A review of the Heteroclinus heptaeolus complex (Pisces: Blennioidei: Clinidae), with three new species and discussion of use of proportions in taxonomic studies Author Hoese, Douglass F. 0000-0003-4380-5450 Australian Museum Research Institute, Australian Museum, 1 William Street, Sydney NSW 2010, Australia Author Hay, Amanda 0000-0002-1335-2342 Australian Museum Research Institute, Australian Museum, 1 William Street, Sydney NSW 2010, Australia Author Dibattista, Joseph D. 0000-0002-5696-7574 Australian Museum Research Institute, Australian Museum, 1 William Street, Sydney NSW 2010, Australia text Zootaxa 2024 2024-04-02 5432 3 301 348 http://dx.doi.org/10.11646/zootaxa.5432.3.1 journal article 292247 10.11646/zootaxa.5432.3.1 9f1bcd58-2ab4-4d2b-ab4b-7f3dd3e29405 1175-5326 10906573 CD1175FB-4CDA-4629-8E6F-75410C302915 Heteroclinus wilsoni ( Lucas, 1891 ) Plate 1C, 1D , Figures 2 , 12–13 , Tables 1–5 , 10–12 Common Name: Wilson’s Weedfish Cristiceps wilsoni Lucas, 1891: 10 , Port Phillip , Victoria . Heteroclinus wilsoni . — Kuiter 1993: 328 (habitat information); Rennis, Hoese & Gomon 1994: 764 (Eden, New South Wales to Kangaroo Island, South Australia ); Hoese, Gomon & Rennis 2008: 714 ( South Australia , Victoria and Tasmania ); Kuiter & Kuiter 2018: 285 ( New South Wales , Victoria and Tasmania ). Neotype : AMS I.20166-001, female 113 mm SL, Stokes Bay , Kangaroo Island , 35°38’S , 137°13’E , 5 March 1978 . Other Material Examined. New South Wales : AMS I.16970-009, 1(58), Boydtown , Nullica Bay , 37°06’S , 149°53’E , 13 March 1972 . Victoria : AMS I.16980-020, 2(50–88), Bells Beach , Surge Channel , 38°09’S , 144°25’E , 13 March 1972 ; AMS I.16984-009, 1(50), 8 km SW of Anglesea , 38°24’S , 144°13’E , 19 March 1972 ; AMS I.16988- 012, 1(55), Childrens Cove , 38°24’S , 142°28’E , 22 March 1972 ; AMS I.19774-004, 1(100), Portsea Pier , Port Phillip , Victoria , 38°19’S , 144°43’E , 9 April 1977 ; AMS I.24051-002, 2(39–41), Port Phillip Bay , 38°09’S , 144°52’E , 1 December 1981 ; NMV A.11357 , 1(68), southern Port Phillip Bay , 38°17– 38°16.8 , 144°37.7– 144°38.3’E ; NMV A.21791, 1(41), Torquay , Point Danger , 38°20’27”S , 144°19’39”E , January 1980 ; NMV A.2533, 3(34–39), Wilsons Promontory , 39°01.50’S , 146°17.5’E , 20 February 1982 ; NMV A.3012, 3(39–55), North shore of Horn Point , Wilsons Promontory , 39°02’S , 146°29’E , 9 February 1982 ; NMV A.7536, 1(70), Gabo Is. , 37°34’S , 149°55’E , 4 April 1989 ; NMV A.29392-001. 1(97), Lorne , 38°32’S , 143°58’E ; WAM P.27123-010, Norman Point , Wilsons Promontory , 38°56’s , 146°22’E , 25 February 1981 . Tasmania : AMS I.17545-007, 2(21–30), Eaglehawk Neck , 43°2’S , 147°56’E , 29 November 1972 ; AMS I.17551-004, 1(20), Rock Pier , Port Arthur , 43°9’S , 147°51’E , 2 December 1972 ; AMS I.17553-004, 1(120), Coles Bay , 42°8’S , 148°17’E , 4 December 1972 ; AMS I.17556-004, 1(98), Granville Harbour , 41°48’S , 145°00’E , 12 December 1972 ; AMS I.17586-002, 1(94), Boat Harbour , 40°56’S , 145°38’E , 10 December 1972 , AMS I.24227-002, 1(34), Eaglehawk Neck , Pirates Bay , 43°01’S , 147°56’E , 30 March 1970 ; AMS I.24287- 001, 1(47), Deal Is. , 39°29’S , 147°21’E , 8 May 1974 ; AMS I.24287-003, 1(100), Deal Is. , 39°29’S , 147°21 E , 8 May 1974 ; CSIRO T .88-02, 1(38), Sea Elephant Reef , King Is. , 39°48’S , 144°11’E , 24 February 1978 ; CSIRO T .1091, 2(99–108), Port Davey , ca. 43°18’S , 145°56’E , 30 January 1982 ; CSIRO T .1646, 1(82), Crayfish