NOAA Technical Report NMFS 140

A Technical Report of the Fishery Bulletin

Quantitative Composition and Distribution

of the Macrobenthic Invertebrate Fauna

of the Continental Shelf Ecosystems

of the Northeastern United States

Roger B. Theroux

Roland L. Wigley

December 1998

U.S. Department of Commerce

Seattle, Washington
Suggested reference

Theroux, Roger B., and Roland L. Wigley. Quantitative composition and

distribution of the macrobenthic invertebrate fauna of the continental

shelf ecosystems of the northeastern United States. U.S. Dep. Commer.,

NOAA Tech. Rep. NMFS 140, 240 p.

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CONTENTS

Introduction 1

Order of Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Previous Studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Macrofauna Samples 5

Sampling Gear . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Sample Processing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Data Treatment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Geographic Areas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Bottom Sediments 9

Bathymetry .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 10

Temperature. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 10

Sediment Organic Carbon. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 11

Description of the Region 11

Topography. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 11

Bottom Sediments 13

Sediment Organic Carbon. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Hydrography. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Zoogeography. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Faunal Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Total Macrobenthos-All Taxonomic Groups Combined 28

Geographic Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 28

Bathymetric Distribution. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 40

Relation to Bottom Sediments 47

Relation to Water Temperature. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 51

Relation to Sedimen t Organic Carbon. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 53

Taxonomic Groups. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 55

Porifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 56

Coelenterata : . . . .. 71

Hydrozoa 71

Anthozoa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 75

Alcyonaria 75

Zoantharia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 79

Platyhelminthes 81

Turbellaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81

Nemertea. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 85

Aschelminthes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 91

Nematoda. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 91

Annelida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 94

Pogonophora 100

Sipunculida 103

Echiura 108

Priapulida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 108

iii
Mollusca 110

Polyplacophora 110

Gastropoda. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 119

Bivalvia 124

Scaphopoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 130

Cephalopoda 135

Arthropoda. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 139

Pycnogonida 139

Arachnida 143

Crustacea 143

Ostracoda. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 144

Cirripedia 145

Copepoda. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 155

Cumacea 155

Tanaidacea 159

Isopoda. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 162

Amphipoda 165

Mysidacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 170

Decapoda 174

Bryozoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 178

Brachiopoda 181

Echinodermata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 185

Crinoidea 186

Holothuroidea 187

Echinoidea 192

Ophiuroidea 199

Asteroidea. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 205

Hemichordata 208

Chordata 210

Ascidiacea 210

Dominant Components of the Macrobenthos 214

Frequency of Occurrence 216

Percentage Composition " 216

Geographic Distribution 216

Selected Genera and Species 217

Phylum Annelida 217

Phylum Mollusca 217

Phylum Anthropoda 217

Phylum Echinodermata 224

Bathymetric Distribution 224

Relation to Bottom Sediments 225

Relation to Water Temperature 225

Relation to Sediment Organic Carbon 226

Acknowledgments 227

Literature Cited 227
Quantitative Composition and Distribution of the

Macrobenthic Invertebrate Fauna of the Continental Shelf Ecosystems

of the Northeastern United States

ROGER B. THEROUX*

ROLAND L. WIGLEY * *

Woods Hole Laboratory

Northeast Fisheries Science Center

National Marine Fisheries Service, NOAA

Woods Hole, Massachusetts 02543

ABSTRACT

From the mid-1950's to the mid-1960's a series ofquantitative surveys ofthe macrobenthic

invertebrate fauna were conducted in the offshore New England region (Maine to Long

Island, NewYork) . The surveys were designed to I) obtain measures of macrobenthic

standing crop expressed in terms of density and biomass; 2) determine the taxonomic

composition of the fauna (ca. 567 species); 3) map the general features of macrobenthic

distribution; and 4) evaluate the fauna's relationships to water depth, bottom type, temperature range, and sediment organic carbon content. A total of 1,076 samples, ranging from 3

to 3,974 m in depth, were obtained and analyzed.

The aggregate macrobenthic fauna consists of 44 major taxonomic groups (phyla,

classes, orders). A striking fact is that only five of those groups (belonging to four phyla)

account for over 80% of both total biomass and number of individuals ofthe macrobenthos.

The five dominant groups are Bivalvia, Annelida, Amphipoda, Echninoidea, and

Holothuroidea.

Other salient features pertaining to the macrobenthos of the region are the following:

substantial differences in quantity exist among different geographic subareas within the

region, but with a general trend that both density and biomass increase from northeast to

southwest; both density and biomass decrease with increasing depth; the composition ofthe

bottom sediments significantly influences both the kind and quantity of macrobenthic

invertebrates, the largest quantities of both measures of abundance occurring in the coarser

grained sediments and diminishing with decreasing particle size; areas with marked seasonal changes in water temperature support an abundant and diverse fauna, whereas a

uniform temperature regime is associated with a sparse, less diverse fauna; and no detectable trends are evident in the quantitative composition of the macrobenthos in relation to

sediment organic carbon content.

Introduction

The broad continental shelf off the northeastern coast

of the United States is a particularly significant topographic feature of the continental margin because ofits

influence on the marine life ofthe region. Water masses

overlying this large shelf, and neritic waters generally,

are noted for their abundance of plankton, fishes, and

associated organisms, some endangered. Noteworthy

of the offshore New England waters, including Georges

Bank, are the rich harvests of fish that have been taken

each year since pre-Colonial days. The marine life inhabiting New England offshore waters has been the

subject ofstudies conducted from time to time throughout the past century. This has resulted in the acquisition of a considerable body of knowledge on the fishes

and plankton in this region, but information about the

benthic invertebrates has been rather limited, espe-

* Present address: P.O. Box 306 East Flamouth , MA 02536.

** Present address: 35 Wilson Road, Woods Hole, MA 02543.
2 NOAA Technical Report NMFS 140

cially regarding quantitative aspects. Because of the key

role played by macrobenthic invertebrates in the ecological dynamics of the marine environment, their usefulness to man as a food resource, their potential as

concentrators of toxic substances that could be transmitted through the food chain, and their usefulness as

indicators of environmental change, the National Marine Fisheries Service (formerly the Bureau of Commercial Fisheries) of the U.S. Department of Commerce, NOAA, in cooperation with the U.S. Geological

Survey and the Woods Hole Oceanographic Institution

conducted a quantitative survey of the benthos of the

entire continental margin of the eastern United States.

The investigation of the macrobenthic invertebrates

was an integral part of a broad program of study of the

Atlantic continental margin (Emery and Schlee, 1963;

Emery, 1966b).

This report is the second of two which describe the

quantitative distribution of macrobenthic invertebrates

of the Atlantic continental shelf and slope. The first

(Wigley and Theroux, 1981) describes the quantitative

distribution of m~or taxonomic groups ofmacrobenthic

invertebrates inhabiting the continental shelf and slope

between Cape Cod, Massachusetts, and Cape Hatteras,

North Carolina. Their distribution in relation to geographic location, water depth, bottom sediments, range

in bottom water temperature, and sediment organic

carbon content is considered.

The present report describes the quantitative distribution ofthe principal groups of macrobenthic invertebrates inhabiting offshore New England waters. The

area studied extends from the mouth of the Bay of

Fundy eastward to Nova Scotia (longitude 64° West)

and southward to central New Jersey. The quantity of

each major taxonomic group is considered in relation

to the same environmental variables. Only the broad

distributional aspects of major groups are presented

and evaluated here. Other aspects of the benthic fauna

derived from these samples, such as community composition, trophic zonation, faunal dominance and diversity, and similar topics will be the subjects of future

reports.

The large database generated by the Continental

Margin Program contains a wealth of valuable geological, faunal, and environmental information of historical as well as current significance. In addition to providing input for a variety of descriptive studies, as described above, the potential exists for information contained in the database for ecosystem modeling tasks;

paleoecological and global climate change studies; and

benthic production estimates (Cohen et al. 1978, 1982;

Cohen and Wright 1979; Warwick 1980; Rowe et al.

1986, 1988; Bourne 1987; Cohen and Grosslein 1987;

Steimle 1987, 1990a, 1990b; Rowe et aI., 1991; and

others) .

Order of Discussion

The first section of this report briefly describes the

principal physical features of the region, providing a

general background for understanding the distribution

of the various faunal groups. This section is followed by

the main body of data describing the quantitative distribution of 44 faunal groups in relation to the five environmental parameters: 1) geography, 2) bathymetry, 3)

bottom sediments, 4) bottom water temperatures, and

5) sediment organic carbon. Quantitative data for geographic distribution are presented at two different levels: a detailed evaluation based on calculations for each

of several hundred unit areas (20 min in latitude by 20

min in longitude); and a less detailed evaluation based

on six large geographical subareas within the region

studied. Faunal groups are chiefly phyla, classes, and

orders ofmacrobenthos presented in phylogenetic order.

The final section is a summary of the environmental relationship of the dominant taxonomic components.

Previous Studies

One of the earliest studies in marine benthic ecology

dealt with populations inhabiting the Woods Hole-Vineyard Sound area offsoutheastern Massachusetts (Verrill

et aI., 1873). This well-known study is not only the first

comprehensive report dealing with the New England

marine benthos but also one of the earliest ecological

accounts of marine zoobenthos in all scientific literature. Included in the report are descriptions of new

species, an annotated catalog of animals found in Vineyard Sound and vicinity, and, significantly, a large part of

the report is devoted to descriptions of the benthic communities and the biotopes they inhabit. Although a small

number of published reports on New England natural

history observations and taxonomic studies were available

as sources ofinformation to supplement theirstudy (Gould

1841,1870; Desor 1851; Stimpson 1851, 1853; Verrill 1867;

and others), by far the bulk of all information contained

in the report by Verrill et al. is based on original collections and observations.

Between 1871 and 1887 nearly 2,000 benthic fauna

samples were collected in waters off the northeastern

United States by the U.S. Fish Commission in cooperation with the U.S. Revenue Service, U.S. Coast Survey,

and zoologists from American universities. Dredging

and trawling were the principal methods of collecting

samples. A large proportion of the samples were collected in coastal areas between New Haven, Connecticut, and Eastport, Maine; only a moderate number of

collections were from offshore areas. Inshore operations were conducted from the vessels Moccasin,

Mosswood, Bache, Speedwell, Blue Light, and to some ex-
_________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 3

tent the Blake and Fish Hawk; however, the latter two

also operated in offshore areas, as did the Albatross and

the chartered fishing schooner,JosieReeves (Packard, 1874,

1876: Agassiz, 1881; Smith and Rathbun, 1882; Tanner,

1882; Smith and Rathbun, 1889; Townsend, 1901).

This early sampling was primarily exploratory in nature. The participating zoologists faced a vast unstudied fauna and a multitude of species new to science.

Scientists most active in this work were chiefly systematists; consequently the results were largely taxonomic

accounts of various groups. The following are typical

examples: Smith, 1879, 1884; Harger,1880, 1883;

Rathbun, 1880; Wilson, 1880; Fewkes, 1881; Verrill ,1881,

1884; Agassiz, 1883, Webster and Benedict, 1884; Bush,

1885; Bigelow, 1891). Professor Addison E. Verrill of

Yale College, who collaborated closely with U.S. Fish

Commission scientists, was undoubtedly the most productive systematist of this, or perhaps any era. He described over one thousand species representing most

major invertebrate groups. A very large percentage of

these new species descriptions was based on specimens

collected off New England. Although several preliminary ecological studies of the offshore benthos were

reported (Smith and Harger, 1874; Verrill, 1874a, 1874b;

Agassiz 1888a, 1888b) and the reports on systematics of

various groups contain ecological information, no comprehensive ecological reports pertaining to the fauna

of this region were published.

The second milestone in ecological research of the

New England marine benthos was a comprehensive

report by Summer et al. (1913). This report is based on

three years of intensive sampling in Vineyard Sound

and Buzzards Bay by the Bureau of Fisheries in 1903,

1904, and 1905. This useful publication not only lists

the species occurring in the Woods Hole region but

includes species distribution charts and discusses some

physical conditions (temperature, depth, and sediments) that influence the distribution of animals. To

this day, this remains the most thorough ecological

study of the New England marine benthos.

After the investigation by Sumner et al. (1913), there

was a 30-year hiatus during which ecological research

on New England marine benthos-particularly that concerned with offshore invertebrates-proceeded at an

exceedingly slow pace. Belding (1914), Allee (1922a,

1922b, 1923a, 1923b, 1923c), Pytherch (1929), Stauffer

(1937), Ayers (1938), and others contributed valuable

information on inshore populations. Rather few ecologically oriented works such as Procter (1933a, 1933b)

and Bigelow and Schroeder (1939) pertaining to offshore zoobenthos appeared during this period. In addition to the foregoing, however, many studies of a taxonomic nature containing valuable ecological information were issued during this time span (Rathbun, 1905,

1925; Koehler, 1914; Nutting, 1915; Pilsbry, 1916; Heath,

1918; Bartsch, 1922; Deichmann, 1930, 1936; and others). Ecological interests of marine scientists conducting field studies in this region centered on plankton and

fishes. It was not until the 1940's that renewed activities in

benthic ecology attained a significant level. Beginning in

that decade a number of investigations were undertaken

concerning inshore populations (Dexter, 1944, 1947; Lee,

1944; Phleger and Walton, 1950; Swan 1952a, 1952b;

Parker, 1952; Pratt, 1953; Burbanck et aI., 1956; Parker

and Athern, 1959; Stickney, 1959; Rhoads, 1963; and others). Ecological studies pertaining to the offshore populations commenced somewhat later, for example the reports by: Parker (1948); Northrup (1951); Phleger (1952);

Clarke (1954); Schroeder (1955, 1958); Taylor et al. (1957);

Wigley (1959); Wieser (1960); Wigley (1960b); Chamberlin

and Stearns (1963); and Wigley and Emery (1968), are

notable examples.

Perhaps the most significant event of this period,

relative to the present work, was the inauguration of

quantitative benthos investigations of the New England

marine fauna (Lee, 1944). Lee's work was a study of the

macrobenthic invertebrate fauna of Menemsha Bight,

an embayment ofVineyard Sound, Massachusetts. Years

later, quantitative studies were made of the benthos of

Long Island Sound (Sanders, 1956; Richards and Riley,

1967), Buzzards Bay (Sanders, 1958, 1960; Wieser, 1960),

Barnstable Harbor (Sanders et. aI., 1962), Greenwich

Bay, Rhode Island (Stickney and Stringer, 1957),

Sheepscot Estuary (Hanks, 1964), Narragansett Bay

(Phelps, 1965), Rand's Harbor, Massachusetts

(Burbanck et aI., 1956), and other locales. In recent

years, due to increased interest in potential impacts of

man's activities in outer continental shelf (OCS) development and exploitation and in understanding the

dynamics of marine ecosystems, quantitative studies of

the benthic fauna in the New England region have

undergone a marked increase, as have studies in other

associated disciplines. Studies such as Wigley (1961 b);

Sanders et. al. (1962); Wigley and McIntyre (1964);

Emery et al. (1965); Nesis (1965); Sanders et al. (1965);

Owen et al. (1967); Wigley and Emery (1967); Wigley

(1968); Mills (1969); Wigley and Theroux (1970);

Haedrich, et al. (1975); Rowe et al. (1975); Wigley et al.

(1975); Uzmann et al. (1977); Pearson and Rosenberg

(1978); Maurer and Leathem (1980, 1981a, 1981b);

Valentine etal. (1980); Magnuson et. al. (1981); Wigley

and Theroux (1981); Maurer and Wigley (1982, 1984);

Steimle (1982); Caracciolo and Steimle (1983); Lear

and O'Malley (1983); Steimle (1985); Rowe et al. (1986);

Maciolek and Grassle (1987); Michael (1987); Theroux

and Grosslein (1987); Langton et al. (1988); Langton

and Uzmann (1988); Sherman et al. (1988); Langton and

Uzmann 1989, Langton et. al. (1990); and Rowe et. al.

(1991), and as well as others have provided much needed

insights into the complex ecosystems of the region.
4 NOAA Technical Report NMFS 140

Several published indexes and bibliographies include

many references to the general literature pertaining to

benthic invertebrates and allied subjects. Many of the

historical as well as the modern reports are included

among the citations in these bibliographies.

The interested reader may wish to consult the following:

1 Fishery Publication Index, 1920-1954. U.S. Fish &

Wildlife Service Circular 36, published in 1955.

2 Publications of the United States Bureau of Fisheries 1871-1940. Compiled by Barbara B. Aller and

published in 1958.

3 A Preliminary Bibliography with KWIC Index on

the Ecology of Estuaries and Coastal Areas of the

Eastern United States. Compiled by Robert

LivingstonJr. and published in 1965.

4 Marine and Estuarine Environments, Organisms and

Geology of the Cape Cod Region, an Indexed Bibliography, 1665-1965. Compiled by Anne E. Yentsch,

M. R. Carriker, R. H. Parker, and V .A. Zullo, published in 1966.

5 Fishery Publication Index, 1955-64. U.S. Fish & Wildlife Service, Bur. Comm. Fish. Cire. 296, published

in 1969.

6 The Effects ofWaste Disposal in the New York Bight.

Compiled by the National Marine Fisheries Service,

Middle Atlantic Coastal Fisheries Center, Sandy

Hook, NewJersey, published in 1972.

7 Coastal and Offshore Environmental Inventory: Cape

Hatteras to Nantucket Shoals. Edited by Saul B.

Saila and published in 1973.

8 Bibliography of the New York Bight: Part I-List of

Citations; Part 2- Indices. Compiled by the National Oceanic and Atmospheric Administration,

Marine Ecosystems Analysis Program, Stony Brook,

N.Y, published in 1974.

9 Fishery Publication Index, 1965-74. Compiled by

M. E. Engett and L. C. Thorson, U.S. Dep. Commerce, NOAA Tech. Rep. NMFS Cire. 400, published in 1977.

10 A Summary and Analysis ofEnvironmental Information on the Continental Shelffrom the Bay ofFundy

to Cape Hatteras (1977). Vol. II, Master Bibliography, Index, Acknowledgements. Prepared for the

Bureau of Land Management by Center for Natural

Areas, published in 1977.

11 The Bay Bib: Rhode Island Marine Bibliography,

Revised Edition. Coordinated by C. Q. Dunn and L.

Z. Hale, edited by A. Bucci, Coastal Resources Center, Northeast Regional Coastal Information Center, Marine Advisory Service, National Sea Grant

Depository, Univ. ofRhode Island Mar. Tech. Rep.

70, published in 1979.

12 An Ecological Characterization of Coastal Maine

(North and East of Cape Elizabeth). Vol. 5, Data

Source Appendix. Compiled by S. E. Fefer and P. A.

Schetting for BioI. Servo Program, Interagency Energy/Environment Res. and Dev. Program, Office

of Res. and Dev., U.S. Environmental Protection

Agency, published in 1980.

13 Benthic Productivity and Marine Resources of the

Gulf of Maine. I. Babb and M. DeLuca (eds.). National Undersea Research Program, Research Report 88-3, published in 1988.

Another result of increased OCS activity is the large

volume of information relating to benthic fauna appearing in the so-called "gray" literature. Included in

this cattgory are completion reports of field study contracts, environmental impact statements, public and private agency investigation reports, annual reports, and other

similar special documents. Many appear in irregular series, or are one-of-a-kind reports, often in photocopied or

mimeographed form and, as such, are not always listed in

the usual literature sources (e.g. Maurer, 1983; Michael

et. aI., 1983; Pratt, 1973; also see Literature Cited).

In addition to Wigley and Theroux (1981) there are

several taxonomically or ecologically oriented reports

based wholly or in part on the samples forming the

basis of the Northeast Fisheries Science Center (NEFC)

benthic database. Such reports include Wigley (1960a,

1960b, 1961a, 1961b, 1963a, 1963b, 1965, 1966a, 1966b,

1968, 1970, and 1973); Pettibone (1961, 1962, 1963);

Chamberlin and Stearns (1963); Emery and Merrill

(1964); Wigley and McIntyre (1964); Emery et al. (1965);

Trumbull (1965); Merrill et al. (1965); Wigley and Shave

(1966); Wigley and Emery (1967); Schopf (1968b);

Haynes and Wigley (1969); Plough (1969); Hazel (1970);

Merrill (1970); Wigley and Theroux (1970); Kraeuter

(1971); Wigley and Burns (1971); Wigley and Theroux

(1971); Bousfield (1973); Cutler (1973, 1977); Wigley

and Stinton (1973); Murray (1974); Wigley et al. (1975);

Wigley and Messersmith (1976); Wigley et al. (1976);

Williams and Wigley (1977); Kinner (1978); Merrill et

al. (1978); Plough (1978); Brodeur (1979); Watling

(1979a); Dickinson et al. (1980); Franz and Merrill

(1980b); Dickinson and Wigley (1981); Franz et al.

(1981); Maurer and Wigley (1982, 1984); Maurer (1983);

Shepard and Theroux (1983); Therouxl ; Theroux and

Wigley (1983); Rowe et al. (1986); Shepard et al. (1986);

Bousfield (1987); Rowe (1987); Theroux and Grosslein

(1987); Langton and Uzmann (1988); Langton et al.

(1988); Rowe et al. (1988); Sherman et al. (1988);

Langton and Uzmann (1989); Langton et al. (1990);

1 Theroux, R. B. 1983. Collection data for the U.S. east coast

gastropod mollusks in the Northeast Fisheries Center Specimen

Reference Collection, Woods Hole, Massachusetts. U.S. Dep.

Commer., NOAA, Nat!. Mar. Fish. Serv., ortheast Fish. Cntr., Woods

Hole Lab. Ref. Doc. No. 8327, 280 p. Unpub!. manuscript.
Composition and Distribution of Macrobenthic Invertebrate Fauna 5

Materials and Methods

Macrofauna Samples

Table 1

Research vessels, cruise numbers, date of collections,

and number of samples obtained.

This report is based on 1,076 quantitative samples of

macrobenthic fauna collected during 22 cruises by 5

research vessels between 1956 and 1965 (Table 1). The

geographic locality of sampling sites is illustrated in

Figure 1, and sampling density is illustrated in Figure 2

in which the number of samples in each geographic

unit area is indicated (dimension of each unit area is 20

minutes latitude by 20 minutes longitude). Collection

data (including cruise, station, and collection numNumber of

Date samples

August 1956 35

August ]957 ]65

August 1959 75

June ]961 75

June 1962 ] 23

April 1963 7

April 1963 3

May 1963 38

May 1963 29

July 1963 1

Auguu]963 93

August-September 1963 32

October 1963 9

October 1963 84

August 1964 72

September 1964 7

April 1964 8

July-August 1964 62

October 1964 24

October-November 1964 10

August 1965 ] 23

1,076

80

10]

59-9

61-10

62-7

10

11

12

]3

20

22

24

28

29

49

51

1

2

64-12

64-13

65-11

Cruise

Vessel number

Albatmss 1/1

Albatmss [1/

Delaware

Delaware

Delaware

Gosnold

Gosnold

Gosnold

Gosnold

Gosnold

Gosnold

Gosnold

Gosnold

Gosnold

Gosnold

Gosnold

Astmas

Astmas

Albatmss [V

Albatross IV

Albatmss IV

Total

change. The results of those programs covered a broad

spectrum of interdisciplinary topics which expanded our

understanding of the marine environment (e.g. Pearce,

1971,1972,1974, 1975; Pratt, 1973; Pearce et aI., 1976a,

1976b, 1976c, 1976d, 1977a, 1977b, 1977c, 1978, 1981;

Caracciolo et aI., 1978; Pearson and Rosenberg, 1978;

Reid et aI., 1979; Steimle and Radosh, 1979; Warwick,

1980; Schaffner and Boesch, 1982; Steimle, 1982; Boehm,

1983; Caracciolo and Steimle, 1983; Lear and O'Malley,

1983; Steimle, 1985; Steimle and Terranova, 1985; Duinker

and Beanlands, 1986; Howart,1987; Neff, 1987; Reid etal.,

1987; Steimle, 1990a, 1990b; Steimle et aI., 1990).

and Rowe et al. (1991); Burns and Wigley2; Wigley et a1. 3;

Theroux and Wigley4; Maurer and Wiglet; Theroux, et

a1. 6; Theroux and Schmidt-Gengenback. 7

Other uses to which the data have proven useful in

the past, as well as in the present, have been varied.

Included have been environmental impact statements

prepared by various public agencies (Dep. Interior,

Minerals Management Service; U.S. Army Corps of Engineers; NOAA, etc.) relating to OCS activities (e.g. oil

and gas exploration, mining, dredging, dumping, etc.);

international litigation (i.e. US/Canada Boundary

Case); marine sanctuary designation proposals (e.g.

Stellwagen Bank, Norfolk Canyon); and others.

Several specially targeted programs initiated in the

latter 1970's and terminated in the mid- to late 1980's

have provided additional impetus for an increase in

attention devoted to the macrobenthos of the region.

During that period many studies were conducted by

public and private agencies and academic institutions

(e.g. NOAA's Northeast Monitoring and Ocean Pulse

Programs; the Northeast Fisheries Center's Marine Resources Mapping, Assessment, and Prediction program

(Sherman, 1980); the Marine Ecosystem Analysis Program (MESA) (see Pearce et aI., 1981); the Woods Hole

Oceanographic Institution's Georges Bank Study program,

and many others). Those studies, in both inshore and

offshore areas, were designed to establish baselines for

assessing environmental quality and to monitor the impacts of present and future activities related to oil and gas

exploration and production, marine mining, ocean dumping, other waste disposal, and natural environmental

2 Burns, B. R., and R. L. Wigley. 1970. Collection and biological

data pertaining to mysids in the collection at the BCF Biological

Laboratory, Woods Hole. U.S. Bur. Comm. Fish. Bio!. Lab. Woods

Hole, Mass., Lab. Ref. 70-3,36 p. (mimeo). Unpub!. manuscript.

3 Wigley, R. L., R. B. Theroux, and H. E. Murray. 1976. Marine

macrobenthic invertebrate fauna of the Middle Atlantic Bight region. Part 1. Collection data and environmental measurements.

Northeast Fisheres Center, Woods Hole Lab. Ref. Doc. 7618, 34 p.

(mimeo). Unpub!. Manuscript.

4 Theroux, R. B., and R. L. Wigley. 1979. Collection data for U.S.

east coast bivalve mollusks in the Northeast Fisheries Center Specimen Reference Collection, Woods Hole, Massachusetts. Northeast

Fisheries Center, Woods Hole Laboratory, National Marine Fisheries Serv., NOAA, Northeast Fisheries Center.,Woods Hole Lab.

Ref. Doc. 79-29, 471 p. (mimeo). Unpub!. Manuscript.

;; Maurer, D., and R. L. Wigley. 1981. Distribution of biomass and

density of macrobenthic invertebrates on the continental shelf off

Martha's Vineyard, Massachusetts. National Marine Fisheries Service,

Northeast Fisheries Center, Woods Hole Laboratory, NOAA, Woods

Hole Lab. Ref. Doc. 81-15, 97 p. (mimeo). Unpub!. manuscript.

6 Theroux, R. B., R. L. Wigley, and H. E. Murray. 1982. Marine

macrobenthic invertebrate fauna of the ew England Region: Collection data and environmental measurements. at. Mar. Fish. Serv., NOM,

Northeast Fish. Center, Woods Hole Lab. Ref.Doc. 82-40, MARMAP

Contrib. MD/NEFC 82-67, 74 p. (mimeo). Unpub!. Manuscript.

7 Theroux, R. B., and]. Schmidt-Gengenbach. 1984. Collection data

and environmental measurements for U.S. east coast Cumacea (Arthropoda, Crustacea) in the Northeast Fisheries Center Specimen Reference Collection Woods Hole, Massachusetts. Nat. Mar. Fish. Serv.,

Northeast Fisheries Center, Woods Hole Lab. Ref. Doc. 84-27, MARMAP

Contr. MED/NEFC 83-46,114 p. (mimeo). Unpub!. Manuscript.
6 NOAA Technical Report NMFS 140

NEW

JERSEY

4·

42·

69"

•

o 120 IL-~_-JI KM

M A I N E

70"

• •

4,·

71·

•

• •

•

•

40·

72·

NEW /

HAMPSHIRE "

/

: PORTLAND

• •

•

•

•

•

•

•

•

39·

•

•

•

•

• •

•

•

•

•

•

•

• •

•

•

•

•

•

38·

•

•

•

•

•

•

• •

•

• •

•

•

•

•

•

•

•

•

••

•

•

•

•

•

74

73

71

72

Figure I

Chart of the study area showing the location of stations where quantitative samples of macrobenthic invertebrates were

obtained.

bers, number of samples, latitude, longitude, date of

sampling, and type of gear used) and environmen tal

measurements (including water depth in meters, bottom type, geographic subarea, temperature range [0C],

and percent organic carbon) for each sampling site are

contained in Theroux et al. Ii).

Sampling stations were located in all sections of the

study area, but somewhat more intensive coverage was

given to the offshore continental shelf region than to

the inshore bays and sounds or to the deep water region beyond the continental shelf. Table 2 lists the

number ofsamples and occurrence frequency for e2.ch

parameter grouping. A moderate number of samples,

however, were taken in the major bays and estuaries,

and in deep water. Ninety-two samples were collected at

depths less than 24 m, and 93 samples from depths

greater than 500 m. The continental rise was only

sparsely sampled because of its great depth and the

correspondingly increased time required to obtain

samples. Minimum and maximum depths at which

samples were taken were 3 and 3,975 m.

Sampling Gear

The samples consisted of bottom sediments with the

constituent fauna collected with a Smith-McIntyre springloaded grab sampler (Smith and McIntyre, 1954) illustrated in Figure 3, or a Campbell grab sampler (Menzies

et aI., 1963) illustrated in Figure 4. The bottom area

sampled by the Smith-McIntyre sampler was 0.1 m2 which

had a capacity ofapproximately 15 liters (L). Area sampled

by the Campbell sampler was 0.56 m2, which had a volume

capacity of about 200 L. The Campbell grab was equipped

with a 35-mm camera and electronic flash, housed within

the buckets, to obtain photographs of the bottom immediately before impact (Emery and Merrill, 1964; Emery et

aI., 1965; Wigley and Emery, 1967; Theroux, 1984).

Sample Processing

Aboard ship, the material obtained at each sampling

site by each sampler was processed and preserved as a
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 7

NUMBER OF SAMPLES

PER UNIT AREA

( 20' x 20')

• 1 - 2

• 3 - 5

• 6 - 10

• ,,- 25

72° 71° 700 69°

, 0 120 . • KM 68°

M A I N E . ,/

. +,

/' ..(

I·-.

Figure 2

Chart showing sampling intensity within each standard unit area (20 min. latitude by 20 min. longitude). All samples

within each unit area have been added to indicate sampling density.

separate sample. The contents of the sampler were

emptied into a bucket or tub calibrated in liters, or

directly into a wash-box (volume measured by means of

a calibrated rule) from which two small subsamples

were removed prior to washing. One of these subsamples

was for meiofauna, and the other for sediment analysis.

Total quantity removed ranged from 0.25 to 1.0 L,

depending upon the total volume of material obtained.

The quantities of both samples and subsamples were

measured and recorded on sample log sheets. Generally, the remaining material was washed through a I-mmaperture mesh-sieving screen. Material remaining on the

screen after washing, consisting of benthic animals, tubes,

shells, shell hash, and coarse sediments, was preserved in a

buffered seawater solution offormaldehyde and brought

to the laboratory ashore for further processing.

Laboratory processing involved separating the preserved

organisms from the mineral debris, sorting them to major

taxonomic groups, identifYing them to the lowest practicable taxonomic level, counting, and weighing. Weights

are damp formalin weight, the "rough weight" ofPetersen

(1918), herein referred to as wet weight inasmuch as the

superficial fluid on the specimens was removed by blotting before being weighed on a Mettler precision balance

to the nearest 0.01 g. Weights include shells and skeletal

materials that constitute an integral part of the living

animal, i.e. shells of living mollusks, brachiopods, and

skeletal structures of bryozoans, barnacles, and similar

organisms. Materials omitted in the weighing procedure

were: tubes of polychaetous worms, gastropod and

scaphopod shells inhabited by pagurid crabs orsipunculid

worms, and othersimilar nonintegral structures or nonliving animal remains. Counts of the number of specimens

were made for all groups. Colonial animals were treated as

individuals; that is, one sponge colony, or a colony of

bryozoans was each counted as an individual specimen;

colonies are much more comparable in size to individuals

of noncolonial animals than are the zooids making up the

colony. Also, the disparity in size from smallest to largest

colonial organisms was only slightly greater than the size

differential between small and large individuals of

noncolonial species.
8 NOAA Technical Report NMFS 140

Specimens of each taxon were bottled separately in

70% ethanol and labeled. Subsequently, specimens were

assembled by taxonomic groups and sent to cooperating systematists for species determinations. There were

more than 40 specialists from the United States and from

other countries cooperating in this part of the study.

Data Treatment

Information pertaining to the location, collecting methods, physical and chemical characteristics of the enviTable 2

umbers ofsamples and occurrence frequency in eac~'l

of the various parameter groupings used in this repor:.

Number Frequency (%)

of of

Parameter samples occurrence

Geographic area

Nova Scotia 85 7.9

Gulf of Maine 303 28.2

Georges Bank 211 19.6

Southern New England Shelf 344 32.0

Georges Slope 52 4.8

Southern New England Slope 81 7.5

Depth range (m)

0-24 92 8.6

25-49 160 14.9

50-99 319 29.6

100-199 246 22.9

200-499 166 15.4

500-999 22 2.0

1000-1999 34 3.2

2000-3999 37 3.4

Sediment type

Gravel 148 13.8

Till 22 2.0

Shell 6 0.6

Sand 455 42.2

Sand-silt 211 19.6

Silt-clay 234 21.8

Temperature range (0C)

0-3.9 335 31.1

4-7.9 158 14.7

8-11.9 336 31.2

12-15.9 157 14.6

16-19.9 62 5.8

20-23.9 28 2.6

Sediment organic carbon (%)

0.00 5 0.5

0.01-0.49 418 38.8

0.50-0.99 167 15.5

1.00-1.49 84 7.8

1.50-1.99 43 4.0

2.00-2.99 13 1.2

3.00-4.99 4 0.4

5.00+ I 0.1

missing data 341 31.7

ronment, and the number and weight of the biological

components of each sample was recorded on preprinted

data forms. The coded information and quantitative

data from the records were entered on automatic dataprocessing cards. Data were summarized by computer

in a form similar to that presented in the tables appearing in the body of this report.

The principal units used for expressing the quantity

of benthic invertebrates (quantity per unit area) are: 1)

density-number of individual specimens per square

meter of bottom area, and 2) biomass-wet weight, in

grams, per square meter of bottom.

Faunal density values used in constructing quantitative

geographic distribution charts for the various taxonomic

groups (Figs. 12,27,33,39,45, etc.) are mean values for

all samples within each unit area as shown in Figure 2.

Qualitative and quantitative differences between seasons and between years were sufficiently small to permit

the consolidation of all samples for purposes of this

report. Some seasonal and yearly differences in taxonomic composition and quantity of animals were detected within specific geographic localities that were

repeatedly sampled. With few exceptions, however, the

dissimilarities were relatively minor in comparison to

the differences from one geographic locality to another. One of the chief reasons for the temporal stability was the presence of many animals having a long

(one year to a century or more) life span. The common

occurrence of sessile forms and nonmigratory motile

forms also contributed to the observed constancy in

biomass. Similar conditions were reported by Zatsepin

(1968, in Steele, 1973) in reference to macrobenthos

samples taken in the Barents Sea and Norwegian Sea

over a 30-year period. He found that a comparison of

samples taken in the same regions in different years

"... showed no substantial changes in the quantitative

distribution of the bottom fauna." Several recent reports also allude to the temporal persistence of certain

dominant components of the macrobenthos of the region (Steimle, 1990a, 1990b; Maurer8; Michael et aI. 9 ).

Geographic Areas

For purposes of detecting and reporting regional differences in faunal composition the region has been

Maurer, D. 1983. Review of benthic invertebrates of Georges

Bank in relation to gas and oil exploration with emphasis on

management implications. Nat!' Mar. Fish, Serv., Northeast Fisheries Center, Woods Hole, Massachusetts, Woods Hole Lab. Ref.

Doc. 83-16, 329 p. (mimeo). Unpub!. manuscript.

9 Michael, A. D., C. D. Long, D. Maurer, and R. A. McGrath. 1983.

Georges Bank benthic infauna historical study. Final report to U.S.

Dep. Interior, Minerals Management Service, Washington, DC,

Rep. 83-1 by Taxon Inc. Salem, MA 01970, 171 p.
Composition and Distribution of Macrobenthic Invertebrate Fauna 9

Figure 3

Side view of the Smith-Mcintyre spring-loaded bottom sampler in the closed position. Lead weights on each

side are set vertically to impede rotation of the sampler during descent and ascent. Vertical distance from

frame base to top plate is 52 em.

divided geographically into six subareas (Fig. 5). These

are: 1) Nova Scotia, containing 44,816 km2 (13,049

mi2)-encompassing southwestern Bay of Fundy, eastern gulf of Maine, Browns Bank, and the Nova Scotian

continental shelf; 2) Gulf of Maine-all of the Gulf of

Maine except the eastern sector encompassing an area

of 80,067 km2 (23,313 mi2); 3) Georges Bank-consisting only of Georges Bank proper with an area of 39,211

km2 (11,417 mi2 ); 4) Southern New England Shelf occupying 73,318 km2 (21,348 mi2)-including the continental shelf from Great South Channel southwestward

to central NewJersey; 5) Georges Slope-the continental slope from Great South Channel northeasterly to off

the Scotian Banks, an area of 50,706 km2 (14,764 mi2);

6) Southern New England Slope-the continental slope

from Great South Channel southwesterly to southwest

ofHudson Canyon, occupying 62,570 km2 (18,218 mi2).

Each subarea has specific biotopic and biogeographic

faunal characteristics. These are discussed in the section entitled "Description of the Region" and in the

"Geographic Distribution" section for each of the major taxonomic groups.

Bottom Sediments

Bottom sediments from the samples have been analyzed for particle size, composition, and color. In addition, a selected series of these samples was further analyzed for carbonate content (Hulsemann, 1966), quantity of organic matter (Hulsemann, 1967) and

minerology (Ross, 1970b). Detailed particle size analyses of approximately 75% of the samples were made by

John Schlee, U.S. Geological Survey (Schlee, 1973).
10 NOAA Technical Report NMFS 140

Figure 4

Bottom view of Campbell Rrab sampler. Camera housing is installed in right-hand bucket and strobe light is in the

left-hand bucket. Shutter tJip weight is in foreground. Width ofthe buckets (vertical dimension in photograph) is 57 cm.

Approximately 20% of the samples were analyzed by

the New York Soil Testing Laboratory (Wigley, 1961a).

The remaining 5% were classified using field techniqces

by K.O. Emery of the Woods Hole qceanographic Institution or by National Marine Fisheries Service personnel. For additional information concerning sediment

analyses, methodology, and detailed results, see references listed by Emery (1966b) and the section of this

report titled "Description of the Region."

Bathymetry

Water depths, 111 meters, were obtained by means of

echo sounders and precision depth recorders and corrected for hydrophone/transducer depth and temperature effects on the velocity of sound in water.

Temperature

Water temperature and salinity data were based primarily on the hydrographic report prepared by John B.

Colton et al. (1968), which gives detailed information

obtained on eight quarterly (March, May, September,

and December) hydrographic survey cruises from 1964

to 1966. Each cruise covered the entire area from Nova

Scotia to New York. We also used several thousand

bottom temperature records obtained on seventeen

bottom trawl survey cruises of the research vessels Albatross Ill, Albatross Iv, and Delaware, conducted by the

Bureau of Commercial Fisheries Biological Laboratory,

Woods Hole, during the years 1956 through 1965. Additional sources of reference and temperature-salinity

data are: Townsend (1901); Sumner, et al. (1913);

Bigelow (1927, 1933); Edwards et al. (1962); Hathaway
_________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 11

NEW

72

71

/'

HAMPSHIRE ,'. /.

:. POIITLANO

71"

M A I N E

69"

0LI~~-,'f...

4'

Figure 5

Chart of the study area showing the location of the six standard geographic subareas used for analytical

purposes: Nova Scotia, Gulf of Maine, Georges Bank, Georges Slope, Southern New England Shelf, and

Southern New England Slope.

(1966); Schopf (1967); and Schopf and Colton (1966);

and Mountain and Holzwarth (1989).

Sediment Organic Carbon

Organic carbon in bottom sediments was measured by

gasometric method in samples after removal of CaC03

by acid treatment. Data are contained in Hathaway

(1971).

Description of the Region _

Topography

Relief of the sea bottom off the New England region

has been studied most recently by the U.S. Geological

Survey and the Woods Hole Oceanographic Institution·

(Austin et al., 1980; Emery, 1965a, 1966b; Emery and

Ross, 1968; Emery and Uchupi, 1972; Gibson et al.

1968; Klitgord and Behrendt, 1979; Klitgord et al., 1982.

Schlee et al., 1976; Sheridan, 1974; Uchupi, 1965b,

1966a, 1966b, 1966c, 1968; UChupi and Emery, 1967;

UChupi et al., 1977; UChupi and Austin, 1979; Valentine, 1981). Figure 6 is based on, and has been derived

from, a much larger more detailed chart by Uchupi

(1965a), USGS Map 1-451, scale 1:1,000,000.

Topographically, the New England offshore area consists of several large, grossly different geological features. The largest and most complex feature is the Gulf

of Maine, an immense, nearly oval-shaped glacially

eroded depression on the continental shelf. The topography in this depression is very irregular, resulting in

numerous basins separated by ridges, swales, and banks.

These topographic irregularities are due in part to

deposition, gouging, erosion, and related actions during the Pleistocene period of glaciation. Greatest depth

in the gulf is 377 m, in Georges Basin; shallowest offshore depth in the gulf is 9 m, at Amen Rock on Cashes

Ledge in the west central part of the Gulf of Maine (see

Ballard and Uchupi, 1975; Austin et al., 1980; Klitgord

et al., 1982; Schlee et al., 1976).

Georges Bank is another striking topographic feature. It is an enormous (120 km by 240 km) submarine

cuestalike bank situated at the mouth of the Gulf of
12 NOAA Technical Report NMFS 140

2000

4'"

I N E

o 120

I I Kill.

'.'"/

- "

"+ .

.'/:i'·

).'

Figure 6

Chart of the study area showing bathymetric and geographical features, Depth contours are in meters (adapted from

Uchupi 1965),

Maine. The bank slopes gently to the southeast and

south and its surface is relatively smooth except for a

series ofsand ridges in the shallow northwest and northcentral sections. The sand ridges are formed by exceptionally strong tidal currents that prevail in this region.

Tidal currents generally flow with greatest velocity in

the northwest and southeast directions, Further details

relating to Georges Bank are contained in Emery and

UChupi (1965); Uchupi et al. (1977); Valentine (1981);

Butman (1982, 1987); Butman et al. (1982, 1987);

Backus (1987); Bourne (1987); Butman and Beardsley

(1987); Cohen and Grosslein (1987); Cooper et al.

(1987); Emery (1987); Flagg (1987); Howart (1987);

K1itgord and Schlee (1987); Maciolek and Grassle

(1987); Michael (1987); Neff (1987); Twichell et al.

(1987); Uchupi and Austin (1987).

Nantucket Shoals is a relatively shallow and topographically uneven area southeast of antucket Island,

Massachusetts. Principal irregularities are large swales

and ridges extending in north-south and northeastsouthwest directions.

The southern New England continental shelf is a

gently seaward-sloping region with rather smooth topography. Width of the shelf is approximately 100 km

and the shelf break occurs at a depth of about 120 m.

See Garrison and McMaster (1966) for more details.

The continental slope is a narrow zone along the

outer margin of the shelf extending from the shelf

break to a depth of 2,000 m. This zone has a comparatively steep gradient, but less than 5°, and the relief is

moderately smooth except where it is cut by submarine

canyons. The continental rise (2,000-6,000 m) is generally similar to the slope in having only gradual changes in

suriicial topography. However, the overall gradient is substantially less than that for the continental slope. Consult

Emery (1965a), Emery and Ross (1968), Gibson et al.

(1968) ,Schlee et al. (1979), Sheridan (1974), and Uchupi

et al. (1977) for details of topography of this region.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 13

72

71

NEW

JERSEY

~GRAVEL

(::':.1 SHELL

~TILL

~~1~N .... SAND

1::::::1 SILTY SAND """

o SILT-CLAY

38'

72'

"

\ ..•.,. NEW / '\ HAMPSHIRE /

/

: PORTLAND

40'

71'

4,"

70"

M A I N E

69"

o 120 ,-'~,---,I KM

42'

4'

Figure 7

Geographical distribution of bottom-sediment types in the study area.

Bottom Sediments

The composition ofsediments blanketing the sea floor

throughout the study area is well known. Detailed studies have included sedimentological aspects of general

lithology, particle size composition, calcium carbonate

content, organic carbon content, nitrogen content,

minerology, sand and gravel fractions, and other components. A representative selection of publications dealing with the bottom sediments of New England marine

waters includes: Shepard, et al. (1934); Shepard and Cohee

(1936); Stetson (1936, 1938, 1949); Shepard (1939);

Hough (1940, 1942); Wigley (1961a); Uchupi (1963,

] 965b, 1966a, 1966b, ] 966c, ] 968, ] 969); Emery (1965a,

1965b, ]966a, 1966b, 1968); Emery et al. (1965); Rvachev

(1965); Garrison and McMaster (1966); Hiilsemann (1966,

]967); McMaster and Garrison (]966); Ross (1967, 1970a,

1970b); Uchupi and Emery (1967); Emery and Ross (1968);

Schlee (1968, 1973); Schlee and Pratt (1970); Emery and

Uchupi (1972); Trumbull (1972); Milliman (1973); Wigley

and Stinton (1973); Sheridan (1974); Austin etal. (1980);

Twichell et a!. (1981); Butman (1982, ]987); Klitgord et.

a!. (1982); and Valentine et al. (1980).

Relict glacial sediments are the major constituents covering most of the study area, particularly on the continental shelf. Quartz and feldspar sands and granite and gneiss

gravels are particularly common in the shallower areas

and on the topographically high elevations in deeper

water. Fine-textured sediments, mainly silts and clays, that

mantle the continental slope, continental rise, and protected pockets and basins on the continental shelf are

predominantly present-day detrital sediments.

Large areas in the deeper part of the Gulf of Maine

are floored with unsorted glacial till, whereas the shallow banks and ridges are commonly covered with gravel

or sand of glacial origin that remained after washing

action removed the finer particles. In some deep parts

of the Gulf, where water currents are minimal, the till is

overburdened with layers of silt and clay. In Long Island Sound, Buzzards Bay, and many of the smaller

bays along the coast, the sediments are composed largely

of silts and clays, with sand and gravel common in the

nearshore zones.

The sediment chart prepared for this report (Fig. 7)

is based on sediment samples taken from the same grab

hauls from which the fauna was obtained.
14 NOAA Technical Report NMFS 140

Sediment organic carbon

The distribution of organic carbon in the bottom sediments of the region is depicted in Figure 8. Values for

sediment organic carbon content from samples were

low to moderate, ranging from less than 0.5% to slightly

over 7% (7.04). The major portion of the continental

shelf contains small amounts « 0.5%) of organic carbon in sediments, with only small, discrete patches,

especially in the Southern New England shelf area, of

slightly greater amounts (0.5-1.99%). Organic carbon

content ofsediments in the two slope subareas, Georges

Slope and Southern New England Slope, was somewhat

higher than on the shelf with values between 0.50 and

0.99% prevailing and with small areas on the Southern

New England slope containing from 1.00 to 1.99% organic carbon. The sediments in both the Gulf of Maine

and Long Island Sound contain comparatively larger

amounts of carbon, primarily in the 1.00 to 1.99%

range over most of their respective areas. Highest organic carbon content (from 2.00 to 7.04%) was almost

exclusively restricted to the major embayments and

estuaries within the study area; only offshore exceptions to this were two small areas on Stellwagen Bank

and in the area known as Georges Basin, where organic

carbon contents in that range were found.

Hydrography

A substantial amount of information has been amassed

over the years concerning the hydrography of the offshore New England region. Some of the first hydrographic data collected were temperature measurements

taken by Benjamin Franklin's nephew in 1789. Since

that time numerous studies have been conducted

primarily by government organizations, such as the U.S.

Fish Commission (subsequently named the U.S.

Bureau of Commercial Fisheries, and currently called

the National Marine Fisheries Service), the U.S. Coast

Survey (now the National Ocean Survey), U.S. Coast

Guard, Tidal Survey of Canada, Biological Board

ofCanada (Fisheries Research Board ofCanada), coastal

states organizations, Bigelow Laboratory, Woods

Hole Oceanographic Institution, Harvard University,

Massachusetts Institute of Technology, University of

NEW

JERSEY

74

71

SEDIMENT

ORGANIC

CARBON

D <0.50

IillTIFnl 0.50 - 0.99

t.:-:.:.:.j 1.00 - 1.99

.2.00 -7.04

38- 39-

72- 71-

41-

700

M A I N E

69"

o 120 LI_~---,I KM.

Figure 8

Geographic distribution of organic carbon in the boltom sediments.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 15

Rhode Island, and other private and governmental

organizations.

One of the most comprehensive reports on this subject is the monograph entitled "Physical Oceanography

of the Gulf of Maine" by Henry B. Bigelow (1927). He

describes the essential features of water temperature,

salinity, tidal and nontidal circulation, and seasonal

variation in these hydrographic features. Much detailed

information was added in succeeding years particularly

by John B. Colton and his associates at the Bureau of

Commerical Fisheries Biological Laboratory at Woods

Hole, Massachusetts, and by Dean F. Bumpus and his

colleagues at the Woods Hole Oceanographic Institution (Stetson, 1937; Bumpus, 1960, 1961; Colton, 1964;

Bumpus and Lauzier, 1965; Bumpus etal., 1973; Butman

et aI., 1980, 1982; Dorkins, 1980; Ramp et aI., 1980,

Moody et. aI., 1984, Mountain and Holzwarth, 1989,

among others). Discussions of early oceanographic research in this region and references to the literature

are given by Colton (1964), Schopf (l968a), and Wright

(1987).

In brief, the main features pertaining to water circulation in the study area are as follows: 1) cold water on

the Nova Scotian Shelf flows southwestwardly along

that feature and turns northward into the GulfofMaine;

2) Gulf of Maine waters form a large non tidal counterclockwise gyre; 3) waters overlying Georges Bank form

a clockwise gyre; 4) nontidal currents generally flow

southwestwardly and westward across Nantucket Shoals

and on the Southern New England continental shelf; 5)

freshwater runoff from land empties by means of large

New England and Canadian rivers into the northern

and western sections of the study area; 6) incursions of

relatively warm high-salinity slope water enter the Gulf

of Maine by way of Northeast Channel; 7) tidal amplitude is exceptionally large in the Bay of Fundy region,

and tidal currents are strong throughout the entire

New England continental shelf area; 8) the Gulf Stream

flows northeastward in deep water south of the New

England continental shelf (usually the Gulf Stream's

northern edge is more than one hundred miles south

of the continental shelf in the region south of Nantucket Island); and 9) below the Gulf Stream in the

vicinity of the ocean bottom, the Western Boundary

Current flows southwestwardly.

Oceanic waters in the vicinity of the Gulf Stream

maintain a relatively constant salinity of about 35%0.

Most of the waters overlying the continental shelf have

a salinity range from 32 to 34%0. Salinity of inshore

waters, which are more strongly influenced by runoff,

fluctuate seasonally and drop to 28%0 in late spring

when river discharge is maximum.

Temperature of water in deep oceanic areas beyond

the continental shelf is typically homostenothermal.

Waters are warm (20°C) at the surface and cold at the

bottom (2.5 to 5°C), and both surface and bottom

temperatures remain relatively stable throughout the

year. Conversely,the inshore waters along the coast are

characteristically heteroeurythermal. They are cold

(O°C) in winter and warm in summer, and because of

the shallowness and general turbulence of the water,

the temperature differential between surface and bottom is relatively small. Also, there is considerable latitudinal effect on inshore waters; in southern areas the

temperature does not drop as low in winter and rises

higher in summer than it does in northern areas.

Midshelf waters-those between the oceanic and inshore zones-are generally intermediate in their temperature regime. Temperature diversity between the

surface and bottom is moderate. Seasonal changes in

temperature are greater in offshore shallow areas (such

as Nantucket Shoals and Georges Bank) than in basins

and other deep water areas, but the range is less than

that in coastal waters. Annual fluctuation in temperature of bottom water is considerably less than that of

surface waters. Latitudinal effect on shelf water masses

is pronounced; ova Scotian water is substantially colder

than other water masses within the study area, and the

temperature generally increases to the west and south

(Bigelow, 1933; McLellan, 1954; Edwards et aI., 1962;

Colton et aI., 1968; Schopf and Colton, 1966; Schopf,

1967; Colton, 1968a, 1968b, 1969; Colton and Stoddard,

1972, 1973; Mountain and Holzwarth, 1989; Colton et

al. 10; Colton et aJ.ll; Colton et al. 12; Colten et al. 13).

Thermal extremes, rather than means, are believed

to have a marked influence on the presence or absence

of various kinds of benthic animals. In order to detect

the possible influence of thermal extremes as a limiting

factor, we have analyzed the invertebrate fauna distribution in relation to the approximate annual minimum and maximum water temperatures, and the range

in water temperature, to which the various taxa are

10 Colton,]. B.,Jr., R. R. Marak, and S. R. Nickerson. 1965a. Environmental observations on continental shelf Nova Scotia to Long

Island, March 1965, Albatross IV cruise 65-3. U.S. Bur. Commer.

Fish. BioI. Lab. Woods Hole, Mass., Lab. Ref. 65-15, 3 p., 9 figs.

(mimeo). Unpubl. manuscript.

II Colton,]. B.,Jr., R. R. Marak, and S. R. Nickerson. 1965b. Environmental observations on continental shelf Nova Scotia to Long

Island, September 1965, Albatross IV cruise 65-12. U.S. Bur.

Commer. Fish. BioI. Lab. Woods Hole, Mass., Lab. Ref. 65-19, 3 p.,

9 figs. (mimeo). Unpubl. manuscript.

12 Colton,]. B.,Jr., R. R. Marak, and S. R. Nickerson. 1966a. Environmental observations on continental shelf Nova Scotia to Long

Island, March 1966, Albatross IV cruise 66-2. U.S. Bur. Commer.

Fish. BioI. Lab. Woods Hole, Mass., Lab. Ref. 66-6, 3 p., 10 figs.

(mimeo). Unpubl. manuscript.

I~l Colton, ]. B., Jr., R. R. Marak, S. R. Nickerson, and R. R.

Stoddard. 1966b. Environmental observations on continental

shelf Nova Scotia to Long Island, May-June 1966. Albatross IV

cruise 66-7. U.S. Bur. Cammer. Fish. BioI. Lab., Woods Hole,

Mass., Lab. Ref. 66-7, 3 p., 10 figs. (mimeo). Unpubl. manuscript.
16 NOAA Technical Report NMFS 140 _

subjected. Charts were constructed to illustrate the isotherms ofmaximum bottom water temperature (Fig. 9)

minimum bottom water temperature (Fig. 10), and

annual range in bottom water temperature (Fig. 11).

Data for these charts were extracted from temperature

records taken during the sampling period when biological data were collected, August 1956 through August 1965, and from the literature (see above citations).

Temperature patterns depicted in these charts are intended to provide a general scheme of annual temperature change. Higher or lower temperatures may have

existed for short periods in some areas and may have

been missed because of the opportunistic nature of the

sampling. Extremes of this kind, however, are not considered usual or of great magnitude.

These charts disclose a wide annual temperature range

in coastal bays and in shallow offshore areas, such as

Georges Bank and Nantucket Shoals. Very little change

occurs in deep water. At depths below 500 meters the

annual variation in temperature is roughly 0-3.9°C.

Bottom water in the Gulf ofMaine is relatively cold, 4 to

8°C and changes very little throughout the year. Bottom water on the Scotian Shelf and Browns Bank is

particularly cold in the spring and warms up only to

moderate levels in the fall and early winter. Annual

average temperature of bottom water for some of the

major areas calculated by Schopf and Colton (1966)

and Schopf (1967) are: Georges Bank 8.6°C, Nantucket

Shoals 7.8°C, Gulf of Maine 5.7°C, Browns Bank 5.0°C,

and the Nova Scotian Shelf 4.6°C.

Zoogeography

The topographic, hydrographic, climatic, and faunal

complexities of the sublittoral portion of the study area

cause considerable difficulty in the definition of definitive zoogeographic boundaries in the Northwest Atlantic. Until recently, the traditional view among biogeographers was that the region embraced portions of two

major zoogeographic provinces: 1) The Boreal Province, sometimes referred to as Acadian or Nova Scotian,

which extends from Newfoundland to Cape Cod, and

2) The Trans-Atlantic or (Warm Temperate) Province

72°

NEW

72

71°

NEW

JERSEY /' HAMPSHIRE /

/

: PORTLAND

71°

41°

70°

M A I N E

69"

o 120 L'_ .........--" KM

42°

8

6

4

4°

Figure 9

Distribution of maximum reponed bottom water temperatures (in degrees Celsius) in the study area.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 17

of which the Virginian subprovince extends from Cape

Cod southward to Cape Hatteras (Ekman, 1953;

Hedgpeth, 1957). Although these views postulated the

highly visible physical features of Cape Cod and Cape

Hatteras as the boundaries between these provinces (a

credible hypothesis topographically and hydrographically), no definitive consensus of opinion among biogeographers of the period prevailed as to the precise

placement of the boundaries in the orthwest Atlan tic.

Indeed, the plethora of varying definitions and terms

led to a rather confusing semantic problem that exists

to this day. Further, these views resulted from studies

based almost solely on biological and physical data

from inshore or nearshore areas.

Hazel (1970) reviewed the historical development of

faunal provinces for North America and Europe based

on the work of 17 authors from 1838 to 1966 and noted

that during that period essentially three biogeographic

schemes evolved to characterize the Northwest Atlantic

down to Cape Hatteras: 1) Cape Cod acts as a boundary

between the cold temperate Nova Scotian or Boreal

Province to the north, and the warm temperate Virginian subprovince to the south, with Cape Hatteras forming the boundary between the Virginian and Carolinian subprovinces, which together formed the TransAtlantic Province down to present day Cape Kennedy;

2) a region of overlap or transition, lacking a unique

fauna of its own (low endemism) with no provincial

status, between the Nova Scotian and Carolinian Provinces; and 3) A cold temperate Boreal Province extending from Newfoundland to Cape Hatteras.

Although more recent biogeographic studies, based

mostly on offshore fauna within the region, such as

those of Bousfield (1960), Coomans (1962), Schopf

(1968b), Franz (1970), Hazel (1970), Bousfield (1973),

Franz (1975), Bowen et al. (1979), Kinner (1978),

Watling (1979), Franz and Merrill (1980a, 1980b), and

Franz et al. (1981) have expressed concern over the

boundary's existence and have attempted to resolve the

semantic problem of terminology through revision and

simplification, they have not, for the most part, significantly altered the three biogeographic concepts of earlier workers. These recent works, however, have provided some new insights concerning the placement of

72' 71' 70' 69"

NEW 0 120 ,

/ 1 I KM JERSEY , NEW '\: HAMPSHIRE ,' . A N E / M I ,

"'-.. : PORTLAND ........ ,":

Figure 10

Distribution of minimum reported bottom water temperatures (in degrees Celsius) in the study area.
18 NOAA Technical Report NMFS 140

NEW

oERSEY

72"

ANNUAL RANGE IN BOTTOM

WATER TEMPERATURE (OC)

o 0- 39 71· EZZi 4- 7.9

8-11.9

o 12-159

• 16-19.9

• 20-239

38· 39·

72·

NEW /'

HAMPSHIRE .-

/

:. PORTLAND

40·

71·

4'·

70·

M A I N E

69·

o 120 ~I_,,--,I KM

42·

Figure II

Distribution of the annual range (difference between maximum and minimum reported values) in bottom water

temperature (in degrees Celsius) in the study area.

more meaningful zoogeographic boundaries for regulating the distribution of benthic taxa within the region.

Boundaries of the geographical area considered in

this report were purposely selected so that they did not

terminate at the margin of a perceived zoogeographical barrier. Cape Cod, lying roughly in the center of the

study area, is of course the main physical feature historically considered to mark the separation between

the Boreal and Trans-Atlantic Provinces. The recent

work ofSchopf (1968b), Hazel (1970), Watling (1979b),

Franz et al. (1981), and other reports (Wigley and

Burns, 1971; Williams and Wigley, 1977; Theroux and

Wigley, 1983; and Theroux and Grosslein, 1987) based

on the same data as, and including, the present report

corroborate the fact that Cape Cod is indeed a zoogeographic boundary. However, the seaward extension of

this boundary, at least as it pertains to benthic animals,

does not traverse the continental shelf over Nantucket

Shoals and the southwestern terminus of Great South

Channel as previously supposed. Rather, the boundary

appears to lie along an easterly path across the northern end of Great South Channel at depths of 50 to 100

m and to continue along the northern margin of

Georges Bank and thence southeasterly along the western boundary of Northeast Channel.

In bathyal and abyssal depths there are at least two

other zoogeographic provinces. Along the continental

slope, at depths between 150 and 2,000 m, is the Atlantic Transitional Province (Cutler, 1977), and at depths

between 2,000 and 4,000 m is the Atlantic BathyalAbyssal Province. Because of the interdigitating distributional patterns resulting from the southward submergence ofBoreal species and the ascendency ofTransitional and Bathyal-Abyssal species in their northward extension, the delineation of these provinces is

imprecise and only partially aligned with topographic

features.

A great deal more work ofa zoogeographic nature on

the many remaining unstudied taxa of benthic invertebrates inhabiting the area needs to be performed before precise zoogeographic boundaries may be drawn,

if at all possible.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 19

Table 3

Rank order of major taxonomic groups according to percentage

composition Df the total macrobenthic fauna in terms of biomass

and number ofspecimens.

cussed in more detail in the following sections. Table 4

lists the components of the macrobenthic invertebrate

fauna inhabiting the New England region, and Table 5

lists the quantitative measures of abundance (mean

and total weights and numbers per square meter), number of samples, and frequency of occurrence for each

taxonomic group considered in this report.

43.4

28.1

10.8

<1.0

82,3

2,9

1.9

1.7

1.5

1.5

1.2

U

1.0

12.8

0,8

0,5

0,5

0.4

0.4

0,3

0,3

0,3

0,2

0,2

0, I

0.1

0,1

0,1

4,3

<0,1

Percentage of

total number

specimens

Total

Ophiuroidea

Echinoidea

Cumacea

Zoantharia

Cirripedia

Gastropoda

Ascidiacea

Bryozoa

Total

Isopoda

Nemertea

Decapoda

Hydrozoa

Sipunculida

Brachiopoda

Scaphopoda

Holothuroidea

Nematoda

Mysidacea

Porifera

Alcyonaria

Polyplacophora

Asteroidea

Total

Turbellaria

Priapulida

Cephalopoda

Echinoidea

Arachnida

Pycnogonida

Ostracoda

Copepoda

Tanaidacea

Crinoidea

Pogonophora

Hemichordata

Echiura

Total

Amphipoda

Annelida

Bivalvia

Taxa

4,0

<0,1

<1.0

3,5

2,3

2,2

1.9

1.8

1.2

1.2

1.2

15.3

0,8

0.7

0.5

0.4

0.4

0,3

0,2

0,2

0,2

0,1

0,1

0, I

44.1

20,0

9,5

7,0

80.6

Percentage

of total

biomass

Bivalvia

Echinoidea

Annelida

Holothuroidea

Total

Taxa

Zoantharia

Amphipoda

Ascidiacea

Cirripedia

Ophiuroidea

Gastropoda

Asteroidea

Porifera

Total

Decapoda

Bryozoa

Brachiopoda

Nemertea

Sipunculida

Hydrozoa

Scaphopoda

Echiura

[sopoda

Alcyonaria

Polyplacophora

Cumacea

Total

Turbellaria

Priapulida

Nematoda

Cephalopoda

Arachnida

Pycnogonida

Ostracoda

Copepoda

Mysidacea

Tanaidacea

Crinoidea

Pogonophora

Hemichordata

Total

Dominance

IV

II

category

1lI

Faunal Composition

The macrobenthic invertebrate fauna of the

New England region is moderate in variety.

A modest number of species (567 in the

present study), in combination with a graded

abundance of individuals composed of a variety of dominants and codominan ts, is characteristic of the fauna, and is generally typical ofBoreal-Temperate faunal assemblages.

Taxa reported on in this study represent 13

phyla and 28 lesser groups such as subphyla,

classes, subclasses, and orders. The majority of

species are Boreal forms, followed closely in

abundance by Virginian (or warm-temperate)

forms. Additionally, there is a small contingent of

Arctic and Subarctic species, particularly in

the Gulf of Maine. Also, a few tropical and

subtropical species occur chiefly in the Southern New England and Georges Bank areas.

The ecological importance of these

groups,judged primarily from their numerical abundance and biomass, ranges from

minor (components that account for less

than 0.1 % in number ofindividuals and biomass) to dominant components that make

up 20% or more in number ofindividuals or

biomass. The 44 major taxonomic groups,

with the percentage of total number of individuals and percentage of total biomass for

each, are listed in Table 3. Also, they are classified into four dominance categories, I to IV.

Over 80% of both the biomass and number of individuals in the macrobenthos is

formed by only five taxonomic groups. These

are classified in dominance category I in

Table 3. Bivalvia is the dominant contributor (44.1 %) to the biomass and is also a

major component (10.8%) in terms ofnumbers ofindividuals. Amphipoda, on the other

hand, is numerically dominant (43.4%) but

contributes only 2.3% of the biomass. Conversely, Echinoidea and Holothuroidea are

important components of the biomass, but

are numerically sparse. Annelida is a major

contributor in both measures of quantity.

Category II, in Table 3, consists of eight

taxa that contribute moderate biomass (1.2

to 2.3% of the total fauna) and number of individuals

(1.0 to 2.9% of the total fauna). Categories III and IV

contain those taxa that contribute small to very small

quantities to the total biomass and density.

The New England region macrobenthos is dominated

by members of four phyla: Annelida, Mollusca, Arthropoda, and Echinodermata. These groups will be dis-
20 NOAA Technical Report NMFS 140

Table 4

List of macrobenthic invertebrate species contained in quantitative samples obtained within the study area.

PORIFERA

Demospongiae

Hadromerida

Suberitidae

Polymastia sp.

COELENTERATA

Hydrozoa

Hydractinia echinata Fleming, 1828

Hydractinia sp.

Anthozoa

Alcyonaria

Alcyonaeea

Alcyonium sp.

Gorgonacea

Acanella sp.

Paragorgia arborea (Linnaeus, 1767)

Primnoa reseda (Pallas, 1766)

Pennatulaeea

Pennatula aculeata Danielssen and Koren, 1858

Pennatula sp.

Stylatula elegans (Danielssen, 1860)

Zoantharia

Zoanthidea

Epizoanthus inerustatus (Verrill, 1864)

Epizoanthus sp.

Aetiniaria

Tealina felina (Linnaeus, 1767)

Edwardsia mlcata (T. Pennant, 1777)

Edwardsia sp.

Aetinostola callosa Verrill, 1882

Antholoba perclix (Verrill, 1882)

Madreporaria

Astrangia sp.

Flabellum goodei Verrill, 1878

Flabellum sp.

Ceriantheria

Cerianthus borealis Verrill, 1878

Cerianthus sp.

Ceriantheopsis mnericanus Verrill, 1866

Annelida

Polyehaeta

Amphinomida

Amphinomidae

Paramphinome jejJreysii (McIntosh, 1868)

Capitellida

Capitellidae

Capitella sp.

MaJdanidae

Asychis biceps (Sars, 1861)

Maldanesp.

Cossurida

Cossuridae

Cossura longicinata Webster and Benedict, 1883

Cossura sp.

Eunicida

Arabellidae

Arabella irieolor (Montaf(u, 1804)

Ambella sp.

Drilonereis longa Webster, 1879

Drilonereis sp.

Notocinus sp.

Eunieidae

Eunice pennata (Muller, 1776)

Eunieesp.

Marphysa sp.

Lumbrineridae

Lumbrinerides awta (Verrill, 1875)

Lumbrineris fmgilis (Muller, 1776)

Lumbrineris sp.

Ninoesp.

Onuphidae

Diopatra wprea (Bose, 1802)

Diopatra sp.

Hyalinoeeia tubieola (Muller, 1776)

Hyalinoecia sp.

Nothria eonehylega Sars, 1835

Onuphis eremita Audoin and Milne-Edwards, 1833

Onuphis opalina (Verrill, 1873)

Onuphis quadriwspis Sars, 1872

Onuphis sp.

Paradiopatra sp.

Flabelligerida

Flabelligeridae

Brada sp.

Flabelligera sp.

Pherusasp.

Opheliida

Opheliidae

Ophelia sp.

Ophelina aulogaster (H. Rathke, 1843)

Ophelina sp.

Travisia carnea Verrill, 1873

Travisia sp.

SeaJibregmidae

Scalibregma inflatum Rathke, 1843

Scalib'regma sp.

Orbiniida

Orbiniidae

Orbinia ornata (Verrill, 1873)

Orbinia swani Pettibone, 1957

Orbinia sp.

Seoloplos mbustus (Verrill, 1873)

Seoloplos sp.

Arieidea jeJJreysii (McIntosh, 1879)

A ricidea sp.

Paraonidae

Paraonis sp.

Oweniida

Oweniidae

Owenia fusi/ormis delle Chiaje, 1844

Oweniasp.

Phyllodoeida

Aphroditidae

Aphr-odita hastata Moore, 1905

Aphrodita sp.

Laetmonice sp.

Glyeeridae

Clyeem americana Leidy, 1855

Clyeem capitata Oersted, 1843

Glyeera dibranehiata Ehlers, 1868

Clycera sp.

conlinued on next page
Composition and Distribution of Macrobenthic Invertebrate Fauna 21

Table 4 (continued)

Goniadidae

Goniada maculata (Oersted, 1843)

Goniada sp.

Goniadella sp.

Ophioglycem gigantea Verrill, 1885

Ophioglycem sp.

Hesionidae

Nereimym punctata (O.F. Mliller, 1776)

Nephtyidae

Aglaophamus circinata (Verrill, 1874)

Aglaophamus sp.

Nephtys bucera Ehlers, 1869

Nephtys incisa Malmgren, 1865

Nephtys picta Ehlers, 1868

Nephtys sp.

Nereidae

Ceratocephale loveni Malmgren, 1867

Ceratocephale sp.

Nereis sp.

Phyllodocidae

Eteone sp.

Eumida sanguinea (Oersted, 1843)

Phyllodoce arenae Webster, 1879

Phyllodoce sp.

Pilargiidae

Ancistrosyllis sp.

Polynoidae

Harrnothoe sp.

Lepidonotus squamatus (Linnaeus, 1758)

Sigalionidae

Leanira sp.

Pholoe minuta (Fabricius, 1780)

Sigalion arenicola Verrill, 1879

Sigalion sp.

Sphaerodoridae

Sphaerodorum gracilis (Rathke, 1843)

SyJlidae

Exogoneverugera (Clarapede, 1868)

Exogone sp.

Tomopteridae

Tomopteris sp.

Sabellida

Sabellidae

Chone inJundibuliformis Kr0yer, 1856

Chone sp.

Euchone sp.

Potamilla neglecta (Sal's, 1850)

Potamilla reniformis (Linnaeus, 1788)

Potamilla sp.

Sabella sp.

Serpulidae

Filograna sp.

Spirorbidae

Spir01bis sp.

Spionida

Chaetopteridae

Spiochaetopterus sp.

Cirratulidae

Chaetozone sp.

Cirratulus sp.

Tharyx sp.

Spionidae

Diospio uncinata Hartman, 1951

Laonice cirrada (Sal's, 1851)

Laonice sp.

Polydora concharum Verrill, 1880

Polydora sp.

Priospio sp.

Spio setosa Verrill, 1873

Spio sp.

Spiophanes bombyx (Clarapede, 1870)

Sternaspida

Sternaspidae

Sternaspis scutata (Renier, 1807)

Sternaspis sp.

TerebeJlida

Ampharetidae

Ampharete acutifrons (Grube, 1860)

Ampharete sp.

Melinna cristata (Sal's, 1851)

Melinna sp.

Pectinariidae

Pectinaria gouldii (Verrill, 1873)

Pectinaria sp.

Terebellidae

Amphitrite sp.

Streblosoma spimlis (Verrill, 1874)

Steblosoma sp.

POGONOPHORA

Siboglinidae

Siboglinum angustum Southward and Brattegard, 1968

Siboglinum atlanticum Southward and Southward, 1958

Sibogliml1n ekmaniJagerston, 1956

Siboglinum hohnei Sou thward ,1963

Siboglinum pholidotum Southward and Brattegard, 1968

Siboglinum sp.

Polybrachiidae

Crassibranchia sandersi Southward, 1968

Diplobmchia similis Southward and Brattegard, 1968

Polybmchia sp.

SIPUNCULIDA

Aspidosiphon zinni Cutler, 1969

Colfingia catharinae (Muller, 1789)

Colfingia elongata (Keferstein, 1869)

GolJingia eremita (Sal's, 1851)

GolfingiaflagriJera (Selenka, 1885)

Golfingia margaritacea (Sal's, 1851)

Colfingia minuta (Keferstein, 1865)

Colfingia murinae murinae Cutler, 1969

Onchnesoma steenstrupi Koren and Danielssen, 1875

Phascolion strombi (Montague, 1804)

Phascolopsis gouldi (Pourtales, 1851)

Sipunculus norvegicus Koren and Danielssen, 1875

ECHIURA

Bonellia thomensis (Gmelin, 1788)

t;chiurus echiurus (Pallas, 1774)

£Chiurus sp.

