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XFWS-A 591 1-47 (1970)
U.S, Fish Wildl, Serv.

591 Spec. Sci. Rep. Fish. AP R 1 : 970 :

DIVISION OF Fic
A Bibliography of the Lobsters,
Genus Homarus

SPECIAL SCIENTIFIC REPORT-FISHERIES No. 591

UNITED STATES DEPARTMENT OF THE INTERIOR ~~

‘U.S. FISH AND WILDLIFE SERVICE “~~
BUREAU OF COMMERCIAL FISHERIES

SPECIAL SCIENTIFIC REPORT--FISHERIES

Robert L. Hacker, Editor

Mary Fukuyama, Associate Editor

PUBLICATION BOARD

John A, Guinan John M, Patton, Jr.
Robert L. Hacker Edward A, Schaefers
John I. Hodges Parker S,. Trefethen
Harvey Hutchings Robert C. Wilson

Leslie W. Scattergood, Chairman

Special Scientific Report--Fisheries are preliminary or progress reports
and reports on scientific investigations of restricted scope. Established as
Special Scientific Reports in 1940, nos. 1 to 67 were issued from that date
to 1949, when the new series, Special Scientific Report--Fisheries, with new
serial numbering, was started.

Special Scientific Report--Fisheries are distributed free to libraries,
research institutions, State agencies, and scientists.

A Bibliography of the Lobsters, Genus Homarus

By

R, D, LEWIS

United States Fish and Wildlife Service
Special Scientific Report--Fisheries No. 591

Washington, D.C.
January 1970

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A Bibliography of the Lobsters, Genus Homarus

R. D. LEWIS, Fishery Biologist

Bureau of Commercial Fisheries Biological Laboratory
West Boothbay Harbor, Maine 04575

ABSTRACT

A total of 1,303 references are given,

INTRODUCTION

This bibliography was begun in the summer
of 1964, It was mimeographed and distributed
at a meeting of United States and Canadian
scientists concerned with the biology of the
American lobster held on November 9-10,
1965, at the Bureau of Commercial Fisheries
Laboratory, West Boothbay Harbor, Maine,

References in the 1965 manuscript were
compiled from the original papers, their cita-
tions, and the bibliographies of Herrick(1911),
Scattergood (1949), and Dawson (1954). The
present list also includes references from
the bibliography of Bergeron (1965).

About 80 percent of the references were
found in the library of the Marine Biological
Laboratory at Woods Hole, Mass., and the
rest at the libraries of the Fisheries Research
Board of Canada, Biological Station, St,

Andrews, New Brunswick; the Bureau of Com-
mercial Fisheries Biological Laboratory, West
Boothbay Harbor, Maine; and the Department
of Interior Library and the Library of
Congress, Washington, D.C,

Lists of references that I had overlooked
in the 1965 manuscript were provided by
H. J. Thomas, Department of Agriculture and
Fisheries for Scotland, Marine Laboratory,
Aberdeen; D. G. Wilder, Fisheries Research
Board of Canada, Biological Station, St.
Andrews, New Brunswick; and R.J. Ghelardi,
Fisheries Research Board of Canada, Biologi-
cal Station, Nanamio, British Columbia.
The library staffs of the Marine Bio-
logical Laboratory at Woods Hole and the
Biological Station at St. Andrews assisted
me,

BIBLIOGRAPHY

A

Abbott, Bernard C., and I. Parnas.

1965. Electrical and mechanical responses
in deep abdominal extensor muscles
of crayfishandlobster. J. Gen. Physiol,
48: 919-931.

Abelous, J.-E., and G, Billard.
1897, De l'action anticoagulante du foie des
crustacés. C, R, Soc, Biol, Ser, 10, 49:
991-993.

Ackerman, Edward A.
1941. New England's fishing industry. Uni-
versity of Chicago Press, Chicago,
303 pp.

Adelman, W. J., Jr., and J. C. Dalton,

1960. Interactions of calcium with sodium
and potassium in membrane potentials
of the lobster giant axon. J. Gen.
Physiol, 43: 609-619.

Alexandrowicz, J. S.

1951. Muscle receptor organs in the abdo-
men of Homarus vulgaris and Palinurus
vulgaris, Quart J. Microsc, Sci. Ser. 3,
92: 163-199,

1952. Receptor elements in the thoracic
muscles of Homarus vulgaris and
Palinurus vulgaris. Quart. J. Microsc,
Sci. Ser. 3, 93: 315-346,

1953, Nervous organs in the pericardial
cavity of the decapod Crustacea. J. Mar.
Biol. Ass. U.K. 31: 563-580.

1967, Receptor organs in thoracic and ab-
dominal muscles of Crustacea. Biol.
Rev. (Cambridge) 42: 288-326.

Alexandrowicz., J. S., and Mary Whitear,
1957, Receptor elements in the coxal region
of decapod Crustacea. J. Mar. Biol.
Ass, U.K. 36: 603-628.

Allen, Edgar J.
1894, Nerve elements of the embryonic
lobster. J. Mar. Biol. Ass. U.K., New
Ser. 3: 208-209.

1894, Studies on the nervous system of
Crustacea. Parts l=35 Quart.) “Ji.
Microsc, Sci. 36: 461-498,

1895. On the nervous system of the em-
bryonic lobster, Rep. Brit. Ass, Advan,
Sci. 65th Meet., pp. 470-471.

1895. The reproduction of the lobster,
J. Mar. Biol. Ass. U.K., New Ser.
4: 60-69.

1895, Additional observations on thenerve-

elements of the embryonic lobster,
J. Mar. Biol. Ass. U.K., New Ser.
A BiOawen

1896. The protection of crabs and lobsters,
J. Mar. Biol. Ass, U.K., New Ser,
4; 182-187.

1897. Studies on the nervous systems of
Crustacea, Part 4, Quart. J. Microsc,
Sci. 39: 33-50.

Alley, Armine,

1934, The effect upon the gastric juice
secretion of various cooked prepara-
tions of haddock (Melanogrammus
aeglefinus) and of lobster (Homarus
americanus), Amer. J. Dig. Dis. Nutr.
1: 182-184,

Allison, James B., and William H. Cole,
1940, The nitrogen, copper, and hemocyanin
content of the sera of several arthro-
pods. J. Biol. Chem. 135: 259-265,

Allison, William S., and Nathan O, Kaplan.
1964, The comparative enzymology of
triosephosphate dehydrogenase. In
Charles A, Leone (editor), Taxonomic
biochemistry and serology, pp. 401-406,
Ronald Press Co., New York,

Altman, Philip L., and Dorothy S,. Dittmer

(editors).
1961. Blood and other body fluids. Federa-
tion of American Societies for Experi-

mental Biology, Washington, D.C.,
540 pp.
Amberson, W. R., H. S. Mayerson, and
W. J. Scott.

1925, The influence of oxygen tension upon
metabolic rateininvertebrates. J.Gen,
Physiol. 7: 171-176.

Anderton, T.

1909. The lobster (Homarus vulgaris).
N. Z. Mar. Dep. Rep. 1908-1909, pp, 17-
238

Appellof, A.
1899, Mittheilungen aus der Lebensweise
des Hummers. Mitt. Deut. Seefisch-
erei-Ver. 15: 99.

1903. Hummerundersggelser i 1902. Norsk
Fiskeritid. 22: 114-132.

1904. Hummerunders¢gelser i 1903. Norsk
Fiskeritid. 23: 112-119.

1909. Undersgkelser over hummeren
(Homarus vulgaris) med saerkskilt,
hensyn til dens optraeden ved Norges
kyster,. Arsberetn. Vedkom. Norg.
Fisk. 1909(1): 1-148.

1909, Untersuchungen ueber den Hummer
mit besonderer Berucksichtigung
seines Auftretens an den norwegischen
Kusten. Bergens Mus. Skr., New Ser.
1(1): 2-79.

Arudpragasam, K, D., and E, Naylor.
1966, Patterns of gill ventialtion in some
decapod Crustacea. J. Zool. (London)
150: 401-411,

Atwood, N. E,
1864, The habits and geographical distri-
bution of the common lobster. Proc,
Boston Soc, Natur. Hist. 10: 11-12,

B

Baird, Frederick T., Jr.
1950. Lobster plugs and their effect on the
meat of the lobster's claw. Maine Dep.
Sea Shore Fish., Res. Bull. 2,12pp.

1953, Probable effects of a size change upon
the lobster industry. Maine Dep, Sea
Shore Fish., Fish. Circ. 9, 10 pp.

Ball, Erik G., and Bettina Meyerhof,

1940, On the occurrence of iron porphyrin
compounds and succinic dehydrogenase
in marine organisms possessing the
copper blood pigment hemocyanin. J.
Biol. Chem, 134: 483-493,

Bales, Henrich,
1930, Wanderungen bei Decapoden (Crusta-
ceen), Ergeb, Biol. 6: 305-326.

1944, Decapoda. In H, G, Bronn's Klassen
und Ordnungen des Tierreichs 1940, III,
Morphologischer Teil. Bd. 5, Abt. 1,
Buch 7: 22-671, Akademische Verlags-
gesellschaft, Geest & Portig K-G., Leip-
zig.

Barnes, Earnest W,
1906. Lobster culture in 1905, R.I, Comm,
Inland Fish., 36th Annu, Rep., pp. 111-
119.

Barnes, Earnest W,.--Con.

1906. Methods of protecting and propagating
the lobster, with a brief outline of its
natural history. R.I. Comm. Inland
Fish., 36th Annu, Rep., pp. 120-152.

1907. Lobster culture at Wickford, R.I. in
1906. R.I. Comm, Inland Fish., 37th
Annu. Rep., pp. 89-94.

1911. Revised edition of ''Methods of pro-
tecting and propagating the lobster, with
a brief outline of its natural history,"'
R.I, Comm, Inland Fish., 4lst Annu.
Rep., Append. A., pp. 83-127.

1911. The season of 1910 at the fisheries
experiment station at Wickford, R.I.
Trans, Amer, Fish. Soc. 40: 365-369.

1939, An analysis of the objectives oflobster
rearing and problems of reinvigorating
the lobsterindustry. Commonw. Mass,
House Doc, 2051, Append. A, pp. 10-32.

1939. The lobster fishery, What maybe done
to benefit it? North Atlantic Confer-
ence on Lobster Conservation, Boston,
Feb, 2, 10 pp.

Barnes, T. C., J. P. Sell, and W. Spofford.
1932. Responses of a teratological crus-
tacean limb to electrical stimulation.
Z. Vergl. Physiol. 18: 280-283.

Bateson, William.

1894, Materials for the study of variation
treated with especial regard to dis-
continuity in the origin of species.
Macmillan and Co., London, 598 pp.

Bean, Tarleton H.
1896. Report on the propagation and distri-
bution of food-fishes. U.S. Comm, Fish
and Fish., Rep. Comm. 1894, Pt, 20:
20-80.

Bell, F. Heward, and Alonzo T. Pruter,

1958. Climatic temperature changes and
commercial yields of some marine
fishes, J. Fish. Res. Bd. Can, 15: 625-
683.

Belloc, Gerard.
1935. Le péche au crustacés. In E. Le
Danois (editor), Manuel des péches
maritimes francaises, No.10: 127-139.

Bennitt, R.
1932, Physiological interrelationship in the
eyes of decapod Crustacea, Physiol.
Zool, 5: 49-64,

Bergeron, Julien.

1961. Rapport préliminaire des travaux sur
le Homard (Homarus americanus) aux
Tles-de-la- Madeleine au cours de la
saison 1960, Sta. Biol, Mar, Grande-
Riviere Que., Rapp. Annu. 1960: 50-58,

1961. Résultats préliminaires de l'emploi
des casiers en aluminium pour la péche
du Homard aux Iles-de-la- Madeleine.
Sta, Biol. Mar. Grande-Riviere Qué.,
Rapp. Annu, 1960: 113-115,

1962. La péche commerciale du Homard aux
Tles-de-la-Madeleine en 1961, sta-
tistiques et échantillonnage. Sta. Biol,
Mar. Grande-Riviere Qué., Rapp. Annu,
1961: 45-49,

1962, Péche expérimentale du Homard dans
la lagune de Havre-aux-Maisons et
dans la baie de Plaisance, Iles-
de-la-Madeleine en 1961. Sta. Biol.
Mar. Grande-Riviere Qué., Rapp. Annu,
1961: 51-61.

1962. Efficacite des casiers en aluminium
pour le péche du Homard aux Iles-de-
la- Madeleine. Sta. Biol. Mar. Grande-
Riviere Qué., Rapp. Annu. 1961: 119-
124,

1963. La péche commerciale du Homard
aux Iles-de-la- Madeleine en 1962,
statistiques et echantillonnage. Sta.
Biol. Mar. Grande-Riviére Qué., Rapp.
Annu, 1962: 147-153,

1963. Taille minimale legale et captures
commerciales de homards (Homarus
americanus) aux Iles-de-la- Madeleine
en 1962. Sta, Biol. Mar. Grande-Riviere
Qué,, Rapp. Annu, 1962: 155-160.

1964, Deuxiéme échantillonnage des cap-
tures completes de homards aux Iles-
de-la- Madeleine-1963. Sta, Biol. Mar.
Grande-Riviére Qué., Rapp. Annu. 1963:
79-84,

1964, La péche commerciale du Homard
aux Iles-de-la-Madeleine en 1963,
statistiques et échantillonnage. Sta.
Biol. Mar. Grande-Riviére Qué., Rapp.
Annu, 1963: 85-94,

1965, Bibliographie du Homard (Homarus
americanus Milne-Edwards et Homarus
gammarus L,), Sta. Biol. Mar. Grande-
Riviére Qué. Cah. Inform. 34, 81 pp.

1966, La péche commerciale du Homard
(Homarus americanus M.Edw.) auxIles-
de-la- Madeleine, au cours dela période
1950-1964, Sta. Biol, Mar. Grande-
Riviere Qué., Cah. Inform, 37, 25pp.

Bergeron, Julien--Con,

1967. La péche commerciale du Homard
(Homarus americanus Milne-Edwards)
au Québec des origines a nos jours,
Sta. Biol. Mar. Grande-Riviere Qué,,
Cah, Inform. 42, 47 pp.

1967. Contribution a la biologie du Homard
(Homarus americanus M.-Edw.) des
Iles-de-la- Madeleine, Natur. Can, 94:
169-207, [Trav. Péche. Qué, 14.]

Bergquist, H.

1963. Kutikula-bildung bei Homarus-
embryonen, eine elektronenmikrosko-
Ppische Studie. Verh, Anat, Ges., Jena,
57: 348-362.

Berrill, N. J.
1961. Maine lobster, In N. J. Berrill, The
living tide, pp. 108-115. Dodd, Mead
and Co., New York,

Bethe, Albrecht.

1930. Studien uber die Plastizitat des
Nervensystems, I. Mitteilung. Arach-
noideen und Crustaceen. Pfluger's
Arch, Gesamte Physiol. Menschen Tiere
224: 793-820.

Bethe, Albrecht, and.E, Berger.

1931. Variationen im Mineralbestand ver-
schiedener Blutarten, Pfluger's Arch,
Gesamte Physiol. Menschen Tiere 227:
571-584,

Bhagvat, Kamala, and Derek Richter.
1938, Animal phenolases and adrenalin,
Biochem, J. 32: 1397-1406.

Biedermann, W.
1901. Uber den Zustand des Kalkes im
Crustaceenpanzer. Biol. Zentralbl. 21:

343-352.

Bilinski, E.

1960. Biosynthesis of trimethylammonium
compounds in aquatic animals. I. For-
mation of trimethylamine oxide and
betaine from C?*-labelled compounds
by lobster (Homarus americanus),
J. Fish, Res. Bd. Can, 17: 895-902.

1961, Biosynthesis of trimethylammonium
compounds in aquatic animals, II. Role
of betaine in the formationoftrimethyl-
amine oxide by lobster (Homarus ameri-
canus).$sJi)\ Fish: Res) “Bdt Can, 18:
285-286.

1962, Biosynthesis of trimethylammonium
compounds in aquatic animals, III. Cho-
line metabolism in marine crustacea,
J. Fish. Res. Bd. Can. 19: 505-510.

Bjerkan, Paul.

1926, Hummerfredninger i England. Norsk
Fiskeritid, 45: 37-41. [Translated by
Arden Nilsen for Bur. Commer. Fish,
Biol, Lab., W. Boothbay Harbor, Maine,]

Blaschko, H., McKeen Cattell, and J. L. Kahn.

1931. On the nature of the two types of

response in the neuromuscular system

of the crustacean claw. J. Physiol.
(London) 73: 25-35,

Blethen, Sandra L., and NathanO, Kaplan,
1967, Purification of arginine kinase from
lobster and a study of some factors
affecting its reactivation. Biochemistry
6: 1413-1421,

Bligh, E. G., W.J. Dyer, and D. C. Horne.
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Annu. Rep., pp. 16-17.

Their

1910. Propagation of lobsters.
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Their 40th

1910. Changes in the lobster rearing ap-
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1911. The propagation of lobsters at the
Wickford experiment station. Their
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1912. The propagation of lobsters at the
Wickford experiment station. Their
42d Annu. Rep., pp. 46-47.

1913. The propagation of lobsters at the
Wickford experiment station, Their

43d Annu. Rep., pp. 30-31.

1914. The propagation ‘of lobsters at the
Wickford hatchery and experiment sta-
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1915. The propagation of lobsters at the
Wickford hatchery and experiment sta-
tion, Their 45th Annu. Rep., pp. 34-35.

