the
Mississippi River and lower parts of its major tributaries from the Ohio and Missouri
rivers southward to the Gulf of Mexico, and in brackish and salt waters along the Gulf
coast from Choctawhatchee Bay, Florida, to northern México (26: 118-120; 69: 5,
6; 34: 2). Gunter recorded Alligator Gars from Copano Bay, Aransas Bay, and from
a Gulf beach, Texas (27: 24), and Reid reported the capture of one from East Bay,
Texas (60 [1956]: 302). Bailey, et a/. gave catch records for the tidewater section of
Escambia River, Florida (5: 117), and specimens frequently on display at the Gulfarium
at Ft. Walton, Florida, are captured in Choctawhatchee Bay.
Synonyms, with References to Occurrence in Salt Water:
Lepisosteus spatula Lacépéde, Hist. Nat. Poiss., 5, 1803: 333 (no local.); Bailey, Winn, and Smith, Proc. Acad.
nat. Sci. Philad., 106, 1954: 117 (tidewater, Florida); Reid, Texas J. Sci., 1956: 302 (salt water, Texas).
Lepisosteus ferox Rafinesque, Ichthyol. Ohiensis, 1820: 73 (Ohio River).
Lepidosteus berlandieri Girard in Pacif. R. R. Surv., Fish., ro (4), 1858: 353 (Tamaulipas, México).
Atractosteus lucius Duméril, Hist. Nat. Poiss., 2, 1870: 360, 364 (Tampico, México).
a
OS
rs.
19.
TEXT AND FOOTNOTE REFERENCES
Aassiz (A.), Proc. Amer. Acad. Arts Sci.,
13, 1878; 14, 1879.
Agassiz (L.), Poiss. Foss., 1833-1843.
Agassiz (L.), Amer. J. Sci., (2) 26, 1853.
Agassiz (L.), Amer. J. Sci., (2) 7, 1854.
Baney, Winn, and Smith, Proc. Acad. nat.
Sci. Philad., 106, 1954.
Balfour, Ganoidei (London), 1881.
Balfour and Parker, Philos. Trans., 773, 1883.
Berg, Class. Fish., Trav. Inst. zool. Acad. Sci.
URSS., 5 (2), 1940; also reprinting by
J. W. Edwards, Ann Arbor, Mich., 1947
(Russ. and Eng.).
Bleeker, Ned. Tijdschr. Dierk. Amst., 4, 1873.
Bohlk, e¢ 2/., Handb. Vergl. Anat. Wirbelt.,
4, 1936.
Bonham, Trans. Amer. Fish. Soc. (1940),
1941.
Bridge, Cambr. Nat. Hist. r, 1904.
Carzssy, Nat. Hist. Carolina, 1754.
Collinge, Quart. J. micr. Sci., N.S. 144 (36),
1894.
Collinge, Proc. Bgham. nat. Hist. Soc., 8, 1893.
Cope, Trans. Amer. philos. Soc., 74, 1871.
Domérn, Hist. Nat. Poiss., 2, 1870.
Err iz Goode, et a/., Fish. Fisher. Industr.
U.S., 2, 1887.
Eddy, How to Know Freshw. Fish. (Iowa),
1957-
87
20.
2I.
22.
39-
40.
4I.
Emelianov, Zool. Jb., (Anat.) 60, 1935.
Eycleshymer, Publ. Univ. Chicago, (Biol.) zo,
1903.
Fowzer, Proc. Acad. nat. Sci. Philad., 62,
IgIo.
Goopricn, Proc. zool. Soc. Lond., 1907.
Gowanloch, Bull. La. Cons. Dep., 23, 1933-
Grassé, et a/., Trait. Zool., 13, 1958.
Gudger, Nat. Hist., N.Y., 49, 1942.
Gunter, Publ. Inst. mar. Sci. Texas, I (1),
1945.
Giinther, Cat. Fish. Brit. Mus., 8, 1870.
Hammarzerc, Acta Zool., 18, 1937.
Hammett and Hammett, Growth, 3 (2), 1939.
Hardwicke, Sci. Gossip, 1873.
Hildebrand and Schroeder, Bull. U.S. Bur.
Fish., 43, 1928.
Hubbs and Lagler, Invest. Ind. Lakes...
Streams, 1943.
Hussakof, Copeia, 1914.
Jeroen: Guide to Study Fish., 1gos.
Jordan, Stanf. Univ. Publ. Biol., 3 (2), 1923.
Jordan and Evermann, Bull. U.S. nat. Mus.,
47 (1), 1896.
Jordan, Evermann, and Clark, Rep. WES:
Comm. Fish. (1928), 2, 1930.
Joseph and Yerger, Pap. oceanogr. Inst. Fla.
St. Univ. Stud., 22, 1956.
Kerr, Proc. zool. Soc. Lond., 1952.
Kilby, Tulane Stud. Zool., 2 (8), 1955.
88
50.
Gis
Memoir Sears Foundation for Marine Research
Lanpacre and Conger, J. comp. Neurol., 23,
1913.
Lankester, Treat. Zool., 9, 1909.
Deleted.
Manx, Bull. Mus. comp. Zool. Harv., 19,
1890.
Matsubara, Fish. Morph. Hierarch., Pt. 1,
1955 (Jap-)-
Mayhew, J. Morph., 38, 1924.
Miller, Copeia, 3, 1954.
Miiller, Abh. Akad. Wiss. Berl., Phys.-Math.
K1., 1844.
Nicxenson, Bull. Mus. comp. Zool. Harv., 24,
1893.
Norman, Synop. Orders, Families, Genera of
Recent Fish. (mimee), 1947.
Preirrer, J. Morph., 54, 1933.
Phillips, Copeia, 1958.
Potter, J. exp. Zool., 49 (1), 1927.
eaeeoue Ichthyol. Ohiensis, 1820.
Rafinesque, J. Acad. nat. Sci. Philad., rz (2),
1818.
57:
58.
59-
60.
6r.
62.
66.
67.
68.
69.
70.
is
72.
73:
74-
75°
76.
Raney, Copeia, 1942.
Regan, Proc. zool. Soc. Lond., 1923.
Regan, Encyc. Britt., ed. 4, 9, 1937-
Reid, Texas. J. Sci., 7 (4), 1955; 8 (3), 1956.
Reissner, Arch. Anat. Phys., 1859.
Romer, Vert. Paleont., ed. 2, 1945.
Scurry, Arch. Naturgesch., 62 (1), 1896.
Smith, N. C. geol. econ. Surv., 2, 1907.
Das Tierreich (Berlin and Leipzig), 1936.
\
Unzer and Lugger, Rep. Comm. Fish. Md.,
1876.
VerrepraTEs of U.S., 1957.
Wacner, Zool. Anz., 39, 1912.
Weed, Leafl. Field. Mus. nat. Hist., 5, 1923.
Wiedersheim, Zool. Jb., Suppl., 7, 1904.
Wilder, Amer. Ass. Adv. Sci., 24 B, 1876.
Wilder, Pop. Sci. Mon., rz, 1877.
Williamson, Philos. Trans., 739 (2), 1849.
Williamson, Philos. Trans., T4I (2), 1851.
Wortman, Amer. Nat., 16, 1882.
Wright, J. Anat., London, 19, 1885.
Order Isospondylt’
COMPOSITE AUTHORSHIP
Characters and Keys to Suborders
and Families
HENRY B. BIGELOW
Museum of Comparative Zoology
Harvard University
Acknowledgments. Special thanks are due the following persons for their invaluable
help and cooperation in preparing the text and keys of this section: William W.
Anderson, ‘fames E. Bihlke, Daniel M. Cohen, F. R. Dymond, Robert H. Gibbs,
Fr., W.A. Gosline, Marion Gray, N. B. Marshall, Giles W. Mead, Fames E.
Morrow, Fr., George S. Myers, Albert E. Parr, and Robert R. Rofen.
Characters of Living Isospondyli. Both premaxillary and maxillary bones are present
in most, but in a few these bones are greatly reduced or lacking (among Argentinoidea,
Part 4). In most the maxillaries form a part of the border of the mouth (see also
pp. 11, 12, 17, 92, 95). The premaxillaries are only slightly protractile, if at all so,
except in the Phractolaemidae,? some Alepocephalidae, and some Stomiatoidea (see
Bertin and Arambourg, 28: 2247), where they are protractile. The bone in the
ethmoid position above the vomer is unpaired in most but paired in a few (pp. 95, 96).
There are four gill clefts, and in a few there is an open slit between the mandible
and the hyoid arch (Malacosteidae).
1. With accounts of the included Suborders, Families, Genera, and Species by various authors as listed in the Table
of Contents. In this series of volumes, the Iniomi, treated as a Suborder of the Isospondyli by some authors, is
given the rank of Order and is dealt with in Memoir I, Part 5.
2. A family of the Chanoidea, or so-called milkfishes, of the tropical Pacific.
89
go Memoir Sears Foundation for Marine Research
Typically the fins have no supporting spines,’ most of the rays being branched;
however, in a few the rays are unbranched at the origin. Typically the pectorals* are
posterior to the gill openings, but in the genus Asguamiceps (Alepocephalidae) they are
well inside the gill cover; in the more primitive genera their line of insertion is low
down on the side near the ventral profile, but in some of the alepocephalids and argen-
tinoids they are inserted higher up on the sides; the pectoral bases are far below the
upper end of the gill openings except in some of the bathylagids, where the upper end
of the gill opening is greatly restricted. The pelvic fins,® if present, are posterior to the
pectorals, their positions ranging from abdominal to thoracic. In most there is only one
dorsal fin with jointed bony or membranous rays (lacking in a few); in some there is
a fleshy “‘adipose’’ dorsal fin with unjointed horny rays between the rayed dorsal and
caudal fins; also, in a few (some Astronesthidae) there is a second adipose fin close in
front of the anus. The rayed dorsal, or the adipose dorsal if present, is separated from
the caudal by a definite gap, and in most the anal fin is similarly separated from the
caudal.® The caudal fin is about the same in width both above and below the projected
longitudinal axis of the trunk,’ and the rear boundary of its fleshy base is symmetrical
relative to the axis of the trunk; in most, the rear outline of the caudal is more or less
deeply forked, or at least concave, but in some it is more nearly truncate transversely,
and in a few it is either rounded or has two or three of the middle rays extending almost
equally beyond the others (genus Pantodon).
The radialia of the pectoral fins are articulated basally with the pectoral girdle.
In the great majority the pectoral girdle has a mesocoracoid element firmly attached
above to the auditory (otic) region of the skull (see also pp. 2, 6). In the pelvic fins
the basal radialia are reduced to a few small nodules, and the pelvics are not attached
to the pectoral girdle. The number of skeletal supports equals the number of rays in
the dorsal and anal fins.
The head, though naked in the great majority, is clothed with smooth scales among
the Esocoidea and in the genus Lepogenys (Alepocephalidae) and with prickly scales
in Gonorhynchus; in no living representative of the Order’ is the head armored with bony
plates. And in no living family are the margins of the caudal fin armed with more than
one enlarged “‘fulcral” scale, such as is characteristic of the caudal of the sturgeons®
3. Among the Idiacanthidae the base of each dorsal and anal ray bears on its anterior side a pair of short sharp
spurs that project through the skin (for details, see Beebe, 3: 152, 213). Among the Sternoptychidae, the rayed dorsal
is preceded by the projecting spine-like tips of approximately 1-7 of the neural processes of an equal number of
vertebrae (for an early account, see Cuvier and Valenciennes, 15: 395, 402, 419).
4. The stomiatoid genera Idtacanthus, Photostomias, and Tactostoma (described by Bolin, 6: 39), are said to have no
pectorals; likewise some specimens of Eustomias and Photonectes.
5. Among many families, such as the salmonids, specimens occasionally lack one or both pelvics, but this is an indi-
vidual anomaly (see Myers, 29: 600-601; 28: 4r).
6. Not in the freshwater Notopteridae of Africa and the East Indies, or in the genus Coi/ia (Engraulidae) of the
tropical warm-temperate Indo-Pacific.
7. Most of the fin rays, however, are “‘attached to the lower spines of the hinder vertebrae, which are greatly
enlarged, and at the same time inclined backwards so as to be more or less parallel to the axis of the body”
(Norman, jr: 61).
8. In the fossil Leptolepidae the dermal bones of the skull were covered with an enamel-like substance.
g. These fuleral scales were “‘present, but usually small” among the fossil Pholidophoridae (Boulenger, 7: 545).
Fishes of the Western North Atlantic gI
and gars (Fig. 3). The body scales, lacking in only a few,!° are thin and overlap each
other in most instances, but in some they are thick and bony; the exposed surface
is smooth or only finely striate except in a few where it is either granular and sculptured
with a network of low ridges or has fine!? or coarse! prickles; in most the exposed
edges are even, or nearly so, but in a few! they are pectinate; the exposed portion of
the scale is not enamel-like in any living family.%
A lateral line is either present or absent, as are luminescent organs.
The swim bladder, if present, is dorsad in position’® and is either closed or con-
nected to the anterior part of the alimentary tract by an open duct; it is not con-
nected to the inner ear by a chain of bonelets (Weberian ossicles). The oviducal
tracts along which the ova pass to the exterior consist either of closed tubes (oviducts
complete) or of membranous channels that are open above but are enclosed by lon-
gitudinal folds of the peritoneum (oviducts incomplete)” The inner wall of the in-
testine in a few has a series of circular ridge-like thickenings which suggest the spiral
valve of elasmobranchs, chimaeroids, and polypteroids; however, they are probably not
homologous (Cohen, 10: 96).
The last few vertebrae are turned upward in a few (Salmonidae). If present, the
lateral processes (parapophyses) that bear the pleural ribs are simply set in pits in
the vertebral centra in some but are fused with the centra in others.
Nomenclature. The names Isospondyli'* and Malacopterygii!® have been used inter-
changeably as ordinal names by various authors for nearly a century. More recently
the name Clupeiformes, proposed for this Order by Berg (4: 216, 417), has been ac-
cepted by Bertin and Arambourg, but in this series of publications the name Isospondyli
is preferred, following Cope (IT: 454, 455).
10. Among the Alepocephalidae, Stomiatoidea, and Galaxiidae.
tr. Among freshwater osteoglossids of South America, Malaya, and Australia.
12. In at least two species of Argentina (see Smitt, 50: fig. 229).
13. In the genus Gonorhynchus of the warm-temperate Indian Ocean and western Pacific. For an excellent illustra-
tion, see Cuvier and Valenciennes (z3: pl. 568).
14. In the genus Brevoortia, p. 342.
15. The exposed surface of the scales was covered with an enamel-like substance (ganoine) in the fossil family Lep-
tolepidae, which was abundantly represented from the Upper Triassic to the Middle Cretaceous, and in the
Pholidophoridae of the Jurassic.
16. The ventral “sole” of Opisthoproctus, interpreted by Trewavas as a swim bladder (53: 610), appears to be part
of a light organ (Bertelsen, 5: 862).
17. The original idea (Rathke, 37) “that the salmonoids have no oviducts and that the ova are deposited free in
the abdominal cavity has been handed down to the present day in all literature pertaining to the subject”’
(Kendall, 24: 190). Similarly, according to Cuvier, the ovaries in Argentina discharge the eggs into the abdom-
inal cavity “comme dans les autres Salmonoides” (14: 410). Trewavas could trace no oviduct in a specimen of
Opisthoproctus cleared in glycerine and caustic potash before dissection (53: 610, 611), nor could Beebe do so
in a mature female of Dolichopteryx (2: 78). It has been reported similarly that the ova are discharged into the body
cavity in the Galaxiidae, the Haplochitontidae, the Hyodontidae, and the Notopteridae. However, Kendall
showed from his own dissections that the actual situation, in both the Salmonidae and the Osmeridae, is as
summarized in the text above, and that the ova cannot be extruded if they be displaced in the abdominal cavity.
Probably this is equally true of the other bony fishes that have been reported as lacking oviducts.
18. Proposed in 1871 by Cope (rr: 454-455).
1g. First proposed in 1738 by Artedi (r: 1), it was given its first post-Linnaean definition in 1893 by Gill (17: 130-
131), as was Cuvier’s “Malacopterygiens abdominaux” (r2: 159), which also included the Ostariophysi. Mala-
copterygii was adopted by Boulenger in 1904 (7: 543).
92 Memoir Sears Foundation for Marine Research
The Order Isospondyli, as defined under Characters of Living Isospondyli, cor-
responds to: the Order Isospondyli of Cope;?® the Suborder Isospondyli in part of
Woodward (54: xxxv1; 55: xvi1); the Suborder Malacopterygii plus the Suborder
Haplomi in part of Boulenger (7); the Suborders Clupeiformes and Esociformes of
Goodrich (25: 386, 397); the Orders Isospondyli (excluding the Mormyriformes) and
Haplomi of Regan (38: 77-78); the Orders Clupeiformes plus Galaxiiformes of Berg
(4: 216, 254, 256, 417, 436, 437); the Order Isospondyli of Romer (45: 581, 584)
plus the Suborder Esocoidea, which he referred to the Order Mesichthyes of Hay
(20: 397); and the Order Clupeiformes of Bertin and Arambourg (z8: 210).
The members of the Order Isospondyli include some of the most generalized of
living bony fishes, and the fossil record for some of them reaches as far back in geo-
logic time as the Lower Cretaceous (p. 15). The Order is accepted here as a matter
of convenience, since the interrelationships of its included units still remain obscure.
Suborders. The heterogeneous assemblage that is grouped together in Part 5 as the
Order Iniomi has been treated as a Suborder of the Isospondyli by some authors
(Schultz and Stern, 47: 233; Bertin and Arambourg, 18: 2269) but as a separate Order
by Regan (38: 77-78; 41: 314), Jordan (22: 153), Norman (30: 317), Berg (4: 242,
256, 429, 437—as Scopeliformes), and Marshall (26: 305-336). In accord with pres-
ent-day tendencies based on evolutionary grounds, discussed below, the Iniomi are
here classed as a separate Order.
In 1929 Regan (41: 313, 314) set the Iniomi apart from the Isospondyli as a
separate Order on the following grounds:
1a. Maxillaries (typically) forming a part of upper border of mouth; pectoral girdle
with mesocoracoid element in the great majority. Isospondyli.
1b. Maxillaries not forming a part of upper border of mouth; pectoral girdle without
mesocoracoid element. Iniomi.
Unfortunately, the separation between Iniomi and Isospondyli is not as clear-cut
as the foregoing implies. Thus the mesocoracoid is lacking: in some of the Argenti-
noidea (53: 609, 612); in the Salangidae (Salmonoidea); in the Retropinnatidae, the
systematic relationships of which remain uncertain; in the Esocoidea (Part 4), the
Bathylaconoidea (35: 52), and the Aplochitonidae (40: 290), which have sometimes
been grouped with the Salmonoidea but which seem to deserve the rank of a separate
Suborder. Furthermore, the Iniomi share the withdrawal of the maxillary bone from
the upper border of the mouth with some fishes that are isospondylous in other respects:
i.e. the genus 4/bula (Albulidae), the Pterothrissidae (here rated as a separate family),
Nematalosa (44: 465, fig. 127, as ““Chaetoessus”), and the genus Chanos.
Such members of the Order Isospondyli as are known to occur in the western
North Atlantic are distributed in Parts 3 and 4 among the Suborders Elopoidea, Clu-
20. The systematic position of Cope’s (rz: 454) Scyphophori (freshwater families Mormyridae—distinguished espe-
cially by the enormous cerebellum—and Gymnarchidae) remains controversial.
Fishes of the Western North Atlantic 93
peoidea, Stomiatoidea (including its subdivisions Gymnophotodermi, Heterophoto-
dermi, and Lepidophotodermi, named by Parr [33: 15-17; 34: 136]), Salmonoidea,
Argentinoidea,#* Esocoidea, and Bathylaconoidea.
These Suborders correspond to: the Suborders Clupeoidei, Salmonoidei, Stomia-
toidei, Opisthoproctoidei, and Esocoidei of Berg (4); the Suborders Clupeoidei, Sal-
moniformes, Opisthoproctoidea, Stomiatoidea, Gymnophotodermoidea, and Esoci-
formes of Schultz and Stern (47); the Suborders Clupeoidea, Stomiatoidea, and Salmo-
noides, plus the Order Haplomi, of Norman (37); the Suborders Elopoidei, Clupeoi-
dei, Salmonoidei, Opisthoproctoidei, Alepocephaloidei, Stomiatoidei, Bathylaconoidet,
and Esocoidei of Bertin and Arambourg (18: 2211); and the Suborders Elopina,
Albulina, Esocina, Clupeina, Chanina, Gonorhynchina, Salmonina, Opisthoproctina,
and Stomiatina of Matsubara (27: 178-227).
One major area of uncertainty as to the relationships among the groups of iso-
spondylous fishes centers around the position of the Elopidae, Albulidae, and Ptero-
thrissidae. Indeed, within recent years it has been questioned whether E/ops is even of
the same lineage as other living teleosts. Thus Saint Seine has derived it from the
Halecostomi (46: 297), a group known otherwise from fossils only (18: 2195-2201),
and Nybelin has emphasized the holostean nature of certain characteristics of E/ops,?®
embodying a similar point of view.
Both the Elopidae and the Albulidae, to be sure, have characteristics that are con-
sidered archaic by both palaeontologists and neozoologists alike; among these are: the
retention of the gular plate and of the numerous branchiostegal rays in E/ops and Tar-
pon; the arrangement of the mucous canals on the top of the snout in E/ops (32: 454,
fig. 1); the presence of close-set teeth on the parasphenoid bone in the roof of the
mouth (Elopidae, Albulidae, and Pterothrissidae); the presence of roofed post-temporal
fossae; and the persistence in A/bula, Pterothrissus, and Megalops (but not in E/ops) of
two rows of valves in the arterial cone of the heart.?%
On the other hand, the elopids and the albulids differ widely and conspicuously
from living members of Lepisosteus and Amia in the narrow rod-like nature of their bran-
chiostegal rays and in the nature of the skeleton of their caudal fin; this latter difference
is a conspicuous one, externally, for while the rear boundary of the fleshy base of the
caudal is symmetrical relative to the main longitudinal axis of the trunk in the elopids
and albulids, it slopes upward-rearward in Lepisosteus (Fig. 3, p. 16) and Amia. Further
features worth mention are: (1) the invariable presence in the elopoids of a median
supraoccipital bone, which, to a greater or lesser degree, is involved in the roof-complex
of the skull; (2) the absence of coronoid teeth in the mandible; (3) a one-to-one rela-
tionship between the principal caudal rays and hypurals of 4mia and Lepisosteus com-
pared with the considerable consolidation of the hypural elements in the elopids; (4)
the absence in elopids of the rigid attachment of the premaxillaries to the skull, which
21. The reason for using this name rather than Opisthoproctoidea is given in Suborder Argentinoidea, Part 4.
22. “Les Elopidae m’apparaissent etre des Holosteens aussi bien qu’Amia ou Lepidosteus’” (32: 458).
23. See Senior for account of the cone for Tarpon (48: 146-151), for Elops (48: 150), and for a summary (49: 83-84).
94 Memoir Sears Foundation for Marine Research
characterizes the living holosteans; and (5) a well-developed articulation with the
autopalatines in elopids. Indeed, in the development of the whole upper jaw mechanism,
the elopoids are considerably in advance of the holosteans.
However, though we accept the traditional view that the elopids and albulids
(with Prerothrissus) do belong among the Isospondyli, there is much uncertainty as to
their disposition within that Order. For example, Jordan and Evermann (23: 407),
followed by Norman (31: 63), rated them as families within the Suborder Clupeoidea;
Berg (4: 221, 222; 419, 420) defined them as superfamilies (Elopoidae and Albuloi-
dae), each to include two families; Fowler (16: 151) and Poll (36: 8) rated them as
families of Isospondyli (Clupeiformes of Poll) without the intervention of Suborders.
However, as early as 1893, Gill (17: 127-128) made them the basis of a separate Sub-
order, Elopoidea, in which he was followed by Bertin and Arambourg (z8: 2211),
who stressed in particular the primitive nature of their caudal fin skeleton.*4 Jordan
(22: 117-118), proceeding a step further, proposed the Suborder Elopoidea for the
elopids and the Suborder Albuloidei for the albulids as contrasted with the clupeoids
and with other Suborders of Isospondyli; Matsubara® has also accepted this arrange-
ment.
Exclusion of the elopids and albulids from the clupeoids clearly seems demanded,
for among living fishes they appear to stand much lower on the evolutionary tree than
any other teleosts that are at all well known. But it still remains an open question whether
the features in which the elopids and albulids agree, balanced against those in which
they differ, are better represented by placing them in two separate Suborders or by
uniting them in one, as is done here.
The isospondylids as here defined include the ladyfishes or bigeye herrings, the
tarpons, the bonefishes, the true herrings, the anchovies, and the salmons, all of which
have been familiar for generations to seaside dwellers in general in one part of the world
or another. They also include such of the pikes as enter brackish or salt water, and
an assemblage of oceanic fishes, many of which have luminescent organs and are
bizarre in appearance; the latter are seldom seen by ordinary seafarers, commercial
fishermen, or anglers, and little is known about their mode of life.
Key to Suborders. Although the general characteristics are now tolerably well
known for the elopoid, herring-like, salmon-like, and pike-like fishes, we still have so
much to learn about the internal anatomy of the less familiar groups of Isospondyli
that their natural affinities remain obscure in many respects. Areas of continuing un-
certainty are, for example: the relative weights to be accorded one character or
another from the evolutionary standpoint; the breadth of coverage to be allotted to
the herring-like fishes and the salmon-like fishes; and the disposition to be made of
various outlying families such as the Bathylaconidae (Part 4), which do not readily fit
in any of the major subdivisions that are recognized generally. Furthermore, Suborders
24. For accounts of the caudal fin skeletons of Elops, Albula, and Megalops, see Regan (39: 355, 356, fig. 1); see Hollister
(21: 260-276) for Elops and Albula.
25. Elopina and Albulina, Suborders of Order Clupeida (equivalent to Isospondyli) (27: 180-182).
Fishes of the Western North Atlantic 95
that are easily separable typically may intergrade with one another marginally. For
instance, the presence of parietal bones among the Clupeoidea and not among the
Stomiatoidea has been invoked as an alternative between these two Suborders, but
this is not strictly so, since some stomiatoids do have parietals (see Regan and Tre-
wavas, 43: figs. 6, 9, 10; Giinther and Deckert, rg: 244). Neither is the arrangement
of the photophores on the sides in definite longitudinal rows reliably diagnostic for the
Stomiatoidea, since they are not so in one species of Neonesthes, while in one other
species the linear arrangement is to be recognized only with difficulty,** as is true of
some species of Astronesthes as well.27 And Mrs. Marion Grey contributes the infor-
mation that some species of Cyc/othone, Gonostoma, and Malacosteus have no photo-
phores at all on the sides of the body.
Under these circumstances the construction of a satisfactory key to the Suborders
of Isospondyli that would withstand the test of time is not to be hoped for at present.
Tentative Key to Suborders of Isospondyli of the Western
North Atlantic
1a. Parasphenoid bone on roof of mouth with low close-set teeth; larval stage ribbon-
shaped (leptocephalus-like). Elopoidea, Part 3, p. 107.
1b. Parasphenoid bone without teeth; larval stage in most not ribbon-shaped.*8
2a. Premaxillary bones minute; maxillaries forming almost entire length of upper
jaw; branchiostegal rays plate-like, the edge of the uppermost attached to
lower margin of subopercular bone and thus taking part in formation of gill
cover. Bathylaconoidea, Part 4.
2b. Premaxillary bones well developed; branchiostegal rays not plate-like; the
edge of upper branchiostegal ray not joining subopercular and interopercular
bones and thus not taking part in formation of gill cover.
3a. Luminescent organs (photophores) absent in most; never present on head,
but present on eye-tube in a few (Do/ichopteryx of Argentinoidea); if
present on sides of body they are in nonlinear arrangement except along
base of anal fin (Binghamichthys of Alepocephalidae); some with a tubular
papilla on each shoulder that connects with a voluminous sac beneath
the skin (Searsiidae).
4a. No adipose fin between rayed dorsal fin and caudal.
sa. Snout noticeably flattened dorsoventrally (depressed), in duck-
billed form; teeth along rear part of lower jaw large, formidable,
conspicuous; premaxillary bones far apart in front; mesethmoid
(proethmoid) bone paired. Esocoidea, Part 4.
26. Information contributed by Robert H. Gibbs.
27. According to Zugmayer (56: 4), the photophores are distinguishable only with difficulty in Astronesthes niger (as
A. myriaster), but Regan and Trewavas (42: 20, fig. r2) picture them clearly for the type specimen of that species.
28. Certain of the Stomiatoidea have ribbon-shaped larvae; Part 4.
96 Memor Sears Foundation for Marine Research
5b. Snout not flattened dorsoventrally, its dorsal contour convex;
teeth along rear part of lower jaw small and inconspicuous; pre-
maxillary bones close together in front; mesethmoid (proeth-
moid) bone not paired. Clupeoidea, Part 3, p. 148.
4b. Adipose fin between rayed dorsal fin and caudal present in most, but
lacking in a few (see 6b).
6a. Swim bladder connected to oesophagus throughout life by
an open pneumatic duct (physostomic); premaxillary bones
well developed; jaws with well developed teeth except in the
Coregonidae, which are either toothless or have minute teeth;
eyes of ordinary type; oviducts incomplete (p. 455), so far as
is known. Salmonoidea, Part 3, p. 455.
6b. Swim bladder not connected to oesophagus during late
stages of growth (physoclystic);2 premaxillary bones small or
lacking; jaws toothless; eyes tubular in some but not in others;
oviducts incomplete in some,?° perhaps complete in others.
Argentinoidea,™ Part 4.
3b. Luminescent organs (photophores) present on head, on body, on tail
sectors, or on all three; their arrangement in longitudinal rows on sides
extending anterior to anus as well as along base of anal fin; no tubular
papilla on shoulder. Stomiatoidea (including Suborders Gymno-
photodermi and Lepidophotodermi*), Part 4.
Families. The Isospondyli plus the Iniomi include about 55 families on a world-
wide basis, about 45 of which are known to occur in the western North Atlantic. At
first glance, this may seem an appalling number among which to choose when one has
to run down a given specimen to its proper family, the more so because there is no
one conspicuous character or combination of characters visible from the outside on
which anyone but a professional taxonomist can rely to tell him whether his fish be-
longs among the Isospondyli or among the Iniomi. The task, however, is not as dif-
ficult as one might expect, for nearly all of the western Atlantic families (or subdivisions
of families) of the two Orders combined are separable, one from another, by features
that are visible from the outside, that are susceptible of rather precise definition, and
that are mutually exclusive, or nearly so.
However, in numerous instances a family, because of the diverse characters of
its genera and species, has been split into two or more parts. Hence the same family
29. Cuvier and Valenciennes seem to have been the first to record this fact for Argentina (14: 411). Trewavas’ interpre-
tation of the ventral “‘sole’’ of Opisthoproctus as a swim bladder is now known to have been erroneous (53: 610).
30. This is the case for Argentina, Opisthoproctus, and Dolichopteryx (Part 4). For Macropinna, however, Chapman
reported that a “‘fine duct” extends from the posterior end of each ovary, but he could not determine whether these
ducts open into the urinary bladder or into the rectum (8: 293).
31. For a detailed comparison of the Argentinoidea and Salmonoidea, see Argentinoidea, Part 4.
32. Proposed by Parr, 33: 15-17.
Fishes of the Western North Atlantic 97
may be referred to in different parts of the same key or in one or more of the other
keys. For example, such of the Bathypteroidae as have no adipose dorsal fin are
found in Key A while those with an adipose fin are found in Key B.
General Key to Families of Isospondyli, Iniomi, and
Giganturoidei of the Western North Atlantic; Adolescents and Adults?
1a. Eyes (if any) of the usual sort, i.e. not at tips of slender stalks or with pupil above
center, and not capable of being directed upward.
2a. No luminescent organs (photophores) either on sides of body, or on head.
3a. No adipose fin between rayed dorsal fin and caudal. Key A, below.
3b. Adipose fin present between rayed dorsal and caudal. Key B, p. 99.
2b, Luminescent organs (photophores) present on head, or on body, or on both;
or with eyes on top of head. Key €; p: 101.
1b. Eyes at tips of slender stalks, or with pupil above the center and capable of being
directed upward. Key D, p. 103.
Key A. No DoRSAL ADIPOSE FIN; NO PHOTOPHORES ON SIDES OF HEAD OR BODY.
1a. Rays in posterior half of dorsal and in rearmost part of anal much longer than
those in anterior part of these fins. Macristiidae.4
1b. Not more than one dorsal ray (if any) and no anal ray much longer than the others.
2a. Bony (gular) plate present in chin region between branches of lower jaw.
Elopidae, Part 3, p. 111.
2b. No bony (gular) plate in chin region.
3a. Upper jaw reaching rearward far beyond eye.
4a. Snout overhanging mouth noticeably.
Engraulidae (western Atlantic saltwater species), Part 3, p. 152.
4b. Snout not overhanging mouth.
ga. Pectoral fin longer than head; either uppermost pectoral rays
or outermost pelvic rays much longer than the others; lower lobe of
caudal much longer than upper in some.
Bathypteroidae in part, Part 5.3°
5b. Pectoral fin shorter than head; none of pectoral or pelvic rays
prolonged; lower lobe of caudal little, if any, longer than upper.
6a. A fleshy ridge present along back anterior to dorsal fin.
Alepocephalidae in part
(Anomalopterus), Part 3, p. 250.35
6b. No fleshy ridge on back anterior to dorsal fin.
33- For larval stages of a few, see Key D, rb. In this key no attempt is made to present the author’s views as to phylo-
genetic relationship.
34. Occur in the vicinity of the Azores, hence may be taken sometime in the western Atlantic.
35- See Key B, 4a for those with adipose fin. 36. See also Key A, 11a; Key C, rsa.
98 Memoir Sears Foundation for Marine Research
37-
39-
40.
7a. Eyes minute, about same diameter as nostril.
Ipnopidae in part (Bathymicrops,
Bathytyphlops), Part 5.37
7b. Eyes well developed, much larger than nostril.
Bathysauridae in part, Part 5.38
3b. Upper jaw reaching rearward little (if any) past eye, and ending con-
siderably short of rear edge of eye in most.
8a. Snout overhanging mouth; parasphenoid bone rearward along mid-
roof of mouth thickly set with low teeth.
ga. Base of dorsal occupying only about 24-2 5°/, of distance between
gill openings and origin of caudal. Albulidae, Part 3, p. 132.
gb. Base of dorsal occupying about 80°/, or more of distance be-
tween gill openings and origin of caudal. Pterothrissidae.?®
8b. Snout not overhanging mouth; parasphenoid bone without teeth.
10a. Rear end of base of dorsal fin separated from upper origin of
caudal by a distance much shorter than head.
11a. Upper jaw not reaching rearward beyond front of eye;
teeth inconspicuous.
Alepocephalidae in part, Part 3, p. 250.4
11b. Upper jaw reaching rearward about as far as rear edge of
eye; teeth along rear half of lower jaw large and formi-
dable. Esocidae, Part 4.
rob. Rear end of base of dorsal fin separated from upper origin of
caudal by a distance at least as long as head, and longer in most.
12a. Midline of abdomen in front of anal fin armed with a
double series of stiff, pointed scales (scutes).
Clupeidae in part (Clupeinae, pp. 274-4113 Pristigaste-
rinae, pp. 411-438; Chirocentrinae, pp. 438-442; Do-
rosomatinae, pp. 443-451), Part 3, Key pp. 259—-262.41
12b. Midline of abdomen in front of anal fin rounded; not
armed with a double series of scutes.
13a. Origin of dorsal fin considerably anterior to mid-
length of trunk; pelvics posterior to dorsal; max-
imum depth of trunk equal to about 30°/, of dis-
tance from gill opening to base of caudal; head about
25°/, as long as distance from snout to origin of
caudal. Clupeidae in part (Dussumierinae,
pp. 262-274), Part 3, Key pp. 259—-262.%
See also Key C, ra. 38. For those with adipose fin, see Key B, 13b.
Sometimes included among the Albulidae. It has not yet been reported for the western side of the Atlantic,
but it may be expected there because it is plentiful at moderate depths along the coast of tropical West Africa.
For details and description, see Poll (36: 16-25, 28, fig. 9).
See also Key A, 6a; Key C, rsa. 41. See also Key A, 13a. 42. See also Key A, 12a.
Fishes of the Western North Atlantic 99
13b. Origin of dorsal fin considerably posterior to mid-
length of trunk; pelvics below or anterior to dorsal ;
maximum depth of trunk only about 15°/, as great
as distance from gill opening to base of caudal;
head between 17—20°/, as long as distance from
snout to base of caudal.
Argentinidae in part (Microstomatinae), Part 4.43
Key B. A DORSAL ADIPOSE FIN PRESENT CLOSE IN FRONT OF CAUDAL; NO PHOTOPHORES
ON SIDES OF HEAD OR BODY.
1a. Well-developed rayed dorsal fin and adipose dorsal fin present.
2a. A narrow, posteriorly-pointing fleshy lobe close above base of pelvic fin on
each side.
3a. Teeth in jaws well developed, easily felt; 19 or more scales in a transverse
series from origin of dorsal to lateral line. Salmonidae, Part 3, p. 457.
3b. Teeth in jaws minute, if present; not more than 13 scales in a transverse
series from origin of dorsal to lateral line. Coregonidae, Part 3, p. 547.
2b. No fleshy lobe on side above base of pelvic fin.
4a. Some of upper pectoral rays longer than others and separate from lower
part of fin, thus forming a separate division.
Bathypteroidae in part, Part 5.44
4b. Pectorals not divided into an upper and lower division.
ga. Dorsal high, sail-like, its base extending from close behind gill
openings nearly to adipose fin. Alepisauridae, Part 5.
5b. Dorsal not high and sail-like, its base occupying not more than
about half the distance between level of gill openings and adipose fin.
6a. Upper jaw not reaching rearward as far as front of eye.
7a. Mouth at least twice as long as eye; eye not more than
about 17°/, of hl (less than this in most); anal base at least
1.5 times longer than base of dorsal; teeth well developed,
long and fang-like in some. Paralepididae, Part 5.
7b. Mouth not longer than eye; eye 27—40°/, of hl; anal base
not appreciably longer than dorsal base; teeth minute, if
detectable at all.
8a. Lateral-line scales not extending out onto caudal fin;
adipose dorsal anterior to rear end of base of anal fin.
ga. Gill membranes separate; branchiostegal rays 5
or 6. Argentinidae in part (Argentininae),
Rarteanss
43. See also Key B, 8b and ga; Key D, 4b. 44. For those without dorsal adipose fin, see Key A, 5a.
45- See also Key D, 4b.
7
100 Memoir Sears Foundation for Marine Research
gb. Gill membranes broadly united; branchiostegal
rays 2. Bathylagidae, Part 4.4
8b. Lateral-line scales extending out onto caudal fin; adi-
pose dorsal fin over or posterior to rear end of base of
anal fin. Argentinidae in part (Nansenia,
Microstomatinae in part), Part 4.47
6b. Rear end of upper jaw reaching beyond front of eye.
10a. Some of jaw teeth large, fang-like, conspicuous.
ita. Eye about 33°/, or more of hl; origin of anal pos-
terior to origin of dorsal by a distance less than
Koo) totale Omosudidae, Part 5.
11b. Eye only about 15°/, of hl; origin of anal posterior
to origin of dorsal by a distance nearly equal to hl.
Evermannellidae in part, Part 5.48
10b. None of jaw teeth much enlarged or fang-like.
12a. Upper jaw extending rearward past eye for a dis-
tance at least twice the length of eye.‘®
13a. Rear end of base of dorsal anterior to origin of
anal by a distance at least 50°/, of hl.
Synodontidae, Part 5.
13b. Rear end of base of dorsal only slightly ante-
rior to origin of anal.
Bathysauridae in part, Part 5.5°
12b. Upper jaw extending rearward past eye for a dis-
tance no longer than length of eye, and relatively
shorter than this in most.
14a. Dorsal profile of head deeply concave close in
front of eye; insertion of pelvics anterior to
origin of rayed dorsal by a distance nearly or
quite as long as eye. Scopelosauridae, Part 5.
14b. Dorsal profile of head close in front of eyes only
weakly concave at most; insertion of pelvics
under or posterior to origin of rayed dorsal.
15a. Upper jaw extending rearward beyond
eye for a distance about as long as eye;
dorsal profile of head weakly concave;
body deepest about at gill opening.
Neoscopelidae in part
(Scopelengys), Part 5.51
46. See also Key A, 13b; Key B, 8b; Key D, 4b. 47. See also Key D, rb.
48. For species having eyes with pupil above center and capable of being directed upward, see Key D, 5b.
49. In small specimens of some species of T'rachinocephalus the upper jaw does not extend rearward as far as this.
50. For those without adipose fin, see Key A, 7b. 51. For those with photophores, see Key C, 14b.
Fishes of the Western North Atlantic IOI
15b. Upper jaw not extending rearward ap-
preciably beyond eye; dorsal profile of
head straight to weakly convex; body
deepest about at dorsal fin.
16a.
16b.
Origin of rayed dorsal nearer to
gill opening than to adipose dorsal
by a distance of slightly less to
considerably -more than head
length.
17a. Pupil round; entire posterior
edge of opercular flap formed
by subopercular bone.
Aulopidae, Part 5.
17b. Pupil keyhole-shaped; pos-
terior edge of opercular flap
formed by opercle above,
and by subopercle below.
Chlorophthalmidae, Part 5.
Origin of rayed dorsal almost or
quite as near to adipose dorsal as
to gill opening, and much nearer
to adipose dorsal in some.
Osmeridae, Part 3, p. 553.
1b. No rayed dorsal fin; only an adipose dorsal fin. Anotopteridae, Part 5.
Key C. PHoToPpHORES PRESENT ON HEAD, OR ON BODY, OR ON BOTH}; OR WITH EYES
ON TOP OF HEAD.
1a. Eyes without lenses, covered by frontal bones, and consisting of large and flat
plates that occupy more than half of dorsal surface of head.
Ipnopidae in part, Part 5.°?
1b. Eyes with lenses, not covered by the frontal bones and, if dorsally directed, oc-
cupying less than half of dorsal surface of head.
2a. One or more barbels on chin, usually long and fleshy but sometimes very
small.
3a. Dorsal fin mostly or wholly anterior to anal, its origin near midlength
of trunk.
4a. Dorsal base extending behind pelvics, its origin behind or only
slightly before pelvic base; first dorsal ray not prolonged.
52. For those without these organs, see Key A, 7a.
Astronesthidae, Part 4.
102 Memoir Sears Foundation for Marine Research
4b. Dorsal base completely in front of pelvic bases; first dorsal ray much
i y P yi
prolonged. Chauliodontidae, specimens up
to about 50 mm long,*? Part 4.
3b. Dorsal fin opposite anal, its origin far posterior to midlength of
trunk.
ga. Sides of body with scales, or marked by hexagonal pigment patterns
resembling scales. Stomiatidae, Part 4.
5b. Sides of body scaleless, either unmarked or with parallel vertical
lines, never with hexagonal areas.
6a. Dorsal fin with more than 50 rays and beginning well before
middle of body; base of each dorsal and anal ray flanked on
either side by a sharp bony spur that penetrates the skin; length
of trunk posterior to head more than 20 times greater than great-
est depth of body. Idiacanthidae in part (adults), Part 4.54
6b. Dorsal fin with 30 rays or less, its origin well behind middle
of body; no bony spurs flanking bases of dorsal and anal rays;
length of trunk posterior to head not more than 15 times greater
than greatest depth of body.
7a. Mouth with a membranous floor; isthmus attached to man-
dibular symphysis; dorsal corner of gill opening behind
posterior margin of eye. Melanostomiatidae, Part 4.
7b. Mouth without a floor; isthmus attached to mandibular
symphysis by a muscular cord; dorsal corner of gill opening
above or in advance of posterior margin of eye.
Malacosteidae in part, Part 4.5°
2b. Chin without fleshy barbel.
8a. Each shoulder with a tubular papilla open at the tip and connected
to a voluminous sac beneath skin. Searsiidae, Part 3, p. 254.
8b. Shoulders without tubular papilla or sac beneath skin.
ga. Dorsal fin preceded by the protruding, spine-like tips of from one
to several of the neural processes of a like number of vertebrae.
Sternoptychidae, Part 4.
gb. Dorsal fin not preceded by the protruding spine-like tips of the
neural processes of any vertebrae.
10a. Branchiostegal rays plate-like, conspicuously exposed, the up-
permost connected to subopercular and interopercular bones.
Bathylaconidae, Part 4.
10b. Branchiostegal rays not plate-like, the uppermost not connected
to subopercular and interopercular bones.
11a. Dorsal adipose fin present.
53. For larger specimens, see Key ©, 12a. 54. See also Key D, 1b.
55- See also Key C, 15b.
Fishes of the Western North Atlantic 103
12a. Teeth enormous, conspicuous even when mouth
is closed; first dorsal ray greatly prolonged.
Chauliodontidae in part, Part 4.56
12b. Teeth small to moderate in size; first dorsal ray
not greatly prolonged.
13a. Upper edge of mouth formed by both max-
illaries and premaxillaries; upper jaw teeth
extending outside lower jaw when mouth is
closed; photophores generally present on
branchiostegal membrane.
Gonostomatidae in part, Part 4.57
13b. Upper edge of mouth formed by premaxil-
laries only; upper jaw teeth contained within
mouth; no photophores on branchiostegal
membranes.
14a. Distance from pelvics to anal origin not
more than twice the length of eye.
Myctophidae in part, Part 5.°°
14b. Distance from pelvics to anal origin at
least 3 times longer than eye.
Neoscopelidae in part, Part 5.°°
t1b. No dorsal adipose fin.
15a. Body with at least 20 longitudinal rows of scales;
mouth not widely distensible; gape not extending
past eye; teeth small.
Alepocephalidae in part, Part 3, p. 250.%°
15b. Body without scales; mouth enormously disten-
sible; gape extending beyond eye by a distance
3 or more times the length of eye; teeth large and
conspicuous. Malacosteidae in part, Part 4.6
Key D. Eyes EITHER TUBULAR (TELEOSCOPIC), OR WITH PUPIL ABOVE CENTER AND CA-
PABLE OF BEING DIRECTED UPWARD, OR WITH EYE SITUATED AT THE TIP OF A
SLENDER STALK.
1a. Eyes not at tips of slender stalks.
56.
57:
58.
60.
2a. Lower lobe of caudal much longer than upper lobe; gill openings minute;
base of pectorals wholly above gill openings. Giganturidae, Part 4.
See also Key C, 4b.
Exceptions: in Ichthyococcus the upper jaw is formed by the maxillaries alone, and in Triplophos it is almost entirely
bordered by the premaxillaries.
See also Key D, rb. 59- See also Key B, 15a.
See also Key A, 6a; Key A, 11a. 61. See also Key C, 7b.
104 Memoir Sears Foundation for Marine Research
2b. Lower lobe of caudal about as long as upper lobe; gill openings broad; base
of pectorals not wholly above gill openings.
3a. Upper jaw falling far short of eye; teeth minute, if detectable at all.
4a. Either an adipose fin between rayed dorsal and caudal, or a dome-
shaped luminescent organ on eye tube. Opisthoproctidae, Part 4.
4b. No adipose fin, and no luminescent organ on eye tube.
Argentinidae in part (Xenxophthalmichthys,
Microstomatinae in part), Part 4.®
3b. Upper jaw reaching rearward beyond front of eye; teeth fang-like, con-
spicuous.
5a. Body strongly compressed sidewise, covered with scales; pectorals
with 19-21 rays, their line of insertion oblique, wholly above artic-
ulation of lower jaw with skull (quadrate bone).
Scopelarchidae, Part 5.
5b. Body not much compressed, without scales; pectorals with only 12
rays, their line of insertion nearly horizontal, close to ventral outline
of body, entirely below articulation of lower jaw.
Evermannellidae in part, Part 5.8%
1b. Eyes at tips of slender stalks.
Stalk-eyed larval stages of Bathylagidae in part (Part 4);°* of Idiacanthidae
in part (Part 4);8° of Myctophidae in part (Part 5);%* and probably others
of unknown parentage.
62. See also Key A, 13b; Key B, 8b; and Key B, ga. 63. See also Key B, 11b.
64. See also Key B, gb. 65. See also Key C, 6a.
66. See also Key C, r4a.
eS
ONC
TS:
16.
Tipe
zé8.
IQ.
20.
2I.
TEXT AND FOOTNOTE REFERENCES
Aareor, Ichthyol., 3, 1738.
Becse, Zoologica, N.Y., 16, 1933.
Beebe, Zoologica, N.Y., 16 (4), 1934.
Berg, Class. Fish., Trav. Inst. zool. Acad. Sci.
URSS, 5 (2), 1940; also reprinting by J. W.
Edwards, Ann Arbor, Mich., 1947 (Russ.
and Eng.).
Bertelsen, Nature, Lond., r8z, 1958.
Bolin, Copeia, 1939.
Boulenger, Cambr. Nat. Hist., 7, 1904.
Cuapman, Ann. Mag. nat. Hist., rr (9), 1942.
Cohen, Zoologica, N.Y., 41 (2), 1956.
Cohen, Bull. Fla. St. Mus., 3, 1958.
Cope, Trans. Amer. philos. Soc., 74, 1871.
Cuvier, Régne Anim., 2, 1817.
Cuvier and Valenciennes, Hist. Nat. Poiss., r9,
1846.
Cuvier and Valenciennes, Hist. Nat. Poiss., 27,
1848.
Cuvier and Valenciennes, Hist. Nat. Poiss., 22,
1849.
Fowzer, Bull. Amer. Mus. nat. Hist., 70 (1),
1936.
Gut, Mem. nat. Acad. Sci., 6, 1893.
Grassé, et a/., Trait. Zool., 13, 1958.
Ginther and Deckert, Zool. Beitr., Berl., N.S.
T (3), 1955.
Hay, Bull. U.S. geol. Survey., 779, 1902.
Hollister, Zoologica, N.Y., 2r (4), 1936.
10S
23.
24.
25.
38.
39-
Techs Stanf. Uniy. Publ. Biol., 3 (2), 1923.
Jordan and Evermann, Bull. U.S. nat. Mus.,
47 (1), 1896.
Kenpatt, Bull. U.S. Bur. Fish., 37, 1921.
Lanxester, Treat. Zool., 9, 1909.
Mansuazt, ‘Discovery’ Rep., 27, 1955.
Matsubara, Fish. Morph. Hierarch., Pt. 1,
1955 (Jap-).
Myers, Copeia, 1946.
Myers, Proc. Calif. Acad. Sci., (4) 33, 1944.
ING Hist. Fish., 1931; and subsequent
editions.
Norman, Synop. Orders, Families, Genera or
Recent Fish. (mimeo), 1947.
Nybelin, Ark. Zool., N.S. To (4-5), 1957-
Parr, Bull. Bingham oceanogr. Coll., 3 (3),
1927.
Parr, Copeia, 1930.
Parr, Copeia, 1948.
Poll, Res. Sci. Exped...Eaux Cétiéres Afr.
Atl. Sud, 4 (2): 1953.
Racurg, Neue Schr. Naturf. Ges. Danzig, Z
(3), 1824.
Regan, Ann. Mag. nat. Hist., (8) 3, 1909.
Regan Ann. Mag. nat. Hist., (8) 5, 1910.
106
40. Regan, Trans. roy. Soc. Edinb., 49 (2), 1913.
41. Regan, Encyc. Britt., 9, 1929, and subsequent
editions.
42. Regan and Trewavas, Dana Rep., 5, 1929.
43. Regan and Trewavas, Dana Rep., 6, 1930.
44. Ridewood, Proc. zool. Soc. Lond., 2, 1904.
45. Romer, Vert. Paleont., ed. 2, 1945.
46. Saint Seine, Nouv. Arch. Mus. Hist. nat.
Lyon, 2, 1949.
47. Schultz and Stern, Ways of Fish., Pt. 1, 1948.
48. Senior, Biol. Bull. Woods Hole, 12, 1900.
49. Senior, Anat. Rec., I, 1907.
50. Smitt, Hist. Scand. Fish., 2, 1895.
5r.
52.
53:
54-
55:
Memoir Sears Foundation for Marine Research
Tcuernavin, Nature, Lond., 757, 1946.
Tchernavin, Feed. Mech. Deep Sea Fish.,
1953-
Trewavas, Proc. zool. Soc. Lond., 1933.
Woopwaro, Cat. Foss. Fish. Brit. Mus., 3,
1895.
Woodward, Cat. Foss. Fish. Brit. Mus., 4,
1gOl.
Zucmayer, Bull. Inst. océanogr. Monaco,
253; 1913.
Suborder Elopoidea’
COMPOSITE AUTHORSHIP
Characters and Key to Families
HENRY B..BIGELOW=2
Museum of Comparative Zoology
Harvard University
Characters. Silvery, herring-like isospondylous fishes with the ParasPpHENOoID
BONE On posterior part of roof of mouth thickly set with low Trrrn,? extending
nearly to posterior end of basioccipital or beyond, without lateral processes for articula-
tion with the mesopterygoid. TeEMporaAL FORAMINA present. Parapopuyses not fused
with vertebral centra, but simply wedged in pits. ProrruMorps not paired (see Eso-
coidea, Part 4). Pecrorat GIRDLE with mesocoracoid element. A Bony (GULAR) PLATE in
chin region between branches of lower jaw in some (Elopidae, including the Megalo-
pidae of Matsubara) but not in others (Albulidae, Pterothrissidae). Luminescent
oRGANS absent. Apipose fin lacking between rayed dorsal and caudal. Prtvic fins
abdominal, standing from about 30 °/, to about 45°/, of distance rearward from gill slit
toward lower origin of caudal. Anat fin originating from about 50°/, to about 68°/, of
distance between pelvics and caudal. Caupat deeply forked. REaRMosT DORSAL RAY
greatly prolonged in some (Tarpon, Megalops); the RrarMost ANAL Ray prolonged as
well in a few (Dixonina); none of the other fin rays prolonged. A single large, nar-
row, fleshy AxILLary scale above pectorals and pelvics. Scares thin, cycloid. Eyes
circular, of the ordinary type. Upper jaw extending rearward far beyond eye in most,
1. With families Elopidae and Albulidae by Samuel F. Hildebrand, pp. 111 and 132, respectively.
2. The assistance of Dr. W. A. Gosline is gratefully acknowledged.
3. Ridewood, who described their skulls in detail, has recorded this peculiar dentition (3: 39, for Elops; 44, for Mega-
lops; 40, for Albula; 53, for Pterothrissus). These observations have been verified for Pterothrissus by Poll (2: 222,
fig. 7), and for Tarpon by original examination.
107
108 Memoir Sears Foundation jor Marine Research
and tip of Lower jaw projecting as far as tip of upper jaw or slightly beyond it; upper
jaw in others (d/bula, Dixonina, Pterothrissus) falling far short of front of eye, and tip
of snout overhanging mouth. Maxittaries forming part of margin of upper jaw in
most but excluded from gape in a few (4/bula, Pterothrissus). BRANCHIOSTEGAL RAYS
slender, rod-like, not forming a part of opercular system. One to three lateral sensory
CANAL-BEARING BONELETS on either side of snout in front of lacrimals. Some with two
rows of valves in cone of Heart (A/bula, Pterothrissus, and Megalops, but not Elops).
There is a ribbon-like (leptocephalus-like) larval stage.
Families. Three families are recognized here within the Suborder as defined
above: Elopidae, Albulidae, and Pterothrissidae, the last of which is extralimital and
is included only in the Key to Families.
Fossil History. The Elopidae are known from as far back in geologic time as the
Lower Cretaceous, the Albulidae (including the Pterothrissidae) from as far back as
the Upper Cretaceous.
Key to Families
1a. A bony (gular) plate in chin region between branches of lower jaw; upper jaw
extending rearward considerably beyond eye; tip of lower jaw projecting at least
as far as tip of upper jaw, or a little farther; 23-35 branchiostegal rays.
Elopidae (including Megalopidae of Matsubara), p. 111.
1b. No gular plate; upper jaw falling short of front of eye; tip of snout overhanging
mouth; only about 11-15 branchiostegals.
2a. Base of dorsal occupying only about 25°/) of distance between gill opening
and base of caudal. Albulidae, p. 132.
2b. Base of dorsal occupying about 80°/, of distance between gill opening and
base of caudal. Pterothrissidae.
Plentiful off tropical West Africa
in depths of 100-250 m and re-
ported down to s500m;? also
Japan.
4. According to Cadenat (r: 365) and Poll (2: 23). Not yet reported for the western Atlantic but may be expected
there.
TEXT AND FOOTNOTE REFERENCES
I. Capenar, Bull. Mus. Hist. nat. Paris, (2) ro, Be Pou, Res. Sci Exped.... Eaux Cétiéres Afr.
1938. Atl. Sud, 4 (2), 1953.
Sh Ripewoon, Proc. zool. Soc. Lond., 1904.
109
IIo Memoir Sears Foundation for Marine Research
Editorial Comments on the Hildebrand Manuscripts
The following manuscripts by Dr. Hildebrand had not been worked on by him
for some years prior to his death in 1949. The Sears Foundation is therefore grateful
to Dr. George S. Myers for having undertaken the task of bringing some manuscripts
up-to-date in such ways as he thought necessary, and to the U.S.Fish and Wildlife
Service for having made this possible through the kind offices of Dr. L.A. Walford.
Dr. Myer’s work, which was both time-consuming and extensive, was confined prin-
cipally to the insertion of recently described genera and species, the rewriting of certain
keys, and research on the classification or nomenclature of certain genera (notably
Pristigaster and Ilisha) which required changes in the names originally used by Dr.
Hildebrand. Revisions that departed radically from what he believed Dr. Hildebrand
would have included have been inserted as initialed footnotes.
Some years ago Dr. John Tee-Van, because of the pressure of administrative duties
at the New York Zoological Society, was no longer able to continue as Editor-in-Chief;
Dr. Bigelow was subsequently chosen by the Editorial Board as his successor. Upon
receipt of the Hildebrand manuscripts from Dr. Myers, Dr. Bigelow made extensive
revisions and additions in many of the accounts, mainly in the sections dealing with life
history. Thus Dr. Bigelow added much information to that originally contributed by
Dr. Hildebrand; his major revisions occur primarily in the sections dealing with the
Elopidae (p. 111), Albulidae (p. 132), and Clupeidae (p. 257). The taxonomy remains
essentially as it was written by Dr. Hildebrand or revised by Dr. Myers, excepting
minor changes for conformance with general format. The value of Dr. Bigelow’s con-
tributions to the Hildebrand papers, and indeed to Part 3 in many other respects as
well, can be measured only by the usefulness of this volume to those who will have
occasion to refer to it. I have made such emendations as seemed desirable for publi-
cation.
Explanations of Dr. Hildebrand’s procedure of measuring and counting are given
on pages 154, 258, 343. Dr. Hildebrand’s taxonomic accounts are as complete as the
material and original sources of information permitted. In most instances they are
based upon his study of extensive material available to him at the U.S. National
Museum and elsewhere, and perhaps equally important, upon his broad and exten-
sive knowledge acquired over the years from firsthand observations in both field and
laboratory. Whether one agrees or disagrees with his treatment of relationships and
such, there can be no question of the care and meticulousness of his basic work.—
Y. H. Olsen.
Family Elopidae’
SAMUEL F. HILDEBRAND?
Characters. Bovy slender, more or less compressed laterally; belly rounded; ordi-
nary scales covering median line. Eye large, with adipose tissue in large examples.
Mourn large, terminal, or superior. Maxittaries large, in about three pieces, ex-
tending far beyond eye in adults. PReMaxiLLarres not protractile. Lower jaw with
an elongate bony plate between its branches. Treru all small, often bluntly villiform;
teeth present on jaws, vomer, palatines, pterygoids, tongue, and basibranchials. Bran-
CHIOSTEGALS about 23-35. GILL MEMBRANES separate, free from isthmus. OPERCULAR
BONES with membranous borders. Git RAKERS moderately long. Last several segments
of SprnAL coLuMN directed upward toward base of upper lobe of caudal. LaTrrat
LINE present. Scaes large or small, with membranous borders; missing on head; the
median row in front of dorsal neither enlarged nor modified. Dorsat fin over or
somewhat behind pelvics. Caupat forked. Pecrorats and Petvics similar, each with
a rather long axillary scale.
Remarks. The members of this family are characterized by the presence of the
bony gular plate between the arms of the lower jaw, the numerous branchiostegals
(23-35), and the large terminal or superior mouth.
The genera Tarpon and Elops (pp. 112, 123) sometimes have been assigned to
two families, Megalopidae and Elopidae. However, the two genera agree in so many
seemingly basic characters—the gular plate, numerous branchiostegals, the dentition,
the large maxillaries, and the primitive character of the caudal skeleton—that it seems
desirable to treat them as one family.
The members of the Elopidae generally are considered to be among the most
archaic of existing teleosts. Their antiquity, particularly as indicated by their skulls,
has been fully discussed (ftn. 3).
1. Edited and emended by George S. Myers, Henry B. Bigelow, and Yngve H. Olsen.
2. August 15, 1883—March 16, 1949.
112 Memoir Sears Foundation for Marine Research
The young pass through a leptocephalus stage, similar to that of the eels.
Range. The members of this family are widely distributed in the warmer seas.
Key to Western Atlantic Genera
1a. Body moderately deep, rather strongly compressed laterally; mouth superior, with
tip of mandible entering dorsal profile; last ray of dorsal and anal produced, fila-
mentous; scales large, about 41-48 in a lateral series; anal with about 22-265 rays.
Tarpon Jordan and Evermann 1896, below.
1b. Body very slender, not greatly compressed laterally; mouth terminal or nearly
so; last ray of dorsal and anal not produced into a filament; scales small,
generally more than roo in a lateral series in American species; anal with fewer
than 20 rays. Elops Linnaeus 1766, p. 123.
Genus Tarpon Jordan and Evermann 1896
Tarpon
Tarpon Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 409; type species by original designation,
Megalops atlanticus Cuvier and Valenciennes, Hist. Nat. Poiss., 79, 1846: 398.
Generic Synonyms:
Clupea Bloch, Naturg. ausland. Fische, Pt. 9, 1794: 32; in part, for C. cyprinoides Bloch; Shaw, Genl. Zool.,
5 (1), 1804: 173, for C. gigantea Shaw.
Megalops Lacépéde, Hist. Nat. Poiss., 5, 1803: 289; in part, for M. giganteus Voigt ix Cuvier, Das Thierreich
(German transl.), 2, 1832: 423.
Characters. Bopy oblong, rather strongly compressed. VERTEBRAE about 55.
PsEUDOBRANCHIAE undeveloped. BrancuiosTEGALs about 23. Mouru quite oblique,
superior, with prominently projecting mandible. Larrrat tine decurved, with branched
tubes. Scaues large, not especially primitive, their characters similar to those of E/ops
(cf. Cockerell, 77: 122); scales extending somewhat onto base of anal and caudal but
not onto dorsal. Dorsat fin high anteriorly, its last ray produced and filamentous. ANAL
similar to dorsal but longer, its last ray produced in large examples, its origin not far
behind dorsal base. Petvics inserted well in advance of dorsal fin. Last several Verre-
BRAE Clearly out of line with main axis, being directed toward the tip of upper lobe of
caudal. Arr BLADDER large and opening into gullet; within bladder a considerable
amount of cellular, lung-like tissue; two large and two small strands adhering to bladder
wall (bladder serving somewhat as an air-breathing organ, much as in gars [pp. 64,
67]; for description and figures of the modified bladder, see Babcock, r [1936]: 50).
ALIMENTARY CANAL much shorter than body; the stomach large, reaching far back
3. The skull has been fully described and figured by Gregory, who regarded it as a little less primitive than that of
Elops (30: 137-142, figs. 31-33). The characters of the tail of Tarpon and related genera have been described by
Regan (59: 354), and in more detail by Hollister (39: 264).
Fishes of the Western North Atlantic reer
into abdominal cavity as a sort of blind sac, the two orifices rather close together.
PyLoric caEcA numerous, short and branched, held together by connective tissue.
Remarks. This genus is close to Megalops Lacépéde, the oxeye herring represented
by M. cyprinoides of the Indian Ocean and adjacent Pacific, but it differs from Megalops
mainly in the more anterior position of the pelvic fins—well in advance of the dorsal
fin in Tarpon but inserted under the base of the dorsal in Mega/ops. Thus the dif-
ference is in the position of the pelvic fins rather than in the dorsal as stated in the
original description of Tarpon. Hollister, who compared specimens, has stated that
Tarpon has only 57 vertebrae whereas Megalops has 68, and that Tarpon has 12-15
dorsal rays whereas Megalops has 19-21 (42: 449). Also, Tarpon grows to a much
larger size. American ichthyologists generally have accepted Tarpon as a valid genus
while Old World students have rejected its separation from Mega/ops. Perhaps in con-
sideration of the additional differences now known, the separation may seem better
founded. Compared to E/ops, Tarpon’s body is not as slender, its scales are larger, and
its dorsal and anal fins have long filaments attached to the last ray of each fin.
Species and Range. A single species is known, ranging from Cape Cod, Massa-
chusetts, to southern Brazil. It has been reported also for tropical West Africa.
Tarpon atlanticus (Cuvier and Valenciennes) 1846
Tarpon, Tarpum, Sabalo Real, Cuffum, Silverfish, Silverking, Grand Ecaille,
Jewfish, Palika
Figure 19
Study Material. A total of 25 specimens: 14, 78-1,175 mm TL (57-837 mm SL),
measured in the laboratory, from Boggy and Boca Grande, Florida; Fajardo, Puerto
Rico; ‘Cuba’; “West Indies”; and ‘Amazon River,” Brazil; 11 adults, 1too—
1,990 mm TL, from Gatun and Miraflores locks, Canal Zone, examined for food, sex,
and spawning condition.
Distinctive Characters. See Remarks, above.
Description. Proportional dimensions in per cent of standard length, and counts,
based on at least 14 specimens in the Study Material unless stated otherwise, spec-
imens 57-837 mm SL.
Body: depth 23.5-29. Scales: 41-48 series.
Head: length 25-31. Gill rakers: 19-21+ 36-40.
Snout: length 4.5—-6.2. Fin rays: dorsal 13-15;
Eye: diameter 5.3-9.5. anal 22—25; pectoral 13 or 14.
Maxillary: length 15.5—-18. Vertebrae: §3-57 (4 specimens).
Anal fin: length of base 18-20.
Pectoral fin: length 19-22.8.
114 Memoar Sears Foundation for Marine Research
Bopy with almost vertical sides; the dorsal outline of head nearly straight and
horizontal, the back somewhat elevated, with ventral outline strongly curved anteriorly ;
depth 3.4-4.3 in SL. Scares large, firm, with crenulate membranous border. LaTERAL
LINE complete, decurved anteriorly, the pores branched. Heap moderately short and
deep, its depth at middle of eye not quite twice its width at the same place, 3.2-4.7
in SL. SNour to rim of upper jaw considerably shorter than eye in small specimens,
gradually becoming equal to and finally longer than eye during growth, 4.5—5.5 in
head. Eve 3.3—-4.7 in head, much nearer to dorsal than to ventral profile, wholly in
anterior half of head in adults. Git: raKers rather long and slender, scarcely dentic-
ef
a Pond
SS q
<—
Ficure 19. Tarpon atlanticus, from off New Jersey, USNM 14924; from Goode. Drawn by H. L. Todd.
ulate. Mourn superior. Manpisxe projecting far beyond the gape, entering dorsal
profile in advance of mouth. Maxittary broad, strongly curved, extending under
posterior part of eye in young (80-100 mm) but far beyond eye in large examples,
1.5—-1.9 in head.
Dorsat fin high anteriorly, its last ray produced in specimens over 100 mm long,
quite long in large specimens, reaching to end of base of anal fin in some specimens
seen; its origin about equidistant between base of caudal and anterior margin of eye.
Caupat deeply forked, the lobes of about equal length, generally somewhat longer
than head. Anat somewhat elevated anteriorly, its last ray produced in adults, some-
times reaching the base of lower rays of caudal, its origin at least an eye’s diameter
behind end of dorsal base, its base 4.6—5.6 in SL. Pexvic rather large, inserted more
than an eye’s diameter in advance of origin of dorsal, and somewhat nearer to base of
pectoral than to origin of anal in large specimens. Pscrorat rather long, extending to
base of pelvic in small specimens, not nearly to pelvic in large ones, 1.2—1.4 in head.
AxILLary scaLe of pectoral rather small, only about a third of the fin length, 2.7—3.4
in head.
Color. Preserved specimens bluish silvery above, sides and lower parts bright to
pale silvery. Pectorals and pelvics pale; other fins all more or less dusky. Longley
Fishes of the Western North Atlantic I15
remarked, ‘“‘The large eye falls below the level of the dorsal green and its iris blends
with the silvery side. As seen by the diver this great fish appears unsubstantial, a gray
ghost floating in gray water” (49: 5).
Size. The Tarpon reaches a very large size; an individual a little over eight feet
having an estimated weight of 350 pounds has been reported for Hillsborough
River Inlet, Florida. A fish up to six feet in length, having a weight of perhaps more
than 100 pounds, is not very rare.
Development. The exceedingly large number of eggs produced by a Tarpon indi-
cates extreme prolificness. For example, a female 80 inches long, weighing 142 pounds
(caught and reported by Babcock, r [1936]: 41), contained 12,201,984 eggs according
to a close estimate made by John T. Nichols of the American Museum of Natural His-
tory. This great number far surpasses the classic example of the prolificness of the
large codfish often mentioned, which contained 1,839,581 eggs.
Ovarian eggs, taken from both unripe and spent fish, measured 0.6—-0.75 mm in
diameter, a small size for so large a fish. They sank slowly when placed in sea water,
suggesting that they may be demersal, not buoyant.
Tarpon eggs that have been spawned naturally and are positively identifiable as
such have not yet been described, nor have the early stages after hatching. But it
has been assumed from analogy with its smaller relative (Megalops cyprinoides) of the
East Indies that the Tarpon, like the ladyfish (E/ops saurus, p. 124) and the bonefish
(Albula vulpes, p. 134), passes through a ribbon-shaped leptocephalus stage during its
larval development. This assumption has been corroborated by the capture at Beaufort,
North Carolina, of a leptocephalus 20 mm long, near the transition stage; this, from
comparison with the corresponding growth stage in E/ops and A/bula (35: 45) and in
the East Indian Mega/ops from Java, probably was a Tarpon.* (One 18 mm long, re-
cently pictured and described by Gehringer, certainly was [27: 290].—H.B.B.)
The smallest fry definitely reported as Tarpon have ranged between 16 and 19.6
mm SL (32: 6, fig. 5, pl. 1; see also pl. 2 for fry up to 36-38 mm; for earlier ac-
counts of 37 and 77 mm fry, see 73: 229 and 63: 72). Breder’s painstaking study of
extensive series of various sizes from the western coast of Florida has shown that little
alteration takes place in the body proportions with subsequent growth.
Rate of Growth. Analyses of the number of checks on the scales of Tarpon from
the western coast of Florida, where it may be assumed that growth is interrupted during
the cold months, suggest that in open waters it averages about 3,050-3,060 mm in
length at one year, about 510-640 mm at two years, and about 1,270-1,525 mm at
three years. Earlier scale studies (z [1936]: 68) suggest that a Tarpon averages about
1,376 mm or four feet six inches when it is nine or ten years old, that a fish of 70 pounds
is about 12 years old, and a fish of 100 pounds is perhaps 13-16 years of age.
Tarpon kept in the old New York Aquarium grew more slowly than those re-
ported above, from less than 500 to only 1,220 mm in five years, perhaps as a result of
4. For an account of the leptocephalus stages of Elops and Albula, see pp. 125 and 136; for the East Indian Megalops,
see especially Chadabaram and Menon (z8: 756-759) and Hollister (42: 449).
8*
116 Memoir Sears Foundation for Marine Research
confinement (z3). Present indications are that a Tarpon does not ordinarily mature
sexually until it is about four feet long, i.e. until after the sixth or seventh winter
(13: 229-247).
Spawning. Ripe or nearly ripe Tarpon have been captured at Boca Grande, Florida,
June 5—22 (z [1936]: 41), and several fish containing large roe have been taken from
the Panama Canal Zone during February, March, and April (37: 25, 30, 35). These
observations indicate that at least some of them spawn there during the spring. Captures
of both green and spent fish have shown that “the time of spawning on the west coast
of Florida is from May to September” (z [1936]: 433; 13: 226).
The general belief that spawning takes place along shore, mostly in shoal water,
is borne out by Babcock’s report that ripe females as well as males ejecting milt were
caught near Boca Grande (z [1936]: 41, 43) and that he received an eyewitness account
of pairs of Tarpon spawning in Charlotte Harbor, near Fort Myers, Florida. The
presence of very young fish at points as widely distributed as Texas, Alabama, Florida,
the Bahamas, Puerto Rico, Haiti, Cuba, and Trinidad seems sufficient evidence that
the breeding range is extensive—along the shores of both continent and islands from
the Gulf of Mexico eastward through the West Indies and Caribbean Sea areas. To
what extent it breeds farther southward is not known. The well-known presence of
young as well as large Tarpon in Lake Nicaragua also makes it likely that this fish
spawns there in fresh water. No evidence, however, has been found that the Tarpon
breeds in the lakes of the Panama Canal (36: 240). It seems probable, therefore, that
the ripe fish there migrates to sea to spawn, using the locks of the Canal as a passageway.
Habits. The Tarpon does not school, at least not habitually like the menhaden, for
example. When it does congregate in considerable numbers, as at the foot of the spill-
ways of the dams in the Canal Zone, it does so presumably because food is abundant
there. In other places it sometimes congregates in pursuit of schools of fish. The fish
apparently is sensitive to cold water and will move away from it if possible. Even in
southern Florida it disappears from the inshore waters during cool weather (34: 211).
If it migrates out to sea, as it probably does at times to escape from cold water, it
seems highly probable that it remains at or near the surface where the water generally
is warmest and where it can utilize the modified air bladder for breathing. The Tar-
pon more or less habitually ascends freshwater streams, and in some places, in the
tropics at least, it occupies freshwater lakes more or less permanently (36: 2403 see
also Habitat).
The rolling movements of the Tarpon at the surface of the water are familiar to all
fishermen. As these movements reveal its presence, anglers watch for this activity.
Why the fish practices such movements habitually was not understood until it was dis-
covered that it possesses an air bladder supplied with a large amount of lung tissue
(36: 246). After the fish has rolled at the surface and descended, bubbles appear at
the surface, these presumably being air exhaled below the surface.
It has been suggested by several writers that the great leaps often executed by
the Tarpon seem to serve a different purpose from rolling movements: to escape from
Fishes of the Western North Atlantic 11g
enemies, to shake off remoras or irritating parasites, or to play. Also, it often executes
great leaps when hooked, evidently trying to free itself; as most Tarpon fishermen
know from experience, these efforts are frequently rewarded with success.
Habitat. The Tarpon, a coastal fish, is seldom if ever seen more than a few
miles out from the land. To be sure, it has been credited with extensive seasonal off-
shore migrations, even across the Gulf of Mexico (p. 118), but not with any definite
supporting evidence. The Tarpon may be taken in either salt or brackish water, and
not infrequently it lives in freshwater streams and lakes. It has long been known to
inhabit Lake Nicaragua, a body of water a hundred feet above sea level, and more
recently it has been reported as plentiful in the rapid San Juan River, via which the
Lake discharges into the Gulf of Mexico (z [1936]: 19-20). This fish is exceedingly
abundant, sometimes at least, in fresh or virtually fresh water at the base of the
spillway of Gatun Dam and also in Gatun Lake below Madden Dam. It occurs
regularly also in other parts of Gatun Lake and in the small freshwater lake be-
tween Pedro Miguel and Miraflores locks (36: 240). To reach these lakes the
fish must pass through the locks of the Canal. The smaller fish also abounds in
fresh and slightly brackish streams, as around southwestern Florida. The young fry
is often reported as occurring in great numbers in very stagnant brackish pools and
lagoons that were not connected to the sea when the collections were made (22: 80;
RMA 5G ASU 2 IGA) §
Migrations. Knowledge of this aspect of the Tarpon’s life is scanty. It is probable
that the great majority of the larval Tarpon passes through the metamorphosis close
to the land, and many do so in estuaries, such as the Grande Afiasco River, Puerto
Rico (21: 146), and the Indian River, Florida (32: 1). However, some drift so far off-
shore that they almost certainly are lost to the Tarpon population: one, for example,
was taken about 150 miles to the east of Brunswick, Georgia, well out in the Gulf
Stream (27: 236).
The reported scarcity of fish less than three feet or so off the open coast, con-
trasted with their abundance in estuarine situations, even in drainage ditches, suggests
that most of them tend at first to work inshore. Indeed, great numbers of small fry
have repeatedly been encountered in brackish pools in mangrove swamps, in landlocked
lagoons, and in small pools around Florida, Puerto Rico, Haiti, Jamaica, and on An-
dros Island in the Bahamas, access to which is possible only during periods of very
high water. Many of the fish that are trapped in this way are no doubt lost to the
population when the pools dry up. (For an interesting summary of recorded instances
of this sort, see Babcock, r [1936]: 24, 32-46.) But many of them probably find their
way out again. It is only in situations of this sort that the Tarpon fry has ever been
taken in any great numbers, at least around Florida (information from Dr. J. E. Ran-
dall). It is to be found in such situations around southwestern and southern Florida
throughout the year. The postlarval stages have recently been taken in such numbers
in the salt marshes of the Indian River on the eastern coast of Florida as to show that
this is an important center of dispersal for them (32: 9). Little Tarpon 1-1.5 feet
118 Memoir Sears Foundation for Marine Research
long are very plentiful in the narrow headwaters of brackish and freshwater streams,
and even in drainage ditches. But the growing fish tends to move down to the broader
waters as it grows.
There is no apparent reason to credit the older Tarpon with any extensive
migrations, whether inshore-offshore or along shore, in the southern part of the range,
for large fish are found there within rivers and along the open coast at all times
of the year.
In the northern part of its range, however, the large Tarpon drops out of sight
with the onset of autumn, not to reappear until the following March or later, according
to locality. It has often been credited with extensive seasonal migrations, “northward
along the Florida coast and down the Texan coast, and back across open waters”
(13: 234). But a more plausible explanation is that the autumnal disappearance of
the Tarpon from the open coasts of the northern-northeastern periphery of the Gulf
of Mexico represents nothing more extensive than a temporary withdrawal; it probably
moves far enough offshore and deep enough to escape winter chilling, such as killed
many of all sizes along the western coast of Florida in 1885, 1894-95, 1905, and 1935
(74: 123; I [1936]: 21; 66: 640). Appearance of the Tarpon off North Carolina in
summer and autumn (as happens more or less regularly), and even farther north
(p. 120), evidently represents so extensive a journey from the nearest wintering ground
available for a warm-water nonoceanic fish that it seems unlikely that any of those
ever find the way back to their southern home.—H.B.B.
Food. The Tarpon, so far as known, is wholly carnivorous. A specimen nearly six
feet long from Boca Grande, Florida, examined by me, had the remains of six marine
catfish in its stomach; and a 12-inch specimen from the same place had ingested a
silverside. The Tarpon has been reported as feeding almost exclusively on cutlassfish,
e.g. in Calcasieu Pass, Louisiana (73: 170-172). At times it gathers in great abun-
dance at the foot of the spillway of Gatun Dam in the Canal Zone and at the spillway
of Madden Dam in the upper part of Gatun Lake, presumably for the purpose of
feeding on small fish that are carried over the spillways. Various observers have
reported that it will follow schools of mullets, anchovies, and other fish, apparently
feeding on them at will. Young Tarpon, ranging from about two to eight inches in
length, taken in an isolated lagoon at Port-au-Prince, Haiti, had fed exclusively on
“aquatic hemiptera which filled the waters of the lagoon” (5: 36). Blue crabs con-
stitute a favorite bait in Florida.
Enemies. The chief natural enemy of the large Tarpon is undoubtedly the shark,
and it has been reported that the porpoise attacks it too (r [1936]: 64); the small
Tarpon quite certainly falls prey to many predators in the sea. The Tarpon’s chief
protection from enemies is apparently its great swimming speed, whereby it escapes.
Variation. Whether geographical variations exist can be determined only from
study of a larger number of specimens, collected from many localities, than have been
available to me.
Relation to Man. As a gamefish, the Tarpon scarcely is surpassed. Its reputation
Fishes of the Western North Atlantic 119
is world-wide and much has been written about the requirements for capturing and
subduing this powerful, hard-fighting fish. Its great strength became evident to the
writer one day when he was helping to land a Tarpon 80 inches long. A man standing
nearby saw the great fish leap ashore from the seine, and wishing to prevent it from
jumping back into the water, he threw himself on it, evidently believing that he had
enough strength and weight to manage the fish. However, with one stroke of the tail the
fish lifted the man, who weighed at least 150 pounds, and threw him a distance of about
12-16 feet. It is understandable, then, why considerable strength, skill, patience, and
determination are required to land a large Tarpon and why Tarpon fishing 1s exciting.
About angling for Tarpon, Miss Francesca LaMonte has provided the following
information (personal communication). The hard-fighting Tarpon, either large or
small, is a prized catch wherever it occurs in western Atlantic waters, from Nova
Scotia to Argentina, or in the eastern South Atlantic off Lagos, Nigeria, and at the mouth
of the Volta River, Ghana. The most famous western North Atlantic grounds for both
numbers of fish and facilities are: the Florida Keys, centering around Islamorada and
Marathon; the western coast of Florida, especially in the passes near Boca Grande and
Useppa Island; the Rio Panuco, México; and the Rio Encantado, Cuba.
The number of Tarpon caught is always small in comparison with the number
present, for the fish puts up a violent and often successful fight. A large one is usually
fished from outboard motor boats (with the motor cut) by casting, trolling, drifting,
or still fishing, on 12-pound line or less, with plugs or with either live or dead bait, and
with or without a drag on the reel. The first problem is to set the hook in the Tarpon’s
hard mouth, then to prevent the fish from throwing it or breaking the line, both of
which it will try to do. First it will make a fast, powerful run and a series of high, twisting
leaps, shaking its head with violence. Failing to throw the hook, it will then try to break
the line either by charging at the boat if the line is too tight, or, if too loose, taking off
into the mangroves where the line will eventually break on the roots. The small young
Tarpon is caught in rivers and canals on fly-fishing or spinning tackle with light mono-
filament line. Though small, it fights just as hard as the adult and is by no means
sport for an inexperienced angler.
There is wide diversity of opinion as to the edible qualities of Tarpon flesh. On
our own shores “large ones at least are not edible’ (Babcock, r [1936]: 63), but in
Panama it is highly esteemed by the natives and West Indian immigrants (36: 239);
there the size of the fish receives no consideration, for an 80-inch Tarpon seemed as
acceptable as a small one; as stated by me in the paper just cited, a small fishery for
Tarpon was in operation in Gatun Lake in 1935. It has been reported that it is highly
prized as food by the Indians of the Caladonia Bay (Panama) region (ro: 152). In
the United States the commercial value of the Tarpon is so slight that it has not
been listed in the statistical reports of the commercial fisheries by the former Bureau
of Fisheries or by the present U.S. Fish and Wildlife Service.
Range and Abundance. The regular range of the Tarpon extends from North
Carolina to central Brazil, and it is probable that the Tarpon frequents the Brazilian
120 Memor Sears Foundation for Marine Research
coast southward to Pernambuco, or even to Bahia,® though the most southerly
definite record is for Lake Papary, Rio Grande do Norte, in the general vicinity of
Natal (65: 8). Thence northward the range is evidently unbroken around the South
American coast, where it has been recorded for: Marajé Island at the mouth of the
Amazon (8: 298); French Guiana (57: 164); Trinidad, where it has been reported
as very plentiful (69: 26); and Lake Maracaibo, Venezuela, and its tributary Rio Con-
cha, where several were seen rolling and a school was reported (62: 33). It is equally
omnipresent in suitable situations all along the Central American littoral, including: the
Panama Canal to Gatun Lake; the San Juan River (p. 117); and Lake Nicaragua,
where it is so plentiful that a recent writer reported seeing “more than a dozen
rolling at the surface at one time” (52: 184). As every angler knows, the Tarpon
abounds near river mouths and in so-called ‘“‘passes” along the coasts of México and
Texas. Indeed, it is not unusual to find a Tarpon 80 or 100 miles up from the sea in
some Mexican rivers (13: 219). In fact, the Bay of Campeche, just west of Yucatan,
was the site of a more or less regular fishery for it as early as about 1675 (Capt. Wm.
Dampier, cited from Gowanloch, 29: 145-146). Proverbial centers of abundance are
the mouth of the Panuco River, near Tampico, México, and Aransas Pass, Texas.
The Tarpon, as Gowanloch (29: 154) expressed it, is to be found near almost any
coastal island or bay of Louisiana, Alabama, the western coast of Florida, and the
eastern coast of Florida, northward in summer about to the vicinity of Daytona. Espe-
cially renowned fishing grounds are: the Calcasieu River, some 30 miles east of the
Texas boundary; Grand Isle, Louisiana; Boca Grande and Captiva, western Florida;
the waterways and rivers among the Thousand Islands, and the interior waterways of
the Florida Keys (see also p. 119).
The Tarpon is widespread but seemingly not very abundant except perhaps locally
throughout the West Indian region in general. More than a century ago it had been
reported in scientific literature for Guadeloupe, Martinique, Santo Domingo, and
Jamaica (20: 399). It has been reported for Haiti (3: 143) and is common around
the Barbados and Puerto Rico. In some rivers of Cuba it is so abundant that a party
of anglers has reported hooking 105 large ones in a day in the Rio Encantado (33:
205). Among the Bahamas a few are seen, and an occasional one is taken off Bimini
(33: 207; also personal communication from Vladimir Walters); it is common around
Andros Island (rr: 66), and is taken or seen occasionally around Bermuda.
The Tarpon ranges in small numbers northward to North Carolina in some sum-
mers, and perhaps every summer; but large schools are sometimes seen as far north
as Cape Lookout. A five-foot fish has been taken off Hatteras, and the North Carolina
State Museum has one of 119 pounds from Beaufort, and another of 176 pounds from
Wilmington (64: 115). But it appears only as a straggler and at irregular intervals
farther north. It has been reported for the lower part of Chesapeake Bay by fisher-
men (38: 80); there are odd records of it for New Jersey, near Sandy Hook (23:
5. The first published account of the Tarpon, by Marcgrave (50), was probably based on examples from Recife (Per-
nambuco) or Salvador (Bahia).—G. S. Myers.
Fishes of the Western North Atlantic 121
264), for the eastern end of Long Island, New York (55: 33), and for Rhode Island
(68: 72). The Tarpon has been reported as being taken in small numbers almost
every year in the vicinity of Woods Hole, Massachusetts (67: 741). But the Tarpon
has been reported only once for the Gulf of Maine, when a specimen was taken at
Provincetown at the tip of Cape Cod on July 25, 1915 (6: 91). Northward from Cape
Cod there are only five records of it, all for Nova Scotia—one from Isaac Harbor
(31: 45), one from Harrigan Cove (70: 53), one of about three feet caught near Ter-
rence Bay, September 6, 1947, and two taken near Halifax on August 3-4, 1953
(information from Dr. A. H. Leim). The Tarpon is common on the coast of tropical
West Africa (25: 1543 43: 106; 1 [1951]: 18-19).
Synonyms and References:®
Clupea gigantea Shaw (in part), Genl. Zool., 5 (1), 1804: 173 (no type or type local. design., diagn., genl. acct.,
habitat).7
Megalops giganteus Voigt in Cuvier, Das Thierreich (German transl.), 2, 1832: 423 (refs.; cf. Amer. M. giganteus
and east. M. filamentosus Lacépéde); ‘Disciples’ edition” of Cuvier’s Régne Anim., 2, 1836: 559
(same as above with somewhat more extended descr.); Eng. ed., 10, 1834: 442 (design. Amer. species
as M. cyprinoides, with ref. to Bloch’s pl. 403; pelvics under anterior part of dorsal are as in Mega/ops
cyprinoides Broussenet as now understood, but 15 rays in dorsal suggest Tarpon).
Megalops atlanticus Cuvier and Valenciennes, Hist. Nat. Poiss., 19, 1846: 398 (orig. descr., type local. Guade-
loupe, Santo Domingo, Martinique, Puerto Rico; types in Paris); Poey, Repert. Fisico-Nat. Cuba, 2, 1868:
423 (diagn., brackish water of Cuban rivers, attains 5 feet); An. Soc. esp. Hist. Nat., 4, 1875: 146 (Cuba,
Jamaica, Puerto Rico, Santo Domingo, Martinique, Trinidad, México, British Guiana); iz Gundlach,
An. Soc. esp. Hist. Nat., 10, 1881: 343 (Puerto Rico, Cuba, Santo Domingo, Martinique, México);
Henshall, Bull. U.S. Fish Comm. (1889), 9, 1891: 385 (effect of cold water); Boulenger, Ann. Mag. nat.
Hist., (6) 20, 1897: 298 (Marajé I., Brazil); Gill, Smithson. misc. Coll., 48, 1905: 35 (descr., habitat,
game qualities, life hist., common names, etc.); Regan, Ann. Mag. nat. Hist., (8) 5, 1910: 354 (caudal
skel. cf. M. cyprinoides); Fowler, Proc. biol. Soc. Wash., 33, 1920: 147 (Cape May, Monmouth and
Ocean counties, New Jersey); Proc. Acad. nat. Sci. Philad., 80, 1929: 609 (Barnegat Inlet, Atlantic
Highlands, and Atlantic City, New Jersey); Copeia, 1931: 46 (sport fish, Port Aransas and Point Isabel,
Texas); Bull. Amer. Mus. nat. Hist., 70 (1), 1936: 154, fig. 61 (African recs.; descr. based on Amer.
specimens); Puyo, Bull. Soc. Hist. nat. Toulouse, 70, 1936: 65, 163 (diagn., habitat in French Guiana);
Cadenat, Rev. Trav. Off. Péches marit., 10, Fasc. 4, Liv. 4, Pt. 2, 1937: 441, figs. 5, 6 (refs., remarks);
Norman and Fraser, Giant Fishes, 1938: 93, fig. 34 (semipopular acct.); Puyo, Faune Emp. Frang.,
12, Poiss. Guijane Frang., 1949: 151, fig. 75 (descr., Fr. Guiana).
Megalops elongatus Girard, Proc. Acad. nat. Sci. Philad. (1858), 1859: 224 (orig. descr., type local. Long Island,
New York; type not preserved).
Megalops thrissoides Giinther (not of Bloch and Schneider, which probably is M. cyprinoides), Cat. Fish. Brit.
Mus., 7, 1868: 472 (diagn., Jamaica, Cuba, Trinidad, Brit. Guiana); Goode, Bull. U.S. nat. Mus., 5,
1876: 68 (rare in Bermuda); Yarrow, Proc. Acad. nat. Sci. Philad., 1877: 215 (Ft. Macon, North
Carolina); Steindachner, Denkschr. Akad. Wiss. Wien, 39, 1878: 69 (Magdalena R., Colombia); Jordan
6. The Tarpon has been discussed in so many popular books and periodicals that it does not seem practical to list all
references to them. Therefore, only some popular accounts are cited, and only those citations in scientific periodicals
that seem to contain substantial information. African records have been omitted except for references to Fowler
(25) and Cadenat (r4), whose publications contain important references to African literature treating the species.”
7. This name was based by Shaw upon Marcgrave (50), who dealt entirely with the Atlantic Tarpon, and upon Clupea
cyprinoides of Bloch, which is a composite of the Tarpon and the Pacific Megalops cyprinoides (Broussonet). Since
Shaw’s account is thus a composite, it devolved upon a subsequent revisor to assign the name gigantea to one of the
included species. This was done by Cuvier (r9: 324), who definitely assigned the name gigantea to the American
Atlantic species. It would thus appear that the correct name of the Tarpon should be Tarpon giganteus (Shaw),
but the name atlanticus is so well established that its conservation, under suspension of the International Rules,
would seem to be worthwhile.—G. S. Myers.
Po Memoir Sears Foundation for Marine Research
and Gilbert, Bull. U.S. nat. Mus., 76, 1883: 262 (descr., distrib.); Willcox, Bull. U.S. Fish Comm.
(1886), 6, 1887: 123 (effects of cold); Henshall, Bull. U.S. Fish Comm. (1894), 74, 1895: 211 (abund.
in Florida; food and game qualities; size; etc.).
Megalops — Gill and Bransford, Proc. Acad. nat. Sci. Philad., 29, 1877: 187 (Toro Rapids, below Lake
Nicaragua; eaten by natives).
Tarpon atlanticus Jordan and Evermann, Bull. U.S. nat. Mus., 47 (Z), 1896: 409 (descr., range, synon.); 47 (4),
1goo: fig. 177; Smith, Bull. U.S. Fish Comm. (1897), 77, 1898: 90 (Woods Hole, Massachusetts) ;
Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 54 (Florida); Linton, Bull. U.S. Fish
Comm. (1899), Ig, 1901: 437 (parasites); Evermann and Marsh, Bull. U.S. Fish Comm. (1900), 20 (1),
1902: 80, fig. ro (descr., small specimens from Puerto Rico); Gregg, Where to Catch Fish. E. Cst. Fla.,
1902: 31, fig. (descr., fishing instruct., nonedible or barely so); Eigenmann, Bull. U.S. Fish Comm.
(1902), 22, 1903: 222 (20-192 mm specimens, no descr., Pinar del Rio, Cuba); Bean, Bull. N.Y. St.
Mus., 60, Zool. 9, 1903: 177 (synon., descr., New York); Hargraves, Fish. Brit. Guiana, 1904: 4
(local name Cuffum; used but not appreciated as food; fresh and brackish water); Smith, N.C. geol. econ.
Surv., 2, 1907: 114, fig. 36 (synon., descr., feed., range, North Carolina); Meek, Field Mus. Publ.,
Zool., 7, 1907: 99, 111 (Lake Nicaragua); Fowler, Proc. Acad. nat. Sci. Philad. (1910), 62, 1911: 599
(New Jersey); Eigenmann, Mem. Carneg. Mus., 5, 1912: 444 (Georgetown, Brit. Guiana); Sumner,
Osburn, and Cole, Bull. U.S. Bur. Fish. (rgr1), 3 (2), 1913: 741 (Woods Hole, Massachusetts);
Cockerell, Bull. U.S. Bur. Fish. (1912), 32, 1913: 122 (scales descr.); Gudger, Proc. biol. Soc. Wash.,
26, 1913: 104 (North Carolina); Halkett, Check List Fish. Canada, 1913: 45 (Isaac’s Harbor, Nova
Scotia); Starks, Stanf. Univ. Publ., Univ. Ser., 1913: 8 (Lake Papary, Rio Grande do Norte, Brazil) ;
Fowler, Proc. Acad. nat. Sci. Philad., 67, 1915: 245 (various Florida locals.; color relative to habitat;
cleaning of “spawning beds”); also 522 (Trinidad, B.W.I.); Coker, Copeia, 1921: 25 (small ones,
Dauphin I., Alabama); Eigenmann, Mem. Carneg. Mus., 9, 1922: 179 (fresh water, Magdalena R.,
Colombia); Meek and Hildebrand, Field Mus. Publ. Zool., 75 (1), 1923: 174 (synon., descr., range,
Panama); Schroeder, Rep. U.S. Comm. Fish. (1923), Append. 12, 1924: 4 (commerc. val., Key West,
Florida); Bigelow and Welsh, Bull. U.S. Bur. Fish. (1924), 40 (1), 1925: 91, fig. 38 (descr., range, rare
in Gulf of Maine); Breder, Zoologica, N.Y., 4 (4), 1925: 140, 152 (numerous at spillway, Gatun Dam,
C. Z.; enters Caladonia Bay, Panama, in schools; highly prized as food by local Indians); Nichols and
Breder, Zoologica, N.Y. (1926), 9 (1), 1927: 33, fig. (distr., life hist., size); Beebe, Bull. N.Y. zool.
Soc., 30 (5), 1927: 141 (young adults from brackish lagoon, Haiti); Hildebrand and Schroeder, Bull.
U.S. Bur. Fish. (1927), 43 (1), 1928: 80, fig. 41 (synon., descr., range, Chesapeake Bay); Beebe and
Tee-Van, Zoologica, N.Y., Io (1), 1928: 33, fig. (descr., Port-au-Prince, Haiti); Whiton and Townsend,
Bull. N.Y. zool. Soc., 3Z, 1928: 170 (feed., food in Calcasieu R., Cameron Parish, Louisiana): Breder,
Field Bk. Mar. Fish. Atl. Cst., 1929: 59, fig. (diagn., distr., habits, food value, size); Jordan, Manual
Vert. Anim. NE U.S., ed. 13, 1929: 37 (diagn., range, size); Nichols, N.Y. Acad. Sci., 10 (2), 1929:
198, fig. (range, Puerto Rico; diagn., habits); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish.
(1928), 2, 1930: 29 (common names, range, synon.); Burr, Bull. Texas Fish Game Comm., No. 5,
1932: 36 (numbers at Port Aransas and Point Isabel, Texas); Beebe and Tee-Van, Field Bk. Shore Fish.
Bermuda, 1933: 33, fig. (descr., uncommon in Bermuda); Breder, Bull. N.Y. zool. Soc., 34, 1933:
65-67 (small ones from pond, Andros I., Bahamas; food); Gregory, Trans. Amer. philos. Soc., 23
(2), 1933: 137-142, figs. 31-34 (skull descr., cf. E/ops and prehist. relatives); Storey and Perry,
Science, 73 (2022), 1933: 284 (young adults, Sanibel I., Florida, and elsewhere); Gowanloch,
Bull. La. Conserv. Dep., 2, 1932: 8, 54-65 (genl. acct.); Bull. La. Conserv. Dep., 23, 1933: 144-
168 (account of 1932 enlarged); Breder, Zoologica, N.Y., 18, 1934: 58, 59 (fresh water, Andros
I., Bahamas); Hildebrand, Copeia, No. 1, 1934: 45 (larval Tarpon descr.); Beebe and Hollister, Zoolo-
gica, N.Y., 19 (6), 1935: 211 (from Union I., Grenadines, B.W.I.); Vladykov and McKenzie, Proc.
N.S. Inst. Sci., rg (1), 1935: 53, fig. 26 (two Nova Scotian recs.); Babcock, The Tarpon, ed. 4,
1936: 1-175, illustrated (good genl. acct. and much scientific data on anat., habitats, breed., respir.,
etc.); Hollister, Zoologica, N.Y., 24 (4), 1936: 263 (caudal skel.); and 24 (4), 1939: 449-475
(caudal skel. and other characters cf. Megalops cyprinoides); Miller, Forest and Stream, 47 (1), 1936:
32-33 (fishery in Rio Frio, tributary of L. Nicaragua); Gudger, Amer. Mus. Novit., 944, 1937: 1,
fig. 2 (albino descr.); Heilner, Salt Water Fishing, 1937: 195-210 (genl. acct.; abund. in Cuba
and elsewhere); Hildebrand, Sci. Month., 44, 1937: 239-248, 4 figs. (Panama Canal; locally prized as
Fishes of the Western North Atlantic 123
food by natives); Kaplan, Big Game Angler’s Paradise, 1937: 89-136 (genl. acct.); Storey, Ecology,
18, 1937: 12 (always hurt by freezes); Breder, Bull. N.Y. zool. Soc., 47, 1938: 24 (irreg. in New York
Harbor); 42, 1939: 154-155 (habitat of small ones at Sanibel I., Florida); Butsch, J. Barbados Mus.
hist. Soc., 7 (1), 1939: 18 (common locally); Heilner, Bk. of Fishes, Nat. Geogr. Soc., 1939: 202,
327-336 (color plate, genl. acct.); Hildebrand, Zoologica, N.Y., 24 (1), 1939: 19, 25, 3 35 (in locks
of Panama Canal; transit through Canal completed); Longley and Hildebrand, Publ. Carneg. Instn.
Wash., 535, 1941: 5 (feed.; descr., especially as seen by a diver; Tortugas, Florida) ; Breder, Zoologica,
N.Y., 27 (1), 1942: 1 (behavior and respir. in confinement, refs. to papers on same subject); Breder,
Zoologica, N.Y., 29, 1944: 217-252 (habitat, body proportions, growth, movements, refs.).
Genus E/ops Linnaeus 1766
Ladyfish
Elops Linnaeus, Syst. Nat., ed. 12, 1766: 518; type and only included species, E/ops saurus Linnaeus 1766.
Generic synonyms:
Argentina, in part, Linnaeus, Syst. Nat., ed. 12, 1766: 519, for 4. carolina Linnaeus.
Mugilomorus Lacépéde, Hist. Nat. Poiss., 5, 1803: 398; type and only included species, Mugilomorus anna-
carolina Lacépéde equals E/ops saurus Linnaeus.
Trachonotus Rafinesque, Anal. Nat. ..., 1815: 88; type species Mugilomorus anna-carolina Lacépéde; Tracho-
notus proposed as a substitute for Mugi/omorus Lacépéde.
Characters® Bopy slender. VERTEBRAE about 75. PsEUDOBRANCHIAE large. Bran-
cHIosTEGALs about 34. Mourn nearly horizontal and almost terminal. LaTgraL LINE
straight, with simple tubes. Scares small, not especially primitive (fully described by
Cockerell, 15: 3; 17: 122), forming a sheath on base of dorsal and anal fins. DorsaL
rather high anteriorly, the last ray not prolonged. Anat similar to dorsal but smaller,
placed far behind dorsal. Petvics inserted near vertical from origin of dorsal. Last several
segments of spinal column directed upward toward base of upper lobe of caudal fin.
Air BLADDER slender, with very transparent walls, occupying full length of ab-
dominal cavity, adhering to abdominal wall dorsally and to alimentary canal ventrally.
A.iMENTARY CANAL, below air bladder, consisting of a straight tube except for a large
elongated blind sac that projects forward to throat below, and parallel to, the main
canal; a mass of coeca (firmly bound together) attached to distal end of blind sac
and forming a close-fitting shield on right side of sac throughout its length; one
lobe of liver forming a hood over blind end of sac, and another lobe shielding left
side of sac.
Remarks. The gular plate distinguishes this genus from d/bula. The very slender
body, the small scales, and the absence of filaments attached to the last ray of the dorsal
and anal fins distinguish it from Tarpon.
Range. About seven species have been recognized, but only E/ops saurus comes
within the range of this work. The species are widely distributed in the warmer seas,
entering estuaries and tidal streams.
8. The characters of the skull have been discussed by Gregory (30: 138), who regarded it as rather more primitive
than that of the tarpon. The ganoid character of the caudal skeleton has been well described and correctly figured
by Hollister (39: 260-263).
124 Memoir Sears Foundation for Marine Research
Elops saurus Linnaeus 1766
Ladyfish,* Bigeye Herring, Tenpounder, Skipjack, Macabi, Matejuelo Blanco,
Matejuelo Real, Long John, Benane
Figures 20, 21
Study Material. Many specimens, including an almost complete growth series
ranging from larvae to adults 550 mm long.
Distinctive Characters. See Remarks, p. 123.
Ficure 20. Elops saurus; U.S. National Museum specimen. Drawn by H. L. Todd.
Description. Proportional dimensions in per cent of standard length, based on
study specimens.
Body: depth 14.5-19.3. Scales: 103-120.
Head: length 25-31. Branchiostegals: 26-32.
Snout: length 5.8—7.0. Gill rakers: 6-8+ 10-15 (excluding ru-
Eye: diameter 5.0—6.3. diments).
Maxillary: length 14.5-17.8. Fin rays: dorsal 21-25; anal 14-17;
Anal fin: length of base 9.8—-11.8. pectoral 16 or 17.
Pectoral fin: length 14-16. Vertebrae: 73-80 (10 specimens).
Bopy very slender, only moderately compressed, its depth 5.2—6.9 in SL. Scares
small, thin, with crenulate membranous border (fully described by Merriman, 57:
113). Larerat tine nearly straight, extending onto base of caudal. Heap low, flat
above, 3.25-4.0 in SL. Snour rather long, usually a little longer than eye, 4.1-4.75
in head. Eye near upper profile, wholly in anterior half of head, 4.55—5.2 in head.
InTERORBITAL flat, 5.7—8.4 in head. Gitt RaKERs about 67°/, of the eye, scarcely
denticulate, their entire surface rough, generally 1 or 2 rudiments in large specimens
g. In scientific literature, as well as in earlier popular books on American game fishes, the name Ladyfish was usually
applied to Albula vulpes (p. 134). But to the angler, who alone takes much interest in either of them, “Ladyfish”
means Elops while Albula is universally termed “Bonefish.”
Fishes of the Western North Atlantic 125
and 2 or 3 in small ones, on upper and lower limbs of first arch; the lower counts
given, for this reason, are invariably based on the smaller specimens. Mourn nearly
horizontal, almost terminal, the lower jaw slightly included. Maxriiary reaching
somewhere under posterior half of eye in young adults, far beyond eye in large exam-
ples, 1.7—2.0 in head.
Dorsat fin rather high anteriorly, its longest rays usually reaching tip of last ray
if deflexed, its outer margin rather deeply concave, its origin about equidistant between
base of caudal and anterior margin of eye. Caupat very deeply forked, the upper lobe
rather longer than the lower one, usually about as long as head. Anat moderately small,
far behind dorsal, its origin usually about equidistant between base of caudal and base
of pelvic, its base 2.4—2.9 in head. Petvic fin not much smaller than pectoral, inserted
under, or slightly in advance of, origin of dorsal, scarcely nearer to origin of anal
than to base of pectoral. Pecrorat fin rather broad, not falcate, reaching scarcely
halfway to pelvic in large examples, somewhat more than halfway in small ones, 1.75—
2.0 in head. AxILLaRy scALE of pectoral extending to about the midlength of fin,
3.0-3.9 in head.
Color. Preserved specimens uniform bluish gray above, silvery elsewhere; dorsal
and caudal more or less dusky; other fins pale.
A large fresh specimen bluish on back, with sides silvery; slightly yellowish below.
Dorsal and caudal dusky and yellowish; pelvics and pectorals yellowish with dusky
punctulations (38: 78).
Size. Although a maximum of 900 mm (36 in.) TL has been reported (45: 410),
the usual length probably is under 500 mm (20 in.). The largest individual seen by
me was 730 mm (610mm SL; 29.2 in.); this large fish occurred among hundreds of
adults stranded in the Gatun Locks of the Panama Canal in 1935 when the Locks were
drained. In my file is a record of another large one, 718 mm (28.75 in.), caught at
Punta Gorda, Florida. Although one of the common names is Tenpounder, weight
records indicate that this fish probably never weighs that much. A specimen 470 mm
long (18.8 in.) weighed a pound, another of 550 mm (22 in.) weighed 1.6 pounds,
and still another 625 mm (25 in.) weighed 2.7 pounds; 82 fish, ranging between 400—
600 mm (16-24 in.), with an average length of about 450 mm (18 in.), weighed
100 pounds.
Development. The eggs and the earliest stages of the leptocephalus are unknown.
Of the many leptocephalus larvae studied, the youngest ones, judging principally by
the development of the fins, are 34, 35, and 37mm long. Although the forked
caudal is well developed, the other fins are virtually undifferentiated; the alimentary
canal is very loosely attached to the abdomen, as in larval herrings and anchovies, and
the vent is less than a head-length from the caudal base.
Two leptocephali, 34.5 and 36.5 mm long, are slightly more advanced; in these
a thickening within the finfold where the bases of the dorsal and anal are developing
has taken place, though rays are not yet definitely formed. The head is strongly depres-
sed, the mouth is terminal and large, and the snout, viewed ventrally, is rather sharply
126 Memor Sears Foundation Jor Marine Research
triangular; these characters at once distinguish the leptocephalus of this species from
that of 4/bula vulpes. The myomeres are mostly quite distinct, 78 having been counted
in one of the specimens. A row of faint dark spots, which are much more distinct in
older leptocephali, is present on each side just above the alimentary canal.
Three leptocephali, 42, 42, and 44 mm long, are the longest larvae in the collec-
tions studied and may represent about the maximum length attained in the leptoceph-
alus stage. Here development has progressed somewhat beyond that of the specimens
described above, for the rays are evident now in the dorsal and anal fins, though an
Ficure 21. Elops saurus. apove, leptocephalus, 35 mm long; BELow, young fish, 20mm TL, recently trans-
formed.
accurate enumeration cannot be made. The anal fin is removed from the base of the
caudal by a distance shorter than the head, and the primitive membrane has remained
between the two fins. The pectoral fins are represented by tufts of membrane and
the pelvics merely by thickened places in the abdominal wall where their bases are
developing. Seventy-seven myomeres were counted in one of the specimens.
In a considerably older specimen, reduced in length to 27 mm, the body, though
still quite flat, has become much more substantial than in the younger stages described.
The dorsal and anal fins are sufficiently developed to permit an accurate enumeration
of the rays, and the anal fin is now well separated from the caudal. The primitive mem-
brane has disappeared behind the fin, though a short section of it remains in front.
The pectorals are rather well developed and show signs of rays. However, the pelvics
are undifferentiated.
A virtually young adult only about 16 mm long (not in good condition) represents
the maximum “‘shrinkage”’ in length during metamorphosis, among specimens studied.
Though the body has become rather robust, the head and especially the snout remain
depressed. The fins, except for the pelvics, are rather well developed. The intestinal
canal remains loosely attached to the abdomen as in younger fish, and the primitive
rows of dark dots on each side of the canal are distinct.
The state of development varies widely among the rather numerous young adults,
18 mm and upward. For example, a specimen scarcely 20 mm long is fully as well
Fishes of the Western North Atlantic 127,
developed as others around 30 mm, and another specimen about 30 mm long is just
as well developed as some between 35 and 40 mm. The small and exceptionally well
developed specimens have advanced especially far in color. A specimen scarcely 20 mm
long already has some of the silvery color of large adults and all the juvenile markings
are missing, whereas a 32-mm specimen among the retarded ones is very pale and still
retains the two rows of dark spots on the abdomen, as in the leptocephali. Even in
the most retarded specimens, the fins are all well developed at a length of 22 mm, and
the outline of the gular plate is visible under magnification. Scales begin to appear in
at least some specimens at a length of 50 mm, and the teeth in the jaws are definitely
in narrow bands. At a length of 60-65 mm, scalation generally is complete, and the
fish are very similar to full-grown adults.
The environment probably affects the development profoundly, for the small,
exceptionally advanced specimens described in the preceding paragraph were taken
in brackish pools at Key West, Florida, whereas the notably larger and more retarded
ones were taken at sea off Ocracoke and Beaufort inlets, North Carolina.
Spawning. There is relatively little information about the actual time and place of
spawning. However, in 1939 I reported (37: 25) that 20 individuals, selected at random
from among the many left stranded when the Gatun Locks (Panama Canal) were
drained in February 1935, consisted of seven ripe or nearly ripe males (56.5—-62 cm
TL), and 13 females (63.5—73 cm TL) with large roe, indicating that spawning time
in that vicinity was at hand. Although leptocephali of 4/bula have been taken in con-
siderable seining during the winter and spring months on the Atlantic coast of Panama,
no young of E. saurus have been reported. On the Pacific coast of Panama, leptocephali
of the closely related E. affinis were taken during February, August, and “autumn.”
In the collections at hand there are leptocephali from Texas, mostly from the vicinity
of Corpus Christi, collected in February, March, April, and November. Young adults
were taken in Aransas Pass, Texas, during June, and at Key West, Florida, in March
and November. Leptocephali are also at hand from the Florida Keys, taken in No-
vember, and from Cuba, caught during May. For Beaufort, North Carolina, there is
a note in my files that a nearly ripe female of 604 mm was taken on October 3, 1912;
leptocephali were collected there during January, February, March, April, May, Oc-
tober, November, and December; rather recently-transformed young adults were taken
in the same vicinity during March, May, June, July, and August. Most of the many
leptocephali studied were taken with tow nets on the bottom, at sea, in comparatively
shallow water, though several captures were made in estuaries. The young adults, ex-
cept for a few lots in the transition stage, were collected with seines in inside waters,
partly in brackish ponds and pools.
Habitat. This is an active little fish, often traveling in schools, frequently skipping
at the surface, and jumping when hooked. Its usual habitat is shallow salt and brackish
water. The young adult has been taken in brackish lagoons and in ponds and pools,
perhaps more or less recently and temporarily separated from the sea. In the Gatun
Locks (see above) it occurred in large numbers and was most numerous in the middle
E28 Memoir Sears Foundation for Marine Research
chamber of the three flights, where the water, at least part of the time, is nearly fresh.
No evidence of its entrance into the strictly fresh water of Gatun Lake was obtained;
it seems improbable, therefore, that it uses the locks as a passageway; more probably
the locks serve merely as a feeding ground. If it migrates to sea to spawn, as might be
supposed since its young pass through a leptocephalus stage, the location of its
spawning ground (or grounds) remains unknown.
Food. Its diet has not been thoroughly studied, but the short alimentary canal
and the well-armored mouth surely suggest a flesh diet. A dozen or so adults taken
in one catch at Beaufort, North Carolina, had fed exclusively on shrimp, and other
examples from the same vicinity had fed on them also (48: 352; 64: 116). It has
been reported that an individual from Haiti had eaten a shrimp and two small fish of
its own species (5: 33). A specimen taken in Connecticut had fed on two small fish,
probably Menidia (51: 113). According to Kaplan, the Ladyfish feeds on shrimp,
shiners, and squid (46: 91).
Parasites. The cestode parasite, Rhynchobothrium bulbifer Linton, has been found
in the viscera of the adult Ladyfish (48: 352).
Relationships. This species is closely related to E. affinis Regan of the Pacific coast
of tropical America, apparently differing only in the number of gill rakers: in 41 spec-
imens of E. saurus examined, 9 had 6 rakers, 16 had 7, and 16 had 8, exclusive of
rudiments, on the upper limb of the first arch; and in 88 specimens, 2 had ro rakers,
7 had 11, 13 had 12, 28 had 13, 35 had 14, and 3 had 15 rakers, exclusive of rudiments,
on the lower limb. In nine specimens of E. affinis examined, 4 had 10 and 5 had 11
rakers on the upper limb, and in 26 specimens, 2 had 16 rakers, 4 had 17, 6 had 18,
11 had 19, and 3 had 20 rakers, exclusive of rudiments, on the lower limb.
Elops saurus has been recorded for West Africa by Fowler (25: 155) and by
others, but Regan (58: 38) has held that African specimens were distinct on the basis
of fewer vertebrae—68 or 69 compared to 78 or 79 for American specimens—and
on the basis of fewer scales—g4—98 for African specimens and 102-118 for American
specimens. According to counts made by me, these differences seem to be constant.
Ten American specimens have 73, 74, 75, 75, 75, 77, 78, 79, 80, and 80 vertebrae,
and nine American leptocephali have 77, 78, 78, 78, 79, 80, 82, 82, and 82 myomeres,
whereas one adult from Ashantee, Africa, has only 67 vertebrae. Counts of scales in
22 American specimens resulted as follows: 1 specimen with 103, 2 with 105, 1 with
106, 2 with 107, 2 with 108, 1 with 109, 2 with 110, I with 111, 2 with 112, 1 with
113, I with 114, 1 with 115, 2 with 116, 1 with 118, and 2 with 120. The three
specimens from Ashantee, Africa, examined by me have 92, 97, and 100 scales.
Although the range of counts for American specimens is rather great, it does not
overlap that given by Regan for African specimens. On the basis of the above data
given by Regan and me, the African representatives, which Regan named Elops senega-
Jensis, may still be considered distinct, though synonomized with E. saurus by Fowler.
Variations. No geographical variation in the number of gill rakers has been dis-
covered, but there appears to be a slight increase in number with age, which seemingly
Fishes of the Western North Atlantic 129
may be attributed to the development of some of the rudiments into rakers as the fish
grows. Thus, in 29 specimens 150 mm SL and under, 5 had 5, 14 had 6, and ro had
7 rakers and 2 or 3 rudiments on the upper limb; among 20 specimens over 150 mm
SL, 2 had 6, 4 had 7, and 14 had 8 rakers and 1 or 2 rudiments. Similarly, among 35
specimens 150 mm SL and under, 5 had 10, 14 had 11, 11 had 12, and 5 had 13 rakers
and 2 or 3 rudiments on the lower limb; among 32 specimens over 150 mm, 4 had 12,
7 had 13, 18 had 14, and 3 had 15 rakers and 1 or 2 rudiments on the lower limb.
Relation to Man. For its size, the Ladyfish affords good sport to the light-tackle
angler, but it is virtually of no value as food, its meat being dry and bony. It is seen
from time to time in the market at Colén, Panama, where it is known as “‘bonyfish”’
to the West Indian Negro immigrants who speak English; however, the demand for
it is small. When the Gatun Locks were drained in 1935, hundreds of Ladyfish
became stranded, and it was estimated that more than 1,000 pounds perished. While
the employees and the native residents eagerly accepted tarpon, crevalle jack (Caranx
hippos), and other species, the Ladyfish was entirely neglected and had to be buried
(37: 25).
Range and Abundance. \ts range extends from Cape Cod to Brazil. The southern-
most locality for which it has been recorded seems to be Rio de Janeiro (60: 90;
26: 132). While E. sawrus is not numerous in the West Indies, it is widespread in
the Caribbean-Gulf of Mexico region, and is common along Atlantic Panama; the
Study Material includes three specimens from the Gulf of Venezuela. It is common
around Florida, especially in enclosed waters such as the Indian River and among the
islands off the southwestern coast. The Ladyfish also occurs around Bermuda, though
not in any great numbers. To the north, it is present irregularly off Beaufort, North
Carolina, and it has been reported as common in the Chesapeake Bay region; farther
to the north its appearances—always in autumn—are sporadic. Considerable numbers
were taken near Sandy Hook in October 1923; for example, 13 out of one trap on
the 19th (55: 33). It has been recorded for the eastern part of Long Island (Orient),
New York, from October 6 to November 4. Also, it has appeared in considerable
numbers at various localities in the Woods Hole region in some autumns (67: 741),
but there is no dependable record for a Ladyfish north of the elbow of Cape Cod—
there is no knowing whether one taken at Chatham, Massachusetts, in 1888 (7: 139)
was from Nantucket Sound or the Gulf of Maine shore of Cape Cod. Nor is there
any possibility of determining whether an “‘4/bula vulpes,” taken at Black’s Harbor
(Charlotte County, New Brunswick) and in poor condition when examined (3I: 45),
was actually a Bonefish or Ladyfish.
Synonyms and References:
Elops saurus Linnaeus, Syst. Nat., ed. 12, 1766: 518 (orig. descr.; type local. “Carolina”; type, a dried skin in
BMNH;; see Goode and Bean, Proc. U.S. nat. Mus. [1885], 8, 1886: 205); Bloch and Schneider,
Syst. Ichthyol., 1801: 430, pl. 82 (descr., refs.); DeKay, New York Fauna, Fishes (4), 1842: 267,
pl. 41, fig. 131 (descr., New York Harbor); Storer, Synop. Fish. N. Amer., 1846: 211 (descr., refs.);
Gronow, Cat. Fish., Gray ed., 1854: 168 (descr., refs., distr.); Poey, Repert. Fisico-Nat. Cuba, 2, 1867:
423 (diagn., Cuba); Giinther, Cat. Fish. Brit. Mus., 7, 1868: 470 (synon., descr., Cuba, Jamaica, St.
9
130
Memoir Sears Foundation for Marine Research
Croix, ‘South America”; also several African and oriental locals. that probably are not this species);
Yarrow, Proc. Acad. nat. Sci. Philad., 1877: 215 (Ft. Macon, North Carolina); Jordan and Gilbert
(part not of Linnaeus), Bull. U.S. nat. Mus., 76, 1883: 261 (diagn., range); Goode and Bean, Proc.
U.S. nat. Mus. (1885), 8, 1886: 205 (type, a dried skin cut in two pieces, in BMNH in Lin-
naean Soc. rooms); Willcox, Bull. U.S. Fish Comm. (1886), 6, 1887: 123 (killed by cold, Florida);
Henshall, Bull. U.S. Fish Comm. (1889), 9, 1891: 373 (leptocephali, Marco, Florida); B.A. Bean,
Proc. U.S. nat. Mus., 74, 1891: 93 (Cape Charles, Virginia); Henshall, Bull. U.S. Fish Comm. (1894),
T4, 1895: 211 (Key West and Tampa, Florida; not as common as 4/éu/a; no econ. import.); Jor-
dan and Evermann (part not of Linnaeus), Bull. U.S. nat. Mus., 47(Z), 1896: 410 (descr., Amer.
range includ. Pacif. cst., which has distinct species, £. affinis Regan); also 47 (3), 1898: 2806 (confusion
of species); also 47 (4), 1900: fig. 178; Jordan and Rutter, Proc. Acad. nat. Sci. Philad., 1897: 94
(Jamaica, “‘sometimes 20 pounds” evidentally an error); Smith, Bull. U.S. Fish Comm. (1897), 17,
1898: go (common in fall, Woods Hole, Massachusetts); Giinther, Proc. linn. Soc. Lond., 1899: 25
(type in Linnaean Soc. coll.); Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 54
(Florida); Evermann and Marsh, Bull. U.S. Fish Comm. (1900), 20 (1), 1902: 81, fig. 11 (descr.,
Puerto Rico); Bean, Bull. N.Y. St. Mus., 60, Zool. 9, 1903: 179 (refs., descr., New York); Schreiner
and Ribiero, Arch. Mus. nac. Rio de J., 12, 1903: 90 (Caravelas and Rio de Janeiro, Brazil; Woods Hole,
Mass.); Hargraves, Fish. Brit. Guiana, 1906: 6 (local name “Long John”); Gill, Smithson. misc. Coll.,
48, 1905: 34 (skull figured, range; E/ops cf. Mega/ops); Jordan and Thompson, Bull. U.S. Bur.
Fish. (1904), 24, 1905: 232 (Tortugas, Florida); Linton, Bull. U.S. Bur. Fish. (1904), 24, 1905: 352
(food, parasites); Bull. Mus. comp. Zool. Harv., 50, 1906: 156 (Gulf of Mexico at Progreso, Yucatan);
Smith, N.C. geol. econ. Surv., 2, 1907: 116, fig. 37 (refs., descr., range, North Carolina); Linton,
Proc. U.S. nat. Mus. (1907), 33, 1908: 86 (Bermuda); Regan, Ann. Mag. nat. Hist., (8) 3, 1909: 37
(distr. limited to Amer. Atlant. cst.; Pacif. cst. represent. E. affimis; Afric. represent. E. senegalensis);
Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 204 (Nantucket, Mass.; “South Carolina”; West Palm
Beach, Florida; Santo Domingo; doubtfully from Jamaica, and Rio de J., Brazil); Weymouth, Proc.
U.S. nat. Mus., 38, 1911: 136 (Cameron, Louisiana); Nichols, Bull. Amer. Mus. nat. Hist., 32, 1912:
181 (common in markets, Cuba); Sumner, Osburn, and Cole, Bull. U.S. Bur. Fish. (1911), 3z (2),
1913: 741 (Woods Hole area, Mass.); Starks, Stanf. Univ. Publ., Univ. Ser., 1913: 8 (coast of Brazil;
Lake Papary, Rio Grande do Norte); Metzelaar, Trop. Atlant. Visschen, 1919: 9 (Curacao, Dutch
W.1., unimport. as food); Fowler, Proc. biol. Soc. Wash., 33, 1920: 147 (Cape May, New Jersey);
Breder, Zoologica, N. Y., 2 (15), 1922: 336 (Sandy Hook Bay); Meek and Hildebrand, Field Mus.
Publ., Zool., 75 (1), 1923: 175 (synon., descr., range, Panama); Linton, Proc. U.S. nat. Mus., 64,
1924: 59 (parasites); Bigelow and Welsh, Bull. U.S. Bur. Fish. (1924), 40 (1), 1925: 90, ftn. (questions
Halkett’s [1913: 45] ident. of New Brunswick specimen): Breder, Zoologica, N.Y., 4 (4), 1925: 141
(common at Gatun Dam Spillway, Canal Zone); Nichols and Breder, Zoologica, N.Y. (1926), 9 (tr),
1927: 33, fig. (Sandy Hook, New Jersey; Orient and New York, N.Y.; Woods Hole, Mass.; a fall visitor) ;
Hildebrand and Schroeder, Bull. U.S. Bur. Fish. (1927), 43 (1), 1928: 79, fig. 40 (descr., range, Chesa-
peake Bay); Beebe and Tee-Van, Zoologica, N.Y., 10 (1), 1928: 32, fig. (diagn., range, “apparently
rather rare” at Port-au-Prince, Haiti); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928),
2, 1930: 39 (common names, range, synon.); Breder, Field Bk. Mar. Fish. Atl. Cst., 1929: 60, fig.
(diagn., range, size); Fowler, Proc. Acad. nat. Sci. Philad., 80, 1929: 609 (Atlantic City, New Jersey);
Jordan, Manual Vert. Anim. NE U.S., ed. 13, 1929: 36 (diagn., range); Nichols, N.Y. Acad. Sci., ro
(2), 1929: 119 (probably not uncommon in Puerto Rican waters; diagn.; excellent sport fish with light
tackle); Fowler, Copeia, No. 2, 1931: 46 (Corpus Christi, Texas; local name skipjack); also Proc. Acad.
nat. Sci. Philad., 83: 1931: 392, fig. 1 (Trinidad); Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda,
1933: 33, fig. (descr., rare in Bermuda); Fowler (part not £. saurus Linn.), Bull. Amer. Mus. nat. Hist.,
70 (1), 1936: 155 (synon., includ. Afric. refs. based on £. senegalensis Regan; descr. based in part on
Afric. specimen); Hollister, Zoologica, N.Y., 21 (4), 1936: 260-263, figs. (caudal skel.); Hildebrand,
Sci. Month., 44, 1937: 243 (abund. in Gatun Locks, Panama Canal, when drained); Kaplan, Big
Game Angler’s Paradise, 1937: 91 (size, food, called Ladyfish in Florida); Storey, Ecology, 18, 1937:
16 (often hurt by freezes); Breder, Bull. N.Y. zool. Soc., 47, 1938: 24 (irregular, New York Harbor);
Hildebrand, Zoologica, N.Y., 24 (1), 1939: 25 (Gatun Locks, Panama Canal, apparently feed. ground);
Merriman, Copeia, 1939: 113, figs. a—p (scales); Bigelow and Schroeder, Copeia, 1940: 139 (Chatham,
Cape Cod, Mass., northernmost definite rec.); Longley and Hildebrand, Publ. Carneg. Instn. Wash.,
Fishes of the Western North Atlantic 131
535, 1941: 4 (rare at Tortugas, Florida); Fowler, Arqu. Zool. estad. Sao Paulo, 3 (6), 1941: 132 (refs.);
Hildebrand, J. Wash. Acad. Sci., 33 (3), 1943: 90-94, 3 figs. (notes on affinity, anat., develop.).
Argentina carolina Linnaeus, Syst. Nat., ed 12, 1766: 519 (orig. descr.; type local. ““Carolina’’; type lost).
Mugilomorus anna-carolina Lacépéde, Hist. Nat. Poiss., 5, 1803: 398 (orig. descr.; type local. “South Carolina”;
type in Paris).
Elops inermis Mitchill, Rep. in part on Fish. New York, 1814: 14 (orig. descr.; type local. New York, specimen
secured in market; type lost); Mitchill, Trans. Lit. philos. Soc. N.Y., 1815: 445 (descr., New York).
Albula vulpes Gregg (not A. vulpes Linnaeus, but E/ops saurus), Where to Catch Fish E. Cst. Fla., 1902: 34
(descr., instruct. for angling, barely edible).
Doubtful Reference:
Elops saurus Halkett, Check List Fish. Canada, 1913: 45 (report. New Brunswick; ident. doubtful).
Negative References :1°
Elops saurus Gregg, Where to Catch Fish. E. Cst. Fla., 1902: 33, fig. (4/éula vulpes, not E. saurus); Bean,
Bahama Islands: Fishes, 1905: 297 (determined as small Fewkinsia /amprotaenia on re-exam. by present
writer; not £. saurus; USNM 53076).
to. To these may be added all mention of Elops saurus based on specimens not taken on the Atlantic coast of America,
of which there are many.
9
Family Albulidae”
SAMUEL F. HILDEBRAND”
Characters. Bopy slender, little compressed; belly rounded; ordinary scales on
median line. Heap rather low; flat above. Snourt conical, projecting far in advance of
mandible. Eye of moderate size, with much adipose tissue in adults. Moutu mod-
erately small, nearly horizontal. Maxittary generally reaching nearly to front of eye,
or to some point underneath it, but never beyond eye. PREMAXILLaRIES not protractile.
Gutar pate lacking. TreTH small, present in bands on jaws, vomer, palatines, ptery-
goids, basibranchials, and sphenoid. BRANCHIOSTEGALS 13 or 14. GILL MEMBRANES
separate, free from isthmus; a membrane across isthmus in front. OPERCULAR BONES
with membranous borders. PsEuDoBRANCHIAE well developed. GiLL RaKERs very short,
stout, tubercular in large examples. Last several segments of SpinaL coLuMN directed
upward toward base of upper lobe of caudal. Larerat LINE present, straight. ScaLes
of moderate size, more or less quadrate, with membranous border, 3 (rarely 4)
nearly parallel basal radii, and 4 or 5 coarse basal lobes (fully described by Cockerell,
I5: 3; 16: 865; 17: 122); scales extending onto the fins; a modified median row of
enlarged scales in front of dorsal fin; no scales on head. Dorsat fin beginning in ad-
vance of pelvics. Caupat forked. Anat very small, placed far behind dorsal. Pecrorats
and Petvics similar, each with an AXILLARY SCALE.
The young pass through a leptocephalus stage like the Elopidae and the eels.
Remarks. The members of this family differ from those of Elopidae principally
in the absence of a gular plate, and in having a conical snout as well as fewer bran-
11. Edited and emended by George S. Myers, Henry B. Bigelow, and Yngve H. Olsen.
12. August 15, 1883—March 16, 1949.
13. The heart of Albula is reminiscent of the ganoids in having a rudimentary conus arteriosus with two rows of
valves (9: 548). Several fossil albulids are known, from as far back as the Eocene (54: 84).—G.S.M.
Hae
Fishes of the Western North Atlantic 133
chiostegals (13 or 14). From members of the Clupeidae they differ in having a well-
developed lateral line and undeveloped gill rakers.
Range. Only two living genera, 4/bula and Dixonina, are recognized.1* The former
has a world-wide distribution in warmer seas, and the latter is known from both coasts
of tropical America.
Key to Western Atlantic Genera
1a. Last ray of dorsal and anal not prolonged; snout only moderately conical, pro-
jecting only about a third of its length beyond mandible; maxillary not reaching
eyes Albula Scopoli 1777, below.
1b. Last ray of dorsal and anal much prolonged; snout sharply conical, projecting
more than a third of its length beyond mandible; maxillary reaching nearly
to middle of eye. Dixonina Fowler 1911, p. 143.
Genus Albula Scopoli 1777
Bonefishes
Albula Scopoli, Introd. Hist. Nat., 1777: 450; no species named, but based on 4/4u/a (name not binomial;
followed by long description), in Gronow, Zoophyl., I, 1763: 102; type and only included species
Albula s.salmo albula Meuschen, in Gronow, 3, 1761, index (not paged), Mediterranean; binomial
name given to Gronow’s (1763) A/éula (Whitley, 72: 303) equals Esox vu/pes Linnaeus, Syst. Nat.,
MAIS Gus
Generic Synonyms:
Esox, in part, Linnaeus, Syst. Nat., I, 1758: 313, for E. eu/pes Linnaeus, Bahamas.
Argentina, in part, Forskal, Descr. Anim ..., 1775: 68, for 4. g/ossodonta Forskal, Arabia.
Amia, in part, Bloch and Schneider, Syst. Ichthyol., 1801: 457, for 4. immaculata Bloch and Schneider, South
America.
Synodus, in part, Bloch and Schneider, Syst. Ichthyol., 1801: 398, for 8. argenteus Bloch and Schneider, Tahiti;
Lacépéde, Hist. Nat. Poiss., 5, 1803: 32, pl. 8, fig. 2, for S. vu/pes Lacépéde.
C/upea, in part, Bloch and Schneider, Syst. Ichthyol., 1801: 427, for C. drasiliensis Bloch and Schneider,
Brazil; Lacépéde, Hist. Nat. Poiss., 5, 1803: 427, for C. macrocephala Lacépéde, Martinique.
Butyrinus Lacépéde, Hist. Nat. Poiss., 5, 1803: 45; type by monotypy, Butyrinus bananus Lacépéde equals
Esox vulpes Linnaeus.
Glossodus Cuvier, Mem. Mus. Hist. nat. Paris, r, 1815: 2323; genotype by monotypy, Argentina glossodonta
Forskal equals Esox vu/pes Linnaeus.
Engraulis Agassiz, L. in Spix and Agassiz, Selecta genera et species piscium...Brazil..., 1829: pl. 23, fig. 2,
for E. sericus Agassiz; pl. 24, fig. 2, for E. dahiensis Agassiz, Brazil.
Conorhynchus Gill, Proc. Acad. nat. Sci. Philad., Suppl., 1861: 55; genotype by monotypy, Butirinus vulpes
Storer equals 4/bula vulpes.
Characters. Dorsat and Anat fins with the last ray not produced. MaxiLiary
falling short of vertical from front border of eye. Snour projecting about a third of its
14. Dr. Hildebrand evidently excluded from the Albulidae the deep-water genus Pterothrissus, which is known from
the coasts of Japan and West Africa and which may eventually be found in the western North Atlantic. Prerothrtssus
differs widely from Albula and Dixonina in the very long, many-rayed dorsal fin.—G. S. Myers.
134 Memoir Sears Foundation for Marine Research
length beyond tip of lower jaw. CAUDAL SKELETON (39: 269-275) exhibiting some
ganoid characters, showing a relationship with that primitive group. VERTEBRAE: last
2 or 3 in 4. vulpes out of line with central axis, directed somewhat dorsally; massive
spines (neurals) attached to them and preceding segment, extending into base of upper
lobe of caudal. ALIMENTARY CANAL appearing as a straight tube in larvae, becoming
modified in the adult with development of a very large stomach that occupies fully
67°/, of the length of abdominal cavity if distended with food; stomach largely a
blind sac; the cardiac and pyloric orifices rather close together. About 13 Pytoric
corca, bound together, forming a sort of sheath over part of stomach. AIR BLADDER
very long, extending backward far beyond vent, its walls very thin, without lung
tissue (see tarpon) except for a pair of very small kidney-shaped cellular bodies
attached ventrally at about midlength.
Remarks. This genus differs from Dixonina chiefly in that the last ray of both
dorsal and anal is short and does not become prolonged in adults; see also Key, p. 133.
From E/ops it is distinguishable by the absence of a gular plate (p. 123).
Range. Only one modern species, 4/bula vulpes, is generally recognized; this
occurs in virtually all warm seas.
Albula vulpes (Linnaeus) 1758
Bonefish, Grubber, Bananafish
Figures 22, 23
Study Material. An almost complete growth series ranging from leptocephali with
virtually undeveloped dorsal and anal fins to adults 430 mm TL, from both Atlantic
and Pacific. As the growth series was not complete in the material from the Atlantic,
some specimens, especially the older stages of leptocephali and first stages of young
adults, from the Pearl Islands in Panama Bay, were used in describing the young.
Distinctive Characters. See Remarks, above.
Description. Proportional dimensions in per cent of standard length, and counts,
based on Atlantic adults only.
Body: depth at origin of dorsal 20-23 in
large specimens, 14-19 in young
adults.
Caudal peduncle: depth 6.9-8.9.
Head: length 27.5-32.
Snout: length 11.5—-13.6 in large speci-
mens, 8.5—10 in young adults.
Eye: diameter 5.5—8.0
Maxillary: length from tip of snout 9.1—
TQ ea73
Anal fin: \ength of base 5.1—-6.75.
Pectoral fin: length 15-19.
Scales: 65-71.
Gill rakers: 7 or 8+ 9 or 10.
Fin rays: dorsal 17 or 18, rarely 19;
anal 8 or 9; pectoral 16 or 17,
rarely 15.
Vertebrae: 72—74 (§ specimens).
Fishes of the Western North Atlantic gis
Bopy slender, rounder and less compressed in large specimens than in young
adults, its dorsal profile more convex than ventral profile, its depth at dorsal origin
4.35—4-9 in SL in large specimens, about 5.25—7.1 in the more slender, recently-meta-
morphosed young. Scates firm, with crenulate membranous edges. Heap low, espe-
cially in large specimens, flat above, its depth exceeding its width at middle of eye
by about diameter of pupil, 3.0-3.8 in SL. Snour rather long, conical, especially in
large specimens, projecting about a third of its length beyond mandible, 2.2-2.5 in
head, about 2.7—3.5 in young adults. Eye moderately small, 4.4—5.5 in head, its
center nearer to margin of opercle than to tip of snout. Maxiiiary not quite reaching to
Ficure 22. A/bula vulpes, from off Newport, Rhode Island; from Goode. Drawn by H. L. Todd.
eye in large specimens, but extending to, or slightly beyond,*anterior margin of eye in
young adults, 2.6-3.15 in head (measured from tip of snout).
Dorsat fin somewhat elevated anteriorly, its longest rays not reaching to tip of
posterior ray if deflexed, its origin a little nearer to tip of snout than to base of
caudal. Caupat deeply forked, the upper lobe somewhat longer than the lower. ANaL
very small, its origin notably nearer to base of caudal than to base of pelvic, its last
ray, though longer than the preceding one, not especially produced, its base 4.7—5.4
in head. Petvic somewhat smaller than pectoral, inserted under or slightly behind
middle of dorsal base. Pecrorat with rounded margin, reaching notably less than half-
way to pelvic, 1.7-2.0 in head. Axittary scaLe of pectoral about half as long as
fin, adherent to body.
Color. Large specimens bluish above; bright silvery on sides and below; dark
streaks between rows of scales, at least on dorsal half of side; dorsal and caudal with
dusky margins, frequently entirely pale in preserved specimens. Very young adults
(completely metamorphosed and about 30 mm) with a double series of dark spots on
the back, each just off the median line; these spots soon uniting to form about nine dark
crossbands on the back, extending down nearly or quite to lateral line; the third band
crossing back at origin of dorsal; next two situated posteriorly under base of dorsal;
136 Memoir Sears Foundation for Marine Research
bands persisting until a length of about 75 mm. The dark longitudinal streaks of the
adults appearing shortly before crossbands become obscure.
Size. Its maximum length is about 770 mm (31 in.) and its weight about 13 or 14
pounds.
Development.* The eggs and the earliest stages of the leptocephali remain unknown.
The youngest at hand, that is, the least developed larva, is apparently somewhat younger
than any yet described. It is 57 mm long ($1 mm SL), was collected at Tortugas, Flor-
ida, and is the only one of the many that has the dorsal and anal fins entirely unde-
veloped. The pectoral fins appear as mere tufts of membrane and the pelvics only as a
thickening of the body wall where the bases are developing. The caudal fin, however,
is well developed with segmented rays, is broadly forked, and is about as long as the
head. This larva is sharply compressed, though not more so than considerably older
ones; its depth increases gradually backward from the head to about the beginning of
the distal fourth of the body, where it is contained 11.8 times in SL. Myomeres are
prominent, except anteriorly and posteriorly, and about 66 may be counted. The head
is low and moderately broad and is contained 16.4 times in SL. The snout is conical
and projects slightly beyond the mandible, the maxillary is not definitely formed, the
slightly oblique gape extends under the anterior part of the eye, and the eye is scarcely
as long as the snout, being contained in the head 4.1 times. The intestinal tract appears
to be a straight tube, is rather loosely attached to the body, has a slight groove on each
side, and extends nearly to the base of the lower lobe of the caudal. This old preserved
specimen, collected in 1919, is now entirely without pigment. Several other specimens
from Tortugas and Cuba are only slightly more developed.
Although leptocephali 75-87 mm long have been reported (28: 40),!6 the largest
one now at hand among several hundred from many localities along the Atlantic and
Pacific coasts of America is just 70 mm long (57 mm SL). In this specimen, one of a
large collection from the Pearl Islands, Panama Bay, the dorsal and anal fins are devel-
oped (showing at least most of the rays), and the fulcra may be fairly accurately enu-
merated. The dorsal fin is placed over about the beginning of the distal fifth of the body.
The anal is scarcely separated from the base of the lower caudal lobe. The pelvic fins
are developed but do not have definite rays and are placed at about midbody length
without the head, a position retained throughout life. The pectoral fins are fairly long,
with definite indications of rays. The body is deepest somewhat in advance of the dorsal
fin, and its depth there is contained 9.25 times in SL; 69 myomeres were counted.
No prominent changes in the head region are evident. The head is contained 14.4
times in the SL. The snout and eye are of about equal length, being 3.6 in head. The
jaws possess minute teeth that project somewhat forward. Two almost continuous dotted
dark lines bound the intestinal tract from behind the pectoral fins to the vent, being
15. A detailed account of the development and distribution of young Albula, based on the DANA collections, has
recently been published by Alexander (DANA Rep., 53, 1961).
16. The fact that d/bw/a has a “‘leptocephalus” larva was first ascertained about 1896 by Prof. C. H. Gilbert of Stan-
ford University. It seems he never published an account himself, but a synopsis of his findings, with his figures of
the developmental stages, was made known by Jordan (44) and Gill (28: 43) — G. S. Myers.
Fishes of the Western North Atlantic E27
interrupted only at the point of insertion of the pelvics; and a vertical series of dark
dots is present on the caudal base. Living larvae of this size, and considerably older
ones as well, are as transparent as glass. They appear as shining objects in the net and
are easily overlooked, only the dark eyes being at all conspicuous.
Older larvae of 45 mm (39 mm SL), though still sharply compressed, are much
more substantial than younger ones at a length of 70 mm or so. The dorsal and anal
fins have moved forward considerably, the anal fin now being separated from the caudal
S
SS
Ficure 23. A/bula vulpes. Pearl Island, Pacific Panama, USNM 128391. a Leptocephalus stage, 68 mm long;
B older larva in transition stage, 76mm TL; c older fry 30 mm TL. Drawn by Nancy D. Patton.
by at least half the length of the head. The greatest body depth, at the origin of the
dorsal, is contained 7.6 times in SL. The myomeres total 70-72. The head remains
rather low and broad and is contained 6.0 times in the length; the snout projects beyond
the mandible more prominently and is now rather longer than the eye, 4.0 times in
head; and the maxillary is fairly well developed, extending well beyond the anterior
margin of the eye, 2.4 in head. The color remains as in the younger fish described,
except that a few dark chromatophores have appeared about the head, and a slightly
broken dark line has developed on both upper and lower lobes of the caudal.
Larvae 32 mm long (26 mm SL) are the smallest in the study collections that may
be classed as leptocephali. The body remains strongly compressed, though of course
more robust and more substantial than in the 45-mm fish described above, and it has
become more elongate, the depth at the dorsal origin being contained 8.4 times in SL.
The dorsal and anal fins have continued to move forward, but they still remain more
posteriorly placed than in the adult; the origin of the dorsal is somewhat nearer to the
base of the caudal than to the tip of the snout and is approximately over the pelvic
138 Memoir Sears Foundation for Marine Research
fins. The intestinal tract has become more fully invaginated and no longer remains
merely attached, except posteriorly, as in younger fish; the vent, instead of being sit-
uated at the origin of the anal, is now well in advance of it, having moved forward
rather more rapidly than the fin. The color markings remain about as in the younger
fish described, but additional chromatophores have appeared, principally at the occiput,
at the base of the dorsal, and on the dorsal.
The smallest specimen that may be classed as a young adult is 28 mm (21 mm SL),
and its length is the least of the many 4/bu/a at hand. This fish is quite a little longer
than the one that diminished from a leptocephalus to a young adult of only 20 mm,
in an aquarium (39: 269; 40: 109). The body remains more compressed in this
and other young adults than in large adults, and it is also more slender, the depth at
the dorsal origin 6.5 times in SL. The head is contained 3.45 times in SL and there-
fore does not differ in this proportion from large adults. The eye is rather small, 5.05
times in head, and is definitely shorter than the snout, which is contained 3.05 times
in the head and projects rather prominently beyond the mandible; the maxillary ex-
tends to the anterior margin of the pupil and is contained 2.9 times in head. The dorsal
and anal fins have continued to move forward, the origin of the dorsal now being equi-
distant between base of caudal and tip of snout; and the attachment of the pelvic fins is
now only a little in advance of the middle of the dorsal base. At this stage, and even in
ones a little older, the two dark lines along the sides of the intestinal tract in the lepto-
cephali persist, two rows of dark spots on the back (described elsewhere) have appeared,
there are indications of some elongate dark spots along the side, and some additional
pigment has appeared on the base of the anal and ventrally on the caudal peduncle.
Scales first appear in young adults about 35 mm long. At a length of 40-45 mm,
scalation generally is complete, except perhaps on the abdomen. At these stages the
lateral line also is well developed. The dorsal and anal fins are situated approximately
as in large adults; that is, the origin of the dorsal is somewhat nearer to the tip of the
snout than to the base of caudal, and the anal is separated from the caudal by a distance
somewhat greater than the length of the snout. The pelvic fins, which, from the time of
their appearance in the leptocephalus, are inserted a little nearer to the head than to
the base of caudal, retain that relative position and are now about under the middle
of the dorsal base. The pectorals are long but not falcate and reach more than halfway
to the pelvics. The vent has continued to move forward and is now in advance of the
anal by a distance nearly equal to the length of the snout.
At 40mm, pigmentation has become general and the nine crossbands on the
back, described elsewhere, are prominent. But dark longitudinal stripes do not appear
definitely until a length of about 75 mm is attained, at which size the crossbands have
become obscure.
Spawning. The spawning season and the spawning grounds remain unknown.
Most of the many larvae and young adults described herein were taken off the coast
of Panama during February and March, but their occurrence at other seasons is not
known, as no year-round collecting has been done there. Growth stages ranging from
Fishes of the Western North Atlantic 139
fairly young leptocephali of 70 mm TL to young adults 52 mm long were exceedingly
numerous among some of the Pacific islands in Panama Bay in March 1937 (especially
in the Pearl Islands). Somewhat similar stages, though less numerous, were taken
in February and March on the Atlantic coast of Panama. Other leptocephali at hand
were taken as follows: January 23 on the Pacific coast of Colombia, February 13 in
Puerto Rico, March 13 in the Virgin Islands, April 8 in Cuba, April 23 and 28 in
Haiti, May 19 in Cuba, June 30 at Beaufort, N.C., July 21 in Bermuda, and August 7
and November 25 at Tortugas, Florida. Beebe and Tee-Van (5: 37) have reported that
on 34 evenings of collecting (between February 26 and April 30) with an electric light
lowered from the gangway of a schooner in Port-au-Prince Bay, Haiti, larvae of 4/bula
were absent only two nights and were very abundant 17 nights. With the capture of
the leptocephali spread over most of the year and at widely spaced localities, there is
little indication of where and when spawning takes place.
Although an advanced leptocephalus transformed rapidly in an aquarium under
observation (39: 269; 40: 109), it cannot be concluded that development before
capture was equally fast, and that the rapid transformation under artificial conditions
was normal, especially in the light of what is known about the slow development of the
leptocephali of freshwater eels. At least the possibility exists that the rather advanced
leptocephali that have been described may have been several months old and were
caught far from the place where they had hatched.
Habits. The adult is usually caught in comparatively shallow water, but to what
extent it enters deep water offshore apparently has not been determined. The larva
has been taken at night at the surface under an electric light, by which it seems to
be attracted, and during the day in the shallow water of bays and estuaries with seines,
at least once in a creek with nearly fresh water (in Panama), and on one occasion in
the surf on the outer shores of Bogue Banks, North Carolina. During March 1937,
the larva and young adult were found in great abundance among islands in Panama
Bay. In one very shallow tide pool with a muddy bottom, off San José Island in the
Pearl Islands group, the water was literally “soupy” with fish and crustaceans. At the
time there was an exceptionally low tide, and the concentration was so great and the
water so shallow and warm that many small shrimp and fish perished, but the greater
part of the aggregation seemed to endure the situation. It was possible to select from
this pool a growth series ranging from leptocephali 70 mm long, the youngest present,
to young adults 52 mm long.
Food. This fish, known as the Grubber in Bermuda (40: 104) grubs up some of
its food from the bottom with its nose and sometimes turns somersaults in the proc-
ess. In the West Indies and Florida it may be seen by day (53: 187), along shallow
sandbanks and among underwater grasses, feeding during incoming tides on worms,
mollusks, and crabs; but in Bermuda it apparently comes into the shallows at night
to feed. Bivalves and small squid have been reported as its food (5: 37). In the Pearl
Islands, young adults were gorged with small shrimp, but a few fish (small atherinids)
had been eaten also.
I40 Memoir Sears Foundation for Marine Research
Relationship and Variation. Only one species has been recognized. Insufficient
material from its wide range has been studied, but the present study of specimens
from the Atlantic and Pacific coasts of America has shown no constant differences or
variations of importance.
Relation to Man. This species is not considered a good foodfish in most of its range
in America. Schroeder, reporting on the fisheries at Key West, Florida, stated that “‘it
is not highly regarded as a foodfish, and its commercial value is negligible” (61: 4).
Similar statements are included in publications about the fishes of the West Indies.
But Hollister, presumably reporting on observations made in Bermuda, stated to the
contrary that “‘it surpasses its famous cousin, the Silver King, or tarpon, in possessing
a combination of qualities, that of gaminess and of being a table delicacy” (go: 104).
From the writer’s own observations in Panama, the Bonefish is considered an inferior
foodfish, whereas the tarpon is highly regarded as food by the native population. Thus
tastes seem to differ in different localities.
However, the Bonefish ranks high with anglers, especially around Florida and
the Bahamas, for it is so wary when feeding on the flats that careful stalking and long
and delicate casting are called for; but once hooked, its runs are longer and swifter
and its resistance far more vigorous and indefatigable than one might expect from such
a comparatively small fish.
Miss Francesca LaMonte of the American Museum of Natural History, long
acquainted with the sport fisheries, has generously offered the following information:
This warm-water fish, which provides splendid angling with simple equipment, 1s
caught on the flats and can be fished there either from a boat or while wading. The
boat should be light as well as flat-bottomed and should be provided with an outboard
motor, oars, and anchors for both bow and stern. The fish is taken by casting or still
fishing, but the favorite method is casting. Light tackle with plug casting or saltwater
fly rod provides good sport. As bait, anglers may use conch, crabs or crayfish, feathers,
lures, or squid. Coming in with the tide, the Bonefish spreads out over the flats and
among the mangrove stands; when the tide ebbs, it seeks deep holes. On the flats,
the angler may chum or get his bait into the middle of a “mud” or group of Bonefish
that is feeding, with tails uppermost while grubbing on the bottom.
When hooked, the Bonefish makes a very fast and strong run, taking out yards
of line. In so doing it varies the straight dash by suddenly turning and coming toward
the boat or by running into the mangroves where the line may be cut on the roots.
After making several of these runs, the Bonefish usually begins to wear out, begins
circling the boat, and can finally be picked up in a landing net.
For many years the angling record stood at 13*/, pounds for one caught on the
famous Bonefish grounds off Bimini in the northern Bahama Islands, but this record
was eventually beaten. The present record is for a fish of 18 pounds 2 ounces, caught
off Kauai, Hawaiian Islands. In the western North Atlantic, the best known Bone-
fish grounds are the Florida Keys near Miami, the Islamorada-Marathon area, and
the northern Bahamas.
Fishes of the Western North Atlantic 141
Range and Abundance. The Bonefish is known from virtually all warm seas. On
the Atlantic coast of America it is plentiful around southern Florida, among the
northern Bahamas, and around Bermuda. But its area of chief abundance evidently
does not reach much farther to the north, for it is rare even at Beaufort, North
Carolina. However, odd specimens have been reported for the vicinity of New York
(2: 183), Newport, Rhode Island (68: 74), the vicinity of Woods Hole, Massachu-
setts (67: 741), and perhaps even Charlotte County, New Brunswick, at the mouth
of the Bay of Fundy (ftn. 15).
To the south it is present more or less regularly in the West Indies and on
the Atlantic coast of Panama, though it does not seem to be numerous; it is known
from as far south as Rio de Janeiro. On the Pacific coast of America it has been re-
ported as occurring from San Francisco Bay, California, to Talara, Peru.
Synonyms and References :17
Esox vulpes Linnaeus, Syst. Nat., ed. 10, 1758: 313 (orig. descr.; based on Vulpes bahamensis Catesby, pre-
Linnaean; type local. Bahamas; no type specimen).
Albula s. Salmo albula (nec Salmo aléula Linnaeus) Meuschen, in Gronow, Zoophyl., 3, 1781: index, unpaged
(on “4/éula” Gronow; type local. Mediterranean; type lost).
Clupea brasiliensis Bloch and Schneider, Syst. Ichthyol., 1801: 427 (orig. descr.; type local. Brazil; based on
Marcgrave’s ss. in Berlin).
Albula conorhynchus Bloch and Schneider, Syst. Ichthyol., 1801: 432, pl. 86 (orig. descr.; based on Gronow,
Plumier, and Marcgrave; no special type local. except Medit. and Amer.); Giinther, Cat. Fish.
Brit. Mus., 7, 1868: 468 (descr., synon., Amer. locals—West Indies, Jamaica, Belize, Puerto Cabello,
Bahia, Natal, and Pacif. cst. of Centr. Amer.); Yarrow, Proc. Acad. nat. Sci. Philad., 1877: 215 (Ft.
Macon, North Carolina).
Albula plumieri Bloch and Schneider, Syst. Ichthyol., 1801: pl. 86 (name on plate only; tarsus for conorhyachus).
Amia immaculata Bloch and Schneider, Syst. Ichthyol., 1801: 451 (orig. descr., after Macadi Parra [1787:
88, pl. 35, fig., nonbinomial]; type local. S. Amer.).
Clupea macrocephala Lacépéde, Hist. Nat. Poiss., 5, 1803: 426 (orig. descr.; type local. Martinique; after
Plumier fig.).
Glossodus forskali Agassiz, in Spix and Agassiz, Pisc. Brasil, 1829: 49; called Engraulis sericus on pl. 23 and
E, bahiensis on pl. 24, fig. 2 (orig. descr.; type local. Bahia, Brazil; type probably in Munich).
Albula parrae Cuvier and Valenciennes, Hist. Nat. Poiss., 19, 1846: 339 (orig. descr.; type local. Bahia and Rio
de Janeiro, Brazil; type in Paris).
Butirinus vulpes Storer, Synop. Fish. N. Amer., 1846: 212 (diagn., refs.).
Albula rostrata Gronow, Cat. Fish., Gray ed., 1854: 189 (orig. descr.; type locals. Amer. and Indian oceans,
Medit. Sea; type probably in London).
Conorhynchus plumieri Poey, Repert. Fisico-Nat. Cuba, 2, 1867: 423 (descr., Cuba).
Albula vulpes Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 258 (descr., range); Bean, Bull. U.S. Fish
Comm. (1888), 8, 1890: 206 (not uncommon at Cozumel, Yucatan); Collins and Smith, Bull. U.S.
Fish Comm. (1891), IZ, 1892: 104 (commerc. catch, Gulf coast, Florida, 1890); Eigenmann, Proc.
U.S. nat. Mus. (1892), 15, 1893: 135 (not abund. at San Diego, California); Henshall, Bull. U.S. Fish
Comm. (1894), 14, 1895: 211 (common at keys, passes, inlets of s. Florida, good food and game fish);
Lénnberg, Ofvers. Svensk. Vet. Akad. Forh., 9, 1895: 662 (Cape Haitien, Haiti); Jordan and Ever-
mann, Bull. U.S. nat. Mus., 47 (2), 1896: 411; 47 (4), 1900: fig. 179 (descr., range, synon.); Jordan
and Rutter, Proc. Acad. nat. Sci. Philad., 1897: 94 (Jamaica); Smith, Bull. U.S. Fish Comm. (1897), 27;
1898: 91 (rare at Woods Hole, Massachusetts); Evermann and Kendall, Rep. U.S. Comm. Fish. (1899),
1900: 55 (Florida); Gilbert, Proc. Wash. Acad. Sci., 2, 1900: 163 (Pernambuco and Maceié, Brazil);
17. Literature dealing with American specimens only, or in part, is cited. It does not seem necessary to cite Old
World references here.
14
Memoir Sears Foundation jor Marine Research
Evermann and Marsh, Bull. U.S. Fish Comm. (1900), 20 (1), 1902: 82 (descr. range, econ. import.,
Puerto Rico); Bean, Bull. N.Y. St. Mus., 60, Zool. 9, 1903: 182 (synon., descr., range, New York and
elsewhere); Bean, Field Mus. Publ., Zool., 7, 1906; 33 (§ Bermuda locals.); Gilbert and Starks, Mem.
Cal. Acad. Sci., 4, 1904: 39 (in Panama City market); Jordan and Thompson, Bull. U.S. Bur. Fish.
(1904), 24, 1905: 232 (Tortugas, Florida; discus.); Gill, Smithson. misc. Coll., 48, 1905: 40 (descr.
and figs. of skull, develop. of leptocephali, habitat, game qualities; C. H. Gilbert discovered leptocephalus
stage); Bean, Bahama Is., Fish., 1905: 297 (Spanish Wells, Bahamas); Smith, N.C. geol. econ. Surv.,
2, 1907: 117 (synon., diagn., U.S. coasts, esp. North Carolina); Coles, Bull. Amer. Mus. nat. Hist., 28,
1910: 344 (Cape Lookout, North Carolina); Cockerell, Smithson. misc. Coll., 56 (3), 1911: 3 (scale
descr.); Bull. U.S. Bur. Fish. (1912), 32, 1913: 122, fig. 2 (scales not different from those of Dixonina;
similar to Syzodus and Esox); Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 204 (Santo Domingo,
St. Martin, Jamaica); Nichols, Bull. Amer. Mus. nat. Hist., 3Z, 1912: 181 (not uncommon in Havana,
Cuba, market); Gudger, J. Mitchell Sci. Soc., 28, 1913: 166 (rare, Beaufort, North Carolina); Starks,
Stanf. Univ. Publ., Univ. Ser., 1913: 8 (Natal, Brazil); Sumner, Osburn, and Cole, Bull. U.S. Bur. Fish.
(1911), 32 (2), 1913: 741 (rare at Woods Hole, Mass.); Halkett, Check List Fish. Canada, 1913: 45
(Bay of Fundy, New Brunswick); Weber and de Beaufort, Fish. Indo-Aust. Archipelago, 2, 1913: 7,
fig. 5 (synon., descr., develop. of leptocephali; figs. of leptocephali labeled “after Gilbert,” but without
citation to Gilbert; figs. of larvae apparently after Gill, 1905); Fowler, Proc. Acad. nat. Sci. Philad., 67,
1915: 522 (Trinidad, B. W.1.); 68, 1916: 397 (Port Limon, Costa Rica); 69, 1917: 128 (Colén,
Panama); Metzelaar, Trop. Atlant. Vissch., rg1g: 8 (St. Martin, and other Windward and Leeward
islands; not common, not much used as food); Metzelaar, Bijdr. Dierk. K. zool. Genoot. Amst., 1922:
134 (Curacao); Meek and Hildebrand, Field Mus. Publ., Zool., 75 (1), 1923: 178 (synon., descr.,
range, Panama); Schroeder, Rep. U.S. Comm. Fish. (1923), Append. 12, 1924: 4 (commerc. value,
Key West, Florida); Mowbray, Book of Fishes, Nat. geogr. Soc., ed 1924: 135-137, color pl. on p. 122;
ed. 1939: 186-187, color pl. on p. 201 (genl. accts.); Breder, Bull. Bingham oceanogr. Coll., 2 (2),
1928: 4 (young adult, Sonora, México; adult from Pearl Is., Panama); Nichols and Breder, Zoologica,
N.Y., 9 (1), 1927: 34 (diagn., distr., life hist., size); Beebe and Tee-Van, Zoologica, N.Y., To (1), 1928:
37 (refs., field characters, size, color, range, uncommon at Port-au-Prince, Haiti); Nichols, N. Y. Acad.
Sci., TO (2), 1929: 200 (distr., not uncommon in Puerto Rican waters, diagn., habits); Breder, Field Bk.
Mar. Fish. Atlant. Cst., 1929: 60, fig. (diagn., abund., range, habits, size); Fowler, Proc. Acad. nat. Sci.
Philad., 80, 1929: 609 (Barnegat Inlet, Atlantic City, first definite recs. for New Jersey); Jordan, Manual
Vert. Anim. NE U.S., 1929: 38 (diagn., range); Fowler, Proc. Acad. nat. Sci. Philad., $2, 1930: 269
(Grenada, B. W. I.); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 39 (range,
synon.); Gowanloch, Bull. La. Conserv. Dept., 21, 1932: 44, fig.; also Bull. La. Conserv. Dept., 23,
1933: 43, fig. (relation., life hist.); Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda, 1933: 34 (descr.,
distr., Bermuda); Gregory, Trans. Amer. philos. Soc., 23 (2), 1933: 144, fig. 37 (relations. as shown
by skull); Breder, Zoologica, N. Y., 28, 1934: 59 (Grassy Creek, Andros I., Bahamas); Beebe and
Hollister, Zoologica, N. Y., Z9 (6), 1935: 211 (leptocephali, adults, Union I., Grenadines, B. W. 1.);
Myers, Copeia, 1936: 83 (cf. Dixonina nemoptera); Hollister, Bull. N.Y. zool. Soc., 39, 1936: 104—
109, figs. (metamorphosis notes); Zoologica, N.Y., 21 (4), 1936: 268-275, figs. (caudal skel.); Far-
rington, Atlant. Game Fishing, 1937: 183-186 (scientific name not given, genl. acct., abund. Bi-
mini, Bahamas); Fowler, Proc. Acad. nat. Sci. Philad., 89, 1937: 304 (Ventnor, New Jersey); Heilner,
Salt Water Fishing, 1937: 174-194 (genl. acct.); Kaplan, Big Game Angler’s Paradise, 1937:
190-195 (genl. acct.); Butsch, J. Barbados Mus. nat. Hist. Soc., 7 (1), 1939: 18 (uncommon at Bar-
bados); Longley and Hildebrand, Publ. Carneg. Instn. Wash., 535, 1941: 5 (color of leptocephali from
Tortugas, Florida, described); Alexander, ‘Dana’ Rep., 53, 1961 (descr. larval develop.; world-wide
distr. and hydrogr.).
Atopichthys esunculus Garman, Mem. Mus. comp. Zool. Harv., 24, 1899: 327, pl. 65, figs. 2, 2a (orig. descr.;
type local. off Acapulco, México; types in MCZ); Jordan, Evermann, and Clark, Rep. U.S. Comm.
Fish. (1928), 2, 1930: 54 (ref. to orig. descr.).
Elops saurus Gregg (not Elops saurus Linnaeus, but Esox vu/pes Linnaeus), Where to Catch Fish. E. Cst. Fla.,
1902: 33, fig. (descr., habitat, feed., instruct. for angling, fairly edible).
Negative Reference:
Albula vulpes Gregg, Where to Catch Fish. E. Cst. Fla., 1902: 34 (not A/bula vulpes Linnaeus, but E/ops
saurus Linnaeus).
Fishes of the Western North Atlantic 143
Genus Dixonina Fowler 1911
Dixonina Fowler, Proc. Acad. nat. Sci. Philad. (1910), 1911: 651; type species by original designation, Dixo-
nina nemoptera Fowler.
Characters. See species account below.
Remarks. Dixonina is easily distinguished from A/bu/a by the produced last ray of
both dorsal and anal fins. Furthermore, Dixonina has a somewhat longer and more
conical snout and a larger mouth than A/bu/a.
Range. A single species is known; this has been reported from the West Indies,
Venezuela, northern Brazil, and from the Pacific coast of México and Central America.
Dixonina nemoptera Fowler 1911
Shafted Bonefish1§
Figure 24
Study Material. One Pacific coast specimen, about 413 mm long, 334 mm SL,
from Acapulco, México, USNM 75547, recorded by Myers.
Ficure 24. Dixonina nemoptera, 413 mm TL, Acapulco, Pacific coast of México, USNM 75547. From
drawing by Ann S. Green.
Description. Proportional dimensions in per cent of standard length, and counts,
based on Pacific specimen 334mm SL.
Body: depth at origin of dorsal 23. Anal fin: length of base 5.5.
Head: length 29.5. Pectoral fin: length 15.6.
Snout: length 10.8.
Eye: diameter 5.4. Scales: 76.
Maxillary: \ength from tip of snout Gill rakers: 5 +10.
13.5. Fia rays: dorsal 19; anal 8; pectoral 16.
Bopy slender, the back somewhat elevated, the ventral outline nearly straight;
depth at origin of dorsal 4.3 in SL. Scatzs firm, with crenulate membranous edges.
18. Name suggested by George S. Myers.
144 Memoir Sears Foundation for Marine Research
Heap long and low, flat above, its depth at middle of eye exceeding its width at the
same place by diameter of pupil, 3.4 in SL. Snour very long, projecting more than
a third of its length beyond mandible, 2.7 in head. Eve moderately small, 5.4 in head,
its center notably nearer to tip of snout than to margin of opercle. MaxrLiary ex-
tending nearly to middle of eye, 2.15 in head (measured from tip of snout).
Dorsat fin moderately high anteriorly, its longest anterior rays about as long as
snout and eye, its last ray filamentous, almost reaching vertical from origin of anal, its
origin about equidistant between tip of snout and caudal base. Caupat deeply forked,
the upper lobe apparently longer than the lower (somewhat damaged). ANAL very small,
its last ray produced, extending nearly to base of caudal fulcra, its origin equidistant
between base of caudal and base of last dorsal ray, its base 3.5 in head. Prtvic not
much smaller than pectoral, inserted about under middle of dorsal. Pecrorat not fal-
cate, extending halfway to pelvic, 1.9 in head. AxrLLary scate of pectoral adherent to
body, not very distinct.
Color. An old preserved specimen, slightly bluish above, silvery below; narrow
dark streaks between rows of scales on upper part of side. Live specimens from the Paci-
fic were described by Beebe (4: 44) as follows:
Brilliant silver, appearing dark above in certain lights with dark green on the head; this dorsal pig-
mentation resolves into eight very dark lines along each side of upper half of the body, covering about
one-fourth of each scale nearest to the adjoining line. Dorsal fin greenish, caudal dusky, anal silvery at base,
a dusky spot at pectoral base, with yellow spot behind; basal membrane of pectoral apple green, rest of fin
dusky. These colors fade at death. Iris silvery.
In the young, at least up to gomm in length, two rows of dark spots extend along the side of body,
the upper close to the mid-back. In a 115 mm fish the spots are less conspicuous and the dorsal dark lines
begin to be distinct. The spots persist after death. In full-grown fish the dark scale lines are fainter than in
fish of medium size.
Relationship. The specimen at hand from the Pacific agrees very well with two
published accounts of Atlantic fish. Some Pacific coast specimens have been described
as D. pacifica by Beebe (4: 43), who considered them specifically distinct from the
Atlantic ones. On the basis of available descriptions (not specimens) of Atlantic fish,
the differences between his Pacific specimens and those from the Atlantic were set forth
as follows:
In the former (D. pacifica) there is a greater number of lateral line scales (81 to 84, as compared
with 76); increased number of vertical lines of scales on dorsal half (9 not 7) and on ventral half (9 not 7);
decrease in predorsal scales (22 instead of 30); small eye (6.8 to 7, not 4.15); longer pelvic fins (2.2 and
not 2.7); greater number of gill-rakers (7 + 11 instead of 4+9).
Most of the apparent distinctions between the Atlantic and Pacific specimens
pointed out by Beebe become obscure or are of doubtful value in the light of measure-
ments and enumerations based on the Acapulco fish. The number of scales in a longi-
tudinal series seems to differ on the opposite sides of this fish; on the right side I counted
76 scales with pores (excluding greatly reduced ones on caudal base) and 79 on the left.
Fishes of the Western North Atlantic 145
Again, I counted 77 oblique series (running upward and backward) just above the
lateral line on the right side and 83 on the left. No variation in the number of longi-
tudinal rows between the lateral line and origin of dorsal, and between the lateral line
and origin of anal, occurs in the specimen studied, as these are respectively 8 and 6
full rows on each side. As to the number of scales in advance of the dorsal, the speci-
men before me has 21 modified scales in the median series, and 31 oblique series
of ordinary scales (running upward and backward) on each side of the median row.
As neither Fowler (24: 652) nor Beebe stated which of the two was counted, I
made inquiry of Mr. Fowler, who re-examined the type and found 21 modified scales
in the median series, and 30-32 ordinary scales in the oblique series. As to the size of
the eye in proportion to the length of the head, the specimen at hand is intermediate,
the eye being contained in the head 5.4 times. The pelvic fin also is intermediate, its
length 2.35 times in head. I have no doubt that the gill rakers in Dixonina become
shorter and obscure with age, as they do in /bu/a, and that in large specimens it ts
difficult without dissection to see all of the bases, for that is all that remains of some of
them. In the specimen from Acapulco, I find 5 +10, including one in the angle with
the lower limb count, a figure intermediate between counts given by Beebe and by
Fowler.
It may be concluded, on the basis of the evidence presented, that D. pacifica for
the present at least is a synonym of D. nemoptera.
Range. This albulid, as here understood, is known from the Atlantic from the
type taken at Santo Domingo and from a specimen collected at Puerto Cabello, Vene-
zuela.1® For the Pacific it is known from a specimen taken at Acapulco, México
(reported by Myers), and from 19 specimens taken off the coast of Costa Rica
(reported as D. pacifica by Beebe). According to Walford (71: 119), the fish from the
Gulf of California figured as A/bula vulpes by Kumada and Hiyama (47: pl. 5) also
is this species.
Synonyms and References:
Dixonina nemoptera Fowler, Proc. Acad. nat. Sci. Philad. (1910), 1911: 652, fig. (orig. descr.; type local.
Santo Domingo, West Indies; ANSP 1597); Myers, Copeia, 1936: 83 (Acapulco, México; relation. to
Albula); Walford, Copeia, 1939: 119 (specimen from Gulf of California, figured as 4/bula vulpes in
47, 1s Dixonina).
Albula nemoptera Metzelaar, Trop. Atlant. Vissch., 1919: 9 (descr.; Puerto Cabello, Venezuela).
Albula vulpes (not of Linnaeus) Kumada and Hiyama, Mar. Fish. Pacif. Cst. Mexico, 1937: 27, pl. 5 in color
(descr., in part at least, and plate based on Dixonina).
Dixonina pacifica Beebe, Zoologica, 27 (8), 1942: 43, figs. 1-5 (orig. descr.; type local. Puerto Culebra, Costa
Rica; SU 46486; cf. D. xemoptera).
19. One seen from Brazil.—G. S. Myers.
Ses Sy Wes
a
2I.
22.
23.
24.
TEXT AND FOOTNOTE REFERENCES
Bascock, The Tarpon, 1936; ed. 5, 1951.
Bean, Bull. N. Y. St. Mus., 60, Zool. 9, 1903.
Beebe, Bull. N. Y. zool. Soc., 30 (5), 1927-
Beebe, Zoologica, N. Y., 27 (8), 1942.
Beebe and Tee-Van, Zoologica, N. Y., 10 (1),
1928.
Bigelow and Welsh, Bull. U.S. Bur. Fish.,
20 (1), 1925.
Bigelow and Schroeder, Copeia, 1940.
Boulenger, Ann. Mag. nat. Hist., (6) 20, 1897.
Boulenger, Cambr. Nat. Hist., 1904.
Breder, Zoologica, N. Y., 4 (4), 1925.
Breder, Bull. N. Y. zool. Soc., 36, 1933-
Breder, Bull. N. Y. zool. Soc., 47, 1939.
Breder, Zoologica, N. Y., 29, 1944.
Gavenat; Rev. Trav. Off. Péches marit., Zo,
Fasc 4, Liv. 4, Pt. 2, 1937.
Cockerell, Smithson. misc. Coll., 56, 1910.
Cockerell, Science, 1911.
Cockerell, Bull. U.S. Bur. Fish. (1912), 32,
1913.
Chadabaram and Menon, Proc. zool. Soc.
Lond., II7, 1947.
Cuvier, Régne Anim., ed. 2, 2, 1829.
Cuvier and Valenciennes, Hist. Nat. Poiss.,
Ig, 1846.
Expman, Copeia, 1960.
Evermann and Marsh, Bull. U. S. Fish Comm.,
20, 1902.
Fowzer, Rep. N. J. Mus. (1906), 1907.
Fowler, Proc. Acad. nat. Sci. Philad., 62, 1910.
146
Fowler, Bull. Amer. Mus. nat. Hist., 70 (1),
1936.
Fowler, Proc. Acad. nat. Sci. Philad., 93, 1941.
Geurincer, J. Fla. Acad. Sci., 27 (3), 1958.
Gill, Smithson. misc. Coll., 48 (1), 1905.
Gowanloch, Bull. La. Conserv. Dep., 23,
133:
Gregory, Trans. Amer. philos. Soc., 23 (2),
1933-
|B aaae Checklist Fish. Canada, 1913.
Harrington, Copeia, 1958.
Heilner, Salt Water Fishing, 1937.
Henshall, Bull. U.S. Fish Comm., 14, 1895.
Hildebrand, Copeia, 1934.
Hildebrand, Sci. Mon., N. Y., 44, 1937.
Hildebrand, Zoologica, N. Y., 24 (1), 1939-
Hildebrand and Schroeder, Bull. U.S. Bur.
Fish., 43, 1928.
Hollister, Zoologica, N. Y., 2, 1936.
Hollister, Bull. N. Y. zool. Soc., 39 (3), 1936.
Deleted.
Hollister, Zoologica, N. Y., 24 (4), 1939.
Tavinz, Fish. Gold Coast, 1947.
Jonas Guide to Study Fish., 1905.
Jordan and Evermann, Bull. U. S. nat. Mus.,
47 (I), 1896.
Kaprax, Big Game Angler’s Paradise, 1937.
Kumada and Hiyama, Marine Fish. Pacif. Cst.
Mexico, 1937.
48.
ae
50.
5 Te
52.
53:
54.
55:
56.
57:
58.
59:
60.
Fishes of the Western North Atlantic
Liston, Bull. U.S. Bur. Fish., 24, 1905.
Longley and Hildebrand, Publ. Carneg. Instn.
Wash., 535, 1941.
Manccrave, Hist.-Nat., Brazil, 1648.
Merriman, Copeia, 1939.
Morden, Nat. geogr. Mag., 86, 1944.
Mowbray, Book of Fishes, 1939.
Myers, Copeia, 1936.
Nicuots and Breder, Zoologica, N. Y., 9 (1),
1927.
Deleted.
Prvo, Bull. Soc. Hist. nat. Toulouse, 70, 1936.
Recay, Ann. Mag. nat. Hist., (8) 3, 1909.
Regan, Ann. Mag. nat. Hist., (8) 5, 1g1o.
Serene and Ribeiro, Arch. Mus. nac. Rio
de J., 12, 1910.
6r.
68.
69.
70.
ie
72.
73-
74-
147
Schroeder, Rept. U.S. Comm. Fish. (1923),
1924.
Schultz, Proc. U.S. Nat. Mus., 99, 1949.
Simpson, Copeia, 1954.
Smith, N. C. geol. econ. Surv., 2, 1907.
Starks, Stanf. Univ. Publ. Biol., 1913.
Storey and Gudger, Ecology, 77, 1936.
Sumner, Osburn, and Cole, Bull. U.S. Bur.
Fish., 32 (2), 1913.
Tracy, 40th Rep. R. I. Comm. inl. Fish., 1910.
\Wittennen Sea Fish. Trinidad, 1gto.
Vladykov and McKenzie, Proc. N. S. Inst. Sci.,
29; 1935-
Wazroro, Copeia, 1939.
Whitley, Rec. Austral. Mus., 17 (6), 1929.
Whiton and Townsend, Bull. N. Y. zool. Soc.,
3I, 1928.
Wilcox, Bull. U.S. Fish Comm., 6, 1887.
10*
Suborder Clupeoidea
COMPOSITE AUTHORSHIP!
Characters and Key to Families
HENRY B. BIGELOW
Museum of Comparative Zoology
Harvard University
Characters. Eyes of the usual sort, i.e. not at the tips of short (tubular) cylinders
or of slender stalks, nor with orbital cavity extended downward. Aprposs fin absent.
Petvic fins at least no farther forward than midlength of body and farther rearward
than this in most, but lacking in a few. Fin rays jointed, and most of them branched.
Scares: the head naked, except in some Alepocephalidae; the trunk scaly in most,
but naked in some of the Alepocephalidae (p. 250); scales large, thin, their free bony
edge entire in most, often with crenulated membranous margin, but more or less
strongly pectinate in a few genera (Brevoortia, p. 342, Fig. 85; and Ethmidium),
exposed surface smooth or only weakly sculptured; midline of abdomen with scales
of the ordinary shape in most, but with a double row of larger, sharp-pointed, and
stiffer scales (scutes) in others (Clupeidae, pp. 257, 259, 260). BarBets absent on chin or
on throat. Luminescent orcans lacking in most, but present in a few (some Alepoce-
phalidae and all Searsiidae). Larrrat Line well developed in most families (Engrau-
lidae, Alepocephalidae, Searsiidae) but not visible in others (Clupeidae) though
continuing rearward below scales to base of caudal except as it may occasionally
perforate a few anterior scales (10: 957). GiLL MEMBRANES wholly free from isthmus
in the great majority.2 Bony GuLarR PLATE absent in chin region between branches of
1. With the late Samuel F. Hildebrand contributing the family Engraulidae as well as the family Clupeidae, except
for the genera Harengula by Luis R. Rivas and Dorosoma by Robert R. Miller.
2. Connected to the isthmus by a thin membrane in the anchovy Cetengraulis (p. 245).
148
Fishes of the Western North Atlantic 149
lower jaw. G1Lt RaKers present in all. PRorrHMorps not paired. Sides of snout without
SENSORY CANAL-BEARING BONELETS. BRANCHIOSTEGAL Rays Slender, rod-like, not forming
part of the opercular system. Tern, if any,? small in most, larger in a few (Chirocen-
tridae); dental patterns various; teeth never present on the parasphenoid bone in mid-
line of roof of mouth rearward. PREMAXILLARIES not protractile, except in some Alep-
ocephalidae and in the Phractolaemidae.* Maxriiaries forming part of upper bor-
der of mouth in the great majority.» Lower jaw without predentary or supra-angular
bones. TEMPORAL FORAMINA present.® BasaL RapIAL1A of the pectoral fin in one row
in most, articulating directly with the pectoral girdle; in two rows in a few (among
the Chirocentridae, rr: 9-11, fig. 23; I: 227, fig. 118, 424). PecroraL GIRDLE with
a mesocoracoid element. Pretvic GrrpLE not connected to the pectoral girdle. Last
few VERTEBRAE not evidently upturned in any; lateral processes bearing pleural ribs
(parapophyses) simply wedged into pits in vertebral centra, not fused with the lat-
ter; a lower series of intermuscular bones well developed in many (notably among
Clupeidae), connected basally to pleural ribs and lying between muscle blocks (myo-
tomes).? Ovaries with complete Ovipucts, so far as known. Swim BLADDER present in
most,® dorsal in position, and connected throughout life to the alimentary tract via
an open pneumatic duct. Sromacu with numerous pyloric caeca. [nrestrne without
spiral or circular folds in most, but perhaps not in all.® ArTERIAL CONE OF HEART
never with two rows of valves. Larvae not leptocephalus-like.
Families. The following families fall within the Suborder as defined above: 1
Clupeidae, including Dussumierinae and Dorosomatinae, p. 257; 11 Alepocephalidae,
p. 250; 11 Searsiidae, p. 254; 1v Engraulidae, p. 152; v Chirocentridae (regarded by
Regan [8: 313-314] and Norman [5: 72] as a subfamily of Clupeidae; by Berg
[z: 227, 422] as a separate Suborder, Chirocentroidei), Indian Ocean, China, and
Japan; v1 Chanidae, tropical Indo-Pacific; vi1 Phractolaemidae, Niger and Congo rivers;
vit Kneridae, rivers of tropical Africa; and 1x Cromeriidae, the Nile.
Clupeidae, Alepocephalidae, Searsiidae, and Engraulidae are represented in the
North Atlantic. The Dolichopterygidae and the Macristiidae have also been referred
to the Clupeoidea by one author or another. But Dolichopteryx (Part 4) falls among the
Argentinoidea, as has long been evident from Parr’s excellent illustration of it (6: 37,
fig. 14). And the affinities of the Macristiidae remain as doubtful today as was the
case in 1911, when Regan (7: 204, 205) redescribed the only specimen of the family
that has yet been seen.
3. The Chanidae lack teeth, at least as adults.
4. A freshwater family.
5. In Nematalosa the mouth is bordered above by the premaxillaries, with the maxillaries covered by the lip, as is
pictured very clearly by Ridewood (9: 465, fig. 127 as “Chaetoessus”).
6. Shown clearly in Ridewood’s illustrations; see ftn. 5.
7. For a diagram showing the relative positions of the pleural ribs and of the lower intermuscular bones, see Ber-
tin (4: 704, fig. 462).
8. Lacking among the Alepocephalidae; probably among the Searsiidae also.
g. A structure suggesting the elasmobranch spiral valve, but probably not homologous, has been reported
in Chirocentrus (see especially Goodrich, 3: 116, fig. 77) and in Alepocephalus (Cuvier and Valenciennes,
2: 176).
I50 Memoir Sears Foundation for Marine Research
Key to Families of Clupeoidea, Known or to be Expected in
the Western North Atlantic
1a. Tip of fleshy snout overhanging mouth. Engraulidae, p. 152.
1b. Tip of fleshy snout not overhanging mouth; at most the upper jaw protruding a
little beyond lower.
2a. Shoulder close behind upper part of gill opening, with a conspicuous pro-
jecting tubular papilla, open at the tip. Searsiidae, p. 254.
2b. Shoulder without projecting tubular papilla.
3a. Rear end of base of dorsal fin in advance of origin of caudal by a distance
as long as head (longer in most); sides silvery; swim bladder well devel-
oped. Clupeidae, p. 257.
3b. Rear end of base of dorsal fin in advance of origin of caudal by a distance
considerably shorter than head; sides not silvery; no swim bladder.
Alepocephalidae, p. 250.
KY
TEXT ANDFFOOTNOTE REFERENCES
: Bere, Class. Fish., Trav. Inst. zool. Acad. Sci.
URSS, 5 (2), 1940; also reprinted by J. W.
Edwards, Ann Arbor, Mich., 1947 (Russ.
and Eng.).
y Cink and Valenciennes, Hist. Nat. Poiss.,
19, 1846.
: Goonnicn, Treat. Zool., 1909.
Grassé, et a/., Trait. Zool., 13 (1), 1958.
- Norman, Synop. Orders, Families, Genera of
o}
Recent Fish. (mimeo.), 1957.
6. Parr, Bull. Bingham oceanogr. Coll., 3 (7),
1937-
Fe Recan. Ann. Mag. nat. Hist., (8) 7, 1911.
8. Regan, Encyc. Brit., 14th ed., 9, 1937.
g. Ridewood, Proc. zool. Soc. Lond., 2, 1904.
IO. Smrirr, Scand. Fish., Pt. 2, 1895.
rr. Starks, Stanf. Univ. Publ. Biol., 6 (2), 1930.
151
Family Engraulidae’
SAM UIE FP? HILDEBRAND?
Characters. Bopy moderately slender, compressed. Curest and ABDoMEN in
American species often carinate, without bony serrae. Mourn horizontal or nearly so.
Maxiitary extending well beyond eye, sometimes nearly or quite to gill opening.
Sout blunt, generally projecting far beyond tip of mandible. Eye placed well forward,
in anterior half of head, sometimes with well developed adipose tissue in adults.
PREMAXILLARIES not protractile, embedded in tissue of snout anteriorly, widely sepa-
rated medianly. Trrrn typically minute (some on jaw enlarged in Lycengraulis); present
on jaws, vomer, palatines, pterygoids, and hyoids. Grit covers generally separate and
free from isthmus (connected by a thin membrane in Cerengrau/is). PsEUDOBRANCHIAE
present. Grit raKERs generally slender, rarely short, broad, and spiny; increasing in
number with age in some species (as in Anchovia). Scares thin, cycloid (usually lost
on preserved specimens), generally if not always forming a sheath on bases of dorsal
and anal fins; an enlarged scale in axil of pectoral and pelvic fins in American species.
LaTERAL LINE absent. Dorsat fin usually median in position, rarely behind mid-
length, with about 12-16 rays in American genera. Apipose Fin lacking. CaupaL
rin rather deeply forked. Anat with various ray counts, about 15-40 in American
species.
Genera. The division of the family into compact and clearly definable genera is
difficult. Thus there is a divergence of opinion among students concerning the status
of described genera. Seven genera have been recognized (3), all confined to American
waters, excepting Exgraulis, which, though not represented in the western North
Atlantic, is very widely distributed in both hemispheres; 75 species have been recog-
1. Edited, with some revision and expansion, by George S$. Myers, Myvanwy M. Dick, Henry B. Bigelow, and
Yngve H. Olsen.
2. August 15, 1883—March 16, 1949.
1$2
Fishes of the Western North Atlantic 153
nized, 44 from the Atlantic and 32 from the Pacific, including one species common
to both coasts of tropical America. For the Atlantic species that come within the scope
of this work, see the Table of Contents.
Commercial Importance. Some anchovies, because of their tremendous abundance,
are of considerable importance. Some are canned whole or are used in the production
of anchovy paste, and larger species are commonly sold fresh in the market, as in
South America; they are also used extensively as bait. But their value as a forage fish
for water birds and creatures that frequent the sea far exceeds their direct value to
man. Many of the larger commercially important fishes, such as the seatrout, bluefish,
and mackerels, feed on them extensively.
Range. Representatives of the family occur in the warmer shore waters through-
out the world. Some of the species ascend freshwater streams, and probably a few live
entirely in fresh water. The Engraulidae are most numerous in tropical regions, but
several species either range into temperate waters or live there permanently.
Key to Genera of the Western North Atlantic
1a. Gill covers not connected across isthmus by a membrane.
2a. Teeth in jaws small or minute, all of about uniform size.
3a. Origin of anal fin posterior to origin of dorsal; tip of pelvic fin usually
below origin of dorsal.
4a. Maxillary more or less sharply pointed distally, generally reaching
well beyond joint of mandible, often to margin of opercle, or
nearly so.
sa. Gill rakers close-set, increasing in number with age, 40—S0 in
young, about 130 in adults, on lower limb of first arch; body
depth usually more than 25°/, of length; maximum size about
250mm. Anchovia Jordan and Evermann 1896, p. 155.
5b. Gill rakers not close-set, only about 15-30 on lower limb, not
known to increase in number with age; body depth usually
less than 25°/, of length; maximum size generally under
150 mm. Anchoa Jordan and Evermann 1927, p. 162.
4b. Maxillary square or rounded distally, rarely reaching joint of man-
dible. Anchoviella Fowler 1911, p. 204.
3b. Origin of anal fin in advance of dorsal origin, rarely under it; tip of
pelvic fin anterior to a vertical from dorsal origin by a distance equal
to about 50°/, of head.
6a. Distance of dorsal fin origin from tip of snout much less than
twice the distance from dorsal origin to caudal base; gill rakers
short and broad, only about 14 on lower limb of first arch; anal
with 30—33 rays. Prerengraulis Giinther 1868, p. 228.
54 Memoir Sears Foundation for Marine Research
6b. Distance of dorsal origin from tip of snout about twice the distance
of dorsal origin from caudal base; gill rakers long, slender, about 33
on lower limb of first arch in young, probably more in adults; anal
with about 25 rays. Hildebrandichthys Schultz 1949, p. 230.
2b. Teeth in jaws, especially the lower ones, notably large, usually unequal in
size. Lycengraulis Giinther 1868, p. 233.
1b. Gill covers broadly connected across isthmus by a thin membrane (easily torn).
Cetengraulis Giinther 1868, p. 245.
Explanation of Measurements and Counts for Engraulidae.
Tora. LENGTH: from tip of snout to vertical at rearmost point of longest caudal
lobe.
STANDARD LENGTH: from tip of snout to base of caudal fin, i.e. to the last vertebra,
as nearly as can be determined without dissection.
CauDAL PEDUNCLE: least depth unless otherwise noted.
LENGTH oF HEAD: from tip of snout to most distal part of bony opercular margin,
regardless of whether the straight line between these points is horizontal or oblique.
Depru oF HEAD: vertical distance at joint of mandible.
PosToRBITAL LENGTH: from posterior rim of eye to most distal part of bony oper-
cular margin.
MaxiLiary: from its anterior embedded end (located by touch) to its posterior
extremity.
CueEEK (smooth bone below and behind eye): distance from rim of eye to its
distal point; the degree of its posterior angle determined by a protractor.
DorsaL anpD ANAL FINS: the counts include all rays, whether simple and un-
segmented, simple and segmented, or branched; the last ray of the anal, even though
split to the base, was counted as one unless definitely separated at base.
LENGTH OF PECTORAL: from base of upper ray to tip of longest ray.
PECTORAL AXILLARY SCALE: distance from base of upper ray of pectoral to tip of
the modified scale or process.
Spiint: adhering to first ray of pectoral; not considered distinct and not counted.
ScaLes oR SCALE PocKETs: counted along the side from the upper anterior angle
of the gill opening to the base of the caudal; as the scales are nearly always lost in
preserved specimens, they are generally not shown in the illustrations, except for a
small patch.
Grit rakeErs: the counts are given as a formula, e.g. 12-16 + 15-18 (as for Anchoa
spinifer), meaning a range of 12-16 rakers on the upper limb of the first branchial
arch and 15-18 on the lower limb. The counts include those rudiments, if present,
that have a free tip but not those that are mere tubercles. The gill raker at the angle of
the first arch generally has a plainly visible root, extending either upward or down-
ward; if the root was directed upward, the raker was included with the upper limb
Fishes of the Western North Atlantic 155
count; if directed downward, it was counted with the lower limb count. Rarely, the
raker at the angle has two roots, one directed upward and one downward; in such
cases it was included with the lower limb count.
VERTEBRAE: total number, including hypural, as determined by dissections.
Genus Anchovia Jordan and Evermann 1896
Anchovia Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 449; type species by original designa-
tion, Exgraulis macrolepidotus Kner and Steindachner 1864.
Characters. Bopy in adults compressed, its depth about 2.6-3.8 in SL. Scates
firm, rather adherent. TrEru small, not disappearing with age. GILL RAKERS numerous,
increasing in number with age, about 40 on lower limb in young, 100 or more in
large specimens. Anat origin somewhere under anterior 66°/, of dorsal base. Verre-
BRAE 41-43.
Size. A length of 175 mm and upward is attained by most of the species.
Species. The genus Anchovia is represented in the Atlantic by 4. c/upeoides and
A. nigra, and in the Pacific by three species.
Range. In the Atlantic, from the West Indies southward to or beyond Pernambuco
(Recife), Brazil, and in the Pacific from the Gulf of California to Ecuador.
Key to Species of the Western Atlantic
1a. Insertion of pelvic fins about equidistant between base of upper ray of pectoral
fin and origin of anal; snout projecting beyond mandible by about half its length;
anal fin with 30-35 rays, most frequently with 33; vertebrae 42 or 43.
clupeoides (Swainson) 1839, below.
1b. Insertion of pelvic fins nearer to base of upper rays of pectoral fin than to origin
of anal; snout projecting beyond mandible by about 67°/, of its length; anal fin
with 28-32 rays, most frequently with 29-31; vertebrae 39-41.
nigra Schultz 1949, p. 158.
Anchovia clupeoides (Swainson) 1839
Sardina Boca Torta, Bocén, Hachudo
Figure 25
Study Material. At least 29 specimens, 75-205 mm TL, from: Puerto Rico;
Jamaica; Cuba; Trinidad; several places on the Atlantic coast of Panama; the Gulf
of Venezuela; Laguna de Tacarigua, Venezuela; and Pard, Rio Grande do Norte, and
Pernambuco (Recife), Brazil. Some small specimens, 48 mm TL and upward, are at
156 Memoir Sears Foundation for Marine Research
hand, but their proportions are not included because they differ considerably from the
larger ones, especially in having a more slender body.
Distinctive Characters. A. clupeoides is closely related to A. rastralis of the Pacific
coast of Middle America, from which it differs in having a slightly longer anal fin,
inserted a little farther forward with respect to the dorsal. In specimens of equal size,
the maxillary is usually a little shorter and tapers more abruptly in c/upeoides, and its
body is somewhat more slender. 4. c/upecides apparently grows to a larger size than
A. rastralis, for none of the latter has been reported to be larger than 110mm TL.
See also 4. nigra, p. 161.
OAS sz
SS GEE
Ficure 25. Axchovia clupecides, 60mm TL, 44 mm SL, from Gulf of Venezuelaj USNM 127552. Drawn
by Louella E. Cable.
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens 75-205 mm TL.
Body: depth 25-31. Scales: ca. 39-43.
Head: length 27.6-31. Gill rakers: ca. 35-40 in younger spec-
Snout: length 3.1-4.3. imens, 75-110 in those of 130 mm
Eye: diameter 6.0-8.3. pil,
Postorbital: distance 15.5—-18.5. Fin rays: dorsal 13-15; anal 30-35;
Maxillary: length 19.5-22. pectoral 13 or 14.
Mandible: length 19-20.5. Vertebrae: 42 or 43 (6 specimens).
Anal fin: length of base 30-33.
Pectoral fin: length 15.4-17.3.
Bopy strongly compressed, its depth increasing with age, about 3.2—-4.0 in SL
in specimens 75mm TL and upward (ca. 4.5 at so mm). Heap 3.2—3.6 in SL, its depth
about equal to its length without snout in specimens 150-160 mm TL; less in smaller
ones. Snour short, projecting about half of its length beyond mandible, 6.0—-8.0 in
head. Eve 3.5—4.5 in head. Maxitiary abruptly pointed, reaching to or beyond joint
Fishes of the Western North Atlantic 157
of mandible, 1.3-1.5 in head. ManpisLe 1.4—1.6 in head. Cueek as long as snout
and eye in large ones, proportionately shorter in small ones, its posterior angle about
35°. Gitt RakeErRs at angle fully as long as eye in large specimens, shorter in young.
Dorsat fin originating equidistant between caudal base and anterior half of eye,
its longest rays often reaching beyond tip of last ray if deflexed. Anat originating about
under middle of dorsal base, its base 3.0—3.5 in SL. Petvic small, reaching only a little
more than halfway to origin of anal, inserted about equidistant between base of pectoral
and origin of anal. Pecrorat fin reaching to or slightly beyond base of pelvic in small
specimens, often not quite to pelvic in large examples, 1.7—1.9 in head. AxILLary
SCALE of pectoral broad, about half as long as fin, 3.0-3.9 in head.
Color. In alcohol, pale; side of head and lower 75°/, of body silvery. Young with
a silvery lateral band, becoming diffuse in specimens about 100 mm long and disap-
pearing entirely with age. Middle of back with dark streak (missing in some, probably
due to fading). Fins unmarked.
Size. The largest specimen examined is 170 mm TL, but 300 mm has been re-
ported (7: 413).
Relation to Man. Its commercial importance is not significant. It occurs oc-
casionally in the Colén market (Panama) and, according to report, in the Havana
market (Cuba).
Range. Its range extends from the West Indies to Panama and at least as far
southward as Pernambuco (Recife), Brazil, probably to Rio de Janeiro and Sao Paulo.
All specimens studied were apparently taken in salt or brackish water, with one
exception; one specimen (CAS 9398) from Lake Rogagua, Bolivia, seems to be this
species.
Synonyms and References:
Engraulis clupeoides Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1839: 388 (orig. descr.; type local. Pernam-
buco, Brazil).
Engraulis productus Poey, Repert. Fisico-Nat. Cuba, 2, 1866: 380 (orig. descr.; type local. Matanzas, Cuba;
type MCZ 17961); Poey, Repert. Fisico-Nat. Cuba, 2, 1868: 423 (Havana, Cuba); Giinther, Cat. Fish.
Brit. Mus., 7, 1868: 388 (descr., Cuba and Jamaica); Tortonese, Bull. Mus. Zool. Anat. comp. Torino,
(3) 47, 1929: 6 (diagn., Puerto Cabello, Venezuela).
Engraulis surinamensis Steindachner (not of Bleeker), Ichthyol. Beitr., 5, 1879: 55 (descr., Bahia, Brazil).
Stolephorus clupeoides Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 447 (descr., range, 8. surina-
mensis probably incorrectly put in synon.; see Cetengraulis edentulus, p. 245); Eigenmann, Rep. Princeton
Exped. Patagonia, 1896-1899, 3 (4), 1910; 45 (Surinam to Rio Grande do Sul; 8. surizamensis Bleeker
apparently incorrectly put in synon.).
Stolephorus productus Jordan and Evermann, Bull. U. S. nat. Mus., 47 (1), 1896: 447 (descr., Cuba and Jamaica) ;
Jordan and Rutter, Proc. Acad. nat. Sci. Philad., 1897: 96 (descr., Jamaica); Evermann and Marsh,
Bull. U.S. Fish Comm., 20 (1), 1900: go (descr., Palo Seco, and Ponce, Puerto Rico); Nichols, Bull.
Amer. Mus. nat. Hist., 37, 1912: 182 (“Havana market”).
Anchovia clupeoides Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 211 (Santo Domingo and Rio Seco, Puerto
Rico; £. productus Poey put in synon.); Starks, Stanf. Univ. Publ. Biol., 1913: 9 (notes, Lake Papary,
Natal, Brazil); Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 412 (descr., range, synon.,
partly incorrect); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 50 (range,
synon., partly incorrect); Fowler, Proc. Acad. nat. Sci. Philad., 83, 1931: 393 (diagn., Quaima R., Tri-
nidad); Howell-Rivero, Bull. Mus. comp. Zool. Harv., 82, 1938: 172 (synon.; the larger of two specimens
158 Memoir Sears Foundation for Marine Research
[MCZ 17961, Poey’s 36] stated to be holotype); Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2),
1943: 27, fig. 9 (descr., range, synon.); Schultz, Proc. U.S. nat. Mus., 99, 1949: 39 (synon., locals.
listed).
Anchovia producta Meek and Hildebrand, Field Mus. Publ., Zool., 15 (1), 1923: 210 (descr., Mindi Cut,
Colén [market], and Porto Bello, Panama); Nichols, Ann. N. Y. Acad. Sci., To (2), 1929: 206 (diagn.,
range, “not uncommon about Puerto Rico’).
Anchovia macrolepidota Fowler (not of Kner and Steindachner), Proc. Acad. nat. Sci. Philad., 1917: 130 (two
from Colén, Panama, found by present writer to be 4. c/upeoides).
