Siamese Fighting Fish (Betta splendens)

Ecological Risk Screening Summary

U.S. Fish and Wildlife Service, June 2019
Web Version, 11/7/2019

Photo: B. Arlandis. Licensed under CC BY-SA 2.0. Available: https://flic.kr/p/eke3fg. (March

2019).

1 Native Range and Status in the United States

Native Range
From Nico and Neilson (2019):

“Tropical Asia. Native to Southeast Asia including the northern Malay Peninsula, central and
eastern Thailand, Kampuchea, and southern Vietnam (Witte and Schmidt 1992).”

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From Vidthayanon (2013):

“Endemic to Thailand, from the Mae Khlong to Chao Phraya basins, the eastern slope of the
Cardamom mountains, and from the Isthmus of Kra.”

Froese and Pauly (2019) report B. splendens as native to Cambodia, Laos, Thailand, and
Vietnam.

Status in the United States

From Nico and Neilson (2019):

“Specimens ranging from 1-3 cm were taken from the Thames River drainage in Connecticut
(Whitworth 1996). Specimens were collected from Lake Worth Drainage District canal L-15,
adjacent to an aquarium fish farm west of Atlantis in Palm Beach County, Florida (Ogilvie 1969;
Courtenay et al. 1974), but have not been found at that locality in subsequent years (Courtenay
and Hensley 1979). A population became established in a canal south of Holmberg Road in
Parkland, northern Broward County. When first discovered in December 1975, it was a dominant
fish; however, the population was killed by extremely cold weather in January 1977 (Courtenay
and Hensley 1979; Courtenay et al. 1984; Courtenay and Stauffer 1990). A single specimen was
collected in the headwaters of Gamble Creek, near Parrish (UF 118475).”

“Failed in Connecticut. Shafland et al. (2008b) list it as a formerly reproducing species based on
a recent survey (Shafland et al. 2008a).”

“A supposed report of Betta splendens in Hiko Spring, Lincoln County, Nevada, was false (W.
Courtenay, personal communication).”

This species is common in trade in the United States. For example, ornamental varieties of Betta
splendens for sale at Petco (2019) include Baby Boy Betta, Baby Girl Betta, male Black Orchid
Betta, male Bumblebee Betta, male Butterfly Betta, male Cambodian Crowntail Betta, male and
female Crowntail Betta, male Copper Betta, male Deltatail Betta, male Doubletail Betta, male
Red Platinum Dragon, male Dragonscale Betta, male and female Elephant Ear Betta, male and
female Halfmoon Betta, male Halfmoon Doubletail Betta, male Halfmoon Doubletail Plakat
Betta, male Halfmoon King Betta, male King Betta, male and female Koi Betta, male Paradise
Betta, male Rose Gold Betta, male Rose Petal Betta, male and female Veiltail Betta, male
Cambodian Veiltail Betta, and male White Opal Betta.

Means of Introductions in the United States

From Nico and Neilson (2019):

“Florida records are the result of probable escapes from local ornamental fish farms (Courtenay
and Stauffer 1990). Connecticut records are probably aquarium releases.”

Remarks
A previous version of this ERSS was published in 2014.

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From International Betta Congress (2018):

“They are often commonly referred to simply as “betta” or “betta fish” [bet-uh].”
“Bettas have evolved far beyond their original wild type appearance, and are now the primary
type of show betta that is seen in pet stores. Sometimes called the “designer fish of the aquatic
world”, bettas have been bred over time into many colors and tail types.”

“The beautiful Betta splendens pet and show fish we see today were originally selectively bred
out of the wild types of Betta imbellis, Betta mahachaiensis, Betta samaragdina, and the
original Betta splendens.”

2 Biology and Ecology

Taxonomic Hierarchy and Taxonomic Standing
From ITIS (2019):

“Kingdom Animalia
Subkingdom Bilateria
Infrakingdom Deuterostomia
Phylum Chordata
Subphylum Vertebrata
Infraphylum Gnathostomata
Superclass Actinopterygii
Class Teleostei
Superorder Acanthopterygii
Order Perciformes
Suborder Anabantoidei
Family Osphronemidae
Subfamily Macropodinae
Genus Betta
Species Betta splendens Regan, 1910”

From Fricke et al. (2019):

“Current status: Valid as Betta splendens Regan 1910. Osphronemidae: Macropodusinae.”

