Gambusia holbrooki (eastern mosquitofish)

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Adult fish

Adult Gambusia holbrooki, the Eastern mosquito fish, from Australia.

Stephen L. Doggett/Dept. of Medical Entomology, Westmead Hospital, Sydney, Australia

Identity

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Preferred Scientific Name

Gambusia holbrooki Girard, 1859

Preferred Common Name

eastern mosquitofish

Other Scientific Names

Gambusia affinis holbrocki (Girard, 1859)
Gambusia affinis holbrooki (Girard, 1859)
Gambusia holbrookii Girard, 1859
Gambusia patruelis holbrooki (Girard, 1859)
Heterandria holbrooki (Girard, 1859)
Heterandria uninotata (non Poey, 1860)
Schizophallus holbrooki (Girard, 1859)
Zygonectes atrilatus Jordan & Brayton, 1878

International Common Names

English: mosquito fish; mosquitofish
Spanish: gambusino

Local Common Names

Albania: barkaleci pikalosh
Australia: eastern gambusia
Iran: gambusia
Italy: gambusia
Portugal: peixe-mosquito
Romania: gambuzie
Sweden: östlig moskitfisk
USA: eastern topminnow

Summary of Invasiveness

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The eastern mosquitofish Gambusia holbrooki is an exotic fish, which is now widespread around the globe and is known to adversely affect native fish through competition and/or predation. G. holbrooki is known to impact upon native fish via competition with similar sized species, predation upon the fry and eggs of native fish, and by attacking all sized fish by aggressive fin-nipping, thereby leaving them susceptible to disease (Arthington, 1991). G. holbrooki can rapidly increase in population size due to its rapid maturation to breeding age (four weeks in summer) and high survival rate of young (Milton and Arthington, 1983; Lloyd et al., 1986; Lloyd, 1990).

Taxonomic Tree

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Domain: Eukaryota
Kingdom: Metazoa
Phylum: Chordata
Subphylum: Vertebrata
Class: Actinopterygii
Order: Cyprinodontiformes
Family: Poeciliidae
Genus: Gambusia
Species: Gambusia holbrooki

Notes on Taxonomy and Nomenclature

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This datasheet focuses upon Gambusia holbrooki, although much of the biology is confused between the previous subspecies and now congeners, G. holbrooki and G. affinis. Therefore, unless specified, reference to mosquitofish refers to both species. Data for both species is also presented because both species were both regarded as Gambusia affinis prior to 1988 and even after this date workers often referred Gambusia affinis as a generic name for both species.

The genus Gambusia comprises about 30 species divided into four sub-genera: Orthophallus, Arthrophallus, Heterophallina,and Gambusia. Other gambusine species rarely coexist with G. affinis, but when they do they are usually ecologically segregated. However, this species will hybridise with some of its congeners, including G. heterochir, G. nobilis, and G. marshi (Hubbs, 1955; Rosen and Bailey, 1963; Hubbs and Perden, 1969; Schoenherr, 1974; Yardley and Hubbs, 1976).

Wotten et al. (1988) formally recognised two species - G. holbrooki and G. affinis - which were previously regarded as sub-species of G. affinis (G. a. holbrooki and G. a. affinis). These species are different in a number of meristic characteristics but are definitively differentiated by the morphological differences of the gonopodium (male anal fin) (see Lloyd, 1990c). Work by Lloyd and Tomasov (1985) showed that the Gambusia in Australia are all G. holbrooki.

Description

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The male G. holbrooki is about 35 mm standard length whereas the female is larger (up to 60 mm) with a deeper body, the anal fin unmodified and when pregnant, a gravid spot is visible just above the vent (Lloyd, 1987). The fish are mostly translucent grey with a bluish sheen on their sides with a silver belly (Lloyd, 1987). The fins are colourless, with transverse rows of black spots. Some male mosquitofish have irregular black blotching, though some largely melanistic male individuals exist but are uncommon in their native range (Sterba, 1962) and are absent from Australia (Lloyd, 1987). On the male, the anal fin is modified to form a long, thin intromitent organ, the gonopodium, used for sperm transfer (Lloyd, 1990c). The body is slightly compressed with a large and flattened head. The eyes are large, and the mouth is small and terminal (Lloyd, 1987).

Distribution

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Mosquitofish have become the most widely distributed freshwater teleosts in the world (Krumholz, 1948) mainly through deliberate human introductions. The native range of G. affinis is throughout the Mississippi Basin and the tributaries to the northern Gulf of Mexico; G. holbrooki ranges from New Jersey in the east to the Gulf of Mexico and northern Florida.

The species (both G. affinis and G. holbrooki) have been introduced across the USA and much of southern Canada, parts of South America, Australia, New Zealand, Papua New Guinea, Indonesia, Many Pacific Islands, SE Asia, China, Japan, India, throughout Europe and the Middle East, South Africa, and parts of Northern Africa (Lloyd, 1987).

Because G. affinis and G. holbrooki were treated as subspecies of G. affinis prior to 1988, it is not always possible to discern which species is referred to in earlier literature. It was confirmed by Pyke (2005) that records across Australia were of G. holbrooki.

Distribution Table

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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.

