363
A REVIEW OF LITERATURE ON MENHADEN WITH SPECIAL REFERENCE TO THE GULF OF MEXICO MENHADEN
SPECIAL SCIENTIFIC REPORT-FISHERIES Na 363
UNITED STATES DERARTMErn^^FJ]H^^ FISH AND WILDLIFE SERVICE
United States Department of the Interior, Fred A. Seaton, Secretary Fish and Wildlife Service, Arnie J. Suomela, Commissioner Bureau of Commercial Fisheries, Donald L.. McKernan, Director
A REVIEW OF LITERATURE ON MENHADEN WITH SPECIAL REFERENCE TO THE GULF OF MEXICO MENHADEN, Brevoortia Patronus GOODE
by
Gordon Gunter and J. Y. Christmas
United States Fish and Wildlife Service flpecial Scientific Report- -Fisheries No. 363
Washington, D.C. October I960 This work was financed by the Bureau of Commercial Fisheries under Contract No. 14-19-9335, with funds made available under the Act of July 1, 1954 (68 Stat. 376), commonly known as the Saltonstall-Kennedy Act.
11 CONTENTS
Page
Int r oduc ti on 1
Gulf menhaden fishery statistics 1
Menhaden biology 3 Taxonomy, distribution and relationships 3 Atlantic nnenhaden 6 Spawning 6 Growth 7 Distribution and migrations 9 Food and feeding 12 Enemies and n^ortality 13 Parasites and diseases 14 Menhaden fishery investigations 14 Gulf of Mexic o menhaden 15 Spawning 15 Gr owth 17 Distribution and migrations 18 Food and feeding 21 Enemies and mortality 21 Parasites and diseases 22 Menhaden fishery investigations 22
Summary 23
Literature cited 25
111 ABSTRACT
Menhaden have been important conrimercial fishes in the United States since colonial days. In this review of the literature on menhaden biology, scattered sources of information are brought together. The chief papers on Atlantic menhaden and practically every piece of biological literature on Gulf of Mexico menhaden are cited. Atlantic and Gulf men- haden literature are treated separately in topical arrangement.
The menhaden fishery developed sporadically in the Gulf of Mexico until 1939. Landings exceeded 200 nnillion pounds for the first time in 1949 and the record catch of 1956 was more than 550 million pounds. Most of the Gulf landings come from Louisiana waters.
There are five North American species of Brevoortia. B, Smithi inhabits both Atlantic and Gulf waters. The Atlantic menhaden fishery is dependent on B. tyrannus and Gulf menhaden fishermen catch B. patronus,
Brevoortia tyrannus spawn at sea, or in high salinities, and the larvae move to low salinity areas which are essential to their development- Spawning time apparently depends on water temperature and occurs off the South Atlantic States in winter and in spring and summer farther north. Menhaden eggs and early larvae have not been reported from the Gulf of Mexico. Collections of larvae and juveniles indicate that BrevOOrtia patronus spawn in high salinities during the fall, winter and spring.
Larvae reach low salinity waters before metamorphosis. On the nursery grounds, body proportions change from the larval stage to the adult at about 30 mm. standard length, and the young fish move toward open waters of higher salinity in the summer and fall. The number of young on the nursery grounds varies greatly from year to year. No long migrations of menhaden are known in the Gulf. There are little data on the growth rate of Gulf menhaden.
Brevoortia patronus is known to feed by filtration and, in turbid estuarine waters, consumes considerable quantities of detritus and sus- pended bacteria in addition to living plankton.
Bluefish, mackerel, sharks and tarpon are the chief enemies of Gulf menhaden, and birds take a toll. The common shore and shallow- water sport fishes seem not to subsist to any great extent on larger menhaden, but predation upon small menhaden in the bays seem to be heavy. A few menhaden have been reported killed in the Gulf by sudden cold waves and the "red tide'* kills numbers of menhaden in Florida waters.
There is no evidence that menhaden are affected by any fatal diseases in the Gulf of Mexico but they are characteristically host to a number of worm and crustacean parasites.
Few fish other than menhaden are taken by menhaden fishermen.
XV A REVIEW OF LITERATURE ON MENHADEN WITH SPECIAL REFERENCE TO THE GULF OF MEXICO MENHADEN, Brevoortla Patronus GOODE
by
Gordon Gunter and J. Y. Christmas
INTRODUCTION
On the Atlantic coast of the United ships of the species of the genus States, nnenhaden have been utilized for Brevoortia. The Atlantic menhaden, food and for fertilizer since the days Brevoortia tyrannus (Latrobe), is dis- of the early colonists (Goode, 1879)- cussed under topic headings. The chief Processing the fish for oil, which in the papers are cited, but no attenipt was beginning consisted nnerely of press- made to cover completely the volu- ing the decomposing raw fish, was minous and sometimes repetitious started in Rhode Island in 1811 (Goode, literature. In the section on the Gulf 1884). In the 1870's the industry was menhaden almost every piece of lit- brought to North Carolina by northern erature pertaining to the biology of soldiers who had noted the abundance this species has been cited. Topical of menhaden in southern waters during arrangement of the material parallels the Civil War (Ellison, 1951). After that of the Atlantic menhaden. some initial failures the menhaden industry became well established in the South Atlantic States in the 1880's. GULF MENHADEN FISHERY STATISTICS Despite the long history of the menhaden fishery in the Atlantic and The development of a menhaden its rapid expansion in the Gulf of fishery in the Gulf States moved from Mexico, extensive studies of the east to west. There was considerable biology of these important commercial production of menhaden on the Florida fishes have not been undertaken until east coast as early as 1923 but con- recently. The only published bibliogra- tinuous production did not start in the phy on biology of the American Men- Gulf States until 1937 (Power, 1958). haden (Reintjes, Christmas, and The production statistics shown in Collins, I960) includes most of the table 1 were taken from various U. S. literature with very brief annotations Fish and Wildlife Service reports. and a subject index. This review of the There are some State statistical re- literature on the biology of menhaden ports, but these are not connparable brings together information from from State to State nor fronn year to somewhat scattered sources. Gulf year. They would be of little value commercial production is discussed except for the years 1946-47 in the first, followed by a summary of the developmental period of the Gulf men- taxonomy, distribution and relation- haden fishery.
Catch records given by Anderson Note.--Dr. Gordon Gunter, Director, and J. Y, Christmas. Fishery Biologist, Gulf Coast Research Lab- and Peterson (1953) show that men- oratory, Ocean Springs, Mississippi. haden were caught off the west Florida coast as early as 1880. From 1902 to be said that the Gulf menhaden fishery 1938 annual production in this area got started in 1939 with the location fluctuated between Z, 000 and 18,815,000 of plants in Mississippi. pounds and the annual average was During the World War II years 7,059.000 pounds for the 13 years re- Gulf production came from Florida ported. Alabama reported 10,000 and Mississippi. By 1948 plants had pounds in 190Z and 4,000 pounds in been located in Louisiana and Texas. 1931. Texas reported 14,118,000 in Landings did not exceed 200 million 1918 and 8,517,000 pounds in 1923. No pounds until 1949- other menhaden catches from the Gulf of Mexico were reported during the Table 2 shows the total catch, years 1880 to 1938. Therefore, it may yearly average and percentage catch
Table 1. --Landings of Gulf of Mexico menhaden by States, 1939-58
[In thousands of poionds]
West coast Total Year Alabama Mississippi Louisiana Texas Source Florida Gulf
1939 2,a49
19A0 267 1941-44
1945 7,166
1946-47
1948 ''No data collected) 1949 24,879
1950 1,534
1951 3,375
1952 10,737
1953 4,031
1954 2
1955 1,935
1956 32 1957 7 1958 9,103 by Gulf States for the years 1949-58, haden fishery more than doubled from inclusive. The Florida catch was neg- 1948 to 1958, the catch per yard in- to ligible, being slightly more than 1 creased from 13.9 thousand pounds percent of the total. No menhaden 19.8 thousand pounds in 1956. The landings were reported from Alabama. catch per yard of seine dropped to 1 1 .6 The major part of the catch, over 60 thousand pounds in 1957 and canne percent, was landed in Louisiana. How- back to 13.9 thousand pounds in 1958. ever, table Z does not fully show the preponderance of Louisiana waters in menhaden production. Statistics are MENHADEN BIOLOGY not available on catch localities, but Texas boats have little Texas water TAXONOMY, DISTRIBUTION in which to fish (Baker, 1955) and the AND RELATIONSHIPS Mississippi coast line is less than 70 miles long. Menhaden belong to a genus of clupeid fishes inhabiting the shore In 1950 the Gulf catch was 32.6 regions of the Atlantic from Nova percent of the United States total (table Scotia to Argentina and, according to 3). In 1955, although the Gulf catch de Buen (1958), Pacific waters off had increased to over 480 million Chile and Peru. Unverified records pounds, the Gulf percentage of the place the genus off the west coast of United States total catch declined to Africa. 25.8. Gulf fishermen took 26.7 percent of the record catch of 1956, when over The genus Brevoortia was es- 2 billion pounds were reported for the tablished by Gill (1861). Brevoortia United States total landings. Total tyrannus (Latrobe) was designated the landings dropped sharply in 1957. genotype. The genus was revised by Power (1959) listed several reasons Goode (1878) and by Hildebrand (1948). for the 1957 decrease. Although the According to Hildebrand (1948) there number of purse seines (table 4) con- were seven species, five in North tinued to increase in the Gulf, the 1957 America and two in South America. Gulf catch was down to 21.5 percent of Hildebrand had no North American total landings. records south of Indian River, the coastal lagoon of east Florida, and Little information on catch per none south of Tannpa on the west Flor- unit effort is available in the Gulf ida coast. The mouth of the Rio Grande menhaden fishery and the period of River was the southern limit in the time during which the fishery has western Gulf. Hildebrand was quite operated at a high level is only 10 firm in asserting, as a result of his years. While the number of yards of sojourn in Dry Tortugas, thit there seine (table 4) for the Gulf coast men- were no menhaden in southern Florida.
