PHILIPP AGRIC SCIENTIST ISSN 0031-7454

Vol. 96 No. 1, 75–85

March 2013

Morphometric and Morphomeristic Variations in Five Populations of

Indigenous Celebes Goby Glossogobius celebius (Perciformes:
Gobiidae) from Southern Luzon, Philippines
Mark Nell C. Corpuz1,*, Ma. Vivian C. Camacho2 and Pablo P. Ocampo2,3
Portion of the Masteral Thesis of the senior author. Funded by the DOST-Accelerating Science and Technology
Human Resources Development Program
1
Southeast Asian Fisheries Development Center/Aquaculture Department, Binangonan Freshwater Station,
Binangonan, Rizal
2
Animal Biology Division, Institute of Biological Sciences, College of Arts and Sciences, University of the Philippines
Los Baños (UPLB), College, Laguna 4031, Philippines
3
UPLB Limnological Research Station, College of Arts and Sciences, University of the Philippines Los Baños,
College, Laguna 4031, Philippines
*
Author for correspondence; e-mail: mark.corpuz@seafdec.org.ph

To elucidate the population structure and pattern of morphological divergence of the indigenous
Celebes goby Glossogobius celebius (Valenciennes, 1837), morphometric and morphomeristic
characters of this species were compared among populations collected from five study sites in
Southern Luzon, Philippines. Univariate analysis of variance confirmed significant differences
among populations for 20 out of 23 allometrically-transformed metric characters (P<0.05), while
nine meristic traits of five populations were comparatively homogeneous (P>0.05). Intrapopulation
variability was very low (6.81–7.21%) for each location. Multivariate analysis of variance
determined a very high significant group variability (P<0.05). Also, principal component analysis
(PC1=36.60%; PC2=18.34%) and canonical variate analysis (CV1=44.59%; CV2=37.30%) identified
the 2nd dorsal fin length, anal fin length, and the 2nd dorsal fin origin as the main significant
contributors of population differentiation. Derived classification functions correctly classified an
average of 68% individuals into their original groups. The canonical variate analysis plot exhibited
high isolation of lacustrine populations (Lake Taal and Laguna de Bay) from riverine populations
(Dampalit River, Alitaaw River and Pansipit River), with the latter having longer metric characters.
Likewise, squared Euclidean distance similarity using metric data clearly produced two clusters
representing both river and lake populations. This morphological divergence is suggested to be
environmentally-induced (phenotypic plasticity) primarily by the differences in flow rate and
temperature of the study sites.

Key Words: Celebes goby, Glossogobius celebius, Laguna de Bay, morphometric, riverine population

Abbreviations: CV – canonical variate, CVA – canonical variate analysis, DOST – Department of Science and
Technology, PC – principal component, PCA – principal component analysis, UPGMA – unweighted pair group
method with arithmetic mean

INTRODUCTION teleost is from the Family Gobiidae. Gobiids comprise

In the Philippines, a biodiversity rich country (Mallari et
al. 2001; Carpenter and Springer 2005), most of the
studies and initiatives on biodiversity had focused on
assessment of the conservation status of most terrestrial
and marine species. Few studies, however, have
evaluated the population status of freshwater fishes,
particularly the many diminutive indigenous and endemic
taxa with little commercial value (Ng et al. 1998; IUCN
2010). One of the most important species of freshwater
the great mass of gobioid fishes among all the freshwater
fish families documented in the Philippines (Herre 1924;
Herre 1927; Vallejo 1985). Among the gobiids in the
Philippines, Celebes goby (Glossogobius celebius) is said
to be one of the most successful species (Herre 1927;
UPLB-LRS 2011), but it is restricted to some isolated
lakes and rivers (UPLB-LRS 2011). The species is
demersal and highly carnivorous and can complete its life
history in freshwater, but its larvae can also occur in
brackish and marine waters (Froese and Pauly 2010). It is

