A new goatfish, Upeneus seychellensis sp. nov. (Mullidae), from the Seychelles Bank, with remarks on Upeneus guttatus and a key to Western Indian Ocean Upeneus species

Figures & data

Figure 1.  A: Upeneus seychellensis sp. nov., holotype, SAIAB 96980, SL 115 mm, and (above) paratype, SAIAB 84280, SL 102 mm, Seychelles Bank (O. Alvheim); B: Upeneus guttatus, SAIAB 84281, female, SL 117 mm, and (below) male, SL 111 mm, Seychelles Bank (O. Alvheim).

Figure 2.  Body depth at anal-fin origin against SL (A), first dorsal-fin height against caudal-peduncle depth (B), anal-fin height against number of pectoral-fin rays (C), and length of anal-fin base against total number of gill rakers (D), in the four Indian Ocean species of the japonicus group (morphometric variables expressed in % SL).

Table I. Measurements and counts for Upeneus seychellensis sp. nov. and the species of the japonicus group (morphometric characters in % SL; main differences with U. seychellensis are emphasized).

	Upeneus seychellensis sp. nov.			U. guttatus			U. pori			U. australiae			U. asymmetricus			U. francisi	U. japonicus
	HT	PT1	PT2	Min	Max	n	Min	Max	n	Min	Max	n	Min	Max	n	n=1	Min	Max	n
SL (mm)	115	102	96	77	146	44	66	110	5	84	130	12	74	100	4	78	104	115	4
Body depth at first dorsal-fin origin	22	21	20	22	26	37	21	24	5	23	26	7	22	24	4	24	23	25	4
Body depth at anal-fin origin	19	18	18	19	23	37	20	22	5	20	23	7	20	22	4	19	19	22	4
Half body depth (from lateral line downwards) at first dorsal fin origin	19	19	17	17	21	33	18	20	5	17	22	8	19	20	4	18	19	21	3
Half body depth (from lateral line downwards) at anal fin origin	15	14	14	14	18	33	14	16	5	16	18	8	14	16	4	15	15	17	4
Caudal-peduncle depth	9.6	9.3	9.2	9.3	11	37	9.1	10	5	10	11	8	8.8	9.9	4	9.9	8.7	10	4
Caudal-peduncle width	3.6	3.8	4.1	3.2	5.1	37	3.5	3.9	5	3.6	5.0	8	3.7	4.8	4	2.3	3.1	5.7	4
Maximum head depth	20	19	18	18	23	37	18	20	5	20	22	8	20	20	4	20	20	21	4
Head depth across a vertical midline through eye	17	15	15	15	19	37	15	16	5	15	19	8	15	16	4	18	16	16	4
Suborbital depth	10	10	9.2	8.7	12	37	8.9	9.7	5	9.5	13	8	9.4	10	4	8.4	9.1	10	4
Interorbital length	7.7	6.8	6.7	7.2	9.0	37	7.4	7.9	5	7.3	9.5	8	8.0	9.0	4	7.6	7.2	8.5	4
Head length	30	28	27	26	30	37	26	28	5	27	30	8	27	29	4	31	28	29	4
Snout length	12	12	11	9.6	13	37	11	11	5	11	13	8	9.9	11	4	10	11	12	4
Postorbital length	12	12	12	10	12	36	11	12	5	11	13	8	11	13	4	13	10	12	4
Orbit length	6.5	6.4	6.0	6.3	8.8	37	6.3	7.8	5	6.1	7.9	8	7.0	7.7	4	8.2	6.8	8.0	4
Orbit depth	6.2	5.6	5.5	5.2	7.6	37	5.7	7.7	5	5.0	6.6	8	5.6	6.5	4	6.7	5.2	7.2	4
Upper-jaw length	11	11	11	9.6	12	37	10	11	5	10	12	8	9.6	11	4	12	10	11	4
Lower-jaw length	11	11	10	9.0	12	37	9.7	10	5	9.2	11	8	9.1	10	4	11	9.6	9.9	4
