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Contents lists available at ScienceDirect

Fisheries Research
journal homepage: www.elsevier.com/locate/fishres

Fish assemblages in Neotropical reservoirs: Colonization patterns,

impacts and management
Angelo A. Agostinho a,b,∗ , Luiz C. Gomes a,b , Natália C.L. Santos a , Jean C.G. Ortega a ,
Fernando M. Pelicice c
a
Programa de Pós-Graduação em Ecologia de Ambientes Aquáticos Continentais, Núcleo de Pesquisa em Limnologia, Ictiologia e Aquicultura, Universidade
Estadual de Maringá, Laboratório de Ictiologia, Av. Colombo, 5.790, Bloco H-90, CEP: 87020-900 Maringá, PR, Brazil
b
Núcleo de Pesquisa em Limnologia, Ictiologia e Aquicultura, Universidade Estadual de Maringá, Maringá, PR, Brazil
c
Núcleo de Estudos Ambientais, Universidade Federal de Tocantins, Rua 3, Quadra 17, Jardim dos Ipês, CEP 77500-000 Porto Nacional, TO, Brazil

a r t i c l e i n f o a b s t r a c t

Article history: Brazil has more than 700 large reservoirs distributed in all of the major river basins of South America.
Available online xxx Most dams were constructed to produce electricity. Although these reservoirs favor the development of
local and regional economies, they seriously impact the aquatic biota. An unavoidable consequence is
Keywords: the change in the composition and abundance of species, with the proliferation of some and reduction
Freshwater fish or even local extinction of others. The intensity and nature of these changes are related to peculiarities
Dam impact
of the local biota and the location, morphometric and hydrological characteristics of the reservoir, dam
Fish management
operation and interactions with other uses of the basin, including other reservoirs. These impacts exhibit
Fish stocking
Fish pass
substantial spatiotemporal variations. The filling phase is marked by abrupt and intense changes in the key
attributes of aquatic habitats, followed by predominantly heterotrophic processes, with possible thermal
stratification and anoxic conditions. Fish richness increases soon after filling and decreases in subsequent
years. Trophic depletion is expected, and diversity gradients are intensified toward more lentic stretches,
the average length of fish decreases, and the fish fauna becomes dominated by species with sedentary
strategies and/or parental care. The virtual absence of species with pre-adaptations to inhabit lentic areas
of large reservoirs leads to a concentration of biomass in shallow littoral areas. Long-distance migratory
species are the most affected, which include larger fish with high market value. Migratory species require
different biotopes to fulfill their life cycles and strongly depend on the seasonal flood regime, which is
altered due to dam operation. In this study, we discuss the details of these trends as well as the mitigation
measures and management actions that are practiced in Brazil. We conclude that these actions have not
promoted the conservation of fish; on the contrary, some of them have generated additional impacts. As
a consequence, the conservation of Neotropical fish and aquatic resources is severely threatened.
© 2015 Elsevier B.V. All rights reserved.

1. Introduction their life cycle in the new environment and take advantage of the
available food resources will achieve their full potential for prolif-
Impoundments lead to extreme changes in fluvial habitats, eration (Agostinho et al., 2007a). The nature of and intensity with
transforming rivers into semi-lentic systems. Animals and plants which the fluvial biota is altered by impoundments are highly vari-
for which these new conditions are restrictive will have their popu- able among reservoirs and must be studied case by case.
lations drastically reduced. However, species that can complete The literature demonstrates that even reservoirs arranged in
series in the same river, with unidirectional interactions from
upstream to downstream, show distinct peculiarities in relation
∗ Corresponding author at: Universidade Estadual de Maringá, Núcleo de Pesquisa to the colonization process and the organization of assemblages
em Limnologia, Ictiologia e Aquicultura, Programa de Pós-Graduação em Ecologia (Agostinho and Gomes, 1997; Petesse and Petrere, 2012). The
de Ambientes Aquáticos Continentais, Laboratório de Ictiologia, Av. Colombo, 5.790, degree of alteration in the structure and dynamics of the local biota
Bloco H-90, CEP: 87020-900 Maringá, PR, Brazil. Tel.: +55 44 3011 4610. depends on several local and regional factors, such as morphome-
E-mail addresses: agostinhoaa@gmail.com, agostinhoaa@nupelia.uem.br
try of the catchment, discharge, patterns of water circulation, depth,
(A.A. Agostinho), lcgomes@nupelia.uem.br (L.C. Gomes), natalia.ictio@gmail.com
(N.C.L. Santos), ortegajean@gmail.com (J.C.G. Ortega), fmpelicice@gmail.com habitat structure, species pool, surface area, the design of the dam
(F.M. Pelicice). and its operational procedures. Thus, a detailed understanding of

http://dx.doi.org/10.1016/j.fishres.2015.04.006
0165-7836/© 2015 Elsevier B.V. All rights reserved.