The Philippine Agricultural Scientist Vol. 96 No. 1 (March 2013) 75

Morphological Variations in Glossogobius celebius Populations
usually included as by-catch and is considered of minor
commercial value because it is rarely seen in the market.
Locals, however, still consume it and utilize it as animal
feed.
There is a paucity of information on the population
of G. celebius mainly due to its remoteness, cryptic
behavior and diminutive adult size (Herre 1927; Larson
and Murdy 2001), hence, the current status of this
population is not fully well-documented (Froese and
Pauly 2010; Paller et al. 2011). Likewise, no study has
been conducted to elucidate the population structure of
this species in the different freshwater ecosystems in
Southern Luzon. Furthermore, continued threats from
pollution, erosion and the introduction of invasive
species, coupled with the lack of local concern, education
and sustainable protection, pose a level of ambiguity on
the survival of this goby (Bagarinao 2001; Guerrero III
2002; Paller et al. 2011). For these reasons, it is has
become increasingly significant to determine the current
status of G. celebius populations.
One very useful method in understanding fish
population structure is the identification of
interpopulation differentiation with different
biogeographical histories (Turan et al. 2004; Mejri et al.
2012). Linear measurements and patterns in body forms
are vital in identifying these discrete phenotypic stocks
(Gunawickrama 2007; Quilang et al. 2007; Hossain et al.
2010). Various studies in morphology such as meristic
and morphometric characters had proven to be effective
parameters to elucidate stock assessments and
relationships among fishes residing in different habitats
(Silva 2003; Coelho et al 2005; Turan et al. 2005, 2006;
Dominguez et al. 2009; Rodriguez et al. 2009; Mejri et
al. 2012). Morphometric variation results from
differences in developmental rates and the population’s
response to varying environmental conditions (Turan et
al. 2004; Burns et al. 2009). Developmental rates also
influence many population characters that are directly
related to population dynamics. Therefore, comparison of
the morphometric and meristic characters among G.
celebius populations with the aid of proper statistical
analyses can be used to clarify intra- and interpopulation
variations. This procedure would shed light on their
population structures and somehow describe the
evolutionary background of test goby and related
congeners. It is also important in defining the appropriate
conservation strategies (e.g., captive breeding program
for stock enhancement) and constructing plans for the
sustainable use of both the species and the environment
(Paller et al. 2011; UPLB-LRS 2011).
Our hypothesis is that morphological plasticity
occurs among the different populations of G. celebius in
five habitats in southern Luzon as a consequence of
geographical separation and exposure to opposing
environmental conditions. This study evaluated the
76
Mark Nell Corpuz et al.
metric and meristic variations of indigenous populations
of G. celebius from five collection sites in Southern
Luzon, Philippines, namely, Alitaaw River in Quezon,
Dampalit River in Laguna, Taal Lake and Pansipit River
in Batangas, and Laguna de Bay in Rizal, and aimed to
provide acceptable explanations for the differences.
MATERIALS AND METHODS
Collection Sites
A total of 179 G. celebius samples were collected from
five sites in Southern Luzon, Philippines, namely,
Dampalit River in Los Baños, Laguna; Laguna de Bay in
Cardona, Rizal; Alitaaw River in Quezon; Taal Lake in
Balete, Batangas; and Pansipit River in Agoncillo,
Batangas (Fig. 1). The geographic coordinates, brief
abiotic and habitat features of the collection sites, and the
composition and sample size of the populations are
presented in Table 1.
The Dampalit River is one of the mountain streams
of the Mount Makiling Protected Landscape. Surface
runoff from the forested surrounding highland and water
seepage are the major sources of water. The river ends in
Laguna de Bay. It has a cataract that is characterized by
clear and cool water flowing down the rapid bouldered
downstream with areas of highly steep walls covered
with epiphyte structures. The site offers ecotourism
potential (Makiling Center for Mountain Ecosystems
2010; personal observation).
Laguna de Bay, the largest inland water body in the
Philippines, has a total area of 900 km2. Its 285 km
shoreline delineates three distinct bays, namely: the west
bay, the central bay and the east bay. The central bay
(collection site) is close to the denuded hills with some
flat areas where rice is grown. It has a total area of 98.38
km2 with an average depth of 3.15 m. Its long shoreline
has numerous fish pens and cages (LLDA 2012;
Punongbayan 2005).
The Alitaaw River in Tayabas City, Quezon Province
is part of the Mount Banahaw-San Cristobal Protected
Landscape. The river initiates in Mount Banahaw,
traverses Tayabas City down to Lucena City before
draining to Tayabas Bay (Joji Roxas 2010, personal
communication). As a typical mountain stream, the water
flows over rocks and boulders, and the area is bordered
by secondary forest growth (higher altitudes), and
residential units (city proper).
Lake Taal is volcanic in origin and the third largest
lake in the Philippines, with a total area of 263.68 km2
and an average depth of 60.1 m (maximum of 198 m)
(Castillo and Gonzales 1976). The collection site is
situated on the eastern part of the lake, where locals
mostly rely on subsistence and artisanal fisheries for their
living.
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 Morphological Variations in Glossogobius celebius Populations Mark Nell Corpuz et al.

Table 1. Abiotic and habitat characteristics, sample sizes and mean standard length (SL) ± SE (range) of Glossogobius
celebius populations in five collection sites.
Abiotic and Habitat Features
Collection Sites
Mean Riffle Sample Mean SL
(Geographic Dominant Vegetation
Temperature Velocity Substrata Sizes (mm)
Coordinates) Observed
(°C) ± SE (m s-1) ± SE
Dampalit River Sandy + rocky +Algae, riparian macrophytes 39 62.80 ± 1.72
25.64 ± 0.28 1.40 ± 0.45
(14°06' N; 121°14' E) boulder perennial weeds (29♀, 10♂) (46.3–98.05)
Laguna de Bay Muddy + Algae + epiphytes + 36 61.11 ± 1.15
30.15 to 33.2* no data
(14°21' N; 121°13' E) biodetritus Eichhornia crassipes (29♀,7♂) (47.05–74.14)
Alitaaw River 36 61.33 ± 1.48
23.17 ± 0.80 1.80 ± 0.85 Boulder + Sandy Mosses, bryophytes
(14°05' N; 121°54' E) (26♀, 10♂) (46.67–82.98)
Lake Taal Sandy to muddy 36 59.99 ± 1.56
29.78 ± 1.95 no data Algae, epiphytes
(14°01' N; 121°05' E) + biodetritus (21♀, 15♂) (45.23–77.55)
Pansipit River Algae, submerged, riparian, 32 62.53 ± 1.58
26.50 ± 1.18 1.10 ± 0.65 Sandy to muddy
(13°55’ N; 120°56' E) and floating macrophytes (27♀, 5♂) (46.7–83.21)
*
Source: LLDA (2012)