Size, Weight, and Age Range

From Froese and Pauly (2019):

“Max length : 6.5 cm TL male/unsexed; [Rainboth 1996]; max. reported age: 2 years [Hugg
1996]”

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Environment
From Froese and Pauly (2019):

“Freshwater; benthopelagic; pH range: 6.0 - 8.0; dH range: 5 - 19. […] 24°C - 30°C [Riehl and
Baensch 1991; assumed to represent recommended aquarium temperature range]”

From Pleeging and Moons (2017):

“Jaroensutasinee and Jaroensutasinee (2001) found that the average water temperature in natural
betta habitats during breeding season is 29.9 °C ± 1.4, and that pH levels are acidic and range
from 5.28–5.80.”

“[…] bettas can survive in water with oxygen levels of as low as 0 - 2 ppm, while other fish die
at this concentration (Moore, 1942; Goldstein, 2004).”

Climate/Range
From Froese and Pauly (2019):

“Tropical; […] 22°N - 8°N, 99°E - 107°E”

Distribution Outside the United States

Native
From Nico and Neilson (2019):

“Tropical Asia. Native to Southeast Asia including the northern Malay Peninsula, central and
eastern Thailand, Kampuchea, and southern Vietnam (Witte and Schmidt 1992).”

From Vidthayanon (2013):

“Endemic to Thailand, from the Mae Khlong to Chao Phraya basins, the eastern slope of the
Cardamom mountains, and from the Isthmus of Kra.”

Froese and Pauly (2019) report B. splendens as native to Cambodia, Laos, Thailand, and
Vietnam.

Introduced
From Froese and Pauly (2019):

“Established in the Magdalena watershed [Colombia].”

“established, natural reproduction […] Species widespread in fish rearing facilities and has
presumably escaped into local waters [in Brazil].”

“Established in Jurong Lake and water bodies in the Sembawang area [Singapore].”

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“During hurricane 'David', B. splendens escaped, presumably from a tropical fish farm or a
private aquarist, into the polluted waters of the Rio Ozama [Dominican Republic] where they are
now abundant [Lever 1996].”

According to Froese and Pauly (2019), B. splendens has also been introduced to Spain, the
Philippines, Malaysia, Indonesia, and Canada; establishment in these locations has not been
confirmed.

From U.S. Fish and Wildlife Service (2007):

“LOCATION […] Thermal spring in Banff National Park, Alberta, Canada”

“EXOTIC SPECIES INTRODUCED INTO HABITAT […] Siamese fighting fish (Betta
splendens)”

Renaud and McAllister (1988) report that B. splendens was introduced into Banff National
Park’s Cave and Basin Hotsprings or nearby. The introduction occurred at some time prior to
1968; according to the authors, it is unknown when the species was last observed there.

From Hammer et al. (2019):

“The species has recently invaded the Adelaide River floodplain, near Darwin in northern
Australia. A very large and extensive population has persisted over consecutive years, with
potential for further spread.”

“[…] introductions have been recorded in at least 10 countries including USA, Brazil, Malaysia
and Singapore, however no major invasive populations have been noted (Froese and Pauly 2018;
Welcomme 1988).”

“The first verified record [of B. splendens in northern Australia] was reported in January 2010,
based on a photograph of a single individual female collected at Fogg Dam, a small modified
area of the Adelaide River floodplain […]. Subsequent anecdotal evidence suggests it may have
been present at the site as early as 2006. Limited searches in Fogg Dam using backpack
electrofishing and baited traps by NT Fisheries, in 2010 and 2011 respectively, were made in
order to evaluate the presence of B. splendens but failed to record any […]. Two individual fish
were again recorded at Fogg Dam through a community report in February 2011. A major shift
in detected abundance coincided with early monsoonal rains followed by moderate flooding in
January/February of 2014 (after two preceding years with below average rainfall and limited
wetland inundation) when large numbers of fish emerged and were readily detected in targeted
dip netting, visual searches and by community reports. […] Lower abundances were observed at
additional sites including a culvert at Beatrice Hill and a flooded car park at the boat ramp on the
Adelaide River proper […] The next period of flooding in the 2016/2017 wet season saw a mass
of fish at the Adelaide River main channel, with continuous chains of fish (around 100 per
minute) close to the shore swimming downstream and high numbers also in flooded edges […].
Two new detection sites were made east of the Adelaide River indicating further spread, namely
a wetland site monitored in 2014 and at Scott Creek via a community report which again has