Anchoviella clupeoides Fowler, Arqu. Zool. estad. Sao Paulo, 3 (6), 1941: 134 (Brazilian rec.).
Doubtful References:
Stolephorus productus Schreiner and Ribeiro, Arch. Mus. nac., Rio de J., 12, 1903: 93 (Rio de Janeiro, without
comment).
Stolephorus clupeoides Eigenmann and Norris, Rev. Mus. paul., 4, 1904: 360 (one from Sao Paulo, Brazil;
proportions given indicate ident. probably not correct).
Anchovia nigra Schultz 1949
La anchoa
Figure 26
Study Material. Many paratypes, 14-76 mm TL (12-60 mm SL), and the type,
125mm TL (95 mm SL), from Lake Maracaibo and its tributary streams.
Distinctive Characters. See Relationship and Variation, p. 161.
Description. Proportional dimensions in per cent of standard length, not including
small examples regarded as juveniles because they would distort the picture; dimen-
sions and counts based on 11 or more specimens from Study Material, 50mm TL
and over.
Body: depth 21-28.9. Pectoral fin: length 16-18.
Head: length 23-31.3.
Snout: length 4.15-4.3. Scales: ca. 40-48.
Eye: diameter 6.1-8.3. Gill rakers: ca. 70+140 in adults, fewer
Postorbital: distance 15-19. in young.
Maxillary: length 18.5—24. Fin rays: dorsal 12-14; anal 29-31;
Mandible: length 17-22. pectoral 12 or 13.
Anal fin: length of base 26-31. Vertebrae: 39-41 (18 specimens).
Bopy strongly compressed, rather deep, its depth increasing with age, 3.5—-4.75
in SL in specimens 60-125 mm TL, about 5.5 in specimens around 25 mm. Heap
3.2-4.35, its depth scarcely as great as length of head without snout. SnouT short,
projecting 66°/, of its length beyond mandible, 6.5—7.1 in head. Eye 3.7—4.3. Posr-
orBITAL length 5.25—6.7. Maxittary abruptly pointed, reaching to or a little beyond
joint of mandible, 1.25-1.4 in head. Manprse 1.4—1.5. CHEEK moderately long and
3. Dr. Leonard P. Schultz has kindly placed his data on this species in my hands. The proportions and counts used
in the Description are his and mine.
Fishes of the Western North Atlantic 159
narrow, about equal to length of snout and eye in adults, its posterior angle approxi-
mately 30°. Git rakers at angle fully as long as eye in adults, proportionately
shorter in young.
Dorsat fin moderately high anteriorly, the longest rays reaching to or beyond tip
of last ray if deflexed, its origin somewhat variable, generally a little nearer to base of
caudal than the tip of snout. Anat fin moderately long, its origin somewhat variable,
sometimes nearly under middle of dorsal base, more frequently anterior to this point,
its base generally about equal to length of head, 3.2-3.85 in SL. Petvic fin small,
Ficure 26. Axchovia nigra, type specimen, 125 mm TL, from Maracaibo Basin, Venezuela, USNM 12176r.
Drawn by A. M. Awl.
reaching a little more than halfway to origin of anal, inserted a little nearer to base of
pectoral than to origin of anal. Pecrorat fin rather large, reaching well beyond base
of pelvic, 1.6—1.9 in head. Axittary scaLe of pectoral fin broad at base, variable in
length, usually reaching to middle of fin or beyond, 3.4—4.1 in head.
Color. In alcohol, back a pale straw color. Sides silvery. Silvery lateral band in
specimens up to 75 mm TL, the band very narrow in 40-mm specimens, becoming
proportionately broader and less well defined ventrally, diffused with silvery color on
lower part of sides in largest example; dark streak present on middle of back in large
examples but scarcely evident in small ones in which the dark punctulations that later
form the band remain separate; scattered punctulations extending down to silvery lateral
band. Dorsal and caudal fins with dark points; other fins plain translucent; small spec-
imens, 75 mm TL and under, with dark dots along base of anal fin and on ventral
surface of caudal peduncle.
Size. The largest specimen examined, the type (USNM 121761), is 125 mm
(5 in.) TL or 95 mm SL. Since a length of 300mm (12 in.) has been reported
for its near relative, clupeoides, it may be assumed that larger examples of xigra will
be found.
160 Memoir Sears Foundation Jor Marine Research
Development. The smallest specimen, only 14 mm, is very slender, its depth 9.15
and the head 4.0 times in SL. In this specimen the mouth is nearly terminal, the lower
jaw is very thin, the gape ends under the eye, and the maxillary is imperfectly developed.
The convoluted intestine, which is usual in larval herring and herring-like fishes, is
externally visible. All fins, however, are sufficiently developed to have at least rudimen-
tary rays. The body is pale, no doubt transparent in life, as general pigmentation has
not taken place. The juvenile markings consist of dark dots, which, exclusive of a few
spots at the nape, are on the lower part of the body; and a few less definite ones are on
the side of the head. An elongate median streak extends backward from the isthmus;
also there are spots along each side of the ventral edge of the abdomen that become
darker and more concentrated along the base of the anal and are reduced to a single
row on the midventral line of the caudal peduncle. A row of indefinite dark dots ex-
tends along the lateral side of the tail from above the anal base to the caudal base; there
are a few more dark dots on the caudal fin.
The body of a fish 20 mm TL is proportionately much deeper than the 14-mm
example, as its depth is contained 5.5 times in SL. The proportion of the head does not
differ greatly from either the small or large ones described, being 3.75 in SL. Much
progress in the development of the mouth parts has been made, however, for the snout
is more pointed and it projects prominently beyond the lower jaw. The maxillary is
definitely visible and extends well beyond the posterior margin of the eye, where it
ends in a rather blunt point. The intestine is no longer visible externally. Pigmen-
tation on the lower part of the body remains about the same as in the smaller ex-
ample, but dorsally, especially on the median line of the back and on the head, the
dark dots are more numerous.
The advances in development with growth between 20-30 mm are not great. The
increase in proportionate depth has continued, with the depth now 5.0 times in SL.
The snout has become more pointed and projects somewhat more strongly, about as in
adults. Some of the gill rakers are developed, 10 + 20 having been counted in one spec-
imen. Pigmentation has advanced somewhat, as dark dots on the back and on the head
have become larger and more numerous. The dark line behind the isthmus has virtually
disappeared. The dark spots on the side of the abdomen, along the base of the anal,
and on the ventral side of the caudal peduncle have become less distinct. A few elongate
dark markings at the base of the outer rays of both lobes of the caudal, already present
in the 20-mm specimen, have become more distinct.
In a fish 40 mm long, the depth is contained 4.5 times in SL, and the head 3.5
times. The cheek and opercular bones are well outlined. And the maxillary, which
now reaches the margin of the opercle, is provided with minute teeth along its free
margin. The gill rakers on the first arch are so numerous and close-set that they are
difficult to count; about 35-45 seem to be present. Pigmentation continues to advance;
the dark dots on the back have become more numerous and those on the lower parts,
exclusive of the ones along the anal base and on the peduncle behind the anal, have
disappeared. The upper parts of the body are slightly straw-colored and the sides
Fishes of the Western North Atlantic 161
of the head are bright silvery; the lower part of the body shows a somewhat silvery
sheen, and there is slight evidence of a silvery lateral band on the anterior half of
the body.
A specimen 70 mm long is shaped essentially like the type, which is 125 mm TL;
the body is proportionately about as deep in a 70-mm specimen as in the type; how-
ever, the gill rakers on the first arch are fewer, as 44 + 70 were counted, whereas about
70+140 seem to be present in the type. Furthermore, the smaller specimen retains
the silvery lateral band which the larger specimen has lost; it also retains the juvenile
dark markings along the base of the anal fin, which are missing in the larger one.
Relationship and Variation. This species is very close to c/upeoides, from which it
differs significantly in the number of vertebrae, the range in 17 specimens of xigra
being 39-41 and in 22 specimens of clupeoides 42 or 43. The anal fin may begin
under the middle of the dorsal base or well in advance of that point in zigra, consequently
it overlaps with c/upeoides, in which this fin begins about under the beginning of the
second third of the dorsal base. Furthermore, xigra generally has fewer rays in the anal
fin than clupeoides, 28-32 (av. around 30) in 38 specimens, and 30-35 (av. 33) in
58 specimens of c/upeoides. The lower average number of rays in the anal of vigra is
reflected in its somewhat shorter base, 3.2—3.85 in SL in 38 specimens of xigra, but
3.0-3.5 in 32 specimens of c/upeoides. The pectoral fin of nigra extends well beyond the
base of the pelvic fin in smaller specimens and only slightly beyond this point in the
largest example, being variable with age in this respect; in c/upevides the pectoral gener-
ally does not extend quite so far back as the pelvic fin; nevertheless, enough overlapping
occurs so that the character is not diagnostic. However, the place of insertion of the
pelvic fin generally is diagnostic; in igra it is nearly always inserted somewhat nearer
to the base of the upper pectoral rays than to the origin of the anal, whereas in c/upeoides
its insertion generally is almost exactly equidistant between the two points mentioned.
Finally, the snout projects about 66°/, of its length beyond the mandible in xigra, and
only about half of its length beyond the mandible in c/upeoides.
A. nigra is also closely related to A. rastralis of the Pacific coast of tropical America.
But in xigra the pelvic fins are inserted somewhat nearer to the pectoral base than to
the anal origin, whereas in rastralis they are inserted notably nearer to the anal origin
than to the pectoral base. The anal in xigra begins at, or more usually in advance of,
the vertical from the middle of the dorsal base, whereas in rastra/is it generally begins
at or behind this point; and zigra has 39-41 vertebrae, whereas rostralis (like clupeoides)
has 42 or 43.
Habitat and Range. A. nigra is known only from the fresh or slightly brackish
water of Lake Maracaibo, Venezuela, and from the lower sections of its tributary
streams. Apparently it does not enter the salt water of the Gulf of Venezuela, where
clupeoides has been taken.
Reference:
Anchovia nigra Schultz, Proc. U.S. nat. Mus., 99, 1949: 39, fig. 4 (orig. descr.; type local. Lake Maracaibo,
Venezuela; type USNM 121761).
162 Memoir Sears Foundation for Marine Research
Genus Anchoa Jordan and Evermann 1927
Anchoa Jordan and Evermann, Proc. Calif. Acad. Sci., 4 (16), 1927: 501; type species by original designation,
Engraulis compressus Girard 1858.
Characters. Bopy usually quite elongate, moderately to strongly compressed.
Maxittary long, reaching to, or more usually beyond, joint of mandible, frequently
to or nearly to margin of opercle, always more or less pointed (except in very young).
GILL RAKERS not numerous, seldom more than 22 on upper limb or more than 28 on
lower limb except in cubana among Atlantic species, not increasing in number with
age and growth. Anat origin under base of dorsal, rarely wholly behind it. VerrrBrarE
38-44, rarely 45 (in Atlantic species).
Remarks. This genus differs from Axchoviella principally in its longer maxillary,
which is somewhat sharply pointed rather than rounded or squared distally. From
Anchovia it differs in the fewer gill rakers, the number of which does not increase with
age. It differs principally from Exgrau/is (not represented in the western North Atlantic)
in the more compressed body (also frequently deeper), and usually in the fewer verte-
brae.
Species and Range. This genus contains many species that range the Atlantic
from Massachusetts (rarely Nova Scotia) to Argentina, and the Pacific from California
to Peru. Some of the species enter brackish or almost fresh water, but none is known
to run far upstream.
Table I. Frequency Distribution of Anal Rays in Axchoa.
SESS INitumalb0 G0) [Pq =e
18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40
SPUMUPED occ ee eek NS). 2
GPRENLEUS oo ce ceeeeeeees Se ees Se ee ee ee
BREE ano oongneocenone =H eS SSS SS SS SS (et SS eee
UINULALIS. oe cece cece See 24 7 5 2————— — — — — —
CAYOTUM oe ceeeeeeeees 3 22 22 12 4 — — — — — — — —
pectoralis ....eeee eee 6 (SSeS Ss Se eee ee ee
mitchilli mitchilli....... — — — — — r 8 22 63 74 51 30 11 FY — — — — — — — — —
mitchilli diaphana ...... — — — — — 2 11 55 83 62 44 14 5 — — — — — — — — — —
POPUL so cddocobonas5ccn —— 1 818 18 302 ee — — — — —
TOLLE GOR BRO ODO ODOC — — — I 312 4m ee
GHOETOSLOM auras sietaleterielels ——— 5416 9 3— — —
TOUT? 06 snacdgeounacoc 2 93017 6 —— — — — — — — — — — — — — — — — —
(TUNA LES OAR OC —— 913 79 4 IH — —
IOLASDUT ET MPa ots oyerereis tale BU ————————————
LamprotaeNid .......0445 — 1 10 31 43 81 5614 3 — — — — — — — — — — — — — —
hepsetus hepsetus ........ 2t 28 4257 20 3 — — — — — — — — — __
hepsetus colonensis....... 341313 3 te — _ — __ __
LV OLEDIStar ter tenctetete sie veto <F — 279 § rem ee
GUYS popcicso cdOGORODO — 1 33 § 2
Fishes of the Western North Atlantic 163
Key to Species of the Western North Atlantic
1a. Anal fin with 25 rays or more (m. mitchilli rarely with 24); anal base 25-35.6°/,
of SL (pectoralis sometimes only 23.4°/o).
2a.
2D.
Origin of dorsal about an eye’s diameter nearer to tip of snout than to base
of caudal; anal with 30 rays or more.
3a. Origin of anal under or somewhat in advance of middle of dorsal base;
32 or more anal rays.
4a. Anal with 35-40 rays; pectoral fin generally reaching far beyond
base of pelvic fin, 5.0-5.7 in SL; margin of caudal black.
spinifer (Cuvier and Valenciennes) 1848, p. 167.
4b. Anal with 32 rays; pectoral fin not reaching far beyond base of pelvic
fin, 6.5 in SL; margin of caudal not black.
argenteus Schultz 1949, p. 169.
3b. Origin of anal behind dorsal base; 30 anal rays.
duodecim (Cope) 1869, p. 170.
Origin of dorsal nearer to base of caudal than to tip of snout; anal usually
with less than 30 rays (mitchilli and trinitatis sometimes with as many as 30).
sa. Origin of dorsal nearer to posterior margin of eye than to base of
caudal; longest rays of dorsal reaching to or beyond tip of last ray when
deflexed; pectoral usually with more than 12 rays (cayorum sometimes
with only 12).
6a. Depth of body 22.2-25.5°/) of SL; gill rakers usually 16-19 + 19—
22 (rarely only 15 on upper limb); anal base 29.4-33.4°/, of SL.
trinitatis (Fowler) 1915, p. 171.
6b. Depth of body 18.2-22.2°/, of SL; gill rakers 12-14+17-19
(cayorum rarely with 15 on upper limb, pectoralis occasionally with
20 on lower limb); anal base 23.5-29.4°/, of SL.
7a. Maxillary narrowly pointed, extending nearly or quite to margin
of opercle, 19.2—22.5°/, of SL; origin of anal well in advance
of middle of dorsal base; pectoral with 12 or 13 rays, generally
failing to reach base of pelvic fin by about half of diameter of
eye, 13.9-16.6°/, of SL. cayorum (Fowler) 1906, p. 173.
7b. Maxillary bluntly pointed, extending only slightly beyond
joint of mandible, 17.7—19.0°/, of SL; origin of anal generally
under middle of dorsal base; pectoral with 15-17 rays, reaching
nearly to base of pelvic fin, 17.8-18.8°/, of SL.
pectoralis Hildebrand 1943, p. 174.
5b. Origin of dorsal nearer to base of caudal than to posterior margin of
eye, very rarely equidistant; longest rays of dorsal usually failing to
reach tip of last ray when deflexed; pectoral with 11 or 12 rays.
mitchilli (Cuvier and Valenciennes) 1848, p. 176.
11*
164 Memoir Sears Foundation for Marine Research
8a. Greatest thickness of body usually exceeding depth of caudal
peduncle; depth 15.8-—25°/, of SL; pectoral often failing to reach
base of pelvic by diameter of eye, 12-18°/, of SL; vertebrae 39-44,
most frequently 42. m. mitchilli Hildebrand 1943, p. 176.
8b. Greatest thickness of body often less than depth of caudal peduncle;
depth 18.2-26.5°/, of SL; pectoral often extending to or nearly to
base of pelvic, 13-20°/, of SL; vertebrae 38—42, most frequently 40.
m. diaphana Hildebrand 1943, p. 176.
1b. Anal fin with less than 25 rays (/amprotaenia occasionally with 25 or 26 rays,
choerostoma rarely with 25); anal base 17.2-25°/, of SL (januaria sometimes 26).
ga. Pectoral without a long, filamentous ray (rarely with a slight filament in
lamprotaenia); origin of dorsal fin usually nearer to base of caudal than to
tip of snout, rarely equidistant.
10a. Origin of anal in advance of base of last dorsal ray; depth of head at
joint of mandible exceeding postorbital length; snout much shorter
than eye; cheek shorter than snout and eye combined; silvery lateral
band much narrower than eye.
11a. Gill rakers on lower limb 2 3—33, on upper limb 1 8-23 (rarely only 17).
12a. Depth of body 18.2-22°/, of SL; pectoral fin 14-18.5°/,
of SL; gill rakers rather robust, not closely crowded, not
difficult to count.
13a. Origin of anal under or near middle of dorsal base;
depth of body 20-22°/, of SL.
14a. Maxillary sharply pointed distally, extending
nearly to margin of opercle, 1.2-1.35 in head;
vertebrae 38—40 (rarely 41).
parva (Meek and Hildebrand) 1923, p. 181.
14b. Maxillary bluntly pointed distally, extending
only to or slightly beyond joint of mandible,
1.3-I.5 in head; vertebrae 41 or 42.
januaria (Steindachner) 1879, p. 183.
13b. Origin of anal far behind middle of dorsal base; depth
of body 18.2—21°/, of SL.
15a. Cheek as long as eye and fully 75°/, of snout
length; pectoral fin 15.6—-18.5°/, of SL; axillary
scale of pectoral about half of length of fin,
3.0-4.0 in head; anal with 22-24 rays.
choerostoma (Goode) 1874, p. 185.
15b. Cheek scarcely longer than eye; pectoral fin
14-14.8°/) of SL; axillary scale of pectoral fully
75°/, of length of fin, 1.9—2.75 in head; anal with
18-22 rays. tricolor (Agassiz) 1829, p. 186.
Fishes of the Western North Atlantic 165
12b. Depth of body 15-18°/, of SL; pectoral fin 12.5-13.7°/o
of SL; gill rakers very slender, close-set, often difficult to
count. cubana (Poey) 1868, p. 188.
11b. Gill rakers on lower limb usually 16-22 (rarely 23 in A. hepsetus),
on upper limb usually 14-20 (rarely 13 or 21).
16a. Depth of body 17.4-18.8°/, of SL; maxillary bluntly pointed
distally, its upper margin rounded, extending to joint of
mandible, 17.2-18.2°/, of SL; vertebrae 44 or 45.
ginsburgi Hildebrand 1943, p. 190.
16b. Depth of body 18.5-23°/, of SL; maxillary usually sharply
pointed distally, generally extending beyond joint of man-
dible, sometimes to or nearly to margin of opercle (mod-
erately short, rather blunt distally, and rounded above in
lamprotaenia), 19-24.5°/, of SL; vertebrae 39-43 (A. hep-
setus sometimes with 44).
17a. Longest rays of dorsal fin usually reaching well
beyond tip of last ray when deflexed; origin of anal
about under beginning of posterior third of dorsal
base; axillary scale of pectoral only a little more than
half of length of fin, 3.0—-3.6 in head; cheek fully as
long as eye and half of snout.
lamprotaenia Hildebrand 1943, p. 192.
17b. Longest rays of dorsal fin failing to reach tip of last
ray when deflexed; origin of anal usually about under
middle of dorsal base, sometimes posterior to middle;
axillary scale of pectoral 2.0-2.5 in head; cheek usually
not much longer than eye.
hepsetus (Linnaeus) 1758, p. 194.
18a. Pectoral fin often failing to reach base of pelvic
by diameter of eye; silvery lateral band usually
75°/, of width of eye.
h. hepsetus Hildebrand 1943, p. 194.
18 b. Pectoral fin frequently extending to or nearly to base
of pelvic; silvery lateral band often scarcely wider
than pupil. 4. colonensis Hildebrand 1943, p. 194.
10b. Origin of anal under or behind base of last dorsal ray; depth of head
at joint of mandible equal to postorbital length; snout scarcely shorter
than eye, 4.3-5.5 in head; cheek about as long as snout and eye
combined; silvery lateral band as broad as eye.
lyolepis (Evermann and Marsh) 1902, p. 200.
gb. Pectoral with a long filamentous ray; origin of dorsal fin usually nearer to
tip of snout than to base of caudal. filifera (Fowler) 1915, p. 202.
166 Memoir Sears Foundation Jor Marine Research
Table II. Frequency Distribution of Gill Rakers on Upper Limb of First Arch
in Anchoa.*
12 eS eA Se T Ge TO la 77)
UH DRA MeO BODCEMOGO DOT TD OC I 2am alate) esd 30 =
GT RORLCUS, 0 o's x 3 ahaie wien Bistsisioveiaiers —- —- — 1 — —
GUOGECIM ae viate since lsat —- — I I —
LHLMELATIS. cidierasslelonina orettorere arene —- —- — I I 7
COYOT UM. «wins 0° hevayntn, 6! eyatoret ave Es I 24 3 => =
(ABT spoon poocasosobnonene I 2 — =
UIECHILISEILILCHILIE tree eerie _ — B 4 63
mitchilli diaphana.......+..0005 —- —- — 9 5 wie
(NATH 0050860 6000000000R0000000 - -—- —- — —
VOTIEE s0a8n6ocacduOesuene non SS SSS SS
CROLL OSLOMGL nary leilets saya (a2 50 - - -—- =—- — I
EEGOLOL MATa eet sta oialove/ era) eevee ce vers\ ere —- -—- -—- -—- SS
GAUTIT8o ooo > HO ODO DORE DEBT ORORO - - -—- =—- — 2
IDUMSBUTD Sexe rolepevate\ uss oy ss ave: oitie\s/eyo\se — —- — I 2
WA TEDLOLACHEGY Mein Noty Nee stal acted ore = — Gp I GP FG} 6
REP SELUSHREPSEL US oe) foseielie i AenkeelAe — — i GY RA
epsetus COLOMENSIS .... 10 ee eneeee —- —- — 3 9 24
UES on 0 Bodo MOBO TOG ooR00056 - -—- — I 5 8
SOUL F ERB oe scaha ys cysks cyaeys msely Seats seats a - - -—- =—- — 5
* When the counts exceed the number of specimens listed in the text, the rakers were
Tior arches.
Table III. Frequency Distribution of Gill Rakers on Lower Limb of First Arch
in Anchoa.*
18
4
19
Lloro La wun | | 4 |
4
20 21
aS
eed UU Wezel rsal eT tT
22
Number of Gill Rakers ——————___—__
23
N
counted on both ante-
Number of Gill Rakers
i lo) LP TS Oy Ie IA Dey VL Wo) Be) OES WS) sf Gr gi BA
SPILL Cf ap stave nin latte chYe ria cicoemoTevetel eats At Oh AS a
AU RERLEUS HT ancrate arate ceili olera ate area Octo pe ee ee oe
MILD. on oacpowooobennacoobodooog ==] SS 9 Si gna es ee ee ee
LTE BIL ALIS Maar ctataratote vos sucvaiee wish ee ———— 1378 ~~~ ~~ ~~ —
GIT shooguaodbDoEscenoogodsobon 18 40 220. —
PECHOIDS 3330100500209000000000000 5¢ Seok SS
TAOANE TOI Oo ononcncponodnoDuSs = = SS 421 45 70 54 21 6 — — — — — — —
mitchilli diaphana .......00++00+0002 — — — — — 12 64 125 92 46 10 3 — — — — — — —
PATOE ov ccccccccc cece eneneeeeeeee ss =e ee — 71014 13 1 ————— —
WM UMUGTIG I caseniten eee — 4 4 5—————— —
WITOPUA oo oncoa0cboonnnoscoosce9 == SS — 1012 9 6 —— — — — — —
WICOLO occ ence eee eee e es See —— Ir 2025 8 2——— — —
GUAT Bis ooeaobowoabbobaeocécospjene = eS SS SS — eit () sess EY of 2 2
BILE cooonoggotonacscovendessess SSS SS 2 % 8 —
HA GONE A TATOO ONO 0 0 DOC OFOE — 2 2694 8231 8 5 ——— — — — — — — — —
BEDSELGSHREPSELUSHETA (oho tele oe eek ——— 92797 93 47 rt — — — — — — —
BEPSELUSNCOLOMEBSES a)ale « « vie1e\s\ e eieletere ee ———— 41525 4—— —
UNGER cost SASHES SOIT CUO SOOT ———— 41114 II § § 4-S— O e
HUPBE ososooosoco vandanbabdasooses ——=—— — 3030°92 Yee
* When the counts exceed the number of specimens listed in the text, the rakers were counted on both
anterior arches.
Fishes of the Western North Atlantic 167
Anchoa spinifer (Cuvier and Valenciennes) 1848
Sardine
Figure 27
Study Material. Many specimens, 30-175 mm TL: from the Atlantic at the
Gatun Locks, Canal Zone; Port-of-Spain, Trinidad; and Bahia, Cachoeira, and Santos,
Brazil; from the Pacific at Pedro Miguel and Miraflores locks, Canal Zone; Panama
Bay; and Guayaquil, Ecuador.
Ficure 27. Anchoa spinifer, go mm TL, from Gatun Locks, Canal Zone, USNM 127560. Drawn by Louella
E. Cable.
Distinctive Characters. The long anal fin, the dorsal’s high rays in the anterior part,
the dorsal’s forward position, the long low head and especially its prolonged postorbital
portion, the long narrow cheek, and the dark margin of the caudal fin distinguish this
species from all of its relatives. Large examples generally are proportionally deeper
than small ones, and generally Atlantic specimens are rather deeper than Pacific ones.
However, there is so much overlapping that, with the material at hand, specimens from
the opposite coasts cannot be separated by this character, nor seemingly by any other.
The pectoral fin also varies greatly in proportional length, but this also seems to be
an individual variation. Nevertheless, the comparison is not satisfactory because nearly
all of the Atlantic specimens are old and faded, not in good condition.
Description. Proportional dimensions in per cent of standard length, and counts,
based on 11 Atlantic specimens and 15 Pacific specimens, 66-175 mm TL.
Body: depth 19.2-25. Anal fin: \ength of base 33.1-36.6.
Head: length 25-27.8. Pectoral fin: length 17.5—20.
Snout: length 3.0-4.3.
Eye: diameter 4.5—-6.25. Scales: ca. 45.
Postorbital: distance 14.9-16.6. Gill rakers: 12-16+15-18.
Maxillary: length 20.8—23.2. Fin rays: dorsal 14-16; anal 35-40;
Mandible: length 19.6—20.3. pectoral 12-14.
Vertebrae: 43-45 (15 specimens).
168 Memoir Sears Foundation for Marine Research
Bopy quite elongate, rather strongly compressed, its depth 4.0-5.2 in SL, in-
creasing somewhat with age and growth. Heap 3.6—4.0 in SL, its depth scarcely
exceeding its postorbital length. Snour projecting nearly its full length beyond man-
dible, 6.6—7.5 in head. Eve small, 4.2-5.0. Posrorsirat part of head long, 1.5—-1.7
in head. Maxittary long and pointed, reaching to or nearly to margin of opercle,
I.1-1.25 in head. Manprsrze 1.22—1.4. Curek in large specimens much longer than
snout and eye, about equal to snout and eye in specimens about 80 mm TL; posterior
angle sharp, about 30°.
Dorsat fin high anteriorly, its longest ray reaching far beyond tip of last ray if
deflexed, its origin generally rather more than an eye’s diameter nearer to tip of snout
than to base of caudal. Anat 2.8—3.0 in SL, its origin generally somewhat in advance
of middle of dorsal base. Prtvic reaching about 66°/, of distance to anal, inserted
about equidistant between origin of anal and base of pectoral. Pecrorat large, reaching
to, and generally beyond, base of pelvic, 1.3-1.5 in head. AxiLLary scaLe of pectoral
broad, reaching somewhat beyond midlength of fin, 2.4—2.9 in head.
Color. In alcohol, pale above. Lower half of side silvery. Young sometimes with a
poorly defined silvery lateral band. Back with dusky punctulations. Fresh specimens
125mm TL and upward (taken in Pedro Miguel and Miraflores locks, Canal Zone)
bright orange, this color contrasting conspicuously with black margin of caudal fin.
Size. The largest specimen seen, about 175mm TL or 7 inches (caudal fin dam-
maged) and 138 mm SL, is probably near the maximum size attained.
Range and Habitat. The range in the Atlantic, so far as known, extends from
Panama to Santos, Brazil. On the Pacific side it is known from Panama Bay to Guaya-
quil, Ecuador. The West Indies has been included in the general range (7: 410), but
I have seen no specimens from that locality and have found no other record. In
rather extensive collecting on the Atlantic coast of Panama (during the dry seasons),
only a single small specimen was secured; but on the Pacific side it was exceedingly
abundant in Pedro Miguel and Miraflores locks when these were drained in 1937;
however, it was not seen elsewhere during four different seasons of collecting. The
water ranges from near oceanic salinity in the lower flights of Miraflores Locks (at sea
level) to fresh in Pedro Miguel Locks.
Synonyms and References:
Engraulis spinifer Cuvier and Valenciennes, Hist. Nat. Poiss., 2Z, 1848: 39 (orig. descr.; type local. Cayenne,
French Guiana); Giinther, Cat. Fish. Brit. Mus., 7, 1868: 394 (descr., type local.); Steindachner,
Ichthyol. Beitr., 8, 1879: 58 (descr.; Guiana; Bahia and Cachoeira, Brazil; w. coast of Panama).
Stolephorus spinifer Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 448 (descr., range); Eigenmann,
Mem. Carneg. Mus., 5, 1912: 449 (synon., descr., Georgetown, British Guiana); Puyo, Bull. Soc. Hist.
nat. Toulouse, 70, 1936: 65, 163 (diagn., habitat French Guiana); Puyo, Bull. Soc. Hist. nat. Toulouse,
50, 1945: 105, fig. 3 (descr., French Guiana); Puyo, Faune Emp. Frang., 72, Poiss. Guyane Frang.,
1949: 156, fig. 79 (descr., French Guiana).
Anchovia spinifera Gilbert and Starks, Mem. Cal. Acad. Sci., 4, 1904: 46, pl. 8, fig. 15 (descr. based on two
from Panama Bay); Meek and Hildebrand, Field Mus. Publ., Zool., 75 (1), 1923: 207 (synon., descr.,
range); von Ihering, Rev. Industr. Anim., Anno 1 (3), 1930: 233 (ref., diagn.); Hildebrand, Zoologica,
N. Y., 24 (1), 1939: 25, 30, 36 (Panama Canal locks).
Fishes of the Western North Atlantic 169
Anchoviella spinifera Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 409 (descr., Cachoeira, Bra-
zil; ““Panama”’ presumably Pacific coast; ‘““West Indies” in range); Jordan, Evermann, and Clark, Rep.
U.S. Comm. Fish. (1928), 2, 1930: 50 (range); Fowler, Arqu. Zool. estad. Sao Paulo, 3, 1941: 134
(refs., Brazil).
Anchoa spinifer Hildebrand, Bull. Bingham oceanogr. Coll., § (2), 1943: 38, fig. 10 (synon., descr., range,
local abund.).
Anchoa argenteus Schultz 1949
Figure 28
Study Material. Known only from the type, ca. 115 mm TL (caudal fin damaged),
g4 mm SL, from Lake Maracaibo, off Pueblo Viejo, Venezuela.
Distinctive Characters. This species is closely related to 4. spinifer, with which it
agrees in the shape and position of the dorsal fin, the long low head, the long narrow
cheek, and in many other respects. But it differs significantly from that species (so far
as can be determined from a single specimen) in having fewer anal rays, in a slightly
greater number of gill rakers on the lower limb, in the longer mandible, in the shorter
pectoral fins, and in the absence of a dark margin on the caudal fin.
Ficure 28. Axchoa argenteus, type specimen, 94mm TL, from Maracaibo Basin, Venezuelaj USNM 121777.
Drawn by A. M. Awl.
Description. Proportional dimensions in per cent of standard length, and counts,
based on the type specimen, 94mm SL.
Body: depth 22.5. Anal fin: length of base 32.
Head: length 26.5. Pectoral fin: length 15.5.
Snout: length 4.25.
Eye: diameter 5.55. Scales: ca. 43.
Postorbital: distance 18.5. Gill rakers: 14419.
Maxillary: length 23.5. Fin rays: dorsal 16; anal 32; pectoral 13.
Mandible: \ength 20.6.
E7O Memoir Sears Foundation for Marine Research
Bony elongate, rather strongly compressed, its depth 4.5 in SL. Heap 3.75, its
depth a little greater than its postorbital length. Snour shorter than eye, projecting
much more than half of its length beyond tip of mandible, 6.25 in head. Eye small, 4.8.
PosrorsitaL length 5.4. MaxiLLtary moderately pointed, reaching margin of opercle,
1.15 in head. ManprsLe 1.3. CHEEK narrow, much longer than snout and eye, its
posterior angle about 30°. Grit RakeRs at angle of first arch about 66°/, of length
of eye, with strong serrae on inner edge, somewhat expanded at tips.
Dorsat fin moderately elevated anteriorly, its margin nearly straight, its longest
rays reaching well beyond the last ray if deflexed, its origin about an eye’s diameter
nearer to tip of snout than to base of caudal. Anat fin fairly long, its origin under middle
of dorsal base, its base notably longer than head, 3.1 in SL. Petvic fin reaching a little
more than halfway to origin of anal, inserted a little nearer to base of pectoral than to
origin of anal, 3.1 in head. Pecrorat fin reaching beyond base of pelvic by about half
of diameter of eye, 6.5 in SL, 1.7 in head. Axitiary scaLe of pectoral extending to
beginning of distal third of fin, 2.55 in head.
Color. In alcohol, somewhat straw-colored on back, changing rather abruptly to
silvery on the side at level of upper margin of eye; back with dusky punctulations, these
extending onto basal 66°/) or so of dorsal fin, and onto outer rays of caudal fin.
Range. Known only from Lake Maracaibo, off Pueblo Viejo, Venezuela.
Reference:
Anchoa argenteus Schultz, Proc. U.S. nat. Mus., 99, 1949: 45, fig. 5 (orig. descr.; type local. Lake Maracaibo,
off Pueblo Viejo, Venezuela; type USNM 121777).
Anchoa duodecim (Cope) 1869
New Jersey Anchovy
Study Material. Known only from the type, 97 mm long (77 mm SL), taken at
Beesley’s Point, New Jersey, ANSP 1363.
Distinctive Characters. This species has been considered identical with 4. mitchilli.
However, it differs strikingly from other local forms in its deep body, high dorsal fin,
the positions of the dorsal and anal fins, and in several other respects. It is of interest
that the species has never reappeared in collections.
Description. Proportional dimensions in per cent of standard length, and counts,
based on the type specimen, 77mm SL.
Body: depth 25. Anal fin: length of base 25.
Head: length 27. Pectoral fin: length 18.2.
Snout: length 4.7.
Eye: diameter 5.8. Scales: partly lost, ca. 38.
Postorbital: distance 16.3. Gill rakers: 14 or 15 +20.
Maxillary: length 22. Fin rays: dorsal 14; anal 30; pectoral
Mandible: length 21. 12,
Fishes of the Western North Atlantic 071
Bopy rather short, deep, and well compressed; dorsal profile more strongly convex
than ventral profile. Heap short, deep, its depth equal to its postorbital length and half
of eye. Snour rather short, projecting a little less than a third of its length beyond
mandible, 5.8 in head. Eye small, 4.7. Posrorsirat 1.6 in head. Maxrtrary fairly
short, reaching joint of mandible, broad and abruptly pointed, 1.25 in head. Man-
DIBLE I.3. CHEEK as long as snout and eye, its posterior angle about 35°. GILL RAKERS
at angle nearly as long as eye.
Dorsat fin high anteriorly, its longest rays reaching far beyond tip of last ray if
deflexed, its origin an eye’s diameter nearer to tip of snout than to base of caudal. Anat
rather long and low, its origin half of an eye’s diameter posterior to end of dorsal base,
and about equidistant between base of caudal and base of pectoral, its base 4.0 in SL.
Petvic fin large, shorter than pectoral by diameter of pupil, extending about 66 °/) of
the distance to origin of anal, inserted a little nearer to joint of mandible than to
origin of anal. Pecrorat reaching base of pelvic, 1.5 in head. AxiLLary scaLe of pec-
toral a little more than half of the fin length, 2.7 in head.
Color. Old preserved specimen grayish brown above. Lower two-thirds of side
silvery; no lateral band and no punctulations visible.
Range. Known only from Beesley’s Point, New Jersey.
Synonyms and References:
Engraulis duodecim Cope, Trans. Amer. philos. Soc., 13 (2), 1869: 405 (orig. descr.; type local. Beesley’s
Point, New Jersey; type ANSP 1363).
Anchovia duodecim Fowler, Rep. N. J. St. Mus. (1905), 1906: 111 (descr.); Fowler, Proc. Acad. nat. Sci.
Philad., 63, 1911: 219; Fowler, Proc. biol. Soc. Wash., 33, 1920: 148.
Anchoa duodecim Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 46 (synon., descr. based on type).
Anchoa trinitatis (Fowler) 1915
Figure 29
Study Material. A total of 19 specimens, including the type, from Trinidad and
Venezuela, ANSP, UMMZ, USNM.
Description. Proportional dimensions in per cent of standard length, and counts,
based on 10 specimens from Study Material, 55-110 mm TL.
Body: depth 22-2 5.6. Anal fin: length of base 29.4—33.2.
Head: length 24.3-27. Pectoral fin: length 16.6—19.
Snout: length 3.9-4.35.
Eye: diameter 6.75—7.6. Scales: ca. 38-42.
Postorbital: distance 11.7—13.3. Gill rakers: 15-19 + 18-22.
Maxillary: length 16.6—22. Fin rays: dorsal 13 or 14; anal 26—30;
Mandible: \ength 16.4-19. pectoral 13 or 14.
Vertebrae: 41 (1 specimen).
Lg 2 Memoir Sears Foundation for Marine Research
Bopy rather strongly compressed, its greatest thickness about equal to depth of
caudal peduncle, its depth 3.9-4.5 in SL. Heap short, 3.7—4.1 in SL, its depth equal
to its postorbital length and about 66°/, of diameter of eye. SNout extending about
66°/, of its length beyond mandible, 5.7-6.6 in head. Eve 3.4—3.7. Posrorsira
1.9—2.1 in head. Maxittary pointed, extending nearly to margin of opercle, 1.2-1.4
in head. Manprse 1.4—1.53. Cueek about as long as eye and half of snout, its
posterior angle about 40°.
Dorsat fin moderately high anteriorly, its longest rays reaching beyond tip of
last ray if deflexed, its origin equidistant between base of caudal and a point at anterior
half of eye. Anat origin under, or somewhat in advance of, middle of dorsal base, its
base 3.0-3.4 in SL. Petvic reaching fully halfway to anal, inserted a little nearer to
origin of anal than to base of pectoral. Pecrorat sometimes not quite reaching to pelvic,
1.35-1.52 in head. Axittary scaLe of pectoral rather broad at base, scarcely extending
to vertical at midlength of fin, 2.8—3.6 in head.
Ficure 29. Axchoa trinitatis, type specimen, from Port-of-Spain, Trinidad, ANSP 45083, modified after
Fowler (1915: 527, fig. 3). Drawn by Alice C. Mullen.
Color. In alcohol, pale above. Side of head and lower part of body silvery. Silvery
lateral band faint, narrower than pupil, missing in specimens presumably preserved
originally in formalin. Back and upper surface of head with dusky punctulations, some-
times forming two almost continuous lines posterior to dorsal fin.
Size. The largest individual seen is 110 mm TL.
Range. So far as known, this species is confined to four localities: Port-of-Spain
and Vessigney, Trinidad; Laguna de Tacarigua (on coast 85 miles east of Caracas),
and Rio Cafio de Sagua, Sinamaica, Venezuela.
Synonyms and References:
Anchovia trinitatis Fowler, Proc. Acad. nat. Sci. Philad., 1915: 527, fig. 3 (orig. descr.; type local. Port-of-Spain,
Trinidad; type ANSP 45083).
Fishes of the Western North Atlantic 173
Anchoviella trinitatis Fowler, Proc. Acad. nat. Sci. Philad., 83, 1931: 392 (Vessigney, Trinidad).
Anchoa trinitatis Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 96, fig. 40 (descr. cf. several
related species, range); Schultz, Proc. U.S. nat. Mus., 99, 1949: 44 (synon., specimens and local. listed;
counts of rays, gill rakers, vert., and scales).
Anchoa cayorum (Fowler) 1906
Key Anchovy, Manjuia (Cuba)
Figure 30
Study Material. A total of 26 or more specimens, 50-85 mm TL, from Tortugas,
Florida; Cabafias Bay, Cuba; Cozumel Island, off Yucatan, México; Belize, British
Honduras; and Hailer’s Rock, Florida Keys (type and paratype).
Ficure 30. Azchoa cayorum, 85 mm TL, 68 mm SL, from Cabafias Bay, Cuba, USNM 82341, with pectoral
fin enlarged. Drawn by Louella E. Cable.
Distinctive Characters. Museum specimens generally have been identified with
A. mitchilli, presumably because of the long anal fin, but cayorum differs in the larger
size it attains (largest seen 85 mm), in the notably fewer gill rakers, and in the much
broader and more prominent silvery lateral band.
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens, 50-85 mm TL.
Body: depth 18-21. Snout: length 4.3-5.0.
Head: length 25-28.5. Eye: diameter 5.8—7.4.
174 Memoir Sears Foundation for Marine Research
Postorbital: distance 13.3-15.3. Scales: ca. 39-42.
Maxillary: length 19.2—22.5. Gill rakers: 13-15 +15—-17 (rarely 18).
Mandible: length 18.5—20. Fin rays: dorsal 13-15; anal 26-28
Anal fin: length of base 25—29.3. (rarely 25 or 29); pectoral 12 or 13.
Pectoral fin: length 13.9-16.6. Vertebrae: 43 (3 specimens).
Bopy slender, moderately compressed, its depth 4.75—5.5 in SL. Heap 3.5—4.0
in SL, its depth equal to its postorbital length and about half of eye. Snour pro-
jecting about 66°/, of its length beyond tip of mandible, 5.0—6.0 in head. Eye 3.5—
4.25. PosrorpiTaL 1.65—2.0 in head. Maxitiary long and pointed, its upper margin
straight, reaching nearly or quite to margin of opercle, 1.1-1.35 in head. MAnpIBLE
1.2-1.4. CHEEK equal to, or a little longer than, eye and 75°/, of snout, its posterior
angle rather sharp, about 40°.
Dorsat fin with the longest rays reaching well beyond tip of last ray if deflexed,
its origin about equidistant between base of caudal and anterior margin of eye. ANAL
fin with origin under beginning of second fourth of dorsal, its base 3.4—4.0 in SL.
Pexvic inserted somewhat nearer to base of pectoral than to origin of anal. PecroraL
falcate, sometimes failing to reach base of pelvic by diameter of pupil, occasionally
reaching nearly to base of pelvic, the first (uppermost) ray generally as long as the
second and about twice the last ray, 1.5-1.9 in head. AxiLLary scaLe of pectoral
scarcely reaching beyond midlength of pectoral, 3.0-3.5 in head.
Color. In alcohol, pale. Side of head silvery. Silvery lateral band bright, about as
wide as eye.
Range and Habitat. The range extends from the Florida Keys, where it is appar-
ently rather rare, to Cuba, Yucatan, and British Honduras. The specimens examined
were apparently taken in shallow water, near the shore.
Synonyms and References:
Stolephorus mitchilli Bean, Bull. U. S. Fish Comm. (1888), 8, 1890: 206 (Cozumel I., off Yucatan, México;
specimens re-exam. and found to be 4. cayorum).
Anchovia choerostoma cayorum Fowler, Proc. Acad. nat. Sci. Philad., 58, 1906: 85, fig. 4 (orig. descr.; type local.
Hailer’s Rock, Florida Keys; type ANSP 30613).
Anchovia cayorum Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 219 (types relisted).
Anchoviella cayorum Jordan, Evermann and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 49 (synon.).
Anchoa cayorum Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 50, fig. 17 (synon., descr., relation.,
range, type material re-exam.).
Anchoa pectoralis Hildebrand 1943
Study Material. Only the type material, 64 specimens, 43-65 mm TL, from
Vigia, Brazil, MCZ 18004, listed for “Brazil, Louis Agassiz.”
Distinctive Characters. This species is closely related to 4. mitchilli diaphana,
A. parva, and A. januaria, from all of which it differs in the more numerous pectoral
rays. Other differences are evident in the descriptions and tables.
Fishes of the Western North Atlantic iii
Description. Proportional dimensions in per cent of standard length based on
7 specimens, the counts based on many more. As the state of preservation is not very
good, some of the proportions may not be entirely true.
Body: depth 19.5—23.0. Anal fin: length of base 23.4-25.6.
Head: length 23.8-26.2. Pectoral fin: length 17.8-18.8.
Snout: length 3.35—4.16.
Eye: diameter 6.7—8.0. Scales: ca. 35—40.
Postorbital: distance 12.9—13.7. Gill rakers: 13 or 14+17—-19.
Maxillary: length 17.7-19.0. Fin rays: dorsal 14-16; anal 25-27; pec-
Mandible: length 15.8—17.6. toral 15-17 (most frequently 16).
Vertebrae: 42 (2 specimens).
Bopy moderately slender, strongly compressed, its depth 4.35—5.1 in SL. Heap
3.8-4.2 in SL, its depth nearly equal to its length without snout. SNour short, pro-
jecting about half of its length beyond mandible, 6.0—7.0 in head. Eye 3.2—3.7. Post-
ORBITAL 1.75—2.0 in head. Maxittary bluntly pointed, its upper margin rounded,
reaching somewhat beyond joint of mandible, 1.2-1.4 in head. Manpiste 1.4—1.6.
Cuzek about as long as eye, its posterior angle broad, about 50°.
Dorsat fin with longest rays reaching to, and occasionally beyond, tip of last ray if
deflexed, its origin generally a little nearer to base of caudal than to middle of eye. ANaL
with origin generally under middle of dorsal, its base 3.9—-4.25 in SL. Petvic reaching
only about halfway to anal, inserted equidistant between origin of anal and base of
pectoral, or slightly nearer the former. Prcrorat falcate, reaching nearly to base of
pelvic, 1.3-1.4 in head. AxiLLary scaLe of pectoral pointed, about 66°/, of length
of fin, 2.5 in head.
Color. Old specimens in alcohol, brownish silvery. A prominent silvery lateral
band present, about as wide as eye. Back with dark dots, in more or less distinct
longitudinal rows. Base of anal with black dots; caudal with dark dots and with a dark
margin.
Range. Known only from the type material from Vigia, Brazil, collected in 1865
or 1866 by the Thayer Expedition of Harvard University. [Stieler’s Atlas lists the
name Vigia for two places in Brazil, one at the mouth of Rio Para (Tocantins), the other
well inland on Rio Jequitinhonha in the Province of Minas Gerais.—y.u.0.].
Synonyms and References:
Anchoviella mitchilli Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 405 (descr., distr., MCZ
18004).
Anchoa pectoralis Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 52, fig. 18 (orig. descr.; type local.
Vigia, Brazil; type MCZ 35276).
176 Memoir Sears Foundation for Marine Research
Anchoa mitchilli (Cuvier and Valenciennes) 1848
Bay Anchovy
Figures 31, 32
Study Material. A total of 100 specimens, 52-102 mm TL; for characters con-
sidered important, such as body depth, pectoral length, number of vertebrae, gill
rakers, and anal rays, many more were used; taken at many places from Woods Hole,
Massachusetts, to Yucatan, México.
Distinctive Characters. The nearest relative is 4. parva, a southern species that
Ficure 31. Axchoa mitchilli mitchilli, 85 mm TL, from Woods Hole, Massachusetts, USNM 125582. Drawn
by Louella E. Cable.
occurs in the West Indies and on the mainland coast from Panama to Venezuela. This
relationship is discussed in the account dealing with parva (p. 181).
Description. Proportional dimensions in per cent of standard length, and counts,
based on Study Material.
Body: depth 16-27. Pectoral fin: length 12-20.
Head: length 22—26.5.
Snout: length 3.7-7.25. Scales: (generally lost in preserved speci-
Eye: diameter 5.8—8.2. mens) ca. 38—44.
Postorbital: distance 11-13.5. Gill rakers: 1§—-19 + 20-26.
Maxillary: length 17.5—21. Fin rays: dorsal 14-16; anal 24-30;
Mandible: length 16.5-18.5. pectoral 11 or 12.
Anal fin: \ength of base 25-30. Vertebrae: 38—44 (1,233 specimens).
Bopy rather slender, moderately compressed, its greatest thickness usually ex-
ceeding the depth of caudal peduncle, its depth 3.65—6.3 in SL. Heap 3.75—4.5, its
depth equal to its length without snout and fourth of eye. Sout short, projecting not
more than a fourth of its length beyond tip of mandible, 5.0—7.0 in head. Eye 2.9—3.9.
Posrorsitat short, 1.8—2.1 in head. Maxitiary pointed, extending nearly to margin of
Fishes of the Western North Atlantic it
opercle, 1.1-1.3 in head. CuzExk short and broad, about as long as eye, its posterior
angle approximately 60°. GiLL RAKERS somewhat shorter than eye.
Dorsat fin rather low, with nearly straight margin, its last ray scarcely longer
than the one immediately before it, its longest rays failing to reach its tip if depressed,
the origin varying between a little nearer to caudal base than to upper anterior angle
of gill opening and equidistant between caudal base and posterior margin of eye. ANAL
usually originating somewhat posterior to dorsal origin, its base 3.3—4.0 in SL. Prtvic
fin very small, not quite reaching halfway to origin of anal, inserted nearer to anal
Ficure 32. Anchoa mitchilli diaphana, 72 mm TL, from Grand Isle, Louisiana, USNM 119790. Drawn by
Louella E. Cable.
origin than to pectoral base by diameter of pupil. Pecrorat variable, failing to reach
base of pelvic by a distance equal to, or even greater than, diameter of eye in northern
specimens, sometimes reaching pelvic base in southern fish, 1.4—1.85 in head. Axtt-
LARY SCALE of pectoral rather narrow, 2.3—3.0 in head.
Color. Preserved specimens, straw-colored above. Lower parts pale silvery. Sides of
head bright silvery. Middle of side with a silvery band, often nearly as wide as eye in
northern specimens, generally narrower in southern ones. Back with dusky punctula-
tions, arranged in two more or less definite rows posterior to dorsal fin; base of anal
with dark spots continued as a single unbroken median ventral line on caudal peduncle;
caudal with many dark points. Color in life, greenish with bluish reflections above;
lower parts pale; abdominal walls translucent.
Size. A total length of 100 mm (4 in.) is seldom exceeded, the usual length being
around 75 mm (3 in.). The largest examples measured were taken in New York, where
this species evidently grows larger than in the southern part of its range, for the largest
specimen among thousands from the Gulf of Mexico was only 75 mm long.
Development and Growth. The egg generally is slightly oblong, but occasionally
round; the major axis is about 0.75 mm and the minor one about 0.62 mm. It is
transparent and has no oil globule. The yolk consists of separate masses that appear
12
178 Memor Sears Foundation for Marine Research
as large cells under the microscope, a character that seems to be common to all species
of Engraulidae whose eggs have been studied. The egg floats at or near the surface
and hatches in about 24 hours at room temperature, at least at Beaufort, North Carolina,
during early summer.*
The newly hatched fish, 1.8-2.0mm long, is rather slender, perfectly trans-
parent, and has no pigment spots. The yolksac is absorbed within about two days, and
the large mouth, which is terminal at this stage, then seems to be functional. The pro-
jecting snout is not developed until the young fish reaches a length of about 20-25 mm.
But the fins are sufficiently developed at a length of about 15 mm to permit a fairly
accurate enumeration of the rays. The number of anal rays and the relative positions
of the dorsal and anal fins are very useful in identifying the postlarvae with the adults.
The early young of the season seem to become sexually mature during their first
summer, for specimens 45-60 mm long that remained quite transparent, taken late
in July and during the first half of August at Beaufort, N. C., contained well devel-
oped roe.
Spawning. In the vicinity of Beaufort, N. C., spawning takes place at least from
early spring to midsummer. The eggs were taken in tows there as early as April 21 (in
1930) and as late as July 15 (in 1929), while gravid fish were seen as late as August 12
(from my unpublished notes).
Food. The food apparently consists mostly of Mysis and copepods, the latter being
the sole food of the young. Other items taken are small fish, gastropods, and isopods
Grane):
Enemies. This fish, which is exceedingly numerous in a large part of its range, is
preyed upon extensively by predatory fish and water birds.
Variation. The variation within this species is so great that I recognized a
northern subspecies, 4. mitchilli mitchilli, and a southern one, A. mitchilli diaphana
(3: 87-94). The specimens from the southern part of the range have rather consistently
deeper and more strongly compressed bodies, fewer gill rakers, fewer vertebrae, a
longer pectoral fin, and a narrower silvery lateral band. These differences are shown by
comparing the characters of a small lot of specimens from Woods Hole, Massachusetts,
with another lot from Grand Isle, Louisiana, which show the widest divergence
(Table 1v). Complete intergradation of the two subspecies occurs in the midsection of
their range, as in North and South Carolina.
Commercial Importance. This Anchovy is used as bait and to a limited extent in
the preparation of anchovy paste. However, it is important chiefly as forage for many
important foodfishes. As this fish usually occurs in schools and is a rather sluggish
swimmer, it is an easy prey of many predators.
Range and Habitat. Cape Cod, Massachusetts (rarely northward into the Gulf of
Maine), and southward to Yucatan, México. Brazilian records probably are referable
either to 4. parva or A. januaria. While A. mitchilli occurs along the outer and exposed
4. The information as to the eggs and young fish is in part from Kuntz (9g: 13-19) and in part from my
field notes.
Fishes of the Western North Atlantic 179
Table IV. Comparison of Divergent Specimens of 4. mitchilli.
Woops Hott, MassacHusETTs Granp Isiz, Louisiana
Eight specimens, 62-85 mm TL Sixteen specimens, 48-76 mm TL.
Depth, 19-21 °/y of SL. Depth, 23-26 °/, of SL.
Greatest thickness of body, usually exceeding depth of Greatest thickness of body, less than depth of caudal
caudal peduncle. peduncle.
Pectoral fin, 13-13.5 °/) of SL. Pectoral fin, 16.5-17.5 °/) of SL.
Gill rakers, 24—28 on lower limb. Gill rakers, 21-23 on lower limb.
Vertebrae, 43 or 44 in four specimens. Vertebrae, 39-41 in nine specimens.
Silvery lateral band about as broad as eye. Silvery lateral band scarcely broader than pupil.
beaches at Beaufort, N.C., at least to a depth of 15—20 fms., it is found more often
in inside waters, especially in areas with muddy bottoms and brackish water. The dis-
tribution is so general that the fish may be expected also in “grassy” areas and along
sandy beaches.
Migrations. It was formerly thought that this species, as well as several other
anchovies, migrated northward and southward with the seasons. This theory was based
on the fact that in the northern part of the range, as at Woods Hole, Massachusetts,
the fish is present only during the summer. However, it has been shown (3: 87—94)
that virtually every section of the coast within the range of mirchilli has a distinctive
population, which suggests that whatever migration takes place is an inshore and
offshore movement.
Details of Occurrence. The most northern records are for Casco Bay, Maine, and
for Provincetown at the tip of Cape Cod, Massachusetts; these have been based on
stray specimens only. But it is described as abundant in the Woods Hole region and
in Rhode Island waters (both adults and larvae), common in New York waters, exceed-
ingly abundant (often in large schools) at many New Jersey localities, and second in
abundance (if not the most abundant) in Chesapeake Bay. It has been recorded for
North Carolina, South Carolina,® and Georgia, and it is at least tolerably plentiful in
the Indian River on the eastern coast of Florida. No certain evidence has been found of
its presence among the Florida Keys, but it has long been known to be plentiful in the
inlets and passes all along the western coast of Florida. It has been recorded for so many
localities in Mississippi, Louisiana (Cameron, Grand Isle, and Lake Ponchartrain),
Texas (Galveston, Corpus Christi, Dickinson Bayou), and México (Vera Cruz, Rio
Panuco, Tampico) as to prove it practically universal in suitable situations all around
the coastline of the Gulf of Mexico to Yucatan. But present indications are that Yucatan
marks the southern boundary of its range, there being no reliable record of its presence
anywhere to the southward around the Central or South American shores of the Carib-
bean.
Reports of it for Cuba (20: 57; I7: 421; 7: 405) on the strength of two
specimens from Poey in the Museum of Comparative Zoology seem to have been
based on a misinterpretation of what Poey wrote about this Anchovy. Actually the
5. The Museum of Comparative Zoology has a specimen from Charleston, collected by Louis Agassiz.
180 Memoir Sears Foundation for Marine Research
locality given in the Museum catalogue for these specimens is “United States,’’ to
which a label in the bottle in my handwriting adds “Sent to Poey by Gill for com-
parison.”
Synonyms and References:
Engraulis mitchilli Cuvier and Valenciennes, Hist. Nat. Poiss., 2Z, 1848: 50 (orig. descr.; type local. New
York; mentions Exgraulis louisiana®); Giinther, Cat. Fish. Brit. Mus., 7, 1868: 391 (synon., descr.,
range).
Stolephorus mitchilli Jordan and Gilbert, Proc. U.S. nat. Mus. (1882), 5, 1883: 248 (descr., synon., includ.
Engraulis duodecim Cope, a valid species; Galveston, Texas; Pensacola, Florida; Woods Hole, Massa-
chusetts); Swain, Bull. U.S. Fish Comm. (1882), 2, 1883: 57 (range; Princetown and Woods Hole,
Massachusetts; Pensacola, Florida; Lake Ponchartrain, Louisiana; Galveston, Texas; and Cuba, which
is probably 4. parva; includ. Engraulis duodecim Cope); Bean, Bull. U.S. Fish Comm. (1887), 7,
1888: 149 (Ocean City, Beesley’s Pt., and Longport, New Jersey; local abund.); Henshall, Bull. U. S.
Fish Comm. (1889), 9, 1891: 373 (very abund. in most inlets and passes of Florida west coast); Smith,
Bull. U.S. Fish Comm. (1892), 72, 1894: 361 (abund. in New Jersey; relation. to commerc. fishes);
Evermann and Kendall, Bull. U.S. Fish Comm. (1892), 12, 1894: 105 (Galveston, Corpus Christi,
Dickinson Bayou, Texas); Moore, Bull. U.S. Fish Comm. (1892), 12, 1894: 359 (near Sea Isle City,
New Jersey); Lonnberg, Oefvers. Svensk. Vet. Akad. Forh., 57, 1894: 115 (St. Petersburg, Florida); Jordan
and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 446 (descr., range, synon., includ. Exgraulis duodecim
Cope); Evermann and Bean, Rep. U.S. Comm. Fish. (1896), 1897: 241 (many at Cocoa and St. Lucia
on Indian R., Florida); Smith, Bull. U.S. Fish Comm. (1897), 77, 1898: 92 (Woods Hole, Massa-
chusetts); Smith and Bean, Bull. U.S. Fish Comm. (1898), 78, 1899: 184 (Potomac R. at Bryants
Pt., Maryland); Evermann, Rep. U.S. Comm. Fish. (1898), 1899: 309 (Baldwin Lodge, Mississippi;
Lake Lapourde, Louisiana); and (1899), 1900: 56 (Florida); Bean, Rep. Forest Comm. N. Y. (1901),
1902: 311 (refs., range, New York, econ. import.); Bean, Bull. N. Y. St. Mus., 60, Zool. 9, 1903:
218 (descr., range, a forage fish; New York); Latham, Copeia, 1917 (Orient, Long Island, New York);
Breder, Zoologica, N. Y., 2, 1922: 338 (Sandy Hook Bay, New Jersey).
Anchovia mitchilli Fowler, Rep. N. J. St. Mus. (1905), 1906: 110, fig. (descr., Delaware R. off Ft. Delaware,
Delaware; Beesley’s Pt. and Ocean City, New Jersey); and (1906), 1907: 267 (Sea Isle City, New Jersey;
taken for bait); Smith, N.C. geol. econ. Surv., 2, 1907: 134 (synon., descr., range, Morehead City,
North Carolina); Jordan and Dickerson, Proc. U.S. nat. Mus., 34, 1908: 12 (diagn., mouth of Rio
Pdnuco, Tampico, México); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7, 1908: 42 (Casco Bay, Maine;
Provincetown and Woods Hole, Massachusetts; Rhode Island); Weymouth, Proc. U.S. nat. Mus., 38,
1910: 136 (Cameron, Louisiana); Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 219 (Woods Hole,
Massachusetts; Seaside Park, Beesley’s Pt., Carson’s Inlet, New Jersey; Ft. Delaware, Delaware; Tol-
chester Beach, Maryland); Sumner, Osburn, and Cole, Bull. U.S. Bur. Fish. (1911), 32 (2), 1913: 743
(Woods Hole, Massachusetts); Kuntz, Bull. U.S. Bur. Fish. (1913), 33, 1914: 13-19, figs. 25-46
(spawn., Beaufort, North Carolina; develop. of eggs and larvae); Bigelow and Welsh, Bull. U.S. Bur. Fish.
(1924), 40 (1), 1925: 124 (descr., size, range, Gulf of Maine); Nichols and Breder, Zoologica, N. Y.,
9, 1927: 44 (diagn., range, Orient, New York; life hist. notes).
Anchoviella mitchilli Jordan and Seale (in part; not of Cuvier and Valenciennes), Bull. Mus. comp. Zool. Harv.,
67 (11), 1926: 405 (synon., includ. Exgraulis duodecim Cope; descr., range; Brazilian and West Indian
specimens not this species); Hildebrand and Schroeder, Bull. U.S. Bur. Fish. (1927), 43 (1), 1928: 109
(synon., descr., food, spawn., eggs descr., size, abund., habitat, Chesapeake Bay); Breder, Field Bk.
Mar. Fish. Atl. Cst., 1929: 71 (genl.); Jordan, Manual Vert. Anim. NE U.S., ed. 13, 1929: 43
(descr., range); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 48 (synon.,
includ. Exgraulis duodecim Cope; range); Hildebrand and Cable, Bull. U.S. Bur. Fish., 46, 1930: 388
(econ. import., spawn., Beaufort, North Carolina; develop. of eggs and young cf. 4. Aepsetus; distinguish.
characters this species and 4. hepsetus; growth rate); Perlmutter, 28th Rep. N.Y. Conserv. Dept.
6. Said to have been described and figured by LeSueur, but since no reference to £. /ouisiana LeSueur is given and
since none can now be found, the name presumably was taken from a manuscript, and Hildebrand, in his review
of the American Engraulidae regarded it as having no standing in modern nomenclature (3).
Fishes of the Western North Atlantic 181
(1938), Suppl., 15 (2), 1939: 18 (around Long Island, New York; spawn.); Greeley, 28th Rep.
N. Y. Conserv. Dept. (1938), Suppl., 15 (2), 1939: 83 (Long Island, New York; a forage fish).
Anchoa mitchilli mitchilli Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 87, fig. 37 (synon., descr.,
cf. 4. mitchilli diaphana, range).
Anchoa mitchilli diaphana Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 91, fig. 38 (orig. descr. of
subspecies 4. m. diaphana; relation., range); Gunter, Publ. Inst. mar. Sci. Texas, © (1), 1945: 33
(depth, temp., salinity, season, spawn., growth, Texas coast).
Negative References:
Stolephorus mitchilli Bean, Bull. U.S. Fish Comm. (1888), 12, 1890: 206 (many at Cozumel Island, Yucatan;
USNM 37053, 37105 are A. cayorum); Schreiner and Miranda-Ribeiro, Arch. Mus. nac., Rio de J.,
I2, 1903: 93 (Rio de Janeiro, where this species does not occur).
Anchoviella mitchilli Fowler, Arqu. Zool. estad. Sao Paulo, 3 (6), 1941: 135 (refs. Brazilian recs., where this
species does not occur).
Anchovia mitchilli von \hering, Rev. Industr. Anim., Anno 1 (3), 1930: 232 (Brazil, where the species does
not occur).
Anchoa parva (Meek and Hildebrand) 1923
Manjuia (Cuba)
Figure 33
Study Material. At least 34 specimens, 35-60 mm TL, and many more used for
some measurements and counts, from the following localities: many from Jamaica;
two from Cuba; many from Coldn and Porto Bello, Panama, and from Gatun Locks,
Canal Zone; one from Laguna de Tacarigua, Venezuela; and one from Trinidad; in
USNM, MCZ, UMMZ, BOC, Museo Poey collections.
Distinctive Characters. A. parva is rather close to both subspecies of mitchilli,
differing principally in having fewer anal rays and more numerous gill rakers, though
there is slight overlapping in each instance (see Tables 1, 1, and 11), in the smaller
anal fin (see Description), in the origin of the anal, which is somewhat farther back
under the base of the dorsal, and in the more strongly projecting snout (Figs. 31-33).
For the relationship of parva to januaria, see p. 183. Its closest relative is curta of
the Pacific coast of Central and South America, from which it is scarcely more than
subspecifically distinct (3: 84).
Description. Proportional dimensions in per cent of standard length, and counts,
based on at least 34 specimens in Study Material, 35-60 mm TL.
Body: depth 20-22. Pectoral fin: length 14.3-17.4.
Head: length 23.3-27.8.
Snout: length 4.0-5.5. Scales: ca. 38-42.
Eye: diameter 6.9-9.5. Gill rakers: 18-20+23-27.
Postorbital: distance 11.1—14.3. Fin rays: dorsal 13-16; anal 21-23
Maxillary: length 17.3-21.7. (rarely as many as 25); pectoral 12
Mandible: length 15-19. or 13.
Anal fin: length of base 21-25. Vertebrae: 38-40, rarely 41 (61 spec-
imens).
182 Memoir Sears Foundation for Marine Research
Bopy moderately deep, strongly compressed, its depth 4.5—5.0 in SL. Heap
3-6-4.3, its depth usually equal to its length without snout. SNout 4.5—6.0 in head,
extending about half of its length beyond mandible. Eyre 3.0-3.5. Posrorsirat 1.8—2.3
in head. Maxittary pointed, reaching to margin of opercle, 1.2-1.6 in head. Man-
DIBLE I.4—1.75. CHEEK scarcely longer than eye, its posterior angle about 60°.
Dorsat fin with the longest rays failing to reach tip of last ray if deflexed, its origin
generally equidistant between base of caudal and about middle of eye. Anat with origin
under, or more frequently slightly anterior to, middle of dorsal base, its base 4.0-4.8
in SL. Petvic reaching scarcely halfway to anal, inserted about equidistant between
origin of anal and base of pectoral. Pecrorat rather variable in length, usually failing
SS
Ficure 33. Azchoa parva, paratype, 59 mm TL, from Fox Bay, Colén, Panama, USNM 79554. Drawn by
Louella E. Cable.
to reach pelvic by diameter of pupil, 1.5-1.7 in head. Axitiary scaLe of pectoral
generally reaching to beginning of distal third of fin, 2.3-3.0 in head.
Color. In alcohol, very pale (presumably preserved originally in formalin). Silvery
lateral band scarcely as wide as pupil (indistinct or missing in some specimens). Back
with dusky punctulations, at least posterior to dorsal fin and extending onto base of upper
rays of caudal; larger dark dots usually present at base of anal and continued as a single
row onto median line of caudal peduncle.
Size. It is apparently small, as the largest specimen seen is only 60mm (2.4 in.) TL.
Range and Habitat. Known from the West Indies and the Atlantic coast of Panama
and Venezuela. Records of 4. mitchilli for the West Indies, in part at least, are refer-
able to this species. This fish inhabits salt and brackish water, and fresh water at least
to a limited extent. It was very numerous in all three flights of Gatun Locks, Canal
Zone, in 1935. The water in the lowermost flight was quite salty, that in the middle one
brackish, and that in the uppermost flight, fresh. It has been reported also as occur-
ring in fresh water in Venezuela.
Synonyms and References:
Anchovia parva Meek and Hildebrand, Field Mus. Publ., Zool., 75 (1), 1923: 202, pl. 10, fig. 2 (orig. descr.;
type local. Porto Bello, Panama; type USNM 81767; also recorded from Colén); Hildebrand, Zoologica,
Fishes of the Western North Atlantic 183
N. Y., 24 (1), 1939: 25, 35 (numerous in Gatun Locks, Canal Zone; erroneously reported from Mira-
flores Locks, Canal Zone, which is 4. curta).
Anchoviella mitchilli Jordan and Seale (in part; not of Cuvier and Valenciennes), Bull. Mus. comp. Zool. Harv.,
67, 1926: 405 (part of Cuban specimens, 4. parva).
Anchoviella parva Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 49 (ref. to orig.
descr.).
Anchoa parva Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 83, fig. 35 (refs., descr., relation., range);
Schultz, Proc. U.S. nat. Mus., 99, 1949: 43 (synon.; Venezuela; notes, counts).
Doubtful Reference:
Anchoviella parva Fowler, Proc. Acad. nat. Sci. Philad., 83, 1931: 392 (98-mm specimen from Trinidad much
larger than any seen by me; also anal rays more numerous).
MY
3
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flora hy Ogg
Ficure 34. Anchoa januaria, 75 mm TL, from Pernambuco, Brazil, MCZ 18012. Drawn by Alice C. Mullen.
Anchoa januaria (Steindachner) 1879
Figure 34
Study Material. A total of 28 specimens, 30-75 mm TL, from Amuay’ and
Salinas bays, Gulf of Venezuela; Pernambuco (Recife), Natal, and Rio de Janeiro,
Brazil; in USNM and MCZ collections.
Distinctive Characters. From both subspecies of mitchilli, januaria differs in a some-
what greater number of gill rakers, fewer anal rays, the more posterior position of the
anal fin, and the more strongly projecting snout. Compared to both mitchilli and parva,
it has a shorter and blunter maxillary; and from parva it differs further in having more
numerous gill rakers on the upper limb and in the slightly more posterior position of
the anal fin. It differs from pectoralis in having fewer pectoral and anal rays and in the
more numerous gill rakers.
7. Schultz (rg: 44) has claimed that the somewhat mutilated specimens from Amuay Bay, Gulf of Venezuela, identified
by me as this species, are 4. blackburni.
184 Memoir Sears Foundation for Marine Research
Description. Proportional dimensions in per cent of standard length, and counts,
based on 15 or more specimens from 33 mm SL upward.
Body: depth 18-20. Anal fin: \ength of base 22-26.
Head: length 22—26.2. Pectoral fin: length 15.6—-17.5.
Snout: length 4.15—5.7.
Eye: diameter 6.9—8.0. Scales: ca. 36-40.
Postorbital: distance 10.5-14.2. Gill rakers: 20-23 + 23-26.
Maxillary: length 16.4-19.3. Fin rays: dorsal 14 or 15; anal 21-24;
Mandible: length 15.3-17. pectoral 12 or 13.
Vertebrae: 41 or 42 (3 specimens).
Bopy strongly compressed, its depth 4.3—5.0 in SL. Heap short, 3.8-4.5 in SL,
its depth equal to its length without snout. SNnour 5.0—6.0 in head, projecting fully
half of its length beyond mandible. Eve 2.9—-3.6 in head. PostorsiTaL 2.0—2.3 in
head. Maxiiiary rather bluntly pointed, its upper margin somewhat rounded, reaching
to, or slightly beyond, joint of mandible, 1.3-1.5 in head. ManprBLe 1.4—1.55. CHEEK
about as long as eye, its posterior angle about 50°.
Dorsat fin with its longest rays failing to reach tip of last ray if deflexed, its origin
equidistant between base of caudal and about middle of eye. Anat low, its origin
under or slightly posterior to middle of dorsal base, its base scarcely shorter than head,
3.8—-4.5 in SL. Prxvic very small, reaching much less than halfway to anal, inserted
about equidistant between origin of anal and base of pectoral. Pecrorat small, failing
to reach ventral by more than half of diameter of eye, 1.4—2.0 in head. AxiLLary
scALE of pectoral about 75°/, of length of fin, 2.4 in head.
Color. In alcohol, pale silvery. Silvery lateral band distinct, about 0.5—0.66 of width
of eye. Back with dark punctulations, arranged in two rows in some specimens, probably
faded in others.
Size. So far as known, this Anchovy reaches a length of only about 75 mm
(3 in.).
Range. It apparently ranges from the Gulf of Venezuela to Rio de Janeiro, Brazil.
Synonyms and References:
Engraulis januarius Steindachner, Ichthyol. Beitr., 8, 1879: 58 (orig. descr.; type local. Rio de Janeiro, Brazil;
cf. E. mitchilli).
Anchovia januaria Starks, Stanf. Univ. Publ., Univ. Ser., 1913: 9 (Natal, Brazil).
Anchoviella januaria Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 206 (descr., Rio de Janeiro,
Brazil, and “Brazil”); Fowler, Arqu. Zool. estad. Sao Paulo, 3 (6), 1941: 134 (refs. to Brazilian
recs.).
Anchoviella mitchilli Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 405 (Pernambuco, Brazil;
MCZ 18012 are 4. januaria).
Anchoa januaria Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 81, fig. 34 (synon., descr., range;
cf. mitchilli, parva, and pectoralis).
Fishes of the Western North Atlantic 185
Anchoa choerostoma (Goode) 1874
Figure 35
Study Material. Many specimens in nine collections from Bermuda.
Description. Proportional dimensions in per cent of standard length, and counts,
based on at least 22 specimens, 55-75 mm TL.
Body: depth 18.2—20.8. Anal fin: \ength of base 20—22.6.
Head: length 27.8—30.2. Pectoral fin: length 15.6—-18.5.
Snout: length 5.1—-6.4.
Eye: diameter 6.58.4. Scales: ca. 38-40.
Postorbital: distance 13.3-15.9. Gill rakers: 17-20+ 23-26.
Maxillary: length 21.2-24.3. Fin rays: dorsal 13-15; anal 22-24
Mandible: length 19.2-22. (rarely 21 or 25); pectoral 12-14.
Vertebrae: 41 or 42 (2 specimens).
Ficure 35. Anchoa choerostoma, 68mm TL, from Bermuda, USNM 21976. Drawn by Louella E. Cable.
Bopy strongly compressed, moderately deep, its depth 4.8—5.8 in SL. Heap
3-3-3-6 in SL, its depth equal to postorbital and about half of eye. Snour short,
projecting rather more than half of its length beyond mandible, 4.7—5.8 in head. Eve
3.34.2. PosrorpitaL 1.85—2.1 in head. Maxitiary reaching nearly to gill opening,
not sharply pointed, its upper margin rounded, 1.2-1.3 in head. ManpiBLe 1.3-1.5
in head. Cueex length equal to eye and fully 75°/, of snout, its posterior angle
about 30°.
Dorsat fin rather high anteriorly, its longest rays reaching to or beyond tip of
last ray if deflexed, its origin about equidistant between base of caudal and anterior
margin of eye. ANAL with origin somewhere under posterior half of dorsal, but well in
advance of the base of last ray, its base 4.4—5.0 in SL. Pexvic failing to reach anal by
nearly an eye’s diameter, inserted a little nearer to origin of anal than to pectoral base.
PecroraL moderately long, reaching nearly or quite to pelvic, 1.5-1.9 in head.
186 Memotr Sears Foundation for Marine Research
AXILLARY SCALE of pectoral reaching somewhat beyond midlength of fin, 3.0-4.0
in head.
Color. In alcohol, generally pale. Side with a silvery band (often missing in speci-
mens preserved a long time in formalin), about 75 °/, of width of eye. Upper surface
of head and back with dusky dots, usually arranged in two longitudinal series
posterior to dorsal fin.
Size. The largest seen is 75 mm (3 in.) TL, which may be near the maximum
size attained.
Reproduction. Specimens taken on June 24, 1927 at Nonsuch, Bermuda contain
well developed eggs, indicating that at least some reproduction takes place there during
the early part of summer.
Range and Habitat. The range seems to be limited to Bermuda, where no other
species of anchovy has been found. West Indian and Panamanian (Atlantic) records
are mostly referable to 4. /yo/epis. The collections studied, according to the information
furnished, were either seined near shore or taken at night under an electric light. This
species has been reported as fairly common about Bermuda (numerous in Hamilton
Harbor). Although it has been reported as not at all common during July, it was abun-
dant in August when seined for bait in Bailey’s Bay and Flatts Inlet.
Synonyms and References:
Engraulis choerostoma Goode, Amer. J. Sci., 8, 1874: 125 (orig. descr.; type local. Bermuda; types lost); Ann.
Mag. nat. Hist., (4) 14, 1874: 380 (as in Goode, 1874).
Stolephorus choerostoma Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 444 (descr. after Goode;
ref.); Barbour, Bull. Mus. comp. Zool. Harv., 46, 1905: 113 (abund. in Bermuda; distrib., includ. Puerto
Rico, where it apparently does not occur; refs.); Bean, T. H., Field Mus. Publ., Zool., 7, 1906: 34
(several locals in Bermuda).
Anchoviella choerostoma Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 48 (ref.,
Hamilton Harbor, Bermuda); Fowler, Proc. biol. Soc. Wash., 43, 1930: 146 (72 from Bermuda).
Anchoa choerostoma Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 71, fig. 28 (refs., descr., distrib.).
Negative References:
Anchoviella choerostoma Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 404 (not 4. choerostoma
[Goode]; descr. evidently based on 4. /yo/epis [Evermann and Marsh]; those from Puerto Rico and e.
coast of Panama are /yo/epis); Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda, 1933: 38, fig. (not
A. choerostoma [Goode]; fig. and descr. at least in part from Beebe and Tee-Van [1928: 47]; distrib.
includ. West Indies, where this species apparently does not occur); Beebe and Tee-Van, Zoologica,
N. Y., 23 (7), 1933: 136 (4. choerostoma var. atlantica Borodin [1928: 7] from e. coast of Panama put
in synon., but 4. choerostoma [Goode] apparently does not occur there. Borodin’s fish not ident. from
his descr.).
Anchoa tricolor (Agassiz) 1829
Piquitinga
Figure 36
Study Material. Many specimens, from: Gulf of Venezuela (1); Natal (8), Rio de
Janeiro (52), Sao Francisco (5), and Sambara River (20), Brazil; Montevideo, Uruguay
Fishes of the Western North Atlantic 187
(1); and Puerto de (or Rio) Quequen, Mar del Plata, Argentina (2), in USNM, MCZ,
CNHM, MHNBA, UMMZ, SU collections.
Distinctive Characters. This species generally has been identified in museum col-
lections with drownii = hepsetus, which it resembles in size and general appearance, but
it differs sharply from it in the shorter and differently shaped maxillary and in the more
numerous gill rakers (Tables 1 and 11).
Ficure 36. Azchoa tricolor, 110 mm TL,'g91 mm SL, from Rio de Janeiro, Brazil, USNM 87721, with pectoral
fin enlarged. Drawn by Louella E. Cable.
Description. Proportional dimensions in per cent of standard length, and counts,
based on at least 41 specimens, 63-118 mm TL.
Body: depth 18.2-21. Anal fin: length of base 17.2—20.
Head: length 23.8—26.7. Pectoral fin: length 14-14.8.
Snout: length 4.15—5.9.
Eye: diameter 6.7—-8.1. Scales: ca. 42-45.
Postorbital: distance 12.5—13.5. Gill rakers: 18-22+24—28.
Maxillary: length 18.2-21.5. Fin rays: dorsal 14-16; anal 18-22;
Mandible: length 15.6—-18.3. pectoral 13-15.
Vertebrae: 40-42 (§ specimens).
Bopy quite slender, moderately compressed, its depth 4.75-5.5 in SL. Heap
3-75—-4.2 in SL, its depth equal to its postorbital length and fully half of eye. Snour
projecting about half of its length beyond mandible, 4.75—5.8 in head. Eve 3.25—3.75.
PostorsiTaL 1.85—2.15 in head. Maxitiary not sharply pointed, its upper margin
rounded, reaching to, or a little beyond, joint of mandible, 1.3-1.4 in head. MANDIBLE
1.31.5. CHEEK rather broad, scarcely longer than eye, its posterior angle about 55°.
188 Memoir Sears Foundation for Marine Research
Dorsat fin with longest rays failing to reach tip of last ray if deflexed, its origin
about equidistant between base of caudal and middle of eye. Anat with origin some-
where under posterior fourth of dorsal, its base 5.0—-5.8 in SL. Petvic reaching scarcely
halfway to anal, inserted about equidistant between origin of anal and pectoral base.
Prcrorat scarcely falcate, failing to reach pelvic by diameter of pupil to 75°/, of dia-
meter of eye, 1.75—2.0 in head. AxrLLary scaLe of pectoral often shorter than fin by
diameter of pupil, 1.9-2.75 in head.
Color. In alcohol, generally rather pale. Side of head silvery. Lateral band bright
silvery, about 75°/, of width of eye.
Size. The largest measured is 118 mm (4.7 in.) TL, which may be near the
maximum length attained.
Commercial Importance. Most of the specimens examined were obtained in markets,
indicating that this Anchovy has at least some direct economic value to man,
Range. The range, so far as known, extends from the Gulf of Venezuela to Mar
del Plata, Argentina, but it apparently does not include streams. This Anchovy’s
frequent appearance in the Rio de Janeiro market (Brazil) seems to indicate that
it is common locally.
Synonyms and References:
Engraulis tricolor Agassiz, in Spix and Agassiz, Pisc. Brasil., 1829: 51, tab. 23, fig. 1 (orig. descr.; type local.
Bahia, Brazil; plate labeled Engrau/is piguitinga); Goeldi, Boll. Museu Paraense, 2 (3), 1898: 466
(names; from type local.).
Stolephorus brownii Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 443 (descr., range, synon.,
includ. £. ¢ricolor Agassiz).
Anchovia brownii Starks, Stanf. Univ. Publ., Univ. Ser., 1913: ro (Natal, Brazil; local abund.; descr. ¢rico/or
by my exam.).
Anchoviella salvatoris Fowler and Bean, Proc. U.S. nat. Mus., 63 (19), 1923: 6 (orig. descr.; type local. Rio
de Janeiro, Brazil; type USNM 83165, in bad condition, apparently ¢ricolor); Fowler, Arqu. Zool.
estad. Sao Paulo, 3 (6), 1941: 134 (refs.).
Anchoviella epsetus Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 396 (Rio de J. and “‘Sambaia”
[probably Sambara R.], Brazil; Montevideo, Uruguay; all exam. and are ¢rico/or); Jordan, Evermann,
and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 48 (includ. ¢rico/or); Fowler, Arqu. Zool. estad.
Sao Paulo, 3 (6), 1941: 134 (refs. to Brazilian recs., in part ¢ricodor).
Anchoviella bonaiensis Marini, Physis, Rev. Soc. Argent. Cienc. nat., II, 1935: 446 (NOMEN NuDUM; type local.
Mar del Plata, Argentina; type in MHNBA; exam. by me and found to be “rico/or).
Anchoa tricolor Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 74, fig. 30 (synon., descr., relation.,
distrib.); Schultz, Proc. U.S. nat. Mus., 99, 1949: 43 (synon.).
Anchoa cuvbana (Poey) 1868
Cuban Anchovy, Bocén, Manjtta (Cuba)
Figure 37
Study Material. A total of 39 specimens, 45-70 mm TL, including two from
Poey’s type material, MCZ 17958, and two paratypes of Stolephorus astilbe Jordan
and Rutter, MCZ 31392; many more for the gill raker counts; from Melbourne Beach
Fishes of the Western North Atlantic 189
and Marco, Florida; Mobile Bay, Alabama; off Grand Isle, Louisiana; Progreso,
Yucatan, México; Puerto Barrios, Guatemala; Cuba; Jamaica; and St. Thomas.
Distinctive Characters. This species differs from 4. hepsetus and from nearly all
other local forms in a very slender body and more numerous gill rakers.
Description. Proportional dimensions in per cent of standard length, and counts,
based on 39 specimens, 45-70 mm TL.
Ficure 37. Anchoa cubana, 63mm TL, 53 mm SL, from St. Thomas, West Indies, USNM 83802. Drawn
by Louella E. Cable.
Body: depth 15-18. Pectoral fin: length 12.5—13.7.
Head: length 22.7—27.8.
Snout: length 3.8 5—4.7. Scales: ca. 40-43.
Eye: diameter 6.1-7.75. Gill rakers: 17-23 + 23-33, usually 19—
Postorbital: distance 10.5-12.5. 22+25-28 (Tables um, 111).
Maxillary: length 18.7-22.2. Fin rays: dorsal 14-16; anal 20-24;
Mandible: length 15.6—18.8. pectoral 13 or 14.
Anal fin: length of base 18.5—22.2. Vertebrae: 42 or 43 (14 specimens).
Bopy slender, rather strongly compressed, with a rather sharp edge on chest and
abdomen, its depth 5.5—6.7 in SL. Heap small, 3.6—4.3 in SL, its depth equal to its
postorbital length and about a third of eye. Snour short, extending about 66°/, of its
length beyond tip of mandible, 4.8-6.5 in head. Eye small, 3.5-4.0. Posrorsirat
short, 1.9—2.3 in head. Maxitrary bluntly pointed, its upper margin nearly straight,
usually extending nearly to margin of opercle, 1.1-1.25 in head. ManpiBte 1.4—1.75.
CurEk somewhat longer than eye, its posterior angle rather broad, about 45°. Git
rakeRS slender, close-set, often difficult to count (probably accounting for wide
range given).
Dorsat fin with the longest rays failing to reach tip of last ray if deflexed, its origin
about equidistant between base of caudal and some point over posterior half of eye.
Anat with origin under or slightly in advance of middle of dorsal base, its base 4.5—
5.4 in SL. Petvic reaching somewhat less than halfway to origin of anal, inserted a
190 Memoir Sears Foundation for Marine Research
little nearer to origin of anal than to pectoral base. Pecrorat failing to reach base
of pelvic by nearly an eye’s diameter, 1.75-1.9 in head. AxrLiary scaLe of pectoral
about 75°/, of length of fin, 2.4—3.0 in head.
Color. In alcohol, rather pale. Side of head silvery. Lateral band bright silvery,
about as wide as pupil (often missing in specimens preserved in formalin). Back usually
with dusky punctulations not arranged in definite longitudinal series.
Size. The largest fish seen was only 70 mm (2.8 in.) long, and most of the numer-
ous specimens examined were under 65 mm TL.
Range and Habitat. A. cubana ranges from both coasts of Florida, along Alabama
and Louisiana, and through the West Indies, to Yucatan, México, and Puerto Barrios,
Guatemala. It is also recorded for Puerto Rico (18: 344) and Grenada (r: 269).
It seems to be common around Jamaica, probably also around Cuba, but apparently
it is rather rare on our southern coast. The specimens from the West Indies, in
part at least, were taken in shallow water, whereas those from the United States were
taken with trawls hauled at depths as great as 30 fms.
Synonyms and References:
Engraulis cubanus Poey, Repert. Fisico-Nat. Cuba, 2, 1868: 420 (orig. descr.; type local. Cuba; MCZ 17958);
Poey in Gundlach, An. Soc. esp. Hist. Nat., ro (4), 1881: 344 (Puerto Rico).
Stolephorus cubanus Jordan and Evermann, Bull. U. S. nat. Mus., 47 (1), 1896: 442 (descr. from orig., range);
Evermann and Marsh, Bull. U.S. Fish Comm. (1900), 20 (1), 1902: 88 (after Jordan and Evermann,
1896).
Stolephorus astilbe Jordan and Rutter, Proc. Acad. nat. Sci. Philad., 1897: 95 (orig. descr.; type local. Kingston,
Jamaica; type SU 4854; paratypes MCZ 31392); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (3);
1898: 2815 (descr. from orig., synon.).
Anchoviella cubana Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 399 (synon., descr., Cuba,
includ. type material, MCZ 17958); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928),
2, 1930: 48 (synon., range); Fowler, Proc. Acad. nat. Sci. Philad., 82, 1930: 269 (Grenada, B.W.1.);
Howell-Rivero, Bull. Mus. comp. Zool. Harv., 82, 1938: 172 (4 specimens in type material, the largest
said to be type, MCZ 17958).
Anchoviella astilbe Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 402 (synon., descr., paratypes
MCZ 31392); Jordan, Evermann, and Clark, Rept. U.S. Comm. Fish. (1928), 2, 1930: 49 (“West
Indies”); Fowler, Proc. Acad. nat. Sci. Philad., 82, 1930: 269 (Grenada, B.W.1.).
Anchovia cubana Nichols, N. Y. Acad. Sci., 10 (2), 1929: 204 (synon., diagn., Cuba and Puerto Rico).
Anchoa cubana Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 76, fig. 31 (synon., descr., extension
of range; cf. paratypes of cubana and astilbe).
Anchoa ginsburgi Hildebrand 1943
Figure 38
Study Material. Type material, 16 specimens, from juveniles 28 mm long to adults
or near adults 63 mm TL, from Gulf of Venezuela.
Distinctive Characters. This species differs from hepsetus chiefly in the shorter and
blunter maxillary and in the more numerous vertebrae when compared with hepsetus
from southern localities, especially from Yucatan and Panama (3: 56, tab. 7). Among
specimens of equal size, this species seems to be more slender.
Fishes of the Western North Atlantic IgI
Description. Proportional dimensions in per cent of standard length, and counts,
based on 13 specimens, 43-63 mm TL.
Body: depth 17.4-18.8. Anal fin: length of base 19.2—22.
Head: length 25-28.5. Pectoral fin: length 13.7-15.3.
Snout: length 4.9-5.9.
Eye: diameter 6.6—9.1. Scales: ca. 42-45.
Postorbital: distance 14.5-15.8. Gill rakers: 16-18 + 20-22.
Maxillary: length 17.2—-18.2. Fin rays: dorsal 14 or 15; anal 18-20;
Mandible: length 15.5—17.8. pectoral 13-15.
Vertebrae: 44 or 45 (2 specimens).
Saat Sere a
SR SSS
Ws
SS
ae
WY aS
SS
Ficure 38. Axchoa ginsburgi, paratype, 60mm TL, 52 mm SL, from Gulf of Venezuela, USNM 119789.
Drawn by Louella E. Cable.
Bopy slender, moderately compressed, its depth 5.3-5.75 in SL. Heap 3.5—4.0
in SL, its depth equal to its postorbital length and about half of eye. Snour short,
projecting a little more than half of its length beyond mandible, 4.5—5.25 in head.
Eye 3.25—4.0. Posrorpitat moderately long, 1.3-1.9 in head. Maxrtary short, its
upper margin rounded, reaching joint of mandible, 1.3-1.5 in head. ManprBLe 1.5—
1.65. CuEEk broad, a little longer than eye, its posterior angle about 45°.
Dorsat fin low, the longest rays failing to reach tip of last ray if deflexed, its origin
usually somewhat nearer to base of caudal than to anterior margin of eye. Anat short,
with origin commonly slightly posterior to middle of dorsal, its base 4.5—5.2 in SL.
Pexvic small, inserted equidistant between origin of anal and base of pectoral, or a little
nearer to the latter. Pecrorat scarcely falcate, failing to reach pelvic by about half of
diameter of eye, 1.7—2.0 in head. AxrLiary scaLe of pectoral moderately pointed, failing
to reach tip of fin by about 75°/, of diameter of eye, 2.8—3.1 in head.
Color. In alcohol pale (originally preserved in formalin). Side of head silvery.
Lateral band bright silvery, about 75 °/, of width of eye.
Range. Known only from the holotype and 15 paratypes, all from the Gulf of
Venezuela, partly from Estauques Bay and partly from Salinas Bay.
192 Memoir Sears Foundation for Marine Research
References:
Anchoa ginsburgi Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 55, fig. 20 (orig. descr.; type local.
Estauques Bay, Gulf of Venezuela; type USNM 119788); Schultz, Proc. U.S. nat. Mus., 99, 1949:
43 (ref., types, locals.).
Anchoa lamprotaenia Hildebrand 1943
Longnose Anchovy, Manjtia (Cuba)
Figure 39
Study Material. A total of 80 specimens, 46-92 mm TL, collected at Palm Beach,
Miami, Key West, and Tortugas, Florida; Progreso, Yucatan, México; Cuba; Puerto
Rico; British Honduras; and the Atlantic coast of Panama.
Distinctive Characters. This species until recently has been consistently identified
with 4. hepsetus in museum collections. Since these species are not easily separated by
any one character, a combination of characters is sometimes necessary. Generally
A. lamprotaenia is distinguished by its longer and more falcate pectoral fin, higher
dorsal fin, shorter and broader axillary scale of the pectoral, and by the shorter and more
bluntly pointed maxillary with its upper rounded margin. The average number of anal
rays is greater and the average number of pectoral rays, gill rakers, and vertebrae is
fewer (see Description and Tables 1-111).
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens, 46-92 mm TL.
Body: depth 19-23. Pectoral fin: length 14.3-18.
Head: length 25-30.
Snout: length 8.1-9.5. Scales: ca. 38-41.
Eye: diameter 7.5-8.3. Gill rakers: 13-18 + 16-20 (usually 14
Postorbital: distance 14.3-16. or 15+18 or 19).
Maxillary: length 21.5-24.5. Fin rays: dorsal 13-15; anal 19-26
Mandible: length 18.5—22. (usually 21-24); pectoral 11-15
Anal fin: length of base 20.5-23.5. (most frequently 12 or 13).
Vertebrae: 39-42 (98 specimens).
Bopy strongly compressed, its greatest thickness about equal to depth of
caudal peduncle, its depth 4.3-5.25 in SL. Heap 3.3-4.0, its depth about equal to
its postorbital length and half of eye. Snour pointed, projecting about 66°/, of its
length beyond tip of mandible, 4.5-5.75 in head. Eye 3.5—4.0 in head. PostorsrTaL
1.75—2.0 in head. Maxittary not sharply pointed, usually distinctly rounded above,
reaching to, or more usually somewhat beyond, joint of mandible, 1.2-1.4 in head.
ManDIBLE 1.35-1.6 in head. Cueek as long as eye and fully half of snout, its
posterior angle about 35°. Gitt RakERs about 75°/, of length of eye.
Dorsat fin rather high anteriorly, its longest rays usually reaching beyond tip of
last ray if deflexed, its origin equidistant between some point over snout or anterior
Fishes of the Western North Atlantic 193
half of eye and base of caudal. Anat fin with origin under or near beginning of posterior
third of dorsal base, its base 4.25—4.9 in SL. Pexvic usually inserted slightly nearer to
origin of anal than to base of pectoral. Pecrorat long, falcate, usually failing to reach
pelvic by only diameter of pupil, its longest ray in a few specimens with a slight filament,
the rays decreasing rapidly in length after the third one, the shortest ray scarcely reach-
ing past midlength of first (uppermost) one, 1.5—1.9 in head. AxILLary scALeE of pec-
toral broad at base, scarcely reaching beyond midlength of fin, 3.0-3.6 in head.
Ficure 39. Axchoa lamprotaenia, paratype, 60mm TL, 48 mm SL, from Key West, Florida, USNM 35000,
with pectoral fin enlarged. Drawn by Louella E. Cable.
Color. Preserved specimens pale. Side of head bright silvery. Silvery lateral band
prominent, not quite as wide as eye. Punctulations on back not prominent; a few dark
dots at base of anal; dorsal and caudal with a few dusky points; other fins unmarked.
Size. A. lamprotaenia apparently attains a smaller size than 4. hepsetus, the largest
seen being only 92 mm (3.7 in.) TL.
Reproduction. Specimens taken at Key West, Florida, June 11, 1936, and at Bush
Key (Tortugas), Florida, July 23, 1937, contained well-developed roe, showing that
at least some spawning takes place in southern Florida during midsummer.
Range and Habitat. From southern Florida through the West Indies to Panama,
and probably to northern Brazil as shown by an imperfect specimen. The collections
studied were apparently taken in shallow water near shore. The species is common in
the Florida Keys, where 4. hepsetus seems to be very rare.
Synonyms and References:
Stolephorus brownii Jordan, Proc. U.S. nat. Mus., 7, 1885: 106 (abund. at Key West, Florida; specimens
USNM 35000 are 4. /amprotaenia); Evermann and Marsh, Bull. U.S. Fish Comm. (1900), 20 (1),
13
194 Memoir Sears Foundation for Marine Research
1902: 88 (descr., range, distr. in Puerto Rico; synon.; specimens from Fajardo [USNM 73755], at
least, are 4. /amprotaenia).
Anchovia brownii Meek and Hildebrand, Field Mus. Publ., Zool., 75 (1), 1923: 204, pl. 14, fig. 1 (synon.,
descr., range, specimens from Colén [CNHM 8217] and Porto Bello [CNHM 8216], Panama, in part
are A. /amprotaenia).
Anchoviella epsetus Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 396 (synon., descr., range;
specimens from Tortugas [MCZ 17944] and Key West [MCZ 18035], Florida, are 4. /amprotaenia);
Pierce, Copeia, 1936: 123 (sample of specimens used in feed. experiments are 4. /amprotaenia); Long-
ley and Hildebrand, Publ. Carneg. Instn. Wash., 535, 1941: 12 (notes on structure; distr. at Tortugas,
Florida; specimens preserved are 4. /amprotaenia).
Anchoa lamprotaenia Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 62, fig. 23 (orig. descr.; type local.
Key West, Florida; type USNM 117661; range cf. 4. hepsetus).
Doubtful References:
Anchovia brownii Jordan and Thompson, Bull. U.S. Bur. Fish. (1904), 24, 1905: 233 (Garden Key [Tortugas],
Florida; probably part, if not all, are /emprotaenia); Fowler, Proc. Acad. nat. Sci. Philad., 58, 1906:
84; and 63, 1911: 216 (from several Florida Keys; probably part, if not all, are /amprotaenia).
Anchoa hepsetus (Linnaeus) 1758
Striped Anchovy (U.S.), Manjua (Cuba)
Figures 40, 41
Study Material. At least 83 specimens, 55-140 mm TL, and many more, taken
at various places from New Jersey southward to Uruguay.
Distinctive Characters. A. hepsetus is related closely enough to 4. /amprotaenia so
that they are not always separable by any one character, a combination of characters
sometimes being necessary. In general, Aepsetus has a shorter and less falcate pectoral
fin, a lower dorsal fin, a longer and narrower axillary scale on the pectoral, and a rather
longer and more sharply pointed maxillary. Also, its average number of anal rays 1s
lower, and its average number of pectoral rays, gill rakers, and vertebrae is higher.
It is related also to 4. ginsburgi, from which it differs especially in the longer and more
sharply pointed maxillary, and generally in having fewer vertebrae.
Description. Proportional dimensions in per cent of standard length, and counts,
based on at least 83 specimens, 55-140 mm TL.
Body: depth 18.5—22. Scales: ca. 37-43.
Head: length 25-30. Gill rakers: 15-20+ 18-24 (usually 16-
Snout: length 4.4—6.3. 19+ 19-22).
Eye: diameter 7.1 5-8.7. Fin rays: dorsal 13-16; anal 18-23;
Postorbital: distance 13.5—15.5. pectoral 13-15 (most frequently
Maxillary: length 19-24.5. Ld) Of 1S):
Mandible: length 16.5-20. Vertebrae: 40-44 (399 specimens), de-
Anal fin: length of base 19-22.5. creasing somewhat in number
Pectoral fin: length 14.5-17.5. southward at least to Panama.
Fishes of the Western North Atlantic 195
Bopy slender, moderately compressed, its greatest thickness generally exceeding
the depth of caudal peduncle, its depth 4.5—5.4 in SL. Heap 3.3—4.0 in SL, its depth
equal to postorbital and about a fourth of eye. SNour projecting about 66°/, of its
length beyond tip of mandible, 4.5—6.0 in head. Eye 3.25—4.0 in head. PosrorsiTaL
1.8—2.1 in head. Maxiitary quite pointed, its upper free margin generally straight,
reaching nearly to margin of opercle, 1.2—1.3 in head. Manprpie 1.4—1.6 in head.
Cueex a little longer than eye, its posterior angle rather broad, about 45°. GILL RAKERS
at angle of first arch about 75°/, of length of eye.
Dorsat fin low, its margin somewhat concave, its longest rays failing to reach tip
of last ray if deflexed, its origin usually about equidistant between base of caudal and
middle of eye. Anat fin with origin under middle to posterior fourth of dorsal base, its
base 4.4—5.2 in SL. Petvic reaching about halfway to origin of anal, inserted nearly
equidistant between origin of anal and pectoral base. Pecrorat not strongly falcate, its
four uppermost rays usually of about equal length, its first or uppermost one some-
times a little shorter than the second one, its last or shortest ray usually reaching well
past midlength of the first one, the fin often failing to reach base of pelvic by diameter
of eye, though sometimes nearly reaching it, its length 1.7-2.0 in head. AxILLary
scaLE of pectoral slender, usually failing to reach tip of fin by less than diameter of
pupil, 2.0-2.5 in head.
Color. Preserved specimens, pale above. Lower parts pale silvery. Side of head
silvery. Middle of side with a bright silvery band, variable in width, often about 75°/,
of width of eye above base of anal, sometimes scarcely wider than pupil.
Size. The largest specimen seen, from Beaufort, North Carolina, is 153 mm
(6.1 in.) TL, which probably is near the maximum size attained. A specimen 150 mm
long, weighing one ounce, has been reported for Chesapeake Bay. The usual size
ranges between about 100-125 mm TL.
Development and Growth. The egg is elliptical, with a major axis of about 1.4—
1.6 mm and a minor one of about 0.7—0.85 mm. It is transparent and has no oil
globule. The yolk, as in other anchovies studied, is separated into masses having
the appearance of large cells under magnification. The egg floats at or near the surface
and hatches within about 48 hours at a water temperature of about 66°—70° F.
The newly hatched fish, very slender and thread-like, is about 3.6—4.0 mm long.
It is transparent, has a slight greenish shade on the head, and has no definite chromato-
phores; the head is slightly concave. The myomeres are very distinct. It is quite
active. The yolksac generally is absorbed within 24 hours after hatching at 65°—70° F.
The fish is then about 4.0-4.5 mm long, at which stage the mouth is open and ap-
parently functional. The fish is still transparent.
The first chromatophores, situated in the midventral line of the chest, appear
when the fish is about 5.0 mm long, and at this stage the mouth is large, terminal,
and oblique. The dorsal and anal fins become more or less definitely differentiated
at a length of 5.0-6.0 mm, but the rays cannot be counted accurately until the fish
is about 10mm, at which length it has become considerably more robust, the
at
13
196 Memor Sears Foundation for Marine Research
notochord is definitely bent upward in the base of the caudal fin, and the caudal has
indefinite rays.
At a length of 15 mm the caudal fin is well developed and forked, the notochord
is no longer visible, and the mouth remains terminal and oblique. Some additional
chromatophores have appeared along the ventral surface, and dark pigment has ap-
peared along the upper margin of the eye.
Ficure 40. Anchoa hepsetus hepsetus, 114mm TL, from Atlantic City, New Jersey, USNM 127525, with
pectoral fin enlarged. Drawn by Louella E. Cable.
Little external development takes place while the fish grows from a length of
15 to 25 mm. However, when the fish reaches 35 mm, the mouth has become almost
horizontal, as in adults, and the conical snout projects prominently. Pigmentation has
increased and the silvery lateral band of the adult, though narrow and indistinct at this
point, has appeared. Although the body has now become fairly robust, it remains more
slender than in adults, its depth being contained in the length about 6.0 times.
When it reaches a length of about 40-45 mm it may be definitely considered
a young adult. Although it remains somewhat more slender than larger fish, it is fully
pigmented, has the characteristic silvery lateral band well developed, is fully scaled,
and is easily identified with the adult (4: 389-393; also unpublished field notes of
the writer).
Spawning. In the vicinity of Beaufort, North Carolina, spawning begins early in
the spring. April 16 is the earliest date on which eggs have been taken. Toward the
middle of June, eggs diminished in number in the tows, though a few were taken
throughout July. The young, 22 mm and less, were collected in Beaufort Inlet as early
as April 11. (4: 388; also unpublished data of the writer).
Food. When the fish is about 20 mm the food consists largely of copepods. As
Fishes of the Western North Atlantic 197
the fish grows, this diet is supplemented by gastropods, foraminifera, an occasional
ostracod, and annelid worms. The diet of the adult is essentially the same (Linton,
TE NAAOST2: 25942015):
Parasites. The principal parasites are nematodes, cestodes (Scolex polymorphus and
Rhynchobothrium sp.), and trematodes (Distomum appendiculatum and Distomum sp.)
(II: 440; 12: 353).
Ficure 41. Anchoa hepsetus colonensis, 70mm TL, 56 mm SL, from Colén, Panama, USNM 117664, with
pectoral fin enlarged. Drawn by Louella E. Cable.
Variation. So great is the variation in this species that certain specimens from
Colén, Panama, and northward to the Gulf of Mexico were recognized (3: 60, fig. 22)
as a subspecies, 4. hepsetus colonensis. The greatest divergence is apparent in speci-
mens from Colon, Panama, with complete intergradation in specimens on the Gulf
coast of the United States. As typical Aepsetus occurs through the range of A. colonensis,
the latter is not a geographical variant but one that occurs within the range of the
typical species.
The variant, or subspecies co/onensis, differs from typical hepsetus in having a some-
what deeper and more strongly compressed body, its greatest thickness scarcely ex-
ceeding the depth of the caudal peduncle. Its pectoral fin is longer and more pointed,
generally extending nearly or quite to the base of the pelvic fin. Its silvery lateral
band is narrower, often scarcely exceeding the width of the pupil.
Commercial Importance. Each year large quantities are marketed as food for man.
In 1939 about 21,000 pounds and in 1940 about 29,000 pounds were handled in the
New York City markets. It is economically important, however, chiefly as a forage fish.
198 Memor Sears Foundation for Marine Research
It is preyed upon by several commercially important fishes, especially the seatrouts
(Cynoscion). Water birds also feed on it extensively.
Range. Plentiful from Chesapeake Bay to the West Indies; north irregularly to
southern New England and as a stray to Maine and the outer coast of Nova Scotia;
south at least as far as Montevideo, Uruguay. It has also been recorded for Cape
Verde, Africa, but that record is in need of verification.
In some years it is numerous at Beaufort, North Carolina, appearing in large
schools, but in other years less abundant. At Beaufort and in Chesapeake Bay it was
taken chiefly with seines near the shore and rarely with trawls in somewhat deeper
water; however, in the Gulf of Mexico it was taken in deeper water more commonly.
Collections made with otter trawls, some of them in water as much as 30-40 fms.
deep, are at hand from off Corpus Christi, Texas, off Grand Isle, Louisiana, and off
St. Vincent Island and Apalachicola Bay, Florida.
Details of Occurrence.* Stray specimens of this Anchovy were reported for Halifax,
Nova Scotia, in 1931 (2r: 3), and for the mouth of the Penobscot River, Maine,
in 1930, when many others were reported as seen (8). The next most northerly
locality of record is the vicinity of Woods Hole, southern Massachusetts, where
it has been reported as abundant, at least in some years. But only one specimen
of this particular Anchovy has been reported for Rhode Island. It is less common
in New York waters than its relative mitchilli (p. 179), but it has been reported as
being locally abundant there; this applies equally to New Jersey waters and to Ches-
apeake Bay as a whole.
Farther to the south, Aepserus appears in large schools in some years along the
coast of North Carolina (see above), where mitchi/li has been reported only once (p. 179).
It is known from Georgia, the Florida Keys region, various localities along the western
coast of Florida from Cape Sable to Pensacola, the coast of Mississippi, Grand Isle
and Cameron, Louisiana, Galveston and Mustang Island, Texas, on the Mexican
coast off Tampico, Vera Cruz, and Yucatan, and the Atlantic coast of Panama. It is
widespread throughout the West Indies also, where it has been recorded for the
northern coast of Cuba, St. Thomas, Puerto Rico, Haiti, Santo Domingo, Jamaica,
Curagao, St. Kitts, Martinique, and Barbados. On the coast of Brazil it is known at
Mamanguape (near Pernambuco), Bahia, and Rio de Janeiro.
Synonyms and References:
Esox hepsetus Linnaeus, Syst. Nat., ed. 10, 1758: 314 (diagn.; type local. ““America’’; refs. to ‘Piquitinga’
Marcgrave, Brazil, and Menidia Browne, Jamaica, nonbinomials).
Atherina epsetus Bonnaterre, Tabl. Encyc. Meth. Ichthyol., 1788: 175 (descr., same fin ray counts as Linnaeus;
ref. to Linnaeus, but apparently not ed. ro).
Atherina brownii Gmelin, Syst. Nat., I, 1788: 1397 (based on Menidia Browne, pre-Linnaean).
Clupea vittata Mitchill, Trans. Lit. philos. Soc. N. Y., , 1815: 456 (orig. descr.; type local. New York; type
lost); De Kay, New York Fauna, Pt. 4: Fishes, 1842: 254 (descr.); Storer, Mem. Amer. Acad. Arts Sci.,
1846: 205, in separate (brief descr., New York).
Engraulis brownii Cuvier and Valenciennes, Hist. Nat. Poiss., 27, 1848: 41 (descr., New York, West Indian
8. Added by H. B. Bigelow.
Fishes of the Western North Atlantic 199
locals., Vera Cruz, Rio de Janeiro; also several Indian O. locals. where this species does not occur);
Giinther, Cat. Fish. Brit. Mus., 7, 1868: 389 (descr., West Indies, Gulf of Mexico, and Bahia; also
from “Libertad,” Pacific Central America, and Ceylon where the species does not occur; synon. includ.
E. tricolor, which is a valid species); Poey, Repert. Fisico-Nat. Cuba, 2, 1868: 419 (descr., Cuba); Poey,
Enumerat., Pisc. Cubana, 1875: 149 (relations., Cuba, Jamaica, and San Domingo); Yarrow, Proc.
Acad. nat. Sci. Philad., 1877: 215 (Ft. Macon, North Carolina; “not common”).
Argentina menidia Gronow, Cat. Fish., 1754: 141 (based on Menidia Browne, pre-Linnaean).
Engraulis vittatus Jordan and Gilbert, Proc. U.S. nat. Mus., 7, 1879: 385 (common, Beaufort Harbor, North
Carolina).
Engraulis hiulcus Goode and Bean, Proc. U.S. nat. Mus., 2, 1880: 343 (orig. descr.; type local. Clearwater
Harbor, Florida; USNM 23632; in bad condition, about 57 mm long, not 47 mm as stated by Goode
and Bean).
Stolephorus perthecata Goode and Bean, Proc. U.S. nat. Mus., 5, 1883: 434 (orig. descr.; type local. Pensacola,
Florida; type USNM 30483); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 442 (descr.).
Stolephorus browni Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 273 (descr., range, synon.); Swain,
Bull. U.S. Fish Comm. (1882), 2, 1883: 56 (descr., several locals. between New York and Brazil;
synon., includ. ¢rico/or); Jordan and Swain, Proc. U.S. nat. Mus., 7, 1885: 230 (Cedar Keys, Florida) ;
Bean, T. H., Bull. U.S. Fish Comm. (1887), 7, 1888: 149 (common around Great Egg Harbor, New
Jersey; eaten by weakfish); Henshall, Bull. U.S. Fish Comm. (1889), 9, 1891: 373 (Cape Sable Cr.,
Cape Romano, and Big Gasparilla, Florida); Bean, B. A., Proc. U.S. nat. Mus., £4, 1892: 93 (abund.,
Cape Charles City, Virginia); Evermann and Kendall, Bull. U.S. Fish Comm. (1892), 12, 1894: 105
(Galveston, Texas); Moore, Bull. U.S. Fish Comm. (1892), 72; 1894: 359 (abund., size, Sea Isle
City, New Jersey); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 443 (descr., range,
synon.); Smith, Bull. U.S. Fish Comm. (1897), 77, 1898: 92 (abund., season, Woods Hole, Massa-
chusetts); Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 56 (Florida locals.); Gilbert,
Proc. Wash. Acad. Sci., 2, 1900: 163 (Mamanguape, Brazil); Linton, Bull. U.S. Fish Comm. (1899),
Ig, 1901: 440 (food, parasites); Bean, T.H., 7th Rep. Forest Comm. N.Y. (1go1), 1902: 310
(synon., forage fish); Evermann and Marsh, Bull. U.S. Fish Comm. (1900), 20, 1902: 88 (descr.,
range, Puerto Rico; synon.); Schreiner and Miranda-Ribeiro, Arch. Mus. nac., Rio de J., 12, 1903:
93 (Rio de Janeiro); Bean, T. H., Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 214 (descr., synon., New
York); Linton, Bull. U.S. Bur. Fish. (1904), 24, 1905: 353 (food, parasites); Barbour and Cole, Bull.
Mus. comp. Zool. Harv., 50, 1906: 156 (Progreso, México); Breder, Zoologica, N. Y., 2 (15), 1922:
338 (Sandy Hook, New Jersey); Metzelaar, Bijdr. Dierk. Amst., 1922: 134 (Curagao).
Anchovia brownii Fowler, Rep. N. J. St. Mus. (1905), 1906: 10g (descr., New Jersey); Fowler, Rep. N. J.
St. Mus. (1906), 1907: 267, pl. 85 (ill.); Smith, N.C. geol. econ. Surv., 2, 1907: 133 (descr., range,
abund., forage fish); Jordan and Dickerson, Proc. U.S. nat. Mus., 34, 1908: 12 (Tampico, México);
Weymouth, Proc. U.S. nat. Mus., 38, 1910: 136 (Cameron, Louisiana); Fowler, Proc. Acad. nat. Sci.
Philad., 63, 1911: 216 (New Jersey, North Carolina, Florida Keys, San Domingo, Puerto Rico,
Rio de Janeiro); Sumner, Osburn, and Cole, Bull. U.S. Bur. Fish. (1911), 32, 1913: 742 (habitat,
spawn. time, food, parasites, Woods Hole, Massachusetts); Fowler, Copeia, 1915: 50 (San Domingo);
Latham, Copeia, 1919: 55 (Orient, Long Island, New York); Meek and Hildebrand, Field Mus. Publ.,
Zool., 5 (1), 1923: 204, pl. 14, fig. 1 (synon., descr., range, Colén and Porto Bello, Panama); Breder,
Zoologica, N. Y., 4 (4), 1925: 152 (large schools, Caledonia Bay, Panama); Nichols and Breder, Zoo-
logica, N. Y., 9 (1), 1927: 43 (descr., range, abund. New York to Woods Hole, Massachusetts; size);
Nichols, N. Y. Acad. Sci., To (2), 1929: 205, fig. 38 (diagn., range, abund., Puerto Rico).
Anchoviella epsetus Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67 (11), 1926: 396 (synon., descr., range;
Rio de Janeiro, “Sambaia” [probably Sambara], Brazil, and Montevideo, Uruguay, specimens are frico/or;
specimens from Key West and Tortugas, Florida, are /emprotaenia); Hildebrand and Schroeder, Bull.
U.S. Bur. Fish. (1927), 43 (1), 1928: 110 (descr., food, spawn. season, size, range, Chesapeake Bay);
Beebe and Tee-Van, Zoologica, N. Y., 10 (1), 1928: 46 (descr., range, Port-au-Prince, Haiti); Breder,
Field Bk. Mar. Fish. Atl. Cst., 1929: 71, text fig. (range, spawn., food, abund.); Jordan, Manual Vert.
Anim. NE U.S., ed. 13, 1929: 43 (descr., range); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish.
(1928), 2, 1930: 48 (synon., range); Hildebrand and Cable, Bull. U.S. Bur. Fish., 46, 1930: 388
(econ. import., spawn., develop. of eggs and young, growth rate, food); Kendall, Bull. Boston Soc.
nat. Hist., 58, 1931: 11 (Portland, Maine); Vladykov, Proc. N. S. Inst. Sci., 79 (1), 1935: 3, 555 fig. 31
200 Memoir Sears Foundation for Marine Research
(Halifax Harbor, Nova Scotia); Hubbs, Publ. Carneg. Instn. Wash., 457, 1936: 175 (discus. ident.,
Rio Champoton and Campeche, Yucatdén); Bigelow and Schroeder, Bull. U.S. Bur. Fish., 48 (20), 1936:
328 (Portland, Maine; Halifax Harbor, Nova Scotia); Howell-Rivero, Handbk. Jamaica, 1936: 1, in
reprint (Port Antonio, Jamaica); Gunter, Ecol. Monogr., 8, 1938: 337 (spawn., abund., Louisiana);
Gunter, Amer. Nat., 72, 1938: 78, 79; Gunter, Ecology, 22 (2), 1941: 203-208 (killed by freeze,
Texas); Fowler (in part, not of Linnaeus), Arqu. Zool. estad. Sao Paulo, 3 (6), 1941: 134 (Brazilian
recs., In part ¢rico/or).
Anchoa hepsetus hepsetus Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 57, fig. 21 (synon., descr.,
range).
Anchoa hepsetus colonensis Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 60, fig. 22 (orig. descr.;
type local. Colén, Panama; type USNM 117664; cf. typical Aepserus; range).
Anchoa hepsetus Gunter, Publ. Inst. mar. Sci. Texas, I (1), 1945: 32 (shallow water, temp., salinity, length
frequencies by months, sex. develop. in spring, Texas).
Doubtful References:
Anchovia brownii Jordan and Thompson, Bull. U.S. Bur. Fish. (1904), 24, 1905: 233 (Garden Key, Florida;
some, if not all, were /emprotaenia); Fowler, Proc. Acad. nat. Sci. Philad., 58, 1906: 84 (Florida Keys;
in part, if not all, /amprotaenia).
Engraulis brownii Lampe, Fisch. Dtsch. Siidpolar Exped., 1901-1903, III, Die Hochsee und Kiisten Fische,
15, Zool. 7, 1914: 216 (Cape Verde, So. Africa; ident. needs verification).
Engraulis hepsetus Fowler, Bull. Amer. Mus. nat. Hist., 70 (1), 1936: 183 (descr. based on New Jersey speci-
mens; rec. for Cape Verde after Lampe).
Negative References:
Stolephorus brownii Jordan, Proc. U.S. nat. Mus., 7, 1885: 106 (abund. Key West, Florida; USNM 35000
listed; these exam. and found to be /amprotaenia); Puyo, Bull. Soc. Hist. nat. Toulouse, 80, 1945:
100, fig. 1 (descr., French Guiana; descr. and ill. appear to be unnamed new species of Aachoviella) ;
Puyo, Faune Emp. Franc., 12, Poiss. Guiane Frang., 1949: 154 (see Puyo, 1945).
Anchovia brownii Starks, Stanf. Univ. Publ., Univ. ser., 1913: 10 (Natal, Brazil; specimens are ¢rico/or).
Anchoa lyolepis (Evermann and Marsh) 1902
Dusky Anchovy, Manjuia (Cuba)
Figure 42
Study Material. At least 22 specimens, 45-70 mm TL; juveniles, including the
type material, also at hand for comparison; from off St. Vincent Island, Florida; Mobile
Bay, Alabama; Mississippi Sound, Mississippi; Grand Isle, Louisiana; Galveston and
Aransas Pass, Texas; several places in Puerto Rico, Cuba, and St. Martin, West Indies.
Distinctive Characters. This species is quite distinctive. It is probably nearest to
lamprotaenia, from which it differs rather prominently in the lower head, the longer
snout, longer and narrower cheek, and in the more posteriorly placed anal fin.
Description. Proportional dimensions in per cent of standard length, and counts,
based on Study Material, 45-70 mm TL.
Body: depth 16.6—20. Eye: diameter 5.5—7.0.
Head: length 27—32.2. Postorbital: distance 15.4—16.7.
Snout: length 5.9—6.6. Maxillary: length 21-27.
Fishes of the Western North Atlantic 201
Mandible: length 20-22.2. Gill rakers: 15-21+19—-26 (most fre-
Anal fin: length of base 19-22.5. quently 18 or 19+ 20-22).
Pectoral fin: length 14-15.4. Fin rays: dorsal 12-15; anal 19-23;
pectoral 12-14.
Scales: ca. 40-44. Vertebrae: 41-43 (9 specimens).
Bopy very slender, rather strongly compressed, its depth 5.0—-6.0 in SL. Heap
3.1-3.7 in SL, its depth about equal to its postorbital length. Snour extending nearly
its full length beyond tip of mandible, 4.3-5.5 in head. Eye scarcely longer than
snout in large specimens, 4.25—5.0 in head. Posrorsgirat part of head slightly longer
Figure 42. Anchoa lyolepis, 63 mm TL, from Hucares, Puerto Rico, USNM 125580. Drawn by Louella
E. Cable.
than snout and eye, 1.7—2.0 in head. Maxixiary rather bluntly pointed, its upper
margin gently rounded, not quite reaching margin of opercle, 1.1§—1.3 in head. Man-
DIBLE I.2$—1.$5. CHEEK narrow, about as long as snout and eye, its posterior angle
acute, about 25°.
Dorsat fin moderately high anteriorly, its longest rays reaching nearly to tip of
last ray if deflexed, its origin usually a little nearer to base of caudal than to tip of
snout. Anat with origin under or slightly behind base of last dorsal ray, its base 4.4 5—
5.3 in SL. Pexvic inserted equidistant between origin of anal and pectoral base, some-
times a little nearer to the latter. Pecrorat rather falcate, failing to reach pelvic by
50-75°/) of diameter of eye, the last ray only about half as long as the first or upper-
most ray, 1.75—2.1 in head. AxiLtary scaLe of pectoral rather short and broad, about
50-75°/) of length of pectoral, 2.9-3.5 in head.
Color. In alcohol pale. Side of head silvery. Lateral band bright silvery, fully as
wide as eye (often dusky in specimens preserved in formalin); back and both dorsal
and caudal fins with dusky dots, these frequently present at base of anal also.
Size. The 70-mm (2.8 in.) specimen measured is the largest one seen, and it 1s
probably near the maximum size attained.
Reproduction. Specimens of two different lots taken during February in Puerto
Rico contain well-developed roe.
202 Memoir Sears Foundation for Marine Research
Variation. Specimens from the Gulf of Mexico have more numerous gill rakers
than those from farther south, but they seem to agree in other respects.
Commercial Importance. So far as known, this Anchovy is not used directly as food
by man. However, where it is common, as in Puerto Rico, it no doubt furnishes con-
siderable forage for larger predatory fishes that are used as food by man.
Range and Habitat. The range extends from the Gulf coast of the United States
through the West Indies to the Gulf of Venezuela. Although some of the specimens
examined were taken with trawls hauled at depths down to 30 fms., most of them
were seined along the shore.
Synonyms and References:
Stolephorus lyolepis Evermann and Marsh, Bull. U.S. Fish Comm. (1900), 20 (1), 1902: 89, fig. 13 (orig.
descr. based on juveniles, 30-37 mm long; type local. Culebra, Puerto Rico; type USNM 49528).
Stolephorus choerostoma Evermann and Marsh, Bull. U.S. Fish Comm. (1900), 20 (1), 1902: 88 (descr., synon.,
specimens in part re-exam., not £. choerostoma Goode).
Anchovia platyargyrea Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 216, fig. 4 (orig. descr.; type local. St.
Martin, West Indies; type ANSP 1416; cf. &. choerostoma Goode; type material, exam. by me, is
8. lyolepis E.and M.).
Anchoviella choerostoma Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 404 (specimens from e.
coast of Panama and Puerto Rico probably this species); Beebe and Tee-Van, Zoologica, N. Y., 10 (1),
1928: 47, fig. (refs., descr., distr., abund. Port-au-Price, Haiti; descr. and fig. show it to be 8. /yolepis
E. and M., not £. choerostoma Goode).
Anchoviella lyolepis Beebe and Tee-Van, Zoologica, N. Y., To (1), 1928: 47, fig. (refs., descr., Port-au-Prince,
Haiti; descr. based on juveniles); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2,
1930: 49 (synon., Puerto Rico only).
Engraulis platyargyreus Fowler, Proc. Acad. nat. Sci. Philad., 80, 1928: 468 (St. Lucia, B.W.1.).
Anchovia choerostoma Nichols, N. Y. Acad. Sci., To (2), 1929: 205, fig. 39 (refs., diagn., Puerto Rico; 8. /yolepis
E.and M., not £. choerostoma Goode).
Anchoviella platyargyrea Jordan, Evermann, and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 49 (ref. to
orig. descr.); Beebe and Hollister, Zoologica, N. Y., 79 (6), 1935: 211 (Union I., Grenadines, West
Indies).
Anchoa lyolepis Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 65, fig. 25 (synon., descr., relation.,
range); Schultz, Proc. U.S. nat. Mus., 99, 1949: 43 (synon.).
Doubtful Reference:
Engraulis (Stolephorus) argentivittatus Regan, Ann. Mag. nat. Hist., (7) 73, 1904: 257 (orig. descr.; type local.
Pas Penas, Jalisco, México; type in BMNH).
Anchoa filifera (Fowler) 1915
Figure 43
Study Material. At least 11 specimens, ranging between 55-75 mm TL. In addi-
tion to the three types from Port-of-Spain, Trinidad, 16 additional specimens from
Kingston, Jamaica, were examined.
Distinctive Characters. This species in general resembles 4. /yo/epis, from which it
differs: in the produced ray of the pectoral, the fin itself being longer; the more anterior
position of the dorsal and anal fins; and the slightly fewer vertebrae. It differs from
Fishes of the Western North Atlantic 203
A. howelli Hildebrand (3: 105), known only from Santos and Rio de Janeiro, Brazil,
in having fewer anal rays, gill rakers, and vertebrae, longer pectoral and pelvic fins,
and a slightly more posteriorly inserted pelvic fin. For details, compare the descriptions
of the two species.
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens, 55-75 mm TL.
Ficure 43. Anchoa flifera, 70 mm TL, from Kingston, Jamaica, USNM 127611. Drawn by Alice C. Mullen.
Body: depth 17.8—20. Pectoral fin: \ength (without filament)
Head: length 28-29.3. 1§.5—-16.6.
Snout: length 5.17—5.85.
Eye: diameter 5.9—7.0. Scales: ca. 39 or 40.
Postorbital: distance 14.7-16. Gill rakers: 17-19 + 20-23.
Maxillary: length 20.5—23. Fin rays: dorsal 13-15; anal 19-23; pec-
Mandible: length 18-21. toral 13 or 14.
Anal fin: length of base 18.5—21. Vertebrae: 39 or 40 (3 specimens).
Bopy strongly compressed, its depth 5.0-5.6 in SL. Heap 3.4—3.6, its depth
slightly exceeding the postorbital length. Snour long, projecting nearly its full length
beyond mandible, only a little shorter than eye, 5.0—5.5 in head. Eve small, 4.2-4.8.
PosrorsiTaL part of head 1.6—1.8 in head. Maxttiary not sharply pointed, its upper
margin rounded, reaching nearly to margin of opercle, 1.25—1.3 in head. Manp1BLe
1.4—1.5. CHEEK narrow, equal to eye and snout, its posterior angle sharp, about 30°.
Dorsat fin rather high anteriorly, its longest rays reaching nearly to tip of last
ray if deflexed, its origin slightly nearer to tip of snout than to base of caudal. ANAL
with origin nearly under the beginning of the last third of dorsal base, the base 4.8—
5.4 in SL. Petvic reaching much more than halfway to anal, inserted a little nearer
to pectoral base than to anal origin. PecroraL with upper ray produced, its filament
(often broken) extending to, and possibly beyond, tip of pelvic in perfect specimens,
the fin, without filament (second ray), reaching base of pelvic, 1.6—-1.8 in head.
204 Memoir Sears Foundation for Marine Research
AXILLARY SCALE of pectoral about 66°/, of length of fin (without filament), 2.8—3.5
in head.
Color. In alcohol pale. Side of head and lower part of side silvery. Silvery lateral
band as wide as eye (often dusky in specimens preserved in formalin). Upper parts of
head and back with numerous punctulations not arranged in longitudinal series.
Size. The longest specimen seen (type) is 75 mm (3 in., 61 mm SL).
Range. This species, as here understood, is known from Puerto Rico, Jamaica,
Haiti, and Trinidad.
Synonyms and References:
Anchovia filifera Fowler, Proc. Acad. nat. Sci. Philad., 1915: 524, fig. 2 (orig. descr.; type local. Port-of-Spain,
Trinidad; type ANSP 45080; cf. 4. platyargyrea = A. lyolepis and A. choerostoma. Gill rakers given as
25 + 22, total anal rays 25, depth 4.75. My counts and measurements give gill rakers 18 + 22, anal
rays 23, depth 5.0. Orig. fig. shows mandible projecting too far in advance of vertical from anterior
margin of eye; anal fin rather too far back, its origin actually well in advance of vertical from base of last
dorsal ray).
Anchoviella filifera Fowler, Proc. biol. Soc. Wash., 43, 1930: 146 (Jamaica); Jordan, Evermann, and Clark,
Rep. U.S. Comm. Fish (1928), 2, 1930: 50 (ref.).
Anchoviella longipinna Beebe and Tee-Van, Zoologica, N. Y., To (1), 1928: 48, fig. (orig. descr.; type local.
Bizoton, Haiti; type NYZS 7460; also Port-au-Prince, Haiti; type material not seen; descr. and fig.
agree well with 4. f/ifera from Jamaica and Trinidad).
Anchoa filifera Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 105, fig. 45 (synon., descr., range).
Genus Anchoviella Fowler 1911
Anchoviella Fowler, Proc. Acad. nat. Sci. Philad., 43, 1911: 211; genotype by original designation, Engraulis
perfasciatus Poey 1861.
Amplova Jordan and Seale, Copeia, 1925: 313 type species Axchovia brevirostris Meek and Hildebrand 1923
equals Amplova balboae Jordan and Seale 1925.
Characters. Bovy usually slender, its depth less than 25°/, of SL (except in pa/-
lida, 26-28.5°/,). Git RaKERs usually fewer than 35 on lower limb (except in pa/-
lida, which may have as many as 45), not known to increase in number with age.
Anat fin rather short, with fewer than 30 rays, its origin usually under or posterior to
middle of dorsal base (somewhat in advance of middle of dorsal in d/ackburni and
jamesi). VERTEBRAE 39-45.
Size. Although a maximum length of 155 mm has been reported for eurystole,
the largest in the collections studied is only 130 mm TL, and the other species are
smaller.
Remarks. Anchoviella obviously is closely related to Anchoa, from which it differs
in the shorter and distally square-to-round maxillary. This structure, variable in length
in both genera, may reach only a little beyond the vertical from the posterior margin
of the eye, as in Anchoviella jamesi, or it may reach to the joint of the mandible, as in
Anchoviella eurystole. Indeed, Jordan and Seale (6: 31) erected a new genus, dmp/lova,
on the basis of the very short maxillary in 4. ba/boae, a species common on the Pacific
Species
THEMED? ooopononocsovceote
PITAR TAA ooo oon opoaonaacc
(NIGIMIDIZ noooovonereqaase
LEGHRD pbooonbooonUCone
pallida
AOR a5 00 go oC ed OHH oDOUS
WEDICENIOSLOLE seckctatntatel-elerelatel ete
(MEET ITRTI 6 3.5 SOR ECB 000 GEC
JET? onc ba opooKDo0Gde0a te - —
* Anal
Fishes of the Western North Atlantic 205
Table V. Frequency Distribution of Anal Rays in Anchoviella.*
of Anal Rays ————————_—___.,
20) 20) 22 nz Geez Ameo m2 ORR 7
Zz,
=
3
o
io
I
wat
cl
~_
al
co
nN i]
nnn
| nono &
LSI A eta es
|
Teele ects lt
aie allies oe aes alle? 3c
| bee lal | | | | |
i]
4
rays given as g—II for cayenmensis, see p. 212.
Table VI. Frequency Distribution of Gill Rakers on Upper Limb in dachoviella.*
Species
-————_ Number of Gill Rakers on Upper Limb of First Arch ———————_.,
TOMI T2 L314 15) LO! 17 18) LO) 20) 20e2223N24e2 haz 27e2SezONszONsTEs 233634
PEGE ononones SS SS SS ae ae a CO
CVG ooo toces SS SSS SS SS SS SS SS SS SS FO SS SS SS
perfasci
i me Bei) G Ga SS SS eS SS
brevirostris ...... —— — 4 2 EE
QUIQNeNSIS 6.66606 —— a ee ee ee OO ee
alleni.
pallida
CLONE GI ar a —— — — — — 7 8B — — — — — —
lepidentostole...... — — — — — — — 3 1—- — eee ee ee eS
blackburni....... Tn ee ee
EERE. oonedecbos a ellie
* When the number of counts exceeds the number of specimens listed in the text, the gill rakers on both an-
terior arches in at least some of the specimens were counted.
Table VII. Frequency Distribution of Gill Rakers on Lower Limb in Anchoviella.*
Species
eurystole.......
estauquae......
perfasciata.....
brevirostris ....
QUIQNENSIS......
alleni
pallida ........
elongata .......
lepidentostole ...
blackburni .....
jamesi
— Number of Gill Rakers on Lower Limb of First Arch ——————————_—_—_,
15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45
* When the number of enumerations exceeds the number of specimens listed in the text, the gill rakers have been counted on both
anterior arches in at least some of the fish.
206 Memotr Sears Foundation for Marine Research
coast of Panama. However, there are species quite intermediate between the type species
of Amplova (balboae) and that of Anchoviella (perfasciata). For that reason Amplova was
reduced to subgeneric rank (3: 108).
The very short maxillary in some species of 4nchoviella furnishes a near transition
from the Engraulidae to the Clupeidae; but other family characters, such as the pro-
jecting snout and the anteriorly more-or-less embedded and widely separated premaxil-
laries, remain as in other Engraulidae.
Range and Habitat. The genus seems to be restricted to the two coasts of America,
ranging in the Atlantic from Woods Hole, Massachusetts, at least to southern Brazil,
and in the Pacific from southern California to northern Peru. Most of the species enter
fresh water. In fact, several of the South American species from the Atlantic drainage
are known only from freshwater lakes and streams, a few of them having been taken
a great distance from the sea.
Key to Species of the Western Atlantic
1a. Maxillary usually extending beyond posterior rim of orbit by a distance equal to
or greater than diameter of pupil. Subgenus Axchoviella.
2a. Origin of anal posterior to middle of dorsal base, sometimes entirely behind
dorsal; not more than 20 anal rays.
3a. Origin of anal clearly posterior to dorsal base, generally under or behind
tips of dorsal rays if deflexed; gill rakers 24-28 + 27-33.
4a. Longest rays of dorsal failing to reach tip of last ray if deflexed;
eye 3.75—4.5 in head, 2.1-2.5 in postorbital; maxillary 15.8-17.8°/,
of SL; postorbital 13.8-15.5°/) of SL.
eurystole (Swain and Meek) 1884, p. 208.
4b. Longest rays of dorsal reaching beyond tip of last ray if deflexed;
eye 4.4-4.8 in head, 2.5—2.75 in postorbital; maxillary 17.5—18.7°/,
of SL; postorbital 15.5—15.8°/) of SL.
estauquae Hildebrand 1943, p. 211.
3b. Origin of anal under or slightly behind base of last 2 or 3 rays of dorsal;
gill rakers usually 13-23 + 19-28.
sa. Only 9-11 anal rays; depth of body less than 14°/, of SL.
cayennensis (Puyo) 1945, p. 212.
5b. 15 anal rays or more; depth of body more than 15°/, of SL.
6a. Pectoral with 14-16 rays; gill rakers 19-23 + 24-28.
perfasciata (Poey) 1860, p. 213.
6b. Pectoral with 11-13 rays; gill rakers 18 or fewer on upper
limb, generally fewer than 25 on lower limb.
7a. Gill rakers 13 or 14+18 or 19; pectoral failing to reach
base of pelvic by diameter of eye, 11.2-13.8°/, of SL,
1.7—2.0 in head. brevirostris (Giinther) 1868, p. 216.
Fishes of the Western North Atlantic 207
7b. Gill rakers 16-18 + 22—26; pectoral failing to reach pelvic
by a distance considerably less than eye, 14.7-16.8°/) of
SL, 1.4-1.6 in head.
8a. Eye 7.0-7.5°/, of SL, 3.2-3.5 in head; cheek fully as
long as eye, with a definite posterior angle of about
60°; axillary scale of pectoral about 50°/, of fin, 2.4
in head; a median dark line posterior to anal fin.
guianensis (Eigenmann) 1912, p. 217.
8b. Eye 5.5-6.25°/, of SL, 3.7—4.1 in head; cheek scarcely
as long as eye, without a definite posterior angle; axil-
lary scale of pectoral about 75°/) of length of fin, 2.0—
2.2 in head; no median dark line posterior to anal fin.
alleni (Myers) 1940, p. 219.
2b. Origin of anal under, or in advance of, middle of dorsal base; 22-29 anal rays.
ga. Anal with 28 or 29 rays; pectoral extending to midlength of pelvic.
nattereri (Steindachner) 1879, p. 220.
gb. Anal with fewer than 28 rays; pectoral usually failing to reach base
of pelvic.
10a. Gill rakers 28-34 + 36-45; depth about 26-28.5°/, of SL; pec-
toral extending slightly beyond base of pelvic, 19—20°/, of SL.
pallida (Starks) 1913, p. 221.
1ob. Gill rakers 18 or fewer on upper limb, 24 or fewer on lower limb;
depth 25°/, of SL or less; pectoral failing to reach base of pelvic,
16.7%), of SL.
11a, Maxillary extending beyond orbit a distance about equal to
diameter of eye, 15-20°/, of SL; mandible 15.8—18.8; gill
rakers 17 or 18+19—24; origin of anal under or slightly
behind middle of dorsal base.
12a. Depth of body 17.5-20°/) of SL; head 25.5-28°/);
postorbital part of head 12—13.3°/); mandible 17.8—
18.8°/, ; silvery lateral band poorly defined, narrower than
eye. elongata (Meek and Hildebrand) 1923, p. 223.
12b. Depth of body 22-25°/) of SL; head 22.7—-23.7°%o;
postorbital 10.7-11°/); mandible 15.8—16.3°/,; silvery
lateral band well defined, broader than eye.
lepidentostole (Fowler) 1911, p. 224.
11b. Maxillary extending beyond orbit a distance equal to only
half of diameter of eye, 12-15°/, of SL; mandible 12.3-14.8;
gill rakers 10-12 + 15-17; origin of anal well in advance of
middle of dorsal base. Jb/ackburni Hildebrand 1943, p. 226.
1b. Maxillary extending a little less than diameter of pupil beyond orbit.
Subgenus dmp/ova.
jamesi (Jordan and Seale) 1926, p. 227.
208 Memoir Sears Foundation for Marine Research
Anchoviella eurystole (Swain and Meek) 1884
Silver Anchovy
Figure 44
Study Material. At least 29 specimens, 55-130 mm, from: Woods Hole and
Menemsha Bight, Massachusetts; Noank, Connecticut; Long Island, New York;
Brigantine Bay, New Jersey; and Beaufort, North Carolina.
Distinctive Characters. A. estauquae is the nearest relative of this species. The rela-
tionship is shown in the account of that species (p. 211).
ZgZ
LE ELLES
Ficure 44. Anchoviella eurystole, 75 mm TL, from Woods Hole, Massachusetts, USNM 77778. Drawn by
Louella E. Cable.
Description. Proportional dimensions in per cent of standard length, and counts,
based on Study Material, specimens 55-130 mm.
Body: depth 15.4—-19. Pectoral fin: length 12.5-13.7.
Head: length 25-28.5.
Snout: length 3.85—5.0. Scales: ca. 40-45.
Eye: diameter 5.7—7.0. Gill rakers: 24-28+27-33 (most fre-
Postorbital: distance 13.8-15.5. quently 25 or 26+28—30).
Maxillary: length 15.8—17.8. Fin rays: dorsal 13-16; anal 15-18
Mandible: length 15-18.5. (rarely 19); pectoral 15 or 16.
Anal fin: length of base 12.5—-16. Vertebrae: 43-45 (3 specimens).
Bopy slender, not strongly compressed, its depth 5.25—6.5 in SL. Heap 3.5—4.0,
its depth equal to length of postorbital part of head. Snour projecting about half of its
length beyond mandible, only a little shorter than eye in large examples, 4.75—-6.5
in head. Eve 3.75—4.5 in head, 2.1-2.5 in postorbital length of head. PosrorsrTat
part of head 1.6—-1.8 in head. Maxittary rather narrow, rounded distally, extending
beyond orbit by a distance equal to about 1.3 diameter of eye, failing to reach joint of
mandible by about half of diameter of pupil, 1.5;—1.7 in head. ManpiBLE 1.45-1.75.
CueEEK only a little shorter than snout and eye, its posterior angle about 45°. GILL
RAKERS about 75°/, of length of eye, not broad or close-set, with moderate serrations
on inner edge.
Dorsat fin low anteriorly, the longest rays failing to reach tip of last ray if de-
Fishes of the Western North Atlantic 209
flexed, its origin usually somewhat nearer to tip of snout than to base of caudal. ANnaL
small, its origin wholly behind dorsal base, most frequently about under tips of longest
deflexed rays, its base 6.25-8.0 in SL. Petvic inserted nearly an eye’s diameter in
advance of dorsal and usually somewhat nearer to pectoral base than to anal origin,
reaching much less than halfway to anal. Pecrorat usually failing to reach pelvic by
more than diameter of eye, 1.8—2.2 in head. AxrLLary scaLe of pectoral long and
narrow, generally failing to reach tip of fin by about half of diameter of pupil, 2.2 5—
3.0 in head.
Color. In alcohol, some rather pale above, others bluish. Side of head and lower
part of side on some silvery. Silvery lateral band about as broad as eye in small speci-
mens, broader and less distinctly outlined ventrally in large ones.
Size. Although a maximum total length of 155 mm (6.2 in.) has been reported,
the largest example in the collections examined in this study is only 130 mm TL.
Development. The information on the development of the eggs and larvae that
follows is based mostly on the work of Kuntz and Radcliffe (ro: 116), who took the
eggs in tows during August 1915 off Gay Head, Massachusetts.
The egg is buoyant, highly transparent, and ellipsoidal, having a long axis of
I.15—I.25 mm, a short axis of 0.55—0.8 mm. The yolk, as usual among anchovies, has
a network of lines on its surface, giving the appearance of large cells. In this anchovy,
as in other species studied, the blastodisc forms at one end of the elongated egg. The
egg is similar to that of dxchoa hepsetus but seems to be slightly shorter though
scarcely narrower. The development, so far as known, is identical in the two species.
The newly hatched fish is about 3.0mm long, being slightly shorter than the
newly hatched larva of 4. hepserus, in keeping with the rather smaller egg. The body is
very slender, and the vent as usual in the larvae of anchovies is situated very far back, in
this species about at the beginning of the distal fourth of the body. Black chromato-
phores occur in a series along the intestine posterior to the yolksac and at the base of
the ventral finfold behind the vent. A day or so after hatching, the yolk is virtually all
absorbed and the fish has grown to a length of about 4.0 mm. The young fish, when
5.2 mm long, has developed a large oblique, terminal mouth, the fins are becoming
differentiated, and the convoluted intestine, incompletely invaginated, is conspicuous.
The chromatophores have become less conspicuous than in the earlier stages. The
development in older stages apparently remains unknown.
Spawning. Although the duration of spawning has not been determined, it is at
its height during July and August (15: 44).
Range and Habitat. This anchovy ranges from Woods Hole, Massachusetts, at
least as far south as Beaufort, North Carolina. The southern limit of its range remains
undetermined, for no specimens of the genus are at hand or seem to be reported for
the coast between Beaufort, North Carolina, and Melbourne Beach, Florida; those from
the last mentioned locality are 4. perfasciata. During some summers this anchovy is
common at Woods Hole; during other summers it seems to be missing. Off Connecti-
cut, New York, and New Jersey it appears only occasionally. It is not recorded for
14
210 Memoir Sears Foundation for Marine Research
Chesapeake Bay; and for North Carolina it is known by only one specimen taken
during the summer of 1913 in inside waters at Beaufort, where intensive collecting was
carried on during many years. It has been stated often that the home of the Silver
Anchovy is in the Gulf Stream. The basis for this statement is unknown and certainly
stands in need of verification, especially as it has been rather definitely established
that the specimens known from North Carolina northward represent a species distinct
from the examples from the West Indies northward to Melbourne Beach, Florida.
All records of argyrophana based on northern specimens apparently apply to eurystole
as herein understood.
Synonyms and References:
Stolephorus perfasciatus Swain (not of Poey), Bull. U.S. Fish Comm. (1882), 2, 1883: 55 (descr. of one from
Woods Hole, Massachusetts; indicated as orig. descr. of 8. eurystole by Swain and Meek); Jordan and
Gilbert (not of Poey), Bull. U.S. nat. Mus., 16, 1883: 273 (descr. of one from Woods Hole, Mass.,
apparently the same one used by Swain; West Indies includ. in the range where ewrysto/e does not occur);
Bean, T. H. (not of Poey), Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 217 (descr., Noank, Connecticut,
and Gravesend Bay, Long Island, New York; in part exam. by me and found to be J. eurystole).
Stolephorus eurystole Swain and Meek, Proc. Acad. nat. Sci. Philad., 1885: 35 (descr. by Swain and by Jordan
and Gilbert above referred to as orig.; type local. Woods Hole, Mass.; type destroyed by fire. This species
was said to differ from Exgraulis perfasciatus Poey chiefly in the greater number of anal rays, a broader
lateral band, and a larger size; this northern species apparently reaches a somewhat larger size, but at most
there is only a small average difference in the number of anal rays, and the width of the lateral band varies
with age. For diagn. char., see Key); Bean, T.H., Bull. U.S. Fish Comm. (1887), 7, 1888: 150,
pl. 3, fig. 19 (Ocean City and Longport, New Jersey).
Stolephorus argyrophanus Jordan and Evermann (not of Cuvier and Valenciennes), Bull. U.S. nat. Mus., 47
(1), 1896: 444 (descr. apparently based on type of £. ewrysto/e; range “Gulf Stream, occasionally north-
ward”); Smith (not of C. and V.), Bull. U. S. Fish Comm. (1897), 77, 1898: 92 (occur. assoc. with other
species round Woods Hole, Massachusetts); Bean, T. H. (not of C. and V.), Bull. N. Y. St. Mus., 60,
Zool. 9, 1903: 216 (descr., Ocean City, New Jersey, and Fire Island, New York).
Anchovia eurystole Fowler, Rep. N. J. St. Mus. (1905), 2, 1906; 109 (descr., Ocean City and Longport, New
Jersey; ident. of this species and Exgraulis argyrophanus C. and V. questioned); Fowler, Rep. N. J.
St. Mus. (1906), 1907: 267, fig. (name); Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 219
(Corson Inlet, New Jersey); Fowler, Proc. Acad. nat. Sci. Philad., 64, 1912: 34 (Long Beach, New
York).
Anchovia argyrophana Sumner, Osburn, and Cole (not of C. and V.), Bull. U.S. Bur. Fish (1911), 3Z (2),
1913: 743 (abund., where and how taken, Woods Hole, Massachusetts; parasites); Kuntz and Rad-
cliffe (not of C. and V.), Bull. U.S. Bur. Fish., 35, 1917: 116 (embryol., larval develop.); Nichols
and Breder (not of C. and V.), Zoologica, N. Y., 9 (1), 1927: 43, fig. (diagn., distr., life hist., size);
Hildebrand (not of C. and V.), Copeia, 1941: 224 (Beaufort, North Carolina).
Anchoviella argyrophana Jordan and Seale (not of C. and V.), Bull. Mus. comp. Zool. Harv., 67, 1926: 402
(descr. of one from Provincetown, Mass.; argyrophana and eurysto/e regarded ident. by Jordan from exam.
of type of former in Paris, which had rg anal rays instead of 17, as stated in orig. descr. But number of
anal rays does not distinguish this species from perfasciata); Breder (not of C. and V.), Field Bk. Mar.
Fish. Atl. Cst., 1929: 70, 71 (defined in key; range; “Gulf Stream species’; eggs, size); Jordan (not of
C. and V.), Manual Vert. Anim. NE U.S., ed. 13, 1929: 43 (descr., range); Jordan, Evermann, and
Clark (not of C. and V.), Rep. U. S. Comm. Fish. (1928), 2, 1930: 48 (range; ewrysto/e in synon.).
Anchoviella eurystole Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 112, fig. 47 (descr., cf. per-
fasciata, range).
Doubtful References:
Anchovia perfasciata Nichols and Breder (probably not of Poey), Zoologica, N. Y., 9 (1), 1927: 42, fig. (diagn.,
N.Y. City area; distingu. char. of this species and one rec. as argyrophana not stated correctly; fig.
Fishes of the Western North Atlantic 211
looks like perfasciata; as it is the same one used by Nichols [1929: 204], it may have been based on a
West Indian specimen).
Anchoviella perfasciata Greeley, 28th Rep. N. Y. Conserv. Dept., Suppl., 15 (2), 1939: 83 (rare at Long Island;
prabably 4. eurysto/e).
Anchoviella estauquae Hildebrand 1943
Figure 45
Study Material. Holotype and 3 paratypes, about 77-92 mm TL (caudal fins
damaged), 65—77 mm SL, collected on a sandy beach in Estauques Bay, Gulf of Vene-
zuela, by the U.S.S. Niacara, on February 20, 1925.
Ficure 45. Anchoviella estaugque, type specimen, 82mm TL, 72 mm SL, from Gulf of Venezuela, USNM
119795. Drawn by Louella E. Cable.
Distinctive Characters. This species is very close to 4. eurystole, from which it is
difficult to separate it with the meager material at hand. The number of gill rakers is
within the range of eurysto/e, but the rakers seem to be a little longer, certainly broader,
apparently closer together, and the serrations on their inner edges are slightly smaller.
The eye appears to be a little smaller and the postorbital part of the head is slightly
longer. The dorsal fin is higher anteriorly and somewhat different in shape; the anal
fin is placed farther back.
Description. Proportional dimensions in per cent of standard length, and counts,
based on type material, 65-76 mm SL.
Body: depth 15.3-19.2. Anal fin: length of base 13.3-15.3.
Head: length 26—27.7. Pectoral fin: length 12.8-14.3.
Snout: length 5.0-5.55.
Eye: diameter 5.7—6.16. Scales: ca. 44-46.
Postorbital: distance 15.5—15.8. Gill rakers: 26-28 + 30-32.
Maxillary: length 17.5-18.7. Fin rays: dorsal 13-15; anal 15-173 pec-
Mandible: length 17.3-18.5. toral 15 or 16.
Vertebrae: 4§ (1 specimen).
212 Memoir Sears Foundation for Marine Research
Bopy not strongly compressed, its depth 5.2-6.5 in SL. Heap 3.6—3.7, its depth
scarcely equal to its postorbital length. Snour projecting rather more than half of its
length beyond mandible, only a little shorter than eye, 5.0—5.3 in head. Eye 4.4—4.8
in head, 2.5—2.75 in postorbital length of head. Posrorsirat part of head long, 1.73-
1.77 in head. Maxitiary rather narrow, rounded distally, extending beyond orbit a
distance equal to about 1.3 diameter of eye, not quite to joint of mandible, 1.5—1.6 in
head. Manp1BLeE 1.48—1.53. CHEEK nearly as long as snout and eye, its posterior angle
about 40°. GILL RaKeERs scarcely as long as eye, broad and close-set, with minute ser-
rations on inner edge.
Dorsat fin with longest rays reaching beyond tip of last ray if deflexed, its origin
about equidistant between tip of snout and base of caudal. Anat with origin well behind
tips of longest deflexed rays of dorsal, its base 6.5—7.5 in SL. Pztvic inserted about
half of diameter of eye in advance of dorsal and nearly equidistant between base of pec-
toral and anal origin, reaching rather less than halfway to anal. Pecrorat usually failing
to reach pelvic by rather more than diameter of eye, 1.9—2.5 in head. AxILLARY SCALE
of pectoral long, narrow, failing to reach tip of fin by a distance equal to or less than
diameter of pupil, 2.3—2.7 in head.
Color. In alcohol, pale (originally in formalin). Sides of head bright silvery.
Lateral band bright silvery, fully as broad as eye.
Range. This species is known only from the Gulf of Venezuela.
References:
Anchoviella estauquae Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 115, fig. 48 (orig. descr.; type
local. Estauques Bay, Gulf of Venezuela; type USNM 119795; cf. 4. eurystole and A. perfasciata);
Schultz, Proc. U.S. nat. Mus., 99, 1949: 47 (ref., type and paratypes listed).
Anchoviella cayennensis (Puyo) 1945
Jamais-gouté
Study Material. None.
Distinctive Characters. The short maxillary, extending about a pupil’s diameter
posterior to a vertical at end of eye, the elongate body, with the depth more than 7
times in SL, and the very short anal, with 9-11 rays and with its origin below that of
the last dorsal ray, show that this species is a very distinctive one, if Puyo’s descriptions
are reliable.
Description. Based on published accounts and figures by Puyo (see References
below). Bopy very elongate, lightly compressed anteriorly and progressively more
rounded posteriorly (in transverse section); depth a little more than 7 times in total
body length (SL). Heap a little more than 5 times in same; a sort of transverse keel
present on superior surface of head. Snour somewhat rounded. Eye equal to snout
(pupil shown as abnormally large in most of Puyo’s drawings, probably in error).
Maxitiary rounded at the end, extending about a normal pupil’s diameter posterior
Fishes of the Western North Atlantic Bio
to vertical at posterior border of eye (in Puyo’s published figure). Lower Jaw slightly
shorter than the upper.
Dorsat fin origin midway between tip of snout and caudal base. Anat fin origin
under base of last dorsal ray. Fin rays: dorsal 12, anal 9-11. Pscrorat fin slightly
less than twice the length of ventral, extending somewhat more than half of the distance
from its origin to that of the pelvic.
Color (from Puyo). On the whole, body clear blackish (apparently in formalin);
opercular bones iridescent. A black longitudinal band on each side; band towards
posterior part of body giving off irregular transverse projections from its lower border;
lateral band merging gradually into an irregular streak on middle caudal rays; dark
band on body bordered below by a silvery band wider than eye diameter. Dorsal and
anal pale. Pectoral and pelvic hyaline. Caudal lighter than the body, its border lightly
tinted with brown.
Size. Puyo’s two type specimens, the only examples so far recorded, were 130 and
135mm SL.
Habits. Said to be confined to estuaries and river mouths.
Range. Known only from the coast of French Guiana. The types were taken in the
Cayenne River, as far up as Macouria,® but fishermen told Puyo they found this species
also in the Mahury River and at the mouth of the Kourou.
References:
Stolephorus cayennensis Puyo, Bull. Soc. Hist. nat. Toulouse, 80, 1945: 101, fig. 1 (descr., Cayenne R., French
Guiana); Puyo, Faune Emp. Frang., 12, Poiss. Guyane Frang., 1949: 157, fig. 80 (descr.).
Anchoviella perfasciata (Poey) 1860
Flat Anchovy, Manjua (Cuba)
Figure 46
Study Material. At least 41 specimens, 60-105 mm TL, from Puerto Rico,
Jamaica, Cuba, and St. Lucia; also some poorly preserved specimens from off Mel-
bourne Beach, Key West, Tortugas, and Pensacola, Florida, and from Mississippi
Sound, probably belonging to this species.
Distinctive Characters. This species differs from 4. eurystole principally in having
a larger eye, a higher dorsal fin, a more anteriorly placed anal fin, a longer pectoral fin,
and fewer gill rakers.
Description. Proportional dimensions in per cent of standard length, and counts,
based on at least 41 specimens, 60-105 mm TL.
Body: depth 15.4—18.2. Snout: length 4.0-5.25.
Head: length 23.3-25.6. Eye: diameter 6.6—7.7.
g. Stieler’s Atlas shows Macouria to be on the coast, as is Kourou.—Y. H. O.
214 Memoir Sears Foundation for Marine Research
Postorbital: distance 11.7—13.3. Scales: ca. 40-44.
Maxillary: length 15.2-17.3. Gill rakers: 19-23+ 24-28 (most fre-
Mandible: length 15-17.2. quently 19-21+ 26 or 27).
Anal fin: length of base 13.7-15.3. Fin rays: dorsal 12-15; anal 15-18; pec-
Pectoral fin: length 12.2-14.2. toral 14-16.
Vertebrae: 42-44 (7 specimens).
Bopy not strongly compressed, its depth 5.5—6.5 in SL. Heap 3.9—4.3 in SL, its
depth slightly exceeding its postorbital length. Svour much shorter than eye, extending
about half of its length beyond mandible, 4.2-—5.75 in head. Eve 3.25—4.0 in head, 1.6—
Ficure 46. Anchoviella perfasciata, 1oomm TL, 83 mm SL, from Cuba, USNM 37472. Drawn by Louella
E. Cable.
2.0 in postorbital length of head. Posrorsirat 1.9—2.1 in head. Maxitary slightly
expanded and rounded distally, generally extending about an eye’s diameter beyond
rim of orbit, failing to reach joint of mandible by nearly diameter of pupil, 1.4-1.55
in head. Manprsiz 1.4—1.66. Cueexk slightly longer than eye, its posterior angle about
60°. Grtt RaKeRs shorter than eye, broad, not close-set, with short coarse serrations
on inner edge.
Dorsat fin with longest rays reaching to, or more usually beyond, tip of last ray if
deflexed, its origin variable, most frequently equidistant between caudal base and tip
of snout. ANat originating under or slightly behind base of last dorsal ray, its base 6.5—
7.25 in SL. Pexvic inserted nearly an eye’s diameter in advance of dorsal and about
equidistant between base of pectoral and anal origin, reaching much less than halfway
to anal. Pecrorat failing to reach pelvic base by about an eye’s diameter, 1.7—2.0 in
head. Axitiary scaLe of pectoral narrow, failing to reach tip of fin by less than dia-
meter of pupil, 1.9—2.3 in head.
Color. In alcohol, pale. Sides of head silvery. Silvery lateral band distinct, generally
nearly as broad as eye; some specimens with dusky punctulations on back extending
onto caudal fin.
Size. A total length of about 105 mm (4.2 in.) is attained.
Variation. Certain poorly preserved specimens from Melbourne Beach, Key West,
Tortugas, and Pensacola, Florida, in the Study Material above, appear to be more or
Fishes of the Western North Atlantic 2s
less intermediate between this species and eurystole. The gill rakers, which are difficult
to count because of the rather small size and poor condition of the specimens, seem to
be rather more numerous than in West Indian specimens. The shape of the dorsal, the
length of the pectoral, and the size of the eye all agree with specimens from the West
Indies, but the position of the anal is rather intermediate.
Range and Habitat. This Anchovy apparently ranges from the West Indies to
Florida and the Gulf Coast, and probably to North Carolina. It has been reported for
New York (15: 42), but none from New York and neighboring states has been avail-
able for study. This fish seems to be common, though probably not numerous, in the
West Indies. Northward it is apparently only a stray. Most of the specimens studied
were taken in shallow water. A few, one from off Melbourne Beach, Florida, and two
from Mississippi Sound, Miss., were taken in somewhat deeper water with trawls. It
is not known to enter brackish water.
Synonyms and References:
Engraulis perfasciatus Poey, Memorias, 2, 1860: 312 (orig. descr.; type local. Cuba; type MCZ 17955; cf.
several species); Giinther, Cat. Fish. Brit. Mus., 7, 1868: 391 (descr., San Domingo, Cuba; doubtfully
distinct from £. argyrophanus Cuvier and Valenciennes).
Stolephorus perfasciatus Swain and Meek, Proc. Acad. nat. Sci. Philad. (1884), 1885: 34 (descr., Key West,
Florida, and Havana, Cuba); Henshall, Bull. U.S. Fish Comm. (1889), 9, 1891: 373 (more abund.
at Marco than elsewhere in w. Florida); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896:
441 (descr., range); Evermann and Marsh, Bull. U.S. Fish Comm. (1900), 20, 1902: 88 (descr.,
range, Aguadilla, Puerto Rico); Metzlaar, Bijdr. Dierk. Amst., 1922: 136 (Curagao, taken with 8.
brownii and Fenkinsia lamprotaenia).
Anchovia perfasciata Jordan and Thompson, Bull. U. S. Bur. Fish. (1904), 24, 1905: 233 (Tortugas, Florida);
Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 211 (Aguadilla, Puerto Rico); Nichols, New York Acad.
Sci., ZO (2), 1929: 204, fig. 34 (diagn., distr., Puerto Rico); Hildebrand, Copeia, 1941: 224 (Beaufort,
North Carolina).
Anchoviella perfasciata Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 394 (descr. of “type” from
Cuba; range, Florida Keys to Cuba); Jordan, Manual Vert. Anim. NE U.S., ed. 13, 1929: 44 (descr.,
range, New York to Cuba); Breder, Field Bk. Mar. Fish. Atl. Cst., 1929: 70, fig. (range; strays north-
ward to N. Y.); Fowler, Proc. Acad. nat. Sci. Philad., 82, 1930: 269 (Grenada, B. W. 1.); Jordan,
Evermann, and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 47 (range, Florida Keys to Jamaica);
Howell-Rivero, Bull. Mus. comp. Zool. Harv., 82, 1938: 172 (types MCZ 17955; 9 specimens, 82—
103 mm, includ. roo-mm specimen as holotype, Poey’s No. 422); Longley and Hildebrand, Publ.
Carneg. Instn. Wash., 535, 1941: 13 (Tortugas, Florida; taxon. notes); Hildebrand, Bull. Bingham
oceanogr. Coll., 8 (2), 1943: 116, fig. 49 (descr., cf. 4. eurystole, range).
Doubtful References:
Engraulis argyrophanus Cuvier and Valenciennes, Hist. Nat. Poiss., 1848, 2Z: 49 (orig. descr.; type local.
“equitorial Atlantic”; cf. 4. epsetus; specimen said to have been taken in equatorial Atlantic on a voy-
age from Europe to Batavia. The specimen, therefore, may not have been taken near America, and the
description is too inadequate for positive identification with any American species. Thus it seems inadvis-
able to use this name. Recent writers generally have referred specimens possessing a short maxillary,
taken between New Jersey and Cape Cod, to argyrophanus, a procedure that in the light of present
knowledge seems to be incorrect).
Stolephorus miarchus Jordan, Proc. U.S. nat. Mus. (1884), 7, 1885: 106 (Key West, Florida; indistingu. from
types of this species; probably young 4. perfasciata).
Anchovia perfasciata Nichols and Breder, Zoologica, N. Y., g (1), 1927: 42, fig. (diagn., vic. of New York
City, range; distingu. characters of this species and of eurystole [argyrophanus] are not stated correctly;
216 Memoir Sears Foundation for Marine Research
the figure looks somewhat like perfasciata; it is the same one used by Nichols [1929: 204] and may have
been based on a West Indian specimen).
Negative References:
Stolephorus perfasciatus Swain, Bull. U.S. Fish Comm. (1882), 2, 1883: 55 (descr. of specimen from Woods
Hole, Massachusetts, which was 4. eurysto/e); Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1883: 272
(descr. of specimen from Woods Hole, Mass., which was 4. eurystole; West Indies includ. in range,
where eurystole does not occur); Bean, T. H., Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 217 (descr. of
specimens from Noank, Connecticut, and Gravesend Bay, Long Island, N. Y.; specimens, in part at least,
exam. by me and found to be 4. eurysto/e).
Anchoviella brevirostris (Giinther) 1868
Study Material. Two small soft specimens, 26 and 24 mm SL, from Lago Max-
imus, far up the Amazon Basin, Brazil, and six specimens that have been more or
less dry, 17-22 mm SL, from Teffé, also far up the Amazon Basin, which may be
of this species, all in MCZ. Another, 40 mm TL, from Lake Rogagua, Bolivia, also
may be of this species (part of CAS 9398). Because of the poor condition and small
size of the specimens and the inadequacy of the original description, identification is
uncertain.
Description. Proportional dimensions in per cent of standard length, based on
two of the larger and better specimens, 26 and 24 mm SL, respectively.
Body: depth 15.3, 16.1. Anal fin: length of base 19.2, 17.5.
Head: length 20.4, 24.4. Pectoral fin: length ?, 11.2.
Snout: length 3.8, 4.15.
Eyes diameter 7-75.7-3; Gill rakers: 13 or 144+19 (4 specimens).
Postorbital: distance 12.7, 12.5. Fin rays: dorsal 12 or 13; anal 16-19;
Maxillary: length 15.4, 12.5. pectoral 11 or 12 (4 specimens).
Mandible: length 15.4, 13.7.
Bopy slender, its depth 6.2, 6.5 in SL. Heap 4.1, 4.9. SNour projecting about
half of its length beyond mandible, 5.5, 6.3 in head. Eye 3.0, 3.2. Posrorsirat part
of head 1.75, 1.9 in head. Maxitrary nearly straight, rounded distally, extending
about an eye’s diameter beyond posterior rim of orbit but not quite to joint of man-
dible, 1.5, 1.75 in head. ManpiBiez 1.57, 1.6.
Dorsat origin about equidistant between base of caudal and tip of snout. ANAL
origin under base of last 2 or 3 rays of dorsal; anal base 5.2, 5.7 in SL. Pecrorat
failing to reach pelvic by an eye’s diameter, 2.0 in head.
Color. Very old specimens, in alcohol, faded, except for the silvery lateral band,
which is somewhat narrower than the eye.
Digest of Giinther’s Description, type 87 mm TL. Heap 4.5, its depth 5.33. Snour
much shorter than eye, projecting only slightly beyond mandible. Eye 3.5 in head.
Fishes of the Western North Atlantic 217
Maxiiiary dilated and rounded distally, not extending to joint of mandible. Dorsat
origin equidistant between base of caudal and tip of snout. Anat origin below last
rays of dorsal. Pecrorat ending “‘a great distance” from the pelvic. Fin rays: dorsal
15, anal 18. Scares 41. Sides with a well-defined silvery band, this band nowhere
wider than eye.
It is most unfortunate that the number of gill rakers was not stated.
Range and Habitat. lf the specimens examined have been correctly identified and
are the same species, the range extends from ‘“‘Caxoeira,!° Province of Bahia,” Brazil,
type locality, to Lago Maximus and Teffé, both far up the Amazon; and probably to
Rio Chapare and Lake Rogagua, Bolivia. This species, then, seems to live in fresh
water. That its distribution is as wide as indicated remains to be confirmed from the
study of more and better specimens.
Synonyms and References:
Engraulis brevirostris Giinther, Cat. Fish. Brit. Mus., 7, 1868: 392 (orig. descr.; type local. “Caxoeira, Province
of Bahia,” Brazil); Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 411 (refer it to dmp/ova,
herein regarded incorrect).
Amplova brevirostris Myers, Proc. Calif. Acad. Sci., 23, 1940: 440 (notes).
Anchoviella brevirostris Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 119 (descr., range).
Doubtful Reference:
Anchoviella carrikeri Fowler, Proc. Acad. nat. nat. Sci. Philad., 92, 1940: 73, fig. 28 (orig. descr.; type local.
mouth of Rio Chaparé, Cochabamba, Bolivia; type ANSP 68980; placed here with considerable doubt
because scales [31-33] and gill rakers on the lower limb [14 or 15] seem to be too few, and the pectoral
fin as shown in figure may be too long).
Negative References:
Anchoviella brevirostris Pearson, Indiana Univ. Stud., 64, 1925: 50 (Lake Rogagua, Bolivia; cf. paratype of
Stolephorus guianensis Eigenmann by Myers and found to be that species; see ref. under 4. guianensis) ;
Fowler, Proc. Acad. nat. Sci. Philad., 92, 1940: 94 (listed, after Pearson).
Anchoviella guianensis (Eigenmann) 1912
Figure 47
Study Material. A paratype, somm TL, from the Demerara River, British
Guiana, CM 2449; and a rather poorly preserved specimen, 62 mm, from Itacoatiara,
Amazonas, Brazil, CNHM 15480.
Distinctive Characters. This species differs from the specimens herein assigned to
A. brevirostris chiefly in the greater number of gill rakers.
Description. Proportional dimensions in per cent of standard length, and counts,
based on Study Material, 50 and 62 mm, respectively.
to. Dr. G.S.Myers has advised that “Caxoeira” and “Cachoeira’’ are alternative spellings in Brazilian Portu-
guese for “rapids,” and that both are pronounced alike. Cachoeira is the generally accepted spelling today.
—Y.H.O.
218 Memoir Sears Foundation for Marine Research
Body: depth 20, 17.4. Anal fin: length of base 16.8, 15.5.
Head: length 23.7, 24.2. Pectoral fin: length 15.2, 16.8.
Snout: length 4.5, 5.0.
Eye: diameter 7.5, 7.0. Scales: ca. 41, 38.
Postorbital: distance 10.7, 14.2. Gill rakers: 16+22, 17+24.
Maxillary: length 12.5, 13.3. Fin rays: dorsal 13, 14; anal 18, 19;
Mandible: length 14, 13.8. pectoral 12, 13.
Bopy compressed, its depth 5.0, 5.7 in SL. Heap 4.2, 4.1 in SL, its depth about
equal to its postorbital part and half of eye. Snour projecting nearly half of its length
Ficure 47. Anchoviella guianensis, 62mm TL, from Amazonas, Brazil, CNHM 15480. Drawn by Louella
E. Cable.
beyond mandible, 5.3, 4.9 in head. Eve 3.2, 3.5. Posrorsirat rather short, 2.2, 1.8
in head. MaxiLiary extending about 75°/, of diameter of eye beyond orbit, but not
to joint of mandible, 1.9, 1.8 in head. Manpisie 1.7, 1.75. CHreK broad, very
slightly longer than eye, its posterior angle about 60°.
Dorsat fin with longest rays reaching somewhat beyond tip of last ray if deflexed,
its origin equidistant between tip of snout and caudal base. Anat originating under,
or a little in advance of, base of last dorsal ray, its base 5.95, 6.5 in SL. Petvic inserted
equidistant between pectoral base and anal origin. Pecrorat failing to reach pelvic by
diameter of pupil, 1.55, 1.45 in head. AxitLary scaLe of pectoral narrow, reaching
well beyond midlength of pectoral, 2.4, 2.45 in head.
Color. In alcohol, pale (probably preserved originally in formalin). Side of head
silvery. Pale streak, probably silvery in life, on side. Back with dark punctulations
arranged in two more or less definite rows behind dorsal fin; dusky punctulations along
base of anal, and a median dark stripe behind anal; paratype with dusky punctulations
present also on dorsal, caudal, anal, and upper rays of pectoral.
Range and Habitat. Its range extends from Trinidad, B.W.I., and Venezuela to
northern Brazil and the upper Amazon Basin in Bolivia. In addition to the holotype
from Bartica Rocks, British Guiana, Eigenmann had paratypes from the Morawhanna
Fishes of the Western North Atlantic 219
and Demerara rivers, British Guiana. It is recorded for La Brea Beach, Trinidad,
and for Cafio de Guanoco, Venezuela, without comment by Fowler (see reference
below). Fresh water seems to be its chief habitat.
Synonyms and References:
Stolephorus guianensis Eigenmann, Mem. Carneg. Mus., 5, 1912: 447, pl. 62, fig. 5 (orig. descr.; type local.
Bartica Rocks, British Guiana; type in CM; paratypes from Morawhanna and Demerara rivers, Brit.
Guiana).
Anchoviella guianensis Fowler, Proc. Acad. nat. Sci. Philad., 83, 1931: 392, 406 (La Brea Beach, Trinidad, and
Cano de Guanoco, Venezuela); Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 122, fig. 52
(synon., descr., relation.); Schultz, Proc. U.S. nat. Mus., 99, 1949: 48 (refs.).
Anchovia brevirostris Pearson (not of Giinther), Indiana Univ. Stud., 11 (64), 1924: 50 (two immatures, 30
and 37 mm, IU 17354, now in CAS, rec. from Lake Rogagua, Bolivia; cf. paratype of Szolephorus
guianensis Eigenmann by Myers and found to be that species).
Amplova guianensis Myers, Proc. Calif. Acad. Sci., (4) 23, 1940: 440 (descr., exam. specimens from Mo-
rawhanna [paratype]; Lagoa Grande, lower Amazon, near Obidas, Brazil; and Lake ‘“Rogoagua,”
Bolivia; those from last mentioned place are the ones ident. as 4. 4revirostris by Pearson; see above).
Anchoviella alleni (Myers) 1940
Study Material. The holotype and 6 paratypes, 80-84 mm TL, from Peruvian
Amazon drainage system; the holotype and 4 paratypes in CAS, 2 paratypes in
USNM.
Distinctive Characters. This species is close to 4. guianensis, from which it differs in
having a somewhat smaller eye, a shorter anal base, a shorter and broader cheek, a
longer axillary scale on the pectoral, and in color.
Description. Proportional dimensions in per cent of standard length, and counts,
based on the type material noted above, 80-84 mm TL.
Body: depth 18.8—20. Anal fin: length of base 13.3-15.
Head: length 22.7-25. Pectoral fin: length 14.7—-16.5.
Snout: length 3.4-4.2.
Eye: diameter 5.5-6.25. Scales: ca. 44-46.
Postorbital: distance 11.3-12. Gill rakers: 16-18 + 23-26.
Maxillary: length 11.8—13. Fin rays: dorsal 14 or 15; anal 18 or 19;
Mandible: length 12.7—13.2. pectoral 12 or 13.
Vertebrae: 42 (1 specimen).
Bopy slender, moderately robust, its depth 5.0—5.3 in SL. Heap 4.0—4.4 in SL, its
depth only slightly less than its postorbital length and eye. SNour projecting about
half of its length beyond mandible, 5.5—6.0 in head. Eve 3.7—4.1 in head, 1.9—2.1 in
postorbital part of head. Posrorgirat 1.9—2.0 in head. Maxiztary slightly expanded
and rounded distally, extending as much as diameter of pupil beyond posterior rim of
orbit, failing to reach joint of mandible by about half of diameter of pupil, 1.7—2.0 in
220 Memoir Sears Foundation for Marine Research
head. ManpisLe 1.63—1.77. CHEEK very short and broad, slightly rounded poste-
riorly, scarcely as long as eye, and scarcely forming an angle.
Dorsat fin with longest rays reaching to or beyond tip of last ray if deflexed,
its origin generally a little nearer to caudal base than to tip of snout. Anat originating
immediately behind vertical from base of last ray of dorsal, its base 6.7—7.4 in SL.
Petyic reaching a little more than halfway to anal, inserted nearer to base of pectoral
than to origin of anal by diameter of pupil. Pecrorat moderately falcate, failing to
reach pelvic by diameter of pupil, 1.4—1.6 in head. Axittary scatz of pectoral reaching
opposite distal third of fin, 2.0—2.2 in head.
Color. In alcohol, pale, probably preserved originally in formalin. Lateral band
bright silvery, nearly as broad as eye, bounded above by a dark line. Tip of snout and
back with dusky punctulations, these sometimes arranged in two more or less definite
rows posterior to dorsal fin. Caudal with dusky markings and sometimes with a broad
dusky margin.
Range. Known only from the Peruvian Amazon country, from Lake Cashiboya,
and Gosulima Cocha, Upper Rio Morona, Peru, 2,000 miles from the sea.
Synonyms and References:
Amplova alleni Myers, Proc. Calif. Acad. Sci., (4) 23, 1940: 441 (orig. descr.; type local. Lake Cashiboya,
Peru; type CAS 6421; cf. Stolephorus guianensis Eigenmann and Engraulis brevirostris Giinther; para-
types from Lake Cashiboya, Rio Ucayale; Gosulima Cocha, Upper Rio Morona; and Rio Morona,
all in Peru); Eigenmann and Allen, Fishes West. S. Amer., Univ. Ky., 1942: 333 (add. specimens from
upper Amazon basin).
Anchoviella alleni Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 124, fig. 53 (descr. of type material;
cf. Stolephorus guianensis Eigenmann).
Anchoviella nattereri (Steindachner) 1879
Study Material. None.
Distinctive Characters. In general, 4. nattereri seems to be related to 4. blackburni
and A. lepidentostole, but it differs from each of them in the rather more numerous anal
rays, the more posterior origin of the anal in relation to the dorsal, and the much longer
pectoral fin, which overlaps the pelvic.
Description. Based on original account, translated and condensed from Stein-
dachner, of a specimen 50 mm long."
11. A description with figure of an anchovy from Iquitos, Peru (upper Amazon Basin), designated Engraulis iquitensis
by Shoji Nakashima (r4: 6), shows some of the characters of 4. mattereri. At least the general shape of the body,
the number of anal rays, and the long pectoral overlapping the base of the pelvic seem to agree. However, Naka-
shima did not describe the maxillary, and its outline is not clearly shown in the figure; the number of gill rakers
is not stated; and the description and figure do not agree in several respects. Therefore, it is impossible to place the
fish in any genus with a degree of certainty, or to determine the validity of the species. As the author designated
no type and failed to compare his fish with any known species, the name probably is without standing and no
further reference will be made to it.
Fishes of the Western North Atlantic 221
Body: depth 4.0. Scales: ca. 40.
Head: length 3.33. Fin rays: dorsal 12; anal 28 or 29.
Bopy with dorsal profile anterior to dorsal fin less convex than ventral profile.
Snour reaching well beyond mandible, 4.0 in head. Eye 3.6. Maxitiary nearly
square distally, not quite reaching to joint of mandible. Longest Gitt RakeRs about
as long as eye.
Dorsat origin slightly nearer to base of caudal than to tip of snout. Anat origin
about under middle of dorsal base. Pecrorat slightly longer than postorbital part of
head, reaching middle of pelvic. The silvery lateral band is definitely marked.
Range. Known only from the type from Belém (Para), Brazil.
Synonyms and References:
Engraulis nattereri Steindachner, Ichthyol. Beitr., 8, 1879: 56 (orig. descr.; type local. Pard, Brazil; cf. Ev-
graulis surinamensis Bleeker = E. edentulus Cuvier).
Anchoviella nattereri Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 133 (descr. after Steindachner;
relation.).
Negative Reference:
Anchovia nattereri Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 413 (descr. based on specimens
from Pard, Brazil, MCZ 18044; exam. by me and found to be dachovia clupeoides, differing from
Steindachner’s descr. of 4. zattereri in the longer pointed maxillary, in the apparently shorter anal
fin and its more posterior origin with respect to dorsal, and in the much shorter pectoral fin, not extending
beyond base of pelvic).
Anchoviella pallida (Starks) 1913
Figure 48
Study Material. Three specimens, 72, 80, and 102 mm TL, the first two being
the holotype and paratype of Anchoviella venezuelae Fowler, the third one, the holotype
of Anchovia pallida Starks, from Cafio de Guanoco, Venezuela, and Para, Brazil, re-
spectively. The specimens agree so well that they almost certainly are one species.
Distinctive Characters. In general shape this species resembles Cetengraulis edentu-
Jus; however, it differs from it in lacking a membrane connecting the gill covers across
the isthmus, and in having a longer and narrower cheek, a higher dorsal, and a longer
pectoral. From the other members of its genus it is well differentiated by the deep body,
long head, long pectoral, and numerous gill rakers.
Description. Proportional dimensions in per cent of standard length, and counts,
based on the study specimens, 72-102 mm TL.
Body: depth 25.6—28.5. Maxillary: length 19-20.
Head: length 28.5—29.4. Mandible: length 18.8—20.
Snout: length 4.5—4.8. Anal fin: length of base 27.7—28.5.
Eye: diameter 6.36—6.9. Pectoral fin: length 18.8—20.
Postorbital: distance 15.8—17.
222 Memoir Sears Foundation for Marine Research
Scales: ca. 36-38. Fin rays: dorsal 12-143 anal 24-26; pec-
Gill rakers: 28-34 + 36-45. toraliieror=r3:
Bopy strongly compressed, its depth 3.5-3.9 in SL. Heap 3.4-3.5, its depth
equal to its postorbital length and about half of eye. Snour projecting about 0.66 of
its length beyond mandible, 6.0—6.4 in head. Eye 4.2—-4.9. Posrorsirat part of head
1.7-1.85 in head. Maxittary somewhat expanded, rounded distally, extending about
1.5 times diameter of eye beyond posterior rim of orbit, not quite reaching joint of
SS
Ficure 48. Anchoviella pallida, type specimen, 102 mm TL, from Pard, Brazil, SU 22216. Drawn by Alice
C. Mullen.
mandible, 1.4-1.55 in head. ManpiBLe 1.43-1.55. CHEEK narrow, equal to length of
snout and eye in large examples, somewhat shorter in smaller ones, its posterior angle
about 35°. Git RAKERS Close-set, numerous, about as long as eye (possibly increasing
in number with age).
Dorsat fin with longest rays reaching beyond tip of last ray if deflexed, its origin
about equidistant between caudal base and tip of snout. ANaL originating about under
middle of dorsal base, 3.5—3.6 in SL. Petvic reaching about 0.66 of distance to anal,
inserted a little nearer to origin of anal than to base of pectoral. Pecrorat reaching
beyond base of pelvic by as much as diameter of pupil in largest specimen examined,
somewhat shorter in smaller examples, 1.4—1.85 in head. Ax1LLary scaLe of pectoral
broad at base, tapering abruptly, about a third to half as long as the fin, 3.3-3.6
in head.
Color. In alcohol, plain. No silvery lateral band evident. Side of head silvery.
Back with dusky punctulations. Caudal fin with a dusky margin in the largest specimen
examined (holotype), but not evident in the others.
Variation. The number of gill rakers in the three specimens examined shows a
rather wide variation, not attributable to age and size; the smallest specimen, according
to my counts, has 34+42 rakers (34 +36 according to Fowler’s original description
Fishes of the Western North Atlantic 223
of A. venezuelae), the largest 34+45, and the intermediate one 33+40 on one side
and 28 +36 on the other.
Range. Known from only three specimens, two from Cafio de Guanoco, Venezuela,
and the type from Para, Brazil.
Synonyms and References:
Anchovia pallida Starks, Stanf. Univ. Publ., Univ. Ser., 1913: 9, pl 1 (orig. descr.; type local. Pard, Brazil;
type SU 22216; cf. Engraulis endentulus Cuvier, E. vaillanti Steindachner and Cetengraulis juruensis
Boulenger).
Anchoviella venezuelae Fowler, Proc. Acad. nat. Sci. Philad., 83, 1931: 406, fig. 6 (orig. descr.; type local.
Cano de Guanoco, mouth of Rio San Juan, Venezuela; type ANSP 53322; cf. Axchovia parva Meek
and Hildebrand).
Anchoviella pallida Fowler, Proc. Acad. nat. Sci. Philad., 93, 1941: 126 (synon., descr., Ceard, Brazil); Fowler,
Arqu. Zool. estad. Sao Paulo, 3 (6), 1941: 134 (ref. Brazilian rec.); Hildebrand, Bull. Bingham oceanogr.
Coll., 8 (2), 1943: 134, fig. 59 (descr., 4. vewexue/ae in synon.); Schultz, Proc. U.S. nat. Mus., 99,
1949: 48 (synon., refs.).
Anchoviella elongata (Meek and Hildebrand) 1923
Figure 49
Study Material. Specimens 65-100 mm: the type material from Mindi, Canal
Zone, one specimen from Porto Bello, Panama, and 22 from Puerto Pilon, Panama.
Distinctive Characters. This species is readily recognized by the slender body,
the very large eye, and the moderately few anal rays and gill rakers. In the length and
shape of the maxillary it approaches the genus Anchoa rather closely.
Description. Proportional dimensions in per cent of standard length, aud counts,
based on at least 14 specimens in Szudy Material.
Body: depth 17.5-20. Anal fin: length of base 22—23.7.
Head: length 25.5—27. Pectoral fin: length 15.3-16.3.
Snout: length 4.5—5.1.
Eye: diameter 8.2—9.0. Scales: ca. 36-40.
Postorbital: distance 12.1-13.3. Gill rakers: 17 or 18+22—24.
Maxillary: length 19-20. Fin rays: dorsal 13 or 14; anal 22-24;
Mandible: length 17.8-18.8. pectoral 12 or 13.
Vertebrae: 39 (2 specimens).
Bopy slender, moderately compressed, its depth 5.0—-5.7 in SL. Heap 3.6—3.9
in SL, its depth equal to postorbital part of head and half of eye. Sour projecting about
half of its length beyond mandible, 5.3—5.8 in head. Eve large, 3.0—3.3 in head. Posr-
ORBITAL part of head 2.0-2.25 in head. Maxittary moderately blunt distally, reaching
nearly or quite to joint of mandible (approaching the genus Azchoa in these respects),
extending nearly an eye’s diameter beyond posterior rim of orbit, 1.3-1.45 in head.
ManpiBle 1.45—1.5. CHEEK about equal to length of eye, its posterior angle approxi-
mately 60°.
224 Memoir Sears Foundation for Marine Research
Dorsat fin moderately high anteriorly, its longest rays reaching beyond tip of
last ray if deflected, its origin about equidistant between caudal base and anterior
margin of eye. ANAL originating under or slightly behind middle of dorsal base, its base
4.2—4.5 in SL. Prtvic generally reaching more then halfway to anal, inserted rather
nearer to pectoral base than to anal origin. Pecrorat usually extending scarcely to
base of pelvic, 1.6—1.75 in head. AxiLLary scaLe of pectoral about half as long as
fin, 2.8—3.75 in head.
Color. In alcohol, pale (originally preserved in formalin). Silvery lateral band
poorly defined, narrower than eye. Upper surface of head and back with dusky punc-
tulations; dark spots at base of anal, and a median dark line posterior to anal.
Size. The largest specimen in the collection, about 100 mm (4 in.) TL, is the
largest individual reported.
Ficure 49. Anchoviella elongata, 101 mm TL, from Porto Bello, Panama, USNM 82093. Drawn by Louella
E. Cable.
Range and Habitat. Known only from the Atlantic coast of Panama, where it ap-
parently is not rare. Specimens have been collected from brackish water at Mindi
(near Gatun), and from Gatun Spillway, Canal Zone; and from Porto Bello and Puerto
Pilon, Panama.
Synonyms and References:
Anchovia elongata Meek and Hildebrand, Field Mus. Publ., Zool., 75 (1), 1923: 198, pl. 12, fig. 2 (orig.
descr.; type local. Mindi, Canal Zone; type USNM 81768; cf. 4. cubana); Breder, Zoologica, N. Y.,
4.(4), 1925: 141 (common at Gatun Spillway, Canal Zone).
Anchoviella elongata Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 49 (ref. to orig.
descr.); Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 126, fig. 54 (descr., generic relations.,
range).
Anchoviella lepidentostole (Fowler) 1911
Figure 50
Study Material. The type, about 96 mm (caudal fin damaged), 84 mm SL, and a
paratype 76 mm long, 63 mm SL, from “Surinam.”
Fishes of the Western North Atlantic 225
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens, 84 and 63 mm SL, respectively.
Body: depth 25, 22.2. Anal fin: length of base 23.8, 23.8.
Head: length 22.7, 23.8. Pectoral fin: length 16.3, 15.8.
Snout: length 3.2, 3.7.
Eye: diameter 6.9, 7.1.
Postorbital: distance 10.7, II.
Maxillary: length 16.4, 15.
Mandible: length 16.4, 15.8.
Scales: ca. 38.
Gill rakers: 17 +19 right side, 18 +20
left side; 17 +19 both sides.
fin rays: dorsal 14; 155 anal 26, 29:
pectoral 12 (?), 14.
Bopy rather robust, its depth 4.0, 4.5 in SL. Heap 4.4, 4.2 in SL, its
depth nearly equal to its length without snout. SNour extending about half of its
Ficure 50. Anchoviella lepidentostole, type specimen, 96mm TL, from “Surinam,” ANSP 1346, modified
after Fowler (1911: 214, fig. 3). Drawn by Alice C. Mullen.
length beyond mandible, 7.0, 5.5 in head. Eve 3.3, 3.3. Posrorsirat part of head
2.1, 2.15 in head. Maxittary not quite square distally, scarcely reaching joint of
mandible, 1.4, 1.6 in head. ManpiBLE 1.35, 1.5. CHEEK as long as eye, the posterior
angle about 50°.
Dorsat fin with longest rays failing to reach tip of last ray if deflexed, its origin
a little nearer to caudal base than to tip of snout. ANAL originating under middle of
dorsal base, its base 4.2, 4.2 in SL. Petvic reaching scarcely halfway to anal, inserted
a little nearer to anal origin than to pectoral base. Pecrorat failing to reach pelvic by
diameter of pupil, 1.45, 1.5 in head. AxiLtary scate of pectoral failing to reach tip
of fin by half of diameter of pupil, 2.0, 2.15 in head.
Color. In alcohol, brown. Lower parts silvery. Silvery lateral band strikingly
prominent, much wider than eye.
Range. Known only from “Surinam.”
Ts
226 Memoir Sears Foundation for Marine Research
Synonyms and References:
Anchovia lepidentostole Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 214, fig. 3 (orig. descr.; type local.
“Surinam”; type ANSP 1346; cf. Exgraulis brevirostris Giinther and E. januarius Steindachner).
Anchoviella lepidentostole jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 49 (ref. to
orig. descr.); Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 127, fig. 55 (descr. of type material).
Anchoviella blackburni Hildebrand 1943
Figure 51
Study Material. Type material, 11 specimens, about 33-38 mm TL (caudal fins
damaged), all from the Gulf of Venezuela. The specimens probably are not full-
Ficure 51. Anchoviella blackburni, type specimen 35 mm TL, from Gulf of Venezuela, USNM 119793.
Drawn by Louella E. Cable.
grown, but the maxillary, which is shorter and blunter in very young than in adults,
probably has acquired the approximate length and shape of a mature fish. At least it
is shorter and much more rounded than in specimens of Auchoa parva and A. mitchilli
of similar size.
Distinctive Characters. The long anal fin, the small number of gill rakers, the short
head, and the very short, blunt snout distinguish this species from related forms.
Description. Proportional dimensions in per cent of standard length, and counts,
based on the 11 type specimens, about 33-38 mm TL.
Body: depth 18.2-20.8. Anal fin: length of base 23.5-26.5.
Head: length 21.7-25. Pectoral fin: length 13.8-16.6.
Snout: length 3.6-3.8.
Eye: diameter 5.0—7.5. Scales: ca, 38-41.
Postorbital:; distance 11-14.3. Gill rakers: 10-12 + 15-17.
Maxillary: length 12-15. Fin rays: dorsal 13-15; anal 25-27;
Mandible: length 12.3-14.8. pectoral 13 or 14.
Vertebrae: 43 (1 specimen).
Fishes of the Western North Atlantic 22
Bopy quite strongly compressed, its depth 4.8—5.5 in SL. Heap 4.0—4.6 in SL, its
depth equal to postorbital part of head and about half of eye. Snour very short,
projecting little beyond mandible, about 5.0—7.0 in head. Eve 3.5—4.5. Posrorsirat
part of head 1.6—-1.9 in head. Maxitrary slightly expanded and rounded distally,
extending about half of diameter of eye beyond posterior rim of orbit, not quite to joint
of mandible, 1.6—2.0 in head. Cueexk only a little longer than eye, the posterior angle
about 60°.
Dorsat fin with the longest rays failing to reach tip of last ray if deflexed, its origin
about equidistant between caudal base and middle of eye. Anat originating about
under origin of second third of dorsal base, its base 3.75-4.25 in SL. Prtvic reaching
more than halfway to anal. Pecrorat failing to reach base of ventral by somewhat less
than diameter of eye, 1.25-1.6 in head.
Color. In alcohol, pale (originally preserved in formalin), with only a suggestion
of a narrow silvery lateral band. A row of dark spots along base of anal, continued as a
median dark line posterior to anal; a similar row along base of dorsal to upper lobe of
caudal.
Range. Known only from the Gulf of Venezuela, where it was taken in Estauques
Bay and Jacque Point.
References:
Anchoviella blackburni Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 129, fig. 57 (orig. descr.;
type local. Jacque Point, Gulf of Venezuela; type USNM 119793; cf. Anchoa parva [Meek and Hilde-
brand] and Anchoviella lepidentostole [Fowler]); Schultz, Proc. U.S. nat. Mus., 99, 1949: 48 (ref.,
counts).
Anchoviella (Amplova) jamesi (Jordan and Seale) 1926
Study Material. Two paratypes, about 47 and 37 mm TL, 36 and 32 mm SL,
from fresh water as listed under Range; MCZ 17783 and 17784, respectively.
Distinctive Characters. This species has an extremely small mouth for an anchovy
and is thus related to 4. ba/boae Jordan and Seale of the Pacific coast of Panama, which
was made the type of the genus 4mplova Jordan and Seale, herein regarded as a sub-
genus. But it differs from 4. ba/boae in having fewer anal rays, fewer gill rakers, a more
posteriorly placed dorsal fin, and apparently a more slender body. 4. balboae has
24-27 anal rays and 19-21 +29-365 gill rakers; its dorsal origin is about equidistant
between the base of caudal and middle of eye; and the depth in specimens 75—100mm
DLs 3-5=4.1 m SE.
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens 36 and 32 mm SL, respectively.
Body: depth 22.2, 23.8. Eye: diameter 5.8, 6.6.
Head: length 22.2, 22.7. Postorbital: distance ?, 12.8.
Snout: length 3.04, 3.1. Maxillary: length 11.6, 11.9.
228 Memoir Sears Foundation for Marine Research
Anal fin: length of base 21, 18.7. Gill rakers: 13 +21, 12 +20.
Pectoral fin: length 16.6, 18.7. Fin rays: dorsal 13, 123; anal 21, 19;
pectoral 11, IT.
Scales: ca. 40. Vertebrae: 40, ?.
Bopy moderately compressed, its depth 4.5, 5.3 in SL. Heap 4.5, 4.4 in SL, its
depth equal to postorbital length and about half of eye. Snour very short, projecting
about a third of its length beyond mandible, 7.2, 7.3 in head. Eye 3.8, 3.5. Post-
ORBITAL part of head 1.85 in head. Maxizrary distally rounded, scarcely reaching
beyond posterior rim of orbit, and not to articulation of mandible, 2.0, 1.9 in head.
Cueek very short and broad, scarcely forming a triangle, much shorter than eye.
Dorsat fin with the longest rays reaching far beyond tip of last ray if deflexed,
its origin nearer to base of caudal than to eye. Anat with a prominent lobe anteriorly,
its origin slightly in advance of middle of dorsal base, its base 4.8, 5.3 in SL. Petvic
reaching more than halfway to anal, inserted about equidistant between base of pec-
toral and origin of anal. Pecrorat nearly reaching base of pelvic, 1.2, 1.25 in head.
AXILLARY SCALE of pectoral missing in specimens examined.
Color. Old specimens, in alcohol, faded, pale. Silvery lateral band rather diffuse,
narrower than eye.
Range. Known from the type material from fresh water, the holotype being from
Jutahy River, the two paratypes from Lago Alexo, Brazil. Two small specimens also
are reported from the Rio Ucayali, in Peru.
Synonyms and References:
Amplova jamesi Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 410 (orig. descr.; type local. Jutahy
R., Brazil; type MCZ 18014; two paratypes listed from Lago Alexo, Brazil); Myers, Proc. Calif.
Acad. Sci., (4) 23, 1940: 441 (descr. after Jordan and Seale; Rio Ucayali, Peru).
Anchoviella jamesi Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 137, fig. 61 (descr., relation. with
Amplova balboae Jordan and Seale).
Genus Prerengraulis Giinther 1868
Pterengraulis Giinther, Cat. Fish. Brit. Mus., 7, 1868: 384, 398; type species by monotypy, C/upea atherinoides
Linnaeus 1758.
Characters. Bopy rather deep, strongly compressed. GiLL RakeERs short and broad,
especially in large examples, only about 11 on upper limb and about 14 on lower
limb. Dorsat origin about equidistant between margin of opercle and base of caudal.
Pexvic inserted about midway between pectoral base and dorsal origin.
Size. A large size is attained, specimens 250 mm long being included in the
collections studied.
Remarks. The anal fin is long, with about 30-33 rays, its origin being under,
or more usually in advance of, the dorsal origin. The teeth are small and of nearly
Fishes of the Western North Atlantic 229
uniform size. The maxillary extends to, or nearly to, the joint of the mandible and is
rounded distally.
Range. Venezuela to northern Brazil.
Species. Only one, P. atherinoides.
Prerengraulis atherinoides (Linnaeus) 1766
Hareng gras Sardinha
Figure 52
Study Material. A total of 20 specimens, 66-250 mm TL, from Cafio de Guanoco
and Rio Apure, San Fernando de Apure, Venezuela; and from the “Amazon River,”
Para, Fortaleza, Cametdé, and Arary, Brazil.
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens, 66-250 mm TL.
Ficure 52. Prerengraulis atherinoides, 245 mm TL, from “Amazon River,” USNM 72551. Drawn by Louella
E. Cable.
Body: depth 22-26. Anal fin: length of base 30—-34.5.
Head: length 22.6—28.5. Pectoral fin: length 20.4—23.7.
Snout: length 2.4—3.1.
Eye: diameter 4.0—5.1. Scales: 42-47.
Postorbital: distance 14.3-16.7. Gill rakers: 10-12 + 12-15.
Maxillary: length 16.5—19.2. Fin rays: dorsal 12-14; anal 29-35
Mandible: length 15.4—-18. (usually 31 or 32); pectoral 13-15.
Vertebrae: 43 and 45 (2 specimens).
Bopy strongly compressed, its depth 3.5-4.4 in SL. Heap 3.8—4.5 in SL, its
depth usually equal to its length posterior to middle of eye. SNour blunt, 8.0—10 in
head. Eye 4.0-6.0. Posrorsitat part of head moderately long, 1.4-1.55 in head.
230 Memor Sears Foundation for Marine Research
MaxiLiary rounded distally, often failing to reach joint of mandible, 1.3-1.5 in head.
MAanpiIBLE 1I.3—1.5. CHEEK increasing in proportionate length with growth, much
longer than eye and snout in large examples, its posterior angle about 40°.
Dorsat fin with the longest rays reaching beyond tip of last ray if deflexed, its
origin about equidistant between caudal base and margin of opercle. ANaL long, its
origin under or more usually somewhat in advance of dorsal origin, its base 2.9-3.3
in SL. Petvic reaching only about a third of distance to anal, usually inserted a little
nearer to base of pectoral than to anal origin. Pecrorat reaching to or beyond middle
of pelvic, 1.0-1.3 in head. AxitLary scaLe of pectoral reaching to or somewhat beyond
midlength of fin, 1.8—-2.1 in head.
Color. In alcohol, pale brownish above. Sides of head and lower parts of body
silvery. Silvery lateral band broad anteriorly, narrower on caudal portion of body, be-
coming diffused with the silvery shade on lower part of body anteriorly in large speci-
mens, apparently persisting posteriorly.
Size. Examples 300 mm TL have been reported.
Commercial Importance. This fish has been reported as very common in the market
at Belém (Pard), Brazil.
Range. Its range extends from Venezuela to northern Brazil, possibly as far south
as Rio de Janeiro. It ascends streams.
