QL

155

.S63

no. 82

11.63

Library

National Wetlands Research Center

U. S. Fish and Wildlife Service

700 Cajundom ii+., ‘.:. .ard

La!zyet!e. La. 7,:~“; Jr;

Biological Report 82 (1 1.63)

August 1988

TR EL-82-4

Species Profiles: Life Histories and

Environmental Requirements of Coastal Fishes

and Invertebrates (Pacific Northwest)

DUNGENESS CRAB

Fish and Wildlife Service

Coastal Ecology Group

Waterwavs ExDeriment Station

U.S. Department of the Interior U.S. Army Corps of Engineers

Biological Report 82(11.63)

TR EL-82-4

August 1986

Species Profiles: Life Histories and Environmental Requirements

of Coastal Fishes and Invertebrates (Pacific Northwest)

DUNGENESS CRAB

Gilbert B. Pauley, David A. Armstrong, and Thomas W. Heun

School of Fisheries and Washington Cooperative Fishery Research Unit

University of Washington

Seattle, WA 98195

Project Manager

Carroll Cordes

Project Officer

David Moran

National Wetlands Research Center

U.S. Fish and Wildlife Service

1010 Gause Boulevard

Slidell, LA 70458

Performed for

Coastal Ecology Group

Waterways Experiment Station

U.S. Army Corps of Engineers

Vicksburg, MS 39180

and

National Wetlands Research Center

Research and Development

Fish and Wildlife Service

U.S. Department of the Interior

Washington, DC 20240
This series may be referenced as follows:

U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life histories

and environmental requirements of coasts fishes and invertebrates. U.S. Fish

Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4.

This profile should be cited as follows:

Pauley, G.B., D.A. Armstrong, and T.W. Heun. 1986. Species profiles: life

histories and environmental requirements of coastal fishes and invertebrates

(Pacific Northwest)--Dungeness crab. U.S. Fish Wildl. Serv. Biol. Rep.

82(11.63). U.S. Army Corps of Engineers, TR EL-82-4. 20 pp.
PREFACE

This species profile is one of a series on coastal aquatic organisms,

principally fish, of sport, commercial, or ecological importance. The profiles

are designed to provide coastal managers, engineers, and biologists with a brief

comprehensive sketch of the biological characteristics and environmental

requirements of the species and to describe how populations of the species may be

expected to react to environmental changes caused by coastal development. Each

profile has sections on taxonomy, life history, ecological role, environmental

requirements, and economic importance, if applicable. A three-ring binder is

used for this series so that new profiles can be added as they are prepared.

This project is jointly planned and financed by the U.S. Army Corps of Engineers

and the U.S. Fish and Wildlife Service.

Suggestions or questions regarding this report should be directed to one of

the following addresses.

Information Transfer Specialist

National Wetlands Research Center

U.S. Fish and Wildlife Service

NASA-Slide11 Computer Complex

1010 Gause Boulevard

Slidell, LA 70458

or

U.S. Army Engineer Waterways Experiment Station

Attention: WESER-C

Post Office Box 631

Vicksburg, MS 39180

iii
CONVERSION TABLE

Multiply

millimeters (mn)

centimeters (cm)

meters (m)

kilometers (km)

square meters (m2)

square kilometers (km2)

hectares (ha)

10.76 square feet

0.3861 square imiles

2.471 acres

liters (1)

cubic meters (mj)

cubic meters

0.2642 gallons

35.31 cubic feet

0.0008110 acre-feet

milligrams (mg) 0.00003527 ounces

grams (9) 0.03527 ounces

kilograms (kg) 2.205 pounds

metric tons (t) 2205.0 pounds

Imetric tons 1.102 short tons

kilocalories (kcal) 3.968 British thermal units

Celsius degrees 1.8("C) + 32 Fahrenheit degrees

inches 25.40 millimeters

inches 2.54 centimeters

feet (ft) 0.3048 ineters

fathoms 1.829 meters

miles (mi) 1.609 kilometers

nautical miles (rmi) 1.852 kilometers

square feet (ft2)

acres

square miles (mi2)

0.0929 square meters

0.4047 hectares

2.590 square kilometers

gallons (gal)

cubic feet (ft3)

acre-feet

3.785 liters

0.02831 cubic meters

1233.0 cubic meters

ounces (02) 28.35

pounds (lb) 0.4536

short tons (ton) 0.9072

British thermal units (Btu) 0.2520

Fahrenheit degrees 0.5556("F

iV

Metric to U.S. Customary

!LY To Obtain

0.03937 inches

0.3937 inches

3.281 feet

0.6214 miles

U.S. Customary to Metric

32)

grams

kilograms

metric tons

kilocalories

Celsius degrees
CONTENTS

Page

PREFACE ................................. iii

CONVERSION TABLE .............................

FIGURES ................................. ;;

ACKNOWLEDGMENTS ............................. vii

NOMENCLATURE/TAXONOMY/RANGE ....................... 1

MORPHOLOGY/IDENTIFICATION AIDS ..................... 1

REASON FOR INCLUSION IN SERIES ..................... 4

LIFEHISTORY .............................. 4

Mating ................................ 4

Eggs and Fecundity .......................... 4

Larvae

Juveniles

............................... 5

............................... 7

Adults. .....

GROWTH CHARACTERISTICS' : : : : : : : : : : : : : : : : : : : : : : : : :

8

9

THEFISHERY ............................... 10

Commercial Fishery. .......................... 10

Sport Fishery ............................. 12

ECOLOGICAL ROLE ............................. 13

ENVIRONMENTAL REQUIREMENTS ....................... 13

Temperature .............................. 13

Salinity ............................... 14

Temperature-Salinity Interactions ................... 14

Substrate ............................... 14

LITERATURE CITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

V
FIGURES

Number

1

2

3

4

5

6

7

8

Dungeness crab. . . . . . . . . .

