Rev Fish Biol Fisheries (2010) 20:457–469

DOI 10.1007/s11160-009-9153-y

RESEARCH PAPER

Patterns in the distribution of fish assemblages in Rı́o

Amacuzac, Mexico: influence of abiotic factors and biotic
factors
Patricia Trujillo-Jiménez • Eugenia López-López •

Edmundo Dı́az-Pardo • Julio A. Camargo

Received: 6 August 2009 / Accepted: 30 November 2009 / Published online: 13 December 2009
Ó Springer Science+Business Media B.V. 2009

Abstract Patterns in spatial and seasonal distribu- the sites in physicochemical and water quality envi-
tion of fish communities were analyzed in the Rı́o ronmental gradients (conductivity, dissolved oxygen,
Amacuzac, Mexico, and their relationship to environ- orthophosphates, ammonium, pH) displaying seasonal
mental variables and habitat characteristics. The PCA variation. Fifteen species were recorded, eight of them
biplot of the study sites and environmental factors are exotic. One new record appeared for the Rı́o
showed the first two axes accounting for 52.93% of the Amacuzac: Pterygoplichthys disjunctivus is exotic.
variance. The diagram showed the study sites ordina- Study sites with highest species richness were: 5, 7 and
tion in environmental gradients. The first axis 9 (twelve species each one); while the study sites with
explained variables related to habitat characteristics low species richness were 1, 2, 3 and 6 (eight species
and temperature (36.30%) and second axis arranged each one). Six of the species were distributed
throughout the whole river. Based on the composition
of the fish fauna, the study sites form two groups and
the analysis of fish species displays three groups
according to the Bray–Curtis index. The diagram of
P. Trujillo-Jiménez (&) the canonical correspondence analysis relates envi-
Centro de Investigaciones Biológicas, Universidad ronmental parameters to the abundance of fish species
Autónoma del Estado de Morelos, Av. Universidad 1001,
and showed that the first two axes exhibit 78.31% of
Col. Chamilpa, CP 62209 Cuernavaca, Morelos, Mexico
e-mail: trujill@uaem.mx the explained variance. Species richness had a spatial
pattern associated to the introduction of exotic species
E. López-López for ornamental uses. According to the results of the
Laboratorio de Ictiologı́a y Limnologı́a, Escuela Nacional
importance value index (IVI), the dominant species
de Ciencias Biológicas, IPN, Prol. de Carpio y Plan de
Ayala, Col. Sto, Tomás, DF 11340, Mexico were the poecilids Poeciliopsis gracilis and Heteran-
dria bimaculata, small fishes that were introduced in
E. Dı́az-Pardo the river. Rı́o Amacuzac has a biotic alteration in the
Facultad de Ciencias Naturales-Biologı́a, Universidad
structure of fish communities due to the invasion of
Autónoma de Querétaro, Av. de las Ciencias s/n,
Juriquilla, Del. Sta. Rosa Jauregui, CP 76230 Santiago de exotic species, representing risks to the integrity of the
Querétaro, Querétaro, Mexico native fish fauna.

J. A. Camargo
Keywords Fish assemblages

Departamento de Ecologı́a, Facultad de Biologı́a,
Universidad de Alcalá, Ctra. Madrid-Barcelona km. 33,6, Distributional patterns
Native species

28871 Alcalá de Henares, Madrid, Spain Exotic species
Rı́o Amacuzac
Mexico

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458 Rev Fish Biol Fisheries (2010) 20:457–469