Point , Taroona , ca. 42°57’E , 147°21’21”E ; CSIRO T .90, 1(56), Ansons Bay , ca. 41°03’S , 148°18’E , 25 March 1979 ; QVM 1972 /5/[ex346], 1(44), Greens Beach , 41°04’55”S , 146°45’18”E ; USNM 214771 , 3 (75–101) Greens Beach , 41°04’55”S , 146°45’18”E ; USNM 216343 , 1 (101), Greens Beach , 41°04’55”S , 146°45’18”E ; WAM P.26569-014, 2(32–44), South Side of Rocky Cape , 40°51’S , 145°31’E , 9 March 1982 ; WAM P.27544-010, 8(27–46), Hope Is. , 43°40’S , 147°03’E , 6 February 1982 ; WAM P.27545-007, 2(19–30), Port Esperance , 43°20’S , 147°03’E , 8 February 1982 ; WAM P.27546-008, 2(25–29), 3 kms N of Esperance Point , 43°20’S , 147°03’E , 9 February 1982 ; WAM P.27547-003, 1(27), Port Esperance , Hope Is. , 43°20’S , 147°03’E , 10 February 1982 ; WAM P.27549-011, 2(32–147), Port Arthur , 43°09’S , 147°51’E , 14 February 1982 ; WAM P.27559-028, 6(33–57.5), St. Helens Point , 41°16’S , 148°22`E , 25 February 1982 ; WAM P.27560-013, 1(32), Bridport , 40°59’S , 147°23’E , 27 February 1982 ; WAM P.27562-014, 1(44), North of Waterhouse Point , 40°49’S , 147°41’E , 2 March 1982 ; WAM P.27577-012, 2(43–90), North of Greens Beach , 41°05’S , 146°45’E , 19 March 1982 ; WAM P.27580-024, 3(29–43), Low Head , 41°04’S , 146°48’E , 21 March 1982 . South Australia : AMS I.17612-005, 2(46–48), Rocky Point , Yorke Peninsula , 35°10’S , 137°41’E , 24 December 1973 ; AMS I.20166-020, 1(103), Stokes Bay , Kangaroo Is. , 35°38’S , 137°13’E , 5 March 1978 ; AMS I.20180-003, 1(104), Penneshaw , Kangaroo Is. , 35°44’S , 137°58’E , 9 March 1978 ; SAM F.1637, 1(117), specific location unknown, 25 November 1931 . Diagnosis. Dorsal fins III, XXV–XXVIII (rarely XXV), 3. A. II, 20–22 (usually 21); pectoral rays 12; gill rakers on outer face of first arch 2–3 + 7–9 = 9–12; circumorbital head pores uniserial (usually 16–l9 pores); orbital tentacle elongate, much longer than wide, with a round to emarginate margin, greater than 1/2 eye diameter in specimens over 50 mm ; nasal tentacle with an expanded tip, weakly to strongly bilobed; middle gill rakers and uppermost rakers on outer face of first arch not branched dorsally; first dorsal fin slightly elevated (second dorsal spine l0.5–l5.4% SL, not showing significant change with size), fin originating over middle to posterior portion of preoperculum, third spine well in front of a vertical from pelvic fin origin; last dorsal ray connected by membrane to posterior part of caudal peduncle; body deep with proportion increasing with increasing size (depth at anal origin 23.8–31.5% SL in specimens 31.5–50 mm SL, 28.3–33.1% SL in specimens 51–83 mm SL, and 28.8–36.4% SL in adults 81–146 mm SL), with proportion increasing with increased size. In life body of adults with a granulated pattern on body, often with minute white dots; a series of 6–7 dark vertical wavy bands on body; a distinct vertical band below the eye and a faint silver bar extending posteriorly from eye. Description. Based on 42 specimens , 21 males and 10 females , 11 undetermined, 20.7–146.5 mm SL. First dorsal III*(42); dorsal rays 3*(42); pelvic rays I,3*(42); segmented caudal rays 10*(20), 11(6), 12(16); vertebrae 14+27(1), 14+28(3), 14+29(2); lower gill rakers on outer face of first arch 7*(6), 8(29), 9(4); pored lateral line scales 21–30 (arched portion of line) + 25–31 (straight portion of line), anterior lateral line scales 21(5), 22(3), 23(3), 24(9), 25*(3), 26(6), 27(2), 30(1); posterior (straight portion) lateral line