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REPRODUCTION of SPECIES of the GENUS Cichla in a RESERVOIR in SOUTHEASTERN BRAZIL

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REPRODUCTION of SPECIES of the GENUS Cichla in a RESERVOIR in SOUTHEASTERN BRAZIL REPRODUCTION OF TWO SPECIES OF Cichla 613 REPRODUCTION OF SPECIES OF THE GENUS Cichla IN A RESERVOIR IN SOUTHEASTERN BRAZIL GOMIERO, L. M. and BRAGA, F. M. S. Departamento de Zoologia, Instituto de Biociências, Universidade Estadual Paulista (UNESP), Av. 24-A, n. 1515, C.P. 199, CEP 13506-900, Rio Claro, São Paulo, Brazil Correspondence to: Leandro Muller Gomiero, Departamento de Zoologia, Instituto de Biociências, Universidade Estadual Paulista (UNESP), Av. 24-A, n. 1515, C.P. 199, CEP 13506-900, Rio Claro, São Paulo, Brazil, e-mail: [email protected] Received January 29, 2003 – Accepted August 22, 2003 – Distributed August 31, 2004 (With 7 figures) ABSTRACT The reproduction of two species of Cichla (Cichla cf. ocellaris and C. monoculus, commonly known in Brazil as tucunaré) introduced in the Volta Grande Reservoir, southeastern Brazil, was analyzed. The period for spawning began at the end of the dry season (September) and extended until the end of the rainy season (January). The length of the first gonad maturation (L50) and the length at which all fish are considered adults (L100) were very similar for both species and coincide with the length of fish at one year of age. The analysis of disassociated oocytes evidenced a partitioned spawning, and that C. monoculus was more prolific than C. cf. ocellaris. The low fecundity, partitioned spawning, bi-parental brood care, and supplying of food for the young are related to the way of life, habitat, and reproductive behavior of these fish. Key words: Cichla, reproduction, fecundity, reservoir, Brazil. RESUMO Reprodução de espécies de Cichla em um reservatório do Sudeste do Brasil Foi analisada a reprodução das espécies de tucunarés (Cichla cf. ocellaris e Cichla monoculus) intro- duzidas no reservatório de Volta Grande, Sudeste do Brasil. A época das desovas teve início no final do período de seca (setembro) e estendeu-se até o final do período chuvoso (janeiro). O comprimento da primeira maturação gonadal (L50) e o comprimento a partir do qual os peixes dessas espécies são considerados adultos (L100) foram muito próximos e coincidem com os comprimentos de peixes com um ano de vida. A análise dos ovócitos dissociados evidenciou a desova parcelada, sendo C. monoculus mais prolífico do que C. cf. ocellaris. A baixa fecundidade, a desova parcelada, o cuidado biparental e o suprimento de alimento para os jovens estão relacionados ao modo de vida, ao habitat e ao comportamento reprodutivo desses peixes. Palavras-chave: Cichla, reprodução, fecundidade, reservatório, Brasil. INTRODUCTION excessive proliferation of some and reduction, or even elimination, of others. The impact level Increasing electricity demand requires the is mainly related to local fauna characteristics, construction of many hydroelectric plants that, location of the dam with respect to population as a result of river damming, have innumerable distribution in the area, morphometrics of the consequences for fish populations. A unavoidable basin, existence of other dams upstream, dam result with respect to aquatic fauna is the design, and operational procedures of the plant alteration in abundance of animal species, with itself (Agostinho, 1994). Braz. J. Biol., 64(3B): 613-624, 2004 614 GOMIERO, L. M. and BRAGA, F. M. S. Introduction of species from other basins into complexity of behavioral interactions between parents reservoirs is a common practice. Since the 60s, and their brood, and the amount of food destined significant transfer of native species from the Amazon for the brood is clearly related to the type and extent basin into fish-rearing centers in northeastern Brazil of this care (Godfray, 1995). Male fish with external has occurred, followed by their subsequent transfer fecundation generally have a high probability of being into southeastern and southern Brazil. From these genetically related to offspring hatching at the rearing centers, the species can easily reach nearby spawning site; this is a precondition to the evolution natural water bodies, which incurs high risk, especially of parental care (Blumer, 1979). Cichlids possess in the case of predatory fish species (Agostinho & fins and muscles associated to control of certain Julio Jr., 1996). When environments are altered, native movements including fanning water, an important species are especially vulnerable (Elvira & requirement in lentic waters since it supplies eggs Almodóvar, 2001). An example is the the Volta with a constant water flux for oxygenation and Grande Reservoir where at least 15 years ago two removal of debris from their surface, as well as species of piscivorous fish easily distinguished by minimizing the chance