Corresponding author: Kōji Yokogawa (
Academic editor: Nina Bogutskaya
Morphological differences, including growthrelated 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.




Postorbital preopercular width  POPW  
Standard length  SL  Upper jaw length  UJL 
Preanus length 

Lower jaw length  LJL 
Body depth  BD 


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 
Predorsal length 

Pectoral fin ray  P_{1}FR 
First dorsal fin (longest spine) length  FDFL  Pelvic fin spine  P_{2}FS 
Second dorsal fin (longest ray) length 

Pelvic fin ray  P_{2}FR 
Caudal fin length  CFL  Pored scale on lateral line  LLS 
Caudal fin notch depth 

Scale above lateral line  SAL 
Anal fin (longest ray) length 

Scale below lateral line  SBL 
Pectoral fin length  P_{1}FL  Upperlimb gill raker  UGR 
Pelvic fin length  P_{2}FL  Lowerlimb gill raker  LGR 
Pectoral scaly area length  PSAL  Total gill raker  TGR 
Head length 

Abdominal vertebra  AV 

Caudal vertebra  CV  
Snout length  SNL  Total vertebra  TV 
Orbital diameter  OD 


Interorbital width  IOW  Dorsocephalic scale row  DSR 
Suborbital width  SOW  Ventromandibular scale row  VSR 
Postorbital 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 finspine (P_{2}FS) and soft rays (P_{2}FRs) showed no variation (P_{2}FS: 1, P_{2}FRs: 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 middorsal aspect of the caudal peduncle (Fig.
For a lengthmeasured dimension (LD), a growthrelated 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 (















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–P_{1}FL/SL  25.19  0.070  0.581  16.98  0.010  0.109  19.79  0.024  0.270 
SL–P_{2}FL/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/SL^{1}  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 interspecific differences in lengthmeasured characters, regressions between SL (or
Regression parameters (slope and intercept) and correlation between logarithmtransformed lengthmeasured characters, together with results of















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 P_{1}FL  0.930  1.38  10.73***  0.990  1.77  1.41  0.976  1.62  3.25* 
ln SL–ln P_{2}FL  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 HolmBonferroni 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







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–P_{1}FR 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* 


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–P_{1}FR 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 


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–P_{1}FR 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 HolmBonferroni 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 lengthmeasured body characters of three
Relationships between standard length and proportions of some lengthmeasured body characters which exhibited prominent plot separation among three
Similar patterns of growthrelated 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 lengthmeasured dimensions (LD) of cephalic characters, SLbased (SL–LD / SL) and
Relationships between standard length or head length and proportions of lengthmeasured cephalic characters of three
Growthrelated proportional change patterns based on SL and
As well as some body characters, specific proportional change patterns were recognized for some characters, e.g., SLbased 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 (postorbital preopercular width) / (snout length) proportions of three
The
Relationships between standard length and some meristic characters which exhibited growthrelated 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
Postjuvenile 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













≤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 


≤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 


≤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 logarithmtransformed lengthmeasured characters by pairwise comparisons for the three
Results of analysis of covariance (ANCOVA) (
Regression  

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 P_{1}FL  6.17***  –  4.24**  –  1.26  12.21*** 
ln SL–ln P_{2}FL  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 HolmBonferroni correction. Bars indicate that calculation was not demonstrated because significance was recognized for the slope and ANCOVA was therein terminated.
Although the MannWhitney
Results of the MannWhitney




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*** 
P_{1}FR 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 HolmBonferroni correction.
Standard errors (SEs) for regression lines between logarithmtransformed SL and lengthmeasured characters, and between SL and meristic characters are summarized in Table
Standard errors for morphological character regressions of three




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 P_{1}FL  0.056  0.063  0.045 
ln SL–ln P_{2}FL  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–P_{1}FR 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 growthrelated proportional change patterns in the three
As in many other fishes (
Growthrelated proportional change patterns of lengthmeasured 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 lengthmeasured 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 growthrelated status of dots / spots on the lateral body region also varied among the three
The proportional growthrelated 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 lengthmeasured cephalic characters of
The UJL and LJL plots for all three species (SLbased 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 lengthmeasured and meristic characters in the original description of
The diagnosis accompanying the original description of
Recent keys for identification of
On the other hand, postorbital 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 lengthmeasured and meristic character regressions, which indicated degrees of morphological variation, were generally lowest in
The present study demonstrated a number of growthrelated morphological changes in the three
a^{1}  Caudal peduncle anterior depth [% of standard length (SL)] > 15%. Snout length (% of SL) > 9%. Upper and lower jaw length [% of head length ( 

a^{2}  Caudal peduncle anterior depth (% of SL) ≤ 15%. Snout length (% of SL) ≤ 9%. Upper and lower jaw length (% of 

b^{1}  Postorbital 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) 

b^{2}  Postorbital 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 (