Ikedella akaeta (Zenkevitch, 1958)

Maxmuelleria lankesteri (Herdman, 1898)

Prometor grandis (Zenkevitch, 1957)

Protobonellia sp.

Sluiterina sibogae (Sluiter, 1902)

SluiteTina sp. continued on next page
22 NOAA Technical Report NMFS 140

Table 4 (continued)

MOLLUSCA

Polyplacophora

Gastropoda

Prosobranchia

Archaeogastropoda

Fissurelidae

Puncttnella noachina (Linnaeus, 1771)

Lepetidae

Lepeta caeca (Muller, 1776)

Trochidae

Calliostoma occidentalis (Mighels and Adams,

1842)

Margarites costalis (Gould, 1841)

Margarites groenlandicus (Gmelin, 1791)

Margarites helicinus (Phipps, 1774)

Margarites sp.

Solariella lamellosa Verrill and Smith, 1880

Solariella obscum (Couthouy, 1838)

Solariella sp.

Mesogastropoda

Littorinidae

Littorina obtusata (Linnaeus, 1758)

Rissoidae

Alvania brychia (Verrill, 1884)

Alvania pelagica (Stimpson, 1851)

Alvania areolata Stimpson, 1851

Alvania sp.

Turritellidae

Tachyrhynchus croms (Couthouy, 1838)

Turritellopsis acicula (Stimpson, 1851)

Cerithiidae

Cerithiella sp.

Diastoma alternatus (Say, 1822)

Epitoniidae

Epitonium dallianum Verrill and Smith, 1880

Epitonium greenlandicum (Perry, 1811)

Melanellidae

Couthouyella striatula (Couthouy, 183Y)

Aclididae

Adis verrilli Bartsch, 1911

Trichotropidae

Trichotropis bonalis Broderip and Sowerby, 1829

Crepidulidae

Crepidula fomicata Linnaeus, 1767

Crepidula plana Say, 1822

Crucibulurn stTiaturn Say, 1824

Aporrhaidae

Aporrhais occidentalis Beck, 1836

Velutinidae

Velutina velutina (Mliller, 1776)

Velutina undata (Brown, 1839)

Velutina sp.

Naticidae

Lunatia heros (Say, 1822)

Lunatia triscriata (Say, 1826)

Lunatia pallida (Broderip, and Sowerby, 1829)

Lunatiasp.

Natica clausa Broderip and Sowerby, 1829

Natica pusilla Say, 1822

Polinices duplicatus (Say, 1822)

Polinices immaculatus (Totten, 1835)

Polinices sp.

Neogastropoda

Muricidae

Boreotmp/wn clathratus (Linnaeus, 1758)

Eupleura caudata (Say, 1822)

Columbellidae

Arnphissa haliaeeti Oeffreys, 1867)

Anachis lafresnayi (Fischer and Bernardi, 1856)

Anachis sp.

Mitrella lunata (Say, 1826)

MitTella pum (Verrill, 1882)

Mitrella rosacea (Gould, 1841)

MitTella sp.

Buccinidae

Buccinurn undatum Linnaeus, 1758

Buccinum sp.

Colus caelatus (Verrill and Smith, 1880)

Colus obesus (Verrill, 1884)

Colus parous (Verrill and Smith, 1882)

Colus pygmaeus (Gould, 1841)

Colus sp.

Neptunea decerncostata (Say, 1826)

Neptunea despecta (Linnaeus, 1758)

Neptunea sp.

Melongenidae

Busycon canaliculatus (Linnaeus, 1758)

Nassariidae

llyanassa obsoleta (Say, 1822)

NassaTius trivittatus (Say, 1822)

Nassarius vibex (Say, 1822)

Cancellariidae

Adrnete couthouyi Oay, 1839)

Turridae

Oenopota decussata (Couthouy, 1839)

Oenopota harpularia (Couthouy, 1838)

Oenopota incisula (Verrill, 1882)

Pleurotornella agassizi agassizi Verrill and Smith,

1880

Pleumtornella blakeana (Dall, 1889)

Pleurotemella curta cUTta (Ve rriII, 1884)

Pteurotornella packaTdi packardi (Verrill, 1872)

Propebela elegans (Moller, 1842)

Propebela examta (Moller, 1842)

Propebela turricula (Montagu, 1803)

Pyramidellidae

Odostornia dealbata (Stimpson, 1851)

Odostornia dux Dall and Bartsch, 1906

Odostomia sp.

TUl·bonilla bushiana Verrill, 1882

Turbonilla elegantula Verrill, 1882

Turbonilla nivea (Stimpson, 1851)

Turbonilla polita (Verrill, 1872)

Tm·bonilla sp.

Opisthobranchia

Acteonidae

Acteon sp.

Ringiculidae

Ringicula nitida Verrill, 1873

Actiocinidae

Acteocina canaliculata (Say, 1822)

Retusa obtusa (Montagu, 1807)

continued on next page
Composition and Distribution of Macrobenthic Invertebrate Fauna 23

Table 4 (continued)

Scaphandridae

Cylichna alba (Brown, 1827)

Cylichna gouldi (Couthouy, 1839)

Cylichna vortex (Dall, 1881)

Cylichna sp.

Seaphander punctostriatus Mighels, 1841

Philinidae

Philine lima (Brown, 1827)

Philine quadrata (S. Wood, 1839)

Philinesp.

Akeridae

Haminoea sp.

Pleurobranchidae

Pleurobranchaea sp.

Nudibranchia

Dendronotidae

Dendronotus jrondosus (Ascanius, 1774)

Bivalvia

Palaeotoxodon ta

Nuculoida

Nuculidae

Nucula delphinodonta Mighels and Adams, 1842

Nucula proxima Say, 1822

Nucula tenuis Montagu, 1808

Nuculasp.

Malletiidae

Malletia obtusa G.O. Sars, 1872

Satumia subovata Verrill, and Bush, 1897

Nuculanidae

Nuculana acuta (Conrad, 1831)

Nuculana pemula (Muller, 1771)

Nuculana tenuisuleata (Couthouy, 1838)

Nuculana sp.

Portlandia jratema (Verrill and Bush, 1898)

Portlandiafrigida (Torrell, 1859)

Portlandia inconspicua (Verrill and Bush 1898)

Portlandia inJlata (Verrill, and Bush, 1897)

Portlandia iris (Verrill and Bush, 1897)

Portlandia lenticula (Moller, 1842)

Portlandia lucida (Loven, 1846)

Yoldia limatula (Say, 1831)

Yoldia myalis (Couthouy, 1838)

Yoldia regularis Verrill, 1884

Yoldia sapotilla (Gould,1841)

Yoldia thraciaejorrnis Storer, 1838

Yoldiasp.

Cryptodonta

Solemyoida

Solemyacidae

Solemya velum Say, 1822

Pteriomorphia

Arcoida

Arcidae

Anadara ovalis (Bruguiere, 1789)

Anadara transversa (Say, 1822)

Bathyarea anomala (Verrill and Bush, 1898)

Bathyarea pectunculoides (Scacchi, 1833)

Bathyarca sp.

Limopsidae

Limopsis ajfinis Verrill, 1885

Limopsis cristataJeffreys, 1876

Limopsis minuta Philippi, 1836

Limopsis suleata Verrill and Bush, 1898

Limopsis sp.

Mytiloida

Mytilidae

Crenella decussata (Montagu, 1808)

Crenella glandula (Totten, 1834)

Crenella sp.

Dacrydium vitreum (Holboll in Moller, 1842)

Modiolus modiolus (Linnaeus, 1758)

Musculus corrugatus (Stimpson, 1851)

Musculus discors (Linnaeus, 1767)

Musculus niger (Gray, 1824)

Musculus sp.

Mytilus edulis Linnaeus, 1758

Mytilussp.

Pterioida

Pectinidae

Chlamys islandiea (Muller, 1776)

Cydopecten pustulosus Verrill, 1873

Placopecten magellanicus (Gmelin, 1791)

Anomiidae

Anommia simplex Orbigny, 1842

Anomia squamula Linnaeus, 1758

Anomia sp.

Limidae

Limatula subauriculata (Montagu, 1808)

Limatula sp.

Heterodonta

Veneroida

Lucinidae

Lucinoma blakeana Bush, 1883

Lucinoma jilosa (Stimpson, 1851)

Lucinoma sp.

Thyasiridae

Thyasira equalis Verrill and Bush, 1898

Thyasira jerruginea Winckworth, 1932

ThyasiraJlexuosa (Montagu, 1803)

ThyasiraJlexuosa forma gouldii Philippi, 1845

Thyasira pygrnaea Verrill and Bush, J898

Thyasira subovataJeffreys, 1881

Thyasira trisinuata Orbigny, 1842

Thyasira sp.

Lasaeidae

Aligenaelevata (Stimpson, 1851)

Leptonidae

l\1ontacuta sp.

Carditidae

Cydocardia borealis Conrad, 1831

Cyclocardia sp.

Astartidae

Astarte borealis (Schumacher, 1817)

Astarte castanea (Say, 1822)

Astarte crenata subequilatera Sowerby, 1854

Astarte elliptiea (Brown, 1827)

Astarte montagui (Dillwyn, 1817)

Astarte nana Dall, 1886

Astarte quadrans Gould, 1841

Astarte smithii Gould, 1841

Astarte undata Gould, 1841

Astarte sp.

continued on next page
24 NOAA Technical Report NMFS 140

Table 4 (continued)

continued on next page

Cardiidae

Cerastodenna pinnulatum (Conrad, 1831)

Laevicardium mortoni (Conrad, 1830)

Mactridae

Mulinia lateralis (Say, 1822)

Spisula solidissima (Dillwyn, 1817)

Mesodesmatidae

Mesodesma sp.

Solenidae

Ensis directus Conrad, 1843

Siliqua costata Say, 1822

Tellinidae

Macoma calcarea (Gmelin, 1791)

Macoma sp.

Tellina agilis Stimpson, 1858

Tellina sp.

Arcticidae

Arctica islandica (Linnaeus, 1767)

Veneridae

Gemma gemma (Totten, 1834)

Ml!1"C1!naria mercenaria (Linnaeus, 1758)

Pitar morrhuanus Linsley, 1845

Myoida

Myidae

Mya arenaria Linnaeus, 1758

Corbulidae

Corbula contracta C.B. Adams, 1852

Corbula sp.

Hiatellidae

Cyrtodaria szliqua (Spengler, 1793)

Hiatella arctica (Linnaeus, 1767)

Hiatella striata (Fleuriau, 1802)

Hiatella sp.

Panomya arctica (Lamarck, 1818)

Panomya sp.

Anomalodesmata

Pholadomyoida

Pandoridae

Pandora gouldiana Dall, 1886

Pandora inflata Boss and Merrill, 1965

Pandora ino'rnata Verrill and Bush, 1898

Pandorasp.

Lyonsiidae

Lyonsia arenosa Moller, 1842

Lyonsia hyalina Conrad, 1830

Lyonsia sp.

Periplomatidae

Periplomafragile (Totten, 1835)

Periploma leanum (Conrad, 1830)

Periploma papyratium (Say, 1822)

Periploma sp.

Thraciidae

Thracia myopsis Moller, 1842

Thracia sp.

Cuspidariidae

C01'diomya permstrata (Dall, 1881)

Cuspidaria glacialis (G.O. Sars, 1878)

Cuspidaria obesa (Loven, 1846)

Cuspidaria paroa Verrill and Bush, 1898

Cuspidaria pellucida Stimpson, 1853

Cuspidaria sp.

Scaphopoda

Dentaliidae

Batlwxiphus ensieulus Oeffreys, 1877)

Cadulus agassizii Dall, 1881

Cadulus cylindratusJeffreys, 1877

CaduluspandionisVerrill and Smith, 1880

Cadulus rushii Pilsbry and Sharp, 1898

Cadulussp.

Dentalium entale stimpsoni Henderson, 1920

Dentalium meTidionale Pilsbry and Sharp, 1897

Dentalium occidentale Stimpson, 1851

Dentalium sp.

Cephalopoda

Octopus sp.

Rossia sp.

ARTHROPODA

Pycnogonida

Achelia scabra Wilson, 1880

Ache/ia spinosa (Stimpson, 1853)

Anoplodaetylus lentus Wilson, 1878

Nymphon brevitarse Kr0yer, 1844

Nyrnphon grossipes (0. Fabricius?) Kr0yer, 1780

Nymphon hirtipes Bell, 1853

Nymphon macrum Wilson, 1880

Nymphon stroerni KJ'0yer, 1844

Paranymphon spinosurn Caullery, 1896

Pycnogonium liUorale (Strom, 1762)

Crustacea

Cirripedia

Balanus sp.

Lepas sp.

Copepoda

Calanus sp.

Caligussp.

Cumacea

Brachydiastylis resima (Kr0yer, 1846)

Campylaspis affinis Sars, 1870

Campylaspis rubieunda (Lilljeborg, 1855)

Cyclaspis longicaudata G.O. Sars, 1864.

Diastylis cornuifer (Blake, 1929)

Diastylis polita S.l. Smith, 1879

Diastylis quadris!Jinosa G.O. Sars, 1871

Diastylis rathkei (Kr0yer, 1841)

Diastylis seulpta G.O. Sars, 1871

Diastylis sp.

Eudorella emarginata (Kr0yer, 1846)

Eudorella hispida Sars, 1871

Eudorella truncatula (Bate, 1855)

Eudorella sp.

Eudorellopsis deformis (Kr0yer, 1846)

Eudorellopsis sp.

Hemilampraps cristata (Sars, 1870)

Lampraps quadriplicata S.1. Smith, 1879

Lamprapsfuscata Sars, 1865

Larnpraps sp.

Leptostylis longimana (Sars, 1865)

Leptostylis rnacrura G.O. Sars, 1869

Leptostylis sp.

Leucon americanus Zimmer, 1943

Leucon nasicoides LiIUeborg, 1855

Oxymstylis smithi CaIman, 1912
Composition and Distribution of Macrobenthic Invertebrate Fauna 25

Table 4 (continued)

Cumacea, continued

Petalosarsia dedivis (G.O. Sars, 1865)

Pseudoleptowma minor (Caiman, 1912)

Isopoda

Calathura branchiata (Stimpson, 1853)

Calathura sp.

Chiridotea arenicola Wigley, 1960

Chiridotea tuftsi (Stimpson, 1883)

Chiridotea sp.

Cirolana concharum (Stimpson, 1853)

Cirolana impressa (Harger, 1818)

Cirolana polita (Stimpson, 1853)

Cirolana sp.

Cyathurapolita (Stimpson, 1855)

C)'athura sp.

Edotea awta (Richardson, 1905)

Edotea triloba (Say, 1818)

Erichsonellafiliformis (Say, 1818)

Idotea phosphorea (Harger, 1873)

Janira alta (Stimpson, 1853)

Pseudarachna sp.

Ptilanthum te/lUis (Harger, 1879)

Ptilanthura sp.

Amphipoda

Caprellidea

Caprellidae

Aeginina longicornis (Kr0yer, 1842-43)

Aeginina sp.

Cap/'ella linemis (Linnaeus, 1767)

Caprella penantis Leach, 1814

Cap/'ella septentrionalis Kr0yer, 1838

CapTella unica Mayer, 1903

Caprella sp.

Luconacia incerta Mayer, 1903

Mayerella limicola Huntsman, 1915

Proaeginina norvegica (Stephensen, 1931)

Hyperiidea

Hyperiidae

Hype/iasp.

Pamthemisto gaudichaudii (Milne-Edwards,

1840)

Parathemisto sp.

Phronima sedentaria (Forskal, 1775)

Vibilia sp.

Gammaridea

Acanthonotozomatidae

Acanthonotozuma serratum (Fabricius, 1780)

Ampeliscidae

Ampelisea abdita Mills, 1964

Ampelisca agassizi Uudd, 1896)

Ampelisea dedivitatus Mills, 1967

Ampelisea eschrichti Kr0yer, 1842

Ampelisea macrocephala Lilljeborg, 1852

Ampelisca vadOTum Mills, 1963

Ampelisca verrilli Mills, 1967

Ampelisca sp.

Byblis gaimaTdi (Kr0yer, 1846)

Byblis serrata (Smith, 1873)

Byblis sp.

Haploops tubicola Lilljeborg, 1856

Haploops sp.

Amphilochidae

Amphilochoides odontyx (Boeck, 1871)

Gitanopsis arctiea G.O. Sars, 1895

Amphithoidae

Amphilhoe rubricala (Montagu, 1808)

Aoridae

Lembos smithi Holmes, 1905

Lembos sp.

Leptocheirus pinguis (Stimpson, 1853)

Leptocheirus sp.

Microdeutopis gryllotalpa Costa, 1853

Pseudunciola obliqua (Shoemaker, 1949)

Unciola dissimilis Shoemaker, 1945

Unciola inermis Shoemaker, 1945

Unciola irrorata Say, 1818

Unciola leucopis (Kr0yer, 1845)

Unciola serrala Shoemaker, 1945

Unciola spicata Shoemaker, 1945

Unciola sp.

Argissidae

Argissa hamatipes (Norman, 1869)

Bateidae

Batea catharinensis Muller, 1865

Calliopiidae

Calliopius laeviusculttS (Kr0yer, 1838)

Halirages fulvocinctus (M. Sars, 1854)

Haliragoides inermis (G.O. Sars, 1882)

Hippomedon serratus Holmes, 1905

Corophiidae

Corophium crassicorne Bruzelius, 1859

Corophium insidiosum Crawford, 1937

Corophium volutator (Pallas, 1766)

Corophium sp.

Siphonocetes smithianus Rathbun, 1908

Siphonocetes sp.

Eusiridae

EttSirus cuspidatus Kr0yer, 1845

Rhachotropis distincta (Holmes, 1908)

Rhaclwtropis inflata (G.O. Sars, 1882)

Rhachotropis owlala (Hansen, 1887)

Gammarus annulatus Smith, 1873

Gammarus pallustTis Bousfield, 1969

Gammarus sp.

HausLOriidae

Aeanthohaustorius intermedius Bousfield, 1965

Aeanthohaustorius millsi Bousfield, 1965

AcanthohaustorittS similis Frame, 1980

Acanthohaustorius spinosttS Bousfield, 1962

Haustorius arenarius (Slabber, 1769)

Haustorius sp.

Parahaustorius attenuatus Bousfield, 1965

PamhaustoTius holmesi Bousfield, 1965

Parahaustorius longimerus Bousfield, 1965

Protohaust01ius deichmannae Bousfield, 1965

Protohaustorius wigleyi Bousfield, 1965

Pseudohaustorius borealis Bousfield, 1965

Ischyroceridae

Ericthonius bTasiliensis (Dana, 1853)

Ericthonius rubricornis Smith, 1873

Ericthonius sp.

Ischyroceros anguipes Kr0yer, 1838

continued on next page
26 NOAA Technical Report NMFS 140

Table 4 (continued)

Ischyroceridae, continued

Ischyroceros megacheir (Boeck, 1871)

Ischyroceros sp.

Lysianassidae

Anonyx debruynii Hoek, 1882

Anonyx lilljeborgi Boeck, 1871

Anonyx nugax (Phipps, 1774)

Anonyx sarsi Steele and BruneI, 1968

Anonyxsp.

Hippomedon propinguus Sars, 1895

Hippomedon serratus Holmes, 1905

Hippomedon sp.

Lysianopsis alba Holmes, 1905

Orchomene groenlandica (Hansen, 1887)

Orchomene minuta Kr0yer, 1846

Orchomene pinguis Boeck, 1861

Orchomene sp.

Psammonyx nobilis Stimpson, 1853

Tmetonyx cicada O. Fabricius, 1780

Tmetonyx sp.

Tryphosella nanoides Lilljeborg, 1865

Melitidae

Casco bigelowi (Blake, 1929)

Eriopisa elongata (Bruzelius, 1859)

Maera danae Stimpson, 1853

Maera loveni (Bruzelius, 1859)

Maerasp.

Melita dentata Kr0yer, 1842

Melita palmata (Montagu, 1894)

Melitasp.

Melphidippidae

Melphidippa goesi Stebbing, 1899

Melphidippa sp.

Oedicerotidae

Bathymedon sausserei (Boeck, 1871)

Monoculodes edwardsi Holmes, 1908

Monoculodes intermedius Shoemaker, 1930

Monoculodes latimanus (Goes, 1866)

Monoculodes longicomis (Boeck, 1871)

Paroediceros sp.

Synchelidium americanum Bousfield, 1973

Westwoodilla megalops (G.O. Sars, 1882)

Paramphithoidae

Epimeria loricata G.O. Sars, 1879

Pardaliscidae

Halice abyssi Boeck, 1871

Pardalisca cuspidata Kr0yer, 1842

Photidae

Gammaropsis maculatus Oohnson, 1827)

Photis dentata Shoemaker, 1945

Photis macrocoxa Shoemaker, 1945

Photis reinhardi Kr0yer, 1842

Photissp.

Podoceropsis nitida (Stimpson, 1853)

Protomediafasciata Kr0yer, 1842

Pleustidae

Neopleustes pulchellus Kr0yer, 1846

Parapleustes sp.

Pleustes panoplus Kr0yer, 1838

Pleustes glaber Boeck, 1861

Stenopleustes gracilis Holmes, 1905

Stenopleustes inermis Shoemaker, 1949

Stenopleustes latipes (M. Sars, 1895)

Podoceridae

Dyopedus articus (Murdoch, 1884)

Dyopedus monacantha (Metzger, 1875)

Dulichia porrecta (Bate, 1857)

Dulichia tuberculata Boeck, 1870

Dulichia sp.

Paradulichia typica Boeck, 1870

Paradulichia sp.

Pon togeneidae

Pontogeneia inennis (Kr0yer, 1842)

Pontogeneia sp.

Pontoporeiidae

Amphiporea gigantea Bousfield, 1973

Amphiporea lawrenciana Shoemaker, 1929

Amphiporea virginiana Shoemaker, 1933

Bathyporeia quoddyensis Shoemaker, 1949

Stegocephalidae

Anadaniopsis nordlandica (Boeck, 1871)

Stegocephalus inflatus Kr0yer, 1842

Stenothoidae

Stenothoe minuta Holmes, 1905

Stenula peltata (Smith, 1873)

Synopiidae

Syrrhoe crenulata Goes, 1866

Syrrhoe spiniferum (Stimpson, 1853)

Mysidacea

Erythrops erythrophthalma (Goes, 1864)

Erythrops sp.

Mysidopsis bigelowi Tattersall, 1926

Neomysis americana (S.l. Smith, 1873)

Neomysis sp.

Decapoda

Caridea

Crangon septemspinosa Say, 1818

Dichelopandalus leptocerus (Smith, 1881)

Eualus pusiolus (Kr0yer, 1841)

Pandalus montagui Leach, 1813 or 1814

Pandalus propinquus G.O. Sars, 1869

Pandalus sp.

Pontophilus brevirostris Smith, 1881

Spirontocaris lilljeborgii (Danielssen, 1859)

Spirontocaris sp.

Astacidea

Homarus americanus H. Milne-Edwards, 1837

Anomura

Axius serratus Stimpson, 1852

Callianassa atlantica Rathbun, 1926

Callianassa biformis Biffar, 1971

Calocaris templemani (Squires, 1965)

Catapagurus gracilis (Smith, 1881)

Munida iris A. Milne-Edwards, 1880

Munida valida Smith, 1883

Pagurus acadianus Benedict, 1901

Pagurus annulipes Stimpson, 1860

Pagurus arcuatus Squires, 1964

PaguruslongicarpusSay, 1817

Pagurus politus (Smith, 1882)

Pagurus pollicaris Say, 1817

Pagurus pubescens Kr0yer, 1838

continued on next page
Composition and Distribution of Macrobenthic Invertebrate Fauna 27

Table 4 (continued)

Anomura, continued

Upogebia afJinis (Say, 1817)

Brachyura

Cancer borealis Stimpson, 1859

Cancer irroratus Say, 1817

Euprognatha rastellifera Stimpson, 1871

Geryon quinquedens Smith, 1897

Hexapanopeus anf!:Ustifrons (Benedict and

Rathbun, 1891)

Hyas coaretatus Leach, 1815

Hyas sp.

Libinia dubia H. Milne-Edwards, 1834

Libinia sp

Neopanope texana sayi (Smith, 1869)

Ocypode sp.

Pelia mutica (Gibbes, 1850)

Pinnixa chaetopterana Stimpson, 1860

Pinnixa sayana Stimpson, 1860

Pinnixasp.

BRYOZOA

Ctenostomata

Alcyonidiidae

Alcyonidium mamillalum Alder, 1857

Alcyonidium sp.

Fl ustrellidae

Fluslrellidra sp.

Cyclostomata

Crisiidae

Crisia cribraria Stimpson, 1853

Crisia denticulata (Lamarck, 1816)

Oncousoeciidae

Oncollsoecia canadensis Osburn, 1933

Oncousoecia diasloporoides (Norman, 1868)

Oncousoecia sp.

Tubuliporidae

fdmonea alianlicaJohnston, 1847

fdmoneasp.

Tubulipora liliacea (Pallas, 1766)

Tubulipora lobulala Hassall, 1841

Tubulipora sp.

Diaperoeciidae

Diaperoecia harmeri Osburn, 1933

Diplosolen obelium (Johnston, 1838)

Entalophora sp.

Frondiporidae

Defrancia sp.

Cheilostomata

Alderinidae

Amphibleslrumflemingii (Busk, 1854)

Amphiblestrum trifolium (Searles Wood, 1850)

Cauloramphus cymbaeformis (Hincks, 1887)

Bugulidae

Bugula elongata Nordgaard, 1906

Bugula lumla (Desor, 1848)

Bugulasp.

Dendrobeania murrayana (Johnston, 1847)

Dendrobeania sp.

Calloporidae

Callopora aurita (Hincks, 1877)

Callopora cralicula (Alder, 1857)

Callopora lineala (Linnaeus, 1767)

Callopora whileavesi Norman, 1903

Callopora sp.

Cellariidae

Cellaria sp.

Celliporidae

Cellepora canaliculala Busk, 1884

Cellepora sp.

Cheilopori nidae

Cl'yplosula pallasiana (Moll, 1803)

Cribrilinidae

Cribrilina punctala (Hassall, 1842)

Cribrilina sp.

Electridae

Electra hastingsae Marcus, 1938

Electra pilosa (Linnaeus, 1767)

PJripora calenularia (Jameson, 1814)

Eucrateidae

Eucralea loricata (Linnaeus, 1758)

Hippoporinidae

Hippodiplosia americana (Verrill, 1875)

Hippodiplosia pertusa (Esper, 1794-97)

Hippoponella hippo-pus (Smitt, 1867)

Hippothoidae

Hippolhoa hyalina (Linnaeus, 1767)

Membrani poridae

Conopeum reticulum (Linnaeus, 1767)

Membranipora sp.

Microporellidae

Microporella ciliala (Pallas, 1766)

Mucronellidae

Mucronella immersa (Fleming, 1847)

Mucronella ventricosa (Hassall, 1842)

Palmicellaria skenei (Ellis and Solander, 1786)

Porella plana (Dawson, 1859)

Parella proboscidea Hi ncks, 1888

Porella propinqua (Smitl, 1867)

Pseudoflustra sp.

Rhamphoslomella bilaminala (Hincks, 1877)

Rhamphoslomella ovata (Smitt, 1867)

Rhamphoslomella sp.

Smittina bella (Busk, 1860)

Smittina reduplicata Osburn, 1933

Smittina rigida Lorenz, 1886

Smittina lrispinosa (Johnston, 1838)

Smittinasp.

Schizoporellidae

Schizomavella auriculala (Hassa!l, 1842)

Schizoporella biaperla (Michelin, 1841-42)

Schizoporella unicornis (Johnston, 1847)

Scrupariinidae

Haplola clavata (Hincks, 1857)

Scrupocellariidae

Caberea ellisii (Fleming, 1828)

Scrllpocellaria scabra (Fabricius, 1780)

Tricellaria gracilis (Van Beneden, 1848)

Tricellaria sp.

Gigan toporidae

Tessaradoma gracile (M. Sars, 1851)

Stomachetosellidae

Escharopsis sarsi (Smitt, 1868)

Stomachetosella sp.

continued on nexl page
28 NOAA Technical Report NMFS 140

Table 4 (continued)

BRACHIOPODA

Terebratulina sp.

ECHINODERMATA

Hololhuroidea

Apodida

Chirodota sp.

Labidoplax buskii (Mcintosh, 1866)

Myritrochus sp.

Synaptasp.

Trochoderma sp.

Aspidochirotida

Astichopus sp.

Dendrochirotida

Cuwmaria planci Marenzeller, 1893

Cucumaria sp.

Havelockia scabra (Verrill, 1873)

Psolus jabricii Duben and Koren, 1846

Psolus phantapus (Strussenfeldt, 1765)

Psolus valvatus 0stergren in Grieg, 1913

Psolus sp.

Stereoderma unisemita (Stimpson, 1851)

Thyone jusus (Muller, 1788)

Thyonesp.

Thyonidium pellucidum Duben and Koren, 1844

Molpadiida

Caudina arenata (Gould, 1841)

Molpadia oolitica (Pourtales, 1851)

Molpadia sp.

Dactylochirotida

EchinoCtlcumis sp.

Echinoidea

Camarodonta

Strongylocentrotus droebachiensis (Muller, 1776)

Clypeastroidea

Echinarachnius panna (Lamarck, 1816)

Echinocyamus grandiporus Mortensen, 1907

Spatangoida

Acesle bellidifera Wyville Thompson, 1877

Aeropsis rostrata Norman, 1876

B,isasterfragilis (Duben and Koren, 1844)

Brissopsis atlantica Mortensen, 1907

Schiwster orbign)'anus A. Agassiz, 1883

Schiwstersp.

Ophiuroidea

Amphilepididae

Amphilepis ingolfiana Mortensen, 1933

Amphiuridae

Amphilimna olivacea (Lyman, 1869)

Total Macrobenthos - All Taxonomic

Groups Combined .

Geographic Distribution

Macrobenthic invertebrates showed clear geographic

trends in abundance (Fig. 12). Both density and biomass of organisms exhibited similar patterns. Density was

generally highest (1,000 to 15,000/m2) in the coastal

regions of the Gulf of Maine, on Georges Bank, and on

Amphilimna sp.

Amphioplus abditus (Verrill, 1872)

Amphioplus macilentus (Verrill, 1882)

Amphioplus sp.

Amphiura oUeri Ljungman, 1871

Amphiurafragilis (Verrill, 1885)

Amphiura grandisquama Lyman, 1869

Amphiura sp.

Axiognathus squamatus (delle Chiaje, 1828)

Axiognathus sp.

Ophiacanthidae

Ophiacantha abyssicola (E. Forbes, 1843)

Ophiacantha bidentata (Retzius, 1805)

Ophiacantha sp.

Ophiochiton tenuispinus (Verrill, 1884)

Ophiocten scutalUm Koehler, 1896

Ophiocten sericeum (Forbes, 1852)

Ophiomusium lymani Thompson, 1873

Ophiura ljungmani (Lyman, 1878)

Ophiura robusta (Ayres, 1852)

Ophiura saTSi Lutken, 1858

0phiurasp.

Asteroidea

Asteriidae

Ast/i1iasjorbesi (Desor, 1848)

Astmas vulgaris (Verrill, 1866)

Astmassp.

Leptasterias sp.

ASlropectinidae

Astropecten am/i1·icana (Verrill, 1880)

Astropecten sp.

Goniopectinidae

Clenodisclts crispatus (Retzius, 1805)

ClenodisC1ls sp.

Echinasteridae

Henricia sanguinolenta (Sars, 1844)

Hemiciasp.

Solasleridae

Solastersp.

HEMICHORDATA

Enleropneusta

Balanoglossus sp.

CHORDATA

Ascidiacea

Amaroucium sp.

Bostrichobranchus sp.

Molgulasp.

most of the continental shelf region off Southern New

England. Density was generally low (less than 100 individuals/ m2) over most of the continental rise, and moderately low (100 to 1,000/m2 ) in the central Gulf of

Maine, on the southeastern Scotian Shelf area, and

along the continental slope. The distribution of biomass was similar to that for density. High biomass (100 to

3,400 g/m2 ) was distributed around the periphery of

the Gulf of Maine, on Georges Bank, on most of the

continental shelf off Southern ew England, and in
Composition and Distribution of Macrobenthic Invertebrate Fauna 29

Table 5

Mean number and weight per square meter, total number and weight, number of samples, and percent frequency of

occurrence of each taxonomic group, based on 1,076 samples.

Mean Total Number Frequency of

of occurrence

Taxon No./m2 Wt./m2 No. Wt. samples %

PORlFERA 1.5 2.24 1,566 2,415.82 71 6.6

COELENTERATA 32.1 7.33 34,513 7,884.63 449 41.7

Hydrozoa 6.4 0.52 6,878 560.32 126 11.7

Anthozoa 25.7 6.81 27,635 7,324.31 323 30.0

Alcyonaria 0.8 0.20 902 219.89 63 5.9

Zoantharia 22.6 6.39 24,322 6,871.85 265 24.6

Unidentified 2.2 0.22 2,411 232.57 100 9.3

PLATYHELMINTHES 0.4 0.01 381 6.44 16 1.5

Turbellaria 0.4 0.01 381 6.44 16 1.5

NEMERTEA 8.2 0.71 8,806 765.70 405 37.6

ASCHELMINTHES 2.8 0.01 3,047 8.19 98 9.1

Nematoda 2.8 0.01 3,047 8.19 98 9.1

ANNELIDA 425.0 17.41 457,283 18,727.37 1,034 96.1

POGONOPHORA 0.6 <0.01 618 3.22 56 5.2

SIPUNCULIDA 5.9 0.75 6,358 802.87 249 23.1

ECHIURA 0.1 0.30 105 327.78 17 1.6

PRIAPULlDA <0.1 <0.01 10 4.60 4 0.4

MOLLUSCA 188.0 83.64 202,250 89,998.88 946 87.9

Polyplacophora 1.5 0.14 1,608 148.66 84 7.8

Gastropoda 17.8 2.23 19,165 2,396.85 470 43.7

Bivalvia 163.1 80.95 175,535 87,105.74 893 83.0

Scaphopoda 5.1 0.32 5,458 339.69 218 20.3

Cephalopoda 0.4 0.01 376 7.34 5 0.5

Unidentified 0.1 <0.01 108 0.60 2 0.2

ARTHROPODA 726.2 9.41 781,348 10,125.53 936 87.0

Pycnogonida 0.3 0.01 369 8.45 25 2.3

Arachnida <0.1 <0.01 3 0.03 1 0.1

Crustacea 725.8 9.40 780,976 10,117.05 910 84.6

Ostracoda <0.1 <0.01 19 0.19 5 0.5

Cirripedia 21.8 3.39 23,511 3,648.00 41 3.8

Copepoda <0.1 <0.01 26 0.18 4 0.4

Cumacea 25.8 0.11 27,758 120.97 390 36.3

Tanaidacea <0.1 <0.01 50 0.66 15 1.4

Isopoda 12.1 0.29 12,966 1,313,24 390 36.3

Amphipoda 655.8 4.16 705,612 4,478.71 862 80.1

Mysidacea 2.5 0.01 2,642 12.33 41 3.8

Decapoda 7.5 1.43 8,039 1,540.72 246 22.9

Unidentified 0.3 <0.01 353 2.07 18 1.7

BRYOZOA 15.7 1.29 16,915 1,391.00 119 11.1

BRACHIOPODA 4.5 0.89 4,793 955.31 54 5.0

ECHINODERMATA 79.3 55.00 85,331 59,182.14 772 71.8

Crinoidea <0.1 <0.01 13 0.18 2 0.2

Holothuroidea 4.3 12.87 4,633 13,849.69 202 18.8

Echinoidea 29.3 36.75 31,512 39,540.94 293 27.2

Ophiuroidea 44.2 3.26 47,565 3,504.18 487 45.3

Asteroidea 1.5 2.13 1,608 2,287.15 144 13.4

HEMICHORDATA 0.1 0.02 101 18.67 4 0.4

CHORDATA 16.3 4.10 17,520 4,415.19 181 16.8

Ascidiacea 16.3 4.10 17,520 4,415.19 181 16.8

U IDE TIFIED 5.8 0.27 6,199 294.42 261 24.3

Total 1,512.2 183.39 1,627,144 197,327.78
30 NOAA Technical Report NMFS 140

NEW

JERSEY

OJ 1- 99

m100-999

.'000- 499

.5000-15,337

INDIVIDUALS

PER SQUARE METER

EXPLANATION

EJ 1.0-49.9

r.:m 500. -999.

11100.0-499.

• 500.0-3444.2

GRAMS

PER SQUARE METER

72·

ALL TAXA COMBINED

70· 69·

Figure 12

Geographic distribution of the density (A) and biomass (B) of all taxonomic groups combined. Density is expressed as number of individuals per square meter of bottom area;

biomass is expressed as wet (damp) weight per square meter of bottom area.

coastal areas south of New York. Low biomass (less than

50 g/m2) was prevalent in the central Gulf of Maine, on

the southeastern Scotian Shelf, and along the continental slope and continental rise.

A few areas were characterized by a very high density

(5,000/m2 or greater) and an unusually high biomass

(500 g/m2 or more). One of these exceedingly rich

areas was located at the mouth of the Bay of Fundy;
___________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 31

'2

71

70

NEW

JERSEY

EXPLANATION

MEAN NUMIER OF ANIMALS

PER SQUARE METER OF BOTTOM,

All TAXA COMIINED.

". " "

M A I N E

. ". L......o....... 'M

A

71

NEW

EXPlAN ATI ON D CRUSTACEA • ECHINODERMATA

ANNELIDA OTHER

rliii1 MOllUSCA

PEICENTAGE COMPOSITION .. B

M A I N E

Figure 13

Quantitative composition of the total macrobenthic invertebrate fauna in

relation to the six standard geographic subareas. A.-Mean number of

individuals per square meter of bottom area; B.-Percentage composition,

by density, of the major taxonomic groups.

another was located in the vicinity of the southern end

of Great South Channel and southwestern Georges

Bank. Several smaller rich areas were encountered in

the coastal region ofRhode Island and New York. Generally, they occurred around the periphery of the Gulf

of Maine and offsouthern New England.

Substantial differences in both biomass and density

existed among the six geographic areas (Tables 6, 7; Fig.

13A). Average density was highest (2,382 and 1,961/m2)

on the Southern New England Shelf and on Georges

Bank, intermediate in Nova Scotia and the Gulf of

Maine, and lowest (about 300/m2) on Georges Slope.
32 NOAA Technical Report NMFS 140

Table 6

Mean number of specimens of each taxon per square meter in relation to geographic area.

Geographic areas

Southern Southern

New England New England

Taxon Nova Scotia Gulf of Maine Georges Bank Shelf Georges Slope Slope All areas

PORIFERA 4.8 2.7 0.6 0.5 0.3 0.2 1.5

COELENTERATA 22.2 9.2 99.7 22.6 6.7 8.5 32.1

Hydrozoa 11.5 3.3 6.8 9.9 0.1 0.8 6.4

Anthozoa 10.7 5.9 92.9 12.7 6.6 7.7 25.7

Alcyonaria 0.7 0.9 1.2 1.0 1.5 0.8

Zoantharia 8.2 3.6 92.5 7.3 3.0 4.3 22.6

Unidentified 1.8 1.4 0.4 4.2 2.6 1.9 2.2

PLATYHELMINTHES 0.2 <0.1 0.2 0.9 0.4 0.4

Turbellaria 0.2 <0.1 0.2 0.9 0.4 0.4

NEMERTEA 3.0 4.1 22.7 6.8 1.2 1.6 8.2

ASCHELMINTHES 0.9 3.1 1.7 4.0 2.3 2.3 2.8

Nematoda 0.9 3.1 1.7 4.0 2.3 2.3 2.8

A NELIDA 648.4 291.3 545.6 530.8 79.9 148.6 425.0

POGONOPHORA 0.3 <0.1 3.1 5.3 0.6

SIPUNCULIDA 9.3 4.6 4.4 7.2 1.2 8.7 5.9

ECHIURA 0.1 0.1 0.4 0.2 0.1

PRIAPULIDA 0.1 0.1 <0.1

MOLLUSCA 77.2 306.2 46.8 244.2 83.1 57.9 188.0

Polyplacophora 1.9 3.6 0.1 0.9 0.6 0.2 1.5

Gastropoda 15.0 15.2 11.2 28.8 8.4 6.9 17.8

Bivalvia 50.6 276.0 34.4 212.3 69.5 45.8 163.1

Scaphopoda 9.6 11.4 1.2 1.0 4.5 4.8 5.1

Cephalopoda <0.1 <0.1 1.0 0.1 0.1 0.4

Unidentified 0.3 <0.1 0.1

ARTHROPODA 329.8 150.4 1,052.4 1,386.0 137.6 21.5 726.2

Pycnogonida 0.8 0.1 0.3 0.6 0.1 0.3

Arachnida <0.1 <0.1

Crustacea 329.0 150.3 1,052.1 1,385.4 137.5 21.5 725.9

Ostracoda 0.1 <0.1 <0.1 <0.1 <0.1

Cirripedia 35.7 6.4 2.7 52.2 21.8

Copepoda <0.1 0.2 <0.1

Cumacea 7.3 15.0 45.0 37.0 2.5 3.0 25.8

Tanaidacea 0.4 0.4 <0.1

Isopoda 3.9 9.5 18.0 17.0 1.3 1.0 12.1

Amphipoda 280.0 118.2 9S2.9 1,269.3 133.7 17.1 655.8

Mysidacea <0.1 0.2 10.6 1.0 <0.1 2.5

Decapoda 2.1 0.5 22.3 8.7 0.1 7.5

Unidentified 0.5 0.5 0.2 <0.1 0.3

BRYOZOA 16.3 6.9 27.9 21.9 0.4 15.7

BRACHIOPODA 22.4 9.5 <0.1 4.5

ECHINODERMATA 23.6 43.3 121.0 122.7 18.8 18.7 79.3

Crinoidea <0.1 <0.1 <0.1

Holothuroidea 2.5 7.8 0.2 4.7 2.2 3.4 4.3

Echinoidea 3.6 4.6 105.6 21.8 0.2 0.3 29.3

Ophiuroidea 17.0 29.5 14.0 93.7 15.8 14.8 44.2

Asteroidea 0.4 1.5 1.1 2.5 0.6 0.2 1.5

HEMICHORDATA 0.3 0.1 0.1

CHORDATA 2.8 2.3 33.8 26.8 2.6 1.3 16.3

Ascidiacea 2.8 2.3 33.8 26.8 2.6 1.3 16.3

U IDE TIFIED 1.7 4.9 4.3 7.6 9.4 7.2 5.8

Total 1,162.6 838.7 1,961.0 2,382.4 347.4 281.9 1,512.2
Composition and Distribution of Macrobenthic Invertebrate Fauna 33

Table 7

The number ofspecimens of each taxon, expressed as a percentage of the total benthic invertebrate fauna, in relation to

geographic area.

Geographic areas

Southern Southern

New England New England

Taxon Nova Scotia Gulf of Maine Georges Bank Shelf Georges Slope Slope All areas

PORIFERA 0.4 0.3 <0.1 0.1 0.1 0.1 0.1

COELENTERATA 1.9 1.1 5.1 1.0 1.9 3.0 2.1

Hydrozoa 1.0 0.4 0.4 0.4. <0.1 0.3 0.4

Anthozoa 0.9 0.7 4.7 0.6 1.9 2.7 1.7

Alcyonaria 0.1 0.1 <0.1 0.3 0.5 0.1

Zoantharia 0.7 0.4 4.7 0.3 0.9 1.5 1.5

Unidentified 0.2 0.2 <0.1 0.2 0.8 0.7 0.2

PLATYHELMINTHES <0.1 <0.1 <0.1 <0.1 0.1 <0.1

Turbellaria <0.1 <0.1 <0.1 <0.1 0.1 <0.1

NEMERTEA 0.3 0.5 1.2 0.3 0.3 0.6 0.5

ASCHELMINTHES 0.1 0.4 0.1 0.2 0.7 0.8 0.2

Nematoda 0.1 0.4 0.1 0.2 0.7 0.8 0.2

ANNELIDA 55.8 34.7 27.8 22.3 23.0 52.7 28.1

POGONOPHORA <0.1 <0.1 0.9 1.9 <0.1

SIPUNCULIDA 0.8 0.6 0.2 0.3 0.3 3.1 0.4

ECHIURA <0.1 <0.1 0.1 0.1 <0.1

PRIAPULIDA <0.1 <0.1 <0.1

MOLLUSCA 6.6 36.5 2.4 10.2 23.9 20.5 12.4

Polyplacophora 0.2 0.4 <0.1 <0.1 0.2 0.1 0.1

Gastropoda 1.3 1.8 0.6 1.2 2.4 2.5 1.2

Bivalvia 4.4 32.9 1.8 8.9 20.0 16.3 10.8

Scaphopoda 0.8 1.4 0.1 <0.1 1.3 1.7 0.3

Cephalopoda <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Unidentified <0.1 <0.1 <0.1

ARTHROPODA 28.4 17.9 53.7 58.2 39.6 7.6 48.0

Pycnogonida 0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Arachnida <0.1 <0.1

Crustacea 28.3 17.9 53.7 58.2 39.6 7.6 48.0

Ostracoda <0.1 <0.1 <0.1 <0.1 <0.1

Cirripedia 3.1 0.8 0.1 2.2 1.4

Copepoda <0.1 0.1 <0.1

Cumacea 0.6 1.8 2.3 1.6 0.7 1.1 1.7

Tanaidacea 0.1 0.1 <0.1

Isopoda 0.3 1.1 0.9 0.7 0.4 0.4 0.8

Amphipoda 24.1 14.1 48.6 53.3 38.5 6.0 43.4

Mysidacea <0.1 <0.1 0.5 <0.1 <0.1 0.2

Decapoda 0.2 0.1 1.1 0.4 <0.1 0.5

Unidentified 0.1 <0.1 <0.1 <0.1 <0.1

BRYOZOA 1.4 0.8 1.4 0.9 0.1 1.0

BRACHIOPODA 1.9 1.1 <0.1 0.3

ECHINODERMATA 2.0 5.2 6.2 5.2 5.4 6.6 5.2

Crinoidea <0.1 <0.1 <0.1

Holothuroidea 0.2 0.9 <0.1 0.2 0.6 1.2 0.3

Echinoidea 0.3 0.5 5.4 0.9 <0.1 0.1 1.9

Ophiuroidea 1.5 3.5 0.7 3.9 4.6 5.2 2.9

Asteroidea <0.1 0.2 <0.1 0.1 0.2 0.1 0.1

HEMICHORDATA <0.1 <0.1 <0.1

CHORDATA 0.2 0.3 1.7 1.1 0.7 0.5 1.1

Ascidiacea 0.2 0.3 1.7 1.1 0.7 0.5 1.1

UNIDENTIFIED 0.1 0.6 0.2 0.3 2.7 2.5 0.4

Total 100.0 100.0 100.0 100.0 100.0 100.0 100.0
34 NOAA Technical Report NMFS 140

Table 8

Mean wet weight ofspecimens of each taxon (grams per square meter) in relation to geographic area.

Geographic areas

Southern Southern

New England New England

Taxon Nova Scotia Gulf of Maine Georges Bank Shelf Georges Slope Slope All areas

PORIFERA 15.49 3.15 0.47 0.09 0.24 0.03 2.24

COELENTERATA 20.23 11.87 3.68 4.62 2.42 0.99 7.33

Hydrozoa 0.49 0.12 1.61 0.41 <0.01 0.03 0.52

Anthozoa 19.74 11.75 2.07 4.21 2.42 0.96 6.81

Alcyonaria 0.03 0.43 0.21 0.14 0.07 0.20

Zoantharia 19.54 10.90 2.04 3.84 1.90 0.70 6.39

Unidentified 0.16 0.41 0.03 0.15 0.38 0.19 0.22

PLATYHELMINTHES <0.01 0.01 <0.01 0.01 0.01 0.01

Turbellaria <0.01 0.01 <0.01 0.01 0.01 0.01

NEMERTEA 0.56 0.34 0.83 1.04 0.11 0.20 0.71

ASCHELMINTHES <0.01 0.01 0.01 0.01 0.01 0.01 0.01

Nematoda <0.01 0.01 0.01 0.01 0.01 0.01 0.01

ANNELlDA 18.50 15.51 7.93 29.60 4.86 4.32 17.41

POGONOPHORA <0.01 <0.01 0.01 0.03 <0.01

SIPUNCULlDA 1.65 0.37 0.46 0.74 1.01 1.83 0.75

ECHIURA 0.01 0.07 5.18 0.40 0.03

PRIAPULlDA 0.01 0.05 <0.01

MOLLUSCA 54.40 31.59 79.54 170.90 2.65 1.18 83.64

Polyplacophora 0.10 0.19 <0.01 U.24 0.01 0.01 0.14

Gastropoda 2.47 0.90 1.98 4.29 0.32 0.05 2.23

Bivalvia 50.81 29.84 77.40 166.34 2.11 1.04 80.95

Scaphopoda 1.03 0.66 0.15 0.02 0.20 0.08 0.32

Cephalopoda <0.01 0.01 0.01 <0.01 <0.01 0.01

Unidentified <0.01 <0.01 <0.01

ARTHROPODA 16.49 2.43 9.75 17.11 0.64 0.13 9.41

Pycnogonida 0.02 0.02 <0.01 <0.01 <0.01 0.01

Arachnida <0.01 <0.01

Crustacea 16.49 2.41 9.75 17.11 0.64 0.13 9.40

Ostracoda <0.01 <0.01 <0.01 <0.01 <0.01

Cirripedia 12.71 0.47 0.35 6.84 3.39

Copepoda <0.01 <0.01 <0.01

Cumacea 0.03 0.05 0.20 0.17 0.04 0.02 0.11

Tanaidacea 0.01 <0.01 <0.01

Isopoda 0.09 0.34 0.30 0.40 0.02 0.02 0.29

Amphipoda 1.36 0.94 5.55 8.34 0.57 0.08 4.16

Mysidacea <0.01 <0.01 0.05 0.01 <0.01 0.01

Decapoda 2.27 0.61 3.29 1.36 0.02 1.43

Unidentified <0.01 <0.01 <0.01 <0.01 <0.01

BRYOZOA 6.32 0.17 2.64 0.71 0.02 1.29

BRACHIOPODA 3.68 2.12 <0.01 0.89

ECHINODERMATA 39.44 56.42 119.99 36.06 3.89 10.01 55.00

Crinoidea <0.01 <0.01 <0.01

Holothuroidea 0.77 27.55 0.50 14.65 1.28 2.75 12.87

Echinoidea 32.75 23.54 117.19 12.89 1.77 4.62 36.75

Ophiuroidea 1.46 3.30 1.26 5.41 0.79 2.62 3.26

Asteroidea 4.45 2.03 1.04 3.11 0.05 0.02 2.13

HEMICHORDATA 0.05 <0.01 0.02

CHORDATA 0.79 2.62 8.41 5.13 0.17 0.11 4.10

Ascidiacea 0.79 2.62 8.41 5.13 0.17 0.11 4.10

UNIDENTIFIED 0.12 0.23 0.08 0.52 0.12 0.17 0.27

Total 177.64 127.03 233.79 266.64 21.34 19.46 183.39
Composition and Distribution of Macrobenthic Invertebrate Fauna 35

Table 9

The wet weight ofspecimens of each taxon, expressed as a percentage of the total benthic invertebrate fauna, in relation

to geographic area.

Geographic areas

Southern Southern

New England New England

Taxon Nova Scotia Gulf of Maine Georges Bank Shelf Geprges Slope Slope All areas

PORIFERA 8.7 2.5 0.2 0.1 Ll 0.2 1.2

COELENTERATA 11.4 9.4 1.6 1.7 11.3 5.1 4.0

Hydrozoa 0.3 0.1 0.7 0.2 <0.1 0.2 0.3

Anthozoa ILl 9.3 0.9 1.5 11.3 4.9 3.7

Alcyonaria <0.1 0.3 0.1 0.7 0.4 0.1

Zoantharia 11.0 8.6 0.9 1.4 8.9 3.6 3.5

Unidentified 0.1 0.3 <0.1 0.1 1.8 1.0 0.1

PLATYHELMINTHES <0.1 <0.1 <0.1 <0.1 0.1 <0.1

Turbellaria <0.1 <0.1 <0.1 <0.1 0.1 <0.1

NEMERTEA 0.3 0.4 0.4 0.4 0.5 1.0 0.4

ASCHELMINTHES <0.1 <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Nematoda <0.1 <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

ANNELIDA 10.4 12.2 3.4 ILl 22.8 22.2 9.5

POGONOPHORA <0.1 <0.1 <0.1 0.2 <0.1

SIPUNCULIDA 0.9 0.3 0.2 0.3 4.7 9.4 0.4

ECHIURA <0.1 <0.1 24.3 2.0 0.2

PRIAPULIDA 0.1 0.2 <0.1

MOLLUSCA 30.6 24.9 34.0 64.1 12.4 6.0 45.6

Polyplacophora 0.1 0.2 <0.1 0.1 0.1 <0. I 0.1

Gastropoda 1.4 0.7 0.8 1.6 1.5 0.3 1.2

Bivalvia 28.6 23.5 33.1 62.4 9.9 5.3 44.1

Scaphopoda 0.6 0.5 0.1 <0.1 0.9 0.4 0.2

Cephalopoda <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Unidentified <0.1 <0.1 <0.1

ARTHROPODA 9.3 1.9 4.2 6.4 3.0 0.6 5.1

Pycnogonida <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Arachnida <0.1 <0.1

Crustacea 9.3 1.9 4.2 6.4 3.0 0.6 5.1

Ostracoda <0.1 <0.1 <0.1 <0.1 <0.1

Cirripedia 7.2 0.4 0.2 2.6 1.8

Copepoda <0.1 <0.1 <0.1

Cumacea <0.1 <0.1 0.1 0.1 0.2 0.1 0.1

Tanaidacea <0.1 <0.1 <0.1

Isopoda <0.1 0.3 0.1 0.2 0.1 0.1 0.2

Amphipoda 0.8 0.7 2.4 3.1 2.7 0.4 2.3

Mysidacea <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Decapoda 1.3 0.5 1.4 0.5 0.1 0.8

Unidentified <0.1 <0.1 <0.1 <0.1 <0.1

BRYOZOA 3.6 0.1 Ll 0.3 0.1 0.7

BRACHIOPODA 2.1 1.7 <0.1 0.5

ECHINODERMATA 22.2 44.4 51.3 13.5 18.2 51.5 30.0

Crinoidea <0.1 <0.1 <0.1

Holothuroidea 0.4 21.7 0.2 5.5 6.0 14.2 7.0

Echinoidea 18.4 18.5 50.1 4.8 8.3 23.7 20.0

Ophiuroidea 0.8 2.6 0.5 2.0 3.7 13.5 1.8

Asteroidea 2.5 1.6 0.4 1.2 0.2 0.1 1.2

HEMICHORDATA <0.1 <0.1 <0.1

CHORDATA 0.4 2.1 3.6 1.9 0.8 0.6 2.2

Ascidiacea 0.4 2.1 3.6 1.9 0.8 0.6 2.2

UNIDENTIFIED 0.1 0.2 <0.1 0.2 0.6 0.9 0.2

Total 100.0 100.0 100.0 100.0 100.0 100.0 100.0
36 NOAA Technical Report NMFS 140

Table 10

Frequency of occurrence (%) of each taxonomic group in the samples in each geographic area.

Geographic areas

Southern Southern

New England New England

Taxon Nova Scotia Gulf of Maine Georges Bank Shelf Georges Slope Slope

PORIFERA 19 9 4 3 10 5

COELENTERATA 61 33 39 43 64 46

Hydrozoa 29 8 15 13 4 I

Anthozoa 32 25 24 30 60 45

Alcyonaria 4 3 7 27 17

Zoantharia 31 22 28 25 17 21

PLAlYHELMINTHES 2 <l 1 3 2

Turbellaria 2 <l 1 3 2

NEMERTEA 28 32 32 52 35 25

ASCHELMINTHES 8 9 5 6 39 15

Nematoda 8 9 5 6 39 15

ANNELIDA 97 98 89 99 96 99

POGONOPHORA 2 1 35 42

SIPUNCULIDA 42 20 13 24 25 36

ECHIURA <l 15 7

PRlAPULIDA 4 3

MOLLUSCA 91 90 73 92 92 95

Polyplacophora 24 11 1 5 15 6

Gastropoda 54 43 35 43 58 53

Bivalvia 80 87 64 89 89 94

Scaphopoda 46 35 7 4 35 33

Cephalopoda 1 <l <l 2 1

ARTHROPODA 95 71 100 95 85 74

Pycnogonida 8 1 3 2 6

Arachnida 1

Crustacea 95 71 ]00 95 85 74

Ostracoda 2 <1 2 ]

Cirripedia 15 4 2 4

Copepoda I 3

Cumacea 31 24 44 49 31 19

Tanaidacea 15 9

Isopoda 35 21 48 48 29 20

Amphipoda 94 58 94 92 73 63

Mysidacea I I 10 4 I

Decapoda 18 6 46 33 3

BRYOZOA 19 12 13 11 6

BRACHIOPODA 21 12 2

ECHINODERMATA 69 79 67 67 81 75

Crinoidea <l 2

Holothuroidea 22 25 2 17 40 27

Echinoidea 26 19 51 28 8 9

Ophiuroidea 55 64 22 35 62 59

Asteroidea 9 13 9 19 14 6

HEMICHORDATA I I

CHORDATA 20 12 14 23 ]4 16

Ascidiacea 20 12 14 23 14 16
___________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 37

"

NEW

JEISEY

EXPLANATION

MEAN WEIGHT OF ANIMALS

PER SQUARE METER Of BOTTOM.

All TAXA COMBINED.

n'

A

,,' '0'

MAl N E

..'

.9' '0'

MAt N E

",

B

,,' 38 39

NEW

JE~~.EY

EXPLANATION D CRUSTACEA • ECHINODERMATA

ANNELIDA D OTHER

'" :~~: , . MOllUSCA

PERCENTAGE COMPOSITION

"

Figure 14

Quantitative composition of the total macrobenthic- invertebrate fauna in

relation to the six standard geographic subareas. A.-Mean wet weight of

animals per square meter of bottom area; B.-Percentage composition, by

biomass, of the major taxonomic groups.

A gradient was evident in the density distribution. In

the three shallow water areas-Nova Scotia, Georges

Bank, and Southern New England Shelf-average density increased in the order listed from the northeast to

the southwest. The Southern New England Shelf area

(2,382/m2) had about twice the density of the Nova

Scotian area (l,163/m2). The Gulf of Maine averaged

839/m2, and the two slope areas had average densities

of about 300/m2.

Biomass distribution followed precisely the same rank

order as density, and the magnitudes of differences in

biomass from one area to another were roughly the
38 NOAA Technical Report NMFS 140

Table 11

Mean numher ofspecimens of each taxon per square meter in relation to water depth.

Water depth (m)

Taxon 0-24 25-49 50-99 100-199 200-499 500-599 1,000-1,999 2,000-3,999 All depths

PORIFERA 1.3 2.8 0.8 1.3 2.5 0.3 0.3 0.1 1.5

COELENTERATA 56.7 96.8 28.6 11.6 9.0 6.0 4.3 1.6 32.1

Hydrozoa 36.8 5.1 5.9 2.5 1.0 0.1 0.1 6.4

Anthozoa 19.9 91.7 22.7 9.1 8.0 6.0 4.2 1.5 25.7

Alcyonaria 0.9 0.9 1.9 Ll 1.2 0.4 0.8

Zoantharia 5.6 91.3 2Ll 6.8 4.4 2.0 0.4 0.2 22.6

Uniden tified 14.3 0.4 0.7 1.4 1.7 2.9 2.7 1.0 2.2

PLATIHELMINTHES 2.6 <0.1 0.3 0.2 0.4

Turbellaria 2.6 <0.1 0.3 0.2 0.4

NEMERTEA 4.2 27.2 7.8 3.8 3.4 0.5 1.5 0.7 8.2

ASCHELMINTHES 6.8 0.8 3.2 1.4 4.4 4.2 1.4 1.7 2.8

Nematoda 6.8 0.8 3.2 1.4 4.4 4.2 1.4 1.7 2.8

ANNELIDA 719.3 436.8 51.9 455.9 240.6 106.5 30.2 9.1 425.0

POGONOPHORA 0.1 1.3 6.8 3.2 3.5 0.6

SIPUNCULIDA 1.8 5.8 6.2 7.5 8.1 1.5 0.7 1.2 5.9

ECHIURA 0.3 0.2 0.5 0.6 0.1

PRIAPULIDA 0.2 0.1 <0.1

MOLLUSCA 570.2 205.0 197.3 135.7 91.0 121.5 55.7 27.2 188.0

Polyplacophora 0.8 4.0 Ll 0.4 2.5 0.2 0.5 0.2 1.5

Gastropoda 64.1 18.4 15.2 11.3 12.1 23.7 3.9 Ll 17.8

Bivalvia 505.1 182.2 179.2 113.1 62.6 86.7 50.1 25.8 163.1

Scaphopoda 0.2 0.4 1.5 9.5 13.8 10.8 1.2 0.1 5.1

Cephalopoda 1.5 0.1 0.4

Uniden tified 0.3 0.1 0.1

ARTHROPODA 1,039.4 1,255.7 1,351.6 168.8 67.0 14.6 13.2 6.6 726.2

Pycnogonida 0.6 0.6 0.3 0.5 0.2 0.1 0.3

Arachnida <0.1 <0.1

Crustacea 1,038.8 1,255.1 1,351.3 168.3 67.0 14.4 13.1 6.6 725.9

Ostracoda <0.1 <0.1 0.1 0.1 0.1 <0.1

Cirripedia 214.2 13.7 2.9 2.3 0.7 21.8

Copepoda <0.1 <0.1 0.5 <0.1

Cumacea 33.1 39.8 50.3 6.8 2.9 1.6 0.7 1.7 25.8

Tanaidacea 0.1 0.1 1.0 <0.1

Isopoda 22.4 37.9 ILl 2.9 3.1 1.3 0.4 1.9 12.1

Amphipoda 746.4 1,148.6 1,273.8 147.4 59.1 ILl 12.0 2.9 655.8

Mysidacea 4.0 6.5 3.8 0.1 <0.1 2.5

Decapoda 18.0 8.2 9.1 8.6 0.3 0.1 7.5

Unidentified 0.6 0.4 0.2 0.2 0.8 0.3

BRYOZOA 38.6 29.4 18.9 7.8 4.3 0.5 15.7

BRACHIOPODA 1.2 6.5 17.0 0.1 4.5

ECHINODERMATA 47.4 133.2 87.4 94.6 48.1 6.5 4.4 7.3 79.3

Crinoidea 0.1 0.1 <0.1

Holothuroidea 1.5 1.3 4.9 4.0 9.8 1.2 1.9 0.7 4.3

Echinoidea 44.4 127.4 19.1 2.0 2.6 0.1 0.3 29.3

Ophiuroidea 0.8 2.0 61.5 86.7 35.0 5.2 2.0 6.2 44.2

Asteroidea 0.7 2.5 1.9 1.8 0.5 0.1 0.3 0.1 1.5

HEMICHORDATA 0.3 <0.1 0.5 0.1

CHORDATA 9.0 32.0 29.8 6.2 2.7 0.1 1.0 1.7 16.3

Ascidiacea 9.0 32.0 29.8 6.2 2.7 0.1 1.0 1.7 16.3

UNIDENTIFIED 6.0 6.6 5.1 5.3 5.6 3.5 5.8 13.1 5.8

Total 2,503.3 2,232.0 2,256.9 906.6 505.1 272.4 122.2 74.9 1,512.2
Composition and Distribution of Macrobenthic Invertebrate Fauna 39

Table 12

The number of specimens of each taxon, expressed as a percentage of the total benthic invertebrate fauna, in relation to

water depth.

Depth zones (m)

Taxon 0-24 25-49 50-99 100-199 200-499 500-599 1,000-1,999 2,000-3,999 All depths

PORIFERA 0.1 0.1 <0.1 0.2 0.5 0.1 0.2 0.1 0.1

COELENTERATA 2.3 4.3 1.3 1.3 1.8 2.2 3.5 2.1 2.1

Hydrozoa 1.5 0.2 0.3 0.3 0.2 <0.1 0.1 0.4

Anthozoa 0.8 4.1 1.0 1.0 1.6 2.2 3.5 2.0 1.7

Alcyonaria <0.1 0.1 0.4 0.4 1.0 0.6 0.1

Zoantharia 0.2 4.1 0.9 0.8 0.9 0.7 0.3 0.2 1.5

Unidentified 0.6 <0.1 <0.1 0.2 0.3 1.0 2.2 1.3 0.2

PLATYHELMINTHES 0.1 <0.1 <0.1 <0.1 <0.1

Turbellaria 0.1 <0.1 <0.1 <0.1 <0.1

NEMERTEA 0.2 1.2 0.4 0.4 0.7 0.2 1.2 0.9 0.5

ASCHELMINTHES 0.3 <0.1 0.1 0.2 0.9 1.5 1.1 2.3 0.2

Nematoda 0.3 <0.1 0.1 0.2 0.9 1.5 1.1 2.3 0.2

ANNELIDA 28.7 19.6 23.0 50.3 47.6 39.1 24.7 12.2 28.1

POGONOPHORA <0.1 0.2 2.5 2.6 4.6 <0.1

SIPUNCULIDA 0.1 0.3 0.3 0.8 1.6 0.5 0.6 1.6 0.4

ECHIURA <0.1 <0.1 0.4 0.8 <0.1

PRIAPULIDA 0.2 0.1 <0.1

MOLLUSCA 22.8 9.2 8.7 15.0 18.0 44.6 45.6 36.3 12.4

Polyplacophora <0.1 0.2 <0.1 <0.1 0.5 0.1 0.4 0.3 0.1

Gastropoda 2.6 0.8 0.7 1.2 2.4 8.7 3.2 1.5 1.2

Bivalvia 20.2 8.2 7.9 12.5 12.4 31.8 41.0 34.4 10.8

Scaphopoda <0.1 <0.1 0.1 1.0 2.7 4.0 1.0 0.1 0.3

Cephalopoda 0.2 <0.1 <0.1

Unidentified <0.1 <0.1 <0.1

ARTHROPODA 41.5 56.3 59.9 18.6 13.3 5.4 10.8 8.8 48.0

Pycnogonida <0.1 <0.1 <0.1 <0.1 0.1 <0.1 <0.1

Arachnida <0.1 <0.1

Crustacea 41.5 56.2 59.9 18.6 13.3 5.3 10.8 8.8 48.0

Ostracoda <0.1 <0.1 <0.1 <0.1 0.1 <0.1

Cirripedia 8.6 0.6 0.1 0.2 0.1 1.4

Copepoda <0.1 <0.1 0.2 <0.1

Cumacea 1.3 1.8 2.2 0.8 0.6 0.6 0.6 2.3 1.7

Tanaidacea <0.1 <0.1 1.4 <0.1

Isopoda 0.9 1.7 0.5 0.3 0.6 0.5 0.3 2.5 0.8

Amphipoda 29.8 51.5 56.4 16.3 11.7 4.1 9.8 3.9 43.4

Mysidacea 0.2 0.3 0.2 <0.1 <0.1 0.2

Decapoda 0.7 0.4 0.4 1.0 0.1 <0.1 0.5

Unidentified <0.1 <0.1 <0.1 <0.1 0.2 0.1 <0.1

BRYOZOA 1.5 1.3 0.8 0.9 0.8 0.7 1.0

BRACHIOPODA <0.1 0.7 3.4 <0.1 0.3

ECHINODERMATA 1.9 6.0 3.9 10.4 9.5 2.4 3.6 9.8 5.2

Crinoidea <0.1 <0.1 <0.1

Holothuroidea 0.1 0.1 0.2 0.4 1.9 0.4 1.5 0.9 0.3

Echinoidea 1.8 5.7 0.8 0.2 0.5 0.1 0.4 1.9

Ophiuroidea <0.1 0.1 2.7 9.6 6.9 1.9 1.6 8.3 2.9

Asteroidea <0.1 0.1 0.1 0.2 0.1 <0.1 0.3 0.1 0.1

HEMICHORDATA <0.1 <0.1 0.2 <0.1

CHORDATA 0.4 1.4 1.3 0.7 0.5 <0.1 0.8 2.2 1.1

Ascidiacea 0.4 1.4 1.3 0.7 0.5 <0.1 0.8 2.2 1.1

UNIDENTIFIED 0.2 0.3 0.2 0.6 1.1 1.3 4.7 17.4 0.4

Total 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0
40 NOAA Technical Report NMFS 140

75

122

272

505

907

2,257

2,232

2,503

EXPlANAT ION

1000+---------1(

D CRUSTACEA

ANNELIDA

.:.:.. - -::... MOllUS,-A ... ~.

II ECHINODERMATA

[;] OTHER

~E~~<:::-t l00:+===:::::~c::;::::::=:::::::::1i~ 200-+-------"'1..

% :zooo+----------\\ t-

..,

Q

CIt

III:

UI

t-

..,

Z

MEAN

NUMBER

PERCENTAGE

COMPOS IT ION

~ooo....l-------------~

Figure 15

Quantitative composition of the total macrobenthic invertebrate fauna in relation

to water depth. Mean number of individuals per square meter of bottom area, and

percentage composition, by density, of the major taxonomic groups are shown.

same as the changes in density (Tables 8, 9; Fig. 14A).

Southern New England Shelf and Georges Bank had

the highest average biomasses (267 and 234 g/m2 );

biomass was intermediate (178 and 127 g/m2) in Nova

Scotia and the Gulf ofMaine, and low (21 and 20 g/m2)

on Georges Slope and Southern New England Slope.

The percentage occurrence of each taxonomic group

in the samples in each geographic area is presented in

Table 10.

Bathymetric Distribution

One ofthe moststriking relationships in the New England

region was the pronounced diminution in quantity of

macrobenthic invertebrates from shallow to deep water

(Tables 11, 12; Fig. 15). In the shallowest waters sampled

(0-24 m) the average density was 2,503/m2, whereas in

the deepest water (2,000-3,999 m) the density averaged

only 751m2. The decrease from one depth range to an-
Composition and Distribution of Macrobenthic Invertebrate Fauna 41

Table 13

Mean wet weight of specimens of each taxon (grams per square meter) in relation to water depth.

Water depth (m)

Taxon 0-24 25-49 50-99 100-199 200-499 500-599 1,000-1,999 2,000-3,999 All depths

PORIFERA 0.06 1.23 2.90 3.08 3.12 0.54 0.02 0.03 2.24

COELE TERATA 3.63 1.49 3.38 18.95 9.13 0.36 0.72 0.69 7.33

Hydrozoa 1.21 0.22 1.17 0.16 0.02 <0.01 <0.01 0.52

Anthozoa 2.42 1.27 2.21 18.79 9.11 0.36 0.72 0.69 6.81

Alcyonaria 0.16 0.33 0.47 0.04 0.25 0.01 0.20

Zoantharia 2.08 1.22 1.99 18.28 8.05 0.19 0.18 0.19 6.39

Unidentified 0.35 0.05 0.06 0.18 0.59 0.13 0.29 0.49 0.22

PLAlYHELMI THES 0.02 <0.01 0.01 <0.01 0.01

Turbellaria 0.02 <0.01 0.01 <0.01 0.01

NEMERTEA 1.00 1.44 0.99 0.23 0.40 0.01 0.06 0.12 0.71

ASCHELMINTHES <0.01 0.01 0.01 0.01 0.01 0.01 0.01 0.01 0.01

Nematoda <0.01 0.01 0.01 0.01 0.01 0.01 0.01 0.01 0.01

A ELIDA 27.22 16.24 25.20 14.76 10.70 4.76 1.41 0.76 17.41

POGONOPHORA <0.01 0.01 0.03 0.02 0.01 <0.01

SIPUNCULIDA 0.22 0.57 0.82 0.77 0.54 3.93 1.38 0.53 0.75

ECHIURA 0.22 0.01 5.04 3.52 0.30

PRIAPULIDA 0.12 0.01 <0.01

MOLL SCA 257.88 106.89 132.14 20.67 10.80 3.26 1.44 0.58 83.64

Polyplacophora 0.84 0.07 0.02 0.02 0.29 0.01 0.01 <0.01 0.14

Gastropoda 4.85 1.82 4.23 1.07 0.18 0.29 0.15 0.17 2.23

Bivalvia 252.18 104.94 127.80 18.56 10.04 2.79 1.07 0.41 80.95

Scaphopoda <0.01 0.07 0.09 0.98 0.29 0.17 0.22 <0.01 0.32

Cephalopoda 0.03 <0.01 0.01

Unidentified <0.01 0.01 <0.01

ARTHROPODA 37.04 15.64 9.31 2.40 3.89 0.09 0.08 0.10 9.41

Pycnogonida 0.01 <0.01 <0.01 0.03 <0.01 <0.01 0.01

Arachnida <0.01 <0.01

Crustacea 37.03 15.64 9.31 2.37 3.89 0.09 0.08 0.10 9.40

Ostracoda <0.01 <0.01 <0.01 <0.01 <0.01 <0.01

Cirripedia 27.08 3.89 0.29 0.10 2.53 3.39

Copepoda <0.01 <0.01 0.10 <0.01

Cumacea 0.08 0.11 0.26 0.04 0.02 0.01 0.01 0.05 0.11

Tanaidacea <0.01 <0.01 0.01 <0.01

Isopoda 0.15 0.66 0.27 0.15 0.42 0.02 0.01 0.02 0.29

Amphipoda 6.39 9.77 6.38 0.97 0.31 0.06 0.03 0.02 4.16

Mysidacea 0.03 0.02 0.02 <0.01 <0.01 0.01

Decapoda 3.32 1.19 2.10 1.12 0.61 0.03 1.43

Unidentified 0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01

BRYOZOA 0.96 0.92 2.88 0.87 0.14 0.02 1.29

BRACHIOPODA 0.11 1.05 3.98 0.01 0.89

ECHINODERMATA 105.93 166.80 33.95 34.23 19.18 1.72 3.16 4.62 55.00

Crinoidea <0.01 <0.01 <0.01

Holoth uroidea 36.59 12.76 19.23 6.47 3.57 0.24 1.24 2.15 12.87

Echinoidea 65.74 153.88 9.24 16.16 11.11 1.13 1.78 36.75

Ophiuroidea 0.29 0.06 2.49 7.50 4.49 1.48 0.75 0.66 3.26

Asteroidea 3.30 0.11 2.99 4.10 0.01 <0.01 0.04 0.04 2.13

HEMICHORDATA 0.05 0.01 0.01 0.02

CHORDATA 3.85 5.20 8.93 1.03 0.72 <0.01 <0.01 0.21 4.10

Ascidiacea 3.85 5.20 8.93 1.03 0.72 <0.01 <0.01 0.21 4.10

UNIDENTIFIED 0.19 0.58 0.28 0.23 0.12 0.19 0.23 0.15 0.27

Total 438.26 317.01 220.95 98.30 62.72 14.91 13.66 11.36 183.39
42 NOAA Technical Report NMFS 140

Table 14

The wet weight ofspecimens of each taxon, expressed as a percentage of the total benthic invertebrate fauna, in relation

to water depth.

