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 States, nnenhaden have been utilized for food and for fertilizer since the days of the early colonists (Goode, 1879)- Processing the fish for oil, which in the beginning consisted nnerely of press- ing the decomposing raw fish, was started in Rhode Island in 1811 (Goode, 1884). In the 1870's the industry was brought to North Carolina by northern soldiers who had noted the abundance of menhaden in southern waters during the Civil War (Ellison, 1951). After some initial failures the menhaden industry became well established in the South Atlantic States in the 1880's. Despite the long history of the menhaden fishery in the Atlantic and its rapid expansion in the Gulf of Mexico, extensive studies of the biology of these important commercial fishes have not been undertaken until recently. The only published bibliogra- phy on biology of the American Men- haden (Reintjes, Christmas, and Collins, I960) includes most of the literature with very brief annotations and a subject index. This review of the literature on the biology of menhaden brings together information from somewhat scattered sources. Gulf commercial production is discussed first, followed by a summary of the taxonomy, distribution and relation- Note.--Dr. Gordon Gunter, Director, and J. Y, Christmas. Fishery Biologist, Gulf Coast Research Lab- oratory, Ocean Springs, Mississippi. ships of the species of the genus Brevoortia. The Atlantic menhaden, Brevoortia tyrannus (Latrobe), is dis- cussed under topic headings. The chief papers are cited, but no attenipt was made to cover completely the volu- minous and sometimes repetitious literature. In the section on the Gulf menhaden almost every piece of lit- erature pertaining to the biology of this species has been cited. Topical arrangement of the material parallels that of the Atlantic menhaden. GULF MENHADEN FISHERY STATISTICS The development of a menhaden fishery in the Gulf States moved from east to west. There was considerable production of menhaden on the Florida east coast as early as 1923 but con- tinuous production did not start in the Gulf States until 1937 (Power, 1958). The production statistics shown in table 1 were taken from various U. S. Fish and Wildlife Service reports. There are some State statistical re- ports, but these are not connparable from State to State nor fronn year to year. They would be of little value except for the years 1946-47 in the developmental period of the Gulf men- haden fishery. Catch records given by Anderson and Peterson (1953) show that men- haden were caught off the west Florida coast as early as 1880. From 1902 to 1938 annual production in this area fluctuated between Z, 000 and 18,815,000 pounds and the annual average was 7,059.000 pounds for the 13 years re- ported. Alabama reported 10,000 pounds in 190Z and 4,000 pounds in 1931. Texas reported 14,118,000 in 1918 and 8,517,000 pounds in 1923. No other menhaden catches from the Gulf of Mexico were reported during the years 1880 to 1938. Therefore, it may be said that the Gulf menhaden fishery got started in 1939 with the location of plants in Mississippi. During the World War II years Gulf production came from Florida and Mississippi. By 1948 plants had been located in Louisiana and Texas. Landings did not exceed 200 million pounds until 1949- Table 2 shows the total catch, yearly average and percentage catch Table 1. --Landings of Gulf of Mexico menhaden by States, 1939-58 [In thousands of poionds] Year West coast Florida Alabama Mississippi Louisiana Texas Total Gulf Source 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 0 9,000 0 0 11,849 Anderson & Peterson (1953) 0 25,195 (No data 0 collected) 0 25,462 Fiedler (1943) 0 57,340 (No data 0 collected) 0 64,506 Anderson & Power (1949) 0 68,636 88,110 28,185 184,931 Anderson & Power (1951) 0 44,579 165,913 41,136 276,507 Anderson & Peterson (1952) 0 69,550 207,755 47,191 326,030 Anderson & Peterson (1953) 0 114,895 209,574 30,121 357,965 Anderson & Peterson (1954) 0 112,890 233,373 52,984 459,984 Anderson & Power (1955) 0 ■ 58,933 307,492 66,589 437,045 Anderson & Power (1956a) 0 79,445 270,094 51,702 401,243 Anderson & Power (1956b) 0 128 , 123 298,309 52,625 480,992 Anderson & Power (1957) 0 172,592 320,521 66,691 559,836 Power (1958) 0 142,124 162,817 57,585 362,533 Power (1959) 0 123,346 241,813 68,559 442,826 C.F.S. No. 2165 Table 2. --Menhaden landings of the Gulf States, 1949-58 [in thousands of pounds] Location Totals Yearly average Percent West Florida Alabana Mississippi Louisiana 55,64-0 5,564 lA 0 0 0 1,046,477 104,648 25.5 2,4-67,661 246,766 60.1 Texas 535,183 53,518 13.0 Total 4,104,961 440,496 100,0 by Gulf States for the years 1949-58, inclusive. The Florida catch was neg- ligible, being slightly more than 1 percent of the total. No menhaden landings were reported from Alabama. The major part of the catch, over 60 percent, was landed in Louisiana. How- ever, table Z does not fully show the preponderance of Louisiana waters in menhaden production. Statistics are not available on catch localities, but Texas boats have little Texas water in which to fish (Baker, 1955) and the Mississippi coast line is less than 70 miles long. In 1950 the Gulf catch was 32.6 percent of the United States total (table 3). In 1955, although the Gulf catch had increased to over 480 million pounds, the Gulf percentage of the United States total catch declined to 25.8. Gulf fishermen took 26.7 percent of the record catch of 1956, when over 2 billion pounds were reported for the United States total landings. Total landings dropped sharply in 1957. Power (1959) listed several reasons for the 1957 decrease. Although the number of purse seines (table 4) con- tinued to increase in the Gulf, the 1957 Gulf catch was down to 21.5 percent of total landings. Little information on catch per unit effort is available in the Gulf menhaden fishery and the period of time during which the fishery has operated at a high level is only 10 years. While the number of yards of seine (table 4) for the Gulf coast men- haden fishery more than doubled from 1948 to 1958, the catch per yard in- creased from 13.9 thousand pounds to 19.8 thousand pounds in 1956. The catch per yard of seine dropped to 1 1 .6 thousand pounds in 1957 and canne back to 13.9 thousand pounds in 1958. MENHADEN BIOLOGY TAXONOMY, DISTRIBUTION AND RELATIONSHIPS Menhaden belong to a genus of clupeid fishes inhabiting the shore regions of the Atlantic from Nova Scotia to Argentina and, according to de Buen (1958), Pacific waters off Chile and Peru. Unverified records place the genus off the west coast of Africa. The genus Brevoortia was es- tablished by Gill (1861). Brevoortia tyrannus (Latrobe) was designated the genotype. The genus was revised by Goode (1878) and by Hildebrand (1948). According to Hildebrand (1948) there were seven species, five in North America and two in South America. Hildebrand had no North American records south of Indian River, the coastal lagoon of east Florida, and none south of Tannpa on the west Flor- ida coast. The mouth of the Rio Grande River was the southern limit in the western Gulf. Hildebrand was quite firm in asserting, as a result of his sojourn in Dry Tortugas, thit there 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 1951 7-45,950 1952 923,606 1953 1,259,031 1954 1,336, 524 1955 1,386,791 1956 1,537,403 1957 1,327,595 326,030 357,965 459,984 437,045 401,243 480,992 559,836 362,533 1,000,498 1,103,915 1,383,590 1,696,076 1,737,767 1,867,783 2,097,239 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 Year Number of purse seines Number of yards of seine Catch per yard in thousands of pounds Source 194-8. 1949. 1950. 1951. 1952. 1953. 1954.. 1955. 1956. 1957. 1958. 34 13,350 13.9 Anderson & Power (1951) 53 20,125 13.7 Anderson & Peterson (1952) 66 24,600 13.3 Anderson & Peterson (1953) 64 24,300 14.8 Anderson & Power (1954) 71 27,000 17.0 Anderson & Power (1955) 74 28,575 15.3 Anderson & Power (1956a) 63 23,502 17.1 Anderson & Power (1956b) 67 26,019 18.5 Anderson & Power (1957) 75 28,225 19.8 Power (1958) 79 31,171 11.6 Power (1959) 80 31,949 13.9 C.F.S. No. 2165 No members of this genus had been reported from South America north of Salvadore (Baia), Brazil or from the West Indies. One exception to the generally ac- cepted western Atlantic distribution of Brevoortia is the statement of Goode (1879) concerning a menhaden in the eastern Atlantic. He said large schools of B. dorsalis of West Africa had been reported to him. Fowler (1936) syn- onym ized the African menhaden with B. tyrannus . Since no specimen was available for check, Hildebrand (1948) doubted the identification. Briggs (1958) listed B. tyrannus from Africa, but gave no records. De Buen (1958) reduced the genus Ethmedium (Thompson, 1916) to sub- generic status in the genus BrevoOTtiOt thereby extending the range of Brevoortia to the Pacific coasts of Peru and Chile. De Buen's arrange- ment places the seven species of Brevoortia recognized by Hildebrand (1948) in a s\xhgen\is, Brevoortia , and adds Brevoortia (Ethmidium) maculata (Valenciennes) 1847 and Brevoortia (Ethmidium) chilcae (Hildebrand) 1946 to the genus Brevoortia. Brevoortia tyrannus of the Atlantic is the most important and abundant menhaden known (Ellison, 1951), and approximately 73 percent (table 3) of all the menhaden landed in the United States from 1950 to 1957 belonged to this species. The Atlantic menhaden has been reported from the Bay of Fundy to Indian River, Florida, and has been fished from Maine toMayport, Florida. Goode (1878) de scribed Brevoortia patronus from specimens collected in the Gulf of Mexico at Brazos Santiago, Texas. Jordan and Evermann (1896) listed the Gulf menhaden a.s BrevOOrtia tyrannus patronus Goode. Regan (1917) placed Brevoortia patronus in synon- ymy with B. tyrannus. Gunter (1945) stated that ' ' Brevoortia patronus Goode, as some authors would have it", had been recog[nized as the only species of menhaden in the Gulf. Since only spe- cific names \were used in his paper, Gunter (1945) referred to E. t. pairOHUS as B. tyranniis- Hildebrand (1948) also noted that B. patroilUS had generally been considered by authors as only subspecifically distinct from B, tyvan- HUS or identical ^with it. However, Hildebrand's diagnosis showed that B. patronus had fewer oblique scales, fewer modified scales in front of the dorsal, fewer vertebrae, fewer ventral scutes and fewer pectoral and dorsal, and more anal rays. The lower caudal, pectoral, maxillary and lower mandible were generally longer. The head was longer and deeper and body depth was greater. B. patronus was accorded full specific rank in Hildebrand's 1948 revision of BrevoOTtia. Meristic char- acters indicated that B. patronilS re- sembled B, tyrannUS of easternFlorida more than it did B. tyranrais farther north. Hildebrand (1948) noted that the lack of a decrease in number of ver- tebrae in southern menhadsn was con- trary to the general observation of lower vertebrae counts for other fishes in warmer waters. Hildebrand (1941) described a small-scaled, nonslimy, yellow-finned menhaden fronn the coast of Carolina and nanned it Sniithi . Ho had pre viously (Hildebrand, 1920) called this fish Brevoortia aurea (Agassiz), which is a South American species. The range was given as Beaufort, North Carolina, to Indian River, Florida. Hildebrand (1941) noted that B. smithi was not abundant, did not school in large num- bers, and was called "yellowfin shad" by local fishermen. He said the fisher- men kept fish of this species aside and carried them home for food. Suttkus (1958) reported B. smithi from the eastern Gulf of Mexico between Cedar Keys and Placida, Florida, noting ". . . that Hildebrand was aware of the pres- ence of Brevoortia smithi in the Gulf of Mexico long before I made my inde- pendent discovery." Gunter (1945) noted a second spe- cies of menhaden in the Gulf which he said was close to if not identical with Brevoortia smithi of the Atlantic Hildebrand (1948) described this as a new species, B. gunteri, which he con- sidered to be a Gulf cognate of B, smilhi. B. smithi and B. gunlcri agreed in having small scales, pointed yellow- ish fins, more silvery color and less green along the back, and in being notably less slimy than B, lyVdlllluS and B. patronus. B. glinteri differed from B. smithi in having a larger head, greater depth and longer maxillaries, mandibles and pectorals. The ventral scutes and vertebrae were fewer in number. Hildebrand (1948) examined specimens from Grand Isle, Louisiana; Aransas Bay, Texas; and the mouth of the Rio Grande, Texas. The fifth North American species of Brevoo^'tia is brevicaudata. Goode (1878) described the species from the only known specimens, collected at Noank, Connecticut, in 1 874. Hildebrand (1948), after re -examining the se speci- mens, listed nine ways, including the short caudal fin, in which they differed from B, tyranmiS. Scalation was simi- lar to that of B. tyrannus. Bigelow and Schroeder (1953) questioned the cor- rectness of these views. The ranges of Brevoortia aurea (Agassiz) and Brevoortia pectinata Jenyns in South America are not well known. Hildebrand (1948) pointed out that the seven American menhaden could be divided into a large -scaled group, B. tyratmiis, B. brevicaudata, B.patromis, B. pectiiiata, andB. aurea , and a small - scaled group, B. smithi and B, gunteri. He further stated that these menhaden could also be divided into two groups according to the shapes of the ventral fins. Brevoortia tyramms, B. brevi- caudata, and B. patronus had rounded fins, while the other four had pointed fins. The two South American species differed from the others by less reduc- tion in size of the scales on the back and on the base of the caudal as com- pared to mid-scales on the sides. Hildebrand (1948) noted that with the exception of B, brevicaudata, North American species of Brevo&rtia could be divided into two closely related pairs on either side of the Florida peninsula. Species of sciaenids and clupeids, which are cognate on the east and west sides of Florida, were listed to support Hildebrand's theory that the paired species of menhaden differ- entiated after the peninsula last rose above the sea. ATLANTIC MENHADEN Spawning Peck (1894) said minute organisms furnish food for menhaden "not only within the limits of these brackish- water inlets and estuaries where the spawn is left to develop" but also in more open waters, apparently having come to this conclusion concerning spawning because small fish were abundant in estuarine situations around Woods Hole, Massachusetts, in the sunnmer of 1893. Peck also pointed out that low salinity areas were im- portant "because they were intrusted with so much embryonic and larval life of the migratory inhabitants of the coast." Smith (1907) said that in New England the spawning of menhaden took place in late spring and early summer, and from Chesapeake Bay southward spawning occurred in late fall and early winter. Fishermen reported large fish with full roe in Novennber and Decem- ber in North Carolina "rivers", but disagreed as to whether the spawning act took place only in the ocean (Smith, 1907). Schools of young were found in- shore in winter. However, Smith was not sure of the spawning location and said "there is some evidence" that spawning took place both in the sea and inside waters. Kuntz and Radcliffe (1918) found some fish ready to spawn and some spent in Woods Hole harbor in August. Eggs and larvae were occasionally taken there during the summer and were abundant off Gay Head in August. Larvae 20 mm. long were taken in the harbor on October 21, 1914. These authors indicated that the main spawn- ing period in the Woods Hole area was June, July, August, "and later". Bigelow and Welsh (1925) found spent and ripe fish in the Gulf of Maine in July and August. They said spawning around Woods Hole took place from June to October. The Grampus (Bigelow and Welsh, 1925) took eggs and larvae in Nantucket Sound and west of Martha's Vineyard in October 1915. Bigelow and Schroeder (1953) reported no eggs and larvae north of Cape Cod but said fry were taken in abundance in Casco Bay in 1900. Warfel and Merriman (1944) found menhaden larvae (less than 23 mm., standard length) in the harbor of New Haven, Connecticut, in July and Sep- tember. The standard length of men- haden collected in October was 30mnn. and a few 38-55 mm. in standard length remained until November. Perlmutter (1939) reported eggs and larvae at 27 of 52 stations in Long Island Sound. He said the spawning season extended from May to October. Westman and Nigrelli (1955) said men- haden in the zero year class appeared in estuaries, tidal rivers, creeks, mooring basins and canals in June 1953 and remained until late September or October. Eggs and larvae were not found in Great South Bay by Perlmutter (1939) although zeros (young-of-the- year) more than an inch in length were usually abundant there inshore. According to Westman and Nigrelli (1955) "The abundance of these young menhaden is difficult to imagine and, unlike the adult population they exhibit no aversion to fresh water." Due to the long spawning season sonne of the young- of-the -year were 7 inches long in October and others were only 2 inches long. Evidently Westman and Nigrelli did not collect larval stages. Radcliffe (Kuntz and Radcliffe, 191 8) took larvae 30 mm. long at the nnouth of the Potomac in February 1914. According to Hildebrand and Schroeder (1928), menhaden in the Chesapeake Bay region seemed to spawn in the fall. Larvae averaged 27.7 mm. long in January, 33.5 mm. in February, 27.3 mnn. in March, 33.0 mm. in April, and 46.0 mm. in May. Pearson (1950) took menhaden larvae at Old Point Comfort, Virginia, during April and May. Massman, Ladd and McCutcheon (1954) caught young menhaden in the brackish and fresh waters of four Virginia Rivers, up to 27 miles above brackish waters, in April and May. These fish were collected in plankton tows and ranged from 24 to 30 mm. standard length. Ninty-eight percent of the 8,000 specimens were found at salinities of 2-3 %o. The ratio of num- bers caught in surface and bottom tows was 200 to 1. In June, menhaden of 40.6 mm. in mean standard length were the most abundant species in the shore zone just above brackish water. Young fish were present in July in the same area. Surface trawls showed essen- tially the same pattern in September when fish averaged 94 mm. standard length. Ellison (1951) quoted Hardcastle (MS) who concluded that, in North Carolina, menhaden probably spawn in late winter near the Gulf stream. Ellison (1951) said the young fish have wide salinity tolerance and live in slightly brackish waters or waters with the salinity of sea water, and quoted Westman and Bidwell (1948) who re- ported young 45 miles up the Hudson River and in Long Island Sound at the same time. Bigelow and Schroeder (1953) said that W. W, Welsh concluded that sexual maturity in menhaden is attained by the third year of life. Westnnan and Nigrelli (1955) wrote that all menhaden 3 years of age or more appeared to be adult fish. Goode (1884) reported that 150,000 eggs were found in one menhaden. Ellison (1951) quoted Hardcastle (MS) who found that 41 percent of mature and immature males were in- fested with a gonadal parasite, Eimevia brevoortia, a sporozoan. The effect of this infestation on sperm production was not determined. Perlmutter (1939) is the only au- thor who has reported salinities at which menhaden spawn. He gave the figure as 84-100 percent (30.5 - 35%o) sea water, at 55°-80°F. All other re- ports of menhaden eggs correspond with high salinity situations. Growth Kuntz and Radcliffe (1918) re- ported that menhaden eggs were 1.4- 1.6 mm. in diameter. Bigelow and Schroeder (1953) gave the measure- ments as 1.5-1.8 mm. and said men- haden produced the largest fish eggs to be found in the Gulf of Maine. McHugh, Oglesby and Pacheco (1959), after studying the Chesapeake Bay pound-net and purse-seine fish- eries, assumed that, in the Chesapeake Bay area, menhaden left the Bay as they approached maturity in late Sep- tember, October and November. Spawn- ing apparently occurred outside the Bay in the fall and continued through the winter. Bimodal distribution of length and weight frequencies in age- group 0 suggested that two spawning peaks may occur in the Chesapeake region. Welsh (Bigelow and Welsh, 1925) found experimentally that hatching took place in less than 48 hours and that the larvae are 4.5 mm. long at that time. Four days after hatching they were 5.7 mm. long. There are no other data on the time elennent and growth of the larvae. The dorsal and caudal fins were visible in the 9 mm. stage and all fins were differentiated at the 23 mm. stage. At this stage the larva was long and slim; when the young menhaden had reached 33 mm. in length it had taken on a more fish-like appearance and acquired scales. June and Chamberlin (1959) dem- onstrated that low salinities are essen- tial to the normal development of Atlantic menhaden larvae. Larvae and metamorphosing individuals from low- salinity areas invariably perished when transplanted to high-salinity rearing ponds, although individuals trans- planted a few weeks after metamorpho- sis survived. Similar phenomena were observed in natural conditions on sev- eral occasions. Larvae reared through metamorphosis in high salinities de- veloped abnormalities while those de- veloping at low salinities were normal. Hildebrand (1948) has noted that all species of young menhaden (aureo and smithi not verified) up to 70 mm. long have minute teeth on the margin of the maxillary which are subsequently lost. Welsh (Bigelow and Welsh, 1925) studied large numbers of menhaden fry and their scales in the Gulf of Maine. Summer hatched fish were found to be 6-8 cm . long the first winter and 16cm. long the second winter. Bigelow and Schroeder (1953) reported young men- haden 9.1-9.9 cnn. long at Woods Hole in late September 1942. Fish taken at Falmouth, Massachusetts, 2 months later were 11.7-12.7 cm. long. Welsh (Bigelow and Welsh, 1925) found as many as nine to ten "winter rings" on the scales of a few of the older fish examined and apparently thought that each ring represented 1 year's growth. Rush (1952) found that fish with one annulus at Beaufort, North Carolina, were approximately 18.9 cm. long (fork length). Three-year fish were 28.7cm. long. The oldest fish, 6 years, was 35.8 cm. long. The growth curve was worked out on 34 specimens. Westman and Nigrelli (1955), who worked on Long Island Sound fish, stated that growth rates determined "upon the basis of many times that number of specimens" were very similar. Fish showing a single annulus averaged 18.4 