% i? NOAA Technical Report NMFS 140 December 199s 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 U.S. Department of Commerce C U.S. DEPARTMENT OF COMMERCE WILLIAM M. DALEY SECRETARY National Oceanic and Atmospheric Administration D.James Baker Under Secretary for Oceans and Atmosphere National Marine Fisheries Service Rolland A. Schmitten Assistant Administrator for Fisheries ^1 ^ NOAA Technical Reports NMFS Technical Reports of the Fishery Bulletin Scientific Editor Dr. John B. Pearce Northeast Fisheries Science Center National Marine Fisheries Service, NOAA 166 Water Street Woods Hole, Massachusetts 02543-1097 Editorial Committee Dr. Andrew E. Dizon National Marine Fisheries Service Dr. Linda L. Jones National Marine Fisheries Service Dr. Richard D. Methot National Marine Fisheries Service Dr. Theodore W. Pietsch University of Washington Dr. Joseph E. Powers National Marine Fisheries Service Dr. Tim D. Smith National Marine Fisheries Service Managing Editor Shelley E. Arenas Scientific Publications Office National Marine Fisheries Service, NOAA 7600 Sand Point Way N.E. Seattle, Washington 981 15-0070 — -^ — LD ■ ru r-=1 - tr <-* m □ L/WHO m — r^ The NOAA Technical Report NMFS (ISSN 0892-8908) series is published by the Scientific Publications Oflice, Na- tional Marine Fisheries Service, NOAA, 7600 Sand Point Way N.E., Seattle, WA 98115-0070. The Secretary of Commerce has de- termined that the publication of this se- ries is necessary in the transaction of die public business required by law of Uiis Department. Use of funds for printing of this series has been approved by the Di- rector of the Office of Management and Budget. The NOAA Technical Report NMFS series of the Fishery Bulletin carries peer-re- viewed, lengthy original research reports, taxonomic keys, species synopses, flora and fauna studies, and data intensive reports on investigations in fishery science, engineering, and economics. The series was established in 1983 to replace two subcategories of the Technical Report series: "Special Scientific Report — Fisher- ies" and "Circular." Copies of the NOAA Technical Report NMFS are available free in limited numbers to government agencies, both federal and state. They are also available in exchange for other scientific and technical publications in the marine 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. Purchasing additional copies Additional copies of this report are available for purchase in paper copy or microfiche from the National Technical Information Service, 5285 Port Royal Road, Springfield, VA 22161; 1-800-553-NTIS; http://vvww.ntis.gov. Copyright law Although the contents of the Technical Reports have not been copyrighted and may be reprinted entirely, reference to source is appreciated. Proprietary products The National Marine Fisheries Service (NMFS) does not approve, recommend, or endorse any proprietary product or proprietary material mentioned in this publication. 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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 Fauna] 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 Sediment Organic Carbon 53 Taxonomic Groups 55 Porifera 56 Coelenterata 71 Hydrozoa 71 Anthozoa 75 Alcyonaria 75 Zoantharia 79 Platyhelminth.es 81 Turbellaria 81 Nemertea 85 Aschelminthes 91 Nematoda 91 Annelida 94 Pogonophora 100 Sipunculida 103 Echiura 108 Priapulida 108 Mollusca HO Polyplacophora I l() ( ■; stropoda II-1 ivalvia 124 Scaphopoda 130 Cephalopoda 135 Arthropoda 139 Pycnogonida 139 Aj achnida 143 Crustacea 143 ( )stracoda 1 44 ( lirripedia 1 43 Copepoda 155 ( lumacea 1 35 Tanaidacea 159 [sopoda 162 Amphipoda L65 Mysidacea 170 Decapoda 174 Bryozoa 178 Brachiopoda 181 Echinodermata 185 Crinoidea 186 Holothuroidea 187 Echinoidea 192 < )phiuroidea '. 199 Asten tidea 205 1 Iemi( hordata 208 Chordata 210 Am idiacea 210 Dominant Components ol the Macrobenthos 214 Frequency of Occurrence 216 Percentage Composition 216 Geographic Distribution 216 Selected ( lenera and Species 217 Phylum Annelida 217 Phylum Mollusca 217 Phylum Anthropoda 217 Phylum Echinodermata 224 Bathymetri( Distribution 224 Relation to Bottom Sediments 225 Relation to Water remperature 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-1950s to the mid-1 960's a series of quantitative surveys of the macrobenthic invertebrate fauna were conducted in the offshore New England region (Maine to Long Island, NewYork). The surveys were designed to 1) 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 tvpe, tempera- ture 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 of the 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 densitv and biomass increase from northeast to southwest; both densitv and biomass decrease with increasing depth; the composition of the bottom sediments significantlv 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 sea- sonal 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 detect- able 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 topo- graphic feature of the continental margin because of its influence on the marine life of the 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 offish that have been taken each year since pre-Colonial days. The marine life in- habiting New England offshore waters has been the subject of studies conducted from time to time through- out the past century. This has resulted in the acquisi- tion 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 Holt-, MA 02543. NOAA Technical Report NMFS 140 daily regarding quantitative aspects. Because of the key role played by macrobenthic invertebrates in the eco- logical dynamics of the marine environment, their use- fulness to man as a food resource, their potential as concentrators of toxic substances that could be trans- mitted through the food chain, and their usefulness as indicators of environmental change, the National Ma- rine Fisheries Service (formerly the Bureau of Com- mercial Fisheries) of the U.S. Department of Com- merce, 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 major taxonomic groups of macrobenthic invertebrates inhabiting the continental shelf and slope between Cape Cod, Massachusetts, and Cape Hatteras, North Carolina. Their distribution in relation to geo- graphic location, water depth, bottom sediments, range in bottom water temperature, and sediment organic carbon content is considered. The present report describes the quantitative distri- bution of the principal groups of macrobenthic inverte- brates 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 com- position, trophic zonation, faunal dominance and di- versity, and similar topics will be the subjects of future reports. The large database generated by the Continental Margin Program contains a wealth of valuable geologi- cal, faunal, and environmental information of histori- cal as well as current significance. In addition to provid- ing input for a variety of descriptive studies, as de- scribed above, the potential exists for information con- tained in the database for ecosystem modeling tasks; paleoecological and global climate change studies; and benthic production estimates (Cohen etal. 1978, 1982; Cohen and Wright 1979; Warwick 1980; Rowe el al. 1986, 1988; Bourne 1987; Cohen and Grosslein 1987; Steimle 1987. 1990a, 1990b; Rowe et al., 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 distri- bution of 44 faunal groups in relation to the five environ- mental parameters; 1) geography, 2) bathymetry, 3) bottom sediments, 4) bottom water temperatures, and 5) sediment organic carbon. Quantitative data for geo- graphic distribution are presented at two different lev- els: a detailed evaluation based on calculations for each of several hundred unit areas (20 min in latitude bv 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 of macrobenthos presented in phylogenetic order. The final section is a summary of the environmental rela- tionship of the dominant taxonomic components. Previous Studies One of the earliest studies in marine benthic ecology dealt with populations inhabiting the Woods Hole-Vine- yard Sound area off southeastern Massachusetts (Verrill et al., 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 litera- ture. Included in the report are descriptions of new species, an annotated catalog of animals found in Vine- yard Sound and vicinity, and, significantly, a large part of the report is devoted to descriptions of the benthic com- munities 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 of information to supplement their studv (Ckaild 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 collec- tions 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 coopera- tion 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 col- lected in coastal areas between New Haven, Connecti- cut, and Eastport, Maine; only a moderate number of collections were from offshore areas. Inshore opera- tions were conducted from the vessels Moccasin, Mosswood, Bachf, Speedwell, Blur Light, and to some ex- Composition and Distribution of Macrobenthic Invertebrate Fauna tent the Blake and Fish Haii'k; however, the latter two also operated in offshore areas, as did the Albatross and the chartered fishing schooner./osw Reeves (Packard, 1874, 1876: Agassiz, 1881; Smith and Rathbun, 1882; Tanner, 1882; Smith and Rathbun, 1889; Townsend, 1901 ). This early sampling was primarily exploratory in na- ture. The participating zoologists faced a vast unstud- ied fauna and a multitude of species new to science. Scientists most active in this work were chiefly system- atists; 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 pro- ductive systematist of this, or perhaps any era. He de- scribed 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 prelimi- nary 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 com- prehensive 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 sedi- ments) 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 con- cerned with offshore invertebrates — proceeded at an exceedingly slow pace. Belding (1914), Allee (1922a, 1922b, 1923a, 1923b, 1923c), Pytherch (1929), Stauffer (1937), Avers (1938), and others contributed valuable information on inshore populations. Rather few eco- logically oriented works such as Procter (1933a, 1933b) and Bigelow and Schroeder (1939) pertaining to off- shore zoobenthos appeared during this period. In addi- tion to the foregoing, however, many studies of a taxo- nomic nature containing valuable ecological informa- tion were issued during this time span (Rathbun, 1905, 1925; Koehler, 1914; Nutting, 1915; Pilsbrv. 1916; Heath, 1918; Bartsch, 1922; Deichmann, 1930, 1936; and oth- ers). Ecological interests of marine scientists conduct- ing 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 al., 1956; Parker and Athern, 1959; Stickney, 1959; Rhoads, 1963; and oth- ers). Ecological studies pertaining to the offshore popula- tions commenced somewhat later, for example the re- ports by: Parker (1948); Northrup (1951 ); Phleger (1952); Clarke (1954); Schroeder (1955, 1958); Taylor etal. (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 of Vineyard 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. al., 1962), Greenwich Bay, Rhode Island (Stickney and Stringer, 1957), Sheepscot Estuary (Hanks. 1964), Narragansett Bay (Phelps, 1965), Rand's Harbor, Massachusetts (Burbanck et al., 1956), and other locales. In recent years, due to increased interest in potential impacts of man's activities in outer continental shelf (OCS) devel- opment 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 (1961b); Sanders et. al. (1962); Wiglev and Mclntyre (1964); Emery etal. (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 Leathern (1980. 1981a, 1981b); Valentine etal. ( 1980); Magnuson et. al. (1981 ); Wigley and Theroux (1981); Maurer and Wigley ( 1982, 1984); Stehnle (1982); Caracciolo and Steimle (1983); Lear and O'Mallevf 1983) ; Steimle ( 1985); Rowe etal. (1986); Maciolek and Grassle (1987); Michael (1987); Theroux and Grosslein (1987); Langton et al. (1988); Langton and Uzmann ( 1988); Sherman etal. (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. 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 Fisher- ies 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 Livingston Jr. and published in 1965. 4 Marine and Estuarine Environments, Organisms and Geology of the Cape Cod Region, an Indexed Bibli- ography, 1665-1965. Compiled by Anne E. Yentsch, M. R. Carriker, R. H. Parker, and V .A. Zullo, pub- lished in 1966. 5 Fishery Publication Index, 1955-64. U.S. Fish & Wild- life Service, Bur. Coram. Fish. Circ. 296, published in 1969. 6 The Effects of Waste 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 1 — List of Citations; Part 2 — Indices. Compiled by the Na- tional 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. Com- merce, NOAA Tech. Rep. NMFS Circ. 400, pub- lished in 1977. 10 A Summary and Analysis of Environmental Informa- tion on the Continental Shelf from the Bay of Fundy to Cape Hatteras (1977). Vol. II, Master Bibliogra- phy, 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 Cen- ter. Northeast Regional Coastal Information Cen- ter, Marine Advisory Service, National Sea Grant Depository, Univ. of Rhode Island Mar. Tech. Rep. 70, published in 1979. 12 An Ecological Characterization of Coastal Maine (North and Fast of Cape Elizabeth). Vol. 5, Data Source Appendix. Compiled by S. E. Fefer and P. A. Schetting for Biol. Serv. Program, Interagency En- ergy/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.). Na- tional Undersea Research Program, Research Re- port 88-3, published in 1988. Another result of increased OCS activity is the large volume of information relating to benthic fauna ap- pearing in the so-called "gray" literature. Included in this category are completion reports of field study con- tracts, environmental impact statements, public and pri- vate agency investigation reports, annual reports, and other similar special documents. Many appear in irregular se- ries, 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. al., 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 Mclntyre (1964); Emery etal. (1965); Trumbull (1965); Merrill etal. ( 1965); Wigley and Shave (1966); Wigley and Emery (1967); Schopf (1968b); Haynes and Wigley ( 1 969) ; Plough ( 1 969) ; Hazel ( 1 970 ) ; 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); Theroux1; Theroux and Wigley (1983); Rowe etal. (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); 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, Natl. Mar. Fish. Sen., Northeast Fish. Cntr., Woods Hole Lab. Ref. Doc. No. N:v_>7. 280 p. Unpubl. manus( ript. Composition and Distribution of Macrobenthic Invertebrate Fauna and Rowe et al. ( 1991 ); Burns and Wigley2; Wigley et al. , Theroux and Wigley4; Manrer and Wigley'; Theroux. et al.h; Theroux and Schmidt-Gengenback. 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 En- gineers; 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 main 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 Re- sources Mapping, Assessment, and Prediction program (Sherman, 1980); the Marine Ecosystem Analysis Pro- gram (MESA) (see Pearce et al., 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 im- pacts of present and future activities related to oil and gas exploration and production, marine mining, ocean dump- ing, other waste disposal, and natural environmental -' Burns, B. R., and R. L. Wigley. 1970. Collection and biological data pertaining to mvsids in the collection at the BCF Biological Laboratory, Woods Hole. U.S. Bur. Conun. Fish. Biol. Lab. Woods Hole. Mass., Lab. Ref. 70-3, 36 p. (mimeo). L'npubl. manuscript. 3 Wigley, R L., R. B. Theroux, and H. E. Murray. 1976. Marine macrobenthic invertebrate fauna of the Middle Atlantic Bight re- gion. Part 1. Collection data and environmental measurements. Northeast Fisheres Center, Woods Hole Lab. Ref. Doc. 7618, 34 p. (mimeo). L'npubl. Manuscript. * Theroux. R. B., and R. L. Wigley. 1979. Collection data for U.S. east coast bivalve mollusks in the Northeast Fisheries Center Speci- men Reference Collection, Woods Hole. Massachusetts. Northeasl Fisheries Center, Woods Hole Laboratory, National Marine Fisher- ies Serv., NOAA. Northeast Fisheries Center.,Woods Hole Lab. Ref. Doc. 79-29, 471 p. (mimeo). L'npubl. 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 Labi irat< iry, NOAA, Woods Hole Lab. Ref. Doc. 81-15, 97 p. (mimeo). L'npubl. manuscript. 6 Theroux, R. B., R. L. Wigley, and H. E. Murray. 1982. Marine mai robenthic invertebrate fauna of the New England Region: Collec- tion data and environmental measurements. Nat. Mar. Fish. Serv., NOAA, Northeast Fish. Center, Woods Hole Lab. Ref.Dor 82-40, MARMAP Contrib. MD/NEFC 82-67. 74 p. (mimeo). Lnpubl. Manuscript. ' Theroux, R. B., andj. Schmidt-Gengenbach. 1984. Collection data and environmental measurements for U.S. east coast Cumace.i (Ar- thropoda, Crustacea) in the Northeast Fisheries Center Specimen Ref- erence Collection Woods Hole, Massachusetts. Nat. Mar. Fish. Serv., Northeast Fisheries Center. Woods Hole Lab. Ref. Doc. 84-27, MARMAP Contr. MED/NEFC 83-46, 1 14 p. (mimeo). Lnpubl. Manuscript. 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 al., 1976a, 1976b, 1976c, 1976d, 1977a, 1977b, 1977c, 1978, 1981; Caracciolo et al., 1978; Pearson and Rosenberg, 1978; Reid et al., 1979; Steimle and Radosh, 19/9; Warwick, 1980; Schaffnerand Boesch, 1982; Steimle. 1982; Boehm, 1983; Caracciolo and Steimle, 1983; Lear and O'Mallev. 1983; Steimle, 1985; Steimle and Terranova, 1985; Duinker and Beanlands, 1986; Howart,1987; Neff, 1987; Reid et al., 1987; Steimle. 1990a, 1990b; Steimle et al., 1990). Materials and Methods Macro fauna Samples 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 l).The geographic locality of sampling sites is illustrated i,i 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 num- Table 1 Researc li vessels, i rnise numbers, date < f collections. and numbei of samples obtained. t l mse Number of Vessel number Date samples Albatross /// 80 Angus! 1956 35 Albatross /// 101 August 1957 165 Delaware 59-9 August 1959 75 Delawan 61-10 June 1961 75 Delaware 62-7 June 1962 123 Cos n old 10 April 1963 7 Com old 1 1 April 1963 3 GosnoM 12 May 1963 38 Gosnold 13 May 1963 29 Gosnold 20 Jul) 1963 1 Gosnold 22 August 1963 93 Gosnold 24 August-September 1963 32 Gosnold 28 October 1963 9 Gosnold 29 October 1963 84 Gosnold 49 August 1964 72 Gosnold 51 September 1964 7 Asterias 1 April 1964 8 Asterias 2 July-August 1964 62 Albatross TV 64-12 October 1964 24 Albatross I\ 64-13 October— Novembe ■ 1964 10 Albatross IV 65-11 August 1965 1 23 Total 1.076 NOAA Technical Report NMFS 140 ' \ y NEW \ NEW JERSEY \ YOfiK yS NEW Figure 1 Chart of the studv 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 environmental measurements (including water depth in meters, bot- tom type, geographic subarea, temperature range [°C], and percent organic carbon) for each sampling site are contained in Theroux et al.1'). Sampling stations were located in all sections of the studv 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 re- gion beyond the continental shelf. Table 2 lists the number of samples and occurrence frequency for each 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 bet attse 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 3,973 m. Sampling Gear The samples consisted of bottom sediments with the constituent fauna collected with a Smith-Mclntyre spring- loaded grab sampler (Smith and Mclntyre, 1954) illus- trated in Figure 3, or a Campbell grab sampler (Menzies et al., 1963) illustrated in Figure 4. The bottom area sampled by the Smith-Mclntyre sampler was 0. 1 m which had a capacity of approximately 1 5 liters (L) . Area sampled by the Campbell sampler was 0.5b m'-', which had a volume capacity of about 200 I .. The Campbell grab was equipped with a 35-mm camera and electronic flash, housed within the buckets, to obtain photographs of the bottom imme- diately before impact (Emery and Merrill, 1964; Emery et al., 1965; Wigley and Emery, 19(17; 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 MUM6ER OF SAMPLES PER UNIT AREA ( 20' x 20' ) • 1 - 2 • 3-5 • 6-10 DEPTH IN METERS Figure 2 Chart showing sampling intensity within each standard unit area (20 min. latitude bv 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 directiv 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 l.O I., depending upon the total volume of material obtained. The quantities of both samples and subsamples were measured and recorded on sample log sheets. Gener- ally, the remaining material was washed through a 1-mm- aperture 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 of formaldehyde 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 practi- cable taxonomic level, counting, and weighing. Weights are damp formalin weight, the "rough weight" of Petersen (1918), herein referred to as wet weight inasmuch as the superficial fluid on the specimens was removed by blot- ting 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 liring animal, i.e. shells of liring 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 or sipunculid worms, and other similar nonintegral structures or nonliv- ing 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. NOAA Technical Report NMFS 140 Specimens of each taxon were bottled separately in 70% ethanol and labeled. Subsequently, specimens were assembled bv taxonomic groups and sent to cooperat- ing 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 meth- ods, physical and chemical characteristics of the envi- Table 2 Numbers of samples and occurrence frequency in each of the various p arameter grot pings used in this report. Number of Frequency (%) ..1 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 10(1-199 246 22.9 L>< II 1-499 106 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-cla) 234 21.8 Temperature ran ge(°C) 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 organi< carbon ('; ) 0.00 5 n -> 0.01-0 19 418 38.8 0.50-0.99 167 15.5 1.00-1.49 84 7.8 1.50-1.99 43 4.0 2 0() 2.99 13 1.2 3.00 1 99 1 114 5.00+ 1 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 data- processing cards. Data were summarized by computer in a form similar to that presented in the tables appear- ing in the body of this report. The principal units used for expressing the quantity of benthic in\ertebrates (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. Fattnal 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 sea- sons and between years were sufficiently small to permit the consolidation of all samples for purposes of this report. Some seasonal and yearly differences in taxo- nomic composition and quantity of animals were de- tected 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 an- other. One of the chief reasons for the temporal stabil- ity 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 re- ports also allude to the temporal persistence of certain dominant components of the macrobenthos of the re- gion (Steimle, 1990a, 1990b; Maurer8; Michael etal.9). Geographic Areas For purposes of detecting and reporting regional dif- ferences in fauna! composition the region has been s Maurer. D. 1983. Review of benthic invertebrates of Georges Bank in relation to gas and oil exploration with emphasis on management implications. Natl. Mar. Fish, Sen.. Northeast Fisher- ies Center, Woods Hole. Massachusetts. Woods Hole Lab. Ret. Doc. 83-16, 329 p. (mimeo). L'npubl. manuscript. " Michael, A. D.. C. D. Long. IX Maurer. and R. A. McGrath. 1983. ( ieorges Bank benthic infauna historical study. Final report to U.S. Dep. Interior, Minerals Management Service, Washington, DC, Rep. 83-1 bv Taxon Inc. Salem. MA 01970. 171 p. Composition and Distribution of Macrobenthic Invertebrate Fauna ■ Figure 3 Side view of the Smith-Mclntyre 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 cm. divided geographically into six subareas (Fig. 5). These are: 1) Nova Scotia, containing 44,816 km2 (13, 049 mi2) — encompassing southwestern Bay of Fundy, east- ern 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 km'2 (23,313 mi2); 3) Georges Bank— consist- ing only of Georges Bank proper with an area of 39,21 1 km2 (11,417 mi2); 4) Southern New England Shelf oc- cupying 73,318 km2 (21,348 mi2) — including the conti- nental shelf from Great South Channel southwestward to central New Jersey; 5) Georges Slope — the continen- tal 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 of Hudson Canyon, occupying 62,570 km2 (18,218 mi2). Each subarea has specific biotopic and biogeographic fauna! characteristics. These are discussed in the sec- tion entitled "Description of the Region" and in the "Geographic Distribution" section for each of the ma- jor taxonomic groups. Bottom Sediments Bottom sediments from the samples have been ana- lyzed for particle size, composition, and color. In addi- tion, a selected series of these samples was further ana- lyzed for carbonate content (Hulsemann, 1966), quan- tity of organic matter (Hulsemann, 1967) and mineralogy (Ross, 1970b). Detailed particle size analy- ses of approximately 75% of the samples were made by John Schlee, U.S. Geological Survey (Schlee, 1973). 