Please cite this article in press as: Agostinho, A.A., et al., Fish assemblages in Neotropical reservoirs: Colonization patterns, impacts and
management. Fish. Res. (2015), http://dx.doi.org/10.1016/j.fishres.2015.04.006
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the context of a particular reservoir is paramount for effective mit- Ichthyofaunal monitoring surveys conducted in 77 reservoirs of
igation measures and/or management actions for the conservation the main river basins in Brazil (Agostinho et al., 2007a) showed that
of fish populations (Weithman and Haas, 1982). A manager should, fish diversity in the impounded area is very low. This study showed
based on local and regional studies, identify any alterations in the that 85% of the reservoirs contain fewer than 40 fish species; reser-
structure of the local fish assemblage and take action to avoid voirs with more than 120 fish species are rare and usually young.
irreversible losses of regional biological diversity and/or natural Forty species can be considered very low if we consider that 80%
resources as a consequence of river damming. of these reservoirs have areas greater than 10 km2 and that a sin-
In general, the fish species most affected by impoundments are gle floodplain lake of much smaller dimensions can harbor from 30
large in size, migrate and have high longevity (k-strategist). In con- species (Paraná River basin; Oliveira et al., 2001) up to 99 species
trast, a massive proliferation of primarily small-sized sedentary (Amazon River basin; Pouilly et al., 2004). In addition, streams and
species (i.e. those that do not migrate) occurs, which have a high rivers in the Neotropical region usually present hundreds of species
reproductive potential and short longevity (r-strategists) and for (Lowe-McConnell, 1999; Agostinho et al., 2007b), e.g. a stream less
which the availability of food resources is high (Agostinho et al., than 10 km long had 108 species (Cancela Stream; Cuiabá River
1999, 2008a; Hoeinghaus et al., 2009). Yet, sedentary species are basin; Mendes et al., 2008). However, species richness in reservoirs
also affected by hydrological alterations and tend to redistribute varies with their surface area, age and, primarily, the basin where
along the river/reservoir gradient (Araújo et al., 2013). In the inner they are located. Thus, reservoirs located in the Amazon basin with
areas of large reservoirs, fish assemblages are profoundly altered areas greater than 500 km2 and less than 15 years old contain more
and composed of a few species with pre-adaptations to live in semi- species than other reservoirs of similar dimensions and age that are
lentic environments (Gomes and Miranda, 2001; Agostinho et al., located in other Neotropical basins. For example, more than 200
2007a). fish species were found in the São Salvador Reservoir, Tocantins
Reservoirs are present in the main river basins in Brazil, and River (104 km2 ; Amazon basin), in the first years after impound-
the principal purpose is the production of electricity. Although ment (Limnobios, 2014). In contrast, 34 species were recorded in
reservoirs are widespread in the country, their distribution is not Segredo Reservoir (85 km2 ; Iguaçu River; Agostinho and Gomes,
homogeneous, e.g. the Upper Paraná River has half of the total 1997) and 107 in Itaipu Reservoir (1350 km2 ; Agostinho et al., 1992)
impounded area and is one of the most regulated rivers in the in a similar time lag. Furthermore, in Capivara Reservoir (576 km2 ;
world (Agostinho et al., 2008a). Even considering the specificity Paranapanema River; Orsi and Britton, 2014) and Salto Osório
of the response of the biota to the impacts generated by each reser- (63 km2 ; Iguaçu River; Baumgartner et al., 2006), both impound-
voir, some patterns can be described based on studies of dozens ments are older than 30 years, were recorded 41 and 23 species,
of reservoirs in Brazil. Therefore, the objective of this paper is to respectively. In fact, there is a consistent decrease in species rich-
review the patterns of fish fauna once a reservoir is formed. First, ness over time (Mol et al., 2007; Orsi and Britton, 2014), i.e. the
we described the variation in fish assemblages over time, from the number of species averages 20 in Neotropical reservoirs older than
filling of the reservoir to the periods in which environmental and 20 years (Agostinho et al., 2007a). This conspicuous decline in
biotic conditions are rearranged and more stable. We categorized species richness is the result of environmental filters that gradually
these variations into phases (heterotrophic, post-heterotrophic and remove pre-existing fluvial species; the new assemblages are com-
trophic equilibrium), considering predicted alterations in produc- posed basically of species that present pre-adaptations to thrive in
tivity. Then, considering the phases, we described broad trends standing waters, with lower dependence on fluvial environments
in fish abundance, species richness, pre-adaptations to pelagic and habitat heterogeneity (Gomes and Miranda, 2001).
environments, and variations in size and reproductive strategies.
Finally, we evaluated management measures presently imple-
mented to mitigate impacts caused by reservoirs on the Neotropical 3. Variation in fish abundance
fish fauna, and we discuss opportunities for improvement as well
as the existing knowledge gaps. As the Upper Paraná River basin is The large release of nutrients resulting from the decomposition
the most dammed in South America as well as the most studied, of organic matter in the flooded area during a reservoir’s early years
we used it as a model to achieve our goals every time an example and the subsequent reduction of nutrients result in wide fluctua-
was necessary. tions in production throughout a reservoir’s history. The nutrient
input increases the production of all trophic levels during a period
known as the “trophic upsurge period” (Kimmel and Groeger, 1986;
Kimmel et al., 1990). This heterotrophic period begins in the fill-
2. Reservoirs and fish diversity ing phase, which is marked by rapid and profound alterations in
the water’s physical and chemical characteristics. During the filling
It is estimated that the number of large reservoirs (dams higher phase, vertical patterns resulting from the expansion of the water
than 15 m; World Commission on Dams, 2000) in South America is column, lentic characteristics and thermal stratification, which
greater than one thousand, and around 50% of them are located affect the sedimentation rate, nutrient cycling and the distribution
within Brazilian territory (Fig. 1). Thirty-seven percent of these of the biota, are added to the predominant transport vector of the
reservoirs produce electricity. Although hydroelectric production river phase. The high concentration of nutrients initially due to the
in dams started in Brazil at the end of the XIX century (Marmelos pulses of litter decomposition and the release of nutrients from
Dam; Paraíba do Sul River; 1889), most of the dams were con- the inundated soil, followed by the decomposition of the leaves of
structed in the second half of the XX century. With regard to the the inundated vegetation (Cunha-Santino et al., 2013), may lead to
area inundated by all reservoirs (>36,000 km2 ), almost half of it stressful conditions for the aquatic biota (e.g. low concentrations of
(47%) is located in the Paraná River, followed by the São Francisco dissolved oxygen, thermal stress, and low pH), especially near the
and Tocantins Rivers (Agostinho et al., 2007a). As potential areas bottom (Agostinho et al., 2008a).
for the installation of new dams in these basins are depleted, there For example, studies conducted in Corumbá Reservoir (located
is a motivation to extend the construction of dams to the Amazon in the Upper Paraná River basin) showed a sharp increment in
basin, especially in the Madeira, Tapajós and Xingú Rivers (Castello primary production after an initial period of increased water
et al., 2013), in addition to the Andean tributaries (Finer and Jenkins, transparency (Secchi depth) due to sedimentation. Thus, the phyto-
2012). plankton productivity that was below 0.17 mgO2 l−1 in the first 10