cm. fork length. Fifty-two males of age class III ranged from 29-5 to 34.0 cm. fork length and 52 females of the same age class ranged from 28.5 to 34.5 cm. long. In age class IV 17 males ranged from 31.5 to 35.0 cm. long and 55 females had a fork length range of 31.5 to 36.0 cm. They stated that the conversion factor from fork to total length was 1.1163. Scattergood, Trefethen and Coffin (1951) reported the conversion factor from standard to fork length to be 1.0459. June and Roithmayr (I960) re- ported results of readings of 13,510 scale samples of Atlantic menhaden taken along the east coast of the United States between Portland, Maine, and Fernandina, Florida, from 1952 through 1956. Scale formation commenced at body lengths between 24 and 30 mm., appearing first in the region of the caudal peduncle and later near the base of the pectoral fins and along the posterior margin of the opercles. Scales were fully formed at body lengths between 30 and 43 mm. Growth of scales of young -of -the -year was traced from May until mid-September. The first age ring was formed sometime after the first summer of life. Ring formation occurred only once each year, between March and May. Marginal scale growth occurred during the warm months of the year. The dis- tance between the last submarginal ring and the margin of the scale reached a maximum in the fall. The distance between the last two adjacent ring modes decreased with age. In general, ring formation was found to occur earliest in southern waters and prog- ressively later farther northward. New scale growth varied considerably among individuals, especially in the younger age groups, and this variability is greatest in those fish occurring in southern coastal waters. Hildebrand and Schroeder (1928) said that sexes could not be differ- entiated externally and that the growth rates were apparently much the same. However, Westman and Nigrelli (1955) analyzed the matter statistically and found that there was a significant dif- ference, the females outgrowing the males . Westman and Nigrelli gave fork- length frequency curves for 1/2-month periods from April to June for fish from Long Island Sound and New Jersey waters. In general, the curves showed a decreasing mode from about 30 cm. in April to 25 cm. in June during 1948 and 1949. Ages of menhaden up to the third year of life may be determined by the length-frequency method or by reading scales (McHugh, Oglesby and Pacheco, 1959). Two groups offish, characterized by their size at the^time of formation of the first annulus, entered the Vir- ginia fishery in 1954 and 1955 suggest- ing that growth rates differ according to circumstances. Mean lengths at the end of each calendar year of life were found to be approximately 125, 200 and 230 mm. Mean weights were about 30, 130 and 200 g. Menhaden reached maturity at about 1^ years. Hildebrand and Schroeder (1928) said that fish varied in fatness during the year and were much fatter in the fall than in spring. They also noted that fatness, and presumably growth rate varied from year to year. Fish, taken in October 1921, 13.0 cm. long averaged 0.78 oz. in weight, and two specimens of the same length, taken a year later, averaged 1.07 oz. In 1921, October fish 23.1 cm. long averaged 4.22 oz. and a similar sample for 1922 averaged 5.17 oz. The October 1922 fish weighed an average 15.5 percent more than fish of the same length taken in October 1921. Hildebrand and Schroeder also gave some tables on weight-length relationships. Fish 10.2 cm. long weighed 0.35 oz., those 20.3 cm. long weighed 3.59 oz. and 35.6cm. long weighed 17.1 oz. Goode (1879) measured 30 fish from the same school taken at Noank, Connecticut, in 1875. They ranged from 12 to 13 inches in length and 10 to 16 ounces in weight. The average length was 12.38 inches. Concerning seasonal changes in fatness, Bigelow and Schroeder (1953) said that menhaden were always thin 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 gallons, and less, per thousand fish in the early part of the season to 14^ in August to 16 and 18 gallons in Septem- ber. They stated that New England fish were larger and fatter than those tothe south. These authors also said that the longest menhaden of their knowledge was a specimen 18 inches long taken at Woods Hole in 1876, but that 29-inch fish had been reported. The heaviest fish reported was one from Orient, New York, weighing 1 pound 13 ounces. Goode (1879) said the largest specimen in the National Museum was a 20-inch cast. These remarks donot correspond to those of Ellison (1951) concerning the abundance of 16- to 20 -inch fish in North Carolina waters in late Novem- ber. June and Reintjes (1959 and I960) presented extensive tables of length and weight frequency distributions of sam- ples of Brevoortia tyrannus from the purse -seine and pound-net fisheries along the Atlantic coast from 1952 through 1956. Comparison of the aver- age length and weight of individual year classes from the middle Atlantic area at different ages showed rather marked variation in fish of comparable age in the different years. The average fork length of age 1 fish ranged from 209 mm. in 1952 to 229 mm. in 1955; of age 2, from 233 mm. in 1952 to 259 rnm. in 1955; of age 3, from 279 mm. in 1955 to 286 mm. in 1956; and of age 4, from 291 mm. in 1955 to 304 mm. in 1954. Five-year-old fish averaged 311 mm. fork length in 1956. The most striking feature was that in every year through 1955 fish of the dominant 1951 year class averaged lighter than ad- jacent year classes. In general, there was an increase in the size and weight of fish of the same age group from south to north. (It should be noted, how- ever, that the fishery starts earlier in the south than it does farther north.) The data show that fish of the domi- nant year class which contributed to the catch in each area in 1956 were shorter and lighter than fish of the same age in 1955. Females were larger than males at older ages. Distribution and Migrations Bigelow and Schroeder (1953) re- ported that spawning probably took place every year from the Long Island Sound area to Florida during the period of their observations, and that in some years it extended into the Gulf of Maine. The eggs were buoyant, accord- ing to Welsh (Bigelow and Welsh, 1925) and were found at the surface in waters of high salinity (Perlmutter, 1939). The larvae moved towards shore, where they reached sheltered and often low-salinity areas at a size of around Z5 mm. standard length. Probably they were mostly at the surface during these movements. There were Sonne indica- tions that the larvae moved inshore shortly after coming into the estuaries and moved back out again as juveniles (Massman, Ladd and McCutcheon, 1954). Apparently this movement con- tinued on out into open waters, for in December Z-inch fish, the zeros of Westman and Nigrelli (1955) showed up regularly in North Carolina mixed in with 10-inch fish (Ellison, 1951). Presumably the 2-inch fish came from the north, possibly Chesapeake Bay, because the North Carolina zeros should not be that size in December (Hildebrand, 1920). McHugh, Oglesby and Pacheco (I959) found that menhaden less than 2 years old were present in Virginia waters throughout the year. Spawning probably took place in the ocean, and the young moved into Chesapeake Bay and its estuaries soon after hatching. As they grew, young menhaden moved back down the estuaries toward the sea. In most seasons mean sizes of fish were progressively greater in a seaward direction. Since the purse - seine fishery exploited fish primarily in their second and third year of life and the pound-net fishery took prin- cipally fish in their first and second year, sampling of the pound-net fishery offered a method of forecasting purse - seine catches at least a year in advance. Seasonal waves of emigration and im- migration to Chesapeake Bay were associated with intense fall and spring spawning periods. Females were slightly larger than males of the same age, but the difference was insignifi- cant in young menhaden. There is some information con- cerning the movements of B. tyrannus up and down the Atlantic coast from observations made by biologists and fishermen. Apparently no menhaden remain in offshore waters north of Chesapeake Bay during the colder months of the year. Hildebrand and Schroeder (1928) found that nnenhaden remained in the deeper parts of Chesa- peake Bay during the winter in reduced numbers, where they were taken in beam trawls; in March the fish came into shallow waters and were then taken in pound nets and haul seines. The large schools migrating up and down the coast did not enter the Bay and the local fishery was not affected by spring, summer and fall "runs" as was the case with the outer shore areas of the Atlantic States. According to Bigelow and Schroe- der (1953) nnenhaden left the Maine coast by the middle of October and Massachusetts Bay by early Novennber, although small ones had been taken there as late as December. Reports from fishermen indicated that the fish went around Cape Cod to the New York region and on southward. Ellison (1951) wrote that the migrating schools were fished from Delaware south to North Carolina as follows: "About October 15 a run of fish appears in North Carolina from the north and is joined by fish from the southern sounds and estuaries. These fish run from 10 to 12 inches in length and are known locally as 'Chesapeake Bay' fish, 'holy jump- ers' or 'forerunners.' They contrib- ute to the fishery about a month and are followed about November 10 by the so-called 'Delaware' fish, which measure 13 to 16 inches. These fish, in turn, are succeeded about Thanks- giving by 16-to-20 inch fish recog- nized as the 'Boston Bay' or the 'Amagansett' fish. All of the fish appearing from October 15 to No- vember are following a north-south migration route." In December schools of small fish 2 to 10 inches long appeared in North Carolina. Their source was unknown, but they probably came from inshore waters. Concerning the spring migration northward Ellison said: "The spring fishery usually starts in May, although sometimes in April. This fishery depends principally upon 10 individuals which run from 6 to 8 inches in length and which are be- lieved locally to come up from Flor- ida. Usually these fish strike shore- ward about the latitude of Fernandina, moving north and paralleling the coast, supplying a good fishery at Mayport, Florida. For the past four years, however, they have scarcely touched Mayport, and snapper fisher- men working 30 miles out have re- ported great schools moving north. In these recent years they have struck first off the South Carolina coast about Georgetown and are called in North Carolina the 'Georgetown-flats' fish. These fish support the fishery in North Car olina until August, when they disappear." Ellison further stated that the fish -eached the Chesapeake region in March and April, the New Jersey and New York region in April and May, and the Maine coast in May and June. These dates are much the same as those given by Goode (1879), who reviewed all available information up to that time. Goode's data included precise dates for a 20-year period from some areas of the Atlantic coast. He did not know whether fish went south in the winter or merely offshore. Menhaden do not always go as far north as the Gulf of Maine in summer, and