Cichla ocellaris (peacock cichlid)

Pictures

Picture	Title	Caption	Copyright
Title Adult Caption Cichla ocellaris (butterfly peacock bass); adult. Captive specimen. ZOO Dvůr Králové, Czech Republic. April, 2010. Copyright Public Domain/via wikipedia/released by Karel Jakubec, Prague, Czech Republic	Adult	Cichla ocellaris (butterfly peacock bass); adult. Captive specimen. ZOO Dvůr Králové, Czech Republic. April, 2010.	Public Domain/via wikipedia/released by Karel Jakubec, Prague, Czech Republic

Identity

Preferred Scientific Name

• Cichla ocellaris Bloch & Schneider, 1801

Preferred Common Name

• peacock cichlid

Other Scientific Names

• Acharnes speciosus Müller & Troschel, 1849

International Common Names

• English: butterfly peacock bass
• Spanish: pavón; sargento; tucunaré

Local Common Names

• Brazil: furiba; peixe moeda; peixe zebra; pitanga; tucunaré comum
• Denmark: aborrecichlide
• Finland: isokikla; toukounaré
• French Guiana: aboné; kunan; malisamba; toekoenari; toukounalé; tukunali
• Germany: Grüner Augenfleck-Kammbarsch
• Hawaii: tucunare
• Puerto Rico: mamito
• Sweden: lukanani

Summary of Invasiveness

C. ocellaris is a large, piscivorous cichlid that is native to large areas of tropical South America. Due to its desirability as a sport-fish, C. ocellaris has been translocated and introduced into aquatic environments of at least 10 countries. This species inhabits rivers, lakes and anthropogenically modified environments including drainage canals and reservoirs. C. ocellaris has also been introduced worldwide as an ornamental aquarium species. C. ocellaris is a highly efficient piscivore that may impact upon ichthyofaunas and aquatic environments through the mechanisms of predation, competition and alteration of food webs. In Lake Gatun, Panama seven of eight native fish species declined by 50–100% and the elimination of small invertebrate-feeding fishes resulted in an increase in mosquito larvae and a higher incidence of malaria around the lake. In contrast to this, in Florida, C. ocellaris has a positive impact where it has been released as a biocontrol agent for invasive species cichlids such as Tilapia mariae.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Chordata
•             Subphylum: Vertebrata
•                 Class: Actinopterygii
•                     Order: Perciformes
•                         Family: Cichlidae
•                             Genus: Cichla
•                                 Species: Cichla ocellaris

Notes on Taxonomy and Nomenclature

Cichla ocellaris Bloch & Schneider 1801, is a piscivorous fish native to large areas of tropical South America (Espínola et al., 2015). Species of Cichla are widely distributed in the Amazon, Tocantins and Orinoco river basins and in the smaller rivers draining the Guianas to the Atlantic Ocean.

Kullander and Ferreira (2006) reviewed the genus and recognised 15 species, including nine new species, by morphology and meristics (principally colour pattern as the most important determinant). These authors split C. ocellaris into several species, including C. kelberi, a species they recognised within the Tocantins–Araguaia Basin.

Willis et al. (2012) conducted genetic analysis of the Cichla genus based on multiple separate sources of molecular data, mtDNA, nuclear sequences and microsatellites and concluded that several of the species described by Kullander and Ferreira (2006) are actually C. ocellaris, a single species with extensive genetic introgression among geographic variants showing varying degrees of morphological differentiation. Willis et al. (2012) recommended that C. ocellaris remain a valid species and C. monoculus, C. nigromaculata, C. kelberi and C. pleiozona are all synonymised with C. ocellaris. Despite this, it has not yet been widely acknowledged and some sources still recognise the distinct species, e.g., Froese and Pauly, (2015).

Due to the taxonomic uncertainty regarding the genus Cichla, USGS NAS (2015) consider that many historical references to C. ocellaris may indeed refer to other species of Cichla, or possibly even hybrids between species. Therefore, in this datasheet C. ocellaris will include the synonymisation of the four other Cichla species listed above and historical references to C. ocellaris will be considered to be based on accurate species identification.

