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PACIFIC Ammodyfes hexapterus WITH NOTESON RELAPSEDAmmodytes SPECIES

L. Jay Fieldl

10ceanAssessment Division, National Oceanic and Atmospheric Administration, 7600 Sand Point Way NE, Seattle, WA 98115

1 BACKGROUND cies from the northwestern Pacific/ and the northwestern and northeastern Atlantic also will be cons~ 1. 1 Literature Search Two species of Ammodyres, AH and A. personarus AP!, havebeen reported from the north- An intensive literature search,except for Japaneseand westernPacific; AH generally is restricted to north of Soviet literature, which was not as thorough, was 45'N latitude Kitaguchi 1979!. The two speciesare completedin December1986, The emphasisof the distinguishedby meristic counts, with AH having the search was on the sand lance of the northeastern higher number of vertebrae and dorsal fin rays Pacific, including the Bering Sea. Lindberg 1937;Kitaguchi 1979!. Both speciesshow a greater number of vertebrae with increasing latitude Kitaguchi 1979!. Andriyashev954! consideredthem 1. 2 Unit Stocks and Their Relationships to be conspecific, and Hashimoto 984a! suggested that Ammodyres may be representedby three sub- The of Ammodyres species is uncertain. speciesalong the coastof Japan. The speciesare primarily distinguishedon the basisof Two other species of Ammodyres, a northern geographicdistribution and meristic counts. However, offshore species,A. dubius AD!, and a southernin- meristic counts frequently exhibit wide variation and shorespecies, A. americanus AA!, havebeen reported latitudinal clines Richards et al. 1963; Reay 1970; from the northwesternAtlantic Reay 1970!. However, Winters 1970; Scott 1972b; Richards 1982!. In this overlap in geographicrange and clines latitudinal and report,the generalclassification scheme proposed by inshore-offshore! in meristic counts that may be Reay 970! will be followed. Ammodyreshexaprerus correlatedwith environmentalconditions suggeststhe AH! is the only sand lance reported from the possibility that only a singlespecies occurs in this area northeasternPacific Hart 1973!. They are found in Richards et al. 1963; Winters 1970; Scott 1972b; shallow nearshore waters froin to the Richards 1982!. Two species,A. robianus AT! andA. Iteaufort Sea Trumble 1973; Craig 1984!. As detailed marinus AM}, are found in the northeastern Atlantic, taxonomic studies have not been carried out, the but only AM will be considered in this report, as AT relationship of AH to sand lance from other areashas does not resembleclosely the speciesfrom other areas not been determined. Consequently,Ammodyres spe- Reay 1970!. 16 / Species Synopses

1.3 The Fishery 1.5 Current Status of Stocks 1.3.1 Relative Size and Importance. Except for and of Management Measures occasional small bait fisheries in and No information is available on the status of stocks of , no fishery currently exists in the AH in the northeasternPacific, since no fishery northeasternPacific, although there is some potential currently exists and AH are not sampled by conven- Trurnble 1973!. In Japan the fishery takes about tional research trawls. 100,000ton pcr year Kitaguchi1977!. No fisheryfor sandlance exists in thenorthwestern Atlantic, although 1. 6 Recruitment Variability feasibility studiesindicate considerable potential R. Smith, Univ. New Hampshire, pers. comm.!. The Evidencefor AH recruitmentvariability is mainly European fishery landed 200,000 mt in 1968 Reay circumstantial. For example, the presenceor absence 1970!, of sandlance in seabirdstomachs has been noted during 1.3.2 Age at Recruitment. In Japan, sand lance trophic studies Vermeer 1979; Drury et al. 1981; AP! usuallyenter the fisheryat age 1, but 0-agefish Springer et al. 1984!. In Japan,there is considerable may enterlate in the season,particularly if the year variability in the percentageof 0-agefish in AH/AP class is strong Kitaguchi 1977!. In the North Sea, catches Hamada 1966a,b,c; Inoue et al. 1967; recruitmentto the AM fishery inay begin at agc 6 Kitaguchi 1977;Nagoshi and Sano 1979!, monthsbut most enter at age 1 Macer 1966;Reay Larval surveys in the northwestern Atlantic indicate 1970!. large interannualvariation in sand lance AA/AD! 1.3.3 Characteristicsof HarvestedFish. Lengthof recruitment. Between 1974 and 1979, abundance of AP/AH in the Japanesefishery rangesfrom 8 to 26 cm larval sandlance increasedby a factor of 20 Smith et Kitaguchi 1977!, In the northeasternAtlantic fishery, al. 1978;Sherman et al. 1981!, which reflects a 50-fold AM rangefrom 5-25 cm in length Macer 1966!. changein adultspawning biomass Meyer et al. 1979; 1.3.4 Typesand Selectivityof Gear, In Japan,a Morse 1982!. variety of seines,lift nets and bottom trawls are used for the sandlance fishery Inoue ct al. 1967!. Thc 1.7 Age Determination and Validation Europeanfishery primarily uses a highopening bottom trawl with a 6 mm cod-endmesh Macer 1966; Macer Age determination for sand lance speciesis basedon andBurd 1970!.The selection length for thisgear was surface readings of whole otoliths Kitakata 1957; estimatedto beapproximately 8-9 cm Reay1970!. Macer 1966; Scott 1968,1973;Reay 1970, 1972, 1.3.5 Distributionof FishingEffort, TheJapanese Kitaguchi1977; Winters 1981!. Validation of ageshas sandlance fishery takesplace in depthsof 80 m or less, beenattempted by observingthe annual pattern of ring in areasof sandand shell bottom Kitaguchi1977!. In formation Kitakata 1957; Reay 1972; Kitaguchi southernareas, the AP fishery runs from mid-March to 1977!. lateJune Inoueet al. 1967!,but at higherlautudes the seasonis somewhatlater, sometimesextending from 1.8 Age Composition of April to December Kitaguchi 1977!. the Population The EuropeanAM fisheryis conductedin depthsof In sandlance fisheries, age composition is apparently lessthan 40 m in sand-bottomareas, primarily on off- subjectto variationwith time of year,area and gear in shorebanks Macer1966!. The fishing season extends additionto interannualdifferences Inoue et al. 1967!. from April to September,with very low catchesat AH collectedby beachseine from the vicinity of other times of the year Macer 1966;Reay 1970!. KodiakIsland, , ranged from age0 to 5, with Fishingtakes place almost exclusively during daylight age 1 fish mostabundant Dick and Warner 1982!. hours Macer 1966!. However, differencesin age composition betweenAH collectedby beachseine and those dug from gravel 1.4 Distribution and Abundance beachesindicate that beachseines were not effective in of the Population samplingolder age classes Dick andWarner 1982!. In northernareas of Japanwhere both AP and AH Sand lance are abundant in shallow ncarshore areas. occur,most of thecatch is composedof ages1-3, and fish overage 4 areuncommon; there is someevidence Pacific Sand Lance / 17

