(DRAFT) - Taxonomy
                           Species mussel, California sea
                                 Species Id M060009
                                   Date 26 AUG 96



TAXONOMY

NAME - mussel, California sea OTHER COMMON NAMES - California sea mussel, rock mussel, big mussel, California mussel and sea mussel ELEMENT CODE - CATEGORY - Aquatic Molluscs PHYLUM AND SUBPHYLUM - , CLASS AND SUBCLASS - Bivalvia - clams or bivalves, ORDER AND SUBORDER - Mytiloida, FAMILY AND SUBFAMILY - Mytilidae, GENUS AND SUBGENUS - Mytilus, SPECIES AND SSP - californianus, SCIENTIFIC NAME - Mytilus californianus AUTHORITY - Conrad, 1837 TAXONOMY REFERENCES - 9 and 186 COMMENTS ON TAXONOMY - The name "California sea mussel" was given to M. californianus because of its restriction mostly to the outer coast *9*. Taxonomy - 1
                                  (DRAFT) - Status
                           Species mussel, California sea
                                 Species Id M060009
                                   Date 26 AUG 96



STATUS

Coded Status Sport Fish Commercial Commercial/consumption Commercial/bait See Comments REFERENCES FOR STATUS - 9 COMMENTS ON STATUS - The California sea mussel, along with the bay mussel, supports a small commercial bait fishery and is cultured for food on a small scale in Tomales Bay and just north of San Diego *9*. SPORT FISHERY A limited sport fishery, where mussels are usually removed from pilings or rocks by hand, now exists during the open season. A daily harvest of 25 pounds is allowed *9*. Status - 1
                               (DRAFT) - Distribution
                           Species mussel, California sea
                                 Species Id M060009
                                   Date 26 AUG 96



DISTRIBUTION

Distribution - 1
     

HABITAT ASSOCIATIONS

HABITAT - AQUATIC REFERENCES FOR HABITAT - 9 LAND USE - Water Bays and Estuaries REFERENCES FOR LAND USE - 9 NATIONAL WETLAND INVENTORY CODES NWI NWICLS NWIMOD NWISPEC Marine Marine, intertidal Estuarine Estuarine, intertidal Marine OW0 REFERENCES FOR NWI - 9 COMMENTS ON HABITAT ASSOCIATIONS - The California sea and bay mussels are relatively common along the rocky coastline and in bays and estuaries. The name "California sea mussel" was given to M. californianus because of its restriction mostly to the outer coast *9*. The eggs are fertilized in the open water. Larvae feed on phytoplankton and are at the mercy of currents. (They may be carried away from setting areas and die). M. californianus sets on barnacles, old mussel shell, and newly exposed hard surfaces. Density of newly settled M. californianus has been found to be highest on filamentous algae. They prefer to set on byssal threads from adults. It probably takes many years for Mytilus to become established in the high intertidal zone at approxiamtely +1.8 to +2.4 m above mean lower low water. Establishment is faster at lower levels. Mytilus californianus from southern California grew 100% faster in height and 50% faster in width at +0.1 m (the plus sign indicates vertical height above mean lower low water) than at +0.3 m where feeding time was shorter *9*. On Washington's exposed outer coast, M. californianus is the dominant competitor for available space and will form dense monospecific stands unless deterred. Mytilus californianus is one of the most consistent occupants of space in the intertidal barnacle-mussel association there. Habitat Associations - 1
                                (DRAFT) - Food Habits
                           Species mussel, California sea
                                 Species Id M060009
                                   Date 26 AUG 96



