Clinus musaicus, Holleman & Heyden & Zsilavecz, 2012

Holleman, Wouter, Heyden, Sophie Von Der & Zsilavecz, Guido, 2012, Delineating the fishes of the Clinus superciliosus species complex in southern African waters (Blennioidei: Clinidae: Clinini), with the validation of Clinus arborescens Gilchrist & Thompson, 1908 and Clinus ornatus Gilchrist & Thompson, 1908, and with descriptions of two new species, Zoological Journal of the Linnean Society 166 (4), pp. 827-853 : 835-843

CLINUS MUSAICUS SP. NOV.

FIGS 1 ; 3A–F ; 9

Holotype

SAIAB 96616 , male 108 mm SL, Windmill , nr Simonstown, South Africa (34°12.025 ′ S, 18°27.390 ′ E), sandy plain with rubble and shell fragments; collected by Guido Zsilavecz (G4), 9 m depth, 25 April 2009.

Paratypes

SAIAB 96614 , female 80.5 mm SL, Windmill, nr Simonstown , South Africa (24°12.025 ′ S, 18°27.390 ′ E), sandy plain with rubble and shell fragments; collected by Guido Zsilavecz (G2), 10 m depth, 25 April 2009 ; SAIAB 96613 , female 145 mm SL, Windmill, nr Simonstown , South Africa (24°12.025 ′ S, 18°27.390 ′ E), sandy plain with rubble and shell fragments; collected by Guido Zsilavecz (G3), 10 m depth, 25 October 2008 ; SAIAB 96615 , male 183 mm SL, Betty’s Bay , Western Cape, collected by Sophie von der Heyden, 2007 ; SAM MB-F037962 , male 84 mm SL, Windmill, nr Simonstown , South Africa (34°12.025 ′ S, 18°27.390 ′ E), sandy plain with rubble and shell fragments; collected by Guido Zsilavecz (G1), 9 m depth, 25 October 2008 .

Diagnosis

A species of the Clinus superciliosus complex with XXXVIII–XL, seven or eight dorsal-fin elements, 15 pectoral fin rays; crest of first three spines, triangular, separated by a deep notch from the fourth spine; origin of dorsal fin over posterior margin of preopercle; dentary pore pattern 2 + 2/2 + 2; palmate orbital and nasal cirri; four white scallop-shaped areas ventrally on the body, with dark reticulate markings, and no pigment on the pectoral or caudal-fin membranes.

Description

Dorsal fin XXXVIII–XL, 7 or 8 (XXXIX, 8); anal fin II, 27–28, usually 28 rays; pectoral fins 15; pelvic fins I, three, inner ray thin and closely applied to second ray, third to half the length of second ray. Vertebrae 18 or 19 + 33–34 (19 + 34). Lateral line scales 38–40 + 32 or 33 (usually 38 + 33). Head length 4.0–4.5 [4.2] in SL; eye diameter 3.0–4.4 [3.7, see note below], maxilla 2.4–2.8 [2.6], snout 3.7–4.1 [3.9] in head length; dorsal-fin origin 1.7–1.9 [1.8] in head length; dorsal fin height 10.6–15.2 [12.6] in SL.

Head profile fairly blunt (~73°), flat in large specimens, slightly rounded in smaller ones. Mouth downturned, maxilla extending to vertical, through anterior margin of pupil. Orbital cirrus a flattened stalk on the upper margin of the orbit, about equal to pupil diameter, palmate, with five ‘fingers’ in smaller specimens, more numerous and longer in larger specimens ( Fig. 3B ), suggesting that, as in C. superciliosus , they become more complex with increasing size; anterior nasal cirrus flat, semi-triangular, and with serrations along the long edge. Crest of males triangular, the first spine the longest and 1.4–1.8 height of dorsal fin, with no apparent difference between males and females. Dorsal-fin spines with single ‘cirrus’ at the tips, which diminish in size posteriorly.