combined data from Bigelow and Welsh ( 1 925) and Bigelow and Schroeder (1953) showed that between 1845 and 1950, inclusive, menhaden had been plentiful to abundant there during only 15 of 66 years reported. It has been generally agreed that temperature governs the north and south migrations of menhaden and that menhaden do not enter waters of tem- peratures less than 50° F. (cf. Ellison, 1951). Goode (1879) collected tem- perature records along the coast and compared them with the time of ap- pearance of the menhaden. This infor- mation led him to state that menhaden appeared after the water temperatures rose to 50° to 51 F. and preferred temperatures between 60° and 70°. Bean (1903) noted that adult menhaden in aquaria died at temperatures lower than 50° F. Bigelow and Schroeder (1953) said their observations cor- roborated the idea that menhaden did not appear in the Gulf of Maine until it was several degrees above 50. Con- versely, falling temperatures drove the fish southward in the fall. Edwards (Kendall, 1910) found that young menhaden could survive cold better than their elders. Fish 2 to 5 inches long survived water tempera- tures of 31.5° F. Better survival of young than of older fish at low tem- peratures was independently observed for other fishes by S. F. Hildebrand and Gordon Gunter following cold spells on the south Atlantic and Gulf coasts. The literature was documented by Gunter (1957). In their long migration from the south Atlantic coast to New England and back, menhaden do not follow pre- cise routes. A "run" may strike shore for many years and then suddenly change and pass as much as 50 miles offshore. Such changes in migration routes may leave established menhaden plants fishless for several years. Elli- son (1951) has given examples. June (1958) studied the variation in meristic characters of young Brevoortia tyrannus to determine whether one or more populations occur in estuarine nursery areas along the Atlantic coast. Vertebrae, ventral scutes, dorsal, and left pectoral fin rays were the meristic characters selected for examination. Counts were made on almost two thousand speci- mens taken from 21 locations between Cape Cod and southern Georgia. Preliminary studies indicated that mean counts do not vary with sex, length of fish over the size range (28-150 mm.) represented, or between right and left pectoral fins. Analysis of variance applied to the meristic data gave no evidence of heterogeneity between the means of successive sam- ples taken in any locality. At least two populations of juvenile menhaden inhabiting the estuarine waters of the Atlantic coast of the 11 United States were indicated, one oc- curring north of Long Island and another south of Long Island. Internningling occurred in the vicinity of Long Island. Difference in mean water temperature during the time of spawning and early larval development may account for the observed differences in mean meristic counts between the two areas. June and Reintjes (1959) noted that the purse-seine fishery depended on the appearance of schools of menhaden at the surface. Fishing started in April on the Florida coast, and by June the fish ranged from northern Florida to the Gulf of Maine. In September and October the schools began todisappear from the most northerly areas, and withdrawal proceeded southward during October. In November and December vast bodies of fish supported a sizable fishery off the North Carolina coast until early January when they once more disappeared fronn coastal sur- face water. Food and Feeding According to Goode (1879) a num- ber of previous workers had observed minute organisms and the presence of chlorophyll in menhaden stomachs. However, other workers he quoted, in- cluding A. E. Verrill, thought such matter was sucked up with the bottom mud. Goode called attention to the fine gill-rakers and the fish's manner of swimming with open nnouth, as if straining the water. The definitive work on feeding and food of the nnenhaden was done by Peck (1894). He showed how the lamel- lae of the gill-rakers overlap "each other in the most perfect manner" and form a fine net for sieving the food from the water. He described a fold of mucous membrane, filled with mucous, which runs along each gill arch at the base of the gill-rakers to form a channel and, presumably, acts as a groove to carry food into the gullet. Peck's statement, "Whether there are definite ciliated tracts with this func- tion of conveying solid particles is not yet known," still holds true. Peck's chief conclusion, which he documented by drawings of plankton in the microscope field, was that menha- den are indiscriminate feeders, and in general take in the same materials in the same proportions as they are found in the water. In the shore waters around Woods Hole, which Peck studied, these materials were predominantly dino- flagellates, diatoms and infusorians, with annelid larvae and crustaceans in lesser abundance. Peck worked mostly with inshore and smaller fish, but found that the young ate the same food as the adults. Peck estimated the volume of water strained by an adult menhaden and arrived at a figure of 7 gallons per minute. He estimated the food material in this amount of water to be 3.4 cc. Hildebrand and Schroeder (1928) summiarized the findings of Dr. Edwin Linton who examined the alimentary contents of 44 fish in Chesapeake Bay, "and found that in most cases they consisted of sandy mud, vegetable debris (mostly algae), and sonne dia- toms, and in a few cases they consisted principally of copepods." The following remarks are taken from Bigelow and Schroeder (1953): "No other Gulf of Maine fish has a filtering apparatus comparable to that of the pogy, nor has it any rival in the Gulf in its utilization of the planktonic vegetable pasture. ". . . And the food eaten at a given locality parallels the general plank- ton content of the water, except that none of the larger aninnals appear in the stomachs of the fish on the one hand, nor the very smallest organ- isms (. . .) on the other." Schroeder (Hildebrand and Schroe- der, 1928) observed that feeding fish swam in circles, rising and falling, renninding him of a whirlwind. Ellison (1951) said the "catholic and non-discriminating taste distin- guishes the menhaden as strongly 12 exceptional, if not unique, among fishes of the sea." Enemies and Mortality The following succinct account is quoted fronn Bigelow and Schroeder (1953): "No wonder the fat oily menhaden, swimming in schools of closely ranked individuals, helpless to pro- tect itself, is the prey of every predaceous animal. Whales and por- poises devour them in large num- bers; sharks are often seen following the pogy schools; pollock, cod, silver hake, and swordfish all take their toll in the Gulf of Maine, as do weakfish south of Cape Cod. Tuna also kill great numbers. But the worst enemy of all is the bluefish, and this is true even in the Gulf of Maine during periods when both bluefish and menhaden are plentiful there. . . Not only do these pirates devour millions of menhaden every summer, but they kill far more than they eat. Besides the toll taken by these natu- ral enemies, menhaden often strand in myriads in shoal water, either in their attempt to escape their enemies or for other reasons, to perish and pollute the air for weeks with the stench of their decaying carcasses ." Hildebrand and Schroeder (1928) said that bluefish migrate up and down the Atlantic coast following schools of menhaden and other fish upon which they feed voraciously. The abundance of menhaden governed the movement of bluefish to some extent. In 1922 young menhaden were plentiful in the lower half of Chesapeake Bay, and the com- mercial catch of bluefish was greater than it had been for many years. Ellison (1951) said the menhaden is "prey to virtually all of those car- nivorous fishes which inhabit the same waters." Goode (1879) was much impressed with predation upon menhaden. He said whales and dolphins ate them by the hogshead. From the air they were attacked by gulls and other sea birds, and the osprey. In fact Goode men- tioned every predator listed in later accounts, including gars and catfish in southern waters. He gave varied ac- counts of attacks by bluefish, and said the menhaden when pursued "often drive in great masses upon the shores ." Goode thought the number destroyed by predators was many times that taken by man. The figure he gave, 3,000 million of millions, equals 3 quadrillion and is certainly a vast overestimate. Ellison (1951) mentioned that blue- fish sometimes ran menhaden into the shallows in such numbers that they died and piled up in windrows 2 feet deep, and polluted the air. Reports of this phenomenon extended from Hatteras to Maine. Local health boards were sometimes required to dispose of the rotting fish. According to Bigelow and Schroeder (1953) dead fish drifted ashore along Massachusetts Bay in 1946 and 1947, and the local health boards had to clean the beaches. However, this kill was attributed to netting by lobster bait fishermen. Westman and Nigrelli (1955) made the following statement concerning an- nual kills of menhaden in New York: "Menhaden appear in the New York bight in commercial quantities during late March or April and, according to reports received, the large fish precede the smaller fish by several weeks. During the latter part of May, or early June, great quantities of menhaden die in these waters and in the waters of western Long Island Sound and litter the beaches in untold numbers. Conse- quently, various agencies call other agencies each year, or write letters, about this phenomenon. These agen- cies, in turn, communicate with other agencies which, in turn, may refer the problem back to the first agency, and so on." Dying fish were called "spin- ners". They were characterized by uncoordinated movements and exop- thalmia. Westman and Nigrelli found 13 hemorrhages caused by gas emboli in capillaries of the gills, eyes and optic lobes of the brain. Bacteria could not be found and the fish entrails were not harmful to cats. Whether the gas emboli were the terminal cause of death or were only an associated con- dition could not be determined. In any case, fish not in distress did not have gas emboli. The authors noted that all mortalities occurred in areas with "markedly varying salinities" and with "organic pollution to a degree where shellfishing is prohibited". Parasites and Diseases Viral and bacterial diseases are unknown in menhaden, and there is no evidence that they are affected by any fatal disease of epidemic proportions. On the other hand menhaden are char- acteristically host to a number of worm and crustacean parasites, which may cause debility but not death. The following parasites of the Atlantic menhaden were listed by West- man and Nigrelli (1955): Parasite Structure attacked Myxosporidian sporozoan: Chloromyxum clupeidae Coccidean sporozoan: Eimeria brevoortiae - - (MS. name) Monogenetic trennatodes: Flesh, Male gonads. (Ellison, 1951) Gills. Stomach. Intestine. Metacercarial cysts in skin. Cysts in viscera. Do. Several ascaroids in intestine and encysted in mesentery. Gills. Body surface. Dn Digenetic trematodes: Podocotyl atomon Trypaborhynchid cestodes: Pterobothrinm heteracanthus Copepods : Bomolochus teres C^/il 'icrtiC nVi ol i fi>')/' ..... — .. — — — — — — — Lernanthropus brevoortiae Lernaeenicus