Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Lepomis cyanellus
(green sunfish)

Toolbox

Datasheet

Lepomis cyanellus (green sunfish)

Summary

  • Last modified
  • 08 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Natural Enemy
  • Host Animal
  • Preferred Scientific Name
  • Lepomis cyanellus
  • Preferred Common Name
  • green sunfish
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Chordata
  •       Subphylum: Vertebrata
  •         Class: Actinopterygii
  • Summary of Invasiveness
  • Lepomis cyanellus is a perciform fish that is native to the central plains of North America, but has been introduced to most of the rest of the USA, and some other countries. Introduced L. cyanellus co...

Don't need the entire report?

Generate a print friendly version containing only the sections you need.

Generate report

Pictures

Top of page
PictureTitleCaptionCopyright
Green sunfish - Lepomis cyanellus
TitleAdult
CaptionGreen sunfish - Lepomis cyanellus
CopyrightPublic Domain - Original artwork from U.S. Fish and Wildlife Service/Duane Raver
Green sunfish - Lepomis cyanellus
AdultGreen sunfish - Lepomis cyanellusPublic Domain - Original artwork from U.S. Fish and Wildlife Service/Duane Raver

Identity

Top of page

Preferred Scientific Name

  • Lepomis cyanellus Rafinesque, 1819

Preferred Common Name

  • green sunfish

Other Scientific Names

  • Apomotis cyanellus Rafinesque, 1819
  • Chaenobryttus cyanellus Rafinesque, 1819
  • Icthelus cyanella Rafinesque, 1819
  • Lepidomus cyanellus Rafinesque, 1819
  • Lepomis cyanella Rafinesque, 1819
  • Lepomus cyanellus Rafinesque, 1819
  • Telipomis cyanelus Rafinesque, 1819

International Common Names

  • English: black perch; blue spotted sunfish; creek perch; goggle eye; green perch; mud bass; perch; pond perch; rice slick; ricefield slick; rubbertail; sand bass; shade perch; sunfish

Local Common Names

  • China: Lan tai yang yu
  • Denmark: gron solaborre
  • Finland: viheraurinkoahven
  • France: crapet vert
  • Germany: grasbarsch; gruner sonnenbarsch; sonnenbarsch
  • Netherlands: groene zonnebaars
  • Norway: gron solabbor
  • Spain: pez sol
  • Sweden: gron solabborre

Summary of Invasiveness

Top of page

Lepomis cyanellus is a perciform fish that is native to the central plains of North America, but has been introduced to most of the rest of the USA, and some other countries. Introduced L. cyanellus compete with and prey on native fish species, reducing their populations, altering their population structure, relative dominance and distribution patterns. Their aggressive nature allows them to compete with larger fish for prey, and prey on eggs and young of other fish species. L. cyanellus are remarkably tolerant of crowding under which conditions they are often stunted in size, creating management problems in small lakes. Large populations often compete with the young of other species for food, or prey upon them causing negative impacts to both game fish and native species. In streams where L. cyanellus have been introduced in California, it is believed to have helped deplete the California roach, Hesperoleucus symmetricus. L. cyanellus along with other predatory fish species is also thought to be responsible for the decline of native frogs and salamanders in the USA. L. cyanellus is listed as a major invasive species in parts of Arizona (USDA, 2012) and California (Dill and Cordone, 1997). New Jersey state authorities list it as a potentially dangerous species due to its ability to outcompete native fish species; it is listed as an invasive species of concern in Georgia and Florida. Marsh (2010), in a discussion on the advantages of genetic biocontrol in the Colorado River basin system, referred to the species as “one of the most invasive, pervasive, and destructive species in the basin, not even recognized as an invasive species by some states”. Olden and Poff (2005), in a study of long-term trends (> 160 years) of fish species distributions in the Lower Colorado River Basin, to identify those native species exhibiting the greatest rates of decline and those non-native species exhibiting the highest rates of spread, found that L. cyanellus was amongst the fastest expanding invaders in the basin and the most invasive in terms of negative impacts on native fish communities. As far as is known, L. cyanellus is not listed as a threatened species in any part of its native range.

Taxonomic Tree

Top of page
  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Chordata
  •             Subphylum: Vertebrata
  •                 Class: Actinopterygii
  •                     Order: Perciformes
  •                         Suborder: Percoidei
  •                             Family: Centrarchidae
  •                                 Genus: Lepomis
  •                                     Species: Lepomis cyanellus

Notes on Taxonomy and Nomenclature

Top of page

Lepomis cyanellus, or the green sunfish as it is commonly known, is a member of the Centrarchidae, the sunfish family. This family includes some of the more attractive and brightly coloured of the freshwater fishes of North America. The scientific name for the green sunfish is derived from the Latin Lepomis meaning “scaled operculum cover“, and cyanellus meaning “blue”.

Description

Top of page

Becker (1983) and Etnier and Starnes (2001) provide detailed descriptions of the anatomical features unique to L. cyanellus. It has an elongate laterally compressed body with a large mouth which extends beyond the broad anterior edge of the eye. The dorsal surface is brown to olive, with small scattered black flecks; the sides are of a lighter coloration, with females having 7-12 vertical bars (Etnier and Starnes, 2001). The two broad dorsal fins are joined, appearing as one; the first dorsal fin has 9-11 spines while the second has 10-12 soft rays. The pectoral fins are short and rounded. The anal fin has 3 spines and 9-10 soft rays, and the pelvic fin a single spine and 5 soft rays. The dorsal, anal and caudal fin margins of males are often bright yellow or orange. A dark spot is present at the rear of the dorsal and anal fins.

Distribution

Top of page

L. cyanellus has a wide distribution range and is native to the central plains of North America between the Appalachian and Rocky Mountains, from Ontario (Canada) and New York state in the north to the Gulf Coast and northern Mexico in the south. It has been so widely introduced in the USA that it is now present in almost every state, including Atlantic and Pacific slope drainages and Hawaii, except for parts of the north-east of the country (Page and Burr, 1991). Populations are stronger within the native range.

According to Ma et al. (2003), L. cyanellus was introduced to China in 1999 for use as food; as far as is known, there have been no further reports on its status there. There is also little information on its presence in the Philippines, where it was introduced from the United States in 1950 for aquaculture purposes, apart from its being listed as present there by Halos et al. (2004).  Although there are reports of an established population in Japan (Welcomme, 1988; Froese and Pauly, 2013), a survey by Mito and Uesugi (2004) on alien species recognised as established or present in the wild (as of 27 October 2004) does not mention L. cyanellus -- L. gibbosus is the only Lepomis species recorded.

There is some confusion as to whether L. cyanellus is present in Germany today. Maitland (1977) (cited in Soes et al., 2011) reported that its introduction by aquarists to the Frankfurt area led to established populations; this appears to contradict an earlier observation in 1959 by Sterba (as cited in Soes et al., 2011), that the species had disappeared from Europe. Lelek (as cited in Arnold, 1990) said there was no evidence for the presence of L. cyanellus in Germany. Elvira (2001), in a report identifying the non-native freshwater fishes established in Europe, lists L. cyanellus not only as an exotic freshwater species introduced to European waters but also as being present and established in Germany. The NOBANIS (European Network of Invasive Alien Species) website lists it as being established in Germany, and first recorded in 1965 (NOBANIS (2005), citing Geiter et al. (2002) (which Nehring (2005) also cites)). An ornamental fish stockist based in Germany lists the green sunfish on its website (www.aquarium_glaser.de) as being suitable for large cold water aquaria and garden ponds, adding that the specimens are German pond-bred and only occasionally found in the trade (Aquarium Glaser, 2013).

Attempts to introduce L. cyanellus in tropical Africa have been largely unsuccessful; where it has established itself, occupying smallish, well-vegetated dams and becoming over-populated, it is regarded a pest, according to Jackson (1976).

Lever (1996) reported the presence of naturalized populations of L. cyanellus in Germany (one population), Asia (2), Africa (4), Brazil (1), and Oceania (1).

