Figures 1–2, 5–6; Tables 1–3
Parapercis binivirgata (not of Waite, 1904): Moreland, 1975: 288 (in part).
Parapercis NFZ1: Clark & Roberts, 2008: Appendix 4, unnumbered.
Holotype. NMNZ P.052461, 113.5 mm, Ranfurly Bank, New Zealand, 37°32.782’S 178°53.415’E, 68–70 m, beam trawl, RV Tangaroa, 30 May 2011.
Paratypes. (n = 17) AMS I.42754-002, 76.9 mm, Wanganella Bank, west Norfolk Ridge, Tasman Sea, 32°36.9417’S 167°35.4167’E, 121–126 m, trawl, NORFANZ survey, 29 May 2003; CSIRO H.6061-01, 3: 56.5–68.9 mm, same data as above; NMNZ P.039278, 81.2 mm, same data as above; NMNZ P.039283, 2: 78.8–81.8 mm, same data as above; NMV A.25154, 76.3 mm, same data as above; NMNZ P.002207, 156.0 mm, Off Mayor Island, Bay of Plenty, New Zealand, 37°20’S 176°18’E, 146 m, trawl, Jul 1957; NMNZ P.036802, 128.6 mm, reef SE of White Island, Bay of Plenty, New Zealand, 37°31’S 177°12’E, 160 m, handline, Aug 1999; NMNZ P.041834, 81.6 mm, SE of Colville Ridge, New Zealand, 37°28.25’S 177°13.065’E, 200– 175 m, epibenthic sled, 12 Nov 2004; NMNZ P.052460, 99.1 mm, Ranfurly Bank, New Zealand, 37°32.782’S 178°53.415’E, 68–70 m, beam trawl, 30 May 2011; NMNZ P.052476, 112.7 mm, Ranfurly Bank, New Zealand, 37°38.783’S 178°56.793’E, 158–160 m, beam trawl, 1 Jun 2011 (genetic sample listed in Table 4); NMNZ P.052477, 81.6 mm, Off East Cape, New Zealand, 37°31.178’S 178°51.833’E, 110–113 m, beam trawl, 1 Jun 2011; NMNZ P.052478, 2: 52.6–67.4 mm, same data as previous; NMNZ P.052492, 75.7 mm, Off East Cape, New Zealand, 37°35.682’S 178°51.907’E, 56–58 m, beam trawl, 1 Jun 2011.
Non-types. (n = 5) NMNZ P.041807, 45.9 mm SL, SE of Colville Ridge, New Zealand, 37°21.305’S 177°06.09’E, 260–280 m, epibenthic sled, 14 Nov 2004; NMNZ P.052456, 58.1 mm, Off East Cape, New Zealand, 37°28.185’S 178°51.78’E, 106–177 m, epibenthic sled, 1 Jun 2011; NMNZ P.052459, 89.5 mm, North Taranaki Bight, NW of Kawhia Harbour, New Zealand, 37°58.302’S 174°04.857’E, 115–118 m, beam trawl, 2 Apr 2011; NMNZ P.052471, 47.8 mm, NW of Three Kings Islands, New Zealand, 33°57.578’S 171°46.343’E, 92–96 m, epibenthic sled, 28 Mar 2011; NMNZ P. 052484, 62.0 mm, same data as previous.
Diagnosis. A species of Parapercis with dorsal-fin rays V, 23; anal-fin rays I, 19; pectoral-fin rays 19–21 (modally 19); lateral-line scales 57–63 (modally 60); gill rakers 5–6 + 8–10 = 13–15 (modally 14); predorsal scales 9 – 11, cycloid; scales on cheek ctenoid (except few anteriorly on preorbital), in about 18 horizontal rows; 6 canine teeth in outer row at front of lower jaw; vomer with 2 rows of robust conical teeth in adults; palatines with 1–2 rows of small teeth; angle of subopercle smooth, lacking spinules; 10 abdominal and 22 caudal vertebrae; pelvic fins reaching between vent and base of second anal-fin ray; and coloration including seven broad reddish-brown vertical bands on upper body between spinous dorsal fin and caudal peduncle, each band except first and last partially bifurcated into two close-set double bars, each bar with a black smudge-like spot at its lower end, membrane of spinous dorsal-fin black.
