LU )7P-OR Sh La Sy S : UCT 02 2007 4) LIBRARIES NOAA Technical Report NMFS SSRF-783 Biomass and Density of Macrobenthic Invertebrates on the U. S. Continental Shelf off Martha’s Vineyard, Mass., in Relation to Environmental Factors Don Maurer and Roland L. Wigley July 1984 U.S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service ty Sys) NOAA TECHNICAL REPORTS National Marine Fisheries Service, Special Scientific Report—Fisheries The major responsibilities of the National Marine Fisheries Service (NMFS) are to monitor and assess the abundance and geographic distribution of fishery resources, to understand and predict fluctuations in the quantity and distribution of these resources, and to establish levels for optimum use of the resources. NMFS is also charged with the development and implementation of policies for managing national fishing grounds, development and enforcement of domestic fisheries regulations, surveillance of foreign fishing off United States coastal waters, and the development and enforcement of international fishery agreements and policies. NMFS also assists the fishing industry through marketing service and economic analysis programs, and mortgage insurance and vessel construction subsidies. It collects, analyzes, and publishes statistics on various phases of the industry. The Special Scientific Report—Fisheries series was established in 1949. The series carries reports on scientific investigations that document long-term continuing programs of NMFS, or intensive scientific reports on studies of restricted scope. The reports may deal with applied fishery problems. The series is also used as a medium for the publication of bibliographies of a specialized scientific nature. NOAA Technical Reports NMFS SSRF are available free in limited numbers to governmental agencies, both Federal and State. They are also available in exchange for other scientific and technical publications in the marine sciences. Individual copies may be obtained from D822, User Services Branch, Environ- mental Science Information Center, NOAA, Rockville, MD 20852. Recent SSRF’s are: 722. Gulf menhaden, Brevoortia patronus, purse seine fishery: Catch, fishing activity, and age and size composition, 1964-73. By William R. Nicholson. March 1978, iii + 8 p., 1 fig., 12 tables. 723. Ichthyoplankton composition and plankton volumes from inland coastal waters of southeastern Alaska, April-November 1972. By Chester R. Mattson and Bruce L. Wing. April 1978, iii + 11 p., 1 fig., 4 tables. 724. Estimated average daily instantaneous numbers of recreational and com- mercial fishermen and boaters in the St. Andrew Bay system, Florida, and adja- cent coastal waters, 1973. By Doyle F. Sutherland. May 1978, iv + 23 p., 31 figs. 11 tables. 725. Seasonal bottom-water temperature trends in the Gulf of Maine and on Georges Bank, 1963-75. By Clarence W. Davis. May 1978, iv + 17 p., 22 figs., 5 tables. 726. The Gulf of Maine temperature structure between Bar Harbor, Maine, and Yarmouth, Nova Scotia, June 1975-November 1976. By Robert J. Pawlowski. December 1978, iii + 10 p., 14 figs., 1 table. 727. Expendable bathythermograph observations from the NMFS/MARAD Ship of Opportunity Program for 1975. By Steven K. Cook, Barclay P. Collins, and Christine S. Carty. January 1979, iv + 93 p., 2 figs., 13 tables, 54 app. figs. 728. Vertical sections of semimonthly mean temperature on the San Francisco- Honolulu route: From expendable bathythermograph observations, June 1966- December 1974. By J. F. T. Saur, L. E. Eber, D. R. McLain, and C. E. Dorman. January 1979, iii + 35 p., 4 figs., 1 table. 729. References for the indentification of marine invertebrates on the southern Atlantic coast of the United States. By Richard E. Dowds. April 1979, iv + 37p. 730. Surface circulation in the northwest Gulf of Mexico as deduced from drift bottles. By Robert F. Temple and John A. Martin. May 1979, iii + 13 p., 8 figs., 4 tables. 731. Annotated bibliography and subject index on the shortnose sturgeon, Aci- penser brevirostrum. By James G. Hoff. April 1979, iii + 16 p. 732. Assessment of the Northwest Atlantic mackerel, Scomber scombrus, stock. By Emory D. Anderson. April 1979, iv + 13 p., 9 figs., 15 tables. 733. Possible management procedures for increasing production of sockeye salmon smolts in the Naknek River system, Bristol Bay, Alaska. By Robert J. Ellis and William J. McNeil. April 1979, iii + 9 p., 4 figs., 11 tables. 734. Escape of king crab, Paralithodes camtschatica, from derelict pots. By William L. High and Donald D. Worlund. May 1979, iii + 11 p., 5 figs., 6 tables. 735. History of the fishery and summary statistics of the sockeye salmon, On- corhynchus nerka, runs to the Chignik Lakes, Alaska, 1888-1956. By Michael L. Dahlberg. August 1979, iv + 16 p., 15 figs., 11 tables. 736. A historical and descriptive account of Pacific coast anadromous salmonid rearing facilities and a summary of their releases by region, 1960-76. By Roy J. Wahle and Robert Z. Smith. September 1979, iv + 40 p., 15 figs., 25 tables. 737. Movements of pelagic dolphins (Srenel/a spp.) in the eastern tropical Pa- cific as indicated by results of tagging, with summary of tagging operations, 1969-76. By W. F. Perrin, W. E. Evans, and D. B. Holts. September 1979, iii + 14p., 9 figs., 8 tables. 738. Environmental baselines in Long Island Sound, 1972-73. By R. N. Reid, A. B. Frame, and A. F. Draxler. December 1979, iv + 31 p., 40 figs., 6 tables. 739. Bottom-water temperature trends in the Middle Atlantic Bight during spring and autumn, 1964-76. By Clarence W. Davis. December 1972, iii + 13 p., 10 figs., 9 tables. 740. Food of fifteen northwest Atlantic gadiform fishes. By Richard W. Langton and Ray E. Bowman. February 1980, iv + 23 p., 3 figs., 11 tables. 741. Distribution of gammaridean Amphipoda (Crustacea) in the Middle At- lantic Bight region. By John J. Dickinson, Roland L. Wigley, Richard D. Bro- deur, and Susan Brown-Leger. October 1980, iv + 46 p., 26 figs., 52 tables. 742. Water structure at Ocean Weather Station V, northwestern Pacific Ocean, 1966-71. By D. M. Husby and G. R. Seckel. October 1980, 18 figs., 4 tables. 743. Average density index for walleye pollock, Theragra chalcogamma, in the Bering Sea. By Loh-Lee Low and Ikuo Ikeda. November 1980, iii + 11 p., 3 figs., 9 tables. A io Nonwuis™ NOAA Technical Report NMFS SSRF- 783 Biomass and density of Macrobenthic Invertebrates on the U. S. Continental Shelf off Martha’s Vineyard, Mass., in Relation to Environmental Factors Don Maurer and Roland L. Wigley July 1984 U.S. DEPARTMENT OF COMMERCE Malcolm Baldrige, Secretary National Oceanic and Atmospheric Administration John V. Byrne, Administrator National Marine Fisheries Service William G. Gordon, Assistant Administrator for Fisheries The National Marine Fisheries Service (NMFS) does not approve, recommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication fur- nished by NMBS, in any advertising or sales promotion which would indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. 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CHE CC mages ocr cvs teres fercces sic avetotasvs se vekch areola ats aie (Goevey oh 6 eseiatesede a tae STI ST MR UST OREICT HRY ECON T SITE oe Maree Crch eee aN Jo TO WAU COTD, <3 ca chal rat RARER ERR RCECRO ISIC OT eC RAL ICRO RE CLD cic eon eke omic rene ci cath oi cry SICAL EI kG oie O eR NEREEE nol io Eso Peeead ERO clO PWN Figures . Location of bottom-sampling stations for fauna and sediment off Martha’s Vineyard, Mass., June 1962.............. . Bottom-sediment types off Martha’s Vineyard, Mass., from samples collected at stations, June 1962................. . Bottom temperature off Martha’s Vineyard, Mass., from measurements taken at stations, June 1962. ............... Quantitative distribution of biomass for all macrobenthic invertebrates combined off Martha’s Vineyard, Mass., from samplesicollectediatistations umes 96257. 5.ccycre ere ce etree eter oie ete NST eS eee Poe eee nt te Quantitative distribution of individuals for all macrobenthic invertebrates combined off Martha’s Vineyard, Mass., fron samplesicollectediat stations; June 1 S62 a racyaes erect secs cesta eee ee TE Teche Ro ona rel RDN eres . Summary of average biomass and number of individuals of major taxa in relation to depth, temperature, median sedi- ment size, and silt-clay, off Martha’s Vineyard, Mass., from samples collected June 1962.....................00000s Table . Comparison of wet weight biomass of macrobenthos in terms of bathymetric stratum of the U.S. northeastern Atlantic ill © OO OO © © & 0 0H 01 WW MWAAAANAAAMUMAUNHFPHPAPNNNNNN w 11 hWhN eS Appendix Tables . Station location and environmental variables off Martha’s Vineyard, Mass., June 1962. ...................0..0002- . List of macrobenthic invertebrates collected off Martha’s Vineyard, Mass., June 1962. .................0 20 ee eeeuee . Wet weight biomass of major taxa of macrobenthic invertebrates per station off Martha’s Vineyard, Mass., June 1962. . Number of individuals per m? of major taxa of macrobenthic invertebrates per station off Martha’s Vineyard, Mass., NUNEWOG2 emieciseictseietcie oe site es nctocinen Gaede hemes teciidee debs dees Raa S38 USS eas See eee 14 16 17 19 Biomass and Density of Macrobenthic Invertebrates on the U.S. Continental Shelf off Martha’s Vineyard, Massachusetts, in Relation to Environmental Factors! DON MAURER? and ROLAND L. WIGLEY? ABSTRACT The mean density and mean biomass of macrobenthic invertebrates on the U.S. continental shelf off Martha’s Vineyard, Mass., were 3,008/m? and 245.7 g/m’, respectively. The latter estimate was considerably higher than values from the North Sea, Scotian Shelf, and Middle-Atlantic Bight. Molluscs (pelecypods and gastropods) and echinoderms (echinoids and ophiuroids) greatly influenced patterns of total biomass distribution. The ocean quahog, Arctica islandica, was the dominant species in terms of biomass. Total density was dominated by crusta- ceans (amphipods), polychaetous annelids, molluscs (small pelecypods), and echinoderms (ophiuroids). Mean density of mollusca was positively associated with sediment size. Mean biomass and density of crustacea were negatively associated with depth, grain size, and bottom temperature, whereas the same param- eters for the Echinodermata were positively associated with those environmental factors. Three faunal assemblages emerged which were analagous to those described from earlier studies on Georges Bank (sand fauna, silt-sand fauna, muddy-basin fauna). The fauna from the Mud Patch most closely resembled the silty-sand fauna. INTRODUCTION Research on the northeastern U.S. continental shelf has ac- celerated because of a wide variety of human activities ranging from fishing and recreation to transportation, mineral recovery, waste disposal, and oil and gas exploration (Grosslein et al. 1979). These activities are not always com- patible, and managers face important decisions attempting to reconcile diverse uses of this valuable resource—the continen- tal shelf. Before decisions can be made, the resource must be assessed in a variety of ways. Any assessment of the shelf should involve benthic in- vertebrates, an important component of the shelf ecosystem (Mills and Fournier 1979). The benthos are important in their own right and as a measure of the health of the ecosystem, and play a critical role in trophic relationships providing a major source of energy to economically and ecologically important groundfish (Cohen et al.