Biological Report 82 1.93Y LU L-82-4,t 93January 1989

Species Profiles: Life Histories andEnvironmental Requirements of Coastal Fishes

r o and Invertebrates (Pacific Northwest)

o GHOST SHRIMP AND BLUE MUD SHRIMP0

ghost shrimp

: blue mud shrimp

" APR 1989

VE

Coastal Ecology Group* Fish and Wildlife Service Waterways Experiment Station

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

__ __ _ __ _ C_ _ 8 9

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

January 1989

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

GHOST SHRIMP AND BLUE MUD SHRIMP

by

Susanna HornigAnita SterlingStanley D. Smith

U.S. Fish and Wildlife ServiceNational Fishery Research Center

Building 204, Naval StationSeattle, WA 98115

Aes :n- For -

Project Officer r F TDavid Moran D'I(T 7.7

U.S. Fish and Wildlife Service dNational Wetlands Research Center .1it fI-----4

1010 Gause BoulevardSlidell, LA 70458

Distribution/

Performed for Avalability Codes

U.S. Army Corps of Engineers and/orWaterways Experiment Station !Dist Special

Coastal Ecology GroupVicksburg, MS 39180

andAr

U.S. Department of the InteriorFish and Wildlife ServiceResearch and Development

National Wetlands Research CeoterWashington, DC 20240

DISCLAIMER

The mention of trade names in this report does not constitute endorsement orrecommendation for use by the Federal Government.

0

This series may be referenced as follows:

U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life historiesand environmental requirements of coastal fishes and invertebrates. U.S. FishWildl. 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: lifenhstorieb 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.

0

PREFACE

---his 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 addo4 as they are prepared.This project is jointly planned and financed by the U.S. Army Corps of Engineersand the U.S. Fish and Wildlife Service.

Suggestions or questions regarding this report should be directe to one ofthe following addresses.

Information Transfer Specialist K?)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

0iii

CONVERSION TABLE

Metric to U.S. Customary

Multiply To Obtain

millimeters (mm) 0.03937 inchescentimeters (cm) 0.3937 inchesmeters (in) 3.281 feetmeters (m) 0.5468 fathomskilometers (km) 0.6214 statute mileskilometers (km) 0.5396 nautical miles

square meters (m2) 10.76 square feet

square kilometers (km2) 0.3861 square mileshectares (ha) 2.471 acres

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

milligrams (mg) 0.00003527 ouncesgrams (g) 0.03527 ounceskilograms (kg) 2.205 poundsmetric tons (t) 2205.0 poundsmetric tons (t) 1.102 short tons

kilocalories (kcal) 3.968 British thermal units

Celsius degrees (°C) 1.8(0 C) + 32 Fahrenheit degrees

U.S. Customary to Metric

inches 25.40 millimetersinches 2.54 centimetersfeet (ft) 0. 3048 metersfathoms 1.829 metersstatute miles mi) 1.609 kilometersnautical miles (nmi) 1.852 kilometers

square feet (ft2) 0.0929 square meterssquare miles (mi2) 2.590 square kilometersacres 0.4047 hectares

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

ounces (oz) 28350.0 milligramsounces (oz) 28.35 gramspounds (lb) 0.4536 kilogramrspounds (1b) 0.0045 metric tonsshort tons (ton) 0.9072 metric tons

British thermal units (Btu) 0.2520 kilocaloriesFahrenheit degrees (OF) 0.5556 (OF - 32) Celsius degrees

iv4

CONTENTS

Page

PREFACE ................................................................. iiiCONVERSION TABLE ........................................................ ivACKNOWLEDGMENTS ......................................................... vi

NOMENCLATURE/TAXONOMY/RANGE ............................................. 1MORPHOLOGY AND IDENTIFICATION AIDS ...................................... 1REASONS FOR INCLUSION IN SERIES ......................................... 5LIFE HISTORY ............................................................ 5Development of Eggs and Larvae ........................................ 6Postlarval Development ................................................ 6Habitat ............................................................... 6

GROWTH CHARACTERISTICS .................................................. 6THE FISHERY ............................................................. 7ECOLOGICAL ROLE ......................................................... 8

Food and Feeding Habits ............................................. 8Cooperation, Competition, and Predatio ................................ 8

ENVIRONMENTAL REQUIREMENTS .............................................. 10

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

v

ACKNOWLEDGMENTS

We are grateful for the reviews and suggestions of David Buchanan of theOregon Department of Fish and Wildlife and Jefferson Gonor of Oregon StateUniversity, 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 otherarthropods, this cephalothorax iscovered by a carapace or exoskeleton