Distribution Table

Top of page

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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ChinaPresentIntroducedMa et al., 2003Introduced for food
JapanPresentIntroducedWelcomme, 1988Introduced for aquaculture
Korea, Republic ofPresentIntroducedWelcomme, 1988Introduced for aquaculture
PhilippinesPresentIntroduced1950Welcomme, 1988; Halos et al., 2004Introduced for aquaculture

Africa

CongoPresentIntroducedWelcomme, 1988; Froese and Pauly, 2013Introduced for aquaculture
KenyaPresentIntroducedWelcomme, 1988; Froese and Pauly, 2013Introduced for aquaculture
MadagascarPresentIntroduced1954Welcomme, 1988; Froese and Pauly, 2013Introduced for aquaculture
MauritiusPresentIntroduced1950Welcomme, 1988; Froese and Pauly, 2013Introduced for aquaculture
MoroccoPresentIntroducedWelcomme, 1988; Froese and Pauly, 2013Introduced for aquaculture
South AfricaPresentIntroduced1939Welcomme, 1988; Froese and Pauly, 2013Introduced for aquaculture
SwazilandPresentIntroduced1939Welcomme, 1988; Froese and Pauly, 2013Introduced for aquaculture
ZambiaPresentIntroducedWelcomme, 1988; Froese and Pauly, 2013Introduced for aquaculture
ZimbabweAbsent, formerly presentIntroduced1940Welcomme, 1988; Froese and Pauly, 2013Introduced for aquaculture; disappeared by 1970.

North America

CanadaPresentPresent based on regional distribution.
-OntarioPresentNative Invasive Meredith and Houston, 1988Native to part of province; introduced elsewhere.
MexicoPresentNativeNatureServe, 2013; USGS, 2013Northern Mexico
USAPresentPresent based on regional distribution.
-AlabamaPresentNativeNatureServe, 2013; USGS, 2013
-ArizonaPresentIntroduced Invasive USDA, 2012; NatureServe, 2013; USGS, 2013
-ArkansasPresentNativeNatureServe, 2013; USGS, 2013
-CaliforniaPresentIntroduced Invasive Dill and Cordone, 1997; NatureServe, 2013; USGS, 2013
-ColoradoPresentIntroduced Invasive NatureServe, 2013; USGS, 2013
-ConnecticutPresentIntroducedNatureServe, 2013; USGS, 2013
-DelawarePresentIntroducedNatureServe, 2013; USGS, 2013
-District of ColumbiaPresentIntroducedNatureServe, 2013; USGS, 2013
-FloridaPresentIntroducedNatureServe, 2013; USGS, 2013Absent from peninsular Florida
-GeorgiaPresentIntroduced Invasive NatureServe, 2013; USGS, 2013
-HawaiiPresentIntroducedNatureServe, 2013; USGS, 2013
-IdahoPresentIntroducedNatureServe, 2013; USGS, 2013
-IllinoisPresentNativeNatureServe, 2013; USGS, 2013
-IndianaPresentNativeNatureServe, 2013; USGS, 2013
-IowaPresentNativeNatureServe, 2013; USGS, 2013
-KansasPresentNativeNatureServe, 2013; USGS, 2013
-KentuckyPresentNativeNatureServe, 2013; USGS, 2013
-LouisianaPresentNativeNatureServe, 2013; USGS, 2013
-MainePresentIntroducedNatureServe, 2013; USGS, 2013
-MarylandPresentIntroducedNatureServe, 2013; USGS, 2013
-MassachusettsPresentIntroducedNatureServe, 2013; USGS, 2013
-MichiganPresentNativeNatureServe, 2013; USGS, 2013
-MinnesotaPresentNativeNatureServe, 2013; USGS, 2013
-MississippiPresentNativeNatureServe, 2013; USGS, 2013
-MissouriPresentNativeNatureServe, 2013; USGS, 2013
-MontanaPresentIntroducedNatureServe, 2013; USGS, 2013
-NebraskaPresentNativeNatureServe, 2013; USGS, 2013
-NevadaPresentIntroduced Invasive NatureServe, 2013; USGS, 2013
-New JerseyPresentIntroduced Invasive NatureServe, 2013; USGS, 2013
-New MexicoPresentNativeNatureServe, 2013; USGS, 2013
-New YorkPresentNativeNatureServe, 2013; USGS, 2013Native to part of state; introduced elsewhere.
-North CarolinaPresentNativeNatureServe, 2013; USGS, 2013
-North DakotaPresentIntroducedNatureServe, 2013; USGS, 2013
-OhioPresentNativeNatureServe, 2013; USGS, 2013
-OklahomaPresentNativeNatureServe, 2013; USGS, 2013
-OregonPresentIntroducedNatureServe, 2013; USGS, 2013
-PennsylvaniaPresentIntroducedNatureServe, 2013; USGS, 2013
-South CarolinaPresentIntroducedNatureServe, 2013; USGS, 2013
-South DakotaPresentIntroducedNatureServe, 2013; USGS, 2013
-TennesseePresentNativeNatureServe, 2013; USGS, 2013
-TexasPresentNativeNatureServe, 2013; USGS, 2013
-UtahPresentIntroducedNatureServe, 2013; USGS, 2013
-VirginiaPresentIntroducedNatureServe, 2013; USGS, 2013
-WashingtonPresentIntroducedNatureServe, 2013; USGS, 2013
-West VirginiaPresentNativeNatureServe, 2013; USGS, 2013
-WisconsinPresentNativeNatureServe, 2013; USGS, 2013
-WyomingPresentIntroducedNatureServe, 2013; USGS, 2013

South America

BrazilPresentIntroducedWelcomme, 1988

Europe

GermanyAbsent, unreliable recordIntroducedLever, 1996; Soes et al., 2011

History of Introduction and Spread

Top of page

Expansion of the range of L. cyanellus in the United States has occurred mainly as a result of accidental stocking. It has been accidentally stocked as bluegill L. macrochirus, with other intended species as a stock contaminant (Dill and Cordone, 1997; USGS, 2013), as forage fish for smallmouth bass (Micropterus dolomieu) in fish farms, and as a sport or game fish. It has also escaped from holding ponds and spread of its own accord through stream channels and over flooded areas (Dill and Cordone, 1997) or via drainage ditches and irrigation waterways connecting river systems.

It has been introduced to other countries primarily as forage for black bass (Micropterus spp.) or for the purpose of aquaculture or sport fishing (Welcomme, 1988). Attempts to introduce it in tropical Africa have been largely unsuccessful; where it has established itself, occupying smallish, well-vegetated dams and becoming over-populated, it is regarded a pest, according to Jackson (1976).

According to Ma et al. (2003), L. cyanellus was introduced to China in 1999 for use as food; as far as is known, there have been no further reports on its status there. It was introduced to the Philippines from the USA in 1950 for aquaculture (Wellcome, 1988).

Maitland (1977) (cited in Soes et al., 2011) reported that the introduction of L. cyanellus by aquarists to the Frankfurt area in Germany led to established populations (Sterba (as cited in Soes et al., 2011) reported that the species had disappeared from Europe, but other sources indicate that it is still present there).

Introductions

Top of page
Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Brazil USA 1930-1939 Aquaculture (pathway cause) Yes Froese and Pauly (2013); Welcomme (1988)
China USA 1999 Yes Ma et al. (2003) Introduced as food
Congo USA  unknown Aquaculture (pathway cause)Froese and Pauly (2013); Welcomme (1988)
Germany USA   Yes Welcomme (1988)
Germany USA   Aquarium trade (pathway cause) Yes Maitland (1977)
Germany USA 1906 Aquarium trade (pathway cause) No Soes et al. (2011)
Hawaii USA  unknown Froese and Pauly (2013); Welcomme (1988)
Japan USA  unknown Aquaculture (pathway cause)Froese and Pauly (2013); Welcomme (1988)
Kenya USA  unknown Froese and Pauly (2013); Welcomme (1988)
Korea, Republic of Japan 1969 Yes Froese and Pauly (2013); Welcomme (1988)
Madagascar USA 1954 Live food or feed trade (pathway cause)Froese and Pauly (2013); Welcomme (1988)
Mauritius USA 1950 Hunting, angling, sport or racing (pathway cause) Yes Froese and Pauly (2013); Welcomme (1988) Stunted population
Morocco USA  unknown Yes Froese and Pauly (2013); Welcomme (1988)
Philippines California 1950 Aquaculture (pathway cause)Froese and Pauly (2013); Welcomme (1988)
South Africa USA 1939 Live food or feed trade (pathway cause) Yes Froese and Pauly (2013); Jackson (1976); Welcomme (1988) Forage fish for bass. Where present in small, stagnant, well-vegetated dams, become overpopulated; regarded as pest
Swaziland South Africa unknown Live food or feed trade (pathway cause) Yes Froese and Pauly (2013); Welcomme (1988) Forage fish for bass
Zambia USA Aquaculture (pathway cause)Unknown Yes Froese and Pauly (2013); Welcomme (1988)
Zimbabwe South Africa 1940 Aquaculture (pathway cause)Froese and Pauly (2013); Welcomme (1988) Forage fish for bass

Risk of Introduction

Top of page

The aggressive and predatory behaviour of L. cyanellus, and its ability to survive in a range of habitats and environmental extremes (low dissolved oxygen, high turbidity, alkalinity and siltation) indicate a high risk of spread and the potential to have a negative impact on native species. New connections such as drainage networks between lakes, rivers, ponds and streams, and flooding, enable its dispersal into water bodies well beyond its native range. Its use as a live bait fish also presents a risk of accidental introduction and subsequent establishment as in the case of the Great Lakes Basin (Mills et al., 1993; Litvak and Mandak, 1999).