Depth zones (m)

Taxon 0-24 25-49 50-99 100-199 200-499 500-599 1,000-1,999 2,000-3,999 All depths

PORIFERA <0.1 0.4 1.3 3.1 5.0 3.6 0.1 0.3 1.2

COELENTERATA 0.8 0.5 1.5 19.3 14.6 2.4 5.3 6.1 4.0

Hydrozoa 0.3 0.1 0.5 0.2 <0.1 <0.1 <0.1 0.3

Anthozoa 0.5 0.4 1.0 19.1 14.6 2.4 5.3 6.1 3.7

Alcyonaria 0.1 0.3 0.8 0.3 1.8 0.1 0.1

Zoantharia 0.5 0.4 0.9 18.6 12.8 1.2 1.3 1.7 3.5

Unidentified 0.1 <0.1 <0.1 0.2 0.9 0.9 2.1 4.3 0.1

PLAlYHELMINTHES <0.1 <0.1 <0.1 <0.1 <0.1

Turbellaria <0.1 <0.1 <0.1 <0.1 <0.1

NEMERTEA 0.2 0.5 0.4 0.2 0.6 0.1 0.4 1.0 0.4

ASCHELM1NTHES <0.1 <0.1 <0.1 <0.1 <0.1 0.1 0.1 0.1 <0.1

Nematoda <0.1 <0.1 <0.1 <0.1 <0.1 0.1 0.1 0.1 <0.1

ANNELIDA 6.2 5.1 11.4 15.0 7.0 32.0 10.3 6.7 9.5

POGONOPHORA <0.1 <0.1 0.2 0.1 0.1 <0.1

SIPUNCULIDA <0.1 0.2 0.4 0.8 0.8 26.3 10.1 4.7 0.4

ECHIURA 0.1 <0.1 36.9 31.0 0.2

PRIAPULIDA 0.9 0.1 <0.1

MOLLUSCA 58.8 33.7 59.8 21.0 17.2 21.9 10.6 5.1 45.6

Polyplacophora 0.2 <0.1 <0.1 <0.1 0.5 <0.1 <0.1 <0.1 0.1

Gastropoda 1.1 0.6 1.9 1.1 0.3 1.9 1.1 1.5 1.2

Bivalvia 57.5 33.1 57.8 18.9 16.0 18.7 7.8 3.6 44.1

Scaphopoda <0.1 <0.1 <0.1 1.0 0.5 1.2 1.6 <0.1 0.2

Cephalopoda <0.1 <0.1 <0.1

Unidentified <0.1 0.1 <0.1

ARTHROPODA 8.4 4.9 4.2 2.4 6.2 0.6 0.6 0.9 5.1

Pycnogonida <0.1 <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Arachnida <0.1 <0.1

Crustacea 8.4 4.9 4.2 2.4 6.2 0.6 0.6 0.9 5.1

Ostracoda <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Cirripedia 6.2 1.2 0.1 0.1 4.0 1.8

Copepoda <0.1 <0.1 <0.1 <0.1

Cumacea <0.1 <0.1 0.1 <0.1 <0.1 0.1 0.1 0.4 0.1

Tanaidacea <0.1 <0.1 0.1 <0.1

Isopoda <0.1 0.2 0.1 0.2 0.7 0.1 <0.1 0.2 0.2

Amphipoda 1.5 3.1 2.9 1.0 0.5 0.4 0.2 0.2 2.3

Mysidacea <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Decapoda 0.8 0.4 1.0 1.1 1.0 0.2 0.8

Unidentified <0.1 <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

BRYOZOA 0.2 0.3 1.3 0.9 0.2 0.2 0.7

BRACHIOPODA <0.1 1.1 6.3 <0.1 0.5

ECHINODERMATA 24.2 52.6 15.4 34.8 30.6 11.5 23.0 40.7 30.0

Crinoidea <0.1 <0.1 <0.1

Holothuroidea 8.4 4.0 8.7 6.6 5.7 1.6 9.0 18.9 7.0

Echinoidea 15.0 48.5 4.2 16.4 17.7 8.2 15.7 20.0

Ophiuroidea 0.1 <0.1 1.1 7.6 7.2 9.9 5.5 5.8 1.8

Asteroidea 0.8 <0.1 1.4 4.2 <0.1 <0.1 0.3 0.3 1.2

HEMICHORDATA <0.1 1.0 <0.1 <0.1

CHORDATA 0.9 1.6 4.0 1.0 1.2 <0.1 <0.1 1.8 2.2

Ascidiacea 0.9 1.6 4.0 1.0 1.2 <0.1 <0.1 1.8 2.2

UNIDE TIFIED <0.1 0.2 0.1 0.2 0.2 1.3 1.6 1.3 0.2

Total 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0
Composition and Distribution of Macrobenthic Invertebrate Fauna 43

98

221

317

438

~~=~~~~ l00+--------=~w-­

200+-----------''l-,

63

1000

UI

ae

1M

tIM

15

!

% 2000 ta.

1M

c 14

ae EXPlANATION iii

tee D CRUSTACEA

ANNELIDA 11

:;:~~.\ MOllUSCA

• ECHINODERMATA

D OTHER

~ooo

MEAN

WET WEIGHT

PERCENTAGE

COMPOSITION

Figure 16

Quantitative composition of the total macrobenthic invertebrate fauna in

relation to water depth. Mean wet weight per square meter of bottom area,

and percentage composition, by biomass, of the major taxonomic groups

are shown.

other was not uniform over the entire depth spectrum.

There was a 60% drop in density from 50-99 m to 100-

199 m and the largest drop in density occurred between

100 and 200 m depth.. Another way of expressing this

change is to say there was a decrease in density per meter

increase in depth. In the vicinity of 100 m the average

density decreased by 18 specimens with each 1-m increase

in water depth. In shallower water the rate ofdecrease was

as high as 10 specimens per 1 m of water depth increase,

but the percentage change was substantially lower. In

deep water (below 200 m) the diminution rate was less

than one specimen per meter change in water depth.

Differences in density in depth ranges between 1,000 and

1,999 m and between 2,000 and 3,999 m decreased an

average of 0.05 per meter increase in depth.

Biomass diminished with depth from an average of

438 g/m2 in shallow water to 11 g/m2 on the continental rise (Tables 13, 14; Fig. 16). The biomass remained

rather high (221 g or more) in shallow water out to 100

meters. In the vicinity of 100 m the biomass was 56%

lower than in the shallowest water, and 76% lower in

the vicinity of 500 m depth.

The relatively high biomass, averaging 11 g/m2, at

water depths between 2,000 and 3,999 m was due in
44 NOAA Technical Report NMFS 140

Table 15

Frequency of occurrence (%) of each taxonomic group in the samples in each depth range class.

Water depth (m)

Taxon 0-24 25-49 50-99 100-199 200-499 500-599 1,000-1,999 2,000-3,999

PORIFERA 5 3 5 8 12 9 12 3

COELENTERATA 36 36 50 40 33 46 65 41

Hydrozoa 21 16 15 10 4 3 3

Anthozoa 15 20 35 30 29 46 62 38

Alcyonaria 5 5 8 23 29 22

Zoantharia 16 21 33 29 19 9 18 8

PLATYHELMINTHES 2 4

Turbellaria 2 4

NEMERTEA 35 51 47 30 28 18 33 24

ASCHELMINTHES 2 6 8 5 15 32 18 35

Nematoda 2 6 8 5 15 32 18 35

ANNELIDA 96 93 96 98 97 100 97 95

POGONOPHORA 5 50 53 43

SIPU CULIDA 3 13 26 32 25 32 21 22

ECHIURA 2 <1 21 19

PRIAPULIDA 9 3

MOLLUSCA 92 82 85 91 89 100 97 87

Polyplacophora 1 4 10 7 11 9 18 8

Gastropoda 52 42 43 37 44 77 74 30

Bivalvia 88 76 81 85 84 100 97 81

Scaphopoda I 1 9 35 43 59 35 3

Cephalopoda 1 1

ARTHROPODA 88 99 100 80 71 82 53 78

Pycnogonida 2 2 2 4 9 3

Arachnida <1

Crustacea 88 99 100 80 71 82 53 78

Ostracoda <1 <I 1 5 3

Cirripedia 13 4 3 3 4

Copepoda I I 5

Cumacea 27 41 56 30 15 36 15 22

Tanaidacea 1 3 35

Isopoda 29 69 46 20 20 18 12 43

Amphipoda 80 95 98 72 58 77 44 54

Mysidacea 13 11 3 <1 1

Decapoda 37 39 35 13 4 3

BRYOZOA 19 13 11 10 12 5

BRACHIOPODA 2 10 14 5

ECHINODERMATA 39 68 71 78 84 64 65 92

Crinoidea 1 3

Holothuroidea 9 8 20 17 33 14 29 27

Echinoidea 21 57 31 16 22 6 16

Ophiuroidea 10 9 40 66 72 59 44 76

Asteroidea 9 9 20 16 6 5 12 5

HEMICHORDATA <1 5

CHORDATA 14 24 19 14 12 5 15 24

Ascidiacea 14 24 19 14 12 5 15 24
Composition and Distribution of Macrobenthic Invertebrate Fauna 45

Table 16

Mean number of specimens of each taxon per square meter in relation to bottom sediments.

Bottom sediments

Taxon Gravel Till Shell Sand Sand-silt Silt-clay All types

PORIFERA 3.9 4.5 0.3 0.4 2.5 0.7 1.5

COELENTERATA 45.0 16.0 7.8 50.0 15.4 6.3 32.1

Hydrozoa 20.3 6.9 5.2 6.0 2.9 1.5 6.4

Anthozoa 24.7 10.9 2.6 44.0 12.5 4.8 25.7

Alcyonaria 1.6 0.2 0.9 1.6 0.8

Zoantharia 13.7 3.4 2.3 43.1 9.3 2.7 22.6

Unidentified 9.3 5.6 0.3 0.7 2.3 0.4 2.2

PLAlYHELMINTHES 1.7 0.2 0.1 0.4

Turbellaria 1.7 0.2 0.1 0.4

EMERTEA 4.9 0.9 27.5 13.3 5.1 3.4 8.2

ASCHELMINTHES 8.7 1.0 1.2 2.4 3.0 2.8

Nematoda 8.7 1.0 1.2 2.4 3.0 2.8

ANNELIDA 504.8 289.5 442.5 558.1 309.7 231.9 425.0

POGONOPHORA <0.1 0.8 1.9 0.6

SIPUNCULIDA 4.0 4.4 6.0 7.1 5.3 5.5 5.9

ECHIURA 0.2 0.3 0.1

PRIAPULIDA <0.1 <0.1 <0.1

MOLLUSCA 83.7 103.0 228.5 98.6 276.0 353.8 188.0

Polyplacophora 2.0 2.5 0.3 3.8 1.3 1.5

Gastropoda 40.3 9.1 82.7 11.2 22.0 11.9 17.8

Bivalvia 39.4 78.3 180.3 85.0 242.1 329.6 163.1

Scaphopoda 2.0 13.1 25.5 1.8 6.4 10.9 5.1

Cephalopoda <0.1 <0.1 1.7 0.4

Uniden tified 0.2 <0.1 0.1

ARTHROPODA 712.0 58.8 124.2 1,336.0 275.5 33.9 726.2

Pycnogonida 2.1 0.1 0.1 0.\ 0.1 0.3

Arachnida <0.1 <0.1

Crustacea 709.9 58.7 124.2 1,335.9 275.4 33.8 725.9

Ostracoda <0.1 0.1 <0.1 <0.1

Cirripedia 28.7 1.0 5.0 16.4 55.5 0.2 21.8

Copepoda 0.1 0.1 0.1 0.1

Cumacea 10.5 1.2 7.5 45.3 18.3 7.0 25.8

Tanaidacea 0.1 0.1 <0.1

Isopoda 5.8 7.1 4.2 22.4 5.2 3.0 12.1

Amphipoda 639.7 49.5 98.8 1,237.6 193.0 23.1 655.8

Mysidacea 0.9 5.0 0.9 0.1 2.5

Decapoda 24.0 8.7 8.7 1.9 0.2 7.5

Unidentified 0.3 0.4 0.6 0.3

BRYOZOA 75.1 5.6 331.0 4.9 5.4 1.5 15.7

BRACHIOPODA 13.6 48.2 37.0 1.2 2.3 2.0 4.5

ECHINODERMATA 23.0 67.0 27.8 94.9 103.7 65.0 79.3

Crinoidea 0.1 <0.1 <0.1

Holothuroidea 3.7 25.4 2.0 2.2 7.4 4.1 4.3

Echinoidea 2.8 3.4 0.3 67.2 0.6 1.5 29.3

Ophiuroidea 15.8 38.0 25.5 23.8 94.3 57.8 44.2

Asteroidea 0.8 0.2 1.8 1.3 1.7 1.5

HEMICHORDATA 0.1 0.2 <0.1 0.1

CHORDATA 29.3 5.1 4.3 22.4 11.4 1.9 16.3

Ascidiacea 29.3 5.1 4.3 22.4 11.4 1.9 16.3

UNIDENTIFIED 2.9 5.6 5.8 6.4 7.0 5.8

Total 1,512.5 609.6 1,236.9 2,243.5 1,002.4 718.2 1,512.2
46 NOAA Technical Report NMFS 140

Table 17

The number ofspecimens of each taxon, expressed as a percentage of the total benthic invertebrate fauna, in relation to

bottom sediments.

Bottom sediments

Taxon Gravel Till Shell Sand Sand-silt Silt-clay All types

PORIFERA 0.3 0.7 <0.1 <0.1 0.2 0.1 0.1

COELENTERATA 3.0 2.6 0.6 2.3 1.5 0.9 2.1

Hydrozoa 1.3 1.1 0.4 0.3 0.3 0.2 0.4

Anthozoa 1.7 1.5 0.2 2.0 1.2 0.7 1.7

Alcyonaria 0.1 <0.1 0.1 0.2 0.1

Zoantharia 0.9 0.6 0.2 2.0 0.9 0.4 1.5

Unidentified 0.6 0.9 <0.1 <0.1 0.2 0.1 0.2

PLATYHELMINTHES 0.1 <0.1 <0.1 <0.1

Turbellaria 0.1 <0.1 <0.1 <0.1

NEMERTEA 0.3 0.2 2.1 0.6 0.5 0.5 0.5

ASCHELMINTHES 0.6 0.2 <0.1 0.2 0.4 0.2

'ematoda 0.6 0.2 <0.1 0.2 0.4 0.2

ANNELIDA 33.4 47.5 34.1 25.4 30.3 32.3 28.1

POGONOPHORA <0.1 0.1 0.3 <0.1

SIPUNCULIDA 0.3 0.7 0.5 0.3 0.5 0.8 0.4

ECHIURA <0.1 <0.1 <0.1

PRIAPULIDA <0.1 <0.1 <0.1

MOLLUSCA 5.5 16.9 22.2 4.5 27.0 49.3 12.4

Polyplacophora 0.1 0.4 <0.1 0.4 0.2 0.1

Gastropoda 2.7 1.5 6.4 0.5 2.2 1.6 1.2

Bivalvia 2.6 12.8 13.9 3.9 23.7 45.9 10.8

Scaphopoda 0.1 2.2 2.0 0.1 0.6 1.5 0.3

Cephalopoda <0.1 <0.1 0.2 <0.1

Uniden tified <0.1 <0.1 <0.1

ARTHROPODA 47.1 9.6 9.6 60.9 27.0 4.7 48.0

Pycnogonida 0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Arachnida <0.1 <0.1

Crustacea 46.9 9.6 9.6 60.9 27.0 4.7 48.0

Ostracoda <0.1 <0.1 <0.1 <0.1

Cirripedia 1.9 0.2 0.4 0.8 5.4 <0.1 1.4

Copepoda <0.1 <0.1 <0.1 <0.1

Cumacea 0.7 0.2 0.6 2.1 1.8 1.0 1.7

Tanaidacea <0.1 <0.1 <0.1

Isopoda 0.4 1.2 0.3 1.0 0.5 0.4 0.8

Amphipoda 42.3 8.1 7.6 56.4 18.9 3.2 43.4

Mysidacea 0.1 0.2 0.1 <0.1 0.2

Decapoda 1.6 0.7 0.4 0.2 <0.1 0.5

Unidentified <0.1 <0.1 0.1 <0.1

BRYOZOA 5.0 0.9 25.5 0.2 0.5 0.2 1.0

BRACHIOPODA 0.9 7.9 28 <0.1 0.2 0.3 0.3

ECHINODERMATA 1.5 11.0 2.2 4.3 10.2 9.0 5.2

Crinoidea <0.1 <0.1 <0.1

Holothuroidea 0.2 4.2 0.2 0.1 0.7 0.6 0.3

Echinoidea 0.2 0.6 <0.1 3.1 0.1 0.2 1.9

Ophiuroidea 1.0 6.2 2.0 1.1 9.2 8.0 2.9

Asteroidea <0.1 <0.1 0.1 0.1 0.2 0.1

HEMICHORDATA <0.1 <0.1 <0.1 <0.1

CHORDATA 1.9 0.8 0.3 1.0 1.1 0.3 1.1

Ascidiacea 1.9 0.8 0.3 1.0 1.1 0.3 1.1

UNIDENTIFIED 0.2 0.9 0.3 0.6 1.0 0.4

Total 100.0 100.0 100.0 100.0 100.0 100.0 100.0
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 47

610

1,513

NUMBER

MEAN PERCENTAGE

ALL TAXA COMPOSITION

i)

fj",,,, {\;{~::." -:::

BOTTOM

TYPE

Gravel

Till

WEIGHT

PERCENTAGE

COMPOSITION

(). . '.' .~ . :.:}:.: .

MEAN

ALL TAXA

181

67

1,237

2,244

1,022

718

Shell

Sand

Sand-silt

Silt-clay

EXPLANATION

223

246

170

86

D CRUSTACEA

ANNELIDA

rn MOLLUSCA

• ECHINODERMATA

OTHER

Figure 17

Quantitative composition of the total macrobenthic invertebrate fauna in

relation to bottom type. Mean number of individuals and mean wet weight,

per square meter of bottom area, and percentage composition, by density

and biomass, of the major taxonomic groups are shown.

large part to occasional large animals. Those groups

that contributed large specimens at those depths were

Sipunculida, Echiura, Echinoidea, and Holothuroidea.

The percentage occurrence of each taxonomic group

in samples in each depth range class is presented in

Table 15.

Relation to Bottom Sediments

A marked disparity in the average density and biomass

of benthic invertebrates was found among the various

kinds of bottom sediments in the New England Region

(Tables 16, 17; Fig. 17). Sand ranked far above the

other sediment types in density, with an average of

2,244 individuals/m2. Three sediment types-gravel,

shelI, and sand-silt-supported a moderate number of

animals; their average densities ranged from 1,022 to

1,513/m2. Lower densities (610 and 718/m2) were found

in tilI and silt-clay sediments.

Distribution of biomass (Tables 18, 19; Fig. 17) was

similar to that of density. Sand and shell supported

high (246 and 223 g/m2) biomasses. Moderate quantities (170 and 181 g/m2) occurred in gravel and sandsilt. Relatively low quantities were encountered in siltclay and tilI sediments.
48 NOAA Technical Report NMFS 140

Table 18

Mean wet weight ofspecimens of each taxon (grams per square meter) in relation to bottom sediments.

Bottom sediments

Taxon Gravel Till Shell Sand Sand-silt Silt-clay All types

PORIFERA 8.62 12.69 5.77 0.34 3.05 0.12 2.24

COELE TERATA 15.29 1.81 0.93 2.00 18.22 3.52 7.33

Hydrozoa 2.60 0.32 0.14 0.34 0.05 0.01 0.52

Anthozoa 12.69 1.49 0.79 1.66 18.17 3.51 6.81

Alcyonaria 0.09 0.22 0.29 0.20 0.20

Zoantharia 12.36 1.09 0.68 1.37 17.25 3.22 6.39

Unidentified 0.24 0.39 0.12 0.07 0.62 0.10 0.22

PLATYHELMINTHES 0.01 0.01 0.01 0.01

Turbellaria 0.01 0.01 0.01 0.01

EMERTEA 0.52 0.06 6.09 0.81 0.83 0.46 0.71

ASCHELMINTHES 0.01 0.01 <0.01 0.01 0.01 0.01

Nematoda 0.01 0.01 <0.01 0.01 0.01 0.01

ANNELlDA 15.52 10.67 15.30 15.00 25.96 16.25 17.41

POGONOPHORA <0.01 0.01 0.01 <0.01

SIPUNCULlDA 0.58 0.29 0.16 0.89 0.81 0.57 0.75

ECHIURA 0.79 0.69 0.30

PRIAPULlDA <0.01 0.02 <0.01

MOLLUSCA 94.44 5.96 167.76 120.99 73.56 18.43 83.64

Polyplacophora 0.72 0.29 0.01 0.06 0.07 0.14

Gastropoda 3.25 0.21 2.13 3.32 1.60 0.22 2.23

Bivalvia 90.22 4.66 165.13 117.44 71.62 17.58 80.95

Scaphopoda 0.25 0.80 0.50 0.22 0.26 0.56 0.32

Cephalopoda <0.01 0.01 0.02 0.01

Unidentified <0.01 <0.01 <0.01

ARTHROPODA 20.32 1.93 6.26 11.92 6.95 0.62 9.41

Pycnogonida 0.05 <0.01 <0.01 <0.01 0.01 0.01

Arachnida <0.01 <0.01

Crustacea 20.27 1.93 6.26 11.92 6.95 0.61 9.40

Ostracoda <0.01 <0.01 <0.01 <0.01

Cirripedia 11.06 0.02 0.45 2.36 4.41 0.01 3.39

Copepoda <0.01 <0.01 <0.01 <0.01

Cumacea 0.06 0.01 0.08 0.20 0.06 0.03 0.11

Tanaidacea <0.01 <0.01 <0.01

Isopoda 0.21 1.36 0.04 0.44 0.14 0.09 0.29

Amphipoda 3.39 0.54 0.92 7.73 1.91 0.18 4.16

Mysidacea 0.01 0.02 0.01 <0.01 0.01

Decapoda 5.56 4.78 1.16 0.41 0.32 1.43

Unidentified <0.01 <0.01 <0.01 <0.01

BRYOZOA 7.39 0.25 ]6.78 0.37 0.05 0.05 1.29

BRACHIOPODA 2.44 15.87 0.22 0.24 0.33 0.28 0.89

ECHINODERMATA 5.66 15.24 3.39 88.43 36.59 42.87 55.00

Crinoidea <0.01 <0.01 <0.01

Holoth uroidea 2.10 4.73 0.40 3.11 25.00 28.74 12.87

Echinoidea 1.98 4.73 2.25 80.68 3.43 7.26 36.75

Ophiuroidea 1.23 5.27 0.74 2.67 5.80 3.26 3.26

Asteroidea 0.34 0.50 1.98 2.27 3.61 2.13

HEMICHORDATA 0.02 0.04 <0.01 0.02

CHORDATA 9.70 1.98 0.09 4.35 2.36 1.95 4.10

Ascidiacea 9.70 1.98 0.09 4.35 2.36 1.95 4.10

UNIDENTIFIED 0.45 0.21 0.23 0.31 0.24 0.27

Total 180.94 66.97 222.75 245.60 169.88 86.10 183.39
Composition and Distribution of Macrobenthic Invertebrate Fauna 49

Table 19

The wet weight of specimens of each taxon, expressed as a percentage of the total benthic invertebrate fauna, in relation

to bottom sediments.

Bottom sediments

Taxon Gravel Till Shell Sand Sand-silt Silt-clay All types

PORIFERA 4.8 19.0 2.6 0.1 1.8 0.1 1.2

COELENTERATA 8.5 2.7 0.4 0.8 10.7 4.1 4.0

Hydrozoa 1.4 0.5 0.1 0.1 <0.1 <0.1 0.3

Anthozoa 7.1 2.2 0.3 0.7 10.7 4.1 3.7

Alcyonaria <0.1 0.1 0.2 0.2 0.1

Zoantharia 6.8 1.6 0.3 0.6 10.2 3.7 3.5

Uniden tified 0.1 0.6 <0.1 <0.1 0.4 0.1 0.1

PLATYHELMINTHES <0.1 <0.1 <0.1 <0.1

Turbellaria <0.1 <0.1 <0.1 <0.1

NEMERTEA 0.3 0.1 '2..7 0.3 0.5 0.5 0.4

ASCHELMINTHES <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Nematoda <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

A ELIDA 8.6 15.9 6.9 6.1 15.3 18.9 9.5

POGONOPHORA <0.1 <0.1 <0.1 <0.1

SIPUNCULIDA 0.3 0.4 0.1 0.4 0.5 0.7 0.4

ECHIURA 0.5 0.8 0.2

PRIAPULIDA <0.1 <0.1 <0.1

MOLLUSCA 52.2 8.9 75.3 49.3 43.3 21.4 45.6

Polyplacophora 0.4 0.4 <0.1 <0.1 0.1 0.1

Gastropoda 1.8 0.3 1.0 1.4 0.9 0.3 1.2

Bivalvia 49.9 7.0 74.1 47.8 42.2 '2.0.4 44.1

Scaphopoda 0.1 1.2 0.2 0.1 0.2 0.6 0.2

Cephalopoda <0.1 <0.1 <0.1 <0.1

Unidentified <0.1 <0.1 <0.1

ARTHROPODA 11.2 2.9 2.8 4.8 4.1 0.7 5.1

Pycnogonida <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Arachnida <0.1 <0.1

Crustacea 11.2 2.9 2.8 4.8 4.1 0.7 5.1

Ostracoda <0.1 <0.1 <0.1 <0.1

Cirripedia 6.1 <0.1 0.2 1.0 2.6 <0.1 1.8

Copepoda <0.1 <0.1 <0.1 <0.1

Cumacea <0.1 <0.1 <0.1 0.1 <0.1 <0.1 0.1

Tanaidacea <0.1 <0.1 <0.1

Isopoda 0.1 2.0 <0.1 0.2 0.1 0.1 0.2

Amphipoda 1.9 0.8 0.4 3.2 l.l 0.2 2.3

Mysidacea <0.1 <0.1 <0.1 <0.1 <0.1

Decapoda 3.1 2.2 0.5 0.2 0.4 0.8

Unidentified <0.1 <0.1 <0.1 <0.1

BRYOZOA 4.1 0.4 7.5 0.2 <0.1 0.1 0.7

BRACHIOPODA 1.4 23.7 0.1 0.1 0.2 0.3 0.5

ECHINODERMATA 3.1 22.8 1.5 36.0 21.6 49.8 30.0

Crinoidea <0.1 <0.1 <0.1

Holothuroidea 1.2 7.1 0.2 1.3 14.8 33.4 7.0

Echinoidea l.l 7.1 1.0 32.8 2.0 8.4 20.0

Ophiuroidea 0.7 7.9 0.3 1.1 3.4 3.8 1.8

Asteroidea 0.2 0.8 0.8 1.3 4.2 1.2

HEMICHORDATA <0.1 <0.1 <0.1 <0.1

CHORDATA 5.4 3.0 <0.1 1.8 1.4 2.3 2.2

Ascidiacea 5.4 3.0 <0.1 1.8 1.4 2.3 2.2

UNIDENTIFIED 0.2 0.3 0.1 0.2 0.3 0.2

Total 100.0 100.0 100.0 100.0 100.0 100.0 100.0
50 NOAA Technical Report NMFS 140

Table 20

Frequency of occurrence (%) of each taxonomic group in the samples in each sediment type.

Bottom sediments

Taxon Gravel Till Shell Sand Sand-silt Silt-clay

PORIFERA 16 41 17 3 8 3

COELENTERATA 47 59 50 38 53 35

Hydrozoa 23 27 33 14 6 4

Anthozoa 24 32 17 24 47 31

Alcyonaria 4 1 10 12

Zoantharia 29 27 33 :Z:Z 35 17

PLAlYHELMINTHES 2 2

Turbellaria 2 2

NEMERTEA 28 14 50 42 47 29

ASCHELMI THES 12 5 4 13 13

Nematoda 12 5 4 13 13

ANNELIDA 97 100 100 94 100 97

POGO OPHORA <I 8 16

SIPUNCULlDA 20 23 17 25 :Z8 18

ECHIURA 3 5

PRlAPULlDA

MOLLUSCA 75 96 lOa 86 94 93

Polyplacophora 14 27 3 11 9

Gastropoda 43 50 83 44 45 42

Bivalvia 68 96 lOa 7'J 92 91

Scaphopoda 17 46 33 10 30 31

Cephalopoda 1 I 1

ARTHROPODA 95 91 lOa 96 il2 68

Pycnogonida 11 5 I 1 I

Arachnida <1

Crustacea 'J5 91 lOa 96 82 68

Ostracoda I 1 <I

Cirripedia 13 9 17 2 3 I

Copepoda <1 I <I

Cumacea 26 14 17 50 34 22

Tanaidacea 2 5

Isopoda 32 50 17 51 28 16

Amphipoda 93 77 100 94 74 51

Mysidacea 4 6 3 I

Decapoda 35 33 37 7 4

BRYOZOA 27 32 50 il 10 4

BRACHIOPODA 14 41 17 2 3 3

ECHINODERMATA 51 X6 50 72 7il 7il

Crinoidea I <I

Holothuroidea 16 50 17 il 36 23

Echinoidea 14 3:Z 17 47 X 15

Ophiuroidea 40 6X 33 :Z'J 64 62

Asteroidea 8 9 14 16 14

HEMICHORDATA <1 <1

CHORDATA 22 3:Z 17 1l.J 15 11

Ascidiacea n 3:Z 17 19 15 11
Composition and Distribution of Macrobenthic Invertebrate Fauna 51

Figure 18

Medium coarse sand bottom at a depth of 87 m on the western Nova Scotia shelf. This locality also

contains some fine angular to rounded gravel to 7 cm in size. Bivalve shell fragments litter the bottom;

an intact mussel valve is visible in the upper right-center portion of the frame. The camera tripping

weight is visible at upper left-center. Photograph was taken at station 1165c, located at lat. 44°09' N.,

long. 66°29' W.

The percentage occurrence of each taxonomic group

in samples in each sediment type is presented in

Table 20.

Photographs of the sea bottom (Figs. 18 to 24) taken

with the photographic system in the Campbell Grab

show the sediment surface and associated fauna in different bottom types in different subareas within the

New England region. Sediment types range from coarse

(gravels and cobbles) to fine (silty sands). The cameratripping weight, visible in each photograph, serves

as a possible indicator of the amount ofsilt·day contained

in the sediment depending on the quantity of material

disturbed and entering into suspension upon disturbance.

Relation to Water Temperature

The abundance of the New England region macrobenthos, in general, was related directly to the annual

range in water temperature. In areas with a small annual range in water temperature, the density of animals

was low and the biomass small. Conversely, where the

annual range in water temperature was large, the density of animals was high (Tables 21, 22; Fig. 25), and

biomass large (Tables 23, 24; Fig. 25). In areas subject

to annual water temperature ranges of less than 4°C,

average density was only 431/m2 and biomass only

46 g/m2. Although there are some inconsistencies, in
52 NOAA Technical Report NMFS 140

Figure 19

Brown silty sand bottom at ] 16 m depth southwest of Grand Y1anan Island. A substantial number

of brittlestars (Ophiuroidea) of various sizes are clearly visible, as are several burrow holes. The

fine nature of the bottom at this locality is indicated by the cloud ofsediment raised by the camera

tripping weight striking the bottom at upper left. Photograph was taken at station] 173, located at

lat. 44°28' , ., long. 67°15' W.

general, density increased to 4,038/m2 where the temperature range was greater than 20°C, and biomass rose

to 467 g/m2 where the range was 16 to 19.9°C. This

close relationship ofsparse fauna in stable temperatJre

areas and rich fauna in unstable areas undergoi ng wide

temperature fluctuations is not necessarily a direct causeand-effect relationship of temperature alone but is the

result of a combination of temperature and other environmental factors. Water masses that remain relatively

constant in temperature and are unchanging in most

other physical and chemical properties also tend to

ha\'e more uniform biological components. In the New

England region these water masses tend to be deeper,

and therefore colder, darker, and lower in nutrients,

plankton, nekton, as well as benthos.

The percentage occurrence of each taxonomic group

in samples in each temperature range class is presented

in Table 25.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 53

Figure 20

Sand and gravel bottom at 194 m depth in the southwestern Gulf of Maine east of Cape Cod.

Sediment contains some pebbles to 8 cm. Five large burrowing sea anemones (Coelenterata,

Ceriantharia) are visible. Camera tripping weight is at extreme right-center. Photograph was

taken at station 1052, located at lat. 42°09' N., long. 69°14' W.

Relation to Sediment Organic Carbon

Two general cause-and-effect relationships were considered in prompting the analysis of organic carbon

content in the sediments. The first was the possibility of

a high standing crop of benthic animals in areas of high

organic carbon, due to the probability of a greater food

supply in those areas; or secondly, the converse could

apply, namely that areas containing large standing crops

would be high in organic carbon due to biogenic activities (fecal deposits, mortality, etc.). Regardless which

prevailed, high organic carbon content would be associated with high abundance.

The analysis did not reveal any clear-cut correlation

between sediment organic carbon content and benthic

faunal abundance. A few faunal groups exhibited good

correlations, some positive and some negative, but by

and large they were exceptional.

Highest average densities (Tables 26, 27; Fig. 26) of

macrobenthos occurred in areas where the percentage
54 NOAA Technical Report NMFS 140

Figure 21

Medium grained brown sand and shell bottom at 88 m depth on east-central Georges Bank. Shell

fraction is composed of whole and broken bivalve and gastropod shells. A large sea scallop

(Placopecten magellanicus) valve is visible in the center of the frame; a small starfish is at lower right;

a small hermit crab (Paguridae) may be seen at lower right-center on a clump of tubes. The

camera trip weight is visible in the upper left corner. Photograph was taken at station 1127,

located at lat. 41 °30' ., long. 66°32' W.

of organic carbon in the sediments ranged from 3.0 to

4.99 (2,588/m2 ) and 2.00 to 2.99 (2,042/m2). In areas

containing from 0.01 to 0.49 and from 1.00 to 1.49

percent organic carbon, densities were about 50% lower

(1,858/m2, and 1,015/m2, respectively). Significantly

lower densities occurred in other ranges; lowest densities (441m2) were recorded in areas containing the

greatest amount of organic carbon (5% and greater).

Density values over the whole range of organic carbon

content as well as between adjacent classes were too

variable to show any definite trends.

Correlations between the distribution of biomass and

sediment organic carbon (Tables 28, 29; Fig. 26) were

somewhat better but not uniform enough to denote any

positive trends. Largest biomass (959 g/m2 ) was recorded

were organic carbon ranged between 2.00 and 2.99%, and

next largest was 809 g/m2 in the 3.00 to 4.99% range, but

biomass was less than 1 g/m2 in ranges of 5% and over.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 55

Figure 22

Coarse brown sand with broken shells and gravel, to 15 em, bottom at 86 m depth on eastern

Georges Bank. Camera tripping weight is visible at upper left-center edge of frame. Photograph

was taken at station 1130, located at lat. 42°01' N., long. 66°31' W.

Biomass of218 g/m2 occurred in the 0.01 to 0.49% range

and became rather uniform (89 to 97 g/m2) in organic

carbon contents from 0.5 to 1.99%.

The percen tage occurrence of each taxonomic group

in samples in each sediment organic content class is

presented in Table 30.

A comparison of the similarities and differences between the macrobenthos of the New England region

and that of the Middle Atlantic Bight region in relation

to the environmental parameters described above is

contained in Sherman et al. (1988).

Taxonomic Groups

This section deals, in turn, with each taxonomic component of the ew England region macrobenthos arranged in the phylogenetic order presented in Table 4.

Included for each taxon are general remarks relating

to overall abundance and frequency of occurrence, as

well as to aspects of the natural history of some of the

common forms encountered; these remarks are followed by discussions dealing with geographic and bathymetric distribution and the quantitative relationship of
56 NOAA Technical Report NMFS 140

Figure 23

Medium to coarse brown sand bottom at 44 m depth on the continental shelf south of Long

Island, New York. Several sand dollars (f:chinarachnius panna) are visible; the camera tripping

weight is in the upper left corner. Photograph was taken at station 1279, located at lat. 40°30' N.,

long. 72°30' W.

this distribution to bottom sediments, bottom water

temperature, and sediment organic carbon content.

This arrangement was chosen for ease of reference

wherein all the information pertaining to a given taxon

is presented in one place, rather than dispersed among

several subsections, each dealing with a single abiotic

parameter. It is our hope that this arrangement will

allow the reader to make comparisons more easily

among the various taxonomic groups.

Porifera

Porifera constituted a moderately small proportion of

the fauna. They were generally uncommon on the continental shelf and uncommon to rare on the continental slope and rise. Specimens in our samples ranged from

small (2-mm) boring species occurring on or in mollusk

shells to moderately large (10 to 20 cm) Polymastia and

Myxilla usually found on cobbles and boulders.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 57

Figure 24

Green silty sand and gravel bottom at 450 m depth on the southern New England continental

slope. Gravel ranged to 2 em. The tubes of two soda straw worms (Hyalinoecia tubicola) are visible at

left edge of the frame, and the tracks created by their movements are also visible. The silty nature

of the bottom is evident by the cloud created by the camera tripping weight at right-center.

Photograph taken at station 1325, located at lat. 39°20' ., long. 72°09' W.

Porifera were most common in the Nova Scotia region and along the coast of Maine. These areas have

stable substrates and moderate to strong water currents.

Colorful species were rare, limited mainly to the red

Microciona. Shades of browns and grays were the most

common colors. The colorless Hexactinellida (glass

sponges) occurred only in deep water, greater than 500 m.

Porifera occurred in 71 samples (7% of total); density

averaged 1.5/m2 and biomass averaged 2.2 g/m2 (Table 5).

Geof!7aphic Distribution

Porifera were clearly more abundant off Nova Scotia

and in the Gulf of Maine than to the south and west

(Fig. 27). They were common along the western coast

of Nova Scotia, particularly in the mouth of the Bay of

Fundy, along the Maine coast, and in the eastern Gulf

of Maine. The scarcity of sponges on the continental

shelf between central Georges Bank and NewJersey was

especially noteworthy.
58 NOAA Technical Report NMFS 140

Table 21

Mean number ofspecimens of each taxon per square meter in relation to the annual range in bottom water temperature.

Annual range in water temperature (degrees Celsius)

Taxon 0-3.9 4-7.9 8-11.9 12-15.9 16-19.9 20-23.9 All ranges

PORIFERA 1.1 1.7 1.1 2.7 1.9 0.4 1.5

COELENTERATA 6.6 14.7 24.0 105.3 14.8 93.9 32.1

Hydrozoa 0.9 5.0 4.5 4.5 36.8 45.9 6.4

Anthozoa 5.7 9.7 19.5 100.8 8.0 48.0 25.7

Alcyonaria 1.1 1.2 0.2 1.7 0.8

Zoantharia 2.4 7.9 18.6 98.4 7.2 4.4 22.6

Unidentified 2.1 0.6 0.7 0.7 0.8 43.6 2.2

PLATYHELMINTHES 0.2 1.0 0.1 <0.1 0.4 0.4

Turbellaria 0.2 1.0 0.1 <0.1 0.4 0.4

NEMERTEA 2.4 4.2 9.0 25.3 4.4 2.9 8.2

ASCHELMINTHES 3.5 2.0 4.3 0.7 0.2 2.8

Nematoda 3.5 2.0 4.3 0.7 0.2 2.8

ANNELIDA 211.5 513.2 568.2 279.8 370.4 1,697.8 425.0

POGONOPHORA 1.5 0.7 0.1 0.6

SIPU CULIDA 4.5 9.0 8.5 2.8 2.6 5.9

ECHIURA 0.1 0.3 0.4 0.1

PRIAPULIDA <0.1 <0.1

MOLLUSCA 84.0 129.2 129.2 344.9 344.5 1,242.2 188.0

Polyplacophora 1.4 0.8 0.8 2.0 7.3 1.5

Gastropoda 9.4 15.1 14.8 13.1 84.9 47.4 17.8

Bivalvia 64.4 102.9 110.1 329.0 252.0 1,194.8 163.1

Scaphopoda 8.9 10.3 2.1 0.8 0.2 5.1

Cephalopoda <0.1 0.1 J.l 0.4

Unidentified <0.1 0.3 0.1

ARTHROPODA 64.9 338.0 1,475.6 768.3 1,040.1 903.5 726.2

Pycnogonida 0.1 0.4 0.7 0.1 0.2 1.5 0.3

Arachnida <0.1 <0.1

Crustacea 64.8 337.6 1,474.9 768.2 1,039.9 902.0 725.9

Ostracoda 0.1 <0.1 <0.1

Cirripedia 0.3 8.7 14.2 14.0 154.4 196.1 21.8

Copepoda 0.1 0.1 <0.1

Cumacea 2.3 9.6 56.3 21.3 43.5 19.0 25.8

Tanaidacea 0.1 0.1 <0.1

Isopoda 2.6 3.8 15.7 25.1 7.0 67.3 12.1

Amphipoda 58.2 311.6 1,371.9 694.7 808.5 598.2 655.8

Mysidacea <0.1 0.8 3.4 6.3 3.4 6.1 2.5

Decapoda 0.9 3.2 13.2 6.4 23.2 13.4 7.5

Unidentified 0.5 0.2 0.4 1.9 0.3

BRYOZOA 3.0 9.5 15.8 35.1 28.3 66.0 15.7

BRACHIOPODA 8.5 5.9 3.0 4.5

ECHINODERMATA 32.9 61.5 104.7 171.2 31.5 21.0 79.3

Crinoidea <0.1 <0.1

Holothuroidea 6.3 4.1 3.7 2.8 1.7 3.5 4.3

Echinoidea 1.4 4.9 40.4 !i3.1 26.8 14.6 29.3

Ophiuroidea 24.8 50.4 57.9 74.1 2.0 2.4 44.2

Asteroidea 0.4 2.2 2.7 1.2 0.9 0.5 1.5

HEMICHORDATA <0.1 0.2 0.2 0.1

CHORDATA 2.0 2.3 35.1 6.0 58.5 4.3 16.3

Ascidiacea 2.0 2.3 35.1 6.0 58.5 4.3 16.3

U IDE TIFIED 5.6 5.7 6.6 4.2 6.6 5.2 5.8

Total 432.2 1,097.9 2,386.~ 1,746.4 1,934.2 4,037.5 1,512.2
Composition and Distribution of Macrobenthic Invertebrate Fauna 59

Table 22

The number ofspecimens of each taxon, expressed as a percentage of the total benthic invertebrate fauna, in relation to

the annual range in bottom water temperature.

Annual range in water temperature (degrees Celsius)

Taxon 0-3.9 4-7.9 8-11.9 12-15.9 16--19.9 20-23.9 All ranges

PORIFERA 0.3 0.2 <0.1 0.2 0.1 <0.1 0.1

COELENTERATA 1.5 1.3 1.0 6.0 2.3 2.3 2.1

Hydrozoa 02 0.4 0.2 0.3 1.9 I.I 0.4

Anthozoa 1.3 0.9 0.8 5.7 0.4 1.2 1.7

Alcyonaria 0.3 0.1 <0.1 0.1 0.1

Zoantharia 0.6 0.7 0.8 5.6 0.4 0.1 1.5

Unidentified 0.5 0.1 <0.1 <0.1 <0.1 I.I 0.2

PLAlYHELMINTHES <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Turbellaria <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

NEMERTEA 06 0.4 0.4 1.4 0.2 0.1 0.5

ASCHELMINTHES 0.8 0.2 0.2 <0.1 <0.1 0.2

Nematoda 0.8 02 0.2 <0.1 <0.1 0.2

A NELIDA 48.9 46.7 23.8 16.0 19.2 42.0 28.1

POGO OPHORA 0.3 0.1 <0.1 <0.1

SIPU CULIDA 1.0 0.8 0.4 0.2 0.1 0.4

ECHIURA <0.1 <0.1 <0.1 <0.1

PRIAPULIDA <0.1 <0.1

MOLLUSCA 19.4 11.8 5.4 19.8 17.8 30.8 12.4

Polyplacophora 0.3 0.1 <0.1 0.1 0.4 0.1

Gastropoda 2.2 1.4 0.6 0.8 4.4 1.2 1.2

Bivalvia 14.9 9.4 4.6 18.8 13.0 29.6 10.8

Scaphopoda 2.0 0.9 0.1 <0.1 <0.1 0.3

Cephalopoda <0.1 <0.1 <0.1 <0.1

Unidentified <0.1 <0.1 <0.1

ARTHROPODA 15.0 30.8 61.8 44.0 53.8 22.4 48.0

Pycnogonida <0.1 <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Arachnida <0.1 <0.1

Crustacea 15.0 30.8 61.8 44.0 53.8 22.3 48.0

Ostracoda <0.1 <0.1 <0.1

Cirripedia 0.1 0.8 0.6 0.8 8.0 4.9 1.4

Copepoda <0.1 <0.1 <0.1

Cumacea 0.5 0.9 2.4 1.2 2.2 0.5 1.7

Tanaidacea <0.1 <0.1 <0.1

Isopoda 0.6 0.3 0.7 1.4 0.4 1.7 0.8

Amphipoda 13.5 28.4 57.5 39.8 41.8 14.8 43.4

Mysidacea 0.1 0.1 0.1 0.4 0.2 0.2 0.2

Decapoda 0.2 0.3 0.6 0.4 1.2 0.3 0.5

Unidentified 0.1 <0.1 <0.1 <0.1 <0.1

BRYOZOA 0.7 0.9 0.7 2.0 1.5 1.6 1.0

BRACHIOPODA 2.0 0.5 0.1 0.3

ECHINODERMATA 7.6 5.6 4.4 9.8 1.6 0.5 5.2

Crinoidea <0.1 <0.1

Holothuroidea 1.5 0.4 0.2 0.2 0.1 0.1 0.3

Echinoidea 0.3 0.4 I.7 5.3 0.6 0.4 1.9

Ophiuroidea 5.8 4.6 2.4 4.2 0.1 O. I 2.9

Asteroidea 0.1 0.2 0.1 0.1 <0.1 <0.1 0.1

HEMICHORDATA <0.1 <0.1 <0.1 <0.1

CHORDATA 0.5 0.2 1.5 0.3 3.0 0.1 I.I

Ascidiacea 0.5 0.2 1.5 0.3 3.0 0.1 I.I

UNIDENTIFIED 1.3 0.5 0.3 0.2 0.3 0.1 0.4

Total 100.0 100.0 100.0 100.0 100.0 100.0 100.0
60 NOAA Technical Report NMFS 140

Table 23

Mean wet weight of specimens of each taxon (grams per square meter) in relation to the annual range in bottom water

temperature.

Annual range in water temperature (degrees Celsius)

Taxon 0-3.9 4-7.9 8-11.9 12-15.9 16-19.9 20-23.9 All ranges

PORIFERA 0.86 2.50 4.63 1.08 0.10 <0.01 2.24

COELENTERATA 9.10 6.74 8.74 2.85 5.15 2.47 7.33

Hydrozoa 0.01 2.02 0.15 0.45 1.54 0.68 0.52

Anthozoa 9.09 4.72 8.59 2.40 3.61 1.79 6.81

Alcyonaria 0.42 O.ll 0.04 0.31 0.20

Zoantharia 8.17 4.59 8.49 2.00 3.57 0.86 6.39

Unidentified 0.48 0.02 0.07 0.09 0.05 093 0.22

PLAlYHELMINTHES 0.01 0.01 <0.01 <0.01 0.09 0.01

Turbellaria 0.01 0.01 <0.01 <0.01 0.09 0.01

NEMERTEA 0.22 0.53 0.89 1.22 1.38 1.11 0.71

ASCHELMINTHES 0.01 0.01 0.01 0.01 <0.01 0.01

Nematoda 0.01 0.01 0.01 0.01 <0.01 0.01

ANNELIDA 9.69 14.75 24.31 13.39 28.48 39.74 17.41

POGONOPHORA 0.01 <0.01 <0.01 <0.01

SIPUNCULIDA 0.73 1.01 0.97 0.34 0.33 0.75

ECHIURA 0.90 0.02 0.39 0.30

PRIAPULIDA 0.01 <0.01

MOLLUSCA 7.57 22.13 128.83 109.06 340.39 87.55 83.64

Polyplacophora 0.12 0.09 0.01 0.04 1.33 0.14

Gastropoda 0.26 1.32 3.75 2.67 4.96 4.09 2.23

Bivalvia 6.76 19.79 124.94 106.26 334.10 83.46 80.95

Scaphopoda 0.43 0.92 O.ll 0.08 <0.01 0.32

Cephalopoda <0.01 0.02 0.02 0.01

Unidentified <0.01 <0.01 <0.01

ARTHROPODA 1.22 3.42 16.45 12.52 16.04 24.65 9.41

Pycnogonida <0.01 0.03 0.01 <0.01 <0.01 0.01 0.01

Arachnida <0.01 <0.01

Crustacea 1.22 3.39 16.44 12.52 16.04 24.64 9.40

Ostracoda <0.01 <0.01 <0.01

Cirripedia 0.05 0.22 6.39 3.52 5.32 20.32 3.39

Copepoda <O.OJ <0.01 <0.01

Cumacea 0.02 0.04 0.26 0.07 0.09 0.06 0.11

Tanaidacea <0.01 <0.01 <0.01

Isopoda 0.27 0.08 0.42 0.34 0.07 0.41 0.29

Amphipoda 0.41 2.02 8.12 5.31 6.27 2.56 4.16

Mysidacea <0.01 <0.01 0.02 0.02 0.03 0.02 0.01

Decapoda 0.47 1.02 1.23 3.24 4.26 1.25 1.43

Unidentified <0.01 <0.01 <0.01 0.02 <0.01

BRYOZOA 0.28 0.56 2.10 2.43 0.87 2.45 1.29

BRACHIOPODA 1.93 0.73 0.57 0.89

ECHINODERMATA 12.24 31.35 35.48 172.83 59.63 263.42 55.00

Crinoidea <0.01 <0.01

Holothuroidea 3.10 7.77 11.75 18.65 18.15 127.96 12.87

Echinoidea 6.00 14.29 17.63 148.09 37.65 134.46 36.75

Ophiuroidea 3.05 5.82 3.28 2.77 0.22 0.53 3.26

Asteroidea 0.09 3.48 2.82 3.33 3.61 0.46 2.13

HEMICHORDATA <0.01 0.04 0.03 0.02

CHORDATA 0.55 4.14 7.28 1.43 14.40 0.56 4.10

Ascidiacea 0.55 4.14 728 1.43 14.40 0.56 4.10

UNIDENTIFIED 0.16 0.22 0.36 0.42 0.26 0.09 0.27

Total 45.48 88.11 230.67 317.62 467.42 422.12 183.39
Composition and Distribution of Macrobenthic Invertebrate Fauna 61

Table 24

The wet weight ofspecimens of each taxon, expressed as a percentage of the total benthic invertebrate fauna, in relation

to the annual range in bottom water temperature.

Annual range in water temperature (degrees Celsius)

Taxon 0-3.9 4-7.9 8-11.9 12-15.9 16-19.9 20-23.9 All ranges

PORIFERA 1.9 2.8 2.0 0.3 <0.1 <0.1 1.2

COELENTERATA 20.0 7.6 3.8 0.9 1.1 0.6 4.0

Hydrozoa <0.1 2.3 0.1 0.1 0.3 0.2 0.3

Anthozoa 20.0 5.3 3.7 0.8 0.8 0.4 3.7

Alcyonaria 0.9 0.1 <0.1 0.1 0.1

Zoantharia 18.0 5.2 3.7 0.6 0.8 0.2 3.5

Unidentified 1.1 <0.1 <0.1 <0.1 <0.1 0.2 0.1

PLATYHELMINTHES <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Turbellaria <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

NEMERTEA 0.5 0.6 0.4 0.4 0.3 0.3 0.4

ASCHELMINTHES <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Nematoda <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

ANNELIDA 21.3 16.8 10.5 4.2 6.1 9.4 9.5

POGONOPHORA <0.1 <0.1 <0.1 <0.1

SIPUNCULIDA 1.6 1.2 0.4 0.1 0.1 0.4

ECHIURA 2.0 <0.1 0.1 0.2

PRIAPULIDA <0.1 <0.1

MOLLUSCA 16.6 25.1 55.8 34.3 72.8 20.7 45.6

Polyplacophora 0.3 0.1 <0.1 <0.1 0.3 0.1

Gastropoda 0.6 1.5 1.6 0.8 1.1 1.0 1.2

Bivalvia 14.9 22.5 54.2 33.4 71.5 19.8 44.1

Scaphopoda 1.0 1.0 <0.1 <0.1 <0.1 0.2

Cephalopoda <0.1 <0.1 <0.1 <0.1

Uniden tified <0.1 <0.1 <0.1

ARTHROPODA 2.7 3.9 7.1 3.9 3.4 5.8 5.1

Pycnogonida <0.1 <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Arachnida <0.1 <0.1

Crustacea 2.7 3.8 7.1 3.9 3.4 5.8 5.1

Ostracoda <0.1 <0.1 <0.1

Cirripedia 0.1 0.2 2.8 1.1 1.1 4.8 1.8

Copepoda <0.1 <0.1 <0.1

Cumacea <0.1 <0.1 0.1 <0.1 <0.1 <0.1 0.1

Tanaidacea <0.1 <0.1 <0.1

Isopoda 0.6 0.1 0.2 0.1 <0.1 0.1 0.2

Amphipoda 0.9 2.3 3.5 1.7 1.3 0.6 2.3

Mysidacea <0.1 <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Decapoda 1.0 1.2 0.5 1.0 0.9 0.3 0.8

Unidentified <0.1 <0.1 <0.1 <0.1 <0.1

BRYOZOA 0.6 0.6 0.9 0.8 0.2 0.6 0.7

BRACHIOPODA 4.2 0.8 0.2 0.5

ECHINODERMATA 26.9 35.6 15.4 54.4 12.8 62.4 30.0

Crinoidea <0.1 <0.1

Holothuroidea 6.8 8.8 5.1 5.9 3.9 30.3 7.0

Echinoidea 13.2 16.2 7.6 46.6 8.1 31.8 20.0

Ophiuroidea 6.7 6.6 1.4 0.9 <0.1 0.1 1.8

Asteroidea 0.2 4.0 1.2 1.0 0.8 0.1 1.2

HEMICHORDATA <0.1 <0.1 <0.1 <0.1

CHORDATA 1.2 4.7 3.2 0.4 3.1 0.1 2.2

Ascidiacea 1.2 4.7 3.2 0.4 3.1 0.1 2.2

UNIDENTIFIED 0.4 0.2 0.2 0.1 0.1 <0.1 0.2

Total 100.0 100.0 100.0 100.0 100.0 100.0 100.0
62 NOAA Technical Report NMFS 140

WEIGHT

TEMPERATURE PERCENTAGE MEAN

RANGE COMPOSITION ALL TAXA

432

1,098

2,386

1,746

1,934

4,038

o to 3.9

4 to 7.9

8 to 11.9

12 to 15.9

16 to 19.9

20 to 23.9

EXPLANATION

D CRUSlACEA MOLLUSCA

ANNELIDA • fCHINOOEIMATA

OTHEI

46

88

231

318

467

422

Figure 25

Quantitative composition of the total macrobenthic invertebrate fauna in

relation to the annual range in bottom water temperature. Mean number

of individuals and mean wet weight, per square meter of bottom area, and

percentage composition, by density and biomass, for the major taxonomic

groups are shown.

A pronounced decrease in abundance of sponges in

each geographic region from ova Scotia southwestward to Southern New England was evident in weight,

number, and frequency of occurrence (Tables 5-10;

Fig. 28). Porifera were particularly scarce in the Southern New England subarea.

Bathymetric Distribution

Porifera were moderately common to scarce on the

continental shelf and upper continental slope (to 500

m),1 to 3 individuals/m2, and were very scarce «0.4/

m2) at depths >500 m (Table 11; Fig. 29). Biomass also

was moderate on the continental shelf and upper slope

and small on the middle and lower slope and continen-
Composition and Distribution of Macrobenthic Invertebrate Fauna 63

c 1

959

17

19

97

96

211

109 •

'

,J[{%?;~ ....:

m.•...... ..•.•.·

WEIGHT

PERCENTAGE MEAN

COMPOSITION ALL TAXA

•

v< ..•.. .'.',

0.00

,ERCENT

ORGANIC

CARlON

44

741

914

1,151

2,511

2,042

HUMIER

MEAN 'EIlCENTAGE

ALL::~A W"

f) 0.01 .. 0."

0.50 to 0.99 9

1.00 to 1.49 •....0 t. 1."

() 2.00 t. 2."

()3.00 ......

Us.oo t. +

EXPLANATION

D (IUSTACfA

ANNELIDA

MOLLUSCA

• ECHINODERMATA

D OTHEl

Figure 26

Quantitative composition of the total macrobenthic invertebrate fauna in

relation to organic carbon in the sediments. Mean number of individuals

and mean wet weight, per square meter of bottom, and percentage composition, by density and biomass, for the major taxonomic groups are shown.

tal rise (Table 13; Fig. 29). In the various depth classes

average biomass ranged from 0.02 to 3.1 g/m2. In deeper

water the average weights were only 0.02 to 0.6 g/m2.

In all depths Porifera accounted for a small percentage «1 %) of the total number of benthic animals

(Table 12). The percentage of the total weight of the
64 NOAA Technical Report NMFS 140

Table 25

Frequency of occurrence (%) of each taxonomic group in the samples in each temperature range class.

Annual range in water temperature (degrees Celsius)

Taxon 0-3.9 4-7.9 8-11.9 12-15.9 16-19.9 20-23.9

PORIFERA 8 15 4 2 8 4

COELENTERATA 36 42 45 45 49 29

Hydrozoa 4 13 15 14 26 18

Anthozoa 32 29 30 31 23 11

Alcyonaria 11 5 2 8

Zoantharia 17 29 30 28 24 14

PLAlYHELMINTHES 3 2 4

Turbellaria 3 2 4

EMERTEA 25 28 49 50 52 11

ASCHELMINTHES 17 6 7 5 2

Nematoda 17 6 7 5 2

ANNELIDA 97 95 97 92 97 96

POGONOPHORA 15 3 <1

SIPUNCULIDA 25 29 29 12 5

ECHIURA 4 3

PRIAPULIDA

MOLLUSCA 91 83 87 87 94 93

Polyplacophora 10 10 6 7 5

Gastropoda 44 46 44 38 57 43

Bivalvia 87 75 81 83 90 93

Scaphopoda 36 37 10 3 2

Cephalopoda <1 2 <1

ARTHROPODA 70 89 98 98 90 86

Pycnogonida 1 4 3 1 2 4

Arachnida 1

Crustacea 70 89 98 98 90 86

Ostracoda 1 1

Cirripedia 2 4 4 4 5 18

Copepoda 1 1

Cumacea 19 37 54 38 31 36

Tanaidacea 4 1

Isopoda 20 27 46 61 31 43

Amphipoda 57 83 97 92 82 71

Mysidacea <1 2 3 9 13 18

Decapoda 6 15 35 31 53 18

BRYOZOA 8 13 11 11 21 18

BRACHIOPODA 10 8 2

ECHINODERMATA 77 78 72 71 45 43

Crinoidea 1

Holothuroidea 25 20 14 18 8 21

Echinoidea 16 26 33 44 24 14

Ophiuroidea 64 62 35 29 15 11

Asteroidea 6 16 21 12 11 4

HEMICHORDATA <1

CHORDATA 12 11 24 17 19 11

Ascidiacea 12 11 24 17 19 11
Composition and Distrihution of Macrobenthic Invertebrate Fauna 65

Table 26

Mean number of specimens of each taxon per square meter in relation to sediment organic carbon content.

Sediment organic carbon content (percent)

Taxon 0.00 0.01-0.49 0.50-0.99 1.00-1.49 1.50-1.99 2.00-2.99 3.00-4.99 5.00+ All ranges

PORIFERA 2.6 2.7 0.5 0.4 0.2 1.7

COELENTERATA 17.8 26.8 10.1 5.3 5.3 17.7

Hydrozoa 8.7 9.2 3.4 0.6 2.5 7.5

Anthozoa 9.1 17.6 6.7 4.7 2.8 10.2

Alcyonaria 0.6 1.2 0.4 0.6

Zoantharia 7.4 6.8 4.4 4.6 2.8 6.6

Unidentified 1.1 9.6 1.9 0.1 3.0

PLA1YHELMINTHES 0.6 0.1 0.9 0.4

Turbellaria 0.6 0.1 0.9 0.4

NEMERTEA 10.8 5.5 4.4 4.4 3.5 1.7 4.9

ASCHELMINTHES 3.l 1.0 2.8 0.4 2.3

Nematoda 3.1 1.0 2.8 0.4 2.3

ANNELIDA 64.0 503.6 234.7 319.1 195.9 406.9 81.3 11.0 395.7

POGONOPHORA 0.2 2.2 1.2 0.1 0.8

SIPUNCULIDA 2.2 6.9 7.3 1.0 0.1 5.7

ECHIURA <0.1 0.3 0.1 0.1

PRIAPULIDA <0.1 <0.1 0.1 <0.1

MOLLUSCA 68.8 139.1 146.7 481.7 602.1 794.7 1,119.5 223.3

Polyplacophora 2.5 0.7 1.2 0.1 1.7

Gastropoda 2.4 20.1 12.6 25.7 44.9 39.4 20.6

Bivalvia 64.2 113.7 125.6 442.3 545.2 750.7 1,119.5 195.5

Scaphopoda 2.2 2.7 5.7 12.5 11.9 4.6 5.0

Cephalopoda 0.1 2.1 0.5

Unidentified <0.1 <0.1

ARTHROPODA 21.0 1,066.3 220.3 86.0 34.4 818.3 1,357.1 22.0 690.4

Pycnogonida 0.4 0.3 0.1 0.3

Arachnida <0.1 <0.1

Crustacea 21.0 1,065.9 220.0 85.9 34.4 818.3 1,357.1 22.0 690.1

Ostracoda <0.1 0.1 <0.1

Cirripedia 19.1 39.3 612.8 83.8 31.1

Copepoda <0.1 <0.1 0.1 <0.1

Cumacea 2.6 25.2 6.1 3.8 6.2 122.8 18.7

Tanaidacea <0.1 0.2 0.1

Isopoda 8.6 13.3 5.0 3.5 0.8 7.9 17.8 9.4

Amphipoda 9.8 1,000.1 164.1 78.0 24.4 66.0 1,255.5 22.0 624.4

Mysidacea 1.6 0.8 1.8 0.9 1.2

Decapoda 6.4 3.4 0.5 1.2 7.9 4.7

Unidentified 0.2 1.0 0.3

BRYOZOA 7.8 21.4 1.2 1.7 35.1 26.5 14.9

BRACHIOPODA 16.0 5.6 5.1 2.8 4.7

ECHI ODERMATA 28.0 57.8 79.6 90.5 33.5 6.6 3.3 63.6

Crinoidea <0.1 <0.1 <0.1

Holothuroidea 18.2 4.2 6.1 7.7 2.2 1.7 3.3 4.9

Echinoidea 2.6 23.8 0.8 0.6 1.0 13.8

Ophiuroidea 7.2 28.7 71.7 80.6 29.9 4.9 43.7

Asteroidea 1.1 1.0 1.6 0.4 1.0

HEMICHORDATA 0.2 0.2 0.1 0.1

CHORDATA 21.3 3.8 6.4 4.4 13.8

Ascidiacea 21.3 3.8 6.4 4.4 13.8

U IDE TIFIED 6.3 6.6 6.8 3.1 3.0 11.0 6.1

Total 221.2 1,858.4 740.7 1,015.2 913.8 2,041.5 2,587.5 44.0 1,446.3
66 NOAA Technical Report NMFS 140

Table 27

The number ofspecimens of each taxon, expressed a~ a percentage of the total benthic invertebrate fauna, in relation to

sediment organic carbon content.

Sediment organic carbon content (percent)

Taxon 0.00 0.01-0.49 0.50-0.99 1.00-1.49 1.50-1.99 2.00-2.99 3.00-4.99 5.00+ All ranges

PORIFERA 1.2 0.1 0.1 <0.1 <0.1 0.1

COELENTERATA 1.0 3.6 1.0 0.6 0.3 1.2

Hydrozoa 0.5 1.2 0.3 0.1 0.1 0.5

Anthozoa 0.5 2.4 0.7 0.5 0.1 0.7

Alcyonaria <0.1 0.2 <0.1 <0.1

Zoantharia 0.4 0.9 0.4 0.5 0.1 0.5

Unidentified 0.1 1.3 0.2 <0.1 0.2

PLATYHELMINTHES <0.1 <0.1 <0.1 <0.1

Turbellaria <0.1 <0.1 <0.1 <0.1

NEMERTEA 4.9 0.3 0.6 0.4 0.4 0.1 0.3

ASCHELMINTHES 0.2 0.1 0.3 0.1 0.2

Nematoda 0.2 0.1 0.3 0.1 0.2

ANNELIDA 28.9 27.1 31.7 31.4 21.4 19.9 3.1 25.0 27.4

POGONOPHORA <0.1 0.3 0.1 <0.1 0.1

SIPUNCULIDA 1.0 0.4 1.0 0.1 <0.1 0.4

ECHIURA <0.1 <0.1 <0.1 <0.1

PRIAPULIDA <0.1 <0.1 <0.1 <0.1

MOLLUSCA 31.1 7.5 19.8 47.5 65.9 38.9 43.3 15.4

Polyplacophora 0.1 0.1 0.1 <0.1 0.1

Gastropoda l.l l.l 1.7 2.5 4.9 1.9 1.4

Bivalvia 29.0 6.1 ]7.0 43.6 59.7 36.8 43.3 13.5

Scaphopoda 1.0 0.2 0.8 1.2 1.3 0.2 0.4

Cephalopoda <0.] 0.3 <0.]

Unidentified <0.1 <0.1

ARTHROPODA 9.5 57.4 29.8 8.5 3.8 40.] 52.5 50.0 47.8

Pycnogonida <0.] 0.1 <0.1 <0.1

Arachnida <0.] <0.1

Crustacea 9.5 57.4 29.7 8.5 3.8 40.] 52.5 50.0 47.8

Ostracoda <0.1 <0.] <0.]

Cirripedia 1.0 5.3 30.0 3.3 2.2

Copepoda <0.1 <0.1 <0.1 <0.]

Cumacea 1.2 ].4 0.8 0.4 0.7 6.0 1.3

Tanaidacea <0.1 <0.1 <0.1

Isopoda 3.9 0.7 0.7 0.3 0.] 0.4 0.7 0.7

Amphipoda 4.4 53.8 22.2 7.7 2.7 3.2 48.5 50.0 43.2

Mysidacea 0.] 0.] 0.2 <0.1 0.]

Decapoda 0.3 0.5 0.] 0.1 0.4 0.3

Uniden tified <0.1 0.1 <0.1

BRYOZOA 3.5 1.1 0.2 0.2 3.9 1.0 1.0

BRACHIOPODA 7.2 0.3 0.7 0.3 0.3

ECHINODERMATA 12.7 3.1 10.7 89 3.7 0.3 0.1 4.4

Crinoidea <0.1 <0.1 <0.]

Holothuroidea 8.2 0.2 0.8 0.8 0.2 0.1 0.1 0.3

Echinoidea 1.2 1.3 0.] 0.1 0.] 1.0

Ophiuroidea 3.3 1.5 9.7 7.9 3.3 0.2 3.0

Asteroidea 0.1 0.\ 0.2 0.1 0.1

HEMICHORDATA <0.1 <0. ] <0.1 <0.1

CHORDATA l.l 0.5 06 0.2 1.0

Ascidiacea l.l 0.5 0.6 0.2 1.0

UNIDENTIFIED 0.3 0.9 0.7 0.3 0.2 25.0 0.4

Total 100.0 100.0 100.0 100.0 100.0 ]00.0 100.0 100.0 100.0
Composition and Distribution of Macrobenthic Invertebrate Fauna 67

Table 28

Mean wet weight ofspecimens of each taxon (grams per square meter) in relation to sediment organic carbon content.

Sediment organic carbon content (percent)

Taxon 0.00 0.01-0.49 0.50-0.99 1.00-1.49 1.50-1.99 2.00-2.99 3.00-4.99 5.00+ All ranges

PORIFERA 0.03 5.19 0.12 0.01 <0.01 2.98

COELENTERATA 6.33 18.48 14.57 1.80 0.06 9.57

Hydrozoa 1.06 0.16 0.01 <0.01 0.02 0.64

Anthozoa 5.27 18.32 14.56 1.80 0.04 8.93

A1cyonaria 0.26 0.32 0.13 0.24

Zoantharia 4.91 17.21 14.07 1.79 0.04 8.42

Unidentified 0.10 0.79 0.36 0.01 0.28

PLATYHELMINTHES <0.01 <0.01 0.19 0.01

Turbellaria <0.01 <0.01 0.19 0.01

NEMERTEA 0.53 0.79 0.79 0.38 0.94 0.48 0.74

ASCHELMINTHES 0.01 <0.01 0.02 <0.01 0.01

Nematoda 0.01 <0.01 0.02 <0.01 0.01

ANNELIDA 7.43 15.93 14.88 12.92 26.92 22.60 11.31 0.11 16.01

POGONOPHORA <0.01 0.01 0.01 <0.01 <0.01

SIPUNCULIDA 0.02 0.93 1.13 0.07 <0.01 0.79

ECHIURA 0.01 1.25 1.35 0.44

PRIAPULIDA <0.01 0.02 0.01 0.01

MOLLUSCA 0.80 132.21 25.23 19.95 13.38 811.54 226.87 99.58

Polyplacophora 0.25 0.03 0.02 <0.01 0.15

Gastropoda 0.02 3.83 0.56 0.28 1.09 10.42 2.59

Bivalvia 0.67 127.81 24.42 19.17 12.05 801.02 226.87 96.54

Scaphopoda 0.11 0.31 0.19 0.48 0.24 0.10 0.29

Cephalopoda 0.01 0.03 0.01

Unidentified <0.01 <0.01

ARTHROPODA 0.31 13.28 6.46 1.22 1.02 18.99 9.44 0.11 9.62

Pycnogonida 0.01 0.01 <0.01 <0.01

Arachnida <0.01 <0.01

Crustacea 0.31 13.27 6.45 1.22 1.02 18.99 9.44 0.11 9.61

Ostracoda <0.01 <0.01 <0.01

Cirripedia 6.45 4.11 14.32 1.10 4.86

Copepoda <0.01 <0.01 <0.01 <0.01

Cumacea 0.03 0.09 0.03 0.02 0.01 0.24 0.07

Tanaidacea <0.01 <0.01 <0.01

Isopoda 0.19 0.27 0.24 0.03 0.14 0.06 0.05 0.23

Amphipoda 0.09 5.29 1.76 0.59 0.17 0.21 8.29 0.11 3.54

Mysidacea 0.01 0.01 0.01 0.01 0.01

Decapoda 1.16 0.30 0.58 0.69 4.15 0.91

Unidentified <0.01 <0.01 <0.01

BRYOZOA 0.23 1.95 0.02 0.06 1.21 0.18 1.19

BRACHIOPODA 1.31 0.96 1.74 0.03 0.95

ECHINODERMATA 6.16 35.22 23.25 44.01 43.35 104.47 561.51 38.82

Crinoidea <0.01 <0.01 <0.01

Holothuroidea 5.80 3.97 11.16 30.95 40.90 104.31 561.51 15.66

Echinoidea 0.29 27.90 5.44 1.33 0.60 17.29

Ophiuroidea 0.07 2.32 5.23 6.28 0.56 0.16 3.26

Asteroidea 1.03 1.42 5.45 1.29 1.61

HEMICHORDATA 0.03 0.03 <0.01 0.03

CHORDATA 4.38 1.88 2.03 0.33 3.15

Ascidiacea 4.38 1.88 2.03 0.33 3.15

UNIDENTIFIED 0.33 0.27 0.29 0.10 0.07 0.11 0.29

Total 16.82 217.56 95.95 96.93 88.72 958.73 809.31 0.33 183.18
68 NOAA Technical Report NMFS 140

Table 29

The wet weight ofspecimens of each taxon, expressed as a percentage of the total benthic invertebrate fauna, in relation

to sediment organic carbon content.