of fungi infection (Zaret, 1980; their external coloration, sagittae, and meristic Blumer, 1986). characters, were introduced (Cichla spp.). Species of the genus Cichla, commonly known Introduced species affect native species through as tucunaré in Brazil, are basically restricted to competition for available resources, predation, pa- transparent waters with high temperatures and access thogen transfer, or simply by habitat alteration. to lentic water habitats for feeding and/or repro- Introduction of alien species and consequent natural duction; in these habitats, they are frequently the habitat loss were the factors responsible for most dominant diurnal piscivores (Winemiller, 2001). animal species extinctions during the twentieth These fish, which lay their eggs on the substrate, century (Elvira & Almodóvar, 2001). Stable in accordance with their reproductive strategy environments are more susceptible to the introduction commonly establish a territory where eggs are laid of new species (specially piscivores) than relatively and guarded. A couple remains together throughout less stable ones; however, every natural system is the parental care period, which extends from the unique and identical changes are rarely possible time of spawning until the immature individuals (Zaret, 1982). Exponential population growth is become totally independent (McKaye, 1977; Zaret, common among introduced predatory species, 1980; Winemiller et al., 1997). bringing with it more intraspecific competition for In species of Cichla reared in captivity, sexu- space and food (Bedarf et al., 2001). As a al maturity occurs between 11 and 12 months of age, consequence, biomass of the introduced species may with the males always being more developed than the decrease (Williams et al., 1998). females (Fontenele, 1950). Before mating, cichlids The family Cichlidae is the most species-rich develop secondary sexual characteristics: post-orbital of all freshwater percoid fish (1,400 species), dis- nuptial humps in the males and bright body colorations playing great variation in reproductive behaviors in both sexes (Fontenele, 1948; Winemiller et al., 1997). (McKaye, 1984; Murray, 2001). Out of the 495 The objective of the present work was to characterize families of bony fish, about 20% exhibit parental the reproduction of Cichla cf. ocellaris (Bloch & care and only two of them, Pomacentridae and Schneider, 1801) and Cichla monoculus (Bloch & Cichlidae, include genera with transitions between Schneider, 1801), as well as its periodicity and kinds of parental care (Gittleman, 1981). The fecundity in the Volta Grande Reservoir. perciforms of the family Cichlidae generally present local sedentary populations, stable food resources, MATERIAL AND METHODS a prolonged breeding season, and parental care, attributes corresponding to an equilibrium strategy The Volta Grande Reservoir, located in a tro- (Winemiller, 1989). Parental care occurs in many pical area (48°25’ and 47°35’W, 19°57’ and different manners, and variation exists as to which 20°10’S), resulted from the damming of the Rio sex takes care of the brood; nevertheless, brood care Grande River. This river is located between the states by both parents is the ancestral condition in relation of Minas Gerais and São Paulo in southeastern to single-parent brood care (Goodwin et al., 1998). Brazil (see Fig. 1). The climate is typically tropical Parental care in animals is determined by the with a mean annual temperature of 22°C and a mean Braz. J. Biol., 64(3B): 613-624, 2004 REPRODUCTION OF TWO SPECIES OF Cichla 615 annual rainfall of 1.635 mm. There are two typical C, mature; and D, spent. The reproductive period was seasonal periods: a warm and rainy season from analyzed according to seasonal variations of the October to April, and a cool and dry season from different maturation stage frequencies during collecting May to September. The Volta Grande Reservoir has periods. The mean length L50 is of the first gonad a flooded area of 221,7 km2 with a water volume maturation at which 50% of the individuals are adult, of 2.268 km3 destined for electric energy production i.e., they have begun the reproductive cycle. The length (Braga & Gomiero, 1997). at which all individuals are apt for reproduction is Collections were made during April and June represented by L100. Both L50 and L100 were estimated 1997, and every month from September 1997 until following Vazzoler (1996). August 1998. Two kinds of fishing equipment were For the study of fecundity, in accordance with used: simple nets with meshes of 2.0, 3.0, 4.0, 5.0, Vazzoler (1996) ovaries in stage C were extracted and 6.0 cm between adjacent knots, and fishing from the visceral cavity and weighted; their mem- rods with natural or artificial bait. Each specific branes were then sectioned longitudinally throughout mesh set was 150 m long and 1.5 m in height,
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