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NOAA Technical Memorandum NMFS-NE-179: Essential Fish Habitat Source Document: Winter Skate, Leucoraja Ocellata, Life History and Habitat Characteristics
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{{#Wiki_filter:C~~tO~ 0 NOAA Technical Memorandum NMFS-NE-179 Ilk'Ars 01F Essential Fish Habitat Source Document: Winter Skate, Leucoraja ocellata, Life History and Habitat Characteristics

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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+ 13 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., 24 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 + 129 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.

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b.I NOAA Technical Memorandum NMFS-NE-179 z m This series represents a secondary level of scientific publishing. All issues.employ QE r - thorough internal scientific review; some issues employ external scientific review. I Reviews are - by design - transparent collegial reviews, not anonymous peer reviews. All issues may be cited in formal scientific communications. EssentialFish HabitatSource 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 19 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 independent data sets from NMFS and several coastal commercial and recreationalfisheries is the continuing states. The species reports are also the source for the loss of marine, estuarine,and other aquatichabitats. current EFH designations by the New England and Mid-Magnuson-Stevens Fishery Conservation and Atlantic Fishery Management Councils, and Management Act (October 11, 1996) understandably have begun to be referred to as the "EFH source documents." The long-term viability of living marine resources NMFS provided guidance to the regional fishery depends on protection of their habitat. management councils for identifying and describing EFH NMFS Strategic Plan for Fisheries of their managed species. Consistent with this guidance, Research (February 1998) the species reports present information on current and historic stock sizes, geographic range, and the period and The Magnuson-Stevens Fishery Conservation and location of major life history stages. The habitats of Management Act (MSFCMA), which was reauthorized managed species are described by the physical, chemical, and amended by the Sustainable Fisheries Act (1996), and biological components of the ecosystem where the requires the eight regional fishery management councils species occur, Information on the habitat requirements is to describe and identify essential fish habitat (EFH) in provided for each'life history stage, and it includes, where their respective regions, to specify actions to conserve available, habitat and environmental variables that control and enhance that EFH, and to minimize the adverse or limit distribution, abundance, growth, reproduction, effects of fishing on EFH. Congress defined EFH as mortality, and productivity. "those waters and substrate necessary to fish for Identifying and describing EFH are the first steps in spawning, breeding, feeding or growth to maturity." The the process of protecting, conserving, and enhancing MSFCMA requires NMFS to assist the regional fishery essential habitats of the managed species. Ultimately, management councils in the implementation of EFH in NMFS, the regional fishery management councils, fishing their respective fishery management plans. participants, Federal and state agencies, and other NMFS has taken a broad view of habitat as the area organizations will have to cooperate to achieve the habitat used by fish throughout their life cycle. Fish use habitat goals established by the MSFCMA. for spawning, feeding, nursery, migration, and shelter, but A historical note: the EFH species reports effectively most habitats provide only a subset of these functions. recommence a series of reports published by the NMFS Fish may change habitats with changes in life history Sandy Hook (New Jersey) Laboratory (now formally stage, seasonal and geographic distributions, abundance, known as the James J. Howard Marine Sciences and interactions with other species. The type of habitat, Laboratory) from 1977 to 1982. These reports, which as well as its attributes and functions, are important for were formally labeled as 'Sandy Hook Laboratory sustaining the production of managed species. Technical Series Reports, but informally known as The Northeast Fisheries Science Center compiled the "Sandy Hook Bluebooks," summarized biological and available information on the distribution, abundance, and fisheries data for 18 economically important species. The habitat requirements for each of the 'species managed by fact that the bluebooks continue to be used two decades the New England and Mid-Atlantic Fishery Management after their publication persuaded us to make their Councils. That information is presented in this series of successors - the 38 EFH source documents - available to 38 EFH species reports (plus one consolidated methods the public through publication in the NOAA Technical report). The EFH species reports are a survey of the Memorandum NMFS-NE series. important literature as well as original analyses of fishery-JAMES J. HOWARD MARINE SCIENCES LABORATORY JEFFREY N. CROSS, (FORMER) CHIEF HIGHLANDS, NEW JERSEY ECOSYSTEMS PROCESSES DIvISION SEPTEMBER 1999 NORTHEAST FISHERIES SCIENCE CENTER

Page iv Page v Contents In trod 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 itat C h a rac te ristic s ...................................................................................................................................................................... 6 S ta tu s o f th e Sto 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 .............. :................... 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' AVERAGE SIZE, MAXIMUM SIZE, AND SIZE AT MATURITY The winter skate [Leucoraja ocellata (Mitchill 1815); formerly Raja ocellata, see McEachran and Dunn (1998); Figure 1] occurs from the south coast of Newfoundland Bigelow and Schroeder (1953) reported winter skate and the southern Gulf of St. Lawrence to Cape Hatteras to have an average size of 76.2-86.4 cm TL. McEachran (Bigelow and Schroeder 1953; McEachran and Musick and Martin (1977) state, that they are one of the larger skates in the Gulf of Maine, with a maximum known size 1975; Scott and Scott 1988; McEachran 2002). Its center of 150 cm TL [Bigelow and Schroeder (1953) give a of abundance is on Georges Bank and in the northern section of the Mid-Atlantic Bight; in both areas it is often maximum length of 106.7 cm TL] with larger individuals more common at higher latitudes. second in abundance to little skate (Leucoraja erinacea), a sympatric species (McEachran and Musick 1975). The size at maturity increases with latitude (McEachran and Martin 1977). On Georges Bank andin Immature winter skate are often confused with the Gulf of Maine, individuals mature between 70-109 cm immature little skate, the distinctions are size-dependent TL. However, in the Gulf of St. Lawrence they mature at (McEachran and Musick 1973; McEachran 2002). a smaller size and do not reach as large a size as other Number of tooth rows, length at maturity, and location of populations (McEachran and Martin 1977). McEachran pelvic denticles are the characters most commonly used to (1973), who studied skates from 1967-1970 that were differentiate the two species (Michalopoulos 1990). 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 LIFE HISTORY mature winter skate was a female 72 cm TL. Bigelow and Schroeder (1953) reported that winter skate does not EGGS mature until at least 63.5-66.0 cm TL. On the eastern Scotian Shelf, Simon and Frank (1998) found that female The single fertilized egg is encapsulated in a leathery, winter skate reached 50% maturity at about 75 cm TL. amber to brown egg capsule which is deposited on the Based on the predictive equations from Frisk et al. bottom (Figure 2). The capsules are rectangular in (2001) and the Northeast Fisheries Science Center outline, with the upper and lower surfaces about equally (NEFSC) survey maximum observed length of 113 cm convex and each corner of the capsule having a long TL, Lmat is estimated at 85 cm TL and Amat is estimated at slender horn (Vladykov 1936; Scott and Scott .1988; 7 years (Northeast Fisheries Science Center 2000b).' 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 FOOD HABITS in width, and are smooth but marked with fine longitudinal striations (McEachran 2002). Generally, polychaetes and amphipods are the most Bigelow and Schroeder (1953) report egg deposition important prey items in terms of numbers or occurrence, to occur during summer and fall off Nova Scotia and, followed by decapods, isopods, bivalves, and fishes quoting Scattergood, probably in the Gulf of Maine as (McEachran 1973; McEachran et al. 1976). Hydroids are well. They also state that egg deposition continues into also ingested (Avent et al. 2001). In terms of weight, December and January off southern New England. 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 JUVENILES and Tsou and Collie 2001a). Bigelow and Schroeder (1953) reported rock crabs and squid as favorite prey, The young are 112-127 mm TL at hatching other items included polychaetes, amphipods, shrimps,. (McEachran 2002) and are fully developed. and razor clams. The fishes that were 'eaten' included smaller skates, eels, alewives, blueback herring, menhaden, smelt, sand lance, chub mackerel, butterfish, ADULTS cunners, sculpins, silver hake, and tomcod. McEachran (1973) studied skates collected from Female winter skates with fully formed egg capsules Nova Scotia to Cape Hatteras during 1967-1970; the are more abundant during the summer and fall but it is following diet descriptions are from him and McEachran possible that reproduction takes place to some degree et al. (1976). throughout the year (Vladykov 1936; Scott and Scott Nephtys spp., Nereis spp., Lumbrineris. fragilis, 1988; McEachran 2002). 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 consisted of crustaceans, with 38-43% of the diet were the most frequently consumed prey in the Gulf of consisting of identifiable amphipods. The most abundant Maine and on the Nova Scotian shelf. amphipod species included Unciola irrorata, Byblis Haustoriids, Leptocheirus pinguis, Monoculodes sp., serrata, and H. serratus. Identifiable decapods made up Hippomedon serratus, ampeliscids, Paraphoxussp., and 23-25% of the diet, most of which were species such as Tmetonyx sp. were the most frequently eaten amphipods C. septemspinosa and C. irroratus. Identifiable over the survey area. Crangon septemspinosa was the polychaete species (9-13% of the diet) included most abundant decapod in the diet. Cancer irroratus, Ampharete arctica. Identifiable isopod species (9% of the Dichelopandalus leptocerus, Pagurus acadianus, and diet) included Cirolana (= Politolana?) polita. Hyas sp. were consistently eaten but in small numbers. Nematodes, bivalves, and fish were included in the "other Among the minor prey items included Cirolana ( prey phyla" category (3-17% of the diet). Politolana?) polita, which was the dominant isopod. For skate 31-40 cm TL, 72-76% of the diet consisted Other isopods eaten included Chiridoteatuftsi and Edotea of crustaceans, with 37-39% of the diet consisting of triloba, but they contributed little to the overall diet. The identifiable amphipods. Major amphipod species included only identifiable bivalves eaten were Solemya sp. and B. serrata, U. irrorata, H. serratus, and several Ensis directus. The most frequently eaten fish was sand unidentified haustoriids. Identifiable decapods made up lance, while yellowtail flounder and longhorn sculpin 17-23% of the diet, most of which were C. septemspinosa were occasionally eaten. and C. irroratus.Identifiable polychaetes (12-17% of the In Smith's (1950) study in Block Island Sound, diet) included Scalibregma inflatum, L. fragilis, and nekton were more important and isopods much less unidentified maldanids. Identifiable isopods (5-8% of the important than in the McEachran (1973) and McEachran diet) included Cirolana (= Politolana?) polita. et al. (1976) studies. The major prey items for winter Miscellaneous items (6-9% of the diet) included skate in Block Island Sound included nekton, L. pinguis, nematodes and bivalves. Among the identifiable fish Nephtys incisa, E. directus, C. septemspinosa, Nereis sp., present in the diet (3-4%) were sand lance, yellowtail C. irroratus,Lumbrineris sp. and Monoculodes edwardsi. flounder, and hakes. Winter skate from Georges Bank had the most The percentage of crustaceans in the diet of winter diverse diet and those from the Mid-Atlantic Bight the skate 41-50 cm TL dropped to 62-69%, although least diverse diet (McEachran 1973; McEachran et al. identifiable amphipods still made up the major portion 1976). There was no significant change in the diet with (33-35%) followed by decapods (14-22%). Identifiable increase in skate size; however, the numbers of polychaetes made up 19-23% of the diet; other prey polychaetes gradually increased and amphipods gradually species (including mollusca), 6-9% of the diet; decreased with increasing skate size. The number of fish identifiable isopods, 7% of the diet; and identifiable fish, and bivalves also increased with predator size and the two 3-8% of the diet. All the major prey species (except for taxa were a major part of the diet of skate > 79 cm TL. the lack of the polychaete S. inflatum) were similar to the The ingestion of decapods was independent of skate size. 31-40 cm TL size class, with the additions of several There was also no indication of either diel or seasonal more Unciola species, L. pinguis (an amphipod), periodicity in intensity of feeding. In Passamaquoddy Bay unidentified pagurid crabs, and nephtyid polychaetes. there were no differences between the diets of small and The percent occurrence of crustaceans in the diet of large winter skate (Tyler 1972). winter skate 51-60 cm TL dropped further, down to 53-The 1973-1990 NEFSC food habits database for 54%, with identifiable amphipods making up only 26-winter skate [Figure 3; see Reid et al. (1999) for details] 32% of the overall diet. Some of the dominant identifiable generally confirms the McEachran (1973) and McEachran amphipods included Psammonyx nobilis, unidentified et al. (1976) studies. Crustaceans made up > 50% of the oedicerotids, H. serratus, and unidentified haustoriids. diet for skate < 61 cm TL, while fish dominated the diet Identifiable decapods made up only 9-12% of the diet; C. of skate > 91 cm TL. Overall crustaceans declined in septemspinosa was again the dominant decapod prey, importance with increasing skate size (includes both followed by C. irroratus and pagurid crabs. Cirolana (= amphipods and decapods) while the percent occurrence of Politolana?) polita was again one of the major polychaetes increased with -increasing skate size until the identifiable isopods, which all together made up 7-12% of skate were about 81 cm TL. Amphipods occurred more the diet. The percent occurrence of identifiable frequently than decapods until the skates were > 71 cm polychaetes continued to increase in the diet, up to 26-TL. Among the most frequently occurring prey species 29%; several of the more numerous species present were for almost all sizes of skate included the decapods C. in the genera Nephtys and Nereis. Identifiable fish also septemspinosa and Cancer and pagurid crabs, the isopod increased in the diet, up to 6-13%, with sand lance the Cirolana (= Politolana?) polita, and sand lance. The dominant species. Other prey phyla, including bivalves following is a detailed description of the diet from the and nematodes, accounted for 9-11% of the diet. NEFSC food habits database broken down by winter The percent occurrence of crustaceans in the diet skate size class (Figure 3). continued to decline for winter skate 61-70 cm TL: down For winter skate 21-30 cm TL, 74-84% of the diet to 38-44%, with identifiable amphipods making up only

