Spanish Mackerel (8

BULLETIN OF MARINE SCIENCE, 41(3): 822-834, ]987

LARVAL KING MACKEREL (SCOMBEROMORUS CAVALLA), SPANISH MACKEREL (8. MACULATUS), AND BLUEFISH (POMATOMUS SALTATRIX) OFF THE SOUTHEAST COAST OF THE UNITED STATES, 1973-1980

Mark R. Collins and Bruce W Stender

ABSTRACT Surface and subsurface ichthyoplankton collections were made from 9 m to beyond the continental shelf(deepest station 3,940 m) in all seasons from Cape Hatteras, North Carolina to Cape Canaveral, Florida. King mackerel spawn from April to at least September, primarily at depths >40 m. Spring spawning activity takes place further offshore than does summer spawning. An apparent concentration of larvae between 32° and 33°N suggests that the area of upwelling associated with the Charleston bump is an important spawning and/or nursery area. Spanish mackerel spawn from May to September in depths <40 m. Larvae were less abundant than those of king mackerel, and no areas of concentration were found. Vertical migration to the surface at night is indicated for both king and Spanish mackerels. Bluefish spawn bimodally from March through at least November in depths >40 m, with the primary spawning peak in spring and the secondary peak in late summer. In spring, larvae were caught most often between 32° and 33°N, but in summer-fall were taken more often at locations further south. Neither vertical migration or visually-cued net avoidance is indicated, but bluefish >4 mm are strongly associated with the surface.

Spanish (Scomberomorus maculatus) and king (S. cavalla) mackerels and bluefish (Pomatomus saltatrix) support large recreational and commercial fisheries along the east coast of the United States. All three species, particularly S, maculatus and P. saltatrix, occur near shore as adults and are available to pier, shore, and small boat anglers. Depletion of stocks, especially of S. cavalla, is of particular region-wide concern. The Gulf and South Atlantic Fishery Management Councils have established a management plan with associated quotas and restrictions for this species; however, management efforts are based upon the limited life history information currently available. Improvements in the data base could enhance future modifications of this plan. Several studies have addressed the distribution of larval S. cavalla and S. maculatus in the Gulf of Mexico (Dwinell and Futch, 1973; Houde et aI., 1979; McEachran et aI., 1980) and bluefish in the Atlantic off the eastern U.S. (Lund and Maltezos, 1970; Norcross et aI., 1974). One or more of these species have been included in faunal surveys along portions of the Atlantic coast (Herman, 1963; Fahay, 1975). In addition Powles (1981), working with a portion (1973- 1976) of the present data, and Kendall and Walford (1979) reported on the occurrence of bluefish larvae in the Atlantic off the southeastern U.S. Most inves- tigations conducted off the southeast coast of the U.S. in the area known as the South Atlantic Bight (SAB) (Blumberg and Mellor, 1983; Weisberg and Pietrafesa, 1983), however, have been limited to a one- or two-year sampling period or restricted in terms of depth range, gear utilization and/or seasonality, Thus, relatively little is known about spawning locations and the distribution of larvae of S. cavalla, S. maculatus and P. saltatrix in this region. The intent of this report is to summarize the distribution, abundance, and occurrence oflarval king mackerel, Spanish mackerel, and bluefish in the SAB from 1973 through 1980.

