D-A188 383 SPECIES PROFILES LIFE HISTORIES AND ENVIRONMENTAL t/1REQUIREMENTS OF COASTAL (U) VIRGINIA POLYTECHNIC INSTAND STATE UNIV BLACKSBURG DEPT OF F C i FAY ET AL

UNCLASSIFIED OCT 83 FWISOBS-82/11 9 F/G 613 NLlllEEEEllhliIIlllEEnlllllIImIIIIE

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OtbFWSIOBS-82111.9 jJE' -TR EL-82-4.7October 1983

Species Profiles: Life Histories andEnvironmental Requirements of Coastal Fishes

-v) and Invertebrates (Mid-Atlantic)

ALEWIFE/BLUEBACK HERRING MAY 2 198-1

Coastal Ecology GroupFish and Wildlife Service Waterways Experiment StationU.S. Department of the Interior U.S. Army Corps of Engineers

87 5 21 i

mFWS/OBS-82/11.9TR EL-82-4October 1983

Species Profiles: Life Histories and Environmental Requirementsof Coastal Fishes and Invertebrates (Mid-Atlantic)

ALEWIFE/BLUEBACK HERRING

by

Clemon W. Fay, Richard J. Neves, and Garland B. PardueDepartment of Fisheries and Wildlife Sciences

Virginia Polytechnic Institute and State UniversityBlacksburg, VA 24061

Project ManagerLarry Shanks

Project OfficerNorman Benson

National Coastal Ecosystems TeamU.S. Fish and Wildlife Service

1010 Gause BoulevardSlidell, LA 70458

This study was conductedin cooperation withCoastal Ecology Group

U.S. Army Corps of EngineersWaterways Experiment Station

Performed forNational Coastal Ecosystems TeamDivision of Biological Services

Fish and Wildlife ServiceU.S. Department of the Interior

Washington, DC 20240

CONVERSION FACTORS

Metric to U.S. Customary

Multiply AX To Obtain

millimeters (mm) 0.03937 inchescentimeters (cm) 0.3937 inchesmeters (m) 3.281 feetkilometers (km) 0.6214 miles

square meters (M2) 10.76 square feetsquare kilometers (km2) 0.3861 square mileshectares (ha) 2.471 acres

liters (1) 0.2642 gallonscubic meters (m3) 35.31 cubic feetcubic meters 0.0008110 acre-feet

milligrams (mg) 0.00003527 ouncesgrams (gm) 0.03527 ounceskilograms (kg) 2.205 poundsmetric tons (mt) 2205.0 poundsmetric tons (Nt) 1.102 short tonskilocalories (kcal) 3.968 BTU

Celsius degrees 1.8(C °) + 32 Fahrenheit degrees

U.S. Customary to Metric

inches 25.40 millimetersinches 2.54 centimetersfeet (ft) 0.3048 metersfathoms 1.829 metersmiles (mi) 1.609 kilometersnautical miles (nmi) 1.852 kilometers

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

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

ounces (oz) 28.35 gramspounds (lb) 0.4536 kilogramsshort tons (ton) 0.9072 metric tonsBTU 0.2520 kilocalories

Fahrenheit degrees 0.5556(F° - 32) Celsius degrees

CONTENTS

Page

CONVERSION TABLE ....... ........................... . . . . iiPREFACE ...... .... ... .... ...................................... ivACKNOWLEDGMENTS ............ ............................. v

PROFILE SCOPE•••............................ 1NOMENCLATURE/TAXONOMY/RANGE . . . .................... 2MORPHOLOGY/IDENTIFICATION AIDS ......... ...................... 2

Alewife. . ............. ............................. 2B1ueback Herring ............ ....................... 2Aids for Species Separation.......... ...................... 2

REASON FOR INCLUSION IN SERIES ...... ........................ 4LIFE HISTORY ........ ........... ..................... 4

Reproductive Physiology/Strategy ......... .................... 4Spawning . . . . . . . . . . . ............... . . ................. 5Eggs ....... ... .... ... ... ... ................................ 6Yolk-Sac Larvae ..... ... .. .... ................................. 7Larvae . . . . . . . . . . .. . .............. . . ................... 7Juveniles .......................................... 7Adults ... ............................ ..................... 9

GROWTH CHARACTERISTICS....... ................................. 9Growth Rates ......... ............... 9Length-Weight Relationships ...................... 12

_ THE FISHERY ...... .......................... 12Comercial Fisheries ......................... 12Recreational Fisheries ....... ... .......................... . 13Population Dynamics ...... .... ... .......................... 13

ECOLOGICAL ROLE ....... ... ... .. ............................. 14Food Habits ............... . . . . . ......................... 14Feeding Behavior ....... ... ..... ............................ 15Competitors .......... .............................. ... 16Predators . . . . . . . . . . . .... . . ........... ..... 16

ENVIRONiENTAL REQUIREMENTS .................................... 16Temperature ...... .... .................................... 16Salinity.. .......................... 18OhrEnvironmental*Factors "'-18Other EnirnretlFcos................... ... ~1Environmental Containants ..... .......... ......... . --. 19

LITERATURE CITED .... .......................... ....... 20

iii

PREFACE

This species profile is one of a series on coastal aquatic organisms,principally fish, of sport, commercial, or ecological importance. The profilesare designed to provide coastal managers, engineers, and biologists with a briefcomprehensive sketch of the biological characteristics and environmental require-nents of the species and to describe how populations of the species may beexpected to react to environmental changes caused by coastal development. Eachprofile has sections on taxonomy, life history, ecological role, environmentalrequirements, and economic importance, if applicable. A three-ring binder isused for this series so that new profiles can be added as they are prepared.This project is jointly planned and financed by the U.S. Army Corps of Engineersand the U.S. Fish and 'Jildlife Service.

Habitat Suitability Index (AISI) models are being prepared by the U.S. Fishand Wildlife Service for the alewife and blueback herring. HSI models aredesigned to provide a numerical index of the relative value of a given site asfish or wildlife habitat.

Suggestions or questions regarding this report should be directed to:

Information Transfer SpecialistNational Coastal Ecosystems TeamU.S. Fish and Wildlife ServiceNASA-Slidell Computer Complex1010 Gause Bc ,levardSlidell, LA 70458

or

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

This series should be referenced as follows:

U.S. Fish and Wildlife Service. 1983. Species profiles: life histories andenvironmental requirements of coastal fishes and invertebrates. U.S. Fishand Wildlife Service, Division of Biological Services, FWS/OBS-82/11.U.S. Army Corps of Engineers, TR EL-82-4.

This profile should be cited as follows:

Fay, C.W., R.J. Neves, and G.B. Pardue. 1983. Species profiles: life historiesand environmental requirements of coastal fishes and invertebrates (Mid-Atlantic) -- alewife/blueback herring. U.S. Fish and Wildlife Service,Division of Biological Services, FWS/OBS-82/11.9. U.S. Army Corps of @Engineers, TR EL-82-4. 25 pp.

iv

ACKNOWLEDGMENTS

We are grateful for the review by Dr. Joseph Loesch, Virginia Institute of MarineScience, Gloucester Point. We appreciate permission from the Canadian Journal ofFisheries and Aquatic Sciences for reprinting Figures 1 and 3. Permission is acknow-ledged from the Southeastern Association of Fish and Wildlife Agencies for reprintingFigure 4.

V

Figure 1. A: alewife; B: blueback herring.

ALEWIFE/BLUEBACK HERRING

PROFILE SCOPE spercies. When available, these differ-ences are addressed by separateThis profile covers life histcry statements for each species. In addi-

and environmental requirements of tion, a special section on the mostboth alewife (Alosa pseudoharenqus) readily distinguishing characteristicsand blueback herring (Alosa aesti- for separating eggs, larvae, andvalis), since their distribution is adults of the two clupeids is pre-overlapping and their morphology, sented in the morphology section ofecological role, and environmental this profile.requirements are similar. Neverthe- Because most of the information

, less, significant differences in certain available concerns the alewife, charac-' physical, physiological, and biological teristics of this species are given pri-"Q ', characteristics exist between the two ority reference in the text. Features

1i

should not be considered similar Alewifein the two species unless noted assuch. In a few studies (particularly Dorsal rays 12-19 (usuallycommercial fisheries statistics), the 13-14), anal rays 15-21 (usuallytwo species are referred to collectively 17-18), scales in lateral series 42-54.as "river herring" or "gaspereau" Prepelvic scutes 17-21 (usually(Canada). 19-20), postpelvic scutes 12-17 (usu-

ally 14-15), gill rakers on first arch38-46. Body strongly compressed,

NOMENCLATURE/TAXONOMY/RANGE deep. Mouth oblique, anterior end oflower jaw thick, heavy, and extendingto middle of orbit. Eye large, diametergreater than snout length. Color:

Scientific names..Alosa pseudoharengus dorsally grey to grey-green, laterally(Wilson)/Alosa aestivalis (Mitchill) silver with prominent dark shoulder

Preferred common names.. Alewife/blue- spot; fins pale, yellow or green.back herring (Figure 1)

Other common ,ldmes .... River herring,gaspereau, oldwife. B!ueback Herring

Clash ..................... OsteichthyesOrder .................... Clupeiformes Dorsal rays 15-20, anal raysFamily ....................... Clupeidae 15-21, scales in lateral series 46-54.