Last updated: 10 Jan 2020

Continent/Country/Region	Distribution	Origin	First Reported	Invasive	Reference	Notes
Africa
Central African Republic	Present	Introduced	1958	Invasive	Froese and Pauly (2008)
Congo, Democratic Republic of the	Present				Alemadi and Jenkins (2007); CABI (Undated)
Côte d'Ivoire	Present	Introduced		Invasive	Froese and Pauly (2008)
Egypt	Present	Introduced	1929	Invasive	Motabar (1978)
Ethiopia	Present	Introduced	1938	Invasive	Froese and Pauly (2008)
Ghana	Present	Introduced		Invasive	Froese and Pauly (2008)
Kenya	Present	Introduced		Invasive	Froese and Pauly (2008)
Madagascar	Present	Introduced	1929	Invasive	Froese and Pauly (2008)
Mauritius	Present	Introduced		Invasive	Froese and Pauly (2008)
-Rodrigues	Present	Introduced		Invasive	Froese and Pauly (2008)
Morocco	Present	Introduced	1929	Invasive	Froese and Pauly (2008); CABI (Undated)
Réunion	Present	Introduced		Invasive	Froese and Pauly (2008)
South Africa	Present	Introduced	1936	Invasive	Fowler (1970); Froese and Pauly (2008)
Sudan	Present	Introduced	1929	Invasive	Motabar (1978)
Zambia	Present	Introduced	1940		Froese and Pauly (2008)	Not established
Zimbabwe	Present	Introduced	1925	Invasive	Froese and Pauly (2008)
Asia
Afghanistan	Present	Introduced	1970	Invasive	Motabar (1978)
Armenia	Present	Introduced		Invasive	Froese and Pauly (2008)
Bangladesh	Present	Introduced		Invasive	Froese and Pauly (2008)
Cambodia	Present	Introduced		Invasive	Froese and Pauly (2008)
China	Present	Introduced	1927	Invasive	Fowler (1970); Froese and Pauly (2008)
Georgia	Present	Introduced	1925	Invasive	Froese and Pauly (2008)
Hong Kong	Present	Introduced		Invasive	Froese and Pauly (2008)
India	Present	Introduced		Invasive	Das and Rampal (1966); Froese and Pauly (2008)
-Jammu and Kashmir	Present	Introduced		Invasive	Froese and Pauly (2008)
Indonesia	Present	Introduced	1929		Froese and Pauly (2008)	Not established
-Irian Jaya	Present	Introduced	1930	Invasive	Glucksman and West (1976)
Iran	Present	Introduced	1928	Invasive	Motabar (1978)
Iraq	Present	Introduced	1928	Invasive	CABI (Undated)	Original citation: Al-Daham et al. (1977)
Israel	Present	Introduced	1924	Invasive	Froese and Pauly (2008)
Japan	Present	Introduced	1916	Invasive	Sasa and Kurihara (1980)
Jordan	Present	Introduced	1930	Invasive	Froese and Pauly (2008)
Kazakhstan	Present	Introduced	1934	Invasive	Froese and Pauly (2008)
Laos	Present	Introduced		Invasive	Froese and Pauly (2008)
Lebanon	Present	Introduced		Invasive	Froese and Pauly (2008)
Malaysia	Present	Introduced		Invasive	Froese and Pauly (2008)
Myanmar	Present	Introduced		Invasive	Froese and Pauly (2008)
Pakistan	Present	Introduced		Invasive	Froese and Pauly (2008)
Philippines	Present	Introduced	1913	Invasive	Seale (1917)
Saudi Arabia	Present	Introduced		Invasive	Froese and Pauly (2008)
Singapore	Present	Introduced		Invasive	Froese and Pauly (2008)
Sri Lanka	Present	Introduced		Invasive	Froese and Pauly (2008)
Syria	Present	Introduced		Invasive	CABI (Undated)	Original citation: Gerberich and Laird (1968)
Taiwan	Present	Introduced	1911	Invasive	Sasa and Kurihara (1980)
Tajikistan	Present	Introduced		Invasive	Froese and Pauly (2008)
Thailand	Present	Introduced	1919	Invasive	Banpot (2004)
Turkey	Present	Introduced	1920	Invasive	Froese and Pauly (2008)
Turkmenistan	Present	Introduced		Invasive	Froese and Pauly (2008)
United Arab Emirates	Present	Introduced		Invasive	Froese and Pauly (2008)
Uzbekistan	Present	Introduced	1930	Invasive	Froese and Pauly (2008)
Vietnam	Present	Introduced		Invasive	Froese and Pauly (2008)
Yemen	Present	Introduced		Invasive	Froese and Pauly (2008)
Europe
Albania	Present	Introduced		Invasive	Froese and Pauly (2008)
Austria	Present	Introduced		Invasive	Fowler (1970)
Bulgaria	Present	Introduced	1924	Invasive	Froese and Pauly (2008)
Cyprus	Present	Introduced	1926	Invasive	CABI (Undated)	Original citation: Gerberich and Laird (1968)
Federal Republic of Yugoslavia	Present	Introduced	1924	Invasive	CABI (Undated)	Original citation: Gerberich and Laird (1968)
France	Present	Introduced	1924	Invasive	Froese and Pauly (2008)
-Corsica	Present	Introduced	1924	Invasive	CABI (Undated)	Original citation: Gerberich and Laird (1968)
Germany	Present	Introduced	1921	Invasive	Fowler (1970)
Greece	Present	Introduced	1928	Invasive	Motabar (1978)
-Crete	Present				Alemadi and Jenkins (2007); CABI (Undated)
Hungary	Present	Introduced	1937	Invasive	Froese and Pauly (2008)
Italy	Present	Introduced	1922	Invasive	Krumholz (1948)
Portugal	Present	Introduced	1921	Invasive	Froese and Pauly (2008)
Romania	Present	Introduced	1921	Invasive	Froese and Pauly (2008)
Russia	Present	Introduced	1925	Invasive	Motabar (1978)
Spain	Present	Introduced			Froese and Pauly (2004)
-Canary Islands	Present	Introduced	1943	Invasive	Krumholz (1948)
Ukraine	Present	Introduced		Invasive	Froese and Pauly (2008)
North America
Canada	Present	Introduced	1924	Invasive	McAllister (1969)
Haiti	Present	Introduced	1983	Invasive	Froese and Pauly (2008)
Mexico	Present	Introduced	1931	Invasive	Froese and Pauly (2008); CABI (Undated)
Puerto Rico	Present	Introduced	1914	Invasive	Froese and Pauly (2008)
United States	Present	Native			Froese and Pauly (2004)
-Florida	Present				Nordlie (2000)
-Hawaii	Present	Introduced	1905	Invasive	Seale (1905)
-Michigan	Present	Introduced	1941	Invasive	Krumholz (1948)
-Utah	Present	Introduced	1927	Invasive	Rees (1934)
Oceania
American Samoa	Present	Introduced		Invasive	Froese and Pauly (2008)
Australia	Present	Introduced			Froese and Pauly (2004); Pyke (2005)
-New South Wales	Present, Widespread	Introduced	1926	Invasive	Wilson (1960)
-Northern Territory	Present, Localized	Introduced	1940	Invasive	Lloyd (1987);
-Queensland	Present, Widespread	Introduced	1925	Invasive	Wilson (1960)
-South Australia	Present, Widespread	Introduced	1926	Invasive	Lloyd (1987)
-Tasmania	Present, Localized	Introduced		Invasive	Keane and Francisco (2004)	First reported: 1990s
-Victoria	Present, Widespread	Introduced	1926	Invasive	Lloyd (1987)
-Western Australia	Present, Localized	Introduced	1934	Invasive	Lloyd (1987)
Cook Islands	Present	Introduced		Invasive	Froese and Pauly (2008)
Federated States of Micronesia	Present	Introduced		Invasive	Froese and Pauly (2008)
Fiji	Present	Introduced	1930	Invasive	Froese and Pauly (2008)
French Polynesia	Present	Introduced		Invasive	Froese and Pauly (2008)
Guam	Present	Introduced		Invasive	Froese and Pauly (2008)
Kiribati	Present	Introduced		Invasive	Froese and Pauly (2008)
Marshall Islands	Present	Introduced		Invasive	Froese and Pauly (2008)
New Zealand	Present	Introduced	1930	Invasive	Allen (1956)
Northern Mariana Islands	Present	Introduced		Invasive	Froese and Pauly (2008)
Papua New Guinea	Present	Introduced	1930	Invasive	Glucksman and West (1976)
Samoa	Present	Introduced		Invasive	Froese and Pauly (2008)
Solomon Islands	Present	Introduced	1930	Invasive	Glucksman and West (1976)
South America
Argentina	Present	Introduced	1943	Invasive	Froese and Pauly (2008)
Bolivia	Present	Introduced		Invasive	Froese and Pauly (2008)
Chile	Present	Introduced	1937	Invasive	Froese and Pauly (2008)
Peru	Present	Introduced	1940	Invasive	Froese and Pauly (2008)

