Corresponding author: Kōji Yokogawa (
Academic editor: Nina Bogutskaya
Morphological differences, including growth-related changes, were examined in three morphologically similar East Asian sea bass species,
Yokogawa K (2019) Morphological differences between species of the sea bass genus
The sea basses of the genus
General aspects of small (fingerling) and large (adult) individuals of
Dots / spots on lateral body regions of
Thus, morphological identifications of the three
Measurements were based on the following
Methods of measurements and counts followed
Characters considered for the analysis.
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Post-orbital preopercular width | POPW | |
Standard length | SL | Upper jaw length | UJL |
Pre-anus length |
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Lower jaw length | LJL |
Body depth | BD |
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Body width | BWT | Dorsal fin spine | DFS |
Caudal peduncle depth | CPD | Dorsal fin soft ray | DFR |
Caudal peduncle anterior depth | CPAD | Anal fin spine | AFS |
Caudal peduncle length | CPL | Anal fin ray | AFR |
Pre-dorsal length |
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Pectoral fin ray | P1FR |
First dorsal fin (longest spine) length | FDFL | Pelvic fin spine | P2FS |
Second dorsal fin (longest ray) length |
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Pelvic fin ray | P2FR |
Caudal fin length | CFL | Pored scale on lateral line | LLS |
Caudal fin notch depth |
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Scale above lateral line | SAL |
Anal fin (longest ray) length |
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Scale below lateral line | SBL |
Pectoral fin length | P1FL | Upper-limb gill raker | UGR |
Pelvic fin length | P2FL | Lower-limb gill raker | LGR |
Pectoral scaly area length | PSAL | Total gill raker | TGR |
Head length |
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Abdominal vertebra | AV |
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Caudal vertebra | CV | |
Snout length | SNL | Total vertebra | TV |
Orbital diameter | OD |
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Inter-orbital width | IOW | Dorsocephalic scale row | DSR |
Sub-orbital width | SOW | Ventromandibular scale row | VSR |
Post-orbital length | POL | First anal pterygiophore | FAP |
Illustrations of
Scale row and paired fin ray counts were made on the left side of the body, whereas gill rakers were counted on the first gill arch on the right side by separating the upper and lower limbs of the gill arch. Because counts of pelvic fin-spine (P2FS) and soft rays (P2FRs) showed no variation (P2FS: 1, P2FRs: 5 in all specimens), these counts were omitted from the statistical analyses. Abdominal and caudal vertebrae were counted, and first anal fin pterygiophore morphology observed from radiographs.
Total numbers of recognizable black or faint spots / dots on the left side of the body and mid-dorsal aspect of the caudal peduncle (Fig.
