Invasion History

First Non-native North American Tidal Record: 1897
First Non-native West Coast Tidal Record: 1930
First Non-native East/Gulf Coast Tidal Record: 1897

General Invasion History:

The Warmouth (Lepomis gulosus) is native to the Great Lakes and Mississippi Basins, from Pennsylvania to Minnesota, south to the Gulf, from Florida to Texas, and Atlantic drainages south of the James River (Jenkins and Burkhead 1993; Lee et al. 1980; Page and Burr 1991). The original range on the Atlantic Coastal Plain is uncertain because of extensive introductions and uncertain early records. It was considered likely native to James and Dismal Swamp, based on Cope's (1869) record, but is probably introduced northward, with some uncertainty due to the lack of early records and spotty present distribution, but is clearly introduced in the Potomac, Susquehanna, Delaware, and Hudson rivers (Jenkins and Burkhead 1993; Horwitz 1986; Daniels et al. 2005). Warmouth were transported and stocked by the US Fisheries Commission and state agencies. On the West coast, it was introduced to the Sacramento-San Joaquin Delta and the Columbia River estuary (Lampman 1946; Dill and Cordone 1997; Moyle 2002). Currently, it has been introduced to non-native waters in 29 states (USGS Nonindigenous Aquatic Species Program 2018). It has also been introduced to Puerto Rico and Mexico (Lever 1996).

North American Invasion History:

Invasion History on the West Coast:

The early history of stocking of the Warmouth on the West Coast is obscure. Early plantings were made in Lake Cuyamaca, San Diego County (400 fish) and the Feather River (100 fish), California, in 1891 and 1895, but details in different reports are contradictory, and it is not clear that any of these fish survived (Smith 1895; Shebley 1917; Dill and Cordone 1997). However, Warmouth were reported to occur in the Central Valley in 1931 and were included in a key for Delta fishes in 1941 (Dill and Cordone 1997). In 1966, Warmouth comprised about 2% of centrarchid fishes collected in the Delta (Turner 1966, cited by Cohen and Carlton 1995). In the freshwater Delta in the 1990s to 2000s, Warmouth were were <1 to 2.2% of the total catch (Feyrer and Healy 2003; Brown and Michniuk 2007; Grimaldo et al. 2012). Single specimens were caught at several stations in the fresh-brackish Suisun Marshes (Matern et al. 2002). Warmouth occur mostly in sloughs of the Delta (Wang 1986).

In the Columbia River basin, there was a stocking of 201 Warmouth in 1892, a release of mixed sunfishes from Illinois in 1893 in the Willamette River at Salem, and a release of mixed Mississippi 'spiny-rayed fishes' displayed at the Lewis and Clark Exposition in Portland in 1905 (Lampman 1946). It is not clear if any of these releases was responsible for the introduction of the Warmouth to the Columbia River system. However, Warmouth were caught in the Columbia River at Kalama in 1930 and in the Willamette River and Columbia Slough near Portland in small numbers (Farr and Ward 1992; Van Dyke et al. 2009).

Invasion History on the East Coast:

Warmouth were collected in the James River near Richmond, Virginia in 1867 (Cope 1869). This fish was probably native in this river, but is introduced in Chesapeake tributaries further north (Jenkins and Burkhead 1993). Warmouth were stocked in Shenandoah River, 1893 (Worth 1895), and was first caught in the Washington DC area in Little River, a Potomac tributary, in 1897. By 1898 it was 'apparently becoming common in a few places' (Smith and Bean 1898). It was collected at Chain Bridge, at the head of tide on the Potomac in 1910 (U.S. National Museum of Natural History 2001). Other tributaries were colonized later, probably by informal stocking. The first verified record of Warmouth in the Rappahannock River was in 1938 (Jenkins and Burkhead 1993), but it was not reported from the tidal river by 1951 (Massman et al. 1952), but was later found there by Maurakis et al. (1987). Warmouth was reported as apparently introduced to the Susquehanna River (Fowler 1919), but was not listed for the river by Denoncourt and Cooper (1975) or Denoncourt et al. (1975b), or for the river below Conowingo Dam by McKeown (1984). However, Aa specimen was caught in Broad Creek, a South River tributary near Annapolis in October 2008 (Rob Aguilar, personal communication).