of interannual variability Kitaguchi 1977!. In con- 2.1.3 Fecundity. Fecundity-lengthrelationships F trast, in the Seto Inland Sea fishery, 0-age fish com- = number of , L = length in cm! have been deter- prise about 80% of the catch and age-2 fish less than mined as follows: 5% Inoue et al. 1967!, whereas in the Harima-Nada and OsakaBay fishery, age-1 AP accountfor 20-77% AA: F = 0.328 L = 3.857 Westin et al. 1979!; of the catch Hamada1966a!. AM: F = 2.046 L = 3.055 Macer 1966!, Macer 966! reported that the percentageof older Sandhnce probably AM! between 120 and 195 mm fish AM! in the catch decreasedduring the last part of standard length! off the Murrnan coast had between the season, and Winslade 974c! suggested this was 3300 and 22,I00 eggs average 6800! per female due to age related differences in timing of over- Andriyashev 1954!. wintering see 2.2.5!. The maximum age of AD in Newfoundlandis age 10 or over Winters 1983!. Ages 2. 2 Distribution and Abundance 3-5 predominatein the survey catches,but the relative abundanceof AD older than age 6 hasbeen increasing 2.2.1 Duration of Spawning. Most information for since the late 1970s Winters 1983!. In the the time of spawning of sand lance is based on the oc- northeasternAtlantic, the maximum age of AM is 9, currenceof early larvae. In ,Washington, but ages 1-3 account for the majority of the catch yolk-sacAH larvaeare mostabundant in late January Reay 1970!. to early March R. Trumble, Washington Dep. Fisheries,pers, comm.! and spawningoccurred in outdoor holding tanks in mid-March Pinto 1984!. Off 2 ADULTS the west coastof VancouverIsland, larval AH beganto occur in February Mason et al, 198la, b!. Larval 2.1 General Description surveys in the Kodiak region indicated late winter February-March! spawning Rogers et al. 1979; 2.1.1 Size and Age Ranges. Based on a small Kendall et al. 1980!. However, Dick and Warner samplesize n=7! of mature fish, AH in the vicinity of 982! reportedintertidal spawning off Kodiak!slandin Kodiak Island, Alaska, matureat ages2-3; the smallest October.Spawning of sandlance AH/AP!in theSoya mature sand lance observed was 128 mm Dick and Strait region of Japan occurred between January and Warner 1982!. In the Soya Strait region of Japan, early May, and there was some evidence of interannual most sand lance AH/AP! spawn at the end of their variation Ki taguchi 1977!. secondyear but some spawnat age 1 Kitaguchi 1979!. 2.2.2 Large-scalePatterns. In general,sand lance Most AP from Ise Bay to the south mature at ages 1-2 occurin shallownearshore areas, usually in depthsless at lengths greater than 110 mm Nagoshi and Sano than100 m, with sandor sand-gravelsubstrates. They 1979!. are abundant from spring to late summer and In the North Atlantic, AA usually are mature 02- uncommonduring the remainder of theyear when they 114 m! at the end of their first year Richards 1982!. presumably are buried in the sand Leim and Scott Scott I968! reportedthat mostAD matureat age2 at 1966; Reay 1970; Trumble 1973!. While inshore- lengths between 180 and 200 mm, whereasWinters offshoremovements have been reported by a numberof 983! found that 50% were matureat age 3 at 180mm authors,there is no evidenceof large-scalemigrations in length. Most AM are matureat age2, but large of any sandlance species Reay 1970!. AH juveniles 00 mm! individualsmay matureat age 1 Macer and adultsare commonly abundantin nearshorewaters 1966!. during spring-summerbut are rarely sampledduring 2.1.2 Typeof SpawningBehavior. Most species of late fall and winter Barton 1978; Fresh 1979; sandlance spawnonce a year Reay 1970!, Unimodal Blackburnand Jackson 1980; Blackbuin et aI. 1980!. distributions of diameters in ovaries of mature AH Accordingto Blackburnand Jackson980!, AH are Pinto 1984!,AD Scott1972a! and AM Macer1966! inactive during the winter and are often found buried in indicatea singlespawning per year, AA thatspawned intertidal sand. In the Bering Sea,they have been in laboratory holding tanks had very few eggs reportedfrom 0- IOOm butare most common in depths remainingin their ovaries,indicating that aIl the eggs lessthan 50 m Shuntov,in Macyet a!. 1978!. to be spawnedin a yearare released in a singlebatch 2.2.3 Small-scalePatterns. All speciesof sand Smigielski et al. 1984!. lanceexhibit schooling behavior, often forming large 18 / Species Synopses

aggregations Reay 1970; Trumble 1973!. Meyer et al. any factors affecting their abundanceand distribution 979! observed schools of AA ranging in number presumablycould affect sandlance feeding. Calanoid from approximatelya hundredto tensof thousands;the copepods also are preyed on to a great extent by largestfish werein the centersurrounded by smaller juvenilePaciTic herring that frequentlyco-occur with fish. sandlance Harris and Hartt 1977a,b; Simenstadet al. 2.2.4 Maximum and Mean Abundances. No 1979!. information on abundance of AH is available. An 2.3.5 Temporal Patternsof Energy Storage. Sea- estimateof spawningbiomass of sandlance AA/AD! sonal variation in fat content has been demonstrated for in the northwestern Atlantic from larval survey data AP Inoue et al. 1967; Sekiguchi et al, 1976! and AM indicated that biomass increased 50-fold between 1974 Kuhl and Luhinan, in Winslade 1974c!, Lowest and 1978 Morse 1982!. values were recorded at the beginning of the feeding 2.2,5 Vertical Distribution. Seasonal and diurnal period and peak fat content is reachedat the end of the variations in the catch of sand lance are thought to be, feeding seasonjust prior ta a prolongedperiod of dor- at least partially, the result of their habit of burrowing mancyduring which the sandlance remain buried in the in the sand when not feeding. On the basis of sand, Fat supply is probably important for main- underwater observations of AH, Hobson 986! tenancemetabolism and gonad maturation Winslade reportedthat samesand lance were buried in the sandat 1974c;Sekiguchi et al. 1976!. all times of the day, but most were feedingin the upper 2,3,6 Evidence of Food Limitation. No data avail- levelsof the water column during the day and buried in able. the sand at night. Trawl catchesof AH were much reducedat night Macer 1966!. 2.4 Predation

2.3 Feeding 2.4.1 Predator Species. A wide variety of marine fish, seabirdsand mammalsare major predatorsof AH 2.3.1 . Sand lance AH! feed primarily in over their entire range Tables 1-3!. the water column, although epibenthic invertebrates 2.4.2 Effect on Spawning Adults. No estimates of occasionallyappear in their diet Simenstad et al. 1979; the importanceof predation have been made for AH, Rogers et al. 1979!. although apparently it is very high, Evidence for spe- 2.3.2 Prey Species. Calanoid capepods are the cies from the northwestern Atlantic Sherman et al. major prey for AH adults andjuveniles from the west- 1981; Winters 1983! and the North Sea Andersen and ern Aleutians Simensradet al. 1978!, Kadiak Harris Ursin 1978! suggestsdecreased levels of predationan and Hartt 1977a,b; Rogerset al. 1979! and Washing- sand lance have led to large increases in sand lance ton Simenstadet al. 1977,1979;Crass et al. 1978!. populations. Other prey are crustaceanlarvae, rnysids, gammarid amphipods,harpacticoid copepods,chaetognaths, lar- 2.5 Other Factors Affecting Adults vaceansand polychaetes. According to Rogerset al. 979!, epibenthic invertebrates,particularly mysids 2.5.1 Biotic. Mass mortalities of sand lance along and gammaridamphipods, inc~ in relative import- the coastof England has been associatedwith blooms ancefor AH during autumn and winter; however,total of Gonyardax sp Adams, in Reay 1970!. See 2.5.2 stomach content weight was much lower. AA made a for laboratory experiments related to food availabihty.! similar shift to benthic prey during early spring when 2.5.2 Abiotic. As a consequence of their habit of zooplanktonabundance was low Richards 1982!. The burrowing into sandy bottoms for extended periods, prey spectra of other speciesof sand lance are very sand lance usually are found in associationwith sand similar ro that of AH see Inoue et al. 1967; Sekiguchi and fine gravel bottoms Reay 1970; Trumble 1973!. et al. 1974,1976; Kiraguchi 1977; and Hashimoto Both particle size and circulation appear ro be import- 1982 for AP; Richards 1965, 1982; and Meyer et al. ant factors, although specific habitat requirementsare 1979 for AA; Scott 1973 for AD and Macer 1966 for unknown. Laboratory experiments showed thar.AH AM!. avoided oiled sand Pearson et al. 1984; Pinto et al. 2.3.3 Prey Density Requirements.Not applicable. 1984!. AH mortality was noted in connectionwith oil 2.3.4 FactorsAffecting Availability of Prey. Since and detergentcleanup in the Torrey Canyon calanoid capepodsare the primary prey of sandlance, Smith, in Reay 1970!. Pacijic Sand Lance / I9

Table 1. Marine fish predatorsof sandlance AH!.