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previously been heavily sampled for aquarium fish and bait. Using all recorded distribution
points […], the extent of occurrence of B. splendens as of February 2017 was 81 km2.”

Means of Introduction Outside the United States

From Froese and Pauly (2019):

“ornamental”

“Species widespread in fish rearing facilities and has presumably escaped into local waters [in
Colombia, Brazil, and the Dominican Republic].”

Wright (2002) reports that the introduction of Betta splendens to Alberta, Canada, was the result
of intentional release by a fish hobbyist.

From Hammer et al. (2019):

“No specific introduction vector was uncovered for B. splendens [in the Adelaide River
floodplain, Australia], and it is unclear exactly how long they have been present in the area, and
if fish were released at Fogg Dam or have dispersed there indirectly from an introduction
elsewhere on surrounding floodplains or semi-naturalised populations such as a ponds [sic]
(Lintermans 2004). Introductions of aquarium fishes are clearly ongoing in the region stemming
from Darwin into peri-urban surrounds, and education and response programs are vital for
limiting further incursions into remote habitats (Duggan et al. 2006).”

Short Description
From Regan (1910):

“Body moderately elongate, more or less compressed. Jaws with fixed conical teeth; palate
toothless. Praeorbital and opercular bones entire. Dorsal without or with a single spine and with
7 to 11 soft rays; origin above the anal; anal without or with 1 to 4 spines and with 20 to 37 soft
rays; pelvics inserted below or a little in advance of the pectorals, of a spine and 5 soft rays, the
outermost of which is more or less produced. Scales large, regularly arranged; lateral line
vestigial or absent.”

“Depth of body 2⅚ to 3⅔ in the length, length of head 3¼ to 3⅗. Snout as long as or shorter than
eye, the diameter of which is 3⅓ to 4 in the length of head; interorbital width 2⅖ to 3 in the
length of head. Maxillary extending to the vertical from between the nostrils. Dorsal I 8-9. Anal
II-IV 21-24. 30 to 32 scales in a longitudinal series. Dark greenish olive above, red below; all the
scales edged with black; a dark oblique stripe from eye to suboperculum; sometimes two dark
longitudinal bands, with a pale band between them, from eye to caudal fin; gill-membranes
blackish; dorsal rays black, membrane greenish with black undulating stripes; caudal rays red,
membrane greenish; pelvics and anal red, with dark edges; pectoral pale.”

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From Pleeging and Moons (2017):

“Wild fish have a brown-green color and both sexes are about 5 – 5.5 cm in size (Smith, 1945;
Jaroensutasinee and Jaroensutasinee, 2001). Domesticated bettas however, are 6 – 6.5 cm
(females are a little smaller), and male fish are bred in almost every imaginable color. Females
have less bright colors than males (Smith, 1945).”

From Snekser et al. (2006):

“B. splendens […] are distinctly sexually dimorphic: adult males are distinguished by larger
bodies and longer fin length from the smaller, shorter finned females (Bronstein and Jones-
Buxton, 1996; Jaroensutasinee and Jaroensutasinee, [2001]).”

Biology
From Snekser et al. (2006):

“Male and female Betta are aggressive towards each other (Goldstein, 1975) and in nature they
disperse to a density of 1.7 fish per m2 (Jaroensutasinee and Jaroensutasinee, [2001]). Male Betta
are well known for their elaborate and stereotyped aggressive displays, which include increased
tail beating, biting, and branchiostegal displaying (raising the opercula) (Simpson, 1968;
Bronstein, 1981, 1983, 1985; Halperin et al., 1998).”