For a length-measured dimension (LD), a growth-related proportional change pattern is given by the relationship between base dimension [e.g., standard length (SL) or head length (
Regression parameters and correlation between standard length (SL) or head length (
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SL– |
64.42 | 0.004 | 0.092 | 74.89 | -0.026 | -0.524 | 63.90 | 0.008 | 0.270 |
SL–BD/SL | 44.23 | -0.108 | -0.735 | 29.94 | -0.029 | -0.379 | 33.03 | -0.021 | -0.240 |
SL–BWT/SL | 8.78 | 0.075 | 0.471 | 10.71 | 0.048 | 0.455 | 8.43 | 0.079 | 0.466 |
SL–CPD/SL | 16.55 | -0.100 | -0.749 | 11.48 | -0.025 | -0.353 | 11.32 | 0.002 | 0.034 |
SL–CPL/SL | 22.33 | -0.007 | -0.069 | 19.83 | 0.019 | 0.216 | 21.55 | -0.010 | -0.115 |
SL–CPAD/SL | 21.12 | -0.091 | -0.686 | 14.36 | -0.014 | -0.220 | 15.05 | 0.009 | 0.140 |
SL– |
44.01 | -0.041 | -0.574 | 39.76 | -0.029 | -0.513 | 45.07 | -0.039 | -0.711 |
SL–FDFL/SL | 22.72 | -0.081 | -0.407 | 12.40 | 0.008 | 0.065 | 22.22 | -0.086 | -0.541 |
SL– |
36.65 | -0.201 | -0.762 | 17.05 | -0.068 | -0.443 | 23.31 | -0.091 | -0.485 |
SL–CFL/SL | 32.62 | -0.085 | -0.472 | 17.40 | 0.008 | 0.056 | 28.45 | -0.055 | -0.445 |
SL– |
9.30 | -0.115 | -0.220 | 2.87 | 0.077 | 0.176 | 25.10 | -0.296 | -0.781 |
SL– |
28.14 | -0.142 | -0.713 | 18.56 | -0.061 | -0.474 | 24.60 | -0.096 | -0.553 |
SL–P1FL/SL | 25.19 | -0.070 | -0.581 | 16.98 | -0.010 | -0.109 | 19.79 | -0.024 | -0.270 |
SL–P2FL/SL | 31.24 | -0.101 | -0.701 | 25.47 | -0.073 | -0.682 | 23.84 | -0.040 | -0.357 |
SL– |
42.88 | -0.054 | -0.677 | 38.39 | -0.036 | -0.629 | 46.25 | -0.066 | -0.836 |
SL–SNL/SL | 8.23 | 0.002 | 0.047 | 11.42 | -0.087 | -0.664 | 10.91 | -0.027 | -0.456 |
SL–OD/SL | 65.54 | -0.431 | -0.958 | 42.67 | -0.364 | -0.945 | 55.60 | -0.368 | -0.963 |
SL–IOW/SL | 7.55 | -0.020 | -0.173 | 9.31 | -0.064 | -0.601 | 7.75 | -0.010 | -0.082 |
SL–SOW/SL | 2.26 | 0.067 | 0.232 | 1.80 | 0.135 | 0.513 | 2.04 | 0.070 | 0.246 |
SL–POPW/SL | 5.47 | 0.045 | 0.423 | 13.03 | -0.094 | -0.741 | 7.21 | -0.008 | 0.066 |
SL–POL/SL | 15.94 | 0.016 | 0.170 | 13.46 | 0.060 | 0.691 | 19.07 | -0.027 | -0.373 |
SL–UJL/SL | 19.09 | -0.061 | -0.706 | 20.81 | -0.083 | -0.778 | 22.01 | -0.071 | -0.778 |
SL–LJL/SL | 20.51 | -0.058 | -0.700 | 22.29 | -0.084 | -0.782 | 21.66 | -0.052 | -0.629 |
SL–PSAL/SL1 | 8.14 | -0.130 | -0.203 | ||||||
SL–POPW/SNL | 71.07 | 0.030 | 0.314 | 90.56 | 0.031 | 0.222 | 65.79 | 0.020 | 0.149 |
20.42 | 0.057 | 0.530 | 28.48 | -0.054 | -0.453 | 24.50 | 0.040 | 0.533 | |
109.60 | -0.400 | -0.946 | 79.68 | -0.338 | -0.945 | 93.74 | -0.323 | -0.950 | |
18.38 | 0.033 | 0.246 | 23.83 | -0.031 | -0.292 | 17.72 | 0.057 | 0.359 | |
5.90 | 0.127 | 0.397 | 5.55 | 0.178 | 0.625 | 4.98 | 0.143 | 0.432 | |
14.81 | 0.090 | 0.690 | 26.66 | -0.022 | -0.240 | 16.41 | 0.061 | 0.418 | |
39.87 | 0.073 | 0.729 | 38.83 | 0.099 | 0.873 | 42.78 | 0.041 | 0.498 | |
44.57 | -0.009 | -0.139 | 51.77 | -0.049 | -0.667 | 47.53 | -0.006 | -0.109 | |
