NOAA Technical Memorandum NMFS-NE-163

Essential Fish Habitat Source Document:

Red Deepsea Crab, Chaceon

(Geryon) quinquedens,

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

January 2001
144. Essential Fish Habitat Source Document: Bluefish, Pomatomus saltatrix, Life History and Habitat

Characteristics. By Michael P. Fahay, Peter L. Berrien, Donna L. Johnson, and Wallace W. Morse. September 1999.

vi + 68 p., 34 figs., 5 tables, 1 app. NTIS Access. No. PB2000-107405.

145. Essential Fish Habitat Source Document: Butterfish, Peprilus triacanthus, Life History and Habitat

Characteristics. By Jeffrey N. Cross, Christine A. Zetlin, Peter L. Berrien, Donna L. Johnson, and Cathy McBride.

September 1999. v + 42 p., 17 figs., 4 tables. NTIS Access. No. PB2000-107404.

146. Essential Fish Habitat Source Document: Longfin Inshore Squid, Loligo pealeii, Life History and Habitat

Characteristics. By Luca M. Cargnelli, Sara J. Griesbach, Cathy McBride, Christine A. Zetlin, and Wallace W. Morse.

September 1999. v + 27 p., 12 figs., 1 table. NTIS Access. No. PB2001-100147.

147. Essential Fish Habitat Source Document: Northern Shortfin Squid, Illex illecebrosus, Life History and

Habitat Characteristics. By Luca M. Cargnelli, Sara J. Griesbach, and Christine A. Zetlin. September 1999. v + 21

p., 7 figs., 1 table. NTIS Access. No. PB2001-100146.

148. Essential Fish Habitat Source Document: Ocean Quahog, Arctica islandica, Life History and Habitat

Characteristics. By Luca M. Cargnelli, Sara J. Griesbach, David B. Packer, and Eric Weissberger. September 1999.

v + 12 p., 3 figs., 1 table. NTIS Access. No. PB2001-100145.

149. Essential Fish Habitat Source Document: Scup, Stenotomus chrysops, Life History and Habitat Characteristics.

By Frank W. Steimle, Christine A. Zetlin, Peter L. Berrien, Donna L. Johnson, and Sukwoo Chang. September 1999.

v + 39 p., 18 figs., 1 table. NTIS Access. No. PB2001-100144.

150. Essential Fish Habitat Source Document: Spiny Dogfish, Squalus acanthias, Life History and Habitat

Characteristics. By Donald G. McMillan and Wallace W. Morse. September 1999. v + 19 p., 10 figs., 3 tables. NTIS

Access. No. PB2001-100148.

151. Essential Fish Habitat Source Document: Summer Flounder, Paralichthys dentatus, Life History and Habitat

Characteristics. By David B. Packer, Sara J. Griesbach, Peter L. Berrien, Christine A. Zetlin, Donna L. Johnson, and

Wallace W. Morse. September 1999. vi + 88 p., 38 figs., 6 tables. NTIS Access. No. PB2001-100149.

152. Essential Fish Habitat Source Document: Tilefish, Lopholatilus chamaeleonticeps, Life History and Habitat

Characteristics. By Frank W. Steimle, Christine A. Zetlin, Peter L. Berrien, Donna L. Johnson, and Sukwoo Chang.

September 1999. v + 30 p., 12 figs., 2 tables. NTIS Access. No. PB2001-100143.

153. U.S. Atlantic and Gulf of Mexico Marine Mammal Stock Assessments -- 1999. By Gordon T. Waring, Debra

L. Palka, Phillip J. Clapham, Steven Swartz, Marjorie C. Rossman, Timothy V.N. Cole, Larry J. Hansen, Kathryn D.

Bisack, Keith D. Mullin, Randall S. Wells, and Neilo B. Barros. October 1999. vii + 196 p., 17 figs., 59 tables, 1 app.

NTIS Access. No. PB2000-102497.

154. Report of the Joint Scientific Review Group Workshop, April 13-14, 1999, Seattle, Washington. By Richard

L. Merrick, compiler. November 1999. v + 22 p., 3 figs., 6 tables, 5 app. NTIS Access. No. PB2000-105393.

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.

Recent Issues in This Series:
U. S. DEPARTMENT OF COMMERCE

Donald L. Evans, Secretary

National Oceanic and Atmospheric Administration

Scott B. Gudes, Acting Administrator

National Marine Fisheries Service

William T. Hogarth, Acting Assistant Administrator for Fisheries

Northeast Region

Northeast Fisheries Science Center

Woods Hole, Massachusetts

January 2001

Essential Fish Habitat Source Document:

Red Deepsea Crab, Chaceon

(Geryon) quinquedens,

Life History and Habitat Characteristics

Frank W. Steimle, Christine A. Zetlin, and Sukwoo Chang

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-163
Editorial Notes on Issues 122-152 and 163

in the

NOAA Technical Memorandum NMFS-NE Series

Editorial Production: For Issues 122-152 and 163, 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 authors and

acknowledgees of each issue, as well as those noted below in "Special Acknowledgments."

Special Acknowledgments: David B. Packer, Sara J. Griesbach, and Luca M. Cargnelli coordinated virtually all aspects of the

preprinting editorial production, as well as performed virtually all technical and copy editing, type composition, and page layout, of

Issues 122-152. David B. Packer coordinated those aspects for Issue 163. Rande R. Cross, Claire L. Steimle, and Judy D. Berrien

conducted the literature searching, citation checking, and bibliographic styling for Issues 122-152. Claire L. Steimle and Judy D.

Berrien conducted those functions for Issue 163. Joseph J. Vitaliano produced all of the food habits figures in Issues 122-152.

Internet Availability: Issues 122-152 and 163 are being copublished, i.e., both as paper copies and as web postings. All web

postings are available at: www.nefsc.nmfs.gov/nefsc/habitat/efh. Also, all web postings will be in "PDF" format.

Information Updating: By federal regulation, all information specific to Issues 122-152 and 163 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 and 163 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., Manning and

Holthius 1989e

, Cooper and Chapleau 1998f, McEachran and Dunn 1998g).

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

Manning, R.B.; Holthius, L.B. 1989. Two new genera and nine new species of geryonid crabs (Crustacea, Decapoda, Geryonidae). Proc. Biol. Soc. Wash. 102:50-77.

f

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.

g

McEachran, J.D.; Dunn, K.A. 1998. Phylogenetic analysis of skates, a morphologically conservative clade of elasmobranchs (Chondrichthyes: Rajidae). Copeia

1998(2):271-290.
Editorial Notes

Species Names: The NEFSC Editorial Office’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).

Statistical Terms: The NEFSC Editorial Office’s policy on the use of statistical terms in all technical

communications is generally to follow the International Standards Organization’s handbook of statistical methods (i.e.,

ISO 1981g

).

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.

g

ISO [International Organization for Standardization]. 1981. ISO standards handbook 3: statistical methods. 2nd ed. Geneva, Switzerland: ISO.
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

31 EFH species reports (plus one consolidated methods

report). The EFH species reports comprise 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 have

understandably 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 31

EFH source documents – available to the public through

publication in the NOAA Technical Memorandum NMFSNE series.

