NOAA Technical Report NMFS 79 July 1989

Species Composition, Distribution,

and Relative Abundance of Fishes

in the Coastal Habitat off the

Southeastern United States·

Charles A. Wenner

George R. Sedberry

U.S. Department of Commerce
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34. Additions to a revision of the shark genus Carcharhinus: Synonymy of Aprionodon and Hypoprion, and description of a new species of Carcharhinus (Carcharhinidae),

hy J. A. F. Garrick. November 1985, 26 p.

35. Synoptic review of the literature on the Southern oyster drill Thais haemastoma

floridana, by Philip A. Butler. November 1985, 9 p.

36. An egg production method for estimating spawning biomass of pelagic fish:

Application to the northern anchovy, Engraulis mordax, by Reuben Lasker (editor).

December 1985,99 p.

37. A histopathologic evaluation of gross lesions excised from commercially important North Atlantic marine fishes, by Robert A. Murchelano, Linda DespresPatanjo, and John Ziskows!d. March 1986, 14 p.

38. Fishery atlas of the northwestern Hawaiian Islands, by Richard N. Uchida and

James H. Uchiyama (editors). September 1986, 142 p.

39. Survey of fish protective facilities at water withdrawal sites on the Snake and

Columbia Rivers, by George A. Swan, Tommy G. Withrow, and Donn L. Park. April

1986, 34 p.

40. Potential impact of ocean thermal energy conversion (OTEC) on fisheries, by

Edward P. Myers, Donald E. Hoss, Walter M. Matsumoto, David S. Peters, Michael

P. Se!d, Richard N. Uchida, John D. Ditmars, and Robert A. Paddock. June 1986,

33 p.

41. A stationary visual census technique for quantitatively assessing community structure of coral reef fishes, by James A. Bohnsack and Scott P. Bannerol. July 1986, 15 p.

42. Effects oftemperature on the biology ofthe northern shrimp, Pandalus borealis,

in the Gulfof Maine, by Spencer Apollonio, David K. Stevenson, and Earl E. Dunton,

Jr. September 1986, 22 p.

43. Environment and resources of seamounts in the North Pacific, by Richard N.

Uchida, Sigeiti Hayasi, and George W. Boehlert (editors). September 1986, 105 p.

44. Synopsis of biological data on the porgies, Calamus arctifrons and C. proridens

(Pisces: Sparidae), by George H. Darcy. September 1986, 19 p.

45. Meristic variation in Sebastes (Scorpaenidae), with an analysis of character

association and bilateral pattern and their significance in species separation, by Lochai Chen. September 1986, 17 p.

46. Distribution and relative abundance of pelagic nonsalmonid nekton off Oregon

and Washington 1979-84, by Richard D. Brodeur and William G. Pearcy. December

1986, 85 p.

47. Reproduction, maturation, and seed production of cultured species: Proceedings

of the twelfth U.S.-Japan meeting on aquaculture, Baton Rouge, Louisiana, October

25-29, 1983, by Carl J. Sindermann (editor). February 1987, 73 p.

48. Widow rockfish: Proceedings of a workshop, Tiburon, California, December

11-12, 1980, by William H. Lenan and Donald R. Gunderson (editors). January 1987,

57 p.

49. Reproduction, movements, and population dynamics of the southern !dngfish,

Menticirrhus americanus, in the northwestern Gulf of Mexico, by Stephen M. Harding

and Mark E. Chittenden, Jr. March 1987,21 p.

50. Preparation of acetate peels of valves from the ocean quahog, Arctica islandica,

for age determinations, by John W. Ropes. March 1987,5 p.

51. Status, biology, and ecology of fur seals. Proceedings of an international

workshop, Cambridge, England, 23-27 April 1984, by John P. Croxall and Roger

L. Gentry (editors). June 1987,212 p.

52. Limited access alternatives for the Pacific groundfish fishery, by Daniel D.

Huppert (editor). May 1987, 45 p.

53. Ecology of east Florida sea turtles: Proceedings of the Cape Canaveral, Florida,

sea turtle workshop, Miami, Florida, February 26-27, 1985, by Wayne N. Witzell

(convener and editor). May 1987, 80 p.

54. Proximate and fatty acid composition of 40 southeastern U.S. fmfish species,

by Janet A. Gooch, Malcolm B. Hale, Thomas Brown, Jr., James C. Bonnet, Cheryl

G. Brand, and Lloyd W. Reiger. June 1987, 23 p.

55. Proximate composition, energy, fatty acid, sodium, and cholesterol content of

finfish, shellfish, and their products, by Judith Krzynowek and Jenny Murphy. July

1987, 53 p.

56. Some aspects of the ecology of the leatherback turtle Dermochelys coriacea at

Laguna Jolova, Costa Rica, by Harold F. Hirth and Larry H. Ogren. July 1987, 14 p.

57. Food habits and dietary variability of pelagic nekton offOregon and Washington,

1979-1984, by Richard D. Brodeur, Harriet V. Lorz, and William G. Pearcy. July

1987, 32 p.

58. Stock assessment of the Gulf of Menhaden, Brevoonia patronus, fishery, by

Douglas S. Vaughan. September 1987, 18 p.

59. Atlantic Menhaden, Brevoonia ryrannus, purse seine fishery, 1972-84, with a

brief discussion of age and size composition of the landings, by Joseph W. Smith,

William R. Nicholson, Douglas S. Vaughan, Donnie L. Dudley, and Ethel A. Hall.

September 1987, 23 p.

60. Gulf Menhaden, Brevoortia patronus, purse seine fishery, 1974-85, with a brief

discussion of age and size composition of the landings, by Joseph W. Smith, Eldon

J. Levi, Douglas S. Vaughan, and Ethen A. Hall. December 1987, 8 p.

61. Manual for starch gel electrophoresis: A method for the detection of genetic variation, by Paul B. Aebersold, Gary A. Winans, David J. Teel, George B. Milner, and

Fred M. Utter. December 1987, 19 p.

62. Fishery publication index, 1980-85; Technical memoradum index, 1972-85, by

Cynthia S. Martin, Shelley E. Arenas, Jac!d A. Guffey, and Joni M. Packard.

December 1987, 149 p.

63. Stock assessment of the Atlantic Menhaden, Brevoonia ryrannus, fishery, by

Douglas S. Vaughan and Joseph W. Smith. January 1988, 18 p.

64. illustrated key to penaeoid shrimps of commerce in the Americas, by Isabel Perez

Farfante. April 1988, 32 p.

65. History of whaling in and near North Carolina, by Randall R. Reeves and Edward

Mitchell. March 1988, 28 p.

66. Atlas and zoogeography of common fishes in the Bering Sea and Northeastern

Pacific, by M. James Allen and Gary B. Smith. April 1988, 151 p.
NOAA Technical Report NMFS 79

Species Composition, Distribution,

and Relative Abundance of Fishes

in the Coastal Habitat off the

Southeastern United States

Charles A. Wenner

George R. Sedberry

July 1989

u.s. DEPARTMENT OF COMMERCE

Robert Mosbacher, Secretary

National Oceanic and Atmospheric Administration

William E. Evans, Under Secretary for Oceans and Atmosphere

National Marine Fisheries Service

James Brennan, Assistant Administrator for Fisheries
The National Marine Fisheries Service (NMFS) does not approve, recommend or endorse any proprietary product or proprietary material mentioned

in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales promotion which would

indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as

its purpose an intent to cause directly or indirectly the advertised product to

be used or purchased because of this NMFS publication.

11
CONTENTS

Abstract 1

Introduction 1

Methods 1

Surveys 1

Biomass and abundance 3

Community analysis 4

Results and discussion 5

Species composition 5

Biomass and abundance 6

Species accounts

Leiostomus xanthurus 8

Micropogonias undulatus 10

Brevoortia tyrannus 12

Anchoa mitchilli 14

Stellifer lanceolatus 16

Cynoscion nothus 18

Prionotus carolinus 20

Larimus fasciatus 22

Chloroscombrus chrysurus 24

Scophthalmus aquosus 26

Anchoa hepsetus 28

Menticirrhus americanus 30

Symphurus plagiusa 32

Opisthonema oglinum 34

Pomatomus saltatrix 36

Community analysis 39

Acknowledgments 44

Citations 44

Appendix 1 48

iii

Species Composition,

Distribution, and Relative

Abundance of Fishes in the

Coastal Habitat off the

Southeastern United Statesl

CHARLES A. WENNER

GEORGE R. SEDBERRY

Marine Resources Research Institute

South Carolina Wildlife and Marine Resources Department

P. O. Box 12559

Charleston. SC 29412-2559

ABSTRACT

Ichthyofauna of the coastal «10 m depth) habitat of the South

Atlantic Bight were investigated between Cape Fear, North

Carolina, and the St. John's River, Florida. Trawl collections

from four nonconsecutive seasons in the period July 1980 to

December 1982 indicated that the fish community is dominated

by the family Sciaenidae, particularly juvenile forms. Spot

(Leiostomus xanthurus) and Atlantic croaker (Micropogonias

undulatus) were the two most abundant species and dominated

catches during all seasons. Atlantic menhaden (Brevoortin tyrannus) was also very abundant, but only seasonally (winter and

spring) dominant in the catches. Elasmobranch fIShes, especially

rajiforms and carcharinids, contributed to much of the biomass

of fishes collected. Total fish abundance was greatest in winter

and lowest in summer and was influenced by the seasonality

of Atlantic menhaden and Atlantic croaker in the catches.

Biomass was highest in spring and lowest in summer, and was

influenced by biomass of spot. Fish density ranged from 321

individuals and 12.2 kg per hectare to 746 individuals and 25.2

kg per hectare. Most species ranged widely throughout the bight,

and showed some evidence of seasonal migration. Species

assemblages were dominated by ubiquitous year-round residents

of the coastal waters of the bight. Diversity (H') was highest

in summer, and appeared influenced by the evenness of distribution of individuals among species.

'Contribution no. 270 of the South Carolina Marine Resources Center.

Introduction _

Investigations of the ichthyofauna ofthe South Atlantic Bight

(SAB, herein defined as the area between Cape Fear, North

Carolina [33.9°N] and Cape Canaveral, Florida [28.5°N])

were initiated by the Bureau of Commercial fisheries during exploratory fishing operations off the southeastern U.S.

coast. The results of these efforts were summarized by

Struhsaker (1969) who presented a generalized habitat

scheme based on bottom type and species composition of the

fishes. The coastal habitat (0-18 m [0-10 fm]) has a sandymud bottom and experiences extensive seasonal temperature

and salinity fluctuations relative to other shelf habitats in the

region (Struhsaker 1969, Mathews and Pashuk 1977, 1986).

Sciaenids are the dominant coastal fishes, and penaeids and

portunids are the most abundant decapods (Struhsaker 1969,

Wenner and Wenner 1989).

Descriptions ofthe fish assemblages of the coastal habitat

are derived from studies ofthe bycatch ofthe penaeid shrimp

fishery (Keiser 1976, Anderson and Gehringer 1965, Knowlton 1972). These accounts were concerned with the annual

cycle of species composition and relative abundance of the

bycatch on a state-by-state basis. This approach, although

useful for a general description of the ichthyofauna of the

inshore zone, has several limitations. First, since vessels in

the fishery use different sized nets, doors, tow times, etc.,

there is a lack of standardization of the fishing gear. This

essentially renders invalid the comparisons of catches from

different vessels over a seasonal cycle. It is also difficult to

quantify data (in terms of catch/hour) from several vessels

that tow differing lengths of time and divide the catch by

time towed to obtain catch/hour. This assumes a linear relationship between tow time and catch. Second, shrimp fishermen confine their efforts to discrete areas where penaeid

shrimp are most abundant. Locations near estuaries are

heavily fished, whereas other areas are very lightly trawled.

Thus there was no random sampling, so conclusions drawn

from these data are limited. The purpose of this paper is to

describe the seasonal species composition, distribution, and

relative abundance of coastal fishes in the SAB from regional,

standardized otter trawl surveys.

Methods _

Surveys

A stratified random sampling design (Grosslein 1969) was

used to determine the seasonal distribution and relative abundance of groundfish in coastal waters of the SAB. Thirteen

strata were erected in depths of 4.6 to 9.1 m from Cape Fear,

North Carolina (33.9°N) to the mouth ofthe St. John's River,

Florida (30.4ON). South of this point to Cape Canaveral,

Florida (28.5°N), an additional five strata were delineated

with the 5.6-km territorial sea boundary as the offshore boundary (Fig. 1) This change in stratum definition was caused

by the steepness of the nearshore shelf in this area. Areas

of strata ranged from 7,486 to 31,661 ha (x = 17,323 ha).
WINTER SPRING

Figure 1

Sampling strata (number in box), number of tows per stratum (number in parentheses in box), bottom temperature, and salinity for each

sampling cruise.

2
(K . M . [0.6H])

10,000 m2/ha

a =

stratified mean catch/tow

= mean catch/tow in numbers or weight in the

h stratum

N = total number of possible sampling units over

all strata

Nh = number of possible sampling units in the h

stratum.

- where Yst

Yh

where a = swept area in hectares

K = speed in meters per hour

M = time in hours fished

H = head rope length in meters.

Biomass and abundance

Catch-per-standard-tow was defined as the combined catch

from both trawl nets fished simultaneously at a given stratified random station. The stratified mean catch-per-tow was

calculated by the expression (Poole 1974):

-

Ysr = ---

N

Frequency distributions of both the number and weight of

fish per tow approximated the negative binomial. Data were

subjected to a naturallogarithrnic transformation to standardize the variances and approximate normality. Appropriate

parametric tests of significance were employed when the

underlying assumptions of the model were met. When these

criteria were not met, nonparametric statistical techniques

were used.

Minimum biomass and density estimates for trawl-caught

groundfish were calculated by the area swept by our trawl

gear. The approximation ofthe area swept was derived from

the equation (Roe 1969):

Number Possible N of

Season Date of tows sample blocks:tows

Winter 7-29 Jan 1982 54 58:1

Spring 28 April-6 June 1981 54 58:1

Summer 15 July-20 Sept 1980 108 29:1

Fall 14 Oct-7 Dec 1982 89 35:1

Table 1

Sampling effort and ratio of the number of possible:actual trawl

tows made during each survey in the survey area.