Distribut ion of the Dungeness crab

Abdominal differences between fema

. . . . . . . . . . . . . .

Life cycle stages of the Dungeness crab. . . . . . . . . . .

Changes in dry weight over time of three age classes of

Dungeness crabs in Grays Harbor, Washington . . . . . . . . .

in the Pacific Northwest .

le and male Dungeness crabs

Growth, based on mean carapace width, of 0+ crabs from _ . .

settlement as 7-mm first instars, in May through November. . .

Dungeness crab landings by season for individual Pacific

Coast States and the Province of British Columbia, 1955-83. . .

Dungeness crab egg brooding periods at various laboratory

seawater temperatures. . . . . . . . . . . . . . . . . . . . .

Page

1

2

3

6

10

10

11

13

vi
ACKNOWLEDGMENTS

We gratefully acknowledge the reviews of this manuscript by David Somerton,

National Marine Fisheries Service, Seattle, and Paul Dinnel, Fisheries Research

Institute, University of Washington, Seattle. We thank Vali Frank for her

assistance with the graphics. Partial support for David Armstrong to acquire and

analyze some of the original data in this report was provided by Washington Sea

Grant.

vii

Figure 1. Dungeness crab.

DUNGENESS CRAB

NOMENCLATURE/TAXONOMY/RANGE

Scientific name . . . Cancer magister

Dana

Preferred common name . . .Dungeness

crab (Figure 1)

Other common names . . .Pacific edible

crab, edible crab, market crab,

commercial crab

Class . .Crustacea

Order . .*. : : : : : : : : . Decapoda

Infraorder . . . . . . . . .Brachyura

Family . . . . . . . . . . .Cancridae

Geographic range: Coastal waters

along the west coast of North

American from Unalaska Island in

the north to Mexico in the south

(Schmitt 1921; MacKay 1943; Butler

1961a; Mayer 1973). The species

ranges from the intertidal zone to a

depth of at least 98 fathoms and

inhabits substrates of mud, mud with

eelgrass (Zostera sp.) and sand

(Schmitt 1921; Butler 1956; Butler

1961a; Stevens 1982). The distribution of the Dungeness crab in the

Pacific Northwest and the ports of

major commercial landings are shown

in Figure 2.

MORPHOLOGY/IDENTIFICATION AIDS

Dorsal and ventral anatomy of a

Cancer crab is shown in Warner (1977).

1
F:igure 2. Distribution of the Dungeness crab in the Pacific Northwest.

I

PACIFIC #

OCEAN ?;x,.;

ga Coastal distribution _:i.._

+ Major commercial

landing ports

NEWP(

IL WASHINGTON

TILLAd

OREGON

CALIFORNIA

CORTES

TLE

HUMBOLDT

- BAY i

,

2
The following morphology and identification aids were taken from Rudy and

Rudy (1979). Size (type specimen):

carapace 120.7 mm long x 177.8 mm

wide. Color: beige to light brown

with blue trim and hue, darkest

anteriorly, often light orange below,

sometimes light gray-purple below;

inner sides of anterior feet and hands

crimson, fingers not dark. Eyes:

eyestalks short, orbits small.

Antennae: antennules folded lengthwise: antenna1 flagella short, more or

less hairy. Carapace: broadly oval,

uneven but not highly sculptured;

granular. Widest at 10th tooth, no

rostrum. Frontal area: narrow with

five unequal teeth, not markedly

produced beyond outer orbital angles;

middle tooth largest, more advanced

than outer pair; outer pair form inner

angles of orbit. Teeth: (anterolateral) ten, counting orbital tooth;

widest at 10th tooth, which is large

and projecting; all teeth pointed,

with anterior separations. Posterolateral margins: unbroken, entire,

without teeth, meets antero-lateral

margin with distinct angle. Abdomen:

narrow in male, broad in female

(Figure 3). Chelipeds: fingers not

dark; dactyl spinous on upper surface;

fixed finger much deflexed; hand

(propod;:ifI-iih six_ carineae on upper

outer ; wrist (carpus) with

strong inner spine. Walking legs:

rough above; broad and flat (especially propodus and dactylus of last

pair).

Juveniles: antero-lateral and

postero-lateral margins meet at a

distinct angle; carapace widest at

10th tooth; postero-lateral margin

entire; carpus of cheliped with

single spine above, fingers light

colored; carapace not as broad as

adults.

The red rock crab, Cancer productus, also has 10 antero-lateral

teeth; frontal teeth are subequal

(not equal) and the frontal area is

markedly pronounced beyond outer

orbital angles. Cheliped fingers are

black; carapace is widest at eighth

antero-lateral tooth.

The rock crab, Cancer

antennarius, like C. productun

dark red with black-tipped chelae, is

widest at the eighth tooth, but is

red-spotted on its ventral surface.

Cancer oregonensis (Oregon Cancer

Figure 3. Abdominal

crabs. Only males,

commercial crabbers.

differences between female (left) and male (right) Dungeness

which possess a slender abdomen, may be kept by sport and

3
crab) is a small, oval crab with 12

teeth. Both Cancer gracilis and

Cancer jordani, two rather uncommon

sDecieS, have nine teeth. Identification keys to the genus Cancer were

prepared by Kozloff (1974) and Carlton

and Kuris (1975).

REASON FOR INCLUSION IN SERIES

The Dungeness crab supports a

valuable commercial and sport fishery

along the west coast of the United

States. It occupies ecological niches

in both marine and estuarine waters

and is important as both predator and

prey at all life stages. Recent

studies on the environmental

consequences of dredging in estuaries

have established a strong probability

that the Dungeness crab population is

likely to be seriously reduced by

habitat alteration from dredging

unless proper precautions are taken to

reduce losses (Armstrong et al. 1982;

Stevens and Armstrong 1984). The loss

of vital estuarine habitat could

significantly reduce recruitment to

the offshore fishery (Armstrong and

Gunderson 1985).