Snout width	9.2	8.0	7.3	7.7	11	35	8.0	10	5	8.6	10	6	8.1	9.3	4	7.0	7.7	9.2	4
Barbel length	22	17	17	16	19	36	16	18	5	17	19	8	18	19	4	18	18	22	4
Maximum barbel width	0.8	0.7	0.8	0.7	1.0	37	0.6	0.7	5	0.8	1.3	8	0.8	1.0	4	0.8	0.6	0.8	4
First pre-dorsal length	39	37	38	33	38	37	34	37	5	34	37	8	37	38	4	38	34	37	4
Second pre-dorsal length	63	64	65	60	67	37	63	65	5	62	64	8	64	66	4	66	63	64	4
Interdorsal distance	12	14	16	13	17	37	13	16	4	13	16	8	14	16	4	15	15	18	4
Caudal-peduncle length	24	24	24	22	26	37	22	23	5	22	26	8	22	24	4	23	23	25	4
Pre-anal length	68	67	65	60	68	37	60	65	5	60	64	8	63	68	4	66	61	67	4
Pre-pelvic length	33	32	30	28	34	37	28	32	5	30	36	8	32	34	4	33	30	33	4
Pre-pectoral length	32	30	28	26	32	37	28	30	5	29	32	8	30	31	4	33	27	33	4
Second dorsal-fin depth (second dorsal-fin origin to anal-fin origin)	19	19	18	19	23	37	20	22	5	20	24	8	19	22	4	20	19	22	4
Pelvic-fin depth (first dorsal-fin origin to pelvic-fin origin)	23	22	21	22	26	37	22	23	5	23	27	8	22	24	4	24	22	25	4
Pectoral-fin depth (first dorsal-fin origin to dorsal origin of pectoral fin)	17	16	17	15	18	37	16	17	5	16	18	8	15	17	4	16	14	18	4
Length of first dorsal-fin base	14	14	14	12	18	37	14	16	4	14	17	8	13	15	4	14	14	17	4
Length of second dorsal-fin base	13	12	12	11	16	37	13	15	5	13	15	8	13	14	4	11	13	15	4
Caudal-fin length (dorsal caudal-fin origin to upper caudal-lobe tip	30	29	28	25	30	37	27	29	4	26	30	7	27	27	4	28	25	27	4
Length of anal-fin base	10	10	9.6	9.3	14	37	12	13	5	11	13	8	10	12	4	12	11	12	4
Anal-fin height	14	15	15	15	19	36	16	17	5	14	17	7	15	16	2	16	15	18	4
Pelvic-fin length	21	20	21	19	22	37	20	23	5	20	22	8	20	21	4	24	21	22	4
Pectoral-fin length	21	21	21	19	22	36	20	22	5	21	24	8	19	21	3	25	24	24	4
Pectoral-fin width (width of pectoral-fin base)	3.9	3.9	4.2	3.7	5.0	37	4.4	5.7	5	4.2	5.9	8	4.1	5.9	4	4.2	3.7	5.8	4
First dorsal-fin height	19	20	20	20	24	33	20	21	4	18	21	8	20	20	3	21	21	22	4
Second dorsal-fin height	17	16	16	14	18	36	15	16	4	14	17	12	15	16	2	16	16	18	4
Pectoral-fin rays	14	15	15	12	14	44	14	14	5	13	15	12	13	13	4	14	13	15	4
Rudimentary gill rakers on upper limb	5	4	4	2	5	44	0	2	5	1	4	12	1	4	4	1	2	3	4
Developed gill rakers on upper limb	2	3	3	2	5	44	6	7	5	2	5	12	3	6	4	8	4	4	4
Developed gill rakers on lower limb	13	13	13	11	14	44	14	18	5	11	14	12	15	17	4	23	12	16	4
Rudimentary gill rakers on lower limb	6	5	5	4	6	44	2	5	5	3	5	12	2	5	4	0	4	7	4
Total gill rakers on upper limb	7	7	7	6	8	44	7	8	5	5	7	12	7	7	4	9	6	7	4
Total gill rakers on lower limb	19	18	18	16	18	44	18	20	5	16	17	12	19	21	4	23	18	20	4
Total gill rakers	26	25	25	23	25	44	26	27	5	21	24	12	26	28	4	32	24	27	4
Scales along lateral line to caudal-fin base	31	29	29	28	31	44	29	30	5	28	31	9	28	29	4	28	28	29	2

