ammodyfes hexapterus with notes on relapsed ammodytes …

PACIFIC SAND LANCE Ammodyfes hexapterus WITHNOTES ON RELAPSED Ammodytes SPECIES

L. Jay Fieldl

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

1 BACKGROUND

1. 1 Literature Search

1. 2 Unit Stocks andTheir Relationships

An intensive literature search, except for Japanese andSoviet literature, which was not as thorough, wascompleted in December 1986, The emphasis of thesearch was on the sand lance of the northeasternPacific, including the Bering Sea.

The taxonomy of Ammodyres species is uncertain.The species are primarily distinguished on the basis ofgeographic distribution and meristic counts. However,meristic counts frequently exhibit wide variation andlatitudinal clines Richards et al. 1963; Reay 1970;Winters 1970; Scott 1972b; Richards 1982!. In thisreport, the general classification scheme proposed byReay �970! will be followed. Ammodyres hexaprerus AH! is the only sand lance reported from thenortheastern Pacific Hart 1973!. They are found inshallow nearshore waters froin California to the

Iteaufort Sea Trumble 1973; Craig 1984!. As detailedtaxonomic studies have not been carried out, the

relationship of AH to sand lance from other areas hasnot been determined. Consequently, Ammodyres spe-

cies from the northwestern Pacific/Japan and thenorthwestern and northeastern Atlantic also will be

cons~Two species of Ammodyres, AH and A.

personarus AP!, have been reported from the north-western Pacific; AH generally is restricted to north of45'N latitude Kitaguchi 1979!. The two species aredistinguished by meristic counts, with AH having thehigher number of vertebrae and dorsal fin rays Lindberg 1937; Kitaguchi 1979!. Both species show agreater number of vertebrae with increasing latitude Kitaguchi 1979!. Andriyashev �954! considered themto be conspecific, and Hashimoto �984a! suggestedthat Ammodyres may be represented by three sub-species along the coast of Japan.

Two other species of Ammodyres, a northernoffshore species, A. dubius AD!, and a southern in-shore species, A. americanus AA!, have been reportedfrom the northwestern Atlantic Reay 1970!. However,overlap in geographic range and clines latitudinal andinshore-offshore! in meristic counts that may becorrelated with environmental conditions suggests thepossibility that only a single species occurs in this area Richards et al. 1963; Winters 1970; Scott 1972b;Richards 1982!. Two species, A. robianus AT! and A.marinus AM}, are found in the northeastern Atlantic,but only AM will be considered in this report, as ATdoes not resemble closely the species from other areas Reay 1970!.

16 / Species Synopses

1.3 The Fishery1.3.1 Relative Size and Importance. Except for

occasional small bait fisheries in Washington andBritish Columbia, no fishery currently exists in thenortheastern Pacific, although there is some potential Trurnble 1973!. In Japan the fishery takes about100,000 ton pcr year Kitaguchi 1977!. No fishery forsand lance exists in the northwestern Atlantic, althoughfeasibility studies indicate considerable potential R.Smith, Univ. New Hampshire, pers. comm.!. TheEuropean fishery landed 200,000 mt in 1968 Reay1970!,

1.3.2 Age at Recruitment. In Japan, sand lance AP! usually enter the fishery at age 1, but 0-age fishmay enter late in the season, particularly if the yearclass is strong Kitaguchi 1977!. In the North Sea,recruitment to the AM fishery inay begin at agc 6months but most enter at age 1 Macer 1966; Reay1970!.

1.3.3 Characteristics of Harvested Fish. Length ofAP/AH in the Japanese fishery ranges from 8 to 26 cm Kitaguchi 1977!, In the northeastern Atlantic fishery,AM range from 5-25 cm in length Macer 1966!.

1.3.4 Types and Selectivity of Gear, In Japan, avariety of seines, lift nets and bottom trawls are usedfor the sand lance fishery Inoue ct al. 1967!. ThcEuropean fishery primarily uses a high opening bottomtrawl with a 6 mm cod-end mesh Macer 1966; Macerand Burd 1970!. The selection length for this gear wasestimated to be approximately 8-9 cm Reay 1970!.

1.3.5 Distribution of Fishing Effort, The Japanesesand lance fishery takes place in depths of 80 m or less,in areas of sand and shell bottom Kitaguchi 1977!. Insouthern areas, the AP fishery runs from mid-March tolate June Inoue et al. 1967!, but at higher lautudes theseason is somewhat later, sometimes extending fromApril to December Kitaguchi 1977!.

The European AM fishery is conducted in depths ofless than 40 m in sand-bottom areas, primarily on off-shore banks Macer 1966!. The fishing season extendsfrom April to September, with very low catches atother times of the year Macer 1966; Reay 1970!.Fishing takes place almost exclusively during daylighthours Macer 1966!.

1.4 Distribution and Abundanceof the Population

Sand lance are abundant in shallow ncarshore areas.

1.5 Current Status of Stocksand of Management Measures

No information is available on the status of stocks ofAH in the northeastern Pacific, since no fisherycurrently exists and AH are not sampled by conven-tional research trawls.

1. 6 Recruitment VariabilityEvidence for AH recruitment variability is mainlycircumstantial. For example, the presence or absenceof sand lance in seabird stomachs has been noted duringtrophic studies Vermeer 1979; Drury et al. 1981;Springer et al. 1984!. In Japan, there is considerablevariability in the percentage of 0-age fish in AH/APcatches Hamada 1966a,b,c; Inoue et al. 1967;Kitaguchi 1977; Nagoshi and Sano 1979!,

Larval surveys in the northwestern Atlantic indicatelarge interannual variation in sand lance AA/AD!recruitment. Between 1974 and 1979, abundance oflarval sand lance increased by a factor of 20 Smith etal. 1978; Sherman et al. 1981!, which reflects a 50-foldchange in adult spawning biomass Meyer et al. 1979;Morse 1982!.

