Marine Biology Research, 2012; 8: 802
814

ORIGINAL ARTICLE

Comparison of different otolith shape descriptors and morphometrics

for the identification of closely related species of Lutjanus spp. from the
Persian Gulf

ZAHRA SADIGHZADEH1*, VICTOR M. TUSET2, TOORAJ VALINASSAB3,

MOHAMMAD R. DADPOUR4 & ANTONI LOMBARTE2
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Marine Biology Department, Faculty of Marine Science & Technology, Islamic Azad University, Tehran, Iran; 2Institut de
Cie`ncies del Mar (CSIC), Barcelona, Catalonia, Spain; 3Department of Resource Management, Iranian Fisheries Research
Organization (IFRO), Tehran, Iran; and 4Department of Horticultural Sciences, Faculty of Agriculture, University of Tabriz,
Tabriz, Iran

Abstract
The anatomical and morphometric (shape indices, contour descriptors and otolith weight) characterizations of sagittal
otoliths were investigated in 11 species of Lutjanus spp. inhabiting the Persian Gulf. This is the first study that compares the
efficiency of three different image analysis techniques for discriminating species based on the shape of the outer otolith
contour, including elliptical Fourier descriptors (EFD), fast Fourier transform (FFT) and wavelet transform (WT). Sagittal
otoliths of snappers are morphologically similar with some small specific variations. The use of otolith contour based on
wavelets (WT) provided the best results in comparison with the two other methods based on Fourier descriptors, but only
the combination of the all three methods (EFD, FFT and WT) was useful to obtain a robust classification of species. The
species prediction improved when otolith weight was included. In relation to the shape indices, only the aspect ratio
provided a clear grouping of species.

Key words: Otolith, morphology, Fourier, wavelets, Lutjanidae, Persian Gulf

Introduction
The Persian Gulf is a semi-enclosed water body
connected to the Oman Sea through the strait of
Hormuz, which is 56 km wide at its narrowest point
and has an average depth of 35 m (Reynolds 1993).
The Oman Sea has an average depth of 700 m and is
connected to the Indian Ocean through the Arabian
Sea (Valinassab et al. 2006). Both are subtropical
water masses located chiefly between 24
308N and
49
628E. The dominant large-scale current is a
counter-clockwise movement, whereby less saline
and less dense water enters at the strait of Hormuz
at the surface and more saline and denser water leaves
the area at the bottom (Hunter 1983; Reynolds
1993). After oil, fisheries represent the second most
important natural resource, and the most important
renewable natural resource in the region (Carpenter

et al. 1997). A total of 12 species of snappers have

been identified along the Iranian coast of the Persian
Gulf and the Oman Sea, including Lutjanus johnii
(Bloch, 1792) and Lutjanus malabaricus (Bloch &
Schneider, 1801), the most abundant species (Valinassab et al. 2006, 2010). They play an important role
in the ecology system of the reefs, mangrove swamps
and seagrass beds (Aiken 1993; Appeldoorn &
Meyers 1993; Baisre 1993; Claro et al. 2001).
However, the biological and ecological knowledge of
snappers of the area is scarce and is mainly focused on
Lutjanus johnii (Valinassab 2006, 2010).
Sharks and other large fishes are probably their
main predators, but due to the lack of adequate
morphological knowledge of their sagittal otoliths it
was not possible to identify them in the stomach
content where they represent the main remnants of

*Correspondence: Zahra Sadighzadeh, Marine Biology Department, Faculty of Marine Science & Technology, Islamic Azad University,
Tehran, 1477893855, Iran. E-mail: zahrasadighzadeh@yahoo.com
Published in collaboration with the University of Bergen and the Institute of Marine Research, Norway, and the Marine Biological Laboratory,
University of Copenhagen, Denmark

(Accepted 5 April 2012; Published online 14 August 2012; Printed 24 August 2012)
ISSN 1745-1000 print/ISSN 1745-1019 online # 2012 Taylor & Francis
http://dx.doi.org/10.1080/17451000.2012.692163

