Content uploaded by Silvia López-Casas
Author content
All content in this area was uploaded by Silvia López-Casas on Jul 13, 2016
Content may be subject to copyright.
Journal of Fish Biology (2016) 89, 65–101
doi:10.1111/jfb.13018, available online at wileyonlinelibrary.com
Freshwater sh faunas, habitats and conservation
challenges in the Caribbean river basins of north-western
South America
L. F. JIMÉNEZ-SEGURA*†, G. GALVIS-VERGARA‡, P. CALA-CALA‡,
C. A. GARCÍA-ALZATE§, S. LÓPEZ-CASAS*, M. I. RÍOS-PULGARÍN‖,
G. A. ARANGO¶, N. J. MANCERA-RODRÍGUEZ**,
F. GUTIÉRREZ-BONILLA†† R. ÁLVAREZ-LEÓN ‡‡
*Universidad de Antioquia, Calle 67 No. 53-108, Medellín, Colombia, ‡Universidad Nacional
de Colombia, Carrera 25 No. 61-20, A-201, Bogotá D.C., Colombia, §Universidad del
Atlántico, Km 7 Antigua vía Puerto Colombia, Barranquilla, Colombia, ‖Universidad
Católica de Oriente, Carrera 46 No. 40b 50, Rionegro, Colombia, ¶Empresas Públicas de
Medellín, Carrera 58 42-125, Medellín, Colombia, **Universidad Nacional de Colombia-Sede
Medellín, Calle 59A No. 63-20, Medellín, Colombia, ††Universidad de Bogotá Jorge Tadeo
Lozano, Carrera 4aNo 22-61, Bogotá, Colombia and ‡‡Fundación Verdes Horizontes,
Carrera 25 No. 61-20, A-201, Manizales, Colombia
The remarkable sh diversity in the Caribbean rivers of north-western South America evolved under
the inuences of the dramatic environmental changes of neogene northern South America, including
the Quechua Orogeny and Pleistocene climate oscillations. Although this region is not the richest
in South America, endemism is very high. Fish assemblage structure is unique to each of the four
aquatic systems identied (rivers, streams, oodplain lakes and reservoirs) and community dynamics
are highly synchronized with the mono-modal or bi-modal ooding pulse of the rainy seasons. The
highly seasonal multispecies shery is based on migratory species. Freshwater sh conservation is a
challenge for Colombian environmental institutions because the Caribbean trans-Andean basins are the
focus of the economic development of Colombian society, so management measures must be directed
to protect aquatic habitat and their connectivity. These two management strategies are the only way
for helping sh species conservation and sustainable sheries.
© 2016 The Fisheries Society of the British Isles
Key words: Caribbean trans-Andean Rivers; sheries; freshwater shes; threats to conservation.
INTRODUCTION
The river basins of northern South America vary widely in their climatic and hydro-
logical conditions, and their geomorphology and soil matrix reveals a dramatic history
of land transformations millions of years ago. Some efforts have already been made to
group them into geographic units using landscape, vegetation type, climate, precipita-
tion, altitude and faunal composition (Hernández-Camacho, 1992; IGAC, 2003).
In South America, Colombia is second, after Brazil, in number of sh species
(Maldonado-Ocampo et al., 2008); however, if the species per unit area of the two
†Author to whom correspondence should be addressed: Tel.: +57 4 2195620; email: luz.jimenez@udea.edu.co
65
© 2016 The Fisheries Society of the British Isles
66 L. F. JIMÉNEZ-SEGURA ET AL.
countries is considered, Colombia is the most diverse. This remarkable sh diversity
is the result of a dynamic geological history caused by plate tectonics, the uplift
of the Andean mountains, marine incursions during the Pleistocene period and the
land-bridge connections with the North America fauna after the rise of the Panama
Isthmus. The long isolated trans-Andean sh fauna is highly endemic, with ancestral
links with the Amazon and Orinoco ichthyofaunas (Rodríguez-Olarte et al., 2011).
Since its separation from the Gondwana continent c. 100 million years ago, aquatic
systems of northern South America evolved from a proto Amazon-Orinoco river that
drained eastward to the western Atlantic to a set of basins that drains to the Caribbean,
because of the rise of various branches of the western, central and eastern Andes
cordilleras, and the Venezuela and Merida branches. Glacial and interglacial periods
of the Pleistocene, caused changes in the sea level and the advance and withdrawal
of salt water into the lowland freshwater systems. Elevation of sea level during the
warmer periods may have generated massive extinctions of freshwater biota and also
the connection of the oodplains of all of these northern basins leading to the potential
dispersion of freshwater faunas along the coast (Rodríguez-Olarte et al., 2011).Today,
the interaction between the Andean mountains and the movement of water vapour
inside the inter-tropical convergence zone (ITCZ), their altitudinal gradients and soil
types caused those north-western basins to have different characteristics of rainfall,
hydrology and water production. Additionally, land use by humans has become
a permanent inuence on freshwater habitats and it has become a threat to biota
conservation. A new extinction period, the Anthropozoic period already began.
Since the 1990s, scientic knowledge about the Colombian sh fauna has
been growing. Checklists have recently been published (Mojica et al., 2006a,b;
Maldonado-Ocampo et al., 2008; Álvarez-León et al., 2013), biogeographic anal-
yses (Rodríguez-Olarte et al., 2011), sheries and biology (Lasso et al., 2011a,
b; Usma-Oviedo et al., 2013), threats to its conservation (Galvis & Mojica, 2007;
Barletta et al., 2010; Baptiste et al., 2010; Mojica et al., 2012) and many other short
publications focused on biology and ecology are compiled in Maldonado-Ocampo
et al. (2005), Maldonado-Ocampo et al. (2013), Usma-Oviedo et al. (2013) and
Álvarez-León et al. (2013), and on the use of cytogenetic and molecular techniques
for the assessment of genetic variability are compiled in Mancera-Rodríguez et al.
(2013).
Here, most of the available knowledge on the sh fauna in some of the river basins of
north-western South America in Colombia draining to the Caribbean sea are reviewed,
to describe species richness, identify life-history patterns, habitat assemblages and the
threats to shes resulting from Colombia’s economic development, and to determine
what is needed to protect the freshwater shes and the associated artisanal sheries.
THE CARIBBEAN RIVER BASINS OF NORTH-WESTERN SOUTH
AMERICA IN COLOMBIA
The Republic of Colombia is located in the north-western corner of South Amer-
ica. It includes a continental area of 1 141 748 km2. Within the Colombian continental
area ve main river basins are recognized: Amazon, Orinoco, Pacic, Catatumbo and
Caribbean. The unit of analyses has followed the basin category and is focused on
the four principal basins north-west of the eastern branches of the Andean mountains:
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 67
12
10
8
6
4
2
807876 74 72
a
b
c
d
e
f
Caribbean Sea
Pacic Ocean
N
F. 1. Map of Colombia and the Caribbean draining trans-Andean River basins. a, Ranchería River; b, Mompo-
sine Depression; c, Magdalena River; d, Cauca River; e, Sinú River; f, Atrato River
Atrato River, Magdalena-Cauca River, Sinú River and Ranchería River (Fig. 1). Table I
shows the characteristics of these river basins.
The Atrato River (5∘28′N; 76∘00′W) ows to the north, between the western
cordillera and the Baudó range, and ends in the Caribbean Sea. It is the main river
in the Chocó biogeographic area, owing to the north. Most of the channel is naviga-
ble (500 km; 67% length) and it is one of the largest producers of water in the world.
The mean annual rainfall is 8 000 mm, and air temperatures range between 23 and 30∘
C. The Atrato River basin has a mono-modal cycle with a rainy season between June
and November, and a dry season from December to May.
T I. Characteristics of some Caribbean basin rivers. Data were taken from Lasso et al.
(2011a,b) and Mojica et al. (2006a,b)
River basin
Characteristic Magdalena Cauca Atrato Sinú Ranchería
Basin area (km2) 262 075 55 599 37 810 18 478 2 338
Maximum altitude (m) 3 685 3 000 3 800 2 300 5 572
Main channel lenght 1 502 1 350 720 336 248
Mean discharge (m3s−1) 7 100 2 407 3 993 486 7·8
Hydrologic pattern Bi-modal Bi-modal Mono-modal Mono-modal Mono-modal
Flooding plane area (Ha) 295 756 950·2 65 000 24 340 510
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
68 L. F. JIMÉNEZ-SEGURA ET AL.
The Sinú River (8∘04′N; 75∘53′W) ows to the north-west between the San Jerón-
imo and Abibe ranges, towards to the Caribbean Sea. Half of its course is navigable.
The range of annual rainfall is between 800 and 3000 mm, and air temperatures range
between 18 and 28∘C. The Sinú River discharge is between 60 and 700m3s−1and it
has a mono-modal cycle with a rainy season between June and November and a dry
season from December to May.
The Magdalena-Cauca Basin is the most populated area in Colombia, most (80%)
of the Colombian population inhabits there. The Magdalena River (1∘32′N; 76∘52′
W) ows to the north-west through an extensive valley between the eastern and central
cordilleras of the Andes. Below 200 m altitude the valley opens to form an extensive
oodplain and the main channel ows through an extensive oodplain (22 000 km2),
the Momposine depression (9∘06′N; 74∘25′W); the Cauca and San Jorge Rivers also
discharge into this depression. The Cauca River runs parallel to the Magdalena River
(1∘13′N; 77∘22′W). Between 1500 and 900 m altitude the river valley widens to form
another great oodplain area. After the 900 m altitude, the valley gets narrower and the
river ows north-east through a deep and narrow canyon until the 400 m altitude; at that
point the Cauca valley opens again to form an extensive oodplain area that nally
discharges into the Momposine depression. Beyond this area the Magdalena-Cauca
River ows to the north-west and discharges into the Caribbean Sea (11∘03′N; 74∘
50′W).
The Ranchería River is a small coastal river formed in the Sierra Nevada de Santa
Marta Mountains (SNSM) (11∘31′N; 72∘54′W). It ows to the north-east and when
it reaches 99 m altitude (11∘11′N; 72∘34′W), the channel turns to the west and
discharges in the Caribbean Sea. The SNSM is a mountainous massif, isolated from
the Andes and Perijá Cordilleras and its lower areas to the north-west form the beaches
of the Caribbean Sea. It is located in the north of Colombia (10∘01′N; 72∘36′W)
and the mean annual temperature ranges between 10 and 32∘C. SNSM forms three
drainage areas: Caribbean, west, and east. The Caribbean basin is formed by the rivers
that ow directly to the sea, covers the north face of the mountain massif and includes
18 rivers, including the Ranchería River. The western basin is formed by six rivers that
drain the western slope and discharge into a brackish water – riverine lake connected
with the Magdalena: Ciénaga Grande de Santa Marta. The eastern basin is formed by
tributaries of the Cesar River: the Badillo, Guatapurí, Cesarito, Los Clavos, Diluvio
and Ariguaní Rivers.
FISH DATA SOURCES
Lists of sh species, data for assemblage analyses, information about sheries and
habitat threats were obtained from different sources. The species list was constructed
with published data (Mojica, 2002; Maldonado-Ocampo et al., 2005, 2008, 2013;
Mojica et al., 2006a,b; Ortega-Lara et al., 2006; Villa-Navarro et al., 2006), Natural
History Museum data les (Universidad de Antioquia, Universidad Católica de Ori-
ente, Universidad del Atlántico) and unpublished data from C. García-Alzate (Cesar
River and lower basin of the Magdalena River). Synonyms and actual classication of
sh groups was veried in the online Fish Catalog (Eschmeyer & Fong, 2016).
Data on species assemblages were only obtained for some aquatic systems (creeks
and rivers) in the western and eastern slopes of the Central Andean Cordillera and
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 69
the oodplain lakes from the Magdalena River valley between the eastern and central
Cordillera. Data were obtained from three rivers: the Manso and La Miel on the eastern
slope (Magdalena Basin) and the Porce on the western slope (Cauca Basin). La Miel
and Porce Rivers were dammed for hydro-power as well as 15 streams that discharge to
reservoirs Porce II and Porce III (Porce River) and 35 oodplain lakes in the Magdalena
River Valley. Samples were taken according to the climatic season (rains or dry) in
each aquatic system and for at least a 1 year period. Fishes in the oodplain lakes
were captured only during the high water season of the sampled years (2008, 2010
and 2012) because in this period most of the sh species inhabit this aquatic system.
These data were published by Granado-Lorencio et al. (2012a,b), Hernández-Serna
et al. (2014), Jiménez-Segura et al. (2014b) and J. Álvarez-Bustamante (unpublished
data). Unpublished data of the Manso River comes from the monitoring programmes
of ISAGEN S.A. E.S.P., who kindly donated these data.
For analysing sh assemblage data, abundance of each sh species in each aquatic
system was normalized. Frequency of each sh species in each aquatic system was cal-
culated as the number of times that the species appears in the total of samples. Relative
abundance (AR) was multiplied by the sh species frequency for ranking the species
importance in the sh assemblage.
Information about sheries yield of Colombian basins was taken from public reports
of Government Agencies (AUNAP, 2014a,b). Although highly criticized because of
changes to their data recording methods, yield reports were made yearly by these Agen-
cies from 1970 until June 2014. The oldest records came from the Magdalena-Cauca
River Basin and since 2006 records have been taken in all continental basins and marine
systems (Pacic Ocean and Caribbean Sea); sadly, records from 2014 were not avail-
able when we get the data. So for comparing between basins only those years with com-
plete data were used. Reports from Lasso et al. (2011a,b) were also used. Knowledge
about habitat threats to sh fauna were compiled from different sources (unpubl. data).
CURRENT STATUS OF THE FISH KNOWLEDGE IN THE CARIBBEAN
NORTH-WESTERN BASINS IN COLOMBIA
PATTERNS OF THE FISH ASSEMBLAGES
Species richness and composition
As research advances the number of freshwater sh species in Colombia increases.
Cala (1987, 2001a,b) mentioned c. 2000–3 000 possible species, Mojica (2002)
838 species, Maldonado-Ocampo et al. (2008) counted 1435 species, and recently
Álvarez-León et al. (2013) found a little more than 1700. For the north-western basins
of the Caribbean Sea, 290 freshwater sh species are listed (Table S1, Supporting
Information). Species are grouped in seven orders and 37 families. The super-
order Ostariophysi dominates with 265 species, with the two most speciose Orders:
Characiformes (134 species; 44% of total) and Siluriformes (117 species, 37%).
Other Orders are Cyprinodontiformes (15 species), Perciformes (eight species) and
Gymnotiformes (13 species). Families with the most species are the Characidae (77
species), Loricariidae (34 species), Trichomycteridae (26 species), Astroblepidae (23
species) and Heptapteridae (11 species). Another eight families are represented just by
one species.