Table 3. — Total landings of Atlantic and Gulf of Mexico menhaden, 1950-57
[In thousands of pounds]
Year Atlantic Gulf Totals
1950 61^,^6S 326,030 1,000,498 1951 7-45,950 357,965 1,103,915 1952 923,606 459,984 1,383,590 1953 1,259,031 437,045 1,696,076 1954 1,336, 524 401,243 1,737,767 1955 1,386,791 480,992 1,867,783 1956 1,537,403 559,836 2,097,239 1957 1,327,595 362,533 1,690,128 Table K. — Number of piirse seines, number of yards of seine, and catch per yard of seine for the Gulf coast for various years
Number Number Catch per yard Year of of yards in thousands Source purse seines of seine of pounds
194-8. 34
1949.
1950.
1951.
1952.
1953.
1954..
1955.
1956. 1957. 1958. -
menhaden in the Gulf. Since only spe- new species, B. gunteri, which he con- cific names \were used in his paper, sidered to be a Gulf cognate of B, Gunter (1945) referred to E. t. pairOHUS smilhi. B. smithi and B. gunlcri agreed as B. tyranniis- Hildebrand (1948) also in having small scales, pointed yellow- noted that B. patroilUS had generally ish fins, more silvery color and less been considered by authors as only green along the back, and in being subspecifically distinct from B, tyvan- notably less slimy than B, lyVdlllluS and HUS or identical ^with it. However, B. patronus. B. glinteri differed from Hildebrand's diagnosis showed that B. smithi in having a larger head, B. patronus had fewer oblique scales, greater depth and longer maxillaries, fewer modified scales in front of the mandibles and pectorals. The ventral dorsal, fewer vertebrae, fewer ventral scutes and vertebrae were fewer in scutes and fewer pectoral and dorsal, number. Hildebrand (1948) examined and more anal rays. The lower caudal, specimens from Grand Isle, Louisiana; pectoral, maxillary and lower mandible Aransas Bay, Texas; and the mouth were generally longer. The head was of the Rio Grande, Texas. longer and deeper and body depth was greater. B. patronus was accorded full The fifth North American species specific rank in Hildebrand's 1948 of Brevoo^'tia is brevicaudata. Goode revision of BrevoOTtia. Meristic char- (1878) described the species from the acters indicated that B. patronilS re- only known specimens, collected at sembled B, tyrannUS of easternFlorida Noank, Connecticut, in 1 874. Hildebrand more than it did B. tyranrais farther (1948), after re -examining the se speci- north. Hildebrand (1948) noted that the mens, listed nine ways, including the lack of a decrease in number of ver- short caudal fin, in which they differed tebrae in southern menhadsn was con- from B, tyranmiS. Scalation was simi- trary to the general observation of lar to that of B. tyrannus. Bigelow and lower vertebrae counts for other fishes Schroeder (1953) questioned the cor- in warmer waters. rectness of these views.
Hildebrand (1941) described a The ranges of Brevoortia aurea small-scaled, nonslimy, yellow-finned (Agassiz) and Brevoortia pectinata menhaden fronn the coast of Carolina Jenyns in South America are not well
and nanned it Sniithi . Ho had pre viously known. (Hildebrand, 1920) called this fish Brevoortia aurea (Agassiz), which is a Hildebrand (1948) pointed out that South American species. The range the seven American menhaden could be was given as Beaufort, North Carolina, divided into a large -scaled group, B. to Indian River, Florida. Hildebrand tyratmiis, B. brevicaudata, B.patromis,
(1941) noted that B. smithi was not B. pectiiiata, andB. aurea , and a small abundant, did not school in large num- scaled group, B. smithi and B, gunteri. bers, and was called "yellowfin shad" He further stated that these menhaden by local fishermen. He said the fisher- could also be divided into two groups men kept fish of this species aside and according to the shapes of the ventral carried them home for food. Suttkus fins. Brevoortia tyramms, B. brevi- (1958) reported B. smithi from the caudata, and B. patronus had rounded eastern Gulf of Mexico between Cedar fins, while the other four had pointed
Keys and Placida, Florida, noting ". . . fins. The two South American species that Hildebrand was aware of the pres- differed from the others by less reduc- ence of Brevoortia smithi in the Gulf tion in size of the scales on the back of Mexico long before I made my inde- and on the base of the caudal as com- pendent discovery." pared to mid-scales on the sides.
Gunter (1945) noted a second spe- Hildebrand (1948) noted that with cies of menhaden in the Gulf which he the exception of B, brevicaudata, North said was close to if not identical with American species of Brevo&rtia could Brevoortia smithi of the Atlantic be divided into two closely related Hildebrand (1948) described this as a pairs on either side of the Florida peninsula. Species of sciaenids and in July and August. They said spawning clupeids, which are cognate on the east around Woods Hole took place from and west sides of Florida, were listed June to October. The Grampus (Bigelow to support Hildebrand's theory that the and Welsh, 1925) took eggs and larvae paired species of menhaden differ- in Nantucket Sound and west of Martha's entiated after the peninsula last rose Vineyard in October 1915. Bigelow and above the sea. Schroeder (1953) reported no eggs and larvae north of Cape Cod but said fry were taken in abundance in Casco Bay ATLANTIC MENHADEN in 1900.
Spawning Warfel and Merriman (1944) found menhaden larvae (less than 23 mm., Peck (1894) said minute organisms standard length) in the harbor of New furnish food for menhaden "not only Haven, Connecticut, in July and Sep- within the limits of these brackish- tember. The standard length of men- water inlets and estuaries where the haden collected in October was 30mnn. spawn is left to develop" but also in and a few 38-55 mm. in standard length more open waters, apparently having remained until November. come to this conclusion concerning spawning because small fish were Perlmutter (1939) reported eggs abundant in estuarine situations around and larvae at 27 of 52 stations in Long Woods Hole, Massachusetts, in the Island Sound. He said the spawning sunnmer of 1893. Peck also pointed season extended from May to October. out that low salinity areas were im- Westman and Nigrelli (1955) said men- portant "because they were intrusted haden in the zero year class appeared with so much embryonic and larval life in estuaries, tidal rivers, creeks, of the migratory inhabitants of the mooring basins and canals in June 1953 coast." Smith (1907) said that in New and remained until late September or England the spawning of menhaden took October. Eggs and larvae were not place in late spring and early summer, found in Great South Bay by Perlmutter and from Chesapeake Bay southward (1939) although zeros (young-of-the- spawning occurred in late fall and early year) more than an inch in length winter. Fishermen reported large fish were usually abundant there inshore. with full roe in Novennber and Decem- According to Westman and Nigrelli ber in North Carolina "rivers", but (1955) "The abundance of these young disagreed as to whether the spawning menhaden is difficult to imagine and, act took place only in the ocean (Smith, unlike the adult population they exhibit 1907). Schools of young were found in- no aversion to fresh water." Due to shore in winter. However, Smith was the long spawning season sonne of the not sure of the spawning location and young- of-the -year were 7 inches long said "there is some evidence" that in October and others were only 2 spawning took place both in the sea inches long. Evidently Westman and and inside waters. Nigrelli did not collect larval stages. Kuntz and Radcliffe (1918) found some fish ready to spawn and some Radcliffe (Kuntz and Radcliffe, spent in Woods Hole harbor in August. 191 8) took larvae 30 mm. long at the Eggs and larvae were occasionally nnouth of the Potomac in February 1914. taken there during the summer and According to Hildebrand and Schroeder were abundant off Gay Head in August. (1928), menhaden in the Chesapeake Larvae 20 mm. long were taken in the Bay region seemed to spawn in the fall. harbor on October 21, 1914. These Larvae averaged 27.7 mm. long in authors indicated that the main spawn- January, 33.5 mm. in February, 27.3 ing period in the Woods Hole area was mnn. in March, 33.0 mm. in April, and June, July, August, "and later". 46.0 mm. in May. Pearson (1950) took Bigelow and Welsh (1925) found menhaden larvae at Old Point Comfort, spent and ripe fish in the Gulf of Maine Virginia, during April and May. Massman, Ladd and McCutcheon ports of menhaden eggs correspond (1954) caught young menhaden in the with high salinity situations. brackish and fresh waters of four Virginia Rivers, up to 27 miles above brackish waters, in April and May. Growth These fish were collected in plankton tows and ranged from 24 to 30 mm. Kuntz and Radcliffe (1918) re- standard length. Ninty-eight percent of ported that menhaden eggs were 1.4- the 8,000 specimens were found at 1.6 mm. in diameter. Bigelow and salinities of 2-3 %o. The ratio of num- Schroeder (1953) gave the measure- bers caught in surface and bottom tows ments as 1.5-1.8 mm. and said men- was 200 to 1. In June, menhaden of haden produced the largest fish eggs 40.6 mm. in mean standard length were to be found in the Gulf of Maine. the most abundant species in the shore zone just above brackish water. Young McHugh, Oglesby and Pacheco fish were present in July in the same (1959), after studying the Chesapeake area. Surface trawls showed essen- Bay pound-net and purse-seine fish- tially the same pattern in September eries, assumed that, in the Chesapeake when fish averaged 94 mm. standard Bay area, menhaden left the Bay as length. they approached maturity in late Sep- tember, October and November. Spawn- Ellison (1951) quoted Hardcastle ing apparently occurred outside the (MS) who concluded that, in North Bay in the fall and continued through Carolina, menhaden probably spawn in the winter. Bimodal distribution of late winter near the Gulf stream. length and weight frequencies in age- Ellison (1951) said the young fish have group suggested that two spawning wide salinity tolerance and live in peaks may occur in the Chesapeake slightly brackish waters or waters with region. the salinity of sea water, and quoted Westman and Bidwell (1948) who re- Welsh (Bigelow and Welsh, 1925) ported young 45 miles up the Hudson found experimentally that hatching took River and in Long Island Sound at the place in less than 48 hours and same time. that the larvae are 4.5 mm. long at that time. Four days after hatching they Bigelow and Schroeder (1953) said were 5.7 mm. long. There are no other that W. W, Welsh concluded that sexual data on the time elennent and growth maturity in menhaden is attained by of the larvae. The dorsal and caudal the third year of life. Westnnan and fins were visible in the 9 mm. stage Nigrelli (1955) wrote that all menhaden and all fins were differentiated at the 3 years of age or more appeared to be 23 mm. stage. At this stage the larva adult fish. was long and slim; when the young menhaden had reached 33 mm. in Goode (1884) reported that 150,000 length it had taken on a more fish-like eggs were found in one menhaden. appearance and acquired scales. Ellison (1951) quoted Hardcastle (MS) who found that 41 percent of June and Chamberlin (1959) dem- mature and immature males were in- onstrated that low salinities are essen- fested with a gonadal parasite, Eimevia tial to the normal development of brevoortia, a sporozoan. The effect of Atlantic menhaden larvae. Larvae and this infestation on sperm production metamorphosing individuals from low- was not determined. salinity areas invariably perished when transplanted to high-salinity rearing Perlmutter (1939) is the only au- ponds, although individuals trans- thor who has reported salinities at planted a few weeks after metamorpho- which menhaden spawn. He gave the sis survived. Similar phenomena were figure as 84-100 percent (30.5 - 35%o) observed in natural conditions on sev- sea water, at 55°-80°F. All other re- eral occasions. Larvae reared through .