Live colour: Above midline a variety of colours ranging from pale grey, olive green, pink, to brown, the colour in ‘lozenges’, with between four and six darker ‘lozenges’ that straddle the midline; below midline white, most often as four or five ‘scallop’ marks with darker reticulations, the entire effect being one of a mosaic; head mostly whitish below eye, and variously spotted and reticulated with darker markings; except for anal fin, fin membranes translucent to transparent, with pastel colours on elements only; anal fin usually white with other colours present on body. Photographs of the species underwater show that the colours approximate habitat colours and render individuals quite cryptic.

Clinus musaicus sp. nov. shows some variation in colour, while retaining a basic pattern of irregular, hourglass-shaped darker bars and four white, scallop-shaped areas below the midline, in pinks, mauves, and other pastel colours.

Colour in preservative: Body pale grey above midline, with darker grey ‘lozenges’ that are darkest above the midline and white below, with faint reticulations; head variously spotted and reticulated in greys and white; crest grey with white spots, remainder of dorsal fin pale grey with darker grey ‘shadows’ of lozenges on side; caudal fin translucent, with dark grey on rays forming three irregular bands, broadest along margin, anal fin white with irregular grey bars, pectoral fins white with dark-grey spots, pelvic fins white with grey bars.

Etymology

‘Musaicus’, the Latin for ‘mosaic’, is given for its colour pattern, the reticulations of which are reminiscent of mosaics.

Comparisons

Clinus musaicus sp. nov. can be distinguished from C. arborescens by colour pattern and number of dorsalfin rays (mean of eight vs. mean of six), from C. ornatus and C. superciliosus by colour patterns and number of dorsal-fin spines (mean of 39 vs. means of 32–36), and from Clinus exasperatus sp. nov. by colour pattern and body depth (mean of 4.6 in SL vs. mean of 3.0 in SL). It can also be distinguished from C. superciliosus by the form of the orbital cirrus, palmate vs. simple.

Distribution

Known from Betty’s Bay, False Bay, and Oudekraal, Western Cape, South Africa ( Fig. 1 ).

GenBank accession numbers

JF320834 – JF320836 (mtDNA control region) ; JF320848 – JF320850 (mtDNA 16S rRNA) ; JF320862 – JF320864 (mtDNA 12rRNA) ; JF320876 – JF320878 (rhodopsin).

CLINUS ORNATUS GILCHRIST & THOMPSON, 1908

FIGS 1 ; 4A–D ; 5B

Clinus ornatus Gilchrist & Thompson, 1908: 116 (type locality, False Bay , Western Cape).

Clinus superciliosus (in part): Smith, 1946; Penrith, 1969: 57, figure 22; Smith in Smith & Heemstra, 1986: 765.

Description

Dorsal fin XXXIII–XXXVI, 7 or 8 (XXXV, 7); anal fin II, 25–27; pectoral fins 14 or 15 (15); pelvic fins I, 3, inner ray minute. Gill rakers 2 or 3 + 6. Vertebrae 16 or 17 + 31 or 32, total 48–49. Lateral line scales 35–40 + 27–31 (36 + 28 or 29). Head length 3.7–4.0 [3.9] in SL; eye diameter 3.0–3.9 [3.5]; maxilla 2.0–2.5 [2.2] and snout 3.8–4.6 [4.2] in head length; dorsal-fin origin 1.7–2.0 [1.9] in head length; dorsal fin height 8.4–10.5 [9.3] in SL.

Head profile blunt (~83°) and slightly rounded. Mouth downturned, maxilla extending to vertical through posterior margin of pupil. Orbital cirrus a flattened stalk with long side branches, in length about equal to pupil diameter ( Fig. 4B ); nasal cirrus similar but smaller with shorter branches. Dorsal-fin origin just posterior to hind margin of orbit; crest of males about 1.2 times height of dorsal fin, first two spines about same length and only slightly longer than third; distance between spines 3 and 4 about equal to distance between spines 1 and 3. Dorsal-fin spines with a single filament at tips for about half the length of fin.