Description

The following description has been compiled using information from Page and Burr (1991), Shafland (1995), Winemiller (2001), Willis et al. (2012), Butler (2015), Froese and Pauly (2015) and ISSG (2015).

C. ocellaris is a large cichlid that may reach 91 cm, though 50-60 cm is the typical adult size. The largest specimen from introduced populations in the USA was 74 cm in length and weighed and 6.8 kg. Males are larger than females. The species has a sloping forehead and an elongate body with a deeply notched dorsal fin. The mouth is large and the lower jaw projects beyond the upper jaw. Many adult fish (primarily males, but also some females) develop a pronounced nuchal hump on their foreheads. Physical traits and colouration vary greatly depending on source location, individual and stage of development. Colouration is olive-green dorsally fading to yellow-white ventrally. An orange-red region may also be present on the ventral surface that includes the anal, pelvic and lower cauda fins. Three broad transverse stripes are usually present on the flanks and between these stripes are a series of irregular dark spots. The first dorsal, upper caudal and pectoral fins are grey or black. White spots are present on the second dorsal and the upper lobe of the caudal fin. There is a diagnostic large black spot encircled by a silver-coloured halo on the caudal fin. Large adults may have a yellow-orange stripe from mouth to caudal fin base. The iris is red. Both sexes have more intense coloration during the reproductive period.

Distribution

C. ocellaris is native to large areas of South America including French Guiana, Guyana, Suriname, Venezuela, Colombia, Peru, Bolivia and Brazil (Winemiller, 2001; Kullander and Ferreira, 2006; Willis et al., 2012; Espínola et al., 2015).

The desirability of C. ocellaris as a sport-fish is the predominant reason for the majority, if not all, introductions of the species around the world; e.g. Panama (Zaret and Paine, 1973), southern Florida (Shafland and Stanford, 1999), Puerto Rico (Erdsman, 1984), Guam (Welcomme, 1988) and Hawaii (Maciolek, 1984). This species is also present in Arizona, USA, the Dominican Republic, Guam, Kenya, Malaysia, Singapore and US Virgin Islands (Welcomme, 1988; Shafland, 1995; Tan and Tan, 2003; Chong et al., 2010; USGS NAS, 2015).

Distribution Table

The 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.

History of Introduction and Spread

USGS NAS (2015) provide a detailed summary of introductions of C. ocellaris to the USA and US territories.

In Florida, there is a long history of stocking C. ocellaris in southern areas of the state. C. ocellaris was introduced into south-eastern Florida in 1964 (Courtenay et al., 1974; Courtenay and Robins, 1989). The Florida Game and Fresh Water Fish Commission obtained breeding stock from several regions of South America and the progeny were released into open waters primarily as a sport-fish, but also as a biological control agent of other introduced cichlids (Courtenay and Robins, 1989; Shafland, 1995). Approximately 10,000 juveniles were released in the Fort Lauderdale area of Dade County in 1964. However, these fish did not survive the cold winter of 1964-1965 (Courtenay et al., 1974; Courtenay and Robins, 1989). From 1984, additional stockings were carried out in canals in Broward and Miami-Dade counties, Florida and have resulted in established populations (Shafland, 1995). Today, C. ocellaris is currently recorded in at least 10 Florida canal systems and from lakes near Miami International Airport and in Miami-Dade County (USGS NAS, 2015). Shafland (1996) reported that fishable populations of C. ocellaris exist in more than 500 km of canals and also numerous urban lakes in the metropolitan Miami-Ft. Lauderdale area.

In Hawaii, C. ocellaris was released into reservoirs on Kauai and Oahu, from 1961, by the Hawaiian Division of Fish and Game (Maciolek, 1984). The broodstock came from Guyana (USGS NAS, 2015). This species has also established in Wahiawa Reservoir, a 350-acre, privately owned irrigation reservoir on Oahu (USGS NAS, 2015).