JEFFREY N. CROSS, CHIEF

ECOSYSTEMS PROCESSES DIVISION

NORTHEAST FISHERIES SCIENCE CENTER
Page v

Contents

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

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

Habitat Characteristics ..............................................................................................................................................................................5

Geographical Distribution .........................................................................................................................................................................7

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

Research Needs .........................................................................................................................................................................................8

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

References Cited .......................................................................................................................................................................................9

Tables

Table 1. Summary of the life history and habitat characteristics of red crab, Chaceon (Geryon) quinquedens ....................................12

Figures

Figure 1. The red crab, Chaceon (Geryon) quinquedens Smith 1879.....................................................................................................13

Figure 2 Summary of the life cycle of the red crab, Chaceon (Geryon) quinquedens [from Hastie (1995)] .........................................14

Figure 3. Distributions of adult red crabs relative to bottom temperature and depth based on spring NEFSC bottom trawl surveys.....15

Figure 4. Distributions of adult red crabs relative to bottom temperature and depth based on fall NEFSC bottom trawl surveys .........16

Figure 5. Seasonal distribution and abundance of juvenile red crabs collected during NEFSC bottom trawl surveys ...........................17

Figure 6. Seasonal distribution and abundance of small adult red crabs collected during NEFSC bottom trawl surveys.......................21

Figure 7. Seasonal distribution and abundance of harvestable-sized red crabs collected during NEFSC bottom trawl surveys.............24

Figure 8. Commercial landings for red crab in the northeast..................................................................................................................27
Page 1

INTRODUCTION

Deep-sea red crabs, Chaceon (Geryon) quinquedens

Smith 1879 (Figure 1), are contagiously distributed along

the continental shelf edge and slope of the western

Atlantic, occurring mostly between 200 and 1800 m from

Emerald Bank, Nova Scotia (and into the Gulf of Maine)

and along the continental slope of the east coast of the

U.S. into the Gulf of Mexico (Pequegnat 1970; Williams

and Wigley 1977; Elner et al. 1987). Previous reports of

the occurrence of Geryon (Chaceon) quinquedens off

West Africa and elsewhere outside of the western North

Atlantic continental shelf edge and slope (including

Bermuda, Brazil, and Argentina) were found to involve

several new geryonid species or mis-identifications

(Manning and Holthuis 1981, 1986, 1989). The species 

distribution in the depths of the Caribbean Sea, around the

West Indies Islands, and off northeastern South America

remains uncertain (R. Manning, National Museum of

Natural History, Smithsonian Institution, Washington DC,

personal communication, January 2000).

The red crab, like most deep-sea organisms, is slow

growing but reaches a maximum size of about 180 mm

carapace width (CW) and may live for 15 years or more,

although precise information on life span is lacking

(Serchuk and Wigley 1982). While catches of red crabs

have been noted since the early days of deep-sea trawling

in the 1880s (McRae 1961), only since the late 1960s and

early 1970s has there been an interest in developing a

commercial fishery for this deep-water crab (Wigley et al.

1975). The interest since the 1970s in deep-sea red crab

as a harvestable species is thought to be the result of a

decline in other coastal and offshore crab and lobster

fisheries (Gerrior 1981; Hastie 1995). A deep-sea red

crab fishery (and for its sibling species, the golden crab,

Chaceon fenneri) also developed slightly later in

southeastern U.S. and Gulf of Mexico waters (Lindberg

and Wenner 1990). To support the development of the

red crab fishery, there have been a number of biological

and technological studies that were summarized by Hastie

(1995). Compared to some other commercially harvested

deep-sea crabs, red crabs are medium sized. However, it

is primarily the crabs’ relatively long (to 60 cm each)

walking legs and attached body sections that reach the

human consumer (Wigley et al. 1975). Larger (> 114 mm

CW) male crabs have traditionally been the target of the

fishery (Serchuk and Wigley 1982). They are harvested

by trap and trawl, although most of the fishery is parttime or a by-catch from the offshore lobster or groundfish

fishery and few fishermen pursue the fishery full time. As

a result of this variable effort, annual landings have varied

by an order of magnitude and also because the fishery is

partially bycatch.

It is important to note that in the information sources

reviewed here the size of the deep-sea red crab is usually

reported as carapace width (CW); however, this

dimension was not measured consistently, and sometimes

included the posterior lateral spines [as on Northeast

Fisheries Science Center (NEFSC) surveys], or

sometimes the space between the notches forward of this

spine, or at time the carapace width limits were

unspecified. The percent contributions of the spines to

carapace width, compared to inter-notch carapace width,

for all size ranges are unknown, at present. Sometimes

carapace length is presented, which is the distance

between the rostral spine to posterior edge of carapace;

this dimension is about 80-82% of the carapace width

(Gerrior 1981).

LIFE HISTORY

A graphic summary of the life cycle of deep-sea red

crabs was developed by Hastie (1995), based on the work

of several researchers (Figure 2). In summary, the red

crab s life cycle is typical of most brachyuran crabs with

male-female coupling when the mature female molts;

fertilized eggs are brooded attached to the undersides of

the females; larvae are released into the water column for

distribution; and after several typical crab larval stages,

juveniles settle to the seabed for a benthic existence

through the rest of its relatively long life.

EGGS

Mature eggs are large and yolky for crustaceans and

range in diameter from 484 to 846 Fm, and are thought to

be the largest eggs known for crabs with planktonic

development (Haefner 1977, 1978; Hines 1982, 1988).

As for most decapod crustaceans, red crab eggs are

ovipositioned and held en mass on the female pleopods

under their abdominal flap for up to nine months until the

eggs hatch and the larvae are released into the water

column (Haefner 1978). The development of the ova was

described by Haefner (1978).

LARVAE

Larval biology for deep-sea red crab is poorly known

as few zoea and megalops have been collected or

identified in zooplankton collections. None were

identified among the 4400 brachyuran crab larvae found

in the 1977-1998 NEFSC MARMAP zooplankton

collections (J. Goulet, NMFS, NEFSC, Narragansett

Laboratory, Narragansett, RI, personal communication,

December 1999). However, Roff et al. (1986) reported

sparse collections (averaging < 10 per 500 m3

 neuston

tow) of the zoeal stage (but not megalops) from off

central Nova Scotia into the outer Gulf of Maine and over

the northeast peak of Georges Bank. No other larval

survey data is known for this species in the northwest

Atlantic.
Page 2

Much of what we know about the development of

this stage comes from eggs that were collected from

captured females and then hatched and raised in

laboratories. Larval development was observed to

consist of four zoeal stages and a final megalopa. The

zoea is typical in appearance to most brachyurans; i.e.,

with a large dorsal spine, and the megalops is also of a

typical crab form (Perkins 1973). The larval stages of this

species are relatively large in size compared to other

brachyuran crabs (Sulkin and Van Heukelem 1980).

Kelly et al. (1982) estimated that, depending upon the

water temperatures and food encountered during

development, the red crab larvae requires about 23-125

days from hatching until the megalops settles. Larvae

have been collected in surface waters in June, according

to Serchuk and Wigley (1982), but details of these

collections were unreported. Roff et al. (1986) also

reported collections of these larvae in surface waters (<

40 m) near Nova Scotia.

JUVENILES

Settling is reported at a relatively large first postmegalop instar stage, about 4 mm CW (Van Heukelem et

al. 1983). This large size at settlement may be an

adaptation to slow post-settlement growth, in that a large

size at settlement will reduce the time and number of

instars required to reach maturity (Hines 1986, 1990).

Growth of juveniles is also partially temperature

dependent, as Van Heukelem et al. (1983) reported that

juvenile crabs maintained at 9-15°C grew six time faster

than those maintained at 6°C, and at least five molts are

required to grow to about 20 mm CW.

ADULTS

Haefner (1978) and Van Heukelem et al. (1983)

suggest that the red crab requires about 18-20 molts

before it reaches its maximum size of about 180 mm CW.

Based on tagging studies, older crabs might molt

infrequently, and intermolt periods can be 6-7 yrs for

larger crabs, > 100 mm CW (Gerrior 1981; Lux et al.

1982). Molting can increase carapace size by about 7-

12% and body weight by about 33% (Serchuk and Wigley

1982). The degree of chitinoclasia (black shell disease)

and barnacle colonization on red crab carapaces (see the

Natural Mortality and Competition sections below) was

suggested as useful indicators of the length of time since a

red crab s last molt (Haefner 1978).

Van Heukelem et al. (1983) believed that red crabs

begin to recruit to the fishery at about 114 mm (CW),

which corresponds to an age of about 5-6 yrs. Van

Heukelem et al. (1983) also reported that male red crabs

reach a maximum size of about 150 mm CW and females

are slightly smaller, about 140 mm CW. Gerrior (1981),

however, reports larger maximum sizes for males, 178

mm CW, but a smaller maximum size, ~ 120 mm CW, for

females.

