NOAA Technical Memorandum NMFS-NE-179

Essential Fish Habitat Source Document:

Winter Skate, Leucoraja ocellata,

Life History and Habitat Characteristics

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
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. January 2000. xiv + 138 p., 1 fig., 7 tables, 2 app. NTIS Access. No. PB2000-106735.

156. Proceedings of the Summer Flounder Aging Workshop, 1-2 February 1999, Woods Hole, Massachusetts. By George

R. Bolz, James Patrick Monaghan, Jr., Kathy L. Lang, Randall W. Gregory, and Jay M. Burnett. May 2000. v + 15 p., 5 figs.,

5 tables. NTIS Access. No. PB2000-107403.

157. Contaminant Levels in Muscle of Four Species of Recreational Fish from the New York Bight Apex. By Ashok D.

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., 1 table, glossary. NTIS Access. No. PB2001-106847.

159. An Overview and History of the Food Web Dynamics Program of the Northeast Fisheries Science Center, Woods

Hole, Massachusetts. By Jason S. Link and Frank P. Almeida. October 2000. iv + 60 p., 20 figs., 18 tables, 1 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. October 2000. 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 Stuart J. Wilk. November 2000. vii + 106 p., 24 figs., 51 tables.

NTIS Access. No. PB2002-105456.

162. U.S. Atlantic and Gulf of Mexico Marine Mammal Stock Assessments – 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. November 2000. ix + 303 p., 43 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.,

1 table. NTIS Access. No. PB2001-103542.

164. An Overview of the Social and Economic Survey Administered during Round II of the Northeast Multispecies Fishery

Disaster Assistance Program. By Julia Olson and Patricia M. Clay. December 2001. v + 69 p., 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. December 2001. viii + 129 p., 1 fig., 81 tables, 4 app. NTIS Access. No.

PB2002-108348.

166. Report on the Third Northwest Atlantic Herring Acoustic Workshop, University of Maine Darling Marine Center,

Walpole, Maine, March 13-14, 2001. By William L. Michaels, editor and coconvenor, and Philip Yund, coconvenor.

December 2001. iv + 18 p., 14 figs., 2 app. NTIS Access. No. PB2003-101556.

Recent Issues in This Series:
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

Essential Fish Habitat Source Document:

Winter Skate, Leucoraja ocellata,

Life History and Habitat Characteristics

David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano

National Marine Fisheries Serv., James J. Howard Marine Sciences Lab., 74 Magruder Rd., Highlands, NJ 07732

This series represents a secondary level of scientific publishing. All issues employ

thorough internal scientific review; some issues employ external scientific review.

Reviews are -- by design -- transparent collegial reviews, not anonymous peer

reviews. All issues may be cited in formal scientific communications.

NOAA Technical Memorandum NMFS-NE-179
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 copublished, i.e., both as paper copies and as Web postings. All Web postings

are available at: www.nefsc.noaa.gov/nefsc/habitat/efh. Also, all Web postings are in "PDF" format.

Information Updating

By federal regulation, all information 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 the use of species names in all technical communications is generally to follow

the American Fisheries Society’s lists of scientific and common names for fishes (i.e., Robins et al. 1991a

), mollusks (i.e.,

Turgeon et al. 1998b), and decapod crustaceans (i.e., Williams et al. 1989c), and to follow the Society for Marine

Mammalogy's guidance on scientific and common names for marine mammals (i.e., Rice 1998d

). 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 1998e

; McEachran and Dunn 1998f).

a

Robins, C.R. (chair); Bailey, R.M.; Bond, C.E.; 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. b

Turgeon, D.D. (chair); Quinn, J.F., Jr.; Bogan, A.E.; 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. Publ. 26; 526 p. c

Williams, A.B. (chair); Abele, L.G.; Felder, D.L.; Hobbs, H.H., Jr.; Manning, R.B.; McLaughlin, P.A.; Pérez Farfante, I. 1989. Common

and scientific names of aquatic invertebrates from the United States and Canada: decapod crustaceans. Amer. Fish. Soc. Spec. Publ.

17; 77 p. d

Rice, D.W. 1998. Marine mammals of the world: systematics and distribution. Soc. Mar. Mammal. Spec. Publ. 4; 231 p. e

Cooper, J.A.; Chapleau, F. 1998. Monophyly and interrelationships of the family Pleuronectidae (Pleuronectiformes), with a revised

classification. Fish. Bull. (U.S.) 96:686-726. f

McEachran, J.D.; Dunn, K.A. 1998. Phylogenetic analysis of skates, a morphologically conservative clade of elasmobranchs

(Chondrichthyes: Rajidae). Copeia 1998(2):271-290.
Page iii 

FOREWORD 

One of the greatest long-term threats to the viability of 

commercial and recreational fisheries is the continuing 

loss of marine, estuarine, and other aquatic habitats. 

Magnuson-Stevens Fishery Conservation and 

Management Act (October 11, 1996) 

The long-term viability of living marine resources 

depends on protection of their habitat. 

NMFS Strategic Plan for Fisheries 

Research (February 1998) 

The Magnuson-Stevens Fishery Conservation and 

Management Act (MSFCMA), which was reauthorized 

and amended by the Sustainable Fisheries Act (1996), 

requires the eight regional fishery management councils 

to describe and identify essential fish habitat (EFH) in 

their respective regions, to specify actions to conserve 

and enhance that EFH, and to minimize the adverse 

effects of fishing on EFH. Congress defined EFH as 

“those waters and substrate necessary to fish for 

spawning, breeding, feeding or growth to maturity.” The 

MSFCMA requires NMFS to assist the regional fishery 

management councils in the implementation of EFH in 

their respective fishery management plans. 

NMFS has taken a broad view of habitat as the area 

used by fish throughout their life cycle. Fish use habitat 

for spawning, feeding, nursery, migration, and shelter, but 

most habitats provide only a subset of these functions. 

Fish may change habitats with changes in life history 

stage, seasonal and geographic distributions, abundance, 

and interactions with other species. The type of habitat, 

as well as its attributes and functions, are important for 

sustaining the production of managed species. 

The Northeast Fisheries Science Center compiled the 

available information on the distribution, abundance, and 

habitat requirements for each of the species managed by 

the New England and Mid-Atlantic Fishery Management 

Councils. That information is presented in this series of 

38 EFH species reports (plus one consolidated methods 

report). The EFH species reports are a survey of the 

important literature as well as original analyses of fisheryJAMES J. HOWARD MARINE SCIENCES LABORATORY

HIGHLANDS, NEW JERSEY

SEPTEMBER 1999 

independent data sets from NMFS and several coastal 

states. The species reports are also the source for the 

current EFH designations by the New England and MidAtlantic Fishery Management Councils, and 

understandably have begun to be referred to as the “EFH 

source documents.” 

NMFS provided guidance to the regional fishery 

management councils for identifying and describing EFH 

of their managed species. Consistent with this guidance, 

the species reports present information on current and 

historic stock sizes, geographic range, and the period and 

location of major life history stages. The habitats of 

managed species are described by the physical, chemical, 

and biological components of the ecosystem where the 

species occur. Information on the habitat requirements is 

provided for each life history stage, and it includes, where 

available, habitat and environmental variables that control 

or limit distribution, abundance, growth, reproduction, 

mortality, and productivity. 

Identifying and describing EFH are the first steps in 

the process of protecting, conserving, and enhancing 

essential habitats of the managed species. Ultimately, 

NMFS, the regional fishery management councils, fishing 

participants, Federal and state agencies, and other 

organizations will have to cooperate to achieve the habitat 

goals established by the MSFCMA. 

A historical note: the EFH species reports effectively 

recommence a series of reports published by the NMFS 

Sandy Hook (New Jersey) Laboratory (now formally 

known as the James J. Howard Marine Sciences 

Laboratory) from 1977 to 1982. These reports, which 

were formally labeled as Sandy Hook Laboratory 

Technical Series Reports, but informally known as 

“Sandy Hook Bluebooks,” summarized biological and 

fisheries data for 18 economically important species. The 

fact that the bluebooks continue to be used two decades 

after their publication persuaded us to make their 

successors – the 38 EFH source documents – available to 

the public through publication in the NOAA Technical 

Memorandum NMFS-NE series. 

JEFFREY N. CROSS, (FORMER) CHIEF

ECOSYSTEMS PROCESSES DIVISION

NORTHEAST FISHERIES SCIENCE CENTER
Page iv 
Page v 

Contents 

Introduction ...................................................................................................................................................................................... 1 

Life History ...................................................................................................................................................................................... 1 

Geographical Distribution................................................................................................................................................................. 5 

Habitat Characteristics...................................................................................................................................................................... 6 

Status of the Stocks........................................................................................................................................................................... 8 

Research Needs................................................................................................................................................................................. 9 

Acknowledgments ............................................................................................................................................................................ 9 

References Cited............................................................................................................................................................................... 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 (Mitchill 1815), male......................................................................................... 16 

Figure 2. Egg case of winter 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 on Massachusetts inshore surveys............ 44 

Figure 21. Seasonal distributions of juveniles relative to temperature, depth, salinity, and DO in the Hudson-Raritan estuary ... 46 

Figure 22. Seasonal distributions of juveniles and adults relative to temperature, depth, salinity, and DO in Delaware Bay ....... 50 

Figure 23. Spring/fall distributions of adults relative to bottom temperature, depth, and salinity based on NEFSC surveys ........ 53 

Figure 24. Distributions of adults relative to bottom temperature and depth based on Massachusetts inshore surveys................. 55 

Figure 25. NEFSC spring survey index of winter skate biomass and commercial landings of the seven species skate complex .. 57 

Page 1 

INTRODUCTION 

 The winter skate [Leucoraja ocellata (Mitchill 1815); 

formerly Raja ocellata, see McEachran and Dunn (1998); 

Figure 1] occurs from the south coast of Newfoundland 

and the southern Gulf of St. Lawrence to Cape Hatteras 

(Bigelow and Schroeder 1953; McEachran and Musick 

1975; Scott and Scott 1988; McEachran 2002). Its center 

of abundance is on Georges Bank and in the northern 

section of the Mid-Atlantic Bight; in both areas it is often 

second in abundance to little skate (Leucoraja erinacea), 

a sympatric species (McEachran and Musick 1975). 

 Immature winter skate are often confused with 

immature little skate, the distinctions are size-dependent 

(McEachran and Musick 1973; McEachran 2002). 

Number of tooth rows, length at maturity, and location of 

pelvic denticles are the characters most commonly used to 

differentiate the two species (Michalopoulos 1990). 

LIFE HISTORY 

EGGS 

 The single fertilized egg is encapsulated in a leathery, 

amber to brown egg capsule which is deposited on the 

bottom (Figure 2). The capsules are rectangular in 

outline, with the upper and lower surfaces about equally 

convex and each corner of the capsule having a long 

slender horn (Vladykov 1936; Scott and Scott 1988; 

Figure 2). The anterior horns are nearly as long as the 

posterior horns and are equal in length to the capsule. The 

capsules range from 55-196 mm in length and 35-53 mm 

in width, and are smooth but marked with fine 

longitudinal striations (McEachran 2002). 

 Bigelow and Schroeder (1953) report egg deposition 

to occur during summer and fall off Nova Scotia and, 

quoting Scattergood, probably in the Gulf of Maine as 

well. They also state that egg deposition continues into 

December and January off southern New England. 

JUVENILES 

 The young are 112-127 mm TL at hatching 

(McEachran 2002) and are fully developed. 