Sediment organic carbon content (percent)

Taxon 0.00 0.01-0.49 0.50-0.99 1.00-1.49 1.50-1.99 2.00-2.99 3.00-4.99 5.00+ All ranges

PORIFERA 0.2 2.4 0.1 <0.1 <0.1 1.6

COELENTERATA 2.9 19.3 15.0 20 <0.1 5.2

Hydrozoa 0.5 0.2 <0.1 <0.1 <0.1 0.3

Anthozoa 2.4 19.1 15.0 2.0 <0.1 4.9

Alcyonaria 0.1 0.3 0.1 0.1

Zoantharia 2.3 18.0 14.5 2.0 <0.1 4.6

Unidentified <0.1 0.8 0.4 <0.1 0.2

PLAlYHELMINTHES <0.1 <0.1 <0.1 <0.1

Turbellaria <0.1 <0.1 <0.1 <0.1

NEMERTEA 3.2 0.4 0.8 0.4 1.1 <0.1 0.4

ASCHELMINTHES <0.1 <0.1 <0.1 <0.1 <0.1

Nematoda <0.1 <0.1 <0.1 <0.1 <0.1

ANNELIDA 44.2 7.3 15.6 13.3 30.3 2.4 1.4 33.3 8.7

POGONOPHORA <0.1 <0.1 <0.1 <0.1 <0.1

SIPUNCULIDA 0.1 0.4 1.2 0.1 <0.1 0.4

ECHIURA <0.1 1.3 1.4 0.2

PRIAPULIDA <0.1 <0.1 <0.1 <0.1

MOLLUSCA 4.7 60.8 26.4 20.6 15.1 84.7 28.0 54.4

Polyplacophora 0.1 <0.1 <0.1 <0.1 0.1

Gastropoda 0.1 1.8 0.6 0.3 1.2 1.1 1.4

Bivalvia 4.0 58.7 25.6 19.8 13.6 83.6 28.0 52.7

Scaphopoda 0.6 0.1 0.2 0.5 0.3 <0.1 0.2

Cephalopoda <0.1 <0.1 <0.1

Uniden tified <0.1 <0.1

ARTHROPODA 1.8 6.1 6.8 1.3 1.2 1.9 1.2 5.2

Pycnogonida <0.1 <0.1 <0.1 <0.1

Arachnida <0.1 <0.1

Crustacea 1.8 6.1 6.8 1.3 1.2 1.9 1.2 33.3 5.2

Ostracoda <0.1 <0.1 <0.1

Cirripedia 3.0 4.3 1.5 0.1 2.7

Copepoda <0.1 <0.1 <0.1 <0.1

Cumacea 0.2 <0.1 <0.1 <0.1 <0.1 <0.1 <0.1

Tanaidacea <0.1 <0.1 <0.1

Isopoda 1.0 0.1 0.3 <0.1 0.2 <0.1 <0.1 <0.1

Amphipoda 0.5 2.4 1.8 0.6 0.2 <0.1 1.0 33.3 1.9

Mysidacea <0.1 <0.1 <0.1 <0.1 <0.1

Decapoda 0.5 0.3 0.6 0.8 0.4 0.5

Unidentified <0.1 <0.1 <0.1

BRYOZOA 1.4 0.9 <0.1 0.\ 1.4 <0.1 0.7

BRACHIOPODA 7.8 0.4 1.8 <0.1 0.5

ECHINODERMATA 36.6 16.2 24.4 45.4 48.8 10.9 69.4 20.7

Crinoidea <0.1 <0.1 <0.1

Holothuroidea 34.5 1.8 11.7 31.9 46.0 10.9 69.4 8.6

Echinoidea 1.7 12.8 5.7 1.4 0.7 9.4

Ophiuroidea 0.4 1.1 5.5 6.5 0.6 <0.1 1.8

Asteroidea 0.5 1.5 5.6 1.5 0.9

HEMICHORDATA <0.1 <0.1 <0.1 <0.1

CHORDATA 2.0 2.0 2.1 <0.1 1.7

Ascidiacea 2.0 2.0 2.1 <0.1 1.7

UNIDENTIFIED 0.2 0.3 0.3 0.1 <0.1 33.3 0.2

Total 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0
Composition and Distribution of Macrobenthic Invertebrate Fauna 69

Table 30

Frequency of occurrence (%) of each taxonomic group in the samples in each sediment organic carbon content class.

Sediment organic carbon content (percent)

Taxon 0.00 0.01-0.49 0.50-0.99 1.00-1.49 1.50-1.99 2.00-2.99 3.00-4.99 5.00+

PORIFERA 20 11 6 2 2

COELENTERATA 53 57 37 26 30

Hydrozoa 19 5 4 5 15

Anthozoa 35 52 33 21 15

Alcyonaria 5 14 8 15

Zoantharia 26 29 23 21 15

PLAlYHELMINTHES 2 8

Turbellaria 2 8

NEMERTEA 40 46 45 39 23 15

ASCHELMINTHES 10 8 17 7

Nematoda 10 8 17 7

ANNELIDA 80 98 99 100 98 100 50 100

POGONOPHORA 2 21 10 2

SIPUNCULIDA 20 31 28 8 2

ECHIURA <1 7 4

PRIAPULIDA <1

MOLLUSCA 100 90 96 98 91 100 75

Polyplacophora 10 12 8 2

Gastropoda 20 51 47 49 61 31

Bivalvia 80 85 94 99 88 100 75

Scaphopoda 20 17 31 32 28 15

Cephalopoda 1 1

ARTHROPODA 100 98 85 67 61 77 50 100

Pycnogonida 3 2 1

Arachnida <1

Crustacea 100 95 83 66 61 77 50 100

Ostracoda 1 1

Cirripedia 6 3 15 25

Copepoda 1 1 1

Cumacea 20 44 29 23 12 31

Tanaidacea 1 7

Isopoda 20 47 27 11 5 15 25

Amphipoda 60 93 72 55 49 62 50 100

Mysidacea 5 3 7 8

Decapoda 32 9 4 5 23

BRYOZOA 20 16 7 4 5 25

BRACHIOPODA 40 6 5 2

ECHINODERMATA 100 70 83 76 63 31 25

Crinoidea <1 I

Holothuroidea 40 17 37 27 16 15 25

Echinoidea 20 34 14 6 12

Ophiuroidea 60 39 70 66 47 23

Asteroidea 14 17 14 7

HEMICHORDATA

CHORDATA 23 19 13 8

Ascidiacea 23 19 13 8
70 NOAA Technical Report NMFS 140

70·

MAl N E

'9· 4"" 4'·

100

GRAMS

PER SQUARE METER

EXPLANATION "','

-, =7~:~ 4:·.: C@ '~, .50.0 - 391.8

G] I - 49

• 50 - 99

• 100 -138

NEW

JERSEY

EXPLANATION

1000 - -.. _ ...... _ .... ,

,

C2J ""

100

...

PORIFERA

Figure 27

Geographic distribution of Porifera: A-number of specimens per square meter of bottom; B-biomass in grams per square meter of bottom.

fauna was moderately high (3-5%) along the outer

continental shelf and upper continental slope (Table

14). On the inner shelf and in very deep water, sponges

made up a small portion of the total biomass.

Sponges were present in 3 to 12% of the collections

within each depth class (Table 15). The highest frequency of occurrence was on the continental slope

(200 to 2,000 m). These sponges were small, and were

frequently attached to polychaete tubes, shell fragments,

corals, and other biogenic substrates.

Relation to Sediments

Porifera were generally more numerous and constituted a substantially larger biomass in coarse substrates
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 71

,~ , \

I \

, \

I \

I \

I \

I \ I

/

I

d

25 50 100 200

WATER DEPTH IN METERS

0-- -0 NUMBER

__ WEIGHT

Figure 29

Density and biomass of Porifera in relation to water depth.

0

.....

..... 1Il0

zm 3

""

~lL

0 0

""cr

g,~

lL"" 0::;:

cr"" ""cr m<l

~::>

::>0

Zlll

cr

""

Q.

°0 10

Figure 28

Density and biomass of Porifera in each of the six geographic

areas.

16

o NUMBER

• WEIGHT - 14

::I ::I

- 12 0 0

... ... ... ",'" ~2 !I-- ::1 0

e lll ...

::I'" - 10 "' ... (30 "'0

!!!O'" "'",

...... 41- ... en ... - 8 ...... ...... z ... 0::1 ~::1

"' ... ...... 3- :-'" ... lIle'" -6 ... ::I=> ... => =>0 ... 0

z'" :-'"

'" 2

I- -4 '" ... ... ... ...

II-- -2

II. 11 n. rL 0 0 NOVA IULf' OF GEORGES SOUTHERN GEORGES SOUTHERtiI

SCOTIA MAINE BANI< NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE

GEOGRAPHIC AREA

(gravel, till, and shell) than in those composed

of fine particles (Tables 16-20; Fig. 30). Also,

the proportion of samples containing Porifera

was higher for the coarse than the fine substrates. Till contained the highest density of

specimens, the greatest weight, and highest frequency of occurrence. Gravel ranked second

in number and shell ranked second in weight.

Sand contained intermediate quantities, and

sand-silt and silt-clay con tained the smallest

quantities.

Relation to Water Temperature

Porifera were generally most numerous in areas where the annual range in bottom water

temperature was moderate-4° to 12°C (Tables

21-25; Fig. 31). Where the annual temperature

excursions were less than 4° or greater than

12°C the average biomass was markedly lower

than at midrange. The frequency ofoccurrence

was highest (15%) in those samples from localities in which the seasonal changes in temperature were between 4° and 8°C.

Two other aspects of the relative density of

Porifera and bottom water temperature that

were examined, but not tabulated or illustrated

here, concern annual maximum and annual

minimum temperature. Sponges were more

plentiful (2 to 7 g/m2 ) in areas where the maximum temperature was moderate, between 6°

and 12°C; they were scarce (<1 g/m2) where

the maximum temperature remained below 6°

or rose above 12°C. Porifera were abundant

(>4 g/m2) where the minimum temperature

was low (0° to 3.9°C) and scarce (<1 g/m2 )

where the minimum temperature was high.

Relation to Sediment Organic Carbon

Porifera were found only where sediments contained low to moderate (0.01 to 1.99%) amounts

of organic carbon (Tables 26-30; Fig. 32). Both

mean density and biomass diminished with increasing organic carbon content. Density

ranged from 3 to 0.2/m2 , and biomass from 5

to <0.01 g/m2.

Coelenterata

The macrobenthic coelenterate fauna of the New England region is composed of members of two classes:

Hydrozoa and Anthozoa. Hydrozoa are treated at the

class level, whereas Anthozoa, composed of members

from seven orders contained in two subclasses, are discussed at the subclass levels: Alcyonaria and Zoantharia.

No discussions are included at the taxonomic levels

Coelenterata or Anthozoa; however, the interested

reader will find summary data, for all treated parameters, for those taxonomic levels in Tables 5-30.

Hydrozoa-Hydrozoans were common in parts of the

New England region, but their limited occurrence and

moderately small size severely restricted their contribution to the total biomass. They made up less than 1% of
72 NOAA Technical Report NMFS 140

Figure 30

Density and biomass of Porifera in relation to bottom sediments.

Figure 31

Density and biomass of Porifera in relation to the annual range

of bottom water temperature.

oL--"O-l.3;-;.9,-----;;4..J,-1;;;.9--8••-!;-11.-;;-9--;;12~-1~5.09---;1"6:'!:'19:;:;.9;=:;;20~'~23".9'---.JO

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

4

o

:IE

o

... ., ... o :IE 0 «Ill

"'IL "0

8 a:

... ......

z ... ,,:IE

6 WL&J

JI",« ... ::>

4 g

... '"...

::E

o

... ., ... ::EO

«ID

4 '" IL "0

~a:... ......

z ... "::E

......

Jla:« ... ::> ... 0 ~.,

a:

...

...

SILT- CLAY

0---0 NUMBER

......... WEIGHT

SANO' SILT

SHELL SANO

BOTTOM SEDIMENTS

GRAVEL TILL

Bathymetric Distribution

Hydrozoans were most abundant in shallow water (0-24

m) where their density averaged 37 colonies/m2 and they

diminished in quantity more or less directly with increased

depth (Tables 11, 12; Fig. 35). In deeper waters (25-200

m) that cover the offshore continental shelf, their average

density was between 2 and 6 colonies/m2. They were

uncommon to rare on the continental slope and rise.

15% in the other shelf areas) to a low frequency (1 to

4%) in the Georges Slope and Southern New England

Slope area.

I

o NUMBER

l- • WEIGHT -I

I-- -I

- I--

-

f-

-

II--

rr -

I 11. o

ffi I

...

:IE

o

...

... lil 4

IL

o

'" 3

IL'" 0:IE

"' ... ... III '"« 2 :IE::> ::>0 Z.,

'"...

...

::E

o

...

... "0ZID

LI.. 4

0 0

.,...

~~ 3

IL'" olE

"' ... 2 ::E::> ::>0 Z.,

Geographic Distribution

Hydrozoans were common in coastal areas and

on offshore banks (Fig. 33). They were absent,

or present in only small quantities, in much of

the central sections of the Gulf of Maine, in

large areas of the continental shelf south of

Rhode Island, and on the continental slope

and rise. High densities, 100 to 500 colonies/

m2, and high biomass, 10 to 45 g/m2, occurr~d

in only a few scattered localities. Low densities,

to 49 colonies/m2, and average weights less

than 1 g/m2 were much more common and

widely distributed.

Three geographic areas contained moderate

to large quantities of hydrozoans: Nova Scotia,

G~orges Bank, and the Southern New England

Shelf (Tables 6, 8; Fig. 34). Average densities in

these areas ranged from 7 to 12 colonies/m2;

biomass averaged from 0.4 to 1.6 g/m2. Georges B2.nk

ranked first in terms of weight, and Nova Scotia ranked

first in number ofspecimens. Small quantities of hydrozoans were found in the Gulf of Maine, Georges Slope,

and Southern New England Slope. Quantities in these

three areas averaged between 3.3 and 0.1 colonies/m2,

and 0.12 and <0.1 g/m2.

The frequency of occurrence of hydrozoans in the

samples (Table 10) indicates the same general trend of

abundance as the average number and weight ofspecimens. Percentages of samples containing hydrozoans

ranged from a high of29% in the Nova Scotia area (8 to

the benthic biomass (Table 3). Members of

this group were most abundant in shallow waters and were attached to firm substrates, frequently where water currents were moderate to

strong.

Hydrozoans in our samples were small in size

and delicately tinted in white, pink, violet, tan,

and brown. They were exclusively carnivores

preying on planktonic crustaceans and other

small animals carried to them by water currents. In turn, hydrozoans are preyed upon by

nudibranchs and, probably, other omnivores

and carnivores.

The most common forms encountered were

Leptomedusae, represented by the genera

Campanularia, Sertularia, Obelia, and others. Less

common were representatives of Anthomedusae, of which Hydractinia is a member.

This hydrozoan occurred on live mollusks and

on dead shell. Some encrusted gastropod shells

were inhabited by hermit crabs.

Hydrozoans occurred in 126 samples (12%

of total); their average density was 6.4/m2 and

biomass averaged 0.5 g/m2 (Table 5).
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 73

r---------------------------,I.O

~-o ...." _ WllIHT

1.0

II: 1.0

1.1

1.0

0- - -~\

\

\

\

\

\

\

\

\

\ 1.0

0.1 1>- __-<>--_

k --0.-

oL...llLo~.o::7I--o.af:-....::.:::::;I;.O:===--"":'l.l:'lI---:I.O-?-,.......,'t..O;;--....lIt;,.o;--.--;;IO!..o.-----...JO

PEI'CENT OftGANIC CAMION

Figure 32

Density and biomass of Porifera in relation to sediment organic carbon.

Biomass trends were the same as those described

above for numerical density, with the exception of unusually large quantities between 50 and 100 meters

(Tables 13, 14; Fig. 35). In the shallow zone (0-24 m)

the average weight was 1.2 g/m2. In deeper water, at a

depth of 1,000 m and greater, the average quantity

decreased to <0.01 g/m2. The extraordinarily large quantity between 50 and 100 m resulted from relatively few

very large colonies.

The frequency of occurrence of hydroids in samples

was highest (21 %) in the shallow water zone (0-24 m),

dropping to 3% at depths below 1,000 m (Table 15).

Relation to Bottom Sediments

Hydrozoans occurred on all sediment types found in

the region; however, they were much more prevalent

on gravels than on other types ofsubstrates (Tables 16,

18; Fig. 36). Both numerical density and weight diminished with decreased sediment particle size. Average

density dropped from 20 colonies/m2 in gravel to 2

colonies/m2 in silt-clay. Wet weight declined from an

average of 2.6 g/m2 in gravel to 0.01 g/m2 in silt-clay.

Intermediate quantities were present on substrates composed of medium-sized particles.

Frequency ofoccurrence also reflected the general trend

of higher densities on coarse substrates and low densities

in fine-grained substrates (Table 20). In coarse sediments

the percentage of samples containing hydroids ranged

from 23 to 33%. The percentage occurrence in sand, an

intermediate grade, was 14%. In the fine-grained sediments the frequency of occurrence was only 4 to 6%.

Hydrozoans constituted only a small part (1.4%) of

the total benthic fauna, even in the sediment type

(gravel) in which they were most abundant (Table 19).

Relation to Water Temperature

The density of hydroids increased as the annual range in

bottom water temperature broadened. Average density
74 NOAA Technical Report NMFS 140

EXPLANATION

,.. GRAMS

PER SQUARE METER

••• 70·

M A IN E

M A IN E

.,.

72· II·

EXPLANATION

100

EEl 1 - 49

IIlIl!I 50 - 99

• 100-497

1000 - __ ~"'"

" 72° 'l,

"

EEl <0.1 - 0.9

iii 1.0- 9.9

• 10.0 - 45.3

1000 - __ ~"'"

" 72° 'l

,

"{:j~c....",,\..•""§j;)

7'·

HYDROZOA

Figure 33

Geographic distribution of Hydrozoa: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

was only 0.9 colonies/m2 where the temperature range

was small, reaching an average of 46 colonies/m2 where

the temperature range was broad (Table 21; Fig. 37), a

marked increase in density occurred between the 12-15.9°C

and 16-19.9°C classes: in the former the average density was

5 colonies/m2; in the latter it was 37 colonies/m2.

The contribution of hydroids to the total faunal density was <0.5% in the middle and low-range classes but

was above 1% in the two high-range classes (Table 22).

Hydroid biomass was moderate or low in all temperature range classes and no consistent trend in relation to

temperature range was evident (Tables 23, 24; Fig. 37).
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 75

Figure 34

Density and biomass of Hydrozoa in each of the six geographic areas.

°0!--....J.10,.--~-~----;"~----;;c~~~500=-~1~00~0~2!::'0~00~4~0';;:OO~0

WATER DEPTH IN METERS

Figure 35

Density and biomass of Hydrozoa in relation to water depth.

.6

e 2 "' ..... ~o

<llll

a:1L

00

za:

- .... .8 ~:;;

o~

........

,.cr<l ..... ::> .... 0

0.4 ,.'"cr

....

a..

::E

e

",5

::Em

1.5 IL

<:>0

z12:

-~

......... 1.0 :I:::E

.... .... 12:

~~

..... 0

0.5 '"

12:

....

0..

0--{) NUMBER

____ WEIGHT

GEORGES SOUTHERN GEORGES SOUTHERN BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELf' SLOPE

GEOGRAPHIC AREA

GUl OF SCOTIA MAINE

combination of light cream (rachis) and deep burgundy (pinnae).

Alcyonarians occurred in 63 samples (6% of total).

Their density averaged 0.8/m2 and biomass averaged

0.2 g/m2 (Table 5).

Geographic Distribution

Alcyonarians were present along the outer margin of

the continental shelf and on the continental slope and

rise. They were sparse in all sections and their occurrence was patchy, especially in the northern section

40r--------------------,

-I

o NUMBER

• WEIGHT

-I.

-

-0

f-

'- -

I II. NOVA F 0

8

o

::E

o

.....

'"ZIIl530 ....

::ElL -0

o

.... cr 0.. ....

"' 20 IL ....

0::E

cr .... .... a: met::>

~g 10 Z a:

....

0..

o

.....

..... "'0 ZIIl

\L 12

<:)0

~~"' ..... IL ....

o~

a: .... .... a: 1Il<l

~::>

::>0

Z",

Although hydroids did not occur in a

large proportion of the samples, there was

a general trend of increasing occurrence

rate from 4% in the low-range temperature

classes to 26 and 18% in the high-range

temperature classes (Table 25).

Relation to Sediment Organic Carbon

Hydrozoan abundance was generally negatively correlated to the quantity of organic

carbon in the sediments (Fig. 38). Density

of hydroids was greatest (9 colonies/m2)

where sediment organic carbon content

was low (<1 %), declining to moderate levels where carbon content was between 1

and 3% (Tables 26, 27). They were absent

in sediments containing the greatest

amounts of organic carbon (>3%).

Biomass of hydrozoans paralleled density,

diminishing with increasing organic carbon

content (Tables 28, 29). Biomass ranged from

slightly over 1 g/m2 to <0.01 g/m2.

Frequency of occurrence of hydroids in

samples ranged from 4 to 19% (Table 30).

The trend differed from density and biomass, however, in being parabolic with lowest occurrence in the middle ranges and

increasing at each extreme.

AnthozoaAlcyonaria-Alcyonarian coelenterates in

our samples were composed of soft corals,

orders Alcyonacea and Gorgonacea, and

sea pens, order Pennatulacea. Because of

the limited occurrence of both groups

«0.1% of the number of all organisms),

and despite their large size, they also constituted <0.1 % of the total benthic biomass

(Table 3). None of the alcyonarians in our

samples were taken from depths less than

50 m. They were most abundant between

200 and 500 m.

Soft corals are typically bush- or treelike

in shape and they attach to hard substrates,

usually rock outcrops or gravel. Soft corals range in

height from a few millimeters to several meters. Trunk

diameters are proportional in size, and in large specimens occasionally exceed 10 cm. Thus, some species of

this group may rank as the largest sessile invertebrates

in this region. Colors are light tan, pink, or various dark

shades of red.

Pennatulacea are feather-shaped animals commonly

10 to 25 cm in length. They characteristically dwell in

soft bottom sediments anchored by a peduncle. Color

ofthe majority ofspecimens in our samples was tan or a
76 NOAA Technical Report NMFS 140 _

Figure 36

Density and biomass of Hydrozoa in relation to bottom sediments.

o

2.8

0.4

2.4

20

2.0 g C/)~

20 0

"al

"'lL

"'0

.6 z 0:::

-",

~~

:1:'"

~2O

.2

~~" "'0

~C/)

0.8 ffi

Q.

BOTTOM SEDIMENTS

GRAVEL TILL SHELL SAND SAND- SILTSILT CLAY

o NUMBER

• WEIGHT

-

-

01- -I

61- -,

2"-

-

BI-

-

4-

o I .. I Il n

(Fig. 39). They were absent in the samples

from Georges Bank. Their average density in

all localities was between 1 and 151m2 . Their

average biomass at all localities was 18.9 g/m2

or less.

Alcyonarians were present in approximately

the same quantities in all standard geographic

areas, except Georges Bank where they were

absent. Average densities in geographic areas

where they were present ranged from 0.7 to

1.5/m2 , and average biomasses ranged from

0.03 to 0.43 glm2 (Tables 6-9; Fig. 40).

In addition to number and weight per unit

area, another reasonably good index of abundance is the percentage of samples containing alcyonarians (Table 10). All shelf areas

(Nova Scotia, Gulf of Maine, Georges Bank,

and Southern New England Shelf) had a frequency of occurrence of 7% or less. The occurrence rates on Georges Slope and Southern New England Slope were 27% and 17%,

respectively. Alcyonarians formed a larger

share of the total benthic fauna in the latter

two (slope) areas than in the shelf areas.

Bathymetric Distribution

Alcyonarians were most common in deep water habitats in the New England region. They

were not present in any samples taken at

depths less than 50 m but were represented in

all depths greater than 50 m (Tables 11-14;

Fig. 41). Although differences in average density and weight from one depth class to another were small, the larger quantities were

most prevalent in deep water (100 to 2,000

m). Highest average density and largest average weight, 1.9/m2 and 0.47 g/m2, respectively, occurred at depths from 200 to 500 m.

The frequency of occurrence of alcyonarians in our shallow water «50 m) samples was

zero (Table 15). The occurrence was 5% at

moderate depths (50 to 200 m) and increased

to moderately high levels (22 to 29%) in deep

water areas.

0--0 NUMBER

20 50 - WEIGHT ::E 0 2.5 0

"'0 /' "'~ ZIIl /' ::EO

'" 40 /' "Ill ::ElL if 2.0 a::lL GO <:>0

"'", / za:: ~~ 30 / -",

lL'" 1.5 ~~

020 ! :1:'" <:>::E

"'''' ILl ILl "'''' 20 Ill" 1.0 ~a::

::E~ CI

~o ~:::>

z'" 10 ~~

'" 0.5 a:: '" ILl Q. D..

0 4-7.9 8-11.9 12-15.9 16,19.9 20-23.9 0

RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 37

Density and biomass of Hydrozoa in relation to the annual range

of bottom water temperature.

Relation to Bottom Sediments

Alcyonarians occurred only in selected types of substrate (Tables 16-19; Fig. 42). The Alcyonacea and

Gorgonacea were collected from gravel and rocky outcrops, in contrast to the Pennatulacea, which inhabited

sand-silt and silt-clay sediments. No alcyonarians were

taken on substrates oftill orshell, and they were presen t

in only 5 samples out of a total of 455 from sand

bottoms. Frequency of occurrence of this faunal group

was highest (10 to 12%) in the silt-elay and sand-silt sediments and substantially lower (4%) in gravel (Table 20).

Relation to Water Temperature

Alcyonarians occurred only in areas where the annual

temperature range was moderate or small, and less

than 16°C (Tables 21-24; Fig. 43). The trends in both

quantitative measures and also in frequency of occurrence exhibited a bimodal relationship.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 77

I 0--0 NUMBER r

o

_WEIGHT

1 10.0 1.0

....... -1

r' \

I \ :lE :lE 8.0 \ 0.8 0 0 I

\ fi)~

~i I :lEO

\ "'lID 1&1 I 0: ... :lE'" \ Co

u O I ~o: \ !:o:

I 1&1 fi)~ 8.0 \ 0.8 l;j ... 1&1 I \ 0:lE ~:lE

0:1&1 I \ 1&11&1

1&10: _0:

me I \ "'I

:lE'j ~'j

1&10 'jO I \ _fi) Zfi)

ffi 4.0 \ 0:

I 0.4 1&1

Q. \ Q.

I

I \

I \

1 \ f\ 2.0 \ / \ 0.2

\ / \

/ \ \ / \ \ / \ li \

0 0 0.01 0.5 1.0 1.5 2.0 3.0 5.0 10.0

PERCENT ORGANIC CARBON

Figure 38

Density and biomass of Hydrozoa in relation to sediment organic carbon.

Alcyonarians were relatively more abundant in both

the narrow (less than 4°C) temperature range class and

where temperatures were moderately broad (12° to

15.9°C) than in areas ofintermediate range. The densities averaged between 1.1 and 1.7/m2 in the narrow

and relatively broad range classes and between 0.2 and

1.2/m2 in the intermediate range classes. Biomass averaged 0.31 to 0.42 g/m2 in the narrow and relatively

broad range classes and only 0.11 g/m2 or less in the

intermediate classes.

Frequency of occurrence of alcyonarians in samples

ranged from 11 to 8% where the temperature was narrow

and relatively broad, and only 2 to 5% in areas where the

temperature range was intermediate (Table 25).

Relation to Sediment Organic Carbon

Alcyonarians occurred only in sediments with low (from

0.01 to 1.5%) organic carbon content; they were absent

in areas with no measurable organic carbon, as well as

in areas with carbon contents exceeding 1.5% (Fig. 44).

The 0.50-0.99% carbon content class contained the

highest average density (Tables 26, 27) as well as highest

average biomass (Tables 28, 29). The only other carbon

content classes in which alcyonarians occurred, 0.01-0.49%

and 1.00-1.49%, had reduced measures of abundance.

Frequency of occurrence in samples showed a trend

similar to density and biomass (Table 30), highest in

the 0.50-0.99% carbon content class and significantly

lower in the two adjacent classes.

Zoantharia-Four morphologically diverse orders of

Zoantharia were represented in our samples from the

New England region: 1) Actiniaria-sea anemones; 2)

Ceriantharia-burrowing anemones; 3) Madreporariastony corals; and 4) Zoanthidea-colonial anemones.

Altogether they contributed 1.5% of the number of

animals and 3.5% ofthe biomass in the total macrofauna

(Table 3).

Our samples contained specimens ranging in size from

small (1 cm or slightly less) Edwardsia and other burrowing anemones to large Cerianthus over 30 cm in length.
78 NOAA Technical Report NMFS 140

8g0

42'

70°

M A IN E

4"

8

400

72°

.r

.,

"

ie""

~l

,

it".""

EXPLANATION

EXPLANAT ION

NEW

JERSEY

IillillJ <0.1 - 0.9

III 1.0 -18.9

[d 1- 9

II 10-15

GRAMS

PER SQUARE METER

72°

72°

ALCYONARIA

Figure 39

Geographic distribution of Alcyonaria: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

These organisms displayed many vanattons of color

pattern and hue. Some small species and many larger forms

had tentacles that were transparent or whitish. Body colors

were mostly tan, flesh colored, or reddish-brown; a few

were orange, red, or another somewhat similar bright color.

Stony corals of both solitary and colonial types occurred but neither type was common.

Commensalism was common among the Actiniaria

and Zoanthidea, particularly in association with hermit

crabs and decapod shrimps.

Occupancy of diverse habitats characterized the New

England zoantharians. They occurred in a wide variety

ofsubstrates, and were distributed over broad ranges of

water depth and temperature.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 79

o .4 .... en .... ~o

.. CD

0:: ...

"0

.3 z 0::

-w

........

J:W

~~

.2

.. .... ::J wo

~en

.1 f5

Q.

.5

GEORGES SOUTHERN GEO GE

BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

NOVA GULF Of

SCOTIA MAINE

0

0 NUMBER

-WEIGHT

-0

-0

51-

-0

01-

-0

51-

0 .. I 0 R S SOUTHERN

.... eno

ZCD

W

... <:>0

Wo::

g;~

~"'" I. o~

o::w

wo::

~~ I. ::Jo Zen

0::

o.

Figure 40

Density and biomass of Alcyonaria in each of the six geographic

areas.

,....--------------------,0.6

Figure 41

Density and biomass ofAlcyonaria in relation to water depth.

o

....

.... "'0

~<D

..

0::'" 0.4 C> 0

zO::

-UJ.... .... w ~~

-UJ

UJo:: .2 ..

.... => UJO

~lIJ

0::

UJ

lL

0---0 NUMBER

_ WEIGHT

25 50

WATER DEPTH IN METERS

Bathymetric Distribution

Zoantharians occurred at depths from 6 to 2,495 m but

were most abundant between 25 and 100 m, where

their density averaged from 21 to 911m2 (Tables 11, 12;

Fig. 47). At other depths their density averaged 0.2 to

6.8/m2. Only a moderate number (5.6/m2) were present

at depths from the shoreline to 24 m. Their density in

deeper water progressively diminished from 6.8/m2 in

outer shelf depths to 0.2/m2 on the continental rise.

In terms of biomass, the relationship between quantity and depth of water was considerably different from

that described above for numbers of specimens. At

depths of 0 to 100 m the average biomass was moderate

Geographic Distribution

Zoantharia were widely distributed over most of

the marine waters off the northeastern United

States (Fig. 45). They were especially common in

coastal areas and on the offshore banks. Low and

moderately low quantities (1 to 49 individuals,

and less than 10 g/m2 biomass) were most prevalent. Intermediate and high quantities (50 to 572

individuals, and 10 to 1,561 g/m2 biomass) occurred in scattered patches.

Average numerical density ranged between 3

and 8.2 specimens/m2 for all standard geographic

areas except Georges Bank where the density averaged 92.5/m2 (Tables 6, 8; Fig. 46). This unusually high abundance on the bank was due mainly

to the presence oflarge quantities ofsmall Ceriantharia

and the moderately common Zoanthidea.

Biomass ofspecimens reflected entirely different patterns of abundance. Average biomass of Zoantharia

within geographic areas was greatest (19.5 g/m2) in

Nova Scotia and diminished rather uniformly to the southwest (Tables 8, 9; Fig. 46), terminating with a low quantity

of 0.7 g/m2 in the southern New England Slope area.

Frequency of occurrence was moderate and fairly

consistent among the various areas. The percentage of

samples containing Zoantharia ranged between 17 and

31 %, with the lowest incidence rates in the two slope

areas (Table 10).

Zoantharians occurred in 265 samples (25% of

the total). Their density averaged 22.6/m2, and

biomass averaged 6.4 g/m2 (Table 5).
80 NOAA Technical Report NMFS 140 _

o NUMBER

_ WEIGHT

- 2f-

- IfI I 0 GRAVEL TILL SHELL SAND SAND- SILTSILT CLAY

o

fIlO

Zal

.... ... (30

.... '" ~~.......

o~

"' .... ale:( .... '" ~::>

::>0

ZfIl

'"....

n.

BOTTOM SEDIMENTS

Figure 42

Density and biomass of Alcyonaria in relation to bottom sediments.

o

fIl~

~O

e:(al

IX ... <:>0

0.6 z '" - .... ~~

:1: ....

~~

04

e:( ~::>

0.2 g

'"....

n.

0--0 NUMBER

_ WEIGHT >- fIlO

Zal

.... ... (30

~~fIl>-

.......

o 2

"' .... .... ale:('" ~::>

::>0 I ZfIl

'"....

n. OL----=o~_3;-:.9:-----:-~----,--h-~L....,--~~~.,.....L.,-----.J 0

ANNUAL

(1.2 to 2.1 g/m2 ). Relatively high quantities

(8 to 18 g/m2) occurred in outer shelf and

upper slope depths (Table 13; Fig. 47). The

proportion of the total benthos contributed

by Zoantharia was low at all depths except

between 100 and 500 m where members of

this group made up 13 to 19% of the total

biomass (Table 14).

Frequency of occurrence of Zoantharia in

samples was moderate (16t033%) at depths

down to 500 m. At depths greater than 500

m they occurred in only 8 to 18% of the

samples (Table 15).

(Tables 21, 22; Fig. 49). Density was highest (981m2 ) in

areas with a moderate (l2°-15.9°C) temperature range,

and densities decreased to about 2 to 41m2 where the temperature range was both narrow «4°C) and broad (>20°C).

The relation between zoantharian biomass and temperature range differed from that of density. Generally

biomass was high (4 to 8 g/m2 ) where the temperature

range was narrow or moderately narrow (<12°C), and

the general trend was a decrease in biomass associated

with a broadening of the temperature range (Table

23). This correlation is revealed more clearly by the

percentage of the total benthic fauna that is made up

by lOantharians (Table 24). Where the tern perature

range was less than 4°C, lOantharians contributed 18%

of the total benthos. Their contribution decreased as

the temperature range broadened, forming only 0.2%

of the benthos where the range was >20°C.

Figure 43

Density and biomass of AJcyonaria in relation to the annual range

of bottom water temperature.

Relation to Sediments

Zoantharians were present In all types of

bottom sediments sampled (Fig. 48). Highest density (431m2 ) occurred in sand sediments, moderate densities (9 to 141m2 ) in

gravel and sand-silt, and low densities (2 to 31

m2) in all other types (Tables 16, 17).

In terms of biomass the relationship was somewhat

different. The largest quantities occurred in sand-silt, which

was due mainly to the presence oflarge burrowing anemones. Rather large quantities, mostly actinarians, were

present in gravel. Relatively small biomasses occurred in

all other types of bottom sediments (Tables 18, 19).

The frequency of occurrence of this diverse group of

animals was moderate and rather uniform in all types of

bottom sediments. The percentage ofsamples con taining Zoantharia in each of the different bottom types

ranged from 17 to 35 (Table 20).

Relation to Water Temperature

The average density of lOantharians in relation to the

annual range of bottom water temperature occurring in

different regions of the study area varied from 2 to 981m2
Composition and Distribution of Macrobenthic Invertebrate Fauna 81

2.0

0---0 NUMBER

_ WEIGHT 0.4

2 0.3 2 0 1.8 0

(I)~ i 2 0

e lll

... a:: ... 0

/p.. "'0

a:: za::

'" /

" -",

... ", 1.0 ~~

/ \ 0.2 02 :.:'" ",2

a::", / \ "'''' lila:: • a::

2 e / \ e ~~ / \ ~~

zg ~~ a:: f \ a::

'" 0.8 / 0.1 '" A. 'h A.

/

/

"

\

/ \

/ '\

0 0 0.8 1.0 1.8 2.0 3.0 8.0 10.0

PERCENT ORGANIC CARBON

Figure 44

Density and biomass of Alcyonaria in relation to sediment organic carbon.

Frequency of occurrence was moderate under all

temperature range conditions. Occurrence rates varied

from 14 to 30% (Table 25). They were higher where

the temperature range was moderate, and lower in the

extreme (lowest and highest) range conditions.

Relation to Sediment Organic Carbon

Zoantharians occurred where sediment organic carbon

content was from 0.01 to 3%; they were absent where

carbon content was 0 or above 3% (Fig. 50). Density

values showed a negative trend of decreasing quantity

with increasing carbon content (Tables 26, 27; Fig. 50)

with mean number ofindividuals ranging from 3 to 71m2.

Biomass values exhibited a somewhat similar trend

with the exception that highest biomass was not in the

lowest carbon content class in which they occurred

(0.01-0.49%). Moderately high biomass occurred in

the two classes between 0.5 and 1.5% with significantly

lower levels above and below these values.

Frequency of occurrence of samples in the carbon

content classes was fairly uniform, ranging from 15 to

29%, with no discernible trend as evidenced by density

and biomass measures (Table 30).

Platyhelminthes

Turbellaria-Tubellarians are free-living members of

the phylum Platyhelminthes. They accounted for a very

small portion of the total New England benthic

macrofauna. In terms of biomass and numbers of individuals they accounted for <0.1 % of the total fauna

(Table 3). They are small in size and those large enough

to be retained on a I-mm mesh sieving screen were present

in very low density. The vast majority ofmarine tubellarians

reported from New England marine waters are less than a

few millimeters in length. Specimens in oursamples ranged

in size from 2 mm to nearly 2 cm in length.

Members of this class of flatworms are free-living,

soft-bodied forms and their shape varies from species to

species, commonly ovoid and dorsoventrally flattened.

Turbellaria occurred in 16 samples (2% of total).

Their density averaged 0.4/m2 , and their biomass averaged 0.01 g/m2 (Table 5).

Geographic Distribution

The few Turbellaria that were present in our samples

were relatively more common in the Southern New

England Shelf area than in any other section (Fig. 51).

Average densities as high as 59 individuals/m2 were

detected in the vicinity of Nantucket Shoals. Elsewhere

densities averaged 91m2 or less. Members of this group

were absent from large portions of the Nova Scotia

Shelf, Gulf of Maine, Georges Bank, and the entire

Southern New England Slope area. Average biomasses

in all localities were 0.3 g/m2 or less.

The density of turbellarians in each geographic area

(Tables 6, 7; Fig. 52) averaged less than 11m2. The
82 NOAA Technical Report NMFS 140 _

EXPLANATION

ill] 1-49

• 50-99

• 100- 572

EXPLANATION

Q

W1 <01- . . 99

IIIB 10.0 - 49.9

• 50.0-1561.3

GRAMS

PER SQUARE METER

39·

ZOANTHARIA

4'·

10·

M A IN E

••• 0'---~___"120 KM

"- "- "-

"

42·

Figure 45

Geographic distribution of Zoantharia: A-number of specimens per square meter of

boltom; B-biomass in grams per square meter of boltom.

southern New England Shelf area contained the highest density (0.9/m2); Georges Slope ranked second with

0.4/m2; and all other areas had 0.2 individual or less

per square meter.

The average biomass of this group of animals in each

of the six geographic areas was nearly the same (Tables

8,9; Fig. 52). The biomasses averaged 0.01 g/m2 or less.

Frequency of occurrence also yielded a low index of

abundance. Percentage of samples containing turbellarians ranged from 0 to 3% (Table 10).

Bathymetric Distribution

Turbellarians were most plentiful (2.6 specimens/m2 )

in shallow water, uncommon at depths from 25 to 500
Composition and Distribution of Macrobenthic Invertebrate Fauna 83

100

0 NUMBER

-WEIGHT

80 0 0

... ... ... en'" 0 ~o

<D ~<D

.... a: .... 0 60 "'0

a: 20 ~a:

W W ... ...... .... w J:W o~ 15 "'~

a:w Ww wa: 40 ~a:

<D~

~::> ... ::>

::>0 wo zen 10 ~en

a: a: w w Cl. 20 Cl.

GEOGRAPH Ie AREA

Figure 46

Density and biomass of Zoantharia in each of the six geographic

areas.

100

r\ ::li

o B I \ ... I \ ~b1LI 11l I \ ::Iiu.. I \ -0 60 u

1LIa: I \ 0-1LI

Ulfu.. 1LI I \ O::li 40 / \ a: 1LI 1LIa:

III <I I ::Ii:::> \ :::>0

ZUl

20 I a:

1LI / 0-

0

0 10 25 50 100 200

c>- -0 NUMBER

_ WEIGHT

1000 2000 4000

WATER DEPTH IN METERS

Figure 47

Density and biomass of Zoantharia in relation to water depth.

meters, and not found in depths greater than 500 meters

(Tables 11,12; Fig. 53).

Average biomass revealed a similar trend (Tables 13,

14; Fig. 53): 0.02 g/m2 in the shallow zone and 0.01 gl

m2 or less in deeper water.

In the depth zones where they were present their

incidence ranged from 1 to 4% (Table 15).

Relation to Sediments

Turbellarians were present in only three of the SIX

major sediment types occurring in the study area. They

were most common in gravel, where they averaged 1.7

individuals/m2. In sand and silt-elay their average densitywas 0.2 and 0.l/m2 (Tables 16,17; Fig. 54).

In terms of biomass they were equally sparse (averaging 0.01 g/m2) in all three sediment types (Tables 18,

19; Fig. 54).

Turbellarians were found in only 1 to 2% of the

samples from the three sediment types in which they

occurred (Table 20).
84 NOAA Technical Report NMFS 140

50

0

>- 4 - >- "'0 ZCD

UJ

~"-

GO 30 ~Q:; ",UJ>- ,,-UJ

O~

Q:;UJ 20- UJQ:;

CD",

~=>

=>0

z",

Q:; 10 UJ

a.

0 GRAVEL

o NUMBER

• WEIGHT

20 0

>- ",>-

~O

",CD

Q:;,,-

15 "'0

~Q:;UJ >->-

:I:UJ

"'~ 10 UJUJ

~Q:;

'" >-=>

UJO

~'"

'"

UJ

a.

0

BOTTOM SEDIMENTS

Figure 48

Density and biomass of Zoantharia in relation to bottom sediments.

100

A 0--0 NUMBER

-.. WEIGHT

::E I \ ::E 0

I 0 f- BO \ f- f- ",f- "'0 I ZCD ::EO

LU \ et lD

::E"- I "',,- 0 0 "'0

LU", 60 I \ a. 12 '" LU "'f- LU

I f-f- ,,-LU \ :I:LU 0::E ~::E

Q:;LU I LULU

LU'" \ lDet 40 ~'" I B et ::E=> f-=> =>0 \ LUo Z'" ~'"

'"

LU '" \ LU 0. 0.

20

/'

/'

.-if

0 0-3.9 4-7.9 8-11.9 12-15.9 16,19.9 20-23.9 0

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 49

Density and biomass of Zoantharia In relation to the annual

range of bottom water temperature.

Relation to Water Temperature

Turbellarians occurred in all temperature range classes

except one (Tables 21-25; Fig. 55). They were absent

from samples representing areas where the temperature change was less than 4°C. Their density, biomass,

and frequency of occurrence in all temperature range

classes in which they occurred were quantitatively low

and roughly equal from class to class.

Relation to Ssediment Organic Carbon

Turbellarians occurred in three organic carbon content classes, two in the low range and one in the moderately high range (Fig. 56). Densities ranged from 0.1 to

0.9 individual/m2 (Tables 26, 27), and biomasses ranged

from <0.01 to 0.19 g/m2 (Tables 28, 29). Frequency of

occurrence in samples ranged from 1 to 8% (Table 30).

In all measures of abundance highest values occurred
Composition and Distribution of Macrobenthic Invertebrate Fauna 85

.- -,20.0

0--<> NUMBER

_ WEIGHT

4.0

16.0

-~

\

\

\

\

\

\

\

\

\

\

90- I ......

I

I

I

I

I

I

I

I

I

I

I

I

I

I

I

2.0

8.0

:I

I-

"'0 zm

IL 6.0

<3 0

lila:

5;~

IL lIl 0:1

a:lIl lila:

m c( 4.0 :I:::>

:::>0 z",

a:

O'----~~::----___:f=__-_;_';;__-__;_'o--*--"L.-*--tf___;~~____;~-----'O o.~ 1.0 I.~ 2.0 3.0 ~.O 10.0

PERCENT ORGANIC CARBON

Figure 50

Density and biomass of Zoantharia in relation to sediment organic carbon.

in the moderately high (between the 2 and 3%) content class.

Nemertea

Nemertines, although widely distributed throughout the

study area, made up a rather small percentage, only 0.5%

of density and 0.4% of biomass, of the total benthic fauna

due to their low abundance and the small size ofthe m~ority

of specimens obtained (Table 3). Their greatest numerical density and biomass occurred on the continental shelf

and along the upper portions of the continental slope.

Members of this group are carnivores which characteristically burrow freely in the substrate.

Specimens in our samples ranged in size from about

1 to more than 25 cm in length; however, some of the

larger ones, although rare, were not whole, representing only part of an obviously larger animal. These softbodied, vermiform organisms easily break during the

collecting process and frequently fragment when placed

in formalin for preservation.

A large proportion of all specimens were uniformly

tan or flesh colored. A few individuals had brownish or

tan bodies with distinctive bands or stripes of white,

yellow, or orange.

Although nemertines were usually represented by

one or a few specimens per sample, some samples contained over 100 individuals. They occurred in 405

samples (38% of total). Their density averaged 8.2/m2,

and their biomass averaged 0.71 g/m2 (Table 5).

Geographic Distribution

Nemertines occurred over nearly the entire study area

(Fig. 57). Their numerical abundance was moderately

low, averaging between 1 and 9 individuals/m2 over

most of their range. An extensive area of moderate

density (10 to 491m2 ) extended along southern Georges

Bank, across Great South Channel, and westward to the

vicinity ofRhode Island. They were absent in a few deep

water sections of the Gulf of Maine and on the continental rise southeast of Long Island, New York. Density

(average number of specimens) was greatest (23 individuals/m2) on Georges Bank (Tables 6, 7; Fig. 58). In

all other areas density averaged between 1.2 and 6.8

individuals/m2.

Over the six standard geographic areas there was a

slight increase in biomass of Nemertea in the shelf

areas from northeast (about 0.6 g/m2 ) to southwest,

with the southern New England Shelf having the largest

biomass (1 glm2). Both slope areas had very small
86 NOAA Technical Report NMFS 140

59·

•

70·

M A IN E

.,.

------~----_/~--------- .... ------.... - ...... 45·

.... ,

,

'00

EXPLANATION

OJ <0.1-0.3

EXPLANATION

OJ 1-9

III 10-59

NEW

JERSEY

GRAMS

PER SQUARE METER

...

...

TURBELLARIA

Figure 51

Geographic distribution of Turbellaria: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

quantities, averaging 0.1 and 0.2 g/m2 (Tables 8, 9;

Fig. 58).

Frequency of occurrence of nemertines was moderate in all geographic areas. They were encountered

most frequently (52% of the samples) in the Southern

New England Shelf. In all other areas they occurred in

25 to 35% of the samples (Table 10).

Bathymetric Distribution

Nemertea were found in water depths ranging from 7

to 3,820 m. Greatest density (27 individuals/m2) was at

depths between 25 and 49 m (Tables 11, 12; Fig. 59).

Substantially lower densities (0.5 to 7.8 individuals/m2)

were found at all other depths, with density generally

decreasing with increased depth. Lowest densities (0.5
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 87

Figure 52

Density and biomass of Turbellaria in each of the six geographic

areas.

to 1.5/m2) occurred at depths greater than

500 m.

The relation between average biomass and

water depth was very similar to that for numerical density, but the range in values was

much more limited. Largest average biomass was 1 g/m2 in the 25-49 m depth class.

At depths greater than 500 m the biomass

was very small, averaging 0.12 g/m2 or less

(Tables 13, 14; Fig. 59).

Frequency of occurrence of nemertines

in samples from the eight depth classes

ranged from 18 to 51 % (Table 15). Higher

rates of occurrence were most prevalent in

the shallow-water classes, and low occurrence was typical of deepwater classes.

:IE

0

>-

>- fl)0

ZlD

:lElL '"<:;0 1.2 "'a:

~~

lL'" O:IE O.B

a:'" ",a:

lD<t

:IE:::l

:::l0 0.4 ZfI)

a:

... '" 0

GEOGRAPHIC AREA

o NUMBER

• WEIGHT

:IE

o

>-

~~<till

a:lL

"'0

za:

-",

>->-

:1:'" ",:IE

0.02 ;

<t >-:::l

001 g

a:

'"

...

WATER DEPTH IN METERS

Figure 53

Density and biomass of Turbellaria in relation to water depth.

0- --0 NUMBER 2:

___ WEIGHT

~b<till

a: lL 0 0

zffi

-Io03i~ OUJ

i£ia:

002 ~~

>-0

UJUl

0.01 a:

'"

Q.

'\

\

\

\

\

\

\

\

Relation to Sediments

Although nemertines were present In all

types of sediments, they were common (13

to 28 individuals/m2) in only two types: shell

and sand (Tables 16, 17; Fig. 60). Densities

of about 3 to 5 individuals/m2 occurred in

gravel, sand-silt, and silt-clay. Lower density

(0.9/m2) was found in till. Biomass of nemertines in shell bottoms was moderately large

in absolute terms (6 g/m2 ), but in relative

terms it was exceptionally large. In all other

sediments types the average biomass was

0.83 g/m2 or less. An unusually small quantity (0.06 g/m2) was present in till substrates

(Tables 18, 19; Fig. 60).

The percentage of samples within each

sediment type in which nemertines occurred

was in close agreement with the quantity

present. Samples from shell bottoms had

the highest incidence of nemertines (50%),

till the lowest (14%); their incidence in

other sediment types was intermediate (28

to 47%) (Table 20).

Figure 54

Density and biomass of Turbellaria in relation to bottom sediments.

BOTTOM SEDIMENTS

0.02

:IE

o

>-

~b<till

a:lL

"'0

~a:

>->-'" :1:'"

"':IE

;~<t >-:::l

",0 0.01 fI)

a:

'"...

o NUMBER

_ WEIGHT

'--~G~R;-;;:AV~E~L-~T:;:IL"7L--S;:-:H"'*E;-;LL-~~---;:-:'~-+~_---..JO

:IE

0

>-

>- fl)0

ZIIl

'"

2lL

0° 2.0

"'a:

... ", fI)>- 1.5 lL'" 02

a:'"

wa: 1.0 lD<t

2:::l

:::l0 ZfI) 0.5

a:

... '" 0

Relation to Water Temperature

The average density of nemertines ranged

from 2.4 to 25.3 individuals/m2 throughout

the entire temperature range of the study

area (Tables 21, 22; Fig. 61). The highest

density occurred in areas with an intermediate annual temperature range of 12° to

15.9°G In areas where the range was either

greater or less than this, the densities were

substantially lower. Where the range was

greater (16° to 23.9°C) density values decreased to 2.9/m2. Where the range narrowed from 11.9° to ooe, the values decreased from 9.0 to 2.4/m2.
88 NOAA Technical Report NMFS 140

::E ::E 0

.... 0---<> NUMBER 0

.... 0.12 I-

"'0 ----. WEIGHT ",IZeD ::EO

'" et eD

::E'" 0:: ...

0° e>o "'0::

Q.", 0.08 ~o:: "' .... 1.0 /\ '" ... '" ........ 0::E :I:'"

/ \ e>::E 0::'" "'''' "'0:: eDet 0.5 / \ 0.04 ~o::

::E:> et :>0 / \ .... :> z", ",0

0:: ~'"

'" 0:: Q.

0 0 '"

Q.

0- .9 4-7.9 8-11.9 12-15.9 16,19.9 20-23.9

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 55

Density and biomass ofTurbellaria in relation to the annual range of

bottom water temperature.

1.0

0--<> NUMBER

_ WEIGHT

0.8 / \ 0.20

I

2 / 2 0

l- I 0

l- Ieno / en I- ZeD

R 2 0

.... 0.6 0.)5 <teD

/ a:1L

u O I \ "0

l&Ja:

Cl.l&J / ~a: I en I- \ l&J

.... 1&1 I \ / 1-1-

J:;l&J 02

I ,,::I

a:l&J \ / l&Jl&J l&Ja: I / 1~0.4 0.10:- \ 1-:::> ::l0 I / l&J0 zen \ :-en a: I \ / a: l&J l&J Cl. I I Cl.

\

0.2 I \ / O.OS

I \ /

I \ /

I \ I

I L I

° 0.01 0.5 - 5.0 10.0 °

Figure 56

Density and biomass of Turbellaria in relation to sediment organic carbon.
___________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 89

72'

72'

EXPLANATION

Lill <0.1

IIllllI 0.1-0.9

• 1.0-35.8

GRAMS

PER SQUARE METER

39'

12'

NEMERTEA

7" 70'

'2'

.9'

Figure 57

Geographic distribution of Nemertea: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

The relation of nemertine biomass to temperature

range differed from that of density, exhibiting a general increase in biomass (from 0.22 to 1.38 g/m2) as the

temperature range broadened (Tables 23, 24; Fig. 61).

Largest average biomass occurred where the range was

16° to 19.9°C, and smallest where it was less than 4°C.

The frequency of occurrence of nemertines in relation to temperature range was similar to their density

distribution. The highest percentage (52 to 49%) of

samples containing specimens occurred in areas with

intermediate temperature ranges. Where the temperature range was narrow or broad, the frequency of occurrence was relatively low (Table 25).

Relation to Sediment Organic Carbon

The average density of nemertines exhibited a rather
90 NOAA Technical Report NMFS 140 _

o

o

....

~b<I%>

Q:o..

<:>0

.2 Z Q: -w

.... .... :t: W <:>~

Ww

O.B :t

.... ::> wO:tV>

Q:

W

ll- 0.4

NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPH Ie AREA

5

0 NUMBER

• WEIGHT

of5f- -I

f- -

f- -

o I I r-. II

2 o

....

.... V>O

ZI%>

W

~o..

U 0 I

~~V> ....

o..w

o~

ffi 10 1%><

~::>

::>0

Z<n

Q:

W

ll- 5

Density and biomass of

areas.

Figure 58

emertea in each of the six geographic

30

~--o NUMBER

0 ....- WEIGHT 1\ 25 1\ 0 / \ 0 f- f- (/)6 .... I \ rno ~m 20 I ~m

... \ <o..

0 0 I \ ~o wa:: I 1.5 za:: ll-w \ _w <Il .... 15 I \ .... ... w f-W

o~ :t:~

a:: w

I sew wa:: 1.0 wa:: met 10 I ~<l ~::J ::J ::JO .... 0 z(/) I ~<Il a::

w I , a:: Q. 5 6 0.5 w

Q.

WATER DEPTH IN METERS

Figure 59

Density and biomass of Nemertea in relation to water depth.

sharp negative correlation to organic carbon content

(Tables 26,27; Fig. 62). Density was highest (111m2 ) in

areas with extremely little or no organic carbon in the

sediment and declined steadily (from 11 to 21m2) as

organic carbon content increased. None were found

where carbon content exceeded 3%.

The relation of nemertine biomass to organic carbon

was distinctly bimodal (Tables 28, 29; Fig. 62). Largest
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 9 I

biomass (0.44 g/m2 ) occurred where organic

carbon ranged from 1.5 to 2%, declined at

higher and moderately lower levels, and

showed another somewhat lower peak (0.8 gl

m2) at levels between 0.01 and 1%. Intermediate biomass occurred where carbon was absent.

Frequency of occurrence of nemertines in

samples also exhibited a generally negative

correlation with organic carbon content

(Table 30). In areas with measurable amounts

of carbon (between 0.01 and 3%), occurrence

diminished from 46 to 15%. In areas with no

measurable amounts, occurrence was moderately high (40%).

Aschelminthes

30 0

ff0

ID

u. 25 0

a:

f-'" u.'" 0:1' 20

a:UJ UJa:

ID" :1'::>

::>0

Z(/) 15

a:

'"

0.

10

l

CJ NUMBER

• WEIGHT

:I'

o

f- (/)f-

:1'0

"ID

a: u. 4 '" 0

Za:

-",

f-fJ:;'"

",:I'

3

f-::>" ",0

~(/)

a:

'"0.

o

f- "'

~O.... «ID

0:: ....

"'0

:!':o::

2 ........ '" I'"

"'~

.... '" ~o::

<t .... => I .... 0

~'"0::

....

n.

D--o NUMBER

__ WEIGHT

relation to bottom

11\

1 \

1 \

1 \ ~-\-~

\

L --0

BOTTOM SEOIMENTS

We found nematodes in 98 samples (9% of total);

their density averaged 2.8/m2, and the biomass averaged 0.01 g/m2 (Table 5).

o'----;;o~.~3.90------;4.-:'.77..9,---o8-::f.1"1.9'---;'02,~,5<;.9'--""7'6;:-'-!;,9"'.9'--"'20'-'±-:23:-:.9"'---.J 0

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN OEGREES CELSIUS

0::

....

0.

Figure 60

Density and biomass of Nemertea in

sediments.

Figure 61

Density and biomass of Nemertea in relation to the annual range

of bottom water temperature.

0::'"

"'0:: ID«5 10

z",

o

ff- 30 (/)0

ZID

'" .... u O

"'0::

g, 20

....

o~'"

Nematoda-Free-living nematodes, members

of the phylum Aschelminthes, are one of the

most numerous animal groups inhabiting the

bottom sediments of the northeastern coast

of the United States. Previous studies conducted by Wieser (1960) in shallow coastal

habitats off southeastern Massachusetts revealed nematode densities of nearly 800,000

individuals/m2. Farther offshore, on the

southeastern coast of Massachusetts at a depth

of 58 m, Wigley and McIntyre (1964) encountered nematodes in densities of nearly 1 million individuals/m2, and this measure did not

include the young stages of numerous species. The vast majority of free-living nematodes are very small, less than 1 or 2 mm in

length; consequently only a small portion (the

large specimens) of the total population was

recovered and treated in the present study.

The bulk of the captured specimens were

between 0.5 and 2 cm long; however, a small

proportion of both larger and smaller specimens was represented. In addition to their

great numerical density, this group of animals is also exceedingly profuse in species

composition.

Three types offeeding habits are commonly

found among free-living nematodes. Some

are saprophagous, feeding on detritus and

dead and decaying animal material; others

are herbivorous, feeding on green plants, particularly diatoms; still others are carnivorous,

ingesting rotifers, tardigrades, small annelids,

other nematodes, and bryozoans.

Because of the incomplete representation of this

group, the quantitative distribution discussed herein

refers only to the exceptionally large species.
92 NOAA Technical Report NMFS 140

~-o NUMBER

_ WEIGHT 1.0

0

eno ZCD

IIJ III ... 13 0

IIJII:

lLlIJ

en~

... IIJ o~

II:IIJ

IIJII: CDC( 10 ~:;)

:;)0

Zen

II:

IIJ

lL

II

\

\

\

\

\

b....

0.8

0.2

o '-------;;o:';;.o::-1--0=-.Il=-------:-'1.'='0---:I'=.Il----;2:';.0;-----3=-.0~"tI---='Il.'='0~i-7.10~.0=----' 0

PERCENT ORGANIC CARBON

Figure 62

Density and biomass of Nemertea in relation to sediment organic carbon.

Geographic Distribution

The distribution oflarge free-living nematodes, although

occurring in all geographic regions, was patchy (Fig.

63). They were present in coastal areas, on the offshore

banks and basins, and on the continental slope and

rise. They were commonly absent in samples from a

number of areas: parts of the ova Scotian shelf, large

portions of the Gulf of Maine and Georges Bank, as well

as many inshore bays and sounds. The pattern of their

distribution suggests that they are less common in substrates where the overlying bottom current or wave

action is strong, for example, the northern section of

Georges Bank and at the mouth of the Bay of Fundy.

The average density of nematodes in each of the six

standard geographic areas was roughly similar, ranging

from 0.9 to 4.0 individuals/m2 (Tables 6, 7; Fig. 64).

Their average biomass was very low, 0.01 g/m2 or less

(Tables 8, 9; Fig. 64).

The percentage of samples containing nematodes

was low (5 to 9%) in the four geographic areas on the

continental shelf (Table 10), but their occurrence in

the two continental slope areas was comparatively high

(15 and 39%).

Bathymetric Distribution

Nematodes were found at depths ranging from 23 to

3,975 m and were present in all depth classes sampled

(Tables 11, 12; Fig. 65). Average density was slightly

higher in the very shallow (0-24 m) and moderately

deep (200-1,000) water than in other zones. Average

density values ranged from 0.8 to 6.8 individuals/m2.

Average biomass values were uniformly low (0.01 g/m2

or less) at all depths (Tables 13,14; Fig. 65).

Frequency ofoccurrence of nematodes in the samples

was low (2%) in shallow water and generally increased

to moderately high levels (35%) in deep water (Table

15) .

Relation to Sediments

Nematodes were present in all bottom types except

shell (Tables 16, 17; Fig. 66). Greatest density (8.7

individuals/m2) occurred in gravel substrates, whereas
Composition and Distribution of Macrobenthic Invertebrate Fauna 93

••0

42°

70°

M A IN E

M A IN E

71°

(8) 100

(JJf[) 100

NEW

JERSEY

EXPLANATI:]{@

121 1-9

• 10-49

• 151

GRAMS

PER SQUARE METER

7'·

7,0

NEMATODA

Figure 63

Geographic distribution of ematoda: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

all other bottom types yielded relatively moderate densities (I to 3 individuals/m2). Average biomass was small

(0.01 g/m2 or less) in all sediment types (Tables 18,19;

Fig. 66).

The frequency of occurrence of nematodes was moderate (12 to 13%) in gravel, sand-silt, and silt-clay but

was low (4 and 5%) in sand and till (Table 20).

Relation to Water Temperature

Nematodes occurred at all temperature ranges except

the highest, 20o-23.9°C (Tables 21, 22; Fig. 67). Their

numerical density was greatest (2.0 to 4.3 individuals/

m2) in the narrow to intermediate temperature ranges

(OO-11.9°C) and declined drastically (0.2 and 0.7/m2)

in the broader ranges.
94 NOAA Technical Report NMFS 140 _

o

I- U)I-

~o

c.'"

11:: ...

"'0

ZII::

-",

1-1-

:<:'"

"'~

"''''

~II::e 1-::>

",0

~U)

II::

'"...

o

IU)I-

~o

"lID

11:: ...

"'0

ZII::

-",

0.01 it;;

"'~

"''''

~II::

e 1-::1

",0

~U)

II::

'"

n.

o

0.01

0.02

o NUMBER

• WEIGHT

<>- -0 NUMBER

_ WEIGHT

500 1000 2000 4000 0

BOTTOM SEDIMENTS

WATER DEPTH IN METERS

10 25 50 100 200

NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTiA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPH Ie AREA

5

0 NUMBER

• WEIGHT

41-

31-

- 21-

IIo I

Figure 65

Density and biomass of Nematoda in relation to water depth.

Figure 66

Density and biomass of Nematoda in relation to bottom sediments.

Figure 64

Density and biomass of Nematoda in each of the six geographic

areas.

Biomass was small, averaging only 0.01

g/m2 or less, in all temperature ranges

(Tables 23, 24; Fig. 67).

The frequency of occurrence of nematodes in samples in the various temperature ranges diminished from a high of 17%,

where the temperature range was narrow,

to zero where the temperature range was

20°C or more (Table 25).

Polychaete worms formed a major component ofthe benthic fauna in terms ofbiomass and numbers of individuals. They were

present throughout the study area and

made up 28% of the total number of individual animals and nearly 10% of the total

biomass (Table 3).

Taxonomically diverse as well as abundant, this group of organisms contributed

over 300 species from among approximately

170 genera to the New England benthic

fauna.

Size differential from the smallest to the

largest specimen was moderate compared

with that for other taxa. The smallest specimens recovered were 3 to 4 mm in length;

the largest were over 200 mm. Although

the vast majority of annelids from these

collections are elongate and cylindrical in

shape (similar to the common earthworm),

the species that is largest in terms ofweight

is Aphrodita hastala, the sea mouse. It is

ovate in shape, ventrally flattened, convex

dorsally, and weighs 75 g or more. The

average wet weight of individual annelids

in the region is less than 0.05 g. A wide

Relation to Sediment Organic Carbon

Nematodes occurred only in the four low

to moderate level organic carbon content

classes (Tables 26, 27; Fig. 68). They were

most abundant (3.1/m2 ) in sediments with

low organic carbon levels (0.01-0.49%) and

least abundant (0.4/m2) at moderate carbon levels (1.5-1.99%).

Biomass was very low «0.01 to only 0.02

g/m2) in all levels ofsediment organic carbon in which they were found (Tables 28,

29; Fig. 68).

Frequency of occurrence in samples in

the four carbon conten t classes ranged from

7 to 17% (Table 30).

Annelida
Composition and Distribution of Macrobenthic Invertebrate Fauna 95

variety ofmorphologically different forms abound. All the

different feeding types--earnivores, suspension feeders,

and selective and nonselective deposit feeders-were represented in our samples. Many representatives from the

two major life modes, the errant (or free living) and the

sedentary (tubicolous) polychaetes were collected.

Coloration of polychaete annelids is extremely diverse, from nearly translucent and white to a dark brown.

Predominant hues are light beige, tan, dark brown, and

various shades of olive and red. Some annelids displayed a cuticular irridescence that greatly enhanced

their appearance. Some forms possessed variegated pat0-- ---<J NUMBER

0 5 ___ WEIGHT 0 ... ... ... ",'" "'0 ::EO ZCIl !\ oqCll .... ... 4 a: ... (30 / \ C>o

.... a: '\ / \ ~a:

g;~ .... 3 \ / \ ...... ....... :<: .... 0::E \ C>::E

a: .... 'y

\~ 0.01 ........ .... a: ~a: CIloq 2 oq ::E::> ... ::> ::>0 .... 0 Z", ~'" a: I a: .... .... a. a.

0 '-- 'r-. 0 0-3.9 4-7.9 B-11.9 12-15.9 16,19.9 20-23.9

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 67

Density and biomass of Nematoda in relation to the annual range of

bottom water temperature.

0.075

0.025

o

0.0ll0

0.01 0.5 1.0 1.5 2.0 3.0 5.0 10.0

PERCENT ORGANIC CARBON

Figure 68

Density and biomass of Nematoda in relation to sediment organic carbon.

0--- -0 NUMBER

... ....--. WEIGHT

... ?\

I \

/ \ - / / \ \

I \ / \

- / \ / \

I \ / \

I \ / \ - / \ / \ - \ / \ I \ / \

- I }J \

I \

I \ - I h.. - I

J - ~"''''''' '" o I ..

0.5

3.0
96 NOAA Technical Report NMFS 140

~~_~

120 KM

M A IN E

EXPLANATION·

[] <0.1-0.9

lTIilll 1.0-49.9

.50.0-324.5

El 1-99

ITIilll 100-999

.1,000-11,804 :.-::,..,.,....,.,.,.,.,.,.,."

NEW

JERSEY

PER

,--__""'-,:---T-__----;7~-~=--- ~77-'-2°----_Q7.:..IO----_____,7~0'--O----76"-90--,

ANNELIDA

Figure 69

Geographic distribution of Annelida: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

terns of color on various parts of the body, including

scarlet branching gills. Tubicolous forms, especially,

exhibited a wide range of color patterns on their tentacles and branchiae.

A few species are commensal on sponges and echinoderms, but more often they themselves are the hosts to

commensal partners, for example the ChaetopterusPinnixa (Decapoda) relationship and the ClymenellaListriella (Amphipoda) relationship.

Annelids occurred in 1,034 samples (96% of total),

their density averaged 425/m2,and their biomass averaged 17.4 g/m2 (Table 5).
Composition and Distribution of Macrobenthic Invertebrate Fauna 97

24

4

8

32

o

28

NOV GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENOLANO SLOPE NEW ENOLANO

SHELF SLOPE

GEOGRAPHIC AREA

0 NUMBER

• WEIGHT

-

'0<:-

-

)0-

-

00f-

-I

oc-

-I

00 -

- )01-

- -

0 A

100

20

o

.....

..... eno

ZeD

lL 5 (30

LUll:

lLLU

en .....

~~

II: LU 4 LUll:

eD4

~;:)

~g

II:

3

7

60

Figure 70

Density and biomass of Annelida in each of the six geographic areas.

Geographic Distribution

Annelid worms were found in all parts of the study area

(Fig. 69). Only the Mollusca, Crustacea, and Echinodermata compare with their widespread distribution,

and only the Crustacea are more numerous.

The density of annelids on the continental shelf and

slope generally averaged between 100 and 1,000 individuals/m2. Over approximately 10% of the shelf and

slope area, the density of annelids was high, 1,000 to

12,000/m2. About 30% of the shelf and slope area

yielded less than 100/m2. In deepwater regions beyond

the continental slope, the density was relatively low,

usually less than 100/m2•

The biomass of annelid worms on the continental

shelf and slope, with few exceptions, averaged between

1 and 50 g/m2 and reached a maximum of 325 gm2.

Among the six standard geographic areas, two slope

areas (Georges Slope and Southern New England Slope)

had the lowest numerical density and smallest biomass

of annelids, 80 to 149 individuals/m2 and about 4 to 5

g/m2 (Tables 6-9; Fig. 70).

In the four continental shelf areas the numerical density of annelid worms was high. Average densities ranged

from 291 to 648 individuals/m2 and greatest density was in

the Nova Scotia area. Biomass in these areas, also, was

moderately high; average values ranged from 8 to 30 g/
98 NOAA Technical Report NMFS 140

5

30

Figure 71

Density and biomass of Annelida in relation to water depth.

°0':---,1~0-----"J2~5-+.50=---,-10:!-:0,-----2::-:!0::::0---=-:!-::--~:-::~~~0

WATER DEPTH IN METERS

0---0 NUMBER

_ WEIGHT

Boor--------------------,

sediments, and the lowest occurred in till. Intermediate

and nearly equal quantities, 15 to 16 g/m2, were present

in the other four bottom types (gravel, shell, sand, and

silt-clay) (Tables 18, 19; Fig. 72).

Annelid worms were present in all samples from till,

shell, and sand-silt sediments; however in 97% ofsamples

in gravel and silt-clay. Although annelids were present

in 94% of the samples from sand sediments, this category of bottom type ranked last in this relationship

(Table 20).

Relation to Water Temperature

Annelids were present in greatest quantities (Table 21;

Fig. 73) where the annual range in bottom water temperature was broadest, and densities generally declined

as the temperature range decreased. Where the temperature range was greater than 20°C, average density

of annelids was 1,698 individuals/m2. At the other extreme, where the temperature range was less than 4°C,

average density was 212/m2. At intermediate levels of

temperature range, density was intermediate, 280 to 568

individuals/m2. The percentage composition of numbers

in this group of organisms in relation to the total fauna in

the various temperature-range classes varied from 16 to

49% (Table 22). Percentages were highest where temperature ranges were extremely high and low.

Biomass of annelids in relation to range of water temperature was similar to that for density. Biomass was large

(about 40 g/m2), not only in relative terms but in absolute

quantities, in areas where the temperature range was broad

::IEsoo

g

<Il~

~~::lEu.. -0

o

UJa:

<l.UJ

en 400

u.. UJ

0::IE

a: UJ UJa:

m<l:

::IE:>

~ga:

200

m2. Largest average biomass occurred in the Southern ew England Shelf area (Tables 6, 8).

The percentage of the total fauna made up by

annelids was high in both biomass and number

of individuals (Tables 7, 10). On a numerical

basis, annelids accounted for 22 to 56% of the

total fauna. In terms of biomass, annelids formed

between 3 and 23% of the total fauna. In the two

deepwater zones, Georges Slope and Southern New

England Slope, the percentage of the total biomass made up by annelids was 22 and 23%, respectively. This is an especially large proportion to be

formed by anyone faunal group and a much higher

proportion than that from the continental shelf

areas where the percentages were only 3 to 12.

Annelid worms were present in an exceptionally large proportion of the samples. In all standard geographic areas, except Georges Bank,

they were present in 96 to 99% of the samples.

On Georges Bank they were present in 89% of

the samples (Table 10).

