Heteroclinus whitleyi, Hoese & Hay & Dibattista, 2024

Hoese, Douglass F., Hay, Amanda & Dibattista, Joseph D., 2024, A review of the Heteroclinus heptaeolus complex (Pisces: Blennioidei: Clinidae), with three new species and discussion of use of proportions in taxonomic studies, Zootaxa 5432 (3), pp. 301-348 : 326-333

Heteroclinus whitleyi , n. sp.

Plates 5 , 6 , Figures 2 , 3 , 10 , 11 , Tables 1–5 , 9 , 11–12

Common Name: Whitley’s Weedfish

Cristiceps wilsoni Lucas, 1891: 10 , fig (in part).

Petraites heptaeolus .— Milward 1967: 5 (description and figure of Western Australian specimens).

Heteroclinus sp. 2 .— Rennis, Hoese & Gomon 1994: 747 (New South Wales and South Australia to Recherche Archipelago, Western Australia); Hoese, Gomon & Rennis 2008: 702 (New South Wales and South Australia to Recherche Archipelago, Western Australia).

Heteroclinus sp. 4 .— Kuiter 1993: 331 (habitat information).

Heteroclinus sp. 3 .— Kuiter & Kuiter 2018: 289 (New South Wales to Western Australia).

Holotype. AMS I.17019-023, 52.9 mm SL female, Long Reef, Sydney , New South Wales, 33°45’S, 151°19’E, 3 m, D. Hoese & party, 27 March 1973.

Paratypes. All from New South Wales : AMS I.16850-022, 1(42), Jervis Bay , 35°08’S, 150°45’E, 9 m, D. Pollard & P. Straw, 30 October 1971 ; AMS I.17019-022, 1(54), Long Reef, Sydney , 33°45’S, 151°19’E, 3 m, D. Hoese & party, 27 March 1973 ; AMS I.15330-036, 1(72), Chinamans Beach, Jervis Bay , 35°01’S, 150°46’E, 3 m, M. Cameron & J. Paxton, 3 January 1969 ; AMS I.16237-030, 6(35–71), Long Reef, Sydney 33°45’S, 151°19’E, 3 m, D. Hoese, B. Congleton & G. McPherson, 29 February 1972 ; AMS I.16467-024, 2(61–64), Minnie Waters , 29°47’S, 153°18’E, 1 m, F. Talbot, 14 February 1965 ; AMS I.16468-018, 5(42–69), Minnie Waters , 29°47’S, 153°18’E, 1 m, F. Talbot, 15 February 1965 ; AMS I.16849-017, 1(45), Jervis Bay , 35°08’S, 150°45’E, 9 m, D. Pollard & P. Straw, 30 October 1971 ; AMS I.16863-014, 1(75), Southwest Bowen Is., Jervis Bay , 35°07’S, 150°46’E, 2 m, D. Pollard & P. Straw, 2-3 May 1973 ; AMS I.18064-012, 1(64), Broken Head, Byron Bay area , 28°43’S, 153°37’E, 0–2 m, D. Hoese, H. Larson & I. Briggs, 2 July 1974 ; AMS I.18336-001 , 2(63–68), Bondi, Sydney , 29°39’S, 15032’E , R. Kuiter , April 1974 ; AMS I.19358-003, 1(58), Clovelly Pool, Sydney , 33°55’S, 15°16’E, 9 m , R. Kuiter , 8 December 1976 ; AMS I.19671-001, 2(44–69), Thompson Bay, Clovelly, Sydney , 33°55’S, 15°16’E, 9 m , R. Kuiter , 28 January 1978 ; AMS I.19758-004, 5(46–60), Long Bay, Sydney , 33°58’S, 151°15’E, 1 m , R. Kuiter , 26 February 1977 ; AMS I.20833-018, 2(55–60), Arrawarra headland, 30°04’S, 153°12’E, 1 m, D. Pollard, 4 May 1977 ; AMS I.24174-001, 1(70), Bondi, Sydney , 29°39’S, 150°32’E, 1 m , R. Kuiter , 25 December 1976 ; AMS IB.7325, 2(63–69), Quarantine Beach, Sydney Harbour , 33°55’S, 151°17’E, 9 m, F. Talbot, J. Yaldwyn & D. Francois, 11 January 1965 ; NMV A.3259, 5(43–60), Arrawarra headland, 30°04’S, 153°12’E, 1 m, J. Paxton and party, 4 May 1977 ; USNM 216337 , 7 (57–75), Nielsen Park, Sydney Harbour , 33°51’S, 151°16’E, 0–3 m, B. Collette .

