ML072070390

NOAA Technical Memorandum NMFS-NE-179: Essential Fish Habitat Source Document: Winter Skate, Leucoraja Ocellata, Life History and Habitat Characteristics

ML072070390
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Site:	Oyster Creek
Issue date:	03/31/2003
From:	Packer D B, Vitaliano J J, Zetlin C A US Dept of Commerce, National Marine Fisheries Service, US Dept of Commerce, National Oceanographic and Atmospheric Administration
To:	Office of Nuclear Reactor Regulation
Davis J NRR/DLR/REBB, 415-3835
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NMSF-NE-179
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{{#Wiki_filter:C~~tO~0 Ilk'Ars 01F NOAA Technical Memorandum NMFS-NE-179 Essential Fish Habitat Source Document: Winter Skate, Leucoraja ocellata, Life History and Habitat Characteristics A§ < -r S- ý --"~- ~ ~U. S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service Northeast Region Northeast Fisheries Science Center Woods Hole, Massachusetts March 2003 Recent Issues in This Series: 155. Food of Northwest Atlantic Fishes and Two Common Species of Squid. By Ray E. Bowman, Charles E. Stillwell, William L. Michaels, and Marvin D. Grosslein. January2000. xiv+ 1 3 8 p., Ifig., 7 tables, 2 app. NTIS Access. No. PB2000-106735. 156. ProceedingsoftheSummer FlounderAgingWorkshop, 1-2 February 1999,Woods Hole, Massachusetts.ByGeorge R. Bolz, James Patrick Monaghan, Jr., Kathy L. Lang, Randall W. Gregory, and Jay M. Burnett. May2000. v + 15p., figs., 5 tables. NTIS Access. No. PB2000-107403. 157' Contaminant Levels in Muscle ofFour Species ofRecreational Fish from the New York Bight Apex. ByAshokD.Deshpande, Andrew F.J. Draxler, Vincent S. Zdanowicz, Mary E. Schrock, Anthony J. Paulson, Thomas W. Finneran, Beth L. Sharack, Kathy Corbo, Linda Arlen, Elizabeth A. Leimburg, Bruce W. Dockum, Robert A. Pikanowski, Brian May, and Lisa B. Rosman. June 2000. xxii + 99 p., 6 figs., 80 tables, 3 app., glossary. NTIS Access. No. PB2001-107346. 158. A Framework for Monitoring and Assessing Socioeconomics and Governance of Large Marine Ecosystems. By Jon G. Sutinen, editor, with contributors (listed alphabetically) Patricia Clay, Christopher L. Dyer, Steven F. Edwards, John Gates, Tom A. Grigalunas, Timothy Hennessey, Lawrence Juda, Andrew W. Kitts, Philip N. Logan, John J. Poggie, Jr., Barbara Pollard Rountree, Scott R. Steinback, Eric M. Thunberg, Harold F. Upton, and John B. Walden. August 2000. v+ 32 p., 4 figs., I table, glossary. NTIS Access. No. PB2001-106847. 159. An Overview and History ofthe Food Web Dynamics Program ofthe Northeast Fisheries Science Center, Woods Hole, Massachusetts. By Jason S. Link and Frank P. Almeida. October 2000. iv + 60 p., 20 figs., 18 tables, I app. NTIS Access. No. PB2001-103996. 160. Measuring Technical Efficiency and Capacity in Fisheries by Data Envelopment Analysis Using the General Algebraic Modeling System (GAMS): A Workbook. By John B. Walden and James E. Kirkley. October2000. iii + 15 p., 9 figs., 5 tables. NTIS Access. No. PB2001-106502. 161. Demersal Fish and American Lobster Diets in the Lower Hudson -Raritan Estuary. By Frank W. Steimle, Robert A. Pikanowski, Donald G. McMillan, Christine A. Zetlin, and StuartJ. Wilk. November2000. vii + 106 p., 2 4 figs., 51 tables.NTIS Access. No. PB2002-105456. 162. U.S. Atlantic and Gulf of Mexico Marine Mammal StockAssessments-2000. Edited by Gordon T. Waring, Janeen M. Quintal, and Steven L. Swartz, with contributions from (listed alphabetically) Neilo B. Barros, Phillip J. Clapham, Timothy V.N. Cole, Carol P. Fairfield, Larry J. Hansen, Keith D. Mullin, Daniel K. Odell, Debra L. Palka, Marjorie C. Rossman, U.S.Fish and Wildlife Service, Randall S. Wells, and Cynthia Yeung. November2000. ix + 303 p., 4 3 figs., 55 tables, 3 app. NTIS Access. No. PB2001-104091. 163. Essential Fish Habitat Source Document: Red Deepsea Crab,Chaceon (Geryon) quinquedens, Life History and Habitat Characteristics. By Frank W. Steimle, Christine A. Zetlin, and Sukwoo Chang. January 2001. v+ 27 p., 8 figs., I table. NTIS Access. No. PB2001-103542. 164. An Overview oftheSocialand EconomicSurveyAdministered duringRound 11 oftheNortheastMultispecies Fishery Disaster Assistance Program. By Julia Olson and Patricia M. Clay. December2001. v +69p.,3 figs., 18 tables, 2 app.NTIS Access. No. PB2002-105406. 165. A Baseline Socioeconomic Study of Massachusetts' Marine Recreational Fisheries. By Ronald J. Salz, David K.Loomis, Michael R. Ross, and Scott R. Steinback. December2001. viii + 1 2 9 p., I fig., 81 tables, 4 app. NTIS Access. No.PB2002-108348. 166. Report on the Third NorthwestAtlantic HerringAcoustic Workshop, University ofMaine Darling Marine Center, Walpole, Maine, March 13-14,2001. By William L. Michaels, editor and coconvenor, and Philip Yund, coconvenor. December2001. iv+ 18 p., 14 figs., 2 app. NTIS Access. No. PB2003-101556. , tAMOS&z m QE r -~rn: b.I NOAA Technical Memorandum NMFS-NE-179 This series represents a secondary level of scientific publishing. All issues.employ thorough internal scientific review; some issues employ external scientific review.Reviews are -by design -transparent collegial reviews, not anonymous peer reviews. All issues may be cited in formal scientific communications. I Essential Fish Habitat Source Document: Winter Skate, Leucoraja ocellata, Life History and Habitat Characteristics David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano National Marine Fisheries Serv., James J. Howard Marine Sciences Lab., 74 Magruder Rd., Highlands, NJ 07732 U. S. DEPARTMENT OF COMMERCE Donald L. Evans, Secretary National Oceanic and Atmospheric Administration Vice Admiral Conrad C. Lautenbacher, Jr., USN (ret.), Administrator National Marine Fisheries Service William T. Hogarth, Assistant Administrator for Fisheries Northeast Region Northeast Fisheries Science Center Woods Hole, Massachusetts March 2003 Editorial Notes on Issues 122-152, 163, and 173-179 in the NOAA Technical Memorandum NMFS-NE Series Editorial Production For Issues 122-152, 163, and 173-179, staff of the Northeast Fisheries Science Center's (NEFSC's) Ecosystems Processes Division have largely assumed the role of staff of the NEFSC's Editorial Office for technical and copy editing, type composition, and page layout. Other than the four covers (inside and outside, front and back) and first two preliminary pages, all preprinting editorial production has been performed by, and all credit for such production rightfully belongs to, the staff of the Ecosystems Processes Division.Internet Availability Issues 122-152, 163, and 173-179 have been copubl ished, i.e., both as paper copies and as Web postings. All Web postings are available at: www.ne/c.noaa.gov/nefsc/habitat/e/h. Also, all Web postings are in "PDF" format.Information Updating By federal regulation, all infornation specific to Issues 122-152, 163, and 173-179 must be updated at least every five years.All official updates will appear in the Web postings. Paper copies will be reissued only when and if new information associated with Issues 122-152, 163, and 173-179 is significant enough to warrant a reprinting of a given issue. All updated and/or reprinted issues will retain the original issue number, but bear a "Revised (Month Year)" label.Species Names The NMFS Northeast Region's policy on tile use of species names in all technical communications is generally to follow the American Fisheries Society's lists of scientific and common names forfishes (i.e., Robins etal. 199 P), mollusks (i.e., Turgeon et al. 19981), and decapod crustaceans (i.e., Williams et al. 1989c), and to follow the Society for Marine M ammalogy's guidance on scientific and common names for marine mammals (i.e., Rice 1 9 9 8 d). Exceptions to this policy occur when there are subsequent compelling revisions in the classifications of species, resulting in changes in the names of species (e.g., Cooper and Chapleau 1998c; McEachran and Dunn 1998').'Robins, C.R. (chair); Bailey, R.M.; Bond, C.F.; Brooker, J.R.; Lachner, E.A.; Lea, R.N.; Scott, W.B. 1991. Common and scientific names of fishes from the United States and Canada. 5th ed. Amer. Fish. Soc. Spec. Publ. 20; 183 p.'Turgeon, t).D. (chair); Quinn. J.F., Jr.: Bogan, A.L.; Coan, E.V.: Hochberg, F.G.; Lyons, W.G.; Mikkelsen, P.M.; Neves, R.J.; Roper, C.F.E.;Rosenberg, G;. Roth. B.: Scheltema, A.; Thompson, F.G.; Vecchione, M.; Williams, J.D. 1998. Common and scientific names of aquatic invertebrates from the United States and Canada: mollusks. 2nd ed. Amer. Fish. Soc. Spec. l'ubl. 26; 526 p.'Williams, A.B. (chair); Abele, L.G.; Felder, D.L.; Hobbs, H.H., Jr.; Manning, R.B.; McLaughlin, P.A.; Pdrez Farfante, 1. 1989. Common and scientific names of aquatic invertebrates from the United States and Canada: decapod crustaceans. Amer. Fish. Soc. Spec. Pub?.17: 77 p."Rice, D.W. 1998. Marine mammals of the world: systematics and distribution. Soc. Maar. Mammal. Spec. Pub/. 4: 231 p."Cooper, J.A.; Chapleau, F. 1998. Monophyly and interrelationships of the farnily Pleuronectidae (Pleuronectiformes), with a revised classification. Fish. Bull. (11S.) 96:686-726. 'Mc~achran, J.D.; l)unn, K.A. 1998. Phylogenetic analysis of skates, a morphologically conservative clade of elasmobranchs (Chondrichthyes: Rajidae). Copeia 1998(2):271-290. Page iii FOREWORD One of the greatest long-term threats to the viability of commercial and recreational fisheries is the continuing loss of marine, estuarine, and other aquatic habitats.Magnuson-Stevens Fishery Conservation and Management Act (October 11, 1996)The long-term viability of living marine resources depends on protection of their habitat.NMFS Strategic Plan for Fisheries Research (February 1998)The Magnuson-Stevens Fishery Conservation and Management Act (MSFCMA), which was reauthorized and amended by the Sustainable Fisheries Act (1996), requires the eight regional fishery management councils to describe and identify essential fish habitat (EFH) in their respective regions, to specify actions to conserve and enhance that EFH, and to minimize the adverse effects of fishing on EFH. Congress defined EFH as"those waters and substrate necessary to fish for spawning, breeding, feeding or growth to maturity." The MSFCMA requires NMFS to assist the regional fishery management councils in the implementation of EFH in their respective fishery management plans.NMFS has taken a broad view of habitat as the area used by fish throughout their life cycle. Fish use habitat for spawning, feeding, nursery, migration, and shelter, but most habitats provide only a subset of these functions. Fish may change habitats with changes in life history stage, seasonal and geographic distributions, abundance, and interactions with other species. The type of habitat, as well as its attributes and functions, are important for sustaining the production of managed species.The Northeast Fisheries Science Center compiled the available information on the distribution, abundance, and habitat requirements for each of the 'species managed by the New England and Mid-Atlantic Fishery Management Councils. That information is presented in this series of 38 EFH species reports (plus one consolidated methods report). The EFH species reports are a survey of the important literature as well as original analyses of fishery-JAMES J. HOWARD MARINE SCIENCES LABORATORY HIGHLANDS, NEW JERSEY SEPTEMBER 1999 independent data sets from NMFS and several coastal states. The species reports are also the source for the current EFH designations by the New England and Mid-Atlantic Fishery Management Councils, and understandably have begun to be referred to as the "EFH source documents." NMFS provided guidance to the regional fishery management councils for identifying and describing EFH of their managed species. Consistent with this guidance, the species reports present information on current and historic stock sizes, geographic range, and the period and location of major life history stages. The habitats of managed species are described by the physical, chemical, and biological components of the ecosystem where the species occur, Information on the habitat requirements is provided for each'life history stage, and it includes, where available, habitat and environmental variables that control or limit distribution, abundance, growth, reproduction, mortality, and productivity. Identifying and describing EFH are the first steps in the process of protecting, conserving, and enhancing essential habitats of the managed species. Ultimately, NMFS, the regional fishery management councils, fishing participants, Federal and state agencies, and other organizations will have to cooperate to achieve the habitat goals established by the MSFCMA.A historical note: the EFH species reports effectively recommence a series of reports published by the NMFS Sandy Hook (New Jersey) Laboratory (now formally known as the James J. Howard Marine Sciences Laboratory) from 1977 to 1982. These reports, which were formally labeled as 'Sandy Hook Laboratory Technical Series Reports, but informally known as"Sandy Hook Bluebooks," summarized biological and fisheries data for 18 economically important species. The fact that the bluebooks continue to be used two decades after their publication persuaded us to make their successors -the 38 EFH source documents -available to the public through publication in the NOAA Technical Memorandum NMFS-NE series.JEFFREY N. CROSS, (FORMER) CHIEF ECOSYSTEMS PROCESSES DIvISION NORTHEAST FISHERIES SCIENCE CENTER Page iv Page v Contents In tro d u c tio n .................................................................. -.......................................................................................... .......................... I L ife H isto ry ...................................................................................................................................................................................... I G eo g rap h ical D istrib u tio n ................................................................................................................................................................. 5 H a b ita t C h a ra c te ristic s ...................................................................................................................................................................... 6 S ta tu s o f th e S to c k s ........................................................................................................................................................................... 8 R e se a rc h N e ed s ................................................................................................................................................................................. 9 A ck n o w led g m en ts ............................................................................................................................................................................ 9 R e feren ces C ited ................................................................................ .............................................................................................. 9 Tables Table 1. Summary of habitat parameters for winter skate, based on the pertinent literature ........................................... 12 Table 2. Summary of habitat parameters for winter skate, based on the most recent NEFSC and state surveys .................. 14 Figures Figure 1. The winter skate, Leucoraja ocellata (M itchill 1815), m ale ..................................................................................... 16 F ig ure 2 .E gg case o f w inter skate ............................................................................................................................................... 17 Figure 3. Abundance (% occurrence) of the major prey items of winter skate collected during NEFSC bottom trawl surveys.. 18 Figure 4. Distribution of juvenile winter skate collected during winter NEFSC bottom trawl surveys ..............