“Male Betta establish territories in which they build bubble nests in which females lay their eggs,
and defend these territories against intrusion by other males (Robertson and Sale, 1974;
Bronstein, 1981). Males are typically more aggressive toward other males than to females
(Simpson, 1968; Robertson and Sale, 1974; Doutrelant et al., 2001), while the converse is true
for females (Braddock and Braddock, 1955; Simpson, 1968; Robertson, 1979; Bronstein and
Jones-Buxton, 1996).”

From Pleeging and Moons (2017):

“Their natural habitat exists of shallow ponds and rice paddy fields with plenty of vegetation.
This vegetation provides cover against fish-eating birds, like egrets, herons and kingfishers
(Jaroensutasinee and Jaroensutasinee, 2001). […] Bettas are carnivorous fish with a diet
consisting of mosquito larvae and other water insects, which form a source of protein and fat. It
is estimated that adult male fish eat about ten to fifteen thousand larvae per year (Smith, 1945;
Goldstein, 2004).”

“Bettas are a member of the family of the Anabantidae, known for their labyrinth organ, which is
a pharyngeal diverticulum that enables consumption of oxygen from the air (Liem, 1963; Kang
and Lee, 2010; Alton et al., 2012). […] The labyrinth organ is also responsible for the production
of air bubbles for nest building.”

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Human Uses
From Froese and Pauly (2019):

“Fisheries: of no interest; aquarium: highly commercial”

From Pleeging and Moons (2017):

“In Thailand, known for its ornamental fish production, Betta splendens represents 10% of the
annual fish export (Wiwatchaisaet, as cited in Meejui et al. 2005).”

“There are two varieties of domestic bettas, the betta bred for fighting and the ornamental variety
(Smith, 1945; Meejui et al., 2005). To the authors’ knowledge, only the ornamental variety with
long and colorful fins is kept in Europe.”

This species is common in trade in the United States. For example, ornamental varieties of Betta
splendens for sale at Petco (2019) include Baby Boy Betta, Baby Girl Betta, male Black Orchid
Betta, male Bumblebee Betta, male Butterfly Betta, male Cambodian Crowntail Betta, male and
female Crowntail Betta, male Copper Betta, male Deltatail Betta, male Doubletail Betta, male
Red Platinum Dragon, male Dragonscale Betta, male and female Elephant Ear Betta, male and
female Halfmoon Betta, male Halfmoon Doubletail Betta, male Halfmoon Doubletail Plakat
Betta, male Halfmoon King Betta, male King Betta, male and female Koi Betta, male Paradise
Betta, male Rose Gold Betta, male Rose Petal Betta, male and female Veiltail Betta, male
Cambodian Veiltail Betta, and male White Opal Betta.

Diseases
No OIE-reportable diseases have been documented for this species.

From Froese and Pauly (2019):

“Fin-rot Disease (late stage), Bacterial diseases

White spot Disease, Parasitic infestations (protozoa, worms, etc.)
Fin Rot (early stage), Bacterial diseases
Bacterial Infections (general), Bacterial diseases
Columnaris Disease (e.), Bacterial diseases
Fish tuberculosis (FishMB), Bacterial diseases
Velvet Disease 2 (Piscinoodinium sp.), Parasitic infestations (protozoa, worms, etc.)
Edwardsiellosis, Bacterial diseases”

From Pleeging and Moons (2017):

“‘Cotton wool’ disease, caused by Flavobacterium columnaris, is a bacterial disease, which is

especially pathogenic to betta species (Goldstein, 2004). Infected fish show ulcerative lesions
with mucus-like filamentous spots (Decostere [et al.], 1998; Goldstein, 2004).”

“Another condition considered to be important for bettas is ‘velvet’ disease (Goldstein, 2004).
This disease is caused by a dinoflagellate, Piscinoodinium spp. Young fry is especially sensitive

8
for infection and the development of clinical disease. The parasite is abundantly present and
flourishes in water of poor quality. Skin and gills become infected, resulting in hyperplasia,
hemorrhage, osmoregulatory compromise and necrosis (Roberts et al., 2009).”

