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Revista de Biología Tropical, ISSN: 2215-2075, Vol. 69(2): 573-587, April-June 2021 (Published Apr. 26, 2021)

Potential distribution in Colombia of the introduced fish

Pangasianodon hypophthalmus (Siluriformes: Pangasiidae)

and implications for endangered native fish

María C. Castellanos-Mejía

Juliana Herrera

Elkin A. Noguera-Urbano

Edison Parra

Luz F. Jiménez-Segura

1. Laboratorio de Ictiología, Instituto de Biología, Universidad de Antioquia, Medellín, Colombia;

mcastel7nbk@gmail.com (*Correspondence), juliana.herrera.p@udea.edu.co, luz.jimenez@udea.edu.co

2. Departamento de Ciencias Biológicas, Escuela de Ciencias, Universidad EAFIT, Medellín, Colombia;

mcastel7@eafit.edu.co

3. Instituto de Investigación de Recursos Biológicos, Alexander von Humboldt, Bogotá, Colombia;

elkalexno@gmail.com

4. Grupo de Investigación en Limnología Básica y Experimental y Biológica y Taxonómica Marina, Instituto de biología,

Universidad de Antioquia, Medellín, Colombia; edisonparra50@gmail.com

Received 15-X-2020. Corrected 05-IV-2021. Accepted 12-IV-2021.

ABSTRACT

Introduction: One of the significant threats to aquatic ecosystems is introducing non-native species in natural

environments. This represents one of the principal causes of biodiversity loss in the world. Pangasianodon

hypophthalmus is an introduced species in Colombia that shares life characteristics and habitat with endangered

native species, such as Ageneiosus pardalis, Sorubim cuspicaudus, and Pseudoplatystoma magdaleniatum.

However, its distribution is little-known, and scientists have barely explored the effects on the native fauna.

Objectives: To evaluate the P. hypophthalmus invasive potential in some of the Colombian basins, which drain

into the Caribbean Sea. Methods: Using records available in various databases, we performed a niche conser-

vatism analysis between the native and introduced records of P. hypophthalmus using the R package Ecospat.

Subsequently, we modeled the potential invasion area of P. hypophthalmus and the distribution areas of three

native species, performing ecological niche modeling (ENM) using the Maxent algorithm. Finally, we calculated

a geographic niche overlap between the non-native and native species. Results: The Ecospat spatial analysis

indicated that P. hypophthalmus retains some environmental niche attributes through space. For this reason,

we can use ENM as an approximation to its range of distribution in the invaded area. Our results using ENM

demonstrated that the four species analyzed prefer low and slightly rocky regions; therefore, the geographical

overlap of the three native species’ ecological niches and the introduced species exceed 80 %. Conclusions:

There are adequate conditions in the study basins to fully establish the species P. hypophthalmus, representing

a high risk for aquatic ecosystems and native ichthyofauna. Knowledge of the potential distribution areas is

essential to implement control of the species.

Key words: niche models; freshwater; invasive species; migratory fish.

Castellanos-Mejía, M.C., Herrera, J., Noguera-Urbano, E.A.,

Parra, E., & Jiménez-Segura, L.F. (2021). Potential

distribution in Colombia of the introduced fish

Pangasianodon hypophthalmus (Siluriformes: Pangasiidae)

and implications for endangered native fish. Revista de

Biología Tropical, 69(2), 573-587. DOI 10.15517/rbt.

v69i2.44223

DOI 10.15517/rbt.v69i2.44223

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Revista de Biología Tropical, ISSN: 2215-2075 Vol. 69(2): 573-587, April-June 2021 (Published Apr. 26, 2021)

Globalization through increased trade,

transport, travel, and tourism will inevitably

increase the intentional or accidental intro-

duction of organisms to new environments

(CABI, 2020) and is still steadily growing

(Hulme et al., 2008; van Kleunen, Schlaep-

fer, Glaettli, & Fischer, 2011). These intro-

ductions occasionally result in alien species’

