746 NOAA Technical Report NMFS SSRF - 746 K^t^-f 0*^00^ Distribution of Gammaridean Amphipoda (Crustacea) on Georges Bank John J. Dickinson and Roland L Wigley June 1981 ,.— . Marine Biological Laboratory ; LIBRARY I OCT 14 1992 Woods Hole, Mass. U.S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service 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, lo understand and predict fluctuations in the quantity and distribution of these resources, and to establish levels for optimum use of the resources. 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Leitzell, Assistant Administrator for Fisheries The National Marine Fisheries Service (NMFS) does not approve, rec- ommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales pro- motion 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. CONTENTS Introduction ' Georges Bank — description of environment 1 Georges Bank— Amphipoda • Methods ' Annotated list of species 2 Discussion ^ Zoogeography ^ Species distribution patterns ^ Northeast Peak ^ Perimeter ° Western Basin ^ Shoals 8 Southern Edge 8 Central 8 Acknowledgments ° Literature cited 8 Table 1 . Six patterns of gammaridean distribution on Georges Bank 7 Figures 1 . Geographical features and bathymetry of Georges Bank 10 2. Geographic distribution of bottom sediment types and station locations on Georges Bank 11 3. Distribution of average bottom-water temperatures during February and September on Georges Bank 12 Geographic distribution of species of gammaridean Amphipoda on Georges Bank: 4. Acanthohauslorius intermedius, A. spinosus, A. similis, A. sp. C, A. sp. D, Acanthonotozoma serratum, Anda- niopsis nordlandica, A mpelisca agassizi 13 5. A mpelisca decliviiatis, A. macrocephala, A. vadorum, Amphilochoides odontyx, Amphiporeia giganlea, A. vir- giniana, Amphithoe rubricata, Anonyx debruyni, A. sarsi, A. liljeborgi j 14 6. Argissa hamatipes, Bathymedon sausserei, Balhyporeia quoddyensis, Sybils gaimardi, B. serrata, Calliopius lae- viusculus, Casco bigelowi, Corophiuin crassicorne 15 7. Dulichia tuberculata, Dyopedos arclicus, D. monacantha, Epiineria loricala, Ericthonius rubricornis, Eriopisa elongata, Eusirus cuspidal us, Ganunaropsis maculatus, Cammarus annulalus 16 8. Gilanopsis arctica, Halice abyssi, Halirages fulvocinclus, Haliragoides inennis, Haploops lubicola, Harpinia pro- pinqua, Hippomedon propinquus 17 9. Hippomedon serralus, Ischyrocerus anguipes, I. megacheir, Leplocheirus pinguis, Melita dentata, Maera danae, Melita sp. A, M. sp. B, Melphidippa goesi 18 10. Monoculodes edwardsi, M. inlermedius, M. latimanus, Monoculopsis longicornis, Neopleustes pulchellus, Orcho- mene mmuta, O. pinguis, Paradulichia typica 19 1 1 . Parahaustorius attenualus, P. holmesi, P. longimerus, Paraphoxus epistomus, Pardalisca cuspidata, Photis den- tata, P. macrocoxa, Phoxocephalus holbolli 20 12. Platylshnopus sp. A, Pleusymles glaber, Pleustes panoplus, Podoceropsis nilida, Pontogeneia Inermis, Protohaustorius deichmannae, P. wigleyl 21 13. Protomedela fasciata, Psammonyx nobilis, Pseudohaustorius borealls, Pseudunciola obliquua, Rhacholropis disllncla, R. inflata, R. oculata 22 14. Siplwnoecetes smithianus, Stegocephalus inflatus, Slenopleustes gracilis, S. inermis, Stenothoe minuta, Stenula peltata, Synchelidium americanum, Syrrhoe crenulata, Tiron spiniferum 23 15. Tmetonyx cicada, Tryphosella nanoides, Unciola dissimilis, U. inermis, U. irrorata, U. spicata, Westwoodilla megalops 24 16. Most common distribution patterns of gammarideans on Georges Bank 25 Distribution of Gammaridean Amphipoda (Crustacea) on Georges Bank JOHN J. DICKINSON' and ROLAND L. WIGLEY' ABSTRACT The distribution of 97 species of gammaridean amphipods is described for the Georges Bank region, based on 379 samples from 326 stations. A wide variety of sampling gear was used, resulting in representation of both infaunal and epifaunal species. Geographic and bathymetric distributions, and sediment preferences are summar- ized for each species. The zoogeography of the gammaridean fauna of Georges Bank is discussed, and the most common geographic distribution patterns are described and related to environmental parameters. INTRODUCTION This report describes distribution patterns for 97 species of gammaridean amphipoda on Georges Banic. Georges Bank is a submerged coastal plain that extends over 300 km east of Cape Cod, Mass., and encompasses an area of approximately 19,000 km^ (Fig. 1). It is one of the most productive fishing grounds in the world yielding thousands of metric tons of finfish and shell- fish annually (Gusey 1977). In the future, Georges Bank might also serve as an offshore source of petroleum. Gammaridean amphipoda are the most numerous group of macroinfaunal organisms inhabiting Georges Bank, composing 49"Vo of the number of specimens in quantitative samples col- lected there (Wigley and Theroux in prep.). Gammarideans are important food for many juvenile bottom- feeding fish. They often comprise 30% or more of the diet (Bowman'). The purpose of this report is to provide information useful to both benthic ecologists and fisheries biologists interested in the Georges Bank ecosystem. Since amphipods are known to be sen- sitive to environmental aheration, the data presented in this report may also be useful as a baseline in the event of petroleum development on Georges Bank. GEORGES BANK— DESCRIPTION OF ENVIRONMENT For purposes of this report, we define Georges Bank as that part of the New England continental shelf between Great South Channel (long. 69 °W) and Northeast Channel at water depths <200 m (Fig. lA). The important physiographic features of Georges Bank include: 1) a steeply sloping northern face; 2) an area of shoals in the north-central part; 3) a gently sloping southern face (Fig. IB). The bank has been shaped by glacial erosion in the past, and recently by the strong tidal currents that run through the shoals (Stewart and Jordan 1964). The sedi- ments of Georges Bank are dominated by sands (Wigley 1961). The central area of the bank including the shoals is chiefly medi- ' Northeast Fisheries Center Woods Hole Laboratory, National Marine Fisher- ies Service, NOAA, Woods Hole, Mass.; present address: National Museum of Canada, Ottawa KIA OM8, Canada. 'Northeast Fisheries Center Woods Hole Laboratory, National Marine Fisher- ies Service, NOAA, Woods Hole, MA 02543. 