-A181 591 SPECIES PROFILES LIFE HISTORIES AMD ENVIRONMENTAL 1/1REQUIREMENTS OF COASTAL.. (U) WASHINGTON COOPERATIVEFISHERY RESEARCH UNIT SEATTLE D J GROSSE ET AL. AUG 6

UNLIFIED FWS-82/ii.9F/O 6/3 NL

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Species Profiles: Life Histories and DTICEnvironmental Requirements of Coastal Fishes ELECTEand Invertebrates (Pacific Northwest)JU197D

AMPHIPODSE

Coastal Ecology Group

Fish and Wildlife Service Waterways Experiment Station

U.S. Department-of the Interior U.S. Army Corps of Engineersrab&MM kidinin b IdIM PA nl Md I

Biological Report 82(11.69)TR EL-82-4August 1986

Species Profiles: Life Histories and EnvironmentalRequirements of Coastal Fishes and Invertebrates (Pacific Northwest)

AMPHIPODS

by

Daniel J. Grosse and Gilbert B. PauleyWashington Cooperative Fishery Research Unit

University of WashingtonSeattle, WA 98195

andDavid Moran

National Wetlands Research Center

Project OfficerJohn Parsons

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 ServiceU.S. Department of Interior

Washington, DC 20240

e

This series may be referenced as follows:

U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life historiesand environmental requirements of coaistal 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:

Grosse, D.J., G.B. Pauley, and D. Moran. 1986. Species profiles: lifehistories and environmental requirements of coastal fishes and invertebrates(Pacific Northwest)--amphip6ds. U.S. Fish Wildi. Serv. Biol. Rep. 82(11.69).U.S. Army Corps of Engineers, TR EL-82-4. 15 pp.

lip

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 add d 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 s~ould be directed to one ofthe following addresses.

S /

Information Transfer Specialist ' , ...National Wetlands Research CenterU.S. Fish and Wfldlife ServiceNASA-Slidell Computer Complex1010 Gause BoulevardSlidell, LA 70458

or

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

Dis fj--' u

, .. . ..

-- V- .,' ""A '

CONVERSION TABLE

Metric to U.S. Customary

Multtply P1 To Obtain

mtlliaeters (wn 0.03937 inchescentimeters (On) 0.3937 Inchesmeters (m) 3.281 feetkilometers (in) 0.6214 miles

square meters (m2) 10.76 square feetsquare kilometers (k02) O.3861 square mileshectares (ha) 2.471 acres

liters (1) 0. 2642 gallonscubic ;meters (m3) 35.31 cubic feetcubic meters 0.0008110 acre-feet

milligrams (ing) 0.00003527 ouncesgrams (g) 0.03527 ounces I.kilograms (kg) 2.205 poundsmetric tons (t) 2205.0 poundsmetric tons 1.102 short tonskilocalories (kcal) 3.968 British thermal units

Celsius degrees 1.8(0C) + 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 (nmn) 1.852 kilometers

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

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

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

Fahrenheit degrees 0.5556(°F - 32) Celsius degrees

iv.- -

CONTENTS

Page

PREFACE. .. .. ........................... iiiCONVERSION TABLE .. .. .......... ....... ....... ivACKNOWLEDGM4ENTS. .. .. . .... ....... ...... ........ vi

NOMENCLATURE/TAXONONY/RANGE. .. ....... ....... ........ 1MORPHOLOGY/IDENTIFICATION AIDS. .. .... ....... ....... ... 4REASON FOR INCLUSION IN SERIES .. .. ...... ....... ......LIFE HISTORY .. .. ...... ....... ....... .......... 5GROWTH CHARACTERISTICS. .. .... .................................. 5POPULATION DYNAMICS AND IMPORTANCE TO FISHERY .. .. ... ....... ... 5ECOLOGICAL ROLE. .. .. ...... ....... ....... ....... 7ENVIRONMENTAL REQUIREMENTS. .. .... ....... ....... ..... 10

Dissolved Oxygen .. .. ....... ...... ....... ...... 10Salinity. .. .... ....... ....... ...... ....... 10Pollution and Dredging. .. .... ....... ...... ....... 10

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

- .

ACKNOWLEDGM4ENTS

We gratefully acknowledge the reviews by Rick Albright, School of Fisheries,University of Washington, Seattle, and Craig P. Staude, Friday HarborLaboratories, University of Washington, Seattle.

i

vii

Pereon

Head---

Rostrum-2

Gland Cone- 31

Flagellum 3 ,

Gnathopods

Antennae Pereopods Pleopods Uropods

Figure 1. A gamaridean amphipod (from Staude et al. 1977).