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
70 L. F. JIMÉNEZ-SEGURA ET AL.
0·1 0·2 0·3 0·4 0·5 0·6 0·7 0·8
Jaccard similarity index
Atrato river
(118)
Magdalena-Cauca rive
r
(224)
Sinu river
(55)
Rancheria river
(50)
0·26
0·22
0·32
Conphenetic correlation= 0·80
F. 2. Similarities in the sh faunas between the Caribbean draining trans-Andean rivers basins. Numbers in
parenthesis are the number of sh species.
Biogeographic regions
Based on sh distributions, Cala (1987) proposed eight biogeographic regions:
Andean altiplano, Atrato River system, Sinú River system, north-western Caribbean
slope and Magdalena River system, Catatumbo River, Orinoco River and Ama-
zonas River. After the analyses of Abell et al. (2008) on freshwater sh distribution
around the world, the Atrato River was joined to North Andean Pacic slopes,
the Magdalena River with the Sinú River and South America Caribbean drainages
were attached to the Trinidad region. Recently, Albert et al. (2011) and Albert &
Reis (2011) further advanced beyond Abell et al. (2008) classication to dene sh
endemism and species densities for 44 South American eco-regions. They found
that although eco-regions in the north of South America are not as diverse as oth-
ers in the Amazonas or Orinoco regions, their sh fauna is highly endemic (50%
endemism).
The basin classication based on the present analysis of species presence or
absence and has an important co-phenetic correlation value (r=0·80) so, the records
used for the construction of the UPGMA dendrogram were an adequate t. The
relationship between the sh faunas showed two large clusters with low similari-
ties: Magdalena-Cauca and Atrato River basins, and Sinú-Ranchería River basins
(Fig. 2). Twenty two species are widely distributed in the analysed basins. The
Magdalena-Cauca and Atrato basins share 75 species and the Ranchería and Sinú
Rivers share 28 species (Table S1).
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 71
120
Fish species
100
80
60
40
20
0
5–100
101–200
201–300
301–400
401–500
501–600
601–700
701–800
801–900
901–1000
1001–1100
1101–1200
1201–1300
1301–1400
1401–1500
1501–1600
1601–1700
1701–1800
1801–1900
1901–2000
2001–2100
2101–2200
2201–2200
>2300
Altitudinal range (m)
Hoplosternum
Prochilodus
Triportheus spp. GymnotiformesChaetostoma spp.
Astyanax spp. Pimelodus spp.
Astroblepus spp.
Trichomycteridae spp.
Grundulus
F. 3. Fish species number across the altitudinal range and endemism percentages in the Magdalena-Cauca River
basin, and some of the key species along the altitudinal range. , endemism percentage; , number of sh
species. Modied from Jiménez-Segura et al. (2014a).
Altitudinal zonation
The number of sh species in the basin of the Magdalena-Cauca River changes along
the altitudinal gradient showing the highest value at lower elevations (Jiménez-Segura
et al. 2014a; Carvajal-Quintero et al., 2015). Even though the number of species is
reduced as the altitude increases, the ß diversity value is high (Carvajal-Quintero et al.,
2015) and the percentage of endemic species is the highest of the analysed basins
(Fig. 3).
Altitudinal distribution of the sh fauna along the river beds in the Andes is
inuenced by the slope, speed and water temperature (Jaramillo-Villa et al., 2010;
Carvajal-Quintero et al., 2015) and the height of the water column; these physical
characteristics of the aquatic systems result in the formation of an altitudinal zonation
of sh assemblages. Aquatic ecosystems of the Magdalena River between 5 and 100m
altitude (i.e. swamps or oodplain lakes) are habitat for 62 species; between 100 and
300 m the number of species reaches 92 and assemblages up to 2300 m may have
between three and six species. Dominant species in the assemblages between 5 and
100 m altitude are Cyphocharax magdalenae (Steindachner 1878), Pimelodus blochii
Valenciennes 1840, Triportheus magdalenae (Steindachner 1878), Hyphessobrycon
proteus Eigenmann 1913, Prochilodus magdalenae Steindachner 1879, Hoplosternum
magdalenae Eigenmann 1913 and Astyanax magdalenae Eigenmann & Henn 1916.
As altitude increases, the Andean rivers assemblage changes and other species
become abundant, such as Chaetostoma spp.,Andinoacara latifrons (Steindachner
1878), Astyanax spp., Bryconamericus spp. and Creagrutus magdalenae Eigenmann
1913. Between 200 and 700 m, Brycon henni Eigenmann 1913 and the genera
Astroblepus,Trichomycterus and Lebiasina characterize the sh fauna.
Although the number of species in the assemblage is negatively correlated with
altitude, this pattern is disrupted over some altitude ranges. Stretches of the channel
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
72 L. F. JIMÉNEZ-SEGURA ET AL.
between 100 and 300 m altitude are the richest in species. These stretches are highly
heterogeneous and different aquatic environments can be found: small streams, creeks,
rivers with wide and deep channels with an extensive lateral oodplain, and large
lowland lakes (Jaramillo et al. 2015a) that are repeatedly ooded and connected
during the two seasons of yearly oods. Although the variety of aquatic environments
in these stretches may be the cause of most species richness, this is a hypothesis that
needs to be veried.
As the altitude increases, the water temperature decreases and water speed increases,
resulting from the increased slope of the channel (Lewis, 2008). These conditions
cause changes in the composition of the sh species assemblage: small species that
have developed body structures and depressed body forms that allow them to live
in such rife-pool, shallow environments are more frequent. In the aquatic habitats
in the altitudinal range between 1000 and 1800 m, there are between 20 and 30
species. Characins like B. henni and species of the genus Creagrutus,Astroblepus
and Trichomycterus are typical. Assemblages over 1800 m are poor in species with
Eremophilus mutissi Humboldt 1805, Lebiasina narinensis Ardila Rodriguez 2002 and
Grundulus species found most frequently. The sucking mouth disc and strong pectoral
and pelvic ns of the Astroblepidae and Loricariidae families are the most obvious
characteristics associated with the type of aquatic environment that these bottom
species inhabit (Maldonado-Ocampo et al., 2005; Carvajal-Quintero et al., 2015).
Aquatic habitats and their sh assemblages
The sh assemblages in the Magdalena-Cauca Basin are unique to each aquatic sys-
tem in the basin. Each aquatic system has a different number of sh species; reservoir
systems were the poorest and the river systems in the low lands were the richest in
species number. In every aquatic system, 70% of the relative abundance is represented
by at least 13 species, all different in each of the aquatic systems (Table II). Inter-
estingly, B. henni was present in all the systems of the western slope of the Central
Cordillera, although its abundance and frequency was not high in the reservoirs or in
the eastern drainages of this basin.
The following sh species were the most representative of the sh fauna in the
Andean streams: B. henni, Guacuco catshes of the genus Astroblepus,A. latifrons
armoured catshes Chaetostoma spp. and the tetra Bryconamericus caucanus Eigen-
mann 1913 comprised most of the total abundance. Habitat characteristics of these
Andean streams are very distinctive. The longitudinal slope of the channel is almost
30% so it forms a serial of cascades, rifes and pools than offer unique conditions for
each sh species. The streams are surrounded by at least a gallery forest fringe that
is the food source for most of the species that inhabit the streams because it offers
seeds, insects and other invertebrates. Also, the rocky substratum favours the growth
of microorganisms in the biolm, one of the most important foods for detritivore
sh species. As previously discussed, Astroblepus and Chaetostoma species also
have body structures that let them attach to the rocky substratum during the ushing
discharges of rainy seasons and during dry seasons, individuals of Trychomycteridae
spp. and Lebiasina spp. dig into the humid bedrock layer awaiting the arrival of the
rains (López-Casas S., pers. observ.); research on these sh strategy is needed.
In the reservoirs on the Porce River, some of the sh species of the original river
assemblage have persisted. Just one sh species, Astyanax microlepis Eigenmann 1913,
comprised 78% of total abundance. Other native species such as H. magdalenae and A.
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 73
T II. Dominant species in each aquatic system in the two slopes of the Andean Central Cordillera. Species in bold form 70% of relative numerical
abundance (AR) and frequency (F)
Central Andes Cordillera
Western Slope: Cauca River Basin Eastern Slope: Magdalena River Basin
Rank
(ARF−1)
Andean Creeks
(550– 900 m)
Dammed River Channel
(650– 900 m) Reservoirs (590 –900 m)
River channel
(200– 700 m)
Dammed River Channel
(145– 200 m)
Floodplain lakes
(70– 215 m)
1Brycon henni Andinoacara latifrons Astyanax microlepis Chaetostoma cf. leucomelas Creagrutus brevipinnis Cyphocharax magdalenae
2Astroblepus unifasciatus Poecilia caucana Hoplosternum magdalenae Creagrutus brevipinnis Gephyrocharax melanocheir Caquetaia kraussii
3Astroblepus trifasciatus Brycon henni Andinoacara latifrons Hemibrycon spp. Crossoloricaria variegata Roeboides dayi
4Andinoacara latifrons Poecilia reticulata Roeboides dayi Prochilodus magdalenae Astyanax liferus Centrochir crocodili
5Chaetostoma cf. leucomelas Astyanax microlepis Coptodon rendalli * Astroblepus guentheri Astyanax fasciatus Hoplosternum magdalenae
6Bryconamericus caucanus Bryconamericus caucanus Brycon henni Astyanax fasciatus Pseudopimelodus bufonius Ctenolucius hujeta
7Chaetostoma cf. thomsoni Chaetostoma cf. leucomelas Oreochromis niloticus * Argopleura magdalenensis Chaetostoma cf. leucomelas Leporinus muyscorum
8Astroblepus chotae Roeboides dayi Poecilia caucana Trichomycterus striatus Dasyloricaria lamentosa Ageneiosus pardalis
9Hemibrycon dentatus Bryconamericus caucanus Hyphessobrycon spp. Geophagus steindachneri Chaetostoma cf. milesi Dasyloricaria lamentosa
10 Poecilia reticulata Lasiancistrus caucanus Parachromis loisellei * Chaetostoma cf. milesi Chaetostoma cf. thomsoni Hypostomus tenuicauda
11 Astroblepus chapmani Chaetostoma cf. thomsoni Oreochromis mossambicus * Chaetostoma cf. scheri Microgenys minuta Gilbertolus alatus
12 Astroblepus grixalvii Hemibrycon dentatus Leporellus vittatus Saccodon dariensis Saccoderma hastatus Eigenmannia humboldtii
13 Astroblepus micrescens Coptodon rendalli * Chaetostoma cf. scheri Leporinus muyscorum Geophagus steindachneri Pimelodus blochii
14 Oreochromis mossambicus * Parodon magdalenensis Poecilia reticulata Characidium phoxocephalum Astyanax magdalenae Curimata mivartii
15 Hemibrycon boquiae Oreochromis mossambicus * Lasiancistrus caucanus Chaetostoma spp. Prochilodus magdalenae
16 Oreochromis niloticus * Hoplosternum magdalenae Trichomycterus banneaui Pimelodella chagresi Astyanax fasciatus
17 Trichomycterus chapmani Hemibrycon boquiae Astyanax magdalenae Ctenoluccius hujeta Astyanax magdalenae
18 Astroblepus frenatus Xiphophorus hellerii * Parodon magdalenensis Brycon fowleri Andinoacara latifrons
19 Poecilia caucana Chaetostoma cf. scheri Trichomycterus caliense Creagrutus magdalenae Hypostomus hondae
20 Xiphophorus hellerii * Oreochromis niloticus * Chaetostoma cf. thomsoni Lasciancistrus caucanus Sternopygus aequilabiatus
Species
number
40 38 14 68 91 53
*, an exotic species.
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
74 L. F. JIMÉNEZ-SEGURA ET AL.
latifrons, are common and abundant. These species are typical of oodplain lakes of the
Magdalena River (Granado-Lorencio et al., 2012a) so the reservoir habitat conditions
may favour their recruitment. Foreign species such as Coptodon rendalli (Boulenger
1897), Parachromis loisellei (Bussing 1989) and Oreochromis spp. were also important
in the reservoir assemblage; they have also been reported in many other reservoirs in
South America (Agostinho et al., 2007). These species are good colonizers of these
articial habitats because they make their nests in the sandy substratum of the reservoir;
they have parental care of their offspring and are mostly omnivores.
The riverine sh assemblage was more diverse than streams or reservoirs. In La
Miel River, downstream of the dam, nine species were the most important in the
sh assemblage: Tetras: Creagrutus brevipinnis Eigenmann 1913, Gephyrocharax
melanocheir Eigenmann 1912, Astyanax liferus (Eigenmann 1913) and Astyanax
fasciatus (Cuvier 1819) and three species of detritivores [Chaetostoma spp., Dasy-
loricaria lamentosa (Steindachner 1878), Crossoloricaria variegate (Steindachner
1879)] and the migratory Prochilodontidae: Bocachico Prochilodus magdalenae
Stenidachner 1879 and the Jetudo Ichthyoelephas longirostris (Steindachner 1879).
This assemblage is seasonally enriched with migratory sh species [Pimelodi-
dae: Pseudoplatystoma magdaleniatum Buitrago-Suarez & Burr 2007, Sorubim
cuspicaudus Littmann, Burr & Nass 2000, Pimelodus spp. and the characiforms
T. magdalenae,Curimata mivartii Steindachner 1878, Cyphocharax magdalenae
(Steindachner 1878)] that move upstream through the Magdalena River from the
oodplain in the lowlands during the dry season. Although the ow in La Miel
River downstream of the dam is inuenced by the water discharge of hydropower
generation, hydro-peaking is damped by the natural ow of its tributaries: the Manso
and Samana Rivers (Jiménez-Segura et al., 2014b; López-Casas 2015). Also, there
are several habitats such as sandy beaches, rocky substratum, pools and rifes that
offer favourable conditions for different sh species. On the other hand, the dammed
channel of the Porce River is species poor if compared with the Manso and La
Miel Rivers and four species are characteristic of the sh assemblage. This is the
result of the isolation of this sector of the river, because the sampling only took
place in the stretch of the Porce River between the discharge of Porce II reservoir
and the main body of the Porce III reservoir. In this sector, due to the lack of sh
pass facilities, there is no possibility of sh species reaching the Porce III reservoir
from downstream, where the ow is free. In this isolated stretch of the river the
substratum is rocky and the ow is mainly inuenced by discharge of hydropower
ows and species typical of riverine systems are found [i.e.B. henni,Chaetostoma
spp., Poecilia reticulata Peters 1859 and Poecilia caucana (Steindachner 1880)] and
from the reservoir (A. microlepis,A. latifrons,C. rendalli and Oreochromis spp.)
