QL 

155 

.S63 

no. 

82-11.9 

FWSlOBS82111.9 

October 1983 

TR EL-82-4 

Species Profiles: Life Histories and 

Environmental Requirements of Coastal Fishes 

and Invertebrates (Mid-Atlantic) 

ALEWIFE/BLUEBACK HERRING 

Coastal Ecology Group 

Fish and Wildlife Service Waterways Experiment Station 

U.S. Department of the Interior U.S. Army Corps of Engineers 

FWS/OBS-82/11.9 

TR EL-82-4 

October 1983 

Species Profiles: Life Histories and Environmental Requirements 

of Coastal Fishes and Invertebrates (Mid-Atlantic) 

ALEWIFE/BLUEBACK HERRING 

Clemon W. Fay, Richard J. Neves, and Garland B. Pardue 

Department of Fisheries and Wildlife Sciences 

Virginia Polytechnic Institute and State University 

Blacksburg, VA 24061 

Project Manager 

Larry Shanks 

Project Officer 

Norman Benson 

National Coastal Ecosystems Team 

U.S. Fish and Wildlife Service 

1010 Gause Boulevard 

Slidell, LA 70458 

This study was conducted 

in cooperation with 

Coastal Ecology Group 

U.S. Army Corps of Engineers 

Waterways Experiment Station 

Performed for 

National Coastal Ecosystems Team 

Division of Biological Services 

Fish and Wildlife Service 

U.S. Department of the Interior 

Washington, DC 20240 
CONVERSION FACTORS 

Multiply 

Metric -- to U.S. Customary 

BY To Obtain 

millimeters (mm) 0.03937 inches 

centimeters (cm) 0.3937 inches 

meters (m) 3.281 feet 

kilometers (km) 0.6214 miles 

square meters (m') 10.76 

square kilometers (km2) 

square feet 

0.3861 square miles 

hectares (ha) 2.471 acres 

liters (1) 0.2642 

cubic meters (m3) 

gallons 

35.31 cubic feet 

cubic meters 0.0008110 acre-feet 

milligrams (mg) 0.00003527 ounces 

grams (gm) 0.03527 ounces 

kilograms (kg) 2.205 pounds 

metric tons (mt) 2205.0 

metric tons (mt) 

pounds 

1.102 short tons 

kilocalories (kcal) 3.968 BTU 

Celsius degrees 1.8((Y) + 32 Fahrenheit degrees 4 

inches 

inches 

feet (ft) 

fathoms 

miles (mi) 

nautical miles (nmi) 

U.S. Customary to Metric - 

25.40 

2.54 

0.3048 

1.829 

1.609 

1.852 

millimeters 

centimeters 

meters 

meters 

kilometers 

kilometers 

square feet (ft2) 0.0929 square meters 

acres 

square miles (mi2) 

0.4047 hectares 

2.590 square kilometers 

gallons (gal) 

cubic feet (ft3) 

3.785 

0.02831 

acre-feet 1233.0 

liters 

cubic meters 

cubic meters 

ounces (oz) 28.35 grams 

pounds (lb) 0.4536 kilograms 

short tons (ton) 0.9072 metric tons 

BTU 0.2520 kilocalories 

Fahrenheit degrees 0.5556(F" - 32) Celsius degrees 
CONTENTS 

CONVERSION TABLE 

PREFACE 

ACKNOWLEDGMEN;S' 

PROFILE SCOPE . 

............................. 

............................. 

............................. 

............................. 

NOMENCLATURE/TAXONOMY/RANGE ....................... 

MORPHOLOGY/IDENTIFICATION AIDS ...................... 

Alewife 

Blueback ;eFring ......................................................... 

Aids for Species Separation ...................... 

REASON FOR INCLUSION IN SERIES ...................... 

LIFEHISTORY ............................... 

Reproductive Physiology/Strategy .................... 

Spawning ................................ 

Eggs .................................. 

Yolk-Sac Larvae ............................ 

Larvae ................................. 

Juveniles ............................... 

Adults ................................. 

GROWTH CHARACTERISTICS .......................... 

GrowthRates .............................. 

Length-Weight Relationships ...................... 

THEFISHERY ............................... 

Commercial Fisheries .......................... 

Recreational Fisheries ......................... 

Population Dynamics .......................... 

ECOLOGICAL ROLE ............................. 

FoodHabits .............................. 

Feeding Behavior ............................ 

Competitors .............................. 

Predators 

ENVIRONMEtiTAL'R;QUIREI;1E~TS : : : : : : : : : : : 

.......................... 

Temperature .............................. 

Salinity ................................ 

Other Environmental Factors ...................... 

Environmental Contaminants ....................... 

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

Page 

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iii 
PREFACE 4 

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 

profile has sections on taxonomy, life 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 1l.S. Army Corps of Engineers 

and the U.S. Fish and Wildlife Service. 

Habitat Suitability Index (HSI) models are being prepared by the U.S. Fish 

and Wildlife Service for the alewife and blueback herring. HSI models are 

designed to provide a numerical index of the relative value of a given site as 

fish or wildlife habitat. 

Suggestions or questions regarding this report should be directed to: 

Information Transfer Specialist 

National Coastal Ecosystems Team 

U.S. Fish and Wildlife Service 

NASA-Slide11 Computer Complex 

1010 Gause Boulevard 

Slidell, LA 70458 

or 

U.S. Army Engineer Waterways Experiment Station 

Attention: WESER 

Post Office Box 631 

Vicksburg, MS 39180 

This series should be referenced as follows: 

U.S. Fish and Wildlife Service. 1983. Species profiles: life histories and 

environmental requirements of coastal fishes and invertebrates. U.S. Fish 

anti Wildlife Service, Division of Biological Services, FWS/OBS-82/11. 

U.S. Army Corps of Engineers, TR EL-82-4. 

This profile should be cited as follows: 

Fay, C.W., R.h. Neves, and G.B. Pardue. 1983. Species profiles: life histories 

and environmental requirements of coastal fishes and invertebrates (MidAtlantic) -- alewife/blueback herring. 1J.S. Fish and Wildlife Service, 

Division of Biological Services, FWS/OBS-82/11.9. U.S. Army Corps of 

Engineers, TR EL-82-4. 25 pp. 

iv 
ACKNOWLEDGMENTS 

We are grateful for the review by Dr. Joseph Loesch, Virginia Institute of Marine 

Science, Gloucester Point. We appreciate permission from the Canadian Journal of 

F-isheries and Aquatic Sciences for reprinting Figures 1 and 3. Permission is acknowiedged from the Southeastern Association of Fish and Wildlife Agencies for reprinting 

Figure 4. 

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

ALEWIFE/BLUEBACK HERRING 

PROFILE SCOPE 

0 

This profile covers life history 

and environmental requirements of 

. both alewife (Alosa pseudoharengus) 

and blueback herring (Alosa aestivalis), since their distribution is 

overlapping and their morphology, 

ecological role, and environmental 

requirements are similar. Nevertheless, significant differences in certain 

physical, physiological, and biological 

w characteristics exist between the two 

species. When available, these differences are addressed by separate 

statements for each species. In addition, a special section on the most 

readily distinguishing characteristics 

for separating eggs, larvae, and 

adults of the two clupeids is presented in the morphology section of 

this profile. 

Because most of the information 

available concerns the alewife, characteristics of this species are given priority reference in the text. Features 

1 
should not be considered similar 

in the two species unless noted as 

such. In a few studies (particularly 

commercial fisheries statistics), the 

two species are referred to collectively 

as “river herring” or “gaspereau” 

(Canada). 

NOMENCLATURE/TAXONOMY/RANGE 

Scientific names. .Alosa pseudoharengus 

(Wilson)/Alosa aestivalis (Mitchill) _.- 

Preferred common names. . Alewife/blueback herring (Figure 1) 

Other common Ildm?S.. . . River herring, 

gaspereau, oldwife. 

Cla_ss.. . . . . . . . . . . . . . . . . . . .Osteichthyes 

Order . . . . . . . . . . . . . . . . . . .Clupeiformes 

Family. . . . . . . . . . . . . . . . . . . . . . .Clupeidae 

Geographical range: The alewife is 

an anadromous species found in 

riverine, estuarine, and Atlantic 

coastal habitats, depending on 

life cycle stage, from Newfoundland (Winters et al. 1973) to 

South Carolina (Berry 1964). 

Landlocked populations are in the 

Great Lakes, Finger Lakes, and 

many other freshwater lakes 

(Bigelow and Schroeder 1953; 

Scott and Crossman 1973). The 

blueback herring is an anadromous species found in riverine, 

estuarine, and Atlantic coastal 

habitats, depending on life stage 

cycle, from Nova Scotia to the 

St. Johns River, Florida (Hildebrand 1963) (see Figure 2 for a 

map of the mid-Atlantic distribution of alewives and blueback 

herring). 

