Gambusia holbrooki (eastern mosquitofish)
Index
- Pictures
- Identity
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Description
- Distribution
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat
- Habitat List
- Biology and Ecology
- Climate
- Latitude/Altitude Ranges
- Water Tolerances
- Natural enemies
- Means of Movement and Dispersal
- Pathway Causes
- Environmental Impact
- Threatened Species
- Risk and Impact Factors
- Uses
- Uses List
- Diagnosis
- Prevention and Control
- References
- Links to Websites
- Contributors
- Distribution Maps
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Top of pageIdentity
Top of pagePreferred 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
Top of pageThe 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
Top of page- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Chordata
- Subphylum: Vertebrata
- Class: Actinopterygii
- Order: Cyprinodontiformes
- Family: Poeciliidae
- Genus: Gambusia
- Species: Gambusia holbrooki
Notes on Taxonomy and Nomenclature
Top of pageDescription
Top of pageThe 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
Top of pageDistribution Table
Top of pageThe 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 Feb 2022Continent/Country/Region | Distribution | Last Reported | Origin | First Reported | Invasive | Reference | Notes |
---|---|---|---|---|---|---|---|
Africa |
|||||||
Algeria | Present | Introduced | 1926 | ||||
Central African Republic | Present | Introduced | 1958 | Invasive | |||
Congo, Democratic Republic of the | Present | ||||||
Côte d'Ivoire | Present | Introduced | Invasive | ||||
Egypt | Present | Introduced | 1929 | Invasive | |||
Ethiopia | Present | Introduced | 1938 | Invasive | |||
Ghana | Present | Introduced | Invasive | ||||
Kenya | Present | Introduced | Invasive | ||||
Madagascar | Present | Introduced | 1929 | Invasive | |||
Mauritius | Present | Introduced | Invasive | ||||
-Rodrigues | Present | Introduced | Invasive | ||||
Morocco | Present | Introduced | 1929 | Invasive | |||
Réunion | Present | Introduced | Invasive | ||||
South Africa | Present | Introduced | 1936 | Invasive | |||
Sudan | Present | Introduced | 1929 | Invasive | |||
Zambia | Present | Introduced | 1940 | Not established | |||
Zimbabwe | Present | Introduced | 1925 | Invasive | |||
Asia |
|||||||
Afghanistan | Present | Introduced | 1970 | Invasive | |||
Armenia | Present | Introduced | Invasive | ||||
Bangladesh | Present | Introduced | Invasive | ||||
Cambodia | Present | Introduced | Invasive | ||||
China | Present | Introduced | 1927 | Invasive | |||
Georgia | Present | Introduced | 1925 | Invasive | |||
Hong Kong | Present | Introduced | Invasive | ||||
India | Present | Introduced | Invasive | ||||
-Jammu and Kashmir | Present | Introduced | Invasive | ||||
Indonesia | Present | Introduced | 1929 | Not established | |||
-Irian Jaya | Present | Introduced | 1930 | Invasive | |||
Iran | Present | Introduced | 1928 | Invasive | |||
Iraq | Present | Introduced | 1928 | Invasive | Original citation: Al-Daham et al. (1977) | ||
Israel | Present | Introduced | 1924 | Invasive | |||
Japan | Present | Introduced | 1916 | Invasive | |||
Jordan | Present | Introduced | 1930 | Invasive | |||
Kazakhstan | Present | Introduced | 1934 | Invasive | |||
Laos | Present | Introduced | Invasive | ||||
Lebanon | Present | Introduced | Invasive | ||||
Malaysia | Present | Introduced | Invasive | ||||
Myanmar | Present | Introduced | Invasive | ||||
Pakistan | Present | Introduced | Invasive | ||||
Philippines | Present | Introduced | 1913 | Invasive | |||
Saudi Arabia | Present | Introduced | Invasive | ||||
Singapore | Present | Introduced | Invasive | ||||
Sri Lanka | Present | Introduced | Invasive | ||||
Syria | Present | Introduced | Invasive | Original citation: Gerberich and Laird (1968) | |||