(J. Álvarez-Bustamante, unpubl. data).
The oodplain lakes in the northern rivers of South America include almost 5092
(Jaramillo et al. 2015) water bodies with a total surface area of 5332 km2(Ricaurte L.,
unpubl. data) whose limits are lost when the entire oodplain is inundated during high
water. Rains, connectivity with the main river channel, shape (perimeter) and area are
all denitive factors that inuence the sh assemblage and the sheries production. Fish
assemblages are dominated by characiforms and siluriforms. The Characidae, Curi-
matidae, Prochilodontidae and Pimelodidae are the most diverse families. Although
these habitats are not the most diverse (nspecies =53) if compared with the La Miel River
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 75
downstream of the dam (nspecies =91), 13 species are characteristic of these aquatic sys-
tems and the importance of iliophagous species as C. magdalenae and P. magdalenae is
remarkable. Other species are also important in the riverine-lake assemblage: the Yel-
low Mojarra Caquetaia kraussii (Steindachner 1878) and migratory sh species such as
Leporinus muyscorum Steindachner 1900, C. mivartii and P. blochii. These oodplain
lakes are an important component of the migratory cycle of some sh species and play
an important role in buffering the highest river discharges in the Magdalena River basin.
These aquatic systems have received much of the scientic research attention
because they are important for sheries sustainability in the Magdalena-Cauca Basin
(Arango-Rojas et al., 2008; Jaramillo-Villa & Jiménez-Segura, 2008; Ríos-Pulgarín
et al., 2008; Jiménez-Segura et al., 2010b; Granado-Lorencio et al., 2012a,b). Cli-
matic seasonality determines the characteristics of the lakes and the behaviour of
the sh fauna is also synchronized with it. Changes in depth, transparency, nutrients,
dissolved oxygen and water temperature are important for habitat and food availability
to shes during the rainy seasons. The sh responses to the change are described
in Jiménez-Segura et al. (2010a). In this sh assemblage there are different groups
of life-history strategies: migrant and non-migrant shes (white and black shes;
Welcomme, 1985). The rst group of shes move between the lake and the river as the
water levels vary and the second group remains in the lakes regardless of this change.
During the dry season, non-migrant sh [i.e. A. latifrons,Synbranchus marmoratus
Bloch 1795, Trachelyopterus insignis (Steindachner 1878) and H. magdalenae] stay
in the lakes. Although little is known about the strategies of these particular species
in these river basins, it is assumed that they may develop strategies similar to those
of their relatives in other basins of South America for breathing air to obtain oxygen,
and can nd food in available habitats in order to survive (Welcomme, 1985; Val
and Randall, 2005). As the rainy season begins, non-migrant shes spawn in the
lake and the inux of river water into the lake triggers the production of plankton,
the main food resource for larvae. Also, as the water level gets high, the growth of
aquatic plants enhances the shelter available for feeding and growth of juvenile sh of
non-migrant and migrant sh species. With the decrease in the intensity of rainfall, the
oodwaters recede, water levels in the lakes drop and the biological cycle starts again
(Jiménez-Segura et al., 2010b). This cycle occurs once in Sinú, Atrato and Ranchería
Rivers (basins with mono-modal hydrological regimen), but in the Magdalena-Cauca
basin occurs twice a year (bi-modal regime).
Although more research is needed on sh biology, some of the described life
strategies conform to the environmental guilds proposed by Welcomme et al. (2006).
There are, however, some important changes in the eupotamonic, pelagophilic and
phytophilic guilds (potamonic communities) proposed, associated with the spawning
and nursery sites. These changes are described.
The sh migrations: the habitat linkage
Potamodromous migration (white sh) is a common strategy of some sh species
in the northern river basins of South America and these movements directly impact
the artisanal sheries. Almost 15% of the sh species in these basins have migratory
behaviour synchronized with the rainy season and mainly associated with spawn-
ing. Of the 15 migratory species living in the Magdalena-Cauca River; most perform
migrations of short (<100 km) or medium lengths (100– 500km) (Usma-Oviedo et al.,
2013). Some of the most famous migratory species in the Magdalena-Cauca River
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
76 L. F. JIMÉNEZ-SEGURA ET AL.
are P. magdalenae,Brycon moorei Steindachner 1878, Salminus afnis Steindachner
1880, C. magdalenae,T. magdalenae,C. mivartii,L. muyscorum,I. longirostris,
Pimelodus blochii Valenciennes 1840, Pimelodus grosskopi Steindachner 1879,
P. magdaleniatum and S. cuspicaudus. Recent estimations of migration distances
made by López-Casas (2015; submitted to Journal of Fish Biology, May 2015)
show that migrations in the Magdalena-Cauca River are from 0·8 to 1223·0 km long;
individuals of P. magdalenae may swim almost 1224 km, S. afnis from 24 to 130 km
and I. longirostris up to 198km.
Rains and water level are the main cues for migratory movements. The hydrological
cycle in the northern river basins is mainly determined by the ITCZ and its interactions
with the Andes. During the 12 months of the year, some basins (i.e. Atrato, Sinú and
Ranchería Rivers) have 6 months of heavy rain (rainy season) and 6 months of reduced
rain (dry season), but the Magdalena-Cauca Basin is bi-modal and has two rainy
(April–June and September–November) and two dry seasons (December–March and
July–August). The rainy cycle of the Magdalena-Cauca is a result of the back and
forth movements across the equator of the ITCZ throughout the year, and the position
of the three cordilleras. During the summer in the northern hemisphere (July and
August) the ITCZ moves to the north, and in December moves to the south. These
two movements of the wet winds over the slopes of the three cordilleras generate this
climatic pattern.
During the last ve decades, knowledge about migration dynamics came from oral
sher information and from sheries report data. Based on 10 years of ichthyoplankton
monitoring in the main channel of the Magdalena River and on migratory routes of
some tagged shes, different phases in the migratory process occur between four main
aquatic systems: oodplain lakes, connection channels between lakes and rivers, main
river channels and tributaries (Fig. 4).
During the dry season, most of the migratory sh (P. magdalenae,T. magdalenae,
C. mivartii and L. muyscorum) move towards the free owing channel because water
conditions in the oodplain lakes became unfavourable (i.e. low dissolved oxygen,
higher water temperatures, area and depth reduction and higher predation). In the rst
phase of the migratory cycle, shes move from the lake to the channel that connects
with the free owing channel; some shes stay in the connection channel and do not
move to the main river. The shes that get out of the lake move to the main river and
begin to move against the ow at the same time that gonad maturation starts (second
phase). As the rainy season begins and the water level rises, the third phase begins:
shes that stayed in the main river channel move to the tributaries to spawn, as do
those that stayed in the tributaries during the dry season. The fourth phase begins
when, as the ows increase, adults and embryos of sh species such as P. magdalenae,
S. cuspicaudus,P. magdaleniatum and Pimelodus spp. drift downstream along the river
channel, as the embryos develop. In the fth phase, with the maximum ows, the river
overows into the oodplain lakes carrying embryos and larvae of these migratory sh
species. Since during high water periods these lakes provide abundant shelter and food,
some of the adults of some species (P. magdalenae and I. longirostris) stay in the trib-
utaries, but most of them return to the lakes and others (mainly pimelodids) stay in the
river channels, where they remain until the next period of low water, when the cycle
starts again.
This migratory cycle associated with reproduction, larvae drift and recruitment
has been described for many tropical rivers around the world (Welcomme, 1985;
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 77
Lateral migration (Connection channels)
Upstream migration (River channel)
Adults ripening (River channel)
Spawning (River channel and tributaries)
Downstream drift and embryo incubation (River and connection channel)
Feeding and growth areas for sh larvae (Floodplain lakes)
Feeding areas for adults (lakes, river channel, tributaries)
Fishery effort
In the river channel and tributaries
in the connection channel
in the oodplain lakes
Jan
Feb
Mar
Apr
May
Jun
Jul
Aug
Sep
Oct
Nov
Dec
River discharge
F. 4. Habitat linkage in sh migration, reproduction and recruitment and sheries in the Magdalena-Cauca
River basin over the annual cycle.
Bialetzki et al., 1999; Lucas & Baras, 2001; Baumgartner et al., 2004), but in the
Magdalena-Cauca River it happens twice in the year with consequences for sh
population recruitment and artisanal sheries.
Feeding groups
Most of the sh species in the northern basins of South America are generalist feeders
and specialized feeding on limited resources is rare. Most of the evidence comes from
research on the ecology of individual sh species and there is little published data on
interaction between sh species or about trophic webs. Matthews (1998) states that
feeding strategies in shes are so diverse that it is hard to dene groups, so simple and
broad trophic groups are proposed here (herbivores, omnivores and carnivores) and
following Bowen (1983) the detritivorous group is included (mud and detritus feeders)
(Table III). In the herbivorous group, shes eat plants, seeds and algae. Carnivores eat
all invertebrate groups (i.e. insects, molluscs and worms) and shes; in this group, are
included the scale eater Roeboides dayi (Steindachner 1878) and the hematophagous
Paravandellia phaneronema (Miles 1943).
Body features of the sh species are important for foraging. Mouth position (ventral
or terminal), body form (depressed or compressed) and caudal peduncle size have been
identied as the main body features than inuence diet (K. Aguirre, unpub. data). In
the herbivorous group, most of the species belong to the family Loricariidae. These
armoured catshes are attened dorso-ventrally and have a ventral sucker-mouth disc
with soft teeth they use to scrape the biolm, thus feeding is done by ploughing along
the substratum and using the thick-lipped, toothy mouth to scrape plant material (l-
amentous algae and diatoms) from hard structures (rocks, trees, roots and boats) or
to suck up ne sediments. The detritivorous group mainly consists of prochilodontids
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
78 L. F. JIMÉNEZ-SEGURA ET AL.
T III. Fish composition in the trophic groups in each aquatic system in the Magdalena-Cauca River basin
Trophic
group Omnivore Carnivore Herbivorous Detritivorous Reference
Andean
streams
Astroblepus spp.,Astyanax spp.,
Brycon spp.,Characidium
spp.,Creagrutus brevipinnis,
Hemibrycon spp.,Saccodon
dariensis,Trichomycterus
spp.,Xiphophorus helleri,
Hyphessobrycon poecilioides,
Bryconamericus spp.
Andinoacara latifrons,
Apteronotus eschmeyeri,
Brachyhypopomus
occidentalis,Par a ch rom i s
loisellei,Trichomycterus
chapmani,Trichomycterus
retropinnis,Parodon
magdalenensis,Poecilia
spp.
Chaetostoma spp.,
Lasiancistrus caucanus
García-Alzate &
Roman-Valencia
(2008), Roman-P
et al. (2014), D.
Restrepo-Santamaria
(unpubl. data).
River
channel
Acestrocephalus anomalus,
Andinoacara latifrons,Brycon
spp.,Astyanax spp.,
Bryconamericus spp.,
Argopleura magdalenensis,
Cetopsis othonops,
Cetopsorhamdia spp.,
Characidium spp., Creagrutus
magdalenae,Cynopotamus
magdalenae,Characidium
spp., Microgenys spp.,
Parodon magdalenensis,
Trichomycterus spp.,
Pimelodus spp., Pimelodella
spp., Roeboides magdalenae,
Saccoderma spp., Saccodon
spp., Xiliphius magdalenae,
Leporinus muyscorum,
Leporellus vittatus,
Triportheus magdalenae
Apteronotus mariae,
Astroblepus homodon,
Caquetaia kraussii,
Caquetaia spp.,
Ctenolucius hujeta,
Eigenmannia virescens,
Gasteropelecus spp.,
Geophagus steindachneri,
Hoplias malabaricus,
Poecilia caucana,Poecilia
spp., Potamotrygon
magdalenae,
Pseudoplatystoma
magdaleniatum,Rivulus
spp., Salminus afnis,
Sorubim cuspicaudus,
Sternopygus aequilabiatus,
Paravendellia
phaneronema
(hematophague)
Ancistrus spp.,
Chaetostoma spp.,
Cordylancistrus spp.,
Hypostomus spp.,
Panaque spp.,
Pterygoplichthys spp.,
Rineloricaria spp.,
Sturisoma spp.,
Spatuloricaria spp.
Ichthyoelephas longirostris,
Prochilodus magdalenae,
Cyphocharax
magdalenae,Curimata
mivartii
Jiménez-Segura et al.
(2014b)
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 79
T III. Continued
Trophic
group Omnivore Carnivore Herbivorous Detritivorous Reference
Floodplain
lake
Brycon spp., Pimelodus spp.,
Hoplosternum magdalenae,
Cynopotamus magdalenae,
Centrochir crocodili,
Leporinus muyscorum,
Astyanax spp., Leporellus
vittatus,Pimelodus spp.,
Pseudopimelodus bufonius
Gilbertolus alatus,Hoplias
malabaricus,
Pseudoplatystoma
magdaleniatum,
Ageneiosus pardalis,
Triportheus magdalenae,
Trachelyopterus insignis,
Abramites eques,
Andinoacara latifrons,
Geophagus steindachneri,
Caquetaia kraussi,
Apteronotus mariae,
Eigenmannia virescens,
Ctenolucius hujeta,
Plagioscion surinamensis,
Sorubim cuspicaudus,
Roeboides dayi
(lepidophague)
Hypostomus spp.,
Crossoloricaria spp.,
Dasyloricaria
lamentosa,
Pterygoplichthys spp.,
Squaliforma tenuicauda,
Sturisoma spp.
Prochilodus magdalenae,
Cyphocharax
magdalenae,Curimata
mivartii
K. Rivera-Coley & D.
Arenas (unpubl..
data), A. Arango
(unpubl. data)
Reservoir Andinoacara latifrons,Astyanax
microlepis,Hoplosternum
magdalenae,Brycon henni
Roeboides dayi
(lepidophague)
Coptodon rendalli
(algivore)
D. Restrepo-Santamaria
(unpubl. data), Y.
Rondon (unplubl.
data)
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
80 L. F. JIMÉNEZ-SEGURA ET AL.
and curimatids. The terminal mouth of I. longirostris (Prochilodontidae) can be pro-
jected forward forming a sucker disc with soft teeth that graze on detritus and biolm
present on hard structures. The migratory P. magdalenae (Prochilodontidae) also uses
its mouth with soft teeth on their lips to scrape biolm from hard substrata in the
rivers it use when it migrates, but this species as well as C. mivartii (Curimatidae), and
C. magdalenae mainly eat mud accumulated on the bottom of the oodplain lakes or
by sucking the roots of aquatic plants (A. Bermúdez, unpubl.data).
