Philippine Journal of Science

142 (1): 55-67, June 2013

ISSN 0031 - 7683
Date Received: ?? Feb 20??

Diversity and Distribution of Freshwater Fish Assemblages

in Tayabas River, Quezon (Philippines)

Vachel Gay V. Paller1, Mark Nell C. Corpuz2,3*, and Pablo P. Ocampo1, 2

Animal Biology Division, Institute of Biological Sciences,

1

College of Arts and Sciences, University of the Philippines Los Baños, College, Laguna
2
UPLB Limnological Research Station, College of Arts and Sciences,
University of the Philippines Los Baños, College, Laguna
3
Southeast Asian Fisheries Development Center, Aquaculture Department,
Binangonan Freshwater Station, Binangonan, Rizal

Three stream sections (upstream, midstream, and downstream) of Tayabas River, Philippines were
surveyed during the wet and dry seasons of 2010 to evaluate the poorly known status of freshwater
fish assemblages. The study collected a total of 1,070 individuals comprising 15 species, 13 genera,
and 8 families. The three most abundant groups were poeciliids (61.85%), gobiids (26.16%), and
cichlid (5.51%). Shannon-Weiner’s diversity indices ranged from 1.270 to 2.171. Relatively high
Shannon evenness indices (0.653–0.846) and low Simpson’s dominance values (0.142–0.322) were
calculated implying a fairly equitable distribution of niche space for dominant and non-dominant
fishes. Significant change on fish assemblage in longitudinal gradient was observed (p<0.05), being
the most diverse fish assemblage registered in the upstream. Species richness is mostly composed
of native fish species (10 species) and mainly represented by stream gobiids (six species). The
downstream, however, had the highest cumulative abundance, in which the larger proportion was
from introduced species. Also, wet season had considerably more fish species and individuals relative
to dry season (p<0.05). This significant spatio-temporal differences in fish assemblage data were
evaluated by multivariate analyses (p<0.05). Canonical correspondence analysis identified the depth
(seasonal water level fluctuations), vegetation growth, and dissolved oxygen concentrations (in order
of importance) as the most influential environmental parameters affecting fish assemblage structure.
Also, climatic stress (prolonged drought) and anthropogenically-induced habitat alteration could
negatively affect the integrity of freshwater fishes within the river. The study suggests extensive
management programs of the river for the protection of native fish species.

Key Words: diversity, fish assemblages, native and introduced fish species

INTRODUCTION
The Mount Banahaw-San Cristobal Protected Landscape
(MBSCPL) that covers an area of about 109 km2 is one of
the few protected areas in Southern Luzon, Philippines.
One of the main aquatic ecosystems of MBSCPL is
*Corresponding author: mark.corpuz@seafdec.org.ph
marknellcorpus.@yahoo.com
the Tayabas River, which harbors diverse biota, but its
ichthyofauna remains poorly evaluated (LABB Inc. 2008).
The Philippine law (National Integrated Protected Area
Law Systems Act of 1992) ensures the protection and
conservation of these areas, particularly the forests and
the diverse flora and faunathey host (PSDN 2012). In
spite of the conservation efforts for MBSCPL, particularly
the forest component, the watersheds and river systems

55
Philippine Journal of Science
Vol. 142 No. 1, June 2013
of Mt. Banahaw are said to be under constant threat due
to human activities including illegal logging, “slash and
burn” (kaingin system), and improper disposal of garbage
and effluents to the river systems. Beyond these factors are
the lack of community-based education and participatory
protection of key aquatic habitats (Diesmos et al. 2004).
Diversity and distribution of riverine fish assemblages are
generally influenced by biotic and abiotic factors (Paul
& Meyer, 2001). These factors include, among others,
stream water levels and flow variability (Bradford &
Heinonen 2008), geo-hydrological feature of the river
(Angermeier & Winston 1999; Angermeier & Davideanu
2004.), microhabitat heterogeneity (Shervette et al. 2007),
and to a certain degree, aggravated by urbanization, habitat
alteration, anthropogenically-induced climate change
(Welcomme 1995; Zampella & Bunnell 1998; Vescovi et
al. 2009; Paller et al. 2011) and presence of invasive alien
fish species (Guerrero 2002).
Riverine ecosystems and its ichthyofauna support a wide
array of indispensable resources (Shinde et al. 2009).
The fish assemblages represent integrative biological
indicators of the stability of fisheries, impacts of habitat
deterioration and climate change, and ecosystem
productivity (Zampella & Bunnell 1998; Cagauan 2006;
Uy 2008).Thus, the use of fish assemblage data could
reflect the environmental and ecological conditions within
the longitudinal gradient of Tayabas River system. The
understanding of the ichthyofaunal population structure
is an important approach for potential sustainable use and
conservation management of both the species and aquatic
habitats (Paller et al. 2011). Moreover, the evaluation of
the poorly known conservation status of native freshwater
fish species will contribute to the development of future
sustainable fisheries and instigate protection from invasive
introduced faunas and beleaguered habitat conditions
(Guerrero 2002; Ong et al. 2002).
Synthetic diversity indices are now commonly applied to
the study of fish composition and distribution, primarily
to assess the health condition of rivers (Begonet al.
1996; Kwak & Peterson 2007). Likewise, appropriate
multivariate gradient techniques are being employed
to correlate the patterns of fish assemblage structure to
different habitat variables at spatial and temporal levels
(May & Brown 2000; Shervette et al. 2007).
There is no known information on the current status of
freshwater fish assemblages in Tayabas River and how
environmental factors influence the spatial and seasonal
conditions related to these assemblage structures. Hence,
the aim of this study was to characterize the freshwater fish
assemblage within the longitudinal gradient of Tayabas
River, comparing fish assemblage between the selected
stream reaches and between the two seasons (wet and dry).
Specifically, we described the spatio-temporal variations
56
Paller et al.: Fish Assemblages in Tayabas River, Philippines
in diversity, abundance, and distribution of the native
and introduced fish assemblages, and their associations
to environmental variables.
MATERIALS AND METHODS
Study site
The study site is relatively dry from November to April
and wet during the rest of the year (PAGASA 2010). Fishes
were collected in Tayabas River in Quezon, Philippines
(14o02’N, 121o31’E) (Figure 1). The river is water-fed by
the springs of Mt. Banahaw and run-offs from highlands
and nearby tributaries. It runs approximately 8.5–9.0
km traversing areas within the cities of Tayabas and
Lucena before draining to Tayabas Bay.It is located in the
southeastern portion of the mountain, approximately 8 km
heading about 120o towards the city of Tayabas. The whole
stretch presents typical mountain stream characteristics.
Upstream and midstream sites are surrounded by primary
and secondary forests. The river banks in either side are
primarily lined by mosses, bryophytes, and other riparian
vegetation, with some steep areas surrounded with coconut
plantations and perennial weeds. Downstream sites are
within the city proper of Tayabas. A large proportion of
inhabitants is engaged in fisheries, farming, and livestock
raising.
Sampling Design
Three stretch sections of the river (upstream, midstream,
and downstream) were used in this study. Three sampling
stations were selected along each stream reach (a total of
nine sampling stations). Each station had approximately
600–700m stream reach with three 200–250m sampling
run/site, which were considered replicates within each
station. Unit area sampled was equal to the maximum
length of seine net (8 m) multiplied to the distance seined
(m). Individual sampling run lasted 40–50 min and were
done during day time. A three-day sampling period was
carried out in February and September 2010 to represent
wet and dry season, respectively. A total of 18 samples
were made (nine sampling sites x two seasons).
Fish specimens were collected using a seine net (1.2
x 1.2-mm mesh), hand nets, fish trap, angling, and
12-v electro-fishing equipment. Captured fish were
immediately counted and identified at lowest possible
taxon. Specimens were either housed in laboratory as
live samples or preserved in 10% buffered formaldehyde
for further documentation and identification. Some
voucher specimens were deposited in the fish collection
at University of the Philippines Los Baños Museum of
Natural History, Philippines.
Philippine Journal of Science Paller et al.: Fish Assemblages in Tayabas River, Philippines
Vol. 142 No. 1, June 2013