1916, Propagation of lobsters at Wickford,
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1917. Propagation of lobsters at Wickford.
Their 47th Annu, Rep., pp. 23-24.

1918. Propagation of lobsters at Wickford,
Their 48th Annu, Rep., pp. 24-25.

1919. Propagation of lobsters at Wickford.
Their 49th Annu, Rep., pp. 26-27.

1920. Propagation of lobsters at Wickford.
Their 50th Annu. Rep., pp. 26-28.

1921. Propagation of lobsters at Wickford,
Their 51st Annu, Rep., pp. 28-29.

1922, Propagation of lobsters at Wickford,
Their 52d Annu, Rep., pp. 24-25.

1923. Propagation of lobsters at Wickford.
Their 53d Annu. Rep., p. 14.

Rhode Island Commissioners of Inland Fish-
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1924, Propagation of lobsters at Wickford.
Their 54th Annu. Rep., pp. 14-15.

1925. Propagation of lobsters at Wickford.
Their 55th Annu, Rep., p. 16.

1926. Propagation of lobsters at Wickford.
Their 56th Annu, Rep., pp. 19-20.

1927. Propagation of lobsters at Wickford,
Their 57th Annu, Rep., pp. 12, 15.

1928. Total number of fourth and fifth stage
lobsters reared each year since 1900,
Their 58th Annu. Rep., p. 15.

1929. Propagation of lobsters at Wickford,
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1930. (Lobster rearing). Their 60th Annu,
Rep., pp. 10, 13.

1931. (Lobster rearing). Their 61st Annu,
Rep-, Pp. 5.

1932. Wickford hatchery. Their 62d Annu,
Rep., p. 24.

1933. Wickford hatchery. Their 63d Annu.
Rep., p. 22.

1934. Propagation of lobsters at Wickford.
Their 64th Annu. Rep., p. 20.

1935. Propagation of lobsters at Wickford.

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Rhode Island Department of Agriculture and
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1936. Its Annu. Rep. 1935, 20 pp.

1937. 1936:

Its Annu.

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1938. Its Annu. Rep. 1937, 30 pp.

1939.
1940.

Rep, 1938,

1939,

26 pp.
26 pp.

Its Annu,

Its Annu, Rep.

1941, Its Annu, Rep. 1940, 21 pp.

1942, Its Annu. Rep. 1941, 24 pp.

1943. Its Annu, Rep. 1941, 12 pp.

1944, Its Annu. Rep. 1943, 17 pp.

1945. Its Annu. Rep. 1944, 20 pp.

1946.
1947.
1948.

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1947, 18 pp.

Its Annu. Rep.

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americanus Milne-Edwards, and Li-
binia emarginata Leach). Cellule 48:
147-176.

Worth, S. G.

1895. Report on the propagation and dis-
tribution of food-fishes. U.S. Comm,
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Pt. 19: 78-138.

Yi

Yeager, J. Franklin, and Oscar E. Tauber.
1935. On the haemolymph cell counts of
some marine invertebrates. Biol. Bull.
(Woods Hole) 69: 66-70.

Yonge, C. M.

1932. On the nature and permeability of
chitin. I. The chitin lining the foregut
of decapod Crustacea and the function
of the tegumental glands. Proc. Roy.
Soc., Ser. B, Biol. Sci. 111: 298-329..-

1936. On the nature and permeability of
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calcified chitin lining the foregut of

Homarus, Proc. Roy. Soc:, Sex. B;
Biol. Sci, 120: 15-41.
1937. The nature and significance of the

membranes surrounding the develop-
ing eggs of Homarus vulgaris and other
decapods. Proc. Zool. Soc. London,
Ser. A 107: 499-517.

1946. Permeability and properties of the
membranes surrounding the developing
egg of Homarus vulgaris. J. Mar. Biol.
Ass, U.K, 26: 432-438,

Young, John.
1879. On the head of the lobster.
Physiol., London 14: 348-350.

Z

J. Anat,

Zandee, D. I.
1964, Absence of sterol synthesis in some
arthropods, Nature (London) 202:
1335-1336.

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Review of Studies of Tuna Food Fees

in the Atlantic Ocean

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SPECIAL SCIENTIFIC REPORT--FISHERI|
EDITORIAL STAFF ae

Robert L, Hacker, Editor is

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Special Scientific Report--Fisheries are distributed free to libraries, research 5
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UNITED STATES DEPARTMENT OF THE INTERIOR

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Dayton L. Alverson, Acting Director, BUREAU OF COMMERCIAL FISHERIES

Review of Studies of Tuna Food
in the Atlantic Ocean

By

ALEXANDER DRAGOVICH

Contribution No, 117, Bureau of Commercial Fisheries
Tropical Atlantic Biological Laboratory
Miami, Florida 33149

United States Fish and Wildlife Service
Special Scientific Report--Fisheries 593

Washington, D.C,
December 1969

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CONTENTS

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Review of Studies of Tuna Food in the Atlantic Ocean

By

ALEXANDER DRAGOVICH, Fishery Biologist

Tropical Atlantic Biological Laboratory
Bureau of Commercial Fisheries
Miami, Florida 33149

ABSTRACT

Published and unpublished reports are reviewed and methods used to evaluate
the data are discussed. A description is presented of the food of seven Atlantic
tunas of commercial importance: little tuna (Euthynnus alletteratus), skipjack tuna
(Katsuwonus pelamis), yellowfin tuna (Thunnus albacares), blackfin tuna (T. atlanti-
cus), the bluefin tuna complex (T. thynnus thynnus and T, maccoyii), bigeye tuna
(I. obesus), and albacore (T, alalunga), Their food consists mainly of pelagic fish
(mostly juveniles, some larvae and adults), crustaceans (mostly macrozooplankton),
and mollusks (chiefly cephalopods), The greatest number offood items are fish taxa
(331), followed by crustaceans (111) and mollusks (74). Prey organisms are listed
alphabetically, according to the tuna species. The food consumed by the species of
tuna was generally similar. Differences in food betweenthe juvenile and adult blue-
fin tuna were pronounced; juveniles fed largely on crustaceans whereas adults fed
primarily on fishes. Seasonal differences were noted in the composition of the food

of skipjack and yellowfin tunas in African waters.

INTRODUCTION

A thorough knowledge of food and feeding of
tunas is important to an understanding of re-
gional and localaggregations of tunas andtheir
behavior (Reintjes and King, 1953; King and
Ikehara, 1956; Alverson, 1963b).

Interest in the food of Atlantic tunas was
evidenced as early as the second half of the
19th century (De Monaco, 1888). French re-
searchers have continued their interestintuna
food studies to the present time. Publications
in English began in the 1920's and are con-
tinuing. Contributions by Soviet, Spanish, Por-
tuguese, and Yugoslavian authors were pub-
lished in the 1960's.

The present paper reviews published and
unpublished reports on the food of tunas in the
Atlantic Ocean. Some of the papers consider
feeding and food in detail; others mentionthem
only incidentally. I do not summarize each
paper individually in this review but discuss
points of interest to a fishery biologist.

I divided the reports that were reviewed into
six categories (number of reports in paren-
theses): descriptive publications (9), notes (9),
reports on data (9), reviews (1), incidental
papers (28), andtaxonomic works (2). The cate-
gories are so designated in the bibliography.
Descriptive papers in general contain detailed

presentations and interpretations of results;
notes are largely brief reports based on lim-
ited observations; reports on data refer to
untreated data in tabular form; reviews contain
information mostly from other papers, which
is summarized and sometimes accompanied
by interpretation and comment; incidental
papers give information on food or feeding
habits in only a casual fashion (titles may be
completely unrelated to the food of tunas); and
taxonomic papers treat taxonomic aspects of
tuna forage in some detail.

The most important commercial species of
tunas dealt with in the papers reviewed were:
little tuna (Euthynnus alletteratus), skipjack
tuna (Katsuwonus pelamis), yellowfin tuna
(Thunnus albacares), blackfin tuna (T. atlanti-
cus), the bluefin tuna complex (T. thynnus thyn-
nus and T, maccoyii), bigeye tuna (T, obesus),
and albacore (T. alalunga). (The synonymy of
each species includes many more names than
those mentioned above.) Albacore received the
most attention in the papers.

The number of stomachs on which different
reports were based varied widely--from one
to several thousand.

Fishing gear used to capture tuna consisted
of trolling lines (with artificial lures), pole
and line, purse seine, longline, spears, rod
and reel, and harpoons.

METHODS USED FOR EVALUATION OF
STOMACH CONTENTS

It seems appropriate to review briefly the
various methods of evaluating food components
that have been used in studies of the stomach
contents of fishes, birds, mammals, and other
animals. The four most common methods are
discussed here and comments are made on
their advantages and disadvantages.

Numerical Method

The numerical method is based on counts of
the food items present; each item is evaluated
as a percentage of the total number of all food
items. The chief disadvantage in this method
is that the size of forage organisms is not
considered. Small organisms suchas megalopa
stages, for example, which occur in tuna
stomachs in large numbers, often rank higher
in apparent importance than do fishes or other
large forage organisms. The method places
undue emphasis on organisms with resistant
parts--crustaceans, shelled mollusks, and
scaled fishes--and thus may give a distorted
picture of food components. Also, itis difficult
and time-consuming to count accurately the
many fragmented organisms foundin stomachs,
and numerous macroplanktonic organisms must
be either counted or estimated from sub-
samples.

Percentage Frequency-of-Occurrence Method

The number of fish that have ingesteda par-
ticular food organism is calculated as a per-
centage of the total number of fish examined.
The method has the same disadvantages asthe
previous one: the number and size of organ-
isms are not considered, small organisms
usually outnumber large ones, and large or-
ganisms lose their importance. Like the pre-
ceding method, this procedure is primarily
useful in evaluating organisms within a simi-
lar size range. The method also allows rough
estimates of availability of foodorganisms and
selectivity by predators.

Volumetric, or Weight, Method

The displacement volume or the weight of
each food item is expressed as a percentage of
the total volume or weight of the stomach con-
tents. This method probably depicts most reli-
ably the relative importance of various food
items.

The volumetric method can be used in
various ways. Two of these--the ''aggregate-
total-volume" and ''average-percentage"' pro-
cedures--were described by Martin, Gensch,
and Brown (1946). In the first version, the
percentage of each kind of food is obtained by
dividing total food of each kind by total volume

of the stomach contents of all fish; the volume
of food from each stomach influences the final
result in direct proportion to that volume.
The method accurately reflects the volumetric
importance of a particular food organism
regardless of how many or how few other
Organisms are present. In the ''average-per-
centage'' version, equivalents are calculated
for each item of food, and each stomach is
taken as 100 percent, regardless of the volume
of its contents; variation in total volume of
food does not influence the results.

In his study of the food of the smallmouth
bass (Micropterus dolomieui), Tester (1932)
combined the volumetric andthe frequency-of-
occurrence methods for a graphic representa-
tion of the food items. Welsh (1949) used an
average percentage rating obtained by aver-
aging (1) the percentage of the total food vol-
ume contributed by the individual food item
(indicating food value), (2) the percentage of
total number of individual animals found in
stomachs (indicating abundance), and (3) the
percentage of the total stomachs in which
organisms were found (indicating availability).
When Welsh's method is used, the final figure
appears to be a combination of differentterms,
but it may serve as a simplified and useful
index.

"Points" Method

Points are assigned to food items, to the
total volumes, and to'the ''degree of fullness"
of each stomach. Assuming that the organ-
isms are of equal size (or allowing for size),
the investigator assigns the greatest number
of points to the most easily digested organ-
isms. The chief disadvantage in this method
is the difficulty of establishing or maintaining
standards on which to base designation of
points throughout extensive surveys.

Comments on Methods Used

In most reports classified in the present
paper as notes, reviews, incidental publica-
tions, and reports on data, evaluation of
stomach contents was qualitative; information
consisted of lists of forage organisms, ac-
companied by general comments. Examples
are: De Monaco, 1888; Chevreux, 1893; Grand-
besancon, 1910; Joubin and Roule, 1918a,
1918b; Le Danois, 1921; Nichols, 1922; De
Buen, 1927; Legendre, 1932, 1933; Long-
ley and Hildebrand, 1941; Priol, 1944; Le
Gall, 1949; Postel, 1950, 1954; Bigelow and
Schroeder, 1953; Rivas, 1954; Postel, 1955b;
Anderson, Gehringer, and Cohen, 1956a, 1956b;
Anderson and Gehringer, 1957a, 1957b, 1958a,
1958b, 1959a, 1959b, 1959c; Krumholz, 1959;
De Sylva and Rathjen, 1961; Klawe, 1961;
Alverson, 1963a; Postel, 1964; Zharov, Zhere-
benkov, Kadil'nikov, and Kuznetsov, 1964; Da
Cruz, 1965; Zharov, 1966; Bogdanov, Korzhova,

Kornilov, Leonova, Liubimova, Obvintsev,
Prosvirov, Sal'nikov, Terekhov, and Khromovy,
1967; Sokolov, 1967a, 1967b.

The numerical method (Legendre, 1934;
Bouxin and Legendre, 1936; Crane, 1936;
Legendre, 1940; Carlson, 1952; Morovic, 1961)
and the volumetric method (Postel, 1963;
Randall, 1967) were used in relatively few
studies.

In the more detailed papers, usually one of
the number methods (the numerical or the
percentage method) was combined with the
volumetric method.

In a study of blackfin and yellowfin tunas
near Bermuda and the West Indies, Beebe
(1936) used a number method in which he listed
the number of stomachs containing each par-
ticular food item--sometimes he listed each
food item in each fish by number and some-
times he used percentages in summations of
broader categories, such as fishes and crus-
taceans.

Various combinations of frequency-of-oc-
currence and volumetric, or weight, methods
have been used by several authors. Bane (1965)
evaluated the forage organisms of blackfin
tuna in the waters of Puerto Rico. Postel
(1955a) evaluated the food of Neothunnus alba-
cora (=Thunnus albacares) in the eastern
tropical Atlantic. De Jager, Nepgen, and Van
Wyk (1963) reported on food of tuna from the
west coast of South Africa. Oren, Ben-Tuvia,
and Gottlieb (1959) described food of T, thyn-
nus, E. alletteratus, and T. alalunga from the
eastern Mediterranean. Penrith (1963) studied
the food of several species of tuna off the
Cape of South Africa. Sund and Richards (1967)
showed the distribution of forage organisms in
the Gulf of Guinea for yellowfin and skipjack

tunas for two different seasons; and Suarez-
Caabro and Duarte-Bello (1961) evaluated the
forage organisms of skipjackandblackfintunas
in Cuban waters.

Soviet scientists (Zharov, 1965; Sokolov,
1967b) often applied the points method (using
an arbitrary scale) to measure the degree of
fullness of tuna stomachs.

RESULTS OF STUDIES

The information in the reports reviewed was
divided into several major topics: food organ-
isms; tunas as collectors of marine organ-
isms; tuna feeding habits; food in relation to
species and size of tunas; and seasonal and
diurnal variations in tuna food and feeding
habits.

Food Organisms

I examined the combined data inthe reviewed
papers from several points of view. It did not
appear that a quantitative treatment of the
data in terms of arealandtemporal distribution

would be particularly worthwhile, since many
of the original data were not quantitative.

Comparison of tuna prey in the stomachs of
fishes caught by various types of gear can be
of considerable value in studies of food selec-
tivity (Blackburn, in press), but data in the
reviewed papers were insufficient for such
a purpose. My compilation is therefore re-
stricted to a taxonomic list of the food items
and a record of their occurrence.

About 500 different forms were identified in
the tuna stomachs. Fishes were represented
by the greatest number of identified kinds (331),
crustaceans were second (111), and mollusks
were third (74). These numbers should not be
compared, since many organisms were identi-
fied only to genus or to family. The list of
identified taxa may reflect the rates of diges-
tion--the axial skeletons and scales of fishes
and the exoskeletons of crustaceans are less
subject to damage by digestive processes (and
therefore easier to identify) than are the
remains of fleshier organisms, Fewer speci-
mens of mollusks were identified than fishes
and crustaceans; most of the mollusks were
cephalopods, whose identifying external char-
acters are the first to be destroyed by the
digestive process. The list of identified taxa
also suggests that taxonomic work done on
fishes was more extensive than that on the
other two groups.

Although most of the studies were qualita-
tive, a general pattern of feeding was evident
for all the species of tuna. Taxonomically,
food consisted mainly of fishes (63 percent of
the total taxa), crustaceans (21 percent), and
mollusks (14 percent), Tunicates (2 percent)
were present but were of minor importance.
Tuna forage organisms (as identified in the
original publications) and the species of tunas
in which they were found are listed in the
Appendix. Fishes are listed alphabetically;
invertebrates also are listed alphabetically
but are grouped under broad categories.

Further considerations of the three main
categories of tuna food revealed that the fishes
eaten by tunas consisted chiefly of pelagic
juveniles and adults, but also included lar-
val forms and (occasionally) benthic fishes.
Crustaceans were largely macrozooplankton
(a great variety of larval decapods and stoma-
topods, and mostly adult hyperiid amphipods,
isopods, copepods, ostracods, euphausiids,
and mysids), and some micronekton (princi-
pally shrimp). Mollusks were mostly cepha-
lopods (principally squid), but heteropods and
pteropods were also prominent.