Intraspecic morphological differences have been described as ‘trophic polymor-
phism’ being related to the structures responsible for food and which can confer advan-
tages in the use of the habitat and specic resources (Olsson & Eklöv, 2005). Roberts
(1974) has noted ve oral and dental polymorphisms for Saccodon dariensis (Meek &
Hildebrand 1913) (Parodontidae) in tributaries of the Rivers Sinú, Magdalena, upper
Cauca and Atrato, and Restrepo-Gómez & Mancera-Rodríguez (2014) suggest that the
coexistence of two of these oral polymorphisms in the Guatape River (Magdalena River
basin) may be related to a trophic polymorphism that confer advantages in different
trophic habits and differential access to the items that constitute their diet.
Although a sh species may be allotted to a trophic group, foraging and feeding
strategies are so diverse because of dynamic environmental conditions and changes in
the diel and annual cycles, that it is necessary to describe its particular feeding strat-
egy. Foraging and diet turnover of sh species during rainy seasons and in response to
the diel cycle are poorly known. It has only been reported in this region for the diet
of the cachegua Trachelyopterus insignis (Steindachner 1878). During the rainy sea-
son, this sh feeds on different sources and is an omnivore, but in the dry season, it
feeds mainly on other shes and hence is a piscivore (S. López-Casas & J. G. Ospina,
unpubl. data). Although the omnivorous group is by far the most diverse, taxa compo-
sition is strongly dependent on the species assemblage in the different aquatic systems
(Table III). In oodplain lakes Hernández-Serna et al. (2015) also found that there is a
foraging segregation during the diel cycle by body size and trophic group: small shes
forage during the day while larger shes do so at night, and carnivores and detritivores
forage during the night and omnivores by day.
The unique characteristics of the aquatic system (depth, substratum structure and
water velocity) may be denitive for trophic group species richness because of food
availability. Higher species composition turnover due to the aquatic system type is
observed in the Loricariidae family. In Andean streams, species of Chaetostoma are
the most important in the herbivore group. Their at bodies, sucker-shaped mouth and
strong pectoral ns let them thrive in this shallow and turbulent aquatic system. In
deeper and quiet oodplain lakes, other Loricariidae species such as Hypostomus spp.
and Pterygoplichthys spp. with higher and shorter bodies are successful. The absence
of substrata for biolm growth in reservoirs may be one of the reasons loricariids are
mostly absent from those habitats. More research is needed as well as further analyses
of carbon ux and food webs to advance knowledge of functional feeding groups in
the region.
Reproductive seasonality
The reproductive season is mostly determined by parental condition related to fat
reserves stored during the previous season of higher food availability, as well as
suitable habitat conditions favourable for egg fertilization and embryo development
and subsequently food availability for larvae (Munro, 1990; Lowe-McConnell 1995;
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 81
Vazzoler 1996; Wootton, 1999). In Andean aquatic systems, the hydrological pattern
and nutrient transport generated by the rainy season are denitive cues for selec-
tion of optimal reproduction timing (Jiménez-Segura, 2007; Kerguelen-Durango &
Atencio-Garcia, 2015). Most of the sh species breed during the rains, a few during
the dry seasons and others throughout the year (Table IV). This variation is mainly
inuenced by the interaction between rains and particular environmental conditions
in each type of aquatic system that offers food for adults and favours spawning
success. Thus, while in oodplains and the main river channel favourable conditions
for reproduction are much more suitable during high waters; in the Andean streams
reproduction is much more common during low waters to avoid egg and larval drift
during the rainy season. Reservoirs are the exception to this pattern, sh species
reproduce throughout the year. The main reason is that there is no drastic change in the
water level as a result of the rain patterns as happens in natural aquatic systems, so sh
species only have to nd favourable habitat conditions to spawning and recruitment.
Rainy season changes in habitat conditions and their direct inuence is greater
in some aquatic systems than in others; ow magnitude, frequency and amplitude
are important. In the rainy season, Andean streams are highly disturbed by frequent
ushing, water turbulence and sediment transport. The cumulative effect of small
streams ooding at the same time, causes the amplitude and magnitude of river ows
to increase, but their frequency diminishes. Finally, when the ow discharge exceeds
the storage capacity of the river channel, the river spills onto the lateral plain and water
enters the oodplain lakes. This physico-chemical stage generated by rains in the
aquatic systems of the Andean mountains causes favourable environmental conditions
for sh reproduction and recruitment in the oodplains.
To reproduce, adults need extra energy for gonad growth and spawning, and
favourable conditions for offspring survival (Munro, 1990). Rains and oods enhance
food for adult shes in every aquatic system (Welcomme, 1985; Junk et al., 1989;
Lowe McConnell, 1995; Jiménez-Segura et al., 2010b), but each sh needs specic
environmental conditions for spawning and maximum offspring survival. Although
rains and oods provide favourable conditions to feed adults and larvae, there is a
time lag after the rains and oods for sh spawning in the Magdalena-Cauca River
(Jiménez-Segura et al., 2010a) and food availability for sh larvae on the oodplains
(Jiménez-Segura et al., 2010b). The hypothetical relationship between spawning area
location for migratory species, optimal drift distances of the larvae related to water
velocity in the river channel and the timing of plankton blooms (food for sh larvae)
in oodplain lakes needs to be veried.
In Andean streams, oods carry nutrients from nearby soils, and rains carry insects,
leaves and seeds from the gallery forest, so extra allochthonous food enters the system,
allowing shes to store energy in adipose tissue that will be used for gonad growth.
In these aquatic systems, some sh species spawn in the dry season and others at the
beginning of the rains (Table IV). To increase embryo and larval survival, some shes
develop parental care and parents carry the eggs (i.e. loricariids) or keep them inside
crevices, submerged trees or make nests to keep them away from the inuence of cur-
rents and safe from predators. When the embryos hatch, in some species the larvae
develop a cephalic sucker to attach to rocky substrata (e.g. B. henni) or stay in pools
to avoid ow. Most of the non-migratory shes in the oodplain lakes of the Mag-
dalena River feed during the oods, their gonads mature in the dry season, they spawn
in the lake as the water level rises, and their larvae nd shelter and food under the
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
82 L. F. JIMÉNEZ-SEGURA ET AL.
T IV. Climatic periods for sh spawning in the aquatic systems in the Magdalena-Cauca River basin
Climatic period
Aquatic system Dry Rains (*) Rains and dry References
Streams Andinoacara latifrons,Characidium
caucanum,Trichomycterus
chapmani,Bryconamericus huilae,
Hemibrycon boquiae,Microgenys
minuta,Creagrutus guanes,
Roeboides dayi
Astroblepus spp., Brachyhypopomus
occidentalis,Chaetostoma spp.,
Characidium grupo zebra,Hemibrycon spp.,
Lasiancistrus spp., Brycon henni,Astyanax
fasciatus,Trichomycterus spp.,Saccodon
dariensis,Bryconamericus caucanus
Astyanax microlepis,Poecilia
caucana,Pimelodus grosskopi,
Astroblepus homodon,
Lasiancistrus caucanus,Astyanax
aurocaudatus
Cala (1997), Román-Valencia & Muñoz
(2001), Román-Valencia et al. (2003),
Román-Valencia & Ruiz (2005),
Román-Valencia & Botero (2006),
Maldonado-Ocampo et al. (2005),
Román-Valencia et al. (2008),
Briñez-Vásquez & Francis-Turner
(2006), Román-Valencia & Samudio
(2007), Torres-Mejía & Ramírez-Pinilla
(2008), Jiménez-Segura et al. (2015), N.
Mancera-Rodríguez et al. (2016), L.F.
Jiménez-Segura (unpubl. data),
River channel Bryconamericus caucanus,Caquetaia
kraussi,Caquetaia umbrifera,
Chaetostoma milesii,Crossoloricaria
variegata,Dasyloricaria lamentosa,
Hypostomus hondae,Trichomycterus
banneaui,Geophagus steindachneri
Characidium phoxocephalum,Leporinus
muyscorum,Pseudoplatystoma
magdaleniatum,Sorubim cuspicaudus,
Parodon magdalenensis,Pimelodus blochii,
Prochilodus magdalenae,Roeboides dayi,
Salminus afnis
Creagrutus brevipinnis,Ctenolucius
hujeta,Trichomycterus caliense,
Trichomycterus chapmani,
Trichomycterus striatus,Poecilia
spp.,Apteronotus magdalenensis
Maldonado-Ocampo et al. (2005),
Rangel-Serpa & Torres-Mejia (2015),
Jiménez-Segura et al. (2009),
Jiménez-Segura et al. (2014b), L.F.
Jiménez-Segura (unpubl. data),
Floodplain lakes Sturisoma panamense Abramites eques,Cyphocharax magdalenae,
Prochilodus magdalenae,Leporinus
muyscorum,Pimelodus blochii,Centrochir
crocodili,Trachelyopterus insignis,Astyanax
caucanus,Astyanax fasciatus,Astyanax
magdalenae,Roeboides dayi,Eigenmannia
virescens,Gilbertolus alatus,Triportheus
magdalenae,Plagioscion magdalenae
Caquetaia kraussi,Andinoacara
latifrons,Geophagus
steindachneri,Hoplias
malabaricus,Rhamdia quelen
Atencio-García et al. (2001),
Maldonado-Ocampo et al. (2005);
Jiménez-Segura et al. (2010b),
Olaya-Nieto et al. (2010).
Reservoirs Brycon henni,Poecilia caucana Andinoacara latifrons,Astyanax
microlepis,Coptodon rendalli,
Oreochromis spp.,Hoplosternum
magdalenae,Roeboides dayi,
Caquetaia kraussi,Caquetaia
umbrifera
Maldonado-Ocampo et al. (2005),
Solano-Peña et al. (2013), J. Londoño
(unpublished data), A. Loaiza
(Unpublished data), L.F. Jiménez-Segura
(unpubl. data),
(*) in the beginning of the rainy period
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 83
140 000
(a)
(b)
120 000
100 000
80 000
60 000
40 000
20 000
0
Yield (t)
Yield (%)
90
60
30
0
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
Year
F. 5. (a) Fisheries yield in Colombia ( , marine landings; , freshwater landings) and (b) percentage of
freshwater yield of each river basin ( , Upper Amazonas; , Atrato River; , Upper Orinoco River; ,
Magdalena-Cauca River) from 1995 to 2014.
aquatic macrophytes (Jiménez-Segura et al., 2010b). Andean reservoirs offer food for
shes throughout the year, allowing native shes that persist in the assemblage to spawn
throughout the year. So, adults of riverine species ripen during the dry season and spawn
at the beginning of the oods, species in the lowland lakes ripen in the dry season, but
spawn after the oods during the high water season, species in streams ripen during
the oods, but spawn during the dry season, and shes in reservoirs spawn any time,
throughout the year.
FISHERIES
Colombia’s shery production comes mainly from marine resources. Freshwater pro-
duction represented 20% (mean value) of the total production in the years 1995– 2014
(Fig. 5). During that time period, catches from trans-Andean rivers represented 80%
of freshwater production. The Magdalena-Cauca Basin is the most productive river for
artisanal sheries; its yield comprised between 72 and 94% of the sheries yield of
trans-Andean basins and it supplies the protein demand of the local population (67%)
and populations in major cities such as Bogotá, Medellin and Barranquilla (Escobar
et al., 2014). Yield data from the Ranchería River do not exist because the shery is
poorly developed and there is no data recording (Mojica et al., 2006a).
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
84 L. F. JIMÉNEZ-SEGURA ET AL.
T V. Fish species and shing gear ranking according their percentage to sheries yield in the Caribbean Basins (Mojica et al., 2006a,b; Gutiérrez
et al., 2011a,b; Gutiérrez, 2011; Escobar et al., 2014). Fishing gear data from AUNAP (2014a,b)
River Basin
Rank Magdalena-Cauca Sinú Atrato Ranchería
Species 1Prochilodus magdalenae Prochilodus magdalenae Prochilodus magdalenae Prochilodus reticulatus
2Pseudoplatystoma
magdaleniatum
Hoplias malabaricus Leporinus muyscorum Ichthyoelephas longirostris
3Pimelodus blochii Cyphocharax magdalenae Hoplias malabaricus Salminus afnis
4Plagioscion magdalenae Leporinus muyscorum Mugil curema
5Pimelodus grosskopi Caquetaia kraussii Centropomus spp.
Fishing gears 1 Gillnet Cast nets Gillnet NR
2 Beach seines Gill nets Hooks and lines NR
3 Cast nets Beach seines Cast nets NR
4 Trawl nets Harpoon Traps NR
5 Surrounding net Hooks and lines NR
6 Hooks and lines Surrounding net
Species number 224 55 118 50
Species in sheries 26 27 40 NR
Fisher number 43 730 3 442 1 242 NR
NR, not reported.
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 85
Fisheries in trans-Andean river basins is artisanal and multispecies (Gutiérrez et al.,
2011a). The sher population was reported as 50 600 individuals (Gutiérrez, 2011;
Gutiérrez et al., 2011a,b). Catch methods are mainly nets (gillnets, cast nets, sur-
rounding nets, trawl nets and beach seines), hooks, lines and traps. Their size, mass and
methods vary locally and depend on the characteristics of the freshwater system and on
the hydrological cycle (Table V). Boat size is highly variable, they may be 2–8 m long
(most frequently 5 m) and 0·4–1 m wide (most frequently 0·8 m), and most of them are
made of wood or breglass.
Prochilodontids are the main source for artisanal sheries in all trans-Andean basins.
Although 45 species are used by shers in the trans-Andean river basins (Lasso et al.,
2011a,b), P. magdalenae is the main target of the artisanal catch and represents
80–95% of catches (Gutiérrez, 2011; Gutiérrez et al., 2011a,b). Pimelodid species
[e.g. P. magdaleniatum,P. blochii,S. cuspicaudus and Pseudopimelodus bufonius
(Valenciennes 1840)] and Bryconidae (e.g. B. moorei,B. henna and Brycon rubricauda
Steindachner 1879) are also used, but their yield does not exceed 30% of the total.
The number of sh species used by shers changes between river basins, e.g. the
Sinú River shery is highly diverse, and shers use 45% of the total number of sh
species, but although some species may be used in all trans-Andean river basins, their
importance differs slightly from one to another (Table V).
