- R1E1 596 SPECIES PROFILES LIFE HISTORIES AND ENVIRONMENTAL /TREQUIREMENTS OF CORSTNL..(U) ASHINGTON COOPERATIVEFISHERY RESEARCH UNIT SEATTLE G B PRULEY ET AL RUG 65

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Species Profiles: Life Histories andEnvironmental Requirements of Coastal Fishesand Invertebrates (Pacific Northwest)

DUNGENESS CRAB

ELECTEJUN 1 9 1987

Coastal Ecology GroupFish and Wildlife Service Waterways Experiment Station

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

Biological Report 82(11.63)TR EL-82-4August 1986

Species Profiles: Life Histories and Environmental Requirementsof Coastal Fishes and Invertebrates (Pacific Northwest)

DUNGENESS CRAB

by

Gilbert B. Pauley, David A. Armstrong, and Thomas W. HeunSchool of Fisheries and Washington Cooperative Fishery Research Unit

University of WashingtonSeattle, WA 98195

Project ManagerCarroll CordesProject OfficerDavid Moran

National Wetlands Research CenterU.S. Fish and Wildlife Service

1010 Gause BoulevardSlidell, LA 70458

Performed forCoastal-Ecology Group

Waterways Experiment StationU.S. Army Corps of Engineers

Vicksburg, MS 39180

and

National Wetlands Research CenterResearch and DevelopmentFish and Wildlife Service

U.S. Department of the InteriorWashington, DC 20240

This series may be referenced as follows:

U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life historiesand environmental requirements of coastiT fishes and invertebrates. U.S. FishWildl. 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., B.A. Armstrong, and T.W. Heun. 1986. Species profiles: lifehistories 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 profilesare designed to provide coastal managers, engineers, and biologists with a briefcomprehensive sketch of the biological characteristics and environmentalrequirements of the species and to describe how populations of the species may beexpected to react to environmental changes caused by coastal development. Eachprofile has sections on taxonomy, life history, ecological role, environmentalrequirements, and economic importance, if applicable A three-ring binder isused for this series so that new profiles can be ad ed as they are prepared.This project is jointly planned and financed by the U. . Army Corps of Engineersand the U.S. Fish and Wildlife Service. /

Suggestions or questions regarding this report sguld be directed to one ofthe following addresses. /

Information Transfer Specialist a - 1 { J "National Wetlands Research CenterU.S. Fish and Wildlife ServiceNASA-Slidell Computer Complex1010 Gause BoulevardSlidell, LA 70458

or

U.S. Army Engineer Waterways Experiment StationAttention: WESER-CPost Office Box 631Vicksburg, MS 39180 Acoession For

NT IS GRA&I-DTIC TABUnannounced QJustificatlon

Distribution/

Availability CodesAvoil and/or

4 Dist Special

iii flo

CONVERSION TABLE

Metric to U.S. Customary

multiply P1 To Obtain

millimeters (m) 0.03937 inchescentimeters (cm) 0.3937 inchesmeters (in) 3.281 feetkilometers (in) 0.6214 miles

square meters (m 2) 2 10.76 square feetsquare kilometers (k) 0.3861 square mileshectares (ha) 2.471 acres

liters (1) 0.2642 gallonscubic meters (m3) 35.31 cubic feetcubic meters 0.0008110 acre-feet

milligrams (iag) 0.00003527 ouncesgrams (9) 0.03527 ounceskilograms (kg) 2.205 pounds v m

metric tons (t) 2205.0 poundsmetric tons 1.102 short tonskilocalories (kcal) 3.968 British thermal units

Celsius degrees 1.8(-C) + 32 Fahrenheit degrees

U.S. Customary to Metric

inches 25.40 millimetersinches 2.54 centimetersfeet (ft) 0.3048 metersfathoms 1.829 metersmiles (mi) 1.609 kilometersnautical miles (nmt) 1.852 kilometers

square feet (ft2) 0.0929 square metersacres 2 0.4047 hectaressquare miles (m) 2.590 square kilometers

gallons (gal) 3.785 literscubic feet (ft3) 0.02831 cubic metersacre-feet 1233.0 cubit meters

ounces (oz) 28.35 granspounds (lb) 0.4536 kilogramsshort tons (ton) 0.9072 metric tonsBritish thermal units (Btu) 0.2520 kilocalories

Fahrenheit degrees 0.5556('F -32) Celsius degrees

ivFarnhi eges0.56°F-3) esusdgIe

CONTENTS

PREFACE.................... . . . ..... . . ..... .. .. .. .. ....CONVERSION TABLE .. .... ..................... ivFIGURES. .. .... ......................... viACKNOWLEDGMENTS .. .. ...... ....... ....... ....... vii

NOMENCLATURE/TAXONMY/RANGE. .. .... ....... ...... ..... 1MORPHOLOGY/IDENTIFICATION AIDS. .. ....... ...... ........ 1REASON FOR INCLUSION IN SERIES .. .... ....... ....... ... 4LIFE HISTORY .. .... ....... ....... ...... ....... 4

Mating. .. ....... ...... ....... ....... .... 4Eggs and Fecundity. .. ....... ...... ....... ...... 4Larvae. .. ....... ...... ....... ....... .... 5Juveniles. .. .... ....... ...... ....... ....... 7Adults. .. .. ...... ....... ...... ....... .....

GROWTH CHARACTERISTICS. .. ....... ...... ....... .... 9THE FISHERY .. .. ...... ....... ....... ...... ... 10

Coinercial Fishery .. .. .... ...... ....... ........ 10Sport Fishery .. .. ...... ....... ...... ........ 12

ECOLOGICAL ROLE .. .. ...... ....... ....... ....... 13ENVIRONMENTAL REQUIREMENTS. .. ....... ....... ......... 13

Temperature .. .. ...... ....... ....... ......... 13Salinity. .. ....... ........................................ 14Temperature-Salinity Interactions. .. .... ....... ....... 14Substrate .. .. ...... ....... ....... ...... ... 14

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

Va

FIGURES

Number Page

1 Dungeness crab . ....... ......................... 1

2 Distribution of the Dungeness crab in the Pacific Northwest . 2

3 Abdominal differences between female and male Dungeness crabs . 3

4 Life cycle stages of the Dungeness crab ..... ............ 6

5 Changes in dry weight over time of three age classes ofDungeness crabs in Grays Harbor, Washington .... .......... 10

6 Growth, based on mean carapace width, of 0+ crabs fromsettlement as 7-m first instars, in May through November. .. 10

7 Dungeness crab landings by season for individual PacificCoast States and the Province of British Columbia, 1955-83. .. 11

8 Dungeness crab egg brooding periods at various laboratoryseawater temperatures ..... ..................... ... 13

I. vi

p *

ACKNOWLEDGMENTS

We gratefully acknowledge the reviews of this manuscript by David Somerton,National Marine Fisheries Service, Seattle, and Paul Dinnel, Fisheries ResearchInstitute, University of Washington, Seattle. We thank Vali Frank for herassistance with the graphics. Partial support for David Armstrong to acquire andanalyze some of the original data in this report was provided by Washington SeaGrant.

vii

1 II' 1 .

w.

Figure 1. Dungeness crab.

- DUNGENESS CRAB 5 -4

NOMENCLATURE/TAXONOMY/RANGE(Schmitt 1921; MacKay 1943; Butler

Scientific name . -Cancer magistero> 1961a; Mayer 1973). The speciesDana ranges from the intertidal zone to a

Preferred common name . . .Dungeness-- depth of at least 98 fathoms andcrab (Figure 1) - inhabits substrates of mud, mud with

Other common names . . >.Pacific edible eelgrass (Zostera sp.) and sandcrab; edible crab; market crab; (Schmitt 1921; Butler 1956; Butlercommercial crab;- 1961a; Stevens 1982). The distri-

Class ........... .7_Crustacea bution of the Dungeness crab in theOrder .............. .. Decapoaia) Pacific Northwest and the ports of ..Infraorder ...... . --. -frahyura major commercial landings are shownFamily _ -- . ......... Cancridae in Figure 2.

