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Redbelly Tilapia (Coptodon zillii) Ecological Risk Screening Summary

U.S. Fish and Wildlife Service, May 2019 Revised, September 2019 Web Version, 11/18/2019

Photo: J. Hoover, Waterways Experiment Station, U.S. Army Corp of Engineers. Public domain.

Available: https://nas.er.usgs.gov/queries/factsheet.aspx?SpeciesID=485. (May 2019).

1 Native Range and Status in the United States Native Range From Froese and Pauly (2019a):

“Africa and Eurasia: South Morocco, Sahara, Niger-Benue system, rivers Senegal, Sassandra,

Bandama, Boubo, Mé, Comoé, Bia, Ogun and Oshun, Volta system, Chad-Shari system [Teugels

and Thys van den Audenaerde 1991], middle Congo River basin in the Ubangi, Uele [Thys van

den Audenaerde 1964], Itimbiri, Aruwimi [Thys van den Audenaerde 1964; Decru 2015], Lindi-

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Tshopo [Decru 2015] and Wagenia Falls [Moelants 2015] in Democratic Republic of the Congo,

Lakes Albert [Thys van den Audenaerde 1964] and Turkana, Nile system and Jordan system

[Teugels and Thys van den Audenaerde 1991].”

Froese and Pauly (2019a) list the following countries as part of the native range of Coptodon

zillii: Algeria, Benin, Cameroon, Central African Republic, Chad, Democratic Republic of the

Congo, Egypt, Ghana, Guinea, Guinea-Bissau, Israel, Ivory Coast, Jordan, Kenya, Lebanon,

Liberia, Mali, Mauritania, Morocco, Niger, Nigeria, Senegal, Sierra Leone, Sudan, Togo,

Tunisia, Uganda, and Western Sahara.

Status in the United States From NatureServe (2019):

“Introduced and established in ponds and other waters in Maricopa County, Arizona; irrigation

canals in Coachella, Imperial, and Palo Verde valleys, California; and headwater springs of San

Antonio River, Bexar County, Texas; common (Page and Burr 1991). Established also in the

Carolinas, Hawaii, and possibly in Florida and Nevada (Robins et al. 1991).”

From Nico et al. (2019):

“Established or locally established in southern Arizona and California, Hawaii, North and South

Carolina, and Texas; reported from several other states. Considered eradicated from all sites in

Florida (Smith-Vaniz, personal communication) and Nevada (Courtenay et al. 1984, 1986). In

California, Oreochromis mossambicus has largely replaced T. zillii in the Salton Sea (and

possibly in coastal southern California) (Swift et al. 1993).”

“Populations introduced into Alabama normally did not survive the winter and required annual restocking (Smith-Vaniz 1968). However, their tolerance to cold temperatures in central

California prompted officials to place the species on the prohibited list for portions of the state

(Shapovalov et al. 1981).”

Very little information is available about current use of C. zillii in the United States. It does not

appear to be common in aquaculture and is not available from major online aquarium and

aquaponics retailers.

From Chapman (2018):

“In the United States, commercial culture of tilapia is concentrated in Arizona, California, and

Florida. It is not clear, however, what species of tilapia are under cultivation. A collection of

hybrid stocks currently constitute the bulk of the commercial production. The hybrids under

cultivation are female mouth-brooders and believed to have originated from genetic crosses of

predominantly blue tilapia (O. aureus) and ancillary O. niloticus, O. mossambicus, and O.

hornorum species. Some evidence of genes from T. rendalli and S. melanotheron are also

apparent.”

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Most States regulate aquaculture species or facilities. The following information on State

regulations is not exhaustive, and focuses on regulations that broadly prohibit importation,

transport, or possession of C. zillii or the genus or family to which it belongs.

Florida Fish and Wildlife Conservation Commission (2019) lists “Tilapia [now Coptodon] zillii

(Redbelly tilapia)” as a Prohibited Nonnative Species.

From Texas Parks and Wildlife (1999):

“The organisms listed here are legally classified as exotic, harmful, or potentially harmful. No

person may possess or place them into water of this state except as authorized by the department

[that is, Texas Parks and Wildlife].”