Figure 3.  Upeneus guttatus: A: SAIAB 84255, female, SL 112 mm, Seychelles Bank (O. Alvheim); B: SAIAB 82714, SL 94 mm, Mozambique (P.C. Heemstra); C: SAIAB 13947, SL 100 mm, Kenya (P.C. Heemstra); D: BPBM 20658, SL 113 mm, Madras, India (J.E. Randall).

Figure 4.  A: Scores of the first and third axis of Principal Component Analysis based on 40 morphometric characters in 37 Upeneus guttatus from different areas in the Indian Ocean, with trends for the highest-contributing characters indicated by arrows; B: caudal-fin length against snout length (both variables expressed in % SL).

Table II. Measurements and counts for the Indian Ocean Upeneus guttatus populations (morphometric characters in % SL).

	Seychelles				East-Africa				Red Sea		Sri Lanka	Madras	Andamans	Malacca (n = 2)		NW Australia
	Min	Mean	Max	n	Min	Mean	Max	n	Suez	Sudan	n=1	n=1	n=1	Min	Max	n=1
SL (mm)	92	107	126	17	87	103	146	19	98	77	84	113	91	97	108	93
Body depth at first dorsal-fin origin	22	23	24	13	22	24	25	16	23	23	24	23	22	24	25	26
Body depth at anal-fin origin	19	20	21	13	19	20	23	16	20	21	20	21	20	21	22	20
Half body depth (from lateral line downwards) at first dorsal fin origin	18	19	21	13	17	19	21	13	18	19	21	21	–	20	20	20
Half body depth (from lateral line downwards) at anal fin origin	15	15	16	13	15	16	18	13	15	16	15	17	–	17	18	17
Caudal-peduncle depth	9.4	10	10	13	9.3	10	11	16	9.5	11	10	10	9.4	10	11	10
Caudal-peduncle width	3.2	3.6	4.2	13	3.5	4.4	5.1	16	3.4	5.1	3.5	3.3	3.4	3.8	4	4
Maximum head depth	19	20	22	13	18	20	21	16	20	19	22	20	20	21	22	23
Head depth across a vertical midline through eye	16	16	19	13	15	16	19	16	15	17	16	16	16	16	16	17
Suborbital depth	9.0	9.6	12	13	8.7	9.8	12	16	8.7	9.9	9.7	11	9.1	9.7	11	10
Interorbital length	7.3	7.6	8.0	13	7.2	7.8	8.4	16	7.3	7.9	8.3	7.8	8.2	8.7	9.0	7.7
Head length	26	27	28	13	26	28	29	16	26	26	29	30	28	28	28	28
Snout length	9.6	10	11	13	9.9	11	13	16	10	11	12	12	11	11	11	11
Postorbital length	10	11	11	13	10	11	12	15	11	11	12	12	11	10	11	11
Orbit length	6.3	6.9	7.8	13	6.4	7.5	8.8	16	7.0	8.2	6.3	6.5	6.3	7.3	7.4	7.2
Orbit depth	5.4	6.0	6.6	13	5.2	6.4	7.6	16	6.1	5.9	6.0	5.8	5.7	6.3	6.9	5.7
Upper-jaw length	9.7	10	12	13	9.6	11	12	16	10	12	11	12	12	11	11	12
Lower-jaw length	9.0	9.7	10	13	9.5	10	12	16	9.6	11	10	11	11	10	11	11
Snout width	7.7	8.5	10	13	7.8	9.0	10	14	8.3	11	10	8.7	10	8.8	10	8.3
Barbel length	16	17	18	13	16	17	19	15	18	16	19	18	18	17	18	19
