Alosa pseudoharengus
(alewife)

Pictures

Picture	Title	Caption	Copyright
Title Artwork of adult fish Caption Alosa pseudoharengus (alewife); artwork of adult fish. Copyright ©Duane Raver/US Fish and Wildlife Service/Bugwood.org - CC BY-NC 3.0 US	Artwork of adult fish	Alosa pseudoharengus (alewife); artwork of adult fish.	©Duane Raver/US Fish and Wildlife Service/Bugwood.org - CC BY-NC 3.0 US
Title Alewife Caption Alosa pseudoharengus (alewife); speciamen from the landlocked population in Lake Ontario, Canada. Copyright ©Theodore Strang/USGS, Oswego, NY, USA.	Alewife	Alosa pseudoharengus (alewife); speciamen from the landlocked population in Lake Ontario, Canada.	©Theodore Strang/USGS, Oswego, NY, USA.

Identity

Preferred Scientific Name

• Alosa pseudoharengus Wilson, 1811

Preferred Common Name

• alewife

International Common Names

• French: gaspareau

Local Common Names

• : bang racer; big-eyed herring; blear-eyed herring; branch herring; ellwife; freshwater herring; glut herring; golden shad; gray herring; grayback; green shad; kiack; kyak; mulhaden; sawbelly; seth; skipjack; spring herring; wall-eyed herring; white herring

Summary of Invasiveness

A. pseudoharengus is native to the western Atlantic Ocean off the coast of North America, where it occurred historically from Labrador, Nova Scotia, and northeastern Newfoundland, Canada south to South Carolina, USA (ASMFC, 2009). It is anadromous, ascending rivers and streams to spawn, and the young naturally occur in many coastal lakes and streams (Scott and Crossman, 1973). Although A. pseudoharengus is a valued fish in its native range, where there are numerous projects aimed at restoring anadromous populations (Maine DMR, 2009; USFWS, 2009), when introduced into lakes and reservoirs, they can become invasive. They can complete their life cycle in freshwater, they are fecund (Norden, 1967; Bronte et al., 1991), they can spawn successfully in a wide variety of habitats (ASMFC, 2009), and, they are relatively long lived with a maximum age of 11 having been reported from the Laurentian Great Lakes of North American (O’Gorman et al., 1987, 1997; Madenjian et al., 2003). In land-locked waterbodies A. pseudoharengus can become overabundant, altering the zooplankton community by size-selective predation (Brooks and Dodson, 1965; Wells, 1970; Warshaw, 1972), affecting native fishes by preying on their young (O’Gorman and Stewart, 1999; Madenjian et al., 2008; O’Gorman et al., in press) and, sometimes, causing a thiamine deficiency among the fishes that eat them (Brown et al., 2005). Dense populations are subject to periodic die-offs and the dead fish washing ashore create odour and sanitation problems (Greenwood, 1970).

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Chordata
•             Subphylum: Vertebrata
•                 Class: Actinopterygii
•                     Order: Clupeiformes
•                         Family: Clupeidae
•                             Genus: Alosa
•                                 Species: Alosa pseudoharengus

Notes on Taxonomy and Nomenclature

The alewife, Alosa pseudoharengus, is one of 216 fishes in the family Clupeidae, a well distributed taxonomic group, mainly tropical and marine, that includes herrings, sardines, menhadens, pilchards, sprats, and shads (UMMZ, 2009). The genus Alosa (river herrings), subfamily Alosinae (shads), has 24 species. Alewives and the blueback herring, Alosa aestivalis, are collectively referred to as river herring in North America perhaps because not only are both species similar in appearance but also because both move into rivers and streams to spawn at about the same time.