1 0 NOAA Technical Report NMFS 140 ' r Figure 4 Bottom view of Campbell grab sampler. Camera housing is installed in right-hand bucket and strobe light is in the left-hand bucket. Shutter trip weight is in foreground. Width of the 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 techniques by K.O. Emery of the Woods Hole Oceanographic Insti- tution or by National Marine Fisheries Service person- nel. For additional information concerning sediment analyses, methodology, and detailed results, see refer- ences listed by Emery (1966b) and the section of this report titled "Description of the Region." Bathymetry Water depths, in meters, were obtained by means of echo sounders and precision depth recorders and cor- rected for hydrophone transducer depth and tempera- ture effects on the velocity of sound in water. Temperature Water temperature and salinity data were based prima- rily 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 Alba- tross III, Albatross TV, and Delaware, conducted by the Bureau ol Commercial Fisheries Biological Laboratory, Woods Hole, during the years 1956 through 1965. Ad- ditional 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 Ges" "slope ■V- -&- -V 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 CaCO, 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; Valen- tine, 1981). Figure 6 is based on, and has been derived from, a much larger more detailed chart by Uchupi (1965a), U.S.G.S. Map 1-451, scale 1:1,000,000. Topographically, the New England offshore area con- sists of several large, grossly different geological fea- tures. The largest and most complex feature is the Gulf of Maine, an immense, nearly oval-shaped glacially eroded depression on the continental shelf. The topo- graphy in this depression is very irregular, resulting in numerous basins separated l>\ i idges. swales, and banks. These topographic irregularities are due in part to deposition, gouging, erosion, and related actions dur- ing the Pleistocene period of glaciation. Greatest depth in the gulf is 377 m, in Georges Basin; shallowest off- shore 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 fea- ture. It is an enormous (120 km by 240 km) submarine cuestalike bank situated at the mouth of the Gulf of 1 2 NOAA Technical Report NMFS 140 Figure 6 Chart of the studv area showing bathymetric and geographical features. Depth contours are in meters (adapted from Uchupi 1965). Maine. The bank slopes gentlv to the southeast and south and its surface is relatively smooth except for a series of sand ridges in the shallow northwest and north- central sections. The sand ridges are formed by excep- tionally 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 Emerv and Uchupi (1965); Uchupi etal. (1977); Valentine (1981); Butman (1982, 1987); Butman et al. (1982. 1987); Backus (1987); Bourne (1987); Butman and Beardsley (19X7); Cohen and Grosslein (1987); Cooper et al. (1987); Emery (1987); Flagg (1987); Howart (1987): Klitgord 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 topo- graphic all\ uneven area southeast of Nantucket Island, Massachusetts. Prim ipal irregularities are large swales and ridges extending in north-south and northeast- southwest directions. The southern New England continental shelf is a gentlv seaward-sloping region with rather smooth to- pography. 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 compara- tively 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 gener- ally similar to the slope in having only gradual changes in surficial topography. However, the overall gradient is sub- stantially less than that for the continental slope. Consult Emery (1965a), Emery and Ross (1968), Gibson et al. (1968) , Schlee etal. (1979), Sheridan (1974), and Uchupi et al. ( 1977) for details of topography of this region. Composition and Distribution of Macrobenthic Invertebrate Fauna NEW \ NEW JERSEY \ YORK/ NEW JTORK GRAVEL SHELL TILL SAND E;;jj;;[ silty sand 2] SILT-CLAY 40° Figure 7 Geographical distribution of bottom-sediment tvpes in the stud) area. Bottom Sediments The composition of sediments blanketing the sea floor throughout the study area is well known. Detailed stud- ies 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 com- ponents. A representative selection of publications deal- ing 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); Wiglev (1961a); Uchupi (1963, 1965b, 1966a. 1966b. 1966c, 1968, 1969); Emery (1965a, 1965b, 1966a, 1966b, 1968); Emerv et al. ( 1965); Rvac hev ( 1 965) ; Garrison and McMaster ( 1 966) ; Hiilsemann ( 1 966, 1967); McMaster and Garrison (1966); 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); Mtlliman (1973); Wigley and Stinton ( 1973); Sheridan ( 1974); Austin et al. ( 1980); Twichell et al. (1981); Butman (1982, 1987); Klitgord et. al. (1982); and Valentine etal. (1980). Relict glacial sediments are the major constituents cov- ering most of the study area, particularly on the continen- tal shelf. Quart/ 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 pro- tected 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 shal- low 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 clav. In Long Is- land 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 sedi- ments 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 car- bon 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 of sediments 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% or- ganic 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 or- ganic carbon content (from 2.00 to 7.04%) was almost exclusively restricted to the major embayments and estuaries within the study area; only offshore excep- tions 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 off- shore New England region. Some of the first hydro- graphic 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 of Canada (Fisheries Research Board of Canada), coastal states organizations, Bigelow Laboratory, Woods Hole Oceanographic Institution, Harvard University, Massachusetts Institute of Technology, L'niversity of 74° ~7~ "v ~7~ ~^T "7^ MAINE TT-- HK v V •■'• NEW \ NEW JERSEY \ YORK/" NEW . \ \ NtW / \ HAMPSHIRE ,' \ V / CONNECTICUT V \ .' PORTLAND MASSACHUSETTS-?*^ "' ^-iip 120 _l KM Figure 8 Geographic distribution <>l organic carbon in the bottom 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 sub- ject 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 bv Dean F. Bumpus and his colleagues at the Woods Hole Oceanographic Institu- tion (Stetson, 1937: Bumpus, 1960, 1961;Colton, 1964; Bumpus and I.auzier, 1965; Bumpus et al., 1973; Butman et al., 1980, 1982; Dorkins, 1980; Ramp et al., 1980. Moodv et. al., 1984, Mountain and Hol/warth. 1989. among others). Discussions of earl) oceanographic re- search in this region and references to the literature are given by Colton (1964), Schopf (1968a). and Wright (1987). In brief, the main features pertaining to water circu- lation 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 Gulf of Maine; 2) Gulf of Maine waters form a large nontidal counter- clockwise 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 ampli- tude is exceptionally large in the Bav of Funch' 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 Nan- tucket Island); and 9) below the Gulf Stream in the vicinity of the ocean bottom, the Western Boundarv Current flows southwestwardly. Oceanic waters in the vicinity of the Gulf Stream maintain a relatively constant salinity of about 35%o. Most of the waters overlying the continental shelf have a salinity range from 32 to 34%o. Salinity of inshore waters, which are more stronglv influenced bv runoff, fluctuate seasonally and drop to 28%o 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 (0°C) in winter and warm in summer, and because of the shallowness and general turbulence of the water, the temperature differential between surface and bot- tom is relatively small. Also, there is considerable latitu- dinal 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 in- shore zones — are generallv intermediate in their tem- perature regime. Temperature diversitv 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 tempera- ture of bottom water is considerably less than that of surface waters. latitudinal effect on shelf water masses is pronounced; Nova 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 al.. 1962; Colton et al., 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 al."; Colton et al.1-'; 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 distri- bution in relation to the approximate annual mini- mum and maximum water temperatures, and the range in water temperature, to which the various taxa are 111 Colton, J. B.,Jr.. R. R. Marak. andS. R. Nickerson. 1965a. Envi- ronmental observations on continental shelf Nova Scotia to long Island, March 196"). Albatross IV cruise 65-3, U.S. Bur. Commer. Fish. Biol. Lab. Woods Hole, Mass., l.ab. Ret. 65-15, 3 p., '.I figs, (mimeo). L'npubl. manuscript. 11 Colton. J. B, Jr., R. R. Marak. and S. R. Nickerson. 1965b. Envi- ronmental observations on continental shelf Nova Scotia to Long Island, September 1965, Albatross IV cruise 65-12. U.S. Bur. Commer. Fish. Biol. Lab. Woods Hole, Mass., Lab. Rel. 65-19, 3 p., 9 figs, (mimeo). L'npubl. manuscript. 12 Colton, J. B.,Jr., R. R. Marak. and S. R. Nil kerson. 1966a. Envi- ronmental observations on continental shell Nova Scotia to Long Island. March 1966. Albatross IV cruise 66-2. U.S. Bur. Commer. Fish Biol. Lab Woods Hole, Mass.. Lab. Ref. 66-6. 3 p., 10 figs, (mimeo). Unpubl. manuscript. 13 Colton. J. 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 l\ iiuise 66-7. U.S. Bur. Commer. Fish. Biol. Lab.. Woods Hole. Mass., Lab. Ref. 66-7. 3 p., 19 figs, (mimeo). Unpubl. manuscript. 16 NOAA Technical Report NMFS 140 subjected. Charts were constructed to illustrate the iso- therms of maximum 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 bio- logical data were collected, August 1956 through Au- gust 1965, and from the literature (see above citations). Temperature patterns depicted in these charts are in- tended to provide a general scheme of annual tempera- ture 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 con- sidered 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 of Maine is relatively cold, 4 to 8°C and changes very little throughout the year. Bot- tom 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 fauna! complexities of the sublittoral portion of the study area cause considerable difficulty in the definition of defini- tive zoogeographic boundaries in the Northwest Atlan- tic. Until recently, the traditional view among biogeog- raphers was that the region embraced portions of two major zoogeographic provinces: 1 ) The Boreal Prov- ince, sometimes referred to as Acadian or Nova Scotian, which extends from Newfoundland to Cape Cod, and 2) The Trans-Atlantic or (Warm Temperate) Province Figure 9 Distribution of maximum reported 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 hydrographi- cally), no definitive consensus of opinion among bio- geographers of the period prevailed as to the precise placement of the boundaries in the Northwest Atlantic. Indeed, the plethora of varying definitions and terms led to a rather confusing semantic problem that exists to this dav. 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 Virgin- ian subprovince to the south, with Cape Hatteras form- ing the boundary between the Virginian and Carolin- ian subprovinces, which together formed the Trans- Atlantic 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 Prov- inces; and 3) A cold temperate Boreal Province extend- ing 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, signifi- cantly altered the three biogeographic concepts of ear- lier workers. These recent works, however, have pro- vided some new insights concerning the placement of Figure 10 Distribution of minimum reported bottom water temperatures (in degrees Celsius) in the study area. 1 8 NOAA Technical Report NMFS 140 Distribution oi 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 regulat- ing 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 zoogeographi- cal barrier. Cape Cod, lying roughly in the center of the study area, is of course the main physical feature his- torically considered to mark the separation between the Boreal and Trans-Atlantic Provinces. The recent work of Schopf (1968b), Hazel ( 1970), Watling (1979b), Fran/ et al. (1981), and other reports (Wigley and Burns, 1971; Williams and Wigley, 1977; Theroux and Wigley, 1983; and Theroux and Grosslein, 1987) based cm the same data as, and including, the present report corroborate the fact that Cape Cod is indeed a zoogeo- graphic 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 north- ern 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 west- ern 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 Atlan- tic Transitional Province (Cutler, 1977), and at depths between 2,000 and 4,000 m is the Atlantic Bathyal- Abyssal Province. Because of the interdigitating distri- butional patterns resulting from the southward sub- mergence of Boreal species and the ascendency of Tran- sitional and Bathyal-Abyssal species in their north- ward extension, the delineation of these provinces is imprecise and only partially aligned with topographic features. A great deal more work of a zoogeographic nature on the many remaining unstudied taxa of benthic inverte- brates inhabiting the area needs to be performed be- fore precise zoogeographic boundaries may be drawn, if at all possible. Composition and Distribution of Macrobenthic Invertebrate Fauna 19 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 va- rietv of dominants and codominants, is char- acteristic of the fauna, and is generally typi- cal of Boreal-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 South- ern New England and Georges Bank areas. The ecological importance of these groups, judged primarily from their numeri- cal abundance and biomass, ranges from minor (components that account for less than 0.1% in number of individuals and bio- mass) to dominant components that make up 20% or more in number of individuals or biomass. The 44 major taxonomic groups, with the percentage of total number of indi- viduals and percentage of total biomass for each, are listed in Table 3. Also, they are classi- fied into four dominance categories, I to IV. Over 80% of both the biomass and num- ber 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 contribu- tor (44.1%) to the biomass and is also a major component (10.8%) in terms of num- bers of individuals. Amphipoda, on the other hand, is numerically dominant (43.4%) but contributes only 2.3% of the biomass. Con- versely, 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 Newr England region macrobenthos is dominated by members of four phyla: Annelida, Mollusca, Arthro- poda, and Echinodermata. These groups will be dis- Table 3 Rank order of major taxonomic groups according to percentage composition of the total macrobent ln< fauna in terms of biomass and number of specimens F ercentage Percentage <>t Dominance ol total total number ( ategory Taxa biomass Taxa specimens I Bivalvia 44.1 Amphipoda 43.4 Echinoidea 20.0 Annelida 28. 1 Annelida 9.5 Bivalvia 10.8 Holothuroidea 7.0 Total 80.6 Total 82.3 II Zoantharia 3.5 Ophiuroidea 2.9 Amphipoda 2.3 Echinoidea 1.9 Ascidiacea 2.2 Cumacea 1.7 Cirripedia 1.9 Zoantharia 1.5 Ophiuroidea 1.8 Cirripedia 1.5 Gastropoda 1.2 Gastropoda 1.2 Asteroidea 1.2 Ascidiacea 1.1 Porifera 1.2 Bryozoa 1.0 Total 15.3 Total 12.8 III Decapoda 0.8 Isopoda 0.8 Bryozoa 0.7 Nemertea 0.5 Brachiopoda 0.5 Decapoda 0.5 Nemertea 0.4 Hydrozoa 0.4 Sipunculida 0.4 Sipunculida 0.4 Hvdiozoa 0.3 Brachiopoda 0.3 Scaphopoda 0.2 Scaphopoda 0.3 Echiura 0.2 Holothuroidea 0.3 Isopoda 0.2 Nematoda 0.2 Alcyonaria 0.1 Mysidacea 0.2 Polyplacophora 0.1 Porifera 0.1 Cumacea 0.1 Alcyonaria 0.1 Polyplacophora 0.1 \sll 1 1 lllll'.l 0.1 Total 4.0 Total 4.3 IV Turbellaria <0.1 Turbellaria <0.1 Priapulida Priapulida Nematoda Cephalopoda Cephalopoda Echinoidea Arachnida Arachnida Pycnogonida Pycnogonida Ostracoda Ostracoda Copepoda ( opepoda Mysidacea Tanaidacea Tanaidacea Crinoidea Crinoidea Pogonophora Pogonophora Hemichordata Hemichordata Echiura Total <1.0 Total <1.0 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), num- ber of samples, and frequency of occurrence for each taxonomic group considered in this report. 20 NOAA Technical Report NMFS 140 Table 4 List of macrobenthic invertebrate species contained in quantitative samples obtained within the study area. PORIFERA Funic id.it' Demospongiae Eunice pennata (Miiller. 1776) Hadromerida Eunice sp. Suberitidae Marphysa sp. Polymastia sp. Lumbrineridae COELENTERATA Lumbrinerides acuta (Verrill, 1875) Hydrozoa Lumbrineris fragilis (Muller. 1776) Hydractinia echinata Fleming, 1828 Lwmbrineris sp. Hydractinia sp Ninoe sp. Anthozoa Onuphidae Alcyoii.ii i.i Diopatra cuprea (Bosc. 181)1') Alcyonacea Diopatra sp. Alcyonium sp. Hyalinoecia tubicola (Muller, 1776) Gorgonai ea Hyalinoecia sp. At anella sp. Nothria conchylega Sars, 1835 Paragorgia arborea (Linnaeus, 1767) Onuphis eremita Audoin and Milne-Edwards, 1833 Primnoa reseda (Pallas. 17(>n> Onuphis iijiiiliiin (Verrill, 1873) Pennatulacea Onuphis quadricuspis Sars, 1872 Pennatula aculeata Danielssen and Koren, 1858 Onuphis */* Pennatula sp Paradiopatra sp. Stylaiula elegans (Danielssen, 1860) Flabelligerida Zoantharia Flabelligeridae Zoanthidea Brada sp. Epizoanthus incrustatus (Verrill, 1864) Flabelligera sp. Epizoanthus sp Pherusa sp. \i liniana Opheliida Tealina felina (Linnaeus. 1767) Opheliidae Edwardsia sulcata (T. Pennant. 1777) Ophelia sp. Edwardsia sp. Ophelina aulogastei (H. Rathke, 1843) Actinostola callosa Verrill, 1882 Ophelina sp. Antholoba perdix (Verrill, 1882) Travisia carnea Verrill, 1873 Madreporaria Travisia sp. Astrangia sp. Si alibi'cgmid.tc Flabellum goodei Verrill, 1878 Scalibregma inflatum Rathke, 184.3 Flabellum sp Scalibregma sp. Cen. inihci i.i Orbiniida Cerianthus borealis Verrill, 1878 Orbiniidae Cerianthus sp. Orbinia ornata (Verrill, 1873) i 'eriantheopsis americanus Verrill, 1866 Oilman nuani Pettibone, 1957 Annelida Orbinia sp. Polychaeta Scoloplos robustus (Verrill, 1873) Amphinomida Scoloplos sp. Amphinomidae Aricidea jeffreysii (M< Intosh, 1879) Paramphinome jeffreysii ( Mcintosh. 1 868 1 Aricidea sp. ( lapitellida Paraonidae ( apitellidae Paraonis sp Capitella sp Oweniida \l. lid. mid. ic Oweniidae [sychis biceps (Sars, 1861) Owenia fusiformis delle Chiaje, 1844 Maldane sp Owenia sp. ( '.osslll 1(1.1 Phyllodot id.i ( OSSIII 111. II' Aphroditidae ( ossura longicirrata Webster and Benedict, 1883 Aphrodita hastata Moore, 1905 Cossurci sp. Aphrodita sp. Eunii id.i Laetmonice sp \i abellidai ( .l\( c i ill. ii Arabella iricoloi (Montagu, 1804) Glycera americana Leidy, 1855 Arabella sp. Churn iit/iilnln Oersted. 1843 Drilonereis longa Webster, IN79 Civ mi dibranchiata Ehlers, 1868 Drilonereh sp. Glycera sp. Notot m us sp. continued on n '\7 /w£l' Composition and Distribution of Macrobenthic Invertebrate Fauna 21 Table 4 (continued) Goniadidae Spionidae Goniada maculata {Oersted. 1843) Diospio uncinata Hartman, 1951 Goniada sp. Laonice cirrada (Sars, 1851) Goniadella sp. Laonice sp. Ophioglycera gigantea Verrill, 1885 Polydora concharum Verrill, 1880 Ophioglycera sp. Polydora sp. Hesionidae Priospio sp Nereimyra punctata (O.F. Mullet, 1776) S/»(> «■/<»« Verrill, 1873 Nephtyidae sy»» sp. Aglaophamus circinata (Verrill, 1874) Spiophanes bombyx (Clarapede. 1870) Aglaophamus sp. Stern aspida Nephtys bucera Ehlers, 1869 Sternaspidae Nephtys incisa Malragren, 1865 Sternaspis scutata (Renter. 18(17) Nephtys picta Ehlers, 1868 Sternaspis sp. Nephtys sp. Terebellida Nereidae Ampharetidae Ceratocephale loveni Malmgren, 1867 Ampharete acutifrons (t. tithe. 1860) Ceratocephale sp. Ampharete sp. Neren sp Melinna cristata (Sars, 185] ) Phyllodocidae Melinna sp. Eteone sp. Pectinariidae Eumida sanguinea (Oersted, 1843) Pectinaria gouldii (Verrill, 1873) Phyllodoce arenae Webster, 1S79 Pectinaria sp. Phyllodoce sp. Terebellidae Pilargiidae Amphitrite sp. Ancistrosyllis sp. Streblosoma spiralis (Verrill, 1874) Polynoidae Steblosoma sp Harmothoesp. POGONOPHORA Lepidonotus squamatus (Linnaeus, 1758) Siboglinidae Sigalionidae Siboglinum anguslum Southward and Brattegard, 1968 Leanira sp. Siboglinwn atlanticum Southward and Southward, 1958 Pholoe minuta (Fabricius, 1780) Siboglinum eAmanz'Jagerston, 1956 Sigalion arenicola Verrill, 1879 Siboglinum holmei Southward .1963 Sigalion sp. Siboglinum pholidotum Southward and Brattegard 1968 Sphaerodoridae Siboglinum sp. Sphaerodorum gracilis (Ratlike. 1843) Polybrachiidae Syllidae Crassibranchia sandersi Southward, 1968 Exogone verugem (Clarapede, ] H tiS ) Diplobrachia snnilis Southward and Brattegard, 1968 Exogone sp. Polybrachia sp. Tomopteridae SIPUNCULIDA Tomopteris sp. Aspidosiphon zinni Cutler, 1969 Sabellida Golfingia catharinae (Mullet. 1789) Sabellidae Golfingia elongata (Keferstein, 1869) Chone infundibuliformis krover. 1856 Golfingia eremita (Sars, 1851) Chone sp. Golfingia flagrifera (Selenka, 1885) Euchone sp. Golfingia margaritaiea (Sars, 1851 ) Potamilla neglecta (Sars, 1850) Golfingia minuta (Keferstein, 1865) Potamilla reniformis (Linnaeus, 1788) Golfingia murinae murinae Cutler, 1969 Potamilla sp. Oncknesoma steenstrupi Koren and Danielssen, 1875 Sabella sp. Phascolion strombi (Montague. 181)4) Serpulidae Phascolopsis gouldi (Pout tales, 1851) Filograna sp. Sipunculus norvegicus Koren and Danielssen. 1875 Spirorbidae ECHIURA Spirorbis sp. Bonellia thomensis (Gmelin, 1788) Spionida Echiurus echiurus (Pallas. 1771) Chaetopteridae Echiurus sp. Spiochaetopterus sp. Ihedella akaeta (Zenkevitrh, 1958) Cirratulidae Maxmuelleria lankesteri (Herdman, 1898) Chaetozone sp. Prometor grandis (Zenkevitch, 1957) Cirratulus sp. Protobonellia sp Tharyx sp. Sluiterina sibogae (Sluiter, 1902) Sluiterina sp. continuedoi next page 22 NOAA Technical Report NMFS 140 Table 4 (continued) MOLLUSCA Neogastropoda Polyplacophora Muricidae Gastropoda Boreotrophon clathratus (Linnaeus, 1758) Prosobranchia Eupleura caudata (Say, 1822) Archaeogastropoda Columbellidae Fissurelidae Amphissa haliaeeti (Jeffreys, 1867) Puncturella noachina (Linnaeus, 1771) Anachis lafresnayi (Fischer and Bernardi, 1856) Lepetidae Anachis sp. Lepeta caeca (Muller, 1776) MitreUa lunata (Say, 1826) Trochidae Mitrellapura (Verrill, 1882) Calliostoma occidentalis (Mighels and Adams, MitreUa rosacea (Gould, 1841 ) 1 842 ) Mllnlla sp. Margarites castalis (Gould, 1841) Buccinidae Margarites groenlandicus (Gmelin, 1791) Buccinum undatum Linnaeus. 1758 Margarita helicinus (Phipps, 1774) Bute, nam sp. Margarites sp. Colus caelatui (Verrill and Smith. 1880) Solarietla lamellosa Verrill and Smith. 1880 Coins obesus (Verrill, 1884) Solariella obscura (Couthouy, 1838) Coins parvus (Verrill and Smith, 1882) Solariella sp Colus pygmaens (Gould, 1841) Mesogastropoda Colus sp. Littorinidae Neptunea decemcostata (Say, 1826) Littorina obtusata (Linnaeus, 1758) Neptunea despecta (Linnaeus, 1758) Rissoidae Neptunea sp. Alvania brychia (Verrill, 1884) Melongenidae Alvania pelagica (Sampson, 1851) Busycon canaliculatus (Linnaeus, 1758) Alvania areolata Stimpson, 1851 Nassariidae Alvania sp. Ilyanassa obsolela (Say, 1822) Turritellidae Nassarius trivittatus (Sa\. 1822) Tachyrhynchus erosus (Couthouy, 1838) Nassarius vibex (Say, 1822) Turritellopsis acicula (Stimpson, 1851) Cancellariidae Cerithiidae Admete couthouyi (Jaw 1839) Cerithiella sp. Turridae Diastema alternatus (Say, 1822) Oenopota decussata (Couthouy, 1839) Epitoniidae Oenopota harpularia (Couthouy, 1838) Epitonium dallianumVerrifl and Smith, 1880 Oenopota mcisula (Verrill, 1882) Epitonium greenlandicum (Perry, 1811) Pleurotomella agassizi agassizi Verrill and Smith, Melanellidae 1880 Couthouyella striatula (Couthouy, 1839) Pleurotomella blah-ana (Dall, 1889) Aclididae Pleurotemella curta rurta (Verrill, 1884) Aclis WCTn'ffiBartsch, 1911 Pleurotomella packardi packardi (Verrill, 1872) Trichotropidae Propebela elegans (Moller, 1842) Tnrhotropn hureahs Broderip and Sowerby, 1829 Propebela exarata (Moller, 1842) Crepidulidae Propebela turricula (Montagu. 1803) Crepidula fomicata Linnaeus. 1767 Pyramidellidae Crepidula plana Say, 1822 Odostomia dealbata (Stimpson, 1851 ) Crucibulum striatum Say, 1824 Odostomia dux Dall and Bartsch. 1906 Aporrhaidae Odostomia sp. Aporrhais occidentalis Beck. 1836 Turbonilla bushiana Verrill, 1882 Velutinidae Turbonilla elegantula Verrill, 1882 Velutina velutina (Muller, 1776) Turbonilla nivea (Stimpson. 1851 ) Velutina undata (Broun, 1839) Turbonilla polita (Verrill, 1872) Velutina sp. Turbonilla sp. Natit nl. it- Opisthobranchia I Amain, linos (Say, 1822) Acteonidae Lunatia triseriata (Say, 1826) Acteon sp. Lunatia pallida (Broderip, and Sowerby, 1829) Ringiculidae Lunatia sp. Ringicula nitida Verrill, 1873 Natica llama Broderip and Sowerby, 1829 Actiocinidae Natica pusilla Say, 1822 Acteocina canaliculata (Sa\, 1822) Polinices duplicatus (Sa\, 1822) Relusa obtusa (Montagu, 1807) immaculatus (Totten, 1835) Polinici > sp. continued on next page Composition and Distribution of Macrobenthic Invertebrate Fauna 23 Table 4 (continued) Scaphandridae Limopsis sulcata Verrill .mil Bush. 1898 Cylichna alba (Brown. 1827) Limopsis sp. Cylichna gouldi (Couthouy, 1839) Mvtiloul.i Cylichna vortex (Dall, 1881) Mvtilidae Cylichna sp. CreneUa decussata (Montagu. 1808) Si aphiiniler punt toslnatus Mighels, 1 84 1 Crenella gtandula (Totten. 18341 Philinidae Crenella sp. Philine lima (Brown, 1827) Dacrydium vitreum (Holboll in Moller. 18121 Philine quadrata (S. Wood. 1839) Modiolus modiolus (Linnaeus, 1758) Philine sp. Musculus corrugatus (Stimpson, 1851 ) Akeridae Musculus discors (Linnaeus, 1767) Haminoea sp. Musculus nigei (Gray, 1824) Pleurobranchidae Musculus sp. Pleurobranchaea sp. Mytilus edulis Linnaeus, 1758 Nudibranchia Mytilus sp. Dendronotidae Pterioida Dendronotus frondosus (Ascanius, 1774) Pectinidae Bivah l.i Chlamys islandica (Muller, 1776) Palaeotoxodonta Cyclopecten pustulosis Verrill, 1873 Nuculoida Placopecten magellanicus (Gmelin, 1791) Nurulidae Anomiidae Nucula delphinodonta Mighels and Adams, 1842 Anommia simplex Orbigny, 1842 Nucula proximo Sav, 18'22 Anomia squamula Linnaeus, 1758 Nucula tenuis Montagu. 18(18 Anomia sp. Nucula sp. Limidae Malletiidae Limatula siihauriculaln (Montagu, 1808) Malletia obtusa GO. Sars, 1872 Limatula sp. Satumia mbovata Verrill, and Bush, 1897 Heterodonta Nuculanidae Veneroida Nuculana acuta (Conrad, 1831) Lucinidae Nuculana pemula (Muller, 1771) Lucinoma hlakeana Bush. 1883 Nuculana tenuisulcata (Couthouy, 1838) Lucinoma filosa (Stimpson, 1851) Nuculana sp. Lucinoma sp. Portlandia fratema (Verrill and Bush. 1898) Thyasiridae Portlandia frigida (Torrell, 1859) Thyasira equalis Verrill and Bush, 1898 Portlandia inconspicua (Verrill and Bush 1898) Thyasira ferruginea Winckworth, 1932 Portlandia mflata (Verrill, and Bush. 1897) Thyasira Jlexuosa (Montagu, 1803) Portlandia iris (Verrill and Bush, 1897} Thyasira flexuosa forma gouldii Philippi, 1845 Portlandia lenticula (Moller, 1842) Thyasira pygmaea Vet rill and Bush, 1898 Portlandia lucida (Loven, 1846) Thyasira subovata Jeffreys, 1881 Yoldia limatula (Saw 1831) Thyasira trisinuata Orbigny, 1842 Yoldia m\aln (Couthouy, 1838) Thyasira sp. Yoldia regularis Verrill, 1884 Lasaeidae Yoldia sapotilla (Gould, 1841 ) Aligena elevata (Stimpson, 1851) Yoldia thraciaeformis Storer, 1838 Leptonidae Yoldia sp. Montacuta sp. Cryptodonta Carditidae Solemyoida Cyclocardia borealis Conrad, 1831 Solemyacidae Cyclocardia sp. Solemya velum Sav, 1822 Astartidae Pteriomorphia Astarte borealis (Schumacher, 1817) Arcoida Astarte castanea (Sav. 1822) Arcidae Astarte crenata subequilatera Sowerby, 1854 Anadara oualis (Bruguiere, 1789) Astarte elliptu a (Brown. 