Introduction Habit et al. 2007). The above environmental vari-

ables, are evidence of the dynamics of ecosystems,
Biological communities change in time and space, as a however, they have little to do with those that humans
result of different habitat characteristics, resource have deliberately or accidentally caused in their
availability, among other (Matthews and Robinson efforts to modify nature for their own benefit. In
1998; Grenouillet et al. 2002). In this respect the study addition to the impacts caused by industrial and urban
of lotic ecosystems deals with the understanding of activities, the introduction of exotic species stands
mechanisms and processes that are responsible for the out because of their permanence and difficult control
differences or similarities between communities and (Contreras-MacBeath et al. 1998). Exotic fish species
their relationship to environmental variables (Torres are widespread in Mexico; over the past century 113
et al. 2006). Longitudinal zonation is one of the most species of freshwater fishes have been introduced to
frequently described distribution patterns for fresh- Mexico (Contreras-Balderas et al. 2008). Most spe-
water fish fauna in river systems (Schlosser 1982). cies were introduced for aquaculture, recreational and
Longitudinal distribution is the result of changes in ornamental purposes, and are the most widely
species richness, abundance, diversity and assem- distributed and are dominant species in many natural
blages composition associated with changes in envi- ecosystems of Mexico. Understanding how distribu-
ronmental spanning from headwater to the river mouth tion of exotic species related to distributional patterns
(Granado 2000; Ostrand and Wilde 2002). of native species is important for conservation and
Ostrand and Wilde (2002) suggested that the management effort (Habit et al. 2007).
assemblage structure is determined more by average In the present study, spatial and temporal distri-
or persistent differences in environmental conditions butional patterns of fish assemblages are examined
among sites, than by seasonal variation in environ- and their relationship with environmental factors in
mental conditions. This is due to the fact that rivers are the Rı́o Amacuzac in Morelos, Mexico.
ecosystems with wide variations between seasons.
During the rainy season, the velocity and flow of the Study area
current increases drastically throughout the whole
system, leading to variation in some physical condi- The Rı́o Amacuzac belongs to the Balsas basin drainage
tions such as temperature, pH and conductivity, among which is one of the largest rivers of southern Mexico that
others, and making more difficult to determine whether flows into the Pacific Ocean (SPP 1988; CNA 1998).
there are changes in distributional patterns (Callow and The most important river within the state of Morelos is
Petts 1994; Rondon and Petrere 2007; Orrego et al. the Amacuzac with 104 km in length. It has its origins in
2009). the Rı́o Chantacotlan and Rı́o San Jeronimo which
Understanding how environmental variables (and emerge from Las Grutas de Cacahuamilpa. The Rı́o
their spatial and temporal variations) shape fish Amacuzac runs through Morelos from the west and
community structure is an important issue for envi- collects the streams of the Rı́os Chalma, Tembembe,
ronmental managers. Numerous studies have tested Apatlaco, Tetlama, Yautepec and Cuautla, and other of
how environmental changes influence fish assem- minor importance (Fig. 1). The climatic regime is hot
blages, mainly in the USA (Gido et al. 1997; Brown subhumid with summer rains (June to October) and dry
2000; Ostrand and Wilde 2002; Quinn and Kwak season (November to May). Mean annual temperature is
2003; Adams et al. 2004; Daniels et al. 2005). Among 21.5°C and mean annual rainfall is 72.2 mm (SMN
comparable studies in Mexico and other tropical 2009). According to the criterion of Strahler (Welcom-
countries, are the studies of Paulo-Maya and Ram- me 1985) it is a third order river. The surrounding
ı́rez-Enciso (1997), Habit et al. (2003), Rondon and vegetation includes tropical dry forest (Aguilar 1999).
Petrere (2007), Fialho et al. (2008), Araújo et al. However, in some slopes of the Rı́o Amacuzac are
(2009) and López-López et al. (2009). isolated patches of semi-deciduous forest. The vegeta-
Environmental variables such as elevational gra- tion on the banks of the river has been modified and
dient, river order and habitat characteristics and removed much of the course for agricultural use,
complexity, have been correlated with distribution preserved in the canyons and ravines of the Rı́o
patterns of fish assemblages (Arunachalam 2000; Amacuzac (Dos Bocas to Huajintlán).

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Rev Fish Biol Fisheries (2010) 20:457–469 459

Methods conductivity meter Conductronic-CL8. To evaluate

water quality, a water sample was taken in a polyeth-
Sampling regime ylene 1,500 ml flask. To quantify total hardness (mg/l)
and carbon dioxide (mg/l), titration techniques pro-
Nine sites were sampled (locality 1 the point of highest posed by APHA (1985) were used. Ammonium (mg/l)
altitude 1,048 masl and 9 the lowest altitude and orthophosphates (mg/l) concentrations were deter-
777 masl), on an 80 km stretch along the Rı́o Amac- mined, using a Hach DR-EL/2000 spectrophotometer.
uzac (Fig. 1). Samplings for nine sites were conducted To characterize the habitat characteristics the follow-
in the 2 years seasons: dry season (November, ing variables were recorded: distance from upper
December 1999; January, March, April and may portion to the lower portion river in km, altitude above
2000) and rainy season (August; September; October sea level (masl), using a geopositioner (GPS 12XL
1999; June and July 2000). Garmin) and the substrate, was divided into four types:
Different environmental factors were recorded at 1 = rocky; 2 = muddy/stony; 3 = sandy/muddy and
each site using YSI 57 sonde: water temperature (°C), 4 = sandy/rocky.
dissolved oxygen (mg/l); pH using a pH-meter Fish were collected with seine net 5 m long and
HANNA, and conductivity (lS/cm) using a 2 m deep, with 1 cm mesh size. A one hour capture