In Texas, C. ocellaris was first released by the Texas Parks and Wildlife Department in 1978 with stock from Colombia, possibly Brazil and from the Florida Game and Fresh Water Fish Commission (USGS NAS, 2015). The species was introduced into five electric power plant reservoirs in Texas between 1978 and 1984; Alcoa Reservoir (Milam County), Lake Bastrop (Bastrop County), Coleto Creek Reservoir (Goliad County), Wilkes Reservoir (Upshur County) and Tradinghouse Creek Reservoir (McLennan County). Some populations persisted briefly though all disappeared by 1992 due to temperature extremes (Howells and Garrett, 1992).

In Arizona, a single C. ocellaris was caught by an angler in the Wellton-Mohawk Canal near Yuma in September 2010. This was considered to be a likely aquarium release or an illegal stocking as no authorized stocking has occurred in Arizona (USGS NAS, 2015).

C. ocellaris was introduced to the Rio Gatuncillo, a small tributary of the Chagres River, Panama, in approximately 1966 (Zaret and Paine, 1973). By 1967, the species had naturally dispersed approximately 8 km of this small tributary to inhabit the Chagres River. By 1970, C. ocellaris was collected approximately 15 km downstream in Lake Gatun (Zaret and Paine, 1973). In 1972, the species was observed on the opposing eastern shore of Lake Gatun, approximately 30 km away. Thus, under favourable conditions, C. ocellaris my naturally disperse very rapidly after introduction. 

Introductions

Risk of Introduction

C. ocellaris is not a popular ornamental species because it grows to a large size and is piscivorous and is therefore definitely not suitable for the standard “community” aquarium. However, C. ocellaris has been widely dispersed, for example, in the USA, as a sport-fish and there is the potential for this to continue (USGS NAS, 2015). Once introduced into an area natural dispersal of this species is likely and can occur very rapidly. However, natural dispersal of C. ocellaris may be constrained by salinity and temperature tolerances.

Habitat

C. ocellaris inhabits rivers and lakes and anthropogenically modified environments including drainage canals (Florida) and reservoirs/lakes (USA and South America) (USGS NAS, 2015). Favoured aquatic habitats of C. ocellaris range from rapids to quiet waters with medium depth (approximately 5 m) and rocky substrates (Froese and Pauly, 2015; ISSG, 2015). As C. ocellaris is a diurnal, visual predator, the species prefers environments with higher water clarity (Winemiller, 2001). The species occurs primarily in freshwater but can tolerate moderate salinities and brackish water (ISSG, 2015).

Habitat List

Biology and Ecology

Genetics

The diploid chromosome number of C. ocellaris is 48 (Brinn et al., 2004; Willis et al., 2012).

Reproductive Biology

C. ocellaris is a biparental substrate spawner. Spawning usually takes place on a flat, horizontal surface which is cleaned of algae or vegetation by the parents during spawning activities. The female lays a single row of eggs and the male follows exuding sperm over each row. Site selection is based on ease of predator deterrence and food supply for the young (Gomiero and Braga, 2004a). Once the eggs have hatched, the parents transport the larvae in their mouths to a shallow depression in the sediment, or “nest” (Zaret, 1980). Although only one nest is used to deposit the larvae, large numbers of nests are built to confuse predators (Braga, 1952). Breeding pairs guard their clutch for up to 10 weeks at which time the juveniles are 60-70 mm (Zaret 1980). Fry then move from open waters to areas of dense vegetation that provide protection from predators and food resources (Schroeder and Zaret, 1977). Breeding C. ocellaris are territorial, aggressive and will be hostile to con and hetero-specifics (Zaret, 1980). Parental care of offspring exhibited by cichlid fishes greatly increases offspring survival rates. In fluvial systems C. ocellaris reproduction occurs during low-water conditions and continues into the flood period (Winemiller, 2001).