FOOD HABITS

TROPHIC LEVEL - SCAVENGER REFERENCES FOR TROPHIC LEVEL - 9 LIFESTAGE FOOD FOOD PART General Microorganisms Not Applicable General Protozoans Not Applicable General Bacillariophyceae Not Applicable General Detritus - Organic Not Applicable General See Comments; Food Not Applicable Larva Phytoplankton Not Applicable REFERENCES FOR GENERAL FOOD - 9 REFERENCES FOR LARVAE FOOD - 9 COMMENTS ON FOOD - Mussels are suspension feeders, are considered to be scavengers, and collect anything in the plankton that is small enough to ingest. Digestion is intracellular *9*. Mussels eat a variety of organisms, such as dinoflagellates, organic particles, small diatoms, zoospores, minute ova and spermatozoa, flagellates and other protozoans, various unicellular algae, and detritus (consisting of particles from the cytolysis of cells of a large variety of plants and animals). Growth rates are related to the abundance of dinoflagellates. Mytilus californianus from southern California grew 100% faster in height and 50% faster in width at +0.1 m (the plus sign here indicates vertical height above mean lower low water) than at +0.3 m where feeding time was shorter. Males grow faster than females the first 1.5 years *9*. FEEDING HABITS Food particles, drawn through an inhalent aperture by ciliary action, are caught on sheets of mucus and carried along the sides of palps to the mouth. Some particles are ingested, but others, if excessive, are discharged from the mantle cavity as pseudofeces. The most common food is the dinoflagellate Prorocentrum micans, at a size of 57 x 30 micrometers *9*. COMMENTS ON LARVAE FOOD - Larvae feed on phytoplankton *9*. Food Habits - 1
                         (DRAFT) - Environment Associations
                           Species mussel, California sea
                                 Species Id M060009
                                   Date 26 AUG 96