Scientific name .......... Callianassa of hard, chitinous material that iscaliforniensis Dana shed (molted) periodically to allow

Preferred common name ... Ghost shrimp for growth. The gills are located in(Figure 1) special chambers at the sides of the

Scientific name ............. Upogebia thorax under the carapace. The bluepugettensis (Dana) mud shrimp has a large rostrum (for-

Preferred common name ....... Blue mud ward extension of the carapace) andshrimp (Figure 2) cylindrical eye stalks; the ghost

shrimp has no rostrum or a small oneOther common names ..... Crawfish, mud and flattened eye stalks. Both have

prawns, ghost shrimp (collectively), external mouthparts (maxillipeds) andburrowing shrimp (Stevens 1928); red antennae. Hair-like structures coverghost shrimp (C. californiensis; much of the shrimps' bodies and servePhillips 1984); orange mud shrimp such functions as receiving sensory(C. californiensis; MacGinitie stimuli, obtaining food, cleaning1§35); mud shrimp (U. pugettensis; self, creating water currents, andHedgpeth 1970). cleaning and carrying eggs (MacGinitie

Class ...................... Crustacea 1934).. Order ....................... Decapoda

Family ................ Callianassidae Both shrimps have five pairs of tho-- racic legs (periopoda). The first

Geographic range:--,The ghost shrimp is pair of legs may be slightly unequalfound in intertidal areas along the and only somewhat pincerlike (sub-west coast of North America from chelate), and the rest, simple as inMutiny Bay, Alaska, to the mouth of the blue mud shrimp; or, the firstthe Tijuana River, San Diego County, pair may be very unequal and veryCalifornia;- MacGinftie (1914 _nd pincerlike (chelate), the second pairRicketts and Calvin (1968) reported also pincerlike, and the fifth pairfinding specimens as far south as El somewhat pincerlike as in the ghostEstuario de Punto Banda, Baja Cali- shrimp (Schmitt 1921). The asymmetryfornia Norte, Mexico. The blue mud of the first pair of legs character-shrimp is found from southeastern istic of the ghost shrimp is moreAlaska to San Quentin Bay (Bahia de pronounced in males, and the largerSan Quentin) in Baja California cheliped (pincer leg) may be on eitherNorte. The general distribution of the left or the right side (MacGinitiethe two species in the Pacific 1934).Northwest is identical (Figure 3).

1 Both shrimps have five pairs ofleaflike abdominal appendages (pleo-

MORPHOLOGY AND IDENTIFICATION AIDS pods) or swimmerets. They also haveflattened tail appendages (uropods)

The head and thorax of the ghost and adapted for swimming. The blue mudblueuwd shrimps are united into a shrimp has a short, square telson

r- r\ y, C. C 0

A Ck V\016A C>~

LA.

a.

largecheliped

first log rostrum

corn ays5 mm

pleopods

uropods' "

telson

Figure 1. Ghost shrimp male (a) showing morphology of leg pairs (actual totallength of specimen from rostrum to telson is 5 cm (2 inches)) and (b) enlargementof head area (dorsal view). Reproduced with permission from Rudy and Rudy 1983(copyright Paul and Lynn Rudy).

2

rostruma. rotuobdomen

eg tels

wlnlgpleopods ods

b.

i C.

V

Figure 2. Ovigerous blue mud shrimp female (a) actual length from rostrum totelson (9 cm; 3.5 inches), (b) enlarged dorsal view of head, and (c) first-stagelarval form (actual total length about 5 mm (0.2 inches)). Reproduced with per-mission from Rudy and Rudy 1983 (copyright Paul and Lynn Rudy).

(terminal segment); the ghost shrimp to pink, red, and orange. Thehas a longer, more pointed one. carapace of the ghost shrimp is often

transparent enough to allow observa-These two shrimps can be distin- tion of its internal organs (Johnson

guished from each other on the basis and Snook 1955), making it anof the differences in the first pair interesting study specimen. There areof legs and color. The blue mud other Callianassa species besides theshrimp is usually dirty blue-green ghost shrimp on the west coast; how-and the ghost shrimp varies from white ever, only one, C. gigas, is similar

3

SrAVNC 1,R CANADA

SSEATTLE

GRAYS* PUGE UNDHARBOR%.

WASHINGTON

* -04

PORTLAND £M8qRIVER

MILES0 0 100

0 so 100K ILOMAET7ERS

BAY

OREGON

CALIFORN IA ---HUMBOLDt

BA Y....

