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1 FVE Biological Report J January 1989 AD-A T/h.824 species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (Pacific Southwest) AM PH IPODS SAN RnMCSCO BAY CALIFORNIA PACIFI OCEAN LOS ANGELES UFish and Wildlife Service U.S. Department of the Interior Coastal Ecology Group Waterways Experiment Station U.S. Army Corps of Engineers f C q)c)z

2 Biological Report 82(11.92) TR EL-82-4 January 1989 Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (Pacific Southwest) AMPHIPODS by #p Daniel J. Grosse and Gilbert B. Pauley Washington Cooperative Fishery Research Unit School of Fisheries University of Washington Seattle, WA Project Officer David Moran Acces n For National Wetlands Research Center NTIS CFA&I U.S. Fish and Wildlife Service Le'IC T Gause Boulevard Slidell, LA J Distribut ionf_ Performed for Avail3hilitY Codes Coastal Ecology Group - i.'lvail1 and/or Waterways Experiment Station Dist Special U.S. Army Corps of Engineers Vicksburg, MS and U.S. Department of the Interior Fish and Wildlife Service Research and Development National Wetlands Research Center Washington, DC

3 This series may be referenced as follows: U.S. Fish and Wildlife Service Species profiles: life histories and environmental requirements of coastal fishes and invertebrates. U.S. Fish Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL This profile,way be cited as follows: Grosse, D.J., and G.B. Pauley Species profiles: life histories and environmental requirements of coastal fishes and invertebrates (Pacific Southwest)--Amphipods. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.92). U.S. Army Corps of Engineers, TR EL pp. 0

4 PREFACE is species profile is one of a series on coast:i 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 the following addresses. one of Information Transfer Specialist C)/ National Wetlands Research Center V U.S. Fish and Wildlife Service NASA-Slidell Computer Complex 1010 Gause Boulevard Slidell, LA or U.S. Army Engineer Waterways Experiment Station Attention: WESER-C Post Office Box 631 Vicksburg, MS ~iii

5 0 CONVERSION TABLE Multiply Metric to U.S. Customary To Obtain millimeters (mm) inches centimeters (cm) inches meters (m) feet meters (m) fathoms kilometers (km) statute miles kilometers (km) nautical miles square meters (m 2 ) square feet square kilometers (km 2 ) square miles hectares (ha) acres liters (1) gallons cubic meters (m 3 ) cubic feet cubic meters (m 3 ) acre-feet milligrams (mg) ounces grams (g) ounces kilograms (kg) pounds metric tons (t) pounds metric tons (t) short tons kilocalories (kcal) British thermal units Celsius degrees (*C) 1.8(OC) + 32 Fahrenheit degrees U.S. Customary to Metric inches millimeters inches 2.54 centimeters feet (ft) meters fathoms meters statute miles (mi) kilometers nautical miles (nmi) kilometers square feet (ft 2 ) square meters square miles (mi 2 ) square kilometers acres hectares gallons (gal) liters cubic feet (ft 3 ) cubic meters acre-feet cubic meters ounces (oz) milligrams ounces (oz) grams pounds (Ib) kilograms pounds (lb) metric tons short tons (ton) metric tons British thermal units (Btu) kilocalories Fahrenheit degrees ( F) ( F - 32) Celsius degrees iv

6 CONTENTS PREFACE... i CONVERSION TABLE... iv FIGURES... vi ACKNOWLEDGMENTS... vii NOMENCLATURE/TAXONOMY/RANGE... 1 MORPHOLOGY/IDENTIFICATION AIDS... 4 REASON FOR INCLUSION IN SERIES... 5 LIFE HISTORY... 5 Reproduction and Fecundity... 5 Growth Characteristics... 5 Importance to Fisheries... 6 ECOLOGICAL ROLE... 8 ENVIRONMENTAL REQUIREMENTS Dissolved Oxygen and Temperature Salinity Pollution and Dredging LITERATURE CITED Page v

7 FIGURES Number 1 A gammaridean amphipod A. Elasmopus sp. and B. Eohaustorius sp., both gammarid amphipods. C. Caprella ferrea, a caprellid amphipod. D. Neocyamus physeteris female, a caprellid amphipod from sperm whale. E. Phronima sedentaria, a hyperlid amphipod that lives inside the tunic of urochordates Distribution of the ubiquitous amphipod suborders Gammaridea and Hyperiidea along the coastal areas of central and southern California Growth of Anisogammarus pugettens is fed Enteroenorpha intestinalis at 10 and 20 'C Fish, avian, and invertebrate predators of the amphipod Corophium salmonisi... 7 Pg vi

8 ACKNOWLEDGMENTS We gratefully acknowledge reviews by Rick Albright, School of Fisheries, University of Washington, Seattle, and Craig P. Staude, Friday Harbor Laboratories, University of Washington, Seattle. vii

9 Pereon -- Pleon -lead-x Rostrum-2 Gland Cone- Accessory.- Flagellum A 4 5 U3 '"Telson Urosome Gnathopods Antennae Pereopods Pleopods Uropds '0 Figure 1. A gammaridean amphipod (from Staude et al. 1977). AMPHIPODS NOMENCLATURE/TAXONOMY/RANGE into Oregon and Washington and southward into Baja California Scientific name... Amphipoda 'farnard 1969a-. - Gammaridea are the Preferred common name... Amphipod most abundant and diverse group of (Figure 1) amphipods, A table of northern and Class... Crustacea southern aphipods was assembled by Subclass... Malacostraca Barnard (1969a). More than 25% of Order... Amphipoda the amphipods in California are of Suborders... Gammaridea, Hyperiidea, unknown geographic affinity. About Caprellidea, Ingolfiellidea (Figure one-third of southern California 2). 1 I species are "cosmopolitan," and 2 one-third of the northern California Geographic range:"this report focuses species inhabit boreal waters of the largely on the suborders Gammaridea eastern and western Pacific. The and Hyperiidea because of their shift from cold-temperate to importance in coastal waters of the warm-temperate environments is Pacific coast region of the South- reflected at Point Conception, which western United States -{ w-e--3): is the northern boundary of many Many of the California amphipod southern species ind the southern species are ubiquitous along the boundary of many northern species Pacific coast and extend northward (Barnard 1969a). -- Tr), -To, WOLP-t VOH, I. CLn A Vc4 k'~r

