Alewife and Blueback Herring

Home , Alewife (fish)

REFERENCE COPY Do Not Rernwe from the Library U. S. Fish and Wildlife Service Nntbnl W- Re-wrrh Cpn~r Biological Report 82(11.111) 700 Cajun Dome Bou teltsl,rd TR EL-82-4 ~ugust1989 Lafayette, Louisiana -2-SC6

Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (South Atlantic)

ALEWIFE AND 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( 11.111) TR EL-82-4 August 1989

Species Profiles : Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (South Atlantic)

ALEWIFE AND BLUEBACK HERRING

Earl L. Bozeman, Jr. and Michael J. Van Den Avyle Georgia Cooperative Fish and Wild1 ife Research Unit School of Forest Resources University of Georgia Athens, GA 30602

Project Officer David Moran U.S. Fish and Wildlife Service National Metlands Research Center 1010 Gause Boulevard Slidel 1 , LA 70458

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

and

U.S. Department of the Interior Fish and Wildlife Service Research and Development National Wetlands Research Center Washington, DC 20240 This series may be referenced as follows: U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life histories and environmental requirements of coas talfishes and invertebrates. U.S. Fish Wildl . Serv. Biol . Rep. 82(11). U.S. Anny Corps of Engineers, TR EL-82-4. This profile may be cited as follows: Bozeman, E.L., Jr., and M.J. VanDen Avyle. 1989. Species profiles: life histories and environmental requirements of coastal fishes and invertebrates (South Atlantic) --alewife and blueback herring. U.S. Fish Wildl . Serv. Biol . Rep. 82( 11.111). U.S. Anny Corps of Engineers, TR EL-82-4. 17 pp. PREFACE

This species profile is one of a series on coastal aquatic organisms, principal ly 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, 1 ife history, ecological role, environmental requirements, and economic importance, if applicable. A three-ring 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 Attenti on: WESER-C Post Office Box 631 Vicksburg, MS 39180 CONVERSION TABLE

Metric to U.S. Customary

To Obtain millimeters (mm) inches centimeters (cm) inches meters (m) feet meters (m) fathoms kilometers (km) statute mi1 es kilometers (km) nautical miles square meters (m2) 10.76 square feet square kilometers (km2) 0.3861 square miles hectares (ha) 2.471 acres liters (1) gal 1ons cubic meters (m3) cubic feet cubic meters (m3) acre-feet milligrams (mg) ounces grams (g) ounces ki1 ograms (kg) pounds metric tons (t) pounds metric tons (t) short tons kilocalories (kcal) British thermal units Celsius degrees (OC) Fahrenheit degrees

U.S. Customary to Metric inches 25.40 millimeters inches 2.54 centimeters feet (ft) 0.3048 meters fathoms 1.829 meters statute miles (mi) 1.609 kilometers nautical miles (nmi) 1.852 kilometers square feet (ft2) square meters square miles (mi2) square kilometers acres hectares gallons (gal) 1i ters cubic feet (ft3) cubic meters acre-feet cubic meters ounces (oz) milligrams ounces (oz) grams pounds (lb) ki1 ograms pounds (lb) metric tons short tons (ton) metric tons British thermal units (Btu) kilocalories Fahrenheit degrees (OF) Celsius degrees CONTENTS

PAGE

PREFACE ...... iii CONVERSION TABLE ...... iv ACKNOWLEDGMENTS ...... v i PROFILE SCOPE ...... NOMENCLATURE. TAXONOMY. AND RANGE ...... MORPHOLOGY AND IDENTIFICATION AIDS ...... Alewife ...... Blueback Herring ...... Aids For Species Separation ...... REASON FOR INCLUSION IN SERIES ...... LIFE HISTORY ...... Spawning ...... Eggs ...... Larvae ...... Juveniles ...... Adults ...... GROWTH CHARACTERISTICS ...... Growth Rates ...... THE FISHERY ...... ECOLOGICAL ROLE ...... Food ...... Comoetitors ...... Predators ...... ENVIRONMENTAL REQUIREMENTS...... Temperature ...... Salinity...... Dissolved Oxygen ...... Substrate and System Features ...... Environmental Contaminants ...... LITERATURE CITED ...... ACKNOWLEDGMENTS

We thank Richard Christie of the Dennis Wildlife Research Center, Bonneau, South Carolina, and Sara Winslow of the North Carolina Division of Marine Fisheries, Elizabeth City, for their reviews of this manuscript. We also appreciate Sue Anthony for typing and preliminary editing. Figure 1. A. alewife; 0. blueback herring.