Please cite this article in press as: Agostinho, A.A., et al., Fish assemblages in Neotropical reservoirs: Colonization patterns, impacts and
management. Fish. Res. (2015), http://dx.doi.org/10.1016/j.fishres.2015.04.006
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Fig. 1. Map showing the distribution of hydroelectric reservoirs in the main river basins of Brazil (Paraná River basin is highlighted due to the high number of dams).

days after the reservoir began filling, reached 0.89 mgO2 l−1 after which the abundance of fish tends to be less variable but usually
39 days (Agostinho et al., 1999). Therefore, due to this increase in higher than in the river before the dam was constructed.
primary productivity, it is common a marked increase in the fish To exemplify this decrease in productivity over time, we used
abundance in the inner areas of the reservoir (Fig. 2). The increase sample data collected in the Itaipu Reservoir (Paraná River basin)
in nutrient concentration is also responsible for the intense pro- from 1983 (one year after filling in 1982) to1997 (15 years after
liferation of floating macrophytes, as verified in Tucuruí Reservoir, filling). In this reservoir, there was a clear temporal decrease in
located in the Tocantins River basin (Tundisi, 1994). The presence fish abundance in number (catch per unit effort, CPUE—number of
of an anoxic layer during filling is also an event common to tropical individuals) and in weight (CPUE—kg), especially in the more inter-
reservoirs and can last for months or even years. However, after nal areas (lacustrine zone). Note that the decreased abundance was
filling, a reduction in fertility due to the loss of organic matter by from two- to four-fold in number and weight, respectively (Fig. 3).
oxidation, sedimentation, biological assimilation and exportation A clear decrease in fish abundance was noted for all zones (it was
is common (Cunha-Santino et al., 2013); this period is known as less noticeable in the transitional zone), but a sharper decrease was
the “post-heterotrophic” or “depression” period (Fig. 2). observed in the inner areas (lacustrine zone) of the reservoir. Fish
Once the phase of high productivity is over, fish species begin to biomass showed the same trends (Fig. 3). These results demon-
adjust to the new environment (Petrere, 1996). The high fish abun- strate that the degree of the impact of the impoundment on fish
dance verified during this phase tends to decrease in the reservoir abundance or biomass has a longitudinal gradient. Orsi and Britton
over time (Fig. 2). This decrease will continue until the reservoir (2014) also reported a sharp decline in native fish abundance 40
reaches a certain trophic equilibrium (at an unknown time), after years after the formation of Capivara Reservoir, which involved

Please cite this article in press as: Agostinho, A.A., et al., Fish assemblages in Neotropical reservoirs: Colonization patterns, impacts and
management. Fish. Res. (2015), http://dx.doi.org/10.1016/j.fishres.2015.04.006
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environment. Therefore, the changes in the physical and chemical

characteristics of the water due to the beginning of reservoir fill-
ing may act as environmental filters, selecting for ecological traits
such as trophic guilds, reproductive strategies and an alteration
of the affinity for habitats (fidelity), which determine the success
of colonization by a particular species. Species that successfully
colonize a reservoir have the ability to search for adequate environ-
ments, such as lotic tributaries or even the littoral areas (Agostinho
et al., 2007a), as well as those that develop strategies different from
those exhibited in the previous lotic environment (Kubečka, 1993).
Species with pre-adaptations to live in lacustrine environments are
Fig. 2. Trends in fish abundance (catch per unit of effort—CPUE) over time in obviously selected to compose the new assemblages (Fernando and
Neotropical reservoirs (modified from Petrere, 1996; Agostinho et al., 2007a). Holcik, 1991; Gomes and Miranda, 2001).
Colonization during the filling of the Salto Caxias (Iguaçu
River Basin) and Corumbá (upper Paraná River Basin) reser-
voirs exemplifies the patterns of occupation of the new floodable
area. To study this phenomenon, we categorized the fish species
according to habitat preference (benthonic, bentho-pelagic and
pelagic) and capture location in relation to the longitudinal gra-
dient (riverine and lacustrine zones) and habitat (littoral, pelagic
or bathypelagic). During the filling phase of these reservoirs,
a greater abundance of benthonic species was observed in the
littoral (Salto Caxias Reservoir—two-way ANOVA; Interaction,
Zones × Habitat Type; F2, 6 = 5.99, p = 0.037; Fig. 4a) and pelagic
habitats (Corumbá—two-way ANOVA; Interaction Zones × Habitat
Type; F1, 8 = 7.90, p = 0.023; Fig. 4b), respectively. In the riverine
zone, as expected, we found the opposite pattern, with benthonic
species occupying deeper strata. A similar pattern was observed
for bentho-pelagic fish, with a greater capture rate in the littoral
(F2, 6 = 5.98, p = 0.037; Fig. 4c) and pelagic (F1, 8 = 14.00, p = 0.006;
Fig. 4d) areas. In contrast, pelagic species were captured in very
low abundance in both reservoirs and did not show any differ-
ences in abundance according to the zone and habitat type (all
possible results with p > 0.05). This result is due to the existence of
few pelagic species in the Neotropical region (Gomes and Miranda,
2001; Araújo et al., 2013).
The results presented for Salto Caxias and Corumbá reveal low
habitat fidelity for benthic species in the inner part of the reser-
voirs during the filling phase. In fact, fish use a habitat according
to physiological convenience, which depends primarily on the con-
centration of dissolved oxygen and water temperature (Prchalová
et al., 2009); their vertical distribution is apparently driven by
restrictions related to thermal and dissolved oxygen stratifica-
tion in the reservoir. This stratification, during the first year after
impoundment, can lead to a chaotic pattern of species occupy-
ing habitats in which they were previously not abundant (e.g.
benthonic species abundant in the littoral or pelagic zones of reser-
Fig. 3. Variations in the catch per unit effort (CPUE; individuals and weight for
voirs). Yet, the reassembly of fish species following changes in
1000 m2 of gillnet in 24 h) in the longitudinal gradient of the Itaipu Reservoir from
1983 (one year after the impoundment) to 1997 (white circles: CPUE in numbers; environmental conditions may occur within the first years after
black circles: CPUE in weight). impoundment, creating new diversity patterns along the reservoir
(Araújo et al., 2013).
After the filling of the reservoir and the beginning of dam opera-
the loss of 27 species; at present, non-native opportunistic species tion, critical conditions of dissolved oxygen may persist, depending
dominate the assemblage in Capivara. on the extension of the anoxic layer and the vertical position of
the water intake for turbines and spillway. These conditions may
4. Fish colonization during the filling phase lead to a narrow oxygenated layer, resulting in instability due to
wind and temperature changes, which can culminate in fish mor-
During reservoir filling, the patterns of vertical colonization of tality concentrated near the margins or the surface (Agostinho et al.,
fish are associated with thermal stratification, an increase in depth 1999).
and a sharp decrease or even the virtual absence of dissolved oxy-
gen. All these factors impose changes on fish distribution patterns.
The increase in water volume and the reduction in water flow lead 5. Heterotrophic and trophic equilibrium phases
to an increase in the area available for colonization (Agostinho et al.,
2008a; Wang et al., 2013). The lack of oxygen in the deeper strata There is evidence from several Neotropical reservoirs that the
(in the bathypelagic zone) may lead species to disperse in vertical species richness increases immediately after the filling phase
and horizontal directions or even upstream, far from the lacustrine (Fig. 5a and b). This increase in species richness is followed by an