ENVIRONMENTAL ASSOCIATIONS

G = General A = Adult LIM = Limiting RA = Resting Adult J = Juvenile FA = Feeding Adult RJ = Resting Juvenile BA = Breeding Adult FJ = Feeding Juvenile P = Pupae L = Larvae E = Egg RL = Resting Larvae FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS G Water Temperature: Greater than 27 degrees C G Water Temperature: Between 21-27 degrees C G Water Temperature: Between 15-21 degrees C G Water Temperature: Below 15 degrees C G Water Temperature: Specified in Comments G Substrate: Specified in Comments G Relation to Substrate: Attached - normally sessile G Relation to Substrate: Specified in Comments G Water Level: Specified in Comments G Human Association: Specified in Comments LIM Currents: specified in comments LIM Water Level: Specified in Comments LIM J Relation to Substrate: Attached - normally sessile J Relation to Substrate: Specified in Comments L Currents: specified in comments L E REFERENCES FOR ENVIRONMENTAL ASSOC_ - 9 REFERENCES FOR LIMITING ENVIRONMENTAL ASSOC_ - 9 REFERENCES FOR JUVENILE ENVIRONMENTAL ASSOC_ - 9 REFERENCES FOR LARVAE ENVIRONMENTAL ASSOC_ - 9 REFERENCES FOR EGG ENVIRONMENTAL ASSOC_ - 9 COMMENTS ON ENVIRONMENTAL ASSOCIATIONS - HUMAN ASSOCIATIONS Man's manipulation of the coastal zone could provide additional habitats for mussels. Piers, floating marinas, jetties, and pilings for oil rigs are examples of surfaces ideal for settling mussels *9*. TEMPERATURE Temperature plays an important role in growth. In one study the growth of M. californianus was most rapid at 17-20 degrees C and decreased sharply above 20 degrees C. Growth continued (but at a lower rate) at 14 degrees C or lower but less rapidly. Feeding continued at 7-8 degrees C and 27-28 Environment Associations - 1 (DRAFT) - Environment Associations Species mussel, California sea Species Id M060009 Date 26 AUG 96 degrees C. Filtration rate and oxygen consumption were highest at 20 degrees C *9*. In southern California growth is fastest during the colder months and slowest (sometimes nil) in mid-summer or above 20 degrees C *9*. TIDAL INFLUENCE Mytilus californianus from southern California grew 100% faster in height and 50% faster in width at +0.1 m (the plus sign here indicates vertical height above mean lower low water) than at +0.3 m where feeding time was shorter *9*. SALINITY In nature, M. californianus is rarely found in bays and estuaries, perhaps because eggs, sperm, and larvae cannot withstand dilutions below 75% sea water *9*. COMMENTS ON LIMITING ENVIRONMENTAL ASSOC_ - CURRENTS Breeding success or failure is frequently determined during the critical larval stage. The larvae are at the mercy of currents and may be carried away from setting areas and die *9*. TIDAL INFLUENCE Mytilus californianus from southern California grew 100% faster in height and 50% faster in width at +0.1 m (the plus sign here indicates vertical height above mean lower low water) than at +0.3 m where feeding time was shorter *9*. SALINITY In nature, M. californianus is rarely found in bays and estuaries, perhaps because eggs, sperm, and larvae cannot withstand dilutions below 75% sea water *9*. WAVE ACTION While Mytilus edulis is sensitive to the effects of waves, M. californianus holds its position more strongly. Mytilus edulis is behaviorally adapted to quiet waters better than M. californianus. Mussels may form clumps up to 25 cm thick. Juvenile M. edulis crawl to the outside of such clumps, which may include M. californianus, but the latter mussel does not. This crawling behavior protects M. edulis from harmful silt that accumulates inside the clumps in quiet waters. This silt may tend to exclude M. californianus there *9*. Wave action or wave-borne logs in exposed areas may remove whole mats of mussels in the intertidal zone. At the edge of a cleared patch, substantial numbers of additional mussels may be removed by wave action. Mussels high in the intertidal zone away from starfish predation may survive for a great many years, until some are killed by wave-borne logs, unusual cold, or other physical stresses *9*. Environment Associations - 2 (DRAFT) - Environment Associations Species mussel, California sea Species Id M060009 Date 26 AUG 96 COMMENTS ON JUVENILE ENVIRONMENTAL ASSOC_ - Mytilus californianus sets on barnacles, old mussel shell, and newly exposed hard surfaces *9*. The cleaning of surface areas ("brushing the flats") has increased recruitment. Also, the placement of pilings, stakes, or poles is a common commercial method of collecting seed mussels. In the Santa Barbara Channel, natural sets of M. californianus are collected from the legs of oil drilling platforms. Setting areas are prepared by cleaning unwanted growth or wrapping areas with special materials that attract wild spat *9*. COMMENTS ON LARVAE ENVIRONMENTAL ASSOC_ - SALINITY In nature, M. californianus is rarely found in bays and estuaries, perhaps because eggs, sperm, and larvae cannot withstand dilutions below 75% sea water *9*. CURRENTS Breeding success or failure is frequently determined during the critical larval stage. The larvae are at the mercy of currents and may be carried away from setting areas and die *9*. COMMENTS ON EGG ENVIRONMENTAL ASSOC_ - SALINITY In nature, M. californianus is rarely found in bays and estuaries, perhaps because eggs, sperm, and larvae cannot withstand dilutions below 75% sea water *9*. Environment Associations - 3
                               (DRAFT) - Life History
                           Species mussel, California sea
                                 Species Id M060009
                                   Date 26 AUG 96