120'50"E 121'17"E 121'40"E

14'24"N

14'09"N

13'50"N

14°02'N

121°31'E

Figure 1. Map outline of study areas of Tayabas River in Quezon, Philippines. Upstream (□), midstream (▲), and downstream (+). Sites
(numbers) within each reach are also included.

Before collection, dissolved oxygen (mgl-1, Hanna HI
3810), water temperature (oC), pH (Oakton pH tester
30), and salinity (ppt, Atago hand refractometer) were
recorded for each site. Geographic position and elevation
were also recorded for each sampling station using a GPS
device (CarNAVi Pro 400). Depth (cm) was measured
using an improvised 100-cm wooden ruler at two or three
points for each site. Water current or stream flow (m s-1)
was measured using a simple float. Dominant bottom
type was recorded and categorized as organic detritus,

57
Philippine Journal of Science
Vol. 142 No. 1, June 2013
silt, mud, sand (0.02–2 mm), gravel (2–64 mm), cobble
(64–256 mm), boulder (˃256 mm) (May &Brown 2000).
Vegetation cover (%) was estimated visually as determined
by the relative amount of submerged and floating aquatic
plants to sampling path as well as those occupying both
sides of the riverbanks.
Data analyses
Species richness was determined by the number of species
present in a community. The relative abundance for each
species was calculated as:
= (ai / A) 100%
where: ai is the number of individuals caught in the ith
species and A is the total number of species collected in
one sampling area during a sampling period. Diversity
index was computed following the Shannon-Weiner
diversity index (H’) (Shannon & Weaver 1949):
S & Verdonschot 1995; Legendre & Legendre 1998).
H’ = - ∑ pi ln pi
i=1
where: s is the number of species; p is the proportion of
individuals found in the ith species and ln is the natural
logarithm. Evenness (J’) was computed following the
Shannon’s diversity index:
J’ = H’ / ln S
where: S is the total number of species. Species dominance
was computed using the Simpson’s index formula (λ)
(Simpson 1949):
S ni(ni1)
λ=∑
i = 1 N(N1)
where: s is the number of species, ni is the number of
individuals in the ith species and N is the total number
of individuals.
Homoscedasticity (Levene’s test) and normality (Shapiro-
Wilk test) of variances were tested for species richness,
log 10 (x+1) transformed abundance data, and water
parameters. Number of taxa and values of Shannon’s
evenness, and Simpson’s dominance for each site in two
seasons were compared to one another. Bootstrapping
method using 1,000 random pairing was used to test the
significance (p<0.05). Comparison of Shannon-Weiner
diversity indices between the two seasons and among the
stream sites were examined using diversity t-test described
by Magurran (1998). Similarity among reaches at two
collection seasons was measured using Morisita-Horn
58
Paller et al.: Fish Assemblages in Tayabas River, Philippines
Index (Wolda 1981) and the unweight pair group average
method (UPGMA) was used to cluster similar group
(sites) according to their composition and log-transformed
abundance data. Pool of log-transformed abundance of
native and introduced fish assemblages between seasons
were compared using Hotelling’s T2 test (a multivariate
analogue to the t-test) (p<0.05). Descriptive statistics were
also computed for each stream reach in two sampling
periods. Pairwise comparison of the mean of each
parameter was tested using Tukey’s honestly significant
differences (p<0.05).
Correspondence analysis (CA) was used to examine
and visualize temporal variation among and between
native and introduced fish assemblages using the pool
of abundance data. Canonical correspondence analysis
(CCA) was used to investigate the association of sampling
sites with environmental and habitat variables Monte
Carlo test with 1,000 random permutations was applied
to test the significance (p<0.05) of the fish assemblage
structure and sites to environmental variables (Ter Braak
All statistical analyses were performed using Statistica
version 7.0.
RESULTS
Composition and Abundance of Fish Assemblages
Our ichthyofaunal survey recorded 431 (dry season) and
639 (wet season) specimens belonging to 15 species,
13 genera, and eight families (Table 1). Fish species
composition in the river system is largely composed of
indigenous species (10 species) and predominated by
gobiids (six species). Although exotic species comprised
a small portion of fish composition (five species), two
of the most dominant fish species were from this group.
The two most abundant species, which consisted at least
half of the total fish specimens were Poecilia sphenops
(wet= 32.55%; dry= 41.30%) and P. reticulata (wet=
26.29%; dry= 26.91%). This is followed by a native
species, Glossogobius celebius (wet= 24.57%; dry=
22.74%). These three most frequent species, jointly
with Orechromis niloticus, Channa striata, and Clarias
batrachus were common residents along the three stream
reaches in either seasons.
Variation in Ichthyofaunal Diversity
Species richness ranging from 6–13 was found to be
significantly different between seasons (U= 60, p<0.05),
with upper stream areas of wet season having the most
numerous taxa (13 species) (Table 2). Furthermore, overall
number of fish caught in the wet season (639) was more
 Table 1. Relative abundances and densities (in parenthesis, fish 10m-1) of freshwater fishes from Tayabas River, Philippines. Us= upstream, Ms= midstream, Ds= downstream.
Wet Season Dry Season Seasons Combined
Fish species Code
Vol. 142 No. 1, June 2013