History of Introduction and Spread

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Mosquitofish have become the most widely distributed freshwater teleosts in the world (Krumholz, 1948) mainly through deliberate human introductions (e.g. Lintermans, 2004). Throughout the world, it has been widely distributed to aid mosquito control in rice paddies and natural waters (Krumholz, 1948; Lloyd et al., 1986; Arthington and Lloyd, 1989). Worldwide introduction of Gambusia has occurred since the first introduction into Hawaii in 1905 (Krumholz, 1948).

It is now known that G. holbrooki was introduced to Mediterranean Europe and Australia, whereas G. affinis was introduced outside its natural range to the western United States, Hawaii, and countries in Africa (Rehage and Sih, 2004).

G. holbrooki were first introduced to Brisbane in 1925 and Sydney the following year (Wilson, 1960). The distribution of the Australian populations has continued to expand through new invasions (such as Northern Tasmania) or in filling of locations within catchments. Gambusia was actively introduced by health authorities to new locations in Australia until the 1990s (Lloyd, 1987).

Only G. holbrooki has been identified in the current field records and museum collections that exist in Australia (Lloyd and Tomasov, 1985). Stocks of mosquitofish have been found in many isolated water bodies in central Australia, suggesting that natural events such as floods aid their dispersal (Lloyd, 1987; Chapman and Warburton, 2006). However, it is human activity that has most aided the dispersal of G. holbrooki, largely through projects aimed at mosquito control on a local, national and global scale (Arthington et al., 1983).

Risk of Introduction

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G. holbrooki is highly invasive because of its ability to move and colonise new habitats, its high fecundity, high survival of juveniles and rapid population growth (Moyle and Light, 1996; Williamson and Fitter, 1996; Alemadi and Jenkins, 2007). Mosquitofish can disperse through waters as shallow as 3 mm, which is only half of average body depth (Alemadi and Jenkins, 2007) and uses drains and natural channels to disperse between water bodies. Rehage and Sih (2004) link dispersal behaviour to the ‘invasiveness’ of a species, with G. affinis dispersing faster than G. holbrooki due to the movement of females into new territories. However, other important characteristics, such as fecundity, and maximum population growth rates suggest that G. holbrooki is ’a superior invader‘ (Rehage and Sih, 2004). Eastern mosquitofish can rapidly increase in population size due to their rapid maturation to breeding age (four weeks in summer) and high survival rate of young (Milton and Arthington, 1983; Lloyd et al., 1986; Lloyd, 1990a).