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Identification of snappers using otoliths

the fish prey (Harkonen 1986; Smale et al. 1995;
Volpedo & Echeverra 2000; Assis 2003; Campana
2004; Tuset et al. 2008).
The otolith is an acellular concretion compound,
mainly of calcium carbonate, crystallized in aragonite
and other inorganic salts, which develop over a
protein matrix (Carlstrom 1963; Blacker 1969; Degens et al. 1969; Cermeno et al. 2006). They are
located in the inner ear of vertebrates in close
association with the sensitive maculae of the labyrinthine compartments (Cordier & Dalcq 1954;
Weichert & Prech 1981; Hildebrand 1988; Jobling
1995). The otolith shape has a high degree of interspecific variation, which can be used to identify the
species (Nolf 1985; Harkonen 1986; Smale et al.
1995; Volpedo & Echeverra 2000; Assis 2003;
Campana 2004; Tuset et al. 2008). Several methods
such as Fourier analysis, wavelets and curvature scale
space analysis have been used for automatic identification of species (Parisi-Baradad et al. 2005; Piera

803

et al. 2005; Mart-Puig et al. 2010; Capoccioni et al.

2011; Tuset et al. in press). Our study intends to
represent a useful method to recognize snapper species
in the stomach content of their predators using
otoliths. Therefore, the aims of this article are (i) to
characterize morphologically and morphometrically
the sagittal otoliths of snappers of Lutjanus spp. of the
Persian Gulf and (ii) to test and compare three
different methods (elliptic fourier descriptors, fast
Fourier transform and wavelet transform) to discriminate species using otolith contour and otolith weight.
Materials and methods
Sampling
Eleven species of snappers of the genus Lutjanus were
collected from the Strait of Hormuz (Figure 1)
with the help of local fisheries. These included
Lutjanus argentimaculatus (Forsskal, 1775) (n 13),
Lutjanus ehrenbergii (Peters, 1869) (n 61), Lutjanus

Figure 1. Map of the Persian Gulf (NE Indian Ocean) showing the area from where Lutjanus spp. were collected.

804 Z. Sagighzadeh et al.

erythropterus Bloch, 1790 (n 9), Lutjanus
fulviflamma (Forsska l, 1775) (n 11), Lutjanus johnii
(n 93), Lutjanus lemniscatus (Valenciennes, 1828)
(n 23), Lutjanus lutjanus Bloch, 1790 (n 23),
Lutjanus malabaricus (n 47), Lutjanus rivulatus
(Cuvier, 1828) (n 12) and Lutjanus russellii
(Bleeker, 1849) (n 32). Total fish length (TL in
cm) was measured for all individuals. Their otoliths
were removed, washed, dried and stored in plastic
vials. They were also weighed (OW) to the nearest 0.01
g. Due to lack of sufficient specimens of Lutjanus
bengalensis (Bloch, 1790), this species was omitted
from the mathematical analysis.

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Morphometry and morphology of otolith sagittae

Otoliths from the left side of the fish were orientated
with the inner side (sulcus acusticus) upwards and
rostrum to the right for digitizing using a microscope
attached to an image analyser with a magnification
depending on the size of the otolith. All images were
captured with an embedded 1-mm scale which was
used to avoid magnification effects in the morphometric calculations. Otolith length (OL in mm),
height (OH in mm), area (OA in mm2) and perimeter
(OP in mm) (Figure 2) were measured using ImagePro Plus version 4.1.0 software (Media Cybernetics
Inc.). Shape indices were obtained by combining size
parameters (Russ 1990; Tuset et al. 2003): aspect
ratio (OH/OL), compactness (OP2/OA) and rectangularity (OA/(OL*OH)), applying an ANOVA followed by Student Newman
Keuls test for multiple
comparisons of means at a 5% significance level
(P B0.05). Descriptive statistics of fish and otolith
variables by species are given in Table I. The otolith
descriptions follow Tuset et al. (2008).

