Alosa pseudoharengus (Wilson, 1811)

Common Name: Alewife

Synonyms and Other Names:

mulhaden, grey herring, golden shad, kyak, sawbelly, grayback, river herring

U.S. Fish and Wildlife ServiceCopyright Info

U.S. Fish and Wildlife ServiceCopyright Info

Identification: The Alewife is a small herring with a dark dorsal side, bluish to greenish, and light sides with horizontal darker stripes. The head is broad and triangular and the body is relatively deep. Eyes are large with adipose eyelids. A dull black spot is located behind the operculum. Scales are easily rubbed off and form scutes on the midline of the belly. Jaw teeth are inconspicuous and tongue teeth are absent. Caudal fin is forked and lacks an adipose fin. Alewife are visually similar to Blueback Herring (Alosa aestivalis), but Alewife has a white peritoneal lining, larger eyes, and a greater body depth than Blueback Herring (Whitehead 1985; Page and Burr 1991; Etnier and Starnes 1993; Jenkins and Burkhead 1994; Scott and Crossman 1998).

Size: Total length up to 38 cm, but inland populations are usually less than 25 cm. Fertilized eggs have a diameter around 0.9 mm and larvae are on average 3.8 mm long at hatch (Henrich 1981).

Native Range: Atlantic Coast from Red Bay, Labrador, to South Carolina; many landlocked populations (Page and Burr 1991).

Great Lakes Nonindigenous Occurrences: Alewife has been recorded in all five Great Lakes, however, the native status of Alewife in Lake Ontario is under debate (Roth et al. 2013).

Table 1. Great Lakes region nonindigenous occurrences, the earliest and latest observations in each state/province, and the tally and names of HUCs with observations†. Names and dates are hyperlinked to their relevant specimen records. The list of references for all nonindigenous occurrences of Alosa pseudoharengus are found here.

Full list of USGS occurrences

State/ProvinceFirst ObservedLast ObservedTotal HUCs with observations†HUCs with observations†
IL194920183Lake Michigan; Little Calumet-Galien; Pike-Root
IN195620142Lake Michigan; Little Calumet-Galien
MI1933201418Betsie-Platte; Betsy-Chocolay; Carp-Pine; Detroit; Fishdam-Sturgeon; Keweenaw Peninsula; Lake Erie; Lake Huron; Lake Michigan; Lake St. Clair; Lake Superior; Ontonagon; Pere Marquette-White; Raisin; St. Clair; St. Marys; Sturgeon; Waiska
MN195620174Baptism-Brule; Beaver-Lester; Lake Superior; St. Louis
NY1868201512Black; Chaumont-Perch; Irondequoit-Ninemile; Lake Champlain; Lake Erie; Lake Ontario; Oak Orchard-Twelvemile; Raisin River-St. Lawrence River; Salmon-Sandy; Saranac River; Seneca; St. Regis
OH193120157Ashtabula-Chagrin; Black-Rocky; Cedar-Portage; Chautauqua-Conneaut; Huron-Vermilion; Lake Erie; Sandusky
PA193120061Lake Erie
VT199720202Lake Champlain; Mettawee River
WI1952201710Beartrap-Nemadji; Door-Kewaunee; Lake Michigan; Lake Superior; Lower Fox; Manitowoc-Sheboygan; Milwaukee; Peshtigo; Pike-Root; St. Louis

Table last updated 5/16/2022

† Populations may not be currently present.

* HUCs are not listed for areas where the observation(s) cannot be approximated to a HUC (e.g. state centroids or Canadian provinces).

Ecology: Alewife is anadromous and euryhaline (prefers <15 psu), but occurs in landlocked water bodies including the Great Lakes (DiMaggio et al. 2016). It exists at various depths ranging throughout the year from littoral to profundal zones depending on the season. In spring, Alewife spreads out across a lake, staying in warmer waters above the thermocline in schools and dispersing shoreward at night to spawn near the surface of open lake shores, bays, harbors, and lower reaches of rivers (O’Gorman et al. 2013). When near shore, Alewife prefers rocky substrates to sandy substrates (Janssen and Kuebke 2004). During summer, young-of-year (YOY) Alewife stays in the warmer epilimnion while older fish will often venture into the thermocline and cooler waters (Wells 1968; Otto et al. 1976; Johannsson and O’Gorman 1991).  In fall, Alewife moves away from shore and into deeper waters as the thermocline descends and weakens and eventually overwinters in the profundal zone. Near the end of winter,  Alewife begins to move shoreward once again to repeat the cycle (Wells 1968; Bergstedt and O’Gorman 1989).