fin origin (PDL2), prepelvic length (PPL), ventral fin to

Fig. 1. Map of Southern Luzon, Philippines showing the
collection sites of Glossogobius celebius:
Dampalit Stream (A), Laguna de Bay (B),
Alitaaw River (C), Lake Taal (D), and Pansipit
River (E).
The Pansipit River is a 9 km river that connects Lake
Taal and Balayan Bay (Mercene and Alzona 1990). Its
river bank is predominantly inhabited by perennial
weeds, riparian vegetation and residents in patchy
housing units. Water and land use patterns include
aquaculture, agriculture, transportation and irrigation.
Sampling
Mature specimens of not less than 45 mm (Larson and
Murdy 2001) were collected using seine net (~8.50 x 2.8
m with 1.2 x 1.2 mm mesh) and hook-and-line between
May 2010 and February 2011. Sexes were verified
through inspection of the genital papilla of the organisms.
The samples were preserved in 10% buffered
formaldehyde and then analyzed at the University of the
Philippines Los Baños-Limnological Research Station
(UPLB-LRS).
Morphometric and Meristic Data
Twenty-five morphometric characters were measured in
32–39 adult fish per population: total length (TL),
standard length (SL), predorsal length (PDL1), 2nd dorsal
anus (VFA), preanal length (PAL), caudal peduncle
length (CPL), caudal peduncle depth (CPD), body depth
at anal fin origin (BDAF), body depth at pelvic fin origin
(BDVF), 1st dorsal fin base (DFB1), 2nd dorsal fin base
(DFB2), pectoral fin length (PFL), anal fin base (AFB),
ventral/pelvic fin length (VFL), dorsal fin length (DFL),
anal fin length (AFL), caudal fin length (CFL) (Fig. 2A),
head length (H), head width (HW), head depth (HD), eye
diameter (E), pre-orbital length (Pre), and post-orbital
length (Post) (Fig. 2C).
The morphometric measurements were taken on the
left side of the body of the fish; otherwise, on the right
side of the body if the left side was damaged. Metric
characters were measured using a digimatic vernier
caliper. Measurements were rounded to the nearest 0.01
mm.
Nine meristic characters were counted, including the
number of longitudinal scale rows (LSR); the number of
pored scales or sensory papillae as described by Miller
(1984), i.e., from the beginning of the lateral line just
behind the head at the top of the operculum to the caudal
base; predorsal scales (scales along the dorsal midline
forward from the origin of the 1st dorsal fin) (PdS); 1st
dorsal fin spine (DS1); 1st dorsal fin rays (DR1); 2nd
dorsal fin spines (DS2); 2nd dorsal fin rays (DR2) anal fin
spines (AS); pectoral fin rays (PR); and caudal fin rays
(CR). Photos of the species and stereomicroscope were
used to count the meristic characters.
Statistical Analyses
Growth patterns in gobiid populations were observed to
be allometric (Mashiko and Yamane 1993) and to have a
strong correlation with fish size (Reist 1986). This size
effect was corrected prior to the comparison of
morphological variations among the different
geographical populations of G. celebius. The formula of
Elliot et al. (1995) was used in allometric adjustment of
the morphometric data set:

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Morphological Variations in Glossogobius celebius Populations
Fig. 2. Morphometric measurements from the left side of
the body (A), fins (B), and head (C) of
Glossogobius celebius (modified pictures from
Herre 1927). A: PDL2 – 2nd dorsal fin origin,
PDL1 – predorsal length, BDVF – body depth at
pelvic fin origin, BDAF – body depth at anal fin
origin, CPD – caudal peduncle depth, PPL –
prepelvic length, VFA – ventral fin to anus, CPL –
caudal peduncle length, PAL – preanal length, SL
– standard length, TL – total length; B: DFB1 –
1st dorsal fin base, DFB2 – 2nd dorsal fin base,
DFL – dorsal fin length, PFL – pectoral fin length,
VFL – ventral/pelvic fin length, AFB – anal fin
base, AFL – anal fin length, CFL – caudal fin
length; C: H – head length, E – eye diameter, Pre
– preorbital length, Post – post-orbital length, HD
– head depth
Mc = M (Ls / SL)b
where Mc is the size corrected measurement, M is the
observed measurement, Ls is the mean of the standard
length for all fish from all samples in each sampling site,
SL is the standard length of the fish, and b is the slope of
the regression of log10 M on log10 SL, for all specimens.
Standard length (SL) was employed as the basis of
transformation since a number of specimens have
damaged caudal fin. Standardization efficiency was
verified using correlation coefficient of multiple
determination (R2) between corrected data and SL
(P<0.05). Combined sexes were utilized in comparing
different populations. Sexes within each population were
78
Mark Nell Corpuz et al.
also compared prior to data analysis.
All data and statistical analyses were conducted
using Paleontological Statistics (PaST) v 2.07 (Hammer
et al. 2001). Assumptions for normality (Kolmogorov-
Smirnov test) and homogeneity (Levene’s test) of
variances were tested. Coefficient of variation in each
group was also determined to evaluate intrapopulation
morphological plasticity. One-way ANOVA was
employed to test if the means of metric measurements
were significantly different among the five G. celebius
populations (P<0.05). Likewise, unplanned multiple
comparison of metric characters was done using Tukey’s
HSD comparison test (P<0.05). Non-parametric Kruskal-
Wallis test was used to compare the median of the actual
distribution of meristic counts of five samples (P<0.05)
and multiple comparison was determined according to
Mann-Whitney test (P<0.05). Metric characters that
showed significance at less than 5% level were subjected
to multivariate analyses and cluster analysis. Descriptive
statistics for metric and meristic variables were also
computed for each site.
Squared Euclidean distance between means of
allometrically-transformed metrics was used as a measure
of (dis)similarity among the five groups. The
relationships between analyzed samples were derived
from the obtained distance matrix using an unweighted
pair group method with arithmetic mean (UPGMA)
clustering method (Sokal and Rohlf 2009).
Comparative analysis of five populations was also
explored using multivariate tests as complement of
univariate analysis. Principal component analysis (PCA)
was used to identify the metric and meristic variables that
contribute the most to the differences and to compare
overall morphological pattern among the five
populations. PCA was performed on 23 size-adjusted
morphometrics using the covariance matrix. Multivariate
analysis of variance (MANOVA) was performed to test
significant overall difference between groups according
to Pillai Trace statistic (P<0.05). Classification success
was derived from CVA and pairwise squared
Mahalanobis distances to assign individual fish to
recognized clustered groups. Group centroids were also
produced by CVA with 5% concentration ellipse level to
visualize associations among groups. Correct
classification rate (jackknifed) was assessed based on the
percentage of individuals correctly classified in the
original population.
RESULTS
Morphometric Measurements
Mean standard length and details of descriptive statistics
for each of the metric variables are presented in Table 2.
No significant correlation was found between size-
The Philippine Agricultural Scientist Vol. 96 No. 1 (March 2013)
Morphological Variations in Glossogobius celebius Populations Mark Nell Corpuz et al.

Table 2. Mean ± standard error of size-corrected metric characters of Glossogobius celebius collected from five study
sites.
Metric Dampalit Laguna Alitaaw Taal Pansipit
F Value
Variables (n=39) (n=36) (n=36) (n=36) (n=32)
In % of SL
H 19.28 ± 0.11a 18.43 ± 0.14b 18.52 ± 0.13b 18.08 ± 0.16b 18.07 ± 0.17b 12.28**
PDL1 24.66 ± 0.13a 23.96 ± 0.14b 24.30 ± 0.19a 23.31 ± 0.19c 24.43 ± 0.17ab 9.59**
PDL2 36.22 ± 0.19ab 35.04 ± 0.15c 35.57 ± 0.17bc 34.84 ± 0.19d 36.59 ± 0.24a 15.56**
PPL 19.83 ± 0.18ab 19.17 ± 0.24bc 19.88 ± 0.19ab 19.06 ± 0.18c 20.18 ± 0.18a 15.84**
VFA 15.01 ± 0.20a 14.73 ± 0.19a 14.80 ± 0.16a 13.98 ± 0.14b 14.63 ± 0.17a 4.94**
PAL 38.92 ± 0.19a 37.38 ± 0.24bc 38.08 ± 0.23ab 36.79 ± 0.24c 37.90 ± 0.25b 12.57**
CPL 15.03 ± 0.17ab 14.75 ± 0.16ab 14.67 ± 0.16ab 14.70 ± 0.17b 15.35 ± 0.19a 2.70*
CPD 6.70 ± 0.08a 6.37 ± 0.09b 6.68 ± 0.08a 6.42 ± 0.07ab 6.68 ± 0.07a 4.46**
BDAF 10.72 ± 0.10ab 10.32 ± 0.11b 10.67 ± 0.08ab 10.40 ± 0.15b 10.85 ± 0.11a 3.88**
BDVF 11.64 ± 0.15a 11.00 ± 0.15b 11.67 ± 0.16a 10.91 ± 0.16b 11.35 ± 0.14ab 5.25**
DFB1 8.44 ± 0.09a 8.24 ± 0.11ab 8.37 ± 0.12a 7.88 ± 0.10b 8.19 ± 0.08a 4.80**
DFB2 12.11 ± 0.16ab 11.38 ± 0.16b 12.18 ± 0.13a 11.50 ± 0.15b 11.97 ± 0.20ab 5.23**
PFL 15.88 ± 0.13a 15.08 ± 0.16b 15.76 ± 0.13a 14.98 ± 0.17b 15.79 ± 0.19a 7.78**
AFB 9.50 ± 0.16a 9.24 ± 0.14a 9.40 ± 0.13a 9.17 ± 0.10a 9.64 ± 0.15a 1.87NS
VFL 12.43 ± 0.18a 12.25 ± 0.19ab 12.78 ± 0.19a 11.56 ± 0.19b 12.22 ± 0.17a 5.72**
DFL 20.42 ± 0.27a 19.83 ± 0.25a 20.78 ± 0.32a 18.57 ± 0.33b 19.81 ± 0.34a 7.94**
AFL 18.29 ± 0.22a 17.95 ± 0.23a 18.08 ± 0.22a 16.98 ± 0.25b 17.61 ± 0.29ab 4.58**
CFL 16.52 ± 0.20a 15.86 ± 0.17ab 16.35 ± 0.21a 15.38 ± 0.18b 15.89 ± 0.23ab 5.22**
HW 13.06 ± 0.11a 12.44 ± 0.09b 13.12 ± 0.12a 12.36 ± 0.13b 12.93 ± 0.16a 9.63**
HD 10.80 ± 0.10a 10.31 ± 0.11b 10.78 ± 0.13a 10.29 ± 0.09b 10.52 ± 0.14ab 4.38**
E 4.15 ± 0.03a 4.07 ± 0.04a 4.04 ± 0.03a 4.09 ± 0.05a 4.060 ± 0.04a 1.02NS
Pre 5.16 ± 0.13a 5.06 ± 0.10a 4.83 ± 0.11a 5.01 ± 0.08a 5.09 ± 0.11a 1.32NS
Post 9.06 ± 0.13a 8.76 ± 0.14ab 8.67 ± 0.10ab 8.02 ± 0.14c 8.40 ± 0.10bc 9.92**
For each variable, means with the same superscript letter are not significantly different; *significant at 5% level; **significant at 1% level; NS not significant
at 5% level
SL – standard length, H – head length, PDL1 – predorsal length, PDL2 – 2nd dorsal fin origin, PPL – prepelvic length, VFA – ventral fin to anus, PAL –
preanal length, CPL – caudal peduncle length, CPD – caudal peduncle depth, BDAF – body depth at anal fin origin, BDVF – body depth at pelvic fin
origin, DFB1 – 1st dorsal fin base, DFB2 – 2nd dorsal fin base, PFL – pectoral fin length, AFB – anal fin base, VFL – ventral/pelvic fin length, DFL – dorsal
fin length, AFL – anal fin length, CFL – caudal fin length, HW – head width, HD – head depth, E – eye diameter, Pre – preorbital length, Post – post-
orbital length

