RESEARCH PAPER

A delineation of Nuclear Middle America biogeographical

provinces based on river basin faunistic similarities
Wilfredo A. Matamoros

Brian R. Kreiser

Jacob F. Schaefer
Received: 28 February 2011 / Accepted: 3 August 2011
Springer Science+Business Media B.V. 2011
Abstract The biogeographical patterns of the obli-
gate freshwater shes of Nuclear Middle America, a
region that expands from southern Guatemala to
northern Nicaragua, are described herein. Historically,
three broad ichthyological provinces have been
assigned to Nuclear Middle America: the Usumacinta,
and the San Juan in the Atlantic slope and the Chiapas-
Nicaraguense in the Pacic slope. With the use of
correspondence analysis and unweighted pair group
method with arithmetic mean cluster analysis of a
presence/absence matrix of 76 obligate freshwater
shes, we identied four ichthyological provinces in
Nuclear Middle America: (1) the Honduras and Guate-
mala Caribbean Highlands Province, (2) the Honduras
and Nicaragua Mosquitia Province, (3) the Chiapas-El
Salvador-Nacaome Province, and (4) the Choluteca and
Nicaragua Pacic Province. Differences between prov-
inces in species composition and species turnover
between provinces were tested by analysis of similarity,
the calculation of beta-diversity indices and an indicator
species analysis. We then further characterized each
province by identifying the number of endemics and
classifying species according to their salinity tolerance.
The most striking patterns of Nuclear Middle America
freshwater sh distribution are its paucity of primary
freshwater shes and limited numbers of endemics. The
four ichthyological provinces are distinct as indicatedby
the ANOSIM and beta-diversity analysis, although one
province showed low beta-diversity values. These
results suggest that, despite of the active geological
history that characterized the region, there has been
limited isolation of species in any given province, and
historical drainage connectivity has been high.
Keywords ANOSIM Beta-diversity
Central America Ichthyological provinces
Indicator species analysis Obligate freshwater shes
Introduction
Central American freshwaters were rst divided into
ichtyological provinces (FIPs) by Miller (1966) and
then Bussing (1976). They identied the following
four FIPs in Central America (1) the Chiapas-
Nicaraguense Province (Pacic slope) extending
from the Tehuantepec River in southern Mexico
south to the Nicoya Peninsula in western Costa Rica;
(2) the Usumacinta Province (Atlantic slope), which
covers the area from the Papaloapan River in
W. A. Matamoros (&)
Louisiana State University/Museum of Natural Science,
119 Foster Hall, Baton Rouge, LA 70803-3216, USA
e-mail: wmatamoros@lsu.edu
W. A. Matamoros
Departamento de Investigacion, Universidad Pedagogica
Nacional Francisco Morazan, Tegucigalpa, Honduras
B. R. Kreiser J. F. Schaefer
Department of Biological Sciences, The University
of Southern Mississippi, 118 College Drive,
Box 5018, Hattiesburg, MS 39406, USA
1 3
Rev Fish Biol Fisheries
DOI 10.1007/s11160-011-9232-8
Author's personal copy
southeastern Mexico to north of the San Juan River in
Nicaragua; (3) the San Juan Province (Atlantic
slope), that includes the Nicaraguan lakes, the San
Juan River basin south to Tortuguero in Costa Rica;
and (4) the Isthmian Province (Atlantic and Pacic
slopes) that includes southeastern Nicaragua, Costa
Rica (except for the small area occupied by the San
Juan Province) and all of Panama. One of the most
important geological units in Central America for
FIPs is Nuclear Middle America. This area is a sub-
region of Central America that is better known in the
geological literature as the Chortis Block; it encom-
passes all of Honduras and El Salvador from the
Motagua fault zone in southern Guatemala through
northern Nicaragua north of the Nicaraguan Lakes
(Rogers et al. 2007). This region includes three FIPs
(Miller 1966; Bussing 1976): the Usumacinta and the
San Juan in the Atlantic slope and the Chiapas-
Nicaraguense in the Pacic slope. Later studies
(Smith and Bermingham 2005; Abell et al. 2008)
utilizing more comprehensive datasets identied
additional biogeographic complexity in Central
America: the four initial FIPs were replaced by 16
smaller provinces (Abell et al. 2008); however, the
three FIPs in Nuclear Middle America remained
unchanged. This stability has more to do with a
paucity of data for Nuclear Middle America than a
comprehensive initial assessment. In Millers (1966)
assessment he identied the need for a more thorough
sampling in Nuclear Middle America; in his map of
FIPs he left the entire Atlantic slope of Honduras and
Nicaragua undesignated. Bussings (1976) biogeo-
graphic analysis of Central America added Martins
(1972) Honduran data to Millers (1966) but this
combination was still quite limited. The incomplete-
ness of these prior analyses is demonstrated by the
checklist of Matamoros et al. (2009). In this checklist
the known species richness of native freshwater shes
for Honduras was increased from the 88 reported by
Martin (1972) to 166. Matamoros et al. (2009) also
increased the distributional ranges of 12 species;
however, this work did not include analyses of FIPs.