Synonyms and References:
Clupea atherinoides Linnaeus, Syst. Nat., ed. 12, I, 1766: 535 (orig. descr.; type local. Surinam; type lost).
Engraulis atherinoides Cuvier and Valenciennes, Hist. Nat. Poiss., 21, 1848: 31 (descr., relations., Surinam);
Giinther, Cat. Fish. Brit. Mus., 7, 1868: 398 (descr., Surinam, British Guiana, and Pard, Brazil);
Steindachner, Ichthyol. Beitr., 8, 1879: 59 (descr., Surinam; British Guiana; Pard, Cametd, Gurupd,
Rio de Janeiro, and Capim R., Brazil).
Pterengraulis atherinoides Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 450 (descr., range);
Schreiner and Miranda-Ribeiro, Arch. Mus. nac., Rio de J., 12, 1903: 94 (range, Guianas to Brazil);
Starks, Stanf. Univ. Publ., Univ. Ser., 1913: 10 (common in market at Pard, Brazil); Jordan and Seale,
Bull. Mus. comp. Zool. Harv., 67, 1926: 386 (descr., Pard, Cametd, Gurupd, and Arary, Brazil); Fowler,
Proc. Acad. nat. Sci. Philad., 83, 1931: 407 (Cafio de Guanoco, Venezuela); Puyo, Bull. Soc. Hist. nat.
Toulouse, 70, 1936: 170, fig. 36 (descr., habitat in French Guiana); Fowler, Proc. Acad. nat. Sci.
Philad., 93, 1941: 126, fig. 3 (Rio Jaguaribe, Ceard, Brazil; refs.); Arqu. Zool. estad. Sao Paulo, 3 (6),
1941: 135 (refs., Brazil); Campos, Arqu. Zool. estad. Sao Paulo, 3 (7) 1941: 200 (synon., descr., range) ;
Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 139, fig. 63 (descr., synon., range); Puyo,
Faune Emp. Frang., 12, Poiss. Guyane Frang., 1949: 158, fig. 81 (descr., habitat, French Guiana);
Schultz, Proc. U.S. nat. Mus., 99, 1949: 48 (San Fernando de Apure, Venezuela).
Clupea pilchardus de Magalhaes (not of Linnaeus), Monogr. Brazil, Peixes Fluviaes, 1931: 194, fig. 104 (descr.,
local name, length, habitat).
Doubtful Reference:
Engraulis atherinoides Knerr, “Novara,” Zool., Fische (1), 1865: 332 (descr. insufficient for ident.; Rio de
Janeiro, Brazil).
Genus Hildebrandichthys Schultz 1949
Hildebrandichthys Schultz, Proc. U.S. nat. Mus., 99, 1949: 49; type species by original designation, Hi/de-
brandichthys setiger Schultz.
Fishes of the Western North Atlantic 2a
Characters. Bopvy slender, rather strongly compressed, its depth about equal to
postorbital part of head. Heap long and low. Eye entirely in anterior half of head.
Maxi.tary rounded posteriorly, not quite reaching joint of mandible. Tgrru all small.
Git. RakerRs slender, those at angle of first arch about as long as eye, about 33 on lower
limb. Dorsat fin far back, its origin about twice the distance from tip of snout as from
caudal base. Anat fin beginning well in advance of origin of dorsal, about 25 rays.
Petvic inserted about equidistant between base of pectoral and anal origin.
Remarks. The numerous long gill rakers and the posterior position of the dorsal
fin are the chief distinguishing characters of this genus. Hi/debrandichthys resembles
Anchovia in the numerous long, slender gill rakers, but it differs in the very rear-
ward position of the dorsal fin. In the position of the dorsal it tends to agree more
or less with Prerengraulis, with which it differs in the numerous, long slender gill
rakers.
This genus is founded on one species, which in turn is based on only a single
small (juvenile) specimen. Though it is quite distinctive, it offers some difficulties be-
cause the adult generic characters apparently are not fully developed. The maxillary, for
example, is short and rounded, which, so far as known, is true of all young Engraulidae;
it may or may not become produced and pointed in the adult, as shown in other generic
descriptions.
Range. Known only from the Maracaibo Basin, Venezuela.
Hildebrandichthys setiger Schultz 1949
Figure 53
Study Material. A single small specimen, about 37 mm TL (caudal damaged),
31 mm SL, from the mouth of Rio Cafio de Sagua, Lake Maracaibo, Venezuela. This
specimen retains some of the juvenile color markings, and the maxillary may not have
acquired its adult length and shape.
Distinctive Characters. The numerous gill rakers, which may increase in number
with age, in general agree with the rakers in Anchovia clupeoides, the posterior position
of the dorsal fin calls to mind Pterengraulis atherinoides. See also Remarks under genus.
Description. Proportional dimensions in per cent of standard length, and counts,
based on young specimen, 31 mm SL.
Body: depth 19.2. Anal fin: length of base 22.6.
Head: length 32. Pectoral fin: length 12.
Snout: length 5.1.
Eye: diameter 8.7. Scales: mostly lost, ca. 38 pockets.
Postorbital: distance 17. Gill rakers: ca. 23 + 33.
Maxillary: length 17. Fin rays: dorsal 123; anal 25; pectoral 13.
Mandible: length 16.
21212 Memor Sears Foundation for Marine Research
Bopy slender, rather strongly compressed, its depth 5.2 in SL. Heap 3.1, its
depth about equal to its postorbital length. Snour projecting about 67°/, of its length
beyond tip of mandible, 6.25 in head. Eye small but longer than snout, 3.7 in head.
Posrorsirat length 5.9. Maxitiary rounded posteriorly, not quite reaching joint of
mandible, 1.9 in head. Manprsie 2.0. Cueexk a little longer than eye, its posterior
angle about 60°. GiLt RakeRs slender, those at angle of first arch about as long as
eye (too close-set for accurate counting), about 23 on the upper limb and about 33
on lower limb.
Dorsat fin small, damaged, apparently with a convex margin, the anterior rays
not reaching beyond tip of last ray if deflexed, the fin placed very far back, its
VEER
Re
Ficure 53. Hildebrandichthys setiger, type specimen, 37 mm TL, from Maracaibo Basin, Venezuela, USNM
121779. Drawn by A. M. Awl.
origin about twice as far from tip of snout as from caudal base. ANat originating about
an eye’s diameter in advance of dorsal origin and about equidistant between joint of
mandible and caudal base, its base 4.4 in SL. Petvic fin noticeably more than halfway
to origin of anal, inserted about equidistant between pectoral base and anal origin,
3-33 in head. Pecrorat fin failing to reach base of pelvic by a little less than diameter
of eye, 8.4 in SL, 2.6 in head. AxiLtary scaLe of pectoral not evident, lost or un-
developed.
Color. This specimen retains some of the juvenile color markings. In alcohol,
rather pale. Side of head and abdomen silvery. A rather faint narrow silvery lateral
band present. Dusky dots present on upper surface of head and snout, along base of
dorsal, and mostly in two rows behind the dorsal; a series of rather large black dots
along base of anal, continued as a single median row behind anal fin (presumably
juvenile markings); dusky dots generally distributed over the caudal fin; other fins
pale or at most with suggestions of dusky markings.
Range. Known only from the mouth of the Rio Cafio de Sagua, Lake Maracaibo,
Venezuela.
Reference:
Hildebrandichthys setiger Schultz, Proc. U.S. nat. Mus., 99, 1949: 49, fig. 7 (orig. descr.; type local. Cafio
de Sagua, Sinamaica, Venezuela; type USNM 121779).
Fishes of the Western North Atlantic 233
Genus Lycengraulis Giinther 1868
Lycengraulis Ginther, Cat. Fish. Brit. Mus., 7, 1868: 385, 399; type species Engraudis grossidens Cuvier 1828,
designated by Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 451.
Characters. Bovy slender, compressed, the depth generally less than 27°/, of SL.
Mourn very large. Maxittary extending far beyond eye, generally to or beyond joint
of mandible, rounded or pointed distally. Manpiste curved upward near tip. GILL
RAKERS variable in length, about 10-20+12-25 on first arch. ANaL originating
definitely posterior to dorsal origin, with about 24-28 rays. Pstvic inserted about
midway between base of pectoral and origin of anal. Young with a silvery lateral band
that disappears with age.
Size. About 250 mm is the maximum size attained in this genus.
Remarks. The genus is distinguished by the prominent teeth in the jaws: un-
equal in size in older specimens but rather even in young; larger in the lower than in
the upper jaw, some of them often canine-like; variable in number among the species,
about 16-25 enlarged teeth in lower jaw and 40-85 in upper jaw.
Species. Eight species have been recognized, one from the Pacific ranging from
Panama Bay to Peru, and seven from the Atlantic ranging from the Gulf of Venezuela
to northern Argentina. Five of the Atlantic species come within the scope of the present
work.
Key to Species of the Western North Atlantic
1a. Gill rakers long and slender, those at angle equal to or longer than pupil.
2a. Gill rakers 13-20+ 17-25 on first arch; cheek scarcely longer than snout and
eye in adults.
3a. Depth of body 23-24.5°/, of SL; maxillary reaching margin of opercle;
pelvic fin usually inserted nearer to origin of anal than to pectoral base,
occasionally equidistant between these; vertebrae usually 43 or 44 (oc-
casionally 42 or 45). grossidens (Cuvier) 1829, p. 234.
3b. Depth of body 21-23°/, of SL in specimens upward of about 100 mm
TL; maxillary not reaching margin of opercle; pelvic fin inserted nearer
to pectoral base than to anal origin; vertebrae usually 41 (occasionally 42).
limnichthys Schultz 1949, p- 237:
2b. Gill rakers g—13+12-15 on first arch; cheek much longer than snout and
eye in adults. batesii (Giinther) 1868, p. 240.
1b. Gill rakers short and broad, the longest scarcely exceeding half of length of pupil.
4a. Maxillary sharply pointed distally, extending well beyond joint of mandible;
origin of dorsal about equidistant between caudal base and middle of eye;
gill rakers 11 or 12+16 or 17. abbotti (Fowler) 1915, p. 242.
4b. Maxillary rounded distally, scarcely reaching joint of mandible; origin of
dorsal much nearer to base of caudal than to posterior margin of eye; gill
rakers 9-II1 + 12—I4. barbouri Hildebrand 1943, p. 243.
234 Memoir Sears Foundation for Marine Research
Table VIII. Comparison of L. grossidens with L. olidus. Proportional Dimensions
in Per Cent of Standard Length.
L. grossidens L. olidus L. grossidens L. olidus
40 specimens 44 specimens 40 specimens 44 specimens
& others (40— (97-240 mm & others (40- (97-240 mm
220mm TL) TL) from 2zomm TL) TL) from
from northern southern Brazil, from northern southern Brazil,
Brazil and Paraguay, Brazil and Paraguay,
Venezuela Uruguay, and Venezuela Uruguay, and
Argentina Argentina
Body: depth 21—24.3* 19.2—22.7 Scales: ca. 40-44 Ca. 40-45
Head: length 22.6-26.6 22.2-25 Gill rakers: 13-17 +17-21 15-19 + 19-23
Szout: length 3.3-4.3 2.7-3-4 (most fre- (most fre-
Eye: diameter 4.8-7.0 4.3-5-7 quently 14+19 quently 16 +21)
Postorbital: or 20)
distance 13-15 1315.5 Fin rays:
Maxillary: length 18.2-21.5 16.5—19.5 dorsal 14 or 15 I4 or 1S
Mandible: length 15.2—-18.2 15.5-17.5 anal 24-28 26-28
Anal fin: \ength pectoral 14 OF 15 14 or 15
of base 23.7-26.5 22-25.5 Vertebrae: 43 or 44 46-48
Pectoral fin: length 16.6—20.0 15.8-17.8 (4 specimens) (3 specimens)
* 23-24.3 im specimens 100 mm upward.
Lycengraulis grossidens (Cuvier) 1829
Sardinha Prata (Brazil)
Figure 54
Study Material. A total of 40 specimens, 40-220 mm TL, and many others as
well [presumably from Rio de Janeiro, Brazil and northward, as implied by Dr. Hil-
debrand under Relationship and Variation—y.u.o.]. Also 44 specimens, 97-240 mm
TL, from southern Brazil, Paraguay, Uruguay, and Argentina, representing L. olidus
(Giinther), used as basis for measurements and counts\in Tables vii—x.
Distinctive Characters. See Relationship and Variation.
Description. Proportional dimensions and counts, based on 40 specimens from
Rio de Janeiro, Brazil, and northward, 40-220 mm TL, in Table vim.
Bopy strongly compressed, increasing in depth with age, its depth 3.85—4.4 in
SL in specimens 100 mm or more. Heap with depth nearly equal to its length with-
out snout, 4.1-4.7 in SL. Snour projecting somewhat less than half of its length beyond
mandible, 5.6—6.9 in head. Eye 4.2-4.9. Posrorsirat part of head 1.6—-1.7 in head.
Maxittary little expanded, pointed distally, extending beyond joint of mandible, often
nearly to margin of opercle, 1.15—1.25 in head. ManpiBLe 1.33—-1.6. CHEEK some-
what longer than eye and snout in large examples, shorter than snout and eye in
specimens less than 100 mm, its posterior angle about 35°. Git RaxkeErs slender, those
at angle fully as long as pupil.
Dorsat fin with the longest rays usually reaching about to tip of last ray if de-
Fishes of the Western North Atlantic 236
Table IX. Frequency Distribution of Gill Rakers in Lycengrau/is.*
7——_ Number of Gill Rakers on Upper Limb of First Arch ————~
Specie 9 10 II 12 13 14 15 16 17 18 19 20
LOT OSSUCE IS ar ehaleserercTotetetehe — — — — 4 15 6 7 I = ee #4,
HU GO CEH OD BORIC — —_— — — — — 3 15 8 4 I 3
GERHIS ge o00g05000668- 3 2 4 5 2 = S| =| =|] =| =| =|
ALOT Sas CoD SODS OOS Se —- — I I -- —_— = = = ay ee a
DALOOUTL ace otic eltveics: I 3 4 — = = =: = —_ eS = =
* When the number of counts exceeds the number of specimens reported in the text, the gill rakers were counted
on both anterior arches of at least some of the fish.
Table X. Frequency Distribution of Gill Rakers in Lycengraulis. *
: Number of Gill Rakers on Lower Limb of JEN a) 9
Species
12 13 14 15 16 17 18 19 20 21 AD 23 24 2G
grossidens . — — — = == I 9 13 15 8 == = a =a
olidus .... = = —_— — — ae 2 5 6 20 7 5 2 I
batesii.... — 3 Wf a
abbotti ... — — a — I — = — — = — — —
barbouri .. I 6 I — — —_ = = — = — re = =
* When the number of counts exceeds the number of specimens reported in the text, the gill rakers were counted
on both anterior arches of at least some of the fish.
flexed, its origin rather variable, most frequently nearer to base of caudal than to poste-
rior margin of eye. ANAL with base 3.75—4.2 in SL, its origin generally under middle
of dorsal base. Petvic small, extending much less than halfway to anal, usually in-
serted nearer to anal origin than to pectoral base. Pecrorat often reaching nearly to
base of pelvic, 1.15—1.35 in head. AxiLLary scaLe of pectoral about 0.75 of length of
fin, 1.$—1.9 in head.
Color. In alcohol, grayish above. Sides of head and lower 67 °/, of body uniform
silvery in adults. Young under 90 mm long with a distinct silvery lateral band nar-
rower than eye, much broader than eye and less distinctly outlined ventrally in speci-
mens 120 mm long, intergrading completely with the silvery color of lower parts of
body in somewhat larger individuals. Margin of caudal blackish in adults.
Size. Examples having a maximum total length of 280 mm (11.2 in.) have been
reported, but the largest in the collections studied is only 220 mm TL.
Relationship and Variation. L. olidus (Giinther) is a very near relative of L. grossi-
dens, the two probably intergrading in southern Brazil. If specimens from Venezuela or
northern Brazil are compared with examples from extreme southern Brazil, Uruguay,
and Argentina, it is at once evident that those from the more southern localities have a
more slender and less strongly compressed body, a shorter and less pointed maxillary,
smaller teeth in the upper jaw but still much larger than those of other American genera,
a greater average number of gill rakers, and more numerous vertebrae. See Table vir
for comparison of L. grossidens with L. olidus.
L. poeyi of the Pacific coast of Panama is also a near relative of the Atlantic species,
2360 Memor Sears Foundation for Marine Research
but it differs in having: a shorter and blunter snout that projects little beyond the
mandible; smaller teeth of more uniform size in the lower jaw; a longer pectoral
that reaches to, or a little beyond, the base of the pelvic and has 16 or 17 rays
(1 or 2 more than in grossidens and olidus); and apparently a dark vertebral band, which
is missing in the L. grossidens and L. olidus examined. In depth of body and in length
and shape of the maxillary it agrees with L. o/idus rather than with L. grossidens.
Commercial Importance. As some of the specimens in the collections studied were
purchased in markets, this species must be of some commercial value.
Ficure 54. Lycengraulis grossidens, 214mm TL, from Porto Inhauna, Brazil, USNM 100835. Drawn by
Louella E. Cable.
Range and Habitat. The range of L. grossidens extends from the Gulf of Venezuela
south about to Rio de Janeiro, Brazil, and of L. o/idus from southern Brazil to northern
Argentina; both species ascend freshwater streams. The rather numerous records of
L. grossidens from Uruguay and Argentina are apparently referable to L. o/idus.
Synonyms and References:
Engraulis grossidens Cuvier, in Spix and Agassiz, Pisc. Brazil. 1829: 50, pl. 24, fig. 1 E (orig. descr.; type
local. Rio de Janeiro, Brazil); Giinther, Cat. Fish. Brit. Mus., 7, 1868: 399 (descr.,synon., British
Guiana).
Engraulis janeiro Agassiz, in Spix and Agassiz, Pisc. Brazil., 1829: pl. 24, fig. 1 (type local. Rio de Janeiro,
Brazil; name and fig. only).
Engraulis dentex Cuvier and Valenciennes, Hist. Nat. Poiss., 27, 1848: 28 (orig. descr.; type local. Rio de Janeiro,
Brazil); Kner, Reise “Novara,” Zool., Fische, 1, 1865: 332 (descr., Rio de Janeiro); Steindachner,
Ichthyol. Notiz., 6, 1867: 35 (descr., Rio de Janeiro and La Plata; the latter undoubtedly referable
to L. olidus).
Lycengraulis grossidens Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 451 (descr., range, synon.);
Gilbert, Proc. Wash. Acad. Sci., 2, 1900: 163 (descr., Pernambuco, Brazil); Starks, Stanf. Univ. Publ.,
Univ. Ser., 1913: 11 (descr., Natal, Brazil); Jordan and Gilbert, Bull. Mus. comp. Zool. Harv., 67,
1926: 383 (descr., Brazil and Uruguay; those from south of Rio de Janeiro, Brazil, probably L. o/idus;
those from Rio Poty, Brazil, are L. darbouri); von Ihering, Rev. Indust. Anim., Anno 1 (3), 1930: 233
(size, diagn.); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 51 (range, synon.);
Fowler, Arqu. Zool. estad. Sao Paulo, 3 (6), 1941: 135 (refs., Brazil); Hildebrand, Bull. Bingham oceanogr.
Fishes of the Western North Atlantic 237
Coll., 8 (2), 1943: 143, fig. 64 (descr., range, synon.); Schultz, Proc. U.S. nat. Mus., 99, 1949:
54 (synon., refs.).
Negative References:
Lycengraulis grossidens Berg, An. Mus. nac., Buenos Aires, 4, 1895: 21 (fairly abund. in markets, Buenos Aires
and Montevideo; L. o/idus); Devincenzi, An. Mus. nac., Montevideo, 2 (5), 1924: 189 (L. olidus);
Thompson, Proc. U.S. nat. Mus., 50, 1916: 405 (numerous from Montevideo, two from Buenos Aires;
in part L. ofidus).
Lycengraulis limnichthys Schultz 1949
Figure 55
Study Material. Type material of many specimens, 24-150 mm TL, 20-116 mm
SL, from Lake Maracaibo, Venezuela, and the lower parts of its tributary streams.
Distinctive Characters. This species is related to L. grossidens and L. olidus, the
latter being extralimital from southern Brazil to Argentina. L. limnichthys differs from
grossidens in having a more slender and apparently less strongly compressed body,
specimens of nearly equal size compared. In /imnichthys the maxillary is a little shorter,
scarcely reaching beyond the joint of the mandible and not to the margin of the opercle,
whereas in adults of grossidens it is longer, reaching well beyond the joint of the mandible
and often to the margin of the opercle. In /imnichthys the pelvic fins are inserted nearer
to the base of the upper pectoral ray than to the anal origin, whereas in grossidens they
are equidistant between these two points, or more usually nearer to the anal origin. And
limnichthys differs from both grossidens and ofidus in having fewer vertebrae.
Description. Propertional dimensions in per cent of standard length, and counts,
based on at least 11 study specimens.
Body: depth 21-23, in specimens 90 mm Pectoral fin: length 16-19.
TL and upward.
Head: length 23-25. Scales: ca. 35-45.
Snout: length 3.0-4.1 5. Gill rakers: 15-18 + 20-23 (rarely 24
Eye: diameter 5.4—6.5. or 25).
Postorbital: distance 12.5—-15. Fin rays: dorsal 13-16; anal 23-25
Maxillary: length 17.2-19.2. (rarely 26 or 27); pectoral 14 or 15
Mandible: length 16.3-18.5. (rarely 13).
Anal fin: length of base 21-23.5. Vertebrae: 41 or 42 (11 specimens).
Bopy moderately compressed, its depth 4.3 5-4.75 in SL in specimens 73-116 mm
SL. Heap 4.0-4.35, its depth slightly exceeding its postorbital length. Snour blunt,
extending about a third of its length beyond tip of mandible, 6.1-7.0 in head. Eve
3-6-4.5. PosrorsiraL length 6.65-8.0 in SL, 1.6-1.8 in head. Maxitiary scarcely
pointed, extending somewhat beyond joint of mandible in the large examples but not
to margin of opercle, 1.25—1.35 in head. ManpiBLe 1.27—1.4. CHEEK moderately short
238 Memor Sears Foundation for Marine Research
and broad, generally about as long as snout and eye, its posterior angle about 30°.
GILL RAKERS moderately slender, those at angle of first arch about as long as pupil,
rather spinulose along inner margin, scarcely expanded at tip.
Dorsat fin with longest rays not reaching tip of posterior ray if deflexed, its margin
generally slightly concave, its origin usually about equidistant between posterior margin
of eye and base of caudal. Anat with a definite lobe anteriorly, its base 4.25—4.75 in
SL, its origin about under middle of dorsal base and equidistant between pectoral base
and caudal base, or a little nearer to the former. Petvic generally extending a little less
Figure 55. Lycengraulis limnichthys, type specimen, from Rio Agua Caliente, above Maracaibo Basin, Vene-
zuelaj USNM 121751. Drawn by A. M. Awl.
than halfway to anal, inserted somewhat nearer to pectoral base than to origin of anal,
2.4-2.8 in head. Pecrorat reaching to or a little beyond base of pelvic in small
specimens, frequently failing to reach pelvic in large ones, 1.25—1.4 in head. Axm-
LARY SCALE of pectoral variable in length, reaching somewhat beyond midlength of
pectoral to beginning of its distal fourth, 1.4—-2.25 in head.
Color. Back pale, with a median dark streak, much more distinct in some speci-
mens than in others. Sides with a broad silvery lateral band in adults, becoming diffused
with the silvery color on the lower part of side in large examples, quite narrow in young.
Dorsal and caudal fins with dusky punctulations, the margin of the latter much darker
in some specimens than in others; other fins plain. Peritoneum silvery.
Size. The largest example in the collection studied is 1;0 mm TL and 116 mm
SL, which may be near the maximum size attained.
Development. The smallest, 19 mm SL, is slender and has all fins well developed.
The caudal fin is as deeply forked as that in the adult, and the other fins have ac-
quired approximately the same shape and relative position as those in adults. The
snout remains shorter and projects less strongly beyond the mandible than in larger
specimens; the maxillary, also much shorter and rounder, fails to reach the joint of the
mandible; and the teeth in the jaws are scarcely evident.
Fishes of the Western North Atlantic 239
The juvenile color of this same specimen and of others of about equal size consists
of a few dark dots over the nostrils, a group of similar dark dots at the occiput, and
more or less definite rows on the back. Scattered dark points, present at and on the
base of the caudal, extend prominently on the outer rays of the fin, which already has
a dusky margin that remains throughout life. A series of prominent black spots along
each side of the anal base continues in a single series on the peduncle behind the anal;
a few scattered dark dots are present on the side of the head and on the isthmus. This
small example, then, is definitely beyond the larval stage and is perhaps a young adult.
The juvenile is slender, as is usual among clupeid fishes. In three specimens, 19,
19.5, and 20mm SL, the depth is respectively 9.0, 9.5, and 9.3 times in SL; in a
group of six, ranging between 27-33 mm SL, the proportionate depth already has in-
creased and is 8.5—6.5 in SL; and in an additional group of five, 32-50 mm SL, the
depth is 6.5—5.45 in SL. Measurements of larger fish indicate that the body tends
to become proportionately deeper as long as the fish grows.
The teeth in the jaws are well developed but remain minute in specimens around
25mm SL. They do not become large (for an anchovy) and unequal in size (the chief
distinguishing characters of Lycengraulis) until the fish attains about 65 mm SL.
In fish 25 mm SL, the snout already projects well beyond the mandible, about
as in the adult, and the maxillary is now somewhat pointed and reaches to a point
approximately opposite the joint of the mandible. The axillary scale of the pectoral,
not detected in the smaller specimens described, is now evident and reaches about to
the midlength of the pectoral; this axillary process, though rather variable in length,
reaches far beyond the midlength of this fin in adults, generally to or beyond the be-
ginning of its distal third.
The juvenile color markings are retained about as described for the 19-mm fish
until a standard length of at least 40 mm is attained. Even then the juvenile markings
remain in large part. However, the sides of the head and the abdomen, in part, have
become silvery. A very narrow indistinct silvery lateral band has become evident, and
this increases rapidly in proportionate width, being prominent and as broad as the eye
in specimens of about 75 mm SL; in examples of about 85 mm SL, the band is much
broader than the eye, and its lower margin generally has become indefinite. In still
larger specimens it usually has become diffused with the silvery color on the lower part
of the side. The dark dots along the base of the anal persist longer than any of the other
juvenile markings and are visible in half-grown fish; in fact, they are visible in some
specimens up to 90 mm SL.
Spawning. Several females with large eggs were taken in Lake Maracaibo on May
16 (1942), indicating that at least some spawning takes place in the spring of the year.
Range. Known only from Lake Maracaibo and from the lower sections of tributary
streams, in fresh to slightly brackish water.
Reference:
Lycengraulis limnichthys Schultz, Proc. U.S. nat. Mus., 99, 1949: 51, fig. 8 (orig. descr.; type local. Rio Agua
Caliente, above Lake Maracaibo, Venezuela; type USNM 121751).
24.0 Memoir Sears Foundation for Marine Research
Lycengraulis batesii (Giinther) 1868
Figure 56
Study Material. A total of 13 specimens, 35-185 mm TL, the larger ones from
Brazil, the smaller ones from Venezuela; also 4 juveniles, 21-31 mm TL, from
Venezuela.
Distinctive Characters. The slender body, the long narrow cheek, and the rather
few and moderately long gill rakers distinguish this species. The statement in the
original description, ‘‘Gill rakers very short, like tubercles,” has been misleading, for
Ficure 56. Lycengraulis batesii, 185 mm TL, 155mm SL, from Teffé, Brazil, MCZ 18006. Drawn by
Alice C. Mullen.
they are fully half of the length of eye, a point that was verified from type specimens
by J. R. Norman. This statement became especially bothersome upon the discovery
of other species that had notably shorter gill rakers.
Description. Proportional dimensions in per cent of standard length, and counts,
based on the Study Material.
Body: depth 16.5-19. Anal fin: length of base 25-26.2.
Head: length 21-23. Pectoral fin: length 15.4—-18.8.
Snout: length 2.75—-3.3.
Eye: diameter 4.2—4.6. Scales: ca. 41-45.
Postorbital: distance 14.3-15.6. Gill rakers: 9-13 + 12-15.
Maxillary: length 17.5-18.2. Fin rays: dorsal 13-15; anal 26-28;
Mandible: length 16.1-18.2. pectoral 13 or 14.
Vertebrae: 47 (2 specimens).
Bopy slender, compressed, its depth 5.25—6.0 in SL. Heap 4.3-4.7 in SL, its
depth about equal to its postorbital length. Snour projecting much less than half of its
length beyond mandible, 7.0-8.0 in head. Eye 4.5—5.4. Posrorsirat part of head
1.4-1.6 in head. Maxitrary slightly expanded, scarcely pointed, reaching nearly or
quite to joint of mandible, 1.2-1.3 in head. ManpipLe 1.27-1.4. CHEEK in large
Fishes of the Western North Atlantic 241
examples much longer than snout and eye, proportionately shorter in small specimens,
its posterior angle about 35°. Gitt RAkeERs at angle fully half of length of eye.
Dorsat fin with the longest rays extending nearly or quite to tip of last ray if
deflexed, its origin much nearer to base of caudal than to eye. Anat base 3.8—4.0 in
SL, its origin slightly in advance of middle of dorsal base. Petvic failing to reach
halfway to anal in adults, extending farther back in young, inserted nearer to pectoral
base than to anal origin. Pecrorat scarcely reaching pelvic in adults, rather longer
in young, I.25—1.5 in head. AxiLLary scat of pectoral about 67 °/, of length of fin,
2.32.7 in head.
Color. Old specimens, in alcohol, grayish above. Side of head and lower 66°/, of
body bright silvery. Small specimens with a broad silvery lateral band and a dark verte-
bral band. The original description mentioned ‘‘an oblong blackish patch on lower half of
the end of the tail,’”’ but such a blackish patch is not present on the specimens studied.
Size. A total length of at least 185 mm (7.4 in.) is attained.
Development. The juvenile, as in other herring and herring-like fishes, is more
slender but rounder than the adult. All juveniles show externally the convoluted
intestine, which apparently has not become fully covered by the abdominal wall. The
snout is very short and blunt, and in the smallest specimen it scarcely projects beyond
the tip of the mandible. The blunt maxillary extends little beyond the posterior margin
of the eye. Both anal fin and vent move forward with age; the anal fin has its origin
under the posterior rays of the dorsal in a 21-mm specimen, about under the middle
of the dorsal base in a 31-mm fish, and in advance of the middle of the dorsal, as in
the adult, in a 35-mm fish.
The smallest fish has a dark line on the isthmus and chest, dark dots along the
base of the anal, and a single continuous dark median ventral line on the caudal peduncle.
The dark line on the isthmus and chest disappears at a length of about 30 mm, but the
markings at the base of the anal and on the peduncle remain until a length of at least
52 mm is attained. The dark vertebral band, described for larger fish, first becomes
partially evident at a length of about 32 mm. There is no indication of a silvery lateral
band up to a length of 52 mm.
Range and Habitat. This species is known from Rio Apure (a tributary of the
Orinoco) at San Fernando de Apure, Venezuela; in Brazil it has been taken from
the Para River, from Teffé on the Solimées River, and from the Jutahy River, the last
two streams being tributaries of the Amazon.
Synonyms and References:
Engraulis batesii Giinther, Cat. Fish. Brit. Mus., 7, 1868: 399 (orig. descr.; type local. Pard R., Brazil;
types in BMNH).
Lycengraulis batesii Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 385 (descr. not based on daresii
throughout; 21 gill rakers on lower limb given whereas dasesii has only 12-15; Rio Grande, Teffé,
Jutahy [River] and Doce R., all in Brazil; Rio Grande specimen is o/idus, the Doce R. specimen schroedert) ;
von Ihering, Rev. Industr. Anim., Anno 1 (3), 1930: 233 (locals., diagn.); Hildebrand, Bull. Bingham
oceanogr. Coll., S (2), 1943: 148, fig. 67 (descr., range); Schultz, Proc. U.S. nat. Mus., 99, 1949: 51
(ref.).
16
24.2 Memoir Sears Foundation for Marine Research
Negative Reference:
Lycengraulis batesii Starks, Stanf. Univ. Publ., Univ. Ser., 1913: 11 (specimen from market at Pard, Brazil, is
L. barbouri Hildebrand).
Lycengraulis abbotti (Fowler) 1915
Figure 57
Study Material. The type and only specimen known, 180 mm long, from Port-of-
Spain, Trinidad.
Ficure 57. Lycengraulis abbotti, type specimen, 180mm TL, from Port-of-Spain, Trinidad, modified after
Fowler, (1915: 523, fig. 1), ANSP 45079. Drawn by Alice C. Mullen.
Distinctive Characters. This fish is related to L. barbouri, having similar short spiny
gill rakers, but it differs in the number of gill rakers, the longer and more pointed
maxillary, and the more anterior position of the dorsal fin.
Description, Proportional dimensions in per cent of standard length, and counts,
based on the type, 180 mm TL.
Body: depth 27. Anal fin: \ength of base 25.6.
Head: length 23.5. Pectoral fin: length 18.2.
Snout: length 3.37.
Eye: diameter 4.95. Scales: ca. 37.
Postorbital: distance 14.8. Gill rakers: 12+16 and I1+17, in-
Maxillary: length 21. cluding rudiments.
Mandible: length 18.3. Fin rays: dorsal 13; anal 25; pectoral 16.
Bopy fairly robust, its depth 3.7 in SL. Heap 4.25 in SL, its depth scarcely
less than its length without snout. SNour projecting less than half of its length beyond
mandible, 7.0 in head. Eye 4.8. Posrorsirat part of head 1.6 in head. MaxiLiary
pointed, extending nearly to margin of opercle, 1.13 in head. Manoprsie 1.3. CHEEK
much longer than snout and eye, its posterior angle about 30°. GiLL RakeRs short,
broad, very spinose, partly rudimentary, none exceeding half of diameter of pupil.
Fishes of the Western North Atlantic 24.3
Dorsat fin with the longest rays not quite reaching tip of last ray if deflexed, its
origin equidistant between base of caudal and middle of eye. Anat origin slightly in
advance of middle of dorsal base, its base 3.9 in SL. Petvic reaching less than halfway
to anal, inserted nearer to base of pectoral than to origin of anal. Pecrorat not quite
reaching base of pelvic, 1.3 in head. Axitiary scate of pectoral fully 75°/, of fin,
1.8 in head.
Color. Grayish above. Sides silvery. No lateral band. Caudal with a narrow dark
margin.
Range. Known only from the type taken at Port-of-Spain, Trinidad.
Synonyms and References:
Anchovia abbotti Fowler, Proc. Acad. nat. Sci. Philad., 1915: 522, fig. 1 (orig. descr.; type local. Port-of-Spain,
Trinidad; type ANSP 45079; cf. Stolephorus surinamensis Bleeker = Engraulis edentulus Cuvier).
Anchoviella abbotti Jordan, Evermann, and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 50 (name, local.).
Lycengraulis abbotti Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 150, fig. 68 (cf. L. darbouri
Hildebrand and L. schroederi Hildebrand).
Lycengraulis barbouri Hildebrand 1943
Figure 58
Study Material. Type material, 3 specimens 237, 230, and 205 mm long, from
the Rio Poty, Brazil, and a fourth specimen, 220 mm long, from the fish market at
Para; holotype and one paratype, MCZ 35277 and 18017, respectively; second paratype
USNM 118997; and fourth specimen SU 22086.
Distinctive Characters. This species differs from L. batesii in having a deeper body,
a somewhat more posteriorly placed anal fin, and much shorter and broader gill rakers;
however, the rakers are about equal in number in the two species. L. schroederi Hil-
debrand, known only from Rio Doce in southern Brazil, is another related species; it
has more numerous but equally short, broad, spiny, partly rudimentary gill rakers
(17 or 18+21 or 22 on the first arch), a larger eye, about 4.9 in head, and a more
posteriorly placed anal, its origin under the posterior third of dorsal base.
Description. Proportional dimensions in per cent of standard length, and counts,
based on the four study specimens, 205-237 mm.
Body: depth 21.2—23.2. Anal fin: length of base 24.3-26.2.
Head: length 22-25.5. Pectoral fin: length 17—18.2.
Snout: length 2.75—3.0.
Eye: diameter 4.25-4.45. Scales: ca. 40-45.
Postorbital: distance 14.5-17.7. Gill rakers: 9-11 + 12-14.
Maxillary: length 18.2—21. Fin rays: dorsal 14 or 15; anal 26-28;
Mandible: length 17.5-20.8. pectoral 14 or 1S.
Bopy rather strongly compressed, its depth 4.3-4.7 in SL. Heap 3.9-4.5 in SL,
its depth about equal to its postorbital length. Snour projecting much less than half
16*
244 Memoir Sears Foundation for Marine Research
of its length beyond mandible, 7.8—9.2 in head. Eve 5.0—5.8. Posrorsitat part of head
1.37-1.5 in head. Maxitiary rounded distally, scarcely reaching joint of mandible,
1.2 in head. Manprsie 1.2—1.27. CHEEK narrow, its length nearly twice snout and
eye, its postorbital angle about 30°. Grizz rakers short, flat, the longest not more than
half of length of pupil. .
Dorsat fin with the longest rays extending to, or a little beyond, tip of last ray if
deflexed, its origin about equidistant between base of caudal and upper anterior angle
of gill openings. Anat with a rather prominent lobe anteriorly, its origin under, or some-
/
Ficure 58. Lycengraulis barbouri, type specimen, 290 mm TL, 237 mm SL, from Rio Poty, Brazil, MCZ
35277. Drawn by Alice C. Mullen.
what in advance of, middle of dorsal base, its base 3.8—4.1 in SL. Petvic reaching much
less than halfway to anal, inserted somewhat nearer to base of pectoral than to origin
of anal. Pecrorat reaching to, or slightly beyond, base of pelvic, 1.3—1.4 in head. Axri-
LARY SCALE of pectoral about 75°/, of length of fin, 1.95—2.1 in head.
Color. Old specimens, in alcohol, grayish above. Lower 75°/, of body and
head plain silvery. Fins unmarked, except for an indication of a black margin on
caudal.
Range and Habitat. Known only from northern Brazil, from the Rio Poty, a trib-
utary of the Rio Parnahyba, and from the Belém (Para) market. [That this species was
obtained in the Belém market, far removed from Rio Poty, seems to indicate that the
species may well be marine.—y. H.0.]
Synonyms and References:
Lycengraulis batesii Starks (not of Giinther), Stanf. Univ. Publ., Univ. Ser., 1913: 11 (specimen from Pard,
Brazil; found to be L. darbouri).
Lycengraulis grossidens Jordan and Seale (in part not of Cuvier), Bull. Mus. comp. Zool. Harv., 67, 1926: 383
(“Rio Puty”= Rio Poty, Brazil; specimens are L. darbouri).
Lycengraulis barbouri Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 151, fig. 69 (orig. descr.; type
local. Rio Poty, tributary of Rio Parnahyba, Brazil; type MCZ 35277; cf. L. datesii).
Fishes of the Western North Atlantic 24.5
Genus Cetengraulis Giinther 1868
Cetengraulis Giinther, Cat. Fish. Brit. Mus., 7, 1868: 383; genotype Exgraudis edentulus Cuvier, first designated
by Jordan and Evermann, Bull. U.S. Nat. Mus., 47 (1), 1896: 450.
Characters. Bopy rather deep in adults but much more slender in young, quite
strongly compressed. Maxittary short, distally rounded or very bluntly pointed,
usually extending about to articulation of mandible. Trrru in jaws all very small (ap-
parently not disappearing with age as has been stated). Grit Raxers long, close-set,
numerous, increasing with age (about 2 5 on lower limb in small specimens to about 60 in
large ones). Petvic inserted under, or a little in advance of, dorsal origin. Silvery lateral
band present only in specimens about 100 mm TL or less.
Remarks. Gill covers connected across the isthmus by a thin transparent membrane
(easily torn); narrow in very young.
Range. The genus is represented on the Atlantic side by two species, ranging
from the West Indies to southern Brazil, and on the Pacific side by one species, C. mysti-
cetus, known from the Gulf of California to Peru.
Key to Atlantic Species
1a. Depth of body 3.0-3.4 in length; head 4.6—5.3; eye 3.7-4.6 in head; pectoral
failing to reach base of pelvic. edentulus (Cuvier) 1829, below.
tb. Depth of body 5.0 in length; head 3.5; eye 5.5 in head; pectoral reaching a little
beyond base of pelvic. juruensis Boulenger 1898, p. 247.
Cetengraulis edentulus (Cuvier) 1829
Bocéon (Cuba)
Figure 59
Study Material. About 70 specimens, mostly small, 35-160 mm TL; from Puerto
Rico; Jamaica; Cuba; and New Grenada; Colén, Panama; Gulf of Venezuela; Rio de
Janeiro, ‘Barro Santas,” ‘““Sambaia,’”’ and Cachoeira, Brazil.
Distinctive Characters. See Key to Atlantic Species, above.
Description. Proportional dimensions in per cent of standard length, based on
20 specimens, 70-160 mm, and most of the counts on many more.
Body: depth 25—34.4. Scales: ca. 40-43.
Head: length 30-34.5. Gill rakers: ca. 30+36 in specimens
Snout: length 3.8-4.75. around 65mm, about 52+ 55 in
Eye: diameter 6.5—8.0. specimens about 130 mm TL.
Postorbital: distance 18.2-21.2. Fin rays: dorsal 14-16; anal 23-26;
Maxillary: length 17.2-19. pectoral 14 or 15.
Mandible: length 17.3-18.5. Vertebrae: 42 (3 specimens).
Anal fin: length of base 21.5-23.
Pectoral fin: length 13.3-16.7.
24.6 Memor Sears Foundation for Marine Research
Bopy strongly compressed, the ventral outline much more strongly convex than
the dorsal outline, the depth increasing with age, 2.9—-3.3 in SL in specimens 75 mm
TL and upward. Hzap 3.0-3.5 in SL. Snour quite pointed, extending nearly its
full length beyond mandible, 5.5—8.0 in head. Eye 3.7-4.8. Posrorsirat part of
head 1.55-1.75 in head. Maxitrary bluntly pointed or rounded distally, 1.72—1.8
in head. Manp1sLe 1.67—1.73. CHEEK nearly or quite as long as snout and eye, its
posterior angle narrow, about 30°. Git raxers long, slender, close-set, increasing
greatly in number with age.
Ss)
= EG
HA
Ficure 59. Cetengraulis edentulus, 120mm TL, 1oomm SL, from Havana, Cuba, USNM 35159. Drawn
by Louella E. Cable.
Dorsat fin with the longest rays failing to reach tip of last ray if deflexed, its
origin generally about equidistant between anterior margin of eye and caudal base.
Anat with origin usually under last fourth of dorsal base, its base 4.3-4.6 in SL.
Perxvic inserted nearer to anal origin than to pectoral base, generally reaching fully
halfway to origin of anal. Pecrorat usually failing to reach base of pelvic, 1.9—2.3
in head. AxiLLaRy scaLE of pectoral broad, with a somewhat thickened ventral edge,
3.25—3.7 in head.
Color. In alcohol, bluish gray above; lower 75°/, of side silvery. Small specimens
with a silvery lateral band, becoming wider and less distinct with age, generally dis-
appearing at a length of about 100 mm.
Size. Specimens as much as 165 mm (6.6 in.) TL have been reported.
Range and Habitat. Reported for the West Indies and the Atlantic coast of
Panama to southern Brazil. It ascends freshwater streams.
Synonyms and References:
Engraulis edentulus Cuvier, Régne Anim., ed. 2, 2, 1829: 323 (orig. descr.; type local. Jamaica); Cuvier and
Valenciennes, Hist. Nat. Poiss., 27, 1848: 51 (descr., abund., distr.).
Cetengraulis edentulus Giinther, Cat. Fish. Brit. Mus., 7, 1868: 383 (descr., range); Jordan and Evermann,
Bull. U. S. nat. Mus., 47 (1), 1896: 450 (descr., range, synon.); Meek and Hildebrand, Field Mus.
Fishes of the Western North Atlantic 24.7
Publ., Zool., 75 (1), 1923: 214 (descr., range; Sto/ephorus garmani Evermann and Marsh and 8. gi/éerti
Evermann and Marsh put in synon.); Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67, 1926: 414
(descr., Cuba and Brazil); Beebe and Tee-Van, Zoologica, N. Y., 10 (1), 1928: 49, fig. (descr., Port-
au-Prince, Haiti); Nichols, N. Y. Acad. Sci., Zo (2), 1929: 206, fig. (synon., diagn., Puerto Rico); Jor-
dan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 51 (range, synon.); Fowler, Proc.
Acad. nat. Sci. Philad., 80, 1931: 393 (Trinidad, B.W.I.); Beebe and Hollister, Zoologica, N. Y., 19
(6), 1935: 211 (Union I., B.W.1.); Fowler, Arqu. Zool. estad. Sao Paulo, 3 (6), 1941: 135 (Brazilian
recs.); Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 155, fig. 71 (descr., synon., range);
Schultz, Proc. U.S. nat. Mus., 99, 1949: 54 (refs.).
Engraulis brevis Poey, Repert. Fisico-Nat. Cuba, 1, 1866: 379 (orig. descr.; type local. Cuba; cf. E. edentulus
Cuvier); Howell-Rivero, Bull. Mus. comp. Zool. Harv., 82, 1838: 172 (holotype, MCZ 24296, Poey’s
No. 716).
Stolephorus robertsi Jordan and Rutter, Proc. Acad. nat. Sci. Philad., 49, 1897: 95 (orig. descr.; type local.
Jamaica; type SU 4853; cf. Sto/ephorus opercularis Jordan and Gilbert, a Pacific coast Cetengraulis);
Jordan and Evermann, Bull. U.S. nat. Mus., 47 (3), 1898: 2815 (descr. after Jordan and Rutter).
Stolephorus gilberti Evermann and Marsh, Rep. U.S. Comm. Fish. (1899), 1900: 352 (orig. descr.; type local.
Palo Seco, Puerto Rico; type USNM 49359; cf. Engraulis productus Poey = Engraulis clupeoides Swain-
son and Szolephorus garmani E. and M.= Engraulis edentulus Cuvier); and Bull. U.S. Fish Comm.
(1900), 20 (1), 1902: 90, fig. 15 (orig. descr.); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (4),
1900: 3146 (descr. after Evermann and Marsh).
Stolephorus garmani Evermann and Marsh, Rep. U.S. Comm. Fish. (1899), 1900: 352 (orig. descr.; type local.
Puerto Real, Puerto Rico; type USNM 49360; cf. Engraulis productus Poey = Engraulis clupeoides
Swainson, and Stolephorus gilberti E. and M.= Engraulis edentulus Cuvier); and Bull. U.S. Fish Comm.
(1900), 20 (1), 1902: 89, fig. 14 (orig. descr.); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (4),
1g00: 3146 (descr. after Evermann and Marsh).
Anchoviella robertsi Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 49 (range, synon.).
Cetengraulis gilberti Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 51 (range, synon.);
Howell-Rivero, Proc. Boston Soc. nat. Hist., 47 (4), 1936: 55 (Cuba).
Cetengraulis garmani Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 51 (range, synon.).
Doubtful References:
Stolephorus surinamensis Bleeker, Tijd. Dierk. Amst. (1865) 3, 1866: 178 (orig. descr. of g6-mm specimen;
type local. Surinam; nothing in descr. inconsistent with C. edentu/us of similar size); Eigenmann,
Mem. Carneg. Mus., 5, 1912: 448 (descr. of 27-and 37-mm specimens from Bartica Rocks, British Guiana) ;
Puyo, Bull. Soc. Hist. nat. Toulouse, 80, 1945: 103, fig. 2 (descr., French Guiana); Puyo, Faune Emp.
Frang., 12, Poiss. Guyane Frang., 1949: 155, fig. 78 (descr., French Guiana).
Stolephorus manjuba Miranda-Ribeiro, Kosmos, Rio de J., Feb., 1908, 5—no pagination (orig. descr. inadequate;
type local., Rio de Janeiro, Brazil).
Anchovia gilberti Fowler, Proc. Acad. nat. Sci. Philad., 1916: 401 (Colén, Panama, without comment).
Anchoviella surinamensis Fowler, Proc. Acad. nat. Sci. Philad., 80, 1931: 392 (three specimens, 34-36 mm
long, from Icacos Beach, Trinidad, B.W.I.; notes suggest C. edentulus).
Cetengraulis juruensis Boulenger 1898
Study Material. None.
Distinctive Characters. See Key to Atlantic Species, p. 245.
Description. Based on original description and figure, which is all that is known.
Bopy depth 20°%/, of SL. Heap 28.5 in SL. Snour projecting strongly (extending
nearly its full length beyond mandible, according to a published figure). Eve 5.5 in
head. Maxiiiary extending to joint of mandible. Gitt rakers long, finely denticulate,
about 40 on lower limb.
24.8 Memoir Sears Foundation for Marine Research
Dorsat origin equidistant between tip of snout and caudal base; 13 rays. ANAL
origin under posterior rays of dorsal; rays 23. Pxtvic fin inserted somewhat nearer to
base of pectoral than to origin of anal. Pecrorat extending a little beyond base of
pelvic, its length a little more than half of length of head. AxitLary scate of pectoral
extending somewhat beyond midlength of fin (according to figure). Scatzs 38.
Color. Olive above, silvery on sides and below. Snout blackish above. Fins pale
orange; caudal rays blackish at tips.
Range and Habitat. Known only from the holotype, a specimen 140 mm TL, from
fresh water, from the Jurua River, a tributary of the Amazon in Brazil.
References:
Cetengraulis juruensis Boulenger, Trans. zool. Soc. London, 14 (10), 1898: 427, pl. 41, fig. 3 (orig. descr.;
type local. Rio Jurud, Brazil; cf. C. edentulus); Eigenmann, Rep. Princeton Exped. Patagonia, 1896—
1899, 3 (4), 1910: 452 (refs. to orig. descr., habitat); Campos, Arqu. Zool. estad. Sao Paulo, 3 (7), 1941:
202, fig. 15 (descr., range); Hildebrand, Bull. Bingham oceanogr. Coll., 8 (2), 1943: 158 (descr. after
Boulenger).
bo
2
Io.
TEXT REFERENCES
[Pasian Proc. Acad. nat. Sci. Philad., 82,
1930.
Fowler, Proc. Acad. nat. Sci. Philad., 83,
1931.
Hupesrano, Bull. Bingham oceanogr. Coll.,
8 (2), 1943.
Hildebrand and Cable, Bull. U. S. Bur. Fish.,
46, 1930.
Hildebrand and Schroeder, Bull. U. S. Bur.
Fish. (1927), 43 (1), 1928.
dee fes and Seale, Copeia, 1925.
Jordan and Seale, Bull. Mus. comp. Zool. Harv.,
67 (11), 1926.
Keone Bull. Boston Soc. nat. Hist., 58,
1931.
Kuntz, Bull. U.S. Bur. Fish., 33, 1914.
Kuntz and Radcliffe, Bull. U. S. Bur. Fish.
(1915-16), 35, 1918.
249
If.
T2.
3.
T4.
T5.
Lintoy, Bull. U. S. Fish Comm. (1899), 79,
gol.
Linton, Bull. U.S. Bur. Fish. (1904), 24, 1905.
Myers, Proc. Calif. Acad. Sci., (4) 23, 1940.
Naxasuima (Shoji), Bol. Mus. Hist. nat.
‘Javier Prado,’ 5 (16), 1941.
Nichols and Breder, Zoologica, N. Y., 9 (1),
1927.
Beeson Indiana Univ. Stud., rz (64), 1924.
Poey, Synop. Pisc. Cubensis, 1868.
Poey in Gundloch, An. Soc. esp. Hist. nat.,
Io, 1881.
Scuutrz, Proc. U.S. nat. Mus., 99, 1949.
Swain, Bull. U.S. Fish Comm., 2, 1883.
Vianyxoy, Proc. N. S. Inst. Sci., 79, 1935.
Interim Account of
Family Alepocephalidae
HENRY 5. BIGELOW
Museum of Comparative Zoology
Harvard University
Characters. Essentially as in the Clupeidae, except with the Dorsat fin consider-
ably posterior to the midlength of body, its base partly or entirely over anal fin and
preceded in a few (Anomalopterus) by a fleshy fold or ridge. LaTeRAL LINE present in
most (described as indistinct or lacking in Xenodermichthys, Leptoderma, and Anomal-
opterichthys). SWIM BLADDER lacking. CoLor ranging between dark blue, purple, and
black, below as well as above; transparent but never silvery. Barsets lacking on chin
and throat. Bony GuLAR PLATE absent in throat region. ExTERNAL TUBULAR PAPILLA
absent on shoulders (present in Searsiidae, p. 254). ABpoMEN rounded, its midline
without keeled scales (scutes). Snour less than twice the length of mouth in most,
not tubular, but tubular and more than twice the mouth in a few (4ulostomatophora).
Movutu ranging from very small, at tip of tubular snout (see above), to a little more
than half as long as head (A4nomalopterus and Bathytroctes). Pecrorat fins very small;
rudimentary in a few (Mirognathus). Prtvic fins present, at least in most. TEETH small,
feeble, present on premaxillaries in most (upper jaw toothless in Axomalopterichthys,
Leptochilichthys, and Asquamiceps), and on maxillaries, palatines, and mandible in some.
ScaLes present or absent; if present, thin, rounded. Luminescent orGaNns (photophores)
lacking in most but present in a few; situated on nodules of raised skin if present.
VENT posterior to midlength of trunk in most (anterior to midlength of trunk in Lep-
toderma). GILL MEMBRANES free from isthmus and overlapping. OperRcuULAR sYSTEM
thin but complete. Girt RaKeRs long, numerous. BRANCHIOSTEGAL RAYS § or more.
PyLoric CAECA 2—21 reported (rudimentary in some).
Genera. Parr, in his recent synopsis of the family, recognized 22 genera, six of
which were described by him as new (4: 1-14); to these he added two more new genera
a year later (5: 263, 265).
250
Lishes of the Western North Atlantic 205
Remarks. The Alepocephalidae differ externally from the true herrings (Clupeidae)
in their very soft texture, in their uniformly dark, dull coloration (below as well as above),
and in the position far rearward of their dorsal fin. They differ internally from the her-
rings in lacking a swim bladder.
Ficure 60. a. Alepocephalus productus, from ALBatross St. 2030, off New Jersey, 39°30’ N, 71°43’W, in 588
fms.; after Goode and Bean. 8. A/epocephalus bairdii, from Grand Banks off Newfoundland, at 200 fms.;
after Goode and Bean. c. A/epocephalus agassizii, from off New Jersey; after Goode and Bean. pv. Xexodermich-
thys copei, from off Chesapeake Bay; after Goode and Bean. £. Aulostomatophora phosphorops, from the Arabian
Sea; after Goode and Bean, based on Alcock.
The members of the family as a whole are creatures of the deep sea, seldom taken
in depths less than 250 fms. or so; a cursory examination of the literature revealed
only two records for shallower depths—20o fms. for the Grand Banks off New-
foundland for Alepocephalus bairdii (I: 55) and 180 fms. north of Celebes for Bathy-
troctes macrolepis (3: 225). The alepocephalids can be described as “universal” in all
oceans, down the lower part of the slopes and on the ocean floors at depths to at
least 3000 fms. (5490 m) or so, and very likely deeper still.
22 Memoir Sears Foundation for Marine Research
Table I. Hauls from off Southern Nova Scotia to Virginia
Containing Alepocephalids
Av. no. Specimens
Depth Total Hauls with race in Hauls with
(fms.) Hauls Alepocephalids Ue peea eee Alepocephalids
50-250 76 ° 2 4
251-300 18 I 6
301-350 22 4 x8 20
351-400 15 6 ae 7
401-450 21 2 38 :
451-500 21 13 62 Be
501-550 8 : We 4
551-600 4 4 a 6
601-650 4 4 mee m3
651-700 2 5 ie e
701-750 2 2 ea ?
During the experimental cruises that were carried out from the Woods Hole
Oceanographic Institution by Mr. William C. Schroeder on the dragger Capn. Brit 11
in 1952-1953 along the continental slope of North America between the offings of
southern Nova Scotia and Virginia, the shallowest haul that brought up an alepoce-
phalid was at 251-350 fms.1 Alepocephalids, however, of one kind or another were
taken in more than a third of the hauls from deeper than 351-400 fms. and in all
of the hauls from deeper than 500 fms., as shown in Table 1, contributed by Mr.
Schroeder.
A detailed account of the family is unavoidably postponed until a subsequent
volume. The general appearance of typical members is represented in the accompanying
outline drawings (Fig. 60, 614).
1. A depth of “68” fms. given by Goode and Bean for Concara macroptera should read 687 fms. (2: 40); see
Townsend (6: 403).
TEXT AND FOOTNOTE REFERENCES
Ei Goopr and Bean, Proc. U. S. nat. Mus., 2, 4. Parr, Amer. Mus. Novit., 1531, 1951.
1879. 5. Parr, Bull. Mus. comp. Zool. Harv., 107 (4),
2. Goode and Bean, Smithson. Contrib. Knowl., 1952.
30, 1895.
3. Giinther, Rep. sci. Res. ‘Challenger,’ Zool.,
22 (57), 1887. 6. "Townsenp, Rep. U. S. Comm. Fish., rgor.
253
Interim Account of
Family Searsiidae
HENRY B. BIGELOW
Museum of Comparative Zoology
Harvard University
Characters. Essentially as in the Alepocephalidae, except that there is a volu-
minous sac extending forward underneath the skin of each shoulder and opening
rearward through a TuBULAR PAPILLA so conspicuous that it affords a convenient and
reliable field mark by which a member of this family can be recognized. This shoulder
apparatus appears to be a secretory organ, probably producing luminous mucus.
Petvic fins are wanting in a few (Platyrroctes).
Remarks. The type genus Searsia was described by Parr in 1937 (1:12), but it
was not until 1951 (2:2) that he proposed and defined the family Searsiidae to include
the genera Searsia Parr 1937, Platytroctes Giinther 1878, Platytroctegen Lloyd 1909,
Holtbyrnia Parr 1937, Barbantus Parr 1951, Normichthys Parr 1951, Pellisolus Parr
1951, and Persparsia Parr 1956. [As this paper goes to press, Parr’s recent extensive
and complete monograph of the known Searsiidae is at hand (3). To the genera already
listed above, he has added Mirorictus Parr 1947, Sagamichthys Parr 1953, Maulisia Parr
1960, and Mentodus Parr 1951, with Mirorictus in the subfamily Mirorictinae, P/aty-
troctes and Platytroctegen in the subfamily Platytroctinae, and all the other genera in the
subfamily Searsinae.—y.H.0. ]
The Searsiidae, like the Alepocephalidae (p. 250), are deep-sea fish, thus far re-
ported in the western North Atlantic for the offing of Delaware Bay; the Caribbean,
the offing of Pernambuco, equatorial Brazil; and the vicinity of Bermuda; also for
the middle and eastern Atlantic off northwestern Africa, Cape Verde, and the Gulf of
Gascony; the offing of southern Africa in the eastern South Atlantic; the Bay of
Bengal; and the eastern equatorial Pacific off the Bay of Panama. The shallowest
haul from which any searsiid has yet been reported was from 500 fms., in the Bay
of Bengal.
254
Fishes of the Western North Atlantic Ale tc
The detailed account of the Searsiidae, like that of the Alepocephalidae (p. 250),
is unavoidably postponed until a subsequent volume. The general appearance of a
member of the Alepocephalidae (Fig. 61a) and two of the included genera in
Searsiidae (Fig. 61 B, c) are shown in the accompanying outline drawings.
Ficure 61. a. Leptoderma macrops (Alepocephalidae), from off northwestern Africa; after Goode and Bean,
based on Vaillant. 8. Searsia koefoedi, from off the Bahamas; after Parr. c. P/atytroctegen mirus, from the Bay
of Bengal; after Lloyd.
TEXT REFERENCES
Te Paze, Bull. Bingham oceanogr. Coll., 3(7), 1937- 2. Parr, Amer. Mus. Novit., 1531, 1951.
3. Parr, ‘Dana’ Rep., 5, 1960.
256
Family Clupeidae’
SAMUEL F. HILDEBRAND?
excepting
Genus Harengula by Luis R. Rivas
Genus Dorosoma by Robert R. Miller
Characters. Bopy usually slender in most but short and deep in a few species;
slightly to excessively compressed in most but almost round in some. ScALEs covering
body; cycloid, serrate, or pectinate; head naked. Chest and abdomen compressed into
a sharp edge, armed with Bony scurTes, except in the Dussumierinae (see 1a in Key to
Genera). Larerat LINE wanting. Mouru moderately large, terminal, superior or infe-
rior. TEETH in jaws small, or canine-like in Chirocentrinae (see 11 b in Key to Genera),
sometimes wanting in large adults of 4/osa. GiLL RAKERsS usually long, slender, nu-
merous, increasing in number with age and growth in some species, in moderate
numbers in others. GILL MEMBRANES not united, free from the isthmus. PsEuDoBRAN-
CHIAE present. Dorsat fin, if present, usually about at midlength of the standard
length, but sometimes posterior (present in American species). AprposE fin absent.
Caupat fin forked. Anat moderately long to very long. Petvic fin moderately large,
small, or absent. Pecrorat fin well developed. VerreBRaz in species of the western
North Atlantic about 39-59.
Remarks. This large family, composed of about 70 genera and perhaps 150 or
more species, of which 16 genera and 36 species come within the scope of this work,
is economically one of the most important groups of fishes in the world. Many of the
species occur in great schools, making it easy for either man or natural predators to
1. Edited, with some revision and expansion, by George S. Myers, Henry B. Bigelow, Mywanwy M. Dick, and Yngve
eee subfamilies recognized herein, particularly the Dussumierinae and Dorosomatinae, have been treated
as separate families in some recent works. Nevertheless their affinities are with the Clupeidae. For practical purposes,
therefore, the characters of the subfamilies have been sufficiently indicated in the Key to Genera (p. 259) for the
major subdivisions to be recognized without further treatment elsewhere.
2. August 15, 1883 — March 16, 1949.
S57)
258 Memoir Sears Foundation for Marine Research
catch them in large quantities; and the supply of some, such as those of the genus
Clupea, seems to be almost inexhaustible. Fresh herring are used extensively as food
by man in nearly all parts of the world, and in many areas large quantities are eaten
after they have been salted and smoked or canned. Great quantities, especially of
Brevoortia, are reduced to oil, fish meal (feed for livestock and poultry), and fertilizer.
The smaller species and the young of the larger ones serve still another useful purpose
in providing extensive forage for larger predatory fishes, many of which enter into
the diet of man. Aquatic mammals, birds, and turtles also feed on them. All this
food is provided for man and beast at virtually no cost, as the herring feed almost
entirely on small organisms that are not suitable food for man or for the other preda-
tors named.
Although the members of the family Clupeidae have no lateral line, nearly all
of the species described in this work have numerous mucous pores and canals about
the head and shoulders that may in part compensate for the absence of a lateral line.
An air bladder with a very thin wall is present in all the species of the genera examined,
namely Clupea, Alosa, Pomolobus, Brevoortia, Opisthonema, Harengula, Sardinella, and
Etrumeus, and it may be assumed that the other members of the family also have an
air bladder. Its walls are so thin and delicate that dissections must be made with care
to find it undamaged.
Range. Herring (Clupeidae) occur in all seas of the world except those of the Arc-
tic and Antarctic regions. Many species ascend freshwater streams to spawn, and
others live in fresh water permanently.
Preparation of Descriptions and Location of Study Material. Nearly all Descriptions
were prepared directly from specimens, but a few are included for which no speci-
mens were available. In such instances it is stated in the accounts that they were
compiled or quoted from publications. The specimens used are in the USNM unless
otherwise stated. Freshwater species occurring in the rivers draining into the western
North Atlantic are not numerous and have been included for the sake of complete-
ness; it is possible that all of these forms occasionally enter brackish or salt water.
Measurements and Counts. The proportions used in Description are based on measure-
ments made with vernier calipers having sharp straight points. Many of the measure-
ments of small fish (sometimes even of larger ones) and of the small structures such
as eye, snout, and maxillary were made under magnification. See also Measurements
and Counts under Brevoortia, p. 343.
ToTaL LENGTH: distance between margin of snout or tip of lower jaw (whichever
is longer), more or less along the body axis to a vertical at tip of longest lobe of
caudal fin.
STANDARD LENGTH: distance between anterior margin of snout and base of caudal.
Bopy pEPTH: greatest depth unless otherwise stated; this may occur over base of
pectoral or as far back as origin of dorsal fin.
Heap: if used without a modifier, the distance between anterior margin of snout
and most distal point of bony (not membranous) margin of opercle.
Fishes of the Western North Atlantic 259
Heap pbepTH: distance from a slight crossgroove at the occiput, between the
naked head and scaly body, to margin of first ventral scute; this line may be vertical
or may slope somewhat backward from the occipital crossgroove.
Snout: distance from anterior margin of snout to anterior margin of orbit.
Eye: horizontal diameter of eyeball, not of orbit.
Maxittary: distance from anterior rim of snout to maxillary’s posterior margin.
Manpis_e: distance from its tip to its joint.
INTERORBITAL: width of bone over or between middle of eyes.
Cueek: the bone, usually with a glossy surface in the herrings, lying below and
usually somewhat behind the eye, its depth measured from lower rim of orbit to its
lowest or deepest point.
PREDORSAL LENGTH: distance between anterior margin of snout and origin of
dorsal.
Caupal Loszs: distance from midbase of this fin to tip of lobe.
Privic AnD Pecrorat Fins: distance from base of outer or upper ray to tip of fin.
Fin ray count: includes all rays, whether rudimentary, simple, or branched;
the last ray, even though nearly double, counted as one.
Scates: range in scale numbers, e.g. 56-62, obtained by counting the oblique
rows (upward and slightly forward) that cross the side between upper anterior angle
of gill opening and base of caudal; any deviation from that procedure is stated.
GILL RAKERS: number present on first gill arch, including rudiments if present;
all enumerations based on adult fish unless stated otherwise.
VENTRAL scuTEs: three sets of counts are given: the range of the total number
present, the number in front of the pelvic fins, and the number behind these fins; the
scute ending nearest to the insertion of the pelvics is included in the number in
advance of these fins; the rest compose the number behind.
VeRTEBRAE: total number, including the hypural.
Key to Western North Atlantic Subfamilies and Genera of Clupeidae?
1a. Body little compressed; abdomen rounded, covered with ordinary scales, without
bony scutes. Subfamily Dussumierinae.
Atlantic coast of North America;
eastern and western Pacific; south-
eastern coasts of Africa and Australia.
2a. Pelvic fin much smaller than pectoral, inserted behind base of dorsal; dorsal
fin with about 17-20 rays; chest round. Etrumeus Bleeker 1853, p. 262.
2b. Pelvic fin nearly as large as pectoral, inserted under base of dorsal; dorsal
fin with only about 9-14 rays; chest slightly keeled.
Fenkinsia Jordan and Evermann 1896, p. 267.
3. This Key is drawn up simply for the ready identification of representatives of the subfamilies and genera that are
known from the western North Atlantic without reference to those outside of this range. Therefore, some characters
are used that apply only to the representatives described in this work and for that reason are not of generic value.
iyfns
260 Memor Sears Foundation for Marine Research
1b. Body moderately to very strongly compressed; chest and abdomen compressed
into a sharp keel armed with bony scutes.
3a. Median line of back in front of dorsal fin with ordinary or modified scales,
thus without a prominent median naked strip; stomach not gizzard-like, but
wall somewhat thickened in some species.
4a. Pelvic fin well developed, inserted under base of dorsal, with 7-9 rays;
mouth gently oblique, the lower jaw usually included but occasionally
projecting; body moderately compressed. Subfamily Clupeinae.
Virtually all temperate and tropical
waters of the world, with some species
entering frigid zones; inhabiting salt,
brackish, and fresh water.
sa. Vertical edge on rim of shoulder girdle (cleithrum) underneath free
edge of gill cover (opercle) smooth or even, without dermal lobes.
6a.
6b.
Back in front of dorsal fin with ordinary scales like those on
rest of body; pelvic with 8 or 9 rays; pectoral fin without a
free axillary process; scales not overlapping deeply, their ex-
posed part not much deeper than long, their margin slightly
irregular, not definitely serrate or pectinate.
7a. Vomer with teeth; greatest depth of body about 4.0-5.0
in SL; ventral scutes weak, numerous, total number about
39-46. Clupea Linnaeus 1758, p. 274.
7b. Vomer toothless; depth of body generally less than 4.0 in
SL; ventral scutes strong, total number about 27-38.
8a. Upper margin of lower jaw within mouth rising gently;
teeth missing in examples 200mm and upward in
length; cheek deeper than long (Fig. 69).
Alosa Link 1790, p. 293.
8b. Upper margin of lower jaw within mouth rising
abruptly; teeth present, at least in lower jaw, at all
ages; cheek not deeper than long, definitely longer than
deep in some species (Fig. 76).
Pomolobus Rafinesque 1820, p. 312.
Back in front of dorsal fin with a row of enlarged modified
scales on each side of median line; pelvic with 7 rays; pectoral
fin with a free axillary process; scales overlapping deeply, their
exposed part much deeper than long, their margin strongly
serrate to pectinate (Fig. 85). Brevoortia Gill 1861, p. 342.
5b. Vertical edge on rim of shoulder girdle (cleithrum) underneath free
edge of gill cover (opercle) with two dermal lobes (Fig. 100).
ga.
Last ray of dorsal greatly produced, filamentous, often reaching
nearly or quite to base of caudal (Fig. 94); anal with 20-25
rays. Opisthonema Gill 1861, p. 380.
Fishes of the Western North Atlantic 261
gb. Last ray of dorsal little if any longer than the preceding ones;
anal with about 15-20 rays.
toa. Last two rays of anal not enlarged; gill rakers about 25-
40 on lower limb; scales about 34—41 oblique series cross-
ing middle of side; pelvic with 8 rays.
Harengula Cuvier and Valenciennes 1847, p. 386.
10b. Last two rays of anal enlarged, finlet-like (Fig. 100); gill
rakers about 70-130 on lower limb; scales about 41-47
oblique series crossing middle of side; pelvics with 9 rays.
Sardinella Cuvier and Valenciennes 1847, p. 397.
4b. Pelvic fin small or wanting, inserted in advance of dorsal fin if present;
mouth strongly oblique to nearly vertical, the lower jaw projecting; body
strongly to excessively compressed.
11a. Teeth in jaws all small, without canines; vomer toothless.
Subfamily Pristigasterinae.
12a. Maxillary with a short sharp spine near anterior end, just
behind premaxillary (Fig. 103); anal fin small, beginning far
behind base of dorsal, with only about 15-18 rays.
Rhinosardinia Eigenmann 1912, p. 411.
12b. Maxillary without a spine; anal fin much longer, beginning
in front of dorsal fin, under it, or immediately behind it,
with more than 30 rays.
13a. Pelvic fin present; dorsal fin moderately large, with
about 15—20 rays. Ilisha Gray 1846, p. 415.
13b. Pelvic fin wanting; dorsal fin with about 10-19 rays.
14a. Body with chest and abdomen greatly dilated,
more or less half-moon shaped (Fig. 108), its
greatest depth about half of its SL; median line of
back in front of dorsal fin with several short spines.
Pristigaster Cuvier 1817, p. 427.
14b. Body elongate, with chest and abdomen not greatly
dilated, its greatest depth less than a third of its
SL; no external spines on median line of back.
15a. Body excessively compressed; dorsal outline
more or less concave over the head; a tooth-
less membranous section between maxillary
and premaxillary, the three elements form-
ing a continuous margin; maxillary greatly
produced in large adults (Fig. 111); anal fin
with about 52-80 rays.
Odontognathus Lacépéde 1800, p. 430.
15b. Body rather strongly but not excessively
262 Memoir Sears Foundation for Marine Research
compressed; dorsal outline convex over head;
maxillary and premaxillary not separated by
a membranous section; maxillary overlapping
distal end of premaxillary, interrupting mar-
gin of upper jaw (Fig. 113); anal fin with
only about 39-48 rays.
Neoopisthopterus Hildebrand 1948, p. 435.
11b. Teeth in jaws of unequal size, both jaws with strong canines (Fig.
115); vomer with teeth. Subfamily Chirocentrinae.
Chirocentrodon Ginther 1868, p. 438.
3b. Median line of back in front of dorsal fin naked, not crossed by scales; stomach
with very thick walls, like the gizzard of a fowl. Subfamily Dorosomatinae.
Dorosoma Rafinesque 1820, p. 443.
Genus Etrumeus Bleeker 1853
Round Herrings
Etrumeus Bleeker, Verh. Batav. Genoot., 25, 1853: 48; genotype by monoytpy, C/upea micropus Temminck
and Schlegel.
Generic Synonym:
Perkinsia Rosa Eigenmann, Amer. Nat., 25, 1891: 153; type species by monotypy, P. othonops Rosa Eigenmann
equals C/upea micropus ‘Temminck and Schlegel.
Characters. Bovy elongate, little compressed, especially in Atlantic species; chest
and abdomen fully rounded. Bony scures absent. ScaLes with smooth edges, deciduous,
covering chest and abdomen. Snovut long, tapering. Mourn terminal. Maxriiary
slender, ending about under anterior margin of eye. TEETH minute, present on jaws,
vomer, palatines, pterygoids, and tongue. Dorsat fin beginning notably nearer to
margin of snout than to base of caudal, with about 17-20 rays. Anat about half as
far from base of caudal as from origin of dorsal, with about 10-12 rays. Petvic fins
inserted well behind base of dorsal, with a shield of enlarged scales covering them in
part, each fin with 8 rays. Pecrorats of moderate size, also shielded in part by enlarged
scales, each fin with about 14-16 rays. Pelvic and pectoral each with a long Axit-
LARY PROCESS.
Range. Represented by one species on the Atlantic and Gulf coasts of the United
States, and by two or more species in the eastern and western Pacific on the coasts of
southeastern Africa and southeastern Australia.
Fishes of the Western North Atlantic 263
Etrumeus sadina Mitchill 1814
Atlantic Round Herring
Figures 62, 63
Study Material. Many specimens, postlarvae to adults, 25-180mm TL, the
specimens from New Jersey northward being adults caught near or on the shore,
those from off Cape Lookout, northern Florida, and the Gulf of Mexico, mostly young,
taken quite a distance offshore in 5-21 fms. depth except for a lot of six small ones
from Beaufort Inlet, North Carolina, taken in water only about 2-3 fms. deep.
Ficure 62. Etrumeus sadina, female, 140 mm TL, 115 mm SL, Woods Hole, Massachusetts, USNM 143574.
Drawn by Ann S. Green.
Distinctive Characters. The very slender and roundish body, the roundish chest
and abdomen covered with ordinary scales, the absence of bony scutes, the small anal
fin situated near the base of the caudal, and the rather small pelvic fin inserted entirely
behind a vertical from the base of the last dorsal ray, readily distinguish this species
from other American clupeoids.
Description. Proportional dimensions in per cent of standard length, and counts,
based on 22 specimens from Study Material, 55 mm (young adults)-180 mm TL,
45-154 mm SL.
Body: depth 16-18.5. Distance from snout to origin of: dorsal
Caudal peduncle: depth 6.7-8.2. 42-46.
Head: length 24-29.
Snout: length 6.0-9.5. Scales: about 48-55, usually lost in pre-
Eye: diameter 6.5—8.0. served specimens.
Interorbital: width 3.0-4.4. Gill rakers: 32-38 on lower limb, ca. 14
Maxillary: length 7.0-10.5. on upper limb.
Anal fin: length of base 5.6—6.9. Fin rays: dorsal 16-20, usually 18 or 19;
Pelvic fin: length 8.9-11.5. anal 10-12, most frequently 11;
Pectoral fin: length 14-17. pectoral 14-16.
Vertebrae: 49 (7 specimens).
264 Memoir Sears Foundation for Marine Research
Bopy almost round, its greatest thickness about 80°/, of its depth, its greatest
depth 5.4-6.25 in SL; ventral profile a little more strongly convex than dorsal pro-
file. CAUDAL PEDUNCLE compressed, its depth 3.3-3.9 in head.
Scates deciduous, usually lost in preserved specimens, very thin, scarcely deeper
than long, with even edges.
Heap low, rather long, 3.45—4.15 in SL. Snour equal to or more usually longer
than eye, tapering, 3.0-3.9 in head. Eye with much adipose tissue in adults, 3.2 5—4.1
in head. Inrerorpirat bone 6.25—7.55. Maxitiary obliquely rounded posteriorly,
reaching about under anterior margin of eye, 2.6—3.0 in head. Manprsie coterminal
with snout, 1.85—-2.0 in head. Grit rakers rather slender, moderately close-set, those
at angle somewhat exceeding half of length of eye; apparently not increasing in number
with age. TrrrH small, in a single series on mandible; a few minute teeth on pre-
maxillaries, and a row on the margin of maxillaries; very small granular teeth on
vomer, palatines, pterygoids, and tongue.‘
Dorsat fin moderately elevated anteriorly, with concave margin, its longest rays
generally not reaching tip of posterior ray if deflexed, its origin about equidistant be-
tween margin of snout and vertical from middle of anal base, its distance from snout
2.15—2.4 in SL. Caupat fin forked, the lobes of about equal length, generally somewhat
longer than head without snout. Anat small, rather feebly developed, placed far behind
dorsal fin, a little nearer to base of caudal than to base of pelvic, its base 3.5—4.5 in
head. Prtvic fin rather short and broad, with nearly straight margin, inserted a little
behind vertical from base of last ray of dorsal, generally a little nearer to base of pec-
toral than to base of caudal, 2.3-3.0 in head; a membranous axillary process present,
about 67°/, of the length of fin. Pecrorat fin moderately developed, reaching notably
less than halfway to base of pelvic, inserted fully an eye’s diameter nearer to tip of
mandible than to base of pelvic, 1.5—1.8 in head; a long membranous axillary process
present about 0.83 of length of fin.
ALIMENTARY CANAL about half of SL. Sromacu with a long blind sac and nu-
merous pyloric caeca. AIR BLADDER long and narrow.
Color. Fresh specimens olive green above, silvery on lower half of side and below.
In old preserved specimens, back generally brownish and lower parts pale (sometimes
silvery).
Size. A length of 200-250 mm (8-10 in.) is said to be attained by this species.
Development. The postlarvae and young adults are very slender, as is usual in
young herrings. In postlarvae 25-28 mm TL (21-24 mm SL) the body depth is
contained about 16-24 times in the SL; the jaws are weakly developed; the teeth
are not evident; the snout is proportionately longer than that in adults, being 1.5—
2.0 times longer than the eye; the pectoral fins are not much more than tufts of mem-
brane; and the pelvics are incompletely developed. The larval pigment spots are still
retained, chiefly in a single slightly broken dark line on the chest, in two diverging
dark lines on the abdomen, in a row of dark spots along each side of the base of
4. To see the teeth clearly it may be necessary to dry the head.
1 WESC AStSS : —————
SS SS
Ficure 63. Etrumeus sadina. a, larva, 27.5 mm TL, near Pensacola, Florida. 8, young, 35 mm TL, near
Pensacola, Florida. c, young, 42.5 mm TL, Beaufort, North Carolina, USNM 143804. All drawn by Ann
S. Green; a and B based on camera-lucida sketches by W. W. Welsh.
anal, and in a single dark median line behind the anal fin; also several dark spots are
present on the base of the caudal.
At about 33-35 mm TL, the fins are all rather fully developed and the body
has become proportionately much deeper, but the pigmentation remains virtually the
same as in smaller examples. Specimens of this size, then, may be classed as young
adults. General pigmentation is delayed and is scarcely complete at 55 mm TL. The
fish continue to increase in relative depth as long as they grow.
Spawning. The smallest specimens at hand, postlarvae 25-28 mm TL, are so
young that they probably were caught not far from where they were hatched; they were
taken at 30°29’ N, 80°37.5’W, off northern Florida. Still smaller larvae, 7.6—21mm,
were listed by the late W. W. Welsh in his unpublished notes, accompanied by seven
camera-lucida sketches. These were taken by the Grampus at the surface in the Gulf
of Mexico at 28°21’N, 89°06’ W, 17 miles or so off the Mississippi Delta. Although
these specimens are apparently no longer extant, the drawings leave no room for doubt
as to their identity. Also in this as well as other collections there are young adults from
northern Florida south to Cape Canaveral and from the Gulf of Mexico off Mobile
Bay. The smallest young (including postlarvae) were taken on January 25 (1940), and
266 Memoir Sears Foundation for Marine Research
other small young adults were taken in March off northern Florida and in the Gulf of
Mexico. The smallest young were caught in water 19 fms. deep, and other small
ones were taken at 21, 20, 16, and 5 fms. From these data it may be concluded that
spawning in the south takes place during the winter, somewhat offshore, and in water
several fathoms deep. If spawning takes place northward it seems not to have been
reported.
Migration and Habitat. In the northern part of its range, as at Woods Hole, Massa-
chusetts, this Round Herring is taken along the shore only during summer. It has not
yet been determined whether this fish migrates northward during summer or whether
it only comes inshore from deep water during warm weather. In southern waters it
is not known to occur along the shore, for there it has been taken only offshore in
water a few to several fathoms in depth. Whether it migrates at all in these waters re-
mains unknown.
Relationships. E. sadina is the only representative of the genus in the Atlantic, but
other species occur in the Pacific: off California, the Philippine Islands, Japan, Hawaii,
and Galapagos Islands. Those from the western Pacific are more compressed and pro-
portionately deeper than the Atlantic species and are therefore less typically “‘round
herrings.”
Abundance. Generally this Round Herring is rare to scarce from New Jersey to
New England, where it occurs along the shore, but occasionally it is common to
plentiful. It was very abundant off Woods Hole, Massachusetts, in 1905 and again in
1908 (123: 741). A number of specimens was also taken at Campobello Island at the
mouth of the Bay in September 1937 (82: 5; 92: 15). Along the coast of eastern Maine,
in August-September 1953, something like 200,000 pounds were landed (174: 194).
But apparently it has been taken only sparingly off the southern Atlantic and Gulf
coast states, though commonly regarded as a warm-water fish.
Range. It is known from Passamaquoddy Bay at the mouth of the Bay of Fundy
and from the eastern part of Maine to northern Florida off Cape Canaveral, and from
the Gulf of Mexico.’ Although it has been reported as not rare southward (72,
1896: 420), no records have been found of its occurrence from Delaware southward;
and there is only one record for the Gulf, the latter being based on specimens taken
from the stomachs of snappers landed at Pensacola, Florida (74: 143). Recent col-
lections by the U.S. Fish and Wildlife Service include: examples from Beaufort Inlet
and off Cape Lookout, North Carolina; four collections of young from off northern
Florida southward to Cape Canaveral; and six collections from the Gulf of Mexico,
five of these from off Mobile Bay and one from south of New Orleans. These rather
recent collections are all from offshore waters of 2-21 fms. depth. Those from New
Jersey northward, all adults, were caught in shallow water near or on the shore.
5. Bertin (r3:9) synonymized £. microps and E. jacksoniensis, previously recognized as valid species, and thereupon
gave the range as the Atlantic coast of the United States, the coast of southeastern Australia, southeastern Japan,
Hawaii, and the southeastern coast of Africa. However, from comparison of specimens from the United States and
Japan, it was at once evident that the latter were different in shape, being especially more compressed. A further
study of specimens from the widely separated regions where the genus occurs would be desirable.
Fishes of the Western North Atlantic 267
Apparently this fish does not occur off southern Florida, for it was not reported
by Longley and Hildebrand from collections made at Tortugas over many years, in-
cluding dredgings at various depths down to about 400 fms (87). This seems to be
another instance of a discontinuous range of the genus such as is reported for Pomo/obus,
Alosa, and Brevoortia (p. 343). However, in Ezrumeus the inhabitants of the Gulf and
the Atlantic appear to be identical, whereas in the other genera named, the inhabitants
of the two areas differ sufficiently to be recognized as distinct species.
Synonyms and References:
Clupea sadina Mitchill, Rep. in part on Fishes of New York, 1814: 21 (orig. descr.; type local. presumably
New York; type lost); Mitchill, Trans. Lit. philos. Soc. N. Y., 7, 1815: 4.57 (descr., New York®).
Alosa teres DeKay, N.Y. Fauna, Fishes, Pt. 4, 1842: 262, pl. 40, fig. 128 (orig. descr.; type local. New York
Harbor; type in State Coll.).
Etrumeus teres Giinther, Cat. Fish. Brit. Mus., 7, 1868: 467 (synon., notes, range); Jordan and Gilbert, Bull.
U.S. nat. Mus., 16, 1883: 263 (diagn., refs.); Jordan and Gilbert, Proc. U.S. nat. Mus. (1883), 6,
1884: 143 (from stomachs of snappers, Pensacola, Florida); Bean, T.H., Bull. U.S. Fish Comm.
(1887), 7, 1888: 148 (Great Egg Harbor Bay, New Jersey); Bean, T. H., Bull. N. Y. St. Mus., 60,
Zool. 9, 1903: 189 (synon., descr., New York); Fowler, Rep. N. J. St. Mus. (1905), 2, 1906: 93
(diagn., synon.); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 36 (refs., New England);
Sumner, Osburn, and Cole, Bull. U.S. Bur. Fish. (1911), 32 (2), 1913: 741 (refs., parasites, Woods
Hole, Massachusetts); Fowler, Proc. biol. Soc. Wash., 33, 1920: 147 (New Jersey); Bigelow and Welsh,
Bull. U.S. Bur. Fish. (1924), 40 (1), 1925: 91, fig. 39 (descr., range, Gulf of Maine); Breder, Field
Bk. Mar. Fish. Atl. Cst., 1929: 63, fig. (diagn., distr.).
Etrumeus sadina Jordan and Evermann, Bull. U. S. nat. Mus., 47 (1), 1896: 420 (descr., range, synon.); Jordan,
Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 46 (synon., range); Bigelow and Schroe-
der, Bull. U.S. Bur. Fish., 48 (20), 1936: 327 (Yarmouth R. and Casco Bay, Maine); Bertin, Bull.
Inst. océanogr. Monaco, 853, 1943: 9, fig. 4 (synon., descr., distr., etc.); Bigelow and Schroeder, Fish.
Bull. (74) U.S. Fish Wildl. Serv., 53, 1953: 87 (Gulf of Maine).
Genus Fenkinsia Jordan and Evermann 1896
Dwarf Herrings
Fenkinsia Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 418; genotype by monotypy, Dussumieria
stolifera Jordan and Gilbert equals C/upea lamprotaenia Gosse.
Characters.? A very small species. Bopy slender, compressed; abdomen rounded;
chest with suggestion of a keel. Bony scutes absent. Scares deciduous, very thin,
with smooth edges, covering abdomen. Snour moderately long, tapering. Mourn
terminal. Maxritary narrowly rounded, terminating about under anterior margin of
pupil. TrerH in a single series on the margin of maxillary and premaxillary; a short
6. Because of such inaccurate statements in the original description as “Mouth. . .toothless,” “Lower jaw projecting,”
and “A. 15,” it was long questioned whether sadina was available for this species. However, the description con-
tains pertinent remarks such as “Form neat, tapering, and slender,” and especially “the belly is not at all serrate,
but quite smooth,” which can scarcely apply to any other clupeoid. Therefore, sadina certainly seems available.
Apparently Alosa teres DeKay, though much better described and rather well figured, must rest in synonymy,
under the rules of zoological nomenclature.
7. Bertin (r3:17) has stated that enkinsia is not separable from Spratelloides Bleeker.—G. S. Myers.
268 Memoir Sears Foundation jor Marine Research
toothless membranous section present between these elements; no teeth on mandible or
vomer; at most a few granular teeth on palatines, pterygoids, and tongue. Dorsat small,
beginning rather nearer to margin of snout than to base of caudal, with about 9-14 rays.
Anat only a little larger than dorsal and placed far behind it, with about 12-16 rays.
Petvic fins nearly as long as the pectorals, each with 8 rays; a scale process present
between fins at base. Pecrorat fins rather small, each with about 10-14 rays. Pelvic
and pectoral both without a free AxILLARY PROCESS.
Remarks. Fenkinsia resembles Etrumeus in having a rather roundish, elongated body,
a rounded abdomen, and no bony scutes on the ventral outline of the chest and abdomen.
However, in Yenkinsia the chest is very slightly keeled. The rather small anal fin, situated
far behind the dorsal, and the rather large pelvic fin, inserted under the anterior half of
the base of dorsal, also help to distinguish this genus. The small size of these herrings,
about 50-70 mm, also separates them from other herrings.
Range. Known from Bermuda, doubtfully from North Carolina, the Florida Keys,
the West Indies, and from some small islands in the Caribbean Sea, off México and
Central America. Only two species are known.
Key to Species
1a. Gill rakers not especially close-set, 19-24 (usually 20 or 21) on lower limb; oblique
series of scales crossing silvery lateral band between margin of opercle and base
of caudal, 33-37. lamprotaenia (Gosse) 1851, below.
1b. Gill rakers close-set, 27—30 on lower limb; oblique series of scales crossing silvery
lateral band between margin of opercle and base of caudal, 39-42.
viridis (T. H. Bean) 1912, p. 272.
Fenkinsia lamprotaenia (Gosse) 1851
Dwarf Herring, Green Fry
Figure 64
Study Material. Many specimens, 28-65 mm TL, 23-55 mm SL, collected at
Key West and Tortugas, Florida, the Bahama Islands, Cuba, Isle of Pines, Jamaica,
Haiti, Puerto Rico, St. Eustatius Island, Old Providence Island, and Yucatan Island
off Cozumel, México.
Distinctive Characters. See Remarks, and Key to Species, above.
Description. Proportional dimensions in per cent of standard length, and counts,
based on 16 specimens, 40-65 mm TL, 32-55 mm SL.
Body: depth 15-20. Snout: length 6.2—8.0.
Caudal peduncle: depth 6.5-8.5. Eye: diameter 7.6—9.7.
Head: length 25-29. Interorbital: width 3.8—5.4.
Fishes of the Western North Atlantic 269
Maxillary: length 9.0-12. Scales: 33-37, generally missing in pre-
Mandible: length 12-14.5. served specimens.
Anal fin: length of base 10-14. Gill rakers: 19-24 (usually 20 or 21) on
Pelvic fin: length 11-15. lower limb.
Pectoral fin: length 14-17. Fin rays: dorsal 9-133; anal 12-15; pec-
Distance from snout to origin of: dorsal toral 10-13.
44-50. Vertebrae: 39 or 40 (6 specimens).
Ficure 64. Fenkinsia lamprotaenia, 65mm TL, 53 mm SL, Haiti, USNM 89600. Drawn by Ann S. Green.
Bopy moderately compressed, its greatest thickness somewhat greater than half of
its depth, its greatest depth 5.0-6.65 in SL; ventral outline more strongly convex
than dorsal outline. CaupaL PEDUNCLE rather strongly compressed, its depth 3.2—-4.1
in head.
Scates generally lost in preserved specimens, very thin, with smooth edges; those
from middle of side below dorsal fin with a wavy ridge on anterior half, about 5 wavy
radii behind it, and generally several shorter ridges at margin of scale.
Heap rather low, 3.4-4.0 in SL. Snour generally shorter than eye, 3.4—4.2 in
head. Eve 2.8—3.4. INTERORBITAL §.5—7.0. MaxiLtary narrowly rounded posteriorly,
reaching to or more usually a little beyond vertical from anterior margin of pupil,
2.25-2.75 in head. Manopiste generally included, not projecting, 1.9-2.2 in head.
GILL RAKERS moderately close-set, those at angle of first arch about 75°/, of length
of eye. TEETH missing on mandible and vomer but present in a single series on
margin of premaxillary and maxillary, the series interrupted by a soft membranous
section between these elements; palatines, pterygoids, and tongue at most with a few
granular teeth.
Dorsat fin moderately elevated, the margin nearly straight, the longest rays reach-
ing well beyond the tip of last ray if deflexed, the fin beginning notably nearer to margin
of snout than to base of caudal, the distance from margin of snout 2.0—2.3 in SL. Caupat
fin moderately forked, the lobes of about equal length, rather longer than head without
snout; each lobe with a greatly elongated scale at base. Anat fin much lower than dorsal,
beginning far behind dorsal fin, about equidistant between insertion of pelvic and base
270 Memoir Sears Foundation for Marine Research
of caudal, its base 1.9—2.6 in head. Petvic fins only a little shorter than pectorals, with
a large scaly process between them, the fins inserted under about 4th—6th dorsal ray,
and generally close to an eye’s diameter nearer to base of caudal than to tip of mandible;
1.9—2.5 in head. Pecrorat fins somewhat pointed, inserted about equidistant between
tip of mandible and base of pelvics, 1.6—1.9 in head.
ALIMENTARY CANAL short. StomacH with a large blind sac. PyLoric cozca large,
about 7. PeR1IToNEum pale, with many dark dots.
Color. In life, faintly greenish above, with a distinct silvery lateral band. Preserved
specimens straw colored, the silvery band persisting. Margin of mandible brown to
black. Well-preserved specimens with an elongated silvery area on abdomen behind
pectoral fin, and a similar though smaller area behind pelvic. Scales above lateral
band with a brownish line on middle of exposed section, paralleling the curve of the
margin of the scale; dark dots also present, especially in and above upper margin of
silvery lateral band; two rows of dark dots on back; generally numerous dots on snout,
including an elongated one in front of lower half of eye; an irregular dark spot at base
of each anal ray, continued as a single median ventral line on caudal peduncle.
Size. The largest example at hand, 65 mm (2.6 in.) TL, may be near the maximum
size attained.
Spawning and Development. Nothing is known definitely concerning reproduction
in this species, but examples collected in 1937 at Kingston, Jamaica, during our winter
months, contain well-developed eggs, indicating that at least some spawning takes place
during that time around Jamaica.
The smallest examples at hand, a little more than an inch (28-30 mm) TL, are
fully developed young adults. They differ principally from large examples from the
same general locality (Tortugas, Florida) in being more slender, the depth being about
6.4 times in SL; in the large adult this proportion varies between about 2.25—5.6.
However, there are large specimens in the collections studied that are just as slender
as the young mentioned. For further discussion of the difference in depth, see Varia-
tions (p. 271).
Migration and Habitat. It seems improbable that this little fish travels long dis-
tances. It does occur in large compact schools that probably consist of millions of fish,
as observed by me at Key West, Florida, and as reported for Tortugas, Florida (49:
156; 2I: 1). It moves about more or less like the young Atlantic menhaden, Brevoortia
tyrannus, without pursuing any definite course.
Food. This species, like many other clupeoids, feeds on plankton, which it screens
from the water as the school girates or mills around at the surface.
Enemies. Many large fish feed on this species. The gray snapper was mentioned
especially by Gudger, who said that the little fish exhibited little fear; it hovered over
the snapper in dense swarms but left a clear space around it approximately equal to the
length of the snapper (49: 156). Whenever the snapper moved, the rather orderly school
broke up but soon re-formed. Longley said, ‘“Exceedingly common at Tortugas (Fla.),
particularly along shore, where predacious fishes often drive them too close for safety,
Fishes of the Western North Atlantic 27
and large numbers are cast upon the windward beaches to spring about until a wave
rescues them, or until they perish” (87: 12).
The copepod, Colliparvus parvus Wilson, was found on the gills of specimens
taken at Nassau, Bahamas (135: 79), and an unidentified trematode was reported as
parasitic on it at Tortugas, Florida (go: 372).
Relationships. Only two species of Fenkinsia are recognized herein. A third nominal
species, stolifera, long was thought to be distinct, but recent works on taxonomy agree
that it and /amprotaenia constitute one variable species; since /amprotaenia is the older
name, stolifera has become a synonym. 7. viridis is closely related to /amprotaenia, as
shown in-the account of viridis (p. 272).
Variations. This species, as indicated, is very variable. While the depth of body
in general increases with age, adult specimens of equal size vary greatly in this respect
among themselves, as is evident from the wide range shown in the Description. And
this in turn is reflected in the wide range in the depth of the caudal peduncle and in
the length of the head. A rather wide variation in the number of dorsal, anal, and pectoral
rays, and in the number of gill rakers, is also shown in the Description. Although some
of the earlier counts may have erred because fins are often broken in preserved specimens
(no doubt so in the original description of sto/ifera), this error does not enter into those
given in the foregoing Description, for ample specimens with undamaged fins were
chosen for the enumerations.
Commercial Importance. Although this little Herring has been reported among the
“white bait” in the market at Port-au-Prince, Haiti (ro: 45), no doubt its chief value
is that of providing food for foodfishes.
Range. Known from the Florida Keys, the Bahamas, throughout the West Indies,
and from Swan Island, Glover Reef, and Cozumel Island in the Caribbean Sea; recently
reported from Venezuela; doubtfully recorded for Beaufort, North Carolina. Appar-
ently it is chiefly a shore species.
Synonyms and References:
Clupea lamprotaenia Gosse, Naturalist’s Sojourn in Jamaica, 1851: 291, pl. 1, fig. 2 (orig. descr.; type local.
Jamaica; types in BMNH; found in schools with an anchovy, Emgraulis brownii = Anchoa hepsetus);
Giinther, Cat. Fish. Brit. Mus., 7, 1868: 465 (descr., Jamaica).
Dussumieria stolifera Jordan and Gilbert, Proc. U. S. nat. Mus. (1884), 7, 1885: 25 (orig. descr.; type local.
Key West, Florida; type USNM 34964; local abund.); Bean, T. H., Bull. U.S. Fish. Comm. (1888),
&, 1890: 206 (Cozumel, Yucatdn).
Fenkinsia lamprotaenia Jordan and Evermann, Bull. U.S.nat. Mus., 47 (1), 1896: 419 (descr., after Giinther);
Evermann and Marsh, Bull. U.S. Fish Comm. (1900), 20 (1), 1902: 84 (descr., refs., Puerto Rico);
Metzelaar, Rapp. Vissch. Curacao, ed. by J. Boeke, 1, W. Indian Fishes, 1919: 9, fig. (refs., descr.,
range; Leeward Is., West Indies); Beebe and Tee-Van, Zoologica, N. Y., 10 (1), 1928: 43, fig. (refs.,
descr., range, F. stolifera synon.; Port-au-Prince, Haiti); Breder, Field Bk. Mar. Fish. Atl. Cst., 1929:
63, fig. (diagn., range); Nichols, N.Y. Acad. Sci., Zo (2), 1929: 201, fig. 32 (refs., distr., diagn.,
Puerto Rico); Parr, Bull. Bingham oceanogr. Coll., 3 (4), 1930: 3 (refs.; discus. ident. of /amprotaenia
and stolifera; Bahamas, Turks Is.); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2,
1930: 46 (range, refs.); Longley and Hildebrand, Publ. Carneg. Instn. Wash., 535, 1941: 12 (discus.
lamprotaenia and stolifera; variability; Tortugas, Florida); Hildebrand, Copeia, 1941: 224 (extension
of range, Beaufort, North Carolina); Fowler, Monogr. Acad. nat. Sci. Philad., 6, 1944: 123, fig. 30
292 Memoir Sears Foundation for Marine Research
(descr., locals., pp. 434, 456); Martin, Mem. Soc. Cient. La Salle, 75, 1955: 184-188 (descr., ills.,
Venezuela).
Fenkinsia stolifera Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 419 (descr., ref. to orig. descr.) ;
Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 55 (refs., Florida); Evermann and
Marsh, Bull. U.S. Fish Comm. (1900), 20 (1), 1902: 84 (descr., refs., Puerto Rico); Bean, B. A. ix
Bahama Islands, Fishes, 1905: 297 (rec. from Spanish Wells); Fowler, Proc. Acad. nat. Sci. Philad.,
58, 1906: 81, fig. 1 (descr.; assoc. with anchovies; Florida Keys); Wilson, Proc. U.S. nat. Mus., 53,
1917: 79 (host of copepod, Colliparvus parvus Wilson; Nassau, Bahamas); Breder, Bull. Bingham oceanogr.
Coll., 7 (1), 1927: 13 (Isle of Pines, Swan I., and Glover Reef off British Honduras); Breder, Amer.
Mus. Novit., 382, 1929: 1-5 (school. behav., Tortugas, Florida); Gudger, Publ. Carneg. Instn. Wash.,
391, 1929: 156, pl. 1, fig. 6 (local abund., habits, Tortugas, Florida); Jordan, Evermann, and Clark,
Rep. U.S. Comm. Fish. (1928), 2, 1930: 46 (range, ref.); Breder, Publ. Carneg. Instn. Wash., 435,
1932: 8 (fed on by needlefish, Strongy/ura; Tortugas, Florida); Gowanloch, Bull. La. Conserv. Dep.,
23, 1933: 214 (in great schools, Gulf of Mexico; but no records for Gulf coast north of extremes
Florida, therefore, apparently not of Louisiana fauna).
Spratelloides lamprotaenia Bertin, Bull. Inst. océanogr. Monaco, 853, 1943: 17 (diagn., revision).
False References:
Fenkinsia stolifera Nichols (not of Jordan and Gilbert), Proc. biol. Soc. Wash., 33, 1920: 63 (Stolephorus
viridis in synon., Bermuda).
Fenkinsia lamprotaenia Beebe and Tee-Van (not of Gosse), Field Bk. Shore Fish. Bermuda, 1933: 37, fig.
(field char., diagn., distr.); Beebe and Tee-Van, Zoologica, N. Y., 73 (7), 1933: 136 (refs., Sto/ephorus
viridis and F. stolifera in synon., Bermuda); Hollister (not of Gosse), Zoologica, N. Y., 2I (4), 1936:
276, figs. 40-44 (caudal skel.).
Fenkinsia viridis (T. H. Bean) 1912
Bermuda Herring, Dwarf Herring, Green Fry
Figure 65
Study Material. A total of 19 specimens, 42-58 mm TL, 32-48 mm SL, as fol-
lows: 4 “‘types,” from Bermuda, presumably from Hamilton, USNM 74084; the holo-
type, USNM 100546; and 14 paratypes of Fenkinsia bermudana Rivas, Grampus St.
10178, off Bermuda, 32°20'N, 64°21’ W.
Distinctive Characters. See Relationships (p. 273) and Key to Species (p. 268).
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens, 32-48 mm SL.
Body: depth 15-17.5. Pectoral fin: length 14-16.
Caudal peduncle: depth 6.7—-8.2. Distance from snout to origin of: dorsal
Head: length 25-27. 44.5—§0.
Snout: length 6.7-8.5.
Eye: diameter 7.0-9.0. Scales: about 39-42, generally lost in
Interorbital: width 3.6—4.75. preserved specimens.
Maxillary: length 9.4-10.8. Gill rakers: 27-30 on lower limb.
Mandible: length 11.6-13.5. Fin rays: dorsal 9-143; anal 14-16; pec-
Anal fin: \ength of base 12-14. toral 12-14.
Pelvic fin: length 12-15. Vertebrae: 40 (I specimen).
Fishes of the Western North Atlantic 278
Bopy shaped in all respects as in /amprotaenia, its greatest depth 5.7—6.6 in SL.
CAUDAL PEDUNCLE 3.2—4.0 in head.
SCALEs as in /amprotaenia, except that there appear to be a few more in a lateral
series.
Heap low, with exceedingly numerous mucous canals and pores, its length 3.7—
4.0 in SL. Snour equal to, or more usually a little shorter than, eye, 3.2—3.8 in head.
Eye 2.9—3.5. INrERORBITAL 5.5—7.1. MaxitLary narrowly rounded posteriorly, gen-
erally reaching vertical from anterior margin of pupil, 2.45-2.6 in head. ManpIBLeE
slightly included, about coterminal with the snout, 2.0—2.2 in head. Git RAKERs very
Ficure 65. Fenkinsia viridis, 45 mm TL, from Bermuda, USNM 100546. After Rivas.
close-set, those at angle rather more than half of length of eye. Teeru apparently in
all respects as in /amprotaenia.
Dorsat fin elevated anteriorly, the longest rays reaching far beyond tip of last
ray if deflexed, the fin generally beginning about half of an eye’s diameter nearer to
margin of snout than to base of caudal, the distance from margin of snout 2.0—2.25 in
SL. Caupat fin as in /amprotaenia, Anat lower than dorsal, its origin about equi-
distant between insertion of pelvic and base of caudal, its base 1.8-2.2 in head. Petvic
fin only a little shorter than pectoral, inserted under the 4th or 5th dorsal ray and
about equidistant between anterior margin of eye and base of caudal, its length 1.5 5—
2.2 in head. Pecrorat fin pointed, inserted a little nearer to tip of mandible than to
base of pelvic fin, 1.55—1.8 in head.
Color. In all respects as in /amprotaenia.
Size. The largest specimens at hand, 55-58 mm TL (2.2-2.3 in.), may be near
the usual maximum length attained.
Food. Although no specific information is at hand, there seems to be no reason to
believe that the food of this species differs from that of /amprotaenia.
Enemies. No enemies have been listed. However, there can be no doubt that
predatory fish and water birds feed on this species as they do on /amprotaenia.
Relationships. As stated in the account of /amprotaenia, the two species of this
18
2a Memoir Sears Foundation for Marine Research
genus recognized herein are closely related. But they differ in the number of gill rakers
on the lower limb of the first arch, viridis having 27-30 while /amprotaenia has only
19-24 (usually 20 or 21). That there is a difference between the species in the inser-
tion of the pelvic fin, as stated by Rivas (r08: 3), I am unable to substantiate, as it
seems to be obliterated by the variations among specimens. However, according to four
viridis and seven /amprotaenia that remain almost fully covered with scales, in the former
39-41 oblique rows of scales cross the lateral band between the margin of the opercle
and the base of the caudal, whereas in the latter only 33-37 series cross this band.
The specimens of viridis at hand have much more prominent mucous canals and pores
about the head and on the anterior part of the body, but they may not be specifically
significant.
It so happens, then, that even though T. H. Bean placed this species in the
wrong genus and family (Sto/ephorus, family Engraulidae), his species still remains
valid (9: 22).
Commercial Importance. This species is not known to be of direct commercial
importance. It has been reported as exceedingly common in Bermuda (9: 123; II: 37),
therefore it must be valuable as a forage fish.
Range. Known only from Bermuda and vicinity. It congregates in schools like
/amprotaenia but ventures some distance offshore, as shown by the Grampus catch noted
above.
Synonyms and References:
Stolephorus viridis Bean, T.H., Proc. biol. Soc. Wash., 25, 1912: 122 (orig. descr.; type local. Bermuda;
types USNM 74084; common).
Fenkinsia stolifera Nichols (not of Jordan and Gilbert), Proc. biol. Soc. Wash., 33, 1920: 63 (Stolephorus
viridis 1M synon.).
‘Fenkinsia lamprotaenia Beebe and ‘Tee-Van (not of Gosse), Field Bk. Shore Fish. Bermuda, 1933: 37, fig.
(field char., diagn., distr.); Beebe and T’ee-Van, Zoologica, N. Y., 13 (7), 1933: 136 (refs., Stolephorus
viridis and F. stolifera in synon.; Bermuda); Hollister, Zoologica, N. Y., 2I (4), 1936: 276, figs. 40-44
(caudal skel.).
Fenkinsia bermudana Rivas, Smithson. misc. Coll., 106 (14), 1946: 1-4, 1 fig., 1 pl. (orig. descr.; type local.
Bermuda; type USNM 100546; cf. /amprotaenia; but 8. viridis Bean apparently overlooked, of which
Lermudana becomes a synon.).
Genus C/upea Linnaeus 1758
Clupea Linnaeus, Syst. Nat., ed. 10, 1758: 317; type species by implication (unquestioned), C/upea harengus
Linnaeus.
Generic Synonym:
Rogenia Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 340; type species by monotypy, Rogenia alba
equals C/upea harengus Linnaeus 1758.
Characters. Bopy quite elongate, compressed. Bony scurgs on median line of
chest and abdomen, weak. ScaLes with crenulate membranous borders. MaxiLlary
with a large paddle-shaped supplemental bone. TrETH on vomer permanent, forming
Fishes of the Western North Atlantic 275
a small patch. Dorsat fin rather small, without a produced ray. Petvic commonly
with 9 rays, rarely with only 8.8 VERTEBRAE 45-57.
Remarks. The very slender body and the presence of vomerine teeth distinguish
this genus from other genera of the western North Atlantic.
Range. North Atlantic and North Pacific, ranging on the American side of the
Atlantic from North Carolina to Greenland and in Europe from the Strait of Gibraltar
to Spitsbergen. In the Pacific, the genus is represented from southern California to the
Aleutian Islands and across to Siberia and Japan. A single species occurs in the western
North Atlantic.® The species generally occurs in schools, which are usually seen at or
rather near the shore.
Clupea harengus Linnaeus 1758
Atlantic Herring, Sea Herring, Herring
Figures 66—68
Study Material. At least 18 specimens, 75-270 mm TL, 63-224 mm SL, and
smaller specimens, including postlarvae; from Kings Creek (tributary to Tangier
Sound), Chesapeake Bay, Maryland; Ipswich River (estuary) and Woods Hole, Mas-
sachusetts; Sheepscot River and Eastport, Maine; St. Lawrence River; Nova Scotia;
New Brunswick; and Newfoundland. This American material was compared with
specimens from Norway and Sweden, and with specimens from the Pacific collected in
California, Oregon, and Alaska.
Distinctive Characters. The following conspicuous field marks separate the post-
larval and adult Atlantic Herring from the shads (4/osa) and from representatives of
Pomolobus (alewives, hickory shad): the point of origin of its dorsal fin is about midway
of its trunk (considerably farther forward in the others); its body is not as deep, a differ-
ence shown better in the illustrations than by words; and the sharp midline of its belly
is less strongly serrate.
[Critical examination is required to distinguish the early stages of clupeoids of the
western North Atlantic, one from another. Larvae of other species with which the
Herring larva might be confused are the launce (¢mmodytes), the so-called rock gunnel
(Pholis), the capelin (Ma/lotus), and the smelt (Osmerus). But the position of the anus,
about 83°/, of the way back along the body, sets the larval Herring apart from all
of these (about 75 °/, of the way back in Ma//otus and Osmerus, about 66 °/, of the
way back in Ammodytes and Pholis).—n. B. B.]
Description. Proportional dimensions in per cent of standard length, and counts,
from specimens in Study Marerial, not including small specimens except for men-
tion of their much more slender body and fewer gill rakers; specimens 63-224 mm SL.
8. Pelvics with 7 or 8 in South American species.—G. S. Myers.
g. The genus Clupea in the present sense is also represented in Australian-New Zealand temperate waters and in
southern South America. Norman has reviewed the South American forms (102: 37).—G. S. Myers.
18*
276 Memoir Sears Foundation for Marine Research
Body: depth 20-2 5.8. Scales: ca. 56-62.
Caudal peduncle: depth 7.2—10. Ventral scutes: ca. 39-46.
Head: length 22.6—26.4. Gill rakers: 37-52.
Snout: length 5.6—7.1. Fin rays: dorsal 16-20, usually 17-19;
Eye: diameter 5.3—7.7. anal 16-20, usually 17 or 18; pec-
Interorbital: width 3.0-4.5. toral 17-19.
Maxillary: length 10.2—12.3. Vertebrae: 55-57 (12 specimens).
Pelvic fin: length 9.4.-11.3.
Pectoral fin: length 14.2-17.7.
Ficure 66. Clupea harengus, 315 mm TL, 12.62 inches, USNM 16667. Drawn by H. L. Todd.
Bopy compressed, its greatest thickness somewhat less than half of its depth, its
depth 3.9-5.0 in SL, much less in very young, 5.2—9.5 in specimens 40-70 mm.
Sca.es only moderately adherent, often lost, especially in small fish; scales with
crenate membranous border; about 16 or 17 longitudinal rows between ventral edge
at base of pelvic and anterior ray of dorsal. VenTRaL scuTEs weak, often difficult to
count, about 26—33 in front of pelvic fin and 13-15 behind.
Heap 3.8-4.4. SNouT 3.25—4.1 in head. Eve 3.4—4.25. INTERORBITAL 3.0—4.5.
Maxittary rounded posteriorly, generally not quite reaching middle of eye, 2.0—2.3 in
head. Gitt RaKERs at angle nearly as long as eye in adults, proportionately shorter in
young; increasing in number with age, the principal increase taking place before the
specimens reach a length of 100 mm; specimens 36—70 mm long with only about
25-36 on lower limb. TEETH missing in upper jaw; the margin of maxillary with minute
serrae; lower jaw anteriorly with about 6 rather prominent teeth on each side; middle
of tongue with an elongated patch of small teeth; vomerine patch elongate, the teeth
rather few and fairly strong.
Dorsat fin little elevated anteriorly, with nearly straight margin, its origin some-
what in advance of pelvics, usually nearer to base of caudal than to tip of snout. CaupaL
with lower lobe rather longer than the upper one, about as long as head. Anat very
low, its origin about equidistant between base of pelvic and base of caudal. Petvic
very near ventral edge, inserted equidistant between base of pectoral and last anal
Fishes of the Western North Atlantic 277
ray, 2.2—3.0 in head. Pecrorat inserted a little higher on side than pelvic, and below
distal margin of opercle, 1.3—1.6 in head.
Color. According to Bigelow and Welsh (16: 93):
Peritoneum quite dusky. Deep steel blue or greenish blue on the back with green reflections; the sides and
belly silvery; the change from dark to pale sides often marked by a greenish band. The gill covers sometimes
glisten with a golden or brassy gloss; indeed, fish just out of the water are iridescent all over with different hues
of blue, green, and violet, beauties that soon fade, however,
leaving only the dark back and silvery sides. The ventral and
anal fins are transparent white; the pectorals, however, are dark
at the base and along the upper edge; the caudal and dorsal
dark grayish or shading into blue or green.
Size. The maximum size attained by the
Atlantic Herring is about 450mm (18 in.),
though the usual size is only about 300 mm
(Ge) se)
Scope of Following Account. Much has been
written about this Herring’s spawning habits, food,
rate of growth, seasonal migrations, and so on,
by both European and American authors. Sum-
maries of the literature about this species on the
American coast, with frequent references to the
European literature, have been given by Bigelow
and Welsh (16: 92-105) and by Bigelow and
Schroeder (15: 88-100), who drew heavily on
important papers by Moore (95: 387-442), Lea
(78: 75-164), and Huntsman (63: 165-171);
furthermore, they were able to add much to the already known facts from their own
studies. A review of all the literature would make this account much too long for
the present work. Therefore, only a brief summary of what is known about its life
history, drawn in large part from the accounts mentioned above (15; 16), is given here.
Nearly all of the papers cited provide bibliographies from which references to literature
containing additional information may be obtained.
Development and Growth. The eggs of the Atlantic Herring, 1.0-1.4 mm in
diameter (6: 98) or 1.2-1.5 mm (29: 362), are heavier than sea water, and adhesive,
adhering singly or in clumps to objects in the water or to the bottom. If they fall on
muddy bottom they probably smother.
The number of eggs produced by a single female is relatively small, and as usual
it varies according to the size of the fish, large fish having many more eggs than small
ones. The number produced by one female may vary between 21,000—47,000 (ZOT:
283) or between 20,000—40,000 (16: 94).
The duration of incubation depends upon the temperature. Norman stated that
eggs kept at 52—58°F hatched in eight or nine days, whereas others held in water at
Ficure 67. Clupea harengus (Europe) eggs
attached to seaweed. After Ehrenbaum.
278 Memotr Sears Foundation for Marine Research
32°F hatched in 47 days (ror: 330). Bigelow and Welsh gave 10-15 days as the
probable average incubation period for the usual temperatures prevailing in the Gulf of
Maine during the hatching season; these seem to vary between about 46—5 5° F (16: 94).
The newly hatched fish, transparent and very slender, vary considerably in length,
with a range of about 5.3-7.0mm (29: 362) or about 5.0-6.0 mm (16: 97). The small
ot
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Ficure 68. Clupea harengus (Europe). a—p, larval stages; A 7mm; B IOmm; c 19 mm; D 29 mm; E young
fry, 41 mm.
yolksac is absorbed in several days, when the larvae have attained a length of 8-romm,
no marked changes take place in the form of the body during this time. The vent is
situated so far back that the length posterior to it is less than a fifth of the total body
length, and the depth at this size is contained about 24 times in the total length. Con-
siderable advancement in the development of the fins was evident in larvae around
19 mm long, taken in the tow at Helgoland; rays definitely were present in the dorsal
and caudal, and the caudal had a concave margin. However, the anal and pelvic fins
did not become differentiated until a length of about 29 mm was attained. In young
about 41 mm long, the proportionate depth of the body has increased greatly, and the
vent as well as the anal and pelvic fins have all moved forward; larvae of this size are
said to be in the transition stage. However, this stage may be attained by the spring fish
at a length of 31-44 mm, and by the fall fish at 44-60 mm (29: 363-366, figs. a—g).
Fishes of the Western North Atlantic 279
Young fish from the Clyde Sea area of England lost their yolksac and began
developing the dorsal fin at about 10 mm; the end of the notochord turned up at about
17 mm; pelvic fins appeared at about 22 mm; the caecal part of the stomach began to
form and the air bladder became prominent at a length of 30-35 mm; and the least
length at which metamorphosis took place was 42 mm (gr: 248, 252). Marshall, et a/.
also recorded a regular weekly increase of 3 mm prior to metamorphosis, at a length of
about 35-40 mm. This rate of growth of the larvae is in fair agreement with that
reported for elsewhere in Europe.
Huntsman observed that, on the American coast, as at Grand Manan Island,
New Brunswick, larvae hatched in late summer and early autumn had attained a length
of 17-20 mm by the end of November or early in December. Fry 50-60 mm long
were abundant in St. Andrews Bay in June. Yearlings were go—125 mm long in the
fall of the year in the Bay of Fundy. This rate of growth is in agreement with that ob-
served by European investigators in examples from Norwegian waters and from the
North Sea. A one-year old Atlantic Herring, then, is about 88-125 mm long.
These fish do not fall into definite size groups when more than one year old. But
their subsequent rate of growth may be determined by segregating the fish into year-
classes through scale reading, for the so-called winter rings are so pronounced that they
can be relied upon with considerable confidence to show the age of the individual
specimen.!° Thus, Meek (93: 76) has indicated the following average lengths for those
caught on the Dogger Banks in the North Sea: 98 mm at one year, 183 at two, 220 at
three, 241 at four, 255 at five, 266 at six, 275 at seven, 281 at eight, 287 at nine, 291
at ten, 293 at eleven, and 295 mm at twelve years of age. The Norwegian fish spawned
in 1899 grew more slowly at first than the Dogger Bank fish but about equaled the
latter in size by their sixth year. Fish up to 19 years old have been found in many
Canadian samples (83: 10) and some 20 years of age have been seen in Europe. How-
ever, the rate of growth after age 12 is extremely slow. According to observations by
Huntsman and by Lea, the rate of growth is about the same for Atlantic Herring both
here and in Europe, with Bay of Fundy fish growing a little faster than those spawned
in the Gulf of St. Lawrence, and with the fish spawned along the outer coast of Nova
Scotia growing a little faster than those produced in the Bay of Fundy.
On the coast of Maine, some individuals reach sexual maturity at a length of about
235 mm (9.5 in.), when three or four years old. Usually they do not spawn until they
are 250-262 mm (10—10.¢ in.) long, at five to six years of age. Most of the individuals
in the schools of spawning fish are 300-325 mm (12-13 in.) long, the majority of
which is then perhaps 10 years of age or more. It is believed that Atlantic Herring,
after reaching sexual maturity, spawn every year as long as they live.
Atlantic Herring may be divided roughly into three growth stages," each with its
10. For a detailed study of the scale of the Atlantic Herring in relation to growth, see Lea (78:81~-93).
rr. Sometimes a greater number of growth stages has been recognized. Thus, Johnson (69:351), following the nomen-
clature of the fishermen of Passamaquoddy Bay, listed “brit”? 80-100 mm, “snippers”’ 110-130 mm, “‘oils” 140—
180mm, and “mustards” 190-230 mm. In New England at least, the term “‘sperling”’ is used for young fish 75—
125 mm long; they therefore come within the range of the sardine group, as used by Bigelow and Welsh (z6:99).
280 Memoir Sears Foundation jor Marine Research
own general distribution and seasonal migration. The first stage, the ‘“‘sardine,”’ consists
of one- and two-year-old fish 45-200 mm long; the second is the “‘fat’”’ stage, con-
sisting of sexually immature fish about two years old, 190-200 mm long; and the
third or “‘spawn”’ stage is a mature fish (see also pp. 283, 284).
Spawning Seasons. Spawning takes place in the spring, summer, or autumn, or
in both spring and autumn, depending on the locality. Apparently there is no time in
the year when spawning, off Europe at least, does not take place somewhere. On the
southern coast of Newfoundland and in the Gulf of St. Lawrence, they are spring
spawners, commencing in May or even as early as the latter part of April. In the southern
and inner part of the Gulf as a whole, the season lasts only a few weeks. But on the
south coast of Newfoundland, and perhaps on the west coast as well, spawning may
continue until July, and some fish spawn in the Gaspé-Bay of Chaleur region in Sep-
tember. To the south, on the Atlantic coast of Cape Breton, Nova Scotia, spawning
occurs chiefly in May or June, but also in August and September; and on the outer
coast of this province to the south of Halifax, from August through September (95:
405; 60: 10; 83: 11, fig. 6).
The Bay of Fundy once harbored both spring and summer-autumn spawning
schools, as did the outer coasts of Nova Scotia (by fishermen’s reports); but they seem
never to have been numerous; it is doubtful whether any of the spring spawners still
exist anywhere south of the Gulf of St. Lawrence. At the mouth of the Bay of Fundy
and off the easternmost part of the Maine coast, the heaviest spawning takes place in
July-August and September in some years, with spawning continuing late in the fall;
but in other years it does not commence until August, to end early in October. Passing
westward—southward, the spawning run is progressively later and shorter: from mid-
August until October in the general vicinity of Mt. Desert Island; in October along
the east coast of Massachusetts (where only a few spawn); and early November near
Woods Hole, Massachusetts.
Off northern Europe, where two principal races are recognized, namely coastal
herring and sea herring, the coastal fish spawn mostly in brackish water and princi-
pally in the spring months, whereas the sea herring spawn in the open sea over widely
distributed times of the year, but mostly in the late summer, autumn, and winter. The
spawning periods of the different stocks are so varied that one may find spawning
fish somewhere at almost any time of the year (2g: 362). The Pacific Herring has
only one spawning period, which may vary between December and June; the earliest
spawning takes place in California and the latest in northern Alaska (rrr: 278).
Spawning Grounds and Depths. In northern European waters, as already indicated,
at least some fish spawn in shallow inshore brackish water whereas others spawn in the
open sea. But in the western side of the Atlantic they are not known to spawn in water
less saline than about 31.9 °/o, or more saline than about 33 °/o9. In the Gulf of Maine,
spawning takes place at temperatures of about 46—55°. For the more northern part
of the American range, however, precise information in this regard is lacking.
Off America, Atlantic Herring spawn in greater or fewer numbers throughout
Fishes of the Western North Atlantic 281
their geographic range, from north to south. Around Newfoundland they spawn chiefly
within the bays. However, from the southern side of the Gulf of St. Lawrence south-
ward as far as southern Maine there is no considerable sector of the coast, except a
part of the Bay of Fundy (p. 290), that does not see a more or less plentiful spawning
run. Moore has spoken of the northern coastline of the Gulf of Maine as “‘a con-
tinuous spawning ground,” succeeded by scattered grounds wherever the bottom is
suitable, extending as far southward as Block Island off the Rhode Island coast (95: 408).
However, local spawning grounds may be abandoned for a term of years—a common
occurrence (95: 430).
In the Gulf of St. Lawrence, Canada, they have been described as spawning in
water so shallow that the eggs are sometimes washed up on the beach by the waves
(95: 411), as happens every year in northern European waters (15: 99). In general,
however, our Atlantic Herring spawn at depths of, say, 2-30 fms. Thus spawning takes
place both along shore and on the various ledges that lie between 5—25 miles off the
coast, where eggs are often found sticking to the anchor lines of fishing vessels. But the
nature of the bottom makes it improbable that any eggs are laid at a depth greater
than 75 fms. on the American side, though spawning at a depth of 100 fms. has been
reported for Scandinavian waters (95: 411). Whether, or how regularly, the fish may
spawn on the shallower parts of the offshore fishing areas, from Georges to Grand
banks, remains to be learned.
Habits." Herring are primarily wanderers of the open sea. While they also enter
bays and estauaries freely, they seem never to have been reported reliably from water
that is more than slightly brackish; along the American coast a salinity of perhaps
2.8 9/59 is about the lower limit of tolerance.
As a rule, all sizes of Atlantic Herring congregate in schools of hundreds or
thousands of individuals, with a given school usually consisting of individuals of about
the same age and size. How long a school may preserve its identity as such is not
known, for while Fridriksson and Aasen have found that specimens tagged and released
together may be recaptured at widely separated localities (¢o: 22), this may not apply
to schools that have assembled under natural conditions.
When a school is at the surface in the daytime, as often happens in calm
weather, it is betrayed by a fine rippling of the water; but the fish do not ordinarily
“fin” or lift their noses above the water, as is characteristic among menhaden
(p. 354). At night they are betrayed at the surface by their luminous trails if the
water is “firing.”
A school of Atlantic Herring is not as easily frightened as a school of mackerel
by an approaching boat. Neither is it usual for them to leap clear of the water, except
when young fry are pursued from below by larger pelagic fish.
A school that is actively feeding is likely to be more or less stationary, drifting as
a whole with the current. At other times schools are seen traveling as if with purpose-
ful intent, all of the members swimming side by side in the same direction. Schools
12. Condensed from Bigelow and Schroeder (r5: 89).
282 Memoir Sears Foundation for Marine Research
of sardine-sized fry may often be seen streaming past some rocky headland, seemingly
in an endless procession. What the directive stimulus may be we have yet to learn, for
as Huntsman has pointed out, “There is no indication that herring swim against
the current unless the water is somewhat turbulent’ (65: 83). Whether they make or
lose headway over the bottom while doing so depends on their swimming speed relative
to the strength of the current.
No information is available as to how rapidly a school may travel at sea. But Frid-
riksson and Aasen have found that specimens held alive in nets swam constantly at a rate
of about 0.2—0.25 sea miles per hour (6-8 m/min.) when not disturbed. It is certain
that they are capable of long journeys, for individuals tagged on the east coast of
Iceland have been recaptured in southern Norway, and vice versa (40: 26, 27).
How active Atlantic Herring are at any time and place depends primarily on
the water temperature. In the Bay of Fundy region, for example, they “move very
sluggishly when the water is coldest, in February and March,” 1.e. when the upper 20
fms. or so have cooled to about 32—36°. They become active again when the water has
warmed to about 40-43°.
While the vertical range of this species extends down to an extreme depth of
about 100 fms., the vast majority lives at depths shallower than 40-50 fms., living
pelagically in the upper water layers for most of the year but close to the bottom during
the coldest months.
Relation to Light. That light is an important factor in controlling their vertical
distribution, at least in southern New Brunswick, was discovered by Johnson in 1939
(69: 349-354). He found that at night, in the absence of appreciable light, all sizes
were quite near the surface. During weak light, as at dawn and dusk, all sizes still were
a foot or so from the surface. However, as the sun rose above the horizon they left
the surface, the largest fish, 190-230 mm long, descending to a depth greater than
10 feet and those 140-180 mm long to a depth of 10 feet or more; the smaller ones in
general were nearer the surface. Also, on cloudy days all sizes were closer to the surface
than on clear days.
Migratory Movements. Because of the economic value of this species, knowledge
of the migrational habits, which would make it possible to predict runs in advance of
their occurrence, has been and continues to be sought on both continents. In Europe
at least, these studies have been complicated by the presence of a large but in-
determinate number of local populations or races (p. 287), each with its own mi-
gratory pattern, a fact not known until comparatively recently and even now not well
understood. As Norman (ror: 263) wrote:
At some seasons herrings may be found in huge numbers in a given locality (in Europe), at others they
will disappear almost entirely; in other places they may be caught all the year round, but the numbers caught
on a given ground may exhibit an immense amount of variation from one season to another.
To explain these long known and very obvious facts, an older view, widely held
until about 1825, was that Atlantic Herring as a whole withdraw in winter to Arctic
Fishes of the Western North Atlantic 2832
waters, to return to more temperate coasts in spring, some of them to the eastern side
of the North Atlantic, others to the western side (for further details, see Meek, 93:
67-87). But Perley wrote as early as 1852 that this idea was “supposed to be wholly
imaginary,” it being generally believed then “that the herring fattens in the depths
of the ocean, and approaches the shore in shoals merely for the purpose of depositing
its spawn” (104: 206). Successive and long continued studies at many hands have
since proven that in reality we are not dealing with any widespread mass migrations,
but with short-range movements (inshore and offshore) of local bodies of fish, each
with its own area of occurrence and each of which may include subpopulations that
spawn at different times of year, as Norman has pointed out. It is equally clear that the
ranges of the races of European fish overlap—highly probable at least that more or
less interchange is constantly taking place between adjacent races.
However this may be, the basic migratory pattern, essentially the same for all of
them, may be conveniently divided into the three successive phases noted previously—
sardine, fat, and spawn.
Those of the sardine stage, 45-200 mm long, tend to remain near their birth-
place. They probably spend the winters on the bottom in a few fathoms, appear in the
inshore waters of New England, New Brunswick, and Nova Scotia in the spring as
one- and two-year-old fish, and remain there through the summer and autumn.
How far they may journey during this period and in what direction depends chiefly
on the movements of the water. For example, every summer untold millions of sardines
congregate in the Passamaquoddy region at the entrance to the Bay of Fundy (p. 290),
and it is now known! that the responsible factor is the prevailing movement shoreward
of the subsurface water in which the sardines are swimming; this shoreward flow is
motivated by the offshore movement of surface flow from tributary streams combined
with superficial offshore currents set up by the wind. Sardine-sized fish, in other
words, drift with the current much as do any planktonic animals, such as the euphausiid
shrimps and copepods on which they feed. In technical language, they are denatant,
which applies equally wherever young Atlantic Herring are produced.
The fat stage consists of sexually immature fish about two years old and 190—
200 mm long that have accumulated a large amount of fat around the viscera and
among the body tissues. During the year’s period that precedes their sexual maturity,
they disperse much more widely than do the sardines. Whether this wide scattering
is brought about wholly by transport of water movements or whether directive swim-
ming plays an important role remains a mystery. In either case they are encountered
anywhere and everywhere throughout the range of this species. In the Gulf of
Maine, for example, where more attention has been paid to the movements of these
fish than has been the case anywhere else in the western North Atlantic, the fat
stage has been encountered indifferently close to shore, as at the mouth of the Bay
of Fundy in the northeast, and offshore in the Massachusetts Bay region in the
13. Studies carried out from the Atlantic Biological Station at St. Andrews, chiefly under Dr. A. G. Huntsman’s
leadership (65:95, 96).
284 Memotr Sears Foundation for Marine Research
southwest, in the central parts of the Gulf, and over Georges Bank. Those picked
up during the warmer months either in mackerel seines or in otter trawls are mostly
very fat and show no signs of approaching sexual maturity. It seems that these “fat’’
fish, like the sardines, merely sink down close to the bottom for the winter wherever
they may be.
About all that is known of the mature spawn stage in the western Atlantic is that
these fish live mostly some distance offshore near bottom and appear in vast num-
bers in their spawning areas at spawning time. In the Gulf of Maine, most of them
seem to drop out of sight after having spawned; the large ones caught there out of
spawning season fall in the fat category. But there is no reason to suppose that they
travel far during the interval (between spawning time and winter) when they are
feeding greedily to recover condition sufficiently for spawning again the next year. In
fact, it is probable that they, like the other two stages in the Gulf of Maine, merely
descend into deeper water to winter, as has long been known to be their custom in
European waters. How deep the great body of them may go is not known, but the
contour of our continental shelf is such that no fish need swim any great distance to
reach water deeper than 50-75 fms. anywhere between Cape Cod and the Gulf of
St. Lawrence. It seems likely also that they spend the winter in deep water in the
southern part of the Gulf of St. Lawrence, but definite information in this regard
is lacking. On the southern and western coasts of Newfoundland, however, where
spawning takes place in the spring, the mature fish ‘‘move off into deeper water in
summer” (125: 35) but return late in the autumn to their natal bays, where they
have long supported important November—January fisheries.
Food and Feeding. Atlantic Herring feed on plankton—first on diatoms and other
unicellular organisms, then on copepods and their eggs following the yolksac stage
(91: 252). In summing up her rather extensive investigations at Plymouth, England,
Lebour (79: 463) listed the following by stages of growth; before disappearance of
the yolksac: larval gastropods, green food (probably diatoms and flagellates), larval
bivalves, nauplii and other young stages of small crustacea, as well as the eggs; after
disappearance of the yolksac up to about 12 mm: the same, but with small adult
copepods added; after 12 mm and probably until metamorphosis: copepods; after
metamorphosis: copepods, decapod crustaceans, amphipods, and fish.
Only two kinds of food were found in about 1500 examples taken off East-
port, Maine (95: 401). Moore said that “One of these [foods] consisted of copepods
(‘red seed’), which appeared to constitute the sole food of the small herring, the so-
called brit, and a considerable portion of that of the larger individuals from 51/, inches
upward.” But the principal food of the larger fish was euphausiid crustaceans
(Meganyctiphanes norwegica), known to local fishermen as “shrimp.” These are so im-
portant in the diet of this species that it seems likely that the appearances and disap-
pearances of large fish in the open waters of the Gulf are correlated with the presence
or absence of euphausiids of one kind or another. Concerning their capture, Moore
remarked:
Fishes of the Western North Atlantic 285
To capture such prey requires some address on the part of the herring; they frequently throw themselves
almost clear of the surface and their splashings... are audible at a considerable distance. When feeding on
copepods the movements of the herring are less impetuous. They swim open-mouthed,™ often with their snouts
at the surface crossing and recrossing on their tracks and eventually straining out the minute crustaceans by means
of their branchial sieves.
The sieves were described by Moore (95: 401) as follows:
An examination of the mouth cavity of the herring will disclose a series of long bristle-like processes,
the gill rakers, projecting from the anterior face of each gill arch, like the teeth of a comb. When the mouth is
opened widely the tips of the gill rakers stand apart, but when it is closed or partly closed they become more
closely approximated and each series is pressed closely against the inner face of the series attached to the arch
next in front. There is thus formed a beautifully fine and effective sieve, capable of retaining small bodies contained
in the water taken in at the mouth and discharged through the gill slits.
The amphipod genus Euthemisto is also an important food for this species off
Europe; that this is not so in the Gulf of Maine no doubt reflects the scarcity of this
pelagic crustacean in our coastal waters. In default of an abundant supply of crustacea,
and sometimes even when these are plentiful, the fish feed on whatever other kinds of
smaller planktonic animals and plants may be available. Small fish of various kinds
have been found in their stomachs: capelin, especially in Newfoundland waters in
winter; and grass shrimps (Crago), schizopod shrimps (Mysis), amphipods (Gammarus),
small squid, annelid worms, and crab larvae at Woods Hole, Massachusetts. But the
smaller planktonic plants, whether diatom or peridonidian, have never been found in
specimens longer than 15 mm, probably because their gill sieves are not fine enough
to retain them.
Enemies. Their chief natural enemies include all kinds of marine predators, such
as cod, pollock, haddock, silver hake, mackerel, dogfish and other sharks, finback
whales, and the common squid (16: 104). An attack on a school of young Atlantic
Herring by squid has been described by Hildebrand and Schroeder (59: 82) as follows:
We had occasion to watch the wholesale destruction of 2 to 4 inch herring during June 1925, on the flats
about Provincetown, Mass. Schools of 10 to perhaps 50 squids circled around a school of herring until they had
bunched their prey into a compact mass. Individual squids then darted in and seized one, sometimes two, and then
darted back for more. Along the beach there remained a silvery streak of dead herring.
A similar firsthand account of silver hake (Mer/uccius bilinearis) driving a school of large
Atlantic Herring up on the beach in Massachusetts Bay has been reported (15: 90).
Parasites. Among the European fish, “‘cestode larvae and trematodes were fre-
quently found as parasites in the gut. The latter were common and the incidence ap-
peared to increase as the herring grew, as many as 29-30 per cent being parasitized
in the large catches of May 21 and 30. Cestode larvae were less numerous, and they
too were more frequent in the older fish” (Marshall, et a/., gr: 253). No fatalities
were specifically ascribed to these parasites; Sindermann and Rosenfield have found
that infestation with trematode larvae may be fatal—at least under experimental con-
14. That the larger herring “‘swim open-mouthed” when feeding on copepods has been questioned by Battle, ef al.,
who are of the opinion that the fish seize the organisms individually (7:411). However, as to the organisms ingested
there is general agreement.
286 Memoir Sears Foundation for Marine Research
ditions—“‘provided they are present in numbers large enough to overwhelm the host”’
(117: 12). Furthermore, it has been reported that nematodes, cestodes, and trematodes
occur in the viscera and sporozoa in the muscles of small fish taken at Woods Hole,
Massachusetts (85: 473).
Mass Mortalities and Diseases. The literature that deals with the Atlantic Herring
contains many accounts of their great destruction on both sides of the North Atlantic.
For June and July 1914, Cox reported widespread mortality on both the southern and
western sides of the Gulf of St. Lawrence (28: 82-85). A few years later, a spectacular
event occurred in Cohasset Harbor, Massachusetts Bay, on October 5, 1920, when an
estimated 20,000 barrels of fish to0o—125 mm long became stranded on the mud with
a falling tide (76: 104). On October 10 a second but smaller run and on October 15
a run as large as the first were stranded similarly. The exact reason for the stranding
and mass destruction was not determined, but fishermen thought that the fish had
been driven into the nearly landlocked harbor by silver hake, Merluccius bilinearis.
Other mass strandings of small individuals have also been reported, for Manchester
Harbor, Massachusetts (14: 327). Pursuit by predatory fishes is the most likely cause
of simultaneous mass strandings within small areas; for an eyewitness account of one
such event, see page 285.
But enough evidence has now accumulated to make it practically certain that
when mortality is very widespread it has been caused by disease of one kind or
another. Thus Cox found that fish from the Gulf of St. Lawrence case of 1914 (above)
were so evidently in a diseased condition and were so heavily infested with a myxo-
sporidian protozoan that it seems practically certain that the latter had been responsible
for the mortality (28: 82-85). It is known now that those along the coasts of New
England, and presumably in more northern waters, suffer from fungous disease caused
by Ichthyosporidium, from ulcer disease caused by protozoa, from pigment spot disease
caused by larval trematode worms, and from tail rot caused by bacteria; it is known
also that fungous disease in particular, and probably the others as well, sometimes
reaches epidemic proportions, with mortality so widespread and so severe that the sardine
fishery is very seriously affected.
Relationships. Although the Pacific herring has been recognized as being specifi-
cally distinct from the North Atlantic Herring and has for many years been designated
Clupea pallasii Cuvier and Valenciennes, it is difficult in the light of rather recent studies
to find distinguishing characters. This difficulty was pointed out by Rounsefell, who
made some studies of the relationship himself and reviewed the findings of several
European investigators (111: 243). The distinguishing character given by Regan in
his revision of the Clupeidae is “Ventral scutes all keeled” in harengus and “Ventral
scutes in front of pelvic fins not keeled” in pal/asii (106: 227). According to the material
now at hand, the scutes in pa//asii are merely less strongly developed.1®
15. For a general survey, with photographs, of diseased fish, and references, see Sindermann and Rosenfield (177).
16. Relationships with the species of temperate southern hemisphere waters have not been adequately worked out.
Regan, the most recent revisor, had inadequate material (106:228). Norman did not attempt to compare the South
Fishes of the Western North Atlantic 287
Variations. It has long been generally appreciated that Atlantic Herring in the east
do not constitute a homogeneous population, since a number of regional races is in-
cluded (pp. 282, 283). Thus “‘it is possible,’’ writes Norman, “‘to recognize North Sea,
Baltic, Norwegian, Icelandic herring” (ror). These races differ from each other in such
morphologic characters as average proportional dimensions and average numbers of
fin rays and vertebrae. But we ought to caution the nontechnical reader that these are
numerical averages for many specimens and are not used for the identification of an
individual specimen, or of even a few fish; only when the number of specimens examined
is large are such figures significant. For example, if the average number of vertebrae
was given as 56.3—56.5 for one locality and as 56.6—56.8 for another, it means simply
that in the first case there were more fish with 56 vertebrae and less with 57; and in
the second case, more fish had 57 and less had 56. No fish ever has a fractional num-
ber of vertebrae.!”
Each of these races has its own chief center of abundance, its own pattern of sea-
sonal migration, and its own spawning season or seasons, for some of the races include
both early and late spawners. The races differ in the average size to which their mem-
bers grow. A knowledge of the times and localities where these populations are catch-
able in greatest amount is of so much commercial importance that these matters are
under continuous survey by the fisheries services of the countries chiefly concerned.
In 1914, Hjort found (60: 9-12) in the western Atlantic Herring: that the average
number of keeled ventral scales is larger (13-14) for the autumn spawners of outer
Nova Scotia and the Gulf of Maine than for the spring spawners of the Gulf of St.
Lawrence; that the number of vertebrae is greatest (55-59, av. 56.8) in those from the
west coast of Newfoundland; that the number of dorsal and anal rays averages higher
in those caught in the open sea than in those from the more enclosed waters of the
Gulf of St. Lawrence and Bay of Fundy; and that, among the spring spawning popula-
tions, the average number of vertebrae, fin rays, and keeled scales is higher in those
from the west coast of Newfoundland than in those from the southern part of the
Gulf of St. Lawrence. But the fishery for ‘‘fat” and for “‘spawn”’ fish plays so small a
role in the general welfare of Canada and the United States (p. 288) that no additional
information seems to have been contributed to the racial question on this side of the
Atlantic. From analyses of extensive data, however, Day, Leim, and Tibbo showed that
Canadian Atlantic Herring represent six more or less distinct populations, differing
from each other in rate of growth and in average number of vertebrae: one in the northern
part of the Gulf of St. Lawrence, a second in the Bay of Chaleur, a third in the southern
part of the Gulf, a fourth off the outer Nova Scotian coast, a fifth along southern New-
foundland, and a sixth along western Newfoundland (see Leim, 83: 11, figs. 6, 12).
Commercial Importance. From an economic standpoint, these fish are perhaps the
most important in the world. In addition to their great value as food for man, many
American forms with the northern ones (102:37). Regan’s Clupea holodon from New Zealand is now known as
Clupea antipodum (Hector).—G. S. Myers.
17. For a general survey of the races of eastern Atlantic Herring, see especially Liibbert and Ehrenbaum (89: 14-19);
for a detailed study of the average number of vertebrae in the different races, see LeGall (80: 167-170).
288 Memoir Sears Foundation for Marine Research
are used for bait, and they constitute an important item in the diet of other fish that
are of great economic importance. The catches have fluctuated greatly from year to year,
but they have not always provided a criterion of the supply available. In America, at
least, the catches have often been governed by demand and profit. Therefore the fishing
efforts from year to year have not been uniform. Through 1952, the annual catches
on the American side have fluctuated between 172,000,000—359,000,000 pounds,"
the major part of the catch from Canada and Newfoundland. The catch in Europe in
1948 was 4,1 50,000,000 pounds (1,881,713,000 kilo; 24: 6), and in 1956, the latest
year for which information is conveniently available, it was 5,858,650,000 pounds
(2,590,715,000 kilo). European waters, in short, have yielded annually something like
15 times the catch (in pounds) taken in American waters. However, the apparent dis-
crepancy in relative productivity does not loom so large when we note that the bulk
of the American catch is concentrated within a latitudinal belt of about 8°30’ at most,
while the European catch is distributed along something like 19°30’, or a distance
nearly 2.5 times greater.
Small fish, 75-125 mm or so in length, are canned in large quantities as sar-
dines, not only in Norway but along the easternmost coast of Maine and in the region
of Passamaquoddy Bay at the mouth of the Bay of Fundy on the New Brunswick side;
these last account for the concentration of Atlantic Herring landings there. Elsewhere
the American catch consists chiefly of the ‘‘fat”’ and “spawn” stages. Some of the large
ones are consumed fresh, others are salted or pickled, some are salted and smoked in
various ways, and still others are canned.
The three principal types of smoked Herring are the kipper, bloater, and red her-
ring. The kipper is split down the back, immersed in brine for a very short period of
time, slightly dried, and then smoked for several hours. The bloater is cured without
splitting, only the belly being cut open for cleaning; it is lightly salted and smoked
only long enough to dry the flesh. The red herring, like the bloater, is not split, but it
is much more heavily salted (buried in salt for at least five days) and then smoked for
ten days or so. Thus the last is a fairly durable product, which can be shipped to distant
markets, whereas the kipper and bloater are perishable. The several curing processes
are practiced much more extensively in Europe than in America and are sometimes
carried out aboard fishing vessels. Large quantities are also frozen, to be used as bait
by the hand-and-long-line fishermen: for example, 32,000,000 pounds in Canada (in-
cluding Newfoundland) in 1954. While corresponding information is not conveniently
available for the United States, this time-honored method of fishing is now far out-
stripped by the otter trawlers. The landings by line fishermen in Maine and Massa-
chusetts in 1954 were 9,000,000 pounds, the greater part of which was hooked on
frozen herring. The offal at the canneries and occasionally surplus catches of whole
fish are reduced to oil, fish meal, and fertilizer.
Fluctuations in Abundance and Catch; Year-Classes. It is notable that a fishery such
as the one for this species in European waters, exploited for centuries, has apparently
18. Statistics from publications issued by the governments of the respective countries named.
Fishes of the Western North Atlantic 289
shown no positive or permanent decline. Of course it has been claimed from time to
time that signs of exhaustion were evident, but such claims seem to have been based
on the scarcity or disappearance of the fish in certain areas, not on the population as a
whole. There certainly is insufficient information concerning the American stocks to
furnish a basis for an answer. If a decline has taken place in American waters, it could
scarcely be ascribed to overfishing, as the supply of fish generally has exceeded the de-
mand. The fluctuations in the annual catches, then, do not necessarily mean that the
fish were scarce when a small quantity was caught, or were abundant when a large catch
was taken. Neither does a small annual catch necessarily indicate that the demand was
light. It may mean only that the fishermen were operating in an area where the fish
are usually abundant but were absent there that year. The fish are influenced in their
migrations by the weather and by the presence of food, which seem to account for their
abundance in one area one year and elsewhere the next. At present it is impossible
to predict the time of arrival of schools in any given area.
From his studies of Atlantic Herring from the English Channel, Ford wrote,
“Sf it could be shown that the variation from season to season followed some rule, it
might yet be possible to discover that rule from the study of past data and hence to predict
what is likely to occur in the future” (36: 369). But all uniformity seems to be lacking.
“Tt can be regarded as certain that fishing results are dependent upon the weather,
but in the present state of meteorological science, we cannot hope to receive much
advanced information regarding weather conditions during a forthcoming season.”
Nevertheless, he found that daily forecasts had some value, for a gale warning not only
indicated an interruption of operations, but it also suggested heavy catches when fishing
was resumed. The prediction of calm weather, on the other hand, suggested lighter
catches.
The many age analyses of Atlantic Herring, chiefly of those from European waters,
have shown that a very large crop of young may survive in some years while very
few do in other years, especially in more northerly spawning regions. Consequently,
those spawned in a favorable year may dominate the population for several years to
come. The classic example is the year-class produced in 1904 that dominated the
Norwegian catches for the next six years and dominated catches on the west coast of
Newfoundland until 1914 and even 1915. In the Gulf of St. Lawrence, of late years,
the broods of 1939, 1941, 1943, and 1944 were highly productive, whereas those of
1940, 1942, and 1945 contributed but little to the population (83: 107). There is
also the well-documented case of a particular body of Bay of Fundy fish that did
not receive any significant recruitment for some 10 years, by which time the few re-
maining fish seem to have dropped out of the picture due to old age (15: 93, 99). Con-
sequently, the successes or failures of reproduction in given years are reflected in the
catches. Thus Ford (36: 370) was able to predict which year-classes would predominate
in the catches of certain years, as well as the probable proportion of large fish, for as
a rule old fish are larger than the younger ones, though the fish in some areas grow
faster than those in others. Ford concluded his discussion on predictions as follows:
SS)
290 Memoir Sears Foundation for Marine Research
In practice it is possible to obtain advance information of the relative richness of broods and hence of the
relative success of future fisheries. But there still remain possible influences which militate against a forecast
of yield based entirely on this evidence, chief among these being the factors which govern migrations of herring.
Methods of Fishing. Most of the fish are caught in traps known as wiers, in floating
traps, stop seines, purse seines, and drift nets; in the American bays and harbors, large
numbers, especially of the smaller sizes, are taken by “‘torching,” i.e. by netting fish
attracted by a light.
Range. Atlantic Herring in American waters range from Greenland to North
Carolina, but usually not in commercial abundance south of New Jersey; they are most
abundant north of Cape Cod. In Europe, where certain races enter brackish water,
they range from Spitsbergen, Novaya Zemlya, the Kara Sea, the White Sea, and
northern Norway south to the Straits of Gibraltar.1®
Occurrence in the Western North Atlantic. Stray specimens have been reported for
as far south as Cape Hatteras, the lower part of Chesapeake Bay, Maryland, and New
Jersey (59: 82; TIQ: 120; 37: 93); and only irregularly do a few visit the coasts of
New York and Rhode Island. But they are seasonally abundant and generally distrib-
uted throughout the Woods Hole region, though not in any great numbers (123: 741).
Although they appear sporadically in the Massachusetts Bay region, they are plentiful
enough in some years to support local fisheries. Along the coast of eastern Maine, and
in the Passamaquoddy region at the mouth of the Bay of Fundy,?° they are not only
far more plentiful but more regular in their comings and goings. But farther up the
Bay of Fundy, on the New Brunswick side and at its head, they are far less plentiful,
probably because of the turbidity of the water there. However, along the Nova Scotian
side, toward the mouth of the Bay, they are sufficiently abundant so that they yielded
between five and six million pounds in 1946 (Annapolis and Digby counties); indeed in
the general vicinity of Yarmouth on the west coast of the province they are so plenti-
ful that great numbers of giant tuna congregate there summer after summer to harry
and destroy the schools in the local tide rips.*! Thence eastward along the outer coast of
Nova Scotia they are abundant and generally distributed; witness a reported catch be-
tween the vicinity of Cape Sable and Cape Breton of something more than 41 million
pounds in 1946, and 21 million pounds in 1956, the latest year for which pertinent
statistics are readily available.—uH. B. B.
There are fewer, it seems, along the Nova Scotian shore line of the Gulf of St.
Lawrence; landings there in 1946 were 1,330,000. But thence westward these fish
are numerous again throughout the southern side of the Gulf,?* around Anticosti
Island (described there as in ‘“‘bancs enormes’’; 716: 285) and along the Gaspé shore
in the lower part of the St. Lawrence estuary. But they are progressively fewer up the
1g. “With the favourable climatic conditions in Arctic regions, the Herring has spread to Jan Mayen, Spitzbergen,
Nowaja Zemlya and Kara Sea” (Jensen, 67:64).
20. For details as to their regional and seasonal occurrence in the Gulf of Maine, see Bigelow and Schroeder (15:93).
21. The annual “Tuna Tournament” is held at Wedgeport, Nova Scotia, not far from Yarmouth.
22. 1946 catches, 36,103,600 for the New Brunswick shore of the Gulf; 6,080,100 for Prince Edward Island; 15,078,000
for the Magdalens; 15,525,000 for Bonaventure and Gaspé counties, Quebec.
Fishes of the Western North Atlantic 291
St. Lawrence, and we find no report of them beyond Trois Pistoles, Quebec, about
opposite the Saguenay River. Although the area of abundance does not extend to the
northern shore of the Gulf, they are widespread there as well as in the Strait of Belle
Isle; the 1946 catch for this entire coastline was only 923,700 pounds. They have
been reported also for the banks in the northeastern corner of the Gulf between New-
foundland and Labrador (125).
They are so plentiful along the Newfoundland shores of the Gulf that they have
long supported important fisheries in and off Bay of Islands and at Bonne Bay. Indeed,
“it seems proven ... that practically all the Newfoundland bays are frequented by
spawning herring often in great numbers” (Thompson, 125: 36). Thus Fortune and
Placentia bays on the eastern part of the southern coast have been the sites of produc-
tive Atlantic Herring fisheries since the days of the early cod fishery; the local catch
is used chiefly as cod bait.
The reported commercial catch of Atlantic Herring in 1946 for Newfoundland as
a whole was only about 65 °/, of that for the southern part of the Gulf of St. Lawrence
on the Canadian side’. This, taken at face value, suggests that the cold waters of eastern
Newfoundland produce fewer Atlantic Herring than the waters of the south and west
coasts. However this may be, they are at least moderately plentiful on the east coast
of Newfoundland right up to the entrance of the Strait of Belle Isle; Jeffers not
only reported them for Rahleigh but stated that they were numerous enough at Quirpon
to supply the local inhabitants with winter food for their sledge dogs (66: 206). They
have also been reported for the northern side of the Strait at Blanc Sablon (r2r: 124),
and repeatedly for the Strait without specified locality.
But a review (perhaps not sufficiently searching) of the various lists of Labrador
fishes has not yielded any reference to their presence anywhere to the north of the
Strait, apart from Stearns’ report that they are abundant “‘north of Blanc Sablon,
growing more and more so all along the Labrador coast, the farther down which are
the greatest catches” (12I: 124).
The offshore boundary of their range follows in general the break in slope at the
outer edge of the American continental shelf. Schools are occasionally seen at the surface
or are picked up by trawlers here or there on Georges and Browns banks, fronting the
Gulf of Maine (75: 95). It seems the situation is similar on the Nova Scotian Banks
eastward to Banquereau and on the Newfoundland Banks, where they are often taken
by otter trawlers in quantities (125: 36). All this has long been common knowledge
among the banks fishermen. During the early years of the Georges Bank cod fishery,
it was common practice to set herring drift nets for bait (16: 102). And it may well
prove that they are much more plentiful on the banks than is now realized, for while
the numbers reported (15: 95, for Georges Bank) have been insignificant, no
serious attempt has yet been made to sample the offshore population with drift nets,
midwater trawls, or bottom trawls of a kind adapted to the capture of this partic-
ular fish.
23. Data on Canadian catches are published annually in “‘Fisheries Statistics of Canada.”
292 Memoir Sears Foundation for Marine Research
It has been known since Fabricius’. day (34) that there were Atlantic Herring in
western Greenland waters, in small numbers. They are now known to be widespread
along the southwestern and western coasts, northward at least to 72°30’ N, though not in
any great numbers if judged by the standards of northern Europe or the American
coast. Their eggs have also been found in the Julianehaab district. The spawning
extends through August and into September, and the presence of young fry is proof
of successful local reproduction. In 1932 Hansen reported the presence of schools on
the eastern coast in the neighborhood of Cape Walloe and Grittenfeldts Island (53: 42);
he also reported the capture of a specimen near Angmagssalik. The evidence, in short,
is strong that this species has expanded its range in Greenland waters during the past
30 years or so, as have other boreal sub-Arctic fishes (for details as to their occurrence
in Greenland waters, see Jensen, 67: 61-64).
Synonyms and References :*4
Clupea harengus Linnaeus, Syst. Nat., ed. 10, 1758: 317 (diagn.); Cuvier and Valenciennes, 20, 1847: 30,
pl. 591 (descr., anat., life hist., distr., etc., includ. review of lit.); Giinther, Cat. Fish. Brit. Mus., 7,
1868: 415 (refs., descr., Europe and N. Amer.); Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1882:
265 (descr.); Goode, e¢ a/., Fish. Fish. Industr. U.S., 1 (3), 1884: 549-568, pl. 204 (abund., import.,
distr., migr., reprod., food, capture, uses); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896:
421; 47 (4), 1900: fig. 18 (descr., synon.); Smith, Bull. U.S. Fish Comm. (1897), 77, 1898: 91,
(spawn., growth rate, Woods Hole, Massachusetts); Linton, Bull. U.S. Fish Comm. (1899), 19,
1901: 437 (food, parasites); Bean, 7th Rep. Forest Comm. N. Y. (1gor), 1902: 298, with fig. (econ.
import., spawn., range); Bean, Bull. N.Y. St. Mus., 60, Zool.9, 1903: 192 (synon., refs., descr.,
import., abund., spawn., distr.); Fowler, Rep. N. J. St. Mus. (1905), 1906: 93, fig. (diagn., New
Jersey); Smith, N.C. geol. econ. Surv., 2, 1907: 120 (descr., range); Ehrenbaum, Eier und Larven Fisch.,
Nord. Plankt., 2, 1909: 361 (develop. of eggs and larvae); Sumner, Osburn, and Cole, Bull. U.S.
Bur. Fish. (1911), 3Z, 1913: 741 (refs., season. abund., spawn., food, parasites, Woods Hole, Massa-
chusetts); Cox, Contr. Canad. Biol. (1914-16), 1916: 81 (disease, Gulf of St. Lawrence, 1914); Meek,
Migrations of Fish, 1916: 67-88 (genl. acct.); Regan, Ann. Mag. nat. Hist., (8) 19, 1917: 227 (diagn.
in key); Lebour, J. Mar. biol. Ass. U. K., N. S. 72 (3), 1921: 459 (food of young clupeoids); Bigelow
and Welsh, Bull. U.S. Bur. Fish. (1924), 40, 1925: 92, fig. 40 (descr., range, breed. habits, growth,
size, season, Movements, food, enemies, destruct. by natural causes, annu. fluct., Gulf of Maine); Jordan,
Fishes, 1925: 275 (distr., abund.); Hildebrand and Schroeder, Bull. U.S. Bur. Fish. (1927), 43, 1928:
81 (refs., descr., spawn., import., rare in Chesapeake Bay); Breder, Field Bk. Mar. Fish. Atl. Cst.,
1929: 64, fig. (range, import., habits, size); Jordan, Manual Vert. Anim. NE U.S., 1929: 39 (diagn.,
range); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 41 (range, synon.);
Perlmutter, 28th Rep. N. Y. Consery. Dept., Suppl., 15 (2), 1939: 17 (spawn., eggs, young caught,
Long Island, New York); Greeley, 28th Rep. N. Y. Conserv. Dept., Suppl., 15 (2), 1939: 82 (Long
Island, New York); Bigelow and Schroeder, Bull. U.S. Bur. Fish., 48, 1940: 327 (range extension;
catch on Georges Bank; distr. of young); Warfell and Merriman, Bull. Bingham oceanogr. Coll., 9 (2),
1944: 51 (near New Haven, Connecticut; sizes and spawn.); Anonymous, Fish. Resources U. S.,
Senate Doc. No. 51, 1945: 61-62, with figs. (commerc. import., range, life hist., fluct.); Bigelow and
Schroeder, Fish. Bull. (74) U.S. Fish Wildl. Serv., 53, 1953: 88-100 (deser., life hist., Gulf of Maine).
24. Numerous accounts pertaining wholly or in part to this Herring have been written, for it has been a principal
subject of extensive study for centuries. A complete bibliography would occupy many pages. Therefore, the refer-
ences given herein are limited to the papers actually quoted (directly or indirectly), to taxonomic works of a general
nature, or to faunal lists of the American coast. Many of these also contain bibliographies. Furthermore, references
are given to publications containing descriptions of the species itself and of several “species” now considered synony-
mous with Clupea harengus. For additional references bearing especially on the biology of this species, see
Scattergood (rr4).
Fishes of the Western North Atlantic 293
Clupea elongata LeSueur, J. Acad. nat. Sci. Philad., r (2), 1818: 234 (orig. descr.; type local. Marblehead
and Sandy Bay, Massachusetts).
Herring, Moore, Rep. U.S. Comm. Fish. (1896), 22, 1898: 387—442 (life hist., econ. import., Passamaquoddy
Bay); Huntsman, Canad. Fish. Exped. (1914-1915), 1919: 165-171 (growth of young); Lea, Canad.
Fish. Exped. (1914-1915), 1919: 75-165 (age and growth, Canad. waters); Norman, Hist. Fishes,
1931: 42, 65, 80, gI, 135, 183, etc. (genl. acct.); Anonymous, Rep. U.S. Tariff Comm., (2) 69, 1933:
78, 106, 117, 143, 149-151, 157, 161, 163-165, 218 (statist. fishery products, Europ. and Amer.
countries); Ford, J. Mar. biol. Ass. U. K., N.S. 19, 1933-34: 305 (econ. of fishery, Plymouth, Eng-
land; life hist. data; forecast. catch); Battle, Huntsman, Jeffers, Johnson and McNairn, J. biol. Bd.
Canad., 2 (4), 1936: 401 (fatness and food of young, Passamaquoddy Bay); Marshall, Nicholls and
Orr, J. Mar. biol. Ass. U. K., N.S. 22, 1938: 245 (growth, feed., Clyde); Johnson, J. Fish. Res. Bd.
Canad., 4, 1939: 349, 392 (effects of light on movements, feed.). Leim, Tibbo, and Day, Bull. Fish.
Res. Bd. Canad., 3, 1957: 1-184 (investig. e. Canada, growth, abund. populations, year-classes).
Doubtful References:
Clupea halec Mitchill, Rep. in part, Fishes of New York, 1814: 20 (orig. descr.; type local. New York; type lost;
possibly a species of Pomolobus); Mitchill, Trans. Lit. philos. Soc. N. Y., 7, 1815: 451 (descr.).
Clupea coerulea Mitchill, Trans. Lit. philos. Soc. N. Y., I, 1815: 457 (orig. descr.; type local. New York;
type lost; possibly a species of Pomolobus).
Genus A/osa Link 1790
Shads
Alosa Link, Mag. Physik. Naturg., Gotha, 6 (3), 1790: 35; type species not specified; Cuvier, Régne Anim.,
ed. 2, 2, 1829: 319; type species C/upea alosa Linnaeus, same as A/osa Link.
Generic Synonym:
Alausa Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 389; type species 4/ausa vulgaris Cuvier and
Valenciennes equals C/upea a/osa, apparently a variant spelling of 4/osa Link.
Characters. Bopy compressed, rather deep to elongate. Bony scures rather promi-
nent, with sharp points; the chest and abdomen with a sharp edge. Curek (bone
below eye) notably deeper than long (Fig. 69). Mourn large. Maxitiary generally
extending beyond middle of eye. Upper jaw with a definite median notch. Lower jaw
largely included in upper one, its upper margin within the mouth rising gradually
(Fig. 69). TEETH in jaws minute or absent (absent in American adults); none on vomer
or tongue in adults. Dorsat with 16-22 rays, its last ray not produced, its origin
nearer to end of snout than to base of caudal. Anat with 18-27 rays (18-22 in Amer-
ican species), the last ones not enlarged. Prtvic usually with 9 rays, inserted under
base of dorsal. VerRTEBRAE about 52-59.
Remarks. This genus is very close to Pomolobus, with some of the species being
difficult to separate generically. The species herein assigned to A/osa have a deep
median notch in the upper jaw; the lower jaw shorter than the upper, with the upper
margin (within mouth) not rising abruptly; the cheek deeper than long; and the adults
without teeth. A combination of these characters is required to identify the species of
Alosa.*®
25. The two “wing-like plicae of skin, covered with elongate scales (a/ae)’’ on the base of the caudal, designated the
leading diagnostic character by Borodin in his revision of the genera Alosa and Pomolobus (18:175), seem unusable
because too often they are indistinct on preserved specimens.
204 Memoir Sears Foundation for Marine Research
Ficure 69. Heads showing shape of mandible and cheek. rert, 4/osa sapidissima, 93 mm TL, Clifton Beach,
Maryland; ricut, Pomolobus aestivalis, 82 mm TL, Smith Point, Maryland. Both drawn by Louise Nash.
Range. Two species occur off Europe, another off northern Africa, and two off
North America, where the recorded range extends from southeastern Newfoundland,
the St. Lawrence River, Quebec, and northern Nova Scotia southward to Indian River
City, Florida, and the Gulf of Mexico westward to the Mississippi River.
Key to North American Species
1a. Lower limb of first gill arch with 59-73 rakers (26-43 in young under 125 mm).
sapidissima (Wilson) 1811, p. 295.
1b. Lower limb of first gill arch with 42-48 rakers in adults (27-33 in young 48—
go mm long). alabamae Jordan and Evermann 1896, p. 308.
Ficure 70. First branchial arch, showing numerous gill rakers: tert, A/osa sapidissima, 550 mm TL, USNM
20969; rIcHT, 4. alabamae (with fewer rakers), 390mm TL, type, USNM 47689. Both drawn by Ann
S. Green.
Fishes of the Western North Atlantic 295
Alosa sapidissima (Wilson) 1811
American Shad, Atlantic Shad, Shad, Common Shad, White Shad
Figures 69-74
Study Material. At least 75 specimens, 35-580 mm TL, 29-475 mm SL, from
St. Andrews Bay, New Brunswick; Flag Island, Eastport, and Portland, Maine;
Holyoke and Woods Hole, Massachusetts; Noank, Connecticut; Long Island, and
Gravesend Bay, New York; Great Egg Harbor, New Jersey; Delaware City, Delaware;
Figure 71. Alosa sapidissima, adult male, 480 mm TL, 19.13 inches, Norfolk, Virginia, USNM 25099.
Drawn by H. L. Todd.
many places in Chesapeake Bay and tributary streams; Avoca and Beaufort, North
Carolina; and New Berlin, Florida.
Distinctive Characters. For the features that distinguish the true shads (4/osa)
from their close relatives, and for the difference between sapidissima and alabamae, see
Key to Species, p. 294, as well as pp. 293, 304, 309, 310.
Unlike the adult with its toothless jaws and a median indentation in the upper jaw,
young of 150 mm and less lack the indentation on the upper jaw and have minute teeth
in the jaws. Therefore the jaw and tooth characters of the adult cannot be used in
identifying the young. However, the deep narrow cheek as well as the slender mandible
with its upper margin not rising abruptly to form an angle in the mouth will aid in
separating young sapidissima from young and adult alewives of the genus Pomolobus
(Fig. 69).
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens 29-475 mm SL.
Body: depth 30.2—36.8. Head: length 23-28.
Caudal peduncle: depth 7.7—10. Snout: length 4.85—-7.1.
296 Memoir Sears Foundation for Marine Research
Eye: diameter 3.8—5.7. Scales: 52-62.
Interorbital: width 4.55—5.95. Ventral scutes: 35-38.
Maxillary: length 11-14. Gill rakers: 59-73 in adults.
Pelvic fin: length 9.9-11.9. Fin rays: dorsal 17-20; anal 20-23; pec-
Pectoral fin: length 15.3-17.7. toral 15-17.
Vertebrae: 55-57 (4 specimens), 54-59
given in literature.
Bopy slender, compressed, its greatest thickness generally about half of its depth;
the depth very variable, increasing with age, 2.75—4.0 in SL.
ScaLes moderately adherent, with crenulate membranous borders, 15 or 16 rows
between base of pelvic fin and anterior dorsal rays. VENTRAL scutes well developed,
20-22 in front of pelvics, 14-17 behind them.
Heap 3.6—4.4 in SL. Snout 3.6—4.6 in head. Eye 4.0—5.9. INTERORBITAL 4.2—
5.8. Mourn moderate, oblique; mandible included. Maxituary broad, its width about
half of length of snout, with a broad supplemental bone, rounded posteriorly; reaching
to middle of eye in young, generally to below posterior margin of eye in adults, 1.9—2.2
in head, GILL RAKERs increasing in number and proportionate length with age, those at
angle shorter than snout in young but longer than snout in large specimens, 26-43
on lower limb in young 29-125 mm SL, 59—73 in specimens 300 mm SL and upward.
TEETH missing in adults though present on jaws and median line of tongue in young.
Dorsat fin somewhat elevated anteriorly, its margin slightly concave, its last ray
a little longer than the preceding one, its origin an eye’s diameter or less in advance
of pelvic fin and about equidistant between tip of snout and vertical from base of last
anal ray. Caupat rather deeply forked, the lobes of nearly equal length and somewhat
shorter than head. Anat low, its origin about an eye’s diameter nearer to base of
pelvic than to base of caudal. Petvic near ventral edge, about equidistant between
base of pectoral and origin of anal, 2.1-2.6 in head; the axillary scale equal to, or a
little more than, half of length of fin. Pecrorat a little higher on side than pelvic,
1.4—1.65 in head.
Color. Fresh specimens greenish to bluish with metallic luster above, gradually
shading into the bright silver on sides. Dark spot on shoulder, sometimes followed
by smaller spots or dots, rarely with a second parallel row of dots below the first. No
dark lines along rows of scales. Fins pale to greenish, the dorsal and caudal somewhat
dusky in large examples, and tips of lobes of caudal dark in some specimens. Peritoneum
mostly pale to silvery.
Size. This species is the largest of the herrings. Females attain a maximum
length of about 600 mm (30 in.) and a weight of 12 pounds in the Atlantic. On the
Pacific coast, where the species was introduced, the maximum weight is even greater;
females of 14 pounds have been reported. In the Chesapeake Bay area the average
weight (and length) of females (roe shad) is probably somewhat under six pounds
(about 57.5 cm; 22.75 in.), that of males (buck shad) about 3.5 pounds (about 50 cm;
20 in.).
Fishes of the Western North Atlantic 297
Development and Growth. The eggs, about 3.5 mm in diameter after they are 1m-
pregnated, are demersal, but they are only slightly heavier than the water in which
they are deposited. As they are not adhesive, they drop loosely and singly to the bottom.
According to Ryder they hatch in 52 hours at an average temperature of 57.2° F, and
in less than 36 hours at an average of 74°F (z12: 796). However, a longer incubation
period was reported by Leim (81: 31), who said that eggs held under artificial conditions
hatched in 12-15 days at 12°C (53.6°F) and in 6-8 days at 17°C (62.6°F).
The newly hatched fish, transparent and extremely slender, are about 9-10 mm
Ficure 72. Alosa sapidissima egg development. Lerr to RIGHT: newly laid egg; embryo in natural position
within enveloping membrane of egg (photograph); egg with embryo nearly ready to hatch. After Leach.
long. The yolk is absorbed in four or five days at a water temperature of 62.5°F and
in about seven days at 53.5°F; the fish have then attained a length of about 12-
15.5 mm (81: 33). In some preserved specimens that show the yolk almost absorbed,
the depth is only about 0.05 of the length, the large dark eyes protrude prominently,
the dorsal fin fold is somewhat developed, and the alimentary canal is discernible,
ending under the beginning of the posterior sixth of the body. During development,
the vent moves forward as in other Clupeidae. The fins are fully developed in 21-
28 days at a length of about 20 mm, when metamorphosis is nearly complete (16: 118).
For a more detailed account of the development of the larvae, see Leim (81: 31-43).
The young remain more slender than the adults until they are nearly grown
fish, but even among sexually mature fish there is great variation in depth, seemingly
not related to sex and only partly related to spawning conditions. In 25 young, 35-—
125 mm long, the depth varied between 26.5-30 °/) of SL, and in a lot of 12, 460—
505 mm long, the depth was 32-48 °/, of SL.
A gradual increase in the number of gill rakers, as already indicated, also takes
place with age and growth. For example, the number on the lower limb of the first
arch varied as follows: in 18 young 35 to 65 mm long, 26 to 31; in 21 examples
am EEL LE ELEL oS
= ee ; SS EEE
: es SK
1 fab Lire Soe ———
Hl LLL LOLTAL LLL LEE LITT ELY (LE EL LEE
MOE E
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Ficure 73. A/osa sapidissima, iarval development. top, immediately after hatching; upPER CENTER, third day
after hatching; LOWER CENTER, five days after hatching; Borrom, 17 days after hatching. After Leach.
70 to 125mm, 34 to 43; in 16 immature fish 190 to 270mm, 48 to 62; and in
32 sexually mature examples 465 to 580mm, 59 to 73.
The young grow rapidly during their first summer while in fresh water. The
total lengths of 769 young taken in the Chesapeake Bay area—mostly from the
Potomac River (in 1912)—ranged between 20-69 mm during June, 30-74 mm during
July, 45-79 mm during August, 50-95 mm during September, 45-99 mm during
October, and 60-119 mm during November (59: 98).
Larger young of 150, 175, and even 225 mm, at an age of seven months, have been
reported. Such fish, in at least one instance, are known to have lived under specially
favorable conditions in a pond well supplied with food (at Washington). Furthermore,
according to Bean: “Nets set off shore in Gravesend bay [New York] in the fall frequently
inclose large quantities of young shad, sometimes a ton and a half at one time,
during the migration seaward... The fish are usually about 6 to 8 inches long” (8: 208).
One may question whether fish of such a large size, taken in such large quantities,
were only six to seven months old. That the young grow faster north of Chesapeake
Bay seems unlikely in the light of a statement by Bigelow and Welsh (z6: 118) con-
cerning this Shad in the Gulf of Maine: ‘The young shad remain in the rivers until
fall when, at a length of 11/, to 41/, inches (37 to 112 mm) and resembling their parents
Fishes of the Western North Atlantic 299
in appearance, they move down to salt water.”” This range in length is close to that
of fish of the same age from Chesapeake Bay and does not indicate more rapid growth
in the more northern part of the range. However, Bigelow and Welsh added: “‘In the
rivers of Maine the fry may be as long as 5 to 7 inches (125 to 175 mm) by the first
week of November.” Included in Hildebrand and Schroeder’s data there were three
specimens from the November collection that ranged somewhere between 145 and
244 mm in length (59); these fish were so much larger than the other 137 specimens
taken in the same month that one is led again to suspect that they may have belonged
to the one-year class.
ww
XS
Ficure 74. Alosa sapidissima, young, 47 mm TL, 1.88 inches, Beaufort, North Carolina. Drawn by Louella
E. Cable.
The growth rate after the young have entered salt water is not well known because
of the lack of specimens available for examination. Hildebrand and Schroeder listed
22 specimens that were spending their first winter in the deeper places of Chesapeake
Bay instead of continuing their migration to sea (59: 98). Eight of these, taken in
January, ranged between 95-124 mm and averaged 108 mm, 3 taken in February
were 95, 105, and 135 mm long, ro taken in March ranged between 105-159 mm and
averaged 142 mm, and 2 taken in May were 145 and 154 mm long. Specimens of a
somewhat similar size were dredged during the winter in the estuary of Newport River,
Beaufort, North Carolina; 18 specimens caught in January ranged between 73-140 mm
and averaged 117mm; 14 caught in February were 100-160 mm and averaged
121 mm; and 3 caught in March were 84, 102, and 120 mm long.
Using mostly specimens from the Connecticut River, Borodin®* (17: 183) deter-
mined the age of older fish from the annuli and transverse grooves on the scales, a
method confirmed as practicable in Barney’s study of otoliths (5: 168). Borodin
provided a list of 10 size groups to which he assigned ages as follows: Group 1 — 58
26. Borodin, in determining the age of Shad from the scales, used the transverse grooves in addition to the annuli,
having discovered that two complete transverse grooves are formed each year. Barney determined their age from
the “‘limy deposits” in the otoliths, which form annual rings; he found that these annuli correspond to the annuli
and crossgrooves on the scales used by Borodin.
300 Memoir Sears Foundation for Marine Research
to 113 mm long, 26 specimens from the river, all less than a year old; Group 2 — 145
to 180 mm long, 4 from salt water, in their second year; Group 3 — 210 to 290 mm,
7 from salt water, in their third year; Group 4 — 300 to 350 mm, Io young males
entering the river, in their fourth year; Group 5 — 360 to 390 mm, 22 young males
entering the river, in their fifth year; Group 6 — 400 to 430 mm, 13 males and 9
females, in their fifth, sixth, and seventh years; Group 7— 440 to 480 mm, 34 males
and 9 females, in their sixth, seventh, and eighth years; Group 8 — 490 to 520 mm,
28 males and 13 females, in their seventh, eighth, and ninth years; Group 9 — 530 to
570 mm, 6 males and 43 females, in their seventh, eighth, ninth, and tenth years; and
Group 10 — 580 to 660 mm long, 1 male and 23 females, in their eighth, ninth, tenth,
and eleventh years.
Young fish from the Bay of Fundy reach a length of 41-45 mm in nine to ten
weeks (81: 74-84). Leim’s calculations, based on scale readings of adult fish, in general
indicate a rather faster rate of growth for his fish than for those of Borodin.
Some of the males, according to Borodin, enter the Connecticut River from the
sea when they are 300-350 mm long and in their fourth year, as shown in the preceding
data. Although the author did not say so, these males undoubtedly were sexually
mature, as migration to the rivers seems to be carried out solely for the purpose of spawn-
ing. The smallest females that returned to the Connecticut to spawn, according to the
same author, were between 400 and 430 mm long and in their fifth, sixth, and seventh
years. Therefore, the males seem to become sexually mature at an earlier age than the
females. On the other hand, females may reach a greater age; among the older fish ex-
amined by Borodin, determined by him to be in their seventh, eighth, ninth, tenth, and
eleventh years, there were only 7 males compared to 86 females. Concerning the Bay
of Fundy fish, Leim said: ‘Most of the shad present on the spawning grounds were
five years old or over and eight or nine years seems to have been the maximum age
represented. Fish under four years of age apparently do not spawn at all” (8r). The
discrepancy in the rate of growth and the age of spawning fish, as determined by
Borodin and by Leim, may have resulted from a difference in the interpretation of the
scale markings, or from a difference in the behavior of two rather widely separated
races that may respond differently to the climatic or general environmental condi-
tions in which they live.
Reproduction. The average number of eggs produced by a single fish varies between
2 §,000—30,000 (11g: 127). In the Manual of Fish Culture (2: 125) it is stated that the
average number is not more than 30,000 but that a single fish has been known to yield
between 60,000 and 115,000 eggs. In 1885, one from the Delaware River yielded
156,000. In this species, as among others, large fish as a rule produce many more
eggs than small ones.
If all of the large number of eggs produced by a female hatched and reached ma-
turity, the progeny would be exceedingly great. However, it is quite certain that nor-
mally only a small proportion of the eggs produces fish that live to maturity. The most
critical time in the life cycle apparently comes when the larvae have absorbed the yolk
Fishes of the Western North Atlantic 301
and must find their own food. If food is found and if other conditions are favorable for
survival, then the spawning season is a successful one. The success or failure of a spawn-
ing season in a river is usually evident from the size of subsequent catches when the
fish hatched in a given year return later to spawn in the same stream. The comparatively
large fluctuations in catches over a period of years no doubt result in large part from
the measure of success of the particular spawnings from which the fish composing the
catches were derived.
Spawning and Migrations. American Shad spawn in fresh water. Some spawn
as soon as they arrive from the sea while others do not spawn until they reach the
upper tributaries of rivers. Virtually all spawning, in this species as in many others,
takes place in the evening between five and ten o’clock. While spawning, the fish are
paired, swimming close together side by side. Since no nests are made, the eggs fall
loosely in the water, and doubtless many are smothered when they fall on muddy bot-
tom. An individual returns to spawn year after year.
The water temperature governs the date when mature American Shad approach
the mouths of the rivers to spawn, about 50-55° being required. Accordingly, they
appear off the Florida coast (St. Johns River) in November, to remain until March,
in the sounds along the Carolina coast in February and March, in the Chesapeake
Bay area in March and April, and farther north in May and June. But the precise
date of arrival at any particular locality along the coast may vary by a couple of weeks
from year to year, depending on the variation from year to year in the vernal warming
of the water.
In large rivers, they. run far upstream if their passage is not blocked. Thus they
still run up 375 miles (or did recently) in the St. Johns River in Florida, 300 miles in
the Altamaha in Georgia, about 200 miles in the St. John in New Brunswick, and at
least 200 miles in the St. Lawrence (122: 11; 127: 129, 130). American Shad as-
cending the estuary of the St. Lawrence may travel as much as 25-50 miles in 24
hours (127: 129).
The spent fish, very thin after spawning, begin their return journey to the sea
immediately afterward; i.e. in March to May in the southern part of the range and
in June to August in the northern part. But the young fry (product of that season’s
spawning) remain in the parent stream until autumn, when the temperature falls to
60° or lower. In the Chesapeake Bay region, their migration out into salt water takes
place principally in October and November; but it is not until late November or De-
cember that all of them are in salt water.
Southward from Cape Cod the spent fish vanish upon re-entering salt water. While
a few young fry in the Chesapeake Bay area spend their first winter in the deeper
parts of the Bay, those from the Hudson may linger nearby for an entire year, for
catches of 6- to 8-inch fish in Gravesend Bay in autumn have been reported (8: 208).
The larger sizes of immature fish vanish as completely from the southern part of the
coast as do the spent fish, until they reappear there for their first spawning.
Spent fish up to ten pounds and averaging about five pounds have been reported
302 Memoir Sears Foundation for Marine Research
as taken in the Gulf of Maine, and schools of them have been seined from time to time
in summer and autumn, even in December, at various places along the Maine coast,
where they have been the object of a frozen-fish industry in some years. Immature
fish (0.5-2.5 pounds) sometimes congregate also along the Maine coast and are more
or less common in the Massachusetts Bay region.
Clearly these Gulf of Maine fish are far too numerous to be derived only from
the streams of the Bay of Fundy, as was formerly considered likely (16: 116). Recently,
extensive tagging experiments (75: 111, ftn. 18-20; 124) have shown that, while
some of them come from as far south as the Altamaha River, Georgia, others come from
as far north as the St. Lawrence estuary, both of these contingents to fatten in the
Gulf. By the end of December, however, they have vanished, not to reappear anywhere
on the coast until the onset of the next spawning season, when the maturing fish return
to the rivers of their native watersheds.
Like salmon and alewives, American Shad spend most of their life in the sea and
make most of their growth there, but where they winter remains to be discovered.
While in the sea, Shad are schooling fish, often in the thousands, and they never re-
enter fresh water until they return to spawn, although they may appear in brackish
estuaries. They have been taken 25-90 miles out off southern New England, 110 miles
out on Georges Bank, 40-50 miles out off the coast of Maine, and 50-60 miles off
eastern Nova Scotia, and as deep as 50-68 fms.
Food. The diet of 14 young fish, 43-60 mm long, from fresh water of the
Potomac River, consisted chiefly of adult and larval insects and ostracods, and in
one instance a small fish (59: 95). Larger juveniles, 100-150 mm long, caught in
Chesapeake Bay, presumably while en route to the sea, had fed mostly on Mysis, although
one stomach contained fragments of a small fish, and another fragments of plant tissue.
Working with young specimens from the Shubenacadie River, tributary to the
Bay of Fundy and its estuary, Leim found that the first food taken by larvae 11 mm
long consisted of midge larvae (Chironomidae), while the somewhat larger larvae had
fed principally on mature and immature copepods (81: 16). In fact, these organisms
constituted the chief food of the young up to the time of transformation, with the
relative abundance of these forms in a particular locality determining which food pre-
dominated. Dr. Leim’s data show also that young adults taken in the same general
vicinity continued to subsist principally on these same organisms. Other foods ingested
consisted of ostracods, amphipods, insects, and fish.
Little or no food has been found in the stomachs of those caught while in fresh
water en route to their spawning grounds, indicating that these fish, like salmon, do
not ordinarily feed then. However, there are some records showing that adults occasion-
ally do take food while in fresh water, at least late during the spawning season. They
will often take a live minnow or an artificial fly when working upstream on their
spawning run.
Many kinds of food have been found in the stomachs of those taken in salt water
in estuaries and bays north of Cape Cod. But southward, examples are rarely obtainable
Fishes of the Western North Atlantic 303
in salt water, except during the migration to and from the spawning grounds. There-
fore it is of interest that an adult female, caught in Chesapeake Bay in December,
was gorged with parts of plants, among which were fragments of a mollusk shell. It
has been reported also that adults taken in the vicinity of Savannah, Georgia, had fed
on plants. Others caught near Fort Lee, New Jersey, had fed liberally on young shrimp,
and still others, taken in Delaware Bay, had eaten small fish.
From an examination of about 350 stomachs of both mature and immature fish
caught in the salt water of Scotsman Bay, Bay of Fundy, Leim found that, while cope-
pods constituted the chief food of the smaller ones, as in fresh water, these crustaceans
were unimportant in fish 400 mm and more in length (87: 68). Mysids, which were
sparingly eaten by small fish, were the chief food of adult fish. In general, about 90 °/,
of the specimens of all sizes from that area had eaten copepods and mysids, with
ostracods, amphipods, isopods, decapod larvae, insects, mollusks, algae, fish eggs, and
fish making up the remainder. After examining many stomachs of specimens taken in
the Bay of Fundy, Willey also concluded that the chief foods consisted of copepods and
mysids, to which he added shrimp and larval stages of barnacles (133: 313-320).
Stomachs of fish from the Gulf of Maine have been found full of copepods (chiefly
Calanus). American Shad also feed greedily on pelagic euphausiid shrimps.
Enemies. No doubt the larger predatory fishes are the principal enemies of the
young while the latter are in the rivers. Although no records at hand show definitely
that this is so, it can scarcely be otherwise, for the young are not rapid swimmers and do
not ordinarily seek protection among plants or other cover. Furthermore, they are small
enough, at least during the early part of summer, to be swallowed by bass and other
carnivorous fishes only an inch or two long. Water birds, turtles, and water snakes
no doubt feed on them too. The toll taken by predatory fishes while the young are en
route to the sea may not be very great, for this migration takes place in the fall when
many marine fishes (at least in Chesapeake Bay) have left the shallow waters. At sea
they doubtless fall prey to all the larger predaceous fishes, and perhaps to finback whales
as well.
The chief enemy of the adults is man, who has not only depleted the stock in many
rivers through overfishing but has cut them off from spawning areas through the
construction of barriers and by polluting the waters with deleterious wastes.
Parasites. The parasites of J. sapidissima have been made known principally through
several papers by Edwin Linton and by Charles B. Wilson, based mostly on specimens
obtained from fish caught in the vicinity of Woods Hole, Massachusetts. The findings
were summarized by Sumner, ef a/. (123: 742) as follows: “Acanthocephala (Linton)
Echinorhynchus acus. Nematodes (Linton) — Ascaris adunca, Ascaris sp. (immature).
Copepods (C. B. Wilson) — Caligus rapax.” To this list was added the copepod Ler-
naeenicus radiatus (135: 59). Working with both young and adults from the Shuben-
acadie River and Scotsman Bay, Leim found no external parasites and only three
internal ones, namely “‘distomes, nematodes, and Acanthocephali” (81: 74). It is not
stated whether any of these parasites were considered seriously detrimental to the host.
304 Memoir Sears Foundation for Marine Research
Relationships. There are two well-marked species of this genus in Europe, 4. alosa
and A. finta, both of which have the deep cheek and notched upper jaw as in American
species. However, 4. a/osa has smaller scales (72-80 lateral series) and more numerous
gill rakers (72—86 on lower limb in adults) than the American species. 4. finta, while
agreeing with the American species as to scales, has only about 24-32 gill rakers on
the lower limb, in grown examples.
The two American species recognized here, namely 4. sapidissima and A. ala-
bamae, differ principally in the number of gill rakers; mature examples of sapidissima
have 59-73 on the lower limb whereas specimens of a/abamae of similar size have
only 41-48. The relationship of these species is discussed further in the account of
alabamae (pp. 309, 310).
Variations. The specimens examined, from various localities within the range
(p. 295) and used in preparing the Description, do not differ specifically or subspecifically.
However, racial differences have been recognized. Vladykov and Wallace have found
slight average differences in the number of vertebrae, pectoral rays, and ventral scutes
between examples from the Bay of Fundy, Delaware Bay, and Chesapeake Bay; these
they regarded as different “populations” (130: 52-66). Investigators of the U.S. Fish
and Wildlife Service, in a special study of 4. sapidissima, found that virtually every
important river system within the range of this fish is inhabited by a different race
(unpublished data).
Commercial Importance. This species has been an important foodfish throughout
its entire range since the earliest settlers arrived, and no doubt these fish were taken
by the Indians long before. Unfortunately, pollution, overfishing, and dams constructed
across the streams (preventing the fish from reaching their spawning grounds) have
caused partial and sometimes serious depletion, or even virtual extermination. So
many streams have been dammed, particularly in New England, that, in the words
of Bigelow and Welsh: “the local stock of shad has diminished until now the Gulf of
Maine stock is but a shadow of its former abundance...” (16: 114). The great decline
in abundance in New England is evident from the published statistics of the former
United States Fish Commission, U.S. Bureau of Fisheries, and the present U.S.
Fish and Wildlife Service;?? the annual New England catch between 1889-1908
seems to have been well above 1.25 million pounds, by 191g it had fallen somewhat
below a million pounds, and between 1924-1940 it has fluctuated around a half
million pounds.
The situation with respect to the American Shad fishery in the Hudson is much
more encouraging, although depletion occurred in that river also; statistics from the
sources previously mentioned show that the catch in that river declined from 3.75
million pounds in 1901 to only 40,000 pounds in 1916 and to 43,000 in 1917; there-
after, between 1918-1935, the catch varied from 94,000 pounds (1924) to 847,000
27. No adjustment for the differences in the number of persons employed, the type and number of vessels, and the
different types of gear used from year to year has been made in the statistics given here and elsewhere. However,
it is highly probable that the “‘fishing effort’? for rooo pounds of fish, for example, has increased.
Fishes of the Western North Atlantic 305
pounds (1935); and in 1936-1941 it varied from 2,467,000 (1938) to 3,270,700
(1939). Although the catch in 1959 was only 1,171,212 pounds, very definite recovery
has been indicated. According to investigators of the U.S. Fish and Wildlife Service,
this recovery is ascribed partially to the elimination of pollution and partially to the
great decrease in fishing effort at the time when depletion made it unprofitable to
fish. The curtailment in fishing permitted the depleted population to reach the
spawning grounds, to return to the sea, and to spawn again and again. Ultimately
this resulted in the rehabilitation of the American Shad in this river system. It is
decidely encouraging to have evidence that a depleted fishery can be restored.
The American Shad fishery in the Potomac, though it has fluctuated greatly, has
also undergone a general decline in recent years, without evident recovery. The catches
in 1896 and 1901 were above 2.5 million pounds whereas in 1909 and 1915 they were
only three-quarters of a million pounds. Then from 1919 to 1923 they ranged above a
million pounds, reaching the impressive totals of somewhat more than two million
pounds in 1919 and over three million pounds in 1922; the latter is the greatest catch
on record. In 1924 the catch dropped to a little over half a million pounds, and since
then it exceeded a million pounds only in 1926, 1928, 1929, 1931, and 1933; in fact,
the catch exceeded two million pounds in 1928, 1931, and 1932. However, between
1934-1941 it has fluctuated only from 631,000 pounds in 1935 to 322,800 pounds in
1940, the smallest catch recorded. Although great fluctuations in the catches, and no
doubt in abundance, have occurred in the Potomac River since 1896, a definite decline
was not indicated by the statistics until 1934. Thereafter, a general decrease in the
catches has reached alarming proportions. In 1959 the catch was only 188,000 pounds.
A decline similar to that in the Potomac River has taken place in the entire Chesa-
peake Bay area, where, exclusive of annual fluctuations in the catch, the general trend
has been downward since 1930. According to statistics, this same situation also applies
to the South Atlantic states, and perhaps to a somewhat smaller extent to the fisheries
of the Middle Atlantic states.
The over-all yield of late has ranged between catches of 8,222,000 and 14,700,000
pounds for the United States (1940, 1945, 1950-1953), and between 1,545,000 and
2,444,000 pounds for Canada (1940, 1945, 1950-1952). The American Shad thus
remains very important economically, yielding the fishermen an annual return of over
a million dollars.
Formerly, when they were abundant, the market for them more limited, transporta-
tion slower, and refrigeration virtually unknown, a considerable portion of the catch
was salted. However, now nearly all of it is marketed in the fresh state. Johnson has
stated, ““During recent years the outstanding problem of wholesale dealers, insofar
as shad are concerned usually has been one of obtaining adequate supplies rather than of
disposing of the catch” (68: 15). In fact, the demand on the Atlantic coast has been
so great that the supply is augmented annually by shipments from the Pacific coast.
The chief markets on the Atlantic coast are New York, Philadelphia, Baltimore, and
Washington.
306 Memoir Sears Foundation for Marine Research
Shad are caught principally with pound nets, gill nets, and seines. Small quantities
are taken with fyke nets, otter trawls, purse seines, traps, and dip nets.
Introductions. In the late 1800’s, many introductions of the fry were attempted
by the former U.S. Fish Commission and U.S. Bureau of Fisheries, but all failed;
large shipments were liberated in the streams tributary to the Gulf of Mexico, in Great
Salt Lake, Utah Lake, and Bear Lake, Utah, and in the Colorado River at the Needles,
Arizona.
In the 1870’s, the New York State Fish Commission introduced numerous fry
in streams tributary to Lake Ontario and in some of the smaller lakes connected to
it. According to Smith (178: 193), 658,000 fry were introduced, and “‘the appearance
of adult shad promptly followed the introduction of young fish in the Genesee River,
and from that time on for a number of years the indications pointed to the probable
successful acclimatization of the species.’’ These fish distributed themselves over the
entire lake and were taken at irregular intervals from 1875 to about 1885; but few,
if any, were seen after 1890. Hubbs and Lagler reported: “‘In the Great Lakes rare,
and confined to the Lake Ontario Basin” (62: 27).
Remarkable success has resulted from stocking waters of the Pacific coast north
of Monterey, California. The first fry, about 12,000, were planted in 1871 in the
Sacramento River by the California Fish Commission, and additional shipments were
sent there by the United States Fish Commission between 1873-1880. On the northern
part of the Pacific coast, fry were introduced in the Willamette, Snake, and Columbia
rivers in 1885 and 1886. From these plantings the species has spread from California
to Alaska, and it has become important commercially from San Francisco northward;
the catch during recent years generally has exceeded two million pounds.
Artificial Propagation. This species received the attention of fish culturists as early
as 1848 when eggs were taken artificially and hatched. However, it was not until 1867
that a really successful apparatus was perfected. This contrivance was known as the
“Seth Green box,” a floating box modified after one used for hatching salmon eggs.
However, when used in tidal waters it was subject to accidents. Thereafter various devices
were brought forward in rapid succession, and finally (in 1882) the “universal jar” or
“McDonald jar,” a modification of the ‘Chase jar” used for hatching whitefish eggs,
was adopted by the U.S. Fish Commission and is still in use. Many millions of fry,
or newly hatched larvae, have been liberated in the streams along the Atlantic coast.
Range. The regular range of the American Shad extends from the lower St. Law-
rence River and Nova Scotian Banks as far south as Indian River City, Florida. The
most northerly record is for one taken in Bulls Bay near St. John’s, Newfoundland
(1: 125). On the Pacific coast it now ranges from about San Pedro, California, to south-
eastern Alaska (131: 221).
Synonyms and References:
Clupea sapidissima Wilson in Rees’s New Cyclopedia, 9, ca. 1811: no pagination (orig. descr.; no local.; probably
Philadelphia, Pennsylvania); Jordan and Gilbert, Bull. U. S. nat. Mus., 16, 1882: 267 (descr., range);
28. For further details concerning their artificial propagation and distribution, see 4 Manual of Fish Culture (2:133-158).
Fishes of the Western North Atlantic 207
Smith, Bull. U.S. Fish. Comm. (1890), 20, 1892: 193, pl. 30 (deposited in small lakes and streams
connected to Lake Ontario); Cheney, Rep. Forest Comm. N. Y. (1885), 1896: 125-134, color pl.
(commerce. import., statist., artif. culture, local spawn., migr.).
Clupea alosa Mitchill (not of Linnaeus), Trans. Lit. philos. Soc. N. Y., 1, 1815: 449 (descr., habits, size).
Alosa preastabilis DeKay, New York Fauna, Pt. 4, Fishes, 1842: 255, pl. 15, fig. 41 (orig. descr.; type local.
New York; cf. European shad).
Alausa sapidissima Gill, Proc. Acad. nat. Sci. Philad., 1861: 54.
Alosa
Shad,
sapidissima Uhler and Lugger iz Rep. Comm. Fish. Md., 1876: 157; 1876: 133 (descr., synon., food
qualities, Maryland); Ryder, Rep. U.S. Comm. Fish. (1881), 1884: 795 (temp. effects on incub. of
eggs); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 427; 47 (4), 1900: fig. 191 (descr.,
range, synon.); Anonymous, Rep. U.S. Comm. Fish. (1897), 1898: 133, pls. 36-39 (artif. propag.
and develop.); Bean, T. H., 7th Rep. Forest Comm. N.Y. (1901), 1902: 305, fig. (synon., range, occur.,
size, food of young in rivers, spawn.); Bean, T.H., Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 204
(synon., refs., decline due to obstructions, food, migr., growth); Fowler, Rep. N. J. St. Mus. (1905),
1906: 99 (descr., import. as food); Smith, N. C. geol. econ. Surv., 2, 1907: 125, pl. 5 in color (synon.,
descr., range, foodfish; rel. size of male and female, migr., statist., etc.); Kendall, Occ. Pap. Boston Soc.
nat. Hist., 7, 1908: 39 (refs., New England); Sumner, Osburn, and Cole, Bull. U. S. Bur. Fish. (1911),
31 (2), 1913: 742 (refs., Woods Hole, Massachusetts, spawn., parasites); Regan, Ann. Mag. nat.
Hist., (8) 28, 1916: 8 (diagn.); Wilson, Proc. U.S. nat. Mus., 53, 1917: 59 (parasites); Leim, Contr.
Canad. Biol., N.S. 2 (1), 1924: 163-284 (life hist., bibliog.); Barney, Trans. Amer. Fish. Soc., 54,
1924: 168, figs. 1-4 (age from otoliths); Borodin, Trans. Amer. Fish. Soc., 54, 1924: 178 (age from
scales); Bigelow and Welsh, Bull. U.S. Bur. Fish. (1924), 40 (1), 1925: 113, fig. 46 (descr., size,
range, Gulf of Maine, migr., food, develop., etc.); Jordan, Fishes, 1925: 276 (relation., quality as food,
spawn.); Nichols and Breder, Zoologica, N.Y., 9 (1), 1927: 40, fig. (distr., New York and New Eng-
land, food, life hist., size); Hildebrand and Schroeder, Bull. U.S. Bur. Fish. (1927), 43 (1), 1928: 93,
fig. 53 (descr., growth of young, food, migr., artif. propag., distr., commerc. import., marketing, size, range);
Breder, Field Bk. Mar. Fish. Atl. Cst., 1929: 66, fig. (abund., decline, migr., food, spawn.); Jordan,
Manual Vert. Anim. NE U.S., ed. 13, 1929: 40 (diagn., range); Truitt, Bean, and Fowler, Bull. Md.
Conserv. Dep., 3, 1929: 50, fig. 16 (descr., Maryland); Jordan, Evermann, and Clark, Rep. U. S. Comm.
Fish. (1928), 2, 1930: 42 (range, synon.); Walford, Fish. Bull., Sacramento, 28, 1931: 48 (diagn.,
distr. Pacific coast, commerc. import.); Greeley and Bishop, 21st Rep. N. Y. Conserv. Dep. (1931), Suppl.,
6, 1932: 78 (rare, Oswegatchie and Black R. systems); Greeley and Bishop, 22nd Rep. N. Y. Conserv.