Geographic range: Coastal waters MORPHOLOGY/IDENTIFICATION AIDS[ along the west coast of North

American from Unalaska Island in Dorsal and ventral anatomy of athe north to Mexico in the south Cancer crab is shown in Warner (1977).

\1

• J44

C,

-=- : -" . , - ..Na

SrTAITOF .RJN

PACIFICOCEANGRY

ARBOR SEATTLE

0 s 10 IB AY WASHINGTON0 so tooA

KILOME TERS

Coastal distribution

*Major commercial TLA PORTLAND d C01 4fvd,landing ports'-RIE

CALIFORNIA

EUHUMBOLDT

BA Y

Figure 2. Distribution of the Dungeness crab in the Pacific Northwest.

2

The following morphology and identifi- fixed finger much deflexed; handcation aids were taken from Rudy and (propodus) with six carineae on upperRudy (1979). Size (type specimen): outer surface; wrist (carpus) withcarapace 120.7 m long x 177.8 - strong inner spine. Walking legs:wide. Color: beige to light brown rough above; broad and flat (espe-with blue trim and hue, darkest cially propodus and dactylus of lastanteriorly, often light orange below, pair).sometimes light gray-purple below;inner sides of anterior feet and hands Juveniles: antero-lateral andcrimson, fingers not dark. Eyes: postero-lateral margins meet at aeyestalks short, orbits small. distinct angle; carapace widest atAntennae: antennules folded length- 10th tooth; postero-lateral marginwise: antennal flagella short, more or entire; carpus of cheliped withless hairy. Carapace: broadly oval, single spine above, fingers lightuneven but not highly sculptured; colored; carapace not as broad asgranular. Widest at 10th tooth, no adults.rostrum. Frontal area: narrow withfive unequal teeth, not markedly The red rock crab, Cancer pro-produced beyond outer orbital angles; ductus, also has 10 antero-lateralmiddle tooth largest, more advanced teeth; frontal teeth are subequalthan outer pair; outer pair form inner (not equal) and the frontal area isangles of orbit. Teeth: (antero- markedly pronounced beyond outerlateral) ten, counting orbital tooth; orbital angles. Cheliped fingers arewidest at 10th tooth, which is large black; carapace is widest at eighthand projecting; all teeth pointed, antero-lateral tooth.with anterior separations. Postero-lateral margins: unbroken, entire, The rock crab, Cancerwithout teeth, meets antero-lateral antennarius, like C. productus, ismargin with distinct angle. Abdomen: dark red with black-tipped chelae, isnarrow in male, broad in female widest at the eighth tooth, but is(Figure 3). Chelipeds: fingers not red-spotted on its ventral surface.dark; dactyl spinous on upper surface; Cancer oregonensis (Oregon Cancer

Figure 3. Abdominal differences between female (left) and male (right) Dungenesscrabs. Only males, which possess a slender abdomen, may be kept by sport andcommercial crabbers.

3-.v

X 11 , 11 ' 11 11' Id , 1 111 11 11,11, 111

NYA9&&%I~~ ~ ~ ~ ~ 91 'uI"1 IIIiI

crab) is a small, oval crab with 12 1966). Approximately 1 h afterteeth. Both Cancer gracilis and molting of the female is completed,Cancer jordani, two rather uncommon mating between the hardshell male andspecies, have nine teeth. Identifi- softshell female occurs. Matingcation keys to the genus Cancer were involves the insertion of the maleprepared by Kozloff (1974) and Carlton gonopods into the spermathecae of theand Kuris (1975). female and the deposition of spermato-

phores. Following copulation, thefemale may be embraced again by the

REASON FOR INCLUSION IN SERIES male for a period of up to 2 days.Both pre- and postmating embraces may

The Dungeness crab supports a serve to protect the female fromvaluable commercial and sport fishery predation, while insuring the matingalong the west coast of the United success of the male by guarding theStates. It occupies ecological niches female against other males (Snow andin both marine and estuarine waters Neilsen 1966). The spermatophoresand is important as both predator and deposited by the male in theprey at all life stages. Recent spermathecae contain sperm that arestudies on the environmental viable for many months (MacKayconsequences of dredging in estuaries 1942), and which may remain viablehave established a strong probability through molting until a second eggthat the Dungeness crab population is extrusion (Orcutt 1978). Eggs arelikely to be seriously reduced by not fertilized until extrusion, athabitat alteration from dredging which time they are attached to theunless proper precautions are taken to female pleopod setae and are carriedreduce losses (Armstrong et al. 1982; beneath the abdominal flap (MacKayStevens and Armstrong 1984). The loss 1942; Wild 1983; Stevens 1982). Eggsof vital estuarine habitat could hatch in 60 to 120 days. ,.significantly reduce recruitment tothe offshore fishery (Armstrong and Eggs and FecundityGunderson 1985).

Eggs are extruded from Septemberto February in British Columbia

LIFE HISTORY (MacKay 1942; Butler 1956), October toDecember in Washington (Cleaver 1949;

Mating Mayer 1973), October to March inOregon (Waldrom 1958), and September

Dungeness crabs mate from April to November in California (Orcutt etto September in British Columbia al. 1976; Wild 1983). An egg mass may(MacKay 1942; Butler 1956), mostly in contain from one to two million eggsMarch and April (David Armstrong, (Wild 1983), and a female may produceUniversity of Washington, pers. comm.) up to five million eggs in three orbut sometimes in May and June in four broods during her lifetimeWashington (Cleaver 1949), and March (MacKay 1942). Eggs are pale white toto July in California (Poole and orange at extrusion, becoming progres-Gotshall 1965). Mating usually occurs sively darker in color as they developin offshore locations. Premolt female (MacKay 1942; Cleaver 1949).crabs are located by adult males formating, possibly through a pheromonal Water temperatures and changeshoming system similar to those used by in water temperatures have consid-other crab species (Knudsen 1964; erable influence on the rate ofEdwards 1966; Hartnoll 1969). The egg development and mortality afterfemale is held by the male in a fertilization and spawning. Whenpremating embrace up to 7 days prior temperatures rise, the rate of eggto her molting (Snow and Neilsen development also rises, but so does

