Alewife/Blueback Herring

Home , Alewife (fish), Blueback herring

~~~~F-?$?~2A~~\ Do Not Remove from the Ubrory U. S. Fish and Wildlife Service I'varfonal wetlands Kesearch Lenter Biological Report 82 (1 1.56) TR EL-82-4 JUIY 1986 700 Cajun Dome Boulevard Lofcryette, Louisiana 70506

Species Profiles: Life Eistories and Environmental Requirements of Coastal Fishes and Invertebrates (North Atlantic)

ALEWIFE/BLUEBACK HERRING

Coastal Ecology Group Fish and Wildlife Service Waterwavs Ex~erimentStation U.S. Department of the Interior U.S. Army Corps of Engineers Biological Report 82(1l.s6) TR EL-82-4 July 1986

Species Prof i1 es : Life Histories and Envi ronmental Requi rements of Coastal Fishes and Invertebrates (North Atlantic)

Dennis M. Mullen, Clemon W. Fay, and John R. Moring Maine Cooperative Fishery Research Unit 313 Murray Hall University of Maine Orono, ME 04469

Project Officer John Parsons National Wet1 ands Research Center U.S. Fish and Wildlife Service 1010 Gause Boulevard Slidell, LA 70458

Performed for Coastal Ecology Group Waterways Experiment Station U.S. Army Corps of Engineers Vicksburg, MS 39180

and

National Wetlands Research Center Research and Development Fish and Wildlife Service U. S. Department of the Interior Washington, DC 20240 This series should be referenced as follows:

U.S. Fish and Wildlife Service. 1983-19-. Species profiles: life histories and environmental requirements of coastal fishes and invertebrates. U. S. Fish Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4.

This profile should be cited as follows:

Mullen, D.M., C.W. Fay, and J.R. Moring. 1986. Species profiles: life histories and environmental requirements of coastal fishes and invertebrates (North Atlantic)--alewife/blueback herring. U. S. Fish Wildl . Serv. Biol . Rep. 82(11.56). U. S. Army Corps of Engineers, TR EL-82-4. 21 pp. PREFACE

This species profile is one of a series on coastal aquatic organisms, principally fish, of sport, commercial, or ecological importance. The profiles are designed to provide coastal managers, engineers, and biologists with a brief comprehensive sketch of the biological characteristics and environmental requirements of the species and to describe how populations of the species may be expected to react to environmental changes caused by coastal development. Each profi 1 e has sections on taxonomy, 1i fe history, ecological role, environmental requirements , and economi c importance, if appl icab1 e. A three-ri ng binder is used for this series so that new profiles can be added as they are prepared. This project is jointly planned and financed by the U. S. Army Corps of Engineers and the U.S. Fish and Wildlife Service.

Suggestions or questions regarding this report should be directed to one of the following addresses.

Information Transfer Specialist National Wetlands Research Center U.S. Fish and Wildlife Service NASA-Slidell Computer Complex 1010 Gause Boulevard Slidell, LA 70458

U. S. Army Engineer Waterways Experiment Station Attention: WESER-C Post Office Box 631 Vicksburg, MS 39180 CONVERSION TABLE

-Metric to U.S. Customary Mu1 tiply & To Obtain mill imeters (mn) inches centimeters ( an) inches meters (m) feet ki1 ometers ( km) mil es 2 square meters (m ) 10.76 square feet square ki1 meters ( km2) 0.3861 square 1ni1es hectares (ha) 2.471 acres liters (1) gal 1ons cubic aeters (m3) cubic feet cubic rneters acre-feet mill igrams (mg) 0.00003527 ounces grams (g) 0.03527 ounces kilograms (k ) 2.205 pounds metric tons qt) 2205.0 pounds metric tons 1.102 short tons ki1 ocal ories ( kcal ) 3.968 British thermal units

Cel sius degrees 1.8(Oc) + 32 Fahrenheit degrees

U.S. Customary to Metric

inches 25.40 mil 1 imeters inches 2.54 centimeters feet (ft) 0.3048 meters fathoms 1.829 meters miles (mi) 1.609 kilometers nautical miles {mi) 1.852 ki1 oveters

square feet (ft2) square meters acres 2 hectares square miles (mi ) square ki1 ometers gal 1ons ( gal ) 1i ters cubic feet (ft3) cubic meters acre- feet cubic meters ounces (02) 28.35 grams pounds (lb) 0.4536 ki1 ograrns short tons (ton) 0.9072 metric tons British thermal units (Btu) 0.2520 ki1 ocal ories Fahrenheit degrees 0.5556("F - 32) Celsius degrees CONTENTS Page PREFACE ...... i i i CONVERSION TABLE ...... i v ACKNOWLEDGMENTS ...... vi PROFILE SCOPE ...... 1 NOMENCLATURE /TAXONOMY /RANGE ...... 1 MORPHOLOGY/IDENTIFICATION AIDS ...... 2 Alewife ...... 2 Blueback Herring...... 2 Aids for Species Separation ...... 2 REASON FOR INCLUSION IN SERIES ...... 2 LIFE HISTORY ...... 5 Reproducti ve Characteristics ...... 5 Fecundity ...... 5 Spawning ...... 6 Eggs ...... 7 Yolk-Sac Larvae ...... 7 Larvae ...... 7 Juveniles ...... 7 Adults ...... 8 GROWTH CHARACTERISTICS ...... 9 Growth Rates ...... 9 Length-Wei ght Re1 ations ...... 9 THE FISHERY ...... 9 Commercial Fisheries ...... 9 Population Dynamics ...... 11 ECOLOGICAL ROLE ...... 11 Food Habits ...... 11 Feeding Behavior ...... 13 Competitors ...... 13 Predators ...... 13 ENVIRONMENTAL REQUIREMENTS ...... 13 Temperature ...... 13 Salinity ...... 15 Other Environmental Factors ...... 15 Envi ronmental Contaminants ...... 16 Entrainment and Turbine Induced Mortality ...... 16 LITERATURE CITED ...... ACKNOWLEDGMENTS

We are grateful for the reviews by Joseph Loesch, Virginia Institute of Marine Science, Gloucester Point, and Lewis Flagg, Maine Department of Marine Resources, Augusta, Maine. Figure 1. A: alewife; B: blueback herring.

ALEWIFEIBLUEBACK HERRING

PROF ILE SCOPE considered similar among the two species unless noted as such. In a This profile describes the life few reports (particularly those on history and environmental requirements comrwrcial fisheries statistics), the of the alewife (Alosa seudoharen us) two species are referred to col- and the blueb~Rdalectively as "river herring" or "gas- aestivalis), since their distribution pereau" (Canada). is overlappi. . nq and their morpholosv, ecological r6l e, and envi ronmental requ irements are similar. NOMENCLATURE /TAXONOMY /RANGE Nevertheless, significant differences in certain physical, physiological, and biological characteristics exist Scientific names . . . . . 9losa between the two species. When seudoharen us (Wilson) or ~lhsa information is available, these '-chill) differences are addressed by separate Preferred common names...... statements for each species. In alewife and blueback herring addition, a special section on the (Figure 1) most readily distinguishing charac- Other common names ...... teristics for separating eggs, larvae, alewife -- freshwater herring, and adults of the two clupeids is grayback, gaspereau, sawbel ly, kyak, presented in the morphology section. branch herri ng; blueback herring -- glut herring, summer herring, black Because most of the information be1 ly, kyack (Bigelow and Schroeder available for the two species concerns (1953) the alewife, characteristics of this Class...... Osteichthyes species are given priority reference Order...... Clupeiformes in the text. Features should not be Fami ly...... C lupei dae Geographical range: The a1 ewife and arch 41-52. Body moderately com- blueback herring are anadromous pressed, elongate, eye diameter small, species common in rivers, estuaries, equal to or less than snout length. and coastal waters of the North Upper jaw with definitive median At1 antic from Newfoundland (Wi nters notch, no teeth on premaxi 11aries . et al. 1973) to South Carolina Color: dorsally, blue to blue-green; (Berry 1964). Self-sustaining, lateral ly, silver with prominent dark landlocked populations have been shoulder spot; fins pale, yellow, or established in the Laurentian Great green. Adults often with rather dis- Lakes, the Finger Lakes of New York, tinct dusky lines along the back as and several other freshwater 1akes shown in Figure 1. (Bigelow and Schroeder 1953; Scott and Crossman 1973). The blueback Aids for Species Separation herring is geographically distri - buted along the Atlantic coast from Eggs. Unferti1 ized blueback Nova Scotia to the St. Johns River, herring eggs amber, alewife eggs F 1orida (Hi 1debrand 1963). The greenish. Oi 1 droplets of fertilized distribution of alewives and blueback herring eggs unequal in size blueback herring from Cape Cod and scattered; those of alewife eggs north to Maine is illustrated in numerous and uniformly small (Kuntz Figures 2a and 2b. and Radcl iffe 1917; Norden 1967).