Reproduction of Alewife is polygynandrous and lasts for around a month in spring once water temperatures surpass ~15ºC (Edsall 1970; Hlavek and Norden 1987). Landlocked populations generally mature a year faster than anadromous populations, taking two years for males and three for females. Females deposit between 10,000 and 360,000 eggs (typically on the lower end) at random on any type of substrate (Hlavek and Norden 1987; Scott and Scott 1988; Scott and Crossman 1998). Eggs are non-adhesive and incubation time varies with temperature, from 15 days at 7.2ºC to 3.7 days at 21.1ºC (Edsall 1970). Larvae are phototropic, pelagic, and begin feeding two days after hatching (Odell 1934; Heinrich 1981).

Alewife can survive in temperatures between 3 to 31ºC, but prefer waters between 16 to 20ºC (Otto et al. 1976; Spotila et al. 1979; Dufour et al. 2008). Water above 31ºC and below 3ºC can cause extreme stress and eventual death (Colby 1973; Otto et al. 1976; McCauley and Binkowski 1982). Lepak and Kraft (2008) found that Alewife experienced sublethal immunosuppression when held in ponds below 2ºC for six weeks. Die-offs of Alewives often occur in the Great Lakes and are typically concurrent with severe winter conditions and/or when fish conditions are poor due to high population densities and increased disease transfer rates (Brown 1972; Colby 1973; Bergstedt and O’Gorman 1989; Lepak and Kraft 2008). Winter temperatures sustained at or below 1ºC can cause osmoregulatory failure due to changes in lipid composition leading to fish mortality. Hence, smaller Alewives with small lipid reserves often experience higher winter mortality than their larger counterparts (Snyder and Hennessy 2003). Therefore, YOY Alewives must be at least 60 mm in total length to successfully overwinter in the Great Lakes, especially in Lake Superior where the growing season is shorter and winter is harsher (Brown 1972; Elrod 1983; O’Gorman and Schneider 1986). As a consequence to the climate in Lake Superior, Alewives there are much fewer in number and much longer than others of the same age in the other Great Lakes due to extreme selection pressure for fast growth (O’Gorman et al. 1997). The warming effects of climate change in Lake Superior are expected to increase the favorable environment for Alewife by reducing the lethality of winter and by promoting food production (Bronte et al. 2003; Hook et al. 2007; O’Gorman et al. 2013).

The diet of Alewife is diverse and impactful, as it is a proficient feeder of eggs, insects, zooplankton, and larval fish. It selects for the largest zooplankton in invaded water bodies and subsequently has altered the species composition of zooplankton in the Great Lakes where Alewife is abundant (Hutchinson 1971; Johannsson et al. 1991). The degree to which Alewife reduces large zooplankton populations is so severe that the seasonal movements of the fish in Lake Ontario have been tracked by the size of zooplankton (O’Gorman et al. 1991). Alewife also preys on larval fish, benefited by its wide-ranging seasonal movements and ability to feed in midwater and low light levels (O’Gorman et al. 2013). Alewife’s predation of fish whose larvae are pelagic can be so severe that their recruitment is drastically reduced (Madenijan et al. 2008). A variety of piscivorous fish consume Alewife in the Great Lakes. Alewife is vital prey for both nearshore (e.g. Yellow Perch (Perca flavescens) and Walleye (Sander vitreus)) and offshore (e.g. Rainbow Smelt (Osmerus mordax), Burbot (Lota lota), and Salmonid) predators (O’Gorman 1974; Elrod et al. 1981; Ridgway et al. 1990).

Means of Introduction: There is apparently disagreement concerning the native status of Alewife in Lake Ontario. Miller (1957) and Smith (1970) point out the first record from Lake Ontario was in 1873. Smith (1970) is of the opinion that it was introduced into the lake. Although Smith (1970) brings up the possibility that Alewife were introduced into Lake Ontario with American Shad stockings in the 1880s, he discounts this possibility in favor of the hypothesis that they reached the lake via the Erie Canal from the Hudson River. He contends that Alewife was only able to invade the lake after the decline of predators such as Lake Trout and Atlantic Salmon in the 1860s. Other authors believe this species was probably native to Lake Ontario (Lee et al. 1980 et seq.) and spread through the Great Lakes via the Welland Canal (Lee et al. 1980 et seq.). The species was first reported from Lake Erie in 1931, Lake Huron in 1933, Lake Michigan in 1949, and Lake Superior in 1954. The spread of Alewives in the upper Great Lakes is thought to be enabled by warmer weather in the 1950’s (O’Gorman and Stewart 1999). The Alewife was intentionally stocked in inland waters. The population in the New River, West Virginia, resulted from stockings in Claytor Lake, New River, Virginia (Jenkins and Burkhead 1994). The recently discovered population in Lake St. Catherine, Vermont, is likely a result of an illegal stocking (Good, personal communication). Lakes in the Adirondack Mountains and Otsego Lake, New York were illegally stocked with Alewife for forage (Smith 1985; Sinnott, personal communication; D. Warner, personal communication).