adjusted data and standard length (P>0.05), which imply
that the allometric standardization method eliminated the
size effect in the data set. Heterogeneity of standard
length was observed, with population from Dampalit
being the largest (62.80 ± 1.72 mm) and the Taal
population being the smallest (59.99 ± 1.56 mm).
Significant variations were found in 20 out of 23
allometrically transformed metric characters (ANOVA,
P<0.05).
Sexual dimorphism within every population showed
significant differences in only 3 out of 23 (DFL, PAL and
AFL) metric characters. Sexes from each group also
failed to deviate in PCA (overlapping scores), and were
significantly comparable in Hotelling’s T2 ­test of
MANOVA (P>0.05). Furthermore, all samples consisted
mostly of females; hence, the sex was not included as an
independent factor in the analysis.
Small coefficients of variation or relative variations
were computed for 23 morphometric measurements
(range: 2.59–15.18%). Furthermore, relative variation
within Taal (7.21%), Pansipit (6.99%), Laguna (6.98%),
Dampalit (6.92%) and Alitaaw (6.81%) were relatively
low and of the same magnitude.
Principal component analysis for significant metric
characters extracted from variance-covariance matrix
showed that PC 1, PC 2 and PC 3 accounted for 36.60%,
18.34% and 8.79% of the variation, respectively, which
gave a cumulative variation of 63.73% for the first three
PCs. The most significant variable weightings on PC 1
(in order of importance) were from DFL, PAL and AFL
(all loaded positively). PC 2 identified DFL, AFL, PAL
and PDL2 as the main contributors of variations, and PC
3 recognized PDL2, CPL, DFL and CFL as its top
loadings (Table 3). PCA plot scores of the first against
the second axes, however, did not clearly visualize the
separation of the groups (Fig. 3), similarly with the third
against the second PC axes (not shown).
Significant morphological differentiation in five
samples (Pillai Trace=1.30, F(80, 632) =3.79, P<0.01) was
determined, with a cumulative variation of approximately
82% for the first two canonical axes. Longer ventral
region, distance from the snout to the 2nd dorsal fin, and
2nd dorsal fin length (PAL, PDL2 and DFL loaded
positively) explained the 44.59% variation in CV1 that
significantly differentiated the stock of Taal and Laguna
from that of Pansipit and Dampalit, with the riverine G.
celebius being larger for such morphometric
measurements. Alitaaw group scores (centroid)
intermediated within the plot (Fig. 4). In CV2, 37.30% of
the total variation was explained primarily by the
relatively longer head measurements and median fin
lengths (H, DFL and AFL loaded positively), which

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Morphological Variations in Glossogobius celebius Populations Mark Nell Corpuz et al.