radiatus- - Gills. Body surface. Gills. L^iuveiiiau bpinusu ---- — ___ Olencira praegustator, a parasitic isopod fronri the mouth, was not listed by Westman and Nigrelli, but is very common in fish in the south (Ellison, 1951). O. praegustator is so charac- teristic that its description by Latrobe, along with the original description of B. t yr annus , was one fact indicating the identity of Latrobe's fish, according to Hildebrand (1948). Menhaden Fishery Investigations Goode (1879) wrote of the conflicts between the menhaden fishery and other fisheries. Smith (1896) reported the first examination of other fishes taken with menhaden. Among almost 28,000,000 menhaden, the other fishes constituted 0.0Z8 percent, exclusive of alewives. Including alewives, the figure 14 was 0.336 percent. Most of the clupeids other than menhaden were taken in a few sets when mistaken for schools of nnenhaden. Smith also noted that "the men- haden is a fish which, as a rule, is found in comparatively close proximity to the land. . ." Most of the fish were taken less than 5 miles from shore. The chief predators were found to be bluefish, sharks and weakfish. Migratory movements were oftenmodi- fied by the presence of "such pre- daceous species as bluefish, sque- teague, and sharks" (Smith, 1896). The autumnal migratory movement began with the fish on the shores of Maine and Massachusetts and continued south- ward. Larger and fatter fish were taken on the New England coast than from areas to the south. New England fish produced more oil in the fall than those from the same area produced in the spring. Examination of ovaries tended to verify the existence of different spawning periods on different parts of the coast. GULF OF MEXICO MENHADEN Spawning Kuntz and Radcliffe (1918) noted that the first menhaden larva identified was a specimen 24 mm. long taken on January 15, 1913 by W. W. Welsh in St. George Sound, Carabelle, Florida. St. George Sound is on the northwest Florida coast and the specimen was doubtless B. patronus . Gunter (1945) reported eight Brevoortia patronus 28-59 mm. total length taken on the Gulf beach of Mustang Island, Texas in March and April. Several larvae, tentatively iden- tified as Brevoortia , taken at the same time, were presumed tobe B.patrotlUS- Baldauf (1954) listed menhaden as being one of the fishes which are to be found in the lower Neches River only "during growth stages". Menhaden were taken in the Neches River at a station approximately 30 nautical miles upstream from Texas Point, at the Sabine Pass entrance to the Gulf. Salinities at stations where menhaden were taken ranged from 0.16 to20.4%o. Surface temperature varied from 14.4° to 31° C. Over 10,000 specimens were taken in 1951, 1952 and 1953. The largest menhaden measured was a specimen, found dead on the river bank, with a standard length of 150 mm. The larger fish were B. patronus but spe- cific identification of young individuals was uncertain. Table 5 gives Baldauf's measurements. Baldauf noted that two incoming populations, November-December and March-April, were not the same and that there may have been two spawning peaks. Table 5. — Numbers and sizes of small menhaden taken in the Neches River, 1952-53 [Based on Baldauf, 1954-] Date Number Ranges ( standard length) Mean July- Aug, . Sept. -Oct. Nov Dec Jan Feb Mar Apr 6 0 69 29 322 307 a/roWJ« in the Aransas River, Texas, in March 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 mm. long, were taken in July at 0.5%o salinity. Gunter (1945) in a discussion of Brevoortia sp. (B. gunteri Hildebrand, 1948) taken in Aransas and Copano Bays, Texas, said: 'Renfro, W. C. 1958. The effect of salinity on the distribution of fishes in the Aransas River. Master's Thesis. Graduate School, The University of Texas. 51 p., 3 tables , 25 figs. "Combined total length-frequency curves for both bays were made. In April, 1941 the only group of men- haden caught were from 88to 133 mm. long with a mode at 1 1 3 and 118 mm . This group persisted until October when it was from 128 to 173 mm. long. In August a second group from 88 to 103 mm. long appeared, which became predominant in November and December. Remnants of the two groups persisted through the winter until May when they became more numerous again. At that time, the smaller group had a mode at 128 to 133 mm. Post-larvae came into the catch in January and they were pre- dominant from then until May, when their size ranged from 23 to 43 mm. in length." Springer and Woodburn (I960) compared data from collections in the Tampa Bay, Florida, area with Suttkus' (1956) Lake Pontchartrain data and stated that B, patronus reached Tampa Bay inland waters. . ."at a smaller size (earlier age?) than it does on the Louisiana coast." It should be noted that Springer and Woodburn's collec- tions were made close to high salinity Gulf waters while the inland waters of Lake Pontchartrain are many miles from the open Gulf and high salinity waters where spawning takes place. The data from both areas indicated ". . .a sudden spurt of growth after May (after June?)" by the difference between the average size of July sam- ples. The growth rate of Florida fish seemed to be slower than fish from Louisiana. Distribution and Migrations According to Goode and Bean ( 1 879) the first menhaden from the Gulf re- ceived by the U. S. National Museum were collected in West Florida in 1 864. Goode (1879) under the heading of "Limits in 1877" quoted two light- house keepers, along the Mississippi and Texas coasts, one of whom was a former Maine pogy fisherman, who said that no menhaden "is found in those waters." However, he quoted a 18 pilot, Capt. Wm. Nichols, who said that the waters of Matagorda Bay and the Gulf were full of them and great quantities of menhaden "drifted upon the beach at Saluria" (Texas) in 1872. Whether the fish were alive or dead at the time they stranded is not clear from the context. In the meantime Goode (1878) described Brevoortia patrorms , based upon specimens from Brazos Santiago, Texas, and the mouth of the Rio Grande. Brazos Santiago remains the southernmost record for B.patrOTlUS in the western Gulf. Jordan and Gilbert (1883) listed the menhaden from Pensacola and Gal- veston. Evermann and Kendall (1894) reviewed previous Texas literature and noted one young specimen from Gal ves - ton. Weymouth (1911) listed ten adults and a "considerable number" of young, the latter being doubtfully referred to Brevoortia tyrannus patronus Goode . Most of the collection was made at Calcasieu Pass, Louisiana, which lies in the range of B. gunteri, and Wey- mouth's doubts may have been well founded. Precise dates of collection were not given, and they were probably not known to Weymouth. The fishes were taken by Milo Spaulding during 1906. Fowler (1931) said that a fishing vessel at Port Isabel, Texas, had a small load of B, patronus in August. Gowanloch (1932. 1933), writing about the fishes of Louisiana, noted the abundance 9f menhaden and said that it "is not commercially fished along the Gulf Coast." In 1932 Gunter began counting the fishes taken by the vessel "Black Mallard" in Louisiana. Five otter- trawl hauls were made monthly from the head of Barataria Bay, to 5 miles offshore from Barataria Pass in the open Gulf. Counts were made from January 193 2 to June 1933; John C. Pearson continued thenn until Decem- ber 1933. The first results were pre- sented in a paper on destruction of fishes by shrimp trawls (Gunter, 1936). The tables were unwisely divided into commercial and noncommercial fishes and table 7 is reworked from tables la, lb, Ila and lib of that paper. Menhaden formed 5.8 percent of the 80,093 fishes taken in the bay during both years and 2.4 percent of the 46,518 fishes taken in the shallow Gulf. Most of the bay menhaden were small, and probably were able to escape the trawl when they moved out into the Gulf as larger 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 1933 Nonedible fishes: Menhaden: 1932 1933 Other nonedible fishes: 1932 1933 Total nonedible fishes: 1932 1933 17,309 2^,917 26,264 11,603 '^3,573 36,529 2,166 130 2,296 1,767 352 2,119 12,637 8,626 21,263 13,A65 11,790 25,255 14,803 8,756 23,559 15,232 12,142 27,374 19 Gunter (1938a) gave a further re- port on the data collected aboard the "Black Mallard", including collections made in 1934. Brevoortia patronus v/as the third ranked fish, taken in the bays, and fourteenth in the Gulf. It made up 14.1 percent of the 144,000 fishes caught. On the basis of collections, gen- eral observations, and evidence from mass mortalities, Gunter (1941b) ranked the menhaden as second "in species mass" among fishes in shallow waters of the northern Gulf, preceded by Anchoa mitchilli and followed by Mugil cephaliiS' Gunter (1945) showed that Bre- VOOrtia ranked twelfth among all species caught in Copano and Aransas Bays and off the Gulf beach of Mustang Island, Texas; it ranked ninth in the bay catches. A comparison of trawl catches in Louisiana bays and Texas bays showed that Brevoortia ranked third in Louisiana catches and seventh in Texas. It seemed to be less common in Texas. Fowler (1945) listed B, patronus from Louisiana and Galveston. Jurgens and Hubbs (1953) included B. gunteri as one of the fishes found in the fresh waters of Texas. Knapp (1953) listed B, gunteri as ascending and B. patronus as possibly ascending Texas rivers. H. H. Hildebrand (1953) said men- haden are not often taken in trawls by shrimpers, but sometimes large schools are encountered which burst the trawls. He reported fifteen speci- mens of B. patronus taken in two trawl hauls at 17 and 18 fathoms off south Texas. Reid (1955a), working in East Bay, Texas, noted that where "considerable" water depth and mud bottom extended to the marsh edge, menhaden and various croakers were more abundant while atherinids and cyprinodontids were less. While sampling the Gulf beach at Gilchrist, Texas, across from East Bay, Reid (1955b) found that more menhaden were taken than all other fishes. The 2,664 B. patronus caught made up 58.7 percent of the catch. Their size range was 89 to 198 mm. standard length but only nine were smaller than the largest bay fish. Breuer (1957) said that Brevoortia sp. were not comnnon in the high- salinity Baffin and Alazan Bays of Texas, but that a few large ones appeared in the fall and some small ones in the summer. Temperature and salinity ranges were not given for species reported in Breuer's paper. Gunter (1950) recorded 95 speci- mens of B. gunteri taken along the Blackjack Peninsula shore (Aransas National Wildlife Refuge) of Mesquite and Ayres Bays, Texas from February to June 1945, where they were the third most abundant fish in minnow seines. The average monthly tem- peratures ranged from 19.0°to32.1°C. and salinity range was 9.5-18.3%o. In the connecting salt-flat ponds, only a few miles away, this species was taken only 8 times among 1,324 fishes, and ranked ninth in numbers caught. The average temperatures in that area during the same period were 19.0°- 32.1° C. and the salinities were 4.4- 12.2%o. Caldwell (1954) listed B. gunteri from a salt-marsh pond on Way Key, (Cedar Keys) Florida. Suttkus (1958) said the fish was B. smithi. Rounsefell (1954) said that several species, including "... the nnenhaden, Brevoortia- ■ ." were "nnore or less independent of the waters between the mainland and the barrier islands." H. H. Hildebrand (1955) reported eight specimens of B. gunteri from two trawl hauls in 7-8 fathoms of water off Punta Morros in the SE portion of the Gulf of Campeche. They were taken February 15, 1951. This is the south- ernmost record for nnenhaden in the Gulf. Hildebrand said he had never heard of menhaden schools south of the Rio Grande. 