Description. Dorsal-fin rays V, 23; anal-fin rays I, 19; all dorsal- and anal-fin rays branched, last to base; pectoral-fin rays 19 (19–21, modally 19, occasionally asymmetrically 19 on one side and 20 on the other), upper ray unbranched, others including lowermost branched; pelvic-fin rays I, 5; branched caudal-fin rays 15; lateral-line scales 61 (57–63), plus usually 2 smaller pored scales on caudal-fin base; scales above lateral line to origin of dorsal fin 8 (7–8), to base of anterior soft rays of dorsal fin 5 ½; scales below lateral line in a oblique row to origin of anal fin 14 (14–15); circumpeduncular scales 24 (24–25); predorsal scales 10 (9–11), extending to (or almost to) a vertical from upper corner of preoperculum; horizontal row of scales from preorbital across cheek to edge of preopercle about 20, vertical rows below middle of eye 7; gill rakers on first arch 5 + 9 = 14 (5–6 + 8–10 = 13–15); branchiostegal rays 6; vertebrae 10 + 22 = 32.
Body depth 6.2 (5.6–6.25) in SL; body subcylindrical, greatest width 0.9 (0.9–1.0) in body depth; head length 3.95 (3.8–4.0) in SL; snout pointed, its length 3.85 (3.9–5.15) in HL; orbital diameter 3.35 (2.75–3.35) in HL; eyes directed more laterally than dorsally, bony interorbital space narrow, 8.25 (6.65–9.65) in HL; caudal-peduncle depth 2.5 (2.65–2.85) in HL; caudal-peduncle length 3.0 (2.75–3.1) in HL.
Mouth oblique, jaws terminal, or upper lip slightly protruding beyond tip of lower jaw; curved canine teeth at front of lower jaw slightly projecting, but not visible when mouth fully closed; upper jaw extending to vertical from middle of eye (middle of eye to just posterior to rear margin of pupil), but fleshy flap extending posteriorly from the tip of the maxilla to vertical between posterior margin of pupil and posterior margin of orbit, upper-jaw length 2.0 (2.0–2.05) in HL; upper jaw with 20–21 (19–22) outer curved canines on each side, first 6–8 clearly the largest, of these the second or third the largest, those following gradually reducing in size posteriorly, broad inner band of small villiform teeth anteriorly, narrowing gradually to form only two rows at rear of jaw; front of lower jaw on each side with 3 enlarged curved canines in distinctly separate outer row, tooth third from symphysis largest, followed in outer row by 4–5 (4–6) small canine teeth, then 3 abruptly larger strongly curved canines of which the middle is distinctly largest, broad inner band of villiform teeth extending posteriorly from symphysis to side of jaw just posterior to largest tooth in outer row, remaining teeth subequal, moderate, in single row of 16 (12–16). Vomer with 2 (1–2, single row only in juveniles) irregular crescentic rows of 6 (4–9) robust conical teeth anteriorly, medial teeth largest, second posterior row shorter, comprised of 7 (0–7) slightly smaller teeth; palatines with 2 (1–2) rows of small subequal teeth, 8 (6–8) in outer row and 3 (0–3) medially in inner row. Tongue spatulate with broadly rounded tip, dorsal surface covered in numerous minute papillae.
Gill membranes united with broad free fold, not attached to isthmus. Gill rakers short, the longest about 3.5 in length of longest gill filament on first gill arch. Anterior nostril small, situated anterior to mid-eye, about half distance from anterior margin of eye to snout tip, with a membranous tube, often lying flat against snout in preserved specimens. Posterior nostril slightly more than half distance from anterior margin of eye to anterior nostril, dorsoposterior to, and about same width as anterior nostril, its opening oval-shaped, with a rounded membranous flap on anteroventral edge, the flap covering most of the nasal opening when depressed; internarial distance about twice width of posterior nostril.
Opercle with distinctly exposed, flattened, blade-like spine; subopercle, including angle, entire; preopercle entire, the margins naked and broadly rounded.
Lateral line continuous, ascending smoothly from opercle to below second or third soft dorsal-fin ray, then approximately following contour of back; scales ctenoid, except for those on nape, on breast and midline of belly, and several anteriormost scales on subopercle; scales on middle of sides with up to about 46 cteni; scales on cheek extending forward to a vertical from anterior margin of orbit in adults to below anterior margin of pupil in juveniles; no scales on dorsal, anal or pelvic fins; up to 10 rows of small feebly ctenoid followed by cycloid scales on base of pectoral-fin rays; elongate ctenoid scales densely arranged on proximal two-thirds of caudal fin in up to 28 rows.