invasions (Lockwood, Hoopes, & Marchetti,

2007). Invasive species have established popu-

lations outside their native distribution ranges

and can spread and affect native ecosystems

(Lockwood et al., 2007). These species can

trigger ecological imbalances, trophic struc-

ture changes, native species displacements,

biodiversity loss, genetic diversity reduction

of native species, and non-native infectious

agents’ transport (Cassemiro, Bailly, da Graça,

& Agostinho, 2018; Hobbs, 2000). Thus, the

introduction and later invasion of non-native

species represent one of the leading causes of

biodiversity loss in the world (Cattau, Martin,

& Kitchens, 2010). The biggest problem asso-

ciated with the introduced species is that they

have been already established and extended

when they are recognized as invasive, making

it almost impossible to eliminate or control

them (Sato et al., 2010). In some cases, native

species declines often co-occur and in the

same place as invasion by non-native species,

leading many conservationists and researchers

to believe that invasions and extinctions are

closely linked (Gurevitch & Padilla, 2004).

For that reason, a prime objective of invasion

biology is predicting which species are likely

to become invaders and where they are likely

to invade even before introduction outside their

native range, which represents a goal of inva-

sion ecologists (Fournier, Penone, Pennino, &

Courchamp, 2019).

The ecological niche modeling (ENM)

use associations between environmental vari-

ables and occurrence locations of the species

to predict the potential presence of the species

in areas that have those ideal environmental

conditions or that are appropriate for the sur-

vival of the species (Raxworthy et al., 2003;

Raxworthy, Ingram, Rabibisoa, & Pearson,

2007). This modeling has many applications,

such as understanding species’ ideal environ-

mental conditions, predicting the existence

of unknown species, inferring aspects of the

biogeography of species, planning conserva-

tion areas, and as in this case, evaluating the

potential areas of invasion of species (Peterson,

2006). In this way, ENM tools generates distri-

bution maps of species, which are usefull for

designing and leading strategies about control

and mitigation of ecological problems gener-

ated by invasion processes, such as establish-

ment, spread, and impact (Jiménez-Valverde

et al., 2011). The invasion of species can be

a highly predictable process if one knows

the environmental conditions in the native

range (Peterson, 2003; Jiménez-Valverde et al.,

2011). The invasive species’ geographic poten-

tial elaborated with ENM techniques is usually

constructed assuming evolutionary conserva-

tism in ecological niche characteristics. This

means that species will follow the same set of

ecological rules on invaded distributional areas

as they do on their native distributional areas

(Peterson, 2006); however, we should take this

assumption with caution because both biotic

and abiotic factors are different in native and

non-native regions (Lockwood et al., 2007).

In recent years, evidence that changes between

the native and non-native niche exist in fish

and other organisms have been found (Lauzeral

et al., 2011). However, the ENM allows valid

estimates of the geographical distribution of

invasive species or their invasiveness (Yiwen,

Bi Wei, & Darren, 2016; Srivastava, Lafond, &

Griess, 2019).

Pangasianodon hypophthalmus (Sauvage,

1878) family Pangasiidae (Siluriformes) is

native to the Mekong and Chao Phraya rivers

basin in Southeast Asia. It carries out long-

distance movements during hundreds of kilo-

meters between ecosystems in high areas and

spawning habitats. This species uses floodplain

lands to feed and breed (FAO, 2010; IAvH, s.f.;

Tarkan et al., 2020). A female in suitable con-

ditions can produce around 100 000 eggs per

kilogram in each oviposition, and its demersal

embryos suggest some parental care (e.g., nests

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on a hard substrate). These hatch in 24 hours,

and the larvae drift in favor of the floods,

searching for growth zones within flooded

areas. Its larvae have an omnivorous diet that

includes algae, seeds, crustaceans, zooplank-

ton, and fish (Van Zalinge, Lieng, Ngor, Heng,

& Valbo-Jørgensen, 2002). As an adult, it is

carnivorous and even cannibalistic. The species

has been reported as a predator of native spe-

cies in some places of Asia (Pallewatta, Reaser,

& Gutierrez, 2003).

Combining the aforementioned biological

characteristics and the escape and introduction

of exemplary species in places other than their

natural distribution justify their classification

as an invasive species or a potentially invasive

species (Gutiérrez, Lasso, Baptiste, Sánchez-

Duarte, & Díaz, 2012; Garcia et al., 2018). Its

broad food spectrum, the ease with which it

gains weight, and its fast growth have made it

possible to introduce this species with produc-

tive purposes in different countries such as

Cuba, Chile, Colombia, Guatemala, México,

the United States, Indonesia, India, Turkey,

Puerto Rico, and The Dominican Republi-

can (IAvH, s.f.). However, fish farming of P.

hypophthalmus presents problems such as the

spreading of pathogens that are difficult to

control (Bigarré et al., 2009; Lakra & Singh,

2010; Mitra, Bandyopadhyay, Gong, Goswami,

& Bhowmik, 2013), and increasing cost for

its production (Lakra & Singh, 2010; Singh

& Lakra, 2011).