1827) Anadara transversa (Say, 1822) Astarte montagui (Dillwyn, 1817) Bathyarca anomala (Verrill and Bush. 1898) Astarte nana Dall, 1886 Bathyarca pectunculotdes (Scacchi, 1833) Astarte quadrans Gould, 1841 Bathyarca sp. Astarte smillin Gould, 1841 Limopsidae Astarte unilata Gould, 1841 Limopsis affinisVeniW, 1885 Astarte sp. Limopsis cristata Jeffreys, 1876 Limopsis minuta Philippi, 1836 i on ti nued on next page 24 NOAA Technical Report NMFS 140 Table 4 (continued) Cardiidae Cerastoderma pinnulatum (Conrad, 1831) Scaphopoda Dentaliidae Laevicardium mortoni (Conrad. 1830) Bathoxiphus ensiculus (Jeffreys. 1877) Ma< tridae Cadulus agassizit Dall, 1881 Mulinia lateralis (Say, 1822) Cadulus cylindratus Jeffrevs, 1877 Spisula solidissima (Dilhwn. 1817) Cadulus pandionis Verrill and Smith, 1880 Mesodesmatidae Cadulus rushii Pilshn and Sharp, 1898 Mesodesma sp. Cadulus sp. Solenidae Dentalium entale stimpsoni Henderson, 1920 l-iius directum Conrad, 1843 Dentalium meridionals Pilsbr) and Sharp. 1897 Siliqua costata Say, 1822 Dentalium occidentals Stimpson, 1851 Tellinidae Dentalium sp. Macoma calcarea (Gmelin, 1791) Cephalopoda Macoma sp. Octopus sp. Tellina agilis Stimpson, 1858 Tellina sp Rossia sp. ARTHROPODA Arcticidae I'm nogonida [rctica islandka (Linnaeus, 1767) Acheha scabra Wilson, 1880 Veneridae Achelia spinosa (Stimpson, 1853) Gemma gemma (Totten, 1834) Anoplodactylus lentus Wilson, 1878 Mercenaria mercenaria (Linnaeus, 1758) Nymphon brevitarseKreyer, 1844 Pilar morrhuanus Linsley, 1845 Nymphon grossipes (O. Fabricius?) Krover, 1780 Myoida Myidae Nymphon hirtipes Hell, 1853 Nymphon macrum Wilson, 188(1 Mya arenaria Linnaeus, 1758 Nymphon stroemi Krover, 1844 Corbulidae Paranymphon spinosum Caullery, 1896 I'nrliiiln contracta C.B. Adams, 1852 Pycnogonium littorale (Strom, 1762) Corbula sp. Crustacea Hiatellidae Cirripedia Cyrtodaria siliqua (Spengler, 1793) Balanus sp. Hiatella arctica (Linnaeus, 1767) Lepas sp. Hiatella striata (Fleuriau, 1802) Copepoda Hiatella sp. Calanus sp Panomya arctica (Lamarck, 1818) Caligus sp. Panomya sp. Cumacea An omalodesmata Brachydiastylis resima (krover. 1846) Pholadomvoida Campylaspis «/ \ _N* W £-~ ■' \ -.CONNECTICUT EXPLANATION MEAN NUMBER OF ANIMALS PES SQUARE METER OF BOTTOM ALL TAXA COMBINED 347 SLOPE ?3* NEW • NEW ..' \ \ JERSEY YORK/ \ V 9&i ' \ \ CONNECTICUT'. V ■jMASSACHUSE ,/\ iOSTi R ■ new y HAMPSHIRE S PERCENTAGE COMPOSITION 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 of Rhode Island and New York. Gen- erally, they occurred around the periphery of the Gulf of Maine and off southern 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. 3 2 NOAA Technical Report NMFS 140 Table 6 Mean number of specimens of each taxon per square meter in relation to geogi aphic area. Geographic areas Southern Southern New England New England Taxon Nova Scotia Gulf of Maine Georges Bank Shell 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 in 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 ANNELIDA 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 PRIAPL'LIDA — — — — 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 1.0 111 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 Ar.u hnida — — <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 ( irripedia 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 280.0 118.2 952.9 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 — 111 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.S 2.6 1.3 16.3 Ascidiacea 2.8 2.3 33.8 26.8 2.6 1.3 16.3 UNIDENTIFIED 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 ,51 2 2 Composition and Distribution of Macrobenthic Invertebrate Fauna 33 Table 7 Tin- number of specimens of each taxon, expressed as a percentage of the total benthic invertebrate fauna, in relation to geographic area. G eographic 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 Hvdrozoa 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 ill <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 ECH1NODERMATA 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 of spe rimens of each taxon (grams pe r square meter) in relation to geographic area. Gei >graphic areas Southern Southern New England New England l.ixon Nova Scotia Gull of Maine Georges Bank Mull G eorges Slope Slope All areas PORIFERA 15 49 3.15 0 47 nun 0.24 0.03 2 24 COELENTERATA 20.23 11.87 3.68 4.62 2.42 0.99 7.33 Hvdrozoa 0.49 0.12 1 i.l (1.41 <0.01 0.03 0.52 Anthozoa 19.74 11.75 2.07 4.21 2.42 0.96 6.81 All \ i maria 0.03 0.43 — 0.21 0.14 0.07 0.20 Zoantharia 19.54 10.90 2.04 3.84 1.9(1 0.70 6.39 Unidentified u |i. 11.41 0.03 0.15 0.38 0.19 0.22 PLATYHELMINTHES • mil 0.01 001 11.01 0.01 — 0.01 Turbellaria <(l.(il II. (II IIOI 0.01 (1.01 — 0.01 NEMERTEA 0.56 0.54 0.83 1.04 0.11 0.20 0.71 ASCHELMINTHES ii.nl 0.01 0.01 o.ol 0.01 0 III 0.01 Nematoda <0.01 0.01 0(11 0.01 0.01 0.01 0.01 ANNELIDA IS Ml 15.51 7.93 _" 11,11 4.86 4.32 17.41 POGONOPHORA ■ 0. Ill <0.01 — — 0.01 0.03 <0.01 SIPUNCULIDA 1.65 0.37 0.46 0.74 fill 1 ,83 0.75 ECHIURA — (1.01 — 0.07 5.18 0.40 0.03 PRIAPULIDA — — — — 0.01 0.05 <0.01 MOLLUSCA 54.40 $1 ".'i 79 ", l 170 on 2.65 1.18 83.64 Polyplacophora 0.10 0.19 IIOI 0.24 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 10 166.34 2.11 1.04 80.95 Scaphopoda 1.03 ii i.i. 0.15 0.02 n -ii DOS 0.32 Cephalopoda <0.01 — 0.01 (I 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 11.01 <0.01 — 0.01 Arai hnida — — <0.01 — — — <(l(ll Crustacea 16.49 2.41 'i 7", 17.11 1)1,1 0.13 9.40 Ostracoda /. 1.1 1.5 0.2 0.3 0.3 0.2 — ■ (1 1 0 1 0.4 Anthozoa 0.8 4.1 1.0 1.0 1.0 2 2 3.5 2d 1.7 Alcyonaria — — <0.1 ii. 1 (1.4 0.4 1.0 0.6 0.1 Zoantharia (1.2 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 <0.1 111 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 11 2.3 0.2 ANNELIDA 28.7 19.6 2". II 5(1.3 47.0 39.1 24.7 12.2 28.1 POGON'OPHOR\ — — — <0.1 0.2 2.5 2.0 4.0 <0.1 SIPUNCULIDA ii 1 0.3 ii I 0.8 1.6 0.5 0.6 1.0 0.4 ECHIURA <0.1 — — <0.1 — — 0.4 (IS <0.1 PRIAPULIDA — — — — — — 0.2 0.1 <0.1 MOLLUSCA 22.8 9.2 8.7 15.0 18.0 44.6 45.0 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 ARTHROPOD A 41.3 56.3 VI 'I 18.6 13.3 5 4 10.8 8.8 48.0 Pycnogonida <0.1 <0.1 <<).] t specimens of each taxon (granu per square meter) in relation to water depth. Taxon Water depth (m) 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 COELENTERATA 3.63 1.49 3.38 18.95 9.13 0.36 (1.72 0.69 7.33 Hvdrozoa 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 0.25 001 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 Pl.ATYHELMINTHES 0.02 <0.01 0.01 — <0.01 — — — 0.01 Turbellaria 0.02 <0.01 0.1)1 — <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 Dill 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 ANNELIDA 27.22 16.24 25.20 14.76 10.70 4.76 1.41 (171. 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 PRIAPUI.IDA — — — — — — 0.12 0.01 <0.01 MOLLUSCA 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 — ■ 11.111 <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 2.10 112 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 1 1 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 Holothuroidea 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 of spe cimens ol each taxon expressed as a percentage of th e total be nthic inverte brate fauna. in relation to water depth. Taxon Depth zones (m) ( )-'.' 4 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 (i.l 0.5 0.2 <0.1 — <0.1 <0.1 0.3 Anthozoa 0.5 0.4 1.0 19.1 14.0 2.4 5.3 0.1 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 <(). 1 <0.1 0.2 0.9 0.9 2.1 4.3 0.1 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 0.5 0.4 0.2 0.0 0.1 0.4 1.0 0.4 ASCHELM1NTHES <0.1 <0.1 <0.1 <0.1 <0.1 0.1 0.1 (I.l <0.1 Neniatoda <0.1 <0.1 <0. 1 <0.1 <0.I 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 1(1.1 4.7 0.4 ECHIL'RA 0.1 — — <0.1 — — 36.9 31.0 0.2 PRIAPULIDA — — — — — — 0.9 (i.l <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 i.l 0.3 1.9 11 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 1 the tota benthic invertebrate fauna in relation to bottom sediments. Taxon 15. tiom sediments Gravel Till Shell Sand 5 and-silt Silt-clay All types PORIFERA 1.8 19.0 2.6 0.1 1.8 o 1 1 2 COELENTERATA 8.5 2.7 0.4 0.8 10.7 1 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 Zoanthai ia 6.8 1.6 0.3 0.6 10.2 3.7 Unidentified 0.1 0.6 <0.1 <0.1 0.4 ■ 0.1 o.l PLATYHELMINTHES <0.1 — — <0.1 — <0.1 <0.1 Turbellaria <0.1 — — «>.l — <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.I <0.1 <0.1 <0.1 Nematoda <0.1 <0.1 — «>.l <0.1 <0.1 <0.1 ANNELIDA 8.6 15.9 6.9 6.1 15.3 18.9 9.5 POGONOPHORA — — — <0.1 <0.1 () 17 :.l 28 16 Amphipoda 93 77 100 94 74 51 Mvsidacea 4 — — (. 3 1 Decapoda 35 — 33 37 7 4 BRYOZOA 27 32 50 8 10 4 brachiopoda II 41 17 2 3 3 ECHINODERMATA 51 86 50 72 78 78 Crinoidea — — — — 1 <1 Holothuroidea 16 30 17 8 36 23 Echinoldea 1 t 32 17 47 8 15 < (phiuroidea Id 68 33 29 (.1 62 Asteroidea 8 'i — 1 l 16 14 HEM1CHORDATA — — — ■ 1 1 <1 ( HORDATA 22 32 17 19 15 1 1 Am idiacea 22 32 17 19 15 11 Composition and Distribution of Macrobenthic Invertebrate Fauna 51 i$& % * t. jr-r, Kt 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 dif- ferent bottom types in different subareas within the New England region. Sediment types range from coarse (gravels and cobbles) to fine (silty sands). The camera- tripping weight, visible in each photograph, serves as a possible indicator of the amount of silt-clay 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 macro- benthos, in general, was related directly to the annual range in water temperature. In areas with a small an- nual 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 den- sity 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/irr and biomass only 46 g/m2. Although there are some inconsistencies, in 52 NOAA Technical Report NMFS 140 .? «r,- «'*• j ' ' m cv Figure 19 Blown silt\ sand bottom .it 1 16 m depth southwest of Grand Marian Island. A substantial number nt brittlestars (Ophiuroidea) oi various sizes an- clearly visible, as are several burrow holes. The tine nature ot the bottom at this locality is indicated by the cloud of sediment raised by the camera tripping weight striking the bottom at upper left. Photograph was taken at station 1 173. located at lat. 44L>S' V, long. 67°15' W. general, density increased to 4,038/m2 where the tem- perature range was greater than 20°C, and biomass rose to 467 g/m2 where the range was l(i to 19.9°C. This close relationship of sparse fauna in stable temperature areas and rich fauna in unstable areas undergoing wide temperature fluctuations is not necessarily a direct cause- and-effe< t relationship of temperature alone but is the result of a combination of temperature and other envi- ronmental factors. Water masses that remain relatively constant in temperature and are unchanging in most other physical and chemical properties also tend to have 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. I he 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 Cull oi Maine cast 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-renter. 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 con- sidered 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 he high in organic carbon due to biogenic activi- ties (fecal deposits, mortality, etc.). Regardless which prevailed, high organic carbon content would be asso- ciated with high abundance. The analysis did not reveal any clear-cut correlation between sediment organic carbon content and benthic faunal abundance. A few fannal 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 mi ^wr m % I * ■ . mt «. ^ ^^^^^^^H'~|^S 4 %«»- ^tf^<^p^pnp^3pj*jHpjjFA ■;, "'%,.-, ; - '. "fc*"^"W JgS ~* '« V « - *» ^ft A^^n-^ . 1^^ 4 4 - ^•■" . ' ■ , .'!.-■ - -^r ;;: ":-- ^ • •c i, -> * '; ji.'^'T** '" . H SI ■ . ' pi » l; m 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' V. 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 densi- ties (44/m2) 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 die 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 c' *&l ( Figure 22 Coarse brown sand with broken shells and gravel, to 15 cm, 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 1 13(1. located at lat. 4201' V. long. 66°31' W. Biomass of 218 g/rrr 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 percentage 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 be- tween 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 com- ponent of the New England region macrobenthos ar- ranged in the phvlogenetic 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 fol- lowed by discussions dealing with geographic and bathy- metric distribution and the quantitative relationship of 56 NOAA Technical Report NMFS 140 mmm/ * ^HnHH|HHH[ If -* JH IB '''*■' * f * ■« k ■ ■ ' MU9 iffflnr 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 (Echinarachnius parma) 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 con- tinental shelf and uncommon to rare on the continen- tal 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 Macrobcnthic Invertebrate Fauna KUHuaowaH ">v X^j? >, *^M*J Figure 24 Green silty sand and gravel bottom at 450 m depth on the southern New England continental slope. Gravel ranged to 2 cm. The tubes of two soda straw worms (Hyalhwecia 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' N., long. 72°09' W. Porifera were most common in the Nova Scotia re- gion 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/nr and biomass averaged 2.2 g/nr (Table 5). Geograph ic Distrib ution 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 New Jersey was especially noteworthy. 58 NOAA Technical Report NMFS 140 Table 21 Mean number of spe ;imens of each taxon per square meter in relation to the annual range in bottom water terr perature. Taxon Annual range in water temperature (degrees Celsius) 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 44.8 93.9 32.1 Hydrozoa 0.9 5.0 1 5 4.5 36.8 45.9 6.4 Anthozoa 5.7 9.7 19.5 100.8 8.0 48.0 25.7 Alcvonaria 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 l).7 0.8 43.6 2.2 PLATYHELMINTHES — 0.2 1.0 111 <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 SIPUNCULIDA 4.5 9.0 8.5 2.8 2.6 — 5.9 ECHIl'RA 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 Polvplacophora 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 11(1.1 329.0 252.0 1,194.8 163.1 Scaphopoda 8.9 10.3 2.1 0.8 0.2 — 5.1 Cephalopoda <0.1 0.1 1.1 — — — 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 11.7 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 — — I) 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 — (12 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 1(14.7 171.2 31.5 21.(1 79.3 Crinoidea <0.1 — — — — — <0.1 I [olothuroidea 6.3 4.1 3.7 2.8 17 3.5 4.3 Echinoidea 14 4.9 40.4 93.1 26.8 14.6 29.3 Ophiuroidea 2 1 8 50.4 57.9 74.1 2.(1 2.4 44.2 Asteroidea 0.4 2.2 2.7 1.2 0.9 0.5 1.5 HEMICHORDATA <0.1 — ill' (12 — — 0.1 ( HORDATA 2.0 2.3 35.1 6.0 58 3 4.3 16.3 Am idiacea 2 11 2.3 35 1 6.0 58.5 4.3 16.3 UNIDENTIFIED 5.6 5.7 6.6 4.2 i, 6 5.2 5.8 Total 432.2 1,097.9 2.386.2 1.746 .4 1,934.2 4,037.5 1,512.2 Composition and Distribution of Macrobenthic Invertebrate Fauna 59 Table 22 The number of specimens of each taxon, expressed as a percentage of the to tal benthic invertebrate fauna. in relation to the annual range in bottom water temperature. Taxon A initial range in water temperature (degrees Celsius) 0-3.9 4-7.9 8-11.9 12-15.9 16-19.9 20-23.9 All ranges PORIFERA 0.3 (1.2 <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 Hvdrozoa 0.2 0.4 0.2 0.3 1.9 1.1 0.4 Anthozoa 1.3 0.9 0.8 5.7 0.4 1.2 1.7 Alcvonaria 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 1.1 0.2 PIATMTELMINTHES <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 0.6 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 o 2 0.2 <0.1 <0.1 — 0.2 ANNELIDA 48.9 46.7 23.8 16.0 19.2 42.0 28.1 POGONOPHORA 0.3 0. 1 <0.1 — — — <0.1 SIPUNCULIDA 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 lis 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 irs 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 Mvsidacea 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 — .(>1 — — <0.01 Cumacea 0.02 0.04 0.26 0.07 0.09 0.06 oil Tanaidarea <0.01 <0.01 — — — — <0.01 Isopoda 0.27 0.08 0.42 0.34 0.07 0.41 o 29 Amphipoda 0.41 2 02 8.12 5.31 6.27 2.56 4.16 Mvsidacea <0.01 <0.01 0.02 0.02 0.03 0.02 001 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 ECH1NODERMATA 12.24 31.35 35.48 172.83 59.63 263.42 55.00 Crinoidea •eO.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 ._, 82 3.33 3.61 0.46 2. 1 3 HEM1CHORDATA <0.01 — 0.04 0.03 — — 0.02 CHORDATA 0.55 4 14 7.28 1.43 11 40 0.56 4 10 Ascidiacea 0.55 4.14 7.28 1.43 14.40 0.56 1 1(1 UNIDENTIFIED 0.16 0.22 0.36 0.42 (1 26 0.09 0.27 Total 45.48 88. 1 1 230.67 317.62 467.42 422.12 183.39 Composition and Distribution of Macrobenthic Invertebrate Fauna 61 Table 24 The wet weight of specimens ol each taxon. expressed as a percentage of the otal benthic invertebrate fauna. in relation to the annual range in bottom water temperature. Taxon Annual range in w ater temperature (degrees Celsius) 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 (1.1 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 <0.1 500 m (Table 1 1; 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 NUMBER PERCENT WEIGHT MEAN PERCENTAGE ORGANIC PERCENTAGE MEAN ALL TAX A COMPOSITION CARBON COMPOSITION ALL TAXA 221 1,838 741 1,015 914 2,042 2,588 44 o.oo 0.01 to 0.49 O.SO to 0.99 1.00 to 1.49 l.SO to 1.99 2.00 to 2.99 3.00 to 4.99 5.00 to ♦ EXPLANATION 17 218 96 97 89 959 809 < 1 CBU5TACEA \\'>S', MOLLUSC A ECHINODERMATA □ 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 compo- sition, bv 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/m . In deeper water the average weights were only 0.02 to 0.6 g/m". In all depths Porifera accounted for a small percent- age (<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 (%) )f each taxonomic group in the samples in each tenipei ature range class. Taxon Annual range in water temperature (degrees Cel sIUS) 0-3.9 4-7. 'J 8-11.9 1 2-15.9 16-19.9 20-23.9 POR1FERA 8 15 4 2 8 4 COELENTERATA 36 4'.' 45 45 49 29 Hydrozoa 4 13 15 14 26 18 Anthozoa 32 29 30 31 23 1 1 Alcyonaria 1 1 5 2 8 — — Zoantharia 17 29 30 28 24 14 PLATVHELM1NTHES — 1 3 1 2 4 Turbellaria — 1 3 1 2 4 NEMERTEA 25 28 40 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 1 — 3 — PRIAPLL1DA 1 — — — — — MOLLUSCA 91 83 87 87 94 93 Polyplacophora in 1(1 6 7 5 — Gastropoda 44 46 4 1 38 57 43 Bivalvia 87 75 8 1 83 9(1 93 Scaphopoda 36 37 10 3 2 — Cephalopoda <1 2 <1 — — — ARTHROPODA 70 89 98 98 90 86 Pvcnogonida 1 4 3 1 2 4 Arachnida — 1 — — — Crustacea 7(1 89 98 98 90 86 Ostracoda 1 1 — — — Cii ripedia 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 1 1 21 18 BRACHIOPODA 10 8 2 — — — ECH1NODERMATA 77 78 72 71 45 43 Crinoidea 1 — — — — — Holothuroidea 25 20 1 I 18 8 21 Echinoidea 16 26 33 44 24 1 1 Ophiuroidea 64 62 35 29 15 11 Asteroidea 6 16 21 12 1 1 4 HEMICHORDATA <1 1 1 — — CHORDATA 12 1 1 24 17 19 1 1 Ascidiacea 12 1 1 24 17 19 11 Composition and Distribution of Macrobenthic Invertebrate Fauna 65 Table 26 Mean number ofspe dmens of eat h taxon p er square metei in relation to sediment organic carb on content. Taxon Sediment organic carbon content (percent) 0.00 0.01-0.49 0.50-0.99 1.00-1.49 ,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 Alcvonaria — 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 PLATYHELMINTHES — 0.6 0.1 — — 0.9 — — 0.4 Tin bellai i.i — 0.6 0.1 — — 0.9 — — 0.4 NEMERTEA 10.8 5.5 4.4 4.4 3.5 1.7 — — 4.9 ASCHELMINTHES — 3.1 1.0 2.8 0.4 — — — 2.3 Nematoda — 3.1 1.0 2.8 0.4 — — — 2.3 ANNELIDA 64.0 503.6 234.7 319.1 195.9 400 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 9 i» 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 Slid 34.4 sis. 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 1.065.9 220.1) 85.9 34.4 SI 8.3 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 Gopepoda — <0.1 <0.1 0.1 — — — — <0.1 Cumacea 2.6 25.2 6.1 3.8 6.2 122.S — — 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.S 21.4 1.2 1.7 35.1 — 26.5 — 14.9 BRAGHIOPODA 16.(1 5.6 5.1 2.8 — — — — 4.7 ECHINODERMATA 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 y y 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 UNIDENTIFIED — 6.3 6.6 6.8 3.1 3.0 — 11.0 6.1 Total 221.2 1.S5S 1 740.7 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 of specimens of each taxon, expressed as a percentage of the total benthic invertebrate fauna, in relation tc sediment organic carbon content. Sediment organic carbon content (percent) Taxon o.oo 0.01-0.49 0.50-0.99 1.00-1.49 1.50-1.99 2.00-2.99 3.00-4.99 All range1' PORIFERA 1.2 0.1 COELENTERATA — 1.0 Hvdrozoa — 0.5 Anthozoa — 0.5 Akvonaria — <0.1 Zoantharia — 0.4 Unidentified — 0.1 PLATYHELM1NTHES — <0.1 Turbellaria — <0.1 NEMERTEA 4.9 0.3 ASCHELMINTHES — 0 2 Nematoda — 0.2 ANNELIDA 28.9 27.1 POGONOPHORA — <0.1 SIPUNCULIDA 1.0 0.4 ECHIURA — <0.1 PRIAPULIDA — <0.1 MOLLUSCA 31.1 7.5 Polyplacophora — 0.1 Gastropoda 1.1 11 Bivalvia 29.0 (1.1 Scaphopoda 1.0 0.2 Cephalopoda — <0.1 Unidentified — <0.1 ARTHROPODA 9.5 57.4 Pvcnogonida — <0.1 Arachnida — .] 47.5 0.1 2.5 43.6 1.2 8.5 <0.1 8.5 <0.1 0.4 0.3 7.7 o.l 0.2 0.3 8.9 0.8 0 1 7.9 0.2 <0.1 in. Ill, 0.7 100.0 <0.1 0.6 o.l 0.5 0,5 <0.1 0.4 II. 1 0.1 21.4 <0.1 <0.1 65.9 <0.1 4.9 59.7 1.3 3.8 3.8 0.7 0 1 2.7 0.2 0.1 3.9 3.7 0.2 o 1 3.3 0.1 0.3 100.0 0.3 0.1 0.1 0.1 <0.1 <0.1 o.l 19.9 38.9 1.9 36.8 40.1 40.1 30.0 6.0 0.4 3.2 <0.1 0.4 0.3 0.1 0.2 0.2 0.2 0.2 100.0 3.1 43.3 43.3 3.3 0./ 48.5 1.0 0.1 o.l 100 II — o.l — 1.2 — 0.5 — 0.7 — <0.1 — 0.5 — 0.2 — <0.1 — <0.1 — 0.3 — 0 2 — 0.2 25.0 27.4 — 0.1 — 0.4 — — <0.1 1.3 — 0.7 50.0 43.2 — 0.1 — 0.3 — <0.1 — in — 0.3 — 4.4 — <0.1 — 0.3 — 1.0 — 3.0 — 0.1 — <0.1 — 1.0 — 1.0 25.0 0.4 100.0 100.11 Composition and Distribution of Macrobenthic Invertebrate Fauna 67 Table 28 Mean wet weight of specimens of each taxon (grams per square meter) in relation to sediment organic carbon content. Sediment organic carbon content (percent) 1 .1X1111 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 rORIFERA 0.03 5.19 COELENTERATA — 6.33 Hydrozoa — 1.06 Anthozoa — 5.27 Alcyonaria — 0.26 Zoantharia — 4.91 Unidentified — 0.10 PLATVHELMINTHES — <0.01 Turbellaria — <0.01 NEMERTEA 0.53 0.79 ASCHELMINTHES — 0.01 N'ematoda — 11.111 ANNELIDA 7.43 15.93 POGONOPHORA — <0.01 SIPUNCULIDA 0.02 0.93 ECHIURA — 0.01 PRIAPL I.IDA — <0.01 MOLLUSCA 0.80 132.21 Polvplacophora — 0.25 Gastropoda 0.02 3.83 Bivalvia 0.67 127.81 Scaphopoda 0.11 0.31 Cephalopoda — Dili Unidentified — — ARTHROPODA 0.31 13.28 Pvcnogonida — 0.01 Arachnida — <0.01 Crustacea 0.31 13.27 Ostracoda — <0.01 Cirripedia — 6.45 Copepoda — <0.01 Cumacea 0.03 0.09 Tanaidacea — <0.01 Isopoda 0.19 0.27 Amphipoda 0.09 5.29 Mvsidacea — 0.01 Decapoda — 1.16 Unidentified — <0.01 BRVOZOA 0.23 1.95 BRACHIOPODA 1.31 0.96 ECHINODERMATA 6. 1 6 35.22 Crinoidea — <0.01 Holothuroidea 5.80 3.97 Echinoidea 0.29 27.90 Ophiuroidea 0.07 2.32 Asteroidea — 1 .03 HEMICHORDATA — 0.03 CHORDATA — 4.38 Ascidiacea — 4.38 UNIDENTIFIED — 0.33 Total 16.82 217.56 0.12 18.48 11.16 18.32 0.32 17.21 0.79 <0.01 <0.01 0.79 <0.01 <0.01 14.88 0.01 1.13 1 .25 0.02 25 23 0.03 0.56 24.42 0.19 11.03 <0.01 6.46 0.01 6.45 <0.01 4.11 <0.01 0.03 <0.01 0.24 1.76 II n I 0.30 <0.01 0.02 1.74 23.25 <0.01 11.16 5.44 5.23 1.42 0.03 1.88 1.88 0.27 95.95 0.01 14.57 0.01 14.56 0.13 14.07 0.36 0.38 0.02 0.02 12.92 0.01 0.07 1.35 0.01 19.95 0.02 0.28 19.17 0.48 1.22 <0.01 1.22 <0.01 0.02 0.03 0.59 0.58 0.06 0.03 44.01 30.95 1.33 6.28 5.45 <0.01 2.03 2.03 0.29 96.93 <0.01 1.80 <0.01 1.80 1.79 0.01 0.94 <0.01 <0.01 26.92 <0.01 <0.01 13.38 <0.01 1.09 12.05 0.24 1.02 1.02 0.01 0.14 0.17 0.01 0.69 1.21 43.35 40.90 0.60 0.56 1.29 0.10 88.72 0.06 0.02 0.04 0.04 0.19 0.19 0.48 22 1. 1 1 811.54 10.42 801.02 0.10 18.99 18.99 14.32 0.24 0.06 0.21 0.01 4.15 104.47 104.31 0.16 0.33 0.33 0.07 958.73 11.31 226.87 226.87 9.44 9.44 1.10 0.05 8.29 0.18 561.51 561.51 809.31 — 2.98 — 9.57 — 0.64 — 8.93 — 0.24 — 8.42 — 0.28 — 0.01 — 0.01 — 0.74 — 0.01 — 0.01 0.11 16.01 — <0.01 — 0.79 — 0.44 — 0.01 — 99.58 — 0.15 — 2.59 — 96.54 — 0.29 — 0.01 — <0.01 0.11 9.62 — <0.01 — <0.01 0.11 9.61 — <0.01 — 4.86 — <0.01 — 0.07 — <0.01 — 0.23 0.11 3.54 — 0.01 — 0.91 — <0.01 — 1.19 — 0.95 — 38.82 — <0.01 — 15.66 — 17.29 — 3.26 — 1.61 — 0.03 — 3.15 — 3.15 0.11 0.29 0.33 183.18 68 NOAA Technical Report NMFS 140 Table 29 The wet weight of specimens of each taxon, expressed as a percei tage of the total bentbic invertebr ate fauna, in relation to sediment organic carbon content. Taxon Sediment organic carbon content (percent) 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 2.0 <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 II 1 — — — — 0.1 Zoantharia — 2.3 18.0 14.5 2.(1 <0.1 — — 4.6 Unidentified — <0.1 0.8 0.4 <0.1 — — — 0.2 PLATYHELMINTHES — <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 1.1 — — — — 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 Polvplacophora — 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 0.2 115 0.3 NEW \ NEW JERSEY i Y0HK GRAMS PER SQUARE METER PORIFERA Figure 27 Geographic distribution of Porifera: A — number of specimens per square meter of bot- tom; 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 fre- quency of occurrence was on the continental slope (200 to 2,000 in). 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 consti- tuted a substantially larger biomass in coarse substrates Composition and Distribution of Macrobenthic Invertebrate Fauna 71 (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 sub- strates. Till contained the highest density of specimens, the greatest weight, and highest fre- quency of occurrence. Gravel ranked second in number and shell ranked second in weight. Sand contained intermediate quantities, and sand-silt and silt-clay contained the smallest quantities. Relation to Water Temperature Porifera were generally most numerous in ar- eas 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 of occurrence was highest (15%) in those samples from lo- calities in which the seasonal changes in tem- perature were between 4° and 8°C. Two other aspects of the relative density of Porifera and bottom water temperature th.it 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 maxi- mum temperature was moderate, between 6° and 12°C; they were scarce (<1 g/m-') 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 con- tained low to moderate (0.01 to 1.99%) amounts of organic carbon (Tables 26-30; Fig. 32). Both mean density and biomass diminished with in- creasing organic carbon content. Density ranged from 3 to 0.2/m2, and biomass from 5 to <0.01 g/m2. Ui IE 3 CD < z => Z CO □ NUMBER ■ WEIGHT I n n I~L n- I" ? (E bj O GEORGES SOUTHERN GEORGES SOUTHERN BAN" NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 28 Density and biomass of Porifera in each of the six geographic areas. 