Us Ms Ds Us Ms Ds Us Ms Ls
Philippine Journal of Science

Cyprinidae (Carps and Minnows)*

Puntius binotatus Pbin 1.56 (0.34) 0.47 (0.1) 0 (0) 0 (0) 0 (0) 0 (0) 0.93 0.28 0
Eleotridae (Sleepers and Gudgeons)*
Butis butis Bbut 0 (0) 0 (0) 0.47 (0.1) 0 (0) 0 (0) 0 (0) 0 0 0.28
Eleotris fusca Efus 0.31 (0.06) 0.47 (0.1) 0 (0) 0 (0) 0 (0) 0 (0) 0.19 0.28 0
Gobiidae (True gobies)*
Glossogobius celebius Gcel 3.91 (0.84) 11.27 (2,4) 9.39 (2.0) 4.18 (0.6) 6.96 (1.0) 11.6 (1.66) 4.02 9.53 10.28
Gobiopterus chuno Gchu 0.78 (0.16) 0 (0) 0 (0) 0 (0) 0 (0) 0 (0) 0.47 0 0
Redigobius bikolanus Rbik 0.94 (0.20) 0.78 (0.16) 0 (0) 0 (0) 0 (0) 0 (0) 0.56 0.47 0
Stiphodon elegans Sele 0.63 (0.14) 0.16 (0.04) 0 (0) 0 (0) 0 (0) 0 (0) 0.37 0.09 0
Stiphodon semoni Ssem 0.47 (0.10) 0 (0) 0 (0) 0 (0) 0 (0) 0 (0) 0.28 0 0
Sycopus sp. Syco 0 (0) 0 (0) 0 (0) 0.23 (0.04) 0 (0) 0 (0) 0.09 0 0
Syngnathidae (Pipefishes)*
Dorichthys martensii Dmar 1.72 (0.32) 1.56 (0.34) 0 (0) 0 (0) 0 (0) 0 (0) 1.03 0.93 0
Channidae (Murrels)
Channa striata Cstr 0.16 (0.04) 0.16 (0.04) 0.16 (0.04) 0.46 (0.66) 0.46 (0.66) 0.70 (0.10) 0.28 0.28 0.37
Clariidae (Freshwater Catfishes)
Clarias batrachus Cbat 0.16 (0.04) 0.31 (0.06) 0.47 (0.10) 1.03 (0.50) 0.70 (0.10) 0.23 (0.04) 0.09 0.47 0.47
Cichlidae (Tilapias)
Oreochromis niloticus Onil 0.78 (0.16) 3.13 (0.66) 1.10 (0.24) 1.86 (0.26) 1.39 (0.20) 3.02 (0.44) 1.21 2.43 1.87
Poecilidae (Livebearers)
Poecilia reticulata Pret 3.13 (0.66) 9.34 (2.00) 13.77 (3.0) 5.57 (0.8) 8.58 (1.24) 12.76 (1.82) 4.11 9.06 13.36
Poecilia sphenops Psph 3.76 (0.8) 10.64 (2.26) 18.15 (3.8) 8.12 (1.16) 9.98 (1.44) 23.2 (3.34) 5.42 10.09 19.81
Total number of species from each site 13 11 7 7 6 6 14 11 7
*native fish family

59
Paller et al.: Fish Assemblages in Tayabas River, Philippines
Philippine Journal of Science Paller et al.: Fish Assemblages in Tayabas River, Philippines
Vol. 142 No. 1, June 2013

Table 2. Abundance, number of taxa, and diversity indices recorded from the three stream reaches of Tayabas River,
Philippines in two sampling seasons. Us= upstream, Ms= midstream, Ds= downstream. H’= Shannon-Weiner diversity
index; J’= Shannon Evenness Index; λ= Simpson’s species dominance index.