“Tilapia, Family Cichlidae

All species of genera Tilapia, Oreochromis and Sarotherodon

Examples: Tilapia [now Coptodon] zillii (Redbelly tilapia), […]”

“Note: Reclassification of a species does not remove the species from the prohibited list.”

From Aquatic Network (2019):

“No tilapia farming, or finfish farming at all is allowed in Alaska.”

“Currently, tilapia zilli, tilapia hornorum, and mossambica are the only species of tilapia allowed

to be cultured in California and they are allowed only in the following counties: Riverside, Los

Angeles, Orange, San Bernardino, San Diego, and Imperial. Please note that you must first

obtain a Restricted Species Permit to possess, import, and transport tilapia [now Coptodon] zilli

[sic].”

“According to the Illinois Department of Natural Resources:

‘It is unlawful for any person(s) to stock Tilapia species in Illinois waters. Tilapia are not on the

Aquatic Life Approved Species List, but are allowed for aquaculture with an Aquaculture Permit

and a Letter of Authorization to possess Tilapia. Aquatic Life Dealers may keep live Tilapia for

sale at a market so long as all fish are dead upon sale.’”

“Tilapia are a restricted species in Nevada and can only be possessed alive in the state for

purposes of commercial aquaculture, or certain other purposes. This requires a Commercial

Possession of Live Wildlife License from the Department of Wildlife. Private possession of live

tilapia for home or personal use is prohibited.”

“The Utah Division of Wildlife Resources (UDWR) has jurisdiction over the species of game

fish one may possess in Utah. In other words, the UDWR must approve any species of game fish

that a private operator may possess in Utah. Their ‘Collection, Importation and Possession of

Zoological Animals’ regulation has classified Tilapia as a nuisance species and it does not allow

importation or possession of the species in Utah.”

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Means of Introductions in the United States From Nico et al. (2019):

“Introduced in most locations by state agencies, universities, or private companies for control of

aquatic plants, to control mosquitoes and chironomid midges, as forage or food fish, and for

aquaculture evaluation (Minckley 1973; Legner and Pelsue 1977; Lee et al. 1980 et seq.;

Shapovalov et al. 1981; Grabowski et al. 1984; McGowan 1988; Courtenay and Robins 1989;

Page and Burr 1991). For example, the species has been stocked into various North Carolina

waters for aquatic plant control by Texas Gulf, Inc., and also by Carolina Power and Light

Company (Courtenay et al. 1986; J. Crutchfield, personal communication). There have been both

authorized as well as illegal releases. Introductions into Dade County, Florida, probably resulted

from escapes from nearby fish farms or aquarium releases (Hogg 1976a, b). Redbelly tilapia and

blue tilapia were inadvertently introduced into Hyco Reservoir in North Carolina in 1984 after a

small number of fish escaped from a holding cage located in the heated discharge area during an

on-site agricultural study (Crutchfield 1995).”

Remarks A previous version of this ERSS was finalized in 2014 under the name Tilapia zillii. Because the

name T. zillii is still commonly used in literature, it was also used when researching in

preparation of this report.

From CABI (2019):

“Tilapia zillii was first described by Gervais in 1848 and was given many synonyms throughout

the nineteenth and twentieth century. Some sources recognize Coptodon zillii as the accepted

name for the species following a molecular phylogenetic study by Dunz and Schliewen (2013).”

From GISD (2019):

“It seems to be a common mistake to omit one of the i's at the end of the word zillii, often

incorrectly being spelled T. zilli.”

2 Biology and Ecology Taxonomic Hierarchy and Taxonomic Standing From Froese and Pauly (2019b):

“Biota > Animalia (Kingdom) > Chordata (Phylum) > Vertebrata (Subphylum) > Gnathostomata

(Superclass) > Pisces (Superclass) > Actinopterygii (Class) > Perciformes (Order) > Labroidei

(Suborder) > Cichlidae (Family) > Pseudocrenilabrinae (Subfamily) > Coptodon (Genus) >

Coptodon zillii (Species)”

From Fricke et al. (2019):

“Current status: Valid as Coptodon zillii (Gervais 1848). Cichlidae: Pseudocrenilabrinae.”

