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Redbreast ( rendalli) Ecological Risk Screening Summary

U.S. and Wildlife Service, 2012 Revised, May and July 2019 Web Version, 9/18/2019

Photo: W. Uys. Licensed under CC BY-NC 4.0. Available: https://www.inaturalist.org/photos/15162853. (May 2019).

1 Native Range and Status in the United States Native Range From Nico et al. (2019):

“Tropical and subtropical Africa. Kasai drainage, upper Congo River, Cunene, Okavango, Limpopo and system, Tanganyika and , east coastal rivers south to Phongolo, and coastal lakes to Sibaya; also occurs in estuaries in and Natal (Thys van den Audenaerde 1964; de Moor and Bruton 1988; Teugels et al. 1991; Skelton 1993).”

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From Konings et al. (2018):

“In central Africa, Coptodon rendalli is naturally known from the Upper Congo basin (Katanga region and Lualaba River) down to Kisangani (Stanleyville) and even to Isangi (Central Congo basin) [Democratic Republic of the Congo].”

“In eastern Africa, this is native to the and Malawi Basins [Burundi, , Democratic Republic of the Congo, , Malawi, Mozambique] and Lakes Chilwa and Chiuta [Malawi, Mozambique] (Tweddle 1979, 1983) and the Shire River (Malawi) (Tweddle and Willoughby 1979).”

“In southern Africa, this species is native to the Cunene, Okavango, Zambezi systems including [Tanzania, Malawi, Mozambique, Zambia, , , Angola, Namibia], and east coastal rivers south to the Phongolo and coastal lakes to Lake Sibaya (Skelton 2001) [Mozambique, , ], as well as estuaries in Mozambique and KwaZulu- Natal [South Africa]. It is found in all Zimbabwean rivers although it was probably absent originally from the upper Save-Runde and scarce on the highveld (Junor 1969).”

“Native: Angola; Botswana; Burundi; Congo, The Democratic Republic of the; Eswatini; Malawi; Mozambique; Namibia (Caprivi Strip); South Africa (Gauteng, KwaZulu-Natal, Limpopo Province, Mpumalanga, North-West Province); Tanzania, United Republic of; Zambia; Zimbabwe”

Status in the United States From Nico et al. (2019):

“Although this species was formerly considered established and spreading in Hawaii (Maciolek 1984), or at least established (Courtenay et al. 1991), its recent status in the state is considered uncertain (Devick 1991a). Established in Puerto Rico.”

Wu and Yang (2012) confirmed establishment of T. rendalli in a reservoir on the Hawaiian island of Oahu through genetic analysis.

From Wu and Yang (2012):

"The results of phylogenetic tree analysis demonstrated that the populations collected from the wild reservoirs included two different species, S. melanotheron and T. rendalli [...]"

There is no indication that this species is in trade in the United States.

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

“This species was brought to Hawaii as a shipment of 52 fish from Africa in 1957; these fish were bred in tanks by the state, and resulting offspring were stocked in Wahiawa Reservoir in 1958 and 1959 and on Maui in 1959. According to Devick (1991b), these intentional 2

introductions were undertaken for aquaculture (aku bait) and weed control. The [sic] were intentionally introduced into Puerto Rico for week [sic] control and angling.”

Remarks A previous version of this ERSS was published in 2012 under the formerly accepted scientific name Tilapia rendalli. Because the acceptance of C. rendalli as the valid scientific name occurred within the past decade and the name T. rendalli is still commonly used in literature, both scientific names were used as search terms to find information for this report.

From Nico et al. (2019):

“Identification of this species in the United States has been problematic and some reports in the literature on nonindigenous species may be misidentifications (Lee et al. 1980 et seq.) or hybrids (Courtenay et al. 1984; Taylor et al. 1986; Howells 1991b). is nearly identical to Tilapia zillii, and the two species frequently hybridize; many reports or specimens of T. rendalli may potentially be T. zillii or their hybrids. Both redbreast and redbelly tilapia are superficially similar to another North American introduced , Tilapia mariae: spotted tilapia lacks the red ventral coloration present in T. rendalli, has lateral vertical stripes that extend onto the , and 5-6 square black blotches along the side (lacking in T. rendalli).”