1.7 Age Determination and Validation

Age determination for sand lance species is based onsurface readings of whole otoliths Kitakata 1957;Macer 1966; Scott 1968,1973; Reay 1970, 1972,Kitaguchi 1977; Winters 1981!. Validation of ages hasbeen attempted by observing the annual pattern of ringformation Kitakata 1957; Reay 1972; Kitaguchi1977!.

1.8 Age Composition ofthe Population

In sand lance fisheries, age composition is apparentlysubject to variation with time of year, area and gear inaddition to interannual differences Inoue et al. 1967!.AH collected by beach seine from the vicinity ofKodiak Island, Alaska, ranged from age 0 to 5, withage 1 fish most abundant Dick and Warner 1982!.However, differences in age composition between AHcollected by beach seine and those dug from gravelbeaches indicate that beach seines were not effective insampling older age classes Dick and Warner 1982!.

In northern areas of Japan where both AP and AHoccur, most of the catch is composed of ages 1-3, andfish over age 4 are uncommon; there is some evidence

Pacific Sand Lance / 17

of interannual variability Kitaguchi 1977!. In con-trast, in the Seto Inland Sea fishery, 0-age fish com-prise about 80% of the catch and age-2 fish less than5% Inoue et al. 1967!, whereas in the Harima-Nadaand Osaka Bay fishery, age-1 AP account for 20-77%of the catch Hamada 1966a!.

Macer �966! reported that the percentage of olderfish AM! in the catch decreased during the last part ofthe season, and Winslade �974c! suggested this wasdue to age related differences in timing of over-wintering see 2.2.5!. The maximum age of AD inNewfoundland is age 10 or over Winters 1983!. Ages3-5 predominate in the survey catches, but the relativeabundance of AD older than age 6 has been increasingsince the late 1970s Winters 1983!. In thenortheastern Atlantic, the maximum age of AM is 9,but ages 1-3 account for the majority of the catch Reay 1970!.

2 ADULTS

2.1 General Description

2.1.1 Size and Age Ranges. Based on a smallsample size n=7! of mature fish, AH in the vicinity ofKodiak Island, Alaska, mature at ages 2-3; the smallestmature sand lance observed was 128 mm Dick andWarner 1982!. In the Soya Strait region of Japan,most sand lance AH/AP! spawn at the end of theirsecond year but some spawn at age 1 Kitaguchi 1979!.Most AP from Ise Bay to the south mature at ages 1-2at lengths greater than 110 mm Nagoshi and Sano1979!.

In the North Atlantic, AA usually are mature �02-114 m! at the end of their first year Richards 1982!.Scott I968! reported that most AD mature at age 2 atlengths between 180 and 200 mm, whereas Winters�983! found that 50% were mature at age 3 at 180 mmin length. Most AM are mature at age 2, but large�00 mm! individuals may mature at age 1 Macer1966!.

2.1.2 Type of Spawning Behavior. Most species ofsand lance spawn once a year Reay 1970!, Unimodaldistributions of egg diameters in ovaries of mature AH Pinto 1984!, AD Scott 1972a! and AM Macer 1966!indicate a single spawning per year, AA that spawnedin laboratory holding tanks had very few eggsremaining in their ovaries, indicating that aIl the eggsto be spawned in a year are released in a single batch Smigielski et al. 1984!.

2.1.3 Fecundity. Fecundity-length relationships F= number of eggs, L = length in cm! have been deter-mined as follows:

AA: F = 0.328 L = 3.857 Westin et al. 1979!;AM: F = 2.046 L = 3.055 Macer 1966!,

Sand hnce probably AM! between 120 and 195 mm standard length! off the Murrnan coast had between3300 and 22,I00 eggs average 6800! per female Andriyashev 1954!.

2. 2 Distribution and Abundance

2.2.1 Duration of Spawning. Most information forthe time of spawning of sand lance is based on the oc-currence of early larvae. In Puget Sound, Washington,yolk-sac AH larvae are most abundant in late Januaryto early March R. Trumble, Washington Dep.Fisheries, pers, comm.! and spawning occurred inoutdoor holding tanks in mid-March Pinto 1984!. Offthe west coast of Vancouver Island, larval AH began tooccur in February Mason et al, 198la, b!. Larvalsurveys in the Kodiak region indicated late winter February-March! spawning Rogers et al. 1979;Kendall et al. 1980!. However, Dick and Warner�982! reported intertidal spawning off Kodiak!sland inOctober. Spawning of sand lance AH/AP! in the SoyaStrait region of Japan occurred between January andearly May, and there was some evidence of interannualvariation Ki taguchi 1977!.

2.2.2 Large-scale Patterns. In general, sand lanceoccur in shallow nearshore areas, usually in depths lessthan 100 m, with sand or sand-gravel substrates. Theyare abundant from spring to late summer anduncommon during the remainder of the year when theypresumably are buried in the sand Leim and Scott1966; Reay 1970; Trumble 1973!. While inshore-offshore movements have been reported by a number ofauthors, there is no evidence of large-scale migrationsof any sand lance species Reay 1970!. AH juvenilesand adults are commonly abundant in nearshore watersduring spring-summer but are rarely sampled duringlate fall and winter Barton 1978; Fresh 1979;Blackburn and Jackson 1980; Blackbuin et aI. 1980!.According to Blackburn and Jackson �980!, AH areinactive during the winter and are often found buried inintertidal sand. In the Bering Sea, they have beenreported from 0- IOO m but are most common in depthsless than 50 m Shuntov, in Macy et a!. 1978!.