~>AMPHIPOD

NOMENCLATURE/TAXONOMY/RANGE northeast Pacific Ocean (Figure 3).Gammaridea are the most abundant and

Scientifc ndiverse of the amphipods. AlthoughScientific nameAmphipoda primarily marine, they are also

(Figure 1) found in freshwater and certainPreferred common name . . ( Amphipods moist terrestrial habitats (ReishClass ...........>rustacea) and Barnard 1979). Marine GammarideaSubclass ......... Malacostraca are ubiquitously distributed. TheyOrder .. ........... .. Amphipoa- are found in all regions, in allSuborders . . Gammaridea, Hyperiidea, habitats, and at most depths. AboutCaprellidea, Ingolfiellidea (Figure 40% of the 80 gammaridean families2). are cosmopolitan in distribution;

the remaining 60% are looselyGeographic range: This report will associated with specific regions or

focus largely on the suborders zones (Barnard 1969; BousefieldGammartdea, Caprellidea, and 1978). Gammaridean distributionsHyperiidea because of their impor- remain poorly known, but more recenttance in coastal areas of the studies (e.g., Barnard 1971) are

,a

D E

Figure 2. A, Elasmopus and B, Eohaustorius, both gammarid amphipods. C,Caprella ferrea, a caprellid amphipod. D, Neocyamus hyseteris (female), acaprellid 7ahipod from sperm whale. E, Phronima sedentaria, a heriid amphipodthat lives inside the tunic of urochordates. (A and B from Barnard 1975; C and 0from McCain 1975; E from Barnes 1974. A-D reprinted with permission from theUniversity of California Press; E reprinted with permission from Saunders CollegePublishing.)

finding more widespread distribu- About 200 gammarid species have beentions than were previously assumed. found in Washington waters (StaudeOff the Oregon Coast, 97 species of et al. 1977). Some gammarid speciesgammarids have been found from the dwell in subtidal or intertidalsurface to a depth of 2900 m environments (Reish and Barnard(Barnard 1971), and 20 species 1979). The suborder Hyperiidea isdivided among 11 families were in entirely marine and pelagic; most -

the upper 200 m (Pearcy 1972). members of the taxon live in the

2

VANCOUVER AI I. -~CANADA

Y

WASHINGTON

PORTLAND~ * 4 4

MILESso too

- -~---~Coastal distribution0 so too ...

KILOMETERSmo

#A Concentrated estuarineareas

OREGON

CALIFORNIA -

Figure 3. Distribution of the ubiquitous amphipod suborders bu.maridea andHyperiidea in the coastal areas of the northeast Pacific Ocean.

3

bathyal zone, and some live in fused with second thoracic segment.coastal waters (Bowman and Gruner Body slender and resembling that of a1973). praying mantis (except for whale

lice). Marine. Includes skeletonshrimp Suborder Caprellidea.

MORPHOLOGY/IDENTIFICATION AIDS2a. Eyes generally large, occu-

Animals of the order Amphipoda pying most of head; coxae of pereopodsare distinguished by sessile, compound small, often fused with the body,eyes, though some species are blind. maxillipeds without palp; last twoA carapace is not present and the abdominal segments fused; body more orfirst, and sometimes the second, less transparent. Marine, and usuallythoracic segments are fused with the planktonic or associated with jelly-head. A "shrimplike" appearance fish or in tunics of dead salps .results from lateral body compression. .Suborder Hyperiidea.Gammarids and hyperiids have threepairs of pleopods (swimmerets); two or 2b. Eyes usually present andthree pairs of uropods on the pleon conspicuous, but not large enough to(abdomen); at least eight pairs of cover most of the head; coxae ofthoracopods, counting the maxilliped; pereopods well developed, usuallyusually seven major leg pairs, called expanded. Marine, freshwater, andpereopods; and five or more pairs of terrestrial . . . Suborder Gammaridea.gills. Males and females often can bedistinguished morphologically. The 2c. Body elongate; coxae small;head has five fused segments, two abdominal segments distinct; all butpairs each of antennae and maxillae, a fourth and fifth pairs of abdominalheavily chitinized mandible, and a appendages vestigial. Marine, inter-limblike maxilliped. There are seven stitial. Rare. . . . Suborderfreely articulated somites on the Ingolfiellidea.thorax (pereon). Coxal platelikelateral extensions of the thoracic There currently exists no concisepereon are developed from the first guide to amphipod species in thesegment of each leg. Branchiae northwest Pacific. Publications of(gills) are fleshy and platelike and the National Museum of Canada, such asare attached medial to the coxae, 2-6 that by Conlan and Bousfield (1982),on each side. The abdominal region will eventually culminate in a compre-consists of three articulating hensive regional handbook on marinesegments on both anterior pleon and gammarideans. Contributions byposterior urosome; the urosome has a Barnard (1975) and Staude et al.terminal telson. (1977) may be useful for identifying

Gammaridea in restricted intertidalThe following key (adapted from regions; the work of Bousfield (1978)

Barnes 1974; Kozloff 1974) is pre- described freshwater Gammaridea.sented as an aid to separate the sub- Kozloff (1974) provided keys to theorders of amphipods: Caprellidea. Hyperiids can be identi-

fied to genus by using the descrip-la. Pereon with seven apparent tions published by Bowman and Gruner

segments, all having well-developed (1973).appendages. Abdomen not vestigial.Body neither slender nor resemblingthat of a praying mantis. ...... 2 REASON FOR INCLUSION IN SERIES

lb. Pereon with six apparent Hyperiid amphipods are the thirdsegments, some may have vestigial most abundant group of coastal marineappendages; abdomen vestigial; head crustacean zooplankton, following