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Size, Weight, and Age Range From Froese and Pauly (2019a):

“Maturity: Lm 7.0, range 20 - ? cm

Max length : 40.0 cm SL male/unsexed; [van Oijen 1995]; common length : 30.0 cm SL

male/unsexed; [van Oijen 1995]; max. published weight: 300.00 g [Ita 1984]; max. reported age:

7 years [Noakes and Balon 1982]”

Environment From Froese and Pauly (2019a):

“Freshwater; brackish; benthopelagic; pH range: 6.0 - 9.0; dH range: 5 - 20; potamodromous

[Riede 2004]; depth range 1 - 7 m [Eccles 1992].”

“Highly adaptable and tolerates varying water qualities [Lamboj 2004]; they are tolerant of a

wide range of temperatures and salinities [Genner et al. 2018], even surviving marine conditions

[Lamboj 2004]. Extended temperature range 6.5 - 42.5 °C, natural temperature range 10.5 - 36°C

[Philippart and Ruwet 1982].”

From GISD (2019):

“While temperatures between 20 to 32 degrees Celsius are optimal for T. zillii, it can tolerate

temperatures between 11 to 36 degrees Celsius, becoming lethargic and vulnerable to predators

and disease below 16 degrees Celsius. Mostly occurring in fresh water, T. zillii are often found in

brackish waters and has occasionally been reported to be found in marine waters; tolerating

salinity levels of up to 29-45 ppt (Costa-Pierce, 2003; FishBase, 2008; GSMFC, 2005).”

From Nico et al. (2019):

“Redbelly tilapia is extremely tolerant of saline conditions, with survival and growth occurring in

salinities up to 40‰ and reproduction occurring through 29‰ (Stickney 1986).”

Climate/Range From Froese and Pauly (2019a):

“Tropical; […] 35°N - 9°S, 17°W - 36°E”

Distribution Outside the United States Native From Froese and Pauly (2019a):

“Africa and Eurasia: South Morocco, Sahara, Niger-Benue system, rivers Senegal, Sassandra,

Bandama, Boubo, Mé, Comoé, Bia, Ogun and Oshun, Volta system, Chad-Shari system [Teugels

and Thys van den Audenaerde 1991], middle Congo River basin in the Ubangi, Uele [Thys van

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den Audenaerde 1964], Itimbiri, Aruwimi [Thys van den Audenaerde 1964; Decru 2015], Lindi-

Tshopo [Decru 2015] and Wagenia Falls [Moelants 2015] in Democratic Republic of the Congo,

Lakes Albert [Thys van den Audenaerde 1964] and Turkana, Nile system and Jordan system

[Teugels and Thys van den Audenaerde 1991].”

Froese and Pauly (2019a) list the following countries as part of the native range of Coptodon

zillii: Algeria, Benin, Cameroon, Central African Republic, Chad, Democratic Republic of the

Congo, Egypt, Ghana, Guinea, Guinea-Bissau, Israel, Ivory Coast, Jordan, Kenya, Lebanon,

Liberia, Mali, Mauritania, Morocco, Niger, Nigeria, Senegal, Sierra Leone, Sudan, Togo,

Tunisia, Uganda, and Western Sahara.

Introduced

From CABI (2019):

“Its non-native distribution includes Antigua and Barbuda, Eritrea, Ethiopia, Guam, Iran, Japan,

Madagascar, Mauritius, Mexico, New Caledonia, Philippines, Saudi Arabia, Russia, Sri Lanka,

Syria, Taiwan, Tanzania, Turkey, UK, […] Australia, Fiji, […] and Thailand (Froese and Pauly,

2014).”

From Gu et al. (2018):

“To date, five tilapia species (O. aureus, O. niloticus, Tilapia zillii, S. galilaeus, and a hybrid of

O. mossambicus and O. niloticus) have been identified in the rivers of South China (Gu et al.

2016). Among these species, O. niloticus and T. zillii are most common, and they were both

introduced to mainland China in 1978 and widely distributed in the study area (Gu et al. 2016,

2018).”

