NOAA Technical Report NMFS 86 February 1990

Benthic Macrofauna and Habitat

Monitoring on the Continental Shelf

of the Northeastern United States

I. Biomass

Frank W. Steimle

U.S. Department of Commerce
NOAA TECHNICAL REPORT NMFS _

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5285 Port Royal Road, Springfield, VA 22161. Although the contents have not been copyrighted and may be reprinted entirely, reference to source is appreciated.

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

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

57 p.

49. Reproduction. movement,. and populatIOn dynamics of the southern kingfish,

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, Arcrica islandica,

for age determinattons. 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.

5:!. Limited access alternatives for the Pacific groundfish fishery, by Daniel D.

Huppert (editor). May 1987,45 p.

53. Ecology of east Florida sea tunles: 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. finfish 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 tunle Dermochelys coriacea at

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

57. Food habits and dietary variability of pelagic nekton off Oregon 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 menhaden, Brevoorria parronus, fishery, by Douglas

S. Vaughan. September 1987, 18 p.

59. Atlantic menhaden, Brevoorria Ivmnnus, purse seine fishery, 1972-84, with a

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

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

September 1987, 23 p.

60. Gulf menhaden, BrevoorrlO parronl/s. 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 ,tarch 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, Jacki A. Guffey, and Joni M. Packard.

December 1987, 149 p.

63. Stock asse>sment of the Atlantic menhaden, Brevoorria ryrarmus, 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.

67. Index numbers and productivity measurement in multispecies fisheries: An

application to the Pacific coast trawl fleet, by Dale Squires. July 1988, 34 p.

68. Annotated bibliography II of the hard clam Mereenaria mereenaria, by JL.

McHugh and Marjorie W. Sumner. September 1988, 59 p.

69. Environmental quality and aquaculture systems: Proceedings of the thirteenth

U.S.-Japan meeting on aquaculture, Mie, Japan, October 24-25, 1984, edited by Carl

J. Sindermann. October 1988, 50 p.

70. New and innovative advances in biology/engineering with potential for use in

aquaculture: Proceedings of the fourteenth U.S.-Japan meeting on aquaculture, Woods

Hole, Massachusetts, October 16-17, 1985, edited by Albert K. Sparks. November

1988, 69 p.

71. Greenland turbot Reinhardrius hippoglossoides of the eastern Bering Sea and

Aleutian Islands region, by Miles S. Alton, Richard G. Bakkala, Gary E. Walters,

and Peter T. Munro. December 1988, 31 p.

72. Age determination methods for northwest Atlantic species, edited by Judy Penttila

and Louise M. Dery. December 1988, 135 p.

73. Marine flora and fauna of the Eastern United States. Mollusca: Cephalopoda.

by Michael Vecchione, Clyde F.E. Roper, and Michael J. Sweeney. February 1989,

23 p.

74. Proximate composition and fatty acid and cholesterol content of 22 species of

northwest Atlantic finfish, by Judith Krzynowek, Jenny Murphy, Richard S. Maney,

and Laurie 1. Panunzio. May 1989, 35 p.

75. Codend selection of winter flounder Pseudoplel/rolleeres americanus, by David

G. Simpson. March 1989, 10 p.

76. Analysis of fish diversion efficiency and survivorship in the fish return system

at San Onofre Nuclear Generating Station, by Milton S. Love, Meenu Sandhu,

Jeffrey Stein, Kevin T. Herbinson, Robert H. Moore, Michael Mullin, and John S.

Stephens, Jr. April 1989, 16 p.

77. Illustrated key to the genera of free-living marine nematodes of the ordet

Enoplida, by Edwin J. Keppner and Armen C. Tarjan. July 1989,26 p.

78. Survey of fishes and water properties of south San Francisco Bay, California,

1973-82, by Donald E. Pearson. August 1989, 21 p.

79. Species composition, distribution, and relative abundance of fishes in the COdSta!

habitat off the southeastern United States, by Charles A. Wenner and George R

Sedberry. July 1989, 49 p.

80. Laboratory guide to early life history stages of northeast Pacific fishes, by Ann

C. Matarese, Arthur W. Kendall, Jr., Deborah M. Blood, and Beverly M. Vinter.

October 1989, 651 p.
NOAA Technical Report NMFS 86

Benthic Macrofauna and Habitat

Monitoring on the Continental Shelf

of the Northeastern United States

I. Biomass

Frank W. Steimle

February 1990

~"'~ OF co U.S. DEPARTMENT OF COMMERCE l'" Il,f+~ Robert Mosbacher, Secretary

.: 1£ ('\ = 1!'l : National Oceanic and Atmospheric Administration

e - - iJ John A. Knauss, Under Secretary for Oceans and Atmosphere

: ~<l k- ~"'~ National Marine Fisheries Service

.rr~TES Of William W. Fox, Jr., 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.

II
CONTENTS

Acknowledgments iv

Introduction 1

Methods 2

Results 3

Chesapeake Bight 4

Station 2 4

Station 3 4

Station 7 5

Station 8 6

Station 9 7

Station 31 7

Station 32 8

New York Bight 8

Station 11 8

Station 12 11

Station 13 11

Station 17 11

Station 33 12

Station 16A 12

Station 16B 13

Station 16C 13

Station 15A 13

Southern New England 15

Station 18 15

Station 34 17

Station 36 17

Station 19 18

Station 20 18

Station 37 18

iii

Georges Bank 20

Station 22 20

Station 23 20

Station 24 21

Gulf of Maine 21

Station 28 23

Station 35 23

Discussion 23

Citations 27
Acknowledgments _

There are many persons who have helped or cooperated in the course of

this study in one way or another. All my thanks and appreciation to S. and

S. Fromm, R. Pikanowski, S. Chang, K. McNulty, D. Radosh, R. Reid,

A. Frame, 1. Lindsay, C. Steimle, J. Berrien, J. Rugg, F. Thurberg,

R. Terranova, R. Theroux, B. Higgins, and reviewers, and especially to

M. Cox for preparing the figures.

IV
Benthic Macrofauna and

Habitat Monitoring on the

Continental Shelf of the

Northeastern United States

I. Biomass

FRANK W. STEIMLE

Sandy Hook Laboratory

Northeast Fisheries Center

National Marine Fisheries Service, NOAA

Highlands, New Jersey 07732

ABSTRACT

Information on long-term temporal variability of and trends in

benthic community-structure variables, such as biomass, is

needed to estimate the range of normal variability in comparison

with the effects of environmental change or disturbance. Fishery

resource distribution and population growth will be influenced

by such variability. This study examines benthic macrofaunal

biomass and related data collected annually between 1978 and

1985 at 27 sites on the continental shelf of the northwestern

Atlantic, from North Carolina to the southern Gulf of Maine.

The study was expanded at several sites with data from other

studies collected at the same sites prior to 1978. Results indicate

that although there was interannual and seasonal variability,

as expected, biomass levels over the study period showed few

clear trends. Sites exhibiting trends were either in pollutionstressed coastal areas or influenced by the population dynamics

of one or a few species, especially echinoderms.

Introduction _

In the northeastern United States, the basic definition and

spatial distribution of benthic communities and dominant taxa

are becoming known (Pratt 1973, Wigley and Theroux 1981,

Theroux and Grosslein 1987); however, temporal variability

and trends have received less attention. There have been few

long-term (>3 years) benthic studies in the northwestern

Atlantic and fewer yet that have considered biomass as a

variable. Benthic community biomass and production have

long been of interest to fishery ecologists, e.g., Peterson

(1918), Blegvad (1928, 1951), and McIntyre (1978), particularly as they relate to the quantity of food available to

fishery species and as a primary component in ecosystem

models. See, for example, Cohen et a1. (1982), Lunz and

Kendall (1982), and Overholtz and Tyler (1986).

Biomass, as with other benthic community-structure variables, can also be used for environmental monitoring. For

example, biomass changes have been related to organic

enrichment, resulting from eutrophication or waste discharges (Pearson and Rosenberg 1978, Cedarwall and

Elmgren 1980). Biomass estimates are usually less affected

than other benthic community-structure variables, e.g.,

species richness or numbers of individuals, by sortingefficiency errors. This is because most biomass is from

larger, more common species (Coleman 1980, Cedarwall and

Elmgren 1980, Glemarec and Menesguen 1980). Biomass

estimates can also be influenced by other factors such as the

intermittent occurrence of rare, large individuals of certain

species. As noted by Coull (1985), long-term data sets may

be necessary, in themselves, to propose credible hypotheses.

Wolfe et al. (1987) and Lewin (1986) also noted that longterm studies are less likely to miss infrequent, but important, perturbations in the ecosystem.

This report summarizes and provides a preliminary

analysis of the range of temporal variability and possible

trends in wet weight biomass for 27 sites, well distributed

on the northeastern continental shelf of the United States.

These were monitored during 1978-85 as part of NOAA's

Northeast Monitoring Program (Reid et al. 1987). This report

also includes results from earlier studies at some ofthe same

sites. This long-term data set augments the extensive, earlier

spatial distribution studies of the area by Wigley and Theroux

(1981), Steimle (1985), and Theroux and Grosslein (1987).

Analysis of other community-structure variables (numerical

abundance, species composition, and richness) is being

prepared, with a more rigorous statistical analysis of habitat

relationships.
y.

72°

.}/... (

o

mlm-i~~ so 100 tSO 200

Kl.OMElERS

42

40°

38°

Figure 1

Location of benthic sampling sites on continental shelf, northeastern United States.

Methods _

The benthic samples used in this analysis were collected during quarterly and annual surveys between 1978 and 1985

(Reid et al. 1987); sampling locations are shown in Figure

1. All samples were collected with a 0.1-m2 SmithMcIntyre grab which provides for a 16-cm maximum

hernicylindrical penetration of the sediment. Five grab

samples were collected at each site per survey, except as footnoted in figure captions. Gaps in the data sets were the result

of samples not being collected or processed. Samples collected before December 1979 were sieved through 1.0-mm

mesh screens and thereafter through 0.5-mm screens. After

sieving, samples were fixed immediately in a 10% buffered

formalin-sea water solution and transferred within 3 days to

a 70% ethyl or isopropyl alcohol/5% glycerin preservative.

After storage for at least 6 months, samples were sorted,

specimens identified to species and enumerated, and each

taxon weighed on a electronic balance to I-mg accuracy,

following a 3-minute blot-drying on absorbent paper toweling. No attempt was made to adjust for weight changes

(possibly ±15 %) resulting from fixation and preservation

(Mills et al. 1982). It is assumed the 6-month delay in processing would make such change relatively consistent.

2

Biomass estimates for the standing stock of benthic organisms based on wet weight are subject to other potential errors

or biases. Many of these problems are due to the variable

contribution to biomass of water and inorganic material (shell

and ash), which can vary for each taxonomic group or

species. Other changes can be caused by the processing of

the samples, i.e., weight changes caused by the preservation media, as noted before, or losses to evaporation during

the weighing. Improvements can be made to the estimates

by using ash-free dry weight values or energy equivalents

for, at the least, dominant species or groups. These conversions can reduce some of the variability in the data because

water and inorganic components are mostly eliminated and

only the variable of primary importance, that related to the

quantity oforganic energy in the benthic sample, is presented.

This makes the benthic data readily applicable in developing energy-based models or budgets for use in fishery

ecology. Because the benthic samples used in this study were

to be archived, a nondestructive approach was taken, i.e.,

use of energy equivalents. The energy equivalency values

from Steimle and Terranova (1985) were used, with 1 Kj

= 0.239 Kcal.
The occurrence, in samples, of a few larger organisms can

obscure basic information or patterns in overall benthic

monitoring data (Buchanan et al. 1978). To address this, the

biomass of individual organisms weighing more than 1.0 g

and occurring in less than 50% of the grab samples in a collection were noted but excluded in the analysis of the data.