2.2.3 Small-scale Patterns. All species of sandlance exhibit schooling behavior, often forming large


aggregations Reay 1970; Trumble 1973!. Meyer et al.�979! observed schools of AA ranging in numberfrom approximately a hundred to tens of thousands; thelargest fish were in the center surrounded by smallerfish.

2.2.4 Maximum and Mean Abundances. Noinformation on abundance of AH is available. An

estimate of spawning biomass of sand lance AA/AD!in the northwestern Atlantic from larval survey dataindicated that biomass increased 50-fold between 1974

and 1978 Morse 1982!.2.2,5 Vertical Distribution. Seasonal and diurnal

variations in the catch of sand lance are thought to be,at least partially, the result of their habit of burrowingin the sand when not feeding. On the basis ofunderwater observations of AH, Hobson �986!reported that same sand lance were buried in the sand atall times of the day, but most were feeding in the upperlevels of the water column during the day and buried inthe sand at night. Trawl catches of AH were muchreduced at night Macer 1966!.

2.3 Feeding

2.3.1 Habitat. Sand lance AH! feed primarily inthe water column, although epibenthic invertebratesoccasionally appear in their diet S imenstad et al. 1979;Rogers et al. 1979!.

2.3.2 Prey Species. Calanoid capepods are themajor prey for AH adults and juveniles from the west-ern Aleutians Simensrad et al. 1978!, Kadiak Harrisand Hartt 1977a, b; Rogers et al. 1979! and Washing-ton Simenstad et al. 1977,1979; Crass et al. 1978!.Other prey are crustacean larvae, rnysids, gammaridamphipods, harpacticoid copepods, chaetognaths, lar-vaceans and polychaetes. According to Rogers et al.�979!, epibenthic invertebrates, particularly mysidsand gammarid amphipods, inc~ in relative import-ance for AH during autumn and winter; however, totalstomach content weight was much lower. AA made asimilar shift to benthic prey during early spring whenzooplankton abundance was low Richards 1982!. Theprey spectra of other species of sand lance are verysimilar ro that of AH see Inoue et al. 1967; Sekiguchiet al. 1974,1976; Kiraguchi 1977; and Hashimoto1982 for AP; Richards 1965, 1982; and Meyer et al.1979 for AA; Scott 1973 for AD and Macer 1966 forAM!.

2.3.3 Prey Density Requirements. Not applicable.2.3.4 Factors Affecting Availability of Prey. Since

calanoid capepods are the primary prey of sand lance,

any factors affecting their abundance and distributionpresumably could affect sand lance feeding. Calanoidcopepods also are preyed on to a great extent byjuvenile PaciTic herring that frequently co-occur withsand lance Harris and Hartt 1977a, b; Simenstad et al.1979!.

2.3.5 Temporal Patterns of Energy Storage. Sea-sonal variation in fat content has been demonstrated for

AP Inoue et al. 1967; Sekiguchi et al, 1976! and AM Kuhl and Luhinan, in Winslade 1974c!, Lowestvalues were recorded at the beginning of the feedingperiod and peak fat content is reached at the end of thefeeding season just prior ta a prolonged period of dor-mancy during which the sand lance remain buried in thesand, Fat supply is probably important for main-tenance metabolism and gonad maturation Winslade1974c; Sekiguchi et al. 1976!.

2,3,6 Evidence of Food Limitation. No data avail-

able.

2.4 Predation

2.4.1 Predator Species. A wide variety of marinefish, seabirds and mammals are major predators of AHover their entire range Tables 1-3!.

2.4.2 Effect on Spawning Adults. No estimates ofthe importance of predation have been made for AH,although apparently it is very high, Evidence for spe-cies from the northwestern Atlantic Sherman et al.1981; Winters 1983! and the North Sea Andersen andUrsin 1978! suggests decreased levels of predation ansand lance have led to large increases in sand lancepopulations.

2.5 Other Factors Affecting Adults

2.5.1 Biotic. Mass mortalities of sand lance alongthe coast of England has been associated with bloomsof Gonyardax sp Adams, in Reay 1970!. See 2.5.2for laboratory experiments related to food availabihty.!

2.5.2 Abiotic. As a consequence of their habit ofburrowing into sandy bottoms for extended periods,sand lance usually are found in association with sandand fine gravel bottoms Reay 1970; Trumble 1973!.Both particle size and circulation appear ro be import-ant factors, although specific habitat requirements areunknown. Laboratory experiments showed thar. AHavoided oiled sand Pearson et al. 1984; Pinto et al.1984!. AH mortality was noted in connection with oilpollution and detergent cleanup in the Torrey Canyonoil spill Smith, in Reay 1970!.

Table 1. Marine fish predators of sand lance AH!.

PE IE

Aleutians

Kodiak

British Columbia

Simenstad et al. 1977

Jewett 1978; Hunter 1979Westrheim and Harling 1983

PaciTic cod

Smith et al. 1978

Simenstad et al. 1977Hunter 1979

Best and Hardman 1982

Halibut

British Columbia

IGxRk

British ColumbiaHarris and Hartt 1977; Hunter 1979Westrheim and Harling 1983

Rock sole

Harris and Hartt 1977; Rogers et al. 1979

Hunter 1979

Westrheim and Harling 1983

Hart 1973

Rosenthal et al. 1981, 1982,

British Columbia

British ColumbiaLingcod

Rockfish black,yellowtail, dusky,widow!