Live colour: Gilchrist & Thompson (1908) described it as ‘... usually vivid. A yellowish or greyish brown, with small dark brown spots; 6 irregular transverse bands of dark brown edged with orange and extending onto dorsal fin; orange blotches and markings on head, body, dorsal and anal fins; pectorals transparent, with 5 rows of brown spots touched with orange; caudal almost transparent with 2 transverse bands of orange near the extremity and spotted with orange and brown; ventrals light green often barred with orange’. Photographs taken by the third author and Carel V.D. Colff show that individuals are variable in colour, blending in with the substrate where they are found. Most characteristic are the many white spots on the upper half of the body and the dark reticulations on the paler lower part of the body and head.

Colour in preservative: The specimens are old and have become pale yellow. A few of the large specimens show reticulate markings on the face and remnants of body patterning similar to C. superciliosus .

Comparisons

Clinus ornatus can be distinguished from all the other species of the complex by the form and position of the dorsal-fin crest. It is the only one of the species in which the three spines of the crest of males are subequal in length, whereas in the other species the first spine (first two spines in C. arborescens ) is longer than subsequent spines.

The origin of the dorsal fin of Clinus ornatus lies close to a vertical through the posterior margin of the eye, whereas in the other species the origin of the dorsal fin lies over or slightly forwards of the posterior margin of the preopercle. Furthermore, the third dorsal-fin pterygiophore of C. ornatus lies over the first vertebra and exoccipitals, whereas the supraoccipital has become concave to accommodate the first two pterygiophores, which have been ‘stretched’ anteriorly ( Fig. 5B ). By comparison, the third pterygiophore of C. superciliosus lies above and posterior to the first vertebra, with the first two pterygiophores over the supraoccipital; the first pterygiophore is almost equilateral in shape ( Fig. 5A ).

Distribution

The species has been found subtidally only in False Bay ( Fig 1 ).

Remarks

Clinus ornatus was described from nine males (there are only eight syntypes in the collections of the SAM, as listed below, and there is a specimen in the BMNH SL; eye diameter 2.9–4.1 [3.2], maxilla 2.4–3.0 [2.7], snout 3.7–5.2 [4.2] in head length; dorsal-fin origin 1.3–1.6 [1.4] in head length; dorsal fin height 12.0– 17.2 [14.8] in SL.

Head profile fairly blunt (77–84°) and rounded in individuals longer than 50 mm SL. Mouth downturned, maxilla extending to vertical through posterior margin of pupil. Interorbital convex. Orbital cirrus varies between a small, flat nubbin and a parallel-sided, flattened stalk with rounded or bifurcate end, in length about equal to pupil diameter ( Fig. 4F ). Dorsal-fin crest of males about 2.3 height of dorsal fin, of females about 1.0–1.3 height of dorsal fin. Dorsal-fin spines with a single filament at the tips, for about half the length of fin.

and one in the MNHN, see below, under C. superciliosus types), with one specimen taken at 24 m near Roman Rock along with the specimens of C. arborescens noted above. The species has however been identified from photographs taken by the third author and others.

Material examined

Lectotype: SAM MB-F038012 (158 mm SL), selected from SAM 9875 .

Paralectotypes: SAM 9874 (138 mm) ; SAM 9875 (6: 68.0–175.0 mm), all from False Bay , Western Cape .

CLINUS SPATULATUS BENNETT 1983

FIGS 1 ; 4E–I ; 9

Clinus spatulatus Bennett, 1983: 2 (type locality Bot River Estuary , Western Cape, South Africa); Smith in Smith & Heemstra, 1986: 764.