Taiwan | Present | Introduced | 1911 | Invasive | |||
Tajikistan | Present | Introduced | Invasive | ||||
Thailand | Present | Introduced | 1919 | Invasive | |||
Turkey | Present | Introduced | 1920 | Invasive | |||
Turkmenistan | Present | Introduced | Invasive | ||||
United Arab Emirates | Present | Introduced | Invasive | ||||
Uzbekistan | Present | Introduced | 1930 | Invasive | |||
Vietnam | Present | Introduced | Invasive | ||||
Yemen | Present | Introduced | Invasive | ||||
Europe |
|||||||
Albania | Present | Introduced | Invasive | ||||
Austria | Present | Introduced | Invasive | ||||
Bulgaria | Present | Introduced | 1924 | Invasive | |||
Cyprus | Present | Introduced | 1926 | Invasive | Original citation: Gerberich and Laird (1968) | ||
Federal Republic of Yugoslavia | Present | Introduced | 1924 | Invasive | Original citation: Gerberich and Laird (1968) | ||
France | Present | Introduced | 1924 | Invasive | |||
-Corsica | Present | Introduced | 1924 | Invasive | Original citation: Gerberich and Laird (1968) | ||
Germany | Present | Introduced | 1921 | Invasive | |||
Greece | Present | Introduced | 1928 | Invasive | |||
-Crete | Present | ||||||
Hungary | Present | Introduced | 1937 | Invasive | |||
Italy | Present | Introduced | 1922 | Invasive | |||
Portugal | Present | Introduced | 1921 | Invasive | |||
Romania | Present | Introduced | 1921 | Invasive | |||
Russia | Present | Introduced | 1925 | Invasive | |||
Slovenia | Present | Introduced | 1927 | ||||
Spain | Present | Introduced | 1921 | ||||
-Canary Islands | Present | Introduced | 1943 | Invasive | |||
Ukraine | Present | Introduced | Invasive | ||||
North America |
|||||||
Canada | Present | Introduced | 1924 | Invasive | |||
Haiti | Present | Introduced | 1983 | Invasive | |||
Mexico | Present | Introduced | 1931 | Invasive | |||
Puerto Rico | Present | Introduced | 1914 | Invasive | |||
United States | Present | Native | |||||
-Florida | Present | ||||||
-Hawaii | Present | Introduced | 1905 | Invasive | |||
-Michigan | Present | Introduced | 1941 | Invasive | |||
-Utah | Present | Introduced | 1927 | Invasive | |||
Oceania |
|||||||
American Samoa | Present | Introduced | Invasive | ||||
Australia | Present | Introduced | |||||
-New South Wales | Present, Widespread | Introduced | 1926 | Invasive | |||
-Northern Territory | Present, Localized | Introduced | 1940 | Invasive | |||
-Queensland | Present, Widespread | Introduced | 1925 | Invasive | |||
-South Australia | Present, Widespread | Introduced | 1926 | Invasive | |||
-Tasmania | Present, Localized | Introduced | Invasive | First reported: 1990s | |||
-Victoria | Present, Widespread | Introduced | 1926 | Invasive | |||
-Western Australia | Present, Localized | Introduced | 1934 | Invasive | |||
Cook Islands | Present | Introduced | Invasive | ||||
Federated States of Micronesia | Present | Introduced | Invasive | ||||
Fiji | Present | Introduced | 1930 | Invasive | |||
French Polynesia | Present | Introduced | Invasive | ||||
Guam | Present | Introduced | Invasive | ||||
Kiribati | Present | Introduced | Invasive | ||||
Marshall Islands | Present | Introduced | Invasive | ||||
New Zealand | Present | Introduced | 1930 | Invasive | |||
Northern Mariana Islands | Present | Introduced | Invasive | ||||
Papua New Guinea | Present | Introduced | 1930 | Invasive | |||
Samoa | Present | Introduced | Invasive | ||||
Solomon Islands | Present | Introduced | 1930 | Invasive | |||
South America |
|||||||
Argentina | Present | Introduced | 1943 | Invasive | |||
Bolivia | Present | Introduced | Invasive | ||||
Chile | Present | Introduced | 1937 | Invasive | |||
Peru | Present | Introduced | 1940 | Invasive |
History of Introduction and Spread
Top of pageG. 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).