822 COLLINS AND STENDER: EAST COAST LARVAL FISHES 823

81. W 80. W 79. W 78. W 77. W

••.• .: .• °0

"P:..... 33. N ... 33. N , .•. , .• '\"...... :"...." " ~l

° ••• 0

32. N 2. N

31. N 31. N

30. N O. H

29. N 29. H

28. N 28. N 82. W 81. W 80. W 79. W 78. W 77. W 76. W Figure 1. Locations of ichthyoplankton collections, 1973-1980.

METHODS

During 1973-1980, the Marine Resources Monitoring, Assessment and Prediction (MARMAP) program of the South Carolina Marine Resources Research Institute conducted quantitative surveys to investigate the distribution and abundance of ichthyoplankton throughout the SAB. A total of 1,163 collections was made from Cape Hatteras, North Carolina to Cape Canaveral, Florida, and from 9 m to beyond the continental shelf over depths as great as 3,490 m (Fig. I). Cruises were carried out aboard the R/V DOLPHIN. Three types of gear were utilized during these cruises: (I) a 1.0 x 0.5 m neuston net with 505 /.1mmesh towed half-submerged, (2) a 2.0 x 1.0 m neuston net with 947 /.1m mesh towed half-submerged, and (3) a bongo frame with 0.6 m diameter nets of 505 /.1mand 333 /.1m mesh towed in double-oblique fashion from 0 to .:5 200 m. For the purposes of this study, no distinction was made between the two surface-towed nets, and only the 505 /.1mbongo sample was sorted for ichthyoplankton. For more specific survey information, see Jossi et al. (1975) and Powles and Stender (1976). Samples were filtered and preserved at sea in 5% buffered formalin and sorted in the laboratory using Bogorov trays under dissecting microscopes. All lengths given refer to notochord length on pre flexion larvae and standard length on flexion and postflexion larvae. The smallest and largest 824 BULLETIN OF MARINE SCIENCE, VOL. 41, NO.3, 1987

Table I. Ichthyoplankton collections 1973-1980 by light phase, gear type and season

Light phase Day Night Dawn/dusk Season Surface Bongo Surface Bongo Surface Bongo Winter 73 72 89 88 39 40 Spring 53 46 42 34 22 21 Summer 117 114 83 82 39 41 Fall 12 10 18 13 II 4

individuals (therefore, minimum and maximum lengths) in each sample were measured to obtain a range oflengths rather than complete length frequency data, and the total number of each species was recorded for each sample. Data on abundance were standardized to concentration (#/1,000 m) using duration, speed, and effective sampling area for surface tows and using calibrated General Oceanics digital flowmeters for bongo tows. When calculating monthly mean catches of bongo collections, abundance data were also standardized to number of larvae under 100 m' of surface area for tabular comparison to concentration values. All references to standardized catch are in terms of concentration. Data were separated for analyses on the basis of gear (bongo vs. neuston), station depth, season, and diel period. Depth zones wcre designated as inner-shelf (s 20 m), middle-shelf (21-40 m), outer- shelf (41-200 m), and off-shelf (> 200 m) areas, and were chosen on the basis of previous hydrographic studies in the SAB (Atkinson et aI., 1985). Seasonal separations were based on generally accepted hydrographic seasons: January-March for winter, April-June for spring, July-September for summer, and October-December for fall. Collections were made in all months except June and December. Dusk and dawn were defined as sunset ± I h and sunrise ± I h, respectively, with night and day as the remaining periods. Table I presents the number of collections made by gear, season, and light phase. Data from all years were pooled for analyses, and all statistical tests were non parametric. The Mann- Whitney (M-W) and Chi-square (x') tests were used to compare standardized catches and frequencies of occurrence, respectively, by shelf area, diel period, and latitude. The Spearman rank correlation coefficient (SR) was used to test for correlations of standardized catch and length with depth and latitude. Minimum and maximum length in each sample were used in all tests involving lengths since length frequency data were not available. For all statistical tests, P was required to be s.05 for significance.

RESULTS Scomberomorus cavalla. -King mackerel larvae (2-14 mm) were taken in 105 collections (Fig. 2), 39 of which were surface tows, for a total of 459 individuals. Larvae occurred from April to September (no June collections) and in November. Individuals ~4 mm in length occurred from May to September. September produced the highest percentage of collections containing larval S. cavalla (24.3%) and, in neuston tows, the greatest monthly mean standardized catch of larvae. Abundance was slightly greater in August than September in bongo collections (Table 2). Collections containing larvae ~4 mm also occurred most frequently in September (21.4%). No larval S. cavalla occurred in the 20 October collections. Frequency of occurrence of S. cavalla was higher in night than in day collections with both bongo and surface nets. There was no significant difference between day and night frequencies for bongo tows, but surface tows produced S. cavalla significantly more often during night than day (x2: P < 0.001). During the day, but not during the night, bongo tows caught S. cavalla more often than surface tows (X2: P < 0.001). King mackerel larvae were taken most often in the outer-shelf area with both gear types. Considering only the months in which larvae occurred, 32% of bongo and 44% of night surface collections from the outer shelf contained larval S. cavalla, while only 2 of the total 175 collections in the inner-shelf area (1.1%) COLLINS AND STENDER: EAST COAST LARVAL FISHES 825