Gulf of Alaska

Rosenthal 1983

Salmon chumsockeye!

W. Aleutians Simenstad et al. 1977

Salmon chinook,coho, sockeye, pink!

British Columbia Beacham 1986

Great sculpin Harris and Hart 1977

Yeilowfin sole

Sand sole

Petrale sole

S.E. Bering SeaW. Aleutians

Kodiak

Gulf of ~Bering Sea

Ou tram and Haegele 1972

Pacijic Sand Lance / I9

Chukchi SeaBlack-leggedkittiwake

Norton Sound

S. Bering Sea%ulfof Alaska

Chukchi Sea

Norton Sound

E. Bering SeaKodiak

Common murre

Chukchi SeaThick-billed murre

Tufted puffin Swartz 1966

Vermeer 1979

Rhinoceros auklet Gulf of Alaska

British Columbia

Washington

Ancient murrelet Scaly 1975

Scaly 1975Sooty shearwater

Pelagic cormorant Swartz 1966

Robertson 1974

Robertson 1974Double-crested

cormorant

British ColumbiaRed-throated loon


Table 2. Marine seabird predators of sand lance AH!.

Chukchi Sea

W. Aleutians

Gulf of Alaska

British Columbia

Washington

Chukchi Sea

Norton Sound

Pribilof Islands

Gulf of Alaska

N. Gulf/S. Bering

British Columbia

British Columbia

Chukchi Sea

British Columbia

British Columbia

Swartz 1966; Springer andRoseneau 1979; Springer et al. 1984Drury et, al. 1981Hatch et al. 1978

Swartz 1966; Springer et al. 1984Drury et al, 1981Ogi and Tsnjita 1973Drury et al. 1981

Swartz 1966; Drury et al. 1981; Springer et al. 1984

Wilson 1977; Wilson et al. 1984

Swartz 1966

Drury et al. 1981Hunt et al. 19&1

Manual and Boersma 1978

Hatch et al, 1978

Leschner in Vermeer 1979

Vermeer 1979

Wilson l977; Wilson and Manuwal 19&6

Reimchen and Douglas 1984

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

E. Aleutians

W. Gulf of Alaska

Kodiak

Fur seal

Gulf of ~

OregonPitcher 1980

Brown and Mate 1983

Harbor seal

Stellar sea lion

Spotted seal

Minke whale

Sei whale

Humpback whale

Gulf of Alaska

E. Bering Sea

N. Pacific

Bering Sea


Taylor et al., in Sirnenstad et al. 1979Wilke and Kenyon 1957; Kajimura 1984Macy et aL 1978

Wilke and Kenyon 1952; Pitcher 1981

Lowry and Frost 1981

Nemoto 1959; Frost and Lowry 1981; Kajimura 1984

Frost and Lowry 1981

Huey, in S imenstad et al. 1979


Temperature and light also appear to be importantfactors. When water temperatures reach 20'C, AP bur-row in the sand and become dormant Nagoshi andSano 1979!. For adult AD, however, increased tem-perature is associated with growth Winters 1983!.White Sea sand lance AM! are reported to burythemselves in response to reduced light levels <0.1Iux! and picdators Girsa and Danilov 1978!.

In laboratory experiments, the activity of adult AM

was directly related to food availability, light intensity,and temperature Winslade �974a,b,c!. If food waspresent, activity was high during the day and low atnight when they mostly remained buried in the sand Winslade 1974a!. When abundant food was available,swimming activity was maximum at light intensitiesof 100 and 1000 lux, much reduced at 10 lux and very

low at I Iux; the threshold level light intensity at 50%maximum activity! was 20 lux Winslade 1974b!.Diurnal and seasonal variations in catch of sand lance

on the fishing grounds correlated with estimated lightintensity of about 100 lux at the bottom Winslade1974b!. Activity during daylight was high at 10 and15'C and much lower at 5'C WinsIade 1974c!. Thepercentage of the annual catch in April the first monthof the fishery! showed a positive correlation withtemperature in April, indicating that initial availabilityin the spring may be more related to temperature thanto food availability Winslade 1974c!. Burying in thesand at the end of the summer may be related to fatcontent, as well as to decreasing levels of foodavailability, light intensity and temperature Winslade1974c!. Older fish are reported to disappear from thecatches earlier in the summer, possibly due to reduced

requirements for growth and/or earlier accumuIation offat Winslade 1974c!.

2,5,3 Total Mortality. For a fished population ofadult AM, total annual mortality was estimated at 65-75% Z=l.2! Macer 1966, cited in Reay 1970!. Basedon age composiuon data from other stocks, Reay�970! suggested fished and unfished stocks have simi-lar mortality rates. Winters �983!, who investigatedAD mortality rates in Newfoundland between 1968 and1979, reported a steady decrease in mortality rates fromZ-values of over 1.0 to less than 0.5. Since there is no

fishery for sand lance in Newfoundland, these rates areconsidered to be natural inortality, The increase in sandlance survival was correlated with a decline in cod

stocks, known to be major predators on AD.

2.6 Laboratory Holding and Rearing

Spawning of adult sand lance in the laboratory has beenreported for AH by Pinto �984!, for AA bySmigielski et al. �984! and for AM by Winslade�971, 1974a,b,c!.

3 EGGS


The sand lance egg, which usualIy contains a singIeyellow oil globule, is nearly spherical, demersal andadhesive Reay 1970; Trurnble 1973!. According toPinto �984!, AH eggs from the northeastern Pacificrange from 0.8-1.22 mm, with a mean diameter of 1.00min. Mean diameters of egg of other Amiriodyresspecies are 0.66 mm for AP, 0.83 mm or 1.00 rnm forAA, 1.00 mm for AD and 1.02 mm for AM Williamset al. 1964; Inoue et al. 1967; Winslade 1971; Scott

1972a; and Smigielski et al. 1984!.