Habitat

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The native habitat of mosquitofish is the lowland ponds, lakes and streams of southern USA (Casterlin and Reynolds, 1977). Mosquitofish are particularly adapted to exploiting inundated floodplains (Ross and Baker, 1983). G. holbrooki is abundant in near-shore environments, close to dense vegetation, and prefers sluggish waters to running water (Lloyd, 1987). Other habitat preferences include: shallow water, dark-coloured substrata, and subsurface vegetation (providing lateral rather than vertical concealment) (Casterlin and Reynolds, 1977; Arthington and Marshall, 1999). They are very adaptable and will live in almost aquatic habitats from fresh to hyper-saline, cold temperate to tropical waters (and artificially heated waters), inland, coastal and estuarine waters, and both still and slow-flowing waters. The species does seem to be poorly adapted to fast flowing waters which inhibits its ability to develop large populations (Lloyd, 1987).

Habitat List

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Category	Sub-Category	Habitat	Presence	Status
Brackish
		Inland saline areas	Principal habitat	Harmful (pest or invasive)

		Estuaries	Secondary/tolerated habitat	Harmful (pest or invasive)
		Lagoons	Principal habitat	Harmful (pest or invasive)
Terrestrial
	Terrestrial ‑ Natural / Semi-natural	Riverbanks	Principal habitat	Harmful (pest or invasive)
	Terrestrial ‑ Natural / Semi-natural	Wetlands	Principal habitat	Harmful (pest or invasive)
Littoral
		Coastal areas	Secondary/tolerated habitat	Harmful (pest or invasive)
		Mangroves	Secondary/tolerated habitat	Harmful (pest or invasive)
		Mud flats	Secondary/tolerated habitat	Harmful (pest or invasive)
		Intertidal zone	Secondary/tolerated habitat	Harmful (pest or invasive)
		Salt marshes	Secondary/tolerated habitat	Harmful (pest or invasive)
Freshwater

		Irrigation channels	Secondary/tolerated habitat	Harmful (pest or invasive)
		Lakes	Principal habitat	Harmful (pest or invasive)
		Reservoirs	Secondary/tolerated habitat	Harmful (pest or invasive)
		Rivers / streams	Principal habitat	Harmful (pest or invasive)
		Ponds	Principal habitat	Harmful (pest or invasive)
Marine
Marine		Inshore marine	Secondary/tolerated habitat	Harmful (pest or invasive)

Biology and Ecology

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Genetics

For more on the genetics of this species please see Yardley and Hubbs (1976) and Wooten et al. (1988).

Associations

Disturbed environments are prone to invasion from this species due to the fact that disturbed habitats can support large populations of invertebrate species (which tend to be pests, e.g. chironomids) and often lack other fish “due to harsh physical conditions” (Lloyd, 1987). Mosquitofish are typically more resistant to pollutants, including organic wastes, herbicides, insecticides, rotenone, phenols, heavy metals and radiation, than most other fish (Lloyd, 1987; Edwards, 2005). The ability of the species to survive genetic ‘bottle-necks’ and their ability to physiologically and genetically adapt to different environments is also important in their spread and success.

Environmental Requirements

Mosquitofish have invaded a wide range of habitats throughout the world including: hot springs, rivers, streams, lakes, swamps, billabongs, cooling pondages, rice fields, ornamental ponds, estuaries, near-shore marine habitats and salt lakes (Lloyd, 1987; Arthington and Lloyd, 1989).

Mosquitofish are found in waters from 0.5°C to 39°C, with a preference for warm waters of about 25°C (Otto, 1974), though Pyke (2005) suggests that mosquitofish prefer water temperatures between 31-35°C. Juvenile mosquitofish are more thermally tolerant than adults, allowing them to colonise and exploit warm patches of the environment with increasing growth, survival, and maturation rate. Mosquitofish can inhabit ice-covered waters (Hirose, 1976;Sasa and Kurihara, 1980) in Japan and hot bores (over 37°C) in central Australia (John Glover, personal communication, cited in Lloyd, 1987).

The broad salinity tolerance of mosquitofish allows them to colonise environments, such as salt lakes, estuaries, near coastal marine environments (Lloyd, 1987). The salinity LD50 for mosquitofish is more than 58g/L and they can tolerate direct transfers to salinity differences of up to seawater (35 g/L) with few mortalities (Chervinski, 1983).

Mosquitofish can tolerate oxygen concentrations as low as 1.3 mg/L, without access to surface water but can withstand virtual anoxia by utilising of the oxygen-rich surface water/air interface because it has a dorso-ventral mouth (Lewis, 1970).