LIFE HISTORY

Mating

Dungeness crabs mate from April

to September in British Columbia

(MacKay 1942; Butler 1956), mostly in

March and April (David Armstrong,

University of Washington, pers. comm.)

but sometimes in May and June in

Washington (Cleaver 1949), and March

to July in California (Poole and

Gotshall 1965). Mating usually occurs

in offshore locations. Premolt female

crabs are located by adult males for

mating, possibly through a pheromonal

homing system similar to those used by

other crab species (Knudsen 1964;

Edwards 1966; Hartnoll 1969). The

female is held by the male in a

premating embrace up to 7 days prior

to her molting (Snow and Neilsen

1966). Approximately 1 h after

molting of the female is completed,

mating between the hardshell male and

softshell female occurs. Mating

involves the insertion of the male

gonopods into the spermathecae of the

female and the deposition of spermatophores. Following copulation, the

female may be embraced again by the

male for a period of up to 2 days.

Both pre- and postmating embraces may

serve to protect the female from

predation, while insuring the mating

success of the male by guarding the

female against other males (Snow and

Neilsen 1966). The spermatophores

deposited by the male in the

spermathecae contain sperm that are

viable for many months (MacKay

1942), and which may remain viable

through molting until a second egg

extrusion (Orcutt 1978). Eggs are

not fertilized until extrusion, at

which time they are attached to the

female pleopod setae and are carried

beneath the abdominal flap (MacKay

1942; Wild 1983; Stevens 1982). Eggs

hatch in 60 to 120 days.

Eggs and Fecundity

Eggs are extruded from September

to February in British Columbia

(MacKay 1942; Butler 1956), October to

December in Washington (Cleaver 1949;

Mayer 1973), October to March in

Oregon (Waldrom 1958), and September

to November in California (Orcutt et

al. 1976; Wild 1983). An egg mass may

contain from one to two million eggs

(Wild 1983), and a female may produce

up to five million eggs in three or

four broods during her lifetime

(MacKay 1942). Eggs are pale white to

orange at extrusion, becoming progressively darker in color as they develop

(MacKay 1942; Cleaver 1949).

Water temperatures and changes

in water temperatures have considerable influence on the rate of

egg development and mortality after

fertilization and spawning. When

temperatures rise, the rate of egg

development also rises, but so does

4
the rate of mortality. In laboratory

tests (Wild 1983) eggs held at 9.4 OC

hatched in 123 days and at 16.7 OC

they hatched in 64 days. At 10 OC,

685,000 larvae were produced per egg

mass, whereas at 16.7 OC, 14,000 larvae were produced per egg mass. In

Similk Bay, Washington, egg mortality

at 15 OC was serious; a major increase

in mortalities was triggered by a

water temperature increase from 10 OC

to 12 OC (Mayer 1973).

Epibiotic fouling of Dungeness

crab eggs has been linked to increased

egg mortality because of mechanical

interference with hatching and oxygen

consumption (Fisher 1976; Fisher and

Wickham 1976, 1977). Waters with high

and rising nutrient levels caused

increased fouling. Egg predation by a

nemertean worm, Carcinonemertes erz, is thought to enhance the foxing of eggs through the liberation of

yolk during feeding and by its own

defecation (Wickham 1979a, 1979b). In

coastal waters near San Francisco, the

estimated average annual mortality

caused by predation of the worm on

Dungeness crab was over 55% in 1974-79

when worm densities were about 14 per

1,000 eggs (Wickham 1979b).

Eggs mature in about 2 to 3

months (Cleaver 1949; Drcutt 1978;

Wild 1983). The hatching season

commonly shortens from north to south

along the Pacific coast. Eggs hatch

in coastal waters from December to

June in British Columbia (but considerably later in Queen Charlotte

Islands) (MacKay 1942; Butler 1956),

January to April + Washington

(Cleaver 1949; ArmstronAnet al. 1981>,

December to April in Oregon (Reed

1969; Lough 1976), January to early

March in northern California (Wild

1983), and late December to early

February in central California (Wild

1983).

Larvae

Larvae emerge as prezoeae and

molt to zoeae within about 1 h

(Buchanan and Milleman 1969). The

duration of the prezoeal period and

the transformation to zoeae vary with

salinity (Buchanan and Milleman 1969).

The larvae progress through five

zoeal stages before molting into

megalopae (Figure 4; Poole 1966; Reed

1969; Lough 1976). Zoeae first appear

within a distance of 5-16 km from

shore (Lough 1976; Orcutt 1977; Reilly

1983a). Offshore movement and

distribution of larvae probably is

regulated by a variety of factors

including depth, latitude, temperature, salinity and ocean currents

(Reilly 1983a, 1985). Using multiple

regression, the most important

independent variable that distribution

offshore is correlated with is depth

(Reilly 1983a, 1985). Distribution is

dependent upon the larval stage and

the larvae show a die1 pattern of

vertical distribution; they are near

the surface at night (Reilly 1983a,

1985). There is considerable offshore

movement of larvae that occurs during

the zoeal stages; the larvae appear to

be transported seaward from the onset

of hatching (Reilly 1983a).

The megalops (advanced) stage of

the Dungeness crab is found from May

to September off the coast of British

Columbia. In Washington waters, the

megalops first appear in April; abundance peaks in May through June. In

Oregon waters, they are most abundant

in April and May (MacKay 1942; Cleaver

1949; Butler 1956; Lough 1976; Stevens

1982). This trend of abundance indicates larval development begins later

proceeding from south to north. Off

Oregon, megalops are carried within 1

km of shore by tidal currents and by

self-propulsion (Lough 1976).