3.2.1 Duration of Egg Stage. In the laboratory,incubation time for AH eggs was approximately 24days at 9'C Pinto 1984!. Japanese sand lance AP!eggs hatched in 33 days at 6.2 C and 13 days at 15.7'C lnoue et al. 1967! and AP or AH eggs hatched inabout 22 days at 6.9' to 10'C Kitaguchi 1977!.According to Yamashita and Aoyama �985!, medianhatching time for AP eggs was 51 days at 6.5'C, 25days at 10.5 C and 20 days at 15.5'C. Averagehatching time for AA eggs was 62 days at 2'C and 30days at 10'C Smigielski et al. 1984!. Experiments onAP eggs Yamashita and Aoyama 1985!, AA eggs Smigielski et al. 1984! and AM eggs Winslade 1971!showed that incubation times were long and variableand that temperature and oxygen were importantfactors.

3.2.2 Large-Scale Patterns. No information on eggdistribution is available, but presumably eggs would befound in the saine general areas as adults since spawn-ing migrations are not known Reay 1970!.

3.2.3 Small-scale Patlerns. Nodataavailable.

3.2.4 Maximum and Mean Abundances. No dataavailable.

3.2.5 Vertical Distribution. Sand lance eggs aredemersal and adhesive, although they are occasionallycollected from the water column, probably as a resultof currents bringing the eggs off the bottom Williamset al 1964; Senta 1965!.

Paciji c Sand Lance I 23

3.3 Feeding

Not applicable.

3.4 Predation

3,4,1 Predator Species. Large numbers of late stageAD eggs were found in stomachs of the yellowtailflounder Limanda ferruginea! in the northwesternAtlantic Scott 1972a!.

3.4.2 Effect on Eggs. No data available.

3.5 Other Factors Affecting Eggs

3,5.1 Biotic, No data available.

3.5.2 Abiotic. According to Inoue et al. �967!, APhad a low hatch rate at 15.7'C and a maximum hatch

rate at 8.2'C, !n a laboratory study on AA, incubationtime and time to hatch completion increased withdecreasing temperatures �0'-2'C! Smigielski et al.1984!, The authors suggest that in the natural envi-ronment mechanical action may be a factor in reducingincubation times. Experiments on AM eggs indicatedhatching time may be affected by factors other thantemperature, e.g., incubation time and mortalityincreased with decreasing oxygen concentrations 9.5-4.0 ppm! Winslade 1971!. No eggs hatched at 2.1ppm; however, early stage embryos were able totolerate low oxygen concentrations for about a weekand to complete development if subsequently shifted tohigher concentrations. Winslade �971! suggested thatthe ability to retard development and survive periods oflow oxygen levels may be of particular adaptive value,since sand lance spawn on sandy bottoms and eggscould become temporarily buried in shifting sand.

3.5.3 Total Mortality. No data available.


Techniques for incubation of sand lance eggs aredescribed by Winslade �971!, Pinto �984! andSmigiclski et al. �984!.

4 LARVAE


Larvae of the five species of Ammodytes consideredherein have been described by the following: Pinto1984 AH!; Kobayashi 1961 AP!; Richards 1965 AA/AD!; Smigielski et al. 1984 AA!; Scott 1972a

AD!; Einarsson 1951, Macer 1966, Winslade 1971 AM!.

AH larvae hatched at 9 C were 5.3 mm in length,had pigmented eyes but did not have a complete gut orfunctional jaws Pinto 1984!. At the end of one week,the yolk sac was not completely absorbed and theintestines and jaws were still not completely developed.

AP larvae hatched at 6.5'C were 4.7 mm in length Yamashita and Aoyama 1985!. Some larvae beganfeeding within 2 days, and 50% of the larvae were feed-ing successfully within 5 days, Yolk-sac absorptionwas 95% complete by 6 days and 100% complete by12 days after hatching. Larvae were able to survivewithout food for a long time after hatching; time to50% mortality was approximately 11, 16 and 21 daysafter hatching at 15.5', 10.5' and 6.5'C, respectively.No clear "point of no return" was noted. The period ofrecoverable starvation from normal onset of feeding toirreversible starvation! was estimated as about 9 days.

AA larvae hatched at four different temperatureregimes �, 4, 7 and 10'C! and had a complete gut andfunctional mouth; some individuals in each of the four

groups began to feed within hours of hatching Smigielski et al. 1984!. At 7 C, 50% of larvae 6.5mm in length were feeding within 2 days. Yolk-sacabsorption took 3 days and oil globule absorption 7days. For AM larvae, yolk-sac and oil globuleabsorption took about 2 weeks at 7'C and slightly lesstime at 10'C Winslade 1971!, After 13 days, 16% oflarvae at 7'C and 58% at 10 C had begun to feed.However, newly hatched larvae were able to survive fora considerable length of time without food: time to50% mortality was 28 days at 7'C and 19 days at 10'C,

Yamashita and Aoyarna �985! determined growthrates of 0,12 mm in length and 4.2% dry weight perday for AP reared in the laboratory at 6.5'C. Growthrates for the two northwestern Aoantic sand lance have

been estimated from length frequency distributions asfollows: AA, 11.7 mm per month Norcross et al.1961! or 10.9 mm per month Smith et al. 1978!; AD,5.9 mm per month Scott 1972a!. Smigielski et al,�984! estimated growth rates for AA to range from2,4-5.62% dry weight per day at 2'C and 10'C, re-spectively, corresponding to length increases of 2.7mm and 11,3 mm per month for 155 days of growth.Buckley �984! estimated minimum growth rates toprevent starvation in AA larvae to be 2.4%, 2.5% and

3.4% change in protein content per day at 2, 4 and 7'C,respectively.