In Colombia, the species started to be used

as an ornamental fish, and in 2008 its meat

began to be imported for national consump-

tion in frozen fillets. Currently, it is sold in

fresh fish marketplaces (Valderrama, Mojica,

Villalba, & Avila, 2016). Pangasianodon hypo-

phthalmus farming has been reported in at least

five states of the country: Valle del Cauca,

Cauca, Huila, Meta and Santander (Gutiérrez

et al., 2012). Since 2015, live specimens have

been captured by fishermen in the aquatic

ecosystems of Magdalena River, in floodplains

of the lower zone, and in main riverbeds

and tributary rivers (Valderrama et al., 2016).

However, the distribution of this species in

Colombia is little-known, and scientists have

barely explored the effects on the native fauna.

In the Caribbean sub-basins such as the Mag-

dalena, Atrato, Sinú, and Catatumbo rivers,

there are other siluriform species that, like P.

hypophthalmus, carry out long-distance migra-

tions between small tributaries and floodplains,

have carnivorous habits, and use floodplains

as breeding and growth areas. Among these

species are Pseudoplatystoma magdaleniatum,

Sorubim cuspicaudus, and Ageneiosus par-

dalis, which are of high importance for com-

mercial and artisanal fishing and are under a

critical conservation status (P. magdaleniatum)

and vulnerable (S. cuspicaudus and A. parda-

lis) (Mojica, Usma, Álvarez-León, & Lasso,

2012). In these sub-basins, only the Magda-

lena river basin contributes 70 % of the gross

domestic product, and 80 % of the population

lives in its territory (The Nature Conservancy,

Fundación Alma, Fundación Humedales, &

AUNAP, 2016). As a result, this basin presents

several environmental conflicts in its aquatic

ecosystems (Barletta et al., 2010; Galvis &

Mojica, 2007; Jiménez-Segura et al., 2016).

Although habitat destruction and human pres-

sures are major causes of this conservation

status, potential competitive interactions could

occur with introduced species, which might

worsen the conservation state of native species.

For this reason, our objective was to evaluate

the invasive potential of P. hypophthalmus in

some of the basins of the Caribbean Sea water-

shed (1) making a comparison between the

native niche and the niche of introduction in

the Caribe watershed, (2) transferring the ENM

calibrated with the native area of this species

in Colombia to calculate the possible area of

invasion and (3) comparing the geographical

distribution of P. hypophthalmus with the dis-

tribution of three native species (P. magdaleni-

atum, S. cuspicaudus, and A. pardalis) present

in rivers of the Caribbean basin.

MATERIALS AND METHODS

Study area: We focused on 5 sub-basins

of Caribbean Basin: Magdalena-Cauca, Atrato,

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Sinú, and Catatumbo. In the Caribbean basin

located North of the Andes, 326 fish spe-

cies were reported, from which 66 % were

endemic (DoNascimiento et al., 2017). Of

these endemic species, 35 were reported based

on specimens captured by artisanal fishermen

(Jiménez-Segura, Gutierrez, Ajiaco-Martínez,

& Lasso, 2020a); 19 of them have migratory

behaviors and move seasonally among low

areas and the Magdalena, Atrato, Sinú and

Ranchería river tributaries (Jiménez-Segura

et al., 2016; López-Casas, Jiménez-Segura,

Agostinho, & Pérez, 2016).

Data of presence: The georeferenced

records of the wildlife of P. hypophthalmus

both in the native and introduced ranges and of

P. magdaleniatum, S. cuspicaudus, and A. par-

dalis were compiled from the review of scien-

tific articles, reports of field observations, and

databases available online as Freshwater Biodi-

versity Data Portal (Biofresh, 2012), the Global

Biodiversity Information Facility (GBIF, 2019;

GBIF, 2020a; GBIF, 2020b; GBIF, 2020c) and

the platform InvBasa through SIB Colombia

(InvBasa UN, 2020; SiB Colombia, 2020).