Description (partially from Bennett, 1983)

Dorsal fin XXXII–XXXV, 6–8 (XXXIV, 7); anal fin II, 23–26 (25 rays); pectoral fins 13–15 (14, rarely 13 or 15); ventral fins I, 2–3, inner ray reduced. Vertebrae 17 or 18 + 28 or 29. Gill rakers 2–4 + 6–8. Lateral line scales 26–31 + 26–37 (usually 27–29 anteriorly and 30–33 posteriorly); anterior portion with double pores for only the anterior half. Head length 3.6–4.1 [4.0] in Live colour ( Fig. 4G–F ): Clinus spatulatus shows little variation in colour pattern, but is the only species that is distinctly sexually dichromatic. Males dark olive green with bright, white spots at the base of the crest, on the opercle and pectoral-fin base, and at the base of each orbital cirrus. The pattern of these spots differs for individual males. Females range from greyish to various shades of green, with indistinct mottling. Individuals living on sandy bottoms photographed in the Bot River estuary show a typical, though paler, ‘superciliosus’ pattern of dark and light areas, which would render them cryptic amongst the sparse vegetation on the sandy substrate.

Colour in preservative: The colour pattern of C. spatulatus is very similar to that of the common, mottled form of C. superciliosus , but lacks the dark bar on the pectoral fin base and the dark spot on the dorsal-fin crest. After some years in preservative the pattern tends to fade: females become a light brown, lightly marbled, with slightly darker dorsal and anal fins; and males become dark brown, with darker dorsal and anal fins. Very dark males have three clearly defined pale areas on the pectoral fin base, four on the opercle, one below the eye at 4 o’clock, and one at the base of each orbital cirrus, and without marks or bars on the lips or chin. There are also several pale spots irregularly along the midline and on the dorsum.

Comparisons

Clinus spatulatus shares with C. exasperatus sp. nov. a very simple orbital cirrus, which is sometimes reduced to a mere nubbin, compared with the more elaborate cirri of the other species. It is, however, the only species that is sexually dichromatic. It differs from C. exasperatus sp. nov. in body depth (mean 4.3 in SL vs. 3.0 in SL), and from all other species in habitat and salinity tolerance. The species is morphologically very close to C. superciliosus , and genetically the two are sister species (see, A genetic perspective).

Distribution

Currently known only from the Bot, neighbouring Kleinmond, and Klein river estuaries, Western Cape Province. A single specimen was picked up on a jetty in Seekoeivlei near Cape Town, possibly dropped by a gull. It is unlikely that a bird would have flown some 50 km to drop its meal, and it is possible that the species occurs in other water bodies as well ( Fig. 1 ); it has not been recorded from Seekoeivlei.

Notes

This is possibly the smallest of the five described species, with a maximum recorded length of 133 mm compared with 183 mm for C. musaicus sp. nov., 175 mm for C. ornatus , 220 mm for C. arborescens , and 300 mm for C. superciliosus (Smith in Smith & Heemstra, 1986). (The maximum length of C. exasperatus sp. nov. is not known; the largest specimen is 127 mm SL.)

Material examined

Holotype: SAM 29449 (78.5 mm SL), female, Bot River estuary, Western Cape, South Africa (34°21 ′ S, 18°71 ′ E), collected by B.A. Bennett, 6 April 1981.

Paratypes: SAIAB 18239 (14: 22.1–113.0 mm SL) ; SAM 29450 (83.0 mm SL) ; SAM 29451 (9: 45.6– 124.0 mm), all collected with holotype from May 1980 to April 1981.

Non-types: SAIAB 83658 (10: 39–83 mm SL), Maanskynbaai, Klein River estuary, Western Cape (34.41°S, 19.33°E), collected by W. Holleman, 10 February 2009 ; SAIAB 83655 (4: 45–50 mm SL), Maanskyn Baai, Klein River estuary, Western Cape (34.41°S, 19.33°E), collected by W. Holleman, 10 February 2009 ; SAIAB 96203 (19.0 and 36.5 mm SL), Fisher Haven, Klein River estuary, Western Cape, collected by S. von der Heyden, 24 October 2008 .