Megalopae often are abundant on the

hydrozoan Velella velella, when they

are scarce or absent elsewhere in the

water column (Wickham 1979c; Stevens

and Armstrong 1985). Wickham (1979c)

suggested that V. velella aids in the

movement and distribution of megalops,

and possibly provides a food source

and protection from predation.

5
Egg from

female

Fertilized egg

Early post- v

Figure 4.

larval crab Y

Life cycle stages of the Dungeness crab: zoea, megolops, postlarva

(juvenile), and adult.

Larvae eat both zooplankton and

phytoplankton, but zooplankton is most

important (Lough 1976). The larvae

capture food items with the natatory

hairs of their maxillipeds, and size

of food is a selection factor (Lough

1976; Armstrong et al. 1981).

Information on larval predators

and predation rates is scarce. Zoeae

are thought to be consumed by numerous

types of planktivores (Stevens 1982);

megalopae are preyed upon by many

fishes, including coho salmon

(Oncorhynchus kisutch) and chinook

salmon (0. tshawytscha), according to

Orcutt et al. (1976) and Reilly

(1983b). Heavy predation by salmon

may have caused the decline of the

Dungeness crab catch in the San

Francisco Bay area (Reilly 1983b).

There appears to be a direct

relationship between coho salmon

hatchery production in Oregon and the

magnitude of predation on the

megalopae in California waters (Reilly

(1983b). In a study of food habits,

the combined stomachs of eight coho

salmon contained 1,061 megalops

(Orcutt 1977); in a separate study
(MacKay 1942) up to 1,500 megalopae

were reportedly removed from a single

fish. Prince and Gotshall (1976)

found Dungeness crab megalops and

instars, the stages between postlarval

molts, to be the most important food

items of copper rockfish (Sebastes

caurinus) in northern California's

Humboldt Bay.

The abundance of a year class

depends primarily on larval survival

to metamorphosis (Peterson 1973;

Wickham et al. 1976; McKelvey et al.

1980). Natural larval mortality is

probably high because of a combination

of predation, excessively high or low

water temperatures and fluctuations,

a scarcity or low quality of food, and

currents affecting distribution (Lough

1976; Armstrong 1983).

Juveniles

Most megalopae molt into

juveniles in August off the coast of

British Columbia (Figure 4; MacKay

1942; Butler 1956), and in April-May

off the coasts of both Oregon (Lough

1976) and Washington (Stevens 1982).

After molting, the juveniles are

found in shallow coastal waters and

estuaries, and large numbers live

among eelgrass (Zostera sp.) or other

aquatic vegetation that provides

protection and substrate, and harbors

food organisms for early instars

(Butler 1956; Orcutt et al. 1975;

Stevens and Armstrong 1984, 1985).

Recently, shells of bivalves such as

Mya arenaria and Crassostrea gigas

have been documented as very important

habitat for young Dungeness crabs

(Armstrong and Gunderson 1985). In

central California there is evidence

that movement of postlarval Dungeness

crabs into the estuaries takes place

in May and June via bottom currents,

where they stay for 11-15 months

(Tasto 1983). Juveniles are common in

estuaries, while subadults and adults

are common offshore. The importance

of estuaries to juvenile Dungeness

crabs has been discussed in detail by

Armstrong and Gunderson (1985) and

Stevens and Armstrong (1985).

Dungeness crab tag recovery data in

California show a regular pattern of

movement of juvenile crabs out of

estuaries and a random movement of

adult crabs in the ocean (Collier

1983). Stevens and Armstrong (1985)

noted that although mating may occur

in the estuary, spawning takes place

offshore, which would be a major

reason for adults' moving out of the

estuary.

Juveniles molt 11 or 12 times

prior to sexual maturity (Butler 1960,

1961b). Carapace width at the first

instar varies from about 5 mm to

greater than 8.5 mm (Cleaver 1949;

Waldrom 1958; Butler 1960, 1961b;

Poole 1967). After 1 year of growth

beyond hatching, most crabs in Bodega

Bay, California, are in their 8th,

9th, or 10th instar (Poole 1967). By

comparison, crabs from Grays Harbor,

Washington, only attain the sixth or

seventh instar by the end of their

first year of life (Stevens and

Armstrong 1984). Carapace width (CW)

after the first year averages 44 mm in

Grays Harbor, while the range is 63-94

mm in Bodega Bay (Poole 1967; Stevens

et al. 1982). The crabs mature after

about 2 years (Butler 1961b) at

carapace widths of about 116 mm for

males and 100 mm for females (Butler

1960).

The diet of juvenile crabs

consists largely of fish, mollusks,

and crustaceans (Butler 1954; Gotshall

1977; Stevens 1982). In Grays Harbor,

Washington, first-year juveniles

~60 mm CW feed primarily on small

mollusks and crustaceans. Secondyear crabs, 61-100 mm CW. feed on

fish and prefer shrimp (Crangon spp.;

Stevens et al. 1982). Fish also are

important to northern California crabs

< 100 mm CW according to Gotshall

(1977), but Butler (1954) reported

that crustaceans were the primary food

among crabs of this size in the Queen

Charlotte Islands, British Columbia.

Cannibalism among Dungeness crabs has

been noted by various authors (MacKay

7
1942 ; Butler 1954; Tegelberg 1972;

Gotshall 1977; Stevens 1982; Stevens

et al. 1982). Cannibalism was most

prevalent among crabs < 60 mm CW which

fed on smaller crabs of the same year

class, probably during molting

(Stevens 1982; Stevens et al. 1982).

Cannibalism is cited as a possible

cause of the dramatic population

cycles characteristic of the Dungeness

crab fishery (Botsford and Wickham

1978).