4 V" • °

the rate of mortality. In laboratory (Buchanan and Milleman 1969). Thetests (Wild 1983) eggs held at 9.4 OC duration of the prezoeal period andhatched in 123 days and at 16.7 OC the transformation to zoeae vary withthey hatched in 64 days. At 10 °C, salinity (Buchanan and Milleman 1969).685,000 larvae were produced per eggmass, whereas at 16.7 *C, 14,000 lar- The larvae progress through fivevae were produced per egg mass. In zoeal stages before molting intoSimilk Bay, Washington, egg mortality megalopae (Figure 4; Poole 1966; Reedat 15 OC was serious; a major increase 1969; Lough 1976). Zoeae first appearin mortalities was triggered by a within a distance of 5-16 km fromwater temperature increase from 10 *C shore (Lough 1976; Orcutt 1977; Reillyto 12 °C (Mayer 1973). 1983a). Offshore movement and

distribution of larvae probably isEpibiotic fouling of Dungeness regulated by a variety of factors

crab eggs has been linked to increased including depth, latitude, tempera-egg mortality because of mechanical ture, salinity and ocean currentsinterference with hatching and oxygen (Reilly 1983a, 1985). Using multipleconsumption (Fisher 1976; Fisher and regression, the most importantWickham 1976, 1977). Waters with high independent variable that distributionand rising nutrient levels caused offshore is correlated with is depthincreased fouling. Egg predation by a (Reilly 1983a, 1985). Distribution isnemertean worm, Carcinonemertes er- dependent upon the larval stage andrans, is thought to enhance the foul- the larvae show a diel pattern ofing of eggs through the liberation of vertical distribution; they are nearyolk during feeding and by its own the surface at night (Reilly 1983a,defecation (Wickham 1979a, 1979b). In 1985). There is considerable offshorecoastal waters near San Francisco, the movement of larvae that occurs duringestimated average annual mortality the zoeal stages; the larvae appear tocaused by predation of the worm on be transported seaward from the onsetDungeness crab was over 55% in 1974-79 of hatching (Reilly 1983a).when worm densities were about 14 per1,000 eggs (Wickham 1979b). The megalops (advanced) stage of

the Dungeness crab is found from MayEggs mature in about 2 to 3 to September off the coast of British

months (Cleaver 1949; Orcutt 1978; Columbia. In Washington waters, the NWild 1983). The hatching season megalops first appear in April; abun- vecommonly shortens from north to south dance peaks in May through June. Inalong the Pacific coast. Eggs hatch Oregon waters, they are most abundant *-' "in coastal waters from December to in April and May (MacKay 1942; CleaverJune in British Columbia (but consid- 1949; Butler 1956; Lough 1976; Stevenserably later in Queen Charlotte 1982). This trend of abundance indi-Islands) (MacKay 1942; Butler 1956), cates larval development begins laterJanuary to April in Washington proceeding from south to north. Off(Cleaver 1949; Armstrong et al. 1981), Oregon, megalops are carried within 1December to April in Oregon (Reed km of shore by tidal currents and by1969; Lough 1976), January to early self-propulsion (Lough 1976).March in northern California (Wild Megalopae often are abundant on the1983), and late December to early hydrozoan Velella velella, when theyFebruary in central California (Wild are scarce or absent elsewhere in the1983). water column (Wickham 1979c; Stevens

and Armstrong 1985). Wickham (1979c)Larvae suggested that V. velella aids in the

movement and distribution of megalops,Larvae emerge as prezoeae and and possibly provides a food source

molt to zoeae within about 1 h and protection from predation.

5

T'4

Egg from

female

0 Fertilized egg

Sperm frommale Zoea

Adult crab

Megalops

Early post-larval crab

Figure 4. Life cycle stages of the Dungeness crab: zoea, megolops, postlarva(juvenile), and adult.

Larvae eat both zooplankton and salmon (0. tshawytscha), according tophytoplankton, but zooplankton is most Orcutt et a]. (1976) and Reillyimportant (Lough 1976). The larvae (1983b). Heavy predation by salmoncapture food items with the natatory may have caused the decline of thehairs of their maxillipeds, and size Dungeness crab catch in the Sanof food is a selection factor (Lough Francisco Bay area (Reilly 1983b).1976; Armstrong et al. 1981). There appears to be a direct

relationship between coho salmonInformation on larval predators hatchery production in Oregon and the

and predation rates is scarce. Zoeae magnitude of predation on theare thought to be consumed by numerous megalopae in California waters (Reillytypes of planktivores (Stevens 1982); (1983b). In a study of food habits,megalopae are preyed upon by many the combined stomachs of eight cohofishes, including coho salmon salmon contained 1,061 megalops(Oncorhynchus kisutch) and chinook (Orcutt 1977); in a separate study

6

0(MacKay 1942) up to 1,500 megalopae Stevens and Armstrong (1985).were reportedly removed from a single Dungeness crab tag recovery data infish. Prince and Gotshall (1976) California show a regular pattern offound Dungeness crab megalops and movement of juvenile crabs out ofinstars, the stages between postlarval estuaries and a random movement ofmolts, to be the most important food adult crabs in the ocean (Collieritems of copper rockfish (Sebastes 1983). Stevens and Armstrong (1985)caurigus) in northern California's noted that although mating may occurHumboldt Bay. in the estuary, spawning takes place

offshore, which would be a majorThe abundance of a year class reason for adults' moving out of the

depends primarily on larval survival estuary.to metamorphosis (Peterson 1973;Wickham et al. 1976; McKelvey et al. Juveniles molt 11 or 12 times1980). Natural larval mortality is prior to sexual maturity (Butler 1960,probably high because of a combination 1961b). Carapace width at the firstof predation, excessively high or low instar varies from about 5 mm towater temperatures and fluctuations, greater than 8.5 mm (Cleaver 1949;a scarcity or low quality of food, and Waldrom 1958; Butler 1960, 1961b;currents affecting distribution (Lough Poole 1967). After 1 year of growth1976; Armstrong 1983). beyond hatching, most crabs in Bodega

Bay, California, are in their 8th,Juveniles 9th, or 10th instar (Poole 1967). By

comparison, crabs from Grays Harbor,Most megalopae molt into Washington, only attain the sixth or

juveniles in August off the coast of seventh instar by the end of theirBritish Columbia (Figure 4; MacKay first year of life (Stevens and1942; Butler 1956), and in April-May Armstrong 1984). Carapace width (CW)off the coasts of both Oregon (Lough after the first year averages 44 mm in1976) and Washington (Stevens 1982). Grays Harbor, while the range is 63-94After molting, the juveniles are mm in Bodega Bay (Poole 1967; Stevensfound in shallow coastal waters and et al. 1982). The crabs mature afteristuaries, and large numbers live about 2 years (Butler 1961b) atamong eelgrass (Zostera sp.) or other carapace widths of about 116 mm foraquatic vegetatlo'n that provides males and 100 mm for females (Butlerprotection and substrate, and harbors 1960).food organisms for early instars(Butler 1956; Orcutt et al. 1975; The diet of juvenile crabsStevens and Armstrong 1984, 1985). consists largely of fish, mollusks,Recently, shells of bivalves such as and crustaceans (Butler 1954; GotshallMya arenaria and Crassostrea gigas 1977; Stevens 1982). In Grays Harbor,have been documented as very important Washington, first-year juvenileshabitat for young Dungeness crabs <60 -m CW feed primarily on small(Armstrong and Gunderson 1985). In mollusks and crustaceans. Second-central California there is evidence year crabs, 61-100 mm CW, feed onthat movement of postlarval Dungeness fish and prefer shrimp (Crangon spp.;crabs into the estuaries takes place Stevens et al. 1982). Fish also arein May and June via bottom currents, important to northern California crabswhere they stay for 11-15 months < 100 - CW according to Gotshall(Tasto 1983). Juveniles are common in (1977), but Butler (1954) reportedestuaries, while subadults and adults that crustaceans were the primary foodare common offshore. The importance among crabs of this size in the Queenof estuaries to juvenile Dungeness Charlotte Islands, British Columbia.crabs has been discussed in detail by Cannibalism among Dungeness crabs hasArmstrong and Gunderson (1985) and been noted by various authors (MacKay