Status: Introduced populations have been established in 22 US states, the Canadian province of Ontario, and throughout the Great Lakes. Introduction to the Youghiogheny River was unsuccessful (Hendricks et al. 1979).

Great Lakes:
Widespread, with populations reproducing and overwintering at self-sustaining levels in all five Great Lakes. However, current populations in the lower Great Lakes have severely declined from peak abundances throughout the 1900s due to salmonid stockings, dreissenid mussel invasion, and food web shifts (O’Gorman et al. 2013).

Great Lakes Impacts: Alosa pseudoharengus has a high environmental impact in the Great Lakes.

Alewife populations grew rapidly in the 1950s and 1960s in Lake Huron, Lake Ontario, and Lake Michigan, until they largely dominated fish communities as populations of top predators declined due to increased commercial fishing pressure (Bogue 2000) and Sea Lamprey (Petromyzon marinus) predation (Grady 2007). Bottom trawls of Lake Michigan revealed that Alewife abundance increased from <500 lbs/hour of trawling in 1963 to as high as 1500 lbs/hour of trawling in 1966 (Brown 1968). As the abundance of Alewife continued to increase in the absence of predators, massive annual die-offs of Alewife began in Lake Ontario, Lake Huron, and Lake Michigan. Beaches and nearshore regions were littered with “huge windrows” of fish (Brown 1968), reportedly removed by bulldozer (Alewife explosion 1967). After the introduction of salmonids in the late 1960s to both control Alewife abundance and create a sport-fishing industry, Alewife populations have decreased steadily over time, with intermittent periods of growth and decline which could have been due to predation pressure, climate, or limited zooplankton availability (Eck and Wells 1987; Rand et al. 1995; Mills et al. 2005; Madenjian et al. 2008).

It was estimated that Alewife populations were responsible for 28% of the total consumption (by wet weight) in Lake Michigan in 1987, and 96% of the total predation on invertebrates in Lake Ontario in 1990 (Rand et al. 1995). The abundance of Alewife combined with a diet preference of zooplankton and larval fishes has been shown to affect both the zooplankton community and certain native fish populations over time (Crowder 1980). Preference for macrozooplankton and microcrustaceans has shifted the zooplankton community structure towards a prevalence of small species. Following an Alewife decline in Lake Michigan in the mid 1970s, Evans (1990) noted a significant increase in abundance of Limnocalanus macrurus and Diaptomus sicilis, two of the largest copepods. Similarly, a 1987-1995 study of Lake Ontario found that abundances of cyclopoids and other larger species of zooplankton increased during this period of Alewife decline (Johannsson et al. 1998). Changes in zooplankton abundance and structure caused by Alewife can lead to changes in the phytoplankton community (Shapiro et al. 1975).

Disappearance of native planktivorous salmonids, such as Lake Whitefish (Coregonus clupeaformis), in the Great Lakes has been attributed in part to the introduction of Alewife because of reduced zooplankton populations (Crowder and Binkowski 1983; Todd 1986; Page and Laird 1993). Crowder (1984) speculated that a Cisco native to Lake Michigan, the Bloater (C. hoyi) evolved fewer and shorter gill rakers, and shifted to benthic habitat and diet as a result of competition with Alewife. Smith (1970) attributed the extermination of the Cisco and decline of chub species in the Great Lakes to the Alewife. Smith (1970) also discussed the various interrelated changes that took place in each of the Great Lakes as Alewife abundance increased. Christie (1972), on the other hand, argued that the Alewife was not responsible for these changes.