Non-type material. New South Wales: AMS I.9265, 2(43–71), Freshwater, Sydney, 33°48’S, 151°17’E ; AMS I.15899-005, 2(15–27), Minnie Water, 29°50’S, 153°18’E, 1 m ; AMS I.1771, 1(79), Sydney Harbour, 33°51’S, 151°16’E ; AMS I.17746-001, 1(46), North Bondi Beach , 33°54’S, 151°17’E, 3 m ; AMS I.19901-026, 1(65), Bottle and Glass Rocks, Nielsen Park, 33°50’51”S, 151°16’15”E, 5 m ; AMS I.19930-002, 2(70–75), Bottle and Glass Rocks, Nielsen Park, 33°50’51”S, 151°16’15”E, 3–5 m ; AMS I.19942-005, 2(60–68), The Basin, Hawkesbury River , 33°38’S, 151°15’E, 2 m ; AMS I.20095-057, 6(54–67), Arrawarra Headland, 30°04’S, 153°12’E, 1 m ; AMS I.20095-059, 17(42–62), Arrawarra Headland, 30°04’S, 153°12’E, 1 m ; AMS I.24175-001, 1(56), near Quarantine Beach, Sydney Harbour, 33°49’S, 161°17’E , AMS I.24491-001, 1(52), Camp Cove, Sydney, 33°50’17” S, 151°16’44” E, 2 m , 31 October 1976; AMS I.31253-043, 2(68–75), Camden Head, 31°38.5’S, 152°50.4’E, 4 m ; AMS I.41831-058, 1(67.5), Iron Peg Point, S of Lennox Head near Boulder Beach, 28°49’21”S, 153°36’24”E ; AMS I.44125-036, 1(67), North Side of Lion Is., Broken Bay , 33°33’19”S, 151°19’01”E ; AMS I.45025-047, 1(34), Collers Beach, Mollymook, 35°20’46”S, 150°28’36”E ; AMS I.45027-034, 2(26-41.5), 35°19’19”S, 150°29’E , Mollymook; AMS I.45633-067 18 (48–71), Washerwomans Beach, Bendalong, 35°14’39”S, 150°32’09”E . South Australia: SAM F.2583, 1(60), no data ; AMS I.17608-006, 18(32–60), Moana, 35°12’S, 138°28’E, 1 m ; AMS I.17614-008 , 14(48–58) and SAM F7694 , 10 (46–56), Tinckera and Point Riley, Spencer Gulf , 33°49’ S, 137°40’E, 1 m. Victoria: NMV A.2531, 1(52), Leonard Bay , cove on southern shore, Leonard Pt., 39°01’S, 146°17’E . Western Australia: WAM P.7226, 1(70), Cape Vlaming, 32°01’S, 115°27’E ; AMS I.19602-025, 2(61–67), 33°37’S, 115°18’E, 0–4 m ; AMS I.19629-030, 1(65), Dunsburg, Geographe Bay . 33°37’S, 115°18’E, 1 m ; AMS I.20216- 009, 4(18–36), Rob Is., Recherche Archipelago, 34°02’S, 122°14’E, 7 m ; AMS I.20219-022, 2(60–66), Rob Is., Recherche Archipelago, 34°02’S, 122°14’E, 7 m ; AMS I.24331-002, 3(50–58), Gannet Rock, Meelup, 33°34’S, 115°04’E, 2 m , AMS IB.5151, 1(41), Rottnest Is., 32°00’S, 115°30’E ; USNM 216338 , 2 (59–71), 30 miles south of Perth , 32°16’S, 115°41’E , 1970; WAM P.6203-001, 1(61), off Yanchep, 31°33’S, 115°25’E ; WAM P.25758-019, 1(40), Fish Hook Bay , Rottnest Is., 32°00’S, 115°30’E, 8 m ; WAM P.25765-010, 2(17–31), Long Is., Recherche Archipelago, 34°03’S, 121°58’E, 8 m ; WAM P.25783-015, 1(30), Rottnest Is., 32°01’S, 115°30’E, 8–10 m ; WAM P.25798-007, 2(58–59), Eagle Bay, Geographe Bay , 33°33’S, 115°04’E, 0–4 m ; WAM P.26007-005, 2(24–42), Rob Is., Recherche Archipelago, 34°20’S, 122°14’E, 10 m ; WAM P.26143-001, 1(25), Dongara, 29°04’S, 114°45’E, 15 m ; WAM P.26541-010, 1(60), Fisherman Is. 30°08’S, 114°56’E, 1–3 m ; WAM P.27273-001, 1(64), outer Geraldton Harbour, 28°46’S, 114°37’E, 0–15 m ; WAM P.27283-006, 2(51–59); Breton Bay , 31°14’S, 115°25’E, 0.5–1.5 m ; WAM P.27644.016 (photo only), Gannet Rock, Meelup, 33°34’S, 115°04’E, 2– 4 m.