ADULTS 

 Female winter skates with fully formed egg capsules 

are more abundant during the summer and fall but it is 

possible that reproduction takes place to some degree 

throughout the year (Vladykov 1936; Scott and Scott 

1988; McEachran 2002). 

AVERAGE SIZE, MAXIMUM SIZE, AND 

SIZE AT MATURITY

 Bigelow and Schroeder (1953) reported winter skate 

to have an average size of 76.2-86.4 cm TL. McEachran 

and Martin (1977) state that they are one of the larger 

skates in the Gulf of Maine, with a maximum known size 

of 150 cm TL [Bigelow and Schroeder (1953) give a 

maximum length of 106.7 cm TL] with larger individuals 

more common at higher latitudes. 

 The size at maturity increases with latitude 

(McEachran and Martin 1977). On Georges Bank and in 

the Gulf of Maine, individuals mature between 70-109 cm 

TL. However, in the Gulf of St. Lawrence they mature at 

a smaller size and do not reach as large a size as other 

populations (McEachran and Martin 1977). McEachran 

(1973), who studied skates from 1967-1970 that were 

collected from Nova Scotia and the Gulf of Maine to 

Cape Hatteras, found that the all specimens > 78 cm TL 

were mature except for a male 88 cm TL; the smallest 

mature winter skate was a female 72 cm TL. Bigelow and 

Schroeder (1953) reported that winter skate does not 

mature until at least 63.5-66.0 cm TL. On the eastern 

Scotian Shelf, Simon and Frank (1998) found that female 

winter skate reached 50% maturity at about 75 cm TL. 

 Based on the predictive equations from Frisk et al. 

(2001) and the Northeast Fisheries Science Center 

(NEFSC) survey maximum observed length of 113 cm 

TL, Lmat is estimated at 85 cm TL and Amat is estimated at 

7 years (Northeast Fisheries Science Center 2000b). 

FOOD HABITS 

 Generally, polychaetes and amphipods are the most 

important prey items in terms of numbers or occurrence, 

followed by decapods, isopods, bivalves, and fishes 

(McEachran 1973; McEachran et al. 1976). Hydroids are 

also ingested (Avent et al. 2001). In terms of weight, 

amphipods, decapods and fish can be most important; fish 

are especially prevalent in the larger winter skate 

(Bowman et al. 2000; see also Garrison and Link 2000a

and Tsou and Collie 2001a). Bigelow and Schroeder 

(1953) reported rock crabs and squid as favorite prey, 

other items included polychaetes, amphipods, shrimps, 

and razor clams. The fishes that were eaten included 

smaller skates, eels, alewives, blueback herring, 

menhaden, smelt, sand lance, chub mackerel, butterfish, 

cunners, sculpins, silver hake, and tomcod. 

 McEachran (1973) studied skates collected from 

Nova Scotia to Cape Hatteras during 1967-1970; the 

following diet descriptions are from him and McEachran 

et al. (1976). 

Nephtys spp., Nereis spp., Lumbrineris fragilis, 

Ophelia denticulata, and maldanids (mostly Clymenella

torquata) were the most abundant polychaetes in the MidAtlantic Bight and Georges Bank stomachs. Nephtys spp., 
Page 2 

Pectinaria sp., O. denticulata, and Aphrodite hastata

were the most frequently consumed prey in the Gulf of 

Maine and on the Nova Scotian shelf. 

 Haustoriids, Leptocheirus pinguis, Monoculodes sp., 

Hippomedon serratus, ampeliscids, Paraphoxus sp., and 

Tmetonyx sp. were the most frequently eaten amphipods 

over the survey area. Crangon septemspinosa was the 

most abundant decapod in the diet. Cancer irroratus, 

Dichelopandalus leptocerus, Pagurus acadianus, and 

Hyas sp. were consistently eaten but in small numbers. 

Among the minor prey items included Cirolana (= 

Politolana?) polita, which was the dominant isopod. 

Other isopods eaten included Chiridotea tuftsi and Edotea

triloba, but they contributed little to the overall diet. The 

only identifiable bivalves eaten were Solemya sp. and 

Ensis directus. The most frequently eaten fish was sand 

lance, while yellowtail flounder and longhorn sculpin 

were occasionally eaten. 

 In Smith’s (1950) study in Block Island Sound, 

nekton were more important and isopods much less 

important than in the McEachran (1973) and McEachran 

et al. (1976) studies. The major prey items for winter 

skate in Block Island Sound included nekton, L. pinguis, 

Nephtys incisa, E. directus, C. septemspinosa, Nereis sp., 

C. irroratus, Lumbrineris sp. and Monoculodes edwardsi. 

 Winter skate from Georges Bank had the most 

diverse diet and those from the Mid-Atlantic Bight the 

least diverse diet (McEachran 1973; McEachran et al. 

1976). There was no significant change in the diet with 

increase in skate size; however, the numbers of 

polychaetes gradually increased and amphipods gradually 

decreased with increasing skate size. The number of fish 

and bivalves also increased with predator size and the two 

taxa were a major part of the diet of skate > 79 cm TL. 

The ingestion of decapods was independent of skate size. 

There was also no indication of either diel or seasonal 

periodicity in intensity of feeding. In Passamaquoddy Bay 

there were no differences between the diets of small and 

large winter skate (Tyler 1972). 

 The 1973-1990 NEFSC food habits database for 

winter skate [Figure 3; see Reid et al. (1999) for details] 

generally confirms the McEachran (1973) and McEachran 

et al. (1976) studies. Crustaceans made up > 50% of the 

diet for skate < 61 cm TL, while fish dominated the diet 

of skate > 91 cm TL. Overall crustaceans declined in 

importance with increasing skate size (includes both 

amphipods and decapods) while the percent occurrence of 

polychaetes increased with increasing skate size until the 

skate were about 81 cm TL. Amphipods occurred more 

frequently than decapods until the skates were > 71 cm 

TL. Among the most frequently occurring prey species 

for almost all sizes of skate included the decapods C. 

septemspinosa and Cancer and pagurid crabs, the isopod

Cirolana (= Politolana?) polita, and sand lance. The 

following is a detailed description of the diet from the 

NEFSC food habits database broken down by winter 

skate size class (Figure 3). 

 For winter skate 21-30 cm TL, 74-84% of the diet 

consisted of crustaceans, with 38-43% of the diet 

consisting of identifiable amphipods. The most abundant 

amphipod species included Unciola irrorata, Byblis

serrata, and H. serratus. Identifiable decapods made up 

23-25% of the diet, most of which were species such as 

C. septemspinosa and C. irroratus. Identifiable 

polychaete species (9-13% of the diet) included 

Ampharete arctica. Identifiable isopod species (9% of the 

diet) included Cirolana (= Politolana?) polita. 

Nematodes, bivalves, and fish were included in the “other 

prey phyla” category (3-17% of the diet). 

 For skate 31-40 cm TL, 72-76% of the diet consisted 

of crustaceans, with 37-39% of the diet consisting of 

identifiable amphipods. Major amphipod species included 

B. serrata, U. irrorata, H. serratus, and several 

unidentified haustoriids. Identifiable decapods made up 

17-23% of the diet, most of which were C. septemspinosa

and C. irroratus. Identifiable polychaetes (12-17% of the 

diet) included Scalibregma inflatum, L. fragilis, and 

unidentified maldanids. Identifiable isopods (5-8% of the 

diet) included Cirolana (= Politolana?) polita. 

Miscellaneous items (6-9% of the diet) included 

nematodes and bivalves. Among the identifiable fish 

present in the diet (3-4%) were sand lance, yellowtail 

flounder, and hakes. 

 The percentage of crustaceans in the diet of winter 

skate 41-50 cm TL dropped to 62-69%, although 

identifiable amphipods still made up the major portion 

(33-35%) followed by decapods (14-22%). Identifiable 

polychaetes made up 19-23% of the diet; other prey 

species (including mollusca), 6-9% of the diet; 

identifiable isopods, 7% of the diet; and identifiable fish, 

3-8% of the diet. All the major prey species (except for 

the lack of the polychaete S. inflatum) were similar to the 

31-40 cm TL size class, with the additions of several 

more Unciola species, L. pinguis (an amphipod), 

unidentified pagurid crabs, and nephtyid polychaetes. 

 The percent occurrence of crustaceans in the diet of 

winter skate 51-60 cm TL dropped further, down to 53-

54%, with identifiable amphipods making up only 26-

32% of the overall diet. Some of the dominant identifiable 

amphipods included Psammonyx nobilis, unidentified 

oedicerotids, H. serratus, and unidentified haustoriids. 

Identifiable decapods made up only 9-12% of the diet; C. 

septemspinosa was again the dominant decapod prey, 

followed by C. irroratus and pagurid crabs. Cirolana (= 

Politolana?) polita was again one of the major 

identifiable isopods, which all together made up 7-12% of 

the diet. The percent occurrence of identifiable 

polychaetes continued to increase in the diet, up to 26-

29%; several of the more numerous species present were 

in the genera Nephtys and Nereis. Identifiable fish also 

increased in the diet, up to 6-13%, with sand lance the 

dominant species. Other prey phyla, including bivalves 

and nematodes, accounted for 9-11% of the diet. 

 The percent occurrence of crustaceans in the diet 

continued to decline for winter skate 61-70 cm TL: down 

to 38-44%, with identifiable amphipods making up only 
Page 3 

13-20% of the diet, while identifiable decapods made up 

11-12%. Major amphipod species included M. edwardsi, 

U. irrorata, H. serratus, and unidentified haustoriids and 

oedicerotids. C. septemspinosa continued to be the 

dominant decapod prey, followed by Cancer and pagurid 

crabs. Identifiable isopods again made up 7-12% of the 

diet; Cirolana (= Politolana?) polita continued to be one 

of the major prey species. The percent occurrence of 

identifiable polychaetes in the diet increased, up to 28-

32%; species in the genera Nephtys and Nereis were again 

dominant. The percent occurrence of identifiable fish in 

the diet continued to increase also, up to 11-24%, most of 

which were sand lance. Nine percent of the diet consisted 

of identifiable mollusks, with bivalves being dominant. 

 While the percent occurrence of crustaceans dropped 

to 29-36% for winter skate 71-80 cm TL, the percent 

occurrence of identifiable decapods was greater than the 

percent occurrence of amphipods: 11-13% versus 7-12%. 

The former were dominated by C. septemspinosa, Cancer

and pagurid crabs, and D. leptocerus, while several 

haustoriid species and U. irrorata were some of the major 

amphipod prey. Identifiable isopods made up 8-9% of the 

diet, the dominant species continued to be Cirolana (= 

Politolana?) polita. Identifiable polychaetes (25-35% of 

the diet) included L. fragilis and several Nephtys and 

Nereis species. The percent occurrence of identifiable fish 

in the diet varied widely between the two sampling 

periods, from 16-36%, although sand lance was still the 

dominant species. Identifiable mollusks made up 9-10% 

of the diet, most of which were bivalves. 

 Fish as prey items became increasingly important for 

winter skate 81-90 cm TL. They made up 29-42% of the 

overall diet. As usual sand lance were the dominant fish 

prey, other species ingested included other skate, 

longhorn sculpin, and silver hake. Crustaceans in the diet 

declined to 19-30%. The major identifiable decapod 

species (8-11% of the diet) continued to be C. 

septemspinosa and Cancer and pagurid crabs as well as 

pandalid shrimp and Ovalipes ocellatus. The major 

identifiable amphipod species (3-8% of the total diet) 

were several haustoriid species. Cirolana (= Politolana?) 

polita was once again the dominant identifiable isopod 

(all isopods together made up 5-7% of the diet). Several 

Nephtys species were the major identifiable polychaetes 

ingested, all polychaetes together made up 22-28% of the 

diet. Bivalves, particularly of the familiy Solenidae, were 

the dominant identifiable molluscan prey ingested, with 

all mollusks together accounting for 7-17% of the diet. 

