Siphamia papuensis, Gon, Ofer, Allen, Gerald R., Erdmann, Mark V. & Gouws, Gavin, 2014

Gon, Ofer, Allen, Gerald R., Erdmann, Mark V. & Gouws, Gavin, 2014, A new species of the cardinalfish genus Siphamia (Perciformes, Apogonidae) from West Papua, Indonesia, Zootaxa 3881 (4), pp. 328-340 : 329-338

Siphamia papuensis , new species

Figures 1 a–d; Table 1

Siphamia species 1 Allen and Erdmann, 2012: 414, Misool Island, West Papua, Indonesia.

Species Voucher specimen Locality GenBank

16S rDNA COI Siphamia argentea SAIAB 194660 Angel’s Window dive site, Lembeh Strait, North Sulawesi, Indonesia KM257072 -

SAIAB 194661 Crinoid City dive site, Milne Bay, Papua New Guinea KM257073 - Siphamia jebbi [KUT 4635] Watering Bay, north side of Yadua Island, Fiji KM257074 - Siphamia papuensis n. sp. AMS I.46420-001 Fiabacet, SE Misool, Raja Ampat Islands, West Papua, Indonesia KM257066 KM357915

AMS I.46420-001 Fiabacet, SE Misool, Raja Ampat Islands, West Papua, Indonesia KM257067 KM357916

BMNH 2014.7.21.1 Nudi Rock dive site, SE Misool Island, Raja Ampat Islands, West Papua, KM257068 KM357917

Indonesia

SAIAB 194663 Nudi Rock dive site, SE Misool Island, Raja Ampat Islands, West Papua, KM257069 KM357918

Indonesia

SAIAB 194663 Nudi Rock dive site, SE Misool Island, Raja Ampat Islands, West Papua, KM257070 KM357919

Indonesia

SAIAB 194704 Pulau Panjang, Fakfak Peninsula, West Papua, Indonesia KM257071 KM357920 Siphamia stenotes WAM P.32799-001 Selat Iris, West Papua, Indonesia KM257075 - Siphamia tubifer SAIAB 194705 Pulau Panjang, Fakfak Peninsula, West Papua, Indonesia KM257076 -

SAIAB 194757 Mergui Archipelago, Myanmar KM257077 -

SAIAB 99441 Socotra, Yemen, Indian Ocean KM257078 -

Outgroups

Apogonichthyoides pharaonis HUJ 19978 Haifa Bay, Israel KM257080 - Fibramia lateralis KU 31867 Davu village, East of Nausori, NE of Suva, Viti Levu, Fiji KM257079 - Holotype. MZB 22100, 27.8 mm, female, Indonesia, West Papua Province, Raja Ampat Islands, SE Misool, Pulau Balbal (02°14.206ʹS, 130°30.673ʹE), 66 m, M. Erdmann, 17 February 2011.

Paratypes. AMS I.46420-001, 2: 21.95–26.5 mm, male and female, respectively, and BPBM 41176, 22.9 mm, male, Indonesia, West Papua Province, Raja Ampat Islands, SE Misool, Fiabacet (02°13.136ʹS, 130°34.063ʹE), 60 m, clove oil and hand net, M. Erdmann, 19 January 2013. BMNH 2014.7.21.1, 27.3 mm, female, Indonesia, West Papua Province, Raja Ampat Islands, SE Misool, Nudi Rock dive site (02°13.196ʹS, 130°33.834ʹE), 60–70 m, M. Erdmann, 14 September 2013. NSMT-P 118258, 26.45 mm, female, ROM 99789, 22.3 mm, male, and SAIAB 194663, 7: 19.35–23.1 mm, males, all collected with BMNH fish. SAIAB 194662, 3: 22.8–26.05 mm, two males and one female, respectively, and USNM 427502, 2: 22.05–27.7 mm, male and female, respectively, collected with AMS fish. SAIAB 194704, 3: 14.6–18.5 mm, 2 females and male, respectively, Indonesia, West Papua Province, FakFak Peninsula, Pulau Panjang (02°57.922ʹS, 131°20.067ʹE), 70 m, rotenone, M. Erdmann and G. Allen, 21 March 2009. WAM P.33386-001, 2: 20.2–24.8 mm, collected with holotype. WAM P.33387-001, 4: 18.45–24.5 mm, Indonesia, West Papua Province, Raja Ampat Islands, SE Misool, Nudi Rock dive site (02°13.196ʹS, 130°33.834ʹE), 66 m, M. Erdmann, 28 February 2011. WAM 34019-001, 2: 17.7–18.8 mm, female and male, respectively, Indonesia, West Papua Province, Triton Bay, Namatote Island (03°46.729ʹS, 133°52.925ʹE), 50 m, clove oil and hand net, M. Erdmann, 10 January 2008.