Prepelvic scutes 18-21, postpelvicscutes 12-16, gill rakers on first arch

Geographical range: The alewife is 41-52. Body moderately compressed,an anadromous species found in elongate, eye diameter small, equal toriverine, estuarine, and Atlantic or less than snout length. Upper jawcoastal habitats, depending on with definitive median notch, no teethlife cycle stage, from Newfound- on premaxillaries. Color: dorsallyland (Winters et al. 1973) to blue to blue-green, laterally silverSouth Carolina (Berry 1964). with prominent dark shoulder spot;Landlocked populations are in the fins pale, yellow or green.Great Lakes, Finger Lakes, andmany other freshwater lakes(Bigelow and Schroeder 1953; Aids for Species SeparationScott and Crossman 1973). Theblueback herring is an anadro- Eggs. Unfertilized blueback her-mous species found in riverine, ring eggs amber, alewife eggs green.estuarine, and Atlantic coastal Oil droplets of fertilized eggs unequalhabitats, depending on life stage and scattered for blueback herring,cycle, from Nova Scotia to the numerous and uniformly tiny for ale-St. Johns River, Florida (Hilde- wife (Kuntz and Radcliffe 1917; Nor-brand 1963) (see Figure 2 for a den 1967).map of the mid-Atlantic distribu-tion of alewives and blueback Larvae. Myomeres between inser-herring). tion of dorsal fin and anal vent 11-13

(mean 11.8) for blueback herring lar-vae, 7-9 (mean 8.0) for alewife larvae

MORPHOLOGY/IDENTIFICATION AIDS (this characteristic is definitiveaccording to Chambers et al. 1976).Larvae less than approximately 15mm

The following information was can be separated using regressions oftaken from summaries in Jones et al. vent to tail distance (mm) and vent to(1978), unless otherwise indicated. urostyle distance (mm), against

125.4 mm 1 inch.

2

N.N.Y.

BALTIMORE

WASHINGTONNN.

PHILADELLHS

KILOETER

BALTIMCARE HA TTERASt~

Figure 2. MdAlniOitibtoRfteaeieEn leakhrig

..... .3

WAHIGTN 00

standard length (SL); presented inChambers et al. (1976).

Adults. Externally, by scaleimbrication patterns and individualscale markings (Figure 3). Scalebaseline and dividing line coincidentalon alewife scales, not on bluebackherring scales (O'Neill 1980; MacLellanet al. 1981). Internally, peritoneallining uniformly dark in blueback her-ring; pale, grey, or silvery with darkpunctations in alewife (Leim and Scott ALEWIFE BLUEBACK HERRING1966; Scott and Crossman 1973).Shape of otolith distinctive (Scott and approx. scale.' 'cmCrossman 1973) and differencesdescribed in Price (1978) and illus-trated in O'Neill (1980). Biochemically / -.distinguished by muscle myogen and 7

retina LDH enzyme migration patterns(elect rophores is) (McKenzie 1973,1975).

Alewives possess fewer verte-brae, dorsal rays, anal rays, and gillrakers on the first arch, in general, W-MW- ,,tJcompared with blueback herring. Eye IT II,, P8 ftw i

diameter to snout length ratios are " S=UI Ws"

also significantly different between the approx. scale 1 1 1... 1 5 mm

two clupeids, but mean values are Figure 3. Scale imbrication patternsclose and ranges overlap (Messieh (top) 3. Sc al catio phology1977). The applicability of these (top) and individual scale morphologymeristic characters for absolute spe- (bottom) used for external discrimina-cies determination is limited since tion of the alewife and blueback her-there is at least some overlap between ring (from MacLellan et al. 1981; withspecies for each characteristic, permission of the Canadian Journal ofFisheries and Aquatic Sciences).

Though dorsal coloration hasbeen cited as species-distinctive in important links in estuarine andfresh specimens (Bigelow and Schroe- marine food webs, between zooplank-der 1953), this character may not be ton and top piscivores. Commercially,reliable. MacLellan et al. (1981) both species have recently (last twofoL d no significant or detectable dif- decades) gained in recognition andference in dorsal coloration and interest as sources of fish meal, fishobserved that such coloration oil, and fish protein, particularly forappeared to vary substantially with the animal food industries.ambient lighting conditions.

LIFE HISTORYREASON FOR INCLUSION IN SERIES

Reprod uctive Ph y siolog y/StrategyThe alewife and blueback herring

are important ecologically and to a Alewives and blueback herringlesser extent as commercial tish spe- are heterosexual, though hermaphro- .cies. Ecologically, these species are ditism in landlocked populations of "V.

4

.7

alewives from Lake Michigan has been (310-mm female) (Loesch and Lundreported (Edsall and Saxon 1968; Hla- 1977). From 10% to 30(o) ofvek and Norden 1977). Females of the initial number of eggs present in, both species are slightly larger and a female blueb.ck herring remainedheavier than males of the same age after spawning. Left ovaries wereCooper 1961; Netzel and Stanek 1966; significantly heavier than right ova-

Marcy 1969; Loesch and Lund 1977). ries, though left ovaries did not con-tain more eggs per gram of ovary

Some male anadromous alewives, than did right ovaries (Loesch andand a slightly smaller percentage of Lund 1977; Loesch 1981). Chesapeakefemales, spawn for the first time at Bay alewives ranged in fecundity fromage 3. Most fish of both sexes have 60,000 to 100,000 eggs per femalespawned once by age 4, and all by (Foerster and Goodbred 1978). Theage 5. Generally, males dominate age fecundity-to-age relationship forclasses 3 to 5 on the spawning Georgia populations of blueback her-grounds, while females dominate age ring did not fit a linear relationshipclasses 7 and older. Blueback her- well (r 2 = 0.42) (Street 1969). It wasring vary more than alewives in age suggested that the age-fecundity rela-of first spawning, though in general tionship is asymptotic for both ale-maturation rates are similar for the wives and blueback herring, and thaitwo clupeids (Joseph and Davis 1965; "fecundal senility" may occur in allLoesch and Lund 1977; O'Neill 1980). long-lived stocks of these species

(Street 1969; Loesch and Lund 1977).Age of first spawning, percent-

age of repeat spawners, and longevity Spawningin populations seem to decrease as oneproceeds from north to south. Anadromous alewives and blue-Spawning populations of alewives in back herring spawn once a year, dur-southern North Carolina comprised ing spring or early summer, in fresh, primarily 3-year-old fish, and no fish or brackish water (Raney and Mass-over 4 years old were found (Tyus mann 1953). Males arrive at mouths1974). In contrast, alewife spawning of spawning rivers earlier thanpopulations in Chesapeake Bay were females (Cooper 1961; Tyus 1971;represented by ages 3 to 8 (Joseph Richkus 1974a). Spawning environ-and Davis 1965), Connecticut River ments vary from streams only a fewstocks by ages 3 to 8 (Marcy 1969; meters (yards) wide and a few centi-Loesch and Lund 1977), and Nova meters (inches) deep to large riversScotian stocks (both alewives and such as the Delaware, Susquehanna,blueback herring) by ages 4-10 and Potomac (Mansueti 1956). Ponds,(O'Neill 1980). Percentage of repeat incluuing barrier beach ponds, withspawners was 60°0 for alewives in Nova an open outlet to the sea, are alsoScotia (O'Neill 1980), 61% in the York used by alewives (Bigelow and WelshRiver, Virginia (Joseph and Davis 1925). Jones et al. (1978) cited1965), and less than 10%o in southern Loesch (1968, 1969) as stating thatNorth Carolina (Tyus 1974). Blue- blueback herring do not ascend intoback herring runs consisted of 65% freshwater as far as alewives onand 75% repeat spawners in the York spawning runs. However, LoeschRiver and Nova Scotian waters, (personal communication) said therespectively (Joseph and Davis 1965; statement pertained to the existingO'Neill 1980). literature at that time, and findings

reported in Loesch and Lund (1977)Fecundity of Connecticut River indicated that upstream distribution

lueback-herring ranged from 45,80- was a function of finding appropriateeggs (238-mm female) to 349,70() eggs spawning habitats.

5

In laboratory tests, adult ana- Blueback herring prefer spawningdromous alewives from Rhode Island sites with fast currents and associatedwaters were capable of distinguishing hard substrates (Loesch and Lundwater of their natal pond from water 1977). Brackish water or standing A Atcollected in nearby ponds (Thunberg water habitats are rarely used. In1971). Olfaction was shown to be the contrast, alewives select a wide vari-major sensory mechanism for homing ety of spawning sites, using standingbehavior. Discriminate function analy- water and oxbows as well as mid-riversis, however, by Messieh (1977) on sites (Kissil 1974). Several studiesSt. Johns River, Florida, spawning have described alewife spawning instocks indicated considerable straying ponds with an open connection tofrom home streams, particularly the ocean (e.g., Havey 1973; Kissilbetween adjacent spawning areas/ 1974), but no observations of pondstocks. Messieh (1977) hypothesized spawning by blueback herring arethat the majority of stock mixing documented. Apparently a consider-occurred during the prespawning able separation, both spatially andperiod (late winter, early spring) temporally, exists for spawning activ-rather than "impulsively" on the ity of anadromous alewives and blue-actual spawning runs. The majority back herring.of spawning alewives in Lake Matta-muskeet, North Carolina, used only Loesch and Lund (1977)one of four available canals for described spawning behavior of blue-spawning migrations (Tyus 1974). It back herring. A spawning groupwas hypothesized that since this canal composed of one female and severalwas the only one available historically males swam in circles for several min-(built in 1907), the alewives may be utes, and males occasionally nudgedexhibiting homing behavior. Tyus the vent of the female. Swimming(1974) did not report, however, that speed increased gradualiy until a deepthis canal was also the shortest and dive occurred with subsequent releasemost direct from bay to lake. of eggs and milt simultaneously, very

near the substrate. Spawning activi-ties of both species occur diurnallyand nocturnally, though the greatestactivity apparently is nocturnal (Gra-

Spawning periods along the ham 1956, Edsall 1964). Both malesAtiantic coast range from late March and females migrate rapidly down-through July, occurring later in the stream after spawning, and totalnorth. Within the mid-Atlantic region, spawning time for a single migratorynearly all alewife and blueback her- group is usually 5 days or lessring spawn from April through mid- (Cooper 1961; Loesch and Lund 1977).July (Hildebrand 1963; Kissil 1969;Loesch 1969; Smith 1971; Tyus 1974;Loesch and Lund 1977). In general, Eggsalewives begin to spawn 3 to 4 weeksbefore blueback herring in sympatric Until water-hardened, eggs ofareas. Spawning peaks are 2 to 3 both species are demersal in stillweeks apart (Jones et al. 1978). Ale- water and adhesive or pelagic inwives began spawning at minimum running water (Loesch and Lund 1977;water temperatures of 10.5'C (510F) Jones et al. 1978). After' water-(Cianci 1969) and blueback herring at hardening (less than 24 hr), eggs lose14'C (57°F) (Loesch and Lund 1977). their adhesive property and enter theBoth species cease spawning when water column, Fertilized, water-hard-water temperature exceeds 27°C (810 ened eggs are green (alewife) toF) (Loesch 1969; Edsall 1970). amber (blueback herring) and contain