“This species is referenced as T. melanopleura in previous reports concerning introduced species (e.g., Maciolek 1984; Devick 1991a, 1991b). A report of T. melanopleura in Alabama by Smith- Vaniz (1968) was based on incorrect identifications of fish later shown to be T. zillii (Smith- Vaniz, personal communication). Barrett (1983) tentatively re-identified some California specimens, previously reported to be T. zillii, as T. rendalli; he based his determination on various morphological characters and color patterns. These fish were taken from near Blythe, Riverside County.”

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

“Biota > Animalia (Kingdom) > Chordata (Phylum) > Vertebrata (Subphylum) > Gnathostomata (Superclass) > Pisces (Superclass) > (Class) > Perciformes (Order) > Labroidei (Suborder) > Cichlidae (Family) > (Subfamily) > Coptodon () > Coptodon rendalli (Species)”

“Status accepted”

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From Fricke et al. (2019):

“Current status: Valid as Coptodon rendalli (Boulenger 1897). Cichlidae: Pseudocrenilabrinae.”

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

“Maturity: Lm 17.7 range ? - ? cm Max length : 45.0 cm TL male/unsexed; [Anonymous 1994]; max. published weight: 2.5 kg [Anonymous 1994]; max. reported age: 7 years [Baensch and Riehl 1991]”

Environment From Froese and Pauly (2019a):

“Freshwater; brackish; benthopelagic; depth range 3 - 8 m [Mundy 2005]. […] 24°C - 28°C [Baensch and Riehl 1991; assumed to be recommended aquarium water temperature range]”

“They are tolerant of a wide range of temperatures (8-41°C) and salinities [Philippart and Ruwet 1982; Skelton 1993; Genner et al. 2018].”

From Konings et al. (2018):

“This species is tolerant of a wide range of temperatures (11–37°C) and salinity to 19 parts per thousand.”

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

“Tropical; […] 20°N - 20°S”

Distribution Outside the United States Native From Nico et al. (2019):

“Tropical and subtropical Africa. Kasai drainage, upper Congo River, Cunene, Okavango, Limpopo and Zambezi system, Lakes Tanganyika and Malawi, east coastal rivers south to Phongolo, and coastal lakes to Lake Sibaya; also occurs in estuaries in Mozambique and Natal (Thys van den Audenaerde 1964; de Moor and Bruton 1988; Teugels et al. 1991; Skelton 1993).”

From Konings et al. (2018):

“In central Africa, Coptodon rendalli is naturally known from the Upper Congo basin (Katanga region and Lualaba River) down to Kisangani (Stanleyville) and even to Isangi (Central Congo basin) [Democratic Republic of the Congo].”

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“In eastern Africa, this species is native to the Lake Tanganyika and Malawi Basins [Burundi, Tanzania, Democratic Republic of the Congo, Zambia, Malawi, Mozambique] and Lakes Chilwa and Chiuta [Malawi, Mozambique] (Tweddle 1979, 1983) and the Shire River (Malawi) (Tweddle and Willoughby 1979).”

“In southern Africa, this species is native to the Cunene, Okavango, Zambezi systems including Lake Malawi [Tanzania, Malawi, Mozambique, Zambia, Zimbabwe, Botswana, Angola, Namibia], and east coastal rivers south to the Phongolo and coastal lakes to Lake Sibaya (Skelton 2001) [Mozambique, Eswatini, South Africa], as well as estuaries in Mozambique and KwaZulu- Natal [South Africa]. It is found in all Zimbabwean rivers although it was probably absent originally from the upper Save-Runde and scarce on the highveld (Junor 1969).”

“Native: Angola; Botswana; Burundi; Congo, The Democratic Republic of the; Eswatini; Malawi; Mozambique; Namibia (Caprivi Strip); South Africa (Gauteng, KwaZulu-Natal, Limpopo Province, Mpumalanga, North-West Province); Tanzania, United Republic of; Zambia; Zimbabwe”

Introduced According to Froese and Pauly (2019a), C. rendalli is established in Brazil, Chad, Colombia, Cuba, El Salvador, , Gabon, , , , , Peru, Rwanda, , , Turkey, and .