Page 3 13-20% of the diet, while identifiable decapods made up with the genus Nephtys the most notable. "Other prey. 11-12%. Major amphipod species included M. edwardsi, phyla" and identifiable mollusks together accounted for U. irrorata,H. serratus, and unidentified haustoriids and 10-12% of the diet, bivalves and nematodes dominated oedicerotids. C. septemspinosci continued to be the this category. dominant decapod prey, followed by Cancer and pagurid Finally, identifiable fish made up > 60% of the diet crabs. Identifiable isopods again made up 7-12% of the of 101-110 cm TL winter skate from .the 1981-1990 diet; Cirolana(= Politolana?)polita continued .to be one NEFSC trawl surveys. Most were sand lance. Mollusks of the major prey species. The percent occurrence of were 14% of the diet, polychaetes were 13% of the diet, identifiable polychaetes in the diet increased, up to 28- and crustaceans were down to 11% of the diet. 32%; species in the genera Nephtys and Nereis were again Using NEFSC data from 1977-1980, Bowman et al. dominant. The percent occurrence of identifiable fish in (2000) found that in terms of percent weight, crustaceans the diet continued to increase also, up to 11-24%, most of were dominant in the diet of skate < 31-50 cm TL, while which were sand lance. Nine percent of the diet consisted fish, mostly sand lance, were dominant in the diet of skate of identifiable mollusks, with bivalves being dominant. 51-110 cm TL. For skate < 31 cm TL, amphipods While the percent occurrence of crustaceans dropped dominated, especially L. pinguis. For skate 31-50 cm TL, to 29-36% for winter skate 71-80 cm TL, the percent decapods dominated, especially C. septemspinosa and C. occurrence of identifiable decapods was greater than the irroratus. On Georges Bank Tsou and Collie (2001a), percent occurrence of amphipods: 11-13% versus 7-12%. using NEFSC dietary data from 1989-1990, also showed The former were dominated by C. septemspinosa, Cancer that fish, especially sand lance, were most important for and pagurid crabs, and D. leptocerus, while several winter skate > 50 cm TL. Other noted fish prey included haustoriid species and U. irroratawere some of the major sliver hake, mackerel, and herring (see also Tsou and amphipod prey. Identifiable isopods made up 8-9% of the Collie 2001b). diet, the dominant species continued to be Cirolana (= Nelson (1993) calculated the predation impact of Politolana?)polita. Identifiable polychaetes (25-35% of winter skate on their Georges Bank prey. Annual the diet) included L. fragilis and several Nephrys and estimates of consumption for winter skate increased as Nereis species. The percent occurrence of identifiable fish they grew larger. Consumption ranged from 1.186 in the diet varied widely between the two sampling kg/fish/year for skate 40-49 cm TL to 5.528 kg/fish/year periods, from 16-36%, although sand lance was still the for skate 90-99 cm TL. The percentage of benthic dominant species. Identifiable mollusks made up 9-10% production consumed by winter skate from 1969-1990 of the diet, most of which were bivalves. ranged from 11-43%. Nelson (1993) suggests that in Fish as prey items became increasingly important for relation to the total macrofauna production on Georges winter skate 81-90 cm TL. They made up 29-42% of the Bank, winter skate (along with little skate) consume < overall diet. As usual sand lance were the dominant fish 0.02% of the total. These results indicate that only a small prey, other species ingested included other skate, to moderate proportion of benthic biomass vulnerable to longhorn sculpin, and silver hake. Crustaceans in the diet skate predation is consumed by both winter and little declined to 19-30%. The major identifiable decapod skate, and their consumptive impact will be dependent on species (8-11% of the diet) continued to be C. the levels of invertebrate biomass and/or production. 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) PREDATORS AND SPECIES were several haustoriid species. Cirolana (= Politolana?) ASSOCIATIONS polita was once again the dominant identifiable, isopod (all isopods together made up 5-7% of the diet). Several Winter skate is preyed upon by sharks, other skates, Nephtys species were the major identifiable polychaetes gray seals, and gulls (Scott and Scott 1988; Kaplan 1999). ingested, all polychaetes together made up 22-28% of the McEachran and Musick (1975) state that winter and diet. Bivalves, particularly of the familiy Solenidae, were little skate co-occurred significantly during 1967-1970 the dominant identifiable molluscan prey ingested, with surveys from Nova Scotia to Cape Hatteras. Although all mollusks together accounting for 7-17% of the diet. winter and little skate are sympatric species with similar Identifiable fish made up >50% of the diet of winter habitat requirements (except perhaps temperature skate 91-100 cm TL. Sand lance was the overwhelming preference), there does not appear to be a high degree of dominant, some of the minor fish prey included silver competitive interaction between them because they are hake, herring, and butterfish. Crustaceans were down to positively correlated by abundance and where the two 12-23% of the diet'. Identifiable decapods made up 5-10% species are most abundant (Georges Bank) they have the of the diet, C. septemspinosa, Cancer and pagurid crabs, most similar diets and highest diversity of assemblages of D. leptocerus, and pandalid shrimp were some of the prey species (McEachran 1973; McEachran and Musick major decapods ingested. Identifiable amphipods made up 1975; McEachran et al. 1976). only 4-5% of the total diet, with few conspicuous species. Also, even though the two species do consume the Identifiable polychaetes accounted for 10-13% of the diet, same large taxonomic groups of benthic fauna

Page 4 (amphipods, decapods, and polychaetes), winter skate 1963-1978. Of the five assemblage species groups or predominately feeds on infaunal organisms while little associations present on Georges Bank in spring and fall skate feeds largely on epifauna (McEachran 1973; throughout the survey period, winter skate belonged to McEachran et al. 1976). McEachran (1973) and the "Intermediate" and "Shallow" assemblage groups. In McEachran et al. (1976) show that large burrowing .the Shallow assemblage the other major species present polychaetes and bivalves were consumed more frequently besides winter skate included Atlantic cod, little skate, by winter skate and epifaunal decapods were eaten more longhorn sculpin, yellowtail flounder, and haddock; in the frequently by little skate. Winter skate ate more Intermediate assemblage, little skate, red and silver hake, burrowing amphipods, especially haustoriids and Atlantic cod, and haddock were some of the other major Trichophoxus epistomus while little skate consumed more species present. Overholtz and Tyler (1985) considered surface dwelling amphipods such as Unciola sp., Dulichia winter skate to be a "resident" species, since they were (= Dyopedos) monacantha, ampeliscids and caprellids. only present in two out of the five assemblages in The division of food resources between the skates is not abundance. The Shallow assemblage covered most of complete because some winter skate ate large numbers of Georges Bank in the spring and was slightly smaller in epifauna and some little skate consumed large numbers of the fall. The Intermediate assemblage occurred mostly infauna. Both species ate considerable numbers of L. south of the Shallow assemblage and inside the southern pinguis and C. septemspinosa. Little skate occasionally edge of Georges Bank; it was somewhat larger in the fall, fed on haustoriids, and deep burrowing polychaetes were suggesting a migration of the species in this area to regular prey items. The infaunal and. epifaunal shallower water as the year progressed. The assemblages preferences of the two skates may be more distinct in in the spring appeared to follow depth contours. areas where they may coexist than in areas where they Garrison (2000) and Garrison and Link (2000b) have occur separately because in Delaware Bay (Fitz and also investigated spatial assemblages and trophic groups Daiber 1963) little skate consumed relatively more from the Georges Bank region. Using 1963-1997 NEFSC infauna than it did in the areas sampled in Smith's (1950) trawl survey data from Georges Bank, as well as the same study or the McEachran (1973) and McEachran et al. NEFSC food habits database discussed above (Garrison (1976) studies. Winter skate does not regularly occur in and Link [2000b] used 1973-1997 data while Garrison Delaware Bay (Fitz and Daiber 1963; see also Delaware [2000] used 1991-1997 data), they found that the major Bay trawl surveys, below). predator groups were consistent across decades, with the