 Identifiable fish made up >50% of the diet of winter 

skate 91-100 cm TL. Sand lance was the overwhelming 

dominant, some of the minor fish prey included silver 

hake, herring, and butterfish. Crustaceans were down to 

12-23% of the diet. Identifiable decapods made up 5-10% 

of the diet, C. septemspinosa, Cancer and pagurid crabs, 

D. leptocerus, and pandalid shrimp were some of the 

major decapods ingested. Identifiable amphipods made up 

only 4-5% of the total diet, with few conspicuous species. 

Identifiable polychaetes accounted for 10-13% of the diet, 

with the genus Nephtys the most notable. “Other prey 

phyla” and identifiable mollusks together accounted for 

10-12% of the diet, bivalves and nematodes dominated 

this category. 

 Finally, identifiable fish made up > 60% of the diet 

of 101-110 cm TL winter skate from the 1981-1990 

NEFSC trawl surveys. Most were sand lance. Mollusks 

were 14% of the diet, polychaetes were 13% of the diet, 

and crustaceans were down to 11% of the diet. 

 Using NEFSC data from 1977-1980, Bowman et al. 

(2000) found that in terms of percent weight, crustaceans 

were dominant in the diet of skate < 31-50 cm TL, while 

fish, mostly sand lance, were dominant in the diet of skate 

51-110 cm TL. For skate < 31 cm TL, amphipods 

dominated, especially L. pinguis. For skate 31-50 cm TL, 

decapods dominated, especially C. septemspinosa and C. 

irroratus. On Georges Bank Tsou and Collie (2001a), 

using NEFSC dietary data from 1989-1990, also showed 

that fish, especially sand lance, were most important for 

winter skate > 50 cm TL. Other noted fish prey included 

sliver hake, mackerel, and herring (see also Tsou and 

Collie 2001b). 

 Nelson (1993) calculated the predation impact of 

winter skate on their Georges Bank prey. Annual 

estimates of consumption for winter skate increased as 

they grew larger. Consumption ranged from 1.186 

kg/fish/year for skate 40-49 cm TL to 5.528 kg/fish/year 

for skate 90-99 cm TL. The percentage of benthic 

production consumed by winter skate from 1969-1990 

ranged from 11-43%. Nelson (1993) suggests that in 

relation to the total macrofauna production on Georges 

Bank, winter skate (along with little skate) consume < 

0.02% of the total. These results indicate that only a small 

to moderate proportion of benthic biomass vulnerable to 

skate predation is consumed by both winter and little 

skate, and their consumptive impact will be dependent on 

the levels of invertebrate biomass and/or production. 

PREDATORS AND SPECIES 

ASSOCIATIONS 

 Winter skate is preyed upon by sharks, other skates, 

gray seals, and gulls (Scott and Scott 1988; Kaplan 1999). 

 McEachran and Musick (1975) state that winter and 

little skate co-occurred significantly during 1967-1970 

surveys from Nova Scotia to Cape Hatteras. Although 

winter and little skate are sympatric species with similar 

habitat requirements (except perhaps temperature 

preference), there does not appear to be a high degree of 

competitive interaction between them because they are 

positively correlated by abundance and where the two 

species are most abundant (Georges Bank) they have the 

most similar diets and highest diversity of assemblages of 

prey species (McEachran 1973; McEachran and Musick 

1975; McEachran et al. 1976). 

 Also, even though the two species do consume the 

same large taxonomic groups of benthic fauna 
Page 4 

(amphipods, decapods, and polychaetes), winter skate 

predominately feeds on infaunal organisms while little 

skate feeds largely on epifauna (McEachran 1973; 

McEachran et al. 1976). McEachran (1973) and 

McEachran et al. (1976) show that large burrowing 

polychaetes and bivalves were consumed more frequently 

by winter skate and epifaunal decapods were eaten more 

frequently by little skate. Winter skate ate more 

burrowing amphipods, especially haustoriids and 

Trichophoxus epistomus while little skate consumed more 

surface dwelling amphipods such as Unciola sp., Dulichia

(= Dyopedos) monacantha, ampeliscids and caprellids. 

The division of food resources between the skates is not 

complete because some winter skate ate large numbers of 

epifauna and some little skate consumed large numbers of 

infauna. Both species ate considerable numbers of L. 

pinguis and C. septemspinosa. Little skate occasionally 

fed on haustoriids, and deep burrowing polychaetes were 

regular prey items. The infaunal and epifaunal 

preferences of the two skates may be more distinct in 

areas where they may coexist than in areas where they 

occur separately because in Delaware Bay (Fitz and 

Daiber 1963) little skate consumed relatively more 

infauna than it did in the areas sampled in Smith’s (1950) 

study or the McEachran (1973) and McEachran et al. 

(1976) studies. Winter skate does not regularly occur in 

Delaware Bay (Fitz and Daiber 1963; see also Delaware 

Bay trawl surveys, below). 

 In addition, differences in the shape and size of the 

mouth and the number of tooth rows between the two 

species were used as evidence by McEachran and Martin 

(1977) to suggest that the sympatric populations of winter 

and little skate underwent character displacement in order 

to avoid direct competition for food resources. In 

sympatric populations, winter skate has greater number of 

tooth rows in the upper jaw and a wider and less arched 

mouth, thus allowing them to feed more efficiently and 

deeper in the bottom than little skate. Little skate has a 

relatively smaller and more arched mouth with fewer 

tooth rows in the upper jaw. 

 Using 1973-1997 NEFSC data from Nova Scotia to 

Cape Hatteras, as well as the same NEFSC food habits 

database discussed above, Garrison and Link (2000a) 

investigated the dietary guild structure of the fish 

community. Both small (10-30 cm TL) and medium (31-

60 cm TL) sized winter skate belonged to the 

“Amphipod/shrimp eaters” group, along with little skate 

and cusk eel; prey included amphipods, polychaetes, 

shrimp, and zooplankton. The largest winter skate (61 to 

> 80 cm TL) were by themselves in a subgroup of 

“Piscivores” because, as described previously under the 

detailed description of the diet using the 1973-1990 

NEFSC food habits database, their diet contained a high 

proportion of sand lance. This again shows that there is a 

trend toward increasing piscivory with size. 

 The resilience of demersal fish assemblages on 

Georges Bank was investigated by Overholtz and Tyler 

(1985) using seasonal NEFSC trawl survey data from 

1963-1978. Of the five assemblage species groups or 

associations present on Georges Bank in spring and fall 

throughout the survey period, winter skate belonged to 

the “Intermediate” and “Shallow” assemblage groups. In 

the Shallow assemblage the other major species present 

besides winter skate included Atlantic cod, little skate, 

longhorn sculpin, yellowtail flounder, and haddock; in the 

Intermediate assemblage, little skate, red and silver hake, 

Atlantic cod, and haddock were some of the other major 

species present. Overholtz and Tyler (1985) considered 

winter skate to be a “resident” species, since they were 

only present in two out of the five assemblages in 

abundance. The Shallow assemblage covered most of 

Georges Bank in the spring and was slightly smaller in 

the fall. The Intermediate assemblage occurred mostly 

south of the Shallow assemblage and inside the southern 

edge of Georges Bank; it was somewhat larger in the fall, 

suggesting a migration of the species in this area to 

shallower water as the year progressed. The assemblages 

in the spring appeared to follow depth contours. 

 Garrison (2000) and Garrison and Link (2000b) have 

also investigated spatial assemblages and trophic groups 

from the Georges Bank region. Using 1963-1997 NEFSC 

trawl survey data from Georges Bank, as well as the same 

NEFSC food habits database discussed above (Garrison 

and Link [2000b] used 1973-1997 data while Garrison 

[2000] used 1991-1997 data), they found that the major 

predator groups were consistent across decades, with the 

boundaries of the assemblages similar to Overholtz and 

Tyler (1985). Garrison (2000) investigated the spatial 

assemblages during spring and autumn. He found that 

during autumn, winter skate was in the assemblage found 

in the deep habitats on southern Georges Bank that also 

included spiny dogfish, butterfish, little skate, red hake, 

fourspot flounder, and yellowtail flounder. The main 

shallow portion of Georges Bank assemblage included 31 

cm to > 80 cm TL winter skate, little skate, spiny dogfish, 

Atlantic cod, windowpane, winter flounder, and sea 

raven. In spring, the main shallow portion of Georges 

Bank assemblage included 31 cm to > 80 cm TL winter 

skate, haddock, Atlantic cod, and spiny dogfish. In terms 

of dietary guilds or trophic groups, the two studies had 

slightly different viewpoints, but the diets of winter skate 

in both studies were the same as discussed in the Food 

Habits section above. In the Garrison and Link (2000b) 

study, winter skate fell into two predator or feeding 

groups. The first was a “Bentho-pelagic” group, which 

included 31-60 cm TL winter skate, little skate, longhorn 

sculpin, and Atlantic cod. The diets of these species were 

the same as that discussed above for winter skate alone: 

shrimp such as pandalids and C. septemspinosa, and 

benthic invertebrates including polychaetes, gammarid 

amphipods, and bivalves. The second group was the 

“Skate” group, consisting of > 80 cm TL winter skate, 

pollock, and windowpane. Their prey was characterized 

by a combination of fish and benthic prey, with a high 

proportion of sand lance during the 1980s. Garrison 

(2000) had slightly different trophic groups. In autumn, 
Page 5 

the largest (61 cm to > 80 cm TL) winter skate were by 

themselves in a subgroup of the “Piscivorous” group, 

feeding on sand lance, silver hake, and Atlantic herring, 

as well as benthic invertebrates. Small and medium (10-

60 cm TL) winter skate were also in the “Demersal 

predators” group, along with flatfish, haddocks, little 

skate, and thorny skate (Amblyraja radiata). Prey 

included gammarid amphipods, polychaetes, isopods, and 

Cancer crabs, as well as C. septemspinosa. During spring, 

10-60 cm TL winter skate were in the “Shrimp/amphipod 

predators” group, along with hakes, longhorn sculpin, 

Atlantic cod, fourspot flounder, little skate, and thorny 

skate. Prey included gammarid amphipods, pandalids and 

C. septemspinosa, polychaetes, and Cancer crabs. Winter 

skate 61 cm to > 80 cm TL were again by themselves in a 

subgroup called the “Generalist” group, consuming 

bivalves, polychaetes, sand lance, and herring. The 

decline in importance of fish prey, 35% fish in the 

autumn and 16% in the spring, is probably related to 

seasonal movements of prey (Garrison 2000). 

 On the Scotian Shelf, Scott (1989), using research 

trawl survey data from roughly 1970-1984 determined 

that winter skate was locally abundant but did not 

associate closely with any other species. However Mahon 

(1997), analyzing trawl survey data for the same region 

from 1970-1993, showed that winter skate was associated 

with such species as longhorn sculpin, sea raven and 

winter flounder in the shallow waters of the Bay of Fundy 

and Sable Island. 

GEOGRAPHICAL DISTRIBUTION 

 In Canada, winter skate are found in the Gulf of St. 

Lawrence, off northeastern Nova Scotia, and the offshore 

banks of Banquereau, Sable Island, and Western Bank 

[Bigelow and Schroeder 1953; McEachran and Musick 

1975; see also Strong and Hanke (1995) for the 1970-

1993 distribution of winter skate in the Scotia-Fundy 

region; further information on winter skate distribution on 

the Scotian Shelf can be found in Simon and Frank (1996, 

1998)]. The population in the southern Gulf of St. 