76. W 82. W 81. W 80. W 79. W 78. W 77. W H. N '. N

33. N 33. N

32. N 32. N

31. N 1. N

30. N O. N

29. N 29. N

28. N 28. N

82. W 81. W 80. W 79. W 78. W 77. W 76. W Figure 2. Locations of ichthyoplankton collections containing Scomberomorus cavalla. con tained larvae. When com pared to the other shelf areas, frequency of occurrence in bongo collections was significantly greater in the outer than inner-shelf area (x2: P < 0.00 I), but was not significantly different from the middle and off-shelf areas. In surface collections, occurrence was significantly greater in the outer than the inner (x2: P < 0.05) and middle-shelf (X2: P < 0.01) areas, but was not significantly different from the off-shelf area. Minimum length was negatively correlated with station depth in spring bongo collections (SR: P < 0.05). The smallest larvae (2 mm) were taken at stations with depths of 32-512 m. The comparison of occurrence (both gears combined) within each degree of latitude to that in the remainder of the study area showed that only between 32°N and 33°N was the occurrence significantly greater than expected (X2: P < 0.05) based on the latitudinal distribution of effort. A repetition of this test omitting the inner-shelf area, since only two larvae were taken there, further supported this observation on latitudinal distribution of larvae (X2: P < 0.01). Standardized 826 BULLETIN OF MARINE SCIENCE, VOL. 41, NO.3, 1987

Table 2. Monthly mean catches as #/1,000 m3 water filtered (V) and # under 100 m2 surface area (A) for bongo collections, and as #/ 1,000 m3 for neuston collections. Months of peak occurrence denoted by asterisk

Pomatomus sa/tatrix ScomberomontS cavalla Scomberomorus maculalus Month Bongo Neuston Bongo Neuston Bongo Neuston

V 0.59 A 2 V A 3 V 4.63 3.18 A 62.50 4 V 44.81 * 45.73 0.59 A 453.12 5 V 53.96 * 41.53 18.36 7.36 2.95 A 395.95 252.52 6 V A 7 V 10.19 15.29 38.41 * A 28.49 55.00 8 V 17.65 29.15 * 17.81 14.47 * 15.73 A 114.58 20.91 9 V 15.61 21.57 28.07 * 25.07 30.68 * 28.24 A 55.01 164.81 38.57 10 V 7.07 A II V 9.50 3.15 0.59 A 48.97 12 V A

catches in bongo collections were also significantly greater in this area than in the rest of the SAB (M-W: P < 0.005). In addition, standardized catches of bongo collections containing larvae :::;4 mm were significantly greater than in the rest of the SAB (M-W: P < 0.05) in summer, the season of greatest abundance and occurrence. Locations of collections containing larvae :::;4 mm are presented in Figure 3. Scomberomorus maculatus. - Ninety-six larval Spanish mackerel (2-9 mm) were taken in 26 collections, 11 of which were surface tows, from May to September (no June collections). Small larvae (:::;4 mm) occurred throughout this period. Month of peak occurrence was September with 16 collections, but the greatest monthly mean standardized catch was in July (Table 2) because the two collections that included S. maculatus involved relatively large standardized catches com- pared to other months. Spanish mackerel larvae were taken only in the inner and middle-shelf areas, with the deepest station at 29 m and shallowest at 11 m. Twenty of the 26 collections (76.9%) were in the inner-shelf area, but within this area there were no significant correlations between length or standardized catch and depth. There was also no significant variation of occurrence or abundance with latitude. Lo- cations of collections of S. maculatus and of those containing S. maculatus :::;4 mm are presented in Figures 4 and 5, respectively. Frequencies of occurrence of S. maculatus larvae in bongo collections were COLLINS AND STENDER: EAST COAST LARVAL FISHES 827