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21 Figure 5. Distribution and abundance of juvenile winter skate collected during spring NEFSC bottom trawl surveys ........... 22 Figure 6. Distribution of juvenile winter skate collected during summer NEFSC bottom trawl surveys ................................ 23 Figure 7. Distribution and abundance of juvenile winter skate collected during fall NEFSC bottom trawl surveys ................ 24 Figure 8. Distribution and abundance of juvenile winter skate in Massachusetts coastal waters ........................................... 25 Figure 9. Distribution and abundance of juvenile and adult winter skate collected in Long Island Sound ............................. 27 Figure 10. Relative abundance catch/tow and percent occurrence for juvenile and adult winter skate in Long Island Sound ....... 28 Figure 11. Seasonal distribution and abundance of juvenile winter skate in the Hudson-Raritan estuary .............. ! ................ 29 Figure 12. Seasonal distribution and abundance of juvenile and adult winter skate in Delaware Bay .................................. ........ 31 Figure 13. Distribution of adult winter skate collected during winter NEFSC bottom trawl surveys ...................................... 35.Figure 14. Distribution and abundance of adult winter skate collected during spring NEFSC bottom trawl surveys ............... 36 Figure 15. Distribution of adult winter skate collected during summer NEFSC bottom trawl surveys ..................................... 37 Figure 16. Distribution and abundance of adult winter skate collected during fall NEFSC bottom trawl surveys .................... 38 Figure 17. Distribution and abundance of adult winter skate in Massachusetts coastal waters ................................................ 39 Figure 18. Seasonal distribution and abundance of adult winter skate in the Hudson-Raritan estuary .................................... 41 Figure 19. Spring/fall distributions of juveniles relative to bottom temperature, depth, and salinity based on NEFSC surveys ... 42 Figure 20, Distributions of juveniles relative to bottom temperature and depth based onMassachusetts inshore surveys ............ 44 Figure 21. Seasonal distributions of juveniles relative to temperature, depth, salinity, and DO in the Hudson-Raritan estuary ... 46 Figure 22. Seasonal distributions of juveniles and adults relative to temperature, depth, salinity, and DO in Delaware Bay ....... 50 Figure 23. Spring/fall distributions of adults relative to bottom temperature, depth, and salinity based on NEFSC surveys ........ 53 Figure 24. Distributions of adults relative to bottom temperature and depth based on Massachusetts inshore surveys ............. 55 Figure 25. NEFSC spring survey index of winter skate biomass and commercial landings of the seven species skate complex.. 57 Page 1 INTRODUCTION' The winter skate [Leucoraja ocellata (Mitchill 1815);formerly Raja ocellata, see McEachran and Dunn (1998);Figure 1] occurs from the south coast of Newfoundland and the southern Gulf of St. Lawrence to Cape Hatteras (Bigelow and Schroeder 1953; McEachran and Musick 1975; Scott and Scott 1988; McEachran 2002). Its center of abundance is on Georges Bank and in the northern section of the Mid-Atlantic Bight; in both areas it is often second in abundance to little skate (Leucoraja erinacea), a sympatric species (McEachran and Musick 1975).Immature winter skate are often confused with immature little skate, the distinctions are size-dependent (McEachran and Musick 1973; McEachran 2002).Number of tooth rows, length at maturity, and location of pelvic denticles are the characters most commonly used to differentiate the two species (Michalopoulos 1990).LIFE HISTORY EGGS The single fertilized egg is encapsulated in a leathery, amber to brown egg capsule which is deposited on the bottom (Figure 2). The capsules are rectangular in outline, with the upper and lower surfaces about equally convex and each corner of the capsule having a long slender horn (Vladykov 1936; Scott and Scott .1988;Figure 2). The anterior horns are nearly as long as the posterior horns and are equal in length to the capsule. The capsules range from 55-196 mm in length and 35-53 mm in width, and are smooth but marked with fine longitudinal striations (McEachran 2002).Bigelow and Schroeder (1953) report egg deposition to occur during summer and fall off Nova Scotia and, quoting Scattergood, probably in the Gulf of Maine as well. They also state that egg deposition continues into December and January off southern New England.JUVENILES The young are 112-127 mm TL at hatching (McEachran 2002) and are fully developed. ADULTS Female winter skates with fully formed egg capsules are more abundant during the summer and fall but it is possible that reproduction takes place to some degree throughout the year (Vladykov 1936; Scott and Scott 1988; McEachran 2002).AVERAGE SIZE, MAXIMUM SIZE, AND SIZE AT MATURITY Bigelow and Schroeder (1953) reported winter skate to have an average size of 76.2-86.4 cm TL. McEachran and Martin (1977) state, that they are one of the larger skates in the Gulf of Maine, with a maximum known size of 150 cm TL [Bigelow and Schroeder (1953) give a maximum length of 106.7 cm TL] with larger individuals more common at higher latitudes. The size at maturity increases with latitude (McEachran and Martin 1977). On Georges Bank andin the Gulf of Maine, individuals mature between 70-109 cm TL. However, in the Gulf of St. Lawrence they mature at a smaller size and do not reach as large a size as other populations (McEachran and Martin 1977). McEachran (1973), who studied skates from 1967-1970 that were collected from Nova Scotia and the Gulf-of Maine to Cape Hatteras, found that the all specimens > 78 cm TL were mature except for a male 88 cm TL; the smallest mature winter skate was a female 72 cm TL. Bigelow and Schroeder (1953) reported that winter skate does not mature until at least 63.5-66.0 cm TL. On the eastern Scotian Shelf, Simon and Frank (1998) found that female winter skate reached 50% maturity at about 75 cm TL.Based on the predictive equations from Frisk et al.(2001) and the Northeast Fisheries Science Center (NEFSC) survey maximum observed length of 113 cm TL, Lmat is estimated at 85 cm TL and Amat is estimated at 7 years (Northeast Fisheries Science Center 2000b).'FOOD HABITS Generally, polychaetes and amphipods are the most important prey items in terms of numbers or occurrence, followed by decapods, isopods, bivalves, and fishes (McEachran 1973; McEachran et al. 1976). Hydroids are also ingested (Avent et al. 2001). In terms of weight, amphipods, decapods and fish can be most important; fish are especially prevalent in the larger winter skate (Bowman et al. 2000; see also Garrison and Link 2000a and Tsou and Collie 2001a). Bigelow and Schroeder (1953) reported rock crabs and squid as favorite prey, other items included polychaetes, amphipods, shrimps,.and razor clams. The fishes that were 'eaten' included smaller skates, eels, alewives, blueback herring, menhaden, smelt, sand lance, chub mackerel, butterfish, cunners, sculpins, silver hake, and tomcod.McEachran (1973) studied skates collected from Nova Scotia to Cape Hatteras during 1967-1970; the following diet descriptions are from him and McEachran et al. (1976).Nephtys spp., Nereis spp., Lumbrineris. fragilis, Ophelia denticulata, and maldanids (mostly Clymenella torquata) were the most abundant polychaetes in the Mid-Atlantic Bight and Georges Bank stomachs. Nephtys spp., Page 2 Pectinaria sp., 0. denticulata, and Aphrodite hastata were the most frequently consumed prey in the Gulf of Maine and on the Nova Scotian shelf.Haustoriids, Leptocheirus pinguis, Monoculodes sp., Hippomedon serratus, ampeliscids, Paraphoxus sp., and Tmetonyx sp. were the most frequently eaten amphipods over the survey area. Crangon septemspinosa was the most abundant decapod in the diet. Cancer irroratus, Dichelopandalus leptocerus, Pagurus acadianus, and Hyas sp. were consistently eaten but in small numbers.Among the minor prey items included Cirolana (Politolana?) polita, which was the dominant isopod.Other isopods eaten included Chiridotea tuftsi and Edotea triloba, but they contributed little to the overall diet. The only identifiable bivalves eaten were Solemya sp. and Ensis directus. The most frequently eaten fish was sand lance, while yellowtail flounder and longhorn sculpin were occasionally eaten.In Smith's (1950) study in Block Island Sound, nekton were more important and isopods much less important than in the McEachran (1973) and McEachran et al. (1976) studies. The major prey items for winter skate in Block Island Sound included nekton, L. pinguis, Nephtys incisa, E. directus, C. septemspinosa, Nereis sp., C. irroratus, Lumbrineris sp. and Monoculodes edwardsi.Winter skate from Georges Bank had the most diverse diet and those from the Mid-Atlantic Bight the least diverse diet (McEachran 1973; McEachran et al.1976). There was no significant change in the diet with increase in skate size; however, the numbers of polychaetes gradually increased and amphipods gradually decreased with increasing skate size. The number of fish and bivalves also increased with predator size and the two taxa were a major part of the diet of skate > 79 cm TL.The ingestion of decapods was independent of skate size.There was also no indication of either diel or seasonal periodicity in intensity of feeding. In Passamaquoddy Bay there were no differences between the diets of small and large winter skate (Tyler 1972).The 1973-1990 NEFSC food habits database for winter skate [Figure 3; see Reid et al. (1999) for details]generally confirms the McEachran (1973) and McEachran et al. (1976) studies. Crustaceans made up > 50% of the diet for skate < 61 cm TL, while fish dominated the diet of skate > 91 cm TL. Overall crustaceans declined in importance with increasing skate size (includes both amphipods and decapods) while the percent occurrence of polychaetes increased with -increasing skate size until the skate were about 81 cm TL. Amphipods occurred more frequently than decapods until the skates were > 71 cm TL. Among the most frequently occurring prey species for almost all sizes of skate included the decapods C.septemspinosa and Cancer and pagurid crabs, the isopod Cirolana (= Politolana?) polita, and sand lance. The following is a detailed description of the diet from the NEFSC food habits database broken down by winter skate size class (Figure 3).For winter skate 21-30 cm TL, 74-84% of the diet consisted of crustaceans, with 38-43% of the diet consisting of identifiable amphipods. The most abundant amphipod species included Unciola irrorata, Byblis serrata, and H. serratus. Identifiable decapods made up 23-25% of the diet, most of which were species such as C. septemspinosa and C. irroratus. Identifiable polychaete species (9-13% of the diet) included Ampharete arctica. Identifiable isopod species (9% of the diet) included Cirolana (= Politolana?) polita.Nematodes, bivalves, and fish were included in the "other prey phyla" category (3-17% of the diet).For skate 31-40 cm TL, 72-76% of the diet consisted of crustaceans, with 37-39% of the diet consisting of identifiable amphipods. Major amphipod species included B. serrata, U. irrorata, H. serratus, and several unidentified haustoriids. Identifiable decapods made up 17-23% of the diet, most of which were C. septemspinosa and C. irroratus. Identifiable polychaetes (12-17% of the diet) included Scalibregma inflatum, L. fragilis, and unidentified maldanids. Identifiable isopods (5-8% of the diet) included Cirolana (= Politolana?) polita.Miscellaneous items (6-9% of the diet) included nematodes and bivalves. Among the identifiable fish present in the diet (3-4%) were sand lance, yellowtail flounder, and hakes.The percentage of crustaceans in the diet of winter skate 41-50 cm TL dropped to 62-69%, although identifiable amphipods still made up the major portion (33-35%) followed by decapods (14-22%). Identifiable polychaetes made up 19-23% of the diet; other prey species (including mollusca), 6-9% of the diet;identifiable isopods, 7% of the diet; and identifiable fish, 3-8% of the diet. All the major prey species (except for the lack of the polychaete S. inflatum) were similar to the 31-40 cm TL size class, with the additions of several more Unciola species, L. pinguis (an amphipod), unidentified pagurid crabs, and nephtyid polychaetes. The percent occurrence of crustaceans in the diet of winter skate 51-60 cm TL dropped further, down to 53-54%, with identifiable amphipods making up only 26-32% of the overall diet. Some of the dominant identifiable amphipods included Psammonyx nobilis, unidentified oedicerotids, H. serratus, and unidentified haustoriids. Identifiable decapods made up only 9-12% of the diet; C.septemspinosa was again the dominant decapod prey, followed by C. irroratus and pagurid crabs. Cirolana (=Politolana?) polita was again one of the major identifiable isopods, which all together made up 7-12% of the diet. The percent occurrence of identifiable polychaetes continued to increase in the diet, up to 26-29%; several of the more numerous species present were in the genera Nephtys and Nereis. Identifiable fish also increased in the diet, up to 6-13%, with sand lance the dominant species. Other prey phyla, including bivalves and nematodes, accounted for 9-11% of the diet.The percent occurrence of crustaceans in the diet continued to decline for winter skate 61-70 cm TL: down to 38-44%, with identifiable amphipods making up only Page 3 13-20% of the diet, while identifiable decapods made up 11-12%. Major amphipod species included M. edwardsi, U. irrorata, H. serratus, and unidentified haustoriids and oedicerotids. C. septemspinosci continued to be