Freshwater sheries yield in the trans-Andean Basins is highly seasonal because it is
based mainly on migratory sh species (Lasso et al., 2011b). The higher shing effort
is during the low water period because shes of migratory species move to the main
river channel from oodplain lakes for upstream migration. The magnitude of sheries
yield differs between the trans-Andean basins and there is a production pattern associ-
ated with the river water level in each basin (Fig. 6). Although there is a clear pattern
between yield and discharge, Escobar et al. (2014) describes a change in sheries yield
associated with the geomorphic characteristics of the river basins. The higher produc-
tion comes from the medium and lower sectors where the lateral oodplain is well
developed, the slope is low, oodplains lakes are numerous and their connection with
the main channels is permanent.
Riverine sheries production in the Magdalena-Cauca River fell from 60 000 t in
1975 to 10 000t in 2014 (Fig. 7). Although the observed pattern should be viewed with
caution because through the years, government agencies have changed the methodol-
ogy used to record landings, shers and environmental agencies recognize a substantial
reduction in landings. Besides the reduction in sher landings, some other worrisome
characteristics were detected: changes in the type of exploited sh species and reduc-
tion in the species size harvested by sheries, variables characteristic of overshing
described by Welcomme (1999) in other multispecies artisanal sheries.
For the last 40 years, the number and composition of species in the catch has changed.
The number of shed species has increased, in the 1970s, populations of ve species
were the most exploited but in the last decade, shers have utilized nine species. Dur-
ing the 1970s, species of high commercial value such as P. magdaleniatum,B. moorei
and S. afnis were the most important species in the sheries catch; during the 1980s,
P. magdalenae became the most important, and in the last decade new species of lower
commercial value such as L. muyscorum,C. mivartii,C. magdalenae,H. malabar-
icus and S. aequilabiatus, and some exotic species such as C. rendalli,Colossoma
macropomum (Cuvier 1816) and Piaractus brachypomus (Cuvier 1818) have become
important in the riverine sheries yield. Most of the important native species used by
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
86 L. F. JIMÉNEZ-SEGURA ET AL.
12 000 (a)
(b)
(c)
10 000
8000
6000
4000
2000
0
1000
800
600
400
200
0
14 000
12 000
10 000
8000
6000
4000
2000
0
250
200
150
100
50
0
600
500
400
300
200
100
0
Jan Feb Mar Apr MayJun
Month
JulAug Sep Oct NovDec
80
60
40
20
0
Discharge (m3 s−1)
Yield (t)
F. 6. Fisheries yield ( ) and monthly mean discharge ( ) for (a) Magdalena-Cauca River, (b) Atrato
River and (c) Sinú River. Yield data were taken from periods 1993–1999, 2006 –2009 and 2012 –2014, and
discharge data were obtained from 2000 to 2013.
artisanal shers make short or medium length migrations, have higher fecundities, per-
form pelagic spawning without parental care, but some of the recent additions to the
shery are non-migratory, build nests and have parental care. Fisheries based on such
kstrategist shes may be a threat to conservation. Also the diversity of catch methods
used by shers makes the sizes of harvested shes vary widely, and many individuals
are harvested at a size below the mean size of rst reproduction (Table VI). These bad
shing practices have also led to the capture size reduction of exploited species; mean
catch size of P. magdalenae has been reduced by 13 cm in the last 30 years (CCI, 2007).
Poverty, few possibilities for economic support and low education level of shers may
be considered as the basic causes of the improper use of the sh resource.
Hydropower development requires the formation of reservoirs, new aquatic habitats
that favour the population growth of some sh species. There are 34 reservoirs in
the trans-Andean river basins, 15 are used only for hydropower (power capacity
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 87
80 000
70 000
60 000
50 000
40 000
30 000
20 000
10 000
0
1975
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
Production (t. year–1)
Year
Pseudoplatystoma magdaleniatum
Brycon moorei
Salminus afnis
Prochilodus magdalenae
Sorubim cuspicaudus
Prochilodus magdalenae
Pseudoplatystoma magdaleniatum
Sorubim cuspicaudus
Brycon moorei
Salminus afnis
Prochilodus magdalenae
Pseudoplatystoma magdaleniatum
Pimelodus blochii
Leporinus muyscorum
Curimata mivartii
Brycon moorei
Sternopygus aequilabiatus
Salminus afnis
Coptodon rendalli
F. 7. Riverine sheries production for the last 40 years in the Magdalena-Cauca River basin and the most
important sh species in landings.
higher than 100 MW), one for crop irrigation, nine for water supply and nine are
multi-purpose (i.e. hydropower, water supply and crop irrigation). Seven sustain
important artisanal sheries (Escobar et al., 2014) that are poorly known because
landings are not monitored. Although some native shes are important in riverine
sheries production, non-native sh species are the mainstay of catches in most of
the reservoirs (Table VII). Life-history strategies of these species (i.e. low fecundity,
parental care, nest building, fast growth and low trophic levels) favours the colonization
and success of these articial aquatic systems (Gomes & Miranda, 2001).
CONSERVATION THREATS
Fifty per cent of the sh fauna of the trans-Andean Rivers has been included in
the Red List of the UICN (Mojica et al., 2012) and depletion in the riverine sh-
eries yield in the Magdalena-Cauca River basin has been observed. This situation is
the result of interacting causes, originating in the non- sustainable success of Colom-
bian society. The growth of the Colombian population, the economic development
of the country based on some legal industries (i.e. oil, mining, hydropower, exten-
sive agriculture and cattle, sh culture to protein supply and enhancement) and illegal
business (i.e. illicit crops) cause ever increasing demands for a large number of envi-
ronmental services (i.e. animal protein, wood and water). So, aquatic habitat change,
water pollution, deforestation, introduced sh species and overshing are considered
the main causes of the observed reduction in sh catches in the trans-Andean river
basins (Mancera-Rodríguez & Álvarez-León, 2005, 2006; Galvis & Mojica, 2007;
Barletta et al., 2010; Gutiérrez et al., 2010; Anderson & Maldonado-Ocampo, 2011;
Jiménez-Segura et al., 2014a). All these causes, common to all trans-Andean river
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
88 L. F. JIMÉNEZ-SEGURA ET AL.
basins, interact synergistically to modify the dynamics, connectivity and structure of
the aquatic systems, reduce their water quality and affect the natural ow pattern.
Growth of Colombian population and sher effort
Population size is now ve times higher than in 1950 so protein demand has increased
proportionately. In 2014 there were 48 813 328 Colombians (DANE, 2015) and sh
consumption per capita in Colombia was estimated between 4 and 5 kg (FAO, 2014).
Although artisanal shers increased their effort shing with non-sustainable techniques
and shing in all aquatic systems throughout the year, their landings fall far short of
satisfying the Colombian demand. Protein supply has been satised instead by the pro-
duction of sh farms in the last decade. In 2011 sh farms produced 82 733 t; sadly,
sh farm production is based mainly on exotic species as Cichlidae, C. macropomum
(Serrasalmidae), Onchorhynchus mykiss (Walbaum 1792) (Salmonidae) and, recently,
Pangasius hypophthalmus (Sauvage 1878) (Pangasiidae).
Sediment yield and deforestation
Erodible soils, slopes steeper than 40∘and the change in the forest cover are one of
the main causes of the higher sediment yield in the Magdalena-River Basin. Restrepo
& Kjerfve (2000) estimated sediment production of 560 t km−2year−1in the basin;
this is higher than numbers reported for the Amazon, Orinoco and Negro Rivers
combined. The deforestation rate in Colombia is one of the highest in tropical basins
around the world (Tucker & Townshend, 2000). Between 1990 and 2010, Colombia
lost 5·8 million ha; in 1990 forest cover was 51·6% of total Colombian land (equal to
64 417 248 ha), in 2010 it was 58 633 631 ha (González et al., 2011). Although there
has been a reduction in the last 4 years, there must be moderate optimism based on
recent data (MADS, 2014).
Because of the high rates of sediment transport in the Magdalena-Cauca River, ood-
plain lakes have been losing depth, their connectivity with the main river has been
reduced and as a result, their buffering capacity during the oods has diminished.
Rather than focusing on the recovery of forest and land use to provide a reasonable
solution to the problem of erosion and sedimentation in the lowlands, the Colombian
Government is directing its resources to specic actions with doubtful effects, e.g.
a long-term solution such as dredging the main channel of the Magdalena River to
increase its depth. Of course, this action has been promoted by the Ministry of Trans-
port to improve transport of larger ships along the river channel and, not by the Ministry
of Environment to protect oodplain lakes and their connectivity channels as the prime
habitats for freshwater fauna, migratory shes and shers.
Water pollution and eutrophication
Ten million people live in Bogotá, the capital city of Colombia. Daily, citizens and
industries discharge wastes into the Bogotá River (a tributary of the Magdalena River)
including among many sewage water, nitrogen and phosphorus-based fertilizers and
xenobiotic substances (e.g. organochlorines, cadmium and lead). Upstream the mouth
of the Bogotá River, the Magdalena River has already received residuals of pesticides
used for insect control in rice crops (e.g. DDT, Lindane, dieldrin, B-BHC, Endosulfan,
Aldrin, Dimethoate, Chlorpyrifos, Malathion and Dazinon) (Villa, 1992) the amounts
of which increase downstream due to pest control on extensive oil palm crops. Added to
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 89
T VI. Some characteristics of the sh species recruited to artisanal sheries in the Magdalena-Cauca River
Fish species LSmax mean LSLSrange LMM % Migratory strategy Mean fecundity References
Pseudoplatystoma magdaleniatum 100 58 20 – 131 80 85 M-Migrant 493 752 Jiménez Segura et al. (2008)
Sorubim cuspicaudus 80 43 6 –93 45 60 M-Migrant 78 943 Jiménez Segura et al. (2008)
Ageneiosus pardalis 44 36 35 44 S-Migrant 21 808 Perez et al. (2005)
Pimelodus grosskopi 22* 23 7– 48 20 22 M-Migrant 28 500 Cala et al. (1993)
Pimelodus blochii 35 15 7–48 18 82 M-Migrant 10 743 Ramírez et al. (2013)
Pseudopimelodus bufonius 47·9* 20 45 100 M-Migrant ?
Hoplias malabaricus 55·2 26 13– 37 25 28 non-migrant ?
Leporinus muyscorum 25·7 24 8 –40 25 52 S-Migrant 63 900 Arguello et al. (2001)
Prochilodus magdalenae 30 23 7– 55 25 67 M-Migrant 53 535 Atencio-García et al. (2013)
Salminus afnis 100 36 35 44 M-Migrant 67 500 Mojica et al. (2012)
Brycon moorei 50 35 35 17 M-Migrant ?
Plagioscion surinamensis 70 32 8– 62 30 49 M-Migrant ?
Caquetaia kraussii 26 15 11 – 24 20 97 non-migrant 1732 Solano-Peña et al. (2013)
LS, max maximum standard length (cm) reported in Froese & Pauly (2015); Mean LS, mean standard length (cm) in sher catch; LMM, mean length (cm) of rst reproduc-
tion; LSrange, standard length range (minimum–maximum); %, percentage of the sher capture under LMM . * Jiménez-Segura & Ortega-Lara (2010) and Jiménez-Segura
& Villa-Navarro (2011).
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
90 L. F. JIMÉNEZ-SEGURA ET AL.
T VII. Characteristics of some Colombian reservoirs and artisanal sheries. Data taken from Jiménez-Segura et al. (2011) and Escobar et al.
(2014). Fish species are rank by their importance in landings
Reservoir & Basin
Urra II Amani Guajaro Betania Prado Tominé Porce II Salvajina
Basin Ranking Sinú Magdalena Magdalena Magdalena Magdalena Magdalena Magdalena Cauca
Area (ha) 7 800 1 230 16 000 7 424 3 410 3 830 890 2 031
Altitude (m) 70 250 9 561 361 2 580 540 1 100
Maximum
depht (m)
73 188 91 90 20 118 148
Yiel d
(t year−1)
75·9 (2012) 11·7 (2012) 387 (2014) 493 (2008) 126 (2010) NR NR 10·9 (1996)
Fished species 1Caquetaia kraussi Oreochromis
niloticus *
Oreochromis
niloticus *
Oreochromis
niloticus *
Caquetaia kraussi Cyprinus
carpio *
Oreochromis
niloticus *
Oreochromis
niloticus *
2Andinoacara
latifrons
Caquetaia
umbrifera
Triportheus
magdalenae
Caquetaia kraussi Chyphocharax
magdalenae
Oreochromis
mykiss *
Coptodon rendalli
*
Pseudopimelodus
buffonius
3Hoplias
malabaricus
Sternopygus
aequilabiatus
Plagioscion
magdalenae
Pimelodus
grosskopf
Caquetaia
umbrifera
Eremophilus
mutissi
Oreochromis spp. *
4Panaque gibbosus Ichtyoelephas
longirostris
Prochilodus
magdalenae
Pimelodus blochii Oreochromis
niloticus *
5Trachelyopterus
badeli
Pimelodus zungaro
6Ageneiosus
pardalis
7Rhamdia spp.
Gears 1 Gillnets Gillnets Gillnets Gillnets Gillnets Hooks and lines Gillnets Gillnets
2 Hooks and lines Hooks and lines Hooks and lines Hooks and lines Hooks and lines Gillnets Hooks and lines Hooks and lines
3 Cast nets Cast nets Cast nets Cast nets
*, no native sh species; NR, not reported.
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 91
this, gold mining wastes (e.g. mercury and arsenic) and heavy metals (e.g. Cd, Pb and
Ni) are being incorporated into sh tissues throughout the food web (Alvarez et al.,
2012). Effects of these substances on freshwater shes are poorly known in Colom-
bia, but some data have been published or compiled in Cala & Sodergren (1999),
Mancera-Rodríguez & Álvarez-León (2013), Marrugo-Negrete et al. (2007, 2008a,b),
Álvarez et al. (2012), Noreña et al. (2012), Trujillo et al. (2010) or are contained in
unpublished technical papers.
Reservoirs and hydropower
Sixty-seven per cent of the energy supply in Colombia comes from hydropower
(Jiménez-Segura et al., 2014a) produced by 32 reservoirs. In the Magdalena-Cauca
River, most of these reservoirs are located in tributaries of the main channel; Urrá
I reservoir was built in the main channel of the Sinú River and in the Atrato River
there are no reservoirs to date. Reservoirs block the upstream– downstream migrations
of some sh species, change natural ow regimes (affecting the environmental cues
for migratory sh species spawning and the seasonal oods important for early stages
recruitment in the oodplain lakes), and reduce the sediment load on the oodplain.
Hence, the future is uncertain for migratory shes and remains a challenge because
government agencies plan to double hydropower production by the year 2027. Further
analyses are detailed in Jiménez-Segura et al. (2014a).