10 A B C Figure 2. A. Elasmopus sp. and B. Eohaustorius sp., both gammarid amphipods. C. Caprella ferrea, a caprellid amphipod. D. Neocyamus physeteris female, a caprellid amphipod from sperm whale. E. Phronima sedentaria, a hyperiid amphipod that lives inside the tunic of urochordates. (A and B from Barnard 1975; C and D from McCain 1975; E from Barnes A-D reprinted with permission from the University of California Press; E reprinted with permission from Saunders College Publishing). 2

11 CAPE MENDOCINO'-, f/. / //, //,7?; I -/', OCEANIC DISTRIBUTION 0, ESTUARY AND K CONCENTRATED.g AREAS OF A DISTRIBUTION SAN FRANCISC& i, N FRANCISCO MON TEREY,, - l". \ NEVADA -.- \ -., 0,,,-. -. CALIFORNIA PACIFIC OCEAN 1;0e /, POINT CONCEPTION AjA #, LOS ANGELES 0 so 100 loop -0 o so' o' 0 * SAN DIEGO KILOME TIE S Figure 3. Distribution of the ubiquitous amphipod suborders Gammaridea and Hyperiidea along the coastal areas of the Pacific Ocean off central and southern California. 03

12 The generic composition of intertidal amphipods in California overlaps thoracic legs (pereopods): the dactyls of the anterior four pair are directed that of similar forms throughout the posteriorly, while the dactyls of the world (60% with Indo-Pacific tropics, posterior legs point anteriorly. 51% with Japan's Okhotsk Sea, and 46% Gills are usually present at the base with the Isle of Man). A total of 174 of pereopods 2-6, protected by the genera and 1,118 species of rocky ventrally expanded coxal plates. intertidal amphipods have been identi- Males and females often can be disfied north of latitude 45 *S. Sixty- tinguished morphologically. The head three genera, or about one-third of has five fused segments, with two the world's intertidal genera, are present in California (Barnard 1969b). pairs each of antennae and maxillae and a heavily chitinized mandible. California's two climates, temperate in the north and subtropical in the There are six or seven freely articu- lated somites on the thorax (pereon). south, are one reason for the many known amphipod taxa in that State. Plates (coxae) are lateral extensions of the thoracic pereon. Gills Another reason is that amphipods have (branchiae) are fleshy and plate-like been studied in far greater detail for and are attached medial to the second many years in California than any through the sixth coxae on each side. other place along the Pacific coast, The abdominal region consists of three so there is a more thorough list of articulating segments on both the taxa. The most abundant species of anterior pleon and posterior urosome. amphipods in California are frequently The urosome has a terminal telson in the most diverse genera (primarily (Figure 1). marine), although amphipods also inhabit freshwater and some moist ter- The following key (adapted from restrial habitats (Reish and Barnard Barnes 1974 and Kozloff 1974) is an 1979). The marine forms live at most aid to separate ampt~ipod suborders: depths, including deep abyssal waters (Hessler et al. 1978), and in a wide la. Pereon with seven apparent segrange of habitats. About 40% of the ments, all having well-developed 80 genera of Gammaridea are common appendages. Abdomen not vestiworldwide, while the remaining 60% are gial. Body neither slender nor loosely associated with specific geo- resembling that of a praying graphical regions or zones (Bousfield mantis ). lb. Pereon with six apparent segfound fudiamotvestigial in almost ments, some appendages; of which may abdomen have Gammarid species are all environments: subtidal, inter- vestigial heafs wit ofrs tidal, freshwater, and terrestrial vestigial; heaedfuor d with first (Reish and Barnard 1979). The and second thoracic segment. Hyprideaar enirly arne ndbody slender and (except for Hyperiidea are entirely marine and whale lice) resembling that of a pelagic (Bowman and Gruner 1973). praying mantis. Marine. Includes MORPHOLOGY/IDENTIFICATION AIDS skeleton shrimp... Caprellidea. Suborder 2a. Eyes generally large, occupying The Amphipoda are distinguished from most of head; coxae of pereopods other crustacea by their unstalked small, often fused with the body; eyes, lack of a carapace, lateral maxillipeds are without palp; compression of the body, and the last two abdominal segments structure of the last three append- fused; body more or less transages (uropods) of the pleon. Amphi- parent. Marine, and usually pods have seven pairs of major planktonic or associated with 4

