Pacific Northwest Aquatic Invasive Species Profile:
Western mosquitofish (Gambusia affinis)
Laura Johnson
FISH 423
December 3, 2008
Figure 1. Western mosquitofish G. affinis (photo source: www.usgs.gov).
Diagnostic information eleven short spines on ray 3 (Page and Burr
1991).
Order: Cyprinodontiformes Until 1988, both the western mosquitofish (G.
Family: Poeciliidae affinis) and eastern mosquitofish (G. holbrooki)
Genus: Gambusia were classified as subspecies of G. affinis. The
Species: affinis classification of each fish as a separate species is
important since they are native to different
Common names: Western mosquitofish, portions of the eastern United States (Wooten et
mosquitofish al. 1988). G. affinis can be distinguished from
G. holbrooki by having six dorsal rays instead of
The western mosquitofish, Gambusia seven, and a lack of prominent teeth on
affinis, is a small (maximum 6.5 cm) gray or gonopodial ray three (Page and Burr 1991).
brown fish with a rounded tail and upturned
mouth (Figures 1 and 2). It may have a large Life-history and basic ecology
dusky to black teardrop marking beneath its eye
(as in Figure 1), but this marking is sometimes Life cycle
reduced (as in Figure 2). G. affinis has a dark G. affinis are ovoviviparous, meaning
stripe along its back to the dorsal fin, yellow and that the young develop within eggs inside the
blue iridescence on transparent silver-gray body mother’s body and are then born live and do not
sides, and six dorsal rays. G. affinis can be receive additional nourishment from the mother
further distinguished from other members of its (Wydoski and Whitney 2003). Newborns are
genus by a gonopodium with an elbow on ray 4a small, and weigh only about 1.2 to 1.3 mg.
composed of two or more segments, and eight to Maturation can occur in as little as 3 to 4 weeks,
although individuals born at the end of the
reproductive season may delay maturity for 6 to
7 months, until the beginning of the following
season. The lifespan of G. affinis differs for
males and females, with females living 6 months
to 1.5 years, and males averaging a much shorter
lifespan, although an accurate estimate is
unavailable (Haynes and Cashner 1995).
Figure 2. G. affinis in a human hand to
demonstrate approximate size (photo source:
www.aqua-fish.net)
Feeding habits anything but a mosquito larvae specialist (Gido
G. affinis is omnivorous and utilizes a and Franssen 2007; Goodsell and Kats 1999).
variety of food sources, which may give it an The predatory feeding behavior of G. affinis
advantage in colonizing new sites (Lockwood et causes serious ecosystem alterations, as shown
al. 2007). It is an aggressive predator and by Hurlbert et al. (1972) and Meffe (1985).
commonly preys on the eggs, juveniles, and Hurlbert et al. (1972) conducted a feeding study
small adults of other fish species. Like many involving G. affinis and found that it reduced
predatory fishes, they have strong, conical teeth crustaceans, insects, and rotifer populations
and short guts, and they consume terrestrial and within experimental pools, which subsequently
aquatic vertebrates, detritus, algae, and vascular caused an increase in phytoplankton
plants (Meffe and Snelson 1989). Grubb (1972) populations. Additionally, Meffe (1985)
also found that G. affinis prey heavily on anuran documented the impact of G. affinis on the
amphibian eggs when they are available (Grubb population of the Sonoran topminnow
1972). Similarly, G. affinis preys heavily on the (Poeciliopsis occidentalis), an endangered fish
larvae of California newts and the tadpoles of native to the southwestern United States. G.
Pacific treefrogs, and may negatively affect their affinis is implicated in the local extirpation of P.
populations (Goodsell and Kats 1999). G. occidentalis in its native habitat, partially due to
affinis also engage in cannibalism, which may predation by G. affinis (Meffe 1985).
provide growth and reproductive benefits for
individuals (Meffe and Crump 1987). Reproductive strategies
Although G. affinis received both its name and G. affinis displays high fecundity and
reputation for its supposed predation on short gestation periods, both of which may
mosquito larvae, studies have demonstrated that confer success in the biological invasion process
when other food sources are available, G. affinis (Haynes and Cashner 1995; Lockwood et al.