0---0 NUMBER 5 • • WEIGHT 3 O - 4 2 t- tr) o z ts 2 CD LU 9 < Su. or u. G° • > ^^"* 8 - j O o rfo: ' '* / 'a z tr LOLU ' — ' ' / ' A (- LU o2 i \ 2^ UJ UJ PiS t 1 * / A * cc £< 6 / \ * \ < ^° i _ r ' '"' \ gg z U} / ^ \ 1 * UJ a. i • i i i i ' i »— i — * ' CL V 10 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 29 Density and biomass of Porifera in relation to water depth. Coelenterata The macrobenthic coelenterate fauna of the New En- gland region is composed of members of two classes: Hvdrozoa and Anthozoa. Hydrozoa are treated at the class level, whereas Anthozoa. composed of members from seven orders contained in two subclasses, are dis- cussed at the subclass levels: Alcvonaria and Zoantharia. No discussions are included at the taxonomic levels Coelenterata or Anthozoa; however, the interested reader will find summary data, for all treated para- meters, 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 contribu- tion to the total biomass. They made up less than 1% of 72 NOAA Technical Report NMFS 140 the benthic biomass (Table 3). Members of this group were most abundant in shallow wa- ters and were attached to firm substrates, fre- quently 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. Thev were exclusively carnivores preying on planktonic crustaceans and other small animals carried to them by water cur- rents. 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, Sertulaiia, Obelia, and others. Less common were representatives of Antho- medusae, 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/m and biomass averaged 0.5 g/mL (Table 5). Geographic Distribution Hydrozoans were common in coastal areas and on offshore banks (Fig. 33). Thev 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/ irr, and high biomass, 10 to 45 g/m2, occurred 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, Georges 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 Bank ranked first in terms of weight, and Nova Scotia ranked first in number of specimens. Small quantities of hydro- zoans 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 of speci- mens. Percentages of samples containing hydrozoans ranged from a high of 29% in the Nova Scotia area (8 to OS UJ Q; OD < 2 to 2 — □ NUMBER ■ WEIGHT I O o si < <* u. o o ? a. IL_ SHELL SAN0 I L SAND- SILT- SILT CLAY BOTTOM SEDIMENTS Figure 30 Density and biomass of Porifera in relation to bottom sediments. O O NUMBER 2 * « WEIGHT O f- \ NEW / «, ^ Y \ \ HAMPSHIRE .' S?0R" N. ^^CONNECT I CUT V V^ EXPLANATION □ <0.l - 0.9 M l0* " 0.0 - 45.3 GRAMS PER SQUARE METER 40° 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 die 12-15.9°C and 1 6-19.9°C classes: in die former the average density was 5 colonies/m2; in the latter it was 37 colonies/m2. The contribution of hydroids to the total faunal den- sity 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 tempera- ture 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 <0 o z o 3> £ u UJ O 2 0: UJ uj or GO < 3 O z tn 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 Hvdrozoan abundance was generally nega- tively 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 lev- els 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 bio- mass, however, in being parabolic with low- est occurrence in the middle ranges and increasing at each extreme. Anthozoa — Alcyonaria — AJcyonarian 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 con- stituted <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 speci- mens 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 of the majority of specimens in our samples was tan or a l I I NUMBER ■ WEIGHT GEORGES SOUTHERN BANK NEW ENGLAND SHELF GEOGRAPHIC AREA Cb_ GEORGES SOUTHERN SLOPE NEW ENGLAND SLOPE 3 O IT u. O O UJ UJ < I- 3 UJ o *<° or UJ Figure 34 Density and biomass of Hydrozoa in each of the six geographic areas. o 0 0 NUMBER > • • WEIGHT 5 O 5 O \ \ tn o 2 §30 2 CD '5 gu. = o \ o o UJ £C * \ *E u \ \ A LU *;U t- UJ \\ \ 10 I 2 os — UJ or W \\ / \ V / \ uj or NUMBE PER SQUAR 5 y- a V / \ 05^ or UJ 0. u0 10 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 35 Density and biomass of Hydrozoa in relation to water depth. combination of light cream (rachis) and deep bur- gundy (pinnae). Alcyonarians occurred in 63 samples (6% of total). Their density averaged 0.8/m 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 occur- rence was patchy, especially in the northern section 76 NOAA Technical Report NMFS 140 (Fig. 39). They were absent in the samples from Georges Bank. Their average density in all localities was between 1 and 15/m . 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 g/m2 (Tables 6-9; Fig. 40). In addition to number and weight per unit area, another reasonably good index of abun- dance is the percentage of samples contain- ing alcyonarians (Table 10). All shelf areas (Nova Scotia, Gulf of Maine, Georges Bank, and Southern New England Shelf) had a fre- quency of occurrence of 7% or less. The oc- currence rates on Georges Slope and South- ern 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. I | NUMBER ■ WEIGHT - 2 4 NUMBER OF SPECIMENS SQUARE METER OF BOTTOM - - WET WEIGHT IN GRAMS SQUARE METER OF BOTTOM OB £ 0. e — - H n n_ 04 4 - L ■ - il 1 1 n. pl GRAVEL TILL SHELL SAND SANO- SILT- SILT CLAY BOTTOM SEDIMENTS Figure 36 Density and biom ass of Hydrozoa in relation to bottom sediments. Bathymetric Distribution Alcyonarians were most common in deep wa- ter 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 den- sity and weight from one depth class to an- other were small, the larger quantities were most prevalent in deep water (100 to 2,000 m). Highest average density and largest aver- age weight, 1.9/m2 and 0.47 g/m2, respec- tively, occurred at depths from 200 to 500 m. The frequency of occurrence of alcyonar- ians 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. o o NUMBER O 50 • • WEIGHT ? ■> 5 o K K tft o J> "f, z m - / 1 °° 5° A ^ 2.0 a. u. o o £« Z (E "f 50 / \ //\ 1.5 UJ 1- 1- O 5 / \ / \ o 2 il UJ / \ i/ \ UJ uj uj cr 20 / \ / \ 1.0 Ser s = Z to / \ / UJ o <£ l0 / \ ^>* i 0.5 a. UJ a. i^^~ i i i i i 0 UJ 0. 0-39 4-79 8119 12-159 16-199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 37 Density and biomass of Hydrozoa in relation to the annua 1 range of bottom water temperature. Relation to Bottom Sediments Alcyonarians occurred only in selected types of sub- strate (Tables 1<>-19; Fig. 42). The Alcyonacea and Gorgonacea were collected from gravel and rocky out- crops, in contrast to the Pennatulacea, which inhabited sand-silt and silt-clay sediments. No alcyonarians were taken on substrates of till or shell, and they were present 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-clay and sand-silt sedi- ments 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 occur- rence exhibited a bimodal relationship. Composition and Distribution of Macrobenthic Invertebrate Fauna 77 s e.o o u. UJ O X a: uj uj a: S3 => o --o NUMBER — • WEIGHT 10 IS 2 0 3.0 PERCENT ORGANIC CARBON S o ? uj cr /A CE U. 04 o O cnH / ' a t- UJ l- uj o 2 / ' \ % 5* £s l0 /-/ V°"A 02*5 |o a \ / V h o =>c/> y S / \f v> i IT fc w o • • WEIGHT 5 £ Z CD < m o w £<* S UJ §fe 0.6 2 (E UJ h- 1- u. UJ O 5 2 04 Z UJ O X IX UJ UJ or \ ^ A UJ u < X 3 => O 1 0? UJ o z <0 \ ^ // N^ * « •***— \ y \* K "- 0 1 1 1 1 \ Jp 0 0-39 4-79 8-119 12-159 I6-F99 20-539 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 43 Density and biomass of Alcvonaria in relation to the annu; il range of bottom water temperature. (Tables 21, 22; Fig. 49). Density was highest (98/m2) in areas with a moderate (12°-15.9°C) temperature range, and densities decreased to about 2 to 4/m where the tem- perature range was both narrow (<4°C) and broad (>20°C) . The relation between zoantharian biomass and tem- perature 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 bv the percentage of the total benthic fauna that is made up by zoantharians (Table 24). Where the temperature range was less than 4°C, zoantharians 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. Composition and Distribution of Macrobenthic Invertebrate Fauna 81 « o Zffi OS (t UJ UJ or IS ig IT £ 0.5 --o NUMBER • WEI8HT 0 15 2 0 3 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 of individuals ranging from 3 to 7/m2. 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 indi- viduals 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 1-mm mesh sieving screen were present in very low density. The vast majority of marine tubellarians reported from New England marine waters are less than a few millimeters in length. Specimens in our samples 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/m , and their biomass aver- aged 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 9/m 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/m''- or less. The density of turbellarians in each geographic area (Tables 6, 7; Fig. 52) averaged less than 1/trr. The 82 NOAA Technical Report NMFS 140 NEW \ NE'W ,.X\ \ ,X JERSEY WORK/ '- \ NEW / -«^» ^CONNECTiCUT'- \ j pooilAnd*— -j. ^f /\. 'V \. \mASSACHUSETTSV^*-^^~ ,) id V— -^ ^>w_ ^>-^\ /• BOSTON £ f****T*' T MAINE 0 120 74° , / ( o ^ w r« >r si — ^ r tooo^-._^^ "s^-. 78° ** > EXPLANATION M 1-49 BSI 50-99 . J ^\g NOVA SCOTIA 7 r" ■ 100-572 ^^^^r* SZ^j^~^~ ~^^X/J/ ^"~-~--^^ INDIVIDUALS PER SQUARE METER \ / /\A S N ' \ \ H<0.l- 9.9 H 10.0-49.9 ■ 50.0-1561.3 GRAMS PER SQUARE METER ZOANTHARIA Figure 45 Geographic distribution of Zoantharia: A — number of specimens per square meter of bottom; B — biomass in grams per square meter of bottom. southern New England Shelf area contained the high- est density (0.9/m'-); Georges Slope ranked second with 0.4/m'-; 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 ttirbel- larians ranged from 0 to 3% (Table 10). Bathymetric Distribution Turbellarians were most plentiful (2.6 specimens/m-') in shallow water, uncommon at depths from 25 to 500 Composition and Distribution of Macrobenthic Invertebrate Fauna 83 GEORGES SOUTHERN GEORGES SOUTHERN SANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 46 Density and biomass of Zoantharia in each of the six geographic areas. A o 0 NUMBER • • WEIGHT s O 80 20 * OF SPECIMENS METER OF BOT o o - / GHT IN GRAMS E METER OF BO Eg / \ 1 \ *§ i- O UJ CO z a. 20 _ 1 5 *cr ' UJ 0. I \ \ °0 10 25 50 100 200 500 I00O 2000 4000 " WATER DEPTH IN METERS Figure 47 Density md h iomass of Z lantharia 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 g/ 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-clay their average den- sity was 0.2 and 0.1 /m- (Tables 16, 17; Fig. 54). In terms of biomass they were equally sparse (averag- ing 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 co < 2o \~~\ NUMBER WB WEIGHT n rn GRAVEL TILL SHELL SAND SAND- SILT- SILT CLAY BOTTOM SEDIMENTS Figure 48 Density and biomass of Zoantharia in relation to bottom sediments. 100 _ 0 0 NUMBER A • • WEIGHT s M 1 £ 80 i o tf> o z m UJ 2 u. ^o: 60 in uj ; x A \ 12 z o: hi h- >- U_ LJ O 5 I UJ O 2 0C UJ UJ w uj (r * (E 2 < 40 2 3 - 8 K§ 3 O ; Vj\ \ uj a z in * w oe rr UJ UJ a. 20 0. 4 y \ ^y^^^^\. ^ lOv i \ \ a. a u ' / \ ^ UJ OL a. ' / \ x^ 2 0 4.0 ' / V. \ '/ ^v \ d \ 1 ' i ' ^*» ' t ' i ' o 0 * 0.01 0.5 10 15 2 0 SO 5 0 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%) con- tent 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 die small size of die majority of specimens obtained (Table 3). Their greatest numeri- cal density and biomass occurred on the continental shelf and along the upper portions of the continental slope. Members of this group are carnivores which charac- teristically 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, represent- ing only part of an obviously larger animal. These soft- bodied, 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 con- tained over 100 individuals. They occurred in 405 samples (38% of total). Their density averaged 8.2/m2, and their biomass averaged 0.71 g/m- (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 49/ m2) extended along southern Georges Bank, across Great South Channel, and westward to the vicinity of Rhode Island. They were absent in a few deep water sections of the Gulf of Maine and on the conti- nental rise southeast of Long Island, New York. Density (average number of specimens) was greatest (23 indi- viduals/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 g/m2). Both slope areas had very small 86 NOAA Technical Report NMFS 140 EXPLANATION □ <0. 1-0.3 GRAMS PER SQUARE METER TURBELLARIA Figure 5 1 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 moder- ate 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 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 nu- merical density, but the range in values was much more limited. Largest average biom- ass 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 occur- rence was typical of deepwater classes. Relation to Sediment*, Although nemertines were present in .ill 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/ in2) was found in till. Biomass of nem- ertines 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/mL or less. An unusually small quan- tity (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). 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 interme- diate annual temperature range of 12° to 15.9°C. 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 de- creased to 2.9/m2. Where the range nar- rowed from 11.9° to 0°C, the values de- creased from 9.0 to 2.4/m2. OF SPECIMENS METER OF BOTTOM o - z o I* cc u o o z cc — UJ h- 1- o z □ NUMBER ■ WEIGHT NUMBER SQUARE o ■ fb o o o o WET WE SQUARE ft * 0 n J m 1 Cc UJ 0. 0 NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 52 Densit) and biomass of Turbellaria in each of the six geo graphic areas. s o 5* — 0 O NUMBER • • WEIGHT z o 1&25 a* \ or it S>£ 20 °5 15 \ \ \ ■" t- •-5 003I2 UMBER SQUARE o \ UJ cc 002 *§ I- O \^ \ uj en Z cc 0.5 — N\ ^^m. 001*^ UJ W*"^^CK\ a^^ UJ 0 i i h< rN-^f^ ^t*^, i f i g ) 10 25 50 100 "200 500 1000 'SOOO 4000 WATER DEPTH IN METERS Figure 53 Density an d biomass of Turbellaria in relation to water depth. z 1 1 NUMBER z o 1- o M WEIGHT w £ z m Ul 11 u «> O Z fe O ° 30 m u; ? CC Ul 1- J- OZ r ui o Z CC UJ w cr io - 0 02 |g £ z 3 t- = zw 05 ■ ■ ■ _ ooi j en rr J cc °- o ■ ri r-i GRAVEL TILL SHELL SAND SAN0- SILT- SILT CLAY BOTTOM SE0IMENTS Figure 54 Density and biomass of Turbellaria in relation to bottom s ediments. 88 NOAA Technical Report NMFS 140 012 5 o t- z o o o NUMBER - u> o z a> UJ 2 u. • • WEIGHT * u. u o NUMBER OF SPE PER SQUARE METER o - o b. o / \ / " 008 0 04 0 ? K UJ 1- t- I UJ UJ uj * K 4 l- z> UJ O * <° K UJ 0. a^~- -^ \ i ¥ ¥ i 0-^.9 4-79 8-119 12-159 16-199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 55 Density and biomass of Turbellaria in relation to the annual range of bottom water temperature. O O NUMBER • • WEIGHT If 0.8 l\ 0.20 X 1 l\ s o 1 I o 1- t- / / \ 1- co O ^ / / \ 2 o Z CD X £ SJU.0.6 - 0.15 < K U. 3> y » o UJ o S I UJ (9 2 OC UJ '' \ // I UJ UJ UJ K S < 0 4 S 3 0.10 * | 1- 3 3 o Z (0 UJ OC UJ 0. a. / \ / / \ 0.2 0.05 / \ ' / \ / ^ ' \ ' ^ / / \ 0 i i a i a ~T "~^^ i A i 1 fi ' 9 i 0 ■ 0.01 0.5 1-0 13 2 0 3.0 5.0 100 PERCENT 0R6ANIC CARBON Figure 56 Densitv and biomass of Turbellaria in relation to sediment organic c arbon. Composition and Distribution of Macrobenthic Invertebrate Fauna 89 "E" \ NEW .-/\ \ / JERSEY ', YORK/ \ \ NEW / \ \ HAMPSHIRE -' _CONNECTICU / ' — \ — ' ~7^\ — ' ^ — ' — ' NEW \ NEW .- \ \ JERSEY '.YORK/ '- \ NEW / \ \ hampshire ^Connecticut" NEMERTEA 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 gen- eral increase in biomass (from 0.22 to 1.38 g/rrr) 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 rela- tion 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 tempera- ture range was narrow or broad, the frequency of oc- currence was relatively low (Table 25). Relation to Sediment Organic Carbon The average density of nemertines exhibited a rather 90 NOAA Technical Report NMFS 140 1 1 NUMBER n ■i WEIGHT OF SPECIMENS METER OF BOTTOM — 00 - " GHT IN GRAMS METER OF BOTTOM NUMBER SQUARE o o CD WET WE SQUARE or UJ °- 5 1 J - 0. Ul 04 "" 0 1 ■ 1 0 NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 58 Density and biomass of Nemertea in each of the six geographic areas. m-~ o o NUMBER 2 • ♦ WEIGHT 25 _ S 5 o O r- r- 5°° 20 S u. - 5 CO gu. G° / \ o o uj or ft 5 1.5 -, a. ^ UJ 05 t- 15 //\\ I- u. B r- UJ o 5 X \. I 2 rr UJ / V o — UJ M8E QUAR 5 / '\ ,0 -g t- o or UJ 0- 5 / \ uj ui * cc 05 £ •^^^°^Sv X^^^-O-.^Ji 0 " 10 25 50 100 200 500 1000 2000 4000 " 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 ( 1 1/m2) in areas with extremely little or no organic carbon in the sediment and declined steadily (from 11 to 2 nrl as organic carbon content increased. None were found where carbon content exceeded 39? . The relation of nemertine biomass to organic carbon was distinctly bimodal (Tables 28, 29; Fig. 62). Largest Composition and Distribution of Macrobenthic Invertebrate Fauna 91 biomass (0.44 g/irr) 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 g/ m2) at levels between 0.01 and 1%. Intermedi- ate 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 moder- ately high (40%). Aschelminthes Nematoda — Free-living nematodes, members of the phvlum Aschelminthes, are one of the most numerous animal groups inhabiting the bottom sediments of the northeastern coast of the United States. Previous studies con- ducted by Wieser (1960) in shallow coastal habitats off southeastern Massachusetts re- vealed nematode densities of nearly 800.000 individuals/m2. Farther offshore, on the southeastern coast of Massachusetts at a depth of 58 m, Wigley and Mclntyre (1964) encoun- tered nematodes in densities of nearly 1 mil- lion individuals/m , and this measure did not include the young stages of numerous spe- cies. The vast majority of free-living nema- todes 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 speci- mens was represented. In addition to their great numerical density, this group of ani- mals is also exceedingly profuse in species composition. Three types of feeding 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, par- ticularly diatoms; still others are carnivorous, ingesting rotifers, tardigrades, small annelids, other nematodes, and bryozoans. Because of the incomplete representation group, the quantitative distribution discussed refers only to the exceptionally large species. 00 .1 5 3 3 O Z CO | — 1 NUMBER M WEIGHT — 6 so — 25 — 20 — — 5 15 — 10 — — 3 — 2 5 — P 1 n. Mi * cr < h- 3 GRAVEL TILL SMELL SAND SAND SILT- SILT CLAY BOTTOM SEDIMENTS Figure 60 Density and biomass <>1 Nemertea in relation to bottom sediments. cr lu u a 00 < Z 3 3 O z en O O NUMBER • • WEIGHT 0-39 3-79 e-119 12-159 16 199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 61 Density and biomass of Nemertea in relation to the annual range of bottom water temperature. of this We found nematodes in 98 samples (9% of total); herein their density averaged 2.8/m2, and the biomass aver- aged 0.01 g/irr (Table 5). 92 NOAA Technical Report NMFS 140 O o NUMBER • • WEIGHT 10 0.8 X o 1- l- V) O Z IB UJ 3 U. 5° U>UJ U. UJ 15 :A/\ - p IGHT IN GRAMS METER OF BOTTOM 3° 10 V \ o WET WE SQUARE IE UJ 0. " Xx- \ re banks and basins, and on the continental slope and rise. Thev were commonly absent in samples from a number of areas: parts of the Nova 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 sub- strates 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 indivi duals/ m2 (Tables 6, 7; Fig. 64). Their average biomass was very low, 0.01 g/m'- or less (Tables 8, 9; Fig. 64). The percentage of samples containing nematodes was low (5 to 9'i ) 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/m-. Average biomass values were uniformly low (0.01 g/m2 or less) at all depths (Tables 13, 14; Fig. 65). Frequency of occurrence 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/m ) occurred in gravel substrates, whereas Composition and Distribution of Macrobenthic Invertebrate Fauna 93 NEW \ NEW .- \ \ JERSEY ', YORK/ N \ NEW / \ \ HAMPSHIRE y v6fi»TS GRAMS PER SQUARE METER NEMATODA Figure 63 Geographic distribution of Nematoda: A — number of specimens per square meter of bottom; B — biomass in grams per square meter of bottom. all other bottom tvpes yielded relatively moderate den- sities (1 to 3 individuals/rrr). Average biomass was small (0.01 g/m2orless) in all sediment types (Tables 18, 19; Fig. 66). The frequency of occurrence of nematodes was mod- erate (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, 20°-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 (0°-11.9°C) and declined drastically (0.2 and 0.7/m2) in the broader ranges. 94 NOAA Technical Report NMFS 140 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 nema- todes in samples in the various tempera- ture ranges diminished from a high of 17%, where the temperature range was narrow, to zero where the temperature range was 20°Cor more (Table 25). Relation to Sediment Organic Carbon Nematodes occurred onlv 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 car- bon levels (1.5-1.99%). Biomass was very low (<0.()1 to only 0.02 g/m2) in all levels of sediment organic car- bon in which they were found (Tables 28, 29; Fig. 68). Frequency of occurrence in samples in the four carbon content classes ranged from 7 to 17% (Table 30). Annelida Polvchaete worms formed a major compo- nent of the benthic fauna in terms of biom- ass and numbers of individuals. They were present throughout the study area and made up 28% of the total number of indi- vidual animals and nearly 10% of the total biomass (Table 3). Taxonomically diverse as well as abun- dant, 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 speci- mens 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 of weight is Aphrodita hastata, 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 5 1 O □ NUMBER O !JO 4 Z CO - ■I WEI6HT 2 u. cr u. 5° £* 3 n z tr if) uj 1- u_ UJ L . ooi £i: O 5 O 5 ■ UJ uj UJ or ■ 1 * cc CO ti ■ ■ « NUM PER SOU o — - i o WET PER SQU NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 64 Density and biomass of Nematoda in each of the six geographic areas. s 0 O NUMBER 3 o •— — • WEIGHT h- 2° 2 CO z, ff> < 0° 8 oO ^ OL ZuJ SjE <\ 1- u. a \ 5Es oS \ Ouj cc £ s§ 4 - \ r~~\ UJ cc 002 $< S 2 => CO " . ^ / ■ A- n uj en 0.01 *„. z cc ^--V/ v v^c °< i i i i i i i i i ) io 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 65 Densi tv and biomass ot Nematoda in relation to water depth. O 1 1 NUMBER z o CIMENS OF BOT o ■ WEIGHT I" or U. o o ftuj a u. UJ OS 6 CC UJ UJ £t §3 * 3° 2(0 2 Or UJ °- 0 - 1 J ill - o o o o WET WEIGHT IN PER SQUARE METER GRAVEL TILL SHELL SAND SAND- SILT- SILT CLAY BOTTOM SEDIMENTS Figure 66 Density aii<\ biom iss of Nematoda in relation to bottom sediments. Composition and Distribution of Macrobenthic Invertebrate Fauna 95 variety of morphologically different forms abound. All the different feeding types — carnivores, suspension feeders, and selective and nonselective deposit feeders — were rep- resented in our samples. Mam1 representatives from the two major life modes, the errant (or free living) and the sedentary (tubicolous) polychaetes were collected. Coloration of polychaete annelids is extremely di- verse, from nearly translucent and white to a dark brown. Predominant hues are light beige, tan, dark brown, and various shades of olive and red. Some annelids dis- played a cuticular irridescence that greatly enhanced their appearance. Some forms possessed variegated pat- X £ ■ t- to o z m UJ 2 u. 4 (OUJ u. UJ J o 2 IE UJ UJ CC m < 2 2 o 3° Z to IT UJ ' a 0 Densi bottoi — o TO WET WEIGHT IN GRAMS £, PER SQUARE METER OF BOTTOM O 0 NUMBER • • WEIGHT A - A / A 0-39 4-79 8-119 12-159 16199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 67 v and biomass of Nematoda in relation to the annual i n water temperature. O 0 NUMBER 3.S 3.0 A . / \ M Z Z o o 1- U |6 IlL / \ / \ DC li- 5° es O a?- z.o 1 \ 1 \ I \ i \ l \ i \ x 1 \ Z K m uj U. UJ OS UJ ii o: uj 1.9 UJ (T - 0.075 £ UJ i| i < t- 3 1.0 K - £8 0.000 * _ K it! UJ a. i \ 0.5 0.025 0 i -r1 * ~1 — "~— J i^t- i v$. i « i » i 0 001 0.S 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. 96 NOAA Technical Report NMFS 140 NEW \ NEW s \ \ JERSEY ', YORK/ '- \ NEW / \ \ HAMPSHIRE •' ncuT\ \. j 00BTL*Ste~3 ^aSSaCHUSETTSV;^^^^::1'- im^? ilfev/^likk ,-'r i > tt^"'V«*^ EXPLANATION X Nl □ --O NUMBER — • WEIGHT 2 O t- S m o o 15 I S * < -3 10 S" 0 10 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 71 Density and biomass of Annelida in relation to water depth. 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% of samples in gravel and silt-clay. Although annelids were present in 94% of the samples from sand sediments, this cat- egory 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 tem- perature was broadest, and densities generally declined as the temperature range decreased. Where the tem- perature range was greater than 20°C, average density of annelids was 1,698 individuals/m2. At the other ex- treme, 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/m . 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 tem- perature ranges were extremely high and low. Biomass of annelids in relation to range of water tem- perature 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 Composition and Distribution of Macrobenthic Invertebrate Fauna 99 5 O CO o 2 IB 5 U. U_ UJ O 2 or uj UJ CE CD < 2 ID => O z to I — 1 NUMBER ■ WEIGHT 300 — 200- — 24 16 .0 z o (- « t; ? ct UJ (- t- X UJ o S UJ ui * or < t- 3 UJ o * GRAVEL TILL SHELL SAND SAND SILT- SILT CLAY 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/irr) 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 es- sentially bimodal relationship to the amount of organic carbon in the sediments (Fig. 74). Greatest density (504/m2) occurred at low organic carbon levels (be- tween 0.01 and 0.49%); another peak in density (407/ irr) 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 sedi- ment 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, rang- ing from 80 to 100% in all classes except one, the 3-5% class, in which only 50%i of the samples contained mem- bers of this group (Table 30). 