Habitat

Top of page

Tolerant of a wide range of habitats, L. cyanellus can be found in small muddy creeks, pools, streams, weedy backwaters with temporary flow to overflow ponds, shallow lakes, impoundments, ponds, springs, areas of rivers with little flow and sometimes the margins of large rivers of low gradient; it is highly adaptable, tending to become abundant in rocky areas of lakes or streams. It is commonly found near the shore and around cover such as stems of vegetation, rocks, or woody debris. Population abundance is positively correlated with percentage vegetative cover (Moyle and Nichols, 1973). It can tolerate turbidity, siltation, intermittent flow, high temperatures, high salt content and low dissolved oxygen. It tolerates crowding, often becoming stunted in size (Wellcome, 1988; Carlander, 1977), resulting in management problems in small lakes. Its aggressive nature allows it to quickly dominate wherever it is introduced, taking over in a very short period, and becoming the only species in a particular area, thus posing a major threat to native species (Etnier and Starnes, 2001). As a pioneering species, it is also the first species to find its way to newly created farm ponds and the first to repopulate streams following periods of drought (Tomelleri and Eberle, 1990; Pflieger, 1975, 1997). In northwestern Ontario it is found in habitats ranging from the shallow regions of moderately-sized lakes to small streams where it can be found in dense growths of vegetation (Crossman, 1976).

Although tolerant of poor conditions, it may prefer cleaner, larger, water bodies, with a moderate flow as growth has been observed to be greater in these areas than in more turbid waters or smaller streams and ponds (Carlander, 1977).

Habitat List

Top of page
CategoryHabitatPresenceStatus
Brackish
Lagoons Principal habitat Harmful (pest or invasive)
Lagoons Principal habitat Natural
Freshwater
Irrigation channels Principal habitat Harmful (pest or invasive)
Irrigation channels Principal habitat Natural
Lakes Principal habitat Harmful (pest or invasive)
Lakes Principal habitat Natural
Ponds Principal habitat Harmful (pest or invasive)
Ponds Principal habitat Natural
Reservoirs Principal habitat Harmful (pest or invasive)
Reservoirs Principal habitat Natural
Rivers / streams Principal habitat Harmful (pest or invasive)
Rivers / streams Principal habitat Natural

Biology and Ecology

Top of page

Genetics

L. cyanellus hybridizes with longear (L. megalotis), orange-spotted (L. humilis) and redbreast (L. auritus) sunfishes, bluegill (L. macrochirus), and pumpkinseed (L. gibbosus) (Scott and Crossman, 1973). Population genetic studies have been carried out on this species along with four other stream fishes (Husemann et al., 2012).  

Reproductive Biology

Age at maturation of L. cyanellus appears to vary depending on location; it has been reported at age one in Missouri, Illinois and Iowa, but not until age three in Michigan (Hubbs and Cooper, 1935; Sprugel, 1955). In Texas, sexual maturity is reached in less than 6 months (White, 1971). Males grow faster than females and appear to live longer (Carlander, 1977), with males and females maturing at minimum lengths of 45 and 66 mm, respectively. In Ontario, L. cyanellus live for 7 to 9 years and may grow to a length of 203 mm (Scott and Crossman, 1973). In Ohio, they may grow to 274 mm and weigh 400 g (Trautman, 1957). A Kansas specimen was recorded at 305 mm, weighing 964 g (Scott and Crossman, 1973). Carlander (1977) reported maximum age, length and weight of green sunfish of about 10 years, 276 mm and 408 g, respectively. In crowded conditions, the fish do not grow as well and stunting may occur (Carlander, 1977; Wellcome, 1988).

In the United States L. cyanellus spawn from late spring to early summer or April to August, when water temperatures range from 15 to 31°C. Spawning occurs from mid-May to early August in Wisconsin (Hunter, 1963) and from June to August in Illinois, Maryland, Michigan and Iowa (Carlander, 1977). There is no information on spawning in the Canadian part of the range, but on the basis of data from the USA, it is believed that it would probably occur in late spring to early summer in Canada as well. Peak spawning activity occurs at 20 to 28° C, and multiple spawnings occur every eight or nine days throughout the season (Hunter, 1963). Temperatures above 24°C for extended periods not only reduce nesting activity but may also stop spawning, and cause gonads to regress (Kaya, 1973).

The males are highly territorial and construct nests in colonies, in sunlit waters of 15 to 25 cm in depth, and sheltered by rocks, logs, clumps of vegetation and trunks (Carlander, 1977). They court females by rushing towards them and returning quickly to the nest. Hunter (1963) has described courtship and parental care in green sunfish in some detail. Females will lay from 2000 to 50 000 eggs (Carlander, 1977). Eggs are yellowish, adhesive, and 1.9 to 1.4 mm in diameter. Once eggs have been laid, the male guards and fans the eggs which hatch in 3 to 5 days (Scott and Crossman, 1973). The larvae are free swimming at 4.2 to 4.7 mm, within two days of hatching, and reach an average size of 23.7 mm by 57 days. Meyer (1970), Taubert (1977) and Auer (1982) have studied the eggs and larval development of green sunfish.

Trautman (1957) observed that growth of fry was rapid and young-of-the-year fish in Ohio reached lengths of 20 to 64 mm by October, and weights of 1.3 to 25 g. Increase in length of approximately 20 mm per year was achieved; no weight data were provided but in Wisconsin increases in weight varied from 10 to 42 g per year, with large increases in the fourth, fifth and sixth years (Carlander, 1977).

Physiology and Phenology

Of the centrarchids found in California, only juvenile L. cyanellus possess a chemical alarm signal similar to those found in cyprinids. When alarm pheromones, released due to mechanical damage to skin, are detected, they remain still with dorsal fins erect (Brown and Brennan, 2000).

Longevity

In Ontario, L. cyanellus have been reported to live for 7 to 9 years (Scott and Crossman, 1973). Applegate et al. (1967) observed that in Bull Shoals reservoir in Arkansas and Missouri, they did not live beyond 6 years of age. In captivity, they lived to 7.5 years (Carlander, 1977). Wang (1986) reported them living 7 to 9 years in California, age at maturity being 1-2 years. Moyle (2002) reported a life span of 10 years with maturity at the beginning of the third year in Californian L. cyanellus.

Activity Patterns

L. cyanellus show strong homing tendencies (Hasler and Wibly, 1958) and are most active at dawn and dusk, preferring to remain hidden in available cover unless feeding (Carlander, 1977).

Population Size and Density

L. cyanellus are remarkably tolerant of crowding, under which conditions they are often stunted, creating management problems in small lakes (Carlander, 1977; Wellcome, 1988).

Nutrition

L. cyanellus is highly predaceous, large populations competing with the young of other species for food or preying directly upon them (Minckley, 1973). Young feed on zooplankton (Carlander, 1977); fry subsequently eat aquatic insects and fish eggs while adults feed on fish eggs, snails, frogs, molluscs, insects and their young, crayfish, small fish and sometimes plant material (Struber et al., 1982). Adults have a larger mouth than most other sunfishes, and thus are able to swallow fish almost half the size of their own body; the adult diet consists of other small fishes and molluscs (Scott and Crossman, 1973). In Ontario, L. cyanellus is usually found in association with carp, Cyprinus carpio, and the smallmouth bass Micropterus dolomieu (Hallam, 1959), falling prey to both species as well as preying on their young. L. cyanellus have also been observed to feed on bryozoans (Fredericella sultana) (Applegate, 1966).

Associations

L. cyanellus is often found in association with other centrarchids, particularly the pumpkinseed (Lepomis gibbosus) and Longear Sunfish (L. megalotis) with which it is known to hybridize (Carlander, 1977). Hallam (1959), describing typical southwestern Ontario habitats for L. cyanellus in Canada, found that it inhabited areas frequented by rock bass (Ambloplites rupestris) and smallmouth bass (Micropterus dolomieu).