This also makes the data more relevant to prey-predator considerations since these large, megafaunal organisms are only

infrequently used as prey.

To evaluate the possible sources of variability in the data,

some factors that can substantially influence results or interpretation are presented: sediment grain size and total organic

carbon (TOC), station location precision, population dynamics ofdominant species, and general environmental or habitat

characteristics for each site, including common benthic

predators found at each site. The sediment TOC (total organic

carbon) data (mg/g dry weight) were determined for 1978-80

samples by chromate oxidation, for 1981-82 samples by

combustion in a furnace, and for 1983 samples by infrared

analysis. The sediment PCB (polychlorinated biphenyl) and

PAH (polycyclic aromatic hydrocarbon) data were from

Boehm (1983), the mean sediment trace-metal data from Reid

et al. (1987); these data are dry weight values. The hydrography was from Ingham (1982), and sediment topography

(i.e., ridge and swale) was based on Swift (1975). Data on

potential benthic predators common at each site were drawn

from records of concurrent trawl collections made at the same

sites as the benthic samples; these records contained notes

on species collected and relative abundance. Stomach contents were examined from a representative sampling of

demersal fish species and lobsters in the trawl collections.

During the study, it became readily apparent that biomass

levels or trends at several sites appeared to be highly influenced by the population dynamics of one or a few species,

especially echinoderms, a factor noted in other benthic studies

(Warwick 1980). To evaluate this factor, the echinoderms

at several sites, where they dominated the biomass, were

measured to provide size-frequency distributions. These

distributions can characterize the structure (number of

cohorts and approximate ages) and dynamics (frequency of

recruitment and approximate growth rates) of populations.

In the case of the sand dollar Echinarachnius parma, this

led to a separate study which included production estimates

(Steimle 1990).

An analysis ofthe significance ofthe change in December

1979 ofsieve mesh size from 1.0 to 0.5 mm was conducted

for two surveys, January and July 1982, by sieving each grab

sample through both screen sizes.

Many of the sites monitored during the 1978-85 period

were originally selected because there was information

available from earlier studies, e.g., at stations 15A (R. Swartz

and F. Cole, U.S. EPA, Corvallis, OR 97330), 18 (Steimle

1982), 16A and B (Steimle 1985), 28 (Maurer 1982), and

three sites from Wigley and Theroux (1981) and Theroux

and Grosslein (1987), that were within a kilometer of stations 8, 22, and 32. The biomass data from these studies have

been included in the results and any differences in methods

3

or location noted. The biomass data are presented as line

graphs instead of histograms, to enhance the definition of

possible long-term patterns. It is understood that this makes

assumptions about the biomass dynamics of the community

between collections, which can be separated by as much as

a year. The graphs were interrupted ifthere were intercollection gaps of more than a year. The results of two studies

that collected benthic biomass data bimonthly at stations 31

and 32 (Howe and Leathem 1984) and near stations 16A-C

(Steimle 1990) show that biomass levels at a site can be either

relatively stable at the bimonthly sampling intensity, disregarding an odd pulse of added biomass, or highly variable.

Because the original sampling design was not planned

specifically for biomass analysis, and considering the

statistical intractability of benthic data from small sample

sizes, detailed statistical analysis was considered unwarranted

or of dubious value. Linear regressions were calculated,

however, for the mean biomass-over-time data for stations

with an apparent trend, assuming equal seasonal contributions, and then tested for a significant difference of the slope

from zero.

Results _

Analysis of the significance of change in screen size from

1.0 to 0.5 mm in December 1979 (Table 1) indicates the

0.5-rom screen contributed, on average, an additional 4%

to total biomass, although this did go as high as 13 %. This

low contribution is consistent with the results of previous

studies on the importance of screen size to biomass, e.g.,

Reish (1959). Thus, differences in screen size are not considered a major factor in evaluating biomass variability.

A comparison of the mean summer-to-winter biomass

values at each site during 1978-83, when seasonal collections were available, shows a majority (20/27) of the sites

did have higher mean summer values (Table 2). In general,

the mean summer biomass levels averaged about 30% higher

than the winter levels (Table 2). Energy equivalents showed

about the same average summer increase (28 %).

The biomass results for each site (Figs. 2-28) show minor

to considerable long-term variability. The results are presented below in regional groups of stations along with

available and relevant environmental information. This

approach is used because each station can be considered

unique, to some degree, due to substantial spatial isolation;

thus individual treatment is warranted. The error bars for

each mean biomass value on Figures 2-28 represent one

standard deviation (SD), as indicated on the vertical axis

legend. For economy of space, some wide error bars are

broken with the actual SD value given in parentheses at the

top of the bar.
Table 1

Significance to total benthic macrofaunal biomass of using O.5-mm

sieve mesh compared with using l.o-mm mesh for winter (January)

and summer (July) 1982 collections. This is based on biomass

retained on the O.5-mm mesh that passed through the l.o-mm mesh,

percent of total biomass this represented, and dominant taxa

retained on the O.5-mm sieve.

Winter Summer

Site Biomass Dominant Biomass Dominant

no. (g/m2

) % Total taxa (g/m2

) % Total taxa

2 0.70 2.36 P,A 2.00 8.20 P.B

3 0.46 1.57 P 3.68 7.39 P,B

7 3.84 4.71 P 1.86 1.65 P,A

8 1.19 3.70 P 0.82 1.39 P

9 1.94 3.43 P 1.29 2.13 P

11 1.71 1.01 P,A,B 0.86 1.33 P

12 2.79 2.58 P,A 1.46 1.99 P,A

13 2.26 1.31 A,P 1.40 0.04 P

15A 1.44 0.63 P,A 1.90 1.11 AP

16A 1.27 2.29 P 2.58 1.68 P

16B 0.28 2.90 B 2.04 6.02 P

16C 4.91 11.88 P,B 0.88 1.87 P

17 0.41 0.10 P 0.62 0.04 P

18 8.55 4.10 A,B 10.94 2.14 A,B,P

19 2.27 3.64 P,A 1.17 0.37 P,A

20 0.74 1.43 P,O 5.44 10.47 0

22 1.83 9.12 A,P 1.61 4.30 P,A

23 1.17 1.52 A,P 3.97 1.03 A

24 0.13 5.40 P 0.13 4.29 P,A

28 0.70 0.97 P,O

33 7.32 6.48 P,A 4.18 7.29 P

34 4.25 6.27 B 3.18 2.03 B,P

35 3.56 3.84 P 23.22 13.07 P

36 3.13 5.11 B 2.84 3.96 B,P

37 2.45 6.88 A,P 3.87 12.26 P

x 2.37 3.73 3.41 4.00

SD 2.12 2.81 4.77 3.86

P polychaete, A amphipod, 8 bivalve, 0 ophiuroid

Chesapeake Bight

The southernmost area, the Chesapeake Bight, contained

seven stations: 2, 3, 7, 8, 9, 31, and 32 (Fig. 1). Data at

stations 2,31, and 32 were discontinuous; however, the other

stations had biomass levels that ranged mostly between 20

and 120 g/m2, with polychaetes, molluscs, and echinoderms

the dominant groups.

Station 2 This station was located off Virginia Beach, VA

(Fig. 1) and actually included two sites. The initial site was

abandoned after a year because of a navigational safety

problem; the second site (lat. 36°49.8'N, long. 75°50.3'W)

was 5 km south of the original. Both sites were about 15 m

deep with medium sand at the initial site and fine to siltyfine sand at the fmal site. Both also had relatively low levels

of trace metals, e.g., <10 ppm dry weight (DW) lead. The

sites were also in the coastal-active hydrographic band that

can be influenced by the Chesapeake Bay plume and shore4

Table 2

Comparison of mean benthic biomass (g/m2

) for 1978-83 summer

and winter collections at each site, estimated energy values

(Kcal/m2

), and mean Kcal:B ratios.

Biomass Energy Kcal:B

Site

no. Summer Winter Summer Winter Summer Winter

2 40.5 14.0 27.5 11.7 0.68 0.84

3 43.3 24.3 34.0 20.0 0.78 0.82

7 74.7 78.0 57.3 55.7 0.77 0.71

8 50.0 34.3 33.0 24.7 0.66 0.72

9 75.7 51.1 60.0 40.0 0.79 0.86

11 90.0 183.3 65.7 126.7 0.73 0.69

12 63.3 70.3 55.0 58.0 0.87 0.83

13 247.7 179.7 184.7 142.7 0.75 0.79

15A 91.0 249.3 64.3 168.7 0.71 0.68

16A 79.7 46.0 76.3 41.0 0.96 0.89

16B 33.0 41.0 34.3 37.3 1.04 0.91

16C 52.7 22.7 46.7 20.0 0.89 0.88

17 605.7 329.0 399.0 218.3 0.66 0.66

18 191.0 167.3 165.0 171.3 0.86 1.02

19 252.0 64.0 197.3 62.0 0.78 0.97

20 83.0 64.0 59.7 44.3 0.72 0.69

22 38.4 21.5 40.6 22.8 1.06 1.06

23 62.0 37.8 56.6 35.3 0.91 0.93

24 24.6 4.5 20.0 4.3 0.81 0.94

28 55.0 45.3 41.0 34.7 0.75 0.77

31 21.5 8.0 17.5 7.0 0.81 0.88

32 37.0 14.0 27.0 13.0 0.73 0.93

33 105.0 122.7 97.7 115.7 0.93 0.94

34 142.7 97.7 100.7 73.0 0.71 0.75

35 154.7 80.0 137.7 70.7 0.89 0.88

36 74.7 79.0 55.0 62.5 0.74 0.79

37 44.3 37.0 40.7 36.5 0.92 0.99

x 104.9 80.2 81.3 63.6 0.81 0.84

zone upwellings. The few collections made at the initial site

(Fig. 2a) are not reliable in any discussion of trends. There

was less than an order of magnitude in mean biomass variability at the second site; this appeared to be related to

seasonal factors. The species that generally dominated the

biomass were the polychaete Glycera americana, and the

molluscs Pandora gouldiana, Tellina agilis, Ensis directus,

and Ilynassa trivittata. Trawls made at both sites indicated

the benthic predators present were weakfish Cynoscion

regalis, croaker Micropogon undulatus, young summer

flounder Paralichthys dentatus, northern searobin Prionotus

carolinus, windowpane Scophthalmus aquosus, and spotted

hake Urophycis regia. Various amphipods, decapods Cancer

sp., Ovalipes sp., Crangon septemspinosus), and the razor

clam Ensis directus, were the benthic prey commonly eaten,

although nonbenthic mysid shrimp and small fish were most

important to some species.

Station 3 This station was located south of the previous

station, offthe Virginia-North Carolina border (lat. 36°34'N,

long. 75°48'W, Fig. 1), near a coastal, shoal-retreat massif

in 20 m with fine-sand sediments which contained <0.4 ppb

PCBs, 12 ppt PAHs, and low levels of trace metals. The
Figure 2

Benthic macrofauna! biomass levels at

station 2. (a) Only four grab samples

available; (b) residua! biomass was

mostly tunicates; (c) one 9.9-g Pagurus

sp. excluded; (d) severa! Encope emarginata (12.0 g) excluded; and (e) one

Mellita quinquiesperforata (8.4 g) and

one Eupleura caudata (1.2 g) excluded•

e

• POLYCHAETES

MOLLUSCS

lIlIII CRUSTACEANS

[] ECHINODERMS

o Residual

d

2/82 8/82 12/82 7/83

®

a

12/79 7/8011/80 7/84 nr--...... t-:-.-"'..-_-..-:.._-.~--'l-.;->-J.1..------....L:-.~-.~-~~-:LI.-~-~ .•..L~----l..---_1]

IX 0 1J21

,...