metamorphosis in high salinities de- taken along the east coast of the United veloped abnormalities while those de- States between Portland, Maine, and veloping at low salinities were normal. Fernandina, Florida, from 1952 through 1956. Scale formation commenced at Hildebrand (1948) has noted that body lengths between 24 and 30 mm., all species of young menhaden (aureo appearing first in the region of the and smithi not verified) up to 70 mm. caudal peduncle and later near the long have minute teeth on the margin base of the pectoral fins and along the of the maxillary which are subsequently posterior margin of the opercles. lost. Scales were fully formed at body lengths between 30 and 43 mm. Growth of Welsh (Bigelow and Welsh, 1925) scales of young -of -the -year was traced studied large numbers of menhaden fry from May until mid-September. The and their scales in the Gulf of Maine. first age ring was formed sometime Summer hatched fish were found to be after the first summer of life.
6-8 cm . long the first winter and 16cm. long the second winter. Bigelow and Ring formation occurred only once Schroeder (1953) reported young men- each year, between March and May. haden 9.1-9.9 cnn. long at Woods Hole Marginal scale growth occurred during in late September 1942. Fish taken at the warm months of the year. The dis- Falmouth, Massachusetts, 2 months tance between the last submarginal ring later were 11.7-12.7 cm. long. Welsh and the margin of the scale reached a (Bigelow and Welsh, 1925) found as maximum in the fall. The distance many as nine to ten "winter rings" on between the last two adjacent ring the scales of a few of the older fish modes decreased with age. In general, examined and apparently thought that ring formation was found to occur each ring represented 1 year's growth. earliest in southern waters and prog- ressively later farther northward. New Rush (1952) found that fish with one scale growth varied considerably annulus at Beaufort, North Carolina, among individuals, especially in the were approximately 18.9 cm. long (fork younger age groups, and this variability length). Three-year fish were 28.7cm. is greatest in those fish occurring in long. The oldest fish, 6 years, was southern coastal waters. 35.8 cm. long. The growth curve was worked out on 34 specimens. Westman Hildebrand and Schroeder (1928) and Nigrelli (1955), who worked on said that sexes could not be differ- Long Island Sound fish, stated that entiated externally and that the growth growth rates determined "upon the rates were apparently much the same. basis of many times that number of However, Westman and Nigrelli (1955) specimens" were very similar. Fish analyzed the matter statistically and showing a single annulus averaged found that there was a significant dif- 18.4 cm. fork length. Fifty-two males ference, the females outgrowing the of age class III ranged from 29-5 to males 34.0 cm. fork length and 52 females of the same age class ranged from 28.5 Westman and Nigrelli gave fork- to 34.5 cm. long. In age class IV 17 length frequency curves for 1/2-month males ranged from 31.5 to 35.0 cm. periods from April to June for fish long and 55 females had a fork length from Long Island Sound and New Jersey range of 31.5 to 36.0 cm. They stated waters. In general, the curves showed a that the conversion factor from fork to decreasing mode from about 30 cm. in total length was 1.1163. Scattergood, April to 25 cm. in June during 1948 Trefethen and Coffin (1951) reported and 1949. the conversion factor from standard to fork length to be 1.0459. Ages of menhaden up to the third year of life may be determined by the June and Roithmayr (I960) re- length-frequency method or by reading ported results of readings of 13,510 scales (McHugh, Oglesby and Pacheco, scale samples of Atlantic menhaden 1959). Two groups offish, characterized by their size at the^time of formation New York, weighing 1 pound 13 ounces. of the first annulus, entered the Vir- Goode (1879) said the largest specimen ginia fishery in 1954 and 1955 suggest- in the National Museum was a 20-inch ing that growth rates differ according cast. These remarks donot correspond to circumstances. Mean lengths at the to those of Ellison (1951) concerning end of each calendar year of life were the abundance of 16- to 20 -inch fish in found to be approximately 125, 200 and North Carolina waters in late Novem- 230 mm. Mean weights were about 30, ber. 130 and 200 g. Menhaden reached maturity at about 1^ years. June and Reintjes (1959 and I960) presented extensive tables of length and weight frequency distributions of sam- Hildebrand and Schroeder (1928) ples of Brevoortia tyrannus from the said that fish varied in fatness during purse -seine and pound-net fisheries the year and were much fatter in the along the Atlantic coast from 1952 fall than in spring. They also noted through 1956. Comparison of the aver- that fatness, and presumably growth age length and weight of individual year rate varied from year to year. Fish, classes from the middle Atlantic area taken in October 1921, 13.0 cm. long at different ages showed rather marked averaged 0.78 oz. in weight, and two variation in fish of comparable age in specimens of the same length, taken a the different years. The average fork year later, averaged 1.07 oz. In 1921, length of age 1 fish ranged from 209 October fish 23.1 cm. long averaged mm. in 1952 to 229 mm. in 1955; of 4.22 oz. and a similar sample for 1922 age 2, from 233 mm. in 1952 to 259 rnm. averaged 5.17 oz. The October 1922 in 1955; of age 3, from 279 mm. in fish weighed an average 15.5 percent 1955 to 286 mm. in 1956; and of age 4, more than fish of the same length taken from 291 mm. in 1955 to 304 mm. in in October 1921. Hildebrand and 1954. Five-year-old fish averaged 311 Schroeder also gave some tables on mm. fork length in 1956. The most weight-length relationships. Fish 10.2 striking feature was that in every year cm. long weighed 0.35 oz., those 20.3 through 1955 fish of the dominant 1951 cm. long weighed 3.59 oz. and 35.6cm. year class averaged lighter than ad- long weighed 17.1 oz. jacent year classes. In general, there was an increase in the size and weight Goode (1879) measured 30 fish of fish of the same age group from from the same school taken at Noank, south to north. (It should be noted, how- Connecticut, in 1875. They ranged from ever, that the fishery starts earlier 12 to 13 inches in length and 10 to 16 in the south than it does farther north.) ounces in weight. The average length The data show that fish of the domi- was 12.38 inches. nant year class which contributed to the catch in each area in 1956 were Concerning seasonal changes in shorter and lighter than fish of the fatness, Bigelow and Schroeder (1953) same age in 1955. Females were larger said that menhaden were always thin than males at older ages. when they arrived in the Gulf of Maine from the south, but put on fat rapidly. In 1894 the yield of oil rose from 12 Distribution and Migrations gallons, and less, per thousand fish in the early part of the season to 14^ in Bigelow and Schroeder (1953) re- August to 16 and 18 gallons in Septem- ported that spawning probably took ber. They stated that New England fish place every year from the Long Island were larger and fatter than those tothe Sound area to Florida during the period south. These authors also said that the of their observations, and that in some longest menhaden of their knowledge years it extended into the Gulf of was a specimen 18 inches long taken Maine. The eggs were buoyant, accord- at Woods Hole in 1876, but that 29-inch ing to Welsh (Bigelow and Welsh, 1925) fish had been reported. The heaviest and were found at the surface in waters fish reported was one from Orient, of high salinity (Perlmutter, 1939). --
The larvae moved towards shore, remained in the deeper parts of Chesa- where they reached sheltered and often peake Bay during the winter in reduced low-salinity areas at a size of around numbers, where they were taken in Z5 mm. standard length. Probably they beam trawls; in March the fish came were mostly at the surface during these into shallow waters and were then taken movements. There were Sonne indica- in pound nets and haul seines. The tions that the larvae moved inshore large schools migrating up and down shortly after coming into the estuaries the coast did not enter the Bay and the and moved back out again as juveniles local fishery was not affected by spring, (Massman, Ladd and McCutcheon, summer and fall "runs" as was the 1954). Apparently this movement con- case with the outer shore areas of the tinued on out into open waters, for in Atlantic States. December Z-inch fish, the zeros of Westman and Nigrelli (1955) showed According to Bigelow and Schroe- up regularly in North Carolina mixed der (1953) nnenhaden left the Maine in with 10-inch fish (Ellison, 1951). coast by the middle of October and Presumably the 2-inch fish came from Massachusetts Bay by early Novennber, the north, possibly Chesapeake Bay, although small ones had been taken because the North Carolina zeros there as late as December. Reports should not be that size in December from fishermen indicated that the fish (Hildebrand, 1920). went around Cape Cod to the New York region and on southward. Ellison (1951) McHugh, Oglesby and Pacheco wrote that the migrating schools were (I959) found that menhaden less than 2 fished from Delaware south to North years old were present in Virginia Carolina as follows: waters throughout the year. Spawning probably took place in the ocean, and "About October 15 a run of fish the young moved into Chesapeake Bay appears in North Carolina from the and its estuaries soon after hatching. north and is joined by fish from the As they grew, young menhaden moved southern sounds and estuaries. These back down the estuaries toward the fish run from 10 to 12 inches in sea. In most seasons mean sizes of length and are known locally as fish were progressively greater in a 'Chesapeake Bay' fish, 'holy jump- seaward direction. Since the purse ers' or 'forerunners.' They contrib- seine fishery exploited fish primarily ute to the fishery about a month and in their second and third year of life are followed about November 10 by and the pound-net fishery took prin- the so-called 'Delaware' fish, which cipally fish in their first and second measure 13 to 16 inches. These fish, year, sampling of the pound-net fishery in turn, are succeeded about Thanks- offered a method of forecasting purse giving by 16-to-20 inch fish recog- seine catches at least a year in advance. nized as the 'Boston Bay' or the Seasonal waves of emigration and im- 'Amagansett' fish. All of the fish migration to Chesapeake Bay were appearing from October 15 to No- associated with intense fall and spring vember are following a north-south spawning periods. Females were migration route." slightly larger than males of the same age, but the difference was insignifi- In December schools of small fish cant in young menhaden. 2 to 10 inches long appeared in North Carolina. Their source was unknown, There is some information con- but they probably came from inshore cerning the movements of B. tyrannus waters. up and down the Atlantic coast from observations made by biologists and Concerning the spring migration fishermen. Apparently no menhaden northward Ellison said: remain in offshore waters north of Chesapeake Bay during the colder "The spring fishery usually starts months of the year. Hildebrand and in May, although sometimes in April. Schroeder (1928) found that nnenhaden This fishery depends principally upon
