QL 

155 

.S63 

no. 82 

11.93 

Llbraxy 

National Wetlands Research Center 

U, S. Fish and Wildlife Service 

700 Cajundome Boulevard 

Lafayette. La. 70506 

Biological Report 82(11.93) TR EL-82-4 

January 1989 

Species Profiles: Life Histories and 

Environmental Requirements of Coastal Fishes 

and Invertebrates (Pacific Northwest) 

GHOST SHRIMP AND BLUE MUD SHRIMP 

blue mud shrimp 

Fish and Wildlife Service 

Coastal Ecology Grou 

Waterways Experiment Statio 

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

Biological Report 82(11.93) 

TR EL-82-4 

January 1989 

Species Profiles: Life Histories and Environmental Requirements 

of Coastal Fishes and Invertebrates (Pacific Northwest) 

GHOST SHRIMP AND BLUE MUD SHRIMP 

Susanna Hornig 

Anita Sterling 

Stanley D. Smith 

U.S. Fish and Wildlife Service 

National Fishery Research Center 

Building 204, Naval Station 

Seattle, WA 98115 

Project Officer 

David Moran 

U.S. Fish and Wildlife Service 

National Wetlands Research Center 

1010 Gause Boulevard 

Slidell, LA 70458 

Performed for 

U.S. Army Corps of Engineers 

Waterways Experiment Station 

Coastal Ecology Group 

Vicksburg, MS 39180 

and 

U.S. Department of the Interior 

Fish and Wildlife Service 

Research and Development 

National Wetlands Research Center 

Washington, DC 20240 
DISCLAIMER 

The mention of trade names in this report does not constitute endorsement or 

recommendation for use by the Federal Government. 

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 may be cited as follows: 

Hornig, S., A. Sterling, and S.D. Smith. 1989. Species profiles: life 

histories and environmental requirements of coastal fishes and invertebrates 

(Pacific Northwest)--ghost shrimp and blue mud shrimp. U.S. Fish Wildl. Serv. 

Biol. Rep. 82(11.93). U.S. Army Corps of Engineers, TR EL-82-4. 14 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 

Metric to U.S. Customary 

Multiply !Y 

millimeters (mm) 0.03937 

centimeters (cm) 0.3937 

meters (m) 3.281 

meters (m) 0.5468 

kilometers (km) 0.6214 

kilometers (km) 0.5396 

To Obtain 

inches 

inches 

feet 

fathoms 

statute miles 

nautical miles 

square meters (m2) 10.76 square feet 

square kilometers (km2) 0.3861 square miles 

hectares (ha) 2.471 acres 

liters (1) 0.2642 gallons 

cubic meters (m3) 35.31 cubic feet 

cubic meters (m3) 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 

metric tons (t) 1.102 short tons 

kilocalories (kcal) 3.968 British thermal units 

Celsius degrees ("C) 1.8(Y) + 32 Fahrenheit degrees 

inches 

inches 

feet (ft) 

fathoms 

statute miles (mi) 

nautical miles (nmi) 

U.S. Customary to Metric 

25.40 

2.54 

0.3048 

1.829 

1.609 

1.852 

millimeters 

centimeters 

meters 

meters 

kilometers 

kilometers 

square feet (ft2) 0.0929 square meters 

square miles (mi2) 2.590 square kilometers 

acres 0.4047 hectares 

gallons (gal) 3.785 liters 

cubic feet (ft3) 0.02831 cubic meters 

acre-feet 1233.0 cubic meters 

ounces (oz) 28350.0 milligrams 

ounces (oz) 28.35 grams 

pounds (lb) 0.4536 kilograms 

pounds (lb) 0.00045 metric tons 

short tons (ton) 0.9072 metric tons 

British thermal units (Btu) 

Fahrenheit degrees (OF) 

0.2520 

0.5556 (OF 

iv 

32) 

kilocalories 

Celsius degrees 
l 

CONTENTS 

Page 

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

CONVERSION TABLE ........................................................ iv 

ACKNOWLEDGMENTS ......................................................... vi 

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

MORPHOLOGY AND IDENTIFICATION AIDS ...................................... : 

REASONS FOR INCLUSION IN SERIES ......................................... 

LIFE HISTORY ............................................................ 5 

Development of Eggs and Larvae ........................................ 6 

Postlarval Development ................................................ 

Habitat ............................................................... : 

GROWTH CHARACTERISTICS .................................................. 6 

THE FISHERY ............................................................. 

ECOLOGICAL ROLE ......................................................... : 

Food and Feeding Habits ............................................... 8 

Cooperation, Competition, and Predation ............................... 

