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Comparison of Different Otolith Shape Descriptors and Morphometrics00
Comparison of Different Otolith Shape Descriptors and Morphometrics00
ORIGINAL ARTICLE
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.
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 24308N and
49628E. 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
*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
803
Figure 1. Map of the Persian Gulf (NE Indian Ocean) showing the area from where Lutjanus spp. were collected.
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)
minmax
423802
146260
316523
176260
167754
298514
153232
235732
405667
150372
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 67.83174.66
18.1540.12 61.57113.06
22.1136.70
30.27313.09
44.4992.36
19.537.35
39.39359.45
98.77175.65
15.5458.73
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
7.4012.53
3.795.68
7.6710.02
4.275.51
5.1015.46
5.878.73
3.935.54
5.9417.31
9.3412.51
3.466.99
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 12.7420.49
6.5410.42
11.6816.32
7.279.82
8.3628.83
10.2415.55
6.839.82
9.2330.16
14.5519.48
6.5012.50
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 37.5061.62
18.8929.24
34.1046.53
20.5226.52
24.1092.74
29.1546.67
19.3728.45
25.9586.19
42.8658.28
18.9333.60
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
0.160.83
0.030.11
0.150.38
0.040.11
0.052.20
0.100.40
0.04 0.11
0.102.45
0.330.86
0.020.16
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
0.550.69
0.510.61
0.610.68
0.530.64
0.530.66
0.510.62
0.540.60
0.570.72
0.630.68
0.470.58
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 20.1025.10
18.6625.74
17.9619.17
18.6821.70
17.1027.47
18.4224.7
18.0122.89
16.6726.87
18.1723.23
19.2224.42
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
0.640.72
0.680.74
0.660.70
0.680.73
0.680.73
0.670.73
0.690.73
0.670.73
0.680.73
0.660.71
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
p2
2
an0 ib0 n
805
Hormonic
Statistical analyses
Fish length
Oij; adj Oij b TLij MTLj ;
Results
Anatomical description
Morphologically, the sagittal otoliths of Lutjanus
spp. are similar with small specific variations
(Figures 35). 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
Figure 3. Sagittal otoliths of Lutjanus spp. from the Persian Gulf. A, L. argentimaculatus (42.363.280.2 cm TL); B, L. bengalensis
(21 cm TL); C, L. ehrenbergii (14.620.326 cm TL); D, L. erythropterus (31.638.752.3 cm TL). Scale bars 1 mm.
Figure 4. Sagittal otoliths of Lutjanus spp. from the Persian Gulf. A, L. fulviflamma (17.621.426 cm TL); B, L. johnii (16.74675.4 cm
TL); C, L. lemniscatus (29.840.551.4 cm TL); D, L. lutjanus (15.319.123.2 cm TL). Scale bars 1 mm.
809
Figure 5. Sagittal otoliths of Lutjanus spp. from the Persian Gulf. A, L. malabaricus (23.545.973.2 cm TL); B, L. rivulatus (40.553.7
66.7 cm TL); C, L. russellii (1525.937.2 cm TL). Scale bars 1 mm.
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).
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
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
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
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
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
813