48.01 | -0.005 | -0.092 | 55.24 | -0.049 | -0.713 | 47.36 | 0.014 | 0.237 |
1 Simple patterned regressions could not be applied for complicated fluctuations in
Characteristics that changed with growth were evaluated so as to determine if the changes were isometric or allometric, i.e., regressions between SL (or
To examine inter-specific differences in length-measured characters, regressions between SL (or
Regression parameters (slope and intercept) and correlation between logarithm-transformed length-measured characters, together with results of
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ln SL–ln |
1.004 | -0.44 | 1.39 | 0.974 | -0.29 | -7.97*** | 1.008 | -0.45 | 3.25* |
ln SL–ln BD | 0.892 | -0.82 | -16.35*** | 0.971 | -1.21 | -5.31*** | 0.979 | -1.11 | -2.87* |
ln SL–ln BWT | 1.075 | -2.43 | 8.05*** | 1.048 | -2.23 | 6.62*** | 1.079 | -2.47 | 6.10*** |
ln SL–ln CPD | 0.900 | -1.80 | -17.04*** | 0.975 | -2.16 | -4.89*** | 1.002 | -2.18 | 0.40 |
ln SL–ln CPL | 0.993 | -1.50 | -1.05 | 1.019 | -1.62 | 2.86* | 0.990 | -1.53 | -1.33 |
ln SL–ln CPAD | 0.909 | -1.55 | -14.28*** | 0.986 | -1.94 | -2.92* | 1.009 | -1.89 | 1.63 |
ln SL–ln |
0.959 | -0.82 | -10.56*** | 0.971 | -0.92 | -7.71*** | 0.961 | -0.80 | -11.72*** |
ln SL–ln FDFL | 0.919 | -1.48 | -6.72*** | 1.008 | -2.09 | 0.85 | 0.914 | -1.50 | -7.45*** |
ln SL–ln |
0.794 | -0.97 | -17.15*** | 0.932 | -1.77 | -6.31*** | 0.909 | -1.46 | -6.42*** |
ln SL–ln CFL | 0.914 | -1.11 | -7.84*** | 1.008 | -1.75 | 0.70 | 0.974 | -1.35 | -2.55 |
ln SL–ln |
0.880 | -2.35 | -3.41** | 1.077 | -3.55 | 2.22 | 0.704 | -1.38 | -13.88*** |
ln SL–ln |
0.858 | -1.27 | -15.17*** | 0.939 | -1.68 | -6.97*** | 0.904 | -1.40 | -7.67*** |
ln SL–ln P1FL | 0.930 | -1.38 | -10.73*** | 0.990 | -1.77 | -1.41 | 0.976 | -1.62 | -3.25* |
ln SL–ln P2FL | 0.899 | -1.16 | -14.81*** | 0.927 | -1.37 | -12.06*** | 0.960 | -1.43 | -4.42*** |
ln SL–ln |
0.946 | -0.85 | -13.87*** | 0.964 | -0.96 | -10.46*** | 0.934 | -0.77 | -17.67*** |
ln SL–ln SNL | 1.002 | -2.50 | 0.67 | 0.913 | -2.17 | -11.57*** | 0.973 | -2.22 | -5.94*** |
ln SL–ln OD | 0.569 | -0.42 | -50.25*** | 0.636 | -0.85 | -37.39*** | 0.632 | -0.59 | -41.41*** |
ln SL–ln IOW | 0.980 | -2.58 | -2.64 | 0.936 | -2.37 | -9.71*** | 0.990 | -2.56 | -0.95 |
ln SL–ln SOW | 1.067 | -3.79 | 3.60** | 1.135 | -4.02 | 7.73*** | 1.070 | -3.89 | 2.94* |
ln SL–ln POPW | 1.033 | -2.84 | 5.68*** | 0.943 | -2.26 | -7.72*** | 0.993 | -2.63 | -0.69 |
ln SL–ln POL | 1.014 | -1.82 | 2.10 | 1.060 | -2.00 | 12.25*** | 0.974 | -1.66 | -4.56*** |
ln SL–ln UJL | 0.939 | -1.66 | -15.04*** | 0.917 | -1.57 | -16.02*** | 0.929 | -1.51 | -14.34*** |
ln SL–ln LJL | 0.942 | -1.58 | -14.74*** | 0.916 | -1.50 | -16.11*** | 0.948 | -1.53 | -9.34*** |
ln SNL–ln POPW | 1.026 | -0.26 | 4.37*** | 1.020 | 0.01 | 1.71 | 1.017 | -0.36 | 4.19*** |
ln |
1.057 | -1.59 | 9.41*** | 0.946 | -1.26 | -6.65*** | 1.040 | -1.41 | 7.28*** |
ln |
0.600 | 0.09 | -44.06*** | 0.662 | -0.23 | -37.38*** | 0.677 | -0.06 | -35.28*** |