10 individuals which run from 6 to 8 than 50° F. Bigelow and Schroeder inches in length and which are be- (1953) said their observations cor- lieved locally to come up from Flor- roborated the idea that menhaden did ida. Usually these fish strike shore- not appear in the Gulf of Maine until ward about the latitude of Fernandina, it was several degrees above 50. Con- moving north and paralleling the versely, falling temperatures drove the coast, supplying a good fishery at fish southward in the fall. Mayport, Florida. For the past four years, however, they have scarcely Edwards (Kendall, 1910) found that touched Mayport, and snapper fisher- young menhaden could survive cold men working 30 miles out have re- better than their elders. Fish 2 to 5 ported great schools moving north. inches long survived water tempera- In these recent years they have tures of 31.5° F. Better survival of struck first off the South Carolina young than of older fish at low tem- coast about Georgetown and are peratures was independently observed called in North Carolina the for other fishes by S. F. Hildebrand 'Georgetown-flats' fish. These fish and Gordon Gunter following cold spells support the fishery in North Car olina on the south Atlantic and Gulf coasts. until August, when they disappear." The literature was documented by Gunter (1957). Ellison further stated that the fish -eached the Chesapeake region in In their long migration from the March and April, the New Jersey and south Atlantic coast to New England New York region in April and May, and and back, menhaden do not follow pre- the Maine coast in May and June. These cise routes. A "run" may strike shore dates are much the same as those for many years and then suddenly given by Goode (1879), who reviewed change and pass as much as 50 miles all available information up to that offshore. Such changes in migration time. Goode's data included precise routes may leave established menhaden dates for a 20-year period from some plants fishless for several years. Elli- areas of the Atlantic coast. He did not son (1951) has given examples. know whether fish went south in the winter or merely offshore. June (1958) studied the variation in meristic characters of young Menhaden do not always go as far Brevoortia tyrannus to determine north as the Gulf of Maine in summer, whether one or more populations occur and combined data from Bigelow and in estuarine nursery areas along the
Welsh ( 1 925) and Bigelow and Schroeder Atlantic coast. Vertebrae, ventral (1953) showed that between 1845 and scutes, dorsal, and left pectoral fin 1950, inclusive, menhaden had been rays were the meristic characters plentiful to abundant there during only selected for examination. Counts were 15 of 66 years reported. made on almost two thousand speci- mens taken from 21 locations between It has been generally agreed that Cape Cod and southern Georgia. temperature governs the north and south migrations of menhaden and that Preliminary studies indicated that menhaden do not enter waters of tem- mean counts do not vary with sex, peratures less than 50° F. (cf. Ellison, length of fish over the size range 1951). Goode (1879) collected tem- (28-150 mm.) represented, or between perature records along the coast and right and left pectoral fins. Analysis compared them with the time of ap- of variance applied to the meristic pearance of the menhaden. This infor- data gave no evidence of heterogeneity mation led him to state that menhaden between the means of successive sam- appeared after the water temperatures ples taken in any locality. rose to 50° to 51 F. and preferred temperatures between 60° and 70°. At least two populations of juvenile Bean (1903) noted that adult menhaden menhaden inhabiting the estuarine in aquaria died at temperatures lower waters of the Atlantic coast of the
11 United States were indicated, one oc- Peck's chief conclusion, which he curring north of Long Island and another documented by drawings of plankton in south of Long Island. Internningling the microscope field, was that menha- occurred in the vicinity of Long Island. den are indiscriminate feeders, and in Difference in mean water temperature general take in the same materials in during the time of spawning and early the same proportions as they are found larval development may account for the in the water. In the shore waters around observed differences in mean meristic Woods Hole, which Peck studied, these counts between the two areas. materials were predominantly dino- flagellates, diatoms and infusorians, June and Reintjes (1959) noted that with annelid larvae and crustaceans in the purse-seine fishery depended on lesser abundance. Peck worked mostly the appearance of schools of menhaden with inshore and smaller fish, but at the surface. Fishing started in April found that the young ate the same food on the Florida coast, and by June the as the adults. fish ranged from northern Florida to the Gulf of Maine. In September and Peck estimated the volume of water October the schools began todisappear strained by an adult menhaden and from the most northerly areas, and arrived at a figure of 7 gallons per withdrawal proceeded southward during minute. He estimated the food material October. In November and December in this amount of water to be 3.4 cc. vast bodies of fish supported a sizable fishery off the North Carolina coast until early January when they once Hildebrand and Schroeder (1928) more disappeared fronn coastal sur- summiarized the findings of Dr. Edwin face water. Linton who examined the alimentary contents of 44 fish in Chesapeake Bay, "and found that in most cases they Food and Feeding consisted of sandy mud, vegetable debris (mostly algae), and sonne dia- According to Goode (1879) a num- toms, and in a few cases they consisted ber of previous workers had observed principally of copepods." minute organisms and the presence of chlorophyll in menhaden stomachs. The following remarks are taken However, other workers he quoted, in- from Bigelow and Schroeder (1953): cluding A. E. Verrill, thought such matter was sucked up with the bottom "No other Gulf of Maine fish has mud. Goode called attention to the fine a filtering apparatus comparable to gill-rakers and the fish's manner of that of the pogy, nor has it any rival swimming with open nnouth, as if in the Gulf in its utilization of the straining the water. planktonic vegetable pasture.
The definitive work on feeding and ". . . And the food eaten at a given food of the nnenhaden was done by locality parallels the general plank- Peck (1894). He showed how the lamel- ton content of the water, except that lae of the gill-rakers overlap "each none of the larger aninnals appear in other in the most perfect manner" and the stomachs of the fish on the one form a fine net for sieving the food hand, nor the very smallest organ- from the water. He described a fold of isms (. . .) on the other." mucous membrane, filled with mucous, which runs along each gill arch at the Schroeder (Hildebrand and Schroe- base of the gill-rakers to form a der, 1928) observed that feeding fish channel and, presumably, acts as a swam in circles, rising and falling, groove to carry food into the gullet. renninding him of a whirlwind. Peck's statement, "Whether there are definite ciliated tracts with this func- Ellison (1951) said the "catholic tion of conveying solid particles is not and non-discriminating taste distin- yet known," still holds true. guishes the menhaden as strongly
12 exceptional, if not unique, among fishes attacked by gulls and other sea birds, of the sea." and the osprey. In fact Goode men- tioned every predator listed in later accounts, including gars and catfish in Enemies and Mortality southern waters. He gave varied ac- counts of attacks by bluefish, and said The following succinct account is the menhaden when pursued "often quoted fronn Bigelow and Schroeder drive in great masses upon the shores ." (1953): Goode thought the number destroyed by predators was many times that taken "No wonder the fat oily menhaden, by man. The figure he gave, 3,000 swimming in schools of closely million of millions, equals 3 quadrillion ranked individuals, helpless to pro- and is certainly a vast overestimate. tect itself, is the prey of every predaceous animal. Whales and por- Ellison (1951) mentioned that blue- poises devour them in large num- fish sometimes ran menhaden into the bers; sharks are often seen following shallows in such numbers that they died the pogy schools; pollock, cod, silver and piled up in windrows 2 feet deep, hake, and swordfish all take their toll and polluted the air. Reports of this in the Gulf of Maine, as do weakfish phenomenon extended from Hatteras south of Cape Cod. Tuna also kill to Maine. Local health boards were great numbers. But the worst enemy sometimes required to dispose of the of all is the bluefish, and this is rotting fish. According to Bigelow and true even in the Gulf of Maine Schroeder (1953) dead fish drifted during periods when both bluefish ashore along Massachusetts Bay in 1946 and menhaden are plentiful there. . . and 1947, and the local health boards Not only do these pirates devour had to clean the beaches. However, this millions of menhaden every summer, kill was attributed to netting by lobster but they kill far more than they eat. bait fishermen. Besides the toll taken by these natu- ral enemies, menhaden often strand in myriads in shoal water, either in Westman and Nigrelli (1955) made their attempt to escape their enemies the following statement concerning an- or for other reasons, to perish and nual kills of menhaden in New York: pollute the air for weeks with the ." stench of their decaying carcasses "Menhaden appear in the New York bight in commercial quantities Hildebrand and Schroeder (1928) during late March or April and, said that bluefish migrate up and down according to reports received, the the Atlantic coast following schools of large fish precede the smaller fish menhaden and other fish upon which by several weeks. During the latter they feed voraciously. The abundance part of May, or early June, great of menhaden governed the movement of quantities of menhaden die in these bluefish to some extent. In 1922 young waters and in the waters of western menhaden were plentiful in the lower Long Island Sound and litter the half of Chesapeake Bay, and the com- beaches in untold numbers. Conse- mercial catch of bluefish was greater quently, various agencies call other than it had been for many years. agencies each year, or write letters, about this phenomenon. These agen- Ellison (1951) said the menhaden cies, in turn, communicate with other is "prey to virtually all of those car- agencies which, in turn, may refer nivorous fishes which inhabit the same the problem back to the first agency, waters." and so on."