does not necessarily prefer mosquito larvae. For 2007). The reproductive season generally lasts
example, Goodsell and Kats (1999) showed that about 7 months, although in warmer climates
even when mosquito larvae were provided as such as Hawaii, reproduction may occur year-
food for G. affinis in a controlled feeding round. In areas where reproductive seasons are
experiment, they still preyed voraciously on shorter, they usually begin in spring and end in
amphibian tadpoles. Furthermore, analyses of fall. The gestation period ranges from only one
stomach contents of wild-caught G. affinis in to three weeks. Females may produce anywhere
two separate studies revealed a variety of prey in from one to seven broods per reproductive
their stomachs, proving that G. affinis is season, and each brood may contain up to about
200 embryos. The fecundity of G. affinis is
high, but ultimately depends on female size and these conditions are lethal to most freshwater
reproductive status as well as geographic fish, G. affinis individuals were able to survive
location. Fecundity tends to decrease from north at least a week in this environment (Hubbs
to south and east to west (Haynes and Cashner 2000).
1995). In addition, female G. affinis are able to
store sperm for extended periods of time, so that Biotic associations
one mating may result in multiple broods (Farr The primary biotic associations of G.
1989). affinis are parasitic, and no commensal or
pathogenic associations are described in the
Environmental optima and tolerances literature. G. affinis individuals commonly
The optimal environment of G. affinis is suffer from black spot disease, which is caused
warm, shallow, slow-moving waters with dense by a parasitic trematode commonly known as a
vegetation, high mineral content, and abundant black grub (Tobler and Schlupp 2008). Black
food organisms. Specifically, they thrive in grubs use freshwater fish as an intermediate
fresh and brackish water, and are found in host, between their definitive host, the belted
ponds, ditches, lakes, creeks, rivers, and springs. kingfisher, and snails. Black grubs enter
They are generally limited by cold temperatures, freshwater fish such as G. affinis by penetrating
and cannot tolerate temperatures below 4 the skin and becoming enclosed in the fish’s
degrees Celsius, which limits the distribution of tissue, where they form pinhead-sized black
G. affinis in the northern United States. They spots, hence the name black spot disease (Figure
are occasionally found in colder climates due to 3). The trematodes can greatly harm the host
annual stocking, but they are not able to fish, but they cannot spread from fish to fish.
establish a self-sustaining population. However, The penetration of the parasites into the skin can
they can tolerate water temperatures up to 38 cause mechanical damage and hemorrhage of
degrees Celsius and thrive in warm climates the host fish, as well as lipid depletion, which
(Wydoski and Whitney 2003). can lead to death (Lane and Morris 2000).
Hubbs (2000) demonstrated the ability of G. Tobler and Schlupp (2008) observed a decrease
affinis to thrive in both high and low in shoaling behavior among G. affinis with black
environmental quality environments. During a spot disease. They hypothesized that individuals
drought in west Texas in 1988, the Diamond/Y with different markings (black spots) caused
Draw drainage experienced extremely poor increased detection and subsequent predation of
environmental quality of 41 ppt salinity, total healthy individuals. Additionally, black spot
ammonia over 10 ppm, nitrates over 100 ppm, disease is associated with increased energy
and dissolved oxygen below 1 ppm. Although demands due to lipid loss, and shoaling with
sp., although the effect of this parasite on natural
populations of G. affinis is not known (Crandall
and Bowser 1981).
Current geographic distribution
G. affinis has established populations in
many states outside of its native range, which is
Figure 3. A fish (bass) exhibiting black spot disease located in the southeastern United States (Figure
(photo source: Fisheries Division, Michigan Dept. of 4). G. affinis exists throughout the Pacific
Natural Resources). Northwest, mostly along river corridors such as
the Columbia, Willamette, Snake, Klamath,
individuals requiring extra resources may reduce Clark Fork, and Flathead (Figure 5). In
food available to other group members. As a Washington State, biologists found G. affinis
result, healthy G. affinis preferred to shoal with overwintering in ponds near the confluence of
other healthy individuals and excluded those the Snake and Columbia Rivers in the 1970s,
with black spot disease (Tobler and Schlupp and it has since spread into other portions of the
2008). Columbia River. In the 1990s, they were
Tapeworms may also infect G. affinis, which captured in the John Day River, Lewis River,
serve as the definitive host for the tapeworm and Yakima River. Generally, they inhabit
Bothriocephalus acheilognathi. Granath and slow-moving backwaters and sloughs in the
Esch (1983) found that infection with B. mid- to lower Columbia River and the lower
acheilognathi caused decreased survival in G. reaches of the Columbia’s tributaries. However,
affinis. Increasing temperatures reduced since they are widely distributed for mosquito
survival in infected fish, as higher temperatures control in private ponds, they may occur in
caused increased growth of the tapeworm almost any water body in the Pacific Northwest
(Granath and Esch 1983). (Wydoski and Whitney 2003).