Figure 3. Map showing geographic distribution of ghost shrimp and blue mudshrimp in Pacific Northwest region in intertidal sand or mudflats of west coastbays and estuaries.

I in distribution, habitat, and habits. intertidal sand or mudflats of westCallianassa gigas is larger than the coast bays and estuaries. Entrancesghost shrimp (125-150 mm (5-6 inches) to ghost shrimp burrows may belong). And although the females and observed in the center of smalljuveniles of these two species are conical hills of sand and smallsimilar in appearance, the large pebbles; those of the blue mud shrimpcheliped of the C. gigas male is are less conspicuous, with muchlonger and narrower than that of the smaller, or absent, surrounding hillsmale ghost shrimp (Morris et al. of sand (Kozloff 1973).1980).

Members of the blue mud shrimp

REASONS FOR INCLUSION IN THE SERIES species nearly always live in male-and-female pairs; each pair inhabitsa smooth-walled, permanent, branching

Althouqh the ghost shrimp and blue burrow extending about 45 cm (18mud shrimp are harvested as bait along inches) below the surface. The burrowthe entire west coast of the United generally has several entrances, eachStates, they are considered by some about 1 cm (0.4 inches) in diameter.to be important pests of commercial The diameter of the tunnel beyond theoyster-growing operations in the opening is too narrow to allow thePacific Northwest (Ricketts and Calvin shrimp to turn around; consequently,1968; McCrow 1972; Buchanan et al. specially enlarged chambers are re-1985). They are believed to destabi- quired. The smooth walls appear to belize the substrate, smother oysters cemented with a secretion produced bywith debris, and drain off water the shrimp. The blue mud shrimp forms(through their burrows) from diked a "mud basket," with its first twooyster beds. pairs of legs, which it uses as a

scoop to transport mud and build itsEither species can alter the burrow, and as a strainer to collect

physical characteristics of the food. The third and fifth pair , ofhabitat it occupies and affect the legs are used for walking, and thecomposition of the intertidal infaunal fourth pair is braced against the bur-community (Brenchley 1981; iosey row walls. The tail-fan can be used1986a). The ghost shrimp (Figure 1) to block the burrow tunnel so effec-is the more active burrower of the two tively that the flow of water isand more severely affects substrate stopped; this may possibly be a pro-consistency (Bird 1982). Both the tective maneuver to ward off attacksblue mud shrimp (Figure 2) and the from the rear (MacGinitie 1934). Theghost shrimp are associated with a species feeds on detritus and planktonvariety of commensal and parasitic strained from seawater, which itspecies (MacGinitie 1930, 1934, 1935; forces through the burrow by using itsMacGinitie and MacGinitie 1968; four pairs of swimmerets (pleopods)Rickets and Calvin 1968; Kozloff (MacGinitie 1930).1973). The ghost shrimp is one of themost abundant residents of marinesloughs or bay mudflats on the west The ghost shrimp also inhabitscoast of North America (MacGinitie burrows, but of a less permanent char-

1934). acter since this species sifts most ofits food directly from the substrateand tunnels almost constantly, rework-

LIFE HISTORY ing the sediment to a depth of about75 cm (30 inches) in search of food.Burrow structures of ghost shrimp

Both the ghost shrimp and the blue are less consistent in pattern thanmud shrimp live in burrows in the those of blue mud shrimp; the ghost

5

shrimp digs tunnels branching in all Blue mud shrimp females are known todirections, forming complex burrows be ovigerous in January, February, andwith various numbers of openings part of March (MacGinitie 1935), but(MacGinitie 1934). The second and larval development of this species hasthird pairs of legs are used for dig- not been studied extensively.ging and the fourth and fifth forcleaning its appendages, gills, and Postlarval Developmentback, and for cleaning and manipulat-ing its eggs (MacGinitie 1934). The Juvenile ghost shrimp are presumedthird, fourth, and fifth pairs of legs to metamorphose rapidly to a stateare used in walking; the fourth pair adapted for life on the bottom justis extended outward against the burrow before recruitment to the estuarywall for support. The swimmerets of (Johnson and Gonor 1982). Mortalitythe ghost shrimp constantly circulate due to predation is probably substan-water through the burrow, facilitating tial during the short period (minutesrespiration. Its tail-fan, like the to hours) between the moment the or-blue mud shrimp's, can be used (proba- ganism drops to the substrate and itsbly protectively) to block the burrow. successful burrowing beneath the sur-The large cheliped of the male is a face (MacGinitie 1934). The lifespanweapon used in disputes over territory of the ghost shrimp has been variouslyand during the mating season estimated at 3-5 years (Bird 1982), 10(MacGinitie 1934). years (MacGinitie 1935), and 15-16

years (Ricketts and Calvin 1968). TheDevelopment of Eggs and Larvae blue mud shrimp is also believed to be

relatively long-lived (MacGinitieFemale ghost shrimp are ovigerous 1930).