C\l '3 E .

c, (J) 100 ...... ,.. til + til ......

E

Cll -

o Cl

iIi~ c: _

Cll Q)

~~

,...

C\J '3 E . c,(J)

...... ,.. til + til ......

E

Cll -

o Cl

iIi~ c:

Cll Q)

~ ~

100

o

• POLYCHAETES

MOLLUSCS

lIlIII CRUSTACEANS

[] ECHINODERMS

o Residual

10n9 7/80 11/80 7/81 2/82 8/82 12/82 7/83 7/84

~e· ~EClCl 50~L.---------·_-.l._·_-··_--_-_-_·-'-"·-_·_---'- __---J -J~05 ... N. : j :~.-.~~~~.-.~:~.-.~:~.-.~~~~.-.:~.-----~:. ~~.~~:~~.~~: t 'oJ

IX 1J21

Figure 3

Benthic macrofauna! biomass levels

at station 3. (a) Only two grab

samples available; (b) one Neverita

duplicata (6.8 g) excluded; and (c)

mostly E. parma.

site was also in a coastal-active band, but only marginally

influenced by the Chesapeake plume. The mean macrofaunal

biomass levels were relatively stable and ranged from 16.3

to 82.2 g/m2 , with seasonal fluctuations apparently responsible for most of the 1978-82 variability (Fig. 3). Biomass

was dominated by the polychaetes Glyeera dibranehiata and

Spiophanes bombyx, and the molluscs Ensis direetus and llynassa trivittata. An expansion of the Atlantic jackknife E.

direetus population was the source of the 1982-83 biomass

increase. As with the previous station, potentially important

benthic predators were windowpane, summer flounder, and

spotted hake. These predators ate predominately Ovalipes

crabs, Crangon shrimp, jackknife clam (Ensis), mysids, or

small fish.

5

Station 7 This station (lat. 36°47.5'N, long. 75°11.7'W)

was on the continental shelf off Virginia (Fig. 1) in 33 m

of water. Sediments were medium sand in a ridge and swale

area, with low levels of trace metals. Hydrography of the

site may be influenced by the development of seasonal,

coastal fronts. The mean macrofaunal biomass at this site

ranged from 19.4 to 114.4 g/m2, without any clear trends

but with an increase between 1980 and 1982 caused by increases in several groups (Fig. 4). Species making major contributions to the biomass were the polychaetes Nephtys pieta

and Spiophanes bombyx, the molluscs Ensis direetus and

Tellina agiUs, and the sand dollar Eehinaraehnius parma.

The influence ofseasonal factors was not evident. Potentially

important benthic predators collected here were the northern
200

......

C\I 'ci E . o,C/)

.......... (/) + (/) .......

tll -

100

iii

c:

Q) Q) tll -

:E

• POLYCHAETES

Il§ MOLLUSCS

ITIlIl CRUSTACEANS

El ECHINODERMS

o Residual

5' " , ; ,

~~o Cl j A--------..---t-------t------:------: .' . . . . -- A .... E 0 . .

IX

12/79 7/8011/80 7/81 2/82 8/8212/82 7/83 7/84

Figure 4

Benthic macrofauna! biomass levels at

station 7. (a) Only four grab samples

available and one AsteriDsforbesi (12.2

g) excluded•

Figure 5

Benthic macrofauna! biomass levels at station 8. (a) Data from a single grab; and (b)

residua! biomass mostly the anthozoan Epizoanthus sp.

7/83

• POLYCHAETES

Il§ MOLLUSCS

ITIlIl CRUSTACEANS

[] ECHINODERMS

o Residual

6/64 12/79 7/8011/80 7/81 2/82 8/82 12/82

.. 5

j r5 ....... (!)Cn ....

g.g: A A···t.. · ·t A t A .... 1; 0 L __----'.=----_-_-_-_--_-....;.:.~_~_~_-_-_--_-_- __--_-_-_-'_.:.=:._:-:..--.--.;-:.-----.------' 0 I.¢

......

N 'ci E . o,C/)

..........

(/) + 100 (/) .......

tll -

g

iii Q)

c:

tll a;

~~

searobin, scup Stenotomus chrysops, black sea bass Centropristis striata, spotted hake, red hake Urophysis chuss, and

horseshoe (Limulus) and calico (Ovalipes) crabs. The rock

crab Cancer irroratus was a common prey ofthe fish, along

with various polychaetes and amphipods.

Station 8 This station (lat. 36°40.7'N, long. 74°45.l'W)

was on the outer shelf, 80 m in depth, off Virginia (Fig. 1)

in a ridge and swale area, with medium sand and low trace

metals. The site was possibly influenced by the shelf-slope

water-mass front and the "cold pool, " as defined by Ingham

(1982). Dominant species contributing to the biomass were

the bivalves Astarte spp., the sea star Astropecten americana,

6
Figure 6

Benthic macrofaunal biomass levels at station 9. (a) This collection might have been

offstation by 2 km; (b) several Arctica islandica (240.5 g) excluded; (c) several AstaT1e sp.

(21.4 g), Buccinium undaJum (49.8 g),

Aphrodite hastala (10.0 g), and A. islandica

(11.9 g) excluded; (d) one A. islandica (23.0

g) and one AstaT1e sp. (10.3 g) excluded; and

(e) one AstaT1e castanea (10.1 g) excluded.

• POLYCHAETES

ffil MOLLUSCS

IIlIIJ CRUSTACEANS

[] ECHINODERMS

o Residual

a

200

o

7/8011180 7/81 2/82 8/8212/82 7/83 7/84

~15j t;;;'!"",,"Ct""C"t:::cctc'" 50!

IX 0 '-- -=- --...J~ 1%

the polychaete Onuphis sp., and the anemone Epizoanthus

sp. The mean biomass levels were relatively constant over

the study period, ranging from 28.0 to 59.8 g/m2 , with

some suggestion ofseasonal fluctuations (Fig. 5). Potentially important predators found at the site were the spotted hake,

sea robins, scup, Cancer crabs, and others to a lesser extent.

Amphipods Ampelisca agassizi and Unciola sp., polychaetes

Lumbrineris sp. and Ophelia denticulata, and decapods

Cancer and Crangon were the most common prey eaten by

the fish.

Station 9 This station (lat. 38°17.3'N, long. 74°17.7'W)

was in a former sewage-sludge disposal area (Fig. 1), used

from 1973 to 1980. Sediments were medium sand in a ridge

and swale area where <2.1 ppb PCBs, 26 ppt PARs, and

low levels of trace metals were measured. The site was 50 m

deep and could have been in the region influenced by the

"cold pool." Biomass was primarily dominated by the sand

dollar Echinarachnius parma and several megafaunal molluscs (Arctica islandica and Astarte spp.) that were excluded

from the data summarized in Figure 6. Mean biomass levels

were fairly constant, ranging from 48.5 to 70.9 g/m2, with

some seasonal influence evident to 1982. Potentially important predators at the site were the little skate Raja erinacea,

windowpane, red hake, Cancer crabs, and other less frequently occurring species. Polychaetes, as well as Cancer

and Crangon, were important prey.

Station 31 This station (lat. 38°44.8'N, long. 75°01'W)

was just below the mouth of Delaware Bay (Fig. 1), 25 m

deep, and in the coastal-active band influenced by the Bay

plume and coastal upwellings. Some summer hypoxia has

been reported in the area. It was in a coastal ridge and swale

area and had medium-to-fine sandy sediments with TOC

levels more variable than usually found (Fig. 7). There were

moderate trace-metal levels, e.g., 6-17 ppm lead, as well.

The species that dominated the biomass were mostly molluscs, including Ensis directus, Nucula annulata, Tellina

agiUs, and Ilynassa trivittata, with patches of blue mussel

Mytilus edulis in 1980. Mean biomass levels for this station

were irregular with an order of magnitude change between

1980-81 (7.6-132 g/m2) and 1983-85 (72.9-1535.3 g/m2)

collection periods (Fig. 7). This change was primarily

because of population dynamics ofthe small nut clam Nucula

annulata that often contributed over 1 kilogram of biomass

per grab sample. The small predatory mudsnail I. trivittata

became abundant in 1984, possibly in response to this large

forage base and perhaps a reasonable explanation for some

ofthe overall decline in biomass levels ofNucula thereafter.

Other, larger, potentially important benthic predators collected at the site included windowpane, silver hake Merluccius bilinearis, spotted hake, horseshoe crabs, and several

less common species. Benthic prey commonly eaten by the

fish were Cancer, Crangon, amphipods (e.g., Unciola sp.),

and polychaetes such as Pherusa affinis.

7
100

.....

C\lE q o,UJ,.... en + en~

«l E -

.s::.

o

iiic: _ 50

«l Q)

~3:

a

b

• POLYCHAETES

MOLLUSCS

[]llI] CRUSTACEANS

El ECHINODERMS 2000

o Residual

1000

10/79 3/807/8011/804/81 9/81 2/82 7/8212/82 7/8312/83

o 0

7/8411/84 6/85

..... 3: 20,.---L._ ___._LU...J---,a----~___I...L_.L..L..l..._l'___ __'_____L.__ ___'__ __'_________'__..L____'____,_1 0

.0 N ~~ W

U'

O~

I- E 0.1..---....-.:...:....;.,;--==---------------=""-------------.......00 IX I~

Figure 7

Benthic macrofaunal biomass levels at station 31. (a) 1980-81 data from Howe and Leathem (1984); and (b) one holothurian

Sclerodactyla briarus (51.8 g) excluded. Note change in scale after 1982.

Station 32 This station (lat. 38°31.4'N, long. 74°7.7'W)

was farther below the mouth of Delaware Bay than the previous site (Fig. 1) and was shallower, 15 m, but with very

similar hydrographic characteristics. It was also in a coastal

ridge and swale zone with coarse-to-medium sands and low

trace-metal levels. The dominant species ofthe biomass were

the bivalve molluscs Spisula solidissma and Tellina agilis and

a mix of polychaete species. Mean biomass levels were low

but fairly stable in the early collections (3.9-15.3 g/m2).

This was punctuated by an odd pulse (52.8 g/m2) of E.

parma in July 1980 (Fig. 8), possibly reflecting some sampling error. The biomass was more variable (9.3-57.7

g/m2) in later collections, but summer increases were clear.

Commonly collected benthic predators included the spotted

hake, a variety of skates and rays, sea robins, young summer

flounder, horseshoe, and other crabs. Decapods, especially

Cancer, Crangon, and Ovalipes, were the most common prey

found in the fish stomachs examined.

New York Bight

The next set of nine stations was in the New York Bight with

stations 11, 12, and 13 on the shelf, stations 17 and 15A in

coastal waters, and stations 33 and 16A-C in the submerged

8

Hudson Shelf Valley (Fig. 1). On the shelf, the mean biomass

levels usually ranged between 50 and 200 g/m2, highly

variable at the coastal sites (because of sand dollar populations), and about 100 g/m2 in the Hudson Shelf Valley.