scales 25(5), 26(7), 28*(6), 29(4), 31(2); total lateral line scales 46, (1), 47(1), 48(2), 49(33), 50(4), 51*(4), 52(7), 53*(9), 55(2); branchiostegal rays 6*(9); other counts are shown in Tables 1–5 . Vomer with conical teeth an inverted V, 1–2 rows in specimens less than 45 mm SL and 2–3 rows in larger specimens; palatine without teeth. Head and body laterally compressed; head length moderate, proportion decreasing with increasing size (23.6– 29.3% SL in adults, 27.6–31.4% SL in juveniles less than 40 mm SL); snout gently curved in side view, snout shorter than eye diameter (about 0.7 of eye length in adults and 0.5 in juveniles), snout length 2.7.–7.1% SL, not changing significantly with size, eye moderate to large, relative size decreasing significantly with growth (8.7–10.6% SL in specimens < 40 mm SL, 7.5–9.3 SL in specimens 41–66 mm SL, 5.1–7.1% SL in specimens 84–147 mm SL); interorbital narrow, about one-half to two thirds eye diameter; mouth short, jaws reaching to below posterior quarter of pupil, upper jaw length, 7.0–10.0% SL, showing slight increase with size (p = 0.01); anterior nostril at end of short tube, about 3 nostril diameters above upper lip, with short spatulate nasal tentacle expanding distally, often with two small lobes distally; 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 slightly shorter and pointed, rakers becoming progressively shorter on following arches; tongue tip broadly pointed; upper jaw with 4–5 rows of incisiformn teeth (slightly compressed), teeth in outer row largest with pointed tips, broadening laterally, usually without lateral tips, 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. Genital valve with irregular margin, sometimes forming up to 4 small lobes, fully covering genital opening in females. Intromittent organ pointed, without lateral lobes. 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 middle of first dorsal fin; scales cycloid overlapping and forming distinct rows, becoming scattered and nonimbricate on caudal peduncle, easily visible giving skin a granulated appearance; pectoral fin base covered with small embedded scales, scales extending onto base of fin to a maximum of basal one-quarter of rays; scales covering proximal bases of dorsal spines in 1–2 rows, not extending onto membranes between spines, not extending onto 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. FIGURE 12. Head of Heteroclinus wilsoni , 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 on QVM 1972/5/[ex346], 44 mm SL. All fin-rays unbranched; first dorsal fin origin, above or just behind posterior margin of 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 connected to body just before second dorsal fin or rarely basally to second dorsal fin; second dorsal origin above a point just before pectoral insertion and behind pelvic insertion; first spine of second dorsal slightly shorter than following spine, spines becoming progressively longer posteriorly, with last spine the longest; dorsal membrane slightly incised, with membrane connecting below tips of spines; last two dorsal rays shorter and widely separated from anterior ray; anal origin below 9 th to 10 th spine of second dorsal fin, anal spines distinctly shorter than rays; posterior rays becoming progressively longer, last two rays shorter, slightly closely spaced and usually widely separate from anterior rays; last anal ray connected along basal half of ray by membrane to or before