13 jellyfish or in tunics of dead LIFE HISTORY salps... Suborder Hyperiidea. Reproduction and Fecundity 2b. Eyes usually present and conspicuous, but not large enough to When mating, the male amphipod holds cover most of the head; coxae of the female in a copulatory embrace pereopods well developed, usually (amplexus). Some species have an expanded. Marine, freshwater, extended precopulatory ritual (preand terrestrial... Suborder amplexus), while others do not Gammaridea. (Borowsky 1984). In swimming species, the male often carries the female 2c. Eyes small; body elongate; small ventrally, or both swim on their coxae; abdominal segments dis- sides. Following ecdysis of the tinct; all but fourth and fifth female (molting of the exoskeleton), pairs vestigial. of abdominal Marine, interstitial, appendages eggs are laid through two ventral pores in the sixth thoracic sternite. Rare... Suborder Ingolfiellidea. Fecundity may exceed 200 eggs per female (Barnard 1969b), but infaunal The most concise identification species tend to have fewer eggs than guides to the marine amphipods of the epifaunal species (Nelson 1980; Van Pacific Southwest region are those of Dolah and Bird 1980). Mature eggs Barnard (1975) and McCain (1975). hatch directly into juveniles that Bousfield (1958) may be useful for resemble adults. These juveniles are identifying freshwater gammaridea. usually held in the brood pouch for a few hours to a few days after hatching, then released. They can REASON FOR INCLUSION IN SERIFS then feed and return to the pouch for protection (Barnard 1969b; Reish and The benthic amphipods, especially Barnard 1979). Gammaridea, are an invaluable food source for many economically important Information on the reproductive fishes (Gerke and Kaczynski 1972; cycles of pelagic species of amphipods Kaczynski et al. 1973; Mason 1974; is scarce and difficult to obtain in Hobson and Chess 1976). Their limited the field. In some hyperiids, the mobility and their sensitivity to male and female apparently cohabit the environmental changes suggest that same medusa prior to copulation their distribution and abundance can (Sheader 1977). Brusca (1967b) be used as an indicator of environ- observed several families of pelagic mental quality (Albright 1982). amphipods off the coast of southern Omnivorous and opportunistic feeders California and found that the highest such as lysianassids (a gammaridean production of ova occurred during the family) and caprellids recycle summer and fall months, and that detritus and play an important role development of the young continued in the ecosystem by scavenging car- through the following spring and casses of large animals following mass summer. mortalities (Keith 1969; Reish and Barnard 1979). Amphipods, in addi- Growth Characteristics tion to being scavengers on fish carcasses, are also predatory to some Amphipod growth rates and lengths degree on small fishes (Westernhagen vary considerably. Like all crus- -i Rosenthal 1976; Hessler et al. taceans, amphipod growth takes place 1978; Stepien and Brusca 1985). at each molt when the old exoskeleton Hyperiid amphipods are one of the most is shed. Amphipods range in length abundant groups of coastal marine from under 1 cm to about 28 cm, the crustaceans (Bowman and Gruner 1973). largest of which is a lysianassid 5

14 photographed in the abyssal Pacific every 20 to 30 days. The average in- Ocean (Hessler et al. 1978). Maximum star (stage of development between growth rates of Anisogammarus successive molts) lasts 15 days. pugettensis were 4.1% of dry weight Gammarids go through at least 12 inper day at 10 'C, increasing more than stars. The maximum life-span estithreefold to 14.3% at 20 0 C (Figure mates are a little more than 6 months 4), with higher growth efficiency at in many species, but some polar 20 0 C (Chang and Parsons 1975). As species are known to live 5 or 6 with most aquatic organisms, tempera- years. Females commonly lay eggs ture has a significant effect on the either during each of the last five or growth rate. Growth in large (10 mg) six instars, or at every other instar individuals of this species was 47% to (Barnard 1969b). 72% of food intake when fed Enteromorpha (Chang and Parsons The Importance to Fisheries growth rate in Gammarus pulex is 63% faster in males than in females, and Amphipods are the main food of females achieve a lower final mean many species of fish (Kaczynski et al. weight (52 mg) than males (65 mg), 1973; Hobson and Chess 1976). Pelagic according to Sutcliffe et al. (1981). species sometimes compose the bulk of the diet of herring, mackerel, and Growth is initially rapid in the Biscayan tunny (Schmitt 1968). Gammaridea; molting may begin shortly Gammarideans, on the basis of an after hatching and continues through Index of Relative Importance (IRI), maturity. As amphipods increase in were the most important food for size, molting usually slows to once nearshore fishes in the Strait of Juan de Fuca. They composed more than half the total food eaten by 38% of the 55 fish species studied (Cross et a' ), and they were the most impot - tant food of tidepool fishes. A tube-dwelling gammaridean, Coro C phium salmonis, is an abundant and preferred prey of chum salmon 10 locmarsh (Oncorhynchus keta) in the Skagit salt in Wasington (Congleton and Smith 1976), as well as in other areas of Puget Sound (Gerke and Kaczynski ). Albright (1982) reported that densities of C. salmonis peaked in the tidal flats of Grays Harbor, Washington, in July and August, where they were the dominant organism on mud and.01- muddy-sand bottoms. Densities as high 'as ,000/M 2 have been observed (Albright and Rammer 1976). Produc- Weeks tion from April through September was g dry weight/m 2. According to Albright (1982), C. salmonis is Figure 4. Growth of Anisogammarus consumed by a large number of fish pugettensis fed Enteromorpha intes- species (Figure 5), including salmon, tinalis at 10 and 20 -C. (Chang and sculpins, sticklebacks, gunnels, Parsons 1975; reprinted with permis- smelts, cod, sole, flounders, and sion from the Journal of the Fisher- pricklebacks. Fish that are predators ies Research Board of CanadaT. on amphipods and other zooplankton in 6

15 Western Sandpiper Pacific Staghorn Sculpin Eogammarus Juvenile Salmon Dunlin Long-billed Sie ec Dowitcher S Perch Starry Flounder English Sole Snake Prickleback Stickleback Juvenile Pacific Tomcod Dungeness Crab Longfin Smelt Saddleback Gunnel Nemertean Worms Crangon Shrimp Figure 5. Fish, avian, and invertebrate predators of the amphipod Corophium salmonisi (from Albright 1982). California have evolved morphologi- for young salmon, but the brine shrimp cally elaborate feeding mechanisms and grew much faster. This gammaridean body forms; the degree of divergence can tolerate wide ranges of temperafrom the basic body plan depends on tures and salinities and eats a wide how extensively they feed on zooplank- variety of plant and animal material, ton (Hobson and Chess 1976). Crusta- in addition to scavenging dead fish cean zooplankton, including amphipods, and uneaten fish food in ponds. may significantly influence nocturnal Gammarus lacustrus, in the shallow versus diurnal distributions and be- prairie lakes of the Hudson Bay drainhavior of nearshore fishes of Cali- age, meets dietary requirements for fornia (Hobson and Chess 1976; Stepien rainbow trout (Salmo gairdneri) that and Brusca 1985). are 5 cm long oronger. These gammarids are easily captured and can be Two gamarid species have been harvested at 1,000 kg wet weight/ha/ examined for their potential in fish yr, are equal to or better than availculture. Mass culture of Anisogam- able commercial trout feeds, and can marus pugettensis was proposed by improve body coloration and market- Chang and Parsons (1975) as an ability of the fish (Mathias et al. alternative to brine shrimp as food 1982). Gammarus tigrinis, a shoreline 7