Diagnosis. Dorsal fins III, XXIII–XXV, 2–3 (usually 3); anal rays II, 15–18 (usually 17); pectoral rays 11–12 (usually 12); gill rakers on outer face of first arch 2–4 + 6–9 = 8–12 (usually 10–11); circumorbital head pores uniserial (12–16 pores); orbital tentacle low, broad, with rounded distal margin about 1–1/2 to 2 times as long as wide; nasal tentacle short, rounded with an expanded tip; middle gill rakers and uppermost rakers on outer face of first arch not branched dorsally; first dorsal fin slightly elevated (second dorsal spine 8.0–11.9% SL, decreasing slightly with increasing growth) originating over middle of preoperculum; third dorsal spine usually slightly in front of a vertical from pelvic origin; last dorsal ray connected by membrane to posterior part of caudal peduncle to upper base of caudal fin; body moderate to deep, proportion increasing significantly with size (depth at anal origin 24.5–31.1% SL in specimens 25.8–51 mm SL, 26.5–32.7% SL in specimens 51–77.7 mm SL). Body brown to reddish brown or purple, often with 3–4 distinct dark, longitudinal bands or large spots forming horizontal rows; a silver or white bar extending horizontally from posterior margin of eye at level of lower pupil margin; dark brown bars below base of dorsal fins broader than light spaces between bars; interspinal membranes of second dorsal fin with no pigment forming a series of clear windows, rarely without clear windows or with windows extending over two spines. Anal fin with similar clear windows between rays.

Description. Based on more than 180 specimens, with data taken for 89 specimens 25–79 mm SL, data taken on 36 males, 45 females, 8 not sexed. First dorsal III* (90); dorsal rays 3*(90); pelvic rays I,3* (80); segmented caudal rays 9(6), 10*(74); vertebrae 13+25(3), 13+26(6), 13+27(2), 14+25*(20), 14+26(21), total 38(2), 39*(25), 40(23); lower gill rakers on outer face of first arch 5(1), 6(10), 7*(42), 8(29), 9(5); circumorbital head pores uniserial (12–16 pores); pored lateral line scales 17–27 (arched portion of line) +19–29 (straight portion of line), anterior lateral line scales 17(1), 18(1), 19(3), 20(4), 21*(16), 22(16), 23(25), 24(15), 25(6), 26(1), 27(1) 19(1), 20(1), 21(2), 22(5), 23(8), 24(8), 25(22), 26*(24), 27(11), 28(3), 29(1); total lateral line scales 38(2), 39(1), 40(1), 41(2), 43(1), 44(9), 45(4), 46(16), 47*(13), 48(11), 49(16), 50(4), 51(4), 52(2), 53(1); branchiostegal rays 6*(30); other counts are shown in Tables 1–5 . Vomer with conical teeth an inverted V, 1 row in specimens less than 45 mm SL and 1–2 rows in larger specimens; palatine without teeth.