Goode (1879) was much impressed Dying fish were called "spin- with predation upon menhaden. He said ners". They were characterized by whales and dolphins ate them by the uncoordinated movements and exop- hogshead. From the air they were thalmia. Westman and Nigrelli found
13 hemorrhages caused by gas emboli in Parasites and Diseases capillaries of the gills, eyes and optic lobes of the brain. Bacteria could not Viral and bacterial diseases are be found and the fish entrails were unknown in menhaden, and there is no not harmful to cats. Whether the gas evidence that they are affected by any emboli were the terminal cause of fatal disease of epidemic proportions. death or were only an associated con- On the other hand menhaden are char- dition could not be determined. In any acteristically host to a number of worm case, fish not in distress did not have and crustacean parasites, which may gas emboli. The authors noted that all cause debility but not death. mortalities occurred in areas with "markedly varying salinities" and with The following parasites of the "organic pollution to a degree where Atlantic menhaden were listed by West- shellfishing is prohibited". man and Nigrelli (1955):
Parasite . .
was 0.336 percent. Most of the clupeids January 15, 1913 by W. W. Welsh in other than menhaden were taken in a St. George Sound, Carabelle, Florida. few sets when mistaken for schools of St. George Sound is on the northwest nnenhaden. Florida coast and the specimen was doubtless B. patronus Smith also noted that "the men- haden is a fish which, as a rule, is Gunter (1945) reported eight found in comparatively close proximity Brevoortia patronus 28-59 mm. total
to the land. . ." Most of the fish were length taken on the Gulf beach of taken less than 5 miles from shore. Mustang Island, Texas in March and April. Several larvae, tentatively iden-
The chief predators were found to tified as Brevoortia , taken at the same be bluefish, sharks and weakfish. time, were presumed tobe B.patrotlUS- Migratory movements were oftenmodi- fied by the presence of "such pre- Baldauf (1954) listed menhaden as daceous species as bluefish, sque- being one of the fishes which are to be teague, and sharks" (Smith, 1896). The found in the lower Neches River only autumnal migratory movement began "during growth stages". Menhaden with the fish on the shores of Maine were taken in the Neches River at a and Massachusetts and continued south- station approximately 30 nautical miles ward. upstream from Texas Point, at the Sabine Pass entrance to the Gulf. Larger and fatter fish were taken Salinities at stations where menhaden on the New England coast than from were taken ranged from 0.16 to20.4%o. areas to the south. New England fish Surface temperature varied from 14.4° produced more oil in the fall than those to 31° C. Over 10,000 specimens were from the same area produced in the taken in 1951, 1952 and 1953. The spring. Examination of ovaries tended largest menhaden measured was a to verify the existence of different specimen, found dead on the river bank, spawning periods on different parts of with a standard length of 150 mm. The the coast. larger fish were B. patronus but spe- cific identification of young individuals was uncertain. Table 5 gives Baldauf's GULF OF MEXICO MENHADEN measurements.
Spawning Baldauf noted that two incoming populations, November-December and Kuntz and Radcliffe (1918) noted March-April, were not the same and that the first menhaden larva identified that there may have been two spawning was a specimen 24 mm. long taken on peaks.
Table 5. — Numbers and sizes of small menhaden taken in the Neches River, 1952-53
[Based on Baldauf, 1954-]
Ranges Date Number Mean ( standard length)
July- Aug, 6 15 Reid (1955a) carried on an inten- 1956 following the closure ofthepassat sive study of East Galveston Bay, Texas Rollover. He concluded in general that from June 1 to July 9, 1954. Young physical characteristics and fauna of B. patvonus were abundant in the bay, the bay had reverted to the 1954 state, and almost 21 percent of seine collec- before the pass was opened. However, tions from the shallow shore zones and "the staggering number of menhaden in about 2 percent of the trawl collections the littoral zone during the 1956 study were this species. Adults were uncom- is not explained." Trawl hauls were mon in the bay. No BrevOOrtia were comparable to previous years, yielding caught in six strikes with a trammel 1,000 menhaden, but 63,000 were taken net. Little menhaden, ranging "in size in seines. The same stations were from 27 to 114 mm., with most of the occupied as in previous years with fish between 27 and 60 mm." (Reid, additional hauls being made in June. 1955b) ranked second in number in The fish ranged from 18 to 60 mm. minnow seine catches and sixth in standard length with a mode at 38 mm. trawls. Gunter (1956a) reported on larval In 20 of 30 trawl hauls, 349 B. and postlarval B. patronus taken in patrotlllS were caught (Reid, 1955b). In White and Grand Lakes, tributary to 10 of 14 seining operations, l,539men- Vermilion Bay in Louisiana. Table 6 haden were captured. Most fish were shows salinity ranges at which men- taken in the low- salinity part of the bay, haden were caught. These fish were at 4-12%o. Some were taken in plankton around 1 inch long. The length frequency tows. Reid (1955a) suggested that the data were lost. The menhaden con- menhaden is a fish spawning offshore, sidered in Gunter 's paper were nearly and that the young come in for only all taken in minnow seines on shore and a period of time. approximately 1 ,800 of them were taken Reid (1956) found that the percent- in February 1952 and January 1953 age of menhaden greatly increased in (Gunter and Shell, 1958). trawl catches in East Bay after a pass Suttkus (1956) said the movement to the Gulf was cut at Rollover, but of larvae of B, patronus , 20 to 30 greatly decreased in seine catches. The mm. standard length, into Lake trawl catch percentage for the 2 years Pontchartrain started in December in was 1.8 and 11.8; for the seine it was 1953 and 1954 and continued into March. 20.7 and 1, respectively. The greatest He assumed that spawning began in number of menhaden were taken in the October and ceased in February. The upper bays. The size range was 21 to text indicates that young fish of the 141 mm. standard length, with the year generally left the area in August greatest number lying between 28 and to October, but according to tables 64 mm. The collections were made in presented in the paper this was not the June 1955. case in 1954. Reid (1957) reported further At the time Gunter (1945) worked studies of East Bay, Texas, nnade in in Copano and Aransas Bays, what he Table 6. --The salinity ranges where small Brevoortia patronus ^yere caught in Grande and White Lakes in Louisiana [Based on Gunter, 1956a] Number Number Average Salinity (^o) caught hauls per haul 0.13-0.29. 10 8 1.3 0.^1-0.93. 1,038 15 69.2 l.U-2.70. 1,175 13 90.4 16 called Brevoortia sp., later named specimens ranged from 20.1 to 31.0 gunteri by Hildebrand (1948), was the mna. standard length, with the average most abundant menhaden in the bays. size, 23.5 mm., coinciding with the mid- Of 1,231 menhaden caught, 66 percent class of the modal frequency. An aver- were taken in Copano Bay and the age standard length of 27.0 mnn. was remainder in Aransas Bay, with one recorded for 400 specimens, 17.5 to batch, probably of the same species, 47.5 mm. standard le'ngth measured in taken on the Gulf beach. Fish largely April. Standard length of May speci- left Copano Bay in midwinter and went mens ranged from 23.5 to 74.5 mnn. into Aransas Bay. One male and one with an average of 29.1 mm. All col- female, with running milt and eggs, lections from February through May were taken in the bays in February and were from Cross Bayou Canal between March. The salinities and tempera- Old Tampa Bay and Boca Ciega Bay tures where these two fish were caught where salinities ranged from 6.6 to were 12.8 and 13.1%o and 10.5° and 18.7%o and temperatures were 20.5° to 14.0° C, respectively. The salinity 27.5° C . During July, a series of speci- range at which this species was taken mens, average standard length 85.1 was 2.0-33.7. The temperature range mm, were taken at Johns Pass, a cut was 9.10-31.0° C. Postlarvae thought between Boca Ciega Bay and the Gulf. to be B. gunteri (as later designated) 21-30 mm. total length appeared in Springer and Woodburn reported Copano Bay in February 1942. They B, smithi and B. patronus were taken were most abundant in April and could together in May. The average size of be followed until May when they were B, smithi, 23.3 mm., in May was about 25-45 mm. total length. A few appeared the same as that of B. patronus in in upper Aransas Bay in April and March. Additional specimens of B. May. All individuals in a school were smithi were reported for the months of near the same size. Fish more than January, June, July and August in 300 mm. in length were taken only in salinity and temperature ranges of June, October and February. The great- fresh water to 31.6%o and 18.0-30.3°C. est abundance was in April and May. Young menhaden were seen at a Gulf The larger fish were most abundant in beach station in March. the lower bay. Smaller fish were taken in Copano Bay from February to Springer and Woodburn (I960) in- August, in Aransas Bay only in April dicated a winter spawning period for and May. Larger fish were in general B. patronus and a somewhat later, found in the higher salinities. The im- spring spawning period for B, Smithi. pression was gained that the spawning season was in late winter and spring. Growth Simmons (1957), referring to the Laguna Madre of Texas, said: "Men- Gunter (1938b), in an analysis of haden, particularly Breyoorija gunteri, species taken in shrimp trawls in were common in waters of moderate Barataria Bay, Louisiana, and adjacent salinity and present in waters of high Gulf waters, wrote that B. patronus salinity. Menhaden definitely spawned ". . . was caught much more in the bay in the area in February 1956. Indi- than outside waters. Individuals from viduals from 15 mm. up were taken in inside were mostly immature and it is March, April and May . " B oth B. ^nteW possible that adults were present in and B. patronus were listed among ani- larger numbers in the Gulf. During the mals