ln |
1.033 | -1.69 | 3.82** | 0.969 | -1.43 | -3.94** | 1.057 | -1.73 | 4.45*** |
ln |
1.127 | -2.83 | 6.52*** | 1.178 | -2.89 | 10.36*** | 1.143 | -3.00 | 5.55*** |
ln |
1.090 | -1.91 | 14.36*** | 0.978 | -1.32 | -3.19* | 1.061 | -1.81 | 5.32*** |
ln |
1.073 | -0.92 | 15.93*** | 1.099 | -0.95 | 23.15*** | 1.041 | -0.85 | 6.62*** |
ln |
0.991 | -0.81 | -2.11 | 0.951 | -0.66 | -11.57*** | 0.994 | -0.74 | -1.27 |
ln |
0.995 | -0.73 | -0.19 | 0.952 | -0.59 | -13.19*** | 1.014 | -0.75 | 2.81* |
Asterisks indicate significance of t values; single, double and triple asterisks indicate 5%, 1% and 0.1% levels, respectively, after Holm-Bonferroni correction by species.
Because some meristic counts tended to increase significantly with growth (Table
Regression parameters (slope and intercept) and correlation between standard length (SL) and meristic counts of three
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SL–DFS counts | -0.00008 | 12.87 | -0.019 | -0.28 |
SL–DFR counts | -0.00081 | 13.13 | -0.130 | -2.05 |
SL–AFR counts | 0.00048 | 7.56 | 0.089 | 1.34 |
SL–P1FR counts | -0.00047 | 16.96 | -0.086 | -1.30 |
SL–LLS counts | 0.01207 | 77.01 | 0.343 | 5.50*** |
SL–SAL counts | -0.00258 | 15.84 | -0.258 | -3.98** |
SL–SBL counts | 0.00057 | 18.57 | 0.046 | 0.68 |
SL–UGR counts | 0.00111 | 8.63 | 0.126 | 1.90 |
SL–LGR counts | -0.00025 | 17.93 | -0.027 | -0.41 |
SL–TGR counts | 0.00086 | 26.56 | 0.073 | 1.10 |
SL–AV counts | 0.00017 | 16.00 | 0.073 | 0.93 |
SL–CV counts | -0.00068 | 20.02 | -0.108 | -1.38 |
SL–TV counts | -0.00051 | 36.02 | -0.083 | -1.80 |
SL–Dot counts | -0.02297 | 12.69 | -0.198 | -2.90* |
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SL–DFS counts | -0.00046 | 12.95 | -0.153 | -2.00 |
SL–DFR counts | -0.00028 | 13.03 | -0.066 | -0.86 |
SL–AFS counts | 0.00008 | 2.98 | 0.104 | 1.36 |
SL–AFR counts | 0.00097 | 7.34 | 0.217 | 2.88 |
SL–P1FR counts | 0.00079 | 16.33 | 0.190 | 2.50 |
SL–LLS counts | 0.00261 | 73.45 | 0.099 | 1.30 |
SL–SAL counts | 0.00008 | 15.52 | 0.009 | 0.24 |
SL–SBL counts | 0.00477 | 18.17 | 0.409 | 5.72*** |
SL–UGR counts | 0.00139 | 6.40 | 0.173 | 2.24 |
SL–LGR counts | 0.00330 | 14.70 | 0.507 | 7.49*** |
SL–TGR counts | 0.00469 | 21.11 | 0.408 | 5.68*** |
SL–AV counts | -0.00026 | 15.97 | -0.135 | -1.67 |
SL–CV counts | 0.00022 | 19.00 | 0.089 | 1.09 |
SL–TV counts | 0.00003 | 34.97 | -0.012 | -0.02 |
SL–Spot counts | 0.02333 | 33.89 | 0.126 | 1.62 |
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SL–DFS counts | -0.00026 | 13.05 | -0.092 | -1.08 |
SL–DFR counts | -0.00041 | 15.11 | 0.011 | -1.20 |
SL–AFS counts | -0.00002 | 3.00 | 0.001 | -0.34 |
SL–AFR counts | 0.00026 | 9.06 | 0.002 | 0.55 |
SL–P1FR counts | -0.00026 | 16.20 | 0.004 | -0.73 |
SL–LLS counts | 0.00264 | 72.91 | 0.169 | 1.99 |
SL–SAL counts | -0.00063 | 13.86 | -0.079 | -0.92 |
SL–SBL counts | -0.00013 | 15.79 | -0.014 | -0.16 |
SL–UGR counts | -0.00045 | 6.83 | -0.072 | -0.83 |
SL–LGR counts | -0.00109 | 17.11 | -0.176 | -2.07 |