middle of caudal peduncle; pectoral fin with rounded posterior margin, central rays longest, reaching to above first to third anal ray; pelvic fins with hidden spine, 3 developed rays; innermost ray about half to slightly more than half length of other two rays, fin extending to just anterior to anus; caudal fin with truncate to slightly rounded posterior margin, length (16.4–22.6% SL), proportion not changing significantly with increasing size; caudal fin with 10 thickened rays and an upper and a lower smaller ray, segmented in about half of the specimens examined, 3–5 upper and lower very short procurrent simple rays difficult to discern. Coloration of freshly collected material ( Plate 1C & D ). Head and body reddish brown or green with 7 or 8 darker brown saddles or vertically elongate spot below dorsal fins, rarely uniformly green without bands; first band below first dorsal fin extending onto operculum, second behind second dorsal fin origin, last below dorsal rays; vertical bands extending dorsally from spot onto dorsal fin, with 3–4 spines between bands; dark brown vertical bands extending ventrally from dorsal spot, first bend extending ventrally from first dorsal fin across anterior operculum, second and third bands on belly, bands 3–8 extending on to anal fin, spaces between brown translucent light brown or green; a dark brown band from ventral eye to end of jaws, with thin dark edges; a small oblique silver spot below posteroventral margin of eye, behind silver spot a thin brown line extending obliquely downward from eye; oblique almost vertical bar sometimes present on preoperculum and operculum; white to light crescent-shaped bar on mid to upper portion of pectoral fin base; large white to silver spot sometimes present posterior to eye; white spots sometimes present on pectoral fin base and belly; first dorsal fin dark, other fins mottled. Live coloration . The coloration is similar to freshly collected coloration, except that the background coloration is lighter, usually light green (see Kuiter & Kuiter 2018 ). Coloration in alcohol. Head and body uniformly light brown to yellowish brown, without distinct marking. Distribution . The species is known from the Eden region in southern New South Wales to Kangaroo Island and Yorke Peninsula, South Australia , including Victoria and scattered localities in Tasmania . The species is associated with algae and kelp on rocky reefs to depths of at least 10 m . Etymology. The species was named after the collector of the original material in 1887–1888, Bracebridge Wilson. Remarks . Heteroclinus wilsoni is most similar to H. colemani in having a high first dorsal fin and in being deeper bodied than other species. Regression analyses ( Table 10 ) indicated a y-intercept, significantly different from 0 for head length (p <0.001), predorsal length (p = 0.002), body depth at anal origin (p <0.001) and eye length (p <0.001). Rank correlation indicated all characters decreased significantly with size, except for body depth at anal origin, which increased with size. Rank correlation tests of three additional characters showed a significant decrease with size including jaw length (p = 0.013), caudal peduncle depth (p = 0.008) and caudal peduncle length (p = 0.014). That analysis also showed the third pelvic ray length increasing with size (p = 0.002). Other characters showed a nonsignificant change with size (p value varying from 0.07 to 0.55). The differences between the two test methods may be due to the small sample size (n=33) for this species. The species reaches a length of 147 mm SL. FIGURE 13. Map showing distribution of Heteroclinus wilsoni based on material examined. TABLE 10. Regression and proportion parameters Heteroclinus wilsoni . 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	33	21–147	0.253	0.240	0.266	1.75	<001	33.7%	26.5%	23.6%	31.4%	decrease	-0.313	0.005