Climate

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Climate	Status	Description
A - Tropical/Megathermal climate	Preferred	Average temp. of coolest month > 18°C, > 1500mm precipitation annually
B - Dry (arid and semi-arid)	Tolerated	< 860mm precipitation annually
C - Temperate/Mesothermal climate	Preferred	Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C
D - Continental/Microthermal climate	Tolerated	Continental/Microthermal climate (Average temp. of coldest month < 0°C, mean warmest month > 10°C)

Latitude/Altitude Ranges

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Latitude North (°N)	Latitude South (°S)	Altitude Lower (m)	Altitude Upper (m)
50	40

Water Tolerances

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Parameter	Minimum Value	Maximum Value	Typical Value	Status	Notes
Dissolved oxygen (mg/l)			>1.3	Optimum
Salinity (part per thousand)			<20	Optimum	0-58 tolerated
Velocity (cm/h)				Optimum	Gambusia prefer slow flowing waters to fast
Water temperature (ºC temperature)	25	31		Optimum	0.5-39 tolerated

Natural enemies

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Natural enemy	Type	Life stages	Specificity	References
Glugea	Parasite	All Stages	to genus	Crandall and Bowser, 1982, recd. 1983
Goussia piekarskii	Parasite	All Stages	to genus	Lom and Dyková, 1995
Kudoa	Parasite	All Stages	not specific	Dyková et al., 1994

Means of Movement and Dispersal

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Mosquitofish have become the most widely distributed freshwater teleosts in the world (Krumholz, 1948) mainly through deliberate human introductions (e.g. Lintermans, 2004). Throughout the world, G. holbrooki and G. affinis have been widely distributed to aid mosquito control in rice paddies and natural waters. (Krumholz, 1948; Lloyd et al., 1986; Arthington and Lloyd, 1989). Worldwide introduction of Gambusia has occurred since the first introduction into Hawaii in 1905 (Krumholz, 1948).

Gambusia are cited to be used in the commercial aquarium industry (www.fishbase.org) but poor sales are likely given its noxious status in many countries, its aggressive behaviour, and its poor appearance. They are likely to be under aquaculture in the USA for use for mosquito control in rice fields. Mosquito control authorities have been known to transfer mosquitofish between locations. Children may also collect mosquitofish for bait and help to distribute it when they move from one place to another.

Pathway Causes

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Cause	Notes	Long Distance	Local
Biological control	For mosquito control	Yes
Flooding and other natural disasters	Natural spread once species is established		Yes
Intentional release	For mosquito control	Yes
Medicinal use	For mosquito control	Yes
Military movements	For mosquito control	Yes
Pet trade			Yes

Environmental Impact

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Impact on Biodiversity

In Australia, and possibly other parts of its introduced range, G. holbrooki faces few predators, parasites, diseases or competitors (Lloyd, 1987). Experiments have shown that several Australian native fish predators actively avoid eating this mosquitofish (Lloyd, 1987).

Rehage et al. (2005a, b) suggest that invasive species of Gambusia (G. holbrooki and G. affinis) are more efficient foragers than their non-invasive relatives (G. geiseri and G. hispaniolae), and that invasive juveniles feed more voraciously and widely than adults. This study suggests that Gambusia, released from their native habitat and therefore native predators, increase in size, and accordingly, have a higher feeding rate which has a greater impact on native species.

Many studies in the USA and Australia have found significant habitat overlap between mosquitofish and native fish throughout all stages of their respective life cycles. These overlaps combined with the superior competitive ability of Gambusia mean that native fish may be lost from waters where Gambusia dominates (Schoenherr, 1974, 1981; Arthington et al., 1983; Lloyd et al., 1986; Lloyd 1987; Arthington, 1989; Arthington and Lloyd, 1989; Lloyd, 1990; Arthington and Marshall, 1999; Rincon et al., 2002; King, 2003; Pyke, 2005).

G. holbrooki is a voracious predator of invertebrates resulting in a possible increase in mosquito populations, as it targets insects, which are natural predators of mosquitoes. Willems et al. (2005) compared two different fish species on their predation of mosquitoes which found that Gambusia were not as effective as controlling mosquito larvae as the native fish in the study. Studies of small native fish and G. holbrooki diets in the River Murray in South Australia (Lloyd, 1987) showed that the eastern mosquitofish was a poor predator of mosquito larvae compared to most of the small native fish species. Ling (2004) showed that G. affinis may inhabit an unoccupied niche in New Zealand environments and a threat to native invertebrate grazers. G. holbrooki has been shown, through gut contents assessment, to be significant predators of native fish in Australia (Ivantsoff and Aarn, 1999). It can also have an impact on the tadpoles of native frogs (Sadlier and Pressey, 1994; Morgan and Buttemer, 1996; Komak and Crossland, 2000; Pyke and White, 2000). The reproductive rituals, breeding success and growth of native fishes can be impacted by gambusia (Howe et al., 1997).

Threatened Species

Gambusia occupy the specialized dystrophic habitats of one restricted and two endangered Australian freshwater fishes - Rhadinocentrus ornatus, Pseudomugil mellis, Nannoperca oxleyana (all found in South-eastern Queensland). It is possible that G. holbrooki and the three species interact and compete for habitat, food and spawning areas (Howe et al., 1997; Arthington and Marshall, 1999; Knight and Arthington, 2008).

Threatened Species

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Threatened Species	Conservation Status	Where Threatened	Mechanism
Nannoperca oxleyana	EN (IUCN red list: Endangered)
Poeciliopsis occidentalis (Gila topminnow)	VU (IUCN red list: Vulnerable); USA ESA listing as endangered species	Queensland	Predation
Pseudomugil mellis	EN (IUCN red list: Endangered)	Queensland	Competition; Predation
Rhadinocentrus ornatus	No details

Risk and Impact Factors

Top of page Invasiveness

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
Benefits from human association (i.e. it is a human commensal)
Long lived
Fast growing
Has high reproductive potential
Gregarious
Has propagules that can remain viable for more than one year
Reproduces asexually
Has high genetic variability

Impact outcomes

Ecosystem change/ habitat alteration
Modification of natural benthic communities
Monoculture formation
Negatively impacts agriculture
Reduced native biodiversity
Threat to/ loss of native species

Impact mechanisms

Competition - monopolizing resources
Competition (unspecified)
Pest and disease transmission
Predation
Rapid growth

Likelihood of entry/control

Highly likely to be transported internationally deliberately
Difficult/costly to control

Uses

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G. holbrooki and G. affinis have been introduced widely for mosquito control. They are used in the commercial aquarium industry (www.fishbase.org) but poor sales are likely given their noxious status in many countries, aggressive behaviour, and poor appearance.