Non-native species introduction
The presence of alien species in the current assemblages in the Caribbean
trans-Andean basins are the result of escapes from sh farms. Escapes from these
farms are common and individuals of these species have colonized the trans-Andean
rivers. Their success in these aquatic systems is so high that these species have been
recently included in the report of the artisanal shery catch. Álvarez-León et al.
(2013) and Mancera-Rodríguez & Álvarez-León (2013) mention that the introduction
of the carnivore O. mykiss is highly correlated with the disappearance of the endemic
trichomycterid Rhizosomichthys totae (Miles 1942) from Andean lakes. Threats to
conservation from alien shes are poorly understood in Colombia. Recently, Gutiér-
rez et al. (2012) compiled a list of 29 alien sh species, including the piscivorous
Arapaima gigas (Schinz 1822) and Micropterus salmoides (Lacépède 1802). The
impact of these alien sh species on native sh assemblages is unknown and research
is needed.
Lost area of the oodplain lakes
Although wetlands are vital for sustaining aquatic biota and associated environmen-
tal services, the aquatic systems in the Caribbean river basins of Colombia are one
of the most threatened habitats as they are the nal receptors of all the changes in
their basin. Mismanagement of the territory by the Colombian Government is the main
cause. At local level, entire villages were built on oodplain areas that are ooded
historically by the river in a recurring ood cycle of c. 50 years. The economic activ-
ities of this population (e.g. crops and cattle) are developed on the surrounding ood-
plains. Although there is no ofcial data, an important area once covered by the lakes
has been lost because many lakes have been drained or their connection channels to
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
92 L. F. JIMÉNEZ-SEGURA ET AL.
40
30
20
10
0
–10
–20
–30
–40
80 000
60 000
40 000
20 000
0
Landing (t) SOI
1975
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
Year
30
25
20
15
10
5
0
2003
2004
2005
2006
2007
2008
2009
2010
2011
111
108
105
102
Individuals m–3
Water level (cm)
(a)
(c)
(b)
F. 8. (a) Southern Oscillation Index (SOI; , mean monthly values; , average for four periods of the monthly
values), (b) weekly ichthyoplankton densities ( ) and water level ( ) between 2004 and 2011 and (c)
riverine sheries landings ( ) of the Magdalena-Cauca River in the last four decades.
the main river were closed to increase croplands and cattle pasture area around them
and, in the last 5 years, illegal gold mining is disruptively modifying the land. At a
regional level, the increase in deforestation rate increases the sediment runoff carried
in the water, so lowland wetlands progressively lose depth. As the oodplain lakes get
shallower, land-owners surrounding these aquatic systems construct levees across the
natural connection channels to stop the inuence of the river ooding into the lake.
So, as the lake loses depth and connectivity with the basin, they dry out and nally
disappear.
Climate change
As sh reproduction and recruitment in tropical areas is highly correlated with rain
patterns and oods, changes in air temperatures and rains associated with ENSO cycles
may have impacts on freshwater sh conservation and artisanal sheries landings. In
Colombia, El Niño causes long periods of low water levels and La Niña long periods
of oods in the trans-Andean rivers (SOI, 2014). Although there is no clear associ-
ation between El Niño periods and the freshwater sher landings (Fig. 8), in some
years, higher landings have occurred during the El Niño period, higher densities of
ichthyoplankton at the beginning of the immediate oods and a new period of high
sher landings, with a positive time lag of one and a half years. If the observed cycles
of droughts-rains-oods change because of variations in the frequency and intensity of
ENSO, population sizes of migratory shes may also change and so impact the artisanal
sheries’ sustainability. Ichthyoplankton monitoring may be a useful biological vari-
able for several research questions about migratory sh recruitment in tropical rivers
and the impact of climatic change.
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 93
FUTURE DIRECTIONS FOR FISH CONSERVATION
The economic activities that have increased the gross domestic product of Colom-
bia are modifying the aquatic ecosystems where the shes live and the livelihoods of
artisanal shers. Modications of the freshwater aquatic systems and their connectiv-
ity to the trans-Andean river basins are the origin of a silent degradation that threatens
freshwater sh conservation in Colombia. Cumulative effects of the human activities
described above may lead to a signicant reduction in the population size of the sh
fauna and perhaps even the extinction of some rare species.
After the Rio Convention in 1992, the Colombian Government created the Minis-
terio de Medio Ambiente y Desarrollo Sostenible MADS (Law 33 of 1993) as the
agency responsible for the environmental management of Colombia. Fish protection
and management are function of the MADS and the Autoridad Nacional de Pesca y
Acuicultura (AUNAP). MADS protects sh diversity and AUNAP the sh as a human
food source. Some MADS advances in environmental protection includes the system of
National Parks, institutions for environmental research (IDEAM, Instituto Alexander
von Humboldt, SINCHI, among others) and the Agencia Nacional para Licenciamiento
Ambiental ANLA. Sadly, although the law, its articles and paragraphs of the Law 33
are well intentioned for the protection of environmental resources, in the real scenario
other ministries related to the economy (Energy, Transport and Agriculture) advance
faster on the exploration and use of resources (water, oil, gas and soil) than MADS
on protecting the terrestrial and aquatic systems as habitat for biota restricted to the
northern regions of South America.
Almost 10% of the terrestrial Colombian area is protected by the Parques Nacionales
Naturales Agency (c. 12 877 086 ha) (Parques Nacionales Naturales, 2015). Fifty per
cent of this protected area is located in the trans-Andean River basins, but most of them
are in the highlands, above 1000m altitude and they do not include the river basin as
a unit. MADS is also constructing a system for compensating the loss of biodiversity
as a result of infrastructure and energy development of the Colombian Government
(MADS, 2012), and after the consequences of the oods by the ENSO cycle of the
year 2009–2011 on human population, this institution is moving to dene the wetland
limits in Colombia (Vilardy et al., 2014).
As already noted, the available information about shes for each river basin is
different; most is from the Magdalena-Cauca with few scattered studies in the other
basins. Those basins, although smaller than the Magdalena-Cauca, are just as impor-
tant for local people as the main basin. More systematic and regional research must be
focused on sh conservation; shes are the source for several environmental services
provided to human populations in Colombia. Although more information is always
needed about shes, there is no time to wait for answers to all the scientic questions
before proposing some simple conservation actions. Freshwater sh conservation
and the artisanal sheries are threatened by the economic development of some
industries. The river basin must be the basic management unit for sh protection.
Actions for protecting and restoring their aquatic habitat quality and their connectivity
must be the main objective for environmental agencies during the next 50 years.
If the aquatic systems are not restored and protected, there is no future for the sh
fauna or for artisanal shers, one of the economically poorest groups in Colombian
society.
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
94 L. F. JIMÉNEZ-SEGURA ET AL.
We are grateful to all our undergraduate and post-graduate students for their enthusiasm for
discovering and knowing more about Colombian shes; this enterprise could not have been done
without them. We also thank the Colombian energy producers ISAGEN S.A. E.S.P. (Agree-
ment 46/3296, Contract No 7/4346) and Empresas Públicas de Medellín EPM (Agreement
2011–00334) for nancial support to collect some of the data included here. We also thank to
D. Taphorn for improving the English language and to unknown reviewers for their opportune
suggestions for improving the manuscript.
Supporting Information
Supporting Information may be found in the online version of this paper:
Table S1. List of freshwater sh species in the Caribbean rivers in the northern of South
America.
References
Abell, R., Thieme, M., Revenga, C., Bryer, M., Kottelat, M., Bogutskaya, N., Coad, B.,
Mandrak, N., Contreras-Balderas, S., Bussing, W., Stiassny, M., Skelton, P., Allen, G.,
Unmack, P., Naseka, A., Sindorf, N., Robertson, J., Armijo, E., Higgins, J., Heibel, T.,
Wikramanayake, E., Olson, D., López, H., Reis, R., Lundberg, J., Sabaj Pérez, M. &
Petry, P. (2008). Freshwater ecoregions of the world: a new map of biogeographic units
for freshwater biodiversity conservation. BioScience 58, 403–414.
Agostinho, A. A., Gomes, L. C. & Pelilice, F. M. (2007). Ecologia e manejo de recursos
pesqueiros em reservatorios do Brasil. Maringa: Editorial da Universidade Estadual de
Maringa.
Albert, J. & Reis, R. (2011). Historical Biogeography of Neotropical Freshwater Fishes.Berke-
ley, CA: University of California Press.
Albert, J., Petri, P. & Reis, R. (2011). Major biogeographic and phylogenetic patterns. In His-
torical Biogeography of Neotropical Freshwater Fishes (Albert, J. & Reis, R., eds), pp.
21–56. Berkeley, CA: University of California Press.
Anderson, E. & Maldonado-Ocampo, J. A. (2011). A regional perspective on the diversity and
conservation of tropical Andean shes. Conservation Biology 25, 30– 39.
Alvarez, A., Kolok, A., Jiménez-Segura, L. F., Granados, C. & Palacio, J. (2012). Mercury con-
centrations in muscle and liver tissue of sh from marshes along the Magdalena River,
Colombia. Bulletin of Environmental Contamination and Toxicology 89, 836 – 840.
Álvarez-León, R., Orozco-Rey, R. H., Páramo-Fonseca, M. E. & Restrepo-Santamaría, D.
(2013a). Peces fósiles y actuales de Colombia: distribución, diagnosis de referencia y
nombres comunes e indígenas. Bogotá: Ecoprints Diseño Gráco y Audiovisual Ltda.
Arango-Rojas, A., Jiménez-Segura, L. F. & Palacio-Baena, J. A. (2008). Variación espacio-
temporal de la asociación de especies de peces en la laguna de cachimbero, un humedal
en la cuenca media del río magdalena, Colombia. Actualidades Biológicas 30, 161–169.
Arguello, L. E., González, H. & Atencio-García, V. (2001). Reproducción inducida de la liseta
Leporinus muyscorum (Steindachner, 1902) con extracto pituitario de carpa (EPC).
Revista MVZ Córdoba 6, 97–101.
Atencio García, V., Bello Sierra, B., Bello Sierra, L. & Espinosa, J. (2005). Producción de alevi-
nos de especies nativas. Revista MVZ-Córdoba 6, 9 – 14.
Atencio-García, V., Kerguelén, E., Naar, E. & Petro, R. (2013). Desempeño reproductivo del
bocachico Prochilodus magdalenae inducido dos veces en un mismo año. Revista MVZ
Córdoba 18, 3304– 3310.
Baumgartner, G., Nakatani, K., Gomes, L. C., Bialetzki, A. & Sanches, P. (2004). Identica-
tion of spawning sites and natural nurseries of shes in the upper Paraná River, Brazil.
Environmental Biology of Fishes 71, 115–125.
Barletta, M., Jaureguizar, A. J., Baigun, C., Fontoura, N. F., Agostinho, A. A., Almeida-Val, V.
M. F., Val, A. L., Torres, R. A., Jiménez-Segura, L. F., Giarizzo, T., Fabré, N. N., Batista,
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 95
V. S., Lasso, C., Taphorn, D. C., Costa, M. F., Chaves, P. T., Vieira, J. P. & Correa, M. F.
(2010). Fish and aquatic habitat conservation in South America: a continental overview
with emphasis on Neotropical systems. Journal of Fish Biology 76, 2118–2176.
Bowen, S. (1983). Detritivory in neotropical sh communities. Environmental Biology of Fishes
9, 137–144.
Briñez-Vásquez, G. N. & Francis-Turner, L. (2006). Aspectos reproductivos de Astroblepus
homodon (Regan, 1914) (Pisces, Siluriformes) en la cuenca del río Coello, Tolima.
Revista Tumbaga 1, 5–20.
Cala, P. (1987). La ictiofauna dulceacuícola: una visión histórica y su estado actual. Revista
Colombiana Ciencias Exactas y Naturales 16, 69–84.
Cala, P. (1997). Espermatogénesis y ciclo anual reproductivo del capaz Pimelodus grosskopi
(Pisces: Pimelodidae) en alto río Magdalena (Colombia). Caldasia 19, 45–53.
Cala, P. (2001a). Ictiofauna neotropical de Colombia en el contexto global neotropical y su
estado actual: una revisión bibliográca. Dahlia 4, 3– 14.
Cala, P. (2001b). Occurrence of mercury in some commercial sh species from the Magdalena
and Meta rivers in Colombia. Dahlia 4, 15–19.
Cala, P. & Sodergren, A. (1999). Occurrence and distribution of organochlorine residues in
sh from the Magdalena and Meta rivers in Colombia. Toxicological and Environmental
Chemistry 71, 185–195.
Cala, P., Pérez, C. & Rodríguez, I. (1993). Aspectos bioecológicos de la población de capaz,
Pimelodus grosskopf (Pisces: Pimelodidae), en el embalse de Betania y parte alta del río
Magdalena, Colombia. Revista de la Academia Colombiana de Ciencias 20, 319– 330.
Carvajal-Quintero, J. D., Escobar, F., Alvarado, F., Villa-Navarro, F. A., Jaramillo-Villa, Ú.
& Maldonado-Ocampo, J. A. (2015). Variation in freshwater sh assemblages along a
regional elevation gradient in the northern Andes, Colombia. Ecology and Evolution 5,
2608–2620.
Galvis, G. & Mojica, J. I. (2007). The Magdalena River fresh water shes and sheries. Aquatic
Ecosystem Health & Management 10, 127–139.
García-Alzate, C. & Roman-Valencia, C. (2008). Biología Alimentaria y Reproductiva de
Hyphessobrycon poecilioides (Pisces:Characidae) en la Cuenca del Río La Vieja, Alto
Cauca Colombia. Revista Museo Argentino de Ciencias Naturales 10, 17–27.
Gomes, L. C. & Miranda, L. E. (2001). Riverine Characteristics dictate composition of sh
assemblages and limit sheries in reservoirs of the upper Paraná River basin. Regulated
Rivers: Research & Management 17, 67–79.
Granado-Lorencio, C., Gulfo, A., Alvarez, F., Jiménez-Segura, L. F., Carvajal-Quintero, J. D.
& Hernández-Serna, A. (2012a). Fish assemblages in oodplain lakes in a Neotropical
river during the wet season (Magdalena River, Colombia). Journal of Tropical Ecology
28, 271–279.
Granado-Lorencio, C., Hernández-Serna, A., Carvajal, J. D., Jiménez-Segura, L. F., Gulfo, A.
& Alvarez, F. (2012b). Regionally nested patterns of sh assemblages in oodplain lakes
of the Magdalena river (Colombia). Ecology and Evolution 2, 1296–1303.
Gutiérrez, F. (2011). Diagnóstico de la pesquería en la cuenca del río Sinú y Canalete. In Cat-
alogo de los recursos pesqueros continentales de Colombia (Lasso, C. A., Gutiérrez, F.