the dominant decapod prey, followed by Cancer and pagurid crabs. Identifiable isopods again made up 7-12% of the diet; Cirolana (= Politolana?) polita continued .to be one of the major prey species. The percent occurrence of identifiable polychaetes in the diet increased, up to 28-32%; species in the genera Nephtys and Nereis were again dominant. The percent occurrence of identifiable fish in the diet continued to increase also, up to 11-24%, most of which were sand lance. Nine percent of the diet consisted of identifiable mollusks, with bivalves being dominant.While the percent occurrence of crustaceans dropped to 29-36% for winter skate 71-80 cm TL, the percent occurrence of identifiable decapods was greater than the percent occurrence of amphipods: 11-13% versus 7-12%.The former were dominated by C. septemspinosa, Cancer and pagurid crabs, and D. leptocerus, while several haustoriid species and U. irrorata were some of the major amphipod prey. Identifiable isopods made up 8-9% of the diet, the dominant species continued to be Cirolana (=Politolana?) polita. Identifiable polychaetes (25-35% of the diet) included L. fragilis and several Nephrys and Nereis species. The percent occurrence of identifiable fish in the diet varied widely between the two sampling periods, from 16-36%, although sand lance was still the dominant species. Identifiable mollusks made up 9-10%of the diet, most of which were bivalves.Fish as prey items became increasingly important for winter skate 81-90 cm TL. They made up 29-42% of the overall diet. As usual sand lance were the dominant fish prey, other species ingested included other skate, longhorn sculpin, and silver hake. Crustaceans in the diet declined to 19-30%. The major identifiable decapod species (8-11% of the diet) continued to be C.septemspinosa and Cancer and pagurid crabs as well as pandalid shrimp and Ovalipes ocellatus. The major identifiable amphipod species (3-8% of the total diet)were several haustoriid species. Cirolana (= Politolana?) polita was once again the dominant identifiable, isopod (all isopods together made up 5-7% of the diet). Several Nephtys species were the major identifiable polychaetes ingested, all polychaetes together made up 22-28% of the diet. Bivalves, particularly of the familiy Solenidae, were the dominant identifiable molluscan prey ingested, with all mollusks together accounting for 7-17% of the diet.Identifiable fish made up >50% of the diet of winter skate 91-100 cm TL. Sand lance was the overwhelming dominant, some of the minor fish prey included silver hake, herring, and butterfish. Crustaceans were down to 12-23% of the diet'. Identifiable decapods made up 5-10%of the diet, C. septemspinosa, Cancer and pagurid crabs, D. leptocerus, and pandalid shrimp were some of the major decapods ingested. Identifiable amphipods made up only 4-5% of the total diet, with few conspicuous species.Identifiable polychaetes accounted for 10-13% of the diet, with the genus Nephtys the most notable. "Other prey.phyla" and identifiable mollusks together accounted for 10-12% of the diet, bivalves and nematodes dominated this category.Finally, identifiable fish made up > 60% of the diet of 101-110 cm TL winter skate from .the 1981-1990 NEFSC trawl surveys. Most were sand lance. Mollusks were 14% of the diet, polychaetes were 13% of the diet, and crustaceans were down to 11% of the diet.Using NEFSC data from 1977-1980, Bowman et al.(2000) found that in terms of percent weight, crustaceans were dominant in the diet of skate < 31-50 cm TL, while fish, mostly sand lance, were dominant in the diet of skate 51-110 cm TL. For skate < 31 cm TL, amphipods dominated, especially L. pinguis. For skate 31-50 cm TL, decapods dominated, especially C. septemspinosa and C.irroratus. On Georges Bank Tsou and Collie (2001a), using NEFSC dietary data from 1989-1990, also showed that fish, especially sand lance, were most important for winter skate > 50 cm TL. Other noted fish prey included sliver hake, mackerel, and herring (see also Tsou and Collie 2001b).Nelson (1993) calculated the predation impact of winter skate on their Georges Bank prey. Annual estimates of consumption for winter skate increased as they grew larger. Consumption ranged from 1.186 kg/fish/year for skate 40-49 cm TL to 5.528 kg/fish/year for skate 90-99 cm TL. The percentage of benthic production consumed by winter skate from 1969-1990 ranged from 11-43%. Nelson (1993) suggests that in relation to the total macrofauna production on Georges Bank, winter skate (along with little skate) consume <0.02% of the total. These results indicate that only a small to moderate proportion of benthic biomass vulnerable to skate predation is consumed by both winter and little skate, and their consumptive impact will be dependent on the levels of invertebrate biomass and/or production. PREDATORS AND SPECIES ASSOCIATIONS Winter skate is preyed upon by sharks, other skates, gray seals, and gulls (Scott and Scott 1988; Kaplan 1999).McEachran and Musick (1975) state that winter and little skate co-occurred significantly during 1967-1970 surveys from Nova Scotia to Cape Hatteras. Although winter and little skate are sympatric species with similar habitat requirements (except perhaps temperature preference), there does not appear to be a high degree of competitive interaction between them because they are positively correlated by abundance and where the two species are most abundant (Georges Bank) they have the most similar diets and highest diversity of assemblages of prey species (McEachran 1973; McEachran and Musick 1975; McEachran et al. 1976).Also, even though the two species do consume the same large taxonomic groups of benthic fauna Page 4 (amphipods, decapods, and polychaetes), winter skate predominately feeds on infaunal organisms while little skate feeds largely on epifauna (McEachran 1973;McEachran et al. 1976). McEachran (1973) and McEachran et al. (1976) show that large burrowing polychaetes and bivalves were consumed more frequently by winter skate and epifaunal decapods were eaten more frequently by little skate. Winter skate ate more burrowing amphipods, especially haustoriids and Trichophoxus epistomus while little skate consumed more surface dwelling amphipods such as Unciola sp., Dulichia (= Dyopedos) monacantha, ampeliscids and caprellids. The division of food resources between the skates is not complete because some winter skate ate large numbers of epifauna and some little skate consumed large numbers of infauna. Both species ate considerable numbers of L.pinguis and C. septemspinosa. Little skate occasionally fed on haustoriids, and deep burrowing polychaetes were regular prey items. The infaunal and. epifaunal preferences of the two skates may be more distinct in areas where they may coexist than in areas where they occur separately because in Delaware Bay (Fitz and Daiber 1963) little skate consumed relatively more infauna than it did in the areas sampled in Smith's (1950)study or the McEachran (1973) and McEachran et al.(1976) studies. Winter skate does not regularly occur in Delaware Bay (Fitz and Daiber 1963; see also Delaware Bay trawl surveys, below)..In addition, differences in the shape and size of the mouth and the number of tooth rows between the two species were used as evidence by McEachran and Martin (1977) to suggest that the sympatric populations of winter and little skate underwent character displacement in order to avoid direct competition for food resources. In sympatric. populations, winter skate has greater number of tooth rows in the upper jaw and a wider and less arched mouth, thus allowing them to feed more efficiently and deeper in the bottom than little skate. Little skate has a relatively smaller and more arched mouth with fewer tooth rows in the upper jaw.Using 1973-1997 NEFSC data from Nova Scotia to Cape Hatteras, as well as the same NEFSC food habits database discussed above, Garrison and Link (2000a)investigated the dietary guild structure of the fish community. Both small (10-30 cm TL) and medium (31-60 cm TL) sized winter skate belonged to the"Amphipod/shrimp eaters" group, along with little skate and cusk eel; prey included amphipods, polychaetes, shrimp, and zooplankton. The largest winter skate (61 to> 80 cm TL) were by themselves in a subgroup of"Piscivores" because, as described previously under the detailed description of the diet using the 1973-1990 NEFSC food habits database, their diet contained a high proportion of sand lance. This again shows that there is a trend toward increasing piscivory with size.The resilience of demersal fish assemblages on Georges Bank was investigated by Overholtz and Tyler (1985) using seasonal NEFSC trawl survey data from 1963-1978. Of the five assemblage species groups or associations present on Georges Bank in spring and fall throughout the survey period, winter skate belonged to the "Intermediate" and "Shallow" assemblage groups. In.the Shallow assemblage the other major species present besides winter skate included Atlantic cod, little skate, longhorn sculpin, yellowtail flounder, and haddock; in the Intermediate assemblage, little skate, red and silver hake, Atlantic cod, and haddock were some of the other major species present. Overholtz and Tyler (1985) considered winter skate to be a "resident" species, since they were only present in two out of the five assemblages in abundance. The Shallow assemblage covered most of Georges Bank in the spring and was slightly smaller in the fall. The Intermediate assemblage occurred mostly south of the Shallow assemblage and inside the southern edge of Georges Bank; it was somewhat larger in the fall, suggesting a migration of the species in this area to shallower water as the year progressed. The assemblages in the spring appeared to follow depth contours.Garrison (2000) and Garrison and Link (2000b) have also investigated spatial assemblages and trophic groups from the Georges Bank region. Using 1963-1997 NEFSC trawl survey data from Georges Bank, as well as the same NEFSC food habits database discussed above (Garrison and Link [2000b] used 1973-1997 data while Garrison[2000] used 1991-1997 data), they found that the major predator groups were consistent across decades, with the boundaries of the assemblages similar to Overholtz and Tyler (1985). Garrison (2000) investigated the spatial assemblages during spring and autumn. He found that during autumn, winter skate was in the assemblage found in the deep habitats on southern Georges Bank that also included spiny dogfish, butterfish, little skate, red hake, fourspot flounder, and yellowtail flounder. The main shallow portion of Georges Bank assemblage included 31 cm to > 80 cm TL winter skate, little skate, spiny dogfish, Atlantic cod, windowpane, winter flounder, and sea raven. In spring, the main shallow portion of Georges Bank assemblage included 31 cm to > 80 cm TL winter skate, haddock, Atlantic cod, and spiny dogfish. In terms of dietary guilds or trophic groups, the two studies had slightly different viewpoints, but the diets of winter skate in both studies were the same as discussed in the Food Habits section above. In the Garrison and Link (2000b)study, winter skate fell into two predator or feeding groups. The first was a "Bentho-pelagic" group, which included 31-60 cm TL winter skate, little skate, longhorn sculpin, and Atlantic cod. The diets of these species were the same as that discussed above for winter skate alone: shrimp such as pandalids and C. septemspinosa, and benthic invertebrates including polychaetes, gammarid amphipods, and bivalves. The second group was the"Skate" group, consisting of > 80 cm TL winter skate, pollock, and windowpane. Their prey was characterized by a combination of fish and benthic prey, with a high proportion of sand lance during the 1980s. Garrison (2000) had slightly different trophic groups. In autumn, Page 5 the largest (61 cm. to > 80 cm TL) winter skate were by themselves in a subgroup of the "Piscivorous" group, feeding on sand lance, silver hake, and Atlantic herring, as well as benthic invertebrates. Small and medium (10-60 cm TL) winter skate were also in the "Demersal predators" group, along with flatfish, haddocks, little skate, and thorny skate (Amblyraja radiata). Prey included gammarid amphipods, polychaetes, isopods, and Cancer crabs, as well as C. septemspinosa. During spring, 10-60 cm TL winter skate were in the "Shrimp/amphipod predators" group, along with hakes, longhorn sculpin, Atlantic cod, fourspot flounder, little skate, and thorny skate. Prey included gammarid amphipods, pandalids and C. septemspinosa, polychaetes, and Cancer crabs. Winter skate 61 cm to > 80 cm TL were again by themselves in a subgroup called the "Generalist" group, consuming bivalves, polychaetes, sand lance, and herring. The decline in importance of fish prey, 35% fish in the autumn and 16% in the spring, is probably related to seasonal movements of prey (Garrison 2000).On the Scotian Shelf, Scott (1989), using research trawl survey data from roughly 1970-1984 determined that winter skate was locally abundant but did not associate closely with any other species. However Mahon (1997), analyzing trawl survey data for the same region from 1970-1993, showed that winter skate whs associated with such species as longhorn sculpin, sea raven and winter flounder in the shallow waters of the Bay of Fundy and Sable Island.GEOGRAPHICAL DISTRIBUTION In Canada, winter skate are found in the Gulf of St.Lawrence, off northeastern Nova Scotia, and the offshore banks of Banquereau, Sable Island, and Western Bank[Bigelow and Schroeder 1953; McEachran and Musick 1975; see'also Strong and Hanke (1995) for the 1970-1993 distribution of winter skate in the Scotia-Fundy region; further information on winter skate distribution on the Scotian Shelf can be found in Simon