In a review of the adverse effects of Alewife on Great Lake fish communities, Madenjian et al. (2008) presented evidence that agreed with Eck and Wells (1987), who stated that Alewife likely has a larger effect on native fish populations through predation of larvae than competition for food resources. Using time-series data for various fish populations along with change point regression analysis, they concluded that predation of larvae by Alewife likely contributed to the decline of Yellow Perch (Perca flavescens), Deepwater Sculpin (Myoxocephalus thompsonii), Burbot (Lota lota), Atlantic Salmon (Salmo salar), Lake Trout (Salvelinus namaycush), and Emerald Shiner (Notropis atherinoides) (Madenjian et al. 2008).

Furthermore, Alewife has an elevated level of thiaminase, an enzyme that can degrade thiamine in those species that prey on Alewife (Tillitt et al. 2005). Alewife has thus been shown to cause thiamine deficiency and, consequently, early mortality syndrome (EMS) in populations of Alewife predators. EMS and its adverse effects on recruitment and fish populations is well-documented for Coho Salmon (Oncorhynchus kisutch), Lake Trout, and Atlantic Salmon (in which it is also referred to as Cayuga syndrome), among other fishes (Fitzsimons et al. 1999; Ketola et al. 2000; Madenjian et al. 2008; Ladago et al. 2020). In a spawning reef in Lake Ontario, 50 to 75% of newly hatched Lake Trout fry were estimated to suffer from EMS from 1992–1999 (Mills et al. 2005).

In high abundances, Alewife could restructure a lake's food web, leaving less food for native species (USEPA 2008). For example, Alewife and Rainbow Smelt predation in Lake Champlain may prevent Mysis diluviana (formerly Mysis relicta) from recovering from pre-1995 (zebra mussel invasion) densities (Ball et al. 2015). In inland lakes, young-of-year Largemouth Bass (Micropterus salmoides) grow slower and have lower trophic position due to the strong effects Alewife has on the zooplankton community (Boel et al. 2018). EMS also has the potential to cause a genetic bottleneck in populations of heavy Alewife predators by increasing fry mortality and inhibiting recruitment (Mills et al. 2005).

Alosa pseudoharengus has a high socio-economic impact in the Great Lakes.

Alewife is a very important species in the history of biological invasions in the Great Lakes. Periodic large-scale die-offs littered the beaches of the Great Lakes with rotting fish in the 1960s. These mortality events happened with such frequency that they became known as “the annual spring and summer die-off” (Brown 1968). Such die-offs cause widespread beach closures and can pose both a nuisance and a health hazard (Becker 1983).

Alosa pseudoharengus has a high beneficial effect in the Great Lakes.

Prompted by calls for Alewife management, Pacific salmonids were introduced to both control Alewife populations and utilize Alewife as a food source for sport fisheries. Non-native salmonids in the Great Lakes now support a multimillion dollar sport fishing economy and have caused Alewife populations to decline to the extent that salmonid stocking has been reduced to bolster Alewife abundance and sustain the sport fisheries (Dettmers et al. 2012). Chinook (Oncorhynchus tshawytscha), Coho (Oncorhynchus kisutch), and Atlantic Salmon (Salmo salar) all rely on Alewife as forage in Lake Ontario (Mumby et al. 2018). In late summer 2016, Alewife dominated Lake Trout (Salvelinus namaycush) diets in northeastern Lake Michigan (Luo et al. 2019).

Indeed, the threat from Alewives now in some cases is their declining abundance. Since there are large predators (e.g., Chinook salmon) that focus on Alewife as prey, these fisheries are heavily reliant on Alewife as a food source, and managers are attempting to balance predator-prey ratios to sustain these populations/fisheries (personal communication, Jesse Lepak, May 21, 2021).

In the Great Lakes, Alewife consumes the invasive cladocerans Bythotrephes and Cercopagis (Keilty 1990; Mills et al. 1992; Bushnoe et al. 2003), with the highest consumption rates nearshore (Keeler et al. 2015).  Alewife also heavily preys upon the invasive bloody red shrimp (Hemimysis anomala) (Boscarino et al. 2020).

Alewife helps sustain populations of the native double crested cormorant (Phalacrocorax auritus), however, at higher than historic abundances cormorants can have significant negative impacts on sport and commercial fisheries, other waterfowl (Madura and Jones 2016), and island plant communities (Boutin et al. 2011).