4

Copepoda and Euphausidea (Bowman and coldest winter months. Females layGruner 1973). The benthic amphipods, eggs during each of the last five orespecially Gammaridea, are an invalu- six molting stages, or at every otherable food source for many economically stage (Barnard 1969).important fish and invertebratespecies. Their limited mobilitysuggests that their distribution and GROWTH CHARACTERISTICSabundance can be used as an indicatorof environmental quality (Albright Growth is initially rapid in1982). Omnivorous, opportunistic Gammaridea; molting initiates withinfeeders such as lysianassids (a several days of hatching and continuesgammaridean family) recycle detritus after maturity, slowing to every 20 toand may help avert pollution by scav- 30 days in the later stages ofenging carcasses of larger animals development. The average instarfollowing mass mortalities (Reish and (stage of development betweenBarnard 1979). successive molts) lasts 15 days.

Gammarids go through at least 12instars; thus, the maximum lifespan

LIFE HISTORY estimates are a little more than 6months, although some polar species

Female Amphipoda spawn via an are known to live 5 or 6 years (Reishamplexus (mating embrace) with males and Barnard 1979).which lasts for hours or days. Inswimming species the female swims with Amphipod growth rates-and lengthsthe male on top, or both swim on their vary considerably. Adult amphipodssides. Following ecdysis (molting) range in size from less than 1 cm toand mating, eggs are laid through two about 28 cm, the largest being anventral pores in the female's sixth undescribed lysianassid that wasthoracic sternite. Eggs can number photographed in the abyssal Pacificfrom 1 to 200 or more. Thin, Ocean (Schmidt 1968). Maximum growthtube-dwelling gammarids have the rates of A. pugettensis, mentionedfewest eggs, which tend to be large or above, were 4.1% per week at 10 °C,contain large amounts of yolk. increasing more than threefold toBecause of the large size of the eggs, 14.3% per week at 20 OC (Figure 4),only one can be carried by some young with higher efficiency at 20 *C.females, while fully mature females Growth relative to food intake incarry three or four. Eggs hatch large (10 mg) individuals of thisdirectly into juveniles resembling species was 47% to 72% when fedadults. In gammarids, one-quarter to Enteromorpha (Chang and Parsons 1975).one-half of the eggs may die beforehatching. Juveniles are generallyheld in the brood pouch for a few POPULATION DYNAMICS AND IMPORTANCE TOhours to a few days after hatching FISHERIESbefore they are released (Barnard1969; Reish and Barnard 1979). Amphipods are the main food item

of many fish species, as well as otherChang and Parsons (1975) found aquatic animals (Figure 5). Some

that the common inshore gammarid pelagic species sometimes comprise theAnisogammarus pugettensis breeds bulk of the diet of herring, mackerel,year round in the Pacific Northwest, and Biscayan tunny (Schmitt 1968).in contrast to beach and some inter- Gammarids, on the basis of the Indextidal amphipods of the cooler North of Relative Importance (IRI), were theAtlantic. Those species either have most important food species forone brood per year or cease their nearshore fishes in the Strait of Juanreproductive activity during the de Fuca (comprising more than half of

5, .

10.0- chum salmon (Oncorhynchus keta) andcoho salmon (Oncorhynchus kisutch) maybe attributable in part to the abun-dance of food organisms in rivers and

20 C estuaries. Abundant populations of1.0 gamuarids in the upper estuary of

F. 10 Hyman Creek, British Columbia, con--stituted the main diet of the fry of

these two salmon species. They alsoconstituted the majority of the diet

~.10- of chum fry at six nearby estuaries atlow tide in the spring.

Corophium salmonis, a tube-dwelling gammarid, is an abundant and

.01 preferred prey organism of chum salmon2 4 10 12 14 in the Skagit River salt marsh in

Washington State (Congleton and SmithWeeks 1976). Though little is known of the

Figure 4. Growthot Anisogarus seasonal abundances of C. salmonis,pugettensis fed Enteromorpha intes- Albright (1982) found peak densitiestinalis at 10 and 20 1C. (Reprinted of the species in tide flats of GraysWipermission from the Journal of Harbor, Washington, in July and Au-the Fisheries Research Board of gust. In the inner half of the bayCanada, from Chang and Parsons 1975. they were the dominant organism on mud

and sandy mud bottoms. Densities ashigh as 57,O00/m2 have been observed(Albright and Rammer 1976). From

the total IRI spectrum for 38X of the April through September production was55 fish species studied) and were the 3.6-10.7 g/m2 , and turnovermost important food item to tidepool (production/mean biomass) was 7.2 tofishes (Cross et al. 1978). For the 8.6 g/m2. In Grays Harbor, C.most part, the gammarids were epiben- salmonis is an important prey item forthic rather than infaunal or pelagic. dunlin (Calidris !), English soleCross et al. (1978) suggested that (Parophyrys vetulus) and starrysince hyperiid populations on which flounder (P hhys stellatus)neritic fishes feed are naturally according to Smith and Mudd (1976) andpatchy, small localized perturbations for other fish species, as well asare likely to create more patchiness, shrimp (Crangon spp.) and DungenessIf adjacent areas remain unaffected, crab (Cancer magister) (Albrightthe neritic fish populations may not 1982). Smith (1980) reportedbe adversely affected. However, similarly high C. salmonis densitiessublittoral fishes, especially juve- and predation on this amphipod bynile fishes, are more dependent on various species in other northwestepibenthic prey, and thus more likely estuaries.to be affected by perturbation,although the amphipod supply is Numerically, amphipods are theoften replenished by tidal action. major component of the fauna ofBecause of their relative isolation, harbor pilings in California. Mosttidepool fishes are most heavily are introduced species that have hadaffected by perturbation (Cross et al. little effect on indigenous amphipods1978). in nearby areas (Barnard 1961; Reish

