A new goatfish of the genus Upeneus (Mullidae) from Australia and Vanuatu, with inter- and intraspecific comparisons

Figures & data

Table I. Measurements and counts, for specimens of 65 mm SL and larger, of Upeneus torres sp. nov., U. australiae, U. guttatus, and U. japonicus.

	Upeneus torres sp. nov.										U. australiae				U. guttatus				U. japonicus
	Torres Strait		Paratypes (NW Australia)				All types > 64 mm SL				Australia				South Africa to Japan				E Malaysia to Korea
	HT	PT	Min	Mean	Max	n	Min	Mean	Max	n	Min	Mean	Max	n	Min	Mean	Max	n	Min	Mean	Max	n
Morphometric characters
SL	81	78	65	82.2	101	9	65	81.7	101	11	72	103.2	128	45	77	103.0	146	84	66	99.4	123	37
In %SL
Body depth at first dorsal-fin origin	25	23	24	25.4	27	9	23	25.1	27	11	23	23.8	27	38	22	23.5	26	73	21	23.5	25	37
Body depth at anal-fin origin	22	21	19	19.9	21	9	19	20.1	22	11	20	21.1	23	38	18	20.1	22	73	18	19.7	22	37
Half body depth at first dorsal-fin origin	21	20	20	20.9	22	4	20	20.8	22	6	18	19.6	22	35	17	19.3	21	63	18	19.7	21	27
Half body depth at anal-fin origin	17	15	15	15.9	17	4	15	16.0	17	6	15	16.3	18	33	14	15.8	18	59	14	16.0	18	26
Caudal-peduncle depth	11	9.8	8.9	9.7	11	9	8.9	9.8	11	11	9.9	10.6	12	45	9.1	9.9	11	73	8.0	9.6	11	37
Caudal-peduncle width	4.2	4.2	3.5	3.7	3.9	9	3.5	3.8	4.2	11	3.2	4.0	5.1	38	3.2	4.0	5.1	73	3.1	3.9	4.8	37
Maximum head depth	21	19	20	21.8	23	9	19	21.4	23	11	20	20.8	22	38	18	20.2	22	73	18	19.8	21	37
Head depth through eye	15	16	17	17.8	19	9	15	17.5	19	11	15	16.4	18	38	15	16.2	18	73	15	15.9	17	37
Suborbital depth	9.8	9.3	9.6	10.5	11	9	9.3	10.4	11	11	9.0	10.2	12	38	8.6	9.7	12	73	8.2	9.4	10	37
Interorbital length	8.2	7.2	7.5	7.9	8.2	9	7.2	7.9	8.2	11	7.1	8.0	9.6	38	7.0	7.8	8.9	73	6.9	7.7	8.7	37
Head length	28	28	29	30.3	31	9	28	29.9	31	11	27	28.9	30	38	26	27.6	30	73	27	28.6	31	37
Snout length	9.8	11	10	11.3	12	9	9.8	11.1	12	11	9.9	11.6	13	38	9.8	10.7	13	73	10	11.0	12	37
Postorbital length	12	10	11	12.3	13	9	10	12.1	13	11	11	11.5	13	38	9.7	11.1	12	73	9.8	11.1	13	37
Orbit length	7.4	7.4	7.5	8.0	8.9	9	7.4	7.9	8.9	11	6.0	6.9	8.0	38	5.9	7.1	8.8	73	6.1	7.4	8.2	37
Orbit depth	6.1	6.2	6.1	6.9	7.3	9	6.1	6.7	7.3	11	5.0	6.0	6.8	38	5.2	6.2	7.6	73	5.2	6.4	7.2	37
Upper-jaw length	11	11	10	11.6	12	9	10	11.4	12	11	9.3	10.9	12	38	9.5	10.9	12	73	9.7	10.6	12	37
Lower-jaw length	10	11	9.3	10.8	12	9	9.3	10.7	12	11	9.0	10.3	11	37	8.7	10.2	12	73	9.2	10.1	11	37
Snout width	8.3	8.5	7.9	9.3	10	6	7.9	9.1	10	8	7.9	8.7	10	35	7.6	8.6	11	70	7.1	8.4	9.8	37
Barbel length	24	24	24	24.4	26	9	24	24.3	26	11	16	18.4	20	45	16	17.9	20	71	18	21.1	23	36
Maximum barbel width	0.7	0.8	0.7	0.8	1.0	9	0.7	0.8	1.0	11	0.8	0.9	1.1	38	0.7	0.8	1.1	73	0.6	0.8	1.0	36
First pre-dorsal length	37	35	36	37.3	39	9	35	37.1	39	11	33	36.3	39	38	33	35.9	38	73	34	36.1	39	37