Fig. 1 Location of study

sites in Rı́o Amacuzac,
Morelos, Mexico. The
number indicate the study
sites: 1 Dos Bocas;
2 Contlalco; 3 Huajintlán;
4 Amacuzac; 5 El
Estudiante; 6 Vicente
Aranda; 7 Los Lagartos;
8 Las Granjas; 9 Las
Huertas

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effort was carried out in each locality; covering about Data were transformed to log (x ? 1 and square
100 m, in order to provide a representative sample of root) for ensuring data normality, changing the
fish fauna (Matthews 1990). All depths of the river weights of different species or variables, and remov-
covered and pools as well as riffles and areas with ing the effect of measurement units.
vegetation were considered. Specimens were fixed in
10% formaldehyde and transported to the laboratory
for taxonomic determination with the aid specialized Results
keys: Meek (1904), Álvarez (1970), Schönhuth and
Doadrio (2003), Miller et al. (2005) and Schmitter- Habitat characterization
Soto (2007). All specimens sampled were deposited
in the Collection aquatic flora and fauna of Centro de The PCA biplot of the study sites and environmental
Investigaciones Biológicas de la Universidad Autó- factors showed the first two axes accounting for
noma del Estado de Morelos (UAEM), identified by 52.93% of the variance (Fig. 2). The diagram showed
the acronym CAFF (New Creation). the study sites ordination in environmental gradients.
The first axis explained variables related to the
Data analysis habitat characteristics and temperature; i.e. vector on
the far right of the diagram: altitude. Along the same
Habitat characterization and assessment of the impor- axis but to the left, are the vectors for distance,
tance of the different environmental variables were hardness, substrate and temperature (Fig. 2a). Study
analyzed with principal components analysis (PCA) sites belonging to the upper portion of the river were
using a matrix, which includes physicochemical related to the altitude vector on the right side of the
parameter (temperature, dissolved oxygen, pH and diagram; on the left are the sites of the lower portion
conductivity), water quality variables (ammonium, of the river (from locality 5 to 9), showed a lower
orthophosphate, hardness) and habitat characteristics altitude, higher temperature and a rocky/sandy and
(distance from upper portion to the lower portion river sandy/muddy substrate (Fig. 2b).
in km, altitude and substrate type). Data were trans- The second axis arranged the sites in physicochem-
formed to log (x ? 1) with the exception of pH. ical and water quality environmental gradients
In order to identify areas along the watercourse with (conductivity, dissolved oxygen, orthophosphates,
characteristic fish communities and to compare the ammonium, pH) displaying seasonal variation. In the
fish community between study sites, we used Bray– high part of the diagram, study sites were clustered and
Curtis similarity index. The study matrix included the belongs to the months of the rainy season (June, July
presence/absence of all species in each site and study and October), when the values of orthophosphates and
month. Species dominance was assessed by impor- conductivity were greater. During this period
tance value index (IVI) as proposed by Krebs (1989), upstream study sites had higher values of dissolved
using the frequency of occurrence and relative abun- oxygen. In the same axis, in the low part, study sites
dance of species. were clustered corresponding to the dry months
Canonical correspondence analysis was used to (January, March, April and November), characterized
evaluate the relationship between fish species, envi- by having more alkaline water (Fig. 2b).
ronmental factors and habitat characteristics. For this, a
matrix of relative abundance of species by site and by Richness
study month was used, transforming values to square
root. The environmental data matrix included physi- A total of 15 fish taxa, representing six orders and
cochemical variables (temperature, dissolved oxygen, eight families were collected. The order Cyprinodont-
pH conductivity), water quality variables (ammonium, iformes had the highest diversity, with two families
orthophosphates, hardness and carbon dioxide), and and six species, followed by the order Siluriformes,
habitat characteristics variables (substrate type, dis- with two families and two species. The Poeciliidae
tance and altitude); all variables were transformed to were the most diverse family, represented by five
log (x ? 1). All statistical procedures were performed species. The second most diverse family was the
with the statistical package XLSTAT (2009). Cichlidae, with four species. The remaining families

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Fig. 2 PCA biplot of (Axes F1 and F2: 52.93 %)