100 NOAA Technical Report NMFS 140 Pogonophora 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 taxo- nomic 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 col- lections 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 frequently of a green- ish-yellow to brownish-green hue. The majority of them exhibited alternating light and dark rings or bands. Embryos were not uncommon in the tubes from our samples. Preserved speci- mens 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 repre- sented in our collections by species from the genera Siboglinum, in the former, and Diplo- branchia 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 com- mon were Siboglinum holmei Southward, S. angustum Southward and Brattegard, and Crassibrachia sandersi Southward. Siboglinum holmei was distinctive for its oc- currence 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 1 171, 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 aver- aged less than 0.01 g/m2 (Table 5). Geographic Distribution Pogonophores were widely distributed along the conti- nental slope and continental rise from New Jersey north- ward to Nova Scotia but rarely occurred on the conti- nental shelf (Fig. 75). In coastal waters they were en- countered 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, 1800 O O NUMBER r • • WEIGHT 1600 - / / 1400 / / / / // / X o t- 1200 t- z m - 40 2 O s u. ft 'ooo co£ - or u. CD o Z EC — UJ *- h u_ UJ x w O 2 /\ 30 o Z a u 800 - j i UJ UJ uj or 7 i * tr m UJ o 0.01 0.5 10 IS 2 0 SO PERCENT ORGANIC CARBON Figure 74 Density and biomass of Annelida in relation to sediment organic carbon. Average density was highest (6.8 individuals/in-) between 500 and 1,000 m. Density was slightly lower (3.2 to 3.5/ m2) in deeper water (1.000 to 2,870 m) but substantially lower in depths less than 500 m (Table 11). Biomass ranged from 0 to 0.03 g/nr, and the trend in relation to water depth was comparable to that de- scribed for density (Table 13). Pogonophores were present in approximately halt 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/ m2) in sand sediments, intermediate in sand-silt, and highest (1.9/nr) in silt-clay (Table 16). Their biomass was very low (<0.01 g/nr) in sand, and only slightly higher in the other fine-grade sediments (Table 18). The occurrence of pogonophores in the samples cor- related very closely with average density. They were exceedingly sparse (occurring in <1 %) 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 of less 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/nr (Tables 21, 23). Even in areas where the temperature range was between 4° and 11.9°C they were exceedingly sparse in both den- sity (0.1 and 0.7/ nr) and biomass (<0.01 g/nr). The areas of small temperature variation (see Fig. 11) correspond to the deepwater regions on the conti- 102 NOAA Technical Report NMFS 140 NEW \ NEW ,./S\ \ JERSEY '.YORK/ \ \ HAMPSHIRE •' CONNECTICUT'* '■ ;' POBIL ^^MASSACHUSETTS' / \ BOSTON 'R i. > INDIVIDUALS PER SQUARE METE NEW \ NEW .- \ \ JERSEY ', YORK/ \ \ NEW / ~) occurred in the Southern New England Slope area; the smallest (0.37 g/m~) was in the Gulf of Maine. Individual sipunculids averaged approximately 0.1 g in weight in all areas except Georges Slope, where their I o o 0 NUMBER o w o • • WEIGHT s ° z m UJ *■ u. 2 u. <° o * a. 0- Pi UJ U)UJ \- h- NUMBER OF SQUARE ME \ \ \ o o o o WET WEIG SQUARE M cr UJ 0. ^~» .L ■2 UJ a. 0 1 1 Y~ — >--Jr, JO JT> 0-39 4-79 8-119 12-159 16199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 79 Density and biomass of Pogonophora in relation to the annual range of bottom water temperature. average weight was nearly forty times greater, 4 g per individual. Frequency of occurrence of sipunculids was moder- ate to moderately low. They were present in 13 to 42% of the samples (Table 10). Highest frequency of occur- rence 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 2/mL. The relation between biomass of sipun- culids and water depth was substantially dif- ferent from that of numerical density. Bio- mass was much greater (averaging about 1 to 4 g/m2) on the middle and lower portions of the continental slope (500 to 2,000 m) than it was on the continental shelf and continen- tal rise (Table 13, Fig. 83). Also, sipunculids formed 10 to 26% of the total benthic biom- ass 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 3.0 O O NUMBER S o 1- »- (ft O Z CD •— — • WEIGHT / \ 2 O uj^ 2.0 6° or u. o o z or — UJ r-|- U. UJ OS / v 19 2 K UJ C0< 10 / \ UJ UJ 0.050 . * = o / \ r- 2 ^ S5 Z V) / \ / \ * " K UJ or 0.025 £ a. / \ 0 |a — Li —~"' 1 1 T & — ' ^1 fi ' 8 ' 0 ' 0 0.01 0.5 1.0 1.5 20 30 50 10.0 PERCENT ORGANIC CARBON Figure 80 Density and biomass of Pogonophora in relation to sediment organic carbon. Composition and Distribution of Macrobenthic Invertebrate Fauna 105 NEW \ NEW ,-^\ \ / JERSEY ; YORK/ \ \ NEW / \ \ hampshire :ticut^ \JrfASSACHUSEr f\ BOSTON 'R I > V INDIVIDUALS PER SQUARE METER NEW " \ NEW JERSEY J YORK/ S\ ' \ \ NEW HAMPSHI SETTSV. y / • 'ViHI.tN'*.. yr MAINE 7 t / ( \ dnnecticut" \ \>IASSACHL / \ BOSTON Vi.) t :/ ^P* ^ / { "V / v v^ i \ ' ^"'O / ( f NOVA / ^*f SCOTIA EXPLANATION □ <0.l-9.9 M 10.0-28.5 o c ^ o ^ GRAMS PER SQUARE METER \ / v j , B / \ \ 39° «o° 41* 42° SIPUNCULIDA 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 7.1/irr. Sand con- tained the highest density. Biomass of sipunculids in relation to various sedi- ment types was considerably more varied than density of sipunculids (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 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 sand- silt; lowest values occurred in shell and silt- clay sediments. Relation to Water Temperature Sipunculid density, biomass, and frequency of occurrence generally tended to decrease as the range in temperature broadened (Fig. 85). None were found where the tem- perature range exceeded 20°C. Density averaged about 5 to 9 individu- als/mJ 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 de- creased at a rather uniform rate as the temperature range increased (Table 22). Biomass of sipunculids was low or mod- erately low, and the changes in quantity in relation to temperature range followed pre- cisely the same pattern as those in relation to density (Tables 23. 24). Biomass aver- aged 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 percentage bio- mass composed of sipunculids also decreased as the temperature range expanded. Frequency of sipunculids in the samples ranged from 0 to 29% (Table 25). Thev 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 occur- rence of sipunculids dropped substantially and thev were absent in samples from ar- eas where the temperature range was more than 20°C. Relation to Sediment Organic Carbon Sipunculids showed a decided preference for sediments with moderately low to low amounts of sediment organic carbon (Fig. Ni>). Both measures of abundance were greatest in the two (lasses between 0.01 s Z O 12 — 1 1 NUMBER _ ■1 WEIGHT o 1 ° z m UJ - JS> 8 z or u, UJ UJ h- K- u. UJ OS i-o a s or uj uj u uj or , n * or ■ S 3 2° L — 05 Z L ■ I * 0) or ■ 1 I or UJ ■ ■ ■ ■ ■ 1 ■ CL NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 82 Density and biomass of Sipunculida in each of the six geographic areas. 0---0 NUMBER » • WEIGHT § o — 4 1- «5 5° 1* 5 u. or u. - O o o uj or 2 X a uj i in h- H l- UJ o 2 8 - ^-\\ \ o 2 - UJ uj or §5 G - i \ S < uj° => Lo * / \ \ uj tn z or 4 — ^ M \ - , -4 1 tr Id / -^-" •l^^ I \ \ UJ a. 2 — d 1 1 1 1 1 1 1 1 J 10 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 83 Density and biomass oi Sipunculida in relation to water depth. s Z o 1 — 1 NUMBER o w o ■ WEIGHT »& z m 10 S oj UJ < Z u. or u. S° 8 CO o 08 _ 0. K z or »UI UJ 1- t- u. UJ 6 O I 1 " j- - 06 I" to S or uj UJ uj K^ 4 1 1 - 04 *5 s = ■ ■ 1- 3 2° ■ U ■ UJ O 2 10 g 1 _ 02 *M or UJ 0. 0 1 1 1 jL 1 1 or UJ a. GRAVEL TILL SHELL SAND SAND- SILT- " SILT CLAY BOTTOM SEDIMENTS Figure 84 Densit \ and biomass of Sipunculida in relation to bottom s< diments. Composition and Distribution of Macrobenthic Invertebrate Fauna 107 i p 10 CO o z o O 5 uj a CD < 3 O --O NUMBER — • WEIGHT L L_ _1_ J... S O 1- -5 |» 10 §£ ? or 0 8 w h- I- i sy o 5 06 uj u * or 1— OTO 1 A\ x 5 Z CD < " 111 o° 1 / / y i j \ x \\ \\ K u. «o ? cr Ui toy OS I- 1- i i \\ 0.8 I UJ (9 S oe ui UJ cc \\ \\ UJ M < h i 1- = 11 i / \\ *" oe A / \\ CC UJ S 20 * ■ ' V \\ Y \ \ \ \ i f — ^ « i >i § i 0.4 0. 0.01 0.5 1.0 15 2 0 3.0 5 0 10.0 PERCENT ORGANIC CARBON Figure 86 0 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 of sipunculids ranged from 0.1 to slightly more than 7 individuals/in2 (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 of occurrence of sipunculids in the samples paralleled the trends established by density and bio- mass. 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 typi- cally burrow into sand and mud. Others are found among rocks and on coral reefs; one southern species uses the tests of sand dollars as a habitat. Their size varies greatly, ranging from tiny 3-mm males to large females over 300 mm in length. Speci- mens in our samples were in the 2 to 8 cm 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 of less than 1.000m. Color of specimens 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). Den- sity averaged 0.1 /m2 and biomass 0.30 g/m2 (Table 5). Geographic 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 onlv 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 aver- age 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). Bathymetric Distribution Echiurids were found at depths ranging from 20 to 3,975 m. Their numerical abundance was greatest, al- beit low (0.5 to 0.6/m2), in water depths greater than 1,000 m and even lower (0.02-0.3/m2) at shelf and inshore depths (Table 1 1, 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 of this 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 0.3/ 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 prefer- ences of echiurids, it is not surprising to find that they also had restricted temperature preferences. They oc- cupied only three temperature ranges; areas with the most stable annual range (0-3. 9°C) contained the low- est density (0.1/m2) but the highest biomass (0.9 g/m2); areas with an annual range of 16-19. 9°C had the high- est 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). Fre- quency 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 1 0 NEW s \ NE'W y^"\ \ JERSEY '.YORK/ \ ....«»__ / INDIVIDUALS PER SQUARE METER \ NEW .- \ \ GRAMS PER SQUARE METER ECHIURA Figure 87 Geographic distribution of Echiura: A — number of specimens per square meter of bot- tom; 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 spe- cies belonging to six genera have been reported (Barnes, 1974). Priapulids contributed <0.1% of the total num- ber of specimens 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 poly- chaete worms. They are typically between 4 to 8 cm. in length. Our specimens were considerably smaller, in the 1 to 2 cm size range. Color of specimens ranges from whitish to flesh colored with some yellow on ap- pendages; some of our specimens were somewhat red- dish brown. 110 NOAA Technical Report NMFS 140 Only 4 (0.4% of total) of our samples yielded specimens (10 individuals weighing a total of 4.60 g) whose mean density was <0.1/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). Mollusca The phylum Mollusca contributes significantly to both measures of abundance (numerical density and biomass) in the New England re- gion as it does in the Middle Atlantic Bight region (Wigley and Theroux, 1981). The mol- luscan fauna comprises five classes: Poly- placophora. Gastropoda, Bivalvia, Scaphopoda, and Cephalopoda. Each of these classes will be discussed separatelv below. For the sake of con- tinuity in the phylogenetic ordering of figures, those figures dealing with phvlum Mollusca (Figs. 99-104, inclusive) are included here, but a de- tailed discussion of the phylum as a whole will be presented in the section "Dominant Com- ponents of the Macrobenthos" below. Polyplacophora — This class of mollusks is also called chitons, which are bilaterally symmetri- cal, have eight overlapping dorsal plates, and a broad, flat, ventral foot. Thev 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/nr. and biomass averaged 0.14 g/m- (Table 5). Geographic Distribution Polyplacophorans, although rather sparse, oc- curred in small to moderately large geographi- NUMBER OF SPECIMENS PER SQUARE METER OF BOTTOM o o o o 1 — 1 NUMBED SB WEIGHT n n 1 WET WEIGHT IN GRAMS PER SQUARE METER OF BOTTOM Densit] areas. NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 88 and biomass of Echiura in each of the six geographic 5 O is 2 O O NUMBER - WEIGHT \ A oj—rir I \ ' 25 50 100 200 500 1000 2000 4O00 WATER DEPTH IN METERS Figure 89 Density and biomass of Echiura in relation to water depth. z O 04 - □ NUMBER 3 o OF SPECIMENS METER OF BOT o o to u. Wt WEIGHT . \ o o OV CD GHT IN GRAMS METER OF SOT NUMBER SQUARE o - o o WET WE SQUARE UJ a. 0 1 l 0 GRAVEL TILL SHELL SAND SAND- SILT- SILT CLAY BOTTOM SEDIMENTS Figure 90 Density and biomass of Echiura in relation to bottom st diments. Composition and Distribution of Macrobenthic Invertebrate Fauna 111 z O O NUMBER s • • WEIGHT — 1.0 K t- w o z m . I ° < a I u. \ - 08 gu. OF SPE METER o \ o / X o 6HT IN METER NUMBER SQUARE o - A x 7Xx UJ uj < UJ o 02 K UJ a UJ a y_ / , \ 0 0-39 4-79 B-II.9 12-159 16199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 91 Density and biomass of Echiura in relation to the annual ange of bottom water temperature. O O NUMBER 1.6 Z r^\ z O 1- J \ 1.2 o K I \ 0.3 / '^ \ < t- 3 UJ O * «o ec CC UJ a. 0.2 0.1 // \ \ 0.4 UJ a. 1 "\ \ u i # i i i v4 i ♦ i • i * i 0 0 0.01 0.5 1.0 1.5 2.0 3.0 3.0 10.0 PERCENT ORGANIC CARBON Figure 92 Density and biomass of Echiura in relation to sediment organic carbon. 112 NOAA Technical Report NMFS 140 NEW \ NEW s\ \ GRAMS PER SQUARE METER PRIAPULIDA Figure 93 Geographic distribution of Priapulida: A — number of specimens per square bottom; B — biomass in grams per square meter of bottom. meter of 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 9/m2 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 1 o I o 1 — 1 NUMBER OF SPECIMENS METER OF 601 o o ■ WEIGHT 006 004 z ° tt u. 43 O ? UJ o * w cr S 0 l l cr UJ NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW EN6LANC SHELF SLOPE GEOGRAPHIC AREA Figure 94 Density and biomass of Priapulida in each of the six geographic: areas. Z o 0---0 NUMBER s o h • • WEIGHT s° CO 0 5 m LU < Su. 11 fe o 32 ~ uj cr Z IE 0- LU — UJ io z CC 0 1 // Yyj - cr LU // \ UJ a. / V 0 10 ~ 25 50 "lOO "200 " 500 "OOO 2000 4000 WATER DEPTH IN METERS Figure 95 Density and biomass of Priapulida in relation to water depth. £ 2 O 1 — 1 NUMBER t- t- ■ WEIGHT cn H « o S2 o 2m 5 "> 2 u. Ct U_ 6° £* Z EC £ w U 1- 1- U. UJ I LU O 3 o z cr uj UJ uj uj cr S IT 13 ■ 0.02 _« IS 0I ■ £8 0 01 * EC LU CL 0 rb rl bl 0 E GRAVEL TILL SHELL SAND SAND- SILT- SILT CLAY BOTTOM SEDIMENTS Figure 96 Densit i and biomass of Priapulida in relation to bottom sediments. 114 NOAA Technical Report NMFS 140 X s o o to o z m O— -o NUMBER • • WEIGHT «> fc; s ° 2 u. £ u. 5° ° o *£ 2 C V) w UJ 1- 1- u. !±> O 2 o S (£ UJ UJ u uj a. oi 5 3 \ 0.01 * 5 < \. UJ o ^ \ * B UJ a 0 1 N= f> f> f jp 0 UJ a. 0-39 4-79 8-119 12-159 16 199 20-23.9 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 97 Density and biomass of Priapulida in relation to the annual range of bottom water temperature. 0 0 NUMBER • • WEIGHT ; j f\ : 0.02 A 0.01 / \ z z / \ 0.008 g H / \ t l- 1 o 2 <*> «> o zS / \ < UJ £ "■ Z u. / \ 0 o 5° SI" \ \ UJ c/i uj »- 1- U.L, °°« \ \ o 006 r uj o z oz / \ UJ u < 1 R\ >- 3 Z D UJ O Z /\\ * • BE ■X £ 0.04 - \ 1 \\ 1 \\ 1 A U 0.004 0. - / »\ «/ \\ 0.02 " / \\ 1 ' \ 0 002 ~ / ^ \ - i" i > i a i g i § i § i o 0 001 0.5 10 15 20 30 50 100 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 \ NEW ." \ \ ,■ JERSEY '.YORK/ V \ NEW / • ^ \ \ HAMPSHIRE -' CONNECTICUT "« *» i »0»Tl«ND' \^»ASSACHUSETT! X\ eosTon f-'R I > V MOLLUSCA Figure 99 Geographic distribution of Mollusca: A — number of specimens per square bottom; B — biomass in grains per square meter of bottom. meter of or less. On Georges Bank, Georges Slope, and the South- ern New England Slope their density averaged 0.1 to 0.6/m , and their weight averaged 0.01 g/m or less. Frequency of occurrence was relatively high ( 1 to 24%) considering the small quantities that were present (Tahle 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 140 300 — to o z m UJ U. UJ OZ a; UJ UJ ct CO < Z => 2° z tn ae UJ a. 200 — ISO — 100 I — I NUMBER ■ WEIGHT 160 120 100 -80 60 X o " J; < » <* u. °o Id (- ►- X UJ (9 Z UJ ui * c < I- 3 ui o * CO a UJ a. 40 20 NOV* SULFOF GEORGES SOUTHERN 8C0R8ES SOUTHERN SCOTI* MAINE BANK NEW ENOIANO SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 100 Densitv and biomass of Mollusca in each of the six geographic areas. m ranged from 0.4 to 4.0/mL. Below 300 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 g/ m2) at depths less than 500 m than in the deeper water where their biomass averaged 0.01 g/irr or less (Table 13; Fig. 107). Frequency of occurrence was low (1 to 18%) and no clear relations with depth were evident. The trend indi- cated 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 wa- ter (>500 m) than on the continental shelf and upper portion of the continental slope. Relation to Sediments Polyplacophorans occurred in all sediments except shell (Table 16; Fig. 108). Highest density (3.8/m2) was en- countered 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 o 0 NUMBER » • WEIGHT 250 S 600 2 O UJ ^ 3 0 \ si * or Z OT 1 \ ' I < £ 400 0. / \ p' > / \ / / UJ 0 * w / \ / / / \ / / K UJ 60 a / \ / / 200 / \ 7 / _/. ■ — '°r~~~~ *----^ / 0 i 1 1 1 1 1 1 1 9 0.01 0.5 1.0 15 2 0 3.0 5.0 10.0 0 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 of occurrence in samples exhibited a trend similar to that of biomass. Incidence was relatively low (ranging from 2 to 12%) and diminished with increas- ing carbon content (Table 30). Gastropoda — Gastropods formed a moderately com- mon component of the New England benthos. They were distributed throughout most of the studv 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 Retusa and Alvania (approximately 2 mm in length ) to large specimens of Neptunea, Colus, Busycon, and Buccinum (ranging up to 13 cm or more). The majority of specimens were between 2 and 30 mm. Shelled gastropods were predominant in our collec- tions, 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 of different taxonomic groups obtain their nourishment by a variety of methods. Feeding types known to be represented in our collections were herbi- vores, 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 8/m2; biomass averaged 2.2 g/m2 (Table 5). 120 NOAA Technical Report NMFS 140 EXPLANATION □ 1-9 10- 99 100-124 INDIVIDUALS PER SQUARE METER NEW \ N7» ,.y'\ \ JERSEY ', YORK/ \ \ NC" / \ \ HAMPSHIRE .* CONNECTlCU GRAMS PER SQUARE METER POLYPLACOPHORA Figure 105 Geographic distribution of Polyplacophora: A — number of specimens per square meter of bottom; B — biomass in grams per square meter of bottom. Geographic Distribution Gastropods were distributed over nearly the entire re- gion (Fig. 111). Moderately low densities (1 to 49 indi- viduals/m2) were widespread over a large part of the study area. Medium and high densities generally oc- curred in limited areas inshore and nearshore. In most areas where gastropods were found, their average biomass was less than 1 g/m . Moderately high biomasses (1 to 25 g/m2) occurred over rather large areas both inshore and offshore, whereas large biomasses (25 to 133 g/m'-') 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 1 1 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 Frequency of occurrence of gastropods was moderately high in all geographic areas. They were present in 35 to 58% of the 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 of small-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 abun- dant (64/m2) in shallow water (0 to 24 in) than at other depths. Their average density (Table 11: Fig. 113) generally diminished with increas- ing water depth, except for a slight reversal oi 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 1/m2). 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 continen- tal shelf and upper slope (Table 15). At mid- and lower-slope depths occurrence was high (74 to 77%). On the continental rise their rate of occurrence diminished to only 30%. Relation to Sediments Although there was no consistent trend in den- sity of gastropods in relation to sediment par- ticle 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 gas- tropods were predators on the bivalves whose shells formed the substrate. Densities were also high (40/m2) on gravel bottoms. In all other sediment types they occurred in only moderate densities (9 to 22 individuals/m2). The unex- pectedly low density of 9/m2 in till substrates indicates that till is more closely allied to silt- clay as a gastropod habitat than it is to gravel. Just the reverse is true for other molluscan groups. o a a uj UJ (T §5 r~] NUMBER ■ WEIGHT 1- D uj O Xt_ GEORGES SOUTHERN GEORGES SOUTHERN BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 106 Density and biomass of Polyplacophora in each of the six geographic areas. o o NUMBER • • WEIGHT a o H06 ? ' uj or — 02 5 0 10 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 107 Density and biomass of Polyplacophora in relation to water depth. 1 o 1 — 1 NUMBER Z o IB WEIGHT »|: CO O ig J6 < "" r,° O O *£ * - z or in £ u. £ O 2 _ - 01 £uj 31 cr id uj cr "b - UJ uj *5 m < 2 S 3 02 ii => a z w - L 0. l 1 1 0. 0 GRAVEL TILL SHELL SAND SAND SILT- SILT CLAY BOTTOM SEDIMENTS Figure 108 Density and bioma ss of Polyplacophora in relation to i )ottom sediments. 122 NOAA Technical Report NMFS 140 o 0 NUMBER s • • WEIGHT 4 2 H * o lO o Z CD UJ A 12 2 "■ < 5° 8 / \ 10 11 U. «> o 10 uj Am ? or UJ u. UJ 6 OS / \\ '. tj 1- h- I UJ CC UJ UJ cc 13 « D.6 UJ ^ S cc < 1 \ 04 UJ o £ 2 in a ui UJ cc «< 2 0 ig \massachuse / \ BOSTO* 'r. i. •> EXPLANATION LI o S a> z m < UJ IT u. Z u O O F, O B 5° — 3 z* ^F. — UJ U_ UJ O 2 - UJ Or — UJ UJ CD < P - - 1- => 1 UJ O 2 |j to (E 30 — K UJ UJ 0. 0. - 20 1 i IC — n - 1 l 1 0 GRAVEL TILL SHELL SAND SAND- SILT- SILT CLAY BOTTOM SEDIMENTS Figure 114 Density and biomass of Gastropoda in relation to bottom sediments. O O NUMBER • • WEIGHT 5 80 O s O 01 O z m / \ 5 " OF SPECIME METER OF O - \A IGHT IN GR METER OF 1 / M UJ y K UJ 40 uj tr — ^\ y * X m < => o z en / ^7 UJ o UJ 0. 20 ^ y_ i 2 ui o-^X 0 T ill1 -1 -0 0-J9 4-79 8-119 12-159 16199 2U-23S 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 UJ 2 I I 5 20.0 !9 NUMBER WEI8HT 0.01 0.8 1.0 1.9 2.0 3.0 PERCENT ORGANIC CARBON =»= — • i 12.0 10.0 z o 8.0 or u. o o z a 6.0 E uj 2* UJ UJ *5 4.0 a. uj o. 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 ivpes, and tem- perature range classes. They were especially plentiful on the continental shelf where their average density commonly ranged from about 50 to 500/m. 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 Nuculanidae. Size of specimens ranged from roughly 15 cm for Modiolus modiolus and Placopecten magellanicus to about 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 nourishment for many marine animals. Mammals, bit els, fishes, and invertebrates all have members that prey heavily upon bivalves. Bivalves occurred in 893 samples (83% of total). Their density averaged 163/m2 and biomass averaged 81 g/ m- (Table 5). C 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/m. High densities (>500/m2) were most common in nearshore areas. Moderate densities (50 to 500 indi- viduals/m2) occurred over extensive areas in the Gulf Composition and Distribution of Macrobenthic Invertebrate Fauna 127 GRAMS PER SQUARE METER BIVALVIA Figure 117 Geographic distribution ol Bivalvia: A — number of specimens per square meter of bottom; B — biomass in grams per square meter oi 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/m'-' 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 CO o Z CD UJ Z u. 5° X* to UJ u. UJ o s CC UJ ui cc CD < Z 3 2 ° z to 160 | 1 NUMBER mt WEIGHT n 80 180 140 5 O — 80 — 60 I UJ o Z UJ w * or < UJ O NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 1 1 8 Density and biomass of Bivalvia in each of tlie six geographic areas. England Slope. Lowest abundance (34/m2) was on Georges Bank, but there the average size was greatest. The biomass of bivalves was exceptionally high (aver- aged 166 g/m2) on the Southern New England Shell (Table 8; Fig. 118). Biomasses were moderate (30 to 77 g/m2) in Nova Scotia, Gulf of Maine, and on Georges Bank. Average biomass of bivalves was smallest (1 to 2 g/m'-') in the Georges Slope and Southern New En- gland Slope areas. Frequency of occurrence was high in all areas, espe- cially in the two slope areas where bivalves occurred in 89 .md 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 great- est 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 26/m2. The decrease in density was quite uniform; the major decre- ments in the trend were at about 25 m and 200 in. Composition and Distribution of Macrobenthic Invertebrate Fauna 129 600 O O NUMBER — 250 ♦ • WEIGHT 500 V \ \ _ 200 s o K>fc"00 Z CD \ \\ \\ s O K S"o > CD UJ \\ < Sfc I50»° o UJ (E \\ zi Q. UJ A * — tij ■"L'OO KUJ 11 -s A A IS f 3 \V \ \ \ — UJ 100*1 f10 200 — r \ UJ(/> * (r UJ £t UJ 0- \\ a. 100 1 \ Nw--^ \ ^ ^ 1 I I I T •-( »_i a i 50 0 10 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 119 Densitv 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/m . 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 silt- clay and decreased as particle size increased, except in shell bottoms where the density of bivalves was moder- ately high (180/m2). Gravels contained the lowest den- sity (39/m2). An entirely different trend was observed for bivalve biomass (Table 18; Fig. 120). The largest quantities (1 17 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. It should be noted that in four of the six sediment types, the biomass of bivalves accounted for over 42% of the total benthic biomass (Table 19). In 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%) of the samples from fine-grained sediments (Table 20). Only a moderately high propor- tion of the samples from sand and gravel contained live specimens of bivalves. 