Bathymetric Distribution

Annelids occurred over the entire depth range

in which samples were collected, 3 to 3,975 m.

Numerical density and biomass were highest in shallow

water and decreased in quantity rather uniformly with

increased water depth (Fig. 71). Average density in the

shallowest depth class was 719 individuals/m2. High

densities (437 to 159/m2) prevailed across the continental shelf and then diminished rather sharply down

the slope and on the continental rise, where the average densi ty was only 9/m2 (Tables 11, 12).

The biomass of annelids followed precisely the same

quantitative trend in relation to water depth as was

exhibited by numerical density. Average biomass was

largest (27 g/m2) in the shallowest depth zone. Biomass was rather high (15 to 25 g/m2 ) on the continental shelf and dropped to 5 g/m2 at midslope and to 0.76

g/m2 on the continental rise (Tables 13, 14).

Annelids were present in a very high percentage of

the samples from all depth classes. Frequency of occurrence among the eight classes ranged from 93 to 100%

(Table 15).

Relation to Sediments

Annelid worms were not only present in all sediment types

sampled but occurred in roughly the same density in each

of the different sediment types (Tables 16, 17; Fig. 72).

Average density from all sediment types ranged from 232

to 558 individuals/m2. Greatest densities were found in

sand and gravel bottoms, lowest densities in silt-clay and till.

Biomass, also, was rather evenly distributed among

the various sediment types. Values ranged from 11 to 26

g/m2. Highest average biomass occurred in sand-silt
Composition and Distribution of Macrobenthic Invertebrate Fauna 99

:::E

o

.... <n ....

:::EO

«al

6

ZCl::

-",

.... ....

:1:'" (!):::E

~~ 2 « .... ::> "'0

~<n

Cl::

'"0..

o

4

28

24

20

8

SAND- SILTSILT CLAYGRAVEL TILL SHELL SAND

0 NUMBER

• WEIGHT

-

-

0-

-

- -I

r- -I

I-

-

r- -

o

100

300

200

400

:::E

o

....

.... <no

Zal

'":::Eu..

Lio

~~<n ....

u..'" 0:::E

Cl::'"

"'Cl::

al«

:::E::>

::>0

Z<n

Cl::

0..

'"

600

50

BOTTOM SEDIMENTS

Figure 72

Density and biomass of Annelida in relation to bottom sediments.

(Tables 23, 24; Fig. 73). Conversely, quantities were smaller

(about 10 g/m2) where the temperature range was small.

Moderate biomasses were encountered in areas where the

temperature range was moderate.

The frequency of occurrence of annelids among the

temperature-range classes was high and fairly uniform.

They occurred in 92 to 97% of the samples (Table 25).

Relation to Sediment Organic Carbon

Annelids of the New England region exhibited an essentially bimodal relationship to the amount of organic

carbon in the sediments (Fig. 74). Greatest density

(504/m2) occurred at low organic carbon levels (between 0.01 and 0.49%); another peak in density (407/

m2 ) occurred at higher levels (between 2 and 3%);

moderate densities prevailed at levels between these

two peaks, with smallest densities occurring in both

lowest and highest levels (0.00 and 5.00+%) of sediment organic carbon (Tables 26, 27).

Annelid biomass was greatest (27 g/m2 ) at organic

carbon levels between 1.5 and 2% (Tables 28, 29; Fig.

74) and gradually diminished at levels both above and

below these values. As with density, lowest biomass was

found in both the lowest (0.00%) and highest (5.00+%)

organic carbon levels.

Frequency of occurrence of annelids in the various

organic carbon content classes was uniformly high, ranging from 80 to 100% in all classes except one, the 3-5%

class, in which only 50% of the samples contained members of this group (Table 30).
100 NOAA Technical Report NMFS 140

Pogonophora

0L-_O-.L3.-.__,.L.7.~'-~8..l."~.'-=12...L.15""'.'''--=16...L.'''"''.'''--=20,....J.2L,.3~.'--.10

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 73

Density and biomass of Annelida in relation to the annual

range of bottom water temperature.

Bathymetric Distribution

Pogonophores were found at depths from 141 to 2,870

m. Not only were members of this phylum restricted to

deep water, but their density, biomass, and frequency

of occurrence were higher in the middle and deeper

portions of their range than in shallow parts (Fig. 77).

:IE

o

....

40 b .. " cr ... 0>0

Zcr

-w

........

X W 30 C)

Ww

~cr.. .... :::>

~g

cr 20

10

between 1 and 511m2. Because of their small size and

low density the average biomass of pogonophores was

very small, less than 0.2 g/m2 in all localities sampled.

Among the six standard geographic areas, pogonophores were present in significant quantities only in

the Georges Slope and Southern New England Slope

areas (Tables 6-9; Fig. 76) where density in those two

areas averaged 3 and 51m2 , respectively, and biomass

0.01 and 0.03 g/m2, respectively.

In view of the relatively recent date (Ivanov, 1963)

this phylum was first reported from the New England

Region, the frequency of occurrence in this series of

samples should be considered unexpectedly high.

Pogonophores occurred in 35% of the samples from

Georges Slope and in 42% of the Southern New England Slope samples (Table 10). In two other geographic areas they were present in 2% or less of the

samples. They were absent in the Georges Bank and

Southern New England Shelf areas.

1800

0--0 NUMBER

..-----. WEIGHT r 1600 I

I

1400 I

I 2; I 0

.... 1200 .... I "'0 z., I W

2;"-

,,0 1000 Wcr I g,~

,,-w I 02;

crw 800 wcr

"<1 I 2;:::> I :::>0

Z",

cr 600

w I Q.

I '00 \ j

\ --- V

200 /

Pogonophora (beard worms) are a minor

consituent of the New England benthos. They

provided less than 0.1 % of the total number of

specimens and biomass of organisms in the study

area (Table 3). Nevertheless, they contributed

some unique records to the study. Chief among

them was a proclivity for deep, cold water. These

unusual animals were one of only a few taxonomic groups that were more abundant in deep

water than in shallow water.

Pogonophores inhabit chitinous tubes buried

in the bottom sediments. The tubes in our collections ranged in length from about 5 to 15 cm

and had diameters from 0.1 to 0.4 mm. The

colors of the tubes varied from very light tan to

dark brown, but were most frequen tly of a greenish-yellow to brownish-green hue. The m~ority

of them exhibited alternating light and dark

rings or bands. Embryos were not uncommon in

the tubes from our samples. Preserved specimens varied in color from whitish to brown; the

most common colors observed were cream to

light reddish tan.

The two existing orders of pogonophores, the

Athecanephria and Thecanephria, are represented in our collections by species from the

genera Siboglinum, in the former, and Diplobranchia and Crassibrachia, in the latter. Six species were

obtained; the most common were Siboglinum ekmani

Jagersten, S. pholidotum Southward and Brattegard, and

Diplobrachia similis Southward and Brattegard. Less common were Siboglinum holmei Southward, S. angustum

Southward and Brattegard, and Crassibrachia sandersi

Southward. Siboglinum holmei was distinctive for its occurrence in the Gulf of Maine and was the only species

of this phylum found in the Gulf. Furthermore, it was

taken in close proximity to land, at the point nearest

land, at station 1171, less than 5 km from Grand Manan

Island and 10 km from the mainland coast of the U.S.

Water depth at this location is 141 m.

Pogonophora occurred in 56 samples (5% of total);

their density averaged 0.6/m2 and their biomass averaged less than 0.01 g/m2 (Table 5).

Geographic Distribution

Pogonophores were widely distributed along the continental slope and continental rise from NewJersey northward to Nova Scotia but rarely occurred on the continental shelf (Fig. 75). In coastal waters they were encountered in only the most northerly part of the study

area, near Grand Manan Island (New Brunswick),

Canada, and Eastport, Maine. Their average density

throughout the study area was low or moderately low,
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 101

!l00

400

I- III 0 ZcD

1&1

l!> 300

1&111::

lLl&I

III I-

... 1&1

o~

11::1&1

1&111::

cDC

~5 200 ZIIl

II::

1&1

lL

100

o

1\

1\

I \

I \

I \

f \

I \

I \

I \

I \

I \

I \

I

I

I

I

I

0---0 NUMBER

_ WEIGHT

\

\

\

\

\

\

\

\

\

\

\

\

\

'\

'\

'\

2!l

OL-----::0~.O..,..I---:0.=-!l--..,.1."=0--...,I"=.!l,-----...,2-:L..O=----='3.'='O---="=---....,IO~.o=------J

PERCENT ORGANIC CARBON

Figure 74

Density and biomass of Annelida in relation to sediment organic carbon.

Average density was highest (6.8 individuals/m2) between

500 and 1,000 m. Density was slightly lower (3.2 to 3.51

m2) in deeper water (1,000 to 2,870 m) but substantially

lower in depths less than 500 m (Table 11).

Biomass ranged from a to 0.03 g/m2 , and the trend

in relation to water depth was comparable to that described for density (Table 13).

Pogonophores were present in approximately half

the samples from depths greater than 500 m. In the two

shallower but adjacent depth classes (100-199 m and

200-499 m), there was a sharp drop in their occurrence

to 5 and 1%, respectively, They were absent in the three

shallow depth classes (Table 15).

Relation to Sediments

Pogonophores were absent in coarse textured sediments

but were increasingly common as the sediment particle

size decreased from sand to silt-clay (Fig. 78). The

density of pogonophores was exceedingly sparse «0.1 I

m2) in sand sediments, intermediate in sand-silt, and

highest (1.91 m2) in silt-clay (Table 16).

Their biomass was very low «0.01 glm2) in sand, and

only slightly higher in the other fine-grade sediments

(Table 18).

The occurrence of pogonophores in the samples correlated very closely with average density. They were

exceedingly sparse (occurring in <l %) in samples from

sand sediments, moderately sparse in sand-silt, and most

common (16%) in silt-clay substrates (Table 20).

Relation to Water Temperature

Pogonophores were restricted in their distribution to

areas that exhibited an annual bottom temperature

range ofless than 12°C (Fig. 79). The vast majority were

obtained where the temperature range varied less than

4°C. In these waters their density averaged 1.5/m2 and

the biomass averaged 0.01 g/m2 (Tables 21, 23). Even

in areas where the temperature range was between 4°

and 11.9°C they were exceedingly sparse in both density (0.1 and 0.7/m2) and biomass «0.01 g/m2).

The areas of small temperature variation (see Fig.

11) correspond to the deepwater regions on the con ti-
102 NOAA Technical Report NMFS 140

NEW

JERSEY

EXPLANATION

[] 1-9

1m 10-51

EXPLANAT'ON

[] <0.1-0.2

GRAMS

PER SQUARE METER

•••

72·

POGONOPHORA

70· ••• '20 '----~_~I KM

Figure 75

Geographic distribution of Pogonophora: A-number of specimens per square meter

of bottom; B-biomass in grams per square meter of bottom.

nental slope and continental rise (see preceding sections on geographic and bathymetric distribution).

Among the three temperature range classes in which

pogonophores were found, their frequency of occurrence was highest (15% of the samples) where the

temperature range was smallest. Frequency of occurrence was very low in the other two temperature range

classes in which they occurred.

Relation to Sediment Organic Carbon

Pogonophores were found where sediment organic carbon content was low to moderate, ranging from 0.01 to

slightly under 2% (Fig. 80).

Highest density (2.2/m2) occurred in carbon contents in the 0.50 to 0.99% range, and lowest (0.11 m2) in

the 1.50 to 1.99% range (Table 26).
Composition and Distribution of Macrobenthic Invertebrate Fauna 103

WATER DEPTH IN METERS

Figure 77

Density and biomass of Pogonophora in relation to water depth.

0.04

:.

0

.... 0.03 "' .... :.0

.. II)

a: .... C>o

:!:a: ... 0.02 ........

:<:'" c>2

......

~a:.. .... :>

0.01 ... 0

~'"a:

...

lL

,

,

,

,

\

tr- -..()

0---<> NUMBER

_ WEIGHT

oL-----':-:=------="~':-----;;-;GEc:-O.==G::::ES:--S::::OU""T:f:H""E.::::.,.......,G==EO~.~GE:::S,..--;:S::!:OU~T~HE!-::."". ---'0

SANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

o NUMBER

• WEIGHT

Figure 76

Density and biomass of Pogonophora in each of the six geographic

areas.

Sipunculids, or peanut worms as they are

sometimes called, are a taxonomically small

group but are found in great abundance in

some habitats and have a wide distribution

in many parts ofthe world ocean. Although

locally abundant in New England waters,

they account for less than 0.5% of the total

benthic fauna in terms of both numerical

density and biomass (Table 3).

Peanut worms are elongate and cylindrical in shape and characteristically burrow

into the bottom sediments where they feed

predominantly on detritus. A few species

utilize abandoned gastropod shells for permanent shelter, and one small species was

found to inhabit empty foraminifera tests.

Specimens in our samples ranged in

length from about 0.5 to 12 cm.

Brown, in various shades, was the predominant color exhibited by ourspecimens;

most often they were tan, flesh, or reddishbrown.

Sipunculids occurred in 249 samples (23%

oftotal) .Their density averaged 5.9/m2, and

biomass averaged 0.75 g/m2 (Table 5).

Sipunculida

Biomass was low (0.01 to <0.01 g/m2 ) in

the four organic carbon range classes in

which they occurred (Table 28).

The greatest frequency of occurrence

(21 %) in samples was in the 0.50 to 0.99%

range class, but frequency diminished rapidly in the higher and lower adjacent classes

(Table 30).

Figure 78

Density and biomass of Pogonophora in relation to bottom sediments.

BOTTOM SEDIMENTS

o

.... en .... ~o

.. lD

a:LL

C>o

:!:a:W

.... .... :I:'"

C>~

"'w

~a:.. .... :> ... 0 ~en

a:

W

lL

0.01

o NUMBER

• WEIGHT

o

....

.... eno

ZlD

w ....

U 0 3

~ffi en .... LLW o 2

a:wwa:

CD ..

~:> ::Jo I z",

a:

W

lL OL--""'GR:-:A~VE:-:L----=TI~L:-:L--:S:-:H"'"EL:-:L----::c~---::":-~---!c:-'!"'---'O

Geographic Distribution

Sipunculids were widely distributed over

the study area (Fig. 81), occurring in low

densities over large areas of banks and basins. Densities generally averaged between

1 and 49 individuals/m2 and exceeded

these values in only three localities. The

maximum density detected was 152 individuals/m2.

Average biomass ofsipunculids was moderately low, usually 1 g or less per square

meter, and with few exceptions was less

than 10 g/m2.

Among the six standard geographic areas there was rather little difference in average density (Table 6; Fig. 82); the range

was about 1 to 9 individuals/m2. Slightly
104 NOAA Technical Report NMFS 140 _

a::

W

ll.

average weight was nearly forty times greater, 4 g per

individual.

Frequency of occurrence of sipunculids was moderate to moderately low. They were present in 13 to 42%

of the samples (Table 10). Highest frequency of occurrence was in the Nova Scotia area and lowest was on

Georges Bank.

Bathymetric Distribution

Sipunculids were taken at depths ranging from 16 to

3,975 m. They were most common at moderate depths,

25 to 500 m, where they averaged about 6 to

8 individuals/m2 (Table 11; Fig. 83). In both

shallower and deeper zones they averaged

nearly 1 or 21m2 .

The relation between biomass of sipunculids and water depth was substantially different from that of numerical density. Biomass was much greater (averaging about 1 to

4 glm2) on the middle and lower portions of

the continental slope (500 to 2,000 m) than

it was on the continental shelf and continental rise (Table 13, Fig. 83). Also, sipunculids

formed 10 to 26% of the total benthic biomass at these depths, compared with less than

1% for the shallower zones and 5% for the

continental rise (Table 14).

The percentage of samples containing

sipunculids was lowest (3 to 13%) at depths

less than 50 m (Table 15). At depths greater

2;

o

... VJ'"

2;0

~1Il

a:: ... Cl O

za::

-w

......

IW

~2;

Ww

~a::

... :> wo

~VJ

-0.01

-0.02

0----0 NUMBER

_ WEIGHT

~,~ 0'---"0--';.3~.9:----;;-4 -',;.7-;;-.9----;e:;-·~'1.;;-9 -=~,2"".,~5.9;;---;,-;:-6 ·-t;,9;-;.9:---;;;20"-~23;-;.9;-----'0

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

a::

W

ll.

Figure 79

Density and biomass of Pogonophora in relation to the annual

range of bottom water temperature.

higher average densities were found in Nova Scotia,

Southern New England, and the Southern New England Slope. The lowest density (1.2/m2) was in the

Georges Slope area.

Differences in average biomass were also comparatively small among the six geographic areas (Table 8;

Fig. 82). Largest average biomass (1.83 glm2) occurred

in the Southern New England Slope area; the smallest

(0.37 g/m2 ) was in the Gulf of Maine.

Individual sipunculids averaged approximately 0.1 g

in weight in all areas except Georges Slope, where their

3.0r------------------------------------.

- 0.050

- 0.025

0---0 NUMSER

.-. WEIGHT

I \ I"

/ \

/ \

I \

/ '\ I \

/ \

/ \

I \

I \

I....... \

L ......__;;;-.,;-d°t:::::::-LI----l,--=:::t;::::=:tO-b~- ....._4~~....,..L.....I~l____....,.LI___4~-1._I--..J o 0.01 0.5 1.0 1.5 2.0 3.0" 5.0 .. 10.0 0

PERCENT ORGANIC CARBON

2

o

I-

(1)1-

2 0 C[III

a::1L

00

za::

-1&.1

1-1-

~I&.I

0 2

1&.11&.1

~a::C[

1-;:)

I&.I~

a::

1&.1

Q,.

Figure 80

Density and biomass of Pogonophora in relation to sediment organic carbon.
Composition and Distribution of Macrobenthic Invertebrate Fauna 105

72·

72°

NEW

JERSEY

EXPLANATION

[ill 1- 49

IIIlI 50 -152

EXPLANATION

[ill <0.1 - 9.9

II 10.0 - 28.5

GRAMS

PER SQUARE METER

,.0

72°

SIPUNCULIDA

4,0

70°

M A IN E

42°

Figure 81

Geographic distribution of Sipunculida: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

than 50 m the percentage of occurrence ranged from

21 to 32%, with the highest rates occurring at depths

<1,000 m and the lower rates at depths >1,000 m.

Relation to Sediments

Sipunculids inhabited all sediment types and occurred

in about the same density in each type (Table 16; Fig.

84). The range in average density in the various types of

bottom sediments was from 4.0 to 7.l/m2. Sand contained the highest density.

Biomass of sipunculids in relation to various sediment types was considerably more varied than density

ofsipuncuiids (Table 18; Fig. 84). Biomass ranged from

low values (0.16 to 0.29 g/m2 ) in shell and till sub-
106 NOAA Technical Report NMFS 140

o

::Ii

.0

",'"

~:il

.5

z'" -",

... ... a

~~.. 0.5 5 ;a",

'"

'"ll.

NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANt< NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

r- o NUM8ER -2 _ WEIGHT

-I

r-

-I.

f--

I -I I o

2

o 12 ...

...

o

CD

IL

o

'"... '" 1L'" 02

"''''

~~ 4 ::Ii::>

::>0

z'"

'"'"ll.

strates to high quantities (0.81 and 0.89 g/

m2) in sand and sand-silt.

Specimens from the shell substrates were

the smallest in individual size.

Frequency of occurrence of sipunculids

in the samples was moderately low and

approximately equal among the different

types of bottom sediments (Table 20).

Range in percentage occurrence was 17 to

28, with highest values in sand and sandsilt; lowest values occurred in shell and siltclay sediments.

WATER DEPTH IN METERS

:::Ii

o

I-

",Io :::Ii 0 .. m Q:lL

<:>0

.8 Z Q:

- ... 1-1-

6

...... .4

1-::1

.2 g

Q:

...

Q,

SAND- SILTSILT CLAY

500 1000 2000 4000 0

SHELL SAND

BOTTOM SEDIMENTS

25 50 100 200

p---q

/' \

0--4 /

/

/

/

/

0'

GRAVEL TILL

0- - -0 NUMBER

__ WEIGHT

D NUMBER

• WEIGHT

-I.

Bf- -p

f- -0

r- -0

21- I -p

• 0

Figure 83

Density and biomass of Sipunculida in relation to water depth.

Figure 82

Density and biomass of Sipunculida in each of the six geographic

areas.

Figure 84

Density and biomass of Sipunculida in relation to bottom sediments.

Relation to Sediment Organic Carbon

Sipunculids showed a decided preference

for sediments with moderately low to low

amounts ofsediment organic carbon (Fig.

86). Both measures of abundance were

greatest in the two classes between 0.01

Relation to Water Temperature

Sipunculid density, biomass, and frequency

ofoccurrence generally tended to decrease

as the range in temperature broadened

(Fig. 85). None were found where the temperature range exceeded 20°C.

Density averaged about 5 to 9 individuals/m2 where the temperature range was

restricted and diminished to zero where

the temperature range was most extensive

(Table 21). The percentage of the total

benthic fauna made up of sipunculids decreased at a rather uniform rate as the

temperature range increased (Table 22).

Biomass of sipunculids was low or moderately low, and the changes in quantity in

relation to temperature range followed precisely the same pattern as those in relation

to density (Tables 23, 24). Biomass averaged about 0.7 to 1 g/m2 in the narrow

temperature range classes and decreased

to zero where the temperature range was

greater than 20°C. The percen tage biomass composed ofsipunculids also decreased

as the temperature range expanded.

Frequency ofsipunculids in the samples

ranged from 0 to 29% (Table 25). They

occurred most frequently, 25 to 29%, in

samples where the temperature range was

less than 12°C. Where the temperature

range was broader than 12°C, the occurrence ofsipunculids dropped substantially

and they were absent in samples from areas where the temperature range was more

than 20°C.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 107

0-- --0 NUMBER

::E _ WEIGHT ::E

0 10 I- ",l- II.O-""'~~ ::EO "'0 ZlXl 4 al

'" B 1.0 0: ... ::IE'" "'0 c:;0

! "'0: ~o: n.", O.B .... '" 6 / 1-1-

:1:'"

l5::E ! ",::E

0.6 0:'" "'''' ~o:

~~ 4 <t ::E::::l I-::::l ::::l0 0.4 ",0 Z", ~'"

0: 2 0:

w .... n. 0.2 n.

0 0 0-3.9 4-7.9 8-11.9 12-15.9 16-19.9 20- 3.9

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 85

Density and biomass of Sipunculida in relation to the annual range

of bottom water temperature.

8.0

O-~ NUMBER

.............. WEIGHT

S

1.2 g fI)~

SO

e(ID

a:1L. (!)o

!a:w ~~

0.8

~~e(

~5

~(I)a:

w 0.4 Q.

_-1

9'"

I

I

I

/

I

/

/

I

/

I

/

I

d

\

\

\

\

\

\

\

\

\

\

\

\

\

\

"-"- "-

oL-4--:~--~----;-l;:---===?~"-=a._::I_.::_-e'--*---'I--~:_.....__:_!_=__--'o 0.01 0.5 1.0 1.5 2.0 3.0 5.0 10.0

PERCENT ORGANIC CARBON

6.0 0

fl)0

ZID

W

~II..

u O

wa: Q.

fI)~w

4.0 11..1&1

O~

a:w

wa:

IDe(

~=>

=>0 ZfI)

a:

w 2.0 Q.

Figure 86

Density and biomass of Sipunculida in relation to sediment organic carbon.

and 0.99% organic carbon. Abundances were low where

carbon was absent, but sipunculids were entirely absent

in sediments whose carbon levels were 2% or greater.

Density ofsipunculids ranged from 0.1 to slightly more

than 7 individuals/m2 (Table 26), and biomass ranged

from <0.01 to a little more than 1.1 g/m2 (Table 28).
108 NOAA Technical Report NMFS 140

Frequency ofoccurrence ofsipunculids in the samples

paralleled the trends established by density and biomass. Range of occurrence was from 2 to 31 % (Table 30).

Echiura

Echiurid worms, like sipunculids and priapulids to which

they are taxonomically allied and which they resemble

somewhat in terms of size and habits, form a rather

obscure and small group and are not very well known in

this region. They are not especially abundant in the

New England region, accounting for only 0.2% of total

biomass and >0.1 % of total density (Table 3).

Echiurids are round, unsegmented worms that typically burrow into sand and mud. Others are found

among rocks and on coral reefs; one southern spec:.es

uses the tests of sand dollars as a habitat.

Their size varies greatly, ranging from tiny 3-rr..m

males to large females over 300 mm in length. Specimens in our samples were in the 2 to 8 em size range.

They are reported to prefer shallow waters; however,

the majority of our specimens were collected from deep

water. Depth range of our samples was from 20 to 3,975

m, but only 5 of the samples were in water depths ofless

than 1,000 m.

Color ofspecimens ranges from drab grays and brown

to green, red, and rose colored, and some were transparent.

Echiurids occurred in 17 samples (1.6% of total). Density averaged 0.l/m2 and biomass 0.30 g/m2 (Table 5).

Geog;raphic Distribution

Echiurids were almost exclusively restricted in their

geographic distribution to the lower continental shelf

and upper continental slope and were found in rather

small, discrete patches (Fig. 87). The only exceptions

were two small areas inshore, one at the mouth of Long

Island Sound and the other near Mt. Desert Island in

the Gulf of Maine. Densities averaged between 1 and 9

individuals/m2, whereas average biomass ranged from

<0.1 to a high of 12.5 g/m2.

Echiurids were absent from Nova Scotia and Georges

Bank and were present in generally equitable densities

in the other standard geographic areas. Highest average densities occurred in the two slope areas (Table 6;

Fig. 88). Biomass was also highest in the slope area;

significantly lower values were observed in the other

areas in which they were found (Table 8; Fig. 88).

Frequency of occurrence of echiurids in samples ranged

from <1 % to 15%. Lowest occurrence was in Gulf of

Maine and highest on Georges Slope (Table 10).

Bathymetri~Distribution

Echiurids were found at depths ranging from 20 to

3,975 m. Their numerical abundance was greatest, albeit low (0.5 to 0.6/m2 ), in water depths greater than

1,000 m and even lower (0.02-0.31 m2) at shelf and

inshore depths (Table 11, Fig. 89).

The relation of biomass to water depth was similar to

that of numerical density. Continental slope depths (1,000

to nearly 4,000 m) provided highest mean biomass ofthis

small group ranging from 3.5 to 5 g/m2. Significantly

lower (0.22 to 0.01 g/m2 ) biomasses occurred in inshore

and midshelf depths, respectively (Table 13; Fig. 89).

Echiurids were found in 19 to 21 % of the samples in

the two depth range classes below 1,000 m, but in only

<1 % of the samples in the 100-199 m range and 2% in

the 0-24 m range class (Table 15).

Relation to Sediments

Echiurids in our samples were rather restrictive in their

choice of sediment, preferring to inhabit only the two

finest-grained types. Both mean density (0.2 and 0.31

m2 ) and biomass (0.79 and 0.69 g/m2 ) were quite evenly

apportioned between sand-silt and silt-clay, respectively

(Tables 16, 18; Fig. 90). Three percent of the samples

in sand-silt and 5% in silt-clay contained specimens

(Table 20).

Relation to Water Temperature

Considering the deep water and fine sediment preferences of echiurids, it is not surprising to find that they

also had restricted temperature preferences. They occupied only three temperature ranges; areas with the

most stable annual range (0-3.9°C) contained the lowest density (0.l/m2) but the highest biomass (0.9 g/m2);

areas with an annual range of 16-19.9°C had the highest density (0.4/m2 ) and also contained the second

highest biomass; and areas where the temperature range

was between 4° and 7.9°C had moderate density and

low biomass (Tables 21, 23; Fig. 91).

Frequency of occurrence ranged from 1 to 4% (Table

25), being highest in the narrowest temperature range.

Relation to Sediment Organic Carbon

Echiurids were also restrictive in relation to organic

carbon content of sediments, occurring only where

amounts ranged from 0.01 to 1.49% (Fig. 92). Density

was greatest in carbon contents 0.50 to 0.99%, falling to

lower levels in areas of both lesser and greater content

(Table 26); conversely, biomass showed an increasing

trend with organic carbon content (Table 28). Frequency of occurrence in samples reflected the trend

for density (Table 30).

Priapulida

Among the invertebrate fauna of the New England

region priapulid worms make up perhaps the rarest
Composition and Distribution of Macrobenthic Invertebrate Fauna 109

u· 100

.,.

..,

- .... ,...... 4'· "-

"

oz·

'0"

II A I HE

II A I HE

.,.

-/...----......--- ------ -," "-

"

,,'

---

...

"

"~--'_J',

•

1000-_.",

~'

100

EXPLANATION

[ill 1-9

EXPLANATION

[])<O.I- 0.9

ml1I 1.0 - 12.5

HEW

JERSEY

GRAMS

PER SQUARE METER

n·

?I'

...

ECHIURA

Figure 87

Geographic distribution of Echiura: A-number of specimens per square meter of bottom; B-biomass in grams per square meter of bottom.

and least known group; they are also uncommon in

other areas of the world ocean as well-only eight species belonging to six genera have been reported (Barnes,

1974). Priapulids contributed <0.1 % of the total number ofspecimens and biomass in the region (Table 3).

Priapulids are cucumber-shaped predacious worms

that burrow into sand and mud and feed upon other

soft-bodied, slow-moving invertebrates, especially polychaete worms. They are typically between 4 to 8 em. in

length. Our specimens were considerably smaller, in

the 1 to 2 em size range. Color of specimens ranges

from whitish to flesh colored with some yellow on appendages; some of our specimens were somewhat reddish brown.
110 NOAA Technical Report NMFS 140

:E

0

.... en .... :EO

<till

a:1L - 08 "'0

za:

-",

- 0.6 .... ....

:1:'" ",2

- 0.4 "''''

~'"<t .... :::> ",0 - 0.2 ~'"

'"'" 0 Q.

0- -<l NUMBER

_ WEIGHT

WATER DEPTH IN METERS

BOTTOM SEDIMENTS

Figure 89

Density and biomass of Echiura in relation to water depth.

o '------;G""RA~V;;:-EL;------;CT,-eeLL----;cSH:;;EccLL;---;;-;SA:7:ND;;-------}SA~N~D'----+.SI ~LT·,----J

SILT CLAY

:E

g

~bw'" :E ....

U 0 08 w'" g;w

IL t;j 06

o:E

a: w w 0.4

~5CIl 02

a:

w

ll.

0 NUMBER

2 • WEIGHT 2 0 -~ 0 .... .... ....

0 "' .... 2° III .. Ill

IL -4 "'IL

° "'0

'" z",

'" -",

.... -3 ........ IL'" 0.4- :1:'" 02 ",2

"'''' "'''' "'''' 0.3- ~'" 1Il<t -2 <t

2:::> ....:::> :::>0

z'" 0.2 i- ~~

'" -I '" '"

Q. '" Il Q.

01'- n .. 0 0 NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

2

0 OAi- 0 NUMBER ....

.... • WEIGHT 0

III

IL 0.3i- 0

'"

'".... IL'" 0.21- 02

"'''' 1Il<t "''''

:E:::> o Ii- :::>0

Zen

'"

'"

Q.

Figure 88

Density and biomass of Echiura in each of the six geographic

areas.

Only 4 (0.4% of total) of our samples yielded

specimens (10 individuals weighing a total of

4.60 g) whose mean density was <0.l/m2 and

whose biomass was <0.01 g/m2 (Table 5).

Priapulids in the New England region were

found in the deep (1,420-2,035 m), cold (0-

3.9°C), sand-silt and silt-clay bottoms of Georges

and Southern New England Slope waters, where

organic carbon levels range from 0.01 to 1.49%,

in very low abundance (Figs. 93-98).

Polyplacophora-This class of mollusks is also

called chitons, which are bilaterally symmetrical, have eight overlapping dorsal plates, and a

broad, flat, ventral foot. They accounted for

less than 1% of the biomass and number of

animals of the total benthic invertebrate fauna

(Table 3).

Specimens ranged in size from 4 mm to 2 cm

and exhibited considerable variation in color.

They were commonly chalky white or various

shades of light gray; a small proportion were

light brown. A few had a dark, nearly black

coating over their plates that contrasted sharply

with the underlying white or light gray.

Chitons were relatively common in the coastal

areas, on relatively shallow offshore banks, on

coarse bottom sediments, and where the water

temperature range was moderately broad.

Polyplacophorans occurred in 84 samples

(8% of total). Their density averaged 1.5/m2,

and biomass averaged 0.14 g/m2 (Table 5).

The phylum Mollusca contributes significantly

to both measures of abundance (numerical

density and biomass) in the New England region as it does in the Middle Atlantic Bight

region (Wigley and Theroux, 1981). The molluscan fauna comprises five classes: Polyplacophora, Gastropoda, Bivalvia, Scaphopoda,

and Cephalopoda. Each of these classes will be

discussed separately below. For the sake of continuity in the phylogenetic ordering of figures,

those figures dealing with phylum Mollusca (Figs.

99-104, inclusive) are included here, but a detailed discussion of the phylum as a whole will

be presented in the section "Dominant Components of the Macrobenthos" below.

Mollusca

Geographic Distribution

Polyplacophorans, although rather sparse, occurred in small to moderately large geographiFigure 90

Density and biomass of Echiura in relation to boltom sediments.
Composition and Distribution of Macrobenthic Invertebrate Fauna III

::IE 0--0 NUMBER ::IE

0 ___ WEIGHT 0 .... 1.0 .... .... "' .... "'0 ::lEO ZIIl W <till

0:: ... ::IE'" O.B

(30 "0

Wo:: ~o:: ""w 0.4 W "' .... 0.6 .... .... ... w :I:W

0::IE ~::IE

o::w /\ Ww wo:: 0.4 ~o::

1Il<t / \ <t ::IE:> 0.2 .... :> :>0 wo z", / \ 0.2 ~'"

0:: 0::

W \ W

"" "" 0 0 0-3.9

ANNUAL

Figure 91

Density and biomass of Echiura in relation to the annual range of

bottom water temperature.

L----4J----I.-....IL-...Jo.1...!l----..,.I"'::.o----,":.!l:--~-2='.0,,----=3'"::.0- .......--:!l...,.0:--.....----,1~0."':'0-----'0

PERCENT ORGANIC CARBON

2

0

lI-

(1)0

ZIII

.... 21k.

u O

.... a: 0.. .... (1)1-

k. .... 0.4 02

a: .... .... a: 1114

2;:) 0.3 ;:)0

Z(I)

a:

....

0.. 0.2

0.1

0

<>- -Q NUMBER

__ WEIGHT

~\ / \

/ \

/ \

/ \

/ \

/ ~, "'-......

Figure 92

Density and biomass of Echiura in relation to sediment organic carbon.

1.6

2

1.2 0

I-

(1)1-

2 0

4 111

a:k. "C)o

~a:.... 1-1-

0.8 :1: .... Cl2

........

~a:4 1-;:)

.... 0 ~(I)

a:

.... 0.4 0..
112 NOAA Technical Report NMFS 140

43°

... 42·

'20 '-~_~, KM

70° ••0

MA IN E

4'· 400

72°

...... _--\

...-....,

....

'''\ ....----- ~-------~------_/ ----, "<D .......... .....

"

EXPLANATION

...

100

1000- __ ~~"",

" 't,

"

',....~,,"'-

EXPLANATION

§

1J.!.llll. 01-10•

1000 - __ ~"'"

" .... t

,

",

,

....

~""'-

12]1-9

NEW

JERSEY

GRAMS

PER SQUARE METER

72°

72°

PRIAPULIDA

Figure 93

Geographic distribution of Priapulida: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

cal areas throughout much of the study area (Fig. 105).

They were most common in the Nova Scotia region, the

periphery of the Gulf of Maine, in deep water south of

Georges Bank, and on the outer shelf south of Cape

Cod, Massachusetts. They were notably absent in the

Georges Bank-Nantucket Shoals area and in deep water

in the western Gulf of Maine. In all but a few locations

their numerical density was less than 91m2 and their

weight averaged less than 1 g/m2•

Three of the six standard geographic areas contained

significant quantities of chitons: Nova Scotia, Gulf of

Maine, and the Southern New England Shelf (Tables 6,

8; Fig. 106). The average number of specimens ranged

from 0.9 to 3.6/m2• Average biomass was small, 0.24 g/m2
Composition and Distribution of Macrobenthic Invertebrate Fauna 113

::IE

e .06 5 «al

~~

Za:: .04 - w .... .... :I:W

"2

~~ .02 IWo

~'"a::

GEORGES SOUTHERN &EOMES SOUTHERN

BANI< NEW ENGLAND SLOPf NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

NO GUL 0 SCOTIA MAINE

0 NUMBER

• WEIGHT -0

-0

0-

-0 5

I 0 VA F F o

:I;

g

.... "'0 Zal

W

:1; .... uO

~~"' .... .... W 0::1: 0.1

a::w

wa::

al«

:1;:::>

=> 0 0.0 Z",

a::

w

Q.

Figure 94

Density and biomass of Priapulida in each of the six geographic

areas.

0-- -0 NUMBER

_ WEIGHT

::E

o

I-

",I-

::Eg

cru.

c>O

02 z cr -UJI- I-UJ

X::E

C>-UJ UJcr 0.1

1-:>

~:UJ

Q.

WATER DEPTH IN METERS

Figure 95

Density and biomass of Priapulida in relation to water depth.

OL.----==-:~----=:7_:______::7:~___;:_:7.;;___='::r1I~-::!:_:_!" IL.-----'O GRAVEL TILL SHELL SAND SAND- SILT- SILT CLAY

::IE

eI-

"'0 Zal

W

::lEu. <:;0

"'a:

3;~

u.'" O::IE

a:w

wa:

al~

::IE:>

~g O,Ha:

W

Q.

o NUMBER

• WEIGHT

- 0.02

-0.01

::IE

o

I-

",I-

::lEo

«al

a:u.

"0

zcr

-w

1-1-

:I: W

"lE

"''''

~a:

1-:>

~~a:

'"

Q.

BOTTOM SEDIMENTS

Figure 96

Density and biomass of Priapulida in relation to bottom sediments.
114 NOAA Technical Report NMFS 140

::E0

...

... "'0 ZID

::E'" '"(jo

"'I>:

CL", "' ... ... ",

0::E

1>:'"

"'I>: 0.1 ID<I

::E:::> :::>0 z",

I>:

'"

CL 0

0----<> NUMBER

----. WEIGHT

00'

L-----"o-=-'·~~.90'-"----;;4~- 7~.9;--BO.~"'11~.9;---;;'2;-,.1;f.5~.9~-';:16"'" ;'~9-;;.9'---;;2"::'0-1_2:P.3"'.9"---.JO ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

::E

o

... ",'" ::EO

<lID

1>: ...

"'0

ZI>:

-",

... ... :1;'"

~::E

"''''

;JI>: <I ... :::> "'0 ;J(J)

I>:

'"

CL

Figure 97

Density and biomass of Priapulida in relation to the annual range

of bottom water temperature.

0.01

1·, 0--0 NUMBER

_ WEIGHT 1

2 2 0.008 0

... ",I- ",I- 2 0

zg <lID

'" a:: ... 2lL "'0. u O ~I>: "'a::

~~ 0.006 '" 1-1-

0.06 :1;'" lL'" ~2 02

"'''' a::", • a:: "'a:: <l ID<l R I-=> 2=> ",0 =>0 / \ .'" z",

/ \ a:: a:: 0.004 0.04 '"

CL

/ \

/ \

/ \ / \

d \ 0.02 / \ 0.002

/ \

/

/

.--~

0 0 0.01 o.~ 1.0 Ul 2.0 3.0 ~.O 10.0

PERCENT ORGANIC CARBON

Figure 98

Density and biomass of Priapulida in relation to sediment organic carbon.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 115

NEW

JERSEY

Lill 1- 49

lillill 50- 499

.500-9453

GRAMS

PER SQUARE

n"

72"

MOLLUSCA

70" .."

Figure 99

Geographic distribution of Mollusca: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

or less. On Georges Bank, Georges Slope, and the Southern New England Slope their density averaged 0.1 to

0.6/m2, and their weight averaged 0.01 g/m2 or less.

Frequency of occurrence was relatively high (1 to

24%) considering the small quantities that were present

(Table 10). This high frequency of occurrence is simply

a reflection of their small size and wide dispersion.

Individuals were especially small in the Georges Slope

area and relatively larger in the Southern New England

Shelf area.

Bathymetric Distribution

Polyplacophorans were present at depths from 16 to

2,840 m. They were more abundant in shallow and

moderate depths than in very deep water (Table 11;

Fig. 107). Average density in depth classes less than 500
116 NOAA Technical Report NMFS 140 _

80

40

60

20

o

20

fIl~

:EO

cal

00

Zo::

-Ill

~~

XIII

~:E

80 III III

.0::

C ~::>

11I0

.fIl

0:: 60 III

Go

40

8ULF OF 8EORGES SOUTHERN GEORGES SOUTHERN MAINE IlANK NEW ENGLAND SLOP£ NEW ENGLAND SHELF SLOPE

GEOGRAPHIC AREA

NDVA

SCOTIA

I

D NUMBER

• WEIGHT

-I

3!10r--

-I

300r-- -I

2!10 f-

-I

ZOO - -

I!lO - -

100 - -

!lO - -

0 .. -

Figure 100

Density and biomass of Mollusca in each of the six geographic areas.

m ranged from 0.4 to 4.0/m2 . Below 500 m the average

density in the various depth classes was 0.2 to 0.5/m2 .

Average biomass of chitons was greater (0.3 to 0.8 gl

m2) at depths less than 500 m than in the deeper water

where their biomass averaged 0.01 g/m2 or less (Table

13; Fig. 107).

Frequency of occurrence wa~ low (1 to 18%) and no

clear relations with depth were evident. The trend indicated a slightly higher rate of occurrence in moderate

(50 to 200 m) and deep (>200 m) water (Table 15).

These results indicate that polyplacophorans are less

abundant but more uniformly distributed in deep water (>500 m) than on the continental shelf and upper

portion of the continental slope.

Relation to Sediments

Polyplacophorans occurred in all sediments exceptshell

(Table 16; Fig. 108). Highest density (3.8/m2) was encountered in sand-silt sediment. Till, gravel, and

silt-clay ranked second to fourth, respectively, and only

very small quantities (0.3/m2) were found in sand

sediments.

The relationship of average biomass to various types

of bottom sediments was quite different from that of

density (Table 18; Fig. 108). Gravel and till yielded the

largest biomasses (0.7 and 0.3 g/m2, respectively),

whereas in all other sediments it averaged <0.07 g/m2.

Frequency of occurrence was highest (11 to 27% of

the samples) in those sediments where chitons were
Composition and Distribution of Macrobenthic Invertebrate Fauna 117

....~---------------------,300

0-- --0 NUMBER

__ WEIGHT

\

\

\

\

\

\

\

\

L_~

"

250

50

WATER DEPTH IN METERS

Figure 101

Density and biomass of Mollusca in relation to water depth.

40

20

60

o

40

20

SAND· SILTSILT CLAY GRAVEL TILL SHELL SAND

0 NU_II

• WEleHT -I

50~ -I

OO~ -I

5C- -I

:oc- -

150- -

100f- -

5Oi- -

0

•

BOTTOM SEDIMENTS

Figure 102

Density and biomass of Mollusca in relation to boltom sediments.
118 NOAA Technical Report NMFS 140

0- --0 NUM8ER

_ WEIGHT

1200 I'

I

I 360 I

I 1000 I

I 320

0 f 0 l- I l- l- (J)l- (J)O 280 ZCIl I ~O

'" <l: CIl

~u.. 800 I Iru.. (30 ClO

"'Ir

0..", I 240 ~lr (J)l- ILl l-l- u..'"

O~ :I: ILl

~~

lr'" ILl ILl "'lr 600 CIl<l: 200 ~lr

~::> <I:

::>0 l-::>

Z(J) ILIa

~(J)

lr lr '" 160 a.. ILl

a..

400

120

/

I 80

200 /

I _ ---0/ 40

0'----.."......:--=----04:-'.7""'.9,...------=8-'-., ':-:.9,...----".,,-2.-!c'5""'.9,...---...,-'6=-·"-:,9:-:.9,...---""20:---·"'23:-:.9,...------'0

RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 103

Density and biomass of Mollusca in relation to the annual range of

bottom water temperature.

most numerous, namely till, gravel, and sand-silt (Table

20). Their occurrence rate was low (3 to 9% of the

samples) in sand and silt-clay.

Relation to Water Temperature

Polyplacophorans were found to inhabit a rather broad

span of annual ranges in water temperature, from 0-

3.9° to 16-19.9°C, inclusive (Table 21, Fig. 109). They

were numerous; average density was 7.3/m2 , where the

water temperature range was greatest. Lower densities

(0.8 to 1.4/m2) were found in the middle and low

temperature range groupings.

Biomass values in relation to water temperature range

rather closely paralleled the trends described above for

density (Table 23; Fig. 109). At the greater temperature

range, 16° to 19.9°C, the biomass averaged 1.3 g/m2,

whereas in the middle and lower range groupings, the

biomass averaged 0.12 g/m2 or less.

Frequency of occurrence was low in all temperature

ranges but was relatively higher (10%) where the fluctuation in temperature was less than 8°C (Table 25). Occurrence rate was relatively moderate (5 to 7%) where the

temperature range was moderate. No chitons were taken

where the temperature range was greater than 20°C.

Relation to Sediment Organic Carbon

Polyplacophorans were rather restricted in their distribution in relation to the quantity of organic carbon in

the sediments, occurring only where the carbon content ranged from 0.01 to 1.99% (Table 26; Fig. 110).

Average density was highest (2.5 1m2) in the lowest

carbon content class (0.01-0.49%), lower still (11m2 )

in the 1 to 1.5% content class; and significantly lower in

the other two classes they occupied.

Average biomass decreased markedly from its highest point (0.25 g/m2) in the lowest content class (0.01-
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 119

1200

~1000 ....

.... eno

ZID

'"

21L

0 0800

'" It:. a.",

en ....

~~

It: '" 600

"'It: IDe 2::)

::)0

zen

ffi 400 a.

200

C>- -0 NUMBER

..--........ WEIGHT

/,

/

/

/

/

/0

/

/

/

./'

/'

/'

rt'

/

/

/

/

/

240

2

180 een .... 2 0

e lD

It: IL C)o

zit:

-",

........

120 :z: '" ~2

"''''

:'11:

e .... ::)

"'0 :'en

It:

60

oL..----'1L--::0~.0:71---=0.'=~----:-I"'=.0~-~I:":.~=----2=-'.'='0--~3'"::.0~----:5:oL:.0=--...........,I~O-::.O-----JO

PERCENT ORGANIC CARBON

Figure 104

Density and biomass of Mollusca in relation to sediment organic carbon.

0.49%) to very small «0.01 g/m2 ) as carbon content

increased to just under 2% (Table 28; Fig. 110).

Frequency ofoccurrence in samples exhibited a trend

similar to that of biomass. Incidence was relatively low

(ranging from 2 to 12%) and diminished with increasing carbon content (Table 30).

Gastropoda-Gastropods formed a moderately common component of the New England benthos. They

were distributed throughout most of the study area, but

because of their generally small size they accounted for

only a small proportion (1.2%) of the total benthic

biomass (Table 3).

These mollusks varied enormously in size, from the

tiny Retusaand Alvania (approximately 2 mm in length)

to large specimens of Neptunea, Colus, Busycon, and

Buccinum (ranging up to 13 em or more). The majority

ofspecimens were between 2 and 30 mm.

Shelled gastropods were predominant in our collections, although some shell-less groups (Nudibranchia

and Aplysiacea) were represented. Nudibranchs were

abundant in a few localized shallow water habitats

but were generally uncommon to rare in the offshore

regions.

Specimens in our samples were usually drab colored,

with various shades or combinations of white, gray, and

brown predominating. The shell-less groups contained

some of the more brightly colored forms. In these

groups light yellow, pink, orange, and rusty-red hues

were common on the dorsal body surface and in the

cerata and tentacles.

Gastropods ofdifferent taxonomic groups obtain their

nourishment by a variety of methods. Feeding types

known to be represented in our collections were herbivores, predacious and nonpredaceous carnivores, and

parasites. Carnivores and scavengers that feed heavily

on bivalve mollusks were the largest and most common

forms encountered. Parasitic species were rare.

Gastropods occurred in 470 samples (44%). Their

density averaged 81m2; biomass averaged 2.2 g/m2

(Table 5).
120 NOAA Technical Report NMFS 140

NEW

JERSEV

1000- .... _ ...... _

,,'

EXPLANATION

o 1-9

III 10- 99

.100-124

,.,

we'")

.......::'i06

1000- __ ...... _

,,'

EXPLANAT ION

EJ <0.1

1l!0.1-0.9

• 1.0-19.3

GRAMS

PER SQUARE METER

...

11'

POLYPLACOPHORA

.,'

70'

M A IN E

... o

~.w

120

.,'

Figure 105

Geographic distribution of Polypl2cophora: A-number ofspecimens per square

meter of bottom; B-biomass in grams per square meter of bottom.

Geographic Distribution

Gastropods were distributed over nearly the entire region (Fig. Ill). Moderately low densities (1 to 49 individuals/m2) were widespread over a large part of the

study area. Medium and high densities generally occurred in limited areas inshore and nearshore.

In most areas where gastropods were found, their

average biomass was less than 1 g/m2• Moderately

high biomasses (1 to 25 g/m2) occurred over rather

large areas both inshore and offshore, whereas large

biomasses (25 to 133 g/m2) occurred in only four

localities.

Gastropods were present in all six of the standard

geographic areas (Tables 6, 8; Fig. 112). Quantities, in

terms of both number of individuals and biomass, were

highest in the four continental shelf areas and lowest in

the Georges Slope and Southern New England Slope

areas. Average density in the shelf areas ranged from 11

to 29 individuals/m2 and average biomass from 0.9 to

4.3 g/m2• In the slope areas the average density was 7 to

8 individuals/m2 and average biomass was less than 0.3

g/m2. Gastropods made up a slightly higher proportion

of the total faunal density in the slope areas than they

did in the continental shelf areas.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 121

Figure 107

Density and biomass of Polyplacophora in relation to water depth.

o

0.8" l3 :lim

$:5 -~0.6 Z er I _ ....

.... :I:::e

0.4 ....

~~.... ::l

.... g 0.2 3: er

....

0.

0-- -<> NUMBER

__ WEIGHT

-------------,1.0

GEORGES SOUTHERIlI GEORGES SOUTHERN

BANt< NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE

GEOGRAPH Ie AREA

WATER DEPTH IN METERS

NOVA Gur..F OF

SCOTIA MAINE

o NUMBER

f-- • WEIGHT -0

3-

-

2f-

-

1-

"- n. II- 0

:lE

., .... zO

.... m:lE .... -0

<.>

.... er 0. .... ., 4

........

0::E

lr .... .... lr m<l

::E 5 2 ::l.,

Z lr

....

0.

Figure 106

Density and biomass of Polyplacophora in each of the six

geographic areas.

Frequency of occurrence of gastropods was

moderately high in all geographic areas. They

were present in 35 to 58% ofthe samples (Table

10). The high frequency of occurrence in the

Georges Slope and Southern New England

Slope areas, in comparison to the low density

and small average biomass in these areas, is

indicative ofsmall-size specimens and of rather

widespread and uniform distribution.

Bathymetric Distribution

Gastropods occurred at water depths ranging

from 3 to 3,310 m. They were far more abundant (641m2 ) in shallow water (0 to 24 m) than

at other depths. Their average density (Table

11; Fig. 113) generally diminished with increasing water depth, except for a slight reversal of

this trend on the upper and middle sections

of the continental slope. Density was roughly

uniform (11 to 24 individuals/m2) between 25

and 1,000 m. In the two deepwater classes the

densities were considerably lower (4 and

11m2 ).

Average biomass also was largest in shallow

water and smallest in deep water (Table 13;

Fig. 113). The average biomass in all depth

classes on the continental shelf was moderate

(1.1 to 4.8 g/m2 ), whereas at all depths greater

than 200 m the average biomass was small (0.15

to 0.29 g/m2 ).

Frequency of occurrence of gastropods was

moderately high (37 to 52%) on the continental shelf and upper slope (Table 15). At midand lower-slope depths occurrence was high

(74 to 77%). On the continental rise their rate

of occurrence diminished to only 30%.

Figure 108

Density and biomass of Polyplacophora in relation to bottom

sediments.

2

o

... "' .... 2 0

.6 <l lD

0: ... <00

ZO:

- ... .4 ",2

......

~lr<l

.2 :;;

0:

...

Cl.

D

BOTTOM SEDIMENTS

GRAVEL TILL SHELL SAND SAND SILT- SILT CLAY

0

o NUMBER

• WEIGHT

-0

I-

-0

I-

-0

o n II r.

2

.... "'0 ZeD

...

2'" ;:;0

... 0: 4 Cl. ... "' .... ......

02

0::'"

"'0:: m <l 2 2::>

::>0

Z",

0::

...

Cl.

Relation to Sediments

Although there was no consistent trend in density of gastropods in relation to sediment particle size, there were several correlative points

of interest (Table 16; Fig. 114). Gastropods

occurred in particularly high densities in shelly

sediments, where their average concentration

was 83 individuals/m2. Presumably these gastropods were predators on the bivalves whose

shells formed the substrate. Densities were also

high (401m 2) on gravel bottoms. In all other

sediment types they occurred in only moderate

densities (9 to 22 individualslm2). The unexpectedly low density of 91m2 in till substrates

indicates that till is more closely allied to siltclay as a gastropod habitat than it is to gravel.

Just the reverse is true for other molluscan

groups.
122 NOAA Technical Report NMFS 140

0-- -0 NUM8ER

:::E _ WEIGHT 1.4

° ::E

ff- °f- (flO 1.2 (flf- Z", ::EO W «'" :::ELL 8 a:: LL <:)0 1.0 C>o wa::

3;~ ~a::W LLW 6 0.8 f-f- o::E IW

c>::E a::w

wa:: Ww 0.6 "'« ~a::

:::E:> 4 « :>0 f-:> Z(fl 0.4 wO

a:: ~(fl

w 2 a:: Q. w 0..--. 0.2 Q.

0 0- 3.9 4-7.9 8-11.9 12-15.9 16,19.9 20-23.9 0

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 109

Density and biomass of Polyplacophora in relation to the annual

range of bottom water temperature.

L..-i--:-0-.l,.0-'-1---0~.Il=----,.JI.'=-O-----:-II::.ll---"""!2~.0~"-----'3:'-.=-O---'---='Il.'=0--"""""IO~.O=-",,-----JO

PERCENT ORGANIC CARBON

2

0

... en'" zi

... 3.0 2'" (30

"'11:

ll;~ 2.11 ......

02

11: ... "'11: IDe( 2.0 2:::) jO

zen

II:

1.11

1.0

0.11

0

~\

/ \

/ \

/ \

/ \

/ \

/ \

/ \

/

I

/

A

/ \

/ \

\

\

\

0---0 NUMBER

_ WEIGHT 0.211

0.20

0.01l

Figure 110

Density and biomass of Polyplacophora in relation to sediment organic carbon.
--------- Composition and Distribution of Macrobenthic Invertebrate Fauna 123

NEW

JERSEY

GRAMS

PER SQUARE METER

1-49

50-99

100-893

12" 10" .."

EXPLANATION

[ill <0.1- 0.9

II 1.0- 24.9

• 25.0 - 132.6

EXPLANATION

!illJ

III

•

GASTROPODA

Figure III

Geographic distribution of Gastropoda: A-numbel' of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

In terms of biomass, the gastropods were most prevalent (3.3 g/m2) in sand and gravel substates (Table 18;

Fig. 114). They occurred in intermediate quantities

(2.1 and 1.6 g/m2) in shell and sand-silt. Low quantities

(0.2 g/m2) were found in till and silt-clay sediments.

Frequency of occurrence was high (83%) only in

samples from shelly sediments (Table 20). In all other

bottom types the occurrence of gastropods was quite

uniform at a moderate (42 to 50%) level.

Relation to Water Temperature

The greatest quantities of gastropods, in terms of both

density and biomass, occurred where the annual range in

temperature exceeded 16°C (Tables 21, 23; Fig. 115).
124 NOAA Technical Report NMFS 140 _

Figure 112

Density and biomass of Gastropoda in each of the six geographic

areas.

o

6

NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

o NUMBER

_ WEIGHT

-

-

of-

-

f-

-

01-

I -IIl 0

0--0 NUMBER

__ WEIGHT

80.----------------------,

o

...

... "'0 ZIIl

'"

~LL

0 0

~:5

"' ... LL'"

o~

ffi 20

IIlCf

~::>

::>0

Zrn

ffi 1 Q.

Both measures indicated a trend ofincreasing

abundance with increased range in temperature. Quantitative values for density were proportionally higher than thos(':Jor biomass, but

the percentage composition of the total fauna

for both measures was moderately low (Tables

22, 24). With the exception of the two broadest

temperature range classes (16°-19.9° and 20°-

23.9°C), where relatively high densities (85

and 47 individuals/m2, respectively) occurred,

gastropod density was fairly uniform at a moderate level of about 10 to 15 individuals/m2.

Gastropod biomass values generally paralleled those of density, but the trend of increased biomass with increased temperature

range was more pronounced. The two largest biomasses (4.96 and 4.09 g/m2) occurred

in the two broadest temperature range

classes. In all other temperature range classes

the biomass was moderate (0.26 to 3.75 gl

m2) and fairly stable.

Frequency of occurrence of these organisms

was moderately high and stable in all temperature range classes (Table 25). They varied only

from 38 to 57%, with no obvious trends.

Figure 113

Density and biomass of Gastropoda in relation to water depth.

::E

o

lI- mO

::E ID

<ILL Ir o 4~zffi

-II-W

:x:::E

3~WWIr

~<I::> 1-0

Wl/)

2 Ir

W

a.

500 1000 2000 4000 a

o

/\

/ \

J \

\

0.......

50 100 200

WATER DEPTH IN METERS

25

'\

\

\

\

\

\

10

of the New England marine benthic fauna. In terms of

number of individuals they provided 10.8% of the total

(Table 3). Weight determinations included shells (see

"Material and Methods"), but even if shells were excluded from weight measurements, the Bivalvia would

still rank first in biomass. In number ofindividuals they

rank third, after Amphipoda and Annelida.

::E 6 o

... ~5WID

~~U

WIr a.",

m I- 40

~~

ffi~ ID Cf ::E:;)

~gIr

W 2 a.

Bivalvia-Bivalvia were inordinately dominant in terms

of wet weight, contributing 44.1 % of the total biomass

Relation to Sediment Organic Carbon

Gastropods displayed a generally bimodal

trend of increasing numerical abundance

and biomass with increasing (to moderate

levels) organic carbon content in the sediments (Tables 26, 28; Fig. 116). Average density ranged from a low of2 individuals/m2 in

areas with no measurable organic carbon to

a high of 451m2 in 1.5 to 1.9% organic carbon. The first peak (201m2 ) occurred in

levels of 0.01 to 0.49% and the second in the

1.5 to 1.9% class. No gastropods were found

where carbon content was greater than 3%.

Biomass values ranged from a low of 0.021

m2 in the absence of organic carbon to a

high of 10.42 g/m2 in the 2.00-2.99% content class. The first peak in moderate biomass (3.8 g/m2 ) occurred at levels of 0.01-

0.49%. Biomass decreased significantly between 0.50 to 1.99% levels and rose dramatically to highest recorded biomass in the highest levels in which they were found.

Frequency ofoccurrence ranged from 20 to 61 % of the

samples (Table 30). Lowest occurrence occurred at both

extremes ofcarbon content, whereas fairly uniform occurrence occurred in the other content classes.
Composition and Distribution of Macrobenthic Invertebrate Fauna 125

::E

o

en~

:1: 0

«al

0:: ... '-'0

ZO::

-W

~~

:z:W

,-,:I:

ww

;,10:: « ~~

~~0::

W

0..

2

o

4

3

SAND· SILT

SILT CLAY GRAVEL TILL SHELL SANO

0 NUMBER

• WEIGHT

OfOf-

-

OfOf- -

Of-

-

o~

Of-

-I

Of0 I I

8

9

7

:I:

o 6

eno

Zal

W

:1: ...

LiO 5

~~en~

... w 0:1:

0:: W 4 WO::

al«

:I:~

~o

Zen

0:: 3 W

0..

BOTTOM SEDIMENTS

Figure 114

Density and biomass of Gastropoda in relation to bottom sediments.

100r------------------------

0----0 NUMBE.R

.--. WEIGHT

::E 80 0

U>O

ZlD

W

::ElL

0 0 60

.... a: 0.. .... U>~

.......

o::E

a:w

wa: 40

al<l:

::E~

~o

zu>

a:

....

0.. 20

/

/

--o---J

~\

I \

I \

I \

I \

/ \

::Ii

o

cn~

~~ 6 a: lL "'0

Za:

- ....

~~

x .... "'::IE

4

<l: ~::>

.... 0 ~cn

a:

....

0..

OL---;:O-.1;-3.-;:9---:;4;-!-~7.-;:9 --'8~-1"1.9;;---'I02 :f'1;;:5.Q9 -"16O:·{;19i<.9"2;r0.-:-.;c23~.9Q--'O

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 115

Density and biomass of Gastropoda in relation to the annual range of

bottom water temperature.
126 NOAA Technical Report NMFS 140

0--0 NUMBER

110.0 - WEIGHT 12.0

P-"

/" 10.0

40.0 / "

/

/ 2

2 0

0 / ... ... B.O ell'" ... 2 0

i 30.0 / cal0: ...

2'" / <:10

0 0

/ ZO: 1&.10: -1&.1 A-I&.I

ell ... 6.0 i l;j l' ... 1&.1 02

02 / 1&.11&.1 0:1&.1 / .0:

~~ 20.0 I~ c

/ ... 2~

~o ~2 Zell / '\'\ / 4.0 0: 0: I '\ / 1&.1

1&.1 AAI \f 10.0 /

/ 2.0

/

/

d

0 0.11 1.0 1.11 2.0 3.0 11.0 10.0 0

PERCENT ORGANIC CARBON

Figure 116

Density and biomass of Gastropoda in relation to sediment organic carbon.

Bivalvia were distributed throughout the entire area

sampled in all water depths, sediment types, and temperature range classes. They were especially plentiful

on the continental shelf where their average density

commonly ranged from about 50 to 500/m2. Biomass

of bivalves averaged 100g/m2 or more over a large

portion of the continental shelf.

A rather wide variety of bivalve species occurs in this

region, and it is estimated that more than 125 speCIes

were present in the samples (Theroux and Wigley, 1983).

Some of the more common families represented were

Astartidae, Veneridae, Mytilidae, and uculanidae.

Size of specimens ranged from roughly 15 cm for

Modiolus modiolus and Placopecten magellanicus to abou t 3

mm for Gemma gemma, Thyasira gouldi, and other small

forms. Large specimens occurred only in shallow and

moderately shallow water, and in medium to coarse

sediments, whereas small specimens were taken at all

depths but mostly in fine-grained sediments.

The color of bivalves in these collections ranged from

white to blackish-brown. The most common colors were

white, light gray, and various shades of brown or olive.

No bright or vividly colored species were represented.

Some of the more colorful forms were Tellina, Thyasira

ovata, and some specimens of Placopecten magellanicus.

In addition to the importance of oysters, soft-shell

clams, quahogs, surf clams, and scallops as food for

man, the bivalves as a group are a major source of

nourishmen t for many marine animals. Mammals, birds,

fishes, and invertebrates all have members that prey

heavily upon bivalves.

Bivalves occurred in 893 samples (83% oftotal). Their

density averaged 163/m2 and biomass averaged 81 g/

m2 (Table 5).

Geographic Distribution

Bivalves were distributed over the entire New England

region (Fig. 117). Average densities per 20-minute unit

area ranged from 1 to nearly 10,000 individuals/m2.

High densities (>500/m2) were most common in

nearshore areas. Moderate densities (50 to 500 individuals/m2) occurred over extensive areas in the Gulf
NEW

JERSEY

Composition and Distribution of Macrobenthic Invertebrate Fauna 127

720 710 100 69°

r---"""-,----';.::..------::"""'---;.:'----..----.------:r--------,,7--------T'-----~

o 120 <M

EXPLANATION

D 1-49

!ill] 50-499

.500-9453

GRAMS

PER SQUARE METER

BIVALVIA

Figure 117

Geographic distribution of Bivalvia: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

of Maine, on the Southern New England Shelf, and

along the continental slope south of Georges Bank and

Nova Scotia.

Bivalves ranked above all other benthic animals in

terms of weight. Average biomass ranged as high as

3,162 g/m2 in some 20-minute unit areas. High biomass

values of bivalves occurred over broad expanses of both

the inshore and offshore sections of the Southern New

England continental shelf, in Great South Channel, on

the southern half of Georges Bank, and in coastal areas

of the Gulf of Maine.

Within the six standard geographic areas bivalves were

most numerous (212 to 276 specimens/m2, respectively)

on the Southern New England shelf and in the Gulf of

Maine (Table 6; Fig. 118). Densities were moderate in all

other areas including Georges Slope and Southern New
128 NOAA Technical Report NMFS 140 _

o

o

o

60

40

180

NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE

GEOGRAPHIC AREA

0 NUMBER

• WEIGHT

-I

Of-

-I

Of-

-I

Of-

-I

Of-

-

Of-

-6

Off- -4

- -2

... ... o 0

BO

40

12

::IE

o

f- 24 f- VlO

Zal

IU

::IE'"- <:;0

5 20 Vlf-

,"-IU

0::IE

a:IU IUa:

al~

::IE::> 16 ::>0

ZVl

a:

IU

a.

32

2B

Figure 118

Density and biomass of Bivalvia in each of the six geographic areas.

England Slope. Lowest abundance (341m2) was on

Georges Bank, but there the average size was greatest.

The biomass of bivalves was exceptionally high (averaged 166 g/m2) on the Southern ew England Shelf

(Table 8; Fig. 118). Biomasses were moderate (30 to 77

g/m2) in ova Scotia, Gulf of Maine, and on Georges

Bank. Average biomass of bivalves was smallest (1 to 2

g/m2) in the Georges Slope and Southern 1 ew England Slope areas.

Frequency of occurrence was high in all areas, especially in the two slope areas where bivalves occurred in

89 and 94% of the samples (Table 10). Georges Bank

had the lowest (64%) rate of occurrence. In all other

areas the rate of occurrence was 80 to 89%.

Bathymetric Distribution

Bivalves were taken at depths from 3 to 3,820 m. They

were common in all depth zones but occurred in greatest density (505 individuals/m2) in shallow water (0 to

24 m) and diminished markedly with increasing water

depth (Table 11; Fig. 119). In the deepest zone samples

(2,000 to 3,999 m) their density was only 261m2. The

decrease in density was quite uniform; the major decrements in the trend were at about 25 m and 200 m.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 129

WATER DEPTH IN METERS

Relation to Water Temperature

Bivalves occurred in significant quantities in all of the

temperature range classes (Tables 21, 23; Fig. 121).

Although the density and biomass exhibited a general

tendency ofincreasing as the temperature range broadened, two major anomalies in this trend were observed

in the two highest range intervals. The 16°-19.9°C class

had a large biomass (334 g/m2 ) and a moderate (252

individuals/m2 ) density, indicating the presence of

larger individuals than in other areas. Conversely, the

200-23.9°C class had a high density (1,195 individualsl

m2) but a small biomass (84 g/m2).

Bivalve biomass was unusually large (106 to 334 g/m2 )

where the temperature range was moderate (8°-19.9°C)

and was comparatively smaller in the low and high

ranges (6.8 and 84 g/m2, respectively).

Frequency of occurrence of bivalves was high and

rather uniform (75 to 93%) among all temperature

range classes (Table 25).

Relation to Sediment Organic Carbon

Bivalves were found in significant quantities in all organic carbon content classes except the highest

(5.00+%). There was a well-defined positive correlation

of increasing density with increasing organic carbon

content (Table 26; Fig. 123). Density of bivalves rose

from moderate levels (641m2 ) in the absence of organic carbon to high levels (1,120/m2 ) where organic

carbon content was between 3 and 5%. Bivalvia is the

only taxonomic group showing such a well-defined trend

in relation to sediment organic carbon. This trend corresponds to that shown for depth distribution, wherein

higher bivalve densities occurred in the shallower estuarine and embayment waters that contained the highest levels of organic carbon.

Although the highest biomasses of bivalves (227 to

801/m2) were found in the higher carbon content

classes (3.00-4.99% and 2.00-2.99%, respectively) the

relationship was not as well-defined as that for density

(Table 28; Fig. 122). Moderately high biomass (1281

m2) also occurred in low carbon levels (0.01-0.49%).

The other carbon content classes contained significantly

lower biomasses.

Frequency ofoccurrence of bivalves in samples in the

various organic carbon content classes was quite high

shelly sediments they formed the exceptionally large

proportion of 74% of the total fauna.

Bivalves were present in a high proportion of the

samples in all sediment types. They occurred in all

samples from shelly bottoms and in a particularly high

percentage (91 to 92%) ofthe samples from fine-grained

sediments (Table 20). Only a moderately high proportion ofthe samples from sand and gravel contained live

specimens of bivalves.

25 50 100 200 500 1000 2000 4000 0

600

0----0 NUMBER 250

__ WEIGHT

500

\ 200

::lil \ ::lil

0 0 \ >- >- ",>- <1)1- 40 \ ~~ ~g

::lil'" \ -0 150"'0 u \ zll: "'Il:

0.'" \ -'"

"'>- 300 >- >-'" ... '" \ J:::lil 0:1: ~", Il:'" \ "'Il: ",Il:

moO: 100~~ :1:5 \ ~'" 200 ~g Il: L~ Il:

'"

0. '" '\ 0.

"'- 50

100 "'- 'o-----~, 'a...... '0

Figure 119

Density and biomass of Bivalvia in relation to water

. depth.

Biomass of bivalves, as with density, was greatest

(252 g/m2 ) in shallow water and decreased sharply with

increasing water depth (Table 13; Fig. 119). On the

continental rise their biomass was only 0.4 g/m2.

Bivalves were present in a high percentage of the

samples from all depth classes. There was, however, a

slightly higher rate (81 to 100%) of occurrence in deep

water (greater than 500 m) than in shallow water, where

the occurrence rate was 75 to 88% (Table 15). Other

characteristics of the bivalve fauna in deep water are

lower average density, lower maximum density, smaller

average biomass, and smaller maximum biomass.

Relation to Sediments

Several clear trends were detected in the correlation

between the quantity of bivalves and the type of bottom

sediments they inhabited (Table 16; Fig. 120). Density

was highest (330/m2) in sediments composed of siltclay and decreased as particle size increased, except in

shell bottoms where the density of bivalves was moderately high (180/m2). Gravels contained the lowest density (391m 2).

An entirely different trend was observed for bivalve

biomass (Table 18; Fig. 120). The largest quantities

(117 to 165 g/m2) were found in sediments of medium

grain size. Smallest quantities (5 to 18 g/m2 ) occurred

in till and silt-clay. Itshould be noted that in four of the

six sediment types, the biomass of bivalves accounted

for over 42% ofthe total benthic biomass (Table 19). In
130 NOAA Technical Report NMFS 140

o

o

o

60

o

:Ii

o

I-

(/)1-

:Ii 0

<tal

00 a:: "- (:10

za::

-LU

1-1-

:z:LU

(:I~

LULU

~a::

<t 1-:::>

LUO

~(/)

a::

LU

Q.

40

20

o SAND- SILTSILT CLAY

GRAVEL TILL SHELL SAND

0 NUMBER

• WEIGHT

-I

-

-I

-

-I

-

-I

-

-8 rl -6

r- -4 -

,- ! -2

I I o •

40

80

240 o

lI- (/)0

Zal

LU

:Ii"- U 0 200

~~(/)1-

,,-LU

o~

160 al<t

~:::>

:::>0

Z(/)

a::

LU

Q. 120

280

320

BOTTOM SEDIMENTS

Figure 120

Density and biomass of Bivalvia in relation to bottom sediments.

and relatively uniform, ranging from 80 to 100%

between the 0.00 and the 2.00-2.99% classes; the

lowest occurrence (75%) was in the 3.00-4.99% class

(Table 30).

Scaphopoda-Scaphopods constituted a minor component of the New England benthic fauna. They accounted

for 0.2% ofthe number of animals and 0.3% of biomass

in the total benthic fauna (Table 3). These samples

included the first representative of the genus Cadulus

known to occur in the Gulf of Maine (Wigley, 1966b).

This same report postulated that the relatively warm

high-salinity bottom water in the larger Gulf of Maine

basins permitted scaphopods from the continental slope

to inhabit deeper sections of the Gulf.

They are burrowing forms that most commonly inhabit medium to fine-grained sediments. The head and

relatively large foot project from the anterior (larger)

aperture of the shell. When buried in the sediment the

an terior end of the shell faces downward and the posterior end is pointed upward. Scaphopods have planktonic eggs and free-swimming trochophoric larvae.

This group of mollusks is taxonomically small. Less

than a score of species and one-half dozen genera are

known from the New England region. Sizes of specimens in our samples ranged from about 1 to 6 cm in

length.

Color of the shells was generally various shades of

white, either uniformly white, or white with partial discolorations of yellow or brown, or occasionally black.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 131

0----<> NUMBER

__ WEIGHT

1200 ?

I

I 360 I

I 1000 I 320 I

I 0 0 l- I l- I- 2BO fill- fIlO I ZCD ~O

W «CD

~"- 800 I a:,,-

GO <.!)o

Wa:

o..w I 240 ~a:

fill- W 1-1- ,,-W :I:W O~ <.!)~

a:w 200 Ww wa: ~a: CD« 600 « ~~ I-~ ~o wo ZUl ~Ul

a: 160 a: w w 0.. 0..