LIFE HISTORY

MORPHOLOGY/IDENTIFICATION AIDS The umbo is at extreme anterior end of the shell. Shell is covered with a heavy, black periostracum. Older specimens, from which this covering is worn off, are blue and have an eroded appearance. Exterior of shell with radiating ridges and concentric growth lines. Interior of shell bluish-black iridescence shows in reflected light. Length to 25 cm. Differs from the bay mussel in having up to a dozen broad radial ribs *9*. SPAWNING The California sea mussel continually spawns throughout the year at a very low level, with breeding peaks in July and December. In a study conducted in northern California, no definite spawning cycle was found. Ripe mussels were seen from February to May and from December to February. Spawning was observed in January, February, May, and December. In the natural environment, temperature does not seem to provide a major stimulus for spawning. During spawning the eggs and sperm are discharged through the excurrent chamber. The eggs, which appear as orange ribbons, are fertilized in the open water *9*. LARVAL STAGE The embryos develop into trochophore larvae (60-80 micrometers) about 24 h after fertilization. The veliger (straight-hinge stage) develops 24 h later. A ciliated velum forms and helps the larva swim and maintain itself in the water column. Larvae feed on phytoplankton and are about 0.17 mm long after 1 week. The veliger, which develops an umbo, may reach a length of 0.20 to 0.24 mm in 2 weeks. Just before metamorphosis, the larva develops an eyespot and foot *9*. The larvae of M. californianus have been reared in the laboratory to metamorphosis in 17-24 days after fertilization. Eyespots were observed when the larvae were 0.23 to 0.24 mm long *9*. Breeding success or failure is frequently determined during the critical larval stage. The larvae are at the mercy of currents and may be carried away from setting areas and die *9*. POSTLARVAE AND RECRUITMENT Mytilus californianus sets on barnacles, old mussel shell, and newly exposed hard surfaces. One study found density of newly settled sea mussels was highest on filamentous algae. Investigators disagree about settling behavior *9*. The cleaning of surface areas ("brushing the flats") has increased recruitment. Also, the placement of pilings, stakes, or poles is a common commercial method of collecting seed mussels. In the Santa Barbara Channel, natural sets of M. californianus are collected from the legs of oil drilling platforms. Setting areas are prepared by cleaning unwanted growth or wrapping areas with special materials that attract wild spat *9*. It probably takes many years for Mytilus to become established in the high intertidal zone at approximatedly +1.8 to +2.4 m above mean lower low water. Establishment is faster at lower levels *9*. MATURITY Sex ratios are generally equal and there is no indication of sex change with Life History - 1 (DRAFT) - Life History Species mussel, California sea Species Id M060009 Date 26 AUG 96 age. Some sexually mature mussels were only 4 months old or 25 mm long. Viable sperm was observed in mussels 30 mm long. Spawning occurred near the end of the first year or when the mussels were 70 mm long *9*. AGE AND GROWTH The age of mollusks is determined by three methods: (1) size-frequency studies; (2) the interpretation of growth interruption lines on the shell; and (3) experimental methods involving the release and recovery of marked individuals. One researcher reported that growth rates of mussels in any given population were so variable that year classes could not be determined from size. Most workers have attempted to age mussels through interpretation of growth interruption lines, although some have studied marked individuals *9*. In southern California, M. californianus sometimes grows as much as 7 mm per month. This species may reach a length of 80-86 mm within a year of settlement, 120 mm after 2 years, and 140-150 mm after 3 years. Maximum growth is 200-250 mm. Growth is fastest during the colder months and slowest (sometimes nil) in mid-summer or above 20 degrees C *9*. COMPETITORS When a resource is potentially limiting (as in the case of living space for sessile organisms in the rocky intertidal zone) one species of competitor may potentially dominate the others in procuring or holding the resource. Predation or physical disturbance may cause mortality of the dominant competitor sufficient to prevent the exclusion of the other competitors, allowing several species to coexist. Such phenomena occur in the rocky intertidal zone of the Pacific coast of the United States. The distribution of M. californianus is controlled by predation and physical disturbance *9*. On Washington's exposed outer coast, M. californianus is the