PE IE

PaciTic cod Aleutians Simenstad et al. 1977 Kodiak Jewett 1978; Hunter 1979 British Columbia Westrheimand Harling 1983

Halibut S.E. Bering Sea Smith et al. 1978 W. Aleutians Simenstad et al. 1977 Kodiak Hunter 1979 Gulf of ~ Best and Hardman 1982 Bering Sea

British Columbia Outram and Haegele1972

Rock sole IGxRk Harris and Hartt 1977; Hunter 1979 British Columbia Westrheimand Harling 1983

Yeilowfin sole Harris and Hartt 1977;Rogers et al. 1979

Sand sole Hunter 1979

Petrale sole British Columbia Westrheimand Harling 1983

Lingcod British Columbia Hart 1973

Rockfish black, Gulf of Alaska Rosenthalet al. 1981, 1982, yellowtail, dusky, Rosenthal 1983 widow!

Salmon chum W. Aleutians Simenstad et al. 1977 sockeye!

Salmon chinook, British Columbia Beacham 1986 coho, sockeye,pink!

Greatsculpin Harris and Hart 1977 20 / Species Synopses

Table 2. Marine seabirdpredators of sandlance AH!.

Black-legged Chukchi Sea Swartz 1966; Springer and kittiwake Roseneau1979; Springeret al. 1984 Norton Sound Drury et, al. 1981 S. Bering Sea%ulf Hatch et al. 1978 of Alaska

Common murre Chukchi Sea Swartz 1966; Springer et al. 1984 Norton Sound Drury et al, 1981 E. Bering Sea Ogi and Tsnjita 1973 Kodiak Drury et al. 1981

Thick-billed murre Chukchi Sea Swartz 1966;Drury et al. 1981; Springeret al. 1984

Tuftedpuffin Chukchi Sea Swartz 1966 W. Aleutians Vermeer 1979 Gulf of Alaska British Columbia Washington Wilson 1977; Wilson et al. 1984

Chukchi Sea Swartz 1966 Norton Sound Drury et al. 1981 Pribilof Islands Hunt et al. 19&1 Gulf of Alaska Manual and Boersma 1978 N. Gulf/S. Bering Hatch et al, 1978

Rhinoceros auklet Gulf of Alaska Leschner in Vermeer 1979 British Columbia Vermeer 1979 Washington Wilson l977; Wilson and Manuwal 19&6

Ancient murrelet British Columbia Scaly 1975

Sooty shearwater British Columbia Scaly 1975

Pelagiccormorant Chukchi Sea Swartz 1966 British Columbia Robertson 1974

Double-crested British Columbia Robertson 1974 cormorant

Red-throated loon British Columbia Reimchenand Douglas 1984 Pacific Sand Lance / 21

Table 3. Marine mammal predators of sand lance AH!.

Fur seal E. Aleutians Taylor et al., in Sirnenstad et al. 1979 W. Gulf of Alaska Wilke and Kenyon 1957;Kajimura 1984 Kodiak Macy et aL 1978

Harbor seal Gulf of ~ Pitcher 1980 Brown and Mate 1983

Stellar sea lion Gulf of Alaska Wilke and Kenyon 1952;Pitcher 1981

Spotted seal E. Bering Sea Lowry and Frost 1981

Minke whale N. Pacific Nemoto 1959;Frost and Lowry 1981;Kajimura 1984

Sei whale Bering Sea Frost and Lowry 1981

Humpbackwhale Huey, in S imenstadet al. 1979 22 / Species Synopses

Temperature and light also appear to be important 2.6 Laboratory Holding and Rearing factors. When water temperatures reach 20'C, AP bur- Spawningof adult sandlance in the laboratoryhas been row in the sand and become dormant Nagoshi and reported for AH by Pinto 984!, for AA by Sano 1979!. For adult AD, however, increased tem- Smigielski et al. 984! and for AM by Winslade perature is associated with growth Winters 1983!. 971, 1974a,b,c!. White Sea sand lance AM! are reported to bury themselvesin responseto reduced light levels <0.1 Iux! and picdators Girsa and Danilov 1978!. 3 EGGS In laboratory experiments, the activity of adult AM 3.1 General Description was directly related to food availability, light intensity, The sand lance egg, which usualIy contains a singIe and temperature Winslade 974a,b,c!. If food was yellow oil globule, is nearly spherical, demersal and present, activity was high during the day and low at adhesive Reay 1970; Trurnble 1973!. According to night when they mostly remained buried in the sand Pinto 984!, AH eggs from the northeasternPacific Winslade 1974a!. When abundant food was available, range from 0.8-1.22 mm, with a mean diameter of 1.00 swimming activity was maximum at light intensities min. Mean diameters of egg of other Amiriodyres of 100 and 1000 lux, much reduced at 10 lux and very speciesare 0.66 mm for AP, 0.83 mm or 1.00 rnm for low at I Iux; the thresholdlevel light intensity at 50% AA, 1.00 mm for AD and 1.02 mm for AM Williams maximum activity! was 20 lux Winslade 1974b!. et al. 1964; Inoue et al. 1967; Winslade 1971; Scott Diurnal and seasonal variations in catch of sand lance 1972a;and Smigielski et al. 1984!. on the fishing groundscorrelated with estimatedlight intensity of about 100 lux at the bottom Winslade 3. 2 Distribution and Abundance 1974b!. Activity during daylight was high at 10 and 3.2.1 Duration of Egg Stage. In the laboratory, 15'C and much lower at 5'C WinsIade 1974c!. The incubation time for AH eggs was approximately 24 percentage of the annual catch in April the first month days at 9'C Pinto 1984!. Japanesesand lance AP! of the fishery! showed a positive correlation with eggshatched in 33 days at 6.2 C and 13 daysat 15.7'C temperature in April, indicating that initial availability lnoue et al. 1967! and AP or AH eggs hatched in in the spring may be more related to temperaturethan about 22 days at 6.9' to 10'C Kitaguchi 1977!. to food availability Winslade 1974c!. Burying in the According to Yamashita and Aoyama 985!, median sand at the end of the summer may be related to fat hatching time for AP eggs was 51 days at 6.5'C, 25 days at 10.5 C and 20 days at 15.5'C. Average content, as well as to decreasing levels of food hatching time for AA eggs was 62 days at 2'C and 30 availability, light intensity and temperature Winslade days at 10'C Smigielski et al. 1984!. Experimentson 1974c!. Older fish are reported to disappear from the AP eggs Yamashita and Aoyama 1985!, AA eggs catches earlier in the summer, possibly due to reduced Smigielski et al. 1984! and AM eggs Winslade 1971! requirementsfor growth and/orearlier accumuIationof showed that incubation times were long and variable fat Winslade 1974c!. and that temperature and oxygen were important 2,5,3 Total Mortality. For a fished population of factors. adult AM, total annual mortality was estimated at 65- 3.2.2 Large-ScalePatterns. No information on egg 75% Z=l.2! Macer 1966, cited in Reay 1970!. Based distribution is available,but presumablyeggs would be on age composiuon data from other stocks, Reay found in the sainegeneral areas as adults sincespawn- 970! suggested fished and unfished stocks have simi- ing migrationsare not known Reay 1970!. lar mortality rates. Winters 983!, who investigated 3.2.3 Small-scale Patlerns. Nodataavailable. AD mortality ratesin Newfoundlandbetween 1968 and 3.2.4 Maximum and Mean Abundances. No data 1979,reported a steadydecrease in mortality ratesfrom available. Z-values of over 1.0 to less than 0.5. Since there is no 3.2.5 Vertical Distribution. Sand lance eggs are fishery for sand lance in Newfoundland, these rates are demersaland adhesive,although they are occasionally consideredto be naturalinortality, The increasein sand collected from the water column, probably as a result lance survival was correlated with a decline in cod of currentsbringing the eggsoff the bottom Williams stocks,known to be major predatorson AD. et al 1964; Senta 1965!. Paciji c SandLance I 23