In the Delaware River, Warmouth was reportedly introduced in the Delaware River (Fowler 1948). Specimens were caught at a power plant in Chester, Pennsylvania River, in 1977 (Horwitz 1986), and in Newcastle County, Delaware County, Augustine Creek (Raasch 1997). In the Hudson River estuary, Warmouth were first collected in 1936, and collected again in the 1970s near Annandale, New York (~River km 160), and near Newburgh (~92 River km) (Smith and Lake 1990), in the tidal freshwater region of the estuary. Daniels et al. (2005) listed the species as 'rare but increasing'. Warmouth may have been introduced as a contaminant with a stocking of Green Sunfish (Greeley 1937, cited by Mills et al. 1997).

Invasion History Elsewhere in the World:

Warmouth were introduced to reservoirs in Puerto Rico in 1916, apparently as a contaminant in a stocking of Largemouth Bass (Micropterus salmoides) and Rock Bass (Ambloplites rupestris). The Rock Bass introduction failed, but in 1971, a Warmouth was caught in Carite Reservoir (Lever 1996; USGS Nonindigenous Aquatic Species Program 2018). They also may have been introduced into reservoirs in the Rio Grande basin in northern Mexico, probably south of their native range (Edwards and Contreras 1991; Lever 1996).


The Warmouth (Lepomis gulosus) is a medium-sized freshwater fish. Fishes of the family Centrarchidae (Sunfishes and Black Basses) have a strongly laterally compressed body. They have a spiny and a soft dorsal fin, which are fused. They have 3–8 anal spines, thoracic pelvic fins, and ctenoid scales. The sunfishes of the genus Lepomis have a shallowly forked tail, a smooth edge to the gill cover, and a fleshy 'ear-flap' projecting from the gill-cover. The gill-cover has a stiff rear edge. Warmouth have a compressed, somewhat rectangular 'bass-like' body, a large mouth (with the upper jaw extending beyond the midpoint of the eye), and short, rounded pectoral fins, which do not extend past the eye when bent forward. The dorsal fin has 9–11 spines and 9–11 rays, while the anal fin has 3 spines and 9–10 rays. There are 36–44 lateral line scales. The maximum length is 310 mm, but a more usual length is 100–180 mm. The back and sides are olive-brown, often with a purple sheen and dark-brown mottling, and dark-brown chainlike bars on the sides, with a white-to-yellow belly. The ear-flap is black, with a yellow edge, marked with a small red spot. Dark Brown lines radiate across the head from the bright red eyes (Hardy 1978; Page and Burr 1991; Jenkins and Burkhead 1993; Moyle 2002).


Taxonomic Tree

Kingdom:   Animalia
Phylum:   Chordata
Subphylum:   Vertebrata
Superclass:   Osteichthyes
Class:   Actinopterygii
Subclass:   Neopterygii
Infraclass:   Teleostei
Superorder:   Acanthopterygii
Order:   Perciformes
Suborder:   Percoidei
Family:   Centrarchidae
Genus:   Lepomis
Species:   gulosus


Lepomis gulosus ((Cuvier in Cuvier and Valenciennes, 1829) 2003-12-02, 1829)
Pomotis gulosus (Cuvier in Cuvier and Valenciennes, 1829 2003-12-02, 1829)
Chaenobryttus gulosus (Cuvier in Cuvier and Valenciennes, 1829 2003-12-02, 1829)
Chaenobryttus coronarius (Bartram, 1791)

Potentially Misidentified Species

Lepomis cyanellus
Lepomis cyanellus (Green Sunfish) differs from the Warmouth in having a more rectangular 'bass-like' body, a large mouth, and dark vertical bars running down the body, and has dark spots on the posterior of the anal and dorsal fins (Page and Burr 1991). The Green Sunfish is native to the Great Lakes-Mississippi-Gulf basins (Page and Burr 1991). It is introduced in the San Francisco and the Columbia River estuaries, and on the Atlantic Slope from South Carolina to Connecticut (USGS Nonindigenous Aquatic Species Program 2018). South of the Potomac River, it appears to be rare or absent in Coastal Plain drainages (Jenkins and Burkhead 1993).

Lepomis gibbosus
Lepomis gibbosus (Pumpkinseed) differs from the Warmouth in having a deeper oval 'typical-sunfish' body, a small mouth, a bright red spot on the ear-flap, and long, pointed pectoral fins. The Pumpkinseed has thin, wavy lines on the posterior of the anal and dorsal fins (Page and Burr 1991). The Pumpkinseed is native to the Atlantic Slope from New Brunswick to South Carolina, the Great Lakes basin, and the Mississippi-Gulf basins from Manitoba to Missouri (Page and Burr 1991). It is introduced to the San Francisco and Columbia River estuaries (USGS Nonindigenous Aquatic Species Program 2018).