Scientists have found G. affinis populations
infected with the microsporidian parasitic History of Invasiveness
Glugea sp. in California. Glugea sp. can occupy
the majority of the abdominal cavity of G. affinis Due to its reputation for biological
with cyst-like structures and cause decreased control of mosquitoes, G. affinis has a long
swimming ability. Mortality may also occur invasion history and has been stocked
when G. affinis is heavily infected with Glugea throughout most of the United States and the rest
Figure 4. The current distribution of G. affinis in the United States. Note that the various shades of pink and red
represent hydrologic units where G. affinis occur (color difference indicates scale of information, not population
density). Light beige represents areas with no known populations of G. affinis, while darker brown indicates its native
range (source: USGS Nonindigenous Aquatic Species program, nas.er.usgs.gov).
were generally unknown or undocumented; thus,
both species are widely distributed around the
world (Fuller et al. 1999).
In 1901, the entomologist Leland Ossian
Howard advocated for the use of Gambusia sp.
in mosquito control after he obtained
information about its feeding habits. Several
years later, in 1905, Gambusia sp. individuals
were transported from North Carolina to
Figure 5. The current distribution of G. affinis in the Camden, New Jersey and released. This is the
Pacific Northwest. Color-coding is the same as in Figure 3 first known introduction of Gambusia sp.
(source: USGS Nonindigenous Aquatic Species program, outside its native range. Several years later, the
nas.er.usgs.gov). mosquitofish was transported to Hawaii and the
of the world. It is important to note that most of Philippines and released for mosquito control,
the introductions of the mosquitofish took place and populations successfully established in each
prior to the distinction of G. affinis and G. location. In the 1920s, Gambusia sp. were
holbrooki, and the origins of introduced stocks introduced into Europe, and shortly thereafter to
Asia and Africa (Krumholz 1948). Now, these populations have displayed the ability to
mosquitofish have established self-sustaining spread from the locations of initial stocking
populations on every continent in the world, (Fuller et al. 1999; Rehage and Sih 2004). G.
except Antarctica (Courtenay and Meffe 1989). affinis display nearly all of the characteristics
commonly associated with successful invasive
Invasion process species, as identified by Lockwood et al. (2007):
abundant and widely distributed in their native
Pathways, vectors, and routes of introduction range, broad environmental tolerances, short
The primary pathway of introduction of generation times, high fecundity, rapid growth,
G. affinis is stocking for biocontrol. Both early sexual maturity, broad diet, and association
citizens and government agencies routinely with human activities. Most likely, all of these
stock G. affinis for predation on mosquito larvae traits influence the successful establishment and
(Fuller et al. 1999). In many states, fish and spread of G. affinis outside its native range.
wildlife departments conduct annual stocking of Furthermore, G. affinis has demonstrated an
G. affinis (Schleier et al. 2007; Fuller et al. ability to live in disturbed, degraded habitats
1999). Additionally, state and local health such as retention ponds and dammed,
departments appear to view G. affinis as an channelized, and diverted rivers (Courtenay and
attractive alternative to pesticides, and Meffe 1989). The ability of G. affinis to survive
encourage citizens to stock western mosquitofish in environmental conditions that are lethal to
in backyard ponds and water features (Fuller et most other freshwater fish may also confer a
al. 1999). Due to the widespread distribution of competitive advantage and allow the species to
G. affinis and stocking activities conducted by withstand environmental events that eradicate
both government agencies and private other species in a given location (Hubbs 2000).
individuals, vectors and routes associated with Thus, as human activity degrades habitats, an
the intentional introduction of G. affinis are ecological niche becomes available which G.
probably numerous but remain unidentified in affinis appear to be more adept at filling than are
the literature. Some introductions of G. affinis native fish (Courtenay and Meffe 1989).