Elliptical Fourier descriptors (EFD)

The otolith outline was extracted by using the
Shape program (Iwata & Ukai 2002). The software
generates 20 harmonics, using elliptical Fourier
descriptors (EFD), whose combinations allow
reconstructing of the otolith contour (Kuhl &
Giardina 1982; Tracey et al. 2006; Farias et al.
2009). Each harmonic was composed of four coefficients and therefore 80 coefficients were obtained for
each individual. The Shape package automatically
normalizes the contour in relation to the first
harmonic and consequently they become invariant
to size, rotation and starting point, which causes the
degeneration of the three first coefficients to fixed
values: a1 1, b1 c1 0 (Kuhl & Giardina 1982;
Iwata & Ukai 2002). For this reason, each otolith
contour was drawn by only 77 coefficients. The first
nine harmonics (FDs 4
12) were used to rebuild the
outline attaining more than 90% of the shape
variation. The minimum number of harmonics is
assembled by plotting the percentage of accumulated
variance explained by the nth harmonic vs. the
harmonic number, n (Stransky et al. 2008).
Fast Fourier transform (FFT)
The otolith silhouette was traced in a counter
clockwise direction and digitized into 512 x
y
equidistant coordinates by using the tip of the otolith
rostrum as a common starting point for the coordinates (Age & Shape Software, Infaimon SL, Spain)
of the software Image Pro Plus. A fast Fourier
transform (FFT) was calculated as a Cartesian
FFT. The Cartesian FFT uses the x
y coordinates
as complex numbers (a  ib), where a is the real
component and ib the imaginary component, representing the amplitudes of the cosine and sine waves,

Figure 2. Mesial surface of sagittal otoliths of Lutjanus spp. illustrating features considered.

Table I. Descriptive statistics of fish and otolith variables for Lutjanids from Persian Gulf. Larg, Lutjanus argentimaculatus; Lehr, L. ehrenbergii; Lery, L. erythropterus; Lful, L. fulviflamma; Ljoh, L.
johnii; Llem, L. lemniscatus; Llut, L. lutjanus; Lmal, L. malabaricus; Lriv, L. rivulatus; Lrus, L. russellii; letters indicate groups from SNK test.
Larg
(n13)

Lehr
(n61)

Lery
(n 9)

Lful
(n 11)

Ljoh
(n93)

Llem
(n23)

Llutj
(n23)

Lmal
(n47)

Lriv
(n12)

Lrus
(n 32)

min
max
423
802
146
260
316
523
176
260
167
754
298
514
153
232
235
732
405
667
150
372
mean9sd
648.2999.5
203.1922.5
370.7961.7
206.3927.9
364.09116.0
379.89 61.7
195.7919.7
317.6985.9
484.9984.0
250.8954.4
Area
min
max 67.83
174.66
18.15
40.12 61.57
113.06
22.11
36.70
30.27
313.09
44.49
92.36
19.5
37.35
39.39
359.45
98.77
175.65
15.54
58.73
mean9sd 123.66931.62 29.5695.16
74.52915.59
27.6795.21
98.13947.59
63.44914.68
29.4495.31
86.33951.00 119.64923.44 34.60911.18
Height
min
max
7.40
12.53
3.79
5.68
7.67
10.02
4.27
5.51
5.10
15.46
5.87
8.73
3.93
5.54
5.94
17.31
9.34
12.51
3.46
6.99
mean9sd
10.4591.62
4.7790.44
8.3990.71
4.7390.50
8.6991.99
7.0690.88
4.8390.44
8.8691.88
10.4191.01
5.1490.88
Length
min
max 12.74
20.49
6.54
10.42
11.68
16.32
7.27
9.82
8.36
28.83
10.24
15.55
6.83
9.82
9.23
30.16
14.55
19.48
6.50
12.50
mean9sd
16.9892.42
8.6390.83
12.8791.40
8.1990.84
15.1193.84
12.7891.50
8.5290.86
13.3293.34
15.9891.49
9.6191.66
Perimeter
min
max 37.50
61.62
18.89
29.24
34.10
46.53
20.52
26.52
24.10
92.74
29.15
46.67
19.37
28.45
25.95
86.19
42.86
58.28
18.93
33.60
mean9sd
52.2697.51
24.6692.34
37.2293.82
23.3592.35
43.27911.37
35.6294.68
24.4792.62
39.82910.07
48.4794.43
26.7194.24
Weight
min
max
0.16
0.83
0.03
0.11
0.15
0.38
0.04
0.11
0.05
2.20
0.10
0.40
0.04
0.11
0.10
2.45
0.33
0.86
0.02
0.16
mean9sd
0.4790.22
0.0690.02
0.2190.07
0.0790.03
0.3490.29
0.1890.08
0.0790.02
0.2990.36
0.4590.17
0.0790.03
Aspect ratio
min
max
0.55
0.69
0.51
0.61
0.61
0.68
0.53
0.64
0.53
0.66
0.51
0.62
0.54
0.60
0.57
0.72
0.63
0.68
0.47
0.58
mean9sd
0.6290.05a
0.5590.02b
0.6590.02d
0.5890.03c
0.5890.03c
0.5590.03b
0.5790.02bc
0.6790.03d
0.6590.02d
0.5490.02e
Compactness min
max 20.10
25.10
18.66
25.74
17.96
19.17
18.68
21.70
17.10
27.47
18.42
24.7
18.01
22.89
16.67
26.87
18.17
23.23
19.22
24.42
mean9sd 22.4591.59a 20.7591.39bc 18.7490.46e 19.8490.83cde 20.0791.75bcde 20.2291.40bcd 20.4991.17bcd 19.2391.92de 19.8591.57cde 21.3491.20b
Rectangularity min
max
0.64
0.72
0.68
0.74
0.66
0.70
0.68
0.73
0.68
0.73
0.67
0.73
0.69
0.73
0.67
0.73
0.68
0.73
0.66
0.71
mean9sd 0.6990.02ab
0.7190.01c
0.6890.01a
0.7190.02c
0.7190.01c
0.7090.02b
0.7190.01c
0.7090.01b
0.7190.01c
0.6890.01a