Uses List

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Environmental

Biological control

General

Laboratory use
Pet/aquarium trade

Diagnosis

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Genetic techniques have been used to distinguish G. holbrooki and G. affinis in Australia.

For information on the morphology of G. holbrooki please see this species' factsheet on FishBase.

Prevention and Control

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

Prevention

SPS measures

All of these measures have been applied to prevent the spread of Gambusia in Australia. The Australian Quarantine Inspection Service (AQIS) has listed G. holbrooki as a high-risk species, highly likely to establish and spread more widely once introduced to new areas (Arthington et al., 1999). Predation by G. holbrooki has been listed as a key threatening process under the New South Wales Threatened Species Conservation Act 1995. The inappropriate spread of Gambusia by humans for the purpose of mosquito control remains a problem in spite of repeated cautionary advice over the past 20 years (Arthington and Lloyd, 1989). Gambusia has been recognised as a potential vertebrate pest in Australia and as such should be managed under the edicts of vertebrate pest policies, programmes, and legislation (Bomford, 2001; Bomford and Glover, 2004).

Early warning systems

As eradication of fish after establishment as breeding populations is usually impossible, alien fish management should concentrate on preventing new introductions and limiting future spread (Lodge et al., 1998; Lodge and Shrader-Frechette, 2003). The resources to monitor and then act upon eradication of new populations are often limited. Involving the community in monitoring activities is one way to achieve early warning advice to management authorities. Involving the community provides people with a sense of ownership of the pest fish issue and an improved understanding of the complexities of managing alien fish species.

Rapid response

Not known to be a feature of Gambusia control but should be a first line of action (McDowall, 2004).

Public awareness

Gambusia is one of about 29 aquarium species listed as harmful to native fishes, invertebrates and aquatic ecosystem integrity in Australia (Arthington et al., 1999). Scientific publications, information papers, pamphlets and posters have all been used to inform the public about the dangers of alien fishes, especially those declared noxious under state and commonwealth legislation, or quarantine regulations such as those of the Australian Quarantine Inspection Service (AQIS). Involving the community in monitoring activities provides people with a sense of ownership of the pest fish issue and an improved understanding of the complexities of managing alien fish species.

Eradication

Attempts have been made to eradicate G. holbrooki from water bodies using the fish poison rotenone. Most native fish are killed by a rotenone concentration of 0.5 ppm but Gambusia can survive this concentration without mortality (Pyke, 2005). Impacts on native fishes and other native fauna have been mitigated by releasing potassium permanganate downstream of the rotenone release point in flowing waterways. Gambusia is more tolerant of the organo-phosphorus pesticide Dursban™ than several native fishes (Pyke, 2005). These observations mean that chemical control methods are highly likely to affect native fish and other aquatic biota well before useful levels of gambusia mortality can be achieved.

Containment/Zoning

May be applicable to Gambusia, but no explicit information available.

Control

Cultural control and sanitary measures

It is well established that many exotic fishes flourish in disturbed habitats, even highly polluted ones in the case of Gambusia. It has been argued that maintaining the natural flow regime and habitat characteristics of aquatic systems should be an effective preventative measure, indeed a management principle (Arthington et al., 1990).

Physical/mechanical control

Gambusia species have been controlled to some extent by draining standing water bodies (especially outdoor ornamental ponds), and by cutting off the pathways for re-colonization (e.g. closing off access drains into and from ornamental ponds). Gambusia is highly invasive and can move through shallow water half their body depth. Barriers to spread must obstruct dispersal pathways (e.g. ditches and channels) as shallow as 3 mm to prevent local spread of Gambusia (Alemadi and Jenkins, 2007).

Movement control

Declaration of noxious status prohibits movement of Gambusia by humans in some countries (e.g. Australia). When Gambusia is collected as by-catch in ecological research programmes it must be disposed of immediately, never transported or introduced to any other water-body or taken to domestic aquaria.

Biological control

Factors limiting natural Gambusia populations (Courtenay and Meffe, 1989) may include endemic parasites and pathogens, as well as predators. Gambusia is reported to host at least 23 parasite species (LN Lloyd, cited in Arthington and Lloyd, 1989), however, most are not host specific. Pathogens may contribute to suppression of Gambusia, yet reliable data on their role are scarce. In April 2000 a fungal infection killed more than 80% of G. affinis in Lake Naini Tal, Uttaranchal, India (Nagdali and Gupta, 2002). Gambusia in its native range may be partially controlled by predators including topminnows of the genus Fundulus. However, there are no reports suggesting that, beyond its native range Gambusia is other than a minor constituent of the diet of piscivores, even other alien fish species such as trout. If Gambusia is a major part of fish diets in some locations then manipulation of aquatic food webs may provide a means to control Gambusia via predation pressure. Unmack (unpublished data) and others have observed that Gambusia and larger galaxiids rarely coexist and galaxiids eagerly devour Gambusia in aquaria. In small water-bodies diadromous galaxiids such as the common galaxiid (Galaxias maculatus) could be introduced at high densities to prey on Gambusia. However, after 2-4 years the galaxiids would die of old age, leaving opportunities for Gambusia to re-establish viable populations. Repeated releases of the galaxiid would be needed to sustain high Gambusia mortality.