Databases were reviewed and curated, apply-

ing verification procedures of duplicate records

or taxonomic uncertainty (Chapman, 2005).

Records that did not coincide with water sourc-

es were relocated to the nearest pixel (Domisch,

Wilson, & Jetz, 2016).

Environmental data: We selected 70

environmental variables from the freshwater

systems that have a relationship with the ana-

lyzed species’ biology (for example, flow, sub-

strate type, and surrounding vegetation (Nori &

Rojas-Soto, 2019). The variables are available

on the EarthEnv website (Domisch, Amatulli,

& Jetz, 2015) and have a 1 km resolution global

coverage. This dataset includes earth covers,

climatic and topographic variables, and soil

characteristics. Besides, we included a flow

layer average for the periods 1960-2015 (Bar-

barossa et al., 2018), which is available global-

ly at a resolution of 1 km (Digital Appendix 1).

The accessible area or mobility (M) can

condition the models’ evaluation methods and

their accuracy (Barve et al., 2011). Therefore,

we defined each species-area by making an

intersection between each species’ presence

records with a map of watersheds HydroBA-

SINS 1.0 Level 6 (Lehner & Grill, 2013). To

evaluate the range of possible invasion of P.

hypophthalmus in Colombia, we selected the

largest M within the native species (A. parda-

lis) (Digital Appendix 2). All variables were

trimmed with the M defined using the raster

package (Hijmans et al., 2015) in the program

R (R Core Team, 2014).

Comparison between the native and

the introduction niche of P. hypophthalmus:

The niche comparisons were made by follow-

ing Broennimann et al. (2012) proposal,which

consisted of getting values of niche similarity

in the environmental space using Schoener’s

D similarity metric (Warren, Glor, & Turelli,

2008). This technique uses the first two axes

of principal component analysis (PCA) that

reduce the 71 environmental variables to know

the environmental conditions occupied by P.

hypophthalmus. We calculated the native niche

stability proportion observed in the exotic niche

and the expansion of the species’ new environ-

ments in the non-native niche. Subsequently,

we tested for niche similarity by assessing

random changes of the niches within available

conditions in the study area, i.e., assessing if

the niche of the species is more similar than

expected by chance under a specific null model

(Warren et al., 2008; Warren, Glor, & Turelli,

2010). We made all analyses using available

tools in the Ecospat package (Di Cola et al.,

2017) in the software R (R Core Team, 2014).

Modeling and projection of the ecologi-

cal niche: We built the ENM for each species

using 71 environmental variables and the Max-

imum Entropy algorithm (Maxent) (Phillips,

Dudík, & Schapire, 2004; Phillips, Anderson,

& Schapire, 2006) through the ENMeval pack-

age (Muscarella et al., 2014) in the R program

(R Core Team, 2014). We selected Maxent

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because of its capability to mitigate redundant

variables’ contributions (Elith et al., 2011;

Feng, Park, Liang, Pandey, & Papes, 2019).

Also, the algorithm has shown to remain stable

both in the precision of the prediction and the

total predicted area present using different

categories of sample sizes (Phillips & Dudík,

2008; Wisz et al., 2008; West, Kumar, Brown,

Stohlgren, & Bromberg, 2016).

In the construction of the P. hypophthal-

mus model, we used the Jackknife method

(Pearson, Raxworthy, Nakamura, & Peterson,

2007) with 31 folds given the low number

of obtained records, while for native species,

we used the random kfold method, 10 000

background points by default, and the combi-

nation of feature classes (Linear, Quadratic,

Linear-Quadratic, Hinge, Linear-Quadratic-

Hinge) with ten regularization values (0.5 to

5, each 0.5). The best model for each species

was selected, bearing in mind that the lowest

ΔAIC (the best models taking into account the

smallest number of parameters, Muscarella et

al., 2014) and the logistic output were used for

all models. Finally, we converted the suitability

map to binary map using the 10P cut-off thresh-

old value (tenth percentile) for the P. hypoph-

thalmus and MTP (minimal training presence)

threshold value for the native species.

Ecological niche overlap of native spe-

cies with non-native species: We estimated

geographical overlap between the P. hypoph-

thalmus and each native species summing

their binary maps of distribution in QGIS soft-

ware (QGIS, 2020). Then, we calculated the

number of pixels corresponding to the spatial

overlap among areas of predicted distribution

of each species.