Second pre-dorsal length	63	66	63	65.2	67	9	63	65.1	67	11	61	63.6	66	38	60	63.5	68	73	61	63.7	67	37
Interdorsal distance	16	18	15	16.3	18	9	15	16.4	18	11	13	14.5	16	38	13	15.7	17	73	14	15.6	18	37
Caudal-peduncle length	23	23	21	23.1	25	9	21	23.1	25	11	22	24.1	26	38	22	23.7	26	73	21	23.2	25	37
Pre-anal length	62	60	64	64.8	66	9	60	64.0	66	11	60	63.4	67	38	60	64.4	68	73	61	65.1	69	37
Pre-pelvic length	32	32	32	34.6	37	9	32	34.1	37	11	30	32.4	35	38	28	31.3	34	73	30	32.5	35	37
Pre-pectoral length	30	31	31	32.4	34	9	30	32.1	34	11	29	30.2	32	38	27	29.5	33	73	29	31.0	33	37
Second dorsal-fin depth	22	21	20	20.9	22	9	20	21.0	22	11	20	21.5	24	38	19	20.8	23	73	18	20.4	22	37
Pelvic-fin depth	25	23	24	25.4	27	9	23	25.1	27	11	22	23.8	27	38	22	23.4	26	73	21	23.4	25	37
Pectoral-fin depth	16	16	16	17.0	18	9	16	16.8	18	11	15	16.7	19	38	14	16.2	19	73	14	16.2	18	37
Length of first dorsal-fin base	13	15	13	14.2	15	9	13	14.1	15	11	13	15.2	17	38	13	14.7	17	73	14	14.7	17	37
Length of second dorsal-fin base	14	12	12	13.3	14	9	12	13.2	14	11	12	13.6	15	38	11	13.5	15	73	12	13.4	15	37
Caudal-fin length	28	–	28	28.5	30	9	28	28.5	30	10	27	29.7	32	35	27	28.9	31	64	25	26.9	29	34
Length of anal-fin base	12	9.8	10	11.0	12	9	9.8	10.9	12	11	10	11.7	13	38	9.5	11.4	14	73	9.9	11.5	13	37
Anal-fin height	17	17	16	18.1	20	9	16	17.9	20	11	15	16.3	18	35	15	16.4	19	72	15	17.3	19	34
Pelvic-fin length	20	21	21	21.8	23	9	20	21.6	23	11	20	21.1	23	38	19	20.6	22	73	19	20.7	23	36
Pectoral-fin length	24	24	24	25.3	26	9	24	25.1	26	11	19	20.7	22	44	19	20.3	22	72	21	22.9	25	36
Pectoral-fin width	4.3	3.7	4.0	4.3	4.7	9	3.7	4.2	4.7	11	4.3	4.6	5.5	45	3.5	4.1	5.0	73	3.7	4.3	5.0	37
First dorsal-fin height	21	22.7	21	22.7	24	7	21	22.5	24	9	18	20.8	23	36	18	21.8	24	62	20	22.0	24	36
Second dorsal-fin height	16	17.5	16	17.2	19	8	16	17.1	19	10	14	15.9	18	36	14	16.0	18	68	15	17.4	19	34
Meristic characters
Pectoral-fin rays	14	14	13	14.0	15	9	13	14.0	15	11	13	14.2	15	45	12	13.2	15	84	13	13.9	15	37
Rudimentary gill rakers on upper limb	3	3	3	3.3	4	9	3	3.3	4	11	1	2.8	4	45	2	3.6	5	84	2	3.2	5	37
Developed gill rakers on upper limb	3	3	2	2.7	3	9	2	2.7	3	11	2	3.5	6	45	2	2.9	5	84	2	3.6	5	37
Developed gill rakers on lower limb	12	12	11	12.6	13	9	11	12.5	13	11	11	12.6	14	45	11	12.6	14	84	11	13.7	17	37
Rudimentary gill rakers on lower limb	5	6	4	4.7	6	9	4	4.8	6	11	3	4.2	6	45	3	4.7	6	83	3	5.3	7	37
Total gill rakers on upper limb	6	6	5	6.0	7	9	5	6.0	7	11	5	6.2	7	45	5	6.5	8	84	6	6.8	8	37
Total gill rakers on lower limb	17	18	16	17.2	18	9	16	17.3	18	11	16	16.8	18	45	16	17.3	19	84	18	19.1	21	37
Total gill rakers	23	24	22	23.2	25	9	22	23.3	25	11	22	23.1	24	45	23	23.8	26	84	24	25.9	28	37
Scales along lateral line	30	30	30	30.0	30	2	30	30.0	30	4	27	28.9	30	26	28	29.5	31	48	29	29.3	30	17