4.2.1 Duration of Larval Stages. AA laboratorystudies show that at 7 C yolk-sac absorption takesplace in 3 days at about 6.5 mm in length, oil globuleabsorption in 7 days at 7.2 mm and 50% first feedingin 2 days at 6.5 mm Smigielski et al, 1984!, Withincreasing temperature, size at yolk-sac absorptiondecreased but no differences in size at ail globuleabsorption or at first feeding were observed. Metamor-phosis to the juveniLe stage occurred at 29 mm inlength, which took 131 days at 4'C and 102 days at7'C. Schooling behavior was abserved at 25-30 mm 90 days at7 C!.

Laboratory-reared AM larvae completely absorbedyolk sacs and oil globules in 14-15 days at 7'C and in12 days at 10'C WinsIade 1971!. AM larvaecompleted metamorphosis at lengths between 30 mmand 40 mm Macer 1966!.

4.2.2 Large-scale Patterns, Sand lance larvae AH!generally occur in shallow less than 200 m! nearshoreareas of the Bering Sea and northeastern Pacific Ocean Trumble 1973; Macy et al. 1978; Rogers et al. 1980!.AH larvae were most abundant in early April in theBering Sea Waldmn and Vinter 1978! and March-Aprilin the Kodiak area Rogers et al. 1979; Kendall et al.1980!. In Washington, small larvae less than 10 mm!were reported from late January to early May in SkagitBay Blackburn 1973! and from mid-January to lateMarch in Puget Sound R, Trumble, Washington Dep.Fisheries, unpublished data!, Recently, hatched larvae AA/AD! that were most abundant in shallow nearshoreareas gradually dispersed offshore in the northwesternAtlantic Richards and Kendall 1973!.

No information is available on interannual varia-

bility in AH larval abundance. However, sand lancelarvae AA/AD! have shown very large interannualchanges in abundance approximately a 20-fold increasebetween 1974 and 1979! Smith et al. 1978; Shermanet al. 1981; Morse 1982!.

4.2.3 Small-scale Patterns. Altukhov �978! re-ported that AM larvae in the White Sea werc concen-trated in areas of "eddying currents."

4.2.4 Maximum and Mean Abundances. In the

eastern Bering Sea, sand lance AH! larvae were thethird most abundant species in bongo net samples�908/10 m ! despite their occurrence in less than 25%of the samples Waldran and Vinter 1978!. AH larvaewere also abundant in ichthyoplankton samples fromthe vicinity of Kadiak Island Dunn and Naplin 1974;Rogers et al. 1979; Kenda11 et al. 1980!. Larval AP

densities in Otsuchi Bay, Japan, averaged 105.6/100m3 in 1981-82 Yamashita et al. 1984b!.

In the Gearges Bank region, concentrations of sandlance larvae AA/AD! were over 100,000/10 m2 Smith et al. 1978!. Mean abundance in this regionranged from ]0 to 1018 pcr 10 m2 between 1974 and1979 and accounted for 55-98% of the larvae in winter

samples Sherman et al. 1981!.4.2.5 Vertical Distribution. In Kodiak Island

samples, AH larvae were most abundant at 10 and 30m depths during the day and were somewhat deeper atnight Rogers et al. 1979!. Inoue et al. �967! andYamashita et al. �985! reported similar results for APlarvae; they were 6-10 m below the surface during theday and deeper at night and also were more concentratedabove thermoclines. Yamashita et al. �985! reportedvertical migratian in larval AP as small as 5-6 mm,although larger larvae had a greater vertical range. AAlarvae also were found to be near the surface during theday and deeper at night Richards and Kendall 1973!.

4.3 Feeding

4.3.1 Habitat, See 4.2.5.!4.3.2 Prey Species. In the Strait of Georgia under

the Fraser River plume!, larval sand lance AH! lessthan 20 mm in length fed on prey less than 500 Ii indiameter, mainly copepod eggs and nauplii LeBrasseuret al. 1969!. Larger larvae �0-40 mm! fed on larger�.5-1.5 mm! zooplankton, primarily species ofMicrocalanus, Oithona and Pseudocalanus and naupliiof larger copepods. AP larvae fed mostly on smallcopepods, copepod nauplii and cladocerans Inoue et al.1967; Sekiguchi et al. 1974!.

According to Covill �959!, the primary prey itemsfor larval AA between 3,2 and 23.1 mm in length werccopepod nauplii and copepods. Phytoplankton, animportant component of the diet for small larvae lessthan 5 mm!, decreased in importance with increasingsize of larvae, Phytaplankton appeared to be mareimportant in winter Dec-Feb.! than in spring March-April! for the same size class of larvae. Ryland �964!noted that AM larvae fed only during daylight.Diatoms and dinaflagcllates were important items inthe diet of AM larvae less than 8 mm in length, andlarger larvae fed mostly on copepad nauplii andappendicularians.

4.3.3 Prey Density Requirements. Laboratory stud-ies of Winslade �971!, Buckley et al. �984! andYamashita and Aoyama �986! showed that newly


hatched AM, AA and AP larvae are capable of surviv-ing up ta 2 weeks without food see also 4.1!.