Farlow (1980) developed the following size-weight

regressions for red crabs:

Males: log10W = 3.09970 log10D - 0.59763 (r =

0.96717, p = 0.00001, N = 140);

Females: log10W = 2.75225 log10D - 0.34986 (r =

0.87320, p = 0.00010, N = 110);

All: log10W = 3.26202 log10D - 0.80350 (r = 0.97126,

p = 0.00001, N = 263);

where W = grams (wet weight) and D = cm (posterior

lateral spine, point-to-point carapace width). It is noted

that this width measurement (D) may be sightly larger

than the carapace widths noted in other studies which did

not include the lateral spines in their measurements; for

example, Gray (1970) measured carapace width between

the notches anterior to this spine. Farlow (1980) found

the size-weight relationship was curvi-linear for both

sexes, with weight gain per growth in width slow at

carapace widths below ~ 80 mm, but above 80 mm CW,

weight increased rapidly with carapace width expansion.

Haefner (1978) reported a similar relationship, as did

Twigg et al. (1997) for the Gulf of Mexico population,

and they also noted that the size and weight structure of

red crabs in the Gulf of Mexico was similar to the

population on the Atlantic slope.

REPRODUCTION

Haefner (1977) reports the size at maturity of red

crabs off Virginia was between 80-91 mm CW. Several

studies reported ovigerous crabs are found primarily at

sizes between 80-130 mm CW (Wigley et al. 1975;

Haefner 1977; Hines 1988), although some egg-bearing

red crabs were observed as small as 61 mm CW (Elner et

al. 1987). Lawton and Duggan (1998) suggest that males

smaller than 115 mm CW may be functionally immature.

Mating behavior is considered typical of other crabs

(Hastie 1995). The amount of time a male encases a

recently molted female can be several weeks (Elner et al.

1987). The opposite sexes probably rely on pheromones

to find each other for mating (Hines 1990), and

pheromones are useful if each sex has different preferred

depths of occurrence during non-mating periods. Egg

mass can be as much as 22% of female body weight,

which is much higher that the average 10% for almost all

other crabs (Hines 1982). Female red crab fecundity has

been estimated to range from 36 x 103

 to 226 x 103 eggs

per female, and is directly related to female body size

(Hines 1988).

Erdman et al. (1991) suggested that the egg brooding

period may be about nine months, at least for the Gulf of

Mexico population, and larvae are hatched in the early
Page 3

spring there. There is no evidence of any restricted

seasonality in spawning activity in any geographic region

of the population, although a mid-winter peak is

suggested as larval releases are reported to extend from

January to June (Wigley et al. 1975; Haefner 1978; Lux et

al. 1982; Erdman et al. 1991; Biesiot and Perry 1995).

Laboratory studies also found hatching to occur from

April to June (Perkins 1973). Gerrior (1981), however,

suggested that red crab egg hatching occurred later,

between July and October, based on the ratio of eggbearing to non-egg-bearing crabs.

Because of the long intermolt period for adult

females (5-7 yrs), and the assumption that like most other

brachyurans fertilization only occurs at molting, it has

been speculated that red crabs may not spawn annually,

although it is possible that sperm could be stored for

intermolt spawning efforts; thus there can be annual

brooding within the population, although not for every

mature individual (Hines 1982; Lux et al. 1982: Erdman

et al. 1991; Biesiot and Perry 1995). Hines (1990) reports

that there is no evidence of senility or any reproduction

capacity decline at the largest sizes for the species.

However, Lux et al. (1982) and Lawton and Duggan

(1998) suggest that there is evidence for a terminal molt;

this may restrict any significant increases in fecundity

with age.

FOOD HABITS

No information is known on the natural diets of red

crab larvae, but it is probably zooplanktivorous, as they

were found to thrive on rotifers, brine shrimp, and

chopped mollusk meats in laboratory cultures (Perkins

1973; Sulkin and Van Heukelem 1980; Van Heukelem et

al. 1983). Sulkin and Van Heukelem (1980) believe that

the ability of red crab larvae to develop with reduced

growth on a relatively poor diet (i.e., rotifers) suggests an

adaptation to patchy and often poor food resources in

slope waters. This can also allow the larvae considerable

flexibility in prey choice.

Farlow (1980) reported that red crabs are

opportunistic feeders, a characteristic of many deep-sea

organisms. He reported that post-larval, benthic red crabs

ate a wide variety of infaunal and epifaunal benthic

invertebrates that they find in the silty sediment or pick

off the seabed surface (Gerrior 1981). This observation is

supported by submersible observations of red crabs being

commonly found in shallow pits and excavations in softer

sediments, and by infaunal bivalve mollusks being found

in red crab stomachs (Valentine et al. 1980). Farlow

(1980) also reported that smaller red crabs ate sponges,

hydroids, gastropod and scaphopod mollusks, small

polychaetes and crustaceans, and possibly tunicates.

Larger crabs ate similar small benthic fauna and larger

prey, such as demersal and mid-water fish (Nezumia and

myctophids were identified in red crab guts), squid, and

the relatively large, epibenthic, quill worm (Hyalinoecia

artifex). Farlow s (1980) finding of sponges, hydroids,

and tunicates in the diets suggests that epifauna attached

to solid surfaces are also preyed upon; such epifauna is

most common within the deeper rocky areas of the Gulf

of Maine and within submarine canyons. Gray (1970)

reported that red crabs held in aquaria ate anemones by

hovering over an anemone until it extended its tentacles,

then it reached down with its chelipeds and gently pulled

off and ate one tentacle at a time until the anemone was

bare of tentacles. Farlow (1980) considered the red crab

the dominant large predator of the biomass on the upper

slope.

They can also scavenge deadfalls (e.g., trawl

discards) of fish and squid, as they are readily caught in

traps with these as bait (Gerrior 1981) and eat them when

held in aquaria (Gray 1970). Olfaction appears to be

important in locating foodfalls as they are reported to be

attracted to bait within a several km radius (Diehl and

Biesiot 1994). They also probably resort to sediment

deposit feeding at times and consume anything else they

might encounter that can provide sustenance, including

man-made artifacts; e.g., wood chips, pieces of yellow

rubber and glass tubing have been found in the stomachs

of some red crabs (Farlow 1980). American lobsters are

also known to eat such man-made artifacts in coastal

waters (Steimle 1994).

Elner et al. (1987) reports that unlike other crabs, red

crabs do not stop feeding during mating; this is possibly

an adaptation to an uncertain food supply and an extended

male-female coupling period. Farlow (1980) estimated

that $average-sized# red crabs fed in laboratory aquaria ate

an average of 0.32 g wet wt of prey per day.

NATURAL MORTALITY

There are few records of predation on red crabs and

these are by several species of gadid fishes. Sedberry and

Musick (1978) reported that the longfin hake (Urophycis

chesteri) ate red crab as a minor prey. A search of the

1973-1990 NEFSC food habits data base (Reid et al.

1999) and shark prey data base found only a few instances

of red crab identified as being eaten by, for example, cod

(Gadus morhua) and red hake (Urophycis chuss), and

there was no evidence of predation by larger sharks (W.

Pratt, NMFS, NEFSC, Narragansett Laboratory,

Narragansett, RI, personal communication, December,

1999). This lack of predation is unexpected as they do

not have any obviously strong defensive mechanisms

(they are non-burrowing, relatively slow moving, and

weakly spined) and, as prey, they have a food energy

value that is mid-range for deep-sea benthic crustaceans

(Steimle and Terranova 1988). Farlow (1980) reports

fresh red crab shell remains in the stomachs of larger

crabs, suggesting a possibility of cannibalism; he

discounted the possibility that these remains were
Page 4

remnants of molted shells. Gray (1970) noted that red

crabs tend to autonomize any legs that are damaged; this

can also be a source of any red crab fragments found in

stomachs.