. In addition, differences in the shape and size of the boundaries of the assemblages similar to Overholtz and mouth and the number of tooth rows between the two Tyler (1985). Garrison (2000) investigated the spatial species were used as evidence by McEachran and Martin assemblages during spring and autumn. He found that (1977) to suggest that the sympatric populations of winter during autumn, winter skate was in the assemblage found and little skate underwent character displacement in order in the deep habitats on southern Georges Bank that also to avoid direct competition for food resources. In included spiny dogfish, butterfish, little skate, red hake, sympatric. populations, winter skate has greater number of fourspot flounder, and yellowtail flounder. The main tooth rows in the upper jaw and a wider and less arched shallow portion of Georges Bank assemblage included 31 mouth, thus allowing them to feed more efficiently and cm to > 80 cm TL winter skate, little skate, spiny dogfish, deeper in the bottom than little skate. Little skate has a Atlantic cod, windowpane, winter flounder, and sea relatively smaller and more arched mouth with fewer raven. In spring, the main shallow portion of Georges tooth rows in the upper jaw. Bank assemblage included 31 cm to > 80 cm TL winter Using 1973-1997 NEFSC data from Nova Scotia to skate, haddock, Atlantic cod, and spiny dogfish. In terms Cape Hatteras, as well as the same NEFSC food habits of dietary guilds or trophic groups, the two studies had database discussed above, Garrison and Link (2000a) slightly different viewpoints, but the diets of winter skate investigated the dietary guild structure of the fish in both studies were the same as discussed in the Food community. Both small (10-30 cm TL) and medium (31- Habits section above. In the Garrison and Link (2000b) 60 cm TL) sized winter skate belonged to the study, winter skate fell into two predator or feeding "Amphipod/shrimp eaters" group, along with little skate groups. The first was a "Bentho-pelagic" group, which and cusk eel; prey included amphipods, polychaetes, included 31-60 cm TL winter skate, little skate, longhorn shrimp, and zooplankton. The largest winter skate (61 to sculpin, and Atlantic cod. The diets of these species were

> 80 cm TL) were by themselves in a subgroup of the same as that discussed above for winter skate alone: "Piscivores" because, as described previously under the shrimp such as pandalids and C. septemspinosa, and detailed description of the diet using the 1973-1990 benthic invertebrates including polychaetes, gammarid NEFSC food habits database, their diet contained a high amphipods, and bivalves. The second group was the proportion of sand lance. This again shows that there is a "Skate" group, consisting of > 80 cm TL winter skate, trend toward increasing piscivory with size. pollock, and windowpane. Their prey was characterized The resilience of demersal fish assemblages on by a combination of fish and benthic prey, with a high Georges Bank was investigated by Overholtz and Tyler proportion of sand lance during the 1980s. Garrison (1985) using seasonal NEFSC trawl survey data from (2000) had slightly different trophic groups. In autumn,

Page 5 the largest (61 cm. to > 80 cm TL) winter skate were by many of these records, as well as those for little skate themselves in a subgroup of the "Piscivorous" group, from the same localities, are suspect. feeding on sand lance, silver hake, and Atlantic herring, . Their range extends from southern New England and as well as benthic invertebrates. Small and medium (10- down the Mid-Atlantic Bight to northern North Carolina 60 cm TL) winter skate were also in the "Demersal (Bigelow and Schroeder 1953; Figures 4-7, 13-16). predators" group, along with flatfish, haddocks, little Previous authors have suggested that winter skate skate, and thorny skate (Amblyraja radiata). Prey undertakes seasonal movements, especially in the included gammarid amphipods, polychaetes, isopods, and southern part of its range, moving shoreward in autumn Cancer crabs, as well as C. septemspinosa. During spring, and offshore in summer (McEachran 1973; McEachran 10-60 cm TL winter skate were in the "Shrimp/amphipod and Musick 1975; McEachran 2002), although this is not predators" group, along with hakes, longhorn sculpin, quite evident from the overall NEFSC bottom trawl Atlantic cod, fourspot flounder, little skate, and thorny surveys (Figures 4-7, 13-16; see descriptions below). In skate. Prey included gammarid amphipods, pandalids and Passamaquoddy Bay, Tyler (1971) reported winter skate C. septemspinosa, polychaetes, and Cancercrabs. Winter present from December to March while Huntsman (1922) skate 61 cm to > 80 cm TL were again by themselves in a stated it was abundant from May to November. subgroup called the "Generalist" group, consuming McEachran (1973) suggests. this disparity may be due to a bivalves, polychaetes, sand lance, and herring. The difference in the areas the two authors sampled. Tyler decline in importance of fish prey, 35% fish in the (1971) sampled the deeper waters of Passamaquoddy Bay autumn and 16% in the spring, is probably related to while Huntsman (1922) did not specify the sampling seasonal movements of prey (Garrison 2000). depths. However Macdonald et al. (1984) determined On the Scotian Shelf, Scott (1989), using research winter skate to be both a regular and occasional resident trawl survey data from roughly 1970-1984 determined in Passamaquoddy Bay, and juveniles were often found at that winter skate was locally abundant but did not beach sites during summer. Merriman and Warfel (1948) associate closely with any other species. However Mahon stated it is a permanent resident off southern New (1997), analyzing trawl survey data for the same region England although there are seasonal fluctuations in from 1970-1993, showed that winter skate whs associated abundance; Bigelow and Schroeder (1953) mention that with such species as longhorn sculpin, sea raven and along the southern coast of New England it comes inshore winter flounder in the shallow waters of the Bay of Fundy near Woods Hole during the colder months. However and Sable Island. 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 GEOGRAPHICAL DISTRIBUTION skate present, but was much less abundant). Schaefer (1967) found winter skate in the surf zone of Long Island In Canada, winter skate are found in the Gulf of St. during May, June, October, and November, while Lawrence, off northeastern Nova Scotia, and the offshore Gottschall et al. (2000), based on surveys from 1984-banks of Banquereau, Sable Island, and Western Bank 1994 (see Habitat Characteristics section, below), found [Bigelow and Schroeder 1953; McEachran and Musick their lowest abundances in Long Island Sound were in the 1975; see'also Strong and Hanke (1995) for the 1970- months of July, August, and September. McEachran and 1993 distribution of winter skate in the Scotia-Fundy Musick (1975) reported winter skate to be more abundant region; further information on winter skate distribution on south of Delaware Bay during the winter, it has also been the Scotian Shelf can be found in Simon and Frank (1996, reported from Chesapeake Bay from December to April 1998)]. The population in the southern Gulf of St. (Hildebrand and Schroeder 1928; Geer 2002). 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 JUVENILES little skate from the Gulf of St. Lawrence and most records of it from northern Nova Scotia probably refer to NEFSC bottom trawl surveys [see Reid et al. (1999) winter skate. for details] captured juvenile (< 84 cm TL) winter skate Winter skate is considered common all around the year-round. (Note that winter and summer distributions Gulf of Maine from Nova Scotia to Cape Cod, except for are presented as presence/absence data, precluding a the deep troughs, and is also common on Georges Bank discussion of abundances.) In winter, juveniles were (Bigelow and Schroeder 1953; McEachran 2002). It has found from Georges Bank to Cape Hatteras, out to the .frequently been reported from the Bay of Fundy, and the 200 m depth contour (Figure 4). Concentrations were coasts of Maine and Massachusetts (Bigelow and found off Long Island and southern New England; they Schroeder 1953) and in Massachusetts Bay (Collette and were almost entirely absent from the Gulf of Maine. In Hartel 1988), as well as along the New Hampshire coast spring they were also found from Georges Bank to Cape (Nelson et al. 1983). However, McEachran (2002) states Hatteras, and were concentrated nearshore throughout the that because of its close resemblance to the little skate, Mid-Atlantic Bight and southern New England as well as