Identification of snappers using otoliths

p2
2
an0 ib0 n

805

respectively (Piera et al. 2005). The resultant 512 set

of complex numbers or descriptors were subsequently normalized for differences in otolith position
by setting the 0th descriptor to 0, and for size and
rotation of the otolith by dividing all the descriptors
with the first descriptor (Smith 1992; Bergenius
et al. 2005). The normalized descriptors (a?ib?)
were used to calculate the absolute value (harmonic)
of each descriptor (Christopher & Waters 1974):

Hormonic

The number of Fourier descriptors required was

determined when accumulated variance explained at
least 90%. Therefore, it was necessary to capture the
first 133 descriptors. Because the Cartesian descriptors are asymmetrical around the middle frequency,
both ends of the array are required in the reconstruction. Consequently, otolith outline shape was reconstructed by using the first two and last two
descriptors, the first three and last three descriptors,
and so on until the first and last 133 descriptors were
used (Bergenius et al. 2005).

Wavelet transform (WT)

The WT is based on expanding the contour into a

family of functions obtained as the dilations and
translations of a unique function known as a mother
wavelet (Mallat 1991). These functions describe
both in space and wave number, the most prominent
features of the curve. The signals of wavelets have
different amplitudes, hence small (low) wavenumbers (frequencies) are associated with a smoothly
varying contour, while large wavenumbers are associated with variation on a small spatial scale (ParisiBaradad et al. 2005, 2010). To obtain the otolith
contour, a total of 512 Cartesian coordinates on
each of the orthogonal projections, which is a
perpendicular image of an object (Schwarzhans
1980; Nolf 1985), were extracted using Age & Shape
program (Infaimon SL, Spain) software Image Pro
Plus. The wavelets four, five and six were selected as
the representatives of the otolith contour.