Head and body laterally compressed; head moderate length 23.4–34.5% SL, proportion decreasing significantly with increasing size 28.4–31.4 % SL in specimens 25.8–39.5 mm SL, 23.7–29.6% SL, in specimens 40–77.7 mm SL; snout gently curved in side view, snout shorter than eye diameter (0.4–7 of eye length), (2.4–4.8% SL), decreasing slightly with increased size; eye moderate, relative size decreasing substantially with growth (8.5–9.3% SL in specimens <40 mm SL, 6.8–8.4% SL in specimens 44–65 mm SL, 5.8–7.5% SL in specimens 66–78 mm SL; interorbital narrow, about one-half to two thirds eye diameter; mouth short, jaws reaching to below middle or just before middle of eye, upper jaw length, 6.4–9.0 % SL in specimens less than 60 mm SL, decreasing with increasing size to 6.4–7.5% SL in specimens 61–77 mm SL; anterior nostril at end of short tube, about 3–5 nostril diameters above upper lip and 3–5 nostril diameter before middle of eye, with short spatulate nasal tentacle expanding slightly distally, with rounded margin; posterior nostril with elevated rim above anterior margin of eye; gill rakers on outer face of first arch, short and slender, much shorter than filaments; rakers on inner face of first arch much shorter, than rakers on outer face of first arch and pointed, rakers becoming progressively shorter on following arches; tongue tip pointed; upper jaw with 4–5 rows of incisiform (slightly compressed) to conical teeth, teeth in outer row largest with pointed tips, without basal lateral cusps, teeth becoming more conical, smaller and pointed in inner rows, rows teeth tapering to one to two rows near end of jaw; teeth in lower jaw incisiform (slightly compressed) to conical without lateral cusps in 4–6 rows ventrally and one row posteriorly.

Genital valve small, composed with smooth to rugose distal margin, fully covering genital opening in females. Intromittent organ pointed, without lateral lobes, with small flap behind anterior tip.

Head pores as shown in Figure 10 ; circumorbital and preopercular pores uniserial.

Head largely naked; body scales small and cycloid extending forward to above operculum below second dorsal spine to just behind first dorsal fin origin; scales cycloid overlapping and forming distinct rows, becoming scattered and non-imbricate on caudal peduncle; pectoral fin base covered with small scales, scales extending onto base of fin to a maximum of basal one-fifth of rays; scales covering proximal bases of some dorsal spines in 1–2 rows, often also found on membranes between spines, not extending on rays of dorsal; scales not extending onto anal fin; scales not extending onto base of caudal rays; lateral line scales extending to caudal peduncle, anterior scales overlapping with a single median posterior pore, posterior scales separate with a median pore at each end.

All fin-rays unbranched; first dorsal fin origin, above posterior preopercular margin to midway between posterior preopercular margin and eye; fin elevated, with second spine longest, fin higher than second dorsal fin; second spine usually longest, with first and third spines subequal in height; membrane from first dorsal fin connects to back or just above base of second dorsal fin; second dorsal origin above a point just before upper pectoral origin and well behind pelvic insertion; first spine of second dorsal slightly shorter than following spine, spines becoming progressively longer posteriorly, with last spine the longest; last two dorsal rays shorter than first ray and widely separated from anterior ray; anal origin below 9 th to 11 th spine of second dorsal fin, two anal spines slightly shorter than rays; posterior rays becoming progressively longer, last two rays shorter, slightly closer spaced and widely separate from anterior rays; last anal ray connected along basal half of ray by membrane to or just before middle of caudal peduncle; pectoral fin with rounded posterior margin, central rays longest, reaching to above or just before anus, rarely reaching to above anal fin origin pelvic fins with hidden spine, 3 developed rays, pelvic rays stout, not reaching to anus, inner ray about 1/2 length of second ray; caudal fin with truncate to slightly rounded posterior margin, 18.0–22.5% SL in specimens less than 40 mm SL decreasing significantly to 15.1–20.3% SL in specimens over 65 mm SL; caudal rays unbranched, 10 thickened rays and an upper and a lower smaller ray, rarely segmented, 3–5 upper and lower very short procurrent simple rays difficult to discern.