*). In addition, the benthos play a sup- porting role in nutrient exchange, providing a mechanism for flux of nutrients initially trapped in sediments to the water col- umn (Zeitzschel 1980). The purpose of this account is to report on the distribution of biomass and density of macrobenthic in- vertebrates off Martha’s Vineyard. Georges Bank off southern New England has been the scene of intensive fishing activity for over 300 yr (Wigley 1961). The Georges Bank area is bounded on the southwest by Martha’s Contribution No. 13 from the Southern California Ocean Studies Consortium. *Southern California Ocean Studies Consortium, California State University, Long Beach, 1250 Bellflower Blvd., Long Beach, CA 90840. *35 Wilson Road, Woods Hole, MA 02543. “Cohen, E. B., M. D. Grosslein, M. P. Sissenwine, and F. Steimle. 1979. An energy budget of Georges Bank. Workshop on Multispecies Approaches to Fish- eries Management, St. John’s, Newfoundland, 26-30 November 1979, p. 1-37. Na- tional Marine Fisheries Service Woods Hole Lab., Woods Hole, Mass. Vineyard and Nantucket Island. Directly south of Martha’s Vineyard is an area about 80-100 m deep termed the Mud Patch, consisting of fine-grain sediment. This sediment type is relatively rare on the northeastern shelf which normally con- sists of sand. The origin of the Mud Patch is uncertain, but it may be an active site of deposition (Milliman et al.*). If so, then processes controlling deposition of fine-grain sediment must be considered in assessing their effect on benthic biota. Presumably, benthos living in the Mud Patch would be exposed to relatively higher levels of particulate contaminants (trace metals, hydrocarbons) than benthos living on the sur- rounding, relatively dynamic, sand bottoms. Assuming this relationship, bioaccumulation of contaminants by the benthos might provide opportunity for biomagnification through food webs. The trophic relationship between demersal fish and ben- thos is well documented for Georges Bank (Wigley 1965). In the early 1960’s benthic research was conducted on the shelf south of Martha’s Vineyard and Nantucket I. Some of these data were reported (Wigley 1963; Wigley and McIntyre 1964; Wigley and Stinton 1973), but the largest portion was placed in a data file report pending further analysis and syn- thesis (Maurer and Wigley*). The report contains maps of biomass and density distribution. With the advent of gas and oil exploration, together with other diverse activities on Georges Bank, documentation of the benthos in and around the Mud Patch would seem to be of special interest to man- *Milliman, J. D., M. H. Bothner, and C. M. Parmenter. 1980. Seston in New England Shelf and Slope Waters, 1976-1977. In J. M. Aaron (editor), Environ- mental geological studies in the Georges Bank area, United States northeastern Atlantic Outer Continental Shelf, 1975-1977, p. 1-1 to 1-73. Final Report submit- ted to the Bureau of Land Management. U.S.G.S., Woods Hole, Mass. *Maurer, D., and R. L. Wigley. 1981. Distribution of biomass and density of macrobenthic invertebrates on the U.S. Continental Shelf off Martha’s Vineyard. Lab. Ref. Doc. 81-15 (unpubl.), 97 p. Northeast Fisheries Center, NMFS, NOAA, Woods Hole, Mass. agers because of its proximity to the coast and its potential as a depositional sink of contaminants. Although there are a number of review documents dealing with benthos off south- ern New England (Pratt 1973; TRIGOM-PARC’; Wigley and Theroux®:’; Maurer'®), no study has featured the fauna of the Mud Patch. MATERIALS AND METHODS Field and Laboratory Samples were collected 11-20 June 1962 by the National Marine Fisheries Service vessel RV Delaware at 64 stations south of Martha’s Vineyard, Mass. (Fig. 1, Appendix Table 1). Stations were spaced at intervals of 16 km on a grid pattern with eight north-south transects at right angles to the depth contours. At each station (except station 7) two quantitative bottom samples were collected with a Smith-MclIntyre grab. This instrument effectively sampled a 0.1 m? area of bottom to a depth of 10-17 cm. At sea, grab samples for macrobenthic studies were washed through a 1.0 mm mesh screen. Macro- benthos remaining on the screen after washing were removed and preserved in a solution of neutral Formalin. In the laboratory macrobenthos were sorted, identified, counted, and weighed. External moisture was removed from specimens by blotting. Shells, internal skeletons, and exo- skeletons were included in the values expressed as wet weight biomass (g/m_’). Sediment samples were collected with a Dietz-LaFond grab at each station and at two localities equally spaced between stations along the cruise. The locations of sediment samples used in conjunction with biological analyses are depicted in Figure 1. Terminology follows the Wentworth Particle Size Classification (Twenhofel and Tyler 1941), and nomenclature follows the classification of Shepard (1954) and Emery (1960). Determinations were made of median sediment size ($), percent sand, percent silt, percent clay, percent Kjeldahl nitrogen, per- cent organic carbon measured, and carbon/nitrogen ratio of the sediments (Hathaway *'). Standard sieving procedures were used to measure sediment size. Based on sediment analyses, a composite sediment-type map was made (Fig. 2). Appendix Table 1 lists environmental data per station. ’TRIGOM-PARC. 1974. A socio-economic and environmental inventory of the North Atlantic Region, Vol. 1, Book 3, 198 p. Report to the Bureau of Land Man- agement, South Portland, Me. ‘Wigley, R. L., and R. B. Theroux. 1976. Macrobenthic invertebrate fauna of the Middle Atlantic Bight region. Part II: Faunal composition and quantitative distribution. Northeast Fisheries Center, NMFS, Woods Hole, Mass., 395 p. °Wigley, R. L., and R. B. Theroux. Reconnaissance survey of the quantitative distribution of macrobenthic invertebrates in the offshore New England region. Manuscr. in prep. Northeast Fisheries Center, NMFS, Woods Hole, Mass. ‘°Maurer, D. 1982. Review of benthic invertebrates of Georges Bank in relation to gas and oil exploration with emphasis on management implications. Report to Northeast Fisheries Center, Woods Hole, Mass., and Sandy Hook Laboratory, Highlands, N.J., 329 p. “Hathaway, J. C. (editor). 1971. Data file, continental margin program, Atlan- tic Coast of the United States. Vol. 2. Sample collection and analytical data. Ref. No. 71-15, 496 p. Woods Hole Oceanographic Institution, Woods Hole, Mass. Analysis Wet weight and number per taxon by station and gear were punched on cards and a computer listing was prepared. Based on the listing, the average biomass and density per station of major taxa (Amphipoda, Pelecypoda, Asteroidea, etc.) were determined and distribution maps plotted by computer (Maurer and Wigley footnote 6). In some cases, maps were also made of particularly important genera and species. Corre- lation coefficients (R) were computed for average weight and number transformed (loge (NV + 1)) of major taxa in relation to environmental variables. ENVIRONMENTAL SETTING Physiography The Georges Bank area off New England is a submerged northeast extension of the Atlantic Coastal Plain (Aaron'). The Bank, which encompasses about 42,000 km?, is covered by up to 200 m of water. The study area for this account (Fig. 1) lies immediately southwest of the Bank. The study area encompasses about 130 km? and extends across the continental shelf to the upper portion of the con- tinental slope. Bottom topography is moderately smooth. Water depths increase gradually and rather uniformly from shore outward to the shelf break, about 120 m. The average gradient of the continental slope off the Middle Atlantic Bight varies from 2° to 7° (Milliman 1973); beyond the shelf break in the study area, the depth gradient is relatively steep, averaging 4° (Wigley and Stinton 1973). The most distinguishing feature on the shelf break is the number of gullies and canyons that transect the slope (Fig. 1). Sediment Composition Six major sediment types occurred in the study area (Fig. 2). Sand, silty sand, and sandy silt occurred over a large area, whereas gravel-sand, sand-silt-clay, and silt were much less widespread. Sand with some gravel (stations 1, 45, 47) covered more than half the area, mainly in shallow water (0 to 60-80 m) except in the eastern sector and in a narrow (6 km) band paral- lel to and just below the outer periphery of the continental shelf. In shallow water the sands were silt free and occasionally mixed with large quantities of shell (mollusks and echinoderm plates). Admixtures of silt occurred with the sand over most of the remaining area. A large area (80 x 100 km) of fine-grain sediment (Mud Patch) occurred in the southwestern sector (Fig. 2). A relative- ly circular area of sand-silt-clay near its center was surrounded by an inner band of sandy silt which grades to an outer band of silty sand. Illite is normally the most important clay mineral, and organic carbon is higher here than in the surrounding sand (Appendix Table 1). This is the largest known natural area of fine-grain sediment on the Middle Atlantic Shelf. Sediments on the continental slope were dominated by silt and clay. "Aaron, J. M. 1980. A summary of environmental geologic studies in the Geor- ges Bank area, United States northeastern Atlantic outer continental shelf, 1975- 1977. Executive Summary of the Final Report submitted to the Bureau of Land Management. U.S.G.S., Woods Hole, Mass., 22 p. Figure 1.