Maximum barbel width	0.7	0.7	0.9	13	0.7	0.8	1.0	16	0.7	0.8	0.8	0.9	0.8	0.8	0.9	0.9
First pre-dorsal length	33	35	37	13	34	36	38	16	35	38	36	37	37	35	36	35
Second pre-dorsal length	60	63	65	13	61	63	67	16	60	65	63	65	63	62	66	64
Interdorsal distance	15	16	16	13	14	16	17	16	13	17	15	16	16	13	17	17
Caudal-peduncle length	22	24	26	13	22	24	26	16	23	26	24	23	22	22	23	25
Pre-anal length	61	65	67	13	60	63	68	16	64	65	63	64	63	63	65	62
Pre-pelvic length	28	31	34	13	28	31	34	16	31	31	33	33	32	30	32	30
Pre-pectoral length	26	28	30	13	26	29	32	16	28	28	30	32	30	28	30	28
Second dorsal-fin depth (second dorsal-fin origin to anal-fin origin)	20	21	22	13	19	21	23	16	19	21	21	22	20	22	22	21
Pelvic-fin depth (first dorsal-fin origin to pelvic-fin origin)	22	23	25	13	22	24	26	16	22	23	23	24	22	24	24	26
Pectoral-fin depth (first dorsal-fin origin to dorsal origin of pectoral fin)	15	17	18	13	15	16	18	16	15	18	16	17	16	17	18	18
Length of first dorsal-fin base	12	15	16	13	12	15	17	15	14	17	15	13	13	15	17	15
Length of second dorsal-fin base	11	13	14	13	12	14	16	16	12	16	14	13	14	13	14	12
Caudal-fin length (dorsal caudal-fin origin to upper caudal-lobe tip	28	29	30	13	25	28	30	16	29	26	27	27	30	25	28	30
Length of anal-fin base	9.3	11	13	13	11	12	14	16	11	13	12	10	12	10	11	11
Anal-fin height	15	16	17	13	15	17	19	15	16	17	17	15	17	15	16	16
Pelvic-fin length	19	21	22	13	19	21	22	16	20	21	20	21	22	20	22	21
Pectoral-fin length	19	20	21	13	20	21	22	15	20	21	20	21	22	21	22	20
Pectoral-fin width (width of pectoral-fin base)	3.7	3.9	4.2	13	3.8	4.4	5.0	16	3.9	4.3	4.3	4.6	3.9	4.0	4.7	4.2
First dorsal-fin height	20	22	24	13	20	22	24	12	22	20	20	21	21	22	23	22
Second dorsal-fin height	14	16	17	13	14	16	18	15	15	16	16	17	17	16	16	15
Pectoral-fin rays	12	13	14	17	13	13	14	19	13	13	13	14	13	13	13	13
Rudimentary gill rakers on upper limb	2	3.7	5	17	2	3.3	5	19	4	4	4	3.0	4	3	3	4
Developed gill rakers on upper limb	2	2.7	4	17	2	3.3	5	19	2	4	3	3.0	2	3	3	2
Developed gill rakers on lower limb	11	12	14	17	11	13	14	19	12	13	12	12	12	13	13	12
Rudimentary gill rakers on lower limb	4	4.8	6	17	4	4.3	5	19	6	4	4	5	5	4	4	5
Total gill rakers on upper limb	6	6.4	7	17	6	6.6	8	19	6	8	7	6	6	6	6	6
Total gill rakers on lower limb	17	17	18	17	16	17	18	19	18	17	16	17	17	17	17	17
Total gill rakers	23	24	25	17	23	24	25	19	24	25	23	23	23	23	23	23
Scales along lateral line to caudal-fin base	29	30	31	11	28	29	31	13	29	29	29	31	–	30	30	30