Coloration of freshly collected material ( Plates 5–6 ). Colour variable. New South Wales ( Plate 5 ): body green, purple, or reddish brown with 7–8 dark saddles along back extending onto dorsal fin, but not extending ventrally to midside; saddles much broader than spaces between saddles; side of body with a series on horizontally elongate silver spots forming two distinct rows, spaces red to brown (darker than body colour); some specimens with large dark spots along midline; head uniformly reddish brown; a large white or silver oblong spot extending horizontally, posteriorly from behind eye; pectoral fin base with a white to light horizontal to crescent-shaped bar on upper half; first dorsal dark, second dorsal fin brown to reddish brown with dark patches extending upward from dorsal saddles; spaces between dark areas with transparent sections (3–8 clear windows in total between some spines, usually with windows separated); caudal fin brown to reddish brown with no pigment on membranes between caudal rays; anal fin with similar coloration to second dorsal fin, but with fewer clear windows and saddles ventrally on body fainter than dorsal saddles; pectoral fin with 2–3 broad dark brown saddles; pelvic fins uniformly brown to reddish brown. South Australia ( Plate 6A & B ): general coloration similar to specimens from New South Wales, differing in having clear windows broader, between 3 dorsal spines, with only one spine in middle of window, with paler pigment than dark areas; one specimen with uniform reddish brownish coloration and clear windows confined to portion of dorsal fin. Western Australia; Saddles dorsally on back connected by a dark brown stripe; saddles about equal to or slightly smaller than spaces between saddles; silver spots on side of body connected to form 4 stripes extending from head to posterior part of body; dorsal fin with 2–3 rows of spots, often interconnected to form oblique bands or stripes; clear windows normally in pairs, sometimes with a single spine in clear area; one female ( Plate 6 C & D ) without silver spots and sides of body with pupil to eye sized black spots scattered over body and short vertical bars on midside; other coloration as in New South Wales specimens.

Live Coloration. Similar to coloration of freshly collected material. Overall head and body green, purple, or reddish brown, with distinct narrow clear windows.

Coloration in alcohol. Head and body uniformly light brown, without distinct marking, dorsal membranes usually with clear or translucent windows, sometimes only slightly lighter than other interspinal membranes.

Distribution. The species is known from Minnie Water, New South Wales to Wilsons Promontory, Victoria and Gulf of St. Vincent to Spencer Gulf, South Australia, and Fisherman Island to Recherche Archipelago, Western Australia, but apparently absent from western Victoria. Associated with small brown algae to depths up to 10 m. Four colour phases were found, with three appearing to be linked to geographical area. However, too few photos are available to be sure the differences are related to specific populations.

Etymology. Named for Gilbert P. Whitley, formerly Curator of Fishes at the Australian Museum.