P., Morales-Betancourt, M. A., Agudelo, E., Ramírez-Gil, H. & Ajiaco-Martínez, R. E.,
eds), pp. 75–100. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander
von Humboldt (IAvH).
Gutiérrez, F. P., Lasso, C. A., Sánchez-Duarte, P. & Gil, D. L. (2010). Análisis de riesgo para
especies acuáticas continentales y marinas. In Análisis de riesgo y propuesta de catego-
rización de especies introducidas para Colombia (Baptiste, M. P., Castaño, N., Cárdenas,
D., Gutiérrez, F. P., Gil, D. L. & Lasso, C. A., eds), pp. 73 – 148. Bogotá: Instituto de
Investigación de Recursos Biológicos Alexander von Humboldt.
Gutiérrez, F., Barreto, C. & Mancilla, B. (2011a). Diagnóstico de la pesquería en la cuenca del
río Magdalena-Cauca. In Catalogo de los recursos pesqueros continentales de Colombia
(Lasso, C. A., Gutiérrez, F. P., Morales-Betancourt, M. A., Agudelo, E., Ramírez-Gil, H.
& Ajiaco-Martínez, R. E., eds), pp. 35–73. Bogotá: Instituto de Investigación de Recur-
sos Biológicos Alexander von Humboldt (IAvH).
Gutiérrez, F., Rivas-Lara, T. & Rincón-López, C. (2011b). Diagnóstico de la pesquería en la
cuenca del río Atrato. In Catalogo de los recursos pesqueros continentales de Colombia
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
96 L. F. JIMÉNEZ-SEGURA ET AL.
(Lasso, C. A., Gutiérrez, F. P., Morales-Betancourt, M. A., Agudelo, E., Ramírez-Gil,
H. & Ajiaco-Martínez, R. E., eds), pp. 103–118. Bogotá: Instituto de Investigación de
Recursos Biológicos Alexander von Humboldt (IAvH).
Gutiérrez, F., Lasso, C., Baptiste, M., Sánchez-Duarte, P. & Díaz, A. M. (2012). Catálogo de la
biodiversidad acuática exótica trasplantada en Colombia: moluscos, crustáceos, peces,
anbios, reptiles y aves. Bogotá: Universidad de Bogotá Jorge Tadeo Lozano, Instituto
de Investigación de Recursos Biológicos Alexander von Humboldt Ltda.
Hernández-Camacho, J. (1992). Caracterización geográca de Colombia. In La diversidad
biológica en Iberoamérica (Halffter, G., ed), pp. 45–54. Ciudad de México: CYTED-B,
Programa iberoamericano de ciencia y tecnología para el desarrollo.
Hernández-Serna, A., Márquez-Velásquez, V., Carvajal-Quintero, J. D., Gulfo, A., Granado-
Lorencio, C. & Jiménez-Segura, L. F. (2014). Length–weight relationships of 38 sh
species of the Magdalena River oodplain lakes. Journal of Applied Ichthyology 30,
549–551. doi: 10.1111/jai.12379
Hernández-Serna, A., Granado-Lorencio, C. & Jiménez-Segura, L. F. (2015). Diel cycle
size-dependent trophic structure of neotropical shes: a three year case analysis from 35
oodplain lakes in Colombia. Journal of Applied Ichthyology 31, 638– 645. doi: 10.1111/
jai.12748
IGAC (2003). Atlas de Colombia. Quinta edición,revisada,actualizada y aumentada. Bogotá:
Imprenta Nacional de Colombia.
Jaramillo-Villa, U., Maldonado-Ocampo, J. A. & Escobar, F. (2010). Altitudinal variation in
sh assemblage diversity in streams of the central Andes of Colombia. Journal of Fish
Biology 76, 2401–2417.
Jaramillo-Villa, U. & Jiménez-Segura, L. F. (2008). Algunos aspectos biológicos de la población
de Prochilodus magdalenae en las ciénagas de Tumaradó (Río Atrato), Colombia. Actu-
alidades Biológicas 30, 55–66.
Jaramillo, U., Cortes-Duque, J. & Flórez, C. (2015). Colombia anbia. Un país de humedales.
Vol I. Instituto de investigación de recursos hidrobiológicos Alexander von Humboldt.
Jiménez-Segura, L. F. (2007). Ictioplancton y reproducción de los peces en la cuenca media del
río Magdalena (sector de Puerto Berrio, Antioquia). PhD Thesis. Universidad de Antio-
quia. Colombia.
Jaramillo, U., Pelaez, S., Aponte, C., Flórez-Ayala, C., Avella, C., Manrique, O., Velasquez,
W., Millán, S. & Rodríguez, A. (2015a). Hacía un inventario completo de humedales. In
Colombia anbia. Un país de humedales. Vol I. (Jaramillo, U., Cortes-Duque, J. & Flórez,
C. eds), pp. 108–109. Instituto de investigación de recursos hidrobiológicos Alexander
von Humboldt. Bogotá, Colombia.
Jiménez-Segura, L. F., Palacio, J. & López, R. (2009). Características biológicas del Blanquillo
Sorubim cuspicaudus Littmann, Burr y Nass, 2000 y Bagre Rayado Pseudoplatystoma
magdaleniatum Buitrago-Suárez y Burr, 2007 (Siluriformes: Pimelodidae) relacionadas
con su reproducción en la cuenca media del río Magdalena (Colombia). Actualidades
Biológicas 31, 53–66.
Jiménez-Segura, L. F., Palacio, J. & Leite, R. (2010a). River ooding and reproduction of migra-
tory sh species in the Magdalena River Basin, Colombia. Ecology of Freshwater Fish
19, 178–186.
Jiménez-Segura, L. F., Carvajal-Quintero, J. D. & Aguirre, N. (2010b). Las ciénagas como
hábitat para los peces: estudio de caso en la ciénaga de Ayapel (Córdoba), Colombia.
Actualidades Biológicas 32, 53–64.
Jiménez-Segura, L. F. & Ortega-Lara, A. (2010). Pseudopimelodus buffonius (Siluriformes,
Pimelodidae). In Catalogo de los recursos pesqueros continentales de Colombia (Lasso,
C. A., Gutiérrez, F. P., Morales-Betancourt, M. A., Agudelo, E., Ramírez-Gil, H. &
Ajiaco-Martínez, R. E., eds), pp. 544–546. Serie Editorial Recursos Hidrobiológicos y
Pesqueros Continentales de Colombia. Instituto de Investigación de Recursos Biológicos
Alexander von Humboldt (IAvH). Bogotá, D. C., Colombia.
Jiménez-Segura, L. F. & Villa-Navarro, F. A. (2011). Pimelodus grosskopi (Siluri-
formes,Pimelodidae). In Catálogo de los Recursos Pesqueros Continentales de
Colombia (Lasso, C. A., Agudelo Córdoba, E., Jiménez-Segura, L. F., Ramírez-Gil, H.,
Morales-Betancourt, M., Ajiaco-Martínez, R. E., Gutiérrez, F. de P., Usma Oviedo, J. S.,
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 97
Muñoz Torres, S. E. & Sanabria Ochoa, A. I., eds), pp. 466–471. Bogotá: Instituto de
Investigación de Recursos Biológicos Alexander von Humboldt (IAvH).
Jiménez-Segura, L. F., Restrepo-Santamaría, D., López-Casas, S., Delgado, J., Valderrama, M.,
Álvarez, J. & Gómez, D. (2014a). Ictiofauna y desarrollo del sector hidroeléctrico en la
cuenca del río Magdalena – Cauca, Colombia. Biota Colombiana 15, 3–25.
Jiménez-Segura, L. F., Maldonado-Ocampo, J. A. & Pérez, C. (2014b). Gradiente de recu-
peración longitudinal en la estructura de la ictiofauna en un río andino regulado. Biota
Colombiana 15, 61–80.
Jiménez-Segura, L. F., Álvarez, J., Ochoa, L. E., Loaiza, A., Londoño, J. P., Restrepo,
D., Aguirre, K., Hernández, A., Correa, J. D. & Jaramillo-Villa, U. (2015). Guía
Ilustrada Peces Cañón del río Porce, Antioquia. Medellín: Empresas Públicas de
Medellín-Universidad de Antioquia.
Junk, W. J., Bayley, P. B. & Sparks, R. E. (1989). The ood pulse concept in river-oodplain
systems. Canadian Special Publication of Fisheries and Aquatic Sciences 106, 110–127.
Kerguelen-Durango, E. & Atencio-García, V. (2015). Environmental characterization of the
reproductive season of migratory sh of the Sinú river (Córdoba, Colombia). Revista
MVZ Córdoba 20, 4766– 4778.
Lasso, C. A., Gutiérrez, F. P., Morales-Betancourt, M. A., Agudelo, E., Ramírez-Gil, H. &
Ajiaco-Martínez, R. E. (2011a). Catálogo de los recursos pesqueros continentales de
Colombia. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander von
Humboldt (IAvH).
Lasso, C., Morales-Betancourt, M. & Sanchez-Duarte, P. (2011b). Recursos pesqueros conti-
nentales de Colombia. In Catalogo de los recursos pesqueros continentales de Colombia
(Lasso, C. A., Gutiérrez, F. P., Morales-Betancourt, M. A., Agudelo, E., Ramírez-Gil, H.
& Ajiaco-Martínez, R. E., eds), pp. 57–68. Bogotá: Instituto de Investigación de Recur-
sos Biológicos Alexander von Humboldt (IAvH).
Lewis, W. M. (2008). Physical and chemical features of tropical owing waters. In Tropical
Stream Ecology (Dudgeon, D., ed), pp. 1–21. London: Academic Press.
López-Casas, S. (2015). Magdalena potadromous migrations: effects of regulated and natural
hydrological regimes. Doctoral thesis. Instituto de Biología, Universidad de Antioquia.
Medellín, Colombia.
Lowe-McConnell, R. H. (1995). Ecological Studies in Tropical Fish Communities. Cambridge:
Cambridge University Press.
Lucas, M. & Baras, E. (2001). Migration of Freshwater Fishes. Oxford: Blackwell Science.
Mancera-Rodríguez, N. J. & Álvarez-León, R. (2005). Estado del conocimiento de las con-
centraciones de hidrocarburos y residuos organoclorados en los peces dulceacuícolas de
Colombia. Dahlia 8, 89–103.
Mancera-Rodríguez, N. J. & Álvarez-León, R. (2006). Estado del conocimiento de las concen-
traciones de metales pesados en los peces dulceacuícolas de Colombia. Acta Biologica
Colombiana 11, 3–23.
Maldonado-Ocampo, J. A., Ortega-Lara, A., Usma, J. S., Galvis, G., Villa-Navarro, F., Vásquez,
L., Prada-Pedreros, L. & Rodríguez, C. A. (2005). Peces de los Andes de Colombia:
guía de campo. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander
von Humboldt.
Maldonado-Ocampo, J. A., Vari, R. & Usma, J. S. (2008). Checklist of the freshwater shes in
Colombia. Biota Colombiana 9, 143–237.
Maldonado-Ocampo, J. A., Usma, J. S., Villa-Navarro, F., Ortega-Lara, A., Prada-Pedreros, S.,
Jiménez-Segura, L. F., Jaramillo-Villa, U., Arango, A., Rivas, T. & Sánchez, G. C. (2013).
Peces Dulceacuícolas del Choco Biogeográco. Bogota: WWF Colombia, Instituto de
Investigaciones Recursos Biológicos IAvH, Universidad del Tolima, Autoridad Nacional
de Acuicultura y Pesca (AUNAP), Ponticia Universidad Javeriana.
Mancera-Rodríguez, N. J. & Álvarez-León, R. (2013). Colombia, la pesca en un país en desar-
rollo. In La Pesca: entre sus circunstancias y consecuencias (Castro-Hernández, J. J.,
ed), pp. 305–341. Las Palmas de Gran Canaria: Colección Textos Universitarios Anroat
Ediciones, S.L.
Mancera-Rodríguez, N. J., Márquez, E. J. & Hurtado-Alarcón, J. C. (2013). Uso de citogenética
y técnicas moleculares en estudios de diversidad genética en peces Colombianos. In Uso
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
98 L. F. JIMÉNEZ-SEGURA ET AL.
de Biología Molecular en Producción Animal y Conservación de Especies Silvestres
(López Herrera, A., ed), pp. 237–312. Medellín: Centro de publicaciones, Universidad
Nacional de Colombia.
Mancera-Rodríguez, N. J., Castellanos-Barliza, J. & Urrego-Ballestas, D. (2016). Biología
Reproductiva de Saccodon dariensis (Teleostei: Parodontidae) en auentes del río
Guatapé, cuenca del río Magdalena, Colombia. Revista de Biología Tropical 64. doi:
10.15517/rbt.v64i2.20691
Marrugo-Negrete, J., Lans, E. & Benitez, L. (2007). Finding of mercury in sh from the Ayapel
marsh, Cordoba, Colombia. Revista MVZ Córdoba 12, 878–886.
Marrugo-Negrete, J., Olivero-Verbel, J., Lans, E. & Benitez, L. (2008a). Total mercury and
methylmercury concentrations in sh from the Mojana region of Colombia. Environ-
mental Geochemistry and Health 30, 21–30.
Marrugo-Negrete, J., Benitez, L. & Olivero-Verbel, J. (2008b). Distribution of mercury in
several environmental compartments in an aquatic ecosystem impacted by gold mining
in northern Colombia. Archives of Environmental Contamination and Toxicology 55,
305–316.
Matthews, W. (1998). Patterns in Freshwater Fish Ecology. New York, NY: Springer.
Mojica, J. I. (2002). Las pesquerías de la cuenca del Magdalena: Ejemplo a no repetir. In Libro
rojo de peces dulceacuícolas de Colombia (Mojica, J. I., Castellanos, C., Usma, S. &
Álvarez-León, R., eds), pp. 35– 41. Bogotá: Instituto de Ciencias Naturales Universidad
Nacional de Colombia, Ministerio del Medio Ambiente.
Mojica, J. I., Usma, J. S., Álvarez-León, R. & Lasso, C. A. (2012). Libro rojo de peces dulceacuí-
colas de Colombia. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander
von Humboldt, Instituto de Ciencias Naturales de la Universidad Nacional de Colombia,
WWF Colombia y Universidad de Manizales.
Mojica, J. I., Castellanos, C., Sánchez-Duarte, P. & Díaz, C. (2006a). Peces de la cuenca del río
Ranchería, La Guajira Colombia. Biota Colombiana 7, 129–142.
Mojica, J. I., Galvis, G., Sánchez-Duarte, P., Castellanos, C. & Villa-Navarro, F. (2006b). Peces
del valle medio del Río Magdalena, Colombia. Biota Colombiana 7, 23– 38.
