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Feeding Morphology of the Neotropical Piscivorous Fish Cichla Kelberi (Perciformes: Cichlidae) Introduced Into an Oligotrophic Brazilian Reservoir

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Feeding Morphology of the Neotropical Piscivorous Fish Cichla Kelberi (Perciformes: Cichlidae) Introduced Into an Oligotrophic Brazilian Reservoir Feeding morphology of the Neotropical piscivorous fish Cichla kelberi (Perciformes: Cichlidae) introduced into an oligotrophic Brazilian reservoir Alejandra Filippo Gonzalez Neves dos Santos1, Luciano Neves dos Santos2 & Francisco Gerson Araújo3 1. Departamento de Zootecnia e Desenvolvimento Agrossocioambiental Sustentável, Universidade Federal Fluminense, Rua Vital Brasil, 64, Faculdade de Veterinária, Niterói, Brasil; [email protected] 2. Departamento de Ecologia e Recursos Marinhos, Universidade Federal do Estado do Rio de Janeiro Fluminense, Avenida Pasteur, 458, Rio de Janeiro, Brasil; [email protected] 3. Departamento de Biologia, Universidade Federal Rural do Rio de Janeiro, Antiga BR 465, Km 47, Seropédica, Brasil; [email protected] Received 13-IX-2010. Corrected 06-I-2011. Accepted 03-II-2011. Abstract: Despite being one of the most well-known cichlid fish of importance to artisanal and sport fishing, and among the largest fishes in the Neotropics, data on digestive tract anatomy of peacock basses (Cichla spp.) are largely lacking, especially for non-native populations. In this paper, we describe for the first time the digestive tract morphology of Cichla kelberi, a voracious piscivore that was introduced in the 1950s into an oligotrophic and physically low-complex impoundment in Brazil. Peacock basses were collected between 1994 and 2002 in Lajes Reservoir, through gillnets (25 to 55mm mesh; 20-50x2m), seines (10x2.5m; 8.0mm mesh), cast nets and angling. All the fishes were kept on ice in the field and then transferred to the laboratory, where they were identified, measured, weighed and dissected for digestive tract analyses. The Index of Relative Importance-IRI was calculated for diet characterization while linear and non-linear regressions were performed to assess growth patterns of four morphological characters related to feeding (e.g. mouth width, mouth height, stomach length and intestine length) and the number of gill rakers during the C. kelberi ontogeny. Most digestive tract structures were directly related to the piscivorous diet of C. kelberi, indicating that peacock bass is a diurnal, bathypelagic and gape-size limited predator that feeds largely on shallow-water prey species within the littoral zone. Mouth width and height grew allometrically (b>1) with the size of peacock bass, broadening the size range in which prey can be eaten, but especially for predators smaller than ~400mm of total length. Differently, stomach and intestine lengths increased isometrically (b=1), which could constrain prey consumption for adult C. kelberi, especially those at advanced stages of gonadal maturation. The presence of longer-drawn, sharp and furcated gill rakers in C. kelberi may be related to increased prey retention in the resource-limited Lajes Reservoir, but fur- ther studies are necessary whether such features are randomly triggered by genetic or phenotypic anomalies, or effectively bring ecological advantages to the predator. In addition to contribute to improve the current biological knowledge on peacock basses, our results can be also useful to further comparisons on whether those morpho- logical features related to feeding will change with transitions on prey consumption by C. kelberi and/or with the particular conditions of the invaded ecosystem. Rev. Biol. Trop. 59 (3): 000-000. Epub 2011 September 01. Key words: Cichlid, peacock bass, ecomorphology, diet, invasive species, reservoir. Peacock basses (Cichla spp.) are the major species are large (>500mm of total lenght), vis- group of piscivorous cichlids in South Ameri- ual-diurnal, and voracious predators, preying ca, which are native of the Amazon, Tocantins largely on small fishes, but occasionally con- and Orinoco river basins, and of the smaller suming shrimp and other aquatic invertebrates rivers draining the Guiana to the Atlantic when adults (Winemiller et al. 2001, Kullander Ocean (Kullander & Ferreira 2006). All Cichla & Ferreira 2006). Although peacock basses Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 59 (3): 000-000, September 2011 1 collectively inhabit a diversity of habitats rang- thresholds and/or transitions (Kováč et al. 1999, ing from streams to large reservoirs, the univer- Hellig et al. 2010). Recent studies have inves- sal requirement of all Cichla species appears tigated the feeding biology of cichlids, par- to be high water transparency, warm tempera- ticularly focusing on the trophic morphology tures, and access to lentic habitats for feeding of little-known species (Gordon & Bills 1999, and/or reproduction (Winemiller et al. 