Larvae. Myomeres between inser- MORPHOLOGY/IDENTIFICATION AIDS tion of dorsaq fin and anal vent 11-13 for blueback herring and larvae, 7-9 The following information was for alewife (this characteristic is taken from morphological summaries definiti ve according to Chambers et prepared by Jones et al. (1978), a1 . C19761). According to Chambers et unless otherwise indicated. al. (1976), larvae less than 15 mm long can be separated by using regres- Alewife sions of vent to tail distance and vent to urostyle distance against Dorsal rays 12-19 (usually standard length (SL). 13-14), anal rays 15-21 (usually 17-18), scales in lateral line series Adults. The two species are 42-54. Belly with scutes forming a morphological ly s imi1 ar but can be keel. Prepel vic scutes 17-21 (usual ly separated by differences in scale 19-20), postpelvic scutes 12-17 imbrication patterns and individual (usually 14-15), gill rakers on first scale markings (Figure 3). Alewives arch 38-46. Body strongly compressed, usually have the fewer vertebrae, deep. Mouth oblique, anterior end of dorsal rays, and gill rakers on the lower jaw thick, heavy; jaw extending f irst arch. to middle of orbit. Eye large, diameter greater than snout length. Color: dorsal ly, gray to gray-green; REASON FOR INCLUSION IN THE SERIES laterally, si1 ver with prominent dark shoulder spot; fins pale, yellow, or green. The alewife and blueback herri ng are important forage and commercial Blueback Herring fishes. Ecologically , they are important energy 1inks between zoo- Dorsal rays 15-20, anal rays plankton and piscivores. In New 15-21, scales in lateral series 46-54. England both species have a long Prepel vic scutes 18-21, postpel vic history of commercial exploitation and scutes 12-16, gill rakers on first are important as sources of fish meal, A TLA N TIC OCEAN

Coastal distribution

MILES

Figure 2a. Distribution of alewives along the U.S. North Atlantic coast.

3 ATLANTIC OCEAN

Coalltal dlstrlbution

MILES

Figure 2b. Distribution of blueback herring along the U.S. North Atlantic coast.

4 among the older fish. This relation suggests that males mature at an earlier age than females but have a shorter life span. The age of first spawning of blueback herring varies more that that of the alewife, yet they mature at about the same age (Joseph and Davis 1965; Loesch and Lund 1977; O'Neill 1980).

Age at f i rst spawning, percentage of repeat spawners, and 1ongevi ty of ALEWIFE BLUEBACK HERRING alewives tend to decrease from north approx. scale: -lcm to south. For example, spawning ale- I-,Arrnrol ulc1v --.--e--.------I wives in southern North Carolina were numerically dominated by Age I11 fish; none were older than Age IV (Tyus 1974). In contrast, alewife spawning populations consisted of age groups I11 to VIII in Chesapeake Bay (Joseph and Davis 1965) and in the Connecticut River (Marcy 1969; Loesch and Lund 1977), and of age groups IV to X for both species in Nova Scotia (O'Neil 1980). The percentages of repeat spawners (fish that spawned during two

ALIma WIrnYICI IIImm or more years) were 60% for alewives approx. scale -5 mm in Nova Scotia (OINeill 1980); 61% in the York River, Virginia (Joseph and Figure 3. Scale imbrication patterns Davis 1965); and less than 10% in (top) and individual scale morphology southern North Carolina (Tyus 1974). (bottom) used for external Repeat spawners among blueback herring discrimination of the alewife and the were 65% in the York River and 75% in blueback herring (from MacLellan et Nova Scotia (Joseph and Davis 1965; al. 1981; used with permission of the O'Neill 1980). Canadian Journal -of Fisheries and Aquatic Sciences). The females of alewives and blueback herring are sli ghtly larger and heavier than males of the same age fish oil, and fish protein, particu- (Cooper 1961; Netzel and Stanek 1966; larly for the animal food industry. Marcy 1969; Loesch and Lund 1977). Alewives also are important as bait in the New England lobster fishery. Fecundity

Fecundity estimates for female LIFE HISTORY alewives in the Parker River, Massa- chusetts, ranged from about 68,000 to Reproductive Characteristics 206,000 eggs per female (Cole et al. 1980). Age group V females averaged A small percentage of anadromous 168,000 eggs whereas females of age alewives spawn for the first time in groups I11 and VII averaged only about their fourth year of life (age group 118,000 eggs (Cole et al. 1980). 111). During spawning, males Fecundity of Chesapeake Bay alewives numerically dominate fish in age ranged from 60,000 to 100,000 eggs per groups I11 and IV and females dominate female (Foerster and Goodbred 1978). The age-fecundity relation may be water flow, suggesting that emigration asymptotic for both alewives and of adults after spawning is a rheotac- blueback herring, and "fecundal tic response (Huber 1978). seni 1ity" may characterize all 1ong-1 ived stocks of these species In laboratory tests, adult (Street 1969; Loesch and Lund 1977). anadromous alewives from Rhode Island waters were capable of distinguishing Females apparently lay nearly all water of their natal pond from water of their eggs during spawning. Spent collected in nearby ponds. Olfaction female alewives leaving the Parker was shown to be the major sensory River contained less than 0.1% of the mechanism for homing behavior average number of eggs in the females (Thunberg 1971). A1 though homing entering the spawning runs. About 7% behavior is strong in alewives, con- of the adult female alewives returning siderable mixing of stocks takes to the sea from the Parker River in place. Discriminant function analyses 1975 and 1976 had not spawned (Cole et by Messieh (1977) for fish of the St. al. 1980). Female blueback herring in Johns River, Florida, indicated that the Connecticut River retained 10% to spawning stocks often strayed from 30% of their full complement of eggs home streams, particularly between after spawning (Loesch and Lund 1977). adjacent spawning areas and stocks. Most of the mixing of stocks was Spawni ng during the prespawni ng period (late winter, early spring) rather than Historically in Maine, alewives during the actual spawning runs. and blueback herring made spawning runs up all or nearly all streams with Alewives spawn along the Atlantic access to lakes, ponds, and backwaters coast from late March through July; (Flagg 1977). Even barrier beach spawning occurs progressively 1ater ponds with an outlet to the sea were from south to north. Alewives spawn used by alewives as spawning grounds from mid-April to mid-May in the (Bigelow and Welsh 1925). Parker River, Massachusetts (Cole et a1 . 1980), and from early May to early The two species spawn once a year June in Maine (Flagg 1977; Libby in spring or early summer in fresh or 1981). In general, spawning of brackish water (Raney and Massmann alewives begins 3 to 4 weeks before 1953). Males enter the mouths of that of blueback herring in the same rivers in which they spawn earlier spawning areas. Spawning peaks of the than females (Cooper 1961; Tyus 1971; two species usually are 2 to 3 weeks Richkus 1974a). Blueback herring do apart (Jones et al. 1978). The not usually swim as far upstream as minimum water temperatures at which alewives do to spawn (Jones et a1 . spawning begins is 10.5 "C in alewives 1978, cited from Loesch 1968, 1969). (Cianci 1969) and 14 OC in blueback Adult alewives move into freshwater herring (Loesch and Lund 1977). Both only during day1 ight hours, primarily species cease spawning when water between 1500 and 1800 h. Be1tz temperature exceeds 27 "C (Loesch (1975) surmised that light acts as a 1969; Edsall 1970). switch, initiating migration in the morning and halting it in the evening. Blueback herring prefer to spawn in fast currents over hard substrate Downst ream movement of adult (Loesch and Lund 1977). In contrast, alewives after spawning is not a ran- alewives select a wide variety of dom event, nor does it depend on the spawning sites, using standing water length of time spent on the spawning and oxbows, as we1 1 as mid-ri ver sites grounds. Downst ream movement appar- (Kissi 1 1974). Several investigative ently is triggered by an increase in studies have described alewife spawning in ponds that have an open Yo1 k-Sac Larvae connection with the ocean (Havey 1973; Kissil 1974). The spawning of Total length (TL) at hatching is anadromous alewives and blueback 2.5 to 5.0 mn and averages 5.1 mn at herring is spatially and temporally yo1 k-sac absorption (Mansueti 1962; different . Norden 1967). Duration of the absorption stage is 2 to 5 days for The spawning behavior of blueback alewives and 2 to 3 days for blueback herring was described by Loesch and herring (Mansueti 1962; Cianci 1969). Lund (1977). A spawning group com- posed of one female and several males Larvae swam in circles for several minutes. The males occasionally nudged the vent The larval stage lasts from of the female. Swi ming speed yo1k-sac absorption unti1 transfor- increased gradually unti 1 a deep dive mation to the juvenile