Wet Season Dry Season

Us Ms Ds Us Ms Ds
Individuals 117c 245b 278a 92b 121a 222 a
Taxa 13a 11ab 7b 7b 6b 6b
H’ 2.171a 1.667b 1.270c 1.518b 1.384c 1.289c
J’ 0.846ab 0.695c 0.653d 0.780b 0.772bc 0.720c
λ 0.142c 0.232b 0.322a 0.261b 0.285ab 0.319a
For each biological index, means with same superscript letter are not significantly different (p>0.05)

than 32% of that in the dry season (431).

Shannon-Weiner’s diversity indices calculated for the
three reachesin both sampling periods varied from 1.270–
2.171 (Table 2). Spatially, the upstream reach showed
significantly highest diversity among the sampling areas
(p<0.05), while downstream sites were the least diverse
(p<0.05). Significant temporal variation in the mean
diversity index was also measured with wet season (1.69)
being more diverse than the dry season (1.35) (Diversity
t= 6.47, p<0.05).
Shannon’s evenness indices exhibited comparable
spatial pattern to that of Shannon-Weiner’s diversity
indices that ranged from 0.653–0.846 (Table 2). Highest
Figure 2. A dendrogram of UPGMA showing the clustering of sites
measurement were calculated in upstream, which was
derived from species composition and log-transformed
significantly different from the middle and downstream abundance data. Upstream at wet season (UW);
reaches (p<0.05). Diversity t-test for evenness, however, midstream at wet season (MW); downstream at wet
showed no significant seasonal differences (wet= 0.644; season (DW); upstream at dry season (UD); midstream
dry= 0.695) (p>0.05). at dry season (MD); downstream at dry season (DD).

Simpsons’ dominance values ranged from 0.142–0.322

and had a definite inverse relationship with diversity and
evenness. Apparently, upstream and downstream had
the significant lowest and highest dominance values,
respectively (p<0.05). Temporally, mean dominance index
were significantly different (wet= 0.24; dry= 0.299) (p<0.05).
Similarity
Analysis on spatial and temporal similarity produced two
conspicuous clusters. The first cluster was composed of
sites from the upstream and midstream during wet season,
and the second one was composed of all sites from dry
season including the downstream of wet season. The first
group registered a similarity of about 92%, which deviated
from the second group with approximately 73% level of
similarity. Interestingly, downstream in wet season was
clustered with the dry season sites at no less than 95%
level of similarity (Fig. 2).
Native and Introduced Fish Species
Although there was more numerous native species,
their cumulative relative abundance was lower than the
introduced fish species at wet season (natives= 34.90%;
introduced= 65.10%) and dry season (natives= 22.97%;
introduced= 77.03%) (Fig.3). Likewise, dominance in
numbers of introduced fishes was observed in dry season,
having no less than 75% of the total relative abundance
(Fig. 3). The only observation in which the relative
abundance of native fish species was higher (56.41%) than
introduced fish species (43.59%) was found in upstream
at wet season. This, however, drastically reduced with

60
Philippine Journal of Science
Vol. 142 No. 1, June 2013
Relative Abundance (%)
Figure 3. Stacked bar of percent representation in abundances of
native (gray bars) and introduced (black bars) freshwater
fishes from the three sampling sites in wet and dry
seasons. Upstream at wet season (UW); midstream at
wet season (MW); downstream at wet season (DW);
upstream at dry season (UD); midstream at dry season
(MD); downstream at dry season (DD).
decreasing altitude (midstream and downstream).
The Hotelling’s T2 test validated the significant variation
on fish assemblage (composition and log-transformed
abundance) between seasons (Hotelling T2= 43.02, F=
CA (18.51%)
Figure 4. Plot and population centroids (5% concentration ellipse level) for CA scores of native and introduced fish
assemblages between seasons. Native species of wet season (NatW, □); introduced species of wet season (IntroW,
∆); native species ofdry season (NatD, ■); introduced species of dry season (IntroD, ▼).
Paller et al.: Fish Assemblages in Tayabas River, Philippines
3.41, p<0.05). This was also visualized by CA plot (Fig.
4). The CA produced two axes that contributed to 58.62%
of the differences in species relative abundance. Loads of
variables and centroids of wet season groups were clearly
separated to that of dry season in Axis 1 (eigenvalue=
0.16) and partially overlapped at Axis 2 (eigenvalue=
0.07) (Fig. 4). Furthermore, native species abundance
also significantly changed across seasons, with higher
abundances on the wet season (HotellingT2= 78.27, F=
4.83, p<0.05). On the contrary, quantities of introduced
fishesdid not changed significantly (HotellingT2= 9.00,
F= 0.56, p>0.05) across seasons.
Canonical Correspondence Analysis
Fish assemblages were significantly different between
the three reaches when using abundance data (p<0.05).
The sites-environment relationships outlined by the
CCA had eigenvalues of 0.22 and 0.07 on the first
two axes, with 79.00% variability in fish assemblages
explained (Table 3). Scores of upstream and midstream
areas were partially separated from the downstream,
and all sites of the dry season (Fig. 5A). CCA also
recognized mean depth, vegetation, and DO (in order
of importance) as the most weighted causative abiotic
parameters structuring the fish assemblage pattern
CA (40.11%)
61
Philippine Journal of Science Paller et al.: Fish Assemblages in Tayabas River, Philippines
Vol. 142 No. 1, June 2013

CCA 2(19.56%)

CAA 1(59.44%)
CCA 2(19.56%)

CAA 1(59.44%)
Figure 5. Canonical correspondence analysis plot scores on the first two axes derived from
sampling sites (A) and species abundance data (B). Group enclosed by continuous
curve line was formed by upstream sites. Group enclosed by dashed lines were
formed by midstream sites.Group enclosed by oval were formed by downstream
sites. upstream (□); midstream (▲); downstream (+). Abbreviations of species
are given in Table 1.