commonly taken at salinities months of June and July in 1932 and from 20 to 60 but rarely above. June 1933 the menhaden was scarce inside and was taken more commonly Springer and Woodburn (I960) re- in the Gulf. In 1932 there was a peak ported on B, patronus taken in the in the bay in January. The following Tampa Bay, Florida, area in 1958. One year the peak came in December and specimen, 22.2 mm. standard length was January and in 1934 it came in January collected in February. In March, 569 and February. This is ample data to 17 . establish the fact that there is an "Combined total length-frequency abundance peak for this fish in the bay curves for both bays were made. In during the winter, which is characteris- April, 1941 the only group of men- tic of the life history of the species. . . haden caught were from 88to 133 mm. The increased numbers caught in mid- long with a mode at 1 1 3 and 118 mm winter. . . are possibly a result of mi- This group persisted until October gration of such individuals from nurs- when it was from 128 to 173 mm. ery grounds near the shore to the open long. In August a second group from bay. They grow rapidly and probably 88 to 103 mm. long appeared, which by midsummer nnost of them pass to the became predominant in November outside as the curve indicates. Fish and December. Remnants of the two taken on the outside were usually adult groups persisted through the winter or nearly adult in size. Smaller indi- until May when they became more viduals were sometimes caught near numerous again. At that time, the the shore." smaller group had a mode at 128 to 133 mm. Post-larvae came into the Baldauf (1954) presented monthly catch in January and they were pre- standard length frequency curves for dominant from then until May, when Brevoortia sp. for November 1952 their size ranged from 23 to 43 mm. through April 1953. The November in length." mode was about 19 mm. and could be followed to a mode of Z5-26 mm. in Springer and Woodburn (I960) March. compared data from collections in the Tampa Bay, Florida, area with Suttkus' Suttkus (1956) gave numerous pro- (1956) Lake Pontchartrain data and portional measurements of young stated that B, patronus reached Tampa B. patronus. These show that the body Bay inland waters. . ."at a smaller shape changes greatly between 20 and size (earlier age?) than it does on the 30 mm. standard length. The meristic Louisiana coast." It should be noted counts fell in the lower ranges of that Springer and Woodburn's collec- Hildebrand's (1948) figures. Suttkus tions were made close to high salinity said serrations on the ventral part of Gulf waters while the inland waters of the maxillary, which disappear with Lake Pontchartrain are many miles age, were not mentioned by Kuntz and from the open Gulf and high salinity Radcliffe (1918) or Hildebrand (1948). waters where spawning takes place. However, Hildebrand called them The data from both areas indicated "teeth", said they were present in all ". . .a sudden spurt of growth after species oi BrevoOTtia, and disappeared May (after June?)" by the difference at about 70 mm. length. between the average size of July sam- ples. The growth rate of Florida fish Renfro ( 1 958)' took five B. />a/roWJ« seemed to be slower than fish from in the Aransas River, Texas, in March Louisiana. and July 1957. One, in March, was 120 nnm. long and was in a salinity of 54.3%o. Another, 32 mm., was taken at a salinity of 47.6. Three others, 52-64 Distribution and Migrations mm. long, were taken in July at 0.5%o salinity. According to Goode and Bean ( 1 879) the first menhaden from the Gulf re- Gunter (1945) in a discussion of ceived by the U. S. National Museum Brevoortia sp. (B. gunteri Hildebrand, were collected in West Florida in 1 864. 1948) taken in Aransas and Copano Goode (1879) under the heading of Bays, Texas, said: "Limits in 1877" quoted two light- house keepers, along the Mississippi and Texas coasts, one of whom was a 'Renfro, W. C. 1958. The effect on of salinity the former Maine pogy fisherman, who distribution of fishes in the Aransas River. Master's Thesis. Graduate School, The University of Texas. 51 p., said that no menhaden "is found in 3 tables , 25 figs. those waters." However, he quoted a 18 -,. pilot, Capt. Wm. Nichols, who said Fowler (1931) said that a fishing that the waters of Matagorda Bay and vessel at Port Isabel, Texas, had a the Gulf were full of them and great small load of B, patronus in August. "drifted upon quantities of menhaden Gowanloch (1932. 1933), writing 1872. the beach at Saluria" (Texas) in about the fishes of Louisiana, noted the the fish were alive or dead at Whether abundance 9f menhaden and said that it from the time they stranded is not clear "is not commercially fished along the the context. In the meantime Goode Gulf Coast." (1878) described Brevoortia patrorms based upon specimens from Brazos In 1932 Gunter began counting the Santiago, Texas, and the mouth of the fishes taken by the vessel "Black Rio Grande. Brazos Santiago remains Mallard" in Louisiana. Five otter- the southernmost record for B.patrOTlUS trawl hauls were made monthly from in the western Gulf. the head of Barataria Bay, to 5 miles offshore from Barataria Pass in the Jordan and Gilbert (1883) listed open Gulf. Counts were made from the menhaden from Pensacola and Gal- January 193 2 to June 1933; John C. veston. Evermann and Kendall (1894) Pearson continued thenn until Decem- reviewed previous Texas literature and ber 1933. The first results were pre- noted one young specimen from Gal ves sented in a paper on destruction of ton. Weymouth (1911) listed ten adults fishes by shrimp trawls (Gunter, 1936). and a "considerable number" of young, The tables were unwisely divided into the latter being doubtfully referred to commercial and noncommercial fishes Brevoortia tyrannus patronus Goode and table 7 is reworked from tables la, Most of the collection was made at lb, Ila and lib of that paper. Menhaden Calcasieu Pass, Louisiana, which lies formed 5.8 percent of the 80,093 fishes in the range of B. gunteri, and Wey- taken in the bay during both years and mouth's doubts may have been well 2.4 percent of the 46,518 fishes taken founded. Precise dates of collection in the shallow Gulf. Most of the bay were not given, and they were probably menhaden were small, and probably not known to Weymouth. The fishes were able to escape the trawl when were taken by Milo Spaulding during they moved out into the Gulf as larger 1906. fish. Table 7. — Numbers of fishes taken in trawls on the Louisiana coast, 1932-33 [As reported by Gunter (1936)] In Bay In Gulf Total Edible fishes: 1932 17,309 26,264 '^3,573 1933 2^,917 11,603 36,529 Nonedible fishes: Menhaden: 1932 2,166 1933 Other nonedible fishes: 1932 1933 Total nonedible fishes: 1932 1933 Gunter (1938a) gave a further re- menhaden were taken than all other port on the data collected aboard the fishes. The 2,664 B. patronus caught "Black Mallard", including collections made up 58.7 percent of the catch. made in 1934. Brevoortia patronus v/as Their size range was 89 to 198 mm. the third ranked fish, taken in the bays, standard length but only nine were and fourteenth in the Gulf. It made up smaller than the largest bay fish. 14.1 percent of the 144,000 fishes caught. Breuer (1957) said that Brevoortia sp. were not comnnon in the high- On the basis of collections, gen- salinity Baffin and Alazan Bays of eral observations, and evidence from Texas, but that a few large ones mass mortalities, Gunter (1941b) appeared in the fall and some small ranked the menhaden as second "in ones in the summer. Temperature and species mass" among fishes in shallow salinity ranges were not given for waters of the northern Gulf, preceded species reported in Breuer's paper. by Anchoa mitchilli and followed by Mugil cephaliiS' Gunter (1950) recorded 95 speci- Gunter (1945) showed that Bre- mens of B. gunteri taken along the VOOrtia ranked twelfth among all species Blackjack Peninsula shore (Aransas caught in Copano and Aransas Bays National Wildlife Refuge) of Mesquite and off the Gulf beach of Mustang and Ayres Bays, Texas from February Island, Texas; it ranked ninth in the to June 1945, where they were the bay catches. A comparison of trawl third most abundant fish in minnow catches in Louisiana bays and Texas seines. The average monthly tem- bays showed that Brevoortia ranked peratures ranged from 19.0°to32.1°C. third in Louisiana catches and seventh and salinity range was 9.5-18.3%o. In in Texas. It seemed to be less common the connecting salt-flat ponds, only a in Texas. few miles away, this species was taken only 8 times among 1,324 fishes, and Fowler (1945) listed B, patronus ranked ninth in numbers caught. The from Louisiana and Galveston. average temperatures in that area during the same period were 19.0°- Jurgens and Hubbs (1953) included 32.1° C. and the salinities were 4.4- B. gunteri as one of the fishes found in 12.2%o. the fresh waters of Texas. Knapp (1953) listed B, gunteri as ascending Caldwell (1954) listed B. gunteri and B. patronus as possibly ascending from a salt-marsh pond on Way Key, Texas rivers. (Cedar Keys) Florida. Suttkus (1958) said the fish was B. smithi. H. H. Hildebrand (1953) said men- haden are not often taken in trawls by Rounsefell (1954) said that several shrimpers, but sometimes large species, including "... the nnenhaden, schools are encountered which burst Brevoortia- ." were "nnore or less the trawls. He reported fifteen speci- independent of the waters between the mens of B. patronus taken in two trawl mainland and the barrier islands." hauls at 17 and 18 fathoms off south Texas. H. H. Hildebrand (1955) reported Reid (1955a), working in East Bay, eight specimens of B. gunteri from two Texas, noted that where "considerable" trawl hauls in 7-8 fathoms of water off water depth and mud bottom extended Punta Morros in the SE portion of the to the marsh edge, menhaden and Gulf of Campeche. They were taken various croakers were more abundant February 15, 1951. This is the south- while atherinids and cyprinodontids ernmost record for nnenhaden in the were less. While sampling the Gulf Gulf. Hildebrand said he had never beach at Gilchrist, Texas, across from heard of menhaden schools south of East Bay, Reid (1955b) found that more the Rio Grande. 