and Frank (1996, 1998)]. The population in the southern Gulf of St.Lawrence may be isolated from populations along the rest of the east. coast (McEachran and Musick 1975).McEachran and Musick (1975) suggest that reports of little skate from the Gulf of St. Lawrence and most records of it from northern Nova Scotia probably refer to winter skate.Winter skate is considered common all around the Gulf of Maine from Nova Scotia to Cape Cod, except for the deep troughs, and is also common on Georges Bank (Bigelow and Schroeder 1953; McEachran 2002). It has.frequently been reported from the Bay of Fundy, and the coasts of Maine and Massachusetts (Bigelow and Schroeder 1953) and in Massachusetts Bay (Collette and Hartel 1988), as well as along the New Hampshire coast (Nelson et al. 1983). However, McEachran (2002) states that because of its close resemblance to the little skate, many of these records, as well as those for little skate from the same localities, are suspect..Their range extends from southern New England and down the Mid-Atlantic Bight to northern North Carolina (Bigelow and Schroeder 1953; Figures 4-7, 13-16).Previous authors have suggested that winter skate undertakes seasonal movements, especially in the southern part of its range, moving shoreward in autumn and offshore in summer (McEachran 1973; McEachran and Musick 1975; McEachran 2002), although this is not quite evident from the overall NEFSC bottom trawl surveys (Figures 4-7, 13-16; see descriptions below). In Passamaquoddy Bay, Tyler (1971) reported winter skate present from December to March while Huntsman (1922)stated it was abundant from May to November.McEachran (1973) suggests. this disparity may be due to a difference in the areas the two authors sampled. Tyler (1971) sampled the deeper waters of Passamaquoddy Bay while Huntsman (1922) did not specify the sampling depths. However Macdonald et al. (1984) determined winter skate to be both a regular and occasional resident in Passamaquoddy Bay, and juveniles were often found at beach sites during summer. Merriman and Warfel (1948)stated it is a permanent resident off southern New England although there are seasonal fluctuations in abundance; Bigelow and Schroeder (1953) mention that along the southern coast of New England it comes inshore near Woods Hole during the colder months. However during August of 1988 Michalopoulos (1990) found that winter skate dominated the inshore skate community off outer Cape Cod (little skate was the only'other skate other skate present, but was much less abundant). Schaefer (1967) found winter skate in the surf zone of Long Island during May, June, October, and November, while Gottschall et al. (2000), based on surveys from 1984-1994 (see Habitat Characteristics section, below), found their lowest abundances in Long Island Sound were in the months of July, August, and September. McEachran and Musick (1975) reported winter skate to be more abundant south of Delaware Bay during the winter, it has also been reported from Chesapeake Bay from December to April (Hildebrand and Schroeder 1928; Geer 2002).JUVENILES NEFSC bottom trawl surveys [see Reid et al. (1999)for details] captured juvenile (< 84 cm TL) winter skate year-round. (Note that winter and summer distributions are presented as presence/absence data, precluding a discussion of abundances.) In winter, juveniles were found from Georges Bank to Cape Hatteras, out to the 200 m depth contour (Figure 4). Concentrations were found off Long Island and southern New England; they were almost entirely absent from the Gulf of Maine. In spring they were also found from Georges Bank to Cape Hatteras, and were concentrated nearshore throughout the Mid-Atlantic Bight and southern New England as well as Page 6 in Cape Cod and Massachusetts Bays (Figure 5). Small numbers were also found along the coast of Maine and southwest Nova Scotia and near Browns Bank.Comparatively few were present in summer, with concentrations on Georges Bank and around Cape Cod (Figure 6). Some were also found near Penobscot Bay, Maine. Winter skate abundances in the fall were not as high as in the spring (Figure 7). In the fall they were collected from Georges Bank to the Delmarva Peninsula and were again concentrated along Long Island, southern New England, around Cape Cod, and on Georges Bank.Small numbers were again found along the coast of Maine and near Browns Bank.Both the spring and fall 1978-2002 Massachusetts inshore trawl surveys [see Reid et al. (1999) for details]show similar abundances and distributions of juveniles (Figure 8). The highest concentrations were found on the Atlantic side of Cape Cod and south and west of Martha's Vineyard (especially in spring) and south and northeast of Nantucket (also in spring). Large numbers were also found near Monomy Point in the fall. Other notable occurrences of winter skate were around Plum Island, Ipswich Bay, north of Cape Ann, near Nahant Bay (especially in the fall), in Cape Cod Bay, and in Nantucket Sound.The distributions and abundances of both juveniles and adults in Long Island Sound (Figures 9-10) as described by Gottschall et al. (2000) will be discussed in the Habitat Characteristics section.Occurrence of juveniles in the Hudson-Raritan estuary appears to have the same seasonal pattern noted by previous authors for other estuaries; i.e., they're generally absent from the estuary during the summer months. Juveniles were fairly well distributed throughout the Hudson-Raritan estuary in winter, spring and fall, and were most abundant in the winter and fall (Figure 11). In summer the few that were left were mostly confined to the deeper and warmer waters of the Ambrose Channel.The 1966-1999 Delaware Bay trawl surveys. (adults and juveniles combined; Figure 12) also confirm the seasonal trends noted previously for winter skate, although they were not very abundant in the Bay overall.They were almost completely absent in summer, and a few were caught in the fall, while the greatest numbers were found in the winter and spring. The skate were most abundant in the center of lower Delaware Bay, near the mouth (Figure 12).ADULTS NEFSC bottom trawl surveys [see Reid et al. (1999)for details] captured adult winter skate (> 85 cm TL)during all seasons (Figures 13-16). The numbers of adults in spring and fall were much lower than for juveniles of the same two seasons (winter and summer distributions are presented as presence/absence data, precluding a discussion of abundances). In winter, they were scattered from Georges Bank to North Carolina; very few occurred in the Gulf of Maine (Figure 13). In the spring, they were also found from Georges Bank to North Carolina but, as with the juveniles, were also distributed nearshore throughout the Mid-Atlantic Bight and along Long Island as well as around Cape Cod and Massachusetts Bays (Figure 14). Small numbers were also found along the coast of Maine and southwest Nova Scotia near Browns Bank. Few occurred in summer, being found mostly on Georges Bank, Nantucket Shoals, and near Cape Cod (Figure 15). In the fall, they were mostly confined to Georges Bank, near Nantucket shoals, and near Cape Cod, with very few found south of those areas (Figure 16).Adult little skate were collected in much fewer numbers than juveniles during the spring and fall Massachusetts inshore trawl surveys I(Figure 17). The greatest numbers were found on the Atlantic side of Cape Cod and, in spring, south of Nantucket. Very few adults were caught in the Hudson-Raritan estuary during spring and fall (Figure 18). Those few that were caught were concentrated around the Ambrose and Chapel Hill Channels.The seasonal distribution and abundance of both adults and juveniles in Delaware Bay were discussed previously (Figure 12).HABITAT CHARACTERISTICS Information on the habitat requirements and preferences of winter skate (based on both the pertinent literature and the most recent NEFSC and state surveys)are presented here and summarized in Tables 1 and 2.Winter skate generally ranges from the shoreline to 371 m, although it is most abundant at depths < 11l m (Bigelow and Schroeder 1953; McEachran 1973;McEachran and Musick 1975; McEachran 2002).Bigelow and Schroeder (1953) suggest that feware caught > 91 m. In the Gulf of Maine, they are most abundant 46-64 m but occasionally occur < I m as well as down to 285 m; they're considered rare at depths < 2-7 m (Bigelow and Schroeder 1953; McEachran 2002). On the Scotian Shelf the winter skate is most frequently caught between 37-90 m (Scott 1982a). That segment of the population residing at depths > 10 m appear to be resident year-round, even though the coastal. edge of the population appears to move shoreward in autumn and offshore in summer (Bigelow and Schroeder 1953).Merriman and Warfel (1948) stated that winter skate is a permanent resident off southern New England between 15-46 m although there are seasonal fluctuations in abundance. The 1978-2002 spring and autumn Massachusetts inshore trawl surveys (see below) show that both juveniles and adults were found between 1-75 m, with most found between 6-25 m (Figures 20 and 24).Edwards et al. (1962) captured it off the Mid-Atlantic states during the winter at depths from 33-113 m. The Page 7 1963-2002 spring and fall NEFSC trawl surveys from the Gulf of Maine to Cape Hatteras (see below) indicated that most juveniles occurred at depths < 70-80 m, although a few occurred as deep as 400 m (Figure 19), while most adults were found < 70 m and a few were as deep as 300 m (Figure 23). In the Hudson-Raritan estuary (see below;Figure 21), juvenile winter skate are found from about 4-22 m, but occur mostly around 5-8 m during a good part of the year. In Delaware Bay (see below; Figure 22)juveniles and adults were found over a range of approximately 7-21 m during winter, spring, and fall.Winter skate has been recorded over a temperature range of -1.2°C tol9°C (Bigelow and Schroeder 1953;Tyler 1971; McEachran 1973; McEachran 2002). On the Scotian Shelf it has been frequently found at temperatures of 5-9°C (Scott 1982a; Scott and Scott 1988). Bigelow and Schroeder (1953) reported them in the Gulf of Maine at around 20'C along the Massachusetts coast in the summer, down to 1-2 C in the coastal belt in winter, and near 0°C in the Bay of Fundy region in some years.McEachran and Musick (1975) reported their temperature ranges at depth of capture were -1.2°C to 4.8°C in the Gulf of St. Lawrence, 1.1-12.7'C off northeastern Nova Scotia, and 2-15°C from southern Nova Scotia to Cape Hatteras. Edwards et al. (1962) captured it off the Mid-Atlantic states during the winter at temperatures from 10-12'C. The 1963-2002 spring and fall NEFSC trawl surveys from the Gulf of Maine to Cape Hatteras (see below) collected juvenile winter skate over a temperatures range of 1-21 'C, with most found between 4-5°C in the spring and about 7-16'C in the fall (Figure 19). Adults were found over a temperature range of 2-19'C with most found around 5'C in spring and between about 1 -15C in the fall (Figure 23).Bigelow and Schroeder (1953) stated that this species is confined to sandy and gravelly bottoms but Tyler (1971) reported it from mud bottoms in Passamaquoddy Bay. In Long Island Sound during the spring, winter skate were most abundant on sand bottoms in the Mattituck Sill and Eastern Basin (Gottschall et al. 2000). On the Scotian Shelf, Scott (1982b) reports that the distribution of winter skate was confined to sand and gravel bottoms and Scott (1982b) suggests that bottom type, rather than depth, appears more important in determining the distributions of winter skate.Winter skates are known to remain buried in depressions during the day and are more active at night (Michalopoulos 1990). This is probably not due to diel foraging, since McEachran et al. (1976) observed no diel periodicity in feeding intensity by winter skate and suggested that they may feed at any time during a 24 hour period.Scott (1982a) mentions that on the Scotian Shelf during the summers of 1970-1979, winter skate was found at preferred salinities of 32-34 ppt.JUVENILES The spring and fall distributions of juvenile winter skate relative to bottom water temperature, depth, and salinity based on 1963-2002 NEFSC bottom trawl surveys from the Gulf of Maine to Cape Hatteras are shown in Figure 19. In spring, they were found in waters between 1-12'C, with the majority at about 4-5°C. Their depth range during that season was between 1-300 m, with most between about 11-70 m. They were found at salinities between 28-35 ppt, with most found between 32-33 ppt. During the fall, juvenile winter skate were caught over a temperature range of 5-21 °C, with most spread between about 7-16'C, and peaks at about 13-15'C. They were found over a depth range of 1-400 m, although most were caught at depths between about 21-80 m. They were found at salinities between 31-35 ppt, with the majority found between 32-33 ppt.The spring and autumn distributions of juveniles in Massachusetts coastal waters relative to bottom water temperature and depth based on .. 