7

Nw

1942; Butler 1954; Tegelberg 1972; Along the coast of northernGotshal1 1977; Stevens 1982; Stevens California, legal-sized and largeet al. 1982). Cannibalism was most sublegal-sized male crabs probablyprevalent among crabs < 60 mm CW which move offshore (often to the south orfed on smaller crabs of the same year north) in late summer, sometimesclass, probably during molting through early winter; sometime in(Stevens 1982; Stevens et al. 1982). winter the direction of movement isCannibalism is cited as a possible probably reversed and the crabs returncause of the dramatic population inshore. Interannual variation in thecycles characteristic of the Dungeness predominant direction of movement iscrab fishery (Botsford and Wickham considerable (Gotshall 1978).1978). Recently, Collier (1983) has shown a

random movement of adult crabs in theJuveniles are captured by a ocean. Many adult female crabs tagged

variety of demersal fishes in the off the coast of northern Californianearshore area with various flatfishes moved relatively little (about 2 km)(starry flounder, Platichthys after 1 year (Diamond and Hankinstellatus; English sole, Parophry 1985). Along the coast of southernvetulus; and rock sole, Lepidopsetta Washington, legal-sized malesbilineata) being the most important generally moved inshore and toward the(Reilly 1983b). Other predators on estuaries in fall (Barry 1985).juvenile crabs are lingcod (Ophiodonelongatus), cabezon (Scorpaenichthys Clams are the most important foodmarmoratus), wolf-eels (Anarrhichthys of adult Dungeness crabs > 151 mm CWocellatus), rockfish (Sebastes spp.), in northern California (Gotshall 1977)and octopus ( us dofleini) and > 166 mm CW in British Columbiaaccording to Waldrom (1958) and Orcutt (Butler 1954). Crustaceans and fish(1977). Predation on Dungeness crabs are valuable foods of the adultmay be seasonal in nature, as observed Dungeness crabs from both Similk Bayin white sturgeons, Acipenser and Grays Harbor, Washington (Mayertransmontanus (McKechnie and Fenner 1973; Stevens et al. 1982). Dungeness1971). Predation on Dungeness crabs crab populations are apparently notmay have a devastating impact as in limited by the abundance or scarcitythe case of sea otters (Enhyd of particular foods; they are somewhatlutris) in Orca Inlet, Alaska (Kimker nonspecific feeders which readily1985b . adjust to various foods (Gotshall

1977). They have developed anAdults evolutionary niche for feeding on

mud-sand substrate (Lawton and ElnerAt about 4 years old, most adult 1985).

Dungeness males in the coastal watersof Washington are of marketable size Crabs of different ages or sizes(> 159 -) (Cleaver 1949; Williams tend to eat different sizes or kinds1979). Marketable crabs usually only of food (Stevens 1982; Stevens et al.molt once a year (MacKay 1942). The 1982). According to Stevens et al.maximum lifespan of Dungeness crabs is (1982), crabs progress from eating8 to 10 years. The maximum size bivalves their first year, to eatingattained is about 218 mm CW in males shrimp (Crangon spp.) their secondand 160 - CW in females at the 16th year, and finally to eating juvenileinstar (MacKay 1942; Butler 1961b). teleost fish in the third year; these

shifts may be caused purely by changesAdult Dungeness crabs are found in mechanisms of food handling, or

primarily in the ocean but are they may have evolved to reducealso abundant in the inland waters competition among age groups ofof Washington and British Columbia. crabs. Crabs display a definite diel

8

activity; they are more abundant by generally, crabs are predicted today in the subtidal area and more reach maturity at the end of theirabundant at night in the intertidal second year after metamorphosis or inarea; the response is positively their third growing season over muchcorrelated with food availability of the coast (Butler 1961b; Cleaver(Stevens et al. 1984). Cannibalism is 1949). While age and size at sexualcommon among adults, but no maturity may not differ substantiallycorrelations have been made between along the coasts, estimates of growththe rate of cannibalism and abundance rates of newly settled 0+ crabs do.(Stevens 1982; Stevens et al. 1982).Sera stds of jvnl

growth rates indicate the process isGROWTH CHARACTERISTICS accelerated in estuaries or within

nearshore coastal embayments whereIn Dungeness crabs, like other water temperatures are relatively

crustaceans, growth proceeds in steps high. Stevens and Armstrong (1984)through a series of molts. The general studied growth of 0+ and 1+ juvenilesprocess of crustacean growth has been in Grays Harbor and found that 6described by Barnes (1974) and Warner months after metamorphosis (May to(1977). The number of molts that a October) 0+ crab averaged 40 -m incrab undergoes before becoming mature carapace width and by 1 year were 50depends upon the increment at each Mm. Crabs aged 0+ in Washingtonmolt and the frequency of molting, coastal estuaries may molt six toboth of which vary among crabs at eight times in the first summer 4different locations. Dungeness crabs growing season after which thegrow in carapace size at molt and gain frequency apparently declines with theweight between molts. In older crabs onset of winter and larger size. Athe growth, as measured by the percent dramatic indication of seasonal growthchange in carapace width, declines as was demonstrated where growth wasthe frequency of molting slows down, based on change in dry weight overbut the rate of weight gain of the time; rapid growth of the 0+ crabs incrabs increases over time. The Grays Harbor resulted in a 280-foldprobability of annual molting in increase in 14 months, and the growthfemale Dungeness crabs declines from rate of all age classes declinedabout 1.0 for crabs of 130-135 mm CW beginning in late summer of 1980.to 0. 0 for crabs of 155 -m CW and Rapid growth in early summer andlarger (Hankin et al. 1985). midsummer of 1980 was not repeated in

summer of 1981 (Figure 5).Among possible attributes of

estuarine residence suggested by Populations of 0+ crabs thatStevens and Armstrong (1984) is an settle directly offshore, as well asenhanced growth rate compared to that 1+ nearshore crabs, grow significantlyof siblings of a year class that slower than those in the estuarysettle offshore. Size attained by (Armstrong and Gunderson 1985).juvenile crabs within certain periods Young-of-the-year crabs in Graysafter metamorphosis seems to be Harbor grew from a first instar sizesomewhat dependent on latitude and on of 7 mm in May to a mean carapacetime of settlement. Extreme estimates width of 38.3 mm (sixth instar) byof age at sexual maturity range from October; 0+ crabs offshore had onlyas long as 4-5 years in British reached a mean width of 18.9 mm inColumbia (MacKay and Weymouth 1935) to November when they were a mixture of1 year in San Francisco Bay, where the third and fourth instars. Mean bottomcrabs reach a carapace width of 100 water temperatures in the estuarymm which is usually associated with were 15-16 0C during this time,sexual maturity (Tasto 1983). More while those offshore were 8.5-10 *C,

9

to o i 2 s..o75 - " .. .9 F

-4550,0

10 V

255- .. .1 ,.so44 20 ,+ ./.+

1.0 •

0 5 7 S 9 to i i

MON"-

Figure 6. Growth, based on meancarapace width, of 0+ crabs fromsettlement as 7-m first instars inMay through November. Bar = ± 1 SD.Grays Harbor population shown ascircles (GH). Offshore population

.02, shown as triangles (OS). Mean bottomwater temperatures at the time of

JJ A SO N 1 JU F M*A dJ-i sampling are also shown for some

1980 1981 samples (Armstrong and Gunderson1980985).

Figure 5. Changes in dry weight overtime of three year classes (1978,1979, and 1980) of Dungeness crabs InGrays Harbor, Washington. Growth However, in general, it is somewhatrates (- k) are expressed as g/g dry slower in the northern part of theweight per unit time. Note the rapid range (Washington and British Colum-growth of 0+ in the summer of 1980 and bia) when compared to the southernthe decline In growth of all year part of the range (California).classes in the fall of 1980 (Stevensand Armstrong 1985).