Table II. Measurements and counts, for specimens smaller than 65 mm SL, of Upeneus torres sp. nov., U. australiae, U. guttatus, and U. japonicus.

	Upeneus torres sp. nov												U. australiae				U. guttatus				U. japonicus
	Paratypes (NW Australia)				Vanuatu				All subadults				NW Australia				NW Australia				Vietnam
	Min	Mean	Max	n	Min	Mean	Max	n	Min	Mean	Max	n	Min	Mean	Max	n	Min	Mean	Max	n	Min	Mean	Max	n
Morphometric characters
SL	33	44.3	61	4	36	51.0	61	12	33	49.3	61	16	24	27.9	33	4	34	40.9	54	8	51	54.9	59	3
In %SL
Body depth at first dorsal-fin origin	23	23.9	24	4	21	22.1	23	12	21	22.5	24	16	23	24.0	25	4	22	24.4	26	8	21	22.3	23	3
Body depth at anal-fin origin	18	18.8	20	4	17	17.4	19	12	17	17.7	20	16	19	20.4	22	4	18	19.3	21	8	18	18.7	20	3
Half body depth at first dorsal-fin origin	–	–	–	0	17	17.7	19	10	17	17.7	19	10	19	19.7	20	3	18	19.0	20	2	–	–	–	0
Half body depth at anal fin-origin	–	–	–	0	12	13.5	15	8	12	13.5	15	8	16	15.9	16	2	–	–	–	0	–	–	–	0
Caudal-peduncle depth	8.9	9.0	9.1	4	8.2	8.6	9.1	12	8.2	8.7	9.1	16	10	10.7	11	4	8.4	9.0	9.6	8	9.3	9.4	9.6	3
Caudal-peduncle width	3.0	3.4	3.9	4	2.8	3.0	3.2	12	2.8	3.1	3.9	16	3.2	3.5	3.7	3	3.1	3.5	3.9	7	3.6	3.8	4.1	3
Maximum head depth	21	21.3	22	4	18	19.9	21	12	18	20.2	22	16	22	22.2	23	4	21	21.7	23	8	18	18.7	20	3
Head depth through eye	17	18.1	19	4	16	17.0	18	12	16	17.3	19	16	19	19.3	20	3	18	18.4	19	8	16	16.2	16	3
Suborbital depth	8.0	8.8	9.7	4	8.2	9.3	9.9	12	8.0	9.2	9.9	16	11	11.1	12	3	7.4	8.9	10	8	8.6	8.8	9.1	3
Interorbital length	7.4	7.8	8.2	4	6.8	7.5	8.0	12	6.8	7.6	8.2	16	8.3	8.6	9.0	4	7.5	8.0	8.3	8	7.2	7.4	7.8	3
Head length	28	31.5	34	4	29	30.3	31	12	28	30.6	34	16	31	31.7	32	4	29	30.6	32	8	29	30.0	31	3
Snout length	10	10.8	11	4	9.8	10.4	12	12	9.8	10.5	12	16	9.0	9.5	10	4	9.1	10.1	11	8	9.9	10.3	11	3
Postorbital length	11	12.7	14	4	11	11.7	12	12	11	11.9	14	16	13	13.9	14	4	11	12.5	13	8	11	11.7	12	3
Orbit length	8.8	9.2	9.7	4	8.3	8.9	9.6	12	8.3	9.0	9.7	16	9.4	9.5	9.8	4	7.8	8.4	9.0	8	8.2	8.6	9.3	3
Orbit depth	7.7	8.2	8.7	4	7.4	7.9	8.3	12	7.4	8.0	8.7	16	8.2	8.6	8.9	4	7.2	7.6	8.5	8	7.5	7.8	8.3	3
Upper-jaw length	11	12.4	13	4	10	10.9	11	12	10	11.3	13	16	11	11.3	12	3	11	12.4	13	8	11	11.2	12	3
Lower-jaw length	10	11.5	12	4	9.3	9.9	11	12	9.3	10.3	12	16	11	10.7	11	3	11	11.4	12	8	10	10.8	11	3
Snout width	8.1	8.4	8.6	4	6.8	8.1	8.8	3	6.8	8.2	8.8	7	8.4	9.0	9.5	3	7.0	8.0	8.9	8	7.6	8.1	8.3	3
Barbel length	25	26.1	28	4	24	26.2	28	12	24	26.2	28	16	16	17.3	19	4	19	19.9	21	8	20	21.1	22	3
Maximum barbel width	0.7	0.8	0.9	4	0.7	0.9	0.9	12	0.7	0.8	0.9	16	0.8	0.9	0.9	3	0.7	0.8	0.9	8	0.7	0.8	0.8	3
First pre-dorsal length	36	37.7	40	4	35	36.5	38	12	35	36.8	40	16	38	38.2	38	3	34	36.3	38	8	35	36.4	38	3
Second pre-dorsal length	62	62.4	64	4	61	64.0	65	12	61	63.6	65	16	65	64.6	65	3	62	63.9	66	8	63	64.4	66	3