—Location of stations of bottom samples for fauna (o) and sediment (x) off Martha’s Vine- yard, Mass., June 1962. Stations 1, 2, 6, and 7 are from the 1977 BLM study. Figure 2.—Bottom-sediment types off Martha’s Vineyard, Mass., from samples collected at stations (o), June 1962. Wane ee ated Ryd ELE 1000 METERS Dis ~ ae a DP pees Vier 1000 METERS BOTTOM SEDIMENTS IZZAGRAVEL-SAND [==JSANDY SILT EZZ4ASAND SAND-SILT-CLAY EZEJSILTY SAND (1) sitt In the fine-grain sediments southwest of Nantucket I., values of 6.0-10 mg/g organic carbon and 0.8-1.2 mg/g nitrogen were recorded (New England OCS'?). Highest concentrations (15,000 g/l) of seston were measured on the shelf south of Nantucket Shoals during the winter (Milliman et al. footnote 5; Aaron et al. 1980). Physical Oceanography Circulation.—The major pattern of circulation on Georges Bank was described earlier by Bigelow (1927) and Bumpus (1973, 1976) and has been updated by Butman et al. (1980). The data of Butman et al. showed a clockwise circulation pat- tern around the Bank. Flow on the southern flank was con- sistently toward the southwest. West of the Great South Chan- nel and south of Nantucket I., mean flow was measured to the west-northwest along contours of local bathymetry (Dorkins 1980). On the eastern side of Great South Channel, mean flow was consistently toward the north at 5-10 cm/s at all depths (But- man et al. 1980). At a station just south of Great South Chan- nel, there was a westerly flow similar to that along the southern ‘New England OCS Environmental Benchmark. 1978. Draft final report, Vol. V, Sects. G-J. Energy Resources Co., Cambridge, Mass., 224 p. flank. On the western side of Great South Channel, little net flow was measured. Great South Channel has been historically considered a ma- jor exit for Gulf of Maine waters (Schlitz in press). However, recent water current measurements indicate that mean flow within the channel is directed mainly toward the Gulf of Maine (Butman et al. 1980). Schlitz et al. (1977) showed that a perma- nent front exists across the channel from Nantucket Shoals toward Georges Bank, separating the Gulf of Maine from the shelf to the south. According to Schlitz {in press), a small amount of water flows southward through Nantucket Shoals, but this probably does not contribute a significant volume. Bottom temperature.—When the study was conducted in June 1962, a cell of cold bottom water (6.1°-6.9°C) extended in an east-west band from the New York Bight eastward to long. 69°30'W (Fig. 3). This cell occurred at depths of 40-80 m, roughly the mid-shelf region. The cold cell was bounded on the north by higher coastal water temperatures (< 12°C) and on the south by values of 10-12°C near the shelf break. Long- term (1940-66) annual maximum and minimum bottom-water temperatures near the shelf break were 16°-17°C and 1°-2°C, respectively (Colton and Stoddard 1973). However, the annual range here is normally 2°C. Offshore shelf waters, particularly in shallow portions, may range from 3°C in February-March to 14°C in September-November (Wigley and Stinton 1973). 100 X : 40°. aS FF ° INE NEA 22 .SS8 Rear Figure 3.—Bottom temperature (°C) off Martha’s 200 Pea eis Oe al alm i) NE, =I Sm nl a a= Vineyard, Mass., from measurements taken at sta- 500 = 5 Dini bas saree Jia tions (0), June 1962. oy eno orev oe WEL oe c 1000 METERS RESULTS Faunal Composition A total 214 taxa were identified (Appendix Table 2). Since some taxa were identified only to phylum (Porifera, Nemertea) and since some taxa identified to genus may include several species, the total number of taxa is conservative. Of the number of taxa, v24.3% were molluscs, 27.1% arthropods, 24.3% polychaetous annelids, 10.7% echinoderms, and 13.6% miscellaneous taxa, including sipunculids, coelenterates, nemerteans, ectoprocts, ascidiaceans, pogonophorans, and hemichordates. Biomass (wet weight g/m’) and density (no./m?) are presented in Appendix Tables 3 and 4. About 190,000 individuals and 15,500 g of specimens were collected. Total Faunal Biomass and Density Biomass and density were distributed among the major taxa as follows: Biomass Density (g/m?) (no./m?) Annelida 10.9% 20.3% Mollusca 56.7% 5.9% Crustacea 4.1% 62.5% Echinodermata 21.3% 7.0% Misc. taxa 7.0% 4.3% NANTUCKET e D oo 1000 METERS 1-99.99/m? VA \00— 999. 99/m2 WY, > 1,000g/m2 Average biomass and density per station for total fauna were 245.7 g/m? and 3,008 individuals/m’, respectively. Biomass in the shallow central stations generally ranged from 100 to 999 g/m? (Fig. 4). Biomass decreased to a range of 1-100 g/m? in slightly deeper water and then increased again to 100-999 g/m? between 60 and 100 m. Below 100 m on the western side of the study area, biomass again decreased to <100 g/m’, but biomass was still in the 100-999 g/m? range on the eastern side to almost 200 m. Molluscs, polychaetes, and shallow-water echinoderms contributed heavily to the highest values of biomass. The density pattern for total fauna was more irregular than for biomass (Fig. 5). A range of 1,000 to >3,000 m? occurred in shallow water. A major portion of the area contained den- sities of 1,000-2,999/m?. Lowest densities (1-999/m?) were recorded below “100 m. Crustacea, primarily amphipods, and polychaetes contributed greatly to the highest density values. Stations were grouped into ranges of depth, temperature, median sediment size, and silt-clay (Maurer and Wigley foot- note 6). Mean biomass and density were calculated for those ranges and summarized. A depth range of 40-80 m, tempera- ture 6.0°-7.9°C, $ 2.0-3.99, and 0.6% silt-clay comprised the main distribution of biomass (Fig. 6). The principal distribu- tion of density was accounted for by a depth range of 0-100 m, temperature 6.0°-9.9°C, $ 1.0-3.99, and 0-40% silt-clay. Mean biomass and density declined with depth but the correlations were low (R = —0.20). Figure 4.—Quantitative distribution of biomass (g/m?’) for all macrobenthic invertebrates combined off Martha’s Vineyard, Mass., from samples collec- ted at stations (0), June 1962. Figure 5.—Quantitative distribution of individuals (no./m’) for all macrobenthic invertebrates com- bined off Martha’s Vineyard, Mass., from samples collected at stations (0), June 1962. 1000 METERS Polychaetous Annelids Many species of polychaetes were identified (Appendix Table 2). The number of species is a conservative estimate because recent work with a 0.5 mm mesh sieve has yielded higher numbers of species (Maurer and Leathem 1980). Mean biomass of polychaetes ranged from 0.1-39 g/m? throughout most of the area to >100 g/m? at a few stations (Maurer and Wigley footnote 6). A biomass range of 40-99 g/m? was measured directly south of Nantucket I. and Nantucket Shoals. Polychaetes were more evenly distributed throughout en- vironmental ranges than any of the major taxa (Fig. 6). Mean biomass and density were negatively associated with tempera- ture (R = —0.39, R = —0.39). A number of polychaete species showed marked differential distribution. Maldanids and Scalibregma inflatum occurred in shallow water and sand. The latter species showed the same relationship over Georges Bank (Maurer and Leathem 1980). In contrast, terebellids and Sternapis scutata occurred in deeper water with fine-grain sediment. Mollusca In terms of mean biomass and density, Mollusca contained Scaphopoda (0.1 and 0.6%, respectively), Gastropoda (4.8 and 15.2%), and Pelecypoda (95.1 and 84.2%). Biomass ranged from 0.1 to >100 g/m? with the highest values directly south of Nantucket Shoals and southwest of Nantucket I. (Maurer and Wigley footnote 6). A depth range of 20-80 m, 6.0°-9.9°C,$ 2.0-3.9, and 0-20% silt-clay comprised the main distribution Ne x Y — aN a Dy oA J ¢ cv Dene ote te NEG COIN — 999.9/m2 V7, >3,000/m2 Ke? of biomass (Fig. 6). Density distribution of Mollusca was very different from biomass distribution because the most numerous molluscan species were smaller than the main contributors to biomass. Densities ranging from 100 to >400/m? occurred in the central part of the study area between 60 and 100 m. This central area was surrounded by densities ranging from 1 to 99/m?. The main distribution of density for molluscs encom- passed 80-100 m, 7.0°-10.9°C, $ 3.0-5.9, and 20-100% silt-clay (Fig. 6). Mean density of mollusca was positively associated with grain size (R = 0.44). Scaphopoda.—The scaphopods Cadulus pandionis, C. ver- rilli, and Dentalium occidentale were sampled only at deep sta- tions (Maurer and Wigley footnote 6). Scaphopods occurred in relatively low biomass (0.05-0.08 g/m?) and density (3.3- 7.5/m?) in water deeper than 200 m. Their distribution mainly encompassed 10.0°-12°C, $ 4.0-5.9, and 40-80% silt-clay (Fig. 6). Gastropoda.—Although biomass and density of gastropods ranged from 0.1 to >100 g/m? and from 1 to >400/m’, re- spectively, there were only a few stations that contained > 100 g/m? or >400/m? (Maurer and Wigley footnote 6). Biomass and density patterns of gastropods were very different (Fig. 6). Biomass patterns were mainly influenced by large-to-inter- mediate size, shallow water, sand-dwelling taxa (Colus, Nassarius, Buccinum, Neptunea, Lunatia), and density was influenced by smaller, deep-water, silt-sand and sandy-silt species (A/lvania carinata). There was a significant negative association between density of gastropods and temperature (R = —0.37). Porifera Coelenterata Nemertea Annelida Pogonophora Sipunculida Mollusca Gastropoda Pelecypoda Scaphapoda Crustacea Cumacea lsopoda Amphipoda Decapoda Echinodermata Holothuroidea Echinoidea Ophiuroidea Asteroidea Ectoprocta Enteropneusta Ascidiacea Miscellaneous Total Depth (m) 2582536 27 3 5 4 ISM 228) 20 29 43 46 23 30 40 728 721 793 456 359 618 698 771 esis ie | Sl 44 4 2. 