dominant competitor for available space and will form dense monospecific stands unless deterred. Mytilus californianus is one of the most consistent occupants of space in the intertidal barnacle-mussel association there, along with the barncles Chthamalus dalli, Balanus glandula, and B. cariosa, and the sea anemone Anthopleura elegantissima *9*. The name "California sea mussel" was given to M. californianus because of its restriction mostly to the outer coast *9*. Mytilus californianus will settle among Balanus and completely cover the barnacles, eventually excluding them. Mytilus californianus denies space to M. edulis on the open coast, but it seems to offer protection from wave action to those M. edulis that do become attached *9*. PREDATORS A number of predators prey on mussels in California. Greater losses from predation have been found in shallow water compared to deep water. Mytilus californianus occupies only about 4% of the space (between about +3 m and a depth of about -27 M) that it would occupy under physically mild and predator-free conditions. The sea star P. ochraceus may clear patches of M. californianus, one of its preferred prey. In an experiment in Washington, P. ochraceus was removed and the area eventually became a monospecific stand of M. californianus. Pisaster ochraceus controls the abundance of the dominant mussel and thus allows other species to coexist. Mytilus californianus escapes predation from whelk predators (Thais spp.) by attaining a variable minimum size of invulnerability. This mussel escapes Life History - 2 (DRAFT) - Life History Species mussel, California sea Species Id M060009 Date 26 AUG 96 predation by P. ochraceus by growing to a minimum size that increases as the size of the sea star increases. Mean density of P. ochraceus may vary locally by at least two orders of magnitufde, affecting local abundance of M. californianus. In Washington, areas of low M. californianus abundance and high P. ochraceus density occur side-by-side with dense stands of large M. californianus that have low P. ochraceus density. The mussels had probably escaped predation by attaining an invulnerable size *9*. Pisaster ochraceus may indirectly enhance recruitment of M. californianus by the following series of events: 1)P. ochraceus consumes limpets (less preferred prey) and may reduce their abundance; 2) as shown experimentally, a reduction in limpet density would consequently enhance the abundance of one of the limpet's major foods, the red alga Endocladia; and 3) mussel recruitment may be enhanced because Endocladia is the major settling substrate for M. californianus larvae *9*. The abrupt lower limit of M. californianus band (about +0.5m) is almost certainly the most conspicuous feature of the rocky intertidal community and is probably set by predation by P. ochraceus *9*. The sea otter (Enhydra lutris) is known to eat both M. californianus and M. edulis, but mussels form only a small portion of its diet in California. Consumption of mussels by sea otters appears to be sufficiently scattered in time and space to preclude the regional depletion of harvestable stocks. The exposure of much of the central California coast to heavy surf probably provides an important refuge for mussels from foraging sea otters. Predation by the black oystercatcher, Haematopus bachmani, and dislodgement by storm waves are probably at least as important as sea otter predation in limiting the availability of mussels for human consumption in California *9*. Diving ducks, especially scoters, are mussel predators. Three species of ducks are responsible for most of the losses: the surf scoter, Melanitta delgandi, white-winged scoter, M. perspicillata, and black scoter, Oidemia amereicana. In feeding, the ducks dive underwater, remove a clump of mussels, return to the surface, shake the clump, and dislodge one mussel and swallow it whole. The other mussels in the clump sink to the bottom *9*. PARASITES AND ENCRUSTING EPIFAUNA In California the parasitic copepod Mytilicola orientales was found in 13 of 20 M. californianus. The gonads of one M. californianus, from Humboldt Bay, were heavily parasitized with trematode cercariae of the family Bucephalida *9*. Subtidal M. californianus may be overgrown by sponges, barnacles, anemones, and other epifauna. Heavily encrusted mussels weighted less (in tissue weight) than those less encrusted *9*. LIFE HISTORY CODES - Foraging Strategy: Scavenging Gestation/Incubation Period: 1-2 days Dispersion: Clumped REFERENCES FOR LIFE HISTORY- 9 Life History - 3
                           (DRAFT) - Management Practices
                           Species mussel, California sea
                                 Species Id M060009
                                   Date 26 AUG 96