stage. Detai led drawings of the developmental occurred. Eggs and mi 1t were released stages of eggs, yolk-sac larvae, and simultaneously near the bottom. Both larvae of both alewives and blueback species spawn mostly at night (Graham herring were pub1 ished by Jones et a1 1956; Edsall 1964). Both male and . (1978). female blueback herring migrate rapidly downstream after spawning. Alewife larvae in two ponds in Few fish stay in the spawning stream the Parker River, Massachusetts, longer than 5 days (Cooper 1961; showed a diurnal change in depth Loesch and Lund 1977). distribution and a random pattern of lateral distribution (Cole et al. 1980). Larval Alosa in Nova Scotian rivers inhabited waters that were relatively shallow (<2 m), sandy, and Unti1 water-hardened, eggs of warm, and were collected in or near both species are demersal in still the spawning grounds (0' Nei 11 1980). water and adhesive or pelagic in running water (Loesch and Lund 1977; Jones et al. 1978). After Ju veni 1es water-hardening (less than 24 h), eggs lose their adhesive property and drift Transformation from the larval to in the water column. Egg diameter the juvenile stage is usually complete ranges from 0.8 to 1.3 mn in alewives when the fish is about 20 mm TL. and from 0.8 to 1.1 mm in blueback Scales first appear on juveniles 25 to herring (Mansueti 1962; Norden 1967). 29 mm TL, and are fully developed at Incubation of blueback herring eggs is 45 mn TL (Hi ldebrand 1963). about 80 to 94 h at 20 to 21 OC and 55 to 58 h at 22 to 24 OC (Cianci 1969; Juveni le alewives and blueback Morgan and Prince 1976). herring in rivers tend to move upstream between June and the start of emi grat ion in October (Bu rbidge 1974; An equation for predicting Warinner et al. 1969). This upstream incubation time for alewife eggs from movement may partly involve juveni les water temperature is from oxbows and tributaries, which gradually move into the main river during summer (Bu rbidge 1974).

Ju veni 1e blueback herring in the where T = time in days and t = James River, Virginia, were more incubation temperature in degrees concentrated near the surface than at Fahrenheit (Edsall 1970). a depth of 5 m throughout their stay in the river (Burbidge 1974). Juvenile Catch data from National Marine alewives in the Potomac River were Fisheries Service trawl surveys along concentrated in the surface waters in the Atlantic coast between Cape September, and in October became more Hatteras and Nova Scotia were abundant at depths of 4.6 m or near summarized annually from 1963 to 1978 the bottom (Warinner et a1 . 1969). (Neves 1981). Samples of fish were taken at depths down to 200 m. Most alewi ves and blueback herring were In most Atlantic coastal popula- caught in water less than 100 m deep. tions, young-of -the-year alewi ves and Average fork lengths (FL) of alewives blueback herring emigrate from and blueback herring ranged from 60 to freshwater and brackish estuaries 350 mm; a1 ewi ves outnumbered blueback between June and November (Burbidge herring about 10:l. Alewives were 1974; Kissil 1974; Richkus 1975; most abundant at depths of 56 to 110 OINeill 1980). Juvenile alewives 32 m, and blueback herring at depths of to 152 mm TL emigrated from Maine 27 to 55 m. Trawl catches over a wide rivers between mid-July and early area indicated that both species were December (Flagg 1977). Their size concentrated in summer and fall in an reportedly depends on the avai 1abi 1i ty area north of 40° N, in or near of food in the river, the total number Nantucket Shoals, Georges Bank, and of young produced in the watershed, the perimeter of the Gulf of Maine. and the length of time in freshwater Winter catches were heaviest between (Flagg 1977). latitudes 40° and 43O N. Spring catches demonst rated that both species Factors initiating emigration were distributed over the entire "waves" (Richkus 1975) that consist of Continental Shelf (Neves 1981). large schools of juvenile alewi ves are: heavy rainfall (Cooper 1961), high water levels (Kissil 1974; In July and October 1964 (the Richkus 1975), and sharp drops in only months sampled), adult alewi ves water temperature (Ri chkus 1975). and blueback herring inhabited only a Richkus (1975) offered three small portion of Georges Bank -- observations: (1) These waves lasted specifically the western slope near two to three days, regardless of the 41° 29' north latitude and 68O 34' duration of the environmental changes ; west longitude (Netzel and Stanek (2) migrations peaked in late 1966). A1 ewi ves outnumbered blueback afternoon; and (3) the magnitude of a herring in these samples. All mature wave was not related to the magnitude age groups were represented, but no of environmental change. About 70% of fish of ages I or I1 were captured. the ju veni les completed emigration in only a few days. Alewives and blueback herring, Adults like other clupeids, are reported to exhi bit seasonal movements in con- junction with changes in water Data on the habits and biology of temperature or avai 1abi 1i ty of forage adult alewife and blueback herri ng (Coll ins 1952; Leggett and Whitney stocks are scarce. More research is 1972); however, sound evidence for needed for identifying inshore and this conclusion is lacking (Richkus offshore stocks, calculating natural 1974b). Feeding and vertical and fishing mortalities, and mi grati on are probably regulated describing movements, migratory largely by 1ight intensity and water patterns, f eedi ng behaviors , and temperature (Richkus and Winn 1979; habitat preferences. Neves 1981). GROWTH CHARACTERISTICS Male blueback: log W=2.904 x log L-4.702 Growth Rates Female blueback: log W=2.472 x log L-3.693 Little is known about growth L = fork length (mm), W = weight (g) rates of either the alewife or blueback herring from the time they hatch to the time they spawn. THE FISHERY Estimates of daily growth of juvenile alewives in two spawning ponds in the Commercial Fisheries Parker River drai nage, Massachusetts, ranged from 0.2 to 0.5 mm (Cole et a1 . Commerci a1 1andings of river 1980). herring (both species combi ned) a1 ong the Atlantic coast of the United Average 1engths of juveni 1e States were 3,783 t .in 1981 and 5,682 emi grants, sampled over three years t in 1982. These landings were worth (1970-72) in Hamilton Reservoir, Rhode $67 1,000 and $1,021,000, respect ively. Island (Richkus 1975), ranged from 25 The 5-year running average of the U.S. to 88 mm SL (30 to 105 mm TL). river herring 1andings (1977-82) was Year1 ing male alewives averaged 147 mm 5,037 tlyr. More than 90% of the U.S. TL by the end of their second summer commercial catch was made within 4.8 in the Connecticut River Estuary km of the coast1 ine (National Marine (Marcy 1969). On Georges Bank, fish Fisheries Service 1983). Commercial of age group 11 of both species were fishing for alewives is primarily about 180 mm TL by the end of their during spawning runs. Weirs and trap third summer (Netzel and Stanek 1966). nets are the most commonly used gears in North Atlantic rivers (Flagg 1977), Mature alewives are usual ly whereas pound nets are commonly used longer than blueback herring of the in the Chesapeake Bay area (Joseph and same age (Table 1). In both species, Davis 1965; Pate 1974). More than 90% males are smaller than females of the of the annual Maine harvest is used as same age. Growth rates for both lobster bait. The rest is used for species level off after reaching trawl bait, human consumption, and for sexual maturity (compared to growth fish meal as a protein supplement in rates of immature fish). Mean weights animal feed. Roe from these species of spawning alewives in Damariscotta is canned and is highly valued (Joseph Lake, Maine, ranged from 153 g (males) and Davis 1965; Street and Davis 1976; and 164 g (females) for age group 111, Merriner 1978). to 325 g (males) and 356 g (females) for age group VII. One female in age The foreign catch of river group VIII weighed 455 g (Walton herring (both species) within the U.S. 1979). Fishery Conservation Zone (FCZ) was 13.9 t in 1981 and 2.0 t in 1982. About 75% of the foreign catch in 1981 Length-Wei ght Re1ati ons was taken by fishermen from Poland; in 1982 fishermen from Italy took the Length-weight relations for largest share (75%). All of the 1981 alewives and blueback herring of the and 1982 foreign catch was taken north St. John River, New Brunswick, were of Cape Hatteras (NMFS 1983). gi ven by Messieh (1977) as follows: The State of Maine has developed Male alewives: an Alewife Management Plan for ana- log W=3.235 x log L-5.420 dromous stocks under its jurisdiction Female alewives: (Walton et al. 1976). Historical log W=3.192 x log L-5.294 landi ngs and management were reviewed Table 1. Ages and fork lengths (mm) of alewives and blueback herring captured during spawning runs in A1 bemarle Sound, North Carol ina (Pate 1974), Chesapeake Bay (Joseph and Davis 1965), the Connecticut River (Marcy 1969), Georges Bank (Netzel and Stanek 1966), and Damariscotta Lake, Maine (Walton 1979).