MORPHOLOGY/IDENTIFICATION AIDS 

The following information was 

taken from summaries in Jones et al. 

(1978), unless otherwise indicated. 

Alewife 

Dorsal rays 12-19 (usually 

13-14), anal rays 15-21 (usually 

17-18), scales in lateral series 42-54. 

Prepelvic scutes 17-21 (usually 

19-20), postpelvic scutes 12-17 (usually 14-15), gill rakers on first arch 

38-46. Body strongly compressed, 

deep. Mouth oblique, anterior end of 

lower jaw thick, heavy, and extending 

to middle of orbit. Eye large, diameter 

greater than snout length. Color: 

dorsally grey to grey-green, laterally 

silver with prominent dark shoulder 

spot; fins pale, yellow or green. 

Blueback Herring 

Dorsal rays 15-20, anal rays 

15-21, scales in lateral series 46-54. 

Prepelvic scutes 18-21, postpelvic 

scutes 12-16, gill rakers on first arch 

41-52. Body moderately compressed, 

elongate, eye diameter small, equal to 

or less than snout length. Upper jaw 

with definitive median notch, no teeth 

on premaxillaries. Color: dorsally 

blue to blue-green, laterally silver 

with prominent dark shoulder spot; 4 

fins pale, yellow or green. 

Aids for Species Separation 

Eggs. Unfertilized blueback herring eggs amber, alewife eggs green. 

Oil droplets of fertilized eggs unequal 

and scattered for blueback herring, 

numerous and uniformly tiny for alewife (Kuntz and Radcliffe 1917; Norden 1967). 

Larvae. Myomeres between i nsertion of dorsal fin and anal vent 11-13 

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

(this characteristic is definitive 

according to Chambers et al. 1976) 

Larvae less than approximately 15 mmi 

can be separated using regressions of 

vent to tail distance (mm) and vent to 

u rostyle distance (mm), against 

l25.4 mm = 1 inch. 

2 
N. Y. 

BALTIMORE JP 

WASHIfiTO%.-I 

‘-’ 

James R. 

ATLANTIC OCEAN 

MILES 

0 50 100 

0 50 100 

KILOMETERS 

ijljjlij Coastal distribution 

Spawning, river/bay 

l Areaof high abundance 

Figure 2. Mid-Atlantic distribution of the alewife and blueback herring. 

3 
standard length (SL); presented in 

Chambers et al. (1976). 

Adults. Externally, by scale 

imbrication patterns and individual 

scale markings (Figure 3). Scale 

baseline and dividing line coincidental 

on alewife scales, not on blueback 

herring scales (O’Neill 1980; MacLellan 

et al. 1981). Internally, peritoneal 

lining uniformly dark in blueback herring; pale, grey, or silvery with dark 

punctations in alewife (Leim and Scott 

1966; Scott and Crossman 1973). 

Shape of otolith distinctive (Scott and 

Crossman 1973) and differences 

described in Price (1978) and illustrated in O’Neill (1980). Biochemically 

distinguished by muscle myogen and 

retina LDH enzyme migration patterns 

(electrophoresis) (McKenzie 1973, 

1975). 

Alewives possess fewer vertebrae, dorsal rays, anal rays, and gill 

rakers on the first arch, in general, 

compared with blueback herring. Eye 

diameter to snout length ratios are 

also significantly different between the 

two clupeids, but mean values are 

close and ranges overlap (Messieh 

1977). The applicability of these 

meristic characters for absolute species determination is limited since 

there is at least some overlap between 

species for each characteristic. 

Though dorsal coloration has 

been cited as species-distinctive in 

fresh specimens (Bigelow and Schroeder 1953), this character may not be 

reliable. MacLellan et al. (1981) 

found no significant or detectable difference in dorsal coloration and 

observed that such coloration 

appeared to vary substantially with 

ambient lighting conditions. 

ALEWIFE BLUEBACK HERRING 

approx. scale: t lcm 

c--------_-e ANTWon MARGIN --+-------------4 

Figure 3. Scale imbrication patterns 

(top) and individual scale morphology 

(bottom) used for external discrimination of the alewife and blueback herring (from MacLellan et al. 1981; with 

permission of the Canadian Journal of 

Fisheries and Aquatic Scienceimportant links in estuarine and 

marine food webs, between zooplan kton and top piscivores. Commercially, 

both species have recently (last two 

decades) gained in recognition and 

interest as sources of fish meal, fish 

oil, and fish protein, particularly for 

the animal food industries. 

LIFE HISTORY 

REASON FOR INCLUSION IN SERIES 

The alewife and blueback herring 

Reproductive Physiology/Strategy 

are important ecologically and to a Alewives 

lesser extent as commercial fish speand blueback herring 

are heterosexual, 

ties . Ecologically, 

though hermaphrothese species are ditism in landlocked populations of & 

4 
alewives from Lake Michigan has been 

reported (Edsall and Saxon 1968; Hlavek and Norden 1977). Females of 

both species are slightly larger and 

heavier than males of the same age 

- (Cooper 1961; Netzel and Stanek 1966; 

Marcy 1969; Loesch and Lund 1977). 

Some male anadromous alewives, 

and a slightly smaller percentage of 

females, spawn for the first time at 

age 3. Most fish of both sexes have 

spawned once by age 4, and all by 

age 5. Generally, males dominate age 

classes 3 to 5 on the spawning 

grounds, while females dominate age 

classes 7 and older. Blueback herring vary more than alewives in age 

of first spawning, though in general 

maturation rates are similar for the 

two clupeids (Joseph and Davis 1965; 

Loesch and Lund 1977; O’Neill 1980). 

Age of first spawning, percentage of repeat spawners, and longevity 

in populations seem to decrease as one 

p roceed s from north to south. 

Spawning populations of alewives in 

southern North Carolina comprised 

primarily 3-year-old fish, and no fish 

_ over 4 years old were found (Tyus 

1974). In contrast, alewife spawning 

populations in Chesapeake Bay were 

represented by ages 3 to 8 (Joseph 

and Davis 1965), Connecticut River 

stocks by ages 3 to 8 (Marcy 1969; 

Loesch and Lund 1977), and Nova 

Scotian stocks (both alewives and 

blueback herring) by ages 4-10 

(O’Neill 1980). Percentage of repeat 

spawners was 60% for alewives in Nova 

Scotia (O’Neill 1980), 61% in the York 

River, Virginia (Joseph and Davis 

1965), and less than 10% in southern 

North Carolina (Tyus 1974). Blueback herring runs consisted of 65% 

and 75% repeat spawners in the York 

River and Nova Scotian waters, 

respectively (Joseph and Davis 1965; 

O’Neill 1980). 

Fecundity of Connecticut River 

blueback herring ranged from 45,800 

eggs (238-mm female) to 349,700 eggs 

(310-mm female) (Loesch and Lund 

1977). F,rom 10% to 30% of 

the initial number of eggs present in 

a female blueback herring remained 

after spawning. Left ovaries were 

significantly heavier than right ovaries, though left ovaries did not contain more eggs per gram of ovary 

than did right ovaries (Loesch and 

Lund 1977; Loesch 1981). Chesapeake 

Bay alewives ranged in fecundity from 

60,000 to 100,000 eggs per female 

(Foerster and Goodbred 1978). The 

fecundity-to-age relationship for 

Georgia populations of blueback herring did not fit a linear relationship 

well (r-2 = 0.42) (Street 1969). It was 

suggested that the age-fecundity relationship is asymptotic for both alewives and blueback herring, and thal 

“fecundal senility” may occur in all 

long-lived stocks of these species 

(Street 1969; Loesch and Lund 1977). 

