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 colemani n. sp. Plate 1A, 1B , Figures 2 , 4 , 5 , Tables 1–6 , 11 , 12 Common Name: Coleman’s Weedfish Cristiceps wilsoni Lucas, 1891: 10 , pl 3 (fig. 1), in part, figured specimen only, Port Phillip, Victoria . Heteroclinus sp. 4 .— Rennis, Hoese & Gomon 1994: 749 ( New South Wales to Tasmania and Kangaroo Island, South Australia ); Hoese, Gomon & Rennis 2008: 703 ( New South Wales to Tasmania and Kangaroo Island, South Australia ). Heteroclinus sp. 1 .— Kuiter 1993: 327 (habitat and distribution); Kuiter & Kuiter, 2018: 284 . Holotype . AMS I.20174-005, a 47 mm SL female, Knob Point , Kangaroo Island , South Australia , 35°36’31”S , 137°13’54”E , 3–8 m , B. Russell , 6 March 1978 . PLATE 1. High dorsal fin species: A) Freshly collected holotype of Heteroclinus colemani AMS I. 20174-005, 47 mm SL female from Kangaroo Island, South Australia; B) Freshly collected paratype of Heteroclinus colemani , AMS I. 24292-001, 37 mm SL female, from Sydney, New South Wales, photo by Rudie Kuiter; C) Heteroclinus wilsoni Neotype AMS I.20166-001, female 113 mm SL, Stokes Bay, Kangaroo Island; D) Heteroclinus wilsoni AMS I. 19774-004, 1(100), Portsea Pier, Port Phillip, Victoria, photo by R. Kuiter. 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. Paratypes . New South Wales : AMS I.16863-008, 1(65), Southwest Bowen Is. , Jervis Bay ,ACT, 0–2 m , 35°07’S , 150°46’E , D. Pollard and P. Straw , 2–3 May 1973 ; AMS I.24292-001, 1(37), Long Bay , Sydney , 33°58’S , 151°15’E , R . Kuiter , 28 December 1976 ; AMS I.44627-046, 3(69–72), Baronda Headland , 36°41’11”S , 149°59’58”E , A. Hay , 8 April 2008 ; AMS I.45633-076, 3(59–60), Washerwomans Beach , 35°14’39”S , 150°32’09”E , M. McGrouther , 16 March 2011 . Tasmania : AMS I.24287-002, 1(46), Deal Is. , 39°29’ S , 147°21’ E , N. Coleman , 8 May 1974 ; NMV A.4271, 1(48), Southport , 43°30’S , 147°00’E , R . Wilson , 27 April 1985 ; Victoria : NMV A.2364 , 1(38), Oberon Bay , Wilsons Promontory , 8–13m , 39°04 ,S , 146°20’E , M. Gomon & J. Jones , 6 February 1982 ; NMV A.24371, 1(46), east of Eagles Nest , 5–11 m , 38°40.76’S , 145°39.24’E ; T . O’Hara et al ., 1 April 1997 ; NMV A.24375, 1(42), off Honeysuckle Hill , 38°40.54’S , 145°37.78’E , T . O’Hara et al ., 1 April 1997 . South Australia : SAM F.3561, 1(66), reef at Port McDonnell , 38°03’15”S , 140°42’E , Dr R .H. Holmes , August 1970 . Non-type material. AMS . B.6629, 1 (83), locality unknown. Diagnosis . Dorsal fins III, XXIV–XXV, 3; anal fin II, 18–19; pectoral rays 13, uppermost ray very short and easily missed; gill rakers on outer face of first arch 3 + 7–8 = 10–11; circumorbital head pores uniserial (13–17 pores); orbital tentacle elongate with a rounded lobe, expanding slightly distally; nasal tentacle simple, round, flap length about equal to tubular base length; middle gill rakers and uppermost rakers on outer face of first arch not branched dorsally; first dorsal fin elevated in both sexes (second dorsal spine 11.6–16.7% SL, not showing significant change with size), fin originating over posterior end of eye to about middle of preoperculum; third dorsal spine in front of a vertical from pelvic 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 25.9–30.6% SL in specimens 36.7–50 mm SL and 30.3–34.6% SL in specimens 51–83 mm SL). Body often red to reddish orange in life; 6–7 dark saddles along dorsolateral surface of trunk and tail extending onto dorsal fin, with 6–8 clear windows between spines, no spines in windows; windows sometimes incomplete, with pigment above and or below clear parts of windows; three silver to white bars with brown margins radiating from eye, the first across snout and middle of upper jaw, the second extending ventrally to end of mouth, the third extending posteriorly from mid posterior edge of eye. Description . Based on 16 specimens , 8 males and 8 females , 36–82.8 mm SL. First dorsal III* (16); dorsal rays 3*(16); pelvic rays I,3* (16); segmented caudal rays 10*(12), 11(2), 12(1); vertebrae 14+2(2); lower gill rakers on outer face of first arch 7*(5), 8(8); circumorbital head pores uniserial (13–17 pores); pored lateral line scales l7–26 (arched portion of line) + 21–26 (straight portion of line), anterior lateral line scales 17(1), 19(1), 22(3), 23*(6), 24(5); posterior lateral line scales 21(2), 22(1), 24(5), 25(2), 26*(6); total lateral line scales 41, (1), 45(3), 46(2), 47(3), 48(3), 49*(3), 50(1); branchiostegal rays 6*(5), other counts are shown in Tables 1–5 . Vomer with conical teeth arranged in 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 moderate, length (26.7–31.8% SL, not showing significant change with increasing size); snout gently convex in side view, snout shorter than eye diameter (0.5–0.7 of eye length, (3.3– 6.4% SL),), proportion not changing significantly with size; eye moderate, relative size decreasing with growth (8.4–8.7% SL in specimens less than 40 mm SL, 7.3–7.9% SL in specimens 44–65 mm SL, 6–6.2% SL in specimens 68–83 mm SL); interorbital narrow, about one-third eye diameter; mouth small, jaws reaching to below anterior edge of pupil to middle of eye, upper jaw length 8.9–12.0% SL, not showing significant change with size; anterior nostril at end of short tube, above upper lip, with short spatulate nasal tentacle expanding distally, without side lobes or branches; posterior nostril with elevated rim above anterior margin of eye; gill rakers on outer face of first arch, long and slender, slightly shorter than filaments; rakers on inner face of first arch and following arches shorter and pointed; tongue tip broadly pointed; upper jaw with 4–5 rows of incisiform teeth (slightly compressed), outer row with pointed tips and very small lateral cusps, teeth becoming more conical and pointed in inner rows, rows teeth tapering to two rows near end of jaw; teeth in lower jaw incisiform (slightly compressed) to conical without lateral cusps. Genital valve composed of three small lobes connected at base in holotype , other specimens with single lobe with irregular margin, fully covering genital opening in females. Intromittent organ pointed, without lateral lobes. Head pores as shown in Figure 4 : circumorbital and preopercular pores uniserial. Head largely naked, body scales small and cycloid extending forward to above operculum below anterior end of first dorsal fin; scales cycloid overlapping and forming distinct rows, becoming scattered and nonimbricate on caudal peduncle; pectoral fin base covered with small embedded scales, scales extending onto base of fin to a maximum of basal quarter of rays; scales covering bases of dorsal spines and membranes between spines, extending to a maximum of basal quarter of fin, sparse or absent on membrane windows, not extending onto dorsal rays; scales not extending onto anal fin; scales extending onto base of caudal rays to about one-tenth of length of fin; scales on fins highly variable and possibly easily lost; lateral line well developed, scales extending to caudal fin; 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 or just behind posterior margin of eye, fin elevated and spines slightly longer than spines in second dorsal fin, second spine usually longest, first and third spines subequal in height; last spine of first dorsal fin connected to basal one-fifth of second dorsal spine; second dorsal origin above a point just behind pectoral fin insertion and behind pelvic fin 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 and widely separated from first ray; anal origin below 8 th to 10 th spine of second dorsal fin, anal spines distinctly shorter than rays; posterior rays becoming progressively longer, last two rays shorter, closely spaced and 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 second or third anal ray; pelvic rays not reaching to anus, inner ray about 1/2 to slightly more than 1/2 length of second ray; caudal fin truncate to slightly rounded, (17.2–21.7% SL, not showing significant change with increasing size), caudal fin with 10 thickened rays and an upper and a lower smaller ray, segmented in 11% of specimens examined, 3–5 upper and lower very short procurrent simple rays difficult to discern. Coloration of freshly collected material ( Plate 1A & 1B ). Head and body brown to reddish or reddish orange with dark brown spots along trunk and tail; 6–7 darker saddles along dorsolateral surface of trunk and tail extending onto dorsal fin; large white to silver spot behind pectoral fin sometimes followed by a series of smaller spots posteriorly and dorsally; 3 silver to white bars with brown margins radiating from eye, the first across snout and middle of upper jaw, the second extending ventrally to end of mouth, the third extending posteriorly from mid posterior edge of eye; orbital tentacle bright red to brown; dorsal and anal fins red to brown interrupted by six clear areas; pectoral, ventral, and caudal fins with a series of red or brown spots forming transverse bands. FIGURE 4. Head of Heteroclinus colemani , 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. TABLE 1 . Number of spines in second dorsal fin of species in the Heteroclinus heptaeolus complex. An asterisk indicates count of holotype or neotype.