Juveniles are captured by a

variety of demersal fishes in the

nearshore area with various flatfishes

(starry flounder, Platichthys

stellatus; English sole, Parophrys

vetulus: and rock sole. Leoidoosetta

bilineata) being the most important

(Reilly 1983b). Other predators on

juvenile crabs are lingcod (Ophiodon

elongatus), cabezon (Scorpaenichthys

marmoratus), wolf-eels (Anarrhichthys

ocellatus), rockfish (Sebastes spp.),

and octopus (Octopus dofleini)

according to Waldrom (1958) and Orcutt

(1977). Predation on Dungeness crabs

may be seasonal in nature, as observed

in white sturaeons. Acioenser

transmontanus (McK&hnie and Fenner

1971). Predation on Dungeness crabs

may have a devastating impact as in

the case of sea otters (Enhydra

lutris) in Orca Inlet, Alaska (Kimker

1985b).

Adults

At about 4 years old, most adult

Dungeness males in the coastal waters

of Washington are of marketable size

(> 159 mm) (Cleaver 1949; Williams

1979). Marketable crabs usually only

molt once a year (MacKay 1942). The

maximum lifespan of Dungeness crabs is

8 to 10 years. The maximum size

attained is about 218 mm CW in males

and 160 mm CW in females at the 16th

instar (MacKay 1942; Butler 1961b).

Adult Dungeness crabs are found

primarily in the ocean but are

also abundant in the inland waters

of Washington and British Columbia.

Along the coast of northern

California, legal-sized and large

sublegal-sized male crabs probably

move offshore (often to the south or

north) in late summer, sometimes

through early winter; sometime in

winter the direction of movement is

probably reversed and the crabs return

inshore. Interannual variation in the

predominant direction of movement is

considerable (Gotshall 1978).

Recently, Collier (1983) has shown a

random movement of adult crabs in the

ocean. Many adult female crabs tagged

off the coast of northern California

moved relatively little (about 2 km)

after 1 year (Diamond and Hankin

1985). Along the coast of southern

Washington, legal-sized males

generally moved inshore and toward the

estuaries in fall (Barry 1985).

Clams are the most important food

of adult Dungeness crabs > 151 mm CW

in northern California (Gotshall 1977)

and > 166 mm CW in British Columbia

(Butler 1954). Crustaceans and fish

are valuable foods of the adult

Dungeness crabs from both Similk Bay

and Grays Harbor, Washington (Mayer

1973; Stevens et al. 1982). Dungeness

crab populations are apparently not

limited by the abundance or scarcity

of particular foods; they are somewhat

nonspecific feeders which readily

adjust to various foods (Gotshall

1977). They have developed an

evolutionary niche for feeding on

mud-sand substrate (Lawton and Elner

1985).

Crabs of different ages or sizes

tend to eat different sizes or kinds

of food (Stevens 1982; Stevens et al.

1982). According to Stevens et al.

(19821, crabs progress from eating

bivalves their first year, to eating

shrimp (Crangon spp.) their second

year, and finally to eating juvenile

teleost fish in the third year; these

shifts may be caused purely by changes

in mechanisms of food handling, or

they may have evolved to reduce

competition among age groups of

crabs. Crabs display a definite die1

8
k

activity; they are more abundant by

day in the subtidal area and more

abundant at night in the intertidal

area; the response is positively

correlated with food availability

(Stevens et al. 1984). Cannibalism is

common among adults, but no

correlations have been made between

the rate of cannibalism and abundance

(Stevens 1982; Stevens et al. 1982).

GROWTH CHARACTERISTICS

In Dungeness crabs, like other

crustaceans, growth proceeds in steps

through a series of molts. The general

process of crustacean growth has been

described by Barnes (1974) and Warner

(1977). The number of molts that a

crab undergoes before becoming mature

depends upon the increment at each

molt and the frequency of molting,

both of which vary among crabs at

different locations. Dungeness crabs

grow in carapace size at molt and gain

weight between molts. In older crabs

the growth, as measured by the percent

change in carapace width, declines as

the frequency of molting slows down,

but the rate of weight gain of the

crabs increases over time. The

probability of annual molting in

female Dungeness crabs declines from

about 1.0 for crabs of 130-135 mm CW

to 0.0 for crabs of 155 mm CW and

larger (Hankin et al. 1985).

Among possible attributes of

estuarine residence suggested by

Stevens and Armstrong (1984) is an

enhanced growth rate compared to that

of siblings of a year class that

settle offshore. Size attained by

juvenile crabs within certain periods

after metamorphosis seems to be

somewhat dependent on latitude and on

time of settlement. Extreme estimates

of age at sexual maturity range from

as long as 4-5 years in British

Columbia (MacKay and Weymouth 1935) to

1 year in San Francisco Bay, where the

crabs reach a carapace width of 100

mm which is usually associated with

sexual maturity (Tasto 1983). More

generally, crabs are predicted to

reach maturity at the end of their

second year after metamorphosis or in

their third growing season over much

of the coast (Butler 1961b; Cleaver

1949). While age and size at sexual

maturity may not differ substantially

along the coasts, estimates of growth

rates of newly settled 0+ crabs do.

Several studies of juvenile

growth rates indicate the process is

accelerated in estuaries or within

nearshore coastal embayments where

water temperatures are relatively

high. Stevens and Armstrong (1984)

studied growth of 0+ and l+ juveniles

in Grays Harbor and found that 6

months after metamorphosis (May to

October) 0+ crab averaged 40 mm in

carapace width and by 1 year were 50

mm. Crabs aged 0+ in Washington

coastal estuaries may molt six to

eight times in the first summer

growing season after which the

frequency apparently declines with the

onset of winter and larger size. A

dramatic indication of seasonal growth

was demonstrated where growth was

based on change in dry weight over

time; rapid growth of the 0+ crabs in

Grays Harbor resulted in a 280-fold

increase in 14 months, and the growth

rate of all age classes declined

beginning in late summer of 1980.