4 20 19 3.4 204 223 247 162 221 432 146 30 44 28 65 204 22i 217 18 192 367 137 100 g/m’. Highest values were recorded directly south of Nantucket Shoals and Nantucket I. (Maurer and Wigley foot- note 6). A range of 20-80 m, 6.0°-9.9°C, $ 2.0-3.9, and 0-20% silt-clay comprised the main biomass distribution (Fig. 6). Density patterns differed from biomass patterns because of size differences. A range of 40-200 m, 6.0°-10.9°C, $ 3.0-5.9, and 20-100% silt-clay comprised the main density distribution. There was a significant negative association between pelecy- pod biomass and temperature and depth, and a positive asso- ciation between density and sediment size. Species of bivalves showed marked differential distribution. Arctica islandica and Cerastoderma pinnulatum mainly occurred at depths shal- lower than 80 m and in sand with <20% silt-clay. In contrast, Cuspidaria striata, C. perrostrata, and Bathyarca pectunculoi- des occurred almost exclusively at depths > 150 m in sediment wit.. >50% silt-clay. Molluscan biomass distribution was chiefly influenced by the distribution of the pelecypod A. islandica. Small species of bivalves contributed significantly to the density pattern of Mollusca. Crustacea In terms of biomass and density, the Crustacea contained Decapoda (5.7 and 0.2%, respectively), Cumacea (1.3 and 2.1%), Isopoda (5.0 and 1.0%), and Amphipoda (88 and 96.7%). Mean biomass and density of crustaceans ranged from 0.1 to >50 g/m? and from | to >1,000/m?. A depth range of 20-80 m, 6.0°-9.9°C, $ 1.0-3.9, and 0-20% silt-clay comprised the main distribution of crustacean biomass and density (Fig. 6). Mean biomass and density were negatively associated with depth (R = —0.53, R = —0.56), sediment size (R = —0.31, R = —0.30), and bottom temperature (R = —0.48, R = —0.50). Decapoda.—Mean biomass and density of decapods ranged from 0.1 to 49 g/m? and from 10 to 25/m?. Their distribution was sporadic (Maurer and Wigley footnote 6), but grab samples are not the most effective way to collect large crusta- ceans. Crangon septemspinosa, Pagurus pubescens, Cancer spp., and Pandalus spp. were collected frequently. Crangon septemspinosa occurred at depths <20 m to >200 m. In con- trast, Hyas sp., Euprognatha sp., Munida sp., and Geryon sp. were generally collected deeper than 100 m. Cumacea.—Mean biomass and density of cumaceans ranged from 0.1 to 1.0 g/m? and from 1.0 to >400/m?. Cumaceans occurred widely and were relatively evenly spread, occurring mainly at lower densities of 1-100/m? (Maurer and Wigley footnote 6). Characteristic species included Diastylis polita, D. quadrispinosa, Eudorella emarginata, Leptostyllis sp., Eu- dorellopsis sp., and Leptocuma sp. A depth range of 20 m, 7.0°-8.9°C, $ 3.0-3.9, and 0-10% silt-clay comprised the main biomass distribution, whereas the main density distribution was most accurately contained in a depth range of 20-40 m, 6.0°-8.9°C, $ 2.0-3.9, and 0-20% silt-clay (Fig. 6). Mean bio- mass and density of cumaceans were negatively associated with depth (R = —0.36, R = —0.52), and density with tempera- ture(R = —0.29). Isopoda.—Mean biomass and density ranged from 0.1 to 10 g/m? and from | to >100/m?. The greatest numbers of isopods occurred between 40 and 60 m throughout most of the study area, but between 60 and 80 m directly south of Nantucket Shoals (Maurer and Wigley footnote 6). Biomass and density patterns were essentially the same. Characteristic species were Cirolana polita, Chiridotea tuftsi, Ptilanthura tenuis, Edotea triloba, and Calathura sp. Mean density of isopods was negatively associated with depth (R = —0.38), sediment size (R = —0.38), and temperature (R = —0.26). Amphipoda.—Mean biomass and density of amphipods ranged from 0.1 to >50 g/m? and from | to >1,000/m?. The shallow water portion mostly contained densities > 1,000/m?. There was a sharp decline in biomass and density below 100 m (Maurer and Wigley footnote 6). Characteristic amphipod taxa were Leptocheirus pinguis, Unciola irrorata, Caprella spp., Coropium spp., ampelescids (including Ampelisca com- pressa, A. macrocephala, and Byblis serrata), phoxocephalids, photids, and haustoriids. A depth range of 20-80 m, 6.0°-9.9°C, $1.0-3.9, and 0-40% silt-clay comprised the main distribution of biomass. These ranges were the same for densi- ty except for depth with a range of 20-100 m (Fig. 6). The dis- tribution of amphipod biomass and density greatly influenced the same patterns for combined Crustacea. Mean biomass and density were negatively associated with depth (R = —0.53, R = —0.54), sediment size (R = —0.31, R = —0.31), and temperature (R = —0.49, R = —0.51). Echinodermata In terms of biomass and density, echinoderms contained Ophiuroidea (21.7 and 78.6%, respectively), Holothuroidea (52.9 and 5.9%), Echinoidea (19.7 and 13.8%), and Asteroi- dea (5.7 and 1.7%). Mean biomass and density ranged from 0.1 to >100 g/m? and from 1 to >1,000/m?. Biomass and density patterns of echinoderms were generally dissimilar (Fig. 6). Mean biomass and density were positively associated with depth (R = 0.27, R = 0.25), sediment size (R = 0.44, R = 0.36), and temperature (R = 0.33, R = 0.42). Ophiuroidea.—Mean biomass and density of ophiuroids ranged from 0.1 to >100 g/m’ and from 1 to >500/m?. However, in general, biomass values of 0.1-49 g/m? were most common, as were densities of 1 to >500/m?’. Characteristic species were Amphilimna olivacea, Amphioplus abditus, A. Sragilis, Axiognathus squamatus, and Ophiura sarsi. In terms of biomass, ophiuroids were mainly restricted to depths >80 m, 10.0°-12.9°C, $ 4.0-5.9, and 20-100% silt-clay, and for density 7.0°-12.9°C and > 3.0-5.9 (Fig. 6). Mean biomass and density were positively associated with depth (R = 0.44, R = 0.40), sediment size (R = 0.36, R = 0.40), and temperature (R = 0.54, R = 0.48). Holothuroidea.—Mean biomass and density of holothur- ians ranged from 0.1 to >100 g/m? and from 1 to 399/m?. Holothurians were almost exclusively collected in the deeper portion of the study area below 80 m (Maurer and Wigley footnote 6). Characteristic species were Synapta sp., Astichopus sp., Molpadia sp., and Havelockia scabra. A depth range of 40-100 m, 6.0°-7.9°C, $ 3.0-4.9, and 20-80% silt-clay comprised the main biomass distribution, whereas a depth range of 80-100 m and 6.0°-10.9°C comprised the main density distribution (Fig. 6). Echinoidea.—Mean biomass and density of echinoids ranged from 0.1 to >100 g/m? and 0.1 to 399/m?. The distribution of echinoids was primarily influenced by the distribution of two species (Echinarachnius parma and Schizaster fragilis) (Maurer and Wigley footnote 6). Echinarachnius parma influenced echinoid biomass and density in shallower water, sand, and low percent silt-clay, and S. fragilis was influenced in deeper water, sandy-silt, and high silt-clay. Asteroidea.—Mean biomass and density of asteroids ranged from 0.1 to >50 g/m? and from 1-49/m?. Their distribution was sporadic (Maurer and Wigley footnote 6), but grab sam- ples are not the most effective way to collect large asteroids. Characteristic species were Asterias vulgaris, Leptasterias sp., Porania sp., Henricia sp., Astropecten americanus, and Astro- pecten sp. Asterias vulgaris represented a shallow-water spe- cies, and Astropecten americanus was a deeper water species. Mean biomass and density were positively associated with sedi- ment size (R = 0.32, R = 0.26). Mud Patch Fauna Earlier in this report, the southwest quadrant of the Martha’s Vineyard-Nantucket Shoals area was referred to as the Mud Patch (Fig. 2). The following stations generally com- prised the Mud Patch: Stations 23-25, 33-36, 38-42, 49-53, and 57-59. Average biomass for these stations was 123.7 g/m’, and density was 1,536.2/m?, which is markedly lower than respec- tive values of 245.7 g/m? and 3,008/m? for the entire Martha’s Vineyard-Nantucket Shoals area. Based on two BLM stations (1 and 2) in the shallow water and clean sand of Martha’s Vineyard-Nantucket Shoals area (Fig. 1), the dominant polychaete species (in abundance and frequency of occurrence) were Exogone hebes, Spiophanes bombyx, Parapionosyllis longicirrata, Sphaerosyllis erinaceus, Owenia fusiformis, Scalibregma inflatum, Aricidea catherinae, and Spio pettiboneae (Maurer and Leathem"*). The Mud Patch was characterized by anthozoans, small-to- medium size bivalves (Bathyarca pectunculoides, Nuculana acuta, Yolida sapotilla, Nucula spp.), small-to-medium size decapods (Catapagurus sharreri, Hyas coarctatus, Euprognatha rastellifera, Munida iris), and a variety of generally small echi- noderms (Amphilimna olivacea, Amphioplus macilentus, Amphiura otteri, Havelockia scabra, Schizaster fragilis). Based on two BLM stations (6 and 7) in sediment containing 32-37% silt-clay near the Mud Patch (Maurer and Leathem footnote 14), dominant species were Paraonis gracilis, Cossura longocirrata, Ninoe nigripes, Aricidea suecica, Nephtys incisa, Tharyx annulosus, Terebellides stroemii, Maldanidae, and Cirratulidae in the deeper water and silty-sand and sandy-silt. Robert Reid’* reported high numbers of the amphipod Ampe- lisca agassizi from the Mud Patch in some 1980 collections to- gether with other species cited above. This fauna contains a high proportion of selective and non-selective deposit feeders, both surface and buried, representing a typical soft-bottom community. Echinoderms were particularly important as con- “Maurer, D., and W. Leathem. 1980. Ecological distribution of polychaetous annelids of Georges Bank. CMS-1-80. College of Marine Studies, Univ. of Dela- ware, Lewes, Del., 181 p. *Robert Reid, Northeast Fisheries Center Sandy Hook Laboratory, National Marine Fisheries Service, NOAA, Highlands, NJ 07732, pers. commun. 