SL–TGR counts | -0.00154 | 23.94 | -0.166 | -1.95 |
SL–AV counts | 0.00004 | 16.03 | 0.018 | 0.22 |
SL–CV counts | -0.00005 | 19.92 | -0.014 | -0.17 |
SL–TV counts | 0.00001 | 35.95 | -0.004 | -0.05 |
SL–Dot counts | -0.06278 | 24.74 | -0.365 | -4.53*** |
Asterisks indicate significance of t values; single, double and triple asterisks indicate 5%, 1% and 0.1% levels, respectively, after Holm-Bonferroni correction by species.
In the above statistical inferences, due to multiple tests being applied simultaneously in each case, multiple comparisons were introduced for the
In the three
Relationships between standard length and proportions of some length-measured body characters of three
Relationships between standard length and proportions of some length-measured body characters which exhibited prominent plot separation among three
Similar patterns of growth-related proportional changes common to the three species were observed for some characters, viz., significant positive allometric growth (proportions increased with growth) in body width and significant negative allometric growth (proportions decreased with growth) in head (
For length-measured dimensions (LD) of cephalic characters, SL-based (SL–LD / SL) and
Relationships between standard length or head length and proportions of length-measured cephalic characters of three
Growth-related proportional change patterns based on SL and
As well as some body characters, specific proportional change patterns were recognized for some characters, e.g., SL-based relationships of POPW, exhibiting isometric growth in
The relationship between SL and pectoral scaly area length (PSAL) in
Relationships between standard length (SL) and pectoral scaly area length (PSAL) proportions for
Plot separation of
Plot separation of first and second dorsal (FDFL and
Upward plot separation of
On the other hand, plot separation between
POPW proportional to SNL is shown graphically in Figure
Relationships between standard length and (post-orbital preopercular width) / (snout length) proportions of three
The
Relationships between standard length and some meristic characters which exhibited growth-related changes in some
Figure
Histograms of meristic characters of three
Some examples of
Relationships between standard length and dot / spot counts on lateral body regions of
Post-juvenile specimens (> ca. 70 mm SL) of the three
Squamation on dorsal head regions of
Some individuals of the three
Frequencies (%) of ventromandibular scale row status in three
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≤100 | 0.0 | 0.0 | 100.0 | 0.0 | 0.0 | 100.0 |
100–200 | 0.0 | 14.3 | 85.7 | 10.7 | 21.4 | 67.9 |
200–300 | 5.0 | 25.0 | 70.0 | 35.0 | 30.0 | 35.0 |
300–400 | 5.3 | 26.3 | 68.4 | 31.6 | 57.9 | 10.5 |
>400 | 25.0 | 50.0 | 25.0 | 37.5 | 62.5 | 0.0 |
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≤100 | 0.0 | 0.0 | 100.0 | 0.0 | 0.0 | 100.0 |
100–200 | 0.0 | 0.0 | 100.0 | 0.0 | 0.0 | 100.0 |