Interdorsal distance	13	14.4	17	4	13	14.4	16	12	13	14.4	17	16	14	14.4	15	3	13	15.1	16	8	16	16.4	17	3
Caudal-peduncle length	20	20.6	21	4	23	24.3	26	12	20	23.4	26	16	25	24.7	25	3	20	22.0	24	8	21	22.2	23	3
Pre-anal length	60	63.6	65	4	63	65.4	69	12	60	64.9	69	16	63	65.0	66	3	60	63.9	66	8	63	63.8	65	3
Pre-pelvic length	34	34.6	35	4	33	36.0	38	12	33	35.6	38	16	35	35.3	36	3	32	33.8	35	8	32	32.9	34	3
Pre-pectoral length	31	32.6	34	4	31	33.3	35	12	31	33.1	35	16	32	33.2	34	3	30	32.4	34	8	32	32.7	33	3
Second dorsal-fin depth	18	18.9	20	4	16	17.2	18	12	16	17.6	20	16	20	20.9	21	3	18	19.3	21	8	18	19.0	20	3
Pelvic-fin depth	23	24.0	25	4	20	21.2	22	12	20	21.9	25	16	23	23.9	25	3	23	24.5	27	8	20	21.2	22	3
Pectoral-fin depth	14	15.5	17	4	13	13.7	14	12	13	14.1	17	16	15	15.6	16	3	15	15.9	17	8	14	14.6	15	3
Length of first dorsal-fin base	14	14.7	15	4	13	14.1	15	12	13	14.2	15	16	14	14.4	15	3	15	15.6	16	8	14	14.4	14	3
Length of second dorsal-fin base	13	13.9	15	4	12	13.1	14	12	12	13.3	15	16	14	13.8	14	3	14	14.1	15	8	13	13.7	15	3
Caudal-fin length	29	31.6	34	4	29	30.7	32	7	29	31.0	34	11	29	30.0	31	3	31	32.4	33	8	29	29.5	30	3
Length of anal-fin base	11	11.7	13	4	11	12.5	13	12	11	12.3	13	16	11	11.6	12	3	12	12.4	13	8	11	11.5	12	3
Anal-fin height	17	18.8	20	4	18	18.5	20	12	17	18.6	20	16	16	16.9	18	3	17	18.4	20	8	17	17.3	17	3
Pelvic-fin length	21	21.8	22	4	21	22.0	24	12	21	22.0	24	16	20	20.8	21	3	21	21.7	23	8	21	21.9	22	3
Pectoral-fin length	25	25.5	26	4	24	24.6	26	12	24	24.8	26	16	21	22.2	23	4	22	22.3	23	8	23	23.2	23	3
Pectoral-fin width	4.0	4.3	4.6	4	3.7	4.0	4.5	12	3.7	4.1	4.6	16	4.0	4.1	4.2	3	3.6	3.9	4.1	8	4.2	4.4	4.7	3
First dorsal-fin height	23	23.5	24	3	20	22.0	24	12	20	22.3	24	15	21	21.8	22	3	19	21.7	24	7	23	23.1	24	3
Second dorsal-fin height	16	18.3	20	4	17	17.4	18	8	16	17.7	20	12	17	17.0	17	3	17	18.1	19	6	18	18.3	19	3
Meristic characters
Pectoral-fin rays	14	14.0	14	4	13	13.6	14	12	13	13.7	14	16	14	14.3	15	4	13	13.0	13	8	13	13.7	14	3
Rudimentary gill rakers on upper limb	2	3.0	4	4	1	2.3	3	12	1	2.4	4	16	0	1.5	4	4	3	3.5	4	8	3	3.3	4	3
Developed gill rakers on upper limb	2	2.5	3	4	3	4.0	5	12	2	3.6	5	16	3	5.3	6	4	2	2.5	3	8	3	3.3	4	3
Developed gill rakers on lower limb	12	12.5	14	4	14	14.6	15	12	12	14.1	15	16	14	14.5	15	4	12	12.5	13	8	13	13.7	14	3
Rudimentary gill rakers on lower limb	5	5.3	6	4	2	3.3	4	12	2	3.8	6	16	2	2.8	3	4	4	4.9	6	8	4	4.7	5	3
Total gill rakers on upper limb	5	5.5	6	4	6	6.3	7	12	5	6.1	7	16	6	6.8	7	4	5	6.0	7	8	6	6.7	7	3
Total gill rakers on lower limb	17	17.8	19	4	17	17.8	19	12	17	17.8	19	16	17	17.3	18	4	16	17.4	18	8	18	18.3	19	3
Total gill rakers	22	23.3	25	4	23	24.1	25	12	22	23.9	25	16	23	24.0	25	4	23	23.4	24	8	25	25.0	25	3
Scales along lateral line	–	–	–	0	29	29.2	30	5	29	29.2	30	5	28	28.0	28	4	–	–	–	0	–	–	–	0