ENVIRONMENTAL REQUIREMENTS .............................................. 1: 

LITERATURE CITED ........................................................ 13 

V 
ACKNOWLEDGMENTS 

We are grateful for the reviews and suggestions of David Buchanan of the 

Oregon Department of Fish and Wildlife and Jefferson Gonor of Oregon State 

University, and for the use of the illustrations by Paul and Lynn Rudy. 

vi 
GHOST SHRIMP AND BLUE MUD SHRIMP 

NOMENCLATURE/TAXONOMY/RANGE cephalothorax. Like that of other 

arthropods, this cephalothorax is 

covered by a carapace or exoskeleton 

of hard, chitinous material that is 

shed (molted) periodically to allow 

for growth. The gills are located in 

special chambers at the sides of the 

thorax under the carapace. The blue 

mud shrimp has a large rostrum (forward extension of the carapace) and 

cylindrical eye stalks; the ghost 

shrimp has no rostrum or a small one 

and flattened eye stalks. Both have 

external mouthparts (maxillipeds) and 

antennae. Hair-like structures cover 

much of the shrimps' bodies and serve 

such functions as receiving sensory 

stimuli, obtaining food, cleaning 

self, creating water currents, and 

cleaning and carrying eggs (MacGinitie 

1934). 

Scientific name . . . . . . . . . . Callianassa 

californiensis Dana 

Preferred common name . . . Ghost shrimp 

(Figure 1) 

Scientific name . . . . . . . . . . . . . Upogebia 

pugettensis (Dana) 

Preferred common name . . . . . . . Blue mud 

shrimp (Figure 2) 

Other common names . . . . . Crawfish, mud 

prawns, ghost shrimp (collectively), 

burrowina ShrimD (Stevens 1928); red 

ghost &imp ‘(c. californiensis; 

Phillips 1984); -orange mud shrimp 

(C. californiensis; MacGinitie 

1935); mud shrimp (LJ. pugettensis; 

Hedgpeth 1970). 

Class . . . . . . . . . . . . . . . . . . . . . . Crustacea 

Order . . . . . . . . . . . . . . . . . . . . . . . Decapoda 

Family . . . . . . . . . . . . . . . . Callianassidae 

Geographic range: The ghost shrimp is 

found in intertidal areas along the 

west coast of North America from 

Mutiny Bay, Alaska, to the mouth of 

the Tijuana River, San Diego County, 

California; MacGinitie (1934) and 

Ricketts and Calvin (1968) reported 

finding specimens as far south as El 

Estuario de Punto Banda, Baja California Norte, Mexico. The blue mud 

shrimp is found from southeastern 

Alaska to San Quentin Bay (Bahia de 

San Quentin) in Baja California 

Norte. The general distribution of 

the two species in the Pacific 

Northwest is identical (Figure 3). 

MORPHOLOGY AND IDENTIFICATION AIDS 

The head and thorax of the ghost and 

blue mud shrimps are united into a 

Both shrimps have five pairs of thoracic legs (periopoda). The first 

pair of legs may be slightly unequal 

and only somewhat pincerlike (subchelate), and the rest, simple as in 

the blue mud shrimp; or, the first 

pair may be very unequal and very 

pincerlike (chelate), the second pair 

also pincerlike, and the fifth pair 

somewhat pincerlike as in the ghost 

shrimp (Schmitt 1921). The asymmetry 

of the first pair of legs characteristic of the ghost shrimp is more 

pronounced in males, and the larger 

cheliped (pincer leg) may be on either 

the left or the right side (MacGinitie 

1934). 

Both shrimps have five pairs of 

leaflike abdominal appendages (pleopods) or swimmerets. They also have 

flattened tail appendages (uropods) 

adapted for swimming. The blue mud 

shrimp has a short, square telson 

1 
h 

first leg ’ ” 

I\ 

corne’a h l ‘ I 

’ eyestalk F \ .s~w*~’ rostrum r: 

Figure 1. Ghost shrimp male (a) showing morphology of leg pairs (actual total 

length of specimen from rostrum to telson is 5 cm (2 inches)) and (b) enlargement 

of head area (dorsal view). Reproduced with permission from Rudy and Rudy 1983 

(copyright Paul and Lynn Rudy). 

2 
, 

rostrum 

Figure 2. Ovigerous blue mud shrimp female (a) actual length from rostrum to 

telson (9 cm; 3.5 inches), (b) enlarged dorsal view of head, and (c) first-stage 

larval form (actual total length about 5 mm (0.2 inches)). Reproduced with permission from Rudy and Rudy 1983 (copyright Paul and Lynn Rudy). 

(terminal segment); the ghost shrimp 

has a longer, more pointed one. 

These two shrimps can be distinguished from each other on the basis 

of the differences in the first pair 

of legs and color. The blue mud 

shrimp is usually dirty blue-green 

and the ghost shrimp varies from white 

3 

to pink, red, and orange. The 

carapace of the ghost shrimp is often 

transparent enough to allow observation of its internal organs (Johnson 

and Snook 1955), making it an 

interesting study specimen. There are 

other Callianassa species besides the 

ghost shrimp on the west coast; however, only one, C. gigas, is similar 
L 

WASHINGTON 

OREGON 

CALIFORNIA 

KILOMETERS 

Figure 3. Map showing geographic distribution of ghost shrimp and blue mud 

shrimp in Pacific Northwest region in intertidal sand or mudflats of west coast 

bays and estuaries. 

4 
in distribution, habitat, and habits. 