“Redbelly tilapia (Tilapia zillii) […] can tolerate colder waters and […] natural populations have

developed in many rivers in South China (He et al. 2013).”

According to Froese and Pauly (2019a), C. zillii is established outside its native range in the

following countries: Antigua, Ethiopia, Guam, Iraq, Israel, Japan, Jordan, Kenya, Madagascar,

Mauritius, Mexico, New Caledonia, Saudi Arabia, Tanzania, Thailand, Turkey, and Uganda.

According to Froese and Pauly (2019a), C. zillii has been introduced but establishment has not

been confirmed in the following countries: Australia, China, Eritrea, Fiji, Iran, Ivory Coast,

Libya, Malaysia, Northern Marianas, Philippines, Russia, Singapore, South Africa, Sri Lanka,

Syria, Taiwan, and the United Kingdom.

Means of Introduction Outside the United States From Khaefi et al. (2014):

“According to local fishermen, T. zillii appeared in this wetland [in Iran] in April or May 2012,

and now it is available in daily catches. According to them, it was deliberately introduced to the

Shadegan wetland to enhance fish production.”

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From Ogutu-Ohwayo (1990):

“[…] Tilapia zillii (Gervais), were introduced into lakes Victoria and Kyoga [East Africa] during

the 1950s and early 1960s. […] T. zillii was introduced to feed on macrophytes which were not

being utilized by other commercially important fishes.”

Short Description From Froese and Pauly (2019a):

“Dorsal spines (total): 13 - 16; Dorsal soft rays (total): 10-14; Anal spines: 3; Anal soft rays: 8 -

10. Diagnosis: A large, deep-bodied species with a narrow head and small strong jaws; generally

has a bright red belly and prominent vertical barring [Genner et al. 2018]. Upper profile of head

not convex; lower pharyngeal bone about as long as broad, and with anterior lamella shorter than

toothed area; median pharyngeal teeth not broadened; dorsal fin with 14-16 spines and 10-14 soft

rays; 8-11 lower gillrakers; dark longitudinal band appears on flanks when agitated; no

bifurcated dark vertical bars on flanks; dorsal and caudal fins not or feebly blotched [Teugels and

Thys van den Audenaerde 2003]. Body brownish-olivaceous with an iridescent blue sheen; lips

bright green [van Oijen 1995; Teugels and Thys van den Audenaerde 2003]. Chest pinkish [van

Oijen 1995]. Dorsal, caudal and anal fins brownish-olivaceous with yellow spots, dorsal and anal

fins outlined by narrow orange band; "tilapian" spot large, extending from last spine to 4th soft

ray and always bordered by yellow band [van Oijen 1995; Teugels and Thys van den

Audenaerde 2003]. Specimens of 2-14 cm standard length with completely yellowish or greyish

caudal fin without dots, but tend to develop a greyish caudal fin with dots of increasing size

during development; above 14 cm standard length, this species has greyish caudal fins with dots

on entire caudal fin [Nobah et al. 2006]. The sexes look very similar, although in a mated pair

the male is usually larger [Genner et al. 2018]. Difficult to distinguish reliably from Coptodon

rendalli, but C. zillii can have a less-steep head profile and more prominent vertical bars; in East Africa, the tailfin of C. rendalli is often divided into a brown/grey upper part and red/yellowish

lower part, whereas the tail of C. zillii is more uniform throughout [Genner et al. 2018].”

Biology From Nico et al. (2019):

“Redbelly tilapia is primarily herbivorous, with aquatic macrophytes, algae, and diatoms

generally comprising >80% of its diet and the remainder including aquatic insects and

crustaceans and fish eggs. Proportion of diet from animal sources is generally size-related, with

larger fish consuming more animal-based food items (Khallaf and Alne-na-ei 1987). […] This

species is a substrate spawner, with fishes forming monogamous pairs and exhibiting biparental

guarding behavior. Nests are primarily small, saucer-shaped depressions in the substrate, but

show some variation in morphology due to environmental conditions (Bruton and Gophen 1992).