Spawning 

Anadromous alewives and blueback herring spawn once a year, during spring or early summer, in fresh 

or brackish water (Raney and Massmann 1953). Males arrive at mouths 

of spawning rivers earlier than 

females (Cooper 1961; Tyus 1971; 

Rich kus 1974a). Spawning environments vary from streams only a few 

meters (yards) wide and a few centimeters (inches) deep to large rivers 

such as the Delaware, Susquehanna, 

and Potomac (Mansueti 1956). Ponds, 

including barrier beach ponds, with 

an open outlet to the sea, are also 

used by alewives (Bigelow and Welsh 

1925). Jones et al. (1978) cited 

Loesch (1968, 1969) as stating that 

blueback herring do not ascend into 

freshwater as far as alewives on 

spawning runs. However, Loesch 

(personal communication) said the 

statement pertained to the existing 

literature at that time, and findings 

reported in Loesch and Lund (1977) 

indicated that upstream distribution 

was a function of finding appropriate 

spawning habitats. 
In laboratory tests, adult anad romou s alewives from Rhode island 

waters were capable of distinguishing 

water of their natal pond from water 

collected in nearby ponds (Thunberg 

1971). Olfaction was shown to be the 

major sensory mechanism for homing 

behavior. Discriminate function analysis, however, by Messieh (19’77) on 

St. Johns River, Florida, spawning 

stocks indicated considerable straying 

from home streams, particularly 

between adjacent spawning areas/ 

stocks. Messieh (1977) hypothesized 

that the majority of stock mixing 

occu r red during the prespawning 

period (late winter, early spring) 

rather than “impulsively” on the 

actual spawning runs. The majority 

of spawning alewives in Lake Mattamuskeet, North Carolina, used only 

one of four available canals for 

spawning migrations (Tyus 1974). It 

was hypothesized that since this canal 

was the only one available historically 

(built in 1907), the alewives may be 

exhibiting homing behavior. Tyus 

(1974) did not report, however, that 

this canal was also the shortest and 

most direct from bay to lake. 

Spawning periods along the 

Atlantic coast ‘range from late March 

through July, occurring later in the 

north. Within the mid-Atlantic region, 

nearly all alewife and blueback herring spawn from April through midJuly (Hildebrand 1963; Kissil 1969; 

Loesch 1969; Smith 1971; Tyus 1974; 

Loesch and Lund 1977). In general, 

alewives begin to spawn 3 to 4 weeks 

before blueback herring in sympatric 

areas. Spawning peaks are 2 to 3 

weeks apart (Jones et al. 1978). Alewives began spawning at minimum 

water temperatures of 10.5”C (51” F) 

(Cianci 1969) and blueback herring at 

14OC (57OF) (Loesch and Lund 1977). 

Both species cease spawning when 

water temperature exceeds 27 O C (81” 

F) (Loesch 1969; Edsall 1970). 

Blueback herring prefer spawning 

sites with fast currents and associated 

:,a;:) substrates (Loesch and Lund 

Brackish water or standing 

water habitats are rarely used. In 

contrast, alewives select a wide variety of spawning sites, using standing 

water and oxbows as well as mid-river 

sites (Kissil 1974). Several studies 

have described alewife spawning in 

ponds with an open connection to 

the ocean (e.g., Havey 1973; Kissil 

1974), but no observations of pond 

spawning by blueback herring are 

documented. Apparently a considerable separation, both spatially and 

temporally, exists for spawning activity of anadromous alewives and blueback herring. 

Loesch and Lund (1977) 

described spawning behavior of blueback herring. A spawning group 

composed of one female and several 

males swam in circles for several minutes, and males occasionally nudged 

the vent of the female. Swimming 

speed increased gradually until a deep 

dive occurred with subsequent release 

of eggs and milt simultaneously, very 

near the substrate. Spawning activities of both species occur diurnally 

and nocturnally, though the greatest 

activity apparently is nocturnal (Graham 1956; Edsall 1964). Both males 

and females migrate rapidly downstream after spawning, and tota I 

spawning time for a single migratory 

group is uslually 5 days or less 

(Cooper 1961; Loesch and Lund 1977). 

Until water-hardened, eggs of 

both species are demersal in still 

water and adhesive or pelagic in 

running water (Loesch and Lund 1977; 

Jones et al. 1978). After waterhardening (less than 24 hr), eggs lose 

their adhesive property and enter the 

water column. Fertilized, water- hardened e99s are green (alewife) to 

amber (blueback herring) and contain 

6 
scattered, unequal (alewife) or 

numerous, small (blueback herring) oil 

droplets (Kuntz and Radcliffe 1917; 

Norden 1967). Egg diameter ranges 

from 0.80 to 1.27 mm for alewives and 

from 0.87 to 1.11 mm for blueback 

herring (Mansueti 1962; Norden 1967). 

Incubation time is approximately 80 to 

94 hr at 20’ to 21°C (68’ to 70” F) 

and 55 to 58 hr at 22” to 24’ C(72O 

to 75OF) for blueback herring eggs 

(Cianci 1969; Morgan and Prince 

1976). Comparative values for alewives are approximately 360 hr. at 

7.2OC (45OF) (Edsall 1970), -178 hr 

at 12.7OC (55OF) (Kellogg 1982), 89 

hr at 21.1”C (7OOF) (Edsall 1970), 

72 hr at 23.8”C (75’ F) (Kellogg 

1982), and 50 hr at 28.9OC (84’ F) 

(Edsall 1970). An equation for predicting incubation time for alewife 

eggs from temperature (Edsall 1970) 

is: 

T = 6.335 x 106’x t 

where T = time in days and t = incubation temperature in degrees F. 

Yolk-Sac Larvae 

Yolk-sac larvae range from 2.5 to 

5.0 mm total length (TL) at hatching, 

and average 5.1 mm TL at yolk-sac 

absorption (Mansueti 1962; Norden 

1967). Duration of this stage is 2 to 5 

days for alewife and 2 to 3 days for 

blueback herring (Mansueti 1962; 

Cianci 1969). 

Larvae 

The larval stage lasts from 

yolk-sac absorption until transformation to the juvenile stage. La rva I 

blueback herring range from 4.0 to 

15.9 mm SL and larval alewives from 

4.3 to 19.9 mm SL (Jones et al. 

1978). Jones et al. (1978) provided 

detailed drawings of the developmental 

stages of eggs, yolk-sac larvae, and 

larvae of both alewives and blueback 

herrina. 

Alosa SPP. larvae, primarily 

those of blueback herring and alewives, were present throughout upper 

Chesapeake Bay from hatching 

(approximately April 15) through June 

(Dove1 1971); Larvae exhibited slight 

downstream movement from presumed 

spawning areas in the bay, and were 

collected only in areas with salinities 

less than 12 ppt. Alosa spp. larvae 

in Nova Scotian rivers occurred in 

areas that were relatively shallow (~2 

m, ~6.6 ft), sandy, and warm, and 

were collected in or near areas of 

spawning adults (O’Neill 1980). 

Juveniles 

Transformation to the juvenile 

stage is gradual, but is completed at 

approximately 20 mm TL. Scales first 

appear on juveniles between 25 and 29 

mm TL, and are fully developed at 45 

mm TL (Hildebrand 1963; Norden 

1967). 

Juvenile blueback herring in the 

James River, Virginia, exhibited a net 

upstream movement between June and 

October, presumably caused by contributions of juveniles from oxbows, 

side channels, and tributaries, which 

gradually moved down into the main 

river through the summer. Densities 

of juveniles were significantly higher 

near the surface than at 5-m (16.4-ft) 

depth throughout their residence in 

the river (Burbidge 1974). 

Warinner et al. (1969) studied 

the distribution of juvenile alewives 

and blueback herring in the Potomac 

River over the first 6 months of life 

(Figure 4). Four important conclusions were evident. 

1) Both species exhibited apparent upstream movement, averaging 24 km (15 mi) over 4 months, 

until the inception of emigration 

in October. 
75 JUNE 

I 

53 

I 25 n 

CC:OaER 

_ 

40 SO F& 70 8; i0 COST4T:CN -RIVER MILE 

Figure 4. Seasonal distribution of 

,juvenile river herring by river mile 

in the Potomac River, Maryland. No 

data are shown for November because 

fish were absent in the study area 

(from Warinner et al. 1969). 

2) Both species were virtually 

absent in the study area by 

November. 

3) Juvenile blueback herring outnumbered alewives 19: 1 overall. 

4) Juvenile alewives were most 

abundant in surface waters 

through September, and 

increased in abundance at 4.6 m 

(15 ft) and on the bottom in 

September and October, prior to 

emigration. In contrast, juvenile 

blueback herring maintained high 

abundance in surface waters 

through October, increased in 

abundance at 4.6 m in November, 

and were never collected in bottom trawls throughout the study 

period. 

Significant diel movements of 

juvenile alewives and blueback herring 

in Virginia rivers were found (Loesch 

et al. 1982a). Fish moved toward the 

bottom during the day and toward the 

surface at night. Blueback herring 

were more sensitive to the sky-opacity 

index and exhibited more extensive 

vertical movement patterns in relation 

to changing light intensity than aiewives. This movement pattern was 

also noted for juvenile alosids in the 

Potomac River (Warinner et al. 1969). 