Species Age group

Location and sex I11 IV V V I VII VIII IX

Alewife

A1 bemarle Sound male f emal e

Chesapeake Bay ma1 e female

Connecticut River ma1 e female

Georges Bank ma1 eifemal e

Damariscotta Lake ma1 e female

Blueback herring

A1 bemarle Sound male 231 241 245 251 258 270 - f emal e 245 252 258 264 268 280 307

Connecticut River male 258 266 280 286 298 - - female 261 277 291 301 311 - -

Georges Bank maleifemale 240 269 281 292 302 313 - and new management recomnendations Love Lake, Maine, were reported by prepared for the alewife runs of each Havey (1973). The number of juvenile coastal county. Catch statistics for emigrants ranged from less than 1,000 each New England State from 1977 to (i.e., 220) in 1962 to 439,000 in 1982 are given in Table 2. 1967. Juveni le emigrants produced per spawning female per year numbered from Popul ati on Dynami cs 12 to 3,209. There were significant 1i near re1ati ons between juveni le Sex ratios/age structure. The emigrant abundance and spawning sex ratio of spawning adults is near population size 4 years later, and 1: 1 in most waters. The percentage of between the log of female escapement males in spawning populations was 56% and the log of juvenile emigrant for alewives in Bride Lake, abundance. The annual adult mortality Connecticut (Kissi 1 1974), 54% for in freshwater averaged 91% and ranged a1 ewi ves and blueback herring in the from 66 to 100% (Havey 1973). The Connecticut River, and 53% (Marcy annual mortal ity for anadromous 1969) and 58% (Loesch and Lund 1977) alewives in Long Pond, Maine was 79% for blueback herring in the Thames between ages V and VI, and 74% between River. Because males tend to mature ages VI and VII (Havey 1961). Fresh- at a younger age than females, there water post-spawni ng mortality for a1 1 is a slight dominance of males in ages in 1954-59 averaged 41% and spawning populations (Kissi 1 1974). ranged from 32% to 67% (Havey 1961). The percentages of males (by different Results from these two studies in age groups) in the 1966 and 1967 Maine and the study by Kissil (1974) spawning populations of a1 ewi ves from in Connecticut indicated that juvenile the Connecticut and Thames rivers , production and adult freshwater Connecticut (data combined) , were 72% mortality may vary considerably among (age group IV), 64% (V), 50% (VI), 34% spawning areas and among different (VII), and 0% (VIII). The percentages years in the same spawning area. of blueback herring males by age group were: 80% (age group 111), 79% (IV), Spawning stocks. A1 though 65% (V), 37% (VI), and 23% (VII) Thunberg (1911) found that olfaction (Marcy 1969). was the basis for homing behavior in alewives, Messieh (1977) reported Abundance and mortality. In considerable mixing among spawning Bride Lake. Connecticut. in 1966. an stocks during the spawning runs. estimated -184,000 adult alew>ves Although most of the alewives homed, a spawned 20.5 billion eggs. About substantial number (determined by 257,000 juveniles were counted as they meristic comparisons) from each emigrated seaward during summer and presumed stock did not (Messieh 1977). fa11. The freshwater mortal ity rate from egg stage to emigration in this Evidence of a nonl andlocked, lake was 99.99%, indicating that about nonmigratory, self-sustaining dwarf three juveniles left the lake for each population of alewives living in the adult female that spawned. Since some mouth of the Susquehanna River was repeat spawning occurs, this level of reported by ~oerster and Goodbred juvenile production seems to be (1978). adequate for sustaining the spawning population in Bride Lake (Kissil 1974). The mortality of spawning ECOLOGICAL ROLE adults in Bride Lake was 57% in 1966 and 49% in 1967. Food Habits

Juvenile production and adult Alewives and blueback herring mortality of the alewife population in feed primarily on zooplankton. Other Tab1 e 2. Annual a1 ewi fe/bl ueback herring 1andi ngs (thousands of pounds) and value (thousands of do1 lars) in New England from 1977 to 1982 (NMFS, unpublished data).

Conn. R. I. Mass. N. H. Maine Year Wt. Value Wt. Value Wt. Value Wt. Value Wt. Value

foods, especially for the larger fish, River showed a strong electivityl for are small fish and insects, and the members of the family Bosminidae, and eggs of fish, insects, and crustaceans moderate electi vity for daphni ds and (Bigelow and Schroeder 1953). Larvae cal anoi d and cyclopoi d copepods begin feeding on relatively small (Domermuth and Reed 1980) . cladocerans and copepods immediately Cladocerans made up 47% of the food after formation of a functional mouth volume (electivi ty was neutral ) . (about 6 mn TL). As they grow, the Copepods made up only 18% of the food larvae eat larger species of these volume and showed strong positive zooplankters (Norden 1968; Ni gro and electi vity . Chi ronomi d 1arvae and Ney 1982). pupae made up 19% and 16% of the food respecti vely, but electivity was strongly negative. Comparisons of these data with similar data for Stomachs from young-of -the-year juvenile American shad (Alosa blueback herring col lected in the and pumpkinseed (Lepomis James River, Virginia, contained that neither blueback primari ly (by volume) the cladocerans, Bosmi na spp., copepod naup 1i i , copepodi tes , and the adult copepods ~urytemora affi nis and Cyclops 'lvlevl s electivity index compares vernal is. Di aphanosoma brachyurum and re1ati ve abundance (percentage of the Canthocamptus robertcokeri we re composition) of a food item in a fish minor food items. Young-of-the-year stomach sample with the relative alewi ves in Hami lton Reservoi r, Rhode abundance of that item in the feeding Island, ate primarily chi ronomid environment. For example, if a food midges in July and cladocerans in item makes up 10% of the food in a August and September (Vigerstad and fish's stomach, but 90% of the Cobb 1978). available food, its electivity index is strongly negative. Electivity is neutral when the proportion of a food Studies of the food of juvenile item eaten is directly proportional to bjueback herring in the Connecticut its avai labi 1ity in the environment. herring nor American shad competed Predators with the benthic-feedi ng pumpkinseed, and that competition between blueback Alewives and blueback herring herring and American shad was contribute substantially to the food inconsequenti a1 (Domermuth and Reed of many riverine, estuarine, and 1980). marine fishes as well as gulls and terns (Commonwealth of Massachusetts Little is known about the food of 1976). Major Dredators are bluefish anadromous adu 1t alewi ves . General ly omat atom us - saltatrix), weakfish they are zooplankti vores, eating (Cynoscion regalis),d striped bass larger and more diverse prey as they (Morone saxati lus) . These pelagic, grow. Large landlocked alewives tend school ing predators commonly prey on to be piscivorous (Kohler and Ney schooling clupeids (Cooper 1961; Tyus 1981). 1974).