ALEWIFE AND BLUEBACK HERRING

PROFILE SCOPE out most of the South Atlantic Region (Cape Hatteras, North Carol ina, to This profile addresses 1 ife histo- Cape Canaveral, Florida) and is ries and environmental requirements of emphasized here. The alewife is more both a1 ewi fe and blueback herring limited in distribution in the South (Figure 1) because the morphology, At1 antic Region, occurring only in ecological role, and environmental waters of North Carolina and northern requirements of the two species are South Carol ina. Most of the similar. The fish are marketed information available on alewife 1 ife together as "river herring" or history is from studies in the Middle "alewife," and are often combined in and North Atlantic Regions or from commercial fishing statistics. The studies of 1andlocked populations in blueback herring is plentiful through- the Great Lakes. Inasmuch as applic- abi 1i ty of some of these data, particu- Alewife 1arly those describing environmental requirements, to southeastern popul a- Dorsal rays 12-19 (usual ly 13-14) ; tions of the alewife is unknown, this anal rays 15-21 (usually 17-18) ; lat- information should be appl ied with eral line scales 42-54. Prepelvic caution. scutes (modified scales along the ventral mid1 ine) 17-21 (usual 1y NOMENCLATURE, TAXONOMY, AND RANGE 19-20); postelvic scutes 12-17 (usually 14-15 P ; gill rakers on first Scientific names...... Alosa arch 38-46. Body strongly compressed seudoharen us (Wi 1son) and7- and deep. Mouth oblique; anterior end -tchi. --. -. - 11 1 of lower jaw thick, heavy, and extend- Preferred common names...... Alewife ing to middle of orbit. Eye large, and blueback herring (Figure 1). diameter greater than snout length. Other common names (both species). .... Color dorsally gray to gray-green; river herring, glut herring, saw- laterally silver with prominent dark be1 ly , goggle-eye, blackbelly, shoulder spot; fins pale yellow to summer herring, kyak, branch her- green. ring, greyback, oldwife, gaspereau. Class ...... Osteichthyes. Bl ueback Herring Order ...... Clupeiformes. Family ...... Clupeidae. Dorsal rays 15-20; anal rays 15-21; lateral line scales 46-54. Geographic range: The alewife is an Prepelvic scutes 18-21; postpelvic anadromous species occurring in scutes 12-16; gill rakers on first riverine, estuarine, and Atlantic arch 41-52. Body moderately com- coastal waters from Newfound1 and pressed and elongate; eye diameter (Winters et al. 1973) to northern small, equal to or less than snout South Carolina (Berry 1964). length. Upper jaw with definitive Reports of the alewife in Florida median notch; no teeth on premaxil- waters are questionable (McLane laries. Color dorsally blue to 1955; Williams and Grey 1975). The blue-green; 1ateral ly si1 ver with Great Lakes and Finger Lakes contain prominent dark shoulder spot; fins 1and1 ocked populations of the pale yellow to green. species (Bigel ow and Schroeder 1953; Scott and Crossman 1973). The Aids for Species Separation blueback herring is an anadromous species occurring in riverine, w. Unfertil ized alewife eggs estuarine, and At1 antic coastal are green, and blueback herring eggs waters from Nova Scotia (Scott and are amber. Oil droplets of fertilized Crossman 1973) to the St. Johns eggs are numerous and uniformly tiny River, Florida (Hildebrand 1963). in the alewife but are of unequal size Landlocked populations of blueback and scattered in the blueback herring herring occur in coastal plain lakes (Kuntz and Radcliffe 1917; Norden and several southeastern reservoirs. 1968). The coastal distributions of the alewife and blueback herring in the Larvae. The number of myomeres South Atlantic Region are shown in between anal vent and insertion of Figure 2. dorsal fin is 7-9 (mean 8.0) in the alewife and 11-13 (mean 11.8) in the blueback herring (Chambers et al. MORPHOLOGY AND IDENTIFICATION AIDS 1976). The following information was Adults. Adults can be distin- taken from Jones et a1. (1978). guished external 1y by individual scale NORTH CAROLINA

SOUTH CAROLINA

A TL AN TIC OCEAN

Alewife a Blueback herring

MILES

KILOMETERS

Figure 2. Coastal distributions of alewife and bl ueback herring in the South Atlantic Region.

3 markings. Scales come together on the Sholar 1977; Bulak and Curtis 1978; dividing line on alewife but not on Hawkins 1979; Fisher 1980). blueback herring (O'Neill 1980; MacLellan et al. 1981). Although Spawning temperature requirements dorsal coloration has been cited as for either species are poorly defined species-distinctive in fresh specimens because spawning runs often coincide (Bigel ow and Schroeder 1953), and most spawning temperatures have MacLellan et al. (1981) found no been recorded for "river herring" detectable difference and observed rather than for each species separ- that dorsal col oration appeared to ately. Marshal 1 (1977) , however, vary with light conditions. Internal- reported ripe alewives at temperatures ly, the peritoneal lining is pale, of 15-20 "C in the Neuse River, North gray, or silvery with dark punctua- Carol ina. Spawning activity of blue- tions in the alewife and uniformly back herring has been observed at dark in the blueback herring (Leim and temperatures as low as 13 "C in the Scott 1966; Scott and Crossman 1973). Neuse River, North Carol ina (Hawkins The shapes of the otol iths are dis- 1979). Spawning activity peaks at tinctive (Scott and Crossman 1973; 17-19 "C in North Carolina and South Price 1978; O'Neill 1980). Carolina (Sholar 1977, Bulak and Curtis 1978) and at 17-20 "C in Georgia and northern Florida (Street REASON FOR INCLUSION IN SERIES 1970; Williams et al. 1975). Both species cease spawning when tempera- Both the alewife and blueback tures exceed 27 "C (Hawkins 1979). herring have decl ined in commercial importance in the South Atlantic Although a variety of spawning Region over the past 15 years (Rul if- habitats are used by both species, son et al. 1982). They are ecolog- blueback herring prefer shall ow areas ically important species due to their covered with vegetation (Frankensteen trophic level. Both species are 1976), old ricefields (Christie 1978), planktivorous and are important links and river swamps and small tributaries between zooplankton and piscivores in above tidal influence (Godwin and estuarine and marine food webs. Adams 1969; Street 1970). Brackish and tidal areas are rarely used by blueback herring for spawning (Loesch and Lund 1977). In contrast, alewives LIFE HISTORY have been reported spawning in barrier beach ponds (Bigelow and Welsh 1925) and brackish streams (Kissil 1974), as Spawning well as at upstream, mid-river sites (Bigelow and Schroeder 1953); they Alewives and blueback herring spawn over a detritus-covered bottom spawn from late winter to early summer with attached vegetation, sticks, or in the South Atlantic Region. other organic matter and occasional ly Marshall (1977) and Sholar (1975) over a hard sand bottom (Cooper 1961). reported spawning runs of alewives Spawning in both species occurs from mid-March to late May in the diurnally and nocturnally, a1 though Neuse River, North Carol ina. Blueback most activity is nocturnal (Graham herring spawn in the St. Johns River, 1956; Edsall 1964). Bl ueback herring Florida, from January to early May make repeat spawning runs and return to (Williams et a1 . 1975). Blueback her- the same river to spawn. Thus, racial ring spawning runs occur later in the differences may exist between rivers season with increasing 1ati tude, and and management of the fishery may need continue into June in North Carolina to be on a river-by-river basis rivers (Street 1970; Sholar 1975; (Christie 1984). 4 The larval stage (from yolk-sac absorption to transformation into the Fecundity estimates of blueback juvenile stage) lasts 2-3 weeks in herring in the Altamaha River, both species. Larval alewives are Georgia, were 120,000-400,000 e gs per 4.3-19.9 mm standard length (SL), and female , and averaged 244,000 qStreet 1arval blueback herring are 4 .O-15.9 1970). Williams et a1 . (1975) esti- mm SL (Cooper 1961; Jones et al. mated that blueback herring in the St. 1978). Jones et a1 . (1978) presented Johns River, Florida, contained detailed drawings of the developmental 150,,000-349,000 eggs (mean, 262,000). stages of eggs, yolk-sac larvae, and There are no reported fecundity advanced 1arvae of both species. estimates for alewife in the South Atlantic Region. Smith (1907) repor- Juveniles ted that alewives in the Potomac River, Virginia, contained an average Transformation to the juvenile of 102,800 eggs per female, and Kissil stage is completed in both species at (1969) estimated that alewives in about 20 mm TL. Scales first appear Connecticut rivers produced 229,000 when juveniles are 25-29 mm TL and are eggs per female. fully developed at 45 mm TL (Hi 1debrand 1963 ; Norden 1968).