In most Atlantic coast populations, juvenile alewives and blueback 

herring emigrate from freshwater/estuarine nursery areas between June 

and November of their first year of 

life (Burbidge 1974; Kissil 1974; Richkus 1975; O’Neill 1980). Juvenile 

river herring in upper Chesapeake 

Bay did not emigrate until early 

spring of their second year, and such 

migrations were comparatively rapid 

once they began (Dove1 1971). This 

was the only report of nonmigratory 

first-year juveniles, except for the 

dwarf, nonmigratory population of 

alewives described by Foerster and 

Goodbred (1978) from the Susquehanna River. 

Stimulatory variables influencing 

the initiation of migratory “waves” 

(Richkus 1975) of juvenile alewives 

from nursery habitats include heavy 

rainfall (Cooper 1961), high water 

(Kissil 1974; Rich kus 1975), and 

sharp declines in water temperature 

(Richkus 1975). Rich kus (1975) 

observed that (I) such waves lasted 2 

to 3 days, regardless of length of 

time of environmental 

migrations peaked in latZhaa?ECnocY~) 

and (3) the magnitude of a migratory 

wave was not related to the magnitude 

of environmental change. Most (608 to 

80%) juveniles emigrated on only a 

small percentage (7% to 8%) of the 

available days (Richkus 1975). 

8 
Significant numbers of juvenile 

alewives and blueback herring were 

captured during winter in the Mullica 

River Estuary, New Jersey, indicating 

the importance of this area for overwintering (Milstein 1981). Blueback 

herring was third in percent representation and alewives seventh among 

53 species collected. Both species 

were captured out to 8 km (5 mi) offshore, which is near the outer limit of 

salinity/temperature influences from 

the Mullica River Estuary. Overwintering fish chose temperatures 

between 4.5”and 6.5*C (40’ zyd 43.7” 

F) and salinities from 29 to 32 ppt. 

Densities of overwintering juveniles 

increased steadily from December to 

March, then declined in April. The 

alewife-to-blueback herring ratio in all 

samples combined was 1:5 (Milstein 

1981). 

Adults 

Little information is available 

concerning the life history or biology 

of alewife and blueback herring stocks 

once the juveniles emigrate to the sea 

at age 0’ or 1’. A literature search 

indicated that much research needs to 

be conducted in stock identification, 

offshore exploitation rates, foreign 

fishing operations, and general life 

history, movement , migration patterns, feeding behavior, and ecology 

of these clupeids in offshore waters. 

Neves (1981) summarized 16 years 

of catch data from National Marine 

Fisheries Service trawl surveys conducted along the Atlantic coast, 

between Cape Hatteras and Nova Scotia. Samples were taken out to depths 

of 200 m (656 ft) . The majority of 

the catch of alewives and blueback 

herring was taken at sampling stations 

where water depth was less than 100 

m (328 ft). Alewives and blueback 

herring collected ranged from 60 to 

350 mm fork length (FL), and alewives 

outnumbered blueback herring 

approximately 1O:l for all samples 

combined. Statistical analysis determined that alewives were most 

abundant at depths between 56 and 

110 m (184 and 361 ft), while blueback herring were most abundant 

between 27 and 55 m (89 and 180 ft). 

In summer and fall, catches of both 

species were confined to the sampling 

area north of 40” north latitude, in 

three general areas: Nantucket 

Shoals, Georges Bank, and the perimeter of the Gulf of Maine. Winter 

catches were between 40” and 43” 

north latitude. Spring catches were 

distributed over the entire Continental 

Shelf in the study region (Neves 

1981). 

Adult alewives and blueback herring chose only a small portion of 

Georges Bank, specifically the western 

slope at 41°, 29’ north latitude and 68’, 

34’ west longitude, as an area of 

residence in July and October, 1964 

(only months sampled) (Netzel and 

Stanek 1966). Alewives outnumbered 

blueback herring in these samples. 

All mature age classes were represented, but age 0+, l+, and 2+ fish 

were not captured. 

Alewives and blueback herring, 

like other clupeids, may exhibit seasonal movements in conjunction with 

preferred isotherms (or preferred 

isotherms of forage organisms) (Collins 1952; Leggett and Whitney 1972); 

however, direct evidence is lacking 

(Richkus 1974b). Feeding and vertical 

migration are probably controlled by 

light intensity patterns within thermal 

preference zones (Richkus and Winn 

1979; Neves 1981). 

GROWTH CHARACTERISTICS 

Growth Rates 

Growth rates (L in mm, wt in g) 

of young-of-the-year blueback herring 

in the James River, Virginia, are 

given below. Blueback herring 

achieved a mean fork length of 35.6 

mm and weight of 3.68 g by 15 
November of their first year (Burbidge 1974) Growth rates of 

young-of-the-year blueback herring in 

the Cape Fear River, North Carolina, 

were similar to those for the James 

River population (Davis and Cheek 

1966). 

Change in L Change in wt 

Period per day (mm) per day (g) 

Mid-June to 

mid-July 0.29 0.03 

Mid-July to 

mid-Aug. 0.12 0.01 

Mid-Aug. to 

mid-Sept. 0.15 0.01 

Mid-Sept.to 

mid-Oct. 0.29 0.05 

Mid-Oct. to 

mid-Nov. 0.32 0.05 

Young-of- the-year alewives. apparently grow faster than blueback 

herring. In the Northeast Cape Fear 

River, North Carolina, young alewives 

reached a mean fork length of 44.0 

mm and 47.8 mm in August of 1964 

and 1965, respectively (Davis and 

Cheek 1966). Certainly some of the 

apparently faster growth of young 

alewives is due to an earlier spawning 

period and therefore a longer growing 

season. Growth of young alewives 

between hatching and fall emigration 

from nursery areas averaged 102 mm 

TL in lower Chesapeake Bay (Joseph 

and Davis 1965), and 113 mm TL in 

the Connecticut River (Marcy 1969). 

Average length of juvenile emigrants, 

sampled daily over three seasons 

(1970-72) in Hamilton Reservoir, 

Rhode Island, ranged from 25 mm SL 

to 88 mm SL (30 mm TL to 105 mm 

TL) (Richkus 1975). 

Little information is available on 

growth rates of these clupeids 

between age 0+ and the time of first 

spawning. Age 1 alewives reached 

147 mm TL by the end of their second 

summer in the Connecticut River 

Estuary; however, only males of that 

age were collected (Marcy 1969). Age 

2 alewives and blueback herring in 

Albemarle Sound, North Carolina, 

reached 153 mm FL and 148 mm FL, 

respectively, by the end of their 

third summer (Kornegay 1978). On 

Georges Bank, age 2 fish of both 

species reached approximately 180 mm 

TL by the end of their third summer 

(Netzel and Stanek 1966). 

Lengths at age for sexually 

mature alewives and blueback herring 

are given in Table 1. In general, 

alewives are longer than blueback 

herring of the same age. Within each 

species males are smaller than females 

of the same age, and growth rates for 

both species level off after reaching 

sexual maturity (compared to growth 

rates of immature fish). Mean weights 

of spawning alewives in Damariscotta 

Lake, Maine, ranged from 153 g (5.4 

oz) (males) and 164 g (5.8 oz) 

(females) at age 3, to 325 g (11.5 oz) 

(males) and 356 g (12.6 oz) (females) 

at age 7. One 8-year-old female 

weighed 455 g (16.0 oz) (Walton 

1979). 

Otolith and scale-aging techniques for alewives and blueback herring from Albemarle Sound, North 

Carolina, 

classes 0, 

agreed nearly 100% for age 

1, and 2, but consistency 

between methods decreased progressively as fish age exceeded 3 years 

(Kornegay 1978). The scale-aging 

technique tended to underestimate the 

proportion of age 4 and 5 individuals 

and overestimate the proportion of age 

6 and 7 individuals for both species. 

Growth rates backcalculated from 

scales tended to be higher than those 

backcalculated from otoliths. Regressions for prediction of fork length 

from scale and otolith measurements 

are available in Kornegay (1978). 

Messieh (1977) gave von Bertalanffy growth equations for alewives 

and blueback herring in the St. John 

River, New Brunswick, as follows: 

10 
Table 1. Length (mm FL) at age (yr) for selected Atlantic coast spawning populations of alewives and blueback herring. 

Locationa Species/sexb 

Length (mm FL) at age (yr) 

3 4 5 6 7 8 9 

AS A/M 

AS A/F 

AS B/M 

AS B/F 

CB A/M 

CB A/F 

CR A/M 

CR A/F 

CR B/M 

CR B/F 

GBC A/MF 

GBC B/MF 

DL A/M 

DL A/F 

236 

254 

231 

245 

229 

239 

258 

261 

270 

240 

260 

273 

249 256 259 268 

261 270 277 283 

241 245 251 258 

252 258 264 268 

239 249 254 259 

249 259 264 274 

265 278 290 301 

284 284 299 308 

266 280 286 298 

277 291 301 311 

284 294 306 316 

269 281 292 302 

278 301 315 318 

283 309 324 333 

279 

287 

270 

280 

282 

324 

327 

313 

356 

307 

330 

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

and Davis 1965); CR = Connecticut River (Marcy 1969); GB = Georges Bank (Netzel 

and Stanek 1966); DL = Damariscotta Lake, Maine (Walton 1979). 