Dep. (1932), Suppl., 7, 1933: 91 (extinct, upper Hudson); Greeley, 24th Rep. N. Y. Conserv. Dep.
(1934), Suppl., 9, 1935: go (Hudson R.); Greeley, 25th Rep. N. Y. Conserv. Dep. (1935), Suppl.,
10, 1936: 78 (Delaware R. within New York); Greeley, 26th Rep. N. Y. Conserv. Dep. (1936), Suppl.,
II, 1937: 91, pl. 3 in color (abund. Hudson R., spawn.); Vladykov and Wallace, Trans. Amer.
Fish. Soc. (1937), 67 (2), 1938: 52-66; also Contr. 17, Chesapeake Biol. Lab. (commerc. import., Chesa-
peake Bay; var. in populations); Greeley, 28th Rep. N. Y. Conserv. Dep. (1938), Suppl., 15 (1), 1939:
393; 15 (2), 1939: 82 (fresh water, Long Island, New York); Greeley, 2gth Rep. N. Y. Conserv. Dep.
(1939), Suppl., 16, 1940: 68 (rare, Lake Ontario); Welander, Copeia, 1940: 221 (introd., distr. Pacific
coast); Hubbs and Lagler, Bull. Cranbrook Inst. Sci., 18, 1941: 27 (range); Fowler, Monogr. Acad.
nat. Sci. Philad., 7, 1945: 45 (synon., refs.); Anonymous, Fish. Resourc. U.S., Senate Doc. No. 51,
1945: ill, 67—68, fig. (migr., fish. management; decline, causes and remedies; range); Warfel and Olsen,
Copeia, 1947: 177-183 (var. in no. of vert. and signif.); Bigelow and Schroeder, Fish. Bull. (74)
U.S. Fish Wildl. Serv., 53, 1953: 108-112 (descr., life hist., Gulf of Maine).
Clark, Rep. U.S. Comm. Fish. (1881), 1884: 783 (amount of water needed to keep eggs and young
alive); Rice, Rep. U.S. Comm. Fish. (1881), 1884: 787 (retard. develop. of eggs); Willey, Contr. Canad.
Biol., N.S. z (16), 1923: 313-320 (food); Johnson, Invest. Rep. U.S. Bur. Fish., 2 (38), 1938: 1-42,
figs. 1-g (nat. hist., sources of fishery, capture, wholesale trade, consump. in homes and public eating
places, nutr. value, recipes, etc.); Hollis, Science, 108 (2804), 1948: 332 (tagging young; return of
mature fish to place of hatch.).
Doubtful References:
Clupea indigena Mitchill, Rep. in part on Fishes of New York, 1814: 22 (orig. descr.; type local. presumably
New York; type lost); Mitchill, Trans. lit. philos. Soc. N. Y., Z, 1815: 454 (descr. inadequate for ident.).
20°
308 Memoir Sears Foundation for Marine Research
Negative Reference:
Clupea sapidissima Jordan and Gilbert (not of Wilson), Proc. U.S. nat. Mus., 5, 1882: 247 (proportions and
counts, showing specimens were 4. a/abamae, not recognized until 1896; Pensacola, Florida).
Alosa alabamae Jordan and Evermann 1896
Alabama Shad, Gulf Shad, Ohio Shad
Figures 70, 75
Study Material. A total of 23 specimens, 175-505 mm TL, 145-395 mm SL,
from the Choctawhatchee River, Pensacola, Florida (including two cotypes of 4. a/a-
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Ficure 75. Alosa alabamae, 380 mm TL, 15 inches, Black Warrior River, Tuscaloosa, Alabama, type, USNM
47689. Drawn by A. H. Baldwin.
bamae); tributary of the Flint River, Georgia; Black Warrior River, Tuscaloosa, Ala-
bama (male and female types); Coosa River, and the Alabama River, tributary of the
Tombigbee River, Alabama; Washita River, near Hot Springs, Arkansas; Ohio River
at the falls, Louisville, Kentucky (including type and several cotypes of 4. ohiensis);
and Fairport, lowa.
Distinctive Characters. See Key to Species, p. 294, and pp. 304, 309, 310.
Description. Proportional dimensions in per cent of standard length, and counts,
based on the 23 study specimens, 145-395 mm SL.
Body: depth 25.5—36. Maxillary: length 11-13.5.
Caudal peduncle: depth 8.0—9.0. Pelvic fin: length 10-11.5.
Head: length 23.5-26.5. Pectoral fin: length 15.3-17.5.
Snout: length 5.5—6.9.
Eye: diameter 4.2—6.5. Scales: ca. §5—60, often lost in preserved
Interorbital: width 4.5—-6.0. specimens.
Fishes of the Western North Atlantic 309
Ventral scutes: 35-38. anal 19-22; pectoral 15-18, usu-
Gill rakers: 42-48 in adults. ally 16 or 17.
Fin rays: dorsal 16-20, usually 17-19; Vertebrae: $5 (1 specimen).
Bopy shaped as in J. sapidissima, the depth in general increasing with age and
growth, 2.8—3.9 in SL in adults. Heap 3.7-4.25 in SL. Snour 3.8—4.4 in head. Eye
4.2—6.5. INTERORBITAL 4.1-6.0. MaxiLLary 1.95—2.15. ScaLes only moderately ad-
herent, about 15 or 16 rows between base of pelvic and anterior rays of dorsal fin.
Petvic fin 2.2—2.5 in head. Pecrorat fin 1.4—1.5.
A. alabamae differs from A. sapidissima in only the few respects shown in the
following comparison:
A. sapidissima A. alabamae
Gill rakers close-set, those at angle of Gill rakers less crowded, those at angle
first arch longer than snout, 59-73 of first arch about as long as snout,
on lower limb in adults 300 mm and 42-48 on lower limb in adults 300
upward. mm and upward in length.
Surface of cheek and opercular bones Surface of cheek and opercular bones not
glossy, generally without conspicu- glossy, covered with conspicuous mu-
ous mucous pores and canals. cous pores and canals.
Axillary scale of pelvic fin generally only Axillary scale of pelvic fin generally about
a little more than half of length of fin. 75 °/ of length of fin.
Size. The Alabama Shad seems to run smaller in size than the American shad; the
largest female seen was 510 mm (20.4 in.) and the largest male 435 mm (17.4 in.)
long, both from the Ohio River at Louisville, Kentucky.
Development. Apparently the eggs and young have not been described. Ten small
specimens, 48-90 mm TL, 36-73 mm SL, all previously misidentified as Pomolobus
chrysochloris (USNM 36424, 36620, 36835, and 62225), are no doubt in their first
year; they were collected from: the Stone River at Nashville and the Clinch River at
Clinton, Tennessee; the Washita River, and the Mulberry River at Mulberry, Arkan-
sas. There is also at hand a specimen 105 mm TL (93 mm SL) taken in brackish water
at Grand Isle, Louisiana. The young differ from their adults principally in having a
more slender body and in possessing notably fewer gill rakers; the depth is 3.6—-4.1 in
SL and the gill rakers on the lower limb vary between 27-33. The increase in the
number of gill rakers seems to be rapid, as five half-grown examples 175-215 mm TL
(145-171 mm SL) from Pensacola already have 38-41 gill rakers.
‘Numerous young, 8 or g inches in length” were reported for Pensacola, Florida,
by Jordan and Gilbert (73: 247) as 4. sapidissima, before A. alabamae had been recog-
nized; five of these (USNM 30809) may be part of that material; they are typical
of the Alabama Shad and are the only half-grown samples at hand. They, like the smal-
ler American shad, are more slender than their adults and have fewer gill rakers; but the
raker count in these is higher than in the smallest examples of a/abamae reported above.
310 Memoir Sears Foundation for Marine Research
Spawning and Migrations. A. alabamae, like its close relative 4. sapidissima, ascends
streams in spring and spawns in fresh water. Sexually mature Alabama Shad, ob-
served at Keokuk Dam, Iowa, in 1915 and 1916 from the early part of May to the
middle of July, were in upstream migration to spawn but were stopped by the dam,
except for a few that might have passed through the lock (27: 169). Coker believed
there were enough there “‘to support a substantial fishery.”” Other runs in the spring
of the year have been reported for the Black Warrior River (Tuscaloosa) and at Mobile,
Alabama (32: 127); the Kanawha River at Montgomery, West Virginia; and the
Ohio River at the falls, Louisville, Kentucky (32: 275-276). An apparently reliable
source reported still other runs to Dr. Evermann—for the Ohio River about 30
miles below Louisville, Kentucky, and for the Mississippi River below Memphis,
Tennessee.
In reply to a letter of inquiry addressed to the State Department of Conser-
vation, Montgomery, Alabama, Mr. Ben. C. Morgan, Director of Conservation, replied
(June 22, 1946):
During the months of May and June, and sometimes earlier, we still have enormous quantities of fresh
water shad which run or attempt to run up our main streams. I say attempt to run because of the fact that the
Coosa, Tallapoosa, Tennessee, and Tombigbee Rivers all have locks and dams which prevent the shad from going
further than these dams. Only last year on the Tennessee they banked below the dams by the millions and could
be scooped with nets, which we do not allow, of course.
It seems highly probable that the young of this species descend the rivers to salt
water at the end of their first summer, and it may be assumed, therefore, that the habits ~
of the Alabama and American shads are very similar.
Food. It is well known that anadromous fishes, while migrating to their spawning
grounds, do not feed regularly, if at all. The Alabama or Ohio Shad is no exception
to this rule, for Mr. Stringham found that 105 of 163 stomachs examined at Keokuk,
Iowa, were empty (27: 171). Others contained small quantities of insect and plant
remains as well as some unrecognizable debris.
Variations. Although adults of a/abamae and sapidissima differ widely in the number
of gill rakers, the number in the very young is so similar in this respect that overlapping
definitely occurs (see Description).
Examples taken from the Ohio River at the falls, Louisville, Kentucky, differed
sufficiently from a/abamae to be recognized by Evermann as a distinct species, which
he named Alosa ohiensis (31: 277). He described ohiensis as differing from a/abamae
principally in a more elongated body and in the greater number of gill rakers on the
first arch. However, the seven large specimens from the Ohio now at hand (including
the type), though representing the extremes in the two characters mentioned, intergrade
with the other 11 sexually mature examples (including the type material of alabamae
from Tuscaloosa) from the several other localities in the Gulf drainage already listed.
Therefore, ohiensis at most is only subspecifically distinct. It is herein regarded as a
synonym of alabamae.
Commercial Importance. This species has never been an important foodfish; only
Fishes of the Western North Atlantic ana
small catches have been recorded a couple of times in the statistical reports of the former
U.S. Fish Commission: for 1889, 6,955 pounds from the Ohio River in Indiana and
Kentucky (1902: 667); and for 1902, 150 pounds from Alabama (1905: 416). No
later catches seem to have been reported. By correspondence with Herman O. Hessen,
U. S. Fish and Wildlife Service, Louisville, Kentucky, it was learned that no one now
(1946) fishes for the ““Ohio Shad” at that place, though a few are taken incidentally
each spring. That these Shad were, and perhaps still are, numerous enough in some
places to be taken in considerable quantities is evident, but no special effort is apparently
made to catch them. In fact, the fish seem to be regarded in some localities as too bony
for human use. At least one fisherman interviewed by Mr. Hessen at Louisville, Ken-
tucky, stated that the fish were so bony that fishermen threw them on the bank and made
no effort to sell them. Nevertheless, the food qualities of this Shad, tested by those
who are familiar with the delicious American shad, have been found not at all inferior
GF: 275; 27: 169).
It seems probable that Alabama Shad constitute at least a small unutilized food
supply, but their actual abundance remains unknown. Evermann indicated that if proper
equipment were used, considerable catches might be taken (317: 279); for example, a
great increase in the catch resulted at Louisville, Kentucky, when surface-fishing seines
were substituted for bottom-fishing seines. However, until people living on the shad
streams of the Gulf drainage recognize its delicious food qualities, there will be little
demand; hence no special effort to catch them will be made. Because of the many dams
constructed across rivers, barring them from the spawning grounds, a decrease in
abundance is no doubt taking place.
Despite an unutilized supply of Alabama Shad in streams tributary to the Gulf
of Mexico, large shipments of American shad fry, 4. sapidissima, were liberated in those
streams (2). However, no American shad are included among the rather numerous
specimens examined, nor has any authentic record of their occurrence in those streams
been found. Indeed, Alabama Shad seem to be referred to as “Potomac shad” by fisher-
men who believe they are the progeny of American shad planted by the former U.S.
Fish Commission.?® But it seems quite certain that the introduction failed.
Range. The Alabama Shad is represented by specimens in the National Museum
and has been recorded for all the principal streams tributary to the Gulf of Mexico
(exclusive of the Pearl River), from the Mississippi eastward to the Choctawhatchee
River, about 50 miles east of Pensacola, Florida. A 1os-mm specimen from Grand
Isle, Louisiana, is also at hand. In the Mississippi Valley it is known from as far west
as Hot Springs, Arkansas, and northward to Fairport, lowa, and Montgomery, West
Virginia.
Synonyms and References:
Clupea sapidissima Jordan and Gilbert (not of Wilson), Proc. U.S. nat. Mus., 5, 1882: 247 (meas., counts
29. Fishermen generally do not seem to distinguish Alabama Shad from skipjack herring, Pomolobus chrysochloris; ac-
cording to Coker (27: 165) they are referred to as “Government shad” by the fishermen at Keokuk, Iowa, apparently
in the belief that they are the result of these plantings.
ee") Memoir Sears Foundation for Marine Research
of half-grown examples from Pensacola, Florida); Bollman, Proc. U.S. nat. Mus., 9, 1886: 462 (two large
ones, Escambia R., Alabama).
Clupea chrysochloris Meek (not of Rafinesque), Bull. U.S. Fish Comm., 14, 1894: 85 (Mulberry, Arkansas;
USNM 62225 4. a/abamae).
Alosa alabamae Jordan and Evermann in Evermann, Rep. U.S. Comm. Fish. (1895), 21, 1896: 203 (orig.
descr.; type local. Black Warrior R., Tuscaloosa, Alabama; types USNM 47689 [9] and 47690 [3];
cf. A. sapidissima); Evermann and Kendall, Bull. U. S. Fish Comm. (1897), 27, 1898: 127 (distr.); Jor-
dan and Evermann, Bull. U.S. nat. Mus., 47 (3), 1898: 2810, figs. 192, 192a (descr., refs.); Regan,
Ann. Mag. nat. Hist., (8) 78, 1916: 9 (refs., diagn.); Jordan, Evermann, and Clark, Rep. U.S. Comm.
Fish. (1928), 2, 1930: 43 (range, ref.).
Alosa ohiensis Evermann, Rep. U.S. Comm. Fish. (1901), 27, 1902: figs. 1 and 2, 9 and ¢ ohiensis; figs. 3 and
4, g and 2 alabamae (orig. descr.; type local. Ohio R., Louisville, Kentucky; types USNM 50469;
cf. alabamae and sapidissima; distr.); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928),
2, 1930: 42 (range, ref.); Coker, e¢ a/., Bull. U.S. Bur. Fish (1929), 45, 1930: 169 (Keokuk, Iowa
1915, 1916; influence of dam on migr.; an unused food).
Alosa alabamae ohiensis Regan, Ann. Mag. nat. Hist., (8) 18, 1916: 9 (ref., diagn.).
Genus Pomolobus Rafinesque 1820
Alewives, River Herring
Pomolobus Rafinesque, West. Rev. Misc. Mag., 2, 1820: 170; Ichthyol. Ohiensis, 1820: 38; genotype by mono-
typy, Pomolobus chrysochloris Rafinesque.
Generic Synonym:
Alausella Gill, Proc. Acad. nat. Sci. Philad., Suppl., 1861: 54; Proc. Acad. nat. Sci. Philad., 1861: 36; genotype
by monotypy, Clupea parvula Mitchill equals probably young of C/upea pseudoharengus Wilson.
Characters. Bopy oblong, compressed. Bony scutes strong, with sharp points; a
sharp edge on median line of chest and abdomen. Cueek generally longer than deep,
about as long as deep in P. mediocris. Moutu large. Maxttiary extending to or
beyond middle of eye. Upper yaw with a more or less definite median notch in adults.
Lower Jaw projecting or included, its upper margin within the mouth rising more or
less abruptly. Teer in jaws not entirely wanting in adults; present at least anteriorly
in lower jaw, though rudimentary in some species; present on median line of tongue;
wanting on vomer. Dorsat with 15-21 rays, the last one not produced, its origin
nearer to tip of snout than to base of caudal. ANAL with 16-23 rays, the last one not
especially enlarged. Petvic with 9 rays. VERTEBRAE 46-55.
Remarks. Two members of this genus, mediocris and chrysochloris, are close to the
American species of A/osa in the general structures about the head, except that the lower
jaw projects beyond the upper one; they differ strongly, however, in having fewer gill
rakers, which Do NoT INCREASE IN NUMBER WITH AGE (18—24 on lower limb). The other
two species, aestivalis and pseudoharengus, herein recognized, agree more or less with the
species of 4/osa in the number of gill rakers (38-51 on lower limb) and in having the
lower jaw included; they differ, however, in having much shallower cheeks, which are
much longer than deep, and in having the upper margin of the lower jaw rising abruptly
within the mouth to form a more or less obtuse angle.
Ficure 76. Pomolobus heads, showing shape of mandible and cheek. a, chrysochloris, 54 mm'TL, USNM 9424;
B, mediocris, 122mm TL (after Hildebrand and Schroeder); c, gestivalis, 45 mm TL, USNM 69834;
pv, pseudoharengus, 48 mm TL, USNM 69833. a drawn by Ann S. Green, B by Louise Nash, c, p by Nancy
D. Patton.
Range. Three species occur in the Atlantic, from Nova Scotia to northern Florida,
and one occurs in the Gulf of Mexico and tributary streams as far west as Corpus
Christi, Texas, ranging northward to Minnesota (see Key).
Key to Species
1a. Lower jaw strongly projecting, extending nearly or quite into dorsal outline of
head (Fig. 76 a, B); only 18—24 gill rakers on lower limb of first arch, not increas-
ing in number with age.
2a. Greatest depth of body usually less than length of head, 3.5—4.1 (usually
3.8-4.0) in SL; teeth present in jaws at all ages, those in lower jaw prominent;
bluish green color of back ceasing abruptly, not blending into the silvery
color of side; no spot at shoulder. chrysochloris Rafinesque 1820, p. 315.
314 Memoir Sears Foundation for Marine Research
2b. Greatest depth of body usually greater than length of head, 3.0-3.75 (usually
3-2—3.6) in SL; teeth missing in upper jaw of adults, those in lower jaw very
small (rudimentary); the grayish green color of back blending rather gradually
into silvery color of the side; a dark spot at shoulder just behind head, often
followed by several smaller spots. mediocris (Mitchill) 1814, p. 319.
1b. Lower jaw included or projecting but little, definitely not extending into dorsal
outline of head (Fig. 76 c, p); gill rakers 38-41 on lower limb in adults, numer-
ous at all ages.
3a. Diameter of eye generally about equal to length of snout in adults; 41-52
(most frequently 44-50) gill rakers on lower limb in adults; peritoneum
usually black, sometimes pale with dusky dots; back in fresh examples bluish.
aestivalis (Mitchill) 1814, p. 324.
3b. Diameter of eye longer than snout at all ages; 38-44 (most frequently 39-41)
gill rakers on lower limb in adults; peritoneum pale, with dusky dots; back
in fresh examples grayish green. pseudoharengus (Wilson) 1811, p. 332.
‘ (Lh, s—
Ficure 77. Pomolob m TL, USNM 125701;
LEFT, mediocris, 310mm TL, USNM 118156; ricut, pseudoharengus, 292 mm TL, USNM 117808. All
drawn by Nancy D. Patton.
Fishes of the Western North Atlantic ues
Pomolobus chrysochloris Rafinesque®® 1820
Skipjack Herring, Skipjack, Golden Shad, Blue or Green Herring, River Herring
Figures 76, 78
Study Material. A total of 32 or more specimens, 115-420 mm TL, 95-341 mm
SL; from Pensacola, Florida; Wetumpka, Montgomery, and Wheeler Reservoir,
Alabama; Pascagula, Chickasawhay River, Pearl River, Eagle Lake, and Vicksburg,
Mississippi; shrimp grounds (trawl) in Louisiana waters, Pass Manchac, and borrow
.
Ficure 78. Pomolobus chrysochloris, 320mm TL, Pensacola, Florida, USNM 30159. Drawn by S. Stearns.
pits in Mississippi River at Lake Providence, Louisiana; Mustang Island (Gulf side),
Texas; St. Francis River, Mississippi River, Miles County, Arkansas; Lexington,
Missouri; Mississippi River, Keokuk, and Fairport, lowa; Cairo and Mississippi River,
Victory, Wisconsin; and Fort Snelling, Minnesota; mostly in USNM, but some
loaned by Miss Fannye A. Cook, Mississippi Game and Fish Commission, Percy
Viosca, Southern Biological Supply Co., and Reeve M. Bailey, University of Michigan.
Distinctive Characters. P.chrysochloris is closely related to P. mediocris, the hickory
shad, of the Atlantic coast; the principal differences are indicated in the account
30. Although this species has long been considered the genotype of Pomolobus, it may be questioned with good reason
whether Rafinesque’s descriptions of the genus and species are based on the Skipjack Herring or the Alabama shad,
Alosa alabamae, or on both. If Rafinesque had both species, as suggested by his accounts, he confused them. In his
generic account he said “Jaws without teeth.”’ This is true of adult Alabama shad but not of the Skipjack. In fact,
the latter has such prominent teeth in the lower jaw that they would scarcely be overlooked. Furthermore, in the
specific account Rafinesque used the local name, Ohio shad, together with Gold shad and Green herring. Present-
day fishermen certainly distinguish the two species and refer to 4. alabamae as the shad and to P. chrysochloris as
the herring (generally with a modifier, such as blue, green, or river).
When listing local names, Rafinesque remarked: “‘A fine fish from twelve to eighteen inches long. Flesh esteemed,
white and less bony than the shad.” The comparison evidently is with the American shad. The remark concerning
the flesh seems to suit the Alabama shad, but not the Skipjack. On the other hand, the size given is more nearly that
of the Skipjack than that of the shad; yet an 18-inch Skipjack would be an exceptionally large one. It is evident,
then, that one cannot be certain as to the species on which Rafinesque’s descriptions were based. Because of the
doubt involved, it seems advisable to continue the use of Pomolobus chrysochloris for the Skipjack Herring. To do
otherwise would lead to great confusion of both generic and specific names.
316 Memotr Sears Foundation for Marine Research
of mediocris (p. 322). Both of these species are at once distinguishable from the other
species of Pomolobus and from Alosa by the much smaller number of gill rakers.
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens, 95-341 mm SL.
Body: depth 24.2—28.4. Scales: 51-60, often difficult to count
Caudal peduncle: depth 7.8-11. accurately.
Head: length 24.6-27.8. Ventral scutes: 33-37, most frequently
Snout: length 4.8-6.3. 24) OF. 3152
Eye: diameter 4.4—7.0. Gill rakers: on lower limb of first arch
Interorbital: width 3.8-4.8. 20-24, most frequently 20-23.
Maxillary: length 10.5—12.5. Fin rays: dorsal 16-21, most frequently
Pelvic fin: length 10-12.6. 18—20; anal 18-21, most frequently
Pectoral fin: length 16-19. 19 or 20; pectoral 16 or 17, rarely 15.
Vertebrae: 53-55 (4 specimens).
Bopy slender, compressed, its greatest thickness generally somewhat less than
half of its depth, its depth increasing with age and growth, usually equal to or less than
length of head, 3.5-4.1 in SL.
Scales moderately adherent, with slightly irregular membranous margins, not
preceded by striae; 15 or 16 longitudinal rows of scales on body between base of pelvic
fin and anterior rays of dorsal. Venrra scutes well developed, 17—21 (most frequently
19 or 20) in advance of pelvics and 14-17 behind them.
Heap 3.6-4.0 in SL. Snour 4.2-4.8 in head. Eye 3.8-—5.9. INTERORBITAL 5.4—
7.5. CHEEK, from lower rim of orbit, somewhat longer than deep. Mourn moderately
oblique. Upper jaw with a definite median notch in adults. Lowsr jaw projecting
strongly and extending into dorsal profile of head, its upper margin within mouth
rising gradually, not forming an angle (Fig. 76 a). Maxittary broad, its width
about 0.67 of length of snout, rounded posteriorly, reaching nearly or quite below
posterior margin of pupil, 2.25-2.4 in head. GiLL RAKERs not increasing in number
with age. Trrru in anterior part of both jaws at all ages, those of lower jaw rather
prominent; sockets present if teeth are broken; teeth on tongue prominent, in a
rather large elongated patch.
Dorsat fin with slightly concave margin, its origin scarcely an eye’s diameter in
advance of pelvic and usually equidistant from margin of upper jaw and a vertical from
last ray of anal; a scaly sheath on dorsal base, composed of a single row of scales and
extending to 4th or sth ray from end of fin. Caupat deeply forked, the lower lobe
rather longer than the upper, about as long as head. Anat low, its margin nearly straight,
its origin usually about equidistant between base of pelvic and base of caudal; a scaly
sheath present on base. Pztvic fin inserted near ventral edge and usually nearer to
base of pectoral than to origin of anal, 2.15—2.4 in head. Pecrorat fin placed some-
what higher on side, 1.4—1.6 in head.
Color. Fresh specimens deep bluish green above, this color ceasing abruptly at
Fishes of the Western North Atlantic QEF
level of upper margin of gill cover. Sides light green, with strong tinge of gold, espe-
cially on head, shading into the white of belly. No dark shoulder spot. Tips of jaws
blackish. Dorsal fin yellowish, more or less dusky at base; caudal yellowish, tips of lobes
dusky; anal and pelvic pale; pectoral pale, a dusky streak on inner side behind first
ray. Peritoneum pale.
Size. Skipjacks are smaller in size than Alabama shad, with which they are more
or less associated in the rivers; their length usually ranges between about 300-450 mm
(12-18 in.).
Development. Although ovarian eggs 0.08—1.11 mm in diameter were observed,
apparently no young under 107 mm (about 4.33 in.) were secured during Dr. Coker’s
extensive investigations at Keokuk, Iowa (26; 27). Each of the smallest specimens at
hand (UMMZ 122510, 128570), from the mouth of Elk River, Wheeler Reservoir,
Alabama, and from the Mississippi River, Miles County, Arkansas, is about 40 mm
(31 mm SL), apparently the smallest known. They differ from larger examples princi-
pally in being more slender, the depth of each being contained 4.75 times in Sie
Furthermore, they have minute teeth on the margin of the maxillary, these persisting
in examples up to 150 mm or so in length. The gill rakers, unlike those in some other
species of the genus Pomo/obus, do not increase in number with age and growth.
Spawning and Migrations. The spawning place and length of spawning are not
fully known. For such information as is available we are indebted to Robert E. Coker,
who, with assistants, carried on an extensive investigation at Keokuk, Iowa (26: 25, 26;
27: 94, 165-169). They reported fish with large roe taken there on: April 29 and
May 23 in 1914; April 29 and 30, May 24, June 5 and 23, July 1 and 2, and
“various other dates” in 1915; and on May 23, June 3 and 17 in 1916. Fully mature
fish, ripe for stripping, apparently were not included. Coker concluded his remarks on
spawning as follows:
Many attempts made during several years to find a particular place and time where and when river herring
were clearly spawning met with no success, although various methods were tried. The problem proved unexpec-
tedly baffling. Our data suggest only that the spawning season is prolonged, possibly beginning early in May;
that the fish do not spawn in large aggregations; that during spawning operations they are not readily captured
by ordinary methods of fishery; and that spawning is ended soon after the first of July.
It has not been proven that Skipjack are anadromous like most of the near rela-
tives, even though Jordan said categorically, “. .. migrating from the Gulf of Mexico”
(71: 40). However, they do enter salt and brackish water. Such a great “run’’ as was
reported at Keokuk, Iowa, in the spring of 1914 as well as smaller runs in succes-
sive years and their apparent disappearance in September of each year do suggest
extensive migration. However, Coker expressed the opinion that the migrations do not
necessarily extend to extreme northern or extreme southern waters. This is in contrast
to the strictly anadromous relatives, Alosa sapidissima, Pomolobus aestivalis, and P. pseudo-
harengus (except where the last species is landlocked), whose young leave the fresh-
water streams at the end of their first summer and do not return until they are ready
to spawn.
318 Memoir Sears Foundation for Marine Research
Evidence has been set forth that chrysochloris reproduced in Lake Pepin after
the closing of Keokuk Dam, an almost complete barrier to them. Dr. Coker said that
without question there was a decided decline in the number of “‘River Herring”’ in this
Lake during the three years immediately following the dam’s construction, but some
commercial catches there in 1926, 12 years after the dam was closed, convinced him
that these fish were not rare; and several commercial fishermen stated that “‘the
herring were coming back.” On the basis of Dr. Coker’s observations, it seems
proper to conclude that P. chrysochloris, although migratory, does not need to have
access to salt water to reproduce, and the species surely is not strictly anadromous.
The presence of immature examples in a collection from brackish water as well as from
the upper Mississippi also may be of some significance.
Food. Skipjack Herring are carnivorous (35: 49); the young feed on insects, the
adults on fish. Coker stated that, of about 150 examples examined for food:
Approximately one-third were empty; a little more than one-third contained fish, chiefly minnows, with some
mooneyes, gizzard shad, and other fish not determinable; and less than one-third contained insects and larvae,
principally May flies, with some caddis flies and others.
Habits. These are more active fish than most other clupeoids; they frequent rivers
with swift currents and often leap from the water, a habit from which they have derived
the name Skipjack. They leap in part for sport and in part for the pursuit of prey (35:
49). Coker remarked that they have, ‘‘a striking habit of leaping from the surface
when feeding upon schools of minnows or in mere play.”
Parasites. We know only that the larvae (glochidia) of the freshwater mussel Fus-
conaia ebena are parasitic on the Skipjack (see below).
Commercial Importance. The value of this species as a foodfish is negligible, for
these fish are bony and not especially well flavored. No catches have been listed in the
statistical reports of the U.S. government. However, they represent a very distinct
economic asset as a host for the larvae of Fusconaia ebena, which has been regarded
as the most valuable of all the pearly mussels of the Mississippi Basin because of its
abundance in all the larger waters of strong current and because it yields a shell of the
best quality for buttons (27: 166).
Range. Skipjack Herring range in the Gulf of Mexico from Pensacola, Florida,
to Corpus Christi, Texas, and sometimes out into the Gulf, as off Breton Island,
Louisiana. They also inhabit larger streams, occasionally lakes, or even borrow pits, of
the Gulf drainage. They commonly range as far north in the Mississippi River as Lake
Pepin, and occasionally to Fort Snelling, Minnesota, to Hudson, Wisconsin, and in
the Ohio River to Pittsburgh, Pennsylvania. Apparently they are landlocked above
Keokuk Dam, Iowa. Although Smith (rr8: 214) said that they were abundant in Lake
Erie, and Jordan and Evermann (72, 1896: 425) as well as several others have stated
that they reached Lake Michigan and Lake Erie through canals, Hubbs and Lagler
(62: 27, ftn.) do not admit them to their list of fishes of the Great Lakes and tributary
waters. The species apparently is most common in swift currents of large streams.
Fishes of the Western North Atlantic 319
Synonyms and References:
Pomolobus chrysochloris Rafinesque, West. Rev. Misc. Mag., 2, 1820: 171; and Ichthyol. Ohiensis, 1820: 39
(orig. descr.; type local. Ohio R.; type lost; length 12-18 in. attained; seldom ascend to Pittsburgh) ;31
Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 425; 47 (4), 1900: fig. 187 (descr., size,
distr., habits, value as food); Jordan and Evermann, Amer. Food Game Fishes, 1g02: ror (descr., range;
Lakes Erie and Michigan erroneously included); Bean, T.H., 7th Rep. Forest Comm. N. Y. (rgor),
1902: 300, fig. (synon., range, habitat; reported to have reached Great Lakes through canals, but occur.
there seems doubtful); Bean, T. H., Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 195 (synon., descr., value
as food, habits, distr.); Forbes and Richardson, Nat. Hist. Surv. Ill., Ichthyol., 3, 1908: 48 (refs., descr.,
range, occur. in Illinois, size, habits); Coker, Rep. U. S. Comm. Fish. (1913), Append. 8, 1914: 25-28
(large schools below Keokuk Dam, Iowa; discus. migr. and ascent up spillways of dam; occur. above
dam; spawn.); Regan, Ann. Mag. nat. Hist., (8) 79 (112), 1917: 299 (refs., descr., range); Jordan,
Manual Vert. Anim. NE U.S., 1929: 40 (descr., range, habitat); Coker, Bull. U.S. Bur. Fish. (1929),
45, 1930: 165, fig. 11 (below and above Keokuk Dam, Iowa, migr., spawn., econ. import.); Hubbs
and Lagler, Bull. Cranbrook Inst. Sci., 18, 1941: 27 (not in list of Great Lakes fishes though reported
there earlier); Gunter, Publ. Inst. mar. Sci. Texas, F (1), 1945: 25 (Texas coast, Colorado R. at Austin).
Meletta suoerii Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 375 (orig. descr.; type local. Wabash
R., tributary of Mississippi R.); Bertin, Bull. Soc. zool. Fr., 66, 1941: 23 (ident. of type).
Clupea chrysochloris Jordan and Gilbert, Proc. U.S. nat. Mus., 5, 1882: 247 (descr.; records two specimens;
from Galveston, Texas; and Pensacola, Florida); Jordan and Gilbert, Bull. U.S. nat. Mus., 76, 1882:
266 (descr., range; Lakes Erie and Michigan apparently includ. erroneously); Jordan and Gilbert,
Proc. U. S. nat. Mus., g, 1886: 11, 14 (Washita R., Arkadelphia, and Saline R., Benton, Arkansas);
Bollman, Proc. U.S. nat. Mus., 9, 1886: 462 (few from Escambia R., Alabama); Gilbert, Bull. U.S.
Fish Comm. (1889), 9, 1891: 158 (near mouth of Escambia R., Alabama); Smith, Bull. U. S. Fish
Comm. (1890), 10, 1892: 214, pl. 31 (Lakes Ontario and Erie); Evermann and Kendall, Bull. U.S.
Fish Comm., 12, 1894: 105 (Red R., Fulton, Arkansas; Galveston, Texas); Meek, Bull. U.S. Fish
Comm., 14, 1894: 78, 82 (White R. at Batesville, Little Red R. at Judsonia, Arkansas).
River herring, Coker, Bull. U.S. Bur. Fish., 45, 1929: 91-104 (discus. abund. below Keokuk Dam, Iowa,
1914-1916; dam a barrier to fish).
Negative References:
Clupea chrysochloris Meek (not of Rafinesque), Bull. U.S. Fish Comm., 14, 1894: 85 (Mulberry, Arkansas,
but specimen USNM 62225 is 4. a/abamae).
Pomolobus mediocris (Mitchill) 1814
Hickory Shad, Hickory Jack, Bonejack, Fall Herring,
Freshwater Taylor, Shad Herring
Figures 76, 77, 79
Study Material. Forty-three specimens, 105-460 mm TL, 82-354 mm SL, from
Provincetown, Massachusetts; Noank, Connecticut; Long Island, New York; Beesley’s
Point, New Jersey; Annapolis, Love Point, Benedict, Barren Island, Lewistown, and
Smith Point, all on or near Chesapeake Bay, Maryland; Potomac River, Washington,
D. C.; Buckroe Beach and Lynnhaven Roads, on Chesapeake Bay, Virginia; Beaufort,
North Carolina; and Brunswick, Georgia.
31. The data are from Richard E. Call’s Ichthyologia Ohiensis (1899), ““A verbatim et literatim reprint of the original,
with a Sketch of the Life, the Ichthyologic Work, and the Ichthyologic Bibliography of Rafinesque,” the original
publications not being available to the author.
320 Memoir Sears Foundation for Marine Research
Distinctive Characters. See Relationships for differences between mediocris and chry-
sochloris (p. 322). From species of A/osa and other Pomolobus species, excepting chryso-
chloris, it is marked by the noticeably smaller number of gill rakers.
Description. Proportional dimensions in per cent of standard length, and counts,
based on 43 specimens, 82-354 mm SL.
Bp
anes tas
ARR » :
yy yy) WN yy eins
Ficure 79. Pomolobus mediocris, 309 mm TL, Brunswick, Georgia, USNM 118156. Drawn by Nancy D.
Patton.
Body: depth 26.7—32.8. Scales: 48-57.
Caudal peduncle: depth 9.7-12. Ventral scutes: 33-38, most frequently
Head: length 24.5-29.6. 6 O&N37: :
Snout: length 5.9—7.5. Gill rakers: on lower limb 18-23, usually
Eye: diameter 4.9—7.1. 20 or 21.
Interorbital: width 3.9-5.4. Fin rays: dorsal 15-20, most frequently
Maxillary: length 10.4-13.4. 17 or 18; anal 19-23, most fre-
Pelvic fin: length 10.6-12.4. quently 20 or 21; pectoral 15 or 16.
Pectoral fin: length 17-21. Vertebrae: 54 or $5 (2 specimens).
Bopy moderately slender, well compressed, its greatest thickness notably less than
half of its depth, its depth rather greater in large than in small specimens, usually
exceeding length of head, 3.0-3.75 in SL.
ScaLes only moderately adherent, with definitely crenulate membranous margin,
preceded by longitudinal striae; about 16 longitudinal rows of scales on body between
base of pelvic and anterior dorsal rays. VENTRAL scuTEs moderately developed, 19-23
(usually 20-22) in front of pelvic fins, 12-17 (usually 13-16) behind them.
Heap 3.4—4.1 in SL. Snour 3.7—5.0 in head. Eye 3.9-5.3. InrERORBITAL 5.5—
7.0. CuEEk generally about as deep as long (Fig. 768). Mourn oblique. Upper jaw
with a definite median notch. Lower jaw projecting strongly and extending into dorsal
profile of head (Fig. 768), its upper margin within the mouth rising gradually, not
forming an angle (Fig. 768). Maxitiary broad, its width somewhat more than half
of length of snout, rounded posteriorly, reaching about to vertical from posterior
Fishes of the Western North Atlantic Bar
margin of pupil, 2.0—2.4 in head. Grit RAkERs apparently not increasing in number
with age. Trrru in the upper jaw absent in large examples 230 mm SL and upward,
but present in smaller ones; present in lower jaw, becoming exceedingly small in large
examples; those on tongue minute, in a small elongated patch.
Dorsat with slightly concave margin, its origin rather less than an eye’s diameter
in advance of pelvic, usually nearer to margin of upper jaw than to vertical from last
anal ray; a sheath of scales on its base, consisting for the most part of more than one
row of scales and extending nearly or quite to base of last ray. Caupat deeply forked,
the lower lobe somewhat longer than the upper, fully as long as head. Anat low, its
margin nearly straight, its origin about equidistant between base of pelvic and base
of caudal; a sheath of scales on its base. Pevic fin inserted near ventral edge and
about equidistant between base of pectoral and origin of anal, 2.0-2.6 in head. Pxc-
TORAL placed somewhat higher on side, 1.3—1.7 in head.
Color. Fresh specimen from Chesapeake Bay grayish green above, shading some-
what gradually into the iridescent silver of sides. Nape green, side of head brassy.
Tip of lower jaw dusky. Dorsal and caudal dusky. Anal and pelvic plain, translucent.
Pectoral slightly dusky. Narrow dark lines along rows of scales on upper part of side;
missing on specimens less than 150 mm SL or so and most distinct in large examples
that have lost their scales, the dark pigment being in the skin underneath the scales.
A dark spot on shoulder; several obscure dark spots behind it, but missing in some
examples. Peritoneum pale, but with scattered dusky punctulations.
Size. Hickory Shad definitely run larger in size than blueback and grayback
herrings but smaller than the shad. A maximum length of 600 mm (24 in.) has been
reported, but the usual size is much smaller. The commercial catches observed by me
at Chesapeake Bay and at Beaufort, North Carolina, consisted mostly of fish varying
between about 300-375 mm (12-16 in.) in length.
Reproduction, Development, and Migrations. Next to nothing is known about the
breeding habits of Hickory Shad. Although it has been stated that they spawn in
North Carolina in the spring, in the headwaters of streams (zrg: 121), spawners have
never been seen in either fresh or salt water. In fact, many subsequent authors (45:
607; 16: 106; 20: 65) have accepted without reservation the opinion that Hickory
Shad, like other local members of that genus, are anadromous. However, the only
supporting evidence is that there are definite spring and autumn runs; in Chesapeake
Bay, only stragglers are seen in the summer; in the spring they are first taken in
the lower part of the Bay in March and progressively later farther up the Bay in late
March and early April. Also, I have observed adults with well-developed roe in the
Potomac above Washington during the spring, but these were not ripe and may have
returned to salt water to spawn. The view that this fish does not enter salt water to
spawn (8: 197; 37: 95) is supported by Hildebrand and Schroeder (59: 84); among
thousands of young clupeids collected during many years in Chesapeake Bay and its
tributaries, in freshwater streams and estuaries, and in salt water out to 15 fms., they
failed to find any Hickory Shad less than 15 mm (6.12 in.) long. Also, in the vicinity
21
a22 Memoir Sears Foundation for Marine Research
of Beaufort, North Carolina, they were absent from extensive collections made by
Hildebrand and his associates with gear of various types, though adults were taken
in the estuaries there in winter and early spring.
In short, Hickory Shad probably spawn in the spring, but it remains to be
learned whether they spawn in fresh or salt water. However this may be, to the north
of New York they are the most plentiful during autumn.
Food. Hickory Shad feed chiefly on small fish of various kinds. Their fish diet
is supplemented by squid, small crabs, various small crustaceans, and by fish eggs
(59: 84).
Enemies and Parasites. Little is known about the enemies of this species; but no
doubt they are eaten by larger fish and are caught in limited quantities by man. Linton
(85: 438) found individuals parasitized with nematodes (Ascaris sp.), cestodes (larval
Scolex polymorphus), and trematodes (Distomum appendiculatum). Wilson (136: 12) found
the Hickory Shad to be a host of the copepod Lernanthropus brevoortiae.
Relationships. Though closely related to P. chrysochloris, Hickory Shad have a
heavier and less shapely (less streamlined) body; the depth in adults is greater than
the length of the head (measured from margin of upper jaw); in chrysochloris, exclusive
of very large or very fat specimens, the depth is equal to or less than the head. In
mediocris the shield of scales on the dorsal fin base consists of more than one row of
scales anteriorly and extends nearly or quite to the last ray of the fin, whereas in chryso-
chloris it is composed of only one row of scales throughout and is missing on the last
four or five rays. The scales on the side of the Hickory Shad have a pronounced crenulate
membranous border preceded by longitudinal striations, while in chrysoch/oris the
border is only slightly irregular and the striations not evident. In mediocris there is
only a slight ridge above the base of the pectoral, which ends at the tip of the axillary
process; in chrysochloris a more pronounced depression (pocket) is provided for the
pectoral fin, and the more prominent ridge above its base extends well beyond the
axillary process of the pectoral.
In large Hickory Shad, 300 mm and upward, teeth in the upper jaw are missing,
but in specimens at least up to 150 mm long they are present there; however, those
in the lower jaw persist, though very small; the patch on the tongue is very small, and
the teeth minute. In chrysoch/oris the teeth are consistently larger, persist in the upper
jaw among adults, are large in the lower jaw, and the patch on the tongue is notably
larger. Although the proportions and counts overlap, it is evident that Hickory Shad
have a proportionately deeper body, deeper caudal peduncle, more anal rays, and
fewer gill rakers than P. chrysochloris.
There are differences in color also, evident in preserved as well as fresh examples.
Hickory Shad have a large dark spot behind the upper part of the opercle, often
followed by several smaller dark spots, and the adults frequently have dark lines along
the rows of scales on the upper part of the side. In P. chrysoch/oris these markings are
missing, and the dark blue or grayish color of the back ends very abruptly, not tending
to shade into the silvery color of the side, as in Hickory Shad.
Fishes of the Western North Atlantic 2.23
Commercial Importance. Hickory Shad are of minor importance as foodfish, mainly
because the meat is bony and is considered inferior in flavor. Their value is somewhat
higher in North Carolina than elsewhere, because they are taken there during the
winter when other fish are scarce. In 1940 or 1941, the following catches were reported:
New England (all reports for Rhode Island and Connecticut), 25,800 pounds; the
Middle Atlantic States (all reports for New Jersey), 4,400 pounds; the Chesapeake
Bay states, 196,200 pounds, of which 22,000 pounds were taken in Maryland and
174,200 pounds in Virginia; and the South Atlantic states (all of which contributed),
371,500 pounds, of which 335,900 pounds were listed for North Carolina. It is
evident, then, that the fishery is centered in Virginia and North Carolina. In the
United States (1940, 1945, 1950-1953), catches have ranged between o and 25,800
pounds for southern New England, 200 and 4,400 pounds for the Middle Atlantic
states, 86,000 and 264,000 pounds for Chesapeake Bay, and 273,000 and 970,000
pounds for the South Atlantic states.
Method of Fishing. The commercial fish are caught principally with seines and pound
nets, the smaller quantities with gill and fyke nets. The fishery is not intense enough
to affect greatly the abundance of these fish in nature. They take the hook occasionally
and offer some sport in the spring, at least at Little Falls on the Potomac River. Anglers
trolling for striped bass (Roccus saxatilis) and mackerel (Scomber) landed a number of
Hickory Shad off the Merrimack River, Massachusetts, in 1932.
Range. The range of Hickory Shad extends from Florida to Maine. They are re-
corded for Campobello Island, New Brunswick (52: 47), but no record of their oc-
currence south of the St. Johns River in Florida has been brought to my attention.
Though rare north of Cape Cod, they are apparently more numerous in southern New
England than in the Middle Atlantic States and are most abundant in Virginia and
North Carolina.%2
Synonyms and References:
Clupea mediocris Mitchill, Rep. in part on Fishes of New York, 1814: 20 (orig. descr.; type local. presumably
New York City; type lost); Mitchill, Trans. Lit. philos. Soc. N. Y., Z, 1815: 450 (descr., New York
Bay along Staten Island); McDonald iz Goode, et a/., Fish. Fish. Industr. U.S., 1 (3), 1884: 607,
pls. 216A, 216B (discus., names, distr., abund., econ. import., migr.).
Clupea mattowaca Mitchill, Trans. Lit. philos. Soc. N. Y., Z, 1815: 451 (orig. descr.; type local. Long Island,
New York; commonly caught in autumn, size).
Alosa mattowacca DeKay, N.Y. Fauna, Pt. 4, Fishes, 1842: 260, pl. 40, fig. 127 (descr., New York; fig.
shows no projecting lower jaw, though descr. says “‘lower jaw longest”).
Alosa lineata Storer, Proc. Boston Soc. nat. Hist., 2, 1848: 242 (orig. descr.; type local. Massachusetts); Storer,
Fish. Mass., 1867: 162, pl. 27, fig. 2 (descr., when and how taken, food value).
Clupea mattowocca Giinther, Cat. Fish. Brit. Mus., 7, 1869: 438 (refs., descr.).
Pomolobus mediocris Uhler and Lugger iv Rep. Comm. Fish. Md., ed. 1, 1876: 159; ed. 2, 1876: 136 (descr.,
synon., Potomac R., size); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 425; 47 (4),
1900: fig. 188 (descr., range); 47 (3), 1898: 2810 (correc. of earlier acct.); Smith, Bull. U.S. Fish.
32. A listing of this herring among the clupeoid fishes of Brazil (25:190) was based on a specimen 170 mm long from
an “‘ancient collection,” without mention of the place of collection. The description offered suits P. mediocris quite
well. However, as this herring does not seem to occur on the Atlantic coast in semitropical waters, as in southern
Florida, one is justified in questioning whether it actually was taken in Brazil.
21*
Bia Memoir Sears Foundation for Marine Research
Comm. (1897), 27, 1898: 91 (Woods Hole, Massachusetts); Smith and Bean, Bull. U S. Fish Comm.
(1898), 18, 1899: 183 (Potomac R., weight); Evermann and Kendall, Rep. U. S. Comm. Fish. (1899),
1900: 55 (refs., Florida); Linton, Bull. U.S. Fish Comm. (1899), 29, 1901: 438 (parasites); Bean,
T.H., 7th Rep. Forest Comm. N. Y. (1901), 1902: 301, fig., also color pl. (synon., names, size, New
York); Jordan and Evermann, Amer. Food Game Fish., 1902: 102, fig. (names, habits, size, descr.); Bean,
T.H., Bull. N.Y. St. Mus., 60, Zool. 9, 1903: 197 (synon., descr., local and techn. names, size, habits,
range); Fowler, Rep. N. J. St. Mus. (1905), 2, 1906: 95, 2 figs. (descr., habits, size, refs.); Smith, N. C.
geol. econ. Surv., 2, 1907: 121, fig. 40 (refs., descr., range, North Carolina, names, econ. import.); Ken-
dall, Occ. Pap. Boston Soc. nat. Hist., 7, 1908: 37 (refs., New Engl. locals.); Halkett, Check List Fish.
Canada, 1913: 47 (Campobello I., New Brunswick); Regan, Ann. Mag. nat. Hist., (8) 29 (112), 1917,
300 (descr.); Bigelow and Welsh, Bull. U.S. Bur. Fish. (1924), 40 (1), 1925: 105 (descr., size, range,
food, Gulf of Maine); Wilson, Proc. U.S. nat. Mus., 64(17), 1925: 12 (host of parasitic copepod);
Hildebrand and Schroeder, Bull. U.S. Bur. Fish. (1927), 43 (1), 1928: 83, figs. 42, 43 (synon., descr.,
distinct. char., food, migr., other habits, Chesapeake Bay); Jordan, Manual Vert. Anim. NE U.S.,
ed. 13, 1929: 40 (diagn., range); Truitt, Bean, and Fowler, Bull. Md. Conserv. Dep., 3, 1929: 49
(descr., Maryland); Breder, Field Bk. Mar. Fish. Atl. Cst., 1929: 65 (relation., range, habits, size);
Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 42 (range, synon.).
Doubtful References:
Clupea pusilla Mitchill, Rep. in part on Fishes of New York, 1814: 20 (orig. descr.; type local. presumably
New York; type lost); Mitchill, Trans. Lit. philos. Soc. N. Y., Z, 1815: 452 (descr.).
Negative Reference:
Pomolobus mediocris Evermann and Hildebrand, young, not of Mitchill, but P. pseudoharengus (Wilson), Proc.
biol. Soc. Wash., 23, 1910: 158 (Chesapeake Bay locals.).
Pomolobus aestivalis (Mitchill) 1814
Blueback Herring, Blueback, Glut Herring, Blue Herring, Summer Herring,
Blackbelly, Alewife, Kyak
Figures 69, 76, 77, 80, 81
Study Material. A total of 75 or more specimens, 33-295 mm TL, 27-240 mm
SL; from off Liverpool, Nova Scotia; Freeport and Portland, Maine; Gloucester,
Cape Cod, and Woods Hole, Massachusetts; Noank, Connecticut; New York Bay,
N.Y.; Washington, D. C.; Chain Bridge, Bryans Point, Plum Point, Havre de Grace,
Patuxent River, Barren Island, and Smith Point, Maryland; Rappahannock River and
Buckroe Beach, Virginia; Avoca and Beaufort, North Carolina; and St. Johns River,
Florida; in USNM collections.
Distinctive Characters. See Relationships, p. 330.
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens, 27-240 mm SL.
Body: depth 21-33.5. Interorbital: width 3.7—6.0.
Caudal peduncle: depth 8.0-10.5. Maxillary: length 9.0-13.2.
Head: length 21.5—30. Pelvic fin: length 9.4—-13.6.
Snout: length 3.8—5.8. Pectoral fin: length 15.5—21.5.
Eye: diameter 5.0-9.5.
Fishes of the Western North Atlantic 225
Scales: 46-54. Fin rays: dorsal 15-20; anal 16-21;
Ventral scutes: 31-36. pectoral 14-16.
Gill rakers: on lower limb 41-51 in Vertebrae: 49-53 (35 specimens).
adults, most frequently 45-50.
Bopy rather slender, moderately compressed, its greatest thickness about 2.5 in
the depth, its depth 3.0—-4.6 in SL.
Scates only moderately adherent, generally lost in young; with crenulate membra-
nous edge preceded by scarcely visible striae: about 13 or 14 longitudinal rows of
7 ES ey
iy ) Woe hy}
Wye sinn
Ficure 80. Pomolobus aestivalis, 265 mm TL, 10.6 inches, USNM 32639.
scales on body between base of pelvic and base of dorsal. VenrraL scures rather
strong, 18—21 in advance of pelvic fins and 12-16 behind them.
Heap frequently equal to depth, 3.33-4.6 in SL. Snour 3.6—5.8 in head. Eye
small, equal to or shorter than snout in adults, 3.0-6.0. INTERORBITAL 4.0—7.0. CHEEK
notably longer than deep (Fig. 76c). Mouru oblique. Upper jaw with a definite
notch in adults. Lower jaw not projecting strongly, partly included, not entering into
dorsal profile, its upper margin within the mouth rising abruptly and forming a
more or less definite obtuse angle (Fig. 76 c). Maxitary broad, its greatest width about
75 °/, of length of snout, rounded posteriorly, reaching to, or more often somewhat
beyond, vertical from middle of eye, 2.0—-2.7 in head. Git rakes at angle of first arch
fully 0.75 of diameter of eye; increasing in number with age: 28-36 on lower limb in
examples 30-49 mm SL, 30-39 in specimens 50-69 mm, 35-41 in specimens 70—
89 mm, 38-44 in specimens 90-109 mm, 42-48 in specimens 110-129 mm, 42—50
in specimens 130-149 mm, and 42-52 in examples 190-260 mm (adults). TEETH on
premaxillary exceedingly minute in young, missing in adults; those on free rim of
maxillary rather prominent in young and often slightly evident posteriorly in adults;
326 Memor Sears Foundation for Marine Research
those present on lower jaw persistent anteriorly, though not with free points in adults,
a permanent narrow band of granular teeth on median line of tongue.
Dorsat fin with a definitely concave margin, its origin scarcely an eye’s diameter
in advance of the pelvic fin and usually a little nearer to margin of upper jaw than to
vertical from last ray of anal; a narrow sheath of scales at base, incomplete, leaving
base of last 4 or 5 rays exposed. Caupat deeply forked, the lower lobe the longer, a
little longer than head. Anat low, with straight margin, its origin about equidistant
between base of pelvic and base of caudal; a narrow sheath of scales at base. Petvic
fin inserted near ventral edge and about equidistant between base of pectoral and
origin of anal, 1.8-2.6 in head. PecroraL somewhat higher on side, reaching about
0.66 of the distance to base of pelvic, 1.2-1.7 in head.
Color. Fresh fish bluish above and sides silvery. Rows of scales on back and upper
part of side with more or less distinct dark lines in adults. Shoulder with a dark spot in
the larger examples, generally missing in individuals less than 100 mm long. Fins plain,
generally greenish or yellowish in life. Peritoneum usually, though not always, dark.
Size. Bluebacks are known to attain a length of about 380 mm (15 in.) anda
weight of approximately 13 ounces. However, the usual size of these fish in commercial
catches as observed in the Chesapeake Bay area is under 300 mm (12 in.) and a half
pound or less in weight.
Development and Growth. The eggs and early young were described by Kuntz
and Radcliffe from material taken at Woods Hole, Massachusetts (76: 123-126).
The eggs are demersal, somewhat adhesive, semitransparent and yellowish, round, and
approximately a millimeter in diameter. The development is rapid; hatching occurs
at ordinary laboratory temperature about 50 hours after fertilization.
The newly hatched larvae (Fig. 81 a) are close to 3.5 mm long and quite slender.
In about four days the yolk is all absorbed and the larvae have grown to a length of
about 5 mm. The vent in these larvae, as in other clupeids, is situated far back, about
at the beginning of the posterior fifth of the body.
The smallest specimen at hand, 20.5 mm long (16.5 mm SL), taken June 17 (1873)
at Washington, D. C., is still very slender (Fig. 81 8), its depth about 7.5 times in
SL. Its eye is rather longer than the snout, the maxillary extends nearly to the pupil,
and the fins are all developed, the pelvic being inserted about under the origin of the
dorsal. In a 25-mm specimen (20 mm SL), the depth is contained 5.0 times in the length,
and the pelvic fin is well behind the origin of the dorsal (Fig. 81 c), as in adults.
The 30-mm fish illustrated by Kuntz and Radcliffe (76: fig. 100) is much deeper than
specimens of somewhat similar size now at hand, its depth being about 3.6 times in
SL; in specimens from the Rappahannock River, Virginia, 34, 35, and 38 mm (28,
29, and 30mm SL), the depth is respectively 4.6, 4.65, and 4.3 times in SL, and
specimens even up to 45 and 50 mm long are more slender than the 30-mm one illus-
trated. If the proportions are correctly shown, it seems highly probable that the
illustrated specimen is not this species.
When the young fish reach a length of about 80-90 mm they are usually pro-
Fishes of the Western North Atlantic 227,
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Ficure 81. Pomolobus aestivalis growth stages; all from Potomac River. a, newly hatched larva, 3.5 mm,
USNM 12830; 8, young, still without scales, 20.5 mm TL, 16.5 mm SL, USNM 12830; c, young with
scales developing, 25mm TL, zomm SL, USNM 12830; p, young adult, 45 mm TL, USNM 69834.
a drawn by Homer Wheelon; B, c by Ann S. Green; p by Nancy D. Patton.
328 Memoir Sears Foundation for Marine Research
portionately about as deep as large examples, though the diameter of the eye continues
to exceed the length of snout until they are nearly full grown. The gill rakers (see
Description) increase in number with age, quite rapidly in the young up to about 100 mm
SL. It has been indicated also that the teeth are more prominent in the young than in
the adult, and that they disappear entirely from the premaxillary with age.
In waters tributary to Chesapeake Bay, in the Potomac River in particular, where
most of the young fish were collected and measured, the rate of growth of the young
during their first summer and subsequently is shown in a table by Hildebrand and
Schroeder (59: 88): 26 taken in June, 30-37 mm; 31 in July, 30-59 mm; g3 in
August, 35-64 mm; Io1 in September, 40-69 mm; 232 in October, 40-74 mm;
and 181 in November, 50-74 mm. Collections of young of the same year-class from
Chesapeake Bay consisted of: 276 taken in January, 65-119 mm long; 280 in Feb-
ruary, 65-119mm; 773 in March, 65-115 mm; and 8 in May, 95-120 mm in
length. These data suggest that the young grow to a length of about 50-75 mm
(2 or 3 in.) while living in fresh water and to a length of 65-120 mm (2.6—4.8 in.)
when a year old.
Some older fish were taken in Chesapeake Bay, often in the same otter trawl hauls
with the younger ones, but they are not separable into year-classes by the lengths
given in the table referred to above. It is highly probable that, with few exceptions,
these older and larger fish are approaching an age of two years. The collections con-
sisted of: 8 taken in January, 160-194 mm in length; 15 in February, 150-209 mm;
and 8 in March, 140-200 mm. If the six largest examples, which may be about
three years old, are excluded, the range for the fish that apparently are two years
old is 140-184 mm. If a length of around 200 mm is attained at three years of age,
it is reasonable to assume that many individuals will attain a length of 250 mm (10 in.)
when four years old. As adult gravid fish often are 250 mm and even less, it is highly
probable that many individuals become sexually mature and perform their first migra-
tion to the spawning grounds at an age of four years.
Spawning, Migrations, and Habitat. Bluebacks are anadromous, but apparently
they do not ascend rivers as far as their near relative, P. pseudoharengus. In the vicinity
of Woods Hole, spawning takes place in fresh to slightly brackish ponds having an
outlet to the sea; also, Bluebacks enter at least the mouths of the larger rivers in
New England.
This species usually arrives in the coastal waters a month or so later than pseudo-
harengus—for example, in the lower parts of Chesapeake Bay during the first half of
April and in the upper sections of the Bay about two weeks later. Hildebrand and
Schroeder (59), in speaking of the migration in Chesapeake Bay, stated: “The peak
of the catch of bluebacks is usually taken between April 1 and 20 in the lower bay.
The numbers decrease throughout May, until after June 1 only stragglers are caught.”
The time of arrival varies from year to year by as muchas a couple of weeks, depending
upon the temperature of the water, which apparently must reach about 70°F before
Bluebacks will enter. Northward, of course, the runs take place correspondingly later.
Fishes of the Western North Atlantic 329
Spawning apparently begins very soon after the fish enter fresh or brackish water.
Judging from the size and development of the young and from what is known concern-
ing their migration locally, spawning in the Potomac seems to be confined principally
to the last half of April and the first half of May. Ripe fish were taken in July (1915)
at Woods Hole, Massachusetts (76: 123); while most of the females were spent,
considerable numbers of each sex were still unripe.
The spent fish, in the Chesapeake Bay area at least, return to the ocean soon after
spawning (59: 86). In the Gulf of Maine, “the spent fish, like alewives, return to the
sea shortly after spawning. Probably these are the bluebacks taken at Woods Hole
and north of Cape Cod in September and October. The winter home of our blue-
backs is unknown; probably like their relatives the sea herring, they move out from
land and pass the cold season near the bottom” (Bigelow and Welsh, 26: 113). The
capture of seven large Bluebacks, up to 375 mm (15 in.) in length, on March 5
(1931) about 70 miles off Barnegat, New Jersey (14: 377), and the acquisition of nine fish
by me January 18 (1926) in the market and reported by the dealer to have been taken
off Beaufort Inlet, North Carolina, with “sink nets’ (often operated in fairly deep
water), tend to substantiate the view that they winter near the bottom. The latter
fish, one female and eight males, all contained well-developed gonads. Bigelow and
Welsh reported that large schools of Bluebacks are often seined off the coast of Maine
“outside the islands” during summer and fall and that they consist mostly of ‘‘2-year-old
fish not yet sexually mature’’ (16).
In the Potomac, where the young are abundant below Little Falls throughout
the summer, fish 20 mm long were taken there as early as June 17, and none less than
30 mm was secured in July. Upon the approach of cool water, October and November
in the Chesapeake Bay area, the young leave the fresh waters; they are then about
five to six months old, and the greater part of them passes through Chesapeake Bay and
out to sea in company with graybacks and shad of a similar age. However, some of them
stop to spend their first winter in the deeper parts of the Bay, and a few seem to spend
their second winter there. However, no young have been listed as occurring off-
shore, from Chesapeake Bay southward. Pearson did list “Pomolobus sp.” as having
been ‘‘observed”’ in trawl catches off the coast of Virginia and North Carolina (103: 18);
whether these fish were young or adults was not stated.
Food. The food (16: 113) consists of plankton, copepods, pelagic shrimp, and
young launce, and no doubt other fish fry. .
Enemies. Undoubtedly the principal enemies of young Bluebacks are the predatory
fishes inhabiting the fresh and brackish water they occupy the first several months
of life. As they school and move into salt water en route to the sea, marine shore
fishes undoubtedly take their toll. Welsh and Breder found that weakfish (Cynoscion
regalis) taken at the head of the Acushnet River, Massachusetts, were gorged with
the young of this species (132: 159). While the schools migrate from the sea to fresh
water, large predatory fishes again must take a toll. Upon arrival in inshore waters
and in bays and streams, large numbers are captured by man.
330 Memoir Sears Foundation for Marine Research
Parasites. The parasitic acanthocephalan Echinorhynchus acus and the nematode
Heterakis foveolata have been listed (123: 742). A specimen with a large colonial
hydroid, Obelia commensuralis, attached to its back has been reported for Beaufort,
N. C. (50: 1-6).
Relationships. This species and pseudoharengus are closely related, but they differ
prominently from chrysochloris and mediocris in the more numerous gill rakers. In general,
aestivalis has a more slender body and a smaller eye than pseudoharengus. However,
extensive overlapping occurs in a series of measurements of specimens from various
localities, even though specimens of equal size are measured. But in any one area, as
in Chesapeake Bay, the separation of the two species by a combination of the two
characters mentioned generally is not difficult.
Northward, as in New England, the species run somewhat more slender, and often
some northern graybacks are quite as slender as some of the Bluebacks from the southern
part of the range. Although the proportionate size of the eye is a somewhat more reliable
character than depth of body, some specimens are difficult to distinguish by this char-
acter also. Fresh examples generally are distinguishable by the color, the Bluebacks
being bluish above, the graybacks grayish green. However, this difference in color
disappears in preserved specimens. The Bluebacks usually have more gill rakers and
vertebrae than the graybacks, but again intergradation occurs in each instance; and the
use of the number of gill rakers is made more complex because of their increase in
number with the age of the fish. The range and intergradation of all the proportions
and enumerations mentioned are shown in the Description.
The peritoneum (lining of abdominal cavity) is said to be black in the Bluebacks
and pale or pale gray in the graybacks. Because this character was supposed to be diag-
nostic, it has been used in keys many times, but in the numerous preserved specimens
at hand, it was found unreliable; Herbert E. Warfel, who has worked with Pomolobus
in Connecticut, has informed me (verbally) that this character is unreliable in fresh
material also. It cannot be trusted, because the peritoneum in the Bluebacks varies all
the way from black to pale or pearly gray.
It is evident from the foregoing discussion that aestivalis and pseudoharengus often
can be distinguished only by a combination of characters, and that, even then, some
troublesome examples occur, with small specimens more difficult to identify than
large ones.
Abundance and Commercial Importance. The relative abundance of the Bluebacks
and graybacks is not definitely known. Although many fishermen recognize the two
species, they are not separated in the market, and the catches are listed merely as
‘‘alewives”’ in the government’s statistical reports. It has been stated that the two species
are about equally abundant in Chesapeake Bay and that each contributes an equal
share to the commercial catch (59: 91); also, the Bluebacks are more abundant than
the graybacks in the southern states and less numerous northward (72, 1896: 426).
For the Gulf of Maine, Bigelow and Welsh found it difficult to arrive at a fair idea
of the relative abundance of the two (16: 112); although it is generally agreed that the
Fishes of the Western North Atlantic peewe
graybacks outnumber the Bluebacks in the northern part of their range, these authors
were certain that large catches of Bluebacks were taken in the Gulf of Maine from time
to time. In Nova Scotia the Bluebacks are rather uncommon and of slight value whereas
the graybacks are common and of considerable economic importance (129: 55). Some
statistics showing the combined catches of the two species, designated “‘alewives,” are
given in the account of the grayback (p. 338).
Fluctuations, Methods of Fishing, and Artificial Propagation. The remarks in the ac-
count of the grayback concerning fluctuations, effects of fishing, methods of fishing,
and artificial culture, also apply in general to the Blueback.
Range. The range of this species extends all the way from Cape Breton, Nova
Scotia (15: 107, ftn. 92), to the St. Johns River, Florida; it is the most numerous
southward.
Synonyms and References:
Clupea aestivalis Mitchill, Rep. in part on Fishes of New York, 1814: 21 (orig. descr.; type local. presumably
New York; type lost); Mitchill, Trans. Lit. philos. Soc. N. Y., Z, 1815: 456 (descr.); Jordan and Gil-
bert, Bull. U. S. nat. Mus., 16, 1882: 267 (cf. C. vernalis Mitchill = C. pseudoharengus Wilson; distinct.
char., range, synon.); Bean, T.H., Proc. U.S. nat. Mus., 6, 1883: 366 (migr. in Chesapeake Bay);
McDonald iz Goode et a/., Fish. Fish. Industr. U.S., Zz, 1884: 579-588, pls. 209, 210 (hist. of
nomencl., abund., geogr. distr., food, reprod., size, uses); Goode, Amer. Fishes, 1888: 393 (discus.
relation. with P. pseudoharengus; names, abund., spawn., etc.).
Pomolobus aestivalis Jordan and Evermann. Bull. U.S. nat. Mus. 47 (1), 1896: 426; 47 (4), 1900: fig. 190
(cf. P. pseudoharengus; descr., synon.); Smith, Bull. U.S. Fish. Comm. (1897), 77, 1898: 91 (Woods
Hole, Massachusetts); Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 55 (St. Johns
R., Welaka, Volusia Bar at Lake George, and Lake Monroe, Florida); Smith, N.C. geol. econ.
Surv., 2, 1907: 124, fig. 42, pl. 4 in color (synon., descr., range, habits, distinct. char.); Kendall,
Occ. Pap. Boston Soc. nat. Hist., 7, 1908: 38 (refs., New England); Evermann and Hildebrand, Proc.
biol. Soc. Wash., 23, 1910: 158 (Bryans Pt., Maryland); Sumner, ef @/., Bull. U.S. Bur. Fish. (1grr),
31 (2), 1913: 742 (refs., Woods Hole, Massachusetts, parasites); Regan, Ann. Mag. nat. Hist., (8) 79,
1917: 300 (descr.); Kuntz and Radcliffe, Bull. U.S. Bur. Fish., 35, 1918: 123, figs. 87-100 (develop.
of egg and young; fig. 100 apparently not this species); Welsh and Breder, Bull. U.S. Bur. Fish. (1923),
39, 1924: 159 (eaten by Cynoscion regalis); Bigelow and Welsh, Bull. U. S. Bur. Fish. (1924), 4o (1),
1925: 110, fig. 45 (descr., size, range, Gulf of Maine); Nichols and Breder, Zoologica, N. Y., 9 (1),
1927: 39, fig. (distr. New York, s. New England, life hist., develop., size); Hildebrand and Schroeder,
Bull. U.S. Bur. Fish. (1927), 43 (1), 1928: 85, figs. 44-50 (synon., descr., cf. P. pseudoharengus,
habits, growth rate of young, commerc. import., range, Chesapeake Bay); Truitt, Bean, and Fowler, Bull.
Md. Conserv. Dep., 3, 1929: 50 (descr., Maryland); Breder, Field Bk. Mar. Fish. Atl. Cst., 1929: 66,
fig. (range, migr., spawn., size); Jordan, Evermann, and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930:
42 (names, synon., range); Greeley, 24th Rep. N. Y. Conserv. Dep. (1934), Suppl., 9, 1935: 89, pl. 1
color (midsection of Hudson R.; cf. P. pseudoharengus); Vladykov and McKenzie, Proc. N.S. Inst.
Sci., I9 (1), 1935: 55, fig. 29 (Nova Scotia); Gudger, Amer. Mus. Novit., 945, 1937: 1-6 (colonial
hydroid on back); Greeley, 26th Rep. N. Y. Conserv. Dep. (1936), Suppl., 11, 1937: 73 (“summer
herring”), 91 (abund. Hudson R., migr. of adults and young); Greeley, 28th Rep. N. Y. Conserv. Dep.
(1938), Suppl., 15 (2), 1939: 82 (Long Island, New York); Bigelow and Schroeder, Bull. U.S. Bur.
Fish., 48 (20), 1936: 327 (adults captured off Barnegat, New Jersey); Anonymous, Fish. Resourc.
U.S., Senate Doc. 51, 1945: iii, 66, fig. (migr., range, commerc. import., methods of capture, causes for
decline, remedies; consid. with P. pseudoharengus, not separately); Bigelow and Schroeder, Fish. Bull.
(74) U.S. Fish Wildlife Serv., 53, 1953: 106-107 (descr., habits, Gulf of Maine).
Pomolobus cyanonoton Bean, T. H., 7th Rep. Forest Comm. N. Y. (1901), 1902: 305, fig. (synon., migr., time
and place of spawn.); Bean, T. H., Bull. N. Y. St. Mus., 60, Zool. 9, 1903: 202 (synon., migr., local
232 Memoir Sears Foundation for Marine Research
occur., common names); Fowler, Rep. N. J. St. Mus., 2, 1905: 98, figs. (male and female fig., dist. char.,
local occur., synon.).
Pomolobus sp. Pearson, Invest. Reps. U.S. Bur. Fish. (1932), Z (10), 1936: 18 (in trawl catch).
Doubtful References:
Clupea fasciata LeSueur, J. Acad. nat. Sci. Philad., 1 (2), 1818: 233 (orig. descr.; type local. Sandwich,
Massachusetts).
Alosa tyrannus De Kay (not of Latrobe), N. Y. Fauna, Pt. 4: Fishes, 1842: 258, pl. 13, fig. 38 (descr., occur.
along coast; plainly not menhaden; descr. and fig. suggest P. aestiva/is).
Alosa cyanonoton Storer, Proc. Boston Soc. nat. Hist., 2, 1848: 242 (orig. descr.; type local. presumably Massachu-
setts); Storer, Fish. Mass., 1867: 161, pl. 27, fig. 1 (descr., names, occur.).
Pomolobus pseudoharengus (Wilson) 1811
Grayback,** Gray Herring, White Herring, Branch Herring, Bigeye Herring,
Freshwater Herring, Sawbelly, Alewife, Kyak, Gaspereau
Figures 76, 77, 82, 83
Study Material. A total of 65 or more specimens, 30-330 mm TL, 23.5—268 mm
SL, from Cape Breton, Nova Scotia; St. Andrews Bay, New Brunswick; Freeport and
Portland, Maine; Falmouth and Woods Hole, Massachusetts; Noank, Connecticut;
Long Island, New York; off Wildwood, New Jersey; and various localities in the
Chesapeake Bay area.
Distinctive Characters. See Relationships under aestivalis, p. 330.
Description. Proportional dimensions in per cent of standard length, and counts,
based on 65 or more specimens, 23.5-268 mm SL.
Body: depth 23.5-35.5. Scales: 42-50.
Caudal peduncle: depth 8.5—12. Ventral scutes: 30-36.
Head: length 22.6—34.8. Gill rakers: on lower limb 38-43 in
Snout: length 4.55—7.6. adults (occasionally 44).
Eye: diameter 5.0-12. Fin rays: dorsal 15-19; anal 17-21;
Interorbital: width 3.45—7.3. pectoral 13-16.
Maxillary: length 10.5—15. Vertebrae: 46-50 (36 specimens).
Pelvic fin: length 10.4-15.8.
Pectoral fin: length 14.6—22.6.
Bopy compressed, its greatest thickness usually about 2.75 in its depth, the depth
2.8—4.25 in SL.
33- Grayback has been chosen as the leading name for P. pseudoharengus and blueback for P. aestivalis because these
names in a measure are descriptive, distinctive, and are perhaps understood by more people within the range of
the species than any others. Alewife certainly is not distinctive, as it sometimes includes both species mentioned,
and in some places it is applied to the menhaden, Brevoortia. Branch herring and glut herring are distinctive in
the Chesapeake Bay area, but these names are not well known northward. However, alewife is used in subsequent
pages to include both of these species in citing commercial statistics, for the species are not separated in the market,
where they are sold either as alewives or as river herring. [The American Fisheries Society recommends “Ale-
wife” for pseudoharengus.—yY.H.0.]
Fishes of the Western North Atlantic 223
Scates only moderately adherent, often lost in preserved specimens; apparently
not different from those of P. aestivalis; 14 longitudinal rows between base of pelvic
and base of dorsal. VENTRAL scuTEs strong, 18-21 in advance of pelvics, 12-16
behind them.
Heap less than depth, often fully by diameter of eye, 2.8-4.2. SNouT 4.1-6.0.
Eye large, longer than snout at all ages, 2.5-4.0. INrERoRBITAL 4.0—7.3. Mourn
oblique, as in P. aestivalis,; margin of lower jaw within mouth with a somewhat
broader angle (Fig. 76D); other characters about the mouth, including Trrrn, as in
Y
Ficure 82. Pomolobus pseudoharengus, 300 mm TL, from a market in Washington, D.C., USNM 25197.
P. aestivalis. MaxtLiary 1.8—2.8 in head. GiLt RakeERs at angle of first arch about 0.66
of diameter of eye, increasing in number with age; 25—33 on lower limb in examples
30-49 mm SL, 32-36 in specimens 50-69 mm, 30—39 in specimens 70-89 mm, 35-38
in specimens 90-109 mm, 36—40 in specimens 110-129 mm, 36—40 in specimens
130-149 mm, and 38—44 in examples 210-270 mm (adults).
Dorsat fin with a straight to slightly concave margin, its origin definitely less than
an eye’s diameter in advance of pelvic fin and usually nearer to margin of upper jaw
than to vertical from last ray of anal. Caupat deeply forked, the lower lobe a little longer
than upper lobe, often exceeding head length by nearly the diameter of eye. Anat low,
with an almost straight margin, its origin about equidistant between base of pelvic
and base of caudal. Petyic fin inserted near ventral edge and usually equidistant be-
tween base of pectoral and origin of anal, 1.75—2.5 in head. Pecrorat fin somewhat
higher on side, reaching about 0.66 of the distance to pelvic in adults and nearly to
pelvic in some small examples, 1.1—2.0 in head.
Color. Fresh specimens, grayish green above. Sides silvery. Rows of scales on
upper part of side sometimes with more or less definite dusky lines in adults. A small
aa4 Memor Sears Foundation for Marine Research
dark spot at shoulder, often missing in specimens less than 100 mm long. Fins all
plain, slightly yellowish to green in life. Peritoneum generally pale or silvery, often
with dark punctulations, and sometimes quite dusky.
Size. The Graybacks, like the bluebacks (qestiva/is), attain a length of about
380 mm (15 in.) and a weight of about 14 ounces; the usual length of market fish, as
observed in the Chesapeake Bay area, averaged only about 275 mm (11 in.) in length
and about a half pound in weight.
Although Graybacks generally grow to a length of 275-300 mm (11-12 in.) in
salt water along the Atlantic coast, their usual length in Lake Ontario (178: 188)
is less than 150mm (6 in.). The same writer mentioned fish only 100-125 mm
(4-5 in.) long with roe as further evidence of stunted growth resulting from the
local conditions under which the fish were living.
Development and Growth. The eggs are 1.25 mm in diameter and very glutinous
when first laid, adhering to brush, rope, stones, piling, and other objects (11g: 123).
The incubation period extends over six days at a mean water temperature of 60°F.
Recently hatched larvae are very slender and extremely transparent, and have
only a row of pigment spots on the side of the tail (173: 506, pl. 1, fig. 8). A two-
day-old fish is about 5.0mm long, and Ryder’s figure shows the greatest depth, in-
cluding the fin folds, to be about ro times in TL, with the vent (as usual in clupeoid
fishes) situated very far back at about the beginning of the posterior fifth of the body.
This agrees very well with a larva of similar age and size described as P. aestivalis
(76: 126). The appendages (pyloric caeca), connected to the intestine near the
stomach, are very small or wanting in the young but become greatly developed with age
(59: 90).
The young reach a length of 15 mm when a month old (26: 109). On the smallest
specimen at hand, 29 mm (22 mm SL), taken on June 18 (1873) at Washington,
D. C., the depth is 4.0 times in SL; in other respects it already resembles the adult;
the eye is notably longer than the snout and remains so throughout life. When the
fish reach a length of about too mm, the proportionate depth is more or less the
same as in adult fish.
The gill rakers (see Description) increase in number with age, this increase being
especially rapid in the young up to about 100 mm SL. In this species, as in P. aestivalis,
the teeth are more prominent in the young than in the adults and disappear from the
premaxillary with age.
The young reach a length of 50-100 mm (2-4 in.) by autumn in streams trib-
utary to the Gulf of Maine (76). This rate of growth is about the same as that
shown by Hildebrand and Schroeder (59: 91) in a table based on collections made in
streams tributary to Chesapeake Bay (especially the Potomac River). This table in-
cludes: 26 young for June, 45-74 mm; 105 for August, 50-84 mm; 134 for Sep-
34. Early larvae and some young adults, supposedly of P. psewdoharengus, were described and figured by Prince
(r05:103-108, pls. 8, ro, in part). It seems doubtful whether his identifications were correct, for his drawings of
young adults, compared with young adults from the Potomac, now before me, show the body to be much deeper
and the mouth nearly vertical.
iS
‘ y
fe
‘ain
949009
NN
4 , I. ERR
ase
i es is we
LRP
i
Ficure 83. Pomolobus pseudoharengus, from Potomac River. Top, young, showing scales developing, 29 mm TL,
22mm SL, USNM 85668; sorrom, young adult, 47.5 mm TL, USNM 69833. upper drawn by Ann S.
Green; Lower by Nancy D. Patton.
tember, 50-79 mm;* 138 for October, 55-94 mm; 221 for November, 55-109 mm;
and 19 for early in December, 60-79 mm TL. In Chesapeake Bay, young that ap-
peared to be in their first winter, as judged from their small size, were 75-140 mm in
January, 80-114 mm in February, and 80-145 mm in March. The young fry taken
in the Bay during the winter varied so widely in length that it is clear they represented
more than one year-class. The 262 older fish taken during the winter ranged between
11$—224 mm, or 115-194 mm, if the three largest examples are excluded.
The foregoing data seem to show that the young of this species grow to a length
of 55-109 mm (2.20—-4.36 in.) while in fresh water, to about 65-145 mm (2.6—
5.8 in.) at one year of age, and to about 115-195 mm (4.6—7.8 in.) at two years of
35- The largest young of the season evidently are not represented in this collection.
336 Memoir Sears Foundation for Marine Research
age. Mature gravid fish often are only 250 mm (10 in.), or even less, and if a length
of 200 mm (8 in.) or more is attained at three years of age, it is probable that many
become sexually mature and perform their first migration to the spawning grounds
when four years old. However, Bigelow and Welsh (16: 110) have reasoned that at
least some of the fish in the Gulf of Maine area reach maturity at three years of age.
This conclusion was reached because the “‘progeny”’ of adults used in restocking streams
returned in the third year after the streams were restocked.
Spawning. This species is very prolific; a total of 644 females from the Potomac
River yielded an average of 102,800 eggs per fish (11g: 123). The time of spawning
varies from some time in March in Chesapeake Bay to late April or May in Maine.
They may spawn in running water or in ponds. Ripe fish have been taken many
times in the Potomac, near Washington, where the eggs have been found adhering
to fish traps and where young occur throughout the summer. Greeley has described
their spawning in dense schools in fast water approximately two feet deep over a
bottom of coarse stones, some sand, and gravel in the Poestenkill, at Troy, New York
(47: 89). Around the Gulf of Maine, however, they spawn more commonly in ponds
(including those behind barrier beaches), though probably not so exclusively as Bigelow
and Welsh believed (16: 109).
According to Greeley (47: 89):
The act of spawning is characterized by the whirling of a pair of fish in a close spiral, usually ending as
they reach the surface with a splash and separate. The whirling is done rapidly and often only a flash of silver
is to be seen. A number of spawning pairs were observed here [presumably the Hudson] by Dr. C. W. Greene
and the writer. The sexes of the fish cannot be determined from the appearance of the fish in the water, but it
was assumed that the fish seen whirling were mated pairs. It is probable that the male is on the outside of the
spiral during the act of spawning and that the rapid circling is adopted for maintenance of contact as the eggs
are shed and fertilized. Many eggs were scattered over the bottom and had drifted downstream where they
were deposited by the current in slack water areas behind obstructions. The temperatures here May 16 (11 A.M.)
were: air, 57 degrees; water 52 degrees.
Migrations and Habitat. This species is strictly anadromous. It has been common
knowledge since colonial times that they run
indifferently up rivers as large as the St. John (New Brunswick), the Merrimac and Potomac, or streams so small
one can almost leap across, and only a few inches deep. In large rivers they may run far upstream ... or their
journey may be only a few yards, as it is in the artificial cuts that are kept open through barrier beaches to allow
the fish access to fresh water ponds behind the latter (15: 103).
The Graybacks arrive in inshore waters a month or so ahead of the bluebacks and
a week or two ahead of the shad. In Chesapeake Bay the first catches are made in late
February or early March. In southern New England (Woods Hole, Massachusetts)
they arrive sometime during March and are caught through March and April
(120: 91). However, they do not reach the rivers of Maine until late April or early
May and, as shown by the commercial catches, they are most numerous there during
the last half of May (z6: 109).
That the young return to the stream of their birth (the so-called “parent stream
Fishes of the Western North Atlantic 227
theory”) was confirmed when the progeny of gravid adults planted in depopulated
streams returned to these rivers three years after hatching (76: 110). This incident
is also cited in a subsequent paragraph as a successful method of restocking streams.
But a much more intensive study of this interesting question is needed before any
categorical statement can be made as to how generally the parent stream theory applies
in the case of the Graybacks (15: 103).
As in the case of bluebacks and shad, relatively little is known about the home
of Graybacks in the sea. The capture of 18 adult Graybacks 70 odd miles off Barnegat,
New Jersey, on March 5, 1931 (14: 327), suggests that the home may be some distance
offshore. Since these adults were taken in March, they may have been en route to some
nearby stream, hence it should not be assumed that the fish actually were taken from their
usual abode in the sea. It has been surmised for some time that the Graybacks, as well
as blueback herring and American shad, live quite a distance offshore from the rivers of
their origin and that their migrations consist of inshore and offshore movements. This
theory finds slight support in the discovery that different races spawn in different
river systems. However, further study is needed.
The young, often with bluebacks and American shad of similar age, have been
seen descending as early as June 15 in the more southerly streams fringing the Gulf
of Maine. Successive companies of fry move out of the places where they were hatched,
run down with the current throughout the summer and are at sea by autumn. In the
Potomac, which the fish cannot ascend beyond Little Falls, the young remain abundant
in the vicinity of Washington, D. C. Here they may be taken in the same seine hauls
with bluebacks and American shad until October, when the main seaward migration
takes place; however, some young remain in the river until November and a few
linger until December. Most of the young apparently pass directly through Chesapeake
Bay and out to sea, but some stop to spend their first winter in the deeper parts of the
Bay, a few seem to remain for their second winter, and very few apparently stay there
for their third winter.
The fish in the freshwater lakes endeavor to carry out the same spawning migra-
tion as those that inhabit the Atlantic coast, for the fish enter the shallow parts of
the lakes and ascend the tributary streams to spawn.
Food. The food of the young while in fresh water apparently consists principally
of diatoms, copepods, and ostracods. After they enter salt water, their chief diet seems
to be shrimps and fish. Young 50-70 mm long, taken in fresh water, had fed principally
on copepods and ostracods; and larger young, 83-178 mm long, caught in Chesapeake
Bay, had continued to feed on copepods, though the larger ones had ingested prin-
cipally mysids and fish (59: 90). Copepods, shrimps, and young squid were listed as
the food of the young by Sumner, et a/. (123: 72). Bigelow and Welsh (16: 110) have
described their chief food as copepods, amphipods, shrimps, and appendicularians, small
fish (such as herring, eel, launce, and cunner) as well as fish eggs and young of their
own. Alewives often contain diatoms, even when adult. It is conceivable that diatoms
could be screened from the water by the rather numerous, close-set gill rakers of adults.
22
338 Memor Sears Foundation for Marine Research
It has long been known that anadromous fishes, as a rule, feed sparingly or not
at all while ascending streams, and Graybacks are no exception to this rule. However,
the spent fish feed ravenously on shrimp on entering brackish water along the coast,
where anglers often hook them on an artificial fly (76).
Enemies. The young are undoubtedly preyed upon by various predatory fishes,
turtles, snakes, and birds while they live in fresh water. When they begin to school and
migrate out to sea, marine fishes and numerous other sea inhabitants very probably
destroy many. When they re-enter shallow shore water as adults in great schools, large
marine fishes of many kinds, finback whales, birds, and turtles, take a further toll.
Upon arriving inshore and in the rivers, man—who captures great quantities for food
—must be their chief enemy.
Parasites. The following have been listed by Sumner, et a/. (123: 742): Acanthoce-
phala (Echinorhynchus acus); cestodes (Rhynchobothrium imparispine) , trematodes (Distomum
appendiculatum, D. bothryophoron, D. vitellosum, and Monostomum sp.); and copepods (4r-
gulus alosae, Caligus rapax, and Lepeophtheirus edwardst).
Variations. Specimens from various localities within the range of this species
along the coast (Study Material, p. 332) show a tendency toward a more slender body
and a proportionately larger head toward the northern part of the range; and a limited
number of specimens from fresh water, Cayuga Lake and Lake Ontario (proportions
not used in Description), show this tendency to be even more pronounced there. In
general, the fish from the northern part of the range (especially from Casco Bay, Maine),
and more particularly those from fresh water, have the appearance of being more or
less emaciated. However, insufficient specimens have been studied to determine the
significance of the apparent differences mentioned. These differences may be of sub-
specific value, but more likely they are of only racial importance. The decrease in
body depth in northern specimens makes their separation from P. aestivalis more
difficult than identification of specimens from the southern part of the range.
Abundance and Commercial Importance. The relative abundance of pseudoharengus
and aestivalis is discussed in the account of the latter (p. 330). It has been claimed
by various writers that Graybacks are far less numerous now than during colonial days,
or even a half century ago. The matter was summed up for the Gulf of Maine by
Bigelow and Welsh (z6: 108) in these words:
During the past two centuries ... its numbers have declined, and its range has been restricted, both by
actual extirpation from certain streams by overfishing, by the pollution of river waters by manufacturing wastes,
and by the erection of dams that it cannot pass. However, the alewife is still a familiar fish all along our coast,
and yields an abundant catch in many of our streams.
These remarks also apply in a measure to fish in the more southern part of the range,
though the decline apparently has been less pronounced southward.
The following statistics give the catches of Graybacks and bluebacks combined,
under the name “‘Alewives.” These records show in part the decline in their abundance
during the past 50 years or so, but they also show that these fish are still abundant
and of great economic importance. There has been no compensation for any variations
Fishes of the Western North Atlantic 220
in the “fishing effort”—the number of men, vessels, and amount and efficiency of the
gear used—but it may be assumed that there was no decrease in this respect. Federal
Government statistics, which have been taken at irregular intervals, report total catches
in round numbers for the Atlantic coast of the United States as follows: 62,067,000
pounds in 1896, 52,061,000 in 1905, 35,809,000 in 1921, 35,290,000 in 1929,
and 29,542,000 in 1941. The greatest proportionate decline took place in New
England, for which the following catches are listed: 12,116,000 pounds in 1896,
8,429,000 in 1905, and 3,975,000 in 1924; however, from 1924 to 1941 the catch
there remained fairly even.
South of New England the decline in abundance began later and was more gradual
and less pronounced. For example, the Chesapeake Bay states (Maryland and Virginia),
which have furnished a larger part of the total catch of ‘‘Alewives”’ than any other section
of the coast since the beginning of this century, yielded a fairly even catch until about
1925. Thereafter a decline has been indicated by the statistics: 25,611,000 pounds in
1925, 21,129,000 in 1930, 18,884,000 in 1937, and 17,012,000 in 1941. The catch
in the South Atlantic states, from North Carolina to Florida, is also very large, has
fluctuated greatly, and has shown a slight downward trend: 15,857,000 pounds for
1897, 11,601,000 for 1902, 15,186,000 for 1918, 7,571,000 for 1923, I1,176,000
for 1929, 6,218,000 for 1937, 11,611,000 for 1938, and 9,116,000 for 1940. The
over-all catch of the two species combined has ranged between 47,300,000 and
57,700,000 pounds for the United States (1940, 1945, 1950-1953), and between
6,250,000 and 41,000,000 pounds for Canada (1940, 1945, 1950-1952).
The Graybacks and blueback herring are still abundant. In fact, every state
along the Atlantic coast exclusive of Pennsylvania, South Carolina, and Georgia, make
substantial contributions to the catch, as do Nova Scotia and New Brunswick, Canada.
Although these species are among the cheapest fish on the market, they furnish a sub-
stantial income because of their abundance. During recent years the annual yield to
the fishermen from this fishery alone has been $ 400,000—800,000 in the United States,
and $ 33,000—350,000 in Canada.
Most of the ‘“‘Alewives’’ caught are salted, but an increasingly large part of the
catch is sold in the fresh state. Although the flesh is quite bony, as in other herrings,
it is delicious and is in demand whether fresh, salted, or smoked. At the salting
houses the roe generally is separated for canning and the offal is made into fish meal
and oil.
Although these fish have become common to abundant in some freshwater lakes,
as in Lakes Ontario and Erie, they are of little economic importance there because of
their small size. However, in 1892 Smith said that they were of more economic impor-
tance in Lake Ontario than is generally realized (118: 189). He pointed out that they
had replaced more important foodfishes (young ciscoes and suckers) as bait for sturgeon
and lake trout in the trawl-line fishery, that the young were used as bait in extensive
sport fishing, and that many tons of those that wash ashore annually are used as
fertilizer.
22*
34.0 Memor Sears Foundation for Marine Research
Fluctuations. Overfishing, pollution, and impassable dams in streams have been the
chief factors in diminishing the abundance of “‘Alewives”’ in many localities, but all of
these causes can be remedied at least in some measure by restocking suitable streams
with gravid fish (16: 110; ef a/.). This has been done in Massachusetts localities, where,
in the third year after the ripe fish were introduced, the progeny appeared in those
streams.
It is not always easy to determine whether an increase or decrease in catch is due
to the amount of fishing, to special conditions created by man, to successful or unsuc-
cessful spawning, or to general conditions obtaining in nature. However, if the increase
or decrease extends over a period of several years, reasonable conclusions may be reached.
For example, the tremendous abundance of ‘‘Alewives” in the Chesapeake Bay area in
1908 and 1909, when the catches were about twice those in preceding and successive
censuses (66,690,000 and 51,425,000 pounds), may be ascribed to especially favorable
conditions during and probably after the spawning season. On the other hand, the
general decline in the fishery in New England (discussed above), despite some
fluctuations, quite surely was the result of adverse conditions created by man. There-
fore, frequent censuses are necessary to show the status of this as well as other fisheries
for the guidance of those entrusted with the promulgation of regulations for the con-
servation of the fishery.
Artificial Cultivation. “ Alewives” have not been artificially propagated regularly be-
cause it has not been deemed necessary. However, in 1882 two million fry hatched
by fish culturists were transported to the Colorado River, Texas (45: 586), an in-
troduction that apparently was not successful. As already stated, another method
of “planting” was successfully used in Massachusetts. Some depopulated streams in
that state, which had been reconditioned and made suitable, were successfully restocked
merely by releasing gravid fish. Graybacks apparently were unintentionally introduced
into Lake Ontario when shad fry were planted there (45: 588; rz8: 188).
Methods of Fishing. “‘ Alewives”’ are easily caught, chiefly with dip nets in New Eng-
land. Pools are constructed about the mouths of rivers and the gravid fish are led into
them by means of nets and stone diversions. Southward, as in the Chesapeake Bay
area and in North Carolina, they are caught chiefly with pound nets. However, they
are taken also with weirs, seines, gill nets, fyke nets, and a few with otter trawls.
Range and Distribution. The range of pseudoharengus extends from the Gulf of St.
Lawrence and Nova Scotia, where it is common (22g: 54), to North Carolina. It has
been recorded for the St. Johns River, Florida, but the first report (45: 336), as
Clupea vernalis, was apparently based on a P. aestivalis (USNM 17929, collected by
Spencer F. Baird), now before me; and the second report, so far as known, by Lénn-
berg (88: 114), may be disregarded, for this species is too difficult to distinguish
from P. aestivalis by a simple “observation.” According to Smith, ‘extensive collec-
tions of fishes in the St. Johns River, in Florida, have failed to disclose its presence”
(11g: 122). Indeed, no specimens of this species from south of the Albemarle Sound
area (where aestivalis is plentiful) have been found in the National Museum, and
Fishes of the Western North Atlantic 341
it was only sparingly represented among many specimens of P. aestivalis collected over
a period of years in the Newport River and its estuary near Beaufort, N. C. Possibly
this species does not occur south of North Carolina.
This species, as stated above under Artificial Cultivation, apparently was uninten-
tionally introduced into Lake Ontario, whence it has spread to Lake Erie and to Lake
Huron. It also inhabits Lakes Seneca and Cayuga, which it probably reached by natural
means from the ocean (45: 116), and the Oswego River system in New York State
(46: 95). It has remained common in Lake Ontario, has become common in Lake
Erie, and has penetrated Lake Huron, where it is still rare (62: 27).
Synonyms and References:
Clupea pseudoharengus Wilson, in Rees’ New Cyclopedia (Art. C/upea), 9, undated, ca. 1811: no pages (orig.
descr.; no local.; probably Philadelphia, Pennsylvania); Smith, Bull. U.S. Fish Comm. (1890), Zo, 1892:
187, pl. 29 (unintent. introd. in Lake Ontario, abund. there, diseases, mortal., uses).
Clupea vernalis Mitchill, Trans. Lit. philos. Soc. N. Y., I, 1815: 454 (orig. descr.; type local. New York;
effects of mill dams on migr. to spawn); Jordan and Gilbert, Bull. U. S. nat. Mus., 16, 1882: 267 (descr.,
range, synon.); McDonald iz Goode, et a/., Fish. Fish. Industr. U.S., 1, 1884: 579-588, pls. 207,
208 (hist. of nomencl., abund., geogr. distr., migr., food, reprod., size, uses); Bean iz Goode, et a/l., Fish.
Fish. Industr. U. S., 1, 1884: 588-593 (Lakes Cayuga, Seneca, and Ontario, origin in lakes, movements,
enemies, fatal., capture); Ryder, Rep. U. S. Comm. Fish. (1885), 1887: 506, pl. 1, fig. 8 (descr. of egg
and newly hatched young); Goode, Amer. Fishes, 1888: 393 (discus. with P. aestiva/is, names, abund.,
spawn.).
Clupea megalops Rafinesque, Amer. Mon. Mag. (1817-1818), 2(3), 1818: 206 (orig. descr.; type local.
Delaware R.).
Meletta venosa Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 374 (orig. descr.; type local. New York);
Bertin, Bull. Soc. zool. Fr., 66, 1941: 23 (ident. of types).
Pomolobus pseudoharengus Gill, Rep. U.S. Comm. Fish. (1871-72), 1873: 811 (names, range); Uhler and
Lugger, Rep. Comm. Fish. Md., ed. 1, 1876: 158; ed. 2, 1876: 135 (descr., synon., habits, Maryland;
pseudoharengus and aestivalis not distingu.); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896:
426; 47 (4), 1900: fig. 189 (descr., range, habits, synon.); Smith, Bull. U.S. Fish Comm. (1897), 17,
1898: gt (Woods Hole, Massachusetts); Bean, T.H., 7th Rep. Forest Comm. N. Y. (1901), 1902:
302, with fig. (synon., names, occur. in lakes, migr., size, methods of capture); Bean, T. H., Bull. N. Y.
St. Mus., 60, Zool. 9, 1903: 199 (synon., descr., names, range, migr., size, growth of young, econ. value,
methods of capture); Fowler, Rep. N. J. St. Mus., 2, 1905: 95 (figs. of male and female, descr., migr.,
synon.); Smith, N. C. geol. econ. Sury., 2, 1907: 122, fig. 44, pl. 3 in color (synon., descr., range, migr.,
reprod., commerc. import., North Carolina, methods of capture); Kendall, Occ. Pap. Boston Soc. nat. Hist.,
7, 1908: 38 (refs., New England); Sumner, Osburn, and Cole, Bull. U.S. Bur. Fish. (1911), 32, 1913:
742 (refs., Woods Hole, Mass., food, parasites); Regan, Ann. Mag. nat. Hist., (8) 19, 1917: 300 (descr.);
Bigelow and Welsh, Bull. U.S. Bur. Fish. (1924), 40, 1925: 107, fig. 44 (descr., size, range, Gulf of
Maine, habits, food, commerc. import.); Nichols and Breder, Zoologica, N.Y., 9 (1), 1927: 38, fig. (distr.
New York, s. New England, food, life hist., size); Hildebrand and Schroeder, Bull. U.S. Fish Comm.
(1927), 43 (1), 1928: 89, figs. 51-52 (synon., descr., relation., growth of young, food, habits, commerc.
import., range, Chesapeake Bay); Greeley, 17th Rep. N. Y. St. Conserv. Dep. (1927), Suppl., 1928: 95
pl. 3 in color (Oswego R. watershed); Jordan, Manual Vert. Anim. NE U.S., ed. 13, 1929: 40 (diagn.,
range); Truitt, Bean, and Fowler, Bull. Md. Conserv. Dep., 3, 1929: 49, fig. 15 (descr., Maryland);
Breder, Field Bk. Mar. Fish. Atl. Cst., 1929: 65 (food, spawn., size); Greeley, 18th Rep. N. Y.
Conserv. Dep. (1928), Suppl., 1929: 167 (common Lake Ontario, enters Niagara R.); Jordan, Evermann,
and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 42 (names, synon., range); Greeley and Greene,
2oth Rep. N. Y. Conserv. Dep. (1930), Suppl., 5, 1931: 82 (St. Lawrence watershed); Greeley and
Bishop, 21st Rep. N. Y. Conserv. Dep. (1931), Suppl., 6, 1932: 78 (common Lake Ontario, St. Lawrence
R., annu. mortal.); McKay, Copeia, 1934: 97 (first rec. for Lake Huron); Vladykov and McKenzie,
Proc. N. S. Inst. Sci., 79 (1), 1935: 54, fig. 28 (Nova Scotia); Greeley, 24th Rep. N. Y. Conserv. Dep.
34.2 Memoir Sears Foundation for Marine Research
(1934), Suppl., 9, 1935: 89 (Mohawk-Hudson watershed, descr. of spawn.); Bigelow and Schroeder,
Bull. U.S. Bur. Fish., 48 (20), 1936: 327 (adults off Barnegat, New Jersey); Greeley, 26th Rep. N. Y.
Conserv. Dep. (1936), Suppl., 11, 1937: 90 (abund. Hudson R.; landlocked in reservoir and lake);
Greeley, 26th Rep. N. Y. Conserv. Dep. (1937), Suppl., 12, 1938: 62 (Allegheny and Chemung water-
sheds); Greeley, 28th Rep. N. Y. Conserv. Dep. (1938), 13, 1939: 39 (in fresh water, Long Island,
New York); Greeley, 28th Rep. N. Y. Consery. Dep. (1938), Suppl., 15 (2), 1939: 82 (salt water,
Long Island, New York); Greeley, 29th Rep. N. Y. Conserv. Dep. (1939), Suppl., 16, 1940: 68 (notes
on alewife), 59-61 (abund. Lake Ontario; stunted growth in fresh water; annu. mortal., spawn.); Hubbs
and Lagler, Bull. Cranbrook Inst. Sci., 18, 1941: 27 (range, Lakes Ontario, Erie, and Huron, where
introd.); Anonymous, Fish. Resourc., U.S. Senate Doc., 51, 1945: iii, 66, fig. (migr. range, commerc.
import., methods of capture, causes for decline, remedies, consid. with P. aestiva/is); Bigelow and Schroe-
der, Fish. Bull. (74) U.S. Fish Wildl. Serv., 53, 1953: 101 (descr., habits, Gulf of Maine).
Pomolobus pseudoharengus lacustris Jordan, Manual Vert. Anim. NE U.S., ed. 1, 1876: 265 (orig. descr.;
type local. Cayuga Lake, New York; landlocked form subspecif. distinct from anadromous one).
Pomolobus mediocris Evermann and Hildebrand (not of Mitchill), Proc. biol. Soc. Wash., 23, 1910: 158 (notes
on young; found later to be pseudoharengus).
Pomolobus sp., Pearson, Invest. Rep. U.S. Bur. Fish., 1 (10), 1932: 18 (in trawl catch, off Virginia capes).
Doubtful References:
Clupea parvula Mitchill, Rep. in part on Fishes of New York, 1814: 21 (orig. descr.; type local. presumably New
York; type lost); Mitchill, Trans. Lit. philos. Soc. N. Y., Z, 1815: 452 (descr.).
Clupea virescens De Kay, New York Fauna, Pt. 4, Fishes, 1842: 252, pl. 13, fig. 37 (orig. descr.; type local.
New York Bay; type lost).
Clupea pseudoharengus Lonnberg, Ofvers. Svensk. Vet. Akad. Forh., 52 (3), 1894: 114 (observed enter. St.
Johns R., Florida in spring; probably P. aestiva/is).
Pomolobus pseudoharengus Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 55 (St. Johns R.,
after Bean 1883 and Lonnberg 1894).
Genus Brevoortia Gill 1861
Menhaden
Brevoortia Gill, Proc. Acad. nat. Sci. Philad., 1861: 37; genotype by original designation, Brevoortia menhaden
Gill equals B. tyrannus (Latrobe).
Characters. Bopy oblong, compressed. Bony scures present, the median line of
chest and abdomen with a sharp edge. Scares adherent, the exposed part much deeper
than long, the margin serrate or pectinate in adults; a series of modified scales pres-
ent next to median line on back in front of dorsal fin. Cuerk deeper than long.
Mourn large. MaxtLiary extending to or beyond middle of eye. Upper jaw with a
distinct median notch. Lower jaw included in the upper one, not projecting, its up-
per margin (within mouth) nearly straight. TerrH wanting in adults. Lower limb of
first GILL aRcH with an obtuse angle. Gitt RakERs long and numerous, increasing in
number with age, those on upper limb of first arch extending downward and over
those on upper part of lower limb. Dorsat with 17-22 rays, the last one not greatly
produced; its origin about equidistant between rim of snout and base of caudal. Ana.
with 17-25 rays, the last one little enlarged. Petvic small, with 7 rays. VERTEBRAE
about 42-50. InTEsTrNE very long. Perrroneum black.
Remarks. The species of this genus generally are recognizable by the strongly
Fishes of the Western North Atlantic 343
serrate or pectinate adherent scales, the exposed part of which is much deeper than
long; the modified series of scales next to the median line of the back in front of the
dorsal fin; the very long gill rakers on the upper limb, which project downward
over the equally long rakers of the lower limb; the very deep cheek; and the rather
small pelvic, composed of only seven rays.
The young, about 60mm TL and less, of North American representatives at least,
have minute teeth on the margin of the maxillary, but these disappear with age. The
gill rakers on the upper limb do not yet extend downward over those on the lower limb,
as in larger fish; the number, as already indicated, increases greatly with age. Further-
more, until the fish reach about 100-125 mm TL, the two series of scales, one on
each side of the median line of the back in front of the dorsal fin, do not become fully
differentiated or modified. Though the exposed margin of the scales on young may
be somewhat uneven, the scales are not definitely serrate; the serrae become somewhat
developed when the fish attain a length of about 100 mm and continue development
until they are comparatively large, but these serrations are not developed uniformly
on all parts of the fish. For some species the length and shape of these serrations or
pectinations are diagnostic characters, but fish of nearly equal size must be compared
and scales from the same part of the body must be used.
The species of Menhaden included in this work fall into two closely related pairs:
tyrannus from the Atlantic and patronus from the Gulf; smithi from the Atlantic and
gunteri from the Gulf—and one odd species, brevicaudata, from Noank, Connecticut,
related to tyrannus. Such a close relationship between species exists in other groups
as well.%
In each pair of fishes named, the range probably was once continuous but became
discontinuous when the last passageway for marine fishes across the Florida peninsula
was closed. At least these fishes do not now occur on the coasts of southern Florida,
indicating that suitable conditions do not exist there. Therefore, the separation between
Atlantic and Gulf representatives is complete. Under this separation, and apparently
under the influence of differences in environment, these fishes have become sufficiently
differentiated to constitute distinct species.
Range. The Atlantic and Gulf coasts of the United States northward to Nova
Scotia; southern Brazil to Argentina; and western Africa.
Measurements and Counts for Brevoortia. For measurements and counts not dis-
cussed below, see p. 258.
Heap beptu: for this genus, measured from the slight crossgroove at occiput to
the keel of the first ventral scute. Although this is not an exact vertical, it does
provide definite and uniform points for measuring.
Scatzs: the number in a lateral series represents the number of oblique series
(running upward and slightly backward) across the middle of the side; these should
36. Among the shad, Alosa sapidissima of the Atlantic and 4. alabamae of the Gulf; also the hickory shad, Pomolobus
mediocris, of the Atlantic and the skipiack herring, P. chrysochloris, of the Gulf and Mississippi Valley. In the
family Sciaenidae, the weakfish, Cynoscion regalis, of the Atlantic and the sand seatrout, C. arenarius, of the Gulf
(Ginsburg, 42: 83); and another pair occurs in the genus Menticirrhus (Ginsburg, unpublished).
344 Memoir Sears Foundation tor Marine Research
be counted uniformly, since they tend to become reduced in size on the back and
larger on the lower part of the side. The comparisons and figures given on these pages
are based on scales from the middle of the side below the anterior dorsal rays.
Caupat Fin: lower lobe measured from middle of caudal base.
Prcrorat FIN: axillary appendage measured from base of upper pectoral ray.
Ficure 84. Brevoortia pelvic fins: a, patronus, 215 mm TL, off Galveston, Texas, USNM 129810, see Fig.
91; B, tyrannus, 320mm TL, Chesapeake Bay, USNM 129809, see Fig. 86; c, guateri, 270 mm TL, Rock-
port, Texas, USNM 129798, see Fig. 93; p, smithi, 295 mm TL, USNM 118723, see Fig. 92; all equally
enlarged. Drawn by Ann S. Green.
Species. Five species fall within the scope of this work (see Key to Species), and
two others, aurea and pectinata, occur in southern Brazil, Uruguay, and Argentina. Still
another one, sometimes identified with syrannus, has been reported for western Africa.
I have seen no specimens of the latter and cannot add any information as regards its
relationship to American forms. In general, the scales of South American representa-
tives are scarcely reduced in size on the back and on the base of caudal, whereas in
North American species they are greatly reduced there.
Key to Species of the Western Atlantic
1a. Scales fairly regularly placed, 35-36 oblique series crossing middle of side; great-
est depth of body 30-40 °/, of SL.
2a. Scales on back and at base of caudal much smaller than those along middle
abe
Fishes of the Western North Atlantic 345
of side; pelvic fin with a definitely convex margin, the innermost ray more
than 67°/, of the length of the outermost one, the fin not pointed when
folded (Fig. 84 a, 8); upper part of opercle with prominent radiating striae;
shoulder spot followed by a variable number of smaller dark spots in adults.
3a. Ventral outline of body only moderately convex anteriorly, usually only
about half of greatest depth below a straight line extending through
lower margin of eye to middle of base of caudal; sheath of scales at base
of dorsal fin composed for the most part of a single row of scales covering
only the basal third of the longest rays when standing erect; pectoral
fin falling far short of base of pelvic fin, 3-7 vertical series of scales
between its tip and base of pelvic, the fin with 16-18 (rarely 15) rays.
4a. Maxillary reaching well beyond vertical from posterior margin of
pupil, 13-16 °/, of SL; mandible 16-19 °/, of SL; pectoral fin
failing to reach base of pelvic fin by less than diameter of eye, 3 or
4 vertical rows of scales between tip of pectoral and base of pelvic,
its length 17-21 °/, of SL; caudal fin with lower lobe about as
long as head, 25-35.5 °/) of SL. tyrannus (Latrobe) 1802, p. 346.
4b. Maxillary reaching only to a vertical from posterior margin of pupil,
12-13 °/, of SL; mandible 15.5-16.5 °/, of SL; pectoral fin failing
to reach base of pelvic fin by a distance exceeding diameter of eye,
5-7 vertical series of scales between its tip and base of pelvic, its
length 15.5-16°/, of SL; caudal fin with lower lobe shorter than
head, 22.5-25 °/) of SL. brevicaudata Goode 1879, p. 363.
3b. Ventral outline of body strongly convex anteriorly, much more than half
of greatest depth below a straight line extending through lower margin
of eye to middle of base of caudal; sheath of scales at base of dorsal fin
composed for the most part of two rows of scales covering basal two-
thirds of shortest rays when standing erect; pectoral fin often reaching
nearly to base of pelvic fin, seldom more than 1 or 2 vertical series of
scales between its tip and base of pelvic, the fin with 14-17 (usually 15
or 16) rays. patronus Goode 1879, p. 365.
Scales on back and at base of caudal not much smaller than those along middle
of side; pelvic fin with a nearly straight oblique margin, the innermost ray
not more than 67 °/, of the length of the outermost one, the fin pointed when
folded (Fig. 84 c, D); upper part of opercle with feeble radiating striae, if any;
shoulder spot not followed by smaller dark spots.
5a. Scales in only 35-46 oblique series crossing middle of side; five longitudinal
rows on side of caudal peduncle; pectoral fin sometimes reaching base
of pelvic fin, occasionally falling short of this point by diameter of pupil,
its length 17.5-21 °/, of SL, o-3 vertical series of scales between its tip
and base of pelvic. pectinata Jenyns 1842.
Rio Grande of Brazil to Bahia
Blanca, Argentina.
34.6 Memoir Sears Foundation for Marine Research
5b. Scales in 48-56 oblique series crossing middle of side: seven longitudinal
rows on side of caudal peduncle; pectoral fin failing to reach base of
pelvic fin by a space varying from half to a full diameter of eye, its length
16.7—18 °/, of SL, 3-6 vertical series of scales between its tip and base
of pelvic. aurea Agassiz 1829.
Bahia (Baia), Brazil, prob-
ably to the Rio de la Plata.
1b. Scales small, irregularly placed, about 60-75 oblique series crossing middle of
side (difficult to count); greatest depth of body 36—45.5 °/) of SL.
6a. Length of head 29—31.5 °/, of SL; maxillary reaching from below middle of
eye to vertical from posterior margin of pupil, 2.0—2.4 in head; pectoral fin
generally failing to reach base of pelvic fin by somewhat more than half of
diameter of eye, 5—8 vertical series of scales between its tip and base of pelvic,
its length 18.5—21 °/, of SL; total number of ventral scutes 30-32 (usually
30); vertebrae 45-47. smithi Hildebrand 1941, p. 372.
6b. Length of head 31-35.5 /) (usually 32-34 °/,) of SL; maxillary reaching to,
or a little beyond, vertical from posterior margin of pupil, 1.8—2.2 in head;
pectoral fin generally failing to reach base of pelvic fin by less than diameter
of pupil, 2-4 vertical series of scales between its tip and base of pelvic, its
length 19—23.5 °/, (usually 20-22 °/,) of SL; total number of ventral scutes
27-30 (usually 28 or 29); vertebrae 42-44.
gunteri Hildebrand 1948, p. 376.
Brevoortia tyrannus (Latrobe) 1802
Atlantic Menhaden, Menhaden, Mossbunker, Bunker, Fatback, Shad, Pogy, Bugfish
Figures 84-89
Study Material. At least 150 specimens, sometimes many more, 70-430 mm TL,
55-355 mm SL, collected at many localities from Massachusetts Bay to Fernandina,
Florida. Some of the data used were accumulated by me over a period of 33 years; in
addition, unpublished data from Dr. A. Bascom Hardcastle and some gathered by the
late William W. Welsh; also juveniles from Woods Hole, Massachusetts; Chesapeake
Bay; and Beaufort, North Carolina.
Distinctive Characters. The close relationship of Brevoortia tyrannus to B. patronus
of the Gulf of Mexico is shown in the account of the latter (p. 370). From B. smithi,
which occupies the same range as the Atlantic Menhaden from North Carolina
southward, tyrannus differs prominently in having larger scales with notably longer pec-
tinations; these species also differ in several other respects as shown in the account of
B. smithi (p. 374).
Description. Proportional dimensions in per cent of standard length, and counts,
based on at least 150 study specimens, 55-355 mm SL.
Fishes of the Western North Atlantic aAF
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Ficure 85. Brevoortia iis from middle of side below anterior dorsal rays. a, patronus, 215 mm TL, see Fig.
QI; B, “yrannus, 320 mm TL, see Fig. 86; c, guateri, 270 mm TL, see Fig. 93; v, smithi, 295 mm TL, see
Fig. 92; all equally enlarged.
Body: depth 30-40, usually 32-37.
Caudal peduncle: depth 8.5—11.
Head: length 29-36, most frequently
31-34.
Snout: length 5.5—9.0, usually 6.0-8.0.
Eye: diameter ca. 5.0—8.0.
Interorbital: width 6.2—-8.2.
Maxillary: length 13-16.
Mandible: length 16-19.
Anal fin: length of base 14-19, usually
15-18.
Pelvic fin: length 8.0-10.
Pectoral fin: length 17-21;
18-20.
usually
Scales: oblique series along middle of
side, 41-55, most frequently 45-
G2s
Modified scales: in a series in front of
dorsal 31-43, most frequently 33-
39-
Ventral scutes:
33:
Fin rays: dorsal 18-22, usually 19-21;
anal 18-24, usually 20-23; pec-
toral 16-18, rarely 15.
Vertebrae: 47-49, very rarely 45, 46, or
50 (194 specimens).
30-35, usually 32 or
348 Memotr Sears Foundation for Marine Research
Bopy elongate, compressed, its greatest thickness varying greatly, from nearly
half of depth in large fat fish in spawning condition to only about a third of depth in
lean half-grown examples; greatest depth generally at, or slightly in advance of, vertical
from tip of pectoral, 2.5—3.3 in SL in specimens 100 mm and upward; ventral outline
moderately convex anteriorly, usually only half of greatest depth below a straight line
through lower margin of eye to middle of base of caudal. CaupaL PEDUNCLE rather
slender, 2.9—4.0, usually 3.2—3.8, in head, and 3.2—4.1, but usually 3.5—4.0, in greatest
depth of body.
Ficure 86. Brevoortia tyrannus, 320mm TL, 247mm SL, Chesapeake Bay, USNM 129809. Drawn by
Ann S. Green.
Sca.es adherent, the exposed part notably deeper than long, the scale itself some-
what deeper than long (Fig. 85 8); margin irregular in young about 60 mm long;
short blunt serrae on margin in examples of about 100 mm, the serrae increasing in
length with age, becoming slender hair-like appendages, often extending nearly across
the exposed portion of the next scale in large specimens; scales greatly reduced in size
on upper part of side, on back, and at base of caudal; in rather regular series on lower
half or so of side; most frequently 3 or 4 vertical rows of scales exposed between tip
of pectoral and base of pelvic. A row of large modified scales on each side of median
line of back in front of dorsal fin, these scales not fully modified in specimens under
about 125 mm TL, rather variable in number, most frequently 34—38 in each series.
VENTRAL scuTEs rather strong, 17—22, usually 18-21, in front of pelvic fins; 10-15,
usually 12-14, behind these fins.
Heap 2.8-3.5, usually 2.9-3.2, in SL, its depth 3.1-4.0, usually 3.3—-3.8.
Snout only moderately blunt, with a prominent median notch in adults, the length
3-9-5.5, usually 4.0-4.7, in head. Eye difficult to measure accurately in adults
because of much adipose tissue, about 4.4—6.2. INTERORBITAL 4.2—5.3, usually 4.4—
5.0. CHEEK deeper than long except in young. Maxitrary rounded, generally scarcely
reaching below posterior rim of eye, the length 1.9—2.7, usually 2.1-2.5, in head.
Manors .e included in upper jaw, its upper margin (within mouth) nearly straight,
Fishes of the Western North Atlantic 34.9
its length to joint 1.7—2.0 in head. Upper section of OpercLe with prominent radiating
ridges. GILL RAKERS very numerous; those on upper limb extending down and over
those on lower limb, the longest about as long as snout in adults; increasing in number
with age and growth, the greatest increase occurring before a 100-mm length is at-
tained: about 60 on lower limb in 60-mm specimens, about 100 in 100-mm fish, about
140 in 200—250-mm examples, and 150-160 in large adults 330-360 mm long.
Trrtu absent except for minute ones on margin of maxillary in young about 60 mm
and less.
Dorsat fin moderately elevated anteriorly, its margin definitely concave, its longest
rays as long as snout and fully half of eye, the last ray somewhat longer than the pre-
ceding ones; origin of fin generally slightly nearer to base of caudal than to margin of
Ficure 87. Brevoortia tyrannus, modified scales in front of dorsal fin, from same specimen shown in Fig. 86.
Drawn by Ann S. Green.
snout; a low sheath on base of fin, composed for the most part of a single series of
scales not extending above the basal third of shortest rays when standing erect. CauDaL
rather deeply forked, the middle rays about as long as eye; the lobes rather short, the
lower one the longer, about as long as head, 2.8—4.0, usually 3.0-3.8, in SL. Anat
low, little elevated anteriorly, the margin nearly straight; its origin under or somewhat
behind vertical from tip of last dorsal ray, its base 5.2—7.0, usually 5.5—6.6, in SL;
a very narrow sheath of scales at base. Prtvic fin small, with a very gently convex
margin, the outermost ray only a little longer than the innermost (Fig. 84 8), the fin in-
serted slightly behind vertical from origin of dorsal, the length 3.1—3.8 in head. Prc-
ToRAL fin slightly falcate, its length 5.0-5.9 in SL in adults, 1.6-1.9 in head, the
longest ray about four times the length of the shortest one, the fin generally failing to
reach base of pelvic in adults by a distance somewhat greater than diameter of pupil.
AXILLARY APPENDAGE of pectoral variable in length, about 75 °/) of the length of fin
in large examples, only about half of length of fin in examples about 100 mm long,
and little developed in young of 50 mm.
Color. In fresh specimens, back dark green to bluish. Sides generally brassy and
sides of head bright silvery to slightly brassy. A large black spot at shoulder a very
short distance behind margin of opercle and well above middle of side, variable in size
and shape among specimens, generally roundish but sometimes vertically elongate, and
about as large as pupil. This spot develops first and is generally evident when the
fish reach a length of about 75 mm (3 in.). The shoulder spot usually followed among
half-grown and large examples, from about 150 mm (6 in.) upward, by a variable
350 Memor Sears Foundation for Marine Research
Ficure 88. Brevoortia tyrannus. a, newly laid egg in advanced stage of cleavage; B, egg with advanced stage
of embryo; c, newly hatched larva, 4.5 mm long. After Kuntz and Radcliffe.
number of smaller dark spots, sometimes only a few small ones and sometimes many,
differing greatly in number, size, and intensity of darkness, these spots sometimes
arranged in two, three, or more indefinite horizontal series on anterior part of side,
seldom extending far beyond vertical from origin of anal; a profusely spotted example
illustrated in Fig. 86. Fins pale yellow to brassy, sometimes with dusky punctulations,;
caudal generally dusky at base and often with a dusky margin. Peritoneum black.
Variation in color, especially in the dark spots on the side, is very great.
Size. The usual size of adult fish at Beaufort, North Carolina, is about 300—
350 mm (12-14 in.) TL. It is generally accepted that the fish grow rather larger in the
northern part of the range than in the southern part. The largest specimen in the U. S.
National Museum, from Long Branch, New Jersey, is 430 mm (17 in.) long, but a
larger one, 480 mm (18 in.), has been reported as taken at Woods Hole, Massa-
chusetts (120: 91), and one of 500 mm (20 in.) was reported to the writer by Dr.
H. F. Prytherch verbally. The last mentioned specimen was taken at Beaufort, North
Carolina, and weighed 3.5 pounds.
Fishes of the Western North Atlantic Rea
Development and Growth. The eggs and larvae have been described by Kuntz and
Radcliffe from material collected at Woods Hole, Massachusetts (76: 119). The eggs
are buoyant and spherical, have a diameter of 1.4—1.6 mm, and are highly transparent;
they contain a small oil globule, and are covered with a thin horny membrane; the
perivitelline space is broad. Incubation occupied “not more than 48 hours.”
The newly hatched fish are “relatively slender” and about 4.5 mm long. The
vent, as usual in clupeoid fishes, is “less than one-fifth the length of the body from the
posterior end.” At four days of age the larvae were 5.7 mm long, and black chromato-
phores were present in a series along the entire digestive tract. At 9.0 mm the dorsal
and caudal fins were at least partly developed, and posteriorly the intestine was convo-
luted. At 23 mm all the fins had become well differentiated, and the body remained
“relatively slender”; black chromatophores were present on the nape, along the margin
of the opercle, near the base of the dorsal, anal, and caudal fins, and in groups posterior
to the dorsal fin and ventral to the pectoral fin; internal black areas could be seen
along the dorsal wall of the abdominal cavity, and a series of dark spots was present
above the notochord.
Compared to the Woods Hole larvae, specimens from Beaufort, North Carolina,
of comparable size agree with them fairly well in development, but in each phase they
are notably more slender. Beaufort examples 9.0 mm long are little compressed and
scarcely thicker than a number-60 sewing thread; the depth is contained about
30 times in TL. Specimens 18 mm long are fully as well developed as those of 23 mm
described and illustrated by Kuntz and Radcliffe, and the 23-mm examples are rather
more advanced than the northern ones of equal length; the body continues to be more
slender; the depth is contained about 21 times in TL. In 25-mm preserved examples
from Beaufort, the convoluted intestine noted in younger fish is scarcely discernible,
and the vent is now situated slightly behind a vertical from the base of the last dorsal
ray, about as in the adult. Some examples, at a length of 30 mm, or even up to 35 mm,
remain very slender, their depth being contained about 13 times in TL; and in these
slender examples the ventral surface of the chest and abdomen remains round to slightly
flat, with indications of a row of bony points along each lateral edge. Juvenile pigment
spots remain about as described for examples of 23 mm or so, but other specimens of
the same length are much more compressed and notably deeper; their greatest depth
is contained only about 7.7 times in TL, or only about 6.7 in SL. In these deeper
and older fish, the two series of lateral bony points noted in the slender fish have
become coalesced to form definite ventral scutes on the median line of the chest and ab-
domen; also, the fish are rather fully pigmented, the sides of the head are bright silvery,
a silvery lateral band is evident, and a sheen of the same color is present on the sides of
the abdomen. It seems, then, that when the fish reach a length of about 30 mm the
increase in length is retarded during metamorphosis from the somewhat roundish
slender larvae to the deeper and more compressed young adults.
Schools of young Atlantic Menhaden were observed repeatedly at Beaufort in
winter and spring, and specimens 24-35 mm, caught from a school on April 23,
Bie2 Memoir Sears Foundation for Marine Research
eee el ae el ee ae
PO i ae ta Ree
: = pecomnnenst aah
ee eS “ 2
EUS aa Sal a Foie os eee ey 3
AES 7 sey Pye ye
ae’
Ficure 89. Brevoortia tyranaus. a, larva four days after hatching, 5.7 mm; B, young, 9 mm; c, 23 mm; D, 33
mm. a, B, c drawn by Albert Kuntz, p by E. B. Decker.
1926, differed little in development; all had remained slender and were more or less in
the larval stage. As seen in the water they were almost transparent, but when removed
from the water, dark pigment spots were evident along the middle of the side in a more
or less definite lateral band.
Specimens 40-45 mm long are shaped very much like large fish, although they
remain rather more slender; their greatest depth of body is contained about 3.6—4.0 in
Fishes of the Western North Atlantic 253
SL. The obtuse angle in the lower limb of the first gill arch is evident, and the gill rakers
on the upper limb of this arch now definitely extend down and over the upper ones on
the lower limb. Striations on the upper plate of the gill covers are evident, but the cheek
remains about as long as deep. Scalation is about complete. The silvery lateral band of
smaller specimens has blended with the silvery abdomen; the dark shoulder spot, always
present in adults, is evident though small; and in some specimens suggestions of sup-
plementary spots are already present.
The gill rakers increase in number and in proportionate length with age (see
Description). Therefore, if their number is to be usable at all for differentiating species,
specimens of equal size must be compared.
In the Gulf of Maine, the fish hatched during the summer reach a length of
60-80 mm during their first winter and average slightly more than 160mm the second
winter, and those hatched in the fall are 30 mm long the first winter and 130 mm the
second winter, with every gradation in size between the two (26: 122).
In Chesapeake Bay, where the fish apparently spawn only during the autumn,
the young are only about 27 mm long the following January and 46 mm the following
April (59: 103). These data, though not complete, are in general agreement with more
extensive data collected by me at Beaufort, where spawning quite certainly occurs only
during late fall and early winter. Seventy-nine larvae, collected there from the last half
of November to and including the first half of March, were between 4.0—20 mm, and
the average increase in length in each month was about 12 mm, excluding December,
when only three larvae were taken (5.0, 5.0, and 7.0mm). Thereafter, more rapid
growth was indicated: seven specimens taken the last half of March were between
22-31 mm and averaged 27 mm; 110 collected in April were 30-40 mm and averaged
about 35 mm; $0 collected during May were 30-41 mm and averaged about 35 mm;
and 48 examples taken in June ranged between 38-56 mm and had an average
length of 45 mm. Lengths of 85-150 mm (3.4—6.0 in.) were attained during their
second winter; the average was about 110 mm (4.4 in.); measurements were based
on 167 fresh examples.
“Sexual maturity is apparently attained in the season following the third winter,
and a few of the older fish Welsh examined showed as many as 9-10 winter rings on
their scales” (Bigelow and Schroeder, 15: 116).
Spawning. The Atlantic Menhaden spawn at sea in saline waters. Many large
fat fish with well-developed roe were observed by me off Beaufort, North Carolina, during
autumn only. Such fish were seen also in the fall in the Chesapeake Bay area (59:
103) and in December off Fernandina, Florida (from unpublished notes by William
W. Welsh). However, off New England, spawning takes place in late spring and through-
out the summer, the height of the spawning season occurring in June; and in 1915
off Woods Hole, Massachusetts, eggs were taken as late as the last week of October
(76: 119).
At Beaufort, spawning apparently takes place only off Beaufort Inlet, for all of
the small ones, 16 mm and less, were taken at sea. This would be expected, for the large
23
354 Memoir Sears Foundation for Marine Research
mature fish in that vicinity seldom enter the harbor or other inshore waters. Northward,
however, as at Woods Hole, Massachusetts, the eggs and larvae were collected in the
harbor (76: 119). The place and season of spawning, then, seem to vary according to
locality.
Habits. Both young and adults, acccording to Bigelow and Schroeder, feed and
travel in compact schools at the surface, swimming close together side by side and
tier above tier (15: I14):
In calm weather they often come to the surface when their identity can be recognized by the ripple they
make, for pogies, like herring, make a much more compact disturbance than mackerel do... Also, pogies as
they feed frequently lift their snouts out of water, which we have never seen herring do, while they break the
water with their dorsal fins, also with their tails. And the brassy hue of their sides catches the eye.
The larvae appear to be more or less solitary, for only one specimen or at most a
few were generally taken in a haul, off Beaufort. They do not stay at or near the surface
as strictly as older ones do, for most of the larvae at hand were taken in nets towed
along the bottom. However, schooling and surface swimming begin early in life, for
many schools of young 25-35 mm long (some just reaching the adult stage) were
observed many times on windy days near shore on the leeward side of Pivers Island
at Beaufort.
Migrations. These fish are present principally in the Gulf of Maine from July to
September, off Woods Hole, Massachusetts, from June to September, and off the coasts
of New York, New Jersey, and Delaware from May to September, being most numer-
ous during midsummer; in Chesapeake Bay small to medium-sized fish are numerous
throughout the summer, and large ones are taken off the mouth of the Bay (or off the
Capes) during October and November; in North Carolina, small and medium-sized
fish are present all summer, the large ones from October to December; and on the
east coast of Florida the fish are numerous from early spring to late fall.
Small to medium-sized specimens are present in Chesapeake Bay throughout the
year but in greatly reduced numbers during the winter. Adults off Beaufort, though
most numerous during late fall, sometimes remain in the nearby offshore waters until
January if the winter is mild. Schools composed of fish that are generally 150 mm
(6 in.) and less are present in Beaufort Harbor and connecting estuaries all winter,
and schools of young, generally under 50 mm in length, ascend freshwater creeks or
enter small drainage ditches in the spring. Unlike adults in northern waters that come
close inshore, into the bays and harbors (16: 122), large fish in North Carolina very
rarely cross Beaufort Inlet to enter the Harbor and adjacent waters.
It is still much of a mystery where the schools of adults go after the fish leave
the waters within 10-15 miles of shore, the usual operating range of the fishing
schooners. Sufficient taxonomic work has been done, however, to indicate that each
section of the coast, that is the New England, Middle Atlantic, and South Atlantic
states, has a more or less distinct population. Therefore, if a north-south migration
does take place, then the fish from each section return to the general vicinity from
which they migrated originally. It seems more probable now that the principal migra-
Fishes of the Western North Atlantic 355
tion is offshore and inshore.?? No doubt temperature is an important factor in the
movements of the fish; it certainly affects the supply of plankton upon which they
feed. In years when inshore runs fail, it is even probable that the fish remain so far
offshore that they are beyond the usual range of the fishing vessels. In slight con-
firmation of this theory, I recall that one autumn, when the usual inshore runs failed
at Beaufort, a progressive operator, upon my suggestion, induced one of his boat
captains to go farther offshore than usual in search of a catch. Fish, indeed, were
found, at a considerable distance offshore, apparently too far off for practical fishing
with the crews and schooners available.
Food. Menhaden feed on small organisms strained from the water by their numer-
ous long, slender, close-set gill rakers, which form an effective strainer. While feeding,
the fish generally swim near the surface and often “break water’’; they whirl around,
sound a short distance, come out of the whirl, and swim up and straight ahead at a
considerable speed for a rather short distance. During this time the mouth is wide
open and the gill covers are lifted, thus making it possible for a fish to filter a great
amount of water with minimum effort. The food that is ingested depends in large
measure upon the organisms that are present where the fish are feeding. Even a
considerable amount of mud and general debris is often swallowed. Included in the
stomach contents examined by various investigators were: numerous small crustaceans,
especially copepods; small annelid worms; rotifers; and unicellular plants, particularly
diatoms and peridinians. The plant organisms, as a rule, constitute the chief food.
This species has a very thick-walled stomach, and a long intestine, as is usual among
fish that feed on vegetable matter. Among 64 fish taken at Beaufort, ranging between
65-214 mm, the intestinal tract increased more or less proportionately with the length
of the fish; generally it remained between 400 and 450°/, of the standard length,
according to A. Bascom Hardcastle (unpublished data).
Enemies. Most predatory animals associated with the sea feed on Atlantic Men-
haden—an easy prey because of their habit of schooling. Their fiercest enemy prob-
ably is the bluefish (Pomatomus), which, it is said, kills many more than it eats. Among
the other fish that feed on them extensively are the cod, pollock, hakes, weakfish,
swordfish, tuna, dolphin, amberjacks, and sharks. Whales and porpoises, as well as
birds, also devour many of them.
Parasites. Atlantic Menhaden so often have a large crustacean parasite in the
mouth that they are sometimes referred to as “‘bugfish.” It is said that in some schools
nearly every fish has a parasite in its mouth. This “bug,” an isopod listed as Olencira
praegustator (Latrobe) (107: 231), has a robust body, generally fully half an inch long
and occasionally approaching an inch in length. Sometimes the mouth of the fish is
deformed because of its presence; thus the amount of water screened for food must be
considerably reduced in fish so infected.
37- According to Rose (zro: 104, 105), the recent investigations by the U. S. Fish and Wildlife Service make it likely
that the north-south migrations of the Menhaden are more regular and extensive than Dr. Hildebrand supposed.
—H. B.B.
230
356 Memoir Sears Foundation for Marine Research
Another large parasite, the copepod Lernaeenicus radiatus (LeSueur), buries its
horns in the flesh on various parts of the fish’s body, most commonly along the back
(137: 480). This parasite sometimes approaches a length of two inches and has a pre-
dominant reddish color, which makes it very conspicuous. Other copepods listed
by Wilson as parasitic on B. tyrannus are: Bomolochus teres Wilson, Caligus schistonyx
Wilson, Caligus chelifer Wilson, Pandarus sinuatus Say, Lernanthropus brevoortiae Rathbun,
and Clavellisa spinosa Wilson.
The following have been listed by Linton (85: 440): Scolex polymorphus Dujardin,
a cestode, small and free in the intestine; cysts and blastocysts of Syxbothrium on the
viscera; and the trematodes Distomum appendiculatum Rudolphi and D. vitellosum Linton;
later he added Dacty/ocotyle sp., and Distomum pyriforme Linton (86: 352).
Hardcastle has reported a sporozoan parasite of the genus Eimeria for the testes
of specimens taken at Beaufort; he stated that a “study of over a hundred individuals
[Menhaden] showed an infection of nearly 50 per cent” (54: 214).
Diseases. No report of a specific disease has been found in the literature, but
there are records of many fish having become stranded along the shore. It has been
suggested that their enemies at times drive them ashore, but my observation is that the
fish make no special effort to flee from enemies. In fact, when caught in a net they
make little effort to escape. It seems more probable, therefore, that the stranded fish
died of a disease. An epidemic has indeed been reported (123: 742).
Variations. Large fish generally are proportionately deeper than small ones, and
those from the southern part of the range are proportionately deeper than northern
ones. However, the variation in depth among fish from any one general region is great
enough to obscure any definite distinctions. Southern fish have a somewhat larger head
and longer fins (especially the pectoral) than those from the northern part of the range,
but the gap is fully bridged by examples from intermediate areas. Contrary to expecta-
tion, data at hand show that the vertebrae do not average fewer in southern specimens.
But further detailed studies are needed to determine the exact characteristics of popula-
tions inhabiting the different general sections of the coast; even now specimens from
the Gulf of Maine are too few in the collections at hand to determine their finer charac-
teristics.
As shown in the Description, the number of gill rakers increases greatly with age,
with the principal increase occurring before the fish reach a length of about loo mm
(4 in.). Also, the scales in small specimens have only irregular margins while those in
larger fish are pectinate. In using scales to differentiate species (their development in
all species being similar), it is necessary to compare scales from examples of about equal
size; and it is preferable to use large specimens, for in them the differences are most
pronounced.
See Co/or for variations in color.
Commercial Importance. These fish, though exceedingly valuable, are not used very
extensively as food by man, mainly because of their bony nature and oiliness. How-
ever, some find the flesh delicious, and those living along the coast, especially the
Fishes of the Western North Atlantic 2157
fishermen, eat them in season as a common article of diet. Considerable quantities are
often ‘“‘corned”’ (salted) for home use during winter, and the writer knows from first-
hand experience that these fish are delicious when smoked. They were canned to a
limited extent for export during the last war, and a small quantity is still canned for
home consumption.
Because of their abundance, large numbers are available for reduction to oil, fish
meal, and fish scrap; the Atlantic Menhaden fishery is one of the most valuable of
our fish industries. The catch for 1945 amounted to 759,073,820 pounds, valued at
$ 11,202,127 (Canned Fish and Byproducts, U.S. Fish Wildl. Serv., 1945: 15); of
this amount, 120,493,920 pounds were credited to Florida, 142,209,510 to North
Carolina, 77,232,910 to Virginia, 280,463,340 to Delaware, and 138,584,140 to New
Jersey. For 1950 through 1953, the approximate catches have been as follows: between
9,304,000 and 39,900,000 pounds for New England, mostly taken south and west
of Cape Cod; between 372,950,000 and 857,584,000 pounds for the Middle Atlantic
states; 92,374,000 for Chesapeake Bay; and between 147,125,000 and 314,841,000
pounds for the South Atlantic states. In 1953 the total catch was 1,259,031,000
pounds.
Because of the great fluctuations in catch from year to year, the catch for a single
year, or even for many successive years, may not indicate their actual abundance.
Furthermore, since the fish are not equally fat every year, or at all times when
available, equal quantities of fish (pounds) do not always yield equal quantities of oil
(gallons), the most valuable product obtained from these fish. Thus the value of a given
catch depends to a large extent upon the amount of oil it yields. In general, large fish
yield more oil than small ones. For example, at Beaufort, North Carolina, the summer
“runs,” consisting of small to medium-sized fish, yield so little oil that they are con-
sidered scarcely worth handling, but the fall runs, consisting of large and fat gravid
fish, make the industry profitable.
In discussing this subject, Greer (48: 24)%* wrote as follows in 1915:
The yield of oil varies greatly on different parts of the coast, and also from year to year and at different
seasons. The fish are always larger and fatter during the fall than at any other season, consequently there is a
greater yield of oil during that period. The fish taken from northern waters produce more than those taken from
southern waters; the writer visited one of the northern factories during August 1913, and was told that a short
time previous to his visit they had made 20 gallons, and that the average yield for July had been 16.5 gallons
per 1,000 fish.8® The maximum for the Middle Atlantic States is about 15 gallons, though it usually is less;
in the early part of the season it is very much less than that amount. The South Atlantic fish rarely produce more
than 8 gallons per 1,000 fish, and frequently it is less than 1 gallon during the early spring fishing. The average
number of gallons of oil made from 1,000 fish during the season of 1912 was as follows: Connecticut 11.73;
New York 11.36; New Jersey 5.08; Delaware 7.72; Virginia 3.72; and North Carolina 1.98.
38. This paper, entitled “The Menhaden Industry on the Atlantic Coast,”’ although written before the manufacture
of fish meal for stock and poultry feed, contains much valuable information concerning the history of the industry,
the construction of factories, the machinery used in these reduction plants, the vessels employed, the methods of
fishing, their reduction, men employed, etc.
39- The fish generally are measured, and the measure used is said to contain 1,000 fish, no account being taken of the
size of fish. Thus, the measure arbitrarily contains 1,000 fish regardless of whether they are 6 or 12 inches long.
In conyerting the number of fish to pounds, the factor 0.67 is used in the statistical division of the U. S. Fish and
Wildlife Service. That is, the number of fish is multiplied by 0.67.
358 Memoir Sears Foundation for Marine Research
As further evidence of the great value of this extensive fishery, additional data
from Greer show that the total catch for 1912, a near banner year, was 1,061,483,750
fish (711,435,312 pounds); this catch yielded 6,651,203 gallons of oil valued at
$ 1,551,990 and 88,520 tons of scrap worth $ 2,138,165; 48 factories, valued at
$ 3,625,983, were in operation; the vessels engaged in the fishery numbered 147,
valued at $ 3,456,792; and there were 2,159 employees in the factories and 3,735
fishermen, drawing a combined wage of $ 1,579,984.
The value of the fish scrap, now that most of it is used as fish meal for poultry
and livestock feed instead of as fertilizer, is considerably greater than it was; this change
in turn produced other changes, for this processing requires quicker and more sanitary
handling of the fish, including cooking before decomposition sets in; some changes at
the factories were also required, for apparatus had to be installed for drying the residue
quickly after the oil had been extracted.
Fluctuations in Abundance. No species has been more erratic in its appearances along
the coast than B. tyrannus. For example, in 1911 and in 1912 large catches were made
at Beaufort, the center of this fluctuating industry in North Carolina, but then these
near banner years were followed by several exceptionally lean years, which forced
some of the local operators into bankruptcy. However, in 1918 the fish were back
again. Even though the annual catches (without adjustment for variations in fishing
effort) have fluctuated since that time from 54,476,000 pounds in 1932 to 181,968,000
pounds in 1939, there has been no complete failure in any year since the period
1913-1916.
Although at least some of the reasons for the fluctuations in abundance of certain
species of fish are known, no explanations have been given for the fluctuations in At-
lantic Menhaden. For the Gulf of Maine, Bigelow and others have stated that these fluc-
tuations date back to the early history of the fishery and have continued since then (15:
116, 1173; 16: 120). Thus, in 1845 they were abundant and in 1847 scarce. Then, for
some years prior to 1875, they were tremendously abundant off the coasts of Massa-
chusetts and Maine every summer, and a considerable fishery for them grew up on the
Maine coast. However, during the cold summer of 1877 few were taken in the Gulf un-
til September and October, when they were reported as “‘about as abundant as normal.”
So few of these fish appeared north of Cape Cod from 1879 through 1885 that the cap-
ture of one caused comment, and many people thought that the fish would never return.
However, in 1886 the fish reappeared and in 1888 they were so abundant that the
fishery was revived. The period of abundance was short, however, for in 1891 the
catch was less than half of that taken the year before, and in 1892 few were seen north
of Cape Cod. Similar periods of scarcity and abundance followed each other until 1904,
when a 20-year period of scarcity began. Then in 1922 the fish returned in such abund-
ance that 18 steamers fished successfully in Massachusetts Bay; further, “upward of
1,500,000 pounds were landed by the larger fishing vessels besides what the small
boats brought in.” From the middle 1920’s until the middle 1940’s there were not
enough of them in the Gulf of Maine to be of any commercial importance there.
Fishes of the Western North Atlantic 359
Although they were plentiful again in the Gulf from 1946 through 1949, fewer were
present from 1950 through 1956.
Comparison of statistics for the past 20 years or so shows no correlation as to the
quantity taken along different sections of the coast. If North Carolina, for example, has
a good run in a given year, it does not follow that Florida, New York, or New Jersey
will have a good run that same year. For example, in 1929 the fishery in North Caro-
lina yielded 173,490,000 pounds of Atlantic Menhaden—considered an unusually large
catch. But in that same year the catch credited to Florida was rather less than average,
50,532,000 pounds, and the combined catch for New York and New Jersey was a
near failure with only 11,092,000 pounds. Again in 1937, when the catch in North
Carolina amounted to only 61,706,000 pounds (next to the smallest during about 20
years), the catch for Florida was 139,788,000 pounds and the combined catch for New
York and New Jersey was 99,684,000 pounds, both well above the average for the
20-year period.
It seems improbable that the commercial catches taken by the fisheries in the
western Atlantic have made any serious inroads on Atlantic Menhaden populations in
any general area. If it is true that the handiwork of man has had no important effect
on the abundance of this species, then the fluctuations must be attributed to natural
phenomena that occur in their enviroment.
Successful and unsuccessful spawning seasons are known to profoundly affect the
abundance of some species and this is probably so in the case of this species also.
However, it does not explain why a fish that may attain an age of 9-10 years
(as concluded from the number of winter rings on the scales) may be abundant one
year in a certain locality but very scarce or even absent there the next year. Furthermore,
since the fish caught during a fishing season in any one area are not all of the same
age and size, the failure of one or even several successive spawning seasons would not
cause an abrupt diminution from one year to the next. The theory that these fish follow
their food and therefore remain in water having an abundant growth of plankton of
suitable size has been advanced. It would follow then that if sufficient food is not
available near the shore, the fish will remain offshore, perhaps beyond the range
of the vessels employed in the fishery. This theory seems to be the most plausible one
advanced to date. It has been suggested also that the temperature of the water is a
factor, which no doubt is true, particularly insofar as it affects the growth of the plank-
tonic food required by these fish. It is obvious that further study is needed to determine
more specifically the reason or reasons for the exceedingly great fluctuations in the runs
of Atlantic Menhaden.
Methods of Fishing. These fish are caught chiefly with purse seines, though smaller
quantities are taken with long haul seines and in pound nets; and minor quantities are
caught with ordinary seines or drag nets and with gill nets.
The vessels employed in the operation of this large industry are a very familiar
sight to those living near the fishing centers, and in some localities, for example
Beaufort and Moorehead City, North Carolina, the economic life of the communities
360 Memoir Sears Foundation for Marine Research
is so dependent upon the fishery that nearly everyone watches the incoming vessels to
see what kind of a catch has been made. If the catch is good, the operator’s flag is flown
from the “‘crow’s nest” (see below), thus notifying the factory to get up steam and for
hands to stand by to take care of the catch. It is a common saying among people of
these communities that the stench from the factories “smells like bread and butter”
to them.
Formerly, sailing vessels were used in this fishery. These were superseded by
auxiliary schooners, which in turn were replaced by power-driven vessels equipped
with diesel engines. All of those used in North Carolina and southward are of nearly
the same design and are capable of carrying 400,000 fish or more. They are ruggedly
built wooden vessels, most of them less than 100 feet long, with a broad beam and a
high bow to make them seaworthy. A tall mast is set in the forepart of the vessel, and
near the top of it is a small platform, generally surrounded by a rail and often partly
enclosed by canvas. This is the crow’s nest from which one or two men watch for
schools of fish.
Each vessel carries two ‘‘purse seine boats’’ which, when not in use, are suspended
from derricks on each side of the mother ship. When a school of fish is sighted, the vessel
approaches it, the purse seine boats with seine aboard are lowered, and the school of
fish, or at least part of it, is surrounded as quickly as possible with the seine. Next the
seine is ‘‘pursed” by drawing in a line (at the bottom of the net) that runs through
rings. After the fish have been concentrated into a more or less compact mass they are
loaded aboard the vessel with a large dip net, generally operated by means of block
and tackle.
Steamers are also used in pursuing these fish, though less so than formerly.
They are larger than the vessels driven by diesel engine, hence they can go farther and
stay out longer. This was advantageous in former days when the scrap was used only
as a fertilizer, but now that it is used mostly as poultry and stock feed and since the fish
preferably should be processed on the day they are caught, the advantage of these
larger vessels has diminished.
General Range. Continental waters of North America, from Nova Scotia to Florida;
taken commercially from eastern Maine to northern Florida, as far south as Mayport.
Stragglers have been taken as far northeastward as St. John Harbor, New Brunswick
(64: 11), and St. Mary Bay, Nova Scotia (16: 120), and as far south as Indian River
City, Florida (33: 56); and large schools have been reported as seen during the winter
off Cape Canaveral and Mosquito Inlet (43: 36). This species has often been reported
for the coasts of the Gulf of Mexico, Brazil, Uruguay, and Argentina, but these reports
are for another species (p. 344; 58: 1-39). Fowler reported it for the “Senegal region”’
of Africa but had no specimens from there (39: 174); also, there are earlier reports
of a Brevoortia from Africa; whether it is tyrannus needs verification.
Details of Occurrence. As discussed under Fluctuations in Abundance (p. 358), it
is well known that Atlantic Menhaden fluctuate widely from year to year and over
periods of years. Prior to 1850 they were reported as plentiful at the mouth of the
Fishes of the Western North Atlantic 361
Bay of Fundy on the New Brunswick side; though they have not frequented these
waters subsequently, except for occasional strays (see above), they have been plentiful
off and on westward. In years when these fish were plentiful, their chief centers of
abundance north of Cape Cod have been the Massachusetts Bay region, the region
of Casco Bay, and among the islands thence eastward to Penobscot Bay, where they may
congregate as much as 40-50 miles out from the land. But they have never been
reported for the central part of the Gulf or for the off-lying fishing banks.
Although the Atlantic Menhaden’s appearances and disappearances are unpredict-
able for any given locality, they are far more regularly and uniformly numerous south-
ward from Cape Cod than northward, not only along the open coast but within the larger
bays and sounds as well. Landings southward from New York for 1954, the most recent
year for which pertinent catch statistics are at hand, were as follows: 136 million
pounds for New York and New Jersey, 306 million for Delaware, 160 million for North
Carolina, 42 million for northeastern Florida, but only four million pounds for Mary-
land and Virginia combined, and about the same amount for South Carolina. While these
data might suggest centers of greatest abundance at some areas along the American
coast compared to others, it seems more likely that they reflect not only the economic
laws of supply and demand, but the wide and varied patterns of distribution that have
been discussed under Fluctuations in Abundance (p. 358).
Synonyms and References:
Clupea tyrannus Latrobe, Trans. Amer. philos. Soc., 5, 1802: 77, pl. 1 (drawn without dorsal; no descr.; type
local. Chesapeake Bay; notes probably applicable to some species of Pomolobus; an isopod, O/encira
praegustator, from mouth descr.).
Clupea menhaden Mitchill, Rep. in part on Fishes of New York, 1814: 21 (orig. descr.; type local. presumably
New York); Mitchill, Trans. Lit. philos. Soc. N. Y., Z, 1815: 453 (descr.; New York; use as fertilizer);
Giinther, Cat. Fish. Brit. Mus., 7, 1868: 436 (synon., descr.).
Clupea neglecta Rafinesque, Amer. Month. Mag., 2 (3), 1818: 206 (orig. descr.; type local. Long Island, New
York).
Alosa menhaden De Kay, New York Fauna, Pt. 4: Fishes, 1842: 259, pl. 21, fig. 60 (descr., import., New York);
Storer, Fishes Mass., 1867: 158, pl. 26, fig. 4 (descr., abund., how caught, uses, Massachusetts Bay).
Alosa sadina De Kay (not of Mitchill), New York Fauna, Pt. 4: Fishes, 1842: 263, pl. 40, fig. 129 (descr.,
fig. clearly shows B. tyrannus).
Clupea carolinensis Gronow in Gray, Cat. Fish. Brit. Mus., 1854: 140 (orig. descr.; type local. S. Carolina).
Brevoortia menhaden Gill, Rep. U.S. Comm. Fish. (1871-1872), 1873: 811 (how and where used, range);
Uhler and Lugger, Rept. Comm. Fish. Md., ed. 1, 1876: 156; ed. 2, 1876: 133 (descr., synon., use,
Maryland).
Brevoortia tyrannus Goode, Proc. U.S. nat. Mus., r, 1879: 5 (establ. validity of Latrobe’s specific name,
tyrannus), 31 (descr.; cf. “var. aurea” and B. patronus; varieties menhaden and aurea reognized;
new variety 4revicaudata named and defined); Goode, Rep. U.S. Comm. Fish. (1877), 1879: ii—xu,
1-529, pls. 1-31 (complete hist. of American menhaden; discus. species and varieties, includ. S. American
ones; full descr. of industry); Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1882: 269 (descr., synon.);
Good iz Goode, et a/., Fish. Fish. Industr. U.S., 1 (3), 1884: 569, pl. 205 (common names, migr.,
distr., food, reprod., enemies, uses); Goode, Amer. Fishes, 1888: 385, fig. (migr., food, etc.); Smith, Bull.
U.S. Fish Comm. (1890), To, 1892: 64 (behavior in lower Potomac R.); Smith, Bull. U. S. Fish Comm.
(1891), IZ, 1893: 191, 195 (small ones enter brackish and fresh water); Peck, Bull. U.S. Fish Comm.
(1893), Z3, 1894: 113-126, pls. 1-8 (food and feed.); Jordan and Evermann, Bull. U. S. nat. Mus.,
47 (1), 1896: 433, fig. 195 (descr., size, range, synon.); Evermann and Kendall, Rep. U.S. Comm.
362 Memoir Sears Foundation for Marine Research
Fish. (1899), 1900: 56 (refs.); Linton, Bull. U.S. Fish Comm. (1899), 19, 1901: 277, 440 (food,
cestode and trematode parasites); Bean, T.H., 7th Rep. Forest Comm. N. Y. (1901), 1902: 309, fig.
(synon., local names, size, range, commerc. uses, migr.); Bean, IT. H., Bull. N. Y. St. Mus., 60, Zool.
9, 1903: 211 (synon., descr., size, range, uses, New York); Linton, Bull. U.S. Bur. Fish. (1904), 24,
1905: 352 (nematode, trematode, and copepod parasites); Richardson, Bull. U.S. nat. Mus., 54, 1905:
231 (isopod parasite); Fowler, Rep. N. J. St. Mus.(1905), 1906: 103, pl.6 (descr., New Jersey);
Smith, N.C. geol. econ. Sury., 2, 1907: 130, figs. 44, 45 (diagn., range, size, common names, spawn.,
econ. import., North Carolina); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7, 1908: 40 (refs., New
England); Sumner, Osburn, and Cole, Bull. U.S. Bur. Fish. (1911), 32, 1913: 742 (reprod., food,
parasites, epidemic, Woods Hole, Massachusetts); Regan (in part of Latrobe), Ann. Mag. nat. Hist.,
(8) 79, 1917: 301 (B. patronus and B. aurea in synon., diagn., range); Kuntz and Radcliffe, Bull. U.S.
Bur. Fish. (1915-1916), 35, 1918: 119-123, figs. 76-86 (eggs, larval develop.); Hildebrand, Rep. U.S.
Comm. Fish. (1918), Append. 6, 1919: 3, 2 figs., 1 pl. (photos of scales, descr., uses, spawn., food,
Beaufort, North Carolina); Huntsman, Contr. Canad. Biol. (1921), 1922: 11 (St. John Harbor, New
Brunswick); Bigelow and Welsh, Bull. U.S. Bur. Fish. (1924), 40 (1), 1925: 118, fig. 48 (descr., size,
range, breed., food, enemies, commerce. import., Gulf of Maine); Jordan, Fishes, 1925 : 277, fig. 211 (econ.
import., habitat); Fish, Bull. U.S. Bur. Fish (1925), 1926: 165-167, 170, 172 (surface catches, Woods
Hole, Massachusetts); Nichols and Breder, Zoologica, N. Y., 9 (1), 1927: 41, fig. (distr., food, life hist.,
size, New York and s. New England); Hildebrand and Schroeder, Bull. U.S. Bur. Fish. (1927), 43;
1928: 102, figs. 55-59 (rets., descr., var., food, spawn., econ. import., range, Chesapeake Bay); Jordan,
Manual Vert. Anim. NE U.S., ed. 13, 1929: 4 (diagn., range, uses, food); Truitt, Bean, and Fowler,
Bull. Md. Conserv. Dep., 3, 1929: 51, fig. 17 (descr., Maryland); Breder, Field Bk. Mar. Fish. Atl.
Cst., 1929: 69, fig. (range, uses, spawn., habits, “races”); Jordan, Evermann, and Clark, Rep. U.S.
Comm. Fish. (1928), 2, 1930: 44 (range, synon.); Wilson, Bull. U. S. nat. Mus., 158, 1932: 381,
401, 404, 437, 451, 480 and 517 (copepod parasites); Greeley, 26th Rep. N. Y. Conserv. Dep. (1936),
Suppl., 11, 1937: 91 (brackish water Hudson R.; numerous at mouth of Mamaroneck R.); Perlmutter,
28th Rep. N. Y. Conserv. Dep. (1938), Suppl., 15 (2), 1939: 17, tab. 4 and 15 (time of occur., Long
Island; spawn.); Greeley, 28th Rep. N. Y. Conserv. Dep., Suppl., 15 (2), 1939: 82 (young at Long
Island, New York; prey of other fish); Anonymous, Fish. Resourc., U.S. Senate Doc. 51, 1945: 71,
figs. (habits, biol., econ. import.); Hildebrand, Smithson. misc. Coll., 107 (18), 1948: 7, fig. 1 (synon.,
relation., descr., var., range); Bigelow and Schroeder, Fish. Bull. (74) U.S. Fish Wildl. Serv., 53, 1953:
113 (descr., life hist., Gulf of Maine).
Brevoortia tyrannus Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1945: 45, 74, 165 and 266 (refs., occur.).
Menhaden, Greer, Rep. U.S. Comm. Fish. (1914), 1915: 5-27, pls. 1-7 (complete acct. of industry);
Hardcastle, J. E. Mitchell Sci. Soc., 57 (2), 1941: 214 (sporozoan parasite); Anonymous, Canned Fish
and Byproducts, Fish and Wildlife Service, CFS 304, 1945: 15 (statist. for 1945).
Doubtful Reference:
Brevoortia tyrannus Fowler, Bull. Amer. Mus. nat. Hist., 70 (1), 1936: 174 (refs. part African; descr. of Amer.
specimens).
Negative References:
Alosa tyrannus De Kay (not of Latrobe), New York Fauna, Pt. 4, Fishes, 1842: 252, pl. 13, fig. 38 (descr.,
probably Pomolobus aestivalis).
Brevoortia tyrannus Henshall (not of Latrobe), Bull. U.S. Fish Comm., 14, 1895: 211 (Tampa, Florida;
presumably B. patronus); Evermann and Kendall, Proc. U.S. nat. Mus., 3Z, 1906: 74 (synon., lit. on
Brevoortia; cf. specimens from various localities, includ. two from Argentina; presumably B. patronus);
von Jhering, Rev. Industr. Anim., 1 (3), 1930: 229 (N. and S. Amer. representatives compared from
lit., econ. import.); Gunter, Publ. Inst. mar. Sci. Texas, r (1), 1945: 29 (habitat, Texas; this is B. pa-
tronus).
Fishes of the Western North Atlantic 363
Brevoortia brevicaudata Goode 1879
Figure 9o
Study Material. Eight specimens, 165-180 mm TL, 126-147 mm SL, all from
Noank, Connecticut, USNM 14044, 14846. This species, as herein understood, is
known only from the above specimens.
Distinctive Characters. See Relationship (p. 364).
Ficure 90. Brevoortia brevicaudata, 180mm TL, 145 mm SL, Noank, Connecticut, type, USNM 129797.
Drawn by Ann S. Green.
Description. Proportional dimensions in per cent of standard length, and counts,
based on specimens 126-147 mm SL.
Body: depth 35-38. Pectoral fin: length 13.5-16.
Caudal peduncle: depth, 9.6—10.5.
Head: length 29-30. Scales: oblique series along middle of
Snout: length 6.4—7.3. side 47-53.
Eye: diameter 5.4—6. Modified scales: in a series in front of dor-
Interorbital: width 6.2—7.0. sal fin, 35-39.
Maxillary: length 12-13. Ventral scutes: 31-33.
Mandible: length 15.5—-16.5. Fin rays: dorsal 19-22; anal 21 or 22;
Anal fin: length of base 17.5—-19. pectoral 17.
Pelvic fin: length 8.2-8.6. Vertebrae: 48 (1 specimen).
Bopy deep and robust, its greatest thickness about a third of its depth, its greatest
depth near midpoint between tip of pectoral and insertion of pelvic, 2.6—2.85 in SL.
CauDAL PEDUNCLE rather deep, 2.8—3.0 in head or 3.4—3.7 in greatest depth of body.
Scares adherent, much deeper than long, with rather short pectinations as in B.
364 Memoir Sears Foundation for Marine Research
tyrannus of similar size, greatly reduced on back and at base of caudal; § or 6 vertical
rows of scales exposed between tip of pectoral and base of pelvic fin. A row of large
modified scales on each side of median line in advance of dorsal. VENTRAL sCUTES
moderate, 18-20 in advance of pelvic fins and 12-14 behind them.
Heap small, 3.3-3.85 in SL, its depth 3.6-3.8. Snour with a definite median
notch, its length 3.9—-4.6 in head. Eve small, 4.7—5.4. INTERORBITAL 4.1—4.7. CHEEK
notably deeper than long. Maxitzary rather short, reaching about to vertical from
posterior margin of pupil, 2.2-2.4 in head. Manprsze short and included, its upper
margin (within mouth) nearly straight, its length to joint 1.8-1.9 in head. Upper
section of Oprrcte with prominent radiating ridges. GiLL RakERs long, and numer-
ous, the longest ones about as long as snout and half of eye, about 115-125 on
lower limb.
Dorsat fin low, its longest rays scarcely longer than the snout, its origin nearer
to margin of snout than to base of caudal by a distance nearly equal to diameter of
eye; its base with a low sheath composed for the most part of a single series of
scales, not extending above the basal third of the rays. Caupat fin very short, the
middle rays about as long as eye; lower lobe somewhat longer than upper one, more or
less damaged, but shorter than head, about 3.7—4.4 in SL. Anat very low, its origin
under, or a little behind, vertical from tip of last dorsal ray, its base 5.25—5.7 in SL;
a very narrow sheath of scales at base. Petvic fin small, with a gently convex margin,
its outermost ray not much longer than innermost ray; the fin inserted about under
origin of dorsal; the length 3.4—3.6 in head. Pecrorat fin very short, not falcate, its
length 7.25-8.25 in SL, or 1.8-2.2 in head, its longest ray about three times longer
than the shortest one, failing to reach base of pelvic by a distance notably greater than
diameter of eye. Ax1LLary appendage of pectoral short and broad, only a little more
than half of length of fin, 3.0-3.8 in head.
Color. Old preserved specimens dark greenish on back and upper part of side.
Lower parts silvery to brassy. Sides of head silvery. A roundish dark spot at shoulder,
about the size of pupil of eye; only one specimen at hand with suggestions of auxiliary
dark spots on side behind shoulder spot. Fins all uniform brownish.
Relationship. It is strange that no additional specimens like the ones herein de-
scribed, collected in 1874, are among the many in the U.S. National Museum from
the general vicinity of Long Island and northward to Woods Hole, Massachusetts.
These specimens differ in so many respects from the more or less typical B. tyrannus
in the same general vicinity (Noank, Conn.) that it seems necessary to recognize them
as representing a distinct species. The apparent distinctions are shown in a comparison
with specimens of B. tyrannus of equal size from the same general vicinity of Noank
(p. 365). It is evident from this comparison that brevicaudata is not a geographical variant.
The eight specimens in the Study Material are, in part at least, those studied by
Goode, who gave measurements of two specimens from USNM 14846 and an accu-
rate description (44: 33, 37); although he gave them the “‘variety” name drevicaudata,
he did not designate a type. Therefore Goode’s variety has been elevated to specific
Fishes of the Western North Atlantic 365
Comparison of B. brevicaudata with B. tyrannus
(in per cent of standard length)
brevicaudata tyrannus brevicaudata tyrannus
Eleapplengthr rn ececee 29-30.5 30.5-33 Manp1Bze: length..... 15.5-16.5 17.5-18.5
Eyr: diameter .....-:.- 5.5— 6.0 6.2— 6.9 Caupat FIN: lower lobe 22.5-27 30-33
Maxixrary: length .... 12-13 I4-15 PrctoraL FIN: length.. 13.5-16 19.5—20
rank, with the designation of a lectotype approximately 180 mm TL (caudal somewhat
damaged) and 145 mm SL, USNM 129797 (58: 10).
Thus the characters in all respects are smaller in drevicaudata than in tyrannus.
The maxillary in the former reaches about to a vertical from the posterior margin of
the pupil instead of well behind it; the caudal’s lower lobe in brevicaudata, if measured
from the middle of the caudal base, is shorter than the head instead of about equal
to it in length; the pectoral in brevicaudata fails to reach the base of the pelvic by a
distance exceeding the eye’s diameter instead of less than its diameter, and it leaves
5-7 vertical series of scales exposed instead of 3 or 4, as in tyrannus.
If the specimens of drevicaudata are compared with examples from the southern
part of the range of ¢yrannus, as from Fernandina, Florida, the differences are even more
pronounced, for the head and eye in such material are relatively larger than those in
northern specimens of tyrannus, and the fins are still longer.
Nothing distinctive is known about the life history of this species.
Range. Known only from Noank, Connecticut.
Synonyms and References:
Brevoortia tyrannus var. brevicaudata Goode, Proc. U.S. nat. Mus., 2, 1879: 34, 37 (orig. descr.; type local.
Noank, Connecticut; cotypes USNM 14846; cf. “normal” ¢yranmus and aurea).
Brevoortia tyrannus brevicaudata Goode, Rep. U.S. Comm. Fish. (1877), Append. A, 1879: 22 (said to vary
from “normal type” with shorter maxillary and mandible, lower anal, and shorter caudal; Noank, Conn.);
Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 634 (cf. aurea, after Goode); Jordan, Ever-
mann, and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 44 (range, synon.).
Brevoortia brevicaudata Hildebrand, Smithson. misc. Coll., 207 (18), 1948: 10, fig. 2 (synon., descr., range,
lectotype USNM 129797).
Brevoortia patronus Goode 1879
Largescale Menhaden, Gulf Menhaden, Alewife, Shad
Figures 84, 85, 91
Study Material. At least 110 specimens, 60-265 mm TL, 45-207 mm SL, col-
lected at many places along the Gulf coast from Apalachicola, Florida, to Brazos
Santiago, Texas, including the types of B. patronus Goode, USNM 892. Some of the
data used were extracted from unpublished tables prepared by the late William W.
Welsh; also a few lots of juveniles.
Distinctive Characters. See Relationship (p. 370).
366 Memoir Sears Foundation for Marine Research
Description. Proportional dimensions in per cent of standard length, and counts,
based on at least 110 specimens, 45-207 mm SL.
Interorbital: width 6.5—8.0.
Maxillary: length 14-17, usually 15-16.
Mandible: length 17-20, most frequently
18 or 19.
Anal fin: length of base 17-21.
Pelvic fin: length 8.5—10.5.
Body: depth 33-45, usually 35-42.
Caudal peduncle: depth, 8.5-12.
Head: length 31-38, usually 32-35.
Snout: length 6.0-9.0, most frequently
7.0-8.0.
Eye: diameter 7.0—9.2.
WITID =e
RU re
a
y
4
SSN
Ficure 91. Brevoortia patronus, 215 mm TL, 164 mm SL, off Galveston, Texas, USNM 129810. Drawn by
Ann S. Green.
Pectoral fin: length 19-23, most fre-
quently 20-21.
Scales: oblique series along middle of
side 36—50, usually 38-46.
Modified scales: in a series in front of dor-
sal 24-33, generally 25-31.
Ventral scutes: 28-32, most frequently
29-31.
Fin rays: dorsal 17-21, most frequently
19 or 20; anal 20-23; pectoral 14—
17, most frequently 15 or 16.
Vertebrae: usually 45-47, rarely 42, 43,
44, or 48 (117 specimens).
Bopy rather deep, strongly compressed, its greatest thickness about a third of its
depth, the greatest depth well in advance of tip of pectoral and often only about an
eye’s diameter behind margin of opercle, 2.2-3.0, usually 2.4-2.8, in SL; ventral
outline very strongly convex anteriorly, notably more than half of greatest depth of
body below a line extending through lower margin of eye to middle of base of caudal.
CauDAL PEDUNCLE rather slender, 3.0—3.9, usually 3.2—-3.7, in head.
Sca.es adherent; the exposed part much deeper than long, the scale itself some-
what deeper than long (Fig. 85 a), the margin definitely serrate in young of about 60 mm,
the serrae gradually increasing in length with age, becoming definitely pectinate, the
pectinations reaching the full length of the otherwise exposed portion of the succeeding
Fishes of the Western North Atlantic 367
scale in specimens 200 mm TL and upward; scales greatly reduced in size on back
and at base of caudal; rarely more than two rows of scales exposed between tip of pec-
toral and base of pelvic. A row of large modified scales on each side of median line of
back in advance of dorsal fin, these scales not fully modified in examples less than about
100 mm TL, rather variable in number, generally about 25-31 in each series, there-
fore usually fewer than in tyrannus. VENTRAL SCUTES quite strong, more prominent in
young than in adults, 18 or 19, rarely 16 or 17, in advance of pelvic fins, and 12 or
13, occasionally 11, behind them.
Heap rather large, 2.6—3.2, usually 2.9-3.1, in SL, its depth 2.7—3.4, most fre-
quently 2.8-3.2. SnouT blunt, with a prominent median notch, its length 3.9—5.0,
most frequently 4.2-4.6, in head. Eye with comparatively little adipose tissue, 4.1—
5.0, usually 4.3-4.8. InTERORBITAL 4.1—5.0, usually 4.3-4.7. CHEEK much deeper
than long in adults. Maxitiary generally reaching vertical from posterior margin of
pupil, 2.0-2.4, usually 2.1-2.2, in head. Manprs_e included in upper jaw, its upper
margin (within mouth) straight, its length to joint 1.75—1.9 in head. Upper section
of OpeRcLE with strong radiating ridges. GILL RAKERs very numerous, those on upper
limb extending down and over those on lower limb, the longest ones rather longer
than snout; increasing in number with age, about 40-50 in specimens 25-40 mm TL,
80-85 in specimens 45-60 mm, 125-130 in specimens 100-130 mm, and 135-150
in examples 200 mm and upward. TrrTu wanting, except for minute ones on margin
of maxillary in small specimens under about 60 mm.
Dorsat fin rather high anteriorly, its margin definitely concave, its longest rays
nearly as long as snout and eye, the last two rays considerably longer than the preceding
ones, its origin about equidistant between margin of snout and base of caudal; a broad
scaly sheath on the base of fin, almost covering the fin except for the last produced
rays if the fin is deflexed, extending well above basal third of the shortest rays when
standing erect. CaupaL moderately forked, the middle rays exceeding length of eye by
nearly diameter of pupil, the lobes rather long, the lower one often longer than head,
2.$—3.2, most frequently 2.8-3.1, in SL. Anat moderately low, somewhat elevated
anteriorly, its margin definitely concave, its origin under, or a little in advance of, tip
of last dorsal ray, its base 4.75-5.9 in SL; a low scaly sheath at base. Petvic fin
small, with a gently convex margin, its length 3.1-3.7 in head, its outermost ray
only a little longer than the innermost ray (Fig. 84); the fin inserted well behind
vertical from dorsal origin. Pecrorat long, somewhat falcate in adults, its length 4.3 5—
5.25, usually 4.75—5.0, in SL, 1.5-1.9, most frequently 1.5—1.7, in head, its longest
rays about four times longer than the shortest one, sometimes nearly reaching base
of pelvic. AxILLARY sCALE of pectoral variable, generally scarcely more than half of
length of fin in large examples, little developed in young.
Color. Back bluish-gray. Sides brassy-green. Sides of head silvery. A large black
shoulder spot, often followed by a single series (rarely by a double series) of smaller
and less definite spots, these being roundish in some and vertically elongate in others;
the shoulder spot generally appearing in specimens between a length of 50 and 75 mm,
368 Memoir Sears Foundation for Marine Research
the auxiliary spots usually appearing later. Fins mostly plain yellowish green; the
caudal with a dusky margin. Peritoneum jet black.
Size. A small size seems to be normal. An example 265 mm (10.6 in.) TL is
the largest now at hand. Goode (44) quoted Silas Stearns as having seen examples 11,
12, and 13 inches long, but these may have been gunteri.
Development and Growth. The postlarvae, 19-24 mm TL and 16-19 mm SL,
are relatively slender, though deeper than young of tyrannus of the same length, their
greatest depth being about 7.4-10.5 in TL and 4.75—-8.0 in SL. Also, these young
fish are more advanced in development, for even the smallest ones have all fins (ex-
clusive of the pectoral) well enough developed to show rays; however, the pelvic
fin is still well in advance of the dorsal. The convoluted intestine, characteristic of
young clupeoids generally, remains evident in the smaller examples but has become
fully invaginated in the larger ones. The snout seems to be rather more pointed in
the postlarvae than in larger fish, and the large rounded maxillary scarcely reaches
beyond the anterior margin of the pupil. The gill rakers of the upper limb do not
yet extend down and over those of the lower limb, and the filaments on the pos-
terior rim of the gill arches are little developed at this size. The ventral scutes in
front of the pelvics are well developed in the largest specimens, but not those behind
them. The smallest specimens have a series of dark chromatophores along the lateral
edge of the abdomen and another along the anal base; also, few dark chromatophores
are present on the opercle, on the base of caudal, and on the chest; some of these
markings have become obscure in the larger specimens of this lot. Dark specks,
present on the caudal and dorsal fins, are decidedly more numerous in the larger
specimens than in the smaller ones.
Specimens of about 30 mm already are shaped much like adults. Scales in part
are developed, and general pigmentation has taken place, the lower part of the sides
already being silvery in some specimens. The dark specks on the dorsal and caudal
fins have increased greatly in number, but the chromatophores along the base of anal
of juveniles remain faintly visible. The gill rakers have increased considerably in pro-
portionate length, and those of the upper limb now extend downward across those of
the lower limb; the filaments, too, are much better developed. The maxillary remains
shorter than in adults, for it scarcely reaches below the middle of the eye. Striations are
faintly visible on the upper part of the opercle, the ventral scutes are fully developed,
and the pelvic fin now is situated under the anterior rays of the dorsal, as in adults.
The dark peritoneum is plainly visible through the abdominal walls at this stage of
development.
Little is known about the rate of growth. Length frequencies, based on 186 fish,
suggest that a maximum length of about 100 mm (4 in.) is attained at one year of age,
probably 150 mm (6 in.) at two years. However, some of the fish in each year-class are
much smaller, some being only about 80 mm long at one year of age and about 125 mm
at two years. The rate of growth seems to be about the same as that of tyrannus at
Beaufort, North Carolina.
Fishes of the Western North Atlantic 369
Reproduction. As in the case of their growth, little is known about reproduction in
the Largescale Menhaden. Goode (45: 577) quoted Silas Stearns as follows:
The first traces of spawn are found in May. But by July it has become sufficiently developed to be noticed
by any person unaccustomed to the examination of such objects. In the latter part of September or first of October,
at which time they are seen in abundance, the ovaries are sufficiently grown to distend the fish’s abdomen, yet
not fully ripe. When they are next caught, in November and December, on the sea-beach, they are without
ovaries and show signs of having spawned.
Since the Gulf species, patronus and gunteri, were not distinguished until recently, one
cannot be sure which species Silas Stearns observed. Even now it is not known whether
the spawning seasons of the two are different.
The large examples at hand were all taken in February and March, and in these
the gonads are in a collapsed state, as though the fish had spawned recently. Further-
more, the juveniles at hand, 28-60 mm TL, were taken in late winter and during the
spring. In fact, there are some postlarvae, 19-24 mm TL, taken at Dead Man’s
Island, Corpus Christi, Texas, March 31, 1926, which are believed to be this species.
Thus it is indicated that spawning takes place during the winter, that is, rather later
in the year than the time suggested by Silas Stearns.
Migrations and Habitat. These Menhaden, like tyrannus, travel in schools while they
feed on plankton at the surface. However, at times they apparently live at or near the
bottom also, for the Bureau of Fisheries schooner, Grampus, took many specimens with
an otter trawl in February and March 1917 off Corpus Christi and Galveston, Texas.
Gunter also reported their capture in the Gulf with an otter trawl (51: 29).
According to Gunter (51), this species was not taken in water having salinities
below 25.0, which agrees in general with the places of collection of the specimens now
at hand. At least all of the larger specimens of patronus were taken either offshore or in
localities where the salinity would be expected to run rather high. However, small to
half-grown examples are at hand that were caught where the water must have been
nearly fresh, as at Manchac Pass on the Illinois Central Railroad, north of New Or-
leans, Louisiana, and in Grand Plains Bayou, near Baldwin Lodge, Mississippi.
Such a habitat, for both young and adults, would correspond to that of tyrannus.
It was stated by Silas Stearns that the fish are not seen in shallow water during
winter but appear in the spring and remain until late fall (## Goode, 45). He said that
in fine weather the fish could be seen approaching the coast in great schools, and that,
once they had reached the inside waters, a large school would break up into smaller
schools. He also said that quiet bayous, creeks, and nooks in the bay were favorite
“feeding or playing grounds.” Furthermore, he noted that small schools feed at the
mouths of rivers, and that upon becoming “‘accustomed to the fresh water,” they move
upstream to quiet places. Again, one does not know whether Stearns observed one or
both species inhabiting the Gulf.
Food. There is every reason to believe that the food of patronus is essentially the
same as that of tyrannus. Both species feed at the surface in dense schools, have a giz-
zard-like stomach, and a long intestine, which suggest a vegetable diet. The presence
24.
370 Memoir Sears Foundation for Marine Research
of mud in the stomach suggests that they also feed on the bottom. Stearns said,
“The Gulf menhaden are a bottom-feeding fish, as their stomachs always contain
soft brown mud, from which I suppose they extract microscopic animal and plant
matter” (45).
Parasites. As in Atlantic menhaden, the isopod O/encira praegustator (Latrobe) is
frequently found in the mouth of Largescale Menhaden (107: 231). Stearns (45) like-
wise reported it, saying:
When the Gulf Menhaden arrive in spring, each one has a parasite in its mouth, a crustacean called
Cymothoa praegustator. This is found always in one position, clinging with its claws to the roof of the fish’s mouth,
with its head looking forward and very near to the jaws of the fish. The parasites remain with the menhaden
as long as the latter is in salt water; in brackish water they are less frequently observed, disappearing altogether
in fresh water. With all the fall fish of this species which I have examined there were no parasites.
Once more it is possible that Stearn’s observations may have been based on one or
both species inhabiting the Gulf.
Two species of copepods, Bomolochus teres Wilson and Lernanthropus brevoortiae
Rathbun, have been reported as parasitic on Gulf Brevoortia (12: 582, 588), but it is
not known whether one or both species served as host.
Relationship. This species, closely related to tyrannus of the Atlantic coast, generally
has been considered identical to it or only subspecifically distinct. However, it differs
in so many respects from syrannus that it apparently must be recognized as a distinct
species. Furthermore, the range of the two species is discontinuous, for neither occurs
in southern Florida, south of the Indian River on the Atlantic or south of Tampa on
the Gulf.
The body depth in patronus compared with tyrannus is a bit greater, the shape is
different, the head and body are deeper anteriorly, and the ventral outline is notably
more strongly convex. The deepest part of patronus’ body is well in advance of the tips
of the pectoral fin and only about an eye’s diameter behind the margin of the opercle,
whereas in ¢yrannus it usually is over the tip of the pectoral and notably more than
an eye’s diameter behind the margin of the opercle. The differences in shape are
plainly evident in Figs. 86 and 91. Also, in parronus the fins generally are longer,
the scaly sheath on the base of the dorsal is broader, the axillary process of the pec-
toral usually is shorter, and the modified scales in front of the dorsal are larger and
have shorter pectinations; in fact, overlapping in the number of modified scales is
slight, occurring in only some of the rather numerous specimens examined.
The extremes of variation, as well as the usual variation, are given in the Descrip-
tion, and from those data it is evident that some overlapping occurs in every proportion
and count. Nevertheless, the average differences are great in some of them. The more
outstanding ones are the following, with those for patronus given first (in per cent of
standard length): depth 38.8, 34.2; head (length) 33.6, 32.2; head (depth) 33.5, 27.8;
pectoral fin 20.9, 18.7; lower lobe of caudal 34.3, 29.4; vertebrae 45.8, 48.0; scales
along middle of side 41.7, 48.2; modified scales on back in advance of dorsal fin 28.0,
36.0; and ventral scutes 29.8, 32.5.
Fishes of the Western North Atlantic PAN
The differences in shape and depth of body, already mentioned, are more out-
standing in young adults than in mature individuals. In fact, the depth in patronus is
already about as great in fish 40-60 mm TL as in large ones; in examples of zyrannus
of similar size the body is notably more slender; for example, in 34 specimens of patronus
falling within the range mentioned, the depth is 36.4-45.0°/, of SL, whereas in
60 examples of tyrannus the range is 25.8—35.8.
Variation. \t is not evident at present that specimens vary between different areas,
as in those from Florida at one extreme of the range and those from Texas at the other.
Sufficient specimens for a thorough study of variation are at hand from Texas only.
The young, as stated elsewhere, acquire the shape of the adult very early in life, and
the pronounced variation in depth of body is therefore an individual variation rather
than one of age. There is also considerable variation in the number and intensity of
the dark spots on the side, with some specimens, including large ones, having only
a shoulder spot (p. 367).
Commercial Importance. Commercial utilization of patronus is a recent development.
Statistics at hand show for 1940 a catch of menhaden for Mississippi only, this being
25,194,000 pounds. For 1950-1953, however, the following catches (probably patro-
nus for the most part but including some gwnteri) were reported for the Gulf coast
(in pounds): 326,030,100 for 1950, 351,965,100 for 1951, 459,983,900 for 1952,
and 437,045,000 pounds, valued at $5,189,000, for 1953.
Range. Gulf coast of Florida to Texas. It was recorded once for as far south on
the western Florida coast as Tampa (55: 211), but at present there are no specimens
in the National Museum collection from farther south than Apalachicola, Florida.
There are many from as far west and south as Corpus Christi, and the types (USNM
892) are from Brazos Santiago, Texas.
Synonyms and References:
Brevoortia patronus Goode, Proc. U.S. nat. Mus., I, 1879: 39 (orig. descr.; “descr.” based on specimens from
Brazos Santiago, Texas [USNM 892]; ‘‘diagnosis” based in part on specimens from “Mouth of Rio
Grande” [USNM 891], which are B. gunteri; “table of measurements” includes both species); Goode,
Rep. U.S. Comm. Fish. (1877), 1879: 26 (after Goode, 1879); Jordan and Gilbert, Bull. U.S. nat.
Mus., 16, 1882: 269 (descr., may be based in part on B. gunteri); Goode in Goode ef a/., Fish. Fish.
Industr. U.S., 1 (3), 1884: 575, pl. 206 (movements, parasites, reprod., food); Richardson, Bull. U.S.
nat. Mus., 54, 1905: 21 (isopod parasite); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928),
2, 1930: 44 (range, synon.); Gunter, Ecol. Monogr., 8, 1938: 355 (season. var. in abund., Louisiana);
Hildebrand, Smithson. misc. Coll., r07 (18), 1948: 13, fig. 3 (synon., relation., descr., range).
Brevoortia tyrannus patronus Evermann and Kendall, Bull. U.S. Fish Comm. (1892), 12, 1894: 105, pl. 21
(Galveston, Texas; differences between Atlantic and Gulf specimens slight); Evermann, Rep. U.S.
Comm. Fish. (1898), 1899: 309 (many young at Grand Plains Bayou, Mississippi); Weymouth, Proc.
U.S. nat. Mus., 38, 1911: 136 (Cameron, Louisiana); Fowler, Monogr. Acad. nat. Sci. Philad., 7,
1945: 365, 372 (refs., occur.).
Brevoortia tyrannus Henshall (not of Latrobe), Bull. U.S. Fish Comm. (1894), 14, 1895: 211 (Tampa, Florida);
Evermann and Kendall, Proc. U.S. nat. Mus., 3Z, 1906: 74 (synon., discus. lit. on Brevoortia; cf.
specimens from various locals.; descr. of two from Argentina); Bere, Amer. Midl. Nat., 17 (3), 1936:
582, 598 (copepods); Gunter, Publ. Inst. mar. Sci. Texas, 7 (1), 1945: 29 (Texas coast habitat).
ae Memoir Sears Foundation for Marine Research
Brevoortia smithi Hildebrand 1941
Yellowfin Shad
Figures 84, 85, 92
Study Material. A total of 18 specimens, 120-315 mm TL, 91-240 mm SL,
collected at Beaufort, North Carolina; Cumberland Sound, Georgia; and at the “mouth
of St. Johns River” and in the “Indian River,” Florida; the type, 295 mm TL, 225 mm
SL, included among the material from Beaufort, USNM 118723. Also, specimens
ane
SON
SOG
Ficure 92. Brevoortia smithi, 295 mm TL, 225 mm SL, Beaufort, North Carolina, type, USNM 118723.
Drawn by Ann S. Green.
from Boca Ciega, Pinella County, and off Englewood, coast of western Florida, that are
intermediate in some respects between B. smithi and B. gunteri, received through the
kindness of William Gosline.
Distinctive Characters. See Relationship.
Description. Proportional dimensions in per cent of standard length, and counts,
based on specimens listed in the Study Material, 91-240 mm SL.
Body: depth 36-43. Scales: oblique series along middle of
Caudal peduncle: depth 9.5—-12. side (too irregular to count accu-
Head: length 29—-32.5. rately) ca. 60-70.
Snout: length 6.88.0. Modified scales: in a series in front of dor-
Eye: diameter 6.1-7.5. sal fin, 39-45.
Interorbital: width 6.2—7.4. Ventral scutes: 30-32.
Maxillary: length 13.5-15. Fin rays: dorsal 18-20; anal 22 or 23;
Mandible: length 17—18.5. pectoral 16 or 17.
Anal fin: length of base 19-21. Vertebrae: 45-47 (9 specimens).
Pelvic fin: length 8.5-11.
Pectoral fin: length 18.5—-21.5.
Fishes of the Western North Atlantic Nae
Bopy deep, rather strongly compressed, its greatest thickness notably less than
half of the depth, its greatest depth at vertical from a point somewhat in advance of
origin of dorsal and about midway between tip of pectoral and base of pelvic, 2.3-2.75
in SL; ventral outline notably more strongly convex than the dorsal outline, more than
half of the greatest depth being below a straight line drawn through lower margin of
eye to middle of base of caudal. CaupaL PEDUNCLE rather deep, 2.6—3.4 in head, and
3.5-4.0 in greatest depth of body.
Scazs closely adherent, the exposed part three or more times deeper than long,
the scales themselves less than two times deeper than long (Fig. 85 p); scales with
prominent serrae ending in blunt points, not long or hair-like and not extending across
exposed part of next scale; the scales greatly reduced in size on back and at base of caudal,
in rather irregular series (difficult to count accurately); about 5-8 rows of scales ex-
posed between tip of pectoral and base of pelvic. A row of enlarged modified scales
on each side of median line of back in front of dorsal fin, the scales variable in number,
39-45 in each series. VENTRAL sCUTES not strong, 18 or 19 in front of pelvics, 12-14,
most frequently 13, behind them.
Heap 3.2—3.4 in SL, its depth 3.25-3.6. Snour compressed, blunt, with a
prominent median notch, its length 3.7-5.0 in head. Eye 4.2-5.2. INTERORBITAL
4.2—4.9. CHEEK much deeper than long. Maxriary broadly rounded, reaching from
under middle of eye to posterior margin of pupil, 2.1-2.3 in head. Manp1ste in-
cluded in upper jaw, its upper margin (within mouth) nearly straight, its length to
joint 1.7—1.9 in head. Upper section of Opercie with very feeble radiating striae.
GILL RAKERS very numerous, those on upper limb extending downward and over
those on lower limb, the longest ones fully as long as snout, 121-149 in adults.
TrEeETH absent.
Dorsat fin rather high anteriorly, the margin rather deeply concave, the longest
rays equal to length of snout and eye to posterior margin of pupil, the last ray consider-
ably longer than preceding ones, the origin about equidistant between margin of snout
and base of caudal; a very low sheath on base of fin, composed of a single row of
scales. Caupat deeply forked, the middle rays about as long as eye, the lobes long, the
lower one slightly the longer, exceeding the length of head, 2.7-3.1 in SL. Anat
notably lower than dorsal, little elevated anteriorly, its margin nearly straight, its
origin under or a little behind tip of last ray of dorsal, its base 4.75—5.25 in SL; a
very narrow sheath at base. Petvic fin with an oblique margin, its length 2.8—3.4 in
head, its outermost ray nearly twice the length of the innermost one, the fin inserted
under or slightly behind vertical from origin of dorsal. Precrorat fin long, scarcely
falcate, its length 4.65—5.4 in SL and 1.45-1.75 in head, its longest ray fully four
times longer than the shortest one, the fin failing to reach base of pelvic by a distance
somewhat greater than half of diameter of eye. AxILLARY APPENDAGE of pectoral vari-
able, reaching to or well beyond midlength of fin, 2.6—4.0 in head.
Color. Back, in fresh specimens, bluish green, lighter than in tyrannus (evident
also in preserved specimens). Sides silvery. A large black spot at shoulder, not followed
374 Memoir Sears Foundation for Marine Research
by smaller dark spots. Fins golden yellow; the margin of caudal paler than rest of fin.
Peritoneum black.
Size. The largest example seen, taken at Beaufort, North Carolina, was 330 mm
(13.2 in.) TL. This is about equal to the usual length of fish that compose the fall and
winter runs of ¢yrannus in the same vicinity.
Reproduction and Development. Nothing is known about reproduction in this species,
notwithstanding the fact that the eggs and young were sought diligently at Beaufort,
North Carolina, for about six years. The several adults taken there during spring and
summer and examined in the laboratory showed no signs of development.
The smallest specimen at hand, 120mm TL, differs from large ones in having
a deeper body and less strongly serrated scales. The last mentioned difference is quite
in keeping with other species of Brevoortia, for the serrations (pectinations) apparently
increase in length with age in all species. It has been pointed out that young adults
of patronus are quite as deep as large ones, which is contrary to the rule in tyranaus.
The very deep-bodied young adults of smithi suggest that the young of this species too
may acquire a deep body at an early age.
Migrations and Habits. Yellowfin Shad were taken at Beaufort, where their oc-
currence was observed several years in succession. They seem to be present there only
during spring and summer and only in the local estuaries. If they occur off Beaufort
Inlet, where large ¢yrannus are taken, they escaped our nets. Although the fish probably
leave brackish water to spawn, the adults, contrary to those of tyrannus, do frequent
brackish water during a portion of the year.
According to fishermen at Beaufort, these Menhaden do not school. This was
confirmed by our investigations, for we took at most 10-12 a day in a pound net
operated for several years in the estuary of Newport River, and rarely were more than
two taken in a single haul with a large collecting seine. In contrast, hundreds of tyran-
nus, if taken at all, generally were present when the pound net was fished.
Yellowfin Shad are much more active fish than tyrannus. When they are caught
in a seine they dash here and there and strike the net again and again in an
effort to escape, whereas syrannus generally strike the net once and then allow them-
selves to be hauled in without making a further effort to flee. The two species, when
once their habits are known, may be identified by their behavior when surrounded
by a seine.
Food. These fish, like tyrannus and other species of menhaden, have a thick-walled
gizzard-like stomach and a long intestine. A fish 330 mm TL and 247 mm SL, for
example, had an alimentary canal 937 mm long; that is 380 °/, of its SL. The stomach
contents of this same specimen consisted of diatoms and spicules of a sponge.
Parasites. Only one parasitic copepod (Lernaeenicus radians)*° has been found on
these Menhaden. The conspicuous isopod Olencira praegustator, very common in the
mouth of ¢yrannus, has not been seen thus far in smithi.
Relationship. The differences between Brevoortia smithi and B. gunteri are given
40. Identification by Paul L. Illg, at the U. S. National Museum.
Fishes of the Western North Atlantic BGs
on p. 379. B. smithi differs from B. tyrannus in having much smaller scales, in less regular
series; notably shorter serrations or pectinations on the scales of specimens of equal
size (Figs. 858 and 85D); more feeble radiating striations on the upper part of the
opercle; and a longer and more pointed pelvic fin. Furthermore, B. smithi has a rather
more strongly compressed and deeper body and a deeper caudal peduncle, a somewhat
smaller head, a rather larger anal fin with a somewhat greater number of rays and a
longer base, and somewhat longer caudal and pectoral fins. Also, B. smithi is lighter
in color, has more definitely yellow fins, and has no small dark spots behind the large
shoulder spot. The two species are separable by touch, as syrannus is very slimy and
slippery whereas smirthi is not.
Variations. The variations among the limited number of specimens known are not
pronounced. The smallest specimen, 120 mm TL, from Cumberland Sound, Georgia,
is the deepest among the 18 at hand, which suggests that in this species, as in pasronus,
young adults are as deep or deeper than large ones, whereas in tyrannus they are more
slender. However, some half-grown fish from the mouth of the St. Johns River, Florida,
are no deeper than large fish from Beaufort. The great difference in the number of gill
rakers (121-149) on the lower limb may be ascribed in part to age and in part to an
error in counting, as it is impossible to count the close-set rakers accurately. The fresh
examples observed in the field were quite uniform in color.
Commercial Importance. Yellowfin Shad are not numerous enough to be com-
mercially important, for only a few are taken at a time. The total annual catch at Beau-
fort probably does not exceed several hundred pounds. Their abundance south of
Beaufort remains unknown.
The fishermen, at least at Beaufort, distinguish these Menhaden from the common
one and refer to them as the “‘Yellowfin Shad.” They claim that the edible qualities are
superior to those of tyrannus, and the few they catch are generally put on their own
table. I like the flavor of both species and could detect no difference except that the
flesh of smithi may be a little less oily.
Methods of Fishing. The fish are caught chiefly in drag nets at Beaufort, the only
place where they seem to be recognized as a distinct species.
Range. Known from Beaufort, North Carolina, to the “Indian River” (probably
Indian River City), Florida, while specimens intermediate between guwxteri and smithi,
from Boca Ciega and from off Englewood on the west coast,"! are nearer to smithi
than to gunteri.
Synonyms and References:
Brevoortia aureus Hildebrand (not of Agassiz), Rep. U. S. Comm. Fish. (1918), Append. 6, 191g: 7, fig., pl. 1,
fig. 2, scale (orig. descr. of B. smithi Hildebrand 1941; habits, food, cf. B. tyrannus; Beaufort, North
Carolina).
Brevoortia smithi Hildebrand, Copeia, 1941: 224 (B. aureus Hildebrand recognized as a distinct species without
a name; type local. Beaufort, N.C.; type USNM 118723); Hildebrand, Smithson. misc. Coll., 207
(18), 1948: 28, fig. 6 (synon., descr., relation., range).
41. Made available for examination through the kindness of Dr. William Gosline.
376 Memoir Sears Foundation for Marine Research
Brevoortia gunteri Hildebrand 1948
Finescale Menhaden
Figures 84, 85, 93
Study Material. A total of 55 specimens, 70-310 mm TL, 53-230 mm SL, from
bays in the vicinity of Corpus Christi, Rockport, and Galveston, Texas, including
several small ones from the mouth of the Rio Grande and from Brazos Santiago,
Ficure 93. Brevoortia gunteri, 270mm TL, 205 mm SL, Rockport, Texas, type, USNM 129798. Drawn
by Ann S. Green.
part of Goode’s type material of patronus, which included two species (44: 39); and
from Grand Isle, Louisiana (one specimen); all in the USNM collections.
Distinctive Characters. See Relationship.
Description. Proportional dimensions in per cent of standard length, and counts,
based on the study specimens, 53-230 mm SL.
Body: depth 37-45, most frequently 38—
41.
Caudal peduncle: depth, 9.5-12.5, most
frequently 10-11.
Head: length 31-35.5, most frequently
3277-34:
Snout: length 7.0-10.
Eye: diameter 6.6-9.5.
Interorbital: width 7.0-8.0.
Maxillary: length 14.5-17.5, most fre-
quently 15.5—16.5.
Mandible: length 17.5-20.5, most fre-
quently 18.5—19.
Anal fin: length of base 19-23, most fre-
quently 20.5-22.5.
Pelvic fin: length 9.3-11.
Pectoral fin: length 19-23.5, most fre-
quently 20-22.
Scales: oblique series along middle of
side (too irregular to count accu-
rately) about 60-75.
Fishes of the Western North Atlantic Be
Modified scales: in a series in front of dor- quently 22-24; pectoral 15 or 16,
sal 35-45, most frequently 38-42. rarely 14.
Ventral scutes: 27-30, usually 28 or 29. Vertebrae: 43 or 44, rarely 42 (22 speci-
Fin rays: dorsal 17-20, most frequently mens).
18 or 19; anal 20-25, most fre-
Bopy deep, strongly compressed, its greatest thickness only about a third of its
depth, its greatest depth, at a vertical slightly in advance of origin of dorsal or the
tip of pectoral, 2.2-2.7 in SL; ventral outline notably more convex than the dorsal
one, much more than half of greatest depth below a straight line extending through
lower margin of eye to middle of base of caudal. CaupAL PEDUNCLE rather deep, 2.7—
3.5 in head, and 3.6—4.0 in greatest depth of body.
Scates closely adherent, the exposed part 3-4 times deeper than long, the depth
of scale itself about twice its length (Fig. 85 c); adults with prominent serrae ending
in somewhat blunted points, not long or hair-like and not extending quite across the
exposed part of the next scale; the scales much reduced in size on back and on base
of caudal, in rather irregular series (difficult to count accurately); about 2—4 vertical
rows of scales exposed between tip of pectoral and base of pelvic. A row of enlarged
modified scales on each side of median line of back in front of dorsal fin, rather variable
in number, 35-45 in each series. VENTRAL scuTES weaker in large examples than in
small ones, 17 or 18, usually 17, in front of pelvic fins, and 10-13, usually 11 or 12,
behind them.
Heap 2.7—3.2 in SL, its depth 2.75-3.4. SNouT compressed, with a sharp median
notch, its length 3.4-4.4 in head. Eye 3.6-4.8, most frequently 4.2-4.7. InTER-
ORBITAL 4.1—4.7. CHEEK deeper than long. Maxitiary broadly rounded, reaching to,
or a little beyond, vertical from posterior margin of pupil, 1.9—2.2, usually 2.0—2.1,
in head. Manpiste included in upper jaw, its upper margin (within mouth) nearly
straight, its length to joint 1.55—1.95 in head. Upper section of Oprercie with very
feeble radiating striae, or none. GILL RAKERS very numerous, those on upper limb ex-
tending downward and across those on upper part of lower limb, the longest ones
somewhat exceeding length of snout; increasing in number with age and growth: 97
on lower limb of first arch in a specimen 70 mm TL, 113-120 in specimens 100—
I1§ mm, and 135-150 in specimens 200-300 mm. TrzTH absent in all specimens
at hand.