1964). Negligible economic loss dueMason (1974) hypothesized that to fouling has resulted (Reish and

delayed seaward migrations of juvenile Barnard 1979). In heavily polluted

6

Eogamma rum

WesernSadpierPacific Staghorn Juvenile Salmon Dni

Sculpin

Long-billed

Dowitcher Shiner Perch

Starry - English SoleFlounder

Corphum Three-spine

Snake Prickleback I I Stickleback

Juvenile Pacific SaddlebackTomcod Gunnel

Dungeness Crab Shrimp

Longfin Smelt Nemertean Worms

Figure 5. Fish and invertebrate predators of the amphipod Corophium salmonis

(from Albright 1982).

sections of harbors, amphipods are It also scavenges dead fishes andreportedly absent both in the benthos uneaten fish food in ponds. However,and on pilings (Reish 1959). its growth is slower than that of

brine shrimp. Gammarus lacustris,Of the pelagic organisms in the found in shallow prairie lakes of the

upper 200 m off the Oregon coast, Hudson Bay drainage, meets dietaryhyperids comprise more than 10% of requirements for rainbow trout

organisms by number; their abundance (Salmo gairdneri) 5 cm or greater, isis known to vary seasonally (Van easiTy captured, and can be harvestedArsdale 1967, cited by Pearcy 1972). at a rate of 1,000 kg per ha per year.

For most food ingredients it isTwo gammarid species have been comparable to or better than commer-

examined for their potential as food cial feeds, and it improves bodyin fish culture. Mass culture of coloration and hence marketability ofAntsogammarus pugettensis was proposed trout (Mathias et al. 1982).by Chang and Parsons (1975) as analternative to brine shrimp culturefor young salmon; A. pugettensis can ECOLOGICAL ROLEtolerate wide ranges of temperaturesand salinities and thrives on a Amphipods are considered the mostvariety of plant and animal material, efficient scavengers of sea bottoms

7

AWC,

and shores, probably clearing up and seven gammarid families (e.g.,recycling more organic shore debris Ampelisca sp. and Photis sp.) con-than any other animal (Schmitt 1968). struct tubes or cradles on soft orGriffiths and Stenton-Dozey (1981) hard substrata, according to Barnarddescribed the importance of the (1969). Corophium sp., common ingammarid Talorchestia capensis in estuaries where silting is heavy,consuming beached kelp in South forms masses of heavy tubes andAfrica. This species and dipteran creates currents with its abdominallarvae consume some 60% to 80% of appendages. The currents are strainedbeached kelp within 2 weeks, and the by fringes of fine hairs on thegammarid is thought to make a signifi- appendages forward of the abdomen;cant contribution (through feces) to then whatever is collected is scrapedorganic enrichment of the inshore into the mouth (Kozloff 1973). Othermarine system. species inhabit dwellings of other

organisms. Many species are burrow-Caine (1980), in an ecological ers, especially in the gammarid

comparison of two littoral species of families Haustoriidae, Oedicerotidae,caprellid amphipods in Washington and Phoxcephalidae. ElongatedState, indicated that each species has setae on the distal articles of thea different effect on its community. posterior pereopods are an adaptationDeutella californica is a predator, for burrowing (Reish and Barnardbut its removal did not alter com- 1979).munity structure, even though itdisplays a preference for the A number of Gammaridea live onepibiotic community of Obelia sedentary invertebrates such asdichotoma. In contrast, Caprella corals, sponges, tunicates, anemones,Taiu sc uIla is a periphyton scraper and polychaetes. Their relationshipsthat has an enormous impact on the with the hosts are not well understoodperiphyton on Zostera marina, and thus (Reish and Barnard 1979).increases avaiiiale light for theseagrass, and permits its growth in Hyperiidea are primarily nek-areas where it would have otherwise tonic. They have well-developedbeen excluded. Observations on swimming devices or buoyancy control,interspecific aggressive behavior or are found in association withindicate that C. laeviuscula is medusae or salps (Reish and Barnarddominant over other caprellids in 1979). Phronima sp., sometimesprotected habitats. Predatory collected in plankton tows or alongcaprellids did not appear to occur docks in the San Juan Islands, istogether where they would compete found in empty salp tests (Kozloffdirectly for food, while filter- 1974). Hyperiids may feed on the veryfeeding caprellids do compete to some organisms that host them, but may alsoextent for food (Caine 1977). use them as a base from which to

forage, or they may feed on foodReish and Barnard (1979) catego- captured by the host (Bowman et al.