Page 6 in Cape Cod and Massachusetts Bays (Figure 5). Small from Georges Bank to North Carolina; very few occurred numbers were also found along the coast of Maine and in the Gulf of Maine (Figure 13). In the spring, they were southwest Nova Scotia and near Browns Bank. also found from Georges Bank to North Carolina but, as Comparatively few were present in summer, with with the juveniles, were also distributed nearshore concentrations on Georges Bank and around Cape Cod throughout the Mid-Atlantic Bight and along Long Island (Figure 6). Some were also found near Penobscot Bay, as well as around Cape Cod and Massachusetts Bays Maine. Winter skate abundances in the fall were not as (Figure 14). Small numbers were also found along the high as in the spring (Figure 7). In the fall they were coast of Maine and southwest Nova Scotia near Browns collected from Georges Bank to the Delmarva Peninsula Bank. Few occurred in summer, being found mostly on and were again concentrated along Long Island, southern Georges Bank, Nantucket Shoals, and near Cape Cod New England, around Cape Cod, and on Georges Bank. (Figure 15). In the fall, they were mostly confined to Small numbers were again found along the coast of Georges Bank, near Nantucket shoals, and near Cape Maine and near Browns Bank. Cod, with very few found south of those areas (Figure Both the spring and fall 1978-2002 Massachusetts 16). inshore trawl surveys [see Reid et al. (1999) for details] Adult little skate were collected in much fewer show similar abundances and distributions of juveniles numbers than juveniles during the spring and fall (Figure 8). The highest concentrations were found on the Massachusetts inshore trawl surveys I(Figure 17). The Atlantic side of Cape Cod and south and west of Martha's greatest numbers were found on the Atlantic side of Cape Vineyard (especially in spring) and south and northeast of Cod and, in spring, south of Nantucket. Nantucket (also in spring). Large numbers were also Very few adults were caught in the Hudson-Raritan found near Monomy Point in the fall. Other notable estuary during spring and fall (Figure 18). Those few that occurrences of winter skate were around Plum Island, were caught were concentrated around the Ambrose and Ipswich Bay, north of Cape Ann, near Nahant Bay Chapel Hill Channels. (especially in the fall), in Cape Cod Bay, and in The seasonal distribution and abundance of both Nantucket Sound. adults and juveniles in Delaware Bay were discussed The distributions and abundances of both juveniles previously (Figure 12). and adults in Long Island Sound (Figures 9-10) as described by Gottschall et al. (2000) will be discussed in the Habitat Characteristics section. HABITAT CHARACTERISTICS Occurrence of juveniles in the Hudson-Raritan estuary appears to have the same seasonal pattern noted Information on the habitat requirements and by previous authors for other estuaries; i.e., they're preferences of winter skate (based on both the pertinent generally absent from the estuary during the summer literature and the most recent NEFSC and state surveys) months. Juveniles were fairly well distributed throughout are presented here and summarized in Tables 1 and 2. the Hudson-Raritan estuary in winter, spring and fall, and Winter skate generally ranges from the shoreline to were most abundant in the winter and fall (Figure 11). In 371 m, although it is most abundant at depths < 11l m summer the few that were left were mostly confined to (Bigelow and Schroeder 1953; McEachran 1973; the deeper and warmer waters of the Ambrose Channel. McEachran and Musick 1975; McEachran 2002). The 1966-1999 Delaware Bay trawl surveys. (adults Bigelow and Schroeder (1953) suggest that feware and juveniles combined; Figure 12) also confirm the caught > 91 m. In the Gulf of Maine, they are most seasonal trends noted previously for winter skate, abundant 46-64 m but occasionally occur < I m as well as although they were not very abundant in the Bay overall. down to 285 m; they're considered rare at depths < 2-7 m They were almost completely absent in summer, and a (Bigelow and Schroeder 1953; McEachran 2002). On the few were caught in the fall, while the greatest numbers Scotian Shelf the winter skate is most frequently caught were found in the winter and spring. The skate were most between 37-90 m (Scott 1982a). That segment of the abundant in the center of lower Delaware Bay, near the population residing at depths > 10 m appear to be resident mouth (Figure 12). year-round, even though the coastal. edge of the population appears to move shoreward in autumn and offshore in summer (Bigelow and Schroeder 1953). ADULTS Merriman and Warfel (1948) stated that winter skate is a permanent resident off southern New England between NEFSC bottom trawl surveys [see Reid et al. (1999) 15-46 m although there are seasonal fluctuations in for details] captured adult winter skate (> 85 cm TL) abundance. The 1978-2002 spring and autumn during all seasons (Figures 13-16). The numbers of adults Massachusetts inshore trawl surveys (see below) show in spring and fall were much lower than for juveniles of that both juveniles and adults were found between 1-75 the same two seasons (winter and summer distributions m, with most found between 6-25 m (Figures 20 and 24). are presented as presence/absence data, precluding a Edwards et al. (1962) captured it off the Mid-Atlantic discussion of abundances). In winter, they were scattered states during the winter at depths from 33-113 m. The

Page 7 1963-2002 spring and fall NEFSC trawl surveys from the JUVENILES Gulf of Maine to Cape Hatteras (see below) indicated that most juveniles occurred at depths < 70-80 m, although a The spring and fall distributions of juvenile winter few occurred as deep as 400 m (Figure 19), while most skate relative to bottom water temperature, depth, and adults were found < 70 m and a few were as deep as 300 salinity based on 1963-2002 NEFSC bottom trawl m (Figure 23). In the Hudson-Raritan estuary (see below; surveys from the Gulf of Maine to Cape Hatteras are Figure 21), juvenile winter skate are found from about 4- shown in Figure 19. In spring, they were found in waters 22 m, but occur mostly around 5-8 m during a good part between 1-12'C, with the majority at about 4-5°C. Their of the year. In Delaware Bay (see below; Figure 22) depth range during that season was between 1-300 m, juveniles and adults were found over a range of with most between about 11-70 m. They were found at approximately 7-21 m during winter, spring, and fall. salinities between 28-35 ppt, with most found between Winter skate has been recorded over a temperature 32-33 ppt. During the fall, juvenile winter skate were range of -1.2°C tol9°C (Bigelow and Schroeder 1953; caught over a temperature range of 5-21 °C, with most Tyler 1971; McEachran 1973; McEachran 2002). On the spread between about 7-16'C, and peaks at about 13-Scotian Shelf it has been frequently found at temperatures 15'C. They were found over a depth range of 1-400 m, of 5-9°C (Scott 1982a; Scott and Scott 1988). Bigelow although most were caught at depths between about 21-80 and Schroeder (1953) reported them in the Gulf of Maine m. They were found at salinities between 31-35 ppt, with at around 20'C along the Massachusetts coast in the the majority found between 32-33 ppt. summer, down to 1-2 C in the coastal belt in winter, and The spring and autumn distributions of juveniles in near 0°C in the Bay of Fundy region in some years. Massachusetts coastal waters relative to bottom water McEachran and Musick (1975) reported their temperature temperature and depth based on .. 1978-2002 ranges at depth of capture were -1.2°C to 4.8°C in the Massachusetts inshore trawl surveys are shown in Figure Gulf of St. Lawrence, 1.1-12.7'C off northeastern Nova 20. In the spring they were found in waters ranging from Scotia, and 2-15°C from southern Nova Scotia to Cape 3-15'C, with the greatest percentages found between Hatteras. Edwards et al. (1962) captured it off the Mid- approximately 8-12'C. Their depth range was from Atlantic states during the winter at temperatures from 10- approximately 6-75 m, with the majority at 6-25 m. 12'C. The 1963-2002 spring and fall NEFSC trawl During the autumn they were found in waters ranging surveys from the Gulf of Maine to Cape Hatteras (see from 5-21 'C; their temperature distribution was below) collected juvenile winter skate over a somewhat bimodal, with the major peaks between about temperatures range of 1-21 'C, with most found between 16-18'C. Their depth range was from 1-65 m, with the 4-5°C in the spring and about 7-16'C in the fall (Figure majority found between 6-25 m. 19). Adults were found over a temperature range of 2- The distributions and abundances of both juvenile 19'C with most found around 5'C in spring and between and adult winter skate in Long Island Sound from April to about 1 -15C in the fall (Figure 23). November 1984-1994, based on the Connecticut Fisheries Bigelow and Schroeder (1953) stated that this species Division bottom trawl surveys, are shown in Figures 9-is confined to sandy and gravelly bottoms but Tyler 10. The following description of their distributions (1971) reported it from mud bottoms in Passamaquoddy relative to depth and bottom type is taken verbatim from Bay. In Long Island Sound during the spring, winter skate Gottschall et al. (2000). were most abundant on sand bottoms in the Mattituck Sill Winter skate were most commonly taken during the and Eastern Basin (Gottschall et al. 2000). On the Scotian spring and late fall, occurring on average in. 16.4% of Shelf, Scott (1982b) reports that the distribution of winter samples during these periods (Figure IOD). Abundance skate was confined to sand and gravel bottoms and Scott was highest during April, and decreased thereafter until (1982b) suggests that bottom type, rather than depth, August when none were recorded in the survey (Figure appears more important in determining the distributions I OA). During the spring, winter skate were most abundant of winter skate. on sand bottom in the Mattituck Sill and Eastern Basin Winter skates are known to remain buried in (Figure 9, Figure 10B). Abundance was similar in most depressions during the day and are more active at night depths, with the exception of depths between 9-18 m, (Michalopoulos 1990). This is probably not due to diel where abundance was lower (Figure 10C). Winter skate foraging, since McEachran et al. (1976) observed no diel abundance increased again in October and November, but periodicity in feeding intensity by winter skate and they were not as concentrated on the Mattituck Sill and in suggested that they may feed at any time during a 24 hour the Eastern Basin as during the spring (Gottschall et al. period. 2000). Scott (1982a) mentions that on the Scotian Shelf The seasonal distributions of juveniles the Hudson-during the summers of 1970-1979, winter skate was Raritan estuary relative to bottom water temperature, found at preferred salinities of 32-34 ppt. 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 fall was spread between 7-11 m and at 13 m and between most caught between 5-8 m. Their salinities ranged 18-19 m. Most were spread between 7-8 m, at 13 m, and between 20-35 ppt, most were found roughly between 23- at 18 m. Their salinities ranged between about 26-32 ppt, 32 ppt. They were found over a range of dissolved *with a few at 16 ppt and 22 ppt. Peaks were at 28 ppt, 30 oxygen levels of between 9-14 ppm with a few at 5 ppm; ppt, and at 32 ppt. They were found over a range of most were found between 10-12 ppm. In spring, juvenile dissolved oxygen levels of between 7-10 ppm, the winter skate were found over a wider temperature range majority were at 9 ppm. 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 ADULTS greater number of trawls done within those temperature intervals. Their depth range was between 4-18 m, with the The spring and fall distributions of adult winter skate majority between 5-8 m. Their salinities ranged between relative to bottom water temperature, depth, and salinity 15-33 ppt, most were found at 25 ppt and between 27-28 based on 1963-2002 NEFSC bottom trawl surveys from ppt. They were found over a range of dissolved oxygen the Gulf of Maine to Cape Hatteras are shown in Figure levels of between 7-13 ppm with most found between 10- 23. In spring, adults were caught at temperatures between 11 ppm. Few were caught in summer; they were found 2-11 'C, with most between 4-6°C and a peak at 5OC. between about 16-21 *C and at depths of 7 m, 18 m, and During that period they were found at a depth range of 1-20 m. Their salinities ranged between 28-29ppt and at 32 300 m, with the majority spread between 31-60 m. They ppt, and they were found over a. range of dissolved were found at salinities of between 30-36 ppt, with the oxygen levels of between 7-8 ppm. In the fall they were majority at 33 ppt. During the fall, they were caught over found between 5-17'C, with most spread between 5- a temperature range of 5-19'C, with most caught between 13 'C. Their depth range during the fall was between 4-21 about I I-15°C and a peak at 14'C. They were found over m, with the majority between 5-8 m. Their salinities a depth range of 11-300 m, with most caught at depths ranged between 17-34 ppt, with most spread roughly between about 21-70 m and peaks at 31-50 m. They were between 23-31 ppt. They were found over a range of found at salinities of between 31-34 ppt, with 80-90% at dissolved oxygen levels of between 6-12 ppm with most 32 ppt. found between 8-9 ppm. Based on the above evidence it The spring and autumn distributions of adults in appears that juvenile winter skate in the Hudson-Raritan Massachusetts coastal waters relative to bottom water estuary are found in warmer waters during the spring and temperature and depth are shown in Figure 24. In the fall as compared to winter, and remain mostly around spring they were found in waters ranging from 2-16'C; depths of 5-8 m during those three seasons. the majority were spread between approximately 6-12'C. The seasonal distributions of both juveniles and During that same season, the adults were found from 1-75 adults in Delaware Bay relative to bottom water m, with most between 6-20 m. In autumn they were found temperature, depth, salinity, and dissolved oxygen based between 5-19'C. The distribution was somewhat on 1966-1999 Delaware Division of Fish and Wildlife bimodal, with a peak at 10C and a minor one around 15-bottom trawl surveys are shown in Figure 22. During the 16'C. The depth range of the adults during autumn was winter they were found between 3-9°C, with the majority aroun'd 1-75 m, with most found between 6-25 m. between 7-8°C. Their depth range during winter was The distributions and abundances of both juvenile between about 7-18 m, with peaks at 14 m, 16 m, and 17 and adult winter skate in Long Island Sound relative to