Please cite this article in press as: Agostinho, A.A., et al., Fish assemblages in Neotropical reservoirs: Colonization patterns, impacts and
management. Fish. Res. (2015), http://dx.doi.org/10.1016/j.fishres.2015.04.006
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Fig. 4. Mean catch per unit of effort (CPUE; log10 x + 1 transformed; vertical lines are the standard errors) for the species categorized as benthonic (a and b), bentho-pelagic
(c and d) and pelagic (e and f) among the distinct reservoir zones (riverine and lacustrine) and types of habitats (littoral, pelagic and bathypelagic). (a), (c) and (e) Filling
phase of the Salto Caxias Reservoir (1998); (b), (d) and (f) Filling phase of the Corumbá Reservoir (1996). Note that in Corumbá, samples were taken only from the pelagic
and bathypelagic regions.

increase in the abundance of fish (Fig. 5c), which is common during cycle (Lowe-McConnell, 1999; Agostinho et al., 2007b; Araújo et al.,
the trophic upsurge period. However, the magnitude of the increase 2013; Franssen and Tobler, 2013).
in abundance varies among species in a new reservoir, and the dom- Abundance follows a similar trend. Upon filling of a reservoir,
inance of certain species with regard to abundance (low evenness) fish abundance increases (Fig. 5c) due to the high input of terres-
causes a continuous decreasing in the species diversity measured trial organic matter, which leads to increased food availability in
by the Shannon index (Fig. 5a). the entire reservoir, especially for omnivorous, herbivorous and
An increase in species richness is expected during filling because insectivorous species. The proliferation of these species causes an
different biotopes, such as wetlands, isolated lakes, lakes perma- increase in food availability for piscivores. However, at the end of
nently or seasonally connected to the river channel and adjacent the heterotrophic phase (see Fig. 2), the abundance of fish decreases
tributaries (river, streams and creeks), are incorporated into the following the decrease in primary production (see Section 3).
new environment. Species associated with these habitats are incor-
porated into the fish fauna of the reservoir, which consequently 6. Constraints and pre-adaptations
increases species richness. The number of species present in a
recent reservoir should not be much lower than the sum of the The virtual absence of natural lakes in Brazil (excepting those
previously existing species in the flooded habitats. However, this associated with fluvial corridors) and the consequent scarcity of
tendency for high richness does not last long (Fig. 5a). The rea- species with pre-adaptations to occupy open areas of reservoirs,
sons for its decrease have been previously discussed and appear to allied with the longitudinal gradients related to the processes of
be related to environmental filters, species sorting and the accom- transport and deposition (e.g. transparency and nutrient loads),
modation of the fish fauna to the new environment, in addition leads to a heterogeneous pattern of the occupation of the new
to trophic depletion and the absence of truly lacustrine species environment. The most important characteristics of truly pelagic
(Agostinho et al., 2007a). Theoretically, the drop in species richness species are their short food chains, high fecundity, pelagic adap-
results from the movement of fish out of the reservoir (upstream tations, and short life cycle, as exhibited by the Clupeiformes
or tributaries) in search of better conditions to complete their life Stolothrissa tanganicae (in Africa) and Dorosoma cepedianum (in

Please cite this article in press as: Agostinho, A.A., et al., Fish assemblages in Neotropical reservoirs: Colonization patterns, impacts and
management. Fish. Res. (2015), http://dx.doi.org/10.1016/j.fishres.2015.04.006
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Fig. 6. Spatial (longitudinal, lateral and vertical gradients) and temporal gradients
in species richness and abundance of fish in the Itaipu Reservoir (Riv = Riverine;
Tra = Transitional; Lac = Lacustrine Source: Agostinho et al., 1999).