Description

A. pseudoharengus has an overall silvery colour with a greyish-green back. A black spot at the eye level is directly behind the head. Adults have longitudinal lines that run along the scale lines above the midline of the body. The large scales are deciduous and the lateral line is not well-developed (Scott and Crossman, 1973). Scales on the midline of the belly form scutes, creating a serrated surface (Trautman, 1957). The body is strongly laterally compressed and relatively deep. Eyes are large. The front of the lower jaw is thick and extends past the upper jaw when the mouth is closed. The maxillary extends to below the middle of the eye. A few small teeth are present on the premaxillary and mandible (Scott and Crossman, 1973). There are more than 30 gill rakers on the lower angle of the first gill arch (Trautman, 1957). The single dorsal fin usually has 13-14 rays but may have 12-16. The caudal fin is forked. The anal fin is short and wide with 15-19 rays (usually 17-18). The pelvic fins are rather small and contain 10 rays. The pectoral fins are low on the sides and they usually have 16 rays but may have as few as 14 (Scott and Crossman, 1973). A. pseudoharengus in landlocked populations become stunted; their maximum total length rarely exceeds 200 mm and average total length ranges from 125 to 175 mm (O’Gorman et al., 1987; Madenjian et al., 2003). Average length usually increases with the size of the waterbody. In contrast, anadromous A. pseudoharengus can grow to 360 to 380 mm (Collette and Klein-MacPhee, 2002).

Distribution

Established populations of A. pseudoharengus in North American waterbodies are mainly in the United States and in the easternmost part of the continent. All but a few are located east of the Mississippi River (USGS, 2009). The lone exceptions are in the north part of the state of Nebraska. Although there are records of alewife in the western state of Colorado (Minckley, 1973; USGS, 2009), the fish was not recently listed as being present in that state (Johnson and Nomanbhoy, 2005).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

The largest geographical spread of alewife occurred in the mid-twentieth century when the fish moved into Lakes Erie, Huron, Michigan, and Superior, four of the five Laurentian Great Lakes of North America (Miller, 1957). A. pseudoharengus had been present in Lake Ontario, the fifth and easternmost of the interconnected Great Lakes, since the mid-1800s and had been abundant there since the late 1870s (O’Gorman and Stewart, 1999).  Although Lake Ontario drains to the Atlantic Ocean through the 500-km long St. Lawrence River, it is generally believed that A. pseudoharengus is not native to Lake Ontario, mainly because of its absence from the historical record until hundreds of years after the arrival of European colonists. Moreover, there is also genetic evidence that suggests alewife were not indigenous to Lake Ontario (Ihssen et al., 1992). There is no agreement on how A. pseudoharengus may have invaded Lake Ontario (Daniels, 2001); some have suggested migration through navigation canals or perhaps inadvertent introduction from planned releases of other fishes (Miller, 1957; Smith, 1970). A. pseudoharengus in Lake Ontario were prevented from moving westward to the other four Great Lakes by Niagara Falls. In the 1920s, improvements to the Welland Canal, a navigation canal which connects Lake Ontario to Lake Erie, apparently allowed A. pseudoharengus to use the canal to bypass Niagara Falls (O’Gorman and Stewart, 1999). A. pseudoharengus were first reported from Lake Erie in 1931, from Lake Huron in 1933, from Lake Michigan in 1949, and from Lake Superior in 1954 (Miller, 1957).