“Finally, mycobacteriosis is the most common cause of death in bettas on breeding farms in
Thailand, and it also poses risks for humans. In fish, it is a slowly progressive disease
accompanied by the formation of granulomas in the liver, spleen and kidneys of infected fish
(Puttinaowarat, 1999; Zanoni et al., 2008). Other symptoms are extreme anorexia, exophthalmia,
keratitis and skeletal deformities (Zanoni et al., 2008; Chansue et al., 2009).”

Threat to Humans
From Froese and Pauly (2019):

“Harmless”

From Pleeging and Moons (2017):

“Finally, mycobacteriosis is the most common cause of death in bettas on breeding farms in
Thailand, and it also poses risks for humans. […] In humans, the main clinical signs of this
zoonotic disease are limited to skin lesions and ulcers (Gray et al., 1990; Speight and Williams,
1997).”

3 Impacts of Introductions
From Nico and Neilson (2019):

“The impacts of this species are currently unknown, as no studies have been done to determine
how it has affected ecosystems in the invaded range. The absence of data does not equate to lack
of effects. It does, however, mean that research is required to evaluate effects before conclusions
can be made.”

From Hammer et al. (2019):

“Betta splendens is rapidly expanding its distribution in the tropical wetlands of the Adelaide
River in the Northern Territory. The biological impacts of the invasion remain to be determined
and ecological studies are warranted to better understand these aspects.”

“Future ecological studies should be focused on potential impacts and understanding dispersal.
The introduced fish could be carriers of disease and parasites (Humphrey et al. 1986).
Aggressive interaction and competition for space and food is likely to occur between B.
splendens and small native fishes and tadpoles in local concentrated refuges, especially
considering the sheer noted abundance.”

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4 Global Distribution

Figure 1. Known global distribution of Betta splendens. Map from GBIF Secretariat (2019). The
following points were excluded from the extent of this map and from the climate matching
analysis: points in India and southeast Australia were excluded because they represent captive
specimens; a point in Russia was excluded due to coordinate error; points in the United States
(Florida), Colombia, and Taiwan were excluded because they do not represent confirmed
established populations of B. splendens.

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5 Distribution Within the United States

Figure 2. Known distribution of Betta splendens in the eastern United States. Map from Nico
and Neilson (2019). All points represent failed populations.

6 Climate Matching
Summary of Climate Matching Analysis
The Climate 6 score (Sanders et al. 2018; 16 climate variables; Euclidean distance) for the
contiguous United States was 0.0, indicating a low overall climate match. Climate 6 scores of
0.005 or below are classified as low match. There were some areas of medium climate match in
southern Florida and Texas, but the rest of the contiguous United States had a very low climate
match.

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Figure 3. RAMP (Sanders et al. 2018) source map showing weather stations in Southeast Asia
and Australia selected as source locations (red; Thailand, Vietnam, Laos, Cambodia, Myanmar,
and Australia) and non-source locations (gray) for Betta splendens climate matching. Source
locations from GBIF Secretariat (2019).

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Figure 4. Map of RAMP (Sanders et al. 2018) climate matches for Betta splendens in the
contiguous United States based on source locations reported by GBIF Secretariat (2019). Counts
of climate match scores are tabulated on the left. 0= Lowest match, 10= Highest match.

The “High”, “Medium”, and “Low” climate match categories are based on the following table:

Climate 6: Proportion of Climate Match

(Sum of Climate Scores 6-10) / (Sum of total Climate Scores) Category
0.000≤X≤0.005 Low
0.005<X<0.103 Medium
≥0.103 High

7 Certainty of Assessment
The biology and ecology of Betta splendens are well-documented. Although this species is
established outside of its native range, there is currently no research available assessing impacts
of its introduction. Further information is needed to adequately assess the risk this species poses
to the contiguous United States, so the certainty of this assessment is low.