Callianassa gigas is larger than the 

ghost shrimp (125-150 mm (5-6 inches) 

long). And although the females and 

juveniles of these two species are 

similar in appearance, the large 

cheliped of the C. gigas male is 

longer and narrower than that of the 

male ghost shrimp (Morris et al. 

1980). 

REASONS FOR INCLUSION IN THE SERIES 

Although the ghost shrimp and blue 

mud shrimp are harvested as bait along 

the entire west coast of the United 

States, they are considered by some 

to be important pests of commercial 

oyster-growing operations in the 

Pacific Northwest (Ricketts and Calvin 

1968; McCrow 1972; Buchanan et al. 

1985). They are believed to destabilize the substrate, smother oysters 

with debris, and drain off water 

(through their burrows) from diked 

oyster beds. 

Either species can alter the 

physical characteristics of the 

habitat it occupies and affect the 

composition of the intertidal infaunal 

;;;;;pity (Brenchley 1981;. Posey 

. The ghost shrimp (Figure 1) 

is the more active burrower of the two 

and more severely affects substrate 

consistency (Bird 1982). Both the 

blue mud shrimp (Figure 2) and the 

ghost shrimp are associated with a 

variety of commensal and parasitic 

species (MacGinitie 1930, 1934, 1935; 

MacGinitie and MacGinitie 1968; 

Rickets and Calvin 1968; Kozloff 

1973). The ghost shrimp is one of the 

most abundant residents of marine 

sloughs or bay mudflats on the west 

coast of North America (MacGinitie 

1934). 

LIFE HISTORY 

Both the ghost shrimp and the blue 

mud shrimp live in burrows in the 

intertidal sand or mudflats of west 

coast bays and estuaries. Entrances 

to ghost shrimp burrows may be 

observed in the center of small 

conical hills of sand and small 

pebbles; those of the blue mud shrimp 

are less conspicuous, with much 

smaller, or absent, surrounding hills 

of sand (Kozloff 1973). 

Members of the blue mud shrimp 

species nearly always live in maleand-female pairs; each pair inhabits 

a smooth-walled, permanent, branching 

burrow extending about 45 cm (18 

inches) below the surface. The burrow 

generally has several entrances, each 

about 1 cm (0.4 inches) in diameter. 

The diameter of the tunnel beyond the 

opening is too narrow to allow the 

shrimp to turn around; consequently, 

specially enlarged chambers are required. The smooth walls appear to be 

cemented with a secretion produced by 

the shrimp. The blue mud shrimp forms 

a "mud basket," with its first two 

pairs of legs, which it uses as a 

scoop to transport mud and build its 

burrow, and as a strainer to collect 

food. The third and fifth pairs of 

legs are used for walking, and the 

fourth pair is braced against the burrow walls. The tail-fan can be used 

to block the burrow tunnel so effectively that the flow of water is 

stopped; this may possibly be a protective maneuver to ward off attacks 

from the rear (MacGinitie 1934). The 

species feeds on detritus and plankton 

strained from seawater, which it 

forces through the burrow by using its 

four pairs of swimmerets (pleopods) 

(MacGinitie 1930). 

The ghost shrimp also inhabits 

burrows, but of a less permanent character since this species sifts most of 

its food directly from the substrate 

and tunnels almost constantly, reworking the sediment to a depth of about 

75 cm (30 inches) in search of food. 

Burrow structures of ghost shrimp 

are less consistent in pattern than 

those of blue mud shrimp; the ghost 
shrimp digs tunnels branching in all 

directions, forming complex burrows 

with various numbers of openings 

(MacGinitie 1934). The second and 

third pairs of legs are used for digging and the fourth and fifth for 

cleaning its appendages, gills, and 

back, and for cleaning and manipulating its eggs (MacGinitie 1934). The 

third, fourth, and fifth pairs of legs 

are used in walking; the fourth pair 

is extended outward against the burrow 

wall for support. The swimmerets of 

the ghost shrimp constantly circulate 

water through the burrow, facilitating 

respiration. Its tail-fan, like the 

blue mud shrimp's, can be used (probably protectively) to block the burrow. 

The large cheliped of the male is a 

weapon used in disputes over territory 

and during the mating season 

(MacGinitie 1934). 

Development of Eggs and Larvae 

Female ghost shrimp are ovigerous 

(capable of producing fertile eggs) 

throughout the year, but the principal 

spawning season is in late June and 

early July (MacGinitie 1935). Intensive breeding probably begins in 

spring, but ovigerous females may 

still be plentiful as late as August. 

Spring warming appears to be the trigger for egg development. Three to 

four broods are produced at about 

6-week intervals. The larvae develop 

as plankton in coastal waters through 

five zoeal stages, which are distinguishable from one another primarily 

on the basis of size (McCrow 1972). A 

total of 6-8 weeks is spent as 

nearshore oceanic plankton (through 

the five zoeal and one megalopal 

stage); zoeal larvae are usually released on ebb tides in June and July, 

and the first megalopae appear in early August. Recruitment to the estuary 

is probably facilitated by flood tides 

occurring in late summer and fall. 