1978-2002 Massachusetts inshore trawl surveys are shown in Figure 20. In the spring they were found in waters ranging from 3-15'C, with the greatest percentages found between approximately 8-12'C. Their depth range was from approximately 6-75 m, with the majority at 6-25 m.During the autumn they were found in waters ranging from 5-21 'C; their temperature distribution was somewhat bimodal, with the major peaks between about 16-18'C. Their depth range was from 1-65 m, with the majority found between 6-25 m.The distributions and abundances of both juvenile and adult winter skate in Long Island Sound from April to November 1984-1994, based on the Connecticut Fisheries Division bottom trawl surveys, are shown in Figures 9-10. The following description of their distributions relative to depth and bottom type is taken verbatim from Gottschall et al. (2000).Winter skate were most commonly taken during the spring and late fall, occurring on average in. 16.4% of samples during these periods (Figure IOD). Abundance was highest during April, and decreased thereafter until August when none were recorded in the survey (Figure I OA). During the spring, winter skate were most abundant on sand bottom in the Mattituck Sill and Eastern Basin (Figure 9, Figure 10B). Abundance was similar in most depths, with the exception of depths between 9-18 m, where abundance was lower (Figure 10C). Winter skate abundance increased again in October and November, but they were not as concentrated on the Mattituck Sill and in the Eastern Basin as during the spring (Gottschall et al.2000).The seasonal distributions of juveniles the Hudson-Raritan estuary relative to bottom water temperature, depth, salinity, and dissolved oxygen based on 1992-1997 Hudson-Raritan trawl surveys are shown in Figure 21.The surveys show that during the winter juveniles were found mostly between 0-7°C, with > 50% at 4-5'C. Their Page 8 depth range during that season was between 4-22 m, with most caught between 5-8 m. Their salinities ranged between 20-35 ppt, most were found roughly between 23-32 ppt. They were found over a range of dissolved oxygen levels of between 9-14 ppm with a few at 5 ppm;most were found between 10-12 ppm. In spring, juvenile winter skate were found over a wider temperature range of between about 2-17'C, with bimodal peaks between approximately 5-9°C and 15-17'C and with most found between 6-9°C. The bimodality may be a function of the greater number of trawls done within those temperature intervals. Their depth range was between 4-18 m, with the majority between 5-8 m. Their salinities ranged between 15-33 ppt, most were found at 25 ppt and between 27-28 ppt. They were found over a range of dissolved oxygen levels of between 7-13 ppm with most found between 10-11 ppm. Few were caught in summer; they were found between about 16-21 *C and at depths of 7 m, 18 m, and 20 m. Their salinities ranged between 28-29ppt and at 32 ppt, and they were found over a. range of dissolved oxygen levels of between 7-8 ppm. In the fall they were found between 5-17'C, with most spread between 5-13 'C. Their depth range during the fall was between 4-21 m, with the majority between 5-8 m. Their salinities ranged between 17-34 ppt, with most spread roughly between 23-31 ppt. They were found over a range of dissolved oxygen levels of between 6-12 ppm with most found between 8-9 ppm. Based on the above evidence it appears that juvenile winter skate in the Hudson-Raritan estuary are found in warmer waters during the spring and fall as compared to winter, and remain mostly around depths of 5-8 m during those three seasons.The seasonal distributions of both juveniles and adults in Delaware Bay relative to bottom water temperature, depth, salinity, and dissolved oxygen based on 1966-1999 Delaware Division of Fish and Wildlife bottom trawl surveys are shown in Figure 22. During the winter they were found between 3-9°C, with the majority between 7-8°C. Their depth range during winter was between about 7-18 m, with peaks at 14 m, 16 m, and 17 m. Their salinities ranged between about 22-30 ppt and 34-35 ppt, most were found at 26 ppt and between 28-29 ppt. They were found over a range of dissolved oxygen levels of between 8-11 ppm, with the majority found between 9-11 ppm. In spring, they were found over a wider temperature range of between 4-17'C, with peaks scattered throughout the range (e.g., 5°C, I l°C, and 13'C). Their depth range was between 7-17 m, with a few at 21 m, and most at 8 m and 12-14 m. Their salinities ranged between 21-33 ppt, with a few at 15 ppt, and peaks scattered throughout with the two highest at 28 ppt and especially 30 ppt. They were found over a range of dissolved oxygen levels of between 7-11 ppm and 13-15 ppm, most were found between 8-11 ppm. In summer, there were too few winter skate caught to plot their distributions relative to the habitat parameters. During fall they were found between 8-13'C, with a few at 16'C;most were between 8-11 'C. Their depth range during the fall was spread between 7-11 m and at 13 m and between 18-19 m. Most were spread between 7-8 m, at 13 m, and at 18 m. Their salinities ranged between about 26-32 ppt,*with a few at 16 ppt and 22 ppt. Peaks were at 28 ppt, 30 ppt, and at 32 ppt. They were found over a range of dissolved oxygen levels of between 7-10 ppm, the majority were at 9 ppm.ADULTS The spring and fall distributions of adult winter skate relative to bottom water temperature, depth, and salinity based on 1963-2002 NEFSC bottom trawl surveys from the Gulf of Maine to Cape Hatteras are shown in Figure 23. In spring, adults were caught at temperatures between 2-11 'C, with most between 4-6°C and a peak at 5OC.During that period they were found at a depth range of 1-300 m, with the majority spread between 31-60 m. They were found at salinities of between 30-36 ppt, with the majority at 33 ppt. During the fall, they were caught over a temperature range of 5-19'C, with most caught between about I I-15°C and a peak at 14'C. They were found over a depth range of 11-300 m, with most caught at depths between about 21-70 m and peaks at 31-50 m. They were found at salinities of between 31-34 ppt, with 80-90% at 32 ppt.The spring and autumn distributions of adults in Massachusetts coastal waters relative to bottom water temperature and depth are shown in Figure 24. In the spring they were found in waters ranging from 2-16'C;the majority were spread between approximately 6-12'C.During that same season, the adults were found from 1-75 m, with most between 6-20 m. In autumn they were found between 5-19'C. The distribution was somewhat bimodal, with a peak at 10C and a minor one around 15-16'C. The depth range of the adults during autumn was aroun'd 1-75 m, with most found between 6-25 m.The distributions and abundances of both juvenile and adult winter skate in Long Island Sound relative to depth and bottom type were discussed previously (Figures 9-10; Gottschall et al. [2000]).Too few adults were found in the Hudson-Raritan estuary to plot their distributions relative to habitat parameters. The seasonal distributions of both juveniles and adults in Delaware Bay relative to bottom water temperature, depth, salinity, and dissolved oxygen based on Delaware Division of Fish and Wildlife bottom trawl surveys were discussed previously (Figure 22).STATUS OF THE STOCKS The following section is based on Northeast Fisheries Science Center (2000a, b).The principal commercial fishing method used to Page 9 catch all seven species of skates [winter, little, barndoor (Dipturus laevis), winter, thorny, clearnose (Raja eglanteria), rosette (Leucoraja garmani), smooth (Malacoraja senta)] is otter trawling. Skates are frequently taken as bycatch during groundfish trawling and scallop dredge operations and discarded recreational and foreign landings are currently insignificant, at < 1%of the total fishery landings.Skates have been reported in New England fishery landings since the late 1800s. However, commercial fishery landings, primarily from off Rhode Island, never exceeded several hundred metric tons until the advent of distant-water fleets during the 1960s. Landings are not reported by species, with over 99% of the landings reported as "unclassified skates." Skate landings reached 9,500 mt in 1969, but declined quickly during the 1970s, falling to 800 mt in 1981 (Figure 25). Landings have since increased substantially, partially in response to increased demand for lobster bait, and more significantly, to the increased export market for skate wings. Wings are taken from winter and thorny skates, the two species currently used for human consumption. Bait landings are presumed to be primarily from. little skate, based on areas fished and known species distribution patterns. Landings for all skates increased to 12,900 mt in 1993 and then declined somewhat to 7,200 mt in 1995. Landings have increased again since 1995, and the 1998 reported commercial landings of 17,000 mt were the highest on record (Figure 25).The biomass for the seven skate species is at a medium level of abundance. For the aggregate complex, the NEFSC spring survey index of biomass was relatively constant from 1968-1980, then increased significantly to peak levels in the mid- to late 1980s. The index of skate complex biomass then declined steadily until 1994, but has recently increased again. The large increase in skate biomass in the mid- to late 1980s was dominated by winter and little skate. The biomass of large sized skates (> 100 cm max. length: barndoor, winter, and thorny) has steadily declined since the mid-1980s. The recent increase in aggregate skate biomass has been due to an increase in small sized skates (< 100 cm max. length: little, clearnose, rosette, and smooth) -primarily little skate.Winter skate abundance is currently about the same as in the early 1970s, and is about 25% of the peak observed in the mid-1980s (Figure 25). Winter skate was, until recently, considered to be overfished (Northeast Fisheries Science Center 2000a, b), but its-status has recently been changed so that it is no longer considered to be in an overfished condition (NMFS 2002).RESEARCH NEEDS Imprecise reporting of fishery statistics where several skate species are lumped together under one category (e.g., "unclassified skates" or "skates spp.") can mask basic changes in community structure and profound reduction in populations of larger, slower growing species (Dulvy et al. 2000; Musick et al., 2000). Thus, it is important to have fishery-independent data on skates where the individual species are reported; it is also necessary to work out any identification problems between winter and little skate.Northeast Fisheries Science Center (2000b) also suggests the following research needs: " More life history studies (including age, growth, maturity, and fecundity studies) are necessary." Studies of stock structure are needed to identify unit stocks.* Explore possible stock-recruit relationships by examination of NEFSC survey data.* Investigate trophic interactions between skate species in the complex, and between skates and other groundfish." Investigate the influence of annual changes in water temperature or other environmental factors on shifts in the range and distribution of the species in the skate complex, and establish the bathymetric distribution of the species in the complex in the northwest Atlantic." Investigate historical NEFSC survey data from the R/V Albatross III during 1948-1962 when they become available, as they may provide valuable historical context for long-term trends in skate biomass.ACKNOWLEDGMENTS The authors thank Barry Shafer, John McCarthy, Tom Finneran, Annette Kalbach, and Meredith Lock for producing the maps and graphics. Thanks also to Claire Steimle and Judy Berrien for literature reviews and interlibrary loans. Frank Almeida and Kathy Sosebee of the NEFSC Woods Hole provided much needed information, input, and reviews.REFERENCES CITED Avent, S.R., S.M. Bollens, M. Butler, E. Horgan, and R.Rountree. 2001. Planktonic hydroids on Georges Bank: ingestion and selection by predatory fishes.Deep-Sea Res II 48: 673-684.Bigelow, H.B. and W.C. Schroeder. 1953. Fishes of the Gulf of Maine. U.S. Fish Wildl. Serv., Fish. Bull. 53.577 p.Bor, P.F.H. 2001 Jan. 22. Table. Egg-capsules of rays and skates. <http://www.raiidae.tmfweb.nl/>. Accessed 2001 Feb. 7.Bowman, R.E., C.E. Stillwell, W.L. Michaels, and M.D.Grosslein. 2000. Food of northwest Atlantic fishes and two species of squid. NOAA Tech. Mem. NMFS-NE-155. 138 p. Page 10 Collette, B.B. and K.E. Hartel. 1988. Annotated list of fishes of Massachusetts Bay. NOAA Tech. Mem.NMFS-F/NEC-51. 70 p.Edwards, R.L., R. Livingstone, Jr., and P.E. Hamer. 1962.Winter water temperatures and an annotated list of fishes -Nantucket Shoals to Cape Hatteras. Albatross III Cruise no. 126. U.S. Fish Wildl. Serv. Spec. Sci.Rep. Fish. 397. 31 p.Frisk, M.G., T.J. Miller, and M.J. Fogarty. 2001.Estimation and analysis of biological parameters in elasmobranch fishes: a comparative life history study.Can. J. Fish. Aquat. Sci. 58: 969-981.Fitz, E.S., Jr. and F.C. Daiber. 1963. An introduction to the biology of Raja eglanteria Bosc 1802 and Raja erinacea Mitchill 1825 as they occur in Delaware Bay.Bull. Bingham Oceanogr. Collect., Yale Univ. 18 (3): 69-97.Garrison, L.P. 2000. Spatial and dietary overlap in the Georges Bank groundfish community. Can. J. Fish.Aquat. Sci. 57: 1679-1691. Garrison, L.P. and J.S. Link. 2000a. Dietary guild structure of the fish community in the Northeast United States continental shelf ecosystem. Mar. Ecol.Prog. Ser. 202: 231-240.Garrison, L.P. and J.S. Link. 2000b. Fishing effects on spatial distribution and trophic guild structure of the fish community in the Georges Bank region. ICES J.Mar. Sci. 57: 723-730.Geer, P.J. 2002. Summary of essential fish habitat description and identification for Federally managed species inhabiting Virginia waters of Chesapeake Bay 1988-1999. Virginia Mar. Res. Rep. VMRR 2001-03, Jan. 2001, Revised.June 2002. 169 p.Gottschall, K, M.W. Johnson, and D.G. Simpson. 2000.The distribution and size composition of finfish, American lobster, and long-finned squid in Long Island Sound based on the Connecticut Fisheries Division Bottom Trawl Survey, 1984-1994. NOAA Tech. Rep. NMFS 148. 195 p.Hildebrand, S.F. and W.C. Schroeder. 1928. Fishes of Chesapeake Bay. Bull. U.S. Bur. Fish. 43(1). 366 p.Huntsman, A.G. 1922. The fishes of the Bay of Fundy.Contrib. Can. Biol. 1922 (3): 49-72 Kaplan, J. 1999. Black-backed gull catches a skate. The Connecticut Warbler 19: 81-82.Macdonald, J.S., M.J. Dadswell, R.G. Appy, G.D.Melvin, and D.A. Methven. 