The spread of A. pseudoharengus through the four Great Lakes in the mid-1900s was facilitated by favourable climatic conditions and the collapse of native piscivorous fishes (O’Gorman and Stewart, 1999). In Lake Superior, the northernmost Great Lake, A. pseudoharengus never became abundant, presumably because the cold temperature regime was unfavourable for successful reproduction (Bronte et al., 1991; O’Gorman et al., 1997). In Lake Erie, the shallowest Great Lake, A. pseudoharengus became abundant, but only in some years; this was most likely due to cold water temperatures in most winters limiting survival (Smith, 1968; Colby, 1973; Ryan et al., 2003). In Lakes Michigan and Huron, however, where temperature regimes were suitable for successful reproduction and where there were relatively warm, deep-water winter refugia, A. pseudoharengus thrived and by the mid-1960s their numbers reached nuisance levels (O’Gorman and Stewart, 1999; O’Gorman et al., in press). Massive die-offs in Lake Michigan left tons of rotting fishes on beaches and caused a public outcry (Greenwood, 1970). Initially, two techniques were employed to reduce alewife numbers, commercial harvest and large-scale stocking of hatchery-reared, Pacific salmon Onchorynchus spp., lake trout Salvelinus namaycush, and brown trout Salmo trutta. Commercial harvest of the low-value alewife was not economically viable whereas stocking of salmon and trout by government agencies created a high-value recreational fishery (Bence and Smith, 1999). Moreover, the hatchery-reared salmonids, particularly Chinook salmon Oncorhynchus tshawytscha, successfully reduced alewife numbers in Lakes Michigan, Huron, and Ontario (O’Gorman et al., in press). Indeed, in Lake Huron, a surge in natural reproduction by Chinook salmon was largely responsible for the collapse of the A. pseudoharengus population (O’Gorman et al., in press). In Lake Erie, where there was limited habitat for salmonids, resurgence of native walleye Sander vitreus contributed to holding the A. pseudoharengus population in check (Knight and Vondracek, 1993; O’Gorman et al., in press).

Risk of Introduction

Introduction of A. pseudoharengus is unlikely without a concerted effort unless the receiving waterbody is in a watershed where A. pseudoharengus are already established. A. pseudoharengus can move within a watershed through rivers and canals, even through canals with navigation locks. There is a large body of literature on the adverse affects of introducing alewife to freshwater ecosystems (O’Gorman and Stewart, 1999; Madenjian et al., 2008; O’Gorman et al., in press) so natural resource management agencies are vigilant against further introductions. The greatest risk for introduction appears to be from illicit releases by individuals seeking to “improve” fishing, followed by accidental introduction by release of A. pseudoharengus, illegally used for live bait. A. pseudoharengus are somewhat fragile and are therefore not used in the aquarium trade or transported live for human consumption.

Habitat

A. pseudoharengus is a pelagic fish that occupies open waters. In the northwestern Atlantic Ocean, A. pseudoharengus favour a depth range of 56 to 110 m (Neves, 1981). In the Laurentian Great Lakes, the depths occupied by A. pseudoharengus change seasonally and vary somewhat among lakes due to differences in bathymetry and temperature regimes (Wells, 1968; O’Gorman and Schneider, 1986; O’Gorman et al., 2000). In general, A. pseudoharengus overwinter offshore in deep water and move shoreward into shallower water in spring to spawn, after which they move back to open waters where they remain throughout the summer, occupying mid to upper levels in the water column (Bergstedt and O’Gorman, 1989; O’Gorman et al., in press). In autumn, as the lakes cool, A. pseudoharengus descend to greater depths. The depth distribution of A. pseudoharengus within a lake can change, however, as it did in Lake Ontario following changes in the lake ecosystem (O’Gorman et al., 2000).

Habitat List

Biology and Ecology

Reproductive Biology

A. pseudoharengus spawn in spring, at night, near the surface, throughout nearshore areas including bays, harbours, and in tributaries (Scott and Crossman, 1973). The minimum temperature at which Atlantic A. pseudoharengus spawn is 10ºC (Cianci, 1965) and spawning peaks at 13-16ºC (Richkus, 1974; Tyus, 1974). In landlocked populations, the spawning period is protracted, lasting more than a month (Odell, 1934; Norden, 1967; Hlavek and Norden, 1978). Mean fecundity of Lake Michigan A. pseudoharengus was 11,150 for fish averaging 160 mm total length, 16,140 for fish averaging 176 mm, and 22,400 for fish averaging 192 mm (Norden, 1967; Hlavek and Norden, 1978). The non-adhesive eggs are demersal and are broadcast at random over any type of bottom (Odell, 1934; Mansueti, 1956). Optimum temperature for egg incubation is 17.8ºC and incubation time varies from 15 days at 7.2ºC to 3.7 days at 21.1ºC (Edsall, 1970). Alewife larvae average 3.8 mm at hatching, 5.1 mm at yolk sac absorbtion, and, when held at 20ºC, they begin feeding two days after hatching (Norden, 1967; Heinrich, 1981). The larvae are positively phototropic and pelagic (Odell, 1934).