THE FISHERY

Commercial Fishery

which may account for the growth Commercial landings of Dungenessdifference (Figure 6). Young-of-the- crab on the Pacific coast haveyear juveniles offshore of San fluctuated widely, almost cyclically,Francisco Bay in the Gulf of the over the past 30 years (Figure 7) andFarallones also grow substantially have been reviewed by Armstrongslower than estuarine crabs (Tasto (1983). According to Peterson (1973),1983): the data show 0+ crabs at commercial landings were highest 1.5about 28-30m, while in the estuary a years after a period of stronggood proportion of this age group were upwelling in California and Oregon,up to 60 mm in width. and 6 months following a strong

upwelling in Washington, although theDungeness crab growth is quite biological sense of this conclusion is

variable along the Pacific coast. much in doubt. Botsford and Wickham

10

I,

15

1955 57 59 61 63 65 67 69 71 73 75 77 79 81 63

19n5 575516-6 6 1 9 7 1 3 7 7

D -'

0 1955 7 59 61 63 65 67 69 71 73 77 79 3

25 CALIORNI

955 5w S'q ;1I 63 69 71 73 75I 77

1955 57 59 61 63 65 67 69 71 73 77 79 3

Figure 7. Dungeness crab landings by season for individual PacificCoast States and the Province of British Columbia, 1955-83 (PacificMarine Fisheries Comission 19583).

(1975) challenged this conclusion by (Eaton 1985; Kimker 1985b; Koenemanusing auto correlation to show that 1985; Merritt 1985), British Columbiacommercial landings are cyclic but (Jamieson 1985), Washington (Barryupwelling is not. 1985), Oregon (Demory 1985), and

California (Oahlstrom and Wild 1983;Another theory to explain catch Warner 1985).

fluctuations suggests that periods ofhigh levels of cannibalism and/or Sport Fisheryinterspecific competition may cause adecline in the fishery 3 or 4 years Sport catch data are scarce andlater (Botsford and Wickham 1978). In according to Barry (1985) thea model predicting recruitment, Washington sport fishery on DungenessMcKelvey et al.(1980) discounted crabs amounts to less than onecannibalism as a factor and contended percent of the annual commercialthat changes in egg and larval harvest. Most of the available sportsurvival have regulated population catch data are from a survey reportedsuccess. Larval survival may be by Williams (1979). He revealedseriously altered by a combination of that from April through August 1974,environmental factors that can cause 471 crabs were taken intertidallyincreased mortality if unfavorable for at Mission Beach, Washington, by 735even short periods of time (Lough sport crabbers. April, May, and1976). Stevens and Armstrong (1981) June produced the best sport catchesindicate that diseases caused by with the highest average catchesvarious organisms (bacteria, Protozoa, occurring on low tides that rangedor fungi) may be responsible for mass from -0.60 to -0.74 m. Aerialmortalities of adult crabs. surveys made over Puget Sound beaches

using Williams' (1979) survey dataPredation may have a profound estimated that the beaches of 0

impact on the Dungeness crab Washington State probably supportedcommercial fishery in certain about 20,000 crabbers during thosegeographic areas (Kimker 1985b). months in 1974. In 1975, the sportReilly (1983b) theorizes that hatchery crab pot fishery alone (other sportreleased coho salmon from the Columbia catch methods are ring nets, dipRiver continue to suppress the nets, and hook and line) accountedDungeness crab fishery through for the harvest of about 300,000extensive predation. Dungeness crabs (Tegelberg 1976).

Only male crabs may be taken in theLandings of Dungeness crab from sport fishery (Figure 3). In

1954 to 1983 are broken down by State Washington there is a minimum sizeand Province in Figure 7. In general, of 6 or 6.25 inches carapace widththe landings in Washington (except for (depending on the area), measuredPuget Sound), Oregon, and California directly in front of the 10thfollow similar trends. Landings from anteriolateral spines. In OregonPuget Sound and British Columbia are the minimum size is 6.25 inches,lower but show less annual variation, measured similarly. The sport catchAlaska landings bear little relation is primarily found in Hood Canal,to other areas of the Pacific Puget Sound, and the major PacificNorthwest. The Alaska catch of 15.6 coast estuaries.million pounds in 1981 was a recordfor Alaska, but Washington coastal The State of Oregon has sought tocrab landings of 2.6 million pounds in limit the conflict between sport andthe same year were the lowest in 30 commercial crabbers by restrictingyears (Demory 1982). Recent reviews commercial crabbing to the middle ofof the commercial Oungeness crab the week and to the use of sport gearfishery have been published for Alaska (Demory 1985).

12 6~

V %

MUM III Il

ECOLOGICAL ROLE varied inversely with seawater

Dungeness crabs consume a wide temperatures of 9 to 17 °C (Figure 8).

variety of food organisms and are p Prolonged egg brooding periods in

to numerous predators. Crabs colder water are consistent with

contribute to sev ral trophic levels prolonged occurrences of ovigerous

as they progress? through successive crabs and cooler ocean temperatures as

life stages. The larvae largely you move progressively northward alongthe Pacific coast (Wild 1983).consume plankton (Lough 1976) and are Hatching success, considered as thepreyed upon by numerous fishes. Hacigscecoidrd sthAdults and juveniles are preyed upon number of larvae that hatch from anby sea otters fishes, and octopuses egg mass, decreased as the temperature(Bster s,54 fishes, and otpuss increased from 10 to 17 °C (Wild(Butler 1954; Waldrom 1958; Stevens 1983). Mayer (1973) found a similar1982; Reilly 1983b; Kimker 1985b). correlation between egg mortality and

Cannibalism is common and probably temperature with 20 mortality at

exercises some control over abundance. t0 wt 20O mortality at

In their various life stages, 10 OC and 100% mortality at 20 OC.

Dungeness crabs feed on a variety ofmollusks, crustaceans, and fishspecies (Stevens et al. 1982). Otherinformation on the ecological role of 13

each life stage is given in the lifehistory section.

ENVIRONMENTAL REQUIREMENTS

TemperatureThe temperature preferences of so •

adult crabs are different among -

seasons (Mayer 1973). They aresomewhat tolerant of abrupt

.temperature and salinity fluctuations(Cleaver 1949), and water temperatures o ,, , 13 ,1 is 16 17

from 3 to 19 °C were listed as normal TEMPt

for the Dungeness crab (Cleaver 1949). Figure 8. Dungeness crab egg brooding

Dungeness crabs have different periods at various laboratory seawater

optimal water temperatures at temperatures (Wild 1983).

different stages. In the laboratory,Des Voigne (1973) reported thatoptimal water temperatures for mating Optimal temperatures for larvaeranged from 12 to 16 °C during long are 10 to 14 OC. Juvenile crabs,photoperlods. Wild (1973) noted an 80 m wide and acclimated to 10.0 OC,apparent trend towards crabs mating have been exposed to water tempera-later in colder water in his tures up to 25.0 °C for 7 days withlaboratory experiments, but noted that little or no mortality (Des Voignemating took place between 10 and 1973); however, an increase of 2.5 0C

17 C. In Washington coastal waters, above 25 C was fatal to i 00 of all

where Dungeness crabs usually mate in crabs tested. In the laboratory,

early spring, the bottom temperatures crabs hd a m axium torlare between 8 and 10 OC (Armstrong, adult crabs had a maximum tolerablepers. com.. According to Wild temperature of 25 C during longp983., orn.). gAccoding torWid photoperiods, which decreased to 20 °C(1983), the egg brooding periods when exposed to short photoperiods

13

(Des Voigne 1973). With adult crabs temperatures, unfavorable salinitiesheld for 8 months, Wild (1983) resulted in complete mortality, butobserved that mortality increased with favorable salinities at unfavorabletemperature from 17% at 10 °C to 58% temperatures allowed some survivalat 13 °C and to 80% at 17 OC, although (Reed 1969). The most obvious effectlaboratory stress probably exacerbated on growth rate occurred at tempera-the effect of high temperatures. tures that resulted in the best

survival. Salinities that favored

Salinity survival generally had little effecton zoeal growth. Survival of zoeae is

Tolerance to salinity varies optimal between the water temperaturesamongetheanife state Dgess of 10.0 and 13.0 *C and salinities ofamong the life stages of the Dungeness 25 and 30 ppt (Reed 1969). The

crab. In general, salinity is not as significant interaction between tem-important as temperature to egg perature and salinity dictates cautiondevelopment and hatching, but the when making statements about eitherlarvae are highly sensitive to changes variable independent of the other one.in salinity (Buchanan and Milleman The effects of temperature or salinity1969). The percentage of eggs alone on C. magister zoeae do nothatching was optimum at 15 ppt, but appear to cause large fluctuations inhatching occurred to some degree over zoeal survival in the ocean (Reed

a wide range of salinities between 10 1969; sugh 1t7e).

ppt and 32 ppt (Buchanan and Milleman 1969; Lough 1976).