are probably escapees from stocking locations, Rehage and Sih (2004) conducted an experiment
in flood events or other natural disasters to test the dispersal behavior of four species of
(Courtenay and Meffe 1989). Gambusia, and hypothesized that species with
greater invasion success such as G. affinis would
Factors influencing establishment and spread display greater dispersal tendencies than
G. affinis has commonly established Gambusia species that were not successful
self-sustaining populations where stocked, and invaders (such as G. geiseri). As expected, G.
affinis were more likely to disperse out of Potential ecological and/or economic impacts
experimental introductory pools, and According to Courtenay and Meffe
subsequently dispersed faster and covered (1989), G. affinis and G. holbrooki have had the
greater distances than did non-invasive greatest ecological impacts of all of the poeciliid
Gambusia species (Rehage and Sih 2004). The (live-bearing) fish. The primary ecological
bold dispersal tendencies exhibited by G. affinis impact is predation on larvae, juveniles, or small
in experimental introductory pools are probably adults of other fish species, and such predation
displayed in wild populations, and assist with has locally extirpated native fish in some regions
the successful spread of introduced populations. (Courtenay and Meffe 1989; Meffe 1985).
The largest limitation to the establishment and Additionally, they have substantial effects on
spread of G. affinis is its intolerance to waters native amphibian populations due to predation
below 4 degrees Celsius. In cold climates, it on eggs and tadpoles (Grubb 1972; Goodsell and
lacks the ability to overwinter and where stocked Kats 1999). Adult G. affinis are extremely
in cold climates, it must be restocked annually aggressive towards other fish, even fish larger
(Wydoski and Whitney 2003). Due to its water than they are. They may attack these fish,
temperature requirements, G. affinis has spread killing them or injuring them enough to cause
throughout temperate and tropical regions of the eventual death (Courtenay and Meffe 1989).
world but has failed to spread in colder regions Potentially, G. affinis may continue to cause
(Fuller et al. 1999). local extirpation of native fish and amphibians,
Additional limitations to G. affinis populations or even cause extinction of some minnow
are heavy flooding, springhead conditions, and species that are already endangered (Meffe
natural predators. Flash flooding in the 1985). Additional negative ecological impacts
southwest United States has reduced or include hybridization with native species and
destroyed populations of G. affinis, while native introduction of parasites, but both of these
fish are able to withstand the high flow potential impacts of G. affinis have not been
conditions. Springhead waters in the southwest thoroughly investigated (Courtenay and Meffe
United States have low pH and high dissolved 1989).
CO2, and have served as a refugia for native fish G. affinis may also be responsible for ecosystem
when threatened with invasion by G. affinis. level impacts. In one study, G. affinis
Finally, predation by fish and aquatic birds may introduction caused large phytoplankton blooms,
limit populations of G. affinis (Courtenay and decreased water clarity, and higher water
Meffe 1989). temperatures (Hurlbert et al. 1972). Another
study found that experimental ponds containing
G. affinis had higher pH and oxygen levels than
those without (Hurlbert and Mulla 1981). plan that applies to both G. holbrooki and G.
Alteration of the ecosystem due to G. affinis affinis. Both G. holbrooki and G. affinis are
invasion may have far-reaching impacts. invasive in Australia, and have had detrimental
Although the literature has not explicitly stated effects on native fish populations there. Their
the economic impact of G. affinis, it is management plan details chemical, biological,
implicated in the reduction of populations of and physical methods that may be used to
largemouth bass due to predation on eggs, control Gambusia. Chemical methods include
larvae, and juveniles (Meffe 1985). Presumably, creation of Gambusia-free environments with
if G. affinis is able to cause a substantial decline the non-specific chemical rotenone prior to
in the populations of popular sportfish, stocking with native fish and amphibians.
economic impacts will be evident. However, due to the non-specific nature of
rotenone, the New South Wales National Parks
Management strategies and control methods and Wildlife Service prefers biological and
physical control methods over chemical ones.