16 amphipod, has been introduced into (Barnard 1969b). Corophium spp., brackish streams as food for fishes common in estuaries where silting is (Reish and Barnard 1979). heavy, form masses of muddy tubes and create currents with abdominal appendages ECOLOGICAL ROLE (Albright 1982). The currents are strained by fringes of fine hairs on appendages forward of the abdomen; Amphipods are considered the most then the selectively collected materefficient scavengers of sea bottoms ial is scraped into the mouth (Kozloff and shorelines, where they probably 1973). clear up and recycle more organic nearshore debris than any other animal Some amphipods inhabit dwellings of group (Schmitt 1968). Griffiths and other organisms (Kozloff 1973). Many Stenton-Dozey (1981) described the species that are burrowers, such as importance of the gammarid, Talor- those of the gammaridean families chestia capensis, in consuming Haustoriidae, Oedicerotidae, and beached kelp in South Africa. This Phoxocephalidae, have elongated spines amphipod and dipteran larvae ate 60% or setae on the distal articles of the to 80% of the beached kelp within 2 posterior pereopods that represent an weeks, and it is thought that they adaptation for burrowing (Reish and make a significant contribution Barnard 1979). (through feces) to organic enrichment in coastal waters. Tube-dwelling amphipods almost never Numerically, amphipods are the major dominate a rocky area pounded by waves. Wave action is usually too component of macrofauna on harbor severe unless protection is given by pilings in California. Most are encrusting organisms. Beds of mussels introduced species (carried into ports of the genus Mytilus serve as excelby foreign vessels) that have had little effect on indigenous amphipods lent protection for amphipods in rocky intertidal areas (Tsuchiya and 0 in nearby water (Barnard 1961; Reish Nishihira 1985). 1964). In heavily polluted harbors, amphipods are scarce in both the About one-third of all amphipod benthos and on the pilings (Reish species in the intertidal areas of 1959). California are tube-dwelling forms, Beachhoppers of the gammaridean compared sediment with only burrowers. 2% that are Phoxocephalids family Talitridae are common on sandy are the major sediment burrowers in intertidal areas, especially among the intertidal zone of southern damp algal debris or wracks (Reish and California (Barnard 1969b). Barnard 1979). These species are locally transitory because of frequent Even among closely related species, changes in the tide and wrack accumu- amphipods have become highly speciallations. The maximum density of other ized (Caine 1980). Many intertidal amphipods of sandy beaches is related and estuarine amphipods appear to to surf intensity and varies with sea- occupy distinct and generally nonson (Hughes 1982). The obligate sand- overlapping niches, which may be burrowing amphipods belong to two separated by one or more environmental major groups withih the family differences (Bousfield 1970; Caine Haustoriidae and are common in south- 1977, 1980; Pinkster and Broodbakker ern temperate waters (Bousfield 1970). 1980; Gunnill 1984). This separation of niches apparently holds true along Some gammarideans, such as Ameplisca the Pacific coast from Washington sp. and Photis sp., construct tubes or (Caine 1980) to California (Gunnill cradles on soft or hard substrates 1984). 8

17 09 Examples of amphipods as indicators habits are poorly understood. Hyperof environmental conditions are iids may feed on the very organisms Pontogeneia and Lysianassa, which are that host them, but probably use them typical of sand-encroachment, and more as a base from which to forage; Parallorchestes, which is typical of or, they may feed on food captured by the wave-dash intertidal zone (Barnard the host (Bowman et al. 1963; Patton 1969b). In areas where these condi- 1968). In one laboratory study of tions mix, these amphipods, along with Lestrigonus sp. and Bougisia sp., food Hyale and Aoroides, are all found was shared with the host, Leptomedusa together. The presence of certain phoxocephalids indicates stratified sp., when supply was adequate, but when it was not, the amphipods fed on and relatively undisturbed sediment. host tissue, starting with the gonads The holdfasts of large kelp (Bowman and Gruner 1973). Parathemisto sp., a free-living hypf id, (Macrocystis) in the subtidal zone preys on other plankters (Bowman host many species of amphipods, some 1960). of which are rare or nonexistent intertidally. An unusually large number of different amphipod species A few nektonic gammarideans live in neritic waters. They are either live exclusively or most frequently on predaceous or are the nektonic mating kelp holdfasts (Barnard 1969b). Hyale or dispersal phases of benthic frequens, the most abundant intertidal gammarids (Reish and Barnard 1979). amphipod in California, lives on or near surf grasses and kelp holdfasts, Chelura terebrans, a wood-borer in or in tidepools (Barnard 1969b). coastal waters principally south of Hyale grandicornis lives among algae San Francisco, is the best known associated with mussels (Mytilus amphipod pest. It enlarges holes in edulis) and barnacles (Chthamalus wood (e.g., boats and pilings) made by challengeri), according to Tsuchiya the isopod Limnora sp. (Reish and and Nishihara (1985). It has been Barnard 1979). observed by Gunnill (1984) that more than one species of amphipod in Swimming among amphipods varies southern California used the brown greatly among the various genera. alga, Pelvetia fastigiata, but that Hyperiidean swimming ranges from the they occupied different niches-- feeble movements of the appendages of Ampithoe tea, A. lindbergii, and A. Cystisoma sp., to the fast swimming pollex lived at the distal ends of the Paraprone spp. which are characterized algal fronds, while Photis spp., by a strong pleonal musculature Corophium spp., and Aorides columbiae (Bowman and Gruner 1973). Most of the lived near the plants' holdfasts. gammarideans--even the burrowing forms--are strong swimmers. The Coralline stands and sedimentary paddling motion of their pleopods is substrates beneath rocks are poor in some cases facilitated by small amphipod habitats. Amphipods that coupling hooks that join the peduncles bu-row into the substrate have of each pair of pleopods. Some definite preferences for habitats and gammarids that live on the sea bottom particle sizes. have elongated pereopods that spread out like a spider to prevent them from Hyperiidea are primarily nektonic, sinking into the mud. Their bodies although Hyperia can be taken in hang upside down, giving them a lower benthic samples. They either have well-developed swimming appendages and center of gravity. This helps avoid displacement by turbulence. Epibenbuoyancy control, or live in associa- thic gammarids may reduce their tion with host medusae or salps (Reish susceptibility to predation by and Barnard 1979). Their feeding swimming. Feller and Kaczynski (1975)