Herein, we use the data of previous works particu-
larly Matamoros et al. (2009) to update the FIPs of
Nuclear Middle America.
Traditionally, biogeographical provinces have
been dened based on visual inspection of the
geographic patterns of species distributions (e.g.
Miller 1966; Martin 1972; Bussing 1976). However,
recent studies have pursued more quantitative
approaches (e.g. Kreft and Jetz 2010), including
ordination analyses (e.g. correspondence analysis
[CA]) and cluster analyses (e.g. Unweighted Pair
Group Method with Arithmetic Mean [UPGMA]).
These approaches have been used successfully in
ichthyological studies at both continental (Unmack
2001; Reyjol et al. 2007) and regional levels (Smith
and Bermingham 2005; Filipe et al. 2009).
Given the complex topography, physiography,
geologic history, and diverse ecosystems that char-
acterize Central America and that have presumably
shaped its biological diversity (Coates and Obando
1996) of which Nuclear Middle America occupies a
signicant portion, it is conceivable that three
provinces do not adequately describe the Nuclear
Middle America ichthyography. Accordingly, our
goal was to investigate the biogeographical patterns
of Nuclear Middle America obligate freshwater shes
by conducting quantitative analyses on the detailed
distributional data now available for the region.
Methods
Data sources
Distributional data for 76 native obligate freshwater
shes found in 29 Nuclear Middle American river
drainages (Fig. 1; Appendix Table 3) were obtained
fromVilla (1982); Bussing (2002); Miller et al. (2005);
Kinh-Pineda et al. (2006), and Matamoros et al. (2009).
The river drainage was used as our geographical
operational unit, since it has been identied as one of
the most important factors in freshwater sh biogeog-
raphy (Gilbert 1980). We followed Myers (1949) for
classication of species as primary and secondary.
Peripheral freshwater shes, which are tolerant of
higher ranges of salinity conditions (Myers 1949) and
can easily disperse among drainages along the coast-
line, were excluded from analysis. The nal data
matrix consisted of the presence/absence of the 76
primary and secondary species across 29 Nuclear
Middle America river drainages (Appendix Table 3).
Data analysis
First, we ran a CA based on the presence/absence data
matrix. Because CAs use a chi-squared metric and as
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such are non-sensitive to zero matches (Hugueny and
Leveque 1994), they are appropriate for ecological
and biogeographical multivariate analysis (Legendre
and Legendre 1998). We created a second data
matrix from the scores of the rst three axes of the
CA and ran a UPGMA on this matrix using
Euclidean distances as a distance measure. Corre-
spondence analysis data transformation reduces noise
associated with the original dataset (Gauch 1982;
Jackson and Harvey 1989; Hugueny and Leveque
1994), as noise is assumed to be uninformative from a
biogeographical perspective (Hugueny and Leveque
1994). In order to test how accurately the dendro-
gram resulting from the UPGMA represented the
original dataset, a cophenetic correlation coefcient
analysis (Farris 1969) was performed. Correlation
results above 0.9 represent a very good t; values
between 0.8 and 0.9 depict a good t; and results
below 0.8 represent a poor t to the data (Rohlf
1997). We used the statistical software package R
2.8.1 (R Development Core Team 2008) to perform
all the above procedures.
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Guatemala
Nicaragua
Panama
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El Salvador
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Fig. 1 A map of Central America showing Nuclear Middle
America and the locality of 29 river drainages used in this
study. 1 R o Polochic-Izabal, 2 R o Motagua, 3 R o Chame-
lecon, 4 R o Ulua, 5 R o Lancetilla, 6 R o Lean, 7 R o Cuero Y
Salado, 8 R o Bonito, 9 R o Danto, 10 R o Cangrejal, 11 R o
Lislis, 12 Bay Islands, 13 R o Aguan, 14 R o Sico-Tinto, 15
R o Platano, 16 R o Patuca, 17 R o Warunta, 18 R o Coco, 19
R o Wawa, 20 R o Prinzapolka, 21 R o Esclavos, 22 Lago
Amatitlan, 23 R o Maria Lucinda, 24 R o Lempa, 25 R o
Goascoran, 26 R o Nacaome, 27 R o Choluteca, 28 R o Negro,
29 R o Estero Real
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Subsequently, we visually searched the resulting
dendrogram from the UPGMA for clusters that
represented species-drainage relationships (i.e. fresh-
water ichthyological provinces). After we established
these provinces, we tested for differences in species
composition with a one-way analysis of similarity
(ANOSIM) as implemented in PRIMER v.6 (Clarke
and Gorley 2006). ANOSIM tests for differences
between and within a priori groupings (Clarke and
Warwick 1994). A test statistic (R) is computed,
which reects the observed differences between
groupings, contrasted with differences within group-
ings. The R statistic ranges between 0 and 1: if R = 1
then all sites within a group are more similar to each
other than any sites from different groups, and if
R = 0 then the similarities between and within
groups are the same on average (Clarke and Warwick
1994).