This pattern can be verified in the Itaipu Reservoir five (1987)

and 15 (1997) years after its formation (Fig. 6). Species richness and
the abundance of fish were considerably higher in the littoral zone,
Fig. 5. Variations in the Shannon diversity index ((a) before and two periods after and this pattern tended to increase over time. After 15 years, 64 out
the formation of Jordão Reservoir; numbers in brackets are species richness and of the 67 species captured in gillnets in the Itaipu were in the littoral
evenness), species richness ((b) before and two periods after the formation of four
zone, whereas in the pelagic and bathypelagic zones, this number
reservoirs in the Upper Paraná River basin) and abundance ((c) catch per unit
effort—CPUE ind. 1000 m2 gillnet in 24 h before and after the formation of four was 22 and 20, respectively (Fig. 6). The proportions of abundance
reservoirs in the Upper Paraná River basin). Modified from Agostinho et al., (2007a). among the zones were 19.4: 1.1: 1.0, respectively (Fig. 6). Moreover,
the riverine zone, in which the processes of transport predominate
over the depositional processes, has higher species richness but
is not the most productive zone (Kimmel et al., 1990; Agostinho
North America) (Gomes and Miranda, 2001). Thus, in reservoirs, et al., 2007a). The upper third of the Itaipu Reservoir harbors all
the colonization success of the species depends on their pre- of the species recorded in the two more internal thirds in addi-
adaptations. In the Upper Paraná River, Gomes and Miranda (2001) tion to those typical to the lotic stretch upstream (the river). The
described that, among the 220 species they analyzed, only approx- higher similarities in flow with the original river, the lower depth,
imately 5% were considered as lacustrine adapted (i.e. Plagioscion the input of allochthonous matter, and predator attraction due the
squamosissimus, Hypophthalmus edentatus). higher abundance of prey species in relation to the upstream stretch
In general, the pelagic areas of the Upper Paraná Reservoirs may explain this pattern (Agostinho et al., 2007a; Araújo et al.,
are inhabited by few fish species, such as the piscivores P. squa- 2013).
mosissimus (a sciaenid introduced from the Amazon basin) and Reproduction, due to its more conservative nature, imposes lim-
Rhaphiodon vulpinus, and the planktivores H. edentatus and Hemio- itations on the occupation of a new reservoir by the river fish fauna;
dus orthonops (the latter was recently introduced through the fish it is probably the main constraint limiting fish fauna reassembly.
passage at Itaipu Dam; Julio et al., 2009; Agostinho et al., 2015). In reservoirs, it is expected that species with higher plasticity in
For example, the success of P. squamosissimus may be attributed the selection of spawning sites have more success in the coloniza-
to its reproductive strategy (Agostinho et al., 1999). This species tion of these environments. Species that demand particular habitats
produces small, pelagic (Fontenele and Peixoto, 1978), and buoyant (e.g. tributaries) or environmental triggers (e.g. hydrological varia-
eggs, spawned in several batches early in the reproductive period tion) may not complete the reproductive process, mainly during
(matching with food availability), and the larvae are also pelagic the years following the impoundment. Medeiros et al. (2014)
(Nakatani et al., 1993). Other important characteristics are morpho- reported failed reproduction for Hemiodontidae after the forma-
logical and related to diet and food capture (Mérona and Vigouroux, tion of Lajeado Reservoir, Tocantins River, an event that changed
2012). Therefore, the low number and uneven distribution of large energy allocation patterns for these species. However, most of
and deep natural lakes in the Neotropical region led to the complete the species that inhabit reservoirs search for lateral tributaries,
absence of a truly pelagic and deep bottom-dwelling species that upstream stretches or other lotic areas for spawning, indicating the
are pre-adapted to occupy open areas of large reservoirs. In general, dependence on riverine habitats to complete their life cycles. In the
species that successfully colonize reservoirs are those that inhabit first years after the formation of a reservoir, internal fecundation
shallow floodplain lakes, which usually occupy the littoral region of appears to be a successful strategy. However, in older reser-
reservoirs (Casatti et al., 2003; Pelicice et al., 2005; Agostinho et al., voirs, species with more elaborate reproductive strategies (usually
2007a). Thus, the greatest abundance of fish species and diversity cichlids with complex mating choice, nest-building and parental
are found in the littoral region. care) have greater occupation success, along with small-sized