Comparative material of Siphamia argentea . Material listed in Gon and Allen (2012), as well as BPBM 40777, 32.0 mm, Vanuatu, Esperitu Santo, Station 36–41, R. Pyle, 2006. WAM P.33477-001, 31.6 mm, Indonesia, Bali, Menjangan Island (08°05.467ʹS, 114°30.131ʹE), 70 m, M. Erdmann, 9 May 2011. SAIAB 194661, 24.5 mm, Papua New Guinea, Milne Bay Province, Crinoid City dive site (10°18.689ʹS, 150°58.494ʹE), 50 m, clove oil and hand net, M. Erdmann and G. Allen, 13 June 2013. SAIAB 194660, 2: 16.4–23.1 mm and WAM P.33892-004, 5: 21.9–29.0 mm, Indonesia, North Sulawesi Province, Lembeh Strait, Angel’s Window dive site (01°30.029ʹN, 125°15.701ʹE), 65–68 m, clove oil and hand net, M. Erdmann, 23–24 July 2013.

Diagnosis. A species of Siphamia with a striated light organ, 6–7 first dorsal-fin spines, 13 (rarely 12) pectoral-fin rays, 2–9 tubed lateral-line scales, 1 developed gill raker on upper limb and 8–9 developed ones on lower limb of first gill arch, 8–17 small serrations around angle of preopercle edge, light organ on caudal peduncle 1.5–2.5 in caudal peduncle length, yellowish green stripe along dorsal edge of light organ and greenish eye.

Colour in life: Body pale, semi-transparent, with about 7 clusters of dark dots in irregular vertical orientation; anterior ones may appear like shallow, forward pointing chevron marks and tend to partially merge, last two reduced to small groups of dots mostly below midline of body ( Figs. 1 a, b). Predorsal area with minute dark dots extending posteriorly below first dorsal-fin base and fading out below second dorsal-fin base. Head with variable amount of black dots on snout, interorbital space, below eye, and gill cover; dots on cheek and opercle become larger ventrally, making lower cheek blackish. Head with several iridescent silver blotches of varying size and shape extending from just behind eye to cheek and opercle. Upper half to two-thirds of eye yellowish green. Darkdotted areas on body and head usually tinged with yellowish green, less noticeable on caudal peduncle. Light organ with dark striation, its dorsal edge with yellowish-green stripe speckled with diffuse dusky spots of various sizes and a narrow, fuzzy, bright yellow area above anal-fin base. Small, faint orange dots scattered on body and upper part of head; faint orange dot at base of dorsal-fins’ spines and rays. Fins usually pale, but first dorsal fin spines sometimes with yellowish-green tinge and pelvic fins with dark dots at base.

Colour in alcohol: Similar to colour in life, but body pale brown with no yellowish green tinge; lower part of iris with varying amount of dark brown pigment over silvery background; upper part of iris mostly blackish, except narrowly dark brown around pupil.

Smallest specimen, a 14.6 mm female from Fakfak Peninsula, and largest specimen, a 30 mm female from Fiabacet Island, SE Misool, both in West Papua Province, Indonesia.