18 believed that in the spring, juvenile Caine 1977, 1980). Whale parasites chum salmon in Puget Sound preferred (Cyamidae) are also in the caprellid harpactacoid copepods because they suborder. The genus Cyamus includes were more easily captured than about 18 host-specific species. As swimming amphipods. non-swimmers, they leave the parental brood pouch and dig into the host with Anisogammarus conferviculus is hooked dactyls (Schmitt 1968). believed to reduce predation, largely by juvenile chum salmon, through It is suggested that the common ecological adaptation. Its avoidance pelagic amphipod species are more behavior includes clumping in refuges with structurally complex habitats, abundant offshore than inshore in oceanic waters and that yearly changes such as bottom vegetation (Levings and in occurrence and abundance of Levy 1976). In Grays Harbor, Washing- hyperiid amphipods inshore may be ton, mature male Corophium salmonis related to coastal upwelling (Lorz and are subject to heavy predation from a Pearcy 1975). Most hyperiid amphipods variety of sources (Figure 5) begin- live in the upper 100 m of the ocean ning in April, when they wander over in the North Pacific central gyre and tidal flats in search of females exhibit diurnal vertical migration (Albright 1982). In addition to being (Schulenberger 1978). Off the coast prey for many fishes and inverte- of southern California, both brates, some pelagic amphipods compose Gammaridea and Hyperiidea have been part of the crustacean diet of whales, and the diet of the grey whale along found at depths greater than 650 m; the depth of their upper limit was the west coast consists largely of six defined by the thermocline and the species of benthic amphipods (Matthews amount of light available (Brusca 1978). Amphipods sometimes are eaten 1967a). Amphipods from this area by British gulls (Larus sp.) according exhibit vertical diurnal movements to Schmitt (1968) and by dunlin (Brusca 1967a). (Calidrus alpina) according to Smith and Mudd (1976). Dogielinotus loquax is a prime target for summer shorebird predation (Hughes 1982). ENVIRONMENTAL REQUIREMENTS Dissolved Oxygen and Temperature Caprellids, of the suborder which includes skeleton shrimp, are largely Pelagic gammarid and hyperiid intertidal. Their preference of amphipods have been collected in deep, substrate is often specific. They poorly oxygenated scattering layers usually cling to living substrate such off southeastern Vancouver Island, as kelp, sea grasses, sponges, British Columbia (Waldichuk and hydroids, and bryozoans; to a lesser Bousfield 1962). Anisogammarus extent, they live on bare sand or mud pugettensis and Allorchestes angustus, bottoms (Keith 1971; Caine 1980). both common inshore gammarid They are relatively motionless and amphipods, survive in water with low feed by grasping food with their free dissolved oxygen as low as 0.04 ppm at anterior legs and antennae, and hold- 12 C near sulfite-rich paper pulp ing their position with their poster- effluent in British Columbia ior legs. Locomotion resembles that (Waldichuk and Bousfield 1962). The of an inchworm with an alternating first species is abundant on the movement of the front and rear legs bottom at 15 to 22 m; the second (Kozloff 1973). They feed on diatoms, species, normally found in shallower small invertebrates, and detritus, and waters, was near the surface, perhaps are in turn the prey of shrimp and seeking more oxygenated water many fishes, including cod, blennies, (Waldichuk and Bousfield 1962). Low and skates (Keith 1969; McCain 1975; oxygen tolerance in either species 10

19 remains to be determined, but Chang preferred a range of 10 to 30 ppt and Parsons (1975) observed that A. (McClusky 1970). Adult C. triaenonyx pugettensis survived for several hours survived in a range of salinities at 20% saturation. They also deter- similar to that at which C. volutator mined a Q1o of 1.6, lower than that of survived (Shyamasundari 1973). other crustaceans, which is generally Although juvenile C. triaenonyx grew near 2. (Q 1 o is a measure of the at salinities of 7.5 to 37.5 ppt, they change in physiological processes with survived and grew best at salinities temperature. If metabolism (and hence of 20 to 32.5 ppt (Shyamasundari oxygen consumption) doubles when the 1973). A. pugettensis, found natutemperature is increased by 10 'C, an rally in salinities of 20 to 28 ppt, organism has a Q1o of 2. If there is cannot survive in freshwater, but can no rate change with temperature, the survive at 11 ppt for at least 1 week animal has a Q1o of 1 and is said to (Chang and Parsons 1975). Some be temperature-independent.) Chang species of gammaridean genera, such as and Parsons (1975) believed this low Gammarus, Hyalella, and Crangonyx, Q1o to be an adaptation of amphipods live in freshwater. for coping with rapidly changing intertidal temperatures. Caprellids In laboratory experiments using leave eelgrass beds in Tomales Bay in estuarine amphipods, Pinkster and droves at night when dissolved oxygen Broodbakker (1980) found that (1) concentrations in the beds drop below survival time of ovigerous females and 2 ppm (Keith 1971). eggs increased with increasing salinity, (2) males survived better at Tolerances to low oxygen concentra- lower salinities than females, (3) tions vary greatly among species. females produced more batches of eggs Many are intolerant of low oxygen at higher salinities, (4) time between concentrations, especially species ovipositions was shorter at higher restricted to waters where dissolved chlorinities, and (5) females produced oxygen usually is high. Groups such more eggs at higher salinities. as phoxocephalids (used as indicators of pollution in bioassays) appear much Pollution and Dredging less tolerant to stressful conditions, such as low oxygen concentrations, Some amphipod species are more tolthan many of the species discussed erant than others of organic polluabove (R. Albright, University of tion, but the reasons why are not Washington; pers. comm.). clear (Reish and Barnard 1979). Allorchestes compressa was found to be Caprella laeviuscula and Meta- the species most sensitive to heavy caprella kennerlyi can survive at metals among the seven species tested temperatures as high as 20 'C, while (Reish and Barnard 1979). Capitella, Caprella striata will only survive at a mari.e polychaete commonly used as a temperatures up to 14 'C (Caine 1980). pollution considered indicator mutually and generally exclusive to Salinity amphipods, has been observed in heavily polluted harbors. Capitella Many species of adult gammarideans also lives in unpolluted deep sea withstand high variations in salinity, waters near coastal California that while some juveniles and embryos are are subject to freshwater inflow-- less capable of doing so (although in habitats where amphipods are notably other species, the juveniles are absent (Reish and Barnard 1979). actually more tolerant than adults). Adults of Corophium volutator, and The construction of harbors has had estuarine species, survived salinities little overall effect on amphipod of 2 to 59 ppt (McClusky 1967), but populations native to California 11