'Raymond Bowman, Fisheries Biologist, Northeast Fisheries Center Woods Hole Laboratory, National Marine Fisheries Service, NOAA, Woods Hole, MA 02543, pers. commun. December 1979. um to coarse sand (Fig. 2A). Gravel covers the Northeast Peak, but fine sands occur around most of the perimeter of the bank. Silts, clays, and fine sands are the dominant components in the Western Basin. Bottom water temperatures on Georges Bank vary a moderate amount during the year usually reaching a maximum in Septem- ber and a minimum in February (Colton and Stoddard 1973). The greatest seasonal changes occur in the shallow central area of the bank which experiences about a 12 °C annual fluctuation ranging from 4° to 16°C(Fig. 3 A, B). The perimeter of the bank has a much smaller annual temperature range fluctuating as little as 2°C in some areas. The coldest year-round bottom waters (6 °-8 °C) occur in the Western Basin and on the North- east Peak. GEORGES BANK— AMPHIPODA The gammaridean amphipod fauna of Georges Bank has been little studied previously. Smith and Harger (1874) collected a few samples from the bank and reported the amphipods which they found, but the taxonomy is now outdated and coverage of the bank was incomplete. Some records of occurrence on Georges Bank are given in papers of Shoemaker (1945b) and Bousfield (1965). Whitely (1948) collected 42 species of epiben- thic and pelagic gammarideans on Georges Bank, but he gave distributional data for only one species, Monoculodes edwardsi. METHODS The distribution patterns described in this report are based on samples collected by the Benthic Dynamics Investigation at the Northeast Fisheries Center at Woods Hole, Mass. These collec- tions were obtained as part of a survey, conducted between 1953 and 1965, to obtain an overview of the general composition and distribution of the macrobenthos on the continental shelf be- tween Nova Scotia and Florida. A total of 379 collections from 326 stations were utilized in charting the distribution of gammarideans on Georges Bank (Fig. 2B). The coverage of the bank was relatively complete except for the Shoals area where fewer samples were taken due to the hazards of navigating there. The samples were collected with a variety of sampling gear, including 1) quantitative grabs (Van Veen, Smith-Mac Intyre, Campbell), 2) epibenthic sled nets (ring and meter), 3) Dibby dredges, 4) otter trawls, and 5) from fish stomachs. The Smith-Mac Intyre grab (160 samples), the ring net (85), and the meter net (50) were the most frequently used samplers. Due to the variety of gear used and the large number of samples collected, we feel that a representative species list was obtained of both infaunal and epifaunal amphi- pods. Estimates of numerical density given in this report were based on samples collected with either the Smith-Mac Intyre (0.1 m-) or the Campbell (0.56 m') grabs. Nearly all the gammaridean species that we collected on Georges Bank are described in previous taxonomic works. Bous- field's (1973) excellent systematic monograph on the shallow- water amphipod fauna of New England described about one- half of the species found on the bank. The next most important reference is Sars (1895) classic work on the Crustacea of Nor- way, which includes about one-third of the species occurring on Georges Bank. Other useful systematic references include Holmes (1905), Shoemaker (1930a, b, 1945a, b), Stephensen (1935, 1938, 1940, 1942), Gurjanova (1951), Barnard (1960, 1969, 1971, 1972, 1979), Bousfield (1965, 1970, 1977, 1978), Mills (1967, 1971), and Laubitz (1977). The species names used in this report primarily follow Bousfield (1973), but recent revisions by Barnard (1969) and Laubitz (1977) have been incorporated. The systematic arrangement of species into families and super- families follows the phylogenetic classification proposed by Bousfield (1978). ANNOTATED LIST OF SPECIES The following list gives a summary of the geographic and bathymetric ranges and sediment preference for each species based on literature sources. Notes on general ecology are also included when information was available. The distribution of each species on Georges Bank is also characterized based on our collections, and a series of charts (Figs. 4-15) show the patterns of occurrence for each species collected. These charts are arranged alphabetically according to the species names. Class Crustacea Subclass Malacostraca Superorder Peracarida Order Amphipoda Suborder Gammaridea Superfamily Phoxocephaloidea Family Phoxocephaidae Harpinia propinqiia G.O. Sars, 1895. Arctic-Boreal regions of North Atlantic; in western Atlantic: Arctic south to Cape Hatteras; 10-1,500 m, sand to silt -clay bottoms, bur- rowing detritivore (Sars 1895; Bousfield 1973; Dickinson et al. 1980). Perimeter of Georges Bank, 84-196 m, gravel to silt-clay bottoms, 4.5°-7.7°C, densities 10-50/m^ Phoxocephalus holbolli Kroyer, 1842. Arctic-Boreal in North Atlantic; in western Atlantic: Arctic south to Vir- ginia; 0-400 m, sand and silty sand bottoms, burrowing detritivore (Bousfield 1973; Dickinson et al. 1980) m'. Perimeter of Georges Bank, 33-179 m, sand and gravelly sand bottoms, 7.1°-12.3°C, densities 10-100/m^ Paraphoxus epistomus (Shoemaker, 1938). (Trichophoxus episiomus in Bousfield 1973. See Barnard 1979 for most recent discussion and proposal to change name to Rhep- oxynius epistomus.) Boreal regions of North Atlantic and North Pacific; in western Atlantic: Maine south to North Carolina; 0-180 m, sand bottoms, burrowing detritivore (Barnard 1960; Bousfield 1973; Dickinson et al. 1980). Central Georges Bank, 33-123 m, gravel and sand bot- toms, 5.4°-12.3°C, densities 10-150/m-. Family Platyishnopidae. Platyishnopus sp. A (undescribed species). Southern edge of Georges Bank, 121 m, sand bottom, 7.5 °C. This species will be described by E. L. Bousfield of the Canadian Na- tional Museum in Ottawa. Superfamily Lysianassoidea Family Lysianassidae Anonyx debniyni Hoek, 1882. Arctic waters of North At- lantic; in western Atlantic: south from Gulf of St. Law- rence to Cape Cod; bathyal depths and cold water (Steele and Brunei 1968). Western Basin of Georges Bank, 183 m, silt-clay bot- toms, 6.6 °C. Anonyx liljeborgi Boeck, 1871. Arctic-Boreal regions of North Atlantic; in western Atlantic: Arctic south to Dela- ware; 0-200 m, sand and silty sand bottoms, epibenthic scavenger (Steele and Brunei 1968; Bousfield 1973). Perimeter of Georges Bank, 60-183 m, sand and silt-clay bottoms, 6.6°-11.6"C. Anonyx sarsi Steele and Brunei, 1968. Circumpolar; in west- ern Atlantic: Arctic south to New Jersey; 0-65 m, sand bot- toms, epibenthic scavenger (Steele and Brunei 1968; Bous- field 1973). Southern edge and Northeast Peak of Georges Bank, 70-97 m, gravel to sand bottoms, 7.4°-7.7°C. Hippomedon propinquus Sars, 1895. Boreal regions of North Atlantic; in western Atlantic: from Nova Scotia south to Cape Hatteras; 15-250 m, gravelly sand to silt-clay bottoms, epibenthic (Sars 1895; Dickinson et al. 1980). Western Basin and perimeter of Georges Bank, 73-196 m, sand to silt-clay bottoms, 6.6°-11.6°C. Hippomedon serratus Holmes, 1905. Western Atlantic: Gulf of St. Lawrence south to North Carolina; 5-90 m, sand and silty sard, epibenthic (Bousfield 1973; Dickinson et al. 1980). Southern Georges Bank, 35-106 m, sand and gravel bot- toms, 7.3°-11.6°C. Orchomene minuia Kroyer, 1846. Arctic-Boreal regions of North Atlantic; in western Atlantic: south from Baffin Island to New Jersey; 0-100 m, sand bottoms, epibenthic (Bousfield 1973; Dickinson et al. 1980). Southern Georges Bank and Northeast Peak, 48-179 m, sand and gravelly sand bottoms, 7.4°-11.6°C, densities 10-100/m^ Orchomene pinguis Boeck, 1861. Arctic-Boreal regions of North Atlantic; in western Atlantic: Arctic south to North Carolina; 0-100 m, sand bottoms, epibenthic scavenger (Stephensen 1938; Bousfield 1973). Perimeter of Georges Bank, 60-168 m, sand and silt-clay bottoms, 6.6°-11.2°C. Psammonyx nobilis Stimpson, 1853. Western Atlantic: Newfoundland to New Jersey; 0-60 m, sand bottoms, bur- rower (Scott and Croker 1976; Dickinson et al. 1980). Northern Georges Bank, 22-128 m, sand and gravelly sand, 6.1°-9.3°C. Tmelonyx cicada O. Fabricius, 1780. Boreal-Arctic regions of North Atlantic; in western Atlantic: Arctic south to New Jersey; 0-2,000 m, sand and silty sand bottoms, epibenthic scavenger (Sars 1895; Stephensen 1935). Perimeter of Georges Bank, 91-183 m, sand and silt-clay bottoms, 6.3°-7.6°C. Tryphosella nanoides Liljeborg, 1865. Eastern North Atlan- tic; not previously reported from western Atlantic; 