1981. tributors to biomass in the Mud Patch. Because of their gener- al mode of feeding, the biota of the Mud Patch would have considerable potential for ingesting pollutants associated with deposition of fine-grain sediment. DISCUSSION Ecological Relationships A summary of mean biomass and number of individuals of major taxa in relation to depth, temperature, median sediment size, and silt-clay is presented in Figure 6. Also contained in Figure 6 age the range (biomass and number of individuals) of each taxon per environmental variable, together with values of average biomass and density plotted for a specific mid-range. Depth.—Biomass and density of polychaetes off Martha’s Vineyard were not significantly associated with depth. However, for the Georges Bank area, the density of polychaetes increased significantly with depth down to +80 m with some indication of reduction in biomass with depth (Maurer and Leathem footnote 14). Off Martha’s Vineyard, biomass and density of molluscs combined were not significantly associated with depth. However, the density of scaphopods increased sig- nificantly with increasing depth and the biomass of pelecypods decreased with depth. Biomass pattern of pelecypods with depth was dominated by the distribution of A. islandica rang- ing between 40 and 60 m. For Georges Bank, density of com- bined molluscs increased with water depth (Maurer'®). For combined crustaceans off Martha’s Vineyard, mean biomass decreased with depth. Mean density of cumaceans and isopods was negatively associated with depth. Mean bio- mass of amphipods decreased with increasing depth. Density pattern of combined crustaceans was dominated by the distri- bution of amphipods, whereas biomass pattern of combined crustaceans was dominated by decapods. For Georges Bank, mean biomass of amphipods decreased in deeper water (Maurer footnote 16). For combined echinoderms off Martha’s Vineyard, mean biomass and density increased with depth. In another study, the bathymetric distribution of echinoderms off the northern Oregon coast was not as uniform, as low wet weights were ob- tained from depths of 86-139 m and 1,189-1,234 m (Alton 1972). Mean biomass and density of ophiuroids increased with depth off Martha’s Vineyard. For Georges Bank, mean densi- ty of combined echinoderms showed no significant relation- ship with depth, but biomass decreased significantly in deeper water (Maurer footnote 16). These relationships reflected the fact that although density was relatively regular throughout the depth range, larger species (sea stars and echinoids) oc- curred in relatively shallow water, with smaller species (brittle stars) in deeper water. Mean biomass and number of individuals of combined taxa were high between 0 and 100 m depths. Biomass and density decreased with increasing depth, although these relationships were not statistically significant. Off southern New England, there was a marked reduction in density and biomass with '*Maurer, D. 1982. Review of benthic invertebrates of Georges Bank in relation to gas and oil exploration with emphasis on management implications. Report to Northeast Fisheries Center, NMFS, Woods Hole, Mass., and Sandy Hook Lab- oratory, Highlands, N.J., 329 p. depth (Wigley and Theroux 1981). Total biomass was domi- nated by molluscs (pelecypods and gastropods) and echino- derms (echinoids and ophiuroids), and density pattern was dominated by amphipods, polychaetes, small bivalves, and ophiuroids. According to Parsons et al. (1977), a number of studies have demonstrated decreased macrofaunal numbers and biomass with increased water depth. Abrupt faunal dis- continuities tended to occur at 100-300 m. These changes gen- erally corresponded to the vertical distribution of other envi- ronmental factors, such as organic carbon, nitrogen, and total particulates which decrease rapidly with increasing depth. Depth-related decreases in macrobenthos are more closely linked to suspended living biomass than to total particulate matter (Parsons et al. 1977). Parsons et al. (1977) cited examples of higher mean biomass in regions of higher phytoplankton production. This relation- ship suggests that benthic biomass in inshore coastal areas may be strongly influenced by sedimentation of organic matter pro- duced during a bloom. This process undoubtedly contributes to macrobenthos at shallow depths of continental shelves. The shallow depths of Georges Bank and Nantucket Shoals fit these conditions. Data from other studies on benthic biomass with depth are presented in Table 1. Mean biomass was generally highest in the 50-99 m stratum. Moreover, mean biomass was higher off New England than the New York Bight. Off the Scotian Shelf, wet weight biomass was 24 g/m? at 0-90 m depth and 22.1 g/m? at 90-180 m (Mills 1980). The inverse relationship between ben- thic biomass and depth is probably more related to the geo- graphic position of zones of primary productivity and sedi- mentation on shelves rather than to absolute values of depth per se. This relationship has been recognized for some time (Rowe 1971; Sokolova 1972). Another aspect related to depth involves substratum stabili- ty. Sediments in shallow coastal waters and the inner shelf are subject to vigorous current action. Wave base to 80-85 m depth is not uncommon during the winter on Georges Bank (Aaron et al. 1980). Accordingly during times of high energy flow, sediment stability decreases, impeding colonization by many infaunal organisms. Specialized species such as haus- toriid amphipods and rapidly burrowing bivalves commonly dominate these sites. Nantucket Shoals represents an area of high seasonal sediment instability. Temperature.—Although there were some quantitative rela- tionships between biomass and density of various taxa and bottom temperature, the presence of a cold-water cell bounded by warmer water on the north and south made it difficult to in- terpret distribution according to temperature (Fig. 3). In this case, there was a shallow-water (0-50 m) and deep-water (80- 100 m) zone both containing water ranging from 7° to 12°C. Although the maximum biomass of certain taxa (Ophiuroidea) was associated with shallow water and deep water (Maurer and Wigley footnote 6), their maximum biomass was similar ac- cording to temperature (11.0°-12.9°C). The Coelenterata, another deep-water taxon, had its highest biomass in 11.0°-11.9°C (Maurer and Wigley footnote 6), temperatures normally associated with depths of about 40 m for this time of year. Thus it is important to bear in mind the relative position of the cold-water cell in relation to depth when comparing dis- tribution according to temperature. Off Atlantic City, N.J., Boesch et al.'’ concluded that tem- perature was the principal hydrographic factor affecting macrobenthic distribution. Temperatures were more variable on the inner and central shelf and more constant on the outer shelf. Differences in temperature regime were probably the prime cause of the sharper faunal change at the outer shelf/ shelf-break transition. Sediment end related environmental variables.—Off Martha’s Vineyard, the pattern of biomass and number of in- dividuals were relatively even throughout a range of median sediment size and percent silt-clay (Fig. 6). For Georges Bank, density of infauna increased significantly with percent gravel (Maurer and Leathem footnote 14). In addition, there were some significant relationships between the density of several dominant polychaete species and sediment parameters (percent silt-clay, percent silt, percent carbon, percent nitrogen, micro- bial biomass, and bacterial biomass) (Maurer and Leathem footnote 14). Off Martha’s Vineyard, polychaetes were ubiquitous in re- gard to sediment type and were major contributors to both average density and biomass of all benthic organisms in each sediment type (Maurer and Wigley footnote 6). This apparent lack of correlation with a sediment type may partly be due to differences in sieve size, wherein some of the smaller taxa known to occur abundantly on the shelf were probably missed with a 1.0 mm mesh net. In a related study off southern New England, greatest amounts of polychaetes were found in shell and sand-gravel (750 and 555/m’, respectively), somewhat lesser amounts in sand, silty sand, gravel, and silt (433, 331, 289, and 118/m7?), and lowest (23 and 9/m*) in sand-shell and clay (Wig- "Boesch, D., J. N. Kraeuter, and D. K. Serafy. 1977. Benthic ecological studies: Megabenthos and macrobenthos. /n Middle Atlantic Outer Continental Shelf En- vironmental Studies, Chap. 6, 111 p. Draft Rep. to Bureau of Land Management. Table 1.—Comparison of wet-weight biomass (g/m’) of macrobenthos (Annelida, Mol- lusea, Crustacea, Echinodermata) in relation to bathymetric stratum at locations off the U.S. northeastern Atlantic coast. Martha’s Vineyard/ _ New York Bight? Nantucket Shoals \(1.0mm_ (0.5 mm Southern Depth Georges Bank' New England? (m) (0.5 mm sieve) (1.0 mm sieve) (1.0 mm sieve) sieve) sieve) 25-49 140.6 308.6 ; 230.4 121.0 78.0 50-99 460.0 230.1 314.3 156.4 96.6 100-199 24.2 60.4 75.6 Pe?) 33.2 ‘Maurer, see text footnote 16. *Wigley and Theroux, see text footnote 8. *Boesch et al., see text footnote 17. 