Environmental Requirements

L. cyanellus is tolerant of a wide range of conditions. It is able to survive in both clear and turbid waters, although high species abundance is positively correlated with moderate (25-100 JTU) turbidities (Trautman, 1957). It is able to tolerate conditions of high turbidity <3500 JTU). Optimal dissolved oxygen requirements have been reported to be more than 5 mg/l (Petit, 1973), with lethal levels being less than or equal to 1.5 mg/l (Moore, 1942). Optimal pH range is believed to be from 6.5 to 8.5 (Stroud, 1967), with mortality occurring at pH £ 4.0 or ³ 10.35 (Ultsch, 1978). Green sunfish in Canada are reported to prefer a pH range of 6.0 to 9.6 and oxygen concentrations of 4 to 8 ppm (Meredith and Houston, 1988). Green sunfish will not tolerate salinities greater than 5.6 ppt (Kilby, 1955), optimal salinities being less than 3.6 ppt (Tebo and McCoy, 1964). Adults prefer 28.2°C, and when possible avoid temperatures above 31°C or below 26°C (Beitinger et al., 1975). There are reports of green sunfish in the field surviving in water temperatures as high as 36°C (Proffitt and Benda, 1971). Laboratory trials suggest that L. cyanellus, when properly acclimated, survives well at water temperatures near 1°C (Cortemeglia and Beitinger, 2008).

According to Childers (1967), optimal temperatures for spawning and successful development range from 20° to 27°C. Spawning will not occur below 19°C or above 31°C (Hunter, 1963). Optimal temperatures for fry range from 18° to 26°C (Coutant, 1977; T. Hardin and K. Bovee, US Fish and Wildlife Service, Instream Flow Group, Fort Collins, Colorado, USA, unpublished data, 1978, cited in Stuber et al., 1982); the range of temperature tolerance is reported to be 10-36°C.

Adults are found in low current velocity areas (Moyle and Nichols, 1973); preferred current velocities are below 10 cm/sec, while velocities of up to 25 cm/sec are tolerated (T. Hardin and K. Bovee, US Fish and Wildlife Service, Instream Flow Group, Fort Collins, Colorado, USA, unpublished data, 1978, cited in Stuber et al., 1982). Though tolerant of high turbidity, L. cyanellus is susceptible to heavy metals and nitrogen pollution, both common conditions in southwestern Ontario (Meredith and Houston, 1988). According to Meredith and Houston (1988), acid rain may have some influence on limiting populations in Ontario. Nests have been reported in polluted and channelized rivers and creeks, where dissolved oxygen levels were as low as 4 ppm (Wang, 1986; Wang and Reyes, 2008).

According to Cortemeglia and Beitinger (2008), the success of L. cyanellus in Great Plains streams and its widespread dispersion throughout North America may have been aided by its temperature tolerance ability.

Natural Food Sources

Top of page
Food SourceLife StageContribution to Total Food Intake (%)Details
fish eggs Adult/Fry
frogs Adult
insect larvae Adult/Fry
insects Adult
juvenile fish Adult
molluscs Adult
plant material Adult
small crayfish Adult
small fish Adult
Zooplankton Fry/Larval

Climate

Top of page
ClimateStatusDescriptionRemark
Am - Tropical monsoon climate Preferred Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
B - Dry (arid and semi-arid) Preferred < 860mm precipitation annually
BW - Desert climate Preferred < 430mm annual precipitation
Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Preferred Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)

Water Tolerances

Top of page
ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Dissolved oxygen (mg/l) 5 Optimum All Stages Lethal levels less than or equal to 1.5 mg/l
Hardness (mg/l of Calcium Carbonate) 900 2000 Optimum Adult
Salinity (part per thousand) 3.6 Optimum All Stages Up to 5.6 tolerated
Spawning temperature (ºC temperature) 19 31 Optimum Broodstock
Turbidity (JTU turbidity) 25 100 Optimum All Stages
Velocity (cm/h) 36000 Optimum Adult Up to 90 000 cm/h tolerated
Water pH (pH) 6.5 8.5 Optimum Adult Mortality may occur at pH less than or equal to 4.0 or pH greater than or equal to 10.35
Water temperature (ºC temperature) 28.2 Optimum Adult Where possible, avoids temperatures >31°C and below 26°C; reported to survive temperatures as high as 36°C and 38°C; spawning requires 19-31°C; tolerances for juveniles may vary
Water temperature (ºC temperature) 15 30 Optimum Broodstock

Natural enemies

Top of page
Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Ameiurus Predator All Stages not specific
Ictalurus punctatus Predator All Stages not specific
Micropterus salmoides Predator All Stages not specific Morris and Mischke, 2000
Pylodictis olivaris Predator All Stages not specific

Means of Movement and Dispersal

Top of page

Accidental introductions of L. cyanellus can occur through the transportation and release of live bait by fishermen and anglers; increases in distribution of the green sunfish in the Great Lakes basin have been attributed to baitfish release (Mills et al., 1993; Litvak and Mandrak, 1999). Many unauthorized introductions of green sunfish across the United States were the result of authorized introductions of bluegill sunfish (L. macrochirus) or largemouth bass (Micropterus salmoides) using stocks which were contaminated with green sunfish (Benson, 1999).

In California, at the beginning of the 20th century, green sunfish were stocked in large numbers through the combined efforts of the California Division of Fish and Game and some sport fish clubs. Some of these green sunfish were raised in rearing ponds, and some were the result of fish rescue from overflow areas (Dill and Cordone, 1997). In 1944, the green sunfish was classified by the California Division of Fish and Game as an undesirable fish (compared with the bluegill) for stocking, although it retained its status as a "gamefish". Since that time it has been exterminated intentionally by the State in some trout waters. According to Dill and Cordone (1997), the introduction of the green sunfish to California was probably a mistake and that the waters of California would be better off without this species.

L. cyanellus were moved to Germany for the aquarium trade and to a number of other countries for aquaculture or as forage for other fish. Introduction to Korea is reported to have been ‘to fill a vacant niche’ (Welcomme, 1988).

Green sunfish have also expanded their distribution outside their native range via the waterways (stream channels, irrigation drainage systems, flooded areas) of the United States.

Pathway Causes

Top of page
CauseNotesLong DistanceLocalReferences
AquacultureIntroduced to the Congo, Zambia, Japan and the Philippines for aquaculture Yes Welcomme, 1988
Flooding and other natural disastersExpanded its range to several US states Yes Yes USGS, 2013
ForageTo Madagascar (1954), South Africa and Swaziland (1939), Zimbabwe (1940), and Brazil (1930s) Yes Welcomme, 1988
Hunting, angling, sport or racingIntroduced to Mauritius (1950) for sport Yes Welcomme, 1988
Intentional releaseRelease as live baitfish probably increased distribution in N. American Great Lakes basin Yes Yes Litvak and Mandrak, 1999; Mills et al., 1993
Interbasin transfersExpanded its range to several US states Yes Yes USGS, 2013
Interconnected waterwaysExpanded its range to several US states Yes Yes USGS, 2013
Pet tradeIntroduced to Germany by aquarists Yes Soes et al., 2011
StockingExpanded its eastward and westward range in the USA mainly due to accidental stocking Yes USGS, 2013

Pathway Vectors

Top of page
VectorNotesLong DistanceLocalReferences
Aquaculture stock Yes Yes Welcomme, 1988
WaterRiver drainage systems Yes Yes USGS, 2013

Impact Summary

Top of page
CategoryImpact
Cultural/amenity Positive
Economic/livelihood Positive and negative
Environment (generally) Positive and negative
Fisheries / aquaculture Positive
Human health Positive
Native fauna Negative

Economic Impact

Top of page

L. cyanellus is problematic for fish management because it often outcompetes native fish.

This species has become an issue in bass-bluegill (Micropterus salmoides-Lepomis macrochirus) population management in the USA. Once introduced into a pond either by natural means (i.e. flooding) or by accident, green sunfish compete with small bass and bluegill; once established they have a tendency to overpopulate and limit food among the other game species present, especially if bass populations decline. They will feed on the eggs and young of bass, reducing bass numbers, and increasing their own numbers and thus the competition (Etnier and Starnes, 2001).

Although L. cyanellus is sometimes deliberately stocked for sport fishing, its high reproductive potential and tolerance for crowding lead to stunted populations which are unattractive to anglers. Precautions should be taken to prevent the introduction of green sunfish into ponds due to their ability to compete with other game fish, small size, and tendency to overpopulate (Etnier and Starnes, 2001).

Experiments on the use of the piscicide antimycin A to remove L. cyanellus from channel catfish (Ictalurus punctatus) production ponds showed that untreated ponds yielded 27.4% fewer catfish than treated ones, and three times as many catfish that were too small for table use (Burress and Luhning, 1969a).