Amacuzac River study sites. (a) 1
a Vectors of the variables
considered (environmental Orthophosphate
factors, water quality and
habitat structure).
b Ordination of study sites. Conductivity
0.5
(Number indicate the study
site (Code of abbreviations
is in Fig. 1) and letter Diss Ox
Altitude
F2 (16.63 %)
month: J January, M March, Ammonium
Ap April, My May, Jn June,
Jl July, A August, S 0 Substrate
Distancie Km
September, O October, N
Temperature
November, D December).
Hardness
Diss Ox dissolved oxygen

-0.5

pH

-1
-1 -0.5 0 0.5 1
F1 (36.30 %)

(Axes F1 and F2: 52.93 %)

(b) 8

6Jl
6

5Jl
8Jl
4

5D 3O
2 8S 7Jn 7D 4Jn 3Jn 3Jl 2Jl 1O
4O 4D 2Jn 2O 1Jl 1D
F2 (16.63 %)

8My
8Jn 9O 7S 5Jn 5O 3My
6My 9Jn 7Jl 4S 2My 3D 1M 1Jn
1J
0 7A 8O 6O 3A4J 2A3J 2J 1My1S
8M 8A 6A 6Jn 9D 6S 6D 5S 5A 4Jl 2S 1A
6J 7J 4A3S 3M 2M
9Jl 9A 6M 5My 9J 8J5J 4M 2Ap 3Ap 1Ap
9M 9Ap 9S 6Ap 7Ap 5M 2N 1N
6N 4N 3N
-2 8Ap 7My 7M 5Ap 4My 7N 5N
8N
9N
-4

-6

-8
-8 -6 -4 -2 0 2 4 6 8
F1 (36.30 %)

had one species each one. One new record appeared Balsas river basin (20%), four were native (26.66%)
for the Rı́o Amacuzac basin; of the family Loricarii- and eight were exotic (53.33%) (Table 1).
dae, formed by the Pterygoplichthys disjunctivus Study sites with highest species richness were: 5, 7
species, also known as plecostoma or Orinoco sailfin and 9 (twelve species each one); while the study sites
catfish. Three of the species were endemic of the with low species richness were 1, 2, 3 and 6 (eight

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species each one). Six of the species were distributed sampled in two sites in the lower portion of the river,
throughout the whole river. The two most important where both share site seven (Fig. 4).
species for fishing, are also native to the basin (mojarra
criolla Cichlasoma istlanum and bagre del balsas Relationship between species physicochemical
Ictalurus balsanus) and were recorded in six and four characteristics and habitat characteristics
localities respectively; nevertheless, it is worth noting
that both species showed low abundance; the rest of The Canonical Correspondence Analysis (CCA)
the species were collected only at one or two localities, diagram, which relates environmental parameters to
also with low abundances (Table 2). the abundance of fish species, showed that the first
The months of September and July were the ones two axes exhibit 78.31% of the explained variance.
that showed the greatest richness of species (13 and Permutation analysis (n = 500) rejected the alterna-
14, respectively), while December and March had a tive hypothesis of no linear relation between sites/
lower number of species (Table 2). According to the species data and sites/variables. Thus, the null
results of the IVI, Poeciliopsis gracilis and Astyanax hypothesis (a linear relation between sites/species
aeneus were the most common and abundant species data and sites/variables data) is accepted at the
in the river, while X. helleri, Aequidens rivulatus and P \ 0.0001 significance level.
Poecilia reticulata were the less common species and In the diagram (Fig. 5) we observed that the study
with low abundance (Table 2). sites were dispersed along component 1, in an altitude
gradient, distance and substrate heterogeneity. Study
Stream zonation by their fish fauna sites with greatest altitude, less distance and rocky
substrates, as well as muddy/stony substrate, are at
Based on the composition of the fish fauna and the extreme left of the diagram, and the ones with
according to the Bray–Curtis analysis, the study sites lower altitude and rocky/sandy and sandy/muddy
form two groups, with a similarity cut-off value of substrates, are at the extreme right (Fig. 5a).
0.7 (Fig. 3). Group I is formed by study sites 2, 1 and In the second component we find a gradient of
4, and they share nine species. Group II is formed by environmental factors with seasonal changes. The dry
the sites of the lower portion of the river. This group season in the lower portion of the diagram, is
is characterized by the presence of four species that characterized by having greater values for ammo-
were not collected in the upper portion of the river. nium, hardness and more alkaline water, while the
rainy season has greater values for temperature,
Fish assemblages carbon dioxide, dissolved oxygen, orthophosphates
and conductivity (Fig. 5b). Three groups of species
Analysis of fish species with the Bray–Curtis index, became evident by their location along this environ-
with the cut made at 1.1 similarity, showed displays mental gradient. In the first group is Astyanax aeneus
three groups (Fig. 4). Group I is an assemblage of and N. moralesi, species that are distributed all along
seven species that are distributed all along the Rı́o the river, but are in greater abundance in the sites
Amacuzac. Group II consists of the four-species with greater altitude and rocky and muddy/stony
assemblages formed by Atherinella balsana and substrates. Second group is formed by H. bimaculata,
Poecilia reticulata that were distributed in one or Poecilia sphenops, Ictalurus balsanus, Ilyodon
two localities of the lower portion of the river, and by whitei, Amatitlania nigrofasciata, Pterygoplichthys
C. istlanum and N. moralesi that were distributed in disjunctivus and X. helleri, species that are related to
localities that were representative of all the sub-basin sites at lower altitude and with rocky/sandy, sandy/
of the Amacuzac. Group III is comprised by Ictalurus muddy and sandy/rocky substrates, as well as with
balsanus and X. helleri. The first species was distrib- high conductivity and orthophosphate values. Third
uted in four sites from the upper portion of the river to group was constituted by C. istlanum, Atherinella
the lower portion, and the second species was only balsana, Poeciliopsis gracilis, Poecilia reticulata,
collected in site 5, where it coexists with Ictalurus Aequidens rivulatus and O. mossambicus, are related
balsanus. It was also represented by Pterygoplichthys to sites that are characterized by high values with
disjunctivus and Aequidens rivulatus, each were respect to ammonium and hardness (Fig. 5b).