In canals in south-east Florida, USA, C. ocellaris spawns from May-June and fecundity estimates range from 4287-9840, with an average of 6560 eggs. Eggs were observed to be elongated, light yellow to yellow-brown and averaged 2.27 mm long and 1.27 mm wide. The smallest mature female and male fish were found to be 256 mm Total Length (TL) and 242 mm TL, respectively. It was found that the percentage of female fish increased with increasing fish size, up to a maximum of 400 mm (no fish larger than 400 mm were sampled for this research) (Shafland, 1999b).

In the Santa Cruz Reservoir, north-eastern Brazil, the sex ratio of the introduced population of C. ocellaris (referenced as C. monoculus) differed from 1:1 with a predominance of males. The size at reproductive maturity was estimated to be 15.65 cm and reproduction occurred throughout the year (de Oliveira Sousa et al., 2015). However, in the Volta Grande Reservoir, south-eastern Brazil, Gomiero and Braga (2004a) investigated the reproduction of introduced populations of C. ocellaris (referenced as Cichla cf. ocellaris and C. monoculus) and found that spawning began at the end of the dry season (September) and extended until the end of the rainy season (January).

Physiology and Phenology

C. ocellaris is diurnal and non-migratory. Shafland (Shafland 1996; Shafland 1999a; Shafland, 1999b; Shafland, 1999c) conducted extensive research on the biology and ecology of C. ocellaris in six canal systems of south-eastern Florida in 1995. The mean TL of the species was 349 mm. Under favourable conditions C. ocellaris can grow rapidly and become sexually mature and attain 250-300 mm in under 12 months (Shafland, 1996). Annual growth estimates for the species in the Tamiami Canal over the years 1-6 were calculated as 204 mm, 327 mm, 418 mm, 480 mm, 531 mm and 592 mm (Shafland, 1999c).

In the Santa Cruz Reservoir, north-eastern Brazil, the standard length (SL) of C. ocellaris (referenced as C. monoculus) ranged between 70- 3877 mm with the most frequent size class being 100-150 mm (de Oliveira Sousa et al., 2015).

Nutrition

C. ocellaris is a diurnal predator and is principally piscivorous (Winemiller, 2001). Prey is typically caught through high-speed pursuit rather than ambush (Shafland, 1995). In Barro Colorado Island, Panama, introduced populations of C. ocellaris were found to consume atherinids, poeciliids, characids, eleotrids and other cichlids (Zaret and Paine, 1973). C. ocellaris was introduced to the canals of south-eastern Florida in an attempt to control small introduced fishes in these habitats (Shafland, 1999a). Here, they fed almost exclusively on fish such as Tilapia mariae (Shafland, 1999b).

Novaes et al. (2004) examined the diet of C. ocellaris (referenced as C. monoculus) from the upper Tocantins River during construction of the Serra da Mesa power plant, in Brazil. Analysis of the stomach contents revealed a piscivorous diet with Tetragonopterin fishes dominating the diet during the filling phase and Cichlidae the dominant prey taxa during the operation phase. Cannibalism was recorded in C. ocellaris in the both the filling operational phases (Novaes et al., 2004).

Environmental Requirements

As C. ocellaris is a diurnal visual predator, it exhibits a strong preference for aquatic environments with high water clarity (Winemiller, 2001). It is euryhaline though occurs primarily in freshwater (ISSG, 2015). An upper salinity tolerance of 18 ppt has been reported (Shafland, 1995). C. ocellaris is a tropical species and is less cold tolerant than other introduced cichlids in the USA. The range of C. ocellaris in Florida is limited by cold winters that restricts it to the southernmost counties and exclude it from much of the Everglades (USGS NAS, 2015). Swingle (1966) reported a lower lethal temperature of 16°C for 80 mm fingerlings and Guest et al. (1979) reported a similar lower lethal temperature of 15.6°C and a higher lethal temperature of 37.9°C for fingerlings between 85-140 mm TL. When salinity was raised to 10 ppt some fish exhibited a lower lethal temperature of 13.5 °C (Guest et al., 1979).