At each randomly selected trawl station within a stratum,

paired four-seam Gulf of Mexico shrimp trawl nets were

towed for 20 min at a speed of 4.4 km/h from either the RIV

Atlantic Sun (summer, spring, and winter surveys) or the

R/V Lady lisa (fall survey). Both vessels were approximately

20 m in length, were formerly involved in the commercial

shrimp fishery, and have outriggers that enabled two nets

to be fished simultaneously. Characteristics of the fishing

gear towed from each outrigger were: 1.5 x 0.8 m wo~en

and chain doors attached with 61-m bridles to the tow wIre;

four-seam (l2.8-m head rope length, 15.8-m foot rope

length) semiballoon trawl net with stretch mesh sizes of 5.1

cm in the wings, 4.4 cm in the body, and 4. 1 cm in the cod

end attached to the doors. A tickler chain was attached

between the doors and adjusted to drag on the bottom 0.6

m in front of the nets. Trawl tows were confined to daylight

hours (l h after sunrise to 1 h before sunset) to eliminate

diel changes in availability and vulnerability of groundfish

to the trawl gear. At the end of each trawl tow, bottom temperature and salinity samples were taken with a Van Dohrn

bottle with an internally mounted stem thermometer. Dates

of the survey and sampling effort are in Table 1.

Contents of each net were processed independently. Fishes

were sorted to species, counted, weighed to the nearest gram,

and either the total catch or a random subsample of very

abundant taxa was measured to the nearest cm total length

(TL) or fork length (FL).

The result (1.12 ha) was multiplied by two since our standard

unit of effort was two nets fished simultaneously. Thus a

standard trawl station sampled 2.24 ha.

3
Community analysis

Species diversity (R') in bits of information per individual

was calculated using the Shannon formula (Pielou 1969). The

species-richness component of diversity was measured using

D = S-l/lnN, where S is the number of species and N is

the number of individuals (Margalef 1968). The evenness

component, i.e. the distribution of individuals among species,

was measured using E = R'/R'max which is the ratio of the

observed diversity (R') of a sample to the maximum R'

the sample could have given its number of species (Pielou

1969). Diversity measures were calculated for pooled trawl

catches in each stratum during a given survey.

Cluster analysis was used to define assemblages of groundfish and to determine the similarity of strata in species composition and relative abundance of species. Prior to cluster

analysis, data from individual trawl tows during a season

were pooled by stratum. In effect, this designates a strata

with number of trawl stations occupied during a survey

as a smgle collection. Pooling was done to reduce the data

matrix to manageable proportions. Because sampling effort

differed among strata, similarity indices for pooled catches

for each stratum were calculated on percent standardized data

(Boesch 1977). Data were edited before cluster analysis to

4

eliminate species that occurred in less than four strata over

the total survey, since these have no discernible distribution

pattern and, therefore, contribute no information to the

analysis (Boesch 1977). The Bray-Curtis coefficient (Clif-

~ord and Stephenson 1975) was used to measure similarity

I~ ~pe~ies composition and relative abundance (percent

slmllanty for standardized data) of collections pooled by

stratu~ a~d cruise. The Bray-Curtis measure is the most appropnate mdex to determine percent similarity (Bloom 1981).

Normal analysis compared similarities among pooled

collections (entities are pooled collections; attributes are the

standardized species abundance scores), and inverse analysis

compared similarities among the distribution patterns of

species (entities are species, attributes are the sites where

they occur) (Clifford and Stephenson 1975, Boesch 1977).

Entities were classified into groups by using flexible sorting

(Lance and Williams 1967), with [3 = -0.25. Nodal analysis

(Boesch 1977) was used to examine the co-occurrence of

species and site groups based on patterns of constancy (frequency of occurrence of members of a particular species

group among the collections of a given site group) and fidelity

(frequency of species of a species group in a particular site

group, relative to other site groups).
Table 2

Top 25 species by numerical abundance for the four seasonal coastal surveys in the South Atlantic Bight, including seasonal

rank and percent of catch.

Total catch Seasonal rank (% catch)

Species Total no. % catch Winter Spring Summer Fall

Leiostomus xanthurus 101,062 29.4 2 (29.9) 1 (55.1) 1 (12.6) 3 (12.0)

Micropogonias undulatus 39,165 11.4 3 (22.8) 2 (8.8) 4 (7.6) 16 (1.4)

Brevoortia tyrannus 36,949 10.7 1 (33.0) 8 (1.5) 45 (0.1) 35 (0.1)

Anchoa mitchilli 25,233 7.3 4 (6.0) 4 (6.8) 8 (5.1) 2 (12.4)

Stellifer lanceolatus 16,154 4.7 5 (2.5) 13 (0.9) 3 (8.6) 4 (8.9)

Cynoscion nothus 14,911 4.3 12 (0.3) 19 (0.5) 10 (2.9) 1 (16.8)

Prionotus carolinus 11,601 3.4 51 «0.1) 3 (7.4) 6 (6.5) 55 «0.1)

Larimus fasciatus 10,262 3.0 16 (0.2) 10 (1.4) 2 (10.2) 11 (2.0)

Anchoa hepsetus 10,233 3.0 14 (0.2) 16 (0.8) 7 (5.2) 6 (7.6)

Menticirrhus americanus 9,346 2.7 11 (0.4) 14 (0.8) 5 (6.9) 7 (4.4)

Symphurus plagiusa 8,780 2.6 9 (0.5) 7 (1.6) 21 (1.1) 5 (8.4)

Chloroscombrus chrysurus 6,341 1.8 72 «0.1) 11 (1.3) 9 (4.9) 10 (2.2)

Scophthalmus aquosus 4,477 1.3 26 «0.1) 5 (2.6) 13 (2.1) 21 (0.6)

Anchoa cubana 4,094 1.2 15 (0.2) 6 (1.8) 14 (1.8) 17 (1.2)

Selene setapinnis 3,910 1.1 72 «0.1) 44 «0.1) 11 (2.7) 9 (2.7)

Anchoa lyolepis 3,727 1.1 44 «0.1) 9 (1.5) 18 (1.3) 13 (1.9)

Trichiurns lepturus 3,580 1.0 25 (0.1) 22 (0.4) 19 (1.3) 8 (3.1)

Cynoscion regalis 3,094 0.9 8 (0.5) 12 (1.0) 23 (0.9) I (16.8)

Etropus crossotus 2,901 0.8 13 (0.3) 27 (0.2) 15 (1.8) 14 (1.5)

Opisthonema oglinum 2,823 0.8 10 (0.4) 21 (0.4) 24 (0.9) 12 (1.9)

Trinectes maculatus 2,147 0.6 46 «0.1) 15 (0.8) 22 (1.0) 20 (0.9)

Centropristis philadelphica 1,857 0.5 30 «0.1) 35 (0.1) 19 (1.4) 19 (1.0)

Arius felis 1,839 0.5 26 (0.2) 12 (2.2) 45 «0.1)

Citharichthys macrops 1,761 0.5 18 (0.1) 25 (0.2) 16 (1.5) 25 (0.5)

Peprilus triacanthus 1,702 0.5 7 (0.6) 17 (0.6) 35 (0.3) 23 (0.5)

Total n 327,949 106,663 91,483 67,701 68,147

Total N 344,116 107,045 93,074 72,844 71,153

Percent nlN 95.3 99.6 98.3 92.9 95.8

Results and discussion _

Species composition

The 305 trawl stations from the four seasonal research survey

cruises in the SAB collected 344,116 fishes in 147 species

(Appendix 1) belonging to 52 families with a total weight

of 12,106.7 kg. The Sciaenidae ranked first by both numbers

and weight and accounted for 56.5 and 66.5% of the total

number and weight, respectively, of fishes taken during all

seasons. Spot (Leiostomus xanthurus) ranked first in total

number and total weight of all species for the four seasons

combined, and was first by weight in all seasons (Tables 2

and 3). Although not included in the top 25 numerically dominant species, rajiform elasmobranchs (Dasyatis centroura,

D. sayi, Rhinoptera bonasus, Raja eglanteria, and Myliobatis

fremenvillei) and the carcharhinid Rhizoprionodon terraenovae were among the 25 most important species by weight

because of their large size relative to the juvenile teleosts

that dominated by number (Table 3). The 25 most numerous

5

species accounted for 95.3 % of the total numbers of fishes

for all surveys, whereas the top 25 species by weight composed 90.6% of the total weight. Seven of the top 25 species

by numbers and weight were sciaenids, a pattern that was

seasonally consistent. Thus, the coastal area of the SAB is

largely a sciaenid habitat.

Some seasonal changes were apparent in the dominant

species. For example, during the winter survey, Atlantic

menhaden (Brevoortia tyrannus) was most numerous, accounting for 33.0% of the total catch (Table 2). The total

number caught during this season was an order ofmagnitude

higher than during the spring, when B. tyrannus was the

eighth most numerous species. In the summer and fall Atlantic menhaden did not rank in the top 25 species. Butterfish

(Peprilus triacanthus), though not as abundant as Atlantic

menhaden, showed a similar seasonal pattern of abundance.

Silver seatrout (c. nothus), striped anchovy (A. hepsetus),

southern kingfish (M. americanus), and weakfish (c. regalis)

were most abundant in the fall, but ranked low in abundance

during other seasons (Table 2).
Biomass and abundance

The stratified mean catch-per-tow for all fishes was highest

by number in the winter and lowest in summer (Table 4),

and this pattern appeared to be greatly influenced by the

seasonality of Brevoortia tyrannus and Micropogonias undulatus (Table 5). Stratified mean catch per tow by weight

was greatest in spring and least in summer. This was due

primarily to large catches of Leiostomus xanthurus, and increased biomass of Cynoscion regalis, Prionotus carolinus,

and Chloroscombrus chrysurus in spring (Table 5). Numerical density estimates ranged from a minimum of 321 to a

maximum of 746 fish/hectare, whereas biomass ranged from

12.2 to 25.2 kg/hectare.

6

Table 4

Seasonal stratified mean catch/tow by numbers and weight, and density estimates based on a swept area of 2.24 hectares during a

standard station.

Stratified mean

catch tow Density

Season n kg n/ha kg/ha

Winter 1,671 51.529 746 23.004

Spring 1,483 56,491 622 25.219

Summer 719 27.355 321 12.212

Fall 793 32.810 354 14.647
Table 5

Stratified mean catch-per-tow and weights (kg) of IS most numerous species caught during the coastal survey.

Winter Spring Summer Fall

Species n weight n weight n weight n weight

Leiostomus xanthurus 547 21.638 629 34.312 92 7.038 91 6.103

Micropogonias undu/atus 422 17.897 ll8 1.946 60 2.287 10 1.434

Brevoortia tyrannus 394 6.7ll 32 1.237 I 0.052 I 0.070

Anchoa mitchilli 103 0.146 III 0.219 26 0.024 89 0.I12

Stellifer /anceo/atus 45 0.398 13 0.280 51 0.650 72 1.046

Cynoscion nothus 8 0.063 9 0.498 22 0.481 143 2.275

Prionotus carolinus * 0.001 150 0.900 42 0.571 * 0.007

Larimus fasciatus 4 0.029 22 0.778 77 2.091 15 0.270

Anchoa hepsetus 5 0.048 14 0.125 44 0.223 67 0.601

Menticirrhus americanus 8 0.607 13 0.889 43 2.251 33 3.038

Symphurus p/agiusa 9 0.235 21 0.700 8 0.226 62 2.012

Ch/oroscombrus chrysurus * ** 24 1.765 35 0.584 15 0.090

Scophtha/mus aquosus I 0.035 38 0.459 15 0.347 6 0.262

Anchoa cubana 4 0.004 29 0.058 14 0.016 9 0.017

Selene setapinnis * ** I 0.031 24 0.Q75 21 0.124

Anchoa /yo/epis * ** 27 0.076 16 0.021 14 0.019

Trichiurus /epturus 7 0.245 7 0.245 7 0.568 30 0.379

Cynoscion regalis II 0.469 14 0.882 4 0.192 10 0.551

Etropus crossotus 5 0.084 4 0.063 13 0.257 12 0.239

Opisthonema oglinum 8 0.051 6 0.145 7 0.I16 19 0.224

Trinectes maculatus * 0.005 15 0.533 7 0.213 7 0.271

Centropristis philade/phica I 0.020 I 0.055 17 0.384 8 0.332

Arius fe/is 0 0.000 3 0.413 14 2.275 * 0.036

Citharichthys macrops 2 0.025 3 0.042 II 0.190 3 0.062

Peprilus triacanthus 9 0.333 8 0.134 2 0.066 4 0.089

Other species 78 2.485 171 9.076 46 5.761 73 13.543

Total 1,671 51.529 1,483 56.491 719 27.355 793 32.810

*<1 individual per tow

**<0.001 kg per tow

7
Leiostomus xanthurus _

Spot ranges from the Rio Grande River to Florida in the Gulf of Mexico and from southern Florida to Massachusetts along

the Atlantic coast of the United States (Fischer 1978). It is one of the most widely occurring and abundant species of fish

in the coastal waters of the SAB (Keiser 1976, Anderson 1968). Spot is an important member of the estuarine fish community

in states throughout the region (North Carolina, Weinstein 1979; South Carolina, Wenner et al. 1982; Georgia, Dahlberg

1972). In continental shelf waters it has been reported from the surf zone (Anderson et al. 1977), coastal waters (Keiser

1976, Struhsaker 1969) to depths as great as 110 m (Wenner et al. 1979a).

We found spot to rank first in both numerical abundance and weight (Tables 2 and 3). This species occurred during all

surveys throughout the SAB regardless ofseason (Fig. 2). There was no significant difference in the frequency of occurrence

of L. xanthurus between seasons at the 95% probability level (X2 = 7.02, df = 3); however, maximum catches occurred

in spring (51,281 individuals) and minimum catches occurred in fall (8,528). Spot were taken at 89% of the 305 stations

made during the four surveys.

The mean catch-per-tow was highest in two strata off Georgia during winter; catches from summer and fall were moderate

throughout the region (Fig. 2). The stratified mean catch/tow both in numbers and weight was highest in spring and lowest

in the fall (Table 5). Length-frequency distributions indicate that two or possibly three age-classes (Young-of-year, I, II)

were present in the SAB (Fig. 3).

8
Spot Leiostomus xanthurus

,,~ ~"?"

0-":) ;:;

OJ ~o

c? .. ;f: .t: . 3/3

.jj .

..C.h.orleS~~(3 0/3

. ... #. ·:r'·

. ' . .. - 3/3

...... ij/ . ~~~."