Description. Proportional measurements of the holotype and selected paratypes are given in Table 1 . Data in parentheses refer to paratypes. Dorsal-fin rays VII + I,9 (VI–VII); anal-fin rays II,8 (8–9); last soft ray of dorsal and anal fins branched to base; pectoral-fin rays 13 (12–13), two upper and lowermost rays unbranched; pelvic-fin rays I,5, all soft rays branched; principal caudal-fin rays 17, upper and lowermost rays unbranched; upper and lower procurrent caudal rays 7 (6–7); tubed lateral-line scales 9 (2–9); vertical scale rows 23–24; median predorsal scales 5 (4–5); circumpeduncular scales 12; developed gill rakers 1 + 8 (1 + 8–9); gill rakers on ceratobranchial 7; branchiostegal rays 7; supraneurals 2. Body deep and compressed, its depth 2.6 (2.3–2.9) in SL and its width 2.2 (1.7–2.2) in depth; head length 2.3 (2.0–2.4) in SL; snout length 5.6 (4.6–6.8), eye diameter 3.7 (2.8–3.8), and interorbital space, slightly convex, 5.2 (3.85–5.7), all in head length.

Mouth terminal, oblique and large; maxilla reaching posteriorly to vertical at posterior edge of pupil (to middistance between rear edges of pupil and eye in several paratypes, and to rear edge of eye in one paratype); upper jaw length 1.95 (1.8–2.0) and lower jaw 1.8 (1.6–1.8) in head length; both jaws with band of small, conical teeth; upper jaw band with 2 series at symphysis, increasing to about 5 series at middle of jaw; lower jaw band with 4 series at symphysis tapering to 1 series posteriorly; inner series teeth of lower jaw somewhat enlarged; vomer and palatines with 1 irregular series of small conical teeth (see Remarks for variation).

Rear nostril oval, close in front of eye, its vertical diameter about 4 times in pupil diameter; anterior nostril round with slightly raised posterior rim, its diameter about half longer axis of rear nostril, and closer to rear nostril than to tip of snout (excluding upper lip) (mostly at about midway between tip of snout and rear nostril in most paratypes).

Preopercular edge armed with 16 (5–17) very small serrations around its angle and the adjacent part of ventral edge; preopercular ridge smooth; exposed edge of posttemporal smooth or with 4 (1–5) tiny serrations.

Body scales large, thin and easily shed; scales spinoid, those along upper and lower margins of body with few minute or no spines; tubed lateral-line scales followed by notched ones; caudal fin with low scaly sheath of 1–2 scale series across its base; other fins without scales or scaly sheath.

Dorsal-fin origin over third lateral-line scale and slightly behind vertical through upper pectoral-fin base; first dorsal spine, when present, usually very small, (4.0–9.5) in second spine; second dorsal spine 5.2 (4.7–6.7) and third dorsal spine longest 3.0 (2.7–4.0) in head length; spine of second dorsal fin 4.1 (3.25–4.15) and longest dorsal soft ray 2.4 (2.05–2.6) in head length; first anal spine 3.7 (2.7–3.55) in second spine, second anal spine 5.0 (4.4–5.7) and longest anal ray 2.65 (2.7–3.9) in head length; pectoral and pelvic fins not reaching beyond anus (reaching over anal-fin origin in smallest paratype); pectoral fin length 4.9 (4.2–5.3) in SL; pelvic fin origin on vertical through upper pectoral-fin base (slightly in front in some paratypes); pelvic fin length 5.6 (4.6–5.6) in SL and pelvic spine length 1.4 (1.3–1.5) in fin’s length; caudal fin forked, 3.6 (3.1–4.0) in SL; length of light organ on caudal peduncle 1.6 (1.5–2.5) in caudal peduncle length and 8.8 (7.6–12.7) in SL.