Lepomis macrochirus
Lepomis macrochirus (Bluegill) differs from the Warmouth in having a deeper oval 'typical-sunfish' body, a small mouth, a bright red spot on the ear-flap, and long, pointed pectoral fins. The Bluegill has a black spot on the rear edge of the dorsal fin (Page and Burr 1991). The Bluegill is native to the Atlantic Slope from North Carolina to Florida, the Great Lakes basin, and the Mississippi-Gulf basins from Quebec and Minnesota to Texas (Page and Burr 1991). It is introduced to the San Francisco and Columbia River estuaries, and widely through the western US, and on the Atlantic Slope, from northern North Carolina to Maine (USGS Nonindigenous Aquatic Species Program 2018).

Lepomis microlophus
Lepomis microlophus (Redear Sunfish) differs from the Warmouth in having a deeper oval 'typical-sunfish' body, a small mouth, longer ear-flap with a bright red spot, and long, pointed pectoral fins (Page and Burr 1991). The Redear Sunfish is native to the Atlantic Slope from Georgia to Florida, and the Mississippi-Gulf basins from Indiana and Illinois and Minnesota to Florida and Texas (Page and Burr 1991). It is introduced to the San Francisco estuary, and on the Atlantic Slope form South Carolina to the Potomac River (USGS Nonindigenous Aquatic Species Program 2018).



The Warmouth (Lepomis gulosus) is a freshwater fish characteristic of lakes, ponds, backwaters, swamps, and slow-moving rivers (Page and Burr 1991; Moyle 2002). Spawning males are bright yellow, with iridescent blue streaks on the gill cover (Hardy 1978). The fish mature at ages of 1–2 years, at 75–100 mm (Hardy 1978; Jenkins and Burkhead 1993). Spawning takes place at ~21 °C (Hardy 1978). Males excavate and guard a nesting site, in sand, rubble, or detritus or leaf mold, often near tree roots, stumps, sticks, etc. Males are territorial and aggressive when defending nesting areas. Nesting males tend to be solitary. Females can carry 4500–63,200 eggs (Hardy 1978; Jenkins and Burkhead 1994). Eggs hatch in 25–45 hours at 25–28 °C. Early maturation and stunting is common in crowded populations, resulting in fish that stop growing at 100–120 mm (Moyle 2002).

Warmouth inhabit a wide range of freshwater habitats, mostly in static and slow-moving waters at low elevation and can tolerate a wide range of temperatures, 4–34 °C (Hardy 1978; Page and Burr 1991; Moyle 2002). It has been collected at salinities as high as 17 PSU, but is rare at salinities above 1.5 PSU (Hardy 1978; Peterson 1988; Jenkins and Burkhead 1993). In estuaries, it seems to be found mostly in sloughs on the edges of estuaries, with few records from brackish waters (Wang 1986; Peterson 1988; Van Dyke et al. 2009). As a relatively small fish with a big mouth, Warmouth are opportunistic feeders, feeding small aquatic insects and crustaceans as juveniles, and incorporating larger adult insects, crayfishes, and small fishes as it grows (Jenkins and Burkhead 1994; Moyle 2002). Predators of Warmouth include larger fishes, such as Largemouth Bass, birds, and humans. However, frequent stunting makes this fish a less desirable catch (Moyle 2002).


insects, snails, crayfish, fishes


fishes, birds, humans


other sunfish species

Trophic Status:




General HabitatFresh (nontidal) MarshNone
General HabitatGrass BedNone
General HabitatCoarse Woody DebrisNone
General HabitatSwampNone
General HabitatNontidal FreshwaterNone
General HabitatTidal Fresh MarshNone
General HabitatSalt-brackish marshNone
General HabitatUnstructured BottomNone
Salinity RangeLimnetic0-0.5 PSU
Salinity RangeOligohaline0.5-5 PSU
Salinity RangeMesohaline5-18 PSU
Tidal RangeSubtidalNone
Vertical HabitatNektonicNone