62
Philippine Journal of Science Paller et al.: Fish Assemblages in Tayabas River, Philippines
Vol. 142 No. 1, June 2013

Table 3. Correlations of significant abiotic variables of the
first two CCA ordination axes. Most significant
weights on CCA 1 and CCA 2 are in bold.
CCA 1
Eigenvalues 0.22
% Variation 59.44
DO 0.35
pH 0.20
Temperature (-)0.33
Depth 0.67
Vegetation 0.51
Velocity (-)0.35
(Table 3). In this analysis, mean depth and vegetation
percentage were highly correlated with the first
axis, while DO and vegetation were the factors most
correlating with the second axis (Table 3).
The first ordination axis (left and right reading) (59.44%)
explained the fish assemblage variation caused by the
difference in longitudinal gradient characteristics. The CCA
diagram identified an aggregation of scores of several native
fishes in Axis 1. The Puntius binotatus, Eleotris fusca,
Gobiopterus chuno, Redigobious bikolanus, Stiphodon
elegans, Stiphodon semoni, and Dorichthys martensii
were most correlated to the upstream and midstream,
while a single native species, Butis butis is correlated to
the downstream (Fig. 5B). The most common riverine
inhabitants, Poecilia reticulata, P. sphenops, Glossogobius
celebius, Channa striata,Clarias batrachus intermediated
within the plot. The second axis (up and down reading)
(19.56%) also explained the gradient of fish assemblage
with B. butis, and Sycopus sp., representing the upper and
lower axes, respectively. The former was the typical species
of the downstream, while the latter was the common species
of the upstream (Figs 5B).
Water Parameters and Habitat Variables
Temporal and spatial comparison habitat and water quality
parameters were presented in Table 4. Significant temporal
CCA 2 differences were recorded in DO and mean depth (p<0.05),
albeit with no significant spatial variation. Aeration brought
0.07 by the water current could be the contributory factor in
19.56 the increase of DO level. Means of water temperature,
0.33 vegetation percentage, and water current were statistically
varied among stream reaches (p<0.05). Water temperature
0.21
was higher in the downstream and water velocity was faster
(-)0.15 in the upstream. Conditions of pH were fairly basic in all
0.27 sites and in two sampling seasons. Upstream and midstream
had considerably deeper water levels as compared to the
(-)0.30
lower stretch. Water depth drastically reduced during the
0.18 dry season. Vegetation growth was significantly denser in
upstream and remained unchanged after the rainy months.
The whole stretch of the river is presented a substrate
composed of sand, gravel, cobble, and boulder, a main
typical feature of a mountain stream.
DISCUSSIONS
The main characteristic of Tayabas River system is its
relatively diverse ichthyofauna (species= 15, H’= 1.55),
mostly represented by native species. The richness and
diversity are relatively higher compared tothe mountain
streams of Mount Makiling Forest Reserve including
Dampalit (species= 12, H’= 1.12) and Molawin (species=
12, H’= 1.19) (Paller et al. 2011), but lower than those have
been calculated in Bulusan River in Sorsogon (species=
16, H’= 2.41) and Pansipit River in Batangas (species=
21, H’= 3.05) (Corpuz et al. 2010; 2011). Although the
mountain rivers are not as diverse compared to estuarine-
linked streams, they remain very important ecosystem for
freshwater biota and for the communities that rely on their
resources (Ong et al. 2002).
The distribution of freshwater fish assemblages showed
low dominance (0.142-0.322) and relatively high evenness

Table 4. Summary of habitat and water quality parameters: mean ± standard deviation collated from three stream sections of Tayabas
River at two sampling periods.
Seasonal Variation Spatial Variation
Wet Dry t Upstream Midstream Downstream F
DO (mg l ) -1
5.96 ± 0.19 5.01 ± 0.10 4.36* 5.13 ± 0.65 a
5.8 ± 0.68
a
5.51 ± 0.33 a
1.67NS
pH 7.34 ± 0.12 7.29 ± 0.07 0.38NS 7.21 ± 0.29a 7.53 ± 0.31a 7.2 ±0.08a 2.94NS
Temperature (oC) 23.77 ± 0.80 24.68 ± 0.85 -0.78NS 21.23 ± 0.69a 26.17 ± 0.96b 25.27 ± 1.58b 31.92*
Depth (cm) 67.33 ± 9.13 41.56 ± 4.32 3.76* 58.5 ± 22.68 a
62.33 ± 16.52 a
42.5 ± 14.94 a
1.97NS
Water velocity (m s-1) 1.10 ± 0.17 1.2 ± 0.30 -0.86NS 1.38 ± 0.28a 0.97 ± 0.08b 1.1 ± 0.11b 8.46*
Vegetation (%) 60.00 ± 9.13 60 .00 ± 9.41 1.06NS 86.83 ± 4.66a 67.00 ± 6.42b 29.17 ± 10.68c 87.23*
*significant at 5% level of confidence, NS not significant at 5% level of confidence
For each abiotic variable, means with same superscript letter are not significantly different at 5% level of confidence