10 ley and Theroux 1981). Wet-weight biomass values between 20 and 30 g/m? occurred in four sediment types: Shell, silty sand, gravel, and sand in order of decreasing amounts. Silt and sand- gravel contained 7 and 116 polychaetes/m?, respectively, and sand-shell and clay had the smallest biomass with 1.7 and 0.45 g/m*. In general, there were no correlations between density of annelids and sediment organic carbon for the southern New England area of the New York Bight. The highest biomass val- ues (45.4 and 37.4 g/m*) occurred in sediment with 1.5-1.9% and 2.0-2.9% organic content, respectively. Off Martha’s Vineyard, mean density of combined Mol- lusca was positively associated with sediment size ($) as was that of pelecypods. For Georges Bank, mean density of com- bined molluscs increased with percent carbon in the sediment (Maurer footnote 16). Franz (1976) reported three molluscan- sediment groups in northeastern Long Island Sound. One group consisted of very fine sand and contained species similar to that of the Mud Patch. A second and third group consisted of medium sand and coarse sand and contained molluscan spe- cies very similar to the sand bottom off Martha’s Vineyard. Based on mean grain diameter, sorting, silt-clay content, and fauna, Driscoll and Brandon (1973) identified four facies in Buzzards Bay. In addition, the density of particular mol- luscan-sediment relationships emerged with certain feeding types. Similar relationships have been reported elsewhere (Maurer 1967a, b). Because of the variety of feeding types in many major taxa, it is difficult to correlate major taxa with sedimentary variables. This exercise is most accurately accom- plished at the species level. Mean biomass and density of combined crustaceans were negatively associated with sediment size off Martha’s Vine- yard. This pattern was primarily influenced by amphipods and, to a lesser extent, isopods. For Georges Bank, mean den- sity and biomass of amphipods were positively associated with percent sand, and mean biomass declined with percent gravel (Maurer footnote 16). The number of amphipod species in- creased significantly with percent sand and decreased with per- cent gravel, percent silt, percent silt-clay, percent carbon, and percent nitrogen. In Long Island Sound, there was a strong correlation in summer between Shannon-Weaver diversity of benthic amphipods and sediment texture, with diversity in- creasing due to decreasing species dominance, and, most im- portantly, to increasing species richness as sediments became coarser (Biernbaum 1979). Increased sediment instability caused by winter storms resulted in marked diversity decrease. The response to seasonal sediment stability by the benthic biota, including amphipods, must be a critical feature influenc- ing recruitment, maintenance, and production on shallow por- tions of Georges Bank, including Nantucket Shoals in this study area. Mean biomass and density of combined echinoderms were positively associated with sediment type off Martha’s Vine- yard. This relationship was recorded for ophiuroids and holo- thurians. According to Tyler and Banner (1977) there was a significant relationship off the Bristol Channel between the density of adult dominant ophiuroids and percent fine materi- al with a weaker relationship between density and organic mat- ter in the sediment. They concluded that distribution of both larvae and adults correlated with the energy distribution of the hydrodynamic regime. In view of the hydrodynamic regime off Martha’s Vineyard influencing, on the one hand, deposition in the Mud Patch, and, on the other hand, extensive scouring on 11 Nantucket Shoals, their findings might be applied to echino- derms and the entire benthic community in the study area. The association between benthic animals and sediment is not a simple causal relationship (Rhoads 1974). Sediment composi- tion and associated physical properties (grain size, sorting, porosity, mechanical strength) are primarily controlled by geo- logic processes. In turn, geologic and physical oceanographic processes exert considerable control over chemical properties of sediment (nutrients, oxygen tension, geochemistry). Finally, chemical properties catalyze and interact with biological prop- erties of sediment (algal sheaths, feces, organic film, bacterial and fungal slime). Because of these properties, sediment pro- vides a substratum for colonization, a medium in which reside temporarily or permanently, material for tube and burrow construction, and a source of nutrition. Thus quantitative rela- tionships between benthos and sediment parameters deserve attention; however, because of the varied histories and origins of the sediment parameters, the relationships are not always immediately obvious in terms of their ecological significance. Without quantitative chemical measures of sediment proper- ties (Johnson 1974), sediment might be considered an integra- tive environmental factor to which the benthos are responding. Faunal assemblage Since identification to species level was not always possible, it was not feasible to quantitatively define communities in the study area. Accordingly, the less formal term ‘‘assemblage’’ was used here to designate a recurring group of organisms liv- ing within broadly defined and repetitive environmental condi- tions. Based on studies in the Gulf of Maine and Georges Bank, four major benthic assemblages were tentatively out- lined (Wigley'*): Sand fauna, silty-sand fauna, gravel fauna, and muddy basin. Pratt (1973) elaborated on Wigley’s scheme and suggested that these assemblages extend along the Middle Atlantic Bight. Characteristic species were recommended for the sand assemblage off the Delmarva Peninsula (Maurer et al. 1976) which confirmed Wigley’s (1968) and Pratt’s (1973) pro- jections. Examination of sediment data (Appendix Table 1), species list (Appendix Table 2), and the distribution maps presented here indicates that almost half the study area con- tained the sand fauna (Echinarachnius parma, Crangon septemspinosa, Chiridotea tuftsi, Pagurus acadianus, Lep- tocuma minor, Haustoriidae, Phoxocephalus holbolli, Para- Phoxus sp., Lunatia heros, Nassarius trivittatus, Spisula soli- dissima, Molgula spp.). The fauna of the southwestern quadrat and south central portion of this study—the Mud Patch—compares well with the silty-sand fauna recognized earlier by Wigley (1968) from other areas on Georges Bank. The silty-sand bottom and Mud Patch both contained Havelockia scabra, ampeliscids, Di- chelopandalus leptocerus, Diastylis spp., Edotea triloba, Scalibregma inflatum, Nephtys incisa, Cerianthus, Nucula spp., Nuculana sp., Amphioplus spp., and Amphilimna oliva- cea. A muddy-basin fauna between fishing banks was also identified earlier (Wigley 1968). The deeper stations in this study (7-10, 21, 22, 37, 38, 52, 53, 56, 57), contained species that are in common with the muddy-basin fauna. For example, "*Wigley, R. L. 1958. Bottom ecology. Jn Annual Report, U.S.D.I., Bur. Comm. Fish., Woods Hole Laboratory, Woods Hole, Mass., p. 55-58. Schizaster fragilis, Ophiura sarsi, Ophiura robusta, Amphiura otteri, Cadulus spp., Dentalium sp., Sternaspis scutata, Am- phitrite sp., Onuphis spp., and Leanira sp. were characteristic of muddy-basin fauna and the deep stations off Martha’s Vineyard. In the most comprehensive benthic survey of the U.S. Mid- dle Atlantic Shelf, five faunal zones were recognized (Boesch et al. footnote 17). Faunal changes were mainly gradual rather than abrupt. The faunal zones included: Inner shelf (to 30 m), central shelf (38-50 m), outer shelf (50-100 m), shelf break (100-200 m), and continental slope (>200 m). According to Boesch et al. (footnote 17), the inner and central shelf assem- blages were relatively similar, and outer shelf assemblages con- tained both inshore and offshore species overlapping in distri- bution. In contrast, shelf break and continental slope assemblages were more discrete. Comparison of the faunal assemblages of the Martha’s Vineyard study with Boesch et al. (footnote 17) is difficult for several reasons. The study by Boesch et al. used a 0.5 mm sieve, and their quantitative (cluster analysis) determination of species and site groups emphasized polychaetes and peracarid crustaceans which were missed or deemphasized in earlier studies with coarser sieves. Another difficulty lies in the presence of the Mud Patch off Martha’s Vineyard. Transects from the east side of the study area (Fig. 2) would be more comparable to transects studied by Boesch et al. (footnote 17). The Mud Patch affords the oppor- tunity for colonization in shallower depths by species normally encountered in mud bottoms at deeper depths further out on the central and outer shelf. Coincidental with this expansion into shallower water is the response of deeper dwelling mud- bottom species to a different temperature regime. Species con- sidered characteristic of a zone normally <11°C (Fig. 3) and with a smaller seasonal range would be colonizing a site with a wider temperature fluctuation. It might be expected that the Mud Patch would consist of a fauna containing both inner and outer shelf components. Based on bathymetry and sediment type, the fauna off Mar- tha’s Vineyard could be conveniently arranged into the faunal zones proposed by Boesch et al. (footnote 17). The added complication of the Mud Patch must also be considered. These qualitative comparisons are primarily offered as suggestions for testing rather than as formal community designations. ACKNOWLEDGMENTS This account was supported in part by a contract with the National Marine Fisheries Service Ocean Pulse Program, and was monitored by John Pearce who provided considerable support and encouragement. LITERATURE CITED AARON, J. M., B. BUTMAN, M. H. BOTHNER, and R. E. SYLVESTER. 1980. Environmental conditions relating to potential geologic hazards. U.S. Northeastern Atlantic Continental Margin. U.S.G.S. Misc. Field Studies Map MF-1193, 3 sheets. ALTON, M.S. 1972. Bathymetric distribution of the echinoderms off the northern Oregon coast. Jn A. T. Pruter and D. L. Alverson (editors), The Colum- bia River Estuary and adjacent ocean waters: Bioenvironmental studies, p. 475-537. Univ. Wash. Press, Seattle. 12 BIERNBAUM, C. K. 1979. Influence of sedimentary factors on the distribution of benthic amphipods of Fishers Island Sound, Connecticut. J. Exp. Mar. Biol. Ecol. 38:201-223. BIGELOW, H. B. 1927. Physical oceanography of the Gulf of Maine. Bull. 40:511-1027. BUMPUS, D. F. 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Distribution of macroscopic remains of recent animals from marine sediments off Massachusetts. Fish. Bull., U.S. 71:1-40. WIGLEY, R. L., and R. B. THEROUX. 1981. Macrobenthic invertebrate fauna of the Middle Atlantic Bight Region: Faunal composition and quantitative distribution. U.S.G.S. Prof. Pap. 529, Chap. N1-198. ZEITZSCHEL, B. 1980. Sediment-water interactions in nutrient dynamics. and B. C. Coull (editors), Marine benthic dynamics, p. 195-218. W. Baruch Symp. Mar. Sci., No. 11. Univ. S.C. Press, Columbia. Science Under- In K. R. Tenore Belle Appendix Table 1. Station Location and Environmental Variables off Martha's Vineyard, Massachusetts. Bottom Sediment Parameters Station Station location Water Water Median Composition Nitrogen Organic Carbon Lat. (N) Long. (W) depth temperature Type diameter Sand Silt Clay (Kjeldahl) carbon Nitrogen m °c o % % % % % 1 40°58" 69°30! 