200–300 | 0.0 | 18.2 | 81.8 | 22.7 | 54.5 | 22.7 |
300–400 | 5.6 | 55.6 | 38.9 | 55.6 | 27.8 | 16.7 |
>400 | 12.1 | 51.5 | 36.4 | 84.8 | 15.2 | 0.0 |
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≤100 | 0.0 | 13.3 | 86.7 | 6.7 | 13.3 | 80.0 |
100–200 | 70.5 | 18.0 | 11.5 | 49.2 | 19.7 | 31.1 |
200–300 | 95.1 | 4.9 | 0.0 | 97.6 | 2.4 | 0.0 |
300–400 | 100.0 | 0.0 | 0.0 | 100.0 | 0.0 | 0.0 |
>400 | 100.0 | 0.0 | 0.0 | 100.0 | 0.0 | 0.0 |
All three
Radiographs of first anal pterygiophores in
Analyses of covariance (ANCOVA) for regressions of logarithm-transformed length-measured characters by pairwise comparisons for the three
Results of analysis of covariance (ANCOVA) (
Regression | ||||||
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Slope | Intercept | Slope | Intercept | Slope | Intercept | |
ln SL–ln |
7.00*** | – | 1.03 | 6.61*** | 7.08*** | – |
ln SL–ln BD | 9.03*** | – | 8.16*** | – | 0.91 | 26.57*** |
ln SL–ln BWT | 2.34 | 6.92*** | 0.22 | 2.58* | 2.22 | 9.23*** |
ln SL–ln CPD | 9.51*** | – | 10.97*** | – | 3.26 | 26.59*** |
ln SL–ln CPL | 2.81 | 2.97* | 0.29 | 9.35*** | 2.69 | 11.00*** |
ln SL–ln CPAD | 9.41*** | – | 10.18*** | – | 2.91 | 36.84*** |
ln SL–ln |
2.22 | 11.60*** | 0.25 | 10.07*** | 1.83 | 21.83*** |
ln SL–ln FDFL | 5.75*** | – | 0.30 | 5.13*** | 5.99*** | – |
ln SL–ln |
8.52*** | – | 6.02*** | – | 1.23 | 18.51*** |
ln SL–ln CFL | 6.05*** | – | 3.45* | – | 1.86 | 16.84*** |
ln SL–ln |
3.99** | – | 3.49* | – | 7.37*** | – |
ln SL–ln |
6.28*** | – | 2.88 | 12.25*** | 2.25 | 11.23*** |
ln SL–ln P1FL | 6.17*** | – | 4.24** | – | 1.26 | 12.21*** |
ln SL–ln P2FL | 3.07 | 9.89*** | 5.18*** | – | 2.96 | 16.28*** |
ln SL–ln |
3.45* | – | 1.82 | 5.42*** | 5.30*** | – |
ln SL–ln SNL | 9.97*** | – | 3.68* | – | 5.53*** | – |
ln SL–ln OD | 5.26*** | – | 4.66*** | – | 0.26 | 28.99*** |
ln SL–ln IOW | 4.29** | 0.73 | 10.95*** | 4.27** | – | |
ln SL–ln SOW | 2.64 | 7.96*** | 0.08 | 5.20*** | 2.15 | 12.35*** |
ln SL–ln POPW | 10.37*** | – | 3.61* | – | 4.15** | – |
ln SL–ln POL | 5.43*** | – | 3.90** | – | 10.54*** | – |
ln SL–ln UJL | 3.42* | – | 1.44 | 25.97*** | 1.55 | 26.46*** |
ln SL–ln LJL | 4.05** | – | 0.79 | 22.93*** | 3.76* | – |
ln SNL–ln POPW | 0.48 | 33.61*** | 0.76 | 27.56*** | 0.18 | 44.42*** |
ln |
11.07*** | – | 1.82 | 23.86*** | 7.76*** | – |
ln |
4.84*** | – | 5.29*** | – | 1.02 | 28.82*** |
ln |
5.47*** | – | 1.52 | 7.78*** | 5.92*** | – |
ln |
1.95 | 9.36*** | 0.46 | 6.42*** | 1.14 | 15.08*** |
ln |
12.17*** | – | 2.40 | 2.74* | 6.34*** | – |
ln |
4.04** | – | 4.15** | – | 7.64*** | – |
ln |
6.89*** | – | 0.38 | 22.63*** | 6.19*** | – |
ln |
7.92*** | – | 2.84 | 3.37** | 9.71*** | – |
Numbers indicate t values given by ANCOVA. Asterisks indicate significance of t vales; single, double and triple asterisks indicate 5%, 1% and 0.1% levels, respectively, after Holm-Bonferroni correction. Bars indicate that calculation was not demonstrated because significance was recognized for the slope and ANCOVA was therein terminated.