4.3.4 Factors Affecting Availability of Prey. Nodata available,

4.3.5 Temporal Patterns of Energy Storage. Natapplicable,

4.3,6 Evidence of Food Limitation. LeBrasseur et

al. �969! reported that over 50% of AH larvae lessthan 40 mm in length in the Strait of Georgia hadempty stomachs and suggested that concentrations ofmicrozooplanklon were limiting survival.

The smallest size group of AA larvae in Long IslandSound had the highest percentage �9%! of emptystomachs Covill 1959!. AA survival to metamor-phosis at food concentrations of 200, 500 and 1000ratifers/liter corresponding to 0,16, 0,40 and 0.80calorie/liter! was estimated ta be 0.12%, 5.75% and11.74%, respectively Buckley et al. 1984!. Sincethese estimates were based on a larval period of fixedlength �02 days!, slower growth at lower foodconcentrations would extend the larval period andincrease mortality Buckley et al. 1984!. Althoughsurvival rates at the above food concentrations are

comparable to those of other species, the caloricrequirements appear to be lower, indicating possibleadaptation to survival at low food concentrations Buckley et al. 1984!. Comparison of RNA-DNAratios in larval sand lance probably AA! collected fromthe northwestern Atlantic indicated that, in 1982, larvaewere generally in poorer condition than larvae collectedin 1981, and that some of the 1982 larvae wereapparently losing weight Buckley 1984!.

4.4 Predation

4.4.1 Predator Species, Marine fish, particularlyjuveniles, are major predators of larval AH, e.g., inBristol Bay, sockeye salmon juveniles Straty 1974!and sockeye aduIts Nishiyama 1974!; in nearshareKodiak, walleye pollock and juvenile pink salmon Rogers et al. 1979!; in Cook Inlet, sockeye and cohosalmon juveniles and stagharn sculpins Blackburn etal 1980!; in Chatam Sound, juvenile coho salmon Manzer 1969!; in the Strait of Georgia and SaanichInlet, juvenile salmonids, including coho, churn,sockeye and king salmon and steelhead trout, Salmogairdnerii, Barraclough 1967; Barraclough et al. 1968;Robinson et al. 1968!; in the Strait of 3uan de Fuca,juvenile coho and, to a lesser extent, juvenile chinookand herring Cross et al. 1978!. Arctic terns were a

major predator of larval AH in Alaska Bent, cited inAinley and Sanger 1979!. See also Tables 1-3.!

Predation by the hyperiid amphipod Pararhemisrojaponica on larval AP Yamashita et al. 1984a, b! wasestimated ta be an important source of morality. Inthe laboratory, both newly hatched and post-yolk sachrvae �3-15 days old; 6.0-6.3 mm standard length!were vulnerable to amph ipad predatian,4.4.2 Effect an Larvae. No data available.

4. S Other Factors Affecting Larvae

4.5.1 Biotic. No data available,

4.5.2 Abiatic. Inaue et al. �967! suggested thatwind is an important factor in larval dispersal. At lowtemperatures, growth is slower and time ta metamor-phosis is longer for AA larvae Smigielski et al.1984!. However, daily mortality rates of AA larvae inthe laboratory are not affected by temperatures between2' and 9 C Buckley et al. 1984!. AM larvae survivelonger without food at lower temperatures Winshde1971!.

4.5.3 Total Mortality, For larval sand lance AA/AD! in the northwestern Atlantic, instantaneous mor-tality rates between 1974 and 1980 ranged from .207-.363, corresponding ta a daily mortality of 6-10% forlarvae of 5-27 mrn Morse 1982!. In laboratoryexperiments, the daily instantaneous mortality rate forAA larvae between hatching and day 16 was 0.01 andwas unaffected by feeding level ar temperature Buckleyet al. 1984!. Mortality rates for older larvae �0-43days! ranged from 0.2-0.02 and decreased withincreasing food level.


Winslade �971! and Smigielski et al. �984! describetechniques for rearing larvae of AM and AA,respectively.

5 JUVENILES

5. 1 Genera! Description

At 7'C, metamorphosis to the AA juvenile phase takesplace at a length of about 29 mm, approximately 102days after hatching Smigielski et al. 1984!, Thejuvenile stage may last fram 1-3 years, depending onthe area see 2.1.1!.



5.2.1 Duration of Juvenile Stage. See 5.1.!5.2.2 Large-scale Patterns. Juveniles commonly co-

occur with adults, although they may also be present ininshore areas where adults are absent Richards et al,1963; Rcay 1970!. Reay �970! suggested that this isdue to the wide dispersal of juveniles rather than toseparate nursery areas. AH juveniles are abundantalong sandy beaches and in shallow nearshore areas ofthe Bering Sea Macy et al. 1978!, Norton Sound Barton 1978!, Kodiak Island Blackburn and Jackson1980! and Cook Inlet Blackbum et al. 1980!, Juve-niles �-age and age 1! exhibit an onshore migrationduring the summer in some areas, and there is someevidence that migration takes place earlier in the sum-mer for age I fish Barton 1978; Blackburn and Jackson1980; Blackburn ct al. 1980!. Large interannual vari-ations have been reported in the catch of 0-age AP inJapan Hamada 1966a, b, c; Inouc ct al 1967; Nagoshiand Sano 1979!.

5.2.3 Small-scale Patterns. Juvenile AH have been

observed schooling with similar-size juvenile herring Hobson 1985!. Richards �976! reported similarobservations for AA. AA schooling behavior begins at25-30 mm, about the time of metamorphosis to thejuvenile stage Smigielski et al. 1984!.