A practical aspect of studies of food organ-
isms is the definition of a relation betweenthe
areas of food abundance (or areas of high
biological productivity) and tuna abundance.
Although the food chain in the ocean has been
studied to some extent (mostly the relation
between primary and secondary producers
and large carnivores), data from the Atlantic

Ocean are scarce. The only reference to a
relation between areas of primary productivity
and abundance of tunas was found in recent
Soviet scientific publications. Soviet investi-
gators who linked the distribution of tuna with
areas of high biological productivity were:
Zharov et al., 1964; Zharov, 1965, 1966, 1967;
Bogdanov et al., 1967; and Sokolov, 1967a,
1967b.

Alverson's (1960, 1963b) studies have shown
that although the presence of anadequate stock
of tuna food in the eastern tropical Pacific
did not necessarily ensure the presence of
tunas, chances of finding the fish were far
greater in areas with abundant food than in
food-impoverished waters. Furthermore, ac-
cording to Blackburn (in press), the correlation
between occurrence of tunas and their prey in
the eastern tropical Pacific was closer than
that between tunas and the organisms (phyto-
plankton and zooplankton) on which tunas do
not prey.

The papers reviewed emphasize the need
for additional studies of tuna food and feeding
in the Atlantic. I believe this need for broader
study must be met through an accumulation of
data collected under contemporaneous long-
term programs that seek to measure zoo-
plankton, micronekton (tuna prey primarily),
large carnivores, and oceanographic condi-
tions.

Tunas as ''Collectors" of Marine Organisms

Tunas are often excellent collectors of
marine species. Numerous organisms de-
scribed in the literature for the first time
were taken from tuna stomachs. Collections
from the stomach contents of Germo alalunga

(=I. alalunga) provided the basis for descrip-
tions of the fauna of the Gulf of Gascony by
Bouxin and Legendre (1936) and Legendre
(1934, 1940). These authors examined 24,293
prey organisms and identified 106 taxa from
stomachs of T. alalunga caught in June, July,
August, and September over 4years (1929-32).
Totals were: invertebrates 68 taxa, 70 per-
cent of the total number of food organisms;
crustaceans 33 taxa, 61 percent; and cephalo-
pods 24 taxa, 9 percent. Of the crustaceans,
57 percent were amphipods. Brachyscellus
crusculum contributed 48 percent of the total
number of crustaceans, and Nematoscelis
megalops 24 percent. The 38 taxa of identified
fish made up 29 percent ofthe total organisms;
57 percent of the fish were Maurolicus muel-
leri.

Until Penrith's (1963) survey, Oreosoma
atlanticum and Tetragonurus cuvieri were
rare in museum collections; his Lestidium sp.
and Taractes sp. represented the first records
of these genera from South African waters; and
Centropholoides falcatus was known from the
type species only. The presence of these spe-
cies in tuna stomachs suggested to Penrith

that the fish might not be as rareas they were
thought to be, but that their apparent rarity
might be linked to existing inadequate methods
for catching midwater organisms.

Russell (1960), used stomach contents of
T. albacares to construct a taxonomic key to
North Atlantic heteropods.

Chevreux (1893) described certain hyperiid’
amphipods on the basis of stomach contents of
albacore caught between France andthe Azores.

Collections of juvenile tunas are valuable
aids in determining the location of thenursery
areas of different species. Klawe (1961) re-
ported juvenile T. atlanticus and Euthynnus
alletteratus, Scomberomorus sp. and Katsu-
wonus pelamis in stomachs of K. pelamis and
E, alletteratus in his studies of young scom-
brids from the waters between Cape Hatteras
and the Bahama Islands.

Feeding Habits

Most of the publications reviewed indicated
that tunas are indiscriminate pelagic feeders;
some of the authors surmised that the fish
also feed near the bottom. Beebe (1936) stated
that yellowfin and blackfin tunas feed near the
bottom at all times. Bane (1965) found that
goatfishes, squirrelfishes, triggerfishes, sur-
geonfishes, and jacks were the most numer-
ous fishes in the stomachs of blackfin tunas
collected near Puerto Rico; most of the food
fish were small (5-7 cm. long) and were com-
mon in nearby reefs and rocky areas. Bane
also found invertebrates in tuna stomachs,
among which were shrimp, squid, larval sto-
matopods, small octopi, lobster and crab
larvae, and gastropod shells. The presence of
benthic crustaceans and cephalopods in the
stomachs of bluefin tuna collectednear islands
in the Adriatic also suggests that tuna some-
times feed near the bottom (Morovi¢, 1961).
Da Cruz (1965) noted large numbers of small
Balistidae in the stomachs of blackfin tunas
and stated that blackfin tunas often must feed
near the bottom because small Balistidae are
usually found on coral bottoms.

Marchal (1959), who studied the food of
Neothunnus albacora (=T. albacares) in the
Gulf of Guinea, divided the forage organisms
into pelagic, bathypelagic, and coastal species.

Food in Relation to Species and Size of Tunas

In one of the more comprehensive studies of
food of several species of tuna, Penrith (1963)
showed important interspecific differences in
feeding habits of tunas off the Cape of Good
Hope, although he did not discuss food inrela-
tion to area or season of capture, or size of
tuna. He found that T, alalunga, T. albacares,
and T. obesus fed mainly on fish, cephalopods,
and shrimp. T. alalunga consumed a wide
variety of fishes andlarge quantities of macro-
plankton. T. albacares fed mainly on large
surface organisms, but took macroplankton

(megalopae) when it was abundant locally.
T. obesus fed mostly on large deep-sea forms.
None of the tuna species seemed to compete
with each other for food.

Zharov et al. (1964), who reviewed the tuna
fishery in the Atlantic Ocean, observed inter-
specific differences in feeding of tunas. Skip-
jack tuna fed on sardines, juvenile fish, squid
and other cephalopods, and small crustaceans;
yellowfin tuna fed on a variety of organisms,
from large plankton to fish; and little tuna fed
on relatively large fish, mainly sardine and
mackerel.

Intraspecific differences infood composition
were noted. In the Mediterranean Sea, food of
adult bluefin tuna differed from that of the
young. Young bluefin tuna ate mostly small
crustaceans (60 percent), fish (20 percent),
mollusks (19 percent), and tunicates (1 per-
cent); most of the fish were larvae and juvenile
sardines and anchovies (Oren, Ben- Tuvia, and
Gottlieb, 1959). Sokolov (1967b) reported that
adult bluefin tuna fed mainly on fishes of vari-
ous sizes and species (anchovy, bonito, mac-
kerel, and small skipjack tuna).

Seasonal and Diurnal Variations in Tuna Food
and Feeding

Information on the seasonal aspect of the
food habits of tunas is given in only a few
papers.

Sund and Richards (1967) found little geo-
graphical variation among tuna forage species
in the Gulf of Guinea, but a difference from
season to season. Anchoviella guineensis was
found only in stomachs of tunas taken during
the ''cool'' season. Trichiurus sp., unidentified
gonostomatids, Oxyporhamphus micropterus,
Atlanta sp., Dlex illecebrosus, and a species
of unidentified salp were present only in sam-
ples taken during the ''warm'' season. The
seasonal distribution of other species couldnot
be delineated because of the large differences
among samples.

Penrith (1963) reported almost no seasonal
variation in food taken by tuna off the Cape of
Good Hope--most species were consistently
present. Funchalia woodwardi was only occa-
sionally important to T. albacares and T. ala-
lunga, and the larvae of Jasus lalandii and
Plagusia chabrus were seasonally important
to T. alalunga and T. albacares, respectively.
Cephalopods were the only other important
food organism with pronounced changes in
seasonal availability.

Information on the diurnal intensity of feed-
ing is very scarce in the literature on Atlantic
tunas. Sokolov (1967b) stated that yellowfin
tuna fed actively in the morning, slackly at
midday, and intensely again in the evening.
Talbot and Penrith (1963) made similar ob-
servations. Suarez-Caabro and Duarte- Bello
(1961) observed no differences in the food
volumes between morning and afternoon.

LITERATURE CITED

(Papers reviewed are marked with asterisk in the literature cited; unmarked papers, which
are not concerned with tuna food in the Atlantic Ocean, were used as references but not

reviewed.)

ALVERSON, FRANKLIN G,

1960. Distribution of fishing effort and re-
sulting tuna catches from the Eastern
Tropical Pacific by quarters ofthe year,
1951-1958. Bull. Inter.-Amer. Trop.
Tuna Comm. 4: 319-446 (English and
Spanish).

*1963a. The food of yellowfin and skipjack
tunas in the Eastern Tropical Pacific
Ocean. Bull. Inter.-Amer. Trop. Tuna
Comm. 7: 295-396. (Incidental publi-
cation)

1963b. Distribution of fishing effort and
resulting tuna catches from the Eastern
Tropical Pacific Ocean, by quarters of
the year, 1959-1962. Bull. Inter.-Amer.
Trop. Tuna Comm, 8: 319-379 (English
and Spanish).

ANDERSON, WILLIAM W., and JACK W. GEH-
RINGER.

*1957a. Physical oceanographic, biological,
and chemical data, South Atlantic coast
of the United States, Theodore N. Gill
cruise 3. U.S. Fish Wildl. Serv., Spec.
Sci. Rep. Fish. 210, 208pp. (Reporton
data)

*1957b. Physical oceanographic, biological,
and chemical data, South Atlantic coast
of the United States, M/V Theodore N.
Gill cruise 4. U.S. Fish Wildl. Serv.,
Spec. Sci. Rep. Fish, 234, 192 pp. (Re-
port on data)

*1958a. Physical oceanographic, biological,
and chemical data, South Atlantic coast
of the United States, M/V Theodore N.
Gill cruise 5. U.S. Fish Wildl. Serv.,

Spec. Sci. Rep. Fish. 248, 220 pp.
(Report on data)

*1958b. Physical oceanographic, biological,
and chemical data, South Atlantic coast
of the United States, M/V Theodore N.
Gill cruise 6. U.S. Fish Wildl. Serv.,
Spec. Sci. Rep. Fish. 265, 99 pp. (Report
on data)

*1959a. Physical oceanographic, biological,
and chemical data, South Atlantic coast
of the United States, M/V Theodore N.
Gill cruise 7. U.S. Fish Wildl. Serv.,
Spec. Sci. Rep. Fish. 278, 277 pp. (Re-
port on data)

*1959b. Physical oceanographic, biological,
and chemical data, South Atlantic coast
of the United States, M/V Theodore N,
Gill cruise 8. U.S. Fish Wildl. Serv.,
Spec. Sci. Rep. Fish. 303, 227 pp.
(Report on data)

*1959c. Physical oceanographic, biological,
and chemical data, South Atlantic coast
of the United States, M/V Theodore N,
Gill cruise 9. U.S. Fish Wildl. Serv.,
Spec. Sci. Rep. Fish. 313, 226 pp.
(Report on data)

ANDERSON, WILLIAM W., JACK W. GEH-

RINGER, and EDWARD COHEN,

*1956a. Physical oceanographic, biological,
and chemical data, South Atlantic coast
of the United States, M/V Theodore N.
Gill cruise l. U.S. Fish Wildl. Serv.,
Spec. Sci. Rep. Fish. 178, 160 pp.
(Report on data)

*1956b. Physical oceanographic, biological,
and chemical data, South Atlantic coast of
the United States, Theodore N. Gill cruise
2. U.S. Fish Wildl. Serv., Spec. Sci.
Rep. Fish. 198,270 pp. (Report on data)

BANE, GILBERT W., JR.

1965. Notes on exploratory fishing and the
biology of the blackfin tuna around
Puerto Rico. Proc. Ninth Int. Game
Fish Conf., Runaway Bay, Jamaica,
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BEEBE, WILLIAM.

*1936. Food of the Bermuda and West Indian
tunas of the genera Parathunnus and
Neothunnus. Zoologica 21: 195-205.
(Descriptive publication)

BIGELOW, HENRY B., and WILLIAM C,

SCHROEDER.

*1953. Fishes of the Gulf of Maine. U.S.
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(Incidental publication)

BLACKBURN, MAURICE.

(In press). Micronekton of the Eastern
Tropical Pacific Ocean: family compo-
sition, distribution, abundance, and rela-
tionships totuna. U.S. Fish Wildl. Serv.,
Fish. Bull. 67.

BOGDANOV, D. V., IU. A. KORZHOVA, N. P.
KORNILOV, L. I, LEONOVA, T. G. LIUBI-
MOVA, A. L. OBVINTSEV, E. S. PROSVIROV,
N. E. SAL'NIKOV, E. S. TEREKHOV, and
N. S. KHROMOV. £

*1967. Meksikanskii zaliv (Spravochnoe po-
sobie dlia rabotnikov rybnoi promy-
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Nauchno-issled. Inst. Morsk. Rybn.
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BOUXIN, J., and R. LEGENDRE.

*1936, La faune pélagique de 1'Atlantique au
large du Golfe de Gascogne, recueillie
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ieme Partie: Céphalopodes. Ann. Inst.
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CARLSON, C. B.

*1952, Exploratory fishing for the little tuna
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(Incidental publication)

CHEVREUX, EDOUARD.

*1893. Quatrieme campagne de 1'Hirondelle
1888. Sur les crustacés amphipodes
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CRANE, JOCELYN.

*1936, Notes on the biology and ecology of
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DA CRUZ, JOAO FRANCISCO.

*1965, Contribuicao ao estudo da biologia
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Norte 2: 33-40. (Note)

DE BUEN, FERNANDO,

*1927. Notes et bibliographie sur la biologie
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DE JAGER, B. VAN D., C. S. de V. NEPGEN,
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*1963. A preliminary report on tuna in west
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DE MONACO, ALBERT (PRINCE).

*1888. Sur l'alimentation des naufagés en
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DE SYLVA, DONALD P., and WARREN F.
RATHI EN.

*1961, Life history notes on the little tuna,
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eastern United States. Bull. Mar. Sci.
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GRANDBESANCON, M.

*1910, La péches au thon dans le Golfe de Gas-
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JOUBIN, L., and L, ROULE.

*1918a. Observations sur la nourriture des
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*1918b. Note sur l'alimentation du thon de
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KING, JOSEPH E., and ISAAC I, IKEHARA,
1956. Comparative study of food of bigeye
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KLAWE, W. L.

*1961. Young scombroids from the wa-
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cation)

KRUMHOLZ, LOUIS A,

*1959. Stomach contents and organ weights
of some bluefin tuna, Thunnus thyn-
nus (Linnaeus), near Bimini, Bahamas.
Zoologica 44: 127-131. (Note)

LE DANOIS, ED.

*1921. Recherches sur le regime des eaux
Atlantiques au large des cotes de France
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mon. (Observations faites pendant la
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(Incidental publication)

LE GALL, JEAN.

*1949, Résumé des connaissances acquises
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LEGENDRE, R.

*1932. La nourriture du germon Germo ala-
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*1933. Présence d'Anotopterus pharao Zug-
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LONGLEY, WILLIAM H., and SAMUEL F.,
HILDEBRAND.

*1941. Systematic catalogue of the fishes of
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Carnegie Inst. Wash. Publ. 535, 331 pp.
(Incidental publication)

MARCHAL, E.

*1959. Analyse de quelques contenus stoma-
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MARTIN, A. C., R. H. GENSCH, “and C. P.
BROWN.

1946, Alternative methods in upland game
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MOROVIC, D.

*1961. Contribution a la connaissance de la
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NICHOLS, J. T.

*1922, Color of the tuna. Copeialll: 74-75.
(Incidental publication)

OREN, O. H., A. BEN- TUVIA, and E. GOTT-
LIEB.

*1959, Experimental tuna fishing cruise in
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PENRITH, M. J.

1963, The systematics and biology of the
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216 pp. (Descriptive report)

POSTEL, E.

*1950. Rapport et note sur quelques poissons
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*1955b. La bonite a ventre raye (Katsuwonus
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*1963. Exposé synoptique sur la biologie du
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*1964. Sur deux lots de germons
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IPIRULOIL I =

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RANDALL, JOHN E.

*1967. Food habits of reef fishes of the
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REINTJES, JOHN W., and JOSEPH E. KING.
1953. Food of yellowfin tuna in the Central
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RIVAS, LUIS RENE.

*1954, A preliminary report on the spawning
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RUSSELL, HENRY D.

*1960. Heteropods
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SOKOLOV, V. A.

Fi967ascGolubei tunets Atlanticheskogo
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SUAREZ-CAABRO, JOSE A., and PEDRO
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*1963. Synopsis of biological data on species
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TESTER, A. L.

1932. Food of the small-mouthed black bass
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1949. A preliminary study of food and feed-
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ZHAROV, V. L.

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*1966. Zavisimost! raspredeleniia skoplenii
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ZHAROV, V., IU. ZHEREBENKOV, IU. KADIL'-
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MS. # 1898

_ Ee a

Appendix Table 1.--Food found in stomachs of seven species of tuna from the Atlantic Ocean

FOOD ITEM Euthynnus
alletteratus

Katsuwonus
pelamis

SPECIES OF TUNA

Thunnus
albacares

Thunnus
atlanticus

Thunnus
thynnus

Thunnus
gbesus

Thunnus.
alalunga

FISHES

Abalistes stellaris (Bloch and
Schneider)

Ablennes hians (Valenciennes)

Acanthurus sp.

Acanthurus chirurgus (Bloch)

Acanthurus coeruleus Bloch and
Schneider

Acanthurus hepatus (Linnaeus)
Acanthurus monroviae Steindachner
Alepisaurus sp.

Alepisaurus ferox Lowe

Allaneta harringtonensis (Goode)

Alutera sp.