Remarks. In having the last 2 dorsal rays widely separated from the first ray, this species is similar to other species of the H. heptaeolus complex. This species differs from H. heptaeolus and H. wilsoni in having fewer spines in the second dorsal fin and fewer anal rays. It is superficially similar to H. heptaeolus in body form and orbital tentacle. However, it has expanded anterior neural spines and an expanded first pterygiophore, similar to those of H. wilsoni and H. colemani . The species is slightly deeper bodied that H. heptaeolus , with considerable overlap between the species. The first dorsal ray is also longer in H. whitleyi ( Figure 21 ), but again with some overlap. Although with different averages, there is some overlap between this species and H. heptaeolus in dorsal spine and anal ray counts. In addition, the coloration is different in live and freshly collected individuals. In H. whitleyi there are about 8 transparent windows between two dorsal spines on the second dorsal fin, resulting in separate distinct windows. In some specimens there may be transparent windows in two adjacent interspinal membranes, resulting in apparent single broad window ( Plate 6A and 6C ), particularly in specimens from South Australia and Western Australia. That coloration superficially resembles the coloration in H. heptaeolus . However, the transparent area is about equal to the dark band on the dorsal spines, with only one spine between in the windowed areas. In H. heptaeolus , the transparent areas is much broader than the dark bands on the dorsal fine, with 2 or 3 spines in the transparent window. There are similar differences in the windows on the anal fin. The body coloration is also distinct. In H. whitleyi there are no vertical bands on the body, but dark stripes or a series of elongated spots. In H. heptaeolus , there are a series of vertical bands on the body. The silver bar behind the eye also differs slightly, with the bar usually horizontal in H. whitleyi and slightly oblique in H. heptaeolus . In preserved material the body and fin coloration fades to a brown or yellowish-brown coloration in both species. However, the clear windows are usually discernible in preserved specimens of H. whitleyi , but not in preserved specimens of H. heptaeolus . Although there are only a few specimens with photographs of fresh specimens, no specimens with uniform body coloration have been found with the confirmed identity of H. whitleyi . Because of the overlap in meristic and morphometric features, identification of the Kalbarri specimen, WAM P.27959-017, is uncertain, other than it could represent H. heptaeolus , H whitleyi , a possible hybrid or a separate unrecognised species.

Regression analyses ( Table 9 ) indicated a y-intercept, significantly different from 0 for head length (p <<0.001), predorsal length (p = 0.005), body depth at anal origin (p <0.001), jaw length (p = 0.002), eye length (p <0.001), second dorsal spine length (p = 0.036), last dorsal spine length (p <0.001), first dorsal ray length (p = 0.002), caudal peduncle length (p = 0.038) and caudal fin length (p <0.001). Rank correlation indicated all characters decreased with size, except for body depth at anal origin which increased. Rank correlation tests for two additional characters showed a significant decrease with size: snout length (p = 0.033) and pectoral fin length (p = 0.043). Regression analysis indicated a significant difference at the p = 0.1 level for these two characters. All other proportions except body depth at anal origin decreased with increasing size. The body depth proportion increased with body size. Changes in the second dorsal spine length proportion was significant in rank correlation (p = 0.009), but the y-intercept was not significantly different from 0. The orbital tentacle proportion also showed significant decrease with increasing size, but regression analysis showed no significant correlation between the orbital tentacle length and standard length. It is likely that measuring error was large and the measurements very small.

Reaches a maximum length of 79 mm SL.

Figures

FIGURE 2. Radiograph of skulls of species of the Heteroclinus heptaeolus complex, showing modification and position of pterygiophores and first 3 dorsal spines on skulls of various species.

FIGURE 3. Radiograph of Heteroclinus whitleyi, n. sp., showing vertebrae and wide separation of the last two dorsal rays from the anterior ray and the less prominent separation of the last two anal rays.

PLATE 5. Freshly collected specimens of Heteroclinus whitleyi from New South Wales: A) holotype, AMS I.17019-022, 52.9 mm SL female from Long Reef, Sydney, photographed several days after capture; B) AMS I.20095-057, male from Arrawarra not measured; C) AMS I.45027-034, male, 41.5 mm SL, Mollymook, photo by M. McGrouther; D) AMS I.45025-047, male, 34mm SL, Mollymook, photo by M. McGrouther.

PLATE 6. Freshly collected specimens of Heteroclinus whitleyi from South Australia and Western Australia: A) AMS17608- 006, male from Moana, South Australia; B) AMS17608-006, female from Moana, South Australia; C) WAM P.25798-007, male, 58mm SL, Geographe Bay, Western Australia; D) WAM P.27644.016, male from Meelup, Western Australia, photo by B. Hutchins.