Until water-hardened, eggs of both Nursery areas for juvenile blue- species are adhesive and will sink back herring in the Neuse River, North unless buoyed by river or tidal Carolina, are characterized by deep, currents. Within 24 h after spawning, black water draining hardwood swamps, the eggs lose their adhesive property with little salinity or current and (Loesch and Lund 1977; Jones et al. with a mud or detritus bottom 1978). Fertilized blueback eggs are (Marshall 1977). Juvenile alewife and ye1 1owi sh and have scattered, unequal - blueback herring were present in South sized oi1 drop1ets, whereas a1 ewi fe Creek estuary, North Carolina, in eggs are amber and have numerous small spring (Rulifson 1985). In the South oil droplets (Kuntz and Radcliffe Atlantic Region, juvenile blueback 1917; Norden 1968). Egg diameters are herring remain in primary nursery areas 0.80-1.27 mm in the alewife and 0.87- until October and then begin migrating 1.11mm in the blueback herring to shallow, high-sal inity estuaries for (Mansueti 1956; Norden 1968). overwintering. These secondary Incubation times for blueback herring nurseries are used unti1 year1 ings eggs are 80-94 h at 20-21 'C and 55- migrate to sea in the spring 58 h at 22-24 'C (Cianci 1969; Morgan (Spi tsbergen and Wol ff 1974). and Prince 1976). Comparative incubation times for alewife eggs are Primary nursery areas for alewives 89 h at 21.1 'C (Edsall 1970) and 72 h are the lower reaches of rivers in at 23.8 'C (Kellogg 1982). brackish water or tidally influenced freshwater. Migration patterns of juvenile alewives are not as clearly Larvae defined as those of blueback herring. The fish migrated from primary Yo1 k-sac 1arvae of both species nursery areas in November in the Cape are 2.5-5.0 mm total length (TL) at Fear River, North Carol ina (Shol ar hatching and average 5.0 mm TL at 1977), but juveniles of 24-105 mm TL yo1 k-sac absorption (Mansueti 1956; were captured in freshwater Lake Norden 1968). This stage lasts 2-5 Ma ttamuskeet, North Carolina, during days in the alewife and 2-3 days in June, November, and January, even the blueback herring (Mansueti 1956; though access to coastal areas was Cianci 1969; Jones et al. 1978). maintained at a11 times (Tyus 1972). Juvenile alewives use high-sal inity November (Davis and Cheek 1966). Mean estuaries as secondary nurseries fork lengths of juveniles in the before migrating to sea in winter and A1 tamaha River, Georgia, increased early spring (Holland and Yelverton from 34.8 mm FL in July to 60.6 mm FL 1973). in November, or a 25.8 mm increase over four months (Godwin and Adams Adults 1969). Juvenile alewife in the Neuse River, North Carol ina, increased from Blueback herring and a1 ewives 35 mm FL in June to 75 mm FL in reach sexual maturity by age 111 or IV November (Hawkins 1979), whereas (Loesch 1969) at about 250 mm TL juveniles in the White Oak, Cape Fear, (Johnson et al. 1978). Females of and Northeast Cape Fear Rivers, North both species are larger than males of Carol ina, increased from 47 mm FL in the same age (Williams et al. 1975; July to 81 mm FL in December (Davis Sholar 1977). Blueback herring sex and Cheek 1966; Sholar 1975). ratios (ma1 e:female) in North Carolina ranged from 1:2.80 in the Northeast Hol 1and and Ye1 verton (1973) Cape Fear River (Fischer 1980) to estimated relations between fork 1:0.65 in the Neuse River (Marshall length and age, and fork length and 1977). Corresponding ranges of sex weight for alewives and blueback ratios for the alewife were from 1:3.0 herring from the Chowan River and in the Cape Fear River (Fischer 1980) offshore North Carol ina (Table 1). to 1:0.45 in the Northeast Cape Fear ,Adult blueback herring and alewives River (Sholar 1977). In offshore attain a maximum size of about 290 mm North Carol ina waters, male blueback FL (females) and 270 mm FL (males) by herring were only slightly outnumbered age VII or VIII (Holland and Yelverton by females, 1:1.02 (Johnson et al. 1973). The oldest reported blueback 1978). herring and alewives (age IX) from the South Atlantic Region were collected After spawning, adults of both in Albemarle Sound (Holland et al. species return to the ocean, where 1975). they inhabit a narrow band of coastal water close to natal estuaries (Jones et a1 . 1978). Distribution of paren- THE FISHERY tal stocks during winter is not well defined, but they are presumed to Blueback herring and alewives are overwinter in offshore waters up to marketed together and labeled as 145 m deep (Bigelow and Schroeder "river herring" or "alewife" in many 1953; Hi1 debrand 1963). fisheries statistics. Both species are sold fresh or salted for human consumption, but most are used for GROWTH CHARACTERISTICS fish meal and fish oil in fertilizer, pet food, and domestic animal food. Growth Rates Some are used for fishing bait, and some are marketed for crab and No published data exist on growth crayfish bait. Roe from these species rates of juvenile alewives or blueback is canned and is highly valued as food herring in the South Atlantic Region, (Joseph and Davis 1965; Pate 1974; but some information is available Street and Davis 1976; Merriner based on average sizes of juveniles at 1978). different times of the year in various rivers. Juvenile blueback herring in U.S. commercial landings of river the Cape Fear River, North Carolina, herring (both species combined) a1 ong grew from 49.3 mm fork length (FL) in the Atlantic coast were 4,949 metric July to a mean of 57.4 mm FL in tons (t) in 1980 and 3,754 t in 1981. Table 1. Fork length (FL; in mm) - age (A; in years) and fork length-weight (W; in grams) relationships of alewife and blueback herring from the Chowan River and offshore North Carolina. Equations reported by Holland and Yelverton (1973).