Origin of dorsal fin just posterior (above or just posterior) to vertical from axil of pectoral fin, the predorsal length 3.65 (3.3–3.75) in SL; dorsal-fin spines progressively longer, the first 9.95 (8.15–13.0) in HL; fifth dorsalfin spine longest, 4.4 (3.75–4.4) in HL; membrane from fifth spine to first soft ray attached near tip of fifth dorsalfin spine and only slightly incised; 20 th (19 th or 20 th) soft dorsal-fin ray the longest, 1.85 (1.65–1.85) in HL; origin of anal fin below base of fifth soft dorsal-fin ray, preanal length 2.2 (2.1–2.2) in SL; anal-fin spine slender, closely attached to first soft ray, 4.9 (4.35–6.3) in HL; longest soft anal-fin ray the 17 th (usually 16 th or 17 th), 1.95 (1.75–2.05) in HL; caudal fin slightly rounded (truncate to slightly rounded), length 5.25 (4.6–5.45) in SL; pectoral fins rounded, 11 th (10 th or 11 th) ray longest, 5.1 (4.15–5.0) in SL, about equal to pelvic fins; origin of pelvic fins in advance of upper base of pectoral fins, on a vertical just anterior to upper corner of opercular opening (upper corner of opercular opening to tip of operculum), prepelvic length 4.0 (3.6–4.05) in SL; pelvic-fin spine closely attached to first soft ray, its termination fleshy and attenuated, reaching about three fourths distance to tip of first ray; fourth soft pelvic-fin ray longest, reaching base of anal-fin spine (to vent in largest paratype, to base of second anal-fin ray in smallest paratype), 4.65 (3.9–4.95) in SL.
Colour when fresh. Upper half of body in holotype (Fig. 1B–C) and paratypes (eg Fig. 1D) pale pinkish orange, with a series of seven reddish brown vertical bands. First band narrower and more diffuse than others, with darker brown blotch restricted to posteroventral portion of band. Second to seventh bands partially bifurcated, forming six pairs of close-set double bars, each with a darker brown smudge-like blotch at its lower end. First three bands extending below lateral line, fourth on or slightly below lateral line, remainder terminating on lateral line. A small dark brown spot on upper base of caudal fin, aligning horizontally with series of bands on body. A broad longitudinal diffuse yellow stripe from upper pectoral-fin base to mid-base of caudal fin, comprised of yellow edges to about four vertical rows of scales, stripe passing directly below and highlighting lower edge of series of dark brown blotches on upper body. Lower body uniformly pale creamish white. Head mostly pinkish yellow, infused with crimson-red on upper half of upper lip, snout, preorbital, interorbital and naked occipital region. Lower half of upper lip bright yellow. Operculum below opercular spine pale yellow. Lower lip and undersurface of lower jaw stark white. Spinous dorsal-fin membrane jet black. Soft dorsal fin translucent, with numerous oblique posteroventrally-directed orange-yellow bands. Basal third of anal fin white, distal two-thirds of fin pale yellow. Caudal fin semi-translucent, with six narrow wavy vertical orange-yellow bands, alternating with six similar bluegrey bands, bands becoming faint in lower quarter of fin. Pectoral fins mainly translucent, but membrane of basal third of fin faintly yellowish. Pelvic fins white, with a faint yellowish blush. Colour in life similar to fresh specimens (Fig. 2).
Colour in alcohol. Head and body of holotype (Fig. 1A) and paratypes pale yellowish cream, with a series of seven vertical brown bands on upper body. First band below posterior portion of spinous dorsal-fin base, faint and narrow above, terminating on midbody below lateral line in a broader dark brown smudge-like blotch. Second band much broader than first, successive bands becoming progressively narrower, each partially bifurcated, forming close-set double bars, each bar forming darker brown smudge-like blotch at its lower end. Lower edge of series of dark blotches in parallel at midbody. Belly pale cream. Tongue creamish white. Peritoneum pale. Spinous dorsal fin membrane black. Soft dorsal, anal and pectoral fins semitranslucent, lacking any distinctive markings. Caudal fin with six narrow wavy greyish bands, the latter fading in lower quarter of fin. Pelvic fins pale cream.
Molecular results. The phylogenetic analyses indicate that P. nigrodorsalis is closely related to (sister taxon), but separate from P. binivirgata and both species form a well-supported clade (posterior probability of 1) with the currently undescribed P. sp. 4 (of Johnson) (Figure 6). The three other species of Parapercis, P. allporti , P. colias and P. gilliesii form the other well supported clade within our phylogeny (Figure 6). In addition to the clear phylogenetic support for the separate but close relationship of P. nigrodorsalis within its clade, it also has an average uncorrected sequence divergence of 5.38% and 7.63% from P. binivirgata and P. sp. 4 (of Johnson) respectively (Table 3). While lower that the overall sequence divergence among all the species of Parapercis in our study (13.49%), these values are higher than those found between P. allporti and P. gilliesii (3.04%); two closely related Parapercis species easily distinguished on morphological characters. Furthermore, the values are higher or comparable to those found among Pinguipes brasiensis, Pinguipes chiliensis and Pseudopercis semifasciata (3.83% – 6.29%, Table 3). Also notable is that sequence divergence within species (where more than one sequence per species available) is almost non-existent (0–0.15%, Table 3).