Larval drifting during this planktonic 

phase very likely serves as a mechanism of genetic exchange among populations in different estuaries (Johnson 

and Gonor 1982). 

Blue mud shrimp females are known to 

be ovigerous in January, February, and 

part of March (MacGinitie 1935), but 

larval development of this species has 

not been studied extensively. 

Postlarval Development 

Juvenile ghost shrimp are presumed 

to metamorphose rapidly to a state 

adapted for life on the bottom just 

before recruitment to the estuary 

(Johnson and Gonor 1982). Mortality 

due to predation is probably substantial during the short period (minutes 

to hours) between the moment the organism drops to the substrate and its 

successful burrowing beneath the surface (MacGinitie 1934). The lifespan 

of the ghost shrimp has been variously 

estimated at 3-5 years (Bird 1982), 10 

years (MacGinitie 1935), and 15-16 

years (Ricketts and Calvin 1968). The 

blue mud shrimp is also believed to be 

relatively long-lived (MacGinitie 

1930). 

Habitat 

Both of these species are commonly 

found in intertidal areas of mixed 

sand and mud. The blue mud shrimp 

lives in muddier areas than does the 

ghost shrimp; observations with respect to tidal height preferences vary 

(Table 1). In Oregon estuaries, ghost 

shrimp were consistently found in 

tideflats closer to the ocean than 

were blue mud shrimp (Bird 1982). 

Both species are common residents of 

eelgrass beds in the Pacific Northwest 

(Phillips 1984). 

GROWTH CHARACTERISTICS 

Typically, length of adults is 

5-10 cm (2-4 inches) in the ghost 

shrimp and 7.5-10 cm (3-4 inches) in 

the blue mud shrimp (MacGinitie and 

MacGinitie 1968). However, length may 

reach 11.5 cm (4.5 inches) in the 

ghost shrimp and 15 cm (6 inches) in 

the blue mud shrimp (Morris et al. 

1980). MacGinitie ( 1930, 1935 ) 

6 

Y 
Table 1. Habitat preferences of ghost shrimp and blue mud shrimp reported at 

different locations. 

Location Ghost shrimp Blue mud shrimp Source 

Pacific NW Muddy sand 

Yaquina Bay, OR 0 to 1 ft 

Muddier sand 

0 to 1 ft 

Oregon estuaries Tideflats close Tideflats further 

to ocean from ocean 

N. California Sandier mid-tidal Lower, muddier 

areas flats 

Tomales Bay & 

Elkhorn Slough, CA 

Monterey Bay, CA 

Muddy sand 

Generally lower 

tidal areas; 

mixed sand and 

mud 

Softer mud 

Generally higher 

tidal areas; mud, 

sandy mud with 

clay 

Kozloff 1973 

Thompson and 

Pritchard 1969 

Bird 1982 

Hedgpeth 1970 

Smith and 

Carlton 1975 

MacGinitie 

1935 

reported finding the largest blue mud 

shrimp in the muddiest, least rocky 

areas. 

Ghost shrimp mature at 18-24 months 

and some reproductive females may be 

less than 3 cm (1.2 inches) long; blue 

mud shrimp take 3 or more years to 

mature and reproductive females 

exceed 6 cm (2.4 inches) in length. 

Estimated growth in length averages 

approximately 15-30 mm/yr (O-6-1.2 

inches/yr) in ghost shrimp and 18-26 

mm/yr (0.7-1.0 inches/yr) in blue mud 

shrimp (Bird 1982). Density within a 

ghost shrimp colony and the colony's 

location appear to influence both 

growth and size at sexual maturity; 

ghost shrimp in the less dense 

colonies closest to the ocean grow 

faster, and the females become sexually mature at larger sizes and produce 

more and larger eggs (Bird 1982). 

Densities of ghost shrimp have been 

estimated at 700-1,400/m2 (2.8-5.6 

million/acre) in Yaquina Bay, Oregon 

(McCrow 1972); 420-770/m2 (1.7-3.1 

million/acre) in high-density areas of 

Sand Lake Estuary, Oregon; and less 

than 300/m2 (1.2 million/acre) in 

other areas on the Oregon coast (Bird 

1982). Blue mud shrimp densities in 

Oregon estuaries range from 330 to 

660/m2 (1.3-2.7 million/acre) (Bird 

1982). Biomass of either species 

sometimes exceeds 2.0 kg/m2 (18,000 

lb/acre (wet weight)). 

THE FISHERY 

Ghost and blue mud shrimp are 

harvested by commercial bait fishermen 

and recreational fishermen in California, Oregon, and Washington. Peterson 

(1977) described a method used in 

southern California in which water is 

pumped into the substrate under pressure, forcing the animals out of their 

burrows; in the area he studied, harvest noticeably reduced the ghost 

shrimp population. In the Pacific 

Northwest, attempts have been made to 

7 
control the shrimp on commercial 

Japanese oyster (Crassostrea gigas) 

grounds with the insecticide Sevin 

(carbaryl). This pesticide has been 

used to control ghost and mud shrimp 

in Washington since 1963 (Washington 

Department of Fisheries and Washington 

Department of Ecology 1985), and although it has been used on oyster 

grounds in Oregon, such use is currently unlawful there (L. Fredd, 

Oregon Department of Fish and Wild- 

?ife, Portland, OR; pers. comm.). 