(capable of producing fertile eggs)throughout the year, but the principal Habitatspawning season is in late June andearly July (MacGinitie 1935). Inten- Both of these species are commonlysive breeding probably begins in found in intertidal areas of mixedspring, but ovigerous females may sand and mud. The blue mud shrimpstill be plentiful as late as August. lives in muddier areas than does theSpring warming appears to be the trig- ghost shrimp; observations with re-ger for egg development. Three to spect to tidal height preferences varyfour broods are produced at about (Table 1). In Oregon estuaries, ghost6-week intervals. The larvae develop shrimp were consistently found inas plankton in coastal waters through tideflats closer to the ocean thanfive zoeal stages, which are distin- were blue mud shrimp (Bird 1982).guishable from one another primarily Both species are common residents ofon the basis of size (McCrow 1972). A eelgrass beds in the Pacific Northwesttotal of 6-8 weeks is spent as (Phillips 1984).nearshore oceanic plankton (throughthe five zoeal and one megalopalstage); zoeal larvae are usually re- GROWTH CHARACTERISTICSleased on ebb tides in June and July,and the first megalopae appear in ear- Typically, length of adults isly August. Recruitment to the estuary 5-10 cm (2-4 inches) in the ghostis probably facilitated by flood tides shrimp and 7.5-10 cm (3-4 inches) inoccurring in .3te summer and fall. the blue mud shrimp (MacGinitie andLarval drifting during this planktonic MacGinitie 1968). However, length mayphase very likely serves as a mecha- reach 11.5 cm (4.5 inches) in thenism of genetic exchange among popula- ghost shrimp and 15 cm (6 inches) intions in different estuaries (Johnson the blue mud shrimp (Morris et al.and Gonor 1982). 1980). MacGinitie ( 1930, 1935 )

6

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 Muddier sand Kozloff 1973

Yaquina Bay, OR 0 to 1 ft 0 to 1 ft Thompson andPritchard 1969

Oregon estuaries Tideflats close Tideflats further Bird 1982to ocean from ocean

N. California Sandier mid-tidal Lower, muddier Hedgpeth 1970areas flats

Tomales Bay & Smith andElkhorn Slough, CA Muddy sand Softer mud Carlton 1975

Monterey Bay, CA Generally lower Generally higher MacGinitietidal areas; tidal areas; mud, 1935mixed sand and sandy mud withmud clay

reported finding the largest blue mud (McCrow 1972); 420-770/M2 (1.7-3.1shrimp in the muddiest, least rocky million/acre) in high-density areas ofareas. Sand Lake Estuary, Oregon; and less

than 300/m 2 (1.2 million/acre) inGhost shrimp mature at 18-24 months other areas on the Oregon coast (Bird

and some reproductive females may be 1982). Blue mud shrimp densities inless than 3 cm (1.2 inches) long; blue Oregon estuaries range from 330 tomud shrimp take 3 or more years to 660/M 2 (1.3-2.7 million/acre) (Birdmature and reproductive females 1982). Biomass of either speciesexceed 6 cm (2.4 inches) in length. sometimes exceeds 2.0 kg/m 2 (18,000Estimated growth in length averages lb/acre (wet weight)).approximately 15-30 mm/yr (0.6-1.2inches/yr) in ghost shrimp and 18-26mm/yr (0.7-1.0 inches/yr) in blue mud THE FISHERYshrimp (Bird 1982). Density within aghost shrimp colony and the colony's Ghost and blue mud shrimp arelocation appear to influence both harvested by commercial bait fishermengrowth and size at sexual maturity; and recreational fishermen in Califor-ghost shrimp in the less dense nia, Oregon, and Washington. Petersoncolonies closest to the ocean grow (1977) described a method used infaster, arid the females become sexual- southern California in which water isly mature at larger sizes and produce pumped into the substrate under pres-more and larger eggs (Bird 1982). sure, forcing the animals out of their

burrows; in the area he studied, har-Densities of ghost shrimp have been vest noticeably reduced the ghost

estimated at 700-1,400/m2 (2.8-5.6 shrimp population. In the Pacificmillion/acre) in Yaquina Bay, Oregon Northwest, attempts have been made to