63
Philippine Journal of Science
Vol. 142 No. 1, June 2013
(0.653 - 0.846), which means that the allocation of niche
space is fairly distributed equitably for most dominant and
non-dominant fish species thriving in the river (Begon et
al.1996). The results indicate that abundance of dominant
species (particularly poeciliids) rose significantly without
measurably displacing harder and robust freshwater fish
species. It is rather assumed that poeciliid populations are
regulated by non-abundant highly effective piscivorous
predators. This can be possible given the fact that
poeciliids (P. sphenops and P. reticulata) are highly
prolific, yet they are diminutive omnivores, and have
pelagic swimming behavior (Burns et al 2009; UPLB-
LRS 2011). This scenario follows the keystone species
concept as characterized by few non-dominant predators
controlling the distribution and abundance of large
numbers of prey species (Mills et al. 1993). In this
riverine system, channids, gobiids, and clariids are the
potential keystone species. This could also be the possible
explanation on fairly high equitability values (J’), in spite
of very high relative abundance of few taxa (P. sphenops
and P. reticulata). Nevertheless, our assumption remains
inconclusive but requires a comprehensive study to
further assess the overall fish fauna compositions of main
watersheds in MBSCPL, and analyze the influence of
introduced fishes towards the dwindling populations of
indigenous fishes.
Higher diversity and equitability calculated in wet season
(Table 1 & 2) serve as an indicator to water and habitat
qualities. The results attribute this fish diversity to the wide
range of riverine habitats formed with the rising water
level (depth) brought by frequent precipitations (Table
2). During rainy months, it provides sufficient volume of
aerated water (DO) to replenish and refill the watersheds
and, thus facilitates the improvement of carrying capacity
for the optimum sustenance of diverse aquatic biota
(Vescovi et al. 2009). Similarly, pooled fish richness and
relative abundances were apparently higher during the
wet season. This supports the assumption that most of the
fish populations are intrinsically increasing during rainy
seasons (spawning season) in response to environmental
stimuli triggered by seasonal changes (Griffin & Ojeda
1992). Environmental stimuli directly affect endocrine
pathways and known to regulate metabolism and
reproduction in ectothermic fishes (Redding & Patiño
1993). During the breeding season, fish prefers suitable
locations (e.g. vegetative and rocky substrates) to deposit,
nest their eggs, and/or shelter themselves. These fish
breeding substrates including vegetation and remnants of
dead plants, rock formations, banks, and shallow pools
are usually found in the sampling stations. As expected,
a number of gravid fish as well as juveniles were found
in the seine sampling (data not shown), which suggests
continuous and successful recruitments within the river.
Temporal and spatial variations of fish assemblage and the
64
Paller et al.: Fish Assemblages in Tayabas River, Philippines
underlying factors that contributed to these variations were
detected by multivariate gradient analyses (Figs. 4, 5A, &
5B). In CCA, direct gradient analysis showed the most
influential environmental variables causing fish assemblage
variability (Figs. 5A & 5B). As reflected in the first
ordination axis, fish assemblage structure was significantly
influenced by mean depth and vegetation growth. These
parameters were regarded as good predictors of gradient
change in fish assemblage structure (Ross 1986; Shervette
et al. 2007; Vorwerk et al. 2007). The second strongest
gradient exhibited by the second ordination axis is best
explained by the DO level (Fig. 5A; Table 3). The species
scores of P. binotatus, E. fusca, G. chuno, R. bikolanus, S.
elegans, S. semoni, and D. martensii that are correlated to
the upstream in Axis 1 represent the pattern of fish species
in the upstream. Central scores from P. reticulata, P.
sphenops, G. celebius, C. striata, and C. batrachus revealed
that such species are the most typical residents along the
stream reaches. Furthermore, B. butis on extreme upper left
represents the species of the downstream.As reflected in
CA, the effect of seasonal and climatic change could also
alter the fish composition and abundance, particularly the
native one (Fig. 4).
Freshwater native fish assemblage is largely composed of
stream gobies that are well-adapted in pristine, vegetated,
fast-flowing, and cool headwater reaches. Unlike the
populations of G. celebius that are distributed throughout
the whole stream sections, all other native species (native
cyprinid, pipefish, and gudgeons) were observed to be
restricted in the upstream and midstream (Table 1). In
ecological viewpoint, the occurrence (also preference)
of these fish populations are generally dependent on the
environmental components and habitat feature of their
ecosystem including vegetation structure (Herre 1927;
Vorwerk et al. 2007), water depth (Ross 1986), elevation
(May & Brown 2000), water velocity (Herder & Freyhof
2006), and bottom substratum type (May & Brown 2000).
Mainly, because of their reliance in habitats in which they
live, these native fish species can be considered asone
of the ecologically important bioindicators of riverine
health status (Zampella &Bunnell 1998; Angermeier &
Davideanu 2004; Cagauan 2007).
The invasion success of introduced fish species occupies
the whole stretch of the study site, albeit the overall pattern
of their abundance is greater in the downstream. The
expansive nature of these fishes is accounted for their high
reproductive capacity and for being habitat generalists
(Guerrero 2005; Cagauan 2006). Their populations have
also the tendency to occupy warmer, shallow low gradient
streams and rivers. This occurrence of introduced species
is often attributed to accidental and deliberate introduction
by the fisherfolks and their escaping from nearby rice
paddies cum ponds (Joji Roxas, personal communication).
As suggested by Kennard et al. (2005) their presence may
Philippine Journal of Science
Vol. 142 No. 1, June 2013
signify both a warning sign and a cause of decline in river
health and the integrity of native fish assemblages.
Significant change on fish assemblage in the stream
longitudinal gradient was observed. In our findings, there
was a gradation in ichthyofaunal composition, with fish
assemblages within the river traversing the downstream
sites or within the cityproper being least diverse (Table 2).
The observations can be associated to the repercussion of
anthropogenic disturbances to the ichthyofaunal diversity.
Species diversity seems to decrease in areas exposed to
different stressors. Moreover, introduced fish species
are more likely to occur in anthropogenically impacted
environments (Kennard et al. 2005; Vescovi et al. 2009).
Secondary forest cover and aquatic vegetation growth
were minimal in the downstream considering most of
its embankments and foreshores were landscaped and
converted into residential areas. Domestic and agricultural
run-offs may also contribute to the pollution that
negatively affects the fish assemblage structure.
All sites exhibited apparent changes on fish assemblage
over the seasonal transition (Fig. 4), which implies a
functional instability of Tayabas aquatic ecosystems
under seasonal stress. It is worth mentioning that Luzon
areas had experienced a long period of drought correlated
to El Niño on the first six months of the year 2010. The
condition resulted to the evident decrease in the water
depth along the stretches of the river and may lead to the
reduction in habitat availability, food production, and
water quality (lesser DO and higher stream temperature)
(Bradford & Heinonen 2008) causing the decline of fish
abundance and compisition during the dry season (Uy