3.3 Feeding AD!; Einarsson 1951, Macer 1966, Winslade 1971 AM!. Not applicable. AH larvae hatched at 9 C were 5.3 mm in length, had pigmentedeyes but did not have a completegut or 3.4 Predation functional jaws Pinto 1984!. At the end of one week, 3,4,1 Predator Species. Large numbers of late stage the yolk sac was not completely absorbed and the AD eggs were found in stomachs of the yellowtail intestinesand jaws were still not completelydeveloped. flounder Limanda ferruginea! in the northwestern AP larvae hatched at 6.5'C were 4.7 mm in length Atlantic Scott 1972a!. Yamashita and Aoyama 1985!. Some larvae began 3.4.2 Effect on Eggs. No data available. feeding within 2 days, and 50% of the larvae were feed- ing successfully within 5 days, Yolk-sac absorption 3.5 Other Factors Affecting Eggs was 95% complete by 6 days and 100% complete by 12 days after hatching. Larvae were able to survive 3,5.1 Biotic, No data available. without food for a long time after hatching; time to 3.5.2 Abiotic. According to Inoue et al. 967!, AP 50% mortality was approximately 11, 16 and 21 days had a low hatch rate at 15.7'C and a maximum hatch after hatching at 15.5', 10.5' and 6.5'C, respectively. rate at 8.2'C, !n a laboratorystudy on AA, incubation No clear "point of no return" was noted. The period of time and time to hatch completion increased with recoverable starvation from normal onset of feeding to decreasing temperatures 0'-2'C! Smigielski et al. irreversiblestarvation! was estimated as about9 days. 1984!, The authors suggest that in the natural envi- AA larvae hatched at four different temperature ronment mechanical action may be a factor in reducing regimes, 4, 7 and 10'C! and had a completegut and incubation times. Experiments on AM eggs indicated functional mouth; some individuals in each of the four hatching time may be affected by factors other than groups began to feed within hours of hatching temperature, e.g., incubation time and mortality Smigielski et al. 1984!. At 7 C, 50% of larvae 6.5 increased with decreasing oxygen concentrations 9.5- mm in length were feeding within 2 days. Yolk-sac 4.0 ppm! Winslade 1971!. No eggs hatched at 2.1 absorption took 3 days and oil globule absorption 7 ppm; however, early stage embryos were able to days. For AM larvae, yolk-sac and oil globule tolerate low oxygen concentrations for about a week absorptiontook about 2 weeksat 7'C and slightly less and to complete development if subsequently shifted to time at 10'C Winslade 1971!, After 13 days, 16% of higher concentrations.Winslade 971! suggestedthat larvae at 7'C and 58% at 10 C had begun to feed. the ability to retard developmentand surviveperiods of However, newly hatched larvae were able to survive for low oxygen levels may be of particular adaptive value, a considerable length of time without food: time to since sand lance spawn on sandy bottoms and eggs 50% mortality was 28 days at 7'C and 19 daysat 10'C, could become temporarily buried in shifting sand. Yamashita and Aoyarna 985! determinedgrowth 3.5.3 Total Mortality. No data available. rates of 0,12 mm in length and 4.2% dry weight per day for AP rearedin the laboratory at 6.5'C. Growth 3.6 Laboratory Holding and Rearing rates for the two northwestern Aoantic sand lance have Techniques for incubation of sand lance eggs are been estimatedfrom length frequencydistributions as described by Winslade 971!, Pinto 984! and follows: AA, 11.7 mm per month Norcross et al. Smigiclski et al. 984!. 1961! or 10.9 mm per month Smith et al. 1978!; AD, 5.9 mm per month Scott 1972a!. Smigielski et al, 984! estimatedgrowth rates for AA to range from 4 LARVAE 2,4-5.62% dry weight per day at 2'C and 10'C, re- spectively, corresponding to length increasesof 2.7 4.1 General Description mm and 11,3 mm per month for 155 days of growth. Larvae of the five species of Ammodytes considered Buckley 984! estimated minimum growth rates to herein have been described by the following: Pinto prevent starvation in AA larvae to be 2.4%, 2.5% and 1984 AH!; Kobayashi 1961 AP!; Richards 1965 3.4%change in proteincontent per day at 2, 4 and7'C, AA/AD!; Smigielski et al. 1984 AA!; Scott 1972a respectively. 24 / Species Synopses

4. 2 Distribution and Abundance densities in Otsuchi Bay, Japan,averaged 105.6/100 4.2.1 Duration of Larval Stages. AA laboratory m3 in 1981-82 Yamashita et al. 1984b!. studies show that at 7 C yolk-sac absorption takes In the Gearges Bank region, concentrations of sand place in 3 days at about 6.5 mm in length, oil globule lancelarvae AA/AD! wereover 100,000/10m2 absorption in 7 days at 7.2 mm and 50% first feeding Smith et al. 1978!. Mean abundance in this region in 2 days at 6.5 mm Smigielski et al, 1984!, With rangedfrom ]0 to 1018pcr 10m2 between 1974 and increasing temperature, size at yolk-sac absorption 1979 and accounted for 55-98% of the larvae in winter decreased but no differences in size at ail globule samples Sherman et al. 1981!. absorption or at first feeding were observed. Metamor- 4.2.5 Vertical Distribution. In Kodiak Island phosis to the juveniLe stage occurred at 29 mm in samples,AH larvae were most abundantat 10 and 30 length, which took 131 days at 4'C and 102 days at m depthsduring the day and were somewhatdeeper at 7'C. Schooling behavior was abserved at 25-30 mm night Rogers et al. 1979!. Inoue et al. 967! and 90 days at7 C!. Yamashita et al. 985! reported similar results for AP Laboratory-reared AM larvae completely absorbed larvae; they were 6-10 m below the surfaceduring the yolk sacsand oil globules in 14-15 days at 7'C and in day and deeper at night and also were more concentrated 12 days at 10'C WinsIade 1971!. AM larvae above thermoclines. Yamashita et al. 985! reported completed metamorphosisat lengths between 30 mm vertical migratian in larval AP as small as 5-6 mm, and 40 mm Macer 1966!. although larger larvae had a greater vertical range. AA 4.2.2 Large-scalePatterns, Sand lance larvae AH! larvae also were found to be near the surface during the generallyoccur in shallow less than 200 m! nearshore day anddeeper at night Richardsand Kendall 1973!. areas of the Bering Sea and northeastern Pacific Ocean Trumble 1973; Macy et al. 1978; Rogers et al. 1980!. 4.3 Feeding AH larvae were most abundant in early April in the Bering Sea Waldmn and Vinter 1978! and March-April 4.3.1 Habitat, See 4.2.5.! in the Kodiak area Rogers et al. 1979; Kendall et al. 4.3.2 Prey Species. In the Strait of Georgia under 1980!. In Washington,small larvae less than 10 mm! the Fraser River plume!, larval sand lance AH! less were reported from late January to early May in Skagit than 20 mm in length fed on prey less than 500 Ii in Bay Blackburn 1973! and from mid-January to late diameter,mainly copepodeggs and nauplii LeBrasseur March in Puget Sound R, Trumble, Washington Dep. et al. 1969!. Larger larvae 0-40 mm! fed on larger Fisheries,unpublished data!, Recently,hatched larvae .5-1.5 mm! zooplankton, primarily species of AA/AD! that were most abundant in shallow nearshore Microcalanus, Oithona and Pseudocalanusand nauplii areasgradually dispersedoffshore in the northwestern of larger copepods. AP larvae fed mostly on small Atlantic Richardsand Kendall 1973!. copepods, copepod nauplii and cladocerans Inoue et al. No information is available on interannual varia- 1967; Sekiguchiet al. 1974!. bility in AH larval abundance. However, sand lance According to Covill 959!, the primary prey items larvae AA/AD! have shown very large interannual for larvalAA between3,2 and23.1 mm in lengthwerc changesin abundance approximately a 20-foldincrease copepod nauplii and copepods. Phytoplankton, an between 1974 and 1979! Smith et al. 1978; Sherman important component of the diet for small larvae less et al. 1981; Morse 1982!. than 5 mm!, decreasedin importance with increasing 4.2.3 Small-scale Patterns. Altukhov 978! re- size of larvae, Phytaplankton appearedto be mare ported that AM larvae in the White Sea werc concen- important in winter Dec-Feb.! than in spring March- tratedin areasof "eddyingcurrents." April! for the samesize class of larvae. Ryland 964! 4.2.4 Maximum and Mean Abundances. In the noted that AM larvae fed only during daylight. eastern Bering Sea, sand lance AH! larvae were the Diatoms and dinaflagcllates were important items in third most abundant species in bongo net samples the diet of AM larvae less than 8 mm in length, and 908/10 m ! despitetheir occurrencein less than 25% larger larvae fed mostly on copepad nauplii and of the samples Waldran and Vinter 1978!. AH larvae appendicularians. were also abundantin ichthyoplankton samplesfrom 4.3.3 PreyDensity Requirements. Laboratory stud- the vicinity of KadiakIsland Dunnand Naplin 1974; ies of Winslade 971!, Buckley et al. 984! and Rogers et al. 1979; Kenda11et al. 1980!. Larval AP Yamashitaand Aoyama986! showedthat newly Pacific Sand Lance / 25