5.2,4 Maximum and Mean Abundances, No esti-

rnates of abundance are available,

5.2.5 Vertical Distribution. Juveniles may be morenumerous than adults in surface waters at night, basedon catches of AP Senta 1965b! and AM Macer 1966!.ln thc laboratory, burrowing behavior in AA juvenilesis first observed when they are between 35 and 40 mmin length Smigiclski ct al 1984!.

5.3 Feeding

5.3.1 Habitat. Juveniles feed almost exclusively inthe water column, but no information is available onfomging depth range.

5.3.2 Prey Species. Studies of the food habits ofjuvenile AH from the Chukchi Sea Springer ct al,1984!, western Aleutians Simenstad et al. 1978!,Kodiak Harris and Hartt 1977a, b; Rogers ct al. 1979!,Cook Inlet Blackburn et al. 1980! and the Strait ofGeorgia Barraclough 1967! agree in thc predominanceof copcpods in the diet. Other prey items of someimportance include other crustacean larvae, larvaceans,cladocerans, chaetognaths, fish larvae, mysids andgammarid amphipods. In the lower Cook Inlet area,

Blackburn ct al. �980! reported that harpacticoidcopepods were important in April, along with shrimpand fish larvae, while calanoid copepods were mostabundant the remainder of summer.

5,3,3 Prey Density Requirements. No data avail-able.

5.3.4 Factors Affecting Availability of Prey, Nodata avaihble,

5.3.5 Temporal Patterns of Energy Storage. Nodata available.

5.3.6 Evidence of Food Limitation, A high nega-tive correlation was found between the size of age 1 APand the abundance of adult fish, indicating possiblecompetition for food Hamada 1967!. Nagoshi andSano �979! reported a similar significant negativecorrelation bctv ccn growth and abundance in 0-age AP.Sekiguchi et al. �976! showed that 0-age AP do notbegin to accumulate fat reserves until attaining 45-50mm in length and that fat content increased withincreasing length; consequently, reduced growth couldaffect survival by reducing energy reserves during thedormancy period.

5.4 Predation

5,4.1 Predator Species. See Tables 1-3 and 4,4.1.!5.4.2 Effect on Juveniles. No data available.

5.5 Other Factors Affecting Juveniles

5.5.1 Biotic. No data available.

5.5.2 Abiotic. Winters �981! reported a significampositive relationship between water temperature andfirst year growth in AD.

5.5.3 Total Mortality. No data available.


Sce 2.6.!

6 CURRENT H YPO TIIESKSON FACTORS AFFECTINGYEAR-CLASS ABUNDANCE

Environmentrti effects: Hamada �966c! correlatedincreased catches of 0-age sand lance AP! with lowwater temperatures during the spawning season and alsowith the number of days with westerly winds duringthe three weeks following peak spawning,

Density-dependent effecrs on recruitment: A strongnegative correlation r = -.87! between the percentage ofage- I AP in thc catch and thc catch of 0-agc fish was


observed by Hamada �966a!. In addition, Hamada�967! reported a negative correlation r = -.81! betweenthc number of eggs spawned and the catch of 0-age AP.

Density-dependent sects on growth: Small size ofage-I AP was correlated with large catches of adult �-3years old! fish Hamada 1967!. Similarly, Nagoshi andSano �979! found a negative correlation r = -.86! be-tween growth of 0-age AP and an index of populationdensity.

l.ood availability: The availability of adequate foodhas frequently been suggested as an important factor inIarvaI mortality. Low food availability may lead toreduced growth and consequently a prolonged larvalphase and increased vulnerability to predation Buckley1984!. Laboratory studies on AA larvae provide esti-mates of minimum growth rates necessary to preventstarvation at a range of temperatures and a model pre-dicting recent larval growih based on RNA-DNA ratiosand temperature Buckley 1984; Buckley et al 1984!.Larvae probably AA! collected during 1981 and 1982indicate differences in larval condition RNA-DNAratio! that could be attributed to food availability B uckley 1984!. Although no direct. evidence was presentcd to indicate diffcrcnccs in larval survival, basedon RNA-DNA ratios, a higher perceniage of larvaewerc in poor condition in 1982, and some had appar-ently been losing weight Buckley 1984!.

Replacement and predation: Andersen and Ursin�978! presented the hypothesis that the reduction ofherring and mackereI stocks have resulted in theirreplacement in the North Sca ecosystem by small, fastgrowing opportunistic species such as sand Iance.However, thc lack of prior data on the abundance ofsand lance makes the hypothesis difficult to test unlessherring and mackerel stocks return to former levels ofabundance,

Sherman et al. �981! presented a similar hypothesisfor changes in the ecosystem structure of the north-western Atlantic. Larval surveys indicated a substantialincrease in the abundance of sand lance AA/AD! larvaebetween 1974 and 1979 which apparently reflects alarge increase in spawning stock bioinass Morse1982!. The authors suggest that this rapid increase insand lance population size was directly related to thereduction of mid-size predators, such as cod, haddock,hake, mackerel and herring.

Both sand lance AD! and capclin are important preyitems for Atlantic cod off Newfoundland Lilly 1982;Lilly and Fleming 1981; Winters 1983!. Winters�983! hypothesized that recent increases in sand lance

AD! abundance were attributable to decreases in codbiomass on the Newfoundland Grand Bank between1968 and 1979. The presence of greater percentages ofolder age groups indicated reduced mortality, and achange in the dominant age group from age 4 to age 3suggested increased recruitment. Using data froin re-search trawl surveys, Winters �983! found significantnegative correlations of sand lance abundance andrecruitment with cod biomass.

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ammodyfes hexapterus with notes on relapsed ammodytes …

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