Station 11 This station (lat. 38°44.6'N, long. 74°02.l'W)

was midshelf off Delaware (Fig. I) in 50 m of water, with

fine-sand sediments and low trace metals, in an area that can

be affected by the "cold pool." During the study period,

mean biomass levels of the site were highly affected by

population dynamics of Echinarachnius parma (Fig. 9),

which will be discussed later. The mollusc biomass was due

partly to the occurrence, in several collections, of Cyclocardia borealis and Astarte spp. Potentially important predators

occurring at the site included four-spot flounder Paralichthys

oblongus, little skate, yellowtail flounder Limanda ferruginea, windowpane, black sea bass, sea robins, red and

spotted hake, Cancer crabs, and a variety of less common

species. Common fish prey were Cancer crabs, Crangon,

polychaetes, amphipods, e.g., Ampelisca sp., and the rhynchocoel Cerebratulus.
0..1..-.........--"'--

I

b a

I

"C

Q)

N

>.

'iii

c

til

oc

I

• POLYCHAETES

MOLLUSCS

IT!llI CRUSTACEANS

[] ECHINODERMS

o Residual "

(\J 3 E .

Olen

'-' ,... II) + II) '-'

tll - E .c o .21 50

iii~ c: _

tll OJ

10/79 3/80 7/8012/804/81 9/81 2/82 8/8212/82 7/83 12/83 7/8411/84 6/85

;ge~E~ 5ol_-=-__---=':.:!!'~':..=' ' _'_'_'..:..'--==='- ---"1

5

0

'V OJ' l __ ····A ··.A A t :(!)~. _ .' ------e -- e------- --'tf-_ ----e---

IX I~

Figure 8

Benthic macrofaunal biomass levels at station 32. (a) 1980-81 data from Howe and Leathem (1984); and (b) one

Neverita duplicata (28.4 g) excluded.

400

• POLYCHAETES

MOLLUSCS

" 300 IT!llI CRUSTACEANS (\j a

E q [] ECHINODERMS

'-'

Olen,... o Residual

II)

II) '-' + tll 200

E .cCl C 0 'w iii

c:

(\l -OJ OJ 100

o..I..-...----......---f"I12/79 7/8011/80 7/81 2/82 8/8212/82 7/83 7/84

Figure 9

Benthic macrofaunal biomass levels at station 11. (a) This collection might have been otT station by 2 km and

includes only three grab samples; (b) one Cancersp. (66.0 g) excluded; (c) one Arctica islandica (107.0 g) excluded;

and (d) one Aphrodite hastata (2.1 g) and an unidentified bivalve (4.9 g) excluded.

9
200

.s:::.

-

Cl

'03 100 3:

-OJ

3:

......

N 3 E .

t5l(J)

'" .... en + en '" tll

E

o

iii

c:

tll

OJ

400

...... ...... C\l

E Q 300

0, (J)

'" .... en + en

tll '"

E .s:::.

-

Cl 0 '03 200 iii 3: c:

OJ

tll -OJ

3:

100

• POLYCHAETES

MOLLUSCS

[J]]ll CRUSTACEANS

[] ECHINODERMS

o Residual

12/79

a

(218)

1

b

d

Figure 10

Benthic macrofaunal biomass levels at station

12. (a) One Cancer irroralus (3.2 g) excluded;

(b) one Astropecten amculaJa (2.9 g) excluded;

(c) one Astarte castanea (12.2 g) excluded, the

high variance due to many small Cyclocardia

borealis in one sample; and (d) one Arctica islondica (11.5 g) excluded, high variance as above.

• POLYCHAETES

MOLLUSCS

[J]]ll CRUSTACEANS

[] ECHINODERMS

o Residual

10/79 7/8011/80 7/81 2/82 8/82 11/82 7/84

Figure 11

Benthic macrofaunal biomass

levels at station 13. (a) Only

three grab samples available;

and (b) high variance because

of several small Arctica islondka and Cyclocardia borealis in

two samples; the mean

biomass of the remaining three

samples was 210 g·m2•

10
Station 12 This station (lat. 38°6.3'N, long. 73°0.7'W)

was on the outer shelf off Delaware (Fig. 1) in 70 m, with

medium sands containing low levels oftrace metals. It could

also have been affected by the "cold pool" and the shelfslope water-mass front. Figure 10 shows polychaetes generally important to the overall mean biomass (with no particular

species dominant), as well as molluscs, mainly Astarte spp.

and Cyclocardia. Predators collected here were four-spot

flounder, red hake, scup, little skate, sea robins, Cancer

crabs, and less common species. The fish ate various polychaetes, including Ninoe sp., along with ampeliscid amphipods and Cancer.

Station 13 This station (lat. 39°20A'N, long. 72°58.9'W)

was in an area on the outer shelf (Fig. 1) and was monitored

because it lies in the subsurface "Baltimore Canyon

Trough", considered to have potential oil or gas reserves.

It was 65 m deep with low trace metals, medium sand, in

or near the "cold pool" and the shelf-slope water-mass front.

Molluscs dominated this site (Fig. 11), again mainly Astarte

spp. and Cyclocardia. Although not an overall major contributor, the amphipod Ampelisca agassizi was conspicuous

in the samples. With the exception of the last two collections (the first biased by a high biomass cluster of the

dominant bivalves in two grabs), the mean biomass was

relatively consistent, averaging just under 200 g/m2 . Potentially important predators collected here were basically the

same as at station 12, but also included yellowtail flounder

and silver hake. Decapods (Cancer, Crangon, and Dichelopandalus) were important food for the hakes and four-spot

flounder, while polychaetes Scalibregma and Lumbrineris

and amphipods Ampelisca and Unciola were important to

yellowtail and scup.

Station 17 This station (lat. 39°35.8'N, long. 73°54.2'W)

was in the area off New Jersey (Fig. 1) defaunated by an

exceptional anoxia event in 1976 (Steimle and Radosh 1979).

The depth was 20 m with medium-to-coarse sand and low

trace metals, in a coastal ridge and swale area, possibly influenced by upwellings and seasonal fronts. The highly

(281)

I

(350)

J

1::

Cl 'iii

;:: 200

N'"' E q c,C/)

-;;; +300 en ....... til

E

o

iii

c:

til Qj

;::

400

500

100

• POLYCHAETES

ml MOLLUSCS

[]lj]] CRUSTACEANS

El ECHINODERMS

o Residual (330)

I

o 12/79 718011/80 7181 2/82 8/8212/82 7183 8/84 6/85 9/85 '"'

'"'0

0

: ...... ;:::= 5j 5<ll

IX~: 0 __..;;.. ... ..~~::....~~: .•_:"-'...

.~~·J·~ ..;-··.l:-.":~-L·.:t::.·.:~~::.:.-· --"""'e-_=-_--=-_._...._.."'......... ·-·-·-·-----------....J~O~· t .:

IJZf

Figure 12

Benthic macrofaunal biomass levels at station 17. (a) Seven Spisula solidissima (148.0 g) from one sample excluded; and

(b) only two grab samples analyzed.

11
• POLYCHAETES

lI§ MOLLUSCS

rnm CRUSTACEANS

[] ECHINODERMS

o Residual

200

......

C\l

E q c,f/)

...... .... (fl

(fl ...... +

as

E .r:.

-

Cl a 0 'Qj iii 3: 100 c:

-as b Q) Q) 3:

o

3: 12/79 7/80 11/80 7/81 2/82 8/8212/82 7/83 8/84 8/85 9/85 ......

..Cl 1OJ -------- .---~ .- / .------~ 1

1

0 ~. 00 .g; .. ... «... . . . . ..... . . ..... . . . . . .... v, f-I~ 0 '--_-_.._.._._.._._.._._._._..... z - 0 CJ

1J/1

Figure 13

Benthic macrofaunal biomass levels at station 33. (a) Two Pilar morrhuanus (28.3 g) and one Asterias vulgarus (49 g)

excluded; (b) two Havelockia scabra (6.4 g) excluded; (c) one Arctica islandica (66.5 g) excluded; and (d) only two grab

samples analyzed.

variable biomass at this site was dominated by Echinarachnius parma (Fig. 12). Major predators were black sea bass,

yellowtail, four-spot flounder, windowpane, sea robins, scup,

and spotted hake. Benthic prey found in their stomachs included the tentacular crowns of the burrowing anemone

Ceriantheopsis americanus, some Crangon, Cancer, a variety of amphipods, polychaetes, and the ribbon worm

Cerebratulus, but not E. parma.

Station 33 This station (lat. 40 001.4'N, long. 73°25.6'W)

was in the Hudson Shelf Valley (Fig. 1) that traverses the

shelf from the mouth of New York Harbor to the Hudson

Canyon on the slope. The site was 62 m deep with silty, fine

sands possibly reflecting a depositional environment. Boehm

(1983) found low levels (6-9 ppb) of PCBs and moderate

levels (about 490 ppt) of PAHs in these sediments. This

possibly reflects some downvalley transport from the highly

contaminated waste-disposal areas just outside the Harbor

(Boehm 1983). This site could be influenced by intrusions

of outer shelf waters. The mean biomass here (Fig. 13) was

dominated by polychaetes, with Nephtys incisa conspicuous;

the amphipod Ampelisca agassizi and several molluscs were

also important. The overall biomass levels appear relatively

stable (57.9-142.0 g/m2), although there may have been a

decline since 1981 (Fig. 13). A variety of predators were

commonly found, including little skate; four-spot and yellowtail flounders, and winter flounder Pseudopleuronectes

americanus; red and silver hake; scup; windowpane; ocean

pout Macrozoarces americanus; Cancer crabs; and American

lobster Homarus americanus. These predators ate a variety

of benthos, particularly Dichelopandalus by the hakes and

four-spot flounder; the nut clam Nucula sp. by lobsters;

and polychaetes Pherusa and Lumbrineris, ampeliscid

amphipods, and Ceriantheopsis by winter and yellowtail

flounders.

Station 16A This station (lat. 40 0

25'N, long. 73°44'W)

was at the northwest edge of a sewage-sludge disposal site

on a shoulder of the upper Hudson Shelf Valley (Fig. 1).

The site contained medium sand in about 25 m of water. PCB

values of about 2.2 ppm have been measured in the sediments, as well as relatively high trace-metal levels, e.g., lead

above 25 ppm. Besides waste disposal, the site can be influenced by the Hudson-Raritan plume and occasional seasonal

hypoxia. Mean biomass was relatively low «50 g/m2) and

consisted mostly of polychaetes (Fig. 14), including a mix

12
200.,---------------------------------------,

8/84 6/85 9/85

o 8/73 12/79 7/80 11/80 7/81 2/82 8/82 11/82 7/83 11/83

..... ~10C' I I I I I I I I

.!,OJ: ~~~-_.---. 0......... • ....----. ' ,

I~ 0 ,~:.-· ..:":"·~·:·-·-.:t·,. .. .. :"~~· ····· ~'>.....

.....

C\j

E q

...., oW..- l/J + l/J ...., til :E 100 E OJ 0 '(jj iii

c:

til a; (l)

::iE a

• POLYCHAETES

MOLLUSCS

IllIIl CRUSTACEANS

ffi] ECHINODERMS

o Residual

Figure 14

Benthic macrofaunal biomass levels at station 16A. (a) Only four grab samples available; (b) residual biomass was mostly anthozoans

and rhynchocoels; (c) one Pilar morrhuana (4.6 g) excluded; and (d) only two grab samples analyzed.

of common sandy-habitat species, e.g., Nephtys picta,

ubiquitous types such as Spiophanes bombyx, and the stresstolerant Capitella capitata. The burrowing anemone Ceriantheopsis americana and the rhynchocoel Cerebratulus were

other major contributors. The aberrant value for August 1982

was mostly that of the easily fragmented, large, predatory

rhynchocoel. Other predators found at this site were essentially the same as for station 33 with the major prey consisting of Pherusa, Ceriantheopsis, and Dichelopandalus.