1984. Fishes, fish assemblages, and their seasonal movements in the lower Bay of Fundy and Passamaquoddy Bay, Canada. Fish. Bull. (U.S.) 82: 121-139.Mahon, R. 1997. Demersal fish assemblages from the Scotian Shelf and Bay of Fundy, based on trawl survey data (1979-1993). Can. Manuscr. Rep. Fish.Aquat. Sci. 2426. 38 p.McEachran, J.D. 1973. Biology of seven species of skates (Pisces: Rajidae). Ph.D. dissertation, Coll. William and Mary, Williamsburg, VA. 127 p.McEachran, J.D. 2002. Skates. Family Rajidae. In B.B.Collette and G. Klein-MacPhee eds. Bigelow and Schroeder's fishes of the Gulf of Maine. 3 rd Edition.p. 60-75. Smithsonian Institution Press, Washington, DC. 748 p. 'McEachran, J.D., D.F. Boesch, and J.A. Musick. 1976.Food division within two sympatric species-pairs of skates (Pisces: Rajidae). Mar. Biol. 35: 301-317.McEachran, J.D. and K.A. *Dunn. 1998. Phylogenetic analysis of skates, a morphologically conservative clade of elasmobranchs (Chondrichthyes: Rajidae).Copeia 1998 (2): 271-290.McEachran, J.D. and C.O. Martin. 1977. Possible occurrence of character displacement in the sympatric skates Raja erinacea and R. ocellata (Pisces: Rajidae).Environ. Biol. Fishes 2: 121-130.McEachran, J.D. and J.A. Musick. 1973. Characters for distinguishing between immature specimens of the sibling species, Raja erinacea and Raja ocellata (Pisces: Rajidae). Copeia 1973: 238-250.McEachran, J.D. and J.A. Musick. 1975. Distribution'and relative abundance of seven species of skates (Pisces: Rajidae) which occur between Nova Scotia and Cape Hatteras. Fish. Bull. (U.S.) 73: 110-136.Merriman, D. and H.E. Warfel. 1948. Studies on the marine resources of southern New England. VII.Analysis of a fish population. Bull. Bingham Oceanogr. Collect., Yale Univ. 11 (4): 131-164.Michalopoulos, C. 1990. A field study on the ecology and behavior of the winter skate (Raja ocellata) and little skate (Raja erinacea) off outer Cape Cod, Massachusetts. M.S. thesis, Southeastern Mass. Univ., North Dartmouth, MA. 71 p.Michels, S.F. and M.J. Greco. 2000. Coastal finfish assessment survey -annual report. Delaware Div.Fish Wildl., Proj. F-42-R-I 1, Dover, DE. 72 p.Murdy, E.O., R.S. Birdsong, and J.A. Musick. 1997.Fishes of Chesapeake Bay. Smithsonian Institution Press, Washington, DC. 324 p.Nelson, J.I., Jr., S. Perry, D. Miller, and G. Lamb. 1983.Inventory of New Hampshire's marine coastal fisheries. N.H. Fish Game Dep., Div. Inland Mar.Fish. 35 p.NMFS. 2002. Annual report to Congress on the status of U.S. fisheries -2001. U.S. Dep. Commerce, NOAA, NMFS, Silver Spring, MD. 142 p.Northeast Fisheries Science Center. 2000a. Report of the 30th Northeast Regional Stock Assessment Workshop (30th SAW): Public Review Workshop. Northeast Fish. Sci. Cent. Ref. Doc. 00-04. 53 p.Northeast Fisheries Science Center. 2000b. Report of the 30th Northeast Regional Stock Assessment Workshop (30th SAW): Stock Assessment Review Committee (SARC) consensus summary of assessments. Northeast Fish. Sci. Cent. Ref. Doc. 00-03. 477 p.Overholtz, W.J. and A.V. Tyler. 1985. Long-term responses of the demersal fish assemblages of Georges Bank. Fish. Bull. (U.S.) 83: 507-520. Page 11 Reid, R., F. Almeida, and C. Zetlin. 1999. Essential fish habitat source document: Fishery independent surveys, data sources, and methods. NOAA Tech.Mem. NMFS-NE-122. 40 p.Schaefer, R.H. 1967. Species composition, size and seasonal abundance of fish in the surf waters of Long Island. N.Y. Fish Game J. 14: 1-46.Scott, J.S. 1982a. Depth, temperature and salinity preferences of common fishes of the Scotian Shelf. J.Northwest Atl. Fish. Sci. 3: 29-39.Scott, J..S. 1982b. Selection of bottom type by groundfishes of the Scotian Shelf. Can. J. Fish. Aquat.Sci. 39: 943-947.Scott, J.S. 1989. Matrix analysis of co-occurrences of fishes of the Scotian Shelf and Bay of Fundy. Can. J.Fish. Aquat. Sci. 46: 191-197.Scott, W.B. and M.G. Scott. 1988. Atlantic fishes of Canada. Can. Bull. Fish. Aquat. Sci. 219. 731 p.Simon, J.E. and K.T. Frank 1996, Assessment of the Division 4VsW skate fishery. DFO Atd. Fish. Res..Doc. 96/105.51 p.Simon, J.E. and K.T. Frank 1998. Assessment of the winter skate fishery in Division 4VsW. Can. Stock Assessment Secretariat Res. Doc. 98/145. 41 p.Smith, F.E. 1950. The benthos of Block Island Sound.The invertebrates, and their quantities and their relations to the fishes. Ph.D. dissertation, Yale Univ., New Haven, CT. 213 p.Strong, M. and A. Hanke. 1995. Diversity of finfish species in the Scotia-Fundy region. Can. Tech. Rep.Fish. Aquat. Sci. 2017. 106 p.Tsou, T.S. and J.S. Collie. 2001a. Estimating predation mortality in the Georges Bank fish community. Can. J.Fish. Aquat. Sci. 58: 908-922.Tsou, T.S. and J.S. Collie. 2001b., Predation-mediated recruitment in the Georges Bank fish community. ICES J. Mar. Sci. 58: 994-1001.Tyler, A.V. 1971. Periodic and resident components in communities of Atlantic fishes. J. Fish. Res. Board Can. 28: 935-946.Tyler, A.V. 1972. Food resource division among northern, marine demersal fishes. J. Fish. Res. Board Can. 29: 997-1003.Vladykov, V.D. 1936. Capsules d'oeufs de raies de I'Atlantique Canadien appartenant au genre Raja.Natur. Can. 63: 211-231. Page 12 Table 1. Summary of habitat parameters for winter skate, based on the pertinent literature. Life Stage Depth Substrate Temperature Juveniles Generally caught at depths from Sand and gravel bottoms but Recorded over a temperature shoreline to 371 m, although most reported from mud bottoms in range of-1.2°C to 19'C.abundant < I l I m. In Gulf of Passamaquoddy Bay. In Long Reported in Gulf of Maine at Maine, most abundant 46-64 m Island Sound during spring 1984- 20'C along Massachusetts coast but occasionally occur < I m and 1994, most abundant on sand in summer, down to 1-2°C in the down to 285 m; they're bottoms in the Matiituck Sill and coastal belt in winter, near 0°C in considered rare at depths < 2-7 m. Eastern Basin. On Scotian Shelf, the Bay of Fundy region in some May be permanent resident off confined to sand and gravel years. Reported at a temperature southern New England between bottoms; bottom type, rather than range of 2-15'C from southern 15-46 m, although there are depth, may be more important in Nova Scotia to Cape Hatteras.seasonal fluctuations in determining distributions of Captured off Mid-Atlantic states abundance. Has been captured off winter skate. during winter at 10-12'C.the Mid-Atlantic states during the winter at depths from 33-113 m. Remains buried in depressions In Long Island Sound during during the day and are more spring 1984-1994, abundance of active at night. This is probably winter skate was similar in most not due to diel foraging since no depths, with the exception of diel periodicity in feeding depths between 9-18 m, where intensity has been observed; they abundance was lower. may feed at any time during a 24 hour period Same as for juveniles. Same as for juveniles. Same as for juveniles. Adults 2 Merriman and Warfel (1948); Bigelow and Schroeder (1953); Edwards et al. (1962); Tyler 1971; McEachran (1973); McEachran and Musick (1975); McEachran et al. (1976); Scott (1982b); Michalopoulos (1990); Gottschall el al. (2000); McEachran (2002).2 Merriman and Warfel (1948); Bigelow and Schroeder (1953); Edwards et at. (1962); Tyler 1971; McEachran (1973); McEachran and Musick (1975); McEachran el al. (1976); Scott (1982b); Michalopoulos (1990); Gottschall et al. (2000); McEachran (2002). Page 13 Table 1. cont'd.Life StageI Prey Predators/Species Associations Juveniles Polychaetes and amphipods most important prey in terms of numbers or occurrence, followed by decapods, isopods, bivalves, fishes. Hydroids also ingested. In terms of weight, amphipods, decapods and fish can be most important; fish are especially prevalent in larger skate. Polychaetes include: Nephtys spp., Nereis spp., Lumbrinerisfragilis, Ophelia denticulata, maldanids (mostly Clymenella torquata), Aphrodite hastata. Amphipods: haustoriids, Leptocheirus pinguis, Monoculodes sp., Hippomedon serratus, ampeliscids, Paraphoxus sp., Tmetonyx sp., Unciola irrorala, Bvblis serrata, oedicerotids. Decapods: Crangon septemspinosa, Cancer irroratus, pagurid crabs, Dichelopandalus leptocerus, pandalid shrimp. Isopods: Cirolana (= Politolana?) polita. Bivalves include Solem*ya sp. and Ensis directus.Sand lance was the most frequently eaten fish; yellowtail flounder, longhorn sculpin, hakes, other skate, herring, butterfish occasionally eaten. Generally, in terms of numbers or occurrence, crustaceans made up> 50% of the diet for skate < 61 cm TL, while fish (and often bivalves) were a major part of the diet of skate >79-91 cm TL.Overall crustaceans declined in importance with increasing skate size (includes both amphipods and often decapods) while polychaetes increased with increasing skate size until the skate were about 81 cm TL.Amphipods occurred more frequently than decapods until the skates were > 71 cm TL.On Georges Bank, decline in importance of fish prey in spring may be related to seasonal movements of prey. In terms of weight, crustaceans were dominant in the diet of skate < 31-50 cm TL, while fish, mostly sand lance, were dominant in the diet of skate 51-110 cm TL. For skate < 31 cm TL, amphipods dominated, especially L.pinguis. For skate 31-50 cm TL, decapods dominated, especially C. septemspinosa and C. irroratus. Predators: sharks, other skates, gray seals, and gulls.Winter and little skate co-occur from Nova Scotia to Cape Hatteras.Although winter and little skate are sympatric species with similar habitat requirements, there's not a high degree of competitive interaction between them because they are positively correlatedby abundance. Also, winter skate feeds largely on infauna, while little skate predominately selects epifauna. Sympatric populations of winter and little skate also undergo character displacement in order to avoid direct competition for food resources. Using 1973-1997 NEFSC data from Nova Scotia to Cape Hatteras and NEFSC food habits database, both small (10-30 cm TL) and medium (31-60 cm TL) sized winter skate belonged to "Amphipod/shrimp eaters" group, along with little skate and cusk eel; prey included amphipods, polychaetes, shrimp, zooplankton. Largest winter skate (61 to > 80 cm TL) by themselves in a subgroup of"Piscivores" because their diet contained a high proportion of sand lance. Again, a trend toward increasing piscivory with size.On Georges Bank, winter skate belongs to spatial assemblages and trophic groups that include Atlantic cod, little skate, longhorn sculpin, yellowtail flounder, red and silver hake, haddock, spiny dogfish, butterfish, fourspot flounder, windowpane, winter flounder, sea raven, thorny skate, Atlantic herring. Also on Georges Bank, winter skate falls into various dietary guilds or trophic groups, depending on the study. Garrison and Link (2000b): "Bentho-pelagic" group included 31-60 cm TL winter skate, little skate, longhorn sculpin, Atlantic cod.Diets of these species included shrimp such as pandalids and C.septemspinosa, and benthic invertebrates including polychaetes, gammarid amphipods, bivalves. "Skate" group included > 80 cm TL winter skate, Oollock, windowpane. Prey was a combination of fish and benthic prey, with a high proportion of sand lance. Garrison (2000): In autumn, 61 cm to > 80 cm TL winter skate by themselves in a subgroup of the "Piscivorous" group, feeding on sand-lance, silver hake, and Atlantic herring, plus benthic invertebrates. 10-60 cm TL winter skate in "Demersal predators" group with flatfish, haddocks, little skate, thorny skate. Prey included gammarid amphipods, polychaetes, isopods, Cancer crabs, C. septemspinosa. During spring, 10-60 cm TL winter skate in "Shrimp/amphipod predators" group with hakes, longhorn sculpin, Atlantic cod, fourspot flounder, little skate, thorny skate. Prey included gammarid amphipods, pandalids, C. septemspinosa, polychaetes, Cancer crabs.Winter skate 61 cm to > 80 cm TL by themselves in a "Generalist" subgroup, consuming bivalves, polychaetes, sand lance, herring.Decline in importance of fish prey, 35% fish in autumn and 16% in spring, probably related to seasonal movements of prey.Same as for juveniles; however, note that Same as for juveniles, but note differences between smaller and larger Adults 2 larger skates consume more polychaetes and skates.'fish while crustaceans decline inthe diet.I Bigelow and Schroeder (1953); McEachran (1973); McEachran and Musick (1975); McEachran et al. (1976); McEachran and Martin (1977); Overholtz and Tyler (1985); Scott and Scott (1988); Kaplan 1999; Bowman et al. (2000); Garrison (2000); Garrison and Link (2000a, b); Avent et al. (2001); (Tsou and Collie 2001a, b); NEFSC 1973-1990 food habits database.2 Bigelow and Schroeder (1953); McEachran (1973); McEachran and Musick (1975); McEachran et al. (1976); McEachran and Martin (1977); Overholtz and Tyler (1985); Scott and Scott (1988); Kaplan 1999; Bowman et al. (2000); Garrison (2000); Garrison and Link (2000a, b); Avent et al. (2001); (Tsou and Collie 2001a, b); NEFSC 1973-1990 food habits database. Page 14 Table 2. Summary of habitat parameters for winter skate, based on the most recent NEFSC and state surveys mentioned in the text.Life Stage [ Survey Depth Temperature Salinity/DO Juveniles 1963-2002 spring and fall NEFSC trawl surveys from Gulf of Maine to Cape Hatteras.Spring: range of 1-300 m, most between about 11-70 m.Fall: range of 1-400 m, most between about 21-80 m.Spring: range of 1-12'C, majority at about 4-5°C.Fall: range of 5-21 'C, most spread between about 7-16'C, peaks at about 13-15'C.Spring: range of 28-35 ppt, most between 32-33 ppt.Fall: range of 31-35 ppt, majority between 32-33 ppt.1978-2002 Spring: range of Spring: range of 3-15°C, Massachusetts approximately 6-75 m, greatest percentages between