23	24	25	26	27	28	Mean	Variance
H. colemani	-	2	14*	-	-	-	24.9	0.117
H. heptaeolus	-	-	5	33	106*	14	26.8	0.343
H. longicauda	-	2	17	12*	-	-	25.3	0.359
H. whitleyi	7	69*	23	-	-	-	24.2	0.280
H. wilsoni	-	-	1	2*	26	16	27.3	0.427

TABLE 2. Number of anal rays of species in the Heteroclinus heptaeolus complex.An asterisk indicates count of holotype or neotype.

15	16	17	18	19	20	21	22	Mean	Variance
H. colemani	-	-	-	4	12*	-	-	-	18.8	0.200
H. heptaeolus	-	-	-	6	28	110*	12	19.8	0.382
H. longicauda	-	-	1	16*	11	-	-	-	18.4	0.312
H. whitleyi	2	6	65*	17	-	-	-	17.1	0.352
H. wilsoni	-	-	-	-	-	6*	27	12	21.1	0.391

TABLE 3. Total number pectoral rays of species in the Heteroclinus heptaeolus complex. An asterisk indicates count of holotype or neotype.

11	12	13
H. colemani	-	-	16*
H. heptaeolus	4	138*	-
H. longicauda	-	1	30*
H. whitleyi	7	82*	-
H. wilsoni	1	41*	-

TABLE 4. Total number of gill rakers on outer face of first gill arch of species in the Heteroclinus heptaeolus complex. An asterisk indicates count of holotype or neotype.

6	7	8	9	10	11	12	Mean	Variance
H. colemani	-	-	-	-	5*	8	-	10.6	0.256
H. heptaeolus	2	15	94*	29	3	-	-	8.1	0.438
H. longicauda	-	-	3	11	9*	1	-	8.9	3.297
H. whitleyi	-	-	2	2*	42	37	10	10.6	0.642
H. wilsoni	-	-	-	3	6	26*	4	10.8	0.536

TABLE 5. Circumorbital pores of species in the Heteroclinus heptaeolus complex. An asterisk indicates count of holotype or neotype.

SPECIES	12	13	14	15	16	17	18	19	20	Mean	Variance
H. colemani	-	2	1	4	5*	1	-	-	-	15.2	1.474
H. heptaeolus	-	-	3	3	10*	11	11	9	3	17.3	2.394
H. longicauda	-	1	3	5	7	2*	2	-	-	15.6	1.726
H. whitleyi	1	7	33	11*	2	-	-	-	-	14.1	0.553
H. wilsoni	-	-	-	2	9	10*	9	4	1	17.2	1.459