Lawrence may be isolated from populations along the rest 

of the east coast (McEachran and Musick 1975). 

McEachran and Musick (1975) suggest that reports of 

little skate from the Gulf of St. Lawrence and most 

records of it from northern Nova Scotia probably refer to 

winter skate. 

 Winter skate is considered common all around the 

Gulf of Maine from Nova Scotia to Cape Cod, except for 

the deep troughs, and is also common on Georges Bank 

(Bigelow and Schroeder 1953; McEachran 2002). It has 

frequently been reported from the Bay of Fundy, and the 

coasts of Maine and Massachusetts (Bigelow and 

Schroeder 1953) and in Massachusetts Bay (Collette and 

Hartel 1988), as well as along the New Hampshire coast 

(Nelson et al. 1983). However, McEachran (2002) states 

that because of its close resemblance to the little skate, 

many of these records, as well as those for little skate 

from the same localities, are suspect. 

 Their range extends from southern New England and 

down the Mid-Atlantic Bight to northern North Carolina 

(Bigelow and Schroeder 1953; Figures 4-7, 13-16). 

 Previous authors have suggested that winter skate 

undertakes seasonal movements, especially in the 

southern part of its range, moving shoreward in autumn 

and offshore in summer (McEachran 1973; McEachran 

and Musick 1975; McEachran 2002), although this is not 

quite evident from the overall NEFSC bottom trawl 

surveys (Figures 4-7, 13-16; see descriptions below). In 

Passamaquoddy Bay, Tyler (1971) reported winter skate 

present from December to March while Huntsman (1922) 

stated it was abundant from May to November. 

McEachran (1973) suggests this disparity may be due to a 

difference in the areas the two authors sampled. Tyler 

(1971) sampled the deeper waters of Passamaquoddy Bay 

while Huntsman (1922) did not specify the sampling 

depths. However Macdonald et al. (1984) determined 

winter skate to be both a regular and occasional resident 

in Passamaquoddy Bay, and juveniles were often found at 

beach sites during summer. Merriman and Warfel (1948) 

stated it is a permanent resident off southern New 

England although there are seasonal fluctuations in 

abundance; Bigelow and Schroeder (1953) mention that 

along the southern coast of New England it comes inshore 

near Woods Hole during the colder months. However 

during August of 1988 Michalopoulos (1990) found that 

winter skate dominated the inshore skate community off 

outer Cape Cod (little skate was the only other skate other 

skate present, but was much less abundant). Schaefer 

(1967) found winter skate in the surf zone of Long Island 

during May, June, October, and November, while 

Gottschall et al. (2000), based on surveys from 1984-

1994 (see Habitat Characteristics section, below), found 

their lowest abundances in Long Island Sound were in the 

months of July, August, and September. McEachran and 

Musick (1975) reported winter skate to be more abundant 

south of Delaware Bay during the winter, it has also been 

reported from Chesapeake Bay from December to April 

(Hildebrand and Schroeder 1928; Geer 2002). 

JUVENILES 

 NEFSC bottom trawl surveys [see Reid et al. (1999) 

for details] captured juvenile (< 84 cm TL) winter skate 

year-round. (Note that winter and summer distributions 

are presented as presence/absence data, precluding a 

discussion of abundances.) In winter, juveniles were 

found from Georges Bank to Cape Hatteras, out to the 

200 m depth contour (Figure 4). Concentrations were 

found off Long Island and southern New England; they 

were almost entirely absent from the Gulf of Maine. In 

spring they were also found from Georges Bank to Cape 

Hatteras, and were concentrated nearshore throughout the 

Mid-Atlantic Bight and southern New England as well as 
Page 6 

in Cape Cod and Massachusetts Bays (Figure 5). Small 

numbers were also found along the coast of Maine and 

southwest Nova Scotia and near Browns Bank. 

Comparatively few were present in summer, with 

concentrations on Georges Bank and around Cape Cod 

(Figure 6). Some were also found near Penobscot Bay, 

Maine. Winter skate abundances in the fall were not as 

high as in the spring (Figure 7). In the fall they were 

collected from Georges Bank to the Delmarva Peninsula 

and were again concentrated along Long Island, southern 

New England, around Cape Cod, and on Georges Bank. 

Small numbers were again found along the coast of 

Maine and near Browns Bank. 

 Both the spring and fall 1978-2002 Massachusetts 

inshore trawl surveys [see Reid et al. (1999) for details] 

show similar abundances and distributions of juveniles 

(Figure 8). The highest concentrations were found on the 

Atlantic side of Cape Cod and south and west of Martha’s 

Vineyard (especially in spring) and south and northeast of 

Nantucket (also in spring). Large numbers were also 

found near Monomy Point in the fall. Other notable 

occurrences of winter skate were around Plum Island, 

Ipswich Bay, north of Cape Ann, near Nahant Bay 

(especially in the fall), in Cape Cod Bay, and in 

Nantucket Sound. 

The distributions and abundances of both juveniles 

and adults in Long Island Sound (Figures 9-10) as 

described by Gottschall et al. (2000) will be discussed in 

the Habitat Characteristics section. 

 Occurrence of juveniles in the Hudson-Raritan 

estuary appears to have the same seasonal pattern noted 

by previous authors for other estuaries; i.e., they’re 

generally absent from the estuary during the summer 

months. Juveniles were fairly well distributed throughout 

the Hudson-Raritan estuary in winter, spring and fall, and 

were most abundant in the winter and fall (Figure 11). In 

summer the few that were left were mostly confined to 

the deeper and warmer waters of the Ambrose Channel. 

The 1966-1999 Delaware Bay trawl surveys (adults 

and juveniles combined; Figure 12) also confirm the 

seasonal trends noted previously for winter skate, 

although they were not very abundant in the Bay overall. 

They were almost completely absent in summer, and a 

few were caught in the fall, while the greatest numbers 

were found in the winter and spring. The skate were most 

abundant in the center of lower Delaware Bay, near the 

mouth (Figure 12). 

ADULTS 

 NEFSC bottom trawl surveys [see Reid et al. (1999) 

for details] captured adult winter skate (> 85 cm TL) 

during all seasons (Figures 13-16). The numbers of adults 

in spring and fall were much lower than for juveniles of 

the same two seasons (winter and summer distributions 

are presented as presence/absence data, precluding a 

discussion of abundances). In winter, they were scattered 

from Georges Bank to North Carolina; very few occurred 

in the Gulf of Maine (Figure 13). In the spring, they were 

also found from Georges Bank to North Carolina but, as 

with the juveniles, were also distributed nearshore 

throughout the Mid-Atlantic Bight and along Long Island 

as well as around Cape Cod and Massachusetts Bays 

(Figure 14). Small numbers were also found along the 

coast of Maine and southwest Nova Scotia near Browns 

Bank. Few occurred in summer, being found mostly on 

Georges Bank, Nantucket Shoals, and near Cape Cod 

(Figure 15). In the fall, they were mostly confined to 

Georges Bank, near Nantucket shoals, and near Cape 

Cod, with very few found south of those areas (Figure 

16). 

 Adult little skate were collected in much fewer 

numbers than juveniles during the spring and fall 

Massachusetts inshore trawl surveys (Figure 17). The 

greatest numbers were found on the Atlantic side of Cape 

Cod and, in spring, south of Nantucket. 

 Very few adults were caught in the Hudson-Raritan 

estuary during spring and fall (Figure 18). Those few that 

were caught were concentrated around the Ambrose and 

Chapel Hill Channels. 

 The seasonal distribution and abundance of both 

adults and juveniles in Delaware Bay were discussed 

previously (Figure 12). 

HABITAT CHARACTERISTICS 

 Information on the habitat requirements and 

preferences of winter skate (based on both the pertinent 

literature and the most recent NEFSC and state surveys) 

are presented here and summarized in Tables 1 and 2. 

 Winter skate generally ranges from the shoreline to 

371 m, although it is most abundant at depths < 111 m 

(Bigelow and Schroeder 1953; McEachran 1973; 

McEachran and Musick 1975; McEachran 2002). 

Bigelow and Schroeder (1953) suggest that few are 

caught > 91 m. In the Gulf of Maine, they are most 

abundant 46-64 m but occasionally occur < 1 m as well as 

down to 285 m; they’re considered rare at depths < 2-7 m 

(Bigelow and Schroeder 1953; McEachran 2002). On the 

Scotian Shelf the winter skate is most frequently caught 

between 37-90 m (Scott 1982a). That segment of the 

population residing at depths > 10 m appear to be resident 

year-round, even though the coastal edge of the 

population appears to move shoreward in autumn and 

offshore in summer (Bigelow and Schroeder 1953). 

Merriman and Warfel (1948) stated that winter skate is a 

permanent resident off southern New England between 

15-46 m although there are seasonal fluctuations in 

abundance. The 1978-2002 spring and autumn 

Massachusetts inshore trawl surveys (see below) show 

that both juveniles and adults were found between 1-75 

m, with most found between 6-25 m (Figures 20 and 24).

Edwards et al. (1962) captured it off the Mid-Atlantic 

states during the winter at depths from 33-113 m. The 
Page 7 

1963-2002 spring and fall NEFSC trawl surveys from the 

Gulf of Maine to Cape Hatteras (see below) indicated that 

most juveniles occurred at depths < 70-80 m, although a 

few occurred as deep as 400 m (Figure 19), while most 

adults were found < 70 m and a few were as deep as 300 

m (Figure 23). In the Hudson-Raritan estuary (see below; 

Figure 21), juvenile winter skate are found from about 4-

22 m, but occur mostly around 5-8 m during a good part 

of the year. In Delaware Bay (see below; Figure 22) 

juveniles and adults were found over a range of 

approximately 7-21 m during winter, spring, and fall. 

 Winter skate has been recorded over a temperature 

range of -1.2EC to19EC (Bigelow and Schroeder 1953; 

Tyler 1971; McEachran 1973; McEachran 2002). On the 

Scotian Shelf it has been frequently found at temperatures 

of 5-9EC (Scott 1982a; Scott and Scott 1988). Bigelow 

and Schroeder (1953) reported them in the Gulf of Maine 

at around 20EC along the Massachusetts coast in the 

summer, down to 1-2EC in the coastal belt in winter, and 

near 0EC in the Bay of Fundy region in some years. 

McEachran and Musick (1975) reported their temperature 

ranges at depth of capture were -1.2EC to 4.8EC in the 

Gulf of St. Lawrence, 1.1-12.7EC off northeastern Nova 

Scotia, and 2-15EC from southern Nova Scotia to Cape 

Hatteras. Edwards et al. (1962) captured it off the MidAtlantic states during the winter at temperatures from 10-

12EC. The 1963-2002 spring and fall NEFSC trawl 

surveys from the Gulf of Maine to Cape Hatteras (see 

below) collected juvenile winter skate over a 

temperatures range of 1-21EC, with most found between 

4-5EC in the spring and about 7-16EC in the fall (Figure 

19). Adults were found over a temperature range of 2-

19EC with most found around 5EC in spring and between 

about 11-15EC in the fall (Figure 23). 

 Bigelow and Schroeder (1953) stated that this species 

is confined to sandy and gravelly bottoms but Tyler 

(1971) reported it from mud bottoms in Passamaquoddy 

Bay. In Long Island Sound during the spring, winter skate 

were most abundant on sand bottoms in the Mattituck Sill 

and Eastern Basin (Gottschall et al. 2000). On the Scotian 

Shelf, Scott (1982b) reports that the distribution of winter 

skate was confined to sand and gravel bottoms and Scott 

(1982b) suggests that bottom type, rather than depth, 

appears more important in determining the distributions 

of winter skate. 