Alutera monoceros (Linnaeus)
Alutera scripta (Osbeck)
Alutera heudelotii Hollard

Ammodytes sp.

Anchoa cubana (Poey)

Anchoa sp.

Anchoviella sp.

Anchoviella guineensis (Rossignol
and Blache)

Anoplogaster cornutus (Cuvier

and Valenciennes)

Anotopterus pharao Zugmayer
Antennarius sp.

Anthias sacer Lowe

Antigonia sp.

Antigonia combatia Berry and Rathjen

Aphanopus sp.

Argentina sp.

Argypropelecus sp.

Argypropelecus aculeatus Cuvier
and Valenciennes

Argypropelecus olfersi (Cuvier)

Ariomma ledanoisi (Belloc)

Arnoglossus sp.

Arnoglossus imperialis Rafinesque
Atherinidae

Atherinomorus stipes (Miller and
Troschel )

Aulopus sp.

Auxis sp.

Auxis thazard (Lacépéede)
Avocettina infans (Giinther)
Balistidae

Appendix Table 1.--Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

FOOD ITEM Euthynnus
alletteratus
FISHES

Balistes sp.

Balistes forcipatus Gmelin
Bathylagus microcephalus Norman
Barathronus parfaiti Vaillant
Belonidae

Belone belone (Linnaeus)

Benthodesmus atlanticus Goode and
Bean

Berycoidea

Blennius ocellaris Linnaeus

Boops vulgaris Bowdich

Bothidae

Box hoops Vinciguerra
Brama sp.

Brama rayi (Bloch)
Bramidae

Brevoortia tyrannus (Latrobe)
Brotulidae

Cantherines pullus (Ranzani)
Canthidermis sufflamen (Mitchill)
Canthigaster rostratus (Bloch)

Capros aper (Linnaeus)
Carangidae

Caranx sp.
Caranx bartholomaei Cuvier
Caranx crysos (Mitchill)

Caranx hippos (Linnaeus)

Caranx latus Agassiz

Caranmx rhonchusGeoffroy St.-Hilaire
Caranx ruber (Bloch)

Caranx trachurus Cuvier

Centropholoides falcatus (Barnard)

Ceratioidei

Ceratoscopelus townsendi (Eigenmann
and Eigenmann)

Chaetodontidae

Chaetodon marleyi Regan

Chaetodon sedentarius Poey
Chaetodon striatus Linnaeus
Champsodon sp.
Chauliodus sloani Schneider
Chiasmodontidae

Chlorophthalmus agassizi Bonaparte
Chlorophthalmus atlanticus Poll
Chloroscombrus sp.

Clinidae

Clupea finta Cuvier

Katsuwonus.
pelamis

SPECIES OF TUNA

Thunnus

albacares

Thunnus
atlanticus

Thunnus
thynnus

Thunnus
obesus

Thunnus
alalunga

Appendix Table 1.--Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

SPECIES OF TUNA

FOOD ITEM Euthynnus Katsuwonus Thunnus Thunnus Thunnus Thunnus Thunnus
alletteratus pelamis albacares atlanticus thynnus obesus alalunga

FISHES

Clupea sprattus Poggi sc -- -- xX -- -- 35

Clupeidae x -- -- a a ss jee

Collybus sp. <= -- -- =o = x

Conger conger (Linnaeus) -- -- -- a5 o> So x

Conger vulgaris (leptocephala) -- -- -- -- =< x
Gtinther

Congermuranea impressa (Poey) xX =o ==
Coryphaena hippurus Linnaeus xX = x
Cubiceps gracilis (Lowe) =e oD x oS So oO
Cyclichthys orbicularis Kaup == 2S xX

x

Cypselurus sp. x oo

Cypselurus furcatus (Mitchill) =- -- -- -- == -- x
Cypselurus heterurus (Rafinesque) aS xX -- -- -- = =
Cypselurus lineatus (Valenciennes) -- -- a5 = 2S as x
Dactyloptena orientalis (Cuvier) -- -- -- -- ae 5A 4
Dactylopteridae -- -- xX == os ae x

Dactylopterus volitans (Linnaeus) -- xX xX

Decapterus macarellus (Cuvier) == oD xX

Decapterus punctatus (Agassiz) x x x

Decapterus ronchus (Geoffroy x OF =o
St.-Hilaire)

Diagramma mediterraneum (Guichenot) -- B= aD == —— ze x

Diaphus sp. -- -- -- = xX x
Diaphus effulgens (Goode and Bean) ae -- -- xX on

Diaphus gemellarii (Cocco) -- -- -- on
Diaphus liitkeni (Brauer) -- -- -- aS mc ae
x

Diaphus rafinesquii (Cocco) er -- So

'
'
'
'

1 OPS PS PS PS

Diaphus theta Eigenmann and -- -- -- = =o Ais x
Eigenmann

Diodon sp. or x xX x -- -- as

Diodon holacanthus Linnaeus om xX x -- -- 2

Diodon hystrix Linnaeus aS -- -- -- X

Diplodus sargus (Linnaeus) So -- -- ao =

Diretmus argenteus Johnson =
Engraulidae

x
Engraulis sp. x == X -- -- -- os
Engraulis encrasicholus (Linnaeus) x Xx

Engraulis hepsetus Linnaeus oo x oo oS OF ae oo

Engraulis japonicus (Hottuyn) aS a5 xX ro xX xX os

Entelurus aeguoreus Linnaeus =o =o a ao ao oO ».¢

Epinnula orientalis Gilchrist and == a co xX a2 =O oR
Von Bonde

Etrumeus teres (De Kay) x x == xX x — ==

Eucinostomus pseudogula Poey =O x xX
Euthynnus alletteratus (Rafinesque) xX ee x 2s ao ie x
Exocoetidae x xk x
Exocoetus sp. »4 om oa oS RO a a
Exonautes rubescens (Rafinesque) oS = es x oD 2S oe

11

Appendix Table 1.--Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

SPECIES OF TUNA

FOOD ITEM Euthynnus Katsuwonus _Thunnus Thunnus Thunnus Thunnus Thunnus
alletteratus pelamis albacares atlanticus thynnus obesus alalunga

FISHES

Fistularia serrata Cuvier == -- -- xX -- = --
Fistularia tabacaria Linnaeus xX == -- xX -- -- ==
Fistularia villosa Klunzinger =- -- X

Fodiator acutus (Valenciennes) xX == X -- -- a ==

Gadidae x me == co co = --
Galeoides polydactylus (Vahl) ».¢ oS
Gempylidae me xX
Gempylus serpens Cuvier oR xX
Gephyroberyx darwini (Johnson) a xX

Gerridae -- x
Gerres cinereus (Walbaum) -- xX --
Gonorynchus gonorynchus (Linnaeus) -- -- xX
Gonostoma sp. ae -- x -- -- -- ==
Gonostomatidae 2 ae xX

Haemulon flavolineatum (Desmarest) xX -- -- -- == =a =2

Halieutea fitzsimonsi (Gilchrist -- -- -- -- 48 nin x
and Thompson)

Harengula sp. -- -- -- == xX ao ae

Helicolenus dactylopterus (De la -- -- am oo — aS) Xx
Roche )

Helicolenus maculatus Cuvier oS -- -- -- X xX xX

Helicolenus porcus (Linnaeus) >
Hemipteronotus sp. is
Hemipteronotus noracula (Linnaeus) xX == oS aS =O —— ==
Hemiramphidae x
xX

Hemiramphus sp.

Hemiramphus balao LeSueur -- xX -- — =
Heterosomata larvae se -- -- xX

Hippocampus sp. B¢ -- x -- xX
Hippocampus brevirostris Valenciennes -- =o -- -- x
Hippocampus erectus Perry =e -- -- ye

Holocentridae D>
Holocentrus Gronow SO
Holocentrus ascensionis (Osbeck)
Holocentrus rufus (Walbaum)

Holocentrus vexillarius(Poey) X -- -- xX -- = cD

Hyporhamphus sp. xX xX -- -- -- -- xX
Hyporhamphus unifasciatus (Ranzani) == co xX -- --

Jenkinsia sp. xX << --
Katsuwonus pelamis (Linnaeus) xX

Lactophrys sp. -- -- --

Lagocephalus sp. -- -- -- --

Lagocephalus laevigatus (Linnaeus) -- -- xX -- --
Lampadena chavesii Collett -- -- -- ==

xX x
Lampanyctodes hectoris Giinther -- -- xX -- x x xX
Lampanyctus sp. -- -- -- -- xX Xe

Appendix Table 1.--Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

FOOD ITEM Euthynnus
alletteratus
FISHES

Lampanyctus alatus (postlarva)
Goode and Bean
--now
Lampanyctus pusillus (Johnson)
Lampanyctus crocodilus (Risso)
Lampanyctus intricarius Taaning
Lampanyctus maderensis (Lowe)
Lampanyctus margaritiferus (Goode
and Bean)

Lamputa umgazi Smith
Laptostomais sp.

Lepidopus sp.

Lepidopus caudatus (Euphrasen)

Lepidotrigla sp.

Leptocephalus (Anguilliformes-larvae)X

Lestidium sp.

Lichia glauca (Linnaeus)
(Probably: Trachinotus glauca
(Linnaeus ))

Liosaccus cutaneus (Giinther)

Lophiidae

Maurolicus sp.

Maurolicus muelleri (Gmelin)
Melanostomiatidae

Merluccius bilinearis (Mitchill)
Merluccius capensis Gastlenau
Merluccius merluccius (Linnaeus)
Micropteryx chrysurus (Linnaeus)

(Chloroscombrus chrysurus)
Molidae

Monacanthus sp.

Monacanthus ciliatus (Mitchill)
Monacanthus hispidus (Linnaeus)
Monacanthus tuckeri Bean
Mullidae

Mulloidichthys martinicus (Cuvier)
Mullus barbatus Linnaeus
Myctophidae

Myctophum coccoi (Cocco)
Myctophum sp.

Myctophum humboldti (Risso)
Myctophum hygomii (Ltitken)
Myctophum punctatum Rafinesque
Myctophum tisso (Cocco)

Naucrates ductor (Linnaeus)

Katsuwonus

pelamis

13

SPECIES OF TUNA

Thunnus
albacares

Thunnus
atlanticus

Thunnus
thynnus

Thunnus
obesus

Thunnus
alalunga

ba a

Appendix Table 1.~-Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

SPECIES OF TUNA

FOOD ITEM Euthynnus Katsuwonus Thunnus Thunnus Thunnus Thunnus Thunnus
alletteratus pelamis albacares atlanticus thynnus obesus alalunga
FISHES
Nemichthys scolopaceus Richardson -- -- -- -- -- -- x
Nesiarchus nasutus Johnson -- -- -- xX -- — a
Nesiarchus sp. -- -- x -- om ~= a>
Notolepis rissoi kroyers (Ltitken) -- -- -- — oD at x
Ogcocephalidae -- -- -- X -- ot md
Oligoplites saurus (Bloch and -- x -- -- a ot as
Schneider)
Omosudis lowii Gtnther -- -- -- x = a0 x
Onos mediterraneus (Linnaeus) -- -- -- -- = oo X
Onos vulgaris Yarrel -- -- -- -- -- -- X
Ophidiidae -- -- -- xX oD -- —
Ophidion barbatum Linnaeus -- -- x -- =o =< at
Ophidion vassali Risso x -- -- st x =5 X
Oreosoma atlanticum Cuvier and -- -- x. -- X X X
Valenciennes
Ostracion sp. -- -- X -- ary a5 X
Ostracion tuberculatus Linnaeus -- -- x -- -- “= =5
Oxyporhamphus sp. -- -- X -- a5 oO a5
Oxyporhamphus micropterus similis a3 xX xX -- -- -- ==
Bruun
Otophidium omostigmum (Jordan and x -- -- -- -- aS es
Gilbert)
Pagellus sp. x -- -- -- -- == =s
Paralepis sp. x -- xX 5'¢ i X -- X
Paralepis coregonoides Risso ae pe = =o = oS Xx
Paralepis coregonoides borealis -- == ao, «= es == X
Reinhardt
Paralepis pseudosphyraenoides Ege -- -- -- -- a a0 xX
Paralepis spesiosus Bellotti ae me == se X = xX
Paralepis sphyraenoides Risso -- =< == cee a5 a0 X
Paranthias furcifer (Valenciennes) = os a xX aS os =
Peprilus alepidotus (Linnaeus) X mr a5 == a a a5
Photichthys argenteus Hutton = a= = as ae = Xx
Plagyodus alepisaurus Lowe -- =e == == = =< X
Planctanthias praeopercularis Fowler -. -- aes == oD os X
Pleuronectoidea X os =o ao —— = =c
Polydactylus virginicus (Linnaeus) =a x Se x oS == SO
Polyipnus spinosus Gtinther om == x =e x X X
Pomadasys sp. Xx ee — = oo OS —
Priacanthidae -- X —— a oo =e =
Priacanthus sp. -- -- X -- == = X
Priacanthus cruentatus (Lacepede) a = x i oH oD co
Priacanthus hamrur Forskal -- == do oS oD oc X
Prionotus sp. x = a om co -- --
Pristopomatides sp. -- -- x == == -- --
Prognichthys gibbifrons (Valenciennes) -- x -- -- -- -- --

Appendix Table 1.--Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

SPECIES OF TUNA

FOOD ITEh Euthynnus Katsuwonus Thunnus Thunnus Thunnus Thunnus Thunnus
alletteratus, pelamis albacares atlanticus thynnus obesus alalunga

FISHES

Psenes sp. -- -- x -- on — X
Psenes cyanophrys Cuvier == -- xX -- == os ae
Pseudopentaceros richardsoni (Smith) -- -- -- == aS oe x

Pseudopriacanthus altus (Gill) xX -- xX =< = = be
Pseudupeneus maculatus (Bloch) x -- mS = a el =

Pseudupeneus prayensis (Cuvier) -- -- xX -- => as ei)
Pteraclidae -- -- =- = =~ a x

Pteraclis sp. -- --

xX
Pterycombus goodei (Jordan) -- x xX -- = ae a
Rhomboplites aurorubens (Cuvier) -- -- -- xX = ts aS

Sardina pilchardus (Walbaum) -- -- -- -- xX mo a

Sardinella sp. xX X

Sardinella anchovia Valenciennes xX -- xX x os = =

Sardinella aurita Valenciennes xX xX

Sardinella eba (Cuvier and -- -- == ze ae ae
Valenciennes)

Sardinella rouxi Whitehead -- == xX

Sardinops ocellata (Pappe) -- -- xX -- X x: xX
Sargus sp. xX

Saurida parri Norman xX -- -- -- -- -- --

Schedophilus enigmaticus Gtinther -- -- -- Sc Do od

Schedophilus medusophagus Cocco -- -- fare 5 ay ey x
Scomber sp. xX

Scomber japonicus Houttuyn x x xX a x
Scomberesox saurus (Walbaum) -- oo xX X
Scomberomorus maculatus (Mitchill) X == St zs ct: eer

Scombridae X BS x Nats ee 2
Selene vomer (Linnaeus) -- =5 o>
Selar crumenophthalmus (Bloch) 54 of x
Serranidae = x Xx
Smaris sp. xX -- -- -- se oe

Soleidae == -- -- -- -- an x

Sparisoma flavescens (Bloch and 6 -- -- -- ed ae
Schneider)

Sphaeroides sp. -- -- -- x ees oS re:

Sphaeroides spengleri (Bloch) -- -- X -- ao aS ir

Sphyraena sp. XK -- x -- =0 aS x

Sphyraena barracuda (Walbaum) -- x = me ae pF:
Spondyliosoma cantharus (Linnaeus) -- -- oc OD a ee

x
Sternoptyx diaphana Herman -- xX -- = ais < x
Stomiatidae -- -- — os ee os x

Strongylura sp. -- -- -- X pe B.. aps

Strongylura marina = -- x == xX oo oe
(dalbaum)

Strongylura timueu(Walbaum) x -- -- co == os oo

Sudis sp. -- -- == ae a5 So

xX
Synagrops microlepis Norman =O == X -- -- -- X
Syngnathidae =< Xx X a oo oD OS

15

Appendix Table 1.--Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

SPECIES OF TUNA

FOOD ITEM Euthynnus Katsuwonus Thunnus Thunnus Thunnus Thunnus Thunnus
alletteratus pelamis albacares atlanticus thynnus obesus alalunga
FISHES

Syngnathus sp. — a= xX x xX -- a4
Syngnathus dunckeri Metzelaar a == -- x: -- -- ==
Syngnathus springeri Herald Xx. -- -- -- -- oe =-
Synodontidae ie -- X -- -- =o ~-
Synodus sp. 7S xX xX -- -- -- -=
Synodus synodus (Linnaeus) -- -- -- -- <= =6 x
Taractes sp. oS -- x -- -- mee xX
Tetragonurus atlanticus Lowe -- -- -- -- == = x
Tetragonurus cuvieri Risso as -- -- -- -- — x
Tetraodontidae -- -- x xX -- oy x
Therapon sp. -- -- -- -- oo 2 x
Thunnus atlanticus (Lesson) mG xX -- -- -- = —
Thyrsites atun (Euphrasen) ae x: -- -- -- a5 x
Trachurus sp. -- -- -- =< =o so x
Trachurus trachurus (Linnaeus) aS AS xX oc xX Sa xX
Trachypterus iris (Walbaum) -- -- = on == Zs x
Trachurus trachurus et -- xX == x x xX

(Linnaeus )
Trichiurus sp. -- x ve = = =e ==
Trichiurus lepturus Linnaeus == xX -- -- -- — De
Trigla gurnardus Linnaeus -- -- a5 =e . —— oe X
Triglidae (Trigla sp.) -- Bo a == =k ae xX
Tripterodon sp. -- -- a os co x xX
Tylosurus acus (Lacépede) X aS Sci au = oa =a
Tylosurus crocodilus Linnaeus x -- — == an = =
Uranoscopus sp. -- -- x — se aH a3
Valenciennellus tripunctulatus -- -- x — oc ae =

(Esmark)
Vomer setapinnis (Mitchill) -- -- x = = on X
Vinciguerria sp. -- == a = = ute x
Vinciguerria sanzoi Jespersen and -- -- -- as me eis X

Taaning
Xanthichthys ringens (Linnaeus) -- -- xX X = = —
Xiphasia setifer Swainson -- -- X os = ae =5
Yozia bicoarctata (Bleeker) -- -- x = “a -: wa
Zeoidei = — aes == anges shy x
Zeus sp. -- -- xX co a 5 =s

INVERTEBRATES

OSTRACODA :

Conchoecia sp. me -- -- -- a = xX

Ostracoda (not further identified) -- -- -- xX a == =5
CEPEPODA :

Calanus finmarchicus (Gunner) = -- -- 5 oD ES x

Copepoda (not further identified) X € -- == me = a

Penella exocoeti (Holten) se -- xX -- a o- =
CIRRIPEDIA:

Lepas anatifera Linnaeus - -- -- -- BS oS xX

16

Appendix Table 1.--Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

FOOD ITEM

INVERTEBRATES

MYSIDACAEA :
Gnathophausia ingens (Dohrn)

ISOPODA :

SPECIES OF TUNA

Euthynnus

alletteratus

Isopoda (not further identified) oC

Idotea metallica Bosc

AMPHIPODA:
Anchylomera blossevillei
H. Milne Edwards

Amphipoda (not further identified) --

Brachyscelus sp.
Brachyscelus crusculum Bate
Cystisoma sp.