Station 16B This station (lat. 40 025'N, long. 73°46'W)

was in the Christiaensen Basin of the upper Hudson Shelf

Valley (Fig. 1). The site was 27 m deep and in silty-fine sands

containing>150 ppb PCBs, >7000 ppt PAHs, and very high

levels of trace metals, e.g., up to 175 ppm lead. It was also

in the coastal-active band, influenced by the Hudson-Raritan

plume and seasonally recurrent hypoxia. Polychaetes were

again the dominant taxon (Fig. 15). There appeared to be

two basic community groups which dominated the biomass:

a Capitella-Cerebratulus group and a CeriantheopsisNephtys incisa-Nucula proxima group. These two groups

could alternately dominate the biomass in separate grabs in

a collection. Potentially important predators were the same

as at stations 16A and 17, and predation on many benthic

species has been previously reported (Steimle 1985). The

relatively low recent biomass levels here suggest a declining trend since 1973; however, most of the 1973 biomass

was that ofthe nut clam, Nucula proxima. The sediment TOC

13

levels were irregular, but this seems to have had little effect

on biomass levels.

Station 16C This station (lat. 400 25'N, long. 73°52'W)

was located north of the New York Harbor dredge-spoil

disposal area (Fig. 1). The site contained mixed sands in

22 m depths of a coastal ridge and swale area possibly influenced by the distribution of dredge spoils during or after

disposal. Less than 0.4 ppm PCBs were measured in these

sediments, but trace-metal levels were relatively high, similar

to station 16A. The site was also in the coastal-active band

influenced by the Hudson-Raritan plume, upwellings, and

seasonally recurrent hypoxia. The benthos had relatively low,

mean biomass (Fig. 16) but with more diversity, e.g., a mix

of polychaetes and the northern dwarf tellin Tellina agilis

being the major contributors. Overall biomass levels suggest a gradual increase during 1980-82, but a decline since.

The peak in biomass in 1980 (Fig. 16) appeared to be related

to a concurrent peak in sediment TOC, the cause of which

is unknown but could be a manifestation of some sampling

error. Potential predators were similar to those found at the

other stations in the area.
200

.t=-

Ol

'Qj

5: 100

o

(135)

J

b

• POLYCHAETES

IHJ MOLLUSCS

!IIIll CRUSTACEANS

[] ECHINODERMS

o Residual

c

8/73 12/79 7/8011/80 7/81 2/82 9/8212/82 7/8312/83 8/84 6/859/65 .....

~~25C~~---~~--~~~-~-=-=-=-~·-~-~--~-~--~---~-~~10~ """'" ", '" ,........... N

o ! --------., ", 'A' £ ~", ,'~ ~~":. en

0 r----.'-'-''-'-'.._-....-.~-'+...::. :...._'<-._.-..-·------·....:·....:'--------------------'-0 cj IX IJ21

Figure 15

Benthic macrofaunal biomass levels at station 16B. (a) Only four grab samples available; (b) most residual biomass was anthozoans,

rhynchocoels, and phoronids; and (c) only two grab samples analyzed•

100

a b

• POLYCHAETES

IHJ MOLLUSCS

!IIIll CRUSTACEANS

[] ECHINODERMS

o Residual

d

8/73 12/79 7/80 11/80 7/81 2/82 8/8212/82 7/83 8/84 6/85 9/85 .....

~~. ~~0l10C:: I ,/'/'\,: : ,~~~~ I I I 1100 (!):~. <~~ , \ .• ~-:-~ '. "'-1 t;' ...:~::.:.:~:. .,;' - ·~~t~~--· .. IX 0 f----''------....s==-==---------------------------

1J21

Figure 16

Benthic macrofaunal biomass levels at station 16C. (a) Only four grab samples available; (b) one Cancer borealis (91.0 g) excluded;

(c) one Euspira heros (1.3 g) excluded, and only two grab samples analyzed.

14
500 (171) • POLYCHAETES I MOLLUSCS

[J]]]J CRUSTACEANS

400 [] ECHINODERMS

~~ b o Residual

E q o,w (243»):

........... 300

C/l

C/l ......

+

ell - E~o .- c .- Q)

[IJ 3: c: 200 ell a; Q) 3: :iE

100

o 12/7 5/73 12/73 12/79 7/8011/80 7/81 2/82 8/8212/82 6/859/85

:!:~ 5j 2 ': I I I I I ': !5

0

:C): UO~ , , >r :PPPPt....t········· ...···········~~~·~ ....~~·~:~·:~~~~· ... I-I~ 0 /I~._....t:!.::-=.:--::-~--=.::-:.:-.:J.~-:.:-.:!-~-:.:-::.--::-:.:-.:.-·!..:.:--:.:-=---::-~·~-=~'-=-__....::..::.I__________ I¢

Figure 17

Benthic macrofaunal biomass levels at station ISA. (a) 1972-73 data from R. Swartz and F. Cole (U.S. EPA, CorvaUis, OR 97330) with

only four grabs collected Dec. 1972 and 14 grabs available for Dec. 1973; (b) only four grabs available; (c) one Astarte castanea (11.9 g)

excluded; and (d) only two grab samples analyzed.

Station 15A This station (lat. 40025.6'N, long. 73° 11. 1'W)

was located off Fire Island, New York (Fig. 1) with mediumsand sediments containing low trace-metal levels in a coastal

ridge and swale area and 30 m deep. The site was also in

the coastal-active band, rarely influenced by the HudsonRaritan plume, but possibly subject to coastal upwellings and

seasonal fronts. This site, like station 17, was also dominated

by a large population of the sand dollar Echinarachnius

parma (Fig. 17). This population appeared to be more persistent than at station 17, being a dominant in 1972-73, with

older cohorts in the population collected in 1979. Mean

biomass levels (Figure 17) generally show late-fall peaks,

in contrast to the summer peak seasonal pattern shown by

most other stations in this study. These peaks mostly reflect

the population dynamics (recruitment, growth, and mortality) of E. parma. A variety of predators were collected ~ere,

including windowpane, yellowtail, four-spot, and wmter

flounders; little skate; scup; and Cancer crabs. Mysids were

an important prey for many of the fish predators, e.g.,

yellowtail and windowpane flounders, and scup; however,

a variety of polychaetes, (e.g., Lumbrineris), amphipods,

(e.g., Ampelisca agassizi and Unciola sp.), and Ceriantheopsis were also very important. Only yellowtail ate E. parma

as a minor diet item.

15

Southern New England

The next group of stations were off southern New England

and included three sites on the shelf (stations 19, 20, and

37) and three in coastal or estuarine areas (stations 18, 34,

and 36) (Fig. 1). The deeper shelf sites, stations 20 and 37,

had relatively low mean biomass levels «100 g/m2) with

higher levels, to 250 g/m2, at the inshore sites.

Station 18 This station (lat. 41 °13.5'N, long. 71 °51.1'W)

was in a 45-m deep bathymetric depression in Block Island

Sound (Fig. 1) characterized by silty-very fine sands with

moderately high levels of trace metals (10-20 ppm lead),

probably a fine sediment depositional area. The site was influenced by the Long Island Sound plume and was in a coastal

mixing front area with strong tidal currents. Several taxa contributed to the biomass here (Fig. 18). Dominant species were

the polychaete Clymenella torquata, the nut clam Nucula

proxima, and the amphipod Ampelisca agassizi. The ocean

quahog Arctica islandica was also relatively common, but

was generally excluded from the analysis because of its large

size. Although there was some intercollection variability,

overall biomass levels here were relatively stable to 1982

(150-220 g/m2), with sampling error probably being the
f

• POLYCHAETES

MOLLUSCS

[Jll]] CRUSTACEANS

EJ ECHINODERMS

o Residual

e

b

r--_--J c

400

......

C\I

E Q 300

..........

oW

en + en ...... ttl - 200 E .s::

Cl 0 '0> iii a c

ttl -

Q) Q) 100

O..L.-......-.....,,/-

:!: 1 0 1,2_

/

J..;_6_9_IL,7-1;/f_

1

:LI1__~-: 7__--JI:L:-__~...J.1/_8-0--7_IL8-1--_----2.J..

/8

__2 -_-:-~L8-_2_~-:L/_8-2-------7--J/--,84101 0 ~(!).W~ ....

gc.> CJ> E .-- ----1- -- .. ,'£'" £ £ £ £.. .£

IX 0

I¢

Figure 18

Benthic macrofaunal biomass levels at station 18. (a) Based on three grab samples (Steimle 1982); (b) one Mercenariil

mercenariil (210.0 g) excluded; (c) one Arctica islandica (440.0 g) excluded; (d) one A. islandica (53.6 g) excluded; (e)

one A. islandica (186.1 g) excluded; and (f) one each Aphrodite hastaJa (3.3 g), A. islandica (357.1 g), and Caudina arenaJa

(4.6 g) excluded•

12/79 7/8011/80 7181 2/82 8/8212/82 7/83 7/84 6/85

~~ 20

1

' _ £ £ t t I i10:(;~ o Cl .---------.- --------.------.----- ---

I- IXE 0 l- 0 I¢

300

.....

C'J c; E .

..........

oW

en + en ......

Ettl -.s:: 200

o Cl iii '0>

c

Q) Q) ttl -

~ ~

100

o

• POLYCHAETES

MOLLUSCS

[Jll]] CRUSTACEANS

EJ ECHINODERMS

o Residual

a

c d

a

Figure 19

Benthic macrofatmal biomass levels at station 34. (a) Residual biomass was mostly ceriantharian anemones; (b) one Mercenariil

mercenariil (17.6 g) excluded; (c) one Havelockia scabra (2.2 g) and Caudina arenata (3.0 g) excluded; and (d) only three

grab samples analyzed.

16
• POLYCHAETES

fIll MOLLUSCS

mm CRUSTACEANS

OJ ECHINODERMS

o Residual

(31»):

a

c

7/8011/80 7/81 2/82 8/8212/82 7/83 7/84 6/85

n,..--1:,--~-~.---.!.:,--~-~~-.--.~--. :...J.~_~~_._._~_-. ~_.:.L...I~-._.-~_-.~---'.i'-.--.~-~:'-'__.1...- ----'-- -----1.----,[

IX 0 1Jl5

Figure 20

Benthic macrofauna! biomass levels at station 36. (a) Residual biomass mostly ceriantharian anemones;

(b) one Busycon canaliculatum (16.5 g) and one Mercenaria mercenaria (57.2 g) excluded; and

(c) an unusually large collection of Leptocheirus pinguis.

cause of the November 1980 aberrant collection (it was extremely difficult to maintain the position of the research

vessel in the strong tidal flows). The potentially important

predators were the same inshore group found as far south

as station 17, i.e., little skate, windowpane, winter flounder,

and red and spotted hake, among others. Several amphipods,

Ampelisca sp., Leptocheirus pinguis, and Unciola sp., were

major prey for most fish species here, but the polychaetes

Nephtys incisa and Pherusa, Cancer crabs, and Crangon

were also important.