Dorsal profile of head straight (slightly convex in some paratypes), predorsal distance 2.5 (2.25–2.5), preanal distance 1.5 (1.4–1.6) and prepelvic distance 2.4 (2.2–2.5) in SL; caudal peduncle depth 1.7 (1.5–2.0) in its length, and the length 1.8 (1.15–1.7) in distance between pelvic-fin base and anal-fin origin; latter distance 3.0 (3.25–4.3) in SL.

Colour of holotype in alcohol: Body pale brown with 7–8 irregular darker brown bars along middle of body; bars less discreet anteriorly, becoming shorter and separated by distinct pale areas posteriorly; last bar reduced to 7–8 dark dots in two series. Upper part of body, from head to below middle of second dorsal-fin base with small dark brown dots, but area immediately below dorsal-fin bases with fewer dots more or less outlining scale pockets; predorsal scales outlined by similar dots. Head below level of middle of eye dark brown, except snout and silvery area on lower cheek, behind maxilla. Jaws with dark brown dots; roof of mouth cavity bordered by vomer and palatines with small black dots extending posteriorly to upper gill chamber. Pelvic-fin spine with single series of small dark dots and pelvic-fin rays with few dark dots proximally; other fins pale, but pectoral-fin base dark brown. Abdominal area of body with dark shade from underlying dark peritoneum. Blackish stripe along upper edge of light organ, tapering posteriorly. Peritoneum with fairly dense, conspicuous blackish dots; stomach and intestine with smaller dark dots; intestine with pale area in middle of its posterior, straight part.

Distribution. Siphamia papuensis is known from the Fiabacet group of small Islets off SE Misool in the Raja Ampat Islands, the Fakfak Peninsula, and Triton Bay, all in Indonesia’s West Papua Province; it has been recorded from depths of 50– 72 m.

Etymology. This species is named papuensis in recognition of its type locality.

Remarks. Siphamia papuensis is unusual among its congeners, and indeed in the Apogonidae , in having variation in the number of first dorsal-fin spines. Of our 36 specimens 11 had seven spines and 25 six spines, a ratio of 1:2.27, respectively. In some of the seven-spined specimens the first spine is very small and can easily be overlooked. There was no relationship between the number of spines and sex, size, depth or locality. In several paratypes the tubed lateral-line scales had a vertical line of minute papillae along the middle of the scale at least above the tube. Two specimens had two developed gill rakers on the upper limb of the first gill arch of the left side, but only one developed gill raker on the right side. The largest females had large gonads with ripe eggs which extended their abdominal region. Consequently, these females had a larger distance from anal-fin origin to anus than observed in males ( Table 1 ). Four specimens, all males, had an unusually short section of the light organ on the caudal peduncle relative to their size. Two of these had parasitic nematodes in their abdominal cavity which may have stunted the fish growth. The other two ( Fig. 1 e, diamond outliers) had no nematode in the abdominal cavity and no evidence of external parasites, and could be a genetic abnormality. Another male had the light organ stunted only on the left side of the caudal peduncle. The light organ develops faster in males, and females of this species are larger than males ( Fig. 1 e). The largest male measured 23.5 mm while the largest female was 30.0 mm. The absence of smaller females is a sampling artefact because only formalin fixed specimens are shown in the graph. Other specimens were fixed in 95% ethanol for genetic work and through random selection smaller females ended up in this group. The iridescent silvery spots and blotches on the head behind the eye are probably due to guanine deposits. In situ observations of numerous specimens by M. Erdmann showed these to be a prominent life colour feature of this rather plain-looking species ( Figs. 1 a–d).

Siphamia papuensis is found in loose aggregations of 10–50 individuals around ledges and crevices on steep, current-swept slopes in depths of 50–72 m (usually below 60 m), and has frequently been observed associating with small gorgonians and crinoids ( Figs. 1 c, d). A number of individuals were observed mouthbrooding around the full moon period.