Figure 1. (a) Upeneus torres sp. nov., holotype (CSIRO H 7202-02, 81 mm SL; Daniel Gledhill); (b) U. torres sp. nov., paratype (CSIRO CA 2067, 98 mm SL; photographer unknown); (c) U. japonicus (KAUM 41736, 119 mm SL; Hiroyuki Motomura); (d) U. australiae (CSIRO H4056-01, 105 mm SL; Alastair Graham); (e) U. guttatus (CSIRO CA 3044, 101 mm SL; photographer unknown); (f) U. guttatus (CSIRO H 7212-02, 121 mm SL, Daniel Gledhill).

Figure 2. Collection localities for specimens of Upeneus torres sp. nov., symbols may represent multiple lots. The approximate distributional ranges, including overlapping ranges, are also depicted for U. australiae, U. guttatus and U. japonicus.

Figure 3. Relationships of morphometric and meristic characters in Upeneus torres sp. nov. and U. japonicus. Specimens of both species from different areas are distinguished by symbols.

Figure 4. Meristic characters for the four Upeneus species, with results of pairwise comparisons indicating significant differences based on χ² test for trends (significance level ≤ 0.01); the letter symbols appearing as the uppermost x-axis labels refer to the respective species below and in the graphs they indicate differences with the respective species.

Figure 5. Depth–size relationships for the four Upeneus species.

Figure 6. Relationships of morphometric and meristic characters in U. australiae and U. guttatus. Specimens of U. guttatus from different areas are distinguished by symbols.