To estimate the rates of species turnover among
FIPs, we ran a beta-diversity analysis (Whittaker
1960, 1972) using the Whittaker index (b
w
) as in
Koleff et al. (2003). Beta-diversity measures the
difference in species composition either between two
or more local assemblages or between local and
regional assemblages (Koleff et al. 2003). To identify
the species that characterized each province we used
an indicator species analysis (ISA; Dufrene and
Legendre 1997). The ISA values are high when
individuals of one species are found in all localities
that comprise a province and not found in any other
province (see Dufrene and Legendre 1997). However,
species that are widely distributed across provinces or
have very small ranges limited to just a few localities
within a province would yield low ISA values. To
implement the ISA, a Monte Carlo test seeded with
1,000 random permutations was used to test the
signicance of the indicator value of each species
within a group. The beta-diversity analysis and ISA
were implemented with the statistical software pack-
age R 2.8.1 (R Development Core Team 2008).
Results
Fish community composition
Seventy six native obligate freshwater shes were
reported inNuclear Middle America (AppendixTable 3).
The family Cichlidae was the most speciose (Fig. 2),
contributing 29 species (38.2% of total). The next most
speciose family was the Poeciliidae (24species31.6%
of total), followed by the Characidae (7 species9.2%
of total), Profundulidae (4 species5.3% of total),
Rivulidae (3 species3.9% of total), Heptapteridae (3
species3.9% of total), Gymnotidae (2 species2.6%
of total), Synbranchidae (2 species2.6% of total),
Lepisosteidae (1 species1.3%of total) and Anablepi-
dae (1 species1.3% of total).
Primary freshwater sh species made up a very
small percentage of the total number of species
(12 species15.8% of total; Appendix Table 3). The
most species (seven) were found within the Characi-
dae, and included Astyanax aeneus, Brycon guatemal-
ensis, Bryconamericus scleroparius, Hyphessobrycon
tortuguerae, H. compressus, H. milleri, and Roeboides
bouchellei. Heptapteridae was represented by three
species: Rhamdia guatemalensis, R. laticauda and
R. nicaraguensis. The Gymnotidae family contributed
two species: Gymnotus cylindricus and G. maculosus
(Appendix Table 3). In any given province, these
primary freshwater sh species contributed between
14.9 and 33.3% of the species present (Fig. 3).
Secondary freshwater sh species were better repre-
sented in each province accounting for 66.7 and 85.1%
of the species (Fig. 3).
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Fig. 2 Bar plot showing Nuclear Middle American number of
obligate freshwater shes per family
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Endemism
Of the 76 obligate freshwater sh species reported to
inhabit Nuclear Middle America only 16 are endemic
to Nuclear Middle America with four pending formal
description. Amphilophus hogaboomorum and Ama-
titlania coatepeque are found in the Pacic Slope of
Nuclear Middle America. Alfaro huberi and Pro-
fundulus portillorum are endemics found in the
Atlantic and Pacic slopes. Finally, the vast majority
of endemics are found in the Atlantic slope of
Nuclear Middle America and include: Archocentrus
spinosissimus, Carlhubbsia stuarti, Heterandria an-
zuetoi, Heterandria milleri, Poecilia sp. 1 (hondur-
ensis), Poecilia sp. 2 (coco), Poecilia sp. 3
(Choluteca), Profundulus sp. 2 (sta. barbara), Ther-
aps microphthalmus, Theraps wesseli, Thorichthys
aureus and Xiphophorus mayae.
Cluster analysis
The results produced by the Euclidean distance-based
UPGMA (Fig. 4; cophenetic correlation coefcient
= 0.92) showed four distinctive clusters which the
ANOSIM indicated were signicantly different
(P = 0.001, R = 0.74). All pairwise comparisons
between clusters (i.e. freshwater ichthyological prov-
inces) were signicant (P B 0.05). These provinces
are described in detail below along with the results of
the beta-diversity analysis and the ISA.
Honduran ichthyological provinces
Honduras and Guatemala Caribbean Highlands
Province (HGCHP)
This province is formed by the Polochic-Izabal River
drainage in the Atlantic slope of Guatemala, the
Motagua River drainage shared by Honduras and
Guatemala, and the Chamelecon, Ulua, Lancetilla,
Lean, Cuero y Salado, Bonito, Danto, Cangrejal, and
Lislis River drainages in the Honduras Atlantic slope.
This province also includes the Bay Islands of
Honduras (Figs. 1, 4). Highly signicant P values
(0.0010.004, Table 1) resulted from the compari-
sons between the HGCHP and all other provinces,
suggesting that the HGCHP support different com-
munities. The ANOSIM R statistic and the beta-
diversity values recovered between the HGCHP and
all other provinces also indicates that obligate
freshwater sh communities are signicantly differ-
ent between provinces. Indicator species for the
HGCHP are: Parachromis friedrichsthali, Ophister-
non aenigmaticum, Poecilia orri, Xiphophorus may-
ae. Ten endemics are found in the HGCHP including
Archocentrus spinosissimus, C. stuarti, H. milleri,
P. sp.1 (hondurensis), P. sp. 2 (coco), P. sp.2 (sta.
barbara), T. wesseli, T. aureus, T. microphthalmus,
and X. mayae.