Nutrition

Land-locked A. pseudoharengus eat zooplankton throughout their life (Morsell and Norden, 1968; Mills et al., 1992, 1995; Stewart et al., 2009). They are size-selective feeders, preferentially eating the largest available zooplankters (Kohler, 1980). Indeed, the movements of A. pseudoharengus around a large lake or the magnitude of an alewife population can sometimes be tracked by the size and species composition of zooplankton (Wells, 1970; Warshaw, 1972; O’Gorman et al., 1991). The first food of larval alewife is cyclopoid copepodites and as the larvae grow they incorporate larger zooplankton in their diet (Heinrich, 1981). When the fish grow to about 110-119 mm they begin to feed on larger invertebrates (Morsell and Norden, 1968) - amphipods, insects, and, in the Laurentian Great Lakes, the opossum shrimp Mysis diluviana and the bloody-red shrimp Hemimysis anomola (Walsh et al., 2008; Stewart et al., 2009; Lantry et al., 2010). A. pseudoharengus are adept at capturing prey in mid-water but have difficulty capturing prey located on or near the lake bottom (Janssen, 1978a). They can feed in the dark (Janssen, 1978b; Janssen et al., 1995; Kelso and Ney, 1983) and they have three modes of feeding: particulate, filtering, and gulping (Janssen, 1976). Nocturnal movement of A. pseudoharengus to nearshore areas of lakes to feed has been recorded (Kelso and Ney, 1983). Conversely, A. pseudoharengus have been found feeding at night far offshore in the Great Lakes on vertically migrating Mysis (Boscarino et al., 2009). A. pseudoharengus also eat their own larvae (Odell, 1934; Rhodes et al., 1974) as well as the larvae of other fishes and small young-of-year fishes (Kohler and Ney, 1980; Brandt et al., 1987; Kreuger et al., 1995).

Environmental Requirements

A. pseudoharengus in Lake Michigan begin spawning when water temperatures reach about 15.6ºC; spawning is interrupted when water temperatures exceed 27.8ºC (Edsall, 1970). Eggs hatch when incubated at 6.9 to 29.4ºC but not at lower or higher temperatures. About 69% of the larvae from eggs incubated at less than 10.6ºC are deformed and not likely to survive. Alewife young-of-year (YOY) have been found at 16 to 29ºC in Lake Michigan (Brandt, 1980; Dufour et al., 2008). Laboratory studies show a YOY temperature preference range of 21 to 31ºC (Otto et al., 1976; Spotila et al., 1979) and critical thermal maximum range of 32 to 34ºC for YOY acclimated to 20-25ºC (Otto et al., 1976). In the Laurentian Great Lakes, A. pseudoharengus must grow to a total length of 60 mm or longer if they are to survive the winter (O’Gorman and Stewart, 1999). A. pseudoharengus are rare in Lake Superior, the northern-most Great Lake, presumably because the short growing season does not allow most young A. pseudoharengus sufficient time to grow to a size that would allow successful overwintering (O’Gorman et al., 1997; O’Gorman et al., in press). In summer, adult A. pseudoharengus in the Great Lakes avoid the cold hypolimnetic waters and occupy the warm epilimnetic waters although under certain conditions they some can be found in the thermocline (Wells, 1968; Olson et al., 1988; Johannsson and O’Gorman, 1991; but see Janssen and Brandt, 1980). Laboratory temperature preferences for adult A. pseudoharengus were 21ºC in spring, 16-19ºC in summer, and 16 -20ºC in autumn (Otto et al., 1976; Spotila et al., 1979). Ultimate upper lethal temperatures were 31-34ºC for adult alewives acclimated to 20-27ºC (Otto et al., 1976; McCauley and Binkowski, 1982).