According to Froese and Pauly (2019a), C. rendalli is probably established in Antigua, Republic of the Congo, and Wallis and Futuna.

According to Froese and Pauly (2019a), C. rendalli is not established or probably not established in Cameroon, Central African Republic, Cote d’Ivoire, Dominican Republic, Panama, Sudan, and .

Capture of over 50 adult T. rendalli from a reservoir in northwest Cameroon, reported by Tombi et al. (2017), supports establishment in Cameroon.

From Konings et al. (2018):

“It has been introduced and translocated in many parts of Africa. In northern Africa, it is recorded from Mauritania (introduced). It has also been introduced in many tropical and subtropical areas around the world.”

“Introduced: Cameroon; Congo; Gabon; Kenya; Mauritania; Rwanda”

From Cassemiro et al. (2018):

“In Brazil, O[reochromis] niloticus and C. rendalli have been recognized as established in the reservoirs of Paraná, São Paulo, and Minas Gerais (Petesse et al., 2007; Novaes & Carvalho, 2012; Ortega et al., 2015; Daga et al., 2016) […]”

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“[…] this species has been recorded primarily in reservoirs of hydroelectric power plants along the Paraná-Paraguay River Basin in Brazil (Luiz et al., 2003; Baumgartner et al., 2006; Rocha et al., 2011; Ortega et al., 2015) and weirs in northeast Brazil (Silva & Araújo, 1996).”

Sherley (2000) reports C. rendalli (as T. rendalli) is established in the Pacific island nations of Wallis, Papua New Guinea, and Guam, and introduced but not established in and .

Pérez-Ponce de León et al. (2000) report C. rendalli (as T. rendalli) is established in Mexico.

Means of Introduction Outside the United States From Konings et al. (2018):

“It has been introduced (as Tilapia melanopleura) for aquacultural purposes, in 1949, from Yangambi, Democratic Republic of Congo, to Yaoundé, Cameroon. According to Thys van den Audenaerde (1966), who observed a few specimens of this species at the fisheries station of Melen near Yaoundé, the specimens from Yangambi originated from Katanga. Its use for aquaculture in Cameroon has been abandoned. It has also been introduced, for aquacultural purposes, in 1953, from the Democratic Republic of Congo to the fisheries station of Djoumouna, Republic of the Congo (Congo basin). According to Moreau et al. (1988) its use in aquaculture was abandoned, but this species probably established in the country [Republic of the Congo]. The introduction of C. rendalli into the Lower Guinean part of the Republic of the Congo is confirmed by a museum record originating from a pond at the station of Dimonika. Coptodon rendalli has also been introduced (as T. melanopleura) around 1950, from Katanga but originating from the fisheries station of Kinshasa (Leopoldville), Democratic Republic of Congo, to Libreville, Makokou, Lebamba, Franceville and maybe other places in Gabon. Its presence in Gabon is confirmed by museum records from the Ogowe River basin.”

“It has been introduced in and many dams and water systems all over the region [Kenya, Tanzania, Uganda], for example the Pangani drainage (including Lake Jipe), Lake Chala and Athi/Sabaki drainage. It has also been introduced in the Tana River system (Mann 1966, 1968, Seegers et al. 2003). According to Welcomme (1988) and Lever (1996) it was introduced from an unrecorded source into Kenya in 1955 for stocking (Seegers et al. 2003). It has also been introduced and is now well settled in the upper and middle Akagera system [Burundi, Rwanda].”

“Its distribution [in Zimbabwe] has been extended by translocations, especially into small farm dams, in an attempt to control plant growth.”