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 Table 1 List of the Amacuzac River fish fauna
Family Species Natural distribution Status

Cypriniformes
Cyprinidae Notropis moralesi (N) de Buen, 195 Papaloapan River, Mexico Threatened (NOM-ECOL-059 2001)
Siluriformes
Ictaluridae Ictalurus balsanus (E) (Jordan & Snyder, 1989) Balsas River, Mexico Vulnerable (Jelks et al. 2008)
a
Loricariidae Pterygoplichthys disjunctivus (I) (Weber, 1991) South America Abundant (Mendoza et al. 2009;
Stabridis et al. 2009)
Cyprinodontiformes
Goodeidae Ilyodon whitei (Meek, 1904) (E) Balsas River, Mexico Vulnerable (Jelks et al. 2008)
Poeciliidae Heterandria bimaculata (I) (Heckel, 1848) Gulf of Mexico from Veracruz Abundant (Contreras-MacBeath
Rev Fish Biol Fisheries (2010) 20:457–469

to the north of Belize et al. 1998)

Poecilia reticulata (I) Peters, 1859 South America Rare (Contreras-MacBeath et al. 1998)
Poecilia sphenops (N) Valenciennes, 1846 Mexico to Colombia Abundant (Contreras-MacBeath
et al. 1998)
Poeciliopsis gracilis (I) (Heckel, 1848) Central America Abundant (Contreras-MacBeath
et al. 1998)
Xiphophorus helleri (I) (Heckel, 1848) Gulf of Mexico from Veracruz Abundant (Contreras-MacBeath
to the north of Belize et al. 1998)
Characiformes
Characidae Astyanax aeneus (N) (Günther, 1860) Grijalva-Usumacinta, Yucatán, Abundant (López-López et al. 2009)
Papaloapan-Coatzacoalcos,
Cuenca del Balsas
Atheriniformes
Atherinopsidae Atherinella balsana (E) (Meek, 1902) Balsas River, Mexico Rare (Contreras-MacBeath et al. 1998)
Perciformes
Cichlidae Cichlasoma istlanum (N) (Jordan & Snyder, 1989) Armerı́a-Coahuayana, Balsas Vulnerable (Jelks et al. 2008)
Amatitlania nigrofasciata (I) (Günther, 1866) Central America Abundant (Contreras-MacBeath
et al. 1998)
Aequidens rivulatus (I) (Günther, 1860) South America Common (Contreras-MacBeath
et al. 1998)
Oreochromis mossambicus (I) (Peters, 1852) Africa Abundant (Contreras-MacBeath
et al. 1998)
(N) Native; (E) Endemic; (I) Introduced
a
New record
463

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Table 2 Spatial and seasonal distribution and importance value index (IVI) (Krebs 1989) of Amacuzac River fish species (** native
or endemic)