A. pseudoharengus are severely stressed by temperatures lower than 3ºC (Colby, 1973). In the Great Lakes A. pseudoharengus are in water at 1 to 3ºC during most winters. In severe winters, as water temperature may be at or below 1ºC (Mortimer, 1971; Rodgers, 1987), mass mortalities of A. pseudoharengus sometimes occur (O’Gorman and Schneider, 1986; Bergstedt and O’Gorman, 1989; O’Gorman et al., in press). Low-temperature mortalities appear to be due to osmoregulatory failure caused by a loss of homeoviscous adaptation (altering the lipid composition of biomembranes to compensate for temperature-induced changes in membrane fluidity; Stanley and Colby, 1971; Snyder and Hennessey, 2003). Homeoviscous adaptation can be influenced by diet (Synder and Murray, 2009) and inter-annual variation in diet may be the reason for discrepancies in the severity of die-offs among years with similar winter weather (O’Gorman et al., in press). When held in ponds where water temperatures were < 2ºC for more than six weeks, A. pseudoharengus experienced sublethal immunosuppression, increasing their susceptibility to disease (Lepak and Kraft, 2008).

Climate

Latitude/Altitude Ranges

Water Tolerances

Pathway Causes

Impact Summary

Economic Impact

Establishment of alewife populations in land-locked waterbodies has had both positive and negative economic impacts. The negative impact most noticeable to the general public has been the periodic die-offs of large numbers of fish. As early as 1892, residents of Burlington, Canada, on Lake Ontario, were complaining of the costs of removing the smelly, dead A. pseudoharengus that washed ashore in summer (Pritchard, 1929). In 1967, a massive die-off of A. pseudoharengus in Lake Michigan (the fifth largest lake in the world) resulted in an estimated loss in excess of 100 million dollars to industry, municipalities, and businesses dependent on recreation (Greenwood, 1970). Other less visible but no less important negative economic impacts include harmful effects on other fishes important to recreational and commercial fisheries.

The largest positive economic impact was the development of multi-million dollar recreational fisheries for hatchery-raised salmon and trout in the Laurentian Great Lakes (Bence and Smith, 1999; Connelly and Brown, 2009) -- A. pseudoharengusprovides an important food source for these fish. These new fisheries are hatchery-dependent, however, and thus require large capital investments to start and continuing investments of capital to maintain.

Environmental Impact

When A. pseudoharengus become established in a waterbody they alter the size and species composition of the zooplankton community by size-selective predation (Brooks and Dodson, 1965; Wells, 1970; Warshaw, 1972; Johannsson, 2003). Kelso and Ney (1983) noted that the high foraging efficiency of A. pseudoharengus on zooplankton may result in trophic competition with juvenile fishes in the littoral zone of a reservoir. Depression of zooplankton can reduce grazing of phytoplankton resulting in decreased water clarity.