m. Their salinities ranged between about 22-30 ppt and depth and bottom type were discussed previously (Figures 34-35 ppt, most were found at 26 ppt and between 28-29 9-10; Gottschall et al. [2000]).

ppt. They were found over a range of dissolved oxygen Too few adults were found in the Hudson-Raritan levels of between 8-11 ppm, with the majority found estuary to plot their distributions relative to habitat between 9-11 ppm. In spring, they were found over a parameters. wider temperature range of between 4-17'C, with peaks The seasonal distributions of both juveniles and scattered throughout the range (e.g., 5°C, I l°C, and adults in Delaware Bay relative to bottom water 13'C). Their depth range was between 7-17 m, with a temperature, depth, salinity, and dissolved oxygen based few at 21 m, and most at 8 m and 12-14 m. Their on Delaware Division of Fish and Wildlife bottom trawl salinities ranged between 21-33 ppt, with a few at 15 ppt, surveys were discussed previously (Figure 22). 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- STATUS OF THE STOCKS 15 ppm, most were found between 8-11 ppm. In summer, there were too few winter skate caught to plot their The following section is based on Northeast Fisheries distributions relative to the habitat parameters. During fall Science Center (2000a, b). they were found between 8-13'C, with a few at 16'C; The principal commercial fishing method used to most were between 8-11 'C. Their depth range during the

Page 9 catch all seven species of skates [winter, little, barndoor reduction in populations of larger, slower growing species (Dipturus laevis), winter, thorny, clearnose (Raja (Dulvy et al. 2000; Musick et al., 2000). Thus, it is eglanteria), rosette (Leucoraja garmani), smooth important to have fishery-independent data on skates (Malacoraja senta)] is otter trawling. Skates are where the individual species are reported; it is also frequently taken as bycatch during groundfish trawling necessary to work out any identification problems and scallop dredge operations and discarded recreational between winter and little skate. and foreign landings are currently insignificant, at < 1% Northeast Fisheries Science Center (2000b) also of the total fishery landings. suggests the following research needs: Skates have been reported in New England fishery " More life history studies (including age, growth, landings since the late 1800s. However, commercial maturity, and fecundity studies) are necessary. fishery landings, primarily from off Rhode Island, never " Studies of stock structure are needed to identify unit exceeded several hundred metric tons until the advent of stocks. distant-water fleets during the 1960s. Landings are not

Explore possible stock-recruit relationships by reported by species, with over 99% of the landings examination of NEFSC survey data.

reported as "unclassified skates." Skate landings reached

Investigate trophic interactions between skate species 9,500 mt in 1969, but declined quickly during the 1970s, in the complex, and between skates and other falling to 800 mt in 1981 (Figure 25). Landings have groundfish.

since increased substantially, partially in response to " Investigate the influence of annual changes in water increased demand for lobster bait, and more significantly, temperature or other environmental factors on shifts to the increased export market for skate wings. Wings are in the range and distribution of the species in the taken from winter and thorny skates, the two species skate complex, and establish the bathymetric currently used for human consumption. Bait landings are distribution of the species in the complex in the presumed to be primarily from. little skate, based on areas northwest Atlantic. fished and known species distribution patterns. Landings " Investigate historical NEFSC survey data from the for all skates increased to 12,900 mt in 1993 and then R/V Albatross III during 1948-1962 when they declined somewhat to 7,200 mt in 1995. Landings have become available, as they may provide valuable increased again since 1995, and the 1998 reported historical context for long-term trends in skate commercial landings of 17,000 mt were the highest on biomass. record (Figure 25). The biomass for the seven skate species is at a medium level of abundance. For the aggregate complex, ACKNOWLEDGMENTS the NEFSC spring survey index of biomass was relatively constant from 1968-1980, then increased significantly to The authors thank Barry Shafer, John McCarthy, peak levels in the mid- to late 1980s. The index of skate Tom Finneran, Annette Kalbach, and Meredith Lock for complex biomass then declined steadily until 1994, but producing the maps and graphics. Thanks also to Claire has recently increased again. The large increase in skate Steimle and Judy Berrien for literature reviews and biomass in the mid- to late 1980s was dominated by interlibrary loans. Frank Almeida and Kathy Sosebee of winter and little skate. The biomass of large sized skates the NEFSC Woods Hole provided much needed (> 100 cm max. length: barndoor, winter, and thorny) has information, input, and reviews. 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. REFERENCES CITED Winter skate abundance is currently about the same as in Avent, S.R., S.M. Bollens, M. Butler, E. Horgan, and R. the early 1970s, and is about 25% of the peak observed in Rountree. 2001. Planktonic hydroids on Georges the mid-1980s (Figure 25). Winter skate was, until Bank: ingestion and selection by predatory fishes. recently, considered to be overfished (Northeast Fisheries Deep-Sea Res II 48: 673-684. Science Center 2000a, b), but its-status has recently been changed so that it is no longer considered to be in an Bigelow, H.B. and W.C. Schroeder. 1953. Fishes of the Gulf of Maine. U.S. Fish Wildl. Serv., Fish. Bull. 53. overfished condition (NMFS 2002). 577 p. Bor, P.F.H. 2001 Jan. 22. Table. Egg-capsules of rays and skates. <http://www.raiidae.tmfweb.nl/>. Accessed RESEARCH NEEDS 2001 Feb. 7. Bowman, R.E., C.E. Stillwell, W.L. Michaels, and M.D. Imprecise reporting of fishery statistics where several Grosslein. 2000. Food of northwest Atlantic fishes skate species are lumped together under one category and two species of squid. NOAA Tech. Mem. NMFS-(e.g., "unclassified skates" or "skates spp.") can mask NE-155. 138 p. basic changes in community structure and profound