7

control the shrimp on commercial manner, there is evidence that it alsoJapanese oyster (Crassostrea gigas) filters detritus and plankton from thegrounds with the insecticide Sevin water moving through its burrow as(carbaryl). This pesticide has been does the blue mud shrimp (Morris etused to control ghost and mud shrimp al. 1980). Rejected material isin Washington since 1963 (Washington deposited outside the burrow.Department of Fisheries and Washington Burrowing activity is heaviest in theDepartment of Ecology 1985), and al- upper 45-50 cm (18-20 inches), wherethough it has been used on oyster the availability of food is greatestgrounds in Oregon, such use is cur- (MacGinitie 1934). The burrowing andrently unlawful there (L. Fredd, feeding hehavior of the ghost shrimpOregon Department of Fish and Wild- is vigorous enough to cause substan-life, Portland, OR; pers. comm.). tial alterations in surface sedimentDuring its use in Oregon, bait fisher- characteristics over time, decreasingmen noted ghost shrimp mortalities in organic content and shifting the par-untreated areas soon after nearby ticle size distribution upwards (Birdoyster grounds were sprayed (Buchanan 1982). Sediment in dense ghost shrimpet al. 1985). beds often has a soft, quicksand qual-

ity (Posey 1985). The burrowingWashington oyster growers estimate activity of both the ghost and blue

that oyster production would drop 70%- mud shrimp aerates the subsurface soil80%, resulting in a $5 million annual (MacGinitie 1930, 1934).loss in Pacific and Grays HarborCounties, without ghost shrimp control The blue mud shrimp is a suspension(Washington Depa'tment of Fisheries feeder, straining detrital particlesand Washington Department of Ecology and plankton from seawater kept moving1985). However, questions have been through its burrow by the action ofraised about the effects of Sevin on its swimmerets. To feed, the animalother organisms, including the positions itself near a burrowcommercially important Dungeness crab entrance and increases the movement of(Cancer magister), and on the the swimmerets to increase the currentestuarine ecosystem as a whole of seawater through the hurrow. The(Lindsay 1961; Stewart et al. 1967; third maxillipeds are used to periodi-Buchanan et al. 1985). Although the cally sweep the food particles col-blue mud shrimp is believed to disturb lected into the animal's mouth.the sediment far less extensively than Particles that are too big are ejectedthe ghnst shrimp (Bird 1982), both (MacGinitie 1930).have been the objects of controlprograms. Cooperation, Competition, and

Predation

ECOLOGICAL ROLE By aerating the subsurface sedimentand digging burrows protected from

Food and Feeding Habits most predators, ghost shrimp and bluemud shrimp provide an environment

The ghost shrimp was once thought to attractive to commensals. Commensalfeed exclusively by sifting organic and parasitic species associated withdetritus from the floor of its burrow these shrimp include a blind goby,through the hairs on the second and three species of pea crabs, twothird pairs of legs, rejecting coarse species of clams, a copepod, a shrimp,material, and then ingesting the re- polynoid worms, and isopods (seetained fine particles by the use of Table 2).the maxillipeds (MacGinitie 1934).And although it is still thought Species that might compete withto obtain most of its food in this these shrimp for either food or space

8

Table 2. Commensal (c) and parasitic (p) species reported in burrows of ghostshrimp and blue mud shrimp (compiled from MacGinitie and MacGinitie 1968;Ricketts and Calvin 1968; Kozloff 1973).

Found with Found withSpecies ghost shrimp blue mud shrimp

GobyClevelandia ios (c) In burrow In burrow

Pea crabsScleroplax granulata (c) Abundant in burrow Abundant in burrowPinnixa franciscana (c) Abundant in burrow ---

P. schmitti (c) In burrow (rare) ---

ClamsPseudopythina rugifera (c) --- Underside of abdomenCryptomya californica (c) Extends syphons Extends syphons

into burrow into burrow

CopepodsHemicyclops callianassae (c) On gills On gillsClausidium vancouverense (c) Underside carapace Under carapace

(common)

ShrimpBetaeus ensenadensis (c) On gills

Polynoid wormsHesperonoe spp. (c) In burrow In burrow

Isopods(unidentified--p) Under carapace ---

Phyllodurus abdominalis (p) --- Underside of abdomen

are rare in ghost shrimp colonies observed to move from one burrow tobecause of the continual reworking of another and were often found withthe sediment by this species. Infauna part of an appendage exposed above theare both more varied and more abundant surface (Posey 1985).in blue mud shrimp colonies becausethis species less severely affects the The seaward boundary of dense shrimpsediment structure (Bird 1982). beds coincided with a fourfold

seaward increase in the density of theAlthough ghost shrimp typically major predator, the Pacific staghorn

inhabit det burrows, they are suscep- sculpin (Leptocottus armatus) in Coostible to predation by other animals Bay, Oregon (Posey 1986b6. Cagingbecause they sometimes venture outside experiments in Coos Bay indicated thattheir burrow entrances. Under test predation by this fish, which was mostconditions, ghost shrimp spent over intense in summer, probably restricts25% of the time within 2 cm of the the seaward distribution of ghostburrow entrance; the shrimp were also shrimp (Posey 1986b).