RESULTS

Data of presence: We analyzed a total of

31 records for the native area of P. hypophthal-

mus and 25 in the non-native area, 57 records

for P. magdaleniatum, 93 for S. cuspicaudus,

and 159 for A. pardalis (Digital Appendix 3,

Digital Appendix 4, Digital Appendix 5, Digi-

tal Appendix 6).

Comparison between the native niche

and introduction niche of P. hypophthalmus:

For the comparisons between the introduc-

tion and the native niche, we obtained PCA

that explains 53.36 % of the variance, with a

Schoener’s D value of 0.125, an expansion of

24 %, and stability of 76 % (Fig. 1A). On the

other hand, similarity tests show that native

and introduced range niches are more similar

to what is expected by chance (P = 0.047; Fig.

1B). Therefore, the niche of P. hypophthalmus

has significantly conserved non-native area.

Ecological niche modeling: P. hypoph-

thalmus is widely distributed in the Asian

tropical region in Laos, Thailand, Vietnam

and Cambodia (Fig. 2A). The ecological niche

model made for its natural distribution zone

gives a probability value of the area under the

curve (AUC) of 0.94, indicating that the model

had a good performance. The selected variables

provided a satisfactory prediction of the species

distribution (Table 1).

TABLE 1

Selected model for each of the species

Species Feature classes

Regularization

multiplier

AICc AUCtrain AUCtest

Number of

parameters

Cut-off

threshold

P. hypophthalmus

L 4.5 619.92 0.9541 0.9426 9 10P

P. magdaleniatum

L 2.5 1048.7 0.9445 0.9001 18 MTP

S. cuspicaudus

H 3.5 1765.6 0.9298 0.8989 24 MTP

A. pardalis

LQH 5 2842.3 0.0495 0.9389 24 MTP

L = Linear, H = Hinge, LQH = Linear-Quadratic-Hinge.

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Seven variables contributed to obtaining

the final model and predicting the geographic

areas with environmental suitability charac-

teristics for P. hypophthalmus in the basin

of Mekong and Chao Phraya rivers (Digital

Appendix 7). Among these variables, the first

one is the elevation (elevation_01) and was the

most influential factor in the model, followed

by the percentage of thick fragments in the soil

(fraction > 0.2 mm; soil_minimum_06) and

the minimum percentage of the substrate type

(R horizon or bedrock; soil_minimum_10).

Therefore, based on the response curves and

geographic representation (Digital Appendix

8), we identified that the species is located in

lower areas of the basin where the composi-

tion of the riverbed is mainly small fragments

(sands, clays, silts). Those conditions represent

a total of 19 938 km

of the basins of its native

distribution. When we projected the model to

Fig. 1. Comparisons of the native and introduced niche of P. hypophthalmus. A. Test’s PCA results proposed by

Broennimann et al. (2012). The shadow in the blue zone represents the species’ density, while the continuous and dotted red

and green contours represent 100 % and 25 % respectively of the available environment in both zones. B. Niche similarity

index histograms, the red bar the value of Schoener’s D. The histogram corresponds to the niche similarity of the non-native

niche in the Colombian region.

Fig. 2. Potential distribution of P. hypophthalmus represented in red lines. A. Its native area in Laos, Thailand, Vietnam and

Cambodia. B. Its non-native area in Colombia. The red dots represent occurrences.

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Colombia (Fig. 2B), we found that the spe-

cies can cover a potential area of 27 801 km

of the Magdalena, Cauca, Atrato, and Cata-

tumbo river basins, representing 48.26 % of

the total extension of water bodies in the area

(57 597 km

In native tree ecological niche models for

native species, all AUC test values are greater

than 0.898, indicating good performance (Table

1). The variable elevation (elevation_01) made

the most significant contribution to the three

models, placing the three species in lowland

areas. The R horizon’s minimum percentage

was the second most important variable for the

S. cuspicaudus and P. magdaleniatum models,

indicating that these species prefer areas with

low percentages of rocky material or bedrock.