Please cite this article in press as: Agostinho, A.A., et al., Fish assemblages in Neotropical reservoirs: Colonization patterns, impacts and
management. Fish. Res. (2015), http://dx.doi.org/10.1016/j.fishres.2015.04.006
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Fig. 7. Pearson correlation coefficient between fish abundance in several size classes
and the age of the Itaipu Reservoir (1983–1997) (Source: Agostinho et al., 1999).
Fig. 8. Ordination from the detrended correspondence analysis (DCA; Axis 1 and
2—DCA1 and DCA2) applied to summarize the data on fish life strategies col-
opportunistic characids that colonize shallow shores (Agostinho lected in the Itaipu Reservoir (S = small-sized species, Ls = 20 cm; m = median-sized
species, Ls = 20–50 cm; l = large-sized species, Ls > 50 cm; adapted from Agostinho
et al., 2007a).
et al., 1999).
A clear consequence of the species sorting process is the
reduction in the mean size of the individuals comprising a fish
assemblage that occupies a reservoir in relation to those that which were the only group of species captured at the end of the
originally comprised the original river. This trend is important study (Fig. 8).
because of its implications concerning the profitability of fisheries Changes in the mean size of the fish fauna after reservoir for-
(Agostinho et al., 2008a), and it is clearly seen in the size distribution mation, as noted above, are mainly a result of the evasion of
of the fish captured in experimental fisheries in the Itaipu Reser- migratory species. These species are generally large-sized carniv-
voir from 1983 to 1997 (Fig. 7). The Pearson correlation between orous or herbivorous fish and are the most preferable for human
the number of fish in several size classes and the reservoir age consumption, therefore demanding a higher market value. Moni-
decreased (or was negative) in the greater size classes, with high toring of the artisanal (commercial) fishery conducted in the Itaipu
positive values for smaller fish. The main reasons for these findings Reservoir revealed the changes in the composition of landed fish
are: (i) long-distance migratory species, usually large-sized, move with reservoir aging (Hoeinghaus et al., 2009). Before the con-
out of the reservoir; (ii) small-sized species (usually r-strategists) struction of the Itaipu Dam, most captured species were large,
are the tolerant species or those with pre-adaptations to thrive migratory fish (Fig. 9). Five years after the reservoir formation, only
in the lentic environment, especially in the littoral zone, where one migratory species captured prior to the dam construction pre-
they are abundant (Agostinho et al., 1999; Hoeinghaus et al., 2009). sented high abundance, but this species was a detritivore (Curimba,
As a consequence, the new fish fauna is essentially composed of Prochilodus lineatus). Fifteen years after the reservoir formation,
species with small body sizes that occupy shallow littoral areas of the artisanal fishery was essentially maintained by non-migratory
the reservoir. or non-native species with low market values. A lotic environ-
ment and the blockage of migration routes are critical aspects for
the maintenance of migratory species stocks in reservoirs, and the
7. Fish assemblage stabilization
replacement of migratory species by coarse fish and usually less
valuable causes substantial losses to the fishing industry. These
The time span after reservoir closure that the fish community
changes clearly illustrate the aging process of the reservoir, in
structure requires for a certain degree of stability varies widely,
which the river system develops into a new and different stable
and no consensus concerning this time span exists in the litera-
state after impoundment, with different productivity levels, species
ture (Petrere, 1996). There are evidences of stabilization of the fish
composition, distribution, biomass and services for society. Future
abundance and species richness between 15 and 40 years after a
research must investigate the time span to reach this new state,
reservoir is formed (Mol et al., 2007; Orsi and Britton, 2014). Several
and the factors responsible for oscillations and stabilization in the
factors may influence this time, such as latitude, hydraulic reten-
structure of the fish fauna.
tion time, morphometry, fish fauna composition before damming,
catchment area, position in the basin, the presence of large tribu-
taries, and the design and operation of the dam (Agostinho et al., 8. Management and impact mitigation
2007a).
Patterns of dam operation and fluctuations in the water level The search for measures to mitigate impacts caused by dams
of the reservoir may cause a constant perturbation, decreasing the and associated reservoirs on fish diversity and fish stocks in Brazil
potential for the fish community to reach stability. These constant began with the construction of the first hydroelectric reservoirs.
perturbations induce oscillations in the abundance of r-strategists The first action taken was the construction of the fish ladder in
species. As noted above, periodic/seasonal species such as long- the Itaipava Dam (Pardo River, Upper Paraná River basin) at the
distance migratory species abandon the reservoir area. Therefore, beginning of the last century. After that initiative, the history of
the fish fauna of these reservoirs, over time, will be dominated by management in Brazil encompassed several phases, with emphasis
opportunistic (r-strategist) sedentary species and/or the ones that on fish stocking, fish farming in cages (in the reservoirs), and fishery
develop parental care (k-strategist). A detrended correspondence control. The results obtained after a century of management were
analysis (DCA) applied to summarize the data on fish life strategies insignificant, leaving key questions unanswered and generating
in the Itaipu Reservoir (Agostinho et al., 1999) serves as an example. great controversy regarding the effects of the fisheries manage-
In the ordination (Fig. 8), long-distance migratory and large-sized ment strategies. However, these results and the controversy are
species (standard length—Ls > 50 cm) were registered in the reser- due to (i) the still incipient knowledge about the Neotropical fish
voir in the first years after impoundment. The opposite trend was fauna, which is characterized by high diversity level in different
verified for sedentary and small-sized species (Ls = 20 to 50 cm), scales; (ii) the absence or inadequate monitoring of the results of

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Fig. 9. Decade tendencies of the dominant species in the landings of the artisanal fishery conducted in the Paraná River before and after the formation of the Itaipu Reservoir.
The years presented are five years before the reservoir was formed and five and 15 years after the formation of the reservoir (adapted from Agostinho et al., 2007a).