Statistical analyses

Normality of variables and homogeneity of variances

for each EFD coefficient, FFT harmonic or WT
point, were examined by using Kolmogorov

Smirnovs and Levenes tests, respectively. Data were
transformed using 1/x or log10, when the variables
did not meet the assumptions. In the case of the WT,
the number of variables was so high that those who did
not meet the assumptions were eliminated. Analysis of
covariance (ANCOVA) test (P B 0.05) was performed

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Fish length

806 Z. Sagighzadeh et al.

to determine the effect of species (the independent
variable) and fish length (the covariate) on EFD
descriptors. Species was considered as the main
factor and fish length as the covariate. When a
significant regression coefficient was detected and
homogeneity was observed among groups, then the
effect of fish length (TL) could be removed from
each measurement using this relationship:

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Oij; adj Oij b  TLij  MTLj ;

where Oij, adj otolith morphological measurement

of ith fish adjusted to mean fish length of group j;
Oij original otolith morphological measurement for
ith fish from group j; b  slope of the relationship Oij;
TLij, common to all groups; TLij fish length of ith
fish in group j; and MTLj average fish length within
group j. If slopes differed among groups, the correction for TL was made separately for each group using
the above equation by replacing the common slope
(b) with the group-specific slope (bj). These corrections were effective for scaling all morphological
variables of all otoliths to their predicted TL group
mean (Campana & Casselman 1993; Bolles & Begg
2000; Begg et al. 2001; DeVries et al. 2002;
Bergenius et al. 2005).
Otolith length was normalized to avoid allometric
effects of this parameter in the otolith weight, taking
into account the allometric relationship. A standardized fish length of 22.0 cm was used as the
reference for all species (Lombarte & Lleonart
1993; Lleonart et al. 2000; Tuset et al. 2006).
Multivariate analysis of variance (MANOVA) was
used to test the hypothesis of no difference in otolith
shape between species. A canonical discriminant
function analysis (DFA) was also carried out to verify
the differences between species using the abovementioned variables. In the first stage a single-factor
ANOVA was carried out to determine which variables
discriminate among species to rank the potential
predictors. For the second stage, multicollinearity
was avoided in the DFA analysis by obtaining a
matrix correlation and then, eliminating the variables
with a high correlation (r 0.5) and small F-score
from ANOVA. The hypothesis that the betweengroup covariances are equal was tested for all models.
The matrices were not homogeneous and the individual within-group matrices were used, thus a quadratic discriminant function was computed (Friedland
et al. 1994; Tuset et al. 2006). The classification
accuracy of each DFA was evaluated by leave-one-out
cross-validation (Nishimoto et al. 2010).

Results
Anatomical description
Morphologically, the sagittal otoliths of Lutjanus
spp. are similar with small specific variations
(Figures 3
5). For example, the otolith shape could
be pentagonal (L. argentimaculatus, L. erythropterus,
L. fulviflamma and L. russellii), elliptical
(L. bengalensis, L. ehrenbergii, L. lemniscatus) or could
change from pentagonal to elliptical (L. erythropterus,
L. johnii, L. malabaricus and L. rivulatus) or vice
versa (L. lutjanus).
In all species, the sulcus acusticus is heterosulcoid,
ostial with median position. The ostium is typically
funnel-like, except for L. bengalensis (rectangular),
L. johnii and L. malabaricus (funnel-like to rectangular).
It can be shorter than the cauda (L. argentimaculatus,
L. bengalensis, L. ehrenbergii, L. erythropterus,
L. fulviflamma and L. lutjanus), or approximately as
long as the cauda (L. johnii, L. lemniscatus,
L. malabaricus, L. rivulatus and L. russellii). The cauda
is tubular ending close to the posterior ventral margin.
The species L. malabaricus, L. johnii and
L. lutjanus show a cauda that is slightly curved, while
in the remaining species it is strongly or markedly
flexed.
The anterior margin noticeably varies and can
change with ageing: round in L. rivulatus, roundangled in L. malabaricus, blunt to peaked in L.
ehrenbergii, peaked in L. lutjanus and L. lemniscatus,
peaked to angled in L. argentimaculatus, peaked to
round in L. erythropterus, L. russellii and angled in L.
bengalensis, L. fulviflamma and L. johnii.
The rostrum is short and broad in all species,
although the tip differs between and within species:
blunt (L. bengalensis, L. ehrenbergii, L. malabaricus and
L. russellii), pointed (L. ehrenbergii, L. erythropterus,
L. johnii, L. lemniscatus and L. lutjanus) round
(L. argentimaculatus, L. erythropterus, L. rivulatus and
L. russellii), oblique (L. fulviflamma) or angled
(L. johnii); with an antirostrum poorly defined or small,
broad and pointed; while the excisura can be wide
without or with a deep and acute notch. Finally, the
posterior region is oblique in most of species but
angled in L. johnii, L. lutjanus and L. malabaricus.
Shape indices
The three shape indices (aspect ratio, compactness and
rectangularity) showed significant differences among
groups of species. The aspect ratio was the index most
variable showing higher mean values in Lutjanus
argentimaculatus, L. erythropterus, L. malabaricus and
L. rivulatus and lowest in L. russellii. By contrast, the
compactness and rectangularity was less apparent for
the groups (Table I, Figure 6).