Breeding season is dependent on climate, with warm, temperature-stable equatorial populations

breeding year-round, and those in areas with more defined seasons breeding during summer

months (Siddiqui 1979; Bruton and Gophen 1992).”

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Human Uses From CABI (2019):

“Redbelly tilapia is an economically important food fish and important to aquaculture and

commercial aquarium trade in its native range (Mehanna, 2004).”

“It provides up to 70% of Egypt’s fish production and is a hardy species, easy to grow and

popular with consumers (white-fleshed and mild-flavoured) (Canonico et al., 2005).”

“It is also an important fish species for recreational fishery (ISSG, 2014). In addition to its value

for commercial fishermen, recreational fishing and tourism may create a demand not only for

food, accommodation and transportation but also for related recreational activities such as

camping, boating, etc. All of these activities may provide economic incomes.”

“Redbelly tilapia is used for controlling species of aquatic plants. It was determined that Chara

sp. and Najasmarina [sic] could be controlled by redbelly tilapia in small lakes and ponds

(Saeed, 1986). It has also been used to control noxious aquatic insects, mosquitos and chrinomid

[sic] midges (Molnar [et al.], 2008).”

Diseases No OIE-reportable diseases (OIE 2019) have been documented for this species.

Poelen et al. (2014) lists the following as parasites of Coptodon zillii:

Acanthogyrus tilapiae, Euclinostomum heterostomum, Posthodiplostomum sp., Amirthalingamia

macracantha, Cichildogyrus tilapiae, Polyacanthorhynchus kenyensis, Cichlidogyrus

amphoratus, Cichlidogyrus digitatus, Cichlidogyrus yanni, Argulus sp., Porrocaecum sp., and

Strigeidae sp. (Strona et al. 2013, Benesh et al. 2017, Smithsonian Institution no date).

From CABI (2019):

“Redbelly tilapia may be infected with a wide range of diseases and parasites, including

Diplozoon paradoxum and Tetraonchus species (Yildirim et al., 2010).”

Threat to Humans From Froese and Pauly (2019a):

“Potential pest”

3 Impacts of Introductions From Nico et al. (2019):

“Juvenile T. zillii were implicated in population declines of desert pupfish Cyprinodon

macularius inhabiting shallow irrigation canals near the Salton Sea, California (Lee et al. 1980 et

seq.; Schoenherr 1985). In Florida, this species was found to be highly aggressive; it is

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considered a serious threat to native aquatic plants and to fish that rely on plants for cover,

foraging, or spawning sites (Courtenay et al. 1974).”

From Crutchfield et al. (1992):

“Redbelly tilapia (Tilapia zilli Gervais) rapidly established a reproducing population in a North

Carolina power plant cooling reservoir after inadvertent introduction in 1984. It eliminated the

submersed macrophyte community, including a 57-ha infestation of Egeria densa (Planch.)

[nonnative] by late 1985.”

From Andreu-Soler and Ruiz-Campos (2013):

“We assessed the effects of the abundance and biomass of four exotic fishes (common carp,

Cyprinus carpio; guppy, Poecilia reticulata; green swordtail, Xiphophorus hellerii; and redbelly

tilapia, Tilapia cf. zillii) on the residual somatic condition (Kr) of endangered Baja California

killifish (Fundulus lima) in two oasis systems of Baja California Sur, Mexico. We used multiple

regressions to analyze relationships between Kr of the killifish and 21 ecological variables.

Biomass of redbelly tilapia and common carp were variables that better explained variation in Kr

among populations of killifish. In both drainages, redbelly tilapia was the dominant fish, which

relegated smaller habitat units to the other coexisting species of fishes, increasing competition

among them and decreasing Kr and abundance of the endemic killifish.”

Most States regulate aquaculture species or facilities. The following States (not an exhaustive

list) have regulations that broadly prohibit importation, transport, or possession of C. zillii:

Alaska, California, Florida, Illinois, Nevada, Texas, and Utah (Texas Parks and Wildlife 1999;

Aquatic Network 2019; Florida Fish and Wildlife Conservation Commission 2019).

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4 Global Distribution

Figure 1. Known global distribution of Coptodon zillii. Map from GBIF Secretariat (2019).