20 Gunter (1956b) recorded B, smithi, B. giinteri and B. patronus as being euryhaline . Hubbs (1957) in a revision of a former checklist included Brevoovtia gunteri among fishes to be found in fresh waters of Texas. Briggs (1958) listed B. smithi and B. tyrannus from the Atlantic coast of Florida and said B. tyrannus was to be found on both sides of the Atlantic. He listed B, patronus from Tampa north- ward and B. gunteri from Cedar Keys (probably on Caldwell's record) to Campeche . Suttkus (1958) discussed the dis- tribution of three species of menhaden in the Gulf. In summary he said: "The two fine-scaled species, Brevoortia gunteri and B. smithi, occur in the western and eastern Gulf respectively. The single large- scaled form, B, patronus, overlaps B. gunteri in the western Gulf from Brazos Santiago, Texas, to Grand Isle, Louisiana, and overlaps B, smithi in the eastern Gulf at Cedar Keys." Springer and Woodburn (I960) found young B. patrOHUS and young B, smithi together in the Tampa Bay, Florida, area. Concerning the move- ment of young B. patronus they said: "In contrast to Suttkus' finding that emigration from estuarine waters took place in July and August, we found no specimens in the bayou after May. . . . The collection from Johns Pass. . . during July does indicate that migration to the Gulf had occur- red at least this early and possibly as early as June." Food and Feeding Gowanloch (1933). Gunter (1945) and Reid (1955a) stated that GuM Br e- VOOrtia are plankton feeders. Darnell (1958) examined the stomach contents of 17 specimens of B. patrOTtUS from Lake Pontchartrain, Louisiana. All contained food nnaterial. Food of small menhaden (38-48 mm.) consisted chiefly of phytoplankton with small amounts of zooplankton and plant fragments. Sev- eral species of Anabaena comprised 77 percent of the total food while the remaining phytoplankton was com- posed of diatoms. Detritus constituted 11 percent of the stomach contents. The small menhaden had obviously been straining plankton at the surface and apparently at depths below the surface . The larger menhaden (85- 103 mnn.) examined by Darnell (1958) contained, in the muscular pyloric region of the stomach, 99 percent ground organic matter and silt with a few diatoms, foramniferans and copepods. Suspen- sions of ground up organic matter were observed, particularly along the south shore of the lake "where wave action was reducing the organic material of the marshy shore to the consistency of coffee grounds." Darnell concluded that menhaden fed by filtration, and that suspended bacteria and material other than living plankton were an important component of the food of menhaden in turbid estuaries. Enemies and Mortality Arthur (1931) commented on men- haden as food of the brown pelican in the following words: "The pelican is a fish eater. While it may be true that the pelican, once in a while, may catch what man terms a 'food fish*, the number of such fish of commercial value it consumes does not justify the prej- udice felt against the big bird. When the World War was on and the Food Administration was calling on the people to 'eat more fish', some fishermen, residents of Florida, asked that an edict be issued calling for the pelican's extermination on the theory that it was to blame for a food-fish shortage. But before this wholesale slaughter was ordered, cooler heads counseled inquiry and in the summer of 1918 the Depart- ment of Conservation conducted a 21 survey of the Brown Pelican breeding places, and secured exact data as to what this particular pelican ate, when fish of all kinds were most plentiful in the Gulf waters. "The Louisiana coast from Pearl River to the Sabine was explored and pelicans and their food collected and sent to Washington, D. C. for identification by the United States Bureau of Fisheries. The result of the examination of many hundreds of stomachs showed that 97 per cent of the fish were menhaden or 'gulf sardines', and 3 per cent were silver- sides - not a single food fish used by man was found!" Lund (1936) reported that about 85 percent of the dead fish reported on the south Texas coast in the sumnner of 1935 were menhaden. Facts reported by Lund and others verbally to the senior author indicate, with little doubt, that this mortality was the result of a "red tide". Gunter {1941a) noted that, although menhaden were listed as a common bay fish in previous papers, only three were observed killed by the hard cold ■wave of January 1940 on the Texas coast. These were seen in Copano Bay and were listed as B. patTOnuS- The specimens could have been B. gunteTi. Gunter (1945) found no menhaden in the stomachs of 23 7 redfish (SciaenopS ocellata) examined. He found three Brevoortia in the stomachs of 153 Cynoscion nehulosus. Gunter, Davis, Williams and Smith (1948) recorded ten B. /JOfronMS killed by the red tide along 20 yards of Fort Myers beach, among 208 other fishes. The species could have been B. Smithi. Reid (1955a) found that 89 percent of "white trout" (Cynoscionaretmrius) stomachs contained fish "and menhaden constituted the greatest part of this item." He indicated that the menhaden soon grow too large to be taken by "trout". He said menhaden were also preyed upon to a lesser extent by the croaker, two nnarine catfishes, the ten- pounder, ElopS saurus, and the lizard- fish, Synodus foetens. It was concluded that predation on young fish, especially menhaden, is severe. Reid (1955b) also pointed out that "birds of the area also exact a toll on these fish." Parasites and Diseases Pearse ( 1 952a) described the para- sitic copepod, Caligiis ventrosetus , from the gills of B. gunteri at Port Aransas, Texas. He also listed Lemanthropus brevoortiae from B. patronus at the same locality. Pearse (1952b) recorded Lemanthropus brevoortiae from the gills of B. patronus in Florida. Causey (1953) reported Lernaeenicus radiatus from B. Louisiana. 'tyrannus" at Grand Isle, Causey (1955) reported the para- sitic copepods, Lemanthropus brevo- ortiae and L. radiatus from B. patronus at Pascagoula, Mississippi. Hargis (1955) listed the monogenetic trema- tode, Mazocraeoides georgei, from the gills of B. patronus at Alligator Harbor, Florida, as a new host record. Koratha (I955) described a new monogenetic trematode, Diclidophora lintoni, from the gills of B. gunteri taken at Port Aransas, Texas. Hargis (1957) listed the following monogenetic trematodes (Family Ma- zocraeidae) as parasitic on Brevoortia patronus: Clupeocotyle brevoortia, C. megaconfibula, Kuhnia brevoortia and Mazocraeoides georgei . Sparks (1958) listed the digenetic tr emai.tod.&. Par ahemiurus merus.irom the gills of B. patronus a.s a new host record. Menhaden Fishery Investigations Miles and Simmons (1950) gave a review of the menhaden industry and fishery. They also summarized certain biological information collected by a series of workers at the Marine Lab- oratory of the Texas Game and Fish Commission and published a series of mimeographed reports by Breuer, 22 Anderson, Knapp, Wilson, and Miles and Simmons. Miles and Simmons, give the following information: 1. Other sports and commercial fishes were taken in very small quan- tities with menhaden. As the result of one study the figure given was one animal to 4,490 menhaden or 0.024 percent, including shrimp and crabs. The fishes caught with 5,326,000 men- haden were 36 gafftopsails (Bagre marina), 75 crabs, 91 jacks (Caranx hippos), 103 croakers {Micropogoti un- dulatus), 191 shrimp, 205 Spanish mackerel (Scontberomorus maculatus) , 242 sand trout (Cynoscion), 304 blue- fish (Pomatomus saltatrix), and a few others, all less than 8 each. The total was 7,589 other fishes. Another study showed that 2,183 other animals, including shrimp, crabs and squid were taken with 2,500,000 menhaden. 2. Mackerel were found with smaller menhaden and bluefish were found with larger menhaden in offshore waters. Bluefish appeared when the snnaller menhaden were replaced by larger ones in July. Crabs, shrimp, sand trout and croakers were usually taken in shallow waters. Bluefish, mackerel, tarpon, jacks and sharks seenned to be the main predators on menhaden. 3. Forty species of fish, com- prising 26,005 individuals were ex- amined for stomach contents. Of these, 13,288 were sea trout, Cynoscionnebu- losus 3,428 were Spanish mackerel and 3,137 were redfish, Sciaenops ocellata. The remainder were 38 other species of fishes. They contained 581 menhaden. The percentages eaten by the common fishes were given. Menhaden were not preferred food of the common sports and comnnercial fishes. 4. Knapp showed that the stomachs of menhaden, mullets and "shad", remaining intact in the stomachs of predators after other parts had been digested, could be identified by their s hape . Knapp (1950) presented some of the same data and came to similar conclusions. Filipich (I947) made five unan- nounced checks of menhaden boat un- loadings at Mississippi factories. In 295 tons of menhaden unloaded, one mackerel and 6 white trout were found. Gowanloch (1949) reviewed the menhaden-sports fisherman contro- versy and concluded that no harm was done to sports fishes by the menhaden industry. Christmas, Gunter and Whatley (i960) reported a study of fishes other than menhaden taken in menhaden purse seines in waters around the mouth of the Mississippi River and in Missis- sippi Sound during the 1958 and 1959 seasons. Samples were taken from catches which totaled nearly 2 million pounds. In numbers of fish, more than 97 percent of the sampled catch were menhaden. The other fishes observed included 62 species. Brevoortia pa- tyonuS was the only menhaden found in the catch. The fishery was generally prosecuted in shallow, low-salinity waters. Seventy percent of the sampled menhaden catches were made in waters of salinities between 5 and 24 parts per thousand. Surface water temperatures at the sampled seine locations varied from 22.6° to 30.5° C. Lower tem- peratures were encountered at the beginning and the end of the fishing season. Fishing started in May when water temperatures rose to About 23° C . and stopped in the fall when surface waters reached approximately the same temperature again. SUMMARY Three of the five North American species of menhaden have been found in the Gulf of Mexico. The present distribution, based on the published record, is: 1. Brevoortia gunteri, Grdiridlsle, Louisiana westward and southward to the Gulf of Campeche. 23 2. Brevoortiapatronns , the Tampa Bay area, Florida to Brazos Santiago, Texas. 