82. W 77. W 76. W 81. W 80. W 79. W 78. W ,. N 3'. N ,!. ,!. ,!.

"',!. ~ .,!. 33. N ,!. 33. N

32. N 32. N

31. N 31. N ,!. ,!.,!. t ,!. ,!.

30. N ,!. 30. N

29. N 29. N

28. N 8. N

82. W 81. W 80. W 79. W 78. W 77. W 76. W Figure 3. Locations of ichthyoplankton collections containing Scomberomorus cavalla larvae ~ 4 mm.

nearly identical in day and night and there was no significant difference in standardized catch. In surface collections, however, occurrence was significantly greater (X2: P < 0.01) in night as none were caught during day. Pomatomus sa/tatrix. - Ninety-three collections, 65 of which were surface tows, produced 3,881 bluefish 2-35 mm in length. Larval P. sa/tatrix were taken in January, March-May, and August-November (no June or December collections were made). Larvae ~4 mm were taken in 32 collections during April, May, August, September, and November. Spring was the peak season in both occurrence and standardized catch (Table 2) for all bluefish larvae (68 spring collections) and for larvae ~4 mm (22 spring collections). Locations of collections containing bluefish are presented in Figure 6, and of collections containing bluefish ~4 mm in Figure 7. 828 BULLETIN OF MARINE SCIENCE, VOl. 41, NO.3, 1987

76. W 82. w 81. W 80. W 79. W 78. lit 77. W ~,. N '. N

~~. N ~. N

~2. N ~2. N

~1. N ~1. N

~O. N O. N

29. N 29. N

28. N 8. N

82. W 81. W 80. W 79. W 78. W 77. W 76. W Figure 4. Locations of ichthyoplankton collections containing Scomberomorus macula/us.

There were no significant differences in abundance or occurrence of bluefish between day and night for either gear. Surface collections produced bluefish significantly more often than bongo (x2: P < 0.001). However, when only those collections containing small (::54 mm) larvae are considered, there was no significant difference in occurrence between gears. The highest frequency of occurrence was in the outer-shelf area with both gears. There was no significant difference in occurrence between outer and off-shelf areas for either gear and occurrence was significantly greater in the outer than inner- shelf area for both gears (X2: P < 0.00 1 for both). In bongo but not surface collections, occurrence was significantly greater in the outer than middle-shelf area (x2: P < 0.001). With one exception, all collections containing larvae ::54 mm were taken outside the inner-shelf area. In surface collections, minimum length was negatively correlated with station COLLINS AND STENDER: EAST COAST LARVAL FISHES 829

82. W 81. W 80. W 79. W 78. W 77. W 76. W H. N ,. N CAPE FEAR '" '" ~ ~~. N '" '" ~~. N

~2. N ~2. N

~1. N ~1. N

~O. N ~O. N

29. N 29. N

28. N 28. N

82. W 81. W 80. W 79. W 78. W 77. W 76. W Figure 5. Locations of ichthyoplankton collections containing Scomberomorus maculatus larvae 054 mm.

depth in fall (SR: P < 0.05), winter (P < 0.05), and spring (P < 0.001). In spring surface tows in the outer-shelf area, maximum length was negatively correlated with latitude in all light phases combined (SR: P < 0.05) and during the day (SR: P < 0.005), and standardized catch was positively correlated with latitude in night collections (SR: P < 0.05). However, when only peak conditions are considered (spring surface collections during the day made at 32°N and above), both standardized catch and maximum length were negatively correlated with latitude (SR: P < 0.05 and P < 0.005, respectively). Frequency of occurrence of larvae ~4 mm between 32° and 33°N was significantly greater than expected in relation to the latitudinal distribution of collections in spring (x2: P < 0.01). Of the 22 spring collections containing larvae ~4 mm, only two were south of 32°N while 8 of 10 summer-fall collections were made south of 32°N. 830 BULLETIN OF MARINE SCIENCE, VOL. 41, NO.3, 1987

82. W 81. W 80. W 3'. N

33. N

32. N

31. N 1. N

30. N 30. N

29. N 9. N

28. N 28. N 82. W 81. W 80. W 79. W 78. W 77. W 76. W Figure 6. Locations of ichthyoplankton collections containing Pomatomus saltatrix.