Additionaly, the second most important vari-

able was the coldest month in the case of the

species A. pardalis. This indicates that higher

values fo temperature (~ 20 °C) favored the

presence of the species; however, a low per-

centage of R horizon values also influenced

the final model (Digital Appendix 9, Digital

Appendix 10, Digital Appendix 11, Digital

Appendix 12, Digital Appendix 13, Digital

Appendix 14).

Ecological niche overlap of native spe-

cies with non-native species: The geographical

overlap analysis shows that the P. hypophthal-

mus distribution area in Colombia covers 91.79

% of the suitable environmental zones of the

A. pardalis; this means that P. hypophthalmus

overlaps 17 400 km

of the native species’

distribution (Fig. 3A, Fig. 3D). In the case of

S. cuspicaudus species, we found an 82.17 %

overlap where both species overlap in 20 045

(Fig. 3B, Fig. 3E), while for P. magdaleni-

atum there is an overlap of 91.72 %, which is

equivalent to 10 342 km

(Fig. 3C, Fig. 3F).

DISCUSSION

Our results suggest a climatic niche con-

servatism in P. hypophthalmus. It shows that

this species is in similar climatic characteris-

tics in both their native and invasive ranges.

We also found that around 45 % of our study

area is susceptible to invasion. These results

have important implications because they are

the first demonstration of a possible invasion

range of P. hypophthalmus, and the risk that

they represent for other species with their

same behavior.

Environmental Niche Models can provide

a realistic proxy about the geographic range of

a fish species, and they can be used for sev-

eral purposes, including conservation action

(Valencia-Rodríguez et al., 2021). In biological

invasions, niche models and their transfers to

non-native geographical zones require caution

in their interpretation (Owens et al., 2013).

Thus, a fundamental assumption for allow-

ing geographic predictability between native

distribution areas and invaded areas is the

niche conservatism analysis of the introduced

species. According to this assumption, the

species usually conserves its ancestral niche

in the invaded regions (Wiens & Graham,

2005; Broennimann et al., 2007; Liu, Wolter,

Xian, & Jeschke, 2020). On the one hand, we

found that P. hypophthalmus shows conserva-

tism of ecological niche; for that reason, we

could transfer the calibrated ENM from the

native to the non-native area with a confidence

approximation of the possible invasion in our

study area. On the other hand, we obtained an

expansion estimation of the niche that could

indicate that P. hypophthalmus occupies new

habitats in Colombia that are not represented

in its native range. These results would imply

that the P. hypophthalmus fundamental niche

is broader than the niche represented in their

native area. By contrast the expansion detected

may also be explained by the low numbers of

records in its native zone, which would lead

to incomplete representation of the niche;

however, the records used to train the P. hypo-

phthalmus model in its native area correspond

with the known distribution area of the species

(FAO, 2010).

The selection of environmental predic-

tors is a fundamental step to produce ENMs;

if predictors are biologically informative, they

represent more realistic species-environment

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relationships (Parra, Graham, & Freile, 2004),

and thus models are projected more accurately

into novel environments. It has even been sug-

gested that elevation, slope, precipitation, soil

features, temperature, and vegetation, define

the type of aquatic ecosystems (Baron et al.,

2003). Therefore the variables selected to build

our models estimate important environmental

preferences of the four species analyzed. Alti-

tude takes on particular importance because

freshwater species are confined to water bod-

ies with specific characteristics. The eleva-

tion is a direct conditioner of characteristics,

such as flow, substrate, or vegetation (Nori &

Rojas-Soto, 2019). Notably, basins with lower

altitude values correspond with high values of

suitability predictions, which represents topo-

graphic variability that facilitate migration and

/ or prevent dispersal process within the hydro-

graphic network (Castillo-Torres, Martínez-

Meyer, Córdova-Tapia, & Zambrano, 2017).

Another important environmental variable that

explained the potential distribution of the four

species was the percentage of coarse frag-

ments in the river bed. This variable is a factor

that determines the nutritional and chemical

Fig. 3. Potential distribution of three native species represented in red lines: A. A. pardalis. B. S. cuspicaudus. C. P.

magdaleniatum. And their areas of distribution overlapping the distribution of P. hypophthalmus: D. A. pardalis. E. S.

cuspicaudus. F. P. magdaleniatum. The red color symbolizes the overlap between the native and non-native species; the

yellow color is the area for the P. hypophthalmus species, and the green color the range of the native species.