hatched AM, AA and AP larvae are capable of surviv- major predator of larval AH in Alaska Bent, cited in ing up ta 2 weeks without food see also 4.1!. Ainley and Sanger1979!. Seealso Tables 1-3.! 4.3.4 Factors Affecting Availability of Prey. No Predationby the hyperiid amphipod Pararhemisro data available, japonica on larval AP Yamashita et al. 1984a, b! was 4.3.5 Temporal Patterns of Energy Storage. Nat estimatedta be an important source of morality. In applicable, the laboratory, both newly hatched and post-yolk sac 4.3,6 Evidence of Food Limitation. LeBrasseur et hrvae 3-15 days old; 6.0-6.3 mm standardlength! al. 969! reported that over 50% of AH larvae less were vulnerableto amphipad predatian, than 40 mm in length in the Strait of Georgia had 4.4.2 Effect an Larvae. No data available. empty stomachs and suggested that concentrations of microzooplanklon were limiting survival. The smallestsize group of AA larvae in Long Island 4. S Other Factors Affecting Larvae Sound had the highest percentage 9%! of empty 4.5.1 Biotic. No data available, stomachs Covill 1959!. AA survival to metamor- 4.5.2 Abiatic. Inaue et al. 967! suggestedthat phosis at food concentrations of 200, 500 and 1000 wind is an important factor in larval dispersal. At low ratifers/liter corresponding to 0,16, 0,40 and 0.80 temperatures, growth is slower and time ta metamor- calorie/liter! was estimated ta be 0.12%, 5.75% and phosis is longer for AA larvae Smigielski et al. 11.74%, respectively Buckley et al. 1984!. Since 1984!. However, daily mortality ratesof AA larvae in theseestimates were basedon a larval period of fixed the laboratoryare not affectedby temperaturesbetween length 02 days!, slower growth at lower food 2' and 9 C Buckley et al. 1984!. AM larvae survive concentrations would extend the larval period and longer without food at lower temperatures Winshde increase mortality Buckley et al. 1984!. Although 1971!. survival rates at the above food concentrations are 4.5.3 Total Mortality, For larval sand lance AA/ comparable to those of other species, the caloric AD! in the northwestern Atlantic, instantaneous mor- requirements appear to be lower, indicating possible tality ratesbetween 1974 and 1980 rangedfrom .207- adaptation to survival at low food concentrations .363, correspondingta a daily mortality of 6-10% for Buckley et al. 1984!. Comparison of RNA-DNA larvae of 5-27 mrn Morse 1982!. In laboratory ratios in larval sand lance probably AA! collected from experiments,the daily instantaneousmortality rate for the northwestern Atlantic indicated that, in 1982, larvae AA larvae betweenhatching and day 16 was 0.01 and were generallyin poorercondition thanlarvae collected wasunaffected by feedinglevel ar temperature Buckley in 1981, and that some of the 1982 larvae were et al. 1984!. Mortality rates for older larvae 0-43 apparentlylosing weight Buckley 1984!. days! ranged from 0.2-0.02 and decreased with increasingfood level. 4.4 Predation

4.4.1 PredatorSpecies, Marine fish, particularly 4.6 Laboratory Holding and Rearing juveniles, are major predators of larval AH, e.g., in Winslade 971! and Smigielski et al. 984! describe Bristol Bay, sockeye salmon juveniles Straty 1974! techniques for rearing larvae of AM and AA, and sockeye aduIts Nishiyama 1974!; in nearshare respectively. Kodiak, walleye pollock and juvenile pink salmon Rogers et al. 1979!; in Cook Inlet, sockeyeand coho salmonjuveniles and stagharnsculpins Blackburn et 5 JUVENILES al 1980!; in Chatam Sound, juvenile coho salmon Manzer 1969!; in the Strait of Georgia and Saanich 5. 1 Genera! Description Inlet, juvenile salmonids, including coho, churn, sockeyeand king salmon and steelheadtrout, Salmo At 7'C, metamorphosisto the AA juvenilephase takes gairdnerii, Barraclough 1967;Barraclough et al. 1968; placeat a lengthof about29 mm, approximately102 Robinson et al. 1968!; in the Strait of 3uan de Fuca, days after hatching Smigielski et al. 1984!, The juvenile coho and, to a lesserextent, juvenile chinook juvenile stagemay last fram 1-3years, depending on and herring Cross et al. 1978!. Arctic were a the area see 2.1.1!. 26 / Species Synopses