rized gammarids and hyperiids by 1963). Their feeding habits arehabitat. Nestlers include beach- poorly understood. In one laboratoryhoppers of the gammarid family study of Lestrigonus sp. andTalitridae, commonly found on sandy Bougisia sp., food was shared withintertidal areas. High numbers occur the host, Leptomedusa sp., when foodunder moisture-maintaining algal supply was adequate, but when it waswrack, as discussed above. These not, the amphipods fed on host tissuespecies must be transitory, the (Bowman and Gruner 1972). Para-authors speculated, because of fre- themisto sp., a free-living hyperiTTd,quent changes in tide and wrack preys on other plankters (Bowmanaccumulations. Species of six or 1960).

8

A few nektonic gammarids are Locomotion is accomplished with afound in neritic waters. These are loop-like movement in which theeither predators or are mating or front legs attach while the reardispersal phases of benthic gammarids ones release and reattach (Kozloff(Reish and Barnard 1979). 1973).

Chelura terebrans, a wood-borerfound in In addition to being prey formphipod pest. It enlarges holes in many fish and invertebrate species,wood made by the isopod Limnoria i some pelagic amphipods comprise part

of the crustacean diet of whales.(Reish and Barnard 1979). Most of the grey whale diet on the

The swimming capability of epi- west coast consists of six species of

benthic gammarids may reduce their benthic amphipods (Matthews 1978).

susceptibility to predation. Feller British gulls are also known to

and Kaczynski (1975) suggest that consume benthic amphipods (Schmitt

harpacticoid copepods were preferred 1968).

over amphipods by juvenile chum salmonin Puget Sound during the spring Locomotion in gammarids is large-because they are relatively easy to ly by swimming; they are poorlycapture. Simenstad (1976) noted the balanced for walking. Even burrowerssame predatory habits for juvenile are strong swimmers. Small couplingpink and chum salmon in Hood Canal, hooks join pleopods, facilitatingWashington, and found that juvenile coordinated paddling motions. Somesalmon, in addition to preferring the softbottom gammarids have elongatedless numerous and smaller harpactacoid pereopods spread out like a spider'scopepods, also consumed gammarid eggs. legs to prevent sinking into the mud.

The body hangs upside down, loweringAnisogammarus conferviculus is the center of gravity. Sediment

believed to defend or "buffer" its burrowers possess strong and denselypopulations against predation by mi- packed spines on their pereopodsgrating fishes, such as juvenile chum (Barnard 1969).salmon, by ecological adaptations.These adaptations decrease the Caprellids, the suborder whichforaging efficiency of the predator includes skeleton shrimp, are largelyand include association with refuges intertidal and shallow subtidal.in vegetation, clumping in refuges, Their preference of substrate in theassociation with structurally complex Pacific Northwest is not specific, buthabitats and distributions related to they do need to cling to something.rlverflow and tides (Levings and Levy Thus, they are found on algae, sea-1976). In Grays Harbor, Washington, grasses, sponges, hydroids, andhowever, mature male C. salmonis are bryozoans, but not on bare sand or mudsubject to heavy predation beginning bottoms. Caprellids feed on diatoms,in April, when they wander over tide- small invertebrates, and possiblyflats in search of females (Albright detritus, and are prey for many fishes1982). (including cod, blennies, and skates)

and also for shrimp (McCain 1975).Hyperiid swimming varies from Whale parasites of the genus Cyamus

feeble movement of appendages in are also in the caprellid suborder.Cystisoma sp. to rapid swimming in This group includes about 18 host-Paraprone sp. which has strong pleonal specific species. They lack a free-musculature (Bowman and Gruner 1973). swimming stage; they leave theCaprellids, attaching with posterior parental brood pouch and dig into thelegs, feed by grasping food with host with hooked dactyls (Schmitttheir free anterior legs and antennae. 1968).

9

ENVIRONMENTAL REQUIREMENTS Salinity

Dissolved Oxygen Adult gammarids found in es-

Pelagic gammarid and hyperiid tuaries are fairly tolerant to a wide

amphipods have been collected from a salinity range while many juveniles

scattering layer in deep, poorly and embryos are not. Adult estuarine

oxygenated waters off southeastern Corophium volutator survived in salin-

Vancouver Island, British Columbia ities of 2 to 59 ppt (McClusky 1967),

(Waldichuck and Bousfield 1962). but preferred a range of 10 to 30 pptAnisogammarus sp. and Allorchestes (McClusky 1970). Adult C. triaenonyx

sp., both common inshore gammarid survived in a similarly wide range of

amphipod genera, were found in low salinities (Shyamasundari 1973),

dissolved oxygen environments (as low though juveniles could develop only at

as 0.04 ppm at 12 °C) near sulfite- salinities of 7.5 to 37.5 ppt. For

rich paper pulp effluent. The former large numbers of individuals to sur-

species was found in high numbers on vive and develop, 20 to 32.5 ppt were

the bottom (15 to 22 m) and the latter required (Shyamasundari 1976).