Management: Regulations (pertaining to the Great Lakes region)
Alosa pseudoharengus is a regulated invasive species in Minnesota (MN Administrative Rules, 6216.0260 Regulated).  New York restricts the use of Alewife as bait in most waters (6 NYCRR Part 19).  While not listed by name, in Ohio it is illegal for any person to possess, import or sell exotic species of fish (including Alosa pseudoharengus) or hybrids thereof for introduction or to release into any body of water that is connected to or otherwise drains into a flowing stream or other body of water that would allow egress of the fish into public waters, or waters of the state, without first having obtained permission (OAC Chapter 1501:31-19). Restricted in Wisconsin under Wis. Admin Code § NR 40.05, making the transport, transfer, introduction, or possession of Alewife illegal without a permit. The import, possession, transport, and release of live Alewife in Manitoba is prohibited under articles 6 to 10 of the Canadian Fisheries Act SOR/2015-121.

Note: Check federal, state/provincial, and local regulations for the most up-to-date information.

The management response to Great Lakes alewife overabundance and recurring die-offs was to invest in sea lamprey (Petromyzon marinus) control and planting of hatchery-reared Pacific salmonids (Oncorhynchus spp.) to re-establish top open-water predators (Kocik and Jones 1999; Hansen and Holey 2002).  Older and larger fish tend to be most heavily affected by piscivores, while smaller and younger fish remain abundant (Hewett and Stewart 1989).  Alewives are now managed in part to support the valuable salmonid fishery.

There are no known physical control methods for this species.

Of the four chemical piscicides registered for use in the United States, antimycin A and rotenone are considered “general” piscicides (GLMRIS 2012).

Increasing CO2 concentrations, either by bubbling pressurized gas directly into water or by the addition of sodium bicarbonate (NaHCO3) has been used to sedate fish with minimal residual toxicity, and is a potential method of harvesting fish for removal, though maintaining adequate CO2 concentrations may be difficult in large/natural water bodies (Clearwater et al. 2008). CO2 is approved only for use as an anesthetic for cold, cool, and warm water fishes the US, not for use as euthanasia, and exposure to NaHCO3 concentration of 142–642 mg/L for 5 min. is sufficient to anaesthetize most fish (Clearwater et al. 2008).

It should be noted that chemical treatment will often lead to non-target kills, and so all options for management of a species should be adequately studied before a decision is made to use piscicides or other chemicals. Potential effects on non-target plants and organisms, including macroinvertebrates and other fishes, should always be deliberately evaluated and analyzed. The effects of combinations of management chemicals and other toxicants, whether intentional or unintentional, should be understood prior to chemical treatment.  Other non-selective alterations of water quality, such as reducing dissolved oxygen levels or altering pH, could also have a deleterious impact on native fish, invertebrates, and other fauna or flora, and their potential harmful effects should therefore be evaluated thoroughly.

Note: Check state/provincial and local regulations for the most up-to-date information regarding permits for control methods. Follow all label instructions.

Remarks: Although there is a report of two small alewives taken from the Colorado River, Texas (Bean 1882), we believe this record is in error. Bean (1882) reported that the specimens were sent to Professor Baird at the National Museum. However, a query of the museum's holdings did not return these specimens. We believe the fish are more likely either misidentified A. chrysochloris or A. sapidissima. Alosa sapidissima were stocked in the Colorado River in 1874 (Bean 1882).

Alewife is one of the most frequently found prey items in the diet of the Double-Crested Cormorant in the southern basin of Lake Michigan (Madura and Jones 2016).

Voucher specimens: Michigan (UMMZ 157215, 160969, 167872, 171308, 170945), Wisconsin (UMMZ 162861, 167945).

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Author: Fuller, P., E. Maynard, D. Raikow, J. Larson, A. Fusaro, M. Neilson, and A. Bartos

Contributing Agencies:

Revision Date: 8/4/2021

Peer Review Date: 8/4/2021

Citation for this information:
Fuller, P., E. Maynard, D. Raikow, J. Larson, A. Fusaro, M. Neilson, and A. Bartos, 2022, Alosa pseudoharengus (Wilson, 1811): U.S. Geological Survey, Nonindigenous Aquatic Species Database, Gainesville, FL, and NOAA Great Lakes Aquatic Nonindigenous Species Information System, Ann Arbor, MI,, Revision Date: 8/4/2021, Peer Review Date: 8/4/2021, Access Date: 5/17/2022

This information is preliminary or provisional and is subject to revision. It is being provided to meet the need for timely best science. The information has not received final approval by the U.S. Geological Survey (USGS) and is provided on the condition that neither the USGS nor the U.S. Government shall be held liable for any damages resulting from the authorized or unauthorized use of the information.