Honduras and Nicaragua Mosquitia Province
(HNMP)
The HNMP includes the Aguan, Sico-Tinto, Platano,
Patuca, Warunta River drainages in Honduras, the
Coco River which is shared between Honduras and
Nicaragua and the Wawa and Prinzapolka River
drainages in Nicaragua (Figs. 1, 4). The mid-level
beta-diversity values between the HNMP and other
provinces are an indicator of a fair amount of species
turnover (Table 1). However, the results of the
ANOSIM (Table 1) indicates that even though
Primary
Secondary
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HGCHP
HNMP CENP CNPP
Ichthyological provinces
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14.9%
85.1% 85.1% 85.1% 85.1%
19.2%
33.3%
20.6%
A A P P
Fig. 3 Stack bar plot showing the percentage of species
contributions per province by species tolerance to salinity.
White represents primary freshwater shes, gray represents
secondary freshwater shes. Each province is labeled with an
A or P to indicate drainages of the Atlantic Slope or Pacic
Slope, respectively. HGCHP Honduras and Guatemala Carib-
bean Highlands Province, HNMP Honduras and Nicaragua
Mosquitia Province, CENP Chiapas-El Salvador-Nacaome
Province, CNPP Choluteca and Nicaragua Pacic Province
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species are shared among these provinces the com-
munities are signicantly different. The HNMP is the
only Nuclear Middle America province that has
contact with all other provinces and this may explain
the mid-level beta-diversity values. Amphilophus
alfari, Parachromis dovii, P. managuensis, and
P. loisellei are species found with signicant indica-
tor species values (Table 2). No endemic species
were found in the HNMP.
Chiapas-El Salvador-Nacaome Province (CENP)
This province spans Pacic slope drainages in several
countries including the Los Esclavos River, Lago
Amatitlan and the Maria Lucinda River of Guate-
mala, the Lempa River of El Salvador and the
Goascoran and Nacaome Rivers of Honduras
(Figs. 1, 4). The cluster analysis placed these river
drainages into a single ichthyological province
(Figs. 4) and was strongly supported by the ANOSIM
with pairwise R statistics ranging from 0.79 to 1.00
(Table 1). Beta-diversity values between CENP and
the two Atlantic provinces were fairly high. However,
Estero Real
Choluteca
Negro
Nacaome
Goascoran
Amatitlan
Maria Lucinda
Esclavos
Lempa
Aguan
SicoTinto
Platano
Coco
Wawa
Prinzapolka
Patuca
Warunta
Bay Islands
Chamelecon
Lancetilla
Cangrejal
Lislis
Lean
Cuero & Salado
Bonito
Danto
PolichIzabal
Motagua
Ulua
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HGCHP
HGCHP
HNMP
HNMP
CENP
CENP
CNPP
CNPP
0.0 0.5 1.0 1.5
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Guatemala
Nicaragua
Panama
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El Salvador
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Fig. 4 Color coded
UPGMA dendrogram and
map of Central America
depicting Nuclear Middle
America ichthyological
provinces. HGCHP
Honduras and Guatemala
Caribbean Highlands
Province, HNMP Honduras
and Nicaragua Mosquitia
Province, CENP Chiapas-El
Salvador-Nacaome
Province, CNPP Choluteca
and Nicaragua Pacic
Province
Table 1 Whittaker beta-diversity index (b
w
) as in Koleff et al.
(2003) is found above the diagonal line
Provinces HGCHP HNMP CENP CNPP
HGCHP 0.51 0.70 0.69
HNMP 0.001 (0.59) 0.67 0.52
CENP 0.001 (0.80) 0.001 (1.00) 0.36
CNPP 0.004 (0.75) 0.006 (1) 0.012 (0.79)
A b
w
value of 0 means that species composition between
provinces is equal and a value of 1 means there are no shared
taxa between provinces. The results of ANOSIM pairwise
comparisons are below the diagonal. The R statistic for each
comparison is reported in parentheses below the P values.
Signicant (P B 0.05) P values are in bold
HGCHP Honduras and Guatemala Caribbean Highlands
Province, HNMP Honduras and Nicaragua Mosquitia
Province, CENP Chiapas-El Salvador-Nacaome Province,
CNPP Choluteca and Nicaragua Pacic Province
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the beta-diversity value between the CENP and the
Choluteca and Nicaragua Pacic Province (CNPP)
was 0.36 (Table 1) indicating that there is a fair
number of species overlapping between these two
provinces. However, the pairwise ANOSIM was
signicant (P = 0.012) for these provinces (Table 1).
The ISA (Table 2) detected ve species with signif-
icant indicator values in the CENP: Cichlasoma
trimaculatum, Poecilia salvatoris, Profundulus gua-
temalensis, Parachromis motaguensis, and G. macu-
losus. Amatitlania coatepeque is the only endemic
found in this province.