46 8.6 Sand- -- -- -- = -- -- -- gravel 2 40°51' 69°31! 46 8.6 Sand 1.00 100.0 0.0 0.0 0.003 0.050 17.0 3 40°40! 69°31! 51 Wot) Sand 2.35 100.0 0.0 0.0 0.010 0.11 Ten 4 40°30! 69°29! 62 7.1 Sand 2.54 100.0 0.0 0.0 0.019 0.13 6.9 BA oul 69°30! 76 9.4 Sand 3.47 SINOm2e7 IGS 0.070 0.46 6.6 6 40°10! 69°31' 91 8.6 Sand 2.23 90.0 10.0 0.0 0.034 0.26 7.7 7 40°00! 69°30! 128 11.9 Sand 2.79 Bass loro. W538 0.031 0.24 7.8 8 39°57! 69°30! 183 11.9 Sand 2.89 86.04 19580) 422 0.022 0.12 5.5 9 39°56! 69°45! 201 8.9 Sand 2.70 85.0 10.7 4.2 0.035 0.22 6.3 10 40°00! 69°45! 139 11.7 Silty- 3.85 57.0 33.4 9.5 0.037 0.61 7.0 Sand 11 40°10" 69°45! 95 73 Silty- 3.70 690" 236) 75 0.064 0.41 6.5 Sand 12 40°20! 69°46! 79 6.7 Sand 3.48 85.5) kOc2 524 0.059 0-41 7.0 13 40°30! 68°45! 73 72 Sand 2.63 9520) 19. Be2 0.026 0.17 6.6 14. 40°40! 69°45! 59 7.5 Sand 2.24 10001. (050) 050 0.004 0.03 7.9 15 40°50! 69°45! 37 9.2 Sand 2.05 100.0 0.0 0.0 0.003 0.02 8.0 16 40°46! 70°00! 38 8.1 Sand 2.67 Toos0). 020" SOLO 0.018 0.03 4.6 17. 40°39! 69°59" 49 7.5 Sand 3.27 STOOLS) “Ses, 0.072 0.39 5.5 18 40°30! 70°00! 73 6.8 Sand 3.65 670) 25632 TET 0.082 0.55 6.8 19 40°20! 49°59! 91 6.7 Sand 3.79 6220, 304) 16.7 0.069 0.48 6.9 20 40°10! 70°00! 117 11.4 Sand 4.12 45.0 44.8 10.2 0.080 0.58 Ted 21 40°00! 70°00! 165 (11.4) Sand 3.35 66.101, n239 Osa 0.058 0.48 8.4 22 40°03! 70°15!" 183 10.8 Silty- 2.18 66)18) 2356 86 0.060 0.59 9.9 5 6 sand 23 40°10! 70°15! 113 11.6 Silty- 4.31 36.2 53.4 10.4 0.092 0.70 7.6 3 a sand 24 ~=40°20! 70°15! 90 7.8 Silty- 4.12 47.0 38.9 14.1 0.133 0.92 6.9 o 5 sand 25 40°30! 70°15!" 70 6.6 Sand 3.83 58.8 30.7 10.5 0.076 0.52 6.8 26 40°40! 70°15" 51 Tse Sand 3.37 85.9) 1022) 358 0.048 0.19 4.0 2740950! 70°15! 44 8.1 Sand 3.43 SG le One 4e5 0.035 0.20 5.7 28 41°00! 70°15! 33 9.4 Sand 3.35 OO 7h maaeoMl Ae, 0.033 0.13 4.0 DOM Alor 70°16" 27 12h Sand 1585 | s100%0)) P40rONOxO 0.011 0.07 6.9 30. 41°10! 70°30! 38 8.3 Sand 203: ._ 100.0. . (Os0N 080 0.018 0.11 6.2 si 41200! 70°30! 48 7.5 Sand 2.81 79.5 14.1 6.4 0.044 0.28 6.3 32. 40°50! 70°30! 59 6.9 Sand 3.30 G75 24ele oes 0.073 0.51 7.0 33 40°40! 70°30! 62 Gey Silty- 3.48 60.0 25.3 14.7 0.137 0.84 6.2 6) % Sand Ba 40e30" 70°30! 73 7.5 Sandy- 4.54 33.2 49.0 17.8 0.148 1.03 6.9 & 5 silt 35 40°20! 70°30! 97 10.0 Sandy- 4.67 se}, Plas) eed! 0.146 ae 7.6 silt 14 Appendix Table 1 (cont.) Bottom Sediment Parameters Station Station location Water Water "Median | Composition Nitrogen Organic Carbon Lat. (N) Long. (W) depth temperature Type diameter Sand Silt Clay (Kjeldahl) carbon Nitrogen m 2% o % % % % % Sqn 40c10! 70°30" 128 12.1 Siltyye 242026: ais2ee) .1l0e2 7.0 0.046 0.41 8.9 3 sand 37. 40°04" 70°29! 220 9.4 Sand TAZ OMMNOSTSIy Piece) | USNO 0.038 0.28 7.5 38 40°02! 70°44! 194 10.8 Silty- 3.61 53.6 31.8 14.6 0.113 0.63 5.6 é 6 sand 39 40°10! 70°45! 132 11.7 Sility— 52020.) aes sboorauy tom 0.183 1.05 5.7 S sand 40 40°20! 70°46! 106 9.2 Sand- 5.01 14.0 61.6 24.5 0.212 1.46 6.9 é = silt-clay 41 40°30! 70°45! 79 6.7 Sandys u4heo) 419m3 | 62.6 18a 0.194 1.03 5.3 silt 42 40°40" 70°45" 66 6.4 silty- BAD MeeS5 a8) |. 29e7 lacs 0.112 0.69 6.2 5 sand 43 40°50! 70°45" 55 6.8 Sand 24371 18545 Toy. Woe 0.054 0.22 Aen 44 41°00! 70°45! 51 6.7 Sand 1248" | -100L0) 0x0, “O%0 0.008 0.07 8.7 45 41°10! 70°45" 38 8.6 Sand- 1.00 100.0 0.0 0.0 0.006 0.06 10.6 gravel 46 41°10! 71°00! 40 9.7 Sand TSO MLOORO) OsONe 0.0 0.007 @acey 2 ai 47. 41°00! 71°00! 51 6.4 Sand- 2.53 ese) ese) 6 ean = = 2 gravel 48 40°50! 71°00! 59 6.1 Sand 230 ese aE | ie) 0.046 0.23 5.0 49 40°40! 71°00" 70 6.3 Sandy- 5.09 16.4 61.0 22.6 0.220 1.09 5.0 silt 50. 40°30! 71°00! 84 6.3 Clayey- 5.84 SET 6943) 2726 0.245 1.24 5.0 silt 51 40°21' 71°00! 99 9.4 Sandy- 4.77 31.2 49.0 19.8 0.150 0.94 6.3 silt 52. 40°10! 71°00! 146 10.8 Silty- 2.35 70M ges) tales 0.067 0.45 6.7 sand 53. 40°06! 71°00! 179 10.8 Silty—| i477, 241i) (54-0, tse 0.124 1.04 8.4 = S sand 54 39°59! 71°00! 366 (6.1) silt SElomiume5eoeossum) © 8s0 0.112 0.88 7.9 55 39°56! 71°00! 567 (6.1) silt AESOmMM A267 S653) 2150 0.160 ileal) 7.3 56 40°03! 71°16" 183 10.3 Sand Dees eorON |) 1Ba4e | VSG 0.050 0.21 4.1 57. 40°10! 71°15! 110 10.8 Silty-" § 2868" | i6or0l 23-60 10n4 0.099 0.74 7.5 rm cS sand 58 40°20! 71°15! 91 9.0 Silty | 35298 5260) B5e3 re slo, 0.136 0.81 5.9 iS 3 sand 59 40°30! 71°15! 77 6.7 Silty= (229569450) 1965 YLT 0 0.086 0.32 357 é 5 sand 60 40°40! 71°15! 62 6.2 Sand Peeysle| aaa) alse | Teas 0.060 0.29 4.9 61 40°50! 71°15! 62 6.4 Sand 3.00 87.9 6.5 5.6 0.044 0.20 4.5 fo} 62 41°01! 71°16! 48 6.7 Sand 1263, 100.0 | 050) -) 0-0 0.009 0.06 Teak 63 41°10! 71°15" 38 8.1 Sand 2.14 100.0 0.0 0.0 0.003 0.06 7.9 64 41°00! 71°30! 55 6.9 Sand 2238) 9a h8944)) 1651s BaeAeS 0.042 0.24 5.6 15 Appendix Table 2.--List of Macrobenthic Invertebrates off Martha's Vineyard, Massachusetts. Porifera Cnidaria Hydrozoa Hydractinia echinata Anthozoa Cerianthus sp. Edwardsta sp. Epizoanthus americanus Pennatula aculeata Stylatula elegans Nemertea Annelida Aglaophamus circinata Ammotrypane aulogaster Amphitrite sp. Anctstrosyllis sp. Aphrodita hastata Arabella tricolor Artcidea jeffreysit Asychis biceps Brada sp. Capitella sp. Ceratocephale Lovent Chaetozone sp. Chone infundibuliformis Cossura Longoctrrata Drilonerets longa Euntce pennata Flabelligera sp. Glycera robusta Glycera tesselata Goniada brunnea Gontada maculata Harmothoe extenuata Hyalinoecia tubtcola Laonice cirrata Leantra sp. Lumbrineris fragilis Lumbrineris tenuis Melinna eristata Nephtys bucera Nephtys incisa Nerets pelagica Ninoe nigripes Notoctrrus sp. Onmuphis conchylega Onuphts opalina Onuphts quadricuspis Orbinia ornata Owenta sp. Paradiopatra sp. Paramphinome pulchella Paraonis neopolitana Phyllodoce mucosa Prionospto sp. Sealtbregma inflatum Sphaerodorum gracilis Spto sp. Sptochaetopterus sp. Sptophanes bombyx Sternaspis scutata Sthenelais limicola Streblosoma spiralis Tharyx sp. Sipunculida Golfingta catherinae Golfingia elongata Golfingia margarttacea Golfingia minuta Golfingia (Phascoloides) sp. Onchnesoma steenstrupt Phascolton strombt Arthropoda Amphipoda Aeginina longicornis Ampelisca compressa Ampelisea macrocephala Anonyx sp. Byblis serrata Caprella sp. Amphipoda (Cont.) Casco bigelowt Corophium sp. Dultichia sp. Erioptsa elongata Harpinta proptnqua Haustoriidae Hippomedon serratus Lembos sp. Levtochetrus pinguis Orchomenella groenlandica Paraphoxrus sp. Photts macrocoza Phoxocephalus holbollt Ptotomedia fasctata Siphonoecetes smithtanus Stenopleustes gracilis Unctola trrorata Unctola Leucopts Decapoda Axius serratus Cancer borealis Cancer trroratus Catapagurus sharreri Crangon septemspinosa Bythocarts nana Dichelopandalus lLeptocerus Europrognatha rastellifera Hyas coarctatus Munida iris Pagurus acadianus Pagurus arcuatus Pagurus politus Pontophilus brevirostris Isopoda Calathura sp. Chiridotea tuftst Ctrolana polita Edotea triloba Ptilanthura tenuts Cumacea Diastylis polita Diastylis quadrispinosa Eudorella emarginata Eudorellopsis sp. Leptocuma minor Leptostylis sp. Petalosarsia declivis Mysidacea Bathymysts renoculata Erythrops erythropthalma Hypererythrops caribbaea Mysts mtxta Neomysts amertcana Cirripedia Balanus sp. Pycnogonida Achelia spinosa Paranymphon spinosum Mollusca Amphineura Chaetoderma nttidulum Pelecypoda Anomta sp. Arctica tslandica Astarte undata Bathyarca pectunculotdes Cerastoderma ptnnulatum Crenella glandula Cusptdarta perrostrata Cuspidaria striata Ensis dtrectus Hiatella sp. Lyonsta arenosa Lyonsia hyalina Maecoa calcarea Mesodesma arctatum Mytilidae Nucula proxima Nucula tenuts Nuculana acuta Pandora gouldiana Pandora inflata Pertploma papyratium Phacoides blakeanus 16 Pelecypoda (Cont.) Phacoides filosus Placopecten mgellanicus Stliqua costata Spisula solidissima Tellina agilis Thracta sp. Thyasira ferruginosa Thyastra gouldt Thyastra ovata Thyasira tristnuata Venericardia borealis Yoldia sapotilla Scaphapoda Cadulus pandionis Cadulus verrillt Dentalium occidentale Gastropoda Alvania carinata Buccinum undatum Colus stimpsoni Crepidula plana Cructbulum striatum Cylichna alba Epttonium dallianum Lunatia heros Lunatia trisertata Nassarius trivittatus Neptunea sp. Polinices sp. Retusa gouldi Sceaphander sp. Echinodermata Asteroidea Astertas vulgaris Astropecten americanus Astropecten sp. Henricta sanguinolenta Leptasterias tenera Porania sp. Echinoidea Schtzaster fragilts Echinarachntus parma Ophiuroidea Amphilimna olivacea Amphtoplus abditus Amphioplus macilentus Amphiura fragilis Amphiura otteri Axtognathus squamatus Ophtura robusta Ophiura sarst Holothuroidea Caudina arenata Chiridota sp. Cuecumaria frondosa Havelockta scabra Molpadia oolitica Psolus fabricti Stereoderma untsemita Bryozoa Dendrobeania murrayana Electra hastingsae Electra pilosa Haplota clavata Hippothoa hyalina Seruparta chelata Ascidiacea Bostrichobranchus ptlularis Ctona intestinalis Cnemidocarpa mollis Heterostigma singulare Molgula eitrina Molgula complanata Molgula stphonalis Pogonophora Siboglinum atlantteum Stboglinum ekmant Hemichordata Enteropneusta Balanoglossus sp. Wet Weight Biomass (g/m?) of Major Taxa of Macrobenthic Invertebrates per Station off Martha's Vineyard, Massachusetts. Appendix Table 3. epodeydess epodddeted epodorz4se9 WOSNTIOW WINONNdIs WHOHdONODOd WdITANNy VaLdadwan VLVWAALNATAOO Wadd luod ToOquinn uoyze3g Siecle atS2O.0y Oy CAALTAANDTAAMMAOAM . eiencecte enie| sje pfecieiie seo i8) eee niertelaee) Vemiewielie T1LoOoNnnr lumuas SOCOHODHOVONOODRLN < +s Ages do mo tos ann ~ os ony ad uw 4 doHioed dN elias Ori Ose 05 eel iea (bef 550.0 On [ eef) SOs O21) alot et} =F} =30 Os} o No oodo no co Ho a a) NHN I (D O1OO1O NA SNOT 00 SENN en SN co as . eiWehierne aifeiteierare cies . ele liiesuemnectesteriecue mM I1aotrrnr 1! onMomMmMnNno od 8 ed owoNnoonornrm + +t AMnSs do oS x DAaAN x ns qd AN dddddians wdddddN dH oO Omg . . . . . UN Ue Det beet 3 S on ° o Yo} Fee | “4 3 o o 3 < OF IROE Irs ad a