20 because the animals are so abundant up are most likely to be affected by oil and down the coast (Reish and Barnard (Baker 1971), as are populations of 1979). Nonetheless, amphipods are subtidal ampeliscids. Dredging is rare in muddy substrates near docks in likely to at least temporarily elimi- Los Angeles, San Francisco, and San Diego (Reish and Barnard 1979). Hownate benthic amphipods that live on or close to the substrate (Albright and ever, delta mudflat areas do contain numerous amphipods; densities of Ramer 1976; Reish and Barnard 1979). However, McCaulley et al. (1977) Corophium salmonis become as high as suggest that when dredging occurs, 120,000/m 2 (Smith 1977). adults of some species are likely to move to nearby unaffected areas and The distribution of Corophium juveniles may rapidly immigrate and salmonis is influenced by sediment repopulate the dredged area. type and depth (it prefers shallow, muddy sand substrates), as well as salinity (Albright 1982). It is A recolonization of benthic orgathought that dredging likely causes a nisms after attempts at pollution net short-term reduction in the control in the consolidated Slip-East numbers of Corophium spinicorne, Basin area (f Los Angeles Harbor was resulting in a substantial impact on described by Reish (1959). Although its fish and invertebrate predators many groups of invertebrates recolo- (Albright and Borithilette 1982). nized the area rapidly, amphipods Other species of Corophium are abun- recovered much more slowly. Albright dant near sewer outfalls, possibly and Borithilette (1982) noted that it because of organic enrichment may take up to a year to repopulate an (Birklund 1977). area with all the invertebrates that Behavioral changes of amphipods were present prior to dredging, but that opportunistic species such as the exposed to sublethal quantities of oil amphipods Corophium spinicorne and C. have been observed. Populations of salmonis may repopulate the area much intertidal gammaridean beachhoppers more quickly. 12

21 LITERATURE CITED Albright, R Population dynam- Barnard, J.L Identification ics and production of the amphipod of gammaridean amphipods. Pages Corophium salmonis in Grays Harbor, in R.I. Smith and J.T. Washington. M.S. Thesis. Univer- Carlton, eds. Light's manual: sity of Washington, Seattle. 76 pp. intertidal invertebrates of the central California coast. Univer- Albright, R., and P.K. Borithilette. sity of California Press, Berkeley Benthic invertebrate studies in Grays Harbor, Washington. U.S. Barnes, R.D Invertebrate Army Corps Eng., Seattle Dist., Con- zoology, 3rd ed. W.B. Saunders Co., tract Rep. DACW67-80-C pp. Philadelphia. 870 pp. Albright, R., and A.D. Ramer Birklund, J Biomass, growth Maintenance dredging and the envi- and production of the amphipod ronment of Grays Harbor, Washington. Corophium insidiosum (Crawford) and Appendix E: The effect of inter- preliminary notes on Corophium tidal dredged material disposal on volutator (Pallas). Ophelia 16(2): benthic invertebrates. U.S. Army Corps of Engineers, Seattle District. 244 pp. Borowsky, B The use of the male's gnathopods during pre- Baker, J.M Growth simulation copulation in some gammaridean following oil pollution. Pages amphipods. Crustaceana (LEIDEN) in E.B. Coweel, ed. The 47(3): ecological effects of oil pollution on littoral communities. Institute Bousfield, E.L Freshwater of Petroleum, London. amphipod crustaceans of glaciated North America. Can. Field-Nat. Barnard, J.L Relationship of 72(2): California amphipod faunas in Newport Bay and in the open sea. Bousfield, E.L Adaptive radi- Pac. Nat. 2: ation in sand-burrowing amphipod crustaceans. Chesapeake Sci. Barnard, J.L. 1969a. Gammaridean 11(3): Amphipoda of the rocky intertidal of California: Monterey Bay to Bousfield, E.L A revised LaJolla. U.S. Natl. Mus. Bull classification and phylogeny of 230 pp. amphipod crustaceans. Trans. R. Soc. Can. Ser. 4(16): Barnard, J.L. 1969b. The families and genera of marine gammaridean Bousfield, E.L Evolution in Amphipoda. U.S. Natl. Mus. Bull. North Pacific coastal marine pp. amphipod crustaceans in G.G.E. * 13