Table 3. Relative eigenvalues, percentage of variation

and weights of significant metric variables of the first
three principal components for total samples of
Glossogobius celebius collected from five sites. Most
significant weights of PC 1, PC 2 and PC 3 are set in
bold font.
PC 1 PC 2 PC 3
Eigenvalues 9.15 4.58 2.20
% Variation 36.60 18.34 8.79
Eigenvector
H 0.20 0.20 0.02
PDL1 0.19 0.26 0.17
PDL2 0.13 0.32 0.52
PPL 0.23 0.27 (-)0.14
VFA 0.16 0.04 0.14
PAL 0.35 0.38 0.04
CPL (-)0.05 (-)0.05 0.46 Fig. 4. Population centroids (5% ellipses) and plot of first
CPD 0.11 (-)0.02 (-)0.07 and second canonical variate (CV) scores of size-
BDAF 0.11 0.08 (-)0.04 adjusted metric characters obtained from five
BDVF 0.22 0.08 (-)0.23 Glossogobius celebius populations. Separation of
DFB1 0.07 (-)0.05 0.11 different stocks (left and right reading) is evident
DFB2 0.22 (-)0.08 (-)0.11 in this plot. Dampalit = Dam (▲); Laguna = Lag
PFL 0.22 (-)0.02 0.27 (▼); Alitaaw = Tay (○); Taal = Taal (□); Pansipit =
VFL 0.28 0.12 (-)0.05 Pan (■).
DFL 0.43 (-)0.61 0.35
AFL 0.33 (-)0.38 (-)0.16 caused divergence in CV scores between Dampalit and
CFL 0.29 (-)0.07 (-)0.35 Pansipit as well as between Taal and Laguna. Pairwise
HW 0.17 0.01 (-)0.11 comparison using squared Mahalanobis distance also
HD 0.16 0.09 (-)0.12 validated this group variability (P<0.05).
Post 0.13 0.11 (-)0.03 The percentage of individuals that were correctly
H – head length, PDL1 – predorsal length, PDL2 – 2nd dorsal
classified into their a priori or original population was
fin origin, PPL – prepelvic length, VFA – ventral fin to anus,
PAL – preanal length, CPL – caudal peduncle length, CPD –
moderately high (mean = 68%) (Table 4), with the
caudal peduncle depth, BDAF – body depth at anal fin origin, highest percentage of correct assignment in the Taal
BDVF – body depth at pelvic fin origin, DFB1 – 1st dorsal fin group (78%). The Laguna, Pansipit, Dampalit and
base, DFB2 – 2nd dorsal fin base, PFL – pectoral fin length, Alitaaw samples had correct classifications rates of about
VFL – ventral/pelvic fin length, DFL – dorsal fin length, AFL – 72%, 69%, 64% and 58%, respectively. Misclassification
anal fin length, CFL – caudal fin length, HW – head width, HD rate was very low (mean = 8%; range 0–25%).
– head depth, Post – post-orbital length A dendrogram from Euclidean (dis)similarity in the
five populations based on transformed metric data set
produced distinct clustering of the populations. The
analysis revealed a strong similarity between the metric
data of the Taal and Laguna populations. These lake
populations were isolated from three other similar
populations residing in riverine habitats, which resulted
in the formation of two distinct clusters (Fig. 5). The
clusters were then represented by lacustrine stocks (Taal
and Laguna) and riverine stocks (Dampalit, Alitaaw and
Pansipit) in another, presenting a degree of variability
between such groups.

Fig. 3. Plot of first and second principal component (PC)
scores for size-adjusted metric characters of
Glossogobius celebius sampled from five study
sites: Dampalit = (▲); Laguna = (▼); Alitaaw =
(○); Taal = (□); Pansipit = (■).
Meristic Counts
Scale counts and frequency of fin rays are given in
Tables 5 and 6, respectively. Longitudinal scales row
(27–31), predorsal scales (15–19), pectoral fin (15–20)
and caudal fin rays (14–19) had the most frequent
observations among the characters, whereas the first
dorsal fin showed consistent number of six spines for all

80 The Philippine Agricultural Scientist Vol. 96 No. 1 (March 2013)

Morphological Variations in Glossogobius celebius Populations Mark Nell Corpuz et al.

Table 4. Direct and percentage of classification (in parentheses) based on canonical variate analysis (CVA) of size-
corrected metric characters of Glossogobius celebius from five different sites. The analysis correctly assigned a mean
of about 68% to their original group.
A posteriori group
A priori group
Dampalit Laguna Alitaaw Taal Pansipit Total
Dampalit 25 (64.10) 5 (12.82%) 5 (12.82%) 1 (2.56%) 3 (7.69%) 39
Laguna 4 (11.11%) 26 (72.22%) 5 (13.89%) 1 (2.78%) 0 (0%) 36
Alitaaw 4 (11.11%) 5 (13.89%) 21 (58.33%) 3 (8.33%) 3 (8.33%) 36
Taal 0 (0%) 4 (11.11%) 2 (5.56%) 28 (77.78%) 2 (5.56%) 36
Pansipit 0 (0%) 0 (0%) 8 (25.0%) 2 (6.25%) 22 (68.75%) 32
Total 33 40 41 35 30 179