Choluteca and Nicaraguan Pacic Province (CNPP)
This province is formed by the Choluteca, Negro, and
Estero Real River drainages (Figs. 1, 4). All com-
parisons between the CNPP and all other provinces
were signicant with P values ranging from 0.012 to
0.006 (Table 1). The beta-diversity analysis (Table 1)
indicates that the amount of species shared between
the CNPP and the two Atlantic slope provinces is
low. However, the beta diversity value for the CNPP
CENP comparison was 0.36, indicating that there is a
fair number of species shared between those two
provinces. A signicant P value of 0.012 and a R
statistic of 0.79 suggest that these two provinces are
statistically different when comparing their obligate
freshwater species composition. The ISA detected
ve species with signicant indicator values in the
CNPP: Archocentrus centrarchus, .A. multispinosus,
A. hogaboomorum and Poeciliopsis turrubarensis.
Amphilophus hogaboomorum is the only endemic
found in this province.
Conclusions
The division of the Nuclear Middle America land-
scape into four smaller ichthyological provinces
(Fig. 4) disagrees with previous ndings (e.g. Bus-
sing 1976), which suggested only three larger
ichthyological provinces. In our work we found two
provinces in the Pacic slope of Nuclear Middle
America, and two more in the Atlantic (Fig. 4). In
addition, each province has unique species assem-
blages. We hypothesize that the larger number of
indicator species and endemics in the HGCHP may
reect the Motagua Rivers position as a biogeo-
graphical transition between regions of higher species
richness to the north and the depauperate species
richness to the south of that zone (Miller 1966; Myers
1966). Additionally, the Motagua and the Polochic-
Izabal River drainages appear to be distinct in the
UPGMA dendrogram, however, because they did not
statistically differ from the remaining drainages that
Table 2 Results of the
indicator species analysis
for the four Nuclear Middle
America ichthyological
provinces
Only species with
signicant value of
P B 0.05 and species
indicator values C0.6 (bold)
are reported
HGCHP Honduras and
Guatemala Caribbean
Highlands Province, HNMP
Honduras and Nicaragua
Mosquitia Province, CENP
Chiapas-El Salvador-
Nacaome Province, CNPP
Choluteca and Nicaragua
Pacic Province
Species HGCHP HNMP CENP CNPP P value
Parachromis friedrichsthalii 0.92 0.00 0.00 0.00 0.001
Ophisternon aenigmaticum 0.75 0.00 0.00 0.00 0.001
Poecilia orri 0.73 0.10 0.00 0.00 0.001
Xiphophorus mayae 0.67 0.00 0.00 0.00 0.005
Parachromis dovii 0.00 1.00 0.00 0.00 0.001
Amphilophus alfari 0.00 0.75 0.00 0.00 0.003
Parachromis managuensis 0.03 0.74 0.00 0.00 0.001
Parachromis loisellei 0.21 0.63 0.00 0.00 0.001
Cichlasoma trimaculatum 0.00 0.00 0.83 0.00 0.001
Poecilia salvatoris 0.00 0.00 0.67 0.00 0.001
Profundulus guatemalensis 0.00 0.00 0.67 0.00 0.002
Parachromis motaguensis 0.04 0.00 0.63 0.07 0.002
Gymnotus maculosus 0.00 0.00 0.60 0.27 0.014
Archocentrus centrarchus 0.00 0.01 0.00 0.89 0.002
Amphilophus hogaboomorum 0.00 0.00 0.00 0.67 0.001
Archocentrus multispinosus 0.00 0.24 0.00 0.62 0.013
Poeciliopsis turrubarensis 0.00 0.00 0.27 0.60 0.013
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formed the HGCHP they were placed within the
HGCHP. Further research should be conducted to test
the hypothesis that these river drainages may be
biogeographically transition zones.
Identication of ner levels of biogeographic
structure is consistent with other recent studies in
the region. Smith and Bermingham (2005) split the
three historically known lower Mesoamerican ich-
thyological provinces in seven smaller provinces and
Abell et al. (2008) split the Central American region
in 16 smaller ichthyological provinces. Low levels of
endemism are one of the most striking patterns of
Nuclear Middle America freshwater sh distribution
(Appendix Table 3; Table 1). Nuclear Middle Amer-
ica rests in a region characterized by an active
geological history that includes faulting, volcanism,
orogeny, and sea level change (Martin 1972), which
is thought to promote speciation (Coates and Obando
1996). However, this active geological history does
not seem to have produced the same extensive
evolutionary diversication of obligate freshwater
shes as it has for other vertebrate taxa in the region
(e.g. amphibians and reptiles; McCranie and Wilson
2002; Wilson and McCranie 2003). Low and mid-
level beta-diversity values between the provinces
may be explained by historical geological events that
promoted drainage connectivity. For example, Pleis-
tocene stream capture has been reported between the
Patuca and the Coco rivers (Rogers 1998; Marshall
2007). Furthermore; connections between Pacic and
Atlantic drainages via the Honduran depression (the
Comayagua graben) may have existed during the
Miocene (Sapper 1902; Olson and McGrew 1941;
Martin 1972).