Climate

Latitude/Altitude

Water Tolerances

Natural Enemies

Notes on Natural Enemies

Little data are available on the natural predators of C. ocellaris, however, fry and juvenile fishes would likely be preyed upon by larger piscivorous fish and animals. A study in Florida found C. ocellaris to be present in the stomachs of Micropterus salmoides (Shafland, 1999b). In Brazil, juvenile C. ocellaris were the major dietary taxa of larger fish (Gomiero and Braga, 2004b). C. ocellaris is also known to be cannibalistic and will consume small fish of the same species (Novaes et al., 2004).

Means of Movement and Dispersal

Natural Dispersal

Upon introduction to favourable environments, populations of C. ocellaris may disperse rapidly through interconnected river systems and aquatic habitats. Three examples include the introduction of the species to Lake Gatun in Panama (Zaret and Paine, 1973), the drainage canals of southern Florida (Shafland, 1995; Shafland, 1996) and tributaries of the Parana River in south-eastern Brazil (Espínola et al., 2015).

Accidental Introduction

Although a number of individuals have apparently claimed responsibility for the introduction of C. ocellaris into Lake Gatun in Panama, the most likely initial introduction of the species was accidental (Zaret and Paine, 1973). Fingerlings were imported from Columbia and placed in an impoundment that was formed by damming a small tributary of the Rio Gatuncillo. It is likely that the impoundment overflowed during the rainy season releasing fish into the tributary and ultimately Lake Gatun (Zaret and Paine, 1973).

Intentional Introduction

C. ocellaris is a desirable sport and table fish and it is likely that the species is regularly released into new waterways. After the introduction and proliferation of C. ocellaris in Panama it has been “casually introduced” throughout the region with no controls or regulation (Zaret and Paine, 1973).

Pathway Causes

Pathway Vectors

Impact Summary

Environmental Impact

C. ocellaris is a highly efficient piscivore that may impact upon ichthyofaunas and aquatic environments through the mechanisms of predation, competition and alteration of food webs. While introductions in Panama (Zaret and Paine, 1973) and Brazil (Gomiero and Braga, 2004c) have resulted in environmental and social impacts, the introduction of C. ocellaris to southern Florida has resulted in greater environmental and social benefits (Shafland, 1999a; Shafland, 1999b).

After the introduction of C. ocellaris into Lake Gatun, Panama, largescale changes in food-web structure and aquatic community composition were recorded (Zaret and Paine, 1973). For example there was a reduction in almost all secondary consumers. The reduction in the planktivore Melaniris chagresi (Atherinidae) resulted in a reduction in tertiary-consumer populations including tarpon, black terns, kingfishers and herons in addition to changes within the zooplankton community. Near Barro Colorado Island in Lake Gatun, seven of eight native fish species declined by 50–100% and sites occupied by C. ocellaris contained seven native fish species in surveys compared with 13 fish species at sites where C. ocellaris was not present (Zaret and Paine, 1973).

In the Parana River in south-eastern Brazil, C. ocellaris, are well dispersed throughout the region and are considered the greatest threat to the native fish diversity (Agostinho et al., 2008; Pelicice and Agostinho, 2009). Kovalenko et al. (2010) conducted mesocosm experiments in sections of the Parana River to determine if the direct and indirect effects of introduced C. ocellaris on native prey were mitigated by the presence of aquatic vegetation. It was concluded that aquatic plants provided very limited protection to native prey and are therefore unlikely to slow down the decline in biodiversity resulting from the introduction and spread of species of Cichla including C. ocellaris (Kovalenko et al., 2010). A study by Pelicice and Agostinho (2009) in the Rosana Reservoir, Brazil, found that due to the introduction of C. ocellaris, the diversity of the reservoir changed dramatically with mean fish density decreasing by 95% and richness by 80%.