Chemical control

Attempts have been made to reduce the abundance of G.holbrooki from water-bodies using the poison rotenone. Impacts on native fishes and other fauna have been mitigated by releasing potassium permanganate downstream of the release point in flowing waterways.

IPM

The integration of science (e.g. alien species biology) with applied management actions will improve environmental outcomes, yet there does not appear to be any example of an integrated management programme to control Gambusia.

Monitoring and Surveillance

The presence and abundance of Gambusia species are being monitored as part of formal stream and river health monitoring programs in Australia (e.g. Kennard et al., 2005).

Ecosystem Restoration

Scientists have long advocated that the maintenance of healthy, functional aquatic ecosystems and the natural processes that support native biodiversity is the best way to suppress alien fishes (Arthington et al., 1990), i.e. maintain ecosystem resistance and resilience. Damaged ecosystems could be restored in ways that are likely to suppress Gambusia, e.g. by protecting riparian corridors, minimizing habitat loss, restoring natural flow regimes or releasing environmental flows to flush out mosquitofish and so constrain their spread and population increase (e.g. Bunn and Arthington, 2002). Issues of alien species being moved about by inter-basin water transfers have been addressed by Todd et al. (2002).

References

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Keane JP, Francisco JN, 2004. First record of mosquitofish, Gambusia holbrooki, in Tasmania, Australia: stock structure and reproductive biology. New Zealand Journal of Marine and Freshwater Research, 38:857-867

Kennard MJ, Arthington AH, Pusey BJ, Harch BD, 2005. Are alien fish a reliable indicator of river health? Freshwater Biology, 50(1):174-193

King A, 2003. Niche overlap between larvae of exotic and native fish in a lowland river. In: Presented at the "ASFB Invasive species: Fish and Fisheries Workshop", Australian Society for Fish Biology, Wellington, New Zealand 29-30 June

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Komak S, Crossland M, 2000. An assessment of introduced mosquitofish (Gambusia affinis holbrooki) as a predator of eggs, hatchlings, and tadpoles of native and non-native anurans. Wildlife Research, 27:185-189

Krumholz LA, 1948. Reproduction in the western mosquitofish Gambusia affinis and its use in mosquito control. Ecol. Monogr, 18(1):1-43

Lewis M, 1970. Morphological adaptations of Cyprinodontoids for inhabiting oxygen-deficient waters. Copeia, 2:319-326

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Lloyd LN, 1987. Ecology and distribution of the small native fish of the lower River Murray, South Australia and their interactions with the exotic mosquitofish, Gambusia affinis holbrooki (Girard). Dept. of Zoology, University of Adelaide

Lloyd LN, 1990. Ecological interactions of Gambusia holbrooki with Australian native fish. In: ASFB Workshop on introduced and translocated fishes and their ecological effects. Bureau of Rural Resources Proceedings No. 8 [ed. by Pollard DA]: AGPS, Canberra

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Lloyd LN, Arthington AH, Milton DA, 1986. The mosquitofish - a valuable mosquito control agent or a pest? In: The ecology of exotic plants and animals: some Australian case studies [ed. by Kitching]: John Wiley and Sons, Brisbane

Lloyd LN, Tomasov JF, 1985. Taxonomic status of the mosquitofish. Gambusia affinis (Poeciliidae), in Australia. Aust. Mar. Freshw. Res, 36:447-51

Lodge DM, Shrader-Frechette K, 2003. Nonindigenous species: ecological explanation, environmental ethics, and public policy. Conservation Biology, 17(1):31-37

Lodge DM, Steen RA, Brown KM, Covich AP, Bronmark C, Gavey JE, Klosiewiwski SP, 1998. Predicting impact of freshwater exotic snails on biodiverstiy: challenges in spatial scaling. Australian Journal of Ecology, 23:53-67

Lom J, Dyková I, 1995. Studies on protozoan parasites of Australian fishes. Notes on coccidian parasites with description of three new species. Systematic Parasitology, 31(2):147-156; 36 ref

McAllister DE, 1969. Introduction of tropical fishes into a hot spring near Banff, Alberta, Canada. Can. Field Nat, 83(1):31-35

McDowall RM, 2004. Shoot first and then ask questions: A look at aquarium fish imports and invasiveness in New Zealand. Journal of Marine and Freshwater Research, 38(3):503-510

Milton DA, Arthington AH, 1983. Reproductive biology of Gambusia affinis holbrooki Baird and Girard, Xiphophorus helleri (Gunther) and X. maculatus (Heckel) (Pisces; Poeciliidae) in Queensland, Australia. Journal of Fish Biology, 23(1):23-41

Morgan LA, Buttemer WA, 1996. Predation by the non-native fish Gambusia holbrooki on small Litoria aurea and L. dentata tadpoles. Australian Zoologist, 30(2):143-149

Motabar M, 1978. Larvivorous fish, Gambusia affinis - a review. Geneva, Switzerland: World Health Organization., 5 pp

Moyle PB, Light T, 1996. Biological invasions of fresh water: empirical rules and assembly theory. Biological Conservation, 78:149-161

Nagdali SS, Gupta PK, 2002. Impact of mass mortality of a mosquito fish, Gambusia affinis on the ecology of a fresh water eutrophic lake (Lake Naini Tal, India). Hydrobiologia, 468:45-52