6

scattered, unequal (alewife) or Alosa spp. larvae, primarilynumerous, small (blueback herring) oil those of blueback herring and ale-droplets (Kuntz and Radcliffe 1917; wives, were present throughout upperNorden 1967). Egg diameter ranges Chesapeake Bay from hatchingfrom 0.80 to 1.27 mm for alewives and (approximately April 15) through Junefrom 0.87 to 1.11 mm for blueback (Dovel 1971). Larvae exhibited slightherring (Mansueti 1962; Norden 1967). downstream movement from presumedIncubation time is approximately 80 to spawning areas in the bay, and were94 hr at 200 to 21C (680 to 700 F) collected only in areas with salinitiesand 55 to 58 hr at 220 to 240 C(72* less than 12 ppt. Alosa spp. larvaeto 750F) for blueback herring eggs in Nova Scotian rivers occurred in(Cianci 1969; Morgan and Prince areas that were relatively shallow (<21976). Comparative values for ale- m, <6.6 ft), sandy, and warm, andwives are approximately 360 hr' at were collected in or near areas of7.2 0 C (45*F) (Edsall 1970), 178 hr spawning adults (O'Neill 1980).at 12.7C (55*F) (Kellogg 1982), 89hr at 21.10C (70*F) (Edsall 1970), Juveniles72 hr at 23.8 0C (750 F) (Kellogg1982), and 50 hr at 28.90C (84 0 F) Transformation to the juvenile

~(Edsall 1970). An equation for pre-dicting incubation time for alewife stage is gradual, but is completed ategg fromntempaturtie fodsalewie 1approximately 20 mm TL. Scales firsteggs from temperature (Edsall 1970) appear on juveniles between 25 and 29

mm TL, and are fully developed at 45

T = 6.335 x 106 'x t mm TL (Hildebrand 1963; Norden1967).

where T = time in days and t = incu-bation temperature in degrees F. Juvenile blueback herring in the

S LJames River, Virginia, exhibited a netYolk-Sac Larvae upstream movement between June and

October, presumably caused by con-Yolk-sac larvae range from 2.5 to tributions of juveniles from oxbows,

5.0 mm total length (TL) at hatching, side channels, and tributaries, whichand average 5.1 mm TL at yolk-sac gradually moved down into the main

absorption (Mansueti 1962; Norden river through the summer. Densities

1967). Duration of this stage is 2 to 5 of juveniles were significantly higher

days for alewife and 2 to 3 days for na the sure thaniat i gherblueack errig (ansuti 162;near the surface than at 5-m (16.4-ft)

blueback herring (Mansueti 1962; depth throughout their residence inCianci 1969). the river (Burbidge 1974).

LarvaeWarinner et al. (1969) studied

The larval stage lasts from the distribution of juvenile alewivesyolk-sac absorption until transforma- and blueback herring in the Potomaction to the juvenile stage. Larval River over the first 6 months of lifeblueback herring range from 4.0 to (Figure 4). Four important conclu-15.9 mm SL and larval alewives from sions were evident.4.3 to 19.9 mm SL (Jones et al.1978). Jones et al. (1978) provided 1) Both species exhibited appar-detailed drawings of the developmental ent upstream movement, averag-stages of eggs, yolk-sac larvae, and ing 24 km (15 mi) over 4 months,larvae of both alewives and blueback until the inception of emigrationherring, in October.

7

7 U NLEE, E I I Significant diel movements ofJUNE -juvenile alewives and blueback herringLii ___ in Virginia rivers were found (Loeschr n -I J et al. 1982a). Fish moved toward the

0 ,bottom during the day and toward ther5 JULY surface at night. Blueback herring

"00 were more sensitive to the sky-opacity21 5 index and exhibited more extensive

o 0 ,0 vertical movement patterns in relation• 8 T7 AUG- UST to changing light intensity than ale-

25 J " n 5 wives. This movement pattern wasC- - 0 " " ,' ,o a also noted for juvenile alosids in the

SEPTMBR" Potomac River (Warinner et al. 1969).SC- 9~0'TCM r!0h

- - In most Atlantic coast popula-,, tions, juvenile alewives and blueback

0 CCE. o herring emigrate from freshwater/es-I S .I. -,'.....Y I tuarine nursery areas between June

25J - Y _ F; and November of their first year of1 _1 __ _ --- ri 1 0life (Burbidge 1974; Kissil 1974; Rich-

STT!CN-,IVER ILE kus 1975; O'Neill 1980) . Juvenileriver herring in upper Chesapeake

Figure 4. Seasonal distribution of Bay did not emigrate until earlyjuvenile river herring by river mile spring of their second year, and suchip the Potomac River, Maryland. No migrations were comparatively rapiddata are shown for November because once they began (Dovel 1971). Thisfish were absent in the study area was the only report of nonmigratory(from Warinner et al. 1969). first-year juveniles, except for the

dwarf, nonmigratory population ofalewives described by Foerster andGoodbred (1978) from the Susque-

2) Both species were virtually hanna River.absent in the study area byNovember.

Stimulatory variables influencing3) Juvenile blueback herring out- the initiation of migratory "waves"numbered alewives 19:1 overall. (Richkus 1975) of juvenile alewives

from nursery habitats include heavy4) Juvenile alewives were most rainfall (Cooper 1961), high waterabundant in surface waters (Kissil 1974; Richkus 1975), andthrough September, and sharp declines in water temperatureincreased in abundance at 4.6 m (Richkus 1975). Richkus (1975)(15 ft) and on the bottom in observed that (1) such waves lasted 2September and October, prior to to 3 days, regardless of length ofemigration. In contrast, juvenile time of environmental change; (2)blueback herring maintained high migrations peaked in late afternoon;abundance in surface waters and (3) the magnitude of a migratorythrough October, increased in wave was not related to the magnitudeabundance at 4.6 m in November, of environmental change. Most (600 toand were never collected in bot- 80%) juveniles emigrated on only atom trawls throughout the study small percentage (7% to 8%) of theperiod, available days (Richkus 1975).

8

Significant numbers of juvenile abundant at depths between 56 andalewives and blueback herring were 110 m (184 and 361 ft), while blue-captured during winter in the Mullica back herring were most abundantRiver Estuary, New Jersey, indicating between 27 and 55 m (89 and 180 ft).the importance of this area for over- In summer and fall, catches of bothwintering (Milstein 1981). Blueback species were confined to the samplingherring was third in percent repre- area north of 40' north latitude, insentation and alewives seventh among three general areas: Nantucket53 species collected. Both species Shoals, Georges Bank, and the perim-were captured out to 8 km (5 mi) off- eter of the Gulf of Maine. Wintershore, which is near the outer limit of catches were between 400 and 43'salinity/temperature influences from north latitude. Spring catches werethe Mullica River Estuary. Overwin- distributed over the entire Continentaltering fish chose temperatures Shelf in the study region (Nevesbetween 4.5°and 6.5°C (40' -,nd 43.7" 1981).F) and salinities from 29 to 32 ppt.Densities of overwintering juveniles Adult alewives and blueback her-increased steadily from December to ring chose only a small portion ofMarch, then declined in April. The Georges Bank, specifically the westernalewife-to-blueback herring ratio in all slope at 41', 29' north latitude and 680,samples combined was 1:5 (Milstein 34' west longitude, as an area of1981). residence in July and October, 1964

(only months sampled) (Netzel andAdults Stanek 1966). Alewives outnumbered

blueback herring in these samples.Little information is available All mature age classes were repre-

concerning the life history or biology sented, but age 0", 1", and 2+ fishof alewife and blueback herring stocks were not captured.once the juveniles emigrate to the seae at age 0+ or 1+. A literature search Alewives and blueback herring,indicated that much research needs to like other clupeids, may exhibit sea-be conducted in stock identification, sonal movements in conjunction withoffshore exploitation rates, foreign preferred isotherms (or preferredfishing operations, and general life isotherms of forage organisms) (Col-history, movement, migration pat- lins 1952; Leggett and Whitney 1972);terns, feeding behavior, and ecology however, direct evidence is lackingof these clupeids in offshore waters. (Richkus 1974b). Feeding and vertical

migration are probably controlled byNeves (1981) summarized 16 years light intensity patterns within thermal

of catch data from National Marine preference zones (Richkus and WinnFisheries Service trawl surveys con- 1979; Neves 1981).ducted along the Atlantic coast,between Cape Hatteras and Nova Sco-tia. Samples were taken out to depthsof 200 m (656 ft). The majority of GROWTH CHARACTERISTICSthe catch of alewives and bluebackherring was taken at sampling stationswhere water depth was less than 100 Growth Ratesm (328 ft). Alewives and blueback --herring collected ranged from 60 to Growth rates (L in mm, wt in g)350 mm fork length (FL), and alewives of young-of-the-year blueback herringoutnumbered blueback herring in the James River, Virginia, areapproximately 10:1 for all samples given below. Blueback herringcombined. Statistical analysis deter- achieved a mean fork length of 35.6

A mined that alewives were most mm and weight of 3.68 g by 15

9

November of their first year (Bur- age were collected (Marcy 1969). Agebidge 1974). Growth rates of 2 alewives and blueback herring inyoung-of-the-year blueback herring in Albemarle Sound, North Carolina,the Cape Fear River, North Carolina, reached 153 mm FL and 148 mm FL,were similar to those for the James respectively, by the end of theirRiver population (Davis and Cheek third summer (Kornegay 1978). On1966). Georges Bank, age 2 fish of both

species reached approximately 180 mmChange in L Change in wt TL by the end of their third summer

Period per day (mm) per day (g) (Netzel and Stanek 1966).