2001). Cochran-Biederman & Winemiller 2010, Hellig Because of their great popularity as sport fishes et al. 2010) or the predation-related impacts or to control undesirable forage fishes, pea- of non-indigenous populations (Bergmann & cock basses are also intentionally introduced Motta 2005, Fugi et al. 2008). Surprisingly, into many tropical and subtropical freshwater studies on the digestive tract morphology of systems around the world (e.g. Brazil, Hawaii, Cichla species are still lacking, despite their Panama, Puerto Rico and Florida), sometimes recognizable value as game fishes and their to the detriment of native fish populations increasing importance as invaders in the trop- (Zaret & Paine 1973, Santos et al. 2001, Latini ics (Espínola et al. 2009, Santos et al. 2009). & Petrere Jr 2004). In this paper, the digestive tract morphology Within the 15 peacock bass species cur- of C. kelberi, a voracious piscivore that was rently recognized by systematists, Cichla kel- introduced in the 1950s into Lajes Reservoir, beri (Kullander & Ferreira 2006) deserves an oligotrophic and physically low-complex special attention since it was introduced into impoundment in Brazil, is first described. In most Brazilian watersheds to enhance game addition to the general hypothesis that mor- and commercial fisheries or to control unde- phological features are close related to diet and sirable fishes (Santos et al. 2008, Espínola et feeding habit of C. kelberi, we expected that al. 2009). Until this recent revision, C. kelberi our study could contribute to improve the cur- was often misidentified as Cichla ocellaris or rent biological knowledge on peacock basses Cichla monoculus, two close congeners that and also to improve the biological knowledge are very similar in body shape and coloura- on non-native C. kelberi populations. tion. After that revision, most peacock bass introductions succeeded in Brazilian reservoirs MATERIALS AND METHODS appear to be of C. kelberi. However, since this species shares many morphometric and Study area: Lajes Reservoir (22o42’ N meristic features with C. monoculus, Cichla - 43o53’ W; 22o50’ N - 44o05’ W) is a 30km2 pleiozona and even C. ocellaris, taxonomic impoundment, located 415m above mean sea confusion probably remains considerable for level in Rio de Janeiro State, south-eastern many non-native populations (Kullander & Brazil. This reservoir was filled between 1905 Ferreira 2006). and 1908 mainly for hydroelectric purposes, The anatomy of the digestive tract in fish damming streams and diverting small rivers is closely related to the diet and type of food of the East Hydrographic Basin (Araújo & consumed (Winemiller 1989, Wainwright & Santos 2001). Lajes Reservoir is oligotrophic Richard 1995). In addition, to help with the (i.e. <20mgL-1 total P; <400mgL-1 total N) and taxonomic differentiation of close related Neo- largely surrounded by well-preserved stretches tropical species (Graça & Pavanelli 2007), of Atlantic Rainforest, but it has experienced provide risk assessment of invasive species on recent trends of nutrient enrichment, due to native fish assemblages (Hill et al. 2004), or increased anthropogenic activities (i.e. diffu- contribute to implement control measures for se pollution) and high water retention time non-native populations (Bergmann & Motta (286 days) (Santos et al. 2004, Guarino et al. 2005, García-Berthou 2007), studies on feed- 2005). Because of vegetation removal prior ing morphology can be also used to identify to reservoir filling and wide water level fluc- the length at which fish undergo life history tuations (i.e. up to 12m per year) from dam 2 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 59 (3): 000-000, September 2011 operation routine, Lajes Reservoir is structura- rakers having a common base being counted lly homogeneous, lacking aquatic macrophytes as a single raker. The intestinal quotient (IQ) and other natural submerged structures (Santos was calculated according to Bertin (1958), by et al. 2008). The non-native peacock bass C. dividing the intestine length (LI) per the total kelberi was introduced into the reservoir at the length of the predator (LT). 1950s to control forage fish overpopulation and improve local angling (Santos et al. 2001). Diet characterization: Food items were Despite the probable adverse impacts on the examined using a stereoscopic microscope and indigenous fish species, legal management identified, counted, weighed (to the nearest actions to protecting C. kelberi (i.e. minimum 10-5g) and measured (to the nearest 10-2mm). capture size, spawning season protection and The Index of Relative Importance-IRI (Pinkas bag limits) already exists in Lajes Reservoir. et al. 1971) was calculated for diet characte- rization with percents by number (%N), by Fish collection: Peacock basses were weight (%W) and frequency of occurrence collected monthly from January to December in (%O), where:
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