The pathology of deep-sea red crabs is also poorly

known. Red crabs have been reported often with dark

lesions, shell disease, or chitinoclasia on their carapace

(Hines 1990; Young 1991). These lesions are thought to

be caused by bacteria that are found attached or adhering

to the crab s carapace; several bacterial taxa have been

isolated from the infected lesions (Bullis et al. 1988). An

association of the lesions with a former deep-water

sewage sludge disposal activity was tentatively implicated

as a cause of these lesions, although there is insufficient

evidence to fully support such an association because a

relatively high incidence of shell disease was also

identified on specimens collected and preserved during

the late nineteenth century, well prior to any substantial

offshore waste disposals (Young 1991; Hastie 1995).

Larger crabs are reported to have more incidences of

lesions, probably because of a longer intermolt period,

and the lesions are thought to disappear with molting

(Young 1991). Diehl and Biesiot (1994) report that there

is a notable relationship between the degree of multi locus

heterozygosity (genetic variability) and shell disease in

individuals of this species that is $novel#. The effect of

the lesions on the health of red crabs is poorly known,

although the lesions can affect marketability, if the crabs

are sold whole. Red crabs do not appear to be susceptible

to gaffkemia, or red tail disease, that occurs in the

American lobster (Cornick and Stewart 1975).

The relatively long life span of red crabs potentially

exposes them over an extended period of time to the

accumulation of contaminants that become available

offshore; e.g., from atmospheric deposition, currents, or

waste disposal. Examinations of red crab tissues,

however, have shown that levels of certain toxic metals

are similar, but no higher, to that found for short-lived,

coastal species (Greig et al. 1976).

COMPETITION

Although observations on competitive interactions of

red crabs with other species are poorly known, Hines

(1990) suggested that the strong-clawed Jonah crab,

Cancer borealis, and the American lobster, Homarus

americanus, may be competing with red crabs on the

continental shelf edge, although the nature of this

competition (food or habitat) has not been observed.

Hagfish (Myxine spp.) and other scavengers can compete

for deadfalls (Lockhart et al. 1990). A variety of

benthivorous demersal fish that occur on the upper slope,

such as those commonly found in red crab or deep-sea

lobster traps, including cusk (Brosme brosme) (Stone and

Bailey 1980), or observed by divers where red crabs

occur, such as eelfish (Aldrovandia affinis), rattails

(Coryphanoides sp.), white hake (Urophycis tenuis), and

galatheid crabs (Munida sp.) (Whitlatch et al. 1990), can

compete with red crabs for benthic prey. Near and south

of Cape Hatteras, a larger geryonid crab – the North

American golden crab, Chaceon fenneri, is also found on

the upper slope and could be competing with C.

quinquedens for some habitat space or other resources if

the two species populations overlap in habitat use

(Lockhart et al. 1990; Hastie 1995).

Although not strictly a competitive relationship, a

number of macroscopic organisms are reported to live

epizoically on the surfaces of the red crab shell; these

commonly include some species of stalked or $goose#

barnacles, which are filter-feeders. A commensal

polychaete worm, Dorveilla geryonicola, has been

reported to inhabit the gill chamber of red crabs, but its

source of food and effect on the crab is unknown (Gaston

and Benner 1981).

MIGRATION

Tagging studies off southern New England by Ganz

and Herrmann (175) and Lux et al. (1982) suggest adult

red crabs move both up and down the continental slope,

covering a range of about 500 m in depth, perhaps as part

their spawning cycle. The adults were also found to move

laterally almost 100 km along the slope, but most tag

returns were from areas within 20 km of their release

location. Movements of the tagged crabs within the ~

200-1000 m depth zone, between Hudson and Block

Canyons, seem to be generally without any predominant

direction. Juvenile red crabs, which are thought to settle

from their larval stage to the seabed in deeper mid-slope

waters, are reported to move gradually upslope into

shallower waters with growth (Wigley et al. 1975), but

this may not be a true migration.

STOCK STRUCTURE

Wigley et al. (1975) reported a bimodal depth

distribution in male and female red crab size frequency

distributions south of Georges Bank, although Farlow

(1980) and Gerrior (1981) found less distinct segregation

of sexes in their studies. Lockhart et al. (1990) suggested

that the habitat use and distribution of the Gulf of Mexico

population of red crabs seems different from what has

been reported for the Middle Atlantic Bight population.

This suggests that there can be physiological tolerance

differences between these two populations. However,

Diehl and Biesiot (1994) reported that red crabs have a

high degree of genetic variability, more so than other

deep-sea crustaceans, which can mask evidence of a

genetic separation trend.
Page 5

HABITAT CHARACTERISTICS

The larvae of this species are pelagic and occur in

warmer and lower salinity surface waters above and

beyond the continental slope habitat frequented by adult

females. Post-larval red crabs are primarily inhabitants of

the silty seabed of the deep cold water on the outer

continental shelf and mid to upper continental slope of the

western North Atlantic, south of the Gulf of Saint

Lawrence (Canada), into the partially rocky Gulf of

Maine, and along the continental shelf edge and slope into

the Gulf of Mexico, and possibly the western South

Atlantic. These crabs are considered part of an

assemblage of deep-water crustaceans that inhabit the mid

to upper continental slope of the northwest Atlantic, and

this assemblage includes a number of smaller shrimp and

crabs (Wenner and Boesch 1979). Salinities on the upper

slope where benthic red crabs occur tend to be stable and

oceanic at about 35-36 ppt (Schmitz et al. 1987). The

thermal regime can be more variable, ~ 4-10°C, and

include the temporary warming effects of the passage of

an inshore loop or gyre of the Gulf Stream along the

upper slope and shelf edge.

EGGS

As for most decapod crustacea, the eggs of the red

crab are attached en mass to pleopods under the

abdominal flap of females until the eggs hatch and the

larvae are released into the water column (Haefner 1978).

Egg-bearing females are most commonly found on the

shallow continental slope, between about 200-400 m,

where temperatures are typically between 4-10°C (Wigley

et al. 1975; Kelly et al. 1982; Lindberg et al. 1990). Gray

(1970) suggested that egg-bearing females are common

within submarine canyons off southern New England.

LARVAE

Sulkin et al. (1980) reported that early larval stages

show positive phototaxis down to light intensities of 2 x

10-2 W/m2

. At lower intensities the majority of the larvae

were indifferent or negatively phototaxic. The larvae

were also found to be able to swim through relatively

sharp thermal gradients; e.g., 10°C. Kelly et al. (1982)

believed that hatched zoea are negatively geotactic and

positively phototactic; i.e., they tend to move upward in

the water column, and somehow swim hundreds of meters

to surface waters. During this ascent, they are thus

exposed to changing thermal and prey and predator

availability conditions. Roff et al. (1986) reported that

the greatest collections of decapod crustacean larvae in

Gulf of Maine and Nova Scotian waters, including that of

the red crab, occurred in neuston samples from surface

waters, < 40 m. Roff et al. also reported that the red crab

larvae were collected in relatively low (averaging less

than 10 per neuston tow) concentrations where surface

salinities ranged between 28.9 and 33.0 ppt, and

temperatures were between 6.0° and 19.5°C. Kelly et al.

(1982) reported that red crab larvae can survive

temperatures as high as 25°C (as found at the summer sea

surface and within parts of the Gulf Stream water mass).

Red crab larvae were also collected between 12 and

270 km offshore in the Nova Scotian-Georges Bank area

and, based on a factorial analysis of crustacean larval

abundance and distribution collection data, were not

strongly associated with concentrations of any other larval

crustaceans (Roff et al. 1986).

Red crab zoeal swimming activity slows down as

they approach metamorphosis into the megalops or first

crab instar stage, which serves to let the megalopa

gravitationally (or actively?) descend to the seabed (Kelly

et al. 1982).