Page 10 Collette, B.B. and K.E. Hartel. 1988. Annotated list of fishes of Massachusetts Bay. NOAA Tech. Mem. Edwards, R.L., R. Livingstone, Jr., and P.E. Hamer. 1962. fishes - Nantucket Shoals to Cape Hatteras. Albatross III Cruise no. 126. U.S. Fish Wildl. Serv. Spec. Sci. Frisk, M.G., T.J. Miller, and M.J. Fogarty. 2001. Can. J. Fish. Aquat. Sci. 58: 969-981. Fitz, E.S., Jr. and F.C. Daiber. 1963. An introduction to Bull. Bingham Oceanogr. Collect., Yale Univ. 18 (3): 69-97. Garrison, L.P. and J.S. Link. 2000a. Dietary guild United States continental shelf ecosystem. Garrison, L.P. and J.S. Link. 2000b. Fishing effects on fish community in the Georges Bank region. ICES J. Geer, P.J. 2002. Summary of essential fish habitat 2001-03, Jan. 2001, Revised.June 2002. 169 p. Gottschall, K, M.W. Johnson, and D.G. Simpson. 2000. American lobster, and long-finned squid in Long Island Sound based on the Connecticut Fisheries Division Bottom Trawl Survey, 1984-1994. NOAA Hildebrand, S.F. and W.C. Schroeder. 1928. Fishes of Chesapeake Bay. Bull. U.S. Bur. Fish. 43(1). 366 p. Contrib. Can. Biol. 1922 (3): 49-72 Kaplan, J. 1999. Black-backed gull catches a skate. The Melvin, and D.A. Methven. 1984. Fishes, fish Mahon, R. 1997. Demersal fish assemblages from the McEachran, J.D. 1973. Biology of seven species of skates (Pisces: Rajidae). Ph.D. dissertation, Coll. William McEachran, J.D. 2002. Skates. Family Rajidae. In B.B. Collette and G. Klein-MacPhee eds. Bigelow and NMFS-F/NEC-51. 70 p. Schroeder's fishes of the Gulf of Maine. 3 rd Edition. p. 60-75. Smithsonian Institution Press, Washington, Winter water temperatures and an annotated list of DC. 748 p. ' McEachran, J.D., D.F. Boesch, and J.A. Musick. 1976. Food division within two sympatric species-pairs of Rep. Fish. 397. 31 p. skates (Pisces: Rajidae). Mar. Biol. 35: 301-317. McEachran, J.D. and K.A. *Dunn. 1998. Phylogenetic Estimation and analysis of biological parameters in analysis of skates, a morphologically conservative elasmobranch fishes: a comparative life history study. clade of elasmobranchs (Chondrichthyes: Rajidae). Copeia 1998 (2): 271-290. McEachran, J.D. and C.O. Martin. 1977. Possible the biology of Raja eglanteria Bosc 1802 and Raja occurrence of character displacement in the sympatric erinacea Mitchill 1825 as they occur in Delaware Bay. skates Raja erinacea and R. ocellata (Pisces: Rajidae). Environ. Biol. Fishes 2: 121-130. McEachran, J.D. and J.A. Musick. 1973. Characters for Garrison, L.P. 2000. Spatial and dietary overlap in the distinguishing between immature specimens of the Georges Bank groundfish community. Can. J. Fish. sibling species, Raja erinacea and Raja ocellata Aquat. Sci. 57: 1679-1691. (Pisces: Rajidae). Copeia 1973: 238-250. McEachran, J.D. and J.A. Musick. 1975. Distribution'and structure of the fish community in the Northeast relative abundance of seven species of skates (Pisces: Mar. Ecol. Rajidae) which occur between Nova Scotia and Cape Prog. Ser. 202: 231-240. Hatteras. Fish. Bull. (U.S.) 73: 110-136. Merriman, D. and H.E. Warfel. 1948. Studies on the spatial distribution and trophic guild structure of the marine resources of southern New England. VII. Analysis of a fish population. Bull. Bingham Mar. Sci. 57: 723-730. Oceanogr. Collect., Yale Univ. 11 (4): 131-164. Michalopoulos, C. 1990. A field study on the ecology and description and identification for Federally managed behavior of the winter skate (Raja ocellata) and little species inhabiting Virginia waters of Chesapeake skate (Raja erinacea) off outer Cape Cod, Bay 1988-1999. Virginia Mar. Res. Rep. VMRR Massachusetts. M.S. thesis, Southeastern Mass. Univ., North Dartmouth, MA. 71 p. Michels, S.F. and M.J. Greco. 2000. Coastal finfish The distribution and size composition of 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 Tech. Rep. NMFS 148. 195 p. Press, Washington, DC. 324 p. Nelson, J.I., Jr., S. Perry, D. Miller, and G. Lamb. 1983. Inventory of New Hampshire's marine coastal Huntsman, A.G. 1922. The fishes of the Bay of Fundy. fisheries. N.H. Fish Game Dep., Div. Inland Mar. Fish. 35 p. NMFS. 2002. Annual report to Congress on the status of Connecticut Warbler 19: 81-82. U.S. fisheries - 2001. U.S. Dep. Commerce, NOAA, Macdonald, J.S., M.J. Dadswell, R.G. Appy, G.D. NMFS, Silver Spring, MD. 142 p. Northeast Fisheries Science Center. 2000a. Report of the assemblages, and their seasonal movements in the 30th Northeast Regional Stock Assessment Workshop lower Bay of Fundy and Passamaquoddy Bay, (30th SAW): Public Review Workshop. Northeast Canada. Fish. Bull. (U.S.) 82: 121-139. Fish. Sci. Cent. Ref. Doc. 00-04. 53 p. Northeast Fisheries Science Center. 2000b. Report of the Scotian Shelf and Bay of Fundy, based on trawl 30th Northeast Regional Stock Assessment Workshop survey data (1979-1993). Can. Manuscr. Rep. Fish. (30th SAW): Stock Assessment Review Committee Aquat. Sci. 2426. 38 p. (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 and Mary, Williamsburg, VA. 127 p. 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 Predators: sharks, other skates, gray seals, and gulls. prey in terms of numbers or occurrence, followed by decapods, isopods, bivalves, Winter and little skate co-occur from Nova Scotia to Cape Hatteras. fishes. Hydroids also ingested. In terms of Although winter and little skate are sympatric species with similar weight, amphipods, decapods and fish can habitat requirements, there's not a high degree of competitive be most important; fish are especially interaction between them because they are positively correlatedby prevalent in larger skate. Polychaetes abundance. Also, winter skate feeds largely on infauna, while little include: Nephtys spp., Nereis spp., skate predominately selects epifauna. Sympatric populations of winter Lumbrinerisfragilis,Ophelia denticulata, and little skate also undergo character displacement in order to avoid maldanids (mostly Clymenella torquata), direct competition for food resources. Using 1973-1997 NEFSC data Aphrodite hastata. Amphipods: haustoriids, from Nova Scotia to Cape Hatteras and NEFSC food habits database, Leptocheirus pinguis, Monoculodes sp., both small (10-30 cm TL) and medium (31-60 cm TL) sized winter Hippomedon serratus,ampeliscids, skate belonged to "Amphipod/shrimp eaters" group, along with little Paraphoxus sp., Tmetonyx sp., Unciola skate and cusk eel; prey included amphipods, polychaetes, shrimp, irrorala,Bvblis serrata,oedicerotids. zooplankton. Largest winter skate (61 to > 80 cm TL) by themselves Decapods: Crangonseptemspinosa, Cancer in a subgroup of"Piscivores" because their diet contained a high irroratus,pagurid crabs, Dichelopandalus proportion of sand lance. Again, a trend toward increasing piscivory leptocerus, pandalid shrimp. Isopods: with size. Cirolana(= Politolana?)polita. Bivalves include Solem*ya sp. and Ensis directus. On Georges Bank, winter skate belongs to spatial assemblages and Sand lance was the most frequently eaten trophic groups that include Atlantic cod, little skate, longhorn sculpin, fish; yellowtail flounder, longhorn sculpin, yellowtail flounder, red and silver hake, haddock, spiny dogfish, hakes, other skate, herring, butterfish butterfish, fourspot flounder, windowpane, winter flounder, sea raven, occasionally eaten. Generally, in terms of thorny skate, Atlantic herring. Also on Georges Bank, winter skate numbers or occurrence, crustaceans made up falls into various dietary guilds or trophic groups, depending on the

              > 50% of the diet for skate < 61 cm TL,       study. Garrison and Link (2000b): "Bentho-pelagic" group included while fish (and often bivalves) were a major  31-60 cm TL winter skate, little skate, longhorn sculpin, Atlantic cod.

part of the diet of skate >79-91 cm TL. Diets of these species included shrimp such as pandalids and C. Overall crustaceans declined in importance septemspinosa, and benthic invertebrates including polychaetes, with increasing skate size (includes both gammarid amphipods, bivalves. "Skate" group included > 80 cm TL amphipods and often decapods) while winter skate, Oollock, windowpane. Prey was a combination of fish polychaetes increased with increasing skate and benthic prey, with a high proportion of sand lance. Garrison size until the skate were about 81 cm TL. (2000): In autumn, 61 cm to > 80 cm TL winter skate by themselves Amphipods occurred more frequently than in a subgroup of the "Piscivorous" group, feeding on sand-lance, decapods until the skates were > 71 cm TL. silver hake, and Atlantic herring, plus benthic invertebrates. 10-60 cm On Georges Bank, decline in importance of TL winter skate in "Demersal predators" group with flatfish, fish prey in spring may be related to haddocks, little skate, thorny skate. Prey included gammarid seasonal movements of prey. In terms of amphipods, polychaetes, isopods, Cancercrabs, C. septemspinosa. weight, crustaceans were dominant in the During spring, 10-60 cm TL winter skate in "Shrimp/amphipod diet of skate < 31-50 cm TL, while fish, predators" group with hakes, longhorn sculpin, Atlantic cod, fourspot mostly sand lance, were dominant in the diet flounder, little skate, thorny skate. Prey included gammarid of skate 51-110 cm TL. For skate < 31 cm amphipods, pandalids, C. septemspinosa, polychaetes, Cancercrabs. TL, amphipods dominated, especially L. Winter skate 61 cm to > 80 cm TL by themselves in a "Generalist" pinguis. For skate 31-50 cm TL, decapods subgroup, consuming bivalves, polychaetes, sand lance, herring. dominated, especially C. septemspinosa and Decline in importance of fish prey, 35% fish in autumn and 16% in C. irroratus. 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 1963-2002 spring Depth Spring: range of 1-300 m, Temperature Spring: range of 1-12'C, Salinity/DO Spring: range of 28-35 ppt, Juveniles and fall NEFSC most between about 11-70 m. majority at about 4-5°C. most between 32-33 ppt. trawl surveys from Fall: range of 1-400 m, most Fall: range of 5-21 'C, most Fall: range of 31-35 ppt, Gulf of Maine to between about 21-80 m. spread between about 7-16'C, majority between 32-33 ppt. Cape Hatteras. peaks at about 13-15'C. 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 Winter: range of about 7-18 m, Winter: range of 3-9°C, Winter: range of about 22-30 Delaware Division peaks at 14 m, 16 m, and 17 majority between 7-8°C. ppt and 34-35 ppt, most at 26 of Fish and Wildlife m. Spring: range of 4-17'C, ppt and between 28-29 ppt / bottom trawl Spring: range of 7-17 m, a few peaks scattered throughout range of 8-11 ppm, majority surveys of at 21 m, most at 8 m and 12- (e.g., 5°C, II°C, and 13'C). between 9-11 ppm. Delaware Bay 14 m. Fall: range of 8-13°C, a few Spring: range of 21-33 ppt, a (juveniles and Fall: range of 7-11 m and at at 16'C; most between 8- few at 15 ppt; peaks scattered adults combined) 13 m and between 18-19 m; 11 *C. throughout with the two most spread between 7-8 m, at highest at 28 ppt and 13 m, and at 18 m. 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 1963-2002 spring Spring: range of 1-300 m, Spring: range of 2-11 'C, most Spring: range of 30-36 ppt, Adults and fall NEFSC majority spread between 31-60 between 4-6°C, peak at 5°C. majority at 33 ppt. trawl surveys from M. Fall: range of 5-19'C, most Fall: range of 31-34 ppt, with Gulf of Maine to Fall: range of 11-300 m, most between I I-15°C, peak at 80-90% at 32 ppt. Cape Hatteras. between about 21-70 m, peaks 14'C. at 31-50 m. 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 See juveniles. See juveniles. See juveniles. Delaware Division of Fish and Wildlife bottom trawl surveys of Delaware Bay (juveniles and adults combined)

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 1981-1990 Amphipoda 37.5% Amphipoda 42.6% Polychaeta Polychaeta 13.1%f Unidentified ft.. Crustacea 1.6% Other prey phyla 17.2% Isopoda 9.4% Other prey ier crustacea 1.6% phyla 3.3% Other crustacea 16.4% Decapoda 23.4% poda a 4.6% n= 10 21-30 cm n= 50 Amphipoda 38.8% Polychaeta 9.40? Isopoda 5.4% Mollusca 4.9%

                                                                                                    'Unidentified Other r                                                                              ,    Crustacea 2.4%

phyla 9

                                                                                       ,da 17.1%    Fish 4.4%

Decapoda 22.9% n=48 31-40 cm n =462 Amphipoda 33.5% Amphipoda 33.1% Polychaeta V Unidentified Crustacea 4.2% Other Fish 3.4% Crustacea 1.1% Isopoda 6.8% Other Crustacea 2.0%

                                                                                                  \ Unidentified Decapoda 22.3%                                                  Fish 7.9% Crustacea 1.8%

n = 53 41-50 cm n = 516 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 ý Unidentified Crustacea 1.6% UU*dPoo I IV-/o 51-60 cm n = 31 n =399 rAmphipoda 20.3% Polychaeta 28.4% _ Crustacea nphipoda 13.2% Shirmp 1.0c Decapoda / 10.5% Othe ./:°Isopoda 12.0% Crustacea 1.6% Mollusca 9.3% Fish 23.9% 61-70 cm n = 78 n = 394 Crustacea Polychaeta 25.2% Polychaeta 34. Shirmp 1.9 Amphipoda 11.8% 6.7% Other prey Unidentified phyla 4.4% Crustacea 1.1 % Mollusca 9.4% Decapoda 12.7% Other 0% a Fish 16.0% Crustacea 2.0% Isopoda 7.9% 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% kmphipoda 7.8% Decapoda Unidentified Crustacea 2.4% 10.5% Other Mollusca 7.2% Crustacea 1.8% Isopoda 7.2% 81-90 cm n = 170 n = 710 Otherprey phyla 10.3% Other S Polychaeta 10.3% 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)

                                      ;~~;?