Speciesa and Sex A1 ewife 3.34 C Offshore. NC W = 2.42 x lom6 FL 0.18 C Offshore, NC A = 190.50 FL Chowan River at Tunis, NC W = 7.49 x 10'~FL~'~~ -6 FL3.13 Chowan River at Tunis. NC W = 7.78 x 10 0.22 Chowan River at Tunis, NC A = 172.70 FL 0.18 Chowan River at Tunis, NC A = 181.40 FL 0.22 Chowan River at Tunis, NC A = 177.70 FL Blueback Herri ng C Offshore, NC W = 4.51 x 10'~FL~.~~ 0.37 C Offshore, NC A = 130.60 FL 3.08 M Chowan River at Tunis. NC W = 9.01 x 10'~FL 2.92 Chowan River at Tunis, NC W = 2.15 x 10'~FL 0.11 Chowan River at Tunis, NC A = 198.40 FL 0.10 Chowan River at Tunis. NC A = 197.90 FL 0.12 F Chowan River at Tunis, NC A = 200.90 FL

------a~ = sexes combined; M = males; F = females

These landings were worth $779.000 and nets, drift gill nets, haul seines, $671,000, respectively (NMFS 1982). and pound nets (Pate 1974). Pound The 1argest river herring fishery in nets recently produced about 95% of the South Atlantic Region is in North the yearly catch (McCoy 1976). In Carolina. From 1972 to 1981, North South Carol ina, the principal comner- Carol ina river herring landings cia1 gears used are haul seines and (31,357 t) accounted for over 97% of dip nets (Bulak et al. 1979); in the total for the South Atlantic Florida, haul seines and occasional 1y Region and were worth about $3 million pound nets are used (Williams et al. (Rulifson et al. 1982). North 1975). There is no commercial exploi- Carolina river herring landings in tation of blueback herring in Georgia, 1985 were 11,548 thousand pounds, the although some of the fish are caught highest since 1972 when the catch was incidentally by fishermen seeking 11,237 thousand pounds (Wins1 ow et a1 . American shad, A. sa idissima, and 1985) hickory shad, -A.-medhl ifson et a1. 1982). Fishing effort for both species is concentrated in rivers during spring A1 ewi fe and blueback herring spawning runs. In North Carolina, populations appear to be declining they are exploited by anchor gill in the South Atlantic Region. Comer- cia1 landings in North Carolina have can accommodate them (Norden 1968; decreased since 1969, and Florida Nigro and Ney 1982). Davis and Cheek landings are no longer reported. In (1966), who compared the food of juve- 1975, South Carolina imposed a quota nile alewife and blueback herring in on commercial landings in an effort to the Cape Fear River, North Carolina, reverse population declines in the reported that blueback herring Cooper River and Lake Moul trie (Curtis selected copepods and dipteran larvae 1976). Several factors seem to be more frequently than did alewives, causing this general decl ine . A1 e- whereas a1 ewives consumed more wives and blueback herring do not ostracods , insect eggs, and insect reach reproductive maturity until age parts. The amounts of crustacean eggs I11 or IV and, unlike American shad in the diets were similar for both that die after spawning once, these species. No benthic organisms or two species rely on repeat spawners to detritus were found in the stomach maintain their population levels. The contents of either species. inshore fishery is based on the exploitation of sexually mature The stomachs of all adult blueback adults; overfishi ng decreases the herring captured in offshore North abundance of older individuals, thus Carolina and examined for food con- decreasing annual spawning potential tained amphi pods, copepods, isopods , (Pate 1974). The offshore North cumaceans , mysids , and decapod larvae; Carol ina fishery was establ ished in none contained fish or fish remains 1967 as a trawl fishery that exploits (Holland and Ye1 verton 1973). A1 ewife sexual ly immature a1 ewives and blue- stomachs examined in the same study back herring (NMFS 1982). The com- contained unidentified fish remains in bined effect of the two fisheries has addition to zooplankton. After apparently played an important role in spawning in freshwater, adult alewives the decline of the North Carolina feed principally on the caddis fly populations (Rul ifson et a1 . 1982). Brachvcentrus sp (Cooper 1961). The status of blueback herring and Alewives feed three ways. Two are alewife fisheries in North Carolina size-selective (in favor of larger and recommendations for management prey): 1) particulate feeding on were summarized by Loesch et al. individual prey; and 2) gulping, in (1977), Johnson et al. (1978), Rul if- which the mouth opens and closes more son et al. (1982), and Winslow et a1. slowly than in particulate feeding. (1985) The third method is filtering with the mouth held agape. The feeding method used depends on an interaction of fish ECOLOGICAL ROLE size with prey size, type, and density (Janssen 1976). Food Competitors The alewife and blueback herring are primarily zoopl anktivores, but Few studies have been conducted on fish eggs, crustacean eggs, insects competitive interactions of alewife and insect eggs, and small fishes may and blueback herring. Some competi- be important food for the larger fish tion for food may occur between the (~igel ow and Schroeder 1953) . Larvae two species due to similarities in begin feeding on zooplankton immedi- diet and feeding behavior. Loesch et ately after the formation of a func- al. (1982a) described a spatial sepa- tional mouth (about 6 mm TL). They ration between young alewives and first rely on small cladocerans and blueback herring in the same habitat, copepods and begin to feed on larger which may lead to reduced competition zooplankton species as their mouths for food. Predators of control eggs (Schubel and Auld 1973). The hatching success of blue- Alewives and blueback herring are back herring and alewife eggs was not prey for many riverine, estuarine, and significantly affected by temperature marine piscivores (Cooper 1961), increases of 6-10 "C for 2.5-60.0 min including gulls and terns (Laridae) , (Schubel 1974). Larvae frorr eggs green herons (Butorides virescens) , stressed by prolonged exposure to otter (Lutra canadensis), and mink elevated temperatures, however, showed (Mustela vison) . Reported fish preda- a variety of deformities, including tors on juvenile alewives and blueback shortened bodies, en1 arged finfolds, herring include American eel, Angui lla and curved or twisted spines. The rostrata, and white perch, Morone magnitude and frequency of deformities americana (Kissil 1969). and chain were directly related to elevated pickerel, ESOX niger, largemouth bass, temperature levels and time of expo- Micropterus salmoides, yellow perch, sure (Koo and Johnson 1978). Incuba- Perca flavescens, and