Life History

Tolerances and Life History Parameters

Minimum Temperature (ºC)4Based on geography, range includes areas with winter ice cover.
Maximum Temperature (ºC)34Hardy 1978
Minimum Salinity (‰)0This is a freshwater species.
Maximum Salinity (‰)17.4Warmouth are usually found at salinities under 1.5 PSU (Hardy 1978)
Minimum Reproductive Temperature21.1Hardy 1978
Minimum Reproductive Salinity0This is a freshwater species.
Minimum Length (mm)75Minimum for reproduction (Hardy 1978)
Maximum Length (mm)284Hardy 1978

General Impacts

The Warmouth Lepomis gulosus) is a popular pan-fish in its native range, but is prone to stunting (Moyle 2002). In introduced estuarine populations on the West and East Coast, it is relatively rare and has few ecological or economic impacts (Cohen and Carlton 1995; Far and Ward 1992; Van Dyke et al. 2009). Hybrids have been noted with Green Sunfish, Bluegills, and Pumpkinseeds (Hubbs 1955; Jenkins and Burkhead 1993; Moyle 2002).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
M130 Chesapeake Bay 1897 Def Estab
P090 San Francisco Bay 1931 Def Estab
M090 Delaware Bay 1977 Def Estab
M060 Hudson River/Raritan Bay 1936 Def Estab
P260 Columbia River 1930 Def Estab
G330 Lower Laguna Madre 0 Native Estab
G320 Upper Laguna Madre 0 Native Estab
G310 Corpus Christi Bay 0 Native Estab
G300 Aransas Bay 0 Native Estab
G280 Matagorda Bay 0 Native Estab
G290 San Antonio Bay 0 Native Estab
G270 Brazos River 0 Native Estab
G260 Galveston Bay 0 Native Estab
G250 Sabine Lake 0 Native Estab
G240 Calcasieu Lake 0 Native Estab
G230 Mermentau River 0 Native Estab
G220 Atchafalaya/Vermilion Bays 0 Native Estab
G200 Barataria Bay 0 Native Estab
G210 Terrebonne/Timbalier Bays 0 Native Estab
G190 Mississippi River 0 Native Estab
G170 West Mississippi Sound 0 Native Estab
G160 East Mississippi Sound 0 Native Estab
G150 Mobile Bay 0 Native Estab
G140 Perdido Bay 0 Native Estab
G130 Pensacola Bay 0 Native Estab
G120 Choctawhatchee Bay 0 Native Estab
G110 St. Andrew Bay 0 Native Estab
G100 Apalachicola Bay 0 Native Estab
G090 Apalachee Bay 0 Native Estab
G086 _CDA_G086 (Econfina-Steinhatchee) 0 Native Estab
G080 Suwannee River 0 Native Estab
G078 _CDA_G078 (Waccasassa) 0 Native Estab
G074 _CDA_G074 (Crystal-Pithlachascotee) 0 Native Estab
G070 Tampa Bay 0 Native Estab
G060 Sarasota Bay 0 Native Estab
G050 Charlotte Harbor 0 Native Estab
G045 _CDA_G045 (Big Cypress Swamp) 0 Native Estab
G040 Rookery Bay 0 Native Estab
G030 North Ten Thousand Islands 0 Native Estab
G010 Florida Bay 0 Native Estab
G020 South Ten Thousand Islands 0 Native Estab
S200 Biscayne Bay 0 Native Estab
S196 _CDA_S196 (Cape Canaveral) 0 Native Estab
S190 Indian River 0 Native Estab
S183 _CDA_S183 (Daytona-St. Augustine) 0 Native Estab
S180 St. Johns River 0 Native Estab
S175 _CDA_S175 (Nassau) 0 Native Estab
S170 St. Marys River/Cumberland Sound 0 Native Estab
S160 St. Andrew/St. Simons Sounds 0 Native Estab
S150 Altamaha River 0 Native Estab
S140 St. Catherines/Sapelo Sounds 0 Native Estab
S130 Ossabaw Sound 0 Native Estab
S120 Savannah River 0 Native Estab
S110 Broad River 0 Native Estab
S090 Stono/North Edisto Rivers 0 Native Estab
S100 St. Helena Sound 0 Native Estab
S070 North/South Santee Rivers 0 Native Estab
S080 Charleston Harbor 0 Native Estab
S060 Winyah Bay 0 Native Estab
S056 _CDA_S056 (Northeast Cape Fear) 0 Native Estab
S050 Cape Fear River 0 Native Estab
S040 New River 0 Native Estab
S030 Bogue Sound 0 Native Estab
S020 Pamlico Sound 0 Native Estab
S010 Albemarle Sound 0 Native Estab
GL-I Lakes Huron, Superior and Michigan 0 Native Estab
GL-II Lake Erie 0 Native Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude


Brown, Larry R.; Michniuk, Dennis (2007) Littoral fish assemblages of the alien-dominated Sacramento-San Joaquin Delta, California, 1980-1983 and 2001-2003., Estuaries and Coasts 90: 186-200

Cavallo, Bradley; Merz, Joseph; Setka, Jose (2013) Effects of predator and flow manipulation on Chinook salmon (Oncorhynchus tshawytscha) survival in an imperiled estuary, Environmental Biology of Fishes 393: 393-403

Chapman, Wilbert M. (1942) Alien fishes in the waters of the Pacific Northwest, California Fish and Game 28: 9-15

Cohen, Andrew N.; Carlton, James T. (1995) Nonindigenous aquatic species in a United States estuary: a case study of the biological invasions of the San Francisco Bay and Delta, U.S. Fish and Wildlife Service and National Sea Grant College Program (Connecticut Sea Grant), Washington DC, Silver Spring MD.. Pp. <missing location>

Cope, Edward Drinker (1869) On the distribution of fishes in the Allegheny region of southwestern Virginia, Journal of the Academy of Natural Sciences of Philadelphia 6(2): 207-249

Daniels, Robert A.; Limburg, Karin E.; Schmidt, Robert E; Strayer, David L.; Chambers, R. Christopher (2005) Changes in fish assemblages in the tidal Hudson river, New York., American Fisheries Society Symposium 45: 471-503

Denoncourt, Robert F.; Robbins, Timothy W.; Hesser, Robert (1975) Recent introductions and reintroductions to the Pennsylvania fish fauna of the Susquehanna River drainage above Conowingo Dam, Proceedings of the Pennsylvania Academy of Science 49: 57-58

Denoncourt, Robert F.; Cooper, Edwin L. (1975) A review of the literature and checklist of fishes of the Susquehanna River drainage above Conowingo Dam, Proceedings of the Pennsylvania Academy of Science 49: 121-125

Dill, William A.; Cordone, Almo J. (1997) History and status of introduced fishes in California, 1871-1996, California Department of Fish and Game Fish Bulletin 178: 1-414

Edwards, Robert J.; Contreras-Balderas, Salvador (1991) Historical changes in the ichthyofauna of the lower Rio Grande (Rio Bravo del Norte), Texas and Mexico, Southwestern Naturalist 36(2): 201-212

Farr, Ruth A., Ward, David L. (1992) Fishes of the lower Willamette River, near Portland, Oregon, Northwest Science 67(1): 16-22

Feyrer, Frederick; Healey, Michael P. (2003) Fish community structure and environmental correlates in the highly altered southern Sacramento-San Joaquin Delta., Environmental Biology of Fishes 66: 123-132

Fowler, Henry W. (1919) A list of the fishes of Pennsylvania, Proceedings of the Biological Society of Washington 32: 49-74

Fuller, P.M., Nico, L.G., Williams, J.D. (1999) Nonindigenous fishes introduced into inland waters of the United States, American Fisheries Society, Bethesda, MD. Pp. <missing location>

Grimaldo, Lenny; Miller, Robert E.; Hymanson, ZacharyPeregrin, Chris M., (2012) Fish assemblages in reference and restored tidal freshwater marshes of the San Francisco estuary, San Francisco Estuary and Watershed Science 10(1):

Hardy, Jerry D., Jr. (1978) Development of fishes of the Mid-Atlantic Bight. Vol. 3. Aphredoderidae through Rachycentridae., In: (Eds.) . , Washington DC. Pp. <missing location>

Horwitz, Richard J. (1986) Fishes of the Delaware estuary in Pennsylvania., In: Majundar, S.K., Brenner, F. J., Rhoads, A. F.(Eds.) Endangered and Threatened Species Programs in Pennsylvania.. , Philadelphia. Pp. 177-201

Hubbs, Carl L. (1955) Hybridization between fish species in nature, Systematic Zoology 4: 1-20

Jenkins, Robert E.; Burkhead, Noel M. (1993) Freshwater Fishes of Virginia, American Fisheries Society, Bethesda, MD. Pp. <missing location>

Keup, Lowell, Bayless, Jack (1964) Fish distribution at varying salinities in Neuse River Basin, North Carolina, Chesapeake Basin 5(3): 119-123