,·-71 3/3

Savannah

WINTER FALL

X Number ITow

Le/ostomus

XBnthuru$

Do

~1-25

26-100

lIT] 101-1000

.,00,.

Figure 2

Seasonal catch rates in

coastal strata. n/N= no.

tows with spot/no. tows in

stratum.

Leiostomus xanthurus

30 Season

>- 1\

0 I

c I \ e-eWinter (n=32.063) Q) .......,. :J I \ ·--·Spring (n=51.288)

0- 20 I 1 .

. \. • ... • Summer (n=8,732) Q) 1 \! .i·

... L 9-"-9 Fall (n=8,745) I i \. u...

" ... I \ . -+- f\ '.. c I I \ .

! \ . Q) f

\ .

0 .. 10 I t . \ . L f • \ ".

Q) I ! : \.

CL I i. .., \ •·.i ii, I ) 0

0 5 10 15 20 25 30

Figure 3

Total Length (em) Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.

9
Micropogonias undulatus _

Atlantic croaker is an abundant sciaenid that occurs in coastal and estuarine waters of the western Atlantic from Argentina

to Massachusetts, USA (Fischer 1978). In the coastal and estuarine waters of the SAB, M. undulatus is one of the most

abundant and widely occurring species of fishes (Wenner et al. 1982, 1984; Dahlberg 1972, Mig1arese et al. 1982). Atlantic

croaker spawn offshore over an extended period (Middle Atlantic Bight off Chesapeake Bay, fall [Hildebrand and Schroeder

1928]; off North Carolina, September to May [Hildebrand and Cable 1930]; South Carolina, October to January [Bearden

1964]; Gulf of Mexico, September to late March [White and Chittenden 1977]). In continental shelf waters of the SAB,

neuston samples in fall (October-November) and winter (February-March) collected larval M. undulatus (Powles and Stender

1976). Larvae move from offshore waters into the estuarine nursery grounds of the SAB (Weinstein et al. 1980, Shenker

and Dean 1979). Small individuals tend to be found in the upper, lower-salinity reaches of estuaries (Chao and Musick 1977,

McGovern and Wenner, in prep.). As M. undulatus grows, it moves down estuary so that increased size is correlated with

increased salinity (Chao and Musick 1977, Bearden 1964). Atlantic croaker leave the estuaries in the fall (Bearden 1964)

and those that have attained sexual maturity move offshore to spawn.

In addition to being an important seasonal component of the estuarine ichthyofauna, M. undulatus constitutes a significant

part of the coastal fish community of the SAB. The data of Anderson (1968) for catches of Atlantic croaker along the South

Atlantic coast indicated marked seasonal changes in the abundance of Atlantic croakers. Catches were very low off all states

during late winter and spring, and reached a maximum during late summer and early fall. This pattern is consistent with

other accounts from this area (Keiser 1976, Knowlton 1972).

In the present study, M. undulatus ranked second by both numerical abundance and weight (Tables 2,3), and composed

11.38% of the total number and 14.34% of the weight of fishes collected. This species occurred in 69% of trawl samples;

however, significant differences in the frequencies of occurrence were noted among seasons (X2 = 18.37, df = 3, p>O.05).

Subsequent analysis showed that M. undulatus was taken significantly more frequently in the summer and fall than during

the winter and spring (x2 = 19.57, df = 1, p>O.05); however, greatest abundance of individuals (24,400) occurred in winter

and least abundance in fall (982 individuals). During the winter and spring surveys, M. undulatus was taken more often

in trawl tows made south of Savannah, Georgia (winter: south 67%, north 45%; spring: south 60%, north 42%) (Fig. 4).

Catch rates of M. undulatus in individual strata were more variable both in number and weight during winter and spring

than summer and fall. The highest mean catch/tow was found in the winter off southern Georgia (Fig. 4). A single haul

in that stratum caught 23,387 Atlantic croaker weighing 1,000 kg. This represented 59.7% of the total number and 57.6%

of the total weight of M. undulatus taken over all seasons. The stratified mean catch/tow, both in numbers and weight, was

highest during winter and lowest during fall (Table 5).

The mean length of M. undulatus during the winter survey was 17 cm TL (range 2-28) (Fig. 5). According to White and

Chittenden (1977) this probably represented three age-classes. The single small individual (2 cm TL) was a young-of-year

(yay) M. undulatus. Generally, yay Atlantic croakers of this size are found in the estuarine habitat in winter. Age-I fishes

appeared to be the most abundant, and less than 1% of the M. undulatus were age II using White and Chittenden's (1977)

length-at-age data. Spring trawl tows caught a greater number of yay Atlantic croaker; this decreased the average length

to 13 cm TL (range 4-32). Summer M. undulatus averaged 17 cm TL (range 6-29), whereas fall fish had a mean length

of 18 cm TL (range 11-29). The largest Atlantic croaker was 32 cm TL. It appears that in the SAB, M. undulatus has similar

population attributes, that is, small size, short life span, and high rates of mortality (White and Chittenden 1977), as do Atlantic croaker from the Gulf of Mexico.

10
Atlantic croaker Micropogonias undulatus

~.

cope J Fear

'~;:3

(. 0/3

'I' Cope

);f?:.

' Fear IJ

, • '. 3/5

.. 11.

X Number/Tow

FALL Mlcropogonlas

undulatus

Do'-25

26-'00

[illj,o,-,ooo

.'OOH

Figure 4

Seasonal catch rates in

coastal strata. nJN=no. tows

with Atlantic croaker/no. in

stratum.

Figure 5

Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.

25 3530

Season

.-.Winter (n=24,400)

• - -. Spring (n=8,218)

.' , , • Summer (n=5,565)

.----. Fall (n=982)

15 20

Total Length (em)

5 10

Mieropogonias

undulatus

lI

,.J. 0-t--r......................J.+-~_+_<....-""""'__r_~~~~ ........ .......................>-+-...,.......,

o

20

30

>-

U

C

Q)

:::J

0-

Q)

L

u..

-+-

C

Q)

10

Q)

0..

11
Brevoortia tyrannus _

Atlantic menhaden occurs along the Atlantic coast of North America from Jupiter Inlet, Florida to Nova Scotia (Fischer

1978). This species apparently spawns in most waters over the continental shelf throughout its range with timing dependent

on latitude (Nelson et al. 1977). In the SAB, spawning occurs from November to March, whereas in higher latitudes spawning occurs later in the year (May-October off New York and New England; March-April and September-October off the

Middle Atlantic States) (Reintjes and Pacheco 1966). Larvae are recruited to estuaries along the coast from Florida to Maine

and, following juvenile growth in these estuarine systems, they leave this habitat in the autumn with the onset of cooler water

temperatures (Kroger and Guthrie 1973).

Studies of the fishes of the nearshore zone along the southeast coast, as derived from the incidental bycatch of the penaeid

shrimp fishery, indicated that Atlantic menhaden were present throughout the year with greatest catch rates in winter months

off South Carolina (Keiser 1976). However, in Georgia waters, no period of marked high abundance was apparent (Knowlton

1972). This was in contrast to another study where increased catch rates of Brevoortia sp. were observed off Georgia during

January, February, and March (Anderson 1968).

In the present study B. tyrannus was the third most numerous species in trawl collections (Table 2) and ranked fourth

in total weight (Table 3). There were seasonal differences in the relative abundance of Atlantic menhaden. In winter, this

species ranked first by number (35,361 individuals) (Table 2). Spring sampling found it to rank eighth, and in the summer

and fall, this species was not in the top 25 most numerous species. Lowest catches were in the summer (70 fish).

Atlantic menhaden were found throughout the survey area during all seasons (Fig. 6), but differed significantly in seasonal

frequency of occurrence ex? = 42.21, df = 3, p>O.05). This species was taken in 65 % of the winter and fall tows, 43%

ofthe spring tows, and only 17% ofthe summer samples. Over all seasons, 36% of collections yielded B. tyrannus. Catch/tow

values for individual strata showed that the highest catch rates (i>I001 individuals/tow) occurred during the winter months

off South Carolina and Georgia (Fig. 6). Standardized catches were lighter during the other seasons with summer and fall

catches being the least. The stratified mean catch/tow was an order of magnitude higher in winter than in spring, and two

orders of magnitude higher than in summer or fall (Table 5).

Atlantic menhaden caught in winter had a modal length of 10 cm FL (range 6-21); only 1.1 % of these fish were greater

than 12 cm FL (Fig. 7). In spring, the modal length was 13 cm FL (range 7-29) and 1.8% of the B. tyrannus were greater

than 18 cm FL. The modal length increased to 14 cm FL in summer (range 5-21) and 17 cm FL in fall (range 9-30).

Data from these surveys indicate that YOY Atlantic menhaden are abundant during winter in the coastal habitat of the

SAB from Winyah Bay, South Carolina, to northern Florida. It must be pointed out, however, that the gear may not be

the most efficient in sampling this species. During winter, we did not observe any surface schools as seen during warmer

months in more northerly waters, so the Atlantic menhaden may either school more deeply or the schools are not as compact

as during other seasons. The question remains as to the source of large aggregations of Atlantic menhaden off the coast in

winter. Do they represent individuals that have left the estuaries of the SAB the previous fall? Are they derived from more

northerly estuaries and have migrated around Cape Hatteras as has been suggested by Kroger and Guthrie (1973)? Are they

from both areas? Kroger and Guthrie (1973) stated that age-I menhaden (the bulk of Atlantic menhaden captured in winter

will be age-I the following spring when the annulus is formed [Kroger et al. 1974]) move northward along the coast with

the largest fish going farther north. Since these age-I fish are a significant part of the commercial catch in Chesapeake Bay

(l.W. Smith, Beaufort Lab., Southeast Fish. Cent., Natl. Mar. Fish. Serv., NOAA, Beaufort, NC 28516, pers. commun.

Sept. 1984), it may be possible to relate the catch-per-unit-effort of YOY Atlantic menhaden along the South Atlantic coast

to landings of age-I fish in the Chesapeake Bay the following fishing season. Such a relationship will require a long-term,

standardized database, but in view of the magnitude of the fishery and the economic importance of this species, it would

be a worthwhile undertaking.

12
Atlantic menhaden Brevoortia tyrannus

~~\1- ,,- Cope

Canaveral \ .~

~".",

,

Cope

~~ Canaveral

FALL

X Number/Tow

Br8lfoortla

tyrannus

Do

[;] '-25

26-'00

[Jj '0'-'000

• '001<

Cape

~~ Canaveral

~,

Figure 6

Seasonal catch rates in

coastal strata. n/N= no.

tows with Atlantic menhaden/no. tows in stratum.

Brevoortia tyrannus

Fork Length (em)

Figure 7

Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.

30 35

Season

25

.-.Winter (n=35,361)

·--·Spring (n=1,425)

.' ..• Summer (n=70)

.----. Fall (n=93)

15 20

•

10

!

• ..... ,: " "

I ll\,/\

:i..~"'V \

: ! \'. ~" 1 .! \. \, / : i \•. \. / :! '. ~,. : ..." ...... ..,(.•.~. / II-:. \, •.

50

>-

u 40

c:

Q)

::J

(J 30

Q)

L

l...L...

-c: 20

Q)

U

L

Q) 10 0....

0 ,. ,

0 5

13
Anchoa mitchilli _

Bay anchovy occurs along the Atlantic coast of the United States from the Gulf of Maine to Florida and throughout the Gulf

of Mexico to Yucatan, Mexico (Hildebrand 1963b). Within this geographical range, this species is one of the most ubiquitous

and abundant coastal and estuarine species.

In the SAB, A. mitchilli has been reported in estuaries .of North Carolina (Weinstein 1979), South Carolina (Wenner et

al. 1982, 1984; Shenker and Dean 1979), and Georgia (Dahlberg and Odum 1970, Dahlberg 1972). All these studies have

indicated that A. mitchilli is a ubiquitous and abundant species that occurs over a broad range ofsubstrate types, water depths,

temperatures (0.7-32.5°C) and salinities (0.0-36%0). In the coastal waters off South Carolina, Keiser (1976) found A.

mitchilli to occur in all months. Bay anchovy accounted for 6.22 % of the total number of fishes in the bycatch of the South

Carolina penaeid shrimp fishery over an annual cycle (Keiser 1976). Anchoa mitchilli is apparently a coastal and estuarine

species since it was not taken in otter trawl surveys in the open-shelf habitat of the SAB (Wenner et al. 1979a,b,c,d, 1980).

We found A. mitchilli to be widely distributed throughout the survey area during all seasons (Fig. 8). Although there was

no significant difference at the 95 % level in the frequency of occurrence of A. mitchilli in trawl tows among seasons (x2

= 7.436, df = 3), this species was caught more frequently in tows made during spring and summer than during other seasons

in the southern section of the survey area (Fig. 8). Winter catch/tow values were generally high except for the three southernmost strata, where this species was absent. In contrast to the spring when catches were greater in magnitude than winter,

those in summer were light. The stratified mean catch/tow was highest in both numbers and weight during the spring and

lowest during summer (Table 5).

The smallest A. mitchilli (2 cm FL) taken was during the summer survey. Although small bay anchovies may not be fully

represented in the samples because of the high probability that many passed through the meshes of the net, the following

length-frequencies are presented for illustrative purposes:

Length Winter

(em FL) 1982

2

3 10

4 ]171

5 3178

6 1563

7 450

8 15

Spring

1981

66

2497

3373

857

26

Summer Fall

1980 1982

6

750 22

1902 2681

881 4584

134 1458

9 70

1

Modal lengths progressed from a low of 4 cm FL in summer to 6 cm FL in Spring.

14
Bay anchovy Anchoa mitchilli

X Number/Tow

FALL Anchoa

mitchllli

Do

W 1-25

25-'00

[illjl0'-400

.40,.

Figure 8

Seasonal caleb rates in

coastal strata. nIN=no. tows

with bay anchovy/no. tows In

stratum.

15
Stellifer lanceolatus _

Star drum is one of the most abundant coastal demersal fishes in the SAB (Welsh and Breder 1923). The primary range

of this species is from the northern part of the Gulf of Mexico to North Carolina (Fischer 1978) although a specimen was

reported along the Virginia coast (Hildebrand and Schroeder 1928). Star drum are common inhabitants of sandy and muddy

bottoms along the South Atlantic coast from the surf zone (Anderson et al. 1977) to depths around 13 m (7 fm) (Welsh and

Breder 1923). In addition to its importance in the SAB coastal fish community, S. lanceolatus has also been found to comprise

a significant part of the ichthyofauna of South Carolina (Wenner et al. 1981, 1982) and Georgia estuaries (Dahlberg 1972).