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

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

'Values in mm TL. 

11 
MA L = 291 &‘(I - e -0.441 (t - O.lQ)) 

FA L = 310.48(1 _ e -0.4OO(t - 0.103) ) 

MB L = 231.33(1 - e-0.59O(t - 0.338) ) 

FB L = 259.85(1 - e -U.469(t - 0.283)) 

where MA is for male alewives, FA is 

for female alewives, MB is for male 

blueback herring, FB is for female 

blueback herring, L = fork length 

(mm) at time t in years, and e is the 

base of natural logarithms. 

Also given were von Bertalanffy 

growth equation parameters and modified Walford plots for four separate 

areas in the St. John River drainage, 

where different spawning stocks were 

suspected. The equations given above 

are for all study areas combined, 

since differences in growth parameters 

among areas were small (Messieh 

1977). 

Length-Weight Relationships 

Length-weight relationships for 

alewives and blueback herring of the 

St. John River, New Brunswick, were 

given in Messieh (1977) as follows: 

Male alewives l0gw=3.23510gL-5.420 

Female alewives logW=3.192logL-5.294 

Male blueback logW=2.904logL-4.702 

Female blueback logW=2.472logL-3.693 

L=Fork length in mm, W=Weight in g 

Condition factors (K) for 

young-of-the-year blueback herring of 

the James River, Virginia, averaged 

1.60, 1.68, 1.57, 1.37, 1.22, and 1.18 

for the months of June, July, August, 

September, October, and November, 

respectively (Burbidge 1974). It was 

hypothesized that a massive flood in 

the study area between August and 

September may have reduced zooplankton availability which, in addition 

to normal seasonal declines in zooplankton populations, resulted in the 

observed deterioration in the condition 

factor (Bu rbidge 1974). 

THE FISHERY 

Commercial Fisheries 

u. s. commercial landings of 

river herring (both species combined) 

along the Atlantic coast were 4,948 mt 

in 1980 and 3,754 mt in 1981. These 

landings were worth $779,000 and 

$671,000, respectively. The current 

5-year running average of river herring landings (1977-81) by U. S. 

fisheries was 5,003 mt/yr. More than 

90% of the U. S. commercial catch 

occurred within 4.8 km (3 mi) of the 

coast (National Marine Fisheries Service, NMFS, 1982). They are commercially fished during spawning runs. 

Pound nets are the most commonly 

used gear (Joseph and Davis 1965; 

Pate 1974). The majority of U. S. 

landings was used for fish meal and 

fish oil to be added to fertilizer, pet 

food, and domestic animal feed. A 

minor portion was 

bait, 

used for fishing - 

and the remainder was sold 

salted or fresh for human consumption . Roe from these species is 

canned and is highly valued (Joseph 

and Davis 1965; Street and Davis 

1976; Merriner 1978). 

The total foreign catch of river 

herring within the U. S. Fishery 

Conservation Zone (FCZ) was 24.6 mt 

in 1980 and 13.9 mt in 1981. Cuba 

took greater than 90% of the 1980 foreign landings, and Poland captured 

approximately 75% of the 1981 foreign 

catch. All of the 1980 and 1981 foreign landings were taken from the 

North Atlantic region (north of Cape 

Hatteras) (NMFS 1982). 

In domestic commercial fisheries 

in Albemarle Sound, North Carolina, 

surveyed in 1972, age 4 and 5 fish 

represented 60% or more of the 

12 
inshore catch of alewives and blueback 

herring (Pate 1974). Significant contributions also occurred from age 3, 

6, and 7 fish (at least 30% of total 

catch) . 

* herring 

Ratio of alewives to blueback 

in the 1972 catch was 2:3, 

and sex ratios within species were 

near 1:l. First - time spawners accounted for 50% and 57%, respectively, of the landings of blueback 

herring and alewives. Historically, 

the Chowan River has been the most 

important inshore fishing grounds in 

Albemarle Sound. Peak landings usually occur in late April, coinciding 

with the spawning runs of the more 

common blueback herring in North 

Carolina (Pate 1974). In the Potomac 

River, Maryland, the 1974 commercial 

catch was dominated by alewives in 

March and early April, while blueback 

herring landings peaked in late April 

and May. The total ratio of alewives 

to blueback herring in the 1974 catch 

was 1:4 (Merriner 1978). 

Total landings and catch per unit 

effort in North Carolina and Virginia 

waters have declined substantially 

over the last decade since peaking in 

& 1969 at 35,302 mt (Street and Davis 

1976; Merriner 1978). The decline 

was due to offshore trawl fisheries, 

which did not begin operating until 

1967 (McCoy 1975). These fisheries 

were not and are not size selective, 

and 65% of the 1975 offshore trawl 

landings of river herring consisted of 

immature fish (Street and Davis 1976). 

In contrast, all inshore commercial 

catches historically were captured 

from sexually mature populations, 

where escapement rates can be controlled to prevent overexploitation 

(McCoy 1975). 

The State of Maine has developed 

an Alewife Management Plan for anadromous stocks within their State 

jurisdiction (Walton et al. 1976). Historical landings and management are 

reviewed and management recommendations presented for the alewife runs of 

each coastal county. Aspects of life 

history, biology, and management of 

Virginia and North Carolina stocks of 

anadromous alewives and blueback 

herring were reviewed by Ruiifson 

and Huish (1982). In addition, recommendations for development of a management plan are presented and discussed. 

Recreational Fisheries 

Recreational fishing for alewives 

and blueback herring is significant 

during spring spawning runs in areas 

such as Delaware Bay, Chesapeake 

Bay, and Albemarle Sound (Pate 

1974). The disposition of the catch, 

however, is not well documented. 

Apparently, most of the recreational 

catch serves as bait for other sportfish (NMFS 1980). The numbers and 

weights of the recreational catch for 

each species are unknown, because 

published surveys lump these two 

species with menhadens, shads, other 

herrings, and sardines. The NMFS 

estimated that 6,169,OOO total “herrings” were captured by Atlantic coast 

recreational fishermen in 1979. 

Population Dynamics 

Sex ratios/age structure. On 

spawning runs, total sex ratios of 

adults are nearly 1 :l in most areas. 

Percentage of male alewives in the 

spawning populations of Bride Lake, 

Connecticut, was 55.6% (Kissil 1974), 

compared to 53.8% and 53.0% for alewives and blueback herring in the 

Connecticut and Thames Rivers (Marcy 

1969), and 58.0% in a later study on 

Thames River blueback herring 

(Loesch and Lund 1977). It has been 

suggested that the slight dominance of 

males in spawning populations is due 

to their earlier sexual maturity compared to that of females (Kissil 1974). 

Sex ratios (% males) by age for 1966 

and 1967 spawning populations of alewives from the Connecticut and 

Thames Rivers, Connecticut, (data 

combined) were 72.3%, 63.7%, 49.7%, 

33.8%, and 0.0% for fish of age 4, 5, 

13 
6, 7, and 8, respectively. Ratios for 

blueback herring were 80.0%, 79.4%, 

64.5%, 36.9%, and 22.6% for fish of 

age 3, 4, 5, 6, and 7, respectively 

(Marcy 1969). 

Life stage abundance/reproductive 

and mortality rates. In Bride Lake, 

Connecticut, an estimated 184,15 

adult alewives spawned 2.05 x lOI a 

e99s in 1966. Subsequently, 257,000 

juveniles were counted as they emigrated during the summer and fall. 

This generated a tota I freshwater 

mortality rate from egg stage to emigration of 99.9987%, and indicated that 

2.88 juveniles left the lake for each 

adult female that spawned. Combined 

with repeat spawning proportions, this 

level of juvenile production seemed 

adequate for sustaining the spawning 

population in Bride Lake (Kissil 1974). 

Tota I freshwater mortality rate of 

spawning adults in Bride Lake was 

reported at 57.4% and 48.6%, in 1966 

and 1967, respectively. 

Havey (1973) investigated juvenile production and adult mortality of 

the alewife population in Love Lake, 

Maine. The number of emigrants 

ranged from 220 to 439,062 fish over 

an 11-year study period. Juvenile 

emigrants produced per female per 

year varied from 12 to 3,209, and 

biomass production of emigrants 

ranged from 0.09 to 21.50 kg (0.2 to 

47.4 lb) per female per year. Significant linear relationships between 

juvenile emigrant abundance and 

spawning population size 4 years 

later, and between the log of female 

escapement and the log of juvenile 

emigrant abundance were found. 