100-700 m (Sars 1895; Stephensen 1935). Northern edge of Georges Bank, 179 m, sand bottom, 7.8 °C. Superfamily Pontoporeioidea Family Pontoporeiidae Amphiporeia giganlea Bousfield, 1973. Western Atlantic: Cape Cod south to New Jersey; 10-40 m, sand bottoms, burrowing detritivore (Bousfield 1973; Glennon 1979). Central Georges Bank, 20-55 m, sand bottom, 7.8°-9.4°C. Amphiporeia virginiana Shoemaker, 1933. Western Atlan- tic: Nova Scotia south to South Carolina; 0-15 m, sand bottoms, burrowing detritivore (Bousfield 1973; Hager and Croker 1979). Central Georges Bank, 33 m, sand bottom, 8.4°C. Balhyporeia quoddyensis Shoemaker, 1949. Western Atlan- tic: Nova Scotia south to Virginia; sand bottoms, burrow- ing detritivore (Bousfield 1973). Central Georges Bank, 33-66 m, sand bottom, 7.6 °C. Family Haustoriidae Acanthohaustorius intermedins Bousfield, 1965. Western Atlantic: Cape Cod to Florida; 0-40 m, sand bottoms, bur- rowing filter feeder (Bousfield 1973; Dickinson et al. 1980). Central Georges Bank, 33-66 m, sand bottom. Acanthohaustorius similis Frame, 1980. Western Atlantic: Cape Cod south to Cape Hatteras; 10-74 m, sand bottoms, burrowing filter feeder (Dickinson et al. 1980; Frame 1980). Central Georges Bank, 33-66 m, sand bottom. Acanthohaustorius spinosus Bousfield, 1962. Western At- lantic: Nova Scotia to New Jersey; 0-200 m, sand bottoms, burrowing filter feeder (Bousfield 1973). Central Georges Bank, 64 m, sand bottom. Acanthohaustorius sp. C (undescribed species). Western Atlantic: Cape Cod to Cape Hatteras; 10-40 m, sand bot- toms (Dickinson et al. 1980). Central Georges Bank, 46-49 m, sand bottom. Acanthohaustorius sp. D (undescribed species). Central and Southern Georges Bank, 46-82 m, sand bottoms. Parahaustorius altenuatus Bousfield, 1965. Western Atlan- tic: Cape Cod south to Virginia; 0-75 m, sand bottoms, burrowing filter feeder (Bousfield 1965, 1973; Dickinson et al. 1980). Central Georges Bank, 33-66 m, sand bottom. Parahaustorius holmesi Bousfield, 1965. Western Atlantic: Cape Cod to Virginia; 20-50 m, sand bottoms, burrowing filter feeder (Bousfield 1965, 1973). Central Georges Bank, 35-42 m, sand bottom, 13.4°C. Parahaustorius longimerus Bousfield, 1965. Western Atlan- tic: Cape Cod Bay south to northern Florida; 0-10 m, sand bottoms, burrowing filter feeder (Bousfield 1965, 1973; Croker 1967). Central Georges Bank, 20-66 m, sand bottom. Protohaustorius deichmannae Bousfield, 1965. Western At- lantic: Maine to South Carolina; 0-40 m, sand, shelly sand, and silty sand bottoms, burrowing filter feeder (Bousfield 1965, 1973; Dickinson et al. 1980). Central Georges Bank, 46-60 m, sand bottom. Protohaustorius wigleyi Bousfield, 1965. Western Atlantic: Maine to North Carolina; 0-150 m, shell and sand bot- toms, burrowing filter feeder (Bousfield 1965, 1973; Dick- inson et al. 1980). Central and southern Georges Bank, 33-95 m, sand and gravelly sand bottoms, 7.6°-12.3°C, densities 10-300/m-. Pseudohaustorius boreatis Bousfield, 1965. Western Atlan- tic: Georges Bank to Virginia; 10-60 m, sand bottoms, bur- rowing filter feeder (Bousfield 1965, 1973; Dickinson et al. 1980). Central and northern Georges Bank, 46-88 m, sand bottoms. Superfamily Gammaroidea Family Gammaridae Gammarus annulatus Smith, 1873. Western Atlantic: Nova Scotia to Long Island Sound; 0-200 m, sand bottoms, epibenthic or pelagic, omnivore (Bousfield 1973; Dickinson et al. 1980). Southwest Georges Bank, 123 m, sand bottom, (it is noteworthy that this species, which reaches densities of 600/m' on Nantucket Shoals (Dickinson et al. 1980), was represented by only a single specimen on Georges Bank.) Superfamily Eusiroidea Family Eusiridae Eusirus cuspidatus Kx6yer, 1 845. Arctic Atlantic; in western Atlantic: Arctic south to Bay of Fundy; 40-400 m, epiben- thic or pelagic, probably carnivorous (Sars 1895; Stephen- sen 1935). Western Basin of Georges Bank, 183 m, silt bottom, 6.6 °C. Rhachotropis distincta (Holmes, 1908). Arctic-Boreal re- gions of North Pacific and North Atlantic; in western Atlantic: Arctic south to Gulf of St. Lawrence, 300-1,200 m, epibenthic carnivore (Barnard 1971; Bousfield 1973). Western Basin of Georges Bank (southern range exten- sion), 157-196 m, silt-clay bottoms, 6.6°-7.7°C. Rhachotropis inflata (G. O. Sars, 1882). Boreal regions of North Pacific and North Atlantic; in western Atlantic: Arc- tic south to Virginia; 20-100 m, epibenthic carnivore (Bar- nard 1971; Dickinson et al. 1980). Southern edge of Georges Bank, 72-144 m, sand and gravel bottoms, 7.2°-12.6°C. Rhachotropis oculata (Hansen, 1887). Pan-Arctic and Arctic-Boreal; in western Atlantic: Greenland south to Cape Cod; 5-l(X) m, epibenthic and pelagic carnivore (Bousfield 1973). Southeast Georges Bank, 82-95 m, sand bottom. Family Pontogeneiidae Pontogeneia inennis (Kr6yer, 1842). Arctic-Boreal in North Atlantic and North Pacific; in western Atlantic: Arctic south to Virginia; 0-80 m, rocky bottoms, epibenthic and pelagic (Bousfield 1973; Dickinson et al. 1980). Central Georges Bank, 20-179 m, gravel to sand bot- toms, 7.0°-11.3°C. Family Calliopudae Calliopius laeviusculus (KreJyer, 1838). Arctic-Boreal in North Atlantic and North Pacific; in western Atlantic: Labrador south to New Jersey; 0-30 m, rock and gravelly sand bottoms, 0°-22°C, pelagic and epibenthic carnivore (Bousfieid 1973; Steele and Steele 1973). Central part of Georges Bank, 20-59 m, sand bottom, 7.6''-ll.rC. Haliragesfulvocinclus (M. Bars, 1854). Circumpolar Arctic; in western Atlantic: Arctic south to New England coast; 10-400 m, gravel and clay bottoms, epibenthic (Sars 1895; Stephensen 1938). Western Basin of Georges Bank, 157-183 m, silt-clay bottom, 6.6°-6.8°C. Haliragoides inermis (C . O. Sars, 1882). Arctic-Atlantic; in western Atlantic: Arctic south to Gulf of St. Lawrence; 30-2,000 m, silt-clay bottoms, epibenthic (Sars 1895; Stephensen 1938). Western Basin and Northern Edge of Georges Bank (southern range extension), 152-196 m, gravel to sih-clay bottoms, 6.6°-7.7°C. Family Paramphithoidae Epimeria loricata G. O. Sars, 1879. Arctic-Boreal regions of North Atlantic; in western Atlantic: Arctic to New Jersey; 150-1,400 m, mud bottoms (Sars 1895; Stephensen 1938; Watling 1979). Western Basin and Northern Edge of Georges Bank, 157-183 m, sand and silt-clay bottoms, 6.6°-7.6°C. Superfamily Oedicerotoidea Family Oedicerotidae Bathyinedon saitsserei (Boeck, 1871). Boreal North Atlan- tic; not previously recorded in western Atlantic; 100-600 m, burrowing detritivore (Sars 1895; Stephensen 1938). Western Basin and Northern Edge of Georges Bank (first western Atlantic records), 154-179 m, sand bottoms, 7.6°C. Monoculodes edwardsi Holmes, 1908. Western Atlantic: Gulf of St. Lawrence south to Florida; 0-80 m, sand bot- toms, burrowing detritivore (Bousfieid 1973; Dickinson et al. 1980). Central Georges Bank, 20-179 m, sand and gravel bot- toms, 7.1°-12.6°C, most abundant species in epibenthic collections. Monoculodes intermedius Shoemaker, 1930. Boreal regions of North Atlantic; in western Atlantic: Gulf of St. Law- rence south to Cape Cod Bay; 5-50 m, sand bottoms, bur- rowing detritivore (Bousfieid 1973). Northeast Peak of Georges Bank, 101 m, gravel bottom, 8.2 °C. Monoculodes lalimanus (Goes, 1866). Boreo-Arctic regions of North Atlantic; in western Atlantic: Arctic south to Gulf of St. Lawrence; 5-100 m, burrowing detritivore (Sars 1895; Stephensen 1938; Bousfieid 1973). Southern Georges Bank, 84 m, gravel bottom. Monoculopsis