Stellifer lanceolatus ranked fifth in abundance (Table 2) but, because of its relatively small size, twelfth by total weight

for all cruises combined (Table 3). Although S. lanceolatus were caught throughout the survey area from Cape Fear, North

Carolina, to Cape Canaveral, Florida, during all seasons (Fig. 9), marked seasonal differences in its frequency of occurrence

in trawl tows were noted. Chi-square analysis showed a significant difference in the frequency of occurrence among seasons

<x2 = 18.81, df = 1, p>O.05). The greatest frequency was found during the fall survey (80% of 89 tows), and the lowest

was during the winter (44% of 54 tows). During the spring and summer surveys, star drum were encountered in 59% of

the tows in each season (spring 54, summer 108).

The greatest mean number per tow of S. lanceolatus (646 individuals) was found in a stratum off southern Georgia during

the winter survey (Fig. 9). The adjacent stratum to the south had a relatively high value, but catches in the remaining strata

were very low or zero. During this period, star drum were absent in 7 of the 18 strata sampled. Spring catch rates for individual strata were moderate and increased during the summer months; the abundance and frequencies of occurrence in

all strata during the fall tended to be higher than any other survey period. Stratified mean catch-per-tow values were 5.5

times greater for number per tow and 2.6 times greater for weight during the fall (Table 5) than the minimum values found

during the spring.

Although the seasonal surveys were not sequential, an "annual" cycle of the length composition and an estimate of the

age structure can be inferred from length-frequency data. Stellifer lanceolatus spawns from late spring through September

in the coastal waters of the SAB (Powles and Stender 1978). During the spring survey, this species had a modal TL of 12

em (range 8-16 em) (Fig. 10). Young-of-year (YOY) star drum were first vulnerable to the gear during the summer; at this

time there were two apparent modes in the length-frequency distribution (8 and 11 em). Small individuals in both fall and

winter surveys suggest continuing recruitment to the gear as would be expected in a species with a protracted (4-5 months)

spawning season. This continual recruitment does not readily allow the breakdown of length-frequency distributions into

age groups. However, the data suggest that two, or at maximum, three age-classes are present in the coastal waters of the SAB.

16
Star drum Stellifer lanceolatus

Cana

Cape ~~" .... ero.1'~

\':""

Cope

Canaveral ."

FALL

X Number ITow

Stelliter

lanceolatu$

Do'-25

26-'00

E]j'O'-400

.400'

Figure 9

Seasonal catch rates in

coastal strata. nlN=110. tows

with star drum/no. tows in

stratum.

Stellifer laneeolatus

0+-

s::

Q)

()

I.-

Q)

0....

30

20

10

Season

e-eWinter (n=2,695)

.--. Spring (n=857)

.' " • Summer (n=6.286)

.-..-. Fall (n=6.316)

/"

'~/.,( .. "\

\;.: . \

\" \

:\ \

'-" . \ "«. 4

'-" \

\\

... \ .\ \ ....

..';.

Figure 10

Seasonal length-frequency

distribution in South Atlan- Jtic Bight coastal waters.

10 15 20

Total Length (em)

5

o+- ...........e:-;~4-~_=___,..~____,,__~~~..,....::::~~ ......__.

o

17
Cynoscion nothus _

Silver seatrout occurs along the northern and eastern Gulf of Mexico and the Atlantic coast of the United States from southern

Florida to Chesapeake Bay (Fischer 1978). This species is rarely taken north of Cape Hatteras (Hildebrand and Schroeder

1928), and in contrast to its congeners along the Atlantic coast (c. regalis and C. nebulosus), C. nothus is restricted in its

estuarine distribution to the higher salinity areas (Schwartz et al. 1982, Dahlberg 1972). Our present knowledge ofthe distribution

and abundance of C. nothus in the SAB results from studies of the bycatch of the penaeid shrimp fishery. Some of this information is suspect, however, because of misidentifications of C. nothus and subsequent lumping with C. regalis. For example,

Knowlton (1972) failed to list C. nothus in his study of the bycatch from coastal waters of Georgia, and Keiser (1976) reported

it as being absent during May through August 1974 in a similar study from South Carolina. Both of these studies are at

variance with other published information and the present study. Anderson (1968) found C. nothus to occur throughout the

SAB in coastal waters during all seasons. Catch rates were an order ofmagnitude higher off Florida, where C. nothus accounted

for 9.6% of the annual fish catch in coastal waters, than off South Carolina and Georgia.

In the present study, C. nothus ranked sixth in numerical abundance and eighth by weight (Tables 2, 3). Silver seatrout

were taken throughout the SAB during each ofthe seasonal cruises (Fig. 11); however, differences in the frequency of occurrence in trawl tows were observed. Cynoscion nothus was taken at significantly more stations (x2 = 11.12, df = 1, p>o.05)

in fall (74% of 89 tows) than during winter (46% of 54 tows), spring (50% of 54 tows), and summer (58% of 108 tows).

Catch-per-tow values in winter were low in all but two strata off northern Florida (Fig. 11). During spring, values were

also low and then increased in summer. The frequency of occurrence in the strata north of Savannah, Georgia, was relatively

low during all seasons except fall when both the catch rates and frequency of occurrence reached a maximum. The stratified

mean catch-per-tow was highest in the fall and lowest in the winter (Table 5).

Cynoscion nothus taken during the winter survey averaged 9 cm TL (range 6-20); the length-frequency distribution during

that season was essentially unimodal (9 cm) (Fig. 12). Cynoscion nothus had a larger mean length (16 cm, range 4-29) during

spring when the frequency distribution had two distinct modes, 13 and 22 cm. Summer length-frequencies also showed two

apparent length groups. Because of the large number of small fishes, the average length decreased to 11 cm (range 4-29).

Fall samples lacked larger sized individuals. The largest silver seatrout we collected (one each in spring and summer) were

29 cm TL (24 cm SL) which is slightly larger than the maximum size reported by DeVries and Chittenden (1982) from the

Gulf of Mexico but smaller than the maximum size found in Georgia waters (31 cm TL) (Mahood 1974).

Although C. nothus has an extended spawning season in the Gulf of Mexico (May through October), two distinct periods

of peak reproductive activity were found in May and August-September (DeVries and Chittenden 1982). Mahood (1974)

indicated that C. nothus spawns during the spring and late summer-early fall off the Georgia coast. Evidence for a spring

spawn was based on the occurrence ofsmall C. nothus in samples taken at that time, whereas late summer-early fall spawning

was determined from gonadal examination and the presence of small individuals in trawl collections from October through

January (Mahood 1974). Our data do not allow the definition ofspecific spawning times throughout the SAB since we collected

relatively small fishes during all surveys.

Available information indicates that large C. nothus disappear during the winter in the Gulf of Mexico off Texas (DeVries

and Chittenden 1982) and along the Georgia coast (Mahood 1974). Our data agree in that fall and winter samples each had

only one C. nothus greater than 22 cm TL. Three possible explanations for this phenomenon are: (1) During fall and winter,

large C. nothus are unavailable to the survey gear because of movement from the limits of the survey area, either south

of Cape Canaveral, Florida or offshore into deeper water; (2) large C. nothus are not vulnerable to the survey gear; (3)

large C. nothus die in fall and winter. Present information does not allow us to rule out the first possibility. However, seasonal

stratified random sampling in offshore waters has collected few C. nothus and these were off Florida in depths from 18 to

27 m (Wenner et al. 1979a,b,c,d, 1980). Since large C. nothus were taken in the survey gear during spring and summer,

trawl avoidance is unlikely. Appearance of some large C. nothus in the early spring indicates that not all large fish die off

in winter, although DeVries and Chittenden (1982) found that C. nothus in the Gulf of Mexico has an annual mortality rate

approaching 100%. It cannot be ascertained that there was a combination of factors, that is, movement offshore or mortality,

and both factors probably contribute to the rarity of C. nothus in the winter.

18
Silver seatrout _ Cynoscion nothus

X Number/Tow

FAll Cynosclon

no thus

Do

G 1-25

26-100

E],0,-400

.401+

Figure 11

Seasonal catch rates in

coastal strata. nlN=00. tows

with silver seatrout/oo. tows

in stratum.

>-

()

C

Q)

:::l

0-

Q)

L

u....

+-

C

Q)

()

L

Q)

0....

30

20

Season

a-aWinter (n=361)

"--"Spring (n=453)

.' . , • Summer (n=2,112)

9----9 Fall (n=11,985)

Total Length (em)

19

Figure 12

Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.
Prionotus carolinus _

Northern sea robin has been reported from the Bay of Fundy to South Carolina (Fritzsche 1978). It is generally found in

coastal and shelf waters, although it is taken, sometimes in large numbers, in estuaries such as the Chesapeake Bay (Hildebrand

and Schroeder 1928). In the present study, P. carolinus was the seventh most numerous species accounting for 3.37% of

the catch (Table 2); it ranked seventeenth by weight (Table 3). Although P. carolinus was taken during each of the cruises

(Fig. 13), there was a significant difference in the frequency of occurrence between seasons (:x2 = 103.36, df = 3, p>O.05).

Northern sea robin was infrequent in winter (7% of tows) and fall (9%) of the trawl samples, whereas in spring and summer

it occurred at 70 and 63 % of the stations, respectively.

The mean catch/tow was very low in the strata where P. carolinus occurred during winter (Fig. 13). The highest catch

rates were found during the spring in strata from Savannah, Georgia, north to Cape Romain, South Carolina. Catches declined

somewhat in summer, and in fall this species was only encountered in the most northerly strata in small numbers. Lengthfrequency distributions (Fig. 14) indicated that there are possibly two age-classes (YOY and age-I) in the coastal habitat.

20
Northern sea robin Prionotus carolinus

Figure 13

Seasonal catch rates in

coastal strata. nIN= no. tows

with northern sea robin/no.

tows in stratum.

Prionotus

c8rolinus

Do

'-25

8BEl 26-100

EiJ]'O'-400

.400_

FALL

,:

cope)~~ Fear '-~

-lJ ' .....

'~

cap~ conav~r~~ r

'\

,-

cope~ ",~~

//~

/~/3

"

~~ Cope

Canaveral, \ ' ,~

,~ '.

Prionotus earolinus

Figure 14

Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.

20 25

Season

e-eWinter (n=5)

.--. Spring (n=6,858)

•...• Summer (n=4,714)

~----~ Fall (n=24)

15

Total Length (em)

r

I \

I \

I \

/ •

I

,

I

I

t

I

I

I

I

I :....--

//.. 5 10

O+--~~_--.~Il-'--"~>--,_____,-~~-__+-~~---__._ ....~~~_,

o

40

10

20

30

-+-

C

Q)

()

L

Q)

D...

()

C

Q)

::J

0-

Q)

L

u...

21
Larimus fasciatus - _

Banded drum is a common coastal sciaenid that occurs on sandy-muddy bottom along the South Atlantic coast (Welsh and

Breder 1923). Although banded drum are occasionally caught north of Cape Hatteras (Hildebrand and Schroeder 1928), the

primary range ofthe species is from North Carolina south along the Atlantic coast to south Florida and throughout the northern

Gulf of Mexico (Fischer 1978). Larimusfasciatus has been taken in salinities ranging from fresh to oceanic waters (Schwartz

et al. 1982); however, when encountered in estuaries along the South Atlantic coast, it occurred most frequently in the polyhaline

and euhaline reaches of estuaries (Wenner et al. 1981, Shealy et al. 1974). In the coastal waters of the SAB, banded drum

have been taken from the surf zone (Anderson et al. 1977) to 19 m (Wilk and Silverman 1976). In this region, however,

the maximum abundance and frequency of occurrence were in depths less than 15 m as shown by data from the bycatch

of the penaeid shrimp fishery. It comprised 1.9% of the discarded weight in South Carolina (Keiser 1976), 3.2% of the

weight in Georgia (Knowlton 1972), and 1.6% of the number of fishes in Florida's bycatch (Anderson 1968).

Plankton tows in the SAB during April-May and August-September collected smal1larvae ofL. fasciatus (Powles and Stender

1978). Presently it is unknown whether this represents two spawning events (spring, fall) as is the case for banded drum

in the Gulf of Mexico (Standard and Chittenden 1984) or is part of an extended spawning season from May to October

(Hildebrand and Cable 1934) along the South Atlantic coast.

Larimus fasciatus ranked eighth in numerical abundance and seventh by weight for fishes taken in the coastal habitat of

the SAB during the four seasonal cruises (Tables 2, 3). This species occurred throughout the survey area during all sampling

periods (Fig. IS); however, the frequency of occurrence in trawl tows was significantly different between seasons (X2 =

18.89, df = 3, p>O.05). Seasonal comparisons showed that L. fasciatus occurred at significantly fewer stations in winter

than in spring (x2 = 5.44, df = 1, p>O.05), summer (x2 = 11.33, df = 1, p>O.05) and fall (X2 = 17.22, df = 1, p>O.05).

Frequencies from seasons other than winter were not significantly different from one another (x2 = 1.57, df = 2). During

those seasons, L. fasciatus was taken in 74.5% of the 251 collections.

Data indicated that banded drum were displaced to the southern part ofthe survey area by the lower winter water temperatures

in the higher latitudes. Their frequency of occurrence in trawl tows south of Savannah, Georgia (approx. 32°N) was significantly

greater than in the northern region (x2 = 15.21, df = 1, p>O.05). Larimus fasciatus became more widely distributed in

the SAB as temperatures increased in the spring (Fig. 15). Maximum catch rates of banded drum were always along the

coast of Florida regardless of the season. The stratified mean catch/tow, both in numbers and weight, was lowest in the

winter and highest during the summer survey (Table 5).