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8 Risk Assessment
Summary of Risk to the Contiguous United States
Betta splendens, the Siamese Fighting Fish, is a small freshwater fish species native to Southeast
Asia. It is ubiquitous in the aquarium trade in the United States and globally. It has been
introduced to several countries outside of its native range, and it is currently established and
reproducing in Australia, Brazil, Colombia, the Dominican Republic, and Singapore. No research
has yet examined the impacts of these introductions, so the history of introduction is “none
documented.” B. splendens has a low overall climate match with the contiguous United States,
with areas of medium climate match in southern Florida and Texas. Further information is
needed to fully evaluate the potential risk this species poses to the contiguous United States, so
the certainty of this assessment is low and the overall risk assessment category is uncertain.

Assessment Elements
 History of Invasiveness (Sec. 3): None Documented
 Climate Match (Sec. 6): Low
 Certainty of Assessment (Sec. 7): Low
 Overall Risk Assessment Category: Uncertain

9 References
Note: The following references were accessed for this ERSS. References cited within quoted
text but not accessed are included below in Section 10.

Fricke, R., W. N. Eschmeyer, and R. van der Laan, editors. 2019. Catalog of fishes: genera,
species, references. Available:
http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp.
(March 2019).

Froese, R., and D. Pauly, editors. 2019. Betta splendens Regan, 1910. FishBase. Available:
https://www.fishbase.se/summary/Betta-splendens.html. (March 2019).

GBIF Secretariat. 2019. GBIF backbone taxonomy: Betta splendens (Regan, 1910). Global
Biodiversity Information Facility, Copenhagen. Available:
https://www.gbif.org/species/2393998. (March 2019).

Hammer, M. P., M. N. S. Simoes, E. W. Needham, D. N. Wilson, M. A. Barton, and D. Lonza.

2019. Establishment of Siamese Fighting Fish on the Adelaide River floodplain: the first
serious invasive fish in the Northern Territory, Australia. Biological Invasions
21(7):2269-2279.

International Betta Congress. 2018. About Betta splendens. Available:

https://www.ibcbettas.org/about-betta-splendens/. (June 2019).

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ITIS (Integrated Taxonomic Information System). 2019. Betta splendens (Regan, 1910).
Integrated Taxonomic Information System, Reston, Virginia. Available:
https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=172
611#null. (March 2019).

Nico, L., and M. Neilson. 2019. Betta splendens Regan, 1910. U.S. Geological Survey,
Nonindigenous Aquatic Species Database, Gainesville, Florida. Available:
https://nas.er.usgs.gov/queries/FactSheet.aspx?SpeciesID=326. (March 2019).

Petco. 2019. Live betta fish. Available:

https://www.petco.com/shop/en/petcostore/category/fish/live-fish/live-betta-fish. (June
2019).

Pleeging, C. C. F., and C. P. H. Moons. 2017. Potential welfare issues of the Siamese fighting
fish (Betta splendens) at the retailer and in the hobbyist aquarium. Vlaams
Diergeneeskundig Tijdschrift 86:213-223.

Regan, C. T. 1910. The Asiatic fishes of the family Anabantidae. Proceedings of the Zoological
Society of London B 1909(4):767-787.

Renaud, C. B., and D. E. McAllister. 1988. Taxonomic status of the extinct Banff longnose dace,
Rhinichthys cataractae smithi, of Banff National Park, Alberta. Environmental Biology
of Fishes 23(1-2):95-114.

Sanders, S., C. Castiglione, and M. H. Hoff. 2018. Risk Assessment Mapping Program: RAMP,
version 3.1. U.S. Fish and Wildlife Service.

Snekser, J. L., S. P. McRobert, and E. D. Clotfelter. 2006. Social partner preferences of male and
female fighting fish (Betta splendens). Behavioural Processes 72:38-41.

U.S. Fish and Wildlife Service. 2007. Kendall warm springs dace (Rhinichthys osculus
thermalis) 5-year review: summary and evaluation. U.S. Fish and Wildlife Service,
Cheyenne, Wyoming.

Vidthayanon, C. 2013. Betta splendens. The IUCN Red List of Threatened Species 2013:
e.T180889A7653828. Available: http://dx.doi.org/10.2305/IUCN.UK.2011-
1.RLTS.T180889A7653828.en. (March 2019).