From Cassemiro et al. (2018):

“In parts of these regions [north Argentina to northeast Brazil], the introduction of C. rendalli occurred with the principal purpose of increasing the fisheries in dammed systems (Agostinho et al., 2007). Thus, this species has been recorded primarily in reservoirs of hydroelectric power plants […]”

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From Espinosa-Pérez and Ramírez (2015):

aureus (Steindachner, 1864), O. mossambicus (Peters, 1852), O. niloticus (Linnaeus, 1758), Tilapia rendalli (Boulenger, 1897), T. zillii (Gervais, 1848) and Hemichromis guttatus Günther, 1862 are African species introduced for aquaculture in Mexico. These species have been progressively introduced in many freshwater bodies of the country, both natural and artificial for aquaculture and commercial catch.”

Short Description From Froese and Pauly (2019a):

“Dorsal spines (total): 15 - 17; Dorsal soft rays (total): 10-13; Anal spines: 3; Anal soft rays: 9 - 10; Vertebrae: 29. Diagnosis: A large, deep-bodied species with a steep head profile, narrow head and small mouth; often appearing brownish with a white belly, some individuals have bright red bellies [Genner et al. 2018]. The sexes look very similar, although males are usually larger [Genner et al. 2018]. Very difficult to distinguish from Coptodon zillii, but C. rendalli usually have a steeper head profile and less prominent vertical bars; in East Africa, the tailfin of C. rendalli is often divided into a brownish upper part ad [sic] yellowish lower part, whereas that of C. zillii is uniform and spotted [Genner et al. 2018].”

Biology From Froese and Pauly (2019a):

“It prefers quiet, well-vegetated water along river littorals or backwaters, floodplains and swamps. […] Forms schools; is mainly diurnal. Juveniles feed on [Lamboj 2004]; adults feed on leaves and stems of underwater plants as well as algae, and vegetative detritus [Lamboj 2004], and . A substrate spawner; male and female form pairs to rear the young; eggs and larvae are usually guarded in a steep-side circular pit dug in the mud [Genner et al. 2018]. Occasionally it spawns in large cave-like structures [Lamboj 2004], e.g. in Lake Malawi they are reported to dig a network of tunnels at some sites [Genner et al. 2018].”

Human Uses From Froese and Pauly (2019a):

“Fisheries: commercial; aquaculture: commercial; gamefish: yes; aquarium: commercial”

From Konings et al. (2018):

“The species is not targeted by the ornamental fish trade, but is an important food fish.”

From Nico et al. (2019):

“Tilapia rendalli is a popular angling species in Africa, is important in aquaculture and fisheries, and also is used for weed control in reservoirs (Skelton 1993). This species is widely targeted by

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anglers in the Humacao Natural Reserve lagoon system in Puerto Rico (Ferrer Montaño and Dribble 2008).”

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

From Froese and Pauly (2019a):

“Anchor worm Disease, Parasitic infestations (protozoa, worms, etc.) Cichlidogyrus Disease, Parasitic infestations (protozoa, worms, etc.) Clinostomum Infestation (metacercaria), Parasitic infestations (protozoa, worms, etc.) Acanthogyrus Infestation, Parasitic infestations (protozoa, worms, etc.) Paradilepis Infestation, Parasitic infestations (protozoa, worms, etc.) […] Eye Infection (Diplostomum sp.), Parasitic infestations (protozoa, worms, etc.)”

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

“Potential pest”

3 Impacts of Introductions From Redding (1989):

“The main negative effects on the environment caused by T. rendalli has been due to its voracious feeding habits. In Mauritius (George 1976), Brazil (Nomura 1976, 1977), and Madagascar (Lamarque et al. 1975) T. rendalli is reported to have upset the ecology of the waters through either overpopulation, destruction of vegetation or through disturbing the ecology. It is reported that in Madagascar T. rendalli was accidentally introduced and proceeded to wipe out 3000 hectares of Ceratophyllum and Nymphaea in three years and, as a result, one valuable species of indigenous fish Paretropus petiti was almost wiped out (Lamarque et al. 1975).”

From Starling et al. (2002):

“We examined whether a large stock of tilapia (>750 kg ha−1, in littoral areas >1300 kg ha−1), mostly Oreochromis niloticus (L.) and Tilapia rendalli (Boulenger), could contribute to the eutrophication of a tropical reservoir (Lago Paranoá, Brasília, Brazil) by enhancing P[hosphorus]‐loading.”