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 of increasing as the temperature range broad- ened, 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 20°-23.9°C class had a high density (1,195 individuals/ m2) but a small biomass (84 g/m2). Bivalve biomass was unusually large (106 to 334 g/m ) 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/m , 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 or- ganic 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 (64/m2) in the absence of or- ganic carbon to high levels (1,120/m ) 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 cor- responds to that shown for depth distribution, wherein higher bivalve densities occurred in the shallower es- tuarine and embayment waters that contained the high- est 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 (128/ m2) also occurred in low carbon levels (0.01-0.49%). The other carbon content classes contained significantly lower biomasses. Frequency of occurrence of bivalves in samples in the various organic carbon content classes was quite high 130 NOAA Technical Report NMFS 140 320 I — I NUMBER ■i WEIGHT S 240 O - en O z m o 5 cr UJ lj or CD < 2 => 3 O 2 V) (JO 1 00 ^ u. (9 O Z 2° Z (/) /\ ^ \ \ UJ w * a. < h- 3 Ct 400 UJ a. 1 \ ^ 1 \ UJ o or / \ / / \ UJ 60 0. / \ / / \ / \ / / \ 200 / o- " *-^^^^ / \ 0 4 l i 1 1 ' ' ' * i 0 001 0.5 10 15 20 SO 50 10 0 PERCENT ORGANIC CARBON Figure 122 Density and biomass of Bivalvia in relation to sediment organic carbon. A comparison of average density and average bio- mass 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 New En- gland 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/m'2) in moderately deep water, 100 to 1,000 m (Table 11; Fig. 1 25) . Increases and decreases in density about their center of abundance were surpris- ingly consistent and well correlated with changes in depth. The biomass of scaphopods (Table 13; Fig. 125) was small (<1 g/m'-') 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 bat In- metric classes were <0.01 g/m2. The average biomass was largest (0.17 to 0.98 g/m2) in moderately deep water (100 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%0 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 sedi- ments sampled, but were clearly more abundant (11 to 26 individuals/m2) in the shelly and Fine-grained sedi- ments 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 den- sity; however, the trend was the same (Table 18; Fig. Composition and Distribution of Macrobenthic Invertebrate Fauna 133 NEW \ NEW S\ \ JERSEY '.YORK/ \ \ NEW / \ \ hampshire ■• :onnecticut\ \ ;' »or'lano» SCAPHOPODA 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 espe- cially 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 12 □ NUMBER OF SPECIMENS METER OF BOTTOM - M WEIGHT I) O O (J) CO ° GHT IN GRAMS METER OF BOTTOM NUMBER SOUARE - UJ uj S or 04 < 1- 3 UJ O * « a UJ 0_ l J 1 02 2 0, 0 1 fl n 1 0 NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 124 Density and biomass of Scaphopoda in each of the six geographic areas. 5 u.a iz o * 2 in o o NUMBER • • WEIGHT 5 O i CD x s o — uj UJ cc s < uj CO 25 50 100 200 500 1000 2000 4000 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 20°-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 gener- ally 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/m-) in the two moderate carbon con- tent classes 1.50-1.99 and 1.00-1.49%, respectively, and Composition and Distribution of Macrobenthic Invertebrate Fauna 135 s 1 t- 1 1 NUMBER 2 £ 20 UJ I- 1- o s ■ — 08 I £ cr uj P-l UJ uj uj or * IT £< io ■ — M 4 5° | - 04 to r-, ,vn K UJ 0 K UJ a o j | J 1 1 GRJWEL TILL SHELL SAND S«N0 SILT- SILT CL4Y BOTTOM SEDIMENTS Figure 126 Density and biomass of Scaphopoda in relation to bottom sed Lments. tr uj UJ a: a) < 3 o 2 en o— -o NUMBER • * WEIGHT 06 cr u. O O 2 cr UJ o 0-39 4-79 8-119 12-15.9 1619. 9 20-53 9 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 127 Density and biomass of Scaphopoda 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% car- bon 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 Cepha- lopoda were not commonly encountered; they ac- counted for less than 0.1% of total biomass and density (Table 3). This apparent rarity is due primarily to sam- pling bias exemplified by the taxa represented in the samples. The class Cephalopoda contains some of the largest, most mobile, and most highly developed ma- rine invertebrates known to science; further, many are semipelagic or pelagic in habit and are therefore se- verely 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 v> o z o u I& 10.0 Si «)£ U. UJ OS IT UJ UJ (C CD < 1 Z> 10.0 9° Z NUMBER WEIGHT _1 I 0 01 0.S 1.0 15 2 0 JO PERCENT ORGANIC CARBON =•=— a — h z o £° <<" * u. 0 3 a 0 x UJ o 2 UJ uj * a < 0.2 t- 2 UJ O * •> » UJ 0 8 Z ° < ■ Z u S "■ 6° ° o s« * K inuj UJ u.Lj °6 — K H os <9 S ct uj UJ u uj cr * St m < §8 °< UJ o z or uj ooi £ C ■ ■ a 02 1 rl III NOVA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 130 Density ind biomass of Cephalopoda in each of the six geographic areas. s O O NUMBER s o j- l- • • WEIGHT S° 9& 5m < PECIM R OF - 0 04 go z£ MBER OF S 0UARE METE A : o o o o T WEIGHT QUARE MET = in UJ CO / \ 001 * rr UJ UJ 0. if i f i 4 i i "~-f^=^ i (j f> i V 10 25 " 50 100 200 500 1000 "2000 4000 " WATER DEPTH IN METERS Figure 131 Density an d 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/ur) in the ranges be- tween 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- 1 1 ,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 S u uj a. It 30 05 z to I 1 NUMBER ■ WEIGHT 1 — 0 02 2 * SAND SILT SILT- CLAY BOTTOM SEDIMENTS Figure 132 Density and biomass of Cephalopoda in relation to bottom sediments. 2 O s O O NUMBER • • WEIGHT y) 0 Z CD S« Hfe IT u. o o 0 15 V K z tr ~ UJ \- t- u. u O 5 10 A en UJ / \ Ul UJ CD < I O 05 z w 002 tig CE Ul °- 0 ^ \ 4 . " cc UJ 0 S 0-39 4-79 8-11.9 12-159 16-199 20-239 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 con- tent 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% of the total num- ber of specimens; 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 treat- ment, 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 NMKs 140 5 O to o Z CD o s cr uj uj O 2 CD toUJ CC UJ uj cc 600 CO O O NUMBER • • WEIGHT tr LLl a 800 600 400 — 200 40 35 30 5 o \- if) o 25 * CD tr IL o O z or UJ h- 20|- Ul X 5 o III UJ rr * < ) i*> h C3 UJ CD * tr hi CL 10 0 10 25 50 100 200 500 1000 2000 4000 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, 0.1/m2 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 dif- ferent temperature range classes. However, the broad- est 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 tem- perature 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 moder- ate 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 1 1 — 1 NUMBER 1200- M WEIGHT _ 20 1000 Z s o o ~~ 16 |_ H 0) fc «>o 80° z 2 5 m Zffi < UJ 9k u. zu. ° O G° UJ ~ z (T 0. * UJ $uj 1- H H ,2 I UJ " 0 Z u_ UJ OS 600 UJ u Or UJ * cr uj or 4 m< ►- 3 13 — UJ o 2° * «0 or or UJ £ 8 0- 400 . I \ 4 200 1 J 1 1 , J 0 0 GRAVEL TILL SHELL SAND S AND- SILT- SILT CLAY 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). \i at Im id. i — 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 be- tween 4° and 7.9°C. Because of the small size of members of the family Halicaridae, only a very small proportion of diem (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 of the macro- 144 NOAA Technical Report NMFS 140 1600 O O NUMBER • • • WEIGHT _ / - 24 '^ / 1400 /\ / / \ / 20 1200 / ^ / 5 O F t 1000 / V / \ 1 \\ / ■ 16 Z o z m UJ 3 m o: u. » O 5 q: OT UJ U, UJ 800 i / \y UJ 1- H I UJ O 5 i 12 o 2 CC UJ UJ uj uj q: % a 00 < 1 < S => i- 3 3 O uj q 2 en ,„„ 600 i / * <" DC IT UJ 1 UJ 0. i 0. 8 l 400 H / 4 200 i i i i i i 0-39 4-79 8-119 12-159 16-199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 138 Density and biomass of Arthropoda in relation to the annual range of bottom water temperature. benthic invertebrate fauna in terms of numerical den- sity. Aniphipoda, in particular, contributes an over- whelming majority of individuals (43%) to total fauna] 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 numeri- cal density. Detailed analysis of this class as a whole will appear below, along with the other dominant taxa, in the sec- tion "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 mem- bers of this group, only a small proportion were re- tained 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 stud)'. 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 1400 1200 2 § 1000 UJ no K 800 u. u oz CE UJ u a = § 60° - -o NUMBER • WEIGHT 1.0 15 2 0 3.0 PERCENT ORGANIC CARBON 20 X o DC UJ a. 100 Figure 139 Density and biomass of Arthropoda in relation to sediment organic carbon. Other studies (Wigley and Mclntyre. 19(34) revealed that ostracods, including specimens as small as 75 mi- crons or less, are not abundant in the offshore South- ern New England region. Their average density there was only slightly more than 1 individual/m . 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 conti- nental slope south of Georges Bank, and the continental rise east of New Jersey. Density of these ostracods averaged <0.1/rrr and their biomass <0.01 g/irr (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 sedi- ments: 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%. Densi- ties were somewhat greater at the higher levels than at the lower ones (Tables 26, 28). Cirripedia — Barnacles were generally sparse and, ex- cept 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 cur- rents, barnacles were common to very abundant (Table 3). Densities of nearly 8,000 individuals/m2 and biom- asses 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 \ NEW ,-/V\ ^\ JERSEY ;york/ \ \ NEW / \ \ hampshire Connecticut' V *■; INDIVIDUALS PER SQUARE METER NEW \ NEW ,'^\ \, JERSEY ; YORK/ \ \ NEW / \ \ HAMPSHIRE •' .CONNECTICU' ' EXPLANATION □ <0.l-0.9 GRAMS PER SQUARE METER PYCNOGONIDA Figure 140 Geographic distribution of Pycnogonida: A — number of specimens per square meter of bottom; B — biomass in grains per square meter of bottom. present in the collections, but those from the latter group were uncommon. The genus Bala mis was over- whelmingly 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 bar- nacles 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 I o o n NUMBER F- F- ■ WEIGHT £ ° 3 m z o < UJ or u. 2 u o o 5° Z IT £a Id ul uj oe — o 2 U. UJ o > - 0 02 UJ UJ or uj 3 o: UJ cr < S? < 04 1- O 2° Z 01 l J 1 001 £8 UJ o- „ 1 I 1 _i o. 0 NOVA GULF OF GEORGES SOUTHERN GEORGES SUU'MtHN SCOTIA MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SNELF SLOPE GEOGRAPHIC AREA Figure 141 Density and biomass of Pvcnogonida in each of the six geographic areas. O---0 NUMBER • • WEIGHT z 3 O 1- Jl — 3 03 k mass of P> cnogonida in relation to water depth. S o 1 — 1 NUMBER 3 F- F- W o ?o M WEIGHT s ° z to Z u. £ u- 5° oo« ° o a> 1.5 * K U>UJ UJ 1- F- u. UJ OS 1 0 — 0 04 O I or uj ■ UJ u uj a: * or o< < I = H 3 5° Z V) 05 - 0 02 u a or c u a. 0 1 r« . !-■ r-B t-l 0 UJ GRAVEL TILL SHELL SAND SANO- SILT- SILT CLAY BOTTOM SEDIMENTS Figure 143 Densit and biomass of Pvcnogonida in relation to bottom sediments. 148 NOAA Technical Report NMFS 140 £ 2 O o O O NUMBER J- (/) O • • WEIGHT »&- 5 a) UJ 5 u- 0= U. o° » o In ^ ; UJ u. uj 02 ,0 cr ui uj or - / / 1- 1- 0 03 x UJ o 5 UJ uj 0 02 * NEW / HAMPSHIRE EXPLANATION □ \ ' V ,- NEW / HAMPSHIRE -' MAINE 0 t20„. F\""" ^B ''••^.CONNECTICUT » \ j portlanojl-^ aJj^^ \fc N. N. N^ASSACHUSETTSV^'^^^Oj T4n ' ^v^ N. N. / \ BOSTON X^VSj-'s i. > t T3° ^- 100 _^ >>>W\^ C l000-__^^ ^~^-^_^ y' ^i^ 7Z» S \ X / > 0 ./ ( NOVA / ^r> scotia EXPLANATION ^s V^_^ O^^^ SI r X'^B ^"X^H^i D 1-49 X-~X_ % \ \ r>0 M 50-99 ■ 100-5074 ^ (_ ^?c INDIVIDUALS PER SQUARE METER x\A • >■ ' 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 of 8 to 329 m. They were substantially more plentiful (214 individuals/m ) in shallow water and diminished sharply in abundance with increasing water depth (Table 1 1; 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 dis- tribution was similar to that of numerical density. Bio- mass was largest (27 g/m'-') in shallow water and de- 152 NOAA Technical Report NMFS 140 2 O CO O Z ID UJ o "' t- U. UJ O 2 cr iij uj or CD < S D 3 O Z CO or UJ 0. 60 50 40 □ NUMBER WEIGHT 20 10- 1_D. — 12 10 X UJ O 2 UJ uj % cr < uj O * « K UJ a NOVA SCOTIA GULF OF MAINE GEORGES SOUTHERN GEORGES SOUTHERN BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 149 Density and biomass of Cirripedia in each <>f th 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 deepvvater 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 moder- ately deep water. Barnacles occurred in only a small proportion of the samples, but they were much more common (139! ) 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 den- sity was, surprisingly, highest (56 individuals/m2) in sand-silt sediments (Table 16; Fig. 151 ). Moderate den- sities (29 and 16 individuals/m ) occurred on gravel and sand bottoms. The other sediment types yielded low (<5/m ) densities. The biomass of barnacles was largest (11 g/m2) on gravel bottoms and was moderate (4.4 and 2.4 g/mL) in sand-silt and sand. Low quantities (<0.5 g/m2) pre- vailed in the other sediments (Table 18; Fig. 151). The unexpectedly high density of cirripedes in sand- silt bottoms resulted from high concentrations of small specimens (average weight <0.1 g) in a small propor- tion (3%) of the samples. Small barnacles densely colo- nize 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 of individual 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 o o NUMBER • • WEIGHT \ ~ A — 30 200 z X IMENS OF BOTTO o \\ \\ \\ - GRAMS OF BOTT o uj (r CL m (0 t- \\ \\ — 5 uj UJ o ■ \ 15 ^w org 100 I 155 iu oe uj < * < CD 3 3 ^ o 1- O =>U? UJ CO z \ 10 * ce o: UJ UJ 11 50 - \ a \V 5 0 |^£— r — ^-*— r.— -t^^i 1 f> 1 f 1 0 3 10 2! 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 150 Density and biom iss of Cirripedia in relation to water depth o s cc ui uj cr CO < S 3 3 o z w I — I NUMBER ■ WEIGHT l~l— Ik 1 SAND- SILT- SILT CLAY BOTTOM SEDIMENTS cc I. ° O Figure 151 Density and biomass of Cirripedia 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 O O NUMBER • • WEIGHT T 20 200 I - 18 180 / ~ 16 £ o z £ 160 to o / 14 o z m J f Z ° UJ I <5 u. a o Ife ,40 U / / 10 £ / / 12 = or UJ b. UJ 120 K K O 5 / / X UJ or uj ,0 O Z UJ CL / / UJ uj i< ,00 4 => a h- 3 z « / / UJ o a. 8 * w UJ 80 — / IT Q. / / UJ a. /\ 1 J 6 60 / \ \ / 1 \ y^ ~ 4 40 / / 20 qL * '-""' — • , i i i 2 0-39 4-79 8-1,9 12-159 16-199 20-239 0 ANNUAL RANGE ,N BOTTOM WATER TEMPERATURE ,N 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/m1' where the temper- attire range was nil or small. Their density steadily increased to 196/irr where the temperature range was greatest. Biomass similarly showed a marked increase in rela- tion to increased temperature range. The range in average biomass was from 0.05 to 20.32 g/m , the ex- treme 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 dasses except the highest (20°-23.9°C), where a mod- erate (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 densitv. Moderate densities (19 and 39 individuals/m2) occurred in the two carbon content range classes be- tween 0.01 and 0.99%; none occurred in content classes between 1.00 and 1.99%; but density increased dra- maticallv to 613/m2 where carbon content was from 2.00 to 2.99%, and was moderately high (83/rrr) in car- bon contents between 3 and 5% (Table 26; Fig. 153). Biomass displaved a similar trend but not as dramati- cally. Moderate biomass (6.5 and 4.1 g/mL>) occurred in the two range classes between 0.01 and 0.99%. with highest biomass (14.3 g/mL>) 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 uj a iS I NUMBER > WEISHT 12 0 O O O z rr UJ UJ l- 3 4.0 I S 20 5 0 PERCENT ORGANIC CARBON Figure 153 Density and biomass of Cirripedia in relation to sediment organi i bon. higher level classes, reflecting the trends established by density and biomass (Table 30). Copepoda — Only four of our samples contained speci- mens 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 or- ganic carbon content classes between 0.01 and 1.49%. Values of copepod biomass and density for each envi- ronmental parameter considered in this report may be found in Tables 6—30. Cumacea — Cumaceans are marine peracarid crusta- ceans that were widely distributed and well represented in New England waters (Theroux and Schmidt- Gengenbach'). Twenty-three species in 13 genera be- longing 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 ex- hibit 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 26/m2, and mean biomasswas0.ll 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 pref- erences 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 Gulf of Maine each contain extensive deep basins floored with fine muds, as well as shallower banks paved with 156 NOAA Technical Report NMFS 140 * — \ — ' /\ ^ K * — 7 V NEW \ N£w / \ \ JERSEY WORK/ *> \ NEW / "E ■■ / EXPLANATION □ 1-49 50-99 100-618 INDIVIDUALS PER SQUARE METER \ NEW .- \ \ EXPLANATION □ <0. 1-0.9 M 1.0-3.7 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. Musi of these areas were devoid of cumaceans (Fig. 154 ) . Average densities ranged from 1 to 618 individuals/ m . The majority of the region contained moderate densities (1-49/m ) with medium size patches of inter- mediate (50— 99/m2) density at continental shelf depths, and high (100-618/irr) 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 50 f^J NUMBER ■ WEIGHT 0 20 £ 4 0 O - 2 0 t- 1— to o i_ 0 16 S ° Z CD < m UJ <* u. 5 u- » O 50 30 — W IV ? or CL * UJ w w 0 12 H ,_ U_ UJ O 2 I UJ 13 5 tr uj 20 uj o: d - LU uj ? or CD < 2 3 0 08 1- § => O LU O z en - n S "> cc - or UJ Q. IC _1 1 1 . LU a 0 04 0 > — NOVA GULF 0^ GEORGES SOUTHERN GEORGES SOUTHERN SCOTifl MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 155 Density and biomass of Cumacea in each of the six geogra phic areas. low biomass (1-4 g/m2) along the shelf break on Georges Bank and the eastern shore of Cape Cod. Among the standard geographic areas, Georges Bank and the Southern New England Shelf yielded the highest mean densities (45 and 37/ m , re- spectively) and biomass (0.20 and 0.17 g/m2, re- spectively). 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 cumaceans (Table 10). 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 signifi- cantly higher mean densities than the deeper zones, ranging from 50 to 33/nr as depth decreased, whereas in the deeper (>100 m) ones they ranged from 7 to 0.7/ m2 as depth increased to 1,999 m. In the deepest zone (2,00-3,999 m) mean density was 2/m2 (Table 11; Fig. 156). The trend for biomass in relation to depth was simi- lar to that for density but was from one to three orders of magnitude lower. Mean biomass ranged from 0.26 to 0.5 O O NUMBER OF SPECIMENS METER OF BOTTOM 0 0 • • WEIGHT M - A \ ^ / A o GHT IN GRAMS E METER OF BOTTOM NUMBER PER SQUARE 1 1 p WET WE PER SQUAR ! ,i.i °— :i=~=^r=f^— 0 1 0 0 ' 10 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 156 Density and biomass of Cumacea in relation to water depth. 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 three between 25 and 200 m and the 500-999 m class, each yielded specimens in over 30% of the samples (range: 30- 158 NOAA Technical Report NMFS 140 50 □ NUMBER ■ WEIGHT S 40 — 0 20 | O in o gg Z O < UJ fc U. IL 0,5 °° 2 a 5° 30 _ — °- fi CO uj o z 2 2 Ui 20 - 0 10 * ft m< 1° z to - 1 rr , respectively), with smallest (1/m2) amounts in till substrates (Table 16; Fig. 157). The trend for biomass was essentially simi- lar 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 cum- aceans ranged from 50 to 26%, in samples from sand, sand-silt, and gravel and was be- tween 14 and 22%< in the other sediment types (Table 20). Relation to Water Temperature Both measures of cumacean abundance showed a bi- modal trend in relation to annual range in bottom water temperature. The greatest numerical abundance ( "><> irr ) and biomass (0.26 g/m2) occurred where tem- perature range was moderate (8-11.9°C), followed by t 0 O NUMBER A • • WEIGHT o 60 trt o z m A - 0 20 O Z^ UJ 50 Z u. 1 u. NUMBER OF SPEC SQUARE METER o o - 0 16 012 0 06 UJ I ui o Z UJ UJ * ir < i- = w O n- 20 cr // ^ Ul UJ 0- o. 004 10 c^ i i i i i 0 0 0-39 4-79 S-ll-9 12-159 16199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 158 Density and biomass of Cumacea in relation to the annual ran ^e of bottom water temp erature. another peak in areas experiencing a somewhat higher ( 16-19. 9°C) range where high density (44/nf) 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 159 125 0 u. u o » c u UJ (C £ < 50.0 3 3 3 O z \ NE" / \ \ HAMPSHIRE -' ^YORKX ~N_.rONN£CTICUT\ \ ;' PORTlANDf INDIVIDUALS PER SQUARE METER ^ — \ — ' 7^\ — ' ^ ' 7 NEW \ NEW ,' \ \ ; JERSEY '.YORK/ \ \ NEW / \ \ HAMPSHIRE .' GRAMS PER SQUARE METER 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 None were present in quantitative samples from the continental shelf even though pre- vious studies and our own nonquantitative samples have revealed their presence in cer- tain habitats in the intertidal and shallow sub- littoral areas along the coast. They appear to be rare or absent from most of the outer conti- nental 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 av- erage biomass of 0.01 g or less/m . Members of this group were present in 15% of the samples from Georges Slope and in 9% of samples from Southern New En- gland 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/m ) in depths greater than 2,000 in than at shallower depths, where they averaged only 0.1 of an individual/irr. Biomass revealed a trend similar to that of numerical density. The biomass averaged <0.01 g/nr 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%< occur- rence in the shallower portion of their bathy- metric range (Table 15). Relation to Sediments Tanaidaceans were found only in soft, fine- grain 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 temperaMre range was less than 8°C (Tables 21, 23, 25; Fig. 164). All of die measures of abundance (density, biomass, and frequencv of occurrence ) were very low in each of die two temperaftire range classes (0°-3.9° and 4°-7.9°C) in which tanaidaceans occurred. I — | NUMBER z o t- h- tn o Z CD UJ ■ WEIGHT z o <" * u. 3; £ u. Q O z ? cr UJ l- h- I UJ cs Z NUMBER PER SQUARE o - \ ~L - 001 UJ uj S cr < UJ o * W CC UJ Q. 02 NOVA SCOTIA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 161 Density and biomass of Tanaidacea in each of the six geographic areas. O— -O NUMBER z o • • WEIGHT Z o »- }- v>0 Sfc zo UMBER OF SPEC 5QUARE METER — ro - l- r 2 - ¥uj uj or - A 001*4 1- O UJ tn Z « «/ / * cc or UJ i g i a i g i gM — o — -t-\f£t — ° i i UJ EL 0. 0 ) 10" 25 " 50 "100 "200 500 "lOOO 2000 4000 " WATER DEPTH IN METERS Figure 162 Density i nd biomass of Tanaidacea in relation to water depth. z o z o 1 — 1 NUMBER t- UJ O 2 m .UJ Z u. 5° ■ WEIGHT cc u. o o OF SPE METER o ro z f f 4? K UJ 0-39 4-79 8-119 12-159 I6-T9.9 20-539 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 164 Density and biomass of Tanaidacea in relation to the annual range of bottom water temperature. s o 0.004 3 1- »- fs O O NUMBER A •— — • WEI8HT i: JL o.a 0.003 50 © 0 a 0: £* U. UJ wt /A 0.002 hi 1- 1- X UJ OS 0 s a: uj w a: A A *£ S < 0.1 _ U.I 0.OOI 4 9° *- 2 UJ O s ° g_ —1 — 4 ' ft • s a • ct w 0.01 03 1.0 " 18 " 20 " 3.0 " 50 " 10.0 a. PERCENT ORGANIC CARBON Figure 165 Density and biomass of Tanaidacea 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 translu- < cm to light tan or medium brown. Approximately 13 species of isopods were represented in the collections, most of which belonged to the fami- lies Cirolanidae and Idoteidae. A new species, Chiridotea arenicolaWigley ( 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% of total). Their density averaged 12.1/m and their biomass averaged 0.29 g/m- (Table 5). Geograph ic I) is tribution 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 conti- nental shelf and rise. Densities between 10 and 50 indi- Composition and Distribution of Macrobenthic Invertebrate Fauna 163 NEW \ NEW s\ \ JERSEY '.YORK/ \ \ NEW / \ \ HAMPSHIRE ,- CONNECTICU' V EXPLANATION □ 1-9 10-49 50-520 INDIVIDUALS PER SQUARE METER v \ ' 7^\ ' ^ ' 7 ^ NEW \ NEW ,' \ \ JERSEY ■ YORK/ '» \ NEW / \ \ HAMPSHIRE -' CONNECT 1 1 1SSACHUSET ISOPODA 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/m extended over large areas of the Gulf of Maine and Southern New England. High densities of 50 to 520 individuals/m2 occurred in reladvely few small areas. Av- erage biomass was commonly less than 1 g/m , 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 major difference was that cumaceans were twice as numerous as isopods, but the relative densities of the two groups corre- sponded 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 South- ern 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 indi- viduals/m2) in the two shallowest depth classes and decreased as water depth in- creased. Lowest density (0.4/m2) was en- countered 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 pat- terns 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/m'J) in water depths less than 500 m than in deeper water where the average biomass ranged from only 0.01 to0.02g/m-. Frequency of occurrence of isopods was highest (46 to 69%) in samples from wa- ter depths ranging between 25 and 100 m. Occurrence rates were intermediate (12-20%) in samples from outer conti- nental shelf and continental slope depths (100-2.000 m) and moderately high (43%) at depths greater than 2,000 in (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/ mJ) in sand substrates than in any other type of bottom. The average density of isopods in the other sediment types was fairly uni- form, ranging only from 3 to 7/nr. The average biomass of isopods was highest (1.36 g/m2) in till substrates (Table 18; Fig. 169). Also, as was the case 5 O OS CE UJ UJ CC CD < 13 2° 2 W i NOVA SCOTIA I — | NUMBER ■I WEIGHT fk rr z o X UJ a z * or * v> GULF OF GEORGES SOUTHERN GEORGES SOUTHERN MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 167 Density and biomass of Isopoda in each of the six geographic areas. O— — O NUMBER • • WEIGHT 5 2 40 O lo 3° ''A - Jl CD GRAMS OF BOTTO UJQ. u-S - / A s / \ zL5 O * 20 04 -£ § 3 - \ / \ lu tn 3 «» - 02 tE UJ a ■ 1 1 1 ~~»-T- £> -T "* 1 0. o 0 i 10 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 168 Density and biomass o Isopoda in relation to water depth. z o 1 — 1 NUMBER ■ WEIGHT z 14 O h- Z GO 25 I ? z ° 2 < D Z U. S u. SPECI tER 0 to O 10 ° 2 CC OB _£ U. UJ |5 O 5 X UJ IS Z 0 6 - CC UJ ijj or .A CD < IU UJ u * at 0 4 |- ? => O . , * w or UJ a 0 n 1 i . 