5. 2 Distribution and Abundance Blackburn ct al. 980! reported that harpacticoid 5.2.1 Duration of Juvenile Stage. See 5.1.! copepodswere importantin April, alongwith shrimp 5.2.2 Large-scalePatterns. Juvenilescommonly co- and fish larvae, while calanoid copepods were most occur with adults,although they may also be presentin abundant the remainder of summer. inshore areas where adults are absent Richards et al, 5,3,3 Prey Density Requirements.No data avail- 1963; Rcay 1970!. Reay 970! suggestedthat this is able. due to the wide dispersal of juvenilesrather than to 5.3.4 Factors Affecting Availability of Prey, No separate nursery areas. AH juveniles are abundant data avaihble, along sandybeaches and in shallow nearshoreareas of 5.3.5 Temporal Patterns of Energy Storage. No the Bering Sea Macy et al. 1978!, Norton Sound data available. Barton 1978!, Kodiak Island Blackburn and Jackson 5.3.6 Evidence of Food Limitation, A high nega- tive correlationwas found between the size of age 1 AP 1980! and Cook Inlet Blackbum et al. 1980!, Juve- niles -age and age 1! exhibit an onshore migration and the abundance of adult fish, indicating possible during the summer in someareas, and thereis some competition for food Hamada 1967!. Nagoshi and evidencethat migration takesplace earlier in the sum- Sano 979! reporteda similar significantnegative correlationbctv ccn growth and abundancein 0-age AP. mer for age I fish Barton 1978; Blackburn and Jackson 1980; Blackburn ct al. 1980!. Large interannualvari- Sekiguchi et al. 976! showed that 0-age AP do not ations have been reported in the catch of 0-age AP in begin to accumulate fat reserves until attaining 45-50 mm in length and that fat content increased with Japan Hamada 1966a, b, c; Inouc ct al 1967; Nagoshi and Sano 1979!. increasinglength; consequently, reduced growth could 5.2.3 Small-scale Patterns. Juvenile AH have been affect survival by reducing energy reserves during the observed schooling with similar-size juvenile herring dormancy period. Hobson 1985!. Richards 976! reported similar observations for AA. AA schoolingbehavior begins at 5.4 Predation 25-30 mm, about the time of metamorphosis to the 5,4.1 PredatorSpecies. SeeTables 1-3 and4,4.1.! juvenile stage Smigielski et al. 1984!. 5.4.2 Effect on Juveniles. No data available. 5.2,4 Maximum and Mean Abundances, No esti- rnates of abundance are available, 5.5 Other Factors Affecting Juveniles 5.2.5 Vertical Distribution. Juveniles may be more numerous than adults in surface waters at night, based 5.5.1 Biotic. No data available. on catchesof AP Senta 1965b! and AM Macer 1966!. 5.5.2 Abiotic. Winters981! reporteda significam ln thc laboratory, burrowing behavior in AA juveniles positive relationship between water temperatureand is first observed when they are between 35 and 40 mm first year growth in AD. in length Smigiclski ct al 1984!. 5.5.3 Total Mortality. No dataavailable. 5.6 Laboratory Holding and Rearing 5.3 Feeding Sce 2.6.! 5.3.1 Habitat. Juvenilesfeed almost exclusively in the water column, but no information is available on 6 CURRENT H YPO TIIESKS fomging depth range. ON FACTORS AFFECTING 5.3.2 Prey Species. Studies of the food habits of YEAR-CLASS ABUNDANCE juvenile AH from the Chukchi Sea Springer ct al, 1984!, western Aleutians Simenstad et al. 1978!, Environmentrti effects: Hamada 966c! correlated Kodiak Harris and Hartt 1977a, b; Rogersct al. 1979!, increasedcatches of 0-age sand lance AP! with low Cook Inlet Blackburn et al. 1980! and the Strait of water temperatures during the spawningseason and also Georgia Barraclough1967! agree in thcpredominance with the number of days with westerly winds during of copcpods in the diet. Other prey items of some the threeweeks following peak spawning, importanceinclude other crustaceanlarvae, larvaceans, Density-dependenteffecrs on recruitment:A strong cladocerans, chaetognaths, fish larvae, mysids and negativecorrelation r = -.87! betweenthe percentageof gammarid amphipods. In the lower Cook Inlet area, age-I AP in thccatch and thc catch of 0-agcfish was Pacific Sand Lance / 27

observedby Hamada 966a!. In addition, Hamada AD! abundance were attributable to decreases in cod 967! reporteda negativecorrelation r = -.81! between biomass on the Newfoundland Grand Bank between thc numberof eggsspawned and the catchof 0-ageAP. 1968and 1979. The presenceof greaterpercentages of Density-dependentsects on growth: Small size of older age groups indicated reduced mortality, and a age-I AP wascorrelated with large catchesof adult -3 changein the dominantage group from age 4 to age 3 yearsold! fish Hamada1967!. Similarly, Nagoshiand suggestedincreased recruitment. Using data froin re- Sano 979! found a negative correlation r = -.86! be- searchtrawl surveys,Winters 983! found significant tween growth of 0-age AP and an index of population negative correlations of sand lance abundance and density. recruitment with cod biomass. l.ood availability: The availability of adequatefood hasfrequently been suggested as an importantfactor in IarvaI mortality. Low food availability may lead to 7 LITERATURE CITED reduced growth and consequentlya prolonged larval phaseand increasedvulnerability to predation Buckley Ainlcy, D.G. and G.A. Sanger. 1979. Trophic rela- 1984!. Laboratory studieson AA larvae provide esti- tions of scabirds in the northeastern Pacific Ocean matesof minimum growth rates necessaryto prevent and Bering Sea. In Conservationof marine starvationat a rangeof temperaturesand a model pre- of northern . U,S. Fish WildI. dicting recentlarval growih basedon RNA-DNA ratios Scrv., Res. Rep. Fish 11: 95-122. and temperature Buckley 1984; Buckley et al 1984!. Altukhov, K.A. 1978, On the reproductionand abun- Larvae probably AA! collectedduring 1981and 1982 dance of the lesser sandlance, A. tnarintts, in the indicate differences in larval condition RNA-DNA White Sea, J. Ichthyol. 18: 560-567. ratio! that could be attributed to food availability Andersen, K.P. and E, Ursin. 1978. A multispecies B uckley 1984!. Although no direct.evidence was pre analysisof the effects of variations of effort upon sentcdto indicate diffcrcnccs in larval survival, based stock composition of eleven North Sea fish on RNA-DNA ratios, a higher perceniageof larvae species. Rapp. P.-v. Reun. Cons. perm. int. werc in poor condition in 1982, and some had appar- Explor. Mer 172; 286-291. ently beenlosing weight Buckley 1984!. Andriyashev, A.P. 1954. Fishes of the northern seas Replacement and predation: Andersenand Ursin of the USSR. Akad. Nauk SSSR, Opred. po. 978! presentedthe hypothesis that the reduction of Faune SSSR 53, 566 p. 964 transL avail., Nat, herring and mackereI stocks have resulted in their Tech. Int. Serv., Springfield, VA, OTS 63- replacementin theNorth Scaecosystem by small,fast 11160!. growing opportunistic species such as sand Iance. Barraclough,W.E. 1967. Number, size, and food of However, thc lack of prior data on the abundance of larval andjuvenile fish caughtwith an IsaacsKidd sandlance makesthe hypothesisdifficult to test unless trawlin the surfacewaters of theStrait of Georgia, herring and mackerel stocksreturn to former levels of April 25-29,1966. Fish.Res. Bd Can,,MS Rep, abundance, Scr, 926, 79 p, Shermanet al. 981! presenteda similarhypothesis Barraclough, W.E., D.G. Robinson, and J.D. Fulton. for changesin the ecosystemstructure of the north- 1968. Datarecord. Number,size composition, western Atlantic. Larval surveys indicated a substantial weight,and food of larvaland juvenile fish caught increasein the abundanceof sandlance AA/AD! larvae with a two-boat surface trawl in Saanich Inlet between1974 and 1979which apparentlyreflects a April 23-July 21, 1968. Fish. Res. Bd Can,, MS large increase in spawning stock bioinass Morse Rep. Ser. 1004, 305 p. 1982!. The authors suggestthat this rapid increasein Barton, L.H. 1978. Finfish resourcesurveys in sandlance population size was directly related to the Norton Soundand KotzebueSound. U.S. Dep. reductionof mid-size predators,such as cod, haddock, Commer.,NOAA, OCSEAPFinal Rep. 4: 75- hake,mackerel and herring. 313. RV0019. Bothsand lance AD! andcapclin are important prey Beacham,T,D. 1986.Type, quantity and size of food itemsfor off Newfoundland Lilly 1982; of Pacific salmon Oncorhynchus! in the Strait of Lilly and Fleming 1981; Winters 1983!. Winters Juan de Fuca, British Columbia, . Fish. 983! hypothesizedthat recent increasesin sandlance Bu11., U.S. 84: 77-90 28 / Species Synopses