9

Mud and ghost shrimps are sometimes below 75% that of seawater (Morriskilled by the leopard shark, Triakis et al. 1980).semifasciata, and by the brown smooth-hound shark, Mustelus henlei. The Oxygen availability is no doubt aleopard shark, whose range extends limiting factor for all intertidalnorth to Oregon, apparently can shovel species, including the ghost and blueor burrow into the substrate to prey mud shrimps (MacGinitie 1935). Thomp-on benthic species (Russo 1975). son and Pritchard (1969) measured oxy-Dungeness crabs are known to eat ghost gen levels in burrows ir Yaquina Bay,shrimp, but the shrimp does not appear Oregon, and found that during ebbto be a major component of the crab's tide, oxygen levels were occasionallydiet (Stevens et al. 1982; Posey zero. They also found that under lab-1985). Sea-run cutthroat trout (Salmo oratory conditions, the ghost shrimpclarki clarki) also commonly eat ghost could survive anoxia (lack of oxygen)shrimp, but are not considered a major for 5.7 days and the blue mud shrimppredator (Posey 1985). Posey also could survive for 3.3 days, far longersuggests that intertidally foraging than they would normally be subjectedbirds may occasionally eat ghost to anoxia in the environment.shrimp.

Although the ghost shrimp has alower normal metabolic rate and sur-

ENVIRONMENTAL REQUIREMENTS vives anoxia and hypoxia (low oxygen)better than does the blue mud shrimp,both appear to have respiratory

The optimal temperature range for adaptations that allow them toghost shrimp appears to be 10 to tolerate the low oxygen conditions13 C, depending on depth below the under which they live. Laboratorysurface. Egg-bearing females seem to experiments have shown that bothprefer the cooler water at the greater species are able to lower theirdepths; immature specimens are found metabolic rates once oxygen levelshigher up in the burrow. Water tem- become critically low. Additionally,perature in ghost shrimp habitat in studies of the ghost shrimp haveYaquina Bay, Oregon, varies season- demonstrated the following adaptationsally from 9 to 15 C (McCrow 1972). to hypoxia/anoxia: when oxygen levelsActivity of ghost shrimp decreased become low, heart rate is loweredslightly with increasing maximum daily (Thompson and Pritchard 1969); aair temperature in an outdoor aquarium respiratory pigment, hemocyanin,(Posey 1987). liberates more bound oxygen to the

tissues (Morris et al. 1980); and theGhost shrimp tend to be most shrimp is able to switch to an

abundant at the seaward end of bays alternate, anaerobic metabolismwith substantial freshwater inflow (Pritchard and Eddy 1979; Morris et(McCrow 1972) and tolerate salinities al. 1980).from about 25% to 125% the salinityof normal seawater (33 ppt). Blood Clifton et al. (1984) studied thesalinity changes along with water effect of spilled oil on ghost shrimpsalinity. In a laboratory test, colonies in Willapa Bay, Washington.salinities of 8-9 ppt were lethal to They concluded that small amounts of75%-100% of ghost shrimp (Posey 1987). oil carried in on the tides and tempo-Activity of ghost shrimp decreased rarily stranded in intertidal areaswith decreasing salinity between 33 are unlikely to have a serious long-and 10 ppt (Posey 1987). The blue term impact. However, stranded oilmud shrimp tolerates salinities as that is buried by a subsequentlow as 10% that of seawater and regu- deposition of oil-free sediment cre-lates osmotically when salinity falls ates a barrier to burrowing activity

10

that can be expected to persist for nature, the ghost and blue mud shrimpyears. They also concluded that the appear to be an integral part of theburrowing activity of the shrimp con- nearshore environments. And fortu-tributes to the introduction of oil nately for the shrimp, their wide-into the sub-surface. spread distribution should allow them

to sustain their populations despiteAlthough their effects on the the current attempts to eliminate them

environment are controversial in locally.