22 Scudder and J.L. Reveal, eds. Anisogammarus pugettensis in rela- Evolution today. Proc. Second tion to its trophic position in International Congress of Syste- the food web of young salmonids. J. matics and Evolutionary Biology. Fish. Res. Board Can. 32(2): Bowman, T.E The pelagic Congleton, J.C., and J.E. Smith. amphipod genus Parathemisto Interactions between juvenile (Hyperiidea:Hyperiidae) in the North salmon and benthic invertebrates in Pacific and adjacent Arctic Ocean. the Skagit salt marsh. Pages Proc. U.S. Natl. Mus. 112(3439):343- in C.A. Simenstad and S.J. Lipovsky, 392. eds. Fish food habits workshop, First Pacific Northwest Technical Bowman, T.E., and H. Gruner Workshop, Astoria, Oregon. Oct. The families and genera of pp. Hyperiidea (Crustacea:Amphipoda). Smithsonian Contrib. Zool. No Cross, J.N., K.L. Fresh, B.S. Miller, 64 pp. C.A. Simenstad, S.N. Steintort, and Bowman, T.E., C.D. Meyers, and S.D. J.C. Fegley Nearshore fish Hicks Notes on the associ- and macroinvertebrate assemblages ations between hyperiid amphipods along the Strait of Juan de Fuca and medusae in Chesapeake and including food habits of common Narragansett Bays and the Niantic nearshore fish. NOAA Tech. Memo., River. Chesapeake Sci. ERL MESA pp. 4(3): Feller, R.J., and V.W. Kacqunski Size selective predation by Brusca, G.J. 1967a. The ecology of juvenile chum salmon (Oncorhynchus pelagic amphipods, I. Species keta) on epibenthic prey in Puget accounts, vertical zonation and Sound. J. Fish. Res. Board Can. migration of amphipods from the 32(8):1' waters off southern California. Pac. Sci. 21(3): Gerke, R.J., and V.W. Kaczynski Brusca, G. J. 1967b. The ecology of Food of juvenile pink and chum salpelagic amphipods, II. Observations mon in Puget Sound, Washington. on the reproductive cycles of Wash. Dep. Fish. Tech. Rep. No. several pelagic amphipods from the 10:1-27. waters off southern California. Griffiths, C.L., and J. Stenton-Dozey. Pac. Sci. 21(4): The fauna and rate of degradation of standard kelp. Estuarine Caine, E.A Feeding mechanisms Coastal Shelf Sci. 12: and possible resource partitioning of the Caprellidae (Crustacea: Gunni1l, F.D Differing dis- Amphipoda) from Puget Sound, USA. tributions of potentially competing Mar. Biol. (Berl.) 42(3): amphipods, copepods, and gastropods among specimens of the intertidal Caine, E.A Ecology of two alga Pelvetia fastigiata. Mar. littoral species of caprellid amphi- Biol. (Berl.) 82(3): pods (Crustacea) from Washington, USA. Mar. Biol. (Berl.) 56(3):327- Hessler, R.R., C.L. Ingram, A.A Yayanos, and B.R. Burnett Scavenging amphipods from the floor Chang, B.D., and T.R. Parsons Metabolic studies on the amphipod of the Philippine Trench. Res. 25(11): Deep-sea 14

23 Hobson, E.S., and J.R. Chess off the Oregon coast. J. Fish. Res. Trophic interactions among fishes Board Can. 32(8): and zooplankters near shore at Santa Catalina Island, California. U.S. Mason, J.C Behavioral ecology Natl. Mar. Fish. Serv. Fish. Bull. of chum salmon fry (Oncorhynchus 74(3): keta) in a small estuary. J. Fish. Res. Board Can. 31(1): Hughes, J.E Life history of the sandy-beach amphipod Dogielino- Mathias, J.A., J. Martin, M. tus loqu~a T ae -from (Crustacea: Dogielinothe outer coast of Yorkowski, J.G.I. Lark, M. Papst, and J.L. Tabachek Harvest Washington, USA. Mar. Biol. (Ber.) and nutritional quality of Gamarus 71(2): lacustris for trout culture. Trans. Kaczynski, V.W., R.J. Feller, J. Am. Fish. Soc. 111(l): Clayton, and R.G. Gerke Trophic analysis of juvenile pink Matthews, L.H The natural and chum salmon in Puget Sound. J. history of the whale. Weidenfeld Fish. Res. Board Can. 30(7):1003- and Nicohson, London. 219 pp McCain, J.C Phylum Arthro- Keith, D.E Aspects of feeding poda:crustacea, Amphipoda: in Caprella californica Stimpson and Caprellidea. Pages in R.I. Caprella equilibra Say (Amphipoda). Smith and J.T. Carlton, eds., Crustaceana 16(2): Light's manual: intertidal invertebrates of the central California Keith, D.E Substrate selec- coast. 3rd ed. University of Calition in caprellid amphipods of fonnia Press, Berkeley. Southern California, with emphasis on Caprella californica Stimpson and McCaulley, J.E., R.A. Parr, D.R. Caprella equilibra Say (Amphipoda). Hancock Benthic infauna and Pac. Sci. 25(3): maintenance of dredging. Water Res. 11(1): Kozloff, E.N Seashore life of Puget Sound, the Strait of Georgia, McClusky, D.S Some effects of and the San Juan Archipelago. Uni- salinity on the survival, moulting versity of Washington Press, and growth of Corophium volutator Seattle. 282 pp. (Amphipoda). J. Mar. Biol. Assoc. U.K. 48(2): Kozloff, E.N Keys to the marine invertebrates of Puget Sound, McClusky, D.S Salinity prethe San Juan Archipelago, and adja- ference in Corophium volutator. J. cent regions. University of Wash- Mar. Biol. Assoc. U.K. 50(3):747- ington Press, Seattle. 226 pp Levings, C.D., and 0. Levy A Nelson, W.G Reproductive "bugs-eye" view of fish predation. patterns of gammaridean amphipods. Pages in C.A. Simenstad and Sarsia 65: S.J. Lipovsky,eds. Fish food habits workshop, First Pacific Northwest Patton, W.K Feeding habits, Technical Workshop, Astoria, Oregon. behavior, and host specificity of Oct pp. Caprella grahami on amphipod commensal with the starfish Asterias for- Lorz, H., and W.G. Pearcy besi. Biol. Bull. (Woods Hole) Distribution of hyperiid amphipods 134(1):