Very few management strategies and The favored biological control method is
control methods are in place for G. affinis in the predation on Gambusia by larger species. The
United States, since many government agencies use of parasites, pathogens, bacteria and viruses
view the fish as beneficial in reducing mosquito to control Gambusia are also an option, but have
populations, despite the fact that studies have not been researched thoroughly at this time. The
shown otherwise. Typically, management preferred physical control method is the draining
consists of a permitting process in which citizens and drying of particular habitats critical to native
desiring G. affinis for backyard ponds must state species followed by reduction of water levels to
that they will not release individuals into prevent Gambusia from re-entering the
connected waterways. Although scientists and waterbodies. Finally, the New South Wales
agencies recognize G. affinis as a highly National Parks and Wildlife Service
invasive species and many studies show its recommends restoring degraded habitats to their
detrimental impacts on other species, original state, since Gambusia thrive in
governments continue to stock them outside disturbed, degraded habitats. Restoration of
their native range for biocontrol of mosquitoes, degraded habitats may give native species a
and allow citizens to follow suit on private long-term competitive advantage over Gambusia
property (Courtenay and Meffe 1989). (New South Wales National Parks and Wildlife
Although not local, the National Parks Service 2003).
and Wildlife Service of New South Wales,
Australia, has created a detailed management
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introduced mosquitofish on Pacific
Courtenay WR Jr, Meffe GK (1989) Small treefrogs and the role of alternative prey.
fishes in strange places: a review of Conserv Biol 13:921-924
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Hall, Englewood Cliffs, pp 319-331 mosquitofish in a predator-free, North
Carolina cooling reservoir. Amer Midland
Crandall TA, PR Bowser (1981) A Nat 110:314-323
microsporidian infection in a natural
population of mosquitofish Gambusia Grubb JC (1972) Differential predation by
affinis. J Fish Diseases 4:317-324 Gambusia affinis on the eggs of seven
species of anuran amphibians. Amer
Farr JA (1989) Sexual selection and secondary Midland Nat 88:102-108
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determinants of male mating success and Haynes JL, RC Cashner (1995) Life history and
the evolution of female choice. In: Meffe population dynamics of the western
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inland waters of the United States.
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Bethesda, MD mosquitofish (Gambusia affinis) predation
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Gido KB, NR Franssen (2007) Invasion of 83:125-151
stream fishes into low trophic positions. Hurlbert SH, J Zedler, D Fairbanks (1972)
Ecol Freshwater Fishes 16:457-464 Ecosystem alteration by mosquitofish
(Gambusia affinis) predation. Science
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western mosquitofish, Gambusia affinis freshwater fishes: North America north of
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Lockwood JL, MF Hoopes, MP Marchetti Schleier JJ, SE Sing, RKD Peterson (2007)
(2007) Invasion ecology. Blackwell Regional ecological risk assessment for the
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holbrooki – the plague minnow.
Other key sources of information and Phone: 360-7539508
bibliographies Email: [email protected]
United States Geological Survey Mosquitofish Allen Pleus
Information: Aquatic Nuisance Species Coordinator
http://nas.er.usgs.gov/queries/FactSheet.asp?spe Washington Department of Fish and Wildlife
ciesID=846 Phone: 360-902-2724
Email:
[email protected]Gambusia Control Homepage:
http://www.gambusia.net/ Pam Meacham
Assistant Aquatic Nuisance Species Coordinator
Fishbase species information page for Gambusia Washington Department of Fish and Wildlife
affinis: Phone: 360-902-2741
http://www.fishbase.org/Summary/speciesSumm Email:
[email protected]ary.php?ID=3215&genusname=Gambusia&spec
iesname=affinis Current research and management efforts
NatureServe Gambusia affinis information page: 1. University of California Davis:
http://www.natureserve.org/explorer/servlet/Nat The UC Mosquito Research Program
ureServe?searchName=Gambusia%20affinis investigates options for controlling mosquito
populations using native fish rather than G.
Michigan Department of Natural Resources affinis.
technical report describing alternatives to
Gambusia for mosquito control: Contact: Gregory Lanzaro, Ph.D.,
http://www.michigandnr.com/PUBLICATIONS/
[email protected]PDFS/ifr/ifrlibra/technical/reports/2003-2tr.pdf
2. University of California Riverside:
Expert contact information in the Pacific Dr. William E. Walton at UC Riverside
Northwest specializes in mosquito biology and ecology,
Kevin Aitkin including control methods involving native fish
Fish Biologist instead of G. affinis.
Invasive Species Lead
Western Washington Office Contact: William E. Walton, Ph.D.,
U.S. Fish and Wildlife Service
[email protected] 3. Edith Cowan University, Centre for
Ecosystem Management, Western Australia:
Dr. Mark Lund conducts wetlands research,
including the effect of G. holbrooki on Western
Australian wetlands.