species, normally found in shallower Anisogammarus pugettensis, found

waters, was observed near the surface, naturally in 20 to 28 ppt salinities,perhaps seeking more oxygenated water cannot survive in freshwater but can

(Waldichuk and Bousfield 1962). Low survive at 11 ppt for at least 1 week

oxygen tolerance in either species (Chang and Parsons 1975). Some

remains to be determined, but Chang species, such as Phoxocephalid spp. or

and Parsons (1975) observed that Ampeliscad spp., may have very narrowAnisogammarus pugettensis survived salinity tolerances. Other amphipods

for several hours at 20% satura- (e.g., Gammarus spp., Hyalella spp.

tion levels. They also determined a and Crangonyx spp.) are found in tQ of 1.6, lower than those of other freshwater.castaceans for which it is around2 (Q is the factor by which themetab 9 ic rate increases after a 100 Pollution and Dredging

increase in temperature). They sug-ges tht tis s a adptaionforReish and Barnard (1979) observedgest that this is an adaptation for that some amphipod species are morecopingtolerant than others to organic pol-tidal temperatures. Caprellids are tknown to leave eelgrass beds "in lution, but do not know what environ-

mental factors cause the differences.droves"It is known that some amphipods are

oxygen levels in the beds drop below sensitive to pollution in harbors.2 ppm. Capitella sp., a marine polychaete

Tolerances to low dissolved which is commonly used as a pollutionoxygen levels vary greatly among indicator and which has a distributionspecies; many are very sensitive to that is often mutually exclusive tolow levels, especially species that of amphipods, is found in heavily

restricted to areas where dissolved polluted harbors. Capitella sp. isoxygen does not historically vary also found in unpolluted areas, suchgreatly. Groups such as phoxo- as deep sea bottoms off the coast of

cephalids (used as indicators of pol- California, which are subject to

lutant levels in sediment bioassays) freshwater inflow -- places whereappear much less tolerant to stress- amphipods are notably absent (Reish

ful conditions than many of the and Barnard 1979).species discussed above (R. Albright,University of Washington; pers. The distribution of Corophiumcomm.). salmonis is influenced by sediment ,

10. .%

type and depth (it prefers shallow, Ampelisca show sensitivity in sub-muddy sand substrates) more than by tidal areas.salinity. Other species ofCorophium exhibit greater produc- Dredging is likely to eliminatetion near sewer outfalls -- an in- benthic amphipods, which live on orcrease which is presumably attribu- close to the substrate (Reish andtable to organic enrichment (Birklund Barnard 1979). However, McCaulley et1977). al. (1977) suggest that in the event

of dredging, adults are likely to moveBehavioral changes in amphipods to nearby unaffected areas or juve-

exposed to sublethal quan'ities of niles may rapidly settle and repopu-oil have been noted and suggest a late the dredged areas (McCaulleysensitivity to fresh oil. Beach- et al. 1977). Crustaceans arehoppers are most likely to be generally very sensitive to pollu-affected by oil due to their occur- tion and, therefore, species depend-rence in the high-tide wrack zone ent on them as food are indirectly(Baker 1971), while species of affected by pollution.

0 1J

I' 6ea

LITERATURE CITED

Albright, R. 1982. Population dynam- Barnes, R.D. 1974. Invertebrateics and production of the amphipod zoology, 3rd ed. W.B. Saunders,Coohu salmonis in Grays Harbor, Philadelphia, Pa. 870 pp.Washington. M. S. Thes is. Univer-sity of Washington, Seattle. 76 pp. Bengston, C., and J. Brown. 1976.

Maintenance dredging and the envi-Albright, R., and A.D. Ramer. 1976. ronment of Grays Harbor, Washington.Maintenance dredging and the Appendix G: Impact of dredging onenvironment of Grays Harbor, Wash- the fishes of Grays Harbor. U.S.ington. Appendix E: The effect Army Corps of Engineers, Seattleof intertidal dredged material District. 125 pp.disposal on benthic invertebrates.U. S. Army Corps of Engineers, Birklund, J. 1977. Biomass, growthSeattle District. 244 pp. and production of the amphipod

Crphiu insidiosum (Crawford) andBaker, J.M. 1971. Growth simu- iiihay notes on Crphiu

lation following oil pollution. volutator (Pallas). Opheliaii1;2)T_-Pages 72-77 in E.B. Cowell, ed. 187203The ecological effects of oilpollution on littoral communities. Bousefield, E.L. 1978. A revisedInstitute of Petroleum, London. classification and phylogeny of

amphipod crustaceans. Trans. R.Barnard, J. L. 1961. Relationship of Soc. Can. Ser. 4(16):343-390.

California amphipod faunas in New-port Bay and in the open sea. Pac. BwaTE 90 h eai

Nat.2: 66-68.aiqphipod genus ParathemistoBarnard, J.L. 1969. The families Hyrida yeidaTiti

and genera of marine gamaridean North Pacific and adjacent Arcticamphpoda U.S Nal. Ms. Bll. Ocean. Proc. U.S. Natl. Mus. 112

271. 535 pp. (49:3332

Barnard, J.L. 1971. Gammaridean Bowman, T.E., and H. Gruner. 1973.amphipoda from a deep-sea transect The families and genera ofoff Oregon. Smithson. Contrib. Hyperiidea (Crustacea: Amphipoda).