Environmental Impact

Top of page

Impact on Biodiversity

Introduced L. cyanellus compete with and prey on native fish species, reducing their populations, altering their population structure, relative dominance and distribution patterns. Their aggressive nature allows them to compete with larger fish for prey, and prey on eggs and young of other fish species. Olden and Poff (2005), in a study of long-term trends (> 160 years) of fish species distributions in the Lower Colorado River Basin, to identify those native species exhibiting the greatest rates of decline and those non-native species exhibiting the highest rates of spread, found that the green sunfish was amongst the fastest expanding invaders in the basin and the most invasive in terms of negative impacts on native fish communities.

According to Moyle and Nichols (1973, 1974), L. cyanellus was responsible for the local extinction of the California roach (Hesperoleucus symmetricus) in California’s San Joaquin Valley as a result of heavy predation on its young. Lemly (1985) noted that the removal of green sunfish from streams resulted in an increase in the biomass and numbers of most native species. In a study of the North Carolina Piedmont streams, he found that green sunfish preyed on minnows and may have been responsible for the elimination of two cyprinid species in the study area.   

Lohr and Fausch (1996) observed that L. cyanellus can reduce or eliminate plains killifish (Fundulus zebrinus) in isolated pools which may explain why both species rarely co-occur in isolated pools. Dudley and Matter (2000) found that young life phases of the Gila chubb (Gila intermedia) do not occur in downstream sections of the Sabino Creek in Arizona inhabited by green sunfish; abundant numbers of the young chubb were reported in upstream sections of the creek devoid of green sunfish. As the green sunfish feeds on all life stages of the Virgin River chubb, Gila seminuda, it is considered a threat to its existence (USFWS, 2013). According to Karp and Tyus (1990), the green sunfish may compete with and adversely affect the endangered young Colorado pikeminnow Ptychocheilus lucius.

In 2002, non-native L. cyanellus were documented for the first time in O’Donnell Creek, a tributary of the Babocomari River in southern Arizona; as a result, numbers of Sonora sucker (Catastomus insignis) and Gila chubb (Gila intermedia) decreased and longfin dace (Agosia chrysogaster) were extirpated. (Treatment with Antimycin A successfully removed all green sunfish). (Blasius, 2002; Clarkson and Marsh, 2010).

L. cyanellus and other introduced predatory centrarchids are also believed to have played a part in the decline of the California tiger salamander Ambystoma californiense in California (Hayes and Jennings, 1986; Dill and Cordone, 1997) and the Chiricahua leopard frog Rana chiricahuensis [Lithobates chiricahuensis] populations in southeastern Arizona (Rosen et al., 1995).

Threatened Species

Top of page
Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Ambystoma californiense (California tiger salamander)VU (IUCN red list: Vulnerable) VU (IUCN red list: Vulnerable); USA ESA listing as endangered species USA ESA listing as endangered speciesArizonaCompetition - monopolizing resources; Predation; Rapid growthDill and Cordone, 1997; Hayes and Jennings, 1986
Gila intermedia (Gila chub)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesArizonaCompetition - monopolizing resources; Predation; Rapid growthBlasius, 2002; Dudley and Matter, 2000; USFWS, 2013
Gila seminuda (Virgin River chub)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); National list(s) National list(s); USA ESA listing as endangered species USA ESA listing as endangered speciesArizona; Nevada; UtahPredation; Rapid growthUSFWS, 2013
Hesperoleucus symmetricusNational list(s) National list(s)CaliforniaCompetition - monopolizing resources; Predation; Rapid growthMoyle and Nichols, 1973; Moyle and Nichols, 1974; USFWS, 2013
Ptychocheilus lucius (Colorado pikeminnow)No DetailsCompetition - monopolizing resources; Predation; Rapid growthKarp and Tyus, 1990; USFWS, 2013
Rana chiricahuensis (Chiricahua leopard frog)VU (IUCN red list: Vulnerable) VU (IUCN red list: Vulnerable)CaliforniaCompetition - monopolizing resources; Predation; Rapid growthRosen et al., 1995
Xyrauchen texanus (razorback sucker)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesColoradoCompetition - monopolizing resources; Predation; Rapid growthDudley and Matter, 2000; Marsh and Langhorst, 1988; USFWS, 2013

Social Impact

Top of page

Although sometimes deliberately introduced for sport fishing, L. cyanellus often has a detrimental impact on populations of other more desirable species (Etnier and Starnes, 2001).

Risk and Impact Factors

Top of page Invasiveness
  • Invasive in its native range
  • 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
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Tolerant of shade
  • Capable of securing and ingesting a wide range of food
  • Highly mobile locally
  • Long lived
  • Fast growing
  • Has high reproductive potential
  • Gregarious
Impact outcomes
  • Altered trophic level
  • Ecosystem change/ habitat alteration
  • Negatively impacts aquaculture/fisheries
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
  • Negatively impacts trade/international relations
Impact mechanisms
  • Competition - monopolizing resources
  • Hybridization
  • Predation
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Highly likely to be transported internationally deliberately
  • Highly likely to be transported internationally illegally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field

Uses

Top of page

Although L. cyanellus is barely suitable as a "pan-fish", and often thought not to be large enough to interest anglers, it can be exciting to catch; it is sought by recreational anglers in some US states (Wang, 1986; USFWS, 2002; Wang and Reyes, 2008). The fish have a tendency to attack almost any bait and provide a tough fight for anglers, given their size (Etnier and Starnes, 2001).

According to White (1971), the golden colour of Texas golden green sunfish (a colour mutation of L. cyanellus) could make it susceptible to predation and therefore a good forage fish. Biggins (1968) reported that the green sunfish was a major food source of largemouth bass, Micropterus salmoides, grown in a desert impoundment; Turner and Summerfelt (1971) also noted that it was a suitable food source for flathead catfish Pylodictis olivaris culture.

It is easily handled, making it a good bioassay specimen (Carlander, 1977).

Green sunfish are kept as aquarium fish by hobbyists in the USA, and possibly Germany and the Netherlands (Soes et al., 2011).

L. cyanellus has been used to control mosquito larvae in ricefields (Davey and Meisch, 1977; Blaustein and Karban, 1986; Grant et al., 1984).

Uses List

Top of page

Animal feed, fodder, forage

  • Bait/attractant
  • Fodder/animal feed
  • Forage
  • Live feed

Environmental

  • Biological control

General

  • Laboratory use
  • Pet/aquarium trade
  • Sport (hunting, shooting, fishing, racing)
  • Sport fish

Human food and beverage

  • Cured meat
  • Fresh meat
  • Live product for human consumption
  • Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)
  • Whole

Detection and Inspection

Top of page

L. cyanellus is easily identified by its large mouth, white- to orange-margined fins, and large dark spot at the base of the second dorsal fin; it can be distinguished from the bluegill L. macrochirus by its large mouth with the jaw extending to the middle of its eye and the lack of vertical bars. There is also a dark "ear" flap on the gill plate in green sunfish. It can be distinguished from the rock bass Ambloplites rupestris by the presence of 3 instead of 5-8 anal spines; its lack of an orange spot on the gill cover distinguishes it from the pumpkinseed L. gibbosus. Pflieger (1975, 1997) has provided details of a key to the identification of L. gulosus, L. cyanellus, L. microlophus, L. gibbosus, L. megalotis, and L. macrochirus.

Similarities to Other Species/Conditions

Top of page

L. cyanellus is similar to the warmouth, L. gulosus, the longear sunfish L. megalotis, and the bluegill L. macrochirus. It can, however be distinguished from these species by its large mouth, short, rounded pectoral fins, the three spines on its anal fin and its teeth-free pterygoids (Sublette et al., 1990). Unlike in the warmouth, there are no teeth on the tongue of the green sunfish. It only has 3 anal spines while the rock bass Ambloplites rupestris has 5-8 anal spines.

Prevention and Control

Top of page

Prevention

Where L. cyanellus has been introduced as a result of an unauthorized fish introduction, a long-term solution would be through education of the public about the negative consequences of transferring fish to new water bodies. In the short term, legislation that makes unauthorized stocking illegal and the imposition of fines can serve as a deterrent. Where green sunfish are deemed a threat to other species in a system, the authorities may have to resort to removal by chemical or mechanical means. Articles in the popular press and in fishing magazines should highlight examples of illegal fish introductions; education can be done at the local level through news releases emphasizing the problems caused by an unauthorized introduction, or by distributing informational literature. Fisheries biologists should be able to suggest ways to humanely dispose of unwanted pet fish or unused baitfish as an alternative to releasing them into local waters (suggestions can be found at the US Geological Survey’s nonindigenous aquatic species information website, http://nas.er.usgs.gov/taxgroup/fish/)

Control

L. cyanellus is difficult to control once it has become established in ponds. The entire fish population must be eliminated with chemicals or the pond drained to eliminate these fish. Sometimes it is possible to control numbers by continuous trapping, or by destroying spawning areas. The removal of vegetation cover will also expose L. cyanellus to predator fish species.