Total adult freshwater mortality averaged 90.7% and ranged from 66% to 

100% over the 11 -year-study period 

(Havey 1973). Total annual mortality 

rates for anadromous alewives in Long 

Pond, Maine, were 78.6% between age 

5 and 6, and 74.4% between age 6 and 

7 (Havey 1961). Freshwater postspawning mortality for all age groups 

averaged 41% and ranged from 32% to 

67% over the 6-year study period 

(Havey 1961). Results from these two 

studies in Maine and the study by 

Kissil (1974) in Connecticut indicated 

that juvenile production and adult 

freshwater mortality may vary considerably among spawning areas and 

among different years in the same 

spawning area. 

Methods of determining true 

spawning population size of alewives, 

by subsampling at various times 

on fishways on the Parker River, 

Massachusetts, were investigated by 

Rideout et al. (1979). Visual counts 

during one lo-min period each hour 

produced estimates (by computer program) with less than 5% error. Visual 

counts during one 5-min period each 

hour or one lo-min period each 1.5 

hours were significantly higher in 

magnitude of potential error (Rideout 

et al. 1979). 

Stock identification. Although 

Thunberg (1971) found that alewives 

were capable of homing behavior 

through olfaction, Messieh (1977) concluded that considerable mixing 

between presumed spawning stocks 

occu r red on the spawning runs. 

Though a majority of the alewives 

exhibited homing, a substantial number (by meristic comparisons) from 

each presumed stock did not home 

(Messieh 1977). 

Evidence for a nonlandlocked, 

non migratory, self-sustaining dwarf 

population of alewives residing in the 

mouth of the Susquehanna River was 

presented by Foerster and Goodbred 

(1978). 

ECOLOGICAL ROLE 

Food Habits 

Alewives and blueback herring 

are primarily zooplanktivores, though 

fish eggs, crustacean eggs, insects 

and insect eggs, and small fishes may 

14 
be important foods in some areas or 

for larger individuals (Bigelow and 

Sch roeder 1953) . Larvae begin feeding 

on zooplankton immediately upon formation of a functional mouth (about 6 

mm TL), concentrating on the relatively small cladocerans and copepods, 

and adding larger species of these 

groups to the diet as their mouths can 

accommodate them (Norden 1968; Nigro 

and Ney 1982). 

Stomachs from young-of-the-year 

blueback herring cot lected in the 

James River, Virginia, contained primarily (by volume) Bosmina spp., 

copepod nauplii, copepodites, and the 

adult copepods Eurytemora affinis and 

Cyclops vernalis. Diaphanosoma brachyurum and Canthocamptus robertcokeri were also minor food items, but 

were not utilized during all seasons 

(Burbidge 1974). Electivity (lvlev 

1961) was strongest for adult copepods; neutral for Bosmina spp., copepodites, and g. brachyu rum; and 

strongly negative for copepod nauplii. 

Daily ration for young-of-the-year 

ranged from 438 g-cal per fish per 

day in July and August to 215 g-cal 

per fish per day in October and 

November (Bu rbidge 1974). 

Young-of-the-year alewives in 

Hamilton Reservoir, Rhode Island, 

consumed primarily chironomid midges 

during July, switching to cladocerans 

in August and September (Vigerstad 

and Cobb 1978). Davis and Cheek 

(1966) compared the food habits of 

young-of-the-year alewives and blueback herring in the Cape Fear River, 

North Carolina. Blueback herring 

selected copepods and dipteran larvae 

more frequently (by percent of stomachs containing items) than did alewives, while alewives consumed more 

ostracods, insect e99s I and insect 

parts than did blueback herring. 

Crustacean eggs in the diets were 

similarly common (‘80% of stomachs) 

for both clupeids. Overlap of either 

diet with that of young-of-the-year 

American shad Alosa sapidissima was 

not extensive ,because of 

range of foods chosen by 

and Cheek 1966). 

the broader 

shad (Davis 

Few direct studies have been 

devoted to food habits of anadromous 

adult alewives and blueback herring. 

In general, they are zooplan ktivores, 

with the size range and diversity of 

available prey increasing as the fish 

g row and can accommodate larger 

items. Considerable piscivority may 

develop in landlocked populations 

(Kohler and Ney 1981). 

Feeding Behavior 

Alewives and blueback herring 

feed in schools of varying size, most 

extensively during daylight. At 

night, schools disperse, and feeding 

activity is negligible or casual, and 

probably by filter feeding (Burbidge 

1974; Janssen 1978). Peak stomach 

fullness occurred at 6 PM and peak 

stomach emptiness occurred at 7 AM in 

blueback herring from the James 

River, Virginia (Burbidge 1974). In 

addition, young-of-the-year blueback 

herring fed more actively near the 

surface than at 5 m (16 ft) depth, 

even though high densities were present at both depths (Burbidge 1974). 

In laboratory tests (Janssen 

1976), alewives exhibited three feeding modes: (1) particulate feeding on 

individual prey, (2) filter feeding 

with mouth agape and rapid swimming 

bursts, and (3) gulping several prey 

at once but not swimming at the rapid 

speed used in the filter-feeding mode. 

Size selectivity of prey items was 

highest in mode (l), moderate in mode 

(3), and negligible in mode (2). Adult 

Lake Michigan alewives and a major 

food organism, Mysis relicta, exhibited 

coincidental, crepuscular vertical 

migrations, from daytime residence 

near the bottom to just below the 

thermocline at night (Janssen and 

Brandt 1980). Such a diel vertical 

migration, though probably not in 

relation to a thermocline, may occur in 

15 
anadromous populations residing in 

estuaries or marine coastal habitats. 

Competitors 

Little study has been devoted to 

competitive interactions of anadromous 

alewives or blueback herring. 

Because of general similarities in diet 

and feeding behavior, some competi - 

tion for food Ii kely occurs between the 

two species. Loesch et al. (1982a) 

described a spatial separation between 

young alewives and blueback herring 

in the same habitat, which may lead to 

reduced competition for food, at least 

among juveniles. 

Predators 

Alewives and blueback herring 

are highly utilized forage species for 

many riverine, estuarine, and marine 

piscivores, including airborne predators such as gulls and terns (Commonwealth of Massachusetts 1976). 

Bluefish (Pomatomus saltatrix), weakfish (Cynoscion regalis), and striped 

bass (Morone saxatilis) are predators 

of these clupeids. Pelagic, schooling 

predators such as these are more 

likely to use schooling clupeids for 

forage compared to a solitary predator 

(Cooper 1961; Tyus 1974). 

ENVI RONMENTAL REQU I REMENTS 

Some research has been conducted to delineate the specific environmental requirements of anadromous 

alewife and blueback herring. Much of 

the available information was derived 

from tests on landlocked populations 

(particularly Lake Michigan alewives). 

Applicability of environmental requirement data for landlocked populations 

to anadromous populations is 

unknown. Since data from landlocked 

populations are major sources of 

information on environmental requirements, they are presented but should 

be interpreted with caution. 

Temperature 

The effects of incubation temperature on alewife eggs from Lake Michigan were studied by Edsall (1970). 

At least some eggs hatched at test 

temperatures between 7’ and 29.5 o C 

(44.6 O and 85.1 OF). Optimum incuba-. 

tion temperature for hatching was 18’ 

C (64OF); 38% hatch was observed. 

Egg mortality over the first 36 hr 

ranged from 22% at temperatures 

between 3.5 O and 6.0°C (38’ and 43’ 

F) to 66% at temperatures between 

25.5’ and 28.5OC (78’ and 83OF). Egg 

mortality rate was directly correlated 

to incubation temperature (Edsall 

1970). An upper lethal temperature 

of 29.7 O C (85’ F) was reported for 

alewife eggs from the Hudson River, 

New York (Kellogg 1982). Maximum 

percentage hatch occurred at 20.8OC 

(69 O F), and at least some eggs 

hatched at test temperatures between 

12.7’ and 26.7OC (55’ and 8OOF). 

Blueback herring eggs collected 

from the Washademoak River, New 

Brunswick, Canada, were subjected to 

time-temperature regimes experienced 

in a powerplant cooling system (Koo 

and Johnston 1978). Compared to 

larval deformity rate, egg mortality 

and hatchability were not good indicators of the effects of temperature 

change. Deformity rate of larvae, 

acclimated at 19OC (66OF) and exposed 

to a 10°C (18OF) increase in temperature for 5 to 180 min, varied from 

0 to 25% (control O-5%). Deformity 

rate increased to 100% under the same 

conditions except with a 15OC (27OF) 

temperature elevation. Deformities 

ranged from minor curvature of the 

spine to complete lack of normal larval 

form or behavior. Deformities were 

permanent and would not have allowed 

such larvae to survive in natural 

environments ( Koo and Johnston 

1978). 