Euthemisto sp.

Euthemisto bispinosa (Boeck)
(Syn. of Parathemisto
guadichaudii (Guerin) )

Euprimno macropaus (Guerin)
--now
Primno macropa (Guerin)

Gammarus sp.

Hyperiidae

Hyperioides longipes (Chevreux)

Hyperia galba (Montegu)
Lanceola sayana Bovallius
Oxycephalus sp.

Parapronoé crustulum Claus

Paraphronima crassipes (Claus)

Parathemisto obliva (Kroyer)
probably Parathemisto
gracilipes (Norman)

Phronima sp.

Phronima atlantica Guérin

Phronima sedentaria (Forskal)

Phronima stebbingii (Vosseler)

Phrosina semilunata Risso
Platyscelus armatus (Claus)
Platyscelus ovoides (Risso)
Platyscelus gerratulus Stebbing
Streetsia sp.

Streetsia challengeri Stebbing
Streetsia pronoides (Bovallius)

STOMATOPODA :

Gonodactylus sp.

Lysiosquilla sp. (larvae)

Stomatopoda (not further
identified)

Squillidae (various types of
larvae)

Squilla sp.

Katsuwonus

pelamis

Thunnus
albacares atlanticus

Thunnus

Thunnus
thynnus

Thunnus
obesus

Thunnus
alalunga

PSPS PS

Pa PS

PS PS Pd PS PS Pd oP

Appendix Table 1.7-Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

SPECIES OF TUNA

FOOD ITEM Katsuwonus

pelamis

Thunnus Thunnus Thunnus Thunnus

Euthynnus AM}
albacares atlanticus thynnus obesus

alletteratus

INVERTEBRATES

EUPHAUSIACEA :
Euphausiacea (not further =F xX Ȣ xX x =n
identified)

Euphausia sp. == -- x -- — ot
Euphausia lucens Hansen == -- xX

Meganyctiphanes norvegica (M. Sars) -- -- -- -- ot 55
Nematoscelis megalops G. O. Sars -- -- -- ac — =
Nematoscelis sp. xX -- -- -= X =a
Nyctiphanes sp. Oe xX -- -- -- =2
Nyctiphanes capensis Hansen a= -- x -- ae a5

Nyctiphanes couchii (Bell) == -- xX -- 55 =s
Stylocheiron abbreviatum G.0. Sars -- -- -- -- = S23
Thysanoessa sp. xX -- -- De x rien
Thysanopoda xX = ois — Xx au

DECAPODA -CRUSTACEA:
Decapoda (not further identified) xX Ȣ xX X == 23

PENAEIDAE:
Aristaeomorpha foliacea (Risso) -- -- -- -- == =
Cerataspis sp. (larvae) -- -- x -- s/ 0 Ss “a
Cerataspis monstrosa Gray =o 25 Xx x
Funchalia villosa (Bouvier) -- -- -- xX — i
Mysis stages

Funchalia woodwardi Johnson -- -- xX -- xX xX

Gennadas (Amalopenaeus) elegans -- -- -- -- =o a
S. I. Smith

Parapenaeus longirostris (Lucas)

Penaeus duorarum Burkenroad xX -- -- -- -- =
Penaeidae (not further identified)

SERGESTIDAE:

Sergestes sp. -- -- = oo za mid
Sergestes arcticus Krtyer -- -- == _— Ars So
Sergestes gloriosus Stebbing -- -- —— aS ee, at
Sergestes phorcus Faxon -- -- = Se AS: sed
Sergestes robustus Smith -- =- as = Te ee

Sergestes splendens Sund -- --

CARIDEA:

Acanthephyra sp. -- -- De = 25 =

Acanthephyra multispina Coutiere -- -- == — oo =e
Syn. of A. pelagica (Risso)

Alpheidae (Diaphorus-larvae) -- == oo oe nS =

Alpheus ruber (larvae Anebocaris) -- -- -- == a --
H. Milne Edwards

Brachycarpus biunguiculatus (Lucas) X -- -- -- xX --
Enoplometopus dentatus Miers -- -- == os 2s ae

Glyphocrangon sp. -- -- oe oo Se oS

Heterocarpus ensifer A. Milne -- -- x = Se eS
Hippolytidae Edwards) __

Thunnus
alalunga

PM

ms

1

Appendix Table 1.--Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

SPECIES OF TUNA

FOOD ITEM Euthynnus Katsuwonus Thunnus Thunnus Thunnus Thunnus Thunnus
alletteratus pelamis albacares atlanticus thynnus obesus alalunga

INVERTEBRATES

Icotopus amphissimus Coutiete xX -- -- = x S3

Leptochela sp. == x
Palaemonidae -- xX == — a2 as ==

Palaemonella sp. oS -- xX X == ae a
Parapasiphae sulcatifrons Smith -- -- -- == BS ors

Pasiphae sp. (?) -- -- == — s a x

Systellaspis debilis -- -- —— ae = <s x
A. Milne Edwards

MACRURA-REPTANTIA:

Axius stirhynchus Leach =5 => w= -- -- -- xX
Hippa cubensis (Saussure) =< aD -- xX -- -- --
Jasus lalandii (A. Milne Edwards) -- =- xX -- xX xX xX
Jasus parkeri Stebbing - Syn. of a cm -- -- om =o X
Projasus parkeri (Stebbing)
Nephrops andamanica (?) == == -- oo = = X
Wood=-Mason
Palinuridae -- »:¢ -- -- oo ats eo
Palinurus sp. == xX -- os = a X
Palinurus regius Brito Capello = os - xX =~ ie xX
Palinurus vulgaris (Phyllosoma) X -- -- a xX sei x
Latreille
Panulirus sp. <n =o -- X on = oo
Phyllosoma -- == x a i as ero
Scyllarides sp. (nisto stage) -- -- -- a5 ae ae x
Scyllarus arctus (Linnaeus) xX om -- == xX =— x
Paguridae (Glaucothée) -- xX xX xX oS “od ——
Pagurus sp. -- -- xX ac oo ae a
Stenopus hispidus (Olivier) -- -- -- x CS uo oo
BRACHYURA :
Brachyrhyncha -- xX -- om oo oo =
Brachyrhyncha-megalopa -- -- x x xX x x
Megalopa (Portunidae and Dromiidae)-- -- xX aes os ue 35
Megalopa a x x x xi x x
Oxyrhyncha -- xX -- xX os ot oD
Plagusia chabrus (Linnaeus) -- -- XK ac ae ai x
Portunas sp. = os -- on x ee oo
Zoea -- xX xX — = “4 X
OCTOPODA:
Allopsus mollis Verrill = oS =e x <5 = CO
Argonauta nodosa Solander a 25 x ms x x x
Argonauta sp. —— Dae X | x x x
Bathypolypus sponsalis pane sie x a us ae nin
P. and H. Fischer
Bolliattaenella (Japetella) a = oS ac = as X
diaphana (Hoyle)
Eledone cirrhosa (Lamarck) me be a5 os Os = xX
Eledone moschata (Lamarck) -- -- -- w= x oo =a
Nautilus sp. -- xX a5 one co = 2c
Octopidae a X a x on = oa

19

Appendix Table 1.--Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

SPECIES OF TUNA

FOOD ITEM Euthynnus Katsuwonus Thunnus Thunnus Thunnus Thunnus Thunnus
alletteratus pelamis albacares atlanticus thynnus obesus alalunga
INVERTEBRATES
Octopus sp. mM Xx a Xx X -- --
Octopus burryi Voss =< x -- X -- = at
Octopus vulgaris Lamarck eS -—- Xx -- -- -- xX
Octopus defilippi Verany co == X -- -- -- ==
Ocythoe tuberculata Rafinesque oc == =- -- -- -- x
Todarodes sagittatus (Lamarck) -- -- xX -- -- -- ven
Tremoctopus violaceus Delle Chiaja -- 2S -- -- -- -- X
Vitreledonella sp. (?) == == -- -- -- -- xX
TEUTHOIDEA :

Abralia gilchristi Robson ob om xX --
Abralia veranyi (Ruppell) == x x -- = oo =
Allotheuthis africana Adam oO -- X -- = = =<
Brachioteuthis (Tracheloteuthis) -- -- -- =o oS a x

riisei (Streenstrup)
Calliteuthis reversa (Verrill) oe -- -- -- = os x
Chiroteuthis veranyi (ferussac) -- -- -- == on = x
Chranchia scabra Leach oS == x TS oo te x x
Ctenopteryx siculus Verany 3 ae a -- -- -- x
Desmoteuthis hyperborea == =o ac os -- -- X

(Steenstrup)
Doryteuthis sp. xX -- -- -- oc =5 a5
Doryteuthis plei (Blainville) oo -- xX -- -- os ——
Galiteuthis armata Joubin -- -- -- -- os =o Xx
Gonatus fabricii (Lichtenstein) -- -- -- -- oo mo x
Heteroteuthis dispar (Ruppell) == -- -- -- -- == xX
Histioteuthis bonelliana (Ferussac )-- -- -- -- == oe xX
Illex coindeti (Verany) -- -- -- = = 2s x
Illex illecebrosus coindetti == x xX -- —— a =

(Verany)
Liocranchia reinhardti (Steenstrup)-- -- -- as oS ot
Loligo sp. x: -- x -- xX so so
Loligo pealei LeSueur -- xX -- co oo =o a5
Loligo reynaudii d'Orbigny -- -- x -- xX x x
Loligo vulgaris Lamarck mo -- -- -- X BO =:
Lolliguncula brevis (Blainville) -- x -- X =o oo ae
Lolliguncula mercatoris Adam == -- xX -- -- = =a
Mastigoteuthis (?) sp. -- -- == =e = ce x
Octopodoteuthis sicula (Ruppell) -- -- x -- xX xX ae
Ommastrephidae -- -- -- a a —— xX
Ommastrephes pteropus Steenstrup a= x xX -- -- -- oO
Ommastrephes sagittatus (Lamarck) -- -- -- == oo ee x
Onychoteuthis banksii (Leach) -- -- xX oD sa os x
Onykia appellofii Pfeffer -- -- x -- -- on ote
Phasmatotenthion richardi (Joubin) -- -- -- == oo oS x

Appendix Table 1.--Food found in stomachs of seven species of tuna from the Atlantic Ocean--Continued

SPECIES OF TUNA

FOOD ITEM Euthynnus Katsuwonus Thunnus Thunnus Thunnus Thunnus' Thunnus
alletteratus pelamis albacares atlanticus thynnus obesus alalunga
INVERTEBRATES
Sepia-sp. >< =o Xx ae ra fost he
Sepictta oweniana g'Orbigny = -- -- oo se aS .
Spirula spirula (Linnaeus) -- -- x -- xX xX xX
Teuthoidea xX xX Xx x Xx a Se
Taoniinae -- _ ee oa Ps BS x
Taonidium pfefferi Russell -- -- -- -- -= mee x
Teuthowenia (Heliocranchia) -- en oe on ak 28 x
pfefferi (Massy)
Todaropsis eblanae (Ball) -- ac x ae Be pe x
GASTROPODA:
Gastropoda (not further identified)-- X x xX -- oo xX
Janthiva sp. a -- -- =: = 2 x
Janthina exigua Lamarck oS -- -- =o — os x
HETEROPODA :
Atlantidae -- Sc) mae ae == os xX
Atlanta sp. -- -- xX 5 -- -- ==
Atlanta peronii LeSueur -- -- om on ed = xX
Heteropoda (not further identifiel) -- xX -- X oe oS oF
Pterotrachea sp. -- -- xX -- X xX X
PTEROPODA :
Cavolinidae -- -- xX -- xX -- xX
Cavolinia sp. -- -- x -- x X X
Clio pyramidata Linnaeus -- a5 sé =: ac — xX
Creseis sp. -- -- -- xX -- =~ --
Cuvierina sp. -- -- xX -- == -= --
Diacria trispinosa (LeSueur) -- -- -- == -- == X
Limacina sp. -- -- ees X oo os ae
Pteropoda (not further identified) -- X aS B55 =e oS a

MISCELLANEOUS :

Chelophyes appendiculatta =o me = =e == 0 xX
(Eschschultz) a0 is rs = ae ae x

Galetta australis? (LeSueur)

Naiades cantrainii (Delle Chiaje) -- os on ae a5 ao xX

Pelagia noctiluca Péron and -- a 7 = == =< X
LeSueur

Pyrosoma atlanticum (Péron) -- == = oe =e == xX

Salpidae -- mae xX = X
Salpa (Iasis) zonaria Pallas -- -- os =o ot ==
Torrea candida (Delle Chiaje) -- -- -- = -- --
Velella velella Linnaeus -- AS os Ac ae ec

Pa PS Pd PS

21

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As the Nation's principal conservation agency, the Depart-
ment of the Interior has basic responsibilities for water,
fish, wildlife, mineral, land, park, and recreational re-
sources, Indian and Territorial affairs are other major
concerns of America's 'Department of Natural Resources,"

The Department works to assure the wisest choice in
managing all our resources so each will make its full
contribution to a better United States -- now and in the future,

UNITED STATES
DEPARTMENT OF THE INTERIOR
U.S. FISH AND WILDLIFE SERVICE
BUREAU OF COMMERCIAL FISHERIES

WASHINGTON, D.C. 20240 POSTAGE ANDIREESIEn

U.S. DEP ARTMENT OF THE INTERIOR

OFFICIAL BUSINESS

Return this sheet to above address, if you do
NOT wish to receive this material [male orif
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XFWS-A 595 1-6 (1970)

US. Fish Wildl, Serv. DIVISION 4 OF F
Spec, Sci, Rep, Fish, U Ss NATIONAL tre

JUL 2 1970

Size, Seasonal Abundance, and
Length-Weight Relation of Some

Scombrid Fishes from Southeast
Florida

CL

-

SPECIAL SCIENTIFIC REPORT-FISHERIES No. 59.5

————— SE EOEEOESeESOOeeEeeEOeeEOeeOEOeeEOeeOeeOEOeOEOeOEOeEOeeeOeeeeese

UNITED STATES DEPARTMENT OF THE INTERIOR mS

ee ee ee eee
p U.S. FISH AND WILDLIFE SERVICE “<
e EEO

SPECIAL SCIENTIFIC REPORT--FISHERIES

EDITORIAL STAFF

Leslie W, Scattergood, Editor

Mary S, Fukuyama, Associate Editor

PUBLICATION BOARD

John A, Guinan

John I, Hodges mm Edward A, Schaefers
Harvey Hutchings Parker S, Trefethen
John M, Patton, Jr. Robert C, Wilson

Leslie W, Scattergood, Chairman

Special Scientific Report--Fisheries are preliminary or progress reports
and reports on scientific investigations of restricted scope, Established as
Special Scientific Report in 1940, nos, 1 to 67 were issued from that date to
1949, when the new Series, Special Scientific Report--Fisheries, with new serial
numbering, was started,

Special Scientific Report--Fisheries are distributed free to libraries, re-
Search institutions, State agencies, and scientists.