During its use in Oregon, bait fishermen noted ghost shrimp mortalities in 

untreated areas soon after nearby 

oyster grounds were sprayed (Buchanan 

et al. 1985). 

Washington oyster growers estimate 

that oyster production would drop 70%- 

80%, resulting in a $5 million annual 

loss in Pacific and Grays Harbor 

Counties, without ghost shrimp control 

(Washington Department of Fisheries 

and Washington Department of Ecology 

1985). However, questions have been 

raised about the effects of Sevin on 

other organisms, including the 

commercially important Dungeness crab 

(Cancer magister), and on the 

estuarine ecosystem as a whole 

(Lindsay 1961; Stewart et al. 1967; 

Buchanan et al. 1985). Although the 

blue mud shrimp is believed to disturb 

the sediment far less extensively than 

the ghost shrimp (Bird 1982), both 

have been the objects of control 

programs. 

ECOLOGICAL ROLE 

Food and Feeding Habits 

The ghost shrimp was once thought to 

feed exclusively by sifting organic 

detritus from the floor of its burrow 

through the hairs on the second and 

third pairs of legs, rejecting coarse 

material, and then ingesting the retained fine particles by the use of 

the maxillipeds (MacGinitie 1934). 

And although it is still thought 

to obtain most of its food in this 

manner, there is evidence that it also 

filters detritus and plankton from the 

water moving through its burrow as 

does the blue mud shrimp (Morris et 

al. 1980). Rejected material is 

deposited outside the burrow. 

Burrowing activity is heaviest in the 

upper 45-50 cm (18-20 inches), where 

the availability of food is greatest 

(MacGinitie 1934). The burrowing and 

feeding behavior of the ghost shrimp 

is vigorous enough to cause substantial alterations in surface sediment 

characteristics over time, decreasing 

organic content and shifting the particle size distribution upwards (Bird 

1982). Sediment in dense ghost shrimp 

beds often has a soft, quicksand quality (Posey 1985). The burrowing 

activity of both the ghost and blue 

mud shrimp aerates the subsurface soil 

(MacGinitie 1930, 1934). 

The blue mud shrimp is a suspension 

feeder, straining detrital particles 

and plankton from seawater kept moving 

through its burrow by the action of 

its swimmerets. To feed, the animal 

positions itself near a burrow 

entrance and increases the movement of 

the swimmerets to increase the current 

of seawater through the burrow. The 

third maxillipeds are used to periodically sweep the food particles collected into the animal's mouth. 

Particles that are too big are ejected 

(MacGinitie 1930). 

Cooperation, Competition, and 

Predation 

By aerating the subsurface sediment 

and digging burrows protected from 

most predators, ghost shrimp and blue 

mud shrimp provide an environment 

attractive to commensals. Commensal 

and parasitic species associated with 

these shrimp include a blind goby, 

three species of pea crabs, two 

species of clams, a copepod, a shrimp, 

polynoid worms, and isopods (see 

Table 2). 

Species that might compete with 

these shrimp for either food or space 

8 
Table 2. Commensal (c) and parasitic (p) species reported in burrows of ghost 

shrimp and blue mud shrimp (compiled from MacGinitie and MacGinitie 1968; 

Ricketts and Calvin 1968; Kozloff 1973). 

Species 

Found with Found with 

ghost shrimp blue mud shrimp 

Goby 

Clevelandia ios (c) In burrow In burrow 

Pea crabs 

Scleroplax granulata (c) 

Pinnixa franciscana (c) 

p. schmitti (c) 

Abundant in burrow Abundant in burrow 

Abundant in burrow -__ 

In burrow (rare) --- 

Clams 

Pseudopythina rugifera (c) ___ Underside of abdomen 

Cryptomya californica (c) Extends syphons Extends syphons 

Copepods 

Hemicyclops callianassae 

Clausidium vancouverense 

Shrimp 

Betaeus ensezadensis (c) 

Polynoid worms 

Hesperonoe spp. (c) 

Isopods 

(unidentified--p) 

into burrow 

On gills 

Underside carapace 

(common) 

into burrow 

On gills 

Under carapace 

On gills ___ 

In burrow In burrow 

Under carapace --- 

Phyllodurus abdominalis (p) ___ Underside of abdomen 

are rare in ghost shrimp colonies 

because of the continual reworking of 

the sediment by this species. Infauna 

are both more varied and more abundant 

in blue mud shrimp colonies because 

this species less severely affects the 

sediment structure (Bird 1982). 

Although ghost shrimp typically 

inhabit deep burrows, they are susceptible to predation by other animals 

because they sometimes venture outside 

their burrow entrances. Under test 

conditions, ghost shrimp spent over 

25% of the time within 2 cm of the 

burrow entrance; the shrimp were also 

9 

observed to move from one burrow to 

another and were often found with 

part of an appendage exposed above the 

surface (Posey 1985). 