Edsall (1970) reported two 

aspects of temperature effects on 

16 
larval alewives from Lake Michigan. 

Survival time of unfed larvae was 3.8 

days at 10.5OC (51°F), 7.6 days at 

14.5’ to 15OC (58’ to 59OF), and 2.4 

& days at 26.5Oto. 28” C (80’ to 82’ F) . 

A functional jaw did not develop in 

fish from eggs/larvae held at or below 

10°C (50°F), even though some eggs 

hatched at such temperatures. Kellogg (1982) reported an upper temperature tolerance of 31°C (88OF) for 

alewife larvae from the Hudson River, 

New York, acclimated to 14OC (57OF). 

Average daily gain in larval weight 

was directly proportional to water 

temperature; higher growth occurred 

at higher temperatures. A maximum 

larval growth rate of 0.084 g/day 

occurred at 29.1°C (84OF), while maximum net gain in biomass (a function 

of both survival and growth) occurred 

at 26.4OC (79.5OF) (Kellogg 1982). 

Young-of-the-year alewives (19 to 

31 mm TL) from the Hudson River, 

New York, preferred a water temperature of 26.3OC (79” F) when given a 

choice in a controlled thermal gradient 

(Kellogg 1982). Young-of-the-year 

& 

alewives from Lake Michigan exhibited 

critical thermal maxima (CTM is the 

mean of temperatures at which experimental fish lose equilibrium) of 28.3OC 

(83OF), 32.7OC (91°F), and 34.4OC 

(94OF) at acclimation temperatures of 

ll°C (52OF), 19°C (66OF), and 25OC 

(77’ F), respectively (Otto et al. 

1976). The equation for predicting 

CTM from acclimation temperature was : 

CTM = 21.9 + 0.5(TA) r2 = 0.96 

where TA = acclimation temperature in 

degrees Celsius. 

Otto et al. (1976) also reported that 

CTM values were 3’ to 6OC (5.4Oto 

10.8OF) higher for young-of-the-year 

alewives than for adults when tested 

at the same acclimation temperatures. 

In laboratory tests, preferred 

(selected) temperatures of juvenile 

alewives and blueback herring (ages 

0+ and 1’) collected from the Delaware 

River, New Jersey, ranged from 20’ 

to 22°C (68’ to 71’ F) at salinities of 4 

to 6 ppt and acclimation temperatures 

from 15’ to 21°C (59” to 70°F) (Meldrim and Gift 1971). Davis and 

Cheek (1966) captured juvenile blueback herring in the Cape Fear River 

seasonally in areas where water temperatures ranged from 11.5’ to 32OC 

(53’ to 89OF). Juvenile alewives in the 

same watershed were captured at temperatures between 13.5’ and 29OC (56’ 

and 84“F). 

School formation patterns and 

daily rhythms of adult Lake Michigan 

alewives were affected by changes in 

temperature in laboratory tanks 

(Colby 1971). As water temperature 

dropped below 6.7OC (44OF), normal 

feeding behavior was disrupted and 

cruising speed of schooling fish 

decreased. Below 4.5’C (40°F), normal schooling behavior was significantly affected. At temperatures 

between 2.0’ and 2.8OC (35.5’ and 37O 

F), alewives lost orientation, swam 

into the sides of the test chamber, 

and ceased feeding and schooling. 

In cold shock tests with adult 

alewives from Lake Michigan, transfers 

to test temperatures less than 3 O C 

(37.4 O F) caused 100% mortality 

regardless of original acclimation temperatures (Otto et al. 1976). Magnitude of temperature-decrease tolerated 

increased gradually with increasing 

acclimation temperature. At least 

some alewives survived a temperature 

decrease of 10’ C (18’F), independently of acclimation temperature, as 

long as the total test temperature did 

not drop below 3OC (37.4OF) (Otto et 

al. 1976). 

Stanley and Colby (1971) investigated electrolyte balance and osmoregulation of Lake Michigan alewives in 

relation to temperature change. 

Transfers of fish acclimated at warm 

temperatures to cold temperatures 

caused levels of Na+, K+, and Ca++ in 

17 
blood and muscle to move towards an 

equilibrium with salinity of the acclimation environment (increased body 

concentrations in salt water, 

decreased body concentrations in 

freshwater). Apparently, the test fish 

temporarily lost the ability to osmoregulate upon exposure to cold, 

independently of the salinity of the 

test environment. 

Salinity 

Though little direct information 

exists, anadromous alewives and blueback are apparently highly tolerant of 

salinity changes (Cooper 1961; Chittenden 1972) . No mortality of adult 

blueback herring from either gradual 

or abrupt changes in salinity, including direct transfers from fresh to salt 

water and the reciprocal, was 

observed by Chittenden (1972). 

Blood and muscle concentrations of the 

electrolytes Na+, K+, and Ca++ were 

similar in fish held in sea water and 

freshwater of the same temperature, 

indicating that after a period of accli-. 

mation, alewives were efficient osmoregulators in either environment 

(Stanley and Colby 1971). 

Other Environmental Factors 

Location of appropriate spawning 

sites and substrates is important not 

only to the perpetuation of each species but also for natural “reproductive 

segregation” between two otherwise 

very similar species. Blueback herring prefer spawning sites with strong 

currents and associated hard substrates (Loesch and Lund 1977). 

They are relatively specialized compared to alewives, which use a wide 

variety of spawning sites, from standing river water, oxbows, coastal 

ponds, and tiny streams to fastwater, mid- river sites. Therefore, 

changes in water currents or substrates in spawning rivers used by 

blueback herring may affect that species more than the alewife, because of 

the more specific spawning site 

requirements of blueback herring. 

Young-of-the-year alewives and 

blueback herring from the Cape Fear 

River system, North Carolina, selected areas where free carbon dioxide 

ranged from 4 to 22 ppm, alkalinity _ 

from 5 to 32 ppm, dissolved oxygen 

from 2.4 to 10.0 ppm, and PH from 

5.2 to 6.8 (Davis and Cheek 1566). 

In an 

‘the effect! 5 

experiment designed to test 

of suspended sediments on 

r I .F hatching ot alewlte eggs (Schubel and 

Wang 1973), a naturally occurring 

fungus in the sediment infected all 

test eggs prior to hatching, and terminated the experiment. Although the 

extent of infection may have been 

enhanced by laboratory conditions, 

the attempt indicated that high levels 

of suspended sediment during or after 

spawning may significantly increase 

infection rates of eggs from naturally 

occurring fungi in sediments (Schubel 

and Wang 1973). Auld and Schubel 

(1978), however, found that suspended sediments in concentrations of 

100 ppm or less had no significant 

effect on hatchability of alewife or’ 

blueback herring eggs. 

The influence of certain environmental variables associated with passage of migratory adult alewives 

through (around) hydrological obstacles has been investigated. Blood lactic acid concentrations, measured in 

alewives moving through a pool-andweir fishway, were representative of 

moderate activity and energy expenditure (Dominy 1971, 1973). Mean levels of blood lactic acid in alewives 

passing through the fishway were less 

than half the levels found for heavily 

exercised fish in the laboratory. Rest 

pools along the course of the fishway 

allowed blood lactic acid levels to drop 

to levels comparable with those for 

alewives in a rested state in the laboratory . 

Upstream migratory patterns of 

adult alewives through a Rhode Island 

18 
river fishway were harmonic with 

diurnal periodicity. Periodicity was 

correlated with magnitude of incident 

solar radiation (Saila et al. 1972). 

k Richkus (1974a) corroborated 

light dependent migratory actlvlty; 

thi; 

also observed that within activity patterns determined by light intensity, 

changes in water temperature strongly 

influenced specific timing of alewife 

upstream movement. Juvenile downstream emigration from Hamilton Reservoi r, Rhode Island, during summer 

and fall was inhibited by the bright 

sunlight-bridge shade interface present at a road bridge on the lower end 

of the reservoir. Higher emigration 

rates under this bridge were observed 

on cloudy days (Richkus 1974a). 

Environmental Contaminants 

The LCSO of total residual chlo9 

rine for blueback herring eggs ranged 

from 0.20 to 0.32 ppm. Larvae from 

eggs exposed to sublethal concentrations of total residual chlorine were all 

deformed (Morgan and Prince 1977). 