2008). This incident caused a profound ichthyofaunal
alteration, which is bias against the more vulnerable
native fish species, following exposure to extreme climatic
stress (Kennard et al. 2005; Xenopoulos et al. 2005).
Conversely, introduced species mainly dominated the fish
abundance during the 2010 El Niño phenomenon (Figs. 4
& 3). Threats like these may pose a level of ambiguity on
the survival of the indigenous fish species. For example,
cryptic genera such as herbivorous gobies (Stiphodon
and Sycopus), diminutive gobies (Gobiopterus and
Redigobius), cyprinid (Puntius), and pipefish (Dorichthys)
and related congeners were once abundant in the river
and adjacent tributaries in 1970s. The species, however,
are now rarely encountered in the Tayabas River system
(Joji Roxas, personal communication). A similar plight
was also documented in the current conservation status
of fishes in the watersheds of Mount Makiling Forest
Reserve (Paller et al. 2011).
Headwaters support higher species richness (represented by
native species) than the downstream. The latter, however,
possessed a greater number of fish individuals, in which
the larger proportion was from the introduced species. The
Paller et al.: Fish Assemblages in Tayabas River, Philippines
diversity and distribution of fish assemblage in Tayabas
River system are affected by interacting environmental
processes, seasonal changes, and anthropogenic pressures.
Water levels, vegetation growth, and DO levels are the
most prominent environmental factors structuring the
ichthyofaunal diversity and distribution within the river.
To some extent, anthropogenic disturbances and climate
change could degrade the riverine habitats impacting
ultimately on diversity loss of more susceptible group, the
native fish species. Thus, proper management strategies
(e.g., urban planning and reforestation) for downstream
sites of MBSCPL are in the utmost importance to
revive the declining populations of native species. This
ichthyofaunal study updated the inventory of freshwater
fishes in Tayabas River and provided essential dataset for
future ichthyofaunal survey and conservation program by
any concerned entities.
ACKNOWLEDGMENT
This paper is a study of FishArk Philippines: Direction
for the Conservation of Native and Endemic Philippine
Freshwater Fishes, which is funded by the Department
of Science and Technology (DOST) through the
Philippine Council for Aquatic and Marine Research and
Development (PCAMRD). The authors would like to
convey their sincerest gratitude to the DOST for funding
the project; University of the Philippines Los Baños
Museum of Natural History; City of Tayabas, Quezon;
Luksong Alyansa para sa Bundok ng Banahaw Inc.;
UPLB Limnological Research Station: BV Labatos,OE
Matalog, RR Alvarez, RL Willy,N Salvador, B Lontoc,
and G Oldan for the logistics during the survey; UPLB
Institute of Biological Sciences: JC Gonzales, R de
Chavez, GR Ugaddan, and MVC Camacho; SEAFDEC/
AQD-Binangonan Freshwater Station: MRR Eguia, MLC
Aralar, E Aralar, and FA Aya.
REFERENCES
ANGERMEIER PL, DAVIDEANU G. 2004. Using
fish community to assess streams in Romania:
initial development of an index of biotic integrity.
Hydrobiologia 511: 65—78.
ANGERMEIER PL, WINSTON MR. 1999. Characterizing
fish community diversity across Virginia landscapes:
prerequisite for conservation. Ecol Appl 9: 335—349.
BEGON M, HARPER JL, TOWNSEND CR. 1996.
Ecology: Individuals, Populations and Communities,
3rd Ed. Blackwell Science, London. 1068p.
65
Philippine Journal of Science
Vol. 142 No. 1, June 2013
BRADFORD MJ, HEINONEN, JS. 2008. Low flows,
instream flow needs and fish ecology in small streams.
Can Water Resources J 33: 165—180.
BURNS JG, DI NARDO P, RODD FH. 2009. The role of
predation in variation in body shape in guppies Poecilia
reticulata: a comparison of field and common garden
phenotypes. J Fish Biol 75: 1144—1157.
CAGAUAN AG. 2007. Exotic Aquatic Species
Introduction in the Philippines for Aquaculture – A
Threat to Biodiversity or a Boom to the Economy? J
Environ Sci Manage 10: 48—62.
CORPUZ MN, PALLER VG, LABATOS BV, OCAMPO
PP. 2010. Ichthyofaunal survey in Bulusan Volcano
Natural Park. Paper presented at the 14th Southern
Luzon Zonal R & D Symposium and Planning
Workshop. Western Philippines University, Puerto
Princesa City, Philippines.
CORPUZ MN, PALLER VG, OCAMPO PP. 2011.
Diversity and distribution of freshwater fishes in Taal
Lake river systems in Batangas, Philippines. Paper
presented at the 15th Southern Luzon Zonal R & D
Symposium and Planning Workshop. Marinduque State
College, Boac, Marinduque, Philippines.
DIESMOS AC, LAGDA MC, TRESNADO NF, BROWN
RM. 2004. The snake from Taal. Haring Ibon 8—12.
GRIFFIN JE, OJEDA SR. 1992. Textbook of Endocrine
Physiology 2nd Ed. Oxford University Press, Inc., New
York. 351p.
GUERRERO RD III. 2002. Invasive aquatic animals in
the Philippines. special report on their impacts and
management. ASEAN Biodiversity (Oct-Dec 2002):
12—15.
HERDER F, FREYHOF J. 2006. Resource partitioning
in a tropical stream fish assemblage. J Fish Biol 69:
571—589.
KENNARD MJ, ARTHINGTON AH, PUSEY BJ,
HARCH BD. 2005. Are alien fish a reliable indicator
of river health? Freshwater Biol 50: 174—193.
KWAK TJ, PETERSON JT. 2007. Community indices,
parameters, and comparisons. In: Guy SC Brown ML
editors. Analysis and Interpretation of Freshwater
Fisheries Data. American Fisheries Society. Maryland.
p. 677—763.
LEGENDRE P, LEGENDRE L. 1998. Numerical
Ecology, 2nd English ed. Elsevier Science, B.V.
Amsterdam. 853p.
MAGURRAN A. 1988. Ecological Diversity and Its
Measurement. Princeton University, Princeton. 179p.
66
Paller et al.: Fish Assemblages in Tayabas River, Philippines
MAY JT, BROWN LR. 2000. Fish community structure
in relation to environmental variables within the
Sacramento River basin and implications for the greater
Central Valley, California. US Geological Survey
Report 00-247. Sacramento, California. 25p.
MILLS LS, SOULE ME, DOAK DF. 1993. The
keystone-species concept in ecology and conservation.
BioScience 43 (4): 219—224.
ONG PS, AFUANG LE, ROSELL-AMBALL RG. 2002.
Philippine Biodiversity Conservation Priorities, a
2nd iteration of the national biodiversity strategy and
action plan: final report. Department of Environment
and Natural Resources, Conservation International
Philippines, Biodiversity Conservation Program,
U.P. Center for Integrated and Development Studies,
Quezon City. p. 1—113.
[PAGASA] Philippine Atmospheric Geophysical
and Astronomical Services Administration. 2010.
Retrieved from http://www.pagasa.dost.org.ph on 02
April 2010.
PALLER VG, LABATOS BV, LONTOC BM, MATALOG
OE, OCAMPO PP. 2011. Freshwater fish fauna in
watersheds of Mt. Makiling Forest Reserve, Laguna,
Philippines. Philipp J Sci 140(2): 195—206.
PAUL MJ, MEYER JL. 2001. Streams in the urban
landscape. Annu Rev Ecol Syst 32: 333—365.
[PSDN] Philippine Sustainable Development Network.
2012. Retrieved from http://www.psdn.org.ph on 21
December 2012.
REDDING JM, PATIÑO R. 1993. Reproductive
Physiology. In: Evans DH. editor. The Physiology
of Fishes. CRC Press Inc. Boca Raton, Florida. p.
503—526.
ROSS ST. 1986. Resource partitioning in fish assemblages:
a review of field studies. Copeia. 352—388.
SHANNON CE, WEAVER W. 1949. The mathematical
theory of communication. Urbana, Univ. Illinois Press.
117p.
SHERVETTE VR, AGUIRRE WE, BLACIO E,
CEVALLOS R, GONZALEZ M, POZO F, GELWICK
F. 2007. Fish communities of a disturbed mangrove
wetland and an adjacent Tidal River in Palmar.
Ecuador. Estuar Coast Shelf Sci 72: 115—128.
SHINDE SE, PATHAN TS, RAUT KS, BHANDARE RY,
SONAWANE DI. 2009. Fish biodiversity of Pravara
River at Pravara Sangam District Ahmednagar, (M.S.)
India. World J Zool 4(3): 176—179.
SIMPSON EH.1949. Measurement of diversity. Nature
163: 688.
Philippine Journal of Science Paller et al.: Fish Assemblages in Tayabas River, Philippines
Vol. 142 No. 1, June 2013