GenBank accession numbers

JF320841 , JF320842 (mtDNA control region); JF320855 , JF320856 (mtDNA 16S rRNA); JF320869 , JF320870 (mtDNA 12S rRNA); JF320883 , JF320884 (rhodopsin).

CLINUS SUPERCILIOSUS (LINNAEUS 1758)

FIGS 1 ; 6A–F ; 9

Blennius superciliosus Linnaeus, 1758: 257 .

Blennius mustelaris Linnaeus, 1758: 257 ; Gronow ed., Grey, 1854: 98 ( mustellaris ).

Blennius punctulatus Lacepède, 1800: 460 .

Blennius mustela Lacepède, 1800: 459 .

Blennius spadiceus Bloch & Schneider, 1801: 172 .

Blennius capensis Forster , in Bloch & Schneide, 1801: 175.

Clinus superciliosus: Cuvier, 1817: 173 ; Valenciennes, in Cuvier & Valenciennes, 1836: 360; Gilchrist & Thompson, 1908: 113; Gilchrist & Thompson, 1918: 149; Barnard, 1927: 855; Smith, 1945: 541, 1949: 354, pl. 77, fig. 986; Penrith, 1969: 57, fig. 22; Smith, in Smith & Heemstra, 1986: 765, pl. 118.

Clinus superciliosus: Swainson, 1839: 276 ; Hubbs, 1952: 106 ( Clinetrachus ).

Blennius versicolor Pappe, 1853: 27 .

Blennius mycterizans Gronow ed., Grey, 1854: 97.

Clinus dubius Castlenau, 1861: 51 .

Clinus pantherinus Castlenau, 1861: 52 .

Non Clinus superciliosus var. arborescens: Gilchrist & Thompson, 1908: 115 .

Non Clinus ornatus: Gilchrist & Thompson, 1908: 116 .

Caboclinus superciliosus: Smith, 1966: 73 .

Doubtful synonym: Blennius varius Seba, 1758: 90 , 93.

Description

Dorsal fin XXXII–XXXVIII, 6–9 (rarely with XXXII spines or 6 rays); anal fin II, 24–28 (rarely 28 rays); pectoral fins 14–18 (rarely 18), number of rays and spines increase from east to west with decreasing water temperature; see section on ‘Water temperature vertebrae numbers’ and Table 2; pelvic fins I, 2. Vertebrae 18 + 30–33. Lateral line scales 29–37 + 26–35 (rarely 37, and usually 31–34 anterior scales; rarely 29 or 35, and usually 29–32 posterior scales); anterior portion with double pores for entire length before curving down. Head length 3.2–4.4 [3.7] in SL; eye diameter 2.6–4.8 [3.5], maxilla 1.9–3.6 [2.6], snout 2.8–4.8 [3.8] in head length; dorsal-fin origin 1.3–1.8 [1.5] in head length; dorsal fin height 9.2–16.7 [11.8] in SL.

Head profile fairly blunt: 74–81° [78°]. Mouth downturned, maxilla extending to vertical through posterior margin of pupil. Orbital cirrus length equal to pupil diameter, flat and at least slightly widened and bifurcate at the end in juveniles, with several short branches in adults ( Fig. 6B ); nasal cirrus a simple flat flap. Dorsal-fin crest of mature males tall and triangular, up to 3 times the height of dorsal fin, of females to 1.7 times the height of dorsal fin. Dorsalfin spines with a single filament at the tips. Scales on dorsal, caudal, and pectoral fin bases.

Live colour: Clinus supersiliosus comes in a wide variety of colours (e.g. Fig. 6C–F ), which are often linked to its immediate environment. Intertidal fish seem invariably to have a ‘default’ coloration of browns and purples ( Fig. 6C ), also seen in some subtidal fish (see the Discussion).