pumpkinseed, tion temperatures below 10.6 "C resul- Lepomis gibbosus (Cooper 1961). ted in 69% deformities in alewife Predators on adul ts are bluef ish . larvae (Edsall 1970). Pomatomus sal tatrix, weakfish; Cvnoscion~- reaal is and stri~ed bass There is no information available mxatilis (Cooper 1961; Tyus on the effects of temperature on juve- 1972). Blueback herrinq play. - an nile blueback herring. Upper 1ethal important ecological roie in the temperature 1imi ts and critical ther- Santee-Cooper System, South Carolina. mal maxima (the mean temperature at Since 1975, an average of 5.3 million which experimental fish lose equilib- herring pass upstream annually through rium) for juvenile alewife collected the Pinopol is Navigation Lock. These from Lake Michigan exceeded those of fish help to maintain an important adults by 3 to 6 "C and increased with striped bass sport fishery in Lakes higher accl imation temperatures. The Marion and Moultrie (Bulak and Curtis preferred temperature was consis- 1978). tently higher for juveniles than for adults (Otto et al. 1976). Some Effects on Freshwater Ecosvstems juvenile alewives survived and fed at temperatures of 34.4-35.0 "C (Dorfman Spawning alewives contribute a and Westman 1970). For a northern substanti a1 net increase in carbon, a1 ewi fe popul ati on, McCaul ey and nitrogen, and phosphorus to small Binkowski (1982) reported an upper streams. Most of the input comes from incipient 1ethal temperature of 31- mortality of the fish. Increased 34 'C for adul ts. Fish were accl imated nutrients from alewives lead to faster first at 27 'C. mi crobi a1 decomposition of 1eaf 1 itter, and probably benefit invertebrates In heat-shock tests with adult that feed on decaying litter. These alewives from Lake Michigan, critical invertebrates are important prey of thermal maxima and upper lethal tem- fishes (Durbin et a1 . 1979). perature 1imi ts increased as accl imation temperatures increased; at equal accl imation temperatures, the critical ENVIRONMENTAL REQUIREMENTS thermal maximum was not affected by fish age. In cold-shock tests with Temperature adult alewives from Lake Michigan, temperatures less than 3 "C caused Hatching success of blueback 100% mortality regardless of the herring eggs exposed to simulated acclimation temperature. Some fish power plant thermal regimes (7-20 "C survived a temperature decrease of 10 above ambient) was 10%-14% below that "C if the lower test temperature was not below 3 "C (Otto et al. 1976). No herring than in the alewife. Blueback information is available on tempera- herring prefer shal low, vegetated ture effects on adult blueback areas with slow current, whereas ale- herring. Alewife and blueback herring wives use a variety of spawning sites, on the open ocean were most frequently from brackish tidal water and barrier caught at 4-7 'C (Neves 1981). beach ponds to upstream mid-river sites. Changes in water currents or Salinity substrates in spawning rivers may affect blueback herring more than Although 1ittle information exists alewives because of the more specific on salinity tolerances of alewives and spawning site requirements of blueback blueback herring, they are apparently herring. efficient osmoregulators in freshwater or saltwater and are highly tolerant Schubel and Wang (1973) found that of sal inity changes. Chi ttenden high levels of suspended sediment (1972) observed no mortality of adult caused a delay in hatching of several blueback herring from either gradual hours. However, Auld and Schubel or abrupt sal inity changes, including (1978) found that 100 mg/L or less of transfers from freshwater to seawater suspended sediment had no effect on and the reverse. Blood and muscle the hatching success of alewife or electrolyte concentrations were blueback herring eggs. simi 1ar in alewives held in seawater and in freshwater at the same temper- Juvenile alewives and blueback ature (Stanley and Colby 1971). The herring in the Cape Fear River, North existence of 1and1 ocked , reproducing Carolina, were found in areas with 4 populations in 1akes and reservoirs to 22 ppm free carbon dioxide, 5 to 32 indicates that neither species ppm a1 kal inity, 2.4-10.0 mg/L requires a saltwater environment to dissolved oxygen, and a H of 5.2 to complete its life cycle. 6.8 (Davis and Cheek 1966r. Dissolved Oxygen In pooled samples taken throughout Mass mortalities of juvenile the year, alewives on the open sea were blueback herring occurred in the lower captured most often at 56-110 m depths, 48 km of the Connecticut River during and blueback herring at 27-55 m. June and July in 1965-67 and 1971, Evidence suggests that both species are when dissolved oxygen concentrations vertical migrators, following the die1 fell below 1.3 m /L at 24.6 "C and 3.6 movements of zooplankton in the water mg/L at 27.6 "C 4 Moss et al. 1976). column (Neves 1981). In 1aboratory studies, juvenile alewives responded to dissolved oxygen Environmental Contaminants concentrations below 2.0 mg/L by moving to the surface of the test The LC-50 (lethal concentration chamber. They can survive for at least for 50% of fish tested) of total 5 min at concentrations as low as 0.5 residual chlorine for blueback herring mg/L if allowed access to an area of eggs in the Potomac River, Maryland, 3.0 mg/L or higher concentration in ranged from 0.20 to 0.32 ppm, and all which to recover (Dorfman and Westman larvae exposed to sublethal concentra- 1970). tions of total residual chlorine were deformed (Morgan and Prince 1976). Substrate and System Features The body tissues of juvenile alewives and blueback herring from the Chicko- Requirements for spawning habitat hominy and James Rivers, Virginia, are more specialized in the blueback contained kepone concentrations greater than 0.3 ppm -- the action level found in fish from the Rappahannock for possible closure of a fishery River, Virginia, or the Potomac River, (Johnson et al. 1978; Loesch et al. Maryland (Loesch et al. 1982b). 1982b). Less than 0.3 ppm kepone was present in young a1 ewives and blueback Status of water qua1 ity and system herring from the Mattaponi and features of major river systems of the Parnunkey Rivers, Virginia, and no South Atlantic Bight are shown in detectable concentration of kepone was Table 2. Nassau I I I 1x1 1x1 I I 1x1 T mxrl I I I 1x1 St. Johns Tomoka 1) e.g.. Heavy metals, organics. 2) Locks and barriers. 3) Agricultural drainage; agricultural and domestic non-point source pollution: overfishing. 4) Agricultural and domestic non-point source pollution.