the samples. Anal fin (I, 7–9) and second dorsal fin (I, 8–
Table 5. Frequency distribution of scale counts of 9) had rather conserved meristics. Results of Kruskal-
Glossogobius celebius from five study sites. The most
Wallis and Tukey’s HSD comparison test for each
frequent (mode) observation is set in bold font.
meristic variable revealed no significant differences
Collection Site
among the five populations (P>0.05). Within-population
DampalitNS LagunaNS AlitaawNS TaalNS PansipitNS variability for meristic characters was very low (mean
Longitudinal Scale Rows coefficient variation = 3.25%; range: 0 –5.87%).
27 1 1 0 0 1
28 8 10 10 3 8
29 14 9 13 16 14 DISCUSSION
30 14 15 12 15 7
31 2 1 0 1 0
There was low intra-population plasticity in the metric
Predorsal Scales
15 5 15 3 10 7 and meristic attributes of G. celebius as inferred by a
16 15 13 25 17 15 small coefficient of variation from every population. This
17 9 7 6 7 6 low variability within samples was also observed by
18 1 0 0 1 2 Quilang et al. (2007) in four silver therapon
19 1 1 0 0 0 (Leiopotherapon plumbeus) populations in the
NS
not significant (P<0.05) in Kruskal-Wallis test by collection sites Philippines.
Table 6. Frequency distribution of fin ray counts of The results of significant metric differences among
Glossogobius celebius from five study sites. The most the five populations and the relatively high correct
frequent (mode) observations are set in bold font. classification percentage indicate inter-population
Roman numerals represent spinous fins, while Arabic divergence, albeit with no pronounced geographical
numerals represent soft fin rays. pattern. We hypothesized that geographic isolation
Collection Site between populations promotes the differentiation in
DampalitNS LagunaNS AlitaawNS TaalNS PansipitNS morphological characters, but discreteness occurred even
in groups thriving in hydro-spatially connected habitats
Caudal Fin such as Pansipit and Taal, as well as in Dampalit and
14 0 0 0 1 0
Laguna. Instead, clustering of the lacustrine populations
15 3 3 4 2 4
16 24 27 25 31 26 from that of the riverine populations was more clearly
17 3 4 2 0 1 evident in the distribution of CV scores (Fig. 4) and in
18 4 2 3 2 0 the Euclidean distance dendrogram (Fig. 5).
19 0 0 1 0 0 The variation in population could be due to drift or to
Pectoral Fin the different environmentally-induced selective regimes
15 2 2 1 4 1 brought about by the type of habitats (Slatkin 1985;
16 33 28 31 25 30 Swain and Foote 1999) where the goby lives. Notably,
17 1 4 1 5 0 the riverine G. celebius is generally adapted to relatively
18 3 2 1 1 1
cold and turbulent waters (Table 1), whereas the goby
19 0 0 1 1 0
20 0 0 1 0 0 populations from Taal Lake and Laguna de Bay are
Anal Fin (Spine/Rays) lacustrine inhabitants and are well-acclimatized in a
I/7 3 1 4 0 0 relatively warm and calm lentic condition (Table 1;
I/8 33 32 31 34 31 Rabanal et al. 1960; Castillo and Gonzales 1976).
I/9 3 3 1 2 1 Hypothetically, the G. celebius populations may have
First Dorsal Fin undergone phenotypic divergence as the species under
VI 39 36 36 36 32 investigation becomes adapted to a variety of diets,
Second Dorsal Fin (Spine/Rays) niches, environmental conditions and related selective
I/8 4 4 2 6 0
pressures in their respective habitats (Mayr 1963; Swain
I/9 33 30 34 28 32