the implemented actions; (iii) the eminently political nature of the to reach the reservoir and dispersed to the upstream stretches of
management decisions, which should be essentially technical; (iv) the watershed (Makrakis et al., 2007a; Julio et al., 2009; Vitule et al.,
insufficient knowledge about the problems to be solved, which led 2012). An emblematic example was H. orthonops, absent from the
to a lack of clarity in the objectives of management actions; and upper Paraná River Basin. The invasion of this species was note-
(v) the naïve belief that impacts caused by impoundments can be worthy for both its fast colonization of the new environment and
reversed or minimized with simple management actions or copied for its abundance, reaching approximately 8% of the total catch
from other part of the world. at the upstream plain, in less than five years (Agostinho et al.,
Concerns with impoundment impacts on migratory species 2015).
populations led to the recommendation to construct fish passages However, effective monitoring of the performance of these pas-
in the dams. This action attempted to facilitate the transit of fish sages began only in the 2000s; even though fishways have been
to their spawning or feeding sites or the dispersal of juvenile fish installed for a century. Yet, most of the current studies are restricted
to downstream stretches of the basin. The installation of these to monitoring species in the fish passage (fish ladders and fish
passages was mandatory for decades (Decree 4390 of 1928), and elevators) with no consideration of the availability of adequate
it is still mandatory in some Brazilian states. However, the con- habitats for the species in the upstream stretches or in the region.
struction requirements and the use of standardized protocols to Tagging studies on fish movements are recent (Hahn et al., 2007;
install fish passages, whose performance depends on the interac- Fontes et al., 2012; Wagner et al., 2012), and these studies have
tion between their technical characteristics and the nature of the not eliminated the controversies regarding the adequacy and the
local ichthyofauna, were at high risk of failure, wasting financial efficiency of fish passages. The controversial aspect on the fish
resources, effort and opportunity (Agostinho et al., 2002; Pelicice passage issue refers mainly to its simplicity and convenience for
and Agostinho, 2008; Pompeu et al., 2012). In fact, some fish pas- management programs that are mandatory in Brazil, but with low
sages were built immediately upstream of natural barriers that significance for conservation, i.e. aiding recruitment of migratory
fish historically did not cross (Charlier, 1957). Other passages fish (Pelicice and Agostinho, 2008; Pompeu et al., 2012).
were highly selective, allowing the passage of large numbers of Other relevant aspects that should be considered in discussions
sedentary species and restricting the passage of the migratory on fish passages are their high selectivity (Agostinho et al., 2007d),
species (Agostinho et al., 2007b; Makrakis et al., 2007a). In addition, the difficulty in controlling which species go through the passage
invasive species previously limited by natural barriers have been (Pompeu et al., 2012), and the absence of downstream movement
reported to be greatly dispersed along river channels, as the case of adults and their offspring (Agostinho et al., 2007c, 2011; Suzuki
of the Itaipu Dam in the Paraná River, where the Piracema Canal is et al., 2011; Pelicice and Agostinho, 2012). Solutions to the existing
located (Julio et al., 2009; Makrakis et al., 2007b; Agostinho et al., bottlenecks concerning recommendations of fish passages as a tool
2015). In this case, the Itaipu Reservoir, filled in 1982, covered the to mitigate impacts on migratory species must address the follow-
Sete Quedas Falls, which was the limit for the distribution of several ing issues: (i) whether the passages are efficient to attract fish and to
species, separating two distinct ichthyofauna provinces, the upper allow free movements; (ii) whether the reservoirs represent a bar-
and the middle Paraná (Bonetto, 1986). After the Itaipu Reservoir rier to downstream movement of adult fish or to the drift of their
was completely filled, several species were able to reach the upper eggs and larvae (Agostinho et al., 2007c; Pelicice et al., in press),
part of the basin (Julio et al., 2009). Some of these species became (iii) whether long-distance migratory species have distinct behav-
abundant and replaced native congeneric species (Agostinho, 2003; iors and the swimming ability to be attracted to and to overcome
Alexandre et al., 2004). During the following 20 years, approxi- the water flow in a fish passage; (iv) whether the passage is safe,
mately 17 species remained restricted to the stretch downstream with low rates of injury or predation (Agostinho et al., 2012), (v)
from the Itaipu Dam. However, after the Canal de Piracema (a whether clear objectives exist (e.g. genetic and/or demographic) to
natural-like fish passage) started operation, other species were able justify the use of fish passes, and (vi) whether the regional context