Dorsat fin rather high anteriorly, its margin rather deeply concave, its longest
rays about as long as snout and half of eye, the last ray considerably longer than the
ones immediately before it, its origin generally about equidistant between the margin
of snout and base of caudal; a very narrow sheath on its base, composed of a single
row of scales. Caupat deeply forked, the middle rays about as long as eye, the lobes
long, the lower one the longer, exceeding length of head, 2.55—3.1, usually 2.8—2.9,
in SL. Anat fin much lower than dorsal, its margin somewhat concave, its longest
rays about 0.33 as long as those of dorsal, its origin under, or a little in advance of,
378 Memoir Sears Foundation for Marine Research
tip of last dorsal ray, its base 4.45-5.25 in SL; a narrow sheath of scales at base.
Petvic fin with a nearly straight oblique margin, its length 2.9-3.4 in head, its
outermost ray somewhat less than twice the length of the innermost ray (Fig. 84), the
fin inserted a little behind vertical from origin of dorsal. Pecrorat fin long, slightly
falcate, its length 4.25-5.25 in SL, and 1.4-1.9 in head, its longest ray fully four
times longer than the shortest one, the fin failing to reach the base of pelvic by a
distance usually a little less than diameter of pupil. AxILLARY APPENDAGE of pectoral
variable in length, apparently increasing in length with age, usually about half of
length of fin in large examples and only about a third in small ones, 2.5—4.6 in head.
Color. Preserved specimens dark gray above a line extending through upper margin
of eye to slightly above middle of base of caudal, changing rather abruptly to the silvery
color of the side. A large black shoulder spot in adults, only faintly visible on those
110-125 mm long, absent in smaller ones, this spot not followed by smaller dark
spots in any specimen at hand. Dorsal and caudal fins somewhat dusky, the margin
of the caudal pale; other fins plain translucent. Gunter, when comparing fresh examples
of this Menhaden with patronus, remarked: “The second species [ gunzeri] . .. was more
silvery and had less green color” (51: 27). Peritoneum black.
Size. The specimens examined run larger in size than those of patronus; quite a
few exceed a length of 230 mm (9.2 in.), and the largest has a length of 310 mm (12.4
in.). An example 313 mm (12.5 in.) long has been reported (57).
Development and Growth. The eggs of this species remain unknown, and very little
has been learned about the development, for no specimens less than 50 mm SL are
included in the collections examined. However, there is no reason to believe that
gunteri and patronus differ essentially in their development. Judging from the smaller
specimens at hand, the indications are that the young of this species, at an early age,
reach a depth of body that is equally as great as that in large adults, as in the case of
patronus. The increase in the number of gill rakers with age and growth is shown in
the Description.
In somewhat more than four months, in 1942, from the time of their capture on
February 25 until July, o-class postlarvae 21-30 mm long (identified by Gunter as
this species) grew to lengths of 25-45 mm (52). Gunter also spoke of a group 88-113
mm long that predominated in Aransas and Copano bays in November and Decem-
ber. Although he did not definitely assign these fish to the o-class, he did conclude
that “The curves indicate that this menhaden attained a length of 88 to 113 mm at
the age of one year.” This is about the same size as that indicated for patronus at a
year of age.
Spawning. Gunter has reported that a 12.5-mm male taken in Aransas Bay on
February 25, 1942, was exuding milt and that a ripe female 15.0 cm long was taken
in Copano Bay on March 29, 1942 (51: 28). From Gunter’s observations it appears
that this species, like patronus, spawns during the winter, and possibly into early spring.
Migrations and Habitat. Nearly all that is known about the migrations and habitats
of B. gunteri has been given by Gunter (51), the only student who seems to have recog-
Fishes of the Western North Atlantic 379
nized this species as distinct from patronus. He found gunteri to be much more common
in Aransas and Copano bays than pafronus, and he expressed the opinion that it prob-
ably is ‘‘completely euryhaline,” though he did add that his data strongly suggest a
midwinter exodus from Copano Bay but not from Aransas Bay. The fact that all speci-
mens of guzteri in the U.S. National Museum collection were collected in inshore
waters and that most of the patronus samples were taken offshore support Gunter’s con-
tention. However, there are some small to half-grown specimens of patronus in the
collections that were taken inshore, where the water must have been decidedly
brackish.
The habitats of guwteri and patronus in the Gulf therefore seem to parallel those
of their close relatives in the Atlantic. However, gunteri has been reported as schooling
by Gunter, and if correct, then gunteri apparently differs in that respect from
smithi, which does not school while inhabiting the estuaries of Beaufort, North
Carolina (p. 374).
Food. B. gunteri also is a plankton feeder, and “it may be seen in schools twisting
and turning through the water, with mouth agape ...” (5Z).
Relationship. This species differs from patronus, its relative in the Gulf, in having
much smaller and more crowded scales, which are placed in less regular series and which
in adults have notably shorter serrations (pectinations). It differs further in having
much more feebly developed radiating striae, or none, on the upper part of the opercle
and in the absence of dark spots behind the black shoulder spot. Gunter also said that
it is not slimy when fresh, like patronus.
Its nearest relative is smithi of the Atlantic. Both of these species have deep,
well-compressed bodies that are almost void of slime, small crowded scales, no dark
spots on the anterior part of the body behind the black shoulder spot, and several other
similar characters. But they differ in that the head is rather longer and deeper in guuteri,
the body is more strongly compressed, the maxillary and mandible are somewhat longer,
as is the pectoral fin, the ventral scutes are nearly always fewer, the scales are propor-
tionately deeper, and the vertebrae, so far as known, are definitely fewer. But the pres-
ence on the coast of western Florida of Finescale Menhaden somewhat intermediate
between gunteri and smithi emphasizes the desirability of a more detailed comparison
of them than has been made as yet.
Variation. The specimens at hand vary considerably in depth, as shown in the
Description; the smaller specimens in general are deeper than the larger ones. The
variation in the number of anal rays is also rather great. The wide range shown for the
number of oblique series of scales along the middle of the side may be ascribed in large
part to the difficulty of counting them accurately. The great difference noted in the
number of gill rakers on the lower limb of the first arch results largely from their in-
crease in number with age. The color is quite uniform among the larger examples, but
the smaller ones are lighter in color, and those less than about 115 mm TL lack the
black shoulder spot.
Commercial Importance. No Menhaden plants are operated at present on the coast
380 Memoir Sears Foundation for Marine Research
of Texas where this species seems to be common, and it is not known whether it is
included among the menhaden processed in Mississippi. Neither is it known to what
extent gunteri may have contributed to the reported Texas catches of 47,190,800 pounds
of ‘‘menhaden” for 1950, 30,121,200 pounds for 1951, 52,983,600 pounds for 1952,
and 589,300,000 pounds for 1953. Because of the extreme thinness of the fish, it
probably would not yield much oil.
Range. Grand Isle, Louisiana, to the mouth of the Rio Grande, Texas.
Synonyms and References:
Brevoortia patronus Goode (in part B. gunteri Hildebrand), Proc. U.S. nat. Mus., 1, 1879: 39 (diagn., tables
based in part on B. gunteri).
Brevoortia sp. Gunter, Publ. Inst. mar. Sci. Texas, I (1), 1945: 27 (second species from Gulf, but not named;
relation with other Brevoortia spp.; habitat, food, habits, spawn.).
Brevoortia gunteri Hildebrand, Smithson. misc. Coll., 107 (18), 1948: 31, fig. 7 (orig. descr.; type local. Aransas
Bay, Texas; type USNM 129798).
Genus Opisthonema 1861
Opisthonema Gill, Proc. Acad. nat. Sci. Philad., 1861: 373; genotype by original designation, O. thrissa Gill
equals O. og/inum (LeSueur).*?
Characters. Bovy elongate, well compressed, its greatest depth exceeding length
of head. Bony scutes with sharp points, about 32-36 on sharp edge of chest and ab-
domen. Cueex notably longer than deep. Vertical part of CLavicuLar MARGIN (under-
neath opercle) with two small projections or lobes. Mourn of moderate size. Maxt1-
Lary failing to reach middle of eye. Upper jaw without definite median notch. Lower
JAW projecting beyond the upper. TzrTH missing. Dorsat with about 17-21 rays,
the last ray greatly produced, often reaching nearly to base of caudal. Anat with about
20-25 rays, the last one not greatly enlarged. Petvic fin with 8 rays, inserted under
base of dorsal fin, generally well behind origin of dorsal. Pecrorat without an axil-
lary process, lying in a slight depression. VERTEBRAE about 46-48.
Remarks. The chief distinguishing characters are the large dorsal fin, with nearly
or equally as many rays as the anal and with the last ray greatly produced, frequently
so that it reaches nearly or quite to the base of caudal; the projecting lower jaw with
its abruptly ascending upper margin within the mouth; the long shallow cheek; the
small pelvic, with 8 rays, inserted behind the origin of the dorsal; and the absence of
a process (scale) in the axil of the pectoral.
Range. The genus is known on the Atlantic coast of America from Cape Cod,
Massachusetts, to southern Brazil, and from the West Indies. On the Pacific coast it
is known to range from the Gulf of California to Peru.
42. It is probable but not certain that Gill intended this generic name for the fish now known as O. oglinum. Clupea
thrissa Linnaeus (84: 318) has been applied to both a Chinese and an American species, which many older authors
considered identical; at the time Gill wrote (1861), they were generally so considered. In fact it was not until 1917
that they were properly separated. Regan (r06: 308) cleared up the confusion, and Herre and Myers (56: 234)
have diagnosed still more carefully the Chinese Clupanodon thrissa.—G. S.M.
Fishes of the Western North Atlantic 381
Opisthonema oglinum (LeSueur) 1817
Atlantic Thread Herring, Hairyback, Shad Herring, Bristle Herring,
Sargo, Sargo de Gato, Sardinha Large, Sardinha Sargo
Figure 94
Study Material. A total of 65 or more specimens, 65-250 mm TL, 50-182 mm
SL, from Massachusetts, Rhode Island, New Jersey, Virginia, North Carolina, Georgia,
Florida (both coasts), and Texas; Atlantic coast of Panama; Colombia; at Recife,
Mat
inca ERR
Ore
SOIR ERO
ii
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Yee
Hoy
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ui
Ficure 94. Opisthonema oglinum, 220mm TL, 167mm SL, Beaufort, North Carolina, USNM 51873.
Drawn by Ann S. Green.
Barro Santos, and Sao Francisco, Brazil; Cuba; Puerto Rico; Jamaica; Haiti; Barbados;
and Trinidad. Also several lots of juveniles from Beaufort, North Carolina; Key West,
Florida; Mustang Island, Texas; Colédn and Portobello, Panama.
Distinctive Characters. See Relationship.
Description. Proportional dimensions in per cent of standard length, based on
study specimens, 1 50-182 mm SL.
Body: depth 30-40. Pelvic fin: length 8.5—-11.5.
Caudal peduncle: depth 8.5-11. Pectoral fin: length 17-21.
Head: length 22-28.
Snout: length 5.3-7.3. Scales: 43-50.
Eye: diameter 6.0-8.5. Ventral scutes: 32-36.
Interorbital: width 5.0-6.7. Fin rays: dorsal 17-21, most frequently
Maxillary: length 8.8-12. 19 or 20; anal 21-25, most fre-
Anal fin: length of base 16.7—21.5. quently 22-24; pectoral 15-17.
Vertebrae: 45-48 (15 specimens).
382 Memoir Sears Foundation for Marine Research
Bopy elongate, rather strongly compressed, its greatest thickness often only a
third of the depth, its depth variable, generally greatest at origin of dorsal, 2.5—3.33
in SL; ventral outline strongly convex, the dorsal outline typically only gently convex,
making the body when viewed from the side rather definitely boat-shaped. CaupaL
PEDUNCLE depth 2.3—2.8 in head.
Scaes only moderately adherent, thin, notably deeper than long, with somewhat
irregular membranous edges; 3 or 4 vertical rows of scales exposed between tip of
pectoral and base of pelvic. VENTRAL scuTEs moderately developed, 18 or 19, occa-
sionally 17, in front of pelvic fins, and 14-16, occasionally 13 or 17, behind them.
Heap small, 3.5-4.5. Snour blunt, without a definite median notch, 3.4-4.5.
Eye moderately large, slightly longer than snout, 3.0-4.0. INTERORBITAL 3.8—5.7.
Cueex much longer than deep. Maxitiary broadly rounded, reaching about to ver-
tical from anterior margin of pupil, 2.15-2.75. Mawnprs_e slightly projecting, its tip
definitely lower than dorsal outline of head, its upper margin (within mouth) ascending
abruptly, forming an obtuse angle. GILL RAKERs very numerous, Close-set, those on
upper limb not extending down and over those on the lower limb, the longest about
0.33 of diameter of eye; increasing in number with age and growth: about 40-50 on
lower limb in specimens 40-60 mm TL and up to 100 or more in adults about 235
mm TL. Teeru not discernible.
Dorsat fin elevated anteriorly, the longest among the anterior rays generally ex-
ceeding length of snout and eye, the last ray filamentous, rather variable in length, most
frequently reaching vertical from base of last ray of anal; origin of fin about half as
far from posterior margin of eye as from base of caudal, the distance from margin of
snout approximately 33 °/, of TL, or 2.3 in SL; a low sheath of scales on base of fin.
Caupat deeply forked, its middle rays about as long as eye, its lobes long, the lower
one 3.0—3.5 in SL. Anat fin scarcely elevated anteriorly, its margin nearly straight,
its last ray slightly enlarged, its origin only about a third as far behind vertical from
last dorsal ray as from base of caudal, its base 1.1-1.6 in head; a low sheath of scales
at base. Petvic fin small, pointed when folded, with a nearly straight oblique margin
when spread, its length 2.1-2.8 in head, its innermost ray about 66 °/, of length of
the outermost one, the fin inserted a little in advance of vertical from middle of base of
dorsal, its distance from origin of anal equal to distance from origin of anal to base of
caudal. Pecrorat fin moderately long, its margin nearly straight, pointed, lying in a
slight depression, its length from base of upper ray to tip of fin, 4.75—5.8 in SL, or
I.I-1.6 in head, its shortest rays only about half as long as the longest ones. AxiL-
LARY PROCESS absent.
Color. Back bluish-green in life, the rest of body silvery. A dark shoulder spot
usually present. Rows of scales on back with more or less definite dark lines. Fins
chiefly translucent, the dorsal and caudal lobes generally with dark tips.
Size. A length of 300 mm (12 in.) has been reported. The size generally is much
smaller, however, the usual length being only about 200-250 mm (8-10 in.).
Development and Growth. The eggs and larvae have not been described. The
Fishes of the Western North Atlantic 383
smallest specimens, about 22—25 mm long, rather certainly identified as this species,
were taken at Key West, Florida, on August 25, 1919, but they are not in good con-
dition, having been partly dried. Although they cannot be used for obtaining propor-
tions, it is evident from these and others that the proportionate depth of the body in
examples under 25 mm is much less than in those 100 mm and more. In fact, those
up to 75 mm long generally are notably more slender than adults, though there is
much individual variation at all sizes.
The chief distinctive marks in the smallest young at hand are the projecting
lower jaw; the long shallow cheek; the shallow concavity in the margin of the opercle
in front of the pectoral fin; the positions of the fins, which correspond to those of the
adult; and the number of dorsal and anal rays, which already can be counted with a
fair degree of accuracy. The last ray of the dorsal becomes prolonged much earlier in
some individuals than in others; for example, it already is somewhat produced in a
specimen 30 mm TL from Mustang Island, Texas, but in others up to 37 mm long
from Beaufort there is no sign of prolongation. Ventral scutes can be accurately counted
when the fish attain a length of about 30 mm, and even earlier in some examples. The
gill rakers, as shown in the Description, increase in number with age and growth.
The color of the young is rather variable. The smallest ones have a silvery lateral
band (dark in some lots of specimens, presumably due to preservative used) that
increases in width with age and merges with the generally silvery color of the side in
some specimens only 35-40 mm long; but it remains visible in others up to a length
of so mm or so. In some young, as in some adults, the shoulder spot is absent whereas
it is quite evident in others at a length of 35 mm. Occasionally there are smaller dark
spots within the lateral band behind the shoulder spots; these may or may not disappear
when the lateral band merges with the silvery color of the side. The black peritoneum
is often visible through the body wall in the young.
Little is known about the rate of growth. A few measurements made at Beaufort
indicate that the fish attain a length of about 35-60 mm at the age of one year and
go-120 mm by the end of their second summer. Sexual maturity, at least in some
individuals, is reached at a length of only 150 mm.
Spawning. At Beaufort, North Carolina, spawning takes place during May and
June, as determined from the gonads of adult fish.
Migrations and Habitat. These Thread Herring often school, but solitary examples
were frequently taken at Beaufort, North Carolina, and on the Atlantic coast of Panama.
They hold more closely to salt water than menhaden and some other Clupeidae and are
not known to migrate inshore to spawn. They are essentially tropical and subtropical
fish, and although they stray northward to Cape Cod, they generally are not numerous
north of North Carolina. In North Carolina, where they usually are abundant during
the summer, they are taken from May through September.
Food. These fish, like most of their relatives, feed largely on small organisms,
which they strain from the water with the numerous close-set gill rakers. Copepods
constituted the chief food of specimens from Chesapeake Bay (59: 101). However,
384 Memoir Sears Foundation for Marine Research
small fish, crabs, and shrimp were also found in the stomachs of fish from Port-au-
Prince, Haiti (10: 42).
The stomach does not have a thick wall, as in the menhaden (p. 355) and the
gizzard shad (p. 443). Neither is the intestine especially long, being only about 1.33 °/,
of TL of adult fish. Possibly to compensate for the relatively short alimentary canal,
the stomach is provided with about 100 pyloric coeca.
Enemies. No doubt they are preyed upon by many predatory fishes, such as the
bluefish, Spanish mackerel, weakfish, and many others. Pelicans also feed on them,
especially when the fish school at the surface.
Relationship and Variation. This species is the only one of the genus recognized
from the Atlantic. However, it has a near relative on the Pacific coast of tropical America,
from which typical specimens of og/inum* differ in having a deeper body with a more
strongly convex ventral outline, longer fins, and no small dark spots behind the shoulder
spot.
: In addition to these differences, there are the variations with age. The body be-
comes deeper as growth proceeds, especially up to a length of too mm or so, and the
gill rakers apparently increase in number as long as the fish grow. Insufficient speci-
mens have been studied critically to determine geographical variations. It would seem
probable from the limited data based on specimens of about equal size that those from
South America and the West Indies generally are rather deeper than those from the
Atlantic coast of the United States and have on an average one more ventral scute.
Commercial Importance. The flesh has been reported as being dry, bony, and of in-
ferior flavor. At Beaufort they are seldom used as food, and during a residence there
for ten summers I saw none in the market. However, they appear rather regularly in
the markets in Colén, Panama, and in the West Indies. At Beaufort and perhaps else-
where on the shores of the southern states, they are sometimes taken with purse seines
by the crews of menhaden boats and delivered to the menhaden reduction plants.
However, the amount of oil yielded is very small. Such catches do not reach the statis-
tical reports, presumably because they are not separated from menhaden.
Range. Middle Atlantic States, sometimes straying northward to Cape Cod; south-
ward to southern Brazil (Sao Francisco); common throughout the West Indies.
Synonyms and References:
Megalops oglina LeSueur, J. Acad. nat. Sci. Philad., 7, 1817: 359 (orig. descr.; type local. Newport, Rhode
Island).
Megalops notata LeSueur, J. Acad. nat. Sci. Philad., z, 1817: 361 (orig. descr.; type local. Guadeloupe, West
Indies; said to have longer dorsal than og/ina, also a shorter produced dorsal ray and different color, etc.).
Chatoessus signifer DeKay, N.Y. Fauna, Pt. 4, Fishes, 1842: 264, pl. 41, fig. 132 (orig. descr.; type local.
New York; type in State Coll.; cf. C. ¢hrissa Bloch and M. oglina LeSueur).
Meletta thrissa Cuvier and Valenciennes (not C/upea thrissa Linnaeus 1758), Hist. Nat. Poiss., 20, 1847: 380
(orig. descr.; type locals. New York, Santo Domingo, Guadeloupe, and Martinique).
43. It is entirely possible that a more critical study than time now permits would reveal the existence of more than
one species on the Atlantic coast of the Americas. Specimens from Bermuda, and some from the Atlantic coast
of Panama, were not used in preparing the Description because it was suspected that they may prove to be separate
and distinct.
Fishes of the Western North Atlantic 385
Alausa striata Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 429 (orig. descr.; type local. Guadeloupe).
Chatoessus eumorphus Gosse, Naturalist’s Sojourn in Jamaica, 1851: 290 (orig. descr.; type local. Jamaica).
Opisthonema thrissa Poey (not C/upea thrissa Linnaeus 1758), Repert. Fisico-Nat. Cuba, 1868: 4.19 (refs., diagn.,
etc.); Enumerat. Pisc. Cubens., 1875: 148 (refs., distr.); Uhler and Lugger iz Rep. Comm. Fish. Md.,
ed. 1, 1876: 158; ed. 2, 1876: 134 (descr., synon., Maryland); Metzelaar, Rapp. Vissch. Curacao, ed.
by Boeke, 1, West Indian Fishes, 1919: 12 (when caught, import. as foodfish, local name, range, Curagao
and St. Eustatius).
Clupea thrissa Giinther (not of Linnaeus), Cat. Fish. Brit. Mus., 7, 1868: 482 (synon., descr., range).
Opisthonema oglinum Henshall, Bull. U.S. Fish Comm. (1889), 9, 1891: 373 (common, w. coast of Florida;
Big Gasparilla and Egmont Key); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 101
(descr., size, range); Jordan and Rutter, Proc. Acad. nat. Sci. Philad., 1897: 94 (refs., Kingston, Jamaica);
Evermann and Bean, Rep. U.S. Comm. Fish. (1896), 1898: 241 (in nest of pelican, counts, Indian R.,
Florida); Smith, Bull. U.S. Fish Comm. (1897), 17, 1898: gt (Woods Hole, Massachusetts) ; Evermann
and Kendall, Rep. U.S. Comm. Fish. (1899), 1g00: 56 (Florida); Evermann and Marsh, Bull. U. S.
Fish Comm. (1900), 20 (1), 1902: 86 (descr., range, synon., Puerto Rico); Schreiner and Miranda-
Ribeiro, Arch. Mus. nac., Rio de J., 22, 1903: 92 (Brazilian recs., notes); Bean, T. H., Bull. N. Y.
St. Mus., 60, Zool. 9, 1903: 209 (synon., descr., range, local occur.); Barbour, Bull. Mus. comp. Zool.
Harv., 46, 1905: 112 (distr., Bermuda); Fowler, Rep. N. J. St. Mus. (1905), 1906: 102 (descr., refs.,
New Jersey); Rep. N. J. St. Mus. (1906), 1907: 266, fig.; Smith, N.C. geol. econ. Sury., 2, 1907:
129 (refs., diagn., size, food value, range, Beaufort, North Carolina); Kendall, Occ. Pap. Boston Soc.
nat. Hist., 1908: 40 (refs., New England); Sumner, Osburn, and Cole, Bull. U.S. Bur. Fish. (1911),
31 (2), 1913: 742 (Woods Hole, Massachusetts); Starks, Stanf. Univ. Publ., Univ. Ser., 1913: 8 (var.
in depth, Natal, Brazil); Regan, Ann. Mag. nat. Hist., (8) 19, 1917: 385 (synon., descr., range); Meek
and Hildebrand, Field Mus. Publ., Zool., 75 (1), 1923: 187 (synon., descr., range, Atlantic, Panama);
Nichols and Breder, Zoologica, N. Y., 9 (1), 1926: 41, fig. (New York and s. New England);
Hildebrand and Schroeder, Bull. U.S. Bur. Fish. (1927), 43 (1), 1928: 101 (refs., descr., food, range,
Chesapeake Bay); Beebe and Tee-Van, Zoologica, N. Y., ro (1), 1928: 42, fig. (refs., descr., size, range,
food, Port-au-Prince, Haiti); Jordan, Manual Vert. Anim. NE U.S., ed. 13, 1929: 41 (diagn., range);
Truitt, Bean, and Fowler, Bull. Md. Conserv. Dep., 3, 1929: 51 (descr., Maryland); Breder, Field Bk.
Mar. Fish. Atl. Cst., 1929: 68, fig. (distr., food, size); Nichols, N.Y. Acad. Sci., 10 (2), 1929: 203,
fig. (distr., esp. West Indies and Puerto Rico, diagn.); Jordan, Evermann, and Clark, Rep. U. S. Comm.
Fish. (1928), 2, 1930: 44 (range, synon.); von Ihering, Rev. Indust. Anim., 1 (3), 1930: 230 (counts,
discus., Brazil); Howell-Rivero, Handbk. Jamaica, 1936: 1, in reprint (small specimens, color, size);
Hollister, Zoologica, N. Y., 2T (23), 1936: 284, fig. 49 (caudal skel., Bermuda); Bigelow and Schroeder,
Bull. U.S. Bur. Fish., 48, 1936: 327 (Monomoy Pt., Cape Cod, Massachusetts); von Ihering, Dict.
Anim. Brazil, 1940: 716 (note, common names); Fowler, Arqu. Zool. estad. Sao Paulo, 3 (6), 1941:
134 (refs., Brazil); Bertin, Bull. Soc. Zool. Fr., 66, 1941: 24 (A/ausa striata C. and V. design. a synon.);
Oliveira, Serv. Agr. Dep. nac. Producao Animal, Caca e Pesca, Rio de J., 1942: 24 (descr.); Gunter,
Publ. Inst. mar. Sci. Texas, r (1), 1945: 25 (Texas).
Genus Harengula Cuvier and Valenciennes 1847 *4
Sardines
By
Luis René Rivas
Harengula Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 277; type species, H. /atu/us Cuvier and Valen-
ciennes (not C/upea /atulus Cuvier 1829) equals C/upea clupeola Cuvier 1829.
Characters. VENTRAL SCUTES 25—32, usually 26-31. ScaLes 38-44, usually 39-43.
Vertical anterior edge of shoulder girdle (CLavicuLar MARGIN) with a bilobed dermal
fold. Upper jaw without distinct median notch. GiLL RaKERs on epibranchial of first
arch not folding downwards over those on the ceratobranchial; 26—40, usually 27-39,
on lower limb of first arch. Last ray of Dorsat fin not produced into a long filament.
Frn rays: dorsal 17-20, usually 17 or 18; anal 15-19, 17-19 in western Atlantic
species, usually 16-18; pelvic 7-9, usually 8; pectoral 13-17, usually 15 or 16.
VERTEBRAE 40-44.
Size. The maximum size is about 170 mm.
Range. Western North Atlantic in coastal waters of Bermuda, and from Florida
southward to Brazil. Pacific coast from the Gulf of California southward to Peru.
Habits. Sardines are plankton feeders, and occur close to the coast, especially in
estuaries, where they usually congregate in compact schools near the surface. Certain
species ascend streams for short distances, but not beyond brackish water.
Abundance and Relation to Man. Sardines are most abundant in bays and estuaries
where they may be captured with beach seines and cast nets. They are used for bait
and food throughout the West Indies and are utilized in the canning industries of
Cuba and Venezuela.
Key to Species
1a. Inner edge of palatines with a row of pointed teeth anteriorly; gill rakers 26-32,
usually 27-31, on lower limb of first arch; ventral scutes 25-29, usually 27 or
28; scales not adherent, easily falling off from either fresh or preserved specimens;
snout 2.$—3.4, usually 2.6—3.3, in distance between origins of pelvic and anal
fins; humeral spot absent. Average length 80-140 mm.
humeralis Cuvier 1829, p. 387.
1b. Inner edge of palatines without a row of pointed teeth anteriorly; gill rakers
44. Contribution No. 44 from the Ichthyological Laboratory and Museum, Department of Zoology, University of
Miami.
386
Fishes of the Western North Atlantic 237
28-40, usually 30-39, on lower limb of first arch; ventral scutes 28-32, usually
29-31; scales adherent, not falling off from either fresh or preserved specimens;
snout 3.4—4.3, usually 3.5—4.2, in distance between origins of pelvic and anal fins;
humeral spot present, sometimes faint. Average length 40-90 mm.
2a. Scales in 42-44, usually 43, transverse rows, and 12 or 13, usually 12,
longitudinal rows; predorsal scales 15-19, usually 16-18.
thrissina Jordan and Gilbert 1882.
Pacific coast of México from the Gulf of
California to Acapulco.
2b. Scales in 38-42, usually 39-41, transverse rows, and 11 longitudinal rows;
predorsal scales 11-14, usually 12 or 13.
3a. Anal with 15-17, usually 16, rays, its base usually slightly shorter than
pelvic fin; tip of dorsal fin blackish. peruana Fowler and Bean 1923.
Pacific coast from Panama to Peru.
3b. Anal with 17-19, usually 18, rays, its base usually slightly longer than
pelvic fin; tip of dorsal fin not blackish.
4a. Gill rakers 28-34, usually 30-32, on lower limb of first arch.
clupeola Cuvier 1829, p. 390.
4b. Gill rakers 30-40, usually 32—39, on lower limb of first arch.
pensacolae Goode and Bean 1879, p. 393.
Harengula humeralis (Cuvier) 1829
Red-ear Sardine, Sardina, Sardina de Ley, Sardine, Loosescale Sardine, Pilchard,
Red-ear Pilchard, Sprat, Whitebill, Pincer
Figure 95
Study Material. Numerous specimens, 44-172 mm SL, from Florida, Bermuda,
West Indies, and the Caribbean coast of Central America, USNM, MCZ, and
UMIM.
Distinctive Characters. The presence of pointed anterior palatine teeth, the reduced
number of gill rakers (26-32) and ventral scutes (25-29), and the nonadherent loose
scales distinguish this species from all the others of this genus.
Description. Proportional dimensions in per cent of standard length, and counts,
based on 577 specimens, 44-172 mm SL.
Body: greatest depth 26.0-30.7; greatest Eye: diameter 8.7—11.3.
width 13.5—-16.0. Interorbital: width 5.5-6.6.
Caudal peduncle: least depth 9.5-11.2. Maxillary: length 12.4-1 5.3.
Head: length 27.1-32.8; depth at occi- Dorsal fin: anterior height 18.5—23.2;
put 22.4-25.9. length 22.2-26.0; length of base
Snout: length 7.7—-9.2. 16.6—-19.0.
Zine
388 Memoir Sears Foundation for Marine Research
Anal fin: anterior height 5.6—9.9; length
18.8-21.3; length of base 14.8—
16.9.
Pelvic fin: length 11.0-14.4.
Pectoral fin: length 18.2—22.1.
Distance from tip of snout to origin of: dor-
78.4; pelvic fin 51.6-55.5; pec-
toral fin 26.2—30.5.
Scales: transverse rows 38—42; predorsal
g-Il.
Ventral scutes: 25-29, usually 27 or 28.
sal fin 43.8-46.3; anal fin 76.2—
Gill rakers: 26-32, usually 27-31, on
lower limb of first arch.
vate EON x
nena Onn
44
up y ii be
=
ASG
FicureE 95. Harengula| humeralis, gravid female, 210 mm TL, 158 mm SL, Tortugas, Florida, USNM 116873.
Drawn by Ann S. Green.
PREDORSAL CONTOUR evenly and moderately convex, the PosrporsaL ConrouR
more or less straight from origin of dorsal fin to caudal fin; VENTRAL conrouR much
more convex than the dorsal, the body axis much closer to dorsal than to pelvic fins.
Dorsal fin originating nearer to tip of snout than to caudal base. Petvic fin inserted
about midway between insertion of pectoral and origin of anal. Pecrorat fin inserted
about midway between tip of snout and insertion of pelvic, or (usually) nearer pelvic.
Color. General coloration silvery, especially on lower half of head and body; the
opercular plates with iridescent or pearlish, metallic reflections. Back brownish gray.
Sides of body with longitudinal streaks. Humeral spot absent. Tip of snout and mandible
dusky. Upper sector of iris dusky. Tip of dorsal fin dusky; pectoral, pelvic, and anal
fins colorless; tip of caudal lobes dusky, the pigmentation extending along their inner
margins.
Size. This is the largest species of the genus. The largest examined measures
m2 mmISL.
Range. Florida Keys, Bahamas, West Indies, and Caribbean coast of Central and
South America, from Yucatan to Venezuela; also Bermuda.
Fishes of the Western North Atlantic 389
Synonyms and References:
Clupea humeralis (not Harengula humeralis Cuvier and Valenciennes 1847) Cuvier, Régne Anim., ed. 2, 2, 1829:
318, ftn. (name only, applied to Duhamel’s fig. 4 of “Sardine des Antilles,” 1776); Jordan, Proc. U.S.
nat. Mus., 9g, 1887: 561 (synon. in part, West Indies).
Harengula maculosa Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 292 (orig. descr., life colors, Martin-
ique); Regan, Ann. Mag. nat. Hist., (8) 79, 1917: 387 (synon., descr., range, Bermuda; Florida;
West Indies); Jordan, Copeia, 1918: 46 (validity); Fowler, Proc. Acad. nat. Sci. Philad., 82, 1930:
269 (Grenada); Proc. biol. Soc. Wash., 43, 1930: 145 (size, Bermuda); Jordan, Evermann, and Clark,
Rep. U.S. Comm. Fish. (1928), 2, App. 10, 1930: 43 (part. synon., West Indies); Storey, Stanf. Ichthyol.
Bull., r (1), 1938: 41 (nomencl.); Longley and Hildebrand, Publ. Carneg. Instn. Wash., 535, 1941:
8, 9 (char. in key, synon., remarks, life colors, char., range, Tortugas, Florida).
Alosa apicalis Miller and Troschel ‘z Schomburgk’s Hist. Barbados, Fishes, 1848: 675 (orig. descr., Barbados);
Hill, Handbook of Jamaica, 1881: 125 (comments, Jamaica); Storey, Stanf. Ichthyol. Bull., z (1), 1938:
41 (nomencl., orig. descr. quoted).
Harengula sardina Poey, Memorias, Cuba, 2 (49), 1860: 310 (orig. descr., Cuba); 2 (50), 1861: 384 (Cien-
fuegos, Cuba); Repert. Fisico.-Nat. Cuba, 2, 1866: 16; 1868: 418 (spec. poison., char., compar., Cuba);
An. Soc. esp. Hist. Nat., 5, 1876: 147 (ref., compar.); Jordan and Bollman, Proc. U.S. nat. Mus., rz,
1889: 550 (Green Turtle Cay, Bahamas); Jordan, Proc. U.S. nat. Mus., 12, 1890: 645 (comments);
Bean, Bull. U.S. Fish Comm., 8, 1890: 206 (length, Cozumel, Yucatdn); Jordan and Thompson,
Bull. U.S. Bur. Fish., 24, 1905: 233 (Tortugas, Florida); Fowler, Proc. Acad. nat. Sci. Philad., 58,
1906: 83, fig. 2 (Hailer’s Rock, Florida); Fowler, Proc. U.S. nat. Mus., 78, 1926: 250 (scales, color,
Boca Grande, Florida); Nichols, N. Y. Acad. Sci., 10 (2), 1929: 202 (part. synon., type local., distr.,
diagn., Puerto Rico); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, App. 10, 1930: 43
(synon. in part, range); Longley, Carneg. Instn. Yearbk., 31, 1932: 299 (synon., Tortugas, Florida);
Beebe and Tee-Van, Field Bk. Shore Fish. Bermuda, 1933: 35, fig. (field char., part. diagn., part.
distr., Bermuda); Storey, Stanf. Ichthyol. Bull., r (1), 1938: 41 (nomencl.); Howell-Rivero, Bull. Mus.
comp. Zool. Harv., 82 (3), 1938: 171 (part.synon., type material, holotype design., Cuba); Butsch,
J. Barbados Mus. Hist. Soc., 7 (1), 1939: 18 (Barbados).
Harengula jaguana Poey, Repert. Fisico-Nat. Cuba, Z, 1865: 189 (comments, orig. descr., Bahia de Jagua,
Cienfuegos, Cuba); 2, 1868: 418 (char., compar., Cuba); An. Soc. esp. Hist. Nat., 5, 1876: 147 (refs.,
type local., char., Bahia de Jagua, Cuba); Jordan, Proc. U.S. nat. Mus., 9, 1886: 33 (compar.); Storey,
Stanf. Ichthyol. Bull., z (1), 1938: 41 (nomencl.).
Clupea macrophthalma (not of Ranzani, 1842), Giinther, Cat. Fish. Brit. Mus., 7, 1868: 421 (synon. in part,
descr., range, West Indies).
Clupea apicalis Giinther, Cat. Fish. Brit. Mus., 7, 1868: 441 (descr., Barbados); Jordan, Proc. U.S. nat. Mus.,
9, 1887: 561 (ref., West Indies); Cockerell, Inst. Jamaica Bull., 1, 1892: 15 (descr., Jamaica).
Harengula macrophthalma (not Clupea macrophthalma Ranzani 1842), Goode, Bull. U.S. nat. Mus., 5, 1876:
10, 13, 69 (use as bait, synon. exclud., range in part, abund., length, Bermuda and West Indies); Prov.
Cat. Fish. Bermuda, 1877: 6 (Bermuda); Barbour, Bull. Mus. comp. Zool. Harv., 46 (7), 1905: 112
(distr. exclud., fin rays, scales, capture, abund., Bermuda).
Harengula callolepis Goode in Goode and Bean, Proc. U.S. nat. Mus., 2, 1879: 152 (orig. descr., compar.,
Bermuda); Storey, Stanf. Ichthyol. Bull., r (1), 1938: 4, 41 (nomencl.); Rivas, Proc. U.S. nat. Mus.,
TOO (3263), 1950: 279, 285, 286, fig. 38 (char. in key, synon., comments, compar., Bermuda).
Clupea sardina Jordan, Proc. U.S. nat. Mus., 7, 1884: 106 (abund., habitat, compar., descr., Key West);
Jordan and Swain, Proc. U.S. nat. Mus., 7, 1884: 230 (relation., scales); Jordan, Proc. U.S. nat. Mus.,
9, 1886: 33 (comments, Havana, Cuba); Rep. U.S. Comm. Fish., App. 2, E (24), 1887: 36, ftn.
(descr., range, compar., synon. in part); Proc. U.S. nat. Mus., 9, 1887: 561 (West Indies).
Clupea callolepis Jordan, Proc. U.S. nat. Mus., 9, 1887: 561 (Bermuda).
Clupea sp. Lee, Rep. U.S. Comm. Fish. (1886), 1889: 672 (Nassau, Bahamas).
Harengula clupeola (not Clupea clupeola Cuvier 1829), Jordan, Proc. U.S. nat. Mus., 12, 1890: 646, 647
(char. in key; Key West; and Havana, Cuba, synon. in part, St. Lucia, West Indies).
Sardinella apicalis Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 428, 429 (char. in key, descr.,
part. synon., Barbados); Rep. U.S. Comm. Fish. (1895), 1896: 282 (range).
Sardinella sardina Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 428, 430 (char. in key, descr.,
range, synon.); Rep. U. S. Comm. Fish. (1895), 1896: 282 (range); B. A. Bean, Bahama Islands, Fishes,
390 Memoir Sears Foundation for Marine Research
ed. Shattuck, 1905: 297 (range, abund., Nassau, Bahamas); Barbour and Cole, Bull. Mus. comp. Zool.
Harv., 50(5), 1906: 156 (Progreso, Yucatén); Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 206
(Hailer’s Rock, Florida); Rosen, Acta Univ. Lund., N.S. 7 (2), 1g11: 48 (Nassau and Green Turtle
Cay, Bahamas); Nichols, Bull. Amer. Mus. nat. Hist., 37 (18), 1912: 182 (market, Havana, Cuba);
Bull. Amer. Mus. nat. Hist., 34 (7), 1915: 141 (San Juan Harbor, Puerto Rico); Bull. Amer. Mus.
nat. Hist., 44 (3), 1921: 22 (Turks I., Bahamas); Metzelaar, Trop. Atlant. Vissch., 1, 1919: II, fig. 3
(part. synon., size, habitat, Fuikbay, Curacao, Aruba); Meek and Hildebrand, Field Mus. Publ., Zool.,
T5 (1), 1923: 183 (synon. in part, descr., range, Atlantic Panama); Beebe and Tee-Van, Zoologica,
N. Y., 20 (1), 1928: 40, fig. (ref., char., size, weight, color, range, distr., abund., capture, food, young,
Port-au-Prince Bay, Haiti); Breder, Field Bk. Mar. Fish. Atl. Cst., 1929, 1948: 68 (range, feed. habits,
size, Florida); Parr, Bull. Bingham oceanogr. Coll., 3 (4), 1930: 2 (Bahama locals.).
Sardinella humeralis (not Harengula humeralis Cuvier and Valenciennes 1847), Evermann and Goldsborough,
Bull. U.S. Fish Comm. (1go01), 1902: 149 (descr., comments, Puerto Morelos, Yucatdn); T. H. Bean,
Field Mus. Publ., Zool., 7 (2), 1906: 34 (Bermuda).
Sardinella macrophthalma (not Clupea macrophthalma Ranzani 1842), Evermann and Marsh, Bull. U.S. Fish
Comm., 1, 1902: 85 (descr., comments, range, synon. in part, Puerto Rico locals.); Breder, Bull. Bingham
oceanogr. Coll., 7 (1), 1927: 12 (Royal I., Bahamas; Point Francis, Isle of Pines; Glover Reef, Brit.
Honduras).
Sardinella macrophthalmus (not Clupea macrophthalma Ranzani 1842), T.H. Bean, Field Mus. Publ., Zool,
7 (2), 1906: 34 (Bermuda).
Sardinella maculosa von Thering, Rev. Industr. Anim. Sao Paulo, 3, 1930: 228 (ref., diagn., comments, Brazil).
Harengula sp. Hollister, Zoologica, N. Y., 21 (4), 1936: 282-284 (caudal skel., Bermuda).
Harengula macropthalma (not Clupea macrophthalma Ranzani 1842) Fowler, Proc. Acad. nat. Sci. Philad., 89,
1937: 309 (Haiti). ,
Harengula humeralis Storey, Stanf. Ichthyol. Bull., r (1), 1938: 13, 15, 21, 23, 24, 28, 39, 50 (char., relation.,
range, key, synon., descr., diagn.; St. Lucia; Glover Reef; Jamaica; Cuba; Florida Keys; Bahamas;
nomencl., orig. descr. quoted); Fowler, Fish Culturist, 2r (9), 1942: 66 (use, size, compar., Cojimar,
Cuba); Fish Culturist, 22 (2), 1942: 9 (size, length vent. fin, Bonacca I., Honduras); Monogr. Acad.
nat. Sci. Philad., 6, 1944: 70, 93, fig. 27, 434, 456 (synon., ref., descr., compar., Serranilla Bank; w.
Caribbean); Rivas, Proc. U.S. nat. Mus., T00 (3263), 1950: 279, 282 (char. in key, synon., comment,
compar., range).
Harengula clupeola (Cuvier) 1829
False Pilchard, Sardina Escamuda, Sprat, Petit Cailleu
Figure 96
Study Material. Numerous specimens, 43-149 mm SL, from Florida, the West
Indies and the Caribbean coast of Central and South America, USNM, BOC, and
UMIM.
Distinctive Characters. The absence of pointed anterior palatine teeth, the small
number of gill rakers (28-35), higher ventral scute count (29-32), adherent scales,
and slender body distinguish this species from all the other western Atlantic forms.
Description. Proportional dimensions in per cent of standard length, and counts,
based on 252 specimens, 43-149 mm SL.
Body: greatest depth 29.5—32.0; greatest Head: length 29.0-32.4; depth at occi-
width 13.7-15.1. put 25.9-27.8.
Caudal peduncle: least depth 10.5-11.2. Snout: length 8.0-9.2.
Fishes of the Western North Atlantic 391
Eye: diameter 9.1-11.5. Pectoral fin: length 21.5-22.9.
Interorbital: width 5.9—-6.5. Distance from tip of snout to origin of: dor-
Maxillary: length 13.6-1 5.2. sal fin 44.3-47.5; anal fin 77.4-
Dorsal fin: anterior height 18.8-20.8; 80.4; pelvic fin 52.8—56.2; pec-
length 21.2-25.2; length of base toral fin 28.1-30.8.
16.3-17.1.
Anal fin: anterior height 5.9—8.2; length Scales: transverse rows 40-43; predorsal
17.5-21.1; length of base 14.2- II—I4.
LOs7% Ventral scutes: 29-32, usually 30 or 31.
Pelvic fin: length 13.0-15.8. Gill rakers: 28-35, usually 30-32, on
lower limb of first arch.
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7 ONO
SABI
Ficure 96. Harengula clupeola, gravid female, 180 mm TL, 138 mm SL, Tortugas, Florida, USNM 116872.
Drawn by Ann S. Green.
PREDORSAL CONTOUR evenly and moderately convex, the PosrporsaL ConTOUR
more or less straight from origin of dorsal to caudal fin; VENTRAL conTrouR much
more convex than dorsal contour, the body axis closer to dorsal than to pelvic fins.
Dorsat fin originating nearer to tip of snout than to caudal base. Petvic fin inserted
nearer to insertion of pectoral than to origin of anal. Pecrorat fin inserted closer to
insertion of pelvic than tip of snout.
Color. General coloration silvery, especially on the lower half of the head and body;
the opercular plates with iridescent or pearlish, metallic reflections. Back dark brownish
or bluish gray. Body with dark, longitudinal streaks more conspicuous on back. Humeral
spot faint. Tip of snout and mandible dusky. Upper sector of iris dusky. Fins colorless
except caudal, which is somewhat pigmented at tip of lobes and along inner margin.
Size. This is the smallest species of the western North Atlantic, seldom reaching
a length of more than 100 mm.
392 Memoir Sears Foundation for Marine Research
Range and Subspecies. H. clupeola is the most widely distributed species and has
almost the same range as Harengula humeralis. It is known from the Florida Keys,
Bahamas, West Indies, and Caribbean coast of Central and South America, from
Yucatan to Brazil. The species does not show variation throughout its range whereby
it can be broken up into geographical subspecies.
Synonyms and References:
Clupea clupeola Cuvier, Régne Anim., ed. 2, 2, 1829: 318, ftn. (name only, applied to Duhamel’s fig. 2 of
“Petit cailleu,” 1776); Régne Anim., Disciples ed., 1843: 274, ftn. (on “Petit cailleu” of Duhamel,
1776); Jordan, Proc. U.S. nat. Mus., 7, 1884: 107 (compar., char.); Rep. U. S. Comm. Fish. (1885),
App. 2, E (24), 1887: 33 (compar., Havana, Cuba); Proc. U.S. nat. Mus., 9, 1887: 561 (part. synon.,
West Indies); Storey, Stanf. Ichthyol. Bull., z (1), 1938: 42 (comments).
Clupea macrophthalma Ranzani, Novi Comm. Acad. Inst. Bonon., 5, 1842: 320, pl. 23, figs. 1-4 (orig. descr.,
Brazil); Cope, Trans. Amer. philos. Soc., 24 (3), 1871: 483 (New Providence, Bahamas); Jordan,
Proc. U.S. nat. Mus., 9, 1887: 561 (synon. and ref. exclud., West Indies); Cockerell, Inst. Jamaica Bull.,
I, 1892: 15 (teeth, color, Jamaica); Storey, Stanf. Ichthyol. Bull., 7 (1), 1938: 44, fig. 15 (char., compar.,
holotype).
Harengula latulus (not Clupea latulus Cuvier 1829), Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847:
280, pl. 595 (orig. descr., Caen; Dieppe); Storey, Stanf. Ichthyol. Bull., z(1), 1938: 3, 36, fig. 15
(comments, nomencl., type local. doubtful, on cotype, cf. H. c/upeo/a).
Harengula clupeola Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 289 (descr., Martinique); Gosse,
Naturalist’s Sojourn in Jamaica, 1851: 210, 289 (used as bait, enemies, behav., Jamaica); Guichenot
in Sagra, Hist. fisica. polit. nat. Cuba, 4 (2), 1853: 230 (diagn., comments, Cuba); Poey, Memorias,
Cuba, 2 (49), 1860: 310 (cf. H. sardina); Memorias, Cuba, 2 (50), 1861: 384, 395 (compar., doubt-
ful in Cuba); Repert. Fisico-Nat. Cuba, r, 1866: 378 (ref., range, Cuba); An. Soc. esp. Hist. Nat., 5,
1876: 147 (ref., comment, range, Cuba); Hill, Handbook of Jamaica, 1881: 126 (Jamaica); Jordan
and Bollman, Proc. U.S. nat. Mus., rz, 1889: 550 (Green Turtle Cay, Bahamas); Jordan, Evermann,
and Clark, Rep. U.S. Comm. Fish. (1928), 2, App. 10, 1930:43 (range, ref., West Indies); Storey,
Stanf. Ichthyol. Bull., r (1), 1938: 3, 13, 15, 23, 24, 29, 42, 49, 50, SI, figs. 1-3, 7-9, II, 14, 16, 17
(relation., range, char., synon., descr., diagn., Panama; Glover Reef; St. Lucia; Puerto Rico; Jamaica;
Cuba; Garden Key, Bahamas; nomencl. on H. c/upeola C. and V. 1847 and on Poey’s No. 377 sp. dubia,
1861); Fowler, Fish Culturist, r9 (4), 1939: 28 (Bimini, Bahamas); Arqu. Zool. estad. Sao Paulo, 3 (6),
1941: 133 (Brazil); Fish Culturist, 22 (2), 1942: 9 (vent. fin; Sheen Cay, Honduras); Monagr. Acad.
nat. Sci. Philad., 6, 1944: 124, 434, 456, fig. 28 (synon., ref., descr., Old Providence I., Bahamas;
w. Caribbean; Honduras); Rivas, Proc. U.S. nat. Mus., 100 (3263), 1950: 280, 289-292, pl. 3,
figs. 5, 6 (char. in key, synon., comments, compar., size, range).
? Alosa bishopi Miller and Troschel ix Schomburgk’s Hist. Barbados..., 1848: 675 (orig. descr.; compar.,
size, use as food, Barbados); Hill, Handbook of Jamaica, 1881: 125 (comments, Jamaica); Storey, Stanf.
Ichthyol. Bull., r (1), 1938: 49 (nomencl., orig. descr. quoted).
Harengula species dubia Poey, Memorias, Cuba, 2 (50), 1861: 384; Repert. Fisico-Nat. Cuba, 2, 1868: 418
(descr., cf. H. clupeo/a, Cuba); Storey, Stanf. Ichthyol. Bull., z (1), 1938: 49 (comments).
Clupea humeralis (not Harengula humeralis Cuvier and Valenciennes 1847), Giinther, Cat. Fish. Brit. Mus.,
7, 1868: 422 (synon., ref., descr., range, all in part).
Harengula pensacolae (not of Goode and Bean 1879), T. H. Bean, U.S. Fish Comm., 8, 1890: 206 (Cozumel,
Yucatan).
Harengula macrophthalma Jordan, Proc. U.S. nat. Mus., 72, 1890: 645, 646 (char. in key, synon. and ref.
in part, Port Castries, St. Lucia); Jordan and Thompson, Bull. U.S. Bur. Fish., 24, 1905: 233 (Garden
Key, Tortugas, Florida); Fowler, Proc. Acad. nat. Sci. Philad., 67, 1915: 257 (St. Vincent, Lesser
Antilles); Regan, Ann. Mag. nat. Hist., (8) 29, 1917: 388 (synon. in part, descr., range, Florida; West
Indies; Bahia; Fernando Noronha exclud.); Jordan, Copeia, 56, 1918: 46 (Atlantic spp.); Fowler, Proc.
Acad. nat. Sci. Philad., 7 (2), 1919: 133 (St. Martin, Lesser Antilles); So, 1928: 462 (Puerto Rico);
Nichols, New York Acad. Sci., ro (2), 1929: 202 (synon. and ref. in part, distr., diagn., remarks, habits,
fig., Puerto Rico); Jordan, Evermann, and Clark, Rep. U. S. Comm. Fish. (1928), 2, App. 10, 1930: 43
Fishes of the Western North Atlantic 393
(range, synon. in part); Fowler, Proc. Acad. nat. Sci. Philad., 82, 1930: 269 (Grenada); 89, 1937: 309
(Haiti); Storey, Stanf. Ichthyol. Bull., z(1), 1938: 43 (nomencl., type local., orig. descr. quoted);
Butsch, J. Barbados Mus. Hist. Soc., 7 (1), 1939: 18 (Barbados); Longley and Hildebrand, Publ.
Carneg. Instn. Wash., 535, 1941: 8—r0 (char. in key, compar., synon. in part, var. in depth accord. to
sex, fin rays, vent. scutes, gill rakers, nomencl., range, Tortugas, Florida).
Clupea bishopi Cockerell, Inst. Jamaica Bull., 1, 1892: 15 (color, Jamaica).
Sardinella clupeola Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 428, 429 (char. in key, descr.
in part, nomencl., ref.); Rep. U.S. Comm. Fish. (1895), 1896: 282 (range); B. A. Bean iz Shattuck,
Bahama Islands, Fishes, 1905: 297 (Green Turtle Cay, Bahamas); Metzelaar, Trop. Atlant. Vissch.,
191g: II, fig. 10 (synon., size, Haiti, drawing erroneously ident. as Sardinella anchovia C. and V. 1847).
Sardinella bishopi Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 428, 430 (char. in key, descr.,
relation.); Rep. U.S. Comm. Fish. (1895), 1896: 282 (range).
Sardinella macrophthalmus Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 428, 430 (char. in key, descr.,
compar., range, synon. in part); Rep. U S. Comm. Fish. (1895), 1896: 282 (range); Jordan and Rutter,
Proc. Acad. nat. Sci. Philad., 49, 1898: 94 (ref., body depth, Kingston, Jamaica); Fowler, Proc. Acad.
nat. Sci. Philad., 52, 1900: 118 (Port Antonio, Jamaica; St. Martin and Santa Cruz, Lesser Antilles);
Nichols, Bull. Amer. Mus. nat. Hist., 37 (18), 1912: 182 (capture, Havana Harbor, Cuba); Metzelaar,
Trop. Atlant. Vissch., 1, 1919: 12, fig. 4 (synon., size, habitat, commerc. import., Curacao; St. Eustatius) ;
Breder, Bull. Bingham oceanogr. Coll., r (1), 1927: 12 (Glover Reef); Beebe and Tee-Van, Zoologica,
N.Y., 20, 1928: 41 (ref, field char., size, weight, range, distr., abund., capture, Port-au-Prince Bay,
Haiti); Breder, Field Bk. Mar. Fish. Atl. Cst., 1929: 68 (range, size); Zoologica, N. Y., 78 (3), 1934: 59
(Andros I., Bahamas); Parr, Bull. Bingham oceanogr. Coll., 3 (4), 1930: 3 (comments, Crooked Is.,
Bahamas only); von Ihering, Rev. Industr. Anim. Sao Paulo, 3, 1930: 229 (ref., diagn., Brazil).
Harengula pensacolae Goode and Bean 1879
Scaled Sardine, Sardine, Pilchard, Alewife, Fatback Minnow,
Shiner, Sprat, Sardina Escamuda, Sardina, Sardinha
Figure 97
Study Material. Numerous specimens, 39-149 mm SL, from Florida, the Gulf of
Mexico, Bahamas, West Indies, Caribbean coast of Central and South America, and
Brazil, USNM and UMIM.
Distinctive Characters. The absence of palatine teeth, the high number of gill rakers
(30-40), the number of ventral scutes, the adherent scales, and the deeper body distin-
guish this species from all other western Atlantic forms.
Description. Proportional dimensions in per cent of standard length, and counts,
based on 1,394 specimens, 39-149 mm SL.
Body: greatest depth 29.5—-34.7; greatest Maxillary: length 12.7-14.3.
width 14.0-15.6. Dorsal fin: anterior height 18.1-20.3;
Caudal peduncle: least depth 10.6-11.4. length 23.6-—25.3; length of base
Head: length 28.0-31.5; depth at occi- 16.4—19.0.
put 26.5—28.8 Anal fin: anterior height 6.3—7.1; length
Snout: length 6.9-8.5. 18.9-21.9; length of base 14.3-
Eye: diameter 9.0-12.0. 16.0.
Interorbital: width 6.0—7.6. Pelvic fin: length 12.9-13.8.
394 Memoir Sears Foundation for Marine Research
Pectoral fin: length 21.8—24.1. Ventral scutes: 30-32; usually 31 in p.
Distance from tip of snout to origin of: dor- majorina; 28—31, usually 29 or 30,
sal fin 42.7-45.8; anal fin 78.6— in other subspecies.
81.7; pelvic fin 53.6—55.6; pec- Gill rakers: 30-40; p. majorina with 30—
toral fin 27.1—30.3. 37, usually 33-35; p. caribbaea
with 31-37, usually 32-35; p. pen-
Scales: transverse rows 39-43, usually sacolae, p. cubana, p. floridana, and
40-42; predorsal 11-14, usually D- pinensis with 34-40, usually 35-
12) OF 13; 39.
L2 Zeer
ee
ie ee
SS.
=.
Ficure 97. Harengula pensacolae, 104.mm SL, Province of Pinar del Rio, Cuba, type of H. p. cubana, USNM
132472. Drawn by Janet Ebeling, after Rivas (108).
PREDORSAL CONTOUR evenly and moderately convex, the PosrpoRsaL CONTOUR
more or less straight or very slightly convex from origin of dorsal to caudal fin; Ven-
TRAL CONTOUR much more convex than dorsal contour, the body axis closer to dorsal
than to pelvic fins. Dorsat fin originating nearer to tip of snout than to caudal base.
Petvic fin inserted about midway between insertion of pectoral and origin of anal, or
nearer insertion of pectoral. Pecrorat fin inserted nearer to insertion of pelvic than
to tip of snout.
Color. General coloration silvery, especially on the lower parts of the head and body;
the opercular plates with iridescent or pearlish, metallic reflections. Back dark brownish
or bluish gray. Body with dark, longitudinal streaks, more conspicuous on back.
Humeral spot variable; conspicuous to faint. Tip of snout and mandible dusky. Up-
per sector of iris dusky. Fins colorless except caudal, which is somewhat pigmented
at tip of lobes and along inner margin.
Size. The largest specimen examined measures 149 mm SL.
Range. Widely and generally distributed from Cape Canaveral, Florida, southward
to Brazil, including the Gulf of Mexico, as discussed under Variation.
Fishes of the Western North Atlantic 395
Variation. H. pensacolae has been subdived into five subspecies, namely floridana,
caribbaea, cubana, pinensis, and majorina, the alternative characters of which were pub-
lished in 1950 (10g: key, 279): floridana Rivas 1950, a large-eyed form from the
Florida Keys; caribbaea Rivas 1950, based on specimens with fewer gill rakers and
a more slender body, from the Caribbean, and Central and South America—Nicaragua
to Colombia, Jamaica, Hispaniola, and Puerto Rico; cubana Rivas 1950 and pinensis
Rivas 1950, for those from northwestern Cuba and Isle of Pines, respectively; and
majorina Storey 1938, for those from the Lesser Antilles, St. Lucia southward, and along
the coast of South America from Venezuela to and including Brazil. These slightly
different forms broadly intergrade with one another. As geographical separation of
them would obviously be purely arbitrary, they are all united here under the specific
name pensacolae.
Synonyms and References:
Harengula humeralis (not Clupea humeralis Cuvier 1829), Cuvier and Valenciennes, Hist. Nat. Poiss., 20,
1847: 293 (descr., comments, local. in part; Guadeloupe, Rio de Janeiro to the West Indies; Brazil;
Bahia; Surinam); Castelnau, Anim. Rares Amer. Sud, 2, 1855: 56 (ref. in part, comments, Bahia, Brazil);
Jordan, Proc. U.S. nat. Mus., 73, 1891: 314 (Bahia, Brazil); Jordan and Thompson, Bull. U. S. Bur.
Fish., 24, 1905: 233 (abund., use as bait, Tortugas, Florida); Fowler, Proc. Acad. nat. Sci. Philad.,
58, 1906: 83, fig. 3 (color, Marquesas Keys and Hailer’s Rock, Florida); 67, 1915: 247 (Clearwater
and Palm Beach, Florida).
Clupea humeralis (not of Cuvier 1829), Giinther, Cat. Fish. Brit. Mus., 7, 1868: 422 (synon., refs., descr.,
range, all in part); Cope, in part, Trans. Amer. philos. Soc., T4 (3), 1871: 483 (ref., St. Croix); Jordan,
Proc. U.S. nat. Mus., 9g, 1887: 561 (synon. exclud., West Indies).
Harengula pensacolae Goode and Bean, Proc. U.S. nat. Mus., 2, 1879: 152 (orig. descr., compar., Pensacola,
Florida); Proc. U.S. nat. Mus., 2, 1880: 343 (char., Clearwater Harbor, Florida); Jordan and Gilbert.
Bull. U.S. nat. Mus., 16, 1882: 268 (descr.); Lonnberg, Ofvers Svensk. Vet. Akad. Forh., 57 (3), 1894:
114 (abund., use, St. Petersburg, Florida); Regan, Ann. Mag. nat. Hist., (8) 9, 1917: 387, 388
(char. in key, synon., ref., descr.; Florida; Trinidad); Fowler, Proc. Acad. nat. Sci. Philad., 77 (2),
1919: 129, 133, 144, 151 (Rio de Janeiro, Brazil; St. Martin, West Indies; St. Croix; habitat, behav.,
capture, Useppa I., Florida); 78, 1926: 250 (size, color., Boca Grande, Florida); 80, 1928: 456, 462
(Haiti; Puerto Rico); Proc. biol. Soc. Wash., 46, 1933: 58 (size, Calcasieu Lake, Louisiana); Proc.
Acad. nat. Sci. Philad., g2, 1940: 2 (size, Boca Grande, Florida); Monogr. Acad. nat. Sci. Philad., 7,
1945: 104, 266 (synon., ref., size, use; Elizabeth City, N. Carolina; Florida locals.); Jordan, Evermann,
and Clark, Rep. U. S. Comm. Fish. (1928), 2, App. 10, 1930: 44 (range, Florida); Longley, Carneg.
Instn. Yearbk., 31, 1932: 299 (nomencl.); Hubbs, Publ. Carneg. Instn. Wash., 457, 1936: 174 (char.,
nomencl., compar., Champoton, Campeche, Yucatdn); Storey, Stanf. Ichthyol. Bull., 7 (1), 1938: 3, 7;
13, 15, 18, 21, 23, 25, 33, 50, figs. 1, 2, 4-11, 13 (relation., range, char., cf. H. majorina, descr.,
diagn., Florida and Texas locals.; Yucatén, México); Hildebrand iz Longley and Hildebrand, Publ.
Carneg. Instn. Wash., 535, 1941: 11 (synon. of H. macrophthalma; gill rakers).
Clupea pensacolae Jordan, Proc. U.S. nat. Mus., 7, 1884: 107 (abund., compar., color, Key West, Florida);
Jordan and Swain, Proc. U.S. nat. Mus., 7, 1884: 230 (color, relation., Cedar Keys, Florida); Jordan,
Rep. U.S. Comm. Fish. (1885), App. 2, E (24), 1887: 36 (N. America); Henshall, Bull. U.S. Fish
Comm., 9, 1891: 385 (abund., range, Florida Keys; w. Florida).
Harengula arcuata (not Clupea arcuata Jenyns 1842), Jordan, Proc. U.S.nat. Mus., 12, 1890: 645, 646
(char. in key, synon. in part; St. Lucia; Florida locals.); Henshall, Bull. U.S. Fish Comm., 9, 1891:
373 (synon. in part, abund., Florida locals.); Evermann and Kendall, Bull. U. S. Fish Comm., 12, 1894:
105 (compar., Texas locals.).
Sardinella humeralis (not Clupea humeralis Cuvier 1829), Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1),
1896: 428, 431 (char. in key, descr., range, synon. and ref. in part, types of H. pensaco/ae design.); Rep.
U.S. Comm. Fish. (1895), 1896: 283 (range, ref.); Fowler, Proc. Acad. nat. Sci. Philad., 52, 1900:
396 Memoir Sears Foundation for Marine Research
118 (Port Antonio, Jamaica); Evermann and Marsh, Bull. U.S. Fish Comm. (1gor), 1, 1902: 85
(char. in key, descr., synon., Puerto Rico locals.); Cockerell, Proc. biol. Soc. Wash., 23, 1910: 63 (scales,
Tampa, Florida); Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 206 (Florida); Copeia, 24, 1915: 50
(Santo Domingo); Proc. Acad. nat. Sci. Philad., 69, 1917: 128 (Colén, Panama); Metzelaar, Trop.
Atlant. Vissch., 1, 1919: 11 (synon., size, doubt. poison., habitat, Curagao; St. Eustatius).
Sardinella macrophthalmus (not Clupea macrophthalma Ranzani 1842), Meek and Hildebrand, Field Mus.
Publ., Zool., 75 (1), 1923: 182, pl. ga (char. in key, synon. in part, descr., comments, range, Toro Point,
Colén, Porto Bello, Panama).
Harengula sardina (not of Poey 1860), Fowler, Proc. Acad. nat. Sci. Philad., 89, 1937: 309 (Port-au-Prince,
Haiti).
Harengula majorina Storey, Stanf. Ichthyol. Bull., z (1), 1938: 7, 18, 21, 23, 25, 32, 42, 44, 50, 51, figs. 1,
8, 9, 17 (char., relation., compar., range, synon., diagn., comments, nomencl., West Indies; St. Lucia;
Santos [Sao Paulo] Brazil); Fowler, Proc. Amer. philos. Soc., 82 (5), 1940: 745 (ill., Rio de Janeiro,
Brazil); Hildebrand iv Longley and Hildebrand, Publ. Carneg. Instn. Wash., 535, 1941: 11 (comments);
Fowler, Arqu. Zool. estad. Sao Paulo, 3 (6), 1941: 133 (ref., Brazil); Monogr. Acad. nat. Sci. Philad.,
6, 1944: 144, 547, fig. 29 (synon., ref., descr., cf. H. pemsacolae, St. Andrews I., local., ref., w. Car-
ibbean).
Harengula macrophthalma (not Clupea macrophthalma Ranzani 1842), Gunter, Publ. Inst. Mar. Sci. Texas,
I(t), 1945: 25, 128 (abund., capture, salinity, temp., size, ‘Texas locals.).
Harengula pensacolae majorina Rivas, Proc. U.S. nat. Mus., 100 (3263), 1950: 292-294, pl. 4, fig. 1 (synon.,
comments, compar., size, range).
Harengula pensacolae caribbaea Rivas, Proc. U.S. nat. Mus., 100 (3263), 1950: 294-296, pl. 4, fig. 3 (synon.,
orig. diagn., comments, compar., size, range).
Harengula pensacolae pensacolae Rivas, Proc. U.S. nat. Mus., 100 (3263), 1950: 281, 296-298, pl. 3, figs.
7, 8 (char. in key, synon., comments, compar., size, range).
Harengula pensacolae cubana Rivas, Proc. U.S. nat. Mus., 100 (3263), 1950: 281, 298-301, fig. 40, pl. 5,
fig. 1-3 (char. in key, orig. descr., compar., size, range, w. Cuba).
Harengula pensacolae floridana Rivas, Proc. U.S. nat. Mus., Too (3263), 1950: 281, 301, 302, pl. 4, fig. 2
(char. in key, orig. diagn., compar., Florida).
Harengula pensacolae pinensis Rivas, Proc. U.S. nat. Mus., 100 (3263), 1950: 281, 302, fig. 41, pl. 5, figs.
4-6 (char. in key, orig. diagn., compar., Isle of Pines, Cuba).
Genus Sardinella Cuvier and Valenciennes 1847
Spanish Sardines
By
Samuel F. Hildebrand
(continued)
Sardinella Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 184.7: 261; genotype 8. aurita Cuvier and Valencien-
nes, first designated by Gill, Proc. Acad. nat. Sci. Philad., 1861; 35.
Generic Synonym:
Sardinia Poey, Memorias, 2, 1860: 311; genotype by monotypy, Sardinia pseudo-hispanica equals 8. anchovia
Cuvier and Valenciennes.
Characters. Bopy quite slender, compressed, its greatest depth equal to or less than
length of head. Bony scutes rather weak, 32-34 along moderately sharp edge of
chest and abdomen. Sca.zs thin, moderately adherent, with one to several vertical
grooves. Vertical part of anterior CLavicuLAR MARGIN (cleithrum) with two mem-
branous projections or lobes, a definite concavity between them (Fig. 100). OprrcLe
without radiating grooves. MoutTu moderately small. Maxittary generally extending
a little beyond anterior margin of eye. Upper jaw without a median notch. Lower
jaw projecting slightly beyond the upper. TEETH absent in upper jaw; several in an-
terior part of lower jaw; minute teeth also on palatines, pterygoids, and tongue but
absent on vomer. GILL RAKERS numerous, close-set, increasing in number with age,
difficult to count (ftn. 47), about 70-130 on lower limb of first arch in adults;
rakers on upper limb not overlapping those on lower limb, decreasing gradually in
length toward anterior part of lower limb. Dorsat with 17-19 rays, the last one not
produced. Anat with 16—20 rays, the last two notably enlarged, finlet-like. Petvic
fin inserted under base of dorsal, almost constantly with 9 rays. Pecrorat fin with-
out a free axillary process, the fin lying in a slight depression.*® VERTEBRAE about
4S—4]s
Remarks. Confusion in regard to the relationship of Sardine/la Cuvier and Valen-
ciennes, Sardina Antipa,‘* and Sardinia Poey has existed and even now has not been
cleared up completely. This relationship has been discussed by Regan (106: 377);
Jordan (70: 46), Hubbs (61: 261-265), and Longley (87: 5), and apparently the con-
clusion may be reached that Sardinella differs from Sardina in the absence of radiating
grooves (or ridges) on the opercle and in the presence of two membranous lobes on the
vertical edge of the clavicular margin (cleithrum). Further, it appears that Sardinia
45. The generic description is based wholly on American specimens, as adequate material from the other side of the
Atlantic is not available.
46. Sardina Antipa 1905 appears to be a synonym of Arengus Cornide 1788 (see 23: 43).—G. S. M.
Soy),
398 Memoir Sears Foundation for Marine Research
Poey is a synonym of Sardinella and that Clupanodon has no American representative.
As the last ray of the dorsal in C/upanodon is prolonged, its relationship is with Opistho-
nema, Dorosoma, and Signalosa (here a synonym of Dorosoma) rather than with Sardinella.
In his discussion, Hubbs (61: 261-265) set up a new genus, Sardinops, for the sardines
of the Pacific and South Africa. This genus differs from Sardine//a principally in the
absence of the fleshy lobes on the cleithrum, in the presence of oblique ridges on the
opercle, and in the overlapping gill rakers, those on the upper arch extending down-
ward over those on the lower limb.
Species. The species of this genus remain somewhat obscure. Among the three
recognized as coming within the scope of this work, pinnu/a is definitely distinct accord-
ing to the specimens at hand; however, specimens from Cuba and Jamaica, not in good
condition, seem to be more or less intermediate between anchovia from the Atlantic
coast of the United States and brasiliensis from Port-of-Spain, Trinidad, and southward.
Such specimens also are included among collections from Rio de Janeiro, Brazil. More
material and further study will be required to determine the exact relationship.
The number of gill rakers on the first arch seems to be the best diagnostic charac-
ter, but the rakers are so numerous, fine, and close-set that they are exceedingly difficult
to count accurately.47 Consequently, the use of this character requires much time and
patience. Its ready use is limited further by the increase in number with age and growth,
making it necessary to compare the number present in specimens of nearly equal size.
Other characters having limited diagnostic value are depth of body, size of head,
eye, and mouth, development of the palatine and pterygoid teeth, the position of dorsal
and pelvic fins, and length of pectoral fin. The color, so far as can be determined from
the preserved specimens at hand, is rather uniform among the species recognized. The
species are carnivorous, occur in schools, and are marine in habitat.
Range. Cape Cod, Massachusetts, to southern Brazil, but not yet known from
México to Colombia. They occur also off Bermuda, Cuba, Jamaica, Trinidad, in the
eastern Atlantic, and in the Indo-Pacific (106: 377).
Key to Species of the Western Atlantic
1a. Distance from margin of snout to origin of dorsal about equal to 33 °/) of TL,
41-43.5 °/, of SL; eye equal to, or usually shorter than, snout, its diameter 5.3—
7.3 %/, of SL; snout 6.0—7.6; gill rakers about 70-100 on lower limb of first
arch (fewer in young); pectoral fin failing to reach base of pelvic by a distance
nearly or quite equal to length of snout and eye.
47. It was found necessary to remove the gill rakers from the arch to make an accurate count. The arch was severed from
the body near the base of the rakers, the membrane was freed from the bony arch at one end, the loosened end was
grasped with forceps, and a gentle pull then freed the membrane with the rakers intact. In a watch glass under low
power of a binocular, the rakers could be separated by ones, twos, or threes, an accurate count thus being obtained.
The counts made previously either with the arch in its natural position or after removal from the body ran much too
high, the error being greatest among those with the highest number of rakers.
2a.
2b.
Fishes of the Western North Atlantic 399
Greatest thickness of body exceeding half its depth; head length greater
than depth of body, 24-26 °/,, and its depth 18-22 °/, of SL, the depth at
vertical from crossgroove at occiput 15.5—17.5 °/, of SL; pectoral failing to
reach base of pelvic by a distance somewhat shorter than snout and eye,
leaving 6 or 7 rows of scales exposed between its tip and base of pelvic, its
length 16-18.5 °/, of SL. pinnula T. H. Bean 1912, below.
Greatest thickness of body generally less than half its depth; head length
about equal to greatest depth of body, 22.3-27.2, and its depth 21-26.9 °/,
of SL, the depth at vertical from crossgroove at occiput 17.6—19.7 °/, of SL;
pectoral failing to reach base of pelvic by a distance generally equal to length
of snout and eye, leaving 6—8 rows of scales exposed between its tip and base
of pelvic, its length 15-17.6 °/) of SL.
anchovia (Cuvier and Valenciennes) 1847, p. 401.
1b. Distance from margin of snout to origin of dorsal notably greater than 33 °/, of
TL, 44-47.5 °/) of SL; eye equal to or longer than snout (rarely slightly shorter),
its diameter 6.9—7.9 °/) of SL; snout 7.3-8.15; gill rakers about 110-130 on
lower limb in specimens 70 mm and upward (fewer in young); pectoral fin failing
to reach base of pelvic by a distance about equal to diameter of eye.
brasiliensis (Steindachner) 1879, p. 407.
Sardinella pinnula 'T. H. Bean 1912
Bermuda Anchovy
Figure 98
Study Material. A total of 41 specimens with caudal fins broken, 74-135 mm SL,
including five specimens of type material, 93-135 mm TL, 74-114 mm SL; no holo-
type designated; all specimens from Bermuda.
Distinctive Characters. This species is very close to the species herein recognized
as anchovia, from which it differs in the more elongated and more robust body and
especially in the lower head.
Description. Proportional dimensions in per cent of standard length, and counts,
based on 24 study specimens within the length ranges given above.
Body: depth 18-22. ' Maxillary: length 9.2-10.5.
Caudal peduncle: depth 6.4—-8.2. Anal fin: length of base 13-16.
Head: length 24-26; depth 15.5- Pelvic fin: length 9.2-10.7.
ge Pectoral fin: length 16-18.5.
Snout: length 7.0—7.6. Distance from snout to origin of: dorsal
Eye: diameter 5.9—7.0. 41-42.5.
Interorbital: width 4.2—5.2.
400 Memoir Sears Foundation for Marine Research
Scales: ca. 43-47, often lost from pre- Fin rays: dorsal 17-19; anal 15-18;
served specimens. pectoral 16 or 17.
Ventral scutes: 32-34. Vertebrae: 46-47 (3 specimens).
Bopy with greatest thickness exceeding half of depth, its greatest depth 4.5—5.5
in SL; ventral outline notably more strongly convex than dorsal outline. CaupaL
PEDUNCLE Slender, its depth 3.0—3.75 in head.
Scates lost in most specimens, apparently not different from those of S. an-
chovia; 6 or 7 rows of scales exposed between tip of pectoral and base of pelvic. VEn-
TRAL SCUTES not strong, 18 or 19 in advance of pelvics and 14 or 15 behind them.
Ficure 98. Sardinella pinnula, about 124 mm TL (caudal damaged), 100 mm SL, Bermuda, USNM 21252.
Drawn by Ann S. Green.
Heap small, slender, 3.85-—4.15 in SL, its depth at vertical from crossgroove at
occiput 5.0-6.1 in SL. Snour moderately long, 3.2—3.4 in head. Eye small, its di-
ameter definitely shorter than snout, 3.55—-4.15 in head. INTERORBITAL 4.7—5.9. CHEEK
longer than deep. Maxrtiary rather broadly rounded posteriorly, scarcely reaching
vertical from anterior margin of pupil, 2.3-2.6 in head. ManprBLe projecting mod-
erately, its margin within mouth rising rather gradually, without a definite angle. Grit
RAKERS long, slender, close-set, difficult to count, those at angle about as long as eye,
the serrations on inner edge of rakers minute; the rakers increasing in number with
age: about 75 on lower limb of first arch in specimens about 85 mm long, about go in
those about 100 mm SL (gill rakers removed for counting). TgeTH as in S. anchovia.
Dorsat fin rather high anteriorly, its margin definitely concave, its longest ray
generally reaching to or beyond tip of last ray if deflexed, its origin rather nearer to
margin of snout than to vertical from base of last anal ray, its distance from margin of
snout 2.3-2.4 in SL; the base with a very narrow sheath of scales. Caupat forked;
broken in all specimens at hand. Anat fin much lower than dorsal, with a slight lobe
anteriorly, its last two enlarged rays nearly as long as the longest ones in anterior lobe
of fin, its origin about equidistant between insertion of pelvic and base of caudal,
its base 1.§5—1.9 in head. Petvic fin reaching a little more than a third of the way
Fishes of the Western North Atlantic 401
to origin of anal, inserted about equidistant between base of pectoral and origin of
anal, 2.4—2.75 in head; the axillary process reaching about to beginning of distal third
of fin, Pecrorat fin rather short, not falcate, its length 5.4—6.25 in SL, 1.35-1.55
in head, failing to reach base of pelvic by a distance generally equal to snout and about
half of eye.
Color. Specimens at hand badly faded; apparently not different from S. anchovia.
Described as “‘steel blue above, silvery below, iridescent. Head, in spirits with golden
tints; no opercular spot” (g: 122).
Size. This, the Anchovy of Bermuda, reaches a length of 6 or 7 inches (II: 35).
The average length of those observed by Goode was about 5 inches (42: 69).
Habits. Like the others of the genus, this little sardine schools, usually in the
offshore waters, and tends to come inshore in large schools “about sunrise or sunset”
(4s Re)
Commercial Importance. This little Sardine was sold “in quantities along the quay”
in March 1872, presumably for human consumption (42: 69). Barbour said that in
1903 it was seined regularly for bait in Hamilton Harbor and Flatts Inlet (4: 112).
Although it was reported as “uncommon close in shore but very abundant in mid-water
in greater depths” (ZI: 35), it was not stated that it is either eaten by man or used as
bait. A small fish occurring in the abundance indicated must be of considerable value
as forage for larger predatory foodfishes.
Range. Known only from Bermuda.
Synonyms and References:
Sardinella anchovia Goode (not of Cuvier and Valenciennes), Bull. U.S. nat. Mus., 5, 1876: 69 (Bermuda,
average size); Barbour, Bull. Mus. comp. Zool. Harv., 46 (7), 1905: 112 (distr.; in part not pinzula;
Hamilton Harbor, Bermuda); Beebe and Tee-Van (in part not of C. and V.), Field Bk. Shore Fish.
Bermuda, 1933: 35 (field char., diagn., distr.); Beebe and Tee-Van (not of C. and V.), Zoologica, N. Y.,
T3 (7), 1933: 135 (8. pivmula synon. with 8. anchovia); Hollister (not of C. and V.), Zoologica,
N. Y., 2Z (4), 1936: 186 (caudal skel.).
Sardinella pinnula T.H. Bean, Proc. biol. Soc. Wash., 25, 1912: 122 (orig.descr.; type local. Bermuda;
types USNM 74086; local name).
Sardinella anchovia (Cuvier and Valenciennes) 1847
Spanish Sardine, Sardina de Espafia, False Sardine, Sardina, Bang
Figures 99, 100
Study Material. Many specimens, from Woods Hole, Massachusetts; Blue Point
Cove, Long Island, New York; Beaufort, North Carolina; and Tortugas, Florida.
Several additional specimens from Cuba, Jamaica, and Rio de Janeiro, Brazil, apparently
belong to this species also.
Distinctive Characters: See Relationship and Variation.
Description. Proportional dimensions in per cent of standard length, based on 32
specimens, 90-155 mm TL, 70-127 mm SL.
26
4.02 Memoir Sears Foundation for Marine Research
Body: depth 21-26.9. Pectoral fin: length 15—17.6.
Caudal peduncle: depth 7.7-10. Distance from snout to origin of: dorsal fin
Head: length 22.3-27.2; depth 17.6— 40-43.5.
19.7.
Snout: length 6.0—7.5. Scales: about 41-46, often lost from pre-
Eye: diameter 5.3—7.3. served specimens.
Interorbital: width 4.2—5.3. Ventral scutes: 32-34.
Maxillary: length 8.75-11. Fin rays: dorsal 17-19, occasionally 20;
Anal fin: length of base 13.7—16. anal 16-18; pectoral 15 or 16.
Pelvic fin: length 8.5—-11. ‘ertebrae: 45-47 (7 specimens).
AACA SGA OR
ROR) V 9
AO YA ORC XX AR
IBAA NIRIDRN ON ONAN)
ROMAN ari iii Sa
Oise
YY
Ficure 99. Sardinella anchovia, about 120 mm TL, 95 mm SL, Woods Hole, Massachusetts, USNM 90233.
Drawn by Ann S. Green.
Bopy rather strongly compressed, its greatest thickness generally less than half of
its depth, its greatest depth 3.7-4.75 in SL; ventral outline notably more strongly
convex than dorsal outline. CaupaL pepuNcLE slender, moderately compressed, its
depth 2.5—3.6 in head.
Scares moderately adherent, those from middle of side below anterior rays of
dorsal scarcely deeper than long, with 3 or 4 more or less complete vertical grooves,
the margin of these scales scarcely indented but scales on back in front of dorsal fin
definitely indented; 6—8 rows of scales exposed between tip of pectoral and base of
pelvic fin. The pair of modified scales at nape small, shorter than eye. VENTRAL
scuTeEs small, 18 or 1g in front of pelvic fins and 14 or 15 behind them.
Heap 3.65—4.35 in SL, its depth at vertical from crossgroove at occiput 4.95—5.I.
SNouT 3.0—3.9 in head, its margin without a median notch. Eye shorter than snout,
3.4—4.5 in head. InreRorsitat bone (over middle of eye) 4.3-6.0. Curek longer than
deep. Maxittary rounded posteriorly, reaching to or a little beyond vertical from
anterior margin of pupil, 2.3-2.7 in head. Manprste projecting only slightly, its
margin without mouth rising rather gradually without a definite angle. GILL RAKERs
long, slender, close-set, those at angle about as long as eye, the serrations on inner edge
minute, the rakers increasing in number with age, difficult to count; about 55—60 on
Fishes of the Western North Atlantic 403
lower limb of first arch in specimens 37-44 mm, about 70-75 in examples 70-80 mm,
and 85-100 in specimens 105-125 mm SIL (gill rakers removed from arch for
counting). Tern absent on upper jaw; some fine points on margin of maxillary,
lower jaw with a few small teeth anteriorly on each side of a median interruption; a
few minute teeth on palatines, pterygoids, and tongue.
Dorsat fin only moderately elevated anteriorly, its margin somewhat concave, its
longest ray usually failing to reach the tip of last one if deflexed, its origin about equi-
distant between margin of snout and vertical from middle of base of anal, its distance
from the margin of snout 2.3-3.5 in SL; its base with a very narrow sheath composed
Ficure 100. Sardinella anchovia. ux¥tT, anal fin showing last two large rays; r1GcHT, left-hand rim of pectoral
girdle (cleithrum), showing two lobes underneath free edge of gill cover; both from specimen in Fig. 99. Drawn
by Ann S. Green.
of a single row of elongated scales, the posterior ones with a free lower margin. CauDAL
fin rather deeply forked, the lower lobe slightly the longer, without scales except for a
few modified ones on its base. Anat fin much lower than dorsal, its last two rays notably
enlarged and scarcely shorter than the longest ones in anterior lobe of fin, its origin
about equidistant between the insertion of pelvic fin and base of caudal, its base
1.45—1.8 in head; a sheath of scales at base somewhat broader than that of dorsal.
Petvic fin with margin scarcely convex when spread, reaching a little more than
halfway to origin of anal, its shortest ray about 75 °/, of the length of longest ray,
the fin inserted about equidistant between base of pectoral and origin of anal, 2.1-3.0
in head; the axillary process reaching to or beyond midlength of fin. Pecrorat fin
scarcely falcate, failing to reach base of pelvic fin by a distance nearly equal to, or quite
equal to, length of snout and eye, its length from base of upper ray to tip of fin
5.7-6.6 in SL, 1.35-1.6 in head.
Color. Bluish black above, changing rather abruptly to bright silvery at level of
upper margin of eye. Head and especially snout with many dusky punctulations; dorsal
and caudal rays with numerous dark dots, these especially crowded on distal parts of
longest rays of dorsal, giving the lobe a blackish appearance when folded; the other fins
plain, translucent, except for dark dots on 2 or 3 of the uppermost rays of the pectoral.
26*
4.04 Memoir Sears Foundation for Marine Research
Live fish have been described as very dark blue above and silvery on the side and
below (87: 7).
Size. The largest specimen at hand, with damaged caudal fin, is about 163 mm
(6.5 in.) TL and 130 mm SL. However, a length of eight inches has been reported,
though the usual length, judging from the examples at hand, probably is about four
or five inches.
Development and Growth. The smallest specimens at hand are 26 mm SL (32 mm
TL), taken at Tortugas, Florida, and 31-44 mm SL, from Beaufort, North Carolina,
taken on June 30, 1913. These young, as is usual among Clupeidae, are notably more
slender than their adults; the depth in the smallest ones (26-31 mm SL) is con-
tained 5.1-6.2 times in SL; at about 60mm SL the fish have attained near adult
proportions.
As already noted (p. 402), the gill rakers increase with age and growth: about
50—60 on the lower limb of specimens 37-44 mm, 70-75 at 70-80 mm, and 85-100
at 105-125 mm SL. The smallest reported for Tortugas, up to 32 mm length, had
only “22 or 23 gill rakers” (87: 8), and the smallest now at hand, 26 mm SL, has
about 36 gill rakers on the lower limb. These data, then, indicate that the greatest
proportional increase in the number of rakers takes place in the young under about
70mm SL.
Spawning. Although Longley did not give the collection dates for the young
26-mm fish referred to above and for young “‘up to 32 mm in length” reported for
Tortugas (87: 8), it is known that he worked at Tortugas only from about the last half
of June to the early part of September. As such small young probably had not strayed
far from the place where they were hatched, it may be assumed that at least some spawn-
ing takes place there during the summer.
Habits. These fish, like most of the herrings, run more or less in schools. In
the fall of the year at Woods Hole, Massachusetts, the fish occur irregularly and in
varying degrees of abundance (120: 91). Since these warm-water fish are not per-
manent residents there and since only adults seem to have been taken, it is surmised
that they may migrate northward in the Gulf Stream. The fish in the vicinity of Tortugas,
at least, seem to be most common some distance offshore where the water is 10-20
fms. deep, although they do come inshore (87: 7). Similar habits have been reported
for the Bermuda sardine (II: 35).
Food. Numerous copepods were found in the alimentary canal of specimens taken
at Woods Hole (85: 438). The numerous close-set gill rakers provide an effective
straining apparatus for extracting minute organisms from the water. Therefore, an
almost exclusively planktonic diet is assumed.
Enemies. Examples of this species were reported among the refuse on the tern
rookery, Bird Key, Tortugas, Florida (87: 8). Other water birds and many predatory
fish no doubt also feed on this sardine.
Parasites. The trematode, Distomum appendiculatum, was reported for the intestinal
tract of specimens from Woods Hole, Massachusetts (85: 438), and the parasitic
Fishes of the Western North Atlantic 405
copepod, Bomolochus eminens Wilson, was found on the gills of specimens taken at Tor-
tugas (134: 370).
Relationship. Longley discovered that the type material of S. anchovia, from Rio
de Janeiro, Brazil, in Paris, consists of two species that are separable by the number
of gill rakers on the lower limb of the first arch (87: 7); he reported that one ex-
ample, 116mm SL, has 105 rakers, while the other four, 114-148 mm SL, have
160-178. Although Longley’s counts are probably not entirely accurate, the difference
may be accepted as valid. Since the specimen with the smallest number of rakers was
mentioned first in the original description, Longley accepted it as the type of S. an-
chovia; this specimen was taken by M. Delalande, the other four by M. d’Orbigny and
M. Gay; Longley assigned the other four specimens to brasiliensis (p. 409) because
they had many more rakers and a more strongly compressed body than the above-
mentioned type of amchovia, specimens of approximately equal length having been
considered.
The number of gill rakers given for this “type” by Longley (105) corresponds
approximately to the number counted in the specimens herein described (85-100),
and since Longley gave no other character that disagrees with these specimens, anchovia
has been accepted as available for this species, but with considerable doubt. This doubt
results from an apparent absence among many Brazilian specimens of fish that have as
low a gill raker count as the northern specimens of anchovia herein described. Indeed,
there are slender specimens from Brazil with fewer rakers than in drasiliensis, as herein
understood, but the raker counts and the relative pectoral fin length are somewhat
intermediate between those for anchovia of the eastern United States coast and the
braziliensis from Trinidad and elsewhere. Therefore, they may be at least subspecifically
distinct from eastern United States specimens described here. If they should prove to
be specifically distinct, then the South American species would stand as auchovia while
Poey’s name, pseudo-hispanica, may be available for the northern species. However,
this cannot be definitely determined from the material at hand. In fact, specimens from
Cuba, presumably identified as pseudoharengus by Poey, seem to belong to brasiliensis
rather than to anchovia.
In the northern part of the range, S. anchovia, most frequently listed in the past
as Clupanodon pseudohispanicus, has frequently been confused with Clupea harengus (the
common Atlantic herring). This has led to some confusion, as pointed out by Longley
(87: 6) and others. The two species, indeed, resemble each other superficially in having
approximately an equally elongated body, but axchovia is easily distinguishable from
C. harengus by the more anterior position of its dorsal fin, absence of vomerine teeth,
fewer ventral scutes, fewer vertebrae, a smaller eye, and the enlargement of the last
two anal rays. These differences are all shown in detail in the descriptions.
S. anchovia of the American coast probably was incorrectly synonomized with S.
aurita of Europe and elsewhere by Regan in his revision of the genus (106: 378). This
tentative conclusion was reached by Longley after he had examined the type material
of both species in Paris (see above). From Longley’s data it appears that aurita grows
406 Memoir Sears Foundation for Marine Research
larger than any American representative of the genus and that it is somewhat inter-
mediate between anchovia and brasiliensis (as herein understood) in the number of gill
rakers. Based upon two specimens of aurita from the Bay of Naples, Italy, the body
is more slender than that of anchovia (comparing specimens of equal size), the pec-
toral fin is somewhat shorter, and the ventral scutes are slightly more numerous, 35
and 37. However, the exact relationship must remain undetermined until more ade-
quate material becomes available for comparison.
S. anchovia and the species herein designated brasiliensis are represented by close
relatives in Cuba and Jamaica even though other specimens, such as those from Woods
Hole, Massachusetts, and from Port-of-Spain, Trinidad, differ rather prominently.
However, the material from Cuba and Jamaica is too meager and too indifferently
preserved to determine the exact relationship (see Re/ationship, pp. 405, 409).
Variation. The increase in body depth and the number of gill rakers have already
been noted (pp. 398, 399); examples of equal size also vary to a considerable degree in
these respects. There is variation also in the position of the dorsal and pelvic fins, rela-
tive size of eye, shape of dorsal, and length of pectoral (see Description, p. 401).
Commercial Importance. These fish are not common enough regularly north of the
Florida Keys to be of commercial importance, but at Tortugas, Florida, particularly
offshore in 10-20 fms., they are common (87: 7, 8). Longley reported that terns fed
upon them. Undoubtedly they are also preyed upon by predacious foodfishes. They
were observed in the market in Havana, Cuba, in February and March 1912, by
Nichols, who said: ‘Abundant in the Havana market’’ (98: 180).
Range. Woods Hole, Massachusetts, to the Florida Keys, and probably south-
ward to southern Brazil. The specimens from the Gulf of Mexico coast seem to be
brasiliensis. Apparently they do not occur regularly north of Florida. In the vicinity of
Woods Hole, where they appear only during autumn (124: 741), they were abundant
in the fall of 1892 at least. It has been said that they are generally common in the
West Indies region.
Synonyms and References:
Sardinella anchovia Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 184.7: 269 (orig. descr.; type local. Rio de
Janeiro, Brazil); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 429 (descr., range);
Meek and Hildebrand, Field Mus. Publ., Zool., 75 (1), 1923: 183 (synon., descr., range); Breder,
Field Bk. Mar. Fish. Atl. Cst., 1929: 67, fig. (abund., food, size); Nichols, N. Y. Acad. Sci., To (2), 1929:
201 (distr., diagn., remarks, habits); Longley and Hildebrand, Publ. Carneg. Instn. Wash., 535, 1941: 7
(synon., relation. with Clupea harengus, 8. brasiliensis, and §. aurita, descr., ‘Tortugas, Florida, range).
Clupea anchovia Ginther, Cat. Fish. Brit. Mus., 7, 1868: 421 (diagn., range).
Clupea pseudohispanicus Kendall and Smith (in part, C/upea harengus), Bull. U.S. Fish Comm. (1894), 74, 1895:
17 (distr., descr., probably two or more species; cf. specimens from diff. locals.).
Clupanadon pseudohispanicus Jordan and Evermann (in part C/upea harengus), Bull. U.S. nat. Mus., 47 (1),
1896: 423 (descr., after Kendall and Smith; relation. to European species); Smith, Bull. U.S. Fish
Comm. (1897), 77, 1898: 91 (Woods Hole, Massachusetts; usually found with young C/upea harengus);
Linton, Bull. U.S. Fish Comm. (1899), 19, 1901: 438 (food, parasites, Woods Hole, Mass.); Evermann
and Marsh, Bull. U.S. Fish Comm. (1900), 20 (1), 1902: 84 (descr., distr., Puerto Rican names, size,
synon.); Wilson, Proc. U.S. nat. Mus., 39, 1911: 370 (parasitic copepod, Bommolophus eminens, on gills);
Nichols, Bull. Amer. Mus. nat. Hist., 32 (18), 1912: 180 (abund. Havana, Cuba, market); Sumner,
Fishes of the Western North Atlantic 407
Osburn, and Cole, Bull. U.S. Bur. Fish. (1911), 3% (2), 1913: 741 (refs., Woods Hole, Mass., food,
parasites); Meek and Hildebrand, Field Mus. Publ., Zool., r5 (1), 1923: 180 (synon., descr., distr.) ;
Beebe and Tee-Van (in part 8. brasiliensis), Zoologica, N. Y., 10 (1), 1928: 39 (refs., field char., discus.).
Sardinella aurita Regan (in part 8. brasiliensis), Ann. Mag. nat. Hist., (8) 79, 1917: 378 (synon., in part not
this species; descr. in part, range); Jordan (in part not of C. and V.), Manual Vert. Anim. NE U.S.,
ed. 13, 1929: 41 (diagn., range); Jordan, Evermann, and Clark (in part not of C. and V.), Rep. U.S.
Comm. Fish. (1928), 2, 1930: 43 (range, synon.).
Sardinia anchovia Jordan, Evermann, and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 43 (range, synon.).
Sardinia pseudo-hispanica Howell, Bull. Mus. comp. Zool. Harv., 82 (2), 1938: 171 (8. pseudo-hispanica Poey
to synon. of 8. azchovia C. and V.; cotypes).
Doubtful References:
Sardinia pseudo-hispanica Poey, Memorias, Cuba, 2, 1860: 311 (orig. descr.; type local. Cuba; cotypes Nos.
17768 and 17771; local name).
Clupea pseudohispanica Giinther, Cat. Fish Brit. Mus., 7, 1868: 442 (descr. after Poey).
Sardinella brasiliensis (Steindachner) 1879
Sardinia, Sardinia de Espafia, Spanish Sardine, Sardinha Verdadeira
Figure 101
Study Material. Numerous specimens, 82-129 mm TL, 64-98 mm SL, from
Port-of-Spain, Trinidad. Also specimens for comparison, from Island of Marguerita,
Venezuela; Cuba; Jamaica; Palmetto Key, and Pensacola, Florida; and many from Rio
de Janeiro, Brazil, not used in preparing the description, since they are extralimital
and may prove to be at least subspecifically distinct; all in the USNM collections.
Distinctive Characters. See Relationship.
Description. Proportional dimensions in per cent of standard length, and counts,
based principally on study specimens, 64-98 mm SL.
Body: depth 22.4-26.9. Pectoral fin: length 17.3-19.5.
Caudal peduncle: depth 7.5-9.5. Distance from snout to origin of: dorsal
Head: length 25.3-28.7; depth 18.722. 44-47.5.
Snout: length 7.3-8.15.
Eye: diameter 6.9—7.9. Scales: 42-47.
Interorbital: width 4.9-5.7. Ventral scutes: 32-34.
Maxillary: length 10.7—-11.7. Fin rays: dorsal 17 or 18; anal 18-20;
Anal fin: length of base 15.1-16.5. pectoral 15 or 16.
Pelvic fin: length 9.9-11.2. Vertebrae: 46 (4 specimens).
Bopy slender, strongly compressed, its greatest thickness about half of its depth,
its greatest depth generally at dorsal origin, 3.7-4.45 in SL; ventral outline much
more strongly convex than dorsal outline. CaupAL PEDUNCLE strongly compressed,
its depth 2.9-3.75 in head.
Scares only moderately adherent; those on middle of side below base of dorsal
scarcely deeper than long, with 3 or 4 partly incomplete vertical grooves, the margin
408 Memoir Sears Foundation for Marine Research
membranous and scarcely indented, but those on back in front of dorsal fin with defi-
nitely indented margin, and no vertical grooves; 4 or 5 rows of scales exposed be-
tween tip of pectoral and base of pelvic. The pair of modified scales at nape shorter
than eye. VENTRAL sCUTES not prominent, 18 or 19 in front of pelvic fins and 14 or
1§ behind them.
Heap with length rather exceeding greatest depth of body, 3.5—3.95 in SL, its
depth at vertical from slight crossgroove at occiput 4.55—5.35 in SL. Snour rounded,
its margin without a median notch, 3.4—3.8 in head. Eye with diameter equal to, or a
Te
DOO SARA ASRS RRR
NNR!
aMauuint SANNA XN yee
OY NS uae
0 Se
Ficure 101. Sardinella brasiliensis, 122mm TL, 94mm SL, Port-of-Spain, Trinidad, USNM 124317.
Drawn by Ann S. Green.
little longer than, snout (rarely slightly shorter), 3.4—3.8 in head. INTERoRBITAL (over
middle of eye) 4.8—5.8. CuzeKk longer than deep. Maxrtiary broadly rounded pos-
teriorly, reaching nearly or quite to vertical from anterior margin of pupil, 2.3-2.5
in head. Manprste projecting slightly, its margin within mouth rising rather grad-
ually without forming a definite angle. Gitt Rakers long, very slender, extremely
close-set, very difficult to count, the longest about as long as eye, the serrations on inner
edge extremely minute; the rakers increasing in number with age and growth: 82 and
83 in specimens 64 and 68 mm SL, 116 in an 89-mm specimen, and 130 and 132 in
specimens g1 and 93mm SL (gill rakers removed for counting). TEETH absent in
upper jaw; some fine points present on margin of maxillary; lower jaw with a row
of 5—7 very small teeth anteriorly on each side of a median interruption; very small
granular teeth present on palatines, pterygoids, and tongue.
Dorsat fin moderately elevated anteriorly, its margin definitely concave, its long-
est rays usually failing to reach the tip of last ray if deflexed, its origin a little nearer
to margin of snout than to vertical from last ray of anal, its distance from margin of
snout 2.15—2.25 in SL; the base of fin with a very narrow sheath composed of a single
row of elongated scales, the posterior ones having a free lower margin. Caupat fin rather
deeply forked, the lower lobe slightly the longer; no small scales, but a few large modi-
Fishes of the Western North Atlantic 4.09
fied ones on base of fin (Fig. 101). Anat much lower than dorsal, its last two rays
considerably enlarged and scarcely shorter than the longest ones in anterior lobe of fin,
its origin generally a little nearer to base of caudal than to base of pelvic, its base
1.7—1.85 in head; a moderately broad sheath of scales at base. Petvic fin with margin
slightly convex when spread, reaching notably less than halfway to origin of anal, its
shortest ray about 75 °/) of the length of the longest one, the fin inserted equidistant
between base of pectoral and origin of anal, or a little nearer to the former, 2.5—2.8 in
head; the axillary process reaching to or beyond midlength of fin. Pecrorat fin slightly
falcate, generally failing to reach base of pelvic fin by a distance nearly equal to di-
ameter of eye, its length from base to tip of longest ray 5.1-5.8 in SL, 1.4—1.55 in head.
Color. Bluish black above, changing abruptly to bright silvery at level of upper
margin of eye. Region of shoulder, upper surface of head, snout, and mandible with
dusky punctulations. Dorsal and caudal more or less dusky, the lobe of the dorsal dusky
when folded, the dusky appearance being caused by dark dots on the rays (missing on
inter-radial membranes); the other fins plain, translucent, except for dark dots on one
or two uppermost rays of pectoral. Peritoneum black.
Size. The largest example included in the collections examined is 180 mm
(7.2 in.) TL, 145 mm SL. However, examples 230 mm (9.2 in.) long have been
reported (25: 189).
Development. No young definitely identified as this species are included in the col-
lections examined, the smallest being 82 mm TL, 64 mm SL. Nevertheless, it is evi-
dent that the remarks on the development of young S. axchovia (p. 404) also apply to
this species, as the smallest examples are the slenderest and have the fewest gill rakers
on the lower limb.
Food. The food of this species has not been listed. However, the very numerous
close-set gill rakers and the stomach with heavy walls and numerous pyloric coeca
suggest a diet consisting of plankton.
Relationship. This species has been here designated as brasiliensis principally be-
cause of Longley’s discovery that the type material of anchovia, in Paris, consisted of
two species (87: 7), as stated in the account of anchovia (p. 405). Accordingly, Longley
assigned the specimens with the more numerous gill rakers to brasiliensis, which may
be correct, but positive proof is lacking, as the original description of brasiliensis gives
neither the number of rakers nor any other diagnostic character as herein understood.
The specimens (or species) with the greater number of rakers were compared with
one identified as “Clupea brasiliensis Steindachner,” also from Rio de Janeiro, Brazil;
they were found to be identical with it.
S. brasiliensis is close to anchovia according to a limited number of rather indif-
ferently preserved specimens, from Cuba and Jamaica. However, specimens from Port-
of-Spain, Trinidad, and from Venezuela are quite distinct from those of our Atlantic
coast, the most outstanding difference being the greater number of gill rakers in brasi/-
iensis; other differences include the position of the dorsal and pelvic fins, the size of
the eye, and the length of maxillary and pectoral fin.
410 Memoir Sears Foundation for Marine Research
The numerous specimens from Rio de Janeiro, Brazil, examined and measured in
part, differ somewhat from those taken at Port-of-Spain, Trinidad, which serve chiefly
as the basis for this description. In fact, the former are less distantly removed from
specimens of azchovia from the eastern United States than from those of Trinidad, as
the eye runs a little smaller (5.75—7.25 °/, of SL), the maxillary is rather shorter (9.5—
11 °/, of SL), the pectoral is a little shorter (16-18 °/) of SL), and the gill rakers are
somewhat less numerous (108-117 in specimens 97—109 mm SL). Therefore, a sub-
specific difference is suggested. Indeed, it seems highly probable that further study
will show some intergradation among all species recognized in these pages.
S. brasiliensis, like S. anchovia, apparently was incorrectly synonomized with
S. aurita of Europe and elsewhere by Regan (p. 405). According to data based on
specimens from Messina, Sicily, its type locality, aurita grows larger than brasiliensis
and has fewer gill rakers in examples of equal size. These species may therefore be
regarded as valid unless further evidence shows this opinion to be incorrect.
Commercial Importance. This sardine is of at least some commercial importance at
Port-of-Spain, Trinidad, where Waldo L. Schmitt of the U.S. National Museum ob-
tained from the market (April 18, 1939) the fine series of specimens used in preparing
the foregoing description. It is abundant and occurs also in large numbers in the market
at Rio de Janeiro, Brazil, but southward in the state of Sao Paulo it is less numerous
(25: 188).%
Range. The west coast of Florida, the West Indies, and Venezuela to southern
Brazil. Not reported for México to Colombia.
Synonyms and References:
Clupea brasiliensis Steindachner, S. B. Akad. Wiss. Wien, So, 1879; Ichthyol. Beitr., 8: 64 (orig. descr.;49 type
local. Rio de Janeiro, Brazil).
Sardinella aurita Regan (in part), Ann. Mag. nat. Hist., (8) 19, 1917: 378 (synon., in part achovia; descr.
in part; range); Campos, Arqu. Zool. estad. Sao Paulo (1941), 3(7), 1942: 188, fig. 2 (synon., descr.,
abund. in market, range, specimens coll. in Brazil); Oliveira (not of Cuvier and Valenciennes), Min.
Agr. Dep. Nac. Pro. Anim., Div. Cacga e Pesca, 1942: 22 (ref., range).
Clupanodon pseudohispanicus Beebe and Tee-Van (in part axchovia), Zoologica, N.Y., 10 (1), 1928: 39 (refs.,
field char., discus.).
Sardinella brasiliensis Longley in Longley and Hildebrand, Publ. Carneg. Instn. Wash., 535, 1941: 7 (cf.
anchovia and aurita).
Doubtful References:
Clupanodon pseudohispanicus Evermann and Kendall, Rep. U.S. Comm. Fish. (1899), 1900: 55 (Pensacola
and Snapper Banks, Florida).
48. The Brazilian Sardinha, or Sardinha Verdadeira (true sardine), is the most important market fish in the area between
Victoria, Espirito Santo, and Santos, Sao Paulo. It is not taken commonly north of this area, and it is to be questioned
that this fish is identical with the populations farther to the north.—G.S.M.
49. Fowler has shown that Clupea brasiliensis Steindachner 1879 is a primary homonym of Clupea brasiliensis Schneider
180r (equals Albula vulpes). Fowler uses the specific name allecia (of Rafinesque), but the latter applies to European
fish, which Fowler, following Regan, would apparently unite with the American form. If the American form is
distinct it would appear to be nameless, unless changes in the International Rules of Zoological Nomenclature,
now said to be imminent, would restore the availability of Steindachner’s name. Until this genus is adequately
revised for the entire Atlantic and the relationships of the American and European populations are worked out,
it would seem premature to disturb the nomenclature.—G. S. M.
Fishes of the Western North Atlantic 411
Sardinella anchovia Schreiner and Miranda-Ribeiro, Arch. Mus. nac., Rio de J., 12, 1903: 91 (Rio de Janeiro,
range, names); Metzelaar (probably not of Cuvier and Valenciennes), Rapp. Vissch. Curagao, ed. by
Boeke, 1, W. Indian Fishes, 1919: 10, fig. 2 (Dutch West Indies: fig. seems based on érasiliensis as
herein understood).
Sardinella aurita von \hering (not of Cuvier and Valenciennes), Rey. Industr. Anim., 1 (3), 1930: 228
(synon., fin rays, scales, ventral scutes, Brazil).
Genus Rhinosardinia Eigenmann 1912
Rhinosardinia Eigenmann, Mem. Carneg. Mus., 5, 1912: 4453 type species by original designation, RAinosardinia
serrata Eigenmann. Myers, Copeia, 1929: 1; Heringia preoccupied in Diptera.
Generic Synonym:
Heringia Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 2073; type species by original designation, C/upea
amazonica Steindachner; name preoccupied in Diptera.
Characters. Bopy moderately elongate, strongly compressed; ventral outline more
strongly convex than dorsal outline. Bony scuTEs on chest and abdomen well devel-
oped, the posterior one ending in a rather long sharp spine. Scares adherent, each
with a prominent vertical groove having more or less horizontal laterals (not verified
in bahiensis). Moutu strongly oblique. Manpise projecting. Maxitiary broadly
rounded posteriorly, with a strong sharp backward projecting spine near its anterior
end (Fig. 103); maxillary-premaxillary margin continuous, composed of a soft mem-
branous section; entire margin toothless. TEETH absent on vomer, but present on
mandible (at least in serrata and bahiensis), palatines, pterygoids, and tongue. GILL
RAKERS rather numerous (about 30-40 on lower limb of first arch in serrata), close-
set. Dorsat fin about over middle of body, with approximately 14-17 rays. ANAL
fin scarcely longer than dorsal and placed well behind it, with about 15-18 rays.
Pertvic well developed, with 8 rays. Pecrorat rather small, more or less equal to
head in length.
Remarks. This genus differs from all others of this family in the presence of a
small sharp spine on the anterior part of the maxillary, close behind the premaxillary.
It differs further from related genera in the small anal fin, which is situated well behind
the dorsal fin.
Range. This genus of small herrings is confined to northeastern and northern
South America, from Rio de Janeiro to Venezuela, in fresh, brackish, and salt water.
Key to Species
1a. Sides with a prominent silvery lateral band, about as wide as eye; origin of dorsal
about over insertion of pelvic fin and nearly an eye’s diameter nearer to tip of
snout than to base of caudal; teeth present on mandible, maxillary, and premaxil-
lary; dorsal with 17 rays; anal with 18; scales 44. dahiensis (Steindachner) 1879.
Coast of Brazil from Bahia (Baia) to
Rio de Janeiro.
412 Memoir Sears Foundation for Marine Research
1b. Sides without silvery band; origin of dorsal definitely behind insertion of pelvic
fin and about equidistant between margin of snout and base of caudal; teeth miss-
ing on maxillary and premaxillary; dorsal with 14 or 15 rays; anal with 15-17
rays; scales 35-38.
2a. Greatest depth of body 3.5-4.0 in SL (75- and 40-mm specimens, re-
spectively); depth of caudal peduncle close to half of greatest depth of body;
head 4.0 in SL; snout 4.0 in head; base of dorsal fin about as long as its
longest rays. amazonica (Steindachner) 1879, below.
2b. Greatest depth of body 3.1-3.4 (60—68-mm specimens) in SL; depth of cau-
dal peduncle definitely less than half of greatest depth of body; head 4.25~-5.1
in SL; snout 4.6—5.5 in head; base of dorsal fin notably shorter than its longest
rays. serrata Eigenmann 1912, p. 413.
Rhinosardinia amazonica (Steindachner) 1879
Study Material. None.
Distinctive Characters. This species is closely related to serrata. However,
amazonica differs from serrata, according to the original description, in the some-
what more slender body, slightly larger head, longer snout, the proportionately deeper
caudal peduncle, in the absence of teeth on the mandible, and in the somewhat dif-
ferent proportion of the longest dorsal rays with respect to the head length and
dorsal base.
Description. Proportional dimensions in per cent of standard length, and counts,
modified after the original description by Steindachner (see Synonyms and Ref-
erences).
Body: depth 27.5 in 75-mm examples, Scales: 37.
about 25 in those 40 mm long. Ventral scutes: 27.
Head: length 25. Fin rays: dorsal 15; anal 16.
Bopy strongly compressed, its ventral outline notably more strongly convex than
dorsal outline, its greatest depth 3.5 in SL in large examples, 4.0 in smaller ones. Cau-
DAL PEDUNCLE deep in comparison with depth of body, close to half of greatest depth.
Sca.es adherent; about 37 in a horizontal row between margin of opercle and base
of caudal; a vertical groove dividing the free (exposed) part of scale from the covered
part, 2 or 3 sharp grooves on the former. VENTRAL scuTeEs on chest and abdomen, 15
in front of pelvic fins and 12 behind them.
Heap 4.0 in SL. Snour 4.0 in head. Eye 3.0. Moutu small. Maxriiary reaching
little beyond anterior margin of eye. Manp1BLe projecting little. TzerH absent on pre-
maxillary, mandible, and vomer.
Dorsat fin with base about as long as longest rays, its origin a little behind vertical
at insertion of pelvic, and equidistant between end of head (snout) and base of caudal.
Fishes of the Western North Atlantic 413
Caupat about 1.5 times length of head, with slender pointed lobes covered with scales
anteriorly.
Color. Body silvery; head yellowish white.
Size. The largest example reported was 75 mm long.
Development. Small specimens (40 mm) are more slender than larger ones (75 mm),
according to the original description, and the ventral outline is strikingly less convex
in the smaller ones. However, the proportionate depth of the caudal peduncle and the
greatest depth of the body are about the same in both small and large specimens.
Range. Known only from Para, Brazil.
Synonyms and References:
Clupea amazonica Steindachner, S. B. Akad. Wiss. Wien, 80, 1879; Ichthyol. Beitr., 8: 65 (orig. descr., type
local. “Amazon R. at Parad”); von Ihering, Dic. Anam. Brasil, 1940: 716 (habitat).
Rhinosardinia amazonica Myers (in part not of Steindachner), Copeia, 1929: 1 (R. serrata thought to bea synon.,
following Regan, which seems to be incorrect).
Sardinella (Rhinosardinia) amazonica von \hering, Rev. Indust. Anim., 1 (3), 1930: 229 (synon., diagn., re-
lation.).
Negative Reference:
Heringia amazonica Regan (not of Steindachner), Ann. Mag. nat. Hist., (8) 9, 1917: 394 (descr. based on
cotypes of R. serrata; synon. apparently incorrect).
Rhinosardinia serrata Eigenmann 1912
Figures 102, 103
Study Material. Five cotypes, 60--68 mm TL, 46-49 mm SL, from Morawhanna,
British Guiana, USNM 66284.
Distinctive Characters. See Relationship.
Description. Proportional dimensions in per cent of standard length, and counts,
based on the five cotypes, 46-49 mm SL.
Body: depth 29.5—32. Pelvic fin: length 15.5—-17.
Caudal peduncle: depth 1.55-1.95. Pectoral fin: length 19.2—20.5.
Head: length 19.5-23.5; depth 20.5- Distance from snout to origin of: dorsal
nie 47-51.
Snout: length 4.1-4.55.
Eye: diameter 6.2—-6.75. Scales: 35-38.
Interorbital: width 4.25-4.8. Ventral scutes: 27-29.
Maxillary: length 9.1—10. Fin rays: dorsal 14 or 15; anal 15-17;
Mandible: length 8.4—9.1. pectoral 11 or 12.
Anal fin: length of base 12.7—14.3. Vertebrae: 29 (105: 394).
Bopy rather deep but not excessively compressed, its greatest thickness about a
fourth of its depth, its greatest depth 3.1-3.4 in SL; dorsal outline gently convex
414 Memoir Sears Foundation for Marine Research
anteriorly, the ventral outline somewhat more strongly convex. CauDAL PEDUNCLE
scarcely as long as deep, its depth definitely less than half of greatest depth of body,
1.55—-1.95 in head.
Scars closely adherent, thin, almost circular, with even margin and with a
prominent, nearly complete vertical groove provided with 1-3 horizontal branches on
Py
ee a
s < i XS) Bas ee ——
\ ae 0
Ns a
\ ass
Ficure 102. Rhinosardinia serrata, 63mm TL, 47 mm SL, Morawhanna, British Guiana, cotype, USNM
66284. Drawn by Ann S. Green.
each side; scales extending well out on caudal fin. VENTRAL scuTeEs well developed,
especially the posterior ones, each with a prominent spine, 16 in advance of pelvic fins
and 11-13 behind them.
Heap rather short and deep, its length 4.25—5.1 in SL, its depth at vertical from
crossgroove at occiput 4.65-4.9. Snour definitely
shorter than eye, 4.6—5.5 in head. Eve 3.0-3.8. In-
TERORBITAL moderately narrow, 4.1—5.25 in head.
Cueek longer than deep. Mourn strongly oblique.
Maxittary broadly rounded posteriorly, reaching
nearly or quite below anterior margin of pupil, 2.1-
2.5 in head. ManpisLe projecting moderately, its
margin within mouth rising strongly, forming a
broadly rounded obtuse angle, 2.15—2.6 in head.
Git RAKERS slender, close-set, very difficult to count,
approximately half as long as eye, about 35-40 on
lower limb. Tzrrx all minute, present on mandible
anteriorly and on palatines, pterygoids, and tongue,
but absent on maxillary, premaxillary, and vomer.
Ficure 103. Rhinosardinia serrata head Dorsat fin well developed, rather high anteri-
showng spine near end of maxillary, : : : :
orly, its margin concave, its longest rays exceeding
from specimen in Fig. 102. Drawn by :
Ann S. Green. the length of base, about as long as head, reaching far
Eee
Fishes of the Western North Atlantic 415
beyond tip of last ray if deflexed, its origin slightly nearer to margin of snout than
to base of caudal, its distance from margin of snout 1.95—2.1 in SL, its base nearly
as long as that of the anal; a narrow sheath of scales on anterior part of base. CauDaAL
fin long, rather deeply forked, the upper lobe exceeding greatest depth of body by
more than diameter of eye. Anat fin small, its origin about half of length of head
behind base of dorsal and a little nearer to base of caudal than to insertion of pelvic,
its base notably shorter than head, 6.9—7.8 in SL; a narrow sheath of scales at base.
Petvic fin long, pointed, inserted about half of eye’s diameter in advance of dorsal
fin and about equidistant between base of pectoral and vent, 1.2-1.5 in head. Pec-
ToRAL failing to reach base of pelvic by diameter of eye, scarcely as long as head,
4.9—5.0 in SL.
Color. Brownish in alcohol, with metallic reflections. Sides of head silvery. Back,
upper surface of head, snout, and mandible with dusky dots; dusky dots also on the
fins, exclusive of pelvics.
Size. The largest specimens reported to date are only 85 mm (3.4 in.) TL.
Relationship. This species was synonymized with R. amazonica by Regan in his
revision of this and related genera (106: 394). A careful rereading of the original de-
scription of amazonica, with syntypes of serrata in hand, reveals some differences that
seem too important to dismiss. Accordingly, the species herein are recognized as distinct.
For a definite determination of the relationship of the two species, a more accurate de-
scription of specimens from the type locality of amazonica is needed.
Range. Known from Venezuela and from Dutch and British Guiana.
Synonyms and References:
Heringia amazonica Fowler (probably not of Steindachner, but serrata Eigenmann), Proc. Acad. nat. Sci. Philad.,
63, 1911: 207 (14 specimens from “‘Surinam’’; made type of a new genus, Heringia; preoccupied in
Diptera); Regan (not of Steindachner), Ann. Mag. nat. Hist., (8) 19, 1917: 394 (Rhinosardinia serrata
Eigenmann apparently incorrectly synon.; descr. based on cotypes of R. serrata).
Rhinosardinia serrata Eigenmann, Mem. Carneg. Mus., 5, 1912: 445, text fig. 39, pl. 62, figs. 3 and 4 (orig.
descr., type local. Morawhanna and Mora Passage, British Guiana; type CM 2443).
Rhinosardinia amaxonica Myers (in part not of Steindachner), Copeia, 1929: 1 (R. serrata thought to be synon.,
following Regan, which seems to be incorrect); Fowler (probably not of Steindachner, but serrata
Eigenmann), Proc. Acad. nat. Sci. Philad., 83, 1931: 406 (name and locals. only; Cafio Guanoco and
Punta Tigre at mouth of San Juan R., Venezuela).
Genus Jlisha Gray 1846
Ilisha Gray in Richardson, Ichthyol. Seas of China, Japan, Rep. Brit. Ass. Adv. Sci. (1845), 1846: 306; type
species by monotypy, I/isha abnormalis Gray equals A/osa elongata Bennett; Myers, Copeia, 1950: 63;
notes on identity with Neosteus and Pseudochirocentrodon.
Generic Synonyms:
Platygaster Swainson, Nat. Hist. Fish. Amphib. Rept., 2, 1839: 294; type species C/upea africana Bloch, first
designated by Jordan, Genera Fishes, 1919: 203; name preoccupied in bees.
Pellona Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 300; type species Pe//ona orbignyana Cuvier
and Valenciennes, first designated by Gill, Proc. Acad. nat. Sci. Philad., 1861: 37; but P. ordignyana
416 Memor Sears Foundation for Marine Research
equals Pristigaster flavipinnis Valenciennes (in d’Orbigny), Voyage Amer. Merid., Poiss., 5 (2), 1847:
8; Atlas, d’Orbigny, 1839: pl. 10, fig. 2.
Zunasia Jordan and Metz, Mem. Carneg. Mus., 6, 1913: 7; type species by monotypy, Pristigaster chinensis
Basilewsky equals 4/osa elongata Bennett.
Pseudochirocentrodon Miranda-Ribeiro, Publ. Comm. Linhas Telegr. Estrat. Matto Grosso Amazonas, 58, 1923
(‘1920’): 8; type species by monotypy, P. amazonicum Miranda-Ribeiro.
Neosteus Norman, Ann. Mag. nat. Hist., (9) IZ, 17, 1923: type species Pe//ona ditchela Valenciennes by sub-
sequent designation of Norman, Zool. Rec., Pisces, for 1923: 25.
Characters. Bopy elongate, rather strongly compressed, its ventral outline more
strongly convex than dorsal outline. Bony scures strong, about 25—37 along sharp edge
on chest and abdomen. Scatzs small to rather large, often missing on preserved
specimens, generally with several irregular vertical grooves. SNour with or without
a definite median notch. Mouru moderate, strongly oblique. Manpis.ez projecting
strongly. Maxitiary generally reaching under anterior half of eye, with a bone or liga-
ment between it and premaxillary. TEETH all small, present in a single series on anterior
part of mandible, on premaxillaries, maxillaries, and usually in bands on palatines,
pterygoids, and tongue, but none on vomer. GILL RAKERs strong, serrate, about 12-25
on lower limb of first arch (in American species). Dorsat fin with about 15-20 rays
(in American species). ANAL fin long, its origin under posterior rays of dorsal or just
behind that fin, with about 36—52 rays (in American species). Petvic fin small (occa-
sionally absent in one species), with 6 or 7 rays (in American species).
Remarks. This genus is characterized by the elongated, compressed body with a
strongly convex ventral outline, by the very long anal fin, and by the small pelvic fin,
inserted in advance of the dorsal fin and consisting of only 6 or 7 rays. Furthermore,
the premaxillary and maxillary are not adjoined, a soft ligament (in J/isha) or a bone
(in Neosteus) being present between them.
Range. Tropical parts of the Atlantic, Indian, and Pacific oceans. Fresh waters of
the Amazon Basin. One species, /. xarragansetae, reported from Rhode Island.
Key to Species of the Western Atlantic and
Tributary Rivers
1a. Ventral scutes 20-25 before and 10-14 behind pelvic fins, total number 32-37.
2a. Gill rakers only 12 or 13 on lower limb; scales 72—75 in a lateral series; depth
of body less than a third (27.5—30 °/,) of SL.
altamazonica Cope 1872, p. 417.
2b. Gill rakers 23-31 on lower limb; scales fewer than 70 in a lateral series; depth
of body about equal to a third (30-37 °/,) of SL.
3a. Dorsal fin with 19 or 20 rays; gill rakers on lower limb 23-26; axillary
process of pectoral reaching middle of first ray.
castelnaeana (Cuvier and Valenciennes) 1847, p. 419.
50. See Myers (97: 63) for discussion.
Fishes of the Western North Atlantic 417
3b. Dorsal fin with 17 or 18 rays; gill rakers on lower limb 30 or 31; axillary
process of pectoral not reaching middle of fin.
flavipinnis Valenciennes 1839.
Buenos Aires, Argentina.
1b, Ventral scutes 19-25 before and only 6 or 7 behind pelvic fins, total number
25-32.
4a. Depth of body less than a third (30 °/,) of SL; scales about 60 in a lateral
series; gill rakers 19 on lower limb; anal with 47—52 rays.
amazonica (Miranda-Ribeiro) 1923, p. 421.
4b. Depth of body about equal to or greater than a third (33-40 °/,) of SL;
scales 33-45 in a lateral series; gill rakers 22-25 on lower limb.
sa. Depth of body greater than a third (37-40 °/)) of SL; ventral scutes
19-21 before and 6 behind pelvic fins; pectoral with 13-15 rays; anal
with 36-42 rays. harroweri Fowler 1917, p. 423.
5b. Depth of body about equal to a third (33 °/,) of SL; ventral scutes 25
before and 7 behind pelvic fins; pectoral with 16 rays; anal with 45 rays.
narragansetae Fowler 1911, p. 426.
Llisha altamazonica (Cope) 1872
Apapa
Figure 104
Study Material. Three specimens, 280, 305, and 360 mm TL, 220, 234, and
275mm SL, from “the Amazon River region, Para to Mandos,” Brazil, USNM
52548.
Distinctive Characters. This species is characterized by the slender body and small
scales. It differs especially from related species in the very few gill rakers on the lower
limb of the first arch.
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens, 220-275 mm SL.
Body: depth 27.5-30. Pelvic fin: length 7.5—8.1.
Caudal peduncle: depth 9.4-9.5. Pectoral fin: length 20.5—22.7.
Head: length 25.5-27; depth 21.5- Distance from snout to origin of: dorsal
igh. 50-52.
Snout: length 5.15—5.5
Eye: diameter 6.2—6.8. Scales: 72-75.
Interorbital: width 12-13.5. Ventral scutes: 32 or 33.
Maxillary: length 12.8-13.5. Fin rays: dorsal 18; anal 37—40; pectoral
Mandible: length 14-15. 16 or 17.
Anal fin: length of base 29-31.
418 Memoir Sears Foundation for Marine Research
- Bopy quite slender, rather strongly compressed, its greatest thickness about 2.5
times in its depth, its greatest depth 3.3-4.65 in SL; the dorsal outline gently convex,
the ventral outline much more strongly convex. CAUDAL PEDUNCLE rather strong, its
depth 2.7—2.8 in head.
Sca.es adherent; those on middle of side below anterior rays of dorsal definitely
deeper than long, without vertical grooves, closely imbricated; the exposed part rather
more than two times deeper than long: the membranous margin only slightly irregular.
VENTRAL scuTEs moderately developed, 22 or 23 in advance of pelvic fins and 10 or
11 behind them.
Ficure 104. L/isha altamazonica, 360 mm TL, 275 mm SL, Amazon River somewhere between Pard and
Mandos, Brazil, USNM 52548. Drawn by Ann S. Green.
Heap 3.7—3.9 in SL, its depth at vertical from slight crossgroove at occiput 4.2 5—
4.65. SnoutT only a little shorter than eye, with a prominent median notch, 2.0-2.05
in head. Eye with much adipose tissue, 3.9-4.3. INTERORBITAL 2.0—2.15 in head.
Cuerek deeper than long. Maxitiary rather narrowly rounded posteriorly, reaching
below middle of eye, 2.0-2.05 in head; a narrow bone with teeth between premaxillary
and maxillary. MAnpIBLE projecting strongly, its tip scarcely below dorsal outline of
head, its margin within mouth rising gently, 1.8—1.85 in head. Git RAKERs at angle
of first arch about 75 °/) of length of eye, only 12 or 13 on lower limb. TzETH small,
in a single row, on anterior part of mandible and on premaxillaries; minute teeth on
margin of maxillary, and still smaller teeth on the bone between premaxillary and
maxillary; bands of minute granular teeth on palatines, pterygoids, and tongue.
Dorsat fin rather high anteriorly, its longest rays reaching far beyond the tip of
last ray if deflexed, its origin equidistant between margin of snout and base of caudal,
or a little nearer to snout, its distance from margin of snout 1.9—2.0 in SL; a sheath
composed of a single row of scales at its base. Caupat fin well forked, the lower lobe
much the longer; small scales extending onto the fin, covering most of it. Anat fin long,
with a definite lobe anteriorly, its origin immediately behind a vertical from last dorsal
ray, only a little nearer to margin of opercle than to base of caudal, its base 3.2—3.4
Fishes of the Western North Atlantic 419
in SL; a sheath composed for the most part of two rows of scales at its base. Petvic fin
small, inserted fully an eye’s diameter in advance of vertical from origin of dorsal and
a little nearer to base of pectoral than to origin of anal, 3.15-3.45 in head; free axillary
process about 66 °/, of the length of fin. Pecrorat fin reaching well beyond base of
pelvic, its length 4.4—4.9 in SL, 1.15—1.3 in head; a free axillary process present along
about 66 °/, of length of fin.
Color. Old preserved specimens grayish brown above, silvery below, with indefi-
nite alternating light and dark streaks along the rows of scales. A slight shoulder spot
present. Interorbital, upper surface of snout, and tip of mandible dark brown. Dorsal,
caudal, and pectoral fins largely dusky, the lower lobe of caudal with a dark area on its
middle rays, this area narrow anteriorly but spreading posteriorly.
Range. Originally described from the Ambyiacu River, a tributary emptying into
the Amazon near Pebas, Peru; also reported from Iquitos, Peru, and the Tocantins
River, Brazil. The three specimens now at hand are from the lower Amazon.
Synonyms and References:
Pellona altamazonica Cope, Proc. Acad. nat. Sci. Philad., 1872: 265 (orig. descr.; type local. Ambyiacu R.,
trib. of upper Amazon).
Ilisha altamazonica Eigenmann, Rep. Princeton Exped. Patagonia, 1896-1899, 3 (4), 1910: 452 (ref., range);
Eigenmann and Allen, Fishes West. S. Amer., 1942: 333 (refs., range); Myers, Copeia, 1950: 64
(synon.).
Neosteus altamazonica Norman, Ann. Mag. nat. Hist., (9) TZ, 1923: 19 (ref., descr.).
Ilisha (Neosteus) altamazonica von Ihering, Rev. Indust. Anim., 1 (3), 1930: 230 (fin rays; note “Amazonas,
rio Tocantins”’).
Ilisha desuratus Nakashima, Bol. Mus. Hist. Nat., “Javier Prado,” 5 (16), 1941: 77, fig. (orig. descr.; type
local., environs of Iquitos, Peruvian Amazon).
Negative Reference:
Llisha altamazonica de Menezes (not of Cope). Publ. Serv. Piscic., Dep. Nac., Ceard, Brazil, 113, 1946: 1
(notes on common name; local. of coll., ident., etc. Later publication [113A, 1947: 1] refers this to
synon. of J. caste/naeana).
Ilisha castelnaeana (Cuvier and Valenciennes) 1847
Cagona, Apapa, Sardinha Grande, Pirapema, Gros Hareng
Study Material. None.
Distinctive Characters. See Relationship.
Description. Proportional dimensions in per cent of standard length, and counts,
after Norman (99: 19), “three specimens 280-390 mm in length,”’ presumably TL.
Body: depth 33.3-37. Scales: 63-66.
Head: length 26.5-27.5. Ventral scutes: 34-37.
Anal fin: length of base 28-29. Fin rays: dorsal 19 or 20; anal 40 or 41.
ring
420 Memoir Sears Foundation for Marine Research
Bopy with greatest depth 3.25—3.33 in SL; the ventral outline more strongly
convex than dorsal outline. Scaes relatively small. VENTRAL scuTES 21—23 in advance of
pelvics and 13 or 14 behind them. Snour a little shorter than eye. Eye 3.8—4.0 in
head. Maxitiary extending to or a little behind middle of eye. ManpiBLe projecting
strongly. GILL RAKERS 23-26 on lower limb of first arch. Dorsat origin nearer to
margin of snout than to base of caudal. Anat with origin behind dorsal base, its base
3-5—3-6 in SL. Petvic longer than eye, inserted much nearer to base of pectoral than
to origin of anal. Pecrorat with axillary process reaching to or beyond middle of first
ray of fin.
Color. According to the original account, greenish above, silvery ventrally. Lower
lobe of caudal with a large dark spot; other fins yellowish.
Size. See Description.
Relationship. This species seems to be closely related to J. flavipinnis, a species
apparently reported only from Buenos Aires, Argentina, with which caste/naeana has
sometimes been considered identical. According to Norman, the latest revisor of this
and related genera, flavipinnis has more gill rakers on the lower limb of the first
arch (30 or 31) and an axillary process of the pectoral fin that is shorter, not reaching
the middle of the first ray (99: 17). These differences seem to be substantiated by two
specimens (USNM 77357) about 220 and 230 mm TL (caudal damaged), 173 and
187 mm SL, from Buenos Aires. Furthermore, the dorsal and anal of flavipinnis seem
to have fewer rays (D. 17 and 18; A. 38 and 39) and a shorter anal fin base (3.45 and
3.55 in SL). In color the specimens of flavipinnis differ from castelnaeana, as described,
in the absence of a dark spot on the lower lobe of the caudal.
Range. “Coast of Surinam and Brazil; Upper Amazons” (Norman).
Synonyms and References:
Pellona castelnaeana Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 306 (orig. descr.; type local., around
mouth of Amazon R.); Fowler, Arqu. Zool. estad. Sao Paulo, 3 (6), 1941: 134 (refs.).
Pellona favipinnis Giinther (not of Valenciennes), Cat. Fish. Brit. Mus., 7, 1868: 454 (descr. based on speci-
mens from Surinam, where species does not occur); Boulenger (not of Valenciennes), Ann. Mag. nat.
Hist., (6) 20, 1897: 298 (Marajé I., Brazil); Goeldi (not of Valenciennes), Bol. Mus. Paraense Hist.
Nat., 2 (3), 1898: 486 (refs., names); Puyo (not of Valenciennes), Bull. Soc. Hist. nat. Toulouse, 70,
1936: 166 (refs., descr., habits, habitat, commerc. import.).
Ilisha flavipinnis Jordan and Evermann (not of Valenciennes), Bull. U.S. nat. Mus., 47 (1), 1896: 435 (descr.
after Giinther, 1868); Eigenmann (in part not of Valenciennes), Rep. Princeton Exped. Patagonia,
1896-1899, 3(4), 1910: 452 (JL. caste/naeana incorrectly synon., range, includ. both species); Eigenmann
(not of Valenciennes), Mem. Carneg. Mus., 5, 1912: 446 (descr., Georgetown, British Guiana, where
species does not occur); Starks (not of Valenciennes), Stanf. Univ. Publ., Univ. Ser., 1913: 8 (Pard,
Brazil, where species apparently does not occur).
Neosteus castelnaeana Norman, Ann. Mag. nat. Hist., (9) IZ, 1923: 19 (synon., descr., range).
Ilishe (Neosteus) castelnaeana von |hering, Rev. Industr. Anim., 1 (3), 1930: 230 (refs., fin rays; ‘““Amazonias
e Guyana; entra no rias”).
Ilisha altamazonica de Menezes (not of Cope), Publ. Serv. Piscic. Dep. Nac., Ceard, Brazil, 113, 1946: 1
(notes on common name; local. of coll., ident., etc.).
Ilisha castelnaeana de Menezes, Publ. Serv. Piscic. Dep. Nac., Ceard, Brazil, 113A, 1947: 1 (J. a/teamaxonica
of de Menezes, 1946, to synon. of this species).
Fishes of the Western North Atlantic 421
Ilisha amazonica (Miranda-Ribeiro) 1923
Apapa, Sardina, Asuna Nahui
Figure 105
Study Material. Type of J. apapae, about 200 mm TL (caudal damaged), 160 mm
SL; exact place of capture unknown, being registered merely as having been collected
in “the Amazon River region, Par4 to Mandos,” Brazil; USNM 52550.
Distinctive Characters. This species differs from the other local species in having a
Ficure 105. Ilisha amaxonica, about 200mm TL (caudal damaged), 160 mm SL, Amazon River, Brazil,
type of J. apapae, USNM 52550. Drawn by Ann S. Green.
ligament between the premaxillary and maxillary; the others have a narrow bone
bearing fine teeth along the margin. Its body is slender as in a/tamazonica, but its scales
are apparently somewhat larger, its dorsal rays fewer, and its anal rays more numerous.
It has more gill rakers than a/tamazonica but fewer than the other species herein de-
scribed. It is nearest to furthii of the Pacific coast of tropical America, which also
has a ligament between the premaxillary and maxillary; but it differs from that species
in having fewer ventral scutes (34-37 in furthii), and in the position of the pelvic
fin—inserted notably nearer to the origin of anal than to the base of pectoral in ama-
zonica but equidistant between these points in furthii; also they differ in several
other respects.
Description. Proportional dimensions in per cent of standard length, and counts,
based on Study Material, specimen 160 mm SL.
Body: depth 30.5. Eye: diameter 6.8.
Caudal peduncle: depth 8.75. Interorbital: width 2.5.
Head: length 25; depth 23.5. Maxillary: length 13.2.
Snout: length 5.6. Mandible: length 14.5.
422 Memoir Sears Foundation for Marine Research
Anal fin: length of base 42. Scales: ca. 60, mostly missing.*
Pelvic fin: length 10.5. Fin rays: dorsal 15; anal 4735 pectoral
Pectoral fin: length 23. 14.
Distance from snout to origin of: dorsal 44. Ventral scutes: 26."
Bopy strongly compressed, its greatest thickness scarcely a third of its depth, its
greatest depth 3.25 in SL; the dorsal outline anterior to dorsal nearly straight, the
ventral outline strongly convex. CAUDAL PEDUNCLE 2.85 in head.
Scatzs mostly lost; the scales on middle of side below base of dorsal scarcely
deeper than long, with 4 or 5 vertical striae, only the posterior one complete; not very
closely imbricated; the exposed part of scale notably greater than half of its depth, the
margin nearly smooth. Venrrat scutes well developed, 20 in advance of pelvic fins
and 6 behind them.
Heap 4.0 in SL, its depth at vertical from slight crossgroove at occiput 4.2.
Snout definitely shorter than eye, without a definite median notch, 4.45 in head.
Eye 2.9. InTERORBITAL 10 in head. CueeK fully as deep as long. Maxittary narrowly
rounded posteriorly, reaching below anterior margin of pupil, 1.85 in head. A soft
ligament between premaxillary and maxillary. Manprsve projecting strongly, its tip
scarcely below dorsal outline of head, its margin within mouth rising rather strongly,
forming an obtuse angle, 1.75 in head. Grit RaKeRs at angle of first arch scarcely half
of length of eye, 19 on lower limb of first arch. TrrrH small; a few in a single row
on anterior part of mandible, a series on premaxillary and on margin of maxillary,
and bands of granular teeth on palatines, pterygoids, and tongue.
Dorsat fin high anteriorly, its longest rays reaching far beyond tip of last ray if
deflexed, only a little shorter than the head, its origin nearer to margin of snout than to
base of caudal by a distance equal to length of snout and eye, its distance from margin
of snout 2.3 in SL. Caupat forked (damaged), the lower lobe evidently the larger; small
scales covering most of fin. Anat fin long, scarcely elevated anteriorly, its anterior rays
not forming a definite lobe, its origin under last ray of dorsal and equidistant between
posterior margin of eye and base of caudal, its base 2.4 in SL; a scaly sheath present.
Petvic fin long (for an J/isha), inserted rather less than an eye’s diameter in advance of
vertical from origin of dorsal and nearer to origin of anal than to base of pectoral, 2.3
in head; a short broad axillary process present. Pecrorat fin reaching beyond base of
pelvic by a distance fully equal to diameter of pupil, 4.3 in SL, 1.05 in head; the
short free axillary process only a little more than a third of the length of fin.
Color. Old preserved specimen grayish above, yellowish to silvery below. Upper
surface of snout and tip of mandible dark brown; a brownish area behind eye. All fins
with dusky punctulations, these few and scattered on pelvic, most numerous on dorsal,
caudal, and upper half of pectoral.
Range. Known only from the Amazon.
sr. If Myers (97: 63-64) is correct in referring I. apapae and I. iguitensis to the synonymy of this species, then the anal
rays vary from 47 (types of amaxonica) to 52 (type of iquitensis), the scales from 57 to about 60, and the total number
of ventral scutes 25 or 26.—G. S. M.
Fishes of the Western North Atlantic 4.23
Synonyms and References:
Pseudochirocentrodon amazonicum Miranda-Ribeiro, Publ. Comm. Linhas Telegr. Estrat. Amazonas, 58, 1923
(“1920”): 8 (orig. descr., type local. Mandos, Estado de Amazonas, Brazil).
Ilisha iquitensis Nakashima, Bol. Mus. Hist. Nat. “Javier Prado,” 5 (16), 1941: 66, fig. (orig. descr.; type
local. environs of Iquitos, Peruvian Amazon).
Ilisha apapae Hildebrand, Smithson. misc. Coll., 70 (9), 1948: 3, fig. 2 (orig. descr.; type local. Rio Amazonas
somewhere between Pard and Mandos, Brazil; type USNM 52550).
Ilisha amazonica Myers, Copeia, 1950: 63 (synon., types of P. amaxonicum and I. apapae).
Ilisha harroweri Fowler 1917
Caille (Trinidad)
Figure 106
Study Material. Twenty-two specimens, 45-155 mm TL, 34-122 mm SL, from
Atlantic coast of Panama (20 specimens) and Port-of-Spain, Trinidad (2). One from
Barro Santos and another from Porto Inhuana, southern Brazil, were compared and
probably belong to this species.
Distinctive Characters. See Relationship.
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens, 34-122 mm SL.
Body: depth 36.5—40. Pectoral fin: length 18-21.5.
Caudal peduncle: depth 9.0-10.4. Distance from snout to origin of: dorsal
Head: length 28-30; depth 28.5—31.5. 47-51.
Snout: length 6.0—7.5.
Eye: diameter 9.75-11.8. Scales: ca. 33-37, usually lost in pre-
Interorbital: width 3.6—4.1. served specimens.
Maxillary: length 14.3-16. Ventral scutes: 25-27.
Mandible: length 15.5-17. Fin rays: dorsal 15 or 16 (rarely 17);
Anal fin: length of base 37-43. anal 36-42; pectoral 13-15.
Pelvic fin: length 7.4-9.0. Vertebrae: 39 or 40 (5 specimens).
Bopy deep, strongly compressed, its greatest thickness definitely less than a third
of its depth, its greatest depth 2.5—2.75 in SL; the dorsal outline only slightly convex,
the ventral outline strongly convex. CauDAL PEDUNCLE strongly compressed, its depth
2.8—3.2 in head.
Scaes mostly lost in preserved specimens; those from middle of side below an-
terior rays of dorsal definitely deeper than long, with about 3-7 vertical grooves, all
except the posterior one interrupted, the margin of these scales irregularly indented.
VENTRAL SCUTES strong, 19-21 (usually 20) in advance of pelvic fins and 6 be-
hind them.
HEaD 3.33—3-55 in SL, its depth at vertical from crossgroove at occiput 3.15—3.5.
Snout shorter than eye, with a slight median notch, 4.15—-5.15 in head. Eve 2.5 5—2.8
424 Memoir Sears Foundation for Marine Research
in head. INTERORBITAL 6.5—8.35. CHEEK fully as long as deep. Maxiiiary rather nar-
rowly rounded posteriorly, generally reaching nearly below middle of eye, 1.8—2.0 in
head; a narrow bone with teeth between premaxillary and maxillary. Manp1BLe pro-
jecting strongly, its tip scarcely below dorsal outline of head, its margin within mouth
rising strongly, 1.7—1.85 in head. Git rakers at angle of first arch only about as long
as pupil, apparently not increasing in number with age, 23-25 on lower limb. TEeTu
small, in a single series on mandible and premaxillary; a row of minute teeth on margin
of maxillary; present in bands on palatines, pterygoids, and tongue.
Ficure 106. Ilisha harroweri, 160mm TL, 123 mm SL, Colén, Panama, USNM 81749. Drawn by Ann
S. Green.
Dorsat fin moderately elevated anteriorly, its longest rays reaching far beyond
tip of last ray if deflexed, its origin a little nearer to margin of snout than to base of
caudal, its distance from margin of snout 1.7—2.1 in SL; a sheath composed of one row
of scales at base. Caupat fin forked, the lower lobe slightly the longer; small scales
extending onto base of fin. Anat fin long, its anterior rays somewhat elevated but
not forming a definite lobe, its margin nearly straight, its origin somewhat variable,
generally under posterior rays of dorsal, usually about equidistant between posterior
margin of eye and base of caudal, its base 2.3-2.7 in SL; the sheath composed of two
rows of scales anteriorly and one row posteriorly. Pre.vic fin small, rarely missing,
reaching halfway or more to origin of anal, inserted under, or more usually somewhat
in advance of, vertical from origin of dorsal, much nearer to origin of anal than to base
of pectoral, 3.1-3.8 in head; no free axillary process. Pecrorat fin moderately
developed, not falcate, reaching base of pelvic in some but failing to reach it in others,
its length 4.7—5.5 in SL, 1.3-1.65 in head; a free axillary process reaching about to
midlength of longest rays.
Color. Bluish gray above. Sides silvery. Upper surface of anterior part of
snout and mandible dusky. Dorsal and anal fins yellowish in life, the tip of the
Fishes of the Western North Atlantic 425
dorsal lobe and generally the margin of the caudal dusky; vertical fins with dusky
punctulations.
Size. The largest example at hand is 155 mm (6.2 in.) TL and 122 mm SL. This
specimen was among the larger ones seen in the Colon, Panama, market.
Development. Nothing concerning reproduction is known, and larval and postlarval
stages have not been described. However, at hand are a few small specimens that are
proportionately quite as deep as large ones. In two specimens, 34 and 50 mm SL, the
depth equals 38.3 and 40 °/, of SL, whereas in the two largest, 115 and 122 mm SL,
the depth equals 39 and 37.6 °/,; furthermore, the ventral profile is much more strongly
convex anteriorly in the small examples. This is contrary to the usual development, as
small specimens of this family ordinarily are slenderer than large ones. A development
similar to that of Aarroweri has been noticed in Brevoortia patronus.
Relationship. It is not possible to distinguish this species from Neosteus ternetzi
as described by Norman (roo: 593) or by a comparison of two specimens from
southern Brazil (Barro Santos, USNM 87718, Porto Inhuana, USNM 100837) with
others from Colén, Panama, and Port-of-Spain, Trinidad. The following proportions
and counts, which are generally in agreement with those given in the original descrip-
tion of fernetzi (from Panama and Trinidad), are based on the two specimens from Bra-
zil, with the figures for the Barro Santos specimen first: Body depth 2.6, 2.75 in SL.
Caudal peduncle depth 2.95, 3.1. Head length 3.4, 3.5; depth at vertical from cross-
groove at occiput 3.15, 3.55. Snout length 5.0, 4.0 in head. Eye diameter 2.5, 2.7.
Interorbital width 6.5, 6.55. Maxillary length 1.85, 1.8. Mandible length 1.85, 1.75.
Anal base 2.4, 2.9. Predorsal distance 2.0, 1.95. Scales lost, about 35. Ventral scutes
18 +6, 19+7. Fin rays: dorsal 16, 16; anal 36, 39; pectoral 15, 15. It seems proper
to conclude, therefore, that sermetzi cannot be distinguished from Aarroweri specifically.
It may be subspecifically distinct, but that can be determined only when more material
becomes available for study.
Variation. The pelvic fins are well developed, but there is at hand a specimen somm
SL from Colon, Panama, in which these fins are missing. Inasmuch as no scar can be
detected, it is assumed that it never possessed the fins. It seems highly probable that
the specimen without pelvics, taken on the Atlantic coast of Panama and described as
a new species by Borodin under the name Pristigaster vanderbilti (1g: 6, pl. 1, fig. 1),
also is this species. The occasional absence of pelvic fins in this species seemingly may
be looked upon as a variation.
Range. Atlantic coast of Panama, Trinidad, and southern Brazil. Not reported as
yet from intermediate points.
Synonyms and References:
Ilisha harroweri Fowler, Proc. Acad. nat. Sci. Philad., 69, 1917: 128, fig. 1 (orig. descr.; type local. Colon,
Panama; type ANSP 46959).
Neosteus harroweri Norman, Ann. Mag. nat. Hist., (9) ZZ, 1923: 19 (descr. after Fowler).
Ilisha argentata Meek and Hildebrand, Field Mus. Publ., Zool., 5 (1), 1923: 190, pl. 9 (orig. descr.; type
local. Fox Bay, Colén, Panama; type USNM 81749).
426 Memoir Sears Foundation for Marine Research
Neosteus ternetzi Norman, Ann. Mag. Nat. Hist., (9) 12, 1923: 593 (orig. descr.; type local. Rio de Janeiro,
Brazil).
Pristigaster vanderbilti Borodin, Bull. Vanderbilt oceanogr. Mus., I (1), 1928: 6, pl. 1, fig. 1 (orig. descr.;
type local. Limon Bay, Colén, Panama; type VOM 12; apparently based on an “abnormal” specimen of
Ilisha harroweri without pelvics).
Ilishe (Neosteus) ternetzi von Vhering, Rev. Industr. Anim., 1 (3), 1930: 230 (fin rays, notes, Rio de Janeiro,
Brazil).
Ilisha narragansetae Fowler 1911
Figure 107
Study Material. None.
Distinctive Characters. This species, according to the original description, seems
to be related to harroweri, having a similar deep body and rather large scales. It seems
to differ, however, in having a greater number of ventral scutes, more numerous anal
and pectoral rays, and a longer pectoral fin.
Description. Proportional dimensions in per cent of standard length, and counts,
condensed from Fowler (38: 208), specimen about 145 mm TL (caudal damaged),
reported as taken off Newport, Rhode Island.
Body: depth 33. Scales: ca. 44.
Head: length 25.5. Ventral scutes: 32.
Fin rays: dorsal 17; anal 45; pectoral 16.
Bopy much compressed, its greatest depth at insertion of pelvic fin, 3.33 in SL;
the dorsal outline slightly convex, the ventral outline more strongly convex anteriorly.
CAUDAL PEDUNCLE compressed, its length a trifle less than its depth, 3.4 in head.
Scares large, cycloid, each with as many as 6 vertical striae. VENTRAL SCUTES
developed, 25 in advance of pelvics and 7 behind.
Heap compressed, slightly concave above, convex below, greatly inclined, 3.5 in
SL. Snour shorter than eye, a median notch scarcely developed, 4.3 in head. Eye 3.0.
InTERORBITAL 6.25. Maxittary reaching somewhat beyond a vertical from anterior
margin of pupil, 1.9 in head. Manprszz strongly projecting, its margin within mouth
well elevated. Girt raxers half of length of eye, 22 on lower limb of first arch.
Trrru small; a series on anterior part of mandible and premaxillaries, a double series
on each palatine; also present on tongue.
Dorsat fin graduated downward from the first branched ray, its origin equi-
distant between tip of mandible and base of caudal. Caupat forked (damaged); small
scales on base of fin. Anat fin low, its margin straight, its anterior rays little elevated,
its origin slightly behind base of dorsal, about equidistant between anterior margin of
eye and base of caudal. Pstvic fin very small, inserted rather less than an eye’s di-
ameter in advance of vertical from origin of dorsal and much nearer to origin of anal
than to base of pectoral. Pecrorat fin falcate, rather broad, reaching far beyond tip
of pelvic fin, 1.3 in head.
Fishes of the Western North Atlantic 427
Color. ““Brownish on back and upper surface of head behind, sides and lower re-
gions silvery white in alcohol; fins all pale brownish.”
Range. Known only from the holotype, purported to have been taken at Newport,
R.I. It is stated in the original account of this species, “it would appear undoubtedly
to have been obtained at Newport as a waif of the Gulf Stream, probably from some
tropical region in America.” This remark dates back to 1911 and no other example like
it has been reported since then. The genus, in fact, is not known from the West Indies
(except Trinidad), nor from any other locality north of the Atlantic coast of Panama.
Ses
Sane
sa
as
fe
Ficure 107. I/isha narragansetae, about 165 mm TL (caudal damaged). Modified by Ann S. Green after
Fowler, type, ANSP 15314.
Synonyms and References:
Ilisha narragansetae Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911: 208, fig. 1 (orig. descr.; type local. New-
port, Rhode Island; type ANSP 15314; cf. Pelloma bleekeriana, Pristigaster flavipinnis, and Pellona
castelnaeana); Fowler, Proc. Boston Soc. nat. Hist., 35 (4), 1917: 112.
Neosteus narragansetae Norman, Ann. Mag. nat. Hist., (9) IZ, 1923: 20 (descr. after Fowler); Jordan, Manual
Vert. Anim. NE U.S., ed. 13, 1929: 42 (diagn.).
Genus Pristigaster Cuvier 1817%?
Pristigaster Cuvier, Régne Anim., ed. 1, 2, 1817: 176; and 4: pl. 10, fig. 33 type species by monotypy,
figured but no specific name given; this same figure repeated in ed. 2 of Régne Anim., in which name
cayanus is provided; generic type by monotypy, Pristigaster sp. Cuvier 1817 equals P.cayanus Cuvier
1829.
Characters. Bopy strongly compressed; the dorsal outline straight to slightly con-
vex; the chest and abdomen greatly dilated, forming a pronounced arc. Springs on
median line between occiput and origin of dorsal, 4 or 5. TrETH small; no canines.
Dorsat fin small, with about 13-15 rays, its origin well in advance of origin of anal.
Anat fin long, its origin under or just behind base of last dorsal ray, with about
46—55 rays. Petvic fin missing.
Remarks. The greatly dilated, rounded, half-moon-shaped outline of the chest
52. Nomenclature and synonymy for both genus and species rewritten by G. S. Myers.
428 Memoir Sears Foundation for Marine Research
and abdomen and the several short spines on the median line of the back between the
occiput and origin of dorsal sufficiently characterize this genus.
Range. Known from a single species, as herein understood, which ranges from the
Guianas to northern Brazil and ascends freshwater streams.
Pristigaster cayana Cuvier 1829
Sardinha d’Agua Doce, Apapa (Brazil)
Figure 108
Study Material. None.
Distinctive Characters. In bodily shape, cayana resembles somewhat J/isha harroweri,
but its chest and abdomen are even more strongly convex, forming a very pronounced
arc (see Fig. 108). The shape strikingly resembles that of the freshwater characin,
Gasteropelecus, but its pectoral fin is much shorter.*? The several spines on the median
Ficure 108. Pristigaster cayana, about 110 mm TL (caudal damaged). Drawn by Ann S. Green after Compos’
P. martii.
line of the back between the occiput and the dorsal fin separate it from all American
species of the family.
Description. Proportional dimensions in per cent of standard length, and counts,
from several published accounts, drawings, and a photograph.
Body: depth 50-56. Gill rakers: on lower limb of first arch,
Head: \ength 25-29. 20-23.
Fin rays: dorsal 13-15; anal 46-55;
Scales: 39-42. pectoral 11.
Ventral scutes: 30-33.
53- Moreover it does not appear to possess the enormous pectoral muscles which enable the characin to buzz its wings
during aerial flight.—G. S. Myers.
Fishes of the Western North Atlantic 429
Bopy much compressed, its ventral outline strongly convex, forming a pronounced
arc, its greatest depth over basal half of pectoral, 1.8-2.0 in SL. CaupaL PEDUNCLE
short, its depth 3.0 in head.
VENTRAL SCUTES prominent, especially the posterior ones. A median series of 5
spines between occiput and origin of dorsal.
Heap 3.65—4.0 in SL. Snour shorter than eye, 3.5 in head. Eye 2.5 in head.
InreRorBITAL half of width of eye. Mouru almost vertical. Maxittary extending
below anterior margin of eye, 2.5 in head. ManprBLe projecting, extending nearly into
dorsal outline of head. Grit raKeERs slender, about 2.25 times in diameter of eye, 20-23
on lower limb.
Dorsat fin high anteriorly, its longest rays reaching far beyond tip of last ray
if deflexed, nearly as long as head, its origin much nearer to margin of snout than to
base of caudal. Caupat fin well forked, the lobes a little shorter than head. Anat long,
its origin under or just behind last rays of dorsal, a little less than 0.5 of SL. Pecrora
a little shorter than head.
Color. After Fowler: ‘‘Color in alcohol with back pale brown and under surfaces
bright shining silvery white. Iris silvery white. Back and occiput with scattered dark
to blackish dots. Dorsal and caudal yellowish buff basally, pale gray terminally. Other
fins dull buff.”
Size. A maximum length of only 110 mm (4.4 in.) has been reported.
Range. Coast of the Guianas to Brazil, ascending streams; reported from the
Ucayali River Basin in the upper Amazon system, Peru.
Synonyms and References:
Pristigaster Cuvier, Régne Anim., ed. 1, 2, 1817: 176 and 4: pl. 10, fig. 3 (diagn. of genus and species combined ;
no specific name given; type local. “Seas of America”); Cuvier, Régne Anim., ed. 2, 3, 1830: pl. 12,
fig. 3 (no specific name on plate, but includ. in 1829 text; see under P. cayanus below).
Pristigaster cayanus Cuvier, Régne Anim., ed. 2, 2, 1829: 321 (two spp. referred to; name cayanus provided
for Amer. species figured in 1817; type local. fixed by specific name as Cayenne, French Guiana; fig. on
1830 plate of ed. 2 Régne Anim. ident. to that in ed. 1); Cuvier and Valenciennes, Hist. Nat. Poiss.,
20, 1847: 334, pl. 597 (descr., discus.); Giinther, Cat. Fish. Brit. Mus., 7, 1868: 463 (synon., descr.,
range); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 438 (descr. after Cuvier and Valen-
ciennes, synon.); Starks, Stanf. Univ. Publ., 1913: 9 (Madeira R., Brazil, 400 miles above mouth);
Norman, Ann. Mag. nat. Hist., (9) 11, 1923: 14 (synon., descr., range); Jordan, Evermann, and Clark,
Rep. U.S. Comm. Fish. (1928), 2, 1930: 45 (range, synon.); von Ihering, Rev. Indust. Anim., 1 (3),
1930: 230 (ref., diagn.); von Ihering, Dic. Anim. Brasil, 1940: 716 (habitat, notes); Eigenmann and
Allen, Fish. West. S. Amer., 1942: 333 (Peruvian Amazon).
Pristigaster martii Agassiz, in Spix and Agassiz, Genera Species Pisc. Brazil., 1829: 55, pl. 24a (orig. descr.;
type local. mouth of Amazon); Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 337 (presumed
differences from P.cayanus; rec. for the mouth of Amazon); Castelnau, Anim. Nouv. Rares Amer.
Sud, Poiss., 1855: 57 (Amazon); Fowler, Proc. Acad. nat. Sci. Philad., gr, 1940: 221, fig. 2 (Rio
Ucayali, Peruvian Amazon, synon., descr.); Campos, Arqu. Zool. estad. Sao Paulo (1941), 3 (7),
1942: 187, fig. 2 (synon., descr., geogr. distr.); Fowler, Arqu. Zool. estad. Sao Paulo, 6, 1948: 16,
fig. 6 (synon.).
Pristigaster americanus Guerin-Meneville, Iconogr. Régne Anim. Cuvier, Poiss., 1829-1844: 33, pl. 57, fig. 3
(type local. coast of S. Amer. in Atlantic).
Pristigaster phaeton Cuvier and Valenciennes, Hist. Nat. Poiss., 20, 1847: 338 (orig. descr., type local. Amazon);
430 Memoir Sears Foundation for Marine Research
Castelnau, Anim. Nouv. Rares Amer. Sud, Poiss., 1855: 57, pl. 28, fig. 3 (based on material of Valen-
ciennes).
Pristigaster cayana Myers, Copeia, 1956: 64 (nomencl., early synon. corrected).
Negative Reference:
Pristigaster martii Fowler, Bull. Amer. Mus. nat. Hist., 70 (1), 1936: 179, fig. 72 (synon. in part incorrect;
descr. after P. do//oi Boulenger, which is not this species).
Genus Odontognathus Lacépéde 1800
Odontognathus Lacépéde, Hist. Nat. Poiss., 2, 1800: 220; type species by monotypy, Odontognathus mucronatus
Lacépéde.
Generic Synonym:
Gnathobolus Bloch and Schneider, Syst. Ichthyol., 1801: 556; type species by monotypy, Odomtognathus mucro-
natus Lacépéde.
Characters. Bopy strongly compressed; the dorsal outline more or less concave
over head, the ventral outline rather strongly convex anteriorly. Bony scuTeEs on sharp
keel of chest and abdomen, the outer margin of the posterior scutes more or less
denticulate. Mourn strongly oblique to nearly vertical, with strongly projecting
lower jaw. Maxitiary greatly produced in large examples; maxillary and premaxillary
separated by a toothless membranous section, forming a continuous (uninterrupted)
margin with maxillary and premaxillary. TzeTH small to minute, present on both jaws,
palatines, pterygoids, and tongue, but wanting on vomer. GILL RAKERS not numerous,
about 19-25 on lower limb of first arch. Dorsat fin rather feebly developed, with about
10-14 rays. ANaL fin long, with about 52-80 rays, beginning far in advance of dorsal
fin, generally well in advance of midlength of SL. Pertvic fins wanting. PEcTroraL
fins generally longer than head.
Remarks. The species of this genus have greatly compressed bodies, a concavity
over the head, a membranous section between the maxillary and premaxillary that
forms a continuous margin with the two elements it separates, a very oblique to nearly
vertical mouth, and a very long anal fin. Several of the posterior ventral scutes generally
are denticulate.
Range. Known from both coasts of Panama, on the Atlantic side southward to
Guiana.
Key to Species
1a. Depth of body 4.65 in SL; anal fin with 79 or 80 rays (71-82 by earlier descrip-
tions); dorsal fin over middle of anal, its origin about twice as far from poste-
rior margin of eye as from base of caudal. = mucronatus Lacépéde 1800, p. 431.
1b. Depth of body 3.5-4.25 in SL in adults; anal fin with only 52-60 rays; dorsal
fin ending over middle of anal, its origin nearly twice as far from margin of snout
as from base of caudal. compressus Meek and Hildebrand 1923, p. 433-
Fishes of the Western North Atlantic 431
Odontognathus mucronatus Lacépéde 1800
Figure 109
Study Material. Two large specimens, 135 and 152 mm TL, 111 and 136mm SL,
from Port-of-Spain, Trinidad, in only fair condition, the fins being more or less
damaged and the color badly faded. Both specimens have the ventral scutes missing
below the pectoral fins, causing a concavity in the ventral outline; this probably is an
abnormality, even though it is present in both specimens.
Distinctive Characters. See Relationship.
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens, 111 and 136 mm SL.
EAST’ oS
Figure 109. Odontognathus mucronatus, about 152mm TL (caudal damaged), 136 mm SL, Gulf of Paria,
Trinidad, USNM 123085. Drawn by Ann S. Green.
Body: depth 20.5-21.5. Distance from snout to origin of: dorsal
Caudal peduncle: depth 5.1—5.4. 70-71.5.
Head: length 17.5-19; depth 15-17.
Snout: length 4.7. Scales: ca. 45-48 pockets, mostly lost in
Eye: diameter 5.0—5.5. preserved specimens.
Interorbital: width 1.5-1.9. Ventral scutes: ca. 21 or 22 (some may
Maxillary: length 14-14.5. be missing because of an abnor-
Mandible: length 11-12.5. mality).
Anal fin: length of base 52-56. Fin rays: dorsal 10-12; anal 79 or 80;
Pectoral fin: length 19-22. pectoral 12.
Bopy with greatest thickness about 0.25 of its depth, its greatest depth 4.6 5—4.8
in SL; the dorsal outline over head concave, the ventral outline moderately convex
anteriorly and then concave (possibly abnormal). CaupaL PEDUNCLE very short, its
depth 3.4—3.5 in head.
Scaes nearly all lost, a few remaining over base of anal, these very thin, notably
deeper than long, with nearly smooth margin and without vertical grooves. VENTRAL
scutes rather prominent, all except the last 3 or 4 ending in prominent spines, the
last ones with nearly smooth margin, missing for a short distance below base of
pectoral (p. 432), forming a slight concavity.
£22 Memor Sears Foundation for Marine Research
Heap small, its length 5.25—5.7 in SL, its depth at vertical from slight cross-
groove at occiput 5.9—6.65. SNourT a little shorter than eye, 3.75-4.05 in head. Eye
3-5—-3-55. INTERORBITAL very narrow, IO—11.5 in head. Cuzex about as long as deep.
Mourn nearly vertical. Maxittary produced into a long narrow process reaching
margin of opercle in larger one, 1.25-1.3 in head. ManpiBLe projecting strongly, its
tip a little below dorsal outline, its margin within mouth rising strongly, forming a
broadly rounded obtuse angle, 2.1-2.2 in head. Git Raxkers fairly strong, only about
half of length of eye, their inner margin rather strongly serrate, 24 or 25 on lower limb
of first arch. TrETu all small to minute, in a single series on mandible, premaxillary, and
maxillary; missing on membranous section between the last mentioned elements; minute
teeth on palatines and pterygoids, but none on vomer; an elongated patch on tongue.
Dorsat fin small, feebly developed (damaged specimens at hand), its origin not-
ably more than length of head behind origin of anal, about twice as far from the
posterior margin of eye as from base of caudal, its distance from margin of snout
1.35-1.45 in SL. Caupat fin damaged, the lower lobe evidently the longer. Anat fin
very long, its origin nearer to tip of mandible than to base of caudal by about twice
diameter of eye, its base 1.8-1.9 in SL. Pecrorat moderately large, a little longer
than head, reaching 4th or sth from last ventral scute, 4.55-5.25 in SL.
Color. Large preserved specimens pale; sides of head silvery. A pale lateral band
(no doubt silvery in life), very narrow (a mere line), anteriorly, becoming fully half as
broad as eye over posterior half of anal. Back with dusky punctulations extending for-
ward onto head and snout and onto tip of mandible; punctulations also on fins.
Relationship. Compared to O. compressus, the body of O. mucronatus is more slender
and apparently less compressed, the anal fin is longer, and the dorsal fin is more pos-
teriorly placed. And in all these respects mucronatus is nearer to the Pacific panamensis
than to compressus; however, it is even more slender than panamensis, and its gill rakers
and anal rays are more numerous. If the interruption in the series of ventral scutes
below the pectorals (p. 431) were normal, which seems doubtful, this would con-
stitute another important difference that separates mucronatus from the other species of
the genus. However, a slight concavity in the ventral outline has been noticed in a large
specimen of O. panamensis and in a much smaller specimen of O. tropicus from northern
Peru, recently referred to a new genus; however, in each instance the scutes are
fully developed in the concavity. Other specimens of the same species have no such
concavity in the ventral outline, nor do any of the numerous specimens of O. compressus
examined. Furthermore, this question of abnormality is not answered in Norman’s
revision (99: 15), wherein he gave no information other than “Ventral scutes 8 + 12,”
which is in approximate agreement with the number of scutes before and after the
interruption at the concavity on the specimens at hand. It may be significant, however,
that in the description of O. panamensis in his revision, the number of ventral scutes is
also written “13 +16’; there certainly is no interruption in the series of scutes in the
specimen of that species before me.
Range. Known from the coast of Guiana and from Trinidad (Port-of-Spain).
Fishes of the Western North Atlantic 4.33
Synonyms and References:
Odontognathus mucronatus Lacépéde, Hist. Nat. Poiss., 2, 1800: 221, pl. 7, fig. 2 (orig. descr. under “Todon-
tognathe aiguillone,” but a footnote gives the binomial Odontognathus mucronatus; type local., Cayenne,
French Guiana); Jordan and Evermann, Bull. U.S.nat. Mus., 47 (1), 1896: 438 (descr., synon.);
Eigenmann, Mem. Carneg. Mus., 5, 1912: 447 (synon., diagn., Georgetown, British Guiana); Norman,
Ann. Mag. nat. Hist., (9) ZZ, 1923: 15 (synon.,descr.); Jordan, Evermann, and Clark, Rep. U.S.
Comm. Fish. (1928), 2, 1930: 45 (ref. to orig. descr.).
Gnathobolus mucronatus Cuvier and Valenciennes, Hist. Nat. Poiss., 27, 1848: 91 (descr.).
Pristigaster mucronatus Ginther, Cat. Fish. Brit. Mus., 7, 1868: 462 (synon., descr.).
Odontognathus compressus Meek and Hildebrand 1923
Figures 110, III
Study Material. Numerous specimens, about 30-145 mm TL (caudal damaged
on 30-mm fish), 23-123 mm SL, including the holotype and 25 paratypes, from the
Atlantic coast of Panama, and a few lots from the Gulf of Venezuela.
Distinctive Characters. The nearest relative of O. compressus is O. panamensis of the
Pacific coast of Panama, but these species differ in having 52—60 and 65-68 anal rays,
respectively. Furthermore, in compressus the dorsal fin is notably in advance of the
middle of the anal base, whereas in panamensis it is about over the middle of the anal
fin. Its relationship with O. mucronatus is shown in the Key to Species (p. 430).
Description. Proportional dimensions in per cent of standard length, and counts,
based on Study Material, specimens 23-123 mm SL.
Body: depth 23.5-28.5. Pectoral fin: length 21-25.
Caudal peduncle: depth 7.25-8.3. Distance from snout to origin of: dorsal
Head: length 18-21; depth 18-22. 65-67.
Snout: length 4.2—$.1.
Eye: diameter 5.25—6.25. Scales: lost, ca. 36-43 pockets.
Interorbital: width 2.0-2.75. Ventral scutes: 25-29.
Maxillary: length 12-16. Vertebrae: 46 or 47 (3 specimens).
Mandible: length 9.3-10.2. Fin rays: dorsal 11-14, usually 11 or
Anal fin: length of base 47-54. 12; anal 52-60; pectoral 12-14.
Bopy elongate, excessively compressed, its greatest thickness about 0.17 of its
depth, its greatest depth 3.5-4.25 in SL; the dorsal outline of head concave in adults;
the ventral outline strongly convex anteriorly. CaupaL PEDUNCLE much shorter than
depth, its depth 2.3-2.9 in head.
Scares mostly lost; very thin, deeper than long, with nearly smooth margin,
without evident vertical striations. VENTRAL scuTEs well developed, the last 8—10 with
serrate margin.
Heap small, its length 4.75-5.5 in SL, its depth at vertical from slight cross-
groove at occiput 4.5-5.55. Snour a little shorter than eye, 3.6—4.7 in head. Eve 2.9—
28
434 Memoir Sears Foundation for Marine Research
3.8. INTERORBITAL very narrow, 7.0—10 in head. Cueex fully as long as deep. Moutu
nearly vertical. Maxittary broad, rounded in young, with an oblique margin in some-
what larger ones, ending under anterior half of eye, becoming produced into a narrow
process in examples 65 mm SL and upward, extending to or beyond margin of opercle
in large individuals, about 2.2 in head in small examples to 1.1 in large specimens.
Ficure 110. Odontognathus compressus, about 105 mm TL (caudal damaged), 88mm SL, Colén, Panama,
type, USNM 79533. Drawn by Ann S. Green.
ManpiBLE projecting strongly, its tip a little below dorsal outline of head, its margin
within mouth rising abruptly, forming an obtusely rounded angle, 1.75—2.0 in head.
Git rakers rather slender, about half as long as eye, with rather well-developed serrae
along inner edge, 19-23 on the lower limb. TEETH small to minute; apparently in a
single series on mandible and on margin of premaxillary and maxillary but absent on
membranous section between the last mentioned elements; minute teeth present on
palatines, pterygoids, and on median section of tongue.
Dorsat fin small, rather feebly developed, more or less damaged in all specimens
at hand, ending over middle of anal fin, its origin more
than a head-length behind origin of anal, not quite
twice as far from margin of snout as from base of cau-
dal, its distance from margin of snout 1.5-1.55 in SL.
Caupat fin rather broadly forked, the lower lobe the
longer, about as long as head. Anat fin long, its origin
about an eye’s diameter nearer to tip of mandible than
to base of caudal, its base 1.8—2.1 in SL. Pecrorat
fin a little longer than head, often reaching opposite to
last ventral scute, sometimes failing to reach this point,
4.0-4.75 in SL; free axillary process about half as
long as fin.
Color. Pale in alcohol. Sides of head silvery. Body
Figure 111. Odontognathus compres-
sus head showing produced maxillar : : :
ponents Aaa a id aloarn a Fig. with a silvery lateral band, about as broad as pupil.
110. Drawn by Ann S. Green. Back with dusky points and a median pale line in ad-
Fishes of the Western North Atlantic ARs
vance of dorsal; and a dark median streak behind it; margin of mandible and usually
the snout behind premaxillary with a dark crossline; base of caudal with a dark
crossline; base of anal with a row of dark dots; upper rays of pectoral with dusky
points.
Size. This is one of the smaller of the excessively compressed herrings. The largest
among the Study Material is 145 mm (5.7 in.) TL, 123 mm SL, which may be near
the maximum.
Development. The body increases greatly in depth with age and growth, as in many
other herrings. The depth in the smallest five examples at hand, 23-38 mm SL, is
contained 4.1—4.7 in SL whereas this proportion in the five largest is 3.5-3.6. Fur-
thermore, the dorsal profile anteriorly is about straight in the small specimens but
concave in the large ones. The maxillary, as indicated in the Description, is short in
small examples but becomes produced in large adults; in the smaller examples it is
broadly rounded posteriorly and reaches under the anterior half of the eye; at about
50 mm SL it is no longer rounded posteriorly and has an oblique margin; as growth
proceeds, the lower angle of the oblique margin becomes longer and sharper, and at
65 mm SL it is already well produced. However, it is only in large examples, upward
of roo mm SL, that the narrow process reaches to, or even slightly beyond, the margin
of the opercle.
Range. Known from the Atlantic coast of Panama, from the Gulf of Venezuela,
and from Trinidad.
Reference:
Odontognathus compressus Meek and Hildebrand, Field Mus. Publ., Zool., 75 (1), 1923: 194 (orig. descr.,
type local. Colén, Panama; cf. O. mucronatus; type USNM 79553).
Genus Neoopisthopterus Hildebrand 1948
Neoopisthopterus Hildebrand, Smithson. misc. Coll., TZ0 (9), 1948: 6; type species by original designation, Odon-
tognathus tropicus Hildebrand.
Characters. Bovy slender, strongly but not excessively compressed; ventral outline
anteriorly much more strongly convex than dorsal outline. Bony scurgs moderately
strong with entire (smooth) margin, the chest and abdomen with a sharp keel. Moutu
moderately oblique, with slightly projecting lower jaw. Maxittary short (with no in-
dication in the rather small specimens that it will become produced in adults, as in
Odontognathus), anteriorly overlapping distal end of premaxillary. Tezru small, present
on jaws, palatines, pterygoids, and tongue, but missing on vomer. GILL RAKERsS about
17-21 on lower limb of first arch. Dorsat fin small, with about 13-16 rays. ANAL
fin moderately long, with about 39-48 rays, beginning a short distance in advance
of origin of dorsal. Petvic fin wanting. VERTEBRAE about 46 or 47.
Remarks. This genus is characterized chiefly by the structure and relative position
of the bones in the upper jaw. In Neoopisthopterus the maxillary and premaxillary over-
28°
436 Memoir Sears Foundation for Marine Research
lap, with the margin of the jaw interrupted at the point where the maxillary passes over
the premaxillary. In related genera (Odontognathus and Opisthopterus) these bones are
separated by a membranous toothless section, with the margin of the upper jaw con-
tinous. The genus differs further from the other genera that lack the pelvic in having
fewer anal rays.
Range. Known from two species, N. ¢ropicus from the Pacific coast of Panama
and Peru and cubanus from Cuba.
Neoopisthopterus cubanus Hildebrand 1948
Figures 112, 113
Study Material. Six small specimens, 43-50 mm TL, 35-41 mm SL, apparently
young adults not fully mature; collected at Havana, Cuba, and sent to the writer by
Dr. Luis Howell-Rivero of the Museo Poey at Havana; USNM 143569 and
o35 10:
Distinctive Characters. N. cubanus is very closely related to N. tropicus Hildebrand
(57: 94, fig. 19), known from Panama and northern Peru, from which it differs only
in having somewhat fewer dorsal and anal rays (cubanus: D.13 or 14, A. 39-433
tropicus: D. 14-16, A 43-48). Also, these fins are somewhat differently placed; in
cubanus the dorsal originates slightly more than an eye’s diameter behind the beginning
of anal and about equidistant between the margin of the opercle and the base of caudal,
whereas in sropicus it begins scarcely an eye’s diameter behind the origin of anal and is
about equidistant between the posterior margin of the eye and the base of caudal. The
average number of gill rakers on the lower limb of the first arch apparently is lower
in cubanus than in fropicus (respectively, 17-19, 18-21). This, then, is another instance
of the rather common occurrence of “‘twin’’ species in the Atlantic and Pacific, gener-
ally found on the opposite coasts of the Isthmus of Panama. As the West Indian and
Atlantic Panamanian faunas are largely identical, cubanus may be expected on the Atlan-
tic coast of Panama and neighboring countries.
Description. Proportional dimensions in per cent of standard length, and counts,
based on study specimens 35-41 mm SL.
Body: depth 17-21. Anal fin: length of base 33-39.
Caudal peduncle: depth 6.5-8.5. Distance from snout to origin of: dorsal
Head: length 21.5-23.5; depth 18.5- 60-64.
20.5.
Snout: length 5.3-6.8. Scales: lost, ca. 43 pockets.
Eye: diameter 5.5—-6.8. Ventral scutes: 23-28.
Interorbital: width 2.6-3.1. Fin rays: dorsal 12-14; anal 39 to 43;
Maxillary: length 13.2-14.8. pectoral 13.
Mandible: length 12.8—13.7. Vertebrae: 47 (1 specimen).
Fishes of the Western North Atlantic 437
Bopy elongate, strongly but not excessively compressed, its greatest thickness
between a third and a fourth of its depth, its greatest depth 4.75-5.75 in SL; the
dorsal outline of head straight to slightly convex; the ventral outline rather strongly
convex anteriorly. CAUDAL PEDUNCLE shorter than deep, its depth 2.6—3.4 in head.
Scates mostly lost; rather large, very thin with smooth edge, and without evident
striations. VENTRAL scuTEs well developed, all with entire (smooth) margin.
Ficure 112. Neoopisthopterus cubanus, 50mm TL, 41 mm SL, Havana, Cuba, type, USNM 143569. Drawn
by Ann S. Green.
Heap fairly short and deep, its length 4.25—4.65 in SL, its depth at vertical from
slight crossgroove at occiput 4.9—5.4. SNouT about as long as eye (in small specimens),
3.34.5 in head. Eye 3.4-4.1. INTERORBITAL very narrow, 7.3—9.0. CHEEK about as
long as deep. Moutu moderately oblique. Maxitiary quite narrowly rounded poste-
riorly, reaching to or somewhat beyond vertical from posterior margin of pupil, 1.5 5—
1.8 in head. Manprste projecting slightly, its margin within mouth rising rather
abruptly, 1.6—1.75 in head. Grit raxkers slender, those at angle of first arch about as
long as pupil, with very fine serrae along the inner edge. Terr small to minute, ap-
parently in a narrow band on anterior part of lower jaw; those on maxillary and pre-
maxillary in a single series, interrupted at point of
overlapping of maxillary and premaxillary; very small
teeth on palatines and pterygoids and on median line
of tongue.
Dorsat fin quite small, somewhat elevated an-
teriorly, its margin convex, its rays rather feeble,
its origin rather more than an eye’s diameter behind
origin of anal and about equidistant between margin of
opercle and base of caudal, its distance from margin
of snout 1.55—1.65 in SL. Cavupat fin rather broadly
forked, the lower lobe slightly the longer, scarcely
as long as head. Anat fin moderately long, its origin Ficure 113. Neoopisthopterus cubanus
about equidistant between posterior margin of eye and head to show overlapping maxillary
: : and premaxillary bones; same as
base of caudal, its base 2.55-3.0 in SL. PEecToRAL
: 5 shown in Fig. 112. Drawn by Ann
fin damaged in all specimens at hand, apparently Ss. Green.
438 Memoir Sears Foundation for Marine Research
rather large, inserted below margin of opercle, and about 1.5 times farther from origin
of anal than from tip of mandible.
Color. Pale in alcohol. Side with a whitish band (no doubt silvery in life) about
half as broad as eye. Upper surface of head brownish posteriorly, with rather large
dusky dots; dusky dots on margin of snout medianly and on mandible anteriorly;
median line of back also with scattered dusky dots; dusky points on base of caudal
forming an oblique dark crossline and extending onto outer rays of fin; base of anal
with a row of dark spots, with fine punctulations on the fin, chiefly near the margin.
Size. This species belongs to a group of herrings that attains a small size, few
exceeding a length of 150 mm (6 in.). The known specimens of cubanus are all small
(apparently immature). A specimen of the closely related Pacific N. ¢ropicus, which
seemingly is mature, is 66 mm SL or about 77mm TL (3.1 in.), the caudal fin
broken. It may be conjectured, therefore, that NV. cubanus attains a length of only about
75-100 mm (3-4 in.).
Range. Known only from the type locality, Havana, Cuba.
Reference:
Neoopisthopterus cubanus Hildebrand, Smithson. misc. Coll., rro (9), 1948: 7, figs. 3 and 4 (orig. descr.; type
local. Havana, Cuba; type USNM 143569; cf. WN. ¢ropicus of the Pacific).
Genus Chirocentrodon Giinther 1868
Chirocentrodon Giinther, Cat. Fish. Brit. Mus., 7, 1868: 463; genotype by monotypy, Chirocentrodom taeniatus
Giinther equals Pe//ona bleekeriana Poey. Myers, Copeia, 1929: 1; Medipe//ona a synonym. Breder,
Copeia, 1942: 134; relationships.
Generic Synonym:
Medipellona Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67 (11), 1926: 417; genotype by original designa-
tion, Pellona bleckeriana Poey.
Characters. Bopy slender, strongly compressed; ventral outline anteriorly much
more strongly convex than dorsal outline. Bony scutes rather strong, about 24-28, the
chest and abdomen with a sharp keel. Mourn oblique, with moderately projecting
lower jaw. TEETH on Jaws, vomer, palatines, pterygoids, and tongue; each jaw (in adults)
with two or more canines anteriorly. Gitt RaKERS few, about 14-18 on lower limb
of first arch. Dorsat fin small, with about 14-16 rays. ANaL fin long, beginning in
advance of origin of dorsal, with about 39-43 rays. Petvic fin very small, with 6
rays. VERTEBRAE about 44 or 45.
Remarks. This genus is characterized by the elongated and strongly compressed
body with a strongly convex ventral outline anteriorly, the vomerine teeth, the canines
anteriorly in each jaw, and the long anal fin beginning in front of the dorsal.
Discussion. Chirocentrodon generally has been placed in the family Clupeidae, but
Jordan and Seale (75: 417), who evidently considered Chirocentrodon taeniatus Ginther
distinct from Pellona bleekeriana Poey (which has been disproved), erected a new genus
Fishes of the Western North Atlantic 439
Medipellona on Pellona bleekeriana and doubtfully assigned it to the family Engraulidae
(anchovies). Myers referred Medipellona to Chirocentrodon (96: 1). There seems to be
no sound reason for removing from the Clupeidae this little herring, which has much
in common with J/isha (and Neosteus), but it apparently should be considered as having
subfamily rank, Chirocentrinae, as already pointed out by Breder, who also discusses
the possible relationship to Chirocentrus, an Asiatic genus (22: 136).54
Range. Known from the West Indies and from the Atlantic coast of South America
from Panama to Sao Paulo, Brazil.
Chirocentrodon bleekerianus (Poey) 1867
Figures 114, 115
Study Material. Twenty-two specimens, ca. 52-107 mm TL (caudal fins dam-
aged), 42-84 mm SL, including the holotype and paratype of J/isha caribbaea Meek
and Hildebrand (equals C. bleekerianus), and a cotype of Pellona bleekeriana Poey, in
poor condition (USNM 120426); from Cuba; Jamaica; Puerto Rico; off Trinidad;
at Fort Sherman (Toro Point) and Porto Bello, Panama; Pt. Macolla, Gulf of Vene-
zuela; and Ubatuba, Sao Paulo, Brazil.
Distinctive Characters. The canine teeth in the anterior part of each jaw (in adults)
are characteristic. The presence of vomerine teeth also distinguishes b/eekerianus from
all American Clupeidae, except the common Atlantic herring, C/upea harengus.
Description. Proportional dimensions in per cent of standard length, and counts,
based on specimens 42—84 mm SL.
Body: depth 17.5-27. Pectoral fin: length 14.8-18.5.
Caudal peduncle: depth 7.8—9.5. Distance from snout to origin of: dorsal
Head: length 22-26; depth 17-22. 58-62.
Snout: length 5.7—8.2.
Eye: diameter 6.0-8.5. Scales: lost, ca. 43 pockets.
Interorbital: width 2.9-4.5. Ventral scutes: 24-28.
Maxillary: length 15-19. Fin rays: dorsal 14-16, usually 15; anal
Mandible: length 13.7-16.9. 39-43, rarely 44 or 45; pectoral
Anal fin: length of base 35-39. 13-15, usually 14.
Pelvic fin: length 3.4-4.9. Vertebrae: 44 or 45 (2 specimens).
Bopy moderately slender, strongly compressed, its greatest thickness about a third
of its depth, its greatest depth 3.85—5.55 in SL; the dorsal outline gently convex, the
ventral contour strongly convex anteriorly. CauDAL PEDUNCLE rather short, strongly
compressed, its depth 2.55—3.7 in head.
54. Following Breder’s excellent paper, there remains little doubt that Cirocentrodon is closely related to the two large
Indo-Pacific species of Chirocentrus and that both genera should be placed together in a separate subfamily or
family.—G. S. Myers.
44.0 Memoir Sears Foundation for Marine Research
Scates lost from specimens at hand; fairly large (judging from pockets). VENn-
TRAL SCUTES strong, 15-19 (most frequently 16 or 17) before pelvic fins and 8-10
(usually 9 or 10) behind them.
Heap 3.85-4.55 in SL, its depth at vertical from crossgroove at occiput 4.55—
5.9. SNout generally about equal to length of eye, without a definite median notch,
Ficure 114. Chirocentrodon bleekerianus, about 87 mm TL (caudal damaged), 70 mm SL, Kingston, Jamaica,
USNM 128274. Drawn by Ann S. Green.
3.25-4.0 in head. Eye 3.0-4.0 in head. InreRorpirat 6.0-8.2. CHEEK triangular,
rather longer than deep. Mouru moderately oblique. Maxitzary rather strongly con-
vex, narrow posteriorly, generally reaching vertical from posterior margin of eye,
1.25-1.55 in head. Manp1BLe projecting moderately, its tip well below dorsal outline
of head, its margin within mouth rising rather gradually, 1.45-1.75 in head. GrLi
RAKERS strongly serrate along inner margin, those at angle of first arch a little shorter
than eye, not increasing in number with age, 14-18 (most frequently 15 or 16) on
lower limb of first arch. TEETH prominent; an outer series of small teeth on mandible,
with I or 2 canines on each side anteriorly forming an incomplete inner series; a
somewhat irregular series on premaxillaries, with some of the anterior teeth (2 or 3 on
each side) somewhat canine-like; a series of rather
prominent slender pointed teeth, generally irregular
in size, on margin of maxillary; a few pointed ones
on vomer laterally; small teeth on palatines and ptery-
goids; and a narrow band of minute teeth on mid-
line of tongue.
Dorsat fin small, not greatly elevated anteriorly,
its longest rays generally reaching nearly or quite to
tip of last ray if deflexed, its origin a little nearer
to base of caudal than to posterior margin of eye,
its distance from margin of snout 1.6-1.7 in SL.
Caupat fin broadly forked, the lower lobe the larger,
nearly as long as head (Fig. 114). Anat fin long, its
Ficure 115. Chirocentradon bleekeria-
nus, enlarged head showing teeth, same
specimen as shown in Fig. 114. Drawn
by Ann S. Green. anterior rays somewhat elevated, its origin about
Fishes of the Western North Atlantic 441
an eye’s diameter in advance of dorsal, its base 2.5—2.95 in SL. Petvic fin shorter
than eye, inserted a little nearer to origin of anal than to base of pectoral, 5.4—6.9 in
head. Pecrorat fin reaching well beyond base of pelvic, 5.4—6.75 in SL, 1.3-1.6 in
head; the free axillary process about a third of the length of the fin.
Color. In alcohol, pale with a fairly definite white lateral band (presumably silvery
in life); nape and tip of snout with dusky punctulations; the more recently preserved
specimens with dusky punctulations on entire median section of back, mostly arranged
in two parallel series behind dorsal fin; dorsal and caudal fins with dusky punctulations;
base of anal generally with a row of small dark spots.
Size. This species is among the smallest of herrings. The largest at hand, about
107mm TL (4.3 in.; 84 mm SL), seems to be the largest example known.
Development. Specimens less than 52 mm TL, 42 mm SL, remain unknown.
Nevertheless, a considerable increase in proportionate depth takes place with age and
growth, as the large specimens at hand are notably deeper than the smaller ones. The
greatest depth in a specimen 42 mm SL is contained 5.7 times in SL whereas in one
84 mm SL this proportion is only 3.7.
Although proportionate increase in the body depth with age and growth is so
usual among Clupeidae that it is expected, a proportionate increase in the size of eye
during development is so unusual as to be subject to question. However, the speci-
mens that are now at hand, suspected at first, did in fact show this characteristic.
Using again the two extremes in size, it is found that the eye in the 42-mm specimen
is contained 3.55 times in the head and is only 7.1 °/, of SL, whereas in the 84-mm
fish it is contained 3.0 times in the head and is 8.3 °/, of SL. From the fairly complete
and graduated growth series within the range already mentioned, it is evident that the
eye does increase in proportionate size with age and growth. It may be assumed, there-
fore, that this development, the reverse of what usually obtains in fishes, is normal in
C. bleekerianus.
Spawning. A female about 107mm TL, containing well-developed eggs fully
0.5 mm in diameter, was taken off Trinidad by the AtBarross, in 31-34 fms. As it
was caught on February 3 (1884), the indication is that C. b/eekerianus spawns during
the northern winter.
Relationship. This species, as now understood, is the only one belonging to the
genus Chirocentrodon. It has been described under at least three different names, as in-
dicated in the Synonyms, but Breder has shown adequately that the three constitute
only one species (22: 133).
Commercial Importance. This little herring was seen occasionally in the market in
Port-au-Prince, Haiti, by Beebe and Tee-Van, who also stated that it is rare in Haiti
(zo: 43). It is not known to be numerous anywhere, therefore its importance as food
for man as well as for fishes and other animals feeding on fish is probably slight.
Range. West Indies, from Cuba, Haiti, Jamaica, Puerto Rico, and Trinidad;
also from the Atlantic coast of Panama, the Gulf of Venezuela, and Ubatuba, Sao
Paulo, Brazil.
44.2 Memoir Sears Foundation for Marine Research
Synonyms and References:
Pellona bleekeriana Poey, Repert. Fisico-Nat. Cuba, 2, 1867: 242 (orig. descr.; type local. Matanzas, Cuba;
cotypes MCZ 17845; one USNM 120426); Poey, Synop. Pisc. Cubens., 1868: 419 (ref. to orig. descr.,
notes); Poey, An. Soc. esp. Hist. Nat., 5 (2), 1876: 180, p.148 in separate (common name, ref.);
Howell-Rivero, Bull. Mus. comp. Zool. Harv., 82 (3), 1938: 171 (cotypes in bad condition).
Chirocentrodon taeniatus Giinther, Cat. Fish. Brit. Mus., 7, 1868: 463 (orig. descr.; type local. Jamaica, West
Indies; types BMNH); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 435 (descr. after
Giinther); Norman, Ann. Mag. nat. Hist., (9) IZ, 1923: 21 (descr., P. b/eekeriana probably not distinct) ;
Beebe and Tee-Van, Zoologica, N. Y., To (1), 1928: 43, fig. (refs., diagn., tables, size, local abund.,
Port-au-Prince, Haiti); Myers, Copeia, 1929: 1 (C. d/eekerianus and C.caribbaeus probably distinct);
Jordan, Evermann, and Clark, Rep. U. S. Comm. Fish. (1928), 2, 1930: 45 (ref.).
Ilisha bleekeriana Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 436 (descr. after Poey); Evermann
and Marsh, Bull. U.S. Fish Comm. (1900), 20 (1), 1902: 86 (diagn.); Nichols, N. Y. Acad. Sci., Zo
(2), 1929: 204 (distr., diagn., Puerto Rico).
Llisha caribbaea Meek and Hildebrand, Field Mus. Publ., Zool., 75 (1), 1923: 191, pl. 10, fig. 1 (orig. descr.;
type local. Porto Bello, Panama; type USNM 81769); Jordan, Evermann, and Clark, Rep. U. S. Comm.
Fish. (1928), 2, 1930: 45 (ref.).
Medipellona bleekeriana Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67 (11), 1926: 417 (descr. after
Poey; relation.).
Medipellona caribbea Jordan and Seale, Bull. Mus. comp. Zool. Harv., 67 (11), 1926: 418 (diagn.).
Chirocentrodon bleekerianus Fowler, Proc. Acad. nat. Sci. Philad., 80, 1928: 456 (descr., taentatus Giinther
synon., Haiti); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 45 (ref.); Breder,
Copeia, 1942: 134, fig. 1 (synon., relation., descr.).
Genus Dorosoma Rafinesque 1820
Gizzard Shads, Threadfin Shad
By
Rosert Rusu MILier
Museum of Zoology
The University of Michigan
Dorosoma Rafinesque, West. Rev. misc. Mag., 2 (3), 1820: 171 (see Fowler, Monogr. Acad. nat. Sci. Philad.,
7, 1945: 6, 8); Ichthyol. Ohiensis, 1820: 39; type species by monotypy, D. wotata equals D. cepedianum
(LeSueur). Below falls of Ohio River.
Generic Synonyms:
Chatoessus Cuvier, Régne Anim., ed. 2, 2, 1829: 320, in part; restricted by Cuvier and Valenciennes to Mega/ops
cepediana LeSueutr.
Signalosa Evermann and Kendall, Bull. U.S. Fish Comm. (1897), 17, 1898: 1273 type species, Signalosa
atchafalayae equals D. petenense (Giinther). Atchafalaya River, Melville, Louisiana.
Characters. Bopy compressed, silvery. The abdomen armed with Bony scurTss;
total 23-32. Scares cycloid, thin, 40-83 along side. Mourn small to moderate in
size, terminal, subterminal, or inferior, the lower jaw included or jaws subequal. SNout
short and rounded. Maxriiary with two supramaxillary bones. Trrrx absent in adult,
but a row of fine teeth on upper jaw in young. Dorsat with last ray prolonged into a
slender filament (absent or inconspicuous in young), thus resembling Opisthonema.
Fin rays: dorsal 9-15, anal 17—38, pelvic 8, pectoral 12-17, caudal 19 (17 branched),
rarely 17 or 18. Sromacu gizzard-like, the Inrzstine long and much convoluted,
with numerous Pytoric cagca. VERTEBRAE (including urostyle) 40-51.
Remarks. Dorosoma may be readily distinguished from Opisthonema in having
(1) the ridge of the back before the dorsal fin naked rather than crossed by scales;
(2) no bilobed dermal fold on the vertical anterior edge of the cleithrum; (3) an axil-
lary scale, the pectoral fins not folding into a groove; (4) the dorsal origin not well in
advance of the insertion of the pelvic fins; and (5) gill rakers 200-400 in Dorosoma,
only about 65-109 in half-grown to adult Opisthonema.
Range. All species except D. smithi are confined to the Atlantic drainage of North
and Middle America, from Canada to Nicaragua. They are found from southern South
Dakota, Nebraska, and Minnesota, from the Great Lakes-St. Lawrence River Basin, and
from about 40 to 41°N on the Atlantic seaboard (New York, New Jersey, and south-
eastern Pennsylvania) southward to Lake Nicaragua, but with a discontinuous distri-
bution from northern Guatemala and British Honduras to Nicaragua. D. smithi is
known only from coastal streams of northwestern México, in Sonora, Sinaloa, and
Nayarit. Only two species, D. cepedianum and D. petenense, have been taken in brackish
and salt water—the former from as far north as Sandy Hook Bay, New York, south-
ward to the vicinity of Tampico, México.
443
444 Memoir Sears Foundation for Marine Research
Key to Species of Dorosoma Known from Brackish or Salt Water
ta. Mouth terminal; ventral edge of upper jaw smooth; fewer than 50 scales in lateral
series, regularly arranged; anal rays 17-27, usually 20-25; prepelvic scutes 14—
17 (rare); vertebrae 40-45. petenense (Giinther) 1866, p. 448.
1b, Mouth subterminal or inferior; ventral edge of upper jaw with pronounced notch
(except in young); more than 50 scales in lateral series, irregularly arranged;
anal rays 25-36, usually 29-35, where sympatric with D. petenense; prepelvic
scutes 17—20; vertebrae 48-51. cepedianum (LeSueur) 1818, below.
Dorosoma cepedianum (LeSueur) 1818
Gizzard Shad, Eastern Gizzard Shad
Figure 116
Study Material. Seventy-four specimens, 78-247 mm SL, for body proportions,
51-200 for meristic data except as otherwise stated, from various parts of its range,
Canada to México.
Distinctive Characters. The comparatively small scales, 52—70 in lateral series
and 36-45 around the body, a higher vertebral count (48-51, usually 50), the longer
anal fin and greater number of rays (2 5-36), and preponderantly 18 + 12 ventral scutes,
distinguish D. cepedianum from its sympatric relative D. petenense. Another relative,
Opisthonema oglinum, may be separated from D. cepedianum by features listed in the
generic account under Remarks (p. 443).
Description. Proportional dimensions in per cent of standard length and head
length, with all measurements “stepped off” with a pair of precision dividers; based
on Study Material, specimens 78-247 mm SL; counts based on 51-200 specimens,
the number of specimens counted being given in parentheses.
Body: depth 2.3-3.1 in SL.
Caudal peduncle: length 2.4-3.5 and
depth 2.45-3.4 in hl.
Head: length 3.0-3.9 in SL; width 1.8—
2.§ in hl.
Snout: length 5.0—6.0 in hl.
Eye: length 3.3—-5.4 in hl.
Interorbital: bony width 3.3-4.5 in hl.
Upper jaw: length 3.5-4.2 in hl.
Mandible: length 2.6—3.3 in hl.
Dorsal fin: base 1.8 5—2.6 in hl; length of
filament 3.1-6.5 in SL, 0.8—2.0
in hl.
Caudal fin: length of lower lobe 0.7—1.1
in hl, typically longer than anal
base (rarely subequal).
Anal fin: length of base 3.2—-3.9 in SL,
0.75—1.25 in hl.
Pelvic fin: length 2.0-2.5 in hl.
Pectoral fin: length 1.15-1.45 in hl.