Best, E.A. and W.H. Hardman. 1982. Juvenile Commer., NOAA, OCSEAP Final Rep., Biol halibut surveys, 1973-1980. Int. Pacif. Halibut Stud. 11: 175-487. RV0237. Commn Tech, Rep, 20,3$ p. Dunn, J.R. and N.A. Naplin. 1974, Fish eggs and Blackburn, J.E. 1973. A survey of the abundance, larvae collected from waters adjacent to Kodiak distribution, and factors affecting distributions of Island, Ahska, during April and May 1972. U.S. ichthyoplankton in Skagit Bay. M.S. Thesis, Dep. Comm., NWAFC, NMFS, NOAA. Univ. Washington,Seattle, 136 p. MARMAP Rep. 12, 61 pSeattle, WA. Blackburn, J.E. and P.B. Jackson. 1980. Seasonal Einarsson, H. 1951, The postlarval stagesof sand composition, abundance and food web eels Ammodytidae! in Faroe, , and west relationshipsof principal juvenile andadult marine waters. Acta nat. islandica 1: 5-75. finfish speciesinhabiting the nearshorezone of Fresh, K.L. 1979. Distribution and abundance of Kodiak Island's eastside. U.S. Dep. Commer., fishes occurring in the nearshoresurface waters of NOAA, OCSEAP Final Rep., 101 p. northern Puget Sound,Washington, M.S. Thesis, Blackburn, J.E., K. Anderson, C.I. Hamilton, and S J. Univ. Washington, Seattle, 120 p. Starr, 1980. Pelagicanddemersal fish assessment Frost, K.J. and L.F. Lowry. 1981. Foods and trophic in the lower Cook Inlet estuary system. U.S. relation-shipsof cetaceansin the Bering Sea. Ln Dep. Commer., NOAA, OCSEAP Final Rep., D.W, Hood and J.A. Calder editors!, The eastern Biol. Stud. 12: 259-602. RV0512. Bering Sea shelf: Oceanographyand resources, Brown, R.F. and B.R. Mate. 1983, Abundance, vol. 2: 825-836. NOAA/Office of Marine Pollu- movements, and feeding habits of harbor seals, tion Assessinent. Distributed by Univ. Wash- Phoca vituliria, at. Netarts and Tillamook Bays, ington Press,Seattle.] Oregon. Fish. Bull., U.S. 81: 291-301. Girsa, I.I. and A.N. Danilov. 1978. The defensive Buckley, L.J. 1984. RNA-DNA ratio: An index of behavior of the White Seasand lanceAmmodytes larval fish growth in the sea. Mar. Biol. 80: 291- hexapterus,J. Ichthyol. 18; 862-865. 298, Hamada, T. 1966a. Studies on fluctuation in the Buckley, L.J., S.I. Turner, T.A. Halavik, A.S. abundance of larval sand-lance in the Harima-Nada Smigielski, S.M. Drew, and G.C. Laurence. and Osaka Bay. I. Relation between the progeny- 1984. Effects of temperatureand food availability abundance and the age composition of parent fish. on growth, survival, and RNA-DNA ratio of larval Bull. Jap. Sac. scient Fish. 32: 393-398. sand lance Ammodytes americartus!. Mar. Ecol. Hamada, T. 1966b. Studies on fluctuation in the Prog. Ser. 15: 91-97. Covill, R.W. 1959, Food and feeding habits of larvae abundance of larval sand lance. II. The distribution and its seasonalchange of larval fish. and post-larvae of Ammadytes americarius, 1952- Bull, Jap. Soc. scient. Fish. 32: 399-405. 1955. In Oceanography of Long Island Sound: 125-146. Bull. Bingham oceanogr. Collect. 17 Hamada, T. 1966c. Studies on fluctuation in the abundanceof larval sand-lance.III. Relationship ! Craig, P.C. 1984. Fish use of coastal waters of the to weather and sea conditions during the breeding Alaskan Beaufort Sea: a review. Trans. Am, season. Bull. Jap. Sac. scienL Fish. 32: 579-5$4. Fish. Soc. 113: 265-282. Hamada, T, 1967. Studies on fluctuation in the Cross, J.N., K,L, Fresh, B.S. Miller, C.A. Simenstad, abundance of larval sand-lance. IV. Relation S.N. Steinfort, and J.C. Fegley. 1978. Nearshore betweenthe numberof eggsand the catchof 0-age fish and macro-invertebrateassemblages along the fish. Bull. Jap. Soc. scient. Fish. 33: 410-416. including food habits of Harris, C.K. and A.C. Hartt. 1977a. Assessment of common nearshore fish. Univ. Washington, Fish. pelagic and nearshorefish in threebays on the east Res. Inst. Ann. Rep. FRI-UW-781$, 188 p. and south coasts of Kodiak Island, Alaska. Univ. Dick, M.H. and I.M. Warner. 1982. Pacific sand Washington, Fish. Res, Inst. FRI-UW-7719, lance, Ammodytes hexapterus Pallas, in the 190 p. Kodiak island group, Alaska, Syesis15; 43-50. Harris, C.K. and A,C. Hartt. 1977b. Assessment of Drury, W.H., C. Ramshell and J.B. French, Jr. 1981, pelagic and nearshorefish in threebays on the east Ecological studiesin the Bering Strait. U.S. Dep. and south coasts of Kodiak Island, Alaska. U.S. Pacific Sand Lance / 29

Dep, Commer., NOAA, OCSEAP Final Rep. U.S. Dep. Commer., NOAA, NMFS. NWAFC RV0485. Proc. Rep. 80-8, 393 pSeattle, WA, Hart, J.L. 1973. Paciflic fishes of Canada. Bull. Fish. Kitaguchi, T. 1977. Some biological observations on Res. Bd Can. 180, 740 p. the fishery of I-KA-NA-GO sand-lance, genus Hashimoto, H. 1984a. Two subpopulations of sand- Ammodytes! in waters around Soya Straits. ecl found off Torttori Pref. Bull. Jap, Soc, scient, Hokkaido Fish. expl. Sta. Rep. 34: 1-12. Fish. 50: 1089-1095. Kitaguchi, T. 1979. A taxonomic study of sand Hashimoto, H. 1984b. Population structure of the lances genus Ammodytes! in the waters of the Ammodytes around Japan. Bull. Jap. northerncoast of Hokkaido. Hokkaido Fish. expl, Soc. scient. Fish. 50: 1357-1366. Sta. Rep. 21: 17-30. Hatch, S,AD,R, Nysewander,A.R. DeGange,M.R. Kitakata, M. 1957. Fishery biological studies of I- Petersen, P.A. Baird, K.D, Wohl, and C,J. KA-NA-GO Sand lance,Ammodytes personatus Lensnik. 1978. Population dynamicsand trophic Girard! in waters around Hokkaido. II. Age and relationshipsof marine birds in the Gulf of Alaska growth. Hokkaido Fish. expl. Sta. 16: 39-48. and southern Bering Sea. U.S. Dep. Commer., Kobayashi, K. 1961. Larvae and young of the sand- NOAA, OCSEAP Ann. Rep. 3: 1-68. RV0341. lance, Ammodytes hexapterus Pallas, from the North Pacific. Bull. Fac. Fish. Hokkaido Univ. Hobson, E.S, 1986. Predation on the Pacific sand lance, Ammodytes hexapterus, Pisces: Ammo- 12: 111-120. Pn Japanese,English abstract.] LeBrasseur, R,J., W.E. Barraclough, D.D. Kennedy, dytidae! during the transition between day and night in southeastern Alaska, U.S.A. Copeia and T.R. Parsons. 1969. Production studies in the Strait of Georgia. Part III. Observationson 1986: 223-226. the food of larval and juvenile fish in the Fraser Hunt, G.L., Jr., B. Burgeson,and G,A, Sanger, 1980, River plume, Feb. to May 1967. J. expl. mar. Feeding of of the eastern Bering Biol. Ecol. 3: 51-61. Sea 4 D.W. Hood and J.A. Calder editors!, The Leim, A.H. and W.B. Scott. 1966. Fishes of the eastern Bering Sea shelf: Oceanography and Atlantic Coast of Canada. Bull. Fish. Res. Bd resources, vol. 2: 629-647. NOAA/Office of Can. 155, 485 p. Marine Pollution Assessment, [Distributed by Lilly, G.R. 1982. Influence of the Labrador Current Univ. Washington Press, Seattle.] on predationby cod on and sandlance off Hunter, M. 1979. Food resourcepartitioning among eastern Newfoundland. NAFO ScienL Coun, demersal fishes in the vicinity of Kodiak Island, Stud. 3: 77-82. Alaska, M,S, Thesis, Univ. Washington,Seattle, Lilly, G.R. atid A.M. Fleming. 1981. Size rela- 131 p. tionships in the predationby Atlantic cod, Gadus Inoue, A., S. Takamori, K. Kuniyuki, S, Kobayashi, morhua, on capelin, Mallotus villosus, and and S. Nishina. 1967. Studies on the fishery sandlance, Ammodytes dubius, in the Newfound- biology of the sand-lance Ammodytes personatus land area, NAFO Scient. Coun. Stud, I: 41-45. Girard, Bull. Naikai reg. Fish. Res. Lab, 25; 1- Lindberg, G.U. 1937. On the classification and 335. distribution of sand-lances genus Ammodytes Jewett, C. 1978. Summer food of the Pacific cod, Pisces!. Bull. Far Eastern Branch Acad, Sci. Gadus macrocephalus, near Kodiak Island, AIaska. USSR 27: 85-93, Fish. Bull., U.S. 76: 700-706. Lowry, L.F. and K.J. Frost. 1981. Feeding and Kajimura, H. 1984. Opportunistic feeding of the trophic relationshipsof phocid sealsand walruses northern fur seal, Callorhinus wisinus, in the in the easternBering Sea. Q D.W. Hood and J,A. easternNorth Pacific Oceanand the easternBering Calder editors!, The eastern Bering Sea shelf: Sea. U.S. Dep. Coinmer., NOAA Tech, Rep, Oceanography and resources, vol. 2: 813-824. NMFS SSRF-779,49 p. NOAA/Office of Marine Poliution Assessment. Kendall, A.W., Jr., J.R. Dunn, R,J. Wolotira, Jr., J.H, [Distributed by Univ. Washington Press, Seattle.] Bowerman, Jr., D.B. Dey, A,C. Matarese, and J.E. Macer, C.T. 1966. Sandeels Ammodytidac! in the Munk. 1980. Zooplankton, including ichthyo- southwesternNorth Sea: their biology and plankton and decapodlarvae, of the Kodiak shelf. fishery. Fishery Invest., Lond., Ser. 2, 24: 1-55. 30 i Species Synopses