The use of antimycin A to remove unwanted L. cyanellus from channel catfish production ponds is well-documented (Burress and Luhning, 1969a, b). Green sunfish were effectively and economically controlled in channel catfish (Ictalurus punctatus) ponds on a Mississippi fish farm with antimycin A treatments. A follow-up application at a higher dose further reduced the green sunfish numbers with no apparent effect on yearling catfish. Untreated ponds yielded 27.4% fewer channel catfish than treated ponds and three times as many catfish that were too small for table use. Precise dosing of Antimycin A has also been used to remove green sunfish from live-haul tanks containing channel catfish following transport and prior to stocking of the latter in production ponds (Lloyd, 1987). This is because LC50 values established for green sunfish show they are 45 times more sensitive to antimycin than catfish (Finlayson et al. 2002). Green sunfish treated for 15 min with a range of doses died at concentrations well below those causing mortality in catfish (Finlayson et al., 2002). According to Lloyd (1987), removing green sunfish before stocking in production ponds results in lower costs and obviates the need for treatment of the entire pond at a later date.

A successful example of removal of L. cyanellus from a natural habitat using Antimycin A occurred in 2002, when the species was documented for the first time in O’Donnell Creek, a tributary of the Babocomari River in southern Arizona. As a result of their illegal introduction, numbers of Sonora sucker (Catastomus insignis) and Gila chubb (Gila intermedia) decreased and longfin dace (Agosia chrysogaster) were extirpated. To prevent extirpation of the Sonora sucker and Gila chubb, the authorities chemically renovated O’Donnell Creek in 2002 with Antimycin A to remove L. cyanellus; this led to their successful 100% removal (Blasius, 2002; Clarkson and Marsh, 2010).

A renovation project carried out in 1999 to remove L. cyanellus from Sabino Canyon (Pima County, Arizona) in an attempt to restore populations of Gila chub (Gila intermedia) was considered a success when repeated monitoring showed no signs of L. cyanellus following fish toxin application (Matter et al., 2001).

Gaps in Knowledge/Research Needs

Top of page

A survey of the status of L. cyanellus outside its native range would help to clear the confusion regarding its status where it has been introduced. Further surveys on its distribution in both its native and introduced range and its impact on native species would also be useful.

References

Top of page

Applegate RL, 1966. The use of a bryozoan, Fredericella sultana, as food by sunfish in Bull Shoals Reservoir. Limnology and Oceanography, 11(1):129-130.

Applegate RL; Mullan JW; Morais DI, 1967. Food and growth of six centrarchids from shoreline areas of Bull Shoals reservoir. Proc S. Assoc Game Fish Comm, 20:469-482.

Aquarium Glaser, 2013. Aquarium Glaser. Rodgau, Germany: Aquarium Glaser. http://www.aquarium-glaser.de/en/index.php

Arnold A, 1990. Eingebürgerte Fischarten (Naturalized fish species). Wittenberg, Germany: A. Ziemsen Verlag, 144 pp. [Die Neue Brehm-Bücherei vol. 62.]

Auer NA, 1982. Identification of larval fishes of the Great Lakes Basin with emphasis on the Lake Michigan drainage. Great Lakes Fish Commission. [Great Lakes Fish Commission Special Publication 82-3.]

Bauer RJ, 1988. 1986 Illinois sport fishing survey. Springfield, Illinois, USA: Illinois Department of Conservation [Special Fisheries report No. 53].

Becker GC, 1983. Fishes of Wisconsin. Madison, WI, USA: University of Wisconsin Press, xii + 1052 pp.

Beitinger TL; Magnuson JJ; Neill WH; Shaffer WR, 1975. Behavioral thermoregulation and activity patterns in the green sunfish, Lepomis cyanellus. Animal Behaviour, 23:222-229.

Benson AJ, 1999. Documenting over a century of aquatic introductions in the United States. In: Nonindigenous freshwater organisms - Vectors, Biology and Impacts [ed. by Claudi R, Leach JH] Boca Raton, Florida, USA: Lewis Publishers, 1-31.

Biggins RG, 1968. Centrarchid feeding interactions in a small desert impoundment. Tucson, Arizona, USA: The University of Arizona, 44 pp.

Blasius HB, 2002. Chemical removal of green sunfish (Lepomis cyanellus) from O'Donnell Creek, Arizona. In: Desert Fishes Council 2002 Annual Meeting, San Luis Potosi, SLP, Mexico.

Blaustein L; Karban R, 1986. Mosquitofish-green sunfish interactions in experimental rice fields: effects on mosquito abundance. Proceedings and Papers of the Annual Conference of the California Mosquito and Vector Control Association, 53:64.

Brown GE; Brennan S, 2000. Chemical alarm signals in juvenile green sunfish (Lepomis cyanellus, Centrarchidae). Copeia:1079-1082.

Brunson M; Robinette H, 1983. Winter growth of bluegills and bluegill x green sunfish hybrids in Mississippi. Proceedings of the Annual Conference of the Southeastern Association of Fish and Wildlife Agencies, 37:343-347.

Burress RM; Luhning CW, 1969. Field trials of antimycin as a selective toxicant in channel catfish ponds. La Crosse, Wisconsin, USA: U.S. Bureau of Sport Fisheries and Wildlife, 12 pp. [Investigations in Fish Control (25).]

Burress RM; Luhning CW, 1969. Use of antimycin for selective thinning of sunfish populations in ponds. La Crosse, Wisconsin, USA: U.S. Bureau of Sport Fisheries and Wildlife, 12 pp. [Investigations in Fish Control (28).]

Carlander KD, 1977. Handbook of Freshwater Fishery Biology, volume two: life history data on Centrarchid fishes of the United States and Canada. Ames, Iowa, USA: Iowa State University Press, 431 pp.

Childers WF, 1967. Hybridization of four species of sunfishes (Centrarchidae). Illinois Natural History Survey Bulletin, 29:159-214.

Clarkson RW; Marsh PC, 2010. Effectiveness of the barrier-and-renovate approach to recovery of warmwater native fishes in the Gila River Basin. In: Proceedings of the Colorado River Basin Science and Resources Management Symposium, Scottsdale, Arizona, USA, 18-20 November 2008 [ed. by Mellis, T. S. \Hamill, J. F. \Coggins Jr, L. G. \Grams, P. E. \Kennedy, T. A. \Kubly, D. M. \Ralston, B. E.]. [US Geological Survey Scientific Investigation Report, 2010-5135.]

Cortemeglia C; Beitinger TL, 2008. Temperature tolerance of green sunfishes Lepomis cyanellus. Texas Journal of Science, 60(3):1-5.

Coutant CC, 1977. Compilation of temperature preference data. Journal of the Fisheries Research Board of Canada, 34:739-745.

Crossman EJ, 1976. Quetico fishes. Toronto, Ontario, Canada: Royal Ontario Museum. [Miscellaneous Life Science Publications of the Royal Ontario Museum.]

Davey RB; Meisch MV, 1977. Control of dark rice field mosquito larvae, Psorophora columbiae by mosquitofish, Gambusia affinis and green sunfish, Lepomis cyanellus, in Arkansas rice fields. Mosquito News, 37(2):258-262.

Dill WA; Cordone AJ, 1997. History and status of introduced fishes in California. Fish Bulletin, California Department of Fish and Game, 178:1-414.

Dudley RK; Matter WJ, 2000. Effects of small green sunfish (Lepomis cyanellus) on recruitment of Gila chubb (Gila intermedia) in Sabino Creek, Arizona. The Southwestern Naturalist, 45(1):24-29.

Dupre HK; Huner JV, 1984. Propagation of black bass, sunfishes, tilapias, eels, and hobby fishes. In: Dupree HK, Huner JV, eds. Third Report to the Fish Farmers. Washington DC, USA: US Fish and Wildlife Service, 119-135.

Elvira B, 2001. Identification of non-native freshwater fishes established in Europe and assessment of their potential threats to the biological diversity. Strasbourg, France: Convention on the Conservation of European Wildlife and Natural Habitats, 35 pp. https://wcd.coe.int/com.instranet.InstraServlet?command=com.instranet.CmdBlobGet&InstranetImage=1338217&SecMode=1&DocId=1464096&Usage=2

Engelhardt T, 1985. Production of hybrid sunfish. In: Proceedings of the 1985 Texas Fish Farming Conference, College Station, Texas, USA.