longicornis (Boeck, 1871). Arctic-Boreal re- gions of North Atlantic; in western Atlantic: Arctic south to Gulf of St. Lawrence; 0-100 m, sand and mud bottoms, burrowing detritivore (Sars 1895; Stephensen 1938). Northern Edge of Georges Bank (southern range exten- sion), 156-168 m, sand bottom. Synchelidium americanum Bousrield, 1973. Western Atlan- tic: Central Maine to Georgia; 0-40 m, sand bottoms, bur- rowing detritivore (Bousfieid 1973; Dickinson et al. 1980). Southern Georges Bank, 73-84 m, sand and gravel bot- toms, 10.5 °C. Westwoodilla megalops (G. O. Sars, 1882). Arctic-Boreal regions of North Atlantic and Arctic Oceans; in western Atlantic: Arctic south to Gulf of St. Lawrence; 10-90 m, burrowing detritivore (Sars 1895; Stephensen 1938). Northeast Peak of Georges Bank (southern range exten- sion), 192 m, sand bottom. Superfamily Leucothoidea Family Pleustidae Neopleustes pulchellus Kr^yer, 1846. Arctic-Boreal regions of North Atlantic; in western Atlantic: Arctic south to Georges Bank; 40-800 m, rock and gravel bottoms, epiben- thic (Sars 1895; Stephensen 1938; Bousfieid 1973). Northeast Peak of Georges Bank, 84-108 m, gravel bottom. Pleustes panoplus Kr«)yer, 1838. Circumpolar; in western Atlantic: Arctic south to New England coast; 10-280 m, sand and gravel bottoms, epibenthic (Sars 1895; Stephen- sen 1938). Southern Georges Bank and Northeast Peak, 71-103 m, gravel to sand bottoms, 7.2°-7.7°C. Pleusymtes glaber Boeck, 1861. Boreal regions of North Atlantic; in western Atlantic: Labrador to Chesapeake Bay; 5-120 m, rocky bottoms (Gurjanova 1951; Bousfieid 1973). Great South Channel and Northeast Peak of Georges Bank, 48-146 m, sand and gravel bottoms, 7.4°-8.9°C. Slenopleusles gracilis Holmes, 1905. Western Atlantic: Cape Cod south to Chesapeake Bay, 5-50 m, sand bottoms (Bousfieid 1973; Dickinson et al. 1980). Great South Channel and Northeast Peak of Georges Bank, 58-103 m, sand and gravel bottoms, 7.1°-8.4°C. Slenopleusles inermis Shoemaker, 1949. Western Atlantic: Gulf of Maine south to Delaware Bay; 5-100 m, sand and silty sand bottoms, epibenthic (Bousfieid 1973; Dickinson et al. 1980). Southern Georges Bank, 66-95 m, sand bottom. Family Amphilochidae A mphilochoides odonlyx (Boeck, 1871). Boreal regions of North Atlantic; in western Atlantic: Arctic south to New Jersey; 20-120 m, sand to silt-clay bottoms (Sars 1895; Stephensen 1938; Watling 1979). Southern Georges Bank, 84 m, gravel bottom, 10.4°C. Gitanopsis arctica G. O. Sars, 1895. Arctic-Boreal regions of North Atlantic; in western Atlantic: Arctic south to Gulf of St. Lawrence; 10-100 m, silt-clay bottoms (Sars 1895; Stephensen 1938). Northern Edge of Georges Bank (southernmost record in western Atlantic), 64 m. Family Stenothoidae Slenothoe minuta Holmes, 1905. Western Atlantic: Cape Cod south to Georgia; 0-40 m, usually associated with hydroids and bryozoans (Bousfieid 1973). Southern Georges Bank, 64-73 m, sand bottom. Slenula peltata (Smith, 1873). Western Atlantic: Arctic to Gulf of St. Lawrence; 50-200 m (Gosner 1971; Bousfieid 1973). Perimeter of Georges Bank, 70-179 m, sand. Superfamily Stegocephaloidea Family Stegocephalidae Andaniopsis nordlandica (Boeck, 1871). Boreal regions of North Atlantic; not previously reported in western Atlan- tic; 40-600 m, epibenthic (Sars 1895; Stephensen 1935). Southern Georges Bank, 84 m, gravel bottom, 10.5 °C. Stegocephalus inflatus Krciyer, 1842. Circumpolar Arctic; in western Atlantic: Arctic south to Rhode Island; 30-600 m, gravel to clay bottoms, epibenthic, predator? (Sars 1895; Stephensen 1935; Bousfield 1973). Perimeter of Georges Bank, 121-196 m, sand and silt- clay bottoms, 6.6°-7.6°C. Family Acanthonotozomatidae AcanthonotozotJia serralum (Vabnc'ms, 1780). Circumpolar Arctic; in western Atlantic: Arctic south to Bay of Fundy; 10-300 m, gravel to silt-clay bottoms, epibenthic (Sars 1895; Stephensen 1938; Gurjanova 1951). Northeast Peak of Georges Bank (southern range exten- sion in western Atlantic); 79-198 m, sand and gravel bot- toms, 7.8°-8.2°C. Superfamily Synopioidea Family Synopiidae Syrrhoe crenulala Goes, 1866. Boreal regions of North At lantic and North Pacific Oceans; in western Atlantic: Arc- tic south to Cape Cod; 40-200 m, epibenthic detritivore (Sars 1895; Barnard 1972). Northeast Peak of Georges Bank, 77-101 m, sand bot- tom, 8.4 °C. Tiron spiniferum (Stimpson, 1853). Circumboreal; in west- ern Atlantic: Arctic south to New England coast; 30-200 m, epibenthic (Sars 1895; Barnard 1972). Northeast Peak of Georges Bank, 93-198 m, gravel and gravelly sand, 7.7°-8.4°C. Family Argissidae Argissa hamatipes {^^oxm&n, 1869). Northern Hemisphere, subarctic to warm temperate; in western Atlantic: Labra- dor south to North Carolina, 4-1,720 m, gravel to silt-clay bottoms (Barnard 1971; Bousfield 1973). Southeast Part of Georges Bank, 60-82 m, sand bottom. Superfamily Pardaliscoidea Family Pardaliscidae Halice abyssi Boeck, 1871. Arctic-Boreal regions of North Atlantic; not previously recorded in western Atlantic; 200-800 m, mud bottoms (Sars 1895; Stephensen 1938). Western Basin of Georges Bank, 168-183 m, silt-clay bottom, 6.6°-7.2°C. Pardalisca cuspidata Krbyer, 1842. Arctic-Boreal regions of North Atlantic; in western Atlantic: Arctic south to New England; 10-750 m, sand to silt-clay bottoms (Sars 1895; Gurjanova 1951; Bousfield 1973). Northeast Peak of Georges Bank, 132 m. Superfamily Ampeliscoidea Family AmpeHscidae Arnpelisca agassizi (Judd, 1896). Atlantic and Pacific coasts of North America; in western Atlantic; Nova Scotia south to Caribbean; 5-450 m, coarse sand to silt-clay bottoms, detritivore, tubedweller (Mills 1971; Dickinson et al. 1980). Southern Georges Bank, 60-170 m, sand bottoms, 6.6°- 12.6°C, often occurs in high densities (100-10,000/m-). Arnpelisca declivitatis Mills, 1967. Northwestern Atlantic: western Greenland south to North Carolina; 100-1,000 m, sand to silt-clay bottoms, probably a tube-dwelling detriti- vore (Mills 1971; Dickinson et al. 1980). Western Basin of Georges Bank, 150-170 m, sand and silty-sand bottoms. Arnpelisca macrocephala Liljeborg, 1852. Arctic-Boreal in North Atlantic, North Pacific, and Arctic Oceans; in western Atlantic: Greenland south to New Jersey; 5-1,700 m, stable sand bottoms, detritus feeding tubedweller (Bar- nard 1971; Mills 1971; Bousfield 1973). Perimeter of Georges Bank, 66-168 m, sand bottoms, 7.3°-12.2°C. Arnpelisca vadonim Mills, 1963. Western Atlantic: Gulf of St. Lawrence to Gulf of Mexico; 0-l(X) m, medium to coarse sand, detritus feeding tubedweller (Mills 1967; Bousfield 1973; Dickinson et al. 1980). North and central Georges Bank, 20-80 m, coarse and medium sand, 7.6°-8.4°C. Byblis gaimardi (Kreiyer, 1846). North Atlantic and Arctic Oceans; in western Atlantic: Greenland south to Cape Cod; 5-575 m, sand bottoms, detritus feeding tubedweller (Mills 1971). Western Basin of Georges Bank, 154 m, sand bottom. Byblis serrata (Smith, 1 873). Western Atlantic: Cape Cod to Cape Hatteras; 0-200 m, medium to coarse sand, detritus feeding tubedweller (Mills 1971; Dickinson et al. 1980). Southern Georges Bank, 44-123 m, sand bottom, 7.3°- 11.6°C, moderately abundant (10-500/m'). Haploops tubicola Liljeborg, 1856. Circumpolar Boreo- Arctic species; in western Atlantic: Arctic south to Gulf of Maine; 10-400 m, sand to silt-clay bottoms, detritus feed- ing tubedweller (Sars 1895; Stephensen 1935; Mills 1971). Western Basin and perimeter of Georges Bank, 79-179 m, sand to silt-clay