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conditions of the rivers, and it characterizes the

structure and dynamics of the aquatic habitat

(Baron et al., 2003). Followed by this, variables

related to temperature, such as the minimum

temperature of the coldest month, influenced

the models, being associated directly with the

organisms’ fitness and, by extension, deter-

mines the place where the species are distrib-

uted and how their community varies. Also,

another variable selected was the precipitation

seasonality which determines the flow pattern

of rivers, lakes, and wetlands, which influence

a large part of the habitat of freshwater species

(Baron et al., 2003).

The presence of P. hypophthalmus in

Colombia has been previously recorded from

field observations (Valderrama et al., 2016).

The individuals of P. hypophthalmus free in

natural environments may be either by acciden-

tal escape from aquaculture or direct introduc-

tions from sport fishers or aquarists (Gutiérrez

et al., 2012; Lasso et al., 2020). Consequently,

in this study, the ENM allowed the identifica-

tion of new areas suitable for the colonization

of this species. The models indicated that the

species’ niche conditions are widely repre-

sented in the Colombian inter Andean valleys.

It corresponds with low areas with sediments

and sedimentary rocks (fragments such as

sand, clay, silt, and consolidations of said

fragments, respectively; Jaramillo, 2002). The

proposed models indicate that the colonization

of P. hypophthalmus in Colombia is possible.

Its survival is confirmed by present records of

P. hypophthalmus in Magdalena-Cauca, Sinú,

Atrato, San Jorge, Cesar, and Catatumbo riv-

ers basins. The spread and colonization of

the aquatic ecosystems in Colombian basins

by the P. hypophthalmus could be possible

because the species has a high tolerance to

changes in environmental conditions, mainly

the resistance to low oxygen levels, salinity,

pH changes, temperature, and turbidity fluctua-

tions (Singh & Lakra, 2011; Faruk, Patwary, &

Hasan 2012; Garcia et al., 2018; Islam, Uddin,

Uddin, & Shahjahan et al., 2019).

Pangasianoson hypophthalmus is a spe-

cies that is easy to farm and popular among

aquaculturists due to its resistance to water

quality variations, fast growth, good survival

rates, and economically attractive market size

(Ali, Haque, & Belton, 2013; Islam et al., 2019).

Given this scenario and its invasive potential,

authorities in countries such as Mexico do not

allow or encourage the introduction and cul-

tivation in their territories (Mendoza-Alfaro,

Luna-Peña, & Arias-Gámez, 2013; Delegación

SADER Tamaulipas, 2017). Pangasianodon

hypophthalmus has a great capacity for dis-

placement due to its migratory nature and high

fecundity. How we illustrate P. hypophthalmus

presents life history characteristics similar to

the native species analyzed here (Maldonado-

Ocampo et al., 2005; Jiménez-Segura, Palacio,

& López, 2009; Jiménez-Segura, Palacio, &

Leite, 2010; Jiménez-Segura et al., 2020b).

These assessments of species’ life features can

offer rigorous scientific support for rapid clas-

sification of extinction or invasion risks and

efforts to prevent invasion because of deter-

mining which species and sources of emerging

invaders are worthy of scrutiny, more attention

to management or policies must be applied

(Liu, Comte, & Olden, 2017). Comparing the

P. hypophthalmus niche with the native species

indicates that the four species share environ-

mental conditions that spatially converge in the

evaluated basins’ lower and middle zones as

high-water temperature and substrates with a

high percentage of small fragments. (e.g. silts,

clays). P. hypophthalmus is distributed almost

entirely in the available space presented by

the three native species, even though the mod-

eled distribution range of P. hypophthalmus in

Colombia is underestimated. Its convergence

could trigger competition for space and food,

and this species can even become a predator of

juvenile and adult organisms of other species

of fish consumed by native species (Gutiérrez

et al., 2012; Raman et al., 2013). Competition

and predation would cause mortality in native

species and could even affect recruitment, and

the result of this situation would be reductions

in the genetic diversity of the native species

(Raman et al., 2013).