1969). When salinity was increased Substratefrom 15 ppt to 32 ppt, the averageprezoeal period was reduced from about Adult crabs are found living over60 min to less than 11 min. At a several substrate types (Schmitt 1921;salinity of 10 ppt, no prezoeae molted Cleaver 1949; Butler 1956), but theyto zoeaea, but 100% molted at 30 ppt prefer sandy-mud bottoms (Karpov(Buchanan and Milleman 1969). The 1982). Early juveniles prefer beds ofhighest survival for larvae was eelgrass, shell, or sandy mud (Stevensbetween salinities from 25 ppt and 30 and Armstrong 1984). This preferenceppt (Reed 1969). Survival decreased may stem from an abundance of foodwith salinity and was poorest at organisms on such substrates orsalinities of 15 ppt (Reed 1969). No perhaps the crabs find shelter fromjuvenile or adult tolerance levels are pehstecrb fid hler roavile in thel teratue ate is predation there (Stevens 1982). Older.available in the literature at this rs sem ls dpnen on.

time. crabs seem less dependent onepibenthic cover and can be found overmore exposed substrates. Most crabs

Temperature-Salinity Interactions remain in the subtidal environment,but may venture into littoral areas at

Salinity and temperature are both high tide (Stevens et al. 1984). Thisrelated to larval survival. Signifi- behavior is enhanced by the presencecant interaction exists between of preferred food items and decreasedthese two factors with salinity during low salinities following heavybuffering temperature. At favorable rains.

14.

. . . A

LITERATURE CITED

Armstrong, D.A. 1983. Cyclic crab fishery. Pages 33-36 in Proceedingsabundance and relationship to of the symposium on Dungeness crabenvironmental causes. Pages 102-110 biology and management. Universityin W. Wooster, ed. From year to of Alaska Sea Grant Rep. 85-3,year: interannual variability of Fairbanks.the environment and fisheries of theGulf of Alaska. University of Botsford, L.W., and D.E. Wickham.Washington Sea Grant Rep. WST-WO 1975. Correlation of upwelling83-3, Seattle. index and Dungeness crab catch.

U.S. Natl. Mar. Fish. Serv. Fish.Armstrong, D.A., and D.R. Gunderson. Bull. 73(4):901-907.

1985. The role of estuaries inDungeness crab early life history: a Botsford, L.W., and D.E. Wickham.case study in Grays Harbor, Washing- 1978. Behavior of age-specific,ton. Pages 145-170 in Proceedings of density-dependent models and thethe symposium on Dungeness crab northern California Dungeness crabbiology and management. University (Cancer magister) fishery. J. Fish.of Alaska Sea Grant Rep. 85-3, Res. Board Can. 35(6):833-843.Fairbanks.

Armstrong, D.A., B.G. Stevens, and Buchanan, D.W., and R.E. Milleman.J.C. Hoeman. 1981. Distribution 1969. The prezoeal stage of theand abundance of Dungeness crab and Dungeness crab, Cancer magisterCrangon shrimp, and dredging-related Dana. Biol. Bull. (Woods Hole)mortality of invertebrates and fish 137(2): 250-255.in Grays Harbor, Washington. U.S.Army Corps of Engineers, Seattle. Butler, T.H. 1954. Food of theTech. Rep. Contract No. DACW67-80-C- commercial crab in the Queen0086. Charlotte Islands region. Fish. Res.

Board Can. Prog. Rep. 99:3-5.Armstrong, D.A., B.G. Stevens, and

J.C. Hoeman. 1982. Distribution Butler, T.H. 1956. The distributionand abundance of aungeness crab and and abundance of early larval stagesCrangon shrimp, and dredging-related of the British Columbia commercialmortality of invertebrates and fish crab. Fish. Res. Board Can. Prog.in Grays Harbor, Washington, U.S. Rep. 107:22-23.Army Corps of Engineers, Seattle.Tech. Rep. Contract No. N-NACW67-80-C0086. 349 pp. Butler, T.H. 1960. Maturity and

breeding of the Pacific edible crab,Barnes, R.D. 1974. Invertebrate Cancer magister Dana. J. Fish. Res.

zoology. W.B. Saunders Co., Board Can. 17(5):641-646.Philadelphia, Pa. 870 pp. Butler, T.H. 1961a. Records of

Barry, S. 1985. Overview of the decapod Crustacea from BritishWashington coastal Dungeness crab Columbia. Can. J. Zool. 39(1):59-62.

15

Butler, r.H. 1961b. Growth and age Des Voigne, D.M. 1973. The influencedetermination of the Pacific edible of temperature and photoperiod uponcrab, Cancer magister Dana. J. Fish. the Dungeness crab, Cancer magister.Res. oard Zan. 18(5):873-889. Ph.D. Dissertation. University of

Washington, Seattle. 84 pp.Carlton, J.T., and A.M. Kuris. 1975.

Keys to decapod crustacea. Pages Diamond, N., and D.G. Hankin. 1985.385-412 in R.I. Smith and J.T. Movements of adult female DungenessCarton, eds. Light's manual: crabs (Cancer magister) in northernintertidal invertebrates of the California based on tag recoveries.central California coast. Can. J. Fish. Aquat. Sci.University of California Press, 42:919-926.Berkley.

Eaton, M.F. 1985. Kodiak IslandCleaver, F.C. 1949. Preliminary commercial Dungeness, Cancer

results of the coastal crab (Cancer magister, fishery. Pages 97-118 inmagister) investigation. Wash. State Proceedings of the symposium onDep. Fish., Olympia. Biol. Rep. Dungeness crab biology and49A:74-82. management. University of Alaska

Sea Grant Rep. 85-3, Fairbanks.Collier, P.C. 1983. Movement and

growth of post-larval Dungeness Edwards, E. 1966. Mating behavior incrabs, Cancer magister, in the San the European edible crab (CancerFrancisco area. Pages 125-133 in pagurus L.). Crustaceana 10(1):P.W. Wild and R.N. Tasto, eds. LiTe 23-30.history, environment, andmariculture studies of the Dungeness Fisher, W.S. 1976. Relationships ofcrab, Cancer magister, with emphasis epibiotic fouling and mortalities ofon the central California fishery eggs of the Dungeness crab (Cancerresource. Calif. Dep. Fish Game Fish magister). J. Fish. Res. Board Can.Bull. 172. 33(12):2849-2853.