400

120

/

/ 80

200 /

/

/ 40 --0

0'-----::---'::-,:----:-4--'=7:-=.9----:8:-·'-:-11.-:-9--:1-=-2-0-1-=-5.-=-9--:-16=-·~19::-.9:::---:2::-::0:--:!:c2 3=-.9:::----'0

RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 121

Density and biomass of Bivalvia in relation to the annual range of bottom water

temperature.

Scaphopoda occurred in 218 samples (20% of total).

Their density averaged 5.1/m2 and biomass averaged

0.32 g/m2 (Table 5).

Geographic Distribution

Scaphopods were present in moderately low densities (1

to 49 individuals/m2) in large portions ofthe Nova Scotia

and Gulf of Maine regions (Fig. 123). Farther south, with

few exceptions, they were present only along the continental slope and the outer portion of the continental

shelf. One area of relatively high density (50 to 106 individuals/m2) was encountered in the north central part of

the Gulf of Maine. The average biomass of scaphopods

was moderately small in all areas, averaging less than

1 g/m2 in more than 50% of their area of occurrence.

Relatively large average biomasses (1 to 6 g/m2) were

common only in the northern part ofthe study area.

Within the six standard geographic areas scaphopods

were most abundant (10 to 11 individuals/m2 ) in the

Nova Scotia and Gulf of Maine area (Table 6; Fig. 124).

On Georges Slope and the Southern New England

Slope their density was about 51m2. Lowest densities,

approximately 11m2, were found on Georges Bank and

the Southern New England Shelf.

The biomass ofscaphopods was largest (0.7 to 1.0 g/m2)

in the northern areas, smallest (0.02 to 0.08 g/m2) in

the southern areas, and intermediate in intervening

areas (Table 8; Fig. 124).
132 NOAA Technical Report NMFS 140 _

1200

~IOOO lI- 1/)0

ZCD

UJ

211L

(30 800

~f5 1/)1-

IL UJ 021

0:: UJ 600

UJO::

CD<t

21::::>

::::>0

ZI/)

f5 400

Q..

200

0--0 NUMBER

__ WEIGHT

/

/'

/'

/'

/

/

/

/

/

/

/

/

/

/

/

o

/

/

~oo

240

21

IBO

1/)1-

21 0

<t CD

O::IL 0 0

Zo::

-UJ

1-1- 120 J: UJ 021

~~<t 1-::::>

UJO

~I/)

0::

60

o LJ-----::0~.0::-c1---;0:l:.5:--------;-l1.~0----:I!;.5;-------;;2l;;.0---3,f.on----..5~.0;-4---1~0~.0;---'0

PERCENT ORGANIC CARBON

Figure 122

Density and biomass of Bivalvia in relation to sediment organic carbon.

A comparison of average density and average biomass in the various areas indicates that scaphopods are

relatively large in Nova Scotia and on Georges Bank,

and relatively small in the Southern New England Slope

area.

Frequency of occurrence was low (4 to 7% of the

samples) on Georges Bank and the Southern ew England Shelf, and moderate (33 to 46% of the samples)

in all other areas (Table 10).

Bathymetric Distribution

Scaphopods were taken at depths ranging from 19 to

2,329 m. They occurred in low density (0.1 and 0.2

individual/m2) in both shallow water and deep water

but were present in relatively high densities (10 to 14

individuals/m2) in moderately deep water, 100 to 1,000

m (Table 11; Fig. 125). Increases and decreases in

density about their center of abundance were surprisingly consistent and well correlated with changes in

depth.

The biomass of scaphopods (Table 13; Fig. 125) was

small (<1 g/m2) in all depth classes, but changes in

abundance relative to changes in water depth were

similar to that described above for numerical density.

Average biomasses in the shallowest and deepest bathymetric classes were <0.01 g/m2. The average biomass

was largest (0.17 to 0.98 g/m2) in moderately deep

water (l00 to 2,000 m). Because of the small size and

low density of this taxonomic group, it contributed a

rather small share (1.6% or less) of the total benthic

biomass in any depth class (Table 14).

Scaphopods occurred in a moderate share (35 to

59%) of the samples collected from depths between

100 and 2,000 m (Table 15). At depths both shallower

and deeper, they occurred in less than 10% of the

samples and in the two shallowest depth classes, they

were present in only 1% of the samples.

Relation to Sediments

Scaphopods were present in all types of bottom sediments sampled, but were clearly more abundant (11 to

26 individuals/m2) in the shelly and fine-grained sediments than in the coarse types (Table 16; Fig. 126).

Sand and gravel bottoms yielded the lowest (1.8 and

2.0/m2 ) densities. The percentage of the total faunal

density composed of scaphopods was small (2.2% or

less) in all types (Table 17).

Differences in biomass from one type of bottom to

another were less pronounced than they were for density; however, the trend was the same (Table 18; Fig.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 133

72·

EXPLANATION

[ill 1- 49

11IIII 50- 99

• 100-106

72·

EXPLANAT ION

0<0.1- 0.9

11IIII 1.0-6.0

GRAMS

PER SQUARE METER

... SCAPHOPODA.,.

70·

M A IN E

Figure 123

Geographic distribution of Scaphopoda: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

126). Relatively large biomasses were found in shell, till,

and silt-clay sediments and small biomasses occurred in

sand, sand-silt, and gravel. Scaphopods made up only a

small portion (1.2% or less) of the total biomass in all

bottom types (Table 19).

Frequency of occurrence of scaphopods was especially low (10 to 17%) in sand and gravel (Table 20). In

shell, till, and other finer sediments they occurred in a

moderate (30 to 46%) proportion of the samples.

Relation to Water Temperature

Scaphopods were most often found in the more stable

environments where the temperature variations were

small. Density and biomass of these organisms were
134 NOAA Technical Report NMFS 140

1.2

0 NUMBER

:E • WEIGHT :E 0 1.0 0 ... ... ... ",'" "'0 ZID :EO ... <lID

:E'" 12 0.8 0:: ...

0 0 c>o

"'0:: ~o:: 5;~ ... ...... ... ", 0.6 X ... 0:E 8 c>:E

0:: ... ... ... "'0:: ;to:: m<l 0.4 <l :E::::l ... ::::l ::::lo ... 0 Z", 4 ~'" 0:: 0:: ... 0.2 ... ll. ll.

0 NOVA GULF OF 0

SCOTIA MAINE

GEOGRAPHIC AREA

Figure 124

Density and biomass of Scaphopoda in each of the six geographic

areas.

0---0 NUMBER

--. WEIGHT

:E

",'"

~g

~~ 16

"'0:: ll. ... (I) ...

... '" 12 0:E

0::'"

... 0::

m:3 B :Eo

~(I)

0::

...

ll. 4

4000 0

WATER DEPTH IN METERS

Figure 125

Density and biomass of Scaphopoda in relation to water depth.

greatest in the 4°-7.9°C range class and diminished

steadily as temperature range broadened (Tables 21,

23; Fig. 127). None were found in the 20o

-23.9°C class.

Quantitative values for density ranged from 0.2 to 10.3

individuals/m2 and for biomass from <0.01 to 0.92 g/m2.

In terms of both density and biomass they provided generally less than 2% of the total fauna (Tables 22, 24).

Scaphopods occurred in 2 to 37% of the samples in

the temperature range classes in which they were found

(Table 25). Frequency of occurrence was highest where

the temperature range was small, and the frequency

rate decreased as the temperature range expanded.

Relation to Sediment Organic Carbon

Scaphopods occurred where organic carbon content

ranged from 0 to 2.99%. They were most abundant (12

to 13 individuals/m2) in the two moderate carbon content classes 1.50-1.99 and 1.00-1.49%, respectively, and
Composition and Distribution of Macrobenthic Invertebrate Fauna 135

0 0 NUMBER 0 .... .... .... • WEIGHT "'0 "' .... ZIIl 30 ~o

'" .qlll

... er ... 0 0 <:>0

"'er zer a-", 20 -", "' .... ........ ... ",

o~ 0.8 :1:'"

<:>~

er'" ",er "'''' ~er lD.q 10 <l: ~::> 0.4 .... ::> ::>0 ",0 Z'" ~'" er er '" '" a- 0 a- GRAVEL TILL SHELL SILT- 0

CLAY

BOTTOM SEDIMENTS

Figure 126

Density and biomass of Scaphopoda in relation to bottom sediments.

,----------------------,1.0

o

....

.... "'0 z m 10

'" ... 0 0

f5 8 .... o~... '"

er '" 6 "'er lD.q

~::>

::>0

Z'" 4

er

'"

a0--0 NUMBER

__ WEIGHT

0.8

....

.qlll

~~

er ... 0.6 <:> 0

zer

-w

.... .... :1:'"

<:>~

.4 l::!

<l: .... ::>

~g

er 0.2

0'-----,;0::'.3i:o.9,----;;4<='-77<.9'----;;8:7.10<1.9'--"'-2.+.,O~.9;--"716i-.~19~.9"""'~20~.fl3~.9,--------JO

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 127

Density and biomass ofScaphopoda in relation to the annual range

of bottom water temperature.

diminished in abundance where carbon content was both

higher than and below those values (Table 26; Fig. 128).

Biomass peaked (0.48 g/m2) in the 1.00-1.49% carbon content class with somewhat smaller amounts (0.31

g/m2 ) in the 0.01-0.49% class (Table 28; Fig. 128);

biomass in the other classes was fairly uniform ranging

from 0.10 to 0.24 g/m2•

Frequency of occurrence was moderately low in the

2.00-2.99% and 0.01-0.49% content classes (15 and

17%, respectively) but rose to moderate levels (20

to 32%) in samples in the other content classes

(Table 30).

Cephalopoda-Representatives of the class Cephalopoda were not commonly encountered; they accounted for less than 0.1 % of total biomass and density

(Table 3). This apparent rarity is due primarily to sampling bias exemplified by the taxa represented in the

samples. The class Cephalopoda contains some of the

largest, most mobile, and most highly developed marine invertebrates known to science; further, many are

semipelagic or pelagic in habit and are therefore severely undersampled by bottom grabs. The abundance

and distribution of the commercially important squids

inhabiting the study area in the order Decapoda, Illex
136 NOAA Technical Report NMFS 140

0---0 NUMBER

_ WEIGHT 0.5

5.0

0.4

0.1

OL..--no~.o:01 --o.~5;;---,JI.;'O ---;Il;.5---;2~.O;---""'3f.o;;--------:l5.'='0~t--:-:10~.0:----l 0

PERCENT ORGANIC CARBON

Figure 128

Density and biomass of Scaphopoda in relation to sediment organic carbon.

illecebrosus, and Loligo pealei, are well documented (Vovk,

1969; Tibbets, 1977; Lange, 1979, 1982; Wigley, 1982;

Murray and Wigley14 and others) in the fisheries literature but are not represented in our grab samples. Nautili and cuttlefish, two other orders of cephalopods that

are wholly pelagic, are extremely rare in this region,

which leaves only members of the order Octopoda, the

octopi, which are almost wholly benthic in habit, and

the smaller and more sedentary squids available to quantitative grab samplers. Thus the representation of cephalopods in our collections does not accurately characterize

this portion of the molluscan fauna.

Our samples, 5 representing 0.5% of the total, contained a total of 376 individual cephalopods from 2

genera, Octopussp. and Rossia sp. (Table 5). Their mean

density was 0.4/m2, and biomass averaged 0.01 g/m2.

14 Murray, H. E., and R. L. Wigley. 1968. Squid catches on three

cruises of Albatross IV: cruise 63-5, July-August 1963; cruise 63-7,

November-December 1963; cruise 64-1, January-February 1964.

Bur. Comm. Fish, BioI. Lab., Woods Hole, Massachusetts Lab. Ref

68-12,16 p. Unpubl. manuscript.

Geographic Distribution

Cephalopods that were sampled occurred in low to

moderate quantities (I to 40 individuals/m2, and <0.1

to 0.5 g/m2) primarily along the offshore fringes of the

study area at the shelf break and deeper from the

northeast peak of Georges Bank westerly to south of

Nantucket Shoals (Fig. 129). One inshore area, on the

eastern Nova Scotian shelf, also yielded specimens of

cephalopods.

Cephalopods occurred in all standard geographic

areas but the Gulf of Maine in small quantities; mean

densities ranged from <0.1 to only 1 individual/m2, and

mean biomass ranged from <0.01 to 0.01 g/m2 (Tables

6,8; Fig. 130).

Frequency of occurrence in the subareas was also low

with <1 to only 2% of the samples containing specimens

(Table 10).

Balhymetric Distribution

Cephalopods were taken at depths ranging from 114 to

320 m, placing them in two of our standard depth
__________________ Composition and Distr!bution of Macrobenthic Invertebrate Fauna 137

4)0

4)°

10°

M A' N E

....----- _/ --

_ ------'<~L -//--------

-- -...... .... ........ '-

.....-...,

,

" '-

EXPLANATION

[]<0.I-O.5

EXPLANATION

[]I 1- 40

'000 - __ '"' ~""

,

" '(

,

,

,

" ,

"'-"\~

..... ..."

" '----

GRAMS

PER SQUARE METER

NEW

JERSEY

12°

14°

)9° 4,°

CEPHALOPODA

Figure 129

Geographic distribution of Cephalopoda: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

range groupings (Tables 11, 13; Fig. 131). Highest mean

density and biomass (1.5/m2 and 0.03/m2, respectively)

occurred in continental shelf depths between 100 and

199 m, \:hereas at deeper depths, 200-499 m, mean

density and biomass were significantly lower (0.l/m2,

and <0.01 g/m2). Only 1% ofthe samples in each depth

range grouping contained cephalopods (Table 15).

Relation to Sediments

Cephalopods were found only in gravel, sand, and sandsilt sediments. Average density was only <0.l/m2 in

both gravel and sand but was 1.71m2 in sand-silt substrates; average biomass, although low, increased with

decreasing particle size ranging from <0.01 to 0.02 g/m2

(Tables 16, 18; Fig. 132).
138 NOAA Technical Report NMFS 140

o

0.01

NOVA GUl.F OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE lW\IK NEW ENGLAND SLOPE HEW EN8lANO

SHELF SLOPE

GEOGRAPHIC AREA

0_11

• WEIIHT

01-

81-

61-

41-

-

2"-

0 r r r r

o.

L

Figure 130

Density and biomass of Cephalopoda in each of the six geographic

areas.

0- --<J NUMBER

- WEIGHT

:::t

o

fenfzO

w m

:::t LL

U 0 2

wa:

Q. w enfLL w o:::t

a: w wa: I met

:::t::>

::>g

z a:

W

Q.

25 50

WATER DEPTH IN METERS

2000 4000 a

Figure 131

Density and biomass of Cephalopoda in relation to water depth.

Frequency of occurrence was low with only 1% of the

samples in each of the three sediment types containing

specimens (Table 20).

Relation to Water Temperature

Cephalopods were found only where the annual range

in temperature was less than 12°C. Density showed a

positive correlation with increasing temperature range

from <0.1 individuals/m2 in the narrowest (0-3.9°C)

range to 1.1 individuals/m2 in the broadest (8-11.9°C)

range in which they occurred (Table 21; Fig. 133).

Biomass was stable (0.02 g/m2) in the ranges between 4° and 11.9°C, and low «0.01 g/m2) in the

narrowest range (Table 23; Fig. 133).

Less than 1% of the samples in the 0-3.9° and 8-

11.9°C range groupings contained cephalopods, whereas

2% of the samples in the 4-7.9°C grouping yielded

specimens (Table 25).
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 139

0 0 NUMBER 0 ... ... (/) ... 2. -WEIGHT (/) ... zlil SO ... e ID

... a: ... 0 0 "'0

"'a: ~a:

g,~ ... ...... ...... x ... O~ I. 0.02 ~~

a: ... ... ... "'a: ;ta: IDe e ~:> ... :> :>0 o.~ 0.01 "'0 z(/) ;t(/)

a: a: ... ... Q. 0 Q.

0 GRAVEL TILL SHELL SAND SAND' SILTSILT CLAY

BOTTOM SEDIMENTS

Figure 132

Density and biomass of Cephalopoda in relation to bottom sediments.

2: 0 0 ~-<> NUMBER ... ....

.... ___ WEIGHT (/) ... (/)0 ~O Zlll e lll

...

2:lL "' ... "'0 0 0 1.5- "'", za: Q, ... - ... "' .... ... ... ...... /\ :1:'" 02: 1.0- "'~ "' ... / \ ...... LU'" :.a: lD<t / \ c

5 0.5- - 1002 .... :> C--I/ \ ... 0

z'" :.(/)

... '" '"

... Q, 0 0 Q.

0-3.9 4-7.9 B-II.9 12,15.9 16-19.9 20-23.9

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 133

Density and biomass of Cephalopoda in relation to the annual range

of bottom water temperature.

Relation to Sediment Organic Carbon

Cephalopods occurred only where organic carbon content ranged between 0.01 and 0.99%. Highest density (2.1

individuals/m2) and biomass (0.03 g/m2) occurred in

sediments with between 0.5 and 0.99% organic carbon;

significantly lower quantities of both measures occurred

at levels between 0.01 and 0.49% (Tables 26, 28; Fig. 134).

Frequency of occurrence of cephalopods in the

samples was only 1% in each of the content classes into

which they were grouped (Table 30).

Arthropoda

Representatives of the phylum Arthropoda in the New

England region, members of the classes Arachnida,

Pycnogonida, and Crustacea, contribute significantly to

both measures of abundance. Among the nine orders

of Crustacea inhabiting the region (see Table 4),

Amphipoda is the overall dominant taxon in terms of

density, contributing slightly over 43% ofthe total number ofspecimens; this is the same as their ranking in the

Middle Atlantic Bight (Wigley and Theroux, 1981).

The classes Arachnida and Pycnogonida and the nine

orders of class Crustacea will be discussed separately

below. In keeping with the phylogenetic order of treatment, the figures relating to Arthropoda (Figs. 135-

139) are presented here, but the detailed discussion of

the phylum, represented almost wholly by the class

Crustacea, will be presented in the section "Dominant

Components of the Macrobenthos" below.

Pycnogonida-The class Pycnogonida, a relatively small

group of marine arthropods containing about 600
140 NOAA Technical Report NM.FS 1'lU

1 C>--<J NUMBER

.....-. WEIGHT

0.002

2;

o

I-

<Ill2;0

q;lD

a::,,-

"'0

0.006 Z a::

-LU

1-1-

ILU

"'~

LULU

~a::q; 1-::>

0.004 g

a::

LU

Q.

0.008

0.01

/~

/ \

/ \

/ \

/ \

/ \

/ \

/ \

/ \

-J \ o L-~~J,-=::..:Q-:l-::---~---4l----~----l~-;;l;;-'-~-;!;;:--~--;1;::--4--~;:;--_-.Jo 0.01 0.5 1.0 1.5 2.0 3.0 5.0 10.0

PERCENT ORGANIC CARBON

1.0

0

lI- <IlO

ZlD

LU

~"-

0 0 3.0 LUa:: Q. LU <Ill-

,,-LU

O~

a::LU

LUa::

lDq;

~::>

::>0 2.0 Z<Il

a::

LU

Q.

Figure 134

Density and biomass of Cephalopoda in relation to sediment organic carbon.

known species (Barnes, 1974), is commonly referred to

as sea spiders. These organisms are found throughout

the world ocean from the Arctic to the Antarctic and

are considered to be common in occurrence. Most

species prefer cold waters, and although some do swim,

most are bottom dwellers and feed on hydroids, bryozoans, soft corals, anemones, and sponges. The sexes are

separate and the males brood the developing eggs on

specialized oviferous legs. They are for the most part

small, ranging in length from 1 to 10 mm; however, a

few species are much larger with body lengths of more

than 6 cm and leg spans to 75 cm. They are found at

nearly all depths but many species are littoral in habits.

Color tends to be drab, but some pycnogonids are

green and a few deepwater species are red.

Sea spiders were not very abundant in the study area,

contributing less than 0.1 % to total biomass and density (Table 3). In spite of this low abundance, a total of

10 species were identified from our samples (see Table

4). They ranged in depth from 18 to 1,420 m and were

mostly 5 to 15 mm in size, with a few to 25 mm in

length. Specimens ranged in color from flesh colored

to dark tan and light brown.

Pycnogonids occurred in 25 samples (2.3% of total)

which yielded a total of 369 individuals; mean number

was 0.3/m2 and biomass 0.01 g/m2 (Table 5).

Geographic Distribution

The distribution ofpycnogonids was quite patchy in the

New England region, exhibiting inshore and offshore

components (Fig. 140). The inshore component was

composed of isolated patches of low density (I-91m2 )

and low biomass «0.1-0.9 g/m2) on the Nova Scotian

shelf and inshore Gulf of Maine and in Long Island

Sound. Patches of moderate density (l0-49/m2) , but

low biomass, were located at the en trances to the Bay of

Fundy and Long Island Sound.

The offshore component contained patches of low

density and biomass on Browns Bank, the Western Basin of the Gulf of Maine, and the periphery of Georges
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 141

:::E

o

f-

<nf-

:::EO

«lIl

O::u.

t.'o

Zo::

-ILl

f-fIUJ

t.':::E

ILl ILl

~o::

« f-::::J

ILl 0

~<n

0::

UJ

CL

2

6

4

o

8

20

NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

0 NUMBER -

• WEIGHT

I-

-I

lf- -I

f-

-

-

-

le-

.. r-L o

200

:::E 1000 o

ff- <no

ZlIl

UJ

:::Eu.

<:)0 800

UJo::

CLUJ

<nfu. UJ

0:::E

5 600

lIl«

:::E::::J

::::Jo

Zen

0::

UJ

CL 400

1400

1200

Figure 135

Density and biomass of Arthropoda in each of the six geographic areas.

Bank out onto Georges Slope. One patch of moderate

density was located in Great South Channel.

Among the standard geographic areas, sea spiders

were found in all but the Southern New England Slope.

The shallow-water shelf areas, Nova Scotia, Southern

New England Shelf, and Georges Bank, yielded higher

mean densities (0.8-0.3/m2 ) than the deeper Gulf of

Maine and Georges Slope localities (Table 6; Fig. 141).

Mean biomass showed a diminishing trend from

northeast to southwest, ranging from 0.02 to <0.01 g/m2

(Table 8; Fig. 141).

Frequency and of occurrence was highest in Nova

Scotia and on Georges Slope and lower in the other

subareas (Table 10).

the other zones. Mean densities were highest (0.6/m2)

in the two zones between 0 and 49 m depth and diminished as depth increased (Table 11; Fig. 142). Lowest

density (0.l/m2 ) occurred in the deepest depth zone

that they occupied (1,000-1,999 m).

Biomass of pycnogonids was low in all depth zones in

which they occurred, ranging from <0.01 to only 0.03

g/m2 (Table 13; Fig. 142). The highest biomass was in

the 100-199 m depth zone.

Frequency of occurrence of sea spiders in samples

was fairly uniform in most depth zones, ranging from 2

to 4%; however, 9% ofsamples in the 500-999 m zone

contained specimens (Table 15).

Bathymetric Distribution

Pycnogonids were absent in the 200-499 m and 2,000-

3,999 m depth zones but present in small quantities in

Relation to Sediments

Shell is the only sediment type in which pycnogonids

were not found. They were most prevalent (by a wide

margin) in both measures of abundance in gravel, where
142 NOAA Technical Report NMFS 140 _

...----------------------------,40

35

30

10

5

o

I- Cl)b

25 II) <l:

Q::u.

C)O

zQ::

-w

20 I- t:i :I:~

~wwQ::

~<l:

15 I- 5

WCl)

Q::

W

a..

\

\

\

\

\

\

\

\

\

600

400

200

0-- -0 NUMBER

.---. WEIGHT

WATER DEPTH IN METERS

Figure 136

Density and biomass of Arthropoda in relation to water depth.

mean density and biomass were 2.1/m2 and 0.05 g/m2,

respectively. Density and biomass in all other sediment

types were significantly lower, 0.1Im2 and <0.01 g/m2 ,

respectively (Tables 16, 18; Fig. 143).

Frequency of occurrence, as might be expected, was

greatest in gravel (11% of samples), intermediate

(5%) in till, and low (1 %) in the other sediment types

(Table 20).

Relation to Water Temperature

Sea spiders occurred in all temperature range classes.

In terms of mean density there was a wide disparity in

the quantities contained; the broadest temperature

range (20-23.9°C) contained from 2 to 15 times more

individuals (1.5/m2 ) than any other range class, which

had ranges from 0.1 to 0.7/m2 (Table 21; Fig. 144).

Biomass was more evenly distributed among the different temperature range classes. However, the broadest temperature range did not contain the highest mean

biomass (only 0.01 g/m2 ), as it did density. The highest

mean biomass occurred in the 4-7.9°C range class which

contained 0.03 g/m2. Mean biomass in the other temperature range classes was 0.01 g/m2 or less (Table 23;

Fig. 144).

Frequency of occurrence of pycnogonids in the

samples in the various temperature range classes was

rather uniformly low, ranging from 1 to 4%, with the

highest incidence occurring in the two range classes

that yielded the highest density and biomass (Table

25).

Relation to Sediment Organic Carbon

Pycnogonids were restricted to areas of low and moderate levels of organic carbon content, being found where

values were between 0.01 and 1.49%. Mean density

decreased from 0.4 to 0.1/m2 as organic carbon con-
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 143

o

o

4

SAND SILT

SILT CLAY

GRAVEL TILL SHELL SAND

0 NUMBER

f-

-WEIGHT

-2

c-

-I

-

-I

Of-

- 0-

- 0-

0

r II..-

20

1200

1000

40

BOTTOM SEDIMENTS

Figure 137

Density and biomass of Arthropoda in relation to bottom sediments.

tent increased, but mean biomass was fairly uniform,

between 0.01 and <0.01 g/m2 (Tables 26, 28; Fig. 145).

Frequency of occurrence of pycnogonids in samples

also diminished as organic carbon content increased,

ranging from 3 to 1% (Table 30).

Arachnida-One specimen of the class Archnida, order

Acarina, family Halicaridae (water mite) was collected

during the course of this study. This specimen was

taken at station 1130, located at a depth of 86 m on the

northeastern edge of Georges Bank. Sediment at this

location was sand, and the temperature range was between 4° and 7.9°C.

Because of the small size of members of the family

Halicaridae, only a very small proportion of them (the

largest specimens) are components of the macrobenthos.

Adjusted statistics for this group are contained in

Tables 3 and 5 for overall faunal relationship and in

Tables 6 through 30 for relationships to the considered

parameters.

Crustacea-The class Crustacea in the New England

region contains representatives from nine orders, each

of which will be discussed separately below. At least

three of these orders, Amphipoda, Cumacea, and

Cirripedia, rank as dominant components ofthe macro-
144 NOAA Technical Report NMFS 140

1600r-------------------------,

0--0 NUMBER

_ WEIGHT

1400

1200

0

lI- 1000 eno

Zal

LU

~~GO

LUQ:

ll.LU

en I- 800 ~LU

o~

Q:LU

LUQ:

alet

~::;)

::;)0

Zen 600 Q:

LU

ll.

400

200

r\

I \

I \

I \

I \

I \

I \

I \

I \

I

I

I

I

I

I

)

/

/

0-3.9 4-7.9 8-11.9 12-15.9 16-19.9 20-23.9

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

24

20

0

I16 en I-

~O

et lD

a::~

(:)0

~a::LU 1-1-

:I:LU

12 (:)~

LULU

~a::et 1-::;)

LUO

~en

Q:

w

ll.

8

4

Figure 138

Density and biomass of Arthropoda in relation to the annual range of bottom

water temperature.

benthic invertebrate fauna in terms of numerical density. Amphipoda, in particular, contributes an overwhelming majority of individuals (43%) to total faunal

density, 1.5 times as many as the second dominant

taxon, Annelida (28%) (Table 3). Cumacea and

Cirripedia each contributed over 1.5% of total numerical density.

Detailed analysis of this class as a whole will appear

below, along with the other dominant taxa, in the section "Dominant Components of the Macrobenthos."

Figure 146, which shows the distribution of density and

biomass of Crustacea, however is included here so as

not to create disorder in the phylogenetic arrangement

of the figures.

Ostracoda-Because of the small size of most members of this group, only a small proportion were retained by the processing techniques used in this study.

Specimens in our samples were approximately 1 to 2

mm in length. The vast majority of ostracods inhabiting

the sediments of this region, however, were smaller

than this and as such belong to the meiofaunal realm

not sampled in this study. Except for references to the

literature, our comments here pertain only to the very

largest species that occur in the New England region.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 145

C>-~ NUMBER

_ WEIGHT

20

1400

/~ / \

1200 / \ 16

\ 2 \ 2

0 0 ... / \ ... ... / 0'" ~glooo \ 2 0

III / e ID

210. \ 12 11:,,-

i:)0 / Clo

11I11: P \ !II: Do. III III 0 ... BOO I \ ...... 10.11I XIII

02 \ Cl2

11:11I I \ 11I11I 11I11: I .11: IDe \ e

~5 600 I 8 ...

11I0 Z0

I \ .0

II: \ II:

I III

\ Do.

400 I \ / \ I 4

\ I \ 200 I \ I \

0 0 0.01 0.11 1.0 1.11 2.0 5.0 11.0 10.0

PERCENT ORGANIC CARBON

Figure 139

Density and biomass of Arthropoda in relation to sediment organic carbon.

Other studies (Wigley and McIntyre, 1964) revealed

that ostracods, including specimens as small as 75 microns or less, are not abundant in the offshore Southern New England region. Their average density there

was only slightly more than 1 individual/m2.

Ostracods were taken at only five stations (0.5% of

total) situated in diverse localities, all in offshore waters

(Fig. 147). Two of the stations were situated on the

Nova Scotia shelf, and one station was situated at each

of the following locations: the Gulf of Maine, the continental slope south of Georges Bank, and the continental

rise east of ewJersey. Density ofthese ostracods averaged

<0. 11m2 and their biomass <0.01 g/m2 (Table 5).

Water depths at which ostracods were found ranged

from 61 to 2,682 m. Their average density was slightly

higher at depths below 200 m than on the continental

shelf (Tables 11, 13).

Ostracods occurred in three types of bottom sediments: gravel, sand-silt, and silt-clay. Their density was

about equal in each type (Tables 16, 18).

Members of this group were found only in areas

where the temperature range was below 8°C. Although

their density in all areas was low, it was slightly higher

where the temperature range was less than 4°C than in

areas where slightly higher (4-7.9°C) ranges prevailed

(Tables 21, 23).

Ostracods occurred where sediment organic carbon

content levels ranged between 0.01 and 0.99%. Densities were somewhat greater at the higher levels than at

the lower ones (Tables 26, 28).

Cirripedia-Barnacles were generally sparse and, except in a few local areas, made up a small proportion of

the total benthic fauna. In some favorable habitats,

such as rocky areas in shallow coastal waters and on

offshore banks subjected to relatively strong water currents, barnacles were common to very abundant (Table

3). Densities of nearly 8,000 individuals/m2 and biomasses of over 1,000 g/m2 were encountered.

Members from two suborders, Balanomorpha (rock

barnacles) and Lepadomorpha (stalked barnacles), were
146 NOAA Technical Report NMFS 140

NEW

JERSEY

.2'

10·

M A I N E

M A IN E

11·

...-....,

~---- !./j,:.;. --------//-------

...-....,

.---

B

100

100

1000 - __ ~_""

" 'l

,

"

" ,,

' .........,

-'--

EXPLANATION

00··;·;< I 9 :;:;::: - rm 10-49

GRAMS

PER SQUARE METER

12·

1000- __ ...... -.. .... ,

"""'" 72° ..... l

,

", EXPLANATION "

[ill <0.1 - 0.9 \'--,-'--

PYCNOGONIDA

Figure 140

Geographic distribution of Pycnogonida: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

present in the collections, but those from the latter

group were uncommon. The genus Balanus was overwhelmingly the dominant form, of which three species

were common.

Rock barnacles were usually 0.5 to 1.5 cm in height

and diameter; however, some newly settled specimens

as small as 1 mm and a few specimens greater than 5 cm

in length and diameter were collected. Stalked barnacles had a more restricted size range; they averaged

0.5 cm in length, with extremes of about 0.25 to 1 cm.

Rock barnacles were most commonly found attached

to rocks, mollusk shells, and shells of other barnacles. A
Composition and Distribution of Macrobenthic Invertebrate Fauna 147

:::E

0

"'0 ZID

:::ElL '"

<:;0

"'It:

... ", 0.8 "'~

IL'" 0:::E

It:'"

"'It: IDel 0.4 :::E:>

:>0 Z",

It:

'"... 0 NOVA

SCOTIA

o NUMBER

• WEIGHT

::I

o

~bel ID

It:IL

"'0

Zit:

-",

~~

x'" ",:::E

0.02 '" '" ~~

~:>

.01 ~g

It:

... '"

GEOGRAPHIC AREA

Figure 141

Density and biomass of Pycnogonida in each of the six geographic areas.

.....--------------------~0.04

o---{) NUMBER

_ WEIGHT

:::E

0.03 (/)0

:::EID

el lL :50

211:: -'" 0.02~ ttl :I:::IE

ILl ~'"It: ;tel:> ~o

ILl(/)

-001 ;t II::

ILl

0..

WATER DEPTH IN METERS

Figure 142

Density and biomass of Pycnogonida in relation to water depth.

2.5i,---------------------_--.

0 NUMBER

• WEIGHT I- ",I-

:1 0 e lll

a: IL 0.06 "0

~a:III ~I-

.04 :Z:III

":I

Will

_a:e ~::l

0.02 1110

-'"a:

III

A.

0

BOTTOM SEDIMENTS

O'L---=GR!cA,JIV!,EL-~,----=-=-~-----!-,JIII-......,L--~--I.--L--....I

Figure 143

Density and biomass of Pycnogonida in relation to bottom sediments.
148 NOAA Technical Report NMFS 140

20

0

... ",>-

20 0 <[ID

a:lL

"'0

~a:w ...... 0.03 XW

"'::I

Ww o.oz ;Ja:<[

>-::> wO 0.01 ;J'"

a:

w

0 Q.

I

/

/

/

0--0 NUMBER

_ WEIGHT

oL---"o.:.1;3-0. 9--4;;".J,1.9;;-----;;S-+.IT;1.g;--""7,""Z.+';15;-;;.9'---7;:16'-!.1~9.;:;-- g-;;z=-o.-!:z=- 3.9=----.J

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

20

o

...

... "'0 ZID

W

20lL

<:;0

wa: 1.5

g,~

lL W

020 1.0 a:wwa:

ID<[

::I::>

g 0.5

a:

w

Q.

Figure 144

Density and biomass of Pycnogonida in relation to the annual

range of bottom water temperature.

-0.025

- 0.050

0--() NUMBER

..- WEIGHT

0,

/ .............

/ "<\

/ \

/ \

/ \

/ \

/ \

/

/"" oL--,j~~=:::tt::=::r;::~-~::;t;==~:::=::~"':"'~-t--::~I~"i----:::1=-I~ ..t--::~I~..t--:-:L"......._--.Jo 0.01 0.5 1.0 1.5 2.0 5.0 5.0 10.0

PERCENT ORGANIC CARBON

Figure 145

Density and biomass of Pycnogonida in relation La sediment organic carbon.

small proportion occurred on stalks of ascidians, on

carapaces of decapod crustaceans, and on a variety of

other hard to moderately hard objects. Stalked barnacles

were attached to similar substrates but were relatively

more common on crustaceans, hydroids, and corals.

The majority of specimens were chalky white. Rock

barnacles from inshore localities sometimes had a film

of yellowish or greenish algae covering their plates, and

in some ridged species a brownish layer occurred in the

grooves of the shelL Very large specimens were commonly light yellow and white, whereas some of the very

small ones were pale gray. Stalked barnacles generally

had white shells on both the capitulum and peduncle,

but a few specimens had delicate shades of rose on the

larger plates. The color of the peduncle ranged from

yellow to grayish brown.

Cirripedia occurred in 41 samples (4% of total).

Their density averaged 2L8/m2. Their biomass averaged 3.39 g/m2 (Table 5).

Geographic Distribution

Barnacles were found at scattered locations along the

entire length of the region's continental shelf (Fig.

148). Coastal regions, shoals, and banks were the most
Composition and Distribution of Macrobenthic Invertebrate Fauna 149

NEW

JERSEY

f9o

m 100-999

.1000-9994

GRAMS

PER SQUARE METER

".

CRUSTACEA

". 70· 69·

o

~K.

120

4'·

..

Figure 146

Geographic distribution ofCrustacea: A-number ofspecimens per square meter

of bottom; B-biomass in grams per square meter of bottom.

commonly inhabited areas. They were absent from almost all of the continental slope and from much of the

central Gulf of Maine. Densities of from 1 to 49 individuals/m2 and biomasses of less than 10 g/m2 were

most prevalent, but large quantities (100-5,000 individuals and 50-612 g/m2 ) were not unusual in the

coastal and nearshore localities.

Barnacles were present in four of the six standard

geographic areas; they were absent from Georges Slope

and the Southern ew England Slope (Table 6;

Fig. 149). The two areas where they were especially

abundant were the Southern ew England Shelf (521

m2

) and Nova Scotia (361m2). Densities in the Gulf of

Maine and on Georges Bank were only 3 and 61m2,

respectively.

Biomass in ova Scotia was larger (13 g/m2 ) than on

the Southern ew England Shelf (7 g/m2) even though

densities were higher in the latter area (Table 8; Fig.
150 NOAA Technical Report NMFS 140

13°

12°

12°

NEW

JERSEY

100

EXPLANATION

[] 1-9

INDIVIDUALS

PER SQUARE METER

NEW

JERSEY

EXPLANAT ION

[ill <0.1

GRAMS

PER SQUARE METER

39°

12°

.....-....,

'-,'-

OSTRACODA

1,°

4,0

10°

MA I NE

M A IN E

4.°

69°

.... "-

"

Figure 147

Geographic distribution of Ostracoda: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

149). Thus the average size of individual barnacles was

larger (0.36 g) in Nova Scotia than on the Southern

New England Shelf (0.13 g). The biomass of barnacles

in the Gulf of Maine and on Georges Bank was less than

0.5 g/m2.

Frequency of occurrence was highest (15% of the

samples) in the Nova Scotia area and substantially lower

(2 to 4%) in the remaining shelf areas. They were

absent in the two slope areas (Table 10).
___________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 151

NEW

JERSEY

.g.

'2·

M A IN E

70·

.,.

....---- -'" --- --- ------

B

72·

....-....\ ..

..... ""........ -/"'------- -------------

0.1- 9.9

10.0-49.9

50.0 -612.0

1-49

50- 99

100- 5074

1000 - __ ~"""

,

't,

\ ,

,

' ....,

......._,

'''-

'00

'00

...

NEW

JERSEY

EXPLANATION

IITJ

II

•

EXPLANATION

lZJ

IIi1

•

INDIVIDUALS

PER SQUARE METER

GRAMS

PER SQUARE METER

72·

CIRRIPEDIA

Figure 148

Geographic distribution of Cirripedia: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

Bathymetric Distribution

Barnacles were limited to the rather shallow depth range

of8 to 329 m. They were substantially more plentiful (214

individuals/m2) in shallow water and diminished sharply

in abundance with increasing water depth (Table 11; Fig.

150). Densities on the outer continental shelf and upper

slope were 2.3 and 0.7 individuals/m2, respectively.

Biomass of barnacles in relation to bathymetric distribution was similar to that of numerical density. Biomass was largest (27 g/m2) in shallow water and de-
152 NOAA Technical Report NMFS 140

o

Ien I-

~o

<l:lD

a::~

ClO

~a::IoJ 1-1-

:I; ILl Cl~

ILlIoJ

~a::<l: I-::J

ILl 0 ~en

a::

ILl

ll.

2

14

2

o

8

4

6

o GEORGES SOUTHERN

SLOPE NEW ENGLAND

SLOPE

GE:lRGES SOUTHERN

BANK NEW ENGLAND

SHELF

GEOGRAPHIC AREA

GULF OF

MAINE

NOVA

SCOTIA

0 NUMBER

- • WEIGHT

-I

r-

-I

- r-

- -

- -

-

r. n. o

10

50

o

lI- eno

ZlD

40

(jO

ILIa::

Il.ILI

en I-

~ILI

o 30

a:: ILl

ILIa::

lD<l:

~::J

::JO

zen

ffi 20 ll.

60

Figure 149

Density and biomass of Cirripedia in each of the six geographic areas.

creased with increasing depth (to 0.1 g/m2 at 100-199

m) (Table 13; Fig. 150). An exception to this trend

occurred in the 200-499 m depth class. In this deepwater

zone the density was low but the biomass was much

larger (2.5 gm2 ) than the general trend would have

indicated. This relatively large value may have been due

to the presence of Balanus hameri in this depth class.

This species is exceptionally large and occurs in moderately deep water.

Barnacles occurred in only a small proportion of the

samples, but they were much more common (13%) in

the shallow depth class (0-24 m) than in the other

classes (3 to 4%). None were present in depths greater

than 500 m (Table 15).

Relation to Sediments

Barnacles were found in all sediment types of the New

England region that were sampled. Their average density was, surprisingly, highest (56 individuals/m2) in

sand-silt sediments (Table 16; Fig. 151). Moderate densities (29 and 16 individuals/m2 ) occurred on gravel

and sand bottoms. The other sediment types yielded

low «51m2 ) densities.

The biomass of barnacles was largest (11 g/m2) on

gravel bottoms and was moderate (4.4 and 2.4 g/m2 ) in

sand-silt and sand. Low quantities «0.5 g/m2) prevailed in the other sediments (Table 18; Fig. 151).

The unexpectedly high density of cirripedes in sandsilt bottoms resulted from high concentrations ofsmall

specimens (average weight <0.1 g) in a small proportion (3%) of the samples. Small barnacles densely colonize occasional mollusk shells and other firm substrates,

often of biogenic origin, but rarely are they able to

attain large size in these habitats. Conversely, on gravel

bottoms the average size ofindividual barnacles was 0.3

to 0.4 g.

Frequency of occurrence of barnacles was low in all

types of bottom sediments. Shell and gravel ranked
Composition and Distribution of Macrobenthic Invertebrate Fauna 153

25(),-~--------------------,

l>- - -<> NUMBER

_ WEIGHT

25

WATER DEPTH IN METERS

30

25f/)~g

«LL :;0

20 z

_.... a:

.... :I:~

1513~«::::l ~o

.... f/)

10 a:

....

lL

5

000 4000 0

Figure 150

Density and biomass of Cirripedia in relation to water depth.

r-----------------------,12

o NUMBER

• WEIGHT

20f- Q.

a:

'".... ... '" 0:IE

a:w",a:

CD .. :IE:>

:>0 z",

6( f---

30f--

- 10

-B

- 6

-4

:IE

o

.... "' .... :lEO

.. CD

a: ... "'0

za:

-w

........

0::'"

"':IE

"''''

~a:.. ....:>

wo

~'"a:

w

Q.

10f- -2

OL---,-L~--~rL~-___:~ GRAVEL TILL SHELLr.'":-_~~___::':_:'~-~~----'o SAND SAND- SILTSILT CLAY

BOTTOM SEOIMENTS

Figure 151

Density and biomass ofCirripedia in relation to bottom sediments.

highest with an incidence rate of 13 to 17%. The

rate was moderately high in till, but in all other types

they were present in less than 3% of the samples

(Table 20).

Relation to Water Temperature

Barnacles occurred throughout the entire temperature

range spectrum of the New England region (Table 21;

Fig. 152). There was a pronounced increase in density as
154 NOAA Technical Report NMFS 140 _

0--0 NUMBER

__ WEIGHT

20

200 18

180 /

/ 16

0 / I- 160 ( 0 I- 14 I- <1)0 <1)1-

W

Zal ~O

.q:a1 ~LL ! GO 140 Q: LL

wQ: 12 (!)O

el- w

I <1)1- ~Q:W LLW 120 1-1- O~ ! xw

Q:W 10 ~~

WQ: I Ww al.q: 100 ~Q: 2::::l .q: :::lo I-:::l Z<I) I wo 8 ~<I) Q:

W 80 Q: el- I W

elI 6

60

! 40 4

I 20

~--J 2

0 --- 0 0- 3.9 4-7.9 8-11.9 12-15.9 16-19.9 20-23.9

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 152

Density and biomass of Cirripedia in relation to the annual range of bottom

water temperature.

the temperature range broadened. This trend was very

pronounced. Their density was 0.3/m2 where the temperature range was nil or small. Their density steadily increased

to 196/m2 where the temperature range was greatest.

Biomass similarly showed a marked increase in relation to increased temperature range. The range in

average biomass was from 0.05 to 20.32 g/m2, the extreme values occurring in the two extreme temperature

range classes (Table 23; Fig. 152).

Frequency of occurrence of cirripedes was rather low

(2 to 5% of the samples) in all temperature range

classes except the highest (200-23.9°C), where a moderate (18%) incidence rate was obtained (Table 25).

Relation to Sediment Organic Carbon

There was a distinct bimodal relationship of cirripede

abundance with regard to organic carbon content of

the sediment. This was especially apparent in terms of

density. Moderate densities (19 and 39 individuals/m2 )

occurred in the two carbon content range classes between 0.01 and 0.99%; none occurred in content classes

between 1.00 and 1.99%; but density increased dramatically to 613/m2 where carbon content was from

2.00 to 2.99%, and was moderately high (831m2 ) in carbon contents between 3 and 5% (Table 26; Fig. 153).

Biomass displayed a similar trend but not as dramatically. Moderate biomass (6.5 and 4.1 g/m2) occurred in

the two range classes between 0.01 and 0.99%, with

highest biomass (14.3 g/m2) in the 2.00 to 2.99% class

(Table 28; Fig. 153). Lowest biomass was in the highest

organic content class.

Frequency of occurrence of barnacles was moderate

(3 and 6%) in the lower level organic carbon content

classes but was moderately high (15 and 25%) in the
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 155

800

0---0 NUIiBER

-. WEigHT

18.0

:IE

o 12.0 ::

"0

~ID "' COO... z'" -",

1-1-

:>:'" 8.0 ~:IE

"'''' .'"C I-~

;~

'" 4.0 '"...

O'-----~:='>::::....:'f=_-_O---~-=-4___;";_---_;;';;__-~-~;;=-'*';__------'O 0.& 1.0 1.8 2.0 3.0 10.0

PERCENT ORGANIC CARBON

Figure 153

Density and biomass of Cirripedia in relation to sediment organic carbon.

higher level classes, reflecting the trends established by

density and biomass (Table 30).

Copepoda-Only four ofoursamples contained specimens of Copepoda, representing only 0.4% of the total

samples. The small size of members of this group in

relation to the sampling methods used in this study led

to incomplete sampling and the attendant extremely

conservative abundance estimates.

Copepoda represented less than 0.1 % of the total

macrofaunal biomass and density (Table 3). A total of

26 specimens was obtained, yielding a mean density of

<0.1/m2 and a mean biomass of <0.01 g/m2 (Table 5).

Samples containing copepods were located in the

Southern New England Shelf and Slope subareas. Depth

ranges occupied were 50-99 m, 200-499 m and 500-

599 m. Copepods were present in three sediment types

(sand, sand-silt, and silt-clay), in two temperature range

classes (0-3.9° and 12-15.9°C), and in the three organic carbon content classes between 0.01 and 1.49%.

Values of copepod biomass and density for each environmental parameter considered in this report may be

found in Tables 6--30.

Cumacea-Cumaceans are marine peracarid crustaceans that were widely distributed and well represented

in New England waters (Theroux and SchmidtGengenbach7). Twenty-,three species in 13 genera belonging to 5 families were identified in our samples

from the New England region. Cumaceans were among

the subdominant taxa in terms of density, providing

1.7% of the total number of specimens, but owing to

their small size, were much less important in terms of

biomass, contributing only 0.1 % of the total (Table 3).

Cumaceans in our collections ranged from 7 to 15

mm in length; most were between 8 and 12 mm long.

Color of our specimens was mostly drab olive to olive

brown with a few lighter in color and mottled by dark spots.

The majority of cumaceans are bottom dwellers that

were found buried in sand and mud, filter feeding or

browsing organic matter from sand grains. Many exhibit diel excursion to the surface or into the water

column where they swarm at night.

Cumaceans occurred in 390 samples (36% of the

total), yielding a total of more than 27,500 specimens

(Table 5). Their mean density was 261m2 , and mean

biomass was 0.11 g/m2•

Geographic Distribution

Although found throughout the study area, cumaceans

showed some interesting patterns of absence, especially

in the Gulf of Maine. These distributional patterns

reflect the rather restricted sediment particle size preferences of cumaceans. They tend to favor sediments of

medium to medium-fine particle sizes that are most

prevalent in the sand fractions, and shun the coarser

(gravels, tills, shelly fractions) and finer (sandy silts,

silts and clays) fractions. The Nova Scotian shelf and

GulfofMaine each contain extensive deep basins floored

with fine muds, as well as shallower banks paved with
156 NOAA Technical Report NMFS 140

69°

oz·

70°

.9° 0'·

EXPLANATION

~<~~=~:~

EXPLANATION

='::~~8

GRAMS

PER SQUARE METER

CUMACEA

Figure 154

Geographic distribution of Cumacea: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

cobbles, gravels, and coarse sands; the central part of

Georges Bank is largely made up of coarse shifting

sands; and Nantucket, Vineyard, and Long Island Sounds

contain large expanses of very fine muds and silts. Most

of these areas were devoid of cumaceans (Fig. 154).

Average densities ranged from 1 to 618 individualsl

m2. The majority of the region contained moderate

densities (I-491m2) with medium size patches ofintermediate (50-991m2) density at continental shelf depths,

and high (100-618/m2 ) density along the southern

edge of Georges Bank and along the coasts of Maine,

Massachusetts, and Rhode Island.

Average biomass was low «0.1-0.9 g/m2 ) over most

of their range with only small patches of moderately
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 157

(II'"

2 0 .. CD 0.2 a: "- ClO

a: z'" -I1-'" I2

",a: ~'"

0.1 ~~

1-0

"'(II

~a:W

Q.

0-- --<l NUMBER

__ WEIGHT

,-.. -----,0.3

0 ~>-:!::-----;&---.l;;______;;!;;;_~~~£;~~~~-~O';;!;;,Jo 0 10 25 50 100 200 500 1000 2000 4000

WATER DEPTH IN METERS

0.08 g/m2 in waters 100 m and less in depth and was

lower still at deeper (>100-1,999 m) sites, ranging from

0.04 to 0.01 g/m2 with increasing depth. The deepest

zone contained a mean biomass of 0.05 g/m2 (Table

13; Fig. 156).

Cumaceans were well represented in the samples in

each depth class. Four depth classes, the th:-ee between

25 and 200 m and the 500-999 m class, each yielded

specimens in over 30% of the samples (range: 30-

Figure 156

Density and biomass of Cumacea in relation to water depth.

T

0 NUMBER

l" • WEIGHT

0 0

40

f- f- enf- f-

"'0 r 0.16 ~o

2lD <l lD

W

3+ a: .... .... "'0 <:>0

wa: a:

e=;~ 0.12 W

f-f-

.... W IW

o~ "'~

a:w Ww

wa: 20 ~a:

lD<l

l" <I

~:;) f-:;)

:;)0 wo 2", ~en

a: a: w w Cl. 10 Cl.

0.04

I

0 NOvA GULF OF 0

SCOTIA MAINE

GEOGRAPHIC AREA

Bathymetric Distribution

Cumaceans were obtained at depths from 4 to

2,840 m. They were most plentiful, however, at

depths shallower than 100 m. The three depth

zones between 0 and 100 m contained significantly higher mean densities than the deeper zones,

ranging from 50 to 331m2 as depth decreased, whereas

in the deeper (>100 m) ones they ranged from 7 to 0.7I

m2 as depth increased to 1,999 m. In the deepest zone

(2,00-3,999 m) mean density was 21m2 (Table 11;

Fig. 156).

The trend for biomass in relation to depth was similar to that for density but was from one to three orders

of magnitude lower. Mean biomass ranged from 0.26 to

Figure 155

Density and biomass of Cumacea in each of the six geographic areas.

low biomass (1-4 g/m2 ) along the shelf break on

Georges Bank and the eastern shore ofCape Cod.

Among the standard geographic areas, Georges

Bank and the Southern New England Shelfyielded

the highest mean densities (45 and 371m2, respectively) and biomass (0.20 and 0.17 g/m2, respectively) .

Lower densities and biomasses occurred in the

Gulf of Maine and Nova Scotia shelf, and lowest

values for both measures occurred in the two slope

areas (Tables 6, 8; Fig. 155).

Frequency of occurrence was moderately high

in all geographic areas with from 19 to 49% of the

samples containing specimens of cumac.eans

(Table 10).
158 NOAA Technical Report NMFS 140 _

o

0.15

0.05

0.10

0.20

GRAVEL TILL SHELL SAND SAND· SILT- SILT CLAY

0

0 NUMBER

• WEIGHT

01- -

0- -

01- -

01- -

0 r-. I II BOTTOM SEDI MENTS

Figure 157

Density and biomass of Cumacea in relation to bottom sediments.

Relation to Water Temperatu-re

Both measures of cumacean abundance showed a bimodal trend in relation to annual range in bottom

water temperature. The greatest numerical abundance

(561m2) and biomass (0.26 g/m2) occurred where temperature range was moderate (8-11.9°C), followed by

56%), while the other depth classes had frequencies over 15% (Table 15).

Relation to Sediments

Cumaceans occurred in all sediments found

in the region but in significantly different

proportions. Sand, sand-silt, and gravel contained the greatest numbers of individuals,

with mean densities of 45, 18, and 111m2 ,

respectively; shell and silt-clay sediments

yielded moderate mean densities (8 and 71

m2, respectively), with smallest (11m 2 )

amounts in till substrates (Table 16; Fig. 157).

The trend for biomass was essentially similar to that of density but at much reduced

levels (Table 18; Fig. 157). Sand, shell, and

sand-silt contained mean biomasses of 0.20,

0.08, and 0.06 g/m2 , respectively, while the

values for gravel, till, and silt-clay were 0.06,

0.01, and 0.03 g/m2, respectively.

The frequency of occurrence of cumaceans ranged from 50 to 26%, in samples

from sand, sand-silt, and gravel and was between 14 and 22% in the other sediment

types (Table 20).

0.28

0- --0 NUMBER

__ WEIGHT 0.24

60 ::E 0 0 .... .... .... 0.20 "' .... "'0 ::EO ZIIl <till '" 50 :lE'" 0: ...

13 0 "'0

"'0:

)\\

0.16 =:0:

3;~ 40 ........ '" ... ", / \ :>0'" 0:lE ",:lE

0:'" I \ 0.12

"'''' "'0: 30 ~o:

1Il<t / \ <t :lE::> >-::> ::>0 I \ ",0 Z", 0.08 ~'"

0: 20 0:

'"

Q. '"

Q.

0.04 10

0 0 0-3.9 4-7.9 8-11.9 12-15.9 16·19.9 20-23.9

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 158

Density and biomass of Cumacea in relation to the annual range of

bottom water temperature.

another peak in areas experiencing a somewhat higher

(16-19.9°C) range where high density (441m2) but

considerably lower (0.09 g/m2) biomass occurred

(Tables 21, 23; Fig. 158). Density was also relatively

high in the two other high temperature range classes

(12-15.9° and >20°C) but fell off considerably in the

more stable temperature regimes with ranges between
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 159

0.211

The highest frequency of occurrence, 44%, was in

the 0.01-0.49% content class followed by the 2.00-

2.99% class, in which 31 % of the samples yielded specimens of cumaceans. Twelve to 29% of the samples in

the other content classes in which they occurred provided specimens (Table 30).

Tanaidacea-Tanaidaceans are peracaridan crustaceans generally considered to be transitory forms between the Mysidacea-Cumacea and the 1sopoda. The

order is made up of four families, but only two, the

Paratanaidae and Neotanaidae, were represented in

our collections.

Although our specimens were collected in relatively

deep water, tanaidaceans can be found from between

tide marks to ultra-abyssal depths exceeding 9,000 m.

They inhabit burrows, tubes, or rock crevices. Some

species are known to inhabit sponges; others may be

found on algae or the shells of bivalve mollusks, barnacles, and other animal groups.

Tanaidaceans attain lengths of 2 cm or more, but

most specimens in our samples were generally small in

size, usually less than 0.5 cm long.

0---0 NUMlER

--.. WEIGHT

1211.0

100.0 0.20

:. :. 0

e 1Il~

:.0

;!i e CII

... 70,0 O.III~

(j0 !cr: 1&.Icr: 1&.1

lLl&.I ~~

1Il~ :%:1&.1

... 1&.1 !!!:. 02 1&.11&.1

11I:1&.1 .11I:

• 110,0

e 0.10:;;

~o • III

Z(I) III:

III: 1&.1

1&.1 lL

lL

211.0 0.011

oand 7.9°C. Biomass values, although at a much lower

level, were similarly affected.

Frequency of occurrence of cumaceans in samples in

the various temperature range classes was fairly equal

(ranging from 31 to 38%) in all but two: the 8-11.9°C

class had a 54% occurrence and the 0-3.9°C class had a

19% rate (Table 25).

Relation to Sediment Organic Carbon

Among the various taxa considered in this study, Cumacea

is one of only a few which showed a marked preference in

relation to organic carbon content. Sediments containing

2.00-2.99% organic carbon were clearly preferred over all

other content classes (Tables 26, 28; Fig. 159). There the

mean density of cumaceans was 123/m2 with a mean

biomass of 0.24 g/m2, reflecting, no doubt, the habit

adopted by many species of browsing organic matter from

sand grains. Another peak in abundance (251m2 and 0.09

g/m2) occurred in the much lower organic carbon content class 0.01-0.49% Significantly lower densities and

biomasses occurred in the other carbon content classes,

and none occurred in levels above 3%.

OL--':"--:O,.':.O"'-I--~0,'=-ll--"7I.1::.0--...,I:.I:.II:----2=".0=-----=sL:,.0---l~--::ll"::.0:--~..."c!o.-=0--...J0

PERCENT ORGANIC CARBON

Figure 159

Density and biomass of Cumacea in relation to sediment organic carbon.
160 NOAA Technical Report NMFS 140

The color ofourspecimens was most commonly whitish

and tinged with light green or light tan. Some specimens

bore small brownish patches in the head region.

Tanaidaceans occurred in 15 samples (1 % of total).

Their density averaged <0.l/m2 and their biomass averaged <0.01 g/m2 (Table 5).

Geographic Distribution

Tanaidaceans were present only on the continental

slope and continental rise south of Georges Bank and

Southern New England (Fig. 160). They occurred in

low density (6 individuals or less/m2), and their biomass was very small «0.01 to 0.10 g/m2 ) in all sectors.

70·

MA I NE

_/ ....------- ----------- --', .,. C2D ",

IOOO- __ .....

'~

"'l

"

"

,

,

" ,

1000- ....... _

'~

""'l

,

",

"""

EXPLANAT ION

Q]O.OI-O.IO

EXPLANATION

I'7:'l tfj I 4 -

mJ 5-6

100

100

NEW

JERSEY

GRAMS

PER SQUARE METER

4' 41·

TANAIDACEA

Figure 160

Geographic distribution of Tanaidacea: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.
Composition and Distribution of Macrobenthic Invertebrate Fauna 161

Figure 162

Density and biomass of Tanaidacea in relation to water depth.

Figure 161

Density and biomass ofTanaidacea in each ofthe six geographic areas.

:IE

o

f-

",f-

:lEO

ocr'"

a: ... C>o

~a:.., f-fx'"

~:IE

..,..,

;loa:ocr f-:>

..,0

;10'"

a:

..,

Q.

-0.01

?

/

/ _d

2000 4000 0

~-o NUMBER

.--. WEIGHT

25 50 100

WATER DEPTH IN METERS

O~ I OL---';;NOt.cVA'---;;CGU-;;-lF;-:O""F--;;;GE~O;;;RG~ES~S;:;;O;;;UT~H;:;ER"'N~G;;;EO~RG!';ES;:---CS~O;;;UT~HE~R""N--'O SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

:IE

o

f- C/)fzO

l&J'"

:IE'" -0o

ffi 2 C/)f-

... l&J

0:IE

a:l&J

l&Jll:: ",ocr

:IE:> I :>0

Zlll

ll::

l&J

Q.

o NUMBER

_ WEIGHT

:IE

o

ff-

"'0

.., Z'"

:IE'"

0 0

~~"'f-

... .., 0:IE

a:..,

..,a:

"'ocr :IE:> :>0

z'"

a: OAf- ..,

Q.

None were present in quantitative samples

from the continental shelf even though previous studies and our own nonquantitative

samples have revealed their presence in certain habitats in the intertidal and shallow sublittoral areas along the coast. They appear to

be rare or absent from most ofthe outer continental shelf areas in the New England region.

Tanaidaceans were present in only two of

the six standard geographic areas: Georges

Slope and Southern New England Slope

(Tables 6, 8; Fig. 161). Average density in

each area was 0.4 individual/m2 with an average biomass of 0.01 g or less/m2.

Members of this group were present in

15% of the samples from Georges Slope and

in 9% of samples from Southern New England Slope (Table 10).

Bathymetric Distribution

In the present study tanaidaceans were found

only in water depths ranging from 366 to

3,820 m (Table 11; Fig. 162). Their density

was low in all depth classes within the range

of their occurrence but was relatively higher

(averaged 1 individual/m2) in depths greater

than 2,000 m than at shallower depths, where

they averaged only 0.1 of an individual/m2.

Biomass revealed a trend similar to that of

numerical density. The biomass averaged

<0.01 g/m2 in the shallower depth classes, and

0.01 g/m2 in deep water (Table 13; Fig. 162).

Frequency of occurrence disclosed trends

similar to both density and biomass. The

occurrence rate of tanaidaceans at depths

greater than 2,000 m was 35%, an unusually

high ratio compared to only 1 to 3% occurrence in the shallower portion of their bathymetric range (Table 15).

Relation to Sediments

Tanaidaceans were found only in soft, finegrain sediments: sand-silt and silt-clay (Tables

16, 18,20; Fig. 163). Density, biomass, and

frequency of occurrence were very low and

approximately equal in both sediment types.

Relation to Water Temperature

Tanaidaceans were encountered only in areas

where the annual temperature range was less

than 8°C (Tables 21, 23, 25; Fig. 164). All of the

measures of abundance (density, biomass, and

frequency ofoccurrence) were very low in each

ofthe two temperature range classes (0°_3.9° and

4°-7.9°C) in which tanaidaceans occurred.

BOTTOM SEDIMENTS

Figure 163

Density and biomass ofTanaidacea in relation to bottom sidements.
162 NOAA Technical Report NMFS 140

0.004 !

(I)~ :.0

0.005 ..

o~

!CII: 1&.1

0.002 ~~

%1&.1

!!a

0.001 ~'IC

~5;(1)

CII:

0 1&.1

A.

-0.01

O-~ NUM8ER

.----. WEIGHT

0--0 NUM8ER

__ WEIGHT

o

"'0 Zoo

'" ....

<;0

"'0::

~~

....

o~'"

0::'"

"'0:: mot

~:>

go 0.1- 0-- -0",,-

0:: e__----<..... '"

lJ.J Z'l h k'I "- o'--n:-'.-o--.-:';",-~'-;;------;c~c;o----,-;~-=--=~-=--..Jo 0- 3.9 4-7.9 8-11.9 12-1~.9 16-19.9 20-~3.9

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

:.

~ii~ o.a u

I&.ICII: Ii 1&.1

lL t; 0.2

0:'

G:I&.I

I&.IG: !i 0.1

15 OL.----=~~:::L..~;-----:-&.:~t-*_4..__:!=_--e'__':.f_:_ .....~l:_......__:_!_=__-.....J A. 0.01 o.a 1.0 1.& 2.0 a.o &.0 10.0

PERCENT ORGANIC CARBON

Figure 164

Density and biomass of Tanaidacea in relation to the annual

range of bottom water temperature.

Figure 165

Density and biomass of Tanaiclacea in relation to sediment organic carbon.

Relation to Sediment Organic Carbon

Tanaidaceans occurred only in areas of low organic

carbon (Tables 26, 28, 30; Fig. 165). Density, biomass,

and incidence of occurrence were all low in two organic

carbon content classes (0.01-0.49 and 0.50-0.99%).

Isopoda-Isopods were moderately sparse but widely

distributed throughout New England waters. Because

of their limited abundance and small size they made up

only a small portion of the total benthic fauna. They

accounted for <1 % of the total number of benthic

animals and only 0.2% of the total biomass (Table 3).

Isopods in our samples ranged from 3 to 20 mm in

length; the majority of specimens were approximately

10 to 15 mm long. Color of most specimens was translucent to light tan or medium brown.

Approximately 13 species ofisopods were represented

in the collections, most of which belonged to the families Cirolanidae and Idoteidae. A new species, Chiridotea

arenicola Wigley (1960a), was described from specimens

found in collections from Georges Bank.

A large proportion of the specimens were adapted

for burrowing in sand or for crawling on sandy or rocky

substrates.

Isopods occurred in 390 samples (36% oftotal) . Their

density averaged 12.1/m2 and their biomass averaged

0.29 g/m2 (Table 5).

Geographic Distribution

Isopods were widely distributed over large portions of

the study area (Fig. 166). They were especially common

on the banks and in coastal regions. They were least

common in the deeper portion of the western Gulf of

Maine, in the vicinity of Nova Scotia, and on the continental shelf and rise. Densities between 10 and 50 indi-
Composition and Distribution of Macrobenthic Invertebrate Fauna 163

72·

73·

72·

NEW

JERSEY

[] 1- 9

II 10- 49

• 50- 520

G!b EXPLANAT ION

[ill <0.1-0.9

lit 1.0-9.9

GRAMS

PER SQUARE METER

72·

ISOPODA

.,.

70·

M A (HE

.2·

.g.

Figure 166

Geographic distribution of Isopoda: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

viduals/m2 extended over large areas ofthe GulfofMaine

and Southern New England. High densities of 50 to 520

individuals/m2 occurred in relatively few small areas. Average biomass was commonly less than 1 g/m2, and the

highest average biomasses were between 1 and 9.9 g/m2.

Among the six standard geographic areas, isopods

were most numerous (18 and 17 individuals/m2) on

Georges Bank and the Southern New England Shelf

(Table 6; Fig. 167). Intermediate densities (3.9 and

9.5/m2 ) were found in the Nova Scotia and Gulf of

Maine areas. Low densities (1.0 and 1.3/m2) occurred

in the two slope areas.

The average biomass of isopods was small (0.4 g/m2

or less) in all areas (Table 8; Fig. 167).

The quantitative geographic distribution of isopods

was very similar to that of cumaceans (see Table 6). The
164 NOAA Technical Report NMFS 140

Figure 168

Density and biomass of Isopoda in relation to water depth.

::E 4 0I- ",I-

::EO

er: eD

II: LL

3 <:> ° ZII:

- ... 1-1-

:1:'"

!:!::E

.2

er: 1-:;)

"'0

~'"II:

...

Q.

0--0 NUMBER

__ WEIGHT

25 50

WATER DEPTH IN METERS

NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENGLAND SLOPE HEW ENGLAND SHELF SLOPE

GEOGRAPHIC AREA

0 NUMBER

-WEIGHT -0.

f--

-0.

f-

-0

f-

-0.

fII n. o 0

20

Figure 169

Density and biomass of Isopoda in relation to bottom sediments.

::E ::E 0 o NUMBER 1.4 0 l- I- I- _ WEIGHT ",I- "'0 25 1.2 ::EO ZIIl <till

::ELL '" 11: ....

(30 1.0 <:>0

"'II: 20 ~II: a. ... 0.8 ... "' .... .... 1- ....... 15 :1:'" O::E ~::E .6 0:: ... ...... "'0:: 10 ~II: Iller: er: ::E:::> 0.4 .... :::> :::>0 "'0 Z'" 5 ~'" 0:: 0.2 II: w ... a. Q.

0 0

BOTTOM SEDIMENTS

Figure 167

Density and biomass of Isopoda in each of the six geographic areas.

major difference was that cumaceans were

twice as numerous as isopods, but the

relative densities of the two groups corresponded rather closely. The geographic

distributions of the two groups were also

similar (see Fig. 154).

Frequency of occurrence of isopods in

the samples was moderate (48% of the

samples) on Georges Bank and the Southern New England Slope. In the other four

geographic areas isopods occurred in 20

to 35% of the samples (Table 10).

Bathymetric Distribution

Isopods occupied a very wide depth range

(5-3,820 m) and specimens were present

in all eight depth classes (Table 11; Fig.

168). Densities were highest (22-38 individuals/m2) in the two shallowest depth

classes and decreased as water depth increased. Lowest density (0.4/m2 ) was encoun tered in water depths between 1,000

and 2,000 m. In samples from the deepest

strata (2,000-4,000 m) the density was

1.9/m2.

Biomass values exhibited similar patterns in relation to water depth (Table

13; Fig. 168). The average biomasses of

isopods were much higher (ranging from

0.15 to 0.66 g/m2) in water depths less

than 500 m than in deeper water where

the average biomass ranged from only

0.01 to 0.02 g/m2.

Frequency ofoccurrence ofisopods was

highest (46 to 69%) in samples from water depths ranging between 25 and 100

m. Occurrence rates were intermediate

(12-20%) in samples from outer continental shelf and continen tal slope depths

(100-2,000 m) and moderately high

(43%) at depths greater than 2,000 m

(Table 15).

Relation to Sediments

Isopods were found in all of the sediment

types occurring in the study area (Table

16; Fig. 169) but were three to seven times

more numerous (22.4 individuals/m2 ) in

sand substrates than in any other type of

bottom. The average density ofisopods in

the other sediment types was fairly uniform, ranging only from 3 to 71m2.

The average biomass of isopods was

highest (1.36 g/m2 ) in till substrates

(Table 18; Fig. 169). Also, as was the case
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 165

0.2

:::E

o

f-

(/)f-

:::EO

<t'"

a::u.

"'0 0.6 Z a::

-w

f-fIW

~:::E

0.4 <t f-:> Wo

~(/)

a::

w

Q.

0--<> NUMBER

......... WEIGHT

occurred in the lower middle range (1.5-1.99%) of

carbon content.

Biomass of isopods was negatively correlated to organic carbon content, tending to decrease as organic

COntent increased (Table 28; Fig. 171). On the whole,

biomass was relatively low, ranging from 0.27 to 0.03 gl

m2. Notable anomalies in the general trend occurred in

the absence of measurable carbon (0.19 g/m2 ) and in a

comparatively sizeable (0.14 g/m2) biomass in the content class that contained the lowest density (1.5-1.99%).

Frequency of occurrence ofisopods in the samples in

the organic carbon COntent classes was moderately high

(47 and 25%) at the extreme ends of the range and

intermediate (5 to 15%) in the intervening COntent

classes (Table 30).

Amphipoda-Amphipods were exceedingly numerous and widely distributed throughout New England

marine waters. They were the most abundant taxonomic group encountered in this study, forming 43%

of the total number ofindividuals in the macrobenthos.

Because of their small size, however, they did not contribute a correspondingly large share (2.3%) of the

total standing crop (Table 3).

The amphipods collected included a wide variety of

taxonomically diverse kinds (Dickinson et aI., 1980;

Dickinson and Wigley, 1981). We estimate that between

100 and 150 different species are represented in the

samples. Four new genera and eight new species of

amphipods were described, at least in part, from specimens collected during the present study. The following

oL--..".o-J,.3:-::.9'------'-4•.1,;7.-=-9--::B:-!·1-:-:1.9,----;1-=-2.~15:-::.9,----;-;:,6:-!. 1-=-9.9::---;;:20;;-"'-!c23;":.9,----'0

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

10

Figure 170

Density and biomass ofIsopoda in relation to the annual range of

bottom water temperature.

70

:::E

60

f- (/)0

Z'" w

:::E u. 50

0 0

Wa::

~~u.. w 40

0:::E

a::w

wa::

'" <t 30 :::E:>

:>0

z(/)

a:: 20 w

Q.

with density, the disparity between the highest and lowest biomass values was quite large.

Average biomass (0.44 to 0.04 g/m2) in the

other five sediment types was nearly 3 to 30

times smaller than in till.

Frequency of occurrence of isopods in

samples from the various sediment types reflected about the same quantitative distribution exhibited by density and biomass combined. Fifty percent or more of the samples

from both till and sand contained isopods,

whereas only 16 to 32% of the samples from

the other sediment types yielded specimens

(Table 20).

Relation to Water Temperature

Among the six temperature range classes two

major concentrations of isopod density were

detected (Table 21; Fig. 170). One COncentration occurred in the broadest (20-23.9°C)

temperature range class, which contained the

highest density (67 individuals/m2). The second concentration occurred where the temperature range was from 8 to 15.9°e. Isopod

densities in this ZOne averaged 16 to 251m2.

Over the remaining temperature ranges, average isopod densities were from 3 to 71m2.

Although the biomass of isopods was small in all

temperature range classes, the quantities varied in approximately direct proportion to their density (Table

23; Fig. 170). Relatively large biomasses (0.34 to 0.42 gl

m2) occurred in the same temperature range classes as

those for high density. Smaller quantities were present

where the density was low.

Frequency of occurrence in each of the temperature

range classes was moderate to moderately low. The

occurrence rates varied directly with density and biomass values. Relatively high percentages ofsamples (43 to

61 %) contained isopods in the intermediate and broadest temperature range classes. Relatively low occurrence

rates (20 to 31 % percent) were found in the other

classes (Table 25).