FIGURE 10. Head of Heteroclinus whitleyi, showing canals (dashed lines) and pores (open circles) and position of dorsal fins; anterior and posterior nostril at end of short tubes elevated above skin, based largely on holotype.

FIGURE 11. Map showing distribution of Heteroclinus whitleyi based on material examined.

FIGURE 21. First dorsal ray length in Heteroclinus whitleyi (black circles) and H. heptaeolus (green triangles).

Tables

TABLE 9. Regression and proportion parameters Heteroclinus whitleyi. P following y-intercept indicated the probability of y-intercept is significantly different from 0. Size trend indicates whether proportion decreases or increases with size (Tau and p for Tau). Size range is mm SL. Shaded probabilities indicate significant differences at or less than 0.05.

Character	n	Size range	Slope	95% confidence		y-intercept	p <>0	Predicted proportion range		Observed proportion range		Proportion trend	Tau	p for Tau
Head length	77	28.5–77.7	0.227	0.212	0.241	2.096	<<0.001	30.0%	25.3%	23.4%	34.5%	decrease	-0.336	<<0.001
Head width	68	28.5–72.5	0.146	0.120	0.171	0.050	0.947	14.7%	14.6%	11.3%	18.3%	ns	0.029	0.725
Predorsal	77	28.5–76.4	0.127	0.111	0.143	1.390	0.005	17.6%	14.5%	11.5%	18.8%	decrease	-0.246	0.001
Body depth anal origin	78	25.8–77.7	0.329	0.310	0.348	-2.198	<0.001	24.4%	30.0%	24.4%	32.7%	increase	0.284	<0.001
Caudal peduncle depth	77	25.8–77.7	0.046	0.041	0.050	0.206	0.126	5.4%	4.9%	4.1%	6.2%	ns	-0.136	0.080
Caudal peduncle length	75	25.8–77.7	0.098	0.087	0.110	0.724	0.038	12.6%	10.8%	9.1%	13.2%	ns	-0.196	0.013
Jaw length	77	25.8–77.7	0.064	0.058	0.070	0.596	0.002	8.7%	7.2%	6.4%	9.0%	decrease	-0.254	<0.001
Eye length	78	25.8–77.7	0.045	0.039	0.052	1.499	<0.001	10.4%	6.5%	5.8%	9.3%	decrease	-0.565	<<0.001
Snout length	77	25.8–77.7	0.029	0.022	0.036	0.370	0.069	4.3%	3.4%	2.4%	4.8%	decrease	-0.166	0.033
Pectoral fin length	75	25.8–77.7	0.145	0.129	0.161	0.842	0.080	17.8%	15.6%	13.7%	19.1%	decrease	-0.161	0.043
Pelvic fin length	73	25.8–77.7	0.160	0.141	0.179	0.015	0.978	16.0%	16.0%	12.6%	19.5%	ns	-0.022	0.782
Third pelvic ray length	75	25.8–77.7	0.066	0.053	0.080	0.321	0.429	7.9%	7.0%	4.4%	9.3%	ns	-0.109	0.167
Second dorsal spine length	77	25.8–77.7	0.087	0.076	0.098	0.714	0.036	11.5%	9.6%	8.0%	11.9%	decrease	-0.203	0.009
Last dorsal spine length	77	25.8–77.7	0.090	0.075	-0.104	1.899	<0.001	16.3%	11.4%	9.6%	15.6%	decrease	-0.374	<<0.001
First dorsal ray length	77	25.8–77.7	0.127	0.112	0.143	1.472	0.002	18.4%	14.6%	12.8%	18.3%	decrease	-0.288	<0.001
Caudal fin length	75	25.8–77.7	0.143	0.160	0.126	2.330	<<0.001	23.4%	17.3%	15.1%	22.5%	decrease	-0.363	<0.001