Munro, A. D. (1990). General Introduction. In Reproductive Seasonality in Teleosts: Environ-
mental Inuences (Munro, A. D., Scott, A. P. & Lam, T. J., eds), pp. 1– 11. Boca Raton,
FL: CRC Press Inc.
Noreña, R. D. A., Arenas, T. A. M., Murillo, P. E., Guío, D. A. J. & Méndez, A. J. J. (2012).
Heavy metals (Cd, Pb and Ni) in sh species commercially important from Magdalena
river, Tolima tract, Colombia. Revista Tumbaga 7, 61–76.
Olaya-Nieto, C., Hernández-Rosso, D. & Ayarza-Pérez, E. (2010). Reproductive Biology of
Liso Rhamdia quelen (Pisces: Heptapteridae) in the Sinu River, Colombia. Acta Biológica
Colombiana 15, 61–74.
Olsson, J. & Eklöv, P. (2005). Habitat structure, feeding mode and morphological reversibil-
ity: factors inuencing phenotypic plasticity in perch. Evolutionary Ecology Research 7,
1109–1123.
Ortega-Lara, A., Usma, J. S., Bonilla, P. & Santos, N. (2006). Peces de la cuenca alta del río
Cauca, Colombia. Biota Colombiana 7, 39–54.
Perez, A. N., Olaya-Nieto, C. W., Segura-Guevara, F. F., Tordecilla-Petro, G. & Bru-Cordero,
S. (2005). Relaciones longitud-peso de la doncella, Ageneiosus pardalis (Pisces:
Auchenipteridae), en la cuenca del río Sinú, Colombia. Dahlia 9, 53–61.
Ramírez, J., Cifuentes, C. Y. & Avilés, M. (2013). Evaluación de la reproducción inducida de
nicuro Pimelodus blochii (Teleostei: Pimelodidae) utilizando diferentes inductores hor-
monales. Revista MVZ Córdoba 18, 3512–3517.
Rangel-Serpa, F. & Torres-Mejia, M. (2015). Reproductive seasonality of Geophagus stein-
dachneri Eigenmann & Hildebrand, 1922 (Perciformes: Cichlidae) in a tropical mountain
river. Neotropical Ichthyology 13, 421– 430. doi: 10.1590/1982-0224-20140091
Restrepo, J. D. & Kjerfve, B. (2000). Magdalena river: interannual variability (1975–1995)
and revised water discharge and sediment load estimates. Journal of Hydrology 235,
137–149.
Restrepo-Gómez, A. M. & Mancera-Rodríguez, N. J. (2014). Trophic ecology of Saccodon
dariensis (Pisces: Parodontidae) in Guatapé River tributaries, Magdalena River Basin,
Colombia. Revista MVZ Córdoba 19, 3930–3943.
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 99
Ríos-Pulgarín, M. I., Jiménez-Segura, L. F., Palacio, J. A. & Ramírez-Restrepo, J. J. (2008).
Comunidad de peces en la Ciénaga de Ayapel, Río Magdalena (Córdoba) Colom-
bia: cambios espacio-temporales en su asociación. Actualidades Biológicas 30,
29–53.
Roberts, T. R. (1974). Dental polymorphism and sistematics in Saccodon, a neotropical
genus of fresh water shes (Parodontidae, Characoidei). Journal of Zoology 173,
301–321.
Roman-P, C., Román-Valencia, C. & Taphorn, C. (2014). Trophic and reproductive ecology of a
neotropical Characid sh Hemibrycon brevispinnis (Teleostei: Characiformes). Caldasia
36, 289–304.
Román-Valencia, C. & Botero, A. (2006). Trophic and reproductive ecology of a species of
Hemibrycon (Pisces: Characidae) in Tinajas creek, Quindío River drainage, upper Cauca
Basin, Colombia. Revista Museo Argentino de Ciencias Naturales 8, 1–8.
Román-Valencia, C. & Muñoz, A. (2001). Ecología tróca y reproductiva de Bryconameri-
cus caucanus (Pisces: Characidae). Bolletino Museum Regionalli Science Naturalli 18,
459–467.
Román-Valencia, C. & Ruiz, R. (2005). Diet and reproduction aspects of Astyanax aurocaudatus
(Teleostei: Characidae) from the upper part of the rio Cauca, Colombia. Dahlia 8, 9–17.
Román-Valencia, C., Botero, A. & Ruiz, R. (2003). Trophic and reproductive ecology of Roe-
boides dayi (Teleostei: Characidae) from upper Rio Cauca, Colombia. Bolletino Museum
Regionalli Science Naturalli 20, 487–496.
Román-Valencia, C. & Samudio, H. (2007). Dieta y reproducción de Lasiancistrus caucanus
(Pisces: Loricariidae) en la cuenca del río La Vieja, Alto Cauca, Colombia. Revista del
Museo Argentino de Ciencias Naturales nueva serie 9, 95– 101.
Román-Valencia, C., Ruiz, R. & Giraldo, A. (2008). Dieta y reproducción de dos especies
sintópicas: Hemibrycon boquiae yBryconamericus caucanus (Pisces: Characidae) en
la quebrada Boquía, río Quindío, Alto Cauca, Colombia. Revista del Museo Argentino
Ciencias Naturales 10, 55– 62.
Solano-Peña, D., Segura-Guevara, F. & Olaya-Nieto, C. (2013). Crecimiento y reproducción
de la mojarra amarilla (Caquetaia kraussii Steindachner, 1878) en el embalse de Urrá,
Colombia. Revista MVZ Córdoba 18, 3525–3533.
Torres-Mejía, M. & Ramírez-Pinilla, M. (2008). Dry-season breeding of a characin in a neotrop-
ical mountain river. Copeia 2008, 99– 104.
Trujillo, F., Lasso, C. A., Diazgranados, M. C., Farina, O., Pérez, L. E., Barbarino, A., González,
M. & Usma, J. S. (2010). Evaluación de la contaminación por mercurio en peces de
interés comercial y de la concentración de organoclorados y organofosforados en el agua
y sedimentos de la Orinoquia. In Biodiversidad de la Cuenca del Orinoco. Bases cien-
tícas para la identicación de áreas prioritarias para la conservación y uso sostenible
de la biodiversidad (Lasso, C. A., Usma, J. S., Trujillo, F. & Rial, A., eds), pp. 175–191.
Bogotá: Instituto de Investigación de Recursos Biológicos Alexander von Humboldt,
WWF Colombia, Fundación Omacha, Fundación La Salle e Instituto de Estudios de la
Orinoquia. Universidad Nacional de Colombia.
Rodríguez-Olarte, D., Mojica, J. I., Corzo, J. & Taphorn, D. (2011). Northern South America,
Magdalena and Maracaibo Basins. In Historical Biogeography of Neotropical Fresh-
water Fishes (Albert, J. & Reis, R., eds), pp. 243–258. Berkeley, CA: University of
California Press.
Tucker, C. J. & Townshend, J. R. G. (2000). Strategies for monitoring tropical deforestation
using satellite data. International Journal of Remote Sensing 21, 1461–1471.
Val, A. L. & Randall, D. J. (2005). The Physiology of Tropical Fishes, Vol. 21. London: Aca-
demic Press.
Vazzoler, A. M. (1996). Biologia da reprodução de peixes teleósteos: Teoria e prática. Nupelia:
DAUFSC, Editorial Universidade de Maringá.
Villa, F. (1992). La variable ambiental en el contexto de la pesquería de la cuenca del Mag-
dalena: Ibagué Fundación Rio Magdalena. Memorias del Seminario Presente y Futuro
del Rio Magdalena.
Villa-Navarro, F., Zúñiga-Upegui, P., Castro-Roa, D., García-Melo, J., García-Melo, L.
& Herrada-Yara, M. (2006). Peces del alto Magdalena, cuenca del río Magdalena,
Colombia. Biota Colombiana 7, 3–21.
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
100 L. F. JIMÉNEZ-SEGURA ET AL.
Welcomme R. L. (1985). River sheries. FAO Fisheries Technical Paper 262.
Welcomme, R. L. (1999). A review of a model for qualitative evaluation of exploitation levels
in multi-species sheries. Fisheries Management and Ecology 6, 1–19.
Welcomme, R. L., Winemiller, K. O. & Cowx, I. G. (2006). Fish environmental guilds as a tool
for assessment of ecological condition of rivers. River Research and Applications 22,
377–396. doi: 10.1002/rra.914
Wootton, R. J. (1999). Ecology of Teleost Fishes. Dordrecht: Kluwer Academic Publishers.
Electronic References
Álvarez-León, R., Orozco-Rey, R.H., Páramo-Fonseca, M.E. & Restrepo-Santamaría, D.
(2013b). Peces fósiles y actuales de Colombia: distribución, diagnosis de referencia
y nombres comunes e indígenas. Bogotá :Ecoprints Diseño Gráco y Audiovisual
Ltda. Available at http://gerenciaecoprints.wix.com/eco-prints#!servicios2/c1m2r (last
accessed 19 March 2015).
AUNAP – Autoridad Nacional de Acuicultura y Pesca (2014a). Servicio Estadístico Pesquero
Colombiano SEPEC. Available at http://sepec.unimagdalena.edu.co/ (last accessed 27
December 2014).
AUNAP – Autoridad Nacional de Acuicultura y Pesca (2014b). Estado de la pesca y la acuicul-
tura 2014. Available at http://www.aunap.gov.co/les/ESTADO_DE_LA_PESCA_Y_
ACUICULTURA_2014_.pdf (last accessed 15 March 2015).
Baptiste, M. P., Castaño, N., Cárdenas, D., Gutiérrez, F. P., Gil, D. L. & Lasso, C. A. (2010).
Análisis de riesgo y propuesta de categorización de especies introducidas para Colom-
bia. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander von Humboldt.
Available at http://www.humboldt.org.co/component/k2/item/235-analisis-de-riesgo-y-
propuesta-de-categorizacion-de-especies-introducidas-para-colombia (last accessed 19
March 2015).
Bialetzki, A., P. Sanches, M. Cavicchioli, G. Baumgartner, R. Pereira, & Nakatani, K. (1999).
Drift of ichthyoplankton in two channels of the Paraná River, between Paraná and Mato
Grosso do Sul states, Brazil. Brazilian Archives of Biology and Technology 42 (1).
10.1590/S1516-89131999000100008
CCI – Corporación Colombia Internacional (2007). Pesca y Acuicultura Colombia 2006. Avail-
able at http://cpps.dyndns.info/cpps-docsweb/planaccion/biblioteca/pordinario/Proceso
%20Ordinario/Fisheries/INCODER2006_InformePescaAcuicultura.pdf (last accessed
11 June 2015).
DANE – Departamento Administrativo Nacional de Planeación (2015). Censo poblacional.
Available at http://www.dane.gov.co/index.php/poblacion-y-demograa/ (last accessed
19 March 2015).
Eschmeyer W. & Fong, J. D. (2016). Catalog of Fishes. Available at http://researcharchive.cal
academy.org/research/ichthyology/catalog/shcatmain.asp (last accessed 27 December
2014)
Escobar, M. D., Olaya, H., Cusva, A., Lasso, C. A. & Londoño, M. C. (2014). Mapa de oferta
potencial de servicios ecosistémicos en complejos de humedales, asociados a la pesca
continental de Colombia. Bogotá: Instituto Alexander von Humboldt.
FAO – Food and Agriculture Organization of the United Nations. (2014). State of the World
Fisheries and Aquaculture. Available at http://www.fao.org/3/a-i3720e.pdf (last accessed
19 March 2015).
Froese, R. & Pauly, D. (2015). FishBase. Available at http://www.shbase.org/ (last accessed
25 March 2015).
González, J. J., Etter, A. A., Sarmiento, A. H., Orrego, S. A., Ramírez, C., Cabrera, E.,
Vargas, D., Galindo, G., García, M. C. & Ordoñez M. F. (2011). Análisis de tenden-
cias y patrones espaciales de deforestación en Colombia. Instituto de Hidrología,
Meteorología y Estudios Ambientales-IDEAM. Gobierno de Colombia. Available
at www.researchgate.net/ …tendencias_y_patrones_espaciales …Colombia/ (last
accessed 19 March 2015)
MADS – Ministerio de Medio Ambiente y Desarrollo Sostenible (2012). Manual para
la asignación de compensaciones por pérdida de biodiversidad. Available at
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
FRESHWATER FISHES OF NORTH-WEST SOUTH AMERICA 101
http://www.tremarctoscolombia.org/pdf/MANUAL_compensaciones%20Final.pdf
(last accessed 23 March 2015)
MADS – Ministerio de Medio Ambiente y Desarrollo Sostenible (2014). Colombia revela
su primera tasa anual de deforestación. Available at https://www.minambiente.gov.co/
index.php/sala-de-prensa/2-noticias/1236-el-uso-sostenible-de-los-bosques-prioridad-
de-minambiente-531 http://www.parquesnacionales.gov.co/portal/colombia-revela-su-
primera-tasa-anual-de-deforestacion/ (last accessed 28 December 2014)
SOI – Southern Oscillation Index (2014). Climate Prediction Center. Available at http://www.
cpc.ncep.noaa.gov/products/precip/CWlink/MJO/enso.shtml (last accessed 19 March
2015).
Usma-Oviedo S., Villa-Navarro, F., Lasso, C. A., Castro, F., Zuñiga, P. T., Cipamocha,
C. A., Ortega-Lara, A., Ajiaco, R. E., Ramirez-Gil, H., Jimenez-Segura, L. F.,
Maldonado-Ocampo, J. A., Muñoz J. & Suarez, J. T. (2013). Peces Dulceacuícolas
Migratorios. In Guía de las especies migratorias de la biodiversidad en Colombia,Vol.
2 (Zapata I.A. & Usma, J.S., eds), pp. 216–485. Bogotá :Ministerio de Ambiente y Desar-
rollo Sostenible. WWF-Colombia. Available at http://awsassets.panda.org/downloads/
migratoriaspeces_42_web_nal.pdf (last accessed 19 March 2015).
Vilardy, S., Jaramillo, Ú., Flórez, C., Cortés-Duque, J., Estupiñán, L., Rodríguez, J. & Aponte,
C. (2014). Principios y criterios para la delimitación de humedales continentales: una
herramienta para fortalecer la resiliencia y la adaptación al cambio climático en Colom-
bia. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander von Humboldt.
Available at https://www.siac.gov.co/documentos/GestCont/Cartilla_humedales_
inteactivo_1.pdf (last accessed 19 March 2015).
© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, 89, 65–101