807

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Identification of snappers using otoliths

Figure 3. Sagittal otoliths of Lutjanus spp. from the Persian Gulf. A, L. argentimaculatus (42.3
63.2
80.2 cm TL); B, L. bengalensis
(21 cm TL); C, L. ehrenbergii (14.6
20.3
26 cm TL); D, L. erythropterus (31.6
38.7
52.3 cm TL). Scale bars  1 mm.

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808 Z. Sagighzadeh et al.

Figure 4. Sagittal otoliths of Lutjanus spp. from the Persian Gulf. A, L. fulviflamma (17.6
21.4
26 cm TL); B, L. johnii (16.7
46
75.4 cm
TL); C, L. lemniscatus (29.8
40.5
51.4 cm TL); D, L. lutjanus (15.3
19.1
23.2 cm TL). Scale bars  1 mm.

809

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Identification of snappers using otoliths

Figure 5. Sagittal otoliths of Lutjanus spp. from the Persian Gulf. A, L. malabaricus (23.5
45.9
73.2 cm TL); B, L. rivulatus (40.5
53.7

66.7 cm TL); C, L. russellii (15
25.9
37.2 cm TL). Scale bars  1 mm.

810 Z. Sagighzadeh et al.

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The FFT was the weakest method for species

discrimination, with a classification success of 47.8%
of cases (Table II). Only the otoliths of Lutjanus
argentimaculatus and L. lemniscatus were identified
above 70%, while L. fluviflamma, L. johnii, L. lutjanus,
L. malabaricus and L. rivulatus were assigned below
40%. The EFD and WT noticeably improved the
identification of the most species increasing the total
percentage to 57.5% and 65.5%, respectively
(Table II); however, it was not the same in L.
lemniscatus, L. erythropterus and L. ehrenbergii. Finally,
the joint application of the three methods allowed a
classification success of 77.6% to be reached. In this
case, the percentage of classification was above 70%,
except for L. fulviflamma where it was 36.4% (Table II).

Discussion

Figure 6. Box plots for the shape indices (aspect ratio, compactness and rectangularity) by Lutjanus spp. from the Persian Gulf.
(see Table 1 for abbreviation explanations).

Otolith weight and contour

For the three methods applied (EFD, FFT and WT)
to characterize the otolith contour, the number of
coefficients implied in the mathematical analyses were
very low. In the case of EFD, the shape variability was
fulfilled only with the first 9 coefficients, while a total
of 53 of 1024 FFT harmonics meet the criteria
adequately. Finally, only 8 of 1536 wavelets were
used, of which most corresponded to the wavelet
number four because wavelets five and six were
characterized by a high variability among species
and fish length. However, the otolith weight was the
most important factor for species discrimination
(ANOVA F 113.21, P B0.001).