UNITED STATES DEPARTMENT OF THE INTERIOR
Walter J. Hickel, Secretary
Leslie L. Glasgow, Assistant Secretary
for Fish and Wildlife, Parks, and Marine Resources
Charles H. Meacham, Commissioner, U.S. FISH AND WILDLIFE SERVICE
Philip M. Roedel, Director, BUREAU OF COMMERCIAL FISHERIES

Size, Seasonal Abundance, and Length-Weight
Relation of Some Scombrid Fishes
from Southeast Florida
By
GRANT L, BEARDSLEY, Jr.,

and
WILLIAM J, RICHARDS

Contribution No. 148, Bureau of Commercial Fisheries Tropical
Atlantic Biological Laboratory, Miami, Fla. 33149

United States Fish and Wildlife Service
Special Scientific Report--Fisheries No, 595

Washington, D.C,
May 1970

~ ee

pag ad eh
: f hia te

CONTENTS

InbrOGUCtIOM. (aici lie iolelereveisnetieliejeyellellene:ejlelelisiie ce
Size distribution and relative apparent abundance,
enpth-weightarelatvonshcveweeheleicteenelenehielenonele
Acknowledpmentsicvericielel eerste lelleletensiotiels) ee) ots

irteraturerciLedunci. fer cmelclien srenete el cuerelemeiterel skere

iii

Oss 3Osee i

Size, Seasonal Abudance, and Length- Weight Relation
of Some Scombrid Fishes from Southeast Florida

By

GRANT L, BEARDSLEY, JR., Fishery Biologist
and

WILLIAM J, RICHARDS, Supervisory Zoologist

Bureau of Commercial Fisheries
Tropical Atlantic Biological Laboratory
Miami, Florida 33149

ABSTRACT

Seven species of scombrid fishes were sampled for length and weight at a taxi-
dermy firm for 1 year (September 1967-September 1968), These data yielded infor-
mation on size distribution and seasonal abundance of the species off south Florida.
In addition, length-weight relations and calculated weights at given lengths are pre-

sented,

INTRODUCTION

Because the southeast coast of Floridafrom
Palm Beach to Key West is one of the major
centers of salt-water sport fishing inthe United
States, samples of a variety of fish are avail-
able for scientific study. In 1967 TABL (Bu-
reau of Commercial Fisheries Tropical At-
lantic Biological Laboratory) in Miami began
sampling several species of scombrid
fishes received from sport fishermen for
mounting by Al Pflueger, Inc., a large taxi-
dermy firm in Hollywood, Fla, The species
sampled were: wahoo, Acanthocybium
solanderi; little tuna, Euthynnus alletteratus;
skipjack tuna, Katsuwonus pelamis; king mack-
erel, Scomberomorus cavalla; Spanish mack-
erel, S, maculatus; cero, S, regalis; and
blackfin tuna, Thunnus atlanticus, This sam-
pling was undertaken to determine the seasonal
presence of scombrids in the area in con-
junction with sampling for their larvae in the
Straits of Florida, Since no commercial fishery
for most of the species exists in this area, we
had to turn to the well-developed sport fishery,

TABL personnel visited the taxidermy plant
each week from September 1967 through Sep-
tember 1968 to examine fishes received the
previous week from the company's various
collecting points along the southFlorida coast,
We did not sample specimens from locations
other than south Florida, We measured length
and weight only, Fork length was recorded to
the nearest millimeter; weight was takentothe
nearest 0,1 pound and was later converted into
kilograms,

Certain problems arise inevaluating samples
of fish from a taxidermy company, Usually
only the largest individuals caught are pre-
served and mounted; size frequencies are
affected accordingly, Apparent seasonal
changes in abundance may reflect merely a
difference in the number of fishermen anda
resulting increase or decrease in the number
of fish received for mounting, Despite these
problems we believe that these data are im-
portant enough to warrant presentation--for
the most part they give a reasonably accurate
picture of seasonal changes in size and abun-
dance of these fishes along the southeast coast
of Florida,

SIZE DISTRIBUTION AND RELATIVE
APPARENT ABUNDANCE

Fork lengths of wahoo, little tuna, skipjack
tuna, king mackerel, and blackfin tuna were
separated by quarters of the year andarranged
into appropriate centimeter groupings, The
length frequencies were plotted, Too few cero
and Spanish mackerel were collected to treat
in this manner,

Wahoo

The wahoo is a highly prized game fish, and
probably a higher percentage are retained for
mounting than any of the other species dis-
cussed; hence seasonal abundance and size
distribution as represented by our samples
are probably more valid for wahoo than for
any of the other species we studied,

Oct-Nov-Dec
N=25

Jan-Feb-Mar,
N-68

NUMBER OF FISH

61-65 81-85 101-105 121-125 141-145 161-165 181-185

FORK LENGTH (CM)

Figure 1,--Length frequency distributions by quarters of
the year for wahoo sampled from a taxidermy plant in
south Florida from September 1967 through September
1968,

Wahoo are most abundant off southern Florida
in winter (fig. 1), LaMonte (1951) reported that
wahoo are taken off Hatteras, N.C., from June
to October and are rare or absent at other
times, Off Bermuda a minor run occurs in
April, and abundance is at a peak in Septem-
ber and October (Mowbray, 1956), In spring a
large number of relatively small (96 to 105
cm.) wahoo appear off the lower Florida
coast, in summer the mode representing this
group of fish (July-September, fig. 1)
progressed slightly, indicating growth of about
3 to 4 cm, per month,

Little Tuna

Little tuna are one of the most common
tunas caught by sport fishermen off southern
Florida and are most abundant in summer
(fig. 2). According to de Sylva and Rathjen
(1961) little tuna are most abundant inthe catch
of sport fish in south Florida from April
through August with a peak in June, Our
samples show that little tuna of 66 to 75 cm,
fork length are dominant in the south Florida
sport-fish catch throughout the year,

Skipjack Tuna

Skipjack tuna, which have become increas-
ingly important to the worldwide commercial
tuna catch in recent years, are most abundant
off Florida in fall and winter, The large mode

in October-December (fig, 3) disappeared in
the next quarter although the single low peak
at 63 to 64 cm, may represent this group of
fish, The peak reappeared in April-June at
about 65 to 68 cm., possibly representing
growth of about 1 cm, per month, Brock (1954)
and Rothschild (1967) showed growth rates for
skipjack tuna in the Pacific of 1.5 to 2,0 cm,
per month, The dominant mode in our samples
in January-March disappeared in April-June,
and a new peak appeared at42to43 cm,, which
indicated that a number of small fish had
moved into the south Florida area,

King Mackerel

The king mackerel, a migratory species, is
most abundant in the winter off southern
Florida where an important fishery for the
king and Spanish mackerels is carried on--in
1967 more than 750 metric tons of king
mackerel and 1,300 metric tons of Spanish
mackerel were landed (Bureau of Commercial
Fisheries, 1968),

The three modes present in the January-.
March quarter (fig, 4) suggest the presence
of only three age groups in the sport-fish
catch off southern Florida, According to a
study of age and growth of king mackerel in
Brazil by Nomura and Rodrigues (1967), how-
ever, the size range represented in figure 4
(January-March) would include their age-
groups IV (62 cm,) through XII (120 cm.),
Catches off Florida are few in summer, and
most king mackerel caught are relatively
small,

Blackfin Tuna

Blackfin tuna, which represent a potentially
valuable commercial resource in the Gulf of
Mexico and Caribbean Sea, are abundant off
southern Florida throughout the fall, winter,
and spring, but sport-fish catches decline
sharply in the summer (fig, 5), The blackfin
is one of the more abundant tunas in the Gulf
and Caribbean but is fished commercially
only in Cuba, where there is an active live-
bait fishery for blackfin and skipjack tunas
off the southern coast, Blackfin tuna make up
about two-thirds of the catch (Suarez Caabro
and Duarte Bello, 1961), and though peak
abundance is from April to July, the fish are
present throughout the year, In our samples
the first mode, in October-December (fig. 5),
appeared to progress from about 47 to 48 cm.
to about 51 to 52 cm, in April-June, which
suggested a growth of slightly under 1 cm,
per month, similar to that indicated for skip-
jack tuna, The second mode in April-June can
possibly be traced back to the large peak at
55 to 56 cm, in October-December, indicating
growth of about 1.5 cm, per month,

NUMBER OF FISH

OCT-NOV-DEC.

JAN- FEB-MAR.
N=105

NUMBER OF FISH

JULY-AUG- SEPT.
N=148

26-27 34-35 42-43 50-51 58-59 66-67 74-75 82-83 90-91

FORK LENGTH (CM)
Figure 2,--Length frequency distributions by quarters of the year for little tuna sampled

from a taxidermy plant in south Florida from September 1967 through September 1968,

OCT-NOV.-DEC.
N-101

JAN- FEB-MAR
N=66

APR-MAY-JUNE

Figure 3.--Length frequency distributions by quar-
ters of the year for skipjack tuna sampled from
a taxidermy plant in south Florida from Septem-
ber 1967 through September 1968,

JULY-AUG-SEPT.

41-42 49-50 57-58 65-66 73-74 81-82

FORK LENGTH (CM)

NUMBER OF FISH

NUMBER OF FISH

OCT-NOV~DEC
N:50

JAN-FEB-MAR

JULY-AUG-SEPT.

56-58 68-70 80-02 92-94 104-106 = NO=118 = 128-130 4421S 2-154

FORK LENGTH (CM)

OCT-NOV.-DEC.

JAN- FEB-MAR.

35-36 43-44 51-52 59-60 67-68 75-76 83-84

FORK LENGTH (CM)

LENGTH-WEIGHT RELATIONS

The relation between length and weight for

each species was calculated by the method of
least squares with the equation

W= aL?
or

log, pW = log, ,a # b log, \L

Figure 4,--Length frequency distributions by quarters of
the year for king mackerel sampled from a taxidermy
plant in south Florida from September 1967 through
September 1968,

Figure 5,--Length frequency distributions by quarters of
the year for blackfin tuna sampled from a taxidermy
plant in south Florida from September 1967 through
September 1968,

where W is weight in kilograms, L is fork
length in centimeters, andaandbareconstants
(table 1), Calculated weights for a given
length--within the length range sampled--are
shown for each species in table 2,

Table 1.--Sample size, size range, and values of the constants a and b in the length-weight

equations for seven species of scombrid fishes sampled from a taxidermy plant in south Florida
from September 1967 through September 1968

Specimens
Length Weight
om kg.

Species

Number Cm.
PROM SOP ANCeT Te = cls. a(arciels (cls 5 3 201 53.4-172.0 0.91-36.29 1.544X10°° 3.29389
Es alletteratuss.... os. 0: 343 23.1-85.8 0.23-8.39 4.956X107 3.26314
PE POP GMT Si. . wcicte'c.c0 ciaieeisbie 244, 37.5-78.0 0.91-9.98 7.927 X10" 3.22750
SMPCAVETI ES coins /cincinaies st 197 58.5-150.0 1.47-32.09 2.701 X10" 3.23000
Smactiietus.ce «ces o00ses 35 33.0-77.0 0.45-4.76 1.053 X10" 2.95842
Senegal Bis <icieedns o's sees 58 213-8325) 7 POSS 22088 web sie 2360Ke 1 OnmmNdoso24911
MepattantScus, sj 110i «siajhies 347 34,5-87.0 0.75-11.79 1.810 X10" 3.02838

Table 2.--Lengths and weights calculated for seven species of scombrid fishes sampled from
a taxidermy plant in south Florida from September 1967 through September 1968

Weight

Fork
length Acanthyocybium Eathynnus Katsuwonus |Scomberomorus | Scomberomorus | Scomberomorus Thunnus
solanderi alletteratus cavalla maculatus regalis atlanticus

@ ----------------------- Kg.------------------------
20 -- 0.11 -- -- -- -- --
30 -- 0.33 -- -- 0.25 0.26 0.54
40 -- 0.84 1.17 -- 0.58 0.60 1.29
50 0.61 11.73 2.41 0.83 1.20 1.15 2.53
60 1.11 3.14 4.35 1.50 1.92 1.96 4.39
70 1.85 5.20 (fea 2.46 3.03 3.08 7.00
80 2.87 8.04 11.02 3.79 4.50 4.55 10.49
90 4.22 11.81 -- 5.54 -- -- 14.97

100 5.98 = = 7.79 - ae ae

110 8.18 -- id 10.59 -- -- --

120 10.91 -- -- 14.04 -- -- --

130 14.18 = = 18.18 ce = =

140 18.10 -- -- 23.09 -- -- --

150 22.72 -- -- 28.86 -- -- --

160 28.10 a se 2c = oe =

170 34.31 -- -- -- -- -- --

180 41.42 -- -- -- -- -- --

ACKNOWLEDGMENTS

Al Pflueger, Jr., and Jerry Webb of Al
Pflueger, Inc., gave us their cooperation and
assistance in obtaining these data,

LITERATURE CITED

BROCK, VERNON E,

1954, Some aspects of the biology of the
aku, Katsuwonus pelamis, in_ the
Hawaiian Islands, Pac, Sci, 8: 94-104,

BUREAU OF COMMERCIAL FISHERIES,

1968, Florida landings 1967, annual sum-
mary, Its Commer, Fish, Statis, 4660,
18 pp.

DE SYLVA, DONALD P,, and WARREN F,
RATHJEN,

1961, Life history notes on the little tuna,
Euthynnus alletteratus, from the south-
eastern United States, Bull, Mar. Sci.
Gulf Carib, 11: 161-190,

LAMONTE, FRANCESCA,

1951. A preliminary survey of marine
angling in North Carolina, In Survey of
marine fisheries of North Carolina,

pp. 251-286, Univ, N.C, Press, Chapel
Hill,
MOWBRAY, LOUIS S,

1956, The gamefishes of Bermuda, Proc,
lst Int, Gamefish Conf,, Nassau, Ba-
hamas, 8 pp. [| Mimeogr.] .

NOMURA, HITOSHI, and MARIA SILVIA DE
SOUSA RODRIGUES,

1967, Biological notes on kind mackerel,
Scomberomorus cavalla (Cuvier), from
northeastern Brazil, Arq, Estag, Biol,
Mar, Univ, Fed, Ceara 7: 79-85,

ROTHSCHILD, B. J.

1967, Estimates of the growth of skipjack
tuna (Katsuwonus pelamis) in the
Hawaiian Islands, Proc, Indo-Pac,
Fish, Counc,, 12th Sess,, Sect. II, pp.
100-111.

SUAREZ CAABRO, JOSE A,, and PEDRO
PABLO DUARTE BELLO,

1961, Biologia pesquera del bonito (Katsu-
wonus pelamis) y la albacora (Thunnus
atlanticus) en Cuba, I, Inst, Cubano
Invest, Tecnol, Ser, Estud, Trab, Invest,
155.151 opps

MS, #1990

GPO 888-347

ne asl

As the Nation's principal conservation agency, the Depart-
ment of the Interior has basic responsibilities for water,
fish, wildlife, mineral, land, park, and recreational re-
sources, Indian and Territorial affairs are other major
concerns of America's "Department of Natural Resources,"

The Department works to assure the wisest choice in
managing all our resources so each will make its full
contribution to a better United States -- now and in the future,

UNITED STATES
DEPARTMENT OF THE INTERIOR
U.S. FISH AND WILDLIFE SERVICE
BUREAU OF COMMERCIAL FISHERIES
WASHINGTON, D.C, 20240

POSTAGE AND FEES PAID
U.S. DEPARTMENT OF THE INTERIOR

OFFICIAL BUSINESS

Return this sheet to above address, if you do
NOT wish to receive this material (ar orif
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a

XFWS-A 598 1-10 969)
U.S, Fish Wildl. Serv.
Spec, Sci. Rep, Fish.

US. NATIONAL MUSEUM

FEB 20 1970

Size Composition, Sex Ratio, and Size
at Maturity of Offshore Northern Lobsters

ay

SPECIAL SCIENTIFIC REPORT-FISHERIES No. 598

UNITED STATES DEPARTMENT OF THE INTERIOR =

U.S. FISH AND WILDLIFE SERVICE =
BUREAU OF COMMERCIAL FISHERIES

~~

SPECIAL SCIENTIFIC REPORT--FISHERIES |
Robert L, Hacker, Editor :

“4
Mary Fukuyama, Associate Editor

PUBLICATION BOARD

John A, Guinan John M, Patton, Jr.
Robert L. Hacker . Edward A. Schaefers
John I, Hodges Parker S. Trefethen
Harvey Hutchings Robert C, Wilson

Leslie W. Scattergood, Chairman

Special Scientific Report--Fisheries are preliminary or progress reports
and reports on scientific investigations of restricted scope. Established as
Special Scientific Reports in 1940, nos. 1 to 67 were issued from that date to
1949, when the new series, Special Scientific Report--Fisheries, with new serial
numbering, was started. ;

Special Scientific Report=--Fisheries are distributed free to Libraries,
research institutions, State agencies, and scientists.