DISCUSSION Scomberomorus caval/a. - Based on occurrence of larvae, summer is the primary spawning season for king mackerel in the SAB, with September the peak month. From data on gonad condition, however, Finucane et al. (1986) concluded that peak spawning activity should occur in July, although the temporal range of activity was identical to that in the present study. In the northeastern (Dwinell and Futch, 1973) and northwestern (McEachran et aI., 1980) Gulf of Mexico, larvae were caught from May to October, and September was the month of greatest abundance. The significantly greater occurrence of king mackerel larvae in bongo tows during the day and in neuston tows at night indicates that vertical migration to the surface at night is taking place. McEachran et al. (1980) found no significant differences in numbers or lengths of larvae between day and night bongo collec- COLLINS AND STENDER: EAST COAST LARVAL FISHES 831

81. W 80. W 79. W 78. W 77. W

33. N 33. N

32. N 2. N

31. N 1. N

30. N O. N

29. N 9. N

28. 28. N

82. W 81. W 80. W 79. W 78. W 77. W 76. W

Figure 7. Locations of ichthyoplankton collections containing Pomatomus sa/tatrix larvae :$4 mm in spring (triangles) and summer-fall (squares). tions, and Fahay (1975) reported that postlarval scombrids were caught in surface nets more often at night. Thus, all available evidence supports vertical migration as the primary factor influencing die) differences in catch of S. caval/a. Based on the distribution of collections containing newly hatched larvae (2 mm), king mackerel spawn over a wide depth range. However, only two larvae were collected in the inner-shelf area, suggesting that the entire larval phase of S. caval/a is spent offshore in the middle-shelf area and beyond. These data support the conclusions of McEachran et al. (1980) that king mackerel larvae are more abundant in depths> 35 m. The negative correlation between standard length (minimum) and station depth was observed only in spring and suggests that spring spawning takes place further offshore than later spawning activity. In addition, of the 61 larvae collected during spring only 3 were taken in depths <329 m. 832 BULLETIN OF MARINE SCIENCE, VOL. 41, NO.3, 1987