Several different methods were tested in the statistical DFA cross-validation procedures to determine
otolith shape identification of species. The results of
the morphological study indicate that otoliths of
snappers from the Persian Gulf are very similar in
morphological terms (shape, type of sulcus acusticus, ostium and cauda), but do show differences in
the anterior and posterior rims of the otolith. Up to
now, only Rivaton & Bourret (1999) had provided
images of otoliths of some species of snappers
(Lutjanus argentimaculatus, L. fulviflamma, L. lutjanus
and L. russellii) from the Indian Ocean. Finally,
although the analysis of the otolith outline revealed a
high variability in the most species, only the combination of all methods provided strong results. The
ability to distinguish between otoliths of these
species is an important finding since species identification has proven difficult when these species cooccur in the stomach contents of predators.
The advantage of this multiscale analysis (WT) is
that it enables us to identify single morphological
points (landmarks) located on the x-axis along the
contour where the rostrum is the origin of the
contour (Parisi-Baradad et al. 2005; Piera et al.
2005; Lombarte et al. 2006). The differences
observed in the ventral margin between rostrum
and post-rostrum and did not depend on the fish
length. On the contrary, EFD and FFT only give a
global approximation of outline variability (ReigBolan os et al. 2010). However, EFD does not
require equal intervals along the outline and therefore can accommodate significantly more complex
shapes than polar Fourier functions, which describe
and characterize outlines better (e.g. Kuhl &
Giardina 1982; Lestrel 1997; Stransky & MacLellan
2005; Tracey et al. 2006; Stransky et al. 2008).
Notwithstanding, Marti-Puig et al. (2010) asserted
that although EFD can represent any contour when

Identification of snappers using otoliths

811

Table II. Results of cross-validation using leave-one-out method for classifying Lutjanus spp. from the Persian Gulf. EFD, elliptical Fourier
descriptors; FFT, fast Fourier transform; WT, wavelets.
Predicted group membership (percentage)
Actual group

Downloaded by [Centre Mediterrani D'investig] at 00:04 21 August 2012

EFD  otolith weight

L. argentimaculatus (Larg)
L. ehrenbergii (Lehr)
L. erythropterus (Lery)
L. fulviflamma (Lful)
L. johnii (Ljoh)
L. lemniscatus (Llem)
L. lutjanus (Llut)
L. malabaricus (Lmal)
L. rivulatus (Lriv)
L. russellii (Lrus)
FFT otolith weight
L. argentimaculatus (Larg)
L. ehrenbergii (Lehr)
L. erythropterus (Lery)
L. fulviflamma (Lful)
L. johnii (Ljoh)
L. lemniscatus (Llem)
L. lutjanus (Llut)
L. malabaricus (Lmal)
L. rivulatus (Lriv)
L. russellii (Lrus)
WT otolith weight
L. argentimaculatus (Larg)
L. ehrenbergii (Lehr)
L. erythropterus (Lery)
L. fulviflamma (Lful)
L. johnii (Ljoh)
L. lemniscatus (Llem)
L. lutjanus (Llut)
L. malabaricus (Lmal)
L. rivulatus (Lriv)
L. russellii (Lrus)
EFD  FFT  WT otolith weight
L. argentimaculatus (Larg)
L. ehrenbergii (Lehr)
L. erythropterus (Lery)
L. fulviflamma (Lful)
L. johnii (Ljoh)
L. lemniscatus (Llem)
L. lutjanus (Llut)
L. malabaricus (Lmal)
L. rivulatus (Lriv)
L. russellii (Lrus)