bottoms, 6.3°-7.4°C. Superfamily Melphidippoidea Family Melphidippidae Melphidippa goesi Stebbing, 1899. Boreal regions of North Atlantic; in western Atlantic: Labrador south to Gulf of Maine; 30-300 m, fUter feeder (Sars 1895; Stephensen 1938; Enequist 1949). Western Basin and Northern Edge of Georges Bank, 157-183 m, sand and silt-clay bottoms, 6.6°-7.6°C. Family Hornelia — Cheirocratus group Casco bigelowi (Blake, 1929). Western Atlantic: Gulf of St. Lawrence south to Maryland; 0-400 m, sand to silt-clay bottoms, epibenthic, omnivore (Bousfield 1973; Dickinson et al. 1980). Perimeter of Georges Bank, 68-179 m, sand and silt-clay bottoms, 5.3°-12.8°C. Superfamily Melitoidea Family Melitidae Eriopisa elongata (Bruzelius, 1859). Arctic-Boreal regions of North Atlantic and North Pacific; in western Atlantic: Cape Cod south to New Jersey; 60-1,200 m, sand to silt- clay bottoms, burrowing deposit feeder (Sars 1895; Ene- quist 1949; Barnard 1971; Dickinson et al. 1980). Southwest Georges Bank, 92 m, sand bottom. Maera danae Slimpson, 1853. Western Atlantic: Gulf of St. Lawrence south to New Jersey; 0-50 m, gravel and silt-clay bottoms, epibenthic omnivore (Bousfield 1973). Northern Edge of Georges Bank, 103 m, sand bottom. Melita dentata Krbyer, 1842. Arctic-Boreal regions of North Atlantic and Arctic Oceans; in western Atlantic: Arctic south to Maryland; 0-300 m, gravel and sand bot- toms, epibenthic omnivore (Bousfield 1973). Perimeter of Georges Bank, 46-183 m, gravel and sand bottoms, 7.1°-11.6°C. Melita sp. A. (undescribed species). Western Atlantic: Cape Cod south to New Jersey; 70-100 m, sand and silty sand bottoms (Dickinson et al. 1980). Southern edge of Georges Bank, 92-114 m, sand bottom. Melita sp. B. (undescribed species). Northeast Peak of Georges Bank, 86-194 m, gravel and gravelly sand. Superfamily Corophioidea Family Photidae Gammaropsis maculatus (Johnston, 1827). Boreal regions of North Atlantic; in western Atlantic: Arctic south to North Carolina; 20-200 m, gravel and gravelly sand, detri- tus feeders (Sars 1895; Bousfield 1973; Fox and Bynum 1975). Northeast Peak of Georges Bank; 79-93 m, gravel bottoms. Photis dentata Shoemaker, 1945. Western Atlantic: Maine south to Florida; 49-92 m, sand bottom, probably a tube- dwelling detritivore (Shoemaker 1945a; Dickinson et al. 1980). Western portion of Georges Bank, 35-124 m, sand and gravelly sand bottoms. Photis macrocoxa Shoemaker, 1945. Western Atlantic: Gulf of St. Lawrence south to Virginia; 0-100 m, sand and silt bottoms, probably a tube-dwelling detritivore (Bousfield 1973). Central and southern Georges Bank; 33-93 m, sand bottom. Podoceropsis nitida (Stimpson, 1853). Boreal regions of North Atlantic; in western Atlantic: Gulf of St. Lawrence south to New Jersey; 0-50 m, primarily on rocky bottoms, probably a detritivore (Bousfield 1973; Watling 1979). Perimeter of Georges Bank, 35-179 m, gravel and sand bottoms, 4.9°-I2.3°C. Protomedeia fasciata Krciyer, 1842. Arctic-Boreal regions of North Atlantic; in western Atlantic: Arctic south to New Jersey; 15-115 m, sand and gravelly sand, burrowing detri- tivore (Sars 1895; Enequist 1949). Perimeter of Georges Bank, 62-146 m, gravel to sand bottoms. Family Ischyroceridae Ericthonius ntbricornis Smith, 1873. Boreal regions of North Atlantic; in western Atlantic: Labrador south to Cape Hat- teras; 0-400 m, gravel to silt-clay bottoms, filter-feeding tubedweller (Enequist 1949; Bousfield 1973; Dickinson et al. 1980). Perimeter of Georges Bank, 46-192 m, gravelly sand and sand bottoms, moderately abundant along southern edge of Georges Bank reaching densities of 500/m! Ischyrocerus angtiipes KreSyer, 1838. Subarctic and Boreal regions of North Atlantic; in western Atlantic: Hudson Strait south to Cape Hatteras; 0-60 m, primarily on hard substratum, tubedweller (Bousfield 1973). Perimeter of Georges Bank, 35-94 m, sand and gravel bottoms. Ischyrocerus megacheir (Boeck, 1871). Arctic-Boreal re- gions of the North Atlantic; in western Atlantic: Arctic south to Gulf of St. Lawrence; 80-1,400 m, gravel and silt- clay bottoms, tube-dwelling detritivore (Sars 1895; Stephensen 1942; Enequist 1949). Northeast Peak and Western Basin of Georges Bank (southernmost records), 112-196 m, gravel and gravelly sand bottoms. Family Amphithoidae Amphithoe rubricaia (Montagu, 1808). Amphi-Atlantic; in western Atlantic: Arctic south to Long Island Sound; 0-40 m, algae bottoms, tubedweller, herbivore (Stephensen 1942; Enequist 1949; Bousfield 1973). Southeastern Georges Bank (unusual record, normally confined to nearshore area), 62 m. FamUy Aoridae Leptocheirus pinguis (Stimpson, 1853). Western Atlantic: Labrador south to North Carolina; 0-250 m, sand to silt- clay bottoms, tubedweller, filter-feeding detritivore (Bous- field 1973; Dickinson et al. 1980). Perimeter of Georges Bank, 44-179 m, sand and gravelly sand bottoms, 7.I°-12.2°C, moderately abundant on Georges Bank often reaching densities of 400/m'. Pseudunciola obliquua (Shoemaker, 1949). Western Atlan- tic: Bay of Fundy south to Virginia; 0-100 m, sand-gravel to silty sand, tubedweller, detritivore (Bousfield 1973; Dickinson et al. 1980). Southern Georges Bank, 64-103 m, sand and shelly sand. Unciola dissimilis Shoemaker, 1945. Western Atlantic: Massachusetts to North Carolina; 0-1, (XX) m, sand and silty sand bottoms, tube-dwelling detritivore (Shoemaker 1945b; Bousfield 1973). Western Georges Bank, 20-135 m, sand bottoms, 4.4°- 11.6°C. Unciola inennis Shoemaker, 1945. Western Atlantic: Bay of Fundy south to North Carolina, 0-200 m, sand and silty- sand bottoms, tube-dwelling detritivore (Shoemaker 1945b; Dickinson et al. 1980). Perimeter of Georges Bank, 57-192 m, sand and gravelly sand bottoms, 7.1 °-11.2°C, high densities between 10 and 2,000/m-. Unciola irrorata Say, 1818. Western Atlantic: Gulf of St. Lawrence south to South CaroHna; 6-500 m, sand-gravel to silt-clay bottoms, tube-dwelling detritivore (Shoemaker 1945b; Bousfield 1973; Dickinson et al. 1980). Central Georges Bank, 35-168 m, gravel and sand bot- toms, 7.1°-12.6°C, moderate densities 10-500/m'. Unciola spicata Shoemaker, 1945. Western Atlantic: Long Island south to Florida; 40-8(X) m, sand and silty sand bot- toms (Shoemaker 1945b; Dickinson et al. 1980). Southern edge of Georges Bank, 114-124 m, sand bottoms. Family Corophiidae Corophium crassicorne Bruzelius, 1859. Arclic-Boreal regions of North Atlantic and North Pacific; in western Atlantic: Arctic south to Virginia, 0-200 m, gravel to silty sand, tubedweller, filter-feeding detritivore (Bousfield 1973; Dickinson et ai. 1980). Perimeter of Georges Bank, 35-146 m, gravel and sand bottoms, 8.3°-12.3°C. Siphonoeceies smilhianus Rathbun, 1908. Western Atlan- tic: Cape Cod south to Maryland, 10-90 m, sand bottoms, tube-dwelling detritivore (Bousfield 1973; Dickinson et al. 1980). Southwest Georges Bank, 66 m, sand bottom. Species Distribution Patterns None of the 97 species collected were ubiquitous over the entire Georges Bank region, all of them were restricted in their distribution to some degree. In comparing the spatial patterns of gammarideans on the bank, it became apparent that many spe- cies had similar patterns. One of si.