Dahlstrom, W.A., and P.W. Wild. 1983. Fisher, W.S., and D.E. Wickham.A history of Dungeness crab 1976. Mortalities and epibioticfisheries in California. Pages 7-24 fouling of eggs from wildin P.W. Wild and R.N. Tasto, eds. populations of the Dungeness crab,U'Tfe history, environment, and Cancer magister. U.S. Natl. Mar.mariculture studies of the Dungeness Fish. Serv. Fish. Bull.crab, Cancer magister, with 74(1):201-207.emphasis on the central Californiafishery resource. Calif. Dep Fish Fisher, W.S., and D.E. Wickham. 1977.Game Fish Bull 172. Egg mortalities in wild populations

of the Dungeness crab in central andDemory, D. 1982. Dungeness crab north California. U.S. Natl. Mar.

fishery. 1980-81. Pac. Mar. Fish. Fish. Serv. Fish. Bull.Comm. Annu. Rep. 34:32-33. 75(1):235-237.

Demory, D. 1985. An overview of Gotshall, D.W. 1977. Stomach -,

Oregon Dungeness crab fishery with contents of northern Calforniamanagement concept for the future. Dungeness crabs (Cancer magister).Pages 27-32 in Proceedings of the Calif. Fish. Game 63(i):43-51.symposium on "ungeness crab biologyand management. University of Gotshall, D.W. 1978. NorthernAlaska Sea Grant Rep. 85-3, California Dungeness crab, CancerFairbanks. magister, movements as shown by

16

tagging. Calif. Fish Game management. University of Alaska64(4):234-254. Sea Grant Rep. 85-3, Fairbanks.

Hankin, D.G., N. Diamond, and M. Mohr. Knudsen, J.W. 1964. Observations of1985. Molt increments, annual the reproductive cycles and ecologymolting probabilities and survival of the common Brachyura and crablikerates of adult female Dungeness Anomura of Puget Sound, Washington.crabs in northern California. Pages Pac. Sci. 18(1):3-33.189-206 in Proceedings of thesymposium on Dungeness crab biology Koeneman, T.M. 1985. A brief reviewand management. University of of the commercial fisheries forAlaska Sea Grant Rep. 85-3, Cancer magister in southeast AlaskaFairbanks. and Yakutat waters, with emphasis on

recent seasons. Pages 61-70 in

Hartnoll, R.G. 1969. Mating in the Proceedings of the symposium on

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Jamieson, G.S. 1985. The Dungeness Kozloff, E.N. 1974. Keys to thecrab, Cancer magister, fisheries of marine invertebrates of Puget Sound,British Columbia. Pages 37-60 in the S nrc e lgo, ndProceedingsthe San Juan Archipelago, andDungeness crab biology and adjacent regions. U. Wash. Press,

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Sea Grant Rep. 85-3, Fairbanks. Lawton, P., and R.W. Elner. 1985. 4.K.A. 1983. Effect of Feeding in relation to morphometrics

Karpov, type on Effect of within the genus Cancer:I"substrate type on survival and evolutionary and ecoT ~l

growth in high density communal codtions. age3780incultures of juvenile Dungeness considerations. Pages 357-380 incurs ofn juvniler. D ges Proceedings of the symposium oncrabs, Cancer m ister. Pages Dungeness crab biology and

311-318 in -P.W. Wild and R.N. Tasto, management. University of Alaska

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crab, Cancer maotster, with emphasis Lough, R.G. 1976. Larval dynamics ofon the central a fornia fishery the Dungeness crab, Cancer magister, ^ NFish Bull. 172. Oregon coast,

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Kimker, A. 1985a. Overview of thePrince William Sound management area MacKay, D.C.G. 1942. The PacificDungeness crab fishery. Pages 77-84 edible crab, Cancer magister. Fish.in Proceedings of the symposium on Res. Board Can. Bull. No. 62. 32ffungeness crab biology and pp.management. University of AlaskaSea Grant Rep. 85-3, Fairbanks. MacKay, D.C.G. 1943. Temperature and

the world distribution of crabs of -,

Kimker, A. 1985b. A recent history the genus Cancer. Ecology 24(1):of the Orca Inlet, Prince William 113-115.Sound Dungeness crab fishery withspecific reference to sea otter MacKay, D.C.G., and F.W. Weymouth.predation. Pages 231-241 in 1935. The growth of the PacificProceedings of the symposiu edible crab, Cancer magister Dana.on Dungeness crab biology and J. Biol. Board Can. i{3)Y1I.D1-212.

17

..N• 1E

Mayer, 0.L. 1973. The ecology and 1981-82. Pages 31-32 in R.G.thermal sensitivity of the Dungeness Porter, ed. Thirty-fifth annualcrab, Cancer magister, and related report of the P.M.F.C. for the yearspecies of its benthic community in 1982. Pacific Marine FisheriesSimilk Bay, Washington. Ph.D. Commission, Portland, Oreg.Dissertation. University ofWashington, Seattle. 188 pp. Peterson, W.T. 1973. Upwelling

indices and annual catches ofMcKechnie, R.J., and R.B. Fenner. Dungeness crab, Cancer magister,

1971. Food habits of the white along the west coast of the Unitedsturgeon, Acipenser transontanus, States. U.S. Natl. Mar. Fish. Serv.in San Pablo and Suisun Bays, Fish. Bull. 22(3):902-910.California. Calif. Fish Game 57(3):209-212. Poole, R.L. 1966. A description of

McKelvey, R., 0. Hankin, K. Yanosko, the laboratory-reared zoeae ofand C. Snygg. 1980. Stable cycles Cancer magister Dana, and megalopaeand C.ltsge 1980.utaent cycels: taken under natural conditionsin multistage recruitment models: (Decapoda, Brachura). Crustaceana

an application to the northern (1):83-97.

California Dungeness crab (Cancer 11(l):83-97.

magister) fishery. Can. J. Fish Poole, R. L. 1967. PreliminaryAquat.Sci. 37(12):2323-2345. results of the age and growth study

Mrritt, M.F. 1985. The lower Cook of the market crab, Cancer magister,Inlet Dungeness crab fishery from in California: the age and growth of

1964-1983. Pages 85-96 in Cancer magister in Bodega Bay.

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Poole, R.L., and D.W. Gotshall. 1965.Orcutt, H.G. 1977. Dungeness crab Regulations and the market crab

research program. Calif. Dep. Fish fishery. Outdoor Calif. 269:7-8.Game Mar. Res. Rep. No. 77-21. 55pp. Prince, E.D., and D.W. Gotshall. 1976.

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research program. Calif. Dep. Fish associated with an artificial reefGame Mar. Res. Rep. No. 75-16. 23 in south Humboldt Bay, California.pp. Calif. Fish Game 64(4):274-285.

Orcutt, H.G., R.N. Tasto, P.W. Wild,C.W. Haugen, and P.C. Collier. Reed, P.N. 1969. Culture methods and1975. Dungeness crab research effects of temperature and salinity

5. Calif. Dep. Fish Game Mar. on survival and growth of Dungenessprogram. crab (Cancer magister) larvae in theRes. Rep. No. 75-12. 77 pp. laboratory. J. Fish. Res. Board

Orcutt, H.G., R.N. Tasto, P.W. Wild, Can. 26(2):389-397.

C.W. Haugen, and E.E. Ebert. 1976.Dungeness crab research program. Reilly, P.N. 1983a. Dynamics ofCalif. Dep. Fish Game Mar. Res. Rep. the Dungeness crab, CancerNo. 76-15. 42 pp. magister, larvae off central and

northern California. Pages 57-84Pacific Marine Fisheries Commission. in P.W. Wild and R.N. Tasto, eds.

1983. Dungeness crab fishery, -Tfe history, environment, and

18

mariculture studies of the Dungeness the southern Washington coast. U.S.crab, Cancer maqister, with emphasis Natl. Mar. Fish. Serv. Fish. Bull.on the central California fishery 79(2):349-352.resource. Calif. Dep. Fish GameFish Bull. 172. Stevens, B.G., and D.A. Armstrong.