 Winter skates are known to remain buried in 

depressions during the day and are more active at night 

(Michalopoulos 1990). This is probably not due to diel 

foraging, since McEachran et al. (1976) observed no diel 

periodicity in feeding intensity by winter skate and 

suggested that they may feed at any time during a 24 hour 

period. 

Scott (1982a) mentions that on the Scotian Shelf 

during the summers of 1970-1979, winter skate was 

found at preferred salinities of 32-34 ppt. 

JUVENILES 

 The spring and fall distributions of juvenile winter 

skate relative to bottom water temperature, depth, and 

salinity based on 1963-2002 NEFSC bottom trawl 

surveys from the Gulf of Maine to Cape Hatteras are 

shown in Figure 19. In spring, they were found in waters 

between 1-12EC, with the majority at about 4-5EC. Their 

depth range during that season was between 1-300 m, 

with most between about 11-70 m. They were found at 

salinities between 28-35 ppt, with most found between 

32-33 ppt. During the fall, juvenile winter skate were 

caught over a temperature range of 5-21EC, with most 

spread between about 7-16EC, and peaks at about 13-

15EC. They were found over a depth range of 1-400 m, 

although most were caught at depths between about 21-80 

m. They were found at salinities between 31-35 ppt, with 

the majority found between 32-33 ppt. 

 The spring and autumn distributions of juveniles in 

Massachusetts coastal waters relative to bottom water 

temperature and depth based on 1978-2002 

Massachusetts inshore trawl surveys are shown in Figure 

20. In the spring they were found in waters ranging from 

3-15EC, with the greatest percentages found between 

approximately 8-12EC. Their depth range was from 

approximately 6-75 m, with the majority at 6-25 m. 

During the autumn they were found in waters ranging 

from 5-21EC; their temperature distribution was 

somewhat bimodal, with the major peaks between about 

16-18EC. Their depth range was from 1-65 m, with the 

majority found between 6-25 m. 

 The distributions and abundances of both juvenile 

and adult winter skate in Long Island Sound from April to 

November 1984-1994, based on the Connecticut Fisheries 

Division bottom trawl surveys, are shown in Figures 9-

10. The following description of their distributions 

relative to depth and bottom type is taken verbatim from 

Gottschall et al. (2000). 

 Winter skate were most commonly taken during the 

spring and late fall, occurring on average in 16.4% of 

samples during these periods (Figure 10D). Abundance 

was highest during April, and decreased thereafter until 

August when none were recorded in the survey (Figure 

10A). During the spring, winter skate were most abundant 

on sand bottom in the Mattituck Sill and Eastern Basin 

(Figure 9, Figure 10B). Abundance was similar in most 

depths, with the exception of depths between 9-18 m, 

where abundance was lower (Figure 10C). Winter skate 

abundance increased again in October and November, but 

they were not as concentrated on the Mattituck Sill and in 

the Eastern Basin as during the spring (Gottschall et al. 

2000). 

 The seasonal distributions of juveniles the HudsonRaritan estuary relative to bottom water temperature, 

depth, salinity, and dissolved oxygen based on 1992-1997 

Hudson-Raritan trawl surveys are shown in Figure 21. 

The surveys show that during the winter juveniles were 

found mostly between 0-7EC, with > 50% at 4-5EC. Their 
Page 8 

depth range during that season was between 4-22 m, with 

most caught between 5-8 m. Their salinities ranged 

between 20-35 ppt, most were found roughly between 23-

32 ppt. They were found over a range of dissolved 

oxygen levels of between 9-14 ppm with a few at 5 ppm; 

most were found between 10-12 ppm. In spring, juvenile 

winter skate were found over a wider temperature range 

of between about 2-17EC, with bimodal peaks between 

approximately 5-9EC and 15-17EC and with most found 

between 6-9EC. The bimodality may be a function of the 

greater number of trawls done within those temperature 

intervals. Their depth range was between 4-18 m, with the 

majority between 5-8 m. Their salinities ranged between 

15-33 ppt, most were found at 25 ppt and between 27-28 

ppt. They were found over a range of dissolved oxygen 

levels of between 7-13 ppm with most found between 10-

11 ppm. Few were caught in summer; they were found 

between about 16-21EC and at depths of 7 m, 18 m, and 

20 m. Their salinities ranged between 28-29 ppt and at 32 

ppt, and they were found over a range of dissolved 

oxygen levels of between 7-8 ppm. In the fall they were 

found between 5-17EC, with most spread between 5-

13EC. Their depth range during the fall was between 4-21 

m, with the majority between 5-8 m. Their salinities 

ranged between 17-34 ppt, with most spread roughly 

between 23-31 ppt. They were found over a range of 

dissolved oxygen levels of between 6-12 ppm with most 

found between 8-9 ppm. Based on the above evidence it 

appears that juvenile winter skate in the Hudson-Raritan 

estuary are found in warmer waters during the spring and 

fall as compared to winter, and remain mostly around 

depths of 5-8 m during those three seasons. 

 The seasonal distributions of both juveniles and 

adults in Delaware Bay relative to bottom water 

temperature, depth, salinity, and dissolved oxygen based 

on 1966-1999 Delaware Division of Fish and Wildlife 

bottom trawl surveys are shown in Figure 22. During the 

winter they were found between 3-9EC, with the majority 

between 7-8EC. Their depth range during winter was 

between about 7-18 m, with peaks at 14 m, 16 m, and 17 

m. Their salinities ranged between about 22-30 ppt and 

34-35 ppt, most were found at 26 ppt and between 28-29 

ppt. They were found over a range of dissolved oxygen 

levels of between 8-11 ppm, with the majority found 

between 9-11 ppm. In spring, they were found over a 

wider temperature range of between 4-17EC, with peaks 

scattered throughout the range (e.g., 5EC, 11EC, and 

13EC). Their depth range was between 7-17 m, with a 

few at 21 m, and most at 8 m and 12-14 m. Their 

salinities ranged between 21-33 ppt, with a few at 15 ppt, 

and peaks scattered throughout with the two highest at 28 

ppt and especially 30 ppt. They were found over a range 

of dissolved oxygen levels of between 7-11 ppm and 13-

15 ppm, most were found between 8-11 ppm. In summer, 

there were too few winter skate caught to plot their 

distributions relative to the habitat parameters. During fall 

they were found between 8-13EC, with a few at 16EC; 

most were between 8-11EC. Their depth range during the 

fall was spread between 7-11 m and at 13 m and between 

18-19 m. Most were spread between 7-8 m, at 13 m, and 

at 18 m. Their salinities ranged between about 26-32 ppt, 

with a few at 16 ppt and 22 ppt. Peaks were at 28 ppt, 30 

ppt, and at 32 ppt. They were found over a range of 

dissolved oxygen levels of between 7-10 ppm, the 

majority were at 9 ppm. 

ADULTS 

 The spring and fall distributions of adult winter skate 

relative to bottom water temperature, depth, and salinity 

based on 1963-2002 NEFSC bottom trawl surveys from 

the Gulf of Maine to Cape Hatteras are shown in Figure 

23. In spring, adults were caught at temperatures between 

2-11EC, with most between 4-6EC and a peak at 5EC. 

During that period they were found at a depth range of 1-

300 m, with the majority spread between 31-60 m. They 

were found at salinities of between 30-36 ppt, with the 

majority at 33 ppt. During the fall, they were caught over 

a temperature range of 5-19EC, with most caught between 

about 11-15EC and a peak at 14EC. They were found over 

a depth range of 11-300 m, with most caught at depths 

between about 21-70 m and peaks at 31-50 m. They were 

found at salinities of between 31-34 ppt, with 80-90% at 

32 ppt. 

 The spring and autumn distributions of adults in 

Massachusetts coastal waters relative to bottom water 

temperature and depth are shown in Figure 24. In the 

spring they were found in waters ranging from 2-16EC; 

the majority were spread between approximately 6-12EC. 

During that same season, the adults were found from 1-75 

m, with most between 6-20 m. In autumn they were found 

between 5-19EC. The distribution was somewhat 

bimodal, with a peak at 10EC and a minor one around 15-

16EC. The depth range of the adults during autumn was 

around 1-75 m, with most found between 6-25 m. 

 The distributions and abundances of both juvenile 

and adult winter skate in Long Island Sound relative to 

depth and bottom type were discussed previously (Figures 

9-10; Gottschall et al. [2000]). 

 Too few adults were found in the Hudson-Raritan 

estuary to plot their distributions relative to habitat 

parameters. 

 The seasonal distributions of both juveniles and 

adults in Delaware Bay relative to bottom water 

temperature, depth, salinity, and dissolved oxygen based 

on Delaware Division of Fish and Wildlife bottom trawl 

surveys were discussed previously (Figure 22). 

STATUS OF THE STOCKS 

 The following section is based on Northeast Fisheries 

Science Center (2000a, b). 

 The principal commercial fishing method used to 
Page 9 

catch all seven species of skates [winter, little, barndoor

(Dipturus laevis), winter, thorny, clearnose (Raja 

eglanteria), rosette (Leucoraja garmani), smooth 

(Malacoraja senta)] is otter trawling. Skates are 

frequently taken as bycatch during groundfish trawling 

and scallop dredge operations and discarded recreational 

and foreign landings are currently insignificant, at < 1% 

of the total fishery landings. 

 Skates have been reported in New England fishery 

landings since the late 1800s. However, commercial 

fishery landings, primarily from off Rhode Island, never 

exceeded several hundred metric tons until the advent of 

distant-water fleets during the 1960s. Landings are not 

reported by species, with over 99% of the landings 

reported as “unclassified skates.” Skate landings reached 

9,500 mt in 1969, but declined quickly during the 1970s, 

falling to 800 mt in 1981 (Figure 25). Landings have 

since increased substantially, partially in response to 

increased demand for lobster bait, and more significantly, 

to the increased export market for skate wings. Wings are 

taken from winter and thorny skates, the two species 

currently used for human consumption. Bait landings are 

presumed to be primarily from little skate, based on areas 

fished and known species distribution patterns. Landings 

for all skates increased to 12,900 mt in 1993 and then 

declined somewhat to 7,200 mt in 1995. Landings have 

increased again since 1995, and the 1998 reported 

commercial landings of 17,000 mt were the highest on 

record (Figure 25). 

 The biomass for the seven skate species is at a 

medium level of abundance. For the aggregate complex, 

the NEFSC spring survey index of biomass was relatively 

constant from 1968-1980, then increased significantly to 

peak levels in the mid- to late 1980s. The index of skate 

complex biomass then declined steadily until 1994, but 

has recently increased again. The large increase in skate 

biomass in the mid- to late 1980s was dominated by 

winter and little skate. The biomass of large sized skates 

(> 100 cm max. length: barndoor, winter, and thorny) has 

steadily declined since the mid-1980s. The recent increase 

in aggregate skate biomass has been due to an increase in 

small sized skates (< 100 cm max. length: little, 

clearnose, rosette, and smooth) – primarily little skate. 

Winter skate abundance is currently about the same as in 

the early 1970s, and is about 25% of the peak observed in 

the mid-1980s (Figure 25). Winter skate was, until 

recently, considered to be overfished (Northeast Fisheries 

Science Center 2000a, b), but its status has recently been 

changed so that it is no longer considered to be in an 

overfished condition (NMFS 2002). 