3. BreVOOrtia smithi, Cedar Keys, Florida southward and presumably around the peninsula to the Atlantic. (R. D. Suttkus, personal communica- tion.) Menhaden fishing in the Gulf of Mexico is dependent on B, patvonus , and is mostly prosecuted in Louisiana waters. Gulf landings of menhaden increased rapidly with the expansion of the fishing until 1957, when landings were only 65 percent of the 1956 catch. However, 1958 landings increased to about 80 percent of the 1956 record. Apparently the 1957 decrease was caused by factors other than depletion of the population. All information agrees with the supposition that Atlantic menhaden spawn at sea, or at least in high salini- ties, and the larvae work in to shore to spend their early life, very often in low salinity or even fresh water. Ap- parently spawning is also governed by temperature. Comparison of surface temperatures (Bumpus, 1957) for re- ported months of spawning along the Atlantic coast indicates that spawning takes place in the Atlantic at about the same temperature in all localities. Thus, the spawning season is during the summer on the New England coast and in the winter off the South Atlantic States. There are no records of spawning B, patvonUS and presumably they spawn at sea. Except for one report of ten- tatively identified Brevoortia larvae on the Gulf beach in March and April in south Texas, there are no reports of menhaden larvae on the beach. On the other hand, vast numbers of larvae and juvenile menhaden have been re- ported in the bay waters of Louisiana and Texas. Low -salinity waters un- doubtedly serve as nursery grounds. Brevoortia gunteri in spawning condition have been reported inCopano and upper Aransas Bays, Texas, in February and March. Spawning B. gunteri have been reported from the Laguna Madre, Texas, in February. While there may be confusion be- tween both the larvae and the young of two species where their range over- laps, reports oi B. patronus 16-25 mm. long extend from November to June, with minimum sizes at both ends of the period. There is some evidence of a double influx of larvae . Thus the spawn- ing season is long, over the fall and winter and into the spring. B. gunteri seem to spawn in the spring in Texas waters. Menhaden eggs and early larvae have not been reported in the Gulf. Presumably, the eggs are bouyant like those in the Atlantic and the larvae remain near the surface, as has been reported in Virginia. The slender larvae which enter estuarine waters undergo metamorpho- sis between 20 and 30 mm. standard length in Lake Pontchartrain. Data on growth rates are scarce. A group thought to be Br evoortia gunteri ranged from 88-133 mm. long in April and 128- 1 73 mm . long in August in Texas waters . A group of Texas Brevoortia 88-103 mm. long in August had a mode of 128- 133 mm. the following May. On the Atlantic coast, menhaden undertake extensive migrations up and down the coast with changes in season, during which they are followed by the fishermen, at least on the trip south. Nothing similar is known in the Gulf, probably because it is an east-west coast. Large menhaden are not comnnon in the bays and evidently the young fish move out, as they grow up, mostly in late summer and fall. Salinity is a critical factor in the development of larval Atlantic menhaden, with low salinity waters being essential to nor- mal development. In Texas, larvae and juveniles 21-48 mm. long have been found at salinities of 2.0-20.4%o and temperatures of 1 5.5°-3 1 .0° C . In White Lake, Louisiana, menhaden larvae have been found in virtually fresh water. 24 The numbers of larvae found in the bays may vary enornnously fronn year to year. There may be sonne con- nection between the heavy 1956 com- mercial production of menhaden and Reid's report of "staggering" num- bers of larvae in East Bay, Texas, in the summer of the same year. There is only one report on the food of Gulf menhaden. It was found that Lake Pontchartrain menhaden fed by filtration and consumed considerable quantities of detritus and suspended bacteria in addition to living plankton. Menhaden in the Gulf of Mexico are not known to be chased ashore by predators or to die in recurring sum- mer maladies from unknown causes as they do in the Atlantic. They have been killed in considerable numbers by the red tide in Texas and Florida but these instances are outside the commercial range. Menhaden seenn to be resistant to sudden cold waves which decimate other fishes in Texas and Florida, and none were reported killed in three studies of cold-killed fishes in Florida and Texas. A few menhaden have been reported killed by the cold on the Texas coast. Menhaden fishermen on the Gulf catch few other fish, comparable to the situation on the Atlantic coast. The chief menhaden enemies seem to be bluefish, mackerel, sharks, and tarpon. The common shore and shallow- water sports and commercial fishes seem not to subsist to any great extent on larger menhaden, but predation upon small menhaden in the bays seems to be very heavy. LITERATURE CITED ANDERSON, ANDREW W., and C. E. PETERSON. 1952. Fishery statistics of the United States, 1949. U. 5. Fish and Wildlife Service, Statistical Digest 25, 298 p. 1953. Fishery statistics of the United States, 1950. U. S. Fish and Wildlife Service, Statistical Digest 27, 492 p. ANDERSON, ANDREW W., and C. E. PETERSON. 1954. Fishery statistics of the United States, 1951. U. S. Fish and Wildlife Service, Statistical Digest 30, 341 p. ANDERSON, A. W., and E. A. POWER. 1949. Fishery statistics of the United States. 1945. U. S. Fish and Wildlife Service, Statistical Digest 18, 372 p. 1951. Fishery statistics of the United States, 1948. U. S. Fish and Wildlife Service, Statistical Digest 22, 304 p. 1954. Fishery statistics of the United States, 1951. U. S. Fish and Wildlife Service, Statistical Digest 30, 341 p. 1955. Fishery statistics of the United States, 1952. U. S. Fish and Wildlife Service, Statistical Digest 34, 345 p. 1956a. Fishery statistics of the United States, 1953. U. S. Fish and Wildlife Service, Statistical Digest 36, 338 p. 1956b. Fishery statistics of the United States, 1954. U. S. Fish and Wildlife Service, Statistical Digest 39, 446 p. 1957. Fishery statistics of the United States, 1955. U. S. Fish and Wildlife Service, Statistical Digest 41, 446 p. ARTHUR, S. C. 1931. The birds of Louisiana. Loui- siana Conservation Department, Bulletin No. 20, 598 p. BAKER, EMMA (Comp.). 1955. Full text of the game fish and fur laws of Texas. Texas Game and Fish Commission, Austin. 319 p. BALDAUF, R. J. 1954. Survey and study of surface and subsurface conditions in and around Beaumont, Texas. Bio- logical survey of the Neches River in the region of Beau- mont, Texas. Texas A. and M. Research Foundation. [Unre- leased mimeographed report.] 184 p. 25 BEAN, T. H. 1903. Catalogue of the fishes of New York. New York State Museum, Bulletin No. 60, Zoology 9, 784 p. BIGELOW, HENRY B., and WILLIAM C. SCHROEDER. 1953. Fishes of the Gulf of Maine. U. S. Fish and Wildlife Service, Fishery Bulletin No. 74, vol. 53, 577 p. BIGELOW, HENRY B., and W. W. WELSH. 1925. Fishes of the Gulf of Maine. Bulletin of the U. S. Bureau of Fisheries, 1924, vol. 40, part 1, 567 p. BREUER, J. P. 1957. An ecological survey of Baffin and Alazan Bays, Texas. Publi- cations of the Institute of Marine Science, University of Texas, vol. 4, no. 2, p. 134-155. BRIGGS, J. C. 1958. A list of Florida fishes and their distribution. Bulletin of the Florida State Museum, Biologi- cal Sciences, vol. 2, no. 8, p. 223-318. BUMPUS, DEAN F. 1957. Surface water temperatures along Atlantic and Gulf coasts of the United States. U. S. Fish and Wildlife Service, Special Scien- tific Report--Fisheries No. 214, 153 p. CALDWELL, D. K. 1954. Addition to the known fish fauna in the vicinity of Cedar Key, Florida. Quarterly Journal of the Florida Academy of Sciences, vol. 17, no. 3, p. 181-184. CAUSEY, D. 1953. Parasitic Copepoda from Grand Isle, Louisiana. Occa- sional Papers of the Marine Laboratory, Louisiana State Uni- versity, No. 7, 18 p. 1955. Parasitic Copepoda from Gulf of Mexico fish. Occasional Papers of the Marine Labora- tory, Louisiana State University, No. 9, 19 p. CHRISTMAS, J. Y., GORDON GUNTER, and EDWARD C. WHATLEY. I960. Fishes taken in the menhaden fishery of Alabama, Mississippi, and eastern Louisiana. U. S. Fish and Wildlife Service, Spe- cial Scientific Report- -Fish- eries No. 339, 10 p. DARNELL, REZNEAT M. 1958. Food habits of fishes and large invertebrates of Lake Pon- chartrain, Louisiana, an es- tuarine connnn unity. Publications of the Institute of Marine Science, University of Texas, vol. 5, p. 353-416. DE BUEN, FERNANDO. 1958u Peces de la superfamilia Clupeoidea en aguas de Chile. Revista de Biologia Marina, vol. 8, no. 1, 2, 3, p. 83-110. ELLISON, W. A., JR. 1951. The menhaden. In Survey of marine fisheries of North Caro- lina, by H. F. Taylor and as- sociates. University of North Carolina Press, Chapel Hill, p. 85-107. EVERMANN, B. W., and W. C. KEN- DALL. 1894. The fishes of Texas and the Rio Grande basin, considered chiefly with reference to their geographic distribution. Bulletin of the U. S. Fish Commission, vol. 12 (1892), p. 57-126. FIEDLER, R. H. 1943. Fishery statistics of the United States, 1940. U. S. Fish and Wildlife Service, Statistical Digest 4, 225 p. FILIPICH, MECO. 1947. Report to the Mississippi Sea- food Connnnission, Biloxi, Mis- sissippi. Menhaden Plants Re- port, 4 p. FOWLER, HENRY W. 1931. A collection of fishes from the Texas coast. Copeia, 1931, no. 2 (July), p. 46-50. 26 FOWLER, HENRY W. 1936. The marine fishes of West Africa. Bulletin of the American Museum of Natural History, vol. 70, part 1, 605 p. 1945. A study of the fishes of the southern Piedmont and coastal plain. Monographs, Academy of Natural Sciences of Philadelphia, No. 7, 408 p. GILL, THEODORE N. 1861. Synopsis of the subfamily Clupeinae, with descriptions of new genera. Proceedings of the Academy of Natural Sciences of Philadelphia, p. 33-38. GOODE, GEORGE BROWN. 1878. A revision of the American species of the genus Brevoortia with a description of a new species from the Gulf of Mexico. Proceedings of the UnitedStates National Museum, vol. 1, p. 30-42. 1879- The natural and economical history of the American men- haden. U. S. Commission of Fish and Fisheries, Part 5, Report of the Commissioner for 1877. Ap- pendix A, The Sea Fisheries, Section I, A History of the Men- haden, p. 1-529. 31 plates. 1884. The food fishes of the United States. In the Fisheries and Fishery Industries of the United States, Section 1, part 3, p. 163- 682. U. S. Comnnission of Fish and Fisheries, Washington. GOODE, GEORGE BROWN, and THEO- DORE H. BEAN. 1879- Catalogue of a collection of fishes sent from Pensacola, Florida, and vicinity, by Mr. Silas Stearns, with descriptions of six new species. Proceedings of the United States National Museum, vol. 2, p. 121-156. GOWANLOCH, J. N. 1932. Sea fishes and fishing in Loui- siana. Louisiana Conservation Department, Bulletin No. 21, 187 p. GOWANLOCH, J. N. 1933. Fishes and fishing in Loui- siana. Louisiana Conservation Department, Bulletin No. 23, 638 p. 1949. Menhaden facts and fallacies. Advertiser Publishing Company, Pascagoula, Miss. 26 p. GUNTER, GORDON. 1936. Studies of the destruction of marine fish by shrimp trawlers in Louisiana. Louisiana Conser- vation Review, vol. 5, no. 4, p. 18-23, 45-46. 1938a. The relative numbers of species of marine fish on the Louisiana coast. American Naturalist, vol. 62, p. 77-83. 1938b. Seasonal variations in abun- dance of certain estuarine and marine fishes in Louisiana, with particular reference to life histories. Ecological Mon- ographs, vol. 8, p. 313-346. 1941a. Death of fishes due to cold on the Texas coast, January, 1940. Ecology, vol. 22, no. 2, p. 203-208. 