Rapid growth in early summer and

midsummer of 1980 was not repeated in

summer of 1981 (Figure 5).

Populations of 0+ crabs that

settle directly offshore, as well as

l+ nearshore crabs, grow significantly

slower than those in the estuary

(Armstrong and Gunderson 1985).

Young-of-the-year crabs in Grays

Harbor grew from a first instar size

of 7 mm in May to a mean carapace

width of 38.3 mm (sixth instar) by

October; 0+ crabs offshore had only

reached a mean width of 18.9 mm in

November when they were a mixture of

third and fourth instars. Mean bottom

water temperatures

were 15-16 OC

in the estuary

during this time,

while those offshore were 8.5-10 OC,

9
-”

l /

Ka.147 4

lgeo

.02

Y

M J J * 8 0 N D’J F M A M J’J*

1960 1961

Figure 5. Changes in dry weight over

time of three year classes (1978,

1979, and 1980) of Dungeness crabs in

Grays Harbor, Washington. Growth

rates (= k) are expressed as g/g dry

weight per unit time. Note the rapid

growth of 0+ in the summer of 1980 and

the decline in growth of all year

classes in the fall of 1980 (Stevens

and Armstrong 1985).

which may account for the growth

difference (Figure 6). Young-of-theyear juveniles offshore of San

Francisco Bay in the Gulf of the

Farallones also grow substantially

slower than estuarine crabs (Tasto

1983): the data show 0+ crabs at

about 28-30mm, while in the estuary a

good proportion of this age group were

up to 60 mm in width.

Dungeness crab growth is quite

variable along the Pacific coast.

45

40

;

E 35.

r

L 30.

;

y 25.

E

& 20.

e

5 15

F

10

5

15.40 I

01

5 6 7 6 9 10 11

FlONTt!

Figure 6. Growth, based on mean

carapace width, of 0+ crabs from

settlement as 7-mm first instars in

May through November. Bar = f 1 SD.

Grays Harbor population shown as

circles (GH). Offshore population

shown as triangles (OS). Mean bottom

water temperatures at the time of

sampling are also shown for some

samples (Armstrong and Gunderson

1985).

However, in general, it is somewhat

slower in the northern part of the

range (Washington and British Columbia) when compared to the southern

part of the range (California).

THE FISHERY

Commercial Fishery

Commercial landings of Dungeness

crab on the Pacific coast have

fluctuated widely, almost cyclically,

over the past 30 years (Figure 7) and

have been reviewed by Armstrong

(1983). According to Peterson (1973),

commercial landings were highest 1.5

years after a period of strong

upwelling in California and Oregon,

and 6 months following a strong

upwelling in Washington, although the

biological sense of this conclusion is

much in doubt. Botsford and Wickham

10
‘::

195557 59 61 6 3 6 5 6 7 6 9 71 7 3 7 5 7 7 7 9 81 8 3

15 BRITISH COLUMBIA

5-b

l’l’l’l’l’l’l’l’~‘l’1’1’1’1 #

1955 57 5 9 61 6 3 6 5 6 7 6 9 71 73 75 77 7 9 81 8 3

J WASHINGTON

15-

I I I ' . I

'63

' 1 I

57 5 9

' 1

61 6 5

'

6 7

' ' I ' I ' ,

1955 6 9 71 7 3 75 77

L

I’I’I’l”‘I”‘l’l’~ ‘rc

1955 57 5 9 61 6 3 6 5 67 6 9 71 7 3 7 7 7 9

25- CALIFORNIA

I

l'l'l'l'l'l't'l'l'l'l'l'~'lr

1955 57 5 9 61 6 3 6 5 6 7 6 9 71 7 3 75 77 7 9 81 8 3

Figure 7. Dungeness crab landings by season for individual Pacific

Coast States and the Province of British Columbia, 1955-83 (Pacific

Marine Fisheries Commission 1983).

11
(1975) challenged this conclusion by

using auto correlation to show that

commercial landings are cyclic but

upwelling is not.

Another theory to explain catch

fluctuations suggests that periods of

high levels of cannibalism and/or

interspecific competition may cause a

decline in the fishery 3 or 4 years

later (Botsford and Wickham 1978). In

a model predicting recruitment,

McKelvey et a1.(1980) discounted

cannibalism as a factor and contended

that changes in egg and larval

survival have regulated population

success. Larval survival may be

seriously altered by a combination of

environmental factors that can cause

increased mortality if unfavorable for

even short periods of time (Lough

1976). Stevens and Armstrong (1981)

indicate that diseases caused by

various organisms (bacteria, Protozoa,

or fungi) may be responsible for mass

mortalities of adult crabs.

Predation may have a profound

impact on the Dungeness crab

commercial fishery in certain

geographic areas (Kimker 1985b).

Reilly (1983b) theorizes that hatchery

released coho salmon from the Columbia

River continue to suppress the

Dungeness crab fishery through

extensive predation.

Landings of Dungeness crab from

1954 to 1983 are broken down by State

and Province in Figure 7. In general,

the landings in Washington (except for

Puget Sound), Oregon, and California

follow similar trends. Landings from

Puget Sound and British Columbia are

lower but show less annual variation.

Alaska landings bear little relation

to other areas of the Pacific

Northwest. The Alaska catch of 15.6

million pounds in 1981 was a record

for Alaska, but Washington coastal

crab landings of 2.6 million pounds in

the same year were the lowest in 30

years (Demory 1982). Recent reviews

of the commercial Dungeness crab

fishery have been published for Alaska

(Eaton 1985; Kimker 1985b; Koeneman

1985; Merritt 1985), British Columbia

(Jamieson 1985), Washington (Barry

1985), Oregon (Demory 1985), and

California (Dahlstrom and Wild 1983;

Warner 1985).