24 Pinkster, S., and N.W. Broodbakker. mental impact of intertidal log The influence of environ- rafting on the Snohomish River mental factors on distribution and Delta. Wash. Coop. Fish. Res. Unit, reproductive success of Eulimno Seattle, Final Rep pp. gammarus obstusatus and other estuarine gammarids. Crustaceana Smith, J.L., and D.R. Mudd Suppl. No. 6: Maintenance dredging and the environment of Grays Harbor, Washington. Reish, D.J An ecological Appendix H: Impact of dredging on study of pollution in Los Angeles- the avian fauna in Grays Harbor. Long Beach Harbors, California. U.S. Army Corps of Engineers, Occas. Pap. Allan Hancock Found. No. Seattle District. 217 pp. 22: Stanhope, M.J., and C.D. Levings. Reish, D.J Studies on the Growth and production of Mylitus edulis community in Alamitos Eogammarus confervicolus (Amphipoda: Bay, California, II. Population Anisogammaridae) at a log storage variations and discussion of the site and in areas of undisturbed associated organisms. Veliger habitat within the Squamish Estuary, 6(3): British Columbia. Can. J. Fish. Aquat. Sci. 42(11): Reish, D.J., and J.L. Barnard Amphipods (Arthropoda: Crustacea: Amphipoda). Pages in C.W. Staude, Cd J.W. Armstrong, R.M. Hart, est~arine ed. invertebrates. Pollution ecology Academic of Thom, illustrated and K.K. key to Chew. the intertidal An Press, New York. gammaridean amphipods of Central Puget Sound. Coll. Fish. Univ. Wash. Seattle, Contrib. No Schmitt, W.L Crustaceans. pp. University of Michigan Press, Ann Arbor. 204 pp. Stepien, C.A., and R.C. Brusca Schulenberger, E Vertical Nocturnal attacks on nearshore distributions, diurnal migrations, fishes in southern California by and sampling problems of hyperiid crustacean zooplankton. Mar. Ecol. amphipods in the North Pacific Prog. Ser. 25(l): central gyre. Deep-sea Res. 25(7): Sutcliffe, D.W., T.R. Carrick, and L.G. Willoughby Effects of Sheader, M Breeding and mar- diet, body size, age and temperature supial development in laboratory- on growth rates in the amphipod maintained Parathemisto guadichoude Gammarus pulex. Freshwater Biol. (Amphipoda). J. Mar. Biol. Assn. 11(2): U.K. 57: Shyamasundari, K Studies on Tsuchiya, M., and M. Nishihira the tube-building amphipod (Coro- Islands of Mytilus as a habitat for phium triaenonyx Stebbing) from small intertidal animals: effect of Visaknapatnam Harbor: effect of island size on community structure. salinity and temperature. Biol. Mar. Ecol. Prog. Ser. 25(1): Bull. (Woods Hole) 144(3): Van Dolah, R.F., and E. Bird Smith, J.E A baseline study A comparison of reproductive patof invertebrates and of the environ- terns in infaunal and epifaunal 16

25 gammaridean amphipods. Estuarine Westernhagen, H.V., and H. Rosenthal. Coastal Mar. Sci. 11: Predator-prey relationship between Pacific herring, Clupea Waldichuk, M., and E.L. Bousfield. harengus pallasi, larvae and a pre Amphipods in low-oxygen datory hyperiid amphipod, Hyperomarine waters adjacent to a sulfite che medusarum. U.S. Natl. Mar. pulp mill. J. Fish. Res. Board Can. Fish Ser. Fish Bull, 74(3): (6): *17

26 REPORT DOCUMENTATION 1. RE NO Recipient' Accession No. PAGE Biological Report 82(11.92)* 4. Title and Subtitle S Repr Dote Species Profiles: Life Histories and Environmental Requirements January 1989 of Coastal Fishes and Invertebrates (Pacific Southwest)--Amphipods g 7. Author(s) S. Performing Organization Rept. No. Daniel J. Grosse and Gilbert B. Pauley 9. Performing Organization Name and Address 10. Proiect/Task/Work Unit No. Washington Cooperative Fishery Research Unit School of Fisheries University of Washington Seattle, WA Contract(C) or Grant(G) No. 12. Sponsoring Organization Name and Address (G) 13. Type of Repor A Penod Covered National Wetands Research Center U.S. Army Corps of Engineers Fish and Wi'dlife Service Waterways Experiment Station U.S. Department of the Interior P.O. Box Washington, DC Vicksburg, MS (c) IS. Supplementery Notes "U.S. Army Corps of Engineers Report No. TR EL Abstract (Umit: 200 words) Species profiles are literature summaries of the taxonomy, morphology, distribution, life history, and environmental requirements of coastal aquatic species. They are prepared to assist in environmental impact assessment. Amphipods are ubiquitous in distribution, but are most abundant in estuarine areas and other high nutrient areas. Hyperiidea are the third most abundant coastal marine crustacean zooplankton, following copepods and euphausiids. Benthic Gammaridea are an invaluable food source for many economically important fish and invertebrate species. Habitat preference and behavior of the major amphipod groups is highly variable. Intertidal California amphipods overlap the distribution of common genera of other regions around the world. Amphipoda are reported to be indicators of heavily polluted areas. They are considered the most efficient of all scavengers on the sea bottom and in shoreline areas. 17. Document Analysis a. Descriptors Amphipods Crustacean zooplankton Sediments Estuaries Harbor pollution Contaminants Fish prey Feeding habits Fisheries Food chains b. ldeitlfiers/ooen.cnd*d Terms Dissolved oxygen requirements Salinity requirements Gammaridea Temperature Ingolfiellidea c.cosati Field/Group Life history Hyperiidea Caprellidea Growth IS. Availability Statement T 12. Security Class (his Rep ) 21. Me. a Pa unclassified 17 Unlimited 0. Se,,curtyCashs Pge)m, Price U nclassified (9 ANSI-Z39 IS) OPTONPAL FORM 272 (4-M7 (Formerly NTI-.3S) Department of Comme"ce

27 As the Nation's principal conservation agency, the Department of 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 national parks and historical places, and providing for the enjoyment of life through outdoor recreation. The Department assesses 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 tinder U.S. administration. U.S. DEPARTMENT OF THE INTERIOR,, FISH AND WILDLIFE SERVICE j i"- TAKE PRIDE in America UNITED STATES DEPARTMENT OF THE INTERIOR POSTAGE AND FEES PAID FISH AND WILDLIFE SERVICE U.S. PARTMENT OF THE INTERIOR National Wetlands Research Center NASA-Slidell Computer Complex 1010 Gause Boulevard Slidell, LA IN1-4 OFFICIAL BUSINESS PENALTY FOR PRIVATE USE, $300