Mid-June tomid-July 0.29 0.03 Lengths at age for sexually

Mid-July to mature alewives and blueback herringmid-Aug. 0.12 0.01 are given in Table 1. In general,

Mid-Aug. to alewives are longer than bluebackmid-Sept. 0.15 0.01 herring of the same age. Within each

Mid-Sept.to species males are smaller than femalesmid-Oct. 0.29 0.05 of the same age, and growth rates for

Mid-Oct. to both species level off after reachingmid-Nov. 0.32 0.05 sexual maturity (compared to growth

rates of immature fish). Mean weightsof spawning alewives in Damariscotta

Young-of-the-year alewives ap- Lake, Maine, ranged from 153 g (5.4faster than blueback oz) (males) and 164 g (5.8 oz)paretly row(females) at age 3, to 325 g (11.5 oz)

herring. In the Northeast Cape Fear (males) and 356 g (12.6 oz) (females)

River, North Carolina, young alewives a age 7. O 8-yearold femalereached a mean fork length of 44.0 at age 7. One 8-year-old femalereahed a7. m n gt of 44 weighed 455 9 (16.0 oz) (Waltonmm and 47.8 mm in August of 1964 1979).

and 1965, respectively (Davis andCheek 1966). Certainly some of theapparently faster growth of young Otolith and scale-aging tech-alewives is due to an earlier spawning niques for alewives and blueback her-period and therefore a longer growing ring from Albemarle Sound, Northseason. Growth of young alewives Carolina, agreed nearly 100% for agebetween hatching and fall emigration classes 0, 1, and 2, but consistencyfrom nursery areas averaged 102 mm between methods decreased progres-TL in lower Chesapeake Bay (Joseph sively as fish age exceeded 3 yearsand Davis 1965), and 113 mm TL in (Kornegay 1978). The scale-agingthe Connecticut River (Marcy 1969). technique tended to underestimate theAverage length of juvenile emigrants, proportion of age 4 and 5 individualssampled daily over three seasons and overestimate the proportion of age(1970-72) in Hamilton Reservoir, 6 and 7 individuals for both species.Rhode Island, ranged from 25 mm SL Growth rates backcalculated fromto 88 mm SL (30 mm TL to 105 mm scales tended to be higher than thoseTL) (Richkus 1975). backcalculated from otoliths. Regres-

sions for prediction of fork lengthLittle information is available on from scale and otolith measurements

growth rates of these clupeids are available in Kornegay (1978).between age 0+ and the time of firstspawning. Age 1 alewives reached Messieh (1977) gave von Berta-147 mm TL by the end of their second lanffy growth equations for alewivessummer in the Connecticut River and blueback herring in the St. JohnEstuary; however, only males of that River, New Brunswick, as follows:

1010 'U'r

Table 1. Length (mm FL) at age (yr) for selected Atlantic coast spawning popula-tions of alewives and blueback herring.

Length (mm FL) at age (yr)Locationa Species/sexb 3 4 5 6 7 8 9

AS A/M 236 249 256 259 268 279 -

AS A/F 254 261 270 277 283 287 -

AS B/M 231 241 245 251 258 270 -

AS B/F 245 252 258 264 268 280 307

CB A/M 229 239 249 254 259 - -

CB A/F 239 249 259 264 274 282 -

CR A/M - 265 278 290 301 - -

CR A/F - 284 284 299 308 324 -

CR B/M 258 266 280 286 298 - -

CR B/F 261 277 291 301 311 - -

GBc A/MF 270 284 294 306 316 327 330

GBc B/MF 240 269 281 292 302 313 -

DL A/M 260 278 301 315 318 - -

DL A/F 273 283 309 324 333 356 -

aAS = Albemarle Sound, North Carolina (Pate 1974); CB = Chesapeake Bay (Joseph

and Davis 1965); CR = Connecticut River (Marcy 1969); GB = Georges Bank (Netzeland Stanek 1966); DL = Damariscotta Lake, Maine (Walton 1979).

bA/M = alewife males; A/F = alewife females; A/MF = alewife males and females;

B/M = blueback herring males; B/F = blueback herr.ing females.

cValues in mm TL.

= 11

MA L = 291. 67(1 - e-0. 4 4 1 (t - 0.142)) plankton populations, resulted in the

F) observed deterioration in the condition

FA L = 310.48(1 - e - 0.4 0 0 (t - 0.103)) factor (Burbidge 1974).

MB L = 231.33(1 - e-0"590(t - 0.338) ) THE FISHERY

FB L = 259.85(1 - e0. 4 69 (t - 0.283))Commercial Fisheries

where MA is for male alewives, FA is

for female alewives, MB is for male U. S. commercial landings ofblueback herring, FB is for female river herring (both species combined)blueback herring, L = fork length along the Atlantic coast were 4,948 mt(mm) at time t in years, and e is the in 1980 and 3,754 mt in 1981. Thesebase of natural logarithms, landings were worth $779,000 and

$671,000, respectively. The currentAlso given were von Bertalanffy 5-year running average of river her-growth equation parameters and modi- ring landings (1977-81) by U. S.fied Walford plots for four separate fisheries was 5,003 mt/yr. More thanareas in the St. John River drainage, 90% of the U. S. commercial catchwhere different spawning stocks were occurred within 4.8 km (3 mi) of thesuspected. The equations given above coast (National Marine Fisheries Ser-are for all study areas combined, vice, NMFS, 1982). They are commer-since differences in growth parameters cially fished during spawning runs.among areas were small (Messieh Pound nets are the most commonly1977). used gear (Joseph and Davis 1965;

Pate 1974). The majority of U. S.Length-Weight Relationships landings was used for fish mc-al and

fish oil to be added to fertilizer, petLength-weight relationships for food, and domestic animal feed. A

alewives and blueback herring of the minor portion was used for fishingSt. John River, New Brunswick, were bait, and the remainder was soldgiven in Messieh (1977) as follows: salted or fresh for human consump-

tion. Roe from these species iscanned and is highly valued (Joseph

Male alewives logW=3.2351ogL-5.420 and Davis 1965; Street and DavisFemale alewives logW=3.1921ogL-5.294 1976; Merriner 1978).Male blueback logW=2.9041ogL-4.702Female blueback logW=2.472logL-3.693 The total foreign catch of riverL=Fork length in mm, W=Weight in g herring within the U. S. Fishery

Conservation Zone (FCZ) was 24.6 mtin 1980 and 13.9 mt in 1981. Cuba

Condition factors (K) for took greater than 90% of the 1980 for-young-of-the-year blueback herring of eign landings, and Poland capturedthe James River, Virginia, averaged approximately 75% of the 1981 foreign1.60, 1.68, 1.57, 1.37, 1.22, and 1.18 catch. All of the 1980 and 1981 for-for the months of June, July, August, eign landings were taken from theSeptember, October, and November, North Atlantic region (north of Caperespectively (Burbidge 1974). It was Hatteras) (NMFS 1982).hypothesized that a massive flood inthe study area between August and In domestic commercial fisheriesSeptember may have reduced zoo- in Albemarle Sound, North Carolina,plankton availability which, in addition surveyed in 1972, age 4 and 5 fishto normal seasonal declines in zoo- represented 60% or more of the

12

inshore catch of alewives and blueback history, biology, and management ofherring (Pate 1974). Significant con- Virginia and North Carolina stocks oftributions also occurred from age 3, anadromous alewives and blueback6, and 7 fish (at least 30% of total herring were reviewed by Rulifsoncatch). Ratio of alewives to blueback and Huish (1982). In addition, recom-herring in the 1972 catch was 2:3, mendations for development of a man-and sex ratios within species were agement plan are presented and dis-near 1:1. First-time spawners ac- cussed.counted for 50% and 57%, respec-tively, of the landings of blueback Recreational Fisheriesherring and alewives. Historically,the Chowan River has been the most Recreational fishing for alewivesimportant inshore fishing grounds in and blueback herring is significantAlbemarle Sound. Peak landings usu- during spring spawning runs in areasally occur in late April, coinciding such as Delaware Bay, Chesapeakewith the spawning runs of the more Bay, and Albemarle Sound (Patecommon blueback herring in North 1974). The disposition of the catch,Carolina (Pate 1974). In the Potomac however, is not well documented.River, Maryland, the 1974 commercial Apparently, most of the recreationalcatch was dominated by alewives in catch serves as bait for other sport-March and early April, while blueback fish (NMFS 1980). The numbers andherring landings peaked in late April weights of the recreational catch forand May. The total ratio of alewives each species are unknown, becauseto blueback herring in the 1974 catch published surveys lump these twowas 1:4 (Merriner 1978). species with menhadens, shads, other

herrings, and sardines. The NMFSTotal landings and catch per unit estimated that 6,169,000 total "her-

effort in North Carolina and Virginia rings" were captured by Atlantic coastwaters have declined substantially recreational fishermen in 1979.O over the last decade since peaking in

10 1969 at 35,302 rnt (Street and Davis Population Dynamics1976; Merriner 1978). The declinewas due to offshore trawl fisheries, Sex ratios/age structure. Onwhich did not begin operating until spawning runs, total sex ratios of1967 (McCoy 1975). These fisheries adults are nearly 1:1 in most areas.were not and are not size selective, Percentage of male alewives in theand 65% of the 1975 offshore trawl spawning populations of Bride Lake,landings of river herring consisted of Connecticut, was 55.6% (Kissil 1974),immature fish (Street and Davis 1976). compared to 53.8% and 53.0% for ale-In contrast, all inshore commercial wives and blueback herring in thecatches historically were captured Connecticut and Thames Rivers (Marcyfrom sexually mature populations, 1969), and 58.0% in a later study onwhere escapement rates can be con- Thames River blueback herringtrolled to prevent overexploitation (Loesch and Lund 1977). It has been(McCoy 1975). suggested that the slight dominance of

males in spawning populations is dueThe State of Maine has developed to their earlier sexual maturity com-

an Alewife Management Plan for ana- pared to that of females (Kissil 1974).dromous stocks within their State Sex ratios (% males) by age for 1966jurisdiction (Walton et al. 1976). His- and 1967 spawning populations of ale-torical landings and management are wives from the Connecticut andreviewed and management recommenda- Thames Rivers, Connecticut, (datations presented for the alewife runs of combined) were 72.3%, 63.7%, 49.7%,each coastal county. Aspects of life 33.8%, and 0.0% for fish of age 4, 5,