Colour in alcohol: Mottled purples on a yellowish background, the mottling with some very regular features:

1. a diffuse to clearly defined dark spot in the centre of the crest;

2. a dark blotch on the dorsum at the base of spines 16–19;

3. often a dark-purple ‘comma’ mark on the opercle, suggesting an incomplete ocellus;

4. a diffuse to well-defined dark bar on the base of the pectoral fin;

5. a small white oval mark at the base of the middle caudal-fin spines.

On the west coast many individuals lose the mottled pattern and become yellow, with fine black speckled over entire body, more distinct in smaller than larger specimens. This has not been observed in specimens from the south and east coasts. Genetically no species-level differences have been detected between west and south coast specimens.

Distribution

Recorded from intertidal pools to a depth of at least 50 m from Rocky Point , Namibia (19°S) , to Kei Mouth on the south-east coast of South Africa.

Comparisons

Clinus superciliosus can be distinguished from C. arborescens , C. ornatus , and C. musaicus sp. nov. by its simple orbital cirrus; from C. exasperatus sp. nov. in body depth (3.7–4.2 vs. 3.0 in SL), number of dorsal-fin spines (almost always <38 vs. 38), and facial pigmentation; and from C. spatulatus by habitat and number of pectoral-fin rays (usually 15 or more vs. usually 14).

However, von der Heyden et al. 2011 recover three distinct genetic lineages within ‘ C. superciliosus ’, with sequence distances equal to recognized species. This suggests that a number of cryptic species might exist, which requires a further, thorough morphological analysis of fishes in this group.

Material examined

Namibia: SAIAB 63525 (43.8 mm), Luderitz; SAIAB 35854 (17: 34.1–60.6 mm) and SAIAB 13719 (5: 74.3– 128.0 mm), Swakopmund; SAIAB 5788 (51.5 and 55.3 mm), Walvis Bay. West Coast, South Africa: SAM 24225 (13: 71.2–113.0 mm), Doring Bay; SAM 25428 (9: 31.9–73.3 mm), SAM 24215 (17: 54.4– 101.0 mm), SAIAB 57087 (29.5 and 33.6 mm), SAIAB 15467 (26: 31.9–39.5 mm), Port Nolloth; SAM 25137 (62.0–130.0 mm) and SAIAB 15460 (3: 50.9– 73.0 mm), Lambert’s Bay. Peninsula, Atlantic Coast: SAM 32907 (8: 78.3–158.0 mm), Strandfontein; SAM 32915 (32: 43.5–138.0 mm), Sea Point; SAM 23214 (5: 128–166 mm), Millers Point; SAIAB 32916 (11: 30.3–92.2 mm), Olifantsbos; SAIAB 32945 (5: 44.7–128.0 mm), Die Mond. False Bay to Bredasdorp: SAM 23904 (7: 105–205 mm), Onrus; SAM 24687 (18: 50.1–233.0 mm), Hondeklip Bay; SAM 25425 (18: 52.3–163.0 mm), Melkbosstrand; SAIAB 32141 (6: 139–180 mm) and SAIAB 32748 (10: 43.7– 70.0 mm), Cape Point; SAIAB 34015 (40.3 and 91.0 mm) and SAIAB 15462 (102 and 110 mm) False Bay; SAM 24547 (7: 100.0–146.0 mm), SAIAB 14188 (13: 51.0–80.0 mm) and SAIAB 15298 (16: 42.7– 150.0 mm), Bredasdorp. Knysna to Tsitsikamma: SAIAB 10855 (10: 45.0– 71.5 mm), SAIAB 8781 (98.0 mm), SAIAB 14219 (44.4 and 100.0 mm), SAIAB 2850 (5: 71.5–176 mm) and SAIAB 10886 (8: 48.9–110.0 mm), Knysna ; SAIAB 32931 (10: 62.1– 125.0 mm) and SAIAB 32943 (36: 36.1–142.0 mm), Buffalo Bay; SAIAB 15437 (113 and 124 mm), SAIAB uncat. (8: 65–136 mm) (Stn 24, 12.8.88), SAIAB uncat. (9: 62.7–131.o mm) (Stn 18, 21.7.88) and SAIAB uncat. (29: 60.5–128.0 mm), Tsitsikamma (Stn 16, 20.7.88). South-East Coast, South Africa: SAIAB 62665 (check no.) (5: 56.6–79.7 mm) Cape St Francis; SAIAB 1646 (3: 79.5–115.0 mm), Sea View; SAIAB 10733 (5: 93–110 mm), Smale 860307 (4: 68.7– 115.0 mm), Smale 860307 (160.0 mm) and Smale 860307 (7: 75.3–115.0 mm), Skoenmakerskop; SAIAB 63181 (9: 51.9–113.0 mm), Cape Recife; SAIAB 11377 (120 mm), SAIAB 32733 (3: 32.5–37.1) and SAIAB 14335 (56.5 and (2.0 mm), Algoa Bay; SAIAB 70871 (33.4–113.0 mm), Bird Island; SAIAB 15445 (78.2 and 104.0 mm), Bushman’s River; SAIAB 32944 (8: 85.6– 127.0 mm), SAIAB 9280 (3: 77–115 mm) and SAIAB 8260 (3: 75–114 mm), Port Alfred; SAIAB 32921 (11: 43.3–110.0 mm), Clayton’s Rocks; SAIAB 75443 (135.0 mm) and SAIAB 15432 (5: 56.6–121.0 mm), Great Fish Point; SAIAB 32756 (7: 36.9–105.0 mm), Hamburg; SAIAB 70363 (4: 93–160 mm) and SAIAB 70374 (146 and 152 mm), Morgan Bay.