1 2 3 4 5 6 7 8 9 S O N D J M Ap My Jn Jl A IVI
X. helleri 0.01
A. rivulatus 0.01
P. disjuntivus 0.01
I balsanus ** 0.03
A balsana ** 0.04
P. reticulata 0.09
C. istlanum ** 0.14
O. mossambicus 0.80
N. moralesi ** 1.33
I. whitei ** 2.97
P. sphenops ** 6.26
A. nigrofasciata 5.99
H. bimaculata 7.08
A. aeneu ** 14.32
P. gracilis 60.92
Number of species 8 8 8 10 12 8 12 9 12 13 10 10 9 10 9 12 12 12 14 12
Native or endemic 5 5 4 6 6 4 5 5 7 7 5 5 5 5 4 7 7 6 7 6
Exotic 3 4 4 4 6 4 7 4 5 6 5 5 4 5 5 5 5 6 7 6

1.2 3

1 2.5
Bray-Curtis Index

Bray-Curtis Index

0.8
2

0.6
1.5

0.4
1

0.2
0.5
0
2 1 4 6 8 7 9 3 5
0
Study Sites
Aaene
Pgrac
Omoss

Anigr
Hbima
Psphe

Preti

Nmora
Ibals

Pdisj
Arivu
Iwhit

Abal

Cistl

Xhell

Fig. 3 Cluster diagram of Amacuzac River sites, based on

presence/absence of fish using Bray–Curtis index of similarity. Species
The dotted line represents the cut at 0.7 similarity. (Code of
abbreviations of study sites is in Fig. 1) Fig. 4 Cluster diagram of fish assemblages defined by the
Bray–Curtis index of similarity. The dotted line represents the
cut at 1.1 similarity. Aaene, Astyanax aeneus; Pgrac, Poecil-
Discussion iopsis gracilis; Omoss, O. mossambicus, Iwhit. Ilyodon whitei,
Anigr, Amatitlania nigrofasciata, Hbima, H. bimaculata;
Psphe, Poecilia shenops; Abals, Atherinella balsana; Preti,
The longitudinal zonation of the river, as well as the
Poecilia reticulata; Cistl, Cichlasoma istlanum; Nmore,
analysis of principal components analysis, allows N. moralesi; Ibals, Ictalurus balsanus; Xhell, X. helleri; Pdisj,
to point out that the Rı́o Amacuzac has two main Pterygoplichthys disjunctivus; Arivu, Aequidens rivulatus

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Fig. 5 CCA biplot. a Sites (Axes F1 and F2: 78.31 %)

and environmental (a) 1
variables, b Environmental
variables and fish species.
(Code of abbreviations of 4Jl 6A 5D 6Jl
study sites is in Fig. 1 and Orthophosphate
code of abbreviations of 4A 5A 5Jn 6O
0.5 Diss Ox
3Jl 4Jn 4O 6Ap 6My 6N
species is in Fig. 4) 1O 3O 5O
5Ap 5S 6D 8S 5Jl
6S
1A 3A 4J 6Jn
4N 5M 4D 5N
4My 6J

F2 (17.90 %)
Altitude 3D 4S 5J6M 8O Dioxide of carbon
1Jn 2Jl
3Jn 4M
Conductivity 8J 7Jl Substrate
3Ap 3J 7D 5My 8Jl
0 Temperature
1D 2A pH 8M 8A 9O Distancie Km
2O 3N 7S
1N 1Jl 7A Ammonium 8N 8Ap
1Ap 1S 2My 3My Hardness
8My 8Jn
1My 1M 1J 2N 2J 3S
7N 7M 9S
2M 2S 3M 7Jn 9Jl 9D 9J
7J 9A 9Ap
-0.5 2Ap 2Jn 7Ap
9M 9N
7My 9Jn

-1
-2 -1.5 -1 -0.5 0 0.5 1 1.5 2
F1 (60.41 %)

(Axes F1 and F2: 78.31 %)

(b) 1.5
Xhell

1
Pdisj

Anigr
F2 (17.90 %)

0.5 Diss Ox
Orthophosphate
Iwhit
Altitude Ibals Psphe Hbima
Aaene Dioxide of carbon
Conductivity Substrate
Nmora
0 Temperature
pH Ammonium D is tancie Km
Hardness
Pgrac
-0.5 Omoss
Cistl

Arivu
Preti

-1 Abals
-2.5 -2 -1.5 -1 -0.5 0 0.5 1 1.5 2
F1 (60.41 %)

regions. The first or upper portion, with a maximum upper and lower portions of the river is high, creates a
altitude of 1,048 masl with shallow, cold, alkaline zone with rapids; this characteristic together with the
and well oxygenated waters, with rocky and muddy/ substrate type and temperature, were the factors most
stony substrate. The second or lower portion is related to spatial changes of the fish community in
characterized by being between 871 and 777 masl, Rı́o Amacuzac.
with higher values of depth, temperature, conductiv- In general, the structure of fish assemblages allows
ity and orthophosphate and lower values of dissolved the prediction of changes occurring from the head
oxygen, transparency, with rocky/sandy and sandy/ waters (upstream) to the plains (downstream) of the
muddy substrates. The altitudinal change between the ecosystem, recording an increase in the number of