The Philippine Agricultural Scientist Vol. 96 No. 1 (March 2013) 81

Morphological Variations in Glossogobius celebius Populations
Fig. 5. A dendrogram from unweighted pair group
method with arithmetic mean (UPGMA)
clustering of squared Euclidean distances
showing the phenetic relationship of five
Glossogobius celebius populations based on
significant size-corrected morphometric
characters. Lacustrine populations were
evidently clustered from the riverine
populations.
and Foote 1999). This concept is in consonance with the
cause of morphological discrepancies observed in the
populations of Liza abu (Turan et al. 2004), Clarias
gariepinus (Turan et al. 2005) and Labeo calbasu stocks
(Hossain et al. 2010).
We predicted that the phenotypic plasticity observed
in G. celebius could be attributed to the flow rate and the
temperature, which are some of the most contrasting
abiotic features between riverine and lacustrine habitats.
In this study, the environmental influence took effect on
the modifications of myo-skeletal developmental patterns
in the median fins in response to environmental
pressures. The longer 2nd dorsal fin (DFL) and anal fin
(AFL) of riverine goby compared with that of lake goby
intrinsically developed as an adaptive mechanism to
improve hydrodynamic efficiency. The elongated and
posteriorly-located 2nd dorsal fin (PDL2) and anal fin
(PAL) play a vital function in stabilizing the fish in mid-
water, and in the swift forward and lateral propulsion
(thrust) against fast-flowing water medium (Lauder and
Drucker 2004; Walker 2004). In fish species, the dorsal
fins also serve as ‘pivot’ for swimming muscles to
facilitate turning or rotating maneuvers (Drucker and
Lauder 2005; Mejri et al. 2012).
82
Mark Nell Corpuz et al.
It is widely known that temperature has a significant
impact in the metabolism and in the development of
muscles and bones of ectothermic species like fish.
(Wimberger 1992; Imre et al. 2002). Moreover,
temperature affects the contractile and erectile responses
of median fins, which have main roles in controlled
locomotive and swimming behavior of fishes (Fuiman
and Batty 1997). Although this study showed
morphometric variations in populations reared in varying
ambient temperatures, we are still inconclusive in the
significant influence of low temperature in elongated and
posteriorly-located median fins of riverine groups. The
thermal conditions of the environments alone may not
supply a comprehensive explanation for the recorded
differentiation, but rather requires further limnological
studies (multiple agents of selection) at larger temporal
scale to assess its effect on the morphology of the fins.
Diet composition analysis should also be integrated to
correlate the ramification of food selection regime on
intra- or inter-population variations.
Another interesting observation is the position of CV
scores and the centroid of the Alitaaw group in the
middle of the CV plot, between riverine and lacustrine
populations from Taal and Laguna. Alitaaw
morphometry seems to partly intermingle with that of
other populations despite the occurrence of geographical
barriers (mountain massifs), oceanic separation and its
distance from the other sites. The geographic isolation of
ancestral taxon caused by volcanic and tectonic
phenomenon (Punongbayan 2005) may account for this
magnitude of phenotypic variation. If this isolation
continues, the Alitaaw group (or also the other groups)
may attain a distinct morphometry and reproductive
behavior evolving into a reproductively isolated
(allopatric) population (Mayr 1963) under different
habitat conditions. This change can happen given the fact
that G. celebius and other gobiids are diminutive, poor
swimmers, benthic dwellers, and have short pelagic
larval stage (Herre 1927; UPLB-LRS 2011; Mejri et al
2012). Future studies regarding the status of Alitaaw
stock should be tackled, which could provide valuable
implications on the possible evolutionary radiation and
biogeographical histories of G. celebius and related
congeners in Southern Luzon.
In spite of slight variations, all the evaluated meristic
parameters were significantly comparable among the
populations (P>0.05) and still in accordance with the
accepted range of counts of G. celebius as reported by
Herre (1927), Akihito and Meguro (1975), and meristic
characteristics presented by Froese and Pauly (2010).
Homogeneity of these taxonomically important
characters was also observed in Arius jella
(Gunawickrama 2007), Algansea tincella (Rodriguez et
al. 2009), Toxotes jaculatrix and T. chatareus (Simon et
al. 2010). Being one of the most conserved characters of
The Philippine Agricultural Scientist Vol. 96 No. 1 (March 2013)
Morphological Variations in Glossogobius celebius Populations
fish, meristic characters are important diagnostic
characters in evaluating taxonomic position. Meristic
attribute is also generally determined at the early stage of
morphogeny and remains stable throughout the fish life
(Camacho et al. 2004). In some studies, however,
environmental factors such as temperature (Sfakianakis et
al. 2011) and salinity (Camacho et al. 2004) could
influence the meristic features of fishes during their
sensitive developmental stages. Small clinal differences
observed in gross fin and scale counts may not open a
room for the revision of its taxonomical position, but
rather a supposition that the larval stages had been
exposed to various environmental conditions (Nakamura
2003; Camacho et al. 2004; Sfakianakis et al. 2011), or
the differences could be due to damage in the fins and
scales during fish handling.
Comparative interpopulation variation of indigenous
G. celebius was investigated in the Philippines for the
first time. This research demonstrated the efficiency of
multivariate morphometric analyses to explain the
population structure of G. celebius in the context of their
morphological differences. The results of the study could
serve as valuable baseline information of G. celebius
populations for further understanding of its biology,
which is essential in stock enhancement program and
related conservation management of this species. For the
purpose of captive breeding, groups that are
morphologically different and tend to be aggressive or
oppressive to other stocks because of these differences
should be managed separately (Silva 2003). This research
should be complemented with cytogenetic, isozyme, and
molecular analyses to further assess the fish
morphological differentiation.
ACKNOWLEDGMENTS
We gratefully acknowledge the funding for this study
given by the Department of Science and Technology and
the Philippine Council for Aquatic and Marine Research
and Development through the Accelarating Science and
Technology Human Resources Development Program.
We thank the municipal governments of Agoncillo,
Belete, Los Baños, Pansipit, and Tayabas for allowing us
to conduct our studies in their areas; OE Matalog, RR
Alvarez and RL Willy for help in catching, measuring,
counting and identifying fishes; UPLB-LRS personnel
for the technical support; the UPLB Institute of
Biological Sciences: MQ Diaz, JC Gonzales, E de
Chavez, GR Ugaddan, and VGV Paller; and SEAFDEC/
AQD: MR Eguia, ML Aralar, E Aralar, FA Aya and J
Valera for logistical supports.
The Philippine Agricultural Scientist Vol. 96 No. 1 (March 2013)
Mark Nell Corpuz et al.
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