RESEARCH NEEDS 

 Imprecise reporting of fishery statistics where several 

skate species are lumped together under one category 

(e.g., “unclassified skates” or “skates spp.”) can mask 

basic changes in community structure and profound 

reduction in populations of larger, slower growing species 

(Dulvy et al. 2000; Musick et al. 2000). Thus, it is 

important to have fishery-independent data on skates 

where the individual species are reported; it is also 

necessary to work out any identification problems 

between winter and little skate. 

 Northeast Fisheries Science Center (2000b) also 

suggests the following research needs: 

• More life history studies (including age, growth, 

maturity, and fecundity studies) are necessary. 

• Studies of stock structure are needed to identify unit 

stocks. 

• Explore possible stock-recruit relationships by 

examination of NEFSC survey data. 

• Investigate trophic interactions between skate species 

in the complex, and between skates and other 

groundfish. 

• Investigate the influence of annual changes in water 

temperature or other environmental factors on shifts 

in the range and distribution of the species in the 

skate complex, and establish the bathymetric 

distribution of the species in the complex in the 

northwest Atlantic. 

• Investigate historical NEFSC survey data from the 

R/V Albatross III during 1948-1962 when they 

become available, as they may provide valuable 

historical context for long-term trends in skate 

biomass. 

ACKNOWLEDGMENTS 

 The authors thank Barry Shafer, John McCarthy, 

Tom Finneran, Annette Kalbach, and Meredith Lock for 

producing the maps and graphics. Thanks also to Claire 

Steimle and Judy Berrien for literature reviews and 

interlibrary loans. Frank Almeida and Kathy Sosebee of 

the NEFSC Woods Hole provided much needed 

information, input, and reviews. 

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skates Raja erinacea and R. ocellata (Pisces: Rajidae). 

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distinguishing between immature specimens of the 

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relative abundance of seven species of skates (Pisces: 

Rajidae) which occur between Nova Scotia and Cape 

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skate (Raja erinacea) off outer Cape Cod, 

Massachusetts. M.S. thesis, Southeastern Mass. Univ., 

North Dartmouth, MA. 71 p. 

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Reid, R., F. Almeida, and C. Zetlin. 1999. Essential fish 

habitat source document: Fishery independent 

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Page 12 

Table 1. Summary of habitat parameters for winter skate, based on the pertinent literature. 

Life Stage Depth Substrate Temperature 

Juveniles 1 Generally caught at depths from 

shoreline to 371 m, although most 

abundant < 111 m. In Gulf of 

Maine, most abundant 46-64 m 

but occasionally occur < 1 m and 

down to 285 m; they’re 

considered rare at depths < 2-7 m. 

May be permanent resident off 

southern New England between 

15-46 m, although there are 

seasonal fluctuations in 

abundance. Has been captured off 

the Mid-Atlantic states during the 

winter at depths from 33-113 m. 

In Long Island Sound during 

spring 1984-1994, abundance of 

winter skate was similar in most 

depths, with the exception of 

depths between 9-18 m, where 

abundance was lower. 

Sand and gravel bottoms but 

reported from mud bottoms in 

Passamaquoddy Bay. In Long 

Island Sound during spring 1984-

1994, most abundant on sand 

bottoms in the Mattituck Sill and 

Eastern Basin. On Scotian Shelf, 

confined to sand and gravel 

bottoms; bottom type, rather than 

depth, may be more important in 

determining distributions of 

winter skate. 

Remains buried in depressions 

during the day and are more 

active at night. This is probably 

not due to diel foraging since no 

diel periodicity in feeding 

intensity has been observed; they 

may feed at any time during a 24 

hour period 

Recorded over a temperature 

range of -1.2EC to 19EC. 

Reported in Gulf of Maine at 

20EC along Massachusetts coast 

in summer, down to 1-2EC in the 

coastal belt in winter, near 0EC in 

the Bay of Fundy region in some 

years. Reported at a temperature 

range of 2-15EC from southern 

Nova Scotia to Cape Hatteras. 

Captured off Mid-Atlantic states 

during winter at 10-12EC. 

Adults 2

Same as for juveniles. Same as for juveniles. Same as for juveniles. 

1 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 et al. (2000); McEachran (2002). 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 et al. (2000); McEachran (2002).
Page 13 

Table 1. cont’d. 

Life Stage Prey Predators/Species Associations 

Juveniles 1 Polychaetes and amphipods most important 

prey in terms of numbers or occurrence, 

followed by decapods, isopods, bivalves, 

fishes. Hydroids also ingested. In terms of 

weight, amphipods, decapods and fish can 

be most important; fish are especially 

prevalent in larger skate. Polychaetes 

include: Nephtys spp., Nereis spp., 

Lumbrineris fragilis, Ophelia denticulata, 

maldanids (mostly Clymenella torquata), 

Aphrodite hastata. Amphipods: haustoriids, 

Leptocheirus pinguis, Monoculodes sp., 

Hippomedon serratus, ampeliscids, 

Paraphoxus sp., Tmetonyx sp., Unciola

irrorata, Byblis serrata, oedicerotids. 

Decapods: Crangon septemspinosa, Cancer

irroratus, pagurid crabs, Dichelopandalus

leptocerus, pandalid shrimp. Isopods:

Cirolana (= Politolana?) polita. Bivalves 

include Solemya sp. and Ensis directus. 

Sand lance was the most frequently eaten 

fish; yellowtail flounder, longhorn sculpin, 

hakes, other skate, herring, butterfish 

occasionally eaten. Generally, in terms of 

numbers or occurrence, crustaceans made up 

> 50% of the diet for skate < 61 cm TL, 

while fish (and often bivalves) were a major 

part of the diet of skate >79-91 cm TL. 

Overall crustaceans declined in importance 

with increasing skate size (includes both 

amphipods and often decapods) while 

polychaetes increased with increasing skate 

size until the skate were about 81 cm TL. 

Amphipods occurred more frequently than 

decapods until the skates were > 71 cm TL. 

On Georges Bank, decline in importance of 

fish prey in spring may be related to 

seasonal movements of prey. In terms of 

weight, crustaceans were dominant in the 

diet of skate < 31-50 cm TL, while fish, 

mostly sand lance, were dominant in the diet 

of skate 51-110 cm TL. For skate < 31 cm 

TL, amphipods dominated, especially L. 

pinguis. For skate 31-50 cm TL, decapods 

dominated, especially C. septemspinosa and 

C. irroratus. 

Predators: sharks, other skates, gray seals, and gulls. 

Winter and little skate co-occur from Nova Scotia to Cape Hatteras. 

Although winter and little skate are sympatric species with similar 

habitat requirements, there’s not a high degree of competitive 

interaction between them because they are positively correlated by 

abundance. Also, winter skate feeds largely on infauna, while little 

skate predominately selects epifauna. Sympatric populations of winter 

and little skate also undergo character displacement in order to avoid 

direct competition for food resources. Using 1973-1997 NEFSC data 

from Nova Scotia to Cape Hatteras and NEFSC food habits database, 

both small (10-30 cm TL) and medium (31-60 cm TL) sized winter 

skate belonged to “Amphipod/shrimp eaters” group, along with little 

skate and cusk eel; prey included amphipods, polychaetes, shrimp, 

zooplankton. Largest winter skate (61 to > 80 cm TL) by themselves 

in a subgroup of “Piscivores” because their diet contained a high 

proportion of sand lance. Again, a trend toward increasing piscivory 

with size. 

On Georges Bank, winter skate belongs to spatial assemblages and 

trophic groups that include Atlantic cod, little skate, longhorn sculpin, 

yellowtail flounder, red and silver hake, haddock, spiny dogfish, 

butterfish, fourspot flounder, windowpane, winter flounder, sea raven, 

thorny skate, Atlantic herring. Also on Georges Bank, winter skate 

falls into various dietary guilds or trophic groups, depending on the 

study. Garrison and Link (2000b): “Bentho-pelagic” group included 

31-60 cm TL winter skate, little skate, longhorn sculpin, Atlantic cod. 

Diets of these species included shrimp such as pandalids and C. 

septemspinosa, and benthic invertebrates including polychaetes, 

gammarid amphipods, bivalves. “Skate” group included > 80 cm TL 

winter skate, pollock, windowpane. Prey was a combination of fish 

and benthic prey, with a high proportion of sand lance. Garrison 

(2000): In autumn, 61 cm to > 80 cm TL winter skate by themselves 

in a subgroup of the “Piscivorous” group, feeding on sand lance, 

silver hake, and Atlantic herring, plus benthic invertebrates. 10-60 cm 

TL winter skate in “Demersal predators” group with flatfish, 

haddocks, little skate, thorny skate. Prey included gammarid 

amphipods, polychaetes, isopods, Cancer crabs, C. septemspinosa. 

During spring, 10-60 cm TL winter skate in “Shrimp/amphipod 

predators” group with hakes, longhorn sculpin, Atlantic cod, fourspot 

flounder, little skate, thorny skate. Prey included gammarid 

amphipods, pandalids, C. septemspinosa, polychaetes, Cancer crabs. 

Winter skate 61 cm to > 80 cm TL by themselves in a “Generalist” 

subgroup, consuming bivalves, polychaetes, sand lance, herring. 

Decline in importance of fish prey, 35% fish in autumn and 16% in 

spring, probably related to seasonal movements of prey. 

Adults2

Same as for juveniles; however, note that 

larger skates consume more polychaetes and 

fish while crustaceans decline in the diet. 

Same as for juveniles, but note differences between smaller and larger 

skates. 

1 Bigelow and Schroeder (1953); McEachran (1973); McEachran and Musick (1975); McEachran et al. (1976); McEachran and 

Martin (1977); Overholtz and Tyler (1985); Scott and Scott (1988); Kaplan 1999; Bowman et al. (2000); Garrison (2000); Garrison 

and Link (2000a, b); Avent et al. (2001); (Tsou and Collie 2001a, b); NEFSC 1973-1990 food habits database. 2 Bigelow and Schroeder (1953); McEachran (1973); McEachran and Musick (1975); McEachran et al. (1976); McEachran and 

Martin (1977); Overholtz and Tyler (1985); Scott and Scott (1988); Kaplan 1999; Bowman et al. (2000); Garrison (2000); Garrison 

and Link (2000a, b); Avent et al. (2001); (Tsou and Collie 2001a, b); NEFSC 1973-1990 food habits database.
Page 14 

Table 2. Summary of habitat parameters for winter skate, based on the most recent NEFSC and state surveys mentioned 

in the text. 

Life Stage Survey Depth Temperature Salinity/DO 

1963-2002 spring 

and fall NEFSC 

trawl surveys from 

Gulf of Maine to 

Cape Hatteras. 

Spring: range of 1-300 m, 

most between about 11-70 m. 

Fall: range of 1-400 m, most 

between about 21-80 m. 

Spring: range of 1-12EC, 

majority at about 4-5EC. 

Fall: range of 5-21EC, most 

spread between about 7-16EC, 

peaks at about 13-15EC. 

Spring: range of 28-35 ppt, 

most between 32-33 ppt.

Fall: range of 31-35 ppt, 

majority between 32-33 ppt.

1978-2002 

Massachusetts 

inshore trawl 

surveys. 

Spring: range of 

approximately 6-75 m, 

majority at 6-25 m. 

Fall: range of 1-65 m, 

majority between 6-25 m. 

Spring: range of 3-15EC, 

greatest percentages between 

approximately 8-12EC. 

Fall: range of 5-21EC, 

temperature distribution 

somewhat bimodal, major 

peak between about 16-18EC. 