Thus, the data indicate spawning beyond the continental shelfin spring, followed by inshore movement. We propose the hypothesis that spawning well offshore during spring may be due to higher water temperatures in the Gulf Stream, and that the warming of inshore waters in summer may reduce the tendency to spawn offshore. Some spawning activity apparently occurs in all parts of the SAB. However, catches were significantly more frequent and larvae significantly more abundant between 32° and 33°N. Significant features of this area include a semi-permanent deflection of the Gulf Stream and a region of upwelling that are produced by the topographic ridge known as the "Charleston Bump" (Atkinson and Targett, 1983; Pietrafesa et aI., 1985). Concentrations of fish larvae have been noted (Fahay, 1975) and fish aggregations acoustically detected (Atkinson and Targett, 1983) in this area. Evidence from the present study suggests that the region between 32° and 33°N, seaward of the inner-shelf area, is either a preferred spawning location or that larvae spawned elsewhere are concentrated in the area by currents. It is likely that the former explanation is at least partially responsible for the concentration since small larvae (=:;4 mm) were significantly more abundant in this area than in the rest of the SAB in summer. Scomberomorus maculatus. - That the major spawning period of Spanish mackerel extends from May to September is corroborated by data on gonad condition offish from the SAB and Gulf of Mexico (Finucane and Collins, 1986). Spawning activities of Spanish and king mackerels are temporally similar but geographically separate. Considering the smallest (2 mm) larvae, there was no depth overlap in collections, with all S. maculatus being taken inshore of the shallowest collection of S. caval/a. Studies in the Gulf of Mexico have also reported larval S. maculatus as most abundant in inshore waters (McEachran et aI., 1980; Dwinell and Futch, 1973; Houde et aI., 1979), but information on their distribution in the SAB is lacking. Spanish mackerel were more common than king mackerel in the eastern Gulf of Mexico (Houde et aI., 1979), but in the western Gulf (McEachran et aI., 1980) and in the present study the reverse was true. However, since the shallowest sample sites in the present study were in approximately 9 m, it is probable that larvae are concentrated further inshore. The variation in occurrence of S. maculatus with gear and light phase was identical to that of S. caval/a. Vertical migration to the surface at night, rather than net avoidance, is indicated as the primary factor affecting diel catch differences. Unlike S. caval/a, there were no significant latitudinal concentrations of larval S. maculatus, nor correlations of length with station depth. The small sample size, probably due to lack of sampling effort between the surf zone and 9 m, may have influenced these conclusions. Additional near-shore ichthyoplankton studies in the SAB would aid in determining whether there are concentrations of Spanish mackerel larvae in waters shallower than those included in the present study. Pomatomus saltatrix. - Both abundance and occurrence of larval bluefish peaked in spring (Table 2), supporting the statement by Kendall and Walford (1979) that a spawning concentration exists during spring in the SAB during northward migration of adult bluefish. A break in spawning activity occurred in early summer, followed by an increase in activity in September with small (:54 mm) larvae taken through November. These results closely agree with the conclusions of Finucane and Collins (in press) based on gonad condition of bluefish from Georgia and the Carolinas. Most (80%) collections of small larvae in summer-fall were south of the area of larval concentration in spring. The summer-fall spawning to the south COLLINS AND STENDER: EAST COAST LARVAL FISHES 833 of the primary spring spawning area may represent a separate population resident to the SAB, as hypothesized by Kendall and Walford (1979), or may represent repeat spawning by the same migratory population. Tagging and reproductive studies during both spring and summer-fall spawning seasons would be of great help in answering this question. Significantly greater occurrence oflarvae :54 mm between 32°and 33°N suggests that the region of upwelling produced by the Charleston bump may be of im- portance as a spawning and/or nursery area for bluefish, as well as for king mackerel. Further support for this hypothesis exists in the form of a positive correlation between standardized catch and latitude for the SAB as a whole but a negative correlation when only the region north of 32°N is considered. Kendall and Walford (1979) reported taking large numbers of larvae in spring between 33° and 34°N, similar to the results of the present study but differing by one degree of latitude. There was no evidence for either vertical migration or diel variation in net avoidance by bluefish. That a strong association with the surface exists (Kendall and Walford, 1979) is confirmed by greater occurrence in surface than in bongo collections. However, the smallest larvae (:54 mm) were caught with equal frequency by both gears, suggesting that recently hatched bluefish utilize a larger portion of the water column than do older larvae. Nearly all spawning activity seems to occur outside the inner-shelf area in all seasons, and collections were most frequent seaward of the 40 m isobath. Powles (1981) stated that the relationship between length and station depth was relatively weak for bluefish, and Kendall and Walford (1979) concluded that larvae from the spring spawning in the SAB are carried northward past Cape Hatteras before moving shoreward. Data from the present study do not support either of these previous conclusions. Length was negatively correlated with station depth in all seasons but summer and the relationship was especially strong (P < 0.001) in spring, indicating shoreward movement with growth. In the outer shelf area, length was negatively correlated with latitude during spring throughout the SAB, as well as in the area north of 32°N, indicating southward movement of bluefish larvae rather than northward. We suggest that the southerly countercurrent located shoreward of the Gulf Stream (Bumpus, 1973) may be the mechanism for this implied movement. No evidence for loss from the SAB of spring-spawned bluefish larvae was found.

ACKNOWLEDGMENTS

We thank the many persons who assisted with ichthyoplankton collections. This study was sponsored by the National Marine Fisheries Service (Southeast Fisheries Center) under Contract 6-35147, and by the South Carolina Wildlife and Marine Resources Department. This is Contribution No. 227 of the South Carolina Marine Resources Center.

LITERATURE CITED

Atkinson, L. P. and T. E. Targett. 1983. Upwelling along the 60 m isobath from Cape Canaveral to Cape Hatteras and its relationship to fish distribution. Deep-Sea Res. 30: 221-226. --, D. W. Menzel and K. A. Bush, eds. 1985. Oceanography of the southeastern U.S. continental shelf. American Geophysical Union, Washington, D.C. 156 pp. Blumberg, A. F., and G. L. Mellor. 1983. Diagnostic and prognostic numerical circulation studies of the South Atlantic Bight. J. Geophys. Res. 88: 4579-4592. Bumpus, D. F. 1973. A description of the circulation on the continental shelf of the east coast of the United States. Prog. Oceanogr. 6: 111-157. Dwinell, S. E. and C. R. Futch. 1973. Spanish and king mackerel larvae and juveniles in the northeastern Gulf of Mexico June through October 1969. Fla. Dept. Nat. Res. Mar. Res. Lab. Leaflet Ser. 5 part I (24): 1-14. 834 BULLETINOFMARINESCIENCE,VOL.41, NO.3, 1987