Station 34 This 32-m station (lat. 41 °24'N, long. 71 °25'W)

was at the mouth of Narragansett Bay (Fig. 1) in silty muds

with moderate levels of trace metals, similar to the last

station. The site was in a coastal-active area influenced by

the Bay plume and upwellings. The biomass here was

dominated by polychaetes (mainly Nephtys incisa early on

and Clymenella torquata later) and molluscs (mainly Nucula

proxima). It was relatively stable until 1982 when an overall

17

increase was evident involving the same species. Seasonal

fluctuations appeared to be a major source of variability until

seasonal collections ceased in 1983 (Fig. 19). Among the

typical group of inshore predators, defined previously and

found here, there was a more abundant lobster population.

A wide variety of prey were found in the fish stomachs examined here, which included several common polychaetes:

Pherusa, N. incisa, and Lumhrineris; ampeliscid amphipods,

Cancer, Crangon, Ceriantheopsis, and a few small bivalves,

e.g., Nucula.

Station 36 This station (lat. 41 °29'N, long. 70 053'W), in

lower Buzzards Bay (Fig. 1), was 23 m deep and in silty

mud with up to 540 ppb PCBs and 560 ppt PAHs, probably

from nearby New Bedford Harbor. The mean biomass levels

ranged from 61.2 to 173.8 g/m2 (Fig. 20) with the

polychaete Nephtys incisa, the nut clam Nucula proxima, and

the anemone Ceriantheopsis americanus being the dominant

species. The increase in 1985 was due to large contributions
400..--------------- ---,

......

C\l

E q 300

o,rn

.........

II) + II) ......

Cll 1: 200 E Cl

'(jj

c: 3:

~ ~ '100

o

b

• POLYCHAETES

IHl MOLLUSCS

[[[Ill CRUSTACEANS

D ECHINODERMS

o Residual

e

f

9/78 12/79 7/8011/80 7/81 2/82 8/8212/82 7/84

~g. ~E~ 101

. l;-.- - .:.:•..:~~.. ~~

~~.---.-------.--- __-.1'.

~~~~ ~_1....

I

.

!'00~'-'

IX 0 ----------- IJZJ

Figure 21

Benthic macrofauna! biolllUllevels at station 19. (a) Only three grabs sampled; (b) one Arctica islandica (300.0 g) excluded;

(c) several Astarte sp. (180.0 g) excluded; (d) two A. islandica (520.7 g) excluded; (e) one A. islandica 038.0 g) excluded;

and (f) two A. islandica 057.7 g) excluded.

by the amphipod Leptocheirus pinguis and the bamboo worm

Asychis elongata. Spider crabs Libinia emarginata, along

with scup, winter flounder, and black seabass, were the common predators. Commonly consumed prey were similar to

the last two inshore stations (18 and 34).

Station 19 This station (lat. 400 41.4'N, long. 71°21'W)

was on the shelf off Rhode Island (Fig. 1) in 62 m of water

with medium-to-fine sand sediments and low trace-metal

levels, probably in the "cold pool." The overall biomass

levels (Fig. 21) were relatively consistent and ranged from

33.1 to 185.6 g/m2 • Seasonal fluctuations appeared to be a

major source of variability. The species which dominated

this biomass were small ocean quahogs Arctica islandica

(larger individuals were collected but excluded) and the amphipod Ampelisca agassizi. Predators collected included

yellowtail flounder, silver and red hakes, among other lesscommon species. A. agassizi was an important diet item for

most predators, as well as the commonly co-occurring amphipod species Leptocheirus and Unciola, the ubiquitous Cancer

and Crangon, and glycerid and lumbrinerid polychaetes.

Station 20 This station (lat. 400 15.2'N, long. 70049.l'W)

was on the outer shelf south of Rhode Island (Fig. 1) at a

depth of 120 m in sandy silt with moderate levels of trace

metals, at the edge of the depositional "Mud Patch" to the

east. It may be influenced by the "cold pool" and the shelfslope water mass front. Biomass levels, which ranged

between 50 and 100 g/m2 , suggest no obvious trends. The

brittlestar Amphioplus abdita was the dominant species and

the source of most of the intercollection variability, with the

18

bivalve Lucinoma sp. also occasionally being important (Fig.

22). Potential predators included silver, red, and white hake;

four-spot flounder; and Cancer crabs, among others. Only

a few fish stomachs were examined, and they suggest various

polychaetes and Crangon may be significant prey.

Station 37 This station (lat. 40 029.7'N, long. 70 012.2'W)

was also midshelf, south of Nantucket Shoals in an area called

the "Mud Patch" (Fig. 1), an extensive silty area, 60 m deep,

with sandy silt sediments that contained up to 8 ppb PCBs

and 150 ppt PAHs. It could have been affected by the "cold

pool" and the shelf-slope water mass front. Benthic collections suggested fairly low but stable levels, between 35.4

and 51.5 g/m2 (Fig. 23). Polychaetes were the dominant

taxa, especially Ninoe nigripes, along with the amphipod

Ampelisca agassizi. The ocean pout Macrozoarces americanus led the list of potential predators, which also included

red and silver hakes; Cancer crabs; and the four-spot and

yellowtail flounders and witch flounder Glyptocephalus

cynoglossus. Polychaetes, especially Pherusa, Nephtys sp.,

Glycera sp., and Lumbrineris, were important in the diets

of many of the fish species common here, but A. agassizi,

Cancer, Crangon, and Dichelopandalus were also commonly

consumed.
100

..... ..., N

E q C,rn

....... .... Ul + Ul ....... eu

E .s::.

-

Cl 0 '(jj iii

c 50 eu -

Q) Q)

(54.8>1 • POLYCHAETES

MOLLUSCS

!IIlIl CRUSTACEANS

EJ ECHINODERMS

o Residual

12/79 7/8011/80 7/81 2/82 8/8212/82 7/84 7/83

201' I I , , , , 1

1

..Cl 1--------.---.-------•------. .•.• 0 ....... i:n~=. ....... Cl . . • . . . . •• ••• . • ••. . . . -----1.. ,.• 0....... . _ ........•......

g L 0 e"

IX 0 1JZ1

Figure 22

Benthic macrofaunal biomass levels at station 20. (a) This collection might have been off station by 2 km;

and (b) one Cancer sp. (13.0 g) excluded.

100....---------------------------.,

..... ..., N

E q C,rn

....... .... Ul + Ul ....... eu - 50 E .s::.

Cl 0 '(jj iii

c:

eu

Q) -

Q)

o

"0 • POLYCHAETES

MOLLUSCS

'" !IIlIl CRUSTACEANS .c

c

0 OJ ECHINODERMS -

!II '" o Residual

0

C

0

a 0

='"0

0

!II

I

"0

!II

'":0

"0

z

7/8011/80 7/81 2/828/8212/82 7/83 7/84 6/85

..

201' I I I! I 1100~:~.... N• ....... Cl -1- . 0....... ......... . '-'~-.~:.',"" .... .-- ---- o . ------.- ~IX 0 L..- _

1JZ1

Figure 23

Benthic macrofaunal biomass levels at station 37. (a) Residual biomass mostly rhynchocoels and ceriantharian

anemones; (b) one holothurian Molpadia oolitica (20.0 g) excluded; (c) another M. oolitica (15.8 g) excluded;

and (d) one Caudina arenata (16.9 g) excluded.

19
• POLYCHAETES

MOLLUSCS

!llIII CRUSTACEANS

EJ ECHINODERMS

o Residual

c

Station Location "a" Station Location "b"

100.----------~------------------___,

1><

0.L~l;JJ~~~ii7i/ij7i9i9/-7-9J--7/80 11/80 7181 2/82 8/8212/82 7184

7/63 1-4...J/7_8_9....J/L78 ~----'- ....JL_-'-__ __-'-_---'-_~ "'-, 10 '-'

=gl-e ~~ 5

0

:::;:' [1:' I 'e---e" :-:- ~'.:. ;.:.-<..........- e---e 10 1J15<.!JCn~. - .------e

,.....

C\I

E q c,C/)

'-' ,.... III +

gJ ::: 50

E a

o Cl

iii

c

«l - (1) (1)

~ ~

Figure 24

Benthic macrofaunal biomass levels at station 22. (a) One grab sample only; (b) only three grab samples available for 1978-79 collections,

except Sept. 1978; and (c) one Cancer irroratus (8.4 g) excluded.

Georges Bank

The next three stations (22, 23, and 24) were on Georges

Bank (Fig. 1) and had relatively low «50 g/m2) mean

biomass levels. The only exception occurred at station 23

because of sand dollar E. parma population dynamics.

Station 22 This station on southern Georges Bank (Fig. 1),

was moved in 1980 from lat. 400 21.5'N, long. 68°29'W

(station 22a), to lat. 400 30'N, long. 68°0<)'W (station 22b).

Both sites were 110 m deep with medium sand at the initial

site and very fine sand, with near undetectable PCB and PAH

levels, at the second site. The shelf-slope front and "cold

pool" could have affected both sites. Biomass levels at both

sites were about equal, 16 to 49 g/m2 , with the second site

showing seasonal variability (Fig. 24). The dominant species

were the tube-dwelling amphipod Ampelisca agassizi and the

polychaete Ninoe gayheadi. The predator guild was similar

to that found at the last two sites, with at least the ubiquitous

Cancer and Pagurus commonly eaten.

Station 23 This central Georges Bank (Fig. 1) station was

located (1at. 40 0

58'N, long. 67°33'W) in 70 m with mediumto-fine sand and low trace metals, in a ridge and swale area.

Ridges were typically 5-15 m in amplitude and 150-175 m

apart. The site could have been in the central Bank mixing

area with strong tidal currents. Biomass levels here were

unusual in that they showed a notable increasing trend after

1980, from <40 to >70 g/m2 (Fig. 25). The increased levels

in July 1981 were partly the result of a large quantity of

epifaunal hydrozoans being collected; however, some of the

increase can be attributed to the amphipod Byblis serrata and

the sand doil?lr Echinarachnius parma, with the razor clam

Siliqua squama adding to levels in July 1982. Yellowtail

flounder was a common predator collected at the site, along

with little skate, red and silver hake, ocean pout, haddock

Melanogrammus aeglefinus, winter flounder, and others. The

most common prey found in the stomachs examined were,

again, the Ampelisca-Leptocheirus-Unciola group of amphipods, the Nephtys incisa-Pherusa polychaete group, and the

decapod group, Cancer-Crangon-Pagurus-Dichelopandalus.

Winter flounder was the only species to use the hydroids as

food, possibly ingesting them secondarily while feeding on

other species hiding within the colonies.

20
• POLYCHAETES

MOLLUSCS

lIIIll CRUSTACEANS

El ECHINODERMS

o Residual

(76.1)

150

2/82 7/82 12/82

b

718011/80 7/81 , , 7/799/79

0

4/78 9/78

~rj ...

•

f-E O IX

,...

C\IE q c,C/)

.........

l/l + l/l ::: 100 III .r:; E 01

'0> c: ;:

III 03

;:

50

Figure 25

Benthic macrofaunal biomass levels at station 23. (a) Only three grab samples available for 1978-79; (b) this

collection might have been off station by 2 km; (c) unusual contribution by hydrozoa; (d) one Arctica islandica

(613.2 g) excluded; and (e) another A. islandica (107.0 g) excluded.

Station 24 This station on the northeast "peak" of Georges

Bank (Fig. 1) also consisted of two sites; the first, station

24a (lat. 41 °50'N, long. 67°51'W) was moved to be consistent with other sampling. Most of the data are from the

second site, station 24b (lat. 42°00'N, long. 67°00'W),

which was 65 m in depth with coarse, gravelly sand containing low trace metals. It was in a dynamic ridge and swale

area influenced by upwellings and strong currents. Biomass

levels at this hydrographically active site were very low, <20

g/m2 (Fig. 26). The overall biomass levels at the site were

very stable, with only one polychaete, Nepthys bucera,

possibly qualified to be considered as a dominant. The

predator guild at this site included all those mentioned at the

last station but with a stronger contribution by the longhorned sculpin Myoxocephalus octodecemspinosus and the

cod Gadus morhua. Basically the same prey were important

here as at the last station.

Gulf of Maine

The final two stations (28 and 35) were in the Gulf of Maine

(Fig. 1) with several species of echinoderms dominating the

biomass. Mean biomass levels varied, but were generally

between 50 and 200 g/m2•

21
Figure 26

Benthic macrofauna! biomass levels at station 24. (a)

Only two grab samples available Apr. 1978-Sept. 1979;

and (b) mean biomass ofthis

collection was 5.8 g' m2 if

the unusual sample of Echinarachnius parma were

excluded.

• POLYCHAETES

MOLLUSCS

lIIlIl CRUSTACEANS

[] ECHINODERMS

D Residual

Station 24b

b

(11 B) Station 24a

9/79,1,2/79 7/8011/80 7181 2/82 7/8212/82 7/83

'----=

=-----__"-=-" .4...=:=..::.'.='==:~_'~'.4. ~_!50 ~... ...... Cl~_ .-- M"__ --_. -- -

IfZJ

100

0

4/78 9/78

~rj .4.

I-EO •

IX

.......

C\l

E Q 0lC/)

...... .... en + en ...... ro

E ..cOJ 0 'm iii 5: c

- 50 ro (1) (1) 5:

Figure 27

Benthic macrofaunal biomass

levels at station 28. (a) Data

from a single grab sample (D.

Maurer 1982) at the same location; (b) one Arctica islandica

(10.7 g) excluded; and (c) one

Astarte borealis (12.0 g) excluded.

• POLYCHAETES (31.6)

MOLLUSCS

lIIlIl CRUSTACEANS

100 [] ECHINODERMS

o Residual

O............V.,L

....... 05: 20[1/77,~~~2J...~7_9 1_1J...~8_0 7..L~_8_1__2.J.~8_2 1_2J...~8_2__7/L8_3 7...L~84110 • • (1) ,! 0 __._-_-_-_-_-_--_-_-_-_--_-_._-_-_..-_._:-_.-._1_~_.-._":-_.•_-_.... _~_.._~_~:._.-_..; J

IfZJ

.......

C\l

E Q 0lC/)

...... .... en + en ...... ro

E ..cOJ 0 'm iii 5: 50 c

ro -(1) (1) 5:

22
300

,...

C\J ci E . OlC/)

'-' ,.... C/) + c/)'-' E1: 200

o Ol

[is

C

t1l Q)

~ ~

100

o

• POLYCHAETES

ffil MOLLUSCS

lllIll CRUSTACEANS

[J ECHINODERMS

o Residual

12/79 7/80 11/80 7/81 2/82 8/82 12/82 7/84 6/85,...

~ge I~EOl: 200IL'..J..~~..~.~..~.~..~ .M~:~'.~~:~~~~~~:~~~_._~_..._._.~_~_ ...._.~.'~''''~~~~~~~~~~~~~~~:~~~~~~~~:~I 0

10

~:Q) ... !:l. 'oJ -------', ._ - _e 'e -- e e----- e--

IfZJ

Figure 28

Benthic macrofaunal biomass levels at station 35. (a) One Molpadia oolitica (16.2 g) excluded.

Station 28 This 60-m deep station (lat. 41°50.6'N, long.

69°30'W) was in the southwest Gulf of Maine (Fig. 1) with

silty-clay sediments and moderately high trace-metal levels.

The collection sequence here was incomplete (Fig. 27) but

suggests a general increase between 1977 «10 g/m2) and

1980-83 (67-122 g/m2). It is obvious that most of this increase was due to echinoderms, primarily two species: the

urchin Brisasterjragi/is and a brittlestar Ophiura sp. Based

on a cursory size-frequency distribution analysis, the major

biomass contribution of each of these species was from older

cohorts, although there was evidence of new Ophiura recruitment almost every year. These recruitments either did not

survive to the next collection or had limited growth, as sizefrequency modal progressions were not evident. Predators

collected at this site included the American plaice Hippoglossoides platessoides, cod, silver hake, and little skate.

Predation data were not available from this station.

Station 35 This station (lat. 42°19'N, long. 70 0

36'W) was

in Massachusetts Bay (Fig. 1) at 60 m in a postglacial depositional area near a mixed waste-disposal site, the "Foul

Grounds;" sediments were silt with up to 30 ppb PCBs and

1200 ppt PAHs. The site was in, or near, an area where

seasonal fronts develop; a general warming trend has been

23

noted in the Gulf of Maine for the last decade. Biomass

levels, 100-250 g/m2, did not suggest any pattern or trend,

except polychaete seasonality (Fig. 28). Most ofthe biomass

came from polychaetes, with Spio filicornis prominent, and

the echinoderms Ctenodiscus crispatus and Molpadia

oolitica. Predators trawled at this site included American

plaice, cod, red and silver hakes, and witch, winter, and

yellowtail flounders. Ophuroids (brittlestars) were important

prey to cod and plaice, pandalid shrimp to the hakes, and

polychaetes, e.g., sabellids and nereids, to the other flounder.

Discussion _

These results suggest several general conclusions: (1) there

was considerable interannual variability in the biomass data

at most sites, as might be expected, but within ranges previously reported for each area; (2) the range of variability

differed at each site and seasonal patterns of fluctuation were

often evident; (3) with only a few exceptions, the general

tendency of the mean biomass, within the time frame and

sampling scale involved, did not usually suggest any clear

increasing or decreasing trends; (4) the population dynamics

of one or a few species were found to be important in the
explanation of some of the variability and apparent trends;

and (5) the relationship of the biomass levels to other environmental factors, e.g., sediment type or quality, could

not be fully evaluated with the available data. These conclusions or general observations and several others are expanded

upon below and, although the relationships may be tentative

at this time, they are noted to suggest hypotheses that could

be explored in the future.

Seasonal cycles of benthic community biomass levels are

a common, if not normal, pattern described frequently (see

Beukema 1974, Buchanan et a1. 1978, Glemarec and

Menesguen 1980). Seasonal fluctuations were evident at

many of the sites, e.g. stations 3, 9, 32, 19, and 22 (Figs.

3,6, 8,21, and 24) with biomass peaks generally occurring

in the summer. Some sites that did not show higher mean

summer values were dominated by sand dollar Echinarachnius parma populations, whose relatively long life-span, 7+

yrs (Steimle 1990), can mask much of the seasonal fluctuations of other species with generally shorter or annual life

cycles.

Review of Figures 2-28 also suggests that biomass levels

at most of the sites appeared to be either relatively constant

(considering normal seasonal fluctuations and disregarding

a few unusual collections), demonstrated no clear increasing or decreasing trends, or the data were too irregular. Some

dispersed sites, e.g., stations 12, 23, 34, 36, and 28 (Figs.

10, 19,20,25, and 27), had biomass levels that did suggest

overall long-term increasing trends. This was supported by

{-tests indicating the slopes were significantly different at

p<0.05. These trends were frequently related to the population dynamics or growth of one or a few dominant species,

e.g., sand dollars at station 23, or possibly to nearby

eutrophic estuarine enrichment, e.g., stations 34 and 36. A

decreasing trend was suggested at stations 16B, 16C, and

33 (Figs. 13, 15, and 16), and to a lesser degree at stations

11, 24b, and 26 (Figs. 9, 24. and 26). The {-tests indicated

that the trends at stations 16B and 33 were significant at

p<0.05. Stations 16B and possibly 33 were in an area considered to be variably affected by pollution from various

sources. No strong reason can be suggested for the decline

at the other sites, although the population dynamics of E.

parma are involved at station 11. The remaining sites, 31

and 32 (Figs. 7 and 8), had collection periods that were either

too brief in duration, discontinuous, or too variable to suggest trends.

Although mean sediment grain size and TOC are only two

of many sediment variables, they can be important to benthic community structure. The sediment data, summarized

also in Figures 2-28, despite being incomplete or discontinuous, suggest these two variables were relatively stable

at most sites. When a sedimentary change was evident,

possibly because of sampling error, there was seldom a

significant, consistent, concomitant response in biomass. The

proximity of many sites, e.g., stations 7, 8, 17, and 15A,

near or in ridge and swale areas suggest a source of some

of the variability in the data at these sites. Schaffner and

Boesch (1982) have demonstrated how benthic community

24

structure can vary significantly relative to location on sand

ridges, slopes, or swales. It is possible that these submerged

sand dunes might move, thus besides the possible variability

caused by minor shifts in sample location in a ridge and swale

environment, future monitoring at these sites needs to take

into account the potential natural changes that may occur at