Zool.61. 8 pp.Smithson. Contrib. Zool. No. 146.Zool.61. 8 pp.64 pp.

Barnard, J.L. 1975. Identificationof gainaridean amphipods. Pages Bowman, T.E., C.D. Meyers, and S.D.314-366 in R.I. Smith and J.T. Hicks. 1963. Notes on the associa-Carlton, eds. Light's manual: tions between hyperiid amphipodsintertidal invertebrates off *the and medusae in Chesapeake andcentral California coast. Narragansett Bays and the NianticUniversity of California Press, River. Chesapeake Sci.Berkeley. 4(3): 141-146.

13

Caine, E.A. 1977. Feeding mecha- Estuarine Coastal Shelf Sci. (1981)nisms and possible resource parti- 12:645-653.tioning of the Caprellidae(Crustacea: Aaphipoda) from Puget Kozioff, E.N. 1973. Seashore life ofSound, U.S.A. Mar. Biol. 42: Puget Sound, the Strait of Georgia,331-336. and the San Juan Archipelago.

Caine, E.A. 1980. Ecology of two University of Washington Press,

littoral species of caprellid Seattle. 282 pp.

amphipods (Crustacea) fromWashington, U.S.A. Mar. Biol. 56: Kozloff, E.N. 1974. Keys to the327-335. marine invertebrates of Puget Sound,

the San Juan Archipelago, and adja-Chang, B.0., and T.R. Parsons. 1975. cent regions. University ofMetabolic studies on the aiphipod Washington Press, Seattle. 226 pp.Ant sogmmrus pugttensits i n

relation to its trophtc position Inthe food web of young salmonids. J. Levings, C.D., and 0. Levy. 1976.Ftsh. Res. Board Can. 32(2):243-247. A "bugs-eye" view of fish predation.

Pages 147-152 in C.A. Simenstad and

Congleton, J.C., and J.E. Smith. S.J. Lipovsky_, eds. Fish food

1976. Interactions between juvenile habits workshop, 1st Pac. N.W.

salmon and benthic invertebrates in Tech. Workshop, Astoria, Oreg. Oct.

the Skagit salt marsh. Pages 31-35 13-15.in C.A. Simenstad and S.J. Lipovsky,e-ds. Fish food habits workshop, 1st Mason, J.C. 1974. Behavioral ecologyPac. N.W. Tech. Workshop, Astoria, of chum salmon fry (OncorhynchusOreg. Oct. 13-15. keta) in a small estuary. J. Fish.

IIs Board Can. 31(1):83-92.Conlan, K.E., and E.L. Bousfield.

1982. Studies on amphipod crus- Mathias, J.A., J. Martin, M. Yorkow-taceans of the northeastern Pacific ski, J.G.I. Lark, M. Papst, and J.L.region 2. Family Amphipodae. Natl. Tabachek. 1982. Harvest andMus. Can. Publ. Biol. Oceanog. No. nutritional quality of Gammarus10: 41-75. lacustris for trout culture.Tirans.

A6l.Fish. Soc. 111(1): 83-89.Cross, J.N., K.L. Fresh, B.S. Miller,

C.A. Simenstad, S.N. Steintort, andJ.C. Fegley. 1978. Nearshore Matthews, L.H. 1978. The naturalfish and macroinvertebrate history of the whale. Weidenfeldassemblages along the Strait of Juan and Nicohson, London. 219 pp.de Fuca including food habits ofcommon nearshore fish. NOAA Tech. McCaln, J.C. 1975. Phylum Arthro-Memo. ERL MESA-32. 1.88 p. poda: Crustacea, Amphipoda:

Caprellidea. Pages 367-376 in R.I.

Feller, R.J., and V.W. Kaczynski. Smith and J.T. Carlton, eds.

1975. Size selection predation of Light's manual: intertidal inver-juvenile chum saloon (Oncorhynchus tebrates of the central Californiauenileo c humsalmon (pryncuet coast, 3rd ed. University of Cali-

keta) on epibenthic prey in Puget fornia Press, Berkeley.'gound. J. Fish. Res. Board Can.32(8): 1419-1429.

McCaulley, J.E., R.A. Parr, and D.R.Griffiths, C. L. , and J. Stenton- Hancock. 1977. Benthic infauna and

Dozey. 1981. The fauna and rate maintenance of dredging. Water Res.of degradation of standard kelp. 11(1): 83-89.