The seaward boundary of dense shrimp 

beds coincided with a fourfold 

seaward increase in the density of the 

major predator, the Pacific staghorn 

sculpin (Leptocottus armatus) in Coos 

Bay, Oregon (Posey 1986b). Caging 

experiments in Coos Bay indicated that 

predation by this fish, which was most 

intense in summer, probably restricts 

the seaward distribution of ghost 

shrimp (Posey 1986b). 
Mud and ghost shrimps are sometimes 

killed by the leopard shark, Triakis 

semifasciata, and by the brown smoothhound shark, Mustelus henlei. The 

leopard shark, whose range extends 

north to Oregon, apparently can shovel 

or burrow into the substrate to prey 

on benthic species (Russo 1975). 

Dungeness crabs are known to eat ghost 

shrimp, but the shrimp does not appear 

to be a major component of the crab's 

diet (Stevens et al. 1982; Posey 

1985). Sea-run cutthroat trout (Salmo 

clarki clarki) also commonly eat ghost 

shrimp, but are not considered a major 

predator (Posey 1985). Posey also 

suggests that intertidally foraging 

birds may occasionally eat ghost 

shrimp. 

ENVIRONMENTAL REQUIREMENTS 

The optimal temperature range for 

ghost shrimp appears to be 10 to 

13 OC, depending on depth below the 

surface. Egg-bearing females seem to 

prefer the cooler water at the greater 

depths; immature specimens are found 

higher up in the burrow. Water temperature in ghost shrimp habitat in 

Yaquina Bay, Oregon, varies seasonally from 9 to 15 OC (McCrow 1972). 

Activity of ghost shrimp decreased 

slightly with increasing maximum daily 

air temperature in an outdoor aquarium 

(Posey 1987). 

Ghost shrimp tend to be most 

abundant at the seaward end of bays 

with substantial freshwater inflow 

(McCrow 1972) and tolerate salinities 

from about 25% to 125% the salinity 

of normal seawater (33 ppt). Blood 

salinity changes along with water 

salinity. In a laboratory test, 

salinities of 8-9 ppt were lethal to 

75%-100% of ghost shrimp (Posey 1987). 

Activity of ghost shrimp decreased 

with decreasing salinity between 33 

and 10 ppt (Posey 1987). The blue 

mud shrimp tolerates salinities as 

low as 10% that of seawater and regulates osmotically when salinity falls 

below 75% that of seawater (Morris 

et al. 1980). 

Oxygen availability is no doubt a 

limiting factor for all intertidal 

species, including the ghost and blue 

mud shrimps (MacGinitie 1935). Thompson and Pritchard (1969) measured oxygen levels in burrows in Yaquina Bay, 

Oregon, and found that during ebb 

tide, oxygen levels were occasionally 

zero. They also found that under laboratory conditions, the ghost shrimp 

could survive anoxia (lack of oxygen) 

for 5.7 days and the blue mud shrimp 

could survive for 3.3 days, far longer 

than they would normally be subjected 

to anoxia in the environment. 

Although the ghost shrimp has a 

lower normal metabolic rate and survives anoxia and hypoxia (low oxygen) 

better than does the blue mud shrimp, 

both appear to have respiratory 

adaptations that allow them to 

tolerate the low oxygen conditions 

under which they live. Laboratory 

experiments have shown that both 

species are able to lower their 

metabolic rates once oxygen levels 

become critically low. Additionally, 

studies of the ghost shrimp have 

demonstrated the following adaptations 

to hypoxia/anoxia: when oxygen levels 

become low, heart rate is lowered 

(Thompson and Pritchard 1969); a 

respiratory pigment, hemocyanin, 

liberates more bound oxygen to the 

tissues (Morris et al. 1980); and the 

shrimp is able to switch to an 

alternate, anaerobic metabolism 

(Pritchard and Eddy 1979; Morris et 

al. 1980). 

Clifton et al. (1984) studied the 

effect of spilled oil on ghost shrimp 

colonies in Willapa Bay, Washington. 

They concluded that small amounts of 

oil carried in on the tides and temporarily stranded in intertidal areas 

are unlikely to have a serious longterm impact. However, stranded oil 

that is buried by a subsequent 

deposition of oil-free sediment creates a barrier to burrowing activity 

10 
c 

. 

that can be expected to persist for nature, the ghost and blue mud shrimp 

years. They also concluded that the 

burrowing activity of the shrimp conappear to be an integral part of the 

nearshore environments. And fortutributes to the introduction of oil nately for the shrimp, their wideinto the sub-surface. spread distribution should allow them 

to sustain their populations despite 

Although their effects on the the current attempts to eliminate them 

environment are controversial in locally. 

11 

LITERATURE CITED 

. 

Bird, E.M. 1982. Population dynamics 

of thalassinidean shrimps and 

community effects through sediment 

modification. Ph.D. Dissertation. 

University of Maryland, College 

Park. 151 pp. 

Brenchley, G.A. 1981. Disturbance 

and cmunity structure: an 

experimental study of bioturbation 

in marine soft-bottom environments. 