The similarity in colour pattern between Siphamia papuensis , S. argentea and S. tubulata (see Comparisons below) could be misleading. Siphamia argentea , originally described from the Philippines, is also known from northwestern Australia and, like S. papuensis , is a deeper water species found at 23–107 m ( Gon & Allen, 2012). During their research for Reef Fishes of the East Indies G. Allen and M. Erdmann collected and photographed specimens of S. argentea in Lembeh Strait ( Figs. 1 f–h) and Bali ( Fig. 2 a), Indonesia, and Milne Bay, Papua New Guinea ( Fig. 2 b). Richard Pyle (BPBM) collected a specimen in Vanuatu ( Fig. 2 c).

Hayashi et al. (1994) collected and photographed Siphamia tubulata in Kashiwa-jima Island, Kochi Prefecture, at a depth range of 18– 44 m. Yoshida et al. (2010) reported it from Yaku-shima Island of the more southern Kagoshima Prefecture on the basis of an underwater photo taken at 25 m and also shown here ( Fig. 2 d). Hayashi et al. (1994) counted 9 developed gill rakers and 13 pectoral-fin rays in their specimens and both counts agree more with those of S. argentea and S. papuensis ( Tables 3, 5) than with S. tubulata (usually 8 and 12, respectively, in the latter; Gon & Allen 2012). Furthermore, a striated light organ is evident on the fish from Yaku-shima Island when the photo of Yoshida et al. (2010) is enlarged, excluding it from the S. tubulata group which has a dark-dotted light organ. The photos in both Japanese papers show an enhanced pattern of irregular bars on the side of the body; distinct dark brown marks on the top of the head, anterior end of first dorsal-fin base, origin and end of second dorsal-fin base, and at lower and upper caudal-fin base; and red spots ventrally along the light organ, and along the back between the dorsal fins and the lateral line. Since these colour features agree with the photos of our specimens of S. argentea from Indonesia, Papua New Guinea and Vanuatu (compare Fig. 2 d with Figs. 1 f–h and Figs 2 a–c) rather than those of S. papuensis ( Figs. 1 a-d), we conclude that the Japanese fish are S. argentea .

While diving in Bali for S. argentea, G. Allen and M. Erdmann photographed a small group of S. stenotes ( Figs. 2 e–g), a species previously known only from the Triton Bay area of West Papua Province in Indonesia. Note the conspicuous orange spot on the chin of the fish. It is not visible in the underwater photo of this species, but present though indistinct in the photo of a fresh specimen in Gon and Allen (2012: figs 28f, g).

The final 16S rDNA alignment was 536 nucleotides in length and is available from TreeBASE (accession number S16805 ). The phylogram (-lnL = 2449.145) from the maximum-likelihood analysis is presented in Figure 3 . This topology was largely identical to the strict consensus of the two equally-parsimonious trees (293 steps, consistency index = 0.587, retention index = 0.698, rescaled consistency index = 0.410), as well as the neighbourjoining tree, with respect to major clades recovered. All topologies retrieved S. papuensis as a distinct, monophyletic clade, clearly separate to that of S. argentea . The former was well-supported (bootstraps ≥ 85%), while the latter was only supported (99% bootstrap) in the neighbour-joining analysis. In turn, the S. papuensis + S. argentea sister-species clade was distinguished from a monophyletic clade containing the remaining species ( S. jebbi , S. stenotes and S. tubifer ), but these clades were not well supported across all analyses. Although the mean uncorrected 16S rDNA sequence divergence between S. papuensis and S. argentea (8.1%; Table 6 ) was less than that observed in all other comparisons among the Siphamia species included (12.6–16.4%), this was greater than the mean sequence divergences observed within each of these species ( S. argentea : 4.2%; S. papuensis : 3.7%). There was also no overlap between the ranges of values observed within each of these species (0–6.9%) and among them (7.6–9.4%). While the latter values were comparable to the intraspecific divergences observed within S. tubifer (5.4–10.0%), the included S. tubifer individuals were from vastly separated localities and these divergences may reflect population level differentiation. Collectively, the genetic evidence substantiates the separation of S. papuensis and S. argentea .