1992-1997 NEFSC 

trawl surveys of the 

Hudson-Raritan 

estuary. 

Winter: range of 4-22 m, most 

between 5-8 m. 

Spring: range of 4-18 m, 

majority between 5-8 m. 

Summer: few caught, found at 

7 m, 18 m, and 20 m. 

Fall: range of 4-21 m, 

majority between 5-8 m.

Winter: range of 0-7EC, > 

50% between 4-5EC. 

Spring: range of 2-17EC, with 

bimodal peaks between 5-9EC 

and 15-17EC, most between 6-

9EC. 

Summer: few caught, found 

between about 16-21EC. 

Fall: range of 5-17EC, most 

spread between 5-13EC.

Winter: range of 20-35 ppt, 

most between roughly 23-32 

ppt / range of 9-14 ppm with a 

few at 5 ppm, most between 

10-12 ppm. 

Spring: range of 15-33 ppt, 

most at 25 ppt and between 

27-28 ppt / range of 7-13 ppm, 

most between 10-11 ppm. 

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. 

Juveniles

1966-1999 

Delaware Division 

of Fish and Wildlife 

bottom trawl 

surveys of 

Delaware Bay 

(juveniles and 

adults combined) 

Winter: range of about 7-18 m, 

peaks at 14 m, 16 m, and 17 

m. 

Spring: range of 7-17 m, a few 

at 21 m, most at 8 m and 12-

14 m. 

Fall: range of 7-11 m and at 

13 m and between 18-19 m; 

most spread between 7-8 m, at 

13 m, and at 18 m.

Winter: range of 3-9EC, 

majority between 7-8EC. 

Spring: range of 4-17EC, 

peaks scattered throughout 

(e.g., 5EC, 11EC, and 13EC). 

Fall: range of 8-13EC, a few 

at 16EC; most between 8-

11EC.

Winter: range of about 22-30 

ppt and 34-35 ppt, most at 26 

ppt and between 28-29 ppt / 

range of 8-11 ppm, majority 

between 9-11 ppm. 

Spring: range of 21-33 ppt, a 

few at 15 ppt; peaks scattered 

throughout with the two 

highest at 28 ppt and 

especially 30 ppt / range of 7-

11 ppm and 13-15 ppm, most 

between 8-11 ppm. 

Fall: range of about 26-32 ppt, 

a few at 16 ppt and 22 ppt; 

peaks at 28 ppt, 30 ppt, and 32 

ppt / range of 7-10 ppm, 

majority at 9 ppm.
Page 15 

Table 2. cont’d. 

Life Stage Survey Depth Temperature Salinity/DO 

1963-2002 spring 

and fall NEFSC 

trawl surveys from 

Gulf of Maine to 

Cape Hatteras. 

Spring: range of 1-300 m, 

majority spread between 31-60 

m. 

Fall: range of 11-300 m, most 

between about 21-70 m, peaks 

at 31-50 m. 

Spring: range of 2-11EC, most 

between 4-6EC, peak at 5EC. 

Fall: range of 5-19EC, most 

between 11-15EC, peak at 

14EC. 

Spring: range of 30-36 ppt, 

majority at 33 ppt. 

Fall: range of 31-34 ppt, with 

80-90% at 32 ppt. 

1978-2002 

Massachusetts 

inshore trawl 

surveys. 

Spring: range of 1-75 m, most 

between 6-20 m. 

Fall: range of around 1-75 m, 

most between 6-25 m. 

Spring: range of 2-16EC, 

majority spread between 

approximately 6-12EC. 

Fall: range of 5-19EC, 

distribution somewhat 

bimodal: peak at 10EC and a 

minor one around 15-16EC. 

Adults

1966-1999 

Delaware Division 

of Fish and Wildlife 

bottom trawl 

surveys of 

Delaware Bay 

(juveniles and 

adults combined) 

See juveniles. See juveniles. See juveniles. 
Page 16 

Figure 1. The winter skate, Leucoraja ocellata (Mitchill 1815), male, from Murdy et al. (1997).
Page 17 

Figure 2. Egg case of winter skate, from Bor (2001).
Page 18 

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]. 

1973-1980 1981-1990

Unidentified 

Crustacea 1.6%

Amphipoda 37.5%

Polychaeta 9.4%

Other prey 

phyla 17.2%

Decapoda 23.4%

Other crustacea 1.6%

Isopoda 9.4%

n = 10 n = 50

Decapoda 24.6%

Other prey 

phyla 3.3%

Polychaeta 13.1%

Amphipoda 42.6%

Other crustacea 16.4%

21-30 cm

n = 48

Amphipoda 36.9%

Unidentified 

Crustacea 5.9%

Fish 2.5%

Isopoda 8.1%

Other 

Crustacea 2.5%

Decapoda 22.9%

Other prey 

phyla 9.4%

Polychaeta 9.4%

Amphipoda 38.8%

Polychaeta 17.1%

Other prey 

phyla 1.1%

Crustacea 

shrimp 6.5%

Other Crustacea 2.2% Decapoda 17.1% Fish 4.4%

Unidentified 

Crustacea 2.4%

Mollusca 4.9%

Isopoda 5.4%

31-40 cm n = 462

41-50 cm

Unidentified 

Crustacea 4.2%

Fish 3.4%

Decapoda 22.3%

Other prey 

phyla 9.0%

Polychaeta 18.9%

Amphipoda 33.5%

Isopoda 6.8%

Other 

Crustacea 2.0%

n = 53

Amphipoda 33.1%

Other 

Crustacea 1.1%

Isopoda 8.2%

Mollusca 4.6%

Unidentified 

Fish 7.9% Crustacea 1.8%

Decapoda 14.3%

Crustacea 

shrimp 4.2%

Other prey 

phyla 1.8%

Polychaeta 23.1%

n = 516
Page 19 

Figure 3. cont’d. 

1973-1980 1981-1990

Decapoda 11.9%

Other prey 

phyla 11.4%

Polychaeta 28.5%

Amphipoda 31.6%

Unidentified 

Crustacea 1.6%

Fish 6.2%

Isopoda 6.7%

Other 

Crustacea 2.1%

51-60 cm

n = 31

Decapoda 9.3%

Other prey 

phyla 0.4%

Polychaeta 26.3%

Amphipoda 26.0%

Unidentified 

Crustacea 1.3%

Fish 12.6%

Isopoda 12.1%

Other 

Crustacea 0.9%

Mollusca 8.2%

Crustacea 

Shirmp 2.9%

n = 399

61-70 cm

n = 78

Other prey 

phyla 4.2%

Polychaeta 31.8% Amphipoda 20.3%

Unidentified 

Crustacea 2.1%

Fish 10.9%

Isopoda 7.0%

Other 

Crustacea 2.7%

Mollusca 9.1%

Decapoda 11.8%

n = 394

Decapoda 

10.5%

Polychaeta 28.4%

Amphipoda 13.2%

Fish 23.9%

Other Isopoda 12.0%

Crustacea 1.6%

Mollusca 9.3%

Crustacea 

Shirmp 1.0%

Other prey 

phyla 4.4%

Polychaeta 34.8%

Amphipoda 11.8%

Unidentified 

Crustacea 1.1%

Fish 16.0%

Isopoda 7.9%

Other 

Crustacea 2.0%

Mollusca 9.4%

Decapoda 12.7%

71-80 cm

n = 149

Decapoda 

10.6%

Polychaeta 25.2%

Amphipoda 6.7%

Fish 35.6%

Isopoda 8.8% Other 

Crustacea 0.9%

Mollusca 10.3%

Crustacea 

Shirmp 1.9%

n = 450
Page 20 

Figure 3. cont’d. 

1973-1980 1981-1990

91-100 cm

n = 91

Amphipoda 5.2%

Unidentified 

Crustacea 2.1%

n = 488

81-90 cm

n = 170

Decapoda 

10.5%

Other prey 

phyla 6.3%

Polychaeta 27.9%

Amphipoda 7.8%

Unidentified 

Crustacea 2.4%

Fish 28.8%

Isopoda 7.2%

Other 

Crustacea 1.8%

Mollusca 7.2%

n = 710

Amphipoda 3.0%

Decapoda 8.1%

Fish 41.9%

Other 

Crustacea 0.9% Polychaeta 22.2%

Isopoda 4.7%

Mollusca 16.9%

Crustacea 

Shirmp 2.2%

Other prey phyla 10.3%

Polychaeta 10.3%

Fish 56.7%

Other 

Crustacea 5.2%

Decapoda 10.3% Decapoda 4.9%

Polychaeta 13.1%

Amphipoda 3.9%

Fish 62.6%

Isopoda 1.6%

Mollusca 11.8%

Crustacea Shirmp 2.0%
Page 21 

Winter Skate

NEFSC Bottom Trawl Surveys

(1964 - 2002)

Number per tow

Winter/juveniles

(<= 84 cm)

Present

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 

Winter Skate

NEFSC Bottom Trawl Surveys

(1968 - 2002)

Spring/juveniles

(<= 84 cm)

NPTOW

1 - 10

11 - 100

101 - 1000

> 1000

Number per tow

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. 

Winter Skate

NEFSC Bottom Trawl Surveys

(1963 - 1995)

Number per tow

Summer/juveniles

 (<= 84 cm)

Present
Page 24 

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]. 

Winter Skate

NEFSC Bottom Trawl Surveys

(1963 - 2001)

Fall/juveniles

(<= 84 cm)

NPTOW

1 - 10

11 - 100

101 - 1000

> 1000

Number per tow
Page 25 

New Hampshire

Cape Ann

Massachusetts

Cape Cod

Martha's

Vineyard

Nantucket

Spring/juveniles

NPTOW

1 - 10

11 - 100

101 - 500

> 500

Winter Skate

Massachusetts Inshore

Trawl Survey

(1978 - 2002)

Spring/juveniles

(<= 84 cm)

 Number per tow

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 

Figure 8. cont’d.

New Hampshire

Cape Ann

Massachusetts

Cape Cod

Martha's

Vineyard

Nantucket

Spring/juveniles

NPTOW

1 - 10

11 - 100

101 - 500

> 500

Winter Skate

Massachusetts Inshore

Trawl Survey

(1978 - 2001)

Fall/juveniles

(<= 84 cm)

 Number per tow
Page 27 

Figure 9. Distribution and abundance of juvenile 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 

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 

Figure 11. Seasonal distribution and abundance of juvenile winter skate in the Hudson-Raritan estuary, based on 

Hudson-Raritan trawl surveys, 1992-1997 [see Reid et al. (1999) for details]. 
Page 30 

Figure 11. cont’d. 
Page 31 

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 

Figure 12. cont’d. 
Page 33 

Figure 12. cont’d. 
Page 34 

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. 

Winter Skate

NEFSC Bottom Trawl Surveys

(1964 - 2002)

Number per tow

Winter/adults

 (> 84 cm)

Present
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 al. (1999) for details].

Winter Skate

NEFSC Bottom Trawl Surveys

(1968 - 2002)

Spring/adults

(> 84 cm)

NPTOW

1 - 10

11 - 100

101 - 1000

> 1000

Number per tow
Page 37 

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.

Winter Skate

NEFSC Bottom Trawl Surveys

(1963 - 1995)

Number per tow

Summer/adults

 (> 84 cm)

Present
Page 38 

Fall/adults

(> 84 cm)

NPTOW

1 - 10

11 - 100

101 - 1000

> 1000

Winter Skate

NEFSC Bottom Trawl Surveys

(1963 - 2001)

Number per tow

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 

New Hampshire

Cape Ann

Massachusetts

Cape Cod

Martha's

Vineyard

Nantucket

Spring/juveniles

NPTOW

1 - 10

11 - 100

101 - 500

> 500

Winter Skate

Massachusetts Inshore

Trawl Survey

(1978 - 2002)

Spring/adults

(> 84 cm)

 Number per tow

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 

Figure 17. cont’d.