\ patterns could be used to characterize nearly all the species (Fig. 16). Although these pat- terns are idealized and not followed exactly by each species, they were useful in simplifying the data and in providing insights into which environmental factors may be important in controlling gammaridean distributions on Georges Bank. A discussion of each of these patterns and the species described by it follows. Family Podoceridae Dyopedos arclicus (Murdoch, 1884). Boreo-Arctic regions of North Atlantic and North Pacific; in western Atlantic: Arctic south to Cape Cod; 10-75 m, filter feeder (Laubitz 1977). Northeast Peak of Georges Bank, 99 m, sand bottom, 8.9°C. Dyopedos inonacaniha (Metzger, 1875). Boreal regions of North Atlantic; in western Atlantic: Arctic south to Vir- ginia; 20-300 m, gravel to silt-clay bottoms, filter feeder (Laubitz 1977; Dickinson et al. 1980). Perimeter of Georges Bank, 35-187 m, sand and gravel bottoms, 6.8°-11.3°C. Dulichia tuberculata Boeck, 1870. Boreo-Arctic regions of North Atlantic; in western Atlantic: Arctic south to Cape Cod; 10-600 m, rock and gravel bottoms, filter feeder (Ste- phensen 1935; Laubitz 1977). Northern Edge of Georges Bank, 156 m, sand bottom. Paradulichia typica Boeck, 1870. Arctic-Boreal regions of North Atlantic; in western Atlantic: Arctic south to Cape Cod; 60-1,100 m, silt-clay bottoms, filter feeder (Gurja- nova 1951; Laubitz 1977). Western Basin of Georges Bank, 157-187 m, sand and silt -clay bottoms, 6.6°-6.8°C. DISCUSSION Zoogeography Georges Bank lies in the latitudes where the Boreal (cold-tem- perate) and Virginian (warm-temperate) faunal provinces over- lap (Bousfield 1973; Briggs 1974). In his study of the inshore gammaridean fauna of the Cape Cod region, Bousfield (1973) found the Virginian forms composed two-thirds of his species list. In contrast, we found that two-thirds of the species on Georges Bank were Boreal forms having their centers of distri- bution in the cold waters north of Cape Cod. However, it should be noted that the shallow central portions of the bank are chiefly inhabited by Virginian species, and that nearly all the Boreal forms are restricted to the perimeter of the bank. Therefore, our results are in agreement with Bousfield's findings. Briggs (1974), in his discussion of the zoogeography of the northwest Atlantic, summarized data for both fish and inverte- brates which showed a clear trend for cold-temperate forms to be amphi-Atlantic while most warm-temperate species are endemic to the western Atlantic. The data presented in Bous- field (1973), Dickinson et al. (1980), and this report indicate that this distributional pattern is also true for gammarideans. Northeast Peak A group of six species (Table 1) showed a strong association with gravel sediments and cold year-round temperatures (6°- 8°C) of the Northeast Peak. This group of species showed a high fidelity to the Northeast Peak with only Anonyx sarsi Table 1. — Six pallerns of gammaridean distribution on Georges Bank. Species in parentheses occurred in only one or two collections. Northeast Peak Acanlhonotozonia serratum A nonyx sarsi (Dyopedos arclicus) Gammaropsis maculalus \felita sp. B Neopleustes pulchellus (Pardalisca cuspidala) Tiron spiniferum ( Tryphosella nanoides) ( H'estwoodi/ta magalops) Perimeter Ampelisca macrocephah Anonyx liljeborgi Casco bigelowi Corophium crassicorne Dyopedos inonacantha Ericihonius ruhricornis Harptnia proptnqua Hippomedon propinquus Leptocheirus pinguis Melita denlala Orchomene pinguis Photis denlala Phoxocephalus holbolli Pleusymies glaher Podoceropsis niiida Prolomedeta fasciala Stenopleusles gracilis Western Basin Ampelisca declivilatis (Anonyx dehruyni) Balhyniedon sausserei {Byblis gaimardi) (Dulichia tuberculata) Epimeria loricata (Eusirus cuspidatus) Halice abyssi Halirages fulvocinclus Haliragoides inerniis Haploops lubicola Ischyrocerus megacheir Melphidippa goesi Monoculopsis longicornis Paradulichia lypica Rhachotropis dtslincla Slegocephalus inflatus Ttnelonyx cicada Shoals Ampelisca vadorutn Amphiporea giganlea Pontogeneta inertnis Psammonyx nobills Southern Edge Ampelisca agassizi Argissa hamatipes (Eriopisa elongata) (Gammarus annulatus) Melita sp. A Orchomene mlnula Pleustes panoplus Rhachotropis inflala Rhachotropis oculata Stenopleusles inerniis (Synchelidium amerlcanum) Central A canthohuustorius intermedius A canthohuustorius similis A canthohuustorius splnosus Acanthohaustortus sp. C. Acanthohaustortus sp. D. Bathyporeia quoddyensls Sybils serrala Hippomedon serralus Monoculodes edwardsi Parahaustorlus holmesl Parahaustorliis longtmerus Paraphoxus epislornus Photis macrocoxa Protohaustorlus deichmannae Protohaustorius wigleyi Pseudohaustorius borealis Pseudunciola obliquua Stenothoe minuta Unclola Irrorata occurring outside this area. These species are all Arctic-Boreal and probably limited to the perimeter by their temperature tolerance, but their preference for gravelly sediments probably accounts for their further restriction to the Northeast Peak. A group of four species represented by single occurrences on the Northeast Peak are also listed in Table 1 as other species which might show this distribution pattern if more data were available. Perimeter A group of 17 species (Table 1) was distributed around the perimeter of the bank. The bottom temperatures around the perimeter generally have an annual range between 4° and 13°C, being warmer in the Great South Channel and cooler along the Northern Edge. The sediments of the perimeter are fine sands except for gravel on the Northeast Peak and medium to coarse sand in the Great South Channel. It is noteworthy that all 17, although having boreal distributions, are reported south of Georges Bank (Dickinson et al. 1980) in contrast to the species restricted to the Northern Edge and Western Basin. It seems likely that this group of species is excluded from the central parts of Georges Bank by the warmer summer temperatures (14°-18°C) and greater annual temperature range (3°-I8°C) found there. It is clear that sediment type may also play a role in restricting the distribution of some of these species since they are absent from the Northeast Peak or Great South Channel por- tions of the perimeter. Western Basin A group of 14 species (Table 1) was restricted to the Western Basin and Northern Edge. This area is characterized by the cold- est year-round temperatures (4°-6°C) and the finest sediments on the bank. Three species in the group (Haploops lubicola, Stegocephalus inflatus, and Tmelonyx cicada) had single records on the Southern Edge of the bank, but their distribu- tions seemed to be better described by a "Western Basin" pat- tern than a "Perimeter" pattern. Only 2 of the 14 species occurred south of Georges Bank, and they (Ampelisca declivita- tis and Tmelonyx cicada) were reported only from bathyal depths (Dickinson et al. 1980). Since these records constitute the southern limit for most of these species, we suspect that temper- ature limits their distribution on the bank to this area of cold year-round bottom temperatures. An additional group of four species are also listed in Table 1 as other species which might show this pattern if more data were available. Shoals Only a small group of species (Table 1) occurs in the gravelly sands of the northern shoals. This area of the bank is character- ized by strong currents that continually rework the sediments. The shoals also experience the greatest annual temperature range (4°-16°C). Three of the four species found in the shoals are known to be associated with coarse sands, and they are usually found in shallow turbulent sand communities of the open coast. Pontogeneia inermis is a pelagic-epibenthic species which is ubiquitous over the central