precise, fixed locations to accurately interpret any later community changes. Generally, the dominant species found at

any site were consistent with the major habitat-related

community types defined by Pratt (1973), considering these

communities are usually not discrete but occur in a variable

continuum (Mills 1969).

Hydrographic conditions at a site can potentially influence

benthic communities, but information on the significance of

this factor in the study area is mostly lacking. These factors

are most likely to be important in coastal areas rather than

offshore. For example, although it is not well documented,

it is very probable that estuarine plumes influence coastal

benthic communities and these plumes can vary in volume,

particulate load, or distribution over time. Chronic seasonal

hypoxia is one possibly plume-related problem that can be

important to benthic communities in some coastal areas, e.g.,

along the New Jersey and Delaware coast (Reid et al. 1987).

Benthic community changes at plume-influenced sites should

be evaluated against any major changes in the plumes, e.g.,

excessive freshwater runoff and particulate load due to major

storm events in the drainage area. The influence of the

variable, coastal-active band is also not well studied, although

a variety of physical and chemical stresses are features of

this band. The potential occurrence and influence of shifting water mass fronts at several sites cannot be truly assessed

in the data presented here. These fronts have been shown

to be important to the shelf benthic community in at least

one study (Magnuson et a1. 1981). Likewise, the influence

of the variably configured, semipermanent "cold pool"

cannot be assessed at the sites likely to be affected, although low temperatures are known to be important to

benthic communities (Buchanan et al. 1978). Tidal velocity

gradients also can be important (Creutzberg 1984), but the

significance of this and other hydrographic factors to the

benthic community have not yet been evaluated in the study

area.

The influence of anthropogenic activities on benthic biomass levels are reported frequently (Pearson and Rosenberg

1978, Cedarwall and Elmgren 1980, Steimle 1985, Pearson

and Barnett 1987). The sediments at several stations were

moderately to highly contaminated by trace metals or toxic

organic chemicals, e.g., stations 31,33, 18,34,20,28,36,

37, 16A-C, and 35. There may be a suggestion of an effect

from this contamination in some of the data examined in this

study, i.e., at stations 33 and 16B. The declining trends at

these two New York Bight apex stations are possibly a

response to the increase in sewage sludge disposal after 1979,

although the actual increase in sewage sludge solids is

reported to be only 5% (Reid et a1. 1987). The biomass data

at other contaminated stations, even those near other dumpsites, e.g., station 35 in Massachusetts Bay and station 9 at
the former dumpsite off Delaware, do not show any clear

response. However, this may be because there is inadequate

information about these sites, e.g., pre-impact data, or

suitable control areas. Alteration of other benthic community

structure variables was reported for the former Delaware

dumpsite (Lear and O'Malley 1983).

Several studies have demonstrated the influence of predators on benthic fauna (McIntyre 1978, Berge 1980, Persson

1981, Choat 1982, Arntz and Rumohr 1986). The predatoroccurrence summaries presented for each site are only gross

indicators of the potential predation pressures at any site.

Although the predator groups were typical of what would

be expected at any site and vary with season and population

dynamics of the species involved (Grosslein and Azarovitz

1982), the limited data on diets at the sites is inadequate to

discuss any more than crude linkages between predators and

the benthic community. However, the data are consistent with

several more comprehensive studies in the region (Langton

and Bowman 1980, 1981; Sedberry 1983). Overall, the

dietary data discussed here, and generally supported by other

studies mentioned above, suggest that despite the diversity

in most benthic communities, much ofthe diet consists quantitatively of a limited number of benthic prey species. These

are generally characterized as being common at the sediment

surface and relatively large or conspicuous, e.g., the tubedwelling amphipods Ampelisca, Leptocheirus, Unciola, the

polychaetes Nephtys, Pherusa, Lumbrineris, the tube-dwelling anemone Ceriantheopsis, and especially the smaller

decapods such as small Cancer, Crangon, Pagurus, and

Dichelopandalus. These motile decapods are most likely not

well-sampled in either grab or trawl collections and their

overall abundance and biomass underestimated. Molluscs and

echinoderms are not generally consumed, despite their

availability, although some predators do eat them as a

relatively common or predominant prey. This suggests that

fish-predation influences to the benthic community, although

variable, are relatively selective. There also is an intermediate

trophic level involved, i.e., many of the "prime" prey listed

above (especially the decapod crustaceans) are themselves

predators of smaller organisms, at least being omnivores

(Commito and Ambrose 1984). What this would mean to

benthic biomass levels and trends can only be speculation.

For example, the low predation pressure on echinoderms and

subsurface dwellers, such as most molluscs and many

polychaetes, may explain why trends can be relatively stable

since these species tend to be the dominant biomass components. However, in terms of value to fishery resources,

the prime prey types listed above must be either very productive or more abundant than their apparent low occurrence

in grab or trawl collections would suggest.

Energy equivalents of the benthic biomass are not presented since they closely parallel the biomass-level patterns

at each site, differing proportionally by the variable mean

Kcal/g ratios defined for each site (Table 2). It had been anticipated that the energy in the benthic pool would be more

conservative, with less variability, since it should reflect the

basic primary production and distribution characteristics of

25

an area; it should also remove some of the variability caused

by the water or shell content of the biomass. However, this

was not evident; some sites had higher winter energy

equivalents that moderated seasonal differences, but other

sites had higher summer equivalents. The minor overall

seasonal difference in energy equivalents, 0.81 to 0.84, is

probably not significant. Because the energy equivalents vary

between benthic groups (Steimle and Terranova 1985), the

proportional contribution of the groups to mean estimated

energy levels of each site changed from that defined by the

original biomass data. Higher energy-equivalent polychaetes

(1.09 Kcal/g wet wt) and crustaceans (1.29 Kcal/g wet wt)

increased their share of the total, while lower-equivalent

molluscs (0.37 Kcal/g wet wt) and echinoderms (0.48 Kcal/g

wet wt) declined. The overall means, 0.81-0.84 Kcal/g, with

a median of 0.82 Kcal/g, were consistent with an earlier 0.8

Kcal/g estimate (Steimle and Terranova 1985), based on

Wigley and Theroux's (1981) summer data. This suggests,

if only wet-weight benthic macrofauna biomass data are

available for the northeast continental shelf, the use of the

0.8 Kcal/g conversion will provide a reasonable (±25%)

estimate ofthe energy pool. Ifspecific areas or communities

are to be considered, then the site-specific data of Table 2

could be used to refine the estimate.

It is apparent that the biomass, at almost all sites, was

dominated by three groups: molluscs, polychaetes, and

echinoderms, with cerianthiid anemones and crustaceans

occasionally being important. This dominance is similar to

the results of Wigley and Theroux (1981) for the Middle

Atlantic Bight, although they show a higher proportion of

molluscs than generally found in this study. This difference

is probably because many of the megafaunal species excluded

in the present analysis were molluscs. Molluscs and polychaetes were notable contributors to the biomass at almost

all sites, with polychaetes being particularly abundant in

siltier sites, e.g., stations l6A-C, 33, 34, 35, 36, and 37.

These sites are all near estuaries or in coastal areas, except

station 37 (the "Mud Patch" off Nantucket). In some cases,

this dominance may reflect some anthropogenic influence as

suggested for other areas (Reise 1982). The echinoderms

were dominated by a few species, e.g., Echinarachnius

parma on Georges Bank and at several Middle Atlantic Bight

sites, and brittlestars or holothurians on deeper areas of

Georges Bank and in the Gulf of Maine basins. The dominant taxa were generally stable in their proportional contribution to the overall biomass at each site. Exceptions occurred

when a community was influenced by the population dynamics of a particularly dominant species, such as the brittlestar

Amphioplus at station 20 or E. parma at several sites.

The separate study of the population dynamics of Echinarachnius parma (Steimle 1990), focusing on four stations (11,

17, 15A, and 23), provided insight on how the recruitment,

survival, and growth of this species can greatly influence

biomass trends. For example, at station 11 there were substantial annual recruitments in 1980-82. This is evident in

the station's biomass (Fig. 9) as the expansion of the echinoderm biomass component (almost entirely the recruitment
and growth ofthese E. parma cohorts) after July 1980, with

a peak in early 1982. The subsequent decline occurred as

recruitment was reduced or ceased in 1983-84 and mortality outpaced growth. At station 17, there were only two

recruitments, one in 1976 after the anoxia event defaunated

the area, and the second in 1978 (Fig. 12). These two

recruitments constitute the entire echinoderm population in

Figure 12. This sand dollar population was not growing any

faster than at the other stations. It also could have beer. subject to a lower mortality because most nonfish predators or

competitors were also removed from the area by the anoxic

conditions. The average life span of this species appears to

be about 8 years in this area (Steimle 1990), and it can be

anticipated that unless there is new recruitment the role of

this species at the site will diminish and other taxa should

increase, as they may be inhibited by the dense E. parma

population. At station 15A, E. parma has been a persistent

part of the community since at least the early 1970s (Fig.

17). There have been annual recruitments between 1978 and

1982 (Steimle 1990) evident as winter/spring peaks to 1982

(Fig. 17). The growth of a dominant 1978 cohort is re~ponsible for the 1983 biomass peak, and mortality in this cohort

is the cause of the subsequent decline. At station 23 a situation similar to station 17 was evident with no sand dollar

population evident in the community studied until recruitments occurred between 1978 and 1982, the last collection

there. The cohorts at this Georges Bank station appeared to

have a lower survival than at the other three sites studied,

since the population did not expand via these frequent recruitments and their expected growth, as they did elsewhere. This

may be related to Collie's (1987) observation of preferential feeding by yellowtail flounder on <12 mm E. parma on

Georges Bank. Much of the gradual increase in community

biomass after 1978 (Fig. 25) was the result of these recruitments and their growth.

In the discussion of biomass variability at each site, the

influence on the biomass from population dynamics of only

a few echinoderm species was considered. There are other

species that need to be considered, e.g., molluscs, as suggested by Nucula and Ilyanassa at station 31. They would

be expected to produce results similar to that of the echinoderms. Several bivalve molluscs are commercially harvested

which also affects their abundance and population dynamics.

Population dynamics of dominant species should be examined

as part of any interpretive monitoring program (Pearson and

Barnett 1987). Life-history summaries (Caracciolo and

Steimle 1983) and studies ofsome dominant species (Steimle

1990) are becoming available and should be used or considered in evaluating the relative status or trending of a

community. The importance of biological interactions to

long-term community stability is well known (Gray 1977).

Some community change can be expected, because of these

interactions and natural environmental factors, including a

change to significantly different or alternate communities.

The definition and understanding of these natural fluctuations or changes are extremely important in detecting and

understanding anthropogenic effects.

26

The mean biomass ranges found at the sites from the 1970s

and 1980s were generally consistent with the values reported

for nearby stations or areas by Wigley and Theroux (1981)

and Theroux and Grosslein (1987), based on extensive collections in the late 1950s and early 1960s. The only notable

exception was the recent data for Buzzards Bay, MA (station 34, Fig. 19), which was generally above 100 g/m2 ,

compared with the Wigley and Theroux (1981) value of

<25 g/m:! for this area. The number of sites monitored in

the present survey were a small fraction (about 3%) of the

earlier 1950s-60s studies; there were small differences in

the methods involved. Other comparisons between the conclusions of these two earlier surveys and the recent series

suggest little difference in overall shelf faunal composition

(bivalves, polychaetes, and echinoderms as biomass dominants), bathymetric trends (general inverse relationship), and

other relationships.

There have been relatively few subtidal benthic macrofaunal studies that have included long-term biomass data, and

these were mostly for European waters as reviewed by Pearson and Barnett (1987). There also have been other studies,

again in the same areas, that have examined relatively limited

data sets separated by several decades (Pearson and Barnett

1987). These studies were often for the purpose of attempting to define changes in the benthic community that could

be related to pollution, especially organic enrichment. In

some cases, changes were found that appeared to support

that relationship (Cedarwall and Elmgren 1980, Pearson et

al. 1985). But these studies are not without the possibility

of alternate interpretation because of a number of confounding factors. These can include differences in methods or other

environmental variables other than pollution, including

changes in the demersal fish populations in the area over the

period being compared (Persson 1981). In other cases, a clear

benthic biomass response to pollution was evident (Pearson

and Rosenberg 1978). In general, these other long-term

studies show results or conclusions similar to those described in this paper: (1) Seasonal biomass fluctuations were

usually evident as late summer-early fall peaks, with late

winter-early spring lows (Buchanan et al. 1978); (2) biomass

levels and trends can be significantly influenced by the

population dynamics of a few larger, long-lived species, frequently echinoderms and molluscs (Pearson et al. 1985);

(3) there can be substantial interannual variability in the

amplitude of the biomass ranges (Blegvad 1951); and

(4) cause-and-effect relationships of any changes or trends

are usually difficult to determine with reasonable confidence

because of the usually large variety of additional factors that

can be involved and need to be monitored (Buchanan et al.

1978). In interpreting causal relationships, the types and

levels of change which are significant for resource management purposes need to be defined (Green 1984, Duinker and

Beanlands 1986). The changes of significance are likely to

be, as McIntyre (1978) states, either a major qualitative

alteration in the biomass or a shift to a new population structure which has significance to predators. Experiments or

studies are needed to defme the term "major" and determine
what benthic population structures are of importance to

fishery resources.

Because the sites examined were not in rocky habitats, the

communities associated with these habitats are not discussed

here or in Wigley and Theroux (1981). These habitats and

their epifauna are important to fisheries, but almost nothing

has been done in the Northeast to quantify the potential forage

value of the community or its natural variability or production for comparison with non-rocky bottom areas. This is

especially important because of the recent interest and increased use of artificial reef habitats as fishery-enhancement

or habitat-loss mitigation tools.

In summary, long-term benthic macrofaunal biomass data

from a number ofsites on the continental shelf of the northeastern United States showed little evidence of clear temporal trends. It appears that most sites had relatively stable

biomass levels, although there were seasonal and annual

variability. The population dynamics of a few species, such

as the sand dollar Echinarachnius parma, were a major factor in variability at several sites. Preliminary examinations

of other potential causes of variability were inconclusive

because of inadequate data but suggested a number of information deficiencies that should be explored in the future. This

study demonstrates the need for caution in using limited benthic biomass data to characterize a station or area, since this

variable is dynamic.

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