TER BRAAK CJF, VERDONSCHOT P. 1995. Canonical

correspondence analysis and related multivariate
methods in aquatic ecology. Aquat Sci 57: 255—289.
[UPLB-LRS] University of the Philippines Los Baños-
Limnological Research Station. 2011. Freshwater
Fishes of Southern Luzon, Philippines. UPLB, Laguna,
Philippines. 100 p.
VESCOVI L, BERTEAUX D, BIRD D, SYLVIE DB.
2009. Freshwater biodiversity versus anthropogenic
climate change. UNESCO-WWAP, France. 18 p.
VORWERK PD, FRONEMAN PW, PATERSON AW.
2007. Recovery of the critically endangered river
pipefish, Syngnathus watermeyeri, in the Karlega
Estuary, Eastern Cape province. S Afr J Sci 103:
199—201.
WELCOMME RL. 1995. Relationships between fisheries
and the integrity of river systems. Regulated rivers:
Research and Management 11: 121—136.
WOLDA H. 1981. Similarity indices, sample size and
diversity. Oecologia 50: 4754.
XENOPOULOS MA, LODGE DM, ALCAMO J,
MARKER M, SCHULZE K, VAN VUUREN D.
2005. Scenarios of freshwater fish extinctions from
climate change and water withdrawal. Global Change
Biology, Vol. 11. Oxford, UK, Blackwell Publishing,
p. 1557—1564.
ZAMPELLA RA, BUNNELL JF. 1998. Use of reference-
site fish assemblages to assess aquatic degradation in
Pinelands Streams. Ecol Appl 8: 645—658.

67