Table 2. System features and water qua1 ity characteristics that may affect populations of alewife or blueback herring in the South Atlantic Bight (from Rulifson et al. 1982b). 12 LITERATURE CITED

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Edsall , T. A. 1964. Feeding by three Holland, B. F., Jr., A. B. Powell, and species of fishes on the eggs of G. F. Yelverton. 1975. Anadromous spawning a1 ewives. Copeia fisheries research program, Northern 1964:226-227. Coastal Region, Offshore N.C. Ann. Prog. Rep. AFCS-8-2, N.C. Dep. Nat. Edsall, T. A. 1970. The effect of Resour. Commer. Dev., Div. Mar. temperature on the rate of Fish. 89 pp. development and survival of a1 ewife eggs and larvae. Trans. Am. Fish. SOC. 9 :376-380. Johnson, H. B., D. W. Crocker, B. F. Holland, Jr., J. W. Gilliken, D. L. Fisher, C. A. 1980. Anadromous Taylor, M. W. Street, J. G. Loesch, Fisheries Research Program. Cape W. H. Kriete, Jr., and J. G. Travel- Fear River System, Phase 11. N.C. stead. 1978. Biology and manage- Dep. Nat. Resour. Commer. Dev., Div. ment of mid-At1 antic anadromous Mar. Fish., Completion Report, Proj. fishes under extended jurisdiction. AFCS-15, 65 pp. NC-VA AFCS 9-2. N.C. Div. of Mar. Fish. and Va. Inst. Mar. Sci. Frankensteen, E. D. 1976. Genus 175 pp. Alosa in a channelized and unchannel ized creek of the Tar River Basin, North Carolina. M.S. Thesis. Jones, P. W., F. D. Martin, and J. D. Eas t Carol ina University, Hardy. 1978. Development of fishes Greenville, N.C. 123 pp. of the mid-Atlantic Bight: an atlas of the egg, larval and juvenile Godwin, W. F., and J. G. Adams. 1969. stages, Vol. 1. U.S. Fish Wildl. Young clupeids of the Altamaha Serv. Biol. Serv . Program River, Georgia. Ga. Game Fish. FWS/OBS-78/12. 366 pp. Comm., Mar. Fish. Div., Contrib. Ser. 15. 30 pp. Joseph, E. B., and J. Davis. 1965. A prel iminary assessment of the river Graham, J. J. 1956. Observations on herring stocks of lower Chesapeake the alewife in freshwater. Univ. Bay. Va. Inst. Mar. Sci. Spec. Sci. Toronto Biol . Ser. No. 62. 43 pp. Rep. No. 51. 23 pp. Hawkins, J. H. 1979. Anadromous Fisheries Research Program-Neuse Kellogg, R. L. 1982. Temperature River. Progress Rep. for Proj. requirements for the survival and early development of the anadromous Loesch, J. G., W. H. Kriete, Jr., and alewife. Prog. Fish-Cult. 44:72-73. E. J. Foell. 1982a. Effects of 1 ight intensity on the catchabil ity Kissil, G. W. 1969. Contributions to of juvenile Alosa species. Trans. the life history of the alewife, Am. Fish. ~ocm:41-44. Alosa pseudoharengus (Wilson), in Connecticut. Ph.D. Thesis. Universi- Loesch, J. G., R. J. Hugget, and E. J. ty of Connecticut, Storrs. 111 pp. Foell. 1982b. Kepone concentration in juvenile anadromous fishes. Kissil, G. W. 1974. Spawning of the Estuaries 5(3) :175-181. anadromous a1 ewi fe, A1 osa pseudoharengus (Wilson), in Bride Lake, MacLellan, P., G.E. Newsom, and P.A. Connecticut. Trans. Am. Fish. Soc. Dill. 1981. Discrimination by 103:312-317. external features between a1 ewife and blueback herring. Can. J. Fish. Koo, T. S. Y., and M. L. Johnson. Aquat. Sci. 38:544-546. 1978. Larva deformity in striped bass, Morone saxati lis (Wal baum) , Mansueti, R. J. 1956. Alewife and blueback herring, A1 osa herring eggs and larvae reared aestival is (Mitchill ) , due to heat successfully in the lab. Maryland shock treatment of developing eggs. Tidewater News 13(1) :2-3. Envi ron. Po1 1ut . 16(2) :137-149. Marshall, M. D. 1977. Anadromous Kuntz, A., and L. Radcliffe. 1917. fisheries research program - Neuse Notes on the embryology and larva1 River. Prog. Rep. for Proj. AFCS- development of twelve teleostean 13-1, N.C. Dep. Nat. Resour. Comer. fishes. U.S. Bur. Fish. Bull. No. Dev., Div. Mar. Fish. 70 pp. 35. 134 pp. McCauley, R.W., and F. P. Binkowski . Leim, A. H., and W. B. Scott. 1966. 1982. Thermal tolerance of the Fishes of the Atlantic coast of alewife, Alosa pseudoharenqus. Canada. Fish. Res. Board Can. Bull. Trans. Am. Fish. Soc. III:389-391. No. 155. 485 pp.