13

6, 7, and 8, respectively. Ratios for 67% over the 6-year study periodblueback herring were 80.0%, 79.4%, (Havey 1961). Results from these two64.5%, 36.9%, and 22.6% for fish of studies in Maine and the study byage 3, 4, 5, 6, and 7, respectively Kissil (1974) in Connecticut indicated(Marcy 1969). that juvenile production and adult

freshwater mortality may vary consid-Life stage abundance/reproductive erably among spawning areas and

and mortality rates. In Bride Lake, among different years in the sameConnecticut, an estimated 184,1 1 spawning area.adult alewives spawned 2.05 x 10eggs in 1966. Subsequently, 257,000 Methods of determining truejuveniles were counted as they emi- spawning population size of alewives,grated during the summer and fall. by subsampling at various timesThis generated a total freshwater on fishways on the Parker River,mortality rate from egg stage to emi- Massachusetts, were investigated bygration of 99.9987%, and indicated that Rideout et al. (1979). Visual counts2.88 juveniles left the lake for each during one 10-min period each houradult female that spawned. Combined produced estimates (by computer pro-with repeat spawning proportions, this gram) with less than 50 error. Visuallevel of juvenile production seemed counts during one 5-min period eachadequate for sustaining the spawning hour or one 10-min period each 1.5population in Bride Lake (Kissil 1974). hours were significantly higher inTotal freshwater mortality rate of magnitude of potential error (Rideoutspawning adults in Bride Lake was et al. 1979).reported at 57.4% and 48.6%, in 1966and 1967, respectively. Stock identification. Although

Thunberg (1971) found that alewivesHavey (1973) investigated juve- were capable of homing behavior

nile production and adult mortality of through olfaction, Messieh (1977) con-the alewife population in Love Lake, cluded that considerable mixingMaine. The number of emigrants between presumed spawning stocksranged from 220 to 439,062 fish over occurred on the spawning runs.an 11-year study period. Juvenile Though a majority of the alewivesemigrants produced per female per exhibited homing, a substantial num-year varied from 12 to 3,209, and ber (by meristic comparisons) frombiomass production of emigrants each presumed stock did not homeranged from 0.09 to 21.50 k9 (0.2 to (Messieh 1977).47.4 Ib) per female per year. Signif-icant linear relationships between Evidence for a nonlandlocked,juvenile emigrant abundance and nonmigratory, self-sustaining dwarfspawning population size 4 years population of alewives residing in thelater, and between the log of female mouth of the Susquehanna River wasescapement and the log of juvenile presented by Foerster and Goodbredemigrant abundance were found. (1978).Total adult freshwater mortality aver-aged 90.7% and ranged from 66% to1001 over the 11-year-study period ECOLOGICAL ROLE(Havey 1973). Total annual mortalityrates for anadromnous alewives in LongPond, Maine, were 78.6% between age Food Habits5 and 6, and 74.4% between age 6 and7 (Havey 1961). Freshwater post- Alewives and blueback herringspawning mortality for all age groups are primarily zooplanktivores, thoughaveraged 41( and ranged from 32% to fish eggs, crustacean eggs, insects

and insect eggs, and small fishes may 0I

14

be important foods in some areas or not extensive .because of the broaderfor larger individuals (Bigelow and range of foods chosen by shad (DavisSchroeder 1953). Larvae begin feeding and Cheek 1966).,on zooplankton immediately upon for-mation of a functional mouth (about 6 Few direct studies have beenmm TL), concentrating on the rela- devoted to food habits of anadromoustively small cladocerans and copepods, adult alewives and blueback herring.and adding larger species of these In general, they are zooplanktivores,groups to the diet as their mouths can with the size range and diversity ofaccommodate them (Norden 1968; Nigro available prey increasing as the fishand Ney 1982). grow and can accommodate larger

items. Considerable piscivority mayStomachs from young-of-the-year develop in landlocked populations

blueback herring collected in the (Kohler and Ney 1981).James River, Virginia, contained pri-marily (by volume) Bosmina spp., Feeding Behaviorcopepod nauplii, copepodites, and theadult copepods Eurytemora affinis and Alewives and blueback herringCyclops vernalis. Diaphanosoma bra- feed in schools of varying size, mostchyurum and Canthocamptus robert- extensively during daylight. Atcokeri were also minor food items, but night, schools disperse, and feedingwere not utilized during all seasons activity is negligible or casual, and(Burbidge 1974). Electivity (Ivlev probably by filter feeding (Burbidge1961) was strongest for adult cope- 1974; Janssen 1978). Peak stomachpods; neutral for Bosmina spp., cope- fullness occurred at 6 PM and peakpodites, and D. brachyurum; and stomach emptiness occurred at 7 AM instrongly negative for copepod nauplii. blueback herring from the JamesDaily ration for young-of-the-year River, Virginia (Burbidge 1974). Inranged from 438 g-cal per fish per addition, young-of-the-year blueback, day in July and August to 215 g-cal herring fed more actively near theper fish per day in October and surface than at 5 m (16 ft) depth,November (Burbidge 1974). even though high densities were pres-

ent at both depths (Burbidge 1974).Young-of-the-year alewives in

Hamilton Reservoir, Rhode Island, In laboratory tests (Janssenconsumed primarily chironomid midges 1976), alewives exhibited three feed-during July, switching to cladocerans ing modes: (1) particulate feeding onin August and September (Vigerstad individual prey, (2) filter feedingand Cobb 1978). Davis and Cheek with mouth agape and rapid swimming(1966) compared the food habits of bursts, and (3) gulping several preyyoung-of-the-year alewives and blue- at once but not swimming at the rapidback herring in the Cape Fear River, speed used in the filter-feeding mode.North Carolina. Blueback herring Size selectivity of prey items wasselected copepods and dipteran larvae highest in mode (1), moderate in modemore frequently (by percent of stom- (3), and negligible in mode (2). Adultachs containing items) than did ale- Lake Michigan alewives and a majorwives, while alewives consumed more food organism, Myss relicta, exhibitedostracods, insect eggs, and insect coincidental, crepuscular verticalparts than did blueback herring, migrations, from daytime residenceCrustacean eggs in the diets were near the bottom to just below thesimilarly common (-80"0 of stomachs) thermocline at night (Janssen andfor both clupeids. Overlap of either Brandt 1980). Such a diel verticaldiet with that of young-of-the-year migration, though probably not inAmerican shad Alosa sapidissima was relation to a thermocline, may occur in

15

anadromous populations residing in Temperatureestuaries or marine coastal habitats.

The effects of incubation temper-Competitors ature on alewife eggs from Lake Mich-

igan were studied by Edsall (1970).Little study has been devoted to At least some eggs hatched at test

competitive interactions of anadromous temperatures between 70 and 29.50Calewives or blueback herring. (44.60 and 85.1°F). Optimum incuba-Because of general similarities in diet tion temperature for hatching was 18*and feeding behavior, some competi- C (64 F); 38% hatch was observed.tion for food likely occurs between the Egg mortality over the first 36 hrtwo species. Loesch et al. (1982a) ranged from 22% at temperaturesdescribed a spatial separation between between 3.50 and 6.00C (380 and 430young alewives and blueback herring F) to 66% at temperatures betweenin the same habitat, which may lead to 25.5* and 28.5'C (780 and 830F). Eggreduced competition for food, at least mortality rate was directly correlatedamong juveniles. to incubation temperature (Edsall

1970). An upper lethal temperaturePredators of 29.7*C (85*F) was reported for

alewife eggs from the Hudson River,Alewives and blueback herring New York (Kellogg 1982). Maximum

are highly utilized forage species for percentage hatch occurred at 20.80Cmany riverine, estuarine, and marine (690 F), and at least some eggspiscivores, including airborne preda- hatched at test temperatures betweentors such as gulls and terns (Coin- 12.70 and 26.70C (550 and 80°F).monwealth of Massachusetts 1976).Bluefish (Pomatomus saltatrix), weak- Blueback herring eggs collectedfish (Cynoscion regalis), and striped from the Washademoak River, Ne,bass (Morone saxatilis) are predators Brunswick, Canada, were subjected toof these clupeids. Pelagic, schooling time-temperature regimes experiencedpredators such as these are more in a powerplant cooling system (Koolikely to use schooling clupeids for and Johnston 1978). Compared toforage compared to a solitary predator larval deformity rate, egg mortality(Cooper 1961; Tyus 1974). and hatchability were not good indica-

tors of the effects of temperaturechange. Deformity rate of larvae,

ENVIRONMENTAL REQUIREMENTS acclimated at 190C (66 0 F) and exposedto a 10°C (188F) increase in tempera-ture for 5 to 180 min, varied from

Some research has been con- 0 to 25% (control 0-5%). Deformityducted to delineate the specific envi- rate increased to 100% under the sameronmental requirements of anadromous conditions except with a 150C (27°F)alewife and blueback herring. Much of temperature elevation. Deformitiesthe available information was derived ranged from minor curvature of thefrom tests on landlocked populations spine to complete lack of normal larval(particularly Lake Michigan alewives), form or behavior. Deformities wereApplicability of environmental require- permanent and would not have allowedment data for landlocked populations such larvae to survive in naturalto anadromous populations is environments (Koo and Johnstonunknown. Since data from landlocked 1978).populations are major sources ofinformation on environmental require- Edsall (1970) reported twoments, they are presented but should aspects of temperature effects onbe interpreted with caution.