Distances: predorsal 1.8 5—2.05; prepelvic
2.0-2.35; anal origin to caudal
base 2.4—2.8, all in SL.
Scales: lateral series 52—70, av. 61.06
(67), the first scale counted being
the first above uppermost corner of
gill opening; between dorsal and
Fishes of the Western North Atlantic 44.5
anal fins 19-24, av. 21.77 (52); 65 mm SL, ca. 350 at 95 mm, and
around body 36-45, av. 41.03 (73), 412 at $7 mm; very numerous and
beginning with first one just in fine (rakers counted ix situ).
front of left pelvic fin; around cau- Fin rays: dorsal 10-13, av. 11.61 (197);
dal peduncle (slenderest part) 16— caudal 19; anal 25-36, av. 31.32
2OWavale-O2), (53): (195); pelvic 8, rarely 7; pectoral
Scutes: prepelvic 17-20, av. 17.99 (196); 14-17, av. 15.52 (2883 144 fish).
postpelvic 10-14, av. 11.76 (197); Vertebrae: 48-51, av. 49.83, including
total ventral 27-32, av. 29.74 (196). urostyle (42).
Gill rakers: on first arch, >300 at
Ficure 116. Dorosoma cepedianum, from Cumberland River, Tennessee, USNM 20034. After Goode, from
drawing by H. L. Todd, somewhat emended.
No external characteristics will reliably distinguish the sexes.
Color. In life, body silvery. Bluish over back and upper sides but milky-white on
abdomen, often with brassy or golden reflections from scales. About 6-8 horizontal,
dark stripes along upper sides above level of middle of shoulder spot, extending from
behind head to base of caudal fin. A large, round dark spot behind opercle lustrous
purple, prominent in young and half-grown. Dorsal fin of adults nearly uniformly
dusky; caudal fin dusky but darkened on its outer third; anal fin with outer 67 °/,
dark, the basal third lighter, with melanophores sprinkled over most of fin; pectorals
and pelvics with their outer halves darkened, grading to pale basally. Top of head,
snout, upper jaw, and upper part of opercle pigmented in young and adult; the rest of
head silvery. In young-of-the-year (up to about 4.5 in. TL), dorsal fin sparsely but
uniformly sprinkled with chromatophores; caudal fin similar but with more pigment
cells; anal, pectoral, and pelvic fins almost unpigmented.
Size. The Gizzard Shad is known to attain a total length of 20.5 inches but does
not commonly grow longer than 10-14 inches. In a letter dated July 14, 1953, Mr.
446 Memoir Sears Foundation for Marine Research
Alfred Larsen (Fishery Biologist, Pennsylvania Fish Commission) wrote that speci-
mens of D. cepedianum killed in Presque Isle Bay, Pennsylvania, early in 1953 ranged
from 4 to 19 inches. Fish 10-13 inches long weigh about one pound, individuals
14-18 inches, 1-3 pounds; the heaviest recorded specimen weighed 3 pounds, 7
ounces (Trautman, 126: 182).
Development and Growth. The prolarvae perform characteristic alternate sinking
and rising movements for the first two days after hatching. The larvae are characterized
by along gut, 1.5—2.0 times the total length of the fish, by the retarded development
of the single dorsal fin, and by the elongated anal fin with 22 rudimentary rays at
17.5 mm and 30-34 at 19-22 mm TL.
In contrast to the deep-bodied and strongly compressed adults, the very young are
slender, cyprinid-like, and nearly cylindrical; the maxillaries have a few teeth on the
lower edge which are lost with age, and unlike the adults with a highly specialized
digestive tract, the young have an almost straight intestine and no pyloric caeca.
By the end of the first summer of life the young possess the specializations of
the adults.
At the end of the first summer, an average total length of about 4 inches is at-
tained in Indiana and Ohio, 4.5 inches in the Chesapeake Bay region, and 5.0 inches
in Tennessee and Oklahoma. The sizes in two Indiana ponds, after the first year, have
been reported as follows: second summer, 7—9 inches, average 7.6; third summer,
g—I1 inches, average 9.5; fourth summer, 9-12 inches, average 10.5; fifth summer,
10-13 inches, average 11.3; and sixth summer, 11-15 inches, average 12.8. Al-
though they may live as long as 10 years, the life span is usually not more than seven
years. In Lake Erie about 75 °/, of the annual growth of the Gizzard Shad takes place
during the summer; in winter they grow little if at all and lose considerable weight.
Maturity typically occurs in the second or third year of life, at about 7-11 in-
ches, but Vladykov (127: 35) has reported a mature female that was only 6 inches
(151 mm) TL.
Spawning. Gizzard Shad are known to spawn only in fresh water, from late winter
(mid-March) through most of the summer (at least to August 20) in sloughs, ponds,
lakes, and large rivers. Spawning generally occurs on a rising temperature, usually be-
tween about 50° and 70°F. The creamy-yellow eggs, demersal and adhesive, are about
0.75 mm in diameter when fertilized and fixed. The embryos hatch after 95 hours of
incubation at 62°F, or after about 36 hours at 80°F.
Precocious females appear to have few eggs. Those belonging to age-group 11
have the most, averaging about 380,000 eggs in five females 291 mm long; egg pro-
duction then declines with successively older groups.
Food. Gizzard Shad are essentially filter feeders. Except for a few weeks after
hatching, those in fresh water are almost entirely herbivorous, feeding heavily on
microscopic plants, phytoplankton, and algae. After they reach a length of about 20-
25mm TL, the diet is remarkably alike at various sizes. The food habits in salt and
brackish water have not been studied.
Fishes of the Western North Atlantic 447
Habitat and Occurrence. Gizzard Shad inhabit large rivers, reservoirs, lakes,
swamps, bays, barrow pits, estuaries, bayous, temporary floodwater pools along large
river courses, sloughs, and similar quiet, open waters that may be clear to very silty.
The very young apparently do not enter brackish water; in coastal rivers of Virginia
the young occur in greatest abundance well upstream. In Chesapeake Bay, Gizzard Shad
are common only during the fall months, occurring principally in brackish water near
the mouths of freshwater streams, but in the rivers of that region they are common
or abundant throughout the year. Along the Texas coast, they frequent the large
brackish-water bays where individuals may be taken throughout the year in waters
that vary in salinity from 2.0 to 33.7 °/)). Thus Gizzard Shad in this area also prefer
brackish water to sea water. In general the smallest fish occur in the freshest water,
with size increasing as salinity increases.
Dr. R. D. Suttkus of Tulane University has taken D. cepedianum at about 20
brackish and saltwater stations in Louisiana, not including nearly 40 separate captures
from Lake Pontchartrain. Most of the 20 records are from along the Gulf of Mexico
coast or within embayments connected with the Gulf. I am grateful to Dr. Suttkus for
these data.
Range. Atlantic drainage of eastern North America south to Rfo Panuco,
México.
Synonyms and References:
Megalops cepediana LeSueur, J. Acad. nat. Sci. Philad., r, 1818: 361 (orig. descr.; markets of Baltimore and
Philadelphia, hence usually given as Chesapeake and Delaware bays).
Clupea heterura Rafinesque, Amer. Month. Mag., 1818: 354 (orig. descr.; Ohio River).
Dorosoma notata Rafinesque, West. Rev. Misc. Mag., 2 (3), 1820: 172 (orig. descr.; type local., falls of
Ohio R.).
Chatossus ellipticus Kirtland, Rep. Zool. Ohio iw Second Rep. geol. Surv. Ohio, Columbus, 1838: 169, 195
(NoMEN NuDUM, Ohio; same as D. xotat2); Boston J. nat. Hist., 4 (2), 1842: 235, pl. 10, fig. 1 (orig.
descr.; compar., occur. in Ohio, habits, mortal., Ohio R. and tributaries).
Megalops bimaculata LeSueur im Cuvier and Valenciennes, Hist. nat. Poiss., 22, 1848: 104. (NOMEN NUDUM,
synon. with Chatoessus cepedianus by Valenciennes).
Chatoessus insociabilis Abbott, Proc. Acad. nat. Sci. Philad. (1860), 72, 1861: 365 (orig. descr.; habits, “sturgeon
pond 2 miles below Trenton, New Jersey”).
Chatoéssus cepediana Giinther, Cat. Fish. Brit. Mus., 7, 1868: 409 (descr., based on specimens from New
York).
Dorosoma cepedianum exile Jordan and Gilbert, Proc. U.S. nat. Mus. (1882), 5, 1883: 585 (orig. descr.; Gal-
veston Bay, Texas).
Dorosoma cepedianum Goode, et al., Fish. Fish. Industr. U.S., 1, 1884: 610 (in brackish or sea water, Atlantic
coast); Higgins and Pearson, Rep. U.S. Comm. Fish. (1927), 1928: 42 (Pamlico Sound, N. Carolina);
Hildebrand and Schroeder, Bull. U.S. Bur. Fish. (1927), 43, 1928: 106 (Chesapeake Bay, rare in strictly
salt water); Fowler, Monogr. Acad. nat. Sci. Philad., 7, 194.5: 165 (Cape I. and rice field near Cordesville,
S. Carolina); Gunter, Publ. Inst. mar. Sci. Texas, r (1), 1945: 30 (brackish and salt water, Texas coast;
prefers brackish to sea water); Massmann, Trans. 18th N. Amer. Wildl. Conf., 1953: 439 (young in
Virginia estuaries); Bailey, Winn, and Smith, Proc. Acad. nat. Sci. Philad., 106, 1954: 120 (common in
fresh tidewater, Escambia R., Florida); Miller, Fish. Bull. (173) U.S. Fish Wildl. Serv., 60, 1960 (review
of systematics and biol.); Minckley and Krumholz, Zoologica, N.Y., 44, 1960 (hybrid. with D.
petenense).
Dorosoma exile Meek, Field Mus. Publ., Zool., 5, 1904: 94 (descr., lowland tributaries of Gulf of Mexico
N. of Veracruz, México, San Juan, Forlén, Valles).
448 Memoir Sears Foundation for Marine Research
Dorosoma petenense (Giinther) 1866
Threadfin Shad
Figure 117
Study Material. Twenty-five specimens, 59-180 mm SL, for measurements, and
186 for meristic data except as otherwise stated, from Florida to Lake Péten,
Guatemala.
Distinctive Characters. The terminal mouth and the smooth ventral edge of the
upper jaw distinguish this species from the other forms of Dorosoma. In addition it is
\\
Ficure 117. Dorosoma petemense, from Atchafalaya River, Alabama, USNM 48790. After Evermann and
Kendall.
readily separated from D. cepedianum in having larger, regularly arranged scales (fewer
than 50 in lateral series), only 40-45 vertebrae, a smaller anal fin with 17-27 rays
(usually 20-25), a longer and more pointed snout, longer upper jaw and mandible,
and usually 16 or 17+9-11 ventral scutes. This species may be confused with
Opisthonema oglinum, since both have a prolonged dorsal filament, but the latter is
distinguished by the features given under Remarks in the generic account (p. 443).
Description. Proportional dimensions in per cent of standard length and head
length, with all measurements “‘stepped off” with precision dividers; based on speci-
mens 59-180 mm SL; counts based on 186 specimens or less, the number of speci-
mens counted being given in parentheses.
Body: depth 2.4—3.0 in SL.
Caudal peduncle: length 2.3-3.4 and
depth 2.25-2.8 in hl.
Head: length 2.95-3.7 in SL; width
1.95—2.4 in hl.
Snout: length 4.1-4.85 in hl.
Eye: length 3.3—4.1 in hl.
Interorbital: bony width 3.5—4.5 in hl.
Upper jaw: length 2.8—3.35 in hl.
Mandible: \ength 2.1-2.5 in hl.
Dorsal fin: base 1.5-2.05 in hl; length
of filament 2.7—3.1 in SL, 0.6—-1.0
in hl.
Caudal fin: length of lower lobe 0.7-0.9
in hl.
Anal fin: length of base 3.5-4.35 in sl.
Fishes of the Western North Atlantic 44.9
Pelvic fin: length 2.05—2.4 in hl. postpelvic 8-12, av. 9.95 (178);
Pectoral fin: length 1.2-1.35 in hl. total ventral 23-29, av. 26.75
Distances: predorsal 1.9-2.25; prepelvic (176).
1.9-2.25; anal origin to caudal Gill rakers: ca. 300 at 66mm SL, 330
base 2.6—3.13 all in SL. at 117mm, 350 at 131 mm, and
about 440 at 180mm; fine and
Scales: lateral series 41-48; between dor- numerous (rakers counted in situ).
sal and anal fins 14-17; around Fin rays: dorsal 11-14, av. 12.13 (178);
body 28-36; around caudal pe- caudal 19; anal 17-27, av. 22.38
duncle 16-20 (all counts based on (179); pelvic 8, rarely 7; pectoral
25 specimens covering the size 12-17, av. 14.39 (354, 177 fish).
range of this species). Vertebrae: 40-45, av. 42.23, including
Scutes: prepelvic 15-18, av. 16.79 (176); urostyle (30).
As in D. cepedianum, there appear to be no reliable external features that will
distinguish the sexes.
Color. In life, body bright silvery, especially on sides, opercles, and underparts,
but back and upper sides bluish-black or dark olivaceous, washed with gold and over-
lain with horizontal rows of dark stripes. A conspicuous, jet-black round or oval spot
behind upper part of opercle. Caudal with middle portion of each fin lobe bright golden
yellow, fading basally and distally, the posterior border dusky; dorsal fin dusky, washed
with yellowish olive; anal fin intensely yellow; paired fins yellow basally. A broad golden
stripe (often represented only by gilt reflections) in the light band between back and
sides, but almost no gilt on sides of head.
Size and Age. The Threadfin Shad is a smaller species than the gizzard shad.
In the northern part of its range, the adult generally does not exceed § or 6 inches
TL; in southern states it may grow to 7 inches, and in Guatemala to nearly 8.75
inches or 180 mm SL (specimen from Laguna de Yalac; UMMZ 143386). The
species has a short life-span, for few individuals appear to exceed two years of age.
Variations in Size with Season. In bays along the Texas coast, adults were most
abundant in August and October while the young were entirely absent; between August
and November the size increased from a mode between 103 and 108 mm TL to
118 mm. In January, only fish with a modal length of 63 mm came into the catch, and
in February the same group was present along with a few fish 114 and 117 mm long.
Possibly early growth takes place only in fresh water. Water temperatures in the bays
of Texas varied from about 48 to 86°F over the year.
Spawning. Threadfin Shad may spawn when they are less than one year old; there
appear to be two spawning peaks—spring and fall. Spawning occurs at about 70° F in
open waters, near or over plants and other objects. The eggs are slightly adhesive.
Habits. Threadfin Shad are generally pelagic; they often feed and migrate in
schools numbering thousands of individuals. The different age-groups usually remain
in separate schools. They are known to concentrate at the downstream faces of dams
29
4.50 Memoir Sears Foundation for Marine Research
and similar obstructions, and large aggregations are seen at reservoir inlets. Water
current obviously attracts large schools; when the electric turbines were reduced or
shut off in the Tennessee impoundments, the Threadfin Shad were difficult to find,
but when the power was turned on they were present in abundance.
Food and Feeding. Like their close relative, these fish are plankton feeders, the major
items of the adult diet consisting of algae (diatoms and unicellular green species), cope-
pods, and cladocerans. Threadfin Shad introduced into the Salton Sea, California, have
a preference for the fry of Bairdiella icistius (Jordan and Gilbert), which suggests that
they may have similar food habits in coastal lagoons and estuaries within their natural
range.
Tolerance to Temperature. As in the case of the gizzard shad, sudden temperature
changes often produce high mortality among Threadfin Shad. Berry et a/. have observed
a spawning mortality for both species during April in Florida lakes. D. petenense cannot
tolerate as low temperatures as D. cepedianum. For Threadfin Shad in the Colorado
River, Texas, temperatures of about 54—58° F may be close to the minimum tolerance.
However, among experimental individuals from the lower Tennessee River, subjected
to water cooled from 50 and 60°F, there was a high mortality below 45°, with very
few fish surviving below 40°.
Habitat and Occurrence. Threadfin Shad seek habitats similar to those of the gizzard
shad and are often associated with D. cepedianum where the two species are sympatric.
They live in bays, large rivers, reservoirs, lakes, estuaries, and other similar situations.
Along the coast of Texas they occur in bays having salinities varying from 3.8 to
26.9 %/y9, the majority having been taken in waters with salinities between 10 and
20 °/,). Apparently these fish show a preference for water that is neither fully marine
nor very fresh, a condition typical in many lagoons along the coast of Texas. Larger fish,
approximately 100-150 mm long, occurred in waters of about 15-30 °/o, whereas
smaller fish, generally below 100 mm, were taken in salinities below 15 °/o9.
Records of Threadfin Shad obtained by Dr. R. D. Suttkus through 1958 indicate
their occurrence at nearly 50 brackish and saltwater stations in Florida, Alabama,
Mississippi, Louisiana, and Texas, not including about 75 captures from Lake Pont-
chartrain.
Range. From the Ohio River of Kentucky and southern Indiana westward and
southward to Oklahoma, Texas, and Florida, thence along the coast of the Gulf of
México to northern Guatemala and British Honduras (Belize River). Their recent in-
vasion of the Ohio River Basin is discussed by Minckley and Krumholz (94).
Synonyms and References:
Meletta petenensis Giinther, Proc. zool. Soc. Lond., 1866: 603 (orig. descr.; Lake Petén, Guatemala).
Chatoessus petenensis Giinther, Cat. Fish. Brit. Mus., 7, 1868: 408 (redescr. of types).
Chatoessus mexicanus Giinther, Cat. Fish. Brit. Mus., 7, 1868: 409 (orig. descr.; México, New Orleans, Louisi-
ana).
Dorosoma mexicanum Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 416 (descr., from Giinther).
Dorosoma petenense Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 417 (descr., from Giinther);
Regan, Biol. Centr. Amer., Pisces, 8, 1906-08: 178 (redescr. of types); Fowler, Proc. Acad. nat. Sci.
Fishes of the Western North Atlantic 451
Philad., 63, 1911: 211 (“Panama,” obviously in error); Bailey, Winn, and Smith, Proc. Acad. nat. Sci.
Philad., 106, 1954: 119 (range; from fresh tidal waters, Escambia R., Florida); Berry, Huish, and Moody,
Copeia, 1956: 192 (spawn. mortal. in Florida, life-hist. notes); Kimsey, Calif. Fish Game, Inl. Fish.
Adm. Rep., 58-16 (mimeo), 1958: 1-21 (life hist., proposed introd. into Sacramento—San Joaquin
Delta); Shapovaloy, Dill, and Cordone, Calif. Fish Game, 45, 1959: 166 (review introd. into California).
Signalosa atchafalayae Evermann and Kendall, Bull. U.S. Fish Comm. (1897), 77, 1898: 127, pl. 7, fig. 4
(orig. descr.; Louisiana and Mississippi; use as bait; type local., Atchafalaya R. at Melville, Louisiana).
Signalosa mexicana Meek, Field Mus. Publ., Zool., 5, 1904: 94 (synon., descr., lowland streams trib. to Gulf
of Mexico, México—Valles, Obispo, El Hule, Perez); Regan, Biol. Centr. Amer., Pisces, 8, 1906-08:
178 (redescr. of types, range); Fowler, Proc. Acad. nat. Sci. Philad., 63, rgt1: 211 (Volusia, Florida);
Regan, Ann. Mag. nat. Hist., (8) I9, 1917: 310 (descr. 9 specimens, includ. types, from Louisiana to
Centr. Amer.); Gunter, Publ. Inst. Mar. Sci. Texas, 1 (1), 1945: 31 (prov. ident., size range, temp.
range, and salinity, Copano and Aransas bays, Texas).
Signalosa petenensis Regan, Ann. Mag. nat. Hist., (8) 19, 1917: 310 (redescr. based on types); Hubbs, in Hubbs
and Allen, Proc. Fla. Acad. Sci., 6, 1943: 116 (8. mexicana and 8. atchafalayae synon. with 8. petenen-
sis); Miller, Proc. Okla. Acad. Sci. (1953), 34, 1955: 33-34 (first rec. for Oklahoma); Riggs and Moore,
Proc. Okla. Acad. Sci. (1957), 38, 1958: 64-67 (Lake Texoma, Oklahoma and Texas); Minckley and
Krumholz, Zoologica, N.Y., 44, 1960 (chars., hybrid. with D. cepedianum, range).
Signalosa mexicana mexicana Weed, Field Mus. Publ., Zool., r2 (11), 1925: 142 (synon., char., basin of Rio
Papaloapan, Veracruz and Oaxaca, México).
Signalosa mexicana campi Weed, Field Mus. Publ., Zool., 12 (11), 1925: 143 (orig. descr.; compar., type local.
Resaca de la Guerra, on or near Media Luna Ranch, Brownsville, Texas).
Signalosa atchafalayae atchafalayae Weed, Field Mus. Publ., Zool., 12 (11), 1925: 144 (synon., char., compar.,
Louisiana).
Signalosa atchafalayae vanhyningi Weed, Field Mus. Publ., Zool., 12 (11), 1925: 145 (orig. descr.; type local.
Prairie Cr., 6 mi. SE of Gainesville, Florida).
Signalosa petenensis vanhyningi Hubbs and Allen, Proc. Fla. Acad. Sci., 6, 1943: 116 (common name, habits,
color, Silver Springs, Florida); Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1945: 266 (synon., refs.,
Volusia, Florida); Parsons and Kimsey, Prog. Fish-Cult., 16 (4), 1954: 179-181, 1 fig. (distr., review
of biol., use as forage fish).
Signalosa petenensis campi Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1945: 22 (Rio Grande).
Signalosa petenensis atchafalayae Fowler, Monogr. Acad. nat. Sci. Philad., 7, 1945: 366, 372 (synon., refs.,
Avery I., Louisiana; Galveston, Texas); Hubbs, Copeia, 1951: 297 (min. temp. tolerance in Colorado
R., Austin, Texas); Chance and Miller, J.’ Tenn. Acad. Sci., 27 (3), 1952: 218 (first publ. rec. for
Tennessee Valley; effect of severe winter, 1950-51); Kimsey, Calif. Fish Game, 40, 1954: 204 (introd.
into California).
Dorosoma petenensis atchafalayae Kimsey, Trans. Amer. Fish. Soc. (1957), 87, 1958: 331, 332 (forage use in
California and Arizona).
Dorosoma petenense atchafalayae Haskell, Copeia, 1959: 298—302 (food habits, Arizona).
Io.
It.
YS So ae
TEXT AND FOOTNOTE REFERENCES
Axoxymous, Rep. Newfoundld. Fish. Res.
Comm., 2 (1), 1933-
Anonymous, Rep. U.S. Comm. Fish. (1897),
1898 (Manual of Fish Culture).
Anonymous, Rep. U.S. Tariff Comm., 68,
1933.
Bassoun; Bull. Mus. comp. Zool. Harv., 46
(7), 1905.
Barney, Trans. Amer. Fish. Soc., 54, 1924.
Deleted.
Battle, Huntsman, Jeffers, et @/., J. biol. Bd.
Canada, 2, 1936.
Bean, Bull. N. Y. St. Mus., 60, Zool. 9, 1903.
Bean, Proc. biol. Soc. Wash., 25, 1912.
Beebe and Tee-Van, Zoologica, N. Y., Io (1),
1928.
Beebe and Tee-Van, Field Bk. Shore Fish.
Bermuda, 1933.
Bere, Amer. Midl. Nat., 77 (3), 1936.
Bertin, Bull. Inst. océanogr. Monaco, 853,
1943-
Bigelow and Schroeder, Bull. U.S. Bur. Fish.,
48 (20), 1936.
Bigelow and Schroeder, Fish. Bull. (74) U.S.
Fish Wild. Serv., 53, 1953-
Bigelow and Welsh, Bull. U.S. Bur. Fish.
(1924), 40 (1), 1925.
Borodin, Trans. Amer. Fish. Soc., 54, 1924.
Borodin, Copeia, 1927.
Borodin, Bull. Vanderbilt océanogr. Mus.,
I (1), 1928.
Breder, Field Bk. Mar. Fish. Atl. Cst., 1929.
Breder, Amer. Mus. Novit., 382, 1929.
Breder, Copeia, 1942.
452
23.
24.
de Buen, Notas Inst. esp. Océanogr., (2) 88,
1935.
Bull. Bur. Stat. Cons. int. Explor. Mer, 33,
1951.
Ganros, Arqu. Zool. estad. Sao Paulo (1941),
3 (7), 1942.
Coker, et a/., Rep. U.S. Comm. Fish. (1913),
1QI4.
Coker, ef a/., Bull. U.S. Bur. Fish. (1929), 45,
1930.
Cox, Contr. Canad. Biol. (1914-1916), 1916.
Enurenzavm, Nordisches Plankton, 2, 1909
(Eier und Larven Fisch.).
Eigenmann, Mem. Carneg. Mus., 5, 1912.
Evermann, Rep. U.S. Comm. Fish. (1gor),
1902.
Evermann and Kendall, Bull. U.S. Fish Comm.
(1897), 17, 1898.
Evermann and Kendall, Rep. U.S. Comm.
Fish. (1899), 1900.
Fasricrus, Fauna Groenl., 1780.
Forbes and Richardson, Nat. Hist. Surv. Illi-
nois, Ichthyol., 3, 1908.
Ford, J. Mar. biol. Ass. U.K. (1933-1934),
N.S. 19 (1), 1935.
Fowler, Rep. N.J. St. Mus. (1905), 1906.
Fowler, Proc. Acad. nat. Sci. Philad., 63, 1911.
Fowler, Bull. Amer. Mus. nat. Hist., 70 (1),
1936.
40.
66.
67.
Fishes of the Western North Atlantic
Fridriksson and Aasen, Rep. Norweg. Fish.
Invest., 9 (1), 1950.
Grunasure, Bull. U.S. Bur. Fish., 45, 1929.
Goode, Bull. U.S. nat. Mus., 5, 1876.
Goode, Rep. U.S. Comm. Fish. (1877), 1879.
Goode, Proc. U.S. nat. Mus. (1878), 2, 1879.
Goode, e¢ a/., Fish. Fish. Industr. U.S., I (3),
1884.
Greeley, 17th Rep. N.Y. St. Conserv. Dep.
(1927), Suppl., 1928.
Greeley, 24th Rep. N.Y. St. Conserv. Dep.
(1934), Suppl., 9, 1935.
Greer, Rep. U.S. Comm. Fish. (1914), Ap-
pend., 3, 1915.
Gudger, Publ. Carneg. Instn. Wash., 391, 1929.
Gudger, Amer. Mus. Novit., 945, 1937.
Gunter, Publ. Inst. mar. Sci. Texas, r (1),
1945-
E lnecess, Check List Fish. Canada, 1913.
Hansen, Beret. K. Vedeor Gronl., I, 1933.
Hardcastle, J. E. Mitchell Sci. Soc., 57 (2),
194I.
Henshall, Bull. U.S. Fish Comm. (1894), r4,
1895.
Herre and Myers, Lingnan Sci. J., TO, 1931.
Hildebrand, Bull. U.S. nat. Mus., 189, 1946.
Hildebrand, Smithson. misc. Coll., 707 (18),
1948.
Hildebrand and Schroeder, Bull. U.S. Bur.
Fish. (1927), 43 (1), 1928.
Hjort, 5th Annu. Rep. Dep. Naval. Serv.
Canada, 1915.
Hubbs, Proc. Calif. Acad. Sci., (4) 18 (11),
1929.
Hubbs and Lagler, Bull. Cranbrook Inst. Sci.,
18, 1941.
Huntsman, Canad. Fish. Exped. (1914-1915),
1919.
Huntsman, Contr. Canad. Biol. (1921), 1922.
Huntsman, James Johnstone Memor. Vol.,
1934.
Ea Contr. Canad. Biol., 7, 1927.
Jensen, Spolia Zool. Mus. Hauniensis, 9, 1948.
68.
76.
77-
78.
79-
80.
8.
82.
83.
84.
85.
56.
87.
88.
89.
90.
gr.
92.
93.
453
Johnson, Invest. Rep. U.S. Bur. Fish., 2 (38),
1938.
Johnson, J. Fish. Res. Bd. Canad., 4 (5), 1939.
Jordan, Copeia, 1918.
Jordan, Manual Vert. Anim. NE U.S., ed. 13,
1929.
Jordan and Evermann, Bull. U.S. nat. Mus.,
47 (1), 1896; 47 (3), 1898.
Jordan and Gilbert, Proc. U.S. nat. Mus., 5,
1882.
Jordan and Gilbert, Proc. U.S. nat. Mus., 6,
1883.
Jordan and Seale, Bull. Mus. comp. Zool.
Harv., 67 (11), 1926
Ker andepadeliteeP nil WeSy Bur Fish
(1915-1916), 35, 1918.
Lacter and Kraatz, Pap. Mich. Acad. Sci.
(1944), 30, 1945.
Lea, Canad. Fish. Exped. (1914-1915), 1919.
Lebour, J. Mar. biol. Ass. U.K., N.S. 12 (3),
1921.
LeGall, Ann. océanogr. Inst. Monaco, 15,
1935.
Leim, Contr. Canad. Biol., N.S. 2 (1), 1924.
Leim, Progr. Rep. Fish. Res. Bd. Canad.,
Atlant. Biol. St., 1937.
Leim, Bull. Fish. Res. Bd. Canad., 3, 1957.
Linnaeus, Syst. Nat., ed. 10, 1758.
Linton, Bull. U.S. Fish Comm. (1899), Z9,
1gol.
Linton, e¢ a/., Bull. U.S. Bur. Fish. (1904),
24, 1905.
Longley and Hildebrand, Publ. Carneg. Instn.
Wash., 535, 1941.
Lénnberg, Ofvers. Svensk. Vetensk.-Akad.
Forh., 52 (3), 1894.
Liibbert and Ehrenbaum, Handb. Seefisch.
Nordeurope, 2, 1936.
Manrer, Amer. Midl. Nat., 38 (2), 1947-
Marshall, Nicholls, and Orr, J. Mar. biol. Ass.
U.K., N.S. 22, 1938.
McKenzie, Proc. N.S. Inst. Sci., 20, 1939.
Meek, Migrations of Fish., 1916.
4.54
94:
95:
96.
97:
98.
99
roo.
Tol.
I0o2.
I03.
104.
I05.
ro6.
I07.
108.
10g.
IIo.
Tit.
Ir2.
Tg"
TT4.
II5.
Memoir Sears Foundation for Marine Research
Minckley and Krumholz, Zoologica, N.Y.,
44 (4)» 1960.
Moore, Rep. U.S. Comm. Fish. (1896), 22,
1898.
Myers, Copeia, 1929.
Myers, Copeia, 1950.
Nicuots, Bull. Amer. Mus. nat. Hist., 37
(18), 1912.
Norman, Ann. Mag. nat. Hist., (9) IZ, 1923.
Norman, Ann. Mag. nat. Hist., (9) 12, 1923.
Norman, Hist. Fishes, 1931; and subsequent
editions.
Norman, “Discovery” Rep., 16, 1937-
Pearson, Invest. Rep. U.S. Bur. Fish., Z (10),
1932.
Perley, Rep. Sea and River Fish. N. Brunsw.,
1852.
Prince, 39th Annu. Rep. Dept. Mar. Fish.
Canada, Suppl., 1907.
Rccan: Ann. Mag. nat. Hist., (8) 19, 1917.
Richardson, Bull. U.S. nat. Mus., 54, 1905.
Rivas, Smithson. misc. Coll., 206 (14), 1946.
Rivas, Proc. U.S. nat. Mus., 200 (3263),
1950.
Rose, Proc. gth Annu. Sess. Gulf Caribb.
Inst., 1956.
Rounsefell, Bull. U.S. Bur. Fish. (1929), 45;
1930.
Ryder, Rep. U.S. Comm. Fish. (1881), 1885.
Ryder, Rep. U.S. Comm. Fish. (1885), 1887.
Scarrercoop, Copeia, 1953.
Scattergood, Res. Bull. Sea Shore Fish. Maine,
26, 1957.
Ir6.
TT7.
riés.
Ig.
120.
I2f.
T22.
123.
124.
125.
126.
PGT
r28.
129.
130.
I3I.
go.
igo:
134.
135.
136.
Ta.
Schmitt, Monogr. Isle d’Anticosti, 1904.
Sindermann and Rosenfield, Res. Bull. Dept.
Sea Shore Fish. Maine, 18 and 21, 1954.
Smith, Bull. U.S. Fish Comm. (1890), Zo,
1892.
Smith, N.C. geol. econ. Surv., 2, 1907.
Smith, Bull. U.S. Fish Comm. (1897), 27,
1898.
Stearns, Proc. U.S. nat. Mus., 6, 1883.
Stevenson, Rep. U.S. Comm. Fish. (1898),
1899.
Sumner, Osburn, and Cole, Bull. U.S. Bur.
Fish., 32 (2), 1913.
"Tarzor and Sykes, Bull. U.S. Fish. Wildl.
Serv., 142, 1958.
Thompson, Rep. Newfoundld. Fish. Res.
Comm., I (1), 1931.
Trautman, Fishes of Ohio, 1957.
Viabynoy, Nat. Canad., 73 (1, 2), 1945.
Vladykov, Nat. Canad., 77, 1950.
Vladykoy and McKenzie, Proc. N.S. Inst.
Sci., T9 (1), 1935-
Vladykov and Wallace, Trans. Amer. Fish.
Soc. (1937), 67 (2), 1938.
Wy craomre Copeia, 1940.
Welsh and Breder, Bull. U.S. Bur. Fish.
(1923), 39, 1924.
Willey, Contr. Canad.
1923.
Wilson, Proc. U.S. nat. Mus., 39, 1911.
Wilson, Proc. U.S. nat. Mus., 53, 1917-
Wilson, Proc. U.S. nat. Mus., 64, 1925.
Wilson, Bull. U.S. nat. Mus., 758, 1932.
Biol., N.S. 1 (16),
Suborder Salmonoidea
COMPOSITE AUTHORSHIP?
Characters and Key to Families
HENRY B. BIGELOW
Museum of Comparative Zoology
Harvard University
Characters. Isospondylous fishes with Petvic Frns abdominal in position, and with
an ApIposE FIN between the rayed dorsal and caudal fin.
Scates. Body and tail sectors of trunk with scales everywhere in most, but naked
in some Salangidae;? head naked; scales thin, rounded. Luminescent orcans (photo-
phores) absent.
Maxitiary bones as well as PRemaxiLiaries taking part in formation of upper
jaw. Teer of moderate size in most, but those on lower jaw in the form of canines in
a few, perforating the upper jaw; teeth present on premaxillaries, maxillaries, palatines,
vomer (except in some Salangidae), dentary portion of lower jaw, and tongue in most;
teeth small or absent among the Coregonidae.
PECTORAL GIRDLE with a mesocoracoid element in most; lacking in the Salangidae.
Last few VERTEBRAE upturned in Salmonidae, Coregonidae, and Thymallidae; a single
vertebral centrum turned upward in the Osmeridae, but conditions in this respect seem
not to have been reported for the Salangidae.
A Swim BLADDER in most, connected to the alimentary canal by a permanently
open duct; no swim bladder in Salangidae. Ovipucts incomplete insofar as known,
1. Characters and Key to Families by H. B. Bigelow; Family Salmonidae and Genus Salmo by J. R. Dymond; Genera
Salvelinus and Cristivomer by H.B. Bigelow; Family Coregonidae by H.B. Bigelow; Family Osmeridae by
H. B. Bigelow and William C. Schroeder.
2. The Salangidae are described (46: 239) as “naked, or with a few exceedingly thin, large, scattered deciduous scales.”
For illustrations of the salangid dentition, see Fang (46: 240, fig. 1; 246, fig. 3; 252, fig. 8; 260, fig. ro).
455
456 Memoir Sears Foundation for Marine Research
being represented by a fold of the peritoneum. Stomach with Pytoric cazca except in
Salangidae.
Remarks. For discussion of the characters that distinguish the Salmonoidea from
other Suborders of the Isospondyli, see Key to Suborders (p. 95) and related text.
Families. Of the six families recognized in the following Key, three are represented
with detailed accounts in this series: Salmonidae (p. 457), Coregonidae (p. 547), and
Osmeridae (p. 553). The Thymallidae (graylings), Plecoglossidae (ayu or sweetfishes),
and Salangidae (icefishes) occur in northern waters of western North America, Europe,
Siberia, northern China, Korea, and Japan. Many of the species are anadromous, as-
cending freshwater rivers to spawn.
Key to Families
1a. A fleshy axillary lobe on each side close above base of pelvic fins.
2a. Rayed dorsal fin, when depressed, shorter than head.
3a. Teeth well developed on jaws, on roof of mouth, and on tongue; at least
19 scales in a transverse series from origin of dorsal fin to lateral line;
parietal bones of skull not meeting in the midline. Salmonidae, p. 457.
3b. Teeth minute or lacking; not more than 13 scales in a transverse series
from origin of dorsal fin to lateral line; parietal bones of skull meeting
in the midline. Coregonidae, p. 547.
2b. Rayed dorsal fin, when depressed, much longer than head.
Thymallidae.
Mostly in clear, cold streams, seldom
entering salt water; western North
; ; America, northern Europe, and Siberia.
1b. No axillary lobe above base of pelvic fins. , sei
4a. Head only slightly depressed, its depth at eyes 67 °/) or more of body depth
at dorsal fin; snout in front of eyes at least no longer than base of dorsal fin;
body not transparent; branchiostegal rays 6-10; swim bladder present;
stomach with pyloric caeca.
sa. Jaw teeth firmly and directly attached to jaw bones; tongue not pleated;
not more than 10-12 pyloric caeca. Osmeridae, p. 553.
sb. Jaw teeth arranged on serrated plates on inner sides of jaws, not directly
attached to jaw bones; tongue pleated; up to 400 pyloric caeca.
Plecoglossidae.*
Running up cold streams to spawn;
coastal waters of Japan and Formosa.
4b. Head strongly depressed, its depth at eyes less than 50°/, of body depth at
dorsal fin; snout in front of eyes about twice the length of base of dorsal fin;
body transparent; branchiostegal rays only 3 or 4; no swim bladder; no
pyloric caeca. Salangidae.*
Ascending rivers to spawn; coastal waters
of Japan, Korea, and northern China.
3. Only known genus, Plecoglossus. 4. Contains only one genus, Salanx.
Family Salmonidae
JOHN R. DYMOND
University of Toronto
Acknowledgment. I am indebted to Henry B. Bigelow for much help in the prepara-
tion of the manuscript, especially concerning conditions in New England and in the
matter of angling.
Characters. Mourn large. Maxitiary reaching to a vertical through the eye or
beyond. Trrru well developed on jaws, vomer, palatines, and in a double series on the
tongue. ParieTALs not meeting in the midline. OrsrrospHENOID, BasIsPHENOID, and
SUPRAOPERCULAR present. Last few VerTEBRAE upturned. Dorsat fin with not more
than 16 rays, 12 or less of them branched. ApiposE porsat fin present. AXILLARY
PROCESS associated with ventral fin. Swim BLADDER connected to alimentary canal
by a permanently open duct. Ovipucts incomplete, being represented by a fold of
the peritoneum. Stomach with Pytoric cagca.
Sexual Dimorphism. This phenomenon occurs in the entire family, especially at
spawning time, with an extreme condition in Oxcorhynchus to comparatively little in
Cristivomer, Salmo occupying an intermediate position. The extent to which these changes
affect the two sexes is discussed in the account of the Atlantic Salmon (p. 463).
Remarks. For characters that separate the family Salmonidae from other families
of the Suborder, see Key to Families (p. 456) and related text.
Range. The Salmonidae, composed of six genera, is an important and widely
distributed family of the Arctic and north temperate zones of the northern hemi-
sphere. Many members are anadromous, living in both fresh and salt water, and
others are confined to only freshwater lakes and rivers (see Key to Genera, p. 458).
The species of Sa/velinus and Salmo are found in both marine and fresh waters of
northern North America, Europe, and Asia. The various species of Oxcorhynchus are
confined primarily to the Pacific, but one or two species range through the Bering
Strait and then for limited distances eastward and westward along the Arctic coasts of
North America and Asia. Three genera are virtually confined to fresh water: Cristi-
vomer of North America, Brachymystax of Asia, and Hucho of Europe and Asia; being
457
458 Memoir Sears Foundation for Marine Research
essentially freshwater fish, they apparently have never been able to spread through salt
water from one continent to another. Members of Sa/velinus live the farthest north,
with S. a/pinus ranging well within the Arctic Circle. [For the ranges of the species
discussed, see pp. 490, 499, 521, 539, and 543.]
© ©
Friceure 118. Outline of caudal fin: Sa/velinus al- Figure 119. Outline of side of head: Sa/velinus
pinus (left); Cristivomer namaycush (right). alpinus (left), Brachymystax (right), illustrating dif-
ference in shape of lower margin of upper jaw
(maxillary bone).
Key to Genera
1a. Lower margin of sides of upper jaw (maxillary bones) weakly and uniformly
concave in outline (Fig. 119).
2a. Anal fin usually with fewer than 13 rays.
3a. A double or zigzag series of teeth on shaft of vomer along midbelt of
roof of mouth, but sometimes lost in adult; sides with black markings.
Salmo Linnaeus 1758, p. 459.
3b. Teeth absent on shaft but present on head of vomer; sides without black
markings.
4a. Teeth on the vomer usually separated from the teeth on the pala-
tines.
sa. Caudal fin only moderately concave (Fig. 118); sides with red-
dish or salmon-colored spots; not more than 50 pyloric caeca.
Salvelinus Richardson 1836, p. 503.
sb. Caudal fin deeply concave (Fig. 118); sides with pale whitish
spots; about 140 pyloric caeca.
Cristivomer Gill and Jordan 1878, p. 542.
4b. Teeth on the vomer connected in an unbroken series with the teeth
on the palatines. Hucho Giinther 1860.
Rivers of central and eastern Europe,
northern Asia, and Japan.
2b. Anal fin with 13-19 developed rays. Oncorhynchus Suckley 1861.
Both sides of northern North Pacific.®
s. The Humpback Salmon (O.gorbuscha) has been introduced in several rivers of Maine and for a time it seemed that
it was established there, but it failed to maintain itself. The Silver Salmon (O. isutch) also has been planted in Maine,
but without result. For further details see Bigelow and Schroeder (z8: 131~-133)-
Fishes of the Western North Atlantic 459
1b. Lower margin of sides of upper jaw (maxillary bone) strongly convex in outline
(Fig. 119). Brachymystax Giinther 1860.
Rivers of Siberia.
Genus Salmo Linnaeus 1758
Salmon, Trouts
Salmo Linnaeus, Syst. Nat., I, 1758: 308; type species, Sa/mo sa/ar Linnaeus, established by Jordan and Gilbert,
Bull. U.S. nat. Mus., 76, 1882: 309. European Ocean.
Generic Synonyms:
Trutta Geoffroy, E. L., Descr. 719 Plantes, 134 Animaux., Paris, 1767: 719.
Fario Cuvier and Valenciennes, Hist. Nat. Poiss., 27, 1848: 227; argenteus equals frutta.
Salar Cuvier and Valenciennes, Hist. Nat. Poiss., 22, 1848: 294; ausomii equals fario.
Species. Salmo, the type genus of the family, contains the species originally named
THE Salmon (8S. sa/ar) and THe Trout (S. ¢rutta). In addition to THe Trout, other black-
spotted trout closely related to it are included. These occur chiefly in Europe and
North America; a few are found in areas that may be regarded as extensions of the
main centers of distribution.
Some isolated landlocked populations have diverged so far from the presumed
parent stocks as to constitute distinct species. In the case of others, the question of
their taxonomic status is still a matter of doubt. This has led to uncertainty as to how
many species should be recognized. Three species fall within the scope of this book:
S. salar, the Atlantic Salmon, which is anadromous and spawns in fresh water (see
also Freshwater Populations, p. 491); S.trutta, the European trout, an introduced
species that is anadromous in Newfoundland and is called the brown trout in America
(p. 498); and S. gairdneri, the rainbow trout, which is indigenous in northwestern
America but which has been widely introduced and is well established in the north-
eastern United States and eastern Canada, occurring mostly in fresh water but being
anadromous locally (p. 499).
Key to Species of the Western North Atlantic
1a. Anal usually with 9 fully developed rays; caudal usually unspotted.
2a. Vomerine teeth little developed, those on shaft few and deciduous; scales
from adipose fin to lateral line, 10-13; maxillary extending to below rear
edge of pupil at length of 150 mm, never far behind in large adults; caudal
weakly concave in adults; black spots on body small, without halos.
salar Linnaeus 1758, p. 460.
2b. Vomerine teeth well developed, those on shaft of bone numerous and per-
460 Memoir Sears Foundation for Marine Research
sistent; scales from adipose fin to lateral line, 12-16; maxillary extending to
below rear edge of eye at 150 mm, farther back in large specimens; caudal
truncate in adults; black spots on body large, many surrounded by halos.
trutta Linnaeus 1758, p. 498.
1b. Anal with 10 or more fully developed rays; caudal spotted.
gairdneri Richardson 1836, p. 499.
Salmo salar Linnaeus 1758
Atlantic Salmon, Sea Salmon
Figures 120-123
Study Material. Ten fresh specimens (7 males, 3 females), about 71-90 cm or
28-35 inches SL, from the Matamek River, Saguenay County, Province of Quebec,
caught towards the end of July 1931 (not preserved); also 87 specimens, fry to
adult, from New Brunswick, Nova Scotia, Newfoundland, and Quebec (including
Ungava Bay).
Distinctive Characters. Salmo salar is distinguished from all species of Oncorhyn-
chus of the Pacific in having fewer than 13 rays in the anal fin (13-19 in On-
corhynchus).
It is readily distinguished from Salvelinus alpinus and fontinalis (Arctic charr
and brook trout) and from Cristivomer namaycush (lake trout) by its black spots (light-
colored in charrs) and by its larger scales (so small as to be readily overlooked in
charrs).
It is separated from Salmo gairdneri (rainbow trout) by the absence of black
spots on its caudal fin (many spots on caudal of gairdneri).
Separation of Salmo salar from Salmo trutta (brown trout) is more difficult than
from the above species, for both sa/ar and sruita lack spots on the caudal fin. How-
ever, in salar the tail is more constricted at the base of the caudal fin than in srutta,
so that in sa/ar the anterior caudal rays form more of a shoulder; consequently, an
Atlantic Salmon does not slip through one’s fingers when grasped round the caudal
peduncle as a brown trout usually does. This difference is reflected in the number of
scales in an oblique row from the posterior edge of the adipose fin (downwards and
forwards) to the lateral line: 1o—13 (usually 11) in sa/ar, 13-16 (rarely 12) in ¢rutia.
Other differences include: dorsal fin rays 10-13 (usually 11) in salar, 13-16 (usu-
ally 14, rarely 12) in srutta; pyloric caeca 50-80 in salar, 30-60 in srutta. The caudal
fin of salar is more widely spread and is more deeply emarginate than that of srusta.
Finally, salar has a smaller mouth, the maxillary in the adult extending scarcely, if
at all, beyond the eye (well beyond the eye in srutra).
Jones (77) has tabulated the differences between the young Atlantic Salmon and
brown trout as follows:
Fishes of the Western North Atlantic
YOUNG ATLANTIC SALMON
461
YOUNG BROWN TROUT
BODY) ass 18 «sient sia ae Usually thinly spotted. Usually much spotted.
Caudal peduncle, depth One-tenth of length of body. One-eighth of length of body.
Operculum .......... Rarely more than three spots. Usually more than three spots.
Maxilla, at length of:
AINCHES See ess ts To center of pupil. To posterior edge of pupil.
Grinc hes mh mseaer. eA). To posterior edge of pupil. To posterior edge of iris.
DORSAL jie ce. se Usually weakly spotted. Usually well-defined spots.
PIGIPOSE Piha ws oes «= Rarely edged with red. Usually edged with red.
UM ED RUS an Ces ox Generally grey. Generally orange-yellow.
187 inst are RA ae Deeply forked; middle rays Shallowly forked; middle rays
not more than 0.6 of length = 0.6 or more of length of
of longest rays. longest rays.
PGPETBAERS stein aol Usually 11-12. Usually 9-10.
It is possible that hybridization may occur between salar and ¢rutta, in which
case the hybrids probably would not possess the distinguishing characters of either
parent.
Description. The adult Atlantic Salmon is a graceful fish, deepening rearward from
a small pointed head to the deepest point under the dorsal fin, then tapering to a slender
caudal peduncle which supports a spreading and slightly emarginate caudal fin. The
mouth is moderately large, with the posterior end of the maxillary extending to, or
slightly beyond, a vertical through the posterior margin of the eye.
Proportional dimensions in per cent of standard length (averages, followed by
ranges in parentheses) of the 1o fresh specimens listed in the Study Material, 28-35
inches (ca. 71-90 cm). Note: Shape and length of head and depth of body varying
with stage of sexual maturity.
Mates FEMALES
Boay-spreatest deptlcl: Seelgne setts cite). Ss 21 (20 —22) 22.6 (21.424)
Sreatest widthivaeises seat a. fas: 12.2 (11.4—13.3) 12.4 (11.3-13.2)
Caudalepeduncle:sdepthi: Sass aes eles 7.9 ( 7.6— 8.6) 8.0 ( 7.0- 8.6)
Plead Mength: Vente ei Moe ee CAMEOS ones 23 (20.8-2 5.9) 20.7 (20.0—21.2)
SHOUT MIENOTE cters as ccsere eee a eres os 9.2 ( 7.2—11.1) Fae Os 77 5)
ive: diameter Of eyeballiyrcn.eiqaaehy <lesst 2.9 ( 2.7— 3.0) 2.8 ( 2.7— 2.9)
Maxillary length) ities soins aes es 12.5 (10.7—14.4) 10.5 (10.0—10.8)
Dorsal fin: longest raywataeea ewe see 3s 11.7 (10.5—13.5) 11.6 (11.3-11.9)
Winal fist: NON Best Gay vein «csi lel Pe slss 10.6 (10.2—11.0) 10.5 (10.1-11.1)
lengthiofsbases. givains meek + 2 9.5 ( 8.9— 9.7) 9.2 ( 8.0—-10.4)
Pelvicifinelengthyen <u). hei. netae Pelee sate 11.0 ( 9.9—11.6) 10.7 (10.5—11.2)
Pectoral jin> lenpth: =e. aac dear nae 13.3 (12.1-13.8) 12.97 (12).5—L3.1)
462 Memoir Sears Foundation for Marine Research
Boru SEXES
Scales: diagonal rows to end of vertebral column 113 (110-118).
Branchiostegals: 11 or 12.
Gill rakers: 18 (17-20).
Dorsal rays: Usually 10 branched rays preceded by one full-length unbranched ray and
one (occasionally two) shorter unbranched rays.
Anal rays: 8 or 9 unbranched rays preceded by one full-length ray and one (occa-
sionally two) shorter unbranched rays.
Ficure 120. Salmo salar, 37.8 inches long, probably from Delaware Bay. After Goode; posterior outline of
fleshy caudal peduncle somewhat emended.
Pyloric caeca: §5.4 (40-74) for Salmon taken in or near the mouth of the Miramichi
River, Quebec (14).
Vertebrae: 58.5 (58-61) for 23 specimens from the Moose River, N.S.
Color. The color of an adult Atlantic Salmon depends on whether it is in salt or
fresh water. When it is at sea, its back is blue-green overlaid with a silvery coating, the
sides are silvery, and the belly is pearly white. There are scattered black, x-shaped, and
round spots on the sides, usually above the lateral line and occasionally below it anteri-
orly; a few round black spots are present on the sides of the head. The dorsal fin is
dusky, with the lower half lighter. The caudal is dusky (darker posteriorly), usually
without spots; and the anal is gray with whitish rays. The pelvics (ventrals) are light-
colored, with a few of the anterior membranes dusky, especially above; the pectorals
are dusky with a whitish base.
When the Atlantic Salmon re-enters fresh water, it loses the silvery guanin coat
and becomes greenish, or reddish brown mottled with red or orange (more especially
in the male), and has large dark spots edged with whitish color.
The parr of this species (p. 467) is bluish or purplish above and has a series of
8—11 or 12 vertical dark crossbars or “‘parr marks”’ along each side; also there are
blackish spots chiefly above the lateral line anteriorly and on the sides of the head, a
red spot between each pair of parr marks, and usually a few other spots.
Fishes of the Western North Atlantic 463
The smolt (p. 467) is silvery due to the deposition of light-reflecting spicules of
guanin in the epidermis, covering the bars and spots of the parr so that it resembles
the adult Salmon at sea.
Changes in Spawning Fish. As spawning approaches, the head (snout and lower
jaw particularly) elongates, and the lengthening of the lower jaw is greater than that
of the skull itself; and in the male the tip of the lower jaw turns upward to form
a prominent hook that fits (when the mouth is closed) into a hollow of the upper jaw.
These changes are more pronounced in the male than in the female, and in both sexes
they are relatively much greater in large specimens than in small ones (127). After
spawning is completed, in those that live, these changes gradually disappear and the
features of the head resume their normal shape and proportions.
Ficure 121. Sa/mo sa/ar, smolt stage, 140 mm TL, from Margaree Harbor, Cape Breton, Nova Scotia, ROMZ
9462. Drawn by Jessie H. Sawyer.
The teeth also undergo very marked changes with each breeding period (126).
The “feeding teeth” are replaced by special “‘breeding teeth” when the fish moves in-
shore towards the river. These are relatively shorter and broader at their bases than the
feeding teeth, and on some bones they are strongly curved; when fully grown in large
males, the breeding teeth are securely fastened to the bones. When spawning is com-
pleted they are apparently replaced again by feeding teeth. In specimens of equal size,
the number of breeding teeth on the maxillaries is higher than the number of feeding
teeth. In the large breeding males examined, the vomerine teeth on the anterior part
of the shaft were in a transverse row. Tchernavin’s observations are supported by those
of Rushton (zz8), who found that well-mended kelts (p. 467) had large teeth, with
small teeth beginning to grow between them. The large teeth showed signs of decay
at their bases and looked as though they would drop out before long. In addition to
the striking changes in head and teeth, the skin of the back becomes thick and spongy
and the scales are embedded.
Size. This species grows larger in European waters than in the western Atlantic,
where very few reach 50 pounds (p. 478). The largest ones that have been caught
in America by angling were a 55-pound fish taken in the Grand Cascapedia River,
Quebec, on July 27, 1939; and two 54-pound fish, one taken there on June 20, 1886
(735: 50), and the other in 1921.® In some of the larger rivers of Newfoundland and in
6. See photo in Atlantic Salmon Journal, Sept. 1959: 24.
4.64 Memoir Sears Foundation for Marine Research
those emptying into the Gulf of St. Lawrence, fish of 40 pounds were not uncommon
years ago, but today an Atlantic Salmon of that size is rare.
The record recognized by the International Game Fish Association as the heaviest
Atlantic Salmon taken by rod and line is a 79-pound 2-ounce fish caught at Tanalev,
Norway, in 1928; also, in 1921, a 74-pound fish was taken in Norway on a fly
(135: 3,4). An 83-pound fish has been reported as being netted in England (78), and
an 84%/,-pound fish in Ireland in 1881 (83). There are reports also of “a one-hundred-
and-three pound, two ounce salmon taken in a net on the Devon River in Scotland”
(148: 23), and of others weighing over 100 pounds (133: 56); however, records such
as these are to be accepted only with extreme caution.
Size-W eight Relationship. The accompanying graph (Fig. 122), based on records for
specimens from several localities, both American and European, is presented as a rough
over-all average of length and weight for the species. A curve for any one locality
would probably differ to some extent from that for any other.
Spawning and Number of Eggs. Atlantic Salmon, following their early growth in the
river and one or more years at sea (see also Spawning Survival, p. 479), return to fresh
water to spawn (p. 473), the spawning time usually occurring in fall or winter according
to locality. In Canada generally, actual spawning occurs in late October and early
November, but in Greenland, near Kapisigdlit in Godthaab Fjord, it is said to occur
in early spring (52). In northern Europe they may spawn as early as September (in
Iceland still earlier) or as late as January, or even later; thus in some Scottish rivers
they do not spawn in numbers until after New Year’s Day (roo), while in southern
England spawning has been observed as late as the third week in March, mainly in the
lower reaches (122). See also Coastal and River Spawning Runs (p. 475).
Any place where the conditions are favorable may be used as a spawning site—
from just above tidal waters to the upper reaches of a river, sometimes as much as
200 miles or more from the sea. The depression, or so-called ‘“‘redd,” in which the
eggs are deposited, is made in shallow sections of the stream where the current is
swift. The most suitable bottom is a coarse gravel, with many stones from two to eight
inches in size, that allows well-aerated water to penetrate. Loose gravel or sand is un-
suitable because of its shifting nature (Zo).
The spawning act has been described repeatedly, but the most careful observations
are those by Jones and King, first made in 1947 and confirmed by them in 1948 (79;
80; 78). For their studies they built a glass-sided observation tank on the banks of the
River Alwen, a tributary of the Welsh Dee. From this they made lateral observations
and took photographs that could be studied later.
Before depositing her eggs, the female cuts a depression in the gravel with her
tail and feels the bed with her anal fin. The male takes no part in making the bed and
is often not present, but he courts the female and drives away all other fish.
Just before the spawning act, the female drops into the bed, opens her mouth,
erects her fins, and bends her tail upwards. Upon this the male immediately joins her
and settles alongside, quivering, with his mouth agape. At this instant the pair spawns
Fishes of the Western North Atlantic 465
almost simultaneously. The complete operation, observed in the case of a nine-pound
female and a five-pound male, consisted of eight emissions in different beds prepared by
the female. After each spawning act, the female moved upstream of the bed and cut
vigorously with her tail, thus covering the eggs with gravel. Each spawning act took
place in 10-15 seconds, and the eggs were completely covered about 30 seconds later.
ae lameal th ie ; i aie
Set = a =|
INCHES
LENGTH IN
| l L ! Jo).
: a a | | 1 tt 1
5 LOM—I Sic Omne Sas On sD) 40 N45 Om SoG Ol 65.070 iy sO
WEIGHT IN POUNDS
Ficure 122. Length-weight relationship for Atlantic Salmon.
In the experimental tank, spawning took place at temperatures varying between
36—42°F. Of the eggs collected from 12 nests, 97.6 °/, had been fertilized.
An interesting feature of the spawning of the Atlantic Salmon is the participation
in it of male parr. That some parr become sexually mature before they leave the river
has long been known. As early as 1686, Willughby wrote that while female parr (salm-
olus) never have ripe eggs, male salmolus fertilize the eggs of the adults (746). These
early observations have been amply confirmed in recent careful studies by Alm (3) and
others (80; 81; 82; rrr). According to Jones, 75 °/, of the male parr become sexually
mature during river life (78: 128). Their sperm is as effective in artificial fertilization
of the eggs as is the sperm stripped from adults. Eggs fertilized by parr sperm
produce normal progeny, and male parr under natural conditions may be responsible
for fertilizing a considerable fraction of the total number of eggs that develop. It has
30
466 Memoir Sears Foundation for Marine Research
been suggested furthermore that the presence of parr is a safety factor. The spawning
males may fail to get their vent deep enough into the bed to have their sperm fertilize
most or all of the eggs. Under such circumstances, the parr are of importance since
their sperm, released very close to the eggs, have a good chance of reaching them.
The number of eggs produced varies with the age and size of the females, as shown
in Table 1. Menzies has given a range of 600-1000 eggs per pound of spawning female
Table I. Average Number of Eggs Produced by Canadian Atlantic Salmon *
r—— First Spawning After ——. Salmon That Have
2 Years at Sea 3 Years at Sea Spawned Before
Numbers fe temalesmeerirstectrstrciclell- 503 15 16
Average weight of female (pounds) ... 10.51 19.13 16.68
Motallmumberjofleges . 2.) ++ -9-- > 8,848 +68 13,883 +483 12,313 +648
Eggs per pound of fish..........-..- 83445 723413 738 + 34
* From Belding (r4).
(r00). Jones and King estimated from the number of eggs recovered from redds made
by their nine-pound experimental female that she had laid 700 eggs per pound of body
weight (79). Large females usually have larger eggs than small ones (14), but they are
fewer (100).
Although Belding found a definite but slight difference in the number of eggs per
pound of fish between females of the Miramichi River (817 fish) and coastal females of
the Bay-of-Chaleur type (854 fish), this difference was not sufficient to warrant its use
in identifying river varieties of Atlantic Salmon at sea (I4).
After being deposited and during incubation, the eggs of the Atlantic Salmon
are subject to many vicissitudes, such as being dislodged by other spawning Salmon
in overcrowded streams, being washed out by floods, and being frozen in the winter
at times of low water.
Incubation Period. The large, thick-shelled eggs, about 6—7 mm in diameter, de-
velop slowly over the winter and hatch in the spring at various times depending on
the temperature and on other conditions at the spawning sites. In hatching experiments
with eggs from specimens of different ages taken in or close to seven rivers of the
three Maritime Provinces of Canada, Belding, et a/. (6) found that the date of hatching
varied from March 15 to May 4 and that the location of the hatchery in which the
eggs were incubated seemed to be an important factor, for it took place on 111/15 at
Prince Edward Island, on 111/21 and 1v/25 at Middleton, N. S., on 1v/11-19 at Mar-
garee, N.S., on 1v/20 at St. John, N.B., on 1v/23 at Bedford, N.S., on 1v/26 at
Antigonish, N. S., and on v/2—4 at Restigouche and Miramichi, N. B. They observed
further that the water temperature “between 34 and 41°F. is not a prominent factor
in determining the length of the incubation period,” whereas temperatures above that
decrease the incubation time markedly; also the water temperature at the end of the
incubation period may determine the hatching date, since the temperatures in the early
hatcheries were only slightly lower at hatching (35°-38°F) than in the late hatcheries
Fishes of the Western North Atlantic 467
(41°-43°F). In another hatchery experiment, hatching occurred much sooner at 42°F
(88 days) than at 33° (191 days).
Names Applied to the Successive Life Stages. The Atlantic Salmon at various stages
of growth has been given the following distinctive names:
Atrvin: the stage from hatching to absorption of the yolk sac, which occurs at a
length of about one inch; often called the yolk-sac stage.
Fry: the stage between absorption of the yolk sac and acquisition of the parr marks.
Parr: the young stage spent entirely in fresh water when it has 8-11 or 12 dark
vertical bars or parr marks on each side, acquired soon after the disappearance of
the yolk sac. Some male parr mature during this stage but the females do not, so far
as is known.
FINGERLING: same as parr.
Smott: A young fish that has acquired a coat of guanine that hides its parr
marks and gives it a shiny, silvery appearance. This change usually occurs after the
fish has spent one to four years, and exceptionally as much as seven years, in the
natal river. Following this transformation the young Atlantic Salmon migrates to the
sea (pp. 468-470).
GritsE: a mature fish that returns to fresh water to spawn after one complete
year at sea; distinguishable from the older fish by its more deeply forked caudal fin
and its smaller size; its average length in Canada is 18-24 inches, its weight 3-5
pounds, with extreme ranges of 2-14 pounds reported.
SaLMon: a mature fish that has spent two or more years in the sea before returning
to fresh water to spawn for the first time; its weight with two years of sea-life, 6.5—16
pounds (usually 10-12 lbs.), and with three years, 20-35 pounds or more; most of
the large fish, 40-50 pounds, are females that have been at sea a long time and have
spawned only once or not at all.
Ketr: a thin adult that has spawned and still shows the dark color assumed
during its freshwater stay; often called a black salmon.
Menopep Ketr: a fish that has recovered from the kelt period by regaining its
silvery color and a more normal plump form.
The Alevin Stage. The newborn alevin, 15—20 mm long, are hatched with a mass
of yolk which nourishes them but which also weighs them down and keeps them from
maneuvering freely; during this time they usually remain on the river bed. The yolk-sac
period for hatchery-reared alevin varies between 29—65 days (usually 39—53 days), with
temperature the controlling factor; as in the hatching of the eggs, the higher the temper-
ature the shorter the yolk-sac period (r6). Vibert’s studies (131) have confirmed earlier
conclusions that alevin hatched under gravel are hardier than those reared in a hatchery,
are better developed, and average 15 °/) heavier. With the disappearance of the yolk, the
little Salmon must begin to seek their own food. They are then gregarious (78), resting
on the stream bed with others and rising to take prey, either individually or with a group.
From Parr to Smolt. Soon after the disappearance of the yolk, which marks the end
of the alevin stage, the young fish develop the characteristic crossbars along each side
30”
468 Memoir Sears Foundation for Marine Research
that distinguish them as parr (p. 467), and they begin to acquire scales. As they
grow, they seek a new habitat, since the area that provided them with food and shelter
when they were younger is no longer adequate. Thus each one establishes a more or
less definite territory of its own as described by Katteberg (85). The behavior pattern
has been described by Huntsman as follows (70): “salmon ... may become related to
particular local environments which may be called their homes. Such relation may
involve having a place or places for rest on the bottom, for stationing in a current,
and for cover, with more or less roaming as well as precise dashes in taking food.”
That parr move about in the parent stream now seems established. Huntsman
(67: 399) found that freshets may greatly alter or destroy their home, and that this
and other factors induce or force the parr to descend from their spawning area to lower
waters of the parent stream, there to populate lakes as well as estuaries that have
fresh or brackish water (but not full sea water, which kills parr; 74: 409). Some may
reascend varying distances upstream and populate places that are accessible; falls as
low as one foot seem to stop parr less than a year old.
Subject as they are to many vicissitudes and to different environmental condi-
tions throughout their range, the populations of young vary widely at all times in abun-
dance as well as in size and growth relative to age (dependent mainly on food and
temperature) and hence in time of smoltification, not only from one general area to the
next and from one river to another in the same general area in different years, but even
from year to year in the same river. Such data as are available indicate that parr
throughout the Atlantic Salmon’s range may spend anywhere from one to seven years
in the river before they become smolts, and that, generally, parr life is longer in northern
than in southern regions (Table 1). Most of the parr in America spend two or three
years in the river, but longer northward. In England and Scotland one- or two-year
smolts are the rule. In southern Norway it is usually three years, but northward, five-,
six-, or even seven-year smolts are common.
In general, the fastest growing parr develop the soonest into smolts, but little is
known about the factors that initiate and control smoltification. According to Pye-
finch (116: 9):
Smoltification is in some way connected with size, but the nature of this relationship is by no means direct.
... It seems possible that rate of growth may be more important than the mere attainment of a critical length.
Reference has already been made to the observation that growth during the first year is an index of growth later
in life. Therefore, assuming that the internal changes which occur during smoltification are seasonal in their
incidence, a rapidly growing fish may have developed far enough for these changes to take place when it is only
a little more than a year old, whereas a fish which grows more slowly is not sufficiently well-developed for these
changes to take place then and so must wait until the following spring.
Elson concluded that, as a general rule, parr that have reached or exceeded a
certain size towards the end of one growing season are likely to become smolts at the
next season of descent (43: 5). Those that have nearly but not quite reached a length
of rocm at the end of the summer will, if they grow fast, reach a much greater length,
say 14—16 cm, at the end of the following summer, and make big smolts. Those that
Fishes of the Western North Atlantic 469
Table II. Percentage Composition of Smolt Population Based on Years Spent as
Parr in Fresh Water
(ee ANS i WE
Nortu AMERICA ; 2 3 4 5 6 7
Penobscot R., Maine (87) .......-. wits majority few
Pollett R., Petitcodiac R. system,
ING BS (CBS 2) eddesodacoossscasdbn mere 90-95 5-10 sir
Marparee Re vINaS:0 (Tz) raecleiersre ers 4 68 25 3
Miramichi R. system, N.B. (43: 1). at: 6a.33 + ca.67 ee is
MiramichiiRe.sN@B21(70))ecmtesvli-lrs afar 15.1 78.1 6.6 0.2
Grand Cascapedia R., Que. (29)... 506 6 58.8 34.1 I.I
Mioisie Re) Ouen (704) * ete ei 0.35 34.45 61.3 3-9 aan
Six west coast rivers of Newfound-
lEnGl (GGA) ce kowsdousonoonoosconoe ae Wel 57-4 29.8 5-4 0.3
East coast rivers of Newfoundland,
including Labrador (20).......... fers 3.4. 26.0 39.1 25.3 cy) 0.5
Greenland!(3))saaece cece sen ace Rohs ae bo 42.8 52.4 4.8 a
Europe
Avon R., Hampshire, Eng. (700)... 60 ae si
Shannon R., Eire (136, 137)T.---- 12.8 79.1 8.1 30 cists
Scotlandu(00) pratense ir scier: few mostly 2—3 years few rare
Rivers of southern Scotland (r00) .. some gOormore some
Tweed R., Scotland (100)......... ane 97 arc ce ate
Norway — Kristiansund distr. (35)tT ae 25.8 64.5 9 0.7
Trondhem district (35)tt --.- Sieys 9.8 69.5 20 0.7
East Finmarken (35) 1909.... a8 2.5 47 43-5 7 ibe ae
Reppertyordelem=sceteste rete ateiels ont a oa 18.2 45.4. 31.8 4.6
* Average 1923 and 1924 collections. T Average 1944-1947.
Tt Average 1908 and 1909. ** Skr. Norske Vidensk., 16, 1941.
have barely attained the required ro-cm size at the right time will, if slow growing,
make small smolts, perhaps only 12 cm long. But Elson has clearly pointed out that
this hypothesis “definitely does not specify that young Atlantic salmon become smolts
as soon as they reach a length of 10 cm, but rather that they transform after reaching
this length in the smolt-running season immediately following.”
Smolt lengths have been reported to be as follows, south to north: rivers tributary
to the Gulf of Maine, 5-6 inches (z8); average for the rivers of Canada, 5—7 inches;
the Little Codroy River, Newfoundland, 5—9 inches; George River, Ungava Bay,
8.5 inches (115).
Only a small percentage of the numerous alevins that are hatched survives
the freshwater stage as parr to become smolts and go to sea, nor is this astonishing
when all the dangers that threaten their continued existence are taken into account:
predators, competitors, disease, food shortage, abnormal water temperatures, and others.
For discussion of survival at different stages, see Length of Life (p. 479).
Downstream movements of smolts result from a change in their behavior asso-
ciated with the physiological changes involved in smoltification. According to Hunts-
man, when they assume the smolt condition they tend to leave the bottom, come nearer
470 Memoir Sears Foundation for Marine Research
the surface, and go or are carried downstream (64). While the resulting downstream
movement is facilitated by freshet conditions and although the peak of the run is usually
associated with a freshet, there is evidence to suggest that some other factor may be
responsible for the initiation of the migration. As long ago as 1926, Bull produced evi-
dence suggesting that floods did not provide the departure stimulus (25). However, his
suggestion that the initiation of the run was not associated with rainfall has not been
accepted. Swain has recently concluded that the immediate stimulus is either a rise in
temperature or some factor associated with a rise in temperature (125). Huntsman has
suggested that freshets appear to affect migration not only through carrying fish down-
stream but through breaking up their “homes” (67: 399; 70: 257).
Normally the main smolt migration occurs in the spring or early summer, chiefly
at night, with minor seaward movements at other times of the year. In the rivers of the
Maritime Provinces of Canada, the main migration takes place in May and June. In
the Little Codroy River, Newfoundland, the run during 1955 took place from May 11
to July 22, with the peak during June 1-15 (5: 63, 64). In the northerly part of their
range, in rivers flowing into Ungava Bay, most of them migrate in June and July,
though smolts may be taken there at all seasons (r15). It is evident from the foregoing
that all members of a year-class do not migrate in the same year, nor at the same time
in any year.
Life at Sea. After moving into salt water, the smolts apparently spend some time
in the estuaries and river mouths and descend to deeper water with the onset of winter
(18). The length of time they remain at sea before returning to fresh water on their first
spawning migration varies from one to four years, and in some areas up to five years.
As in river life, there is variation in growth and size at sea brought about by various
factors. Thus fish of the same age from different rivers often grow at different rates
while at sea, and members of one year-class may reach weights that are different from
those of another year-class from the same river with the same length of sea-life. Gen-
erally, the longer they continue to feed at sea, the larger they are when they return,
growing more rapidly in summer than in winter. Those that spend two or more con-
tinuous years at sea before spawning will in most instances be larger fish than those
of a similar age from the same river that have survived the ordeal of spawning, since
the growth of the latter has been interrupted by fasting for a longer or shorter time,
depending on how long they have been in fresh water before spawning.
Salmon develop rapidly during their sea life. Thus smolts may grow from only
one or two ounces to as much as six or seven pounds and a length of 16 inches or
more in their first year at sea.
There is comparatively little certain knowledge about the habits of Atlantic Salmon
when they are in salt water. Most of what we know about their movements in the
sea is derived from data on fish caught in nets set by commercial fishermen. Since such
nets are fished only in certain restricted areas, there is little knowledge of this species in
places where no fishing has been carried on. We have no knowledge, for instance, as
to whether or not they are to be found in midocean, far from any shore, because there
Fishes of the Western North Atlantic 471
has been little or no fishing there. In Canada, except for limited offshore fishing by
surface drift nets, commercial fishing is practically restricted to fixed traps and gill nets
operated near shore, nowadays only in tidal waters. Occasionally Salmon are taken in
herring weirs, in trawling for ground fish on offshore banks, and in line fishing, for
instance in the Gulf of Maine and the Bay of Fundy (78).
Even when they are taken in salt water there is no way of telling how far they
have traveled or from where they have come unless they bear a tag or mark and unless
the previous whereabouts of an individual fish so identified has been recorded. (Tagging
consists of attaching numbered tags to the fish; marking is the clipping of a fin or a
combination of fins.) In America, tagging and marking of specimens have been carried
out mainly in Canada, and to a lesser extent in Maine. Although river fish, both kelts
and smolts, have been used in most cases, some Atlantic Salmon caught at sea have
also been tagged.
Numerous specimens tagged in their river of origin have been recaptured there
later, and although it is usually assumed that they return to their natal river, it is
known that they sometimes enter and subsequently leave rivers other than their own.
If tagged specimens are recaptured in the sea at some distance from the place of release,
it is usually assumed that they are returning to their river of origin; this may be true
if they have the ability to direct their course; otherwise they may be merely wandering.
Studies indicate that some of them remain relatively close to their river of origin,
whereas others travel various distances to feeding grounds, often hundreds of miles
away. Such a variable pattern of dispersal has been shown by results from the tagging
and release of nearly 12,000 kelts by the Canada Department of Fisheries between
1913 and 1936 in 12 localities of the Maritime Provinces. Of those recaptured, some,
notably those of the St. John River, N.B., were recaptured within or close to the
estuary, and others were taken at various distances from the point of release, even as far
as Newfoundland and Labrador (62: 96; 67: 311; 58). The meaning of these results
is not clear. We can only guess how many of those found in or near the streams in
which they had been tagged may have been to some distant point (such as Newfound-
land) and returned. That not all had undertaken far journeys is suggested by the fact
that none of the St. John fish was recaptured more than 50 miles from the St. John
River mouth.
A wide distributional pattern has also been shown by the marking of smolts from
a number of rivers and by their subsequent recapture at sea as adults. Thus Atlantic
Salmon that developed from hatchery stock planted in the Pollett River (Petitcodiac
River system at the head of the Bay of Fundy) were taken in quantity by fishermen
over a wide area, including trap nets around Newfoundland, and drift nets off Mirami-
chi, northern New Brunswick.
Based on smolts marked in the Miramichi River, N.B., and recaptured as adults,
it has been estimated from one experiment that approximately one-third of the total
catch of Miramichi stock was taken off Newfoundland and Labrador in commercial
nets, about three-fifths in the Miramichi area in commercial nets and by freshwater
472 Memoir Sears Foundation for Marine Research
anglers, and the remainder elsewhere in the Maritime Provinces in commercial salt-
water nets. However, more were taken in the freshwater section of other rivers than
in that of the Miramichi (C. J. Kerswill, personal communication).
A considerable number of marked fish originating in the Port Daniel River,
Quebec, has also been taken around Newfoundland (4), as have specimens marked in
Maine rivers (45).
In contrast to the wide distribution of Atlantic Salmon from these rivers, only
one out of 31,359 Margaree River smolts was recaptured on the east coast of New-
foundland; they were taken mostly in the waters of western Cape Breton and the
neighboring mainland of Nova Scotia (66: 381).
Specific examples of results from taggings at sea are as follows:
Off Bonavista on the coast of eastern Newfoundland (27):
Of 386 grilse tagged, 140 were recovered—g3 °/, in Newfoundland waters, 7 °/,
in Canadian mainland waters.
Of 68 older specimens tagged, 28 were recovered—about 57°/, in Newfoundland
waters, 40°/, in Canadian mainland waters, and somewhat less than 4 °/, off
Labrador.
In Cabot Strait between Newfoundland and Nova Scotia (r4):
Recaptured along the eastern part of the north shore in the Gulf of St. Lawrence
and in waters off the northern part of western Newfoundland.
In the eastern end of Belle Isle Strait (14):
Recaptured in parts of the Gulf of St. Lawrence where fish tagged in Cabot Strait
did not go.
The results of the Cabot Strait and Belle Isle taggings were interpreted by Belding
(14) as indicating two distinct stocks.
Since Atlantic Salmon do not originate in Massachusetts rivers, specimens taken
from time to time off that coast probably come from the rivers of eastern Maine or
even from the St. John River. Other examples of rather extended movements are af-
forded by the capture of Salmon by trawlers 902-100 miles seaward from the outer coast
of Nova Scotia, and 100 miles from the nearest land, on Georges Bank (z8).
European findings also support the conclusion that some of them make extended
journeys (up to 1,600 miles), while others appear to move relatively short distances
(r02). Especially definite are Alm’s observations (2). Young fish from rivers tributary
to the Baltic and Gulf of Bothnia migrate to the southern part of the Baltic, where they
feed and then return to their rivers as grilse or older fish. Smolts from the rivers of
western Finland and from the northern and middle parts of Sweden move down the
Swedish coast to the southwestern corner of the Baltic or travel across the mouth of
the Gulf of Bothnia and then down to the Polish and Pomeranian coasts. Fish from
Estonia and Latvia find an adequate quantity of food comparatively close to their
native rivers.
Apart from the capture of one Scottish fish in Greenland waters, there is no
Fishes of the Western North Atlantic Ano
evidence that Atlantic Salmon travel from one side of the Atlantic to the other. This
fish, tagged at Loch na Croie, Blackwater River, Ross-shire, Scotland, on November
23, 1955, was recaptured in Eqalug Fjord, south of Sukkertoppen, Greenland (64°
56.5’N, 52°02’W) on October 15, 1956 (105). It is a matter of speculation whether
Atlantic Salmon from Europe and eastern North America intermingle in the mid-
Atlantic, as do salmon of Asian and North American origin over a broad area of the
Pacific high seas, at least between 160°W-170°E (6).
The farthest any American Atlantic Salmon is known to have traveled is from
the Annapolis River on the Bay coast of Nova Scotia to Ramah in northern Labrador,
a journey of about 1,900 miles accomplished in nearly two years (62: 105); there are
also many other records, though not as spectacular, of fish traveling hundreds of miles
both here and in Europe (z8). Blair has reported a distance of 792 miles for a grilse
and of 785 miles for an older Salmon tagged at sea off Bonavista, Newfoundland (27).
In Europe, two fish from Norway swam impressive distances of 1,600 and nearly 1,200
miles round the North Cape to the White Sea and to the Pechora River (102), which
is believed to be the eastern limit for the distribution of Sa/mo salar. In the Baltic,
Atlantic Salmon cover as much as 800 miles from their native rivers to their feeding
grounds (r02). The one tagged in Scotland and recovered in Greenland had traveled
1,730 miles in less than one year.
There appears to be general agreement that Atlantic Salmon feed in the mid-
depths (z8); that is, the bulk of them is neither on the bottom nor at the surface; how-
ever, some have been taken in both situations. In February and June 1948, 130 full-
grown Salmon averaging 11 pounds were netted by mackerel fishermen about 100
miles west of Land’s End, Cornwall, England, within 10 fms. of the surface (22).
Evidence of their occurrence on bottom is found in reports that trawlers have taken
strays on offshore fishing banks at depths of 50 fms. (r8) and 70 fms. (3). Dr. Hunts-
man was told by a fisherman of Sandy Cove, Digby Neck, Nova Scotia, that he once
caught an Atlantic Salmon at 35 fms., on a handline, when he expected to take a
pollock. Menzies has reported the capture of smolts in 65—70 fms. of water by trawlers
off Scotland (r00). There is no factual support, however, from fish actually taken,
for Roule’s belief, often quoted, that Salmon pass their growth period at sea, deep down
the continental slope.
Huntsman says that after Atlantic Salmon have become fat and when the water
is not too cold, they roam to and fro near the surface; it is during these movements
that they are taken in drift nets (69).
Return of Salmon to their Natal Rivers. The belief that Salmon return to their natal
river is very old. Isaac Walton reported that many in his day believed that ‘every
salmon usually returns to the same river in which it was bred.’ Scheer (rrg) has con-
cluded: ‘‘The hypothesis most compatible with these facts is that anadromous salmon
hatched and reared in a particular region will, when seeking freshwater, seek out and
return in the great majority of cases to the same region, even from considerable dis-
tances.”” From what distance, under what conditions, and to what extent Atlantic Salmon
474 Memoir Sears Foundation for Marine Research
find their way back to their home stream is an aspect of their sea-life about which a
great deal more information is needed before definite conclusions can be reached. That
most of those that actually spawn do so in the river where they have spent their
early life appears to be accepted by most observers who have investigated their life
history. Some investigators, however, believe that Atlantic Salmon that go to distant
places in the sea do not usually get back and thus may be regarded as lost (61: 18).
That Atlantic Salmon are able to return to their natal stream from a distant
place is proved by finding one in the Margaree River that had been marked there
two years before; in the interval, this fish had been caught, identified, and released at
Bonavista on the coast of Newfoundland; the return of this fish involved a round trip
of about 550 miles (66). Whether the performance of this one fish was exceptional
or whether it is usual for Salmon to undertake such migrations and even longer ones
and get back to their home stream must await the results of further research. In
studies carried out on the Miramichi River, N.B., over a five-year period, it was
found that smolts produced in two tributaries and marked at two-way counting traps
returned as adults to the trap of their natal stream with over 98 °/, accuracy (C. J. Kers-
will, personal communication). It is also known that Atlantic Salmon often wander in
and out of estuaries other than their own and that marked fish are occasionally caught
in strange rivers.
Views as to the means by which Salmon return to their river of origin tend in
two extreme and opposite directions. One extreme view pictures them as going uner-
ringly from their river to a predetermined feeding place from which, on the approach
of sexual maturity, they return unerringly to their river of origin, as if drawn by a magnet.
According to the other extreme view, they are carried to feeding grounds by ocean cur-
rents, and the only ones that find their natal rivers again are those whose wanderings,
with cessation of feeding on the approach of sexual maturity, perhaps aided by currents,
take them to the coasts, which they follow until they find themselves within the guiding
influence of their home stream.
Huntsman’s studies have led him to believe that Salmon, when outside of the
influence of their natal river, find their way back, if they do, through a process of
wandering until they come within the influence of the native river (73). Homing, he
says, is the end of wandering rather than a directive factor. He believes that the shore
or sea coast and transportation by currents are definitely directive factors for Salmon
movement (71).7 Toner, who has reached a similar conclusion, says that “salmon
approach the Irish coasts in a haphazard fashion and only when they are close inshore
do they make a definite search for the river of their origin” (428). Nearly a hundred
years ago, Austin’s observations led him to a somewhat similar view (8). He wrote:
A prominent feature in the migration of these fish to and from the sea is, that they always . . . hug the shore on
their way up and down. They do not, asserts Mr. Russell in his work on the salmon, lie off in mid-ocean, and then
as with a needle and compass, steer right into the river’s mouth; but they feel, or, as Sir Humphrey Davy ex-
pressed it, scent their way along the shore for many miles and follow closely the indents of the land.
7. For a discussion of the relation of currents to Salmon movements, see Bigelow and Schroeder (r7: 970).
Fishes of the Western North Atlantic ARS
There is a good deal of evidence that these fish in some way, probably through
their olfactory sense, are able to recognize the water of the river in which they spent
the early part of their life. Hasler has reviewed the results of observations and experi-
ments on the perception of pathways by fishes in migration (54). He concluded that
there is in river and creek water some characteristic odor to which the young become
conditioned while they are in the stream and which they recognize and to which they
orient upon coming in contact with it. Hasler, et a/. (55) have carried out experiments
that support the theory that fish, when out of touch with the shore or of the water
of their natal stream, have a sun-compass mechanism for direction-finding similar to
that found in birds (97) and in bees (49).
Coastal and River Spawning Runs. A spawning run usually occurs in two stages,
the first being the coastal run, the second the river run. During the coastal run, the fish
enter the river mouth or estuary and remain there for varying lengths of time before
they actually enter the fresh water on their river run. The river run may take place
shortly after the coastal run, or the two stages may be so widely separated that “‘early
running” and “late running” have little meaning as applied to a particular lot of fish.
If the fish appear early off the river mouth they may be referred to as “early running”’
by the commercial fishermen who catch them there, and if their actual entrance into
the river is long delayed, they may be “late running”’ to the anglers who take them in
the river.
With reference to the coastal run, no Atlantic Salmon move in from the sea off
Canadian rivers when the water is coldest, from January to April, with the probable
exception of the southern outer coast of Nova Scotia. Most of the fish that are
heading for the rivers of the Maritime Provinces (New Brunswick, Nova Scotia, and
Prince Edward Island) appear in coastal waters sometime between late May and July.
In Newfoundland, where there is a major and minor coastal run each year (95), the
fish of the major run appear in the latter half of May off St. John’s (in the southeast)
and progressively later northward, with the fish appearing at Cartwright (Labrador)
late in June; this coastal run in the spring is short, and profitable fishing can be pursued
for only about three weeks in any one locality; the minor run in the winter is much
more restricted, and evidence of it is confined almost entirely to the Avalon Peninsula
in the southeast, where some fish are taken in November and December. To the
south, in the St. John River, N.B., it has also been observed that some fish, mostly
large female grilse, enter the estuary in November and December, where they remain
until the following May before ascending the river; however, they do not spawn until
October (59).
Two of the conditions that determine how long Atlantic Salmon remain in or
near the river mouth before beginning their ascent of the river appear to be water
temperature and volume of river discharge, with high temperatures and the absence
of freshets being nonconducive to their movement (89). The Margaree River in
northern Nova Scotia presents a striking case of the action of these two factors (63).
The Northeast Margaree, which has cool spring water that travels from high land
476 Memoir Sears Foundation for Marine Research
through a deep gorge, provides angling from the time the fish start to wander in the
adjacent sea, from June onwards through the summer and fall. But the Southwest
Margaree, which arises from the large shallow Lake Ainslie in relatively low country,
has warm water and, in contrast to the Northeast Margaree, is populated by alewives,
eels, and white perch; here Atlantic Salmon sometimes provide a little angling at the
beginning of the season; otherwise they are not seen ascending the river until the water
becomes cool in the fall, toward spawning time.
It has long been known that runs of certain stream fishes are more or less definitely
associated with floods, freshets, or spates; sometimes artificial freshets have been used
to bring Atlantic Salmon up from the sea. For example, in 1888, in the Grimersta
River, Scotland, water from Loch Langabaht was held by a six-foot dam at the mouth
of the lowest of a series of four lochs. For a long time the fish had gathered at the head
of tide (in a very dry season), and when the dam was broken they swarmed up the
river (27). In the Moser River (Nova Scotia), during the dry summer of 1942, sharp
but not large artificial freshets brought double the expected number of Atlantic Salmon
and brook trout from the sea (70: 257). Artificial freshets,* however, have not been
used regularly, for this procedure is rarely feasible.
The spring runs in the Maritime Provinces of Canada are usually at their height
in June.® Most of the rivers tributary to the Bay of Chaleur and around the Gaspé
Peninsula have early runs, as do those on the northern shore of the St. Lawrence, al-
though the runs there are slightly later.1° The Miramichi in central New Brunswick
“thas both early and late runs. The small rivers of the coast of eastern New Brunswick,
the Nova Scotia rivers of the Northumberland Straits and the rivers of Prince Edward
Island have fall runs only” (15: 225-230). In the Koksoak River, tributary to Ungava
Bay, northern Quebec, a summer river run has been reported as occurring sometime
between July 25 and the end of August, and sometimes there is a late run in September
(39). At or near their northern limit in the western Atlantic, they enter a river near
Kapisigdlit (¢nner Godthaab Fjord) in southwestern Greenland in both July and Oc-
tober (53). In England, gravid females may ascend rivers from late autumn until
after Christmas, and some of them may remain in ice-free rivers for 12 months or
more prior to spawning (121; 122).
The distance they run upstream to spawn depends on a number of factors such
as the length of the river, presence of insurmountable falls and man-made obstructions
(p. 485), available spawning sites, etc. The fish that enter the river the earliest usually
go the farthest upstream before spawning. In large rivers, as in the St. John system,
this may mean an upstream run of more than 200 miles and of many months spent in
fresh water before spawning. Fish entering later travel lesser distances, and very late
8. Hayes (56) has discussed the results of observations and experiments on artificial freshets and other factors as con-
trolling the ascent and populations of Atlantic Salmon in the La Have R., Nova Scotia.
g. It has been reported that, in the latter part of the eighteenth century, when Atlantic Salmon were still running
in the Connecticut River, the river migration began about April 1 and was at its height in May, though occa-
sional fish were taken as early as mid-January (23).
to. The early-run fish are much more desirable for the angler than the late-run fish, both because they rise more
freely to the fly and because they are much more desirable on the table.
Fishes of the Western North Atlantic 477
fish often spawn only a short distance above the head of tide. It has also been observed
that most of the rivers having only fall runs are small, with their spawning grounds
relatively short distances from the river mouth. The Miramichi is the only large
Canadian river that has a fall run.
As they gradually work their way upstream, from one resting place to another,
they assemble in the deeper and stiller parts; these are known as salmon pools to
anglers. In some rivers they must ascend through long stretches of turbulent rapids,
some of which culminate in falls. Whether or not the fish can surmount falls depends
on the configuration and height of the falls and on the amount of water at the base;
the deeper the water the higher they can leap, but to—12 feet seems to be the maximum.
Salmon leap" not only in attempting to surmount falls but in the quieter stretches of
the river, and at sea.
Differences and Changes in Composition of River Runs. Salmon entering each river
system have certain characteristics, such as smolt age, size, length of sea-life, and time
of running (early or late), that distinguish them from those of other rivers; and
these characteristics, with minor variations from year to year (g: 304), tend to persist
over long periods of time. For Newfoundland waters it has been demonstrated (95)
that there are no fewer than 25 classes of fish with different life histories, and 33
types are listed as likely to be found in these waters. According to the laws of permu-
tations and combinations, a very large number of types could and possibly do occur,
some of them rarely.
Spawning runs consist of both grilse and older Atlantic Salmon of all sizes and
ages with two to four or more years of sea-life behind them; and, whether grilse or older
fish, those returning early in the season are generally smaller than those returning
later because their growth period at sea has been shorter. Grilse, ready to spawn, return
to fresh water during their second summer at sea, and these are mostly males; for
example, in the Little Codroy River, Newfoundland, only 15 °/, of the spawning grilse
taken were females, and in Norway only 19 °/. But in certain localities of Scotland,
females may constitute more than 50 °/, of the grilse (100: 79).
In some rivers grilse are the more numerous in a run and in other rivers the
older fish predominate.!? Rivers reported as having a high proportion of grilse in-
clude: The Minas system, northern Bay of Fundy, 89 °/, (59); the Little Codroy,
southeastern Newfoundland, 64 °/, in 1954, 74 °/) in 1955 (106). For grilse in other
rivers, the following percentages have been reported: six western Newfoundland
rivers, 34.3 °/, (ZZ); an unidentified Greenland river, 33.4°/, (53). In the Moisie and
Grand Cascapedia (north shore of the Gulf of St. Lawrence and Chaleur Bay, respec-
tr. It is from its habit of leaping that the Salmon gets its Latin name. Both Salmo and salar are apparently from the
same root, salire, to leap.
12. Estimates of the composition of a catch or population of fish may be subject to large error, as in the case of esti-
mating the proportions of grilse and older Salmon. In commercial fishing, the gear used may take a selective rather
than a representative sample of the run, being designed to take only the larger individuals. Often the fishing effort
is concentrated on the main run and is not carried on throughout the year. Even in experimental fishing, a good
representative sample can be obtained only if the fishing is continuous and is done with gear designed to sample
accurately the various sizes.
478 Memoir Sears Foundation for Marine Research
tively) and in New England rivers, grilse have been reported as being seldom seen
(10I; 104; 29) or rare (7).
Rivers having a high proportion of fish that have been at sea for three years are
known as “large salmon”’ rivers, two of the most notable being the Moisie and Grand
Cascapedia rivers, for which the following percentages have been given for two- and
three-year sea-life fish, respectively: 18.3 and 58.8 for the Moisie(z2); 6 and 58.8 for the
Grand Cascapedia (29). The tabulated data to the left, giving the percentage of Atlantic
Salmon of 30 pounds and over in the Moisie run, suggest that
1900-04.... 11.78 there has been a decline in the proportion of heavier fish in
190$—09.... 5.86 that river (12).
IQIO—-14.... 3.96 The average weights (in pounds) of Atlantic Salmon re-
I915-19.... 5.69 ported for various rivers are: for those two years at sea—10.8
1920-24.... 2.44 for the Restigouche in New Brunswick, 8.9 for the Godbout
1925-29.... 2.33 in Quebec, and 11.2 for the Wye in England, averaged from
1930-34.... 5.68 records of more than 20 years (9); 11.7 for the Grand Casca-
pedia, 10 for the Miramichi, and 11.4 for the east coast
of Newfoundland, including Labrador (20); for those three or more years at sea—
Grand Cascapedia, 23.6 and 37 for 3 and 3+ fish, respectively (29); Miramichi, 19.5
for 3 and 3+ fish combined;?* and the coast of eastern Newfoundland, including Lab-
rador, 19.5 for 3 and 3+ combined (20).
It has been suggested for Newfoundland Atlantic Salmon that those having a short
river life tend to stay longer at sea before they return to spawn (95). On the other
hand, the opposite suggestion is implied in Dahl’s statement that findings in Norway
do not appear to support the idea that the longer parr remain in the river the less
time they will spend in the sea (37). For Scotland, Calderwood found that smolts
that matured rapidly and well in the river were likely to stay away from fresh water
for some years after leaving it, while smolts that remained in the river for more than
two years and developed slowly were likely to return to it soon. However, he found
that this relationship did not apply to the Grand Cascapedia, Quebec. A tabulation
of the characteristics of Atlantic Salmon from various sections of the Gulf of St.
Lawrence does not show a consistent relation between those factors (12).
There is no agreement from either observations or experiments concerning the
relative extent to which heredity or environment affect the early or late running char-
acteristics of fish of the different rivers (144; 26: 22), or the marked changes that
have sometimes occurred in the proportion of grilse running to a number of streams,
or other differences. The nature and extent of these changes are indicated in the fol-
lowing examples.
A serious and almost catastrophic decline in the stock of grilse along the coast
and in the rivers of Scotland during the present century (100: 79), more especially after
about 1908, has prompted the question as to whether selective fishing or some other
factors may be responsible for these changes.
13. Blair, M. A. thesis, University of Toronto (1932).
Fishes of the Western North Atlantic 4.79
Following construction of a hydroelectric plant on the Shannon River, Ireland,
there was a marked change in the composition of the incoming runs (138: 789). Taken
as representative of the pre- and post-period of hydroelectric operation, about 27.5°/,
of the total run occurred in June of 1928 compared with 70 °/, that month in 1941;
in 1928, 24.7°/, of the run consisted of grilse, and in 1941, 75°/). The age of the fish
in 1941 averaged approximately one year less than in 1928, and of course the average
size was smaller. This change, due to a proportionate rather than an actual increase in
the number of grilse, arose from a reduction in the number of age groups spending
two or more years feeding at sea. Hydroelectric operations resulted in a reduction of
the Atlantic Salmon stocks in the river to about 30-40 °/, of those of prehydroelectric
days. The Mulcair River, which was the major spawning ground for the smaller fish,
became the major spawning tributary for both large and small fish.
In the Miramichi, regulations to restrict fishing to the early run and encourage
reproduction of the late run (natural as well as artificial) have not succeeded materially
in either reducing the early run or increasing the late run, although this unconscious
experiment had been carried on for more than 80 years (58).
Length of Life and Spawning Survival. During all stages, from egg to fry, fry to
smolt, smolt to adult, and after spawning, there are wide differences in the number of
Atlantic Salmon that survive, not only from river to river but in the same river from
year to year, depending on meteorological conditions, population density of Atlantic
Salmon as well as other species, abundance of predators, disease, and man’s activities.
In considering these differences, factors responsible for discrepancies in estimates of
the composition of the runs should be kept in mind (ftn. 12).
Kerswill has found in experiments in progress on the Pollett River, N. B., that
survival of fry from egg deposition (three light seedings) had averaged 6°/, in July
(1957 Mss.). “Under favourable conditions, each 100 square yards of stream is cap-
able of producing 5 or 6 smolts.... These will arise from 10 or 12 large parr, and
they in turn from 30 to 40 hatchery fingerlings planted in late summer.’’ About 200
eggs, it was estimated, should produce this number of fish (5: 49). Another estimate of
survival from fingerling to smolt stage, based on observations for five successive years,
is 8 °/, (go).
Percentage survival from smolts to returning adults is difficult to determine; the
following figures are not all directly comparable. For example, it has been estimated
from the counting and marking of descending smolts and from the trapping and
counting of returning adults in several rivers of the Maritime Provinces that ‘about
8°/, of the smolts survive the rigors of life in the sea and return to fisheries and
rivers as mature salmon” (43: 25; 91: 825). On the other hand, the contribution to
various fisheries by the rivers was found to be about 2°/, of the marked descending
smolts (5).
The following figures on smolt survival have been published: Little Codroy
River, Newfoundland, 3.3 °/, (107); Pollett River, 0.5 °/, (88); Great Britain, 1 °/,
(116); Tay River, Scotland, nearly 2 °/, (100); Sweden, 5-10 °/, (32).
480 Memoir Sears Foundation for Marine Research
Data on survival after spawning—more adequate than data on survival during earlier
stages of life history—show that the percentage of Atlantic Salmon that survive spawn-
ing is usually rather small but is higher in North America than in Europe. The strain
of spawning leaves the fish thin and exhausted. Although some of the spawned-out
kelts descend to the sea in the autumn after spawning, others winter in the river and
migrate downstream the following spring, sometimes along with the migrating smolts.
That comparatively few are seen dead in the rivers suggests that most of those that
die regain salt water first. The high death rate probably results chiefly from a combi-
nation of exhausting travel up the river, prolonged fasting en route to the spawning
site, and production of the spawning products.
The percentage of fish that survive after spawning to the stage when they are
ready to spawn again is reflected in the percentage of such fish in the catch of adults
participating in the run to a given area or river. The following are examples of such
percentages: for all the Canadian rivers investigated to 1934, 3-26 °/, of the total
spawning populations of the rivers concerned (10); Grand Cascapedia, Quebec, 34 °/,
in 1926, and 22 °/, (29; 23); Miramichi, N. B., 12.8 °/, (79); Upsalquitch, N. B.,
5 lo (13); Moisie, Quebec, 16.7 °/) for 1922 and 1923 combined (zor), and 22.3 °/,
(12); Little Codroy, Newfoundland, 6°/, for 1954, 5°/) for 1955 (106); eastern New-
foundland, including Labrador, 16.5 °/, (20); Greenland, 14 °/, (53); rivers on the
east coast of Scotland, about 4°/, (100); two rivers on the west coast of Scotland,
II % and 34 °/) (zz6); all rivers of Ireland, about 5°/,, or a range of 1-15°/, (129);
Adour River, France, 3.8 °/, (130).
Of 6,475 kelts tagged and released, only 2.12 °/, were subsequently recaptured
(ror 2en):
Of those that spawn a second, third, or more times, Menzies observed that in
one year, of 63 previous spawners in the Moisie River, 35 had spawned once before,
I1 twice, 16 three times, and one four times (ror). On the coast of eastern Newfoundland
(including Labrador), in 1939, 15.9 °/, had spawned once, 0.5 °/) twice, and 0.08 °/,
three times (20). In six small rivers on the coast of western Newfoundland, of 70 previous
spawners, 15 had spawned twice or more, two had spawned three times, and one had
spawned four times (ZZ).
In Ireland, among nearly 50,000 fish, the scales of which were examined, 3 °/,
had two spawning marks, but only six fish had three marks (129). In Scotland only
0.2 %/, and o.1°/, of fish from Spey and Aberdeenshire Dee, respectively, on the east
coast, had spawned twice before capture. In the Add River, 6.5 °/, had two spawning
marks, 2.9 °/), three. A fish with four spawning marks was taken in the Add in r1gr4,
and another was taken in the Conon in 1920 (Z00).
The percentage of Atlantic Salmon that survive spawning is [higher among
females than among males. A male that returns to spawn a second time is a rarity.
According to Menzies, there is no record of a male spawning three times (oT: 338),
in spite of a report that six of the 16 Moisie fish that had spawned three times were
males; Menzies was inclined to question the identification. However, if they were
Fishes of the Western North Atlantic 481
males it suggests the possibility that males may live longer in the western Atlantic than
in the eastern Atlantic.
Fourteen years is the greatest age reported for an Atlantic Salmon. A fish from
the Moisie River, Quebec, that had spawned in four consecutive seasons is reported
to have attained that age (28). Another Moisie River Salmon weighing 38.5 pounds,
which had also spawned four times, is reported to have been 13 years of age (ZOZ).
Menzies has also referred to a 13-year-old Scottish Atlantic Salmon (z00). All of these
very old fish were females.
Tolerance to Temperature. It is evident, from their northerly range, that Atlantic
Salmon prefer cool waters, and from their observed behavior in streams, that water
temperature is certainly one of the most influential of environmental factors, as is illus-
trated in the two branches of the Margaree River (p. 475). Egg development, incu-
bation time, and the rate of growth of fry are definitely linked to temperature condi-
tions (p. 466).
The lethal temperature for any given individual varies with its thermal history.
Experimental fingerlings that had been living at 25°C began to die when exposed
for any length of time to a temperature of 28.5°C; and planted fingerlings died when
the stream temperature reached 30.5°C (98). Parr tolerate temperatures as high as
28-29.5°C (82-85°F), at least temporarily, but they die under experimental condi-
tions at temperatures of 32.9 and 33.8°C (90.2 and 91.8°F). The older fish die when
the water temperature rises as high as 28-29.5°C, which happens in some of the
smaller streams of New Brunswick and in some of the rivers of outer Nova Scotia during
periods of very low water resulting from deficient rainfall and protracted hot weather
(65: 485; 67: 476).
The success or failure of commercial fishing and of angling is closely associated
with the temperature of the water, and it is common knowledge among anglers that
Atlantic Salmon do not rise freely to the fly if the water is warmer than about 70°F.
Food. Studies of specimens from many parts of their range indicate that the same
items—May fly nymphs, caddis larvae, and particularly chironomid larvae—usually pre-
dominate in the food of parr; all live in the bottom. However, as the fish grow, surface
food assumes greater importance. Allen has given a comprehensive review of the feeding
habits of young Atlantic Salmon (1: 47-76).
Studies in eastern Canada have shown that chironomid larvae consitute the chief
food of parr that are beginning to feed (139: 499-506); later, especially during mid-
summer, May fly nymphs become important. Toward the end of the summer growing
season, caddis larvae are taken in increasing numbers. Some Plecoptera (stone flies)
as well as simulids and tipulids have been found in small or limited quantities in the
stomachs of parr examined in early summer and in the fall.
The feeding by parr in the River Dee was found to be indiscriminate. A single
meal might include representatives of a dozen or more species taken at random from
under stones (insect larvae), in open water (Cladocera and small beetles), or from sur-
face driftage (aerial and terrestrial arthropods); the large number of dipteran flies
31
482 Memoir Sears Foundation for Marine Research
(uniform in species) sometimes found in a single fish suggests that it might have
snapped at swarms in the air (33).
Because adults do not feed while they are in fresh water and have usually ceased
feeding by the time they reach inshore waters, where most of them are caught com-
mercially, comparatively little is known about what they eat when they actively feed at
sea. Among the food items most commonly mentioned are herring, capelin, and sand
launce; small mackerel are also frequently mentioned.
Many of those taken in traps inshore are empty, probably because they share
with many other fishes the habit of disgorging their food out of fright, or digesting
it before they are removed from the traps (84: 165-166). In spite of this, there are
numerous records of the Atlantic Salmon’s diet inshore, where they have been re-
ported as preying on small fishes of various kinds as well as crustaceans. For example,
many of those from the north shore of the Gulf of St. Lawrence, opened by Comeau,
contained herring, small mackerel, and sculpins (cottids), and some had gorged on
capelin (34: 184-188). Kendall found alewives (Pomo/obus) in specimens from the St.
John River, New Brunswick, and smelts in Penobscot River fish (87: 32, 34). Ungava
Bay specimens have contained Arctic cod (Boreogadus saida) and sculpins (39).
Stomachs of this species from off Norway are sometimes packed full of herring;
these as well as launce are common food in the Baltic; Day saw 22 entire sprats (C/upea
Sprattus) up to seven inches in length taken from the stomach of a 12-pound fish caught
in the tidal portion of the Severn River, England (38). Haddock, eels, small trout
(Salmo trutta), flatfishes, and other species have also been found in Atlantic Salmon.
A hook-and-line fishery using herring as bait is (or was) carried on in the Baltic, and
hooks baited with launce (4mmodytes) or with pieces of mackerel sometimes catch them
in British waters. Trout, charr, and salmon parr have been reported as being taken
from stomachs of Salmon caught by anglers in Scotland (30: 73).
Atlantic Salmon from the Penobscot, Maine, have been full of “shrimp” (probably
euphausiids),'* and grilse have been described as feeding on euphausiid shrimps and
pelagic amphipods (36). Sand fleas (gammarid crustaceans) rank with launce and herring
as food in the North and Baltic seas; crabs have also been found in Salmon.
In Iceland, and in Great Britain and elsewhere in Europe, they are caught by
anglers using natural bait (prawns, angleworms) and artificial minnows, not only in the
estuaries and lower reaches of the rivers, but also in lakes near the sea; no doubt this
would be the case in Canada also if bait-fishing were allowed there.
Adults normally eat very little, if at all, after they have run up well beyond the
head of tide. Accordingly, their stomachs seldom contain anything except a little yel-
lowish fluid. Of the numerous fish from fresh water on the north shore of the Gulf
of St. Lawrence that were opened by Comeau (34), only four contained recognizable
items: two taken by anglers had eaten insects; one speared in November had parts of
a mouse in its stomach; and a kelt contained organic fragments, apparently of some
14. For a survey, with references, of the diet of Salmon in general and in the Gulf of Maine in particular, see Kendall
(87: 33-34); for diet lists for the Baltic and North Sea Salmon, see Eichelbaum (41).
Fishes of the Western North Atlantic 483
bird. The energy to maintain them during their stream life when they return to spawn
and the resources for the development of either eggs or milt have been built up and
stored in their bodies during the feeding stage in salt water.
Why Atlantic Salmon in fresh water so often rise freely to artificial flies has
been widely argued and still is. An experienced angler has suggested verbally that on
sighting the fly an occasional fish suffers an attack of temporary insanity; this ex-
planation is perhaps as satisfying as any other that has been proposed.
Predators. Atlantic Salmon are preyed on throughout their life, in both stream
and sea, and this factor, probably as much as any other, is generally responsible for
the small percentage that survives to spawn (pp. 479, 480). Most of what we know
about this subject is confined to reports for fresh water; as Pyefinch has aptly observed
(116): “The depredations of birds and other fish in fresh or estuarine waters may be
impressive only because they are visible and are noticed, they may actually be much
less significant than the effects of other predators present, for example, in the sea.”
The extent of predation today relative to that under primitive conditions is not known
(142; 143).
Atlantic Salmon are probably preyed on throughout their range by one or another
of the predatory fish that cohabit the same waters. In fresh water, birds (mergansers,
cormorants, gulls, and herons) destroy significant numbers of young. In the Maritime
Provinces of Canada, mergansers (especially Mergus merganser) and the belted king-
fisher (Megaceryle alcyon) are often destructive; control of these in the Pollett River
(Petitcodiac R. system) is believed to have increased smolt production eightfold. Follow-
ing control of mergansers!® on the Northwest Miramichi (but not on the Dungarvon
which is also tributary to the Miramichi system and has a similar run of adults),
production of large parr increased more than fourfold (44). Mergansers are also serious
predators on young Atlantic Salmon in Sweden (93). Sa/velinus fontinalis (brook trout)
in America and Sa/mo trutta in Europe eat young Atlantic Salmon in the streams;
and in some areas eels reduce their number (43: 15).
In marine situations, both harbor seals and gray seals are reported as frequently
preying on Atlantic Salmon when these gather near river mouths. Indeed, depredations
by seals, chiefly gray seals, cause substantial losses to the fishermen who operate traps
and drift nets in the estuary of the Miramichi and within the mouth of the river (47:
28). Very little information is available about the predation that obviously must occur at
sea. Ten Salmon, 10-12 pounds each, are reported to have been found in the stomach of
a shark about 25 feet long, caught eight miles east by south of Peterhead, Scotland.
One has also been reported from the stomach of a shark taken in Greenland (p. 490).
An inspection in both 1956 and 1957 of adults caught in the estuary of the Mira-
michi River, N. B., showed that 0.46 and 0.66 °/,, respectively, either had small living
lampreys attached to them or bore lamprey scars; the corresponding figures for those
in fresh water in the Northwest Miramichi were 0.15 and 0.25°/,; no fish were
found with large adult lampreys attached. These records indicate that lampreys are not
15. White has also studied the feeding by mergansers on Salmon fry (139, 140, 142, 143):
484. Memoir Sears Foundation for Marine Research
a serious menace to adult Atlantic Salmon in the Miramichi (M.H.A. Keenleyside,
personal communication).
Parasites and Diseases. Numerous parasites and a few diseases have been reported
for Atlantic Salmon, but it is not known to what extent they affect the population. For
the most part, ways and means of controlling them have not been found. Among the
external parasites, sea lice (Lepeophtheirus salmonis) and gill maggots (Sal/mincola salmonea)
are common and widespread, and five others have been reported as sometimes occurring.
Of the diseases, ““Salmon disease’’ and furunculosis are the more common.
Sea lice, which occur on Atlantic Salmon throughout their range in both North
America and Europe,” are parasitic copepods frequently found attached to fish that
have recently entered fresh water from the sea. Sea lice usually die within a week of
being brought into fresh water, but sometimes they live longer. Most of those attached
to Atlantic Salmon are females about three-quarters of an inch long. White observed
that some of the fish that entered the Moser River in 1939 were so heavily infested
with them and had so much of the skin in the occipital region eroded away that the
fish died after entering fresh water (141). The large fish were not so heavily infested
as grilse, some of which had an almost complete layer of lice extending over the dorsal
part of the body from the posterior edge of the eyes to the caudal peduncle, and a
few lice around the anal and ventral fins. Before the skin sloughs away there is a
distinct white area over the regions affected; White believes this is the condition de-
scribed as “white spot.’’?”
Gill maggots (Sa/mincola salmonea) are smaller than sea lice (0.2 5—0.33 in.), and
most of those on Atlantic Salmon are females. Although they are able to breed only in
the river, numbers survive in salt water. Maggots attach themselves to the gills of
the fish in fresh water, only after the latter have been in salt water and have returned.
They are not found on young that have remained in fresh water (31: 16). Maggots
have been found on fish that have been in fresh water for as much as a year and in
salt water for two additional years.
Internal parasites of Atlantic Salmon are reported under more than a hundred
names, but until careful taxonomic studies have been made it is impossible to say how
many of these represent valid species. Twenty-five names, for example, have been re-
ported for trematodes or flukes, 38 for cestodes or tapeworms, 25 for nematodes or
round worms, 22 for acanthocephalous or thorn-headed worms, and 8 for Protozoa.
Furunculosis, the ulcer or boil disease, caused by the bacterium Bacillus salmonicida,
has occasionally assumed epizootic proportions in hatcheries, whence it has spread to
natural waters; it has also reached similar proportions at times in southern England.
A second bacterial disease (Salmon disease), caused by Bacillus salmonis pestis, has
occasionally assumed serious proportions in southern England.
16. Lists of parasites reported for the Atlantic Salmon were provided by the Parasitology Division, Ontario Research
Foundation; Animal Disease and Parasite Research Division of the U.S. Department of Agriculture; and Institute
of Parasitology, McGill University.
17. Sea lice also affect brown trout (Salmo trutta), brook trout (Salvelinus fontinalis), Dolly Varden (Salvelinus alpinus
malma), and two Pacific Salmon (Oncorhynchus tshawytscha and O. gorbuscha) (120; 51; 147).
Fishes of the Western North Atlantic 485
Decline in Abundance. Atlantic Salmon were extremely abundant in early colo-
nial days from New England northward to the northern shores of the Gulf of St.
Lawrence and Labrador in every river that was not barred to them by impassible falls
and where spawning conditions were suitable. The few taken by the natives merely
reduced overcrowding at the spawning sites; and the waters ran clear and clean, deeply
shaded by virgin forest growth down to the stream and river banks. The advances in
colonization and expansion, however, with concomitant increases in man’s exploitation
of both the land’s resources and the fishery, soon began to deplete the Salmon pop-
ulation.
In 1849, Perley (122) wrote:
The quantities of salmon in the River Restigouche and Miramichi, at the first settlement of the country
were perfectly prodigious; although many are yet taken annually, the supply diminishes from year to year. And
this is not surprising when it is considered that many of the streams formerly frequented by salmon are now
completely shut against them by mill dams without “‘fishways”—that in the branches of the large rivers, as also
in the smaller rivers, nets are too often placed completely across the stream, from bank to bank, which take every
fish that attempts to pass—that “‘close time” in many of the rivers is scarcely, if at all, regarded—and that, besides
the improper use of nets at all seasons, fish of all sizes, are destroyed by hundreds in the very act of spawning,
by torchlight and spears at a time when they are quite unfit for human food.18
Perhaps the most destructive practice in the early days, especially in New England,
was the construction of dams in connection with the production of power through
water wheels, as typified in the Connecticut and Merrimack rivers. The Connecticut
River, extending 407 miles northward into northern New England, supported a large
run until the end of the eighteenth century. In four years, however, following the con-
struction of a dam 16 feet high across the river at Miller’s Run in 1798, there were
few fish left, and in 12 years there was none (23). The fate of this species in the Mer-
rimack River in northeastern Massachusetts (110 miles long and extending into New
Hampshire) also typifies the history of Atlantic Salmon in New England rivers from
which they have been barred (z8). They spawned plentifully in this river’s upper
reaches as late as 1793; in 1790, for example, 60-100 fish a day was the usual catch
with a go-yard seine, near the river’s mouth; but the completion in 1847 of a dam at
Lawrence, Massachusetts, completely barred the spawning areas of the river to them.
For some years thereafter those that had been produced in the upper reaches gathered
below the dam in spring and summer, “‘vainly endeavoring to ascend.”’ There has
been no run in the Merrimack since 1859 or 1860, when the last fish hatched above
the dam had lived their span of life, and there has been no spawning there except by
a few that may have been lifted over the dam.
The spawning areas in only a few of the numerous rivers in New England that
supported Salmon runs in the early days were still open to them by the end of the past
century, and by 1925 only two of the Maine rivers, the Dennys and the Penobscot,
saw regular runs, aided in part by the artificial propagation carried on by the U.S.
Bureau of Fisheries. Since then, however, the situation has improved slightly in Maine.
18. See also 108; 92; 7.
486 Memoir Sears Foundation for Marine Research
Canada has always had more Atlantic Salmon than New England, and the area
occupied by the Canadian fishery has seen less industrial development than the area
originally frequented by Atlantic Salmon in New England. It was stated in 1931 that de-
pletion in the Maritime Provinces of Canada was particularly evident in rivers emptying
into the Bay of Fundy, and in some cases it had progressed to total extinction (59).
The Petitcodiac River system in New Brunswick afforded an example of a stop just
short of extinction. However, in 1958 it was reported that Atlantic Salmon in fluctu-
ating numbers occur in 300 rivers of Canada’s Atlantic provinces (45: 19), with 75 °/,
of the population being restricted to six principal river systems of New Brunswick and
Newfoundland.
The largest recorded commercial catch in Canadian waters, exclusive of New-
foundland, was taken in 1873 and amounted to 6,698,200 pounds. The largest for
United States waters was 205,679 pounds in 1888. At present the total Canadian catch,
including Newfoundland, totals up to 3,000,000 pounds a year, one-half to two-thirds
of which comes from Newfoundland, including Labrador. U.S. commercial production
is now less than 1,000 pounds. From the 1888 high in New England, the catch de-
clined so that about f0 years later (1932-1938) it varied between 16,000 and 40,000
pounds; and during the next ten years it averaged only 3,600 pounds, less than 2 °/,
of what it had been about 60 years before.
The total Canadian angling catch in recent years has been 75,000 Salmon per
year. Of this, the Miramichi River, N.B., reputed to be the greatest salmon river in
the world, contributed 30,000, followed by the Restigouche with 3,000; all the New-
foundland rivers combined yielded 20,000 fish, and the Quebec rivers 12,000 (gZ).
In Maine, the combined angler’s catch for 1956 was only 278 fish (148: 212).
Lumbering and agriculture, by removing natural soil cover, have exposed large
areas to erosion, thus creating several conditions detrimental to this species, including
silting of stream beds, floods, and summer low water. Floods wash out nests and eggs
and destroy the homes of the fry and fingerlings. Low water not only enables predators
to more readily see and capture the young, but it results in higher summer tempera-
tures that sometimes reach the lethal point for Salmon. Spraying forests with ppT
and other chemicals to control insects (such as the spruce bud-worm and others) has
recently created another danger. Not only is ppr harmful to many of the insects on
which the young feed, but it is directly destructive to the fish themselves (4; 86; 75).
Pollution of stream waters by poisonous chemicals and other wastes has also been in-
jurious, sometimes directly, and sometimes indirectly through the removal of oxygen
from the water.
The more recent use of rivers for the development of hydroelectric power has
added other destructive conditions; either the spawning grounds are eliminated through
the creation of large lakes, or the young are destroyed in going over the spillway or
through the turbines. Irrigation projects involving dam construction and creation of
storage basins also obstruct their ascent and eliminate spawning grounds.
Whether Salmon predators, for example mergansers and kingfishers, are more or
Fishes of the Western North Atlantic 487
less destructive today than under primitive conditions has not been determined. Their
relative effect on Atlantic Salmon populations now as compared with those in earlier
times may be due to either or both of the changes that affect the ability of the predators
to reproduce or to prey on the fish (142; 143).
Overfishing has often been blamed for the decline noted. Anadromous fish are
more vulnerable to overfishing than species that remain at sea throughout their entire
life because they concentrate within a small area when they run up rivers to spawn;
thus it is much easier for fishermen to take a high percentage of the stock. The high
price that Atlantic Salmon command encourages intense exploitation of the fishery.
However, present evidence indicates that the fishery as now permitted by law does
ATLANTIC SALMON
2)
os
=)
D4 LANDINGS
6 3
BY i2
°
|
a
=
1870 1880 1890 1900 1910 1920 1930 1940 1950
Figure 123. Commercial landings of Atlantic Salmon from the Gulf of St. Lawrence area (Cape Gaspé to
Cape Breton) for the period 1870-1954 (Elson and Kerswill, 44).
not reduce the spawning population below what is necessary to provide all the young
that the streams, in their present condition, can support (42). In other words, the
decline of the fishery has been due to the creation of unfavorable conditions in
streams rather than to overfishing (see also Maintenance and Restoration of the Fishery,
p- 488-490).
Fluctuations in Abundance. It is reasonable to assume that the Atlantic Salmon
population has always fluctuated more or less in abundance, but it is only for recent
years that it has been possible to estimate with any degree of certainty from the catch
statistics the extent and nature of the fluctuations. Although catch cannot be regarded
as an accurate measure of actual population, it is the only available measure we have.
On the basis of such statistics it must be concluded that populations not only have
fluctuated widely from time to time but have declined seriously since 1930. However,
since 1955 there has been evidence of some improvement.
The extent and nature of these fluctuations, illustrated in Fig. 123, show three
important features: (1) the great difference between neighboring highs and lows, for
instance between 1873 and 1881 and between 1924, 1928, and 1930; (2) the over-all
steady decline since 1930, with recent low catches persisting much longer than in earlier
years; and (3) the tendency of the peaks of production to occur about every 10 years.
This periodicity was noted almost simultaneously around 1930 by several investigators.
Griswold’s discovery was based on a study of the records of the Cascapedia and Resti-
488 Memotr Sears Foundation for Marine Research
gouche Salmon clubs and of New Brunswick and Quebec fishery statistics (50). Phelps
and Belding corroborated Griswold’s findings from the records of the Restigouche
Club, noting a periodicity of nine or ten years extending back to 1879 (zzr4). Hunts-
man used for his study the fishery statistics of the Maritime Provinces of Canada, which
extended back to 1869 (58: 9-14). Huntsman attributed periodicity in Atlantic Salmon
to periodicity in rainfall (60: 17-28; 72: 43-53). Griswold related it to a nine-year
cycle of tides, believing that coastal currents move offshore and onshore in cycles
of nine years. Phelps and Belding believed that an oceanic—or possibly a river—
condition affected the migrating smolts periodically. What connection, if any, there
may be between such periodicity and the nine- or ten-year periodicity in terrestrial
animals such as the varying hare (Lepus americanus), rufted grouse (Bonasa umbellus),
and some others is not known (40: 1-34).
Atlantic Salmon catches in Great Britain and Ireland also fluctuate and show a
striking similarity to those in America (88). Svardson, while noting a tendency toward
parallel trends in the yields of Atlantic Salmon fisheries in Great Britain, Norway,
Sweden, and Holland, has pointed out that they are not parallel every year (124: 226—
262). He believes that he found a statistically significant positive correlation between
the area of ice cover in the Baltic and the Atlantic Salmon catch five years later, sug-
gesting, in explanation, that severe winters cause the death of large numbers of the
common porpoise, which preys on them. Lindroth, however, questioned Svardson’s
conclusions, arguing that the short-term fluctuation is only an artifact created by the
long-term fluctuation, which is dependent on climate (94).
Maintenance and Restoration of the Fishery. The problem of maintaining or re-
storing the species is largely one of assuring that conditions necessary for successful
spawning, hatching, survival, and growth in fresh water are maintained or restored.
There are no reasons for believing that the sea could not support as large populations
today as it did hundreds of years ago; therefore it is in the rivers that remedial action
is necessary.
Salmon thrive best under the primitive conditions of northern regions. They
require cold water and, for reproduction, clean gravel. The changes incidental to settle-
ment and industrial development that destroy these conditions have already been
mentioned under Decline in Abundance. The problem of preserving Salmon is that of
halting these injurious forces and, when possible, of restoring favorable conditions.
The failure of experiments to reintroduce Atlantic Salmon into a stream flowing into
Lake Ontario where they formerly spawned illustrates the detrimental effects of man’s
activities (98). Apart from mill dams that formerly barred the fish from reaching the
spawning grounds, the changes brought about in the conditions of this stream were
due to lumbering, which removed the tree cover, and to farming, which exposed the
soil to erosion. High lethal temperatures eliminated the planted fry from some sections,
and over the balance of the planted part, the greatest mortality occurred soon after
planting when heavy predation by other species of fish occurred. The extent of preda-
tion was related largely to the amount of suitable shelter for the fry. This was limited
Fishes of the Western North Atlantic 489
generally to gravelly ripples where the degree of bottom silting determined the amount
of shelter.
Measures necessary to improve streams for Atlantic Salmon include: better land-
use practices to reduce erosion, floods, and low summer flow; overcoming pollution
and the detrimental effects resulting from measures undertaken for the benefit of other
resources (e. g. spraying forests to control injurious insects); and means of assuring
reproduction in streams used in the development of hydroelectric power or irrigation.
The cost involved in adopting remedial measures will determine the extent to which
suitable streams will be saved for Atlantic Salmon production. If society values this
species highly enough, its future can be assured through wise management based on
research. Overcoming pollution, for instance, is largely a matter of deciding on the
relative importance or value of Atlantic Salmon compared with the value of industries
that cause pollution. If the cost of disposing of polluting substances by means other
than putting them into streams will be greater than the value placed on the fishery,
then the latter will be sacrificed; and, in considering the worth of this species, values
other than those of economics must be considered, such as the sports value.
In the past, relatively too much emphasis has been placed on restricting the catch
and too little on maintaining suitable stream conditions. Recent studies indicate that, in
many streams, enough spawners escape capture to provide all the young that can
survive and grow in the streams in their present condition (42). That this number is
not large is shown by the results of studies in four streams of the Maritime Provinces,
where it was found that between 40 and 50 pounds of adult females per mile of stream
10 yards wide are enough to maintain stocks (43: 23).
The role of hatcheries in maintaining Salmon populations is not as great as it once
was believed to be, but hatcheries have an important place if the young fish produced
in them are used intelligently. If the eggs normally deposited by the adults at the
spawning grounds are adequate to produce all the young that can survive and grow in
the stream, the planting of more young is not only wasteful but may actually be harm-
ful by increasing the competition for limited living room and food. However, in
streams that lack suitable grounds but where conditions are suitable for the survival
and growth of young to the smolt stage, the planting of hatchery-reared young may
be beneficial. Also, in streams barred by dams over which it is not practical to pass
spawning adults, the planting of hatchery-reared young may be advantageous if means
can be provided for them to reach salt water. Where the use of streams by adults and
young has been made impossible, a fishery may be preserved by raising young to the
smolt stage under artificial conditions and planting them directly in an estuary or in
the sea. Another promising method of providing artificial propagation is in creating
artificial spawning beds to replace those eliminated by power development or other
detrimental factors.
In Sweden the experimental planting of marked smolts has given a return of 5—
10 °/) in adults, a higher percentage than is usual; this suggests that artificial propa-
gation may be sound both biologically and economically (32). Efforts to rehabilitate
490 Memoir Sears Foundation for Marine Research
this species in Maine have met with some success; at least seven rivers there (versus
two in 1925) now have Salmon populations, with sizable runs in four or five of them (45).
Relation to Man. The Atlantic Salmon has long been highly esteemed as a table
fish; in fact, its bones have been found commonly in caves occupied by prehistoric man
in western Europe. Ausonius, a Latin poet of the fourth century, was the first to
describe fishing for this species—in the Moselle River, a tributary of the Rhine.
Since the Atlantic Salmon is prized as a game fish as well as a commercial species,
it is difficult, if not impossible, to estimate its true monetary value. Angling is one of
the most popular of outdoor recreations, and the Atlantic Salmon’s value is indicated
by the relatively large sums of money that sportsmen are willing to spend to fish for it.
Salmon fishing ranks with the higher satisfactions of life for which some men are willing
to pay handsomely after their physical needs have been satisfied. The value of the
Canadian commercial catch is in the neighborhood of a million dollars a year, and it
has been estimated that the angling fishery is worth more than this amount (44).
Range of Sea-going Populations. The general range of Salmo salar includes the
coasts of northern Europe and North America, from the Arctic to Portugal in Europe
and to New England in America. In Europe the northern limit is the Tschernaja River,
which flows into the White Sea east of the Pechora. The southern limit is the Mifio
River at the boundary between Spain and northern Portugal; perhaps a few reach the
Douro River midway of Portugal (123: 18; 24: 353; 109: 3543; ITO: 355; 96: 155).
There are many good Atlantic Salmon rivers in Iceland.
The Atlantic Salmon is relatively scarce in Greenland; it is known mainly from
the southwest coast. Before the 1920’s it was known from small numbers at only
Kapisigdlit (Godthaab Fjord, 64°10’ N) and Amerdlok Fjord (near Holsteinsborg, just
within the Arctic Circle). However, during the 1920’s, according to Jensen, it increased
off western Greenland with rising sea temperatures (76: 17, 18). Towards the end of
that decade a migration occurred in October and November in the Sukkertoppen
district (65°22’N), and later, especially in 1935 and 1936, the Salmon occurred in
large numbers at several places in the same district from October on into December.
In 1952 is was reported for Amerdlok Fjord, Ikerasak (Sukkertoppen district), and
Kapisigdlit (39: 92). In 1951 an Atlantic Salmon was found in the stomach of a shark
taken in the Umanak River (about 70°20’N), north of Disko Island; it must have
been devoured only a short time before the shark was caught because the Salmon
was so fresh that the natives ate it. The northernmost river in which the Atlantic
Salmon was known to spawn in 1953 was near Kapisigdlit (53: 79-81).
On the North American mainland the Atlantic Salmon ranges north to Hudson
Strait. There was a commercial fishery in Ungava Bay from 1881 till the early 1930’s
(39), and there still are spawning migrations into the larger rivers on the eastern half
of the Bay between and including the George and Koksoak rivers. The latter is at
present the western limit of its normal distribution in this area. According to native
report, it is occasionally taken as a stray in the Leaf River, farther west; and a fishing
camp for anglers has recently been established on the George River (148: 215).
Fishes of the Western North Atlantic 491
The northernmost Salmon rivers on the Atlantic coast of America are “just north
of Hopedale, about 350 miles north of the Strait of Belle Isle; the Adlok near Hopedale
has a run of salmon which will average well over ten pounds” (148: 216).
Southward there are regular spawning runs in all the larger rivers of Newfound-
land, including outer Labrador, Quebec as far into the Gulf of St. Lawrence as Bay
St. Paul and Kamouraska (1732), Prince Edward Island, New Brunswick, and Nova
Scotia. The northern limit of intensive commercial exploitation for it is Indian Harbor
on the north shore of Hamilton Inlet, about 54°N (z8).
In New England during colonial days the Atlantic Salmon occurred in every suit-
able stream southward to Connecticut, but after the early part of the past century, dam
construction (p. 485) was responsible for the elimination of the Atlantic Salmon from
many of the rivers. In the decade 1850-1860 it still entered the St. Croix, Dennys,
East Machias and Machias, Penobscot, Sheepscot, Kennebec, and Androscoggin. At
present the St. Croix, Dennys, Machias and East Machias, Narraquagus, and Pleasant
rivers alone have regular runs large enough to attract some anglers, and a few still
enter the lower reaches of the Penobscot, where a Federal hatchery is in operation.
Southward, enough Atlantic Salmon to yield a supply of eggs for artificial hatching con-
tinued to enter the lower reaches of the Merrimack until 1896 at least (doubtless
wanderers from the north), and while none has been reported for the Merrimack since
1901, it is not unusual for some, straying from the north, to be reported off Massa-
chusetts. Thus in 1928, 1,600 or so fish (16,050 pounds) were taken offshore there
(on long lines and in otter trawls), while in 1937 the floating traps along the north
shore of Massachusetts Bay picked up 4,400 pounds. A few are taken in Cape Cod
Bay in most years; there are records for the Woods Hole region, as well as for New
Jersey; and in April 1893, one of 16 pounds was taken with a school of mackerel off
Delaware, where it is said to have been introduced (48). However, it no longer runs
in any river south of Maine.
Freshwater Populations. In addition to anadromous populations, there are in both
Europe and North America populations consisting of fish that spend their entire life
in fresh water; the counterpart of the anadromous Salmon’s sea phase is passed in some
lake. They are sometimes referred to as “landlocked,” but this term is not accurately
descriptive, for they could all go to sea, but many of them could not return to their natal
rivers because of impassible falls. However, the Shubenacadie Lake population (Nova
Scotia) can freely pass to and from salt water. Lake Salmon is a more appropriate name
than landlocked Salmon for these nonmigratory populations (68: 289).
There is no agreement as to whether their habit of remaining in fresh water is
hereditary or not. Rounsefell has found that various species of salmonid fishes are
characterized by different degrees of anadromy (r1r7). Several species besides the At-
lantic Salmon, notably Salmo gairdneri and Salvelinus alpinus, have given rise to lake
populations. In experiments with marked young of both native Lake and Atlantic Salmon
planted in Lake Shubenacadie, Huntsman observed no differences in their migratory
behavior; however, the fish used were too few to provide final conclusions. Wilder
4.92 Memoir Sears Foundation for Marine Research
has suggested that the observed differences in coloration, spots, and flesh color in Lake
and Atlantic Salmon result solely from environmental differences (145).
Some of the lake populations were once regarded as being composed of distinct
species, but they are now generally considered subspecific. Instead of being distin-
guished from anadromous Atlantic Salmon by the scientific names Salmo sebago, S.
ouananiche, and S. hardinii, they are usually referred to now as Salmo salar sebago, etc.
Lake populations in North America are now found in Vermont, New Hampshire,
Maine, New Brunswick, Nova Scotia, Quebec, Newfoundland, and Labrador, and
formerly in Lake Ontario. They occur also in Norway, Sweden, and Russia.
Synonyms and References :1®
Salmo salar Linnaeus, Syst. Nat., IT, 1758: 308 (European seas); Fabricius, Fauna Groenl., 1780: 170 (rep. w.
Greenland; not seen); Bloch, Naturg. Fisch. Dtsch., 7, 1782: 128, 2 pls. (descr., habits, German fishery);
Bonnaterre, Tab]. Encyc. Meth. Ichthyol., 1788: 158, tab. 1 (descr.); Walbaum, J. F. Kleinii Ichthyol.
Enodata, 1793: 109 (Klein’s [1749: 109] Trutta (3) dentata ident. as Sa/mo salar); Bloch and Schneider,
Syst. Ichthyol., 1801: 398 (descr.); Donovan, Nat. Hist. Brit. Fish., 1802-1808: plates only; Lacépéde,
Hist. Nat. Poiss., 5, 1803: 159 (genl.); Turton, C. Linné, Genl. Syst. Natur., Amend., 2, 1806: 846
(descr., habits, n. Europe); Mitchill, Trans. Lit. philos. Soc. N.Y., Z, 1815: 435 (extinction from
Connecticut R., casual occur. Hudson R.); Davy, H., Salmonia, 1828: 273 pp. (descr., habits); Fleming,
Hist. Brit. Anim., 1828: 179 (descr., life hist.); Jenyns, Manual Brit. Vert. Anim., 1835: 421 (descr.,
habits); Richardson, Fauna Boreal-Amer., Pt. 3, 1836: 145-157, pl. 91, fig. 1 (distr., descr., life hist.) ;
Shaw, Edinb. New Philos. J., 22, 1836: 99 (expts., observ. proving parr young salmon); Yarrell, Brit.
Fish., ed. 1, 2, 1836: 1, 6 figs. (descr., excell. ill., habits, abund., size, 83-lb. female); Henry, Trans.
Lit. Hist. Soc. Quebec, (1) 3, 1837: 347 (habits, Canada); Kroyer, Danmark’s Fiske, 2, 1838: 53-540—
581 (descr., meas., genl. acct.; Danish); Parnell, Mem. Werner. Soc. Edinb., 7, 1838: 278, pl. 32,
fig. 1, pl. 30 (descr., size, habits, life hist., diet, Scotland); Jardine, Brit. Salmonidae, 1 and 2, 1839-1841
(color ills.; not seen); Shaw, Trans. roy. Soc. Edinb., 74, 1840: 546, 2 pls. (parr growth, transform. to
smolts); De Kay, Zool. N.Y., 3, 1842: 241, 242, pl. 38, fig. 122 (descr., former abund. L. Ontario
and trib. of Hudson R.); Hamilton, Brit. Fish., 2, 1843: 116 (good genl. acct., parr growth, after
Shaw); Lilljeborg, Ofvers. Vet. Akad. Forh. Stockh., 1849: 35, 36 (European trout cf. 8. ruta);
Perley, Fish. Gulf St.Lawrence, 1849: 126 (genl.); Fitzgibbon (Ephemera), Book of Salmon, 1, 2,
1850: 157-242, 3 pls., 5 figs. (fishing, flies, life hist., size, artif. propag., color ills. parr, smolt); Herbert
(Frank Forester), Fish Fish. U.S. and Brit. Prov., Canada, ed. 2, 1850: 54, 1 pl., 4 figs. (descr., life hist.);
Perley, Sea and River Fish. N. Brunsw., ed. 2, 1850 (status of fishery); Gosse, Nat. Hist. Fish., 1851:
232, I fig. (genl., fly fishing in sea, size); Perley, Cat. Fish. N.Brunsw. and Nova Scotia, 1852: 199 (life
hist.); Nilsson, Skand. Fauna, Fisk., 4, 1855: 370 (refs., descr., genl. acct., Scandinavia, lgth.-wet.
relation.); Thompson, Nat. Hist. Ireland, 4, 1856: 143 (food, large specimens abund., fishery, Ireland,
parr descr.); Adamson, Canad. J. Industr. Sci. Art, (N.S.) 2, 1857: 1-7, 1 pl. (decrease, restor., preserv.,
Canada); Davy, J., Trans. roy. Soc. Edinb., 2Z, 1857: 253 (doubts small salmonids, closely re-
sembling parr, a distinct species); Nettle, Salmon Fish. St. Lawrence, 1857: 1-129 (descr. fishery); Storer,
Mem. Amer. Acad. Arts Sci., 6 (2), 1858: 320, pl. 25, fig. 2 (refs., descr., former abund. Merrimack R.);
Alexander, Salmon-fishing Canada, 1860: 1-286 (acct. of salmon rivers) App. 1 = Adamson, 1857, App.
11 = Henry, 1837; Jardine, Edinb. New Philos., (N.S.) 15, 1862: 203-231 (fishery in England, Wales);
Roosevelt (Barnwell), Game Fish. north. states Amer. and Brit. Prov., 1862: 88-115 (descr., habits,
salmon rivers, life hist.); Schlegel, Natuurl. Hist. Nederl., Vissch., 1862: 126, pl. 13, fig. 1 (genl.;
19. References to the Atlantic Salmon are so numerous that only part of them are included here. Those listed provide
information about the Salmon in the western North Atlantic and in fresh water of North America, but some
European contributions dealing with taxonomy, distribution, and life history that are of particular importance are
also included. Publications repeating essentially the same information or results have been omitted for the most
part. References to Salmon are also included in most natural histories and in many annual reports or documents
on fish or fisheries by government departments of the United States, Canada (particularly the Fisheries Research
Board of Canada), and European countries where Salmon occur.
Fishes of the Western North Atlantic 4.93
Dutch); Widegren, Ofvers. Vet. Akad. Forh. Stockh., 1862: 541, tab. 4, figs. 3, 4 (descr., meas., ills.
of parr, cf. lake salmon [8. /acustris] and trout; Danish); Lemoine, Les Péch. Canada, 1863: 40-46,
64-76 (salmon rivers n. shore St. Lawrence, genl. acct.); Knight, Fish. N. Brunsw., 1863 (descr.
fishery); Couch, Fish. Brit. Isles, 4, 1864: 163, pl. 211 (life hist., hist. of fishery); Russell, A., The
Salmon, Edinburgh, 1864: 234 pp. (life hist., value, decrease, legislation); Norris, Amer. Angler’s Book,
ed. 2, 1865: 206, 3 figs. (nat. hist., art of taking); Austin, Trans. Lit. Hist. Soc. Quebec, (N.S.) 4,
1866: 115 (St. Lawrence R.); Blanchard, Poiss. des eaux douces... France, 1866: 448 (genl. acct.);
Gilpin, Proc. N.S. Inst. Sci., r (4), 1866: 76 (descr., life hist.); Giinther, Cat. Fish. Brit. Mus.,
6, 1866: rr (synon., refs., descr., life hist.); Hamilton, Naturalist’s Library, ed. Jardine, 37, 1866:
116-127 (summ.); Knight, T’., Descr. Cat. Fish. Nova Scotia, 1866: 10 (salmon); Fisher. Nova Sco-
tia, 1867: 18 (life hist.); Fish Fisher. Nova Scotia, 11, Shore and deep sea fisheries, 1867: 56, 57 (cap-
ture, curing, export); 11, River fisheries, 1867: 4—8 (life hist.); Storer, Fish. Mass., 1867: 142, pl. 25,
fig. 2 (see Storer, 1858); Norris, Amer. Fish Cult., 1868: 102 (cf. salmon of diff. rivers, former abund.,
causes of decline, migr.); Gilpin, Ann. Rec. Sci., 1872: 446 (winter quarters Nova Scotian salmon);
Atkins, Rep. U.S. Comm. Fish. (1872-1873), 2, 1874: 226-288 (artif. cult.), 289-335 (hist. of fish-
ing New England rivers, growth, migr.); Hamlin, Rep. U.S. Comm. Fish. (1872-1873), 2, 1874: 338
(landlocked salmon); Lanman, Rep. U.S. Comm. Fish. (1872-1873), 2, 1874: 223 (time of river en-
trance, spawn.); Stone, Rep. U.S. Comm. Fish. (1872-1873), 2, 1874: 216 (date of runs, spawn., size,
Miramichi R.); Suckley, Rep. U.S. Comm. Fish. (1872-1873), 2(B), 1874: 104 (doubts ident. of
Amer. species with that of Europe); Collett, Norges Fiske, 1875: 155 (genl., variations); Liitken, Cat.
Fish. Groenl., revised, iz Jones, Manual Nat. Hist. Geol. Greenl., 1875 (not seen); Scott, Fishing Amer.
Waters, 1875: 215-251 (fishing, St. John R., Gaspé, Quebec); Malm, Goteborg Fauna, 1877: 534—
535 (varieties or races); Hallock iz Goode, Game Fish. United States, 1879: 17-49, 1 fig. (genl., Amer.
rivers); Houghton, Brit. Freshw. Fish., 1879: 83, 1 pl. (life hist.); Jones, Trans. N.S. Inst. Sci., 5, 1879:
92 (Nova Scotia, yearly var. in abund.); Giinther, Guide to Study Fish., 1880: 644 (descr.); Hind,
Forest and Stream, 74, 1880: 126 (movements in sea); Day, Fish. Gt. Brit. Ireland, 2, 1880-1884:
63, pl. rr0, figs. 1, 2 (ill. old and young, genl., bibliogr.); Buckland, Nat. Hist. Brit. Fish., 1881: 2g2—
295 (lists enormous salmon); Moreau, Hist. Nat. Poiss. Fr., 3, 1881: 525 (genl., rivers, France); Gilpin,
Proc. N.S. Inst. Sci., 5 (1), 1882: 38, 1 pl., 4 figs. (descr., life hist.); Jordan and Gilbert, Bull. U.S.
nat. Mus., 16, 1882: 312 (descr.); Lord, Bull. U.S. Fish Comm., 7, 1882: 274 (wgts., Restigouche,
1879); Smitt, Ofvers. Vet. Akad. Forh. Stockh., 39 (8): 1882: 32 (distr., White Sea salmon named var.
brevipes); Miescher-Riisch, Contr. Biol. Rhine Salmon, Rep. U.S. Bur. Fish. (1880), 8, 1883: 427 (early
studies, body changes in fresh water, age of first spawn., and frequency); Goode, et a/., Fish. Fish. In-
dustr. U.S., 7, 1884: 468-474; 2, 1887: pl. 186 (distr., abund., habits, life hist.); Joncas, Fisher. Canada,
1884: 28-30 (val. of fishery, import. of conserv.); Garman, 1gth. Rep. Mass. Comm. inl. Fish., 1885:
8, figs. 2-4 (descr., good line ills.); Sundman and Reuter, Finlands Fiskar, 1885: 11, tab. 18 (excell.
color ill., adult; Sweden, England); Pennell, Sport. Fish Gt. Brit., 1886 (list of “proved facts”; summ.
to date); Lilljeborg, Sverige Norg. Fisk., 2, 1886: 551 (genl. acct.; Sweden); Seeley, Freshw. Fish.
Europe, 1886: 267-278, 2 figs. (genl.); Whiteaves, Cat. Canad...., Fish., 1886: 1-30 (local. recs.);
Day, Brit. Irish Salmonidae, 1887: 51-142, pls. 3, 4, 6 figs. (excell. genl. acct., many refs., color ills.
parr, smolt, adults); Goode, Amer. Fish., 1887: 441, 4 figs. (genl., abund. early days); Day, Trans. linn.
Soc. London, Zool. 2 (2), 1888: 447 (breed. parents never descended to sea); Sage, Restigouche and
Salmon Fisher. (1882), 1888: 275 pp. (fishery, Restigouche R., Canada; not seen); Traherne, Habits...
Salmon, 1889: 1-159 (genl., Gt. Brit.); Fatio, Fauna Vert. Suisse, 5 (Poiss.), 1890: 298 (descr., racial
diffs., Switzerland); Hallock, Amer. Game Fish., 1892: 17-49, 1 fig. (habits, habitat, angling); Smith,
Bull. U.S. Fish Comm., 14, 1895: 95 (captures at sea and coast. waters of e. states); Smitt, Hist. Scand.
Fish., 2, 1895: 827, pl. 37, figs. 3, 4, pl. 38, fig. 1 (descr., refs., Scandinavia, 8. ¢rutta consid. var. ot
8. salar); Chambers, Ouananiche and Canad. Environ., 1896: 1-x11+1-329 (descr., habits, distr.
in freshw., salmon, Lake St. John, Quebec); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896:
486 (descr., distr.); Montpetit, Poiss. d’eau douce Canada, 1897: 305-330, 2 pls., 3 figs. (distr.,
abund. in Canada, migr., life hist.); Smith, Bull. U.S. Fish Comm., 77, 1898: 113-124 (Maine);
Paton, Rep. Fish. Bd. Scotl. Salmon Fish. (1898), 1899: 1-174 (Scottish fishery, 1898); Ehrenbaum, in
Rémer and Schaudinn, Fauna Artica, 2, Fische, 1901: 128, 129 (e. Arctic Atlant. to n. Scandinavia,
Murman cst., White Sea, as 8. sa/ar var. nobilis); Calderwood, Rep. Fish. Bd. Scotl. Salmon Fish., 1g01-
1926 (migr., life hist.); Barrett-Hamilton, Ann. Mag. nat. Hist., (7) 9, 1902: 106-120 (life hist., nup-
494 Memoir Sears Foundation for Marine Research
tial and sex. ornament., develop.); Bean, 7th Rep. Forest Comm. N.Y. (1901), 1902: 344, 1 fig.; John-
son, Fish. New England Canada, I, 1902: 1, 4 pls. (descr., range, life hist., habits); Paton and New-
begin, Proc. roy. Soc. Edinb., 23, 1902: 44—54 (changes in wgt., muscles, gonads in fresh water, phos-
phorous content, pigment sources); Bean, Bull. N.Y. St. Mus., 60, Zool. 9, 1903: 244, I fig. (descr.,
life hist., early reps. for Hudson R., New York); Goode, Amer. Fish., ed. 2, 1903: 441-453, 4 figs.
(habits, life hist.); Stafford, Zool. Anz., 28, 1904: 681 (four spp. trematodes from Canad. salmon);
Wilson, Proc. U.S. nat. Mus., 28, 1905: 640 (Lepeophtheirus salmonis, parasite); Johnson, Rep. Fish. Bd.
Scot., 1905-1910 (life hist. through scale studies); Cornish, Contr. Canad. Biol. (1902-1905), I0, 1907:
83 (Canso, Nova Scotia); Stafford, Contr. Canad. Biol. (1902-1905), IO, 1907: g1 (four spp. trem-
atodes in salmon); Calderwood, Life of Salmon, ed. 2, 1908: 160 pp., 7 pls. (salmon protection, life
hist.); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 44 (distr., New England); Malloch, Life
Hist. and Habits Salmon. .., 1910: 1-124, 130, fig. (life hist., sea movements, diseases); Dahl, Age
and Growth ... Norway, 1911 (genl. life hist., bibliogr.); Regan, Fresh Water Fish. Brit. Isles, rg11:
26-53, pl. 2, fig. 5 (distr., descr., life hist.); Halkett, Check List Fish. Canada, 1913: 51 (distr.); Master-
man, Bd. Agric. Fish., Fish. Invest., London, (1) 2, 1913: 1-111 (early rep., scale study); Dahl, Salmon
and Trout, Handbk., 1914: 1-57 (genl. life hist., bibliogr.); Regan, Ann. Mag. nat. Hist., 13, 1914:
405-408 (systematics, Salmonidae); Meek, Migrations of Fish, 1916: 115-122 (migr.); Heitz, Arch.
Hydrobiol., 72, 1920: 311-373, 485-561 (parasites, salt and fresh water); Calderwood, Salmon Rivers
and Lochs Scotland, ed. 2, 1921 (artif. freshets to bring salmon up from sea); Kendall, Bull. U.S. Bur.
Fish., 37, 1921: 183-208, fig. 7 (relation of ovaries to peritoneal membranes, §.sa/ar); Huntsman,
Contr. Canad. Biol. (1921), 3, 1922: 11-12 (Bay of Fundy); Hutton, Life-Hist. Salmon, 1924: 56 pp.,
17 pls. (summ., life hist. through scale studies; not seen); Sage, Salmon and Trout, 1924: 1-36, 1 pl.
(hist., habits); Bigelow and Welsh, Bull. U.S. Bur. Fish., 4o (1), 1925: 130-138, 1 fig. (descr., life
hist., Gulf of Maine); Hewitt, Secrets of Salmon, 1925: 3-23, 4 figs. (life hist., habits); Jenkins, Fish.
Brit. Isles, 1925: 215-227, pls. 87—g2 (life hist., color ills. of diff. stages, great wgts., Gt. Brit.); McFarland,
Salmon of Atlantic, 1925: 139 pp., 10 pls. (hist., anat., physiol., disease, habits); Menzies, The Salmon,
1925 (life hist., migr.); Calderwood, Proc. roy. Soc. Edinb., 45, 1926: 142 (growth, spawn. frequency);
Proc. roy. Soc. Edinb., 45, 1926: 334-345 (Moisie R. salmon, Canada); Rushton, Salm. Trout Mag.,
44, 1926: 275 (teeth of freshw. salmon, kelts); Calderwood, Proc. roy. Soc. Edinb., 47, 1928: 142—
147, 2 pls. (Grand Cascapedia R. salmon, Canada); Menzies and Macfarlane, Proc. roy. Soc. Edinb.,
47, 1928: 359-363 (Moisie R. salmon, Canada); Palmer, Salmon Rivers Nfdld., 1928: 21-246 (New-
foundld. rivers, with angling characterized for each); Breder, Field Bk. Mar. Fish. Atl. Cst., 1929: 73
(distr.); MacFarlane, Proc. roy. Soc. Edinb., 48, 1929: 134-139, 1 pl. (Moisie R. salmon, Canada);
Regan, Encyc. Brit., ed. 14, 9, 1929: 889-890 (genl., includ. taxon.); Scheuring, Ergbn. Biol., 5, 1929:
405 (summ., discus. of lit. on migr.; not seen); Bere, Contr. Canad. Biol. Fish., 5 (13), 1930: 423
(parasitic copepods, Passamaquoddy, New Brunswick region); Calderwood, Salmon and Sea Trout,
1930: I-85, 114-232, 11 pls. (life hist., habits, management); Survey Salmon Fisher. e. Canada,
1930; Notes on Salmon Fisheries Newfoundld., 1930; Griswold iz Griswold and Hume, Life Hist.
Canadian Salmon, 1930: 75-84 (demonstrates g-10 year periodicity in catch at Cascapedia and
Restigouche Salmon clubs and in commerc. catch in New Brunswick and Quebec, numbers, sizes, diff.
rivers); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 56 (distr., checklist);
Phelps and Belding, Invest. Conditions Affect. Salmon Fisher., Bay Chaleur and Adj. Waters, 1930;
Calderwood, Salmon Hatching ... Migration, 1931: 4 pls.; Huntsman, Bull. biol. Bd. Canad., 27,
1931: 99 pp-, 29 figs. (g- or 10-year periodicity; summ. Canad. salmon); Norman, Hist. Fish., 1931
(refs.); Phelps and Belding, Statist. Study Salmon Fishing Restigouche, 1931 (9- or 10-year periodicity
extend. back to 1879); Belding, Trans. Amer. Fish. Soc., 62, 1932: 304 (ocean. feed. grounds, wgts.,
2 sea-year Canadian and English salmon); Belding, Pender, and Rodd, Trans. Amer. Fish. Soc., 62, 1932:
211 (early parr growth, Canada); Berg, Zoogeographica, I (2), 1932: 112 (see Smitt, 1882); Lindsay
and Thompson, Rep. Nfdld. Fish. Comm., 2, 1932: 7-80, 8 pl., 1 chart, 7 figs. (salmon in New-
fdld.); Nobre, Biol. Min. Agric. Lisboa, (1) 13 (2), 1932: 351 (50-cm specimen, Rio Leca, N. of
Oporto, Spain); Allen, Trans. N.S. Inst. Sci., 8, 1933: 34-49 (phys. changes, early develop.); Hunts-
man, Progr. Rep. Biol. Bd. Can., Atl. St.,6, 1933: 7-10 (Nov. entrance in St. John R., New Brunswick,
estuary, spawn. after Oct.); Roule, Fishes, Journeys and Migrations, 1933: 63-66 (assumed migr.);
Alm, Rapp. Cons. Explor. Mer, 92, 1934: 1 (Baltic); Belding, Trans. Amer. Fish. Soc., 64, 1934: 211
(spawn. habits); Trans. Amer. Fish. Soc., 64, 1934: 219-222 (mortal. causes after spawn.); Belding
Fishes of the Western North Atlantic 495
and Kitson, Trans. Amer. Fish. Soc., 64, 1934: 225-230 (races); Dymond and Vladykov, Proc. 5th
Pan-Pacif. Sci. Congr., 5, 1934: 3743 (distr.); Huntsman, Trans. Amer. Fish. Soc., 64, 1934: 351—
355 (effect of temp., river flow, wind, water movement on return from sea); Kuitunen, Contr. Canad.
Biol. Fish., 8 (6), 1934: 89 (cestode parasite); Kobbs, Salmon Tactics, 1934 (genl.); White, Trans.
Amer. Fish. Soc., 64, 1934: 360-362 (does presence of young cause adults to select spawn. stream ?);
Belding, Trans. Amer. Fish. Soc., 65, 1935: 157-160 (parr growth); Blair, J. Biol. Bd. Canad., 7, 1935:
159-169 (ages at migr., Miramichi R., Canada); Kendall, Mem. Boston Soc. nat. Hist., g (1), 1935:
17-103, pls. 1-4 (max. sizes, range, captures at sea, feed., life-cycle, angling New England); Nobre,
Fauna Marinha Port., r, Vert., 1935: 344 (Rio Leca specimen, after Nobre, 1932); Huntsman, Bull.
Biol. Bd. Canad., 51, 1936: 22 pp., 10 figs. (return from sea); Melangon, Poiss. de nos Eaux, 1936:
60-66, 1 fig. (life hist.); Siedlecki, Rapp. Cons. Explor. Mer, Zor (3) 9, 1936: 10 pp., 6 figs. (fluct.,
strength of year-classes, Europe); White, J. Biol. Bd. Canad., 2, 1936: 299-309, 2 fig. (kingfishers and
mergansers as predators); J. Biol. Bd. Canad., 2, 1936: 391-400, 4 figs. (homing of salmon); J. Biol.
Bd. Canad., 2, 1936: 499-506, 2 figs. (food of fry); Belding, Trans. Amer. Fish. Soc., 66, 1937: 211
(Newfoundld. parr); Allen, Rep. Freshw. Biol. Ass. Brit. Emp., 5, 1937: 31 (biol. early stages); Dahl,
Salm. Trout Mag., 86, 1937: 10-13 (parr life); Calderwood, Salm. Trout Mag., 88, 1937: 207-213
(homing instinct); Huntsman, Science, 85, 1937: 313-314, 582-583 (homing and races); Trans. roy.
Soc. Canada, 32 (V), 1937: 17-27 (causes of periodic scarcity); Russell, Biol. Rev., 12, 1937: 329-331
(migr.); Tchernavin, Salm. Trout Mag., 88, 1937: 235-242 (descr. skull); Belding, Trans. Amer. Fish.
Soc., 67, 1938: 195-206 (salmon in Moisie R., Canada); Belding and Clark, Trans. Amer. Fish. Soc.,
67, 1938: 184 (parr in Margaree R., Canada); Calderwood, Salmon Experiences Reflections, 1938:
175 pp.) 15 pls. (genl.); Huntsman, Nature, Lond., 141, 1938: 421, 422 (homing instinct); Rapp. Cons.,
Explor. Mer, ror (4), 1938: 9-15 (strength of year-classes, N.Amer.); J. Fish. Res. Bd. Canad., 4,
1938: 96-135 (sea movements, Canadian Atlant. kelts); Salm. Trout Mag., go, 1938: 24 (case against
heredit. instinct); Trans. roy. Soc. Canad., 32, 1938: 63-70 (life hist., mortal. in fresh water and sea);
Orton, Jones, and King, Proc. roy. Soc. London, B 125, 1938: 103-114 (male sex. stage in parr, lit.);
Regan, Fishing Gazette, 726, 1938: 298 (ripe female parr); Schrenkeisen, Field Bk. Freshw. Fish. N.
Amer., 1938: 41-43 (occur.); Schultz, Smithson. Rep. (1937), 1938: 365-376 (breed. habits); T’cher-
navin, Trans. zool. Soc. London, 24, 1938: 103-184, 5 pls., 25 figs., 12 graphs (changes in skull); White
and Huntsman, J. Fish. Res. Bd. Canad., 4, 1938: 1-18, 5 figs. (no observ. diff. in behav. between
progeny of early- and late-run salmon planted in same river); Belding, Publ. Amer. Ass. Adv. Sci., 8,
1939: 32 (migr., conserv.); Carpenter, Nature, Lond., 143, 1939: 336 (parr food); Dahl, Salm. Trout
Mag., 94, 1939: 19-26, I fig. (homing instinct); Hoar, J. Fish. Res. Bd. Canad., 4, 1939: 441-460
(Igth.-wet. relation); J. Morph., 65, 1939: 257-295 (Atlant. salmon thyroid); Huntsman, Salm. Trout
Mag., 96, 1939: 233-237 (races, homing instinct); Bull. Fish. Res. Bd. Canad., 57, 1939: 75 pp-s
34 figs. (descr., salmon for angling in Margaree R., Canada); Publ. Amer. Ass. Adv. Sci., 8, 1939: 32-44,
8 figs. (migr., conserv.); Huntsman and Hoar, J. Fish: Res. Bd. Canad., 4, 1939: 409-411 (parr’s resist.
to sea water); Jensen, K. Dansk Vidensk. Selsk. Skr., Biol., 14, 1939: 17-18 (effect of climatic change
on distr., Greenl. salmon); King, Jones, and Orton, Nature, Lond., 143, 1939: 162 (spawn. of male parr);
Menzies, Publ. Amer. Ass. Adv. Sci., 8, 1939: 13, 11 figs. (migr., Europe); Scheer, Quart. Rey. Biol.,
I4, 1939: 408-430 (summ. parent stream theory); Tchernavin, Proc. phys. Soc. Edinb., 23, 1939:
73-78 (summ. parr spawn.); Salm. Trout Mag., 95, 1939: 120-140 (marine or freshw. origin); White,
Bull. Fish. Res. Bd. Canad., 58, 1939: 30 pp-, 13 figs. (increase of smolts after control of kingfishers,
mergansers); White, J. Fish. Res. Bd. Canad., 4, 1939: 323 (factors infl. descent of smolts); Allen, J.
Anim. Ecol., 9, 1940: 1-23 (biol., early stages); Belding, Trans. Amer. Fish. Soc., 69, 1940: 285 (river
var.); Trans. Amer. Fish. Soc., 69, 1940: 290-295 (migr., Gulf of St. Lawrence); Calderwood, Salm.
Trout Mag., 100, 1940: 196-207 (marking, scale stud.); Jones and Orton, Proc. roy. Soc. London, B 128,
1940: 485-499, 3 figs. (over 50°/, male parr in Gt. Brit. sex. mature and spent before leaving river);
Rounsefell and Bond, Trans. Amer. Fish. Soc., 78, 1940: 189 (size expectancy of young); White, J. Fish.
Res. Bd. Canad., 5, 1940: 172-175 (sea lice causing death); Allen, J. Anim. Ecol., 10, 1941: 47-76 (feed.
habits, parr); Dymond, Canad. Field Nat., 55, 1941: 19 (Ungava Bay salmon); Friend, Trans. roy. Soc.
Edinb., 60, 1941: 503 (life hist., ecol. of gill maggot); Herrington and Rounsefell, Trans. Amer. Fish. Soc.,
70, 1941: 123-127 (restor., New England); Huntsman, J. Fish. Res. Bd. Canad., 5, 1941: 227-235 (rela-
tion of birds to cyclical abund.); Miller, Canad. J. Res. D., 79, 1941: 28 (trematodes); Elson, Trans.
7th N. Amer. Wildl. Conf., 1942: 202-210 (behav., survival rate, artif. planting, fingerlings, Nova Scotia) ;
4.96
Memoir Sears Foundation for Marine Research
Hoar, J. Fish. Res. Bd. Canad., 6 (1), 1942: 90 (diurnal feed.); Huntsman, Science, 95, 1942: 381-383
(return of marked Salmon from distant places); Trans. 7th N. Amer. Wildl. Conf., 1942: 195-201
(planting results); Orton, Nature, Lond., 150, 1942: 21-22, 1 fig. (temp. rel. to spawn.); White, J. Fish.
Res. Bd. Canad., 6, 1942: 24-29 (life hist. Lepeophtheirus salmonis); J. Fish. Res. Bd. Canad., 6, 1942:
37-44, 2 figs. (artif. spawn. beds); Alm, K. Landtbr. Styr. Medd., Stockh., 22, 1943: 1-40 (male parr for
fertil. eggs); Blair, Res. Bull. Newfdld. Dept. nat. Resources, £3, 1943 (Newfoundland and Labra-
dor salmon, 1939); Tchernavin, Proc. zool. Soc. London, B 173, 1943: 206-232 (skull changes in breed.
salmon, bibliogr.); Went, Proc. roy. Irish Acad., 49, 1943: 151-175 (changes, Shannon R. stocks 1929—
1941, Ireland); Huntsman, J. Fish. Res. Bd. Canad., 6, 1945: 311-325 (var. seaward migr.); J. Fish.
Res. Bd. Canad., 6, 1945: 399-402 (parr migr.); La Monte, N. Amer. Game Fish., 1945: 11, color. ill.
(distr., descr., food); Went, Sci. Proc. roy. Dublin Soc., 24, 1945: 1-8, 1 fig. (°/) of previously spawned
fish, Irish rivers); Huntsman, Salm. Trout Mag., 178, 1946: 234-257 (netting rel. to angling); Dymond,
Trans. roy. Soc. Canad., 41 (V), 1947: 1 (fluct. animal populations); Jones, Proc. zool. Soc. London, 117,
1947: 708, 1 pl., 1 fig. (distinct. between young salmon and trout); Huntsman, Science, 105, 1947:
289-290 (are Lake Salmon hereditarily distinct?); Menzies, Salm. Trout Mag., 120, 1947: 130-132
(spawn.); Regan, Encyc. Brit., ed. 14, 1947, 9: 314 (genl.); rg: 889; Huntsman, Trans. Amer. Fish.
Soc., 75, 1948: 257-266 (freshets effect. migr.); Salm. Trout Mag., 123, 1948: 153-158 (sea migr.);
Nature, Lond., 161, 1948: 300-305 (migr.); Jensen, Spolia Zool. Mus. Hauniensis, 9, 1948: 65-67
(Greenland); Power, Arctic, IT, 1958: 86—92 (evol. freshw. salmon); Dawson, The Salmon (Sa/mo salar)
in Game Fish. of World, 1949: 1-7 (genl., life hist.); Heller, Can. J. Res. D., 27, 1949: 243 (7 spp.
parasites, Canad. salmon); Jones and King, Proc. zool. Soc. London, 17g, 1949: 33-48, 2 pls., 3 figs,
(spawn. expts., review of spawn. behav.); Menzies, Stock of Salmon, 1949: 96 pp., 18 figs. (migr.,
preserv., improvement); Merriman and Jean, Copeia, 1949: 220-221 (Connecticut R.); Saemundsson,
Marine Pisces iv Zool. Iceland, IV (72), 1949: go (genl., Iceland); Huntsman, Salm. Trout Mag.,
I30, 1950: 227-239 (factors of migr.); Jones and King, Proc. zool. Soc. London, 120, 1950: 317-323
(further spawn. observ.); Salm. Trout Mag., 128, 1950: 24-26 (parr sperm prod. norm. progeny);
McCrimmon, Trans. Amer. Fish. Soc., 78, 1950: 128 (reintrod. in Lake Ontario); Vibert, Ann. St.
Centr. Hydrobiol. appl., 3, 1950: 27-149 (age, growth, spawn. cycle, races); Ackerman, Handbk. Fish.
Atlant. Seaboard, 1951: 106-107 (genl.); Hasler and Wisby, Amer. Nat., 85, 1951: 223-238 (discrimina-
tion of stream odors, parent stream behav.); Hewitt, Salm. Trout Mag., 132, 1951: 40-44 (salmon
rivers, Spain); Hoar, Publ. Ont. Fish. Res. Lab., 7Z, 1951: 1-51 (hormones); Menzies, Rep. Atl.
Salmon Fisher. Canada, Atl. Salmon Ass., 1951 (conserv. problems); Wolf, Salm. Trout Mag., 737,
1951: 68-76 (causes of decline, also papers in later numbers Salm. Trout Mag.); Collins, Fish. Bull.,
U.S. Fish Wildl. Serv., 73, 1952: 373-396 (orient. of migr. anadrom. fish.); Dunbar and Hilde-
brand, J. Fish. Res. Bd. Canad., 9, 1952: 92—94 (life hist., commerc. import. Ungava Bay; also refs. to
earlier authors); Hoar, Trans. roy. Soc. Canad., 46, 1952: 39 (thyroid function, anadromy); Huntsman,
Salm. Trout Mag., 736, 1952: 185-191 (wandering vs. homing); J. Cycl. Res., 1, 1952: 43-53 (causes
of cycles); Jones, Nature, Lond., 169, 1952: 882 (spawn. parr with female); Jones and King, Proc. zool.
Soc. London, 122: 1952: 615-619 (spawn. parr with female); Rostlund, Freshw. Fish Fishing N. Amer.,
1952: 4, 25, 26, 52, 258 (refs.); Bigelow and Schroeder, Fish. Bull. (74) U.S. Fish Wildl. Serv., 53,
1953: 121-131, 1 fig. (Gulf of Maine); Brainard, Atl. Salm. J., 3, March, 1953: 74—76 (Connecticut
R.); Harriman, Atl. Salm. J., Aug. 1953: 108 (restor. in Narraguagus R., Maine); Hansen and Hermann,
Danm. Fisk. Havunders., 15, 1953: 78-81 (Greenland); Hayes, Bull. Fish. Res. Bd. Canad., 99, 1953: 47
pp, 26 figs. (artif. freshets, etc., La Have R. populations); Hoar, Biol. Rev., 28, 1953: 437-452 (control.
fish migr.); Johnson, Salm. Trout Mag., 738, 1953: 135-153 (early and late runs); Rostlund, Copeia,
3, 1953: 192-193 (discus. Henry Hudson’s rep. of salmon in Hudson R., 1609); Sedgwick, Salm. Trout
Mag., 137, 1953: 77-82 (relation. of parr age and early or late return for first spawn.); White, Bull. Fish.
Res. Bd. Canad., 97, 1953: 44 pp., 117 figs. (kingfisher predation); Anon., Bull. U.S. Fish Wildl. Serv.,
88, 1954: 1-12 (genl. acct.); Huntsman, Nature, Lond., 774, 1954: 215-217 (water movements
re migr.); Jones and Ball, Brit. J. Anim. Behaviour, 2, 1954: 103-114 (spawn. behav.); McCrimmon,
J. Fish. Res. Bd. Canad., rr, 1954: 362-403 (planting results); Vibert, ‘Trans. Amer. Fish. Soc., 83,
1954: 194-201 (effect of sun and gravel on develop.); Wisby and Hasler, J. Fish. Res. Bd. Canad., rz,
1954: 472-478 (effect of olfactory occlusion on migr.); Wooding, Canad. Dept. Fish., 1954: (genl.
acct.); Alm, Rep. Inst. Freshw. Res., Drottn., 36, 1955: 13 (artif. hybridization); Carlin, Rep. Inst.
Freshw. Res., Drottn., 36, 1955: 57-74 (tag. smolts, Sweden); Elson, Prog. Rep. Fish. Res. Bd. Canad.,
Salmo
Salmo
Salmo
Salmo
Salmo
Fishes of the Western North Atlantic 497
Atl. St., 63, 1955: 13-15 (overfish.); Elson and Kerswill, Trans. 2oth N. Amer. Wildl. Conf., 1955:
415-426, 4 figs. (fluct. in catch, util., bird predation, effect of DDT spray., managemt.); Everhart, Watson,
and Cutting, Maine Atl. Salm. Comm. Dept., 1955: 14 pp. multilith (restor., Penobscot R.); Kerswill,
Atl. Salm. J., Z, Jan. 1955: 26, 3 figs. (recent develop. in research); Atl. Salm. J., 7, Jan. 1955: 30 (effect
of black salmon [kelt] on angling); Lindroth, Rep. Inst. Freshw. Res., Drottn., 36, 1955: 126-132
(mergansers as predators); Parson, Atl. Salm. J., 3, Sept. 1955: 16 (distance, speed of travel); Pyefinch,
Scott. Home Dept., Freshw. Salmon Fish. Res., 9, 1955: 1-24 (reviews biol.); Sedgwick, Salm. Trout
Mag., 143, 1955: 31-33 (runs in ice-free rivers any month of year); Svardson, Rep. Inst. Freshw. Res.,
Drottn., 36, 1955: 226-262 (fluct. in Baltic); Weeks, Atl. Salm. J., z, Jan. 1955: 3-45 3, Apr.
1955: 33 (record wets.); Hasler, Quart. Rev. Biol., 32, 1956, 200-209 (perception of pathways by migr.
fish.); Blair, J. Fish. Res. Bd. Canada, 13, 1956: 219-232 (tag., Newfoundld.); Hurley, Effects of
Aerial Spraying ... on Food, Young Salmon (Sa/mo salar L.), New Brunswick Streams, Ph. D. Thesis,
Univ. Toronto, 1956; Kerswill, Atl. Salm. J., 1, Feb. 1956: 23-24 (1955 scarcity); Anon., Atl. Salm.
J., I, Feb. 1957: 11 (artif. propag., Sweden); “Avondhu,” Salm. Trout Mag., 150, 1957: 156 (large
Irish salmon); Blair, J. Fish. Res. Bd. Canad., 74, 1957: 135-140 (tag., Labrador); Elson, Canad.
Fish. Cult., 27, 1957: 1-6 (size in smoltification); Canad. Fish. Cult., 27, 1957: 7-17 (use of hatchery-
reared young); Canad. Fish. Cult., 2z, 1957: 19-23 (no. needed to maintain stocks); Canad. Fish.
Cult., 27, 1957: 25-32 (role of hatchery); Ide, Trans. Amer. Fish. Soc., 86, 1957: 208-219 (effect
of DDT on aquatic insects of streams); Hansen, Almind. Beret. Ved. Gronl. Admin. (1956), 1957:
57 (managemt., Greenland); Lindroth, Rep. Inst. Freshw. Res., Drottn., 38, 1957: 109-130 (fluct.
in Baltic); Menzies and Shearer, Nature, Lond., 779, 1957: 790 (salmon tagged in Scotland re-
cov. in Greenland); Rounsefell, Fish Bull. U.S. Fish Wildl. Serv., 57, 1957: 45 pp. (var. in fecund.
with pop., size, age); Svardson, Rep. Inst. Freshw. Res., Drottn., 38, 1957: 357-384 (climate);
Toner, Nature, Lond., 780, 1957: 190 (Irish salmon); Went, Nature, Lond., 180, 1957: 190 (change
in char. of Shannon salmon, Ireland); White, Bull. Fish. Res. Bd. Canad., 116, 1957: 63 pp., 8 figs.
(food, nat. hist. mergansers on salmon waters, Maritime Provinces); Backus, Bull. Amer. Mus. nat. Hist.,
IT3, 1958: 287 (Labrador); Burridge, Trade News, Canad. Dept. Fish., ro (12), 1958: 19-23 (invest.
managemt.); Everhart, Atl. Salm. J., 1, Mar. 1958: 19 (managemt., Maine rivers); Hasler, Horrall,
Wisby, and Braemar, Limnol. Oceanogr., 3 (4), 1958: 353 (sun orient. and homing); Katteberg,
Rep. Inst. Freshw. Res., Drottn., 39, 1958: 55 (compet. of juv. salmon and trout, Sweden); Kers-
will, Atl. Salm. J., 3, Sept. 1958: 28 (research program); Kerswill, Elson, and Keenleyside, Trade News,
Canad. Dept. Fish., To (12), 1958: 4-18 (research program); Murray, Prog. Rep. Fish. Res. Bd. Canad.,
Atl. St., 68, 1958: 20-27 (biol., Newfoundld.); Prog. Rep. Fish. Res. Bd. Canad., Atl. St., 70, 1958:
16-21 (survival, Newfoundld.); Power, Arctic, IZ, 1958: 63-64 (Ungava Bay salmon); Arctic, rz,
1958: 86-92 (evol., freshw. salmon); Rounsefell, Fish. Bull. U.S. Fish Wildl. Sery., 131, 1958 (anad-
romy); Vladykov, Atl. Salm. J., 4, Nov. 1958: 35 (Quebec primer on salmon); Wulff, Atlant. Salmon,
1958: 1-222 (life hist., angling, Canada, Newfoundld., photos., salmon rivers descr., Maine to Labra-
dor); Jones, The Salmon, 1959: 1-192, 27 photos., 24 diagr. (genl.); Keenleyside, Canad. Fish. Cult.,
24, 1959: 17-22 (effect of spray.); Margolis, Canad. J. Zool., 36, 1959: 889 (taxon. review shows
salmonis only Lepeophtheirus from salmonids in N. Atlant. and N. Pacif.); Weeks, Atl. Salm. J., z, March
1959: 3-8 (record salmon); Kerswill, Resources for Tomorrow Conf., background papers, 2, 1961
(review of salmon research).
nobilis Olafsen, Island Reise, 1, 1787: 65 (Iceland); Pallas, Zool. Rus. Asiat., 3, 1814: 342 (diagn.,
refs., distr.; for date of Publ., see Sherborn, Ibis, [13] 4, 1934).
samulus Turton, Brit. Fauna, 1806: 104 (= parr); Parnell, Mem. Werner. Soc. Edinb., 7, 1831-1837:
298, pls. 30, 32, fig. 1, also sep. 1838 (parr distinct from salmon); Jenyns, Manual Brit. Vert. Anim.,
1835: 426 (descr. parr, sep. spp., abund., Britain); Fries, Handl. K. Vet-Akad. Stockh., 1837: 3,
pl. 1 (young; not seen); Thompson, Nat. Hist. Ireland, 4, 1856: 143 (= young 8. sa/ar).
hamatus Cuvier, Régne Anim., ed. 2, 5, 1829: 254 (old male of 8. sa/ar; not pl. 102, fig. 1, which is
of 8. schiefemulleri, or La Truite de Mer); Cuvier and Valenciennes, Hist. Nat. Poiss., 27, 1848: 212,
pl. 615 (sex, maturing, male §. sa/ar with hooked lower jaw); Heckel and Kner, Siisswasserfische, 1858:
276 (old male).
salmo Cuvier and Valenciennes, Hist. Nat. Poiss., 27, 1848: 169, pl. 614 (distr., French rivers).
immaculatus Storer, J. Boston Soc. nat. Hist., 6, 1850: 364 (young 8. sa/ar, type in MCZ; Red Bay,
n. shore Strait of Belle Isle).
32
498 Memoir Sears Foundation for Marine Research
Salmo sebago Girard, Proc. Acad. nat. Sci. Philad., 6, 1854: 380 (Sebago Lake, Maine); Jordan and Ever-
mann, Amer. Food Game Fish., 1902: 168-170, 1 pl. (orig. distr., size, habits); Kendall, Occ. Pap.
Boston Soc. nat. Hist., 7 (8), 1908: 44 (distr., New England); Mem. Boston Soc. nat. Hist., 9 (1),
1935: 104-157, pls. 5-11 (cf. lake and sea salmon, color. ills., distr., habits, size, New England;
S. ouananiche consid. ident. to 8. sebago).
Salmo gloveri Girard, Proc. Acad. nat. Sci. Philad., 7, 1856: 85 (in trib. of Union R., e. Maine); Norris,
Amer. Angler’s Book, 1864: 248 (angling); Gilpin, Trans. N.S. Inst. Sci., Z (4), 1866: 86 (lakes, around
Halifax); Adams, Field Forest Rambles, 1873: 214, 3 figs. (descr., N. Brunswick); Jones, Trans. N.S.
Inst. Sci., 5, 1879: 93 (distr.).
Trutta salar von Siebold, Amtl. Ber. 35, Vers. Dtsch. Naturf. (1860), 1861: 74—77?° (relations discus.); Siiss-
wasserfische Mittel Europa, 1863: 292 (genl., good ills. of vomerine teeth); Malmgren, Finlands Fisk-
fauna, 1863: 58-60 (refs., Finland, name J’. re/icta proposed for salmon of Lake Ladoga; Sweden;
also German transl. by C. F. von Frisch); Steindachner, S.B. Akad. Wiss. Berl., 54 (7), 1866: 19; Brito-
Capello, Cat. Peix. Portugal, Mem. Acad. Sci., Lisboa, 1880: 35 (n. Portugal).
Salmo gracilis Couch, Rep. roy. Cornwall Polytec. Soc., 1859: 1-29 (genl.); Fish. Brit. Isles, 4, 1867: 216,
pl. 216, fig. 1 (genl.).
Trutta relicta Malmgren, Finlands Fiskfauna, 1863: 59 (name proposed for salmon of Lake Ladoga).
Salmo argenteus Ginther, Cat. Fish. Brit. Mus., 6, 1866: 86 (kelt).
Salmo hardinii Ginther, Cat. Fish. Brit. Mus., 6, 1866: 107 (Lake Vener, Sweden).
Salmo ouananiche McCarthy, Forest and Stream, 42 (10), 1894: 206 (Lake St. John [Quebec], trib., and Sague-
nay R.; name commonly given to lake salmon of n. Quebec, Labrador, Newfoundld.); Jordan and
Evermann, Amer. Food Game Fish., 1902: 170-174 (genl.).
Salmon, Pennant, Brit. Zool., 3, 1776: 284, pl. 58 (Greenland, Newfoundld., n. N. Amer.); Arctic Zool., 2,
1792: 392 (n. Amer., none beyond New York); DeWitt Clinton, Trans. Lit. philos. Soc. N.Y., 1,
1815: 147 (Lake Ontario, trib. rivers); Russell, The Salmon, 1864: 234 (genl.); Couch, Fish. Brit. Isles,
4, 1865: 163, pl. 211 (genl., Gt. Brit.); Prince, 31st Ann. Rep. Canad. Dep. Mar. Fish. (1898), Ap-
pend., 1899: lxxi—Ixxi (early abund. Lake Ontario, later decrease; artif. propag. L. Ontario, n. shore
Gulf of St. Lawrence); Comeau, Life and Sport ... Lower St. Lawrence R. and Gulf, 1909: 175-190
(movements, food, salt water, and rivers), 359-367 (local salmon rivers), 308-316 (recs., Gulf of
St. Lawrence rivers); Griswold, Fish Facts and Fancies, 1926: 29-77 (genl.); Huntsman, J. Fish. Res.
Bd. Canad., 5, 1942: 485—-so1; 6, 1946: 476-479 (death from high temp.).
Salmo trutta Linnaeus 1758
Brown Trout (North America); the Trout, Lochleven Trout (British Isles);
European Trout (Continental Europe)
Figure 3
The Brown Trout, introduced from Europe to North America, is similar in general
appearance to the Atlantic Salmon (8S. sa/ar) but is distinguished from it by the characters
given on pp. 459-461. It is easily separable from the brook trout (Sa/velinus fontinalis)
and from the Arctic charr (Sa/velinus alpinus) by its much larger scales, its black-
spotted sides, and the presence of one zig-zag row (or two alternating rows) of sharp,
well-defined teeth (easily felt) along the shaft of the vomer in the midline of the roof
of the mouth. As in other salmonids, the Brown Trout in the sea is silvery in color.
In Europe, where this species is indigenous, anadromous populations were formerly
20. The reference in Neaves Nomenclator to Geoffrey, 1764, is erroneous, based on a mistake of Sherborn’s. The
work was in fact by Garsault, and is not binomial; this information is contributed by Dr. E.Trewavas of the
British Museum (Natural History), who is preparing a note about it for Bull. Zool. Nomencl.
Fishes of the Western North Atlantic 499
considered specifically distinct from the strictly freshwater type, but no specific difference
is now recognized.
The Brown Trout is believed to have been first brought to North America in 1883.
In that year eggs were sent from Germany to New York. The next year a shipment
was sent from England. In 1884, the Lochleven trout, now considered to be the same
as the trout in other parts of Great Britain and continental Europe, was introduced in
Newfoundland. Another variant known as the German Brown Trout was brought to
Newfoundland in 1892.
Anadromous populations are well established on the Avalon Peninsula in southern
Newfoundland, where this species has spread through salt water to many streams,
some as far as 60 miles from the four original centers of introduction. H. W.
Walters, Director of the Wildlife Division of the Newfoundland Department of Mines
and Resources, believes that the native brook trout (S. fontina/is) is declining in numbers
in streams resorted to by the Brown Trout and that the latter will continue to spread to
still more rivers around Newfoundland. The anadromous Brown Trout is also reported
for the Guysborough River watershed, Nova Scotia, where it is taken chiefly in the
10-12 mile long estuary (information from James Cott), and a few are taken in some
streams and rivers of Maine, Massachusetts, Connecticut, New York, and New Jersey.
Salmo gairdneri Richardson 1836
Rainbow Trout, Steelhead Trout
This species has been transplanted artificially from western to eastern North
America. In the west there are both anadromous and freshwater populations. Some of
the latter have diverged far enough from the parent stock to be regarded by some as
specifically distinct while the taxonomic status of others is still in doubt. The only
evidence that the Rainbow resorts to salt water in eastern North America is based on its
presumed natural extension from Crooked Creek, New Brunswick, to a few neighboring
streams, apparently by way of the Shepody River estuary into which Crooked Creek
flows. The sea-going individual is silvery when in salt water, but upon returning to
fresh water it becomes greenish with a pinkish or reddish band along the side, hence
the name “Rainbow.”
ay So Hoes
23
24.
TEXT AND FOOTNOTE REFERENCES
Auten, J. Anim. Ecol., Zo, 1941.
Alm, Rapp. Cons. int. Explor. Mer, 92, 1934.
Alm, Landtbr. Styr. Medd., Stockh., 22, 1943.
Anonymous, Annu. Rep. Fish. Res. Bd. Canada,
1955:
Anonymous, Annu. Rep. Fish. Res. Bd. Canada
(1956-57), 1957-
Anonymous, Annu. Rep. Fish. Res. Bd. Canada
(1957-58), 1958.
Atkins, Rep. U.S. Comm. Fish. (1872-73), 2,
1874.
Austin, Trans. Lit. Hist. Soc. Quebec, N.S. 4,
1866.
Sato Trans. Amer. Fish. Soc., 62, 1932.
Belding, Trans. Amer. Fish. Soc., 64, 1934.
Belding, Trans. Amer. Fish. Soc., 66, 1937.
Belding, Trans. Amer. Fish. Soc., 67, 1938.
Belding, Publ. Amer. Ass. Adv. Sci., 8, 1939.
Belding, Trans. Amer. Fish. Soc., 69, 1940.
Belding and Kitson, Trans. Amer. Fish. Soc.,
64, 1934.
Belding, Pender, and Rodd, Trans. Amer. Fish.
Soc., 62, 1932.
Bigelow and Schroeder, Bull. U.S. Bur. Fish.,
40 (2), 1927.
Bigelow and Schroeder, Fish. Bull. (74) U.S.
Fish. Wildl. Serv., 53, 1953.
Blair, J. biol. Bd. Canada, r, 1935.
Blair, Res. Bull. Nfdld. Dept. Nat. Resources,
13, 1943.
Blair, J. Fish. Res. Bd. Canada, 13, 1956.
Blond, Great Migrations (transl. Frances
Frenaye). McMillan, New York, 1956.
Brainard, Atl. Salmon J., 1953 (March).
Brito-Capello, Mem. Acad. Sci., Lisboa, 1880.
500
25.
26.
45.
Bull, H. O., Rep. Dove Mar. Lab. (1930),
N.S., 29, 1931.
Burridge, Trade News, Canad. Dept. Fish.,
To (12), 1958.
Cinercon! Salmon Rivers and Lochs of
Scotland, ed. 2, London, 1g2r.
Calderwood, Proc. roy. Soc. Edinb., 45, 1926.
Calderwood, Proc. roy. Soc. Edinb., 47, 1928.
Calderwood, Salmon and Sea Trout, 1930.
Cameron, Parasites and Parasitism, 1956.
Carlin, Rep. Inst. Freshw. Res. Drottn., 36,
1955:
Carpenter, Nature, London, 143, 1939.
Comeau, Life and Sport North Shore Lower
St. Lawrence and Gulf, 1go9.
ten Age and Growth, Salmon and Trout
in Norway, IgIt.
Dahl, Salmon and Trout, Handbk., 1914.
Dahl, Salmon Trout Mag., 86, 1937.
Day, British and Irish Salmonidae, 1887.
Dunbar and Hildebrand, J. Fish. Res. Bd.
Canada, g (2), 1952.
Dymond, Trans. roy. Soc. Canada, 41 (v),
1947-
Excuersaum, Rapp. Cons. int. Explor. Mer,
2I, 1916.
Elson, Progr. Rep. Fish. Res. Bd. Canada, Atl.
Cst. St., 63, 1955.
Elson, Canad. Fish. Cult., 27, 1957.
Elson and Kerswill, Trans. 20th N. A. Wildl.
Conf., 1955.
Everhart, Atl. Salmon J., 1958 (March).
46.
47-
48.
49-
50.
75:
Fishes of the Western North Atlantic
Fiaye; Sinensia, 4 (9), 1934.
Fisher, Atl. Salmon J., 3, 1955 (Sept.).
Fowler, Rep. N. J. State Mus., 1905.
Frisch, von, Study of Orientation of Honey
Bee (Eng. transl.), Bull. Anim. Behav., 9,
IQSt.
(Greve in Griswold and Hume, Life Hist.
Canadian Salmon. Privately printed, Nor-
wood, Mass., 1930.
Gurney, Brit. Freshw. Copepoda, 23, 1933-
iB lam, Almind. Beretn. vedror. Gronl.
Admin. (1956), 1957.
Hansen and Hermann, Danm. Fisk. Havy-
unders., I5, 1953.
Hasler, Quart. Rev. Biol., 3z, 1956.
Hasler, Horrall, Wisby, and Braemar, Limnol.
Oceanogr., 3 (4), 1958.
Hayes, Bull. Fish. Res. Bd. Canada, 99, 1953.
Hewitt, Salmon Trout Mag., 132, 1951.
Huntsman, Bull. biol. Bd. Canada, 27, 1931.
Huntsman, Progr. Rep. Fish. Res. Bd. Canada,
Atl. Cst. St., 6, note 25, 1933.
Huntsman, Trans. roy. Soc. Canada, 31 (v),
1937-
Huntsman, Science, 85, 1937.
Huntsman, J. Fish. Res. Bd. Canada, 4, 1938.
Huntsman, Bull. Fish. Res. Bd. Canada, 57,
1939-
Huntsman, Publ. Amer. Ass. Adv. Sci., 8, 1939
Huntsman, J. Fish. Res. Bd. Canada, 5, 1942.
Huntsman, Science, 95, 1942.
Huntsman, J. Fish. Res. Bd. Canada, 6, 1945.
Huntsman, Science, 105, 1947.
Huntsman, Salmon Trout Mag., 123, 1948
(May).
Huntsman, Trans. Amer. Fish. Soc., 75, 1948.
Huntsman, Salmon Trout Mag., 130, 1950
(Sept.).
Huntsman, J. Cycl. Res., Z, 1951.
Huntsman, Salmon Trout Mag., 736, 1952
(Sept.).
Huntsman and Hoar, J. Fish. Res. Bd. Canada,
4, 1939-
es Trans. Amer. Fish. Soc., 86, 1957.
76
100.
Tol.
102.
103.
. Kerswill, Resources for
501
: eeu K. Dansk. Vidensk. Selsk. Skr., Biol.
Medd., r4, 1939.
. Jones, Proc. zool. Soc. Lond., 177, 1947.
. Jones, The Salmon, 1959.
. Jones and King, Proc. zool. Soc. Lond., r7g,
1949.
. Jones and King, Proc. zool. Soc. Lond., 120,
1950; or Salmon Trout Mag., 728, 1950.
. Jones and King, Proc. zool. Soc. Lond., 122,
1952.
. Jones and Orton, Proc. roy. Soc. Lond., B 128,
1940.
. Jordan, Guide to Study Fishes, 2, 1905.
. Jordan and Evermann, Amer. Food Game
Fishes, New York, 1go02.
: | Grama. Rep. Inst. Freshw. Res. Drottn.,
39, 1958.
. Keenleyside, Canad. Fish. Cult., 24, 1959.
- Kendall, Mem. Boston Soc. nat. Hist., 9 (1),
1935:
- Kerswill, Atl. Salmon J., 1955 (Jan.).
- Kerswill, Atl. Salmon J., 1956 (Feb.).
. Kerswill, Atl. Salmon J., 1958 (Sept.).
Tomorrow Conf.
background papers, 2, 1961.
. Knight, Fishes New Brunswick, 1863.
5 eke, Rep. Inst. Freshw. Res., Drottn.,
36, 1955.
. Lindroth, Rep. Inst. Freshw. Res., Drottn.,
38, 1957.
. Lindsay and Thompson, Rep. Nfdld. Fish. Res.
Comm., I, 1932.
. Lobo, Mem. Acad. Real. Cien. Lisboa, 1879.
: IMererre Bird Migration, Cambr. Monogr.
exp. Biol., 3, 1955.
. McCrimmon, J. Fish. Res. Bd. Canada, rz,
1954-
. McFarland, Salmon of the Atlantic, 1925.
Menzies, The Salmon, 1925.
Menzies, Proc. roy. Soc. Edinb., 45, 1926.
Menzies, Publ. Amer. Ass. Adv. Sci., 8, 1939.
Menzies, The Stock of Salmon. Arnold, Lon-
don, 1949.
502
To4.
T05.
I06.
107.
Ios.
I09.
IIo.
Tit.
Tae)
TIF:
IT4.
THEE
rr6.
TG
Irs.
TI9Q.
120.
TATE
122.
123)
Memoir Sears Foundation for Marine Research
Menzies and Macfarlane, Proc. roy. Soc.
Edinb., 47, 1928.
Menzies and Shearer, Nature, London, 179,
1957-
Murray, Progr. Rep. Fish. Res. Bd. Canada,
Atl. Cst. St., 68, 1958.
Murray, Progr. Rep. Fish. Res. Bd. Canada,
Atl. Cst. St., 70, 1958.
Nerrzz, Salmon Fish. St. Lawrence, 1857.
Nobre, Biol. Minist. Agric. Lisboa, ano 13,
No. 2 (1 ser.), 1932.
Nobre, Fauna Marin. Portugal, z, Vert.,
nO Sir
Oircran: Jones, and King, Proc. roy. Soc.
Lond., B 725, 1938.
Pee Rep. Fisher. Gulf of St. Lawrence,
ed. 1, 1849.
Perley, Rep. Sea River Fisher. New Brunsw.,
ed. 2, 1852.
Phelps and Belding, Statist. Study Recs.
Salmon Fish. Restigouche River, 1931.
Power, Arctic, IZ, 1958.
Pyefinch, Freshw. Salmon Fish. Res., Scott.
Home Dept., 9, 1955.
Roonserex, Fish. Bull. U.S. Fish Wildl.
Serv., 131, 1958.
Rushton, Salmon Trout Mag., 44, 1926.
SaaS Quart. Rey. Biol., 74, 1939.
Scott, British Parasit. Copepoda, I, 1913.
Sedgwick, Salmon Trout Mag., 137, 1953.
Sedgwick, Salmon Trout Mag., 143, 1955.
Steindachner, S. B. Akad. Wiss. Berl., 54 (7),
1866.
124.
125.
T26.
127.
128.
129.
130.
T3I.
132.
ine yey
134.
135.
136.
137.
738.
139.
140.
It.
T42.
T43.
T44.
T45.
146.
147.
r48.
Svardson, Rep. Inst. Freshw. Res., Drottn.,
36, 1955.
Swain, Nature, London, 180, 1957.
Teche Trans. zool. Soc. Lond., 24,
1938.
Tchernavin, Proc. zool. Soc. Lond., B 173,
1943.
Toner, Nature, London, 780, 1957.
Twomey, Nature, London, 180, 1957.
\ see Ann. St. Centr. Hydrobiol. appl.,
3, 1950.
Vibert, Trans. Amer. Fish. Soc., 83, 1954.
Vladykov, Fishes of Quebec, Atl. Salmon,
Dept. Fish., Quebec (undated).
Vladykov and McKenzie, Proc. N. S. Inst.
Sci., I9, 1936.
Vladykov and McKenzie, Atl. Salmon J.,
1955 (Jan.).
\W ape Atl. Salmon J., Z, 1955.
Went, J. Anim. Ecol., 15, 1946.
Went, J. Cons. int. Explor. Mer, 16, 1950.
Went, Nature, London, 180, 1957.
White, J. biol. Bd. Canada, 2, 1936.
White, Bull. Fish. Res. Bd. Canada, 58,
1939-
White, J. Fish. Res. Bd. Canada, 6, 1942.
White, Bull. Fish. Res. Bd. Canada, 97,
1953.
White, Bull. Fish. Res. Bd. Canada, r7r6,
1957-
White and Huntsman, J. Fish. Res. Bd.
Canada, 4, 1938.
Wilder, Canad. J. Res., D 25, 1947.
Willughby, Hist. Pisc., 1686.
Wilson, Proc. U.S. nat. Mus., 28, 1905.
Wulff, The Atlantic Salmon, 1958.
Genus Sa/velinus Richardson 1836
By
Henry B. BicELow
Museum of Comparative Zoology
Harvard University
Salvelinus Richardson, Fauna Boreal.-Amer., 3, 1836: 169; first use in the singular (by implication) of the
group name Salmones salvelines of Nilsson, Prod. Ichthyol. Scand., 1832: 7; type species, Sa/mo salvelinus
Linnaeus, Syst. Nat., ed. 10, Z, 1758: 309, Austria, designated by Jordan, Genera Fish., 2, 1919: 186.
Generic Synonyms:
Salmo (in part) Linnaeus, Syst. Nat., ed. 10, 7, 1758, for 8. 2/pinus Linnaeus 1758: 309, Lapland, Alpine lakes
in England; 8. sa/ve/inus Linnaeus, 1758: 309, Austria; 8. sa/marinus Linnaeus, 1758: 310, “Habitat
tridente in fluviis frigidis’; and 8. umé/a Linnaeus, 1758: 310, Switzerland.}
Baione, DeKay, Zool. N. Y., Fishes, 4, 1842: 244; type species, B. fontinalis DeKay 1842; Rockland County,
New York.
Umébla, von Rapp, Jheft. Ver. Vaterl. Naturk. Wiirttemberg (1854), To, 1856: 171; type species, Sa/mo umbla
Linnaeus, Syst. Nat., ed. 10, Z, 1758: 310; Lakes of Switzerland and Italy.
Not Cristivomer Gill and Jordan iz Jordan, Manual Vert. north. U.S., ed. 2, 1878: 356, 359; type species,
Salmo namaycush Walbaum, P. Artedi Genera Pisc., Ichthyol., Emend., 3, 1792: 68, by ref. to “the
Namaycush,” Pennant, Arctic Zool., r, Introd., 1784; cxci; lakes far inland from Hudson Bay.
Characters. Essentially as in the genus Sa/mo, but with: Scares very small,
scarcely visible—at least 195 rows and commonly 200 or more (120-130 in Salmo);
TerrtH lacking on shaft of vomerine bone along midline of roof of mouth (one or two
more or less complete rows in the young of all species of Sa/mo, persisting throughout
life in some); the teeth on premaxillaries usually forming a continuous series around
front of mouth (for exception, see Fig. 127). Caupat fin only moderately concave.
Bone above vomer (EtHMorp of Kendall, Mrserumorp of Regan, Prorrumorp of Starks)
more than 38 °/, as wide as long. FRONTAL AREA OF SKULL not flat, usually with a median
longitudinal ridge. Not more than 50 Pytoric cazca.
Sexual Dimorphism. Normally, in the mature male the head is noticeably larger
than in the female, as is the mouth, the teeth are somewhat larger, and the pectoral
and pelvic fins are somewhat longer. In the male at spawning time the lower jaw be-
comes slightly hooked, as in Fig. 124 (less so than in the salmon), a thick mucous
layer develops, nearly or quite concealing the scales, and the colors intensify, especially
the reddish hues, with the white margins of the pectoral and pelvic fins standing out
more conspicuously. These seasonal characters disappear soon after spawning has been
completed (45: 85; 62: 7; 9: 8).
Remarks. The restriction of the genus Sa/velinus to species having a nearly truncated
1. Salmo lacustris Linnaeus, 1758: 309, appears to have included more than one species. For an early discussion, see
Heckel (31: 353, pl. 7, figs. 4, 5)-
$93
504. Memoir Sears Foundation for Marine Research
caudal fin (at most a moderately concave one), a relatively broad bone in the ethmoid
position, a more or less rounded frontal region in the skull, not more than 50 gastric
caeca, and various skull characters as well as differences in the caudal skeleton,? ex-
cludes zamaycush Walbaum (North American lake trout), for which Gill and Jordan
founded the genus Cristivomer. Viadykov has revived Baione DeKay as a subgenus of
Salvelinus for the species fontinalis (67:
928), which, according to him, is charac-
terized by a “‘little forked tail, black stripe
on lower fins, the absence of hyoid teeth,
and an uninterrupted row of vomerine
and palatine teeth.’ But Walters has
found that in some specimens of the com-
panion species a/pinus the vomerine and
palatine teeth also form an uninterrupted
row (70: 258, 260, fig. 14). Neither do
small differences in color seem to me to
; provide a sufficient basis for subgeneric
Roun 24 Sling bi Hed fron malt separation in a genus where local popu
land. After Jensen. lations vary as widely in this respect as
they do in Sa/velinus; nor does the pres-
ence or absence of hyoid teeth serve any practical purpose in this case, whatever the
phylogenic implication of differences in this respect.
Species. The species remaining in Sa/velinus after Cristivomer namaycush is excluded
fall in two divisions, the one typified by what may be termed the “‘fontinalis complex,”
the common brook trouts of North America (p. 525), the other by the “‘a/pinus com-
plex,” the Arctic-subarctic charrs (p. 507). Regan characterized these two divisions as
follows (55: 408):
S. alpinus division: head of the vomer with posterior process little developed. No
dark spots or markings. Circumpolar.