JUVENILES

The benthic settlement of the first $crab# stage of red

crabs is reported to occur in mid-slope water (~ 1000 m),

but the growing juveniles may gradually move upslope to

warmer, upper slope waters (Wigley et al. 1975). Wigley

et al. (1975) reported that $young# red crabs were only

collected in cold water, 4.4-5.5°C, which is

characteristically found below the ~ 500 m lower limit of

the shelf break $warm band# inhabited by tilefish,

Lopholatilus chamaeleonticeps, and associated species

(Steimle et al. 1999). This upslope movement can expose

the juveniles to warmer water, and juvenile red crab

growth rates were reported to increase as they are

gradually exposed to warmer waters in the laboratory

(Van Heukelem et al. 1983); this can be a survival

enhancement adaptation. Availability of benthic prey

may also be a factor in the upslope movement (if it is

real), because the mean biomass of benthic infauna

increases from 1000 m to 100 m in the Middle Atlantic

Bight and New England (Wigley and Theroux 1981;

Theroux and Wigley 1998). The juveniles were collected

or observed on smooth or dimpled surfaced, silt-clay

sediments (Auster et al. 1991), but might occur on other

sediment types, especially in the heterogeneous sediment

types of the Gulf of Maine and submarine canyons. Gray

(1970) reported that the results of his trawl surveys

suggested that submarine canyons off southern New

England held concentrations of small crabs of both sexes.

The NEFSC collections of juvenile red crab from the

bottom trawl surveys that are accompanied by

hydrographic data are too few to summarize and draw any

conclusions relative to their depth and temperature

preferences and thus are not presented.

ADULTS

Like juveniles, adult red crabs are reported associated

with smooth or dimpled, unconsolidated and consolidated
Page 6

silt-clay sediments (Haefner and Musick 1975; Wigley et

al. 1975; Valentine et al. 1980; Lindberg et al. 1990),

although Schroeder (1959) reported that there were good

catches from both hard and soft bottom grounds.

However, Herrmann (1974) reported that pot (trap)

catches on hard bottoms were small compared to pot

catches on softer sediments. Haefner and Musick (1974)

reported that catches were lower within Norfolk

Submarine Canyon (off Virginia) than on the slope

adjacent to the canyon; their report is consistent with

Gray (1970) who also noted this inter-canyon habitat

preference for adults (especially males) off southern New

England. However, Valentine et al. (1980) reported that

red crabs were common in Oceanographer Canyon on the

southern edge of Georges Bank. Auster et al. (1991)

reported submersible observations showing that red crab

seem to have a completely random distribution across the

seabed, at a depth of about 700 m, without any apparent

aggregation nodes.

The upper thermal limits for the benthic red crab

have been reported to be about 10-12°C (Haefner 1978),

which is also typical of the shelf edge $warm band#

favored by tilefish (Steimle et al. 1999), but red crabs

were reported to exhibit thermal stress at temperatures

above 10°C (Gray 1970; Serchuk and Wigley 1982). The

NEFSC spring and fall bottom trawl survey data for small

and large (i.e., harvestable-sized) adult red crabs collected

at depths less than 400 m shows that, in the shallow edge

of their known depth distribution, both size groups were

most commonly collected at bottom temperatures of about

11°C in during the February-March spring surveys

(Figure 3) and at slightly cooler areas of about 7-9°C

during the September-November fall surveys (Figure 4).

Both size groups were mostly collected at depths greater

than 300 m in both seasons. They were also collected in

minor quantities at a greater range of temperatures (3-

14°C) and at depths as shallow as 80 m.

The upper depth limits of red crab occurrence along

the slope overlaps the deep-sea oxygen minimum zone,

which occurs at about 200-400 m along the slope; this

zone is where dissolved oxygen concentrations (DO) can

be as low as 3 ml/l, while DO concentrations above or

below this minimum zone are usually about 5-7 ml/l

(Flagg 1987; Schmitz 1987). The ability of red crab to

tolerate hypoxic conditions reported by Henry et al.

(1990a) and Erdman et al. (1991) seems an adaptation for

adults to transcend or inhabit the upper slope oxygen

minimum depth zone. Laboratory studies have found that

red crabs are intolerant of salinities below 20 ppt (Henry

et al. 1990b); however, these low salinity levels are

unlikely to occur in the depths of the edge of the

continental shelf.

Hastie (1995) believes that several submersible

studies (e.g., Valentine et al. 1980; Whitlatch et al. 1990)

suggest that red crabs can be locally important

bioturbators of surface sediments (and associated infaunal

communities) by their sediment digging and benthic

foraging activities. This bioturbation was speculated to

play an important role in maintaining deep-sea

biodiversity.

Although red crabs are most often found in outer

continental shelf and slope depths > 100 m, they are also

reported at depths as shallow as 40 m in the Gulf of

Maine (Lux et al. 1982). Red crabs have not been

reported to be a common member of the tilefish pueblo

village, burrow, or canyon community, although red crab

and tilefish depth distribution zones overlap (Grimes et al.

1987; Cooper et al. 1987). Red crabs, however, can be

common in some canyon systems, such as Oceanographer

Canyon and others along southern New England (Gray

1970; Valentine et al. 1980).

Haedrich et al. (1980) found red crab the second most

abundant megafaunal species in the 653-1290 m depth

zone off southern New England, after the northern

cutthroat eel, Synaphhobranchus kaupi. Hecker (1990)

also included three other fish species as representative

members of the megafaunal assemblage on the mid- to

upper slope off southern New England: a grenadier

(Nezumia spp.), the longfin hake (Phycis chesteri), and

the witch flounder (Glytocephalus cynoglossus). To this

assemblage that includes red crab, Farlow (1980) added

offshore hake (Merluccius albidus) and the wolf eelpout

(Lycenchelys verrilli).

Sexual separation in habitat use has been reported,

with females being more abundant in shallower and

warmer slope waters (Schroeder 1959; Haefner and

Musick 1974; Wigley et al. 1975; Haefner 1978; Stone

and Bailey 1980; Lux et al. 1982; Lindberg et al. 1990).

Although the portion of the NEFSC bottom trawl survey

results for red crab that includes sexual differentiation and

temperature data is small (181 samples between 1963-

1999), it suggests that within the relatively shallow, <400

m survey depth range, females were more commonly

collected on about a 2:1 ratio than males, and females

were slightly more common in the fall than in the spring.

Because of the limited depth scope and sample size of the

survey results, these tentative conclusions can be biased.

Known large scale disturbances on the continental

slope can affect the red crab population, as well as most

other benthic species that live there. One such

disturbance is that in the past areas of the continental

slope with soft sediments have become dislodged and

slumped into deeper waters (Tucholke 1987). This can

undoubtedly have an effect on biological populations in

and near the slump zone. Another such disturbance was

that which caused the mass mortlities of tilefish and

associated organisms in 1880 (Steimle et al. 1999).

Although the actual cause of the massive tilefish mortality

is not known, the oceanographic or geological factors

involved (assuming either of these was involved) in that

event, which basically decimated the tilefish population

and most other organisms associated with the warm band

in the Middle Atlantic Bight for over a decade, could

reoccur and also have an effect on the red crab

population. A recent analysis suggests that anomalously

cold water was in the area of the 1880s tilefish kill and
Page 7

probably responsible for the mortality event (Marsh et al.

1999).

GEOGRAPHICAL DISTRIBUTION

Post-larval red crabs are inhabitants of the seabed of

the mid to upper continental slope of the western North

Atlantic, Gulf of Mexico and possibly the northwestern

South Atlantic, and the larvae inhabit the water column

generally above and near the adult distributions. The

NEFSC bottom trawl surveys collect small quantities of

benthic red crabs, but these surveys are typically

restricted to depths less than 400 m on the upper slope,

although they occasionally trawl in deeper canyons (Reid

et al. 1999). Fishery independent information of red crab

distributions below this NEFSC bottom trawl survey

depth limit are only available from infrequent, special

surveys, such as those reported by McRae (1961),

Haefner and Musick (1974), or Wigley et al. (1975).

EGGS

Egg-bearing females are generally found distributed

on shallow slope waters of less than 400 m (Kelly et al.

1982) and from Nova Scotia to at least the Gulf of

Mexico. As for most decapod crustacea, eggs are held en

mass on the pleopods under the female s abdominal flap

until the eggs hatch and the larvae are released into the

water column (Haefner 1978).

LARVAE

Larvae have been collected from off Nova Scotia

(Roff et al. 1986), but have not yet reported from other

areas where adults occur, including the Gulf of Mexico.