Feeding Behavior ENVIRONMENTAL REQUIREMENTS In laboratory tests, alewives showed three feeding modes: (1) Some research has been conducted particulate feeding on individual prey to delineate the specific environ- organism, (2) filter feeding (mouth mental requi rements or preferences of agape during rapid bursts of anadromous a1 ewi ves and blueback swimming), and (3) gulping of several herring. However, much of the prey organisms at once (but not avai lab1e informat ion on a1 ewi ves was swimming at the rapid speed used in derived from tests on landlocked filter-feeding) (Janssen 1976). Size populations, particularly in Lake selectivity of prey was highest in Michi gan. The differences in particulate feeding, moderate in prey envi ronmental requ irements of gulping, and negligible in filter 1and1 ocked populations and anadromous feedinq. Adult Lake Michiqan alewives populations is unknown. Since many of and their major food, ~~s';'srelicta, the available data are from studies of mak e coincidental. cre~uscular landlocked populations, the following vertical migrations, from nea;. bottom envi ronmental requi rements refer during daylight to just below the largely to them. thermocl ine at night (Janssen and Brandt 1980). This die1 vertical Temperature migration, not necessarily related to the thermocline, also may occur in The effects of water temperature anadromous populations 1i vi ng in on the incubation of alewife eggs from estuaries or marine coastal habitats. Lake Michi gan were studied by Edsal 1 (1970). Eggs hatched at test temperatures between 7 and 29.5 OC. At the Cornpeti tors optimm temperature for hatching (18 OC) 38% of the eggs hatched. Egg The degree of compet it ion between mortality over the first 36 h ranged anadromous alewives and blueback from 22% (at temperatures of 3.5 herring is not known. Because of to 6.0 OC) to 66% (at temperatures of general similarities in their diets 25.5 to 28.5 OC). Egg mortality was and feeding behavior, competition for directly correlated with incubation food is probable. Spatial separation temperature (Edsall 1970). An upper between young alewives and blueback lethal temperature of 29.7 OC was herring in the same habitat, which may reported for alewife eggs from the lead to reduced competition for food Hudson River, New York (Kel logg 1982). among juveniles, was reported by Most eggs hatched at a water Loesch et a1 . (1982a). temperature of 20.8 OC; only a few hatched at temperatures of 12.7 and tenperature at which experimental fish 26.1 OC. lose equilibrium) of 28.3 OC, 32.7 OC, and 34.4 OC at acclimation tempera- Blueback herring eggs col lected tures of 11 OC, 19 OC, and 25 OC, from the Washademoak Lake, Canaan respectively (Otto et al. 1976). The River, New Brunswi ck , Canada, were equation for predicting CTM from time-temperature tested in a power acclimation tenperature was CTM = 21.9 plant cooling system (Koo and Johnston + 0.5(TA), where TA = acclimation 1978). These tests proved that larval temperature in degrees Celsius deformity rates were better than egg (correlation coefficient r2 = 0.96). mortalities as indicators of the effects of temperature change. For alewives tested at the same Deformity rates of larvae, acclimated accl imation temperatures, CTM values at 19 OC and exposed to 20 OC water were 3 to 6 OC higher for for 5 to 180 min, ranged from 0 to 25% young-of-the-year than for adults. (control 0-5%) . Deformity rate (Otto et al. 1976). increased to 100% when the temperature was increased to 34 OC. Deformities In laboratory tests, preferred ranged from minor curvature of the (selected) water temperatures of spine to completely abnormal larval juvenile alewives and blueback herring form or behavior. Deformities were (age groups 0 and I) collected from permanent and few such larvae would the Delaware River, New Jersey, and have survived in natural environments accl imated to water temperatures of 15 (Koo and Johnston 1978). to 21 OC ranged from 20 to 22 OC at salinities of 4 to 6 ppt (Meldrim and Two effects of temperature on Gift 1971). Davis and Cheek (1966) larval alewives from Lake Michigan captured juvenile blueback herring in were reported by Edsall (1970). the Cape Fear River seasonally in Survival time of unfed larvae was 3.8 areas where water temperatures ranged from 11.5 to 32 OC. Juvenile alewives days at 10.5 OC, 7.6 days at 14.5 to in the same watershed were captured at 15 OC, and 2.4 days at 26.5 to 28 OC. A functional jaw did not develop in water temperatures of 13.5 to 29 OC. fish from eggs or larvae kept at or below 10 OC, even though some eggs School formation patterns and hatched at those temperatures. An daily rhythms of adult alewives in upper temperature tolerance of 31 OC Lake Michigan were affected by changes for alewife larvae from the Hudson in water temperature in laboratory River, New York, acclimated to 14 OC, tanks (Colby 1971). As water was reported by Kellogg (1982). temperatu re dropped be1 ow 6.7 OC, Average daily gain in larval weight normal feeding behavior was disrupted was di rectly proportional to water and cruising speed of schooling fish temperature. The maximum larval decreased. Below 4.5 OC, normal growth rate was 0.084 g/day at 29.1 school ing behavior was significantly OC; net gain in biomass (a function of reduced. At temperatures between 2.0 survival and growth) was highest at and 2.8 OC, alewives lost orientation, 26.4 OC (Kel logg 1982). swam into the sides of the test chamber, and ceased feeding and Young-of-the-year alewives (19 to schooling. 31 mm TL) from the Hudson River, New York, prefer water temperatures near In laboratory cold shock tests 26 OC when given a choice in a with adult alewives from Lake Michi- control 1ed thermal gradient (Kel 1ogg gan, transfers to temperatures of less 1982). Young-of -the-year alewi ves than 3 OC caused 100% mortality, from Lake Michigan exhibited critical regardless of accl imat ion temperatures thermal maxima (CTM; the mean (Otto et al. 1976). The magnitude of the temperature decrease tolerated by with strong currents and hard sub- alewi ves increased gradual ly with strates (Loesch and Lund 1977). Their increasing accl imation temperature. spawning requirements are thus more Some alewives survived a temperature speci a1 ized than those of a1 ewi ves, decrease of 10 OC, regardless of which use a wide variety of spawning acclimation temperature, if the tem- sites. Alewives may spawn in standing perature did not drop below 3 OC (Otto river water, oxbows, coastal ponds, et al. 1976). small streams, or fast mid-river currents. The electrolyte balance and osmoregul ation of alewives in Lake Michigan changed with a change in Young-of -the-year alewives and temperature (Stanley and Col by 1971). blueback herring from the Cape Fear Transfer of fish acclimated at high Rivery North Carol inay were abundant temperatures to low cold temperatures in water where free carbon dioxide caused levels of Na+, K+, and Ca++ in ranged from 4 to 22 ppm, alkalinity blood and muscle to move toward an from 5 to 32 ppm, dissolved oxygen equilibrium with the salinity of the from 2.4 to 10.0 ppm, and pH from 5.2 accl imat ion envi ronment ( body to 6.8 (Davis and Cheek 1966). concentrations increased in saltwater and decreased in freshwater). An experiment by Schubel and Wang The fish temporarily lost their (1973), designed to test the effects abi 