J. Mar. Res. 39:767-790. 

Buchanan, D.V., D.L. Bottom, and D.A. 

Armstrong. 1985. The controversial 

use of the insecticide Sevin in 

Pacific Northwest estuaries: its 

effects on Dungeness crab, Pacific 

oyster, and other species. Pages 

401-407 in Proceedings of the 

symposium Tn Dungeness crab biology 

and management, Oct. 9-11, 1984, 

Anchorage, AK. Alaska Sea Grant 

Rep. No. 85-3, University of Alaska, 

Alaska Sea Grant College Program, 

Fairbanks. 

Clifton, H.E., K.A. Kvenvolden, and 

J.B. Rapp. 1984. Spilled oil and 

infaunal activity--modification of 

burrowing behavior and redistribution of oil. Mar. Environ. Res. 

ll:lll-136. i 

Hedgpeth, J.W. 1970. Introduction to 

seashore life of the San Francisco 

Bay region and the coast of northern 

California. University of California Press, Berkeley. 136 pp. 

Johnson, G.E., and J.J. Gonor. 1982. 

The tidal exchange of Callianassa 

californiensis (Crustacea, Decapoda) 

larvae between the ocean and 

the Salmon River Estuary, Oregon. 

Estuarine Coastal Shelf Sci. 14:501- 

516. 

Johnson, M.E., and H.J. Snook. 1955. 

Seashore animals of the Pacific 

coast. Dover Publications Inc., New 

York. 659 pp. 

Kozloff, E.N. 1973. Seashore life of 

Puget Sound, the Strait of Georgia, 

and San Juan Archipelago. 

University of Washington Press, 

Seattle. 282 pp. and 28 plates. 

Lindsay, C.E. 1961. Pesticide tests 

in the marine environment in the 

state of Washington. Proc. Natl. 

Shellfish. Assoc. 52:87-97. 

MacGinitie, G.E. 1930. The natural 

history of the mud shrimp Upogebia 

pugettensis (Dana). Ann. Mag. Nat. 

Hist., Ser. 10, 6:36-44. 

MacGinitie, G.E. 1934. The natural 

history of Callianassa californiensis Dana. Am. Midl. Nat. 15:166- 

177. 

MacGinitie, G.E. 1935. Ecological 

aspects of a California marine 

estuary. Am. Midl. Nat. 16:629-765. 

MacGinitie, G.E., and N. MacGinitie. 

1968. Pages 284-293 in Natural 

history of marine animaly, 2nd ed. 

McGraw-Hill, New York. 

McCrow. L.T. 1972. The ahost shrimp. 

Callianassa californiensis Dana; 

1854, in Yaquina Bay, Oregon. M.S. 

Thesis. Oregon State University, 

Corvallis. 56 pp. 

Morris, R.H., D.P. Abbott, and 

E.C. Haderlie. 1980. Intertidal 
invertebrates of California. Stanford University Press, Stanford, 

California. 690 pp. 

Peterson, C.H. 1977. Competitive 

organization of the soft-bottom 

macrobenthic communities of southern 

California lagoons. Mar. Biol. 

(Berl.) 43:343-359. 

Phillips, R.C. 1984. The ecology of 

eelgrass meadows in the Pacific 

Northwest: community profile. 

U.S. Fish W?ldl. Serv. FWS/OBS84/24. 85 pp. 

Posey, M.H. 1985. The effects upon 

the macrofaunal community of a 

dominant burrowing deposit feeder, 

Callianassa californiensis, and the 

role of predation in determining its 

intertidal distribution. Ph.D. 

Dissertation. University of Oregon, 

Eugene. 119 pp. 

Posey, M.H. 1986a. Changes in a 

benthic community associated with 

dense beds of a burrowing deposit 

feeder, Callianassa californiensis. 

Mar. Ecol. Prog. Ser. 31:15-22. 

Posey, M.H. 1986b. Predation on a 

burrowing shrimp: distribution and 

community consequences. J. Exp. 

Mar. Biol. Ecol. 103:143-161. 

Posey, M.H. 1987. Effects of lowered 

salinity on activity of the ghost 

shrimp Callianassa californiensis. 

Northwest Sci. (in press). 

Pritchard, A.W., and S. Eddy. 1979. 

Lactate- formation in Callianassa 

californiensis and Upogebia pugettensis (Crustacea: Thalassinidea). 

Mar. Biol. (Berl.) 50:249-253. 

important invertebrate animals. 

U.S. Fish Wildl. Serv. Biol. Serv. 

Program FWS/OBS-83/16. 225 pp. 

Russo, R.A. 1975. Observations on 

the food habits of leopard sharks 

(Triakis semifasciata) and brown 

smooth-hounds (Mustelus henlei). 

Calif. Fish Game 61:95-103. 

Schmitt, W.L. 1921. The marine 

decapod crustacea of California. 

Univ. Calif. Publ. Zool. 23:1-470. 

Smith, R., and J.T. Carlton. 1975. 

Light's manual: intertidal invertebrates of the central California 

coast. University of California 

Press, Berkeley. 716 pp. 