“We took advantage of an extensive fish kill [mostly Oreochromis niloticus (L.) and Tilapia rendalli (Boulenger)] (>150 tons removed) during May–August 1997 in a hypereutrophic branch of the reservoir to compare water quality characteristics 1 year before and after this event by means of BACI statistics. We also measured P‐excretion rates in laboratory trials to assess the P‐ loading of the reservoir by the tilapia relative to tributary inputs and loading from a sewage treatment plant.”

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“Concentrations of chlorophyll a (decline from 84 to 56 μg L−1, P = 0.018) and total P (decline from 100 to 66 μg L−1, P < 0.001) decreased significantly in the branch of the reservoir affected by the fish kill, compared with a similar but unaffected branch that served as a control. Because P‐loading by both a sewage treatment plant and tributaries remained high after the incidence, the fish kill was likely to contribute to the observed water quality improvement.”

From Wager and Rowe-Rowe (1972):

“Wager (1968) lists many floating and submerged aquatic, shallow water and verge plants that are entirely consumed by T. rendalli. Junor (1969) gives a similar list, including many different species. T. rendalli avidly feeds on fry of other fish, notably of largemouth bass, Micropterus salmoides.”

“Most harm is done to a fresh water by T. rendalli.”

From Canonico et al. (2005):

“In Lake Alaotra, the progressive introductions of different species – carp first, followed by several species of in 1954 (T. rendalli), 1958 (O. macrochir), and 1961 (O. niloticus and O. mossambicus) – have also induced a drastic decline of native fish (Lévêque, 1997). Lévêque noted the quick proliferation of each of the tilapias since the first introduction in 1954, attributed to their high fecundity and ability to occupy empty niches.”

From Ramanantsalama et al. (2018):

dambabe is a local endemic fish species only found in Kinkony Lake (in north- western wetland and protected area) and two surrounding small lakes, Andranobe and La Digue [Madagascar]. […] Environmental variables and the presence of the Tilapiinae do not yet affect the presence of P. dambabe in the Kinkony Lake. But, the abundance of this endemic fish is negatively and significantly influenced by the abundance of the common carp Cyprinus carpio, Ambassis sp., sleepy goby biocelatus, G. giuris, longfin tilapia Oreochromis macrochir, tilapia O. niloticus, Mozambic tilapia O. mossambicus, redbreast tilapia Tilapia rendalli and redbelly tilapia T. zillii.”

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4 Global Distribution

Figure 1. Known global distribution of Coptodon rendalli, reported from Africa and the Americas. Map from GBIF Secretariat (2019). Occurrences in Alabama (United States), central Brazil, the Dominican Republic, and the Ivory Coast were excluded from the climate matching analysis because they do not represent established populations of C. rendalli.

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5 Distribution Within the United States

Figure 2. Known distribution of Coptodon rendalli in the United States. Map from Nico et al. (2019). Yellow diamonds represent established populations; the orange diamond represents a specimen with a status of “collected”.

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 0.101, indicating a medium overall climate match. A Climate 6 score between 0.005 and 0.103 indicates a medium climate match. The southern United States had the highest climate match: individual State Climate 6 scores were high in Alabama, Arizona, Florida, Georgia, Louisiana, Mississippi, North Carolina, New Mexico, and Texas; and medium in California and South Carolina. Locally, high matches occurred in peninsular Florida, along the Gulf Coast, in western Texas, and in southern New Mexico. The northern United States had a generally low climate match, except around Puget Sound, where the climate match was medium.

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Figure 3. RAMP (Sanders et al. 2018) source map showing weather stations selected as source locations (red; southern and central Africa [including Madagascar], Brazil, Colombia, the United States [Puerto Rico and Hawaii], and Mexico) and non-source locations (gray) for Coptodon rendalli climate matching. Source locations from GBIF Secretariat (2019).