Larg

Lehr

Lery

Lful

Ljoh

55.7

6.6
44.4
18.2
9.8

24.6
22.2
45.5

1.6
22.2

Llem

Llut

Lmal

Lriv

3.3

1.6
11.1

6.6

Lrus

7.7

92.3

8.7
8.5
8.3

4.3
21.7
2.1
16.7
3.2

56.5
4.3
25.5

3.3
66.7
9.1
5.4

56.5
6.4
8.3

7.7
1.6

16.4
36.4
6.5

37.0
73.9

4.3

26.1
4.3
8.3

27.3
2.2
4.3
6.4
66.7

17.4
27.7
16.7

3.2

2.2
26.1
4.3
74.2

8.2

1.6

4.9
11.1

3.3

9.1
8.7
4.3
30.4
2.1

9.1
28.3

12.0

1.1
17.4

34.0
41.7

23.4
25.0

2.1
8.3
58.1

3.3
11.1

3.3
11.1

1.6

1.1
17.4

11.1

4.3
26.1
2.1

19.6
4.3
13.0
46.8

22.6

92.3
60.7
11.1
36.4
1.1

9.1
10.9

55.4

38.7

100.0
47.5
18.2
2.2
8.7
13.0

3.2

3.3
55.6
9.1
4.3
4.3
4.3
6.4

26.2
11.1
45.5

3.2

3.2

4.3

8.2
11.1
81.5
4.3
4.3
10.6
8.3

3.3

3.3

9.1

18.2
2.2

5.4

3.3

47.8
10.6
8.3

13.0
59.6
16.7

13.0
8.5
66.7

65.2

19.4

4.3
71.0

92.3

7.7
75.4

11.5

1.6

9.8

77.8
36.4
2.2
13.0

11.1
36.4
1.1

81.5
4.3

18.2
2.2
78.3
6.4

12.8

8.3

a large number of coefficients are used, but in

practice the application of the number of descriptors
is limited to a certain low value (e.g. Parisi-Bardad
et al. 2005; Tracey et al. 2006; Burke et al. 2008;
present study) representing an approximation to the
original silhouette to a lesser degree than other
analyses. This becomes obvious in the presence of
a close similarity between species. Our results
support the hypothesis that Fourier transcription
methods are more efficient when discriminating
between groups or stocks of fish, but are not when

3.2

10.9

2.2
13.0

82.6

4.3
4.3

1.6
11.1
9.1

12.9

70.2
8.3

4.3
2.1
83.3

4.3
83.9

the aim is to identify and classify closer species.

However, the statistical results of the study showed
that combinations of these methods could be very
useful to discriminate species.
Otolith shape is affected by ecological, ontogenetic,
environmental and genetic factors (Morales-Nin
1987; Castonguay et al. 1991; Gago 1993; Friedland
& Reddin 1994; Lombarte et al. 2003; Cadrin &
Friedland 2005). In this framework, juvenile and
subadult fishes of many snappers of the Persian Gulf
and the Oman Sea exhibit a high grade of adaptability

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812 Z. Sagighzadeh et al.

to different environments such as mangroves, seagrass beds, freshwater streams or brackish water.
They migrate to rocky environments or coral reefs
when they are adults and then take up permanent
residence there (Druzhinin 1970; Cervigo n 1993;
Cocheret et al. 2003). These species differ in behaviour (nocturnal or diurnal), coloration patterns on
body, habitat and sociality (solitary or groups)
influencing their behavior and intra- and interspecific communications (Lewis & Pring 1986;
Sainsbury 1987; Williams & Russ 1992; Sommer
et al. 1996; Kailola et al. 1993; Lieske & Myers 1994;
Kuiter & Tonozuka 2001). In fact, the anatomical
affinity
narrower otoliths and deeper notch of
excisura ostti between rostrum and antirostrum of
the otolith of Lutjanus ehrenbergii, L. fulviflamma and
L. lutjanus
is reasonable taking into account that
these species inhabit coastal environments and have a
similar coloration and body shape, which may be
interpreted as a common phylogenetic origin where
L. fulviflamma might be the ancestral species.
Our study is the first investigation that compares
actual data with related mathematical protocols by
several methods and provides results of a high degree
of reliability. At this point, we would like to comment
on the phenomenon of multicollinearity among
variables. Elimination of these variables did not
significantly increase the percentage of classification.
It is true that the number of wavelets used in the
discriminant analysis was low, but that was due to the
phenomenon of multicollinearity among the closer
wavelets. Moreover, our study demonstrates that the
variance homogeneity among species is smaller in
wavelets with a low frequency, which show more
details of the otolith profile, which we consider a
relevant result for future studies using wavelets.
Finally, several studies have shown that otolith
weight is a more powerful discriminator and has less
variation than Fourier descriptors (Forsberg & Neal
1993; Tuset et al. 2006). Our results support these
findings and reinforce the fact that future research
should be focused on developing three-dimensional
techniques to help researchers in the proper identification of species.

Acknowledgements
This work has been supported by the MICIN
CTM2010-19701 (AFORO3D project). The anonymous reviewers provided many helpful comments
on both the structure and analysis of an earlier
version of the report. Their comments resulted in a
much improved manuscript and their assistance is
appreciated. Dr Victor Tuset is worker of CSIC
within the modality JAE-Postdoc of Programme

Junta para la Ampliacio n de Estudios co-funded by

the European Social Foundation.
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