1i ty to osmoregul ate when exposed of suspended sediments on the hatchi ng to cold, regardless of the salinity of of a1 ewife eggs, was terminated the water. because a naturally occurring fungus in the sediment infected all test eggs Salinity before they hatched. Although the extent of infection may have been A 1though supporting data are enhanced by 1aboratory conditions , the sparse, anadromous alewives and attempt indicated that high levels of blueback herring are highly tolerant suspended sediment during or after of salinity changes (Cooper 1961; spawning may cause high excessive Chittenden 1972). No mortality of mortality of eggs in some natural adult blueback herring from either waters. In contrast, suspended gradual or abrupt changes in salinity, sediments in concentrations of 100 ppm including di rect transfers from fresh- or less produced no significant effect water to saltwater and vice versa, was on egg mortality in either of alewives observed by Chittenden (1972). Blood or blueback herring (Auld arid Schubel and muscle concentrations of Na+, K+, 1978). and Ca++ were similar in fish kept in seawater and freshwater of the same temperature -- indi cating that, after Blood 1act ic acid concent rations , a period of accl imati on, alewi ves were measured in alewives moving through a efficient osmoregul ators in either pool -and-wei r f ishway, indicated environment (Stanley and Col by 1971). moderate acti vity and energy expenditure (Domi ny 1971, 1973). The Other Envi ronmental Factors mean levels of blood lactic acid in alewi ves passing through the f ishway The location of appropriate were less than half the levels found spawning sites and substrates is in heavily exercised fish in the important not only to the perpetuation laboratory. The use of resting pools of each species but also for natural along the course of the fishway "reproduct ive segregati on'' between two allowed blood lactic acid levels in otherwise closely similar species. fishes to drop to levels observed in Blueback herring prefer spawning sites rested fish in the laboratory. Periodicity of upstream migratory the James and Chickahominy Rivers, movements of adult alewives through a Virginia (Johnson et al. 1978; Loesch Rhode Island river fishway was et a1 . 1982b). Kepone (in concentra- correlated with the magnitude of tions less than 0.3 ppm) was present incident solar radiation (Saila et al. in young alewives and blueback herring 1972). Richkus (1974a) corroborated from the Mattaponi and Pamunkey this 1i ght-dependent migration; Rivers, Virginia, but was not present however, he also observed that within in detectable quantities in fish from activity patterns determined by 1ight the Rappahannock River, Virginia, or intensity, changes in water the Potomac River, Mary1 and (Loesch et temperature strongly influenced the al. 1982b). specific timing of upstream movement of a1 ewi ves . Ju veni 1e downst ream Entrainment and Turbine Induced emi grat ion from Hami 1ton Reservoi r, Mortal ity Rhode Island, during summer and fall was inhibited somewhat by the sunlight Young alewives and blueback and shade interface present at a herring in the Hudson River are highway bridge at the lower end of the susceptible to power plant entrainment reservoir. More fish passed under for up to 63 days after spawning this bridge on cloudy than on sunny (older ones usual ly escape) (Boreman days (Ri chkus 1974a). et al. 1981). Entrainment mortality calculated for three power plants on Environmental Contaminants the river was 3.5 to 4.1% in 1974 and 6.1 to 11.2% in 1975. Juveniles had The LC (concentration needed to higher mortality than eggs and kill 50% o?Othe test fish) of total sac-f ry . residual chlorine for blueback herring eggs ranged from 0.20 to 0.32 ppm. A1 1 The mortal ity of blueback herring larvae from eggs exposed to sublethal passing through the 17 MW Kaplan concentrations of total residual turbine at Holyoke Dam, Massachusetts, chlorine were deformed (Morgan and varied with the level of power gener- Prince 1977). Concent rations of ation (Knapp et al. 1982). Mortality Kepone greater than 0.3 ppm (termed was lowest when the operating eff ici- the "action level" for possible clo- ency of the turbine was highest. sure of a fishery) were found in body Mortality ranged from 62% at an output tissues of young-of-the-year alewives of 16.5 MW to 82% at outputs of 12.0 and blueback herring collected from and 5.5 MW. LITERATURE CITED

Auld, A. H., and J. R. Schubel. 1978. herring in the James River. Effects of suspended sediment on ~irginia. Trans. Am. Fish. Soc. fish eggs and larvae: a 103: 297-311. 1aboratory assessment. Estuarine Coastal Mar. Sci. 6: 153-164. Chambers, J. R., J. A. Muscik, and J. Davis. 1976. Methods of Beltz, J. R., 1975. Movement and distinguishing larval alewife behavior of adult anadromous from larval blueback herring. alewives, Alosa seudoharen us Chesapeake Sci . 17: 93-100. (Wilson) inthe g_S;arker River ~assachusetts. M.S. ~hesis Chittenden, M. E. 1972. Salinity Univ. Massachusetts, Amherst. 66 tolerance of young blueback PP herring. Trans. Am. Fish. Soc. 101: 123-125. Berry, F. H. 1964. Review and emendat ion of f ami ly C 1upei dae. Copeia 1964: 720-730. Cianci, J. M. 1969. Larval development of the alewife and the glut Bigelow, H. B., and W. C. Schroeder. herring. M.S. Thesis. University 1953. Fishes of the Gulf of of Connecticut, Storrs. 62 pp. Maine. U.S. Fish Wildl. Serv. Fish. Bull. No. 74. 577 pp. Cole, C. F., R. J. Essig, and 0. R. Sainelle. 1980. Biological Bigelow, H. B., and W. W. Welsh. investigation of the alewife 1925. Fishes of the Gulf of population, Parker River, Maine. U. S. Bur. Fish. Bull. No. Massachusetts. U.S. Natl. Mar. 40. 567 pp. Fish. Serv. Final Rep. No. 11, 884-32-50-80-CR. 31 pp. Boreman, J., L. W. Barnthouse, 0. S. Vaughan, C. P. Goodyear, S. W. Colby, P. J. 1971. Alewife dieoffs: Christenson, K. D. Kumar, B. L. why do they occur? Limnos 4(2): Kirk, and W. Vanwinkle. 1981. 18-27. The im~act of entrainment and inpingement on fish populations in the Hudson River Estuary. Collins, G. B. 1952. Factors Vol. I. Ent rainment inpact influencing the orientation of estimates for six fish migrating anadromous fishes. U.S. populations inhabiting the Hudson Fish Wildl. Serv. Fish. Bull. 73: River Estuary. ORNL/NUREC/- 37 3-396. TM-385/VI and NUREC/CR-2220. Oak Ridge National Laboratory, Oak Commonwealth of Massachusetts. 1976. Ridge, Tenn. 416 pp. Anadromous fish of Massachusetts. Mass. Dep. Fish., Wildl. and Burbidge, R. G. 1974. Distribution, Recreational Vehicles. Oiv. Mar. growth, selecti ve feedi ng and Fish. Proj. Rep. No. AFCS-14-1, energy t ransformat ions of Public information pamphlet. 20 young-of-the-year blueback PP Cooper, R. A. 1961. Early life Foerster, J. W., and S. L. Goodbred. history and spawning migration 1978. Evidence for a resident of the alewife. M.S. Thesis. alewife population in the Uni versity of Rhode Is1and, northern Chesapeake Bay. Kingston. 58 pp. Estuarine Coastal Mar. Sci. 7: 437 -444. Davis, J. R., and R. P. Cheek. 1966. Distribution, food habits and Graham, J. J. 1956. Observations on the alewife in freshwater. Univ. growth of young clupeids, Cape Toronto Biol. Ser. No. 62. 43 Fear- River system, North Carolina. Paper presented at PP Annu. Meet. South. Div. Am. Fish. Havey, K. A. 1961. Restoration of Soc., Asheville, N.C., 1966. 