Another puzzling feature of the Honduran ichthy-
ofauna is the extreme paucity of primary freshwater
shes. Myers (1966) discussed the overall scarcity of
primary freshwater shes in Central America, which
is most prominent in the area of Nuclear Middle
America. To explain the lack of primary freshwater
shes in this region, Myers (1966) suggested that the
most feasible theory is that the invasion of these taxa
in the region coincided with the lifting of the
Panamanian isthmus approximately 3.3 Mya. Conse-
quently, there has been insufcient time for extensive
speciation. This theory, however, is not congruent
with recent molecular data that date the arrival of
several primary freshwater shes in Central America
as occurring approximately 47 Mya (Bermingham
and Martin 1998; Perdices et al. 2002; Perdices et al.
2005; Concheiro Perez et al. 2007; Ornelas-Garcia
et al. 2008). To date, the timing of the arrival of
primary freshwater shes in Central American
remains unresolved. Perhaps more interesting is the
question as to why the primary freshwater families
(i.e. catshes, characids and gymnotids) in Nuclear
Central America are distinctly depauperate in species
richness compared to southern Central America
(Angermeier and Karr 1984) and South America
(Ouboter and Mol 1993; Hardman et al. 2002), where
these three are among the most speciose groups.
Molecular systematic and phylogeographic studies
may provide additional insight into the biogeography
of Nuclear Middle America freshwater shes. For
example, Perdices et al. (2002) in a phylogenetic
analysis of the genus Rhamdia in Central America
found that R. guatemalensis from the Lempa River in
the Pacic slope of Honduras was most closely
related to individuals from the Patuca and Aguan
Rivers, which are located in the Atlantic slope of the
country. Similarly, they also found that R. laticauda
from the Choluteca River (Pacic slope) was most
closely related to individuals from the Ulua and
Patuca Rivers (Atlantic Slope). At least for the genus
Rhamdia, a variety of historical drainage connections
between the Atlantic and Pacic slopes appears to
have facilitated the dispersal of these freshwater
shes across Nuclear Middle America. Additional
studies on other wide-ranging taxa may further
characterize these types of geologic events that have
shaped the distribution of freshwater shes in Nuclear
Middle America.
The Nuclear Middle America landscape was
formerly presented as being mostly homogeneous
with ichthyological differences existing between the
Pacic and Atlantic slopes only and consequently
disregarded the complexity of the region landscape
from east to west. This research lls in the previous
gaps in species distribution data resulting in analyses
capable of producing new river drainage-species
relationships at a ner scale and, as such, reveals a
region much more ichthyologically complex than
previously demonstrated. The dividing of Nuclear
Middle America Atlantic and Pacic slopes into four
smaller distinctive ichthyological provinces is con-
gruent with modern biogeographical inference, in
which an analytical approach is applied and the
results are interpreted in light of the physiographic,
Rev Fish Biol Fisheries
1 3
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ecological and geological features of landscape
(Unmack 2001; Smith and Bermingham 2005). From
a conservation perspective, ichthyological provinces
derived from a more nely scaled approach can
provide NGOs and other agencies a more useful
framework for prioritizing conservation planning
efforts in a region (Higgins 2003).
Acknowledgments Funding for this study was provided by
the Critical Ecosystems Partnership Fund grant # 51962. The
World Wildlife Fund W F Russell E. Train Education for
Nature fellowship, and the United States Agency for
International Development project (USAID/MIRA). We
would especially like to thank Jorge Ivan Restrepo, Fredy
Membreno and Gunther Suarez from the Instituto Regional
para la Biodiversidad and the Instituto Zamorano de
Biodiversidad for their timely and altruistic cooperation.
Pepe Herrero of project USAID /MIRA provided crucial
logistic support for sampling in the north coast of Honduras.
We would also like to thank the undergraduate students from
the Universidad Nacional Autonoma de Honduras who helped
in the eld while collecting samples for this study: Marcela
Matamoros, Alejandra Sanchez, Melissa Medina, Jonathan
Hernandez, Fausto Elvir, and Hermes Vega. The Honduran
biologists Hector Portillo and Juan Carlos Carrasco also
volunteered and spent much time in the eld with our team.
We are also very grateful to Luis Morales from Direccion
General de Pesca, and Claudia Carcamo, Ivonne Oviedo and
Andres Alegr a from the Instituto de Conservacion Forestal for
their help in obtaining collection permits in Honduras. Finally,
anonymous reviewers are gratefully acknowledged for
comments on a draft of the manuscript.
Appendix
See Table 3.