inshore trawl majority at 6-25 m. approximately 8-12'C.surveys. Fall: range of 1-65 m, Fall: range of 5-21 'C, majority between 6-25 m. temperature distribution somewhat bimodal, major peak between about 16-18'C.1992-1997 NEFSC Winter: range of 4-22 m, most Winter: range of 0-7°C, > Winter: range of 20-35 ppt, trawl surveys of the between 5-8 m. 50% between 4-5°C. most between roughly 23-32 Hudson-Raritan Spring: range of 4-18 m, Spring: range of 2-17'C, with ppt / range of 9-14 ppm with a estuary. majority between 5-8 m. bimodal peaks between 5-9°C few at 5 ppm, most between Summer: few caught, found at and 15-17'C, most between 6- 10-12.ppm. 7 m, 18 m, and 20 m. 9°C. Spring: range of 15-33 ppt, Fall: range of 4-21 m, Summer: few caught, found most at 25 ppt and between majority between 5-8 m. between about 16-21 *C. 27-28 ppt / range of 7-13 ppm, Fall: range of 5-17'C, most most between 10- 11 ppm.spread between 5-13'C. Summer: few caught, between 28-29 ppt and at 32 ppt /between 7-8 ppm.Fall: range of 17-34 ppt, most spread roughly between 23-31 ppt / range of 6-12 ppm, most_between 8-9 ppm.1966-1999 Delaware Division of Fish and Wildlife bottom trawl surveys of Delaware Bay (juveniles and adults combined)Winter: range of about 7-18 m, peaks at 14 m, 16 m, and 17 m.Spring: range of 7-17 m, a few at 21 m, most at 8 m and 12-14 m.Fall: range of 7-11 m and at 13 m and between 18-19 m;most spread between 7-8 m, at 13 m, and at 18 m.Winter: range of 3-9°C, majority between 7-8°C.Spring: range of 4-17'C, peaks scattered throughout (e.g., 5°C, II°C, and 13'C).Fall: range of 8-13°C, a few at 16'C; most between 8-11 *C.Winter: range of about 22-30 ppt and 34-35 ppt, most at 26 ppt and between 28-29 ppt /range of 8-11 ppm, majority between 9-11 ppm.Spring: range of 21-33 ppt, a few at 15 ppt; peaks scattered throughout with the two highest at 28 ppt and especially 30 ppt / range of 7-11 ppm and 13-15 ppm, most between 8-11 ppm.Fall: range of about 26-32 ppt, a few at 16 ppt and 22 ppt;peaks at 28 ppt, 30 ppt, and 32 ppt / range of 7-10 ppm, majority at 9 ppm. Page 15 Table 2. cont'd.Life Stage Survey Depth T Temperature Salinity/DO Adults 1963-2002 spring and fall NEFSC trawl surveys from Gulf of Maine to Cape Hatteras.Spring: range of 1-300 m, majority spread between 31-60 M.Fall: range of 11-300 m, most between about 21-70 m, peaks at 31-50 m.Spring: range of 2-11 'C, most between 4-6°C, peak at 5°C.Fall: range of 5-19'C, most between I I-15°C, peak at 14'C.Spring: range of 30-36 ppt, majority at 33 ppt.Fall: range of 31-34 ppt, with 80-90% at 32 ppt.1978-2002 Spring: range of 1-75 m, most Spring: range of 2-16'C, Massachusetts between 6-20 m. majority spread between inshore trawl Fall: range of around 1-75 m, approximately 6-12'C.surveys. most between 6-25 m. Fall: range of 5-19'C, distribution somewhat bimodal: peak at 10°C and a minor one around 15-16°C.1966-1999 Delaware Division of Fish and Wildlife bottom trawl surveys of Delaware Bay (juveniles and adults combined)See juveniles. See juveniles. See juveniles. Page 16 Figure 1. The winter skate, Leucoraja ocellata (Mitchill 1815), male, from Murdy et al. (1997). Page 17 Figure 2. Egg case of winter skate, from Bor (2001). Page 18 1973-1980 Amphipoda 37.5%1981-1990 Polychaeta Polychaeta 13.1%f Unidentified ft..Crustacea 1.6%Isopoda 9.4%Other prey ier crustacea 1.6% phyla 3.3%Other prey phyla 17.2%Polychaeta 9.40?Other r phyla 9 Polychaeta V Decapoda 23.4%n= 10 poda n=21-30 cm Amphipoda 42.6%Other crustacea 16.4%a 4.6%50 Amphipoda 38.8%Isopoda 5.4%Mollusca 4.9%'Unidentified , Crustacea 2.4%,da 17.1% Fish 4.4%=462 Amphipoda 33.1%Other Crustacea 1.1%Decapoda 22.9%n=48 31-40 cm n Amphipoda 33.5%Unidentified Crustacea 4.2%Fish 3.4%Isopoda 6.8%Other Crustacea 2.0%Decapoda 22.3%n = 53\ Unidentified Fish 7.9% Crustacea 1.8%n = 516 41-50 cm Figure 3. Abundance (% occurrence) of the major prey items of winter skate collected during NEFSC bottom trawl surveys from 1973-1980 and 1981-1990. Methods for sampling, processing, and analysis of samples differed between the time periods [see Reid et al. (1999) for details]. Page 19 1973-1980 1981-1990 Amphipoda 31.6%Amphipoda 26.0%Polychaeta ýUnidentified Crustacea 1.6%I IV-/o 51-60 cm n = 31 n =399 rAmphipoda 20.3% Polychaeta 28.4% _Crustacea Shirmp 1.0c Decapoda /10.5%nphipoda 13.2%Othe ./:°Isopoda Crustacea 1.6%Mollusca 9.3%12.0%Fish 23.9%61-70 cm n = 78 n = 394 Polychaeta 25.2%Polychaeta 34.Crustacea Shirmp 1.9 Amphipoda 11.8%6.7%Other prey phyla 4.4%Decapoda 12.7%Other 0% a Crustacea 2.0% Isopoda 7.9%Unidentified Crustacea 1.1 %Mollusca 9.4%Fish 16.0%71-80 cm n = 149 n = 450 Figure 3. cont'd. Page 20 1973-1980 1981-1990 Crustacea 1% 'Shirmp 2.2%Polychaeta 27.9%Other prey phyla 6.3%Decapoda 10.5%Other Crustacea 1.8%Isopoda 7.2%kmphipoda 7.8%Unidentified Crustacea 2.4%Mollusca 7.2%81-90 cm n = 170 Otherprey phyla 10.3%S Polychaeta 10.3%n = 710 Other Unidentified Crustacea 2.1%Fish 56.7%Fish 91-100 cm n = 91 n = 488 Figure 3. cont'd. Page 21 Figure 4. Distribution of juvenile winter skate collected during winter NEFSC bottom trawl surveys [1964-2002, all years combined; see Reid et al. (1999) for details]. Survey stations where juveniles were not found are not shown. Page 22 Spring/juveniles (<= 84 cm)Number per tow 1 -10 0 11-100 0 101-1000 0 > 1000 Winter Skate NEFSC Bottom Trawl Surveys (1968 -2002)Figure 5. Distribution and abundance of juvenile winter skate collected during spring NEFSC bottom trawl surveys[1968-2002, all years combined; see Reid et al. (1999) for details]. Page 23 Figure 6. Distribution of juvenile winter skate collected during summer NEFSC bottom trawl surveys [1963-1995, all years combined; see Reid et al. (1999) for details]. Survey stations where juveniles were not found are not shown. Page 24 FaIlljuveniles (<= 84 cm)Number per tow 1 -10 0 11 -100 0 101 -1000 0 > 1000 Winter Skate NEFSC Bottom Trawl Surveys (1963 -2001)Figure 7. Distribution and abundance of juvenile winter skate collected during fall NEFSC bottom trawl surveys [1963-2001, all years combined; see Reid et al. (1999) for details]. Page 25 Figure 8. Distribution and abundance of juvenile winter skate in Massachusetts coastal waters collected during the spring and autumn Massachusetts inshore trawl surveys [1978-2002, all years combined; see Reid et al. (1999) for details]. Page 26 Winter Skate Massachusetts Inshore Trawl Survey (1978- 2001);~ ~;?.J /I Fall/juveniles (<= 84 cm)Number per tow* 1-10* 11-100* 101 -500 D > 500 Massachusetts 0 0 V Figure 8. cont'd. Page 27 TUAfS=1114 Sum = 436 Max > 12 Mill -1 X=O Figure 9. Distribution and abundance ofjuvenile and adult winter skate collected in Long Island Sound, based on the finfish surveys of the Connecticut Fisheries Division, 1984-1994 [from Gottschall et al. (2000)]. Circle diameter is proportional to the number of fish caught, and is scaled to the maximum catch (indicated by "max=" or "max>").-Collections were made with a 14 m otter trawl at about 40 stations chosen by stratified random design. Page 28 04~0.3)0 ICV a 0.2~0i bMonth Do Percent Occurrencel by Month 30 20[!? .)) 0.00 CO 3, tip.. ....... .......... 10o 005 02 74... .. .... .... ..... .... 0 .._ _.... .- .....SE. F- P-eierent Occurrence I by Month and Bottom Type SO~ --...... ....................................... .Abundance iby Month and Bottom Type 1 2 ......-. .......i a C o.40 D40 ja , 0.0e o 0 o 1 .0. 3 0 o o.o0 o..0 is mud 10. 01 77 1 3,5 1,, .Ao 0,0 o0.0 6.2 7.9.c. .. -w, 2 , ý.o : 0.0 0 2o,6 142 1 as 04 i so________ G __ 'D0oW G4ooo .1 1 1.8 32,6 7.7 00 LI ý 82 2 III' .....undance Ii Percent Occurrence 1by Month and Depth Interval iby Month and Depth Intervalt II.0.4 4, -- 4 2D o.0 0.10 4'-~ --- -0 0----- I-6.6__ 011 1 4 1 2 I 1 001 0.0 0.00 002 .0210 o 0.7 ...... ..O. .... .1. .... ...... .. .... ...... ... .... .................................. ...................... .. ...... .......Figure 10. Relative abundance (geometric mean catch/tow) catch/tow and percent occurrence (proportion of samples in which at least one individual was observed) for juvenile and adult winter skate in Long Island Sound by month, month and bottom type, and month and depth interval. From Gottschall et al. (2000). Page 29 74" 15" 740 10.WINTER SKATE--Juveniles (<= 84 cm)Winter (92 -97)Hudson/Raritan Trawl Survey .Fish Caught/station (excluding 0 catch): Mean: 3 Min: 1 Max: 18 74' 05"............... 4- 7 ........ -r 740 00"..... ...... ... ... ....... .. ... ? ...... .....-\-Hudson River'<N New York Length (cm): Min: 25 Max: 84 Staten Island y .~0. '0 C,' 0 C,,....---.*/ 0 C .. 0'/C> .0.1 .S' ~. b 4 0.o.* 0* * *~Og.'0.S.73" 55........... I IIII_35".4Q0 30")0 Number/Tow 1-2* 3-5-0 6-10 f400 25.>10 New Jersey 74 15" 740 10'WINTER SKATE--Juveniles (<= 84 cm)Spring (92 -97)Hudson/Raritan Trawl Survey Fish Caught/station (excluding 0 catch): Mean: 2 Min: 1 Max: 13 Length (cm): Min: 21 Max: 84 Staten /, Island 74005"/..A-74° 000" 73' 55" Hudson River"" New York \\-- 40 35-/N'C ., C,, *0~*0 40° 30'csi New Jersey 0 P",'Number/Tow 1-2* 3-5* 6-10 40*25">10 Figure 11. Seasonal distribution and abundance ofjuvenile winter skate in the Hudson-Raritan estuary,based on Hudson-Raritan trawl surveys, 1992-1997 [see Reid et al. (1999) for details]. Page 30 74O 15 ' 74' 10" WINTER SKATE--Juveniles (<= 84 cm)Summer (92 -96)Hudson/Raritan Trawl Survey Fish Caught/station (excluding 0 catch): Mean: 2 Min: 1 Max: 3 Length (cm): Min: 39 Max: 61 74° 05" 740 00'730 55--Hudson River New York--..... -40*35-V 7, Staten Island 40° 30 'fý/.1 9--.d'1-011 New Jersey 74° 15'" 740 10 740 05' 740 00 A -H u d sn.......... I........ ............... .......... R e .......... r", --L,-Hudson River Number/Tow 1-2* 3-5 -* 6-10 -i40 25">10 730 55" v York i- 0":., 40°35 WINTER SKATE--Juveniles (<= 84 cm)Fall (92 -96)Hudson/Raritan Trawl Survey Fish Caught/station (excluding 0 catch): Mean: 3 Min: 1 Max: 14 Length (cm): Min: 25 Max: 83 Staten Island N Ne~e/* .. 0*

• 4 0 *0* .... .0.* 0,* *** .0 0~.* S.,.a 0 S.. *0.*.5* 6 0ý40° 30'I S.o 6 P ,n ~ .N'V Number/Tow
• 1-2 3-5 -* 6-10 .40°25>10 New Jersey Figure 11. cont'd.

Page 31.750 30 '750o10 750 00' I 740 50.WINTER SKATE All Size Classes Combined Winter (1966 -1999)Delaware Bay Trawl Survey Average Number of Fish Caught and Percent Occurrence

390 30".390 20'4w.NJ A 0.00 0%0.07/'7%C-s'0.33 20%0.23.14%0.00 0%' 390 10/</.3* / r ..* h:39° O0 6 ape May# Times Station Sampled 0.60 13%0.00 0%0.69 25%1.20 33%3.29.29%0.33 33%0.00 00/0 DE"*, 0.40 lO0%/1-5 10-22.31 -40 o 41-50 o 51-61~380 50 Data Source: Delaware Division of Fish & Wildilfe Cape/Henlopenr.

Figure 12. Seasonal distribution and abundance of juvenile and adult winter skate in Delaware Bay, based on Delaware Division of Fish and Wildlife bottom trawl surveys from 1966-1999 (all years combined). Surveys were conducted monthly at 9-14 fixed stations, using a 9.1 m otter trawl towed for 20-30 min (for methods see Michels and Greco 2000). Page 32 750 30 '750 20 " 750 10 '750 00 " 740 50 '~-- -y WINTER SKATE All Size Classes Combined Spring (1966 -1999)Delaware Bay Trawl Survey Average Number of Fish Caught and Percent Occurrence NJ/" K"

• 390 30" 390 20 1390 10'0.00 0 0%'N-0.00 0 0%0.15 0 7%0.30 D E DE 0.31 0 17%0.11 0 7% 0.85 35%0.25 0.41 14% 22%0.00 00/%0.60 60%4.50 100%/ .i/ I Ca# Times Station Sampled-- 1-5 0.4550" 18%open;10-22* 31-40 o 41-50* 51-61 Data Source: Delaware Division of Fish & Wildlife Cap(Henlh Figure 12. cont'd.

Page 33 7, 50 30" 75o 20" 75- 10 75 00" 740 50 WINTER SKATE All Size Classes Combined 30 Summer (1966 -1999)-A,, Delaware Bay Trawl Survey Average Number of Fish Caught and Percent Occurrence 0.00 ,ýp _ 0,,.0 ,d! --39° 20" 0% ,f" 0%0.00 0.00 0% 0%/ 39° 00 0% +~o/ 0 0.0 0.00 0% 0 0% #Times Station Sampled 1-5 DE \ 0.00"\* 0.00 10-22 -3850".31-40 Data Source: / o 41-50 HenlopenH Delaware Division of Fish & Wildilfe e 51-61.. .' ......... ......... .........Figure 12. cont'd. Page 34 750 30 " 750 20 '750 10 '750 00'WINTER SKATE All Size Classes Combined Fall (1966 -1999)Delaware Bay Trawl Survey 740 50;390 30~.390 20 Average Number of Fish Caught and Percent Occurrence ('J390 10 x J 0.05 0 2%0.jo 0%0.00 0%0.06 0 ,.{ 4%DE 39° 00 0.00 0%0.34 14%0.13 3%0.26 0 7%/0.20 20%0.00 0%0.06 Data Source: Cape/Delaware Division of Fish & Wildilfe Henlopen'...... .!. .. .......... ...... ... .......... .... ........ .. .... ........... ..... ..................... ............ .# Times Station Sampled-- 1-5 10-22* 31-40 o 41-50 e 51-61 0=138° 50'-L.. ...]Figure 12, cont'd. Page 35 Figure 13. Distribution of adult winter skate collected during winter NEFSC bottom trawl surveys [1964-2002, all years combined; see Reid et al. (1999) for details]. Survey stations where adults were not found are not shown. Page 36 Figure 14. Distribution and abundance of adult winter skate collected during spring NEFSC bottom trawl surveys [1968-2002, all years combined; see Reid et a. (1999) for details]. Page 37 cm).Present Winter Skate NEFSC Bottom Trawl Surveys (1963-1995) Figure 15. Distribution of adult winter skate collected during summer NEFSC bottom trawl surveys [1963-1995, all years combined; see Reid et al. (1999) for details]. Survey stations where adults were not found are not shown. Page 38 Figure 16. Distribution and abundance' of adult winter skate collected during fall NEFSC bottom trawl surveys [ 1963-2001, all years combined; see Reid et al. (1999) for details]. Page 39 Figure 17. Distribution and abundance of adult winter skate in Massachusetts coastal waters collected during the spring and autumn Massachusetts inshore trawl surveys [1978-2002, all years combined; see Reid et al. (1999) for details]. Page 40 New Hampshire / Winter Skate Massachusetts Inshore Alý Trawl Survey (7 (1978 -2001): "" r " Fall/adults (> 84 cm)Number per tow* 1-10 , 11-100.101-500.->500.Massachusefts

• • * *
• Nantucket Figure 17. cont'd.