New Hampshire

Cape Ann

Massachusetts

Cape Cod

Martha's

Vineyard

Nantucket

Spring/juveniles

NPTOW

1 - 10

11 - 100

101 - 500

> 500

Winter Skate

Massachusetts Inshore

Trawl Survey

(1978 - 2001)

Fall/adults

(> 84 cm)

 Number per tow
Page 41 

Figure 18. Seasonal distribution and abundance of adult winter skate in the Hudson-Raritan estuary, based on HudsonRaritan trawl surveys, 1992-1997 [see Reid et al. (1999) for details]. 
Page 42 

0

10

20

30

40

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)

Percent

Trawls N=10537

Occurrence N=2296

Catch N=35461

0

5

10

15

20

25

30

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

Bottom Depth (m)

Percent

Trawls N=12193

Occurrence N=2744

Catch N=46565

0

10

20

30

40

50

26 27 28 29 30 31 32 33 34 35 36 37

Salinity (PPT)

Percent

Trawls N=1959

Occurrence N=507

Catch N=4900

Winter Skate

NEFSC Bottom Trawl Survey 

Spring/Juveniles 

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 

0

5

10

15

20

2 4 6 8 10 12 14 16 18 20 22 24 26 28

Bottom Temperature (°C)

Percent

Trawls N=11844

Occurrence N=1954

Catch N=20568

0

5

10

15

20

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

>500

Bottom Depth (m)

Percent

Trawls N=13543

Occurrence N=2210

Catch N=25045

0

20

40

60

27 29 30 31 32 33 34 35 36 37

Salinity (PPT)

Percent

Trawls N=1956

Occurrence N=404

Catch N=4255

Winter Skate

NEFSC Bottom Trawl Survey 

Fall/Juveniles 

Figure 19. cont’d. 
Page 44 

0

10

20

30

40

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16

Bottom Temperature (°C)

Percent

Trawls N=2312

Occurrence N=1310

Catch N=86254

0

10

20

30

40

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2385

Occurrence N=1342

Catch N=90236

Winter Skate

Massachusetts Inshore Trawl Survey 

Spring/Juveniles

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 

0

5

10

15

20

25

4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

Bottom Temperature (°C)

Percent

Trawls N=2067

Occurrence N=958

Catch N=46556

0

10

20

30

40

50

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2156

Occurrence N=1017

Catch N=52536

Winter Skate

Massachusetts Inshore Trawl Survey 

Fall/Juveniles

Figure 20. cont’d. 
Page 46 

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. 

Winter 

0

10

20

30

40

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=462

Occurrence N=205

Catch N=562

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=468

Occurrence N=205

Catch N=562

0

10

20

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

Salinity (PPT)

Percent

Trawls N=466

Occurrence N=205

Catch N=562

0

10

20

30

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=265

Occurrence N=125

Catch N=292

Winter 

0

10

20

30

40

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=462

Occurrence N=205

Catch N=562

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=468

Occurrence N=205

Catch N=562

0

10

20

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

Salinity (PPT)

Percent

Trawls N=466

Occurrence N=205

Catch N=562

0

10

20

30

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=265

Occurrence N=125

Catch N=292
Page 47 

Figure 21. cont’d.

Spring 

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=326

Occurrence N=113

Catch N=261

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=330

Occurrence N=114

Catch N=262

0

10

20

30

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

Salinity (PPT)

Percent

Trawls N=326

Occurrence N=113

Catch N=261

0

10

20

30

40

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=177

Occurrence N=57

Catch N=105

Spring 

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=326

Occurrence N=113

Catch N=261

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=330

Occurrence N=114

Catch N=262

0

10

20

30

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

Salinity (PPT)

Percent

Trawls N=326

Occurrence N=113

Catch N=261

0

10

20

30

40

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=177

Occurrence N=57

Catch N=105
Page 48 

Figure 21. cont’d.

Summer 

0

10

20

30

40

50

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=352

Occurrence N=4

Catch N=7

0

10

20

30

40

50

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=356

Occurrence N=4

Catch N=7

0

10

20

30

40

50

60

70

80

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

Salinity (PPT)

Percent

Trawls N=347

Occurrence N=4

Catch N=7

0

10

20

30

40

50

60

70

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=215

Occurrence N=3

Catch N=5

Summer 

0

10

20

30

40

50

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=352

Occurrence N=4

Catch N=7

0

10

20

30

40

50

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=356

Occurrence N=4

Catch N=7

0

10

20

30

40

50

60

70

80

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

Salinity (PPT)

Percent

Trawls N=347

Occurrence N=4

Catch N=7

0

10

20

30

40

50

60

70

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=215

Occurrence N=3

Catch N=5
Page 49 

Figure 21. cont’d. 

Fall 

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=570

Occurrence N=224

Catch N=583

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=586

Occurrence N=229

Catch N=590

0

10

20

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

Salinity (PPT)

Percent

Trawls N=532

Occurrence N=209

Catch N=563

0

10

20

30

40

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=458

Occurrence N=183

Catch N=509

Fall 

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=570

Occurrence N=224

Catch N=583

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=586

Occurrence N=229

Catch N=590

0

10

20

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

Salinity (PPT)

Percent

Trawls N=532

Occurrence N=209

Catch N=563

0

10

20

30

40

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=458

Occurrence N=183

Catch N=509
Page 50 

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. 

Winter 

0

10

20

30

40

50

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=166

Occurrence N=24

Catch N=100

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=166

Occurrence N=24

Catch N=100

0

10

20

30

40

50

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

Salinity (PPT)

Percent

Trawls N=157

Occurrence N=24

Catch N=100

0

10

20

30

40

50

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=125

Occurrence N=21

Catch N=90

Winter 

0

10

20

30

40

50

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=166

Occurrence N=24

Catch N=100

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=166

Occurrence N=24

Catch N=100

0

10

20

30

40

50

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

Salinity (PPT)

Percent

Trawls N=157

Occurrence N=24

Catch N=100

0

10

20

30

40

50

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=125

Occurrence N=21

Catch N=90
Page 51 

Figure 22. cont’d. 

Spring 

0

10

20

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=373

Occurrence N=49

Catch N=102

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=378

Occurrence N=50

Catch N=103

0

10

20

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

Salinity (PPT)

Percent

Trawls N=374

Occurrence N=50

Catch N=103

0

10

20

30

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=312

Occurrence N=39

Catch N=75

Spring 

0

10

20

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=373

Occurrence N=49

Catch N=102

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=378

Occurrence N=50

Catch N=103

0

10

20

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

Salinity (PPT)

Percent

Trawls N=374

Occurrence N=50

Catch N=103

0

10

20

30

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=312

Occurrence N=39

Catch N=75
Page 52 

Figure 22. cont’d. 

Fall 

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=422

Occurrence N=16

Catch N=39

0

10

20

30

40

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=405

Occurrence N=16

Catch N=39

0

10

20

30

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

Salinity (PPT)

Percent

Trawls N=399

Occurrence N=16

Catch N=39

0

10

20

30

40

50

60

70

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=341

Occurrence N=14

Catch N=37

Fall 

0

10

20

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Depth (m)

Percent

Trawls N=422

Occurrence N=16

Catch N=39

0

10

20

30

40

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

Temperature (0C)

Percent

Trawls N=405

Occurrence N=16

Catch N=39

0

10

20

30

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

Salinity (PPT)

Percent

Trawls N=399

Occurrence N=16

Catch N=39

0

10

20

30

40

50

60

70

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

DO (PPM)

Percent

Trawls N=341

Occurrence N=14

Catch N=37
Page 53 

0

10

20

30

40

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)

Percent

Trawls N=10537

Occurrence N=779

Catch N=2458

0

5

10

15

20

25

30

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

Bottom Depth (m)

Percent

Trawls N=12193

Occurrence N=1025

Catch N=3562

0

20

40

60

80

26 27 28 29 30 31 32 33 34 35 36 37

Salinity (PPT)

Percent

Trawls N=1959

Occurrence N=66

Catch N=96

Winter Skate

NEFSC Bottom Trawl Survey 

Spring/Adults

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 

0

5

10

15

20

25

30

2 4 6 8 10 12 14 16 18 20 22 24 26 28

Bottom Temperature (°C)

Percent

Trawls N=11844

Occurrence N=754

Catch N=3142

0

10

20

30

40

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

>500

Bottom Depth (m)

Percent

Trawls N=13543

Occurrence N=932

Catch N=4639

0

20

40

60

80

100

27 29 30 31 32 33 34 35 36 37

Salinity (PPT)

Percent

Trawls N=1956

Occurrence N=104

Catch N=228

Winter Skate

NEFSC Bottom Trawl Survey 

Fall/Adults

Figure 23. cont’d. 
Page 55 

0

5

10

15

20

25

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16

Bottom Temperature (°C)

Percent

Trawls N=2312

Occurrence N=407

Catch N=2902

0

10

20

30

40

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2385

Occurrence N=409

Catch N=2912

Winter Skate

Massachusetts Inshore Trawl Survey 

Spring/Adults 

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 

0

5

10

15

20

25

4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

Bottom Temperature (°C)

Percent

Trawls N=2067

Occurrence N=217

Catch N=1520

0

10

20

30

40

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2156

Occurrence N=221

Catch N=1548

Winter Skate

Massachusetts Inshore Trawl Survey 

Fall/Adults 

Figure 24. cont’d.
Page 57 

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. 

Gulf of Maine, Georges Bank, Southern New England, Mid-Atlantic Bight

0

2000

4000

6000

8000

10000

12000

14000

16000

18000

196019621964196619681970197219741976197819801982198419861988199019921994199619982000

Year

Landings (metric tons)

0

2

4

6

8

10

12

14

16

18

Stratified Mean Weight/Tow (kg)

Skate complex, commercial landings (mt)

Survey index of winter skate biomass

Publishing in NOAA Technical Memorandum NMFS-NE

Jon A. Gibson, Biological Sciences Editor & Laura S. Garner, Editor

Manuscript Qualification

This series represents a secondary level of scientific publishing in the National Marine Fisheries Service (NMFS). For all

issues, the series employs thorough internal scientific review, but

not necessarily external scientific review. For most issues, the

series employs rigorous technical and copy editing. Manuscripts

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be initially submitted to one of NMFS’s primary series (i.e.,

Fishery Bulletin, NOAA Technical Report NMFS, or Marine

Fisheries Review).

Identical, or fundamentally identical, manuscripts should

not be concurrently submitted to this and any other publication

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series strictly because of geographic or temporal limitations may

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Manuscripts by Northeast Fisheries Science Center (NEFSC)

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authors who are not federal employees will be required to sign a

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Send all materials and address all correspondence to:

Jon A. Gibson (Biological Sciences Editor)

NMFS Northeast Fisheries Science Center

166 Water Street

Woods Hole, MA 02543-1026 USA.

NORTHEAST FISHERIES SCIENCE CENTER

Dr. John G. Boreman, Acting Science & Research Director

Capt. John T. Moakley, Operations, Management & Information Services Chief

Teri L. Frady, Research Communications Chief
Research Communications Unit

Northeast Fisheries Science Center

National Marine Fisheries Service, NOAA

166 Water St.

Woods Hole, MA 02543-1026

Publications and Reports

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