Live coloration . Similar to freshly collected coloration, except one aquarium specimen with brownish, rather than reddish body. Second dorsal fin with 5–8 distinct transparent windows, first usually behind fifth spine and last after 23 rd or 24 th spine and a large window between first and second rays and small window on membrane behind last ray. Anal fin with similar windows, spaced irregularly. TABLE 6. Regression and proportion parameters of Heteroclinus colemani . 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 range of proportion		Size trend	Tau	p
Head length	16	36–82.8	0.270	0.238	0.302	1.210	0.173	30.3%	28.4%	26.7%	31.8%	ns	-0.167	0.368
Head width	14	36–82.8	0.118	0.093	0.142	1.209	0.093	15.1%	13.2%	11.7%	16.8%	ns	-0.297	0.139
Predorsal	16	36–82.8	0.131	0.110	0.153	1.467	0.023	17.2%	14.9%	13.9%	18.6%	decrease	-0.383	0.019
Body depth anal origin	16	36–82.8	0.362	0.321	0.404	-2.896	0.020	28.2%	32.7%	25.9%	34.6%	increase	0.467	0.006
Caudal peduncle depth	15	36-82.8	0.057	0.043	0.070	-0.060	0.873	5.5%	5.6%	4.7%	6.6%	ns	0.105	0.586
Caudal peduncle length	15	36–82.8	0.127	0.061	0.194	-0.498	0.783	11.4%	12.1%	8.8%	16.0%	ns	0.162	0.400
Jaw length	15	36–82.8	0.087	0.060	0.114	0.507	0.492	10.1%	9.3%	12.0%	8.9%	ns	-0.105	0.586
Eye length	15	36–82.8	0.045	0.033	0.058	1.514	0.001	8.8%	6.4%	8.7%	6.0%	ns	-0.657	<0.001
Snout length	15	36–82.8	0.052	0.032	0.071	-0.282	0.598	4.4%	4.8%	3.3%	6.4%	ns	0.143	0.458
Pectoral fin length	15	36–82.8	0.220	0.175	0.265	-1.443	0.253	18.0%	20.3%	14.4%	21.9%	ns	0.067	0.729
Pelvic fin length	15	36–82.8	0.205	0.165	0.245	-1.443	0.200	16.5%	18.8%	14.7%	20.4%	ns	0.333	0.083
Third pelvic ray length	15	36–82.8	0.101	0.071	0.132	-0.629	0.449	8.4%	9.4%	6.4%	11.0%	ns	0.238	0.216
Second dorsal spine length	15	36–82.8	0.112	0.079	0.144	1.554	0.095	15.5%	13.1%	11.6%	16.7%	ns	-0.314	0.102
Last dorsal spine length	15	36–82.8	0.112	0.074	0.150	1.573	0.148	15.6%	13.1%	12.1%	17.7%	ns	-0.429	0.026
First dorsal ray length	14	36–82.8	0.142	0.109	0.174	1.242	0.182	17.6%	15.7%	14.5%	17.9%	ns	-0.187	0.352
Caudal fin length	14	36–82.8	0.194	0.160	0.228	-0.065	0.943	19.2%	19.3%	17.2%	21.7%	ns	0.077	0.702

FIGURE 5. Map showing distribution of Heteroclinus colemani based on material examined. Coloration in alcohol . Head and body uniformly light yellowish brown, without distinct marking. Clear windows on interspinal membranes in life appearing as slightly lighter areas than remaining interspinal membranes. Distribution . The species is known from Sydney, New South Wales to Kangaroo Island, South Australia , including Victoria and scattered localities in Tasmania . It has been recorded as usually associated with red algae on rock faces around rocky reefs from 5–15 m ( Kuiter 1993 ). Etymology. Named for Neville Coleman, who collected material of this species and other clinid species studied. Remarks . The species is distinctive from other species in the complex in usually having a reddish body coloration; orbital tentacle expanding distally, 3 bars radiating from the eye and combination of pectoral, caudal, dorsal and anal ray and gill raker counts. The sample size for this species was small with measurements obtained from 14– 16 specimens for various measurements. Regression analyses indicated a y-intercept, significantly different from 0 for predorsal length (p = 0.023), body depth at anal origin (p = 0.020) and eye length (p = 0.001). Rank correlation indicated all characters decreased with size, except for body depth at anal origin which increased. Characters where the regression analyses showed a y-intercept not significantly different from 0, also showed no significant values for the rank correlation of the proportions tested with Kendall’s Tau ( Table 6 ). In coloration and deep body, the species is most similar to H. wilsoni . Heteroclinus colemani has 13 pectoral rays (versus 12) and lower second dorsal spine and anal ray counts. Too few specimens were available to determine colour variation. However, in the two fresh specimens of this species the dark bar below the eye is much narrower than the pupil diameter, while in the two fresh photos of H. wilsoni the bar is about as large as the pupil diameter. The species has a higher first dorsal fin than other species in the complex.