S. fontinalis division: head of the vomer with posterior process well developed.
Blackish or dark olivaceous markings on back and on dorsal and caudal fins. North
America.
The shape of the head of the vomer, however, is not a convenient aid for identi-
fication, for careful examination of the lower surface of the anterior part of the
skull is required. But the color pattern does afford a reliable field mark, not only for
freshwater populations of a/pinus and fontinalis but even for those taken in brackish
or salt water; specimens of S. fontinalis always retain some trace of the wavy dark
markings on the dorsal and upper part of caudal, even though similar marks may dis-
appear entirely from the back and sides when the fish leave fresh water, or shortly
before. Further, the a/pinus division, compared to the fontina/is division, has the smaller
2. For further details, including the history of the case, see Kendall (46: 78-81), and especially Vladykov
(67: 932).
Fishes of the Western North Atlantic 505
mouth and the less blunt snout; although the maximum numbers of dorsal and anal
rays and of gill rakers average more in a/pinus than in fontinalis,® the overlap here is
too wide for these numerical differences to serve as a reliable basis for separation.
Neither is the shape of the caudal fin a safe clue to identity, for while the concavity
is somewhat less deep on the average among /ovtinalist than among sea-run a/pinus, it
is deeper in some freshwater trout from Labrador (seen by me) than in some a/pinus
from eastern Greenland.
Both the fontinalis complex and the a/pinus complex include populations that
spend their entire life in fresh water, as well as populations that spawn in fresh water
but spend a part of the year in brackish or salt water, just as salmon do. It is with
these migratory populations alone that we are concerned here.
Both complexes tend to vary widely in color and body proportions and in the
sizes to which they grow in different localities, as is reflected in the names under
which they have appeared in scientific literature (see Synonyms, pp. 523, 541). In
the case of fontinalis, these local varieties seem so clearly to be a result of en-
vironment that only one of the forms in question, agassizii Garman 1885, has been
recognized as a separate species, by Jordan, et a/. (43: 60), but doubtfully so in my
opinion.
“The various forms of charr allied to Sa/velinus alpinus (Linnaeus) present a...
confused picture to the systematist, for a myriad of forms, described and undescribed,
exist” (Walters, 72: 274); views have differed widely as to how many of these
deserve recognition as distinct species or as subspecies. On the one hand, six separate
species were listed for North America by Jordan, et a/. (43: 60-61), 15 for the British
Isles by Regan (54: 82-85), and 16 for the Soviet Union and adjacent countries by
Berg (in 71: 274). On the other hand, Goode (26: 501), following von Siebold’s lead
(59: 285-288) and finding it “difficult to believe that every little lake or group of lakes
in Europe possesses a well characterized species of fish,” tentatively regarded the charrs
of Europe as representing a single highly variable species. This viewpoint is further
exemplified by Smitt’s union of all charrs of Scandinavia, from both fresh and salt
water, in the single species, umb/a Linnaeus, with a/pinus Linnaeus and sa/velinus
Linnaeus rated as synonyms (6I: 841).
In view of this diversity of opinion as to the systematic status of the freshwater
charrs of the a/pinus complex from different parts of the world, it is fortunate for pres-
ent purposes that sea-going populations of the western North Atlantic appear clearly
to represent a single species that seems to be identical to the common sea-run charr
of Greenland. This was described by Fabricius in 1780 as Salmo carpio (22: 170);
but this name had been previously applied by Linnaeus to one of the European
freshwater charrs (1758: 309). To replace carpio for our fish, Kendall (44: 507),
3- S. fontinalis: dorsal rays 9-14, anal rays 8-12, gill rakers on first arch 4-8+ 7-9; alpinus: dorsal rays 9-15, anal rays
8-14, gill rakers usually 6-8+ 12-16, with a total of 13-22 reported. For meristic counts for sea-run trout of the
Moser River, Nova Scotia, compared with several populations of freshwater fontinalis, see Wilder (78: 186, tab.8);
for fontinalis compared with alpinus, see Vladykov (67: 909-912).
4. The anglers’ name, Square-tail Trout, is based on the contrast between fontinalis and Cristivomer namaycush (p. 453).
506 Memoir Sears Foundation for Marine Research
following Regan (55: 408), proposed a/pinus Fabricius 1780 (22: 173) on the assump-
tion that Fabricius’ a/pinus is not separable specifically from his carpio, an assumption
that seems fully justified, so closely do Fabricius’ accounts of the two agree. The name
alpinus has accordingly been used for the sea-run Arctic charr by a majority of the
authors who have studied it within the last few years. In this connection, see especially
Vladykov (67: 929) and Walters (77: 274).
The name stagnalis Fabricius 1780 (p. 175) has sometimes been applied to the
sea-run Arctic charr (for instances, see References, pp. §23, 524), but it is not an ap-
propriate choice, for as Kendall has emphasized (44: 507), Fabricius gave this name to a
large Greenland trout (17.3 in.), of remote mountain waters, that never descends to the
sea. If Fabricius was correct in using for his Greenland charr the name a/pinus of Linnaeus
(1758: 510), it follows further that the charr with which we are dealing here is not
separable specifically from the Arctic charr of Lapland or perhaps from the charr
(or one of the charrs) of England. Indeed, a specimen about 215 mm TL from Novaya
Zemlya does not differ from my smaller specimens from Greenland and Labrador in
any evident respect that might be regarded as specific, unless perhaps it differs in a more
slender trunk and in a wholly plain coloration with blue-gray back, bright silvery sides,
and white lower surface. Present indications are that the red-spotted charr of the
alpinus group, named malma by Walbaum in 1792 (69: 66) and which is sea-run
along both coasts of the northern North Pacific, deserves recognition as at least a
subspecies of a/pinus (Dymond, 17: 39), if not as a separate species, as it is classed
by Delacy and Morton (12: 81-90).
The freshwater populations of northeastern continental America have been given
separate specific names, 1. e. oguassa Girard 1854 for those of the Rangeley Lakes
region in northern Maine, aureo/us Bean 1887 for those of the cold lakes of northern
New England in general, and marstoni Garman 1893 for those known in similar situa-
tions in northern Quebec. It remains an open question, however, whether any of these
named forms, based chiefly on color and size, differ consistently enough from the typical
sea-running a/pinus to call for specific or subspecific recognition in zoological nomen-
clature; this applies equally to the names that have been proposed for the freshwater
populations of Arctic Canada and northward; for a list of these, see p. 524.
Key to Species of Sa/velinus, Taken in Brackish or Salt Water,
in the Western North Atlantic
1a. Dorsal fin and upper corners of caudal with dark wavy or vermiculated markings;
color pattern on back, if visible, in the form of darker and paler marblings; length
of head from tip of snout to rear margin of gill cover about 3.8—4.4 in SL; anterior
contour of head bluntly rounded; pectoral fin broadly rounded (Fig. 128).
fontinalis (Mitchill) 1815, p. 525.
1b. Dorsal and caudal fins with no dark markings; back without darker and paler
marblings; length of head from snout to rear edge of gill cover 4.6—5.3 in SL,
Fishes of the Western North Atlantic 507
but 4.5 in larger males where heads are relatively larger; anterior contour of head
more narrowly tapered; pectoral fin pointed (Figs. 125, 126).
alpinus (Linnaeus) 1758, below.
Salvelinus alpinus (Linnaeus) 1758
Arctic Charr
Figures 3, 118, 119, 124-127
Study Material. 1. Sea-run populations. More than 31 specimens: 15, 78—ca.
600 mm SL, from eastern Greenland, Thule in northwestern Greenland, and Godthaab
in western Greenland; 3, from the Fraser River near Nain, Labrador (ca. 56°37'N);
2, from Novaya Zemlya, and others from the northwestern coast of North America and
from Sakhalin Island, MCZ; 11, 305-ca. 600 mm SL, from Baffin Island, Ungava
Bay (Turner Coll.), and Island Harbor near Mokovik, Labrador, USNM.
II. Freshwater populations. Numerous specimens of the a/pinus complex from
various freshwater lakes in northern New England, northern Quebec (Lac de Marbré,
and a lake near Lake St. John), Norway, Switzerland, and Austria.
Distinctive Characters. The only saltwater fishes for which Sa/velinus alpinus might
be mistaken in eastern North American waters or around Greenland are Salmo salar
(Atlantic salmon) of corresponding sizes, Sa/velinus fontinalis (sea-run brook trout), or
possibly Cristivomer namaycush (lake trout), which have sometimes been known to stray
out into brackish or salt water;° nor is there much danger of confusing a/pinus with any
one of these. It is readily separated from fontinalis by its wholly plain color, or at most
by its faintly spotted dorsal and caudal fins, by the spotted (not vermiculated) color
pattern (if any) on its back, and by its noticeably more pointed head. It is set apart
from Sa/mo salar old enough to have dropped some of the vomerine teeth by its very
much smaller scales and the lack of black markings on its sides; and the toothless
nature of the shaft of its vomer is an additional point of distinction between a/pinus and
a very small S. sa/ar. Neither, I fancy, would anyone at all acquainted with the salmon-
like fishes be likely to mistake a/pinus for C. namaycush, so much more deeply forked is
the caudal fin and so much longer relatively is the head of the latter (cf. Figs. 125, 129).
Description. Based on six sea-run specimens, 360-600 mm SL, from Mokovik
Bay, Fraser River, and Ungava Bay, Labrador; Baffin Island; Godthaab, western Green-
land; and eastern Greenland; in USNM and MCZ; and on other available data.®
Trunk fusiform, its maximum thickness about 11-16 °/, its maximum depth
about 22—26 °/,, of SL, depending on the fatness of the specimens; dorsal profile weakly
and about equally convex both forward from dorsal fin to snout and rearward to adipose
fin. Depth of CaupaL PEDUNCLE between 0.33 and 0.5 of length of head. Sca.es present
5- In northern Labrador, in the northern part of Hudson Bay, and in Bathurst Inlet, Arctic coast of Canada. For a
recent list of such happenings, see Walters (7r: 275).
6. For proportional dimensions and meristic counts for sea-run a/pinus from various localities in Arctic-subarctic Canada
and western Greenland, see Garman (25: 80-81), Dresel (r4: 255-258), Henn (32: 2), Vladykov (67: 909-913), and
Backus (3: 288-293).
Memoir Sears Foundation for Marine Research
508
“(uonsaT[op sou) 6Ez+E WNSN “eg PABBUL] WoIy ‘afeur ‘sHusd7y snuraajog *Qz1 AMMO]
“9fESE WNS ‘purfuccd£y Jo IsvOd ysvo WIZ “opeUTay
‘uu oot ‘snuidjp snuiaayyg “StI aXnoIy
Fishes of the Western North Atlantic 509
on body and tail sectors of trunk; head and fins naked; scales minute, as is charac-
teristic of the genus Sa/velinus; 200-235 reported. Lareral LINE represented as having
120-130 scales perforated.
Heap 22 °/, of SL, noticeably less blunt anteriorly than in fontinalis (Fig. 128).
SNnouT narrowly rounded, its length in front of eye 29-30 °/, of head. Tip of Lower
jaw about even with tip of Upper jaw, or extending slightly beyond it; rear end of
upper jaw reaching to a perpendicular varying in position from about abreast of mid-
point of pupil to slightly behind eye in young specimens and in larger females (Fig.
125), and to behind eye by a distance about equal to length of eye in mature males
(Fig. 126). Rear edge of Eve about 35-43 °/, of distance rearward from snout to rear
margin of gill cover. BRaANCHIOSTEGAL rays IO—I1 reported. GILL RAKERS 13 and 16
on lower limb and 10 and 10 on upper limb on two fish—from western Greenland and
eastern Greenland; 11-18 reported on lower arch and 6-10 on upper limb; the total
number in nine fish from northern Canada 17-27, average 23.2 (67: 910, tab. 4).
TeerH. Premaxillaries, maxillaries, palatines, and dentary portion of lower jaw
each with a single series of sharp, slightly recurved teeth, smaller on the whole and
hence less conspicuous than in /fontinalis; a patch of similar teeth on head of vomer
anteriorly on the midroof of mouth; a single row around the tongue; also a patch of
minute but sharp teeth on the bones at base of tongue (no teeth there in fontinalis),
one such patch farther rearward below, in the pharynegeal region, and two such pat-
ches in pharyngeal region above, one of these along the base of each fourth gill arch.’
Rayed Dorsat fin rhomboid, its corners angular or very slightly blunted, its
base 1.7—2.0 in head, its origin a little less than midway rearward (about 47-48 °/,)
from tip of snout to upper origin of caudal fin, with 10-12 rays® (9-15 reported),
the first 1-3 not branched, the longest ray 0.9-1.1 times as long as base. Dorsat
Avipose fin with shape and relative size as shown in Figs. 125, 126, its origin 56-61 °/,
of distance rearward from rear end of rayed dorsal base to upper origin of caudal.
CaupaL with upper and lower margins very weakly convex, the upper and lower
corners abruptly rounded, the rear contour weakly concave, the breadth of fin, when
widespread, about 1.5 times length of its upper and lower margins. Anat rhomboid,
with angular corners, of the shape shown in Figs. 125, 126, its base about 84-95 °/,
of base of dorsal, its origin about midway between a perpendicular at midpoint of
dorsal and lower origin of caudal, with
II-I2 rays (8-14 reported), the first 2 Table I. Frequency Distribution of Dorsal (17 fish)
and Anal Rays (18 fish) in Sea-run a/pinus from
not branched, the longest ray about 0.88— Arctic’ Canean®
1.2 times longest dorsal ray. Petvics with
2 A is Number of rays....... 9 10 II
midpoint of base about at a perpendicular Dorsal, no. of fish
at midpoint of base of rayed dorsal or a Anal, no. of fish ....... 8 7 3
little anterior thereto, the longest ray about * From Vladykov, 67: tables 8, 9.
=
>
=
ONAN
7. For photographs of the tongue teeth, hyoid teeth, and pharyngeal teeth of the a/pinus species-complex (as represented
by the freshwater S. marstoni and S. aureolus), of S. fontinalis, and of Cristivomer namaycush, see Vladykov (67: 920,
924, figs. 6-11).
8. Precise counting is made difficult by the thickness and fleshy nature of the anterior basal part of the fins.
510 Memoir Sears Foundation for Marine Research
as long as longest ray of rayed dorsal; the fleshy appendage beside base of pelvics a
little longer than eye. Pecrorats noticeably narrower and with more acute tip than
in fontinalis, their shape, when spread, as shown in Figs. 125, 126, their origin a
little anterior to rearmost edge of gill cover (as in fontinalis, Fig. 128), the longest
ray a little longer than longest ray of rayed dorsal.
VERTEBRAE 61-69 _ recorded; see
Table 1 for their relative frequency.
Baie tment coer cs en Pytoric CAECA, average 39.1 reported.
Number of fish.. 6 26 151 213 125 31 6 ee Dimorphism. See p. 503.
Percent /of total’. Me) oe Ma5 3B 22) 6 x Color.2 While the young a/pinus so-
journ in fresh water, their back and the
upper part of their sides are of some shade
of brown or blackish green with about ten well-marked darker crossbars, the so-
called “‘parr marks.”’ These bars are conspicuous until the little fish have grown to a
length of at least 150 mm, and they continue discernible up to 240 mm or 50, 1. e.
until after the fish have moved out into brackish or salt water on their first seaward
migration.
While they are in salt water and until the time arrives for their return to fresh
water, their back and the upper part of their sides have been variously described as
metallic blue, greenish blue, sea green, or steel grey, with metallic reflections; the dark
of the back extends forward onto the head and downward there to the tip of the upper
jaw. The sides as a whole are pale yellowish, tinged with a fleshy color in some, and
marked with a basic pattern of many small spots, yellow or faintly pinkish, and indis-
tinctly outlined; the spots vary widely in size from specimen to specimen—in regularity
of distribution, in color, and especially in intensity—being so faint in some as to be
scarcely, if at all, distinguishable; they are the most numerous above the lateral line
anteriorly, but below the lateral line posteriorly; and they are confined to the upper part
of the sides above a line extending from the base of the pectorals to the origin of the
lower side of the caudal fin. In salt water, the lower part of the sides becomes overlaid
in some specimens with a silver coating so dense that these areas are as silvery as in
Atlantic salmon (pp. 462, 463); but others show hardly a trace of silver; the reasons
for this difference still remain a subject for speculation. The lower surface is white,
sometimes tinged faintly with yellow, without luster. The dorsal fin ordinarily is either
colorless or of a pale shade of the back color, sometimes tinged posteriorly with yellowish
red or in some cases with purplish black (see below). The caudal is either dusky, purplish
black (see below), or colorless, with the outermost rays (below as well as above) yellowish
red; and an occasional fish may show irregular dark spots on both dorsal and caudal.
The pectorals and pelvics are creamy white. The anal is either creamy white or some-
what dusky centrally, tinged with red both anteriorly and posteriorly.
Table II. Number of Vertebrae in 558 Arctic Charr
from the Northeastern Labrador Coast*
* From Andrews and Lear, 2: 854, tab. 8.
g. I have no color notes from life. This color account is based on the Study Material, in comparison with published
descriptions; see especially Richardson (56: pl. 8), Garman (25: 81, fig. 19), Kendall (44: 505), Johansen (40: 671,
pl. 45, figs. 6, 7), Weed (72: 133), Jensen (39: 68, 69), and Backus (3: 289, 290).
Fishes of the Western North Atlantic spi
The following description of the color of a/pinus from the Gander River, New-
foundland, contributed by G. B. Wiggins, is especially instructive, because it was
based on specimens that were still alive when removed from the gill net:
The fish had indistinct flesh-colored spots on the sides only and there were no red or blue spots. The flesh-
coloured spots were almost completely obscured by a purplish-black metallic colour all over the sides. There
were no markings on the back, the colour there being a solid purple-black like the sides. The abdomen was
white with greyish blotches. The dorsal and caudal fins
were solid purple-black and had no markings. The anal, ‘Table III. Percentages of Fish with White, Pink, and
abdominal and pectoral fins had a pink tinge. Theanal_ Req Meat among §97 Arctic Charr from Represent-
and abdominal fins had a distinct white border, while ative Localities, South to North, along the North-
the white border of the pectoral fins was less distinct. eastern Labrador Coast, 1953 *
: Locality White Pink Red No. of Fish
When Arctic Charr re-enter freshen ly. 9 84 7 89
water, the silvery sheen disappears from Nain No. r.. Fi 43 50 31
their sides, their back changes toa neutral Nain No.2.. ° 39,6 117
greenish brown, the color pattern of their amieeach ie e i me
upper parts becomes greatly accentuated, Ramah...... ° 64 36 120
the spots on their sides assume a stronger * From Andrews and Lear, 2: 856, tab. ro.
whitish yellow or orange-red shade, their
ventral fins become more or less suffused with a reddish color, and the color of their
bellies changes to a strong orange yellow. The alteration is much more spectacular for
maturing males than for immature specimens or for maturing females.
In northeastern Greenland, and presumably elsewhere also, the nuptial livery of
Arctic Charr is at its height in November, but the brilliant red-orange hues have en-
tirely faded once more by the time the fish move downstream again during the fol-
lowing May or June. We have yet to learn whether their sides turn silvery again just
before they actually re-enter salt water, or just afterward.
Color of Flesh. The color of the flesh of a/pinus ranges between white and red, with
recorded percentages as given in Table 111.
In western Greenland the commercial catch is sorted similarly into white and red
meat, with the latter fetching the higher price (39: 71). But the mutual percentages at a
given locality may differ considerably from
Table IV. Percentages of Arctic Charr with Pink, year to year, as demonstrated in Table iv.
White, and Red Meat in the Commercial Catches at General experience with other sal-
Okkak Bay, Labrador, during the Seaward Migra-
vempiyy cy Os monids suggests that its food determines
whether a given fish will be white-
Flesh 1944 1945 1946 1947 1948 1949 5 :
White gr) are) eek ka Mateo EE fleshed, pink-fleshed, or red-fleshed, with
Payer 27. 28 25 26 29 «+55 a crustacean diet favoring the last state;
Red ...... 65 49 44 59 53 18 no definite evidence, however, seems yet
* From Andrews and Lear, 2: 857, tab. 11. to have emerged in this regard for a/pinus.
Size. The averages given in Table v
show roughly the relationship between length (mm) and weight (lbs) at four localities.
It appears from Table v that the Arctic Charr of Hudson Bay, of northeastern Labra-
ne Memoir Sears Foundation for Marine Research
Table V. Averages (in pounds) Showing the Relationship between Length and Weight
of Arctic Charr from Northern North America and Spitsbergen
a Weight (1b) ———______—_
Length (mm) Frobisher Bay Northeastern Western Southern
Baffin Islandt Labradortt Hudson Bay* Spitsbergen**
250-295 ...... 0.5 =
ZOC— alc iaeeris 0.6 - -
350-395 ...... I.I 1.5 1.5 Tas
400-445 ...... Tey 1.9 2.4 2.2
450-495 ...... 2.4 Ag 3.3 3-3
500-545 ...... 3.3 3.8 3.75 aos
550-595 ...... 4.4 5-1 5.2 4.8
600-645 ...... 5.8 a7, 7.0 5.5
650-695 ...... 752 6.9a 8.34 -
7OO-745 ...... 8.4 = - 8.8
T5O—7.9 Sue ise 10.2 - jeeyy -
+ Grainger, 28: 351. tt Andrews and Lear, 2: 853, tab.7. * Sprules, 62: 6, tab.1. ** Dahl, rz: 6.
a One fish only.
dor, and of Spitsbergen average slightly heavier at equal lengths than those of Baffin
Island. The general run, as caught, is reported as averaging about 4.3 pounds for
the northwestern coast of Hudson Bay, with 56 °/, of the catch falling between 3-4
pounds. In Ungava Bay, the most common size is between 2—8 pounds. For southern
Baffin Island the average is about 3.5 pounds. The bulk of the catch for northeastern
Labrador runs from a pound or two up to eight or even ten pounds per fish;
spawning fish in poor condition, caught in a brook tributary to Anatalek Bay, weighed
close to five pounds. Most of the western Greenland fish weigh between 5.5—7.7 pounds.
The maximum weights (in pounds) that I have found reliably recorded have
been 16 (7.3 kilo) for Hudson Bay; 9 for the south coast of Baffin Island and 11.9
for Frobisher Bay; 10 for northeastern Labrador; 8.5 for western Greenland; and 8.8
(4 kilo) for Novaya Zemlya. However, occasional a/pinus grow much larger than the
majority; specimens of 20 and even 26 pounds have been reported for the north-
western Hudson Bay area (62: 5; 35: 24), and a giant of 35 pounds (16 kilo) has been
reported for southern Novaya Zemlya (Yessipov, 79: 70), though not on wholly con-
clusive evidence.
Development and Rate of Growth. The young fry under hatchery conditions have
been reported as averaging about 17 mm in length at hatching. At room temperature
(at Ostanback, Sweden) they averaged 27 mm after 19 days, a rate of growth no doubt
more rapid than that for “wild” fish exposed to considerably lower temperatures in
nature; for example, Nystrom reported that fry artificially hatched in January in
Sweden grew so slowly thereafter in the cold spring-fed fish ponds that they were
only a little more than an inch (about 25 mm) long the following August (i7 61: 847).
Average lengths for hatchery fish have been reported as 97 mm at the end of their
first year, 123 mm at the end of their second year, and 137-216 mm at the end of
their third year (61: 847). These lengths seem somewhat greater (thanks to warmer
Fishes of the Western North Atlantic 518
Table VI. Average Lengths (mm)* of S. a/pinus at Different Ages for Localities in
North America, Greenland, and Spitsbergen
Labrador Herschel I. Frobisher Bay ae)
Ages** aes (Hebron to Arctic Baffin MG cir ea
eons ih Ramah)tt Canada% Island Oe ee
Gi riaphe oats 382 — 246+ 130 201-224 369
Ohta sists 408 _— 362+ 139 237-258 412
Tan ee oe 446 412-449 407+ 172 275-310 483
Seis 468 423-557 425t 302 268-525 533
Qielsxsia vais 494 442-576 421+ 337 430-546 611
LO everccesuets 524 472-584 524+ 363 480-554. 670
TDs cisiciscayats 567 478-626 557+ 414 489-571 7oot
Teas cotta 597 495-626 — 425 519-550 —
ING fst anal sets 638 503-656 —_— 484 5 30-606 ao
Wiselc Caade 636 608+ _ 510 — —
nGesaepoee --- 671+ — 553 — —
TOmets ccc s 682+ 764+ — 557 605 _
Ween teels 791 — — 559 = _
Ws coaoeer 750+ _ _— 600 — —
H@sadoosor — —_ = 613 — —_
AO berms aes _ — _ 649 — —
Wai sre leteve a — — 642 — —
ara Noitets 822+ —- — 655 — —
2B ietereiais arece — _ — 678 — —
DAreieseieiess, <= — —- — 678 “= —
QAst ae le sts — —_— — 685 — os
* Measurements for Labrador and Frobisher Bay fish were given as fork length (snout to fork of caudal
fin); those for Herschel Island, originally given as standard length, have been adjusted here to fork length by
adding 9°/o of the standard length given. The standards of measurement were not stated for either western
Greenland or Spitsbergen.
** Ages for Hudson Bay, Labrador, western Greenland, and Spitsbergen were given in years, those for
Frobisher Bay and Herschel Island in “winters.” Those for Labrador, Frobisher Bay, Herschel Island, and
western Greenland were derived from studies of the otoliths, those for Hudson Bay and Spitsbergen from
studies of the scales.
+ Sprules, 62: 8, tab.2. + Andrews and Lear, 2: 851. «a Grainger, 28: 341-348. A Hansen
in 28: 354, 369. £6 Dahl, rr: 1-12. + One fish only.
water and to a more abundant food supply) than those to be expected among the anadro-
mous populations with which we are immediately concerned.
The average lengths reached at equal ages by the Arctic Charr at different local-
ities are summarized in Table vi. Although the data are not strictly comparable (see
Table v1, footnotes), it seems evident from these data, added to other available informa-
tion, that: (1) “the char[r] grow very slowly and may reach an age of more than
24 years” (28: 327); (2) the rate of growth varies widely from place to place, even
within short distances, both for Labrador and for the western Greenland coast; and
(3) sea-run Arctic Charr grow considerably faster along the Atlantic coast of Labrador,
in the Hudson Bay region, near Herschel Island, along the coast of western Greenland,
and at Spitsbergen than they do in Frobisher Bay, Baffin Island. Grainger has pointed
out also that those within a given age class may vary widely in size, irrespective
33
514 Memoir Sears Foundation for Marine Research
of locality (28: 348). Fish of 600 mm, “for example, are found in all age groups
between 13 and 24 winters.” The over-all picture for the western North Atlantic area
as a whole, and for Hudson Bay, is of a very long-lived and slow-growing fish, with
the relationship between age and size averaging roughly as shown in Table vir.
Dahl’s observation that such of the
Table VII. Approximate Average Lengths of 4- to a/pinus of Spitsbergen as run down to the
ce ae sea after four years in fresh water grow
Bee Fork Length Age _ Fork Length larger than those that remain in fresh
(years) Ce) Ve) ) water for two or three years (Table vir)
ARS 35. go-201 II 414-626 ; -
pure T genase Bs 425-625 Suggests that the longer an a/pinus is a
GAA ice 123-408 13 484-656 parr, the more rapid is its subsequent
aa ae 14 510-636 growth in the sea.
severe eies 208-557 15 553-571 : F
neh nope se aera The populations of alpinus that re
Towa 363-584 main permanently in fresh water grow
more slowly than those of the same region
that go to sea. Sprules (62: 9), for example, has pointed out that landlocked a/pinus
in the northwestern Hudson Bay region average only about 12.3 inches and about
nine ounces when ten years old, whereas the sea-run Arctic Charr of that same age
average about 20.6 inches and 3.75 pounds. In western Greenland, similarly, the
landlocked fish found above falls are smaller than
those that are sea-run (52: 294; 39: 74). Table VIII. Lengths (mm) of Sea-run
Survival Rate and Longevity. The great ma- 5. 4/pinus of Different Ages, as Related to _
wey fa pe f theast leah the Number of Years Spent in Fresh Wa-
EA ect TAPP ens Roe Men ea Se et abra~ ter Previous to their First Migration to
dor are between 7+ and 12+ years old, with a Salt Water*
pronounced peak at 8+ to 11+ years and a rapid
: 7 Age at First Migration —
falling off after 13+ years, as appears from Table Years
ix. Also, the mean age (in years) of the samples _ in sea yi adie ae
studied showed an increase from south to north 2...... 340 361 390
as follows: Adlok, 8.5; Nain, 8.9; Okkak, 10.2; 3°": rT: 377 493
Ant tererts 446 450 487
Hebron, 10.5; Ramah, 10.5. et 516 £38 £70
In Spitsbergen waters, according to Dahl, 6...... 590 603 610
84 °/, of the fish examined were between about ~* After Dahl, zz: 7.
4-6 years old and about 7~9 years old, including
2—4 years passed as parr in fresh water (I: 6); and an occasional fish had even
spent as much as eight years in fresh water. In Frobisher Bay, Baffin Island, ac-
Table IX. Number of Arctic Charr of Different Ages among 487 Fish from the Coast
of Labrador, Adlok to Ramah Combined t
Age (years)* ... 7+ 8+ 9* 10+ Ir+ 12+ 13+ 14+ 15+ 16+
No. of fish..... 47 69 108 99 66 44. 30 5 5 2
Percentages .... 10 14 22 20 14 9 6 I I 0.3
+ From Andrews and Lear, 2: 849, tab. 3. * Based on a study of the otoliths.
Fishes of the Western North Atlantic SiS
cording to Grainger’s age determinations (28: 341), the great majority of 1,566
specimens measured during 1948, 1950, and 1951 were 300-350 and 650-695 mm
long, with a few longer than 795 mm; these sizes correspond roughly to ages of 9-10
and 19-24 years, respectively, with a few somewhat older still. A similar picture
emerges for the northwestern part of Hudson Bay, where only seven fish among 96,
for which Sprules determined the ages, were less than seven years old whereas 70 °/,
were between 8 and 11 years old (62: 6, tab. 1); and while the survival rate dropped
off abruptly after 11 years, one of the specimens was 22 years old and weighed 16
pounds. Thus, the sea-run a/pinus are not only much longer-lived than the sea-run
fontinalis but their rate of survival is much higher.
Spawning and Reproduction. Spawning may take place either in streams or in lakes,
at depths ranging from within a foot or two of the surface down to 15 feet or so.
Sprules reported that, in lakes, the eggs are deposited on gravel beds in 6-15 feet of
water (62: 12). In rivers they are laid in shallow pools below rapids in about two feet
of water, where the current is strong and the bottom is covered mostly with ‘‘round
stones just too large to be carried along by the water.”’ Here the fish lie ‘‘apparently
motionless, ... and ... return to the same places almost immediately after being
frightened away” (Weed, 72: 132). The females of the landlocked fish of Lake
Windermere, England, have been described as making shallow depressions for the
reception of their eggs in the gravel or stony bottom (Jackson iz 61: 847), as other
salmonids commonly do, but this habit has not been recorded for the sea-running
populations.
Normally, a/pinus, like fontinalis, spawn in fresh water, but Weed has written that
“the fishermen along the Labrador coast believe that some of the sea trout spawn in
the sea” (72: 132); and he reports the capture in salt water of females with free eggs
in the oviducts. There seems to be no reason, however, to believe that any eggs would
survive that might be deposited in salt water.
Arctic Charr have been described as spawning for the first time in Spitsbergen
when they are 5+ years old (zr: 8), in Icelandic waters probably at about 6 years
(Saemundssen iz 28: 363), and around Novaya Zemlya at 6 or 7 years of age (79:
69, 70), but not until about their 12th year or later in Frobisher Bay, Baffin Island
(28: 363). Thus, if these estimations are even approximately correct, a/pinus is very
much slower than fontinalis in reaching sexual maturity (p. 531). It appears also from
evidence assembled for Baffin Island by Grainger (28: 363) that spawning may not
be a regular annual event for every adult, though it seems hardly likely to be a bien-
nial one regularly, as has been suggested for Arctic Charr of Novaya Zemlya (79).
Labrador females of 465-665 mm contained 2,173-7,223 ovarian eggs in dif-
ferent stages of development, a western Greenland female of 520 mm contained 4,620
(28: 364; 39: 69), and Novaya Zemlyan fish averaged about 3,500 eggs (79: 69-70).
The eggs of freshwater Arctic Charr of Scandinavia are 4-5.5 mm in diameter. |
have found no record of the size of naturally spawned eggs of the sea-run popula-
tions of alpinus.
hs
516 Memoir Sears Foundation for Marine Research
Under hatchery conditions, incubation of the eggs of freshwater Arctic Charr
has been found to occupy between 60-70 days at a temperature of 40°F (4.4°C)
(in 61: 847). Eggs that were artificially fertilized at Baldernas in Dalsland, Sweden, at
the end of October did not begin to hatch until the beginning of the following March
(Malm iz 6r: 847). In the northwestern Hudson Bay area “the eggs are still visible
at break-up [of the ice] in the following spring, probably hatching soon after open
water appears” (Sprules, 62: 12).
Migrations. The migratory schedule of the sea-running populations of a/pinus
consists of a downstream movement early in summer out into brackish or salt water;
a sojourn there until late summer or early autumn, during which time they make
most of their year’s growth; a return movement in late summer or early autumn to
fresh water, with the immature fish participating as well as those that are maturing
sexually; and a sojourn in fresh water until the following June or July, the early part
of which covers the spawning period.
Arctic Charr of Frobisher Bay, Baffin Island, are reported as carrying out their
first downstream migration after they have spent 5—7 winters in fresh water, those
of Spitsbergen migrate after 2, 3, or 4 years, and those of Novaya Zemlya after 3 or 4
years. Those of northeastern Labrador, however, may descend after only 2 or 3 years to
judge by the presence in the bays there of fish not more than three or four inches
long, sometimes several miles away from the nearest spawning stream (72: 132).
The spawning migration into fresh water takes place between late July and the
last half of September, according to locality, with the precise date governed to a con-
siderable degree by the height of the water in the streams. Heavy rainfalls, for example,
swell the streams so that Arctic Charr can ascend falls that would otherwise bar their
passage, for they do not have the leaping powers of salmon and often are “stopped
by falls with a vertical drop of not more than a foot” (Weed, 72: 132). For this same
reason also, their ascent is much easier during periods when the tides are at their
highest (spring tides) than when they are at their lowest (neap tides).
Along Southhampton Island they congregate at the head of tide by late August,
to ascend either with the first spring-tide period or during the first heavy rainfall, which-
ever comes the first. In Ungava Bay, in both the George River region and the Kok-
soak River, Arctic Charr enter fresh water by the latter part of July in most years and
during the early part of August in others, with the height of the run lasting from
eight to fifteen days (Turner iz 16: 99, ftn.). In the vicinity of Nain, on the coast of
northeastern Labrador, in the one year of record (1928), they had entered the streams
by August. In the Sylvia Grinnell River, Frobisher Bay, Baffin Island, the mouth
of which is barred to them by falls that are passible by fish only at the time of spring
tides, the fish were ascending by the last week of July, in 1950; in 1948, however, few
ascended until August. In western Greenland they have been described as ascending
from late July through August until the streams freeze once more; in northeastern
Greenland (Denmark Bay) the run continues from August until about mid-September,
when the local rivers freeze; and the run covers about this same period in Novaya Zemlya.
Fishes of the Western North Atlantic 517
It seems that the fish generally tend to continue upstream until they meet some
obstacle that bars their further progress. In the larger streams or rivers this is likely to
be the first waterfall; in many cases, however, the governing factor lies in the length
of the stream in which they spawn. In western Greenland, for example, the narrow-
ness of the ice-free belt limits the maximum distance of their upstream migrations to
25-30 miles. For more extreme cases it seems sufficient to refer the reader to
Dunbar and Hildebrand’s report of spawning fish taken in the lowest half mile of a
stream tributary to the estuary of the George River (Ungava Bay, northern Quebec),
beyond which further progress is barred by falls (16: 99); and to Weed’s account
of alpinus spawning about five miles up a brook tributary to Anatalek Bay, near Nain,
Atlantic coast of northern Labrador (72: 131, 132). Turner, on the other hand, reported
the presence of a/pinus more than 110 miles up the Koksoak River, tributary to Ungava
Bay, and farther still up a tributary of the Koksoak (in 16: 99, 100, ftn.)—if indeed
the fish in question were a/pinus, not Atlantic salmon, which run up the Koksoak
regularly in considerable numbers (p. 476).
The descent to salt water usually takes place at about the time the river ice breaks.
At Southhampton Island, northwestern Hudson Bay, for example, the descending fish
have congregated within the mouth of Brooks River by the last part of June, and
catches have been made in salt water nearby as early as July 8 in some years (e. g.,
1934) but not until after July 15 in other years (47: 128, 129). In Frobisher Bay,
Baffin Island, in 1951 (the only year of record), the river ice broke on June 10 and
Arctic Charr were caught outside the mouth of the river in salt water shortly thereafter
(28: 355). I find no definite information as to the date of the downstream migration
for Labrador. Local ice conditions suggest, however, that it may fall at about the same
time there as in Frobisher Bay, though in western Greenland the outlets of the fresh-
water lakes remain frozen until July, when there is a rush of a/pinus to the sea (McMil-
lan iz 32: 1). Similarly, in northeastern Greenland, where the river ice persists late
into the season, it was not until the first week of July that Johansen’s nets, set in the
outlet of a lake tributary to Denmark Harbor, took this species (¢0: 669). Arctic Charr
in Novaya Zemlya are described as moving out into salt water during June and the
first half of July (79: 69, 70).
It is not yet known whether every adult returns to fresh water to spawn every
year, whether every fish that has spawned in a given autumn invariably returns to sea
the following summer, or whether any of them winter in the sea with any regularity
in any given locality.
The few accounts that have appeared concerning the distribution of a/pinus while
they are in salt water, based on their numerical distribution there, are unanimous to
the effect that they remain for the most part close to the mouth of their native rivers,
or at least within the bays into which these open. Perhaps the most definite evidence
in this regard is Manning’s observation that, while they are regularly caught at Igloolik
Island, three miles off the mainland of Southampton Island, nortwestern Hudson
Bay, none was caught “‘at Walrus Island, 30 miles off Southhampton Island, although
518 Memoir Sears Foundation for Marine Research
a net was kept out there for a week in early August 1936” (47: 128). It seems well
established, also, that their wanderings along shore are so brief that but little inter-
mingling takes place between local populations. The supporting evidence for this is
of two sorts: (1) the development of an intensive fishery has, in at least two documented
instances, been followed within a few years by a precipitous decline in the number of
fish taken; (2) it has been found by Andrews and Lear that the number of fin rays
and of vertebrae average progressively higher from south to north along the north-
eastern Labrador coast (2: 854, tab. 8, 856, tab. 9, 860), which would not be the case
if much intermingling took place between the populations that are produced in the
various rivers along this stretch of coast.
Food. Sea-run Arctic Charr, like the closely related sea-run fontinalis (p. 536),
feed on active organisms; to all intents and purposes this limits the diet to fishes smaller
than themselves and to crustacea, supplemented on occasion by nereid worms. In Un-
gava Bay, for example, a/pinus have been reported as feeding chiefly, if not exclusively,
on the crustacean genera Gammarus and Pseudaliprotus, fish not having been mentioned
at all (r6: 100). On the coast of southern Spitsbergen also, large gammarids (Gam-
marus locusta) as well as mysid shrimps provide their chief sustenance, with fishes
(capelin, small cottids, small liparids) of secondary importance (zr: 8). Similarly,
in Frobisher Bay, Baffin Island, the gammarid genera Pseudaliprotus and Gammara-
canthus, together with liparids, were dominant items in their diet; among the 34 food
species listed was a variety of small crustaceans (copepods, euphausiid shrimps, clad-
ocerans, and decapod larvae), nereid worms, and fishes (such as Eumicrotremus spinosus,
small cottids, liparids, launce, and Boreogadus saida, together with young of their own);
even a few insects were included (28: 366, 367).
In Hudson Bay and along northeastern Labrador the recorded diet has been divided
in varying proportion between fish (chiefly launce, Lumpenus, capelin, cottids, and
Cyclopterus) and amphipods and euphausiids. Table x, showing the percentages of
different items in the stomach contents, illustrates the variation to be expected from
place to place.
Table X. Percentages, by Volume, of Different Species in the Stomach Contents of
Arctic Charr Taken at Successive Localities from South to North along the Coast of
Northeastern Labrador*
77. Fish ———_,
antes )
Locality Capelin Launce Sculpins Amphipods and
Euphausiids
IAGlOkire sewers vee 88.0 3.0 fo) °
Nain, specimen 1 .... 75.2 10.7 ° 13.5
Nain, specimen 2 .... 15.0 12.7 61.2 II.I
OkkaksBaye cs iricisiecie 99-3 0.2 ) 0.5
ILE DEOU a etcn etoteteletsiel- 2.9 0.3 14.9 81.3
PRATT A Lo aten os ctoteyerere et 3.9 13.5 24.7 56.2
* From Andrews and Lear, 2: 858.
Fishes of the Western North Atlantic 519
Finally, Yessipov’s report (79: 69, 70) that the sea-run Arctic Charr of Novaya
Zemlya, where 75 °/, of their food consists of crustaceans and worms, eat the young
of the Atlantic cod (Gadus morrhua) in greater amount than any other fish is especially
interesting, for young cod are not mentioned at all in any of the other diet lists that I
have seen.
Parasites. Sea-run alpinus of northeastern Labrador have been reported as being
heavily parasitized by the tapeworm Exbothrium, and lightly so by the round worm
Philomena (2: 858, 859). It seems astonishing that none of the accounts of this species
has mentioned any infestation by the so-called sea lice (parasitic copepods, genus
Lepeophtheirus) that are so often found clinging to the sides of Atlantic salmon (p. 484)
and sea-run brook trout (p. 537) when they re-enter fresh water.
Variations. Variations have been reported in the colors of sea-run a/pinus, the size
of the mouth, the length of the pectorals, the average number of vertebrae, the
number of fin rays, and dentition. However, the relative degree of “‘silveryness’’ of the
Arctic Charr caught during their marine sojourn together with the distinctness of their
color pattern seems clearly to be associated with the relative salinity of the water where
they are taken combined with the length of time since they last left fresh water. The
difference in the relative length of the pectoral fins between sexually mature freshwater
alpinus of Labrador (average 17-19 °/, of TL) and those taken in salt water (not in
breeding condition) does not bear directly on the racial problem of saltwater fish (3: 290).
The south-to-north increase in the average number of both fin rays and verte-
brae that has been reported for the coast of northeastern Labrador (2: 854-856)
evidently is a function of latitude, a relationship paralleled among sundry other coastal
fishes of boreal-Arctic waters; thus it is environmental, not genetic. The small-mouthed
fish reported by Weed for the vicinity of Nain, Labrador (72: 131), doubtless rep-
resented the female portion of the pop-
ulation, the very large-mouthed ones
“with strongly hooked jaws” the ma-
ture males. While the sea-run Arctic
Charr from Collinson Point, Alaska
(70: 258, fig. 14), included some that
had an upper pattern of dentition that
is generally characteristic of Sa/velinus
(p. 527, Fig. 127), others had a pat- Ficure 127. Salvelinus alpinus dentition.
tern characteristic of the genera Hucho
and Brachymystax (pp. 458, 459, Fig. 1278, c); in this case there can be no question
of regional varieties, since both types were taken at the one locality.
In short, the regional differences reported thus far for the populations of sea-
run a/pinus do not seem either wide enough or consistent enough to justify their
formal recognition in systematic nomenclature for a/pinus anywhere between Novaya
Zemlya in the east and northwestern Alaska in the west. Farther westward the typical
alpinus is replaced, at least predominantly, by the red-spotted form ma/ma, which
520 Memoir Sears Foundation for Marine Research
probably does represent a separate subspecies (p. 519).1° Neither has my own rather
cursory examination brought to light any differences that seem taxonomically significant
between my Spitsbergen, Greenland, and North American specimens of a/pinus and
others from Sakhalin Island, Gulf of Tartary, with which I have compared them.
Numerical Abundance and Relation to Man. Attention has been called repeatedly
to the great abundance of the Arctic Charr off the stream mouths in Arctic America.
The famous explorer, Capt. James C. Ross, reported in 1835 the capture of 3,378 fish
ranging from 2 to 14 pounds in weight and averaging something more than 4 pounds,
off a stream in the Boothia Peninsula (58: lvi, ftn.). Bryant described the “sea trout”
similarly as being so numerous in Komaktorvik Bay, northern Labrador, that ‘“‘you
could almost walk across the rivers without wetting your feet.... They were only
about one or two feet apart all over the shoal places” (Kendall, 44: 503); and Welch
has recently reported “incredible numbers” in the Sylvia Grinnell River, Frobisher
Bay, Baffin Island (73: 14).
In northeastern Labrador, 264,000 pounds of salted fish were marketed in the
Hopedale-Hebron district in 1948, and 224,620 pounds in 1953, corresponding to
something like 62,000—71,000 fish if these averaged between three and four pounds
each; and a single schooner took 79,200 pounds, or something like 20,000—26,000
fish, in Nachvak Bay between early July and mid-August in 1948 (2: 845). On the
coast of western Greenland, in 1914, about 24,250 pounds of Arctic Charr, or
about 6,000 fish if these averaged four pounds, were netted at the head of one of
the fjords near Godthaab; and for Denmark Harbor, northeastern Greenland, they
were described by Johansen as being so numerous that “as many as 50-100 could be
taken in one net in one day” (40: 669).
It is no wonder that an excellent food, so abundant locally and so readily available
close inshore, should be an important item in the economy of the native inhabitants,
not only for human consumption but for dogfood as well. The reported catch, for ex-
ample, averaged about 125,620 pounds (salted) for the Hopedale-Hebron section
of Labrador for the 10-year period 1944-1954, with maximum catches of 264,000
pounds and 224,620 pounds in 1948 and 1953, respectively (2: 845). On the north-
western Hudson Bay coast, gill nets yielded an average of about 150 pounds per 100
yards at a locality about 150 miles north of the Churchill River (62: 2). A yearly
commercial catch ranging between 100 and 800 barrels (2,200 and 176,000 pounds,
assuming the weight of a barrel to have been 220 pounds) has been reported for
western Greenland (39: 71). And Yessipov reported a yearly catch ranging between
14 and 80 Russian tons (79: 63), corresponding to 31,000-176,000 pounds," for
Novaya Zemlya.
Between Hopedale and Hebron as a whole, the local stock of Arctic Charr did not
show any signs of depletion during the ten years covered by Andrews and Lear’s report;
ro. On the Arctic coast of Alaska, malma is reported from as far East as Point Hope and Cape Lisburne near 166°W
(Evermann and Goldsborough, 21: 264), nominally even from Herschel Island, Arctic Canada (139°W), but with-
out supporting evidence as to identity.
tr. The Russian metric ton equals 2,204.6 pounds.
Fishes of the Western North Atlantic Ret
in fact, the average for the last four-year period (145,000 pounds) was more than
twice that for the first four-year period (67,320 pounds). Andrews and Lear have
pointed out, however, that the trend of production may not represent a corresponding
trend in the numbers of fish, since the intensity of the fishery there depends on whether
it is the more profitable to fish for cod or Arctic Charr (2: 845).
However this may be, the intensive exploitation of a more or less isolated stock
has resulted in unmistakable depletion in two well-documented cases: in Nachvak Bay,
northern Labrador, where the catch made by the same schooner decreased from 79,200
pounds during July-August of 1948 to 11,000 in 1950, when the project was abandoned
(2: 845); and in western Greenland, where the catch at a certain cannery at Godt-
haab fell from 24,955 pounds in 1915 to 12,540 in 1917, when the cannery was
closed (39: 72).
Although Arctic Charr may seem almost unbelievably plentiful when crowded to-
gether off the mouth of some stream, and while they may in fact be plentiful enough
locally to withstand a moderate amount of fishing, it appears that the populations that
are more or less isolated geographically cannot long withstand intense exploitation.
“Even on the normal Eskimo scale of fishing, streams may become seriously depleted
and require a rest for some years before their numbers are restored” (Dunbar and
Hildebrand, 76: 100). Dunbar has reached this same conclusion for those of Frobisher
Bay, where, fortunately for the fish, many of the “charr streams were not touched at
all by the Eskimo” (15: 178-190).
The catch of Arctic Charr is made chiefly in gill nets or in stake nets, close to
the land in water only a few feet deep, and during the months when they are in the sea
(p. 516). Some are also caught through the ice in winter while they are in fresh water,
at least in the Hudson Bay region.
The sea-running Arctic Charr have not yet attracted the attention of anglers in
general, nor are they likely to do so unless the Arctic-subarctic regions become more
easily accessible; but visitors to the airfield at Frobisher Bay report having great sport
with them there (5: 333 73: 4).
General Range. The alpinus species complex (freshwater and sea-running popula-
tions combined) is circumpolar and widespread in Arctic-subarctic regions, with iso-
lated (doubtless relict) populations in some cold lakes far south of the boundary of
their general range. Their area of more or less continuous occurrence in America
embraces Arctic Canada and Arctic Alaska, and the Arctic Archipelago at least as far
northward as Discovery Harbor, Grinnell Land, 82°34’N (30: 294, 295), which is
the northernmost locality whence any salmonid has been reported. S. a/pinus range
southward along the Atlantic coast to the northern side of the Gulf of St. Lawrence,
to eastern Newfoundland, and to both coasts of northern Hudson Bay; and along
the Pacific coast to northern California. There are isolated populations also in many
of the deep, cold lakes of Quebec, northern Maine, northern New Hampshire, and
northern Vermont.
Arctic Charr occur around Greenland from the north coast down the west coast
522 Memoir Sears Foundation for Marine Research
and up the northeast coast at least to 77°N. Their range also embraces Iceland, Spits-
bergen, Bear Island, and Novaya Zemlya; northern Europe in general, including
various alpine lakes southward to the British Isles, Austria, and Switzerland; the Arctic
coast of Siberia, with outlying populations as far south as Lake Baikal; and the
coastal belt of northeastern Asia southward to Kamchatka, the Sea of Okhotsk, Sak-
halin, and northern Japan.
Occurrence of Sea-run Populations in the Western North Atlantic, with its Arctic Trib-
utaries, in Hudson Bay, along Arctic Canada and Alaska, and in the northern North Pacific.
The mouth of the Gander River on the coast of eastern Newfoundland is the southern-
most point on the Atlantic coast where a sea-run population of a/pinus is known to be
established definitely.% In the northern side of the Gulf of St. Lawrence, Arctic Charr
have been reported to occur as far west as Trinity River and Bay, tributary to the
Saguenay River (10: 352-355, as S. oguassa; 18: 185); and by common report, they
occur generally off the river mouths thence eastward along the north shore, though
they are so overshadowed there in public interest by the Atlantic salmon that precise
information as to their local status is lacking. I have found no report of them for the
west coast of Newfoundland, though here, as in the case of the east coast, I suspect
that a/pinus as well as fontinalis may be represented among the larger “‘sea trout”
taken there.
Arctic Charr are recorded for the St. Mary’s River, Hawke Harbor, and Domino
Island along the southern part of the Atlantic coast of Labrador. Although the pub-
lished record does not suggest that they are plentiful anywhere south of the region
of Hamilton Inlet, Arctic Charr have been reported (or are represented in museum col-
lections) for every locality farther northward whence saltwater fishes have been listed:
along the Labrador coast in the vicinities of Mokovik Bay, Hopedale, Nain, Cape
Mugford, the Okkak Islands, Hebron, Ramah, Nachvak and Komaktorvik fjords,
and Ryans Bay near Cape Chidley; the recorded localities also include the Ungava
Bay region, in general, including Port Burwell; Hudson Strait (reported for Notting-
ham Island and Wakeham Bay); and Foxe Basin on the coast of western Baffin Island
as well as Frobisher Bay and Cumberland Sound on the east coast. Around Green-
land, sea-run Arctic Charr are known to range northward to 77°N, at least on the
east coast; to Etah (78°20’N) on the west coast (in fresh water to Inglefield Land,
79°); and even to the north coast, where a small specimen was taken by Lange Koch
in 1917 (39: 68).
Sea-running populations of a/pinus are similarly widespread around the coast of
Hudson Bay southward as far as the vicinity of Great Whale River, in the east, and
of Fort Churchill, in the west; Boothia Peninsula, Wallaston Peninsula, Victoria Island,
and Herschel Island of Arctic Canada; Camden Bay, Point Barrow, and Collinson
Point of Arctic Alaska; and various rivers of Arctic Canada and Arctic Alaska.
Along the northwest coast of America from the region of Cape Barrow south-
ward to northern California, in salt water, S. a/pinus is represented by a red-spotted
12. The color description on pp. 510, 511 is based on Gander River specimens.
Fishes of the Western North Atlant 523
charr, the so-called “Dolly Varden,” which has commonly been ranked as a separate sub-
species, malma Walbaum 1792 (pp. 519, §25). They range southward on the Asiatic
side to Kamchatka, the Sea of Okhotsk, Sakhalin Island, and northern Japan.
Synonyms, with References for Sea-run Populations of Greenland and America westward to the International
Boundary between Canada and Alaska:
Salmo alpinus Linnaeus, Syst. Nat., ed. 10, I, 1758: 309 (Lapland, alpine lakes, England); Fabricius, Fauna
Groenl., 1780: 173 (descr., range as for §.carpio Fabricius, i.e. lakes, rivers, river mouths, Green-
land); Johansen, Medd. Gronl., 45 (12), 1912: 668-694 (size, color, migr., season, abund. Denmark
Harbor, ne. Greenland).
Salmo carpio Fabricius (not of Linnaeus, 1758), Fauna Groenl., 1780: 170 (detail. descr., diet, lakes, rivers, sea
at river mouths, Greenland).
Salmo salar Pennant (not of Linnaeus, 1758), Arctic Zool., Introd., 2, 1784: cxcr (abund. along beaches, w.
side Hudson Bay N. of Churchill R.; Arctic charr evidently mistaken for Atlantic salmon).
Salmo rossii Richardson in Ross, Narr. Second Voy. ... North-west Passage, Append., Nat. Hist., Fish., 1835
(descr., abund., Boothia Penin., Arctic Canada); Fauna Boreal.-Amer., 3, 1836: 163, pl. 80 (descr.,
color. ill., abund., Boothia Penin., Arctic Canada).
Salmo immaculatus Weiz (not of Storer, 1850), Proc. Boston Soc. nat. Hist., 10, 1866: 274 (Hopedale, Labrador) ;
Packard, Labrador Coast, 1891: 400 (Hopedale, Labrador).
Salmo stagnalis Garman (not of Linnaeus, 1780), rgth Rep. Comm. inl. Fish. Mass. (1884), 1885: 80, fig. 19
(descr., color, good ill., Godthaab, Greenland; in MCZ).
Salvelinus stagnalis Dresel, Proc. U.S. nat. Mus., 7, 1885: 255-258 (descr., color, meas., Disco I., w.Green-
land); Kendall, Proc. Portland Soc. nat. Hist., 2 (8), 1909: 228, 230, 231, 232, 234, 236 (early Labrador
rec.); Proc. U.S. nat. Mus., 38, 1911: 503-510 (color, meristic counts, discus. of ident. with Greenland
and Ungava Bay populations, util., abund. Komaktorvik Bay, n. Labrador); Henn, Mem. Carnegie Mus.,
12 (2 [3]), 1932: 1-2 (meas., Southampton I. specimen, migr., seasons, Greenland); Hildebrand, Medd.
Gronl., 125 (1), 1939: 5 (e. Greenland locals.; also Foxe Basin and Melville Penin., Arctic Canada).
Salvelinus alpinus stagnalis Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 510 (descr., refs., Green-
land, Boothia Penin.).
Salvelinus umbla stagnalis Dahl, Skr. Svalb. Ishavet, 7, 1926: 1-12 (age at first sea migr., growth, size at ma-
turity, food, Spitsbergen).
Salvelinus alpinus Halkett, Bull. nat. Mus. Canada, 53 (Biol. Ser., 15), 1928: 117, 118 (abund. s. Baffin I.,
discus. subsp. stagnalis, alipes, naresii, arcturus, oguassa); Soper, Bull. nat. Mus. Canada, 53 (Biol. Ser.,
15), 1928: 116 (s. Baffin I.); Dymond, Canad. Field Nat., 46, 1932: 185 (n. shore Gulf of St. Law-
rence); Vladykov, Contr. Canad. Biol., N.S. 8 (2), 1933: 19, 20, 39 (spawn. season, sizes, capture, util.,
Hudson Bay and Strait, consid. distinct from Greenland and Arctic forms); Weed, Copeia, 1934: 131-133
(size, migr., spawn., color, food, cf. fortina/is, ne. coast Labrador); Manning, Canad. Field Nat. (1942),
56, 1943: 128—129 (season, habits, sizes, abund., fishery, Taverner Bay, w. Baffin I., Melville Penin.,
Arctic Canada, in fresh and salt water); Hinks, Fishes Manitoba, Dept. Mines nat. Resources, 1943: 24
(genl. range, habits, abund., max. size); Jensen, Spolia zool. Mus. Hauniensis, 9, 1948: 67-78 (abund.,
local distr., w. and e. Greenland, color, size, secondary sex. chars., migr. schedule, util., fishery, cf. freshw.
populations; discus. forms named by Fabricius, 1780; genl. range, growth rate cited from Hansen, Beretn.
verdror. Gronl. Styr., 1940 [1]: 362-363; not seen); Sprules, J. Fish. Res. Bd. Canada, g (1), 1952:
I-15 (sizes, growth rate to 22 years, dominant ages, migr., spawn. season, second. sex. chars., abund.,
fishery, w. coast Hudson Bay); Dunbar and Hildebrand, J. Fish. Res. Bd. Canada, 9 (2), 1952: 97-100
(sizes, fecund., diet at sea, extent and dates of upstream migr., spawn. season, fishery, Ungava Bay region,
n. Labrador); Grainger, J. Fish. Res. Bd. Canada, ro (6), 1953: 326-370 (fishery, descr., ills. of otoliths,
age-length relation. based on otoliths, weights, growth rate for other locals., migr., egg sizes, fecund., diet,
Frobisher Bay, Baffin I.; also list of sp. and subsp. names for a/pizus species complex in Greenland, N.
America, and n. Asia by ref. to Dymond and Vladykov, Proc. 5th Pacif. Sci. Congr., Canada [1933], Fish.,
1934: 3741); Walters, Amer. Mus. Novit., 1643, 1953: 2 (food, color, persist. of parr marks, var. in
dentition, Mould Bay, Prince Patrick I., Arctic Canada); Bull. nat. Mus. Canada (1951-1952), 128,
1953: 258, fig. 14, 260 (var. in dentition, Collinson Point, Alaska; Bernard Harbor, Arctic Canada;
524 Memotr Sears Foundation for Marine Research
locals. in Arctic Canada, Arctic Alaska); Andrews and Lear, J. Fish. Res. Bd. Canada, 13 (6), 1956: 843-
860 (fishery, age determ., growth rate, sizes, meristic counts, color of flesh, diet, parasites, ne. Labrador).
Salvelinus rossii Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 60 (in checklist, Green-
land, Boothia Penin., refs.).
Salvelinus alpinus forma stagnalis Pfaff, Rep. 5th Tuure Exped. (1921-1924), Zool. 2 (7), 1937: 1-3, 5, 6
(abund., Gore Bay, Fury and Hecla Strait, Canada; var. in length — maxillaries, pectorals; dentition).
Arctic Trout, Lowe, Rep. Dominion Exped. ... Neprunz (1903-1904), 1906: 11 (abund., Cumberland Gulf,
Baffin I.).
Synonyms for Freshwater Populations of Greenland and America westward to the International Boundary be-
tween Canada and Alaska:
Salmo stagnalis Fabricius, Fauna Groenl., 1780: 175 (detail. descr., remote mountain waters, not descending
to sea, Greenland).
Salmo rivalis Fabricius, Fauna Groenl., 1780: 176 (brief descr., small streams, stagnant waters, not descending,
Greenland).
Salmo alipes Richardson iz Ross, Narr. Second Voy. ... North-west Passage, Append., Nat. Hist., Fish., 1835:
tvuI (brief descr., abund., Boothia Penin., Arctic Canada); Fauna Boreal.-Amer., 3, 1836: 169, pl. 81
(descr., color. ill., streams trib. to Prince Regent Inlet, Boothia Penin., Arctic Canada); Giinther, Proc.
zool. Soc. London, 1877: 476 (lakes near Discovery Bay, Grinnell Land, Arctic Canada).
Salmo nitidus Richardson iz Ross, Narr. Second Voy... . North-west Passage, Append., Nat. Hist., Fish., 1835:
ty (brief descr., in lake, Boothia Penin., Arctic Canada); Fauna Boreal.-Amer., 3, 1836: 171, pl. 82,
fig. 1, pl. 86, fig. 2 (descr., color, color. ill., in lake, Boothia Penin., Arctic Canada).
Salmo oguassa Girard, Proc. Boston Soc. nat. Hist., 4, 1854: 262 (type local., Oquassa or Oquossoc Lake; Range-
ley Lakes region, Maine).
Salmo arcturus Giinther, Proc. zool. Soc. London, 1877: 294, pl. 32 (descr., color. ill., Discovery Bay, Grinnell
Land, Arctic Canada, near 87°30'N in fresh water).
Salmo naresii Giinther, Proc. zool. Soc. London, 1877: 476 (descr., color, Discovery Bay, Arctic Canada).
Salvelinus aureolus Bean, Proc. U.S. nat. Mus. (1887), ro, 1888: 628 (descr., meas., Sunapee Lake, New
Hampshire).
Salmo (Salvelinus) marstoni Garman, Science, July 14, 1893: 23 (descr., color, type local. Lac de Marbré,
Ottawa, Quebec; widespread, lakes of n. Quebec).
Salvelinus alpinus Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 508 (descr., cold lakes, mountain
streams, n. Europe and ne. Amer.; lists probable synon.).
Salvelinus alpinus alipes Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 509 (descr., refs., lakes,
Greenland, Boothia Penin.).
Salvelinus alpinus arcturus Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 510 (descr. by ref. to
Giinther, 1877, see above; Victoria Lake, Floeberg beach, Discovery Bay, Grinnell Land, Arctic
Canada).
Salvelinus alpinus aureolus Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 511 (descr., discus., refs.,
lakes, n. New Hampshire and Maine).
Salvelinus oquassa Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 514 (descr., refs., Rangeley Lakes,
Maine).
Salvelinus oguassa naresti Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 515 (descr., refs., lakes,
Arctic Canada).
Salvelinus alpinus marstoni Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 515 (descr., ref., Lac
de Marbré, Quebec).
Salvelinus stagnalis Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 60 (in checklist,
refs., Greenland, Boothia and adj. regions).
Salvelinus arcturus Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 61 (in checklist,
ref., Arctic Amer., 82°34'N).
Salvelinus naresii Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 61 (in checklist, lakes,
Discovery Bay, Cumberland Gulf, Arctic Canada, ref.).
Salvelinus alpinus complex, Walters, Bull. Amer. Mus. nat. Hist., 106 (5), 1955: 274-275 (list. freshw. locals.,
Arctic Alaska).
Fishes of the Western North Atlantic 525
Synonyms for the a/pinus species complex of the Pacific Coast of North America westward to Japan:
Salmo malma Walbaum, P. Artedi Genera Pisc., Ichthyol., Emend., 3, 1792: 66 (Kamchatka, based on ma/ma
of Pennant).
Salmo curilus Pallas, Zoogr. Rosso-Asiat., 3, 1831: 351 (Kurile Is.).
Salmo callaris Pallas, Zoogr. Rosso-Asiat., 3, 1831: 353 (Bering Sea).
Salmo laevigatus Pallas, Zoogr. Rosso-Asiat., 3, 1831: 385 (Kurile Is.).
Salmo nummifer Cuvier and Valenciennes, Hist. Nat. Poiss., 27, 1848: 365 (Kamchatka).
Salmo erythrorynchos Cuvier and Valenciennes, Hist. Nat. Poiss., 27, 1848: 366 (Kamchatka).
Salmo spectabilis Girard, Proc. Acad. nat. Sci. Philad. (1856), 8, 1857: 218 (Fort Dalles, Oregon).
Salmo bairdii Suckley, Ann. N.Y. Lyc., 7, 1861: 309 (Flat Head R. syst., Montana).
Salmo parkei Suckley, Ann. N.Y. Lyc., 7, 1861: 309 (Kootenay R., B.C., Canada).
Salmo lordii Giinther, Cat. Fish. Brit. Mus., 6, 1866: 148 (Skagit R., Alaska).
Salmo pluvius Hillgendorf, Mitt. dtsch. Ges. Ostasiens, rz, 1876: 26 (Japan).
Salvelinus spectabilis Jordan, Proc. U.S. nat. Mus. (1878), I, 1879: 79 (refs., in synop.).
Salvelinus bairdii Jordan, Proc. U.S. nat. Mus. (1878), 2, 1879: 82 (in synop.).
Salvelinus malma Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1882: 319 (in synop.); also many subsequent
authors.
Salvelinus fontinalis (Mitchill) 1815
Brook Trout, Salter, Squaretail Trout, Sea Trout, Salmon Trout, White Sea Trout
Figure 128
Study Material. 1. Sea Run: A total of 33 specimens: 11,8 46-150mm TL,
from the intertidal zone almost 200 feet below the high-water mark, near White Brook
in the vicinity of Jonesport, Maine; 6, about 222-262 mm TL, 209-247 mm SL,
from the Wilmot River estuary, south shore of Prince Edward Island; 16, about
156-380 mm TL, from near the mouth of the Little River Codroy estuary, the west
coast of Newfoundland.
Il. Fresh Water: about 300 specimens, large and small, from Labrador, Island of
Anticosti, Nova Scotia, New Brunswick, Maine, New Hampshire, Massachusetts,
New York, Pennsylvania, North Carolina, Georgia, Lake Superior, Wisconsin, North
Dakota (introduced), and California (introduced); MCZ.
Distinctive Characters. The only fish for which sea-run fontinalis is likely to be
mistaken are young Sa/mo salar (Atlantic salmon, p. 460); Salmo gairdneri (rainbow
trout, p. 499), an introduced species from the west that seems to have established a
sea-run population in the east; Sa/mo trutta (brown trout, p. 498), a second widely in-
troduced species (from Europe) that is known to run down to salt water in New-
foundland;18 or Sa/velinus alpinus (Arctic charr, p. 507). And there is not much danger
of confusion in any of these cases. Thus, the dark vermiculated markings on the dorsal
and caudal fins of fontinalis and the conspicuously blunter head and broader pectorals
13. Received through the courtesy of Keith A. Havey, Maine Department of Fisheries and Game.
14. Received through the kindness of the late A. H. Leim, and Marden Smith, Fisheries Research Board of Canada.
15. Received through the kindness of W. Templeman and A. R. Murray, Fisheries Research Board of Canada.
16. A few summers ago we saw (and handled) a brown trout that was found on the beach at Cohasset, Massachusetts.
And a 6-7 pound fish, caught in Witless Bay, coast of eastern Newfoundland, in 1935, was identified as such by
Harold Thompson (Frost, 24: 13).
526 Memoir Sears Foundation for Marine Research
(cf. Figs. 128, 125) are enough to set it apart at a glance from a/pinus. Its minute
scales (large and conspicuous in Sa/mo salar) and the total lack of black markings on
its sides distinguish it from S. salar of any size. Reliable field marks that distinguish
it from both Salmo gairdneri and S.trutta are the minuteness of its scales and the re-
striction of the teeth on the roof of its mouth to a cluster in front of the vomer; in both
Salmo gairdneri and S. trutta the teeth extend in a row rearward along the shaft of the
bone. There should be no danger of confusing fontinalis with Cristivomer namaycush
(lake trout), which has been reported occasionally as taken in brackish water in
Hudson Bay, Ungava Bay, and along the coast of northeastern Labrador (77: 275;
Figure 128. Sa/velinus fontinalis, 236 mm, Little Codroy River, Newfoundland, MCZ 40006.
3: 295), for the caudal fin of C. namaycush is much more deeply forked than that of
fontinalis, its head is noticeably narrower, and it has many more gastric caeca.
Description of Sea-run Specimens. Based on six fish, about 223 to about 262 mm
TL and 209-247 mm (about 8.25 to about 9.75 in.) SL, from the Wilmot River
Estuary, on the south coast of Prince Edward Island (p. 525), and on other available
information.
Trunk fusiform, its maximum thickness about 14-16 °/, of SL, its maximum
depth about 19-23 °/,; the depth of trunk exclusive of head about 1.2-1.4 times
greater than maximum thickness, the precise relationship depending on the relative
fatness of a particular specimen; the dorsal profile weakly convex, the ventral profile
anterior to anal fin only slightly more so, except in especially well-fed specimens
or in sexually ripe individuals in which the sex organs have enlarged. Depth of
CauUDAL PEDUNCLE about 10, about twice its thickness. ScaLes present on body and
tail sectors everywhere; head and fins naked; the scales rounded, entirely enclosed in
skin, and so minute that they are likely to be overlooked; about 230 along the LaTErat
LINE (reported).
Heap about 23-25 °/, of SL, its dorsal profile weakly concave between perpen-
diculars at upper end of gill opening and posterior edge of eye in some specimens,
Fishes of the Western North Atlantic R27
but not appreciably so in others, thence increasingly convex forward. SNouT 24—30°/,
of length of head, bluntly rounded. Tip of Lower jaw almost even with tip of Upper
Jaw, or extending very slightly beyond it; rear end of upper jaw reaching to a per-
pendicular that varies in position from only slightly behind rear edge of eye to behind
eye by a distance about as long as eye or a little longer than eye. Eve 16-19 °/ of
head, but much larger relatively in fingerlings, being 24 °/, in one 49 mm SL, its rear
edge about 24—25°/, of distance from the snout rearward to rear edge of gill cover.
PosrorsiTat part of head 53-59 °/) of head. Moutu moderately oblique. GiLL RAKERS
4-8 on lower limb of first arch; 7-9 on upper limb (reported). BrancurosTEGAL rays
g—12 (reported).
TeretH. Upper and lower jaw bones (premaxillaries, maxillaries, palatines, and
dentary portion of mandibles) each with a single row of sharp, slightly recurved teeth
of moderate size; also a patch on anterior end (head) of vomer on the midroof of mouth
(none, however, rearward along shaft of vomer); a single row of smaller teeth around
tip of tongue; also a patch of sharp but minute teeth in the pharyngeal region below,
and two such patches above, side by side. No teeth on bones at base of tongue.
Rayed Dorsat fin rhomboid and with angular or slightly blunted corners, its
base 50-60 °/, of length of head, its origin about midway (0.47—-0.50) from tip of
snout to caudal base, its longest ray 0.9—1.0 times as long as the base. ADIPOSE DORSAL
fin shaped as in Fig. 128, the midpoint of its base about 60°/, of distance from rear
end of rayed dorsal base to origin of upper side of caudal. Caupat with upper and
lower corners abruptly rounded, the rear contour weakly concave, varying from double-
sinuous to uniformly arcuate, its breadth when widespread about twice the length of
upper and lower margins. Anat rhomboid, its anterior corner usually bluntly rounded,
its posterior corner angular, its outer margin weakly concave, at least in most speci-
mens (Fig. 128), its base 0.7—0.8 as long as dorsal base, its origin about midway
between origin of lower side of caudal and a perpendicular at dorsal origin, its longest
ray about as long as longest dorsal ray. Petvics with midpoint of base at a perpen-
dicular about under midpoint of dorsal base, the longest ray about as long as longest
dorsal ray; the fleshy appendage beside base of pelvics conspicuous, about as long
as eye. Pecrorats with shape, when well spread, as shown in Fig. 128, the origin a
little anterior to rear edge of gill cover, the longest ray a little shorter than the longest
dorsal ray.
Fin rays: dorsal 11-14 reported, anal 9-12 for Moser River Salters (Table xz).
VERTEBRAE 58—62 reported; average 59.5. PyLoric cagca, average 38.4 reported.
Color. On fontinalis from fresh water, the back and upper part of the sides are some
shade of olive, conspicuously marbled with paler and darker markings; the lower part
of the sides is marked with many pale yellowish spots interspersed with a smaller number
of vermillion dots encircled (typically) with blue. The lower surface varies from grayish
blue through shades of pale orange and a fleshy color (reddening at spawning time) with
a narrow white midline. The dorsal fin is a paler olive than the back, with coarse blackish
vermiculations. The caudal is a darker olive (reddening more or less at spawning
528 Memoir Sears Foundation for Marine Research
time) with darker wavy crossbars, especially on the upper and lower corners. The
pectorals, pelvics, and anal are some shade of pink, orange, or crimson, the pec-
torals and pelvics dusky; the first ray of the pectorals, pelvics, and anal is white
or cream-colored, conspicuously edged rearward with black. Fingerlings up to 5 or
6 inches or so are more or less conspicuously marked on the sides with a series of
7-11 (av. 9) dark, vaguely outlined
Table XI. Dorsal and Anal Rays of 8. fontinalis* crossbars or blotches, known as
INo# ofirays yy ts1.vetiacers OPRLOMATTADS (12 e0T 31) ex4! “parr marks,”’
Dorsal, no. of specimens. — — 40 120 50 I The young Brook Trout that
Anal, no. of specimens... I 47 123 38 — — _ are destined to move out into salt
* After Wilder, 78: 186, tab. 8. water cannot be distinguished at first
by their color from those that are
to remain in fresh water. This continues true in some localities until after they have
actually left the stream mouths, for example on the coast of eastern Maine (see Study
Material, p. 525). In other localities, however, as in the Moser River, Nova Scotia,
the sides of the future Salters, now known as “smolts” in conformance with salmon
nomenclature, have turned partially silvery, through the deposition of guanin crystals,
by the time they are ready to migrate downstream, although they still show their parr
marks. It seems that this difference in color between different populations is associated
with the age at which they move out into brackish water, for while some of the Maine
Salters (p. 525) do this as early as their first summer, and many by their second sum-
mer, the smallest sea-run Moser River fish were in their third summer. In any case,
the development of the Salter coloration evidently is a reversible process, for Wilder
found that Moser River smolts whose sides had already turned partially silvery lost
the silver if they were retained in fresh water (78: 173).
After the Brook Trout move out into brackish or salt water, their sides tend to
become increasingly silvery, the greenish-blue-to-bottle-green marbling of their upper
parts becomes obscured, their bellies become paler, and their pectoral, pelvic, and
anal fins lose color. In extreme cases, their sides may turn as silvery as those of Atlantic
salmon, with the pale yellow spots and crimson dots showing only faintly and the
marblings hardly visible. Their pelvic and anal fins are white, perhaps faintly tinged
with pink, and show no trace whatsoever of the black edging on the first ray that is
so conspicuous on freshwater fish. But the dark wavy markings on the dorsal fin and
on the upper part of the caudal continue to be conspicuous.
The published descriptions of the color of sea-run fontinalis have in most cases
been based on specimens of this sort, but the individual Salters may vary widely in
color, even though caught side by side. One of the Prince Edward Island specimens
listed above is pale (Fig. 128), although its sides, after preservation in formalin, now
show no trace of silver. On another, however, the back and the upper part of the
sides are much darker, the marblings, pale spots, and crimson dots are much more
evident, as are the dark markings on the dorsal and caudal fins, the lower part of
the sides is gray, and the first ray of the pectoral, pelvic, and anal fins is outlined as
Fishes of the Western North Atlantic 529
conspicuously rearward with black as it is in freshwater specimens. The four other
Prince Edward Island specimens are intermediate in coloration between these extremes.
But none of the Newfoundland specimens (see Study Material, p. §25) approaches the
extreme sea-run coloration, though they were taken only one mile upstream from where
the river empties into the open Gulf of St. Lawrence.”
It seems, as Smith remarked a century and a quarter ago (60: 354, 355), that the
most silvery of the Salters are the ones that pass their marine sojourn in the saltiest water
and are taken soon after they leave the stream mouths, whereas the least silvery are
those that remain within the estuaries, where the water is more or less brackish.
When the Salters re-enter fresh water, the back and the upper sides soon darken,
the silver of the sides fades out, the pale spots on the sides become more intensely
orange, the pectoral, pelvic, and anal fins gradually assume the white-black and orange-
to-reddish pattern typical of the freshwater fish, the pure white on the lower part of the
sides develops a pinkish band in the males, and the belly tends to become gray. Brook
Trout fresh-run from the estuary that were contained in pens in the Moser River under-
went this alteration between July 15 and September 10, by which date “‘it was impos-
sible to distinguish sea and fresh water trout by color” (Wilder, 78: 174). And the
Salters of Cape Cod may lose their sea coloring within 10-14 days after they re-enter
fresh water (9: 8).
Size. Brook Trout are about 14 or 15 mm (0.56 in.) long when hatched, and there
is no reason to suppose that fontinalis fry destined to run down into salt water differ
in size at hatching from those that are destined to spend their life in fresh water.
Two or three years later, when they migrate downstream, the Salters average about
175mm in length (6.9 in.) in Nova Scotian waters, and about 178 mm (7 in.) on the
coast of western Newfoundland, a growth rate that agrees closely with 6.5 inches at
two years and about 8 inches at three years as reported for hatchery fish (perhaps
more abundantly fed) at Caledonia Springs Hatchery, New York (29: 312).
The maximum size to which foxtinalis grow in fresh water varies from one stream
to another and from one pond or lake to another, even within short distances. A 1.5-
pound fish, generally speaking, is a large one in any small water, and a 5-pound fish
is a large one in most of the larger waters. In some northern streams, however, and in
northern lakes, many of them grow far larger than that. Goode et a/., for example, men-
tion one of 11 pounds that was received by Louis Agassiz from the Androscoggin
River in northwestern Maine in 1860, and a Rangeley Lakes fish that weighed 10
pounds after it had been in captivity for three years (26: 499, 500). Kendall found
records of more than 60 fish heavier than 9 pounds, 15 of 10-11 pounds, and 4
of 12—12.5 pounds, which had been taken at one time or another in the Rangeley Lakes,
Maine (45: 90-95); he also reported one of 11 pounds for Belgrade Lake, Maine, and
one of ten pounds for Square Lake, Maine. The record (weighed) weight is 14 pounds
8 ounces, for one caught July 1916 by J. W. Cook in the Nipigon River, tributary
to the north shore of Lake Superior, where Hewitt (34: 170) also took one 26 inches
17. Information contributed by A. R. Murray, Fisheries Research Board of Canada. 18. Ibid.
530 Memoir Sears Foundation for Marine Research
long having a girth of 20 inches; this fish must have weighed close to 13 pounds, as
calculated by the accepted formula W = (L.G$)/1000, where W equals weight in ounces,
L the length in inches from eye to base of caudal fin, and G? the cube of the girth in
inches. Dunbar and Hildebrand’s report (16: 95, 96) of “speckled trout” weighing up
to ten pounds, taken near Fort Chimo, northern Labrador (Ungava Bay), shows fur-
thermore that fontinalis may grow equally large right up to the northernmost limit of
their geographic range.
Sea-run fontinalis in general grow larger than those that remain in fresh water;
not very much larger, however, except in situations where the growth rate of the fresh-
water fish is low. On the south shore of Cape Cod, for example, where few of the native
fish that remain in fresh water are heavier than 0.5-—0.75 pound, the largest sea-run
fish reported by Smith (in 1833) among many hundreds were one of 2.5 pounds, two
that weighed close to 3 pounds each, and three that together weighed 8.25 pounds
(60: 353). The largest, taken at the mouth of a privately owned brook tributary to
Buzzards Bay, where a continuous record has been kept since 1870, weighed 3.75
pounds, and the largest recently taken there was 2.75 pounds (information contributed
by C. P. Lyman). Rumors of 5-pound Salters on Cape Cod or on Long Island, New
York, seem not to have been supported by concrete evidence.
The average for the largest ten Salters from the Moser River, Nova Scotia, measured
by Wilder (78: 189, tab. 10, 191), was close to 15 inches (381 mm) and 13.4 ounces
(381 g), whereas the average for the largest ten freshwater specimens was only 11.12
inches (282 mm) and 12 ounces (340 g); and one Salter of 18.7 inches (about 476 mm),
which must have weighed close to three pounds, has been reported for the Moser River
(9: tab. 1). The heaviest that has been reported to me for the waters of Nova Scotia
was eight pounds.!® On the other hand, no sea-run fish longer than ten inches were
seen off the mouth of a small brook in eastern Maine, where a special study of them
was carried out by Charles F. Ritzi,?° though conditions in the marine environment
there would seem to be favorable for growth to a large size.
Available evidence suggests that the Salters may average somewhat larger along
the southern side of the Gulf of St. Lawrence than in Nova Scotian waters, for Goode
et al. described them as usually weighing ‘‘two and one-half pounds,” adding that “‘they
are not seldom caught as heavy as six or eight” (26: 499). This accords with Perley’s
description of catching five-pounders himself on the north shore of Prince Edward
Island (57: 132) and with W. P. Templeman’s report of Sa/ve/inus weighing up to eight
pounds in the Bay of Chaleur (letter). A Brook Trout weighing six pounds was taken in
Grand River Codroy, west coast of Newfoundland, by Sir Bryan Leighton in 1904
(48: 81), and fish of seven or eight pounds have been reported for West Brook, some
15 miles north of Bonne Bay, Newfoundland (50: 24), if these were not Arctic charr
(p. 521). However, catches of 33 fish averaging 3.3 pounds off the mouth of Grandys
Brook near the Bureo Islands on the coast of southwestern Newfoundland (48: 68)
rg. Unsigned statement received in reply to an inquiry as to the status of sea-run fontinalis in Nova Scotian waters.
20. Information contributed by Charles F. Ritzi.
Fishes of the Western North Atlantic Bow
Table XII. Average Total Lengths; of Sea-run and Freshwater Brook Trout of
the Moser River in Five Seasons, May 18—August 4.* In Parentheses, Number of
Fish Examined
Seasons 3rd 4th sth 6th 7th
Sea-run trout (mm)......... 213 262 290 354 420
Sea-run trout (in.).........-. 8 3/8 105/16 11 7/16 14 16 9/16
(94) (103) (307) (7) (1)
Freshwater trout (mm)...... 164 189 237 296
Freshwater trout (in.)....... 6 7 9/16 9 5/6 II 3/16
(51) (123) (46) (2)
+ Standard lengths converted to total lengths by the addition of 6°/,, a conversion factor determined from
the Prince Edward Island Salters listed (p. 525).
* After Wilder, 78: 191, tab. rr.
and of many of 3—-3.5 pounds in Newman’s Sound on the eastern coast (34: 4) are
more nearly representative of the largest Salters that are commonly taken in New-
foundland waters, where the great majority are nearer one pound.
Salters ‘‘up to 7 and 10 pounds” are often taken along the Atlantic coast of Labra-
dor (48: 68), but some of these “outsize” fish may have been Arctic charr (p. 521).
At any rate, it seems that the Labrador Salters do not average any larger than the Nova
Scotian fish, for those caught along the southern section of Labrador “‘usually run
from one pound to four pounds in weight.’ Those reported as being taken near
Nain (about 56°30’N) weigh up to only about two pounds (72: 130), while Arctic
charr there weigh up to 8-10 pounds or more. Even the largest sea-run fontinalis that
have ever been weighed anywhere, or reliably reported, have fallen far short of the
largest of the freshwater fontinalis from various Maine and Canadian waters (p. 529).
Rate of Growth. In the Mashpee River, on the south shore of Cape Cod, the Salters
average about 7.4 inches (188 mm) TL in their second season, about 8.8 inches
(223 mm) in their third season, about 9.9 inches (249 mm) in their fourth season,
and about 12.3 inches (312 mm) in their fifth (9: tab. 1). The average lengths at cor-
responding ages for Salters of the Moser River, Nova Scotia, are about the same as these,
as appears from Table xu, based on Wilder’s observations (78: 191, tab. 11); and a
yearly increment of 1.4 inches, reported for the fish that sojourned the longest in the
sea off the coast of eastern Maine, is of this same order of magnitude. Table x11 tends
to support the general view that the Salters grow somewhat faster than the freshwater
fontinalis in the same stream.
Survival Rate. Information about the rate of survival for sea-going fontinalis in
different localities shows that only 127 (10.4 °/)) of the 1,220 Salters marked in the
Moser River system were retaken in the traps there on their return from the sea, with
the percentage greater for those larger than 220 mm (31-35 °/,) than for smolts of
180-205 mm (8.5 °/,); and it seems that the yearly mortality for sea-run fontinalis
is about the same on the coast of eastern Maine, where few survive for more than three
21. Information from Newfoundland Ranger S. M. Christian.
34
532 Memoir Sears Foundation for Marine Research
growing seasons. Wilder’s data show that the survival rate declines abruptly for both
Salters and nonmigratory fish of the Moser after they pass their fourth season and that
very few of either survive for as long as six years (78), though it is known that an
occasional fontinalis may survive into its eighth year elsewhere (g: 14).
Spawning and Reproduction. The migratory populations of S. fontinalis, like the
nonmigratory fish, spawn exclusively in fresh water in autumn at temperatures ranging
from about 49°F (9.4°C) down to about 40°F (4.4°C). In general, spawning by fox-
tinalis takes place earlier in the season to the north than to the south, but the precise
spawning period at any particular locality is governed by the rapidity with which the
water cools with the onset of autumn rather than by the latitude. Brook Trout
spawn from mid-October until early December on Cape Cod (probably), from mid-
October on into December to the north in New Hampshire (26: 499), from mid-
October into November in the Rangeley region of Maine (45: 84), from late Oc-
tober through December into January (exceptionally into February) on Prince Ed-
ward Island (74: 356, 357), during October in the Moser River, Nova Scotia (75:
185), and from mid-October to middle or late November on the coast of eastern
Newfoundland (24: 8). In shallow, rocky streams, however, in the Maritime Provinces
of Canada, where the water chills very rapidly, they may commence to spawn as soon
as late September (74: 356, 357). Hence the spawning dates given by Vladykov for
the high-lying Laurentide Park lakes, north of Quebec City (68: 800), probably are not
applicable to the coastal streams along the north shore of the Gulf, where it is more
likely that foxtinalis spawn from mid-October to mid-December, as they do in southern
Ontario (57: 99). No information is available in this regard for the outer coast of
Labrador. Along the southern shoreline of Ungava Bay, however, “‘it is probable
that spawning takes place in September or October,” to judge from the state of the
gonads (Dunbar and Hildebrand, 16: 96).
S. fontinalis, like Salmo salar, spawn on bottoms of sand or gravel in shallow water,
in streams, ponds, or lakes, where either the current or the inflow from some spring
keeps the eggs clear of detritus. Here the females smooth out a shallow depression
(the so-called redd) into which the eggs settle with the milt from the attendant male,
and immediately after the spawning act, the eggs are covered with gravel. The eggs, aver-
aging about %/,,of an inch (about 5 mm) in diameter, do not hatch until the following
April or May, the precise date depending on the temperature of the water; 53°F
(11.7°C) is about the upper limit for their development (1g: 281-289); and the eggs
develop normally in water as cold as 35°F (1.7°C). The fry, about 0.5 inch (14 or
15 mm) long at hatching, carry a large yolk sac at first, as do species of all the Sal-
monidae; and they tend to remain close to the redd until they have grown to a length
of 1.5—2.0 inches (about 38-50 mm).
Some of the sea-going fontinalis of the Moser—probably this is true of the majority
—spawn in the autumn of the same year in which they make their first migration to the
sea. Others, like young Atlantic salmon (grilse), do not spawn until the second autumn
following their original descent to salt water.
Fishes of the Western North Atlantic 533
Nonmigratory fontinalis ordinarily spawn every year once they have reached sexual
maturity, but it seems that some of the Salters do not, for the upstream runs, at least
in the Moser River, Nova Scotia, include some large fish that show no signs either
of approaching sexual maturity or of having spawned during the previous autumn
(75: 179). It appears, too, that some of the mature Salters may occasionally remain
in fresh water throughout an entire twelve-month period (78: 195); hence it would
not be astonishing if some of them fail to run up into fresh water at all in some
years.
It appears that the majority of the Moser River Salters spawn three or four times
during their life and that a few may do so five or even six times, assuming that they
spawn every year (which some do not) and that they continue fecund throughout their
life (which they may not), all of which probably applies to populations of Salters
elsewhere.
Habitat and Migrations. In the Moser River, Nova Scotia, about 79 °/, of the
smolt carry out the initial downstream migration in the second spring when they
average about 170 mm TL, and the remainder in their third spring (75: 179); this
applies also to Brook Trout artificially stocked in Cape Cod waters (49: 10). On the
coast of eastern Maine, however, some move out into salt water in their first spring,
when they are only about 46-52 mm or 1.75-—2.0 inches long (see Study Material,
p- 525); Weed has reported similarly that in northern Labrador, in the vicinity of
Nain, “many go down to the edge of salt water on the tidal flats during their first or
second summer” (72: 130).
The migratory schedule of the sea-running /fonsina/is, after their initial descent to
brackish or salt water, consists of an upstream migration in late spring or summer by
fish destined to spawn that autumn as well as by smaller immature fish, and, after the
spawning season, of a downstream migration by smolts, immature larger fish, and
kelts (spent fish) to brackish or salt water, where the kelts recover condition during
the winter.
The upstream run by the fat fish takes place from late May through June on
Cape Cod, with a few entering in September and perhaps even as late as November.”
On the coast of eastern Maine, the principal run is from May until early August, with
some irregular movement to and fro in the autumn;?3 in Nova Scotian waters, as illus-
trated by the Moser River, the chief run is from about mid-June into August, as in
1939 when 93 °/, of the run there was in July (75: 178, 179).
The Salters appear along the shore during July in the Mira River district of Cape
Breton (Breck, 8: 97), and at about this same time they are running upstream in force
in the estuary of the Little River Codroy, Newfoundland.*4 But along the southern
sector of the outer Labrador coast (according to Blair, ftn. 26) they are moving
upstream in September. Large runs of fontinalis are recorded for July and August
22. Mullan (49: rr) has mentioned a probable instance of upstream migration in the Mashpee River in November.
23. Information contributed by Charles F. Ritzi.
24. Information contributed by A. R. Murray, Fisheries Research Board of Canada.
534 Memoir Sears Foundation for Marine Research
for Hamilton Inlet and Sandwich Bay on the outer coast of Labrador (3: 294), and the
seasonal schedule is about the same as this near Nain (about 56°30’N), where “brook
trout weighing up to almost two pounds are found in the sea from the time when the
ice goes out early in June” (Weed, 72: 130).
On Cape Cod, Massachusetts, the return migration downstream takes place soon
after the spawning season has ended; i. e. in November and December, perhaps com-
mencing even in October (7: 140; 6: 121; 9: 26); and seemingly the seasonal schedule
is similar to this in the western side of Hudson Bay where Doan has observed that
the spent fish apparently winter in the estuaries of the large rivers and then cruise
along the coast (73: 6).
On the coast of eastern Maine, however, and to the north in general, the great
majority of the kelts and smolts, with the immature fish that have run up during the
previous summer, spend the winter in fresh water. The principal downstream move-
ment by the Brook Trout takes place the following spring; for example, in the Moser
River, Nova Scotia, the smolts move down in April and May and the kelts do so until
well into June when the temperature of the water has risen to 64°—65°F or 18°C (76).
On the coast of western Newfoundland, as represented by the Little River Codroy,
the chief downstream run “‘begins in late April or early May, to cease around mid-
June,” although it is preceded by some movement “down the estuary as early as
January and February” (Weed, 72: 130). While they have been reported by P. S. Mar-
tin as being in salt water from November on along the north shore of Prince Edward
Island (as they are on Cape Cod also), they enter the sea around the middle of
May? in the southern section of Labrador; and it is not until early June when the
river ice opens that they are found in the sea in the vicinity of Nain, northern Labrador
(72: 130). Thus the annual sojourn in the marine environment (as determined by
tagging experiments) is usually 30-60 days on the coast of eastern Maine, with the
recorded maximum 106 days and the minimum 1-5 days; the average is 65 days
for the larger of the Moser River fish and 71 days for the smolts (76: 263). In
northern Labrador they are in salt water from early June to late summer, but precise
dates have not been given.
Since mature sea-running fish eat very little while they are in fresh water, the feed-
ing period throughout the northern part of their geographic range is thus condensed
within only a little more than two months out of the twelve. While they may sojourn
in salt water for as much as six months (November to June) in the southernmost part of
their range (Cape Cod), the period during which they feed actively probably is no
longer there than it is in Nova Scotia—perhaps not so long—for the surface temperature
in the bays and estuaries of Cape Cod where they winter usually continues below 40°F
from sometime in December until mid-April (23: figs. 4, 5 [100]), with the water
usually chilling to within a degree or two of the freezing point of salt water during
25. Information based on the results of tagging, contributed by A. R. Murray, Fisheries Research Board of
Canada.
26. Information from W. A. Blair, in charge of Salmon investigations for Newfoundland.
Fishes of the Western North Atlantic a5
January and February and with ice forming in greater or lesser amounts during the
coldest part of the winter.
Wanderings in Salt Water. The movements of the Salters after they have dropped
downstream to the head of tide vary widely from place to place. At the one extreme, as
was formerly illustrated along the southern shore of Cape Cod and is illustrated today
by the estuary of the Little River Codroy on the coast of western Newfoundland and by
the Wilmot River estuary on the south shore of Prince Edward Island, many Brook
Trout (perhaps a majority) may remain with the river influence during the entire interval
between their downstream and upstream runs. At the opposite extreme, they may leave
the mouths of the rivers, even within the first days after the ice has broken, so that
nets set a mile or two offshore will catch more of them than nets set near river mouths;
this is the case in the Moser River, Nova Scotia (75: 180), and off the southern part
of the outer coast of Labrador.?” We suspect that regional variation in the available
food supply rather than the saltiness of the water is the determining factor here, for
fontinalis in splendid condition are caught in distinctly brackish water as well as in
water of salinity characteristic of the open coast nearby.
In any case, sea-run Brook Trout tend to hold very close to the immediate coastline
and to water only a few feet deep. Along the south shore of Cape Cod, for example,
in the days when Salters were caught there in large numbers, they were taken chiefly
in the salt creeks and in the estuaries into which these open, not far out in the bays.
Off the Moser River, Nova Scotia, they are seen similarly ‘‘in schools in water five
to ten feet deep (1.5 to 3 m) around the inner islands and also around some of the small
private wharves ... when lobster pots are brought in and cleaned, or when fish are
being dressed,” with those of different sizes tending to school separately (White, 77:
471, 472). They are also found indifferently along sandy beaches, over muddy bottom,
and around rocky shores.
The only precise information as to how far offshore they are to be found regularly
is that they are taken in drift nets set for Atlantic salmon over the shoals in the Bay
of Chaleur,** and that they are caught on hook-and-line as much as seven or eight miles
offshore, in the western side of Hudson Bay off the Nelson River (13: 6). How much
farther out to sea they may stray occasionally is anyone’s guess.
The most extensive journey along shore that has been reported so far for any of
the many Brook Trout marked either in the Moser River, Nova Scotia (77: 471), or
on the south shore of Cape Cod has been eight miles (49: 10); in both cases the fish
in question were recaptured in streams other than the ones where they had been
marked. For Salters to reach Woods Hole on the southern coast of Massachusetts, as
they sometimes do, does not require a swim of more than seven or eight miles from
the nearest home stream. On the southern part of the outer coast of Labrador, however,
fontinalis have been caught in nets set on headlands ten miles away from the nearest
river where Salters have been taken.?”? Furthermore, during the period when Salters
27. Information from Newfoundland Ranger S. M. Christian.
28. Information contributed by W. P. Templeman.
536 Memoir Sears Foundation for Marine Research
would ordinarily remain in salt water, or at least in brackish water, they may return
temporarily to the stream mouth in pursuit of prey. Thus they re-enter the Fox
Island River, Newfoundland, in June and July, apparently to feed on the smelt
that spawn there, and return to the sea afterwards with the spent smelt;*® also, Brook
Trout that have wintered in the sea revisit the Dunk River on the north shore of
Prince Edward Island in April, or a little afterwards, when the smelt enter the brooks
to spawn.°°
Relation to Temperature. S. fontinalis feed most actively at temperatures of about
45°-65°F (7.2°-18.3°C); they do not thrive in water warmer than 68°F (20°C);
and they cannot long survive a temperature higher than about 70°F (21.1°C) unless
the water is especially well aerated, when they can withstand 75°F (23.9°C) for a short
time. At the other extreme, they become inactive and refuse food if the temperature
falls much below about 38°F (2°C), which accords with White’s observation that the
annual downstream movement of fontinalis in the Moser River system is interrupted
if the temperature of the water temporarily falls that low (76: 260, fig. 2).
Food. The diet of the fingerling Salters is the same while they are in fresh water as
that of their nonmigratory relatives; i. e. chiefly the aquatic stages of various insects
such as midges (chironomids), caddis flies (Trichoptera), May flies (Ephemeridae), and
black flies (Simulidae). They also eat snails and freshwater gammarid crustaceans if
these are available, besides terrestrial insects that may fall into the water.
During their sojourn in brackish or salt water they feed chiefly on small fishes,
of whatever kinds are the most easily available locally, or on small crustaceans. Ever-
mann, for example, found Brook Trout in the tidal waters of Casco Bay, Maine, “gorged
with young eels of the translucent stage” (20: 105). The diet list reported for Salters of
the Moser River includes young eels, young hake (Urophycis), young searavens (Hemi-
tripterus), killifishes (Fundulus), rock eels (Pholis), sticklebacks (Gasterosteus), and ale-
wife fry (Pomolobus); and smelts (Osmerus) as well as silversides (Menidia) have been
reported as being eaten by them at Prince Edward Island (75: 184; 77: 472). They
feed greedily on decapod shrimps, amphipods, and isopods, which, together with
Fundulus, are their chief food in the Cape Cod region. Weed (72: 130) has written
that Salters caught in the bays of Labrador had their stomachs crammed with beach
fleas (Gammarus). Others had been feeding on the tiny swimming larvae of the rock
crab (Cancer). And they have been found gorged with sea worms (Nereis) at Prince
Edward Island.
They eat very little, if anything, however, after they re-enter fresh water, probably
because of the scarcity there of prey of the sizes to which they have become accustomed
in salt water. And it is likely that such of the spent fish as winter in fresh water (p. 534)
continue on starvation rations, if they feed at all, until they move down again to the
head of tide in the spring.
Predators and Parasites. The smaller Salters are preyed upon, no doubt, by the
larger predaceous fishes, such as eels, striped bass (Roccus saxatilis), pollock (Pollachius
29. Information contributed by W. P. Templeman. 30. Information contributed by P. S. Martin.
Fishes of the Western North Atlantic 537
virens), silver hake (Merluccius bilinearis), and perhaps spiny dogfish (Squalus acanthias),
and by cormorants and gannets. The larger Salters are so active that they are not likely
to suffer seriously from these predators, but seals doubtless take a toll of them. During
their sojourn in fresh water they suffer from attacks by herons and eels, as shown by
the scars sometimes seen on their sides (75: 182); mink and otter are also poten-
tial enemies.
N. C. White has contributed the information that when the Salters re-enter the
streams they are heavily infested with sea lice (parasitic copepods, genus Lepeophtheirus)
similar to those that infest fresh-run Atlantic salmon, but these drop off soon after the
fish have entered fresh water.
Variations. Anglers have long been aware that the populations of fontinalis in dif-
ferent lakes, ponds, and streams differ widely in color, relative size of head, depth of
body, size at maturity, and maximum size of growth. Hatchery experience, however,
has proven that the rate of growth depends primarily on the supply of food available
to each individual fish combined with the yearly duration of the season when the water
is neither too cold nor too warm for them to feed actively.
Fat fontinalis are clearly deeper-bodied than lean ones, and Ricker has pointed
out that the maximum size to which fontinalis grow seems correlated with the size of
the body of water in which they live, and with the presence of suitable large food,
i.e. fish or crayfish (57: 73). These generalizations find their expression in the large
sizes to which they tend to grow in large streams and lakes in the northern part of
their range, contrasted with the stunted but overplentiful populations that inhabit
many of the smaller mountain streams of southern North Carolina and northern Georgia,
where fontinalis “rarely pass the dimensions of fingerlings” (Jordan, 47: 112) though
they are self-sustaining. Also it seems from casual firsthand observation that the heads
average larger, relatively, among the slow-growing populations than among fast-
growing fish, quite apart from sexual dimorphism.
In general, fontinalis average paler when living over a pale sandy bottom in brightly
illuminated waters than over a dark bottom in shady situations, but their nuptial
coloration usually is more brilliant in the second case than in the first. But this rule is
not invariable (for an interesting exception among Labrador fish, see Weed, 72: 131).
Brook Trout that are dark when fresh-caught soon fade after death. In short, it seems
unlikely that any of the local populations of fotinalis represent recognizable subspecies,
except perhaps those in Dublin and Center ponds and Christmas Lake, New Hamp- .
shire, which were made the basis of a separate species (agassizii) by Garman (25: 78).
The fontinalis that are taken in salt water in estuaries or at sea differ so widely
in their pale coloration and silvery sides from the freshwater Brook Trout that they
have been mistaken by various authors (60: 141, 1423 5I: 131, 1323 4: 206; 53:
2555) for the sea-run form of Sa/mo trutta, the brown trout (p. 498); and those from
Hudson Bay, Labrador, and from Newfoundland were described by Suckley (64: 310)
as a separate species, Salmo hudsonicus, which has been accepted as a subspecies by
31. For historical account, diagnostic characters, and excellent colored figures of agassizit, see Kendall (45: 56-68, pl.6).
538 Memoir Sears Foundation for Marine Research
Hubbs (36: 17; 37: 258). Anyone, for that matter, who has compared them is likely
to have noticed that the adults average thicker but less deep among the sea-run /fon-
tinalis than among freshwater fish of a corresponding size (78: 181, tab. 3, 185,
187). Due to their stoutness, Salters average somewhat heavier than freshwater fish
at a given length; Smith, for example, remarked in 1833 that while freshwater fontinalis
of one pound are about 14 inches long if in good condition, Salters may weigh one
pound when only about 11 inches long (60: 350). Correspondingly, a Newfoundland
Salter of 14 inches TL (see Study Material, p. §25) weighs about 2.25 pounds after
preservation in formalin. Wilder, however, has shown that these differences are not
statistically significant for the females, and that the differences in shape of body and
size of head are “‘less than those between progeny of one pair of trout raised at different
temperatures, and much less than the differences found among four widely separated
populations of freshwater trout.” The readiness, too, with which hatchery fish of
strictly freshwater parentage assume the sea-running habit “refutes any claims for a
genetic difference between anadromous and freshwater brook trout” (4g: 8-11). Thus
Jordan’s dictum* of 1905, that the seagoing fontinalis are merely sea-run brook trout,
has been verified conclusively.
Numerical Abundance. The only precise evidence available as to how many Salters
a given stream may harbor nowadays is as follows: 1,220 taken in 1939 in one of the
counting traps in the Moser River, Nova Scotia, on their descent to salt water (75);
between 500 and 1,000 smolts counted on their way downstream from late April
through May in the Little River Codroy, coast of western Newfoundland;3* and 200—
400 larger fish taken there from July through August on their return.
The evidence at hand does not suggest that any general alteration has taken place
one way or the other in the abundance of Salters in Nova Scotian waters, or to the
north. In the Cape Cod region, however, Salters are far less numerous now than they
were formerly, if, indeed, any of the undiluted native strain still exists there. A con-
crete example of their early abundance is that in April 1829 two anglers, in five
days, caught 296 (averaging about 0.5 pound) at one of the well-known fishing points
in Waquoit Bay (Smith, 60: 365%). In Mullan’s opinion this decline has been due prima-
rily to the effects of the Cape Cod Canal, the cranberry industry along the streams,
the ditching of marshes for mosquito control, and rural development in general in
reducing the area of suitable freshwater habitat (49: 21, 22). Catches at the mouth
of one privately owned brook tributary to Buzzards Bay suggest furthermore that the
disappearance of the eelgrass (Zostera marina) in 1932-1933 was an additional
calamity for the local Salters; for while the average number of sizeable fish taken there
yearly was about 116 for the period 1928-1932, the largest catch for any subsequent
year down to 1954 was only 45.
Results of Stocking with Hatchery Fish. The localities of recapture of 92,100 marked
32. Information contributed by Charles F. Ritzi.
33- Information contributed by A. R. Murray, Fisheries Research Board of Canada.
34. Misprinted as p. 265. 35- Information from C. P. Lyman.
Fishes of the Western Ni orth Atlantic 539
fish that were released by the Division of Fisheries and Game of Massachusetts in
Cape Cod streams in 1949-1956 have shown that hatchery fish of freshwater par-
entage readily adopt the sea-running habit and wander as far afield as wild Salters
are known to do anywhere (49: 8-11); tagging experiments have yielded similar
evidence for the coast of eastern Maine.** ’
Relation to Man, Sea-run Brook Trout have long been favorites with anglers, for
they take a brilliant artificial fly readily, put up a strong resistance, and are excellent
fish on the table, firm and pink-meated and of delicious flavor. Waquoit and Poponesset
bays, for example, on Cape Cod, with their tributary streams, were famous Brook
Trout waters more than a century anda quarter ago (60: 349-351); and anglers as early
as 1851 resorted ‘‘annually during the month of June for sea trout’ to the estuary
of the Philip River on the Northumberland Strait shore of Nova Scotia (Perley, 51:
131). Brook Trout fishing, indeed, continues a favorite sport today wherever they are
plentiful in localities that are easily accessible and where they are not so overshadowed
by the Atlantic salmon that they are looked upon as a nuisance rather than as a sporting
asset. They are netted, also, in some numbers for local consumption wherever there are
enough of them to make this worthwhile. But they are not plentiful enough anywhere
to be of commercial importance.
General Range. Eastern and north-central North America, in cold waters, from
northernmost Labrador, the southern part of Hudson Bay, and the tributaries of James
Bay, southward coastwise to northern New Jersey, and thence inland along the Allegheny
Mountains to North Carolina and northern Georgia; westward about as far as the
western slope of the Alleghenies in the southern part of the range; to northeastern Iowa,
Wisconsin, southeastern Minnesota, Lake Superior with its northern tributaries, and
to eastern Saskatchewan in the northern part; also widely and successfully introduced
in high altitude streams and lakes in the Rocky Mountains of the United States,
California, British Columbia, and southern Alaska; likewise in various localities in
South America, South Africa, and northern Europe.
Occurrence of Sea-run Populations. Migratory populations formerly existed as far
southward as Long Island, New York, where Herbert described them as weighing
up to five pounds in the salt creeks (33: 303) and where Goode again reported them
in 1884 (26); a century ago they abounded also in suitable places at the head of Buzzards
Bay and along the southern shores of Cape Cod in southern Massachusetts.? But
native Salters have been nearly, if not completely, wiped out on Long Island by over-
fishing combined with intermixture with hatchery fish; although a few are still taken
in the salt creeks and off their mouths along the southern and northern shores of Cape
Cod, it is doubtful whether any of the pure native strain still exists there, so widespread
and long continued has been the introduction of hatchery stock.
Salters seem never to have been known anywhere between the northern drainage
36. Information contributed by Charles A. Ritzi.
37- For an interesting account of the status of the Brook Trout in the Cape Cod region early in the past century,
see Smith (60: 348-394).
540 Memoir Sears Foundation for Marine Research
of Cape Cod, Massachusetts, and Cape Elizabeth, Maine, unless perhaps in one or two
small streams tributary to Ipswich Bay, Massachusetts. However, they do populate
many of the small tributaries of Casco Bay, Maine (20: 105; 45: 49, table), and there
are sea-going populations in the area near Jonesport (see p. 534 and Study Material,
p- 525); they probably occur elsewhere as well along the eastern part of the Maine
coast; but these can hardly be plentiful, otherwise anglers would be familiar with them.
Huntsman found no evidence of them in either salt or brackish water along the New
Brunswick shore of the Bay of Fundy (38: 60), but White has reported them for the
river estuaries at the head of the Bay (75: 180); according to local information, sea-run-
ning fontinalis still exist along the Nova Scotian shore of the Bay, notably in Salmon River.
Detailed information is lacking as to their status along the outer Nova Scotian
coast west of Halifax, but east of Halifax there are Salters in practically every trout
stream that is not blocked by high falls:** all around Cape Breton, along the entire
southern coastline of the Gulf of St. Lawrence, around Prince Edward Island, the
Magdalens, and the Island of Anticosti. Palmer has also reported them as being
plentiful enough to be worth the attention of anglers at the mouth of some 26 rivers
and small streams along the west coast of Newfoundland, about 39 along the south
coast, and about 25 along the east coast (50).
Published information about the Salter situation along the north shore in the inner
part of the Gulf of St. Lawrence is scanty, doubtless because they are so greatly over-
shadowed there by Atlantic salmon. But A. A. Blair has contributed the information
that all rivers and brooks along this coast are well stocked with them; and Storer noted
more than a century ago that Brook Trout were seined by cod fishermen at the brook
mouths there (63: 264). Blanc Sablon Bay, just within the Strait of Belle Isle on the
Labrador side, has been described by Barteauas “filled with sea trout from June through
July” (48: 90); they are also reported in numbers for Barge, Wreck, Red, Chateau,
and Temple bays within the Strait; St. Lewis Bay and Capelin bays northward; and
Hawke Bay, for which Barteau has reported a catch of 33 fish that weighed 137
pounds (48: 90, 91). Backus has also reported large runs of migrating fontinalis for
July and August in the Hamilton Inlet-Sandwich Bay region (3: 294). The vicinity
of Nain (about 56°30’ N) is the next region to the north whence they have been reported
in scientific literature (72: 128), and their presence along the intervening coast was
established by my own local inquiry during the summer of 1900.
Nutak Harbor, on the northwesternmost of the four Okkak Islands (57°28’N), is
the northernmost station of recent record for sea-run /fontinalis on the Atlantic coast
of Labrador, though Gordon and Backus think it likely that “‘small populations are
more or less continuously distributed around Cape Chidley and into Ungava Bay’”’
(27: 17). There are sea-going populations of fontina/is in the rivers that drain into the
southern part of Ungava Bay (16: 95-97) as well as in the southern part of Hudson
Bay, especially around the shores of James Bay, where they are widespread in large
numbers (66: 40).
38. For regional details, see especially Breck (8).
Fishes of the Western North Atlantic 541
Synonyms and References for Brackish or Salt water:°9
Salmo fontinalis Mitchill, Trans. Lit. philos. Soc. N.Y., 2, 1815: 435 (descr., color, Long Island, New York);
Perley, Rep. Fish. Bay of Fundy, 1851: 128 (color, abund., New Brunswick and Nova Scotia, descent to
sea); Storer, J. Boston Soc. nat. Hist., 6, 1857: 264 (abund., s. Labrador coast); Garman, rgth Rep.
Comm. inl. Fish. Mass. (1884), 1885: 76, figs. 14-15 (early refs., good descr., ills.; not fig. 16—a speci-
men of a/pinus species complex, probably subsp. aureo/us, from New Hampshire).
Salmo alleganiensis Rafinesque, Ichthyol. Ohiensis, 1820: 44 (brooks trib. to Allegheny and Monongahela
rivers, Ohio; not seen).
Salmo nigrescens Rafinesque, Ichthyol. Ohiensis, 1820: 45 (Pennsylvania; not seen).
Salmo fario Smith, Nat. Hist. Fish. Mass., 1833: 141 (size, color, in fresh water).
Salmo trutta Smith, Nat. Hist. Fish. Mass., 1833: 140-141 (river mouths, small rivers, Massachusetts); Herbert,
H.W., Frank Forester’s Fish and Fish. U.S., 1850: 120-125 (Gulf of St. Lawrence, diff. from sea-run
Sontinalis of Long Island and Massachusetts); Perley iz Herbert, H. W., Frank Forester’s Fish and Fish.
U.S., 1850: 123-124 (abund., size, capture, Gulf of St. Lawrence); Rep. Fish. Bay of Fundy, 1851: 131
(abund., sea-run, n. shore Nova Scotia, New Brunswick, Prince Edward I., and Magdalens; angling);
Bell, Canad. Nat. Geol., 4, 1859: 206 (abund., Gaspé, Quebec); Reeks, Zoologist, London, 2 (6), 1870:
2555 (abund., sea-run, capture, w. coast Newfoundld.).
Salmo canadensis Smith in Cuvier, Anim. Kingd., Pisces, Griffith ed., 10, 1834: 474, pl. 41 opp. p. 174 (color,
color. ills, Canada); Gilpin, Proc. N.S. Inst. Sci., r (4), 1866 (descr., Nova Scotia).
Baione fontinalis DeKay, Zool. N.Y., 4, 1842: 244 (descr., Rockland County, New York).
Salmo hudsonicus Suckley, Ann. N.Y. Lyc., 7, 1862: 310 (descr., color, cf. fowtinalis, Hudson Bay and tribu-
taries, Labrador, Newfoundld.); Weiz, Proc. Boston Soc. nat. Hist., 10, 1866: 275 (brackish water,
Square I., Okkak, n. Labrador).
Trutta argentina Scott, G.C., Fishing in Amer. Waters, 1869: 256 (descr., sea-run, Canada, Nova Scotia, New-
foundld.; not seen).
Trutta marina Scott, G.C., Fishing in Amer. Waters, 1869: 256 (see preceding ref.).
Salmo symmetrica Baird, Rep. U.S. Comm. Fish. (1872—1873), 2, 1874: 372 (color, cf. “brook trout,” Monad-
nock Lake, New Hampshire).
Salmo immaculatus Lanman, Rep. U.S. Comm. Fish. (1872-1873), 2, 1874: 221 (descr. after Storer, 1857;
name immacudatus applied to sea-run fowtina/is of s. Gulf of St. Lawrence; actually a synon. of a/pinus).
Salvelinus fontinalis Jordan, Proc. U.S. nat. Mus., 2, 1878: 80, 82 (fontinalis a Salvelinus, not a Salmo, distinct.
chars. in key); Goode, e¢ a/., Fish. Fish. Industr. U.S., r (3), 1884: 497, 502, pl. 192 (distr., size, habits,
spawn., early develop., in salt creeks, Long Island, New York); Jordan and Evermann, Bull. U.S. nat. Mus.,
47 (1), 1896: 506 (descr., range, in salt water, synon.); Schmitt, Monogr. Isle d’Anticosti, 4, Zool., Poiss.,
1904: 285 (abund., estuary of Riv. aux Canards); Evermann, Rep. U.S. Bur. Fish. (1904), 1905: 105
(salt water, food, Casco Bay, Maine); Kendall, Proc. Portland Soc. nat. Hist., 2 (8), 1909: 210, 228-242
(saltwater locals., Labrador, for 1891; refs. to other Labrador recs.); Kendall, Mem. Boston Soc. nat.
Hist., S (1), 1914: 75-103, table p.g9, pl. 7 (genl. acct., color. ill., tidewater, Casco Bay, Maine);
Huntsman, Contr. Canad. Biol. (1921), 3, 1922: 60 (no report of sea-run populations, New Brunswick,
shore, Bay of Fundy); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 60 (range,
synon., checklist); White, Trans. Amer. Fish. Soc., 60, 1930: 101-105 (spawn., freshwater and sea-
run populations, Prince Edward I.); Vladykov, Contr. Canad. Biol., N.S. 8 (2), 1933: 18 (sea-run,
size, abund., se. Hudson Bay, James Bay); Weed, Copeia, 1934: 131-133 (sea-run, color, food, vic.
of Nain, n. Labrador); White, Trans. Amer. Fish. Soc., 64, 1934: 356-357 (spawn. season rel. to temp.);
Vladykov and McKenzie, Proc. N.S. Inst. Sci., 79 (1), 1935: 57 (sea-run, size, Nova Scotia); Frost, Fish.
Bull., Dept. nat. Resources Newfoundld., 9, 1940: 10 (spawn., e. coast Newfoundld.); White, J. Fish. Res.
Bd. Canada, 5 (2), 1940: 176-186 (migr., food, sea-run population, Moser R., Nova Scotia, cf.
salmon); J. Fish. Res. Bd. Canada, 5 (3), 1941: 258-264 (migr., time in sea, survival rate, Moser R.,
Nova Scotia); J. Fish. Res. Bd. Canada, 5 (5), 1942: 471-473 (sea life, off Moser R., Nova Scotia);
Doan, Bull. Fish. Res. Bd. Canada, 79, 1948: 1-12 (sea-run, w. coast Hudson Bay); Wilder, J. Fish. Res.
39- The Brook Trout (really a charr) is so great a favorite with anglers that the literature treating of it has grown to
enormous proportions, both in scientific publications and in sportsmen’s magazines. As the present account is
centered about fontinalis in brackish or salt water, the following list includes only those references that touch
directly on the sea-going populations, with a few general works included.
542 Memoir Sears Foundation for Marine Research
Bd. Canada, 9 (4), 1952: 169-203 (sea-run trout, Moser R., Nova Scotia, cf. freshw. trout, color,
migr.); Dunbar and Hildebrand, J. Fish. Res. Bd. Canada, 9 (2), 1952: 95-96 (sea-run, Ungava Bay,
n. Labrador, size of “spotted trout” in trib.); Vladykov, J. Fish. Res. Bd. Canada, rr (6), 1954: go4—
932 (det. cf. 8. a/pinus species complex and Cristivomer); Gordon and Backus, Copeia, 1957: 17 (sea-
run, Nutak Harbor, Labrador, 57°28’N); Backus, Bull. Amer. Mus. nat. Hist., 773 (4), 1957: 292-
295 (abund. sea-run, Hamilton Inlet-Sandwich Bay area, Labrador, July-Aug., freshw. Labrador
locals.); Bridges and Mullan, Fish. Bull. Mass. Fish. Game, 23, 1958: 38 pp. (genl. life hist., ecol., e.
fontinalis, present sea-run, Massachusetts, refs.); Mullan, Fish. Bull. Mass. Fish. Game, 17, 1958:
25 pp. (present status sea-run fomtinalis, Cape Cod, effects of hatchery fish).
Salmo agassizii Garman, 19th Rep. Comm. inl. Fish. Mass. (1884), 1885: 78, fig. 17 (cf. fontinalis, good ill.,
Dublin Pond, New Hampshire, not fig. 18 —a specimen of a/pinus species complex, probably subspp.
aureolus ox oguassa, local. not given).
Salvelinus fontinalis agassizii Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 507 (color, cf. typ-
ical fontinalis, Lake Monadnock, New Hampshire).
Salvelinus fontinalis hudsonicus Hubbs, Misc. Publ. Mus. Zool., Univ. Mich., 15, 1926: 19 (cf. typical fow-
tinalis, sea-run, Gulf of St. Lawrence); Copeia, 194.3: 258 (subsp. of fontinalis).
Salvelinus agassizii Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 60 (listed, Dublin
Pond, New Hampshire).
Salvelinus (Baione) fontinalis Vladykov, J. Fish. Res. Bd. Canada, rr (6), 1954: 928, 930 ( fontinalis regarded
as sep. subgen. Baione).
Sea trout, Smith, Nat. Hist. Fish. Mass., 1833: 348-394 (abund., sea-run populations, angling, catches, Cape
Cod, Massachusetts); Millais e¢ a/., Newfoundld. Guide Bk., 1905: 65—g1 (sizes, abund., sea-run, New-
foundld., Labrador coast N. to Sandwich Bay); Breck, Edw., Sport. Guide to Nova Scotia, Halifax,
190g: 1-101 (Nova Scotia, sea-run, season); Palmer, Salmon Rivers Newfoundld., 1928: 271 pp.
(details, sea-run, rivers and streams, Newfoundld.).
Trout, Herbert, H. W., Frank Forester’s Field Sports, Append. A, 1849: 303 (abund., size, salt creeks, Long
Island, New York).
Sea trout or Canadian trout, Norris, Thad., Amer. Angler’s Bk., 1864: 239-247 (sea-run, abund., weights,
as game fish, n. and s. shores Gulf of St. Lawrence).
Brook trout, Green, Amer. Angler, 7, 1885: 312 (size at hatching, early growth, size, Long Island, New
York, saltwater occur. implied).
Genus Cristivomer Gill and Jordan 1878
Lake Trout
By
Henry B. BicELow
Museum of Comparative Zoology
Harvard University
Cristivomer Gill and Jordan iz Jordan, Manual Vert. north. U.S., ed. 2, 1878: 356, 3593 type species, (Sa/mo)
namaycush Nalbaum; Lakes of Rocky Mountains to Maine and northward.
Generic Synonym:
Salmo, in part, Walbaum, P. Artedi Genera Pisc., Ichthyol., Emend., 3, 1792: 68, for 8. zamaycush Walbaum;
and various subsequent authors.
Characters. Essentially as in Sa/velinus (p. 503), but with: the frontal area of
Sku t flat; rear contour of Caupat fin deeply forked (Fig. 129); two longitudinal rows
of TzeTu on tongue parallel and close together (wide apart and forming a triangle in
Fishes of the Western North Atlantic 542
Salvelinus); and usually more than 120 Gastric cozca (av. only 38 or 39 in Sa/velinus)
(67: 928).
Size. The Lake Trout is by far the largest of the small-scaled salmonoids of North
America. One of 63 pounds was taken on rod and reel by Miss L. L. Hayes in Lake
Athapapuskow, Manitoba, in August 1930, and fish even up to 120-125 pounds have
been reported by commercial fishermen, but without supporting evidence.
Range. Lakes and some streams of North America, from the northern United
States north to the Arctic Circle, sometimes occurring as a stray in salt or brackish
water of Arctic Canada.
Ficure 129. Cristivomer namaycush, 290 mm SL, St. Lawrence basin, Canada, MCZ 33163.
Remarks. The Lake Trout of Lake Superior was described by Louis Agassiz in
1850 as a separate species, Salmo siscowet (I: 336), which was reduced to the rank
of subspecies by Jordan and Evermann with the remark “‘that it is probably a local va-
riety” (42: 505). Neither do I find anything in Agassiz’ original description to justify
any formal recognition of siscower in ichthyological nomenclature.
Species. One species only is known, the common Lake Trout (C. zamaycush) of
northern North America, which perhaps includes one recognizable subspecies, siscowet
L. Agassiz 1850, in Lake Superior.
Cristivomer namaycush (Walbaum) 1792
Lake Trout, Mackinaw Trout, Togue, Gray Trout
Figures 118, 129
Cristivomer namaycush is typically a freshwater fish; the only reason for mentioning
it here is that stray specimens have been reported for Hudson Bay, for Ungava Bay in
northern Quebec, and for the northeastern Labrador coast. Should one be taken in salt
water, the minuteness of its scales would show at a glance that it is not Sa/mo salar
(Atlantic salmon, p. 460) or any other member of the genus Sa/mo for that matter
544 Memoir Sears Foundation for Marine Research
(pp: 459-499); and the rear contour of its caudal fin is much more deeply concave than
that of the sea-run Sa/velinus fontinalis (Fig. 128). The danger might be greater of
mistaking a stray for Sa/velinus alpinus (Arctic charr), for while the head is somewhat
longer and the caudal fin more deeply concave in the Lake Trout than in the Arctic
charr, the contrasts between the two fish in these respects are not wide (Fig. 118; also
cf. Figs. 125, 126, 129). In case of doubt, cut open the body cavity and estimate
(even roughly) how many gastric coeca there are, for while the Lake Trout has about
120, the average for the Arctic charr is only about 39 (p. 510).
Synonyms:
Salmo namaycush Walbaum, P. Artedi Genera Pisc., Ichthyol., Emend., 3, 1792: 68 (brief descr. based on
namaycush of Pennant, Arctic Zool., Introd., z, 1784: cxcr; lakes far inland from Hudson Bay).
Salmo salar Williams, Nat. Civil. Hist. Vt., 1794: 121, 122 (Lake Champlain, other Vermont locals.).
Salmo symmetrica Prescott, Amer. J. Sci., (2) IZ, 1851: 340 (Lake “Winnipiscogee,” New Hampshire, and con-
necting waters).
Salmo toma Hamlin ix Holmes, Ezekial, 2nd Rep. nat. Hist. Geol. Maine, 2, 1862: 10g (great lakes, deep
mountain tarns, Maine; New Brunswick, Canada).
Cristivomer namaycush Evermann and Kendall, Rep. U.S. Comm. Fish. (1894), 20, 1896: 591 (Lake Champlain,
L. Memphremagog, Vermont).
Salvelinus namaycush Walters, Bull. Amer. Mus. nat. Hist., 106 (5), 1955: 275.
Probable Synonym:
Salmo tructa Belknap, J. Hist. N.H., 3, 1792: 179 (“Winnipisogee” Lake and stream, New Hampshire).
References for Salt Water:
Weed, Copeia, 1934: 128 (rec. for Adlok Bay, near Hopedale; for Upaqtik Bay between Hopedale and Nain,
Atlant. coast of Labrador); Dunbar and Hildebrand, J. Fish. Res. Bd. Canada, 9 (2), 1952: 95 (rep. by
natives for brackish water, George and Koksoak rivers, tributary to Ungava Bay, n. Quebec; but no report
of it for salt water there); Walters, Bull. nat. Mus. Canada, Biol. Ser., 128, 1953: 261 (brackish water, Hudson
Bay; river mouths, Coronation Gulf); Bull. Amer. Mus. nat. Hist., 106 (5), 1955: 275 (Bathurst Inlet, Arctic
Canada, near mouth of Burnside R.).
SA SSS
Io.
II.
I2.
Ti)
14.
T5.
16.
I.
Ts.
19.
20.
TEXTOAND EOQOTNOTLE, REBFERENGES
Aassiz, L., Lake Superior... 1850.
Andrews and Lear, J. Fish. Res. Bd. Canad., 73
(6), 1956.
Brenus Bull. Amer. Mus. nat. Hist., 773 (4),
1957-
Bell, Canad. Nat. Geol., 4, 1859.
Betz, Outdoor Life, N.Y., 122 (2), Aug.1958.
Bigelow and Schroeder, Fish. Bull. (74) U.S.
Fish Wildl. Serv., 53, 1953.
Bigelow and Welsh, Bull. U.S. Bur. Fish.
(1924), 40 (1), 1925.
Breck, Edw., Sport. Guide to Nova Scotia,
1909.
Bridges and Mullan, Fish Bull. Mass. Fish.
Game, 23, 1958.
Comeau, Life and Sport N. Shore Lower
St. Lawrence and Gulf, 1909.
Din, Skr. Svalb. Ishavet, 7, 1926.
Delacy and Morton, Trans. Amer. Fish. Soc.,
72, 1942.
Doan, Bull. Fish. Res. Bd. Canad., 79, 1948.
Dresel, Proc. U.S. nat. Mus., 7, 1885.
Dunbar, Arctic, 9, 1956.
Dunbar and Hildebrand, J. Fish. Res. Bd.
Canad., 9 (2), 1952.
Dymond, Bull. Biol. Bd. Canad., 32, 1932.
Dymond, Canad. Field Nat., 46, 1932.
Emsony, Trans. Amer. Fish. Soc., 64, 1934.
Evermann, Rep. U.S. Bur. Fish. (1904),
1905.
545
2I.
Evermann and Goldsborough, Bull. U.S. Bur.
Fish. (1906), 26, 1907.
| eereree Fauna Groenl., 1780.
Fish, Bull. U.S. Bur. Fish. (1925), 42, 1926.
Frost, Fish. Bull., Dept. Nat. Res. Newfoundld.,
9, 1940.
Garman, tgth Rep. Comm. inl. Fish. Mass.
(1884), 1885.
Goode, et a/., Fish. Fish. Industr. U.S., r (3),
1884.
Gordon and Backus, Copeia, 1957.
Grainger, J. Fish. Res. Bd. Canad., ro (6), 1953.
Green, Amer. Angler, 7, 1885.
Giinther, Proc. zool. Soc. Lond., 1877.
Hecxer, S.B. Akad. Wiss. Wien, Math Nat.
K1., 8 (3), 1852.
Henn, Mem. Carnegie Mus., 12 (2 [3]), 1932.
Herbert, H. W., Frank Forester’s Field Sports,
1849.
Hewitt, Telling on Trout, 1930.
Hinks, Fishes Manitoba, Dept. Mines nat. Re-
sources, 1943.
Hubbs, Misc. Publ. Mus. Zool., Univ. Mich.,
15, 1926.
Hubbs, Copeia, 1943.
Huntsman, Coutr. Canad. Biol. (1921), 3, 1922.
5 eo Spolia zool. Mus. Hauniensis, 9, 1948.
Johansen, Medd. Gronl., 45 (12), 1912.
Jordan, Guide to Study Fish., 2, 1g05.
35
546
42.
43:
46.
47-
48.
49-
50.
5I.
52.
53:
54:
55:
56.
57:
58.
Jordan and Evermann, Bull. U.S. nat. Mus.,
47 (Z), 1896.
Jordan, Evermann, and Clark, Rep. U.S. Comm.
Fish. (1928), 2, 1930.
| apie Proc. U.S. nat. Mus., 38, 191t.
Kendall, Mem. Boston Soc. nat. Hist., 8 (1),
1914.
Kendall, Copeia, 1919.
Manaine, Canad. Field Nat. (1942), 56 (8
and 9), 1943.
Millais, e¢ 2/., Newfoundld. Guide Bk., 1905.
Mullan, Fish. Bull. Mass. Fish. Game, 17,
1958.
leaner Salmon Rivers Newfoundld, 1928.
Perley, Cat. [in part] Fish. N. Brunsw. and
Nova Scotia iv Rep. Fisher. Bay of Fundy,
1851.
Porsild, Medd. Gronl., 60, 1921.
Reexs, Zoologist., London, (2) 6, 1870.
Regan, Freshwater Fish. Brit. Isles, grt.
Regan, Ann. Mag. nat. Hist., (8) 73, 1914.
Richardson, Fauna Boreal.-Amer., 3, 1836.
Ricker, Publ. Ont. Fish. Res. Lab., Toronto
Stud., Biol., 36 (44), 1932.
Ross, Narr. Second Voy.... North-west Pas-
sage, Append., Nat. Hist., Fish., 1835.
59:
60.
6r.
62.
63.
64.
65.
66.
67.
Memoir Sears Foundation for Marine Research
Stesoxp, von, Siisswasserfische Mitteleuropa,
1863.
Smith, Nat. Hist. Fish. Mass., 1833.
Smitt, Hist. Scand. Fish., ed. 2, 2, 1895.
Sprules, J. Fish. Res. Bd. Canad., 9 (1), 1952.
Storer, J. Boston Soc. nat. Hist., 6, 1857.
Suckley, Ann. N.Y. Lyc., 7, 1862.
Sumner, Osburn, and Cole, Bull. U.S. Bur. Fish.
(1911), 32 (2), 1913.
Viapyxov, Contr. Canad. Biol., N.S. 8 (2),
1933-
Vladykov, J. Fish. Res. Bd. Canad., rr (6),
1954-
Vladykov, J. Fish. Res. Bd. Canad., 13 (6),
1956.
WW nares P. Artedi Genera Pisc., Ichthyol.,
Emend., 3, 1792.
Walters, Bull. nat. Mus. Canada (1951-1952),
Biol. Ser., 128, 1953.
Walters, Bull. Amer. Mus. nat. Hist., 706 (5),
1955-
Weed, Copeia, 1934.
Welch, Outdoor Life, N.Y., 122 (5), 1958.
White, Trans. Amer. Fish. Soc., 64, 1934.
White, J. Fish. Res. Bd. Canad., 5 (2), 1940.
White, J. Fish. Res. Bd. Canad., 5 (3), 1941.
White, J. Fish. Res. Bd. Canad., 5 (5), 1942.
Wilder, J. Fish. Res. Bd. Canad., 9 (4), 1952-
Yessirov, Trans. arct. Inst. Leningr., 77, 1935
Family Coregonidae
HENRY B. BIGELOW
Museum of Comparative Zoology
Harvard University
Characters. The same as those in the Salmonidae, but with: Trerru minute, if
present at all; Scares relatively large, not more than 13 scales in a transverse series
from origin of dorsal fin to lateral line (at least 19 among Salmonidae); and Pariera
songs of skull meeting in the midline (not meeting among Salmonidae).
Remarks. The Whitefishes are widespread in middle and high latitudes in lakes
and streams throughout Eurasia and North America. They spawn in fresh water and are
confined there throughout the greater part of their geographic range, though in many
places they run down into brackish water and out into salt water.
Genera. ““The taxonomy of this group has been particularly confusing” (8: 276)
and seems likely to continue so, as to both genera and species. Thus, Jordan, er a/.,
reported five genera: Stenodus, Leucichthys (with three subgenera), Coregonus, Pro-
sopium, and Trillion for North America (4: 61-66). Norman, however, recognized
only Stenodus and Coregonus on a world-wide basis (7: 104), an extreme simplifica-
tion that does not accord with the weight of present-day opinion.
Key to Genera Recognized for North America
1a. Teeth distinct, in bands on jaws and palatines; a patch on vomer and another
on tongue. Stenodus Richardson 1936.
Sheefish or Inconnu.
Fresh and brackish waters of northern
Europe and Asia, from White Sea
(Mezen R.) eastward to western Be-
ting Sea drainages (Bay of Korf); Kus-
35
547
548 Memoir Sears Foundation for Marine Research
1b. Teeth vestigal or wholly lacking.
kokwim River, Alaska; western Arctic
Canada southward to northern British
Columbia (Teslin and Kluane lakes)
and to Fort Smith on Slave River be-
tween Great Slave Lake and Lake
Athabasca (9: 9-10).
2a. Base of adipose fin nearly or quite as long as base of anal fin.
Trillion Jordan 1918.
Rivers of Oregon.
2b. Base of adipose fin not more than about half as long as base of anal.
3a. A single fleshy flap on either side on snout between nasal openings; body
rounded; gill rakers short, knob-shaped.
Prosopium Milner 1878.
Round Whitefishes.
Northern Siberia; North American Arctic
coasts eastward to Bathurst Inlet, Hudson
Bay, Ungava Bay, and Labrador Peninsula;
southward in the east to the Great Lakes
region, Quebec, northern New York, and
Connecticut; in the west to Washington,
northern Idaho, western Montana, Wyoming,
Utah, and Nevada. Not reported in northern
or western Ontario, interior Manitoba, Sas-
katchewan, or Alberta.
3b. Two fleshy flaps on either side on snout between nasal openings; body
laterally flattened in most; gill rakers
slender, pointed.
4a. Margin of upper jaw noticeably sinuous in contour, the premaxillary
bones wider than long; tip of upper jaw extending beyond tip of
lower jaw; front of snout bluntly rounded; gill rakers fewer than 28,
the longest not more than a third as long as lower limb of arch.
Coregonus Linnaeus 1758, p. 549.
4b. Margin of upper jaw only weakly sinuous in contour, the premaxillary
bones longer than wide; tip of lower jaw extending at least as far as
tip of upper jaw; front of snout
tapering; gill rakers usually more
than 31, the longest nearly or quite half as long as lower limb of
gill arch. Leucichthys! Dybowski 1870.
Chubs,
Ciscoes, and Tullibees.
Alaska in general; Arctic coast of Canada and Hud-
son Bay; North-Central Canada in general, south-
ward to the Great Lakes region, and to northern
and central New York. Not reported for Newfound-
land, or for northern or eastern Labrador.
1. Walters considers Leucichthys a subgenus of Coregonus (8: 279, 281).
Fishes of the Western North Atlantic 549
Genus Coregonus Linnaeus 1758
Coregoni, Linnaeus, Syst. Nat., ed. 10, I, 1758: 310, for Sa/mo /avaretus Linnaeus. Europe.
Generic Synonyms:
Tripteronotus Lacépéde, Hist. Nat. Poiss., 5, 1803: 47; type and only included species, T. Aautin Lacépéde
equals Sa/mo /avaretus Linnaeus 1758.
Coregonus Lacépéde, Hist. Nat. Poiss., 5, 1803: 239; type (first mentioned) species, C. /evaretus Lacépéde equals
Salmo lavaretus Linnaeus 1758.
Characters. Base of Apipose fin only about half as long as base of anal. Bopy
laterally flattened, at least in most. SNout with two fleshy flaps on either side between
nasal openings, bluntly rounded in front. Tip of Upper jaw projecting a little beyond
lower jaw, its margin conspicuously sinuous. PREMAXILLARY BONES wider than long.
GILL RAKERs on first arch fewer than 28, the longest not more than a third as long as
lower limb of gill arch.
Size. Coregonus often weigh up to five pounds, and occasional specimens are much
heavier.
Relation to man. The Lake Whitefishes are highly esteemed market fish wherever
they are plentiful enough to support a commercial fishery; in the Great Lakes region
they are the most valuable of the foodfishes.
Species. The genus Coregonus, as defined above, is represented in North American
waters by two rather sharply defined species: C. xasus (Pallas) 1776, with the gill rakers
shorter than the diameter of the eye (9g: 13), occurs in Siberia, Alaska, and Arctic
Canada; and C. clupeaformis, possibly identical to C. /avaretus (Linnaeus) 1758 of
Europe, is the common Lake Whitefish of North America. A third form, xe/soni Bean
1884, also with fewer gill rakers than c/upeaformis, has been reported repeatedly for
Alaska as a separate species, but it is only doubtfully distinguishable from C. pidschian
(Gmelin) 1788? of northern Asia. Our only direct concern here is with C. clupeaformis.
Coregonus clupeaformis Mitchill 1818
Lake Whitefish
Figure 130
Study Material. About 20 specimens, from Lake Superior, Michigan, New York,
New Hampshire, Maine, and the Saskatchewan River.
Distinctive Characters. Should a coregonid of this or any other species be taken
in brackish or salt water, it is not likely that it would be mistaken for another fish, for
while its adipose fin and general make up would evidently relate it to the salmonids
and osmerids, its very small mouth with tiny teeth (if it has any) would mark it off at
a glance from all of these. There might be more chance of mistaking a Lake Whitefish
for an argentinid were it not that the argentinid’s pelvic fins are posterior to the rear
2. For a recent discussion of this matter, see especially Walters (8: 284-288).
550 Memoir Sears Foundation for Marine Research
end of the base of the rayed dorsal fin rather than under the midpoint of the base of the
dorsal, as in the Whitefishes. If a specimen of this family is taken anywhere along
the Atlantic coast of eastern North America and if it does not agree in detail with
the following illustration of the common Lake Whitefish (Fig. 130), it is probably a
Round Whitefish (genus Prosopium, p. 548).
Description. As the Lake Whitefish is not a regular member of the saltwater fish
fauna anywhere on the Atlantic coast of North America, the accompanying illustration
seems sufficient for identification without detailed description.
Suan,
ae x) Dns uM Ryn
any PM Ha a Mis I ae
eS yy iy yy
yyy I) Soe
si mins V5 i Wire oy ee
yi oy ee Sse) ua Jy Meee
- Cc se ee on My TOE
Peay 30) pie wis
= oe Rye
LES
pias my nh? ae Ne
Yo )
Figure 130. Coregonus clupeaformis, from Ecorse, Michigan, USNM 10300, after Goode. Drawn by
H. L. Todd.
Subspecies. C. clupeaformis, like most of the coregonids, is extremely variable, so
much so that Koeltz, in his detailed study (5: 370-381), recognized seven subspecies,
four of which, c. c/upeaformis (Mitchill) 1818, c. stanleyi Kendall 1904, c. neo-hauto-
niensis (Prescott) 1851, and c. gu/liveri Koeltz 1931, are reported for the Atlantic
watershed of eastern North America. Discussion of these would be out of place here,
for C. clupeaformis, so far as is known, occurs only as a stray in salt water in the
western North Atlantic, though it does occur more commonly in such situations in
Hudson Bay.
Range in North America. Arctic coasts and drainage basins of Alaska and Canada,
including the Ungava region of northern Quebec, and southward generally to Nova
Scotia (see below), northern New England in general, northern and central New York,
south-central Ontario (Peterborough County), the Great Lakes region, northern
Manitoba, central and southern British Columbia and Montana, and to the Bristol Bay
drainage basin on the Bering Sea coast of Alaska.
Occurrence in the Western North Atlantic. The only published record for a coregonid
of any sort in salt water on the Atlantic coast of North America is of two that were
Fishes of the Western North Atlantic Bee
taken in ‘‘full salt water’® in midchannel at the mouth of the Sissibo River, St. Mary
Bay, Nova Scotia, in 1919 (3: 59). These specimens, no longer in existence, were doubt-
fully identified as guadrilateralis, the Round Whitefish, which is commonly placed in
the genus Prosopium; but doubtless they were c/upeaformis, for R. A. McKenzie has
written that all of the whitefishes from that general part of the Province received at
the St. Andrews Laboratory of late years have been c/upeaformis.t W. B. Scott has also
reported that an additional c/upeaformis was caught off a wharf at Wedgeport, Nova
Scotia, in July 1954, and that others were taken or seen there during the ensuing
month by W. L. Klawe.® But there seems to be no reason to suppose that any
regular sea-run populations of whitefish occur anywhere on the Atlantic coast of
America or among the Arctic islands to the north.
In Hudson Bay, however, in the vicinity of Churchill on the west coast and in
James Bay northward to Great Whale River in the east, both Lake Whitefish (Coregonus)
and Round Whitefish (Prosopium) as well as one of the Ciscoes (Leucichthys artedi, sub-
species not recorded) appear to run down regularly into salt water (r: 12).
3- Personal communication from A. G. Huntsman; in Gunter (2: 313).
4. Specimens of c/upeaformis that were taken in fresh water in the headwaters of the LaHave River, Canada, were in-
advertently included among a list of marine records (McKenzie, 6: 43).
5. From a letter to J. R. Dymond by W. B. Scott.
TEXT AND FOOTNOTE REFERENCES
: Dymonp, Contr. Canad. Biol., N.S. 8(1), 1933- 6. Mexenzie, Proc. N.S. Inst. Sci., 20 (2),
1940.
; Gonrer, Amer. Midl. Nat. (1942), 28 (2),
1943.
7. N ORMAN, Synop. Orders, Families, Genera of
. Huntsman, Contr. Canad. Biol. (1921), 3; Recent Fish. (mimeo), 1957.
1922.
: Tisoxs, Evermann, and Clark, Rep. U.S.
Comm. Fish. (1928), 2, 1930. 8. Wares Bull. Amer. Mus. nat. Hist., 706
(5)> 1955-
g. Wynne-Edwards, Bull. Fish. Res. Bd. Canad.,
Kosxrz, Pap. Mich. Acad. Sci., 23, 1931. 94, 1952.
55
Family Osmeridae
HENRY, B. BIGELOW
Museum of Comparative Zoology
Harvard University
and
WILLIAM C. SCHROEDER
Woods Hole Oceanographic Institution+
Characters. Essentially as in the Salmonidae (p. 457), but with: sides lacking the
FLESHY APPENDAGES above bases of pelvics that characterize Salmonidae; last few
VERTEBRAE definitely not upturned; neither an OrsirosPpHENOID bone separating the
orbital depressions of the skull nor a Posroric in the auditory region;? the Bony PLaTE
on dorsal side of skull above vomer divided longitudinally by a median suture in some
(Osmerus, Allosmerus, Thaleichthys, and Spirinchus) but not in others (Ma//otus),? nor
among the Salmonidae;* only a few PyLtoric carca or none (not more than 7 in any
western North Atlantic species).
Remarks. In their general appearance the smelt and capelin closely resemble
trout and young salmon, but there is no danger of mistaking the one for the other,
for the members of the smelt tribe are slimmer-bodied than the salmons, have more
deeply forked tails, and their pelvic fins stand farther forward relative to their dorsal
fin (see Key, p. 554); they differ further from the salmons in lacking the fleshy append-
ages above the pelvic bases.
Genera. Six genera seem to deserve recognition within the Osmeridae as defined
above, only two of which occur in the western North Atlantic, the others being con-
fined to the northern Pacific. Perhaps we should remark in passing that the ‘‘smelt”’
of New Zealand and of New South Wales, Australia, is not a true smelt, but belongs
to the related family Retropinnidae, the dorsal fin of which is far posterior to the pelvics.
- Contribution No. 1149 from the Woods Hole Oceanographic Institution.
. For additional details as to skull characters, see Berg (6: 240, 428).
. For proethmoid bones, see Chapman (rr: 280-291, figs. 3, 4) and Starks (ro6: 151, fig. 3, 335)-
. For dorsal view of the skull of the salmons, see especially Parker (74: 144, pl. 7, fig. 1) and Gregory (32: 154, fig. 48 A,
dermethmoid bone).
PWN H
553
isn Memoir Sears Foundation for Marine Research
For a recent account of the occurrence of the New Zealand smelt, with illustrations,
see Graham (32: 113). The so-called “herring smelt’ of the northern North Atlantic
belongs to the Argentinidae, a family differing widely from the true smelts in having
a very small mouth and a structure in the intestine reminiscent of the spiral valve of
elasmobranchs but probably not homologous with it (Part 4).
Key to Genera of Osmeridae®
1a. Scales small, in 170-220 transverse series. Mallotus Cuvier 1829, p. 573.
1b. Scales large, fewer than 80 along lateral line.
2a. Teeth on vomerine bone (roof of mouth) and on tip of tongue large, canine-like.
3a. Pelvic fins inserted under or posterior to origin of dorsal fin.
Osmerus Lacépéde 1803, below.
3b. Pelvic fins inserted definitely in advance of origin of dorsal fin.
4a. Head, as seen from above, sharply pointed; gill covers only weakly
striated; gill rakers 10 or 11 +22—26 on first arch; anal fin with only
1$—17 rays. Allosmerus Hubbs 1925.
Coast of northern California.
4b. Head, as seen from above, bluntly rounded; gill covers strongly
striated; gill rakers only 4-6+13-16 on first arch; anal fin with
20-22 rays. Thaleichthys Girard 1858.
Alaska to Oregon.
2b. Teeth on roof of mouth small, not canine-like.
sa. Pectoral fins considerably shorter than head; base of anal fin shorter than
head, its maximum height less than height of dorsal; upper jaw reaching
only a little past anterior edge of eye. Hypomesus Gill 1862.
Alaska to middle California;
also Kamchatka and Japan.
5b. Pectoral fins about as long as head; base of anal fin about as long as head,
its vertical height considerably greater than height of dorsal; upper jaw
reaching back as far as rear margin of eye.
Spirinchus Jordan and Evermann 1896.
Pacific coast of America, Middle California to Alaska.
Genus Osmerus Lacépéde 1803
Smelts
Osmerus Lacépéde, Hist. Nat. Poiss., 5, 1803: 229; type species, Sa/mo eper/anus Linnaeus, Syst. Nat., ed. 10,
T, 1758: 310. “Habitat in Europa.”
5. The so-called “‘ghost smelt,’ Therobromus callorhini Lucas 1898, which is known only from bones found in the
stomachs of fur seals, has been included in the Osmeridae by Jordan, et a/. (52: 68). But Hubbs has suggested that it
may not be an osmerid at all (43: 49, ftn.), while Chapman has concluded definitely that it is not (rr: 295). In any case,
it cannot be included in this Key for want of information about its external features.
Fishes of the Western North Atlantic 555
Generic Synonyms:
Salmo, in part, Linnaeus, Syst. Nat., ed. 10, , 1758: 310, for 8. eper/anus Linnaeus; Pallas, Zoogr. Rosso-Asiat.,
3, 1831: 386, 387, for 8. (Osmerus) eperlanus and 8. (Osmerus) spirinchus.
Eperlanus Gaimard, Voy. Islande et Groenl., Atlas, Zool., Poiss., 1851: pl. 18, fig. 2; type species Sa/mo eper-
Janus Linnaeus 1758.
Atherina Mitchill, Trans. Lit. philos. Soc. N.Y., I, 1815: 446; type species, 4.mordax Mitchill 1815. New
York.
Not Arherina Linnaeus, Syst. Nat., ed. 10, ZT, 1758: 315; type species, 4.hepsetus Linnaeus; Mediterranean; a
member of the Atherinidae (silversides), a family of percoid fishes.
Characters. Bopy slender, laterally flattened. Heap pointed, about 0.2 of length
to base of caudal fin. Mourn wide. Upper jaw reaching rearward beyond center of
eye. Tip of Lower jaw projecting a little beyond upper. Snour in front of eye about
as long as eye. BRANCHIOSTEGAL RAYS 7—8. GILL RAKERS about 67 °/, as long as eye,
about 25-37 on first arch.
Dorsat fin small, its origin at or near midlength of trunk, with 10-11 rays,
the first 2 or 3 rays not branched. Apipose fin well developed. Caupat fin deeply
forked. Anat with more rays than dorsal, but 17 at most, the first 3 (rarely 2) not
branched. Petvics under or a little posterior to origin of dorsal. PEcrorats about as
long as upper jaw.
Sca.es in less than 80 transverse series; thin, transparent, smooth-edged, the ex-
posed surface sculptured with many fine concentric striations, and wholly without
pigment; those perforated by the lateral line with a low median ridge, and some weakly
indented posteriorly in the midline;® scales elsewhere without median ridge and with
evenly rounded rear outline. Visible LarrraL Line not extending beyond origin of
dorsal fin.
TeetH. Roof of mouth with a few large canine-like teeth (not shed in spawning
season) on lateral tips of U-shaped vomerine bone; teeth also present on tip of tongue;
smaller teeth present on the jaws, the palatine bones on roof of mouth, and the pharynx.
Bony plate on dorsal side of skull above vomer (PRoETHMorp of Starks) with a longi-
tudinal median suture.
VERTEBRAE $5—64. PyLoric CAECA 7 at most.’
Sexual Dimorphism. At spawning time, large numbers of so-called “nuptial”
tubercles develop on the scales of the body and along the rays of the dorsal and pectoral
fins of the males, making them so rough to the touch that it is easy to tell the males
from the females (59: 141; 42: 86; 89: 27, 28).
Range. Coastal waters, estuarine situations, and fresh water in lower courses of
streams and rivers of the western North Atlantic from the outer coast of Labrador
(about 54°N; head of Hamilton Inlet) and the Strait of Belle Isle south regularly to
the tidal reaches of the rivers tributary to Delaware Bay (p. 568); the eastern Atlantic
from southeastern Norway and the Gulf of Bothnia southward to the Loire River, and
6. Smitt has described and pictured the lateral-line scales as being deeply incised posteriorly (ro4: 865, fig. 218). But
our own experience with the scales, of both American and European smelts, suggests that this appearance was
the result of mutilation.
7. For other anatomical details, see especially Smitt (104: 864, 865) and Chapman (zz: 280).
556 Memoir Sears Foundation for Marine Research
reported for Vigo in northern Spain (5: 132); the Arctic coasts of Eurasia, Alaska,
and Canada eastward as far as the estuary of the Mackenzie River; the western
Pacific southward to northern Japan and northern China; and the eastern Pacific to
Yakutat Bay, Alaska. They are landlocked in many lakes of Europe, including the
Volga River system and the basin of the Caspian Sea (57: 241, 308, 315, 323, 3463
49: 1023; 95: 649), eastern North America, eastern Asia, and the White Sea-Kola
Peninsula region. We should perhaps emphasize that, while distributed widely,
smelts are unknown along the Arctic coast of Canada east of the Mackenzie,
in Hudson Bay, Greenland, Iceland, Spitsbergen, and along the coast of western
Norway.
Species. The members of the genus Osmerus as limited above were originally
described under two names: (1) Salmo eperlanus Linnaeus 1758 for the smelts of
Europe, with which spirinchus Pallas 1811 (73: 387), from the lakes and rivers of
Russia,® and vulgaris Gaimard 1838 (29: pl. 18, fig. 2), from Iceland, clearly are syno-
nyms; and (2) Atherina mordax Mitchill 1815 for western Atlantic smelts. S. eper/anus
and 4. mordax (as viridescens LeSueur 1818) were retained as separate species, first by
Cuvier and Valenciennes, chiefly, it seems, on the assumption that LeSueur would have
recognized at first sight a fish with which he had doubtless been familiar since child-
hood (7g: 388), and then by Giinther (34: 166, 167). The list was soon enlarged with
sergeanti, proposed by Norris in 1868 for the smelts of the Schuylkill River (Delaware),
of Brandywine Creek (Pennsylvania), and of the tidal part of the Delaware River (70:
93); dentex, proposed in 1870 by Steindachner and Kner for those of northern China
(107: 429); abbotti and spectrum in 1871 by Cope for those of Wilton Pond and
““Cobessicontic”” Lake, Maine (16: 490); and dvinensis in 1882 by Smitt for those of
the White Sea (102: 34; 104: 867, ftn. k).
So far as we can learn, Rice’s comparison of the number of transverse rows of
scales on smelts from various American localities with others from Liverpool, England
(86: 91), was the earliest comparison of the sort that has been made; he concluded
that the American smelts are identical to the European eper/anus. The next landmark
in smelt taxonomy was Smitt’s (103: 168-188) very detailed comparison of the pro-
portional dimensions for the smelts of Scandinavia (as eper/anus), the White Sea (as
spirinchus), and the North Pacific (as dentex); from this he concluded that the three
represented three distinct species. In 1895, however, he followed Rice in uniting the
smelts on the two sides of the Atlantic under the older name eper/anus (104: 869).
Since that time opinions have differed widely as to how many recognizably distinct
units (species or subspecies) the eper/anus-dentex-mordax group actually includes.
One extreme among students concerned primarily with the smelts of North America
may be represented by Jordan, et a/., who recognized four distinct species (52: 67):
eperlanus (tacitly), mordax (with spectrum and abbotti as subspecies), dentex, and sergeanti.
The other extreme may be illustrated by Fowler’s union of mordax, including sergeanti,
8. The Smelt from Kamchatka which Pallas reported as spirinchus belong to the form that has subsequently been named
dentex.
Fishes of the Western North Atlantic 557
spectrum, and abbotti, with the European eper/anus, based on his comparison of smelts
from France with those from New York and from various localities in New England
(28: 555). As an example of extreme subdivision of the smelts of the eastern Atlantic,
Europe, and the Arctic Ocean, it seems enough to cite the following names applied
by Berg (5: 132): Osmerus eperlanus eperlanus, northern Spain to Scandinavia and the
Baltic; O. eper/anus dentex, White Sea and Arctic coast; O. eperlanus eperlanus natio
ladogensis, Russian lakes; O. eperlanus morpha spirinchus, lakes of northern Europe;
and O. eperlanus dentex natio dvinensis morpha spirinchus, lakes of the White Sea region
and Kola Peninsula.
The differing views in this regard after 1895 seem not to have reflected any
extensive contributions to recorded fact until 1934, when Petrow showed (77: 179, 182)
that smelts of the Arctic coasts of
Eurasia and of the northwestern Table I. Osmerus. Scales Penetrated by the Lateral Line in:
Pasihe: bh | 5 ip A, II specimens, 145-174 mm SL, from Nova Scotia and New
aetac : VCR AT a ee England (mordax), MCZ; B, 11, 72-206 mm, from Bering Sea
lateral-line scales (14-30) than (dentex), USNM; C, smelts from the White Sea, Arctic coast
smelts of Europe (4-14), that they of Siberia, and eastern Siberia, according to Petrov; D, 6, 67—
174mm, from northern Europe (eper/anus), MCZ; and E,
have somewhat larger teeth, and 4 : Be ieee
2 smelts from Russia (eper/anus), according to Petrow (77).
that on the average they have a
A A B iC D E
longer interspace between the bases
3 Scalestteyrsier. ssa 15-26 17-25 14-30 7-II 4-16
of the rayed dorsal fin and adipose Extreme manger SS go eee
fin. Consequently he recognized
two subspecies, eper/anus and dentex, not three, as Berg had done (4: 100, 102).
Unfortunately, Petrow did not extend his comparison in this regard to the smelts
of the western North Atlantic. But our own examination of the series listed (Tables 1
and 11) has shown that in this respect mordax agrees more closely with dentex than with
eperlanus, a result at which Hubbs had already arrived from his study of the number
of gill rakers and the height of the anal fin (43: 52). It also appears that the contrast
in number of scales is a decidedly consistent one, although we have found nothing to
suggest that the western Atlantic, eastern Atlantic, and Bering Sea smelts differ signifi-
cantly one from another in any of the other features that seem the most likely to be
significant from the species standpoint among salmon-like fishes as a whole.
If we can accept the number of lateral-line scales as our criterion, the smelts of the
western Atlantic are more closely related to the Arctic-North Pacific smelts than to the
European smelts, though the reverse might have been expected on purely geographic
grounds. And the separation between the two divisions seems sufficient to warrant the
recognition of two corresponding subspecies within the species eper/anus, but not of
three (eperlanus, mordax, and dentex), as by Wynne-Edwards (118: 17). Admittedly
the basis for this conclusion is so weak that future examination of more extensive series
of specimens from key localities may prove that it is not valid. But our provisional
adoption of it here (Key, p. 558) has at least the practical advantage of freeing us
from the need to expand the present account of the western Atlantic smelts to cover
the eastern Atlantic smelts in corresponding detail.
558 Memoir Sears Foundation for Marine Research
Table II. Osmerus. Proportional Dimensions and Meristic Characters for Three Major
Geographic Areas: A, 9 Western Atlantic Specimens from Cape Breton, Halifax, St.
John Harbor (Canada); Massachusetts Bay, Nahant, Charles River, Boston Harbor, and
Edgartown (Massachusetts); and New York. B, 19 Alaskan and Japanese Specimens
from Bristol Bay, St. Michaels, Port Clarence, and Wainwright Inlet, Alaska; Petro-
paulski Harbor, Kamchatka; and Mororan, Japan. C, 7 European Specimens from
the Elbe River; Le Havre, France; and Lake Oppmanna, Sweden.
Averages in Parentheses.
A B €
MAVEN TIF eb ode GOO bin DOOR BEE 97-198 (167) 104-206 (173) 67-174 (139)
Per cent of standard length:
[Hah 8 leplil oac6 odo baobOSaEODE 13.4—23.4 (17.9) 15.2—20.0 (18.3) 17.2—20.9 (19.3)
Head: length........ WSR ACES AE 20.3-24.9 (23.1) 22.2-26.9 (25.0) 23.8-26.6 (25.1)
Distance from:
snout to dorsal fin ........... 49.2—52.2 (50.9) 48.5-52.5 (50.1) §0.0-54.5 (51.7)
dorsal to adipose fin.......... 21.7-24.2 (23.3) 20.8—27.3 (23.4) 20.4—23.1 (21.8)
adipose to caudal fin ......... 8.7-10.8 (9.7) 7.8-10.7 (9.2) 9-5—10.9 (10.0)
analtolcaucdalfinliy- tris stere) as 6.0— 8.0 (7.1) 6.5-8.8 (7.5) 6.7— 8.7 (8.0)
Per cent of head length:
TBO CHESTER 66 soc o0a0cODEAC oT 18.9-23.3 (21.4) 17.8-—23.2 (19.6) 18.2—20.3 (18.8)
Distance from:
SNC 10) Gh oA WS snWod oo00 500 26.3-31.1 (29.2) 25.0-34.5 (29.3) 25.0-31.6 (28.9)
eye tommostril epyrerrctee sicratels aie Q.7—-12.2 (10.4) 9.3-12.7 (10.3) 7.9- 8.8 (8.5)
Oppersjaco wengihyeres sae ele: 44.8-54.2 (49.6) 47-5—59-8 (51.0) 43-8—-52.2 (50.1)
Counts:
Scales: perforated by lateral line. . 15—26* 17-25 3-10
total longitudinal series ... 64-69 64-69 59-63
Branchiostegals rays........0005. 7-8 7-8 7
Gill rakers: upper, first arch...... 8-11 g-10 g-I1
lower, first arch...... 17-21 17-22 17-21
DY LTA SESE) cosatdooed SAhOne IO-II 10-12 IO-II
ARALUITE SLAY Steretete ich ie loareierehala\= I5—17 15-17 IS—I7
WGCBIR ALAA: LAYS chiar. otaeterstatte tele 8 8 8
Pectoral frie 2\ TAY 83) sie ote isyas axatate + (ai II-12 II-I3 II-I2
* Only 4 specimens countable.
Key to Subspecies of Osmerus eperlanus
1a. Scales perforated by lateral line, 14—30. mordax Mitchill 1815, p. §59.
tb. Scales perforated by lateral line, only 4-16. eperlanus Linnaeus 1758.
Northern Europe.
Fishes of the Western North Atlantic 559
Osmerus eperlanus (Linnaeus) 1758
mordax Mitchill 1815
American Smelt, Icefish
Figures 131, 132
Study Material. Many specimens: 80, 67-230 mm SL, from various coastal
and estuarine situations along the coast of eastern North America from Cape Breton
(Nova Scotia) southward to New Jersey; numerous specimens from various land-
Ficure 131. Osmerus eperlanus mordax, adult, about 260 mm long (10.4 in.), from Woods Hole, Massachusetts.
Drawn by H. L. Todd.
locked situations in New England and New York state; 19, 104-206 mm long, from
Bristol Bay, St. Michaels, and Port Clarence in Alaska, Kamchatka, and northern Japan.
Also, for comparison, 7 specimens of O. eper/anus eperlanus from Le Havre, France;
the Elbe River; and Lake Oppmanna, southern Sweden; all in MCZ and USNM.
Distinctive Characters. Western North Atlantic fishes with which O. e. mordax
might be confused are Mallotus villosus (capelin, p. 576), a near relative, Argentina
(argentine, Part 4), or one of the silversides (family Atherinidae), to which it is only
remotely related. Its much larger scales, smaller adipose fin, and less projecting lower
jaw are enough to mark it off from M. villosus; its very much larger mouth and the
more forward position of its pelvic fins separate it from Argentina; and even the most
cursory examination should be enough to separate it from the silverside, for the latter
has two dorsal fins with the first supported by spines, no adipose fin, pelvics standing
considerably anterior to the first dorsal fin, and a mouth much smaller than that of
O. e. mordax.
Description. Based on 10 specimens: 9, 150-230 mm SL, and 1, 97 mm SL, from
several localities between Cape Breton (Nova Scotia) and southern Massachusetts.
Bopy elongated, moderately compressed (Fig. 131), the dorsal and ventral profiles
of head and trunk slightly convex, the dorsal profile a little more so than ventral profile;
560 Memor Sears Foundation for Marine Research
its depth 4.3—7.5 in SL, its greatest thickness about 1.7—2.4 times caudal peduncle depth
or 1.4—2.0 times distance from snout to eye. Least depth of CaupaL PEDUNCLE 1.0—
1.4 times in snout. ScaLes everywhere on trunk, extending onto base of caudal but not
onto dorsal or anal; 14.5 scales in oblique series from origin of pelvics to midline of
back; cycloid, thin, transparent, deciduous, with closely spaced circuli; scales along
midsection of sides with vertical axis about 1.5 times the horizontal axis and about equal
to diameter of pupil, those toward head and tail somewhat smaller and more roundish;
rear margin rounded, most abruptly so at horizontal axis posteriorly; head scaleless.
MobpIFIED LATERAL-LINE SCALES 14—30 (Table 1, p. §57), beginning at upper margin of
gill opening and extending rearward in a nearly straight line, ending somewhere be-
tween perpendiculars from rear part of pectorals and a little in front of dorsal; lateral
line ending with most posterior modified scale.
Heap 4.1—4.6 in SL, its dorsal profile straight or nearly so. SNour pointed, 3.2—
3.8 in head. Lower Jaw projecting. Eye 4.3—5.3 in specimens about 150 mm long
or larger, its rear margin about halfway between tip of snout and rear margin of opercle.
INTERORBITAL space very gently rounded, 3.5—4.3 (2.6 in 97-mm specimen). Post-
ORBITAL part of head 1.7—2.0 in head. Nosrrits situated in line with tip of snout and
upper edge of eye, and the two openings close together, the posterior one the larger,
its anterior rim bearing a flap, its distance from eye 8.1-12.2 in head. Opercie with
broadly rounded margin. Margin of Preoperc te about halfway between eye and margin
of opercle. BRANCHIOSTEGALS 7 or 8. GILL RAKERs very slender, the longest 1.4-1.9
in eye; 8-11 on upper limb of first arch, 17-21 on lower limb, total 25-32.
Mourtu oblique, the gape reaching almost or quite to a perpendicular from anterior
edge of eye. Maxittary beginning a little in advance of anterior margin of nostril,
widening rearward, its posterior margin broadly rounded and reaching from opposite
rear edge of pupil to just beyond eye. PremaxiLiary extending a little past origin of
maxillary. Manp1BLe 1.5—1.7 in head.
TeretTH. Lower jaw with one or two rows of small canines anteriorly, but one row
posteriorly where they are largest; maxillaries and premaxillaries with very small teeth,
unevenly spaced; vomer with one to two pairs of canines (sometimes 3 or § teeth),
these being among the largest in the mouth; palatines and mesopterygoid with a row
of low rather blunt teeth; tongue boat-shaped with fleshy lateral margin, with 1 or 2
strong fang-like teeth at the tip, followed by several progressively smaller teeth, and by
still smaller teeth behind these.
Dorsat with outer margin straight or nearly so, its base 1.2—1.5 times snout, its
origin about midway between tip of snout and caudal base, with 10 or 11 rays, the
first 2 unbranched, the longest 1.5—-1.8 times length of base. Apipose fin with outer
margin rounded, its base 2.2—3.4 times in base of dorsal, its origin over about middle
of anal base, it height 1.2-2.0 times in the base. Caupat deeply forked, its tips
pointed. Anat with outer margin straight or weakly convex, its base 1.3—-1.6 times
dorsal base, its origin about equidistant between origin of pelvics and base of caudal,
with 15-17 rays, the first 3 unbranched, the longest 0.6—0.7 of length of base. PEtvics
Fishes of the Western North Atlantic 561
with rear margin straight or nearly so, their origin opposite dorsal origin, with 8 rays,
the first (outer) unbranched (2 reported), the longest ray 3.3—4.0 times length of base.
Pecrorats with rear margin straight or slightly convex, the angles rounded, their
origin under rear margin of opercle, with 11 or 12 rays (13 reported), the first (upper)
ray unbranched, the longest ray 3.7—4.5 times the base.
VERTEBRAE 62—64 (ten specimens counted, but not the above described specimens).
Color.* Back and upper part of sides translucent, yellowish olive or bottle green,
sometimes with a brassy sheen; skin of upper parts, including fins, usually more or less
speckled with minute purplish black dots that show through the scales. Sides with a
broad bright silvery belt enclosing the lateral line, and outlined above by a narrow,
vaguely outlined dusky purplish band; lower part of sides less brilliantly silvery. Belly
and lower side of head as a whole, white. All the fins transparent, or nearly so; dorsal
with a paler shade of same color as back; the caudal similar but more or less dark-edged;
anal and pelvics whitish; pectorals colorless or nearly so, except for a greenish margin
anteriorly. The lining of body cavity silvery.
Size and Weight. American Smelts from salt water average about 7-9 inches long
when full-grown, and about 12—13 inches is the usual maximum. As caught, they
ordinarily run between 1 and 4 ounces, depending on size and fatness, and very large
ones may weigh as much as 6 ounces. In Great Bay, New Hampshire, the largest of
many measured was about g inches long. In the Bay of Chaleur, Quebec, 9- to 9.5-inch
fish averaged a little more than 0.2 of a pound, while a 10.5-inch fish (270 mm)
weighed a little more than 0.25 of a pound (122.5 g).
In some lakes where they are landlocked they commonly run up to 10-15
inches and (by report) up to as much as two-thirds to one pound or so in weight,
as in Sebago Lake and North Belgrade Lake in southern Maine and in some of the
Nova Scotian lakes, Canada. Other lakes, however, harbor stunted races, e.g. Grand
Lake in the St. Croix watershed of eastern Maine (56: 277), Lac des Isles in western
Quebec, and Lake St. John, tributary to the Saguenay.
Odor. Accounts have repeatedly credited the smelts with a “cucumber’’ odor,
especially smelts of Europe, which have often been described as strong-smelling. But
in the American populations this odor is so faint that we have not noticed it, though
we have caught and handled many.
Development and Growth. Females weighing no more that two ounces may produce
as many as 40,000—50,000 eggs (85: 188); one 232 mm (9.12 in.) long, taken in
Crystal Lake, Michigan, contained 43,125 eggs (59: 142). The eggs, which range in
diameter from 0.6 mm to about 1.2 mm in different waters and according to different
authorities, sink to the bottom where they stick to each other in clusters or cling to
any object on which they settle. In European waters, the eggs hatch in 8-27 days,
depending on the temperature of the water. In Massachusetts they have been reported
as hatching in 13 days.
The larvae are about 5—6 mm long when they hatch and are perfectly transparent
g- From market specimens in excellent condition that we have examined recently.
36
562 Memoir Sears Foundation for Marine Research
at first. Once hatched they rise close to the surface and drift downstream. On the average,
they grow to 17 or 18 mm during their first month, to 27-34 mm during the second
month, and to about 40 mm after 3.5 months, when we have seined several hundred
fry of 30-40 mm at Mt. Desert Island, Maine; and the schedule is about the same
in European waters, the precise rate of growth depending on temperature.
By the time larvae of the Elbe River in Europe have grown to 8 mm, the yolk sac
is mostly absorbed; at 15 mm all the fins are more or less developed, but the caudal
is still rounded; and by 45 mm the formation of scales has begun. We see no reason
to suppose that development proceeds otherwise in America.1°
In their second spring, when one year old, the fry average about 86 mm (data for
Great Bay, New Hampshire). From scale studies it appears that they average as follows:
a Sars ces =e
Ficure 132. Osmerus eperlanus mordax, larva, 26mm long, from Portland Harbor, Maine, July 31, 1912.
Drawn by Mrs. E. B. Decker.
at two years about 145 mm (5.7 in.) and about 0.6 of an ounce, at three years 171 mm
(6.7 in.) and about 1.1 ounces, at four years 220 mm (8.7 in.) and about 2 ounces.
The largest measured was 230 mm (about g in.). At this locality at least, four or more
year-classes are represented in the commercial catches. Those taken in Crystal Lake,
Michigan, where they were introduced, grow somewhat faster, reaching 6—8 inches
at two years, up to 9 inches at three years, up to ro inches at four years, and 10.75-
12 inches at five years (17: 401-424), with a maximum recorded length of 14 inches,
probably for a six-year-old. Doubtless they grow much more slowly in other lakes that
harbor stunted races.
Spawning Habits. The marine fish normally spawn in fresh water, and as a rule
they do not travel far upstream; they may go only a few hundred yards above the head
of tide. Some spawn in the tidal zone and some even spawn in brackish water behind
barrier beaches. But they never spawn in salt water, for flooding with the latter, as some-
times happens, kills the eggs. They spawn on pebbly bottom where there is a current,
and often in water only a few inches deep. Most often the spawners are two years old,
or older. Spawning takes place in late winter or early spring, depending on the temper-
ature of the water. According to data from hatchery operations, the chief production
of eggs takes place in temperatures of 50°—57° F in Massachusetts, and of about 45°—
so°F in Grand River, Quebec, representative of the northern part of their range.
The spawning period lasts 1o-14 days and is completed ordinarily by mid-May in
10. For excellent accounts and illustrations of the smelt of the Elbe River, see Ehrenbaum (25: append. 3, pl. 1), whose
illustrations have been reproduced by Kendall (56: fig. 25). For the embryology and early larval development of
the American Smelt, see Rice (86: 57-76, pls. 1-5).
Fishes of the Western North Atlantic 563
Massachusetts, not until late May along the coast of Maine, and not until some time
in June in the Gulf of St. Lawrence.
The spent fish—except those that die, as many do—drop down again to brackish
or salt water immediately after spawning, so that all of them have deserted fresh
water by the middle of May in Massachusetts, nearly as early in Maine (where we
have seen a bushel of large fish taken as early as May 4 from a weir near Cutler at
the mouth of the Grand Manan Channel), and by the end of June at the latest in the
Gulf of St. Lawrence.
Habitat and Migrations. Smelt, whether American or European, make their growth
in brackish or salt water if they are not landlocked. During the marine phase of their
life they are confined to so narrow a coastal belt that none has ever been reported
more than six miles or so out from the land and seldom below 2-3 fms.; the deepest
record for them is 9-10 fms. at the mouth of Port-au-Port Bay on the west coast
of Newfoundland, where 185 specimens were taken in a half-hour haul with an otter
trawl. Many of them spend their entire growth period in estuarine situations, including
the tidal reaches of rivers.
Their habitat in the summer along any particular section of the coast appears
to depend chiefly on the temperature of the water. From Massachusetts southward,
most of them (though not all) desert the harbors and similar situations during the
warmest season, moving, it seems, only far enough out and deep enough to find slightly
cooler water. Along the coasts of Maine and the Maritime Provinces of Canada, how-
ever, where water temperatures rule lower, they are found in the harbors, bays, and
estuaries all summer.
With the onset of autumn, those that have moved out to sea re-enter the harbors
and estuaries, so that by mid-October, or early November at the latest, practically the
entire population is concentrated there. In our experience the smaller ones tend to
reappear the earliest, but reports are contradictory in this respect. At any rate, by
December some have even worked up into stream mouths to the head of tide. But the
fish fated to breed that season, most of which are two years old or older, do not actually
enter fresh water until late winter or early spring when the water off the mouth of the
stream has warmed to at least 39°-42° F (4°-5.5° C), if it has chilled below that figure,
and until the ice has gone out of the stream, if any has formed there during the preceding
winter.
The movement of the maturing fish into fresh water commences late in February
along the southern coast of New England and southward, some time in March along
northern Massachusetts, seldom until April along the eastern part of the Maine coast,
and not until the latter half of May along the southern shores of the Gulf of St. Lawrence.
This species, though confined to very shoal water, is not a bottom fish but tends
to hold position at some intermediate level. The small ones, and probably the large ones
also, gather and travel in schools that are composed for the most part of fish of about
the same size, i. e. the product of one year’s hatch. In the smaller harbors they tend to
move in and out with the tide, especially if the tidal flow is strong. But while they
Bor
564 Memoir Sears Foundation for Marine Research
may be plentiful close in to the tideline, few of them run up into salt marsh creeks
unless these creeks receive some inflow of fresh water; we have never heard of them
being trapped on the flats by the outgoing tide, as happens now and then to herring.
Although they are not Arctic fish, we have never heard of their being killed by winter
chilling, as in the case of some other fishes from time to time.
Their actions, if these are watched from some raft or float, suggest that they travel
but little during their inshore sojourn, except as they may be carried to and fro by
the tide, as just remarked. Anglers have long known that American Smelt do not dart
at bait, but approach it slowly and appear to suck it into the mouth rather than seize
it, as most of the predaceous fishes do.
Food. This species is carnivorous and predaceous. In salt and brackish water, shrimps
(decapod and mysid) probably are their chief support on the Massachusetts coast; sim-
ilarly, the stomach contents of those in the Gulf of St. Lawrence have consisted chiefly
of copepods, amphipods, and mysids, with algal debris probably taken incidentally.
In some localities small fish rank next. We have found them packed full of young
Atlantic herring, on the coast of Maine, and a wide variety of fishes has been recorded
as occurring in their stomachs, at Woods Hole. They also take small shellfish, small
squid, annelid worms, and small crabs as occasion offers. But they cease to feed during
the spawning season, as many other fishes do.”
Enemies, Disease, and Mass Fatalities. Doubtless the marine population is preyed
upon by a wide variety of the larger predaceous fishes, but we have no definite informa-
tion in this regard for the American coast (see 63: 236 for the European smelt). In
American lakes they are perhaps the favorite prey of landlocked salmon and lake trout
and are commonly used as bait for these.
In European harbors, mass destruction of them, from some unknown cause, takes
place from time to time (63: 39-42).
A mortality of smelt, affecting all sizes and both sexes, occurred in Lake Huron
and in Lake Michigan during the fall and winter of 1942-1943 (72: 310-337), with
the first indication of a recovery being apparent in 1945. It was concluded that this
mortality, which all but wiped out the populations in these two lakes, could be ex-
plained only as resulting from a communicable disease (bacteria or virus). We have
found no record of any happening of this sort in brackish or salt water along the
American coast.
Variations. A separate specific name, sergeanti, was given years ago to American
Smelt of the streams tributary to the head of Delaware Bay because of their smaller size
and a slight difference in color (Norris, 69: 59; 70: 93). Hubbs recognized sergeant
as at least a ‘“‘race’’ because of the fewer gill rakers on the first gill arch and because of
a slightly higher anal fin than on the more northerly fish (43: 52). Similarly, Rice
long ago reported small differences in color between those of the Raritan River, New
Jersey, and those of more northern localities (86: 79), though he did not consider
those differences wide enough for recognition in nomenclature.
11. For their diet in fresh water, with extensive tables for many different lakes, see Kendall, 56: 281-287.
Fishes of the Western North Atlantic 565
It has been found that American Smelts average somewhat larger at maturity in the
Bay of Chaleur, Quebec, than in the estuary of the St. Lawrence River (64: 71-85),
and that those of the Bay of Chaleur have a larger number of vertebrae on the
average (62.18—62.9 according to locality) than those from various places between
Matane on the southern side of the estuary and Quebec City (60.32-61.92). The
first of these differences may be associated with the food supply; the second illus-
trates the long-known rule that among fish species that occur over a wide range
of temperatures, the number of vertebrae average the most numerous among popu-
lations that pass their growth stages in colder than in warmer water. Nothing else
is known in this regard for any other parts of the American coast. For the American
Smelt in fresh water we know only that they run much larger in some lakes than in
others.
Racial studies of the subspecies mordax of the White Sea region, Arctic Asia,
and eastern Siberia by Petrow (as dentex) did not reveal any regional differences call-
ing for recognition in nomenclature (77: 177-188). For the European subspecies
eperlanus, Hubbs (43: 52) has pointed out that the gill rakers average more numerous
in specimens from Scandinavia (32-37) than in those from the English Channel
(25-32). And Jensen has reported slight differences, based on extremely detailed com-
parisons of smelts from different lakes in Denmark, especially in the number of verte-
brae (49: 73—109).1?
Abundance. In colonial days, according to Captain John Smith, American Smelt in
1622 were in “‘such abundance, that the Salvages doe take them up the rivers with
baskets, like sives” (56: 244). While it seems certain that they were far more plentiful
along the coasts of New York and Maine up to the early part of the nineteenth
century than they are today, by 1849 a decrease was reported for northern New
Jersey, and by 1885 for the streams of Long Island, New York. By the end of
the century, at Cold Spring Harbor, the New York State Hatchery, which planted
48 million fry in 1898, produced only a few million in 1899 for lack of ripe females
and eggs from nearby waters, and after 1900 its operations were discontinued. Many
reports of their depletion in the waters of Massachusetts have been received also;
there the number of eggs from the Weir River, tributary to Boston Harbor, fertilized
at the Palmer Hatchery, fell from around too and 11o million in 1918 and 1919 to
32 million in 1920, after which Smelt hatching was given up.'® A general decrease
seems also to have taken place along the coast of Maine since the end-of the last century,
and from the same causes that produced the depletion southward, i. e. from contraction
of favorable spawning grounds by dams close to the mouth of streams, and perhaps
from overfishing. In 1889, for example (15: 79), United States waters yielded 1,209,385
pounds of American Smelts in contrast to a yearly average of only 473,550 pounds
for the period 1951-1954. But the catch records do not suggest any extensive or
12. For a recent survey of variations in the taxonomic features of fishes in general, as associated with regional dif-
ferences in their environment, see Vladykov (rrj: 122).
13- For details and for general information as to the smelt, see 84.
566 Memoir Sears Foundation for Marine Research
general alteration in their abundance during the past 15 years or so for either Maine
or Canada, except locally, as in the Miramichi area, Canada, where their access to
fresh water has been hindered by obstructions in the streams. On the contrary, the
yearly Canadian catch increased from an average of only 8,000,000 pounds for the
20-year period 1918-1937 (65) to 21,295,000 pounds for the later period 1951-
1954. This increase doubtless is indicative of an increasing demand for them rather
than of any corresponding increase in their abundance.
Artificial Propagation. American Smelt have been a favorable subject for artificial
propagation.'* Many million fry were hatched in past years at the Cold Spring Harbor
Hatchery, New York, as well as at the Palmer Hatchery, Massachusetts (p. 565). The
results have been widely heralded, for great catch increases were reported for streams
where fry were released. The most notable example is that 32 million eggs were col-
lected in 1885 from a New York stream where there had been no Smelts for at least
some years previous (85: 188). And a similar example, though less spectacular, was
reported for Massachusetts.
Relation to Man. The American Smelt is a favorite among the market fish, deli-
cious when fresh-caught or even after being iced properly, and great numbers, especially
from the Gulf of St. Lawrence, are marketed. The average landings reported for the
four-year period 1951-1954 were 5,323,000 pounds for the Canadian Atlantic coast
and 150,700 pounds for the United States coast, or a total of 5,473,700 pounds; that
is, 55 million individuals if these ran say, 10 to the pound, all marketed for human
consumption. Years ago they served as cod bait in the Gulf of St. Lawrence (60: 224),
and large quantities were used yearly as manure along the Gulf of St. Lawrence
shores of New Brunswick (76: 135).
These fish also have great recreational value. As many as 2,336 anglers have been
counted ‘‘smelting’’ about Houghs Neck in Boston Harbor at one time, and the same
sort of thing is to be seen on a smaller scale throughout their range. On occasion, espe-
cially on a rising tide, the fish bite greedily. Sea worms (Vereis) are the best bait, espe-
cially for larger fish, salt-marsh shrimps (Crago) a second best, and small fish a poor
third. They have also been reported as taken with a small red artificial fly in the Gulf
of St. Lawrence (60: 225), and doubtless they might be taken with a fly elsewhere. On
the other hand, anglers prefer American Smelt to other bait for landlocked salmon in
lakes of northern New England and of the Maritime Provinces, Canada, wherever they
are to be had.
Methods of Capture. The commercial catch is made chiefly in nets of various kinds,
but partly with hook-and-line. For example, of the 1954 Maine catch, which may serve
as a representative year, about 31 °/, were taken in haul-and-stop seines, about 26 °/,
in gill nets, about 21 °/, in bag nets operated through the ice, about 4°/) in weirs, and
about 18 °/, on hook-and-line. In New Hampshire, however, in that same year, 3 °/) of
the commercial catch was made in bag nets, 23 °/, in weirs, and 74 °/, on hook-and-
line. Along the New Brunswick shoreline of the Gulf of St. Lawrence, where the
14. For methods employed, see especially Rice, 86: 41-56; also 85: 188.
Fishes of the Western North Atlantic 567
catch is much larger than anywhere in United States waters, about 95 °/) are taken in
winter through the ice, in trap nets of one kind or another.
General Range. Western North Atlantic coast, from the outer coast of Labrador
at about 54°N, and northern Newfoundland, southward to the head of Delaware Bay;
also (as dentex) in the White Sea, Arctic coasts of Europe and Asia at scattered localities,
northwestern Pacific south to northern Japan and northern China (type locality of
dentex), Pacific Bering Sea and Arctic coasts of Alaska northward from Yakutat Bay,
northern British Columbia (36: 55) to Bering Strait, and eastward thence on the
Arctic coast of Canada to the delta of the Mackenzie River, where they are reported
to be plentiful at the mouth of the Arctic Red River. They are also landlocked in many
lakes and rivers.
The European O. eper/anus eperlanus range from Vigo in northern Spain north-
ward to southeastern Norway, and up the Baltic to the Gulfs of Finland and Bothnia,
chiefly in the watersheds of the Baltic and North seas. They have not yet been reliably
reported for Iceland. Like some American Smelts, they are landlocked in many lakes
and river systems of foreign places.¥®
Occurrence in the Western North Atlantic. The most northerly locality of positive
record for American Smelts in eastern North America is the mouth of Northwest
River, tributary to Lake Melville at the head of Hamilton Inlet on the Atlantic coast of
Labrador (about 54°N), where Low described them as “abundantly taken” in Novem-
ber and early December (62: 329); and they have recently been recorded for various
places in the Hamilton Inlet-Lake Melville region (3: 295). The next most northerly
locality is the Newfoundland side of the Strait of Belle Isle, where, writes Jeffers,
“a few were generally observed each summer” (47: 207). We have found no proof
of their presence anywhere, passing westward, along the north shore of the Gulf of
St. Lawrence until nearing the Bersimis River, whence small commercial catches are
reported. But they are plentiful near Quebec City, and at least one specimen has been
reported to be as far up the St. Lawrence River as Lake St. Pierre (18: 20). They
are caught all along the south shore of the St. Lawrence estuary. They have been
reported for Anticosti Island and are taken regularly in commercial quantities
around the Gaspé Peninsula and Magdalen Islands. In the shallow, semi-enclosed
bays of Prince Edward Island and on the north shore of New Brunswick Harbor there
occur what are probably the greatest concentrations of American Smelt in the western
side of the Atlantic. Many years ago they were described as ascending the smaller
streams of Newfoundland “in thousands as soon as the ice disappears” (Reeks, 82:
2556), and they have recently been reported by the Newfoundland Fishery and Research
Commission as plentiful there in the shallow bays and river mouths on both the
west coast at the mouth of the Humber River and Port-au-Port Bay and the east coast
in the rivers of Notre Dame Bay. They were also described long ago as common
around the wharves at Cape Breton, Nova Scotia.
American Smelt are familiar thence southward along the coasts of outer Nova
15. For recent summary, see Jensen, 49: 102.
568 Memoir Sears Foundation for Marine Research
Scotia and northern New England wherever local surroundings are favorable, and they
are widespread along the shores of southern New England, as well as New York,
where the stream mouths of Long Island formerly were centers of abundance (p. 565).
Their range also includes northern New Jersey—the Raritan River in particular; and
there is a reliable old record of them in the lower reaches of streams and rivers of
Delaware and Pennsylvania, at the head of Delaware Bay (Schuylkill River and Brandy-
wine Creek in particular), and as far up the Delaware River as the swift water at Trenton,
New Jersey (70: 94); but it 1s questionable whether there are any survivors today
of this southernmost population. We have found no reliable basis for the repeated
statements that they range southward to Virginia.1®
They differ widely in numbers from place to place, even within short distances,
depending on how well the local surroundings meet their rather exacting requirements.
Thus Great Bay, New Hampshire, is a center of abundance, perhaps second only—
considering its size—to the much more extensive shoal bays on the north shore of New
Brunswick. But the inner part of the Bay of Fundy on the New Brunswick side is a
region of great scarcity.!”
There are progressively fewer American Smelts southward along the coast as a
whole. This gradation may be illustrated for recent years by average landings that
were about 25-40 times greater for the Gulf of St. Lawrence shore of New Brunswick
than for the longer Maine shoreline. For 1899, when the status of American Smelts
represented natural conditions more nearly than their present status, the catch for
Maine was 1,055,000 pounds, but only 154,000 pounds thence southward along the
coast of New England. Westward and southward from Connecticut the commercial
catches are negligible, and seemingly always have been.
Occurrence in Fresh Water. \t has long been a matter of common knowledge that
some American Smelts are also landlocked. In eastern North America there were in-
digenous populations in Long Lake, near Plymouth, Massachusetts (whence the Mu-
seum of Comparative Zoology received 10 specimens in 1875).!8 They occur in some
lakes in New Hampshire (Squam, Winnisquam, and Winnipesaukee) and in many
of the coastal lakes and ponds of all the principal river basins in Maine.!* There
are native Smelt in: many of the Nova Scotian (61: 29) and New Brunswick lakes;
Lake Memphremagog (partly in Vermont and partly in Quebec); Lake Champlain
(in abundance); Lake St. John, eastern Quebec; Lac des Isles, Green Lake, and
Lac Brulé in western Quebec northwest of Montreal; and in Muskrat and Doré lakes
of Ontario, which are tributary to the upper Ottawa River, at an altitude of about
3,000 feet above sea level (doubtless as a “relic of the time following the retreat of the
last ice sheet, when an arm of the sea extended up the Ottawa Valley to the head of
Lake Timiskaming”; Dymond, 23: 59). No doubt there are native Smelts in various
16. Based seemingly on Jordan and Gilbert (53: 293).
17. For discussion, see Jeffers, 46 (1932): 27.
18. Perhaps elsewhere on Cape Cod (locality not specified), to judge from an article in Forest and Stream for April 18,
1889.
tg. For the Smelt lakes of Maine, see Kendall, 56: 273-278.
Fishes of the Western North Atlantic 569
other Canadian lakes, but we have not chanced to hear of them. We have not found
any report of landlocked Smelt for Newfoundland.
There are smelts of the subspecies mordax in lakes of the White Sea region, the
Kola Peninsula (5: 132, as eperlanus dentex natio dvinensis morpha spirinchus), and
eastern Siberia. In Europe the subspecies eper/anus is widespread in lakes of Germany,
Denmark, southern Norway, Sweden, Finland, and Russia; even in the Volga River
and the Caspian Sea there are smelts, as has long been known (57: 241).
Artificial Introductions. It has been a matter of scientific record for more than a
century and a half that the introduction of smelts from one lake to another is pos-
sible if the new situation is favorable for them (8: 182), and on various occasions
they have been introduced successfully from salt to fresh water. The literature on this
species includes repeated references to such introductions on both sides of the At-
lantic, either by the transfer of fertilized eggs or by the transport and release of adult
fish. In America, the earliest recorded case was near the close of the 18th century,
when they were successfully introduced into Jamaica Pond near Boston (83: 20),
whence the Museum of Comparative Zoology received a series in 1875. As our concern
here is primarily with the marine form, it is enough to add that introductions? account
for their presence today in certain lakes of New Jersey, the Rangeley Lakes, Moose-
head Lake, the upper Penobscot River, and Eagle Lake in Maine, and the Great
Lakes region. They are now (1957) so plentiful in Lake Erie that catches taken there
are regularly on sale in Boston markets.
Synonyms and Selected References:
Salmo eperlanus Pennant, Arctic Zool., r, Introd., 1784: 127 (name, Kamchatka); Mitchill, Rep., in part, on
Fishes of New York, 1814: 12 (New York; earliest rep. in e. N.Amer.); Trans. Lit. philos. Soc. N.Y.,
T, 1815: 435 (color, anal rays, Passaic R., New Jersey); Gmelin, Naturg., Fische, 1818: 298 (freshw.
lakes, Prussia).
Salmo (Osmerus) eperlanus (in part) Pallas, Zoogr. Rosso-Asiat., 3, 1831 :*1 386 (descr., Baltic and White seas,
Kamchatka, Sea of Okhotsk; includ. subspp. eper/anus and mordax).
Salmo (Osmerus) spirinchus (in part) Pallas, Zoogr. Rosso-Asiat., 3, 1831 :242 387 (descr. includ. smelts of Europ.
Russia and e. Siberia, subspp. eper/anus and mordax).
Atherina mordax Mitchill, Trans. Lit. philos. Soc. N.Y., I, 1815: 446 (descr., adipose fin noted).
Salmo eperlano marinus Gmelin, Naturg., Fische, 1818: 299 (North Sea, Baltic).
Osmerus viridescens LeSueur, J. Acad. nat. Sci. Philad., 1818: 230 (descr., Boston, Massachusetts, to Newport,
Rhode Island); DeKay, Zool. N.Y., 3, 1842: 243; 4, 1842: pl. 39, fig. 124 (descr., New York); Storer,
Mem. Amer. Acad. Arts Sci., N.S. 2, 1846: 449, 450; also separate, Cambridge, 1846: 197, 198 (brief
descr., cf. Europ. smelt; Maine, Massachusetts, New York); Cuvier and Valenciennes, Hist. Nat. Poiss.,
21, 1848: 388-389 (diagn., New York, cf. Europ. eper/anus); Perley, Cat. [in part] Fish. N. Brunsw.,
Nova Scotia iv Rep. Fisher. Bay of Fundy, ed. 1, 1851: 1353; Descr. Cat. [in part] ..., ed. 2, 1852: 202;
Descr. Cat. [in part] ..., as separate, ed.2, 1852: 24 (size, abund., odor, cod bait and manure, feed.,
capture); Rep. Fisher. Gulf of St. Lawrence, ed. 2, 1852: 22, 23 (rivers, abund., capture, spawn., 2 spp.);
Bell, Canad. Nat. Geol., 4, 1859: 206 (Gulf of St. Lawrence); Fortin, Rep. Magistr., Exped. Protect.
Fisher., Gulf of St. Lawrence (1861, 1862), 1863: 120 (Quebec City; not seen); Norris, Proc. Acad.
nat. Sci. Philad. (1861), 1862: 58-59 (diffs. between smelt of Schuylkill R. and north. viridescens);
Giinther, Cat. Fish. Brit. Mus., 6, 1866: 167 (Boston and New York; scarcely distinct from eper/anus) ;
Storer, Fish. Mass., 1867: 149, pl. 25, fig. 4 (descr., size, abund., Massachusetts); Reeks, Zoologist,
20. For details, see Kendall (56: 351), Dymond (24: 12, 13), and Van Oosten (71: 64, 65).
21. Date of publication probably was 1814, according to Sherborn (99: 164-167).
570 Memor Sears Foundation for Marine Research
London, (2) 6, 1871: 2556 (abund., Newfoundld.); Adams, A. L., Field Forest Rambles, London,
1873: 243 (spawn., distr., abund., New Brunswick coast, streams, food for pigs); Cox, Bull. nat. Hist.
Soc. New Brunsw., 3 (13), 1895: 66 (abund. New Brunswick, landlocked in lakes); Montpetit, Poiss. d’eau
douce Canada, 1897: 6, 477 (Quebec City, Trois Riviéres, Quebec; not seen).
Osmerus eperlanus (in part) Smith, Nat. Hist. Fish. Mass., 1833: 147 (habitat, abund.); Richardson, Fauna
Boreal.-Amer., 3, 1836: 185 (Halifax, N.S., New England; like Europ. smelt, but origin of pelvics
farther rearward, under 6th D. ray; descr. Europ. smelt); Storer, J. Boston Soc. nat. Hist., 2, 1838:
432 (descr., abund. Massachusetts); Rep. Fish., Reptiles, Birds, Mass., 1839: 108 (descr., local abund.,
refs.); Liljeborg, Svenska VetenskAkad. Handl. (1850), 1851: 304 (Archangel); Perley, Rep. Fisher. G. of
St. Lawrence, ed. 2, 1852: 23 (see under O. viridescens above) Rice, Rep. Comm. Fish. Md., 1878: 41-94
(hatchery methods, develop., Raritan R., New Jersey, variations, scales show Amer. smelt same as Europ.
smelt, refs.); Smitt, Hist. Scand. Fish., ed. 2, 2, 1895: 869, 872 (mordax synon. of eper/anus); Knipo-
witsch, Ann. Mus. Zool., Acad. imp. Sci. St. Petersb., 2, 1897: 153 (White Sea region); Fowler, Proc.
Acad. nat. Sci. Philad., 63, 1912: 555 (mordax united with eper/anus, cf. French and Amer. specimens).
Osmerus spirinchus Liljeborg, Svenska VetenskAkad. Handl. (1850), 1851: 304 (White Sea); Smitt, Svenska
VetenskAkad. Handl. (1885), N.F., 2z [8], 1886: 168, 189 (White Sea; cf. proport. dimens. for spi-
rinchus, dentex, eperlanus); Hist. Scand. Fish., ed. 2, 2, 1895: 867 (name for Arctic smelts), 868 (distinct
from eper/anus and dentex).
Osmerus mordax Gill, Proc. Acad. nat. Sci. Philad. (1861), Append., 1862: 53 (in synop., fishes e. coast N.
Amer.); Adams, Field Forest Rambles, London, 1873: 306 (freshw., New Brunswick); Baird, Rep.
U.S. Comm. Fish. (1871-1872), 1873: 826 (Woods Hole, Massachusetts); Gill, Rep. U.S. Comm.
Fish. (1871-1872), 1873: 810 (e. coast U.S.); Lanman, Rep. U.S. Comm. Fish. (1872-1873), 2,
1874: 224, 225 (season, abund., Bay of Chaleur, n. New Brunswick, cod bait, capture with artif. fly);
Jordan, Manual Vert. north. U.S., ed. 1, 1876: 261; and later edits. (descr., range); Goode and Bean,
Bull. Essex Inst. Salem (1879), rz, 1880: 23 (abund., season, Essex Co., Massachusetts; cf. Europ.
eperlanus); Bean, Proc. U.S. nat. Mus. (1880), 3, 1881: 105 (Maine, Massachusetts, Connecticut);
Jordan and Gilbert, Bull. U.S. nat. Mus., 76, 1882: 293 (descr., Virginia northward); Goode, et a/.,
Fish. Fish. Industr. U.S., 1 (3), 1884: 543, pl. 199 (range, habits, early abund., fishery); Stearns,
Proc. U.S. nat. Mus. (1883), 6, 1884: 124 (abund., C. Breton, Nova Scotia); Bean, Fish. Penn., 1893:
64 (Pennsylvania); Jordan and Evermann, Rep. U.S. Comm. Fish (1895), 1896: 294 (Virginia to Gulf
of St. Lawrence; also Lake Champlain and L. Memphremagog); Bull. U.S. nat. Mus., 47 (1), 1896: 523
(descr., range); Low, Ann. Rep. geol. Surv. Canada, App. 111 to Rept. L, N.S. 8, 1896: 329 (Northwest
R., Lake Melville, head of Hamilton Inlet, Labrador); Anon., Rep. U.S. Comm. Fish. (1897), Append.,
1898: 188 (no. and descr. of eggs, culturing procedure, artif. propag., Cold Spring Harbor, Long Island,
New York); Bean, 7th Rep. Forest Comm. N.Y. (1gor), 1902: 256, 351 (New York locals., spawn.,
eggs, food); Bull. N.Y. St. Mus., 60, Zool. 9, 1903: 282 (descr., size, distr., bait and food for larger
salmonoids, habits, season in N.Y., artif. propag. New York, New Jersey, Pennsylvania; lakes); Schmitt,
Monogr. Isle d’Anticosti, 4, Zool., Poiss., 1904; 285 (common, Anticosti; plent. n. coast, Esquimaux Pt.);
Fowler, Rep. N. J. St. Mus. (1905), 1906: 115 (descr., ill., New Jersey, Delaware; sergeanti Norris
1868 a synon.); Tracy, 36th Rep. R.I. Comm. inl. Fish., 1906: 57 (season, abund., food, max. size,
Rhode Island); Evermann and Goldsborough, Proc. biol. Soc. Wash., 20, 1907: 105 (St. Lawrence R.
to Quebec City, Trois Riviéres; landlocked, New Brunswick and Quebec); Fowler, Amer. Nat., 47, 1907:
7 (Schuylkill R. and Delaware R.); Kendall, Occ. Pap. Boston Soc. nat. Hist., 7 (8), 1908: 48 (New
England locals.); Proc. Portland nat. Hist. Soc., 2(8), 1909: 221 (C. Breton, Nova Scotia, by ref. to
Stearns, 1883); Tracy, goth Rep. R.I. Comm. inl. Fish., rg1o: 84 (descr., Rhode Island); Sumner, Os-
burn, and Cole, Bull. U.S. Bur. Fish., 3r (2), 1913: 743 (Woods Hole region, parasites); Halkett,
Check List Fish. Canada, 1913: 55 (genl. range, Lac des Isles, Gatineau R. system, Quebec); Cockerell,
Bull. U.S. Bur. Fish. (1912), 32, 1914: 136 (scales descr.); Fowler, Proc. Boston Soc. nat. Hist., 35,
1917: 112 (Maine, Massachusetts locals.); Latham, Copeia, 1917: 18 (season, Orient, New York);
Wilson, Proc. U.S. nat. Mus., 53, 1917: 59 (copepod parasite, Woods Hole, Massachusetts); Hunts-
man, Trans. roy. Soc. Canada, (3) 12 (4), 1918: 63 (Gulf of St. Lawrence, warm surface water layer);
Fowler, Proc. biol. Soc. Wash., 32, 1919: 56 (tidal streams, head of Delaware Bay); Proc. biol. Soc. Wash.,
33, 1920: 149 (New Jersey locals., lakes where introduced); Huntsman, Contr. Canad. Biol. (1921),
3, 1922: 60 (Bay of Fundy locals., landlocked, Chamcook and Utopea lakes, New Brunswick); Linton,
Proc. U.S. nat. Mus., 64, 1925: 71, 102 (cestode parasites); Bigelow and Welsh, Bull. U.S. Bur. Fish.