Loesch, J. 1969. A study of the McCoy, E. G. 1976. Assessment of blueback herring, Alosa aestival is North Carolina's river herring (Mitchill). in Connecticut waters. fishery. N.C. Dep. Nat. Econ. Res., P~.D. ~hesis. University of Con- Div. Mar. Fish., Mimeo. Rep. 13 pp. necticut, Storrs. 78 pp. McLane, W. M. 1955. Fishes of the St. John's River System. Ph.D. Loesch, J. G., W. H. Kriete, Jr., H. Thesis. University of Florida, B. Johnson, R. F. Holland, and M. W. Tallahassee. 361 pp. Street. 1977. Biology and management of mid-At1 antic anadromous Merriner, J. V. 1978. Anadromous fishes under extended jurisdiction. fisheries of the Potomac Estuary. Proj. NU. NC-VA AFCS 9-1, N.C. Div. Va. Inst. Mar. Sci. Contrib. No. Mar. Fish. and Va. Inst. Mar. Sci. 696. 4 pp. Proj. Rep. 1977. 183 pp. Morgan, R. P., 11, and R. D. Prince. Loesch, J., and W. A. Lund, Jr. 1977. 1976. Chlorine toxicity to A contribution to the life history estuarine fish eggs and larvae. of the blueback herring, A1 osa Chesapeake Biol. Lab. Univ. Ed. Ctr. aestivalis. Trans. Am. Fish. Soc. Environ. Estuarine Stud. Ref. No. 106:583-589. 76-116 CBL. 122 pp. Moss, S. A., W. C. Leggett, and W. A. Price. W. S. 1978. Otolith comoari- Boyd. 1976. Recurrent mass son of Alosa pseudoharengus and mortal ities of the blueback herring , Alosa aestivalis. Can. J. Zool. Alosa aestivalis, in the lower Con- 56:1216-1218. necticut River. -In D. Merrirnan and L. M. Thorpe, eds. The Connecticut Rulifson, R.A. 1985. Distribution and River ecological study, the impact abundance of fishes in tributaries of of a nuclear power plant. Am. Fish. South Creek Estuary, North Carolina, Soc. Monogr. No. 1:227-234. U.S.A. J. Elisha Mitchell Sci. Soc. 101:160-176. National Marine Fisheries Service (NMFS). 1982. Fisheries of the Rulifson, R. A., M. T. Huish, and R. United States, 1981. U. S. Dep. W. Thoesen. 1982a. Anadromous fish Comer. Curr. Fish. Stat. No. 8200. in the Southeastern United States 131 pp. and recommendations for development of a management plan. U.S. Fish Neves, R.J. 1981. Offshore Wildl. Ser., Fish. Res., Region 4, distribution of alewife, Alosa Atlanta, Ga. 525 pp. pseudoharenaus, and blueback herring, Alosa aestival is, along the Atlantic coast. U.S. Natl . Mar. Fish. Serv. Rulifson, R.A., M.T. Huish, and R.W. Fish. Bull. 79:473-486. Thoesen. 1982b. Status of anadromous fishes in southeastern Nigro, A. A., and J. J. Ney. 1982. U.S. estuaries. Pages 413-425 &I Reproduction and early 1 ife accom- V.S. Kennedy, ed. Estuarine Com- modations of 1 andlocked alewives to pari sons. Academic Press, New York. a southern range extension. Trans. Am. Fish. Soc. 111:559-569. Schubel, J. R. 1974. Effects of exposure to time-excess temperature Norden, C. R. 1968. Morphology and histories typically experienced at food habits of the larval alewife in power plants on the hatching success Lake Michigan. Proc. Conf. Great of fish eggs. Estuarine Coastal Lakes Res. 11:103-110. Mar. Sci. 2:105-116.