Wright, D. 2002. Intentional introductions of alien species of fish: have we learned from our
mistakes?. Pages 201-217 in R. Claudi, P. Nantel, and E. Muckle-Jeffs, editors. Alien
invaders in Canada’s waters, wetlands, and forests. Canadian Forest Service, Science
Branch, Natural Resources Canada, Ottawa, Ontario.

15
10 References Quoted But Not Accessed
Note: The following references are cited within quoted text within this ERSS, but were not
accessed for its preparation. They are included here to provide the reader with more
information.

Alton, L. A., S. J. Portugal, and C. R. White. 2012. Balancing the competing requirements of air-
breathing and display behavior during male–male interactions in Siamese fighting fish
Betta splendens. Comparative Biochemistry and Physiology Part A: Molecular &
Integrative Physiology 164(2):363-367.

Braddock, J. C., and Z. I. Braddock. 1955. Aggressive behavior among females of Siamese
fighting fish, Betta splendens. Physiological Zoology 28:152-172.

Bronstein, P. M. 1981. Commitments to aggression & nest sites in male Betta splendens. Journal
of Comparative and Physiological Psychology 95:436-449.

Bronstein, P. M. 1983. Onset of combat in male Betta splendens. Journal of Comparative

Psychology 97:135-139.

Bronstein, P. M. 1985. Predictors of dominance in male Betta splendens. Journal of Comparative

Psychology 99:47-55.

Bronstein, P. M., and R. A. Jones-Buxton. 1996. Sensitization of escape in female Betta

splendens. Aggressive Behavior 22:431-435.

Chansue, N., A. Sermwatanakul, and K. Anekthanakul. 2009. Detection of Mycobacterium in

Betta splendens excreta by DNA analysis. The Annual Conference on Fisheries 2009,
Department of Fisheries, Bangkok, Thailand.

Courtenay, W. R., Jr., and D. A. Hensley. 1979. Survey of introduced non-native fishes. Phase I
report. Introduced exotic fishes in North America: status 1979. Report submitted to
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Florida.

Courtenay, W. R., Jr., D. A. Hensley, J. N. Taylor, and J. A. McCann. 1984. Distribution of

exotic fishes in the continental United States. Pages 41-77 in W. R. Courtenay, Jr., and J.
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Courtenay, W. R., Jr., H. F. Sahlman, W. W. Miley, II, and D. J. Herrema. 1974. Exotic fishes in
fresh and brackish waters of Florida. Biological Conservation 6(4):292-302.

Courtenay, W. R., Jr., and J. R. Stauffer, Jr.. 1990. The introduced fish problem and the
aquarium fish industry. Journal of the World Aquaculture Society 21(3):145-159.

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Decostere, A., F. Haesebrouck, and L. A. Devriese. 1998. Characterization of four
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Veterinary Microbiology 62:35-45.

Doutrelant, C., P. K. McGregor, and R. F. Oliveira. 2001. The effect of an audience on

intrasexual communication in male Siamese fighting fish, Betta splendens. Behavioral
Ecology 12:283–286.

Duggan, I. C., C. A. Rixon, and H. J. MacIsaac. 2006. Popularity and propagule pressure:
determinants of introduction and establishment of aquarium fish. Biological Invasions
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Froese, R., and D. Pauly, editors. 2018. FishBase. Available: http://www.fishbase.org.

Goldstein, S. R. 1975. Observations on the establishment of a stable community of adult male

and female Siamese fighting fish (Betta splendens). Animal Behavior 23:1179-1185.

Goldstein, R. 2004. The betta handbook. Barrons Educational Series Inc., New York.

Gray, S. F., R. S. Smith, N. J. Reynolds, and E. W. Williams. 1990. Fish tank granuloma. British
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Halperin, J. R. P., T. Giri, J. Elliott, and D. W. Dunham. 1998. Consequences of hyper-

aggressiveness in Siamese fighting fish: cheaters seldom prospered. Animal Behavior
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