UNITED STATES DEPARTMENT OF THE INTERIOR

Walter J. Hickel, Secretary
Russell E. Train, Under Secretary
Leslie L. Glasgow, Assistant Secretary
for Fish and Wildlife, Parks, and Marine Resources
Charles H. Meacham, Commissioner, U.S. FISH AND WILDLIFE SERVICE

Dayton L. Alverson, Acting Director, BUREAU OF COMMERCIAL FISHERIES

Size Composition, Sex Ratio, and Size at
Maturity of Offshore Northern Lobsters

By

BERNARD E, SKUD and HERBERT C, PERKINS

United States Fish and Wildlife Service

Special Scientific Report--Fisheries No, 598

Washington, D.C,
December 1969

:
es?

af TAY i Mees Sac 2nTATS at Me Ais
WwW

4 hint ; i Oss

CONTENTS

INET OGUCELON swawenetameiteS oie lhe l/oNel(oielsehieneieieliemom sy) ene
Sarmplin pewewente teres eiecl eiisilcioficwe: i eienst ois uel stents

Composition of catches from offshore canyons,

Hudson Canyon, ......
VieatchiiGany.onteme rece cle
Oceanographer Canyon ,
Lydonia Canyon.,.....
Corsair Ganyoni. cnc exe <
Molting a crenerelereiereiteheikelene
Maturity ewovet iehenoneledshop enon
DUTTA Vaweveu Oelonenoleneheieh he

Tite raturexvcitedic cis. sieweltehione

iii

Page

Size Composition, Sex Ratio, and Size
at Maturity of Offshore Northern Lobsters

By

BERNARD E, SKUD and HERBERT C, PERKINS, Fishery Biologists

Bureau of Commercial Fisheries Biological Laboratory
W. Boothbay Harbor, Maine 04575

ABSTRACT

The fishery for northern lobsters, Homarus americanus, inthe offshore waters
of the North Atlantic is developing rapidly andis expected to alter certain biological
characteristics of the stocks, Samples of lobsters have been taken during research
cruises and from commercial catches to document the existing size composition,
sex ratio, and size at maturity, These records will provide useful indices of popula-
tion changes as fishing effort increases,

The data in this report are from five of the major fishing areas--
Hudson, Veatch, Oceanographer, Lydonia, and Corsair Canyons, Lobsters from
canyons 200 km, from shore were substantially smaller than those from areas
farther offshore, The size oflobsters generally increased with depth and the propor-
tion of females generally increased with increasing size, Females dominated the
samples and sometimes were 70 percent of the catch, The numbers of egg-bearing
females are reported, The smallest female with external eggs was 8.0 cm, in car-

apace length,

INTRODUCTION

Landings of northern lobsters (Homarus
americanus) from the offshore trawl fishery
of the North Atlantic have been increasing
rapidly during the past 10 years, Otter trawls
took 484 metric tons of lobsters in 1958 and
over 2,500 metric tons in 1968, As the fishery
develops, it is important to document the
changes in the size composition and sex ratio
of the stocks, These data, together with catch
and effort statistics, will be needed to assess
the effect of fishing on the abundance of off-
shore lobsters, Sampling of the catch was too
limited in earlier years to detect changes in
population structure, During the past 3 years,
however, sampling of commercial landings
and of catches from research cruises has been
more extensive and has provided information
by fishing area that will be a base for future
comparison,

The early history of the offshore fishery was
described by Firth(1940) and Schroeder (1959).
The first reports of lobsters caught by trawling
date back to the early 1900's, when beam and
otter trawls were introduced to American
waters, These lobsters were incidental catches
in the fisheries for various groundfish, Inci-
dental catches continue but usually are not

included in the catch statistics of the lobster
fishery. The earliest otter trawl catch of
lobsters that is recorded was 4 metric tons
landed in New York in 1921; the exact location
of the catch is unknown, Even through 1960,
little information was gathered on the location
of the offshore catches,

Statistics of the Bureau of Commercial
Fisheries do not separate inshore and off-
shore catches of lobsters; however, practically
all of the fishing inshore is done with pots
whereas otter trawls are used offshore, Trawl
catches from the New England region to the
Chesapeake date back to the late 1920's; and
though a few inshore catches are included,
these data serve to document the increase in
offshore landings that has occurred over the
years and show that these catches remained
less than 45 metric tons annually through 1946,
Skud (in press) described the increased catches
of recent years, From 1947, the landings
increased steadily and approached 500 metric
tons for the first time in 1955 (table 1), This
growth continued and the peak offshore land-
ing--2,500 metric tons in 1968--accounted
for 16.9 percent of the total U.S, catch of
lobsters, Landings from the offshore fishery
contribute substantially to the total landings in
Massachusetts, Rhode Island, New York, and

Table 1.--U.S. catch of lobsters by the inshore
pot fishery and offshore trawl fishery

Pot Trawl Percentage

Year | fishery | fishery | TOt@l | of catch
by trawls
----- Metric tons- - - - Percent

1943 7,420 30 7,450 0.4
1944 8,095 317, 8,132 0.5
1945 10,275 34 10,309 0.3
1946 10, 840 35 10,875 0.3
1947 10,802 51 10,853 (0) 45)
1948 9,417 59 9,476 0.6
1949 11,102 124 alabyAs) algal
1950 10 , 394 129 10,523 2,
alesenk 11,680 89 11,769 0.8
1952 dlabealte)s) 159 11,354 ee.
1953 12,479 275 12,754 2.2
1954 12,078 386 12,464 Shae
1955 12,649 482 aleyo dksye S\O/
1956 alae): 510 12,025 4.2
1957 13,316 2517 13,673 2.6
1958 11,858 484 12,342 Soe)
1959 12,589 592 13,181 4.5
1960 ale }es jal 810 14,121 Disc,
1961 11,621 1,056 12,677 8.3
1962 12,124 1,236 13 , 360 9.2
1963 12,369 1,366 13',.735 9.9
1964 aba AbeArd 1,855 14,032 13.2
1965 gee 2,481 13,702 18.1
1966 11,612 1,774 13,386 als} 3}
1967 10,083 1,949 12,032 16.2
1968* 12,300 2,500 14,800 16.9

*Preliminary.

New Jersey. None of the Maine catch includes
offshore lobsters,

The fishery operates at the edge of the
Continental Shelf, 100 to 300 km, from shore,
mainly in depths between 200 and 500 m, The
vessels, which are usually about 30 m, long,
operate on a year-round basis, The average
fishing trip lasts 10 days; the lobsters are
held alive in tanks until the vessel returns to
port, Catches during a trip may range from
1 to 9 metric tons, but the daily average is
about one-half a metric ton, Lobsters weighing
less than 2,25 kg, are termed selects and
command a premium price; larger lobsters
are called jumbos and sell for 50¢ to $1.00
less per kilogram, The average annual price
to the fisherman is about $1.75 per kilogram,
for selects and $1.25 for the jumbos, but the
price of selects has been as high as $2.50 per
kilogram in late winter and early spring,

The gear used in this offshore fishery is
the standard bottom trawl described by Bruce
(1967), If the bottom is rough, rollers are
added to the foot rope of the nets to help pre-
vent snagging. The boats fish aroundthe clock,
and tows last 3 to 4 hours,

Skud (1966) reported on preliminary sam-
pling of the offshore fishery but didnot discuss
the differences among individual fishing
grounds, The objectives of the present paper
are to present the results of research cruises
and sampling of the commercial catch, and
to compare size composition, sex ratio, and
size at maturity of the lobsters caught in off-
shore fishing areas,

SAMPLING

Most of the samples of lobsters were taken
from catches at submarine canyons along the
edge of the Continental Shelf (fig. 1). As the
sampling stations had marked differences, we
discuss each of the canyon areas separately,

Samples from the commercial catch were
taken throughout the year, At least 100 lob-
sters, representing all size categories, were
measured (carapace length) and sexed from
each of the landings sampled, In addition, data
on the vessel, gear, date sailed, date landed,
days fished, fishing days lost, area and depth
of catch, and total catch were obtained from
interviews with the vessel captains,

During research cruises aboard the Alba-
tross IV and the Delaware, more detailed
data on tows were obtained, including the time
and distance as well as the range in depth of
each tow, Hydrographic data were collected
after each tow, Sampling during the research
cruises was concentrated in particular fishing
areas to determine seasonal changes in size
composition of sex ratio. Since these same
areas were also regularly fished by the com-
mercial fleet, we could compare the catches
of commercial and research vessels, The sex,
carapace length, and weight of each lobster
were recorded, The number of females with
external eggs (berried) was also noted, A
summary of the research cruises is presented
in table 2,

The research cruises also provided infor-
mation on sublegal or prerecruit lobsters
which was not available from commercial
landings because of minimum-size restrictions
(7.8-8.1 cm, carapace lengthin various States),
Though the fishery operates in international
waters, which have no restrictions on lobster
fishing, State laws apply to the landings; con-
sequently the offshore fishery is subject to
the same size regulations that control the in-
shore fishery, All of the New England States
also prohibit the landing of berried lobsters,

COMPOSITION OF CATCHES FROM
OFFSHORE CANYONS

The five canyon areas discussed in this
paper were selected as sampling sites because
of their commercial importance or their geo-
graphic separation, to provide a composite of

ae.

GEORGES BANK

Figure 1.--Sampling locations for lobsters on the Continental Shelf off southern New England.

Table 2.--Lobster research cruises:
catch, 1965-67

locations, fishing data, and

2 D
pea) ae ae pan noye | lOtal| Depth | Lobster
Cruise and vessel Canyon | +OWS | time | range | catch
ee
Number Hours Meters Number

6 6 113-146 108

2 2 126-182 5

2 2 160-237 16

3 6 171-271 30

5 11 273-455 174

1 1 337 26

Bb wm 124-173 «4159

3 3 106-109 77

5 5 91-273 0)

2 2.5 200-246 104

4 4 118-206 168

2 2 209-273 12

10 10 109-237 314

6 6 127-182 1

7 7 135-246 217

10 10 167-237 140

10 10 146-355 Wl

10 10 126-260 81

4 4 109-218 47

5 5 118-228 38

2 2 109 439

6 6 91-127 23

4 4 136-300 88

“4 4% 109-364 515

11 10.5 109-182 548

9 8.5 136-164 323

9 8.5 118-218 268

Hudson 6 9.25 109-127 279
Veatch 5 8 113-135 422
Oceanographer 6 8 100-136 54

the offshore fishery, There are other offshore
grounds that are fished by commercial ves-
sels, These include an area 35 km, southwest
of Hudson Canyon between the 200-and400-m,
contours, and the Southwest Part, Southeast
Part, and north central areas of Georges Bank,

The oceanographic features of the area have
been reported by Colton, Marak, Nickerson,
and Stoddard (1968), The temperature at the
depths where lobsters are caught differs as
much as 5.0° C, between canyons, Generally
there is a cooling trend from west to east,
Periodically a tongue of water from the Gulf
Stream intrudes into the area of Oceanographer
and Lydonia Canyons, warming the bottom
water to somewhat higher temperatures than
occur westward at Veatch Canyon, Corsair
Canyon, far to the east, is consistently the
coldest area,

Hudson Canyon

Hudson Canyon (lat, 39°35! N., long.
72°25! W.) lies 100 km, south of eastern Long
Island, N.Y., and is the farthest west and south
of the sampled areas, The canyon was sampled
during research cruises in April, June, and
October 1967. Tows made at various depths
between 100 and 215 m, took a total of 907
lobsters (table 3),

During the June cruise, six tows in the Hud-
son Canyon area caught 470 lobsters--an
average of 78 per 1-hour tow, In October, 279

Table 3.--Catch of lobsters by research vessels, Hudson Canyon,

1965-67
| 7 ] | Females
Carapace | | | =a a Se |
length | Females Males | Total || Percentage
| | of total |Berried| Berried
| | | | lobsters
Cn. Number Number Number Percent Number Percent
4 1 1 2 50.0 -- -~
5 24 9 33 72.7 -- --
6 87 28 115 75.7 -- --
7 167 75 242 69.0 -- --
8 158 72 230 68.7 6 3.8
9 119 31 150 79.3 49 41.2
10 73 12 85 85.9 38 52.1
il 16 v 23 69.6 12 75.0
12 9 1 10 90.0 3 33.3
als} 6 1 7 85.7 5 83.3
14 1 1 2 50.0 ab 100.0
1 3 10) 5) 100.0 ie) 0.0
16 1 1 2 50.0 ie) 0.0
17 1 0 1 100.0 e) 0.0
18 ie) al 1 0.0 (0) 0.0
19 ie) 0 e) 0.0 ie) 0.0
20 ie} 1 1 0.0 ie) 0.0
Total 666 241 907 73.4 114 17.1

lobsters were taken in six tows, averaging
30 per 1l-hour tow, Soft or recently shed lob-
sters were taken in June but not inOctober,

The October catch included 189 lobsters with
carapace lengths of 8 cm, or larger (i.e.,
above the legal minimum), Of these, 160 were
females and 99 (62 percent) of the females were
berried, The females were from 8 to 15 cm,
long and the berried individuals averaged
9.8 cm. Eggs of the berried lobsters had been
extruded recently and were in prenaupliar
condition,

Females dominated the catch in all but the
largest size categories and accounted for 70
percent of the total, but the ratio of females
to males decreased with depth, In a regres-
sion analysis in which the ratio of females to

males was a dependent variable on depth,
there was a significant negative relation
(b = =0.298, Sp = O.1 11; b/s, = 2.685,

P = 0,02), Though the sex ratio changed, the
mean size of lobsters did not increase sig-
nificantly with increasing depth,

Veatch Canyon

Veatch Canyon (lat. 40°00! N,, long. 69°35!
W.) is 130 km, south of Nantucket Island,
Mass, It was fished more frequently during re-
search cruises than any other of the off-
shore areas, Catches here are usually good,
and the area receives considerable effort
from commercial fishing vessels, The canyon
was sampled in April, June, and October
1965; April, June, July, and October 1966;
and April, June, and October 1967, Table 4
Summarizes data obtained from these re-
search cruises, and table 5 summarizes the
samples taken from catches by commercial
vessels from December 1965 through June
1967.

Table 4.--Catch of lobsters by research vessels, Veatch Canyon,

1965-67
Females
pera Females | Males Total ||Percentage
of total | Berried | Berried
lobsters
Cn. Number Number Number Percent Number Percent

4 1 iz 8 12.5 -- --

5 62 71 133 46.6 -- --

6 162 208 370 43.8 -- --

7 286 351 637 44.9 -- --

8 264 327 591 44.7 te) --

9 194 226 420 46.2 25 12.9
10 94 102 196 48.0 25 26.6
iat 47 62 109 43.1 11 23.14
12 33 23 56 58.9 8 24.2
3 21 b 36 58.3 5 23.8
14 7 6 3 53.8 1 14.3
15 14 5 19 73.7 1 Tel
16 al, 8 9 11.1 to) 0.0
17 2 7 9 22.2 1 50.0
18 1 5 6 16.7 ie) --
19 1 3 4 25.0 0 --
20 (e) ie) (0) -- -- --
21 ie) 2 2 -- -- -

Total 1,190 1,428 2,618 45.5 7 6.5

Table 5.--Samples of the commercial catch of
lobsters from Veatch Canyon, December 1965
to June 1967

Gerne Females | Males | Total Poeaen
Cm. Number Number Number’ Percent
8 334 283 617 5401
9 825 620 1,445 PHfodk
10 583 SyAl 954 61.1
an 364 243 607 60.0
12 369 163 532 69.4
alge) 298 120 418 YAlo3}
14 177 95 272 65.1
15 107 80 187 Dilise,
16 65 76 141 46.1
17 Al 59 100 41.0
18 21 64 85 24.7
19 17 35 52 SY207/
20 6 29 3D aoa
21 0 8 8 --
22 0) 3 3} --
Total 3,207 2,249 5,456 58.8
During research cruises, more lobsters

were caught per hour in shoaler water: 49 per
hour in 100 to 182 m, (55-100 fathoms) and 12
per hour in depths greater than 182m. (fig. 2).
Greater depths yielded fewer but larger lob-
sters; lobsters taken below 182 m, averaged
10 cm, in carapace length and 900 g. (2 pounds),
and those taken in shallower water averaged
8 cm, and 450 g, (fig. 3). Commercial catches
generally produced a higher yield in weight of

HUDSON VEATCH OCEANOGRAPHER LYDONIA CORSAIR
41-50- (91 meters)

31-60
61-70 i
7I-80-
RES

eal (182 meters)

101-110

i)
1
i
I
1-120 |
121-130 t
' i
|
1
i]
I
I
1
i]
I
—_—
1
t
I
i]
'
1
i]
>

(--------

141-150 (273meters)

131- mal ————_—S>- |
[

1SI-I60

T
(Sirocco

DEPTH (FATHOMS)

161-170)

171-180
18!-190-
191-200; (364meters)
al

211-2207

221-230

231-240

el (455meters) dt (a tebe ee eee (ere ee ay
25 0 25 20 ° 20 10 0 10 5 0 5 15 o. 8615

PERCENT OF TOTAL

------—------@-------

Figure 2,--Percentage of lobsters caught at various depths in the offshore fishery,

oy 182 METERS or LESS
N= 2374

PERCENTAGE OF TOTAL CATCH

CARAPACE LENGTH (cm)

Figure 3,--Length distributions of lobsters caught above and below 182 m. by research vessels at Veatch Canyon,

catch per day in waters deeper than 180 m,
than in shoaler waters, This evidence suggests
that waters between 100 and 182 m, are the
realm of most prerecruit lobsters at Veatch
Canyon and that lobsters move gradually to
deeper waters as they grow, The catch per
hour in shoaler waters was greater during
daylight (0600-1800 hours) than at night
(1800-0600), Catches averaged 65 lobsters per
hour during the day and 28 at night, In depths
greater than 182 m, there was no appreciable
difference; day tows caught 11 lobsters per
hour, night tows 13, Time of day had no effect
on the variation of carapace length orthe ratio
of females to males,

The depth of fishing at Veatch Canyon has a
direct effect on the composition of the catch,
Tows during research cruises were made at
depths from 100 to 450 m, (55-250 fathoms),
As depth increased, the ratio of females to
males increased, The linear regression of
percentage of females dependent on the depth
fished is highly significant (b = 0.131,
Sp = 0.035, b/s, = 3.743, P< 0,01). The same
level of significance was obtained withthe data
from commercial catches, The meancarapace
length of lobsters increased with depth, but
preliminary results from a tagging study sug-
gest seasonal changes in the relation of size to
depth, Not enough tows have been made in all
months at all depths to show these differences,