Although the Mann-Whitney
Results of the Mann-Whitney
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DFS counts | 0.37 | 3.00* | 3.64** |
DFR counts | 0.12 | 16.22*** | 15.60*** |
AFS counts | 0.00 | 1.29 | 0.64 |
AFR counts | 1.39 | 14.64*** | 14.11*** |
P1FR counts | 5.69*** | 10.62*** | 5.77*** |
LLS counts | 11.53*** | 13.74*** | 0.89 |
SAL counts | 2.04 | 11.50*** | 11.47*** |
SBL counts | 3.57** | 14.43*** | 13.88*** |
UGR counts | 14.31*** | 14.58*** | 0.65 |
LGR counts | 15.45*** | 8.83*** | 11.76*** |
TGR counts | 16.54*** | 15.13*** | 7.81*** |
AV counts | 4.23*** | 0.64 | 4.15*** |
CV counts | 13.58*** | 0.01 | 13.45*** |
TV counts | 14.82*** | 0.73 | 14.09*** |
Asterisks indicate significance of z vales; single, double and triple asterisks indicate 5%, 1% and 0.1% levels, respectively, after Holm-Bonferroni correction.
Standard errors (SEs) for regression lines between logarithm-transformed SL and length-measured characters, and between SL and meristic characters are summarized in Table
Standard errors for morphological character regressions of three
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ln SL–ln |
0.024 | 0.029 | 0.015 |
ln SL–ln BD | 0.057 | 0.050 | 0.043 |
ln SL–ln BWT | 0.080 | 0.064 | 0.077 |
ln SL–ln CPD | 0.051 | 0.046 | 0.036 |
ln SL–ln CPL | 0.055 | 0.060 | 0.044 |
ln SL–ln CPAD | 0.055 | 0.044 | 0.031 |
ln SL–ln |
0.033 | 0.033 | 0.020 |
ln SL–ln FDFL | 0.103 | 0.084 | 0.068 |
ln SL–ln |
0.094 | 0.096 | 0.084 |
ln SL–ln CFL | 0.085 | 0.095 | 0.068 |
ln SL–ln |
0.273 | 0.299 | 0.119 |
ln SL–ln |
0.079 | 0.079 | 0.074 |
ln SL–ln P1FL | 0.056 | 0.063 | 0.045 |
ln SL–ln P2FL | 0.058 | 0.053 | 0.054 |
ln SL–ln |
0.034 | 0.031 | 0.022 |
ln SL–ln SNL | 0.044 | 0.067 | 0.027 |
ln SL–ln OD | 0.074 | 0.087 | 0.053 |
ln SL–ln IOW | 0.066 | 0.059 | 0.065 |
ln SL–ln SOW | 0.160 | 0.155 | 0.140 |
ln SL–ln POPW | 0.050 | 0.058 | 0.060 |
ln SL–ln POL | 0.057 | 0.044 | 0.035 |
ln SL–ln UJL | 0.035 | 0.046 | 0.029 |
ln SL–ln LJL | 0.034 | 0.046 | 0.033 |
ln SNL–ln POPW | 0.052 | 0.097 | 0.068 |
SL–DFS counts | 0.515 | 0.424 | 0.299 |
SL–DFR counts | 0.649 | 0.607 | 0.420 |
SL–AFS counts | 0.000 | 0.109 | 0.086 |
SL–AFR counts | 0.626 | 0.629 | 0.581 |
SL–P1FR counts | 0.624 | 0.589 | 0.432 |
SL–LLS counts | 3.828 | 3.725 | 1.623 |
SL–SAL counts | 1.117 | 0.614 | 0.837 |
SL–SBL counts | 1.394 | 1.516 | 1.009 |
SL–UGR counts | 1.020 | 1.131 | 0.659 |
SL–LGR counts | 1.073 | 0.804 | 0.644 |
SL–TGR counts | 1.366 | 1.507 | 0.963 |
SL–AV counts | 0.155 | 0.279 | 0.191 |
SL–CV counts | 0.420 | 0.370 | 0.336 |
SL–TV counts | 0.426 | 0.414 | 0.333 |
The present study revealed that most body proportions of the three
On the other hand, taxonomic and related literature on
Proportional range comparisons of head length [
Differing growth-related proportional change patterns in the three
As in many other fishes (
Growth-related proportional change patterns of length-measured cephalic characters (based on SL and
The proportional values (percentages) of proportions subject to allometric growth are correlated with the base dimension (e.g., SL and