Comparisons. See Tables 3–5 for frequency distributions of pectoral rays, lateral-line scales and gill rakers in species of the Siphamia tubifer group with 13 pectoral-fin rays. The striated light organ of S. papuensis places it in the Siphamia tubifer group of species. Within this group, the count of 13 pectoral-fin rays separates S. papuensis from all other species, except S. argentea , S. jebbi and S. stenotes which share the same count ( Gon & Allen, 2012). The latter two species are easily distinguished by having two developed gill rakers on the upper limb of the first gill arch ( Table 3) and a single supraneural. They also have different colour patterns, with S. stenotes exhibiting two dark stripes along the body ( Figs. 2 e-g) and S. jebbi having no dark stripes or bars, but series of reddish orange spots and dashes on its body ( Fig. 2 h). Siphamia argentea differs in having a complete lateral line and although it may occasionally have a dull, yellowish green appearance (e.g. Fig. 1 g) similar to S. papuensis , its colour pattern is usually markedly different. The markings on the side of the body of S. argentea are in a reticulated pattern rather than barred, more brownish ( Figs. 1 h, 2d) and sometime reddish ( Figs. 1 f, 2b), and stand out against a whitish background. Furthermore, S. argentea has orange dots or short, vertical bars along the light organ, and reddish brown dots on the upper part of the body and head; it also has blackish marks on nape, the origin and usually end of both dorsal fins and sometimes anal fin, and at upper and lower caudal-fin base ( Figs. 1 f-h and 2a-d); these marks are absent in S. papuensis ( Fig. 1 a-d). S. argentea seems to be a larger species reaching over 40 mm while the largest S. papuensis measured 30 mm. The pattern of irregular bars and associated greenish colour or the greenish to reddish brown reticulated pattern of S. papunesis and S. argentea , respectively, are also present in S. tubulata , S. corallicola and S. elongata , but these three species have a dotted light organ, 11–12 pectoral-fin rays and usually 6 gill rakers on the ceratobranchial ( Gon & Allen, 2012).

1 2 7 8 9 10 n S. papuensis 34 2 28 8 36 S. argentea 27 1 1 18 9 28 S . jebbi 35 5 23 7 35 S. stenotes 30 1 29 30 0 1 2 3 4 5 6 7 8 9 10 11 23 24 n S. papuensis 1 1 3 5 6 3 2 3 24 S . argentea 3 10 13 S . jebbi 11 13 5 1 30 S. stenotes 2 6 8 16 Species Number of rays

12 13 14 n S. papuensis 1 28 29 S . argentea 1 22 23 S . jebbi 3 31 1 35 S. stenotes 1 23 6 30

Though S. argentea in particular shares similar habitat preferences to S. papuensis (associating with crinoids or small sea fans along deep current-swept reef slopes in the 50–70 m depth range), our in situ observations of the former species in Lembeh Strait, North Bali and Milne Bay suggest that it is less gregarious than S. papuensis , normally occurring in smaller groups of only 2–8 individuals. Siphamia tubulata has been observed on sandy bottoms with green algae and seaweed at a shallower depth range of 14–32 m ( Gon & Allen, 2012).

Figures

FIGURE 1. (a – b) Siphamia papuensis, holotype, Pulau Balbal, Indonesia, 66 m (M. Erdmann). (c) Siphamia papuensis, Fiabacet, Indonesia, 60 m (G. Allen). (d) Siphamia papuensis, Fiabacet, Indonesia, 70 m (M. Erdmann). (e) Light organ length in males and females of Siphamia papuensis (polynomial fitting). Outlying individuals (diamonds) are males with a very short light organ (see Remarks for details). (f – h) Siphamia argentea, Lembeh Strait, Indonesia, 70 m (M. Erdmann & G. Allen).

FIGURE 2. (a) Siphamia argentea, North Bali, Indonesia, 70 m (G. Allen). (b) Siphamia argentea, Milne Bay, Papua New Guinea, 50 m (M. Erdmann). (c) Siphamia argentea, Vanuatu (R. Pyle). (d) Siphamia argentea, Yaku-shima Island, Japan, 25 m (S. Harazaki). (e – g) Siphamia stenotes, NW Bali, Indonesia (G. Allen). (h) Siphamia jebbi, Cenderawasih Bay, W Papua Province, Indonesia, 14 m (G. Allen).