Oceanographer Canyon

Oceanographer Canyon (lat, 40°25' N., long,
68°10' W.) is 30 km, south of the Southwest
Part of Georges Bank and 100 km, east north-
east of Veatch Canyon,

The catches of lobsters during research
cruises at Oceanographer Canyon in April and
June 1965, April and June 1966, and June and
October 1967 are summarized in table 6, Tows
were made in depths of 100 to 337 m, (55-185
fathoms),

Differences were not significant between the
average numbers of lobsters caught per hour
in daylight and at night, Tows in water 182 m,
deep or less produced five lobsters per hour
during the day and six at night; tows made in
deeper water produced 31 per hour during the
day and 24 at night, The increase in catch
with depth, however, was significant, Here,
as in the other areas, the ratio of females to
males increased with depth of capture, The
increase held between 100 and 235 m, but did
not extend beyond 235 m, where percentage
females in both the research and commercial
samples varied between 65 and 75,

The most striking difference between Ocean-
ographer and Hudson and Veatch Canyons is
the preponderance of large lobsters, 12 cm,
and more in carapace length, This difference
was also evident in commercial samples
(table 7),

Table 6.--Catch of lobsters by research vessels, Oceanographer
Canyon, 1965-67

= | 7 | = -
| Females
STE Females | Males | Total Per rcentage
| | of total |Berried| Berried
lobsters
Cn. Number Number Number Percent Number Percent
5 3 e) 3 100.0 -- --
6 2 9 11 18.2 -- --
7 8 15 23 34.8 -- --
8 10 17 27 37.0 1 10.0
9 19 16 35 54.3 3 15.8
10 40 13 53 75.5 6 15.0
ll 34 3 47 72.3 10 29.4
12 44 21 65 67.7 4 9.1
13 62 12 74 83.8 13 21.0
14 43 15 58 T4.1 8 18.6
15 26 9 35 74.3 - 15.4
16 27 e) 36 75.0 7 25.9
17 8 1) 23 34.8 ie} --
18 10 8 18 55.6 1 10.0
19 9 5 14 64.3 1 11.1
20 1 4 5 20.0 ie} --
21 ie) 5 5 -- -- ==
22 ie) 1 1 -- -- --
23 ie) 1 1 -- -- ==
Totals 346 188 534 64.8 58 16.8

Table 7.--Samples of the commercial catch of
lobsters from Oceanographer Canyon, February
1966 to June 1967

aca Females | Males | Total ee
cm, Munber Number Number Percent
8 16 n mp Bi).
9 67 55 122 54.9
10 OL 4B ee 8 Gl
11 168 49 217 = “FIA
12 213 68 281 75.8
13 260 «89 349) 8 7s
4 24 «77 291 «73.5
15 9 O77 1760 Sone
16 ay Ge) 123) en
17 9149 140 65.0
18 65 | 47 12. «58.0
19 Bu ha3 67 50.7
20 12 21 33) masons
21 A NG 18 11.1
22 0 5 5 0.0
Totals 1,416 699 Bpals) 67.0

At Oceanographer Canyon, as at Veatch
Canyon, greater fishing depths produced larger
lobsters, The mean carapace length of lob-
sters caught was 11 cm, at depths of 182 m.
or less, and 13 cm, at depths of 182 m, or
more (fig. 4).

182 METERS OR LESS
N=88

A

KN >182 METERS.
| N Neda

PERCENTAGE OF TOTAL CATCH
ey tn Feed :

4 6 8 10 Me

12 D Ye
CARAPACE LENGTH (cm)

Figure 4.--Length distributions of lobsters caught above
and below 182 m. at Oceanographer Canyon.

Lydonia Canyon

Lydonia Canyon is at lat, 40°30' N., long.
67°40! W., 35 km. east of Oceanographer
Canyon, The area was sampled during research
cruises in April 1965 and in April, June, July,
and October 1966. Tows were made at depths
90 to 355 m, (50-195 fathoms), The dominance
of large lobsters here, as at Oceanographer
Canyon, is apparent both in the researchcatch
(table 8) and the commercial samples (table 9),

Depth did not have a significant effect onthe
ratio of females to males, or the mean size of
the catch, and the proportion of females did
not increase with the mean carapace length
of lobsters in a tow,

In depths of 182 m., or less, the difference
between numbers of lobsters caught per hour
in daytime (14) and at night (18) was not sig-
nificant, In depths greater than 182 m., more
lobsters were caught per hour in daylight (33)
than at night (14) but the sample size was

Table 8.--Catch of lobsters by research vessels, Lydonia Canyon,

1965-67
| |
| | Females
Soe Females | Males | Total ||Percentage
| of total | Berried| Berried

| | | | lobsters
a Number Number Number Percent Number Percent
5 0 1 1 0.0 -- --
6 2 2 4 50.0 -- --
tf 3 9 12 25.0 -- --
8 S) 10 19 47.4 0 --
9 i 21 32 34.4 al 9.1
10 23 21 a 52.3 te 30.4
i 29 24 53 54.7 5 17.2
i2 58 12 70 82.9 15 25.9
pis) 5 26 82 68.3 4 25.0
uw 50 27 67 59.7 12 30.0
15 a 22 43 48.8 4 19.0
16 7 13 20 35.0 1 14.3
17 16 18 34 47.1 3 18.8
18 9 9 18 50.0 3 33.3
19 5 13 12 27.8 1 20.0
20 0 9 9 -- -- =
21 0 5 5 -- -- =o
22 8) 3 3 -- -- os
2 is) 1 ab -- ~~ ere

re)
E
D

x

oO
3S
¥
Wn
w
Ww
Ww
R
oO
iN)
ie)
a

Table 9.--Samples of the commercial catch of
lobsters from Lydonia Canyon, December 1965
to July 1966

Carapace Percentage

een Females | Males Sekronales

Cm. Number Number Number Percent
8 22 21 43 51.2
9 52 60 112 46.4
10 63 45 108 Bie555}
ala 86 57 143 60.1
12 91 45 136 66.9
1B 98 61 159 61.6
14 75 48 123 61.0
15 57 44 101 56.4
16 33 57 90 36.7
17 30 47 LHL 39.0
18 Pail, 42 63 333
19 9 44 53 17.0
20 5) 30 35 14.3
al 2 19 21 9.5
22 fe) 7 7 0.0
Totals 644 C2 eee 50.7

small and more data are needed to substan-
tiate this difference, When catches from all
tows in 182 m. or less were combined and
compared with those in all tows made at
greater depths, significant difference in catch
per hour was apparent,

Corsair Canyon

The area sampled is at lat, 41°15'N., long.
66°22' W., 154 km, northeast of Lydonia
Canyon and 15 km, southwest of Corsair
Canyon, The area was sampled during re-
search cruises inApril, June, and October 1965
and in April, June, and July 1966, Table 10

Table 10.--Catch of lobsters by research vessels, Corsair Canyon,

1965-66
Females
eae ean Females Males | Total || Percentage
of total |Berried| Berried
lobster
Cn. Number Number Number Percent Number Percent
©) (0) 3 3 0.0 -- --
10 2 1 3} 66.7 1 50.0
11 7 3 10 70.0 3 42.9
12 24 8 32 75.0 13 54.2
13 18 15 33 54.5 7 38.9
4 30 11 41 732 15 50.0
15 30 8 38 78.9 19 63.3
16 28 18 46 60.9 16 57.1
17 19 4 23 82.6 5 26.3
18 20 8 28 71.4 12 60.0
19 1 6 7 14.3 at 100.0
20 2 ab 3 66.7 2 100.0
21 (0) 3 3 -- -- --
Totals 181 89 270 67.0 94 51.9

summarizes the results, Catches averaged be-
tween 8 and 12 lobsters per hour in this area
in April and June 1965 and June 1966; but only
one lobster was taken in a total of 11 tows
made in October 1965 and April and July 1966.
The results of sampling are presented pri-
marily to show the great size of lobsters in
the area,

MOLTING

Only a few soft (recently molted) lobsters
have been taken during the research cruises
(table 11), Soft lobsters are knownto seek pro-
tective areas and would not be expected to
enter the catch in substantial numbers, The
percentage of soft lobsters was highest at
Veatch Canyon in July 1966, when 39.6 percent
of 47 lobsters collected were soft, Though we
can be reasonably sure in assigning July and
perhaps August as peak months of molting at
Veatch Canyon, there is also a suggestion that
molting occurred later in the eastern areas
where the winter temperatures are lower,

Table 11.--Percentage soft (recently molted) lobsters in the catch
by research vessels

seers Hudson Veatch | Oceanographer | Lydonia | Corsair

Percent Percent Percent Percent Percent
April 1965 -- 0.9 0.0 0.0 0.0
April 1966 0.0 0.0 0.0 0.3 0.0
April 1967 0.0 2.8 -- a =
June 1965 -- 1.7 0.0 -- 0.0
June 1966 -- 1.8 0.0 0.0 0.0
June 1967 0.9 7.4 2.2 -- os
July 1966 -- 39.6 0.0 2.6 --
Oct. 1965 -- 1.3 -- -- 0.0
Oct. 1966 -- 0.9 -- 4.3 —_
Oct. 1967 0.0 0.2 4.8 0.0 --

MATURITY
Table 12 shows the number of female

lobsters and the number and percentage ber-
ried from the five canyon areas, The smallest
berried lobster had a carapace length of 8cm.
Between carapace lengths of 10 and 16 cm,
only 30 percent of the females were berried,
though all females within this size range were
considered to be sexually mature,

Templeman (1935, 1944) reported a method
for determining maturity of females by the in-
crease in the width of the abdomen relative
to total length, We measured the width of the
second abdominal segment and carapace length
of 1,700 female lobsters from the five can-
yons, The lengths were 4 to 20 cm, In our
analysis we included all females, whether
berried or not, and used both the actual width
of the abdominal segment and its ratio to the
carapace length (Perkins and Skud, 1966).

Table 12.--Percentage of berried females by
size groups, all canyons combined

Bere p ace Females B i

length erried females
Cn. Number Number Percent

8 441 V 1.6

9 343 78 Reel:

10 232 77 33.2

lat, 133 41 30.8

12 168 43 25.6

13 163 44 27.0

14 121 37 30.6

15 94 28 29.8

16 64 24 37.5

17 46 9 19.6

18 40 16 40.0

19 16 3 18.8

20 3 2 66.7

Total 1,864 409 2L69

The relation of the actual width ofthe second
abdominal segment to carapace length repre-
sents a significant curvilinear regression
which is best fitted by the cubic regression
equation Y=a+t+bxtc +dx?, For con-
venience the carapace lengths were grouped
into 5-mm, divisions and the mean width of
the segments within each of these divisions
was considered as the corresponding de-
pendent variable, Figure 5 shows the second
abdominal width-carapace length ratios and
the corresponding carapace lengths,

From observations of berried lobsters, we
know that maturity is attained by some females
that have a carapace length of 8 cm, The data
plotted in figure 5 show an inflection at 7.7
cm., and an asymptote is reached at about
10 cm, This indicates that the morphometric
changes are associated with maturity and sug-
gests that all females above 10 cm, are
mature,

To test this hypothesis, 54 female lobsters
were examined to determine the condition of

a

g

ABDOMINAL WIDTH
CARAPACE LENGTH

&

8
Issa a pe aes Delle ss
o

4 1 n L n 1 1 1 yt
6 7 8 3 10 " rR s 4 6 16 v 1 ’ 20

CARAPACE LENGTH (CM)

&

Figure 5.--Ratio of the width of the second abdominal
segment to carapace length.

their ovaries and internal egg size, These
lobsters, captured in May and June 1967 from
Hudson, Veatch, and Oceanographer Canyons,
had carapace lengths of 7.6 to 10.7 cm, The
developmental stage of the ovary was cate-
gorized according to (1) color (white indicated
immature ovaries; yellow, maturing; and green,
mature) and (2) mean egg diameter (less than
0.4 mm, indicated an immature ovary; 0.4 to
0.8 mm,, a maturing ovary; and larger than
0.8 mm.,, a mature ovary), Egg diameter was
determined from a sample of 10 eggs taken
from the surface of the ovary, The width and
length of each egg were measured, and an
average of the two measurements was used as
the ''size,'' Lobsters with immature eggs had
carapace lengths of 7.6 to 8.4 cm.; females
with developing eggs were 7.6 to 9,1 cm.; and
those with mature eggs were 8.0 to 10.7 cm,

Measurements of the width of the second
abdominal segment had been taken for 41 of
these females from Hudson, Veatch, and Ocean-
ographer Canyons, When the measurements of
the ovarian eggs of these 41 individuals were
plotted against the corresponding width of the
abdominal segment, a definite relation was
apparent (fig, 6), The highly significant corre-
lation between the egg size and the abdominal
width (r = 0.897, P < 0,01), and between egg
size and carapace length confirms the relation
of morphometric changes and maturity, Though
exceptions can be anticipated, on the basis of
our samples all females over 10 cm, in length
(carapace) and/or with an abdominal width
greater than 6 cm, are mature,

20

is

SIZE OF EGGS IN OVARY(MM)

WIDTH OF SECOND ABDOMINAL SEGMENT (MM.)

Figure 6.--Relation between width of second abdominal

segment and size of internal eggs.

SUMMARY

The trawl-fishery for lobsters in the off-
shore waters of the North Atlantic is de-
scribed and the increase inlandings during re-
cent years is discussed, This fishery con-
tributed less than 1 percent to the U.S, lobster
landings before 1948, but accounted for 17
percent of the total in 1968,

Samples of lobsters were obtained during
research cruises andfrom commercial catches
in five fishing areas--Hudson, Veatch, Ocean-

ographer, Lydonia, and Corsair Canyons, The
size composition, sex ratio, and number of
egg-bearing lobsters are summarized by can-
yon,

Lobsters were smaller and apparently more
numerous in the areas of the Continental Shelf
west of and including Veatch Canyon than in
areas to the east, At several canyons the
average size of lobsters and the ratio of fe-
males to males increased with depth, and at
some canyons there were striking differences
between the numbers of lobsters caught in
shoal waters (100-182 m.) and those taken in
deep waters (over 182 m.), At Veatch Canyon,
more lobsters were taken in shoal waters,
whereas at Lydonia catches were much better
in deep water than in shoal areas, There were
also differences of day and night catches at
the various canyons,

Females dominated the catch, at times ac-
counting for 70 percent of a sample, The ratio
of females to males was about 1:1 at sizes
below 8 cm, carapace length, From 9 to 15
cm, the proportion of females increased but
declined thereafter; at the largest sizes, 19 to
21 cm., males predominated, Skud (in press)
examined the size composition and sex ratio by
canyon area and hypothesized that the dif-
ferences were due mainly to the effects of
fishing,

Molting occurred from spring through fall
in most areas, July appeared to be the month
of peak molting at Veatch Canyon, and there
was an indication that greatest molting activity
occurs later in the areas east of Veatch Can-
yon,

The number of egg-bearing females in each
size group was tallied onthe researchcruises,
and these data, coupled with measurements
of the second abdominal segments and ovarian
eggs, showed that maturity may be attained at
a carapace length of 8,0 cm, and indicated that
all females are mature at a carapace length
of 10.0 cm, or at an abdominal width (second
segment) greater than 6 cm,

LITERATURE CITED

BRUCE, ROBERT A,
1967. North Atlantic trawl nets. U.S, Fish
if Wildl, Serv., Fish, Leafl, 600, 23 pp.
COLTON, JOHN B. JR., ROBERT R. MARAK,
SAMUEL R, NICKERSON, and RUTH STOD-
DARD,
1968, Physical, chemical, and biological ob-
servations onthe Continental Shelf, Nova
Scotia to Long Island, 1964-66, U.S, Fish
Wildl. Serv., Data Rep. 23, 189 pp.
FIRTH, FRANK E,

1940, Giant lobsters, New Engl, Natur. 9:

11-14,
PERKINS, HERBERT C,, and BERNARD E,
SKUD,

1966, Body proportions and maturity of fe-
male lobsters. Amer. Zool. 6: 615.

SCHROEDER, WILLIAM C,

1959, The lobster, Homarus americanus,
and the red crab, Geryon quinquidens,
in the offshore waters of the western
North Atlantic. Deep Sea Res. 5: 266-
282.

SKUD, BERNARD E,

1966, Size composition, sex ratio, and mat-
urity of offshore lobsters,
6: 362-363,

In press. Giant lobsters, In Frank E, Firth
(editor), Encyclopedia of Marine Re-
sources, pp, 353-357. Van Nostrand
Reinhold Company, New York,

10

Amer, Zool,

In press. The effect of fishing on size com-
position, and sex ratio of offshore
lobster stocks, Fiskeridir, Skr. Ser.
Havunders,

TEMPLEMAN, WILFRED.

1935, Local differences in the body propor-
tions of the lobster, Homarus amer-
icanus, J. Biol, Bd. Can. 1:213-226,

1944, Abdominal width and sexual maturity
of female lobsters on Gandian Atlantic
coast, J, Fish, Res. Bd, Can. 6: 281-
290,

MS. #1936

GPO 882-648

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As the Nation's principal conservation agency, the Depart-
ment of the Interior has basic responsibilities for water, ;
fish, wildlife, mineral, land, park, and recreational re-
sources, Indian and Territorial affairs are other major
concerns of America's 'Department of Natural Resources,"

The Department works to assure the wisest choice in
managing all our resources so each will make its full
contribution to a better United States -- now and in the future,

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