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Figure 4. Map of RAMP (Sanders et al. 2018) climate matches for Coptodon rendalli in the contiguous United States based on source locations reported by GBIF Secretariat (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 Climate Match (Sum of Climate Scores 6-10) / (Sum of total Climate Scores) Category 0.000≤X≤0.005 Low 0.005

7 Certainty of Assessment Information is available on the biology, ecology, and distribution of Coptodon rendalli. Multiple introductions of this species outside of its native range have been documented, although identification to the species level can be difficult at times. Negative impacts of introduction of this species have been documented, but only one study attributed impacts solely to C. rendalli and not to a group of introduced species. Further information is needed to be confident in the assessed risk, so the certainty of this assessment is medium.

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8 Risk Assessment Summary of Risk to the Contiguous United States Coptodon rendalli, the Redbreast Tilapia, is a fish species native to a wide range of fresh and brackish waters in tropical and subtropical Africa. It has been introduced widely outside its range in Africa for aquaculture and weed control, and it is also a commercial and recreational fishing target. C. rendalli has been introduced to parts of Africa outside the native range, Central and South America, Thailand, Turkey, and some Pacific islands. In the United States, C. rendalli is established in Hawaii and Puerto Rico. In Madagascar, extreme population decline of an endemic fish is attributed to the impacts of C. rendalli herbivory on its habitat. Because most introductions of C. rendalli have occurred in locations where other nonnative species introductions have also occurred, the impacts of C. rendalli often cannot be separated from impacts of other nonnative species. For these reasons, the history of invasiveness is high for C. rendalli, but the certainty of the assessment is medium. C. rendalli has a medium climate match with the contiguous United States, with the highest match occurring in the southern United States. The overall risk assessment category is high.

Assessment Elements  History of Invasiveness (Sec. 3): High  Climate Match (Sec. 6): Medium  Certainty of Assessment (Sec. 7): Medium  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.

Canonico, G. C., A. Arthington, J. K. McCrary, and M. L. Thieme. 2005. The effects of introduced tilapias on native biodiversity. Aquatic Conservation: Marine and Freshwater Ecosystems 15:463-483.

Cassemiro, F. A. S., D. Bailly, W. J. da Graça, and A. A. Agostinho. 2018. The invasive potential of tilapias (Osteichthyes, Cichlidae) in the Americas. Hydrobiologia 817(1): 133-154.

Espinosa-Pérez, H., and M. Ramírez. 2015. Exotic and invasive in Mexico. Check List 11(3):1627.

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).

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Froese, R., and D. Pauly, editors. 2019a. Coptodon rendalli Boulenger, 1897. FishBase. Available: https://www.fishbase.de/summary/Coptodon-rendalli.html. (May 2019).

Froese, R., and D. Pauly, editors. 2019b. Coptodon rendalli (Boulenger, 1897). In World Register of Marine Species. Available: http://www.marinespecies.org/aphia.php?p=taxdetails&id=1013588. (July 2019).

GBIF Secretariat. 2019. GBIF backbone : Coptodon rendalli (Boulenger, 1897). Global Biodiversity Information Facility, Copenhagen. Available: https://www.gbif.org/species/9312593. (July 2019).

Konings, A., A. Awaïss, A. Azeroual, A. Getahun, M. Hanssens, P. Lalèyè, B. Marshall, T. Moelants, G. Natakimazi, and D. Tweddle. 2018. Coptodon rendalli. The IUCN Red List of Threatened Species 2018: e.T60690A47209450. Available: https://www.iucnredlist.org/species/60690/47209450. (May 2019).

Nico, L., M. Neilson, and B. Loftus. 2019. Tilapia rendalli (Boulenger, 1897). U.S. Geological Survey, Nonindigenous Aquatic Species Database, Gainesville, Florida. Available: https://nas.er.usgs.gov/queries/FactSheet.aspx?SpeciesID=48. (May 2019).

OIE (World Organisation for Health). 2019. OIE-listed diseases, infections and infestations in force in 2019. World Organisation for Animal Health, Paris. Available: http://www.oie.int/animal-health-in-the-world/oie-listed-diseases-2019/. (July 2019).