FIGURE 3. Maximum likelihood phylogram (- lnL = 2449.145), based on the analysis of a 16 S rDNA sequence fragment, depicting relationships among Siphamia papuensis, S. argentea, S. jebbi, S. tubifer and S. stenotes. Numbers on the branches indicate bootstrap support for the nodes from the likelihood, parsimony and neighbour-joining analyses (in the order parsimony / likelihood / neighbour-joining). The tree was rooted using Apogonichthyoides pharaonis and Ostorhinchus lateralis as outgroups. Broad collection localities are indicated to the right of the terminals. Asterices indicate those sequences that have been designated as paragenetypes of S. papuensis.

Tables

TABLE 1. Proportional measurements of selected type specimens of Siphamia papuensis expressed as percentages of the standard length. Absent first dorsal spine refers to specimens with six spines in the first dorsal fin.

Holotype	Paratype	Paratype	Paratype	Paratype	Paratype
MZB	WAM	WAM	WAM	WAM	NSMT-P
22100	P.33387-001	P.33386-001	P.33387-001	P.33387-001	118258
Standard length (mm) 27.8	18.45	20.2	22.3	24.5	26.45
Head length (mm) 12.0	8.5	8.9	9.75	10.4	11.5
Body depth 38.1	39.0	38.4	37.7	34.1	38.6
Body width 17.1	17.9	19.1	21.1	19.6	20.0
Head length 43.2	46.1	44.1	43.7	42.45	43.5
Snout length 7.7	6.8	7.4	9.2	7.8	9.3
Eye diameter 11.7	13.55	12.6	12.8	12.65	13.2
Interorbital width 8.3	8.1	9.9	8.3	7.8	10.2
Upper jaw length 22.1	23.6	22.3	21.5	20.8	23.4
Lower jaw length 24.3	26.0	25.7	26.5	24.1	27.6
Maxillary width 5.2	4.6	4.95	6.3	6.3	6.8
First dorsal spine Miniscule	Absent	Absent	0.9	Absent	Absent
Second dorsal spine 8.3	7.6	7.2	6.5	8.4	Damaged
Third dorsal spine 14.2	14.4	14.4	13.45	13.5	14.7
Spine of second dorsal fin 10.6	11.65	Damaged	10.5	10.6	11.3
Longest soft dorsal ray 18.2	18.4	18.3	16.6	17.1	19.3
First anal spine 2.3	3.25	3.0	2.7	3.1	2.65
Second anal spine 8.6	8.9	7.9	7.6	8.6	9.1
Longest soft anal ray 16.2	11.9	14.4	Damaged	13.5	14.9
Pectoral fin 20.3	21.1	21.8	20.4	18.8	22.7
Pelvic fin 18.0	20.3	18.8	17.9	18.0	20.2
Pelvic fin spine 12.95	14.1	14.1	12.3	12.9	13.2
Caudal fin 27.5	30.6	28.5	26.5	26.1	26.5
Length of caudal peduncle 18.2	19.5	20.3	17.9	16.9	22.7
Depth of caudal peduncle 10.8	11.65	9.9	10.5	9.0	12.5
Length of light organ on caudal 10.1	7.9	8.2	11.2	10.2	12.3
peduncle
Anus to anal-fin origin 6.1	0.5	1.5	1.1	4.1	3.4
Pelvic fin to anal fin origin 33.1	24.7	23.3	23.5	29.4	29.7
Predorsal distance 40.65	44.4	42.1	43.05	40.4	43.1
Preanal distance 68.5	64.0	65.6	66.6	68.2	67.5
Prepelvic distance 41.0	40.1	41.8	44.6	40.2	42.3
Sex Female	Male	Male	Male	Female	Female