Although the seasonal surveys were not sequential, it is possible to trace modes in the length-frequency distribution through

an "annual" cycle to estimate the age and longevity of L. fasciatus in the SAB. Because of the limited sample size in winter

collections, only a single mode of 7 cm TL can be inferred from the data (Fig. 16). This can be traced through the other

seasons (spring, 9 cm TL; summer, 10 cm TL; fall, 13 cm TL). Interpretation of this is that the 7-cm winter peak represents

the modal length of young-of-year banded drum that were spawned during the previous spring and summer. During the period

between winter and spring, they increased 2 cm TL in modal length and we estimate that they are approximately age-I by

summer. The second summer mode (17 cm TL) probably represents age-II fish; the possibility of older age-classes cannot

be clearly inferred from the data. It appears that possibly three age-classes of L. fasciatus are in the SAB. These estimates

of the length-at-age are in close agreement with von Bertalanffy predicted length-at-age for this species in the northwestern

Gulf of Mexico (I = 13 cm TL; II = 18 cm TL) (Standard and Chittenden 1984). Ross (1978, cited in Standard and Chittenden 1984) obtained the following length-at-age values for L. fasciatus collected off North Carolina: I = 153; II = 188;

ill = 209 mm TL; these are comparable to our estimates.

22
Banded drum Larimus fasciotus

SPRING

Figure 15

Seasonal catch rates in

coastal strata. nlN=no. tows

with banded drum/no. tows

in stratum.

Larlmus

fasc/atus

Do

~1-25

26-100

[ill} 101-400

.401+

X Number/Tow

FALL

,

Cope

Fear

) 1/3

0/3

",,~ ji , 0/3

Savann~h

WINTER

\'

cope~ .:- Fear

..... . . . "," 1·'3

]

' 0/3

,,-<- ;i:-?

&;;

"5 ~o

G? 4. t:' 0/3

ChOriestonfL 0/3

~>

';J;!.' 1/3 . >

...5 <I

. '" 2/3

..... .,,,~ji •

,·n 0/3

Savonn~h

Larimus faseiatus

40

Total Length (em)

Season

.--Winter (n=184)

• - -. Spring (n= 1,266)

.- - - • Summer (n=7,410)

• _ ..-. Fall (n= 1,402)

Figure 16

Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.

5 10 15 20 25

10

20

30

>-

U

C

Q)

:::J

0-

Q)

u....

0+-

C

Q)

U

Q)

a..

23
Chloroscombrus chrysurus _

Atlantic bumper is a schooling species found along the Atlantic coast of North and South America from Massachusetts to

Uruguay, in Bermuda, and possibly throughout the West Indies (Fischer 1978). In the north central Gulf of Mexico, C. chrysurus

is most abundant in depths from 2 to 18 m, and catch rates decline with increasing depth (Klima 1971). Atlantic bumper

are generally rare beyond the 91-m (50-fm) curve, although it was taken to the 1830-m (IOOO-fm) curve (Klima 1971).

Chloroscombrus chrysurus has been found in both the high-salinity portions of gradient estuaries (Wenner et al. 1982, Anderson 1968, Dahlberg 1972) and in euhaline estuaries (Cain and Dean 1976), but greatest abundances have been reported from

coastal waters of the SAB, primarily as a bycatch species of the penaeid shrimp fishery (Keiser 1976). Juvenile Atlantic

bumper (20-54 mm SL) have been taken in the surf zone off South Carolina during summer (Anderson et al. 1977) to offshore

waters of the SAB (Wenner et al. 1980). Small, pelagic juveniles (18-33 mm SL) have been found commensally with sea

nettles (Chrysoura quinquecirrha) and Stomolophus meleagris during July, August, and October in the northern Gulf of Mexico (Phillips et al. 1969). Anderson(l968) found this species to be abundant in the SAB during exploratory surveys of the

coastal habitat. Highest catch rates of Atlantic bumper were during the summer months off South Carolina, late summer-early

falloff Georgia, and fall-early winter off Florida (Anderson 1968).

During the present investigation, Atlantic bumper ranked twelfth in numerical abundance and thirteenth by weight (Tables

2, 3). This species occurred at 60% of stations made during all seasons; however, there was a marked difference in occurrences. During the winter survey, only a single individual was taken, and it occurred at the southernmost station (Fig. 17).

The spring, summer, and fall surveys found C. chrysurus to be widespread throughout the SAB. No significant difference

was found in the frequency of occurrence among these three seasons (x2 = 2.382, df = 2), when Atlantic bumper were

taken at 73 % of 251 stations.

During the spring and summer, maximum catch rates were in the strata off Georgia and Florida (Fig. 17). Atlantic bumper

were widespread throughout the SAB in fall with the highest mean catch/tow in the two most southern strata. The stratified

mean catch/tow by number was highest during the summer survey and lowest during fall (excluding the extremely low value

for winter) (Table 5). The spring survey had the highest value for biomass/tow.

Examination oflength-frequency distributions of the three surveys that caught fish showed that spring sizes were unimodal

with a mean length of 17 cm FL (range 13-26) (Fig. 18). Summer trawl tows had two modal length groups (6 and 17 cm

FL) whereas fall fish had a single modal length (7 cm FL). Atlantic bumper probably spawn in spring and summer along

the southeastern coast of the United States (Fischer 1978), and fish in the smaller modal length group are probably YOY

Atlantic bumper. The larger mode probably represents age I+ fish. This interpretation of the ages is tentative, however,

until age determination using the analysis of hard parts is conducted.

24
Atlantic bumper Chloroscombrus chrysurus

~' Cape

, ,~~b

'f~3 /'

(/

"."''''

·\L("{

Cope

Canoveral \ ,~

'~~

Cope '\t~ Canaveral

FALL

Cape'\i' Fear

..-=1)

X Number/Tow

Chloroscombru$

chrysurus

Do

0'-2526-50

El] 5'-'00

• '0"

Figure 17

Seasonal catch rates in

coastal strata. nIN= no. tows

with Atlantic bumper/no.

tows in stratum.

Season

Figure 18

Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.

20 25 30

• - -. Spring (n= 1.238)

.' , , • Summer (n=3.564)

.-..-. Fall (n=1.538)

10 15

Fark Length (em)

5

ft

! / \

t, I \

i \ I \

i \ / \ i~, { \ i: '..:~ I \

!: \ I

t '\ I \

(\ (\

;: i f \

i ' I \

:.; ;,\ I

·t ' " •'. :/ '.\ I~' '.\

.) '. \ 1,' ..f' ., ::-a.<..'-:. .. ....

>-

30

()

c

a>

::J

0-

a> 20 Ll..L.

C

a>

()

L- 10 a>

CL

0

0

25
Scophthalmus aquosus _

Windowpane have been reported from the Atlantic coast of North America from the Gulf of St. Lawrence to Florida (Gutherz

1967). In the SAB, a few S. aquosus were taken occasionally in otter trawl tows in the open-shelf habitat in depths of 9-22

m (Wenner et al. 1979a,b,c,d, 1980). However, most accounts indicated that S. aquosus were shallow-water residents along

the South Atlantic coast. Keiser (1976) found small numbers in all months but November and December in the bycatch of

the penaeid shrimp fishery off South Carolina. Exploratory trawling along the Georgia coast caught S. aquosus during every

month with highest catch rates of 13 and 26 fish per hour in early spring (Anderson 1968). Although reports have indicated

that S. aquosus was primarily a coastal bothid, it was collected in the high-salinity portions of estuaries in North Carolina

(Weinstein 1979), South Carolina (Wenner et al. 1982), and Georgia (Dahlberg 1972). A single report indicated that S. aquosus

ranged from freshwater to oceanic salinity (Schwartz et al. 1982); however, this species has been found almost exclusively

at high salinities.

Scophthalmus aquosus was more abundant during the present study then was found in the literature. This species was the

thirteenth most numerous fish for all four surveys, but because of its small size, it was less important by weight (Tables

2,3). Scophthalmus aquosus was taken in 57% of the 305 trawl samples and there was a significant difference in the frequency of occurrence between seasons (x2 = 30.308, df = 3, p>O.05). It had the following frequencies of occurrence: winter

26% (N = 54); spring 80 % (N = 54); summer 62 % (N = 108); fall 56 % (N = 89). When the survey area was divided

into northern and southern sections at Savannah, Georgia, and the presence-absence data were pooled for each area over

the four seasons, there was a significant difference in the frequency of occurrence between areas (x2 = 46.001, df = 1,

p>o.05). Scophthalmus aquosus was taken significantly more frequently in the northern (78% of 139 tows) than in the southern

area (39% of 166 tows) of the SAB.

Mean catch/tow values for individual strata were very low during the winter survey (Fig. 19). Spring sampling showed

reasonably consistent and large values in the strata north of Savannah, Georgia. This was followed by a general decline in

mean values for strata in summer and fall. The stratified mean number/tow ranged from a high of 38 individuals in spring

to a low of 1 in winter (Table 5). These two seasons also had the highest and lowest kg/tow values (Table 5).

Length-frequency distributions showed two apparent modes in winter (5 and 15 cm TL) (Fig. 20). During the following

seasons, the smaller mode progressed to 9 cm TL in spring, 11 cm TL in summer, and 15 cm TL in fall. The large mode

was not discernible in other seasons; however, fishes from 16 to 20 cm TL were present in spring and a few individuals

greare' than 20 cm TL were found in fall. Comparisons of these length-frequency distributions of S. aquosus with length-age

inforr'ation from southern New England indicated this species attains a much larger size in the northern part of its range.

Age determinations by otoliths in fishes from southern New England (Moore 1947) gave the following 1ength-at-age (cm

TL) values: I, 6.3; II, 11.7; III, 19.4; IV, 24.7; V, 27.7; VI, 29.6; VII, 31.5. In southern New England, male S. aquosus

matured at age-III whereas some females were mature at age-III and all were mature at age-IV (Moore 1947). Data from

the present study indicated that S. aquosus was not as long-lived and reached a smaller maximum size in the SAB than individuals

north of Cape Hatteras.

26
Windowpane _ Scophthalamus aquosus

,:

cope J Fear

'~

( 0/3

FALL

X Number/Tow

Scophthslmus

aquosus

Do

[;J'-25

26-50

EJ 5'-15

.16'

Figure 19

Seasonal catch rates in

coastal strata. nlN=00. tows

with windowpane/no. tows

in stratum.

Seophthalmus aquosus

Toto I Length (em)

Figure 20

Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.

20 25

Season

.-.Winter (n=49)

·--·Spring (n=2.451)

•...• Summer (n= 1.518)

.----. Fall (n=459)

.. 5 10 15

To i\.

i \. -"\ . : f 'yo I'

f It i \

f ~\ f \

f \ i

f \ i

f \ i

f ,~i

I \ i

I \ !

I • V'

I ./ O+--~~.---.~"'--,- .......__q=o_ _<l_=o____<._........."'"* ='t___lJ=~

o

30

20

10

>-

(J

C

Q)

::J

{J

Q)

L

0+-

C

Q)

(J

L

Q)

a....

27
Anchoa hepsetus _

Striped anchovy occurs along the Atlantic coast from Nova Scotia to Florida, throughout the Gulf of Mexico, and along

the South American coast to Uruguay (Hildebrand 1963b, Whitehead et al. 1988). In Chesapeake Bay, Hildebrand and Schroeder

(1928) found A. hepsetus to be abundant during warmer months in the southern part of the bay. Striped anchovies were yearround residents in estuaries and coastal waters of North Carolina (Schwartz et al. 1982), and have been taken in estuaries

of Georgia during warmer months of May to November (15.7-30.8°C) (Dahlberg 1972). In the coastal waters of the SAB,

A. hepsetus was the ninth most numerous species offishes, comprising 2.65%, in the incidental bycatch ofthe penaeid shrimp

fishery off South Carolina (Keiser 1976).

In the present study, we found A. hepsetus to be the ninth most numerous species for all four surveys combined (Table

2). Because of the small size of the striped anchovy, it ranked only 23rd by weight (Table 3). During the spring, summer,

and fall surveys, A. hepsetus was found throughout the SAB, but in winter this species was taken only in trawl tows off

the Florida coast (Fig. 21). Anchoa hepsetus was in 65% of the 305 trawl samples; however, there was a significant difference in its frequency of occurrence between seasons (x2 = 47.94, df = 3, p>O.05). Striped anchovies were in only 24%

of the winter tows, whereas in spring, summer, and fall they occurred at 74, 72, and 75 % of the stations, respectively.

In winter, the mean catch/tow of A. hepsetus was low for all strata except for one off central Florida (Fig. 21). Spring

samples showed that striped anchovies had spread throughout the Bight with catches being generally low, except off northern

and central South Carolina and southern Georgia. Summer and fall values were highest in the southern part of the survey

area. The stratified mean catch/tow was highest in the fall for both numbers and weight and lowest in winter (Table 5).

The length composition of A. hepsetus from the surveys was:

Fork length

(em)

3

4

5

6

7

8

9

10

11

12

13

14

15

Winter

3

60

132

38

13

Spring

47

50

8

8

196

306

106

28

Summer Fall

21 29

97 557

255 324

627 13

980 1

795 83

539 1332

382 2071

112 739

10 262

1 31

From these data, it appears that A. hepsetus may live 1+ years.

28
Striped anchovy _ Anchoa hepsetus

it Number/Tow

FALL Anchoa

haps.tus

Do'-25

I8SllI 26-'00

EJ '0'-200

.20'+

Figure 21

Seasonal catch rates in

coastal strata. nlN= DO. tows

with striped anchovy/no.

tows in stratum.

29
Menticirrhus americanus _

Southern kingfish ranges from New York to Texas and is the most abundant of three species of the genus occurring in nearshore waters of the SAB from Cape Fear, North Carolina, to Cape Canaveral, Florida (Hildebrand and Cable 1934, Bearden

1963, Dahlberg 1972). Menticirrhus americanus is a demersal species that occurs over a wide variety of substrates ranging

from mud to sandy-mud mixtures. Adults prefer sand bottoms of ocean beaches and the mouths oflarge coastal sounds (Bearden

1963). Southern kingfish are purported to make seasonal movements from the estuarine and nearshore oceanic waters «9

m) to deeper offshore waters when water temperatures dropped below 10°C (Bearden 1963), yet otter trawl surveys have

failed to document this event (Wenner et al. 1979b). A protracted spawning season extends from April to September, and

juvenile growth rates are rapid (Hildebrand and Cable 1934, Bearden 1963).

Menticirrhus americanus occurred from Cape Fear, North Carolina, to Cape Canaveral, Florida, during each ofthe seasonal

surveys (Fig. 22). However, a highly significant (P<0.01) seasonal difference in the frequency of occurrence in trawl tows

was found (x2 = 35.43, df = 3). Frequency of occurrence was highest during summer and fall when M. americanus was

found in 93 of 108 and 87 of 89 trawl collections, respectively. In winter, southern kingfish were encountered in 32 of 54

tows, and the spring survey found this species in 41 of 54 tows. Southern kingfish ranked tenth in numerical abundance

overall and third by weight for the survey (Tables 2, 3).