GenBank accession numbers

JF320837 , JF320838 , JF320843 (mtDNA control region), JF320851 , JF320852 , JF320857 (mtDNA 16S rRNA); JF320865 , JF320866 , JF320871 (mtDNA 12S rRNA); JF320879 , JF320880 , JF320885 (rhodopsin).

Figures

Figure 1. Distribution of the six species of the Clinus superciliosus complex.

Figure 3. A, Clinus musaicus, holotype, SAIAB 96616, male, 108 mm SL, Windmill, False Bay (drawn by Elaine Heemstra); B, orbital cirri; C, Clinus musaicus, photograph of holotype (G. Zsilavecz); D–F, Clinus musaicus underwater, habitat and different colour forms, False Bay (G. Zsilavecz).

Figure 4. A, Clinus ornatus, Lectotype, SAM MB-F038012, male, 158 mm SL (drawn by Elaine Heemstra); B, orbital cirri; C, D, Clinus ornatus underwater, False Bay (C, G. Zsilavecz; D, unknown); E, Clinus spatulatus, SAIAB 93658, male, 84.6 mm SL, Klein River estuary, aquarium photograph (W. Holleman); F, orbital cirri (from Bennett 1983); G, Clinus spatulatus underwater, male; H, female, Bot River estuary; I, female, Klein River estuary (G. Zsilavecz).

Figure 5. Posterior of skull with first three dorsal-fin spines and supporting pterygiophores: A, Clinus superciliosus; B, Clinus ornatus.

Figure 6. A, Clinus superciliosus, SAIAB 8260, male, 78 mm SL (drawn by David Voorveld); B, orbital cirri; C–F, Clinus superciliosus underwater, different colour forms and habitats, False Bay; C is the ‘default’ colour form (G. Zsilavecz).

Figure 9. A, Bayesian tree of the combined mtDNA and nDNA fragments with a total of 2070 bp. Numbers at nodes are support values (ML above and Bayesian PP and MP below); values less than 50% are marked with - -; B, ML tree of the control region only showing the monophyly of the Clinus superciliosus and C. spatulatus clades. Only ML support values are shown.