Schubel, J. R., and A. H. Auld. 1973. O'Neill, J. T. 1980. Aspects of the Hatching success of blueback herring 1 ife histories of anadromous alewife and striped bass eggs with various and the blueback herring, Margaree time vs temperature histories. In River and Lake Ainsle, Nova Scotia, J. W. Gibbons and R. R. Sharitc 1978-1979. M.S. Thesis. Acadia eds. Thermal ecology, AFC Symp. University, Wolfville, Nova Scotia, Ser. (Conf. 730505), 1973. Canada. 306 pp. Schubel, J. R., and J. C. S. Wang. Otto, R. G., M. A. Kitchel , and J. 0. 1973. The effects of suspended Rice. 1976. Lethal and preferred sediments on the hatching success of temperatures of the a1 ewife (Alosa ye1 1ow perch, white perch, striped seudoharen us) in Lake Michigan. bass, and alewife eggs. Ichthyol . kE7dkh.SOC . 105 :96-106. Assoc. Spec. Rep. No. 30, Ref. 73-3. 77 PP. Pate, P. P. 1974. Age and size com- position of commercial catches of blueback herring in A1 bemarl e Sound, Scott, W. B., and E. J. Crossman. North Carolina, and its tributaries. 1973. Freshwater fishes of Canada. Rep. N.C. Dep. Nat. Econ. Res., Div. Fish Res. Board Can. Bull. 184. Comer. Sport Fish. 10 pp. 966 pp. 16 Sholar, T. M. 1975. Anadromous of SA6. ICNAF Res. Doc. 76/VI/61. fisheries survey of the New and 7 PP. White Oak River Systems. N.C. Div. Mar. Fish., Compl. Rep. Oct. 73 - Jun. 75, Proj. AFC-9. 54 pp. Tyus, H. M. 1972. Notes on the 1 ife history of the alewife, Alosa Sholar, T. M. 1977. Anadromous seudoharen us, in North ~aro- fisheries research program, Cape itche ell Sci. Soc. Fear River system, Phase I. Prog. 88(4) :240-243. Rep. for Proj. AFSC-12. N.C. Dep. Nat. Resour. Comer. Dev., Div. Mar. Williams, R., and W. Grey. 1975. Fish. 63 pp. Stream survey section of anadromous fish project. 13 pp. In R. Smith, H. M. 1907. The fishes of Williams, W. Grey, and J. Hufceds. North Carolina. N.C. Geol. Econ. Study of andromous fishes in Surv. 2. 453 pp. Florida. MS. completion report NMFS Grant-in-aid Program, AFSC-5. Fla. Spitsbergen, D. L., and M. Wolff. Dep. Nat. Resour., Mar. Resour. 1974. Survey of nursery areas in Lab., Mimeo. western Pam1 ico Sound, North Carolina. Compl . Rep. Proj. No. Williams, R., W. Grey, and J. Huff. 2-175-R. N.C. Dep. Nat. Econ. Res., 1975. Study of anadromous fishes Div. Com. Sport Fish. 80 pp. of Florida. MS. Compl . Rep. NMFS Grant-in-aid Prog., AFCS-5. Fla. Stanley, J. G., and P. J. Colby. Dep. Nat. Resour., Mar. Resour. Lab. 1971. Effects of temperature on 160 pp. electrolyte balance and osmoregu- lation in the alewife in fresh and Winslow, S. E., S. C. Mozley, and R. sea water. Trans. Am. Fish. Soc. A. Rulifson. 1985. N.C. Anadromous 100 :624-638. fisheries management program. Com- pletion Report AFCS-22. N.C. Dep. Street, W. M. 1970. Some aspects of Nat. Resour., Community Develop., the life histories of hickory shad, Div. Mar. Fish. Alosa mediocris [Mitchilll . and blueback herring, ~losaaeifival is (Mitchill) , in the A1 tamaha River, Winters, G. H., J. A. Moores, and R. Georgia. M.S. Thesis. University Chaul k. 1973. Northern range of Georgia, Athens. 89 pp. extension and probable spawning of gaspereau (A1 osa pseudoharengus) in Street, M. W., and J. Davis. 1976. the Newfoundtd area. J. Fish. Notes on the river herring fishery Res. Board Can. 30:860-861. 50272 -101 REPORT DOCUMENTATION 1. REPORT NO. 2. 3. Rec8p8ent'a Acceauon No. PAGE Biological Report 82(11.111) *

Species Profiles: Life Histories and Environmental Requirements I kugust 1989 of Coastal Fishes and Invertebrates (South Atlantic)--Alewife Is. imd Rll~~haCkH~rnng. I 7. Author(a1 I. Pertorrnlng Organizat!on Rept. No. Earl L. Bozeman, Jr., and Michael J. Van Den Avvle 9. Pedorming Organization name and Address 10. Pro~ect/Tash/Worh Unlt No. Georgia Cooperative Fish and Wild1 ife Research Unit School of Forest Resources 11. Contrac~(Clor Granl(G1 NO. University of Georgia (c) Athens, GA 30602 (GI

12. Soonwrtng Organiratlon Name and ~ddress 1 13. 710. oi R~WR6. Permd Covered U.S. Department of the Interior U.S. Amy Corps of Engineers Fish and Wildlife Service Waterways Experiment Station National Wetlands Research Center P.O. Box 631 14. Washinaton. DC 20240 VickJura. MS 39180 IS. Supplementary Notes *U.S. Army Corps of Engineers Report No. TR ED-82-4

16. Abstract (Limit: 200 rorda) Species profiles are summaries of the literature on taxonomy, life history, and environmental requirements of coastal fishes and aquatic invertebrates. They are prepared to assist with impact assessment. The alewife (Alosa pseudoharengus) and blueback herring (A. aestival is) are morphologically and ecologically similar anadromous species of cluneids. The blueback herring is common throughout the South Atlantic Region, but the alewife occurs primarily in North Carolina and northern parts of South Carolina. These species spawn in spring in freshwater or brackish, tidally influenced portions of coastal rivers. Blueback herring initially use freshwater habitats for nursery areas, and then migrate downriver to brackish estuaries, where they overwinter prior to migrating to sea the following spring. Alewives use brackish water or tidal freshwater as nursery areas until they migrate to coastal waters in winter or the following spring. Landlocked populations of bl ueback herring occur in several southeastern reservoirs. Both species are ecologically important by serving as prey for many other fishes; they are economically important because they support commercial inshore and offshore fisheries . Little information is available on environmental factors that limit these species in the South Atlantic Region. Adults of both species have broad salinity tolerances, but bl ueback herring appear to require access to freshwater for successful reproduction.

17. Document Anelpis a. Deurrmon Estuaries Growth Coastal rivers Feeding Coastal areas Spawning Cl upei dae b. Idmtlt3enlOp.n.End.d Terms A1 ewi fe Habitat requirements A1 osa pseudoharengus Bl ueback herring Alosa aestival is c. COUTl FieldlGmup

18. Aw~llabilltySlatem.... 1 19. S*curnty Class (Thm Rewnl I 21. No. oi Pal-8 / Unclassified 1 17 -- Unl imi ted 8 m s.c rut~Clja nhi Pa~e) I 22. Pnce i ~ncYass~fied I

5.. In.rruct,ons on R*r.rr. OPTIONAL FORM 272 (1-71 (Formerly NTIS-351 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 resources, protecting our fish and wildlife, preserving the environmental and cultural values of our national parks and historical places, and providing for the enjoy- ment of life through outdoor recreation. The Department assesses our energy and mineral resources and works to assure that their development is in the best interests of all our people. The Depart- ment also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S. administration.

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