Points in Angola, Australia, Botswana, Eritrea, Guyana, Mozambique, Réunion, the Seychelles,

South Africa, Syria, the United Kingdom, Zambia were excluded from climate matching because

the establishment status of C. zillii has not been confirmed in these countries. Points in the

contiguous United States that were inconsistent with the established U.S. range of C. zillii as

reported by Nico et al. (2019; Figure 2) were also excluded from climate matching analysis. No

occurrence data were available for parts of the species established range in Antigua, Iraq, New

Caledonia, Saudi Arabia, and Turkey.

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5 Distribution Within the United States

Figure 2. Known distribution of Coptodon zillii in the United States. Map from Nico et al.

(2019). Yellow diamonds represent established occurrences; orange diamonds represent all other

status designations (failed, collected, extirpated, unknown, or stocked). Point in Idaho was

excluded from climate matching analysis because it is located in a geothermal water body.

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 0.347, indicating a high overall climate match. (A Climate 6 score

of 0.103 or greater is classified as high.) The climate match was high in peninsular Florida, the

Mid-Atlantic Region, much of Texas, the Southwest, California, and scattered areas in the

Pacific Northwest. Most of the rest of the contiguous United States had a medium climate match.

There were areas of low climate match in the Pacific Northwest, New England, interior

Southeast, and the north-central United States from the western Great Lakes to the eastern Rocky

Mountains. Individual States had medium or high climate scores except for the following States

that had low climate scores: Alabama, Colorado, Iowa, Maine, Michigan, Minnesota,

Mississippi, North Dakota, Nebraska, New Hampshire, Rhode Island, South Dakota, Vermont,

Wisconsin, and Wyoming.

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Figure 3. RAMP (Sanders et al. 2018) source map showing weather stations selected as source

locations (red; southern United States and Hawaii; Mexico; West, Central, and East Africa

including Madagascar; scattered locations in North Africa; Israel; Lebanon; Jordan; Thailand;

China; Japan) and non-source locations (gray) for Coptodon zillii climate matching. Source

locations from GBIF Secretariat (2019) and Nico et al. (2019).

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Figure 4. Map of RAMP (Sanders et al. 2018) climate matches for Coptodon zillii in the

contiguous United States based on source locations reported by GBIF Secretariat (2019) and

Nico et al. (2019). 0 = Lowest match, 10 = Highest match.

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

Climate 6: Proportion of

(Sum of Climate Scores 6-10) / (Sum of total Climate Scores)

Climate Match

Category

0.000≤X≤0.005 Low

0.005<X<0.103 Medium

≥0.103 High

7 Certainty of Assessment There is adequate information available about the biology and distribution of Coptodon zillii. Its

invasion history, both in the United States and globally, has been well-documented. Negative

impacts of this species’ introduction have been reported from multiple geographic locations in

multiple studies in the scientific literature. Certainty of this assessment is high.

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8 Risk Assessment Summary of Risk to the Contiguous United States Coptodon zillii, the Redbelly Tilapia, is a fish species native to Africa and the Middle East. It is

adaptable to a wide range of environments and has been introduced in many countries for

aquaculture, recreational fishing, forage, and control of aquatic macrophyte or insects. C. zillii is

also used for commercial fishing and in the aquarium trade. Many of these introductions have

resulted in population establishment. C. zillii is not heavily in use in the United States currently,

but it has become established in the southern United States through a history of deliberate release

and accidental escape from captivity. Several States regulate importation, transport, or

possession of C. zillii. Declines of native fish species, such as the endangered Baja California

Killifish and the desert pupfish Cyprinodon macularius, have been attributed to C. zillii. C. zillii

also eliminated the aquatic macrophyte community of the Hyco Reservoir in North Carolina.

History of invasiveness is high. This species has a high climate match with the contiguous

United States, especially in the southern United States where it is already established. Certainty

of this assessment is high because negative impacts of this species’ introduction have been

clearly documented in the scientific literature in multiple locations. The overall risk assessment

category is high.