Please cite this article in press as: Agostinho, A.A., et al., Fish assemblages in Neotropical reservoirs: Colonization patterns, impacts and
management. Fish. Res. (2015), http://dx.doi.org/10.1016/j.fishres.2015.04.006
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(i.e. distribution of critical habitats) supports the use of fishways to stock (Flagg and Nash, 1999); (v) the compatibility of the quantity
achieve conservation goals (Pelicice and Agostinho, 2008; Pompeu of fish, size, place and timing of the release with the distribution
et al., 2012). While these aspects are neglected, the decision on the and structure of natural populations (Molony et al., 2003); and (vi)
construction of fish passages will remain nebulous, with the risk of monitoring of the stocking and wild populations. In fact, monitor-
causing further impacts and complicating alternative conservation ing programs should be an integral and indissoluble component of
efforts. stocking, and the results obtained should be the base for adjusting
Another controversial management action has been fish stock- or even halting the procedures.
ing. The first stocking initiatives were conducted with non-native Aquaculture, in a strict sense, is not considered a management
species in northeastern Brazil and were successful in producing activity destined to mitigate impoundment impacts. However,
self-sustaining populations and improving fishing yields (Paiva aquaculture has been conducted under the argument that it mini-
et al., 1994). This experience, especially with non-native species, mizes fishing pressure on wild stocks, either by the involvement
spread to other regions of Brazil and was the main fishery of the fishermen in production activities (farming) or by a reduc-
management activity conducted by the Brazilian fishery-related tion in the demand for wild fish (Agostinho et al., 2007a). Although
institutions and by power companies. Until 1990, non-native considered an important food production activity, aquaculture, as
species were emphasized in stocking programs developed in south- any other production method, affects the environment with an
eastern and southern Brazil. Some of the stocked non-native species intensity that varies according to the type (intensive or extensive)
were successful colonizers and are currently widespread in many and the species farmed. Such impacts are evident in the intensive
basins (e.g. silver croaker P. squamosissimus; peacock basses Cichla farming conducted in caging nets installed in Brazilian reservoirs,
spp.). Other species were successful in some reservoirs, where they which has received subsides from the governmental financial agen-
appear in high abundance (e.g. tilapias—mainly Oreochromis niloti- cies related to fish production (Agostinho et al., 2008b; Lima et al.,
cus, oscar Astronotus ocellatus, and freshwater sardine Triportheus 2012). Although fish farming in cages has not been adequately mon-
angulatus). Although stocking activities currently emphasize native itored, preliminary studies already indicate some distortions with
species, monitoring of commercial (artisanal) fishery outputs regard to proposed objectives, conflicts among users, profitability,
shows that stocking has not been efficient and might even repre- introduction of species, and aquaculture as a source of water quality
sent an additional source of impacts (Agostinho et al., 2004, 2010). degradation (Agostinho et al., 2007a; Strictar-Pereira et al., 2010;
There are no relationship between stocking efforts and captures Azevedo-Santos et al., 2011; Pelicice et al., 2014). Given the com-
(landed fish) in artisanal fisheries in the reservoirs of southeast- mon occurrence of escapes in aquaculture, the use of non-native
ern and southern Brazil, where stocking was more intense. Some of species was prohibited in reservoirs where these species were not
the stocked species were never captured in the fisheries (AES-Tietê, established. However, this restriction was removed by a Federal
2007). Decree, which provided the status of “native” to several species
Stocking programs were historically conducted based on a pre- from other continents (i.e. tilapias species), as a mean of fostering
carious knowledge of the system to be managed, of the species aquaculture in large reservoirs (Vitule et al., 2012; Pelicice et al.,
to be released, and of the real need for the action. In addition, 2014). This decision may increase non-native dispersion across
inexperience regarding how to conduct stocking (which species, South American basins; it is well known that aquaculture is the
the necessary quantities, the appropriate location, the size of the main source of non-native species to Neotropical reservoirs (Ortega
fish, and the time of release, among others) led to the practice et al., 2015). In addition, fish farming of native species in cages
of “trial and error.” Furthermore, stocking has been conducted conducted by traditional fishermen has not been promising due to
without monitoring and without learning from the past actions, the high costs of production, difficulties in commercialization, and
which can be helpful in avoiding future mistakes (Gomes et al., small-scale production stemming from the investment capacity of
2004; Agostinho et al., 2007a; Pelicice et al., 2009). For example, the fishermen. Regardless of these negative points, aquaculture
knowledge of the carrying capacity of the receptor environment in public waters (reservoirs) may be environmentally sustainable
and the size of wild stocks are fundamental assumptions in stock- and may promote social development, generating income and jobs.
ing for supplementation (Cowx, 1999), and such knowledge has However, such a system requires a program with ample interaction
been ignored in the stocking programs conducted in Brazil. Addi- with other activities related to fishery resources, created with rig-
tionally, the processes used to rear fish for stocking programs are orous planning and sustained by technical studies on production,
frequently the same used to produce fish for farming; in fact, fishes impacts and marketing (Agostinho et al., 2008b). Unfortunately,
for both purposes have been reared together in fish farms. Thus, aquaculture in Brazilian reservoirs does not follow these high
by ignoring the genetic quality of the brooders and other possi- environmental standards, and constitutes an additional source of
ble negative impacts on natural populations, stocking became a disturbance to wild freshwater fish (Agostinho et al., 2007a; Pelicice
potential and constant threat to local populations and to the fishery et al., 2014).
itself, although such consequences were never empirically studied Control of the fishery activity is an ongoing alternative to man-
(Agostinho et al., 2010). It should be highlighted that stocking pro- aging reservoirs in Brazil. However, there are also huge practical
grams are supported by society, based on the naïve belief that fish and conceptual difficulties to overcome. In general, fishing in reser-
populations were impacted, declined and must be recomposed in voirs has already begun in the heterotrophic phase just after filling,
the reservoirs; stocking, in this sense, is a valid compensation or when the harvest is high. In this phase, a great number of fisher-
mitigation measure to address the impact of impoundments on fish men engage in the activity. Thus, the fishermen who traditionally
diversity and fishery resources (Agostinho et al., 2007a). This is a fished in the river are included among the unemployed people
common-sense explanation used in legislative initiatives to make who worked in the construction of the dam and the farmers who
stocking mandatory in the entire country, recently approved in Law lost part of their land to the impoundment and who need a com-
Project 5989-09 (Lima et al., 2012; Pelicice et al., 2014). Ideally, plementary source of income for subsistence. With the natural
stocking strategies should consider (i) the need for stocking, based decrease in the harvest after the “trophic upsurge” period, fish
on detailed information about the environment, the target species, stocks do not support the fishery pressure, causing poverty in
and the intensity of the exploration; (ii) an understanding of the the area. This type of fishery is not characterized by initial plan-
processes that drove the wild stock to depletion; (iii) the establish- ning, and control becomes virtually impossible due to the high
ment of clear and quantifiable objectives; (iv) the capacity to rear demand for a scarce resource. Overfishing is constant and acts syn-
and distribute fries with a genetic quality equivalent to the wild ergistically with other disturbances such as those resulting from

Please cite this article in press as: Agostinho, A.A., et al., Fish assemblages in Neotropical reservoirs: Colonization patterns, impacts and
management. Fish. Res. (2015), http://dx.doi.org/10.1016/j.fishres.2015.04.006
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impoundment (e.g. the reduction in large-sized migratory species), Acknowledgments

leading to a severe depletion of stocks (Agostinho et al., 2007a).
Brazilian fishery legislation imposes temporal restrictions (periods The authors are grateful to the power companies (Itaipu, Copel,
of reproductive migration), spatial restrictions (places where fish Furnas Centrais Elétricas, Eletrobrás, and CESP) that financed sev-
stocks are more vulnerable to capture or nurseries), restrictions on eral projects and Nupelia for the infrastructure that enabled the
fishing gear and methods (gear with low selectivity or that captures development of the projects. The authors also thank CNPq for the
many fish), and restrictions on the size of the landed fish (capture fellowships awarded to JCGO and NCLS. AAA, LCG and FMP are “Bol-
of juveniles). In addition, entrance to the fishery requires a license, sista de Produtividade” of CNPq. Jaime Luiz Lopes Pereira made all
which should theoretically control access. However, enforcement the figures.
efforts are minimal, and access to the fishery is facilitated by fed-
eral laws. Furthermore, there is no monitoring of the effectiveness
of legal restrictions. It is clear, then, that the rigor of the law is not References
sufficient, except for a certain reduction in fishing effort during the
AES Tiête, 2007. Programa de manejo e conservação de bacias hidrográficas e reser-
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Please cite this article in press as: Agostinho, A.A., et al., Fish assemblages in Neotropical reservoirs: Colonization patterns, impacts and
management. Fish. Res. (2015), http://dx.doi.org/10.1016/j.fishres.2015.04.006
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Please cite this article in press as: Agostinho, A.A., et al., Fish assemblages in Neotropical reservoirs: Colonization patterns, impacts and
management. Fish. Res. (2015), http://dx.doi.org/10.1016/j.fishres.2015.04.006