Sport Fishery

Sport catch data are scarce and

according to Barry (1985) the

Washington sport fishery on Dungeness

crabs amounts to less than one

percent of the annual commercial

harvest. Most of the available sport

catch data are from a survey reported

by Williams (1979). He revealed

that from April through August 1974,

471 crabs were taken intertidally

at Mission Beach, Washington, by 735

sport crabbers. April, May, and

June produced the best sport catches

with the highest average catches

occurring on low tides that ranged

from -0.60 to -0.74 m. Aerial

surveys made over Puget Sound beaches

using Williams' (1979) survey data

estimated that the beaches of

Washington State probably supported

about 20,000 crabbers during those

months in 1974. In 1975, the sport

crab pot fishery alone (other sport

catch methods are ring nets, dip

nets, and hook and line) accounted

for the harvest of about 300,000

Dungeness crabs (Tegelberg 1976).

Only male crabs may be taken in the

sport fishery (Figure 3). In

Washington there is a minimum size

of 6 or 6.25 inches carapace width

(depending on the area), meas;;;;

directly in front of the

anteriolateral spines. In Oregon

the minimum size is 6.25 inches,

measured similarly. The sport catch

is primarily found in Hood Canal,

Puget Sound, and the major Pacific

coast estuaries.

The State of Oregon has sought to

limit the conflict between sport and

commercial crabbers by restricting

commercial crabbing to the middle of

the week and to the use of sport gear

(Demory 1985).

12
ECOLOGICAL ROLE

Dungeness crabs consume a wide

variety of food organisms and are prey

to numerous predators. Crabs

contribute to several trophic levels

as they progress through successive

life stages. The larvae largely

consume plankton (Lough 1976) and are

preyed upon by numerous fishes.

Adults and juveniles are preyed upon

by sea otters, fishes, and octopuses

(Butler 1954; Waldrom 1958; Stevens

1982; Reilly 1983b; Kimker 198513).

Cannibalism is common and probably

exercises some control over abundance.

In their various life stages,

Dungeness crabs feed on a variety of

mollusks, crustaceans, and fish

species (Stevens et al. 1982). Other

information on the ecological role of

each life stage is given in the life

history section.

ENVIRONMENTAL REQUIREMENTS

L

Temperature

The temperature preferences of

adult crabs are different among

seasons (Mayer 1973). They are

somewhat tolerant of abrupt

temperature and salinity fluctuations

(Cleaver 1949), and water temperatures

from 3 to 19 OC were listed as normal

for the Dungeness crab (Cleaver 1949).

Dungeness crabs have different

optimal water temperatures at

different stages. In the laboratory,

Des Voigne (1973) reported that

optimal water temperatures for mating

ranged from 12 to 16 OC during long

photoperiods. Wild (1973) noted an

apparent trend towards crabs mating

later in colder water in his

laboratory experiments, but noted that

mating took place between 10 and

17 OC. In Washington coastal waters,

where Dungeness crabs usually mate in

early spring, the bottom temperatures

are between 8 and 10 OC (Armstrong,

pers. comm.). According to Wild

(1983), the egg brooding periods

varied inversely with seawater

temperatures of 9 to 17 OC (Figure 8).

Prolonged egg brooding periods in

colder water are consistent with

prolonged occurrences of ovigerous

crabs and cooler ocean temperatures as

you move progressively northward along

the Pacific coast (Wild 1983).

Hatching success, considered as the

number of larvae that hatch from an

egg mass, decreased as the temperature

increased from 10 to 17 OC (Wild

1983). Mayer (1973) found a similar

correlation between egg mortality and

temperature with 20% mortality at

10 OC and 100% mortality at 20 OC.

9 10 I, I2 13 14 1 5 16 17

TEMP %

Figure 8. Dungeness crab egg brooding

periods at various laboratory seawater

temperatures (Wild 1983).

Optimal temperatures for larvae

are 10 to 14 OC. Juvenile crabs,

80 mm wide and acclimated to 10.0 OC,

have been exposed to water temperatures up to 25.0 OC for 7 days with

little or no mortality (Des Voigne

1973); however, an increase of 2.5 OC

above 25 OC was fatal to 100% of all

crabs tested. In the laboratory,

adult crabs had a maximum tolerable

temperature of 25 OC during long

photoperiods, which decreased to 20 OC

when exposed to short photoperiods

13
(Des Voigne 1973). With adult crabs

held for 8 months, Wild (1983)

observed that mortality increased with

temperature from 17% at 10 OC to 58%

at 13 OC and to 80% at 17 'C, although

laboratory stress probably exacerbated

the effect of high temperatures.

Salinity

Tolerance to salinity varies

among the life stages of the Dungeness

crab. In general, salinity is not as

important as temperature to egg

development and hatching, but the

larvae are highly sensitive to changes

salinity (Buchanan and Milleman

G69). The percentage of -eggs

hatching was optimum at 15 ppt, but

hatching occurred to some degree over

a wide range of salinities between 10

ppt and 32 ppt (Buchanan and Milleman

1969). When salinity was increased

from 15 ppt to 32 ppt, the average

prezoeal period was reduced from about

60 min to less than 11 min. At a

salinity of 10 ppt, no prezoeae molted

to zoeaea, but 100% molted at 30 ppt

(Buchanan and Milleman 1969). The

highest survival for larvae was

between salinities from 25 ppt and 30

ppt (Reed 1969). Survival decreased

with salinity and was poorest at

salinities of 15 ppt (Reed 1969). No

juvenile or adult tolerance levels are

available in the literature at this

time.