Fahay, M. P. 1975. An annotated list oflarval and juvenile fishes captured with surface-towed meter net in the South Atlantic Bight during four R/V DOLPHINcruises between May 1967 and February 1968. NOAA Tech. Rep. NMFS SSRF-685. 39 pp. Finucane, J. H. and L. A. Collins. 1986. Reproduction of Spanish mackerel, Scomberomorus maculatus, from the southeastern United States. Northeast Gulf Sci. 8: 97-106. --- and ---. In Press. Reproductive biology of bluefish, Pomatomus saltatrix, from the southeastern United States. Northeast Gulf Sci. ---, ---, H. A. Brusher and C. H. Saloman. 1986. Reproductive biology of king mackerel, Scomberomorus caval/a. from the southeastern United States. Fish. Bull. U.S. 84: 841-850. Herman, S. S. 1963. Planktonic fish eggs and larvae of Narragansett Bay. Limnol. Oceanogr. 8: 103- 109. Houde, E. D., J. C. Leak, C. E. Dowd, S. A. Berkeley and W. J. Richards. 1979. Ichthyoplankton abundance and diversity in the eastern Gulf of Mexico. Rep. Bur. Land Management Cont. No. AA550-CT7-28, June 1979. 546 pp. Jossi, J. W., R. R. Marak and H. Peterson, Jr. 1975. MARMAP survey I manual. At sea data collection and laboratory procedures. MAR MAP Prog. Off., NMFS, Washington, D.C. 113 pp. Kendall, A. W., Jr. and L. A. Walford. 1979. Sources and distribution of bluefish, Pomatomus saltatrix, larvae and juveniles off the east coast of the United States. Fish Bull. U.S. 77: 213-277. Lund, W. A., Jr. and G. C. MaItezos. 1970. Movements and migration of the bluefish, Pomatomus saltatrix, tagged in waters of New York and southern New England. Trans. Am. Fish. Soc. 99: 719-725. McEachran, J. D., J. H. Finucane and L. W. Hall. 1980. Distribution, seasonality and abundance of king and Spanish mackerel larvae in the northwestern Gulf of Mexico (Pisces, Scombridae). Northeast Gulf Sci. 4: 1-16. Norcross, 1. J., S. L. Richardson, W. H. Massmann and E. B. Joseph. 1974. Development of young bluefish (Pomatomus saltatrix) and distribution of eggs and young in Virginian coastal waters. Trans. Am. Fish. Soc. 103: 477-497. Pietrafesa, L. J., G. S. Janowitz, and P. A. Wittman. 1985. Physical oceanographic processes in the Carolina capes. Pages 23-32 in L. P. Atkinson, D. W. Menzel and K. A. Bush, eds. Oceanography of the southeastern U.S. continental shelf. American Geophysical Union, Washington, D.C. Powles, H. 1981. Distribution and movements of neustonic young of estuarine dependent (Mugil spp., Pomatomus saltatrix) and estuarine independent (Coryphaena spp.) fishes off the southeastern United States. Rapp. P.-v. Reun. Cons. Int. Explor. Mer., 178: 207-209. --- and B. W. Stender. 1976. Observations on composition, seasonality and distribution of ichthyoplankton from MARMAP cruises in the South Atlantic Bight in 1973. S. C. Mar. Resour. Center Tech. Rep. No. II: 1-47. Weisberg, R. H. and L. J. Pietrafesa. 1983. Kinematics and correlation of the surface wind field in the South Atlantic Bight. J. Geophys. Res. 88: 4593-4610.

DATEACCEPTED: March II, 1987.

ADDRESS: South Carolina Wildlife and Marine Resources Department, Marine Resources Research Institute, P.O. Box 12559, Charleston, South Carolina 29412.