Concentrations of kepone greater than 

0.3 ppm (termed the “action level” for 

possible closure of a fishery) were 

found in body tissues of young-ofthe-year alewives and blueback herring collected from the James and 

Chickahominy Rivers, Virginia (Johnson et al. 1978; Loesch et al. 

1982b). Kepone was also present in 

young alewives and blueback herring 

from the Mattaponi and Pamunkey Rivers, Virginia (in concentrations less 

than 0.3 ppm), but was not present 

in detectable quantities in fish from 

the Rappahannock River, Virginia, 

and the Potomac River, Maryland 

(Loesch et al. 1982b). 
LITERATURE CITED 

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373-396. -i 

Berry, F. H. 1964. Review and emendation of family Clupeidae. Copeia 1964: 720-730. 

Commonwealth of Massachusetts. 1976. 

Anadromous fish of Massachusetts. Mass. Dep. Fish., 

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Bigelow, H. B., and W. C. Schroeder. 1953. Fishes of the Gulf of 

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Fish. Bull. No. 74. 577 pp. 

Bigelow, H. B., and W. W. Welsh. 

1925. Fishes of the Gulf of 

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No. 40. 567 pp. 

Cooper, R. A. 1961. Early life history 

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58 PP. 

Burbidge, R. G. 1974. Distribution, 

growth, selective feeding and 

energy transformations of 

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103: 297-311. 

Davis, J. R., and R. P. Cheek. 1966. 

Distribution, food habits and 

growth of young clupeids, Cape 

Fear River system, North Carolina. Paper presented at Annu. 

Meet. South. Div. Am. Fish. 

sot., Asheville, N. C., 1966. 20 

PP. 

Chambers, J. R., J. A. Musick, and 

J. Davis. 1976. Methods of distinguishing larval alewife from 

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Dominy, C. L. 1971. Changes in blood 

lactic acid concentrations in alewives during passage through a 

pool-and-weir fishway. J. Fish. 

Res. Board Can. 28: 1215-1217. 

Chittenden, M. E. 1972. Salinity tolerance of young blueback herring. Trans. Am. Fish. Sot. 

101: 123-125. 

Dominy, C. L. 1973. Effect of 

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Edsall, T. A. 1970. The effect of 

temperature on the rate of 

20 
development and survival of alewife e99s and larvae. Trans. 

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b Edsall, T. A., and M. I. Saxon. 

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PP. 

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& 434-437. 

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1972-1975. 

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23 PP. 

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44: 63-73. 

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Kissil, G. W. 1974. Spawning of the 

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Fish. Sot. 103: 312-317. 

Kohler, C. C., and J. J. Ney. 1981. 

Consequences of an alewife dieoff 

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110: 360-369. 

21 
Koo, T. S. Y., and M. L. Johnston. 

1978. Larval deformity in striped 

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137-149. 

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valis. M. S. Thesis. University 

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Loesch, J. G. 1969. A study of the 

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43(2): 77-79. 

Loesch, J. G., and W. A. Lund. 

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Loesch, J. G., W. H. Kriete, Jr., 

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Loesch, J. G., R. J. Hugget, and E. 

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MacLellan, P., G. E. Newsome, and 

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544-546. 

Mansueti, R. J. 1956. Alewife herring 

eggs and larvae reared successfully in lab. Maryland Tidewater News 13(l): 2-3. 

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with comments on its ecology in 

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173-205. 

Marcy, B. C., Jr. 1969. Age determination from scales of Alosa pseu- ___- 

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in Connecticut waters. Trans. 

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65-68. 

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22 
Meldrim, J. W., and J. J. Gift. 1971. 

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b 

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Bank, July and October, 1964. 

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5 PP. 

Neves, R. J. 1981. Offshore distribution of alewife and blueback herring along the Atlantic coast. U.S. 

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111: 559-569. 

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behavior and growth of juvenile 

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Fish. Sot. 104: 483-493. 

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1971. Effects of temperature on 

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Street, M. W., and J. 

Notes on the river 

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Tyus, H. M. 1974. Movements and 

spawning of anadromous alewives 

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103: 392-396. 

24 
Vigerstad, T. J., and J. S. Cobb. 

1978. Effects of predation by 

sea-run juvenile alewives on the 

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Resou r. Res. Ref. Dot. No. 

79/23. 75 pp. 

Walton, C. J., M. E. Smith, and D. 

B. Sampson. 1976. Alewife management plan. Maine Dep. Mar. 

Res. Job Compl. Rep. No. 

AFSC-13/,FWAC-2. 37 pp. 

Warinner, J. E., J. P. Miller, and J. 

Davis. 1969. Distribution of juvenile river herring in the Potomac 

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23: 384-388. 

Winters, G. H., J. A. Moores, and R. 

Chaulk. 1973. Northern range 

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30: 860-861. 

25 
3ulll-I”I 

REPORT DOCUMENTATION ~ 1. REPORT NO. 

~ 

~ 2. 

i 

3 Reclplent’s Accesslo” NO 

PAGE FWS/OBS-82/11.9* 

4. iltle and Subtitle 

Species Profiles: 

5. Report Date 

Life Histories and Environmental Requirements of 

Coastal Fishes and Invertebrates (Mid-Atlantic) -- Alewife/Blue_ Is, 

October 1983 

back Herring 

7. Author<%, ) 8. PerformIng Organization wept. NO. 

Clemon W. Fay, Richard J. Neves, and Garland B. Pardue 

i. Performing Organlzatlo" Namu and Address 10. Pro,ect/TasklWork “n,t No. 

Department of Fisheries and Wildlife Sciences 

Virginia Polytechnic Institute and State University 11. Contract(C) or GrantG, No. 

Blacksburg, VA 24061 (C) 

(Gl 

N’~t~~,‘,“‘i”“~~~~io”~~m,~~~tAed~~*STeam 

U.S. Army Corps of Engineers 

13. Type of Report & Period Coverqd 

Fish and Wildlife Service Waterways Experiment Station 

U.S. Department of the Interior P.O. Box 631 

Washington, DC 20240 Vicksburg, MS 39180 14. 

__-___ 

15. Supplementary Notes 

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

16. Abstract(Limlt:zoowords) 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 as 

links between the zooplankton they consume and top piscivores. Both anadromous and landlocked populations exist. Some individuals mature by age 3, and all mature by age 5. Repea 

spawning is common. Spawning environments range from streams only a few centimeters deep tc 

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 mid-Atlantic region; the 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 between June and 

November of 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 the Gulf of Maine are important overwinter 

ing grounds. Commercial and limited recreational fisheries for these species occur; total 

U.S. landings in 1981 were 3,754 mt, while foreign landings were 13.9 mt. Some eggs can 

hatch at water temperatures between 7" and 29.5"C, but temperatures above 29.7"C are lethal. 

Larvae need temperatures greater than 10°C for proper development; upper lethal temperature ~___~. 

17. Dacvment Analysis a. Descr,,,fors 

Rivers 

-ts-srfte;-- 

Growth 

Anadromous Feeding 

Estuaries 

Fish 

b. Identlf,ers/Open-Ended Terms 

Alewife Temperature requirement 

Alosa pseudoharengus Salinity requirements 

Blueback herring Spawning 

Alosa aestivalis 

c. COSATI F&d/Group 

__.__. ~_____ 

I’. i\“aliaDIl,ty Statement / 19. Security Class VhlS Report, 

: Unclassified 

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20. Security Class (This Page) . z.Pze ~--.-- 

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Species profiles : 

Fay, Cienon k. 

Headquarters, Division of Biological 

Services, Washington, DC 

Eastern Energy and Land Use Team 

Leetown, WV 

National Coastal Ecosystems Team 

Slidell, LA 

Western Energy and Land Use Team 

Ft. Collins, CO 

Locations of Regional Offices 

REGION 1 

Regional Director 

U.S. Fish and Wildlife Service 

Lloyd Five Hundred Building, Suite 1692 

500 N.E. Multnomah Street 

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REGION 2 

Regional Director 

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P.O. Box 1306 

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REGION 3 

Regional Director 

U.S. Fish and Wildlife Service 

Federal Building, Fort Snelling 

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Regional Director 

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One Gateway Center 

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REGION 6 

Regional Director 

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P.O. Box 25486 

Denver Federal Center 

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REGION 7 

Regional Director 

US Fish and Wildlife Service 

1011 E. Tudor Road 

Anchorage, Alaska 99503 
DEPARTMENT OFHE INTERIOR 

U.S. FISH AND WILDLIFE 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 resources, protecting our fish and wildlife, 

preserving theenvironmental and cultural values of our national parks and historical places, 

and providing for the enjoyment 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 Department also has a major responsibility for 

American Indian reservation communities and for people who live in island territories under 

U.S. administration.