16

larval alewives from Lake Michigan. 0* and 1+) collected from the DelawareSurvival time of unfed larvae was 3.8 River, New Jersey, ranged from 20days at 10.5*C (510 F), 7.6 days at to 22°C (680 to 71'F) at salinities of 414.5* to 15*C (580 to 590F), and 2.4 to 6 ppt and acclimation temperaturesdays at 26.5 0 to 280C (80 to 820 F). from 15° to 21'C (590 to 70'F) (Mel-A functional jaw did not develop in drim and Gift 1971). Davis andfish from eggs/larvae held at or below Cheek (1966) captured juvenile blue-100C (500 F), even though some eggs back herring in the Cape Fear Riverhatched at such temperatures. Kel- seasonally in areas where water tem-logg (1982) reported an upper tern- peratures ranged from 11.5 ° to 32'Cperature tolerance of 310C (880F) for (530 to 89'F). Juvenile alewives in thealewife larvae from the Hudson River, same watershed were captured at tem-New York, acclimated to 140C (57 0 F). peratures between 13.5* and 290C(560Average daily gain in larval weight and 84'F).was directly proportional to watertemperature; higher growth occurred School formation patterns andat higher temperatures. A maximum daily rhythms of adult Lake Michiganlarval growth rate of 0.084 g/day alewives were affected by changes inoccurred at 29.1C (84*F), while max- temperature in laboratory tanksimum net gain in biomass (a function (Colby 1971). As water temperatureof both survival and growth) occurred dropped below 6.7*C (44F), normalat 26.4oC (79.5 0 F) (Kellogg 1982). feeding behavior was disrupted and

cruising speed of schooling fishYoung-of-the-year alewives (19 to decreased. Below 4.5*C (40°F), nor-

31 mm TL) from the Hudson River, mal schooling behavior was signifi-New York, preferred a water tempera- cantly affected. At temperaturesture of 26.3*C (79 0 F) when given a between 2.0* and 2.80c (35.5* and 370choice in a controlled thermal gradient F), alewives lost orientation, swam(Kellogg 1982). Young-of-the-year into the sides of the test chamber,alewives from Lake Michigan exhibited and ceased feeding and schooling.critical thermal maxima (CTM is themean of temperatures at which experi- In cold shock tests with adultmental fish lose equilibrium) of 28.30C alewives from Lake Michigan, transfers(83*F), 32.70C (91 0 F), and 34.4'C to test temperatures less than 3C(940F) at acclimation temperatures of (37.4 * F) caused 100% mortality110C (520F), 190C (660F), and 250C regardless of original acclimation tern-(770 F), respectively (Otto et al. peratures (Otto et al. 1976). Magni-1976). The equation for predicting tude of temperature-decrease toleratedCTM from acclimation temperature was: increased gradually with increasing

acclimation temperature. At leastCTM = 21.9 + 0.5(TA) r 2 = 0.96 some alewives survived a temperature

decrease of 100C (18*F), indepen-where TA = acclimation temperature in dently of acclimation temperature, asdegrees Celsius. long as the total test temperature did

not drop below 30C (37.4 0 F) (Otto etOtto et al. (1976) also reported that al. 1976).CTM values were 30 to 60C (5.4 0 to10.8*F) higher for young-of-the-year Stanley and Colby (1971) investi-alewives than for adults when tested gated electrolyte balance and osmoreg-at the same acclimation temperatures. ulation of Lake Michigan alewives in

relation to temperature change.In laboratory tests, preferred Transfers of fish acclimated at warm

(selected) temperatures of juvenile temperatures to cold temperaturesalewives and blueback herring (ages caused levels of Na , K , and Ca in

17

blood and muscle to move towards an Young-of-the-year alewives andequilibrium with salinity of the accli- blueback herring from the Cape Fearmation environment (increased body River system, North Carolina, select-concentrations in salt water, ed areas where free carbon dioxidedecreased body concentrations in ranged from 4 to 22 ppm, alkalinityfreshwater). Apparently, the test fish from 5 to 32 ppm, dissolved oxygentemporarily lost the ability to os- from 2.4 to 10.0 ppm, and pH frommoregulate upon exposure to cold, 5.2 to 6.8 (Davis and Cheek 1966).independently of the salinity of thetest environment.

In an experiment designed to testSalinity the effects of suspended sediments on

hatching of alewife eggs (Schubel andThough little direct information Wang 1973), a naturally occurring

exists, anadromous alewives and blue- fungus in the sediment infected allback are apparently highly tolerant of test eggs prior to hatching, and ter-salinity changes (Cooper 1961; Chit- minated the experiment. Although thetenden 1972). No mortality of adult extent of infection may have beenblueback herring from either gradual enhanced by laboratory conditions,or abrupt changes in salinity, includ- the attempt indicated that high levelsing direct transfers from fresh to salt of suspended sediment during or afterwater and the reciprocal, was spawning may significantly increaseobserved by Chittenden (1972). infection rates of eggs from naturallyBlood and muscle concentrations of the occurring fungi in sediments (Schubelelectrolytes Na + , K , and Ca + were and Wang 1973). Auld and Schubelsimilar in fish held in sea water and (1978), however, found that sus-freshwater of the same temperature, pended sediments in concentrations ofindicating that after a period of accli-- 100 ppm or less had no significantmation, alewives were efficient os- effect on hatchability of alewife or-moregulators in either environment blueback herring eggs.(Stanley and Colby 1971).

Other Environmental Factors The influence of certain environ-mental variables associated with pas-

Location of appropriate spawning sage of migratory adult alewivessites and substrates is important not through (around) hydrological obsta-only to the perpetuation of each spe- cles has been investigated. Blood lac-cies but also for natural "reproductive tic acid concentrations, measured insegregation" between two otherwise alewives moving through a pool-and-very similar species. Blueback her- weir fishway, were representative ofring prefer spawning sites with strong moderate activity and energy expendi-currents and associated hard sub- ture (Dominy 1971, 1973). Mean lev-strates (Loesch and Lund 1977). els of blood lactic acid in alewivesThey are relatively specialized com- passing through the fishway were lesspared to alewives, which use a wide than half the levels found for heavilyvariety of spawning sites, from stand- exercised fish in the laboratory. Resting river water, oxbows, coastal pools along the course of the fishwayponds, and tiny streams to fast- allowed blood lactic acid levels to dropwater, mid-river sites. Therefore, to levels comparable with those forchanges in water currents or sub- alewives in a rested state in the labo-strates in spawning rivers used by ratory.blueback herring may affect that spe-cies more than the alewife, because ofthe more specific spawning site Upstream migratory patterns ofrequirements of blueback herring, adult alewives through a Rhode Island

18

river fishway were harmonic with rine for blueback herring eggs rangeddiurnal periodicity. Periodicity was from 0.20 to 0.32 ppm. Larvae fromcorrelated with magnitude of incident eggs exposed jo sublethal concentra-solar radiation (Saila et al. 1972). tions of total residual chlorine were allRichkus (1974a) corroborated this deformed (Morgan and Prince 1977).light-dependent migratory activity; he Concentrations of kepone greater thanalso observed that within activity pat- 0.3 ppm (termed the "action level" forterns determined by light intensity, possible closure of a fishery) werechanges in water temperature strongly found in body tissues of young-of-influenced specific timing of alewife the-year alewives and blueback her-upstream movement. Juvenile down- ring collected from the James andstream emigration from Hamilton Res- Chickahominy Rivers, Virginia (John-ervoir, Rhode Island, during summer son et al. 1978; Loesch et al.and fall was inhibited by the bright 1982b). Kepone was also present insunlight-bridge shade interface pres- young alewives and blueback herringent at a road bridge on the lower end from the Mattaponi and Pamunkey Riv-of the reservoir. Higher emigration ers, Virginia (in concentrations lessrates under this bridge were observed than 0.3 ppm), but was not presenton cloudy days (Richkus 197 4a). in detectable quantities in fish from

the Rappahannock River, Virginia,Environmental Contaminants and the Potomac River, Maryland

(Loesch et al. 1982b).The LC 50 of total residual chlo-

19

LITERATURE CITED

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setts. Mass. Dep. Fish.,Berry, F. H. 1964. Review and emen- Wildl., and Recreational Vehicles,

dation of family Clupeidae, Cop- Div. Mar. Fish. Proj. Rep. No.eia 1964: 720-730. AFCS-14-1, Public information

pamphlet. 20 pp.Bigelow, H. B., and W. C. Schroe-

der. 1953. Fishes of the Gulf of Cooper, R. A. 1961. Early life historyMaine. U. S. Fish Wildl. Serv. and spawning migration of theFish. Bull. No. 74. 577 pp. alewife. M. S. Thesis. Univer-

sity of Rhode Island, Kingston.Bigelow, H. B., and W. W. Welsh. 58 pp.

1925. Fishes of the Gulf ofMaine. U. S. Bur. Fish. Bull. Davis, J. R., and R. P. Cheek. 1966.No. 40. 567 pp. Distribution, food habits and

growth of young clupeids, CapeBurbidge, R. G. 1974. Distribution, Fear River system, North Caro-

growth, selective feeding and lina. Paper presented at Annu.energy transformations of Meet. South. Div. Am. Fish.young-of-the-year blueback her- Soc., Asheville, N. C., 1966. 20ring in the James River, Vir- pp.ginia. Trans. Am. Fish. Soc.103: 297-311. Dominy, C. L. 1971. Changes in blood

lactic acid concentrations in ale-Chambers, J. R., J. A. Musick, and wives during passage through a

J. Davis. 1976. Methods of dis- pool-and-weir fishway. J. Fish.tinguishing larval alewife from Res. Board Can. 28: 1215-1217.larval blueback herring. Chesa-peake Sci. 17: 93-100. Dominy, C. L. 1973. Effect of

entrance pool weir elevation andChittenden, M. E. 1972. Salinity tol- fish density on passage of ale-

erance of young blueback her- wives in a pool-and-weir fishway.ring. Trans. Am. Fish. Soc. Trans. Am. Fish. Soc. 102:101: 123-125. 398-404.

Cianci, J. M. 1969. Larval develop- Dovel, W. L. 1971. Fish eggs and lar-ment of the alewife and the glut vae of the upper Chesapeakeherring. M. S. Thesis. Univer- Bay. Univ. Maryland, Nat.sity of Connecticut, Storrs. 62 Resour. Inst. Spec. Rep. No. 4.pp. 71 pp.