Stevens, B.A. 1928. Callianassidae 

from the west coast of North 

America. Publ. Puget Sound Biol. 

(Berl.) Stn. 6:315-369. 

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

Cusimano. 1982. Feeding habits of 

the Dungeness crab Cancer magister 

as determined by the Index of 

Relative Importance. Mar. Biol. 

72:132-145. 

Stewart, N.E., R.E. Millemann, and 

W.P. Breese. 1967. Acute toxicity 

of the insecticide Sevin and its 

hydrolytic product 1-naphthol to 

some marine organisms. Trans. Am. 

Fish Sot. 96:25-30. 

Thompson, R.K., and A.W. Pritchard. 

1969. Respiratory adaptations of 

two burrowing crustaceans, 

Callianassa caiiforniensis and 

Upogebia pu)gettensig:;ol~(Deca;~;~l 

Thalassinidea . 

(Woods Hole) 136:274-287. 

Ricketts, E.F., and J. Calvin. 1968. 

Between Pacific tides. Revised by 

J.W. Hedgpeth, Stanford University 

Press, Stanford, CA. 614 pp. 

Rudy, P., Jr., and L.H. Rudy. 1983. 

Oregon estuarine invertebrates: an 

illustrated guide to the common and 

Washington Department of Fisheries and 

Washington Department of Ecology. 

1985. Final environmental impact 

statement: use of the insecticide 

Sevin to control ghost and mud 

shrimp on oyster beds in Willapa Bay 

and Grays Harbor. 82 pp. plus 

appendices. 

14 
c 

l 

0272 -101 

REPORT GOCUMENTATION 1. .RE’-=-. NO- 2. 3. RKiPient’s Accession No. 

PAGE Biological Report 82(11.93)* 

@. Title l n! Subtitle 

Species Profiles: Life Histories and Environmental Requirements of 

5. R*wfi Da10 

Coastal Fishes and Invertebrates (Pacific Northwest)--Ghost Shrimp _a 

January 1989 

and Blue Mud Shrimp 

I. Au(horB) a PerforminS Organization fkpt. N, 

Susanna Hornig, Anita Sterling, and Stanley D. Smith 

9. Pwforming Organization Name and Address 10. PmieCt/l~sh/WorL Unit No. 

U.S. Fish and Wildlife Service 

National Fishery Research Center 11. Contract(C) or Grant(G) No. 

Building 204, Naval Station Puget Sound 

Seattle, WA 98115-5007 

(C) 

-- 

12. SpwwwinS OrSanlUtlon Name and Addnss 

(0) 

U.S. Department of the Interior U.S. Army Corps of Engineers 13. TYly of RwJott 6 Pwiod Cover. 

Fish and Wildlife Service Waterways Experiment Station 

National Wetlands Research Center P.O. Box 631 

Washington, DC 20240 Vicksburg, MS 39180 

F 

14. 

IS. Supplwnentary Notes 

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

Ia Abenet (Limit: 200 words) 

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

history, and environmental requirements of coastal aquatic species. The profiles are 

prepared to assist in environmental impact assessments. The ghost shrimp 

(Callianassa californiensis) and blue mud shrimp (Upogebia pugettensis) are common 

residents of intertidal mudflats of the Pacific Northwest, as well as of the entire 

west coast of the contiguous United States. These species are decapod crustaceans, 

but not true shrimp. They are harvested as bait by recreational and commercial 

oyster-growing operations. Ghost shrimp larvae develop in summer in nearshore coastal 

waters and settle to the substrate surface, where they rapidly metamorphose; the life 

cycle of the blue mud shrimp is presumed to be similar. Both species spend their 

lives in burrows in the mudflat, where the ghost shrimp is primarily a deposit feeder 

and the blue mud shrimp is a suspension feeder. 

It. Dowm*nt Analysis a. D*scrip~on 

Fishery Salinity Sediments 

Growth Temperature Life Cycles 

Estuaries Oxygen Shrimps 

Feeding habits Depth Oil spills 

b. Identlfl8n/~.Ewhd Tarns 

Ghost Shrimp Callianassa californiensis Dana 

Predators 

Competitors 

Commensals 

Blue Mud Shrimp Upogebia pugettensis (Dana) 

c. COSATI FieldlGmup 

19. Security Class (This Repwt) 21. No. of PaIUnclassified 14 

Unlimited release 

. .._. ___ _- 

20. Security Clas.8 This Pace) 

Unclasslfled 

P. Price 

--.-.... -______ ._ . 
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the Interior has responsibility for most of our nationally owned 

public lands and natural resources. This includes fostering the 

wisest use of our land and water resources, protecting our fish 

and wildlife, preserving the environmental and cultural values of our 

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our energy and mineral resources and works to assure that their 

development is in the best interests of all our people. The Department also has a major responsibility for American Indian reservation 

communities and for people who live in island territories under U.S. 

administration. 

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FISH AND WILDLIFE SERVICE 

US I . 

TAKE PRIDE 

in America 

UNITED STATES 

DEPARTMENT OF THE INTERIOR 

FISH AND WILDLIFE SERVICE 

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