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fish species in the downstream of the river. This The dominance of some species is a dynamic
pattern is frequently attributed to a general increase in process; temporal variations of biotic and abiotic
the size and heterogeneity of the habitat, together conditions lead to other species becoming dominant,
with a decrease in speed of current, allows for an allowing the species with similar ecological require-
increase in the variety of habitats, abundance and ments to coexist (Lowe-McConnell 1987). In the
diversity of food and refuge, and also to the fact that present study, and according to the results of the IVI,
some species reproduce in this part of the ecosystem the dominant species were the poecilids Poeciliopsis
(Matthews 1998; Habit et al. 2003). gracilis and Heterandria bimaculata, small fishes that
In the Rı́o Amacuzac there is a spatial pattern, were introduced in the Rı́o Amacuzac. Authors like
since the richness of species increased downstream, Johnson and Hubbs (1989), Meffe and Snelson
however, this increase in the richness of the upper (1989), stated that members of the Poeciliidae family
portion, compared to the lower, is not natural, since occupy a large variety of tropical and temperate
only the native species are uniformly distributed habitats, in freshwater and brackish water, and have a
along the whole river. The lower portion increase in great capacity for colonization, high genetic plastic-
species richness is associated with anthropogenic ity, are viviparous and have a broad trophic spectrum,
impacts by the introduction of exotic species with makes them a highly successful group.
ornamental uses, as is the case of X. helleri and This is what occurs in the Rı́o Amacuzac with
Pterygoplichthys disjunctivus. P. gracilis, is an ecologically tolerant species, living
The presence of Atherinella balsana in the lower in waters with currents as well as in pools, in clean to
portion of the Rı́o Amacuzac, is possibly due to a turbid or very muddy waters (Meffe and Snelson
migratory event, since was reported in the Rı́o 1989; Miller et al. 2005). In this study it was detected
Amacuzac by Huerta and Castañeda (1987) (In its dominance throughout the whole river, as well as
Contreras-MacBeath et al. 1998); however, Contre- that of H. bimaculata (is dominant in a locality in the
ras-MacBeath et al. (1998) mentioned that due to the upper portion and in two of the lower portion).
absence of specimens of this species in this river Another dominant species of the Rı́o Amacuzac was
during the 10 years of sampling, as well as the lack of Amatitlania nigrofasciata (dominant in the down-
preserved specimens in the scientific collections, stream), an exotic and very tolerant species that was
there is doubt the validity of the previous records. liberated into the Rı́o Chalma, is a tributary of the
Nevertheless, in the present study, Atherinella bal- Amacuzac; this was recorded as being a dominant
sana specimens were collected in localities eight and species in the aforementioned rivers, as well as in the
nine of the Rı́o Amacuzac, with a higher temperature, Rı́o Tembembe, another tributary of the Amacuzac
more alkaline waters and a decrease in dissolved (Contreras-MacBeath 1991). Among the dominant
oxygen. These are characteristics that are apparently native species, this Astyanax aeneus, it is distributed
preferred by this species, as was reported by Paulo- along the Rı́o Amacuzac and was dominant in the
Maya and Ramı́rez-Enciso (1997) in the Rı́o Cutza- upper portion where the main substrate is rocky,
mala in Michoacán, Mexico. abundant currents, colder waters and with well
The addition of species to the lower portion of the oxygenated. Its dominance is possibly due to the fact
Rı́o Amacuzac, by populations established in the that it is a species a fast swimming and with an
tributaries, is possibly by drift transportation during evident ability to adapt to different environments.
the rainy season, or were displaced towards the main López-López et al. (2009) reported that Astyanax
current by a colonization mechanism. This could have aeneus in the Rio Champotón, Campeche, México is
happened with Poecilia reticulata, since this species is common, abundant and permanent river species and
sampled to be widely distributed in two of the main is characteristic the freshwater region.
tributaries of the Rı́os Amacuzac, Yautepec and Currently, the dispersion of exotic species is
Apatlaco (Contreras-MacBeath et al. 1998), converge considered to be one of the most serious conservation
in the Amacuzac in study site seven where the species problems of ecosystems (Ruiz et al. 2000). Rahel
was collected. Also, X. helleri was collected during (2000) points out that the adverse effects of the exotic
2 months of the rainy season in locality five, where a species may be manifested in diverse ways: when
tributary (Rı́o Chalma) joins the Amacuzac. these have reached a broad distribution, they generate