Fall/juveniles

                                 .J   /

I (<= 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 Do Percent Occurrencel bMonth by Month 04 30

                 ~0.3
                 )0                                                                                                     20[

ICV a 0.2

                ~0i                                                                                                     10o
                                                          !?.))     0.00       CO       3, tip                    .. .....
                                                                                                                                   . ....                                 005              002

_.... . 74 ._ .- Abundance SE. F- P-eierent Occurrence iby Month and Bottom Type Iby Month and Bottom Type SO~ - . -....... ..................................... . 12 . . . . ..-. . . . . . . . i C a o. 40 D40 ja

                         ,     0.0eo    0o    .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 2                , ý.o :                           0.0 .c. 0       .. -w,                       2o,6
                                                                                                                                 *.*      142 1 as                        04

i so

________ G'D0oW __ G4ooo .1 1 1.8 32,6 7.7 00LI ý 82 2 III' ..... undance Ii Percent Occurrence 1by Month and Depth Interval iby Month and Depth Intervalt 0.4 II. 4, -- 4 2D 4'o.00.10

                                                                             -  ~     -     ---0                          0-----                                                                   I-2      I      1                001      011 0.0     0.00     002 . 6.6__ *170 o                    .. 0210
                                                                                                                 .....                    0.7 .                       O. .                ...              . 1..   .
                                                                                                       .......                                            1 4        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. 74' 05"

                                                                    ....................... 4-7-r              740 00"
                                                                                                                                   .....?......

73"........... 55 I IIII_ WINTER SKATE--Juveniles (<= 84 cm) -\-Hudson River Winter (92 - 97) Hudson/Raritan Trawl Survey .

                                                                                                                          '<NNew York Fish Caught/station (excluding 0 catch):                                                                                                           -=°**40° 35" Mean: 3 Min: 1 Max: 18 Length (cm): Min: 25 Max: 84                                                                                                   S.

0. '0 C,' 0 C,,.

                                                ...---.*/             0 C                                                     .. 0' Staten           C>
                                                .1
                                                    /
                                                                                                .       0 y   *!f' Island                                                           S
                   .~
                                                                                                  '       ~.           b 4            0.                                                                                          4Q0 30" o.

0* * *~Og

                                                                                    .  '0.

Number/Tow

                                                                                                                                )0                      1-2

3 New Jersey 0 6-10 f400 25.

                                                                                                                                                        >10 74 15"               740 10'                         74005"                                    74° 000"                                     73' 55" A- Hudson River WINTER SKATE--Juveniles (<= 84 cm)

Spring (92 - 97) Hudson/Raritan Trawl Survey "" New York \\ Fish Caught/station (excluding 0 catch): /.. - /N

                                                                                                                                                              -- 40 35 Mean: 2 Min: 1 Max: 13 Length (cm): Min: 21 Max: 84                          'C          .                 ,

C,, *0~ Staten /, Island *0 40° 30' Number/Tow 1-2 0 P",'

3-5 csi New Jersey
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" 74° 05" 740 00' 730 55 WINTER SKATE--Juveniles (<= 84 cm) -- Hudson River Summer (92 - 96) Hudson/Raritan Trawl Survey New York Fish Caught/station (excluding 0 catch): -- *:/*> ..... -40*35 Mean: 2 Min: 1 Max: 3 -V 7, Length (cm): Min: 39 Max: 61 Staten Island 40° 30 '

9--.d' Number/Tow fý 1-2 1-011

3-5 -

                          .1 New Jersey

6-10 -i40 25"

                                                                                                                                   >10 740 10                  740 05'                             74000 74° 15'"                                                          A -H...............

I........ud sn.......... R

                                                                                                   .......... er
                                                                                                            ..........                730 55"
                                                                              ", --L,-Hudson River WINTER SKATE--Juveniles (<= 84 cm)

Fall (92 - 96) Hudson/Raritan Trawl Survey Ne~ N v York i- 0

                                                                                                                                              -*: 40°35 Fish Caught/station (excluding 0 catch):                                    e Mean: 3 Min: 1 Max: 14                                                                /

Length (cm): Min: 25 Max: 83

                                                                                       *              ..                 0 Staten                                                  4 Island

0 0,*

                                                                                                                      *0 0.
                                                                                             *            .           0 S..

0~.

0.

                                                   *.5                                                                                            ý40° 30'

S.,.
6 a 0 0 S

                                                                               .o I

6 Number/Tow

                                                                                                                                 *1-2 P
                                                                   ,n     ~          .

3-5 - N

                                                                                                                   'V New Jersey

6-10 .40°25

                                                                                                                                  >10 Figure 11. cont'd.

Page 31. 750 30 ' 750o10 750 00' I 740 50

                                                                .WINTER SKATE                              :390 30" All Size Classes Combined Winter (1966 - 1999)

Delaware Bay Trawl Survey Average Number of Fish Caught and Percent Occurrence

                                      '4w.
                                                                                                           .390 20 A

NJ 0.00 0% 0.07 ' 390 10

                     /'7%

0.33 20% C-s'

/<

0.23. 0.00 * / /.3 r .. 14% 0%

                                                                                      *              *" h:39° O0 0.60                              0.33 13%        0.00                               6 ape 0%       1.20        33%          May 33%

3.29. 0.00 0.69 00/0 # Times 25% 29% Station Sampled DE 1-5

                                                   "*,    0.40                           10-22            ~380 50 lO0%
                                                                                      . 31 -40 Cape/
                                                                   /                  o 41-50 Data Source:

Delaware Division of Fish &Wildilfe Henlopenr. o 51-61 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 ' WINTER SKATE

390 30" All Size Classes Combined Spring (1966 - 1999)

Delaware Bay Trawl Survey Average Number of Fish Caught and Percent Occurrence

 ~--  -y 390 20 NJ 0.00 0                 'N-0%
                                                              /"             K
                                                      " 'N_*.>*    *..*

0.00 0 0% 1390 10'

            ,---%fv*

0.15 0 7% 0.00 i/

M/* . 0.31 0 17% 00/% ICa 0.30 0.60 0.11 0 7% 0.85 60% 35% D E 4.50 0.25 0.41 100% # Times 14% 22% Station Sampled DE -- 1-5 0.45 10-22

                                                                                                      *380 50" 18%

31-40 o 41-50 Data Source: Cap(

Delaware Division of Fish &Wildlife Henlhopen;

51-61 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
             ,ýp    0,,. _

0.00 0 ,d! --39° 20" 0% ,f" 0% 0.00 0.00 0% 0% 0% +~o/

                                                                                 /            39° 000 0.0                 0.00 0%         0        0%      #Times Station Sampled 1-5 DE                       \        0.00
                                                "\* 0.00                       10-22      -3850"
                                                                            . 31-40
                                                            /              o 41-50 Data Source:                                   HenlopenH Delaware Division of Fish &Wildilfe             e                       51-61 Figure 12. cont'd.

Page 34 750 30 " 750 20 ' 750 10 ' 750 00' 740 50 WINTER SKATE All Size Classes Combined ;390 30~ Fall (1966 - 1999) Delaware Bay Trawl Survey Average Number of Fish Caught and Percent Occurrence

                                                                                                                  .390 20

(' J390 10 x 0.05 0 2% 0.jo 0.00 J 0% 0% 39° 00 0.06 0 0.20

                                            ,.{4%          0.00 0%       0.13          20%

3% 0.26 0.00 0.34 0 # Times 7%/ 0% 14% Station Sampled DE 0.06

                                                                                                 -- 1-5          =138° 50' 0

10-22

31-40 Data Source: o 41-50 Cape/

Delaware Division of Fish &Wildilfe Henlopen' e 51-61

                                                                                 .                             -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 74* 05 " _74,' 00 73' 55 WINTER SKATE--Adults (>= 85 cm) \ -~---Hudson River Spring (92 - 97) Hudson/Raritan Trawl Survey New York ', Fish Caught/station (excluding 0 catch): > 4Q0 35' Mean: 1 Min: 1 Max: 2 -J#

Length (cm): Min: 85 Max: 92

/-/

Staten W/V Island 4Q0 30" Number/Tow 4ý 1-2

3-5 New Jersey 66-10 04"25

                                                                                                                                      >10
                                                                       ,                 -j 74' 15"              740 10"                  74005"                       740 00                                        730 55" i......... ÷.........
                                                                              "'* Hud,                                                          f WINTER SKATE--Adults (>= 85 cm)                                                son River Fall (92 - 96)

Hudson/Raritan Trawl Survey NewYork Ne, - Fish Caught/station (excluding 0 catch): 400 35 Mean: 1 Min: 1 Max: 1 Length (cm): Min: 87 Max: 94

"/

I Staten Island L 52 4400 30" Number/Tow

                                                                                                       ,01 1-2
                             ~5J/'                              -5   -,,.