Table 3 List of obligate freshwater shes found in 29 Nuclear Middle America river drainages
Sal. Family Ichthyological
provinces
HGCHP HNMP
Species/river
drainages
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
I Characidae Astyanax aeneus 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1
I Characidae Brycon
guatemalensis
1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0
I Characidae Bryconamericus
scleroparius
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1
I Characidae Hyphessobrycon
compressus
1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
I Characidae Hyphessobrycon
milleri
1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
I Characidae Hyphessobrycon
tortuguerae
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1
I Characidae Roeboides
bouchellei
0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1
I Gymnotidae Gymnotus
cylindricus
1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1
I Gymnotidae Gymnotus maculosus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
I Heptapteridae Rhamdia
guatemalensis
0 1 1 1 0 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1
I Heptapteridae Rhamdia laticauda 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1
I Heptapteridae Rhamdia
nicaraguensis
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1
II Lepisosteidae Atractosteus tropicus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Anablepidae Anableps dowei 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Amatitlania
coatepeque
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Amatitlania
nigrofasciata
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
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Table 3 continued
Sal. Family Ichthyological
provinces
HGCHP HNMP
Species/river
drainages
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
II Cichlidae Amatitlania siquia 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1
II Cichlidae Amphilophus alfari 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1
II Cichlidae Amphilophus
hogaboomorum
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Amphilophus
longimanus
0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1
II Cichlidae Amphilophus
macracanthus
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Amphilophus
robertsoni
1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 0 0 0 0
II Cichlidae Archocentrus
spinosissimus
1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Archocentrus
centrarchus
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0
II Cichlidae Archocentrus
multispinosus
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1
II Cichlidae Cichlasoma bocourti 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Cichlasoma
trimaculatum
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Cichlasoma
ufermanni
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Cichlasoma
urophthalmus
1 1 1 1 1 1 1 1 0 0 1 0 0 0 0 0 0 1 1 1
II Cichlidae Criptoheros cutteri 0 1 1 1 1 1 1 1 1 1 1 0 1 1 0 1 0 0 0 0
II Cichlidae Criptoheros spilurus 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Hypsophrys
nicaraguensis
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0
II Cichlidae Parachromis dovii 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1
II Cichlidae Parachromis
friedrichsthalii
1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0
II Cichlidae Parachromis loisellei 0 1 1 1 1 1 1 0 0 1 0 0 1 1 1 1 1 1 1 1
II Cichlidae Parachromis
managuensis
0 0 1 1 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1
II Cichlidae Paraneetroplus
guttulatus
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Paraneetroplus
maculicauda
1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1
II Cichlidae Parachromis
motaguensis
0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Rocio octofasciata 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Theraps
microphthalmus
0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Cichlidae Theraps wesseli 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0
II Cichlidae Thorichthys aureus 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Alfaro cultratus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1
II Poeciliidae Alfaro huberi 0 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1
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Table 3 continued
Sal. Family Ichthyological
provinces
HGCHP HNMP
Species/river
drainages
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
II Poeciliidae Belonesox belizanus 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1
II Poeciliidae Brachyrhaphis
holdridgei
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1
II Poeciliidae Carlhubbsia stuarti 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Gambusia luma 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Gambusia
nicaraguensis
1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1
II Poeciliidae Heterandria
anzuetoi
0 1 1 1 1 1 0 0 0 1 1 0 1 1 1 1 1 1 0 0
II Poeciliidae Heterandria
bimaculata
1 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1
II Poeciliidae Heterandria
litoperas
1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Phallichthys amates 0 1 1 1 1 1 1 0 0 1 1 0 1 1 1 0 1 1 1 1