Colby, P. J. 1971. Alewife dieoffs: Edsall, T. A. 1964. Feeding by threewhy do they occur? Limnos 4(2): species of 'ishes on the eggs of18-27. spawning alewives. Copeia 1964:

226-227.Collins, G. B. 1952. Factors influenc-

ing the orientation of migrating Edsall, T. A. 1970. The effect ofanadromous fishes. U. S. Fish temperature on the rate of

20

, II

development and survival of ale- Janssen, J., and S. B. Brandt. 1980.wife eggs and larvae. Trans. Feeding' ecology and verticalAm. Fish. Soc. 99: 376-380. migration of adult alewives in

Lake Michigan. Can. J. Fish.Edsall, T. A., and M. I. Saxon. Aquat. Sci. 37: 177-184.

1968. Two hermaphroditic ale-wives from Lake Michigan. Copeia Johnson, H. B., D. W. Crocker, B.1968: 406-407. F. Holland, Jr., J. W. Gilliken,

D. W. Taylor, M. W. Street, J.Foerster, J. W., and S. L. Goodbred. G. Loesch, W. H. Kriete, Jr.,

1978. Evidence for a resident and J. G. Travelstead. 1978.alewife population in the northern Biology and management of mid-Chesapeake Bay. Estuarine Atlantic anadromous fishes underCoastal Mar. Sci. 7: 437-444. extended jurisdiction. N. C. Div.

Mar. Fish. and Va. Inst. Mar.Graham, J. J. 1956. Observations on Sci. Rep. NC-VA AFSC 9-2. 175

the alewife in freshwater. Univ. PP.Toronto Biol. Ser. No. 62. 43pp. Jones, P. W., F. D. Martin, and J.

D. Hardy, Jr. 1978. DevelopmentHavey, K. A. 1961. Restoration of of fishes of the mid-Atlantic

anadromous alewives at Long Bight: an atlas of the egg, larvalPond, Maine. Trans. Am. Fish. and juvenile stages, Volume I. U.S.Soc. 90: 281-286. Fish Wildl. Serv. Biol. Serv.

Program FWS/OBS-78/12. 366Havey, K. A. 1973. Production of PP-

juvenile alewives at Love Lake, Joseph, E. B., and J. Davis. 1965. AWashington County, Maine. preliminary assessment of theTrans. Am. Fish. Soc. 102: river herring stocks of lower434-437. Chesapeake Bay. Va. Inst. Mar.

Hildebrand, S. F. 1963. Family Clu- Sci. Spec. Sci. Rep. No. 51.peidae. Pages 257-385, 397-442, 23 pp.and 452-454 in Fishes of the Kellogg, R. L. 1982. Temperaturewestern North Atlantic. Sears requirements for the survival andFound. Mar. Res. Mem. 1(3). early development of the anadro-

mous alewife. Prog. Fish-Cult.Hlavek, R. R., and C. R. Norden. 44: 63-73.

1977. Two hermaphroditic fresh-water alewives from southern Kissil, G. W. 1969. Contribution toLake Michigan. Prog. Fish-Cult. the life history of the alewife,39(2): 104-105. Alosa pseudoharengus, in Con-

necticut. Ph.D. Thesis. Univer-Ivlev, V. S. 1961. Experimental ecol- sity of Connecticut, Noank. 11

ogy of the feeding of fishes. Yale pp.University Press, New Haven,Conn. 302 pp. Kissil, G. W. 1974. Spawning of the

anadromous alewife in BrideJanssen, J. 1976. Feeding modes and Lake, Connecticut. Trans. Am.

prey size selection in the alewife. Fish. Soc. 103: 312-317.J. Fish. Res. Board Can. 33:1972-1975. Kohler, C. C., and J. J. Ney. 1981.

Consequences of an alewife dieoffanssen, J. 1978. Will alewives feed in to fish and zooplankton in a res-

the dark? Environ. Biol. Fishes ervoir. Trans. Am. Fish. Soc.3: 239-240. 110: 360-369.

21

Koo, T. S. Y., and M. L. Johnston. Loesch, J. G., W. H. Kriete, Jr.,1978. Larval deformity in striped and E. J. Foell. 1982a. Effects ofbass and blueback herring due to light intensity on the catchabilitbheat shock treatment of develop- of juvenile Alosa species. Trans.ing eggs. Environ. Pollut. 16: Am. Fish. Soc. 111: 41-44.137-149.

Loesch, J. G., R. J. Hugget, and E.Kornegay, J. W. 1978. Comparison of J. Foell. 1982b. Kepone concen-

aging methods for alewife and tration in juvenile anadromousblueback herring. N. C. Dep. fishes. Estuaries 5(3): 175-181.Nat. Resour., Div. Mar. Fish.Spec. Sci. Rep. No. 30. 51 pp. MacLellan, P., G. E. Newsome, and

P. A. Dill. 1981. DiscriminationKuntz, A., and L. Radcliffe. 1917. by external features between ale-

Notes on the embryology and lar- wife and blueback herring. Can.val development of twelve teleos- J. Fish. Aquat. Sci. 38:tean fishes. U. S. Bur. Fish. 544-546.Bull. No. 35. 134 pp.

Mansueti, R. J. 1956. Alewife herringLeggett, W. C. G., and R. R. Whit- eggs and larvae reared success-

ney. 1972. Water temperature and fully in lab. Maryland Tidewa-the migrations of the American ter News 13(1): 2-3.shad. U. S. Nati. Mar. Fish.Serv. Fish. Bull. No. 70. 670 Mansueti, R. J. 1962. Eggs, larvaepp. and young of the hickory shad,

with comments on its ecology inLeim, A. H., and W. B. Scott. 1966. the estuary. Chesapeake Sci. 3:

Fishes of the Atlantic coast of 173-205.Canada. Fish. Res. Board Can.Bull. No. 155. 485 pp. Marcy, B. C., Jr. 1969. Age determi-

nation from scales of Alosa pseu-Loesch, J. G. 1968. A contribution to doharengus and Alosa aestivalis

the life history of Alosa aesti- in Connecticut waters. Trans.valis. M. S. Thesis. University Am. Fish. Soc. 98: 621-630.of Connecticut, Storrs. 31 pp.

McCoy, E. G. 1975. Statement ofLoesch, J. G. 1969. A study of the North Carolina Division of Marine

blueback herring in Connecticut Fisheries concerning river her-waters. Ph.D. Thesis. University ring. Rep. N. C. Dep. Nat.of Connecticut, Storrs. 78 pp. Econ. Resour. 5 pp.

Loesch, J. G. 1981. Weight relation McKenzie, J. A. 1973. Comparativebetween paired ovaries of blue- electrophoresis of tissues fromback herring. Prog. Fish-Cult. blueback herring and gaspareau.43(2): 77-79. Comp. Biochem. Physiol. 44B:

65-68.Loesch, J. G., and W. A. Lund.

1977. A contribution to the life McKenzie, J. A. 1975. Retina-specifichistory of the blueback herring. LDH isozyme in blueback herringTrans. Am. Fish. Soc. 106: and alewife. Anim. Blood Groups583-589. Biochem. Genet. 6: 245-247.

22

Meldrim, J. W., and J. J. Gift. 1971. back and. alewife from GeorgesTemperature preference, avoid- Bank, July and October, 1964.ance and shock experiments wit+h ICNAF Res. Bull. No. 3, 1966., estuarine fishes. lchthyol. Assoc. 5 pp.Bull. No. 7. 75 pp. Neves, R. J. 1981. Offshore distribu-

Merriner, J. V. 1978. Anadromous tion of alewife and blueback her-fishes of the Potomac Estuary. ring along the Atlantic coast. U.S.Va. Inst. Mar. Sci. Contrib. Natl. Mar. Fish. Serv. Fish.No. 696, Gloucester Point. 4 pp. Bull. 79: 473-485.

Messieh, S. N. 1977. Population Nigro, A. A., and J. J. Ney. 1982.Reproduction and early-lifestructure and biology of alewives accommodations of landlocked ale-

and blueback herring in the Saint aoten ange e--John River, New Brunswick. wives to a southern range exten-

JonR iver Ne sBuswick.:sion. Trans. Am. Fish. Soc.Environ. Biol. Fishes 2(3): 111: 559-569.

195-210.

Milstein, C. B 1981 Abundance and Norden, C. R. 1967. Development andi"stin Bunde and identification of the larval alewifedistribution of juvenile Alosa in Lake Michigan. Proc. 10th

species off southern New Jersey. in L e Michigan. : 0th

Trans. Am. Fish. Soc. 110: Conf. Great Lakes Res.: 70-78.

306-309. Norden, C. R. 1968. Morphology and

food habits of the larval alewifeMorgan, R. P., II, and R. D. Prince. in Lake Michigan. Proc. 11th

1976. Chlorine toxicity to estua- Conf. Great Lakes Res.: 103-110.rine fish eggs and larvae. Ches-apeake Biol. Lab. Univ. Maryland O'Neill, J. T. 1980. Aspects of theCent. Environ. Estuarine Stud. life histories of anadromous ale-Ref. No. 76-116 CBL. 122 pp. wife and the blueback herring,

Margaree River and Lake Ainsle,Morgan, R. P., 11, and R. D. Prince. Nova Scotia, 1978-1979. M. S.

1977. Chlorine toxicity to eggs Thesis. Acadia University, Wolf-and larvae of five Chesapeake ville, Nova Scotia, Canada. 306Bay fishes. Trans. Am. Fish. PP.Soc. 106: 380-385.

Otto, R. G., M. A. Kitchel, and J.National Marine Fisheries Service 0. Rice. 1976. Lethal and pre-

(NMFS). 1980. Marine recreational ferred temperatures of the alewifefishery statistics survey, Atlantic in Lake Michigan. Trans. Am.and gulf coasts, 1979. U. S. Fish. Soc. 105: 96-106.Dep. Comm. Current FisheriesStatistics No. 8063. 139 pp. Pate, P. P. 1974. Age and size com-

position of commercial catches ofNational Marine Fisheries Service blueback herring in Albemarle

(NMFS). 1982. Fisheries of the Sound, North Carolina, and itsUnited States, 1981. U. S. Dep. tributaries. Rep. N. C. Dep.Comm. Current Fisheries Statis- Nat. Econ. Resour., Div. Com-tics No. 8200. 131 pp. mercial Sport Fish. 10 pp.