In the northwest Atlantic, Kelly et al. (1982) suggested

that because the early red crab larval stages seek surface

waters and grow faster in warm water, the movement of

the Gulf Stream [and its meanders and gyres] south of

Georges Bank can have a role in larval dispersal. Larvae

can tolerate water temperatures as high as 25°C, as would

be found in Gulf Stream or South Atlantic Bight/Gulf of

Mexico surface waters, as well as < 10°C water

temperatures found on the continental slope seabed. With

offshore Nova Scotia being the northern boundary of

where they are known to be found, if the Gulf Stream is a

significant red crab larvae dispersal factor, it is possible

that a red crab larval source from as far south as the South

Atlantic Bight can be important to juvenile recruitment in

the New England area. However, there is general

southerly shelf water movement inshore of the Gulf

Stream affected area, and occasional southward moving

Gulf Stream gyres or eddies can distribute the larvae from

southern New England and Georges Bank spawning

populations south to near Cape Hatteras, too (Kelly et al.

1982). Kelly et al. (1982) also estimated that the range of

larval dispersal can vary with their position in the water

column and season, and dispersal might range from about

150 to almost 1000 km from the hatching area. Kelly et

al. (1982) also presented a larval dispersal model that

suggested that recruitment to southern New England red

crab stocks might be variable, while recruitment to the

stocks off Virginia might be more predictable.

JUVENILES

Juvenile red crabs were reported or presumed to have

the same latitudinal distribution as adults; i.e., occurring

from Nova Scotia to at least the Gulf of Mexico.

However, their distribution along the continental slope

depth gradient within this range may differ from that of

adults. Wigley et al. (1975) and Haefner (1977) believed

that metamorphosing megalopa settle to the seabed at

mid-slope depths (~ 1000 m) as first instar juveniles and

move upslope gradually as they grow. This pattern of

settlement and gradual redistribution was supported by an

inverse relationship between body size and depth reported

by Wigley et al. (1975). They also report that only small

specimens are usually collected in deep water, 740-1051

m, off southern New England. On the Scotian shelf,

however, a wider range of sizes were found at the 720-

900 m depths (Stone and Bailey 1980), and no clear

relationship between red crab individual weight and depth

was found by Farlow (1980). McRae (1961) noted that

most of the large numbers of red crabs collected in a

1955-1957 survey of the Gulf of Maine (i.e., generally at

depths of less than 300 m) were $small#; but this shallow

depth distribution of presumed juvenile red crabs could be

also the result of an early 1950s strong year class using

most acceptable habitats, or an unusual pattern of settling

larvae.

In the ~ 400 m depth-limited 1964-1999 NEFSC

bottom trawl surveys, juvenile red crabs (equal or less

than 7 cm CW) were collected in small quantities (less

than 6 per tow) mostly within the western Gulf of Maine

and even less were collected on the continental shelf edge

between southern Georges Bank and near Toms Canyon

(Figure 5). Because of the depth limitations of the

NEFSC surveys, their assessments of the distribution of

red crabs are incomplete, and juveniles (and adults) that

are known to occur deeper on the continental slope were

not surveyed.

ADULTS

In the Northeast, adult red crabs occur along the

continental shelf edge and upper slope from the Scotian

Shelf and the Gulf of Maine to Cape Hatteras (Serchuk

and Wigley 1982). The species also occurs south of Cape
Page 8

Hatteras into at least the Gulf of Mexico at similar depths.

For an examination of the 400 m depth-limited 1964-1999

NEFSC bottom trawl survey data, the adult red crabs were

segregated into two groups for analysis: below

harvestable-size (small) adults (8-11 cm CW; Figure 6)

and large adults at or above the commonly used

harvestable size of 11 cm CW (Figure 7). The small

adults were not collected during the winter bottom trawl

surveys and only a few were collected in the summer

surveys within and around the perimeter of the Gulf of

Maine, but during the spring and fall surveys, they were

collected in minor to moderate numbers both within the

western Gulf of Maine and along the outer continental

shelf between southern Georges Bank and Norfolk

Canyon (Figure 6). The larger, harvestable adults were

collected in a similar pattern, although fewer of the larger

crabs were collected within the Gulf of Maine during the

spring and fall trawl surveys (Figure 7).

As previously noted, there may be an inverse

relationship between body size and depth (Wigley et al.

1975). Stone and Bailey (1980) reported that large crabs

were only collected at 180-360 m depths on the Scotian

Shelf. In the Gulf of Mexico, red crabs were not

commonly collected above the 677 m depth zone;

temperature or bottom sediment type could be prime

factors controlling their distribution in the Gulf of Mexico

(Lindberg et al. 1990; Lockhart et al. 1990).

From around Cape Hatteras into the Gulf of Mexico,

red crabs may partially overlap the distribution of a larger

sibling species, the golden crab (C. fenneri, previously

called Geryon affinis), that was noted to occur

uncommonly off southern New England, too (Wigley et

al. 1975).

STATUS OF THE STOCKS

Although occasional surveys of the relative

abundance and distribution of this species have been

conducted since the 1950s in some areas (Schroeder 1959;

McRae 1961; Haefner and Musick 1974; Wigley et al.

1975; Haefner 1978; Gerrior 1981; Stone and Bailey

1980; McElman and Elner 1982) there have not been any

broad-scale, long-term, fishery-independent surveys for

this species to document any trends in relative population

abundance and distribution (F. Almeida, NMFS, NEFSC,

Woods Hole Laboratory, Woods Hole, MA, personal

communication, December 1999). Broad-scale pot

surveys have not been done, either, although pots may be

better collectors of red crabs than trawls (P. Gerrior,

NMFS, NEFSC, Woods Hole Laboratory, Woods Hole,

MA, personal communication, August 2000). Hastie

(1995) summarized a number of previous stock

assessments and density estimates for the species to the

early 1990s, but these also included information from

south of Cape Hatteras. Annual northeast landings were

reported since the 1970s but the data were not

comprehensive until the early 1980s (C. Yustin, NMFS,

NERO, Gloucester, MA, personal communication,

January 2000). Landings between 1982 and 1998

averaged overall about 1800 MT, but reported landings

annually varied between a range of 0.25 and 3800 MT.

Peak harvests were reported in 1984 and 1988, and an

almost insignificant harvest was reported in 1994 (Figure

8); CPUE data are not available.

The slow growth and maturation rates of red crabs,

and possible infrequent recruitment, suggests that the high

yields of the 1970s-1980s will likely be unsustainable

(Hastie 1995).

RESEARCH NEEDS

It is obvious more needs to be known about the

distribution of the species at all life stages, the variability

and trends in population abundance and dynamics, and a

way to adequately sample larval stages. The studies on

red crabs summarized above include some that present

differing results or conclusions about the biology or

habitat use of this species; e.g., do megalops settle in

deeper water and move upslope with growth and are sexes

distributed separately (Hines 1990)? These areas of

conflicting information need to be resolved by further

research or data analysis. Further research or data is also

needed on: age-size relationships, lengths of intermolt

periods for all size classes, mortality rates for various size

classes (especially for discards), yield per recruit, upslope

and lateral migrations (including for spawning or related

to size class), Middle Atlantic Bight larval dispersal

patterns and the testing of larval dispersal models, genetic

comparison of the Northwest Atlantic and Florida/ Gulf of

Mexico populations, and where and when females

primarily spawn and the possibility of sperm storage by

intermolt females (Kelly et al. 1982; Armstrong 1990;

Lindberg and Wenner 1990; Lockhart et al. 1990; Hastie

1995). In the northeast the use of submarine canyons as

preferred spawning and juvenile red crab nursery areas, as

suggested by Gray (1970), needs to be re-examined and

verified for the potential of this habitat being critical to

the life history of the species. There is also a need to find

a cost-effective approach to conducting stock assessment

surveys of this and other harvestable deep-water species.