parts of Georges Bank. It is probable that other gammarideans may be associated with the shoals, but we may have missed them in our limited collections from this part of the bank. Southern Edge A group of eight species (Table 1) was restricted in their distribution to the Southern Edge of the bank. This area is char- acterized by fine sands and a narrow annual temperature range fluctuating between 8° and 11°C. Since these species are all cold-water forms, it seems Hkely that distribution around the perimeter of Georges Bank is Umited by their preference for fine sands. Three additional species are listed in Table 1 since we sus- pect they might show this pattern if more data were available. Central A group of 19 species (Table 1) is associated with the medium sand sediments and warmer summer temperatures (12°-14°C) of the central portions of the bank. However, this group of species seems to avoid the northern shoals area of the bank. These species all have Virginian distributions and are endemic to the western Atlantic. Some species in this group such as mem- bers of the family Haustoriidae have limited distributions on the bank, probably reflecting their need for special sediment charac- teristics. Other species in this group such as Unciola irrorata and Monoculodes edwardsi are broadly distributed in the central region of the bank. In general, we suspect that this species group is restricted from the perimeter of the bank by the cold year- round temperatures and from the shoals by the turbulence. ACKNOWLEDGMENTS We thank our associates at the Northeast Fisheries Center in Woods Hole, particularly Bruce Burns, Gilbert Chase, Evan Haynes, Tom Morris, Betty Murray, Ruth Stoddard, and Henry Jensen for assistance in collecting and processing samples; Roger Theroux and Jacqueline Murray for their help in data processing; John Lamont for his assistance in drafting the figures; and Richard Langton for his critical reading of the man- uscript. We would also like to thank E. L. Bousfield and Diane Laubitz of the National Museum of Canada, Ottawa, for their taxonomic assistance. LITERATURE CITED BARNARD. J. L. 1960. The amphipod family Phoxocephalidae in the Eastern Pacific Ocean, with analyses of other species and notes for a revision of the family. Allan Hancock Pac. Exp. 18:175-375. 1969. The families and genera of marine Gammaridean Amphipoda. U.S. Natl. Mus. Bull. 271, 353 p. 1971. Gammaridean Amphipoda from a deep-sea transect off Oregon. Smithson. Contrib. Zool. 61. 86 p. 1972. A review of the family Synopiidae ( = Tironidae). mainly distributed in the deep sea (Crustacea:Amphipoda). Smithson. Contrib. Zool. 124, 94 p. 1979. Revision of American species of the marine amphipod genus Para- phoxus (Gammaridea: Phoxocephalidae). Proc. Biol. Soc. Wash. 92:368-379. BOUSFIELD, E. L. 1965. The Haustoriidae of New England (Crustacea: Amphipoda). Proc. U.S. Natl. Mus. 117 (3512):159-239. 1970. Adaptive radiation in sand-burrowing amphipod crustaceans. Chesapeake Sci. 11:143-154. 1973. Shallow-water gammaridean Amphipoda of New England. Cor- nell Univ. Press, Ithaca. N.Y., 312 p. 1977. A new look at the Systematics of Gammaroidean Amphipods of the World. Crustaceana Suppi. 4:282-316. 1978. A revised classification and phylogeny of Amphipod Crustaceans. Trans. R. Soc. Can., Ser. IV, XVI:343-390. BRIGGS, J. C. 1974. Marine zoogeography. McGraw Hill Co., N.Y.. 475 p. COLTON, J. B., Jr., and R. R. STODDARD. 1973. Bottom-water temperatures on the Continental Shelf, Nova Scotia to New Jersey. U.S. Dep. Commer., NOAA Tech, Rep. NMFS Circ. 376, 55 p. CROKER. R. A. 1967. Niche diversity in five sympatric species of intertidal amphipods (Crustacea: Haustoriidae). Ecol. Monogr. 37:173-2(X). DICKINSON, J. J.. R. L. WIGLEY, R. D. BRODEUR, and S. BROWN- LEGER. 1980. Distribution of gammaridean Amphipoda (Crustacea) in the Middle Atlantic Bight region. U.S. Dep. Commer., NOAA Tech. Rep. NMFS SSRF-741, 46 p. ENEQUIST. P. 1949. Studies on the soft-bottom amphipods of the Skagerak. Zool. Bidr. Upps. 28:297-492. FOX. R. S., and K. H. BYNUM. 1975. The amphipod crustaceans of North Carolina estuarine waters. Chesapeake Sci. 16:223-237. FRAME. A. 1980. Two new species of sand burrowing Amphipod crustaeceans from Long Island Sound and the New York Bight (Amphipoda: Haustoriidae). Estuaries 3:75-83. GLENNON, T. A. 1979. Description of the male of Amphlporeia gigantea Bousfield (Amphipoda, Haustoriidae). Crustaceana 37:304-310. GOSNER, K. L. 1971. Guide to identification of marine and estuarine invertebrates. Cape Hatteras to the Bay of Fundy. Wiley-Interscience, NY., 693 p. GURJANOVA, E. F. 1951. Amphipoda-Gammaridea of the seas of the USSR and adjoining waters. Keys to the Fauna of the USSR. (In Russ.] Zool. Inst. Acad. Sci. USSR 41, 1029 p. GUSEY, W. F. 1977. The fish and wildlife resources of the Georges Bank Region. Shell Oil Co., Houston, Tex., 553 p. HAGER, R. P., and R. A. CROKER. 1979. Macroinfauna of Northern New England sand. IV. Infaunal ecology of Amphiporeia virgmiana Shoemaker, 1933 (Crustacea:Amphi- poda). Can. J. Zool. 57:1511-1519. HOLMES, S. J. 1905. The Amphipoda of Southern New England. Bull. (U.S.] Bur. Fish. 24:459-529. LAUBITZ, D. R. 1977. A revision of the genera Dulchia Krbyer and Paradulichia Boeck (Amphipoda, Podoceridae). Can. J. Zool. 55:942-982. MILLS, E. L. 1%7. A re examination of some species of Ampehsca (Crustacea: Amphi- poda) from the east coast of North America. Can. J. Zool. 45: 635-652. 1971. Deep-sea Amphipoda from the western North Atlantic Ocean. The Family Ampeliscidae. Limnol. Oceanogr. 16:357-386. SARS, G. O. 1895. An account of the Crustacea of Norway (with short descriptions and figures of all the species.) Vol. 1, Amphipoda. Alb Cammermeyers, Copenhagen, 71 1 p. SCOTT. K. J., and R. A. CROKER. 1976. Macroinfauna of northern New England marine sand. III. The ecology of Psammonyx nobths (Stimpson), 1853 (Crustacea: Amphipo- da). Can. J. Zool. 54:1519-1529. SHOEMAKER. C. R. 1930a. The lysianassid amphipod crustaceans of Newfoundland, Nova Scotia, and New Brunswick in the United States National Museum. Proc. U.S. Natl. Mus. 77(2827), 19 p. 1930b. The Amphipoda of the Cheticamp Expedition of 1917. Contrib. Can. Biol. 5:221-359. 1945a. The amphipod genus Photis on the east coast of North America. Charleston Mus. Leafi. 22:1-17. 1945b. The amphipod genus Unciola on the east coast of America. Am. Midi. Nat. 34:446-465. SMITH, S., and O. HARGER. 1874. Report on the dredgings in the region of St. George's Banks, in 1872. Trans, Conn. Acad. Arts Sci. 3:1-57. STEELE. D. H., and P. BRUNEL. 1968. Amphipoda of the Atlantic and Arctic coasts of North America: /lnon>':v (Lysianassidae). J. Fish. Res. Board Can. 25:943-1060. STEELE, D. H.. and V. J. STEELE. 1973. Some aspects of the biology of Calliopius laeviusculus (Krbyer) (Crustacea, Amphipoda) in the northwestern Atlantic. Can. J. Zool. 51:723-728. STEPHENSEN, K. 1935. The Amphipoda of N. Norway and Spitsbergen with adjacent waters. Troms* Mus. Skr. III(I):1-140. 1938. The Amphipoda of N. Norway and Spitsbergen with adjacent waters. Tromsb Mus. Skr. lll(ll):14I-278. 1940. The Amphipoda of N. Norway and Spitsbergen with adjacent waters. Tromsb Mus. Skr lII(lll):279-362. 1942. The Amphipoda of N. Norway and Spitsbergen with adjacent waters. Troms PWx'^' 'f€^ ^ ' >> J • J / "* v ^ • : • ^^ >' 100 M-^^ ^y^M^" 200 M-, ,.,-S' ^~' ^^ Aconthohausforius similis 1 1 1 - 42° 40° 100 M ^Oo200M,, Acanthohaustorius sp C / \ T L (r i \^ >'" "■i ' _^.. L a\\ "' 'U^ ^^ ^^' 'y 3 1 •^.