Macer, C.T. and A.C. Burd. 1970. Fishing for Norcross, J. J., W.H. Massman, and E.B Joseph, sandeels. Lowestoft Fish. Lab. Leafl. No. 21, 9 1961. Investigations of inner continental shelf p waters off lower ChesapeakeBay. Part EL Sand Macy, P.T., J.M. Wall, N.D, Lampsakis, and J.E. lance larvae,Ammodytes ameri canus. Chesapeake Mason. 1978. Resources of non-salmonidpelagic Sci. 2: 49-59. fishes of the Gulf of Alaska and eastern Bering Ogi, H. and T. Tsujita. 1973. Preliminary exami- Sea, U.S. Dep. Commer., NOAA, NMFS, nation of stomachcontents of murres spp.! NWAFC Final Rep. OCSEAP Task A-7. Parts I from the easternBering Seaand Bristol Bay, June- and II, 714 p. August, 1970 and 1971. Japan J, Ecol. 23: 201- Manuwal, D.A. and D. Boersma, 1978, Dynamicsof 209. marine populations on the Barren Islands, Outram, D.N. and C. Haegele. 1972. Food of Pacific Alaska. U.S. Dep. Commer., NOAA, OCSEAP hake Merlucci us productus! on an offshore bank Ann. Rcp, 3: 575-679. RV0341. southwest of Vancouver Island, British Columbia, Manzer, J,l. 1969. Stomach contents of juvenile J. Fish. Res. Bd Can. 29: 1792-1795. Pacific salmon in Chatam Sound and adjacent Pearson,W,H,, D.L. Woodruff and P.C. Sugarman. waters. J. Fish. Res. Bd Can, 26; 2219-2223. 1984. The burrowing behavior of sand lance, Mason, J.C., O.D. Kennedy, and A.C. Phillips. Amrnodytes hexapterus: effects of oil-contami- 19&la. Canadian Pacific coast ichthyoplankton nated sediment. Mar. envir. Res. 11: l.7-32. survey: 1980, Ichthyoplankton, Cruise Two, Pinto, J, 1984, Laboratory spawning of Ammodytes 1980 February 13-20!. Can. Data Rep. Fish. hexapterus from the Pacific coast of North aquat. Sci. 276. America with a description of its eggs and early Mason, J.C., O.D. Kennedy, and A.C. Phillips, larvae. Copeia 1984:242-244. 1981b. Canadian Pacific coast ichthyoplankton Pinto, J.M., W,H. Pearson and J.W. Anderson. 1984. survey: 1980. Ichthyoplankton, Cruise Three, Sedimentpreferences and oil contaminationin the 1980 March 12-20!. Can. Data Rep. Fish. aquat. Pacificsand lance Ammodytes hexapterus. Mar. Sci. 277. Biol. 83: 193-204. Meyer, T.L., R.A. Cooper, and R,W, Langton. 1979. Pitcher, K.W. 1980. Food of the harbor seal, Phoca Relative abundance, behavior, and food habits of vitulina richardsi, in the Gulf of Alaska. Fish. the American sandlance, Ammodytes americanus, Bull., U.S. 78: 544-549. from the Gulf of Maine. Fish. Bull., U.S. 77: Pitcher, K.W. 1981. Prey of the Stellar sea lion, 243-253. Eumetopiasjubatus, in the Gulf of Alaska. Fish. Morse, W. 19&2. Spawning stock biomassestimates Bull., U,S. 79: 467472. of sand lance, Ammodytes sp., off northeastern Reay, P.J. 1970. Synopsisof biological data on North United States, determined from MARMAP Atlantic sandeelsof the genusAmmodytes A. plankton surveys, l974-1980. Int., Cons. Explor. tobianus, A. dubius, A, americanus, and A. Sca, C.M. 1982/G: 59, marinus!. FAO Fish. Biol. Synop. Na. 82. Nagoshi,M. andM. Sano. 1979. Populationstudies Reay, P.J, 1972. The seasonalpattern of otolith of sand-ecl,Ammodytes personatus, in lsc Bay.I. growth and its application to back-calculation Growth and it relation to population density. studies. of Ammodytes tobianus L. J, Cons. Japan J. Ecol. 29: 1-10. perm. int. Explor, Mer 34: 485-504, Ncmoto, T. 1959. Food of baleen whales with Reimchen,T.E, and S. Douglas. 1984. Feeding reference to whale movements. Whales Res. Inst. schedule and daily food consumption in red- Sci. Rep. 14: 149-290. throatedloons Gaviastellata! over the prefiedgling Nishiyama, T. 1974. Energy requirementsof Bristol period, 101: 593-599. Bay sockeyesalmon in thecentral Bering Sea and Richards,S.W. 1965.Description of thepostlarvae of Bristol Bay. + D.W. Hood and E.J. Kefiy the sandlance Ammodytes!from the eastcoast of editors!,Oceanography of the Bering Seawith North America. J, Fish. Res. Bd Can. 22: 1313- emphasis on renewable resources: 321-343. Univ. 317. Alaska, Fairbanks, Inst. Mar. Sci. Occas. Publ. Richards,S.W. 1976. Mixed speciesschools of No. 2. postlarvaeof Ammodyteshexapterus and Clupea Pacific Sand Lance / 31

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