20 anadromous alewives at Long PP Pond, Maine. Trans. Am. Fish. SOC. 90: 281-286. Dominy, C. L. 1971. Changes in blood 1act ic acid concentrations in Havey, K. A. 1973. Production of alewives during passage through a juvenile alewives at Love Lake, pool-and-weir f ishway. J. Fish. Washington County, Maine. Trans. Res . Board Can. 28: 1215-1217. Am. Fish. Soc. 102: 434-437. Dominy, C. L. 1973. Effect of Hildebrand, S. F. 1963. Fami ly entrance pool wei r elevation and Clupeidae. Pages 257 -385, fish density on passage of 397-442, and 452-454 in Fishes of alewi ves in a pool -and-wei r the western North Atlantic. Sears fishway. Trans. Am. Fish. Soc. Found. Mar. Res. Mem. l(3). 102: 398-404. Huber, M. E. 1978. Adult spawning success and emigration of Domermuth, R. B., and R. J. Reed. juveni le a 1ewi ves (Alosa 1980. Food of iuvenile American pseudoharengus) from the Parker shad, Alosa sapiiissima, juvenile River, Massachusetts. M.S. bluebar herring, Alosa Thesis. Uni v. Massachusetts, aesti val is, and pumpkinseed, Amherst. 75 pp. Lepomi s gibbosus, in the Connecticut R iver be1 ow Hol yoke Janssen, J. 1976. Feeding modes and Dam, Massachusetts. ~stua'ries prey size selection in the 3(1): 65-68. alewife. J. Fish. Res. Board Can. 33: 1972-1975. Edsall, T. A. 1964. Feeding by three species of fishes on the eggs of Janssen, J., and S. B. Brandt. 1980. spawning alewives. Copeia 1964: Feeding ecology and vert ical 226-227. migration of adult alewives in Lake Michigan. Can. J. Fish. Edsall, T. A. 1970. The effect of Aquat. Sci . 37: 177-184. temperature on the rate of Johnson, H. B., D. W. Crocker, B. F. development and su rvival of Holland, Jr., J. W. Gilliken, D. alewife eggs and larvae. Trans. W. Taylor, M. W. Street, J. G. Am. Fish. Soc. 99:376-380. Loesch, W. H. Kriete, Jr., and J .G. Travelstead. 1978. Biology Flagg, L. N. 1977. The alewife and management of mid-At lantic fishery of Maine. Report to anadromous fishes under extended Department of Marine Resources, jurisdiction. N.C. Div. Mar. State House, Augusta, Maine. 5 Fish. and Va. Inst. Mar. Sci. PP- Rep. NC-VA AFSC 9-2. 175 pp. Jones, P. W., and F. D. Martin, and Leggett, W. C. G., and R. R. Whitney. J.D. Hardy, Jr. 1978. 1972. Water temperature and the Development of fishes of the migrations of the American shad. Mid-Atlantic Bight: an atlas of U.S. Natl. Mar. Fish. Serv. Fish. the egg, larval and juvenile Bull. No. 70. 670 pp. stages, Volume I. U.S. Fish. Wildl . Serv. Biol . Serv. Program Libby, D. A. 1981. Difference in sex FWS/OBS-78/12. 366 pp . ratios of the anadromous alewife, Alosa pseudoharengus, between the top and bottom of a fishway at Joseph, E. B., and J. Davis. 1965. A Damariscotta Lake, Maine. U.S. preliminary assessment of the Natl. Mar. Fish. Serv. Fish. river herring stocks of lower Bull. 79(1):207-271. Chesapeake Bay. Va. Inst. Mar. Sci. Spec. Sci. Rep. No. 51. 23 Loesch, J. G. 1968. A contribution to PP- the life history of Alosa aestivalis. M.S. Thesis. Uni ver- Kellogg, R. L. 1982. Temperature sity of Connecticut, Storrs. 31 requirements for the survival and PP early development of the a nadro- mous alewife. Prog. Fish-Cult. Loesch, J. G. 1969. A study of the 44: 63-73. blueback herring in Connecticut waters. Ph .D. Thesis. University Kissil, G. W. 1974. Spawning of the of Connecticut, Storrs. 78 pp. anadromous alewife in Bride Lake, Connecticut. Trans. Am. Fish. Loesch, J. G., and W. A. Lund. 1977. SOC. 103: 312-317. A contribution to the life history of the blueback herring. Knapp, W. E., B. Kynard, and S. P. Trans. Am. Fish. Soc. 106: Gloss. 1982. Potential effects 583-589. Of Kaplan' Osbergers and Loesch, J.G.,W.H.Kriete,Jr.,and Turbines on anadromous fishes of the Northeast United States. U.S. E. J. Foell. 1982a. Effects of Fish. Wildl. Serv., Biol. Serv. 1 ight intensity on the Program FWS/OBS-82/62. 131 pp. catchabil ity of juveni le Alosa species. Trans. Am. Fish. -Soc. Kohler, C. C., and J. J. Ney. 1981. Consequences of an a1 ewif e dieoff Loesch, 3. G., R. J. Hugget, and E. J. to fish and zooplankton in a Foell. 1982b. Kepone concen- reservoir. Trans. Am. Fish. Soc., trati on in juveni 1e anadromous 110: 360-369. fishes. Estuaries 5(3): 175-181.

Koo, T. S. Y., and M. L. Johnston. MacLellan, P., G. E. Newsome, and 1978. Larval deformity in P.A. Dill. 1981. Discrimination striped bass and blueback herring by external features between due to heat shock treatment of alewife and blueback herring. developing eggs. Environ. Pol lut. Can. J. Fish. Aquat. Sci. 38: 16: 137-149. 544-546.

Kuntz, A., and L. Radcliffe. 1917. Mansueti, R. J. 1962. Eggs, larvae Notes on the embryology and and young of the hickory shad, 1 arval development of twel ve with comments on its ecology in teleostean fishes. U.S. Bur. the estuary. Chesapeake Sci. 3: Fish. Bull. No. 35. 134 pp. 173-205. Marcy, B. C., Jr. 1969. Age alewives to a southern range determi nation from scales of extension. Trans. Am. Fish. Soc. A1 osa pseudoharengus and Alosa 111: 559-569. aesti val is in Connecticut waters. Trans. Am. Fish. Soc. 98: Norden, C. R. 1967. Development and 621-630. identification of the larval Meldrim, J. W., and J. J. Gift. 1971. alewife in Lake Michigan. Proc. Temperature preference, avoidance 10th Conf. Great Lakes Res.: and shock experiments with 70-78. estuarine f ishs. Ichthyol. Assoc. Bull. No. 7. 75 pp. Norden, C. R. 1968. Morphology and food habits of the larval alewife Merriner, J. V. 1978. Anadromous in Lake Michigan. Proc. 11th fishes of the Potomac Estuary. Conf. Great Lakes. Res. 103-110. Va. Inst. Mar. Sci. Contrib. 696, Gloucester Point. 4 pp. O'Neill, J. T. 1980. Aspects of the 1 ife histories of anadromous Messieh, S. N. 1977. Population alewife and the blueback herring, structure and biology of alewives Margaree River and Lake Ainsle, and blueback herring in the Saint Nova Scotia, 1978-1979. M. S. John River, New Brunswick. Thesis. Acadi a Uni versi ty , Envi ron. Biol. Fishes 2(3): Wolfville, Nova Scotia, Canada. 195-210. 306 pp. Morgan, R. P., 11, and R. D. Prince. 1976. Ch 1ori ne toxicity to Otto, R. G., M. A. Kitchel, and J. 0. estuarine fish eggs and larvae. Rice. 1976. Lethal and Chesapeake Biol. Lab. Univ. preferred temperatures of the Mary1 and Cent. Envi ron. Estuarine alewife in Lake Michigan. Trans. Stud. Ref. No. 76-116 CBL. 122 Am. Fish. Soc. 105:96-106. PP Pardue, G.B. 1983. Habitat suitabi l- Morgan, R. P., 11, and R. D. Prince. ity index models: alewife and 1977. Chlorine toxi city to eggs blueback herring. U.S. Fish and larvae of five Chesapeake Bay Wi1 dl . Serv. FWS/OBS-82/10.58. fishes. Trans. Am. Fish. Soc. 22 PP 106: 380-385. National Marine Fisheries Service Pate, P. P. 1974. Age and size (NMFS). 1983. Fisheries of the comosition of commercial catches United States. 1982. U.S. Dep. of blueback herring in A1 bemarl e Commer., Curr. Fish. Stat. No. Sound, North Carolina, and its 8300. 117 pp. tributaries. Rep N. C. Dep. Nat. Econ. Resour., Div. Comner. Netzel , J., and E. Stanek. 1966. Some Sport Fish. 10 pp. biological characteristics of blueback and alewife from Georges Raney, E. C., and W. H. Massmann. Bank, July and October, 1964. 1953. The fishes of the ICNAF Res. Bull. No. 3. 