^ v / •_ If / • • • 100 M^^^^ ^-— -^C^ ^^ zooM.r^^T^^r _ Aconthohausforius sp 1 1 D 1 - 42° 40° 40°- 100 M^^ ^ ^ .,^J"-^'"'' 200 M-, • Acanthonotozoma serratum ▲ Andaniopsis nordlandica 70° 68° 66° 1 • / /" ) / 4 ttf^ p^ ,' 100 M^^^ FaP" 200 M-^^^.,,' ^-' - Ampelisca 1 ogassizi 1 1 70° 68° - 42° - 40' 66° Figure 4. — Geographic dislribution of species of gammaridean Amphipoda on Georges Banii. 13 42= 40= 100 M 200 M- ▲ Ampelisca declivitatis • Ampelisca macrocephala 70° 68° 66° 1 1 I /• • •• •.. / / 100 M-^^ ^^y^jj>"'' 200M-, •k^"**" — Ampelisca vadorum 1 1 1 - 42° 40° 42° 40°- 100 M 200M.,_ Amphilochoides odontyx • Amphiporeia giganteo 1 ( /' " J' • 100 M /-— -""^^A^-^'"'' 200M-, __ _,_,r~"' • Amphiporeia virginiana A Amphithoe rubricata 1 42° 40° 42° U 1 1 100 M,..,^^^^^-^yU^'"'' 40° 200 M-, ,.,-.r~"' A /Inonyx debruyni 1 ^ 1 1 70° 68° 66° 100 M 200 M,, Anonyx liljeborgi _L 70° 68° - 42° - 40° 66° Figure S. — Geographic distribution of species of gammaridean Amphipoda on Georges Bank. — Continued. 14 70° 68° 66° 42' 40° 100 M ZOOM .^J-^'-"' • Argissa hamafipes ^ Bathymedon sausserei 70° 68° 66° 1 -'^ 1 J'- 100 M.^^^^ ^ — y^^^jj^"' zooM-T^T^r" — Bathyporeia 1 quoddyensis 1 1 42" 40" 100 M 40O200M-, A SyMs galmardi Byblis serrata 100 M ZOOM,, Calliopius laeviusculus 42' 40° 42° /i; 1 1 40° 100 M,,^^ ^/-^-'O-'^-^''' zooM-,_ _,_,r"' ' Cosco bigelowi 1 1 1 100 M ZOOM Corophium crassicome _l I - 42° - 40" 70° 68° 66° 70° 68° 66° Figure 6.— Geographic distribudon of species of gammaridean Amphipoda on Georges Banlt.— Continued. 15 42= 100 M 40°^°°^-- ,\jJ'-"' Dulichio tuberculata Dyopedos arcticus. - 42' 100 M 200 M Dyopedos monocanfho I I - 40° 42" 40' /I- 1 / 1 ( i" /f // 100 M ^.-^^jj'-' • 200M-, _ ,.j'"' — Epimeria loricata 1 1 1 100 M 200 M-, Ericfhonius rubricomis I l__ 42' 40" Erioplsa elongata Eusirus cuspidafus 70° 68° 66° C'^J'>"' 100 M. 200 M-^ • Gammaropsis maculatus A Gammarus annulatus I 1 70° 68° - 40° 66° Figure 7. — Geographic distribution of species of gammaridean Amphipoda on Georges Banli. — Continued. 16 42' 100 M 40° 200M., A Gitanopsis arctica • Halice obyssi L 100 M 200 M-, .^J'>"' Hallrages fulvoclnctus _l I 42° 40" 42° i; 1 1 \ ^ ^ \ \ 1 1 J 1 *^ \y /J ( /" 100 M^^ ^—^^,^-^-^'" • ^4 40° 200 M,^ _,-J'^~' Hallragoides Inermis 1 1 1 /U I 1 k V ^^"~""\ ^ M^^ J / "^^^ Vy J/ ^ •- 100 M — -^ ,/^J- '' 200M,,^ _,-,r~"' _ Haploops tubicola 1 1 1 - 42° 100 M 40°-?°°'^- Harpinia propinqua _J L 70° 68° 66° 100 M 200 M., Hippomedon propipquus 70° 68° 42° - 40° 66° Figure 8. — Geographic distribution of species of gammaridean Ampliipoda on Georges Banli. — Continued. 17 42= 40° - 100 M ZOOM- • Ischyrocerus angulpes A Ischyrocerus megacheir 42' - 40" 42' 40°- ^ ^.^ 1 1 • ^7-''"' 100 M,...^^^ J^- 200 M-, c^'*" — Leptocheirus pinguis 1 1 1 40" 100 M 200 M- A Maera danae • Me///o sq A 70° 68° 66° /i; 1 1 "^ 1 J // /^' 100 M,,.^^ ^'—^.'^■^■^^' ,/ 200M-, _,.,r~ _ • MeMo sp B ▲ Melphidip'pa goesi 1 i .._ 1 70° 68° 42° - 40° 66° Figure 9. — Geographic distribution of species of gammaridean Amphipoda on Georges Bank. — Continued. 18 42° 70° 68° 66° ^/f^ 1 "'* "^'^ / 100 M-...^^ ^ ^^^.f-^'"' 40° 200 M-,^ _,-^r~~' Monoculodes edwardsl 1 1 1 70° 68° 66° 1 /I' 1 ' ^)" '^-A \y 100 M..^^^^^ — ^J\J-^'-''' • 200 M-, ,.,-,r~~' • Monoculodes intermedius A Monoculodes latlnnanus 1 42° - 40' 40' 100 M, 200M,. A Monoculopsis longlcornis • Neopleustes pulchellus 42° /i; 1 I '^^^ \y • ./ _f_y^^ 100 M^^^^^^--^,,^^*>"' 40° 200M-,_ ,.,-S'"' Orchomene pinguis 1 1 1 70° 68° 66° /i; 1 1 m?.f^ \ » J / ^ \y 100 M.,^^^ ^ — ^ ,,^_/-^'"' 200M-, _.^r~"' — Parodul.ichio fypica 1 1 1 70° 68° - 42° - 40° 66° Figure 10. — Geographic distribution of species of gammaridean Amphipoda on Georges Bank. — Continued. 19 70° 68° 66° /i; 1 1 42° -- * v/ • /■' 100 M,.,^^^ ^.,—y^^}^j>''' 40° ZOOM,, _ _,.^r~~' Parahaustorius attenuafus 1 1 1 100 M 200 M-, A Parahaustorius holmesi • Parahaustorius longimerus - 42° - 40° Paraphoxus epistomus 100 M 200 M A Pardalisca cuspidate • Photis denfata - 42° - 40° 40° P/^o//s macrocoxa 70° 68° 66° V - 42° 100 M 200 M Phoxocephalus holbolli 70° 68° - 40° 66° Figure 11. — Geographic distribution of species of gammaridean Amphipoda on Georges Banlt. — Continued. 20 70° 68° 66° 42<^ 40° 100 M- 200 M-.^ A Platyishnopus sp. A • Pleusymtes glaber 100 M 200 M-^ Pleusfes panoplus 42° 40° 42° • • t V 1 100 M-,^^^ ^ — ^''^-'"■'''' --•^ 200M-,_ ,.-— .''~"' — Podoceropsis nitida 1 1 1 iia 1 ^ ) ) / / "^^ \ . If •• • ,/ • *i4i ^ ^ 100 M^..,^^ ^^^ ^y 200M-, _ _,_,r~' - Pontogenela inermis 1 1 42° - 40° 40°- 100 M 200 M Protohaustorius deichmonnae _J \ L 70° 68° 66° 100 M 200 M-. Protohaustorius wigleyl 70° 68° 42° - 40° 66° Figure 12.— Geographic distribution of species of gammaridean Amphipoda on Georges Bank.— Continued. 21 42° 40° - 70° 68° 66° L ('r ... 1 • • • ?•■■ ^-A \y • • / ^ 100 M — • y^y 200 M-, ,_,-S'^ _ Psammonyx 1 nobilis 1 1 - 42° - 40° 42° l\) 1 1 k CV .--.'X ~ V , V' 100 M..^^^^^^ ^ %J-'>'-''' 40° 200 M-. ,-— r~~' Pseudohausfohus borealis 1 1 i 40° 100 M, 200M, A Rhachotropis distlncta • Rhachotropis Inflata 70° 6B° 66° Figure 13. — Geographic distribulion of species of gammaridean Ampliipoda on Georges Bank. — Continued. 22 70° 68° 42° 100 M 40c200M, A Siphonoecetes smithianus • Stegocephalus mflatus 66° 68° 100 M 200 M-, V...J--"' Stenopleusfes gracilis _J \ 66° - 42° 40° Stenopleusfes inermis _l \ i; 1 \ \ 1 J / 100 M,^^^^ ^ y^J\J'-^'- 200 M,, .,-J'~~ _ • Stenothoe minuta A Stenula peltafa 1 - 42° - 40° • Synchelidium omencanum ▲ Syrrhoe crenulota 70° 68° 66° 100 M 200 M- r/Aon splniferum 70° 68° 42° - 40° 66° Figure 14. — Geographic distribution of species of gammaridean Amphipoda on Georges Bank. — Continued. 23 70° 68° 66° L 1^' 1 1 100 M„.^^ — - ^,%j> 200M-, ,„-S'~' — Tryphosella nanoides 1 1 1 - 42° - 40° 42° 100 M.,^^^ — -^ ^y\j->'-''' 1 / 40° 200 M-^ ,.--~r~~' Unciola dissimilis 1 1 1 - 42° - 40° 42° 40° 70° 68° 66° Figure 15. — Geographic distribulion of species of gammaridean Ampliipoda on Georges Banli. — Continued. 24 66° 42' 40' 100 M 200 M-, Northeast Peak M^/ ^fTl)- 100 M-,,^^^^^ ^ .^y^j^'''' ZOOM-, _,_,r''"' Central 1 1 1- 70° 42« - 40° 68° 66° Figure 16.— Most common dislribulion patterns of gammarideans on Georges Bank. 25 5 WHSE 04526 NOAA TECHNICAL REPORTS NMFS Circular and Special Scientific Report— Fisheries Guidelines for Contributors CONTENTS OF MANUSCRIPT First page. Give the title (as concise as possible) of the paper and the author's name, and footnote the author's afnUation, mailing address, and ZIP code. Contents. Contains the text headings and abbreviated figure legends and table headings. Dots should follow each entry and page numbers should be omitted. Abstract. Not to exceed one double-spaced page. Foot- notes and literature citations do not belong in the abstract. Text. See also Form of the Manuscript below. Follow the U.S. Government Priming Office Style Manual, 1973 edi- tion. Fish names, follow the American Fisheries Society Special Publication No. 6, A List of Common and Scientific Names of Fishes from the United States and Canada, third edition, 1970. Use short, brief, informative headings in place of "Materials and Methods:' Text footnotes. Type on a separate sheet from the text. For unpublished or some processed material, give author, year, title of manuscript, number of pages, and where it is filed — agency and its location. 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