Winter abundance, as indicated by the mean catch/tow was greatest toward the south (Fig. 22). In winter, the transformed

values (1n [number + 1]) of the mean catch/tow were significantly correlated with the mean bottom temperatures of the strata

(r = 0.611, p<0.05, df = 16). During spring, southern kingfish spread throughout the Bight and reached their greatest

frequency of occurrence and abundance in fall trawl tows.

The stratified mean catch/tow by number was lowest in the winter and highest in the summer (Table 5) when its value

was five times greater than the winter minimum. Larger, heavier M. americanus were captured in fall, thus making this

season first by weight.

Menticirrhus americanus moves from high-salinity estuaries and coastal waters of the northern parts of the SAB to the

central Atlantic coast of Florida during winter. A four-year average of commercial landings for "king whiting" (mostly

M. americanus) in the Cape Canaveral area showed that 88% ofthe 112,084 kg landed on average were made between November

and February (Anderson and Gehringer 1965). Maximum landings occurred at the height of the shrimp season and coincided

with increased effort. Fisheries independent data from 1933-1935 showed lowest catches in the spring (2-3 fish/hour), moderate

summer (40-274 fish/hour) and winter catches (50-188 fish/hour), and maximum catches in the fall (44-680 fish/hour) off

central Florida (Anderson and Gehringer 1965, see Table 9). The recent increase in Florida landings probably resulted from

increased winter effort in the penaeid shrimp fishery along the Florida coast coupled with the "concentrating" effect ofseasonal

hydrographic conditions.

Although M.americanus was a year-round, shallow-water resident of the SAB, seasonal changes in abundance were readily

apparent. Bearden (1963) obtained the lowest catch/tow in southern South Carolina estuaries and coastal stations from January

through April; maximum values were from August through November. We observed the same seasonal changes in abundance

with few M. americanus captured north of Savannah, Georgia, during winter. Since abundance was correlated with temperature,

it appears that most fishes move into warmer waters with the cooling of coastal water in winter. It was assumed that during

this period of cold water temperatures, M. americanus moves offshore to deeper, warmer water (Bearden 1963). This does

not appear to be the case since this species was encountered in only 14% of the stratified random trawl tows (N = 35) in

deeper water during winter months (Wenner et al. 1979b). We also failed to observe any seasonal trend in the frequency

of occurrence, and all captures were in depths from 9 to 20 m, mostly along the coast.

Seasonal length-frequency distributions indicated that two age-classes, in addition to young-of-the-year fish, were present

in the catches, based on age data from Smith and Wenner (1985) (Fig. 23).

30
Southern kingfish Menticirrhus americanus

20

Menticirrhus americanus

20

FALL

X Number/Tow

Mentlcirrhus

amerlcanus

Do1-25

26-50

EJ 51-100

.,01+

Figure 22

Seasonal catch rates in

coastal strata. nlN=00. tows

with southern kingfish/no.

tows in stratum.

>-

U

15

:J

0-

l.L 10

+-'

E

QJ

Q) Summllra... Spawned yay

Winter (n~392)

SpringSpawned YCff

10 '5 20 25 30 35 40 45

15

Summer (n=4,998)

20

>-

U

c

QJ 15

:J

0-

QJ

10

+-'

C

QJ

U

"-

QJ

0..

0

0 '0

Ago 1

Spring (n=800)

15 20 25 30 35 40 45

Total Length (em)

20

15

10 Fall (n=3.1 69)

Ago 1

15 20 25 30 35 40 45

Total Length (em)

31

Figure 23

Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.
Symphurus plagiusa _

Blackcheek tonguefish is found along the Atlantic and Gulf coasts of the United States, the Bahamas, and in the Greater

Antilles (Fischer 1978). It is the most common species of Symphurus on the Atlantic and Gulf coasts of the United States

in inshore waters (Ginsburg 1951). Symphurus plagiusa is the most euryhaline of the Atlantic cynoglossids, being found

from freshwater in estuaries (Schwartz et al. 1982) to hypersaline waters in Florida (Roessler 1970).

Along the southeastern coast ofthe United States, S. plagiusa has been reported from the estuaries of North Carolina (Tagatz

and Dudley 1961, Schwartz et al. 1982), South Carolina (Wenner et al. 1982) and Georgia (Dahlberg 1972). In nearshore

waters off South Carolina, S. plagiusa was most abundant December-April and composed 2.05 % ofthe total number offishes

in the bycatch of the penaeid shrimp fishery (Keiser 1976). In the Gulf of Mexico and Middle Atlantic Bight, this species

spawns during spring and summer (Topp and Hoff 1972, Olney and Grant 1976).

In the present study, S. plagiusa was the eleventh most numerous species and ranked fifth by weight (Tables 2, 3). This

species was found at 78% of stations from all cruises, and there was a significant difference in the frequency of occurrence

of S. plagiusa between seasons (x2 = 41.998, df = 3, p>O.05). It occurred less frequently during winter (48% of all tows)

than in spring (85%), summer (76%), or fall (94%). Seasonal samples found S. plagiusa to be widespread throughout the

survey area (Fig. 24). Catch-per-tow values by stratum were generally low in winter, with maximum catches off southern

South Carolina and central Georgia (Fig. 24). During spring, mean values were lowest in southern strata and highest off

the northern Georgia coast. Summer catches were relatively consistent throughout the SAB, and catches in fall were highest

in the more northern strata.

The length composition of the trawl-caught S. plagiusa was consistent from season to season, with means of 14-15 cm

TL (Fig. 25). The smallest individual was 5 cm TL from the summer, whereas the largest specimens were 21 cm TL from

the fall. The fact that fish less than 10 cm TL were rare in the samples indicated that either .S. plagiusa inhabit a different

habitat as juveniles or they are not recruited to the fishing gear until 10 cm TL.

32
Blackcheek tonguefish _ Symphurus plagiusa

Symphurus

plaglusa

Do

8'-25

26-50

Will 51-100

.,01+

X Number/Tow

FALL

'\ Cope

Feat

'\'

cope~ '~.. Feor , ..

: . 2/3

(1 0/3

1/3

, '. , Figure 24 ~, .~ '''', \t'" .~

\t~ Seasonal catch rates in

Cape copb '"'i\\ Cope coastal strata. nIN=00, tows Canaveral \ Canaveral Canoveral Canaveral with blackcheek tonguerlShl \\~\

"'"

~, no. tows in stratum.

Symphurus plagiusa

>-

0 30 30

c:

Q) Wlntor (n=566) :J

tT

Q) 20 20 '-

u-

-c:

CD ,0 10 0

'-

Q)

ll.

0 0

0 5 10 15 20 25 0 10 15 20 25

>-

0 30

c:

Q)

:J

tT

Q) 20 '-

uCQ) ,0 0

'-

Q)

ll.

0

0 10 '5 20

Total' Length (em)

25

Total Length (em)

Figure 25

Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.

33
Opisthonema oglinum _

Atlantic thread herring is a pelagic clupeid that occurs primarily in coastal waters from the Gulf of Maine to Brazil (Fischer

1978). Occasionally this species will form dense nearsurface schools that are harvested by the same purse seine nets as those

used in the menhaden fishery (Fuss et al. 1969). In the eastern Gulf of Mexico, gill net catches of O. oglinum were highest

between 27 and 29°C in coastal waters less then 18.3 m deep (Kinnear and Fuss 1971). Along the South Atlantic coast,

thread herring was reported as a ubiquitous, transient species that occurred over a broad range oftemperature (0.7-31.0°C)

and salinity (0.5-360100) in the Cape Fear River estuary of southern North Carolina (Schwartz et al. 1982). In the coastal

waters of South Carolina, O. oglinum was collected in the surf zone (Anderson et al. 1977) to locations as far as 20 Ian

offshore (Wenner et al. 1979d). These latter few specimens were taken incidentally to demersal species in bottom trawl samples.

Opisthonema oglinum was found during all seasons in the bycatch of fishes of the South Carolina penaeid shrimp fishery

(Keiser 1976), and Dahlberg (1972) occasionally collected O. oglinum in the higher-salinity waters of lower estuaries and

ocean beaches along the Georgia coast. Small numbers were reported throughout the year in the bycatch from inventories

of the penaeid shrimp resource in coastal and estuarine areas of Georgia and Florida (Anderson 1968).

During the present study, O. oglinum ranked 20th in numerical abundance but only 34th by weight, since they were generally

small fishes that contributed only 0.36% to the total catch.

Opisthonema oglinum was taken in 105 of the trawl tows made during the survey. Although this species was encountered

throughout the study area, seasonal differences in the frequencies of occurrence and distribution patterns were apparent (Fig.

26). There was a significant difference in the frequency of occurrence between seasons (x2 = 28.70, df = 3, p >0.05).

Opisthonema oglinum occurred in significantly fewer trawl tows during winter than in spring (x2 = 17.00, df = 1, p>O.05),

summer (x2 = 23.88, df = 1, p>O.05) or fall (x2 = 4.01, df = 1, p>O.05). Frequencies of occurrence in spring and summer collections were not significantly different (x2 = 0.18, df = 1), whereas both of these seasons had significantly greater

frequencies than during the fall (spring-fall X2 = 4.64, df = 1, P >0.05; summer-fall X2 = 9. 17, df = 1, p >0.05).

Seasonal catches indicated that during the winter months the distribution of O. ogli'1um was restricted primarily to the

coastal waters off Florida (Fig. 26). With the onset of spring, thread herring dispersed throughout the Bight; however, catch

rates and frequency of occurrence were still greater in the southern strata. In summer, O. oglinum was fairly ubiquitous

throughout the survey area. With the cooling of the coastal waters in fall, this species migrated to the south. The seasonal

changes in this species' distribution pattern inferred from trawl catches is supported by additional evidence from tagging

studies. Pristas and Cheek (1972) tagged fish off Beaufort, North Carolina, during September and recoveries showed that

by November these individuals had migrated south 592 Ian to the coastal waters off northern Florida.

The stratified mean catch/tow (in number) was lowest in the spring and highest in the fall (Table 5). The fall value was

highest because of elevated catch rates in the southern part of the survey area during this season.

Opisthonema oglinum has a protracted spawning season (April-September) in the eastern Gulf of Mexico (Houde 1977).

A more restricted period was reported off Beaufort, North Carolina (Hildebrand 1963a). Initial estimates of length-at-age

for O. oglinum from commercial purse catches in the eastern Gulf of Mexico were: age-I 146 mm FL; age-IT 149 mm FL.

Estimates of age-ill were considered unreliable because of the small sample size (Fuss et al. 1969). The methodology was

not validated and sampling was biased by the gear. We did not age or document the reproductive biology of this species

during this study; however, some very tentative inferences can be made about these aspects of the natural history of O. oglinum

from the seasonal length-frequencies of the trawl catches.

The length composition of the fish caught during the winter survey had one apparent mode (8 cm FL, Fig. 27). Two modes

(10 cm FL and 14 cm FL) were observed during spring, and summer samples had smaller fish (4-6 cm FL) that were absent

during other seasons. These small fish were probably the result of a spring spawning event in the SAB and represented the

size of first recruitment to the fishing gear. The fall survey had length frequencies with modes at 8 cm FL and 15 cm FL,

and lengths in winter trawl tows were also bimodal (8 and 15 cm FL). We interpret these data as indicating that during the

spring, age-I fish ranged in length between 9 and 13 cm FL, while age-IT were from 13 to 16 cm FL. The smaller fish taken

during the summer survey were young-of-year. Further estimates from these data could be misleading because of the nature

of the distribution in the larger length classes and the possibility of size-related changes in migratory patterns and vulnerability to the gear.

34
Atlantic thread herring Opisthonema oglinum

X Number/Tow

FALL Opisthonema

oglJnum

Do

c:3 '-20

21-50

.51+

Figure 26

Seasonal catch rates in

coastalstrata. nIN= 110. tows

with Atlantic thread berringI

no. tows in stratum.

Fork Length (em)

Season

·-·Winter (n=452)

• - -. Spring (n=362)

.- - .• Summer (n=630)

.----. Foil (n=1.379)

Figure 27

Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.

15 20

-

•.

5 10

Opisthonema

40 o~in~m

10

50

20

30

>-

()

c

Q)

0-

Q)

L

l.J....

--C

Q)

()

L

Q)

CL

35
Pomatomus saltatrix _

Bluefish is a pelagic schooling species of commercial and recreational importance that occurs throughout the Gulf of Mexico,

along northern Cuba, Bermuda, and the Atlantic coast of the United States to Nova Scotia (Fischer 1978). This species is

also found along the eastern coast of South America, in the eastern Atlantic and Mediterranean Sea, and in the Indo-West

Pacific (Fischer 1978). Along the Atlantic coast of the United States, bluefish are highly migratory. A generalized movement

pattern has emerged from ichthyoplankton and tagging studies, but specific movements from all areas are unknown. This

is especially true for juvenile bluefish from the coastal and estuarine habitats of the SAB. The seasonal movements of adult

fishes (north in spring, south in fall) are well documented (Lund and Maltezos 1970, Wilk 1977, Kendall and Walford 1979).

During fall, cooling of waters on the continental shelf in the northern part ofthe species range along the eastern U.S. stimulates

the movement of adult bluefish to warmer waters offshore and south (Lund and Maltezos 1970). Some individuals migrate

to waters off southeastern Florida (Wilk 1977), whereas others from southern New England apparently move southeast to

the warmer waters of the upper continental slope in the vicinity of Hudson's Canyon where they have been taken in otter

trawl gear (Lund and Maltezos 1970, Bigelow and Schroeder 1953). In the spring, bluefish migrate from shelf waters off

southeastern Florida to higher latitudes (Wilk 1977).

Ichthyoplankton surveys have found two distinct periods and areas of peak reproduction of bluefish along the Atlantic coast.