Head width	20	39–118	0.156	0.137	0.175	0.29	0.420	16.3%	15.8%	13.8%	18.0%	ns	-0.251	0.066
Predorsal	32	21–147	0.131	0.120	0.143	1.46	0.002	20.0%	14.1%	12.4%	20.2%	decrease	-0.357	0.002
Body depth anal origin	40	21-147	0.350	0.334	0.367	-2.93	<<.001	21.1%	33.0%	23.8%	36.4%	increase	0.569	<<0.001
Caudal peduncle depth	28	21–118	0.058	0.053	0.062	-0.18	0.279	4.9%	5.6%	4.8%	6.4%	ns	0.329	0.008
Caudal peduncle length	26	21–118	0.095	0.082	0.109	0.11	0.782	10.1%	9.6%	8.4%	13.3%	ns	-0.342	0.014
Jaw length	32	21-147	0.080	0.074	0.086	0.41	0.086	9.9%	8.3%	7.0%	10.0%	ns	-0.276	0.013
Eye length	33	21–118	0.043	0.038	0.047	1.89	<<.001	13.3%	5.9%	5.1%	10.6%	decrease	-0.781	<<0.001
Snout length	32	21–147	0.041	0.035	0.047	0.32	0.153	5.6%	4.4%	2.7%	7.1%	ns	-0.159	0.224
Pectoral fin length	31	21–147	0.208	0.192	0.224	-0.79	0.187	17.0%	20.3%	16.2%	23.9%	ns	0.202	0.110
Pelvic fin length	30	21–147	0.191	0.180	0.201	-0.31	0.432	17.6%	18.9%	14.5%	20.8%	ns	0.172	0.181
Third pelvic ray length	30	21–147	0.113	0.102	0.123	-0.65	0.096	8.2%	10.8%	7.9%	13.2%	ns	0.370	0.002
Second dorsal spine length	35	21–147	0.132	0.121	0.144	-0.11	0.786	12.7%	13.2%	10.5%	15.4%	ns	0.071	0.551
Last dorsal spine length	34	21–147	0.127	0.115	0.139	0.46	0.290	14.9%	13.0%	10.6%	16.2%	ns	-0.061	0.361
First dorsal ray length	26	21–118	0.158	0.146	0.170	-0.13	0.768	15.2%	15.7%	13.0%	17.3%	ns	0.194	0.165
Caudal fin length	26	21–118	0.192	0.178	0.206	0.62	0.213	22.3%	19.7%	16.4%	22.6%	ns	-0.108	0.220

Neotype selection . The description of Clinus wilsoni by Lucas indicated that he had multiple species. The counts given for the dorsal spines and rays were 27–28 for the second dorsal fin and 3 rays, with a window between the first and last two rays, bands on the body and the dorsal fin membrane attaching to the base of the caudal fin. That combination indicates the species here is treated as Heteroclinus wilsoni or H. heptaeolus . However, it is apparent that he had multiple specimens and species and indicated that the head coloration was variable. He mentioned that the spines in the first dorsal fin were rather higher than the anterior rays of the second dorsal fin, suggesting the species treated here as H. wilsoni . In addition, he described and figured one specimen , which he regarded as slightly different, with 24 spines in second dorsal fin and 17 anal rays and 3 dorsal rays and first dorsal spines not longer than anterior spine of second dorsal fin. Those features would suggest Heteroclinus heptaeolus . However, Lucas regarded the coloration as reddish, unusual for H. heptaeolus and the body is shown to be very deep, more characteristic of H. colemani or H. wilsoni . To fix the identity of the species we select AMS I.20166-001, from Kangaroo Island, South Australia as the neotype of the species. Although originally described from near Port Phillip heads, Victoria , we select a specimen with the best photo showing fresh coloration. No type specimens have been found in museums in Australia for the species described by Lucas (1891) .