Because most of the length-measured characters of the three
Counts of pored scales on the lateral line (LLSs) and scales above the lateral line (SALs) tended to increase and decrease with growth, respectively, in
The growth-related status of dots / spots on the lateral body region also varied among the three
The proportional growth-related change pattern of pectoral scaly area length (PSAL) in
The CPAD proportion may be a useful feature for specific identification, since it can also be determined from illustrations and photographs of
In addition to caudal peduncle stoutness in
Caudal fin notch depth (
Among the length-measured cephalic characters of
The UJL and LJL plots for all three species (SL-based relationships) were well clustered around their regression curves (high negative allometry), but could not be distinguished from one another vertically (Fig.
The original description of
In addition to length-measured and meristic characters in the original description of
The diagnosis accompanying the original description of
Recent keys for identification of
On the other hand, post-orbital preopercular width (POPW) is a notable dimension, showing a contrasting pattern to SNL, i.e., plots of proportional POPW in small (< 200 mm SL)
Proportional differences between
Although
On the other hand, caudal and total vertebral counts (CV and TV, respectively), in which dominant counts were almost completely replaced between
Although
A morphological difference in the first anal pterygiophore (FAP) between
Standard errors (SEs) for the length-measured and meristic character regressions, which indicated degrees of morphological variation, were generally lowest in
The present study demonstrated a number of growth-related morphological changes in the three
a1 | Caudal peduncle anterior depth [% of standard length (SL)] > 15%. Snout length (% of SL) > 9%. Upper and lower jaw length [% of head length ( |
|
a2 | Caudal peduncle anterior depth (% of SL) ≤ 15%. Snout length (% of SL) ≤ 9%. Upper and lower jaw length (% of |
|
b1 | Post-orbital preopercular width (POPW) [% of snout length (SNL)] < 90% [POPW (% of SL) < 7.5% in specimens ≤ 200 mm SL; SNL (% of SL) > 7.7% in specimens > 200 mm SL]. Caudal vertebrae 20 (usually)–21 [rarely 19 (13.5%)]; total vertebrae 36 (usually)–37 [rarely 35 (13.5%)]. First anal pterygiophore modestly arched in specimens ≥ 90 mm SL. Spots / dots absent on body in specimens > 260 mm SL (although some specimens ≤ 260 mm SL have some dots restricted to upper part than lateral line) |
|
b2 | Post-orbital preopercular width (POPW) [% of snout length (SNL)] ≥ 90% [POPW (% of SL) ≥ 7.5% in specimens ≤ 200 mm SL; SNL (% of SL) ≤ 7.7% in specimens > 200 mm SL]. Caudal vertebrae 18–19 (usually) [rarely 20 (9.2%)]; total vertebrae 34–35 (usually) [rarely 36 (6.6%)]. First anal pterygiophore straight. Usually many clear black spots on lateral and dorsal body regions (usually even on lower part than lateral line) |
|
The author is grateful to Drs. Hiroshi Senou (