14

McClusky, D.S. 1967. Some effects of juvenile salmonids in Hoodof salinity on the survival, moult- Canal, Washington. Pages 163-176ing and growth of Corophum voluta- in C.A. Simenstad and S.J. Lipovsky,tor (Amphipoda). J. aN . OlT. eds. Fish food habits workshop, 1stAssoc. U.K. 48(2): 607-617. Pac. N.W. Tech. Workshop, Astoria,

Oreg. Oct. 13-15.McClusky, 0.S. 1970. Salinity pre-

ference in Corph um volutator. J. Shyamasundari, K. 1973. Studies onMar. Biol. Assoc- U.K.5(3): 747- the tube-building amphipods752. (Corophium triaenonyx) (Stebbing)

from Visaknapatnam Harbor: effect ofPark, T.S. 1961. Tentative keys salinity and temperature. Biol.

to the gamarid amphipods of the San Bull. (Woods Hole) 144(3): 503-510.Juan area. Unpubl. manuscript,University of Washington. Friday Shyamasundari, K. 1976. Effects ofHarbor Laboratory, Friday Harbor. salinity and temperature on the

development of eggs on thePearcy, W.G. 1972. Distribution and tube-building amphipod Corophium

ecology of oceanic animals off Ore- triaenonyx (Stebbing). Biol. Bull.gon. Pages 351-377 in A.T. Pruter TWoods Hole) 150(2):286-293.and 0.L. Alverson, eds. TheColumbia River Estuary and adjacent Smith, J.E. 1980. Seasonality,ocean waters: bioenvironmental spatial dispersion patterns andstudies. University of Washington migration of benthic invertebratesPress, Seattle. 868 pp. in an intertidal marsh-sandflat

system of Puget Sound, Washington,Reish, D.J. 1959. An ecological and their relation to waterfowl,

study of pollution in Los Angeles- foraging and for feeding ecology ofLong Beach Harbors, California. staghorn sculpin, LeptocottusOccas. Pap. Allan Hancock Found. armatus. Ph.D. Dissertation.No. 22: 1-119. iTviiiity of Washington, Seattle.

177 pp.Reish, D.J. 1964. Studies on theMylitus edulis community in Alamitos Smith, J.L. and O.R. Mudd. 1976.ay, a-iornia. II. Population Maintenance dredging and the envi-variations and discussion of the ronment of Grays Harbor, Washington.associated organisms. Veliger 6: Appendix H: Impact of dredging on202-207. the avian fauna in Grays Harbor.

U.S. Army Corps of Engineers,Reish, D.J., and J.L. Barnard. 1979. Seattle District. 217 pp.

Chapter 11. Amphipods (Arthropoda:Crustacea: Amphipoda). Pages Staude, C.P., J.W. Armstrong, R.M.345-700 in C.W. Hart, ed. Thom, and K.K. Chew. 1977. AnPollution ecology of estuarine illustrated key to the intertidalinvertebrates. Academic Press, New gammaridean amphipods of centralYork. Puget Sound. Coll. Fish. Univ.

Wash. Contrib. No. 466: 1-27.Schmitt, W.L. 1968. Crustaceans.

University of Michigan Press, Ann Waldichuk, M., and E.L. Bousfield.Arbor. 204 pp. 1962. Amphipods in low-oxygen

marine waters adjacent to a sulfiteSimenstad, C.A. 1976. Prey orga- pulp mill. J. Fish. Res. Board Can.

nisms and prey comunity composition 19(6): 1163-1165.

15

0m

m. P...,,,~W.* U

alashi ngto Af im- i Ir b Wetlands Rsac Center ___no.P" IBiological Report 82(11 .69)*1

Unvrst of Washington111 101 Gause BoulvarSeale, WA~j 98195e~ Slidell, LA 70458

Species Profiles: Life Histories and Environmental August 1986Requirements of Coastal Fishes and Invertebrates(Pacific Northwest)--Amphipods

*71 5 Army Cop of Enier Reor No.-* TRatf OWL no.IL lMa~(imi aaSmi~s~m nm

Danies pros e Gilbert B. Pauleya, and Davidthranh PWory and eirnd e mentsofco asta aqu ac s Ty arereare to

aWashington Cooperative Fishery National Wetlands Research Center ioResearch Unit NASA-Slidell Computer Complex rus1t.aa o antm, orfollown

-University of Washington 1010 Gause Boulevard

Seattle WA 98195 Slidell a LA 70458

National Wetlands Research Center U.S. Amy Corps of Engineers I the mOr afmt*hdFish and Wildlife Service Waterways Experiment StationU.S. Department of the Interior P.O. Box 631Washington, DC 20240 Vicksburg, MS 39180

IL Suipint Na.fDe

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

Species profiles are literature summaries of the taxonomy, morphology, range, lifehistory, and environmental requirements of coastal aquatic species. They are prepared toassist in environmental impact assessment. Amphipods are ubiquitous in distribution.Hyperiidea are the third most abundant coastal marine crustacean zooplankton, following

acopepods and euphausids. Benthic Gamridea are an invaluable food source for manyeconomically important fish and invertebrate species. Lifestyles of the major amphipodgroups are varied. On the basis of the Index of Relative Importance (IRI), they comprisemore than hlf of the total IRI spectrum for 38 of 55 fish species in the Strait of Juande Fuca. They are reported to be indicators of heavily polluted areas.

17. OeTaIONit AnUlO a. 2(

Fisheries Food chains Oxygen

Estuaries Life cycles Growth

Salinity Water pollution Feeding Habits

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Dissolved oxygen requirements Life historySalinity requirements Hyperiidea ,

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