Relation to Sediment Organic Carbon

Isopod density in the various organic carbon content

classes exhibited a trend similar to that in relation to

water temperature in that two major concentrations

were detectable (Table 26; Fig. 171). The highest concentration (18 individuals/m2) occurred in the next to

highest content class (3.00-4.99% organic carbon);

whereas, the second highest concentration (131m2 )

was in the class containing the lowest amounts of organic carbon detected (0.02-0.49%). Interestingly, a

moderate density (91m2) of isopods occurred in areas

of undetectable organic carbon. Lowest density (<11m2)
166 NOAA Technical Report NMFS 140 _

0---() NUMBER

_ WEIGHT

20.0

0.3

I- (/)1-

:10

~ID

et: 1L C)o

Zet:

0.2 %:\&1

~:I

~~

I-~

1&10 .(1)

0.1 ffi

Q.

/~ / \

/ \

/ \

/ \

/ \

/ \

/ \

/ \

? \

/ \

/ \

\

\

\

\

\

\

\

\

\

\

\

\

0.... .......

"

~IM

lI- (/)0

ZID \&I

:IlL

13 0

~et:

(/)~IO.O

IL\&I

0:1

et:\&I

\&let:

ID~

:I~

~oz(/)

ffi 11.0 Q.

OL...---,:-l::--~::__-___:'~--_:l:_-=--__::1:=__-____::~--;I;;_~j-__::!_;:_---.Jo 0.01 O.ll 1.0 loll 2.0 3.0 10.0

PERCENT ORGANIC CARBON

Figure 171

Density and biomass of Isopoda in relation to sediment organic carbon.

new forms were described by Edward L. Bousfield

(1965), National Museum of Canada: Protohaustorius

wigleyi, Parahaustorius longimerus, P. holmesi, P. attenuatus,

Pseudohaustorius borealis, Acanthohaustorius millsi, A. intermedius, and A. spinosus. At least three suborders are

represented in the collections: Gammaridea, Caprellidea, and Hyperiidea. Both the species and the number of individuals of the first group were considerably

more numerous than in the latter two groups.

Body size of amphipods was somewhat limited. Small

species, which were common in the families Metopidae

and Stenothoidae, were 1 to 2 mm in length, or slightly

more. The largestspecies in our collections, the caprellid

Aeginina longicornis, had a body length of more than 2

em. Gammaridea larger than 1.5 em (Casco, Maera, and

a few others) were uncommon.

Color of amphipods ranged from light cream or nearly

white (Lysianopsis, Ampelisca, and many others) to moderately dark brown (Leptocheirus, Melita, and others).

No brightly colored specimens were detected. Some of

the more colorful genera were Stenothoe, Listriella, and

Amphiporeia, which have red eyes that contrast with the

cephalon, and some members have contrasting colors

on the thoracic and abdominal plates.

Quite a few species of New England gammaridean

amphipods are tube dwellers. The tubes are usually

elongate, cylindrical, or laterally flattened structures

constructed of sand grains or clay particles cemented

together. It may be significan t that at least two of the

most common genera (Ampelisca and Unciola) are tube

dwellers. Also Haploops, which is one of the few amphipods that was relatively common in the deeper waters

of the Gulf of Maine, is tubicolous.

Amphipods occurred in 862 samples (80% of total),

their density averaged 656/m2, and their biomass averaged 4.16 g/m2 (Table 5).

Geographic Distribution

Amphipods were extensively distributed throughout the

New England region (Fig. 172). They were particularly

abundant on the continental shelf, except for the deeper

parts of the Gulf of Maine and the southwestern part of

the Nova Scotian shelf. Amphipods were an exceptionally abundant group and densities in the coastal areas

and on the offshore banks commonly averaged between 100 and 1,000 individuals/m2. High density (1,000

to 8,900 individuals/m2) areas were not uncommon in

this region. Densities of 1 to 100/m2 were typical in the
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 167

[] 1-99

Illiill 100-999

.1000-8946

EXPLANAT ION

EJ<0.1-0.9

lTIillI 1.0-9.9

.10.0-49.2

NEW

JERSEY

PER

72·

" " NEW /'

'\ '" HAMPSHIRE " . , , / <·".'·CONNECTICUT '\.. '",- :

........ ..' ,,>MASSACHUSEn'

,.::. "R:.~~',) BOSTON"

7'· 70·

M A IN E

M A IN E

69·

~~~''20 KM

AMPHIPODA

Figure 172

Geographic distribution of Amphipoda: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

deeper parts of the Gulf of Maine and on the continental slope and rise. The biomass of amphipods in the

coastal areas and on the offshore banks generally averaged between 1 and 50 g/m2. In the central Gulf of

Maine and on the continental slope and rise, the biomass of amphipods averaged less than 1 g/m2.

Two of the standard geographic areas, Georges Bank

and the Southern New England Shelf, contained very

high average densities (953 and 1,269 individuals/m2,

respectively) of amphipods (Table 6; Fig. 173). Three

areas (Gulf of Maine, Georges Slope, and Nova Scotia)

had intermediate densities (118 to 280/m2). The South-
168 NOAA Technical Report NMFS 140 _

o

m~

~o.q12l

D::a..

ClO

~D::IU ~~

:I:IU

Cl~

lUlU

~D::.q

~:;)

1U0

~m

D::

IU

ll.

7

8

9

2

5

6

3

4

o NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLANO

SHELF SLOPE

GEOGRAPH Ie AREA

0 NUMBER 0'- • WEIGHT

-

0-

-

01- -

- 0-

-

01-

-

I-

-

'--

-I

r rI ro..

400

200

o

100

mo

ZI2l

IU

~a..

U O

~:5 80

"'~ a.. IU O~

D::W

IUD::

12l.q

~:;) 60 :;)0

Zm

D::

IU

ll.

120

140

Figure 173

Density and biomass of Amphipoda in each of the six geographic areas.

ern New England Slope area yielded an average of only

17 amphipods/m2.

Average biomass was very closely correlated with number of specimens and ranged from 8.34 g/m2 in the

Southern New England Shelf area to 0.08 g/m2 in the

Southern New England Slope area (Table 8; Fig. 173).

Frequency of occurrence of amphipods was generally

high; they were found in 58 to 94% of the samples. In

the three shallow shelf areas their occurrence ranged

from 92 to 94% of the samples. Occurrence was lowest

(58 to 73%) in the Gulf of Maine and the two slope

areas (Table 10).

Bathymetric Distribution

Amphipods occurred at depths of 5 to 3,975 m. Their

densities were highest on the continental shelf and

decreased with increasing water depth (Table 11; Fig.

174). Usually high average densities (1,149 and 1,274

individuals/m2) were encountered at depths between

25 and 100 m. In water depths greater than 100 m,

amphipod densities dropped sharply. The average value

for the lower continental slope area was 121m2, and on

the continental rise their density declined to an average

of only three specimens/m2.

Biomass values revealed distribution patterns in relation to depth similar to those described above for density (Table 13; Fig. 174). At depths less than 100 m, the

average biomass values were between 6 and 10 g/m2 . In

water deeper than 100 m, a pronounced decline in biomass occurred; values ranged from 0.97.g/m2 on the outer

continental shelf to 0.02 g/m2 on the continental rise.

Frequency of occurrence was moderate to high. In

the shallow and moderately shallow water (0-100 m)

amphipods occurred in 80 to 98% of the samples, but

in deeper water they were present in 44 to 77% of the

samples (Table 15).

Relation to Sediments

Amphipods were encountered In all of the different

bottom types and revealed marked changes in density
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 169

2

10

0- --<) NUMBER

__ WEIGHT

Amphipod biomass in relation to temperature range

paralleled the same trends as those revealed by numerical density (Table 23; Fig. 176). Greatest biomass (S.l

g/m2 ) occurred in an intermediate temperature range

class (SO-11.9°C). Moderate values occurred in adjacent broader and narrower range classes. The smallest

(0.41 g/m2 ) biomass was found where the temperature

range was less than 4°C.

The frequency of amphipods in our samples was

moderate to high in all temperature range classes (Table

25). The highest rates of occurrence (92 to 97%) were

encountered where the temperature range was moderate. Somewhat lower incidence rates occurred where

the ranges in temperature were slightly narrower and

slightly broader. Lowest occurrence rates prevailed in

those areas where the temperature range was lowest

(less than 4°C) and highest (more than 20°C).

Relation to Sediment Organic Carbon

Amphipods occurred in all sediments containing organic carbon (Table 26; Fig. 177). Two abundance

peaks were clearly evident. Densities were very high

(between 1,000 and 1,256 individuals/m2) in both low

(0.01-0.49%) and high (3.00-4.99%) concentrations

of organic carbon and were much lower (24 to 164/m2)

where organic carbon content was between these exa: W wa:

lD cf

~~ 600 ~o

zen

a:

w

a. 400

200

1400

o

1200 eno

ZlD

W

~llU 0 1000

~a:

en~

~~ 800

Figure 174

Density and biomass ofAmphipoda in relation to water depth.

°0!---1:l=0-----::2~5-~50::----:I~OO=--200~--=5=!OO~t=:10::t:00::=*20=:'OO~L4000~0

WATER DEPTH IN METERS

from one type to another (Tables 16, 17; Fig. 175).

Density was exceptionally high (1,23S individuals/m2)

in sand and contributed over 56% of the total number

of animals in this type ofsediment. Density was moderately high in gravel; intermediate in sand-silt, till, and

shell; and relatively low (231m2) in silt-clay.

Biomass trends were the same as those for density (Table

IS; Fig. 175). Sand sediments contained the largest biomass, an average of7.7 g/m2• Intermediate quantities (0.5

to 3.4 g/m2) were present in gravel, till, shell, and sandsilt. The lowest biomass (O.lS g/m2) was found in silt-clay.

The occurrence of amphipods in different types of

sediments ranged from very high to moderate. It was

high to very high (93 to 100%) in sand, shell, and gravel

types; intermediate in till and sand-silt; and moderate

(51 %) in silt-clay (Table 20).

Relation to Water Temperature

Amphipods occupied the entire spectrum of temperature range variations occurring in the study area (Table

21; Fig. 176). Highest average density (1,372 individuals/m2) was encountered where the temperature was

intermediate (SO-11.9°C). Densities were moderately

high (59S-S09/m2) in areas where the range in temperature was broad, but were substantially lower (5S311/m2) where the temperature range was nil or small.
170 NOAA Technica.l Report NMFS 140

o

.... "'

~o.... «al

a:: ..... Cl O

Za::

-w

.... .... :r:w

~~

Ww

~a::

« .... :::> Wo

~",

a::

8

7

6

SAND- SILTSILT CLAY

GRAVEL TILL SHELL SAND

0 NUM8ER

0-

-WEIGHT -

0-

-

0-

o~ -

c-

-5

0-

01- -4

01-

--'3

0"-

- -2

f-

-I

lII I 1-' 0

60

o

100

90

40

200

100

a::

w

Cl.

300

110

120

o

....

.... '" 0 80 Zal

W .....

<:;0

~~ 70

"' .... ..... w o~

a::w

wa::

al«

::1::::>

g 50

BOTTOM SEDIMENTS

Figure 175

Density and biomass of Amphipoda in relation to bottom sediments.

tremes. Lowest density occurred in the absence of measurable organic carbon.

Biomass followed the pattern established by density

(Table 28; Fig. 177). Biomass was high (8.3 to 5.3 g/m2 )

at high and low carbon concentrations, fell to much

lower levels (0.2 to 1.8 g/m2) in intermediate levels,

and was lower still where carbon was absent or at the

highest levels measured.

Frequency of occurrence was high (93 to 100% ofthe

samples) in the organic carbon content classes containing the lowest and highest concentrations measured

but dropped to moderate levels (49 to 72%) in all other

classes (Table 30).

Mysidacea-Mysids constituted a minor portion of

the total benthos, 0.2% of the number of individuals

and less than 0.1 %of the biomass (Table 3). They were

small in size, their geographic distribution was limited,

and their numerical density was generally low. The

average number of specimens usually was less than 51

m2 and average biomass less than 0.01 g/m2 . Some

species of mysids characteristically make diurnal excursions from the sea bottom to the upper water layers, at

which time they become members ofthe plankton community; however, when they retreat to the ocean floor

to feed or to excavate in the bottom sediments, they are

considered an integral part of the benthos. Since bottom grabs do not sample mysids well, our data should not

be taken as indicative of actual distribution or abundance.

Individual specimens in our collections ranged in

body length from 3 to 12 mm. The large inshore species

Mysis stenolepis and Praunusflexuosuswere not present in

our quantitative samples.

The color of the majority of specimens was white or

translucent with small areas of brownish to nearly black
Composition and Distribution of Macrobenthic Invertebrate Fauna 171

o1----o=-.-':3;-';.9,----74 .-':;7;-;:;.9:------;:8;-.~11.-;:;9-..1ns.9-:-::16~-1!-=9-::.9-:::;20:;---;;.2:;-3.=-9---'0

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

2

Average density was low (1-49 individuals/m2) in all

localities except for two small patches of moderate

density (50-991m2 ) south of the northern edge on

Georges Bank and a patch of moderately high density,

between 100 and 187 individuals/m2, on south-central

Georges Bank.

Mysid biomass was typically low «0.1-0.9 g/m2 ) in

all areas of occurrence except for one small patch in

the region of highest density where biomass barely

exceeded 1 g/m2.

Georges Bank was dominant among the standard

geographic areas in both measures of mysid abundance

(Tables 6, 8; Fig. 179), containing an average density

and biomass of 10.6 individuals/m2 and 0.06 g/m2, respectively. The other geographic areas contained significantly lower amounts, ranging from 10 to 100 times less in

terms of density and 6 times less in terms of biomass.

10

<r- -{) NUMBER

_ WEIGHT

1400

8

1200

0 0 l- I- enlenoI- 1000 ~o

e:[CIl

W

ZCIl 6 a:: II.-

:Oil.. ClO

u O

wa:: ~a::

ll.w w en I- 800 1-1- :I:W ... w Cl~ o~ Ww a::w ~a:: wa:: e:[ IDe:[ 4 1-::> ~::> 600 w O ::>0

ZtJ) ~en

-~

a:: w w 11. 11.

400

200

Figure 176

Density and biomass of Amphipoda in relation to the annual range of bottom

water temperature.

pigmentation on the body and appendages. The eyes of

most species were brownish-black. The most colorful

members of the group were specimens of Erythrops

erythrophthalma, which had red eyes and yellow and orange color patches on the body.

Mysids occurred in 41 samples (nearly 4% of the

total). Their density averaged 2.5/m2, and biomass averaged 0.01 g/m2 (Table 5).

Geographic Distribution

Mysids were sparsely distributed in the study area occurring in widely separated patches mainly along the

Southern New England, Long Island, and New Jersey

shores (Fig. 178), and on Georges Bank. Two small

enclaves also occurred inshore on the central Maine

coast and in the mouth of Saint Mary Bay in western

Nova Scotia.
172 NOAA Technical Report NMFS 14U _

1400r ----------------- ...,

O-~ NUMBER

_____ WEIGHT

1200

2

~IOOO

11)0

ZCD

ILl

21&.

0 0 800

ILIa:: IL ILI II)~

I&. ILl

02

a:: ILl 600

ILIa::

CDc 2::)

::)0

ZII)

ffi 400

IL

200

r

I \

I \

I \

I \

I \

I \

I \

I \

I

I

I

I

Figure 177

Density and biomass of Amphipoda in relation to sediment organic carbon.

10.0

8.0

2

o

II)~

2 0

e CD

a:: I&.

6.0 C) 0

Za::

-ILl

~~

% ILl

02

ILl ILl

4.0:' ~::)

ILI~ :. a:: ILl

IL

2.0

Frequency of occurrence of mysids in samples was

low in all areas ranging only from 1 to 10% (Table 10).

Georges Bank had the highest occurrence frequency.

Bathymetric Distribution

Mysids occurred in the somewhat limited depth range

of 9 to 292 m. They were most abundant (densities of

3.8 to 6.5 individuals/m2 ) at depths less than 100 m

(Table 11; Fig. 180). At depths greater than 100 m,

their density was only 0.1 or lesslm2•

Biomass, although very small, revealed a rather constant diminution in quantity from shallow to deep water

(Table 13; Fig. 180).

Mysids occurred in 13% of the samples from the

shallowest depth class, and their rate of occurrence

dropped with increasing water depth to 1% or less in

the deepwater classes (Table 15).

Relation to Sediments

Mysids were found in four of the six bottom sediment

types (Table 16; Fig. 181). Their density (5 individualsl

m2) in sediments composed of sand was substantially

higher than in the other sediment types. In gravel,

sand-silt, and silt-clay the densities were less than 11m2 .

Biomass was very low (0.02 g/m2) even in sand sediments

where mysids were most abundant (Table 18; Fig. 181).

Values for the other bottom types were 0.01 g or less/m2.

Mysids were present in a higher proportion of the

sampIes (6%) from sand sedimen ts than from the other

bottom types (Table 20).

Relation to Water Temperature

Mysids exhibited a general trend of increasing in both

density and biomass as the annual range in temperature broadened (Tables 21, 23; Fig. 182). Average numerical density increased from <0.l/m2 where the temperature range was <4°C, to 6.1/m2 in localities where

the temperature range was over 20 QC. Biomass ranged

from <0.01 to only 0.02 g/m2•

Frequency of occurrence values varied in a similar,

but more consistent, manner to those of density and

biomass. The percentage ofsamples yielding specimens
Composition and Distribution of Macrobenthic Invertebrate Fauna 173

NEW

JERSEY

GRAMS

PER SQUARE METER

1- 49

50- 99

100-187

69°

o 120 '--~_~. KM

70·

,..-----

/ ----....... , 4'· .... .....

"

•••

"'.-----

/ ---- ..... ,....... 4'· .....

"

MA I NE

.,.

".

---------

'------..,.-

72°

39"

EXPLANAT ION

[ill <0.1-0.9

II 1.0- 1.2

EXPLANATION

[ill

m

•

72°

72°

MYSIDACEA

Figure 178

Geographic distribution of Mysidacea: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

ranged from less than 1% in the lowest range class to

18% in the highest range class (Table 25).

Relation to Sediment Organic Carbon

Mysids occurred in four of the eight organic carbon

content classes (Table 26; Fig. 183). Average densities

(ranging from 0.8 to 1.8 individuals/m2) were quite

similarly distributed among the four classes, exhibiting

a bimodal trend, one in the lower carbon content classes

(between 0.01 and 0.99%) and the other in the intermediate (1.5 to 2.99%) classes.

Biomass values were uniformly low (0.01 g/m2 ) in all

organic carbon classes in which they occurred (Table

28; Fig. 183).
BOTTOM SEDIMENTS

Figure 180

Density and biomass of Mysidacea in relation to water depth.

Frequency of occurrence of mysids was

higher (7 and 8%) in the intermediate carbon content classes than in the lower ones (3

and 5%) in which they occurred (Table 30).

Decapoda-Decapods, although well represented in terms of the number of taxa that

were found in our quantitative samples (two

suborders and three sections contained 24

genera and 34 species, Table 4), did not rank

as highly in their contributions to the total

number of specimens (0.5%), or to total biomass (0.8%) (Table 3). The apparent poor

showing of the class, the largest among all

classes of crustaceans, is misleading, precisely

because of the large size and mobility of many

of the representative species. These large,

highly mobile forms are more effectively

sampled by qualitative gear types, such as

dredges and trawls, than by quantitative grab

samplers. Thus the estimates of density and

biomass presented in this report, for this class

of crustaceans, should be considered to be

very conservative at best. Indeed, the taxonomic list of decapods obtained by means of

qualitative sampling gears in our databases is

considerably more expansive than the one

included here in Table 4. Nevertheless, the

quantitative data reported in our report contain a fair representation of the major forms

present in the region and constitute the most

extensive and complete set (known to us) upon

which to base our estimates.

Decapods were not found in most of the

Gulf of Maine. For the most part, they are

restricted to the Southern New England continental shelf and upper slope and to Georges

Bank; some occurred on the western portion

of the Nova Scotia shelf and the western basin

of the Gulf of Maine. Average densities were

low, ranging from 1 to 49 individuals/m2, over

most of their range in the study area. Size of

captured specimens tended to be smaller than

the overall average for this class, reflecting the

bias imparted by the relatively small area

sampled by the quantitative grabs used in our

study. The smaller members of this group were

less adroit at avoiding capture. The average size

of caridian shrimps ranged from 20 to 40 mm;

occasional larger specimens (40-60 mm) were

captured. The latter were usually representatives of more sedentary, less active taxa, such as

the burrowing sand shrimp Crangon septemspinosus. Average anomuran size ranged between

4 and 20 mm carapace length. Most frequently

captured were representatives of the relatively

o

....

~6el lll

"'''- "'0

z",

-w

........

::I: W

"'~ ww

~a::

el

0.01 5

~'"

'"

W

Q.

0.02

o NUMBER

• WEIGHT

0- --<l NUMBER

....- WEIGHT

6 ....

....

o

III

"-o

'"

W

....

W

W

a:: 2 el

::>

o

'"'"

W

Q. Ol-----,G:-!R,.,JAV~El,-L---;T~1L7L-~S;:;-H;;:E;-:LL-~S!:;A~ND~-----;;~L-----!.~L-------'0

Figure 181

Density and biomass of Mysidacea in relation to bottom sediments.

Figure 179

Density and biomass of Mysidacea in each of the six geographic

areas.

174 NOAA Technical Report NMFS 140

12

0 NUMBER

• WEIGHT

10

0 0 .... ....

.... "'

~o.... "'0 ZIIl B el lll

W

~"- "'''- "'0 ijO

w'" 0.05 ~'"

5i~ W........ ,,-w 6 ::I:W

o~ 0.04 "'~

"'W Ww

wa:: ~'"el lIlel 0.03 .... ::> ~::> 4 wO ::>0

Z", ~'"a::

'" 0.02 W W Q.

Q.

0.01

0 0

GEOGRAPHIC AREA
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 175

Figure 182

Density and biomass of Mysidacea in relation to the annual range

of bottom water temperature.

0

... Ul'"

~o

0.03 <tID

a: .... <!lo

~a:

......'" 0.02 :1:'"

~~

"''''

~a:<t ... => "'0 0.01 ~Ul

a:

'"

Q.

0

0-- -0 NUMBER

_ WEIGHT

J\

/ \

/

/

the high annual yields of lobsters and shrimps. However, low densities were recorded from the northeast

corner of the Gulf at the en trance to the Bay of Fundy,

the Western Basin section north of Great South Channel, and some inshore localities. Low densities (1-49

individuals/m2) also prevailed over most of their range

elsewhere on the continental shelf in the study area.

Moderate (50-99/m2 ) and high (100-266/m2 ) average

densities were· restricted to small patches on Georges

Bank and in Nantucket and Vineyard Sounds and at the

head of Long Island Sound.

Average biomass of decapods was low «0.1 to 9.9 g/

m2) over the major portion of their range; there were

only a few small patches of moderate (10-50 g/m2 )

8,-------------------------,

OL---;::;O~'3i<.9.--....... 4:!;.7;;;.9-....,8o:.+011.~9 -""12c-!'1!-;-5.~9 -----C1C::- 6 -+'19"".9:;---;;:20""'-!:23"".9:;-----.J

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

a:

'"t;j 4

a:

'"

Q.

....

o

'"

a:

<t

=>

o

Ul

o

...

...

o 6 ID

slow-moving pagurid hermit crabs. Size of brachyuran

crabs averaged between 15 and 20 mm carapace width;

the two species of Cancer did provide some larger specimens in the 50 to 60 mm range, and Pinnixaspp. provided

some of the smaller specimens in the 5 to 6 mm range.

Colors of decapods in our samples ranged from the

nearly transparent or translucent white of Crangon to

the dark reddish browns and blacks of Hyas. A veritable

spectrum of colors was represented between these extremes, ranging from the delicate flesh and pink hues

of the pandalid shrimps, to the tans, greens, blues, and

grays and muted reds of the pagurid and brachyuran

crabs. Most colorful were the bright red-orange Geryon

quinquedens.

The decapods as a group are similar to

bivalves in providing a broad spectrum of

prey to a variety of predators. In addition to

man, whose harvests of lobsters, shrimps,

and crabs are well known, many other marine animals (including other invertebrates,

mammals, birds, and fishes) depend on decapod prey for a substantial portion of their

sustenance.

Decapods occurred in 246 samples (23%

of the total). Their density averaged 8 individuals/m2, and their biomass 1.32 g/m2

(Table 5).

Geographic Distribution

The most striking feature of decapod distribution revealed by our samples was their

apparen t absence from large portions of the

Gulf of Maine (Fig. 184). This artifact is due

primarily to sampling gear bias since, traditionally, this region has been well known for

2

2 2.0 0

IUlI- I- 0--0 NUMBER 2 0 CDO Q. .---. WEIGHT e:t: lII ZlII

11.1 5\ 1.5 - / \ a: ... 2'" / \ 00

u O

/ \ za: 11.1 a: / \ \ -11.1 ~II.I / \ / 1-1- CD I- 1.01- \ :J:II.I ... 11.1 / \ / \ - 0.050 0:2

02

/ \ / I&ll&l a:1&l ~a:

~a: \ e:t:

S~ 0.51- / "- / \ - 0.025l;j g ::::10

ZU) "- / "- / \

.1 a:

a:

- \1 ___ I Q I I&l I&l 0 0 0.01 0.5 1.0 1.5 2.0 3.0 5.0 10.0

PERCENT ORGANIC CARBON

Figure 183

Density and biomass of Mysidacea in relation to sediment organic carbon.
176 NOAA Technical Report NMFS 140

.2·

M A IN E

.,.

EXPLANAT ION

[ill <0.1 - 9.9

mB 10.0-49.9

• 50.0-64.8

---....-

EXPLANATION

[ill 1- 49

mB 50- 99

.100-266

70·

GRAMS

PER SQUARE METER

72·

72·

DECAPODA

Figure 184

Geographic distribution of Decapoda: A-number of specimens per square meter of

boltom; B-biomass in grams per square meter of boltom.

biomass, mostly in inshore regions near Cape Cod and

Long Island. One small area of moderately high biomass (50-65 g/m2 ) occurred on the southeast part of

Georges Bank.

Decapods occurred in five of the six standard geographic areas; they were absent only in the Georges

Slope area. Highest average density (221m2) occurred

on Georges Bank followed by significantly lower densities on the Southern New England Shelf and off Nova

Scotia (9 and 21m2, respectively). Density was below II

m2 in the Gulf of Maine and on the Southern New

England Slope (Table 6; Fig. 185).
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 177

..-------------------,4

Figure 186

Density and biomass of Decapoda in relation to water depth.

o

... en'"

~o

el lD

It: IL <:>0

Zit:

-w

......

:E:W

<:>~

Ww

~It:

eI ...::;)

wo

~en

It:

W

ll.

o

(>--0 NUMBER

.--. II/EIGHT :::E

o

... 3 en b :::ElIJ

eI

l;15

zit:

-~ 2 ... w :E::::E

~w

Wit:

~eI::;)

I t;:;g

It:

W

ll.

25 50 100 200 500 1000 2000 4000 0

WATER DEPTH IN METERS

GEOGRAPHIC AREA

NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

m2; however, sand contained nearly five times less biomass (1.16 g/m2 ) than gravel. Sand-silt and silt-clay, the

finest sediments, yielded lower density and biomass.

Frequency of occurrence of decapods was moderate

and uniform in samples in gravel, shell, and sand, ranging from 33 to 37%, but was relatively low in sand-silt

and silt-clay, 7 and 4%, respectively (Table 20).

Relation to Water Temperature

Average density and biomass of decapods showed a tendency to increase with broadening temperature range to

5

0 NUMBER

• WEIGHT

0-

-4

5f-

-3

Of- -

5f- -I

0 I J

o 10

20

:::E

o

...

b 16 wlIJ

~15 o

It ffi 12 en ...

~~

ffi B lIJeI

~i5

Z 4

W

ll.

2

2 o

...

... eno

Zen

W

~IL

0 0 I

Wit:

5;~

ILW

O~

It:W

Wit:

enel

~::;)

::;)0

Zen

It:

W

ll.

Figure 185

Density and biomass of Decapoda in each ofthe six geographic areas.

Average biomass ranged from slightly over

3 g/m2 on Georges Bank to 0.02 g/m2 on

the Southern New England Slope (Table 8;

Fig. 185). In the other areas, biomass ranged

from 0.6 to slightly over 2 g/m2.

Frequency ofoccurrence ofdecapods in the

samples was moderately high on Georges Bank,

Southern New England Shelf, and off Nova

Scotia, ranging from 46 to 18% (Table 10).

Their occurrence in samples from the Gulf of

Maine and Southern New England Slope was

considerably lower, 6 and 3%, respectively.

Bathymetric Distribution

Decapods were almost wholly restricted to

water depths of less than 500 m and showed

a general trend ofdiminishing in abundance

as water depth increased (Table 11; Fig. 186).

Average density was highest (18/m2 ) in the

shallowest depth-range class (0-24 m) and

dropped to 50% and less of this value in the

continental shelf depth classes between 25

and 200 m. Density in the upper slope depth

class (200-499 m) was low (0.3/m2 ) and

very low (0.l/m2) in the only deepwater

depth class (1,000-1,999 m) in which they

occurred.

Average biomass generally followed the

trend established for density (Table 13; Fig.

186). In the shallow-water depth class, average biomass was nearly one and one half to

three times (3.3 g/m2 ) higher than that in

the continental shelf depth classes where it

ranged from 1.1 to 2.1 g/m2• Biomass was

0.61 g/m2 in the upper slope depth class

(200-499 m) but only 0.03 g/m2 in the deepest class in which decapods occurred (1000-

1999 m).

Decapod frequency of occurrence in the

samples was fairly uniform at moderate levels (35 to 39%) in the three depth classes

<99 m, moderately low (13%) in the shelf

edge class, and low (3 and 4%) in the other

two classes they occupied (Table 15).

Relation to Sediments

Decapods were present in all sediment types except till.

Both density and biomass diminished with decreasing

sediment particle size (Tables 16, 18; Fig. 187). Gravel

bottoms contained the greatest average number (241

m2 ) of decapods as well as greatest biomass (5.56 gl

m2). A drop in density occurred in shell and sand, each

ofwhich yielded an average of 91m2, nearly three times

fewer decapods, but biomass diminution was not as

dramatic in shell which contained an average of 4.78 gl
178 NOAA Technical Report NMFS 140

BOTTOM SEDIMENTS

Figure 187

Density and biomass of Decapoda in relation to bottom sediments.

Figure 188

Density and biomass of Decapoda in relation to the annual range

of bottom water temperature.

:a;

o

... ",'"

20 0 e(al

O::IL

"'0

Zo::

- ... ... ... "' ... ~2O

~~e(

... :::> "'0

~'"0::

...

Q.

2

5

;t5

e('"

4 0:: IL 00

ZO::

- ...

3 ti

0 2

......

~~ 2 ... :::>

~g

0::

...

Q.

o

<>- -<> NUMBER

___ WEIGHT

/

/

/

"

-j/

GRAVEL TILL SHELL SAND SAND· SILT· SILT CLAY

'---no-=-'.3".9.-----;;:4:';.7;Q.9:--........-:B.+.IIO.9-"12;::.+-150.9--;;;16,...:·1!;;-9.-;;"9---':2;;;0-!.2l-;30.9----I O

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

5

o NUMBER

• WEIGHT

-6

-

-5

5- -4

- -3

-2

-

I -III o •

ffi~ 10 ale(

:a;:::>

:::>0 z",

0::

...

Q.

:a; 20 o

...

...

o

al

IL

o

0::

...

... IL'" 020

0::'"

... 0:: ale( 10 2:::>

:::>0

z'"

0::

... 5 Q.

0

2

25 ... "'0

Zal

... 20

~~"' ... LL l.IJ 15 02

Relation to sediment organic carbon

Decapods were absent in areas where no measurable organic carbon occurred in the sediments, as well as in areas with the highest

recorded amounts. They were present in the

five organic carbon content classes between

0.01 and 2.99% where average density and biomass described U-shaped distributions (Tables

26,28; Fig. 189). Average density ranged from

nearly 8 to 0.5 individuals/m2 and average biomass from 4.15 to 0.30 g/m2. Values were highest at the extremes ofthe carbon content classes

in which they occurred and fell to the lowest.

levels in the middle carbon content classes,

slightly biased toward the higher end.

Frequency of occurrence of decapods in

the samples described a distribution similar

to density and biomass. Occurrence was moderate, ranging from 4 to 32%, but in this

instance was biased slightly toward the lower

end of the content range (Table 30).

19.9°C; beyond this, in the 20.0-23.9°C range

class, both measures declined (Tables 21, 23;

Fig. 188). Average density and biomass (0.9/m2

and 0.47 g/m2, respectively) were lowest in the

narrowest temperature range class (Q-Q3.9°C)

and generally increased with broadening temperature range, peaking at 23.2/m2 and 4.26 gl

m2 in the 16.Q-19.9°C range class. Intermediate

values of both measures occurred where the

temperature range was broadest.

The frequency of occurrence of decapods

in the samples in the various temperature

range classes parallels the trend established

for density and biomass, ranging from 6% in

the narrowest range class to 53% in the 16-

9.9°C class, and dropping to intermediate (18%)

levels in the broadest range class (Table 25).

Bryozoa

Bryozoans are sessile colonial animals most frequently

found attached to rocks, shells, ship bottoms, pilings,

firm outer surfaces of other animals, and other similar

hard substrates. Their distribution in the study area was

somewhat patchy owing to their requirement for a firm

substrate and moderate to strong water currents. The

currents transport to them their main food supply,

minute plankton, principally diatoms.

These organisms contribute a significant number

(more than one hundred) of species to the New England benthic fauna. The majority of these species belong to the class Gymnolaemata, order Cheilostomata.

The calcareous encrusting forms were especially numerous, but the chitinous foliaceous types were usually

the largest specimens encountered.

Although the Bryozoa constitute a major phylum in

species diversity and are a major contributor to the

biomass in certain localized habitats, their small size

and patchy distribution rather severely limited their

contribution to the total benthic fauna. In terms of numerical density, ectoprocts made up 1% of the total fauna

and contributed only 0.7% of the total biomass (Table 3).

Bryozoans occurred in 119 samples (11 % of total).

Their density averaged 15.7/m2. Their biomass averaged 1.29 gl m2 (Table 5).
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 179

10.0 1l.0

1.0

I- IIlI-

:Ii crill° '.0 l!>

ZII:

-101

1-1-

%101

!!!:Ii

101 101 .11:

2.0 I1010

• III

II:

101

A.

4.0

~\

I \

I \

I \

I \

I \

I \

I

I \

I \\ I \

I \

I \

I

2.0

8.0

oL..-......---=7.--~0;f,.1l=-----:-'1.0=------,J1.':-1l---=2";;.0---:,:'::.0=-- ....-,1l:1.0=--.....----:-:!10'-::.0,....--.....J0

PERCENT ORGANIC CARBON

Figure 189

Density and biomass of Decapoda in relation to sediment organic carbon.

Geographic distribution

Bryozoans were distributed in somewhat scattered tracts

in nearly all sections of the study area (Fig. 190). They

occurred most commonly in coastal areas and on offshore banks. Specimens were noticeably scarce in the

central part of the Gulf of Maine, over large portions of

the eastern Nova Scotian Shelf, from offshore parts of

the Southern New England Shelf, and on the continental slope and rise. Dense assemblages of over one

hundred colonies per square meter were present in

small areas dispersed throughout the banks and coastal

areas.

In the six standard geographic areas, bryozoans were

most prevalent, on the average, on Georges Bank (28

colonies/m2) and on the Southern New England Shelf

(22 colonies/m2) (Table 6; Fig. 191). They were moderately common in the Nova Scotia and Gulf of Maine

areas and scarce or absent in the two slope areas.

The largest average biomass, 6.3 g/m2, occurred in

the Nova Scotia area (Table 8; Fig. 191). Georges Bank

ranked second with 2.6 g/m2, and all other areas contained less than 1 g/m2 .

Incidence of occurrence in the four continental shelf

areas generally diminished from a high of 19% in the

northeast to a low of 11 % in the southwest. Their occurrence was even lower, 6 and 0%, in the two slope areas

(Table 10).

Bathymetric Distribution

Bryozoans were taken at water depths ranging from 8 to

3,820 m. There was a very pronounced decrease in the

density as water depth increased (Table 11; Fig. 192).

The average number of colonies in the shallowest (0-

24 m) depth zone was 391m2 and decreased steadily to

an average of 4.3/m2 on the upper continental slope at

a depth of 500 m. Below 500 m they were absent or

present in very low (0.5 colony/m2) quantities.

The average biomass of bryozoans was higher in shallow water than in deep water. This relationship

was similar to that described above for density, except

for an unusually large average biomass of 2.9 g/m2

at depths between 50 and 99 m. This was the largest

average biomass from anyone depth class (Table 13;

Fig. 192).
180 NOAA Technical Report NMFS 140

12°

,,0

NEW

JERSEY

EXPL4N4TION

Ell 1-49

II 50-99

.100-798

EXPLANATION

[ill <0.1- 9.9

fIBJ 10.0- 49.9

• 50.0- 225.9

GRAMS

PER SQUARE METER

...

,,0

..,.

BRYOZOA

,,0

.,0

M A I NE

M A I NE

••0

o

~'M

120

.,0

Figure 190

Geographic distribution of Bryozoa: A-number ofspecimens per square meter

of bottom; B-biomass in grams per square meter of bottom.

The percentage ofsamples containing bryozoans generally decreased with increasing water depth. The highest incidence of occurrence (19%) was in the shallowest depth class. At depths greater than 500 m, they

occurred in from 0 to 5% of the samples (Table 15).

Relation to Sediments

Bryozoans were especially common in the hard, coarse

substrates that afforded suitable surfaces for attachment.

In addition, they were also found in smaller quantities in

the soft fine-grain sediments but attached to hard biogenic materials, occasional pieces of gravel, or man-made

debris (Tables 16, 18; Fig. 193). Shell substrate seemed

particularly suitable in as much as it yielded an average of

over 300 colonies/m2 and an average biomass of nearly 17

g/m2. Moreover, bryozoans made up over 25% of the

total number ofspecimens and 7.5% ofthe biomass ofthe

total benthic fauna in shell substrates (Tables 17, 19).

Gravel substrates ranked second in quantity, with an average density of75 colonies/m2 and a biomass of 7.4 g/m2.

Quantities were low (less than 6 colonies and 0.4 g/m2) in

the remaining four sediment types.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 181

3.0

Brachiopoda

7

6

o

::Ii

o

.... Ul~

5 CD

a:: .... "'0

~a::

4 ........ '"

:z:'" ",::Ii

~~ot .... ::> ",0

;JUl

a::

'"

0..

0.5

.... 2.5 Vl b::Om

<I:

0:: .... ",0

2.0 z 0::

-UJ.... .... UJ :Z:::O

1.5 ~W

Wo::

~<I:

.... 5 1.0 Vl

0::

W

0..

WATER DEPTH IN METERS

NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENGLANO SLOPE NEW ENGLAND

SHELF SLOP(

GEOGRAPHIC AREA

0---0 NUMBER

__ WEIGHT

Brachiopods, commonly known as lamp shells, are sessile

organisms normally found attached to rocks and other

firm substrata at continental shelf depths, usually in

cold water. These requirements limited the scope of

their distribution to the northeastern sectors where

suitable habitats were more prevalent than elsewhere in

10

o NUMBER _

• WEIGHT

-

-

o~ -

- 0"-

-

oIL -I

0 I

Figure 192

Density and biomass of Bryozoa in relation to water depth.

Figure 191

Density and biomass of Bryozoa in each ofthe six geographic areas.

Incidence of occurrence was highest (50%)

in shell, moderately high in till and gravel, and

low (10% or less) in the fine-grain sediments

(Table 20).

Relation to Water Temperature

Although bryozoans were rather severely limited in distribution by specific substrate requirements, water temperature range appeared to

playa lesser role in inhibiting abundance. Members of this phylum occurred in all temperature range classes but revealed a pronounced

trend of increasing density with a broadening

of the temperature range (Table 21; Fig. 194).

The density of bryozoans averaged only 3 colonies/m2 where the temperature range was less

than 4°C. Their density increased to an average

of66 colonies/m2 where the temperature range

was greater than 20°C.

Biomass values also exhibited a general upward trend (0.28-2.45 g/m2 ) as the temperature range broadened. This increase, however,

was less consistent than that exhibited by numerical density (Table 23; Fig. 194).

The occurrence of bryozoans in the samples

ranged from 8 to 21 %. Generally, incidence of

occurrence was low where the temperature

range was narrow, and high where the temperature range was broad (Table 25).

Relation to Sediment Organic Carbon

The relationship of bryozoans to sediment organic carbon was not nearly as well defined as

that in other parameters because no orderly

trend or pattern was discernible. Relatively high

average densities, between 21 and 35 individual

colonies/m2, occurred in widely separated organic carbon content classes in the low, middle,

and higher regions of the content spectrum

(Table 26; Fig. 195). Significan tly lower average densities, ranging between 0 and 81m2,

occurred in adjacent carbon content classes,

effectively separating and isolating the higher

values.

Average biomass was distributed in a manner

similar to that for density but was not as pronounced. Highest bryozoan biomasses (1.95 and

1.21 g/m2) occurred in the low and upper middle

carbon content range, interspersed with significantly

lower values (Table 28; Fig. 195).

The percentage of samples containing bryozoans

ranged from 0 to 25% (Table 30). Incidence of occurrence showed a general trend of decreasing (20 to 0%)

as organic carbon content increased from 0 to 2.99%,

but it shot up to 25% in the 3.00-4.99% class.
182 NOAA Technical Report NMFS 140

6

B

o

2

4

SAND· SILTSILT CLAY

GRAVEL TILL SHELL SAND

0 NUMBER

• WEIGHT

01-

-I

I- -I

f-- -I

f-- -I

-I

I-

-

'-

-

- r- -

I- -

.... .. r-t o

40

160

80

120

200

32

240

360

280

BOTTOM SEDIMENTS

Figure 193

Density and biomass of Bryozoa in relation to bottom sediments.

0

1

'-----;0:-.;-3.9;;------:4;-'.:;-;7.9;;-----;;B-+.I7";1.9;;----'1"'"2-+.15=-:.9;;----;-1=-6'+:'9:-::.9-=-20'-.!..,23,.-,.9----10

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 194

Density and biomass of Bryozoa in relation to the annual range of

bottom water temperature.

eo

0--0 NUMBER

lE ----.. WEIGHT 0

ff0 60 al

LL

0

Q:

"I

.... 40 LLW

OlE

Q:W

wQ:

al<t

lE:>

:>0

Z", 20

Q:

"I

l1.

lE

o

.... "' .... lEO

<tal

Q:LL

"'0

ZQ:

-"I

.... .... 2 :I: "I ",lE

"1"1 :t:Q: <t f-:>

"10

:t:'"Q:

"I

l1.

the New England region. Although often

locally significant in terms of density and

biomass, their contribution to the total fauna

was limited by the patchiness ofsuitable habitats. Brachiopods accounted for only 0.3%

of the total number of specimens and 0.7%

of the total biomass (Table 3).

Brachiopod diversity was also noteworthy.

Although the phylum is not noted for large numbers of species, there being only about 280

known living species, our samples contained

only members of one genus, Terabratulina.

Size ofspecimens ranged from about 5 mm

to some large specimens of about 30 mm

length. Color was typical for the genus with

dull white and silvery gray hues predominating; many specimens were fouled by other

sessile forms such as bryozoans, hydroids, and

encrusting sponges of various hues, but tans,

browns, yellows, and grays predominated.
_________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 183

40.0

10.0

0--0 NUMBER

_ WEIGHT

/~\

/ \

/ \

/ \

/ \

/ \

/ \

/ \

/ \

/ \

I \

Z.O

1.6

0.4

oL-""""7.l:-:--__::1=_J:=::r;:~____;~-__::1=_~~:I.::_-__:~~__,~----JO 0.01 O.!l 1.0 I.!l Z.O 3.0 10.0

PERCENT ORGANIC CARBON

Figure 195

Density and biomass of Bryozoa in relation to sediment organic carbon.

Brachiopods occurred in 54 samples (5% of total).

Their numerical density averaged 4.5 individuals/mz,

and their biomass averaged 0.89 g/mz (Table 5).

Geographic Distribution

Brachiopods were restricted to the northeastern sector

of the study area, the only region in which suitable

habitats were found (Fig. 196). Numerical density was

usually low «49 individuals/mZ) over most of their

range as was biomass (<10 g/mz); small areas of moderate (between 50 and 100/mz) density and biomass (10

to 63 g/mz) occurred in south-central Gulf of Maine

adjacent to the northern edge of Georges Bank and on

the Nova Scotian shelf. Significant densities (between

100 and 490/mz) did occur in a few places, notably at

the mouth of the Bay of Fundy.

Among the six standard geographic areas brachiopods were restricted to Nova Scotia, the Gulf of Maine,

and Georges Slope. Largest average density (22/mz)

and biomass (3.68 g/mz) occurred off Nova Scotia followed by Gulf of Maine (9.5/mz and 2.12 g/mz);

Georges Slope contained insignificantquantities (Tables

6, 8; Fig. 197).

Twenty-one percent of the samples off Nova Scotia

contained brachiopods versus 12% in the Gulfof Maine;

only 2% of the Georges Slope samples yielded specimens (Table 10).

Bathymetric Distribution

Brachiopods were taken at depths between 51 and 690

m. Very few occurred in water deeper than 499 m

(Tables 11, 13; Fig. 198). Largest average density and

biomass were found at depths between 200 and 499 m

in so-called upper slope depths; there density averaged

17 individual/mz and biomass nearly 4 g/mz. Both

average density and biomass decreased significantly with

decreasing depth above 200 m.

Frequency of occurrence of brachiopods in the

samples was moderate to low. The highest frequency

was in the 200-499 m depth class at 14%; the rate
184 NOAA Technical Report NMFS 140

"-

"

19·

42·

70"

M A IN E

4,°

1-49

50-99

100-490

59°

100

1000- __ ...... _ ..........

'" ,,

.... t

,

",

,

....,

...........,

~",

EXPLANATION

LIl <0.1 - 9.9

m10.0-62.8

1000--.. _ ...............

NEW

JERSEY

EXPLANATION

[]

o

•

GRAMS

PER SQUARE METER

74°

72°

BRACHIOPODA

Figure 196

Geographic distribution of Brachiopoda: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

decreased to 5% in deeper water classes and to 10 to

2% in the shallower water depth classes (Table 15).

Relation to Sediments

Brachiopods occurred in all sediment types found in

the region but showed a pronounced preference for

the coarser types over the finer ones (Tables 16, 18; Fig.

199). Till sediments ranked first in preference, containing an average of 48 individuals/m2 and an average

biomass of nearly 16 glm2.

Interestingly, although shell sediments ranked second in terms of density (371m2), they ranked last (0.22

g/m2) in terms of biomass. Gravel, with a mean density

and biomass of 141m2 and 2.44 g/m2, respectively,
Composition and Distribution of Macrobenthic Invertebrate Fauna 185

Figure 198

Density and biomass of Brachiopoda in relation to water

depth.

:E

o

.... "' .... :EO

.. a> a:u.

"0

~a:

........ '" :x:'" ,,:E

"''''

~a:.. ....~

",0

~'"a:

'"

n.

4

-4

-3

-2

-I

000 4000 0

o NUMBER

• WEIGHT

0- -0 NUMBER

_ WEIGHT

25 50

WATER DEPTH IN METERS

201..-----------------------,

O'-----';N~OV:='A--;;:G;;;UL~FO~F-G;;<'~O';;-;G,'Os----"SO;;CUT"t;:;H<-":;;"N--;;:G'~O'::::G '~S----"SO;;;-UT¢;;H<-":;;"N --'0

SCOTIA MAINE BANI( NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

ber of organisms in the region, and Holothuroidea

rank fourth with a 7% contribution (see Table 3).

Ophiuroidea and Asteroidea are subdominants, providing 1.8 and 1.2%, respectively, of the total number

of organisms. In terms of biomass, however, none of

the four rank among the top dominants, but

Ophiuroidea and Echinoidea are subdominants providing 2.9 and 1.9%, respectively, of the total. Holothuroidea and Asteroidea contributions to biomass,

0.3 and 0.1 % respectively, are significantly below their

showing in terms of density (see Table 3).

Figure 197

Density and biomass of Brachiopoda in each of the six geographic areas.

25

:E 20f0

....

.... "'0 za>

:lEu. '"

<3 0 151-

"'a:

g;~

u.'" 0:E

a:", 101- "'a: a> .. :E~

:>0

z'"

a:

Relation '"

n. sI- to Water Temperature

Brachiopods were quite restricted in their relation to the annual range in water temperature

and showed a very strong tendency of decreasing

abundance with increasing temperature range

(Tables 21, 23; Fig. 200). None were found where

the annual temperature range exceeded 11.9°C.

They were most plentiful (average density of 9/m2

and biomass of 1.93 g/m2) in the narrowest (0-

3.9°C) temperature range and declined rapidly and

steadily as the temperature range broadened.

The frequency of occurrence of brachiopods in

samples in the temperature range groupings was

low (10 to 2%) and followed the trend established

in the abundance measures (Table 25).

ranked third. Both mean density (range 0 to 2/

m2) and mean biomass (range 0.24 to 0.33 g/m2)

were significantly lower in sand, sand-silt, and siltclay, each of which offered very limited attachment potential.

Samples in till substrates yielded the highest occurrence frequency of specimens (41 %), shell and

gravel were about even, but considerably lower in overall frequency than till (17 and 14%, respectively; Table

20). Only 2 to 3% ofsamples in the other sediment

types provided brachiopod specimens.

Relation to Sediment Organic Carbon

Brachiopods preferred low levels of sediment organic carbon (Tables 26, 28; Fig. 201), being absent at levels above 1.49% carbon content. Numerical density was 3 to 5 times (range 16 to 3/

m2) higher where no measurable carbon was found

than where small amounts occurred. Biomass was

greatest (1.74 g/m2) in the 0.5 to 0.99% carbon

content class versus a slightly lower biomass (1.31 g/

m2) in the 0% grouping. Significantly lower levels

occurred in the other carbon content groupings.

Frequency of occurrence of brachiopods in

samples declined rapidly from a high of 40% in

the 0% organic carbon level grouping to only 6%

in the next (0.01-0.49%) grouping, then more slowly

to 2% in the 1.00 to 1.49% grouping (Table 30).

Echinodermata

The phylum Echinodermata is represented, in the New

England region, by members of five classes: Crinoidea,

Holothuroidea, Echinoidea, Ophiuroidea, and

Asteroidea. All but Crinoidea provide significant contributions to the total benthic fauna. In terms of contribution to overall density, members of Echinoidea are

second in dominance, providing 20% of the total num-
186 NOAA Technical Report NMFS 140

20

0 NUMBER

-WEIGHT 162:

0 2: .... 0 .... 14 .... UlO Ul ....

ZeD 2: 0

'" <>:eD 2:"-

<3 0 12 <.00 "'''- "'", 60

~'" Q.", "' .... ,,-'" 10 ........ '" 02: :r'" <.02:

"'''' eD<>:

2:::> "'''' 40 8 "''''

~'"<>: ::>0 .... ::> ZUl ",0

'" ~'"

'" 6

'" Q. '"

Q.

20 4

O'---"*'~'---;CT~'L~L--7.SH~E~L~L-~S:::;:AN~D~----;:S~AN:I!D~------:S~,L=!T·---.JO SILT CLAY

BOTTOM SEDIMENTS

Figure 199

Density and biomass of Brachiopoda in relation to bottom sediments.

2_0

2: 10 C>- -0 NUMBER 2: 0 0 .... _ WEIGHT .... .... 1.6 (1)0 "' .... 2: 0 ZeD B <teD '"

2:"- "'''-

(30 <.00

"'", '-2 z'" g,~ 6 -",

........ ,,-'" :r'" 02: <.02:

"'''' O_B .... '" "'''' 4

eD<>: ~'"<>: 2:::> .... ::> ::>0 Z", 0.4 ~~

'" '" .... '" Q. Q.

0 0 0- 3.9 4·7.9 8-1'-9 12-15.9 16- 9.9 20-23.9

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 200

Density and biomass of Brachiopoda in relation to the annual

range of bottom water tempereature.

Following the procedure established above for the

other major taxa, the figures relating to Echinodermata (Figs. 202-207) will be presented here, and a

detailed discussion of the phylum will be presented in

the section "Dominant Components of the Macrobenthos" below. Discussion of the five classes of echinoderms follows.

Crinoidea-Crinoids formed a minute portion of the

New England benthic fauna. Members of this group

were obtained at only two stations. Six specimens that

totaled 0.16 g in weight were taken at station 1038 in

the Gulf of Maine. Water depth at this locality was 203

m; bottom sediments were of sand-silt. Two specimens

that weighed 0.02 g were taken at station 2185 in the

Georges Slope area. Water depth at this locality was

1,420 m; bottom sediments were composed of silt-clay.

The annual range in bottom water temperature at each

of these sites was less than 4°C. Organic carbon content

at station 1038 was 0.40% and at station 2185, 0.55%.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 187

20.0

S.O

\

\

\

\

\

\

\

\

h-_ -~

"- "-

0--0 NUMBER

__ WEIGHT 2.0

1.6

0.4

o L----;;O~.07'"1--o;!:.s;----;-ll.o::--..Jt::=-:'I.I:-S~~""'2J,;.O,...------:5:1:.0o--...-s::.l..0=--.....--:-:!IOL,.0,...--....Jo

PERCENT ORGANIC CARBON

Figure 201

Density and biomass of Brachiopoda in relation to sediment organic carbon.

Holothuroidea-Holothurians formed a moderate component of the New England benthos in terms of biomass, but made up only a minor portion of the total

number ofspecimens (Table 3). This was due mainly to

the relatively large size of individual specimens. Members of this group averaged 3 g each, which is a size

unsurpassed by any other major faunal group collected

in this study. The larger specimens, Cucumaria and

Molpadia, were 10 to 14 cm long, and 2 to 4 cm in

diameter. The smallest specimens were juvenile Psolus

about 4 mm in length.

Five orders of holothurians were represented in the

samples (Table 4). The dominant group, from the standpoint of abundance and taxonomic diversity, was the

Dendrochirotida. Few species and specimens were taken

belonging to the orders Molpadiida, Apodida,

Aspidochirotida, and Dactylochirotida.

Color of specimens in this region was generally uniform over the body surface (except for the contrasting

light colored tube feet in some species), which usually

was reddish-brown, tan, or occasionally light cream.

One small species was very dark violet and black.

Holothurians occurred in 202 samples (19% of total). Their density averaged 4.3/m2 and their biomass

averaged 12.9 g/m2 (Table 5).

Geographic Distribution

Holothurians were prevalent over large sections of the

Gulf of Maine and in the offshore Southern New England

Shelf area (Fig. 208). They occurred on Georges Bank in

low densities and in very few samples. Relatively high

densities (25 to 88 individuals/m2) occurred over rather

large portions of the central Gulf of Maine and along the

southern end of Great South Channel. In most localities,

however, the average densities were less than 9 dividuals/

m2• In terms of biomass, holothurians were present in

substantial quantities, usually greater than 1 g/m2, and

not uncommonly in quantities of 10 to over 50 g/m2.

Among the six standard geographic areas, holothurians were present in largest quantities in the Gulf of
188 NOAA Technical Report NMFS 140

OJ

III 50- 99

.100-751

GRAMS

PER SQUARE

...

". ". 70·

M A IN E

...

ECHINODERMATA

Figure 202

Geographic distribution of Echinodermata: A-number ofspecimens per square

meter of bottom; B-biomass in grams per square meter of bottom.

Maine area (Tables 6, 8; Fig. 209), where the average

density was nearly 8 individuals per1m2 and the biomass averaged more than 27 g/m2. The Georges Bank

area had the lowest average density, 0.2/m2, and an

average biomass of only 0.5 g/m2. The other four areas

had intermediate quantities.

Holothurians accounted for a small proportion of

the total number ofspecimens in all areas, but made up

a substantial part of the total weight in two areas (Table

9). In the Gulf of Maine area they accounted for 22% of

the total faunal weight and on the Southern New England

Slope they made up 14% of the total faunal weight.

The occurrence of holothurians in the samples was

moderately low (17 to 40%) in all areas except Georges

Bank, where they were present in only 2% ofthe samples.

They were present in a slightly higher proportion of the

samples from the two slope areas than from the shelf

areas (Table 10).
Composition and Distribution of Macrobenthic Invertebrate Fauna 189

0

ICIlI-

~O

«CD

ll:lI...

ClO

Z ll:

LU l- I- I LU Cl

LU LU

ll:

« I- ::l LU 0

CIl

ll:

LU

Q.

o

o

20

00

20

60

40

GEORGES SOUTHERN

SLOPE NEW ENGLAND

SLOPE

GULF OF

MAINE

NOVA

SCOTIA

I

0 NUMBER

• WEIGHT

-

-I

.--

-8

I--

I-- -

I--

-

f--

- I-- .-- .--

• I o GEORGES SOUTHERN

BANK NEW ENGLAND

SHELF

GEOGRAPHIC AREA

Figure 203

Density and biomass of Echinodermata in each of the six geographic areas.

20

120

40

140

o

lI- CIlO

Z CD 100 LU

~lI...

0 0

LUll: Q. LU CIl I- 80

lI... LU O~

ll:LU

LUll:

CD«

~:l 60 :l0 ZCIl

ll:

LU

Q.

,.....- ---;; --,200

40

160

0---{) NUMBER

__ WEIGHT

'0-- --\\

\

\

\

\

\

\

o 10 100 200 ~OO 1000 2000 4000 0

WATER DEPTH IN METERS

Figure 204

Density and bomass of Echinodennata in relation to water

depth.
190 NOAA Technical Report NMFS 140 _

90

80

o

o

20

70

80 e

;b

c ID

a:,,-

"'0

!lO ! a:

~~'" :z:'" ",2

40

c ~:::> ",a .'"a:

'"

...

GRAVEL TILL SHELL SAND SANO SILT SILT CLAY

o HUMBER

• WEIGHT

-

-

0,- - ,

l -

1 -

0- -

0- -

- -I

I • -

4

20

o

12

~lil ...

2"-

(j 0 100

~~

"'~

~~

a:... 8 "'a: IDe

2:::>

:::>0

Z'"

a: 6

'"...

BOTTOM SEDIMENTS

Figure 205

Density and biomass ofEchinodermata in relation to bottom sediments.

Bathymetric Distribution

Holothurians were collected at depths ranging from 6

to 3,820 m, and were present in all depth classes over

this broad depth range (Table 11; Fig. 210). Densities

were highest (average 4 to 10 individuals/m2) at intermediate depths (50 to 500 m) and somewhat lower (0.7

to 1.9/m2 ) in both shallower and deeper bathymetric

classes.

Biomass distribution of holothurians differed substantially from the depth-density relationship (Table

13; Fig. 210). Highest biomass averages (13 to 37 g/m2)

were found in depths less than 100 m. Lowest biomass

(0.2 g/m2) occurred at 500 to 999 m; intermediate

quantities (l to 6 g/m2 ) were found in other depth

classes.

Individual holothurians from shallow water (0 to 24

m) were larger, averaging nearly 25 g each, and size

decreased with increasing depth to less than I g each at

depths greater than 500 m.

The frequency ofhoiothurian occurrence was higher

in samples from deep water than in those from shallow

water (Table 15). At depths less than 50 m they were

present in 8 to 9% of the samples, whereas, in water

depths greater than 50 m they occurred in 14 to 33% of

the samples.

Relation to Sediments

Holothurians were relatively numerous in till substrates

and much less common in all other bottom types (Table

16; Fig. 211). Their average density in till was 25 individuais/m2, whereas, in the other types of sediments

their density was only 2.0 to 7.4/m2.

The relationships of holothurian biomass to sediments were entirely different from those pertaining to

density. The biomass was high (25 to 29 g/m2) in sandsilt and silt-clay (Table 18; Fig. 211). In fact, holothurians accounted for from 15 to over 33%, respectively, of

the total benthic biomass in those two sediment types

(Table 19). In other types of sediments their biomass

was moderate to small (4.7 to 0.4 g/m2).

The presence ofrelatively fewer but larger specimens

in soft sediments and numerous small specimens in till

sediments accounts for the disparity between the biomass and density values in these substrates. In the other

sediment types they were generally more equally distributed in density and biomass.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 191

r------------------------,280

240

200 ::IE

0

Il/ll-

::lEO

«ID

a: a.. 160 ClO

za:

-w

1-1-

X W Cl::IE

120 WW

~a:« 1-::> wO

~l/l

a:

80 W

ll.

40

L.-.---;;--7;~-~:;:;----;;B;--I~.1.~9---;1~2~.1-:;-5.~9-7.16:--7.19~.9~-::2~0--;.2!:-:3:"':.9:---..J

RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

180

160

::IE

0

II- 140 l/l0

ZID

w

::lEa..

u O 120 wa: ll.w

l/lla.. w o::IE 100

a:w

wa:

ID« ::IE::> 80 ::>0 Zl/l

a:

w

ll. 60

40

20

0

0---0 NUMBER

__ WEIGHT

Figure 206

Density and biomass of Echinodermata in relation to the annual range of

bottom water temperature.

Frequency of occurrence was moderate to low in all

sediment types, and the relationship of holothurians

with various sediments was similar to that described for

numerical density (Table 20). Occurrence was highest

(50%) in till, lowest (8%) in sand, and intermediate

(16 to 36%) in the other sediment types.

Relation to Water Temperature

The relation of holothurian numerical density to the

range in water temperature was the opposite of that

exhibited by their biomass. Density generally decreased

as the temperature range increased (Table 21; Fig.

212). The average density of holothurians was about 6

individuals/m2 where the temperature range was less

than 4°C and decreased to about 2 individuals/m2 where

the temperature variation was 16° to 19°C.

Conversely, the biomass of holothurians increased as

water temperature ranges increased (Table 23; Fig. 212).

Their biomass was relatively small (3 g/m2) where the

temperature range was small «4°C). With increased

annual temperature range, the biomass of holothurians increased substantially to an average of 128 g/m2

where the range was greatest (20 0-23.9°C).

Size disparity of specimens in the various temperature range classes was the principal cause of the reversed trends. The size of specimens increased markedly in waters having a broad temperature range,

whereas the numerical density diminished slightly.

Holothurians contributed a moderately large proportion (4-9%) of the total biomass in five of the six

temperature range classes. In the other class, where the

temperature variation was high (>20°C), they contributed an extraordinarily large share (>30%) of the biomass (Table 24).

Frequency of occurrence was quite uniform at a moderate level (14-25%) in all temperature range classes

except one (16°-19.9°C), in which holothurians contributed only 8% of the samples (Table 25).
192 NOAA Technical Report NMFS 140

100

0---4 NU_

_ WlIIHT

100 100

/\

// \ 10 tI' \ 100

/ \ Ii e~

/ \ .0 I~ c· / u \ I~ fll: / \ ZII: e'" / \ -Ill

~I .0 I 100 ~~z:'"

II / \ !!-

\ "'''' / .~

/ \ tij II: \ •

f / \ II:

40 / 100 f

/

,J \

\

\ 10 \ 100

-0..... '-

oL-...-:.-::-!:':----::::-----:':;----:':::----::!';;.--*--o:f;;---.::..e.........,~-----'o 1.0 1.5 2.0 5.0 5.0 10.0

P[I'C[NT OftGANIC C""ION

Figure 207

Density and biomass of Echinodermata in relation to sediment organic carbon.

Relation to Sediment Organic Carbon

Holothuroidea was one of only a very few taxonomic

groups for which definite, consistent trends were clearly

demonstrated in relation to the amount of organic

carbon in the sediments.

The trend was the reverse of that pertaining to water

temperature. The organic carbon-density relationship

was in general negatively correlated, whereas the organic carbon-biomass relationship was essentially positively correlated. Seven of the eight carbon content

classes were occupied; the one exception was the highest class (5.0+%).

Holothurian average density was highest (18 individuals/m2) where no measurable organic carbon was

found (Table 26; Fig. 213). Much lower average densities (ranging from 8 to 21m2) prevailed in the other

carbon content classes with a general tendency of decreasing with increasing organic carbon content.

Average biomass, on the other hand, showed a trend

that was the reverse of the one for density (Table 28; Fig.

213). Average biomass was lowest (6 g/m2) in the 0%

carbon content class and steadily increased with increasing organic carbon content, culminating in exceptionally large biomasses in the two highest classes occupied. There was nearly a threefold increase between

the 1.50 and 1.99% class and the 2.00 and 2.99% class

(41 vs. 104 g/m2 ) and a fivefold increase from the latter

class to the 3.00-4.99% class (562 g/m2 ).

Frequency of occurrence was quite uniform at a moderate level (15 to 40%) in all organic carbon content

classes (Table 30).

Echinoidea-Sea urchins are the second largest (after

bivalves) contributors to the New England benthic biomass, providing 20% of the total (Table 3). This large

contribution was made by a group with low taxonomic

diversity. Fewer than six species contributed over 95%

of the specimens. The major contributors were sea

urchins, heart urchins, and sand dollars.

The feeding habits of echinoids are varied. Most are

bottom feeders (carnivores, herbivores, or omnivores),

but some common species are plankton feeders, and
___________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 193

12"

12°

NEW

JERSEY

EXPLANATION

[ill 1-9

II 10-24

• 25-88

EXPLANAT ION

[ill <0.1 - 9.9

liliiii 10.0- 49.9

• 50.0- 379.8

GRAMS

PER SQUARE METER

12"

HOLOTHUROIDEA

.,0 .."

.."

Figure 208

Geographic distribution of Holothuroidea: A-number of specimens per square meter

of bottom; B-biomass in grams per square meter of bottom.

the distinction is not always maintained. When the preferred food types are unavailable, echinoids may may

revert to other food sources, or in some species, to a

different mode of feeding.

Echinoids in turn are preyed upon by a variety of

benthic and nektonic animals. They have been observed

in the diet of crabs, starfish, finfish, lobsters, birds, and

mammals, including man. The quantity utilized for

human consumption in recent years has increased with

an annual harvest worth several million dollars.

Coloration of most of our echinoids was rather drab

consisting largely of grays, brown, reddish-browns, and

brownish-violet. In many species the color was gray or

brown with suffusions of white, green, pink, or violet.
194 NOAA Technical Report NMFS 140

30

o NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA .....INE BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELf StOPE

GEOGRAPHIC AREA

o NUMBER

• WEIGHT

-2

B- -2

6'- -I

- -I

2- -

.. ...- • I o

Figure 209

Density and biomass of Holothuroidea in each of the six geographic areas.

2:

40 fflb

2:al

cl ... ffio 30 z a:: -w

1-1-

J:w

,,::li:

20 - w Wa::

~<l

I-~o 10 ffl

a::

w

ll.

0-- -0 NUMBER

--. WEIGHT

R

I \ I ,

I '

I \ I ,

I ,

I ' I \

, \ ....--6 \\

,

\0-_---0-""

25 50 100 200 500 1000 2000 4000 0

WATER DEPTH IN METERS

Figure 210

Density and biomass of Holothuroidea in relation to water depth.

Sizes ranged from 2 mm in diameter, in juvenile specimens, to large adults having a test diameter of 5 em. The

majority ofspecimens ranged between 0.5 and 3 em.

Echinoids occurred in 293 samples (27% of the total). Their density averaged 29.3/m2 and their biomass

averaged 36.8 glm2 (Table 5).

Geographic Distribution

Echinoids occurred extensively throughout most of the

study area (Fig. 214), but their distribution pattern was

somewhat patchy and irregular in density. Intermediate and high densities (10 to 1,051 individuals/m2 )

were common from Georges Bank to the Southern New

England Shelf areas. The biomass of echinoids was

especially high (100 to 1,027 g/m2) on Georges Bank

where they made up 50% of the total benthic biomass

in Northeast Channel, and in coastal areas of central

and Southern New England (24%).

Among the six standard geographic areas, the average density of echinoids was moderately high (over 100

individuals/m2) on Georges Bank, intermediate (4 to

221m2 ) in the Nova Scotia, Gulf of Maine, and Southern ew England Shelf areas; and low (0.2 and 0.3/m2)

in the two slope areas (Table 6; Fig. 215).

Biomass was moderate to very high in all areas. It was

especially high (averaging 117 g/m2) in the Georges
Composition and Distribution of Macrobenthic Invertebrate Fauna 195

2~

0 NUM8ER

• WEIGHT

:Ii

0

Ienl- 30 :Ii 0

«Ill

0:"-

"'0

2~ zo:

-",

1-1-

20 :1:'" ",:Ii

"'''' ~o:

I~ « 1-::;)

~g

10 0:

'"

CL

0

BOTTOM SEDIMENTS

Figure 211

Density and biomass ofHolothuroidea in relation to bottom sediments.

140

<>---0 NUM8ER

-... WEIGHT 120

:Ii :Ii 0 0 l- II- 100 en I- eno ~O ZIIl «al

'" 0:: ... 2'"

;:;0 "'0

"'0: 80 ~o:: CL", w en I- 1-1-

... w :z;w o~ 8 "'~

o::w 60 Ww wo:: ~o::

1Il« 6

'" cr :Ii::;) 1-::> ::;)0

" 40 w O Zen "'" ~lf)

0:: 4 'o.-----o-.. __ 0::

w wCL CL

20

0 0 4-1.9 8-11.9 12·1~.9 16-19.9 20-23.9

RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 212

Density and biomass of Holothuroidea in relation to annual range of

bottom water temperature.

Bank area. Moderately high (13 to 33 g/m2 ) biomasses

of echinoids were found off Nova Scotia, in the Gulf of

Maine, and on the Southern New England Shelf. Lowest biomass averages occurred in the slope areas (Table

8; Fig. 215).

Echinoids were present in only moderate to small

percentages of the total number ofsamples. Frequency

of occurrence was moderate (19 to 51 %) in the four

continental shelf areas, and low (8 to 9%) In the two

slope areas (Table 10).

Bathymetric Distribution

Echinoids occurred in water depths ranging from 7 to

2,950 m. Average densities were highest (127 individuals/m2) near mid-shelf depths (25 to 49 m) and diminished in both shallower and deeper regions (Table 11;
196 NOAA Technical Report NMFS 140 _

.------------------------------,800

100

800

400 too

CIl too

SOcal

ac ... 1:)0

~acIII

300~ l:i

!!!S

III III

.acc too:>

III 0

.CIl

ac 200

q

\

\

\

\

\

\

\

\

\

\ ....... \ ./ \

\ /F \

iT/ \

\

\ ....... A ....... "- --~ ""- oL....a:=='~~:::::::x;:=-___:"J,,_-___:J,,_-__=J,,_-__=:l:__-__=:l:__~"-~+,...--..J 0 0.01 O.!l 1.0 I.!l 2.0 3.0 !l.0 10.0

PERCENT ORGANIC CARBON

!l.0

O-~ NUMeER

- WEIGHT

Figure 213

Density and biomass of Holothuroidea in relation to sediment organic carbon.

Fig. 216). At depths greater than 100 m, their density

was drastically reduced (2.0 or fewer individuals/m2).

The pattern of biomass distribution was quite similar

to that for density. Average biomass was exceptionally

high (154 g/m2) near mid-shelf depths (25 to 49 m)

and decreased in both shallower and deeper regions

(Table 13; Fig. 216). The decrease was only moderate

in the shallower depth class but was severe (0 to 1.8 g/

m2) in depths greater than 500 m.

Occurrence of echinoids in the samples was moderate to low in all depth classes and followed the same

trend as density and biomass. They were present in 57%

of the samples near the mid-shelf depths and decreased

in samples from both shallower and deeper bathymetric classes (Table 15).

sity averaged 67 individuals/m2 in sand but only 3.4 or

less per square meter in other sediments.

Precisely the same pattern was revealed for biomass.

The average biomass in sand sediments was 81 g/m2; in

all other sediments the biomass of echinoids averaged

7.3 g or less/m2 (Table 18; Fig. 217). The proportion of

the total benthic biomass that was formed by echinoids

in the sand sediments was 33%, which is an exceptionally large contribution for one taxon (Table 19).

The occurrence of echinoids in the samples was moderate to low in all sediment types (Table 20). As

expected, it was highest (47%) in sand substrates. Somewhat unexpectedly, the incidence rate was lowest

(8%) in sand-silt sediments and relatively high (32%)

in till.

Relation to Sediments

The correlation of echinoids with sand substrates was

exceedingly high (Table 16; Fig. 217). Although they

were present in all other types ofsediments, their denRelation to Water Temperature

Echinoids were most abundant in terms of both density

and biomass in areas where the annual range in bottom

water temperature was moderate, 12° to 15.9°e.
Composition and Distribution of Macrobenthic Invertebrate Fauna 197

72°

72°

NEW

JERSEY

EXPLANATION

W 1- 9

if] 10- 99 C1!])

• 100-1051

INDIVIDUALS

PER SQUARE METER

EXPLANATION

[] <0.1 - 49,9

50 0 - 99 9 r---..

.. 100',0- 10~7,0<.J

GRAMS

PER SQUARE METER

72°

""

~"'-J\ ,

•

B

40°

ECHINOIDEA

71°

4,°

70°

M A IN E

69°

~~~120KM

42°

Figure 214

Geographic distribution of Echinoidea: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom,

Echinoid density averaged 93 individuals/m2 at midrange (Table 21; Fig. 218). From this peak the average

density diminished to 1.4/m2, where the temperature

range was less than 4°C, and to about 15/m2 where the

temperature change was 20°C or more.