The introduction of C. ocellaris to southern Florida is the only documented example of positive environmental impacts following introduction of the species. It is important to note that the success of the introduction and the limitation of deleterious environmental impacts are largely due to the confinement of C. ocellaris to artificial drainage canals and lakes in the southern extremity of Florida where temperatures restrict the spread of this species. Shafland (1999b) determined that C. ocellaris primarily consumed the introduced cichlid Tilapia mariae and there was little dietary overlap between C. ocellaris and the native predator Micropterus salmoides. C. ocellaris also feeds on other non-native species present in the waters and the impact on native species is limited.

Social Impact

Following the introduction of C. ocellaris into Lake Gatun, Panama, radical changes in community composition and the structure of the aquatic food web occurred (Zaret and Paine, 1973). As such, the elimination of small invertebrate-feeding fishes as a consequence of C. ocellaris introduction resulted in more mosquito larvae and a higher incidence of malaria around the lake (Zaret and Paine, 1973).

Risk and Impact Factors

Impact mechanisms

• Competition - monopolizing resources
• Filtration
• Predation
• Rapid growth

Impact outcomes

• Altered trophic level
• Damaged ecosystem services
• Ecosystem change/ habitat alteration
• Negatively impacts cultural/traditional practices
• Negatively impacts human health
• Reduced native biodiversity
• Threat to/ loss of native species

Invasiveness

• Abundant in its native range
• Benefits from human association (i.e. it is a human commensal)
• Fast growing
• Gregarious
• Has a broad native range
• Has high reproductive potential
• Highly adaptable to different environments
• Highly mobile locally
• Long lived
• Proved invasive outside its native range

Likelihood of entry/control

• Difficult to identify/detect in the field
• Highly likely to be transported internationally deliberately
• Highly likely to be transported internationally illegally

Uses

Economic Value

The desirability of C. ocellaris as a sport-fish is the predominant reason for the majority, if not all, introductions of the species around the world; e.g. Panama (Zaret and Paine, 1973), southern Florida (Shafland and Stanford, 1999), Puerto Rico (Erdsman, 1984), Guam (Welcomme, 1988) and Hawaii (Maciolek, 1984). In Florida, a study ascertained that the total value of the canal sport-fisheries were US $15.5 million annually and estimated that the annual asset value for C. ocellaris was $6.6 million (Shafland and Stanford, 1999).

Environmental Services

C. ocellaris was extensively released not only to create a sport-fishery, but also to control introduced fishes such as Tilapia mariae, the spotted Tilapia (Shafland, 1999b; Robins, 2015). C. ocellaris may also be caught for their meat.

Uses List

Environmental

• Biological control

General

• Pet/aquarium trade
• Research model
• Sport (hunting, shooting, fishing, racing)

Human food and beverage

• Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)

Similarities to Other Species/Conditions

In the USA, species of Cichla are unlikely to be misidentified with native fishes due to their distinctive morphology and colouration. However, the accurate identification of individual Cichla species may be problematic because most members of the genus exhibit somewhat similar morphology and colouration.

Identification can be more difficult because of variations in morphology and colouration within species and potential hybridisation between species (USGS NAS, 2015). Thus, as outlined above, the systematics of the genus are contested.

Prevention and Control

Prevention

SPS Measures

In Brazil, the translocation and release of species of Cichla is illegal. However, due to a lack of knowledge of potential ecological impacts and their appeal as a sport-fish, introductions regularly occur all over the country (Pelicice and Agostinho, 2009).

Gaps in Knowledge/Research Needs

In order to fully understand the impacts of species of Cichla, there is a need for greater documentation of the invasion processes and the consequences for native communities and ecosystems (Espínola et al., 2015).

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Principal Source

Draft datasheet under review

Contributors

13/03/2016 Original text by:

Mark Maddern, University of Western Australia, Australia

Distribution Maps

• = Present, no further details
• = Evidence of pathogen
• = Widespread
• = Last reported
• = Localised
• = Presence unconfirmed
• = Confined and subject to quarantine
• = See regional map for distribution within the country
• = Occasional or few reports

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