1984. Distribution, abundance and

Reilly, P.N. 1983b. Predation on growth of juvenile Dungeness crabs,

Dungeness crabs, Cancer magister, in Cancer gister, in Grays Harbor

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L-1fe history, environment, and 82(3):469-483.

mariculture studies of the Dungeness Stevens, B.G., and D.A. Armstrong.crab, Cancer magister, with emphasis 1985. B.g, andonth enra alfrna ihey 1985. Ecology, growth, andon th tr Califia fishery population dynamics of juvenileresource. Calif. Dep. Fish Game Fish Dungeness crab, Cancer magisterBull. 172. Dana, in Grays Harbor, Washington,

Reilly, P.N. 1985. Dynamics of 1980-81. Pages 119-134 inDungeness crab, Cancer magister, Proceedings of the symposium on

larvae off central and northern Dungeness crab biology and manage-

California. Pages 245-272 in ment. University of Alaska Sea

Proceedings of the symposium on Grant Rep. 85-3, Fairbanks.

Dungeness crab biology andmanagement. University of Alaska Stevens, B.G., D.A. Armstrong, and R.Sea Grant Rep. 85-3, Fairbanks. Cusimano. 1982. Feeding habits of

the Dungeness crab Cancer magister

Rudy, P., Jr., and L.H. Rudy. 1979. as determined by the index of rela-

Oregon estuarine invertebrate tive importance. Mar. Biol. (Berl.)

animals. Oregon Institute of Marine 72(1):135-145.

Biology, University of Oregon,Charleston. U.S. Fish Wildi. Serv. Stevens, B.G., D.A. Armstrong, andContract No. 79-111. 131 pp. J.C. Hoeman. 1984. Diel activity

of an estuarine population ofSchmitt, W.L. 1921. The marine Dungeness crabs, Cancer magister, indecapod crustacea of California. relation to feeding andUniv. Calif. Pub1. Zool. No. 23. environmental factors. J.470 pp. Crustacean Biol. 4(3):390-403.

Snow, C.D., and J.R. Neilsen. 1966.Premating and mating behavior of the Tasto, R.N. 1983. Juvenile DungenessDungeness crab (Cancer magister crab, Cancer magister, studies inDana). J. Fish. Res. Board Can. the San Francisco Bay area. Pages

23(9):1319-1323. 135-154 in P.W. Wild and R.N. Tasto,eds. Life history, environment, and

Stevens, B.G. 1982. Distribution, mariculture studies of the Dungeness

abundance, and food habits of the crab, Cancer magister, with emphasis

Dungeness crab, Cancer magister, in on the central California fishery

Grays Harbor, W-shiTngton. Ph.D. resource. Calif. Dep. Fish Game

Dissertation. University of Fish Bull. 172.Washington, Seattle. 213 pp. ,nTegelberg, H. 1972. Dungeness crab

Stevens, B.G., and D.A. Armstrong. study. Pages 5-16 in Annual1981. Mass mortality of female progress report, Wash. State Dep.Dungeness crab, Cancer magister, on Fish. Proj. No. 1-76-R.

19

J

Tegelberg, H.C. 1976. Crabbing in Wickham, D.E. 1979c. The relation-Washington. Wash. State Dep. Fish. ship between megalopae of theRep. 11 pp. Dungeness crab, Cancer magister, and

the hydroid, Velella veiell, andWaldrom, K.D. 1958. The fishery and its influence on abundance estimates

biology of the Dungeness crab of C. magister negalopae. Calif.(.Cancer sacister Dana) in Oregon Fisth-Gamn 65(3):184-186.

waters. Oreg. Fish. Comm. Contrib.No. 24: 1-43. Wickham, D.E.., R. Shieser, and A.

Schuur. 1976. Observations on theWarner, G.F. 1977. The biology of inshore population of Dungeness

crabs. Van Nostrand Reinhold Co., crabs in Bodega Bay. Calif. FishNew York. 202 pp. Game 62(1):89-92.

Warner, R.W. 1985. Overview of the Wild, P.W. 1983. The influence ofCalifornia Dungeness crab, Cancer seawater temperature on spawning,magister, fishery. Pages 11-8 n egg development, and hatchingProceedings of the symposium on success of the Dungeness crab,Dungeness crab biology and Cancer magister. Pages 197-214 inmanagement. University of Alaska .WW ld and R.N. Tasto, eds. LifeSea Grant Rep. 85-3, Fairbanks. history, environment, and

mariculture studies of the DungenessWickham, D.E. 1979a. Carcinonemertes crab, Cancer magister, with emphasis

errans and the fouling of eggs of on the central California fisherytheungenes; crab, Cancer magister. resource. Calif. Dep. Fish GameJ. Fish. Res. Board a-n.-36(1):1319- Fish Bull. 172.1324.

Williams, J.G. 1979. Estimation ofWickham, D.E. 1979b. Predation by intertidal harvest of Dungeness

the nemertean Carcinonemertes errans crab, Cancer magiste r, on Pugeton eggs of the Dungeness crab, Sound, Washington, beaches. U.S.Cancer magister. Mar. Biol. (Berl.) Natl. Mar. Fish. Serv. Fish. Bull.55(11: 45-53. 77(1): 287-292.

20

LIAM 2L

M&MW DOCUMENATION IL WONT NO- L. sr'a. Rtciolent' Accession Heo.PAGE Biological Report 82(11.63)*

4. 1MM Was SUIRM 5. Ilamae DateSpecies Profiles: Life Histories and Environmental August_____1986 _

Requirements of Coastal Fishes and Invertebrates (PacificgNorthwest)--Dungeness crab

GletB. Pauley 9 David A. Arstrongba and Thomas W. HeunaSPwenuOgnitinat.i.S. P*Vf Oftengatign aMe a"~ Aurda 10. Preiec/Tlask/Wert Unit iMe.a Washington Cooperative Fishery b School of Fisheries

Research Unit University of Washington I .Cna C rGetG o

University of Washington Seattle, WA 98195 (Seattle, WA 98195 GM2 senarln Oqaentlan NRno. andA*e"_______________

1I. Ty"e Of MOP*"t 4Ptted Covered

National Wetlands Research Center U.S. Army Corps of Engineers

U.S. Department of the Interior P.O. Box 631 14.Washington, DC 20240 Vicksburg, MS 39180

*U.S. Army Corps of Engineers Report No. TR EL-82-4ItMaa (,. Umift M10 Wards)

Species profiles are literature summaries of the taxonomy, morphology, range, lifehistory, and environmental requirements of coastal aquatic species. They are designed toassist in environmental impact assessment. The Dungeness crab (Cancer magister) is foundof f the coasts of Washington, Oregon, and southern British ColiiET, -as well as in theestuarine waters of this geographic area. It is a shellfish highly prized and soughtafter by both commercial and sport fishermen. In Washington and Oregon, only male crabsmay be retained by sport and commercial fishermen. Commuercial crab catches are highlyvariable from year to year, but the catches from Washington and Oregon follow a verysimilar pattern. The highest sport catches take place on low tides ranging from -0.60 to-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 watertemperature increases from 10 to 17 OC; the optimal temperature for larval crabs isbetween 10 and 14 OC. Salinity is not as important to egg development and hatching astemperature, but optimum hatching occurs at about 15 ppt.

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Temperature Feeding habitsEstuaries CrabsShellfish Life cyclesSalinity Temperature

Dungeness crab Ecological roleCancer magister Dana Environmental requirements -

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