Netzel, J., and E. Stanek. 1966. Some Price, W. S. 1978. Otolith comparisonbiological characteristics of blue- of Alosa pseudoharengus and

22

Alosa aestivalis. Can. J. Zool. Schubel, J. R., and J. C. S. Wang.56: 1216-1218. 1973. The effects of suspended

sediments on the hatching suc-Raney, E. C., and W. H. Massmann. cess of yellow perch, white

1953. The fishes of the tidewater perch, striped bass and alewifesection of the Pamunkey River, eggs. Ichthyol. Assoc. Spec.Virginia. J. Wash. Acad. Sci. Rep. No. 30, Ref. 73-3. 77 pp.43(12): 424-432.

Scott, W. B., and E. J. Crossman.Richkus, W. A. 1974a. Factors influ- 1973. Freshwater fishes of Can-

encing the seasonal and daily ada. Fish. Res. Board Can.patterns of alewife migration in a Bull. No. 184. 966 pp.Rhode Island River. J. Fish.Res. Board Can. 31: 1485-1497. Smith, B. A. 1971. The fishes of four

low salinity tidal tributaries ofRichkus, W. A. 1974b. Influence of the Delaware River Estuary. M.S.

environmental variables on the Thesis. Cornell University,migratory behavior of adult and Ithaca, N. Y. 304 pp.juvenile alewives. Ph.D. Thesis.University of Rhode Island, Stanley, J. G., and P. J. Colby.Kingston. 225 pp. 1971. Effects of temperature on

electrolyte balance and osmoregu-Richkus, W. A. 1975. Migratory lation in the alewife in fresh and

behavior and growth of juvenile sea water. Trans. Am. Fish.anadromous alewives in a Rhode Soc. 100: 624-638.Island drainage. Trans. Am.Fish. Soc. 104: 483-493. Street, M. W. 1969. Fecundity of the

blueback herring in Georgia. Ga.Richkus, W. A., and H. E. Winn. Game Fish Comm. Mar. Fish.

1979. Activity cycles of adult and Div., Contrib. Ser. 17. 15 pp.juvenile alewives, recorded bytwo methods. Trans. Am. Fish. Street, M. W., and J. Davis. 1976.Soc. 108: 358-365. Notes on the river herring fish-

ery of SA6. ICNAF Res. Doc.Rideout, S. G., J. E. Johnson, and 76/VI/61. 7 pp.

C. F. Cole. 1979. Periodic countsfor estimating the size of the Thunberg, B. E. 1971. Olfaction inspawning population of alewives, parent stream selection by theEstuaries 2: 119- 123. alewife. Anim. Behav. 19:

217-225.Rulifson, R. A., and M. T. Huish.

1982. Anadromous fish in the Tyus, H. M. 1971. Population size,Southeastern United States and harvest and movements of ale-recommendations for development wives during spawning migrationsof a management plan. U. S. to Lake Mattamuskeet, NorthFish Wildl. Serv., Region 4, Carolina. Ph.D. Thesis. NorthAtlanta, Ga. 525 pp. Carolina State University,

Raleigh.Saila, S. B., T. T. Polgar, D. J.

Sheehy, and J. M. Flowers. Tyus, H. M. 1974. Movements and1972. Correlations between alewife spawning of anadromous alewivesactivity and environmental vari- at Lake Mattamuskeet, Northables at a fishway. Trans. Am. Carolina. Trans. Am. Fish. Soc.Fish. Soc. 101: 583-594. 103: 392-396.

24

Vigerstad, T. J., and J. S. Cobb. Res. Job Compl. Rep. No.1978. Effects of predation by AFSC-13/FWAC-2. 37 pp.sea-run juvenile alewives on thezooplankton community at Hamil- Warinner, J. E., J. P. Miller, and J.ton Reservoir, - Rhode Island. Davis. 1969. Distribution of juve-Estuaries 1(1): 36-45. nile river herring in the Potomac

River. Proc. Annu. Conf. South-Walton, C. J. 1979. Growth of Alosa east. Assoc. Game Fish Comm.

pseudoharengus from two Maine 23: 384-388.watersheds. Maine Dep. Mar.Resour. Res. Ref. Doc. No. Winters, G. H., J. A. Moores, and R.79/23. 75 pp. Chaulk. 1973. Northern range

extension and probable spawningWalton, C. J., M. E. Smith, and D. of gaspareau in the Newfoundland

B. Sampson. 1976. Alewife man- area. J. Fish. Res. Board Can.agement plan. Maine Dep. Mar. 30: 860-861.

25

50272 -101

REPORT DOCUMENTATION 1. REPORT NO. 2. 1 3. trt" ,on N.

PAGE FWS/OBS-82/119*4. Title and Subtitle 5 epor Dat

Species Profiles: Life Histories and Environmental Requirements of! October 1983Coastal Fishes and Invertebrates (Mid-Atlantic) -- Alewife/Blue- 1.back Herring _

7. Author(s) 8. Perforrning Organization Rapt. No

leiFon-..1Fay. Richard J. Neves, and Garland B. Pardue9. Performing Organization Name and Address 10. Proiect/Task,'Work Unit No.

Department of Fisheries and Wildlife SciencesVirginia Polytechnic Institute and State University 1t. Contract(C) or Grant(G) No

Blacksburg, VA 24061 '(C)

2. Sponsoring Orgarizatgn. and Addressea U 13. Type of Report & Period Coveredational Coastal cosystem'ssseam U.S. Army Corps of Engineers

Fish and Wildlife Service Waterways Experiment StationU.S. Department of the Interior P.O. Box 631 .

Washington, DC 20240 Vicksburg, MS 39180

15. Supplementary Notes

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

16. Abstract (Limit: 200 words) Species profiles are lierature summaries of th'e--Ta xonoy,-6oph--og-y.-range, life history, and environmental requirements of coastal aquatic species. They areprepared to assist in impact assessment. The alewife and blueback herring, Alosa pseudo-harengus and Alosa aestivalis, are important species in estuarine and marine ecosystems aslinks between the zooplankton they consume and top piscivores. Both anadromous and land-locked populations exist.) Some individuals mature by age 3, and all mature by age 5. Repeaspawning is common. 4S-wning environments range from streams only a few centimeters deep tolarge rivers. Alewives will also spawn in ponds with an open connection to the sea. Blue-alewives select a wider variety of sites, from standing water and oxbows to mid-river areas.

Spawning occurs from April to July in the mid-Atlantic region; the onset and peak of alewifespawning precede those of blueback herring by 2 to 3 weeks. Larvae and juveniles remain inor near areas spawned before emigrating (as juveniles) to coastal areas between June andNovember of their first year.' Emigration is apparently triggered by heavy runoff from rain

and/or sharp decreases in water emperature. Adults overwinter offshore to depths of atleast 110 m. Nantucket Shoals, Georges Bank, and the Gulf of Maine are important overwinter.ing grounds. Commercial and limited recreational fisheries for these species occur; totalU.S. landings in 1981 were 3,754 mt, while foreign landings were 13.9 mt. Some eggs canhatch at water temperatures between 70 aLV 29.5I

, but temperatures above 29.7"t are lethal.

Larvae need temperatures greater than 10C for proper development; upper lethal temperature- 17o's __ .utTu-O-erptrs- _... - - 31 q ,

River r Growth (PhS;o/-Anady umous Feeding, 0711)Estuaries) . -

_>4 -Fish*

b. Identifiers/Open Ended Terms

Alewife Temperature requirementAlosa pseudoharengus alinity requirements.W6,. __.Blueback herring SpawningAlosa aestivalis

C. COSATI Field/GroupI'- ,-ailatddy Statement 19. Security Class (This Report) 21. No ot Pages

Unclassified 25Unlimited 20. Security Class (This Page) 22. Pri

Unclassified(See AN SI-Z39.18) OPTIONAL FORM 272 (4-77)

(Foreerly NTIS-35) 0Department Of Commerce

2.4

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Hawaiian ani

SHeadquarters,. Ovsion of BiologicalServices Washington. DC

X Eastern Energy and Land Us. TeamL e e to w n" --

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*National Coastal Ecosystems Team

* Western Energy and Land Use TeamFt Collins. CO

• Locations of Regional Offices

REGION I REGION 2 REGION 3Regional Director Regional Director Regional DirectorU.S. Fish and Wildlife Service U.S. Fish and Wildlife Service U.S. Fish and Wildlife ServiceLloyd Five Hundred Building, Suite 1692 P.O. Box 1306 Federal Building, Fort Snelling500 N.E. Multnomah Street Albuquerque, New Mexico 87103 Twin Cities, Minnesota 55111Portland, Oregon 97232

REGION 4 REGION 5 REGION 6Regional Director Regional Director Regional DirectorU.S. Fish and Wildlife Service U.S. Fish and Wildlife Service U.S. Fish and Wildlife ServiceRichard B. Russell - ilding One Gateway Center P.O. Box 2548675 Spring Street, S.W. Newton Corner, Massachusetts 02158 Denver Federal CenterAtlanta, Georgia 30303 Denver, Colorado 80225 ..

REGION 7Regional DirectoiU,S. Fish and Wildlife Service0 I11 L. Tudor Road

Anchorage, Alaska 99503

......._

.iI

Nv-m

DEPARTMOT OF THE INTERIORU.S. FISH AND WILDLIFE SERVICE

As the Nation's principal conservation agency, the Department of the Interior has respon-sibility for most of our nationally owned public lands and natural resources. This includesfostering the wisest use of our land and water resources, protecting our fish and wildlife,preserving the environmental and cultural values of our national parks and historical places,and providing for the enjoyment of life through outdoor recreation. The Department as-sesses our energy and mineral resources and works to assure that their development is inthe best interests of all our people. The Department also has a major responsibility forAmerican Indian reservation communities and for people who live in island territories underU.S. administration.

NOW WIRNIMPOWMENIONEEMEMP ML"dmmmmov