02 cr UJ 0- 0 GRAVEL TILL SHELL SAND SAN0- SILT- SILT CLAY BOTTOM SEDIMENTS Figure 169 Densiiv and biomass of Isopoda in relation to bottom sed iments. Composition and Distribution of Macrobenthic Invertebrate Fauna 165 en o z CD u. UJ O 5 or u uj or CD '^4 i 0 0.01 0.9 10 1.9 2.0 3.0 9 0 10.0 PERCENT ORGANIC CARBON Figure 171 Density . md biomass of Isopoda in relation to sediment organic carbon. new forms were described by Edward L. Boitsfield (1965), National Museum of Canada: Protohaustorius wigleyi, Parahaustorius longimerus, P. holmesi, P. attenuatus, Pseudohaustorius borealis, Acanthohaustorius millsi, A. in- termedins, and A. spinosus. At least three suborders are represented in the collections: Gammaridea, Caprel- lidea, and Hvperiidea. Both the species and the num- ber 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 largest species in our collections, the caprellid Aeginina longicornis, had a body length of more than 2 cm. Gammaridea larger than 1.5 cm (Casio, Maera, and a few others) were uncommon. Color of amphipods ranged from light cream or nearly white (Lysianopsis, Ampelisca, and many others) to mod- erately dark brown (Leptocheims, 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 significant that at least two of the most common genera (Ampelisca and Unciola) are tube dwellers. Also Haploops, which is one of the few amphi- pods 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 aver- aged 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 exception- ally abundant group and densities in the coastal areas and on the offshore banks commonly averaged be- tween 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 GRAMS PER SQUARE METER 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 continen- tal slope and rise. The biomass of amphipods in the coastal areas and on the offshore banks generally aver- aged between 1 and 50 g/m2. In the central Gulf of Maine and on the continental slope and rise, the biom- ass 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 (1 18 to 280/m2). The South- 168 NOAA Technical Report NMFS 140 □ £ 1000 in o z ID OS Q: UJ lj UJ * 1 UJ i- . F i. w 6 i-hj o5 eoo 6 \ 1 \ 1 I2 Q- UJ \ 1 UJUI SO: \ 1 So: m< \ 1 < i§ 600 4 |1 o: or UJ UJ 0. 400 — \l CL V 2 200 0 10 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 174 Density and biomass oi Amphipoda in relation to water depth. from one type to another (Tables 16, 17; Fig. 175). Density was exceptionally high (1,238 tndividuals/m2) in sand and contributed over 56% of the total number of animals in this type of sediment. Density was moder- ately high in gravel; intermediate in sand-silt, till, and shell; and relatively low (23/irr) in silt-clay. Biomass trends were the same as those for density (Table 18; Fig. 175). Sand sediments contained the largest bio- mass, an average of 7.7 g/m2. Intermediate quantities (0.5 to 3.4 g/m2) were present in gravel, till, shell, and sand- silt. The lowest biomass (0.18 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 tempera- ture range variations occurring in the study area (Table 21; Fig. 176). Highest average density (1,372 individu- als/m2) was encountered where the temperature was intermediate (8°-11.9°C). Densities were moderately high (598-809/m2) in areas where the range in tem- perature was broad, but were substantially lower (58- 31 1/irr) where the temperature range was nil or small. Amphipod biomass in relation to temperature range paralleled the same trends as those revealed by numeri- cal density (Table 23; Fig. 176). Greatest biomass (8.1 g/mL) occurred in an intermediate temperature range class (8°-11.9°C). Moderate values occurred in adja- cent broader and narrower range classes. The smallest (0.41 g/m ) 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 moder- ate. 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 or- ganic carbon (Table 26; Fig. 177). Two abundance peaks were clearly evident. Densities were very high (between 1,000 and 1,256 individuals/in-') 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 ex- 1 70 NOAA Technical Report NMFS 140 5 O w o 800 — Z CD £ H 3, 500- I — 1 NUMBER tm WEIGHT i n. is UJ * K UJ o GRAVEL TILL SHELL SAND SAND SILT- SILT CLAY BOTTOM SEDIMENTS Figure 175 Density and biomass of Amphipoda in relation to bottom sediments. tremes. Lowest density occurred in the absence of mea- surable 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/m'-') 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% of the samples) in the organic carbon content classes contain- ing 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 5/ m2 and average biomass less than 0.01 g/m . Some species of mysids characteristically make diurnal excur- sions from the sea bottom to the upper water layers, at which time they become members of the plankton com- munitv; 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 bot- tom 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 to O O NUMBER 1400 - 1200 - k 6 '/ \ X s o o t- 1- 1000 _ 95 to o Z CD UJ - 6 Z u. a O OF SPECI METER 0 CD o o 5 o: UJ I UJ O 5 UJ UJ NUMBER SQUARE O O - / \ 4 < UJ O * » CE CE UJ UJ 0. a. 400 - 2 200 J i 1 1 1 1 1 n 0 0-3.9 4-79 8-119 12-159 16-19.9 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS 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 or- ange color patches on the body. Mysids occurred in 41 samples (nearly 4% of the total). Their density averaged 2.5/m2, and biomass av- eraged 0.01 g/m2 (Table 5). Geographic Distribution Mysids were sparsely distributed in the study area oc- curring 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. Average density was low (1-49 individuals/m ) in all localities except for two small patches of moderate density (50-99/m2) 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, re- spectively. The other geographic areas contained signifi- cantly lower amounts, ranging from 10 to 100 times less in terms of density and 6 times less in terms of biomass. 172 NOAA Technical Report NMFS 140 1400 1200 - OI000 to o z m Ul S U. r>° 800 ft1.*, ff> w H II "J o * CC UJ 600 UJ CE a < » 3 3 O Z (0 400 200 O O NUMBER •— — • WEIGHT - 10.0 8.0 2 o K u. 6.0 ° ° Z CC 1 £ UJ Ul 4.0*5 t- 2 8 IE Ul a 2.0 0.01 0.5 0 15 2 0 3.0 PERCENT ORGANIC CARBON 100 Figure 177 Density and biomass of Amphipoda in relation to sediment organic carbon. 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 less/m2. Biomass, although very small, revealed a rather con- stant 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 individuals/ 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 l/m\ 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 samples (6%) from sand sediments 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 tempera- ture broadened (Tables 21, 23; Fig. 182). Average nu- merical density increased from <0.1/m2 where the tem- perature range was <4°C, to 6.1/m2 in localities where the temperature range was over 209C. 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 of samples yielding specimens Composition and Distribution of Macrobcnthic Invertebrate Fauna 173 NEW \ NEW JERSEY ; Y0RK \ NEW .- \ \ V NEW / \ \ HAMPSHIRE nciir'* N^tASSACHUSEr /^\ BOSTON -R I > T v ' -r*r =■ r ■; 7 NEW \ NEW S \ \ JERSEY ; YORK/ V \ NEW / \ \ hampshire .' .Connecticut'*. \ • pobtlano# EXPLANATION D 8 — < Sfc oo u &;5 6 VA^ 003 ?jS n- UJ 4 ./X \ 002 iS NUMBE >ER SOUAR ro - v. - o o WET W PER SQUA I ill n— 1 — n__ T\, l (l f 1 °o~ 10 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 180 Density and biomass of Mysidacea in relation to water depth. 1 6 1 — 1 NUMBER 2 O o m u. O 4 cr UJ i- Ul s ■ WEIGHT o o GHT IN GRAMS METER OF BOT UJ tr 2 O en ft UJ a- _ rl rl j o WET WE PER SQUARE GRAVEL TILL SHELL SAND SAND- SILT- SILT CLAY BOTTOM SEDIMENTS Figure 181 Density and biomass of Mysidacea in relation to bottom sediments. Frequency of occurrence of mysids was higher (7 and 8%) in the intermediate car- bon content classes than in the lower ones (3 and 5%) in which they occurred (Table 30). Decapoda — Decapods, although well rep- resented 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 bio- mass (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 taxo- nomic 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 con- tain 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 con- tinental 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/m , 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 representa- tives of more sedentary, less active taxa, such as the burrowing sand shrimp Crangon septem- spinosits. Average anomuran size ranged between 4 and 20 mm carapace length. Most frequently captured were representatives of the relatively Composition and Distribution of Macrobenthic Invertebrate Fauna 175 slow-moving pagurid hermit crabs. Size of brachyuran crabs averaged between 15 and 20 mm carapace width; the two species of Cancel- did provide some larger speci- mens in the 50 to 60 mm range, and Pinnixa spp. 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 ex- tremes, 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 ma- rine animals (including other invertebrates, mammals, birds, and fishes) depend on de- capod prey for a substantial portion of their sustenance. Decapods occurred in 246 samples (23% of the total). Their density averaged 8 indi- viduals/m2, and their biomass 1.32 g/m2 (Table 5). Geographic Distribution The most striking feature of decapod distri- bution revealed by our samples was their apparent absence from large portions of the Gulf of Maine (Fig. 184). This artifact is due primarily to sampling gear bias since, tradi- tionallv, this region has been well known for the high annual yields of lobsters and shrimps. How- ever, low densities were recorded from the northeast corner of the Gulf at the entrance to the Bay of Fundy, the Western Basin section north of Great South Chan- nel, and some inshore localities. Low densities (1-49 individuals/ m ) 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) O O NUMBER WEIGHT _s^r i z o 5 ° 003 1 m o 5 * a: < kj o 0 01 * EXPLANATION □ <0.l - 99 M 10.0-49.9 ■ 50.0-64.8 GRAMS PER SQUARE METER DECAPODA Figure 184 Geographic distribution of Decapoda: A — number of specimens per square bottom; B — biomass in grams per square meter of bottom. meter of biomass, mostly in inshore regions near Cape Cod and Long Island. One small area of moderately high biom- ass (50-65 g/irr) occurred on the southeast part of Georges Bank. Decapods occurred in five of the six standard geo- graphic areas; they were absent only in the Georges Slope area. Highest average density (22/m2) occurred on Georges Bank followed by significantly lower densi- ties on the Southern New England Shelf and off Nova Scotia (9 and 2/m2, respectively). Density was below 1/ 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 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 of occurrence of decapods 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. Bath y metric Distrib it tio n Decapods were almost wholly restricted to water depths of less than 500 m and showed a general trend of diminishing in abundance as water depth increased (Table 1 1 ; Fig. 186). Average densitv 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 (0.1/rrr) in the only deepwater depth class (1,000-1,999 m) in which they occurred. Average biomass generally followed the trend established for densitv (Table 13; Fig. 186). In the shallow-water depth class, aver- age 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 deep- est class in which decapods occurred ( 1000- 1999 m). Decapod frequency of occurrence in the samples was fairly uniform at moderate lev- els (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 (24/ m2) of decapods as well as greatest biomass (5.56 g/ m2). A drop in density occurred in shell and sand, each of which yielded an average of 9/m2, nearly three times fewer decapods, but biomass diminution was not as dramatic in shell which contained an average of 4.78 g/ 25 [ — | NUMBER m WEIGHT Z 20 o - z o 1- 4 <"S (/> o s £ Z ID Z ahl in uj 0: lu ■ UJ U, * 5 15 z o S to < o o X Z — UJ uj a: $« I- O uj en * 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 186 Density and biomass of Decapoda in relation to water dt pth. mz; however, sand contained nearly five times less bio- mass (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, rang- ing 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 ten- dency to increase with broadening temperature range to 178 NOAA Technical Report NMFS 140 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 (0-y3.9°C) and generally increased with broadening tem- perature range, peaking at 23.2/m'-' and 4.26 g/ m2 in the 16.0-19. 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). Relation to sediment organic carbon Decapods were absent in areas where no mea- surable organic carbon occurred in the sedi- ments, 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 bio- mass described U-shaped distributions (Tables 26, 28; Fig. 189). Average density ranged from nearly 8 to 0.5 individuals/m2 and average bio- mass from 4.15 to 0.30 g/m . Values were high- est at the extremes of the 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 mod- erate, ranging from 4 to 32%, but in this instance was biased slightly toward the lower end of the content range (Table 30). tr uj uj ct CD < 3° 2 V) 1 — 1 NUMBER IB WEIGHT l 1 J 6 5 4 3 2 0 < J- 3 UJ o SAN0- SILT SILT- CLAY GRAVEL TILL SHELL SAND BOTTOM SEDIMENTS Figure 187 Density and biomass of Decapoda in relation to bottom sediments. 0 O NUMBER s • • WEIGHT 2 5 O o 1- 1- 25 m •- r- £ O trt O I m z m < UJ 4 o: u n 2° - //\\ o o OF SPE METER UJ _ / 1 V 6HT IN METER NUMBER SQUARE o - /x/ / V \ WET WE R SQUARE £ 5 a. — <7 i uj r- 1 1 1 1 1 0-39 4-79 8-11.9 12-15.9 16-199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 188 Density an d biomass of Decapoda in relation to the annua range of bottom water temperature. Bryozoa Bryozoans are sessile colonial animals most frequentlv 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 En- gland benthic fauna. The majority of these species be- long to the class Gymnolaemata, order Cheilostomata. The calcareous encrusting forms were especially nu- merous, 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 patchv distribution rather severely limited their contribution to the total benthic fauna. In terms of nu- merical 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 aver- aged 1.29 g/m2 (Table 5). Composition and Distribution of Macrobenthic Invertebrate Fauna 179 10.0 6.0 1 O I- w o Zo III X u. o° <** tow U. UJ OS tt uj UJ CE CD 3 4 0 I! z in — o NUMBER — • WEIGHT 4.0 ± a: u ►- >- I UJ O 3 UJ u * E 2.0 ,_ 5 UJ o * K UJ a 10 IS 2 0 SO 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 off- shore 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 continen- tal 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 mod- erately 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 con- tained 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 1 1 % in the southwest. Their occur- rence 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 39/m2 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 shal- low 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 any one depth class (Table 13; Fig. 192). 1 80 NOAA Technical Report NMFS 140 NEW ' \ NEW ,.^\ \ y / 7^. 0 7 JERSEY '.YORK/ ^ \ Ne* r^3 *&. ,—f \ \ HAMPSHIRE MAINE ' ( <■ X. \J§j|*, \.MASSACHUSETrt)^--*~'"^£__V*i*^__, /^•s. ^!5^JSife S\ BOSTON /_ . "\ / ^\itV _/-'r i > XT?" >~iiP \ \ ^^^vr^ o 7\ JZik^ « l2)o / J NOVA W) SCOTIA EXPLANATION vv V^~ ^^ r^- ^*^Hs_i B 1-49 s^\^ X i\ o H 50-99 ^-^V ■ 100-798 _ N<^- |— i — VH^*-"^. J ' A L> INDIVIDUALS PER SQUARE METER v f y^ ' ^ ' >• \ JERSEY ', rORK/ '- \ «» / .j» ^ __/ \ \ HAMPSHIRE BRYOZOA Figure 190 Geographic distribution of Bryozoa: A — number of specimens per square i of bottom; B — biomass in grams per square meter of bottom. The percentage of samples containing bryozoans gen- erally decreased with increasing water depth. The high- est incidence of occurrence (19%) was in the shallow- est depth class. At depths greater than 500 m, they occurred in from 0 to 5% of the samples (Table 15). Relation to Sediments Brvozoans 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 bio- genic 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 of specimens and 7.5% of the biomass of the total benthic fauna in shell substrates (Tables 17, 19). Gravel substrates ranked second in quantity, with an aver- age density of 75 colonies/nr and a biomass of 7.4 g/ni". Quantities were low (less than 6 colonies and 0.4 g/nr ) in the remaining four sediment types. Composition and Distribution of Macrobenthic Invertebrate Fauna 181 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 Watei Temperature Although bryozoans were rather severely lim- ited in distribution by specific substrate require- ments, water temperature range appeared to play a lesser role in inhibiting abundance. Mem- bers of this phvlum occurred in all tempera- ture range classes but revealed a pronounced trend of increasing densitv with a broadening of the temperature range (Table 21; Fig. 194). The density of bryozoans averaged only 3 colo- nies/m2 where the temperature range was less than 4°C. Their density increased to an average of 66 colonies/m2 where the temperature range was greater than 20°C. Biomass values also exhibited a general up- ward trend (0.28-2.45 g/m2) as the tempera- ture range broadened. This increase, however, was less consistent than that exhibited by nu- merical 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 tem- perature range was broad (Table 25). Relation to Sediment Organic Carbon The relationship of bryozoans to sediment or- ganic carbon was not nearly as well defined as that in other parameters because no orderly trend or pattern was discernible. Relativelv high average densities, between 21 and 35 individual colonies/ m'-, occurred in widely separated or- ganic carbon content classes in the low. middle, and higher regions of the content spectrum (Table 26; Fig. 195). Significantly lower aver- age densities, ranging between 0 and 8/ni'-', 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 pro- nounced. 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 occur- rence 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. to o Z ffi Id 3 !!; o° u_ LU O S CE UJ UJ 01 CD < 3° Z CO 5 0 I — I NUMBER I uj O GEORGES SOUTHERN GEORGES SOUTHERN BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 191 Density and biomass of Bryozoa in each of the six geographic areas. O 0 NUMBER • • WEIGHT — 3 0 2 O gjjj A s o 25co£ 2 CD UJ tr u. -O / \ o o ° 00 — ^ / \ 20ZLt CL Lu — UJ i — 0 °0 10 25 50 100 200 500 1000 2000 4O00 WATER DEPTH IN METERS Figure 192 Density and biomass of Bryozoa in relation to water depth. Brachiopoda 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 182 NOAA Technical Report NMFS 140 5 O U) O Z CD UJ 2 u. 5° OjUJ U. UJ O S or UJ uj a CD < 2 3 3° Z 05 or UJ a 360 320 280 240 200 160 120 80 40 I — I NUMBER HI WEIGHT TILL SHELL SAND- SILT- SILT CLAY — 18 16 s o t- 14 f- %a or u_ z ce UJ T- UJ UJ UJ * or < UJ O *<" DC UJ a. BOTTOM SEDIMENTS Figure 193 Density and biomass of Bryozoa in relation to bottom sediments. 80 o o NUMBER CIMENS OF BOTTOM o • • WEIGHT O _ / / / GRAMS OF BOTTOM NUMBER OF SPE PER SQUARE METER ro t> O O / WET WEIGHT IN PER SQUARE METER 1 1 1 1 1 1 0-39 4-79 8-119 12-159 16199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 194 Density and biomass of Bryozoa in relation to tlie annual ra nge of bottom water temperature. 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 of suitable habi- tats. 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 num- bers of species, there being only about 280 known living species, our samples contained only members of one genus, Terabratulina. Size of specimens 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 predominat- ing; many specimens were fouled by other sessile forms such as bryozoans, hydroids, and encrusting sponges of various hues, but tans, browns, yellows, and grass predominated. Composition and Distribution of Macrobenthic Invertebrate Fauna 183 400 s o u. u OS (E Ul Ul or § < 20.0 50 z co IE Ul a. 10.0 O O NUMBER • • WEIGHT 2 0 1.6 »5 5 CD < 1.2 a 11 19 0 ? or UJ 1- 1- I UJ 0 2 UJ uj * or 0 8 < LU O * » 0.4 0 01 0.5 10 15 20 * 30 PERCENT ORGANIC CARBON Figure 195 Density and biomass (if Bryozoa in relation to sediment organic carbon. Brachiopods occurred in 54 samples (5% of total). Their numerical density averaged 4.5 individuals/m2, and their biomass averaged 0.89 g/m- (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/m ) over most of their range as was biomass (<10 g/m2); small areas of moder- ate (between 50 and 100/m2) density and biomass (10 to 63 g/m2) 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/ m2) did occur in a few places, notably at the mouth of the Bav of Fundy. Among the six standard geographic areas brachio- pods were restricted to Nova Scotia, the Gulf of Maine, and Georges Slope. Largest average density (22/m2) and biomass (3.68 g/m2) occurred off Nova Scotia fol- lowed by Gulf of Maine (9.5/m2 and 2.12 g/m2); Georges Slope contained insignificant quantities (Tables 6, 8; Fig. 197). Twenty-one percent of the samples off Nova Scotia contained brachiopods versus 12% in the Gulf of Maine; only 2% of the Georges Slope samples yielded speci- mens (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/m2 and biomass nearly 4 g/m2. 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 1 84 NOAA Technical Report NMFS 1 40 74° 73" 72° NEW \ NEW ,-yr\ \ JERSEY . YORK/ \ \ NEW / \ \ HAMPSHIRE •" V /a EXPLANATION □ 1-49 H 50-99 ■ 100-490 INDIVIDUALS PER SQUARE METER NEW \ NEW ,' \ \ / JERSEY ', YORK/ \ \ N£w / \ \ HAMPSHIRE -' ONNECTlCUT^ \ ;' POBTLANOf \"ASSACHUSETTS" . BOSTON R . I. > GRAMS PER SQUARE METER 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 icoarsei types over the finer ones (Tables 16? 18; Fig. 199). Till sediments ranked first in preference, contain- ing an average of 48 individuals/irr and an average biomass of nearly 16 g/m2. Interestingly, although shell sediments ranked sec- ond in terms of density (37/m2), they ranked last (0.22 g/m'-') in terms of biomass. Gravel, with a mean density and biomass of 14/m2 and 2.44 g/m2, respectively. Composition and Distribution of Macrobenthic Invertebrate Fauna 185 ranked third. Both mean density (range 0 to 2/ m2) and mean hiomass (range 0.24 to 0.33 g/m2) were significantly lower in sand, sand-silt, and silt- clay, each of which offered very limited attach- ment potential. Samples in till substrates yielded the highest oc- currence frequency of specimens (41%), shell and gravel were about even, but considerably lower in over- .tll frequency than till (17 and 14%, respectively; Table 20). Only 2 to 3% of samples in the other sediment types provided brachiopod specimens. Relation to Water Temperature Brachiopods were quite restricted in their rela- tion 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. The\' were most plentiful (average density of 9/nr 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). Relation to Sediment Organic Carbon Brachiopods preferred low levels of sediment or- ganic carbon (Tables 26, 28; Fig. 201), being ab- sent at levels above 1 .49% carbon content. Nu- merical 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 slighdy lower biomass (1.31 g/ m ) 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, bv members of five classes: Crinoidea, Holothuroidea, Echinoidea, Ophiuroidea, and Asteroidea. All but Crinoidea provide significant con- tributions to the total benthic fauna. In terms of contri- bution to overall density, members of Echinoidea are second in dominance, providing 20% of the total num- CO 0 z m UJ Z . q: uj uj a 13 I — I NUMBER ■ WEIGHT 2 - I UJ UJ uj * a: < i- id UJ o * "> GEORGES SOUTHERN GEORGES SOUTHERN BAN" NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 197 Density and biomass of Brachiopoda in each of the six geo- graphic areas. 20 ENS BOTTOM cri t 0 o NUMBER fo\ • • WEIGHT 4 I O t- "o I CO < <*£. - O / \ o O OF SPEC METER CD PJ '- / \ 2 .-£ cr£ 6 \ uj U ct * Hi 10 1.9 2 0 3 0 PERCENT ORGANIC CARBON 10 0 Figure 201 Density and biomass of Brachiopoda in relation to sediment organic carbon. Holothuroidea — Holothurians formed a moderate com- ponent of the New England benthos in terms of bio- mass, but made up only a minor portion of the total number of specimens (Table 3). This was due mainly to the relatively large size of individual specimens. Mem- bers 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 stand- point 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 uni- form 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 to- tal). Their density averaged 4.3/mL' 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 Shell 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/ irr. 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/m'-. Among the six standard geographic areas, holothuri- ans were present in largest quantities in the Gulf of 188 NOAA Technical Report NMFS 140 -too r iooo EXPLANATION □ 1-49 50-99 100-751 INDIVIDUALS PER SQUARE METER ECHINODERMATA Figure 202 Geographic distribution of Echinodermata: A — number of specimens 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 per/m2 and the bio- mass 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 <>l the total number of specimens in all areas, but made up bstantial pan of the total weight in two areas (Table 9). In the Gulf of Maine area they accounted for 22% of the total fauna! weight and on the Southern New England Slope the\r made up 14% of the total fauna! 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% of the 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 o CO o Z CD ?fe OS CE UJ uj cr CD < 2D 60 Z> O z wi I — | NUMBER ■ WEIGHT 120 100 aJ s o — 40 e> 5 * o: < uj o * en ?0 GULF OF GEORGES SOUTHERN GEORGES SOUTHERN MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 203 Density and biomass of Echinodermata in each of the six geographic areas. z£ UJ 1 ^ feS 80 I"" O O NUMBER • • WEIGHT 0L-~~—L UJ q: 80 * 3 win 500 1000 2000 4O00 WATER DEPTH IN METERS Figure 204 Densin and bomass of Echinodermata in relation to water depth. 