582

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The finding of conserved climatic niche

for invasive species has major implications

for predicting future invasion risks. Because

it permits a model transferability using niche

models. This kind of study represents a cost-

effective strategy focused on identifyinging

climatic conditions occupied in their native

range but (not yet) colonized in their intro-

duced range (Liu et al., 2020). We identify

that P hypophthalmus present a niche conser-

vatism, and for that reason we identify areas

with a high risk of being invaded. That is why

we call on environmental authorities because

it is important to develop short-term actions

to avoid this species’ dispersal. In this case,

it is necessary to control its cultivation and

use in aquariums. Although they may seem

drastic measures, they should be considered

by the Colombia’s environmental and fisheries

authorities. Posing that the legalization of the

use of the species is the only way to control its

current illegal exploitation is not an appropriate

justification since the control measures for fish

farmers and aquarists have historically been

non very effective. Eradication through hunting

and the control prohibition of its eventual cul-

tivation directly in rivers must be accompanied

by educational programs to help consumers

understand the impact non-native species have

on aquatic biodiversity, and to encourage fish

farmers to use good practices to reduce the

impact of their activities on natural aquatic

environments; also, invest more resources in

research for developing fish culture with native

species. This is an environmental decision; the

precautionary principle should always prevail,

in the absence of more robust information on

the invasive capacity of P. hypophthalmus in

Colombia’s river networks and on its real effect

on a highly endemic aquatic biodiversity.

Ethical statement: authors declare that

they all agree with this publication and made

significant contributions; that there is no con-

flict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgements

section. A signed document has been filed in

the journal archives.

ACKNOWLEDGMENTS

We thank Juan Luis Parra and Daniel

Valencia for their support, suggestions and dis-

cussions. To the Humboldt Institute, InvBasa,

Margarita Roa, Claudia Castellanos, and fish-

ermen from Magdalena river basin for facili-

tating the systematization and compilation of

the species presence in Colombia. To María

Piedad Baptiste and Jose Manuel Ochoa from

the Humboldt Institude for facilitating this col-

laboration between the University of Antioquia

and Humboldt institutions. To Wyatt Madden-

son and Carolina Soto for English grammar

support, and finally, we would like to thank

our reviewers and Octavio Rojas for improving

our manuscript.

RESUMEN

Distribución potencial en Colombia del pez

introducido Pangasianodon hypophthalmus

(Siluriformes: Pangasiidae) e implicaciones para

los peces nativos en peligro de extinción

Introducción: Una de las amenazas importantes para

los ecosistemas acuáticos es la introducción de las especies

no nativas en ambientes naturales. Esto representa una

de las principales causas de pérdida de biodiversidad en

el mundo. Pangasianodon hypophthalmus es una especie

introducida en Colombia que comparte características de

vida y hábitat con especies nativas en peligro de extin-

ción como: Ageneiosus pardalis, Sorubim cuspicaudus y

Pseudoplatystoma magdaleniatum. Sin embargo, poco se

conoce de su distribución y los efectos en la fauna nativa

han sido poco explorados. Objetivo: Evaluar el potencial

invasivo de P. hypophthalmus en algunas de las cuencas

colombianas que desembocan en el Mar Caribe. Métodos:

Utilizando registros disponibles en varias bases de datos,

realizamos un análisis de conservadurismo de nicho entre

los registros nativos e introducidos de P. hypophthalmus

usando el paquete de R Ecospat. Posteriormente, modela-

mos el área potencial de invasión de P. hypophthalmus y

las áreas de distribución de tres especies nativas realizando

modelos de nicho ecológico (MNE) utilizando el algoritmo

de Maxent. Finalmente, calculamos una superposición de

nicho geográfico entre las especies nativas y no nativas.

Resultados: El análisis espacial de Ecospat indica que

P. hypophthalmus conserva algunos atributos del nicho

583

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ambiental a través del espacio. Por esta razón, podemos

usar los MNE como una aproximación a su rango de dis-

tribución en el área invadida. Nuestros resultados usando

MNE demostraron que las cuatro especies analizadas

prefieren regiones bajas y ligeramente rocosas; por ello,

la superposición geográfica de los nichos ecológicos de

las tres especies nativas y la especie introducidas superan

el 80 %. Conclusiones: Existen condiciones adecuadas en

las cuencas de estudio para el establecimiento completo

de la especie P. hypophthalmus, lo que representa un alto

riesgo para los ecosistemas acuáticos y la ictiofauna nativa.

El conocimiento de las áreas de distribución potencial es

fundamental para implementar controles sobre la especie.

Palabras clave: modelos de nicho; agua dulce; especies

invasoras; peces migratorios.

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