Pérez-Ponce de León, G., L. García-Prieto, V. León-Règagnon, and A. Choudhury. 2000. Helminth communities of native and introduced fishes in Lake Pátzcuaro, Michoacán, México. Journal of Fish Biology 57:303-325.

Ramanantsalama, R. V., R. H. Andriamasimanana, A. Andrianarimisa, and R. Oliarinony. 2018. Relationship between the abundance and distribution of the endemic fish Paretroplus dambabe in the north-western wetland of Madagascar and the effect of environmental parameters and the presence of other fish species. Biodiversity International Journal 2(2):165-171.

Redding, T. A. 1989. Report on the biology and ecology of the introduced tilapia Oreochromis mossambicus (Peters) (Pisces: Cichlidae) in the Sepik River, Papua New Guinea, and the social and economic impact of its introduction. PNG/85/001 Field Document no. 10. Food and Agriculture Organisation of the United Nations, Rome.

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

Sherley, G., editor. 2000. in the Pacific: a technical review and draft regional strategy. South Pacific Regional Environment Programme, Apia, Samoa.

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Starling, F., X. Lazzaro, C. Cavalcanti, and R. Moreira. 2002. Contribution of omnivorous tilapia to eutrophication of a shallow tropical reservoir: evidence from a fish kill. Freshwater Biology 47:2443-2452.

Tombi, J., J. M. Tchiegno, and J. F. Akoumba. 2017. Microecology of monogenean gill parasites of Tilapia rendalli Boulenger, 1897 from Bamendjing Lake, Cameroon. Journal of Environment and Ecology 8(1):43-54.

Wager, V. A., and D. T. Rowe-Rowe. 1972. The effects of Tilapia rendalli and T. mossambica on aquatic macrophytes and fauna in five ponds. South African Journal of Science 68(10):257-260.

Wu, L., and J. Yang. 2012. Identifications of captive and wild tilapia species existing in Hawaii by mitochondrial DNA control region sequence. PLoS ONE 7(12):e51731.

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.

Agostinho, A. A., L. C. Gomes, and F. M. Pelicice. 2007. Ecologia e manejo de recursos pesqueiros em reservatórios do Brasil. EDUEM, Maringá, Brazil.

Anonymous. 1994. Atlas pesquero de México. Instituto Nacional de la Pesca, Mexico City, Mexico.

Baensch, H. A., and R. Riehl. 1991. Aquarien Atlas, volume 3. Mergus, Verlag für Natur-und Heimtierkunde, Melle, Germany.

Barrett, P. J. 1983. Systematics of fishes of the genus Tilapia (Perciformes: Cichlidae) in the lower Colorado River basin. Master’s thesis, Arizona State University, Tempe, Arizona.

Baumgartner, D., G. Baumgartner, C. S. Pavanelli, P. R. L. Silva, V. A. Frana, L. C. Oliveira, and M. R. Michelon. 2006. Fish, Salto Osório Reservoir, Iguaçu River basin, Paraná State, Brazil. Check List 2:1-4.

Courtenay et al. 1984 [Source material did not provide full citation for this reference.]

Courtenay, W. R., Jr., D. P. Jennings, and J. D. Williams. 1991. Appendix 2: exotic fishes. Pages 97-107 in C. R. Robins, R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. B. Scott, editors. Common and scientific names of fishes from the United States and Canada, 5th edition. American Fisheries Society Special Publication 20. American Fisheries Society, Bethesda, Maryland.

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Daga, V. S., T. Debona, V. Abilhoa, É. A. Gubiani, and J. R. S. Vitule. 2016. Non-native fish invasions of a Neotropical ecoregion with high endemism: a review of the Iguaçu River. Aquatic Invasions 11:209-223. de Moor, I. J., and M. N. Bruton. 1988. Atlas of alien and translocated indigenous aquatic in southern Africa. South African National Scientific Programmes Report 144. Foundation for Research Development and Council for Scientific and Industrial Research, Pretoria, South Africa.

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