3-5 New Jersey -{40025"
6-10

                                                                                                                                      >10 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 [I Trawls N=10537 0 Occurrence N=2296

      -   30 a)                                                                                          o] Catch N=35461 2 20 o-   10 0

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) 0 Trawls N=12193 30 25 0 Occurrence N=2744 E 20 [3 Catch N=46565 15 a 0 o o 0 0 0 0 0 0 0 0 0 0 0 0 000 0 .. r , .I ......... ...... Bottom Depth (m) [I Trawls N=1959 50 40 0 Occurrence N=507 a 30 0l Catch N=4900 20 10 0. 26 27 28 29 30 31 32 33 34 35 36 37 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- 0 Trawls N=11844 15 N Occurrence N=1954 El Catch N=20568 2 10 0~ [IJUhiirn i,111 nn n n n ] 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Bottom Temperature (°C)

Trawls N=1 3543 20
Occurrence N=2210 15 0 Catch N=25045 0 fL I

              -    N. CO    O LA  0   N   CO     O  0     (N   O    CO Co    0   0    0   0     0 o
                   ý        'T 0 0?  ?       c0 0    T0   00    N   0    0       0    0    c)  0)    0 Boo     DIeIT9                             OA (N    C-) -

Bottom Depth (in) El Trawls N=1956 60 N Occurrence N=404

      ~401                                                                                          [0 Catch N=4255
   *_  20  -

0 27 2 27 29 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 [I Trawls N=2312 0 Occurrence N=1 310 2 .20 o Catch N=86254

       ~                  1    2     3    4    5   6     7    8     9  10    11   12  13   14   15   16 Bottom Temperature ('C) o3 Trawls N=2385 40 N Occurrence N=1342 30                                                                                    EoCatch N=90236 C:

8 20 a) 10 0 4? 0 I$) O In 0 I ) 0 In L0 0 LnO 0 IO 0 In

                         -   -t               *

I IO 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 0 Trawls N=2067 20

Occurrence N=958 c 15 EOCatch N=46556 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) [o Trawls'N=2156 50

Occurrence N=1017 40 0 Catch N=52536 30 20 a20~

1D-10 0 N NO 0o co to 0 Depo i LO (0 (o0 Bottom Depth (m) Figure 20. cont'd.

Page 46 Winter 30 C 20 1 Li Trawls N=468 0) C-, 0 Occurrence N=205 a, a- 10 - h 07Catch N=562 0 III II:l II:i

                            - .1 1.1 1.1 lit

[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 E  Trawls N=466 C

0) C., 10 I Occurrence N=205 a, E1Catch N=562 0~ 0 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 Salinity (PPT) C 20 El Trawls N=265 0, 0 E Occurrence N=125 a, 11 Catch N=292 0~ 10-0- 0 1 2 3 4 5 6 7 r,-I]L 8 9 10 11 12 I[ 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 0I-

                          -I"     t1T1-,f    -.1                                                             I Trawls N=330 I Occurrence N=1 14
                                                                                                             - Catch N=262 0      2     4     6   8   10    12    14   16    18        20       22        24    26  28 30                                       Depth (m) 20                                                                                                 I:] Trawls N=326 a)
     .-)

o Occurrence N=1 13 n 10 D]Catch N=261 0 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Temperature (0C) 30 "E 20 ED Trawls N=326 m Occurrence N=1 13 0- 10 ]I Catch N=261 0 0 rL El Ill Imlll* iI11 Im[ Illll[ An Ill I* 0 22, .4 .4 *66 88 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 12 14 Salinity (PPT) 40.- 30 - E:Trawls N=1 77 C M Occurrence N=57 F (D 20 E.- El Catch N=105 10 0 0 1 2 3 4 5 6 nU, 7

                                                         ý]ii 8     9  10        11       12       13    14 15 DO (PPM)

Figure 21. cont'd.

Page 48 Summer 50 40 LwTrawls N=356

    ,. 30 (D                                                                                o Occurrence N=4 20                                                                          EoCatch N=7 10 0                                           1'-,r 0     2      4   6   8  10   12    14   16 18      20   22  24 26  28 Depth (m) 50 40 C

Trawls N=352 30 U1K U) 0
Occurrence N=4 U) 20 L, Catch N=7 0~

10 n 0 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Temperature (0C) 80 70 60 E] Trawls N=347 C 50 C.) '40

Occurrence N=4 30 EoCatch N=7 20 10 0

H 1[1 11 r11

                                                                       ,I]1 0   2    4     6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 Salinity (PPT) 70 60 50                                                                         EoTrawls N=215 E

2 40 a Occurrence N=3 U)_ F,I 30 El Catch N=5 rPH 20 10 n 0 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 Fall

         "*t*

20- Li Trawls N=586

Occurrence N=229 a 10
Catch N=590 0 II 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Depth (m) 30 20 El Trawls N=570 (D
Occurrence N=224
Catch N=583 0

0 2 4 [I1~I]~ 6 8 fi1r FJ 10 12 14 1I 16 18 20 22 24 26 28 Temperature (°C) LU Ei Trawls N=532 C a, C.) 10 0 Occurrence N=209 a) LI Catch N=563 lil IIiIii

                                                 -   JI    I              -          -

0 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 Salinity

                                                     ý]  - CIA, (PPT) 40 a,
        .30
                                                                                              *I Trawls N=458 H 20

Occurrence N=183 LI Catch N=509 10 0

0 1 2 3 4 5 6 7 ir 8 9 10 11

                                                                              -oil 12 13   14 15 DO (PPM)

Figure 21. cont'd.

Page 50 Winter 30 20- El Trawls N= 166 0 Occurrence N=24 C.) El Catch N=1 00 10 0 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Depth (m) 50 - 40 L- Trawls N=166 30 a) 20 Ii n Occurrence N=24 11 Catch N=100 a-10 0 IL 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Temperature (°C) 50 40 C

Trawls N=157 a) 30 C.)
Occurrence N=24 a) 20
Catch N=100 10 0 0---

2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 Salinity (PPT) 50 - 40 [] Trawls N=1 25 30 20 0 Occurrence N=21 m Catch N=90 10 0 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 30 20 11 Trawls N=378 C: D 0 Occurrence N=50 1U_ X UiCatch N=103 10 0 -i-F 1'I 11LII, n -L 0 2 4 6 8 10 .12 14 16 18 20 22 24 26 28 Depth (m) 20 C

                                                                                        *iTrawls N=373 o.)  10

Occurrence N=49 (D

(L ri Catch N=102 0 0 2 [ýIII 4 6 B 3~ ýýiJrrfn 8 10 12 14 16 18 20 22 24 26 28 Temperature (°C) 10 Li Trawls N=374 a)

Occurrence N=50 I0

                                                                                        *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) rIrI 30 F 20 Li Trawls N=312 U.) 0 Occurrence N=39 a) n- Catch N=75 Cn 10 0 mu ,I EL 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 DO (PPM) Figure 22. cont'd.

Page 52 Fall. 30 C 20 3 .Trawls N=422 a, 0 U Occurrence N=1 6 a)

Catch N=39 ci. 10 0

0 2 4 6 8 1.0 12 14 16 18 20 22 24 26 28 Depth (m) 40 30 Li Trawls N=405 20- UOccurrence N=16

0. 13 Catch N=39 10 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Temperature (°C) 30

        -201I                                                                           Lj Trawls N=399 I

1I I a) ° .I I N Occurrence N=16

        **10-El Catch N=39 0   2   4    6  8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 Salinity (PPT) 70-60-50-                                                                         HI Trawls N=341 C

a, 40- II Occurrence N=14 C., a, 30 - Catch N=37 0~ 20 - 10-0 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 [I Trawls N=10537 0 Occurrence N=779 30 EoCatch N=2458 220 10 0 1 2 3 4 5 6 7 8 9 1011121314151617181920212223 Bottom Temperature (°C) [I Trawls N=12193 30 M Occurrence N=1 025 215 EoCatch N=3562

          *0 o   o   o   0    0      0     0     0    0    0  0   0 0   0    0     0     0     0 a- 1                                    (

c';(C JC? i 4 Bottom Depth (Mn) [3 Trawls N=1 959 80 0 Occurrence N=66 L)40 a) I o Catch N=96

      -201 0                      ---     ,=3--T-ý T-              ..              I0     -T-      *1ý __

26 27 28 29 30 31 32 33 34 35 36 37 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 [I Trawls N=11844 25 0 Occurrence N=754

     " 20

[3 Catch N=3142 2 15 _ 10 5 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 Bottom Temperature (°C) [I Trawls N=13543 40 30

*1

Occurrence N=932 0 Catch N=4639 2 20

a. 10 0

o 0 0 0 0 0 0 0 C0 U) D CO 1? 0? cý ( ~-0 a0 C c" 0 c>~0 0C0 0 0

                                                          -     - - -- --    (          tMC
                                                                                          )   ý Bottom Depth (in) 100                                                                                        (3 Trawls N=1956 0 Occurrence N=104 80 i]                                                                                   E3Catch N=228 a,-60 40 20 0

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 0 Trawls N=2312 20- E Occurrence N=407 15 a, - 0 Catch N=2902 S10 n-5- 1 2 3 4 5 6B 7 8 9 10 11 12 13 14 15 16 Bottom Temperature (°C) [3 Trawls N=2385 40 0 Occurrence N=409 C 30 El Catch N=2912 20 a) n 10 0 U9 0 ) 0CD 0 UD) 0 UD 0 C) 0 U) 0 C3 0 V) w--J CN C) C) I i U? U? I? I? 1 1-iZ ? 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 0 Trawls N=2067 N Occurrence N=217 0 Catch N=1520 Bottom Temperature (°C) 40 0 Trawls N=2156

Occurrence N=221 10 0 Catch N=1548 5

1011 i S20

                             ~FJlrim                 L 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     T- CO
                                                                       ý     CO    -C
                                                                                    ?

(0J R (' F (F) A :ý U0 G ) O CL 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 16000 0Skate

                         --          complex, commercial landings (mt)
                                                                               ,-,                              16
                      -       Survey index of winter skate biomass             "O 14000                                                                                                 14,,
        " 12000                                                                                              -- 12   .

10000 E w 8000 8

      "    6000                                                                                                 6 4000                                                                                     :%'4 2000                                    ""                                             -             22 0                                                                                                  0 C:    N     T (0   co    0:    N~ --t    0 CD 0      0It(00 N     C,   (0    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.}}

ML072070390

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