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uniformity in the distribution patterns—‘‘global threatened according to the protection criteria set by
homogenization’’—modifying what was once the Mexican legislation NOM-059-ECOL (2001).
distinctive biota of each region (Sousa et al. 2008). In spite of the fact that the Rı́o Amacuzac is
They also affect native communities through inter- considered by the National Commission for Knowl-
specific relations with the indigenous biota. In the edge and Use of Biodiversity (CONABIO) as the Rı́o
same way, certain exotic species become invasive Amacuzac—Lagunas de Zempoala Prioritary Hydro-
when they displace the native ones by direct compe- logical Region RTP-67 (Arriaga-Cabrera et al. 1998),
tition, depredation, transmission of diseases, modifi- and is part of the Huautla Sierra Biosphere Reserve
cation of the biogeochemical cycles, and they even (Taxco-Huautla Sierras Prioritary Terrestrial Region
affect the structure of the trophic system, in conse- RTP 120) (Arriaga-Cabrera et al. 2000), and it
quence they are capable of generating adverse effects belongs to the Importance Area for the Conservation
in the invaded habitat (Davis and Thompson 2000; of Birds AICA C-21 (Benı́tez et al. 1999), this
Mack et al. 2000; Sax et al. 2002). The invading ecosystem shows an increase in the dominance of
species have been identified as a second cause of exotic species, which is a symptom of stress in the
biodiversity loss, at a global level (Vitousek et al. aquatic ecosystems (Rapport 1992).
1996; Leung et al. 2002). Eradication of exotic species from the Rı́o Amac-
Meek (1904) recorded that native fish community uzac is perhaps no longer possible, given their wide
in the Rı́o Amacuzac sub-basin was constituted by distribution and dominance; however, measures
eight species, of them only Poeciliopsis balsas should be sought to control the populations of these
(endemic of the Balsas basin) was not collected in exotic species, and to avoid the introduction of
the present study. Like this, we recorded that there is others, since if this is not done, the native fishes that,
still not a serious loss of species richness in this has already been damaged, could be declined more
ecosystem, in spite of the large increase in the drastically. An option for exotic species management
number of fish species introduced as the result of could be what has been implemented in the state of
human activities. However, the structure of the Michoacan (Mexico), at the Infiernillo Dam reservoir,
community has been affected and we observed a where technologies are being developed to capture
remarkable dominance of the exotic species, record- and industrialize the devil fish. For this, it has been
ing that only three of the eight exotic species reached determined that this species contains a high nutri-
a dominance of 73.99%. This has produced a strong tional value and it may be commercialized as a
ecological impact in the native species, as records protein concentrate to enrich foods. Also, subprod-
show a strong displacement by Amatitlania nigrofas- ucts may be generated to industrialize the organic
ciata, of the two most exploited species in fishing residues of these species, to be used as protein
activities: the native ‘‘mojarra criolla’’ C. istlanum products in the feed of pigs and cows; this would
and the bagre del balsas Ictalurus balsanus, both generate an important reduction in the costs produc-
contributing significantly to the diet of the inhabitants tion (Martı́nez-Palacios 2007; Stabridis et al. 2009).
of the areas adjacent the river. Contreras-MacBeath Another strategy could be to carry out environ-
et al. (1998) according to research done in 1990, mental education campaigns for the general public
which aimed at evaluating the distribution and and mainly for the Association of Ornamental Fish
ecological impact of Amatitlania nigrofasciata rivers Producers in the state of Morelos, where emphasis
in the State of Morelos, this species was found to be should be placed on the danger involved in putting
dominant in the Rivers Amacuzac, Chalma and exotic fish in the natural ecosystems, as well as
tembembe and accounted for approximately 50% of setting forth preventive and remediation measures for
the total fish community as based upon biomass and this fish pest in Morelos.
abundance. The survey also showed a clear displace-
ment of the native C. istlanum and the complete Acknowledgments This project was financed by the National
Commission for Knowledge and Use of Biodiversity
absence of Ictalurus balsanus. According to Jelks
(CONABIO–Project S150). We express our gratitude to Joel
et al. 2008 these species and Ilyodon whitei are Paulo Maya and Topiltzin Contreras-MacBeath for his
vulnerable, while N. moralesi is considered whit comments and suggestions.

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