20 . . . Gunter (1956b) recorded B, smithi, contained food nnaterial. Food of small B. giinteri and B. patronus as being menhaden (38-48 mm.) consisted chiefly euryhaline of phytoplankton with small amounts of zooplankton and plant fragments. Sev- Hubbs (1957) in a revision of a eral species of Anabaena comprised former checklist included Brevoovtia 77 percent of the total food while the gunteri among fishes to be found in remaining phytoplankton was com- fresh waters of Texas. posed of diatoms. Detritus constituted 11 percent of the stomach contents. Briggs (1958) listed B. smithi and The small menhaden had obviously B. tyrannus from the Atlantic coast of been straining plankton at the surface Florida and said B. tyrannus was to be and apparently at depths below the found on both sides of the Atlantic. He surface listed B, patronus from Tampa north- ward and B. gunteri from Cedar Keys The larger menhaden (85- 103 mnn.) (probably on Caldwell's record) to examined by Darnell (1958) contained, Campeche in the muscular pyloric region of the stomach, 99 percent ground organic Suttkus (1958) discussed the dis- matter and silt with a few diatoms, tribution of three species of menhaden foramniferans and copepods. Suspen- in the Gulf. In summary he said: sions of ground up organic matter were observed, particularly along the south "The two fine-scaled species, shore of the lake "where wave action Brevoortia gunteri and B. smithi, was reducing the organic material of occur in the western and eastern the marshy shore to the consistency of Gulf respectively. The single large- coffee grounds." Darnell concluded scaled form, B, patronus, overlaps that menhaden fed by filtration, and that B. gunteri in the western Gulf from suspended bacteria and material other Brazos Santiago, Texas, to Grand than living plankton were an important Isle, Louisiana, and overlaps B, component of the food of menhaden in smithi in the eastern Gulf at Cedar turbid estuaries. Keys." Springer and Woodburn (I960) Enemies and Mortality found young B. patrOHUS and young B, smithi together in the Tampa Bay, Arthur (1931) commented on men- Florida, area. Concerning the move- haden as food of the brown pelican in ment of young B. patronus they said: the following words: "In contrast to Suttkus' finding that "The pelican is a fish eater. While emigration from estuarine waters it may be true that the pelican, once took place in July and August, we in a while, may catch what man found no specimens in the bayou after terms a 'food fish*, the number of May. . . . The collection from Johns such fish of commercial value it Pass. . . during July does indicate consumes does not justify the prej- that migration to the Gulf had occur- udice felt against the big bird. When red at least this early and possibly the World War was on and the Food as early as June." Administration was calling on the people to 'eat more fish', some fishermen, residents of Florida, Food and Feeding asked that an edict be issued calling for the pelican's extermination on Gowanloch (1933). Gunter (1945) the theory that it was to blame for and Reid (1955a) stated that GuM Bre- a food-fish shortage. But before this VOOrtia are plankton feeders. Darnell wholesale slaughter was ordered, (1958) examined the stomach contents cooler heads counseled inquiry and of 17 specimens of B. patrOTtUS from in the summer of 1918 the Depart- Lake Pontchartrain, Louisiana. All ment of Conservation conducted a 21 . survey of the Brown Pelican breeding pounder, ElopS saurus, and the lizard- places, and secured exact data as to fish, Synodus foetens. It was concluded what this particular pelican ate, when that predation on young fish, especially fish of all kinds were most plentiful menhaden, is severe. Reid (1955b) also in the Gulf waters. pointed out that "birds of the area also exact a toll on these fish." "The Louisiana coast from Pearl River to the Sabine was explored and pelicans and their food collected Parasites and Diseases and sent to Washington, D. C. for identification by the United States Pearse ( 1 952a) described the para- Bureau of Fisheries. The result of sitic copepod, Caligiis ventrosetus , from the examination of many hundreds of the gills of B. gunteri at Port Aransas, stomachs showed that 97 per cent of Texas. He also listed Lemanthropus the fish were menhaden or 'gulf brevoortiae from B. patronus at the sardines', and 3 per cent were silver- same locality. Pearse (1952b) recorded sides - not a single food fish used by Lemanthropus brevoortiae from the man was found!" gills of B. patronus in Florida. Causey (1953) reported Lernaeenicus radiatus Lund (1936) reported that about 85 from B. 'tyrannus" at Grand Isle, percent of the dead fish reported on the Louisiana. south Texas coast in the sumnner of 1935 were menhaden. Facts reported Causey (1955) reported the para- by Lund and others verbally to the sitic copepods, Lemanthropus brevo- senior author indicate, with little doubt, ortiae and L. radiatus from B. patronus that this mortality was the result of a at Pascagoula, Mississippi. Hargis "red tide". (1955) listed the monogenetic trema- tode, Mazocraeoides georgei, from the Gunter {1941a) noted that, although gills of B. patronus at Alligator Harbor, menhaden were listed as a common Florida, as a new host record. Koratha bay fish in previous papers, only three (I955) described a new monogenetic were observed killed by the hard cold trematode, Diclidophora lintoni, from wave of January 1940 on the Texas the gills of B. gunteri taken at Port coast. These were seen in Copano Bay Aransas, Texas. and were listed as B. patTOnuS- The specimens could have been B. gunteTi. Hargis (1957) listed the following monogenetic trematodes (Family Ma- Gunter (1945) found no menhaden zocraeidae) as parasitic on Brevoortia in the stomachs of 23 7 redfish patronus: Clupeocotyle brevoortia, C. (SciaenopS ocellata) examined. He found megaconfibula, Kuhnia brevoortia and three Brevoortia in the stomachs of Mazocraeoides georgei 153 Cynoscion nehulosus. Sparks (1958) listed the digenetic Gunter, Davis, Williams and Smith tr emai.tod.&. Parahemiurus merus.irom (1948) recorded ten B. /JOfronMS killed the gills of B. patronus a.s a new host by the red tide along 20 yards of Fort record. Myers beach, among 208 other fishes. The species could have been B. Smithi. Menhaden Fishery Investigations Reid (1955a) found that 89 percent of "white trout" (Cynoscionaretmrius) Miles and Simmons (1950) gave a stomachs contained fish "and menhaden review of the menhaden industry and constituted the greatest part of this fishery. They also summarized certain item." He indicated that the menhaden biological information collected by a soon grow too large to be taken by series of workers at the Marine Lab- "trout". He said menhaden were also oratory of the Texas Game and Fish preyed upon to a lesser extent by the Commission and published a series of croaker, two nnarine catfishes, the ten- mimeographed reports by Breuer, 22 . , . Anderson, Knapp, Wilson, and Miles Knapp (1950) presented some of and Simmons. Miles and Simmons, give the same data and came to similar the following information: conclusions. 1. commercial Other sports and Filipich (I947) made five unan- in very small quan- fishes were taken nounced checks of menhaden boat un- the result of tities with menhaden. As loadings at Mississippi factories. In the figure given was one one study 295 tons of menhaden unloaded, one menhaden or 0.024 animal to 4,490 mackerel and 6 white trout were found. percent, including shrimp and crabs. The fishes caught with 5,326,000 men- haden were 36 gafftopsails (Bagre Gowanloch (1949) reviewed the marina), 75 crabs, 91 jacks (Caranx menhaden-sports fisherman contro- hippos), 103 croakers {Micropogoti un- versy and concluded that no harm was dulatus), 191 shrimp, 205 Spanish done to sports fishes by the menhaden mackerel (Scontberomorus maculatus) industry. 242 sand trout (Cynoscion), 304 blue- fish (Pomatomus saltatrix), and a few Christmas, Gunter and Whatley others, all less than 8 each. The total (i960) reported a study of fishes other was 7,589 other fishes. than menhaden taken in menhaden purse seines in waters around the mouth of Another study showed that 2,183 the Mississippi River and in Missis- other animals, including shrimp, crabs sippi Sound during the 1958 and 1959 and squid were taken with 2,500,000 seasons. Samples were taken from menhaden. catches which totaled nearly 2 million pounds. In numbers of fish, more than 2. Mackerel were found with 97 percent of the sampled catch were smaller menhaden and bluefish were menhaden. The other fishes observed found with larger menhaden in offshore included 62 species. Brevoortia pa- waters. Bluefish appeared when the tyonuS was the only menhaden found in snnaller menhaden were replaced by the catch. The fishery was generally larger ones in July. Crabs, shrimp, prosecuted in shallow, low-salinity sand trout and croakers were usually waters. Seventy percent of the sampled taken in shallow waters. Bluefish, menhaden catches were made in waters mackerel, tarpon, jacks and sharks of salinities between 5 and 24 parts per seenned to be the main predators on thousand. Surface water temperatures menhaden. at the sampled seine locations varied from 22.6° to 30.5° C. Lower tem- peratures were encountered at the 3. Forty species of fish, com- beginning and the end of the fishing prising 26,005 individuals were ex- season. Fishing started in May when amined for stomach contents. Of these, water temperatures rose to About 23° C 13,288 were sea trout, Cynoscionnebu- and stopped in the fall when surface losus 3,428 were Spanish mackerel and waters reached approximately the same 3,137 were redfish, Sciaenops ocellata. temperature again. The remainder were 38 other species of fishes. They contained 581 menhaden. The percentages eaten by the common fishes were given. Menhaden were not SUMMARY preferred food of the common sports Three of the five North American and comnnercial fishes. species of menhaden have been found in the Gulf of Mexico. The present 4. Knapp showed that the stomachs distribution, based on the published of menhaden, mullets and "shad", record, is: remaining intact in the stomachs of predators after other parts had been 1. Brevoortia gunteri, Grdiridlsle, digested, could be identified by their Louisiana westward and southward to s hape the Gulf of Campeche. 