DUS CPS ITE Ts 1) iC et ete TTT (ESS Vet oT Lim t d best} = Ter} WOANNHTNEFDEFOFDMNDODTAMAMOMAMEHAN awe Cua Ds Onn On . . eset aeSnermelkelbelMelteney")s AMOCDNNAPTHNOODMONTDHADHANNNOMNAMO N OtNd A ANA tN HANrEr Ow 4d N NARN Am ain © Nd tos COMNHaAN omy co og O80. [Eevovventie> [pu |inlwene: . oe OMHAO rs ood ” o o oon NOANMHA® N qd FANANDNWOOMNG wd ™ NO AHAN OL (iO CO 2 ODE OS OED Ome [) aT] el (pe Taf ps Tk 0:06 O80 q MDOHWOOHNCO CO N HM oooONM qd AAW a ” on o) cae . BO DU UT UR USU SUPA Salley es CU UE Ue US Ui Ur Ue 808.0 808.0 oe) LT SOON ROU CMLA ALO Ste and ia) ONOFTDUODHDAN Od xs oMiteillesie Oy OF [Lm] > -00- [hp 2a O00 0 UO SOU) 0 "8 Amonmodomaa NO a OMANMWONT a) ad taal N aq a a 1.0 SESS GU GERD (Ra CaaS) wo . eels HICH SSI Simian es a N 4d N al a a - d Hs dq i ert, dq Oa hse) 0 Ie Ot" ets 0 Tee its Ui S0= Jato 0") Jone) oO Calta) oO Om 2 O20) oO Gal ad dN Ue eit (ist Drs lise (ete sho Desire Sah Neth oroeioteti) oo FIAMNAAMNVDOAOBTMNOMOWODNANAANNTAMEF Aw emerge, ome iy Orv Ginie ge . Oe {poe ONANAHANANASTONONANTRACHDAMNAMNDAD AM MAMNANT AT ANATMNAAITMNSTON dn asm onn Ad AAN AOKnA AAA aH wn eyeerus 20s OFF] . i sOO: On 0 f] Ono) {0 1) . ooo ° a ON Ao ooo onoo ooo (= oo al otn~r te MON NONDAHM MaAQo Ont ea sor oN} ooo ik te DA ha) O00 0 100 . N NEO FH MNO SARANG HARROW qd Loa N ete Us Wet Deu Pave a Doe Oa UE De 0 et a a 1) =) ANMAMORADANDOHANTNORDHOHAMTINWE AD MMMM MM Pt tt tt te tt ttt 19 19 1 Ln tn Ln in tn ti Galt vs ete |=2e) Sco N N N Ne} NNO aq . lee oor oO MUNHD wo woe (j220 Sof N N N Ke) 17 Appendix Table 3 (cont.) TWLOL PooeTpTosy WLSNANdOMALNA WLOOUdOLOG Paptoisezsy eaptoazntydo eaptoutyog PoptoznyyOTOH WLWWaadONTHOd epodesaq epodtyduy epodost Pooeund VaoOvLsndo Zaqunn uotze4S N UU Defh Te Oe O20 dist) soc UR th Uestee °o 4d Od qd doa eer Peer el od (0 bee x) Oo (==) wo a fon) d OU ete De Ue TO Lemay) aS faa °o (sa) ° d OOS ied Pees} ° Des tee et st Mind st © OS os =| on st ~ of Ding ce emcee AU cee seit azien' |. Teller NoHo (2) oO; ‘Oo = d ta) N iva) oNnw PAP Ty (bet bee Ot PTET ESO 00 ona eet tT a fel ” nNOo Nd KFonotMdttMNHA wo } [2.6505 ODO LOR DMO CORO EUs el bal | NAAR HAOTANDO oT d oOo Ord AN dq Md AHN wn [in PCIE [Pest [Ea SUIS at aL iO Fi Lia} Sh ~wo (of) = 25506 615.9 562.4 byes) 284.0 CA +tHoOnAMNO wo iO 0 00" O20 U0 WHOrNnOoONnr ts DAr-MNMNarsto ww N ACOH wo = fo} s+ © Osco) 0. Ue 1 020 d NN qd N LFie(hrUnoth STE SU Cha 1 qd dw Uae This {ea UO Fis SOs 0.577) o ow ad dq Sees Pete thes “et Co a ros ope aloit SN Mh healt wonnm Cd St taal Ie OCU eee es ns a Now oprorseett eit) IR le ul oNW ett = wonn tm od arlene. Jule en cera 10 «© © as a oma el in Na Te USO Ee Leet etl) i) od SON dat aM dd ad AOFM DNTFDANNODDNALPMNMNAGCAMNNWHAAVNGCVOTHOR ONO" OOO LO 3 CA t0 200" OD" 0 O50 00500 . 00-7070 WOOTMODEFAROAHSTOOCOMNANUVHOAMNADHANNDOR BPADLHPDONNANROMNOGCVOTARANOMNOFMAMTAN A tNAAN d at Nd doa d N oN ANWH So) ord ann dd OT PUT Up 60 V0 1S SO ses Oo Pecl he (rxeD-moimt)) | FSW Ae ent oe (=) IO) eit) nnd No oman ~m s Od qd mo q et Ua ie AOS) ike elise het ih OnOct) D Otel Def U nN °o >) OM She vata a etre OS Oe dhe t a0 tends Teeth Tathe oO x wu ow . doHwo mm wo ci cede ete is ie ie is Oe et Carte oe ou) iiss (i= oct) <0 oo ic) Coon OnS o Ya) NADDANOMO «A Am MMAMM INN Wea Wecioctegters ee oes opie ih ous [ma ly lle Nhe Wie Nieamey 9, om. one oucceierer ane’ 1 SAMNHAOHMH Oo oHom-orco®n re a a tmMHn 1M HoH MM A oO a4 d nO a Coal ViecsOoa Fm Jer bape) Finer ret nt [ar Jie fel Gist TR ec ee Ul a Meal PPS ates OS DST Shel To wo Oo ~ AM ° ~ MN qd Ano tN + raOarwo N a} ee Ot COTS One al ORS Leal eel el es Teepe ce O Oona hess 0 flo Tl jtHo OM N DOINMMN ° nd wo 4d AN qa aN d ANcCoOrAUuTNEN A dA cAAnnNotmMor ae egies’ WalGote: oometest emer froress it. iii ef fo} te teciemters | mRPONnWvmNtTaOtONR OO °o SomMModmMondaAN ndAomMmst mnM ONdwo AHAMnr a~NA ON dq d dq Cs (PST Seay eT fis PI Diath Po TSS eR TTS Te Stine heed tio Peer ST (heath fT (=) N AEE) CONGO LEN e119 Let COL fs ral Tei I Ol Co se teva IRN ce ole oy 0s te, ei te aire) cey emia’ fel einen enige deere NGAUWASSSSTUHMNTSENDELMSNDSODCSCCO® wo ANAN ~m > si rd HAwodtno qd qd qd qd OT aG (Pedy Sate se rai Om Oreo sone (Pesor [poe thane [ff [) = 20 o oo N i=) ONOOMO Oo o o o NANA Gl feb aah al Goss Ted d qd etre Mets et rentals | aie . jeeoeo-(} Tb i eon 7-0 coooo0o°o ooc000K0G00 oo oO Oo char) e) Melitomothelce. seime, . sience mlomielileimelue ASAwASSNSHUMINEIMIMoMadcccKCCS ANANHs 31 37 DHNOHANMNYTNOFADAHOAN ba i i i i i i iva) 18 Appendix Table 4. Number of Individuals per mof Major Taxa Per Station. < oO < é = a x o wo < 3 oO Hy oO <= 2 a ees » say 2 Se ae Sige) Sas ine 3 ai ae WEOKS yD Role OUR e ¢ se & a = = Slum 3 canes s $$ Bieter sa: 5 = apes = 3 a SS a Rep asec os oS aibeaeoe Sous fe KS Satnaks ee A eet Moe ea ets a 2 8 1 1 TI a, Lae 22 011) 21 =" “W192 SS Sab aA eRL LS =a - - 1,932 - - 4,324 2 1 - - 53 - - - - - 73 = 32 31 11 = - - - 11 - - 148 3 - - 32 42 - - Ls Re 170 rahul 138 - 21 St phy cs - - ob 287 4 - 144 21 1,360 - 41 S7aelle p26) Rm 2 379) 82) 36) “2252; 710 135 - 1355 - - 11 - 111 4,139 5 - 16" 166) 1,129) T 179 16 163 - 10,957 47 21 10,868 21 32 =) 1 21 - - - 12,340 6 - 10 62 955 - - 279 - 279 - 8,970 16 27 8,916 11 225% 36) = 189 - - - 62 10,563 8 - 21 - 572 - 62 14. -- «114 ~«- 31 lo - 21 - 489 - - 489° - - - - 1,289 3 1 140 ll 483 - ll 166 - 123 43 222 = Abt 211 - 32 — 32) - =" 21) «1/087 10 - 51 10 170 - 10 GY Glee 36 1039 = 26 - 379 - 10 339 30 10 o) bs 727 11 - 23-28 500 - 28 491129479. — 564 35 ll 518 - 1,096 81 - 1,015 - - =" 23) 25753 12 - lo 15 678 - 10 159 10 149 - 9,353 10 10 9,333 - 4. 31 - lo - - 51 - 10,317 13 - io 10 1,346 - 20 255 10 245 - 19,083 71 10 18.992 10 30 lo - 20 - - - 41 20,795 14 - lo 26 820 - 31 201 One lONe ik 371 374 37: 297 - 412 - 392 10 10 10 = LOW ln 7LO 15 - - 10 260) == 20m 2 524 Sega SV ALON i cy i - - - - - 581 16 - - - 456 - - 21 — ee — 631 16 32 583 - ll - au - - ll - 196 1,326 17 - 10 21 1,082 - 26 235', 47> (68) =" 3/031! 62 62 2,907 - 172 =) 172" > = - - - 31 4,488 18 - lo 415 651 - 10 308 40 268 - 3,030 20 10 3,000 = - - - - - - 10 4,034 19 - - 20 394 - 51 230) 230) e257 08 =~ =) 525-708) - 861 200 - 651 10 - - 10 4,274 20 - 200 41 - 8 2 ent ee 84 12a 720~«—- 3440 - 10 324 10 = - - 534 2) — ee py W209 1539153 )an 25 lo - 15) == 335\0 a= 15M 310810 = = = 705 27 — 2 = 199) freq Tht eer ple 41 = ha (Als se 41610 — 406 - - - - 788 23~«C 35 20 379 - 101 81 ee sly = 50 20 20 10) = 57 Ome LON 556 10 - - 51 1,293 2 20 30 31s) 5s 25 me 25 lll = - lll - 283 olen 192 - - - 787 25a = 25 838s moe 576 20 556 - 343 20 - 313) (10s a = = = - - - = 1,782 26 = lo 15 1,184 - 49 7122) 520-9702, = 8a 15312 66 10 1,236 - os 3 = = - 10 - = 3,263 27% i= 31 10 750 ae 357 esa | e999 5601-20! e923) = 15! 15 - = - - - 3,162 ag) = LOMMHLONIE225¢0 = a>y B25 g1 0720) See 37: 10 222 1,484 21 165 a 165 = - 10 - - 3511 29° = 20 «21 867. 829 7200) 162), 511326) 36 42 1,238 10 220 = 22 - 31 - 541 2,910 30 - = 16 993 - ag 37 =) 375 awe 409m 159510956 758 «= 22 225 = = - - 49 2,574 31 = 10 10 833 - 109 239957 56):183) 438839 13719) 23),173)1) 20a — 10. - 10 10 - Ope 025 32) = = 25 24Tae ae 1010 810559) e— 76 10 - 66 = 104 = = = 10 a = = 459 cee é 15 490 - _ 457 354 121 - 152 51 20 81 - 20} 810) 10a = - - - 1,152 34 30 10 22790 162i eit = 853 35 10 788i n2Ou 12) eS Ge S6u as - - - 1,414 35 i 30 20 379 - 49 1,030 404 626 - 131 20 - lll = Th SO5iates - - - 2,206 36 = 76 «010 39 Ger— y 30 - 30 eee 50 10 - 40, - 626.00 = A2bs <= - 5 shee alpsily 37) = = 10 61 - 13iteelO, 121 = 30 = - 30, = 368540) S10) S18 — - - 20 620 Appendix Table (cont.) < © < < & © BE & a) < = = vc. oc wo < a a c 7 = 2 2 o < > Ps 2 5 < 3 8 a 3 o f 9 Se pcg 0) eesetwe ad 5 a & Ps a x 5 oS a a ° ° 3 © ° 7 a 3 _ ° a ° a iS) on m 2 & nl fo) o a ° > a= < o 7. a ° oS <= ° & ° = re) < “Oo mh mw = ~ z z —) Me ro] a & ] ° ot a z “ ce > h a & a - oor ~ a a o —) a vo ev i) n a a. o- i] i) ° ~ ~~ oe o wh ” < se = w = z =) a a a = & > € ° a ca) x= = x a ) & 5 oO & »3s 0 ° cs] Zz o mH (>) a o ov = a ao § cy Oo ° uv a 2 oO a ° nz & Oo =z < a n = o a n oO oO Lal a nm = mw o < m we =< id 38 - 42 - 965 - 62 82 10 62 10 139 - 10 129 - 20 10 - 10 = - = = 1,310 39 10 177 - 91 20 71 - 20 - - 20 - 717 - 10 707 - - 106 1,121 40 — 51 10 61 - - 570 30 540 - 166 10 10 146 - 1,606 20 778 808 - - - 51 2,512 41 - 30 = 182 - - 51 - 51 - 15393 40 - L303 - 30 - - 20 10 - - 81 1,767 42 - 10 10 1,020 = - 133 102 31 - 1,571 10 - 1,561 - - - - - - - - - 2,744 43 = = 10 763 - - 46 31 15 - 1,960 10 10 1,940 - 10 - - 10 - - - - 2,789 4&& - - 10 959 - - 46 20 26 - 2,800 31 102 2,667 - - - - - - =~ = = 3,815 45 = = - 841 - -= - - - - 5,623 10 10 5,603 = - - - - - - - 98 6,562 46 = = 31 619 - = 58 21 37 - 4,381 68 21 4,292 - 16 - 16 - - - - 73 5,178 47 - - 20 444 - - - - - - 3,075 30 81 2,964 - - - - - - - - 20 3,559 48 - - 16 1,804 - = 198 73 125 - 3,651 276 1643), 338 21 16 - - - 16 = — = 5,685 49 - 15 10 1,116 - - 177 81 96 - 813 - - 813 - 30 - - - 30 - - - 2,161 50 - 10 - 333 - - 252 10 242 - 40 20 - 20 - 125 15 - 101 10 = - 51 812 51 - 51 20 273 - 10 394 30 364 - lll 10 30 71 - 747 45 - 702 - - 30 71 1,707 52 - 77 10 979 - - 280 10 270 - 10 - - 10 = 461 10 - 461 - - 10 = 1,837 53 10 10 10 328 - 20 273 - 273 - 25 - - 25 - 222 10 - 212 - - - - 898 54 - 20 - 803 Sil 9 7, 91 25 51 15 25 - 10 15 = 35 10 - 25 - - = = 1,222 55 - = - 432 36 - 30 10 20 - 10 - - 10 = 20 - - 20 - - 10 = 538 56 = 51 10 503 - - 133 10 123 - 116 - 10 106 - 72 - - 62 10 - - - 885 57 - 10 - 259 - 36 71 - 71 - 66 20 - 46 - 550 10 - 540 - - - 15 1,007 58 - 35 Sp e374 - - 576 10 566 - 51 - - 51 - 697 - = 687 10 - - 61 1,809 59 — 35 10 490 - - 56 - 56 - 15 - - 15 - 126 - = 126 - - = 354 1,086 60 - - 15 938 - 15 525 20 505 - 1,555 50 10 1,495 - - - - - - - = 126 3,174 61 - = 1579 6} - 10 122 20 102 - 3,442 162 30 3,250 - 10 - - = 10 = = = 4,355 62 = 12 o 871 - 53 62 De 10 - 4,980 12 72 4,896 - 10 - 10 = - - - 54 6,042 63 - = - 380 - 21 - - = - 1929: 73 26 1,830 = 135 - 125 10 = = = = 2,465 64 - - - 439 - 63 83 10 73 - 3,681 177 10 3,483 ll 10 - 10 = - = = 10 4,286 20 NOAA TECHNICAL REPORT NMFS Guidelines for Contributors CONTENTS OF MANUSCRIPT First page. 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