South ofCape Hatteras, N.C. (approx. 35°N), major concentrations oflarval P. sa/tatrix were found inside the 183-m (1oo-fin)

curve from 33°N to 34°N during May (Kendall and Walford 1979) and offshore of the continental shelf north of 32° in

April and May (Collins and Stender 1987). Powles and Stender (1976) and Collins and Stender (1987) found larval P. sa/tatrix

east of the 1oo-fin curve in the SAB in significant numbers during May. In the Middle Atlantic Bight, maximum larval abundance was found in midshelf waters from 38°N to 40 0 N during August (Kendall and Walford 1979). It should be noted,

however, that spawning activity as demonstrated by the presence of larvae occurs winter, spring, and autumn (Powles and

Stender 1976, Kendall and Walford 1979, Collins and Stender 1987) in the South Atlantic and June through September in

the Middle Atlantic Bight (Kendall and Walford 1979).

With the exception of the schools of adult fishes that occur around artificial reefs and jetties during early spring, the bluefish

found in the coastal habitat ofthe SAB are juveniles. Kendall and Walford (1979) present a generalized scheme of recruitment

pathways of larval and juvenile bluefish into estuarine nursery areas, but there is an obvious lack of information on juvenile

bluefish in the nursery areas of the SAB. Larval bluefish are primarily neustonic and have a life-cycle involving spawning

in warm, saline, offshore waters and movement to cooler, less saline, nearshore waters (Powles 1981, Kendall and Walford

1979). Powles (1981) found the smallest larval bluefish nearest the 180-m contour in the SAB, and Collins and Stender (1987)

reported maximum abundance of larvae ~4 mm over the outer continental shelf. Larval size increases with proximity to

shore (Powles 1981, Collins and Stender 1987). Collins and Stender (1987) suggested that a southerly flowing countercurrent

(Bumpus 1973) may provide a mechanism for movement of larvae to nearshore waters within the SAB.

Along the South Atlantic coast, P. sa/tatrix has been found from the surf zone (Anderson et al. 1977) to offshore waters

at the outer continental shelf (37-55 m) (Wenner unpubl. data). This species occurs in all estuarine systems of the area from

the Cape Fear River estuary in North Carolina (Schwartz et al. 1982), South Carolina (Wenner et al. 1982), and Georgia

(Mahood et al. 1974a,b,c). Bluefish have been found in the bycatch of the commercial shrimp fishery off South Carolina

(Keiser 1976), Georgia (Knowlton 1972), and Florida (Anderson and Gehringer 1965). In South Carolina coastal waters,

young bluefish accounted for 0.49% and 0.89% ofthe number and weight, respectively, offishes in the bycatch over a 14-month

period (Keiser 1976). Seventy-seven percent of the total number and 65.7% of the weight were found during June, July,

and August (Keiser 1976). Only six specimens weighing 0.3 kg were collected from December to May (Keiser 1976). In

Georgia, P. sa/tatrix was a minor constituent ofthe bycatch, comprising <0.1 (July) to 2.5 % (November) of the total weight

of the bycatch of fishes (Knowlton 1972). Catch rates were 0 or very low «0.1 kg/h) from December to April. Bluefish

were taken in trawl tows during all months in estuarine systems of Georgia (Mahood et al. 1974a,b,c). Largest catches occurred

in April, May, and June; these accounted for 55 % of the total weight of bluefish taken during the annual survey.

In the present study, bluefish ranked 30th by numerical abundance (980 individuals, 0.28% of the catch) and 18th by weight

(Table 3) for the four surveys. Forty-one percent ofthe 305 total trawl collections caught bluefish. Sampling showed seasonal

changes in the distribution, frequency of occurrence, and abundance in the SAB. Bluefish were in only a single trawl haul

(southern South Carolina) along the coast of the Carolinas during winter (Fig. 28), but were more abundant in strata to the

south. During the other seasons, this species was widely distributed throughout the Bight. There was a significant difference

in the frequency of occurrence of bluefish between seasons (x2 = 19.91, df = 3,p>O.05). Winter sampling found P. sa/tatrix

in significantly fewer trawl collections (11 of 54) than spring (33 of 54; X2 = 18.56, df = 1, p>O.05), summer (40 of 108;

X2 = 4.63, df = 1, p>O.05), or fall tows (40 of 89; X2 = 7.57, df = 1, p>O.05). Seasonal values of the stratified mean

catch-per-tow for bluefish were:

36
Bluefish Pomatomus saltatrix

Season

Winter

Spring

Summer

Fall

Yst no.

3.7

4.4

1.5

3.5

Yst kg

0.15

0.43

0.14

0.47

Although bluefish may forage throughout the water column, they are primarily a strong-swimming pelagic species (Wille

1977). The present study reports the catches of a four-seam, demersal shrimp trawl net. Therefore, these catch rates with

their means should be considered relative.

Although the seasonal surveys were not sequential, it is possible to follow the length composition of bluefish in the coastal

habitat of the SAB over an "annual cycle" (Fig. 29). Wilk (1977) presented the following mean-Iength-at-age estimates for

P. saltatrix aged by scales: 121 cm FL; II 35 cm FL; III 41 cm FL. He determined that the annulus was laid down during

spring by marginal increment analysis. Based on this information, bluefish collected during the winter survey (x FL = 16 cm; mode = 15 cm; range 12-23 cm) were young-of-year (yay) probably spawned during the late winter or spring

(Powles and Stender 1976, Kendall and Walford 1979, Collins and Stender 1987). The spring survey collected some small

individuals (7-10 cm FL) that may have resulted from the fall spawn in the SAB (Kendall and Walford 1979). The overall

mean length was 19 cm FL (mode 18 cm; range 7-31 cm), and according to Wilk (1977) these were almost 1 year-old, i.e.,

the first annulus was about to be laid down. Bluefish from the summer survey had a large length range (12-33 cm FL;

x 18 cm) and were probably yay and age-I fish. Fall fish averaged 21 cm FL (range 9-30 cm), and these were probably

yay and age-I fish as in the summer collections.

From these data, it appears that the coastal waters of the SAB provide habitat for a significant number of yay and age-I

bluefish. In addition, personal observations indicate that bluefish of sizes similar to those captured in the coastal surveys

inhabit the estuaries of South Carolina during the warmer months of the year. The relative abundance of larger fish (ages

II and older) during the year does not reflect the abundance of juvenile bluefish in South Carolina waters. This strongly

suggests that with increased length and age, bluefish that utilized both the estuaries and coastal waters ofthe SAB as "nursery

areas" joined the bulk of the larger fish that have large seasonal migrations along the Atlantic coast. The relative contribution

of this "stock" to the western Atlantic population has not been documented and in most cases it is ignored (Wilk 1977).

However, estuaries in coastal South Carolina and Georgia are extensive, and bluefish from these areas may make a significant

contribution to the harvest in northern waters as they mature.

37
Pomatomus saltatrix _

FALL

X Number/Tow

Pomstomus

ss/tstrlx

Do

~1-10

~11-20

.20+

Figure 28

Seasonal catch rates in

coastal strata. nlN= no. tows

with bluefish/no. tows in

stratum.

30

Pomatomus

Figure 29

Seasonal length-frequency

distribution in South Atlantic Bight coastal waters.

30 35

•

Season

25

e-eWinter (n=151)

.--. Spring (n=269)

•...• Summer (n=144)

0----0 Fall (n=407)

\

,

\

15 20

Fork Length (em)

5 10

saltatrix

o+--.~~--,-~~-t'~f--....~~....,..,:.:~_,__:;..:.;..--,-::,.>_r_~~~~~.•:..._,

o

20

-+-

C

Q) 10

o

L

Q)

0...

>-

o

c

Q)

::J

0-

Q)

L

l..L.

38
Community analysis

Normal cluster analysis formed 11 groups of sites, and

clusters primarily reflected the similarity ofspecies composition within the season of collection. Geographic area appeared to have a secondary influence on the cluster results

(Fig. 30). Group 1consisted mainly of collections from North

Carolina through Georgia. Many ofthese entities were united

because they consisted of trawl collections off Georgia in

spring. Northerly stations that were sampled in summer and

fall also occurred in this group, perhaps indicating seasonal

migration of similar species between these regions. Collections in Groups 2 and 3 consisted of stations off South

Carolina that were sampled in all seasons except winter. Most

of these stations were sampled in spring and summer. Collections in Groups 4 and 5 were all made in fall, and most

were from South Carolina and Florida waters. Group 6

stations were primarily from northern Florida and southern

Georgia in summer. Group 7 stations were primarily southern

stations sampled in spring and fall. Station Group 8 included

collections from Florida in winter and South Carolina in summer and fall, indicating occurrence ofsimilar species in these

relatively widely separated geographic areas during different

times of the year. Group 9 consisted of stations that were

located throughout the SAB during the cooler seasons of

winter and spring. Group 10 comprised stations from Florida

and Georgia sampled during the warmer seasons, and Group

11 comprised mainly stations from Georgia and South

Carolina waters sampled in winter.

Inverse cluster analysis formed 14 groups ofspecies with

similar distributions (Table 6, Fig. 31), some of which

showed high constancy at several site groups (Fig. 32). In

general, however, fidelity values were low, indicating broad

latitudinal distribution of species groups over the four

seasons. Species in Group A were small, benthic species that

were found most often off Florida and Georgia, generally

during the warmer seasons. Group A species were most frequent at stations in site Group 10 (Florida and Georgia in

summer and fall), and showed moderate fidelity to that site

group.

Group B included some schooling species (S. vomer, S.

maculatus, S. aculeatus, R. terraenovae). These species were

frequently taken from site Group 3, spring and summer collections off South Carolina. Collins and Wenner (1988) found

increased abundance of S. maculatus in trawl catches in the

South Atlantic Bight in summer, and clustering of Group B

species may reflect the abundance of newly recruited

juveniles to coastal waters of the Bight. Large numbers of

other Group B species, such as S. aculeatus, R. terraenovae,

and P. carolinus, occur in shallower waters in summer (Fritzsche 1978, Sedberry and Van Dolah 1984, Parsons 1983),

reflecting seasonal migration and recruitment in these species.

Group B species showed low fidelity to all site groups. Group

C species showed high fidelity to site Group 11 (winter stations off South Carolina and Georgia). Squalus acanthias,

a member of Group C, is commonly found in coastal waters

of the southeast only in winter (Struhsaker 1969), as are

39

migrating American shad (Leggett and Whitney 1972, Van

Dolah, et al. 1987).

Species Groups D,J,K,L,M, and N comprised species that

appeared to be ubiquitous, year-round residents of coastal

waters, based on within-group uniformity of constancy levels

and a concomitant low fidelity for all site groups. Species

in Group D were relatively rare species that showed low constancy to all site groups, whereas Groups J-N were frequently

taken (i.e., high in constancy) at most site groups. These

groups included very abundant species such as those in Group

L, which demonstrated very high constancy at all site groups,

as well as other species collected throughout the Bight.

Species in Groups J and M were high in constancy at all site

groups except Group 11, which were mainly winter stations

off South Carolina and Georgia. Many of these species were

sciaenids known to migrate to offshore waters and to the

south in winter (Chao and Musick 1977) as well as other

species known to have similar winter movements. It is

noteworthy that spotted hake, U. regia, occurred in Group

J. Based on reported seasonal offshore-inshore migrations

for this species, it should occur relatively frequently in coastal

waters in winter (Sikora et al. 1972, Burr and Schwartz

1986). Apparently spotted hake had moved farther inshore

into high-salinity estuaries during the winter sampling period.

January movement into estuaries is well-documented for

spotted hake (Barans 1972, Sikora et al. 1972, Burr and

Schwartz 1986). Species in Group N were high to very high

in constancy at all site groups with the exception of Group

5 that consisted of all fall collections off Florida.

Groups E through I, unlike many other species groups,

demonstrated moderate to high fidelity for at least one site

group. Species in Group E were moderately faithful to site

groups from summer and fall southern strata (Georgia and

Florida). Group F species were moderately faithful to sites

from South Carolina in spring and summer. Group G consisted mainly of elasmobranch species that were relatively

rare in collections, but occurred frequently in fall samples

(site Groups 4 and 5), probably reflecting coastal migratory

patterns exhibited in many coastal elasmobranch species

(Smith and Merriner 1986). The occurrence of many species

of large elasmobranchs to the exclusion of most small teleosts

has been noted previously, and may indicate competition

between the two groups or avoidance of predation by the

smaller teleosts (Longhurst 1969). Species Groups H and I

were also frequently taken in fall samples (site Groups 4 and

5).

Highest diversity (H') was in summer (Fig. 33), and H'

diversity values appeared to be more influenced by the evenness of distribution of individuals among species (Fig. 34)

than by species richness (Fig. 35). The overwhelming

dominance of spot (Table 2), which was at least one order

of magnitude more abundant than all other species, contributed to the low evenness values, and low H' values often

coincided with high abundances of spot. Lowest values of

H' were found in winter and spring when evenness and, particularly in winter, species richness values were low. Low

diversity values in winter probably reflect migration offshore
:::> Z I-- w 0 SIMILARITY < I-- (/)

[[ < < 0.8 0.6 0.4 0.2 0.0 -0.2 -0.4 -0.6 -0.8 I-- I-- W (/) (/) (/) I I I I I I I I I

35 GA SP

37 GA SP

39 GA SP

45 se SP

31 GA SP

55 SeiNe SP

53 se FA

55 seINe FA

55 seINe su

47 se SP

41 se FA

51 se su

53 se SP

43 se su

51 se SP

47 se su

41 se SP

49 se SP

45 se su

49 se su

41 se su

43 se FA

45 se FA

47 se FA

49 se FA

35 GA FA

22 FL FA

26 FL FA

24 FL FA

28 FL FA

30 FL rA

28 FL SU

30 FL SU

31 GA SU

33 GA SU

35 GA SU

31 GA FA

37 GA SU

39 GA SU

22 FL sp

30 FL sp

37 GA FA

39 GA FA

30 FL WI

24 FL WI

26 FL WI

22 FL WI

53 se su

51 se FA

26 FL SU

28 FL WI

31 GA WI

43 se WI

28 FL SP

41 se WI

55 seiNe WI

24 FL SP

26 FL SP

33 GA SP

22 FL SU

24 FL SU

33 GA FA

35 GA WI

49 se WI

47 se WI

51 se WI

53 se WI 11

37 GA WI

45 se WI

39 GA WI

43 se sp

33 GA WI

Figure 30

Dendrogram depicting normal cluster analysis of trawl collections pooled by stratum within season. Strata

numbers correspond to those in Figure 1.