Etnier D; Starnes W, 2001. The Fishes of Tennessee. Knoxville, Tennessee, USA: The University of Tennessee Press.

Finlayson BJ; Schnick RA; Cailteux RL; DeMong L; Horton WD; McClay W; Thompson CW, 2002. Assessment of antimycin A use in fisheries and its potential for reregistration. Fisheries (Bethesda), 27(6):10-18.

Froese R; Pauly D, 2013. FishBase. http://www.fishbase.org

Geiter O; Homma S; Kinzelbach R, 2002. Bestandsaufnahme und Bewertung von Neozoen in Deutschland (Stock-taking and evaluation of neozoa in Germany). Berlin, Germany: Umweltbundesamt, 290 pp. [Texte 25/02.]

Grant CD; Combs JC; Coykendall Rl; Lusk EE; Washino RK; Blaustein L; Karban R, 1984. Continuing studies of Gambusia affinis, Lepomis cyanellus and their interactive effects on mosquito abundance in experimental rice plots. Proceedings and Papers of the Fifty-second Annual Conference of the California Mosquito and Vector Control Association, 29 January-1 February, 1984, Long Beach, California, USA, 120 pp.

Hallam JC, 1959. Habitat and associated fauna of four fish in Ontario streams. Journal of the Fisheries Board of Canada 16, 16:147-173.

Halos S; Vina AG La; Lim MS; Caleda MJ, 2004. Developing the National Biosafety Framework for the Philippines. Quezon City, Philippines: Department of Environment and Natural Resources-Protected Areas and Wildlife Bureau, 164 pp.

Hasler AD; Wibly WJ, 1958. The return of displaced largemouth bass and green sunfish to a "home" area. Ecology, 39:289-293.

Hayes MP; Jennings MR, 1986. Decline of Ranid Frog Species in Western North America: Are Bullfrogs (Rana catesbiana) Responsible? Journal of Herpetology, 20:490-509.

Heidinger RC, 1975. Growth of hybrid sunfishes and channel catfish at low temperatures. Transactions of the American Fisheries Society, 104:333-334.

Hubbs CL; Cooper GP, 1935. Age and growth of the long-eared and the green sunfishes in Michigan. Papers of the Michigan Academy of Science, Arts and Letters, 20:669-696.

Hunter J, 1963. The reproductive behaviour of the green sunfish, Lepomis cyanellus. Zoologica, Scientific Contributions of the New York Zoological Society, 48:13-23.

Husemann M; Ray JW; King RS; Hooser EA; Danley PD, 2012. Comparative biogeography reveals differences in population genetic structure of five species of stream fishes. Biol. Linn. Soc. Lond, 107(4):867-885.

Jackson PBN, 1976. Water resources and freshwater fishes in southern Africa. In: Resources of Southern Africa, today and tomorrow [ed. by Baker, G.]. Johannesburg, South Africa: Association of Scientific and Technical Societies of South Africa, 196-207.

Juliano RO; Guerrero R III; Ronquillo I, 1989. The introduction of exotic aquatic species in the Philippines. In: De Silva SS, ed. Proceedings of the Workshop on Introduction of Exotic Aquatic Organisms in Asia: The Asian Fisheries Society, 83-90.

Karp CA; Tyus HM, 1990. Behavioral interactions between young Colorado squawfish and six fish species. Copeia, 1:25-34.

Kaya CM, 1973. Effects of temperature and photoperiod on seasonal regression of gonads of Green Sunfish, Lepomis cyanellus. Copeia, 2:369-373.

Kilby JD, 1955. The fishes of two gulf coast marsh areas of Florida. Tulane Studies in Zoology, 2:175-247.

Kurzawski K; Heidinger R, 1982. The cyclic stocking of parentals in a farm pond to produce a population of male bluegill x female green sunfish F1 hybrids and male redear sunfish x female green sunfish F1 hybrids. North American Journal of Fisheries Management, 2:188-192.

Lemly AD, 1985. Suppression of native fish populations by green sunfish in first-order streams of Piedmont North California. Transactions of the American Fisheries Society, 114:705-712.

Lever C, 1996. Naturalized fishes of the world. California, USA: Academic Press, 408 pp.

Litvak MK; Mandrak NE, 1999. Baitfish trade as a vector of aquatic introductions. In: Nonindigenous freshwater organisms: vectors, biology and impacts [ed. by Claudi, R. \Leach, J. H.]. Boca Raton, Florida, USA: Lewis Publishers, 163-180.

Lloyd SW, 1987. Use of antimycin A in live-haul tanks to remove scaled fish from fingerling channel catfish populations. The Progressive Fish-Culturist, 49:131-133.

Lohr SC; Fausch KD, 1996. Effects of green sunfish (Lepomis cyanellus) predation on survival and habitat use of plains killifish (Fundulus zebrinus). The Southwestern Naturalist, 41(2):155-160.

Ma X; Bangxi X; Yindong W; Mingxue W, 2003. Intentionally introduced and transferred fishes in China's inland waters. Asian Fisheries Science, 16:279-290.

Maitland PS, 1977. Elseviers gids voor de zoelwatervissen ([English title not available]). Amsterdam, Netherlands: Elsevier.

Marsh PC, 2010. The Colorado River basin, its Native and Non-native Fishes, and the Potential for Application of Genetic Biocontrol to the Conservation of an Imperiled Fauna. In: International Symposium on Genetic Biocontrol of Invasive Fish, Minnesota, USA, 21-24 June 2010. http://www.seagrant.umn.edu/downloads/biocontrol/Colorado_River_Basin_and_Their_Invasives-Paul_Marsh.pdf

Marsh PC; Langhorst DR, 1988. Feeding and fate of wild larval razorback sucker. Environmental Biology of Fishes 21, 21:59-67.

Matter W; Romero S; Blasius H, 2001. Sabino Canyon. In: Reducing the exotic aquatic species threat in Pima County. Proceedings, 27 February 2001. Tucson, Arizona, USA: Pima Association of Governments, 9-11.

McKechnie RJ; Tharratt RC, 1966. Green sunfish. In: Inland Fisheries Management [ed. by Calhoun, A.]. Sacramento, California, USA: California Department of Fish and Game, 399-402.

Meredith GN; Houston JJ, 1988. Status of the green sunfish, Lepomis cyanellus in Canada. Canadian Field Naturalist, 102(2):270-276.

Meyer FA, 1970. Development of some larval centrarchids. Progressive Fish Culturist, 32:130-136.

Mills EL; Leach JH; Carlton JT; Secor CL, 1993. Exotic species in the Great Lakes: A history of biotic crises and anthropogenic introductions. Journal of Great Lakes Research, 19(1):1-54.

Minckley WL, 1973. Fishes of Arizona. Arizona Fish and Game Department. Phoenix, AZ, USA: Sims Printing Company, Inc.

Minckley WL; Clarkson RW, 1979. Fishes. In: Resource inventory for the Gila River complex, eastern Arizona [ed. by Minckley, W. L. \Summerfeld, M. R.]. Tempe, Arizona, USA: Arizona State University, 510-531. [Final Report to U.S. Bureau of Land Management, Safford District, Contract No. YA-512-CT6-216.]

Mischke CC; Morris JE, 1997. Out-of-season spawning of sunfish Lepomis spp. in the laboratory. Progressive Fish-Culturist, 59(4):297-302.

Mito T; Uesugi T, 2004. Invasive alien species in Japan: the status quo and new regulations for prevention of their adverse effects. Global Environmental Research, 8(2):171-191.

Moore WG, 1942. Field studies on the oxygen requirements of certain freshwater fishes. Ecology, 23:319-329.

Morris JE; Mischke CC, 2000. A white paper on the status and needs of sunfish aquaculture in the North Central Region. North Central Region Aquaculture Center, Michigan, USA. 17 p.

Moyle PB, 2002. Inland fishes of California. Berkeley, USA: University of California Press, 502 pp.

Moyle PB; Nichols R, 1973. Ecology of some native and introduced fishes of the Sierra-Nevada foothills in central California. Copeia:478-490.

Moyle PB; Nichols R, 1974. Decline of native fish fauna of the Sierra-Nevada foothills, central California. American Midland Naturalist, 92:72-83.

NatureServe, 2013. NatureServe Explorer: An online encyclopedia of life. Arlington, Virginia, USA: NatureServe. http://www.natureserve.org/explorer/

Nehring S, 2005. International shipping - risk for aquatic biodiversity in Germany. Neobiota, 6:125-143.

NOBANIS, 2005. Lepomis cyanellus. http://www.nobanis.org/NationalInfo.asp?countryID=DE&taxaID=5587

Olden JD; Poff NL, 2005. Long-term trends of native and non-native fish faunas in the American Southwest. Animal Biodiversity and Conservation, 28:75-89.