MANAGEMENT PRACTICES

RESULT MANAGEMENT PRACTICE Beneficial Constructing/maintaining piers Beneficial Constructing/maintaining moring piles, dolphins, and bouys Beneficial Constructing/maintaining jetties, groins, and breakwaters Beneficial Controlling pollution [thermal, chemical, physical] Beneficial Other management practices [specified in comments] Existing Regulating harvest of species being described Existing Transplanting wild animals Existing Stocking captive-reared wild-strain animals Existing Regulating harvest - setting bag/creel limits Existing Mariculture activities Existing Other management practices [specified in comments] Adverse Other management practices [specified in comments] REFERENCES FOR BENEFICIAL MANAGEMENT PRACTICES - 9 REFERENCES FOR ADVERSE MANAGEMENT PRACTICES - 9 REFERENCES FOR EXISTING MANAGEMENT PRACTICES - 9 COMMENTS ON MANAGEMENT PRACTICES - THE FISHERY The California sea mussel, along with the bay mussel, supports a small commercial bait fishery and is cultured for food on a small scale in Tomales Bay and just north of San Diego *9*. Mussels are an important food item throughout the world. There is potential for culturing the California sea mussel for food more extensively in California. The cleaning of surface areas ("brushing the flats") has increased recruitment. Also, the placement of pilings, stakes, or poles is a common commercial method of collecting seed mussels. In the Santa Barbara Channel, natural sets of M. californianus are collected from the legs of oil drilling platforms. Setting areas are prepared by cleaning unwanted growth or wrapping areas with special materials that attract wild spat *9*. It probably takes many years for Mytilus to become established in the high intertidal zone at approximatedly +1.8 to +2.4 m above mean lower low water. Establishment is faster at lower levels *9*. PROBLEMS WITH THE FISHERY Problems associated with culturing the mussel for food are seasonal closures caused by pollution and paralytic shellfish poisoning (PSP), finding appropriate culture sites, and obtaining public acceptance to use of these sites *9*. No mussels can now be sold for human consumption from May 1 to October 31 because of the presence of paralytic shellfish poisoning *9*. The consumption of mussels that have concentrated large amounts of the poison-producing microscopic dinoflagellate Gonyaulax catenella sometimes causes serious illness. In 1980, for example, poisoning was recorded in 98 residents of Marin and Sonoma counties, California, of whom two died *9*. POLLUTION Management Practices - 1 (DRAFT) - Management Practices Species mussel, California sea Species Id M060009 Date 26 AUG 96 Because mussels are distributed along the coast, they are often subjected to sewage and other kinds of pollution. The mussels may be unsafe to eat and must be depurated before marketing *9*. Mussels concentrate hydrocarbons in their tissues. The hydrocarbons are rapidly taken up by the gill tissues and eventually are deposited in high concentrations in the alimentary canal. Although oil is only slightly toxic to mussels, it may affect the marketing of the animals by tainting them. One study reported that the uptake and loss of petroleum hydrocarbons was related to the magnitude of exposure. When mussels were placed in clean water, most hydrocarbons were lost, but signigicant quantities of No.2 fuel oil and outboard motor oil remained for as long as 35 days. Fuel oil and outboard motor oil may inhibit byssal thread formation, but at Long Beach Harbor, California, motor boat activity posed little threat *9*. A number of trace metals are found in the soft tissues of mussels, including AG, Cu, Cr, Mn, Pb, Cd, and Zn *9*. HUMAN ASSOCIATIONS Man's manipulation of the coastal zone could provide additional habitats for mussels. Piers, floating marinas, jetties, and pilings for oil rigs are examples of surfaces ideal for settling mussels *9*. SPORT FISHERY A limited sport fishery, where mussels are usually removed from pilings or rocks by hand, now exists during the open season. A daily harvest of 25 pounds is allowed *9*. AQUACULTURE Four companies have shown interest in farming mussels for human consumption in Tomales Bay,California, because the demand for a year-round supply of quality mussels has outgrown the supply from native beds. They hang ropes from longlines supported by floats to collect a natural set. The seed is then placed in plastic netting, which is then hung from the longlines. One company cultures mussels from the legs of oil drilling platforms in the Santa Barbara channels. Seed, placed in plastic netting, is wrapped around the legs. When the mussels reach maturity divers scrape them off the legs. The mussels are conveyed to the surface through suction hoses and are cleaned, packaged, and shipped fresh to market *9*. Management Practices - 2
                                   (DRAFT) - References
                              Species mussel, California sea
                                    Species Id M060009
                                      Date 26 AUG 96



     

References

9* Shaw, W., Hassler, T., Moran, D. 1988. Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (Pacific Southwest)--California sea mussel and bay mussel. U.S. Fish and Wildlife Service Biol. Rep. 82(11.84) pp 16. 186* Turgeon, D.D., A.E. Bogan, E.V. Coan, W.K. Emerson, W.G. Lyons, W.L. Pratt, C.F.E. Roper, A. Scheltema, F.G. Thompson, J.D. Williams. 1988. Common and scientific names of aquatic invertebrates from the United States and Canada: mollusks. American Fisheries Society Special Publication 16. (ed.):277. References - 1