Impact on Biodiversity

Alewife populations in landlocked waterbodies negatively impact biodiversity in three ways. First by size-selective predation on zooplankton they can eliminate the largest species and shift dominance to the smallest species (Brooks and Dodson, 1965; Wells, 1970; Warshaw, 1972; Johannsson, 2003). Second, by preying on pelagic larvae of native fishes with no evolutionary exposure to A. pseudoharengus, they can sharply curtail recruitment, depressing populations of native fishes (Madenjian et al., 2008; O’Gorman et al., in press). Third, by causing a thiamine deficiency among some salmonines that eat mainly A. pseudoharengus, they can lower reproductive success of ecologically and economically important top predators. A. pseudoharengus contain high concentrations of thiaminase, an enzyme that breaks down thiamine (Gnaedinger, 1964). Female salmonines that eat mostly A. pseudoharengus can produce thiamine-deficient eggs (Marcquenski, 1996; Brown et al., 1998, 2005) and the young that emerge from those eggs develop Early Mortality Syndrome (EMS), a syndrome characterized by abnormal behaviour, physical abnormalities, and death (McDonald et al., 1998). EMS is thought to be one of the major impediments to restoring the native lake trout Salvelinus namaycush to many areas of the Laurentian Great Lakes (Madenjian et al., 2008). The final elimination of the native, land-locked Atlantic salmon Salmo salar from Lake Ontario in the late 1800s, soon after alewife proliferated, may well have been due to thiaminase-induced thiamine deficiency (Ketola et al., 2000; Madenjian et al., 2008).

Risk and Impact Factors

• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Highly adaptable to different environments
• Is a habitat generalist
• Capable of securing and ingesting a wide range of food
• Highly mobile locally
• Long lived
• Fast growing
• Has high reproductive potential
• Gregarious

• Damaged ecosystem services
• Ecosystem change/ habitat alteration
• Negatively impacts cultural/traditional practices
• Negatively impacts animal health
• Negatively impacts aquaculture/fisheries
• Reduced native biodiversity
• Threat to/ loss of native species

• Competition (unspecified)
• Herbivory/grazing/browsing
• Predation

• Difficult/costly to control

Uses List

Animal feed, fodder, forage

• Bait/attractant
• Forage

Human food and beverage

• Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)

Similarities to Other Species/Conditions

A. pseudoharengus is similar in appearance to the blueback herring Alosa aestivalis. Indeed, due to their similarity, the two species are often harvested and managed together as “river herring” in the USA or as “gaspereau” in Canada (Mullen et al., 1986; NOAA, 2009). The eye of A. pseudoharengus is large with a diameter greater than the length of the snout whereas the eye of the blueback herring is small with a diameter equal to or less than the snout length (Mullen et al., 1986). Internally, the peritoneum of the alewife is pearly to white, sometimes with grey flecks, whereas that of the blueback herring is sooty to black (Scott and Crossman, 1973; Owens et al., 1998). Landlocked populations of A. pseudoharengus are common and occur across a broad geographic range whereas landlocked populations of blueback herring are uncommon and are confined to the southern USA (Owens et al., 1998).

Prevention and Control

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Once established in a waterbody, an alewife population cannot be eradicated although it can be reduced by increasing the number of piscivorous fishes (O’Gorman et al., in press). Invasive populations of alewife were established by movement of alewife through navigation canals (O’Gorman and Stewart, 1999) and by introduction, either accidental or intentional. Stocking of alewife was once done to augment prey fish populations but once the deleterious effects of establishing a population became known, it was no longer sanctioned. Illicit, intentional introductions are now the primary means of movement and they are most likely done by anglers who view A. pseudoharengus favourably because of the well-advertised, large size of piscivorous fishes in some lakes where A. pseudoharengus are abundant (Marsden and Hauser, 2009). Populations may also become established by anglers releasing the fish they are using for bait, some of which may be A. pseudoharengus. The state of New York, as well as other states, prohibit or greatly limit the use of A. pseudoharengus for live bait in inland waters.

References

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CTDEP, 2007. Connecticut Department of Environmental Protection. Prohibition on the taking of alewife and blueback herring from Connecticut waters extended for another year. Connecticut Department of Environmental Protection. Prohibition on the taking of alewife and blueback herring from Connecticut waters extended for another year. http://www.ct.gov/dep/cwp/view.asp?Q=335656&A=2794

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Links to Websites

Contributors

13/04/10 Original text by:

Robert O'Gorman, Lake Ontario Biological Station, USGS7 Lake Street, Oswego, NY 13126, USA

Distribution Maps