Nordlie FG, 2000. Patterns of reproduction and development of selected resident teleosts of Florida salt marshes. Hydrobiologia, 434:165-182

Otto RG, 1974. The effects of acclimation to cyclic thermal regimes on heat tolerance of the western mosquitofish. Trans. Fish. Soc, 103(2):331-335

Pusey BJ, Kennard MJ, Arthington AH, 2004. Freshwater Fishes of North-Eastern Australia. Collingwood, Victoria, : CSIRO Publishing

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Pyke GH, White AW, 2000. Factors influencing predation on eggs and tadpoles of the endangered Green and Golden Bell Frog Litoria aurea by the introduced Plague Minnow Gambusia holbrooki. Australian Zoologist, 31(3):496-505

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Ross ST, Baker JA, 1983. The response of fishes to periodic spring floods in a southeastern stream. The American Midland Naturalist, 109(1):1-14

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Seale A, 1905. Report of Mr. Alvin Seale, of the United States Fish Commission, on the introduction of top-minnows to Hawaii from Galverston, Texas. Hawaiian Forestry and Agriculture, 2:364-367

Seale A, 1917. The mosquitofish, Gambusia affinis (Baird & Girard), in the Philippine islands. Philipp. J. Sci., 12(3):177-187

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Distribution References

Alemadi S D, Jenkins D G, 2007. Behavioral constraints for the spread of the eastern mosquitofish, Gambusia holbrooki (Poeciliidae). Biological Invasions.

Allen K R, 1956. The geography of New Zealand's freshwater fish. New Zealand Science Review. 14 (3), 3-9.

Anon, 2008. FishBase. In: FishBase, [ed. by Froese R, Pauly D]. http://www.fishbase.org

Banpot P, 2004. Extension Bulletin. Food & Fertilizer Technology Center, Thailand: 11 pp.

CABI, Undated. Compendium record. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

Das S M, Rampal C, 1966. On the life history of Gambusia affinis in Kashmir, India. Indian Science Congress, Proceedings. 53 (31), 376-377.

Fowler J, 1970. Introduction to panel: a review of Gambusia effectiveness. P.P.A.C.C.M.C.A. 72.

Froese R, Pauly D, 2004. FishBase. http://www.fishbase.org

Glucksman J, West G, 1976. The introduced fishes of Papua New Guinea with special reference to Tilapia mossambica. Biological Conservation. 37-44.

Keane J P, Francisco J N, 2004. First record of mosquitofish, Gambusia holbrooki, in Tasmania, Australia: stock structure and reproductive biology. New Zealand Journal of Marine and Freshwater Research. 857-867.

Krumholz L A, 1948. Reproduction in the western mosquitofish Gambusia affinis and its use in mosquito control. Ecological Monographs. 18 (1), 1-43.

Lloyd L N, 1987. Ecology and distribution of the small native fish of the lower River Murray, South Australia and their interactions with the exotic mosquitofish, Gambusia affinis holbrooki (Girard). Adelaide, Australia: Dept. of Zoology, University of Adelaide.

McAllister D E, 1969. Introduction of tropical fishes into a hot spring near Banff, Alberta, Canada. Canadian Field Naturalist. 83 (1), 31-35.

Motabar M, 1978. Larvivorous fish, Gambusia affinis - a review. In: Larvivorous fish, Gambusia affinis - a review. Geneva, Switzerland: World Health Organization. 5 pp.

Nordlie F G, 2000. Patterns of reproduction and development of selected resident teleosts of Florida salt marshes. Hydrobiologia. 165-182. DOI:10.1023/A:1004073125007

Pyke G H, 2005. A review of the biology of Gambusia affinis and G. holbrooki. Reviews in Fish Biology and Fisheries. 15 (4), 339-365. http://www.springerlink.com/link.asp?id=100215 DOI:10.1007/s11160-006-6394-x

Rees D H, 1934. Notes on Mosquitofish in Utah, Gambusia affinis (BAIRD & GIRARD). Copeia. 157-159.

Sasa M, Kurihara T, 1980. The use of poeciliid fish in the control of mosquitoes. In: Biocontrol of medical and veterinary pests. [ed. by Laird M]. Praeger.

Seale A, 1905. Report of Mr. Alvin Seale, of the United States Fish Commission, on the introduction of top-minnows to Hawaii from Galverston, Texas. Hawaiian Forestry and Agriculture. 364-367.

Seale A, 1917. The mosquitofish, Gambusia affinis (Baird & Girard), in the Philippine islands. Philippine Journal of Science. 12 (3), 177-187.

Wilson F, 1960. Tech. Commun. Commonw. Inst. Biol. Control, Ottawa, viii + 102.

Links to Websites

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Website	URL	Comment
Gambusia control homepage	http://www.gambusia.net
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gateway	https://doi.org/10.5061/dryad.m93f6	Data source for updated system data added to species habitat list.
Global register of Introduced and Invasive species (GRIIS)	http://griis.org/	Data source for updated system data added to species habitat list.

Contributors

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08/08/08 Original text by:

Angela Arthington, Australian Rivers Institute, Room 1.09A, Environ. 2, (Building N13), Griffith School of Environment, Griffith University, Nathan QLD 4111, Australia

Lance Lloyd, Lloyd Environmental Pty Ltd, PO Box 3014, Syndal, Victoria, 3149, Australia

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Datasheet

Gambusia holbrooki (eastern mosquitofish)

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