Assessment Elements History of Invasiveness (Sec. 3): High

Climate Match (Sec. 6): High

Certainty of Assessment (Sec. 7): High

Overall Risk Assessment Category: High

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.

Andreu-Soler, A., and G. Ruiz-Campos. 2013. Effects of exotic fishes on the somatic condition

of the endangered Killifish Fundulus lima (Teleostei: Fundulidae) in oases of Baja

California Sur, Mexico. The Southwestern Naturalist 58(2):192-201.

Aquatic Network. 2019. United States aquaculture by regions. Available:

https://www.aquanet.com/us-regulations. (September 2019).

CABI. 2019. Tilapia zillii (redbelly tilapia) [original text by A. S. Tarkan]. In Invasive Species

Compendium. CAB International, Wallingford, U.K. Available:

https://www.cabi.org/ISC/datasheet/61147. (May 2019).

Chapman, F. A. 2018. Culture of hybrid tilapia: a reference profile. CIR1051. University of

Florida/Institute of Food and Agricultural Sciences, Gainesville, Florida.

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Crutchfield, J. U., Jr., D. H. Schiller, D. D. Herlong, and M. A. Mallin. 1992. Establishment and

impact of redbelly tilapia in a vegetated cooling reservoir. Journal of Aquatic Plant

Management 30:28-35.

Florida Fish and Wildlife Conservation Commission. 2019. Prohibited nonnative species list.

Florida Fish and Wildlife Conservation Commission, Tallahassee. Available:

https://myfwc.com/wildlifehabitats/nonnatives/prohibited-species-list/. (May 2019).

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.

(May 2019).

Froese, R., and D. Pauly, editors. 2019a. Coptodon zillii Gervais, 1848. FishBase. Available:

https://www.fishbase.de/summary/Coptodon-zillii.html. (May 2019).

Froese, R., and D. Pauly, editors. 2019b. Coptodon zillii (Gervais, 1848). FishBase. In World

Register of Marine Species. Available:

http://www.marinespecies.org/aphia.php?p=taxdetails&id=317356. (September 2019).

GBIF Secretariat. 2019. GBIF backbone taxonomy: Coptodon zillii (Gervais, 1848). Global

Biodiversity Information Facility, Copenhagen. Available:

https://www.gbif.org/species/2370703. (September 2019).

GISD (Global Invasive Species Database). 2019. Species profile: Tilapia zillii. Invasive Species

Specialist Group, Gland, Switzerland. Available:

http://www.iucngisd.org/gisd/speciesname/Tilapia+zillii. (May 2019).

Gu, D. E., F. D. Yu, M. Xu, H. Wei, X. D. Mu, D. Luo, Y. X. Yang, Z. Pan, and Y. C. Hu. 2018.

Temperature effects on the distribution of two invasive tilapia species (Tilapia zillii and

Oreochromis niloticus) in the rivers of South China. Journal of Freshwater Ecology

33(1):511-524.

Khaefi, R., H. R. Esmaeili, H. Zareian, and S. Babaei. 2014. The first record of the redbelly

tilapia, Tilapia zillii (Gervais, 1848), in freshwaters of Iran. Turkish Journal of Zoology

38:96-98.

NatureServe. 2019. NatureServe Explorer: an online encyclopedia of life, version 7.1.

NatureServe, Arlington, Virginia. Available: http://explorer.natureserve.org. (May 2019).

Nico, L., M. Neilson, and B. Loftus. 2019. Tilapia zillii (Gervais, 1848). U.S. Geological Survey,

Nonindigenous Aquatic Species Database, Gainesville, Florida. Available:

https://nas.er.usgs.gov/queries/factsheet.aspx?SpeciesID=485. (May 2019).

Ogutu-Ohwayo, R. 1990. The decline of the native fishes of lakes Victoria and Kyoga (East

Africa) and the impact of introduced species, especially the Nile perch, Lates niloticus,

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and the Nile tilapia, Oreochromis niloticus. Environmental Biology of Fishes 27(2):81-

96.

OIE (World Organisation for Animal Health). 2019. OIE-listed diseases, infections and

infestations in force in 2019. World Organisation for Animal Health, Paris. Available:

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