(1924), 40 (1), 1925: 143 (descr., ill., habits, abund., develop., commerc. import., Gulf of Maine);
Fishes of the Western North Atlantic 571
Creaser, Pap. Mich. Acad. Sci., 5, 1925: 405 (introd. into Great Lakes, tributary waters; spawn., growth,
food, reactions to light and current); Hubbs, Proc. biol. Soc. Wash., 38, 1925: 52 (cf. Pacific dentex);
Manter, Illinois biol. Monogr., 10, 1926: 8 (trematode and nematode parasites); Kendall, Bull. U.S.
Bur. Fish. (1926), 42, 1927: 217-385 (genl. acct., special ref. to freshw., name); Nichols and Breder,
Zoologica, N.Y., 9 (1), 1927: 47 (abund., New York; Woods Hole); Breder, Field Bk. Mar. Fish. Atl.
Cst., 1929: 75 (listed, ill.); Scheuring, Ergebn. Biol., 5, 1929: 654 (summ. of nat. hist. w. N. Atlan-
tic); Bere, Contr. Canad. Biol., N.S. 5 (13), 1930: 423 (copepod parasite, Bay of Chaleur, Quebec);
Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 67 (Virginia to Gulf of St.
Lawrence); Creaser, Copeia, 1932: 157 (recent spread, Lakes Huron and Michigan); Wilson, Bull.
U.S. nat. Mus., 158, 1932: 17, 377, 480 (copepod parasites, Woods Hole); Anon., Rep. Nfdld. Fish.
Res. Comm., I (4), 1932: 107 (Newfoundld.); Anon., Rep. Nfdld. Fish. Res. Comm. (1932), 2 (1),
1933: 125 (depth of capture, numbers, w. Newfoundld.); Mason, Copeia, 1933: 34 (Lake Ontario,
1931); Vladykov, Trans. Canad. Inst., 20, 1934: 122 (landlocked, Ontario); Langlois, Copeia, 1935: 141
(spawn., size, no. eggs, Crystal Lake, Michigan); Vladykov and Tremblay, Nat. Canad., 62, 1935: 79
(Trois Pistoles, Quebec); Hoover, Copeia, 1936: 85 (spawn., habits, sex ratio, nuptial tubercles of male,
Lake Winnisquam, New Hampshire); Vladykov and McKenzie, Proc. N.S. Inst. Sci., 19 (1), 1936: 58
(distr., abund., commerc. catches, Nova Scotia); Dymond, Copeia, 1937: 59 (Ontario lakes); Pigeon and
Vallée, Nat. Canad., 64 (2), 1937: 38 (food, Canada); Van Oosten, Copeia, 1937: 64 (early recs.,
Lake Erie); Bailey, Fish iz Surv. Merrimack Watershed, Rep. N. H. Fish Game Dept., 3, 1938: 162
(Merrymeeting and Squam lakes, New Hampshire); Schrenkeisen, Field Bk. Fresh-w. Fish. N. Amer.,
1938: 71 (diagn., ill., habits, landlocked, subspp.); Dymond, Contr. R. Ontario Mus. Zool., 75, 1939:
18 (native, Ontario lakes); Chapman, J. Morph., 69, 1941: 279-301 (skel. chars., rel. of Osmeridae) ;
Linton, Proc. U.S. nat. Mus., 88, 1942: 34, 68, 126, 156 (trematode parasites; Woods Hole); Beckman,
Copeia, 1942: 120 (length-weight, age, sex ratio, food, Crystal Lake, Michigan); Gunter, Amer. Midl.
Nat., 28 (2), 1942: 313 (fresh and salt water, e. coast N. Amer.); Richardson, Copeia, 1942: 27 (nuptial
tubercles of male, Lac Brulé, Quebec); Warfel, Frost, and Jones, Trans. Amer. Fish. Soc. (1942), 72;
1943: 257 (age-groups, growth, sex. dimorph., fishery methods, Great Bay, New Hampshire); Dymond,
Canad. Field Nat., 58, 1944: 12 (introd. and spread in Great Lakes, refs.); La Monte, N. Amer. Game
Fishes, 1945: 152, pl. 67; Cuerrier, Fry, and Préfontaine, Nat. Canad., 73, 1946: 20 (Lake St. Pierre,
St. Lawrence R.); Marcotte, Rapp. Sta. biol. St-Laurent, 5, 1946: 75-82 (size, larval growth, migr.,
vert. counts, Quebec); Van Oosten, Trans. Amer. Fish. Soc., 74, 1947: 310 (mortal., Lakes Huron and
Michigan); Marcotte, Rapp. Sta. biol. St.-Laurent, 6, 1947: 45, 56 (size, spawn., Grand R., cf. smelts
of St. Lawrence R. estuary); Contr. Sta. biol. St-Laurent, 18, 1948: 1-107 (descr., distr. Quebec, size,
breed., habits, develop., racial diffs.); Scattergood, Copeia, 1948: 142 (taken in herring fishery, Maine);
Heller, Canad. J. Res., Ottawa, 27 D (5), 1949: 248, 256 (parasites, Bay of Chaleur, Quebec); Baldwin,
Trans. Amer. Fish. Soc. (1948), 78, 1950: 176 (Lake Huron); Bigelow and Schroeder, Fish. Bull. (74)
U.S. Fish Wildl. Serv., 53, 1953: 135 (see Bigelow and Welsh, 1925); Livingstone, Proc. N.S. Inst.
Sci. (1950-1951), 23 (1), 1953: 29 (Nova Scotian lakes where native); Legendre, Freshw. Fishes, Univ.
Montreal, ZT, 1954: 27, fig. 29 (key, ill., size, wgt.); Backus, Bull. Amer. Mus. nat. Hist., 173 (4), 1957:
295 (Hamilton Inlet—Lake Melville region, Labrador).
Osmerus sergeanti Norris, Proc. Acad. nat. Sci. Philad., 20, 1868: 93 (descr., cf. north. viridescens, Schuylkill R.,
Brandywine Cr., Delaware R. to Trenton, New Jersey); Jordan, Evermann, and Clark, Rep. U.S.
Comm. Fish. (1928), 2, 1930: 67 (checklist).
Osmerus dentex Steindachner and Kner, S.B. Akad. Wiss. Wien, Math. Nat. 62, 1870: 429 (descr.,n. China, lat.-
line scales, dentex and eper/anus); Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1882: 294 (diagn., Port
Clarence, Alaska); Goode, e¢ a/., Fish. Fish. Industr. U.S., 1 (3), 1884: 544, pl. 199 (Kamchatka, Port
Clarence, Alaska); Murdoch, Rep. intern. Polar Exped. Pt. Barrow, Alaska, 1885: 131 (abund., Wain-
wright Inlet, Alaska); Smitt, Svenska VetenskAkad. Handl. (1885), N.F. 21 (8), 1886: 166-188 (cf.
eperlanus and spirinchus [name used for White Sea smelt]); Turner, L. M., Contr. nat. Hist. Alaska, 1886:
102, pl. ro (ill., abund. St. Michaels, Alaska, use by Eskimos); Jordan, Rep. U.S. Comm. Fish. (1885),
13, 1887: 830 (Alaska); Nelson, Rep. nat. Hist. Collect. Alaska, 3 (Fish.), 1887: 313 (color, abund.,
Kotzebue Sd., St. Michaels, Alaska); Smitt, Hist. Scand. Fish., ed. 2, 2, 1895: 868 (fewer gill rakers than
in eperlanus); Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 524 (descr., Bering Sea coast,
Alaska; w. coast N. Pacific S.to n. China); Jordan and Gilbert iz Jordan, et a/., Fur Seals... Islands
N. Pacif. (1896-1897), 3, 1899: 439 (Kamchatka, Alaska); Scofield iz Jordan, ef a/., Fur Seals . . . Islands
N. Pacif. (1896-1897), 3, 1899: 497 (Port Clarence, n. Alaska, cf. Kamchatkan specimens); Jordan
Coe Memar Sears Foundation for Marine Research
and Snyder, Proc. U.S. nat. Mus., 23, 1901: 349 (Jap. specimens “‘not evidently different from Alaskan”) ;
Jordan and Snyder, Proc. U.S. nat. Mus., 24, 1902: 587 (descr., color, Kushiro, n. Japan); Schmidt, P.
IU., Pisces Mar. Orient. imp. Rossici, 1904: 278-281, 335, 356, 396 (cf. other named species, distr., e.
Siberia; Russ.); Smith and Pope, Proc. U.S. nat. Mus., 3, 1907: 463 (Shiogama, Japan); Evermann
and Goldsborough, Bull. U.S. Bur. Fish., 26, 1907: 269 (ill., Alaskan locals., refs.); Preble, Biol. Surv.,
U.S. Dept. Agric., N. Amer. Fauna, 27, 1908: 513 (Arctic Red R., Mackenzie R. Delta); Pavlenko,
Trud. Obshch. estest., Kazan Univ., 42 (2), 1910: 15 (Peter the Great Bay, e. Siberia; Russ.); Snyder,
Proc. U.S. nat. Mus., 42, 1912: 403 (n. Japan, in markets); Jordan and Metz, Mem. Carneg. Mus., 6
(1), 1913: ro (ill., Korea); Halkett, Check List Fish. Canada, 1913: 55 (Naas R., Brit. Columbia, Red
R. of North, Mackenzie Delta, Arctic Canada); Hubbs, Proc. biol. Soc. Wash., 38, 1925: 52 (N. Pacific,
Arctic coast of Alaska, cf. Atlantic smelt); Jordan and Hubbs, Mem. Carneg. Mus., ro (2), 1925:
149 (Hokkaido, Japan); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish (1928), 2, 1930: 67 (in
checklist, Alaska and nw. Pacific, S. ton. China); Tanaka, Fish. Japan, 48, 1930: 947, pl. 188, fig. 515
(descr., ill., refs., Hokkaido, Japan); Schrenkeisen, Field Bk. Fresh-w. Fish. N. Amer., 1938: 72 (possible
occur. landlocked, Pacific coast); Gunter, Amer. Midl. Nat., 28 (2), 1942: 308 (fresh and salt water,
w. coast, N. Amer.); Nakamura, Contr. Centr. Fisher. Sta. Japan, 1 (43), 1946-1948: 259 (factors affect.
hatch. of eggs).
Osmerus abbotti Cope, Proc. Amer. philos. Soc., 12, 1871: 490 (Cobessicontic Lake, Maine); Kendall, Occ. Pap.
Boston Soc. nat. Hist., 7 (8), 1908: 49 (Cobessicontic Lake, Maine).
Osmerus spectrum Cope, Proc. Amer. philos. Soc., 1z, 1871: 490 (Wilton Pond, Maine); Kendall, Occ. Pap.
Boston Soc. nat. Hist., 7 (8), 1908: 48 (Wilton Pond, Maine).
Osmerus mordax var. spectrum Jordan, Manual Vert. north. U.S., ed. 1, 1876: 262 (proport. dimens., Wilton
Pond, Maine); Jordan and Gilbert, Bull. U.S. nat. Mus., 16, 1882: 294 (diagn., Wilton Pond, Maine).
Osmerus mordax var. abbotti Jordan, Manual Vert. north. U.S., ed. 1, 1876: 262 (proport. dimens., Cobessi-
contic Lake, Maine); Jordan and Gilbert, Bull. U.S. nat. Mus., 76, 1882: 294 (diagn., Cobessicontic
Lake, Maine).
Osmerus eperlanus dvinensis Smitt, Ofvers. VetenskAkad. Forh., Stockh. (1882), 39 (8), 1883: 34 (name applied
to White Sea smelts); Berg, Freshw. Fish. Russia, 1923: 102 (refs., Russ.).
Osmerus dvinensis Smitt, Hist. Scand. Fish., ed. 2, 2, 1895: 867, ftn. x, 868, table (name applied to White Sea
smelts, by ref. to Smitt, 1882; sep. from O. e. eper/anus by length of longest anal ray and gill-raker counts).
Osmerus mordax abbotti Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 524 (descr., Cobessicontic
Lake, Maine); Jordan, Manual Vert. Anim. NE. U.S., 1929: 54 (size of eye and mouth, Cobessicontic
Lake, Maine); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 67 (landlocked,
Lake Cobessicontic, Maine).
Osmerus mordax spectrum Jordan and Evermann, Bull. U.S. nat. Mus., 47 (1), 1896: 523 (descr., Wilton Pond,
Maine); Jordan, Manual Vert. Anim. NE. U.S., 1929: 54 (size of eye and mouth, Wilton Pond, Maine);
Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 67 (Wilton Pond, Maine).
Osmerus eperlanus var. dvinensis Knipowitsch, Ann. Mus. Zool. Acad. imp. Sci. St. Pétersb., 3 (2), 1898: 5, 6,
11 (cf. typical eper/anus, listed for Murman coast, White Sea).
Osmerus eperlanus dentex Berg, Freshw. Fish. Russia, 1923: 102 (diagn., ill., refs., Russ.); Berg, Zoogeographica,
Jena, 1, 1933: 132 (White Sea, Arctic Siberia); Petrow, Zool. Anz., 107, 1934: 179, 182 (dentex a
recognizable subsp.); Wynne-Edwards, Bull. Fish. Res. Bd. Canada, 94, 1952: 17 (Arctic Canada, abund.
Mackenzie Delta).
Osmerus sergeanti Jordan, Manual Vert. Anim. NE. U.S., 1929: 54 (species or subsp., S. of New York).
Osmerus eperlanus dentex natio dvinensis morpha spirinchus Berg, Zoogeographica, Jena, I, 1933: 132 (lakes,
White Sea region and Kola Penin.).
Osmerus sergeanti Hubbs, Proc. biol. Soc. Wash., 38, 1925: 52 (species, possibly race).
Osmerus eperlanus mordax Wynne-Edwards, Bull. Fish. Res. Bd. Canada, 94, 1952: 17 (mordax subsp. of
eperlanus).
Smelt, Perley, Rep. Sea River Fisher. N. Brunsw., ed. 2, 1852: 37, 62, 81, 82 (cod bait, abund., taken with
fly); Chambers, E.T.D., Forest and Stream, 61 (20), 1903: 380 (Lakes Memphremagog and St. John,
Quebec; stunted race in St. John); Stevenson, Canad. Field Nat., 58, 1944: 128 (status in Lake Huron
in 1943); McKenzie, Progr. Rep. Fish. Res. Bd. Canada, Atl. Sta., 35, 1944: 24 (decrease in nos., Mira-
michi region; favorable results of stream clearance); McKenzie, Bull. Fish. Res. Bd. Canada, 70, 1946:
3-20 (hist., fishery methods, catches, ne. New Brunswick).
MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE
CAMBRIDGE 38, MASSACHUSETTS
““Tirg AGasstz Museum’’
To: Dr. Columbus O'D. Iselin
Woods Hole Oceanographic Institution
Woods Hole, Massachusetts;
with the compliments of the Fish Department
Henry B. Bigelow, Professor of Zoology, Emeritus
William C. Schroeder, Honorary Associate in Ichthyology
Giles W. Mead, Curator of Fishes
Myvanwy M. Dick, Research Assistant in Fishes
Associates in Ichthyology
Richard H. Backus (Woods Hole Oceanographic Institution)
Robert H. Gibbs, Jr. (United States National Museum)
N. B. Marshall (British Museum, Natural History)
PAG Diet oe
“3 * i) ‘i j “ ie.
Cs beadivrlul } hy Occ by adasi
eaiiceae iu? ; aor > eal e.
Ser? fetine } letaABM a4
Fishes of the Western North Atlantic Se
Genus Ma//otus Cuvier 1829
Capelin
Mallotus Cuvier, Régne Anim., nouv. ed., 2, 1829: 305, 306; type species, C/upea villosa Miiller, O.F., Zool.
Danicae Prod., 1777: 245. Greenland.
Generic synonyms:
Clupea Miller, Zool. Danicae Prod., 1777: 245; for C. villosa Miiller.
Sa/mo Fabricius, Fauna Groenl., 1780: 177, for 8. arcticus Fabricius, Greenland; Bloch, Naturg. auslind. Fische,
8, 1794: 99, pl. 381, fig. 1 for 8. groen/andicus Bloch, Greenland; Pallas, Zoogr. Rosso-Asiat., 3, 1831:
389, for 8. socia/is Pallas, islands between Asia and America.??
Osmerus Nilsson, Prod. Ichthyol. Scand., 1832: 11, for O. arcticus Nilsson, Arctic Ocean and northern Norway;
Cuvier and Valenciennes, Hist. Nat. Poiss., 27, 1848: 385, pl. 621, for O. microdon Cuvier and Valen-
ciennes, Norway.
Not Sa/mo Linnaeus 1758; type species, 8. sa/ar Linnaeus 1758.
Not C/upea Linnaeus 1758; type species, C. Aarengus Linnaeus 1758.
Not Osmerus Lacépéde 1803; type species, Sa/mo eper/anus Linnaeus 1758.
Characters. Essentially as in Osmerus (p. 555), except as follows: ScaLes much
smaller and much more numerous, 170—220 transverse rows along lateral line,?* each
scale with only about 5-6 concentric striae posterior to the so-called nucleus and 3
anterior to it, the visible lateral line extending to base of caudal fin; all TrerH so small
as to be easily overlooked, there being no conspicuous fangs either on roof of mouth
or on tongue; the Aprpose fin about half as long at its base as the anal, and shaped
as in Figs. 133, 1343 the Pecrorat fins broadly rounded (rhomboid in smelts; cf. Figs.
131, 134); rays of the pectorals (15-20) and Anat (20-21) more numerous (Osmerus
with 13 at most in pectorals, 17 in anal; Table 1); and the plate on dorsal side of skull
above vomer (PRorTumorp of Starks) not divided by a longitudinal median suture
(p- 555)-"4
Sexual Dimorphism. The two sexes differ so conspicuously, one from the other,
that male and female capelin are separable at a glance. The base of the anal fin is
strongly convex in males but nearly straight in females. The least depth of the caudal
peduncle is greater in males (about 80-85 °/, of distance from tip of snout to eye)
than in females (about 60-75 °/,). The scales on the males are lanceolate along a
band close above the lateral line from the gill opening to the base of the tail, and along
a second similar band low down on the side extending from the pectorals to the pelvics
and thence to the anal, but on females they are rounded. The dorsal fin usually originates
a little farther forward in males (about 52-54 °/, of SL from snout to dorsal) than in
females (about 55-60 °/,). The 5th—12th anal rays are thicker in males than in females.
The margin of the anal is weakly convex, indented opposite the 9th and roth segmented
rays in mature males but about straight in females. The pectoral fins are longer in males
(about 80-85 °/, as long as head) than in females (about 60-65 °/,) and much wider
(Figs. 133, 134).
22. For date of publication, see Sherborn (99: 167).
23. A total of 150 according to Jordan and Evermann (51), followed by Hubbs (43: 51) and Chapman (rr: 299).
24. For additional skeletal characters, see Chapman rr: 291-293, figs. 2, 5, 6, 13, 14.
574 Memor Sears Foundation for Marine Research
The difference in appearance between the sexes becomes greatly accentuated with
the approach of spawning time, when the skin on maturing males thickens along the
bands of modified scales, each of which develops a finger-like extension clothed with
epidermis (Fig. 135 G—1), so that the nuptial bands then appear as two furry or plush-
like ridges (Figs. 133, 135 F). Templeman has reported (110: 146) that “while maturing
males showed only faint traces of these ridges on May 2oth all the males near the
beaches have well developed spawning ridges a month later’; this illustrates the
rapidity with which this nuptial development may take place. Small roundish so-
called nuptial tubercles also develop along the rays of the pectoral, pelvic, dorsal,
and anal fins (Fig. 135 c); these are most prominent and the most numerous on
the paired fins, less so on the dorsal, and usually sparsest and smallest (or lacking)
on the anal. Scattered tubercles also develop on the lower side of the head (39: 417-
420). In maturing males the outer ends of the terminal branches of the rays along the
midsection of the anal fin fuse together at their tips to form a plate-like structure
(Fig. 135 D; TI4: 200).
Finally, in Pacific specimens the vertebrae are reported to be more numerous on the
average in males (av. 65.65) than in females (av. 65.13) (39: 419), but in Atlantic
specimens as more numerous in females (av. 65.65) than in males (av. 65.52) (110: 148),
a puzzling situation that we are not in a position to clarify.
Remarks. Capelin are small, slender, silvery, smelt-like fishes, growing to a max-
imum length of about 7.5—8 inches (p. 579). Though marine in habit, they run up
into very shallow water to spawn, and occasionally into the mouth of rivers. They
are North Boreal to Arctic in nature and are present in countless multitudes in their
centers of abundance. They are distinguishable from Osmerus by the differences given
on p. 554.
Range. North Atlantic and tributary parts of the Arctic; in the eastern Atlantic,
from Spitsbergen and Jan Mayen (48: 90), southeastern Greenland, Iceland, White
Sea, and northern Norway southward to Trondheim Fjord in abundance, occasionally
to Oslo Fjord and the Faeroes; in the western Atlantic, from southwestern Green-
land, Hudson Bay, and northern Labrador, southward to Newfoundland (including
the islands of St. Pierre and Miquelon), the Gulf of St. Lawrence, northern Nova
Scotia, and occasionally to the eastern part of the Gulf of Maine; Arctic coasts of
Alaska and Canada (Bathurst Inlet, Coronation Gulf, and Welcome Gulf) (87:
710; 88: 187, 188; Bean im 58: 135); in the North Pacific southward to the Strait
of Juan de Fuca in the east and to Korea (66: 1) and northern Japan (43: 55) in
the west.
Species and Subspecies. The capelins of the North Atlantic and North Pacific were
originally described as two separate species, the former as vi//osus Miller 1777 (67:
245), the latter first as catervarius Pennant 1784°6 (75: cxxvii) and later as socialis
25. Jensen (48: go, ftn.) has reported four specimens (in the Museum in Copenhagen) for the Faeroes, taken in 1870,
and A.Vedel Taning informs us that 50 were trawled among the Faeroes in 1938.
26. This account was evidently taken from Steller (r08: 149), as pointed out by Schultz (98: 15). For the nomenclatural
validity of this name, see Hubbs and Chapman (44: 296).
ee te
a
Fishes of the Western North Atlantic 575
Pallas 1814 (73: 389). These early names for the Pacific capelin seem then to have
been overlooked; at least all the subsequent references to them seem to have been under
the name vil/osus down to 1937 (Soldatov and Lindberg, 105: 57; Hart, 39: 417-420);
and this same course was followed by Saemundsson as recently as 1949 (93: 93). In
1937, however, Schultz (98: 13-20) reported that the capelin of the Bering Sea region
differ statistically from those of the Murman coast and of Newfoundland in the number
of scales and pectoral fin rays, in the distance from the tip of the snout to the insertion
of the pectoral fins, and in what he termed the “character index,” i. e. the number
of dorsal rays, plus the number of anal rays, plus the number of pectoral rays, plus the
number of scales below the lateral line, minus the total number of gill rakers on the first
arch (see Key, below). He also reported that
Pacific capelin average smaller at maturity than Table III. Relative Frequency in the Num-
the Atlantic fish. ber of Fused Rays in the Anal Fin (in per
: : ; for Pacifi Atlantic Capelin.
Accordingly, Schultz revived catervarius as a Se ore atc and Vande Copelin
distinct species, in which he has been followed eae aL SEs ASG
; used Rays Pacific Atlantic
by Clemens and Wilby (173: 23, 97) as well as by bs ide ae 38
Hubbs and Chapman (44: 296); and Vladykov comeseanel Ne 66 54
(714: 201) has contributed the additional infor- 6.222.222 eee 13.5 5
mation that the number of fused rays in the anal
fin tend to be greater in Pacific capelin than in Atlantic specimens (Table 11). On
the other hand, Vladykov, while adding support to the impression generally held that
Pacific capelin average smaller than Atlantic fish, has pointed out that the greatest
length credited to a capelin anywhere was 10 inches or less for the Arctic coast of
Alaska (26: 105), where catervarius rather than villosus is to be expected on geographic
grounds, if the two are separable.
All in all, our present state of knowledge seems best expressed by rating catervarius
as a subspecies of vi//osus, as Rumjanev has done (97), at least until someone has an
opportunity to study adequate series of capelin from the Arctic coast of Canada; for
example, from Bathurst Inlet, or from Coronation Gulf, where they were reported
many years ago (p. 574), or even from Hudson Bay—localities that are intermediate
between those of record for vi//osus on the one hand and for catervarius on the other.
Key to Subspecies of Ma/lotus villosus
1a. Character index (see above) 30 or more (30-38 recorded).
villosus Miller 1777, p. 576.
1b. Character index (see above) not more than 29 (22-28 recorded).
catervarius Pennant 1784.
Both sides of the North Pacific,
southward to northern Japan and
Korea in the west; to the Strait
of Juan de Fuca in the east.
576 Memoir Sears Foundation for Marine Research
Mallotus villosus (Miller) 1777, subspecies villosus
Capelin
Figures 133-136
Study Material. Over 100 specimens, 115-165 mm SL, from Newfoundland (in-
cluding Labrador), St. Pierre and Miquelon, Halifax, Nova Scotia, Grand Manan,
New Brunswick, and northern Norway, MCZ; also many handled both alive and
freshly caught on the Atlantic coast of Labrador many years ago.
tS
AX SSS
Wa
Ficure 133. Mad/otus villosus villosus, sexually mature male, from comparison with specimens 150-160 mm SL,
from St. Pierre and Miquelon, south of Newfoundland. After Cuvier and Valenciennes as emended by Jessie
H. Sawyer.
Distinctive Characters. The only fishes of the North Atlantic with which the adult
of this species is at all likely to be confused are Osmerus eperlanus (American smelt,
p: 559), Argentina (argentine, Part 4), or, in the southernmost part of the capelin’s range,
the silverside (Menidia). It takes little more than a glance to separate it from all of these,
for its scales are very much smaller than those of O. eper/anus and its adipose fin is of
a different shape (cf. Figs. 131, 134); its mouth is much larger but its scales much
smaller than in Argentina; and it is only in its slender form and silvery color that it
resembles the silverside, which has a much smaller mouth and a rayed (not adipose)
second dorsal fin. For differences between the larvae of M. v. vi//osus and those of other
fishes with which they are likely to be taken, see p. 581.
Description. Based on five males, 129-160 mm, and five females, 129-153 mm SL,
from Newfoundland (including Labrador), St. Pierre and Miquelon, and Grand Manan.
Bopy elongated, moderately compressed, its depth 5.8—6.2 in SL, its greatest thick-
ness about 1.2—1.7 times distance from snout to eye; the dorsal and ventral profiles of
head and trunk slightly convex, but males with contour more abruptly rounded along
base of anal fin. Least depth of CaupaL PEDUNCLE in males 15.2—16.7 times in SL, in
females 19.0-21.8 times, thus noticeably deeper in males. ScaLzs present everywhere
on trunk as well as on base of caudal, but not on dorsal or anal; 177-205 (170-220
reported); 32-37 in oblique series from origin of pelvics to midline of back—16-20
ee
Fishes of the Western North Atlantic 577
above lateral line (18-23 reported) and 16-19 below (18-23 reported); scales very
small, cycloid, thin, not as deciduous as in Osmerus; the largest scales below lateral
line, those of the females mostly ovoid with vertical axis about 1.5 times horizontal axis;
the scales above lateral line smaller and more nearly round; mature males with a prom-
inent band of lanceolate scales, about 5 or 6 scales wide, immediately above lateral line
from gill openings to base of caudal; the scales narrow and very elongated posteriorly
(Fig. 135 G—1); other differentiated scales extending along lower sides from near tip
of pectorals to near anal base, of various shapes, some lanceolate (between pectorals and
pelvics) and others more or less as shown in Fig. 135 (opposite anal), the apexes of some
XY RASS
Figure 134. Ma//otus villosus villosus, female, about 175 mm TL (7.75 in.), from Grosswater Bay, southeastern
Labrador. After Goode; drawn by H. L. Todd.
pointing obliquely downward. Scales along upper sides and back in males ranging in
shape from roundish, but with longest axis horizontal, to pointed or flap-like rear-
ward. A row of Mopiriep LATERAL-LINE SCALES in both sexes, from gill opening to
base of caudal, with a median horizontal channel that is connected here and there
from scale to scale. Most of the scales, particularly above the lateral line, with some of
the circuli complete, but others, chiefly below the lateral line, with all of the circuli in-
terrupted dorsoventrally; the scales above lateral line with fewer circuli than those below.
Heap with dorsal profile straight or very gently convex, 4.3-4.7 in SL. Snour
pointed, 2.9—3.5 in head. Eye 3.9—-4.7, its rear margin midway between tip of snout
and rear margin of opercle, or a little closer to opercle. INTERORBITAL space straight
or slightly concave, 3.5-4.6. Posrorsirat part of head 2.0—2.3 in head. Nostrits
in line with end of snout and upper edge of eye, the two openings on each side close
together, oval-shaped, the posterior one with a flap, the distance from eye 7.0-8.5 in
head. Operce with broadly rounded margin. Margin of Preoprrcte about 0.33 of
distance between perpendiculars from rear margin of eye and margin of opercle.
BraNCuHIiosTeGaLts 8. GILL RaKERS very slender, the longest 1.2-1.6 in eye; 8-10
rakers on upper limb of first arch (11 reported), 24-27 on lower limb (31 reported),
total 32-37 (42 reported).
Movutu oblique, the gape reaching to a perpendicular from nostrils. Lower Jaw
37
578 Memoir Sears Foundation for Marine Research
P5292d)a2RH)
29522 Fal CE
DH SEES
ioe SSeS 3525
1322 dy B35
ENA NON
SION)
dss)
YY YD
yy yyy ay X¥)
yy)
yy)
Figure 135. Madlotus villosus villosus. a, pectoral fin of sexually mature male, about 175 mm Abs tedoaneisiz,
Pierre and Miquelon Island; fin widely spread to show distribution of nuptial tubercles on rays; about x1.3.
B, pelvic fin of same specimen; about xt. c, distal portion of pelvic rays of same specimen as seen by reflected
light; drawn to larger scale to show nuptial tubercles. p, middle sector of anal fin (from gth—14th rays) showing
bony distal plate developed in breeding males; from Vladykov. z, scales along side above first three split anal
rays; about x3. F, scales on side below dorsal fin, from close above to close below upper nuptial ridge; about
<3. c, scale from midbelt of upper nuptial band. u, scale from close beside upper nuptial band. 1, scale from close
above middle of base of anal fin. c-1 from camera lucida drawings; about x13. Drawn by Jessie H. Sawyer.
projecting. Maxittary beginning about halfway between anterior nostril opening and
tip of snout, widening rearward, its posterior margin rounded and reaching to about
opposite center of pupil. PremaxiLiary extending a little past origin of maxillary, its
length about equal to distance between tip of snout and nostril. ManprBLE 1.6—2.0
in head.
Treru. Lower jaw, maxillaries, and premaxillaries with a row of minute teeth,
close-set, unevenly spaced. Vomer with a row of about 3 very small teeth on each side
of a very short, toothless, median space. Palatines with one or two rows of very
small teeth. Tongue boat-shaped, with numerous small retrorse teeth, but with no
fangs as in Osmerus.
ee
Fishes of the Western North Atlantic 579
Dorsat with outer margin straight or slightly convex, its base 1.5—2.0 times
snout, its origin a little closer to base of caudal than to tip of snout, its distance from
latter about 52—54°/, of SL in males and 55—60°/, in females, with 12-14 rays,?’ the
first 2 or 3 unbranched, the longest ray o.g—1.3 times length of base. Dorsat ap1PosE
fin with outer margin straight or slightly convex anteriorly and straight or slightly
concave posteriorly, situated behind dorsal, its base 1.7—1.8 times in dorsal base, its
origin ranging from about over origin of anal fin to about over middle of anal base,
its height 3.0-5.8 times in its base. Caupat deeply forked, its tips pointed in some
but rounded in others. Anat with outer margin weakly convex in mature males and
indented opposite gth and roth segmented rays but about straight in females, its base
1.5—1.7 times dorsal base in males but 1.2—1.4 times in females, its origin from tip
of snout a distance of 70-73 °/) of SL in males but 75-79 °/, in females, and its
origin from origin of pelvics a distance of 1.$—1.7 times in distance between anal
origin and caudal base in males but 1.2—-1.4 times in females, with 19-21 rays (17-22
reported),”” the first 3 or 4 unbranched, the longest ray 2.3-3.1 times in anal base.
Pexvics with rear margin weakly convex, their origin varying from slightly in advance
of, to slightly behind, dorsal origin, with 8 rays, the first (outer) ray unbranched, the
longest ray 1.4—1.8 times in head. Pectorats with rear margin broadly rounded, their
area, when spread, larger in males than in females of about the same size, their origin
under rear margin of opercle, with 18—20 rays, the first (upper) unbranched, the longest
ray I.5—1.7 times in head in males but 1.8—2.1 times in females.
VERTEBRAE 64—70 reported.
Sexual Dimorphism. See p. 573.
Color, In life, translucent, olive to bottle green above, with metallic reflections
like the American smelt (p. 561) but with the back and head darker at spawning time.
Sides a uniform silvery color below lateral line. Scales dotted with very small to minute
dusky specks. Belly white. Lining of body cavity a dusky silvery color.
Size. The largest standard lengths (snout to base of caudal fin) that we have found
to be definitely recorded for any Atlantic Capelin are 215 mm for males and 230 mm
for one female, from the east coast of Newfoundland, where spawning males range
from 130 to 215 mm and spawning females from 110 to 210 mm TL.* An average of
146.7 mm SL or 164.4 mm TL reported by Vladykov for 63 specimens (sex not speci-
fied) from the inner part of the Gulf of St. Lawrence and Cape Breton, Nova Scotia
(114: tab. 2, 202), suggests that Capelin may tend to mature at a somewhat smaller size
in higher than in lower temperatures. The weights of Newfoundland Capelin at different
total lengths are given in Table rv.
Development and Growth. Newfoundland Capelin produce about 15,000—52,000
eggs, according to size.® And it is not likely that they are either much more productive
or much less so in other parts of the Atlantic. The eggs of Newfoundland fish (Fig.
27. Last two rays of dorsal and anal, branching from a common base, were counted as one.
28. Totals of 59,042 males and 23,567 females measured by Templeman (zzro: 145).
29. Counts by W. P. Templeman.
580 Memoir Sears Foundation for Marine Research
Table IV. Average Weights of Newfoundland Capelin at Different Total Lengths.f
Length 7— Weight —~ Length az Weight — Length 7— Weight ~
(mm) (0z.) (mm) (0z.) (mm) (0z.)
9¢ 3d 99 3d 99 3d
TAO) 2 0.48 _ WS) 507 0.91 0.96 LOGuce 1.40 Teo,
nilstey Hae 0.63 _ TSO Nese 1.03 1.13 ZOO ne _— 1.61
TOOM rr. 0.71 0.78 NGO Gar 1.29 1.47 Aglare -- Te7i5e
t After Templeman, rro: 125. * Four fish.
136 A-D) average 0.96—0.97 mm in diameter, are spherical, transparent, and of a
reddish or reddish yellow hue, and have a moderately broad perivitelline space. They
have no oil globule, though the yolk contains many minute globules that tend to unite
as development proceeds. The eggs sink and are so sticky that they cling to each other
or to anything on which they chance to come to rest.
Jeffers has observed that incubation occupies 55 days at a temperature of 0° C, 30
days at 5°C, and 15 days at 10°C, with about 12° the upper maximum, though the eggs
may hatch in water as warm as 15° and in salinity as low as 7 °/o) (46 [1931]: 17, 18).
| SPEDES LF SP BY SY SEF SS 2) OF 9 SF I 757 SY PF STD BY EI ES TOR 3
Sangueumsuumamuaccuaccsssst<:”
LY SANA AU BAY AC GASR BOE SS NSES ot
one 2a fa Be
Ficure 136. Mallotus villosus villosus. a—c, egg in segmentation stages. p, larva before hatching. rk, newly
hatched larva, 6 mm long. F, larva g mm long. c, larva 16 mm long. a—k, after Sleggs (106); F—G, after Temple-
man (ZIO).
Fishes of the Western North Atlantic 581
The larvae at hatching are 6-7 mm long (Fig. 136 £),?° very slender, and wholly
transparent except for a double ventral row of black pigment spots between the heart
and the yolk sac, thence rearward a single row to the base of the future caudal fin, and
a larger spot above the anus; and soon an additional row develops a little farther up
along each side. The pectoral fins are already formed at hatching; the yolk sac has been
absorbed at a length of about 7 mm; at 9 mm (Fig. 136 F) the first indications of the
dorsal and adipose fins are apparent; at 20-30 mm all of the fins are clearly outlined;
and at 30-40 mm the young fry begin to assume the outlines of the adult. It seems
that most of the larvae are 20-40 mm long by their first winter, 40-65 mm early in
the following spring, and 60--80 mm
by midsummer, when they are about
one year of age. Table v gives the
subsequent relation between age and
Table V. Total Lengths (mm) of Newfoundland Capelin at
Two through Five Years of Age, Taken Inshore in Late
June and July.*
Males Females—
(ae ray (a3 \
total length for Newfoundland fish Ages Range Ay. Range Av.
taken inshore in late June and July, 2 years... 140-180 159.7 11s—165 141.8
according |tosTLempleman’s detailed » 3 Y@™2-- «130-200 eS 130-195 Oia
: : 4 years... 155-210 = 189. 150-205 176.7
studies of the scales and otoliths of e yer ait fe ieee
extensive samples (zz0). These values
correspond closely to lengths of 60—
80mm at one year, 135-150 mm for males and 105-125 mm for females at two
years, and 155-205 mm for males and 130-195 mm for females at three years, as re-
ported in 1933 for Newfoundland Capelin by Sleggs from length-frequencies combined
with scale studies (100: 37). Pitt found much the same rate of growth for Grand
Bank Capelin (79: 295-311).
Comparison of Capelin Larvae With Other Larvae. Larvae of Clupea harengus (At-
lantic herring) might be mistaken for those of v. vi//osus if taken together. But the
position of the yolk sac, with its anterior end at least as far forward as the origin of
the pectorals in C. harengus and definitely posterior thereto in v. vi/losus, will serve to
distinguish them from each other until the yolk has been absorbed. During the earlier
stages also, the anus is about 83 °/, of the way back along the body in C. harengus but
only about 75°/) of the way back in v. villosus. At lengths greater than 15-20 mm
the presence of the adipose fin marks v. villosus as such.
Should larvae of Osmerus eperlanus (American smelt) be taken with those of v. vi/-
Josus, an event not reported as yet, the two can be separated up to a length of about
15 mm by the presence in O. eper/anus of a ventral pair of black spots anterior to the
pectoral fins. At larger sizes the very evident air bladder of O. eper/anus sets it apart
from v. villosus.
Other northern fish larvae that parallel the larvae of v. vil/osus in their slenderness
are those of Pholis gunnellus (rock gunnel) and Ammodytes (launce). But there should
be no danger of confusion here, for the anus in these is near the middle of the body,
as compared with about 75°/, of the way back in v. villosus (II0: 68, 69).
* After Templeman, rzro.
30. For excellent illustrations of Newfoundland larvae, see Dannevig (zo: pl. 3, figs. 26, 27); for descriptions and illustra-
tions, see Templeman (rz0: 70-75, figs. 18-20); for Icelandic larvae, see Schmidt (96: 16-18, pl. 1, figs. 24-28).
582 Memoir Sears Foundation for Marine Research
Age at Spawning. Sleggs (100: 21-30) and Templeman (zr0: 87-106) have
shown from length-frequencies, scales, and otoliths that a great majority of the New-
foundland Capelin spawn when they are three years old, and this is also believed to
be the case around Iceland as well as off northern Norway. But a few Newfoundland
females do not spawn until they are four or even five years old, and Hansen has recently
reported, from a study of otoliths combined with length-frequencies, that in western
Greenland waters 48-69 °/, (according to locality) do not spawn until the fourth or
fifth year (37: 121-124). On the other hand, Rachmanowa (80: 120) concluded that
most of the spring-spawning fish on the Murman coast are only two years old (zwei-
jahrigen), and most of the summer spawners only one year old (einjahrigen). But we
are not in a position to judge the validity of these results, for no illustrations of the scales
on which the observations were based are included in the account.
In either case, most of them die shortly after they have spawned once, though
some females may spawn twice or even three times during their life.*
Spawning Season. In the western Atlantic the Capelin are spring-summer spawners,
with the season opening some weeks later in the northern part of the breeding range
than in the southern part. In Hudson Bay they have been reported as spawning from
late July into early August (rr2: 21). For the years 1941 and 1942, when detailed
information was obtained for many localities in Canada (110: 56, 57, figs. 14, 15),
they were spawning from July 25 to August 20 along Labrador north of Hamilton
Inlet, from June 18 to August 25 on the outer Labrador coast south of that point,
from June 1 to August 25 along the coast of eastern Newfoundland, from June 1 to
August 4 along the western part of the coast of southern Newfoundland, from June 6
to July 2 along the northern part of the coast of western Newfoundland, and from
June 1 to August 12 in the Strait of Belle Isle. The chief production took place during
the last three weeks of June and first three weeks of July in eastern and southeastern
Newfoundland, and some weeks later in outer Labrador. In 1931 and 1932 the season
seems to have been earlier, at least in Newfoundland waters, for the latest dates re-
ported by Sleggs were June 28 to July 3. But year-to-year variations of this sort
seemingly are not wide, for Anspach, writing in 1819, described the Capelin as coming
into Conception Bay about the 20th of June, to continue “‘on this coast for about six
weeks” (2: 400, 401). Templeman has also reported that on rare occasions Capelin
“may return again to the beaches to spawn” some weeks after the main spawning
period has ended. And it seems that Capelin larvae taken around Newfoundland in
April and May are more likely to be the product of late spawning inshore than of
spawning that may commence as early as February offshore, as has been suggested
(roms 355)).
Along the coast of western Greenland the Capelin spawn from May through June
and less commonly into July (27: 177, 178; 90: 135), from early March into June
around southwestern and southern Iceland, and until August around the northwestern,
northern, and eastern coasts (92: 413). The spawning season lasts from late April or
31. See Templeman (rro: 62-64) for the evidence upon which this statement rests.
Fishes of the Western North Atlantic 583
early May through June in northern Norwegian waters (14: 152); but on the Murman
coast the fish are inshore from March to July; and Rachmanowa’s observations (1928)
suggest that there may be two reproduction peaks, an early one in March and April
and a late one in June and July.
Spawning Grounds. Capelin spawn on the bottom close inshore and on the off-
shore banks as well (see below). It has been a matter of common knowledge for genera-
tions that most of the hordes of Capelin of Newfoundland (including Labrador) spawn
right up on the beaches, even in the wash of the waves;3? and multitudes spawn simi-
larly along the intertidal zone in Greenland waters. But considerable spawning also takes
place near shore, from 2 or 3 fms. down to 20 fms. or so; occasionally even at 40-50
fms. ;°3 and at 25-35 fms. on the Grand Bank, where Pitt has recently reported the
widespread presence of spawning populations (79: 279-281). In Icelandic waters
they ‘‘most frequently breed in rather deep water.’’*4 Around northern Norway also,
they have been reported as spawning mostly at depths of 4-20 fms. (94: 172); if
beach spawning takes place at all there, this is not in an amount sufficient to have
attracted the attention of either local fishermen or scientists. In the Barents Sea “eggs
are most often caught ... above depths from 50-100 meters” (Rass, 81: 31).
Type of Bottom for Spawning. When Capelin spawn on the beach, they usually
select stretches of gravel for the purpose rather than sandy stretches where the eggs
are more apt to be buried by the action of the waves. The chief production of eggs on
the Newfoundland beaches occurs where the grains of gravel run about 5—15 mm in
diameter (1r0: 41), and on the Grand Bank where the sand grains are almost 0.5—
2.2 mm (7g: 280). Pacific Capelin, which are supposed to average smaller than those
of the Atlantic, are reported as spawning chiefly where the gravel size is 1-5 mm
(40: 24). But the fish do sometimes deposit their eggs on rocks, on sand, and even
on algae, more commonly so, it seems, on the western Greenland coast than on the
Newfoundland coast (48: 82). And when they spawn regularly below the intertidal
zone, as seems the general rule in Norwegian waters, they may do so on sand, or without
regard for the nature of the bottom.
Spawning Activities. The schools that come in on the beach consist solely of mature
males and mature or maturing females; the juvenile fish remain somewhat farther out,
where quantities of them are often seen, most of them deeper down. On the whole the
larger fish spawn earlier than the smaller ones.
“The spawning act,” writes Templeman, “has been observed by a few scientists,
and by tens of thousands of Newfoundlanders,” for Capelin spawning on the beach
are in such dense schools that they are in plain view from the strand (zr0: 33).
So spectacular are their spawning activities that these have been described in vivid
terms,* the earliest, it seems, for the Pacific subspecies catervarius Pennant, in 1784
(75: CXxvI1; 73: 389), and time and again for the Atlantic vé//osus.
32. Anspach, for example, wrote in 1819 (2: 401) of Capelin depositing their eggs ‘‘on the sand” in Conception Bay.
33- For details, see especially Templeman (zro: 59-61).
34- Information contributed by A.Vedel Taning, from observations by H. Einarsson.
35- The accounts by Perley (76: 135) and Lanman (60: 225), often quoted, evidently were taken (partly verbatim)
584 Memoir Sears Foundation for Marine Research
As the fish run in on the beach, either two males may squeeze a female between
them or a single male may keep in close contact with a female. The spawning trios or
pairs, having run up the beach as far as they can, deposit their sperm and eggs and then
separate to regain deeper water again if they be fortunate; so many of them are not
that the beaches where Capelin are spawning usually are lined with windrows of dead
fish. Thus Anthonie Parkhurst described them as early as 1600 as “‘driven drie by the
surge of the Sea... you can take up with a shove-net as plentifully as you do wheate in
a shovell”’ (in 35: 133). The mass strandings of Capelin have been described repeatedly
since then, as by Anspach in 1819 for Conception Bay (2: 400). Even if the Capelin
do escape in the wash of the waves, they are so exhausted and the fins of the males so
battered that multitudes of the dead and dying are to be seen floating close inshore.
It appears that nearly all of those remaining must perish soon thereafter, so unusual 1s
it for a Capelin to spawn more than once during its lifetime.
Spawning is said to take place chiefly in cloudy weather or at night, without regard
for the stage of the tide, and it is likely that the course of events is similar for Capelin
that spawn in deeper water where they do not come under direct observation.
The eggs spawned on the beach are soon mixed with gravel by the waves, resulting
in “a quivering mass of gravel and attached eggs” (Templeman, rro: 41). But mul-
titudes of those that have failed to adhere there drift offshore to sink to the bottom,
where they too stick to anything with which they come in contact, often massing on
nets and anchor lines.
Relation of Spawning to Temperature. In Newfoundland coastal waters, the Capelin
spawn chiefly when and where the temperature is between 6° and about 8 or 9°C, but
never in water warmer than about 10.5°; on the Grand Bank they spawn at 2.8—
4.7°C (100; 110; 83), along western Greenland in water as cold as 3—5°C,** and
in the Barents Sea at 2° (81: 29-33).
Habits and Migrations. The migratory pattern of the Capelin consists essentially
of a passive dispersal of the larvae followed by the wanderings of the juveniles in search
of food; this tends to bring them inshore and near the surface in early summer but
offshore and into deeper water in autumn, leading finally to the inshore migration of
the mature or maturing fish toward their spawning grounds in spring and summer.
There is no direct evidence from actual captures that Capelin as a whole spread
very far during their period of growth, at least in the western side of the Atlantic.
The greatest distance from land where immature fish have actually been taken, off
Labrador or Newfoundland, is only about 75 miles for fry small enough to have been
picked up in tow nets, and 150 miles for larger (hence older) juveniles trawled at or
near the bottom (p. 585, Fig. 137). Around Iceland, where young fry abound close
inshore, Jespersen has recorded captures of fry 60-70 miles offshore to the west,
with a few taken at distances up to 200 miles or so toward the southeast (50: 12,
from the earlier accounts by Chappell (r2: 131-134), as Kendall (55: 29) and Hardy (38: 4-8) have pointed out.
For recent accounts, see Sleggs (100: 38-40) and Templeman (rro: 32-39).
36. Information contributed by A. Vedel Taning.
Fishes of the Western North Atlantic 585
50°
|
|
|
|
a2 = 50°
|
| e
|
x
e
ST, PIERRE
hele GRAND BANKkK.:
e
45° r sls 45°
15
“taps e
. e
e
60° 55° 50°
Ficure 137. Recorded localities off Newfoundland where Capelin have been small enough to be taken in tow
nets (x) or large enough to be trawled on or near the bottom (@). From Rep. Newfoundland Fish. Res. Comm.
fig. 8, and 15, fig. 10). A. Vedel Taning has informed us that young fry from the Ice-
landic area are sometimes carried as far southward as the Faeroes, or even farther. While
the fish of northern Norway and the White Sea region may disperse more generally
in the Barents Sea, recent studies have not supported an older view that the center of
population for the Capelin of the North Atlantic lies in the northern part of the Nor-
wegian Sea, whence Capelin fan out, as it were, to northern Scandinavia, Spitsbergen,
Jan Mayen, Iceland, Greenland, and the coast of North America.
Some, perhaps a majority, of the Capelin that are hatched well up in the ex-
586 Memoir Sears Foundation for Marine Research
60°
Ficure 138. Distribution of the coldest water, irrespective of depth, from southern Newfoundland to the Gulf
of Maine, for May rg15. After Bigelow, somewhat emended.
tensive bays of Newfoundland and Labrador pass the winter there, near bottom; they
have been found repeatedly in the stomachs of winter-caught cod, and recently in the
stomachs of seals taken in winter from Lake Melville above the head of Hamilton Inlet,
Labrador (3: 295).
Those that are produced along the outer coasts, and some of the Bay Capelin as
well, come under the influence of the general pattern of coastal circulation that 1s
characteristic off western Greenland and Newfoundland (including Labrador) during
the part of the year when Capelin are near the surface. On this basis the hosts of Cape-
lin on the southern part of the west coast of Greenland (with some recruits from the
southeast coast) may be expected to tend northward, perhaps as far as Disko.
There is nothing in the published record to suggest that Capelin are ever carried
across the northern part of the Labrador Sea from Greenland to America, or that they
survive the journey if they are. But those hatched on the outer coast of Labrador may
be expected to come under the influence of the inner edge of the ice-laden Labrador
Current until the latter slackens, in autumn (Fig. 138). Also, ‘‘a smaller branch of the
Arctic current enters the Gulf of St. Lawrence (but not as a steady stream) through the
Strait of Belle Isle on its north side” (Huntsman, 45: 277, 278; Fig. 139); this
doubtless brings some Capelin with it besides the icebergs that it sometimes carries as
far as the Island of Anticosti. But there is no knowing how important a role this cur-
rent may play in maintaining the stock of Capelin in the inner part of the Gulf of
St. Lawrence.
Fishes of the Western North Atlantic 587
X
‘Nie
LABRADOR \ SS * —
QUEBEC
A
7
ay Bee CCL x
SHALLOWS & ¥
) x
Ficure 139. Prevailing circulation, spring and summer, around Newfoundland, in the Gulf of St. Lawrence
and along the Atlantic Coast of Nova Scotia. After Huntsman.
Capelin hatched along the northern part of the coast of western Newfoundland,
together with the older juvenile fish, may similarly be expected to drift at first north-
ward and then outward along the southern side of the Strait of Belle Isle, if they travel
that far. Off the coast of eastern Newfoundland, however, where the main drift of
the Labrador Current lies well out from the coast (usually skirting the eastern edge
of the Grand Bank rather than flooding the latter), the Capelin might be expected
to either pass their juvenile years nearby or disperse over the Bank. This expecta-
tion is borne out by the localities where the larvae and older juveniles have been taken
(Fig. 137).
Presumably the waves of Capelin that occasionally invade the Bay of Fundy are
carried thither by the ice-chilled Nova Scotian Current which brings water colder than
3°C (37°-38° F) along the Nova Scotian coast and into the eastern side of the Gulf
of Maine at some time in the spring in most years (Fig. 138).*” It is not likely that
any Capelin or their offspring ever find their way north again, if any succeed in repro-
ducing that far south.
37- For additional details, see Bigelow, 7: 831, 832; Taylor, et a/., rog: 311, fig. 14.
588 Memoir Sears Foundation for Marine Research
Food. Capelin feed chiefly on euphausiid shrimps as well as various isopod-, gam-
marid-, and copepod-crustaceans. The stomachs of Capelin taken from among the
spawning schools have been found full of Capelin eggs—evidence that they may not
fast as rigorously at spawning time as various other fishes do.
Enemies. “A list of the animals that prey on the Capelin would include the names
of almost all fish, sea birds and sea mammals that inhabit the parts of the world where
capelin are to be found” (Templeman, rzo: 132). Seals, for example, subsist largely
on them; finback whales have often been found full of them; during their periods of
abundance they are one of the chief items in the diet of the white whale (De/phinapterus)
in the estuary of the St. Lawrence River and no doubt elsewhere; they are also eaten
by the common porpoise of the north (Phocaena) (33: 47). The nesting colonies of puf-
fins, murres, guillemots, gulls, terns, and shearwaters all take heavy toll of them.
And the list of fishes known to prey regularly on them includes spiny dogfish (Squalus
acanthias), Greenland shark (Somniosus microcephalus), Atlantic salmon (Salmo salar),
Arctic charr (Sa/velinus alpinus), Atlantic herring (Clupea harengus) which often gorge
on Capelin larvae, sculpin (Myoxocephalus), eelpout (Macrozoarces americanus, in American
waters), wolffish (4uarrhichas), flounders (Pseudopleuronectes), Greenland halibut (Rein-
hardtius hippoglossoides), and Greenland cod (Gadus ogac). But their worst enemy by
far on both sides of the Atlantic is the Atlantic cod (Gadus morrhua), which not only
devour immense numbers of Capelin when the latter come in to spawn, but also harry
the young in deeper water offshore, seemingly without letup.
Other serious losses to the potential population of the Capelin result from the
drying out of great numbers of eggs spawned on the beaches, and from “‘immense
destruction of the exposed spawn [that] is wrought by maggots or small beach flies of
the genus Fuce/laria” (Sleggs, T00: 44).
Variations. Neither the average numbers of vertebrae that have been reported for
various localities (377: 121-1243; IITO: 109-1153; 79: 285-290) nor other numerical
and proportional characters (68: 64) have suggested any consistent racial differences
between the Capelin of different regions.
Numerical Abundance. Attention has been called time and again to the multitudes
in which Capelin gather when they come inshore. Perhaps it will suffice here to refer
to Anspach’s description of them in 1819 in Conception Bay, Newfoundland, as being
present in schools so immense as to color the surface of the sea and to enable two
persons to fill a common-sized boat with them in less than two hours (2: 400), and to
Hardy’s description of the water surface as appearing like “‘a living mass as far as the
eye can reach” (38: 8, 9).
However, the only precise available information as to their actual numbers is
Slegg’s calculation based on the counts of measured samples of beach material (roo:
42, 43): The number of eggs per cubic foot was between six and seven million, at
which rate “it would require about a third of a million females to furnish the spawn
on a hundred yards of beach and a like number of males. When we take into account
the spawn below the tidal zone, it seems safe to say that a million capelin spawned upon
a ae
Fishes of the Western North Atlantic 589
each hundred yards of beach” on this particular occasion. Templeman has written simi-
larly of ‘“‘possibly hundreds of millions’? spawning at Grand Beach on the east coast
of Newfoundland (rro: 52). With this same sort of thing taking place here and there
from Saglek Bay in northern Labrador to southeastern Newfoundland and St. Pierre
Island, along western Greenland, and along at least the coast of southern Iceland,
(where Jespersen has recorded a catch of 5,900 larvae in one 20-minute haul with a
“young fish” trawl [50: 11]), the total number of Capelin involved must surpass human
reckoning.
Periodic Fluctuations in Abundance. It has long been known that Capelin may be
decidedly erratic from year to year in their appearances at particular localities within
short distances; for example, around the southwestern angle of Newfoundland, where
they may even fail to spawn at all, as seems to have happened locally in 1941 and 1942
(110: 55-58, figs. 14, 15), and around Ungava Bay near the northern boundary of their
range, where they were reported as far more plentiful in 1884 than in 1947-1949
(22: 104). Capelin also appear from time to time in schools far to the south of their
regular range, as in the Gulf of Maine in the west and off Norway in the east (179:
15). The fish also range much farther eastward along the Murman coast in some
years than in others (So: 119). Catches of 4,800,000 pounds of Capelin in the Gulf
of St. Lawrence in 1919, 1,990,000 in 1951, 677,000 in 1952, 147,000 in 1964,
and 497,000 in 1956 suggest that the numbers of Capelin declined greatly there
during recent years. But the decline suggested by these figures seems as likely to have
resulted from a decrease in the demand for Capelin as from any general reduction
in their numbers. We question also whether the poundage credited to Newfoundland
for 1952%8 is realistic, for we have been informed by letter from John D. Kaylor of
the General Foods Corporation, a fisheries expert of long experience, that ‘‘on the
spot inquiries in many places, plus personal observation during the height of the
capelin run [of 1958] show a total catch for a// uses at less than 5 million pounds for
all Newfoundland.”
It is said that Capelin have not visited the coast of western Greenland in such
hordes of late years as they did during the colder-water period from about 1885 down
to the 1920’s, but they may have continued equally abundant in deeper water, farther
out from the immediate tideline.
Relation to Man. The Capelin, a delicious morsel as we can bear witness, is an
important fish, not only in the natural economy of northern seas but in that of the
native inhabitants of Greenland, where great quantities are dried in good Capelin years
during the spawning run, to be used for human food during the winter. So vital an item
is the Capelin in the winter diet of the Greenland Eskimos that they suffer great hard-
ship in any particular district in a year when the run fails. The Capelin are used
there for dog food as well.
In Newfoundland, great quantities were netted for human consumption during
38. We are greatly indebted to the late A. H. Leim and W. P. Templeman of the Fisheries Research Board of Canada,
and to F. B. Clark, Canadian Consul (commercial) at Boston, for information regarding the commercial landings
of Capelin in Newfoundland for recent years.
590 Memoir Sears Foundation for Marine Research
the first quarter of the 19th century, to be dried for shipment to London, to be exported
“by the thousands of barrels to the eager French fishermen on the Bank” as bait
(Hardy, 38: 8), and to be used locally as manure. Then there followed a period lasting
down through the first quarter of the present century when Capelin were in so little
demand in Newfoundland that Sleggs, writing in 1933, estimated the local consumption
of them to be not ‘‘more than a few pounds per head, annually” and the exports for
some time past as “insignificant” (roo: 57). Capelin were in greater demand, however,
along the Quebec shore of the Gulf of St. Lawrence than in Newfoundland (see Periodic
Fluctuations in Abundance, p. 589); 130,268 pounds of dried Capelin plus 29,117 pounds
of fresh and salted fish were exported from the Island of St. Pierre to France in 1924;
and 116,855 pounds (frozen) were shipped to the United States in 1925 (ro: 151).
Of late years Capelin have been taken in much larger quantities in Newfoundland
than had been the case for many years previous to the Second World War, sure evi-
dence of an increase in the demand (for catches 1952, 1954, see p. 589). We owe to
F. B. Clark, of the Canadian Consulate General, the information that 634,000 pounds
(2 °/,) of the Newfoundland catch for 1954 were used for human consumption (fresh,
dried, or smoked), and the remainder (98 °/,) for other purposes, such as fresh and
frozen bait for cod fishermen, fertilizer, etc. In addition, it has been reported that
Capelin are utilized as fish meal, as food (lightly salted and dried) for dog teams, as
canned cat food, and as a source for a high quality marine oil (zzz: 7, 8, 18).°9
Methods of Capture. Wherever Capelin are caught they are taken chiefly in cast
nets or dip nets, but sometimes in beach seines and even in tubs, buckets, and hand
scoops, as has been described in vivid terms by Hardy for Conception Bay, Newfound-
land (38: 8). We have never heard of one being caught on hook-and-line.
General Range and Occurrence in the Western North Atlantic. The general range of
the Capelin is outlined on p. 574. In Greenland, Capelin are known on the east
coast northward to Scoresby Sound (about 70°N) and on the west coast northward
to Thule (76°30’N); on the west coast they did not occur of old with any regularity
north of Vaigat (about 70°N), but by the 1930’s they were abundant in Northeast
Bay, and by 1935 they had spread to the northern part of the Upernavik district
(about 74°25’N); one was even taken at Thule that year, and a second was taken in
1936 (48: 87, 88).
On the American side of the Labrador Sea, the most northerly point where we
find them reliably reported on the Atlantic coast of northern Labrador is Saglek Bay
between 57 and 58°N (41: 5). Capelin have not been recorded for Baffin Island, but for
Ungava Bay a few have been reported recently though larger numbers were recorded
for 1884 (22: 104). They have also been recorded for many localities in the northern
part of Hudson Bay southward to James Bay, sometimes in great abundance (112: 20),
for Melville Peninsula in the Arctic (78: 7), for Bathurst Inlet (87: 710), and for
39. We are indebted to Leslie W. Scattergood, U.S. Fish and Wildlife Service, for bringing this reference to our at-
tention, and to W. P. Templeman, St. Johns, Newfoundland, and John D. Kaylor, General Foods Corporation,
for personal communications regarding the utilization of Capelin.
Fishes of the Western North Atlantic 591
the mouth of the Great Fish River, tributary to Bathurst Inlet, where Richardson
found them present “in multitudes” (88: 187). For records of the Capelin farther
westward along the Arctic coast of America, see Walters (116).
Southward from Saglek, Labrador, Capelin have been reported by fishermen for
so many points as to show that they visit every suitable spawning beach in greater or
lesser numbers all along the outer coast; they have been taken even as far inland as
Lake Melville above the head of Hamilton Inlet.4? They are reported to be widespread
along the coast of eastern Newfoundland and along the eastern part of the south
coast; and it was known that Capelin swarm about the islands of St. Pierre and
Miquelon long before Duhamel, writing in 1769, called their abundance on the Grand
Bank to scientific attention. It seems that the waters along the western part of the
southern coast of Newfoundland and along the southern part of the western coast have
become too warm for regular spawning by the time the Capelin are sexually ready to
spawn (110: 54—57). But they do spawn yearly along the northern part of Newfound-
land’s west coast as well as in the Strait of Belle Isle, though perhaps not as abun-
dantly in the Strait as on the east coast of Newfoundland; this, it seems, applies also
to the north shore of the Gulf of St. Lawrence in general.
In good Capelin years, considerable numbers of them appear on the south shore
of the St. Lawrence River estuary and at the entrance to the St. Lawrence River; for
instance, the reported catch for the Gaspé region (Gaspé and Bonaventure counties)
and for Rimouski County, Quebec, in 1919 was 3,635 barrels whereas in 1929 it was
only 240 barrels; Capelin have been reported to occur as far up the St. Lawrence River
estuary as Trois Pistoles, about opposite the Saguenay River (115: 79). They are well
known around Anticosti Island (97: 285; see also Study Material, p. 576). But we find
no definite evidence that Capelin visit the southern part of the Gulf in any numbers,
or with any regularity, if they occur there at all.
To the south of Cabot Strait, Capelin occur only sporadically? and in limited
areas. ‘“The southeastern corner of Cape Breton is the center of such an area, where
large quantities were taken in 1917. Halifax is the center of another area’”’ where
Capelin were reported for about 1853 (Hardy, 38: 5, ftn.) and were abundant in 1919.
On rare occasions they appear for a year or two in the Bay of Fundy in great num-
bers; Perley, for example, writing in 1852, reported them for a number of localities
near St. John, New Brunswick; they were next reported in the Bay in 1903; a few were
taken in 1915; in the autumn of 1916 they were so plentiful that a catch of 3,000
pounds of fish in the Passamaquoddy region included 2,000 pounds of Capelin. They
have also been reported for various other Fundian localities. In the spring of 1917
they reappeared on the Nova Scotian side of the Bay, and in 1919 they were taken
40. Backus (3: 295) has reported Capelin found in the stomachs of cod taken there in winter. And one has been
reported for Northwest River, at the head of Lake Melville (54: 210).
41. The late A. H. Leim informed us, from his personal observations, that the fish reported in the commercial
statistics for 1930 and preceding years as Capelin from Prince Edward Island “doubtless were silversides (Menidia)
which are known there as ‘capelin.’”
42. Letter from A. G. Huntsman.
592 Memoir Sears Foundation for Marine Research
not only in the deep water off Passamaquoddy Bay through the late winter and early
spring but to the west as well, far up the Penobscot River. There is even one specimen
labeled Woods “Holl,” Massachusetts, in the collection of the Academy of Natural
Sciences of Philadelphia (28: 555), whether correctly so or not. None seems to have
been seen since 1919 in the Gulf of Maine, nor have Capelin ever been reported as
spawning south of the Gulf of St. Lawrence, so far as we can learn.
A final item, remarked on first by Cuvier and Valenciennes in 1848 (19: 408),
is that on the shores of the White Sea, spawned-out Capelin that are washed up on the
shore often become enveloped in nodules of beach material, within which their skeletons
persist, perfectly preserved, until the nodules fall apart. This type of subfossilization,
commented on in greater detail by Collett (14: 161), has also been reported for Iceland
(r: 42), and for Greenland, where Quaternary “‘clay plates are not rarely found
which ... contain a skeleton or impression of capelin....” Such clay balls containing
remains of Capelin are well known from the Quaternary deposits in Norway also
(48: 92), and from Bosnia at an elevation of 580 m above the sea (g).
Synonyms and Selected References:
Clupea villosa Miller, Zool. Danicae Prod., 1777: 50 (Greenland; not seen); Gmelin iv Linnaeus, Syst. Nat.,
ed. 13, Z(3), 1793: 1409 (north. seas; for publ. date, see Hopkinson, Proc. zool. Soc. Lond., 1907:
1035-1037).
Salmo arcticus Fabricius, Fauna Groenl., 1780: 177 (descr., spawn. habits, season, Greenland).
Salmo groenlandicus Bloch, Naturg. auslind. Fische, 8, 1794: 99, pl. 381, fig. 1 (refs., descr., ill., spawn., food,
use in Greenland, vernac. names); Shaw, George, Genl. Zool., 5, 1804: 70 (descr., abund., spawn.
migr., food, use, Greenland, Iceland, Newfoundland); Richardson, John, zv Franklin, Narr. . . . Journey
to Polar Sea, Append. 6, Fishes, 1823: 710 (abund., spawn., Bathurst Inlet, Arctic Canada).
Clupea lodna Hermann, Observ. Zool., posth. ed., F. L. Hammer, Paris, 4, Pisces, 1804: 290-328 (316) (see
Hubbs, Copeia, 1936: 124-125; “the capelin, Ma//otus villosus, is called Clupea lodna [from Iceland,
described from plate by Olafsen and Povelsen],” Reise Island, 2, 1774-1775: pl. 28).
Salmo villosus Miller, Zool. Danica, 4, 1806: 45, pl. 160 (descr., ill, Greenland).
Mallotus villosus Cuvier, Régne Anim., nouv. ed., 2, 1829: 306 (diagn., cod bait); Storer, Mem. Amer. Acad.
Arts Sci., N.S. 2, 1846: 454; also separate, Cambridge, Mass., 1846: 229 (descr., refs., Greenland, New-
foundland); Cuvier and Valenciennes, Hist. Nat. Poiss., 27, 1848: 392 (sp. Ma/ottus), pls. 622, 623
(descr., ills. male and female, habits, food, abund., season, St. Pierre and Miquelon, use, early refs.,);
Gaimard, Voy. Islande et Groenl., Atlas, Zool., Poiss., 1851: pl. 18, fig. r (ill., male, Iceland); Agassiz,
L., Proc. Boston Soc. nat. Hist., 3, 1851: 42 (modern subfossil., Iceland); Perley, Cat. [in part] Fish. N.
Brunsw., Nova Scotia iv Rep. Fisher. Bay of Fundy, ed.1, 1851: 135; descr. Cat. [in part]..., ed. 2,
1852: 203; (descr. Cat. [in part] ..., as separate, ed. 2, 1852: 25 (never farther S. than N. Brunswick,
spawn., odor, export, cod bait, import. food in Greenland, never enters freshw. streams); Rep. Fisher.
G. of St. Lawrence, ed. 2, 1852: 7—g (cod bait and manure, rel. to cod fishery); Storer, J. Boston Soc.
nat. Hist. (1850-1857, 6, 1857: 265 (N. shore, Gulf of St. Lawrence); Bell, Canad. Nat. Geol., 4, 1859:
207 (Gulf of St. Lawrence); Gill, Proc. Acad. nat. Sci. Philad. (1861), Append., 1862: 53 (catalog., e.
coast N. Amer.); Canad. Nat. Geol., N.S. 2, 1865: 259 (in synop., Gulf of St. Lawrence, Bay of Fundy);
Weiz, Proc. Boston Soc. nat. Hist. (1864-1866), 10, 1866: 275 (spawn., season, abund. of fry, Square
I., near Okkak, n. Labrador); Hardy, Proc. N.S. Inst. Sci., r (2), 1867: 4-13 (n. Nova Scotia and Halifax,
abund., spawn., enemies, use, Newfoundland); Jones iz Hardy, Proc. N.S. Inst. Sci., r (2), 1867: 5-6
(descr.); Reeks, Zoologist, London, (2) 6, 1871: 2556 (abund., spawn., Newfoundland); Lanman, Rep.
U.S. Comm. Fish. (1872-1873), 2, 1874: 225 (spawn. activ., after Chappell, 1818: 132-133); Bean iz
Kumlien, Bull. U.S. nat. Mus., 15, 1879: 135 (Bathurst Inlet, Arctic Canada); Jones, Proc. N.S. Inst.
Sci. (1879-1882), 5, 1882: 93 (S. to Halifax, Nova Scotia, in cold years); Stearns, Proc. U.S. nat.
Mus. (1883), 6, 1884: 124 (n. Gulf of St. Lawrence, no def. locals.); Goode et a/., Fish. Fish. Industr.
Fishes of the Western North Atlantic 593
U\S., 1 (3), 1884: 544, pl. 201 (abund., Grand Bank, Newfoundland, Labrador, spawn., largely after
earlier accounts, ill.); Dresel, Proc. U.S. nat. Mus. (1884), 7, 1885: 255 (w. Greenland); Smitt, Svenska
VetenskAkad. Handl. (1885), N.F. 2r (8), 1886: 189-196 (vi//osus, Scandinavia, cf. arcticus, Green-
land); Jordan, Manual Vert. Anim. north. U.S., ed. 5, 1888: 76; and subsequent eds. (diagn., Arctic
S. to Maine); Packard, Labrador Coast, 1891: 401-402 (se. Labrador); Smitt, Hist. Scand. Fish., ed. 2,
2, 1895: 876, pl. 41, figs. 2, 3 (refs., descr., ills., range, genl. nat. hist., ercticus of Greenland not separate
from vi//osus of Scandinavia); Bay, Medd. Gronl., 19, 1896: 57 (e. Greenland); Collett, Forh. Vidensk-
Selsk., Christiania, 9, 1903: 147-162 (habits, develop. stages, food, distr., Norway, Novaya Zemlya, Spits-
bergen, subfossil.); Schmitt, Monogr. Isle d’Anticosti, 4, Zool., Poiss., 1904: 285 (season, cod bait, Anti-
costi); Kendall, Schmidt, Medd. Komm. Havunders. Kbh., Fisk. 2 (4), 1906: 16-18, pl. 1, figs. 24-28
(descr., ills., larvae, around Iceland); Proc. Portland Soc. nat. Hist., 2 (8), 1909: 210 (Labrador locals.,
refs.); Kendall, Proc. U.S. nat. Mus., 38, 1g11: 508 (adults, larvae, dates, Labrador); Fowler, Proc.
Acad. nat. Sci. Philad., 63, 1912: 555 (specimen labeled Woods “Holl” Massachusetts); Meek, Migra-
tions of Fish., 1916: 138 (spawn. migr., abund., periodicity, enemies, range); Kendall, Copeia, 1917: 28
(Dennys R., e. Maine in 1916); Dannevig, Garedt Fish. Exped. (1914-1915), Dept. Naval Serv., 1919:
30, fig. 21, pl. 3, figs. 26, 27 (larvae, Newfoundland, Gulf St. Lawrence, excell. ills.); Kendall, Copeia,
1919: 70 (Penobscot R., Maine, in 1919); Halkett, 53rd Rep. Fish., Br. Naval Serv., Canada (1919),
1920: 56 (Hudson Bay); Jespersen, Medd. Komm. Havunders. Kbh., Fisk. 6 (1), 1920: 11-16 (post-
larvae, around Iceland); Cox, Contr. Canad. Biol. (1918-1920), 11, 1921: I10 (rare at Magdalens,
one specimen, Cheticamp); Huntsman, Contr. Canad. Biol. (1921), 3, 1922: 60 (Bay of Fundy); Bigelow
and Welsh, Bull. U.S. Bur. Fish. (1924), 40 (1), 1925: 140 (descr., ill, Gulf of Maine); Bolkay, Novit.
Mus. Sarajevo., 3, 1925: 1-4 (fossil, Bosnia; not seen; see summ., ieee! Lond., 177, 1926: 136, 139);
Hubbs, Proc. biol. Soc. Wash., 38, 1925: 55 (range); Seemendeed Rit. Visind. intends (Soc. Sci. Islandica),
2, 1927: 41 (abund., season, beach spawn., age and death rel. to spawn., food, enemies, juv., Iceland);
Rachmanowa, Zool. Anz., 78, 1928: 119-123 (spawn. season, age, size at spawn., varietal stud., food,
increased range in w. Atlant.); Schroeder, Bull. U.S. Bur. Fish., 46, 1930: 71 (prey of cod); Schnaken-
beck, Faune Ichthyol. N. Atlant., Cons. int. Explor. Mer, 1931: pl. unnumbered (genl. acct., ill., fin-ray
and scale counts); Freund, Cetacea iz Grimpe and Wagler, Tierwelt Nord- u. Ostsee, 12 x, 1932: 44,
47 (prey of white whale, Be/uga, and north. porpoise, Phocaena); Jeffers, Contr. Canad. Biol., N.S. 7,
1932: 207 (abund., season around Newfoundland); Sleggs, Rep. Nfdld. Fish. Res. Comm., I (3), 1932:
66 pp., 2 pls., 2 charts (spawn. season and process, age of spawners, offshore recs., larvae, use, refs., New-
foundland); Martinsen, Trans. oceanogr. Inst. Moscow, 4 (1), 1933: 37-50, 15 charts (distr., Barents
Sea coast; Eng. summ.); Anon., Rep. Nfdld. Fish. Res. Comm. (1932), 2 (1), 1933: 125 (where taken
in trawl); Nikforovsky, Trans. oceanogr. Inst., Moscow, 4 (1), 1933: 54-64 (racial stud., Murman coast;
Eng. summ.); Rass, Trans. oceanogr. Inst., Moscow, 4 (1), 1933: 31 (depth and temp. of spawn., Barents
Sea); Vladykov, Contr. Canad. Biol., N.S. 8 (2), 1933; 8-12, figs. 2, 41 (Hudson Bay locals., abund.,
spawn. season); Awerinzew, J. Cons. int. Explor. Mer, 9 (1), 1934: 79-82 (distr., season, Barents Sea,
Novaya Zemlya); Anon., Rep. Nfdld. Fish. Res. Comm. (1933), 2 (2), 1934: 115 (where taken in trawl
and pelagic); Vladykov and Tremblay, Nat. Canad., 62, 1935: 79 (listed, Trois Pistoles, Quebec); Liib-
bert and Ehrenbaum, Handb. Seefisch. Nordeuropas, 2, 1936: 42 (descr., ill., develop., habits, food,
enemies, distr., methods of capture); Pfaff, Rep. 5th Thule Exped. (1921-1924), 2 (7), Zool. 11,
1937: 7 (Melville Penin., Arctic Canada); Schultz, Proc. U.S. nat. Mus., 85, 1937: 19 (cf. Capelin of n.
Pacific); Hildebrand, Medd. Gronl., 125 (1), 1939: 5 (e. Greenland, n. Labrador); Chapman, J. Morph.,
69, 1941: 279-301 (skel. chars.); Hoygaard, Skr. norske VidenskAkad. (1940), Mat.-Nat. Kl. 9, 1941:
17, 42 (abund., season, e. Greenland); Vladykov, Nat. Canad., 68, 1941: 193 (sex. dimorph. of anal
fin), 201 (cf. M. catervarius, refs.); Hansen, Ann. Biol., Cons. int. Explor. Mer, I, 1943: 121-124
(length-age relation., no. vertebrae, age at spawn., e. and w. Greenland); Breder, Field Bk. Mar. Fish.
Atl. Cst., 1948: 74 (ill., range, habits); Jensen, Spolia zool. Mus. Hauniensis, 9, 1948: 79-92 (habits,
spawn., abund., distr., w. and e. Greenland); Templeman, Bull. Nfdld. Govt. Lab., 17, 1948: 151 pp.
(genl. life hist., Newfoundland, refs.); Saemundsson, Zool. Iceland, 4 (72), 194.9: 93 (no subspp. recogn.);
Dannevig, Ann. Biol., Cons. int. Explor. Mer, 8, 1951: 15—16 (nos. distr. larvae, n. Norway, expansion
southward, 1915); Dunbar and Hildebrand, J. Fish. Res. Bd. Canad., 9 (2), 1952: 103 (Ungava Bay);
Bigelow and Schroeder, Fish. Bull. (74) U.S. Fish Wildl. Serv., 53, 1953: 134 (descr., ill., Gulf of
Maine); Backus, Bull. Amer. Mus. nat. Hist., 773 (4), 1957: 295 (Labrador, L. Melville, from seal
stomachs).
38
594 Memoir Sears Foundation for Marine Research
Osmerus arcticus Nilsson, Prod. Ichthyol. Scand., 1832: 11 (diagn., spawn., food of cod, Arctic seas ton. Norway,
refs.); Nilsson, Skand. Fauna, Fisk., 1855: 441 (refs., descr., habits, distr.).
Mallotus groenlandicus Cuvier, Anim. Kingd., Pisces, Griffith ed., 10, 1834: 420 (diagn.).
Salmo (Mallotus) villosus Richardson, Fauna Boreal.-Amer., 3, 1836: 187-190 (genl. range, habits, abund.,
mouth of Back R., trib. to Bathurst Inlet, Arctic Canada, descr., meas., fin-ray counts).
Ma/lotus arcticus Kroyer, Danmarks Fiske, 3, 1846-1853: 23 (descr., ill., habits, range; in Danish); Rae, White
Sea Peninsula, 1881: 61 (Murman coast); Smitt, Svenska VetenskAkad. Handl. (1885), N.F. 27 (8),
1886: 189-196 (aercticus of Greenland cf. vi//osus of Scandinavia).
Osmerus microdon Cuvier and Valenciennes, Hist. Nat. Poiss., 27, 1848: 385, pl. 621 (descr., Norway).
Mal/lotus villosus (in part) Giinther, Cat. Fish. Brit. Mus., 6, 1866: 170 (refs., descr., Greenland, Hudson Bay,
but subsp. catervarius includ. by ref. to Kamchatka); Jordan, Rep. U.S. Comm. Fish. (1885), 1887: 830
(Greenland, Alaska); Jordan and Evermann, Rep. U.S. Comm. Fish. (1895), 21, 1896: 294 (Arctic
Greenland S. to C. Cod; Alaska; Kamchatka); Bull. U.S. nat. Mus., 47 (1), 1896: 520; 47 (4), Ig00:
pl.85, fig. 225 (descr., ill., distr.; socia/is Pallas 1814 includ. as synon.); Halkett, Check List Fish. Canad.,
1913: 55 (Greenland, Labrador to C. Cod; refs. to Kamchatka and Brit. Columbia for subsp. cater-
varius); Jordan, Evermann, and Clark, Rep. U.S. Comm. Fish. (1928), 2, 1930: 66 (refs., range, socia/is
Pallas, islands between Asia and Amer. includ.).
Mallotus villosus villosus Rumjanev; Bull. Pacif. sci. Inst. Fish., Vladivostok, 22, 1947: 35-75 (Eng. summ.;
not seen).
Anchovas or Capelinas, Parkhurst iz Hakluyt, Voy. Navig. Traff. Discov. English Nation, 3, 1600: 133 (abund.,
strandings, pursued by cod, Newfoundland).
Capelin de l’Amerique septentrionale, Duhamel (collab. de La Marre), Traité Péch., 2 (1), 1769: 149, pl. 26,
figs. 1-8 (descr., ills., sex. dimorph., abund., method of capture, palatability, Newfoundland Bank).
Capelan, Pennant, Arctic Zool., Suppl., 2, 1787: 141; also 2, 1792: 349 (refs., descr., abund., use, Iceland,
Greenland, Newfoundland); Sars, Rep. U.S. Comm. Fish. (1880), 8, 1883: 167-187 (habits, migr.,
spawn., abund., season, n. Norway to White Sea); Hjort, Rapp. Cons. int. Explor. Mer, 20, 1914: 113—
114 (migr., Barents Sea); Bronkhorst, Notes Off. Péch. Marit., Paris, 53, 1927: 75, 151 (abund., St.
Pierre and Miquelon, cod bait, exports to France, to U.S.); Vladykov, Chasse, Biol. Val. Econ. Marsouin
blanc... Fleuve Golfe Saint-Laurent, Dept. Péch., Quebec, 1944: 117 (leading item in diet).
Capelin, Chappell, Voy. Rosamonp to Newfoundld., Labrador, London, 1818: 131-134 (spawn. activ., abund.,
Newfoundland); Anspach, Hist. Nfdld., 1819: 400 (abund., method of capture, Conception Bay, New-
foundland); Perley, Rep. Sea River Fisher. N. Brunsw., ed. 2, 1852: 30, 34, 37, 46-49, 52, 62 (cod
bait and manure, rel. to cod fishery, abund.); Rink, Danish Greenland, 1877: 135 (abund., spawn.
season, uses w. Greenland; Eng. transl.); Hind iz Goode, et a/., Fish. Fish. Industr. U.S., 1 (3), 1884:
545, 546 (winter. grounds, Labrador n. limit); Atwood, Fish. Fish. Industr. U.S., 4, Fishermen, 1887:
151 (spawn., bait, s. Labrador); Thompson, Rep. Nfdld. Fish. Res. Comm., I (1), 1930: 43-44 (spawn.
temp., possible commerce. exploit., import. as food for cod, Newfoundland); Jeffers, Ann. Rep. biol. Bd.
Canada (1930), 1931: 17-18 (age at spawn., incubation rel. to temp.); Sleggs, Rep. Nfdld. Fish. Res.
Comm. (1931), I (4), 1932: 34, 35 (summ. of invests., season, temp., predom. of 3-year fish, spawn.,
larvae).
Capeling, Audubon, Ornithol. Biogr., 2, 1834: 526 (abund., n. shore Gulf of St. Lawrence).
Lodde, Hansen and Hermann, Danm. Fisk. Havunders., 15, 1953: 85-88 (genl. habits, range, abund., com-
merc. import., Greenland; in Danish).
eee ee
NOK to
coms
Io.
IT.
I2.
Te
T4.
T5.
16.
17.
18s.
19.
TEXT AND FOOTNOTE REFERENCES
Avcassiz, L., Proc. Boston Soc. nat. Hist. (1848—
1851), 3, 1851.
Anspach, Hist. Newfoundld., London, 1819.
Backus, Bull. Amer. Mus. nat. Hist., 773 (4),
1957.
Berg, Freshwater Fishes of Russia, ed. 2, 1923.
Berg, Zoogeographica, Jena, I, 1933.
Berg, Class. Fish., Trav. Inst. Zool. Acad. Sci.
URSS, 5, 1940; reprinted by J. W. Ed-
wards, Ann Arbor, Mich., 1947 (Russ. and
Eng.).
Bigelow, Bull. U.S. Bur. Fish., go (2), 1928.
Bloch, Oecon. Naturg. Fische Dtsch., 1, 1782.
Bolkay, Novit. Mus. Sarajevo., 3, 1925; not
seen, but see Nature, Lond., 117, 1926:
136, 139.
Bronkhorst, Notes Off. Péch. Marit., Paris, 53,
1927.
Cuapman, J. Morph., 69, 1941.
Chappell, Voy. Rosamonp to Newfoundld., and
Labrador, London, 1818.
Clemens and Wilby, Bull. Fish. Res. Bd. Canad.,
68, 1949.
Collett, Forh. Vidensk. Selsk., Christiania
(1903), 9, 1903.
Collins and Smith, Bull. U.S. Fish Comm.
(1890), Io, 1892.
Cope, Proc. Amer. philos. Soc., rr, 1871.
Creaser, Pap. Mich. Acad. Sci., 5, 1925.
Cuerrier, Fry, and Préfontaine, Nat. Canad.,
73> 1946.
Cuvier and Valenciennes, Hist. Nat. Poiss., 27,
1848.
595
20.
2I.
34:
35:
36.
Dasnevic, Canad. Fish. Exped. (1914-1915),
Dept. Naval Serv., 1919.
Duhamel (collab. de La Marre), Trait. Péch.,
2 (1), 1769.
Dunbar and Hildebrand, J. Fish. Res. Bd.
Canad., 9 (2), 1952.
Dymond, Copeia, 1937.
Dymond, Canad. Field Nat., 58, 1944.
Enrennavum, Mitt. dtsch. Seefisch. Verein,
Beilage, 10, 1894.
Evermann and Goldsborough, Proc. biol. Soc.
Wash., 20, 1907.
Fasrrcrus, Fauna Groenl., 1780.
Fowler, Proc. Acad. nat. Sci. Philad., 63, 1912.
Garmaro,Voy. Islande et Groenl., Atlas, Zool.,
Poiss., 1851.
Goode, et a/., Fish. Fish. Industr. U.S., 1 (3),
1884.
Graham, D. H., Treasury of New Zealand
Fishes, 1953.
Gregory, Trans. Amer. philos. Soc., 23 (2),
1933.
Grimpe and Wagler, Tierwelt Nord- u. Ost-
see, 12 K, 1932.
Ginther, Cat. Fish. Brit. Mus., 6, 1866.
Haxzvvr, Voy. Navig. Traff. Discov. English
Nation, 3, 1600.
Halkett, Check List Fish. Canad., 1913.
38°
596
37:
Hansen, P. M., Ann. Biol., Cons. int. Explor.
Mer, I, 1943.
Hardy, Proc. N.S. Inst. Sci., r (2), 1867.
Hart, J. biol. Bd. Canad., 3 (5), 1937.
Hart and McHugh, Bull. Fish. Res. Bd. Canad.,
64, 1944.
Hildebrand, Medd. Gronl., 725 (1), 1939.
Hoover, Copeia, 1936.
Hubbs, Proc. biol. Soc. Wash., 38, 1925.
Hubbs and Chapman, Copeia, 1951.
Huntsman, Oceanography iz Handb. of Canada,
Univ. of Toronto Press, 1924.
: Teeree, Ann. Rep. biol. Bd. Canad. (1930),
19313 (1931) 1932.
Jeffers, Contr. Canad. Biol., N.S. 7 (16),
[A 13], 1932.
Jensen, Spolia zool. Mus. Hauniensis, 9, 1948.
Jensen, Vidensk. Medd. dansk naturh. Foren.,
Kbh., 171, 1949.
Jespersen, Medd. Komm. Havunders. Kbh.,
Fisk., 6 (1), 1920.
Jordan and Evermann, Bull. U.S. nat. Mus., 47
(1), 1896.
Jordan, Evermann, and Clark, Rep. U.S.Comm.
Fish. (1928), 2, 1930.
Jordan and Gilbert, Bull. U.S. nat. Mus., 76,
1882.
Kenpaut, Proc. Portland Soc. nat. Hist., 2(8),
1909.
Kendall, Copeia, 1917.
Kendall, Bull. U.S. Bur. Fish. (1926), 42,
1927.
Kessler, Fishes Aralo-Caspian Exped., 1877.
Kumlien, Bull. U.S. nat. Mus., 75, 1879.
Lanetors, Copeia, 1935.
Lanman, Rep. U.S. Comm. Fish. (1872-1873),
2, 1874.
Livingstone, Proc. N.S. Inst. Sci. (1950-1951),
23 (1), 1953.
Low, Annu. Rep. geol. Surv. Canad., App. 11
to Rept.1, N.S. 8, 1896.
Liibbert and Ehrenbaum, Handbk. Seefisch.
Nordeuropas, 2, 1936.
64.
65:
66.
67.
68.
69.
70.
fake
72.
Memoir Sears Foundation for Marine Research
Mascorrz, Contr. Sta. biol. Saint-Laurent,
18, 1948.
McKenzie, Bull. Fish. Res. Bd. Canad., 70,
1946.
Mori, J. Chosen nat. Hist. Soc., 11, 1930.
Miller, Zool. Danicae Prod., 1777.
N IKFoROvsKY, Trans. oceanogr. Inst. Moscow,
4 (1), 1933-
Norris, Proc. Acad. nat. Sci. Philad. (1861),
1862.
Norris, Proc. Acad. nat. Sci. Philad., 20, 1868.
Oocsren, van, Copeia, 1937.
Oosten, van, Trans. Amer. Fish. Soc. (1944),
74; 1947-
P allas, Zoogr. Rosso-Asiat., 3, 1831.
Parker, Philos. Trans. (1873), 163 (1), 1874.
Pennant, Arctic Zool., r, Introd., 1784.
Perley, Cat. [in part] Fish. N. Brunsw., Nova
Scotia iz Rep. Fisher. Bay of Fundy, 1951.
Petrow, Zool. Anz., 107, 1934.
Pfaff, Rep. 5th Thule Exped. (1921-1924),
2 (7), Zool. 11, 1937.
Pitt, J. Fish. Res. Bd. Canad., 75 (3), 1958.
eexcrmancnsiae Zool. Anz., 75, 1928.
Rass, Trans. oceanogr. Inst. Moscow, 4 (1),
1933 (English Summary).
Reeks, Zoologist, London, 2 (6), 1871.
Rep. Mass. Comm. Fish. (1868), 1869.
Rep. Mass. Comm. Fish. Game (1916-1917),
52nd, 1918; (1917-1918), 53rd, 1919;
(1918-1919), 54th, 1920; (1919-1920),
55th, rg2t.
Rep. U.S. Comm. Fish. (1897), Append., 1898.
Rice, Rep. Comm. Fish. Md., 1878.
Richardson, John, iv Franklin, Narr. ... Jour-
ney to Polar Sea, Append. 6, Fishes, 1823.
Richardson, Fauna Boreal.-Amer., 3, Fish.,
1836.
Richardson, Copeia, 1942.
Rink, Danish Greenland, 1877.
Rumjaney, Bull. Pacif. sci. Inst. Fish., Vladivos-
tok, 22, 1947 (not seen).
92.
93-
94-
95-
96.
97-
9s.
99-
Too.
Tor.
102.
103.
To4.
I05.
roo.
107.
Tos.
Fishes of the Western North Atlantic
SaEMuNDssON, Rit. Visind. Islend., 2, 1927.
Saemundsson, Zool. Iceland, 4 (72), 1949-
Sars, Rep. U.S. Comm. Fish. (1880), 8, 1883.
Scheuring, Ergebn. Biol., 5, 1929.
Schmidt, Medd. Komm. Havunders. Kbh.,
Fisk. 2 (4), 1906.
Schmitt, Mongr. Isle d’Anticosti, 4, Zool.,
Poiss., 1904.
Schultz, Proc. U.S. nat. Mus., 85, 1937.
Sherborn, Ibis, 4, 1934.
Sleggs, Rep. Nfdld. Fish. Res. Comm. (1931),
I (3), 1932.
Sleggs, Rep. Nfdld. Fish. Res. Comm. (1931),
T (4), 1933.
Smitt, Ofvers. VetenskAkad. Forh., Stockh.
(1882), 39 (8), 1883.
Smitt, Svenska VetenskAkad. Handl. (1885),
N.F. 2z (8), 1886.
Smitt, Hist. Scand. Fish., ed. 2, 2, 1895.
Soldatov and Lindberg, Rev. Fishes Far East,
Bull. Pacif. sci. Inst. Fish., Vladivostok, 5,
1930 (Russ.).
Starks, Stanf. Univ. Publ. Biol., 4 (3), 1926.
Steindachner and Kner, S.B. Akad. Wiss.
Wien, Mat-Nat., 62, 1870.
Stellers, Beschreibung... Kamtschatka, 1774.
T09.
Ito.
ITl.
If2.
IT3.
IT4.
IT5.
Ir6.
II7.
rr8s.
Dy
aytor, Bigelow, an raham, Fish. Bull.
ly Bigel d Graham, Fish. Bull
(115) U.S. Fish. Wildl. Serv., 57,
1957-
Templeman, Bull. Newfoundld. Govt. Lab.,
17, 1948.
Trade News, Canad. Fish., 8 (5), 1955.
Wiaanykoy: Contr. Canad. Biol., N.S. 8 (2),
1933-
Vladykov,
1934-
Vladykov, Nat. Canad., 68, 1941.
Vladykov and Tremblay, Nat. Canad., 62,
1935-
Trans. Canad. Inst., 20 (1),
Warrers, Bull. Amer. Mus. nat. Hist., 706
(5) 1955.
Wiborg, Ann. Biol., Cons. int. Explor. Mer
(1851), 8, 1852.
Wynne-Edwards, Bull. Fish. Res. Bd. Canad.,
94, 1952.
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INDEX OF COMMON NAMES
Principal references are given in parentheses. The letters that accompany some page numbers indicate
references to the following: bait (B), disease (D), parasites (P), predators (E), or prey (F).
Arrican catfish, Sc
Alabama shad, (308), 315, 317.
Alewife, 275, 302, (312), (324), (332), (365),
(393), 476, 482F, 536F.
Alligator gar, 66, 67, (83).
Amberjack, 355 E.
American shad, 39, (295), 309-311, 315, 336,
337s 340-
American smelt, 275, (559), 576, 581.
Anchoa, la, (158).
Anchovas, 594.
Anchovy, 94, 118F, 439.
Bay, 79F, (176).
Bermuda, (399).
Cuban, (188).
Dusky, (200).
Flat, (213).
Key, (173).
Longnose, (192).
New Jersey, (170).
Silver, (208).
Striped, (194).
Anglerfish, 3, 7, 9-
Angleworm, 482B.
Apapa, (417), (419), (421), (428).
Arctic charr, or charr, 15, 460, 482, 498, 5o4—
506, (507), 525, 531, 544, 588 E.
Arctic cod, 482F.
Arctic trout, 524.
Asuna Nahui, (421).
Atlantic cod, 588E.
Atlantic herring, (275), 343, 405, 439, 581, 588E.
Atlantic menhaden, 270, (346), 370.
Atlantic round herring, (263).
Atlantic salmon, 52, 457, 459, (460), 498, 507,
517s 519, 522, $25, 528, 532, 535, 537, 539
54, 543, 588E.
599
Atlantic shad, (295).
Atlantic sturgeon, 39, 45, (46).
Atlantic thread herring, (381).
Ayu, 456.
Bacrerua, 286P.
Bananafish, (134).
Banded killifish, 44 F.
Bang, (401).
Barnacle, 303 F.
Bass, 303 E.
Striped, 323, 536E.
Batfish, 7.
Bay anchovy, 79F, (176).
Beach flea, 536F.
Beach fly, 588P.
Beetle, 481 F.
Belted kingfisher, 483 E.
Benane, (124).
Bermuda anchovy, (399).
Bermuda herring, (272).
Bermuda sardine, 404.
Big sturgeon, 46.
Bigeye herring, 94, (124), (332).
Bird, 31 F, 62, 338E, 483E, 497E.
Water, 153E, 178E, 198E, 303E, 355E,
404E.
Bivalve, 44F, 52F, 139F, 284F (larva).
Blackbelly, (324).
Black fly, 536F.
Black Sea sturgeon, 29.
Blenny, 6, 7.
Blue crab, 67F, 74F, 79F, 86F, 118F.
Blue herring, (315), (324).
Blueback herring, or blueback, 321, (324), 332,
334, 336-339-
600 Index of Common Names
Bluefish, 153E, 355E, 384E.
Bocén, (155), (188), (245).
Bonefish, 15, 94, I15, 124, 129, (133), (134)-
Shafted, (143).
Bonejack, (319).
Bowfin, 13.
Branch herring, (332).
Bristle herring, (381).
Brook trout, 460, 483E, 484, 498, 507, 519, (525).
Brown trout, 460, 461, 484, (498), 525.
Bugfish, (346).
Bunker, (346).
Burrowing May fly, 44 F (larva), 52F (larva).
Capois fly, 318F, 481F (larva), 536F.
Cagona, (419).
Caille, (423).
Camus, 41.
Capelan, 594.
Capelin, 275, 285F, 482F, 518F, 553, 559,
(573), (576).
Capelin de I’Amerique septentrionale, 594.
Capelinas, 594.
Capeling, 594.
Catfish, 5, 6, r18F.
African, 5.
Sea, 79F.
Characin, 428.
Charr (or Arctic charr), 15, 460, 482F, 498, 504—
506, (507), 525, 531, 544, 588E.
Red-spotted, 506, 522.
Chub, 548.
Chub mackerel, 7.
Cisco, 339, 548.
Cod, 285E, 355E, s19F, 586F, 593 E.
Atlantic, 588E.
Arctic, 482 F.
Greenland, 588E.
Colonial hydroid, 330P.
Common shad, (295).
Common sturgeon, (46), (56).
Conch, 140B.
Cormorant, 483E, 537E.
Crab, 139F, 140B, 285F (larva), 322F, 384F,
482F, 564F.
Blue, 67F, 74F, 79F, 86F, 118F.
Rock, 536F.
Crayfish, 140B, 537F.
Crevalle jack, 129F.
Cuban anchovy, (188).
Cuffum, (113).
Cunner, 337F.
Cutlassfish, 118 F.
Disease, Boil, 484D.
Fungous, 286D.
Furunculosis, 484 D.
Pigment spot, 286D.
Salmon, 484D.
Tail rot, 286D.
Ulcer, 286D, 484D.
White spot, 484 D.
Dogfish, 285 E.
Spiny, 537E, 588E.
Dolly Varden, 484, 523.
Dolphin, 355E.
Duck, 86F.
Dusky anchovy, (200).
Dwarf herring, (267), (268), (272).
Eastern gizzard shad, (444).
Ecailles, 46.
Eel, 5, 6, 12, 132, 139, 337, 476, 482F, 4835,
536F, 537E.
Gulper, 2.
Eelgrass, 538 F.
Eelpout, 588 E.
Eggs (copepod), 284F.
Eggs (fish), 303F, 322F, 337F.
Escargot, 41, 46.
Esturgeon d’eau salée, 46.
Esturgeon jaune, 41.
Esturgeon noir, 46.
European sturgeon, 58.
European trout, 10, 459, (498).
Faw herring, (319).
False pilchard, (390).
False sardine,(40r).
Fatback, (346).
Fatback minnow, (393).
Filefish, 7.
Finback whale, 285 E, 303E, 338E, 588E.
Finescale menhaden, (376).
Index of Common Names 601
Flat anchovy, (213).
Flatfish, 482 F.
Flea, Beach, 536F.
Sand, 482F.
Florida gar, 67, 75, (81).
Florida spotted gar, (81).
Flounder, 588 E.
Fluke, 484 P.
Fly, Beach, 588P.
Black, 536F.
Burrowing May, 44F (larva), 52F (larva).
Caddis, 318F, 481F (larva), 536F.
May, 318F, 481 F (nymph), 536F.
Stone, 481 F.
Freshwater herring, (332).
Freshwater taylor, (319).
Fry, Green, (268), (272).
Gasyer, 537.L-
Gar or garfish, 6, 7, 13-16, 62-67, (68), 91,
2s
Alligator, 66, 67, (83).
Florida, 67, 75, (81).
Longnose, 15, 66, 67, (75).
Spotted, 66, (71).
Gaspereau, (332).
German brown trout, 499.
Ghost smelt, 554.
Giant tuna, 290E.
Gill maggot, 484P, 495.
Gizzard shad, 318F, (443), (444), 449, 450.
Glut herring, (324), 332.
Goby, 3.
Gold or golden shad, (315).
Grand ecaille, (113).
Grass shrimp, 285 F.
Gray herring, (332).
Gray snapper, 270E.
Gray seal, 483E.
Gray trout, (543).
Grayback, 321, 329-331, (332)-
Grayling, 8, 456.
“Green food,” 284F.
Green fry, (268), (272).
Green herring, (315).
Greenland cod, 588E.
Greenland halibut, 588E.
Greenland shark, 588E.
Gros hareng, (419).
Grubber, (134).
Guillemot, 588E.
Gulf menhaden, (365).
Gulf of Mexico sturgeon, (56).
Gulf shad, (308).
Gull, 62, 483E, 588E.
Gulper eel, 2.
Gunnel, Rock, 275, 536 F, 581.
Hacuvpo, (155).
Haddock, 285, 482F.
Hairyback, (381).
Hake, 355E, 536F.
Silver, 285E, 286E, 537E.
Halibut, Greenland, 588 E.
Harbor seal, 483 E.
Hareng gras sardinha, 229.
Herring, or Atlantic herring, 7, 15, 94, 251, 258,
(275) 337F, 354, 482F, 564F.
[For discussion of commercial or fishery terms—
bloater, brit, fat, kipper, mustard, oil, red herring,
sardine, snipper, spawn—see pp. 279, 280, 284,
288.]
Atlantic, (275), 343, 405, 439, 581, 588E.
Atlantic Round, (263).
Bermuda, (272).
Bigeye, 94, (124), (332)-
Blue, (315), (324).
Blueback, 321, (324), 332, 334, 336-339.
Branch, (332).
Bristle, (381).
Dwarf, (267), (268), (272).
Fall, (319).
Freshwater, (332).
Glut, (324), 332.
Gray, (332).
Grayback, 321, 329-331, (332):
Green, (315).
River, (312), (315), 332-
Round, 98, (262).
Sea, (275).
Shad, (319), (381).
Skipjack, 311, (315), 343-
Summer, (324).
White, (332).
Heron, 483E, 537E.
Herring smelt, 8, 554.
602 Index of Common Names
Hickory jack, (319).
Hickory shad, 275, 315, (319), 343-
Holosteen, 93.
Humpback salmon, 458.
Hydroid, Colonial, 330P.
Lcerisu, 456, (559).
Ichthyostés, 14.
Inconnu, 547.
Insect, 31F, 52F, 302F, 302F (larva), 303F,
310F, 318F, 318F (larva), 481F (larva),
482F, 486 F, 494, 518F, 536F.
ee Crevalle, 129 F.
Hickory, (319).
Jamais-gouté, (212).
Jewfish, (113).
Key anchovy, (173).
Killifish, 536F.
Banded, 44F.
Kingfisher, 486E, 495E, 496E.
Belted, 483 E.
Kyak, (324), (332).
Lapvrisx, 15, 94, (123), (124).
Lake salmon, 493.
Lake sturgeon, 31, (41), 47, 53, 54-
Lake trout, 460, 493, 504, 507, 526, (542), (543),
564.
Lake whitefish, (549).
Lamprey, 483 E.
Largescale menhaden, 79F, (365).
Launce (or Sand launce), 52F, 275, 329F, 337F,
482F, 518F, 58r.
Lice, Sea, 484P, 519P, 537P.
Lochleven trout, (498).
Lodde, 594.
Long john, (124).
Longnose anchovy, (192).
Longnose gar, 15, 66, 67, (75).
Loosescale sardine, (387).
Lungfish, 2-4, 9, 10, 12.
Macasr, (124).
Mackerel, 7, 153, 281, 285E, 323, 354, 473,
482F.
Chub, 7.
Spanish, 384 E.
Mackinaw trout, (543).
Maggot, 588P.
Gill, 484P, 495 P.
Manjtia, (173), (181), (188), (192), (194), (200),
(213).
Matejuelo blanco, (124).
Matejuelo real, (124).
May fly, 318F, 481 F (nymph), 536F.
Menhaden, 116, 281, 332, (342), (346), 384.
Atlantic, 270, (346), 370.
Finescale, (376).
Gulf, (365).
Largescale, 79 F, (365).
Merganser, 483E, 486E, 495E, 497E.
Midge, 302F (larva), 536F.
Milkfish, 15, 89.
Mink, 537E.
Minnow, 302F, 318F.
Fatback, (393).
Mooneye, 318F.
Mossbunker, (346).
Mouse, 482F.
Mullet, 86F, r18F.
Murres, 588 E.
Mussel (freshwater), 318 P (larva).
Nezprerisn, 272E.
New Jersey anchovy, (170).
New Zealand smelt, 553, 554.
Ono shad, (308), 315.
Otter, 537E.
Paciric salmon, 484.
Paddlefish, 5, 6, 8, 11-14, 26.
Palika, (113).
Pelican (bird), 384E.
Pelican (sturgeon), 46.
Penguin, 62.
Perch, white, 476.
Petit cailleu, (390).
Index of Common Names 603
Bikesess 75104.
Pilchard, (387), (393)-
False, (390).
Red-ear, (387).
Pincer, (387).
Pipefish, 9.
Piquitinga, (186), 198.
Pirapema, (419).
Plover, 62.
Pogy, (346), 354-
Pollock, 285E, 355E, 536E.
Porcupinefish, 7.
Porpoise, r18E, 355E, 488E, 588E, 593E.
Potomac shad, 311.
Prawn, 482B.
Puffin, 588E.
Rarwsow trout, 8, 459, 460, (499), 525-
Ray, 5, 7, 8.
Torpedo, 6.
“Red seed” (copepod), 284 F.
Red-ear pilchard, (387).
Red-ear sardine, (387).
Red-spotted charr, 506, 522.
Remora, 6, 117.
River herring, (312), (315), 332-
Rock crab, 536F.
Rock gunnel, 275, 536F, 58r.
Round herring, 98, (262).
Atlantic, (263).
Round whitefish, 548, 550, 551.
Roundworm, 484P, 519P.
Ruffed grouse, 488.
Sasato real, (113).
Sailfish, 6.
Salmon, 3, 5, 7, 94, 302, (459), 482F, 503, 516,
541, 553, 564, 566.
Atlantic, 52, 457, 459, (460), 498, 507, 517,
519, 522, 525, 528, 532, 535, 537» 539, 540,
543, 588E.
Humpback, 458.
Lake, 493.
Pacific, 484.
Sea, (460).
Silver, 458.
White Sea, 493.
Salmon trout, (525).
Salter, (525).
Sand flea, 482F.
Sand launce, or launce, 52F, 275, 329F, 337F,
482F, 518F, 58r.
Sand seatrout, 343.
Sardina, (387), (393), (401), (421).
Sardina boca torta, (155).
Sardina escamuda, (390), (393).
Sardina de Espafia, (401).
Sardina de Ley, (387).
Sardine, (167), (386), (387), (393).
Bermuda, 404.
False, (401).
Loosescale, (387).
Red-ear, (387).
Scaled, (393).
Spanish, (397), (401), (407).
Sardinha, (393).
Sardinha d’agua doce, (428).
Sardinha grande, (419).
Sardinha large, (381).
Sardinha prata, (234).
Sardinha sargo, (381).
Sardinha verdadeira, (407).
Sardinia, (407).
Sardinia de Espafia, (407).
Sargo, (381).
Sargo de gato, (381).
Sawbelly, (332).
Scaled sardine, (393).
Sculpin, 482F, 518F, 588E.
Sea catfish, 79F.
Sea herring, (275).
Sea lice, 484P, 519P, 537P.
Sea salmon, (460).
Sea sturgeon, (46).
Sea trout, (525).
White, (525).
Sea worm, 536F, 566B.
Seahorse, 9.
Seal, 31 F, 537E, 586E, 588E, 593 E.
Gray, 483E.
Harbor, 483E.
Searaven, 536F.
Seatrout, 153E, 198E.
Sand, 343.
Shad, 44F, 55, 275, (293), (295), 321, 329 3375
343» (346), (365).
604 Index of Common Names
Shad (cont.)
Alabama, (308), 315, 317-
American, 39, (295), 309-311, 315, 336, 337,
340-
Atlantic, (295).
Common, (295).
Eastern gizzard, (444).
Gizzard, 318F, (443), (444), 449, 450-
Gold or Golden, (315).
Gulf, (308).
Hickory, 275, 315, (319), 343.
Ohio, (308), 315.
Potomac, 311.
Threadfin, (443), (448).
White, (295).
Yellowfin, (372).
Shad herring, (319), (381).
Shafted bonefish, (143).
Shark, 4, 6, 8, 118E, 285E, 355E, 483E, 490E.
Greenland, 588E.
Sharpnose sturgeon, 46.
Shearwater, 588E.
Sheefish, 547.
Shellfish, 564F.
Shiner, 128, (393).
Shortnose sturgeon, (36), 45, 47-
Shovelnose sturgeon, 27.
Shrimp, 26 (penaeid), 52F, 128F, 139F, 283,
284F, 303F, 303 F (euphausiid), 329 F, 337F,
338F, 384F, 482F, 564F, 566B.
Grass, 285 F.
Silver anchovy, (208).
Silver hake, 285 E, 286E, 537E.
Silver salmon, 458.
Silverfish, (113).
Silverking, (113), 140.
Silverside, 118F, 536F, 555, 559, 576, 591-
Skate, 5.
Skipjack, (124), (315).
Skipjack herring, 311, (315), 343-
Sludgeworm, 39F, 52F.
Smelt, 482F, 536F, (553), (554)-
American, (559), 576, 581.
Ghost, 554.
Herring, 8, 554.
New Zealand, 553, 554.
White Sea, 572.
Smoothback, (41).
Snail, 536F.
Snake, 338E.
Water, 303 E.
Snapper, 266E, 267E.
Gray, 270E.
Spanish mackerel, 384 E.
Spanish sardine, (397), (401), (407).
Speckled trout, 530.
Spiny dogfish, 537E, 588E.
Sponge, 374F.
Squid, 128F, 139F, 140B, 285 FE, 322F, 337F,
564F.
Spotted gar, 66, (71).
Sprat, (387), (390), (393), 482F.
Squaretail trout, 505, (525).
Steelhead trout, (499).
Stickleback, 536F.
Stone fly, 481F.
Striped anchovy, (194).
Striped bass, 323, 536E.
Sturgeon, 3, 7, 8, 11-16, (26), go.
Atlantic, 39, 45, (46).
Big, 46.
Black Sea, 29.
Common, (46), (56).
European, 58.
Gulf of Mexico, (56).
Lake, 31, (41), 47, 53, 54.
Sea, (46).
Sharpnose, 46.
Shortnose, (36), 45, 47-
Shovelnose, 27.
White, 31, 51.
Sucker, 339.
Summer herring, (324).
Sweetfish, 456.
Swellfish, 7.
Swordfish, 6, 355 E.
"Tapeworm, 484P, 519P.
Tarpon, 15, 94, III, (112), (113), 129, 134, 140.
Tarpum, (113).
Taylor, freshwater, (319).
Tenpounder, (124).
Tern, 404E, 406 E, 588E.
Thorn-headed worm, 484 P.
Threadfin, 7.
Threadfin shad, (443), (448).
‘Toadfish, 9.
——
Index of Common Names
Togue, (543).
Torpedo ray, 6.
Triggerfish, 7.
Trout, 3, 5, (459), 482F, 497, (498).
Arctic, 524.
Brook, 460, 483E, 484, 498, 507, 519, (525).
Brown, 460, 461, 484, (498), 525.
European, 10, 459, (498).
German brown, 499.
Gray, (543).
Lake, 460, 493, 504, 507, 526, (542), (543),
564.
Lochleven, (498).
Mackinaw, (543).
Rainbow, 8, 459, 460, (499), 525-
Salmon, (525).
Sea, (525).
Speckled, 530.
Squaretail, 505, (525).
Steelhead, (499).
White Sea, (525).
Trunkfish, 7.
Tullibees, 548.
Tuna, 355E.
Giant, 290E.
Turkey, Water, 86F.
Turtle, 303E, 338E.
Viswine hare, 488.
605
Warer bird, 153E, 178E, 198E, 303E, 355E,
404E.
Water snake, 303E.
Water turkey, 86F.
Weakfish, 199 E, 329E, 343, 384.
Whale, 355E.
Finback, 285 E, 303E, 338E, 588E.
White, 588E, 593E.
White bait, 271.
White herring, (332).
White perch, 476.
White Sea salmon, 493.
White Sea smelt, 572.
White Sea trout, (525).
White shad, (295).
White sturgeon, 31, 51.
White whale, 588E, 593E.
Whitebill, (387).
Whitefish, 8, 547.
Lake, (549).
Round, 548, 550, 551.
Wolffish, 588E.
Worm, 52F, 139F, 519F.
Sea, 536F, 566B.
Sludge, 39F, 52F.
Thorn-headed, 484 P.
Yeviowrn shad, (372).
INDEX OF SCIENTIFIC NAMES
Principal references are given in parentheses. The letters that accompany some page numbers indicate
references to the following: bait (B), disease (D), parasites (P), predators (E), or prey (F).
Assorti, Anchovia, 243.
Anchoviella, 243.
Lycengraulis, 233, 235, (242).
Osmerus, 556, 557, 572-
abbotti (subsp.), Osmerus mordax, 556, 572.
abbotti (var.), Osmerus mordax, 572.
abnormalis, Ilisha, 415.
acanthias, Squalus, 537E, 588.
Acanthocephala, 303P, 338P.
Acanthocephalan, 330P; 484 P (worm).
Acanthocephali, 303 P.
Acanthodii, 2.
Acipenser, 3, 27, (28).
anasimos, 46.
anthracinus, 46.
atelaspis, 46.
baeri, 32.
baeri x A. ruthenus, 32.
bairdi, 56.
brevirostris, 24, 29, 32-34, (36), 41, 42, 47, 48,
55+
brevirostris x A. sturio, 54.
brevirostrum, 40.
buffalo, 46.
carbonarius, 46.
cayennensis, 54, 50.
cincinnati, 46.
copei, 46.
dabryanus, 32.
dekayi, 41.
fulvescems, 24, 28-34, 36-38, 40, (41), 47, 48,
53> 54-
kirardi, 56.
glabra, 28.
guldenstadti, 29, 32, 58.
giildenstadti x A. ruthenus, 32.
helops, 28.
606
holbrooki, 56.
huso, 28.
kennicotti, 56.
kikuchit, 32.
kirtlandi, 46.
laevis, 46.
lamarii, 46.
lecontei, 56.
(Huso) lesueurii, 41.
liopeltis, 46.
macrohinus, 50.
macrostomus, 4.6.
maculosus, 46.
medirostris, 32, 33, 30.
megalaspis, 46.
microrhynchus, 41.
milberti, 56.
mitchilli, 56.
multiscutatus, 32.
nacaril, 32.
nertinianus, 46.
nudiventris, 29, 30, 32.
ohiensis, 46.
oxyrhynchus, 15, 25, 27, 29-34, 36-45, (46).
oxyrhynchus de sotoi, 57.
oxyrhynchus desotoi, 24, 36, 47, 54, (56).
oxyrhynchus oxyrhynchus, 24, 36, (46).
paranasimos, 46.
platyrhinus, 46.
primigenius, 32.
rafinesquii, 46.
rauchi, 46.
rhynchaeus, 46.
richardsoni, 46.
rosarium, 46.
(Huso) rostellum, 41.
rubicundus, 46.
Index of Scientific Names 607
rupertianus, 46.
ruthenus, 28, 29, 31, 32.-
ruthenus x A. baeri, 32.
ruthenus x A. giildenstadti, 32.
schrencki, 32.
schypa, 28.
serotinus, 28, 46.
sinensis, 32.
(Huso) sinus, 41.
stellatus, 28, 29, 32.
storert, 56.
sturio, 25, 27-29, 325 35> 471 49s 54, 50-58.
sturio x A. brevirostris, 54.
sturio oxyrhynchus, 56.
transmontanus, 29, 31-33, 36, 5I-
vultharius, 28.
Acipenserid, or Acipenseroid, 2, 5, 7, 8, 11-13.
Acipenseridae, 24, (26).
Acipenserini, 27.
Acipenseroidei, 14, 16, (24).
Actinopteri, 13.
Actinopterygia, 13.
Actinopterygian fishes, 5, ro.
Actinopterygii, 2, (11).
acus, Echinorhynchus, 303 P, 330P, 338P.
Acus maxima squamosa viridis, 79
adamantinus, Litholepis, 69.
adunca, Ascaris, 303 P.
aestivalis, Clupea, 331.
Pomolobus, 294, 312-314, 317, (324), 332-334,
338, 340-342, 362.
affinis, Elops, 127, 128, 130.
africana, Clupea, 415.
agassizii, Alepocephalus, 251.
Cylindrosteus, 75.
Salmo, 505, 537, 542-
Salvelinus, 542.
agassizit (subsp.), Salvelinus fontinalis, 542.
Agnatha, 2, 4.
Agrostichthyes, 7.
alabamae, Alosa, 294, 295, 304, (308), 315,
343-
ohiensis, Alosa, 312.
Alausa, 293.
sapidissima, 307.
striata, 385.
vulgaris, 293.
Alausella, 312.
alba, Rogenia, 274.
Albula, 92-94, 107, 108, 115, 123, 124, 127, 130,
132, (133), 143, 145.
s. Salmo albula, 141.
conorhynchus, 141.
nemoptera, 145.
parrae, 141.
plumieri, 141.
rostrata, 41.
vulpes, 115, 124, 126, 129, 131, 133, (134), 145,
410.
albula, Albula s. Salmo, 141.
Salmo, 141.
Albulid, 93, 94.
Albulidae, 14, 15, 92, 93, 98, 107, 108, 110, (132).
Albulina, 93, 94.
Albuloidae, 94.
Albuloidei, 94.
album, Scaphirhynchus, 28.
alcyon, Megaceryle, 483E.
Alepisauridae, 99.
Alepocephalid, go.
Alepocephalidae, 14, 89-91, 95, 97, 98, 103, 148-
150, (250), 254.
Alepocephaloidei, 93.
Alepocephalus, 8, 149.
agassizil, 251.
bairdit, 251.
productus, 251.
Aleuteridae, 7.
alipes, Salmo, 524.
alipes (subsp.), Salvelinus alpinus, 523, 524.
allecia, Clupea, 410.
alleganiensis, Salmo, 541.
alleni, Amplova, 220.
Anchoviella, 205, 207, (219).
Allosmerus, 553, 554.
Alosa, 257, 258, 260, 267, 275, (293), 312, 316,
320.
alabamae, 294, 295, 304, (308), 315, 343.
alabamae ohiensis, 312.
alosa, 304.
apicalis, 389.
bishopi, 392-
cyanonoton, 332.
elongata, 415, 416.
Jinta, 304.
lineata, 323.
mattowaccta, 323.
menhaden, 361.
608 Index of Scientific Names
ohiensis, 308, 310, 312.
preastabilis, 307.
sadina, 361.
sapidissima, 39, 294, (295) 309-312, 317, 343-
teres, 267.
tyrannus, 332, 362.
alosa, Alosa, 304.
Clupea, 293, 3°7-
alosae, Argulus, 338P.
alpinus, Salmo, 503, 505, 506, 523.
Salvelinus, 15, 458, 460, 491, 498, 504-506,
(507), 525, 526, 541, 542, 588E.
alipes, Salvelinus, 523, 524.
arcturus, Salvelinus, 524.
aureolus, Salvelinus, 524, 541, 542.
malma, Salmo, 523.
malma, Salvelinus, 484, 506, 519, 520.
marstoni, Salvelinus, 524.
naresii, Salvelinus, 523.
oquassa, Salvelinus, 523, 542.
stagnalis, Salvelinus, 523.
stagnalis (forma), Sa/velinus, 524.
altamazonica, Ilisha, 416, (417), 420, 421.
Ilisha (Neosteus), 419.
Neosteus, 419.
Pellona, 419.
amazonica, Clupea, 411, 413.
Heringia, 413, 415.
Ilisha, 417, (421).
Rhinosardinia, (412), 415.
Sardinella (Rhinosardinia), 413.
amazonicum, Pseudochirocentrodon, 416, 423.
americanus, Lepus, 488.
Macrozoarces, 588E.
Pristigaster, 429.
Amia, 62, 65, 93, 133-
immaculata, 133, 141.
Amiid, or Amioid, 13, 64, 65.
Amiidae, 12, 64.
Amiiformes, 64.
Amioidea, 13.
Ammodytes, 52¥, 275, 482 FB, 581.
Amphibia, 3, 62.
Amphiboidei, ro.
Amphipnoid, 6.
Amphipod, 44F, 52F, 284F, 285F, 302F, 303F,
3378, 482F, 518F, 536F, 564F.
Amplova, 204, 206, 217.
alleni, 220.
balboae, 204, 206, 228.
brevirostris, 217.
fuianensis, 219.
Jamesi, 228.
Amplova (subgenus), 206, 207.
jamesi, Anchoviella, (227).
Anarrhichas, 588E.
anasimos, Acipenser, 46.
Anchoa, 153, (162), 204, 223.
argenteus, 162, 163, 166, (169).
blackburni, 183.
brownii, 187.
cayorum, 162, 163, 166, (173), 181.
choerostoma, 162, 164, 166, (185), 204.
cubana, 162, 165, 166, (188).
curta, 181, 183.
duodecim, 162, 163, 166, (170).
filifera, 162, 165, 166, (202).
ginsburgi, 162, 165, 166, (190), 194.
hepsetus, 165, 187, 189, 190, 193, (194), 209,
Aiils pic
hepsetus colonensis, 162, 165, 166, 197, 200.
hepsetus hepsetus, 162, 165, 166, 200.
howelli, 203.
januaria, 162, 164, 166, 174, 178, 181, (183).
lamprotaenia, 162, 164-166, (192), 194, 199,
200.
Lyolepis, 162, 165, 166, 185, 186, (200), 202,
204.
mitchilli, 79F, 163, 170, 173, (176), 181-184,
198, 226.
mitchilli diaphana, 162, 164, 166, 174, 178, 181.
mitchilli mitchilli, 162, 164, 166, 178, 18r.
parva, 162, 164, 166, 174, 176, 178, 180, (181),
183, 184, 226, 227.
pectoralis, 162, 163, 166, (174), 183, 184.
spinifer, 162, 163, 166, (167), 169.
tricolor, 162, 164, 166, (186), 199, 200.
trinitatis, 162, 163, 166, (171).
Anchovia, 152, 153, (155), 162, 231.
abbotti, 243.
argyrophana, 210.
brevirostris, 204, 219.
brownii, 188, 194, 199, 200.
cayorum, 174.
choerostoma, 202.
choerostoma cayorum, 174.
clupeoides, (155), 159, 161, 221, 231.
cubana, 190.
Index of Scientific Names 609
duodecim, 171. lyolepis, 202.
elongata, 224. mitchilli, 175, 180, 181, 183, 184.
eurystole, 210. nattereri, 207, (220).
Sillifera, 204. pallida, 205, 207, (221).
gilberti, 247. parva, 183.
januaria, 184. perfasciata, 205, 206, 209-212, (213).
lepidentostole, 226. platyargyrea, 202.
macrolepidota, 158. robertst, 247.
mitchilli, 180, 181. salvatoris, 188.
mattereri, 221. spinifera, 169.
nigra, 155, 156, (158). surinamensis, 247.
pallida, 221, 223. trinitatis, 173.
parva, 182, 223. venexuelae, 221, 223.
perfasciata, 210, 215. Anchoviella (subgenus), 206.
platyargyrea, 202, 204. Anhinga anhinga, 86¥.
producta, 158. anna-carolina, Mugilomorus, 123, 131.
rastralis, 156, 161. Annelid (worm), 197F, 285F, 355F, 564F.
Spinifera, 168. Anomalopterichthys, 250.
trinitatis, 172. Anomalopterus, 97, 250.
anchovia, Clupea, 4.06. Anotopteridae, ror.
Sardinella, 393, 398-400, (401), 409-411. Antaceus, 28.
Sardinia, 397, 407. anthracinus, Acipenser, 46.
Anchoviella, 153, 162, 200, (204). antipodum, Clupea, 287.
abbotti, 243. apapae, Ilisha, 421-423.
alleni, 205, 207, (219). apicalis, Alosa, 389.
argyrophana, 210. Clupea, 389.
astilbe, 190. Sardinella, 389.
balboae, 227. Aplochitonidae, g2.
blackburni, 204, 205, 207, 220, (226). Apodes, 6.
bonaiensis, 188. Appendicularian, 337 F.
brevirostris, 205, 206, (216). appendiculatum, Distomum, 197P, 322P, 338P,
carrikeri, 217. 356P, 404P.
cayennensis, 206, (212). arcticus, Mallotus, 594.
cayorum, 174. Osmerus, 573, 593-
choerostoma, 186, 202. Salmo, 573, 592.
choerostoma var. atlantica, 186. arcturus, Salmo, 524.
clupeoides, 158. Salvelinus, 524.
cubana, 190, 224. arcturus (subsp.), Sa/velinus alpinus, 524.
elongata, 205, 207, (223). arcuata, Clupea, 395.
epsetus, 188, 194, 199. Harengula, 395.
estauquae, 205, 206, 208, (211). arenarius, Cynoscion, 343.
eurystole, 204—206, (208), 211-213, 215, 216. Arengus, 397.
Silifera, 204. argentata, Ilisha, 425.
Luianensis, 205, 207, (217), 219. argenteus, Anchoa, 162, 163, 166, (169).
Jamesi, 204, 205, 207, 228. Salmo, 498.
(Amplova) jamesi, (227). Salmo or Fario, 459.
Januaria, 184. Sarchirus, 80.
lepidentostole, 205, 207, 220, (224), 227. Synodus, 133.
longipinna, 204. Argentina, 8, 91, 96, 123, 133, 559, 576.
39
610 Index of Scientific Names
carolina, 123, 131. Tarpon, (113).
glossodonta, 133. Atopichthys esunculus, 142.
menidia, 199. Atractosteus bocourti, 70.
silus, 8. lucius, 86.
argentina, Trutta, 541. Atractosteus (subgenus), 69, 70.
Argentinid, Argentinoid, or Argentine, 12, 90, 549, Aulopidae, 15, 101.
559, 576. Aulostomatophora, 250.
Argentinidae, 8, 99, 100, 104, 554. phosphorops, 251.
Argentininae, 99. aurea, Brevoortia, 344, 34.6, 362, 365.
Argentinoidea, 8, 12, 89, 92, 93, 95, 96, 149. aurea (var.), Brevoortia tyrannus, 361.
argentivittatus, Engraulis (Stolephorus), 202. aureolus, Salvelinus, 506, 509, 524.
Argulus alosae, 338P. aureolus (subsp.), Sa/velinus alpinus, 524, 541, 542.
argyrophana, Anchovia, 210. aureus, Brevoortia, 375.
Anchoviella, 210. aurita, Sardinella, 397, 405-407, 410, 411.
argyrophanus, Engraulis, 210, 215. ausonii, Salmo or Salar, 459.
Stolephorus, 210. ayresii, Lepidosteus, 81.
artedi, Leucichthys, 551.
Arthropod, 481 F.
Ascaris adunca, 303 P. Bacittus salmonicida, 484.D.
sp. 303 P, 322P. salmonis pestis, 484D.
Asquamiceps, 90, 250. baeri, Acipenser, 32.
astilbe, Anchoviella, 190. Acipenser x A. ruthenus, 32.
Stolephorus, 188, 190. bahamensis, Vulpes, 141.
Astronesthes, 95. bahiensis, Engraulis, 133, 141.
myriaster, 95. Rhinosardinia, 411.
niger, 95. Baione, 503.
Astronesthidae, 90, ror. Sontinalis, 503, 541.
Astroscopus, 12. Baione (subgenus), 542.
atchafalayae, Signalosa, 443, 451. bairdi, Acipenser, 56.
atchafalayae, Signalosa, 451. Bairdiella icistius, 450F.
vanhyningi, Signalosa, 451. bairdii, Alepocephalus, 251.
atchafalayae (subsp.), Dorosoma petenense, 451. Salmo, 525.
Dorosoma petenensis, 451. Salvelinus, 525.
Signalosa atchafalayae, 451. balboae, Amplova, 204, 206, 228.
Signalosa petenensis, 451. Anchoviella, 227.
atelaspis, Acipenser, 46. Balistid, 7.
Atherina, 555. Balistidae, 7.
browntii, 198. bananus, Butyrinus, 133.
epsetus, 198. Barbantus, 254.
hepsetus, 555. barbouri, Lycengraulis, 233, 235, 236, 242, (243).
mordax, 555, 556, 569. bartonii, Cylindrosteus, 75.
Atherinid, 139F. batesit, Engraulis, 241.
Atherinidae, 555, 559. Lycengraulis, 233, 235, (240), 243, 244.
atherinoides, Clupea, 228, 230. Bathylaconidae, 94, 102.
Engraulis, 230. Bathylaconoidea, 16, 92, 93, 95-
Pterengraulis, (229), 231. Bathylaconoidei, 93.
Athrodiri, 2. Bathylagid, go.
atlantica (var.), Anchoviella choerostoma, 186. Bathylagidae, 100, 104.
atlanticus, Megalops, 112, 121. Bathymicrops, 98.
Index of Scientific Names 611
Bathypteroidae, 97, 99-
Bathysauridae, 98, roo.
Bathysaurus, 11.
Bathytroctes, 250.
macrolepis, 251.
Bathytyphlops, 98.
Batoid, 4, 6.
Beluga, 593E.
berlandieri, Lepidosteus, 83, 86.
bermudana, Fenkinsia, 274.
bilinearis, Merluccius, 285 E, 286E, 537E.
bimaculata, Megalops, 447.
Binghamichthys, 95.
bishopi, Alosa, 392.
Clupea, 393.
Sardinella, 393-
bison, Lepidosteus, 79.
blackburni, Anchoviella, 204, 205, 207, 220, (226).
bleekeriana, Ilisha, 442.
Medipellona, 442.
Pellona, 427, 438, 439, 442.
bleekerianus, Chirocentrodon, (439).
Blenniidae, 7.
bocourti, Atractosteus, 70.
Bomolochus eminens, 405P, 406P.
teres, 356P, 370P.
bonaiensis, Anchoviella, 188.
Bonasa umbellus, 488.
Boreogadus saida, 482F, 518F.
Lothryophoron, Distomum, 338P.
Brachymystax, 457-459, 519.
Brachyopterygil, rr.
brasiliensis, Clupea, 133, 141, 409, 410.
Sardinella, 398, 399, 405, 406, (407).
brevicaudata, Brevoortia, 343, 345, (363).
brevicaudata (subsp.), Brevoortia tyrannus, 365.
brevicaudata (var.), Brevoortia tyrannus, 361, 365.
brevipes (var.), Salmo salar, 493.
brevirostris, Acipenser, 24, 29, 32-34, (36), 41, 42,
47, 48, 55.
Acipenser, x A. sturio, 54.
Amplova, 217.
Anchovia, 204, 219.
Anchoviella, 205, 206, (216).
Engraulis, 217, 220, 226.
brevirostrum, Acipenser, 40.
brevis, Engraulis, 247.
Brevoortia, 91, 148, 258, 260, 267, 332, (342).
Brevoortia aurea, 344, 346, 362, 365.
aureus, 375.
brevicaudata, 343, 345, (363).
Sunteri, 3435 3445 346, 347, 369, 371, 372, 3745
(376).
menhaden, 342, 361.
patronus, 79¥F, 343-3475 361, 362, (365), 374-
376, 378-380, 425.
pectinata, 344, 345.
smithi, 3435 3445 346, 347, (372), 379-
sp., 380.
tyrannus, 270, 342-345, (346), 363-365, 368—
VES SISO
tyrannus var. aurea, 301.
tyrannus brevicaudata, 365.
tyrannus var. brevicaudata, 361, 365.
tyrannus var. menhaden, 301.
brevoortiae, Lernanthropus, 322P, 356P, 370P.
browni, Stolephorus, 199.
brownti, Anchoa, 187.
Anchovia, 188, 194, 199, 200.
Atherina, 198.
Engraulis, 198, 200, 271.
Stolephorus, 188, 193, 200, 215.
buffalo, Acipenser, 46.
bulbifer, Rhynchobothrium, 128 P.
Butirinus vulpes, 133, 141.
Butyrinus, 133.
bananus, 133.
Cazanus, 303F.
Calichthyid catfishes, 5.
Caligus chelifer, 356P.
rapax, 303P, 338P.
schistonyx, 356P.
callaris, Salmo, 525.
Callinectes sapidus, 67F, 74F, 79¥, 86F.
callolepis, Clupea, 389.
Harengula, 389.
callorhini, Therobromus, 554.
campi (subsp.), Signalosa mexicana, 451.
Signalosa petenensis, 451.
canadensis, Salmo, 541.
Cancer, 536F.
Caranx hippos, 129.
carbonarius, Acipenser, 46.
caribbaea, Ilisha, 439, 442.
caribbaea (subsp.), Harengula pensacolac, 394-396.
caribbaeus, Chirocentrodon, 4.42.
39
612 Index of Scientific Names
caribbea, Medipellona, 442. insoctabilis, 447.
carolina, Argentina, 123, 131. MeXiCanus, 450.
carolinensis, Clupea, 361. petenensis, 450.
carpio, Salmo, 505, 506, 523. signifer, 384.
carrikeri, Anchoviella, 217. thrissa, 384.
Cartilaginous fishes, 2, 4, 5, 7, 8. Chatoéssus cepediana, 447.
castelnaeana, Ilisha, 416 (419). Chatossus ellipticus, 447.
Ilishe (Neosteus), 420. Chauliodontidae, 102, 103.
Neosteus, 420. chelifer, Caligus, 356P.
Pellona, 420, 427. Chimaeroid, 4—8, 91.
castelnaudi, Cylindrosteus, 83. chinensis, Pristigaster, 416.
catervarius, Mallotus, 574, 575, 593- Chirocentridae, 149.
catervarius (subsp.), Madlotus villosus, 575, 583; Chirocentrinae, 98, 257, 262, 439.
594. Chirocentrodon, 262, (438).
cayana, Pristigaster, (428). bleekerianus, (439)-
cayanus, Pristigaster, 427, 429. cartbbaeus, 442.
cayennensis, Acipenser, 54, 56. taeniatus, 438, 442.
Anchoviella, 206, (212). Chirocentroidei, 149.
Stolephorus, 213. Chirocentrus, 8, 149, 439.
cayorum, Anchoa, 162, 163, 166, (173), 181. Chironomid, 39F (larva), 44 F (larva), 52 F (larva),
Anchovia, 174. 481F (larva), 536.
Anchoviella, 174. Chironomidae, 302 F.
cayorum (subsp.), Anchovia choerostoma, 174. Chlorophthalmidae, ror.
cepediana, Chatoéssus, 447. Choanichthyes, ro.
Megalops, 443, 447. choerostoma, Anchoa, 162, 164, 166, (185), 204.
cepedianum, Dorosoma, 443, (444), 448-450. Anchovia, 202.
exile, Dorosoma, 447. cayorum, Anchovia, 174.
cepedianus, Chatoessus, 447. Anchoviella, 186, 202.
cephalus, Mugil, 86F. Engraulis, 186, 202.
Cepola, 3. Stolephorus, 186, 202.
Cestode, 128P, 197P, 285 P (larva), 286P, 322P, Chondrichthyes, 2, 4.
338P, 356P (larva), 362P, 484P, 495 P, Chondrostei, 13.
570P. Chondrosteidae, 26.
Cetengraulis, 148, 152, 154, (245). Chondrosteoidea, 13.
edentulus, 157, 221, (245), 24.8. chrysochloris, Clupea, 312, 319.
armani, 247 Pomolobus, 309, 311-313, (315), 320, 322, 339,
gilberti, 247. 343.
juruensis, 223, 245, (247). Cichlid, 6.
mysticetus, 245. cincinnati, Acipenser, 40.
Cetomimidae, 12. Cladocera, 481F.
Chaetoessus, 92, 149. Cladoceran, 450F, 518F.
Chanidae, 15, 149. Clavellisa spinosa, 356P.
Chanina, 93. clintonii, Lepidosteus, 81.
Chanoidea, 89. Clupanadon pseudohispanicus, 406.
Chanos, 92. Clupanodon, 398.
Characid, 8. pseudohispanicus, 405, 410.
Chatoessus, 443. thrissa, 380.
cepedianus, 447. Clupea, 112, 133, 258, 260, (274), 573-
cumorphus, 385. aestivalis, 331.
Index of Scientific Names 61
africana, 415.
allecia, 410.
alosa, 293, 307.
amazonica, 411, 413.
anchovia, 406.
antipodum, 287.
apicalis, 389.
arcuata, 395.
atherinoides, 228, 230.
bishopi, 393-
brasiliensis, 133, 141, 409, 410.
callolepis, 389.
carolinensis, 361.
chrysochloris, 312, 319.
clupeola, 386, 389, 392.
coerulea, 293.
cyprinoides, 112, 121.
elongata, 293.
Sasciata, 332.
gigantea, 112, 121.
halec, 293.
harengus, 274, (275), 405, 406, 439, 573, 581.
heterura, 447.
holodon, 287.
humeralis, 389, 392, 395-
indigena, 307.
lamprotaenia, 267, 271.
latulus, 386, 392.
lodna, 592.
macrocéphala, 133, 141.
macrophthalma, 389, 39°, 392, 396.
mattowaca, 323.
mattowocca, 323.
mediocris, 323.
megalops, 341.
menhaden, 361.
micropus, 262.
neglecta, 361.
pallasii, 286.
paroula, 312, 342.
pensacolae, 395.
pilchardus, 230.
pseudoharemgus, 312, 331, 341, 342.
pseudohispanica, 407.
pseudohispanicus, 406.
pusilla, 324.
sadina, 267.
Sapidissima, 306, 308, 311.
sardina, 389.
W
sp., 389.
Sprattus, 482F.
thrissa, 380, 384, 385.
tyrannus, 361.
vernalis, 331, 340, 341.
villosa, 573, 592.
VIFeSCENS, 3.42.
vittata, 198.
clupeaformis, Coregonus, (549).
clupeaformis, Coregonus, 550.
gulliveri, Coregonus, 550.
neo-hautoniensis, Coregonus, 550.
stanley, Coregonus, 550.
clupeaformis (subsp.), Coregonus clupeaformis, 550.
Clupeid, or Clupeoid, 7, 8, 94.
Clupeida, 94.
Clupeidae, 14, 15, 98, 110, 133, 148-150, 206,
250, 251, (257).
Clupeiformes, 91, 92, 94.
Clupeina, 93.
Clupeinae, 98, 260.
Clupeoidea, 14, 92-96, (148).
Clupeoidei, 93.
clupeoides, Anchovia, (155), 159, 161, 221, 231.
Anchoviella, 158.
Engraulis, 157, 247.
Stolephorus, 157, 158.
clupeola, Clupea, 386, 389, 392.
Harengula, 387, 389, (390).
Sardinella, 393.
Coelacanth, 2-5, 7-11.
coerulea, Clupea, 293.
Coilia, 90.
Colliparvus parvus, 271 P, 272P.
colonensis (subsp.), Anchoa hepsetus, 162, 165, 166,
197, 200.
commensuralis, Obelia, 330P.
compressus, Engraulis, 162.
Odontognathus, 430, 432, (433).
Concara macroptera, 252.
Conorhynchus, 133.
plumieri, 141.
conorhynchus, Albula, 141.
copei, Acipenser, 46.
Lepidosteus, 81.
Xenodermichthys, 251.
Copepod, 178F, 196F, 271 P, 272P, 283-285F,
302, 303 RH, 322.P5) 32458513298, 33725
338P, 355F, 356P, 362 P, 370P, 371, 374P,
614 Index of Scientific Names
383 F, 404F, 405 P, 406P, 450F, 484 P, 494 P,
518F, 519P, 537P, 564F, 570P, 571 P, 588F.
Coregoni, 549.
Coregonidae, 8, 14, 96, 99, 455, 456, (547).
Coregonus, 547, 548, (549)-
clupeaformis, (549).
clupeaformis clupeaformis, 550.
clupeaformis gulliveri, 550.
clupeaformis neo-hautoniensis, 550.
clupeaformis stanley, 550.
lavaretus, 549.
nasus, 549.
nelsoni, 549.
pidschian, 549.
Cottid, 482 F, 518F.
Crago, 52¥F, 285F, 566B.
Craniata, 2.
crassus, Lepidosteus, 79.
Cristivomer, 455, 457» 458, 503, 504, (542).
namaycush, 458, 460, 505, 507, 509, 526, (543).
Cromeriidae, 149.
Crossopterygidae, 10.
Crossopterygii, 10.
Crustacea(n), 31 F, 39F, 79F, 284F (nauplii,
young), 285F, 322F, 355 FP, 370P, 482F,
518F, s19F, 536F.
cubana, Anchoa, 162, 165, 166, (188).
Anchovia, 199.
Anchoviella, 190, 224.
cubana (subsp.), Harengula pensacolae, 394-396.
cubanus, Engraulis, 190.
Neoopisthopterus, (436).
Stolephorus, 190.
curilus, Salmo, 525.
Curinata, 8.
curta, Anchoa, 181, 183.
cyanonoton, Alosa, 332.
Pomolobus, 331.
Cyclopterus, 518F.
Cyclostome, 2, 4, 9.
Cyclothone, 95.
Cylindrosteus, 69.
agassizii, 75.
bartonii, 75.
castelnaudi, 83.
megalops, 80, 83.
productus, 75.
Cylindrosteus (subgenus), 69.
Cymothoa praegustator, 370P.
Cynoscion, 198E.
arenarius, 343.
regalis, 329E, 331E, 343.
cyprinoides, Clupea, 112, 121.
Megalops, 113, 115, 121, 122.
Dasryvanus, Acipenser, 32.
Dactylocotyle sp., 356P.
Dallia, 2.
dauricus, Huso, 29.
Decapod, 284F, 303F (larva), 518F (larva); 536F
and 564F (shrimp).
dekayi, Acipenser, 41.
Delphinapterus, 588.
dentex, Engraulis, 236.
Osmerus, 556, 557, 57° 571-
dentex (subsp.), Osmerus eperlanus, 557, 572.
dentex natio dvinensis morpha spirinchus, Osmerus
eperlanus, 557, 569, 572.
de sotoi (subsp.), Acipenser oxyrhynchus, 57.
desotoi (subsp.), Acipenser oxyrhychis, 24, 36, 47, 54,
(56).
desuratus, Ilisha, 419.
diaphana (subsp.), Anchoa mitchilli, 162, 164, 166,
174, 178, 18r.
diaphanus, Fundulus, 44F.
Diatom, 284¥, 285F, 337F, 355F, 374F, 450F.
Dinectus truncatus, 46.
Diodontid, 7.
Diplobatis, 6.
Dipneusti, 9.
Dipnoa, 3.
Dipnoi, 2-5, 7-9, 11-13.
Diptera, 411.
Dipteran (flies), 481 F.
Distome, 303 P.
Distomum appendiculatum, 197P, 322P, 338P,
356P, 404P.
Lothryophoron, 338P.
pyriforme, 356P.
sp., 197P.
vitellosum, 338P, 356P.
ditchela, Pellona, 416.
Dixonina, 107, 108, 133, 134, 142, (143).
nemoptera, 142, (143).
pacifica, 144, 145.
Dolichopteryx, 91, 95, 96.
dolloi, Pristigaster, 430.
Index of Scientific Names 615
Dorosoma, 148, 257, 262, 398, (443).
cepedianum, 443, (444), 448-450.
cepedianum exile, 447.
exile, 447.
mexicanum, 450.
notata, 443, 447.
petenense, 443, 444, (448).
petenense atchafalayae, 451.
petenensis atchafalayae, 451.
smithi, 443.
Dorosomatinae, 98, 149, 257.
duodecim, Anchoa, 162, 163, 166, (170).
Anchovia, 17%.
Engraulis, 171, 180.
Dussumieria stolifera, 267, 271.
Dussumierinae, 98, 149, 257, 259.
dvinensis, Osmerus, 556, 572.
dvinensis (subsp.), Osmerus eperlanus, 572.
dvinensis (var.), Osmerus eperlanus, 572.
dvinensis (natio) morpha spirinchus, Osmerus eperlanus
dentex, 557, 569, 572.
Esena, Fusconaia, 318P.
Echinorhynchus acus, 303P, 330P, 338P.
edentulus, Cetengraulis, 157, 221, (245), 248.
Engraulis, 221, 223, 243, 245-247.
edwardsi, Lepeophtheirus, 338P.
Eimeria, 356P.
Elasmobranch, 4-8, 91, 554.
elisabeth, Lepidosteus, 81.
ellipticus, Chatossus, 447.
Ellops, 28.
elongata, Alosa, 415, 416.
Anchovia, 224.
Anchoviella, 205, 207, (223).
Clupea, 293.
elongatus, Megalops, 121.
Elopid, or Elopoid, 93, 94.
Elopidae, 12, 14, 15, 93, 97, 107, 108, 110, (111),
032.
Elopina, 93, 94.
Elopoidae, 94.
Elopoidea, 14, 92, 94, 95, (107).
Elopoidei, 93.
Elops, 93, 94, 107, 108, III, 12, 115, 122, (123).
affinis, 127, 128, 130.
inermis, 131.
Saurus, 115, 123, (124), 142.
senegalensis, 128, 130.
eminens, Bomolochus, 405P, 406P.
Engraulidae, 14, 90, 97, 148-150, (152), 274, 439.
Engraulis, 133, 152, 162.
(Stolephorus) argentivittatus, 202.
argyrophanus, 210, 215.
atherinoides, 230.
bahiensis, 133, 141.
batesii, 241.
brevirostris, 217, 220, 226.
brevis, 247.
brownti, 198, 200, 271.
choerostoma, 186, 202.
clupeoides, 157, 247.
compressus, 162.
cubanus, 190.
dentex, 236.
duodecim, 171, 180.
edentulus, 221, 223, 243, 245-247.
eurystole, 210.
grossidens, 233, 236.
hepsetus, 200.
hiulcus, 199.
iquitensis, 220.
janeiro, 236.
Januarius, 184, 226.
louisiana, 180.
macrolepidotus, 155.
mitchilli, 180, 184.
nattereri, 221.
perfasciatus, 204, 210, 215.
piquitinga, 188.
platyargyreus, 202.
productus, 157, 247.
Sericus, 133, 141.
spinifer, 168.
surinamensis, 157, 221.
tricolor, 188, 199.
vittatus, 199.
eperlano-marinus, Salmo, 569.
Eperlanus, 555.
vulgaris, 556.
eperlanus, Osmerus, 556-558, (559), 576, 581.
dentex, Osmerus, 557, 572-
dentex natio dvinensis morpha spirinchus, Osmerus,
557 569, 572.
dvinensis, Osmerus, 572.
dvinensis (var.) , Osmerus, 572.
616 Index of Scientific Names
eperlanus, Osmerus, 557-559, 565, 567-
eperlanus, Salmo (Osmerus), 569.
eperlanus natio ladogensis, Osmerus, 557.
mordax, Osmerus, 557, 558, (559).
Salmo, 554-556, 569, 573.
Salmo (Osmerus), 555, 569.
spirinchus (morpha), Osmerus, 557.
eperlanus (subsp.), Osmerus eperlanus, 557-559, 565,
567.
ladogensis (natio), Osmerus eperlanus, 557.
Salmo (Osmerus) spirinchus, 569.
Ephemeridae, 536F.
epsetus, Anchoviella, 188, 194, 199.
Atherina, 198.
erythrorynchos, Salmo, 525.
Esocidae, 98
Esociformes, 92, 93.
Esocina, 93.
Esocoidea, 7, 90, 92, 93, 95, 107.
Esocoidei, 93.
Esox, 69, 133, 142.
hepsetus, 198.
lucius, 5.
asseus, 69, 79-
virdis, 79.
vulpes, 133, 141, 142.
estauguae, Anchoviella, 205, 206, 208, (211).
esunculus, Atopichthys, 142.
Ethmidium, 148.
Etrumeus, 258, 259, (262), 267, 268.
Jacksoniensis, 266.
microps, 266.
sadina, (263).
teres, 267.
Euacipenser, 29.
Eubothrium, 519P.
Eumicrotremus spinosus, 518¥F.
eumorphus, Chatoessus, 385.
Euphausiid, 283 F (shrimp); 284F; 303 F (shrimp);
482F; 518F and 588F (shrimp).
Eurypharynx, 2.
eurystole, Anchovia, 210.
Anchoviella, 204-206, (208), 211-213, 215,
216.
Engraulis, 210.
Stolephorus, 210.
Eustomias, 90.
Euthemisto, 285 P.
Evermannellidae, 100, 104.
exile, Dorosoma, 447.
exile (subsp.), Dorosoma cepedianum, 447.
Fanio, 459.
fario, Salmo, 541.
Salmo or Salar, 459.
fasciata, Clupea, 332.
felis, Galeichthys, 79¥.
ferox, Lepisosteus, 69, 86.
Silamentosus, Megalops, 121.
jilifera, Anchoa, 162, 165, 166, (202).
Anchovia, 204.
Anchoviella, 204.
Jinta, Alosa, 304.
Flagellate, 284F.
flavipinnis, Ilisha, 417, 420.
Pellona, 420.
Pristigaster, 416, 427.
fioridana (subsp.), Harengula pensacolae, 394-396.
fontinalis, Baione, 503, 541.
agassizit, Salvelinus, 542.
hudsonicus, Salvelinus, 537, 542.
Salmo, 541.
Salvelinus, 460, 483E, 484, 498, 499, 504-507,
509, 510, 515, 518, 522, 523, (525), 544.
Salvelinus (Baione), 504, 542.
Foraminifera, 197F.
forskali, Glossodus, 141.
foveolata, Heterakis, 330P.
Fucellaria, 588P.
fulvescens, Acipenser, 24, 28-34, 36, 38, 40, (41),
47, 48, 53, 54.
Fundulus, 536F.
diaphanus, 44F.
Surthit, Ilisha, 421.
Furunculosis, 484 D.
Fusconaia ebena, 318P.
Ganus morrhua, 519F, 588E.
ogac, 588E.
gairdneri, Salmo, 8, 459, 460, 491, (499), 525.
Galaxiidae, gt.
Galaxiiformes, 92.
Galeichthys felis, 79F.
Gammaracanthus, 518F.
Gammarid, 482F, 518F, 536F, 588F.
Index of Scientific Names 617
Gammarus, 285F, 518F, 536F. gulliveri (subsp.), Coregonus clupeaformis, 550.
locusta, 518F. gunnellus, Pholis, 581.
Ganoid, 14, 62, 64, 132. gunteri, Brevoortia, 343, 344, 346, 347, 369, 371,
Ganoidea, 14. 372, 374, (376).
Ganoidei, 3, 10, 13, 14. Gymnarchidae, 92.
garmani, Cetengraulis, 247. Gymnophotodermi, 93, 96.
Stolephorus, 247. Gymnophotodermoidea, 93.
Gasteropelecus, 428.
Gasterosteus, 536F.
Gastropod, 44F, 52F, 178F, 197F, 284F (larva).
gavialis, Lepisosteus, 69, 79. Huzec, Clupea, 293.
gigantea, Clupea, 112, 121. Halecostomi, 93.
giganteus, Megalops, 112, 121. Halosaurid, 13.
Tarpon, 121. hamatus, Salmo, 497.
Gigantura, 11. Haplochitontidae, gr.
Giganturidae, 103. Haplomi, 92, 93.
Giganturoid, 5-7, 16. hardinii, Salmo, 492, 498.
Giganturoidel, 14, 16, 97. Harengula, 148, 257, 258, 261, (386).
gilberti, Anchovia, 247. arcuata, 395.
Cetengraulis, 247. callolepis, 389.
Stolephorus, 247. clupeola, 387, 389, (390°).
Ginglymodi, 64. humeralis, 386, (387), 392, 395-
ginsburgi, Anchoa, 162, 165, 166, (190), 194. Jaguana, 389.
girardi, Acipenser, 56. latulus, 386, 392.
glabra, Acipenser, 28. macrophthalma, 389, 39°, 392, 395, 396.
gladius, Psephurus, 26. maculosa, 389.
Gladostomus, 29. majorina, 396.
glossodonta, Argentina, 133. pensacolae, 387, 392, (393)-
Glossodus, 133. pensacolae caribbaea, 394-396.
Sorskali, 141. pensacolae cubana, 394-396.
gloveri, Salmo, 498. pensacolae floridana, 394-396.
Guathobolus, 430. pensacolae majorina, 394-396.
mucronatus, 433. pensacolae pensacolae, 394, 396.
Gonorhynchina, 93. pensacolae pinensis, 394-396.
Gonorhynchus, 90, 91. peruana, 387.
Gonostoma, 95. sardina, 389, 392, 396.
Gonostomatidae, 103. Sp.» 390.
gorbuscha, Oncorhynchus, 458, 484. species dubia, 392.
gracilis, Lepidosteus, 79. thrissina, 387.
Salmo, 498. harengus, Clupea, 274, (275), 405, 406, 439, 573,
groenlandicus, Mallotus, 593. 581.
Salmo, 573, 592. harlani, Lepidosteus, 81.
grossidens, Engraulis, 233, 236. harroweri, [lisha, 417, (423), 428.
Lycengraulis, 233, (234), 237, 244. Neosteus, 425.
guianensis, Amplova, 219. hautin, Tripteronotus, 549.
Anchoviella, 205, 207, (217), 219. Helops, 28.
Stolephorus, 217, 219, 220. helops, Acipenser, 28.
giildenstddti, Acipenser, 29, 32, 58. Hemiptera, 118F.
Acipenser, x A. ruthenus, 32. Hemitripterus, 536F.
40
618 Index of Scientific Names
hepsetus, Anchoa, 165, 187, 189, 190, 193, (194),
209; 2055 271.
colonensis, Anchoa, 162, 165, 166, 197, 200.
hepsetus, Anchoa, 162, 165, 166, 200.
Atherina, 555.
Engraulis, 200.
Esox, 198.
hepsetus (subsp.), Anchoa hepsetus, 162, 165, 166,
200.
Heringia, 411, 415.
amazonica, 413, 415.
Heterakis foveolata, 330P.
Heterophotodermi, 93.
heterura, Clupea, 447.
Hexagenia, 44¥, 52¥F.
Hildebrandichthys, 154, (230).
setiger, 230, (231).
Hippocampus, 9.
hippoglossoides, Reinhardtius, 588E.
hippos, Caranx, 129.
hiulcus, Engraulis, 199.
holbrooki, Acipenser, 56.
holodon, Clupea, 287.
Holostean, 93, 94.
Holostei, 13, 64.
Holtbyrnia, 254.
horatii, Lepidosteus, 81.
howelli, Anchoa, 203.
Hucho, 457, 458, 519.
hudsonicus, Salmo, 537, 541.
hudsonicus (subsp.), Sa/velinus fontinalis, 537, 542.
humeralis, Clupea, 389, 392, 395-
Harengula, 386, (387), 392, 395.
Sardinella, 390, 395.
huronensis, Lepisosteus, 79.
Huso, 27-29, 32.
dauricus, 29.
huso, 29, 31.
huso, Acipenser, 28.
Huso, 29, 31.
Hyodontidae, gr.
Hypomesus, 554.
Texuocorna, 28
Ichthyococcus, 103.
Ichthyosporidium, 286P.
icistius, Bairdiella, 450F.
Idiacanthidae, 90, 102, 104.
Idiacanthus, 90.
Ilisha, 110, 261, (415), 439.
abnormalis, 415.
altamazonica, 416, (417), 420, 421.
(Neosteus) altamazonica, 419.
amazonica, 417, (421).
apapae, 421-423.
argentata, 425.
bleekeriana, 442.
caribbaea, 439, 442.
castelnaeana, 416, (419).
desuratus, 419.
flavipinnis, 417, 420.
Surthii, 421.
harroweri, 417, (423), 428.
iquitensis, 422, 423.
narragansetae, 416, 417, (426).
Llishe (Neosteus) castelnaeana, 420.
(Neosteus) ternetzi, 426.
immaculata, Amia, 133, 141.
immaculatus, Salmo, 497, 523, 541.
imparispine, Rhynchobothrium, 338P.
indigena, Clupea, 307.
inermis, Elops, 131.
Iniomi, 11, 14-17, 89, 92, 96, 97.
insociabilis, Chatoessus, 447.
Ipnopidae, 98, ror.
iguitensis, Engraulis, 220.
Tlisha, 422, 423.
Trillion, 547, 548.
Isopod, 52F, 178F, 303F, 355P, 361P, 362P,
370P, 371P, 374P, 536F, 588F.
Isospondyli, 8, 14-17, 20, (89).
Istiophoridae, 12.
Istiophorus, 6.
i pooeeeee Etrumeus, 266.
jaguana, Harengula, 389.
jamesi, Amplova, 228.
Anchoviella, 204, 205, 207, 228.
Anchoviella (Amplova), (227).
janeiro, Engraulis, 236.
januaria, Anchoa, 162, 164, 166, 174, 178, 181,
(183).
Anchovia, 184.
Anchoviella, 184.
januarius, Engraulis, 184, 226.
Fenkinsia, 259, (267).
Index of Scientific Names
bermudana, 274.
lamprotaénia, 131, 215, (268), 273, 274.
stolifera, 271, 272, 274.
viridis, 268, 271, (272).
iuruensis, Cetengraulis, 223, 245, (247).
Kewnicortt, Acipenser, 56.
kikuchii, Acipenser, 32.
kirtlandi, Acipenser, 46.
kisutch, Oncorhynchus, 458.
Kneridae, 149.
koefoedi, Searsia, 255.
Lacusrais, Salmo, 493, 503.
lacustris (subsp.), Pomolobus pseudoharengus, 342.
ladogensis (natio), Osmerus eperlanus eperlanus, 557.
laevigatus, Salmo, 525.
laevis, Acipenser, 46.
lamarit, Acipenser, 46.
Lepidosteus, 81.
lamprotaenia, Anchoa, 162, 164, 166, (192), 194,
199, 200.
Clupea, 267, 271.
Fenkinsia, 131, 215, (268), 273, 274.
Spratelloides, 272.
Latimeria, 2-4, 7, 10.
latulus, Clupea, 386, 392.
Harengula, 386, 392.
lavaretus, Coregonus, 549.
Salmo, 549.
lecontei, Acipenser, 56.
Lepeophtheirus, 519P, 537P.
edwardsi, 338P.
salmonis, 484P, 494P, 496P, 497 P.
lepidentostole, Anchovia, 226.
Anchoviella, 205, 207, 220, (224), 227.
Lepidophotodermi, 93, 96.
Lepidosteiformes, 64.
Lepidosteini, 64.
Lepidosteoidei, 64.
Lepidosteus, 65, 69, 93.
ayresii, 81.
Lerlandieri, 83, 86
bison, 79-
clintonti, 81.
copei, 81.
Crassus, 79.
elisabeth, 81.
gracilis, 79.
harlani, 81.
horatit, 81.
lamarii, 81.
leptorhynchus, 75, 79.
lesueurit, 81.
louisianensis, 81.
milberti, 81.
osseus, 81.
otarius, 79.
piquotianus, 81.
rostratus, 79.
Semiradiatus, 79.
smitht, 81.
thompsont, 81.
trecult, 81.
troostit, 81.
Lepisostei, 14, 16 (61).
Lepisosteid, or Lepisosteoid, 6, 7, 13, 65.
Lepisosteida, 64.
Lepisosteidae, 61, (68).
Lepisosteoidea, 13.
Lepisosteus, 15, 61, 65, 68, (69), 93.
Sferox, 69, 86.
gavialis, 69, 79.
huronensis, 79.
lineatus, 79.
longirostris, 79.
oculatus, 61, 67, 70, (71), 75, 76, 81-83.
osseus, 15, 61, 66, 69-74, (75), 81, 83.
oxyurus, 79.
platostomus, 69, 71, 83.
platyrhincus, 61, 67, 70, 71, 75, (81), 83.
sinensis, 67.
619
spatula, 61-63, 67, 70, 71, 75, 76, 80, 81, (83).
stenorhynchus, 81.
tristeochus, 70.
tropicus, 67, 70.
Lepisosteus (subgenus), 69, 70.
Lepogenys, go.
Leptochilichthys, 250.
Leptoderma, 250.
macrops, 255.
Leptolepidae, go, 91.
leptorhynchus, Lepidosteus, 75, 79-
Lepus americanus, 488.
Lernaeenicus radians, 374P.
radiatus, 303P, 356P.
40
620 Index of Scientific Names
Lernanthropus brevoortiae, 322P, 356P, 370P.
lesueurit, Acipenser (Huso), 41.
Lepidosteus, 81.
Leucichthys, 547, 548.
artedi, 551.
Leucichthys (subgenus), 548.
limnichthys, Lycengraulis, 233, (237).
Limnodrilus, 52F.
lineata, Alosa, 323.
lineatus, Lepisosteus, 79.
Lioniscus, 28, 29.
liopeltis, Acipenser, 46.
Liparid, 518F.
Litholepis, 69.
adamantinus, 69.
Litholepis (subgenus), 69.
locusta, Gammarus, 518F.
lodna, Clupea, 592.
longipinna, Anchoviella, 204.
longirostris, Lepisosteus, 79.
Lophius, 2, 35 7 9:
Lophobranch, 5, 7.
Lophobranchii, 3.
Lophotid, 13.
lordit, Salmo, 525.
Loricariid, 5.
loricatus, Macrognathus, 79.
louisiana, Engraulis, 180.
louisianensis, Lepidosteus, 81.
lucius, Atractosteus, 86.
Lumpenus, 518F.
Lycengraulis, 152, 154, (233).
abbotti, 233, 235, (242).
barbouri, 233, 235, 236, 242, (243).
batesit, 233, 235, (240), 243, 244.
grossidens, 233, (234), 237, 244.
limnichthys, 233, (237).
olidus, 234-237, 241.
poeyi, 235.
schroederi, 241, 243.
lyolepis, Anchoa, 162, 165, 166, 186, (200), 202, 204.
Anchoviella, 202.
Stolephorus, 202.
Lyomeri, 5, 12, 14-17, 20.
Lyomerid, 4, 5, 12, 16.
Macaisrnpag, 97, 149.
macrocephala, Clupea, 133, 141.
Macrognathus loricatus, 79.
macrohinus, Acipenser, 56.
macrolepidota, Anchovia, 158.
macrolepidotus, Engraulis, 155.
macrolepis, Bathytroctes, 251.
macrophthalma, Clupea, 389, 390, 392, 396.
Harengula, 389, 39°, 392; 395, 396.
Sardinella, 390.
macrophthalmus, Sardinella, 390, 393, 396.
Macropinna, 8, 12, 96.
macrops, Leptoderma, 255.
macroptera, Concara, 252.
macrostomus, Acipenser, 46.
Macrozoarces americanus, 588E.
maculosa, Harengula, 389.
Sardinella, 390.
maculosus, Acipenser, 46.
majorina, Harengula, 396.
majorina (subsp.), Harengula pensacolae, 394-396.
Malacopterygiens abdominaux, gI.
Malacopterygii, 91, 92.
Malacosteidae, 89, 102, 103.
Matlacosteus, 95.
Mallotus, 275, 553, 554, (573):
arcticus, 594.
catervarius, 574, 575, 593-
groenlandicus, 593.
socialis, $74, 594.
villosus, 559, 574s 575, (57).
villosus catervarius, $75, 583, 594.
villosus villosus, 575, (576).
malma, Salmo, 506, 523, 525.
Salvelinus, 519, 520, 525.
malma (subsp.), Salmo, 523.
Salvelinus alpinus, 484, 506, 519, 520.
manjuba, Stolephorus, 247.
marina, Trutta, 541.
Zostera, 538.
marstoni, Salmo (Salvelinus), 506, 524.
Salvelinus, 506, 509.
marstoni, Salmo (Salvelinus), 506, 524.
marstoni (subsp.), Sa/velinus alpinus, 524.
martit, Pristigaster, 428-430.
mattowaca, Clupea, 323.
mattowacca, Alosa, 323.
mattowocca, Clupea, 323.
Maulisia, 254.
mediocris, Clupea, 323.
Pomolobus, 20-23, 312-315, (319), 33% 342, 343-
Index of Scientific Names 621
Medipellona, 438, 439.
bleekeriana, 442.
caribbea, 442.
medirostris, Acipenser, 32, 33, 36.
Megaceryle alcyon, 483E.
megalaspis, Acipenser, 46.
Megalopidae, 107, 108, 111.
Megalops, 93, 94, 107, 108, 112, 113, 115, 122,
130.
atlanticus, 112, 121.
bimaculata, 447.
cepediana, 443, 447.
cyprinoides, 113, 115, 121, 122.
elongatus, 121.
Sfilamentosus, 121.
giganteus, 112, 121.
notata, 384.
oglina, 384.
thrissoides, 121.
megalops, Clupea, 341.
Cylindrosteus, 80, 83.
Meganyctiphanes norwegica, 284F.
Melanostomiatidae, 102.
Meletta petenensis, 450.
suoerit, 319.
thrissa, 384.
VENOSa, 341.
menhaden, Alosa, 361.
Brevoortia, 342, 361.
Clupea, 361.
menhaden (var.), Brevoortia tyrannus, 36%.
Menidia, 198.
Menidia, 128F, 536F, 576, 591.
menidia, Argentina, 199.
Menticirrhus, 343.
Mentodus, 254.
Mergus merganser, 483E.
Merluccius bilinearis, 285 E, 286E, 537E.
Mesichthyes, 92.
mexicana, Signalosa, 451.
campi, Signalosa, 451.
mexicana, Signalosa, 451.
mexicana (subsp.), Signalosa mexicana, 451.
mexicanum, Dorosoma, 450.
mexicanus, Chatoessus, 450.
miarchus, Stolephorus, 215.
microcephalus, Somniosus, 588E.
microdon, Osmerus, 573, 594.
microps, Etrumeus, 266.
micropus, Clupea, 262.
microrhynchus, Acipenser, 41.
Microstomatinae, 99, 100, 104.
milberti, Acipenser, 56.
Lepidosteus, 81.
Miripinnati, 7.
Mirognathus, 250.
Mirorictinae, 254.
Mirorictus, 254.
mirus, Platytroctegen, 255.
mitchilli, Acipenser, 56.
Anchoa, 79F, 163, 170, 173, (176), 181-184,
198, 226.
Anchovia, 180, 181.
Anchoviella, 175, 180, 181, 183, 184.
Engraulis, 180, 184.
Stolephorus, 174, 180, 181.
diaphana, Anchoa, 162, 164, 166, 174, 178, 181.
mitchilli, Anchoa, 162-164, 166, 178, 181.
mitchilli (subsp.), Anchoa mitchilli, 162-164, 166,
178, 181.
Mollusk, 31 F, 44F, 52F, 139F, 303F.
Monognathidae, 12.
Monostomum sp., 338P.
mordax, Atherina, 555, 556, 569.
Osmerus, 556, 557, 570.
abbotti, Osmerus, 556, 572.
abbotti (var.), Osmerus, 572.
spectrum, Osmerus, 572.
spectrum (var.), Osmerus, 572.
mordax (subsp.), Osmerus eperlanus, 557, 558, (559)-
Salmo (Osmerus) spirinchus, 569.
Mormyridae, 92.
Mormyriformes, 92.
morrhua, Gadus, 519F, 588E.
mucronatus, Gnathobolus, 433.
Odontognathus, 430, (431), 433, 435-
Pristigaster, 433.
Mugil, 86F.
cephalus, 86F.
Mugilomorus, 123.
anna-carolina, 123, 131.
multiscutatus, Acipenser, 32.
Muraenidae, 6.
Myctophidae, 15, 103, 104.
Myoxocephalus, 588E.
myriaster, Astronesthes, 95.
Mysid (shrimp), 303 F, 337 F; 518F and 564F.
Mysis, 178F, 285 F, 302F.
622 Index of Scientific Names
mysticetus, Cetengraulis, 245.
Myxosporidian (protozoan), 286P.
N ACARI, Acipenser, 32.
namaycush, Cristivomer, 458, 460, 505, 507, 509,
526, (543).
Salmo, 503, 504, 542, 544.
Salvelinus, 544.
Nansenia, 100.
naresit, Salmo, 524.
Salvelinus, 524.
naresii (subsp.), Salvelinus alpinus, 523.
Salvelinus oguassa, 524.
narragansetae, Ilisha, 416, 417, (426).
Neosteus, 427.
nasus, Coregonus, 549.
nattereri, Anchovia, 221.
Anchoviella, 207, (220).
Engraulis, 221.
neglecta, Clupea, 361.
nelsoni, Coregonus, 549.
Nematalosa, 92, 149.
Nematode, 197 P, 286P, 303 P, 322P, 330P, 362P,
484.P, 571P.
nemoptera, Albula, 145.
Dixonina, 142, (143).
Neoceratodus, 10.
neo-hautoniensts (subsp.), Coregonus clupeaformis, 550.
Neonesthes, 95.
Neoopisthopterus, 262, (435)-
cubanus, (436).
tropicus, 436, 438.
Neopterygii, rr.
Neoscopelidae, 100, 103.
Neosteus, 415, 416, 439.
altamazonica, 419.
castelnaeana, 420.
harroweri, 425.
narragansetae, 427.
ternetzi, 425, 426.
Nereid (worm), 518F.
Nereis, 536F, 566B.
virens, 52¥F.
nertinianus, Acipenser, 46.
niger, Astronesthes, 95.
nigra, Anchovia, 155, 156, (158).
nigrescens, Salmo, 541.
nitidus, Salmo, 524.
nobilis, Salmo, 497.
nobilis (var.), Salmo salar, 493.
Normichthys, 254.
norwegica, Meganyctiphanes, 284F.
notata, Dorosoma, 443, 447.
Megalops, 384.
Notopteridae, go, 91.
nudiventris, Acipenser, 29, 30, 32.
nummifer, Salmo, 525.
Osetia commensuralis, 330P.
oculatus, Lepisosteus, 61, 67, 70, (71), 75, 76, 81— 83.
Odontognathus, 261, (430), 435, 436.
compressus, 430, 432, (433).
mucronatus, 430, (431), 433, 435-
panamensis, 432, 433.
tropicus, 432, 435.
ogac, Gadus, 588E.
Ogcocephalidae, 7.
oglina, Megalops, 384.
oglinum, Opisthonema, 380, (381), 444, 448.
ohiensis, Acipenser, 46.
Alosa, 308, 310, 312.
ohiensis (subsp.), Alosa alabamae, 312.
Olencira praegustator, 355 P, 361 P, 370P, 374P.
olidus, Lycengraulis, 234-237, 241.
Oligochaete (worm), 31F, 52F.
Omosudidae, 100.
Oncorhynchus, 457, 458, 460.
gorbuscha, 458, 484.
hkisutch, 458.
tshawytscha, 484.
opercularis, Stolephorus, 247.
Ophiocephalidae, 12.
Opisthomi, 6.
Opisthonema, 258, 260, (380), 398, 443.
oglinum, 380, (381), 444, 448.
thrissa, 380, 385.
Opisthoproctidae, 104.
Opisthoproctina, 93.
Opisthoproctoidea, 93.
Opisthoproctoidei, 93.
Opisthoproctus, 91, 96.
Opisthopterus, 4.36.
Opsanus, 9.
oguassa, Salmo, 506, 524.
Salvelinus, 522, 524.
naresii, Salvelinus, 524.
Index of Scientific Names 623
oguassa (subsp.), Salvelinus alpinus, 523, 542.
orbignyana, Pellona, 415.
Osmerid, 549.
Osmeridae, 14, 91, 101, 455, 456, (553).
Osmerus, 275, 536F, 553, (554)s 573 574, 577,
578.
abbotti, 556, 557, 572.
articus, 573, 593-
dentex, 556, 557, 572% 571.
dvinensis, 556, 572.
eperlanus, 556-558, (559), 576, 581.
eperlanus dentex, 557, 572.
eperlanus dentex natio dvinensis morpha spirinchus,
557» 569, 572.
eperlanus dvinensis, 572.
eperlanus var. dvinemsis, 572.
eperlanus eperlanus, 557-559, 565, 567.
eperlanus eperlanus natio ladogensis, 557.
eperlanus mordax, 557, 558, (559).
eperlanus morpha spirinchus, 557.
microdon, 573, 594.
mordax, 556, 557, 570.
mordax abbotti, 556, 572.
mordax vat. abbotti, 572.
mordax spectrum, 572.
mordax var. spectrum, 572.
sergeanti, 556, 564, 570-572.
spectrum, 556, 557, 572.
spirinchus, 556, 570, 571.
viridescems, 556, 569, 571.
osseus, Esox, 69, 79.
Lepidosteus, 81.
Lepisosteus, 15, 61, 66, 69-74, (75), 81, 83.
Ostariophysi, 91.
Ostariophysid, 6.
Osteichthyes, (2).
Osteoglossid, gr.
Ostraciodontid, 7.
Ostracod, 197F, 302F, 302F (larva), 303F,
337F.
otarius, Lepidosteus, 79.
othonops, Perkinsia, 262.
ouananiche, Salmo, 492, 498.
Oxyeleotris, 3.
oxyrhynchus, Acipenser, 15, 25, 27, 29-34, 36-45,
(46).
de sotoi, Acipenser, 57.
desotoi, Acipenser, 24, 36, 47, 54, (56).
oxyrhynchus, Acipenser, 24, 36, (46).
oxyrhynchus (subsp.), Acipenser oxyrhynchus, 24, 36,
(46).
Acipenser sturio, 56.
oxyurus, Lepisosteus, 79.
Pactrica, Dixonina, 144, 145.
Palaeoniscoid, 4.
Palaeopterygii, 11.
pallasti, Clupea, 286.
pallida, Anchovia, 221, 223.
Anchoviella, 205, 207, (221).
panamensis, Odontognathus, 432, 433-
Pandarus sinuatus, 356P.
Pantodon, 90.
Paralepididae, 99.
paranasimos, Acipenser, 46.
Parascaphirhynchus, 27.
parkei, Salmo, 525.
parrae, Albula, 141.
parva, Anchoa, 162, 164, 166, 174, 176, 178, 180,
(181), 183, 184, 226, 227.
Anchovia, 182, 223.
Anchoviella, 183.
parvula, Clupea, 312, 342.
parvus, Colliparvus, 271 P, 272P.
patronus, Brevoortia, 79¥, 343-347, 361, 362,
(365), 374-376, 378-380, 425.
pectinata, Brevoortia, 344, 345.
pectoralis, Anchoa, 162, 163, 166, (174), 183, 184.
Pellisolus, 254.
Pellona, 415.
altamazonica, 419.
bleekeriana, 427, 438, 439, 442.
castel/naeana, 420, 427.
ditchela, 416.
flavipinnis, 420.
orbignyana, 415.
Penaeid (shrimp), 26.
pensacolae, Clupea, 395.
Harengula, 387, 392, (393):
caribbaea, Harengula, 394-396.
cubana, Harengula, 394-396.
SJloridana, Harengula, 394-396.
majorina, Harengula, 394-396.
pensacolae, Harengula, 394, 396.
pinensis, Harengula, 394-396.
pensacolae (subsp.), Harengula pensacolae, 394, 396.
Percesoces, 8.
624 Index of Scientific Names
Percoid, 555.
perfasciata, Anchovia, 210, 215.
Anchoviella, 205, 206, 209-212, (213).
perfasciatus, Engraulis, 204, 210, 215.
Stolephorus, 210, 215, 216.
Peridinian, 355 F.
Peridonidian, 285 F.
Perkinsia, 262.
othonops, 262.
Persparsia, 254.
perthecata, Stolephorus, 199.
peruana, Harengula, 387.
pestis (subsp.), Bacillus salmonis, 484 D.
petenense, Dorosoma, 443, 444, (448).
atchafalayae, Dorosoma, 451.
petenensis, Chatoessus, 450.
Meletta, 450.
Signalosa, 451.
atchafalayae, Dorosoma, 451.
atchafalayae, Signalosa, 451.
campi, Signalosa, 451.
vanhyningi, Signalosa, 451.
phaeton, Pristigaster, 429.
Phallostethid, 8.
Philomena, 519P.
Phocaena, 588E, 593E.
Pholidophoridae, go, gr.
Pholis, 275, 536¥F.
gunnellus, 581.
phosphorops, Aulostomatophora, 251.
Photonectes, 90.
Photostomias, 90.
Phractolaemidae, 89, 149.
pidschian, Coregonus, 549.
pilchardus, Clupea, 230.
pinensis (subsp.), Harengula pensacolae, 394-399.
pinnula, Sardinella, 398, (399).
piquitinga, Engraulis, 188.
piquotianus, Lepidosteus, 81.
Pisces, (2).
Pisidium, 52¥.
Placoderm, 2, 4.
Placodermi, 2.
platostomus, Lepisosteus, 69, 71, 83.
platyargyrea, Anchovia, 202, 204.
Anchoviella, 202.
platyargyreus, Engraulis, 202.
Platygaster, 415.
Platygobius, 3.
platyrhincus, Lepisosteus, 61, 67,70, 71,75, (81), 83.
platyrhinus, Acipenser, 46.
Platytroctegen, 254.
mirus, 255.
Platytroctes, 254.
Platytroctinae, 254.
Plecoglossidae, 456.
Plecoglossus, 6, 4.56.
Plecoptera, 481 F.
Plectognath, 7.
plumieri, Albula, 141.
Conorhynchus, 141.
pluvius, Salmo, 525.
Pneumatophorus, 7.
poeyi, Lycengraulis, 235.
Pollachius virens, 536E.
Polychaete (worm), 52F.
polymorphus, Scolex, 197P, 322P, 356P.
Polynemidae, 7.
Polyodon, 30.
spathula, 26.
Polyodontid, 5, 6, 8, 11-13.
Polyodontidae, 26.
Polypteridae, 64.
Polypterini, 64.
Polypteroid, 2, 3, 6-8, 10-14, 62, gr.
Pomatomus, 355E.
Pomocentrid, 6.
Pomolobus, 258, 260, 267, 275, 293, (312), 361,
482F, 536F.
aestivalis, 294, 312-314, 317, (324), 332— 3345
338, 340-342, 362.
chrysochloris, 309, 311-313, (315), 320, 322,
33° 343.
cyanonoton, 331.
mediocris, 20-23, 312-315, (319), 330 342, 343.
pseudoharengus, 312-314, 317, 328, 330, 331,
(332).
pseudoharengus lacustris, 342.
SP+» 329, 3325 342-
Potomabranchii, 3.
praegustator, Cymothoa, 370P.
Olencira, 355P, 361 P, 370P, 374P.
preastabilis, Alosa, 307.
primigenius, Acipenser, 32.
Pristigaster, 110, 261, (427).
americanus, 429.
cayana, (428).
cayanus, 427, 429.
Index of Scientific Names 625
chinensis, 4.16.
dolloi, 430.
flavipinnis, 416, 427.
martit, 428-430.
mucronatus, 433.
phaeton, 429.
SP.» 427.
vanderbilti, 425, 426.
Pristigasterinae, 96, 261.
producta, Anchovia, 158.
productus, Alepocephalus, 251.
Cylindrosteus, 75.
Engraulis, 157, 247-
Stolephorus, 157, 158.
Prosopium, 547, 548, 550, 551.
quadrilateralis, 551.
Protozoa(n), 286PD, 484 P.
Psallisostomus, 69.
Psephurus gladius, 26.
Pseudaliprotus, 518F.
Pseudochirocentrodon, 415, 416.
amazonicum, 416, 423.
pseudoharengus, Clupea, 312, 331, 341, 342.
Pomolobus, 312-314, 317, 328, 330, 331, (332)-
Sardinella, 405.
lacustris, Pomolobus, 342.
pseudohispanica, Clupea, 407.
pseudo-hispanica, Sardinia, 397, 407.
Sardinella, 405, 407.
pseudohispanicus, Clupea, 406.
Clupanadon, 406.
Clupanodon, 405, 4.10.
Pseudopleuronectes, 588E.
Pseudoscaphirhynchus, 27, 28.
Prerengraulis, 153, (228), 231.
atherinoides, (229), 231.
Pterothrissidae, 92, 93, 98, 107, 108.
Prerothrissus, 93, 94, 107, 108, 133.
pusilla, Clupea, 324.
pyriforme, Distomum, 356P.
Qwapaiareratis, Prosopium, 551.
Raptans, Lernaeenicus, 374P.
radiatus, Lernaeenicus, 303P, 356P.
rafinesquit, Acipenser, 46.
rapax, Caligus, 303P, 338P.
rastralis, Anchovia, 156, 161.
rauchi, Acipenser, 46.
regalis, Cynoscion, 329 E, 331E, 343.
Reinhardtius hippoglossoides, 588E.
relicta, Trutta, 498.
Retropinnatidae, 92.
Retropinnidae, 533.
Rhinosardinia, 261, (411).
Rhinosardinia amazonica, (412), 415.
bahiensis, 411.
Serrata, 411, 412, (413).
Rhipidistia, 9.
Rhomboganoidea, 64.
rhynchaeus, Acipenser, 46.
Rhynchobothrium bulbifer, 128P.
imparispine, 338P.
sp. 197 P.
Rhynchoceratias, 7.
richardsoni, Acipenser, 46.
rivalis, Salmo, 524.
robertsi, Anchoviella, 247.
Stolephorus, 247.
Roccus saxatilis, 323, 536E.
Rogenia, 274.
alba, 274.
rosarium, Acipenser, 46.
rossit, Salmo, 523.
Salvelinus, 524.
rostellum, Acipenser (Huso), 41.
rostrata, Albula, 141.
rostratus, Lepidosteus, 79.
Rotifer, 355 F.
rubicundus, Acipenser, 46.
rupertianus, Acipenser, 46.
ruthenus, Acipenser, 28, 29, 31, 32.
Acipenser, x A. baeri, 32.
Acipenser, x A. gildenstddti, 32.
SaccopHaRyNcors, 6.
sadina, Alosa, 361.
Clupea, 267.
Etrumeus, (263).
Sagamichthys, 254.
saida, Boreogadus, 482F, 518F.
Salangidae, 92, 455, 456.
Salanx, 456.
Salar, 459.
salar, Salmo, 459, (460), 498, 507, 523, 525, 526,
5435 544, 573-
626 Index of Scientific Names
brevipes (var.), Salmo, 493.
nobilis (var.), Salmo, 493.
sebago, Salmo, 492.
trutta (var.), 493.
Trutta, 498.
Salariinae, 6.
salmarinus, Salmo, 503.
Salmincola salmonea, 484P.
Salmo, 455, 457, 458, (459), 503, 541-543, 5555
573-
agassiZit, 505, 537, 542.
albula, 141.
alipes, 524.
alleganiensis, 541.
alpinus, 503, 505, 506, 523.
alpinus malma, 523.
arcticus, 573, 592.
arcturus, 524.
argenteus, 498.
(or Fario) argenteus, 459.
(or Salar) ausonii, 459.
bairdii, 525.
callaris, 525.
canadensis, 541.
carpio, 505, 506, 523.
curilus, 525.
eperlano-marinus, 569.
eperlanus, 554-556, 569, 573.
(Osmerus) eperlanus, 555, 569.
(Osmerus) eperlanus eperlanus, 569
erythrorynchos, 525.
fario, 541.
(or Salar) fario, 459.
Sontinalis, 541.
gairdneri, 8, 459, 460, 491, (499), 525, 526.
gloveri, 498.
gracilis, 498.
groenlandicus, 573, 592-
hamatus, 497.
hardinii, 492, 498.
Audsonicus, 537, 541.
immaculatus, 497, 523, 541.
lacustris, 493, 503.
laevigatus, 525.
lavaretus, 549.
lordii, 525.
malma, 506, 523, 525.
(Salvelinus) marstoni, 506, 509, 524.
namaycush, 503, 504, 542, $44.
naresti, 524.
nigrescens, S41.
nitidus, 524.
nobilis, 497.
oguassa, 506, 524.
ouananiche, 492, 498.
parkei, 525.
pluvius, 525.
rivalis, 524.
rossil, 523.
salar, 459, (460), 498, 507, 523, 525, 526, 543,
544) 573-
salar var. brevipes, 493.
salar var. nobilis, 493.
salar sebago, 492.
salmarinus, 503.
salmo, 497.
salvelinus, 503, 505.
samulus, 497.
schiefmulleri, 497.
sebago, 492, 498.
siscowet, 543.
socialis, 573.
spectabilis, 525.
spirinchus, 556.
(Osmerus) spirinchus, 555, 569.
(Osmerus) spirinchus eperlanus, 569.
(Osmerus) spirinchus mordax, 569.
stagnalis, 506, 523, 524.
symmetrica, 541, 544.
toma, 544.
tructa, 544.
trutta, 10, 459-461, 482F, 483E, 484, 492, 493.
(498), 525, 526, 537, 541.
(or Fario) trutta, 459.
umbla, 503, 505.
villosus, 592.
(Mallotus) villosus, 594.
salmo, Salmo, 497.
salmonea, Salmincola, 484P.
Salmones salvelines, 503.
salmonicida, Bacillus, 484D.
Salmonid, or Salmonoid, 6-8, 90, 91, 549.
Salmonidae, 14, 91, 99, 455, 456, (457), 547, 553-
Salmoniformes, 93.
Salmonina, 93.
salmonis, Lepeophtheirus, 484P, 494P, 496P, 497P.
pestis, Bacillus, 484D.
Salmonoidea, 14, 92, 93, 96, (455)-
Index of Scientific Names 627
Salmonoidei, 93.
Salmonoides, 93.
salvatoris, Anchoviella, 188.
salvelines, Salmones, 503.
Salvelinus, 455, 457, 458, (503)5 541-543-
agassiZit, 542.
alpinus, 15, 458, 460, 491, 498, 504-506, (507),
525, 526, 541, 542, 588E.
alpinus alipes, 523, 524.
alpinus arcturus, 523, 524.
alpinus aureolus, 524, 541, 542.
alpinus malma, 484, 506, 519, 520.
alpinus marstoni, 524.
alpinus naresii, 523.
alpinus oquassa, 523, 542.
alpinus stagnalis, 523.
alpinus forma stagnalis, 524.
arcturus, 524.
aureolus, 506, 509, 524.
bairdti, 525.
fontinalis, 460, 483E, 484, 498, 499, 504-507,
509, 510, 515, 518, 522, 523, (525), 544.
(Baione) fontinalis, 504, 542.
Sontinalis agassiZit, 542.
Sontinalis hudsonicus, 537, 542.
malma, 519, 520, 525.
marstoni, 506, 509.
namaycush, 544.
naresit, 524.
oguassa, 522, 524.
oquassa naresii, 524.
rossit, 524.
spectabilis, 525.
stagnalis, 523, 524.
umbla stagnalis, 523.
salvelinus, Salmo, 503, 505.
samulus, Salmo, 497.
sapidissima, Alausa, 307.
Alosa, 39, 294, (295), 309-312, 317, 343-
Clupea, 306, 308, 311.
sapidus, Callinectes, 67 F, 74¥, 79F, 86F.
Sarchirus, 66, 68, 69.
argenteus, 81.
vittatus, 69, 79-
Sarcopterygii, 10, IT.
Sardina, 397.
sardina, Clupea, 389.
Harengula, 389, 392, 396.
Sardinella, 389.
Sardinella, 258, 261, (397)-
(Rhinosardinia) amazonica, 413.
anchovia, 393, 398-400, (401), 409-411.
apicalis, 389.
aurita, 397, 405-407, 410, 411.
bishopi, 393-
brasiliensis, 398, 399, 405, 406, (407).
clupeola, 393.
humeralis, 390, 395.
macrophthalma, 390.
macrophthalmus, 390, 393, 396-
maculosa, 390.
pinnula, 398, (399)-
pseudoharengus, 405.
pseudo-hispanica, 405, 407.
Sardina, 389.
Sardinia, 397-
anchovia, 397, 407.
pseudo-hispanica, 397, 407-
Sardinops, 398.
saurus, Elops, 115, 123, (124), 142.
saxatilis, Roccus, 323, 536E.
Scaphirhynchini, 27.
Scaphirhynchus, 27, 28.
album, 28.
schiefmulleri, Salmo, 497.
Schipa, 28.
schistonyx, Caligus, 356P.
Schizopod (shrimp), 285 F.
schrencki, Acipenser, 32.
schroederi, Lycengraulis, 241, 243.
schypa, Acipenser, 28.
Sciaenidae, 343.
Scolex polymorphus, 197 P, 322P, 356P.
Scomber, 7, 323-
Scombridae, 7.
Scopelarchidae, ro4.
Scopelengys, 100.
Scopeliformes, 92.
Scopelosauridae, 100.
Scyphophori, 92.
Searsia, 254.
Searsia koefoedi, 255.
Searsiidae, 14, 95, 102, 148-150, 250, (254).
Searsinae, 254.
sebago, Salmo, 492, 498.
sebago (subsp.), Salmo salar, 492.
Semionotidae, 65.
Semionotoidea, 64.
628 Index of Sctentific Names
semiradiatus, Lepidosteus, 79.
senegalensis, Elops, 128, 130.
sergeanti, Osmerus, 556, 564, 570-572.
sericus, Engraulis, 133, 141.
serotinus, Acipenser, 28, 46.
Serrasalmus, 5.
serrata, Rhinosardinia, 411, 412, (413).
setiger, Hildebrandichthys, 230, (231).
Signalosa, 398, 443.
atchafalayae, 443, 451.
atchafalayae atchafalayae, 451.
atchafalayae vanhyningi, 451.
mexicana, 451.
mexicana campi, 451.
mexicana mexicana, 451.
petenensis, 451.
petenensis atchafalayae, 451.
petenensis campi, 451.
petenensis vanhyningi, 451.
signifer, Chatoessus, 384.
silus, Argentina, 8.
Simulid, 481 F.
Simulidae, 536F.
sinensis, Acipenser, 32.
Lepisosteus, 67.
sinuatus, Pandarus, 356P.
sinus, Acipenser (Huso), 41.
siscowet, Salmo, 543.
smithi, Brevoortia, 343, 344, 346, 347, (372), 379-
Dorosoma, 443.
Lepidosteus, 81.
socialis, Mallotus, 574, 594.
Salmo, 573.
Somniosus microcephalus, 588E.
spathula, Polyodon, 26.
spatula, Lepisosteus, 61-63, 67, 70, 71, 75, 76, 80,
81, (83).
spectabilis, Salmo, 525.
Salvelinus, 525.
spectrum, Osmerus, 556, 557, 572-
spectrum (subsp.), Osmerus mordax, 572.
spectrum (var.), Osmerus mordax, 572.
spinifer, Anchoa, 162, 163, 166, (167), 169.
Engraulis, 168.
Stolephorus, 168.
Spinifera, Anchovia, 168.
Anchoviella, 169.
spinosa, Clavellisa, 356P.
spinosus, Eumicrotremus, 5181.
Spirinchus, 553, 554-
spirinchus, Osmerus, 556, 570, 571.
Salmo, 556.
Salmo (Osmerus), 555, 569.
eperlanus, Salmo (Osmerus), 569.
mordax, Salmo (Osmerus), 569.
spirinchus (morpha), Osmerus eperlanus, 557.
Osmerus eperlanus dentex natio dvinensis, 557, 569,
572.
Sporozoa, 286P, 356P, 362 P.
Spratelloides, 267.
lamprotaenia, 272.
sprattus, Clupea, 482F.
Squalus acanthias, 537K, 588E.
stagnalis, Salmo, 506, 523, 524.
Salvelinus, 523, 524.
stagnalis (subsp.), Sa/velinus alpinus, 523.
Salvelinus umbla, 523.
stagnalis (forma), Sa/velinus alpinus, 524.
stanley (subsp.), Coregonus clupeaformis, 550.
stellatus, Acipenser, 28, 29, 32.
Stenodus, 547.
stenorhynchus, Lepisosteus, 81.
Sterleta, 28.
Sterletus, 28.
Sternoptychidae, go, 102.
Stolephorus, 274.
argyrophanus, 210.
astilbe, 188, 190.
browni, 199.
brownii, 188, 193, 200, 215.
cayennensis, 213.
choerostoma, 186, 202.
clupeoides, 157, 158.
cubanus, 190.
eurystole, 210.
Larmani, 247.
gilberti, 247.
guianensis, 217, 219, 220.
lyolepis, 202.
manjuba, 247.
miarchus, 215.
mitchilli, 174, 180, 181.
opercularis, 247.
perfasciatus, 210, 215, 216.
perthecata, 199.
productus, 157, 158.
robertst, 247.
spinifer, 168.
Index of Scientific Names 629
Surinamemsis, 157, 243, 247.
viridis, 272, 274.
stolifera, Dussumieria, 267, 271.
Fenkinsia, 271, 272, 274.
Stomiatidae, 102.
Stomiatina, 93.
Stomiatoid, 5, 7, 20, 90, 95.
Stomiatoidea, 89, 91, 93, 95, 96.
Stomiatoidei, 93.
storeri, Acipenser, 56.
striata, Alausa, 385.
Strongylura, 272.
Sturio, 28.
vulgaris, 28.
sturio, Acipenser, 25, 27-29, 32, 35, 47, 49, 545
56-58.
oxyrhynchus, Acipenser, 56.
Acipenser, x A. brevirostris, 54.
Stylophorid, or Stylophoroid, 4, 7.
suoerii, Meletta, 319.
surinamensis, Anchoviella, 247.
Engraulis, 157, 221.
Stolephorus, 157, 243, 247.
Symbranchii, 12.
symmetrica, Salmo, 541, $44.
Synbothrium, 356P.
Syngnathid, 13.
Syngnathus, 9.
Synodontidae, roo.
Synodus, 133, 142.
argenteus, 133.
vulpes, 133.
"Tacrosroma, go.
taeniatus, Chirocentrodon, 438, 442.
Tarpon, 93, 107, IIT, (112), 123.
atlanticus, (113).
giganteus, 121.
Teleostea, 4.
Teleostei, 3, 13, 14.
Teleostomi, 3, 13.
Teleostomii, 4.
teres, Alosa, 267.
Bomolochus, 356P, 370P.
Etrumeus, 267.
ternetzi, Ilishe (Neosteus), 426.
Neosteus, 425, 426.
Tetraodontid, 7.
Tetrapod vertebrates, 11.
Thaleichthys, 553, 554.
Therobromus callorhini, 554.
thompsont, Lepidosteus, 81.
thrissa, Chatoessus, 384.
Clupanodon, 380.
Clupea, 380, 384, 385.
Opisthonema, 380, 385.
Meletta, 384.
thrissina, Harengula, 387.
thrissoides, Megalops, 121.
Thymallidae, 455, 456.
Thymallus, 8.
Tipulid, 481F.
toma, Salmo, 544.
Trachinocephalus, 100.
Trachonotus, 123.
Trachypteroid, 4, 7.
transmontanus, Acipenser, 29, 31-33, 36, SI.
treculi, Lepidosteus, 81.
Trematode, 197 P, 271 P, 285 P, 285 P (larva),
286P, 286P (larva), 322P, 338P, 356P,
362 P, 404P, 484P, 495 P, 571 P.
Triacanthidae, 7.
Trichoptera, 536F.
tricolor, Anchoa, 162, 164, 166, (186), 199,
200.
Engraulis, 188, 199.
trinitatis, Anchoa, 162, 163, 166, (171).
Anchovia, 172.
Anchoviella, 173.
Triplophos, 103.
Tripteronotus, 549.
hautin, 549.
tristoechus, Lepisosteus, 70.
troostit, Lepidosteus, 81.
tropicus, Lepisosteus, 67, 70.
Neoopisthopterus, 4.36, 438.
Odontognathus, 432, 435.
tructa, Salmo, 544.
truncatus, Dinectus, 46.
Trutta, 459.
argentina, $41.
marina, S41.
relicta, 498.
salar, 498.
trutta, Salmo, 10, 459-461, 482F, 483E, 484, 492,
493, (498), 525, 526, 537, 541.
Salmo or Fario, 459.
630 Index of Scientific Names
tshawytscha, Oncorhynchus, 484.
tyrannus, Alosa, 332, 362.
Brevoortia, 270, 342-345, (346), 363-365, 368—
371, 373-375.
Clupea, 361.
brevicaudata, Brevoortia, 365.
aurea (var.), Brevoortia, 361.
brevicaudata (var.), Brevoortia, 361, 365.
menhaden (var.), Brevoortia, 361.
Unarsus, Bonasa, 488.
Umbla, 503.
umbla, Salmo, 503, 505.
stagnalis, Salvelinus, 523.
Uranoscopidae, 12.
Uranoscopus, 12.
Urophycis, 536¥.
Vanpereitt, Pristigaster, 425, 426.
vanhyningi (subsp.), Signalosa atchafalayae, 451.
Signalosa petenensis, 451.
venezuelae, Anchoviella, 221, 223.
venosa, Meletta, 341.
vernalis, Clupea, 331, 340, 341.
villosa, Clupea, 573, 592.
villosus, Mallotus, 559, 574, 575, (576)-
Salmo, 592.
Salmo (Mavlotus), 594.
catervarius, Mallotus, 575, 583, 594.
villosus, Mallotus, 575 (576).
villosus (subsp.), Ma/lotus villosus, 575, (576).
virens, Nereis, 52¥.
Pollachius, 536E.
virescens, Clupea, 342.
viridescens, Osmerus, 556, 569, 571.
virdis, Esox, 79.
viridis, Fenkinsia, 268, 271, (272).
Stolephorus, 272, 274.
vitellosum, Distomum, 338P, 356P.
vittata, Clupea, 198.
vittatus, Engraulis, 199.
Sarchirus, 69, 79.
vulgaris, Alausa, 293.
Eperlanus, 556.
Sturio, 28.
Vulpes bahamensis, 141.
vulpes, Albula, 115, 124, 126, 129, 131, 133, (134),
145, 410.
Butirinus, 133, 141.
Esox, 133, 141, 142.
Synodus, 133.
vultharius, Acipenser, 28.
Xenovermicuruys, 250.
copei, 251.
Xenophthalmichthys, 104.
Xiphias, 6.
Zosrera marina, 538.
Zunasia, 416.
Fishes of the
Western North Atlantic
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