The biomass of urchins was distributed in essentially

the same manner as their numerical density among the

various temperature range groupings, with one exception (Table 23; Fig. 218). Large biomass (148 g/m2)

occurred in the mid-range; moderate biomass (about

14 to 38 g/m2) in intermediate ranges; and small biomass (6 g/m2 ) in the stable areas. The one exception was

a relatively large biomass (135 g/m2) in the broadest

(200-23.9°C) temperature range class.
198 NOAA Technical Report NMFS 140

o

::I:

o

... Ul'"

::1: 0

00 < ell a:,,-

ClO

za:

- .... ......

x .... CI::I:

........

~a:

< ... ::;)

~g

a:

....

20

o

20

40

80

GEORGES SOUTHERN GEORGES SOUTHERN

BANK NEW ENGLANO SLOPE NEW ENGLANO SHELF SLOPE

GEOGRAPHIC AREA

GULF OF

MAINE

NOVA

SCOTiA

0

0 NUMBER

• WEIGHT

Of-

-I

01-

-I

,CI-

-

-6

0-

-

Of-

-

I I I 0 - •

6

2

8

4

12

10

Figure 215

Density and biomass of Echinoidea in each of the six geographic areas.

140r-----------------------~

160

0-- -0 NUMBER

_ WEIGHT

10 25 50 500 1000 2000 4000 0

20

120

::I:

~51°O

.... eIl

~~.... a: 80 11. ....

Ulll5~6

ffi~ m<l

::I:::::l

g 40

a::

....

a.

WATER DEPTH IN METERS

Figure 216

Density and biomass ofEchinoidea in relation to water depth.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 199

80..------------------------,

o NUMBER

• WEIGHT

80

60

0 0 VJ ~.

l/lo ~o

zm ot m .... 0::1&- ~I&- 60 0 0 (30

.... 0:: ~o::

ll. .... .... l/l~ ~~

40 :1: .... 1&- .... O~ O~

0:: .... ........ .... 0:: "0::

mot ot 40 ~:::l ~:::l .... 0 :::lo Zl/l "l/l

0:: 0::

-1"

.... .... ll. ll.

20

BOTTOM SEDIMENTS

Figure 217

Density and biomass of Echinoida in relation to bottom sediments.

The proportion of the number of individuals in the

total macrobenthos made up by urchins was low to

moderate (0.3-5%, Table 22), but in terms of biomass

they contributed extraordinarily large amounts (8-47%,

Table 24) of the total.

Echinoids were present in 14 to 44% of the samples

in the various temperature range classes. Frequency of

occurrence, as for density and biomass, was highest in

the mid-range classes and decreased in both the broader

and narrower range classes (Table 25).

Relation to Sediment Organic Carbon

Echinoids showed a marked preference for low levels of

sediment organic carbon and were entirely absent where

levels of 2% or more prevailed. Their density was highest, but only at moderate levels (24 individuals/m2), in

the lowest carbon content class (0.01-0.49%) and fell off

drastically where organic carbon was absent (2.6/m2), as

well as in the classes between 0.50 and 1.99% where

density ranged from only 0.6 to 11m2 (Table 26; Fig. 219).

Biomass was similarly distributed among the carbon

content classes. The lowest content class contained

nearly 28 g/m2 and biomass values fell off markedly in

adjacent classes (Table 28; Fig. 219).

Frequency of occurrence of echinoids in the samples

was generally low, ranging between 6 and 34%, and

paralleled the trend exhibited by density and biomass

measures (Table 30).

Ophiuroidea-Ophiuroids formed a moderately small

butsignificant component ofthe New England benthos.

On the average they made up about 3% of the number

of animals and 2% of the biomass in the total

macrobenthos.

Two orders ofophiuroids (Euryalae-basketstars, and

Ophiurae-brittlestars) inhabit the study area. The

basketstars have a limited distribution within the region

and are relatively rare. Brittlestars, however, are widely

distributed and form the bulk of all ophiuroids in our

collections. They have the greatest diversity of species

of all echinoderm groups; more than a score ofspecies

have been identified within the region.

The size of brittlestars in our samples ranged from

large specimens of Ophiomusium, with disc diameters of
Geographic Distribution

Brittlestars occurred over approximately three-fourths

of the study area (Fig. 220). Their average density over

most of their range was between 1 and 49 individuals/

m2• High densities (100 to 680/m2) were widespread

along the outer continental shelf south of Nantucket

Shoals. Brittlestars were absent from large portions of

central Georges Bank, Nantucket Shoals, and much of

the New York and NewJersey region.

Biomass distribution of ophiuroids tended to parallel

their density distribution. Moderate (1-10 g/m2 ) and

large (l0-80 g/m2 ) biom,sses were widespread off

Southern New England on the outer continental shelf,

and in the eastern Gulf of Maine.

The average density of ophiuroids was moderate to

moderately high in all six standard geographic areas

(Table 6; Fig. 221). Highest average density (94/m2 )

Ophiuroidea occurred in 487 samples (45% of the

total). Their density averaged 44.2/m2 and their biomass averaged 3.26 g/m2 (Table 5).

200 NOAA Technical Report NMFS 140

100

O--~ NUMBER

- WEIGHT

BO 160

20 20 0 0

cno cn~

20 0 Zal

a..! «al

20LL 60 120 a:LL

u O ClO

a..! a: lLa..! ~a:

cn~ a..! ~~ LLa..! :z;a..! 020 Cl20

a: a..! a..! a..! a..! a: ~a: al« 40 80 « 20::> ~::> ::>0 a..! 0 Zcn ~cn

a: a: a..! a..! lL a.

20 40

o'----;:;:h;---4A:-:I77."9---:8;;-:.-hII~.9-"712;;---h15;-;.9:--"716;:--+'19;-;:9:---;:"20;:---!:-2=-3.=-9 ----10

RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 218

Density and biomass of Echinoidea in relation to the annual range of bottom

water temperature.

about 25 mm and arm lengths over 12 cm, to small

specimens of Amphipholis and Ophiura, with disc diameters of less than 2 mm and arms 10 mm long.

The coloration exhibited by ophiuroids in oursamples

was varied. Some genera, such as Ophiura and

Ophiomusium, were uniformly whitish or light gray. Others were more colorful because of their mottled patterns of contrasting hues, including dark red, pink,

brown, and orange. Among the more brightly colored

genera were Ophiopholis, Amphiura, and Ophioscolex.

Ophiuroids obtain their food by a variety of different

feeding methods; feeding types represented in our

samples were carnivores, detritus feeders, filter feeders,

and omnivores. A large share of the New England species generally combines the ingestion of bottom material with selective carnivorous feeding. The diet thereby

consists of detritus, diatoms, and other small-size foods,

as well as polychaete worms, crustaceans, bivalve mollusks, and other similar types of organisms. Brittlestars,

in turn, are preyed upon by other echinoderms, but

most significantly by demersal fishes.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 201

o-~~R

_WfIGHT 50.0

1.0

10.0

tlI.o

eo.o;c·

11: ... ClIO

!~

......

:z;w

11.0 a

.~

iiII:

II

1\

I \

I \

I \

I \

I \

I \

\

\

\

\

\

\

\

I \

I \

\

\

b----ool--.......-O'='O!:-:I---to.a::----:I;.I;.O.---:....--;I:I:.lI--~I.O~ ..~I.~o-""""":IIt..O....-·IO~.O....- .....o

HItCENT OMANIC CMIlOM

1.0

10.0

20.0

Figure 219

Density and biomass of Echinoidea in relation to sediment organic carbon.

occurred on the Southern New England Shelf, and a

moderately high density (30/m2 ) occurred in the Gulf

of Maine. In the four remaining areas the density was

moderate (14 to 17 individuals/m2) and about equal.

The biomass of ophiuroids, also, was relatively uniform

from one area to another; total range was 0.8 to 5.4 g/m2

(Table 8; Fig. 221). Relatively large biomasses (3.3 and 5.4

g/m2) were encountered on the Southern New England

Shelf and in the Gulf of Maine. Smallest (0.8 g/m2)

biomass was on Georges Slope. Although the average

ophiuroid biomass on the Southern New England Slope

was 2.6 g/m2, an intermediate quantity, the proportion of

the total fauna it made up was 13.5%, a much higher

proportion than that for any other area (Table 9).

Frequency of occurrence ofthese organisms was moderately low (22 to 35%) in the samples from Georges

Bank and, surprisingly, on the Southern New England

Shelf. Ophiuroids were present in 55 to 64% ofsamples

from all other areas (Table 10).

Bathymetric Distribution

Ophiuroids were taken at depths ranging from 13 to

3,820 m. Their density distribution revealed a pronounced zone of high abundance (35 to 87 individuals/m2) at depths between 50 and 500 m (Fig. 222).

Lower densities (0.8 to 6.2/m2) prevailed in both deeper

and shallower water. The lowest density occurred in the

shallowest depth zone, 0 to 24 m.

The biomass of ophiuroids was more uniform among

the various depth classes than was density; however, the

same general trend was clearly evident (Table 13; Fig.

222). Biomass was relatively large (2.5 to 7.5 g/m2) at

depths between 50 and 500 m, and smaller in both

deeper and shallower water.
202 NOAA Technical Report NMFS 140

NEW

JERSEY

EXPLANATION

EJ 1- 49

II 50- 99

.100-680

lTI]<0.1- 0.9

mil 1.0- 9.9

• 10.0- 80.3·

GRAMS

PER SQUARE METER

OPHIUROIDEA

Figure 220

Geographic distribution of Ophiuroidea: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

Occurrence of ophiuroids was low (9 to 10%) in

samples from the inner continental shelf, at depths less

than 50 m (Table 15). At depths of 50 to 500 m, where

ophiuroids were most abundant, their occurrence in

the samples was substantially higher, 40 to 72%. In

water deeper than 500 m, ophiuroids were present in a

slightly higher proportion ofsamples (44 to 76%). This

indicates that these organisms were more uniformly

distributed at a lower density in deepwater regions than

they were in shallow water.

Relation to Sediments

Ophiuroids were rather plentiful in all types of bottom

sediments, but trends in density and biomass in the
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 203

o

o 6 Ien I-

~O

<l:lD

a::u.

ClO

za::

-ILl

4 tJ Cl~

ILl ILl

~a::

<l: 1-::>

ILl 0

~en

2 ffi

ll.

GEORGES SOUTHERN

SLOPE NEW ENGLANO

SLOPE

GEORGES SOUTHERN

BANK NEW ENGLANO

SHELF

GEOGRAPHIC AREA

GULF OF

MAINE

NOVA

SCOTIA

r- 0 NUMBER

-WEIGHT

Of-

-

Of-

-

Of-

-

O~

- r0 I

10

6

o

lI- (/)0

ZlD

ILl

~u.

U 0 6

ILIa:: ll.1LI

(/)1-

u. 1LI o~

ffi 4 lD<l:

~::>

::>0

zen

a::

ILl

ll. 2

Figure 221

Density and biomass of Ophiuroidea in each of the six geographic areas.

different types were evident. Densities were low

(16 to 26 individuals/m2) in gravel, sand, and shell;

intermediate (38 and 581m2) in till and silt-clay; and

high (941 m2) in sand-silt (Table 16; Fig. 223).

The trend of biomass in relation to sediment

type was nearly the same as that revealed by density. Small biomasses of ophiuroids occurred in

gravel, sand, and shell; intermediate quantities

were found in silt-clay; and largest biomasses (5.3

and 5.8 g/m2) occurred in till and sand-silt (Table

18; Fig. 223).

Occurrence of ophiuroids in the samples revealed a pattern similar to those of both density

and biomass. They were present in a relatively

small proportion (29-40%) of the samples from

gravel, shell, and sand, and they were present in a

substantially larger share (62-68%) ofthe samples

in till, sand-silt, and silt-clay (Table 20).

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

0---0 NUMBER

- WEIGHT B

2; B I 2; 0 0 l- I I-

~b / \ 6 Ul b ",Ill 2; III

2; .... ? \ <I ....

0 06 ~o

"'1I: I \ ZlI: CL",

\ Ull- -'"I-

.... '" I 1-'" 02; \ 4:J:2i

lI:~ 40 I \ !2",

"'lI:

~~ I /, ;J<I

2;0 1-::>

~Ul I \ ",g

\ ;J lI: lI: .... I .... CL \ CL

20 I \

I \

h----0-----

00 10 25 50 100 200 500 1000 2000 4000

Relation to Water Temperature

Ophiuroid density, biomass, and frequency of occurrence all conformed generally to the same

trend of high abundance where the temperature

range was less than 16°C, and low abundance

WATER DEPTH IN METERS

Figure 222

Density and biomass of Ophiuroidea in relation to water

depth.
204 NOAA Technical Report NMFS 140

o SAND· SILTSILT CLAY

GRAVEL TILL SHELL SAND

0 NUMBER

• WEIGHT

- -

-

-

f-

-

fo I

80 o

lIlO

zm101

~lL

U 0 60

lola:: Q. w lIl~

lL W

O~

ffi 40

lIJet

~::;)

::;)0

Zlll

a::

101

Q. 20

100

BOTTOM SEDIMENTS

Figure 223

Density and biomass of Ophiuroidea in relation to bottom sediments.

Figure 224

Density and biomass of Ophiuroidea in

range of bottom water temperature.

0'-----"0,--'.3ic.9;;------:;4,.-'.7"'.g;-----;;-e-+.II;-;.9;----""1;;-2.-!-;15;-;.9;----~~=~~,.------...l0

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

0---0 NUMBER

--.. WEIGHT

relation to the annual

density declined in both lesser and greater amounts of

organic content, with the lowest density (51m2 ) occurring in the highest content class they occupied (2.00-

2.99%).

Biomass showed a similar trend but at considerably

lowered magnitude (Table 28; Fig. 225). Biomass was

from 5 to 6 g/m2 in areas of organic carbon content

between 0.5 and 1.49% and fell offsharply to consider-

::IE

en b eo ZlIl

::IE'" '"

u O

60

en~

...'" 0::IE 40

a:",

"'a: lIler

::IE::>

g 20

a:

'"

0..

where the range was greater than 16°e. Average density of ophiuroids increased from 25

to 74 individuals/m2 as the range in temperature increased from less than 4°e to 12°-

15.9°e, then dropped precipitously to less than

31m2 in the two broadest temperature range

classes (Table 21; Fig. 224).

Biomass values were an order of magnitude

lower than those of numerical density, but

they revealed a general decline associated with

an increased range in temperature (Table 23;

Fig. 224). Where the temperature range was

less than 12°e, the average biomass was about

3 to 6 g/m2. Where the temperature range

was greater than 16°e, the biomass averaged

about 0.5 g or less/m2.

Frequency of occurrence diminished rather

consistently with increased range in water temperature (Table 25). Ophiuroids occurred in

64% of the samples in areas where the temperature range was less than 4°e and declined steadily

to 11 % in areas where the range in temperature was

200e or more.

Relation to Sediment Organic Carbon

Ophiuroids were numerically most abundant (72 to 81

individuals/m2) where the organic carbon content was

between 0.50 and 1.49% (Table 26; Fig. 225). Average
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 205

100

/~ ,/ \

\

/ \

/ \

/ \

/ \

/ \

/ \

/ \

l

/ \

/ \

/ \

I \\

0--0 NUMBER

--- WEIGHT

B.O

o

.... 6.0 fI) ....

~o

e CD

a::~ C)o

~a::ILl

4.0 i

C)~

~~c .... 5 ~fI) 2.0 a::

ILl

CL

o L-----1L--=-l:::7"""--~0.~5----:-I.I:.0:---~I:-I:.5:----2::.1.'='0=....~3t:.0-..---:5J.,.0,..--..~1-!0-=.o---Jo

PERCENT ORGANIC CARBON

Figure 225

Density and biomass of Ophiuroidea in relation to sediment organic carbon.

ably lower levels (0.1-2 g/m2 ) in classes on either side

of these.

Occurrence of ophiuroids in the samples was moderate to moderately high (66-70%) in the two mid-range

content classes and in the absence of measurable carbon, and moderately low (23 to 47%) in the other

content classes they occupied (Table 30).

Asteroidea-Starfishes constituted a moderately small

portion of the New England benthos. Their contribution to the total number of specimens was only 0.1 %,

but they provided 1.2% of the total biomass (Table 3).

Representatives of three orders of the Asteroidea

were present in our samples: Phanerozonia, Spinulosa,

and Forcipulata. All three orders contained species that

were common in the samples.

Starfishes exhibited considerable variation in color, both

inter- and intraspecifically. The most brightly colored species encountered was Porania insignis, which commonly

was a deep, rich claret color with patches of whitish and

yellowish papillae. Other specimens of the same species

and otherspecies within the same genus were much paler,

even a drab brownish-gray. Other colorful genera encountered were Hippasterias and Solaster. Many of the more

common genera, such as Asterias, Astropecten, and

Ctenodiscus, were predominantly tan, brown, or olive.

Sizes ranged from juvenile specimens of various species that were 2 to 3 mm in radius to large specimens of

Asterias and Solasterwith radii of over 18 cm. Specimens

with radii over 8 cm were uncommon.

Average weight of individual specimens was 1.4 g,

which was large compared with other taxonomic groups.

The majority of starfishes in this region are carnivores, particularly the selective, predatory type of carnivore. Deposit feeding and filter feeding are adaptations of

only a few species, some of which are locally abundant.

Bivalve mollusks appear to be the principal food of New

England starfishes, although a variety of small invertebrate species, as well as dead fish, are consumed. A few

species ofstarfishes are serious predators ofoysters, clams,

mussels, and other commercially valuable mollusks.

Asteroids occurred in 144 samples (13% ofthe total).

Their density averaged 1.5/m2 and their biomass averaged 2.1 g/m2 (Table 5).
206 NOAA Technical Report NMFS 140

Geographic Distribution

Starfishes had a moderately broad areal distribution in

the study area (Fig. 226). They occurred in an especially large proportion of the samples from offshore

Southern New England and the southern part of

Georges Bank but were noticeably absent or sparse in

the central Gulf of Maine, central Georges Bank, off

NewJersey and along most of the continental rise.

Asteroid density was generally low throughout the

ew England region, averaging only 1 to 191m2. The

slightly higher densities of 10 to 191m2 occurred only

in the coastal and near-coastal zones. Biomass of starfishes was relatively high compared with their numerical abundance, and although they averaged less than 10

g/m2 over most of their range, there was a substantial

number of localities where the average biomass was

between 10 and 105 g/m2 . The rather high biomass (10

to 50 g/m2 ) along the outer margin of the continental

shelf off Southern ew England corresponds to the

distribution ofsand in that area (see Fig. 7).

Differences in starfish density among the six standard geographic areas was moderate-extremes ofmean

density were 0.2 and 2.5/m2 (Table 6; Fig. 227). Densities ofstarfishes were generally higher in the continental shelf areas than they were on the continental slope.

Indications of a north-south trend were revealed by a

low density in the Nova Scotian shelf area, intermediate

values in the Gulf of Maine and Georges Bank region,

and relatively high density in Southern New England.

Biomass distribution was similar to density. There

were only two geographic areas in which the biomass

was unusually small (0.02 and 0.05 glm2), the two slope

areas (Table 8; Fig. 227). Relatively high biomasses (1.0

to 4.5 g/m2) occurred in the continental shelf areas.

Disparity in the average size of starfishes from the

different areas was substantial. For example, in the

Nova Scotia area the average weight ofindividuals specimens was 1 g, whereas in the slope areas, they averaged

only 0.1 g each.

The frequency of occurrence ofstarfishes was low (6

to 19%) in all geographic areas (Table 10).

Bathymetric Distribution

Asteroids occurred at depths ranging from 13 to 2,329

m, and at a mean depth of 184 m. They occurred in

substantially higher densities (1.8 to 2.5/m2) at depths

between 25 and 200 m than at other depths (Table 11;

Fig. 228). In very shallow water (less than 25 m) and in

depths beyond the continental shelf their density was

0.1 to 0.7/m2. Densities were generally lowest in the

deepest water. The maximum density of 167/m2 was

encountered at a depth of 49 m.

Starfish biomass varied considerably from one depth

class to another, but the distributional pattern was similar to that described above for density (Table 13; Fig.

228). The average biomass on the continental shelf was

3 to 4 g/m2 (except in the 25-49 m depth class where

the biomass was unusually small). In depths below 200

m, the biomass averaged 0.04 g or less per square meter

but was slightly higher on the lower continental slope

and continental rise than on the middle and upper

continental slope.

The proportions of samples in the various depth

classes that contained starfishes ranged from 5 to 20%

(Table 15). Their frequency of occurrence was slightly

higher between 50 and 200 m than at other depths.

Relation to Bottom Sediments

Starfishes were taken in all sediment types except shell,

and there were marked differences in quantity between

samples from fine-texture sediments compared with

those from coarse sediments (Table 16; Fig. 229). Largest quantities were from fine-grain sediments.

Average density in the coarse substrates (till and

gravel) was 0.2 and 0.8/m2, respectively. In the finer

substrates (sand, sand-silt, and silt-clay) asteroids averaged 1.3 to 1.8/m2.

Asteroid biomass was 0.5 g or less per square meter in

the coarse sediments and 2.0 to 3.6 g/m2 in the finegrain sediments.

Starfishes occurred in a markedly higher proportion

(14 to 16%) of samples from sediments composed of

fine particles than of samples composed of gravel and

till (8 and 9%) (Table 20).

Relation to Bottom Temperature

Asteroids were found in moderate to low amounts in all

temperature range classes (Table 21; Fig. 230), but they

were more abundant where the temperature range was

moderate rather than extreme.

Average densities (0.4 to a.9/m2 ) of starfishes in the

extreme temperature range classes (0°_3.0° and 16°-

23.9°C) were considerably below those found in the midranges (4°-15.9°C) where values of 1.2 to 2.7/m2 occurred. In terms of total faunal density, asteroids provided only 0.2% or less of the total number ofspecimens

in each of the six temperature range classes (Table 22).

The average biomass of starfishes in the intermediate

temperature range classes was quite stable, varying only from

2.8 to 3.6 g/m2; the two extreme classes, however, yielded

only 0.1 to 0.5 g/m2, respectively (Table 23; Fig. 230). Starfishes in the various temperature range classes represented

from 0.1 to 4% of the total faunal biomass (Table 24).

The proportion of samples that contained asteroids

was lowest (6--4%) in the two extreme temperature

range classes and the highest (21 %) in the 8°-11.9°C

temperature range class (Table 25).

Relation to Sediment Organic Carbon

Asteroids were present in only four of the eight organic
Composition and Distribution of Macrobenthic Invertebrate Fauna 207

...

...

NEW

JERSEY

EXPLANATION

Llli r7:l I 9 -

11IIII10-19

EXPLANAT ION

[ill <0.1 - 9.9

11IIII 10.0- 49.9

• 50.0 - 105.3

GRAMS

PER SQUARE METER

,..

ASTEROIDEA

70· .,.

Figure 226

Geographic distribution of Asteroidea: A-number of specimens per square meter

of bottom; B-biomass in grams per square meter of bottom.

carbon content classes. The four classes in which they

occurred ranged from 0.01 to 1.99%. Average density

was fairly low but relatively uniform in the three classes

between 0.01 and 1.49% carbon content, ranging from

1.6 to 1.0 individuals/m2 (Table 26; Fig. 231). The 1.50

to 1.99% class, however, contained only 0.4 individuals/m2.

The distribution of biomass was parallel to that of density, highest (5.5 g/m2) in the 1.00-1.49% carbon content

class and dropping off significantly in higher and lower

adjacent content classes where values were uniformly low,

ranging from 1.03 to 1.42 g/m2 (Table 28; Fig. 231).

Frequency of occurrence of asteroids in the samples

in the organic carbon content classes ranged from 14 to
208 NOAA Technical Report NMFS 140 _

Bathymetric Distribution

Acorn worms occurred in three depth range

classes: 50-99 m, 100-199 m, and 500-599

m, where their mean density ranged from

<0.1 to 0.5 individual/m2 and their mean

biomass was 0.01 g/m2 in the two deepwater

ranges and 0.05 g/m2 in the mid-shelf range

(Tables 11, 13; Fig. 233). Five percent of the

samples in the 500-999 m depth range

yielded specimens, whereas the occurrence

in the other two was from <1 to 1% (Table

15). The depth range of our samples W.1S

from 79 to 567 m.

Hemichordata

Geographic Distribution

The geographic distribution of acorn worms

was restricted to the Southern New England

Shelf (3 stations) and Southern New England

Slope (l station). Mean densities were 0.3

individual/m2 or less and mean biomass 0.06

glm2 or less (Tables 6 and 8; Fig. 232). Frequency of occurrence of hemichordates in

the samples was a very low 1% (Table 10).

Representatives ofthe phylum Hemichordata

are all from one class (Enteropneusta, acorn

worms) and one genus (Balanoglossus). A total of 101 specimens, weighing 18.7 g, were

collected at 4 stations (0.4% of total); average density (0.1 individuals/m2 ) and biomass

(0.02 g/m2) were very low (Table 5).

17% in the three classes between 0.01 and

1.49%, and was 7% in the 1.50 to 1.99% content class (Table 30).

:IE

o

.... "' .... s

"'LL

4 '" 0 Z",

-- .... ........ 3 " .... ~:IE

~~« .... ::> .... 0

~'"

'"....

Q.

o

:IE

o

.... 4",1-

:::EO «00

",LL

"'0

3 Z '" -- .... .... t;:;

ili:::E 2- ....

....

~«'" .... 5 I~'"

'"....

Q.

0---<:> NUMBER

_ WEIGHT

500 1000 2000 4000 0

NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

0 NUMBER

31- • WEIGHT -

--

21--

--

- II--

I n -I

I n

Figure 227

Density and biomass of Asteroidea in each of the six geographic

areas.

Figure 228

Density and biomass of Asteroidea in relation to water depth.

r--~----------------___,5

WATER DEPTH IN METERS

Figure 229

Density and biomass ofAsteroidea in relation to bottom sediments.

Relation to Bottom Sediments

Acorn worms were found in low abundance

in sand, sand-silt, and silt-clay substrates.

Mean density ranged from <0.1 to 0.2 individual/m2 and mean biomass from <0.01 to

0.04 g/m2 (Tables 16, 18; Fig. 234). Frequency of occurrence in the samples was <1

to only 1% (Table 20) .

Relation to Bottom Temperature

Hemichordates were restricted to areas where

the annual range in water temperature was

less than 16°C. Mean density and biomass

were very low «0.l/m2 and <0.01 g/m2) in

the lowest temperature range class; mean densities of 0.2/m2 occurred in each of the two

:::E

o

.... "' .... ~~

"'LL

"'0

4 '"

.... ........ " .... "':::E

.... .... 2 '"« .... ::> .... 0

~'"

.... '"Q. o SAND· SILTSILT CLAY

SHELL SAND

BOTTOM SEDIMENTS

GRAVEL TILL

0 NUMBER

• WEIGHT

-

I-

- If11. ill

:::E

o

....

.... "'0 Zoo

....

:::ELL

0 0

~ffi "' .... LL.... 2 O:::E

"' .... .... al«'" :::E::>

::>0

Z'"

'" o
___________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 209

2

0

fofo0

III

IL

0

a:

'"f0-

IL'"

0:E

a:",

"'a: 1Il<t

2:> :>0 z'"a:

'"

a.

D

0---0 NUMBER

-. WEIGHT

Figure 230

Density and biomass ofAsteroidea in relation to the annual range

of bottom water temperature.

.--- --,6

0---0 NUMBER

_ WEIGHT

OL---4I----:~--....,O:!-..5:--------:-1 '=.O-----;-'1.5;;------;2;';.O;---'....----=3"".O~.-....,5:':.O;;-~.~-;cIO:!c.O=----.JO

PERCENT ORGANIC CARBON

Figure 231

Density and biomass of Asteroidea in relation to sediment organic carbon.

::I ::I 4 0 0 l- I- ml- ml- ::1 0

ffi~ e ID

a::lL ::IlL ClO 13 0

za:: 1&1 a::

ll;~ -1&1

3 1-1-

lLI&I :rl&l

0::1 C)::I

a:: 1&1 1&11&1

1&1 a:: ~a::

IDe e ::Ij 1-;:)

;:)0

Zm ;&1 a:: a:: 2 1&1 2 lL

range classes between 8 and 15.9°e. Mean biomass ranged

from 0.03 to 0.04 g/m2 (Tables 21, 22; Fig. 235). Frequency of occurrence was in the <1 to 1% range in the

three range classes (Table 25).

Relation to Sediment Organic Carbon

Distribution of hemichordates was limited to the three

organic carbon content classes between 0.01 and 1.49%.

Mean densities were between 0.1 and 0.2 individual/m2
210 NOAA Technical Report NMFS 140 _

WATER DEPTH IN METERS

Figure 233

Density and biomass of Hemichordata in relation to water depth.

Geographic Distribution

Ascidians were distributed over large portions of the study area (Fig. 237), but

their occurrence was patchy, with density

changing abruptly from one locality to

another. They were most common in the

coastal areas and on the offshore banks

and were sparse or absent from the following areas: the deeper part of the western Gulf of Maine, north-central Georges

Bank, and parts of the continental shelf

off Connecticut, New York, and New JerAscidiacea-Ascidians were moderately

common in the New England benthos and

constituted 1.1 % of the total number of

specimens and 2.2% of the total biomass

(Table 3).

The size ofsolitary tunicates in our collections ranged from 4 mm to more than

20 cm. Although colonial forms are known

to attain lengths greater than 50 cm in

this region, the specimens in our samples

were smaller (12 cm or less in length)

than the largest solitary forms.

Interspecific variation in color of New

England tunicates was rather broad, and

in some groups the intraspecific variation was also considerable. The majority

ofspecies were dull olive, tan, or brown.

A rather large number covered their tests

with sand grains or silt that very effectively camouflaged them. A few groups

contained beautifully colored species.

Their coloration consisted of a blending of cream and light yellow with various shades of orange and red.

In order to control their rapid colonization on hard substrates and fouling of

man-made structures, such as ship bottoms, pipes, buoys, and similar structures,

the maritime industry incurs high economic costs. Ascidians occurred in 181

samples (17% of the total). Their density

averaged 16.3/m2 and their biomass averaged 4.1 g/m2 (Table 5) .

and mean biomass between <0.01 and 0.03

g/m2 (Tables 26, 28; Fig. 236). Only 1%

of the samples in the three carbon content classes contained specimens of acorn

worms (Table 30).

Chordata

::Ii

o

.. ~~<lOO

0: ...

"'0

20:

-",

....

I'" ",::Ii

10.110.1

~a:

<l .. ::> 10.10

~(J)

a:

10.1

Q.

:::li

o

.. (J)"

:::li0

<[lD

a: ... "'0

2a:

-10.1

.. .. IW

~:::li

Ww

~a:<[ .. :::> Wo

~(J)

a:

10.1

Q.

0.04

o

.. .06 <1)5

~CD

<[

a:'" ClO

za: 0.04 - w

.. .. ",

I::li

CI

-wwa:

~<[

002 .. ::> ",0

~C/l

0:

W

Q.

- 0.04

- 0.02

- 0.06

o NUMBER

• WEIGHT

o NUMBER

• WEIGHT

(}-- -0 NUMBER

_ WEIGHT

BOTTOM SEDIMENTS

1\

/ \

/ \

/ \

/ \ / \ / \

/ \

oOL.-.--'10-4----'25~W--'5-0-...L-...L-¥-l'--1-00LO--'W-2-.JOOLO-400...LO--J0

::E

o

.... (J)O

200

'"::E ... <:;0

~o:

(J)'"

lL. O.3~ o::li

0:'"

UJ Q:: 02f- OO<l

::E:::>

:::>0

Zoo Qlf- n

o,'------cc~-~__=__:c:____===_:_:,.".__::_:'_~'="::____oc~=___='I"""'" NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN .~---JO SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

Figure 234

Density and biomass of Hemichordata in relation to bottom sediments.

::li

0

.... (J)O

2lD

W

:::li'"

u O

Wa: Q. w (J) ..

... w o:::li 0.2

o:w ",a:

lD<[

:::li::>

::>0 01 2(J)

a:

wQ.

0

Figure 232

Density and biomass of Hemichordata in each of the six geographic

areas.
Composition and Distribution of Macrobenthic Invertebrate Fauna 211

::IE

0

I-

",I-

::lEO

... '" a: ... 0.04 "0

za:

-w

0.03 1-1-

:t:'" ,,::IE

0.02 w'"

~a:

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0.01 ",0

~'"a:

0 '"

Q.

0---<> NUMBER

___ WEIGHT

II

I

'I

f

4·7.9 8-11.9 12-15.9 16'19.9 20-23.9

RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

::IE

0

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'"

0

a:

w

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a:",

wa:

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a:

w

Q.

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Figure 235

Density and biomass of Hemichordata in relation to the annual

range of bottom water temperature.

I

0.2110

0.01l0

(0.0321

J.

,---<>\

I \

I \

I \

I

I

I

I

I

I

I

I

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I

I

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0--0 NUMBER

__ WEIGHT

0.01

0.008

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00

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o'--.....---;0~.0::71---;0:':.1l;-----;-1.';;-0---;';1.11;--".--;;2:';;.0;-- ....--;'3.';;'0-.--;1l;':.0;;--....---;;10t:.0;;------' 0

PERCENT ORGANIC CARBON

Figure 236

Density and biomass of Hemichordata in relation to sediment organic carbon.

sey. Densities throughout most of their range averaged

less than 50 individuals/m2. In a few localities, however,

their density averaged between 100 and 900/m2. Their

biomass was moderate (less than 10 g/m2) over most of

their range and large quantities (50 to 288/m2) were

encountered in several different areas.

The average density of ascidians was moderately high

(34 and 27 individuals/m2) in only two of the six stan-
212 NOAA Technical Report NMFS 140

EXPLANATION

1m 1- 49

• 50- 99

.100-905

EXPLANAT ION

[J <0.1 - 9.9

III 10.0 - 49.9

• 50.0 - 248.4

GRAMS

PER SQUARE METER •

'0°

ASCIDIACEA

.,. '0·

69°

Figure 237

Geographic distribution of Ascidiacea: A-number of specimens per square meter of

bottom; B-biomass in grams per square meter of bottom.

dard geographic areas, Georges Bank and the Southern New England Shelf (Table 6; Fig. 238). In these two

areas the biomass also was large (5.1 and 8.4 g/m2)

(Table 8; Fig. 238). In the four other areas their average density was low (2.8 or less/m2), and the biomass

was relatively small (0. I to 2.6 g/m2). Biomass was especially small (0.11 and 0.17 g/m2) in the two slope areas.

Ascidians occurred in a moderately small proportion of

samples (12 to 23%) and were present in approximately

the same proportion ofsamples from all areas (Table 10).

Bathymetric Distribution

Ascidians occurred at water depths ranging from 13 to

3,080 m, (Table 11; Fig. 239). Relatively high densities
Composition and Distribution of Macrobenthic Invertebrate Fauna 213

WATER DEPTH IN METERS

Figure 239

Density and biomass of Ascidiacea in relation to water depth.

Figure 238

Density and biomass of Ascidiacea in each of the six geographic

areas.

o

o

8

~~.. 1O

a: .... "'0

za:

-",

1-1-

x'"

"'~

4 "''''a:

.. 1-::>

",0

~'"a:

'"

Cl.

NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN

SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND

SHELF SLOPE

GEOGRAPHIC AREA

I

0 NUMBER

• WEIGHT

-

r-

-6

f-

-

f---

-

II I r1. Il

10

0--0 NUMBER

___ WEIGHT

35

't--- 8

30 I 0 0 l- I- I-

~b / "'0

/ ~ID WID

.... 25 :LL I -0 6 ClO u

wa:

Q. / zffi w -I- "'I- 20 / I- W ....

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/ ~::> 1-0 ::>0 w'" z'"a: I ~'" '" 10 d w Q. Q.

o

40

25 50 100 200

Frequency of occurrence of ascidians was moderately

low in all temperature range classes. Their occurrence

rate, however, was somewhat high (17 to 24%) where

the temperature range was moderate, and lower (11 to

12%) where the temperature range was very small and

very large (Table 25).

30

I-

"'0 ZlO

'" .... <:>0

~~en.... 20

o~.... '"

a:'" "'a: 1O .. ~::>

::>0

Z",

e::i 10

Cl.

(6 to 32 individuals/m2 ) were encountered on

the continental shelf; low densities (0.1 to 2.71

m2) occurred on the continental slope and

continental rise.

Biomass of ascidians, also, was larger on the

continental shelf than in deep water beyond

the shelf (Table 13; Fig. 239). Average biomass

at water depths less than 200 m ranged between 1.0 and 8.9 g/m2, whereas at depths

greater than 200 m the biomass averaged less

than 0.7 g/m2•

Ascidians were present in the samples from

nearly all depth classes at about the same low

range of 5 to 24% (Table 15).

Relation to Water Temperature

Although the trends of ascidian density and

biomass relative to the annual range in bottom

water temperature were not consistent, generally the quantities were larger where the temperature range was moderate, and smaller at

both temperature extremes (Tables 21, 23; Fig.

241). In areas where the temperature range

was moderate, from 8° to 19.9°C, the average

density of ascidians ranged from 6 to 591m2

and the biomass ranged from 7 to 14 g/m2.

Where water temperature ranges were extreme, less

than 8°C and more than 20°C, the average density was

low (2-4/m2 ) and the biomass was small (0.5-4.1 gl

m2). Two temperature range classes (8°-11.9°C and

16°-19.9°C) contained significantly greater quantities

than did the other classes.

Relation to Sediments

Ascidians occurred in all six types of bottom sediments; however, their density and biomass varied

substantially from one type to another (Table 16;

Fig. 240). Densities were relatively high (29 and

221m2) on gravel and sand bottoms, and intermediate (111m2) in sand-silt. Densities were low

(5.1 or less/m2) in till, shell, and silt-clay.

Biomass of ascidians followed the same trend

as density, but the differences were less pronounced (Table 18; Fig. 240). Average biomass

was largest (9.7 to 4.4 glm2) on gravel and

sand, in termediate in sand-silt, and small «2

g/m2) in the remaining sediment types.

Frequency of occurrence in the samples was

moderately low (11 to 32%) in all types of

bottom sediments (Table 20). In the majority

ofsediment categories, the small differences in

occurrence rate are correlated directly with ascidian density and biomass. Shell and till values, which are based on a small number of

samples, are incongruous.
214 NOAA Technical Report NMFS 140

2

o

o

2

2

o B I- 11I1-

20

4 11I

11: .... 00

ZII:

-lU

6 i:;;

02

lUlU

~~

1-;;)

~~ 4 II:

lU

Q.

SAND· SILT

SILT CLAY

SHELL SAND

BOTTOM SEDIMENTS

GRAVEL TILL

I

0 NUMBER

• WEIGHT

-I

0- -

!If-

-

Of-

!If- -

Of-

-

!If-

- r -

II:lU

lUll:

11I4 :IE;;)

;;)0

ZIIl

II:

lU

Q.

o

:IEe 3

I- 11I0

ZIII

lU

:IE ....

0 0 2

~ffi 11I1-

.... lU 0:IE 2

Figure 240

Density and biomass of Ascidiacea in relation to bottom sediments.

Relation to Sediment Organic Carbon

A general trend of diminishing density and biomass

with increasing sediment organic carbon content was

exhibited by New England region ascidians (Tables 26,

28; Fig. 242), and lhey were restricted to the low to midrange carbon content classes between 0.01 and 2.99%.

Mean density ranged from 21 to 4.4 individuals/m2,

and mean biomass from 4.4 to 0.2 g/m2.

Frequency of occurrence in samples was low and

paralleled the trend ofdensity and biomass, ranging from

23% to 8% as carbon content increased (Table 30).

Dominant Components of the

Macrobenthos

This section identifies and defines the dominant faunal

constituents of the New England region macrobenthos

and their relationship to each of the abiotic parameters

considered in the treatment of each taxonomic group in

the preceding sections. "Dominance," as used in this report, refers to the taxonomic group that mathematically

contributed the highest number ofindividuals or greatest

total accumulated wet weight. Results are expressed in

both measures of abundance because of the marked differences that existed between them.

In spite of individual disparity in rank order within

each measure of abundance, members of four taxonomic groups, collectively, made up the bulk of the

macrobenthic invertebrate fauna of the New England

region. The four major taxonomic components are 1)

Annelida; 2) Mollusca, comprising Bivalvia, the chiefcomponent, as well as Gastropoda, Scaphopoda, Polyplacophora, and Cephalopoda; 3) Crustacea, with

Amphipoda the chief component of this group, followed

by Cumacea, Isopoda, Decapoda, Cirripedia, Mysidacea,

Tanaidacea, Ostracoda, and Copepoda in progressively

smaller proportions; and 4) Echinodermata composed of

Ophiuroidea, Echinoidea, Holothuroidea, Asteroidea, and

Crinoidea in diminishing proportions. Table 3 lists the

contributions of each ofthe above taxa to the total density

and biomass of the ew England macrobenthic fauna.
Composition and Distribution of Macrobenthic Invertebrate Fauna 215

0--0 NUMBER

60 _ WEIGHT 16

0 0 l- I- ",l- I- 50 "'0 ~o ZcD <cD ... 12 lrl&. ~l&. C>o ijO

"'lr 40 :!:lr 0.. ...

j\ ... "'I- 1-1- l&. ... :I:'" o~ 30 L \ B C>~

lr'" ...... "'lr \ ~lr

cD< < ~~ \ I-~

~o 20

/ ... 0 Z", \ ~'" lr / 4 lr ... ... a.. 10 a..

I

-----J 0 0-3.9 4-7.9 B-II.9 12-15.9 16-19.9 20-23.9 0

ANNUAL RANGE IN BOTTOM WATER TEMPERATURE

IN DEGREES CELSIUS

Figure 241

Density and biomass of Ascidiacea in relation to the annual range of

bottom water temperature.

20.0

0--0 NUMBER

__ WEIGHT 0.0

o L----;~---;o.to;-----;-1.1;;:0----;.l;1.1I;--.....~2!;;.0:---~5.1;;0~i--;lI;1:.0;;--.....--::10!-:.0;;-----J0

PERCENT ORGANIC CARBON

20.0 4.0

\ 2 I 2 e I 0

l- I-

~i I w ... 2 0 1&1 <cD

~~IO.O I 5.0

rQ: I !Q: en 1&1

... I 1&1

l&. ... ... ... 02 I :I: 1&1

!!!2

;~ \ 1&1 ... .Q:

2~ \ < ig 10.0 2.0 ... &

Q: \ ~en

... I Q:

Q. 1&1

Q.

I

\ 0.0 \ / / 1.0

V

Figure 242

Density and biomass of Ascidiacea in relation to sediment organic carbon.
216 NOAA Technical Report NMFS 140 _

Frequency of Occurrence

Geographic Distribution

Figure 243

Percentage composition of the dominant components of the macrobenthic

invertebrate fauna expressed in terms of number of individuals and wet

weight of the total fauna.

NUMBER WEIGHT

Among the four dominant taxa,

Annelida was the most ubiquitous

in distribution, occurring in 96% of

all stations sampled (Table 5). Next

in order were members of Mollusca,

which occurred in 88% of all

samples, followed by Crustacea in

85% of the samples; echinoderms

ranked fourth with a 72% occurrence rate.

For comparative purposes, some

nondominant taxa showed intermediate frequencies; among these

were Coelenterata, which occurred

in 42% of the samples, and Nemertea and Sipunculida with frequencies of 34% and 23%, respectively.

Ascidiacea, Bryozoa, Aschelminthes, Porifera, Brachiopoda, Pogonophora, Turbellaria, and Hemichordata were encountered with

diminishing frequencies ranging

from 17% to <l %.

Percentage Composition

The four dominant macrobenthic components collectively contributed more than 90% of the total number

of individuals, and 90% of the accumulated wet weight

of the macrobenthos sampled within the New England

region. The relative contribution and ranking of the

four, however, differed markedly between the two measures of abundance (Fig. 243).

Crustacea, with Amphipoda providing the single largest amount ofindividuals, was the highest ranking taxonomic component in terms of density, accounting for

nearly one half (48%) of all individuals within the

region. Annelida ranked second, providing slightly more

than 28% of the total number of individuals encountered; Mollusca and Echinodermata followed, contributing somewhat more than 12% and 5% of the total

number of individuals, respectively.

The biomass ranking was nearly the reverse of that

for density. Mollusca, with Bivalvia predominating,

ranked first in biomass versus third in density, contributing 46% of the total weight. Echinodermata ranked

second versus fourth; Annelida ranked third versus second; and Crustacea, first in density, ranked fourth in

biomass.

The differences in rank order between the two measures of abundance are attributable to size differences

within the various taxonomic groups.

The geographic distribution of density and biomass of

each of the four dominant components of the

macrobenthos of the New England region is depicted

in Figures 69, 99,146, and 202. Within the subareas of

the region, the rank order of the four dominant taxa

varied in terms of density and biomass (Figs. 13B, 14B;

Tables 6, 8). In the Nova Scotia subarea, Annelida

ranked first in density (648/m2) but third in biomass

(19 g/m2 ); Crustacea was second in density (329/m2 )

but fourth in biomass (17 glm2); Mollusca, third in

density (771m2 ), was first in biomass (54 g/m2 ); while

Echinodermata fourth in density (241m2 ), was second

in terms of biomass (39 g/m2 ).

In the Gulf of Maine, mollusks (306/m2) replaced

annelids (291m2 ) as the dominant taxon in terms of

density, the latter occupying second place, whereas

mollusks were second (32 g/m2) and annelids third

(16 g/m2), in biomass. Crustaceans (150/m 2) and echinoderms (431m2 ) ranked third and fourth, respectively,

in density but were fourth (2 g/m2) and first (56 g/m2),

respectively, in biomass.

On Georges Bank, the crustaceans (1052/m2 )were

nearly twice as abundant as the annelids (546/m2),

almost nine times denser than echinoderms (12l/m2),

and 22 times denser than mollusks (471m2). In terms

of biomass, however, the echinoderms (120 g/m2 ) outweighed mollusks (80 g/m2 ) by 1.5 times, and crustaceans (10 g/m2 ) and annelids (8 g/m2 ) by 12 and 15

times, respectively.
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 217

In the Southern New England Shelfsubarea, density

was dominated by crustaceans (1385/m2), followed by

annelids (531/m2 ), mollusks (244/m2 ) and echinoderms (123/m2 ) in terms of density. Mollusk biomass

(171 g/m2) was dominant in this subarea followed by

significantly smaller amounts of echinoderm (36 g/m2 ),

annelid (30 g/m2 ), and crustacean (17 g/m2) biomass.

The two continental slope subareas contained lower

densities and biomasses of all components than the

shelf subareas. The mean densities and biomass on

Georges Slope for the major taxa were as follows: Crustacea, 138/m2 and 0.6 g/m2; Mollusca, 831m2 and 2.7

g/m2; Annelida, 801m2 and 4.9 g/m2; and Echinodermata, 191m2 and 3.9 g/m2. Southern New England

Slope values were Annelida, 149/m2 and 4.3 g/m2;

Mollusca, 581m2 and 1.2 g/m2; Arthropoda, 221m2

and 0.1 g/m2; Echinodermata, 191m2 and 10 g/m2.

Selected Genera and Species-This section deals with

the geographic distribution of 24 selected genera and

species of macrobenthic invertebrates. These particular forms were selected because of their common occurrence, regional ubiquity, or distinctive distribution.

Figures 244 to 249 depict the distributions of the selected forms.

Phylum Annelida

Aphrodita hastala (Moore) (Fig. 244). Commonly

known as the sea mouse, this polychaete bristle worm of

the family Aphroditidae may attain lengths up to 150

mm (6 inch); smaller individuals are often found in

haddock, cod and red hake stomachs. It commonly

inhabits mud bottoms, or mixed bottoms with a high

mud content.

Scalibregma inflatum (Rathke) (Fig. 244). This medium-sized (1-5 cm) polychaete is a member of the

family Scalibregmidae. An important food of many demersal fish, this species inhabits silty sand substrates.

Sternaspis scutata (Renier) (Fig. 244). This moderately small (1 cm), burrowing polychaete is stout in

appearance and is a member ofthe family Sternaspidae;

it is found in the diet of winter flounder. It commonly

inhabits silty sediments.

Phylum Mollusca

Arctica islandica (Linnaeus) (Fig. 244). This rather

large (18-15 cm) commercially harvested bivalve, known

as the ocean quahog, mahogany quahog, or black clam,

belongs to the family Arcticidae. It is a very slow-growing species that occurs very abundantly in some localities on the continental shelf. Small tv medium-sized

individuals are preyed upon by cod and several species

ofstarfish. It usually inhabits muddy sand bottoms.

Astarte undata Gould (Fig. 245), the common wavy

astarte of the family Astartidae, is a medium-sized (2.5-

3.8 cm) bivalve. In the New England region, it is most

prevalent at mid-shelf depths (50-99 m) in sand and till

substrates. Although juvenile specimens are occasionally found in fish stomachs, it is not a major prey item of

demersal fishes.

Cerastoderma pinnulatum (Conrad) (Fig. 245), the

northern dwarf cockle, is a common, moderately small

(1 cm) bivalve belonging to the family Cardiidae. This

species is infrequently found in fish stomachs. It prefers

sandy substrates but does occupy, in lower abundance,

other types ofsediments.

Cyclocardia borealis (Conrad) (Fig. 245), the northern

cyclocardia, is a medium-sized (3-5 cm) bivalve of the

family Carditidae. Broadly distributed throughout the

region, it prefers sand and till substrates but does occur in

other sediments as well. It is not common in fish diets.

Modiolus modiolus (Linnaeus) (Fig. 245), the northern horse mussel, is the largest (5-15 cm) and most

common mussel of the offshore New England region; it

is a member of the family Mytilidae. It is found on the

periphery of the Gulf of Maine and on Georges Bank,

and extends onto the Southern New England Shelf. It

prefers sand and sand-shell substrates.

Placopecten magellanicus (Gmelin) (Fig. 246), the sea

scallop, is one of the most valuable commercial shellfish resources of the U.S. East Coast, especially in the

New England region. It is a large bivalve (12-20 cm) of

the family Pectinidae, found most abundantly on coarse

sandy bottoms. In addition to harvest by man, juveniles

of this species are found in the diets of some demersal

fishes, principally haddock, and ocean pout.

Buccinumspp. (Fig. 246) is represented in our samples

by four species in the family Buccinidae. The species

represented are B. elatius, B. gouldi, B. hydrophanum,

and B. undatum. Among the four species, the moderately large (5-10 cm) B. undatum, the waved whelk, is

overwhelmingly the most common form. They are typically found at mid- to lower-shelf depths in sand and

coarser grained sediments.

Neptunea decemcostata (Say) (Fig. 246), the wrinkled

whelk, is a moderately large (7-11 cm) gastropod belonging to the family Buccinidae. This species typically

inhabits hard bottoms ranging from coarse sand to

gravels at mid- to lower-shelf depths.

Phylum Arthropoda

Ampelisca agassizi (Judd) (Fig. 246), this gammaridean

amphipod of the family Ampeliscidae is a mediumsized (4-7 mm) tube dweller. It is the most abundant

and common species of amphipod in the southwestern

half of the study area; in some localities it is exceptionally

abundant. It prefers a sandy substratum. This species is a

very common prey in the diet of many demersal fish.

Leptocheirus pinguis (Stimpson) (Fig. 247), a species

of gammaridean amphipod, family Aoridae, is a moder-
218 NOAA Technical Report NMFS 140

..:...~!t'::

.. ...:.:

. .. Aphrodita hastata

..

.

•

(f:"

\

\

Scalibregma inflatum

Stemaspis scutata , Arctica islandica

Figure 244

Geographic distribution of three selected species of Annelida, and one species of Bivalvia.
Composition and Distribution of Macrobenthic Invertebrate Fauna 219

Astarte undata

,

.

.

•

.'

,

.

•

eye/acardia borealis (

•

•

,,:../

."

Modio/us modiolus

Figure 245

Geographic distribution of four selected species of Bivalvia,
220 NOAA Technical Report NMFS 140

(/ Buccinum sp.

"

t

•.i;

AmpeJisca agassizi

••••

•

.• • .-.-

• •• • ••••

.~~:/f . ~:

,

.

..

••

./

'" ./ ..../

Ii

( .., Neptunea d,,,emco"ala

Figure 246

Geographic distribution of one selected species of Bivalvia, one genus and one species of Gastropoda, and one

species of Arnphipoda.
Composition and Distribution of Macrobenthic Invertebrate Fauna 221

Leptocheirus pinguis

•

r UncioJa irrorata

Homarus americanus

Figure 247

Geographic distribution of two selected species of Amphipoda, and two of Decapoda.
222 NOAA Technical Report NMFS 140

,

.

Hyas coarctatus

..

.,- ••-7·,'/

.::J,I" Pagurus spp.

.. " .. •

Asterias vulgaris

.' ..

Girolana spp.

• •

•

.• •...• /~

... .. .. 'It ,-w I

, . • : • • 'ltei ·... , ' ..' , ••• • d ..... '--"""-----.: .... / . · .~.,-

.,-

Figure 248

Geographic distribution of one species and one genus of Decapoda, one genus of Isopoda, and one species of

Asteroidea.
Composition and Distribution of Macrobenthic Invertebrate Fauna 223

,

.

.

•

.<',<' ,<' •

Leptasterias spp.

Strongylocentrotus

drobachiensis ('

Echinarachnius parma,

.'

..

Ophiura spp.

Figure 249

Geographic distribution of one genus of Asteroidea, two species of Echinoidea, and one genus of Ophiuroidea.
224 NOAA Technical Report NMFS 140 _

ately large (10-17 mm) and abundant tube-dwelling

inhabitant of continental shelf sandy substrates. It is

very important in demersal fish diets.

Unciola irrorata (Say) (Fig. 247), another moderate-sized

(10-13 mm) tube-dwelling gammaridean amphipod of

the family Aoridae, is abundant in the sands of Georges

Bank and is also very important in demersal fish diets.

Crangon septemspinosa Say (Fig. 247), the sevenspined

bay shrimp, a moderately small (5-8 em) caridean

shrimp of the family Crangonidae. It is typically found

in sandy sediments of the region in both inshore and

continental shelf waters, and in certain localities is very

abundant. This shrimp is a very important prey to nearly

all demersal fishes.

Homarus americanus H. Milne-Edwards (Fig. 247), the

American lobster, is, together with the sea scallop, one

ofthe most commercially valuable invertebrate resources

ofthe northwest Atlantic. It is widely distributed throughout the New England region from inshore bays and

sounds to the offshore canyons bisecting the edge of

the continental shelf. Size of mature lobsters ranges

from around 25 em average length, for individuals captured for market by the inshore fishery, to very large

(sometimes in excess of 80 em), for specimens in the

offshore stock; minimum legal size for capture is currently 8.13 em (3.25 inch). Lobsters are scavengers and

inhabit a variety ofsubstrates.

Hyas coarctatus Leach (Fig. 248), the arctic lyre crab,

is a moderately small (to 31 mm) spider crab in the

family Majidae. This species is common throughout the

New England region on muddy and pebbly bottoms.

Small individuals have been reported to occur occasionally in the diet of long-horned sculpin.

Pagurus spp. (Fig. 248) comprise seven species of

pagurid hermit crabs resident in the New England region. The species of the genus Pagurus in the family

Paguridae represented in this study include P. acadianus,

P. annulipes, P. arcuatus, P. longicarpus, P. politus, P. pollicaris,

and P. pubescens. The represented species range from

small to medium in size (9-31 mm, carapace length), are

ubiquitous throughout the region in nearly all substrate

types, and are preyed upon by bottom fishes. The most

common and broadly distributed species is P. acadianus.

Cirolana spp. (Fig. 248), comprise three species of

the isopod crustacean family CiroJanidae resident in

the study area (c. concharum, C. impressa, and C. polita)

along with several others identified only to the generic

level. These moderately small (16-23 mm) crustaceans

are fairly common on muddy and sandy bottoms in the

Gulf of Maine and on Georges Bank. They are prey to a

variety of demersal fishes.

Phylum Echinodermata

Asterias vulgaris Verril (Fig. 248), the northern starfish or purple star, is one of the most common starfishes inhabiting the offshore waters of the New England region and is a member of the family Asteriidae.

This is a large species commonly between 15 em and 30

em (6-12 inch) in diameter; some specimens up to 42.5

em (17 inch) have been reported from the northern

limits of its range. It is normally found on sandy bottoms where it is a very important predator of bivalve

mollusks. Juvenile specimens are occasionally encountered in fish stomachs.

Leptasterias spp. (Fig. 249), which represent several

species of the genus Leptasterias, also of the family

Asteriidae, are common inhabitants of the New England Region on sandy bottoms. These brightly, but

variably, colored species are of moderate size (5-10

em) and in some localities are very abundant. Small

specimens are occasionally preyed upon by some species of groundfish.

Echinarachnius parma (Lamarck) (Fig. 249), the northern sand dollar, is the most abundant urchin (class

Echinoidea, family Scutellidae) of the New England

region; it is so abundant in some localities of Georges

Bank that the bottom resembles a mosaic pavement. As

its common name implies, it is a sand dweller. Sand

dollars ofthe region are typically 7.5 em in diameter. They

are a common prey offlounders, haddock and cod.

Strongylocentrotus droebachiensis (Muller) (Fig. 249),

another ubiquitous echinoid (family Strongylocentrotidea), the green sea urchin, is a hard bottom dweller

for whose popular roe a commercial fishery, inactive

since the 1930's and 1940's, is reemerging in northeastern U.S. and Canadian waters. Size ranges from 5 em to

nearly 9 em. Haddock and American plaice prey on this

spiny morsel.

Ophiura spp. (Fig. 249) comprise three species and

some undetermined specimens of this genus of brittle

stars (family Ophiolepididae) inhabiting the New England region; included are 0. lj"ungmani, 0. robusta, O.

sarsi, and unidentified species. Members ofthis group are

widely distributed and occur in most sediment types. Size

of the central disc ranges from 10 to 38 mm. They are

common in the diets of haddock and American plaice.

Bathymetric Distribution

In the New England region density and biomass of the

major taxa generally decreased with increasing water

depth (Tables 11, 13; Figs. 15, 16). Crustacea was the

dominant component of the fauna, in terms of density,

in shallow and continental shelf depths, ranging from

1,351 to 169 individuals/m2. Substantially lower densities occurred in waters deeper than 200 m. Annelida

had the next highest densities in shallow waters (7191

m2) and at continental shelf depths (519-437/m2).

Moderate numbers (241-107/m2 ) occurred at conti-
__________________ Composition and Distribution of Macrobenthic Invertebrate Fauna 225

nental slope depths, and low densities (9-30/m2) were

encountered at lower slope and upper rise depths. Mollusca showed a similar pattern of density distribution

with depth; density was greatest (570/m2 ) in shallow

water, moderate (l36-205/m2) at shelf depths, and

moderately low in deeper waters. However, mollusks

were numerically dominant at the deeper depths. Echinoderm density was greatest (l33/m2) at mid-shelf

depths between 25 m and 49 m, moderate (87-95/m2 )

in deeper shelf water, and decreased with increasing

depth beyond 200 m. Moderately low densities (471m2)

occurred in shallow (0-24 m) water.

Mollusca was the dominant faunal component of

biomass at nearly all depths. Greatest biomass (258 gl

m2) occurred in shallow water (0-24 m) with values

moderately high (132-21 g/m2 ) at shelf depths and

diminishing rapidly at depths below 200 m. Echinoderm biomass was greatest (167-106 g/m2 ) in waters of

49 m and less, decreased to 34 g/m2 at outer shelf

depths, but dominated at depths beyond 200 m (1.7-19

g/m2). Annelida biomass was highest (27 g/m2 ) in

shallow (0-24 m) water and nearly equal (25 g/m2) in

water depths between 50 m and 99 m. Values ranged

between 15 g/m2 and 16 g/m2 at other continental

shelfdepths but decreased rapidly with increasing depth

beyond 200 m. Crustacea, although numerically dominant, ranked fourth in biomass at nearly all depths

except the shallowest one (0-24 m) where a value of 37

g/m2 placed them third. Biomass of crustaceans ranged

from 2 to 16 g/m2 at continental shelf depths (25-199

m), with a rapid decrease from 4 g/m2 at 200-499 m to

approximately 0.1 g/m2 in slope and rise waters.

Relation to Bottom Sediments

Numerical abundance ofthe four dominant faunal components in relation to the six major sediment types

encountered in the New England region did not exhibit any trend as dramatic as that for depth (Tables 16,

18; Fig. 17). Annelids seemed to prefer sand (558/m2),

gravel (505/m2 ), and shell (443/m2) bottoms but were

moderately abundant in till, sand-silt, and silt-clay bottoms as well. Mollusks were generally more abundant in

silt-clay (354/m2 ), shell (229/m2), and sand-silt (2761

m2) but were found in somewhat lower abundance in

other sediments also. Crustacea were most abundant in

sand (l,336/m2), gravel (710/m2), and sand-silt (2751

m2) and were found in diminishing amounts in shell

(l24/m2 ), till (591m2 ), and silt-clay (341m2 ). Sand-silt

(l04/m2) and sand (951m2) contained the most echinoderms, followed by till and silt-day (671m2 and 651m2,

respectively), then shell (281m2) and gravel (231m2).

Density rank order in the various sediments listed by

decreasing particle size was as follows: gravel: Crustacea, Annelida, Mollusca, Echinodermata; till: Annelida,

Mollusca, Echinodermata, Crustacea; shell: Annelida,

Mollusca, Crustacea, Echinodermata; sand: Crustacea,

Annelida, Mollusca, Echinodermata; sand-silt: Annelida,

Mollusca and Crustacea equal, Echinodermata; silt-clay:

Mollusca, Annelida, Echinodermata, Crustacea.

The distribution of the biomass of the major taxa

among the various sediment types was fairly even. The

annelids showed the greatest uniformity with the smallest biomass (11 g/m2) in till and largest (26 g/m2) in

sand-silt. Biomass ranged from 15 to 16 g/m2 in the

four other types. Mollusks showed some variability, with

shell bottoms containing the largest biomass (168 gl

m2) and till the smallest (6 g/m2). Molluscan biomass

in gravel was 94 g/m2, in sand 121 g/m2, in sand-silt 74

g/m2, and in silt-clay 18 g/m2• Crustacean biomass was

20 g/m2 in gravel and 12 g/m2 in sand; 7 g/in2 and 6 gl

m2 in sand-silt and shell, respectively, and 2 g/m2 and

0.6 g/m2 in till and silt-clay ,respectively. Echinoderm

biomass was greatest in sand (88 glm2), 43 glm2 in siltclay, and 37 g/m2 in sand-silt. Median amounts occurred in till (15 g/m2 ) and lower amounts in gravel (6

g/m2 ) and shell (3 g/m2 ). Biomass rank order in bottom sediments was as follows: gravel: Mollusca, Crustacea, Annelida, Echinodermata; till: Echinodermata,

Annelida, Mollusca, Crustacea; shell: Mollusca, Annelida, Crustacea, Echinodermata; sand: Mollusca, Echinodermata, Annelida, Crustacea; sand-silt: Mollusca,

Echinodermata, Annelida, Crustacea; and silt-clay: Echinodermata, Mollusca, Annelida, Crustacea.

Relation to Water Temperature

Among the four dominant taxa there were no clear-cut

trends discernible with regard to the annual range in

bottom water temperature (Tables 21, 23; Fig. 18).

Where ranges of temperatures were between 8 and

19.9°C, Crustacea was the numerically dominant taxon,

with densities ranging from 768 to 1,475 individuals per

m2, whereas annelids dominated in areas exhibiting

rather stable annual temperature regimes, between 0

and 7.9°C (212-513 individuals per m2), and in areas

experiencing the broadest temperature range of 20-

23.9°C, where mean densities of 1,698 individuals per

m2 were found. Densities of Mollusca and Echinodermata were fairly consistent at moderate levels (84-3451

m2 for Mollusca and 21-171/m2 for Echinodermata)

throughout the temperature range spectrum. Mollusca,

however, did make a strong showing (1,242 individuals/m2) where the range in annual temperature was

broadest.

Rank order of dominance for the major taxa in the

six annual temperature range classes in terms of density was as follows: 0-3.9°C: Annelida, Crustacea, Echi-
226 NOAA Technical Report NMFS 140 _

nodermata, Mollusca; 4-7.9°C: Annelida, Crustacea,

Mollusca, Echinodermata; 8-11.9°C: Crustacea,

Annelida, Mollusca, Echinodermata; 12-15.9°C: Crustacea, Mollusca, Annelida, Echinodermata; 16-19.9°C:

Crustacea, Annelida, Mollusca, Echinodermata; 20-

23.9°C: Annelida, Mollusca, Crustacea, Echinodermata.

The relationship of the dominant taxa biomasses to

annual range of bottom water temperature was similar

to that of density in that no definite trends were evident. However, a marked change in dominance ranking prevailed, wherein the density dominants (crustaceans and annelids) were replaced by echinoderms and

mollusks as the leading contributors to biomass in nearly

all temperature range regimes. Echinodermata dominated biomass in four of the six temperature range

classes, including the narrowest and broadest ranges;

their mean biomass ranged from 12 to 263 g/m2. Mollusk biomass, second to that of echinoderms in most

temperature ranges, was clearly dominant where temperature ranges of 8-11.9°C and 16-19.9°C prevailed;

their mean biomass was 129 g/m2 in the former and

340 g/m2 in the latter. The contributions of the other

two dominant taxa, annelids and especially crustaceans,

due to their small size, were clearly subordinate in all

temperature regimes. Annelid biomass ranged from 10

to 40 g/m2, and crustacean biomass from 1 to 25 g/m2.

Rank order of dominance for the major taxa in the

six annual temperature range classes in terms of biomass was as follows: 0-3.9°C: Echinodermata, Annelida,

Mollusca, Crustacea; 4-7.9°C: Echinodermata, Mollusca,

Annelida, Crustacea; 8-11.9°C: Mollusca, Echinodermata, Annelida, Crustacea; 12-15.9°C: Echinodermata,

Mollusca, Annelida, Crustacea; 16-19.9°C: Mollusca,

Echinodermata, Annelida, Crustacea; 20-23.9°C: Echinodermata, Mollusca, Annelida, Crustacea.

Relation to Sediment Organic Carbon

As mentioned above (see section ''Total Macrobenthos")

there was no clear-cut correlation between sediment

organic carbon content and faunal abundance except

in a few exceptional cases (Tables 26, 28; Fig. 26).

The numerical abundance of the dominant taxa varied widely in relation to organic carbon content for all

except Echinodermata. This taxon ranked fourth in all

organic carbon content classes except two (0.00% and

1.00-1.49%) where it ranked third, slightly ahead of

Crustacea. Density of echinoderms was moderately low,

ranging from only 3-91 individuals per m2• Crustacean

densi ty varied widely among the various carbon conten t

classes, ranging from 21 to 1,357/m2. Greatest abundances occurred in carbon content levels of 0.01-0.49%

(1,066/m2) and 3.00-4.99% (l,357/m2), with moderately low to fairly high densities occurring in carbon

content levels between these two. Lowest densities (21-

221m2 ) prevailed in areas where no measurable carbon

existed as well as in areas where the greatest amounts of

carbon were measured. Mollusca, was one exception, showing a positive correlation of generally increasing density

with increasing carbon, ranging from 691m2 in areas

devoid of carbon to 1,120/m2 where carbon was between

3.0% and 4.9%. No mollusks occurred where carbon

content exceeded 5%. Annelida were present in all organic carbon content classes. Their density was significantly lower at both extremes of the carbon content spectrum (between 111m2 and 811m2) compared with their

abundance (l96-504/m2 ) in areas containing low (0.01-

0.49%) to moderate (2.0-2.99%) amounts of carbon.

Rank order of the numerical abundance ofthe dominant taxa with regard to organic carbon content was as

follows: 0%: Mollusca, Annelida, Echinodermata, Crustacea; 0.01-0.49%: Crustacea, Annelida, Mollusca, Echinodermata; 0.50-0.99%: Annelida, Crustacea, Mollusca;

Echinodermata; 1.00-1.49%: Mollusca, Annelida, Echinodermata, Crustacea; 1.50-1.99%: Mollusca, Annelida,

Crustacea, Echinodermata; 2.00-2.99%: Crustacea,

Mollusca, Annelida, Echinodermata; 3.00-4.99%: Crustacea, Mollusca, Annelida, Echinodermata; 5.00%+:

Crustacea, Annelida; no Mollusca or Echinodermata

were found in this class.

Similar to numerical abundance, biomasses of dominant taxa showed no clear-cut correlation to the organic carbon content of the bottom sediments. Most

notable was the considerable echinoderm biomass in

all but the highest carbon content classes, compared

with its low numerical density. Highest mean biomasses

(105 g/m2 and 562 g/m2) occurred in areas with moderately high carbon contents (between 2% and 4.99%),

and lowest (6 g/m2 ) occurred in areas devoid of measurable organic carbon. Moderate biomasses, ranging

from 23 to 44 g/m2, occurred in areas with low to

intermediate carbon content levels (0.01-1.99%). Mollusca, also absent where the highest measures of organic carbon occurred, nevertheless showed a preference for some organic carbon content, with highest

biomasses (812 g/m2 and 227 g/m2) occurring in the

two carbon content classes between 2.0 and 4.99%.

However, moderately high biomass (132 g/m2) was also

found where carbon levels were only between 0.01 and

0.49%. Lowest biomass (only 0.8 g/m2) occurred in

sediments devoid of carbon. Moderate levels of biomass (25-13 g/m2) occurred in organic carbon levels

that ranged from 0.50 to 1.99%. The mean biomass of

Annelida was fairly consistent at moderate levels ranging between 11 g/m2 and 27 g/m2 in areas of organic

carbon content ranging between 0.01 and 4.99%. Lowest mean biomass of annelids (0.11 g/m2) occurred in

the highest carbon content class (5+%) and intermediate amounts (7 g/m2) were found where measurable
Composition and Distribution of Macrobenthic Invertebrate Fauna 227

carbon was absent in the sediments. Crustacean mean

biomass ranged from a low of 0.11 g/m2 in areas of

highest organic carbon content to a high of 19 g/m2

where organic carbon was between 2% and 2.99%.

Moderately low biomasses (between 1 g/m2 and 9 g/

m2) occurred in the other carbon content classes. Areas

devoid of organic content also contained low mean

biomass (0.31 g/m2).

Rank order of the mean biomass of the dominant

taxa in terms of organic carbon content was as follows:

0%: Annelida, Echinodermata, Mollusca, Crustacea;

0.01-0.49%: Mollusca, Echinodermata, Annelida, Crustacea; 0.50-0.99%: Mollusca, Echinodermata, Annelida,

Crustacea; 1.00-1.49%: Echinodermata, Mollusca,

Annelida, Crustacea; 1.50-1.99%: Echinodermata,

Annelida, Mollusca, Crustacea; 2.00-2.99%: Mollusca,

Echinodermata, Annelida, Crustacea; 3.00-4.99%: Echinodermata, Mollusca, Annelida, Crustacea; 5.00%+:

Annelida and Crustacea were equal, whereas Mollusca

and Echinodermata were absent in this class.

Acknowledgments _

The authors are grateful to the many persons who

provided assistance in the various phases of this study.

We are especially indebted to Herbert W. Graham,

Robert L. Edwards, and K. O. Emery for their assistance

and support in the planning and organization of the

study. Northeast Fisheries Center personnel who assisted with the collection and processing of biological

samples included Bruce R. Burns, Gilbert L. Chase,

Philip H. Chase Jr., Evan B. Haynes, Henry W. Jensen,

Lewis M. Lawday, Arthur S. Merrill, Harriet E. Murray,

Clifford D. Newell, Timothy Robbins, Carol Schwamb,

and Ruth Stoddard Byron.

Appreciation is due the personnel of the NEFC ADP

unit for assistance in processing the voluminous numerical database generated by the study; Edward M.

Handy, Katherine Payne, Philip H. ChaseJr., Margaret

E. Cory, Johnny Blevins, and Francis W. Tinker, for

assisting with coding, data entry, programming, plotting, and data processing. Drafting assistance was provided by Frank A. Bailey, Herbert A. Ashmore, and

John R. Lamont.

Scientists from the U. S. Geological Survey and Woods

Hole Oceanographic Institution marine geology group

who provided sedimentological information or participated in shipboard work were K. O. Emery, John C.

Hathaway, Jobst Hulsemann, Frank Manheim, Robert

H. Meade, Richard M. Pratt, David Ross,John S. Schlee,

James V. A. Trumbull, and Elazar Uchupi.

Those who generously provided taxonomic assistance

were Edward L. Bousfield,John C. McCain, Edward B.

Cutler, Lion F. Gardner, Porter M. Kier, Peter Kinner,

Louis S. Kornicker, John M. Kraeuter, Don Maurer,

Arthur S. Merrill, Roy Olerod, David L. Pawson, Frank

Perron, Marian H. Pettibone, Thomas Phelan, Harold

H. Plough,Johanna Reinhart, Howard L. Sanders, Thomas]. M. Schopf, Eve C. Southward,]. H. Stock, Lowell

P. Thomas, Ruth D. Turner, Bertn Widersten, Austin B.

Williams, Lev A. Zenkevitch, and Victor A. Zullo.

We also wish to thank Marvin Grosslein, Kenneth

Sherman, Robert Reid, and Frank Steimle for their

critical review of the manuscript and their many helpful suggestions to improve it.

It is our pleasure to acknowledge the wholehearted

cooperation of the officers and crews of the research

vessels Albatross III (Capt. Emerson Hiller), Albatross IV

(Capt. Walter E. Beatteay), Delaware (Capt. John ].

Walsh), Asterias (Capt. Arthur D. Colburn Jr.), and

Gosnold (Capt. Harry Seibert).

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