II Poeciliidae Poecilia gillii 0 1 1 1 0 0 0 0 0 1 1 0 1 1 1 1 1 1 1 1
II Poeciliidae Poecilia marcellinoi 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Poecilia orri 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0
II Poeciliidae Poecilia Rositae 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Poecilia salvatoris 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Poecilia sp. 1
(hondurensis)
0 0 1 1 1 1 1 1 1 1 1 0 1 1 0 0 0 0 0 0
II Poeciliidae Poecilia sp. 2
(coco)
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0
II Poeciliidae Poecilia sp. 3
(choluteca)
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Poeciliopsis
pleurospilus
1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Poeciliopsis
turrubarensis
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Xiphophorus helleri 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Xiphophorus
maculatus
1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Poeciliidae Xiphophorus mayae 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0
II Profundulidae Profundulus
guatemalensis
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Profundulidae Profundulus labialis 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Profundulidae Profundulus
portillorum
0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Profundulidae Profundulus sp. 2
(sta. barbara)
0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
II Rivulidae Kryptolebias
marmoratus
0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0
II Rivulidae Rivulus isthmensis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1
II Rivulidae Rivulus tenius 0 1 1 1 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0
II Synbranchidae Ophisternon
aenigmaticum
1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0
II Synbranchidae Synbranchus
marmoratus
0 1 1 1 0 1 1 1 1 0 1 0 1 1 1 1 1 1 1 1
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Table 3 continued
Sal. Family Ichthyological provinces CENP CNPP
Species/river drainages 21 22 23 24 25 26 27 28 29
I Characidae Astyanax aeneus 1 1 1 1 1 1 1 1 1
I Characidae Brycon guatemalensis 0 0 0 0 0 0 1 0 0
I Characidae Bryconamericus scleroparius 0 0 0 0 0 0 0 0 0
I Characidae Hyphessobrycon compressus 0 0 0 0 0 0 0 0 0
I Characidae Hyphessobrycon milleri 0 0 0 0 0 0 0 0 0
I Characidae Hyphessobrycon tortuguerae 0 0 0 0 0 0 1 0 0
I Characidae Roeboides bouchellei 1 1 1 1 1 1 1 1 1
I Gymnotidae Gymnotus cylindricus 0 0 0 0 0 0 1 0 0
I Gymnotidae Gymnotus maculosus 1 1 1 1 1 1 1 1 0
I Heptapteridae Rhamdia guatemalensis 1 1 1 1 1 1 1 1 1
I Heptapteridae Rhamdia laticauda 1 1 1 1 1 1 1 1 1
I Heptapteridae Rhamdia nicaraguensis 0 0 0 0 0 0 0 1 1
II Lepisosteidae Atractosteus tropicus 1 1 1 1 0 0 0 0 1
II Anablepidae Anableps dowei 1 1 1 1 1 1 1 1 1
II Cichlidae Amatitlania coatepeque 0 0 0 1 0 0 0 0 0
II Cichlidae Amatitlania nigrofasciata 1 1 1 1 1 1 1 0 0
II Cichlidae Amatitlania siquia 1 1 1 1 1 1 0 0 0
II Cichlidae Amphilophus alfari 0 0 0 0 0 0 0 0 0
II Cichlidae Amphilophus hogaboomorum 0 0 0 0 0 0 1 1 0
II Cichlidae Amphilophus longimanus 1 0 1 0 0 1 1 1 1
II Cichlidae Amphilophus macracanthus 0 0 1 1 1 0 0 0 0
II Cichlidae Amphilophus robertsoni 0 0 0 0 0 0 0 0 0
II Cichlidae Archocentrus spinosissimus 0 0 0 0 0 0 0 0 0
II Cichlidae Archocentrus centrarchus 0 0 0 0 0 0 1 1 1
II Cichlidae Archocentrus multispinosus 0 0 0 0 0 0 1 1 1
II Cichlidae Cichlasoma bocourti 0 0 0 0 0 0 0 0 0
II Cichlidae Cichlasoma trimaculatum 1 1 1 1 1 0 0 0 0
II Cichlidae Cichlasoma ufermanni 1 1 1 0 0 0 0 0 0
II Cichlidae Cichlasoma urophthalmus 0 0 0 0 0 0 0 0 0
II Cichlidae Criptoheros cutteri 0 0 0 0 0 0 1 0 0
II Cichlidae Criptoheros spilurus 0 0 0 0 0 0 0 0 0
II Cichlidae Hypsophrys nicaraguensis 0 0 0 0 0 0 0 0 0
II Cichlidae Parachromis dovii 0 0 0 0 0 0 0 0 0
II Cichlidae Parachromis friedrichsthalii 0 0 0 0 0 0 0 0 0
II Cichlidae Parachromis loisellei 0 0 0 0 0 0 0 0 0
II Cichlidae Parachromis managuensis 0 0 0 0 0 0 0 0 0
II Cichlidae Paraneetroplus guttulatus 1 1 1 0 0 0 0 0 0
II Cichlidae Paraneetroplus maculicauda 0 0 0 0 0 0 0 0 0
II Cichlidae Parachromis motaguensis 1 1 1 1 1 1 1 0 0
II Cichlidae Rocio octofasciata 0 0 0 0 0 0 0 0 0
II Cichlidae Theraps microphthalmus 0 0 0 0 0 0 0 0 0
II Cichlidae Theraps wesseli 0 0 0 0 0 0 0 0 0
II Cichlidae Thorichthys aureus 0 0 0 0 0 0 0 0 0
II Poeciliidae Alfaro cultratus 0 0 0 0 0 0 0 0 0
II Poeciliidae Alfaro huberi 0 0 0 1 0 0 1 0 0
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Sal. Family Ichthyological provinces CENP CNPP
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Roman numbers represent the families salinity tolerance (Sal.) based on Myers (Myers 1949)
Ichthyological provinces abbreviated as follows: HGCHP Honduras and Guatemala Caribbean Highlands Province, HNMP Honduras and
Nicaragua Mosquitia Province, CENP Chiapas-El Salvador-Nacaome Province, CNPP Choluteca and Nicaragua Pacic Province. Arabic
numbers represent the river drainages: 1 R o Polochic-Izabal, 2 R o Motagua, 3 R o Chamelecon, 4 R o Ulua, 5 R o Lancetilla, 6 R o Lean, 7
R o Cuero y Salado, 8 R o Bonito, 9 R o Danto, 10 R o Cangrejal, 11 R o Lislis, 12 Bay Islands, 13 R o Aguan, 14 R o Sico-Tinto, 15 R o
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