Temperature-Salinity Interactions

Salinity and temperature are both

related to larval survival. Significant interaction exists between

these two factors with salinity

buffering temperature. At favorable

temperatures, unfavorable salinities

resulted in complete mortality, but

favorable salinities at unfavorable

temperatures allowed some survival

(Reed 1969). The most obvious effect

on growth rate occurred at temperatures that resulted in the best

survival. Salinities that favored

survival generally had little effect

on zoeal growth. Survival of zoeae is

optimal between the water temperatures

of 10.0 and 13.0 OC and salinities of

25 and 30 ppt (Reed 1969). The

significant interaction between temperature and salinity dictates caution

when making statements about either

variable independent of the other one.

The effects of temperature or salinity

alone on C. magister zoeae do not

appear to cause large fluctuations in

zoeal survival in the ocean (Reed

1969; Lough 1976).

Substrate

Adult crabs are found living over

several substrate types (Schmitt 1921;

Cleaver 1949; Butler 1956), but they

prefer sandy-mud bottoms (Karpov

1982). Early juveniles prefer beds of

eelgrass, shell, or sandy mud (Stevens

and Armstrong 1984). This preference

may stem from an abundance of food

organisms on such substrates or

perhaps the crabs find shelter from

predation there (Stevens 1982). Older

crabs seem less dependent on

epibenthic cover and can be found over

more exposed substrates. Most crabs

remain in the subtidal environment,

but may venture into littoral areas at

high tide (Stevens et al. 1984). This

behavior is enhanced by the presence

of preferred food items and decreased

during low salinities following heavy

rains.

14
LITERATURE CITED

Armstrong, D.A. 1983. Cyclic crab

abundance and relationship to

environmental causes. Pages 102-110

in W.

year:

Wooster, ed. From year to

interannual variability of

the environment and fisheries of the

Gulf of Alaska. University of

Washington Sea Grant Rep. WST-WO

83-3, Seattle.

Armstrong, D.A., and D.R. Gunderson.

1985. The role of estuaries in

Dungeness crab early life history: a

case study in Grays Harbor, Washington. Pages 145-170 in Proceedings of

the symposium onDungeness crab

biology and management. University

of Alaska Sea Grant Rep. 85-3,

Fairbanks.

Armstrong, D.A., B.G. Stevens, and

J.C. Hoeman. 1981. Distribution

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shrimp, Crangon and dredging-related

mortality of invertebrates and fish

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J.C. Hoeman. 1982. Distribution

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Crangon shrimp, and dredging-related

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15
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1 7
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i?fe history, environment, and

18
mariculture studies of the Dungeness

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20
\r

1111-10,

lEPORT DOCUMENTATION 1. REWRT No. 2.

PAGE Biological Report 82(11.63)*

. . Title and Subtitle

Species Profiles: Life Histories and Environmental

iequirements of Coastal Fishes and Invertebrates (Pacific

Vorthwest)--Dungeness crab

_.-_

Author(s)

Bilbert B. Pauleya, David A. Armstrongb, and Thomas W. Heuna

3. ReCwl.nt’s Accessmn No.

5. Rw-art Date

August 1986

6.

8. Pwfoming Organization Rcpt. N

. Perfomina Organization Name and Address

Washington Cooperative Fishery

Research Unit

University of Washington

Seattle, WA 98195

2. Sponsoring Orcaniratlon Name and Address

National Wetlands Research Center

Fish and Wildlife Service

U.S. Department of the Interior

Washington, DC 20240

10. PmjedlTasklWork Unit No.

bSchool of Fisheries

University of Washington 11. Contract(C) or Grant(G) No.

Seattle, WA 98195 (C)

(G)

13. Type of Report 6 Permd Cover.,

U.S. Army Corps of Engineers

Waterways Experiment Station

P.D. Box 631

Vicksburg, MS 39180

E

14.

5. Supplcm.ntary Natea

*U.S. Army Corps of Engineers Report No. TR EL-82-4

6. Abstract (Limit: 200 words)

Species profiles are literature summaries of the taxonomy, morphology, range, lift

history, and environmental requirements of coastal aquatic species. They are designed tc

assist in environmental impact assessment. The Dungeness crab (Cancer magister) is fount

off the coasts of Washington, Oregon, and southern British Columbia, as well as in the

estuarine waters of this geographic area. It is a shellfish highly prized and soughi

after by both commercial and sport fishermen. In Washington and Oregon, only male crab!

may be retained by sport and commercial fishermen. Commercial crab catches are high11

variable from year to year, but the catches from Washington and Oregon follow a ver)

similar pattern. The highest sport catches take place on low tides ranging from -0.60 tc

-0.74 m. Dungeness crab go through a life cycle that involves several metamorphic stages

zoea, megalops, postlarval crab, and adult crab. Hatching success decreases as wate!

temperature increases from 10 to 17 OC; the optimal temperature for larval crabs i:

between 10 and 14 OC. Salinity is not as important to egg development and hatching a!

temperature, but optimum hatching occurs at about 15 ppt.

1. Doc"ment Analysis . . oesccnpton

Temperature

Estuaries

Shellfish

Salinity

Feeding habits

Crabs

Life cycles

Temperature

- -

b. IdentlRen/Opcn.Ended Terms

Dungeness crab

Cancer magister Dana

Lifehistory

Ecological role

Environmental requirements

c. COSATI Field/Group

8. Avmlabiiity Statement

Unlimited

19. Sccurvty Class (This Rep-art)

Unclassified

Sccuruty Class (This Page)

Unclassified

2,. No. of Pager

20 _-

22. Price

ee ANSI-239.18) CPTIONAL FORM 272 M-77)

(Formerly NTIS-351

oeprrtmen+ of Commcr+. L
TAKE PRIDE

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U.S. FISH AND WIlDLIfE SERVICE

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fostering the wisest use of our land and water resources, protecting our fish and wildlife,

preserving theenvironmental and cultural values of our national parks and historical places,

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