Page 41 74' 15 74' 10 WINTER SKATE--Adults (>= 85 cm)Spring (92 -97)Hudson/Raritan Trawl Survey Fish Caught/station (excluding 0 catch): Mean: 1 Min: 1 Max: 2 Length (cm): Min: 85 Max: 92 74* 05 " _74,' 00 73' 55\ -~---Hudson River New York ',> 4Q 0 35'/-J#4Q0 30"/-/Staten Island W/V 4ý"--New Jersey , -j 740 05".. .................. ................. "' Hud, N N 74' 15" 740 10" WINTER SKATE--Adults (>= 85 cm)Fall (92 -96)Hudson/Raritan Trawl Survey Fish Caught/station (excluding 0 catch): Mean: 1 Min: 1 Max: 1 Length (cm): Min: 87 Max: 94 umber/Tow 1-2* 3-5 6 6-10 04"25>10 740 00 730 55".. ............ i ......... ÷......... ...... ...... .... ...... ... .. .. ....... --son River f Ne, NewYork -400 35 I 4400 30" 52 Staten Island"/L~5J/'-5 -,,.,01 Number/Tow 1-2* 3-5* 6-10>10-{400 25" New Jersey Figure 18. Seasonal distribution and abundance of adult winter skate in the Hudson-Raritan estuary, based on Hudson-Raritan trawl surveys, 1992-1997 [see Reid et al. (1999) for details]. Page 42 Winter Skate NEFSC Bottom Trawl Survey Spring/Juveniles 40-30 a)2 20 o- 10 0[I Trawls N=10537 0 Occurrence N=2296 o] Catch N=35461 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 Bottom Temperature (°C)30 25 E 20 15 a 0 o o 0 0 0 0 0 0 0 0 0 0 0 0 000 0 r , I .. .......... ......Bottom Depth (m)0 Trawls N=12193 0 Occurrence N=2744[3 Catch N=46565 50 40 a 30 20 10 0.26 27 28 29 30 31 32 33 34 35 36 37[I Trawls N=1959 0 Occurrence N=507 0l Catch N=4900 Salinity (PPT)Figure 19. Spring and fall distributions of juvenile winter skate and trawls relative to bottom water temperature, depth, and salinity based on NEFSC bottom trawl surveys (1963-2002; all years combined). White bars give the distribution of all the trawls, black bars give the distribution of all trawls in which winter skate occurred, and gray bars represent, within each interval, the percentage of the total number of winter skate caught. Page 43 Winter Skate NEFSC Bottom Trawl Survey Fall/Juveniles 20-15 2 10 0~i,111 [IJUhiirn n n n n n 0 Trawls N=11844 N Occurrence N=1954 El Catch N=20568 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Bottom Temperature (°C)]20 15 0 fL Iý o 'T 0 ? 0 ? c0 T 0 0 N 00 0 0 0 0 c) 0) 0-N. CO O LA 0 N CO O 0 (N O CO Co 0 0 0 0 0 Boo DIeIT9 OA (N C-) -Bottom Depth (in)* Trawls N=1 3543* Occurrence N=2210 0 Catch N=25045 60~40120 -0 27 2 27 29 El Trawls N=1956 N Occurrence N=404[0 Catch N=4255 30 31 32 33 34 35 36 37 Salinity (PPT)Figure 19. cont'd. Page 44 Winter Skate Massachusetts Inshore Trawl Survey Spring/Juveniles 40-30-2 .20~-10-1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Bottom Temperature ('C)[I Trawls N=2312 0 Occurrence N=1 310 o Catch N=86254 40 30 C: 8 20 a)10 0 o3 Trawls N=2385 N Occurrence N=1342 Eo Catch N=90236 4? 0 I$) O I n 0 I ) 0 In L0 0 LnO 0 IO 0 In-O -t I I I P c, Bottom Depth (m)Figure 20. Spring and fall distributions of juvenile winter skate and trawls relative to bottom water temperature and depth based on Massachusetts inshore trawl surveys (1978-2002, all years combined). White bars give the distribution of all the trawls, black bars give the distribution of all trawls in which winter skate occurred, and gray bars represent, within each interval, the percentage of the total number of winter skate caught. Page 45 Winter Skate Massachusetts Inshore Trawl Survey Fall/Juveniles 25 20 c 15 10 0 1 ------4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 Bottom Temperature (°C)0 Trawls N=2067* Occurrence N=958 EO Catch N=46556[o Trawls'N=2156

• Occurrence N=1017 0 Catch N=52536 50 40 30 20 a20~1D-10 0 N NO o co 0 0 LO (0 (o0 to De po i Bottom Depth (m)Figure 20. cont'd.

Page 46 Winter 30 20 1 C 0)C-, a, a-Li Trawls N=468 0 Occurrence N=205 07 Catch N=562 h 10 -0 III II:l II:i lit-.1 1.1 1.1 [E aO EL EL 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Depth (m)40 30 7-n 3[] Trawls N=462 20 0 Occurrence N=205 L a- Catch N=562*0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Temperature (°C)v 20 C 0)C., a, 0~10 E Trawls N=466 I Occurrence N=205 E1Catch N=562 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 Salinity (PPT)0 0 2 4 C 0, 0 a, 0~20 10-0-El Trawls N=265 E Occurrence N=125 11 Catch N=292 r,-I]L I[0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 DO (PPM)Figure 21. Seasonal distributions of juvenile winter skate and trawls relative to bottom water temperature, depth, salinity, and dissolved oxygen based on NEFSC Hudson-Raritan estuary trawl surveys (1992-1997; all years combined). White bars give the distribution of all the trawls, black bars give the distribution of all trawls in which winter skate occurred, and gray bars represent, within each interval, the percentage of the total number of winter skate caught. Page 47 Spring 30 20 10 0 30 20 a).-)n 10 0-- -I" t 1 --T 1 ,f .1 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Depth (m)0I-0 I Trawls N=330 I Occurrence N=1 14-Catch N=262] Trawls N=326 o Occurrence N=1 13 D] Catch N=261 I: 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Temperature (0C)30"E 20 0- 10 0 ED Trawls N=326 m Occurrence N=1 13]I Catch N=261 rL i I11 An El Ill Im[ Illll[ Ill 0 2, .4 8 10 12 14 0 2 .4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 Salinity (PPT)C (D E.-40.-30 -20 10 E: Trawls N=1 77 M Occurrence N=57 El Catch N=105 n U, ý] ii F 0 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 DO (PPM)Figure 21. cont'd. Page 48 Summer 50 40 ,.(D 30 20 10 0 2 4 6 8 10 12 14 18 20 Lw Trawls N=356 o Occurrence N=4 Eo Catch N=7 1 '-,r 16 22 24 26 28 0 Depth (m)C U)0 U)0~50 40 30 20 10 0 U 1K* Trawls N=352* Occurrence N=4 L, Catch N=7 n 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Temperature (0C)C C.)80 70 60 50'40 30 20 10 0 E] Trawls N=347* Occurrence N=4 Eo Catch N=7 1 [1 H 11 r 11 ,I]1 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 Salinity (PPT)E U)_70 60 50 40 30 20 10 0 rPH F 2 ,I Eo Trawls N=215 a Occurrence N=3 El Catch N=5 n i -0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 DO (PPM)Figure 21. cont'd. Page 49 Fall20-a 10 0 0 30 20 (D 0 Li Trawls N=586* Occurrence N=229* Catch N=590 2 I I 4 6 8 10 12 14 16 18 20 22 24 26 28 Depth (m)[I1~I]~El Trawls N=570* Occurrence N=224* Catch N=583 fi r 1 FJ 1 I 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Temperature (°C)C a, C.)a)LU 10 JI Ei Trawls N=532 0 Occurrence N=209 LI Catch N=563 0ý] -CIA, lil I IiIii-I --0 2 4 6 8 10 12 14 16 18 20 22 Salinity (PPT)24 26 28 30 32 34 36 38 40.30 a, H 20 10 0 ir*I Trawls N=458* Occurrence N=183 LI Catch N=509-oil 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 DO (PPM)Figure 21. cont'd. Page 50 Winter 30 C.)20-10 0 50 -El Trawls N= 166 0 Occurrence N=24 El Catch N=1 00 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Depth (m)L- Trawls N=166 Ii n Occurrence N=24 11 Catch N=100 IL 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Temperature (°C)a)a-40 30 20 10 0 C a)C.)a)50 40 30 20 10 0* Trawls N=157* Occurrence N=24* Catch N=100 2 4 6 8 10 12 14 16 18 20 22 24 26 28 0---30 32 34 36 Salinity (PPT)50 -40 30 20 10 0[] Trawls N=1 25 0 Occurrence N=21 m Catch N=90 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 DO (PPM)Figure 22. Seasonal distributions of juvenile and adult winter skate and trawls relative to bottom temperature, depth, salinity, and dissolved oxygen based on Delaware Division of Fish and Wildlife trawl surveys from 1966-1999 (all years combined). White bars give the distribution of all the trawls, black bars give the distribution of all trawls in which winter skate occurred, and gray bars represent, within each interval, the percentage of the total number of winter skate caught. Page 51 Spring C: D 1U_30 20 10 X 1' I F 11LI I, n -L 11 Trawls N=378 0 Occurrence N=50 Ui Catch N=103 0 -i-0 20 2 4 6 8 10 .12 14 16 18 20 22 24 26 28 Depth (m)C o.)(D (L 10[ýIII B 3~*i Trawls N=373* Occurrence N=49 ri Catch N=102ýýiJrr fn 0 0 2 4 6 8 10 12 14 16 18 20 22 Temperature (°C)24 26 28 a)I0 10 Li Trawls N=374* Occurrence N=50*i Catch N=103 U 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 Salinity (PPT)30 20 U.)a)Cn 10 0 F mu rIrI Li Trawls N=312 0 Occurrence N=39 n- Catch N=75 ,I EL 0 1 2 3 4 5 6 7 8 DO (PPM)9 10 11 12 13 14 15 Figure 22. cont'd. Page 52 Fall.C a, 0 a)ci.30 20 10 3 .Trawls N=422 U Occurrence N=1 6* Catch N=39 0 0 2 4 6 8 1.0 12 14 16 18 20 22 24 26 28 Depth (m)40 30 20-0.10 0 2 4 6 Li Trawls N=405 UOccurrence N=16 13 Catch N=39 8 10 12 14 16 18 20 22 24 26 28 Temperature (°C)30-201I a) ° I .I I**10-1 I I 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 Salinity (PPT)Lj Trawls N=399 N Occurrence N=16 El Catch N=39 I Trawls N=341 I Occurrence N=14-Catch N=37 C a, C., a, 0~70-60-50-40-30 20 0 H I 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 DO (PPM)Figure 22. cont'd. Page 53 Winter Skate NEFSC Bottom Trawl Survey Spring/Adults 40 30 220 10 0[I Trawls N=10537 0 Occurrence N=779 Eo Catch N=2458 1 2 3 4 5 6 7 8 9 1011121314151617181920212223 Bottom Temperature (°C)30 215 o o o 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 a- 1 (c';(C J C? i 4 Bottom Depth (Mn)80 L)40 a) I-201 0 ---T- ,=3--T-ý 1ý .. I0 -T- __26 27 28 29 30 31 32 33 34 35 36 37[I Trawls N=12193 M Occurrence N=1 025 Eo Catch N=3562[3 Trawls N=1 959 0 Occurrence N=66 o Catch N=96 Salinity (PPT)Figure 23. Spring and fall distributions of adult winter skate and trawls relative to bottom water temperature, depth, and salinity based on NEFSC bottom trawl surveys (1963-2002; all years combined). White bars give the distribution of all the trawls, black bars give the distribution of all trawls in which winter skate occurred, and gray bars represent, within each interval, the percentage of the total number of winter skate caught. Page 54 Winter Skate NEFSC Bottom Trawl Survey Fall/Adults 30 25" 20 2 15_ 10 5 0[I Trawls N=11844 0 Occurrence N=754[3 Catch N=3142 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Bottom Temperature (°C)40 30 2 20 a. 10 0 100 80 a,-60 40 20 0*1 o 0 C0 D 0 0 0 0 0 0 a0 0 0C0 1? 0? cý U) CO c" c>~0 ( ~- C 0 0 0-------( MC ) t ýBottom Depth (in)[I Trawls N=13543* Occurrence N=932 0 Catch N=4639 (3 Trawls N=1956 0 Occurrence N=104 E3 Catch N=228 i]27 29 30 31 32 33 34 35 36 37 Salinity (PPT)Figure 23. cont'd. Page 55 Winter Skate Massachusetts Inshore Trawl Survey Spring/Adults 25 20-a, 15 -S10 n-5-1 2 3 4 5 B 6 7 8 9 10 11 12 13 14 15 16 Bottom Temperature (°C)0 Trawls N=2312 E Occurrence N=407 0 Catch N=2902 40 30 C 20 a)n 10 0[3 Trawls N=2385 0 Occurrence N=409 El Catch N=2912 U9 0 ) 0 CD 0 UD ) 0 UD 0 C) 0 U ) 0 C3 0 V)w--J CN C) C) I i U? U? I? I? 1 iZ 1- ? 0?C otm M It U) I n -Bottom Depth (in)Figure 24. Spring and fall distributions of adult winter skate and trawls relative to bottom water temperature and depth based on Massachusetts inshore trawl surveys (1978-2002, all years combined). White bars give the distribution of all the trawls, black bars give the distribution of all trawls in which winter skate occurred, and gray bars represent, within each interval, the percentage of the total number of winter skate caught. Page 56 Winter Skate Massachusetts Inshore Trawl Survey Fall/Adults 25 Bottom Temperature (°C)40 10 5 S20 1011 i ~FJlrim L 0 Trawls N=2067 N Occurrence N=217 0 Catch N=1520 0 Trawls N=2156* Occurrence N=221 0 Catch N=1548 U 0 ) 0 LO 0D M) 0= O 0D M 0 U) C= 0 U =) 0 CO O CO C? O Ci O Ci O -CO T ý CO -C ?(0J (' R F (F) A :ý U0 G ) CL O CO ;z Iro-Bottom Depth (m)Figure 24. cont'd. Page 57 Gulf of Maine, Georges Bank, Southern New England, Mid-Atlantic Bight 18000 18 0--0Skate complex, commercial landings (mt)16000 ,-, 16-Survey index of winter skate biomass "O 14000 14,," 12000 --12 .10000 E w 8000 8 8" 6000 6 4000 :%'4 2000 2 "" -2 0 0 C: N T (0 co 0: N~ --t 0 CD 0 N (0 C, 0It(00 C) CO'tO( 0 0C) 0) 0) 0") 0) 0) 0) 0) 0) 0C 0D 0) 0) 0) 0) 0) 0") 0 0) 0-- -------------,- ~- ---N Year Figure 25. NEFSC spring survey index of winter skate biomass and commercial landings of the seven species skate complex from the Gulf of Maine to the Mid-Atlantic Bight.}}