Page LM; Burr BM, 1991. A field guide to freshwater fishes of North America north of Mexico. Houghton Mifflin Company, Boston, 432 pp.

Petit GD, 1973. Effects of dissolved oxygen on survival and behaviour of selected fishes of western Lake Erie. Ohio Biological Survey Bulletin, 4(4):1-76.

Pflieger W, 1975. Sunfishes. In: The fishes of Missouri. Jefferson City, Missouri, USA: Missouri Department of Conservation, 249-275.

Pflieger WL, 1997. The fishes of Missouri. Missouri Department of Conservation, Jefferson City, MO. 372 pp.

Proffitt MA; Benda RS, 1971. Growth and movement of fishes, and distribution of invertebrates, related to a heated discharge into the White River at Petersburg, Indiana. Bloomington, Indiana, USA: Indiana University Water Resources Department, 94 pp. [Indiana University Water Resources Department Investigation 5.]

Rahel FJ; Thel LA, 2004. Plains killifish (Fundulus zebrinus): A technical conservation assessment. USA: USDA Forest Service, Rocky Mountain Region, 49 pp. http://www.fs.fed.us/v2/projects/scp/assessments/plainskillifish.pdf

Rosen PC; Schwalbe CR; Parizek DAJ; Holm PA; Lowe CH, 1995. Introduced aquatic vertebrates in the Chiricahua region: effects on declining ranid frogs. In: Biodiversity and Management of the Madrean Archipelago: The Sky Islands of Southwestern United States and Northwestern Mexico [ed. by DeBano, L. F. \Gottfried, G. J. \Hamre, R. H. \Edminster, C. B. \Folliott, P. F. \Ortega-Rubio, A.]. Fort Collins, Colorado, USA: Rocky Mountain Forest and Range Experiment Station, 251-261.

Scott WB; Crossman EJ, 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada, Bulletin, No. 184:966 pp.

Sigler WF; Miller RR, 1963. Fishes of Utah. Utah, USA: Utah State Department of Fish and Game, 203 pp.

Soes DM; Cooke SJ; Kleef HH van; Broeckx PB; Veenvliet P, 2011. A risk analysis of sunfishes (Centrarchidae) and pygmy sunfishes (Elassomatidae) in The Netherlands. Netherlands: Bureau Waardenburg Bv, 110 pp.

Sprugel G Jr, 1955. The growth of green sunfish (Lepomis cyanellus) in Little Wall Lake, Iowa. Iowa State College Journal of Science, 29:707-719.

Sterba G, 1959. Süsswasserfische aus aller Welt (Freshwater fishes of the whole world). Zimmer & Herzog Verlag.

Stroud RH, 1967. Water quality criteria to protect aquatic life: a summary. In: A Symposium on Water Quality Criteria to Protect Aquatic Life. 33-37. [American Fisheries Society Special Publication 4.]

Stuber RJ; Gebhart G; Maughan OE, 1982. Habitat suitability models: green sunfish. Washington, DC, USA: US Department of the Interior, Fish and Wildlife Service, 32 pp. [FWS/OBS-82/10.16.] http://www.nwrc.usgs.gov/wdb/pub/hsi/hsi-015.pdf

Sublette JE; Hatch MD; Sublette M, 1990. The fishes of New Mexico. Albuquerque, New Mexico: University New Mexico Press, 393 pp.

Taubert BD, 1977. Early morphological development of the green sunfish, Lepomis cyanellus, and its separation from other larval Lepomis species. Transactions of the American Fisheries Society, 106(5):445-448.

Tebo LB; McCoy EG, 1964. Effects of seawater concentration on the reproduction of largemouth bass and bluegills. Progressive Fish Culturist, 6(3):99-106.

Thomas C; Bonner TH; Whiteside BG, 2007. Freshwater Fishes of Texas: A Field Guide. Texas, USA: Texas A&M Press.

Tidwell JH; Webster CD, 1993. Effects of stocking density and dietary protein on green sunfish (Lepomis cyanellus) bluegill (L. macrochirus) hybrids overwintered in ponds. Aquaculture, 113(1/2):83-89; 18 ref.

Tidwell JH; Webster CD; Clark JA, 1992. Growth, feed conversion, and protein utilization of female green sunfish male bluegill hybrids fed isocaloric diets with different protein levels. Progressive Fish-Culturist, 54(4):234-239; 26 ref.

Tomelleri JR; Eberle ME, 1990. Fishes of the Central United States. Lawrence, Kansas, USA: University of Kansas Press, 226 pp.

Trautman MB, 1957. The fishes of Ohio with illustrated keys. Columbus, Ohio, USA: Ohio State University Press.

Turner PR; Summerfelt RC, 1971. Food habits of adult flathead catfish, Pylodictus olivaris (Rafinesque), in Oklahoma reservoirs. In: Proceedings of the Annual Conference of the Southeastern Association of Game and Fisheries Commission 24, 1970. 387-401.

Ultsch GR, 1978. Oxygen consumption as a function of pH in three species of freshwater fishes. Copeia:272-279.

USDA, 2012. Review and assessment of programs for invasive species management in the Southwestern region, 2012. USA: USDA Forest Service, Southwestern Region, 33 pp. [TR-R3-16-25.]

USFWS (US Fish and Wildlife Service), 2002. 2001 National Survey of Fishing, Hunting, and Wildlife-Associated Recreation. Washington, DC, USA: US Department of the Interior, 170 pp.

USFWS (US Fish and Wildlife Service), 2013. Environmental Conservation Online System. http://ecos.fws.gov

USGS, 2013. USGS Nonindigenous Aquatic Species Database. Gainesville, Florida, USA: United States Geological Survey. http://nas.er.usgs.gov

Wang JCS, 1986. Fishes of the Sacramento-San Joaquin Estuary and Adjacent Waters, California: A Guide to the Early Life Histories. Berkeley, USA: Digital Library Project, ix + 680 pp. [Interagency Ecological Program Technical Report No. 9].

Wang JCS; Reyes RC, 2008. Early Life Stages and Life Histories of Centrarchids in the Sacramento-San Joaquin River Delta System, California. Denver, Colorado, USA: US Department of the Interior, Bureau of Reclamation, 112 pp. [Tracy Fish Facilities Study Series Vol 42.]

Wang Ning; Hayward RS; Noltie DB, 1998. Effect of feeding frequency on food consumption, growth, size variation, and feeding pattern of age-0 hybrid sunfish. Aquaculture, 165(3/4):261-267; 17 ref.

Webster CD; Tidwell JH, 2002. Centrarchids: hybrid bluegill (Lepomis cyanellus × Lepomis macrochirus). In: Nutrient requirements and feeding of finfish for aquaculture [ed. by Webster, C. D.\Lim, C.]. Wallingford, UK: CABI Publishing, 381-387. http://www.cabi.org/CABeBooks/default.aspx?site=107&page=45&LoadModule=PDFHier&BookID=86

Webster CD; Tiu LG; Tidwell JH, 1997. Growth and body composition of juvenile hybrid bluegill Lepomis cyanellus x L. macrochirus fed practical diets containing various percentages of protein. Journal of the World Aquaculture Society, 28(3):230-240; 32 ref.

Welcomme RL, 1988. International introductions of inland aquatic species. FAO Fisheries Technical Paper, No. 294:x + 318 pp.

White GE, 1971. The Texas golden green: a color mutation of the green sunfish. Progressive Fish Culturist, 33(3):155.

Wills PS; Paret JM; Sheenhan PJ, 1994. Pressure induced triploidy in hybrid Lepomis. Journal of the World Aquaculture Society, 25(4):507-511.

Yamamoto MN, 1992. Occurrence, distribution and abundance of accidentally introduced freshwater aquatic organisms in Hawaii. State of Hawaii, Federal Aid in Sportfish Restoration, Dingell-Johnson JOR. Freshwater Fisheries Research and Surveys, Project No. F-14-R-16.

Links to Websites

Top of page
WebsiteURLComment
Aquarium-Glaserhttp://www.aquarium-glaser.de/en/index.php
European Network on Invasive Alien Species (NOBANIS)http://www.nobanis.org/
FishBasehttp://www.fishbase.org/search.php
United States Geological Survey Nonindigenous Aquatic Specieshttp://nas.er.usgs.gov/taxgroup/fish/

Contributors

Top of page

Main Author
Uma Sabapathy Allen
Human Sciences, CAB International, Wallingford, Oxon, OX10 8DE, UK

Distribution Maps

Top of page
You can pan and zoom the map
Save map