I0

0 11•

LITERATURE CITED

Bird, E.M. 1982. Population dynamics Estuarine Coastal Shelf Sci. 14:501-of thalassinidean shrimps and 516.community effects through sedimentmodification. Ph.D. Dissertation. Johnson, M.E., and H.J. Snook. 1955.University of Maryland, College Seashore animals of the PacificPerk. 151 pp. coast. Dover Publications Inc., New

York. 659 pp.Brenchley, G.A. 1981. Disturbance

and community structure: an Kozloff, E.N. 1973. Seashore life ofexperimental study of bioturbation Puget Sound, the Strait of Georgia,in marine soft-bottom environments, and San Juan Archipelago.J. Mar. Res. 39:767-790. University of Washington Press,

Seattle. 282 pp. and 28 plates.

Buchanan, D.V., D.L. Bottom, and D.A. Lindsay, C.E. 1961. Pesticide testsArmstrong. 1985. The controversial in the marine environment in theuse of the insecticide Sevin in state of Washington. Proc. Natl.Pacific Northwest estuaries: its Shellfish. Assoc. 52:87-97.effects on Dungeness crab, Pacific Shlfs.Aoc528-7oyster, and other species. Pages MacGinitie, G.E. 1930. The natural401-407 in Proceedings of the history of the mud shrimp Upogebiasymposium on Dungeness crab biology pugettensis (Dana). Ann. Mag. Nat.and management, Oct. 9-11, 1984, Hist., Ser. 10, 6:36-44.

Anchorage, AK. Alaska Sea Grant

Rep. No. 85-3, University of Alaska, MacGinitie, G.E. 1934. The naturalAlaska Sea Grant College Program, history of Callianassa califor-Fairbanks. niensis Dana. Am. Midl. Nat. 15:166-

Clifton, H.E., K.A. Kvenvolden, and 177.

J.B. Rapp. 1984. Spilled oil and MacGinitie, G.E. 1935. Ecologicalinfaunal activity--modification ofburrowing behavior and redistri- aspects of a California marinebution of oil. Mar. Environ. Res. estuary. Am. Midl. Nat. 16:629-765.11:111- 136. MacGinitie, G.E., and N. MacGinitie.

1968. Pages 284-293 in NaturalHedgpeth, J.W. 1970. Introduction to history of marine animals, 2nd ed.

seashore life of the San Francisco McGraw-Hill, New York.Bay region and the coast of northernCalifornia. University of Cali- McCrow, L.T. 1972. The ghost shrimp,fornia Press, Berkeley. 136 pp. Callianassa californiensis Dana,

1854, in Yaquina Bay, Oregon. M.S.Johnson, G.E., and J.J. Gonor. 1982. Thesis. Oregon State University,The tidal exchange of Callianassa Corvallis. 56 pp.californiensis (Crustacea, Decapoda)larvae between the ocean and Morris, R.H., D.P. Abbott, andthe Salmon River Estuary, Oregon. E.C. Haderlie. 1980. Intertidal

13

invertebrates of California. Stan- important invertebrate animals.ford University Press, Stanford, U.S. Fish Wildl. Serv. Biol. Serv.California. 690 pp. Program FWS/OBS-83/16. 225 pp.

Peterson, C.H. 1977. Competitive Russo, R.A. 1975. Observations onorganization of the soft-bottom the food habits of leopard sharksmacrobenthic communities of southern (Triakis semifasciata) and brownCalifornia lagoons. Mar. Biol. smooth-hounds (Mustelus henlei).(Berl.) 43:343-359. Calif. Fish Game 61:95-103.

Phillips, R.C. 1984. The ecology of Schmitt, W.L. 1921. The marineeelgrass meadows in the Pacific decapod crustacea of California.Northwest: a community profile. Univ. Calif. Publ. Zool. 23:1-470.U.S. Fish Wildl. Serv. FWS/OBS-84/24. 85 pp. Smith, R., and J.T. Carlton. 1975.

Light's manual: intertidal inverte-Posey, M.H. 1985. The effects upon brates of the central California

the macrofaunal community of a coast. University of Californiadominant burrowing deposit feeder, Press, Berkeley. 716 pp.Callianassa californiensis, and therole of predation in determining its Stevens, B.A. 1928. Callianassidaeintertidal distribution. Ph.D. from the west coast of NorthDissertation. University of Oregon, America. Publ. Puget Sound Biol.Eugene. 119 pp. (Berl.) Stn. 6:315-369.