190 NOAA Technical Report NMFS 140 I o I- 120 iooI- 5 * C u 80 — S 3 3 O z in \i I □ NUMBER ■ WEIGHT K U. 40 U u < *■" 30 a: GRAVEL TILL SMELL SANO SAND- SILT- SILT CLAY BOTTOM SEDIMENTS Figure 205 Density and biomass of Echinodermata 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/ m ) at inter- mediate 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 sub- stantially 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 (1 to 6 g/m2) were found in other depth classes. Individual holothurians from shallow water (0 to 21 m) were larger, averaging nearly 25 g each, and size decreased with increasing depth to less than 1 g each at depths greater than 500 m. The frequency of holothurian occurrence was highei 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 indi- viduals/m2, whereas, in the other types of sediments their density was only 2.0 to 7.4/m2. The relationships of holothurian biomass to sedi- ments were entirely different from those pertaining to density. The biomass was high (25 to 29 g/m2) in sand- silt and silt-clay (Table 18; Fig. 211). In fact, holothuri- ans 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 of relatively fewer but larger specimens in soft sediments and numerous small specimens in till sediments accounts for the disparity- between the biom- ass and density values in these substrates. In the other sediment types they were generally more equally dis- tributed in density and biomass. Composition and Distribution of Macrobenthic Invertebrate Fauna 191 280 O o NUMBER . • • WEIGHT J leo - A / " 240 160 z / _ 200 2 o t- : /M / : O Y- 140 w o Z OD UJ 2 u. cc u. SPECII TER 0 o 160 a° z cc — UJ 1- t- U_ UJ O 2 100 / / V / I UJ CC UJ 120 S ^ UJ CC CD < ? Z> 80 / / \\ / '*" $ CC t- 3 2° Z in / / \\ / uj o CC CC UJ p 1 \ 1 UJ Q. 60 so a 40 ' y/ / k - 40 20 I 1 III II 0 0-39 4-79 8-119 12-159 16-19.9 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 206 Density and biomass of Echinodermata in relation to the annual i ange 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/m 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/m') where the temperature range was small (<4°C). With increased annual temperature range, the biomass of holothuri- ans increased substantially to an average of 128 g/m2 where the range was greatest (20°-23.9°C). Size disparity of specimens in the various tempera- ture range classes was the principal cause of the re- versed trends. The size of specimens increased mark- edly in waters having a broad temperature range, whereas the numerical density diminished slightly. Holothurians contributed a moderately large pro- portion (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 contrib- uted an extraordinarily large share (>30%) of the bio- mass (Table 24). Frequency of occurrence was quite uniform at a mod- erate level (14-25%) in all temperature range classes except one (16°"19.9°C), in which holothurians con- tributed only 8% of the samples (Table 25). 192 NOAA Technical Report NMFS 140 O O NUMMft •— • WEI«HT f 100 600 - |- - 400 2 is G5 / '•• . Ss 8« \ Z K U fc¥ •» - aoo£t fa EjK U kJ ■a * « NUM PER SOU / > < too £ J W \ / to Y \ 100 _/ \ 1 ^^* V~ 1 0 •""l 1 1 1 1 1 '"^-i i 0 001 0J 10 1.9 2 0 JO 5 0 " 10 0 PERCENT ORGANIC CARBON 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 or- ganic carbon-biomass relationship was essentially posi- tively correlated. Seven of the eight carbon content classes were occupied; the one exception was the high- est class (5.0+%). Holothurian average density was highest (18 indi- viduals/m ) where no measurable organic carbon was found (Table 26; Fig. 213). Much lower average densi- ties (ranging from 8 to 2/m'J) prevailed in the other carbon content classes with a general tendency oi de- creasing 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. !13). Average biomass was lowest (6 g/m ) in the 0% carbon content class and steadily increased with in- creasing organic carbon content, culminating in excep- tionally large biomasses in the two highest classes occu- pied. 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 mod- erate 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 bio- mass, 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 E* \ new y./\ \ \ \ PER SQUARE METER NEW \ NEW .- \ \ JERSEY WORK/ '« \ NEW / \ \ HAMPSHIRE ..' CONNECT rfASSACHUSET EXPLANATION □ <0.l - 9.9 H 10.0- 49.9 ■ 50.0-379.8 GRAMS PER SQUARE METER HOLOTHUROIDEA 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 pre- ferred 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 u. Ul OS a uj UJ cr ISS I — I NUMBER ■ WEIGHT GEORGES SOUTHERN GEORGES SOUTHERN SANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 209 Density and biomass of Holothuroidea in each of the six geo- graphic areas. s 0 0 NUMBER X o 1- • • WEIGHT o 1- co o «»S Z IT u. O O — 20 o s UJ uj I- 3 UJ O Density and biomass of Holothuioidea in relation to bottom sediments. O O NUMBER • • WEIGHT cr uj UJ CE co < 5 o 3 o z in 1 o 8- <* u. u o -80 Z cc -100 I 120 0-3.9 4-79 8-119 12-159 16-19.9 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 212 Density and biomass of Holothuioidea 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. Low- est biomass averages occurred in the slope areas (Table 8; Fig. 215). Echinoids were present in only moderate to small percentages of the total number of samples. 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 individu- als/m2) near mid-shelf depths (25 to 49 m) and dimin- ished in both shallower and deeper regions (Table 1 1; 196 NOAA Technical Report NMFS 140 600 0 0 NUMBER • • • WEIGHT 11 - A - 500 / 1 - a o £20.0 / 1 - 400 o 1- Id / \ , o 5fe <\ j \ _ < £ uj i \ <° o I- Ul u. uj O X 15.0 - joo£ t (9 3 £t UJ uj a: \ / \ UJ u tiD < * ) 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 in. Occurrence of echinoids in the samples was moder- ate to low in all depth classes and followed the same trend as densitv and biomass. They were present in 57% of the samples near the mid-shelf depths and decreased in samples from both shallower and deeper bathymet- ri< c lasses (Table 15). sity averaged 67 individuals/m 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/irr; in all other sediments the biomass of echinoids averaged 7.3gorless/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 exception- ally large contribution for one taxon (Table 19). The occurrence of echinoids in the samples was mod- erate to low in all sediment types (Table 20). As expected, it was highest (47%) in sand substrates. Some- what unexpectedly, the incidence rate was lowest (S'V ) 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 of sediments, their den- Relation to Water Temperature Ki hinoids 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°C. Composition and Distribution of Macrobenthic Invertebrate Fauna 197 NEW \ NEW ,,/ \ JERSEY '.YORK/ J^BS*,0"" \ ^v— CONNECT 1 ^- iSO — ^-^V__ \ NEW / \ \ HAMPSHIRE .' ^UT^ ' : P0«TL4ND*-— \massachusettS^^V-^^^ ) /\ Bosicr* ^ ' M A 1 N E ^ a / 120 .. / <* A \ EXPLANATION ^ii. H 10-99 (— -n ■ 100-1051 INDIVIDUALS PER SQUARE METER NOVA SCOTIA \ , NEW \ NEW ,.-/ \ \ JERSEY . york/ K- \ NEW / \ \ HAMPSHIRE .' .CONNECTICUT**, \ ; post-land iassachusett: BOSTO' 'r I •> 7 GRAMS PER SQUARE METER ECHINOIDEA 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 mid- range (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/m where the temperature change was 2()°C or more. The biomass of urchins was distributed in essentiallv the same manner as their numerical density among the various temperature range groupings, with one excep- tion (Table 23; Fig. 218). Large biomass (148 g/m2) occurred in the mid-range; moderate biomass (about 14 to 38 g/m ) in intermediate ranges; and small biom- ass (6 g/m ) in the stable areas. The one exception was a relatively large biomass (135 g/m2) in the broadest (20°-23.9°C) temperature range class. 1 98 NOAA Technical Report NMFS 140 120 | — | NUMBER HI WEIGHT 100 - 120 OF SPECIMENS METER OF BOTTOM O O - - 3 8 GHT IN GRAMS METER OF BOTTOM NUMBER PER SQUARE o — o WET WE PER SQUARE — 40 20 1 1 20 0 rl rl 1 . . 0 NOVA SCOTIA GULF OF GEORGES SOUTHERN GEORGES SOUTHERN MAINE BANK NEW ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA Figure 215 Density i ind biom ass of Echinoidea in each of the six geographic areas. 140 o -o NUMBER • • WEIGHT 160 120 t— 7\ s 1 \ z o 1- if> fc 100 _ 1 \ W o ,z, £ 120 Z O SIS _ 3 fc o o z K CL ui W K 1- ui °5 60 II I — 80 I Z ££ Jl UJ LE IS 40 \ S < ui m z $ i — 40 (E 20 \ \^-«-^^ °o - 10 25 50 100 200 500 1000 2000 4000 WATER DEPTH IN METERS Figure 216 Density ; nd hi imass i >l Echinoidea in relation to water depth. Composition and Distribution of Macrobenthic Invertebrate Fauna 199 5 O z CD U. UJ O 5 cr uj uj cr CD o Z 10 OE / / \\ / 1- o uj o UJ a. / / \ w UJ a 20 / / \ ~ 40 1^ 0 ? 1 1 1 1 1 0 0-39 4-79 8-119 12-159 16-199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 218 Density and biomass of Echinoidea in relation to the annual range ( f bottom water temperature. about 25 mm and arm lengths over 12 cm, to small specimens of Amphipholis and Ophiura, with disc dia- meters of less than 2 mm and arms 10 mm long. The coloration exhibited bv ophiuroids in our samples was varied. Some genera, such as Ophiura and Ophiomusium, were uniformly whitish or light gray. Oth- ers were more colorful because of their mottled pat- terns of contrasting hues, including dark red, pink, brown, and orange. Among the more brightlv colored genera were Ophiopholis, Amphiura, and Ophioscolex. Ophiuroids obtain their food bv 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 spe- cies generally combines the ingestion of bottom mate- rial with selective carnivorous feeding. The diet thereby consists ol detritus, diatoms, and other small-size foods, as well as polychaete worms, crustaceans, bivalve mol- lusks, and other similar upes of organisms. Brittlestars. in turn, are preyed upon bv other echinoderms, but mosi significantly by demersal fishes. Ophiuroidea occurred in 487 samples (45% of the total). Their density averaged 44.2/m2 and their biom- ass averaged 3.26 g/m (Table 5). Geograph ic 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 New Jersey region. Biomass distribution of ophiuroids tended to parallel their density distribution. Moderate (1—10 g/m ) and large (10-80 g/mL>) 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) Composition and Distribution of Macrobenthic Invertebrate Fauna 201 58 20 0 B ?o o K u IB.O II So N ■E Q E O O NUMBER • • WEIGHT too" o <• "o X u W.O S u w * DC a u 5 * •> K E so i.o i* ta ».o KRCENT OMANIC CAMOM 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/m-, 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 of these organisms was mod- erately 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% of samples 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 pro- nounced zone of high abundance (35 to 87 individu- als/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 \ NEW ,-X\ ^\ JERSEY '.YORK/ N \ NEW / \ \ HAMPSHIRE .' 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 10c? ) 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 of samples (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 203 100 5 o o z in 80 20 — LaJLal I I NUMBER ■1 WEIGHT JJ GULF OF GEORGES SOUTHERN GEORGES SOUTHERN MAINE BANK NE* ENGLAND SLOPE NEW ENGLAND SHELF SLOPE GEOGRAPHIC AREA 5 O or u. o o 1 s UJ UJ * CC < I- => UJ O 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/ nr) in gravel, sand, and shell; intermediate (38 and 58/m2) in till and silt-clay; and high (94/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 den- sity. Small biomasses of ophinroids 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 ophinroids in the samples re- vealed 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%) of the samples in till, sand-silt, and silt-clay (Table 20). Relation to Water Temperature Ophiuroid density, biomass, and frequency of oc- currence all conformed generally to the same trend of high abundance where the temperature range was less than 16°C, and low abundance ,-,° 60f- 1° o o NUMBER • • WEIGHT _ 6£g o O z a * < 500 1000 2000 4000 WATER DEPTH IN METERS Figure 222 Density and biomass of Ophiuroidea in relation to water depth. 204 NOAA Technical Report NMFS 140 2 O CO O Z CD Id o V) w U_ LU O 5 Or LU w or 40 CD < S =) => o Z CO ^ □ NUMBER ■I WEIGHT \ 111 GRAVEL TILL SHI i 2 O X LJ C£ 5 UJ u, * or S CC UJ UJ uj UJ CC CD < X 3 uj O * ■ BOTTOM SEDIMENTS Figure 229 Density and biomass of Asteroidea 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 indi- vidual/m2 and mean biomass from <0.01 to 0.04 g/m2 (Tables 16, 18; Fig. 234). Fre- quency 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 (<0.1/m2 and <0.01 g/m2) in the lowest temperature range class; mean den- sities of 0.2/m2 occurred in each of the two Composition and Distribution of Macrobenthic Invertebrate Fauna 209 2 O r- 3 O— -o NUMBER i o t- • • WEIGHT z m ijj ,\ * u. 4 <= O ? Z 01 uj O // "-\ _ , *- DC 0. UJ 0 #ll| , 0. 0 0-39 4-79 8-119 12-159 16 199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 230 Densit \- and biomass of Asteroidea in relation to the annual range of bottom water temperature. o 0 NUMBER 6 . • • WEIGHT 5 ■ o 1- z m J> AMS BOTTOM IlL s« u. UJ OS _ 01 GHT IN GR METER OF CC UJ is ii UJ uj £8 v 2 - - UJ 0 l ^~~~*W i i \ - 0 Ul 0. 0-39 1-: o z m o* (E UJ UJ IE (D < 2 3 3 O Z (O I — I NUMBER ■ WEIGHT A 2 O »o a: u. o o z a UJ UJ *5 K UJ TILL SHELL SAND BOTTOM SEDIMENTS SANO SILT- SILT CLAY 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 they were restricted to the low to mid- range 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 of density 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 fauna! 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 re- port, refers to the taxonomic group that mathematically contributed the highest number of individuals or greatest total accumulated wet weight. Results are expressed in both measures of abundance because of the marked dif- ferences that existed between them. In spite of individual disparity in rank order within each measure of abundance, members of four taxo- nomic 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 chief com- ponent, as well as Gastropoda, Scaphopoda, Poly- placophora, 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 of the above taxa to the total density and biomass of the New England macrobenthic fauna. Composition and Distribution of Macrobenthic Invertebrate Fauna 215 O— -o NUMBER 60 — • • WEIGHT 16 I z O ih o J- z o 1 \ Ul 9° 40 / \ 12 2 „, 5oc R OF S E METE o A / \ - CD EIGHT E METE uj a- * oc 4 lg 20 _ /i \ \ / \\ UJ O Z (0 oc y i \ \ \\ 4 or UJ "■ 10 / 1 XJ \ UJ a. i i i i i T 0 0-39 4-79 8-119 12-159 16199 20-239 ANNUAL RANGE IN BOTTOM WATER TEMPERATURE IN DEGREES CELSIUS Figure 241 Density and biomass of Ascidiacea in relation to the annua range of bottom water temperature. 25.0 20 0 ss U. UJ O 3 OC uj * § l0 ° Z tft 8.0 - NUMBER WEIGHT 001 0.5 10 l» 2.0 3.0 PERCENT ORGANIC CARBON 50 -•— ^ 3.0 OC o u. o z oc UJ H 1- T UJ O Z UJ * oc < l.U \- s * in a. UJ 100 Figure 242 Density and biomass of Ascidiacea in relation to sediment organic carbon. 216 NOAA Technical Report NMFS 140 Frequency of Occurrence 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% occur- rence rate. For comparative purposes, some nondominant taxa showed inter- mediate frequencies; among these were Coelenterata, which occurred in 42% of the samples, and Nemer- tea and Sipunculida with frequen- cies of 34% and 23%, respectively. Ascidiacea, Bryozoa, Aschelmin- thes, Porifera, Brachiopoda, Pogo- nophora, Turbellaria, and Hemi- chordata were encountered with diminishing frequencies ranging from 17% tol Asteroidea. Composition and Distribution of Macrobenthic Invertebrate Fauna 223 Leptasterias spp. Strongylocentrotus drobachiensis Echinarachnius parrna 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 cm) 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 americanusH. Milne-Edwards (Fig. 247), the American lobster, is, together with the sea scallop, one of the most commercially valuable invertebrate resources of the northwest Atlantic. It is widely distributed through- out 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 cm average length, for individuals cap- tured for market by the inshore fishery, to very large (sometimes in excess of 80 cm), for specimens in the offshore stock; minimum legal size for capture is cur- rently 8.13 cm (3.25 inch). Lobsters are scavengers and inhabit a variety of substrates. 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 occa- sionally in the diet of long-horned sculpin. Pagurus spp. (Fig. 248) comprise seven species of pagurid hermit crabs resident in the New England re- gion. The species of the genus Pagurus in the family Paguridae represented in this study include P. acadianus, P. anmdipes, P. arcuatus, P. longicarpus, P. politus, P. polMcaris, 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 Cirolanidae resident in the study area (C. concharum, ('.. impressa, and C. polila) 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 star- fish or purple star, is one of the most common star- fishes inhabiting the offshore waters of the New En- gland region and is a member of the family Asteriidae. This is a large species commonly between 15 cm and 30 cm (6-12 inch) in diameter; some specimens up to 42.5 cm (17 inch) have been reported from the northern limits of its range. It is normally found on sandy bot- toms where it is a very important predator of bivalve mollusks. Juvenile specimens are occasionally encoun- tered in fish stomachs. Leptasierias spp. (Fig. 249), which represent several species of the genus Leptasierias, also of the family Asteriidae, are common inhabitants of the New En- gland Region on sandy bottoms. These brightly, but variably, colored species are of moderate size (5-10 cm) and in some localities are veiy abundant. Small specimens are occasionally preyed upon by some spe- cies of groundfish. Echinarachnius parma (Lamarck) (Fig. 249), the north- ern 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 of the region are typically 7.5 cm in diameter. They are a common prey of flounders, haddock and cod. Strongylocentrotus droebachiensis (Miiller) (Fig. 249), another ubiquitous echinoid (family Strongylocen- trotidea), 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 northeast- ern U.S. and Canadian waters. Size ranges from 5 cm to nearly 9 cm. 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 En- gland region; included are O. ljungmani, O. robusta, O. sarsi, and unidentified species. Members of this 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. Bathymetrie 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/m . Substantially lower densi- ties occurred in waters deeper than 200 m. Annelida had the next highest densities in shallow waters (719/ m-') 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. Mol- lusca showed a similar pattern of density distribution with depth; density was greatest (570/m2) in shallow water, moderate (136-205/m'-') at shelf depths, and moderately low in deeper waters. However, mollusks were numerically dominant at the deeper depths. Echi- noderm density was greatest (133/ 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 (47/m2) occurred in shallow (0-24 m) water. Mollusca was the dominant fauna] component of biomass at nearly all depths. Greatest biomass (258 g/ in'-') 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. Echino- derm 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 shelf depths but decreased rapidly with increasing depth beyond 200 in. Crustacea, although numerically domi- nant, 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 of the four dominant faunal com- ponents in relation to the six major sediment types encountered in the New England region did not ex- hibit 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 bot- toms as well. Mollusks were generally more abundant in silt-clay (354/m2), shell (229/m2), and sand-silt (276/ m2) but were found in somewhat lower abundance in other sediments also. Crustacea were most abundant in sand (1,336/m2), gravel (710/m2), and sand-silt (275/ m2) and were found in diminishing amounts in shell (124/m2), till (59/m2), and silt-clay (34/m2). Sand-silt (104/m2) and sand (95/m2) contained the most echi- noderms, followed by till and silt-clay (67/m2 and 65/m2, respectively), then shell (28/m2) and gravel (23/m2). Density rank order in the various sediments listed by decreasing particle size was as follows: gravel: Crusta- cea, 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 small- est 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 g/ 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/m2 and 6 g/ ur 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 g/m2), 43 g/m2 in silt- clay, and 37 g/m2 in sand-silt. Median amounts oc- curred in till (15 g/m2) and lower amounts in gravel (6 g/m2) and shell (3 g/m2). Biomass rank order in bot- tom sediments was as follows: gravel: Mollusca, Crusta- cea, Annelida, Echinodermata; till: Echinodermata, Annelida, Mollusca, Crustacea; shell: Mollusca, Anne- lida, Crustacea, Echinodermata; sand: Mollusca, Echi- nodermata, Annelida, Crustacea; sand-silt: Mollusca, Echinodermata, Annelida, Crustacea; and silt-clay: Echi- nodermata, 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 m , 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 Echinoder- mata were fairly consistent at moderate levels (84-345/ m2 for Mollusca and 21-171/m2 for Echinodermata) throughout the temperature range spectrum. Mollusca, however, did make a strong showing (1,242 individu- als/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 den- sity 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; 12-1 5. 9°C: Crus- tacea, 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 hiomasses to annual range of bottom water temperature was similar to that of density in that no definite trends were evi- dent. However, a marked change in dominance rank- ing prevailed, wherein the density dominants (crusta- ceans and annelids) were replaced by echinoderms and mollusks as the leading contributors to biomass in nearly all temperature range regimes. Echinodermata domi- nated 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. Mol- lusk biomass, second to that of echinoderms in most temperature ranges, was clearly dominant where tem- perature 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 bio- mass was as follows: 0-3. 9°C: Echinodermata, Annelida, Mollusca, Crustacea; 4—7. 9°C: Echinodermata, Mollusca, Annelida, Crustacea; 8-11.9°C: Mollusca, Echinoder- mata, Annelida, Crustacea; 12-1 5. 9°C: Echinodermata, Mollusca, Annelida, Crustacea; 16-1 9. 9°C: Mollusca, Echinodermata, Annelida, Crustacea; 20-23. 9°C: Echi- nodermata, 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 fannal abundance except in a few exceptional cases (Tables 26, 28; Fig. 26). The numerical abundance of the dominant taxa var- ied 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 m . Crustacean density varied widely among the various carbon content classes, ranging from 21 to 1,357/ m2. Greatest abun- dances occurred in carbon content levels of 0.01-0.49% (1,066/m2) and 3.00-4.99%, (1,357/m2), with moder- ately low to fairly high densities occurring in carbon content levels between these two. Lowest densities (21- 22/m2) 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, show- ing a positive correlation of generally increasing density with increasing carbon, ranging from 69/ m2 in areas devoid of carbon to 1,120/m where carbon was between 3.0% and 4.9%. No mollusks occurred where carbon content exceeded 5%. Annelida were present in all or- ganic carbon content classes. Their density was signifi- cantly lower at both extremes of the carbon content spec- trum (between 11/m2 and 81/m2) compared with their abundance (196-504/ m2) in areas containing low (0.01- 0.49%) to moderate (2.0-2.99%) amounts of carbon. Rank order of the numerical abundance of the domi- nant taxa with regard to organic carbon content was as follows: 0%: Mollusca, Annelida, Echinodermata, Crus- tacea; 0.01-0.49%: Crustacea, Annelida, Mollusca, Echi- nodermata; 0.50-0.99%: Annelida, Crustacea, Mollusca; Echinodermata; 1.00—1.49%: Mollusca, Annelida, Echi- nodermata, Crustacea; 1.50-1.99%: Mollusca, Annelida, Crustacea, Echinodermata; 2.00-2.99%: Crustacea, Mollusca, Annelida, Echinodermata; 3.00-4.99%: Crus- tacea, Mollusca, Annelida, Echinodermata; 5.00%+: Crustacea, Annelida; no Mollusca or Echinodermata were found in this class. Similar to numerical abundance, biomasses of domi- nant taxa showed no clear-cut correlation to the or- ganic 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 mod- erately high carbon contents (between 2% and 4.99%), and lowest (6 g/m2) occurred in areas devoid of mea- surable 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%). Mol- lusca, also absent where the highest measures of or- ganic carbon occurred, nevertheless showed a prefer- ence 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 biom- ass (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 rang- ing between 1 1 g/m2 and 27 g/m2 in areas of organic carbon content ranging between 0.01 and 4.99%,. Low- est mean biomass of annelids (0.11 g/m2) occurred in the highest carbon content class (5+%) and intermedi- ate 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/m'-' where organic carbon was between 2% and 2.99%. Moderately low biomasses (between 1 g/m2 and 9 g/ m ) 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, Crus- tacea; 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%: Echi- nodermata, Mollusca, Annelida, Crustacea; 5.00%+: Annelida and Crustacea were equal, whereas Mollusca and Echinodermata were absent in this class. 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, Tho- mas J. M. Schopf, Eve C. Southward, J. 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 help- ful 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 J. Walsh), Astenas (Capt. Arthur D. Colburn Jr.), and Gosnold (Capt. Harry Seibert). Literature Cited and Selected References 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 as- sisted 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 nu- merical 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, plot- ting, and data processing. Drafting assistance was pro- vided bv 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 partici- pated in shipboard work were K. O. Emery, John C. Hathaway, Jobst Hiilsemann, Frank Manheim. Robert H. Meade, Richard M. Pratt, David Ross, John S. Schlee, James V. A. Trumbull, and Elazar Uchupi. 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