Beside these research needs, there is a need to

standardize size measurements on red crabs, such as

carapace widths, which should not include damageable or

erodible rostral or lateral spines in the measurements, as

was done in some studies. The carapace width between

the notches forward of the outermost spine would provide

a more consistent and reliable measure of size, and

because the shell is relatively soft and can be bent with

pressure, the measuring caliphers should just touch the

shell (Gray 1970).
Page 9

ACKNOWLEDGMENTS

Assistance with obtaining literature sources for this

review was provided by Claire Steimle and Judy Berrien.

Assistance with searching databases and retrieving data in

tabular or graphical form was also provided by Joe

Vitaliano, John Witzig, Charles Yustin, Annette Kalbach,

and Frank Almeida. Useful comments or information

were provided by Pat Gerrior and Fred Serchuk.

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Page 12

Table 1. Summary of the life history and habitat characteristics of red crab, Chaceon (Geryon) quinquedens. (CW=

carapace width.)

Life Stage: Eggs Larvae Juveniles Adults Spawning

Adults

Time of Year

Fall-spring. January-June. Throughout. Throughout. Fall-spring.

Size and

Growth

Fecundity: up to

~ 226,000

eggs/female.

4 instar stages,

then megalops

stage settles to

seabed.

4-70 mm CW;

molting period

extends with

size.

80-180 mm

CW molts

every 6-7 yrs;

CW increases 7-

12% per molt.

Egg-bearing

from ~ 70 mm

to 140 mm CW.

Geographic

Location

Nova ScotiaGulf of Mexico.Nova ScotiaGulf of Mexico.Nova ScotiaGulf of Mexico.Nova ScotiaGulf of Mexico.Nova ScotiaGulf of Mexico.

Habitat

Attached to

female abdomen

until hatching.

Water column,

outer shelf and

slope.

Seabed, 700-

1800 m.

Seabed, 200-

1300 m.

Seabed, upper

slope.

Substrate

Attached to

female abdomen

until hatching.

Water column,

surface to

bottom.

Smooth-surface

silt-clay.

Silty to hard

sediments.

Silt-clay.

Temperature

~ 4-12°C. ~ 4-25°C. ~ 4-10°C. ~ 5-14°C. ~ 4-12°C

Salinity

~ 35 ppt. 29-36 ppt. ~ 35 ppt. ~ 35 ppt. ~ 35 ppt.

Dissolved

Oxygen

~ 3-8 ml/l. ~ 5-8 ml/l. ~ 3-7 ml/l. ~ 3-8 ml/l. ~ 3-8 ml/l.

Prey

Yolk. Probably

planktivorous.

Small benthic

invertebrates

and detritus.

Benthic

organisms and

detritus, carrion.

Benthic

invertebrates

and detritus.

Predators

Unreported. Unreported. Longfin hake,

other gadids.

Same as for

juveniles (?)

Unreported.

Notes

Attached to

female abdomen

until hatching.

Early stages rise

to surface; later

stages gradually

settle to the

seabed.

May move

upslope with

growth.

Movements are

generally

limited to < 50

km.

Occurs when

female molts,

inter-molt sperm

storage possible.
Page 13

Figure 1. The red crab, Chaceon (Geryon) quinquedens Smith 1879.
Page 14

Figure 2. Summary of the life cycle of the red crab, Chaceon (Geryon) quinquedens [from Hastie (1995), used with

permission].
Page 15

Figure 3. Distributions of small and harvestable-sized (large) adult red crabs relative to bottom water temperature and

depth based on spring NEFSC bottom trawl surveys (1968-1999; all years combined). White bars give the distribution

of all the stations, and black bars represent, within each interval, the percentage of the total number of red crab caught.

0

10

20

30

40

50

60

70

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

Temperature (0C)

Percent

Stations=53

Catches=418

0

10

20

30

40

50

60

70

80 120 160 200 240 280 320 360 400 440

Depth (m)

Percent

Stations=60

Catches=441

0

10

20

30

40

50

60

70

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

Temperature (0C)

Percent

Stations=26

Catches=204

0

10

20

30

40

50

60

80 120 160 200 240 280 320 360 400 440

Depth (m)

Percent

Stations=35

Catches=227

Small Adult

Small Adult

Large Adult

Large Adult
Page 16

Figure 4. Distributions of small and harvestable-sized (large) adult red crabs relative to bottom water temperature and

depth based on fall NEFSC bottom trawl surveys (1963-1999; all years combined). White bars give the distribution of

all the stations, and black bars represent, within each interval, the percentage of the total number of red crab caught.

0

10

20

30

40

50

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

Temperature (0C)

Percent

Stations=43

Catches=181

0

10

20

30

40

80 120 160 200 240 280 320 360 400 440

Depth (m)

Percent

Stations=51

Catches=255

0

10

20

30

40

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

Temperature (0C)

Percent

Stations=40

Catches=170

0

10

20

30

40

80 120 160 200 240 280 320 360 400 440

Depth (m)

Percent

Stations=43

Catches=186

Small Adult

Small Adult

Large Adult

Large Adult
Page 17

Figure 5. Seasonal distribution and abundance of juvenile red crabs (< 7 cm CW) collected during NEFSC bottom trawl

surveys (1963-1999, all years combined).

RED DEEPSEA CRAB – Juveniles (< 7 cm CW)

Winter (1964 – 1999)

NEFSC Bottom Trawl Surveys

Crabs caught/station (excluding null stations):

Mean: 5 Min: 5 Max: 5

Length (cm):

Mean: 6 Min: 5 Max: 7
Page 18

Figure 5. cont’d.

RED DEEPSEA CRAB – Juveniles (< 7 cm CW)

Spring (1968 – 1999)

NEFSC Bottom Trawl Surveys

Crabs caught/station (excluding null stations):

Mean: 2 Min: 1 Max: 4

Length (cm):

Mean: 6 Min: 4 Max: 7
Page 19

Figure 5. cont’d.

RED DEEPSEA CRAB – Juveniles (< 7 cm CW)

Summer (1964 – 1995)

NEFSC Bottom Trawl Surveys

Crabs caught/station (excluding null stations):

Mean: 2 Min: 2 Max: 2

Length (cm):

Mean: 7 Min: 6 Max: 7
Page 20

 Figure 5. cont’d.

RED DEEPSEA CRAB – Juveniles (< 7 cm CW)

Fall (1963 – 1999)

NEFSC Bottom Trawl Surveys

Crabs caught/station (excluding null stations):

Mean: 2 Min: 1 Max: 4

Length (cm):

Mean: 6 Min: 1 Max: 7
Page 21

Figure 6. Seasonal distribution and abundance of small adult red crabs (8-11 cm CW) collected during NEFSC bottom

trawl surveys (1963-1999, all years combined; none were collected in winter).

RED DEEPSEA CRAB – Early Adults (8-11 cm CW)

Spring (1968 – 1999)

NEFSC Bottom Trawl Surveys

Crabs caught/station (excluding null stations):

Mean: 7 Min: 1 Max: 264

Length (cm):

Mean: 10 Min: 8 Max: 11
Page 22

Figure 6. cont’d.

RED DEEPSEA CRAB – Early Adults (8-11 cm CW)

Summer (1964 – 1995)

NEFSC Bottom Trawl Surveys

Crabs caught/station (excluding null stations):

Mean: 1 Min: 1 Max: 2

Length (cm):

Mean: 10 Min: 8 Max: 11
Page 23

Figure 6. cont’d.

RED DEEPSEA CRAB – Early Adults (8-11 cm CW)

Fall (1963 – 1999)

NEFSC Bottom Trawl Surveys

Crabs caught/station (excluding null stations):

Mean: 5 Min: 1 Max: 47

Length (cm):

Mean: 10 Min: 8 Max: 11
Page 24

Figure 7. Seasonal distribution and abundance of harvestable-sized red crabs (> 11 cm CW) collected during NEFSC

bottom trawl surveys (1963-1999, all years combined; none were collected in winter).

RED DEEPSEA CRAB –

Harvestable Adults (> 11 cm CW)

Spring (1968 – 1999)

NEFSC Bottom Trawl Surveys

Crabs caught/station (excluding null stations):

Mean: 6 Min: 1 Max: 90

Length (cm):

Mean: 13 Min: 12 Max: 21
Page 25

Figure 7. cont’d.