Pocey, M.H. 1986a. Changes in a Stevens, B.G., D.A. Armstrong, and R.benthic community associated with Cusimano. 1982. Feeding habits ofdense beds of a burrowing deposit the Dungeness crab Cancer magisterfeeder, Callianassa californiensis. as determined by the Index ofMar. Ecol. Prog. Ser. 31:15-22. Relative Importance. Mar. Biol.

72:132-145.Posey, M.H. 1986b. Predation on a

burrowing shrimp: distribution and Stewart, N.E., R.E. Millemann, andcommunity consequences. J. Exp. W.P. Breese. 1967. Acute toxicityMar. Biol. Ecol. 103:143-161. of the insecticide Sevin and its

hydrolytic product 1-naphthol toPosey, M.H. 1987. Effects of lowered some marine organisms. Trans. Am.

salinity on activity of the ghost Fish Soc. 96:25-30.shrimp Callianassa californiensis.Northwest Sci. (in press). Thompson, R.K., and A.W. Pritchard.

1969. Respiratory adaptatior, ofPritchard, A.W. , and S. Eddy. 1979. two burrowing crustaceans,

Lactate formation in Callianassa Callianassa californiensis andcaliforniensis and Upogebia puget- Upogebia pugettensis (Decapoda,tensis (Crustacea: Thalassinidea). Thalassinidea). Biol. Bull.Mar. Biol. (Berl.) 50:249-253. (Woods Hole) 136:274-287.

Ricketts, E.F., and J. Calvin. 1968. Washington Department of Fisheries andBetween Pacific tides. Revised by Washington Department of Ecology.J.W. Hedgpeth, Stanford University 1985. Final environmental impactPress, Stanford, CA. 614 pp. statement: use of the insecticide

Sevin to control ghost and mudRudy, P., Jr., and L.H. Rudy. 1983. shrimp on oyster beds in Willapa BayOregon estuarine invertebrates: an and Grays Harbor. 82 pp. plusillustrated guide to the common and appendices.

14

30272 -1t

REPORT DOCUMENTATION I. REPORT NO. 2. 3. ReciPent's Accession No.PAGE Biological Report 82(11.93)*

4. Title and Subtitle S. Report DatsSpecies Profiles: Life Histories and Environmental Requirements of January 1989Coastal Fishes and Invertebrates (Pacific Northwest)--Ghost Shrimp 6and Blue Mud Shrimp

7. Auth.ons) 6. Performing Organization Rept. No.Susanna Hornig, Anita Sterling, and Stanley D. Smith

4. Performing Organization Name and Address 10. Project/Task/Work Unit No.U.S. Fish and Wildlife ServiceNational Fishery Research Center 11. conct(c) or Grnt(G)No.Building 204, Naval Station Puget Sound (C)Seattle, WA 98115-5007

12. Sponsoring Organization Name and Address

U.S. Department of the Interior U.S. Army Corps of Engineers TypeofRport4PeriodCover"

Fish and Wildlife Service Waterways Experiment StationNational Wetlands Research Center P.O. Box 631Washington, DC 20240 Vicksburg, MS 39180

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

15. Abstrsct (Limit: 200 words)

Species profiles are literature summaries of the taxonomy, morphology, range, lifehistory, and environmental requirements of coastal aquatic species. The profiles areprepared to assist in environmental impact assessments. The ghost shrimp(Callianassa californiensis) and blue mud shrimp (Upogebia pugettensis) are commonresidents of intertidal mudflats of the Pacific Northwest, as well as of the entirewest coast of the contiguous United States. These species are decapod crustaceans,

but not true shrimp. They are harvested as bait by recreational and commercialoyster-growing operations. Ghost shrimp larvae develop in summer in nearshore coastalwaters and settle to the substrate surface, where they rapidly metamorphose; the lifecycle of the blue mud shrimp is presumed to be similar. Both species spend theirlives in burrows in the mudflat, where the ghost shrimp is primarily a deposit feederand the blue mud shrimp is a suspension feeder.

17. Domment Analysis a. Oescriptors

Fishery Salinity Sediments PredatorsGrowth Temperature Life Cycles CompetitorsEstuaries Oxygen Shrimps CommensalsFeeding habits Depth Oil spills

b. Identlfleirs/Open-Ended Terms

Ghost Shrimp Callianassa californiensis Dana

Blue Mud Shrimp Upogebia pugettensis (Dana)

c. COSATI ileld/Group

1. Availability Statement IS. Security Class (This Report) 21. NO. of Pagse

Unclassified 14Unlimited release 20 Security Class rhis Page) 2. Price

Unclassified(See ANgI-29.1I) OPTIONAL FORM 272 (4-77)

(Formerly NTIS.-35)Ospartint of Commerme