23 , 2. Brevoortiapatronns , the Tampa gunteri have been reported from the Bay area, Florida to Brazos Santiago, Laguna Madre, Texas, in February. Texas. While there may be confusion be- 3. BreVOOrtia smithi, Cedar Keys, tween both the larvae and the young of Florida southward and presumably two species where their range over- around the peninsula to the Atlantic. laps, reports oi B. patronus 16-25 mm. (R. D. Suttkus, personal communica- long extend from November to June, tion.) with minimum sizes at both ends of the period. There is some evidence of a Menhaden fishing in the Gulf of double influx of larvae . Thus the spawn- Mexico is dependent on B, patvonus ing season is long, over the fall and and is mostly prosecuted in Louisiana winter and into the spring. B. gunteri waters. Gulf landings of menhaden seem to spawn in the spring in Texas increased rapidly with the expansion of waters. the fishing until 1957, when landings were only 65 percent of the 1956 catch. However, 1958 landings increased to Menhaden eggs and early larvae about 80 percent of the 1956 record. have not been reported in the Gulf. Apparently the 1957 decrease was Presumably, the eggs are bouyant like caused by factors other than depletion those in the Atlantic and the larvae of the population. remain near the surface, as has been reported in Virginia. All information agrees with the supposition that Atlantic menhaden The slender larvae which enter spawn at sea, or at least in high salini- estuarine waters undergo metamorpho- ties, and the larvae work in to shore sis between 20 and 30 mm. standard to spend their early life, very often in length in Lake Pontchartrain. Data on low salinity or even fresh water. Ap- growth rates are scarce. A group parently spawning is also governed by thought to be Brevoortia gunteri ranged temperature. Comparison of surface from 88-133 mm. long in April and 128- temperatures (Bumpus, 1957) for re- 1 73 mm . long in August in Texas waters . ported months of spawning along the A group of Texas Brevoortia 88-103 Atlantic coast indicates that spawning mm. long in August had a mode of 128- takes place in the Atlantic at about the 133 mm. the following May. same temperature in all localities. Thus, the spawning season is during On the Atlantic coast, menhaden the summer on the New England coast undertake extensive migrations up and and in the winter off the South Atlantic down the coast with changes in season, States. during which they are followed by the fishermen, at least on the trip south. There are no records of spawning Nothing similar is known in the Gulf, B, patvonUS and presumably they spawn probably because it is an east-west at sea. Except for one report of ten- coast. tatively identified Brevoortia larvae on the Gulf beach in March and April in Large menhaden are not comnnon south Texas, there are no reports of in the bays and evidently the young menhaden larvae on the beach. On the fish move out, as they grow up, mostly other hand, vast numbers of larvae in late summer and fall. Salinity is a and juvenile menhaden have been re- critical factor in the development of ported in the bay waters of Louisiana larval Atlantic menhaden, with low and Texas. Low -salinity waters un- salinity waters being essential to nor- doubtedly serve as nursery grounds. mal development. In Texas, larvae and juveniles 21-48 mm. long have been Brevoortia gunteri in spawning found at salinities of 2.0-20.4%o and condition have been reported inCopano temperatures of 1 5.5°-3 1 .0° C . In White and upper Aransas Bays, Texas, Lake, Louisiana, menhaden larvae have in February and March. Spawning B. been found in virtually fresh water. 24 The numbers of larvae found in ANDERSON, ANDREW W., and C. E. the bays may vary enornnously fronn PETERSON. year to year. There may be sonne con- 1954. Fishery statistics of the nection between the heavy 1956 com- United States, 1951. U. S. Fish mercial production of menhaden and and Wildlife Service, Statistical Reid's report of "staggering" num- Digest 30, 341 p. bers of larvae in East Bay, Texas, in the summer of the same year. ANDERSON, A. W., and E. A. POWER. 1949. Fishery statistics of the There is only one report on the food United States. 1945. U. S. Fish of Gulf menhaden. It was found that and Wildlife Service, Statistical Lake Pontchartrain menhaden fed by Digest 18, 372 p. filtration and consumed considerable 1951. Fishery statistics of the quantities of detritus and suspended United States, 1948. U. S. Fish bacteria in addition to living plankton. and Wildlife Service, Statistical Digest 22, 304 p. Menhaden in the Gulf of Mexico 1954. Fishery statistics of the are not known to be chased ashore by United States, 1951. U. S. Fish predators or to die in recurring sum- and Wildlife Service, Statistical mer maladies from unknown causes Digest 30, 341 p. as they do in the Atlantic. They have 1955. Fishery statistics of the been killed in considerable numbers United States, 1952. U. S. Fish by the red tide in Texas and Florida and Wildlife Service, Statistical but these instances are outside the Digest 34, 345 p. commercial range. Menhaden seenn to 1956a. Fishery statistics of the be resistant to sudden cold waves which United States, 1953. U. S. Fish decimate other fishes in Texas and and Wildlife Service, Statistical Florida, and none were reported killed Digest 36, 338 p. in three studies of cold-killed fishes 1956b. Fishery statistics of the in Florida and Texas. A few menhaden United States, 1954. U. S. Fish have been reported killed by the cold and Wildlife Service, Statistical on the Texas coast. Digest 39, 446 p. 1957. Fishery statistics of the Menhaden fishermen on the Gulf United States, 1955. U. S. Fish catch few other fish, comparable to and Wildlife Service, Statistical the situation on the Atlantic coast. Digest 41, 446 p. The chief menhaden enemies seem to be bluefish, mackerel, sharks, and ARTHUR, S. C. tarpon. The common shore and shallow- 1931. The birds of Louisiana. Loui- water sports and commercial fishes siana Conservation Department, seem not to subsist to any great extent Bulletin No. 20, 598 p. on larger menhaden, but predation upon small menhaden in the bays seems BAKER, EMMA (Comp.). to be very heavy. 1955. Full text of the game fish and fur laws of Texas. Texas Game and Fish Commission, Austin. LITERATURE CITED 319 p. ANDERSON, ANDREW W., and C. E. BALDAUF, R. J. PETERSON. 1954. Survey and study of surface 1952. Fishery statistics of the and subsurface conditions in and United States, 1949. U. 5. Fish around Beaumont, Texas. Bio- and Wildlife Service, Statistical logical survey of the Neches Digest 25, 298 p. River in the region of Beau- 1953. Fishery statistics of the mont, Texas. Texas A. and M. United States, 1950. U. S. Fish Research Foundation. [Unre- and Wildlife Service, Statistical leased mimeographed report.] Digest 27, 492 p. 184 p. 25 BEAN, T. H. CHRISTMAS, J. Y., GORDON GUNTER, 1903. Catalogue of the fishes of New and EDWARD C. WHATLEY. York. New York State Museum, I960. Fishes taken in the menhaden Bulletin No. 60, Zoology 9, 784 p. fishery of Alabama, Mississippi, and eastern Louisiana. U. S. BIGELOW, HENRY B., and WILLIAM C. Fish and Wildlife Service, Spe- SCHROEDER. cial Scientific Report- -Fish- 1953. Fishes of the Gulf of Maine. eries No. 339, 10 p. U. S. Fish and Wildlife Service, Fishery Bulletin No. 74, vol. 53, DARNELL, REZNEAT M. 577 p. 1958. 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BUMPUS, DEAN F. EVERMANN, B. W., and W. C. KEN- 1957. Surface water temperatures DALL. along Atlantic and Gulf coasts of 1894. The fishes of Texas and the the United States. U. S. Fish and Rio Grande basin, considered Wildlife Service, Special Scien- chiefly with reference to their tific Report--Fisheries No. 214, geographic distribution. Bulletin 153 p. of the U. S. Fish Commission, CALDWELL, D. K. vol. 12 (1892), p. 57-126. 1954. Addition to the known fish fauna in the vicinity of Cedar FIEDLER, R. H. Key, Florida. Quarterly Journal 1943. Fishery statistics of the of the Florida Academy of United States, 1940. U. S. Fish Sciences, vol. 17, no. 3, p. and Wildlife Service, Statistical 181-184. Digest 4, 225 p. CAUSEY, D. 1953. Parasitic Copepoda from FILIPICH, MECO. Grand Isle, Louisiana. Occa- 1947. Report to the Mississippi Sea- sional Papers of the Marine food Connnnission, Biloxi, Mis- Laboratory, Louisiana State Uni- sissippi. Menhaden Plants Re- versity, No. 7, 18 p. port, 4 p. 1955. Parasitic Copepoda from Gulf of Mexico fish. Occasional FOWLER, HENRY W. Papers of the Marine Labora- 1931. A collection of fishes from the tory, Louisiana State University, Texas coast. Copeia, 1931, no. 2 No. 9, 19 p. (July), p. 46-50. 26 FOWLER, HENRY W. GOWANLOCH, J. N. 1936. The marine fishes of West 1933. Fishes and fishing in Loui- Africa. Bulletin of the American siana. Louisiana Conservation Museum of Natural History, vol. Department, Bulletin No. 23, 70, part 1, 605 p. 638 p. 1945. A study of the fishes of the 1949. Menhaden facts and fallacies. southern Piedmont and coastal Advertiser Publishing Company, plain. Monographs, Academy of Pascagoula, Miss. 26 p. Natural Sciences of Philadelphia, No. 7, 408 p. GUNTER, GORDON. 1936. Studies of the destruction of GILL, THEODORE N. marine fish by shrimp trawlers 1861. Synopsis of the subfamily in Louisiana. 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Proceedings 1956a. Some relations of faunal dis- of the United States National tributions to salinity in estuarine Museum, vol. 2, p. 121-156. waters. Ecology, vol. 37, no. 3, p. 616-619. GOWANLOCH, J. N. 1956b. A revised list of euryhaline 1932. Sea fishes and fishing in Loui- fishes of North and Middle siana. Louisiana Conservation America. American Midland Department, Bulletin No. 21, Naturalist, vol. 56, no. 2, p. 187 p. 345-354. 27 GUNTER, GORDON. HILDEBRAND, SAMUEL F. 1957. Temperature. In Treatise on 1941. An annotated list of salt and marine ecology and pale- brackish water fishes, with a oecology, vol. 1, Chapter 8, p. new name for a menhaden, found 159-184. The Geological Society in North Carolina since the pub- of America, Memoir 67, New lication of "The fishes of North York. Carolina" by Hugh M. Smith in 1907. Copeia, 1941, no. 4 GUNTER, GORDON, C. C. DAVIS, R.H. (No- vember), 220-232. WILLIAMS, and F. G. WALTON SMITH. p. 1948. review of 1948. 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