Lampman, Ben Hur (1946) Coming of the Pond Fishes, Binfords & Mort, Portland, OR. Pp. <missing location>

Lee, David S.; Norden, Arnold; Gilbert, Carter, R.; Franz, Richard (1976) A list of the freshwater fishes of Maryland and Delaware, Chesapeake Science 17(3): 205-211

Leidy, R. A. (2007) <missing title>, San Francisco Estuary Institute, Oakland. Pp. <missing location>

Lever, Christopher (1996) Naturalized fishes of the world, Academic Press, London, England. Pp. <missing location>

Massmann, William H.; Ladd, Ernest C.; McCutcheon, Henry M. (1952) A biological survey of the Rappahannock River, Virginia, Virginia Fisheries Laboratory Special Scientific Report 6: 1-152

Matern, Scott A.; Moyle, Peter; Pierce, Leslie C. (2002) Native and alien fishes in a California estuarine marsh: twenty-one years of changing assemblages, Transactions of the American Fisheries Society 131: 797-816

Maurakis, Eugene; Woolcott, William S.; Jenkins, Robert E. (1987) Physiographic analyses of the longitudinal distribution of fishes in the Rappahannock River, Virginia, ASB Bulletin 34(1): 1-14

McIvor, Carole C.; Odum, William E. (1988) Food, predation risk, and microhabitat selection in a marsh fish assemblage, Ecology 69(5): 1341-1351

McKeown, Paul E. (1984) Additions to ichthyofauna of the Susquehanna River with a checklist of fishes of the Susquehanna River drainage below Conowingo Dam, Proceedings of the Pennsylvania Academy of Science 58: 187-192

Mills, Edward L.; Scheuerell, Mark D.; Carlton, James T.; Strayer, David (1997) Biological invasions in the Hudson River: an inventory and historical analysis., New York State Museum Circular 57: 1-51

Moyle, Peter B. (2002) Inland Fishes of California, revised and expanded, University of California Press, Berkeley CA. Pp. <missing location>

Page, Lawrence M.; Burr, Brooks M. (1991) Freshwater Fishes: North America North of Mexico, Houghton-Mifflin, Boston. Pp. <missing location>

Peterson, Mark S. (1988) Comparative physiological ecology of centrarchids in hyposaline environments, Canadian Journal of Fisheries and Aquatic Sciences 45: 827-833

Raasch, Maynard S. (1996) Delaware's Freshwater and Brackish-water Fishes: A Popular Account, T.F.H. Publications, Neptune, NJ. Pp. <missing location>

Shebley, W. H. (1917) Introduction of food and game fishes into the waters of California., California Fish and Game 3(1): 1-12

Smith, C. Lavett, Lake, Thomas R. (1990) Documentation of the Hudson River fish fauna, American Museum Novitates 2981: 1-17

Smith, Hugh M. (1895) A review of the history and results of the attempts to acclimatize fish and other water animals in the Pacific states., Bulletin of the U. S. Fish Commission 15: 379-472

Smith, Hugh M.; Bean, Barton A. (1898) List of fishes known to inhabit the waters of the District of Columbia and vicinity., Bulletin of the U. S. Fish Commission 18: 179-187

Sytsma, Mark D.; Cordell, Jeffrey R.; Chapman, John W.; Draheim, Robyn, C. (2004) <missing title>, Center for Lakes and Reservoirs, Portland State University, Portland OR. Pp. <missing location>

USGS Nonindigenous Aquatic Species Program 2003-2024 Nonindigenous Aquatic Species Database.

Van Dyke, Erick, S. Storch, Adam J. Reesman, Martyne J. (2009) Seasonal composition and distribution of fish species in the lower Columbia slough; Completion Repor, City of Portland, Bureau of Environmental Services, Portland OR. Pp. <missing location>

Waldman, John R.; Lake, Thomas R.; Schmidt, Robert E. (2006) Biodiversity and zoogeography of the fishes of the Hudson River watershed and estuary, American Fisheries Society Symposium 51: 129-150.

Wang, Johnson, C. S.; Kenehahan, Ronnie (1979) Fishes of the Delaware estuaries - a guide to the early life histories, EA Associates, Towson MD. Pp. <missing location>

Worth, S. G. (1895) A review of the history and results of the attempts to acclimatize fish and other water animals in the Pacific states, In: (Eds.) Report of the United States Commission of Fish and Fisheries for 1893. , Washington D.C.. Pp. 78-138