40
Table 6

Species groups resulting from inverse cluster analysis of

trawl data by season, pooled by stratum. 0.6

I

0.4

I

0.2

I

SIMILARITY

0.0 -0.2

I I

-0.4

I

-0.6

I

-0.8

I

Group A

Eucinostomus argenteus

Eucinostomus gula

Syacium papillosum

Etropus cyclosquamus

Centropristis striata

Group B

Selene vomer

Chilomycterus schoepfi

Citharichthys spilopterus

Scomberomorus maculatus

Stenotomus aculeatus

Rhizoprionodon terraenovae

Prionotus evolans

Ancylopsetta quadrocellata

Paralichthys dentatus

Group C

Squalus acanthias

Clupea harengus

Menticirrhus saxatilis

Alosa sapidissima

Group D

Trachurus lathami

Urophycis jloridana

Diplectrum formosum

Decapterus punctatus

Hippocampus erectus

Dasyatis sayi

Urophycis earlli

Hypsoblennius hentzi

Etrumeus teres

Group E

Narcine brasiliensis

Paralichthys albigutta

Porichthys plectrodon

Sphyrna tiburo

Caranx hippos

Caranx crysos

Paralichthys squamilentus

Sphyrna lewini

Trachinotus carolinus

Group F

Monacanthus hispidus

Syngnathus louisianae

Scomberomorus cavalla

Ophidion marginatum

Rhinobatus lentiginosus

Astroscopus y-graecum

Group G

Dasyatis americana

Gymnura micrura

Myliobatis freminvillei

Dasyatis centroura

Ophidion grayi

Scorpaena calcarata

Rhinoptera bonasus

Group H

Sphyraena guachancho

Menticirrhus littoralis

Harengula jaguana

Sardinella aurita

Anchoa sp.

Lagodon rhomboides

Group I

Etropus microstomus

Menidia menidia

Mustelus canis

Syngnathus fuscus

Paralichthys lethostigma

Membras martinica

Sphoeroides maculatus

Dasyatis sabina

Ogcocephalus rostellum

Group J

Cynoscion regalis

Trinectes maculatus

Peprilus triacanthus

Pomatomus saltatrix

Raja eglantaria

Prionotus tribulus

Bairdiella chrysoura

Urophycis regia

Group K

Centropristis philadelphica

Peprilus alepidotus

Prionotus salmonicolor

Prionotus scitulus

Orthopristis chrysoptera

Citharichthys macrops

Etropus crossotus

Synodus foetens

Arius felis

Chaetodipterus faber

Group L

Leiostomus xanthurus

Micropogonias undulatus

Brevoortia tyrannus

Group M

Larimus fasciatus

Menticirrhus americanus

Chloroscombrus chrysurus

Selene setapinnis

Opisthonema oglinum

Trichiurus lepturus

Anchoa hepsetus

Stellifer lanceolatus

Cynoscion nothus

Group N

Anchoa lyolepis

Anchoa cubana

Prionotus carolinus

Scophthalmus aquosus

Symphurus plagiusa

Anchoa mitchilli

Figure 31

Dendrogram depicting inverse cluster analysis of trawl coUections pooled by stratum

within season. Species groups correspond to those in Table 6.

41
SITE GROUPS SITE GROUPS

< 1 VerYILow

::: 1 Low

2 Moderate

4 Very High IJZ]

3 High 0

FIDELITY

A

B

C

o

E

Ff.f.i;!;I;;;;b"",j~!"""

Gp""''''9=''f'''~

Hf--j--+---l'''~=

I

J

K

L

M

N""",=""",,,,,,,,,,,

< 0.1 Very Low

::: 0.1 Low

::: 0.3 Medium

CONSTANCY

::: 0.7 Very High [IT]

:::0.5 High 0

.2

A

B

C

0

E

F

G

H

I

J

K

L

M

N

I

.2

-.8

>- -.6 !:::

a::

< -.4

...J

-.2

(ii

SIMILARITY

I , I I I I I I I I

-.8 -.6 -.4 -.2 0

C/)

a..

::::>

o

a::

"

C/)

W

(3

w

a..

C/)

Figure 32

Nonnal and inverse cluster hierarchies and nodal constancy and fidelity diagrams for trawl collections pooled by stratum within season.

5 5

Winter Summer

4 • 4 • •• • • • • • • • • • • • • • • • • • 3 • • 3

• • • • 2 • • • 2

• • • ".-... • I 1 1

• • '--"

>- 0 0 ......

en

l0-

a>

.2:

0 5 5

Spring Fall

4 • 4 • • • •• • • • • • •• • • • • • 3 • • • • 3 • • • • • • • • 2

•• 2

• • • 1 1

• 0 0

20 25 30 35 40 45 50 55 60 20 25 30 35 40 45 50 55 60

Strata

Figure 33

H' diversity values for collections pooled by stratum within season. Strata numbers correspond to those in Figure 1.

42
1.0 1.0

Winter Summer

0.8 • 0.8 • • • • • • • • • • • • • • • • • 0.6 • • • • 0.6 • • • • • • • • r--.. 0.4 • 0.4 - • • -,

'-" 0.2 • 0.2

Vl • • Vl

Q) 0.0 0.0

C

C

Q)

>

W 1.0 1.0

Spring Fall

0.8 0.8 • • • • • • • • •• • • 0.6 • 0.6 • • • • • • • • • • • • • • • 0.4 • 0.4 • • • • • • 0.2 0.2

• 0.0 0.0

20 25 30 35 40 45 50 55 60 20 25 30 35 40 45 50 55 60

Strata

Figure 34

J' evenness values for collections pooled by stratum within season. Strata numbers correspond to those in Figure 1.

8 8 Summer

7 Winter 7 • • • • 6 6 • • • • • • • • • • • • 5 • • 5 • • • 4 • • 4

• • • • • Vl 3 • • 3 Vl • •• • <l> 2 • • 2 c

£ 1 1

U 0 0

0:::

Vl

<l>

U

<l> 8 8

0.. Fall • (f') 7 Spring 7 • • • • • • • 6 6 • • • • • 5 • • • 5 • • • • • • •• • • • • • • • 4 • 4 • • • • 3 3

2 2

1 1

0 0

20 25 30 35 40 45 50 55 60 20 25 30 35 40 45 50 55 60

Strata

Figure 35

Species richness values for collections pooled by stratum within season. Strata numbers correspond to those in Figure 1.

43
or to the south of the study area of many species found in

coastal waters during warmer seasons, and an overwhelming dominance (62.9% of all individuals) of the fauna by only

two species, spot and Atlantic menhaden (Table 2). In spring,

the fauna was largely dominated by spot (55.1 % of all individuals); evenness and, therefore, H' values were low. In

summer and fall, the two most abundant species comprised

only 22.8% and 29.2 % ofthe total individuals, respectively,

and there was greater evenness of distribution of individuals

among species, in addition to greater species richness.

There was considerable variation in diversity among collections within seasons and strata. Diversity values were

similar in range to those reported from other habitats in the

SAB (Sedberry and Van Dolah 1984, Wenner 1983).

In conclusion, the coastal habitat of the South Atlantic

Bight supports a fish fauna dominated by the family

Sciaenidae, with many other higWy migratory species of

fishes. Many of the genera of sciaenids, as well as other

genera, are typical of those found in coastal soft-bottom

habitats in other parts ofthe subtropical and temperate Atlantic (Lowe and Longhurst 1961, Lowe 1962, Longhurst 1969).

Families, genera, and many species that form the coastal fish

community in the SAB are identical to those found in similar

habitats in the Gulf of Mexico, off South America and the

Atlantic coast of Africa (Longhurst 1969), and familial composition is similar to that found in soft-bottom estuarineinfluenced coastal habitats in other oceans (Longhurst 1969).

Patterns of biomass, abundance, and community structure

reflect the seasonal migration patterns of the dominant

species, as well as recruitment of juveniles to the coastal

nursery area.

Acknowledgments _

We thank all who assisted in field work, especially W.A.

Roumillat, D. Stubbs, C.W. Waltz, and C.A. Barans.

MJ. Clise and W.P. Coon assisted with data processing and

K. Swanson prepared many ofthe figures. C.A. Barans and

M.R. Collins reviewed the manuscript. This work was supported by the MARMAP contract between the National

Marine Fisheries Service, NOAA, and the Marine Resources

Research Institute of the South Carolina Wildlife and Marine

Resources Department.

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Appendix 1

Seasonal presence-absence of species and families of fishes collected during the coastal survey. Plus sign (+) indicates presence.

Family Family

Species Winter Spring Summer Fall Species Winter Spring Summer Fall

CARCHARHINIDAE GOBIESCOCIDAE

Carcharhinus acronotus + + Gobiesox strumosus +

Carcharinus limbatus + OGCOCEPHALIDAE Mustelus canis + + +

Rhizoprionodon terraenovae + + + Ogcocephalus comiger +

Ogcocephalus cubifrons +

SPHYRNIDAE Ogcocephalus rostellum + + +

Sphyraena lewini + + + GADIDAE Sphyraena tiburo + + + + Urophycis earlli + + + +

SQUALIDAE Urophycis floridana +

Squalus acanthias + Urophycis regia + + +

RHINOBATIDAE OPHIDIIDAE

Rhinobatos lentiginosus + + + Ophidion beani +

TORPEDINIDAE Ophidion grayi +

Narcine brasiliensis + + + Ophidion holbrooki +

Ophidion marginatum + + + +

RAJIDAE EXOCOETIDAE Raja eglanteria + + + + Hyporhamphus unifasciatus + +

DASAYATIDAE ATHERINIDAE Dasyatis americana + + + + Membras martinica + Dasyatis centroura + + Menidia menidia + + Dasyatis sabina + + + +

Dasyatis sayi + + + + SYNGHATHIDAE

Gymnura micrura + + + Hippocampus erectus + + + +

MYLIOBATIDAE Sygnathus fuscus + +

Myliobatis freminvillei Sygnathus louisianae + + + + + + + Sygnathus pelagicus + Rhinoptera bonasus + +

ELOPIDAE PERCICHTHYIDAE

Elops saurus Morone americana + +

OPHICHTHIDAE SERRANIDAE

Ophichthus ocellatus Centropristis ocyurus +

+ Centropristis philadeliphica + + + +

CLUPEIDAE Centropristis striata + + + +

Alosa mediocris + Diplectrum formosum + +

Alosa sapidissima + Serraniculus pumilio + +

Brevooria tyrannus + + + + PRIACANTHIDAE Clupea harengus + Priacanthus arenatus + Dorosoma cepedianum +

Etrumeus teres + POMATOMIDAE

Harengula jaguana + + + + Pomatomus saltatrix + + + +

Opisthonema oglinum + + + + RACHYCENTRIDAE Sardinella aurita + + + Rachycentron canadum + +

ENGRAULIDAE ECHENEIDAE Anchoa cubana + + + + Echeneis naucrates Anchoa hepsetus +

+ + + +

Anchoa lyolepis + + + + CARANGIDAE

Anchoa mitchilli + + + + Alectis ciliaris +

Engraulis eurystole + Caranx crysos + + +

SYNODONTIDAE Caranx hippos + + +

Synodus foetens Chloroscombrus chrysurus + + + +

+ + + + Decapterus punctatus + +

ARIIDAE Selene setapinnis + + + +

Arius felis + + + Selene vomer + + +

Bagre marinus + + Trachinotus carolinus + +

BATRACHOIDIDAE Trachurus lathami + +

Opsanus pardus + + + LUTJANIDAE

Opsanus tau + Lutjanus analis +

Porichthys plectrodon + + + Lutjanus synagris +

48
Appendix 1 (Continued)

Family Family

Species Winter Spring Summer Fall Species Winter Spring Summer Fall

GERREIDAE STROMATEIDAE

Eucinostomus argenteus + + Peprilus alepidotus + + + +

Eucinostomus gula + + + Petrilus triancanthus + + + +

HAEMULIDAE SCORPAENIDAE

Orthopristis chrysoptera + + + + Scorpaena agassizi +

SPARIDAE Scorpaena brasiliensis + +

Archosargus probatocephalus + Scorpaena calcarata + +

Calamus leucosteus + Scorpaena plumieri +

Diplodus holbrooki + TRIGLIDAE

Lagodon rhomboides + + + + Prionotus carolinus + + + +

Stenotomus aculeatus + + + Prionotus evolans + + + +

SCIAENIDAE

Prionotus ophryas + +

Bairdiella chrysoura

Prionotus salmonicolor + + + +

+ + + + Prionotus scitulus + + + + Cynoscion nebulosus + Prionotus tribulus + + + + Cynoscion nothus + + + +

Cynoscion regalis + + + + BOTHIDAE

Larimus fasciatus + + + + Ancylopsetta quadrocellata + + + +

Leiostomus xanthurus + + + + Citharichthys macrops + + + +

Menticirrhus americanus + + + + Citharichthys spilopterus + + + +

Menticirrhus littoralis + + + + Etropus crossotus + + + +

Menticirrhus saxatilis + + + Etropus sp. A + + +

Micropogonias undulatus + + + + Etropus microstomus + + +

Pareques acuminatus + Paralichthys albigutta + + +

Stellifer lanceolatus + + + + Paralichthys dentatus + + + +

Umbrina coroides + + Paralichthys lethostigma + + + +

MULLIDAE Paralichthys squamilentus + +

Upeneus parvus + Scopthalmus aquosus + + + +

Syacium gunteri +

EPHIPPIDAE Syacium papillosum + + + +

Chaetodipterus faber + + + + SOLEIDAE

LABRIDAE Gymnachirus melas + + +

Hemipteronotus novacula + Trinectes maculatus + + + +

Tautoga onitis + + CYNOGLOSSIDAE

SPHYRAENIDAE Symphurus plagiusa + + + +

Sphyraena guachancho + + + BALISTIDAE

URANOSCOPIDAE Aluterus schoepfi + +

Astroscopus y-graecum + + + Balistes capriscus +

BLENNIIDAE Monacanthus hispidus + + + +

Hypleurochilus geminatus + + TETERAODONTIDAE

Hypsoblennius hentzi + + + + Lagocephalus laevigatus +

GOBIIDAE Sphoeroides maculatus + + +

Gobiosoma bosci + DIODONTIDAE

Gobiosoma ginsburgi + Chilomycterus antillarum +

TRICHIURIDAE Chilomycterus schoepfi + + +

Trichiurus lepturus + + + +

SCOMBRIDAE

Scomberomorus cavalla + +

Scomberomorus maculatus + + +

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