Etymology. From the Latin nigro for black and dorsalis for dorsal fin, in reference to the distinctive black spinous dorsal-fin membrane of the species.
Distribution and abundance. Known on rocky reef, gravel and rubble bottom on Wanganella Bank, western Norfolk Ridge, Tasman Sea, south to the North Island of New Zealand, from North Taranaki Bight, north to the Three Kings Islands and south-east to the Bay of Plenty and Colville Ridge, in depths of 56 to 280 m (Fig. 5).
It is difficult to draw conclusions on the abundance of P. nigrodorsalis on the Norfolk Ridge from NORFANZ results, as few trawls were undertaken in the depth range that specimens were collected (over 90 percent of trawl and sled tows were in depths greater than 200 m). However, in New Zealand it appears to be relatively common where suitable habitat exists, with 14 specimens collected by trawl in depths of 56– 280 m. The species has also been observed and photographed off the North Island of New Zealand by divers using rebreather apparatus in depths of 55–71 m in the Poor Knights Islands Marine Reserve, at locations known locally as Ngaroimata Point, Northern Arch, Serpent Rock and Landing Bay Pinnacle (Fig. 2). A large individual was identified from an underwater photograph taken by Kendall Clements in 2010 at “The Archways”, Burgess Island, Mokohinau Group, outer Hauraki Gulf, at a depth of 16– 20 m. The latter is the shallowest confirmed record for the species.
Discussion. Parapercis nigrodorsalis (Figs. 1–2) is most similar to P. binivirgata (Figs. 3–4), sharing similar morphology, meristic formulae, and coloration, including numerous dark transverse bars on the upper sides. It differs most obviously in coloration, having seven broad bands across the upper body, the first originating below the posterior portion of the spinous dorsal fin and the second to seventh bifurcated to form close-set double bars with darker smudge-like blotches below (versus 13 narrower, distinctly separate dark bars on the upper body, lacking darker spots or blotches below, and the first bar originating on the nape). In addition, the spinous dorsal-fin membrane is jet black, the pelvic fins lack dark pigmentation, the upper pectoral-fin base lacks a red blotch and there are six alternating orange-yellow and blue-grey bands on the caudal fin in P. nigrodorsalis (versus spinous dorsal-fin membrane semitranslucent to faintly reddish, or greyish, pelvic fins dusky on the basal half to two-thirds of medial rays, upper pectoral-fin base with a conspicuous red blotch, and seven alternating bands on the caudal fin, respectively, in P. binivirgata). Although meristics for both species strongly overlap, there are modal differences in counts of pectoral-fin rays (19–21, modally 19 in P. nigrodorsalis, versus 19–21, modally 20 in P. binivirgata), lateral-line scales (57–63, modally 60, versus 59–67, modally 64) and total gill rakers (13–15, modally 14, versus 12–15, modally 13) (Table 2). The two species are sympatric in the region off the eastern side of the North Island of New Zealand, but P. nigrodorsalis has not been recorded elsewhere within the known range of P. binivirgata.
Among other species of Parapercis with numerous transverse bands on the upper body, P. nigrodorsalis is readily distinguished from P. decemfasciata by the number of bands and the dorsal-fin spine and ray counts (10 narrow pale reddish bars, extending well below midbody, and dorsal fin IV, 25, versus seven broader bands not extending below midbody, and dorsal fin V, 23 in P. nigrodorsalis); from P. multifasciata by coloration (10 narrow dusky bands extending below midbody and several broad wavy yellow bands across the head and nape, versus seven broader bands not extending below midbody and no yellow bands on head and nape in P. nigrodorsalis); and from P. allporti by a lower dorsal fin-ray count, absence of palatine teeth and different configuration of dark bars on the upper body (dorsal-fin rays 21, palatine teeth absent, and five to seven principal dark bars alternating with numerous narrower secondary bars, extending below midbody, versus dorsal-fin rays 23, palatine teeth present and seven broad bands not extending below midbody in P. nigrodorsalis).
A successful genetic sequence was only generated from a single specimen of the new species (paratype NMNZ P.052476). Unfortunately this specimen was in relatively poor condition and no images of it in fresh coloration were available, hence it was unsuitable for designation as the holotype. NMNZ P.052476 was collected from a location closely adjacent to that of the holotype and has almost identical meristic and morphometric values and preserved coloration to the holotype, leaving no doubt that the two are conspecific.