Risk of Introduction
Top of pageHabitat
Top of pageThe 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
Top of pageCategory | Sub-Category | Habitat | Presence | Status |
---|---|---|---|---|
Brackish | Inland saline areas | Principal habitat | Harmful (pest or invasive) | |
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) | |
Littoral | Mangroves | Secondary/tolerated habitat | Harmful (pest or invasive) | |
Littoral | Mud flats | Secondary/tolerated habitat | Harmful (pest or invasive) | |
Littoral | Intertidal zone | Secondary/tolerated habitat | Harmful (pest or invasive) | |
Littoral | Salt marshes | Secondary/tolerated habitat | Harmful (pest or invasive) | |
Freshwater | ||||
Freshwater | Irrigation channels | Secondary/tolerated habitat | Harmful (pest or invasive) | |
Freshwater | Lakes | Principal habitat | Harmful (pest or invasive) | |
Freshwater | Reservoirs | Secondary/tolerated habitat | Harmful (pest or invasive) | |
Freshwater | Rivers / streams | Principal habitat | Harmful (pest or invasive) | |
Freshwater | Ponds | Principal habitat | Harmful (pest or invasive) | |
Brackish | ||||
Brackish | Estuaries | Secondary/tolerated habitat | Harmful (pest or invasive) | |
Brackish | Lagoons | Principal habitat | Harmful (pest or invasive) | |
Marine | Inshore marine | Secondary/tolerated habitat | Harmful (pest or invasive) |
Biology and Ecology
Top of pageGenetics
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).
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
Top of pageClimate | Status | Description | Remark |
---|---|---|---|
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
Top of pageLatitude North (°N) | Latitude South (°S) | Altitude Lower (m) | Altitude Upper (m) |
---|---|---|---|
50 | 40 |
Water Tolerances
Top of pageParameter | Minimum Value | Maximum Value | Typical Value | Status | Life Stage | 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
Top of pageNatural enemy | Type | Life stages | Specificity | References | Biological control in | Biological control on |
---|---|---|---|---|---|---|
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
Top of pageMosquitofish 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
Top of pageCause | Notes | Long Distance | Local | References |
---|---|---|---|---|
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
Top of pageImpact 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).
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
Top of pageThreatened Species | Conservation Status | Where Threatened | Mechanism | References | Notes |
---|---|---|---|---|---|
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- 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
- Ecosystem change/ habitat alteration
- Modification of natural benthic communities
- Monoculture formation
- Negatively impacts agriculture
- Reduced native biodiversity
- Threat to/ loss of native species
- Competition - monopolizing resources
- Competition (unspecified)
- Pest and disease transmission
- Predation
- Rapid growth
- Highly likely to be transported internationally deliberately
- Difficult/costly to control
Uses
Top of pageG. 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.
Diagnosis
Top of pageGenetic 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
Top of pageDue 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
Rapid response
Public awareness
Eradication
Containment/Zoning
Control
Cultural control and sanitary measures
Physical/mechanical control
Movement control
Biological control
Chemical control
IPM
Monitoring and Surveillance
References
Top of pageAllen KR, 1956. The geography of New Zealand's freshwater fish. New Zeal. Sci. Rev, 14(3):3-9
Arthington AH, Mitchell DS, 1986. Aquatic invading species. Ecology of biological invasions, 34-53
Edwards TE, 2005. Environmental influences on mosquitofish reproduction. University of Florida
Fowler J, 1970. Introduction to panel: a review of Gambusia effectiveness. P.P.A.C.C.M.C.A, 38:72
Froese R, Pauly D, 2004. FishBase DVD. Penang, Malaysia: Worldfish Center. Online at www.fishbase.org
Froese R, Pauly D, 2008. FishBase. http://www.fishbase.org
Hubbs C, Peden AE, 1969. Gambusia georgi sp. nov. from San Marcos, Texas. Copeia, 2:357-364
Hubbs CL, 1955. Hybridisation between fish species in nature. Syst. Zool, 4(1):1-20
Rees DH, 1934. Notes on Mosquitofish in Utah, Gambusia affinis (BAIRD & GIRARD). Copeia, 4:157-159
Sterba G, 1962. Freshwater Fishes of the World. London, : Vista
Williamson MH, Fitter A, 1996. The characteristics of successful invaders. Biological Conservation
Distribution References
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
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
Rees D H, 1934. Notes on Mosquitofish in Utah, Gambusia affinis (BAIRD & GIRARD). Copeia. 157-159.
Wilson F, 1960. Tech. Commun. Commonw. Inst. Biol. Control, Ottawa, viii + 102.
Links to Websites
Top of pageWebsite | 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
Top of page08/08/08 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
Distribution Maps
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