5 pp. tidewater section of the Parnunkey River, Virginia. J. Wash. Acad. Neves, R. J. 1981. Offshore distri- Sci 43(12): 424-432. bution of alewife and blueback . herring along the Atlantic coast. Richkus, W. A. 1974a. Factors U.S. Natl. Mar. Fish. Serv. Fish. influencing the seasonal and Bull. 79: 473-485. daily patterns of alewife Nigro, A. A., and J. J. Ney. 1982. migration in a Rhode Island Reproduction and early-life River. J. Fish. Res. Board Can. accommodations of 1and1 ocked 31: 1485-1497. Richkus, W. A. 1974b. Influence of fishery of SA6. ICNAF Res. Doc. environmental variables on the 76/VI/61. 7 pp. migratory behavior of adult and juvenile alewives. Ph.D. Thesis. Thunberg, B. E. 1971. Olfaction in University of Rhode Island, parent stream selection by the Kingston. 225 pp. alewife. Anim. Behav. 19: 217-225. Richkus, W. A. 1975. Migratory behavior and growth of juveni 1e Tyus, H. M. 1971. Population size, anadromous alewives in a Rhode harvest and movements of Island drainage. Trans. Am. Fish. alewives during spawning mi gra- SOC. 104: 483-493. tions to Lake Mattamuskeet, North Carolina. Ph .D. Thesis. North Richkus, W. A., and H. E. Winn. 1979. Carolina State University, Activity cycles of adult and Ral eigh. juveni le a1 ewi ves, recorded by two methods. Trans. Am. Fish. Tyus, H. M. 1974. Movements and SOC. 108: 385-365. spawning of anadromous alewives at Lake Mattamuskeet, North Saila, S.B., T. T. Polgar. D. J. Carolina, Trans. Am. Fish. Soc. Sheehy, and J. M. Flowers. 1972. 103: 392-396. Correlations between alewife activity and environmental Vigerstad, T. J., and J. S. Cobb. variables at a fishway. Trans. 1978. Effects of predation by Am. Fish. Soc. 101: 583-594. sea-run juvenile alewives on the zooplankton community at Hami lton Schubel, J. R., and J. C. S. Wang. Reservoi r, Rhode Island. Estuaries l(1): 36-45. 1973. The effects of suspended sediments on the hatching success of ye1 low perch, white perch, Walton, C. J. 1979. Growth of Alosa striped bass and alewife eggs. seudoharengus from two Maine Ichthyol. Assoc. Spec. Rep. No. eatersheds. Maine Dep. Mar. 30, Ref. 73-3. 77 pp. Resour. Res. Ref. Doc. No. 79/23. 75 PP-

Scott, W. B., and E. J. Crossman. Walton, C. J., M. E. Smith, and D. 8. 197 3. Freshwater fishes of Sampson. 1976. Alewife management Canada. Fish. Res. Board Can. plan. Maine Dep. Mar. Res. Job Bull. No. 184. 966 pp. Compl . Rep. No. AFSC-13/FWAC-2. 37 PP. Stanley, J. G., and P. J. Colby. 1971. Effects of temperature on Warinner, J. E., J. P. Miller, and J. electrolyte balance and Davis. 1969. Distribution of osmoregulation in the alewife in juvenile river herring in the fresh and sea water. Trans. Am. Potomac River. Proc. Annu. Conf . Fish. Soc. 100:624-638. Southeast. Assoc. Game Fish Comm. 23: 384-388. Street, M. W., 1969. Fecundity of the blueback herring in Georgia. Ga. Winters, G. H., J. A. Moores, and R. Game Fish Comm. Mar. Fish. Div., Chaul k. 1973. Northern range Contrib. Ser. 17. 15 pp. extension and probable spawning of gaspareau in the Newfoundland Street, M. W., and J. Davis. 1976. area. J. Fish. Res. Board Can. Notes on the river herring 30: 860-861. 10172 .I01 --- '- I 3. Iocwasnl'a ACCI.VO~ NQ I REPORT WUMENTATION I I. RimRrNo. I I. I 1 Bioloqical Report 82(1156)* 1 -- I L noon p.10 Species Profiles: Life Histories and Environmen$al Requirements of July 1986 Coastal Fishes and Invertebrates (North Atlantic)--Alewife/Blue- -6 back Herring 7. AdhoH.1 Dennis M. Mullen, Clemon W. Fay, and John R. Moring 9. hrlormm. OI.on8rotnon Norno .nd Addnls Maine Cooperative Fishery Research Unit 313 Murray Hall, University of Maine --- Orono, ME 04469 (a

National idetland; Research Center U.S. Army Corps of Engineers Fish and Wildlife Service Waterways Experiment Station U.S. Department of the Interior P.O. Box 631 Washington, DC 20240 Vicksburg, MS 39180

I 1% -0(onun101y Nd.. *U.S. Army Corps of Engineers Report No. TR EL-82-4.

I6 *k(~(Urnlt I#) reds) Species profiles are literature summaries of the taxonomy, morphology, range, life history, and environmental requirements of coastal aquatic species. They are prepared to assist in impact assessment. The alewife and blueback herring (Alosa pseudoharengus and Alosa aestivalis) are important species in estuarine and marine ecosystems, though both anadromous and landlocked populations exist. Some individuals mature by age 3, and all mature by age 5. Repeat spawning is common. Spawning environments range from streams only a few centimeters deep to large rivers. Alewives will also spawn in ponds with an open connection to the sea. Blueback herring prefer spawning sites with fast currents and associated hard substrates, while alewives select a wider variety of sites, from standing water and oxbows to mid-river areas. Spawning occurs from April to July in the north Atlantic region;'t.he onset and peak of alewife spawning precede those of blueback herring by 2 to 3 weeks. Larvae and juveniles remain in or near areas spawned before emigrating (as juveniles) to coastal areas in their first year. Emigration is apparently triggered by heavy runoff from rain and/or sharp decreases in water temperature. Adults overwinter offshore to depths of at least 110 m. Nantucket Shoals, Georges Bank and tile Gulf of Maine are important overwintering grounds. Commercial and limited recreational fisheries for these species occur; total U.S. landings in 1982 were 5,682 t, while foreign landings within the U.S. Fiahery Conservation Zone were 2.0 t. Someoeggs can hatch at water temperatures between 7 and 29.5'~, but tem~eraturesabove 29.7 C are 17- -m-tAnalml* .. o*=n~~t- lethal. Larvae need temperatures greater than 10'~ for proper 'devel opment . Rivers Growth l e~~lperature COII~JIII~ 11a11ts lisltcries Feedillghabits sa~iIlity Life cycles Estuaries Ilerrings Animal migrations

'm! ?I!!%?uonlOD.n.Cndod Torms Alewife A1 osa pseudoharengus Blueback herring Spawning Alosa aestival is

c. COSATI Floldltrrrue

P -. 1L Ava&labllity Slatomont 19. Ssurlfr Class IThas a.mn1 21. wo of roeol Unclassified 21 , ------Unlimited Release 2U. Scurstl Class IThm Panel 22. hco Unclassified So# ANSIY9.1Bl O.?IO:.AL FORU 211 (r-771 ,rar,v.olv Ml$S-351 r:,, , -,.. ... REGION 1 REGION 2 REGION 3 Regional Director Regional Director Regional Director U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service Lloyd Five Hundred Building, Suite 1692 P.O. Box 1306 Federal Building, Fort Snelling 500 N.E. Multnornah Street Albuquerque, New Mexico 87 103 Twin Cities, Minnesota 55 1 1 1 Portland, Oregon 97232

REGION 4 REGION 5 REGION 6 Regional Director Regional Director Regional Director U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service Richard B. Russell Building One Gateway Center P.O. Box 25486 75 Spring Street, S.W. Newton Corner, Massachusetts 02158 Denver Federal Center Atlanta, Georgia 30303 Denver, Colorado 80225

REGION 7 Regional Director U.S. Fish and Wildlife Service 101 1 E. Tudor Road Anchorage, Alaska 99503 TAKE PRIDE inAmerzcd DEPARTMENT OF THE IWTERIOR U.S. FISH AND WllDLlA SERVICE

As the Nation's principal conservation agency. the Department of the Interior has responsibility for most of our .nationally owned public lands and natural resources. This includes fostering the wisest use of our land and water resourcm, protecting our fish and wildlife, preserving thsenvironmental 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 in the best interests of all our people. The Department also has a major responsibility for American Indian reservation communities and for paople who live in island territories under U.S. administration.