RED DEEPSEA CRAB –

Harvestable Adults (> 11 cm CW)

Summer (1964 – 1995)

NEFSC Bottom Trawl Surveys

Crabs caught/station (excluding null stations):

Mean: 1 Min: 1 Max: 2

Length (cm):

Mean: 12 Min: 12 Max: 14
Page 26

Figure 7. cont’d.

RED DEEPSEA CRAB –

Harvestable Adults (> 11 cm CW)

Fall (1963 – 1999)

NEFSC Bottom Trawl Surveys

Crabs caught/station (excluding null stations):

Mean: 4 Min: 1 Max: 58

Length (cm):

Mean: 13 Min: 12 Max: 22
Page 27

Figure 8. Commercial landings for red crab in the northeast.

0

0.5

1

1.5

2

2.5

3

3.5

4

198219841986198819901992199419961998

Year

Landings (metric tons, 000’s)
Publishing in NOAA Technical Memorandum NMFS-NE

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

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 that may warrant a primary level of scientific publishing

should 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

series. Manuscripts which have been rejected by any primary

series strictly because of geographic or temporal limitations may

be submitted to this series.

Manuscripts by Northeast Fisheries Science Center

(NEFSC) authors will be published in this series upon approval

by the NEFSC's Deputy Science & Research Director. Manuscripts by non-NEFSC authors may be published in this series if:

1) the manuscript serves the NEFSC’s mission; 2) the manuscript meets the Deputy Science & Research Director’s approval; and 3) the author arranges for the printing and binding

funds to be transferred to the NEFSC’s Research Communications Unit account from another federal account. For all manuscripts submitted by non-NEFSC authors and published in this

series, the NEFSC will disavow all responsibility for the manuscripts’ contents; authors must accept such responsibility.

The ethics of scientific research and scientific publishing

are a serious matter. All manuscripts submitted to this series are

expected to adhere -- at a minimum -- to the ethical guidelines

contained in Chapter 1 (“Ethical Conduct in Authorship and

Publication”) of the CBE Style Manual, fifth edition (Chicago,

IL: Council of Biology Editors). Copies of the manual are

available at virtually all scientific libraries.

Manuscript Preparation

Organization: Manuscripts must have an abstract, table of

contents, and -- if applicable -- lists of tables, figures, and

acronyms. As much as possible, use traditional scientific

manuscript organization for sections: “Introduction,” “Study

Area,” “Methods & Materials,” “Results,” “Discussion” and/or

“Conclusions,” “Acknowledgments,” and “References Cited.”

Style: All NEFSC publication and report series are obligated to conform to the style contained in the most recent edition

of the United States Government Printing Office Style Manual.

That style manual is silent on many aspects of scientific manuscripts. NEFSC publication and report series rely more on the

CBE Style Manual, fifth edition.

For in-text citations, use the name-date system. A special

effort should be made to ensure that the list of cited works

contains all necessary bibliographic information. For abbreviating serial titles in such lists, use the most recent edition of the

BIOSIS Serial Sources (Philadelphia, PA: Biosciences Information Service). Personal communications must include date of

contact and full name and mailing address of source.

For spelling of scientific and common names of fishes,

mollusks, and decapod crustaceans from the United States and

Canada, use Special Publications No. 20 (fishes), 26 (mollusks),

and 17 (decapod crustaceans) of the American Fisheries Society

(Bethesda, MD). For spelling of scientific and common names

of marine mammals, use Special Publication No. 4 of the

Society for Marine Mammalogy (Lawrence, KS). For spelling in

general, use the most recent edition of Webster’s Third New

International Dictionary of the English Language Unabridged

(Springfield, MA: G.&C. Merriam).

Typing text, tables, and figure captions: Text, tables,

and figure captions must be converted to the NOAA-wide

standard of WordPerfect. In general, keep text simple (e.g.,

don’t switch fonts and type sizes, don’t use hard returns within

paragraphs, don’t indent except to begin paragraphs). Also,

don't use the WordPerfect automatic footnoting function; all

notes should be indicated in the text by simple numerical

superscripts, and listed together in an "Endnotes" section prior

to the "References Cited" section. Especially, don’t use the

WordPerfect graphics function for embedding tables and figures

in text.

Tables may be prepared either with WordPerfect text or

with the WordPerfect table formatting function. If text is used,

then data should be assigned to columns by using all tabs or all

spaces, but not a combination of the two.

Figures must be original (even if oversized) and on paper;

they cannot be photocopies (e.g., Xerox) unless that is all that is

available, nor be on disk. Except under extraordinary circumstances, color will not be used in illustrations.

Manuscript Submission

Authors must submit one paper copy of the double-spaced

manuscript, one disk copy, and original figures (if applicable).

NEFSC authors must include a completely signed-off “NEFSC

Manuscript/Abstract/Webpage Review Form.” Non-NEFSC

authors who are not federal employees will be required to sign

a “Release of Copyright” form.

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. Michael P. Sissenwine, 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

NOAA Technical Memorandum NMFS-NE -- This series is issued irregularly. The series includes: data reports of longterm or large area studies; synthesis reports for major resources or habitats; annual reports of assessment or monitoring programs;

documentary reports of oceanographic conditions or phenomena; manuals describing field and lab techniques; literature surveys of

major resource or habitat topics; findings of task forces or working groups; summary reports of scientific or technical workshops; and

indexed and/or annotated bibliographies. All issues receive internal scientific review and most issues receive technical and copy

editing. Limited free copies are available from authors or the NEFSC. Issues are also available from the National Technical

Information Service, 5285 Port Royal Rd., Springfield, VA 22161.

Fishermen's Report and The Shark Tagger -- The Fishermen's Report (FR) is a quick-turnaround report on the

distribution and relative abundance of commercial fisheries resources as derived from each of the NEFSC's periodic research vessel

surveys of the Northeast's continental shelf. There is no scientific review, nor any technical or copy editing, of the FR; copies are

available through free subscription. The Shark Tagger (TST) is an annual summary of tagging and recapture data on large pelagic

sharks as derived from the NMFS's Cooperative Shark Tagging Program; it also presents information on the biology (movement,

growth, reproduction, etc.) of these sharks as subsequently derived from the tagging and recapture data. There is internal scientific

review, but no technical or copy editing, of the TST; copies are available only to participants in the tagging program.

Northeast Fisheries Science Center Reference Document -- This series is issued irregularly. The series

includes: data reports on field and lab observations or experiments; progress reports on continuing experiments, monitoring, and

assessments; background papers for scientific or technical workshops; and simple bibliographies. Issues receive internal scientific

review but no technical or copy editing. No subscriptions. Free distribution of single copies.

The mission of NOAA's National Marine Fisheries Service (NMFS) is "stewardship of living marine resources for the

benefit of the nation through their science-based conservation and management and promotion of the health of their

environment." As the research arm of the NMFS's Northeast Region, the Northeast Fisheries Science Center (NEFSC)

supports the NMFS mission by "planning, developing, and managing multidisciplinary programs of basic and applied

research to: 1) better understand the living marine resources (including marine mammals) of the Northwest Atlantic,

and the environmental quality essential for their existence and continued productivity; and 2) describe and provide to

management, industry, and the public, options for the utilization and conservation of living marine resources and

maintenance of environmental quality which are consistent with national and regional goals and needs, and with

international commitments." Results of NEFSC research are largely reported in primary scientific media (e.g.,

anonymously-peer-reviewed scientific journals). However, to assist itself in providing data, information, and advice to

its constituents, the NEFSC occasionally releases its results in its own media. Those media are in three categories:

Publications and Reports

of the

Northeast Fisheries Science Center

To obtain a copy of a technical memorandum or a reference document, or to subscribe to the fishermen's report,

write: Research Communications Unit, Northeast Fisheries Science Center, 166 Water St., Woods Hole, MA

02543-1026. An annual list of NEFSC publications and reports is available upon request at the above address.

Any use of trade names in any NEFSC publication or report does not imply endorsement.

STANDARD

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