1941b. Relative nunnbers of shallow water fishes of the northern Gulf of Mexico, with some records of rare fishes from the Texas coast. American Midland Naturalist, vol. 26, no. 1, p. 194-200. 1945. Studies on marine fishes of Texas. Publications of the In- stitute of Marine Science, Uni- versity of Texas, vol. 1, no. 1, p. 1-190. 1 950. Distributions and abundance of fishes on the Aransas National Wildlife Refuge, with life history notes. Publications of the In- stitute of Marine Science, Uni- versity of Texas, vol. 1, no. 2, p. 89-101. 1956a. Some relations of faunal dis- tributions to salinity in estuarine waters. Ecology, vol. 37, no. 3, p. 616-619. 1956b. A revised list of euryhaline fishes of North and Middle America. American Midland Naturalist, vol. 56, no. 2, p. 345-354. 27 GUNTER, GORDON. 1957. Temperature. In Treatise on marine ecology and pale- oecology, vol. 1, Chapter 8, p. 159-184. The Geological Society of America, Memoir 67, New York. GUNTER, GORDON, C. C. DAVIS, R.H. WILLIAMS, and F. G. WALTON SMITH. 1948. Catastrophic mass mortality of marine animals and coincident phytoplankton bloom on the west coast of Florida, November 1946-August 1947. Ecological Monographs, vol. 18, p. 309-327. GUNTER, G., and W. E. SHELL, JR. 1958. A study of an estuarine area with water-level control in the Louisiana marsh. Proceedings of the Louisiana Academy of Sciences, vol. 21, p. 5-34. HARGIS, W. J., JR. 1955. Monogenetic trematodes of Gulf of Mexico fishes. Part VII. The super-family Diclid- ophoroidea Price, 1936 (con- tinued). Quarterly Journal of the Florida Academy of Sciences, vol. 18, no. 2, p. 113-119. 1957. The host specificity of mono- genetic trematodes. Experinnen- tal Parasitology, vol. 6, p. 610- 625. HILDEBRAND, H. H. 1953. A study of the fauna of the brown shrimp {Penaeus aztecus Ives) grounds in the western Gulf of Mexico. Publications of the Institute of Marine Science, University of Texas, vol. 3. no. 2, p. 228-366. 1 map. 1955. A study of the pink shrinnp [Penaeus duorarum Burkenroad) grounds in the Gulf of Cannpeche, Publications of the Institute of Marine Science, University of Texas, vol. 4, no. 1, p. 169-302. HILDEBRAND, SAMUEL F. 1920. Two species of menhaden oc- curring on the coast of North Carolina. U. S. Bureau of Fish- eries, Report of the United States Commissioner of Fisheries for the fiscal year 1918, Appendix 6, 8 p. HILDEBRAND, SAMUEL F. 1941. An annotated list of salt and brackish water fishes, with a new name for a menhaden, found in North Carolina since the pub- lication of "The fishes of North Carolina" by Hugh M. Smith in 1907. Copeia, 1941, no. 4 (No- vember), p. 220-232. 1948. A review of the American menhaden. Genus Brevoortia, with a description of a new spe- cies. Smithsonian Miscellaneous Collections, vol. 107.no. 18, 39 p. HILDEBRAND, SAMUEL F., and W. C. SCHROEDER. 1928. Fishes of Chesapeake Bay. Bulletin of the U. S. Bureau of Fisheries, 1927, vol. 43, part 1, 366 p. HUBBS. CLARK. 1957. A checklist of Texas fresh- water fishes. Texas Game and Fish Commission, IF Series No. 3, 11 p. JORDAN, DAVID STARR, and B. W. EVERMANN. 1896. The fishes of North and Mid- dle America. United States Na- tional Museum, Bulletin No. 47. 4 vols. JORDAN, DAVID STARR, and C. H. GILBERT. 1883. Notes on fishes observed about Pensacola, Florida and Galves- ton, Texas, with descriptions of new species. Proceedings of the United States National Museum, vol. 5, no. 282, p. 241-301. JUNE, FRED C. 1958. Variation in meristic charac- ters of young Atlantic menhaden, Brevoortia tyrannus. Rapports et Proces-Verbaux des Reunions. Conseil Permanent International pour 1 'Exploration de la Mer, vol. 143, part 2, p. 26-35. JUNE, FRED C, and L. LOCKWOOD CHAMBERLIN 1959. The role of the estuary in the life history of the Atlantic men- haden. Proceedings of the Gulf and Caribbean Fisheries Insti- tute, Eleventh Annual Session, 1958, p. 41-45. 28 JUNE, FRED C, and JOHN W. REINTJES. 1959. Age and size composition of the menhaden catch along the Atlantic coast of the United States, 195Z-55; with a brief re- view of the commercial fishery. U. S. Fish and Wildlife Service, Special Scientific Report--Fish- eries No. 317, 65 p. 1960. Age and size composition of the menhaden catch along the Atlantic coast of the United States, 1956; with a brief review of the commercial fishery. U.S. Fish and Wildlife Service, Spe- cial Scientific Report--Fish- eries No. 336, 38 p. JUNE, FRED C, and CHARLES M. ROITHMAYR. I960. Determining age of Atlantic menhaden from their scales. U. S. Fish and Wildlife Service, Fishery Bulletin No. 1 71, vol. 60, p. 323-342. JURGENS, K. E., and CLARK HUBBS. 1953. A checklist of Texas fresh- water fishes. Texas Game and Fish, vol. 11, no. 4, p. 12-15. KENDALL, W. D. 1910. Effects of the menhaden and mackerel fisheries upon the fish supply. Bulletin of the U. S. Bu- reau of Fisheries, 1908, vol.28, part 1, p. 279-293. KNAPP, F. T. 1950. Menhaden utilization in rela- tion to the conservation of food and game fishes of the Texas Gulf coast. Transactions of the Annerican Fisheries Society, vol. 79, p. 137-144. 1953. Fishes found in the fresh- waters of Texas. Ragland Studio and Litho Printing Co., Bruns- wick, Georgia. 166 p. KORATHA, K. J. 1955. Studies on mohogenetic tre- matodes of the Texas coast. II. Descriptions of species from marine fishes of Port Aransas. Publications of the Institute of Marine Science, University of Texas, vol. 4. no. 1, p. 250-278. KUNTZ, A., and L. RADCLIFFE. I9I8. Notes on the embryology and larval development of twelve teleostean fishes. Bulletin of the U. S. Bureau ofFisheries, 1915- I9I6, vol. 35, p. 87-134. LUND, E. J. 1936. Some facts relating to the oc- currence of dead and dying fish on the Texas coast during June, July and August, 1935. Annual Report of the Game, Fish and Oyster Commission of Texas, 1934-35, p. 47-50. McHUGH, J. L., R. T. OGLESBY, and A. L. PACHECO. 1959. Length, weight, and age com- position of the menhaden catch in Virginia waters. Limnology and Oceanography, vol. 4, no. 2, p. 145-162. MASSMANN, W. H., E. C. LADD, and H. N. McCUTCHEON. 1954. Postlarvae and young of the menhaden (Brevoortia tyrannus) in brackish and fresh waters of Virginia. Copeia, 1954 no. 1 (February), p. 19-23. MILES, D. W., and E. G. SIMMONS. 1950. The menhaden fishery. Texas Garne, Fishand Oyster Commis- sion, Bulletin No. 30, 28 p. PEARSE, A. S. 1952a. Parasitic Crustacea from the Texas coast. Publications of the Institute of Marine Science, Uni- versity of Texas, vol. 2, no. 2, p. 5-42. 1952b. Parasitic crustaceans from Alligator Harbor, Florida. Quarterly Journal of the Florida Academy of Sciences, vol. 15, no. 4, p. 187-243. PEARSON, J. C. 1950. The young of some marine fishes taken in lower Chesapeake Bay, Virginia, with special ref- erence to the gray sea trout, Cynoscion regalis (Bloch).U.S. Fish and Wildlife Service, Fish- ery Bulletin No. 36, vol. 50, p. 79-102. 29 PECK, J. I. 1894. On the food of the menhaden. Bulletin of the U. S. Fish Com- mission, 1893, vol. 13, p. 113- 126. 8 plates. PERLMUTTER, A. 1939. An ecological survey of young fish and eggs identified from townet collections. In A biologi- cal survey of the salt waters of Long Island. Part II (1938), New York State Conservation Depart- ment, p. 11-71. POWER, E. A. 1958. Fishery statistics of the United States, 1956. U. S. Fish and Wildlife Service, Statistical Digest No. 43, 476 p. 1959. Fishery statistics of the United States, 1957. U. S. Fish and Wildlife Service, Statistical Digest No. 44, 429 p. REGAN, C. TATE. 1917. A revision of the clupeoid fishes of the genera PomolohlCS , Brevoortia and Dorosoma, and their allies. Annals and Maga- zine of Natural History, 8th Series, vol. 18, p. 297-316. REID, G. K., JR. 1955a. A summer study of the biol- ogy and ecology of East Bay, Texas. Texas Journal of Science, vol. 7, no. 3, p. 316-343. 1955b. A summer study of the biol- ogy and ecology of East Bay, Texas. Part II. The fish fauna of East Bay, the Gulf beach and summary. Texas Journal of Science, vol. 7, no. 4, p. 430-453. 1956. Ecological investigations in a disturbed Texas coastal estuary. Texas Journal of Science, vol. 8, no. 3, p. 296-327. 1957. Biologic andhydrographic ad- justment in a disturbed Gulf coast estuary. Limnology and Oceanography, vol. 2, no. 3, p. 198-212. REINTJES. JOHN W., JAMES Y. CHRISTMAS, JR., and RICHARD A. COLLINS. I960. Annotated bibliography on bi- ology of American menhaden. U. S. Fish and Wildlife Service, Fishery Bulletin No. 170, vol. 60, p. 297-322. ROUNSEFELL, G. A. 1954. Biology of the commercial fishes of the Gulf of Mexico. In Gulf of Mexico, its origin, waters, and marine life. U. S. Fish and Wildlife Service, Fish- ery Bulletin No. 89, vol. 55, p. 507-512. RUSH, W. A. 1952. Observations of age and growth in the menhaden (Bve- voortia tyrannus) as determined by scale examination. Copeia, 1952, no. 3 (September), p. 208- 209. SCATTERGOOD, L. W., P. S. TRE- FETHEN, and G. W. COFFIN. 1951. Notes on the size of menhaden taken in Maine during 1949. Copeia, 1951 no. 1 (March), p. 93-94. SIMMONS, E. G. 1957. An ecological survey of the upper Laguna Madre of Texas. Publications of the Institute of Marine Science, University of Texas, vol. 4, no. 2, p. 156-200. SMITH, H. M. 1896. Notes on an investigation of the menhaden fishery in 1894, with special reference to the food-fishes taken. Bulletin of the U. S. Fish Commission, 1895, vol. 15, p. 285-302. 1907. The fishes of North Carolina. North Carolina Geological and Economic Survey, vol. 2, 453 p. 30 SPARKS, A. K. 1958. Some digenetic trematodes of fishes of Grand Isle, Louisiana. Proceedings of the Louisiana Academy of Sciences, vol. 20, p. 71-77. SPRINGER, V. G., AND K. D. WOOD- BURN I960. An ecological study of the fishes of the Tampa Bay area. Florida State Board of Conserva- tion, Marine Laboratory, Pro- fessional Papers Series No. 1, 104 p. SUTTKUS, R. D. 1956. Early life history of the Gulf menhaden, Brevoortia patronus, in Louisiana. Transactions of the 21st North Annerican Wild- life Conference, p. 390-407. 1958. Distribution of menhaden in the Gulf of Mexico. Transactions of the 23d North American Wild- life Conference, p. 401-410. THOMPSON, WILL F. 1916. Fishes collected by the United States Bureau of Fishes steamer ''Albatross" during 1888 be- tween Montevideo, Uruguay, and Tome, Chile, on the voyage through the Straits of Magellan. Proceedings of the United States National Museum, vol. 50, no. 2133, p. 401-426. U. S. FISH AND WILDLIFE SERVICE 1959. Gulf fisheries, 1958, annual summary. U. S. Fish and Wild- life Service, Current Fishery Statistics No. 2165, 12 p. WARFEL, H. E.,andD. L. MERRIMAN. 1944. Studies on the marine re- sources of southern New Eng- land. An analysis of the fish of the shore zone. Bulletin of the Bingham Oceanographic Collec- tion, vol. 9, art. 2, p. 35-42. WESTMAN, J. R., andM. H. BIDWELL. 1948. Waste disposal and fisheries of the salt waters adjacent to the Greater New York metropolitan area. New York State Depart- ment of Conservation. [Mime- ographed report.] WESTMAN, J. R.. and R. F. NIGRELLI. 1955. Preliminary studies of men- haden and their mass mortalities in Long Island and New Jersey waters. New York Fish and Game Journal, vol. 2, no. 2, p. 142-153. WEYMOUTH, F. W. 1911. Notes on a collection of fishes from Cameron, Louisiana. Pro- ceedings of the United States National Museum, vol. 38, no. 1734, p. 135-145. 31 MS #972 GPO 902933 MBL WMOl [ ilir.-irv Scri.ils ilill IJIIIIJii 5 WHSE 01491